Neanderthal

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How to read a taxoboxNeanderthals
H. neanderthalensis La Ferrassie 1
H. neanderthalensis La Ferrassie 1
Conservation status
Prehistoric
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Genus: Homo
Species: H. neanderthalensis
Binomial name
Homo neanderthalensis
King, 1864
Neanderthal range
Neanderthal range
Synonyms

Palaeoanthropus neanderthalensis
H. s. neanderthalensis

Neanderthal or Neandertal is a relatively recent extinct member of the Homo genus that inhabited Europe and parts of western Asia in the middle to late Pleistocene, first appearing in the fossil record some 200,000 to 400,000 years ago and disappearing about 30,000 years ago. They are a sister group of present-day humans, with whom they came in contact during the later part of their history, from at least 80,000 years ago, and by whom they were displaced. They are variously classified as a species (Homo neanderthalensis) or subspecies (H. sapiens neanderthalensis).

Sequencing of the Neanderthal genome has suggested that Neanderthals, modern humans, and another hominid known as Denisovans descended from a common ancestor several hundred thousand years ago (perhaps 350,000 to 500,000 years ago). The branch giving rise to the Neanderthal/Denisovan lineage is theorized to have migrated from Africa and shortly thereafter split into Neaderthals (which settled in Europe and Western Asia) and Denisovans (which settled further to the east). Later, anatomically modern humans left Africa (perhaps as recently as 50,000 to 100,000 years ago). Distinct Neanderthal DNA found in the genome of living humans has suggested subsequent interbreeding among anatomically modern humans and Neanderthals.

The fossil record shows that Neanderthals lived in Europe and Western Asia, as far south as the Middle East, as far east as Siberia, and as far west as Great Britain. In addition to the discovery of fossil bones, various tools have also been discovered and some evidence has been presented that Neanderthals intentionally buried their dead.

Contents

Overview of human evolution

The earliest delineated member of the genus Homo is H. habilis, which lived from 2.33 to 1.44 million years ago, although some authorities do not consider it should be included in Homo, considering it more worthy, for example, to be retained in Australopithecus (Wood and Richmond 2000). Homo erectus is considered to have arrived around 1.8 million years ago, with fossils supporting its existence to 143,000 years ago. Homo ergaster is another early Homo species that has been delineated, and traced to about 1.8 to 1.3 million years ago. H. ergaster is possibly ancestral to or shares a common ancestor with H. erectus, or is the African variety of H. erectus; it is widely considered to be the direct ancestor of later hominids such as Homo heidelbergensis, Homo sapiens, Neanderthals, Denisovans, and even Asian Homo erectus. Homo erectus and H. ergaster were the first of the hominina known to leave Africa. For example, H. erectus is known to have spread as far as Georgia, India, Sri Lanka, China, and Java.

There is also support for the idea that that the numerous distinct species being recognized in the fossil record, such as H. erectus and H. habilis, are actually just morphological variation among members of a single evolving lineage among early members of the Homo genus, and that perhaps even only one species with a lot of variability emerged from Africa (Wilford 2013a; Watson 2013; Lordkipanidze et al. 2013).

Modern human beings, Neanderthals, and Denisovans are believed to have shared a common ancestor about 400,000 to 500,000 years ago (Marshall 2013; Green et al. 2010). One theory is that these three groups all descended from Homo heidelbergenesis, which lived between 600,000 to 250,000 years ago (Marshall 2013) (other species suggested as ancestral are H. rhodesiensis and H. antecessor). One branch of H. heidelbergenesis are theorized to have left Africa about 400,000 years ago and split shortly thereafter to become Neanderthals, which settled in West Asia and Europe, and Denisovans, which settled farther to the east (NG 2013).

Neanderthals are considered to have lived from perhaps 400,000 years ago, with their appearance in the European fossil record variously put at 200,000 (Zimmer 2013) to 400,000 years ago (Green et al. 2010). Neanderthals disappeared from the fossil record about 30,000 years ago. Based on the DNA sequences for the nuclear genome of Neanderthals and modern humans, the population split between Neanderthals and modern humans took place 270,000 to 440,000 years ago (Reich et al. 2010).

Archaic Homo sapiens, the forerunner of anatomically modern humans, appeared between 400,000 and 250,000 years ago (O'Neil 2013b). Anatomically modern humans are believed to have evolved from archaic Homo sapiens in the Middle Paleolithic, about 200,000 to 130,000 years ago (SA 2005; NG 2013), then migrated out of Africa some 50,000 to 100,000 years ago (Recent African Ancestory Theory) and replaced local populations of H. erectus, H. floresiensis, H. heidelbergenesis, and the Denisovan and Neanderthal populations. Cro-Magnon Man is a name applied to the earliest known European examples of Homo sapiens sapiens, modern human beings. Cro-Magnons lived from about 40,000 to 10,000 years ago in the Upper Paleolithic period of the Pleistocene epoch.

The transition to behavioral modernity for Homo sapiens with the development of symbolic culture, language, and specialized lithic technology happened around 50,000 years ago according to many anthropologists (Mellars 2006b), although some suggest a gradual change in behavior over a longer time span (Mcbrearty and Brooks 2000). Until about 50,000 to 40,000 years ago, the use of stone tools seems to have progressed stepwise: Each phase (habilis, ergaster, and neanderthal) started at a higher level than the previous one, but once that phase had started, further development was slow. After 50,000 years ago, in what Jared Diamond, author of The Third Chimpanzee, and other anthropologists characterize as a "Great Leap Forward," human culture apparently started to change at much greater speed: "Modern" humans started to bury their dead carefully, made clothing out of hides, developed sophisticated hunting techniques (such as pitfall traps, or driving animals to fall off cliffs), and made cave paintings. This speed-up of cultural change seems connected with the arrival of modern humans, Homo sapiens sapiens. Additionally, human culture began to become more technologically advanced, in that different populations of humans begin to create novelty in existing technologies. Artifacts such as fish hooks, buttons, and bone needles begin to show signs of variation among different population of humans, something what had not been seen in human cultures prior to 50,000 BP.

Fossil discoveries and history

Location of Neander Valley, Germany. (The highlighted areas are the modern federal states of North Rhine-Westphalia and Rhineland-Palatinate.)

Fossils of Neanderthals were first found in the eighteenth century prior to Charles Darwin's 1859 publication of The Origin of Species, with discoveries at Engis, Belgium in 1829, at Forbes Quarry, Gibraltar in 1848, and most notably a discovery in 1856 in Neander Valley in Germany, which was published in 1857. However, the earlier findings prior to 1856 were not recognized as belonging to archaic forms, but were widely misinterpreted as skeletons of modern humans with deformities or disease (Gould 1990).

The August day in 1856 when a fossil was discovered in a limestone quarry in Germany is heralded as the beginning of paleoanthropology as a scientific discipline (Kreger 2005). This discovery of a skullcap and partial skeleton in a cave in the Neander Valley (near Dusseldorf) was the first recognized fossil human form, although the prior two discoveries were subsequently recognized as the first early human fossils found (Smithsonian 2014b).

The type specimen, dubbed Neanderthal 1, consists of a skull cap, two femora, three bones from the right arm, two from the left arm, part of the left ilium, fragments of a scapula, and ribs. The workers who recovered this material originally thought it to be the remains of a bear. They gave the material to amateur naturalist Johann Karl Fuhlrott, who turned the fossils over to anatomist Hermann Schaffhausen. The discovery was jointly announced in 1857.

Type Specimen, Neanderthal 1

These, and later, discoveries led to the idea that these remains were from ancient Europeans who had played an important role in modern human origins. The bones of over 400 Neanderthals have been found since.

The term Neanderthal Man was coined by Irish anatomist William King, who first named the species in 1863 at a meeting of the British Association, and put it into print in the Quarterly Journal of Science the following year (Kreger 2005). The Neanderthal or "Neander Valley" itself was named after theologian Joachim Neander, who lived there in the late seventeenth century.

"Neanderthal" is now spelled two ways. The spelling of the German word Thal, meaning "valley or dale," was changed to Tal in the early twentieth century, but the former spelling is often retained in English and always in scientific names. The original German pronunciation (regardless of spelling) is with the sound /t/. When used in English, the term is usually anglicized to /θ/ (as in thin), though speakers more familiar with German use /t/.

Classic Neanderthal fossils have been found over a large area, from northern Germany, to Israel to Mediterranean countries like Spain and Italy, and from England in the west to Uzbekistan in the east. This area probably was not occupied all at the same time; the northern border of their range especially would have contracted frequently with the onset of cold periods. On the other hand, the northern border of their range as represented by fossils may not be the real northern border of the area that they occupied, since artifacts indicative of the Middle Paleolithic have been found even further north, up to 60° on the Russian plain (Pavlov et al. 2004).

In Siberia, Middle Paleolithic populations are evidenced only in the southern portions. Teeth from Okladniko and Denisova caves have been attributed to Neanderthals (Goebel 1999), although subsequent DNA analysis showed that some findings attributed to Neanderthals actually belong to a distinct hominid, the Denisovans (Reich et al. 2010). Notably, a phalanx and two tooth found in the Denisova Cave were distinct from the Neanderthals and attributed to Denisovans. On the other hand, the discovery of a toe bone in 2011 in the Denisova Cave was preliminarily determined to belong to a Neanderthal, not a Denisovan (Gibbons 2011). In 2013, mitochondrial DNA from a 400,000 year old hominin femur bone from Spain, which had been seen as either Neanderthal or Homo heidelbergensis, was found to be closer to Denisovan mtDNA than to Neanderthal mtDNA (Callaway 2013).

Timeline

Skull, found in 1886 in Spy, Belgium
Frontal bone of a neanderthal child from the cave of La Garigüela
Skull from La Chapelle aux Saints
Semi-frontal view of a neanderthal skull from Gibraltar
  • 1829: Neanderthal skulls were discovered in Engis, in present-day Belgium.
  • 1848: Neanderthal skull Gibraltar 1 found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time.
  • 1856: Johann Karl Fuhlrott first recognized the fossil called "Neanderthal man", discovered in Neanderthal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany.
  • 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
  • 1886: Two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft with numerous Mousterian-type implements.
  • 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones.
  • 1899: Sand excavation workers found bone fragments on a hill in Krapina, Croatia called Hušnjakovo brdo. Local Franciscan friar Dominik Antolković requested Dragutin Gorjanović-Kramberger to study the remains of bones and teeth that were found there.
  • 1905: During the excavation in Krapina more than 5 000 items were found, of which 874 residue of human origin, including bones of prehistoric man and animals, artifacts.
  • 1908: A nearly complete Neanderthal skeleton was discovered in association with Mousterian tools and bones of extinct animals.
  • 1925: Francis Turville-Petre finds the 'Galilee Man' or 'Galilee Skull' in the Zuttiyeh Cave in Wadi Amud in The British Mandate of Palestine (now Israel).
  • 1926 Skull fragments of Gibraltar 2, a four-year-old Neanderthal girl, discovered by Dorothy Garrod.
  • 1953–1957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in the Kurdistan region of northern Iraq.
  • 1975: Erik Trinkaus' study of Neanderthal feet confirmed they walked like modern humans.
  • 1987: Thermoluminescence results from Israeli fossils date Neanderthals at Kebara to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by electron spin resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP).
  • 1991: ESR dates showed the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh.
  • 1993: A 127.000 years old DNA is found on the child of Sclayn, found in Scladina (fr), Belgium.
  • 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley (Kriegs et al. 1997).
  • 1998: A team led by pre-history archeologist João Zilhão discovered an early Upper Paleolithic human burial in Portugal, at Abrigo do Lagar Velho, which provided evidence of early modern humans from the west of the Iberian Peninsula. The remains, a largely complete skeleton of an approximately 4-year-old child, buried with pierced shell and red ochre, is dated to ca. 24,500 years BP (Duarte et al. 1999). The cranium, mandible, dentition, and postcrania present a mosaic of European early modern human and Neanderthal features (Duarte et al. 1999).
  • 2000: Igor Ovchinnikov, Kirsten Liden, William Goodman et al. retrieved DNA from a Late Neanderthal (29,000 BP) infant from Mezmaiskaya Cave in the Caucasus (Ovchinnikov et al. 2000).
  • 2005: The Max Planck Institute for Evolutionary Anthropology launched a project to reconstruct the Neanderthal genome. In 2009, the Max Planck Institute announced the "first draft" of a complete Neanderthal genome is completed.
  • 2010: Draft sequence of the Neanderthal genome, taken from three individuals, is published (Green et al. 2010). Comparison of this draft Neanderthal genome with modern humans from Africa and Eurasia shows that 1–4% of modern non-African human genome might come from the Neanderthals (Green et al. 2010).
  • 2010: Discovery of Neanderthal tools believed to be far away from the influence of H. sapiens suggest that the species might have been able to create and evolve tools on its own, and therefore be more intelligent than previously thought. Furthermore, it was proposed that the Neanderthals might be more closely related to Homo sapiens than previously thought and that may in fact be a subspecies of it. However, evidence of a more recent nature suggest that these artifacts are probably of H. sapiens sapiens origin (Benazzi et. al. 2011).
  • 2012: Charcoal found next to six paintings of seals in Nerja caves, Malaga, Spain, has been dated to between 42,300 and 43,500 years old, making them perhaps the oldest known cave paintings. José Luis Sanchidrián at the University of Cordoba, Spain believes the paintings are more likely to have been painted by Neanderthals than early modern humans (MacErlean 2012).
  • 2013: A jawbone found in Italy had features intermediate between Neanderthals and Homo sapiens suggesting it could be a hybrid. The mitochondrial DNA is Neanderthal (Viegas 2013).
  • 2013: An international team of researchers reported evidence that Neanderthals practiced burial behavior and intentionally buried their dead (Wilford 2013b).
  • 2013. The complete genome of a toe fossil of a 130,000-year-old Neanderthal, taken from a Siberian cave, is sequenced to a high degree of accuracy and published in Nature (Zimmer 2013; Prüfer et al. 2013).

Specimens

The Ferrassie skull.
  • Neanderthal 1: The first Neanderthal specimen found during an archaeological dig in August 1856. It was discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, three right arm bones, two left arm bones, ilium, and fragments of a scapula and ribs.
  • La Chapelle-aux-Saints 1: Called the Old Man, a fossilized skull discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth, with evidence of healing. For him to have lived on suggests that someone process his food for him, one of the earliest examples of Neanderthal altruism (similar to Shanidar I.)
  • La Ferrassie 1: A fossilized skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
  • Le Moustier: A fossilized skull, discovered in 1909, at the archaeological site in Peyzac-le-Moustier, Dordogne, France. The Mousterian tool culture is named after Le Moustier. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a somewhat less developed brow ridge and occipital bun as might be expected in a juvenile.
Type Specimen, Neanderthal 1
  • Shanidar 1: Found in the Zagros Mountains in (Iraqi Kurdistan); a total of nine skeletons found believed to have lived in the Middle Paleolithic. One of the nine remains was missing part of its right arm, which is theorized to have been broken off or amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One of the skeletons was buried with flowers, signifying that some type of burial ceremony may have occurred.

Chronology

Bones with Neanderthal traits in chronological order. (Sorted by time)

Mixed with H. heidelbergensis traits
  • > 350 ka: Sima de los Huesos c. 500:350 ka ago (Bischoff et al. 2003; Arsuaga et al. 1997)
  • 350–200 ka: Pontnewydd 225 ka ago.
  • 200–135 ka: Atapuerca (Kreger 2009), Vértesszőlős, Ehringsdorf, Casal de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade
Typical H. neanderthalensis traits
  • 135–45 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov Altai, Pech de l'Azé, Tabun 120 ka – 100±5 ka (Mcdermott et al. 1993), Qafzeh9 100, Shanidar 1 to 9 80–60 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 53±8 ka (Rink et al. 2002; Valladas et al. 1999), Teshik-Tash.
  • 45–35 ka: Le Moustier 45 ka, Feldhofer 42 ka, La Quina, l'Horus, Hortus, Kulna, Šipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, Arcy-sur-Cure (Rincon 2006).
  • < 35 ka: Châtelperron, Figueira Brava, Zafarraya 30 ka (Rincon 2006), Vogelherd 3?,Template:Vs (Conard et al. 2004) Vindija 32,400 ± 800 14C B.P. (Higham et al. 2006) (Vi-208 31,390 ± 220, Vi-207 32,400 ± 1,800 14C B.P.) (Higham et al. 2006), Velika Pećina,
Homo sapiens with some neanderthal-like archaic traits
  • < 35 Pestera cu Oase 35 ka, Mladeč 31 ka, Pestera Muierii 30 ka (n/s) (Hayes 2006), Lapedo Child 24.5 ka.

DNA analysis

Determining the DNA sequence of mitochondrial DNA (mtDNA) and nuclear DNA in organisms provides a useful tool to elucidate the evolutionary relationships among species. In general, closely related organisms have a high degree of agreement in the molecular structure of these substances, while the molecules of organisms distantly related usually show a pattern of dissimilarity. Mitochondrial DNA in hominids is inherited from the mother (maternally inherited) and there is usually no change in mtDNA from parent to offspring, although it does recombine with copies of itself within the same mitochondrion and there is a mutation rate, which is generally higher than that of nuclear DNA. The mtDNA is useful for tracking ancestry through females (matrilineage). Nuclear DNA is inherited from both parents and genes are rearranged in the process of recombination. Conserved sequences, such as mitochondrial DNA, are expected to accumulate mutations over time, and assuming a constant rate of mutation provides a molecular clock for dating divergence The DNA sequence of mtDNA and nuclear DNA has been determined from a large number of species (including some organisms that are extinct), and the comparison of those DNA sequences represents a mainstay of phylogenetics.

However, for most fossils of ancient hominin species, the DNA cannot be recovered, because it degrades over the long time periods, and this degradation increases with temperature and such conditions as acidic soil. Most early hominin fossils are from tropical and equatorial regions where conditions for survival are poor. Thus, DNA sequences to date have not been recovered from Homo erectus, H. heidelbergensis, or H. antecessor (Krause et al. 2010).

However, mitochondrial DNA and nuclear DNA has been obtained from Neanderthals, as well as Denisovans and modern humans. This allows insight on their evolutionary relationships.

Early investigations concentrated on mitochondrial DNA (mtDNA). Owing to the strictly matrilineal inheritance and subsequent vulnerability to genetic drift of mtDNA, it is of limited value in evaluating the possibility of interbreeding of Neanderthals with modern human beings (specifically, with []Cro-Magnon]] people, the name applied to the earliest known examples of modern human beings, Homo sapiens sapiens, which lived from about 40,000 to 10,000 years ago in the Upper Paleolithic period of the Pleistocene epoch).

In 1997, geneticists were able to extract a short sequence of maternally inherited mitochondrial DNA (mtDNA) from the Neanderthal type specimen found at Feldhofer from 30,000 years ago (Brown 2008; Krings et al. 1997). Comparison with the mtDNAs of modern humans suggested that the Neanderthal mtDNA shared a common ancestor with present-day human mtDNA about half a million years ago (Krings et al. 1997; Krings et al. 1999). Subsequently, many additional mtDNA sequences were retrieved and analyzed from Neanderthal fossils (Green et al. 2006). Based on these findings and morphological evidence, Green et al. concluded in 2006 that "if any genetic contribution of Neanderthals to modern human occurred, it was of limited magnitude" (Green et al. 2006). In 2010, Ovchinnikov et al. reported the extraction of mtDNA from a specimen in which they concluded the sequence showed no sign of modern human descent from Neanderthals (Ovchinnikov et al. 2000).

In 2006, Green, Krause, Ptak, et al. completed a preliminary sequencing (one million base pairs) of the nuclear DNA from a 38,000-year-old Neanderthal fossil, which was the bone fragment of a femur found in 1980 at Vindija Cave, Croatia. They also examined the mitochondrial DNA. These researchers were from the Max-Planck Institute for Evolutionary Anthropology (Germany), 454 Life Sciences (United States), and the Institute of Quaternary Paleontology and Geology, Croatian Academy of Sciences and Arts (Croatia). They reported that comparison with the genome of modern humans (and chimpanzees) revealed that Neanderthal and modern human DNA sequences diverged about 500,000 years ago. Neanderthals and modern humans were also estimated to share about 99.5% of their DNA (Green et al. 2006). Notable, other researchers suggested a time of divergence based on fossil records of 400,000 years ago (Wade 2006a) and a 2007 study pushed the point of divergence back to around 800,000 years ago (Pennisi 2007).

Scientist at the Max Planck Institute for Evolutionary Anthropology extracting the DNA.

In July 2006, the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany and 454 Life Sciences in Connecticut, USA announced that they would sequence the Neanderthal genome over the next two years. It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains (Moulson 2006). The study was lead by Svante Pääbo, who was director of the Department of Genetics at the Max Planck Institute for Evolutionary Anthropology since 1997.

In 2008, Green et al. from Max Planck Institute for Evolutionary Anthropology published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested "Neanderthals had a long-term effective population size smaller than that of modern humans" (Green et al. 2008). They also determined based on the mtDNA comparisons between modern humans and Neanderthals a "divergence date between the two mtDNA lineages of 660,000 ± 140,000 years" (Green et al. 2008).

In February 2009, at the Annual Meeting of the American Association for the Advancement of Science (AAAS), it was announced that the Max Planck Institute for Evolutionary Anthropology, in collaboration with the 454 Life Sciences Corporation, had completed the first draft version of the Neanderthal genome.

In May 2010, the above-noted draft sequence of the Neanderthal genome was published in the journal Science by Green et al., who noted that the draft sequence was composed of more than 4 billion nucleotides from three individuals. An analysis of this first draft of the Neanderthal genome indicated that interbreeding between Neanderthals and modern humans may have occurred. "Those of us who live outside Africa carry a little Neanderthal DNA in us," said Pääbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School, who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea. The finding is that about 1 to 4 percent of the genes of the non-Africans came from Neanderthals, compared to the baseline defined by the two Africans (Green et al. 2010).

This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa. No evidence for gene flow in the direction from modern humans to Neanderthals was found. Gene flow from modern humans to Neanderthals would not be expected if contact occurred between a small colonizing population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonization process (Green et al. 2010).

While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely related to Neanderthals than other Africans were, due to ancient genetic divisions within Africa (Green et al. 2010).

In 2013, Prüfer et al. reported the sequencing to a high degree of accuracy of the complete genome of a toe fossil of a 130,000-year-old Neanderthal, taken from a Siberian cave (Zimmer 2013; Prüfer et al. 2013). In addition, a low coverage sequencing was done for the genome of a Neanderthal from the Caucasus. Prüfer et al.(2013) reported that a comparison with available genomes of archaic and present-day humans revealed several gene flow events among early modern humans, Denisovans, and Neanderthals. They also were able to establish a "definitive list of substitutions that became fixed in modern humans after their separation from the ancestors of Neanderthals and Denisovans."

Anatomy

Neanderthal cranial anatomy
Comparison of crania, sapiens (left) and neanderthalensis (right)

Neanderthals had a compact body of short stature. On average, Neanderthal males stood about 1.64 meters tall (5' 5") and were heavily built with robust bone structure (Smithsonian 2014b). Females were about 1.55 meters tall (about 5'1") (Smithsonian 2014b). Males were an estimated average of 65 kg (143 lb.), and females averaged an estimated 54 kg (119lb) (Smithsonian 2014b).

Neanderthal brain sizes have been estimated to be larger than modern humans, although such estimates have not been adjusted for their more robust builds; their brain sizes are proportional to their more robust frame (Smithsonian 2014b).. Neanderthals also had a large cranial capacity, estimated at 1500cc, which is slightly larger on average than modern humans (1350 to 1450cc range).

Neanderthals also differed from modern humans in that they had a low forehead, double-arched brow ridge, larger nasal area, projecting cheek region, weak chin, obvious space behind the third molar, heavily-built bones, broad scapula, short lower leg and arm bones relative to the upper portions, occasional bowing of the limb bones, the hip joint rotated outward, a long and thin pubic bone, and large joint surfaces of the toes and long bones (Smithsonian 2014b).

The following is a list of physical traits that distinguish Neanderthals from modern humans; however, not all of them can be used to distinguish specific Neanderthal populations, from various geographic areas or periods of evolution, from other extinct humans. Also, many of these traits occasionally manifest in modern humans, particularly among certain ethnic groups. Nothing is known about the skin color, the hair, or the shape of soft parts such as eyes, ears, and lips of Neanderthals (Carey 2005).

Neanderthal physical traits
Cranial Sub-cranial
Suprainiac fossa, a groove above the inion Considerably more robust
Occipital bun, a protuberance of the occipital bone that looks like a hair knot Large round finger tips
Projecting mid-face Barrel-shaped rib cage
Low, flat, elongated skull Large kneecaps
A flat basic cranium Long collar bones
Supraorbital torus, a prominent, trabecular (spongy) browridge Short, bowed shoulder blades
1200-1750 cm³ skull capacity (10 percent greater than modern human average) Thick, bowed shaft of the thigh bones
Lack of a protruding chin (mental protuberance; although later specimens possess a slight protuberance) Short shinbones and calf bones
Crest on the mastoid process behind the ear opening Long, gracile pelvic pubis (superior pubic ramus)
No groove on canine teeth
A retromolar space posterior to the third molar
Bony projections on the sides of the nasal opening
Distinctive shape of the bony labyrinth in the ear
Larger mental foramen in mandible for facial blood supply
A broad, projecting nose

Neanderthals appear to have had many adaptations to a cold climate, such as large brain cases, short but robust builds, and large noses.

Classification and origin

For many years, professionals have vigorously debated whether Neanderthals should be classified as Homo neanderthalensis or as Homo sapiens neanderthalensis, the latter placing Neanderthals as a subspecies of Homo sapiens.

The original reconstruction of Neanderthal anatomy was flawed and exacerbated the distinction between Neanderthals and humans. Based on a nearly complete skeleton of an elderly male found in France, the reconstruction showed bent knees and a slouching gait (Smithsonian 2014b). This image, which led to a standard and lingering view of crude cavemen, was mistaken, as Neanderthals apparently walked fully upright without a slouch or bent knees. They also had a larger cranial capacity than modern humans and were culturally sophisticated in terms of tool making, symbolic ritual, and seemingly burying their dead (Smithsonian 2014b).

However, there are many differences between Neanderthals and modern populations. They had a low forehead, double-arched brow ridge, larger nasal area, projecting cheek region, heavily-built bones, short lower leg and arm bones relative to the upper bones, and so forth.

The first proto-Neanderthal traits appeared in Europe as early as 350,000 years ago (Bischoff et al. 2003). By 130,000 years ago, full blown Neanderthal characteristics were present. Both the Neanderthal's place in the human family tree and their relation to modern Europeans have been hotly debated ever since their discovery. A common perspective among scientists, based on ongoing DNA research, is that Neanderthals were a separate branch of the genus Homo, and that modern humans are not descended from them (fitting with the single-origin thesis).

Comparison of the DNA of Neanderthals and Homo sapiens suggests that they diverged from a common ancestor between 350,000 and 500,000 years ago. This was probably Homo heidelbergensis. Heidelbergensis originated between 800,000 and 1,300,000 years ago, and continued until about 200,000. It ranged over east and South Africa, Europe and west Asia. Between 350,000 and 400,000 years ago the African branch is thought to have started evolving towards modern humans and the European branch towards Neanderthals. Mayr claims that Neanderthals arose from Homo erectus, arguing, "There is little doubt that…the western populations of H. erectus eventually gave rise to the Neanderthals" (2001).

Scientists do not agree when Neanderthals can first be recognized in the fossil record, with dates typically ranging between 200,000 and 300,000 years before present (Smithsonian 2014b; Stringer 2011; Johansson and Edgar 2006) and Green et al. (2010) stating that the first European fossils of Neanderthal date to 400,000 years ago.

Habitat and range

Sites where typical Neanderthal fossils have been found

Early Neanderthals lived in the Last glacial period for a span of about 100,000 years. Because of the damaging effects the glacial period had on the Neanderthal sites, not much is known about the early species.

Classic Neanderthal fossils have been found over a large area of Europe and Western Asia, including as far south as the Middle East and far east as southern Siberia (Green et al. 2010). Countries where Neanderthal remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north, including most of Western Europe, including the south coast of Great Britain (Dargie 2007), Central Europe, the Carpathians, and the Balkans (Durham 2008), some sites in Ukraine and in western Russia. Neanderthal remains also have been found east of Europe in Siberia to the Altai Mountains and south through the Levant to the Indus River. Sites have been found in northern Germany, Israel, Spain (Arsuaga et al. 1989), Italy (Mallegni et al. 1987), Portugal, and Uzbekistan, and east into southern Siberia's Altai Mountains (Wade 2007b; Ravilious 2007). On the other hand, in 2013, mitochondrial DNA was extracted from a 300,000- to 400,000-year-old sliver of hominin femur bone from Spain, which had been seen as either Neanderthal or Homo heidelbergensis, and the retrieval of an almost complete mitochondrial genome yielded the surprising result of ancestral Denisonian DNA (Callaway 2013; Gibbons 2013).

This area probably was not occupied all at the same time. The northern border of their range, in particular, would have contracted frequently with the onset of cold periods. On the other hand, the northern border of their range as represented by fossils may not be the real northern border of the area they occupied, since Middle Palaeolithic-looking artifacts have been found even further north, up to 60° N, on the Russian plain (Pavlov et al. 2004). It is estimated that the total Neanderthal population across this habitat range numbered at around 70,000 at its peak (O'Neil 2013a).

Neanderthal fossils have not been found to date in Africa, but there have been finds rather close to North Africa, both on Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains, in fact, date from after the same sites were vacated by modern humans. Mammal fossils of the same time period show cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than modern humans and at times displaced them in parts of the Middle East when the climate got cold enough (Jordan 2001).

Homo sapiens sapiens appears to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived south-west of present-day Israel. When further climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before modern people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before modern people totally replaced the Neanderthal, despite the recent evidence of some successful interbreeding (Jordan 2001).

There were separate developments in the human line, in other regions such as Southern Africa, that somewhat resembled the European and Western/Central Asian Neanderthals, but these people were not actually Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis) who existed long before any classic European Neanderthals, but had a more modern set of teeth, and arguably some H. rhodesiensis populations were on the road to modern H. sapiens sapiens. At any rate, the populations in Europe and West/Central Asia underwent more and more "Neanderthalization" as time went on. There is some argument that H. rhodesiensis in general was ancestral to both modern humans and Neanderthals, and that at some point the two populations went their separate ways, but this supposes that H. rhodesiensis goes back to around 600,000 years ago.

To date, no intimate connection has been found between these similar archaic people and the Western/Central Eurasian Neanderthals, at least during the same time as classic Eurasian Neanderthals, and H. rhodesiensis seems to have lived about 600,000 years ago, long before the time of classic Neanderthals. This said, some researchers think that H. rhodesiensis may have lived much later than this period, depending on the method used to date the fossils, leaving this issue open to debate. Some H. rhodesiensis features, like the large brow ridge, may have been caused by convergent evolution.

It appears incorrect, based on present research and known fossil finds, to refer to any fossils outside Europe or Western and Central Asia as true Neanderthals. While they had a known range that extended as far east as the Altai Mountains, it was not found to extend farther to the east and apparently not into Africa. At any rate, in North-East Africa the land immediately south of the Neanderthal range was possessed by modern humans Homo sapiens idaltu or Homo sapiens, since at least 160,000 years before the present. About 160,000-year-old hominid fossils at Jebel Irhoud in Morocco were previously thought to be Neanderthal, but it is now clear that they are early modern humans (Max Planck Institute 2007).

Tools, burial, and other cultural aspects

Neanderthals made finer tools than earlier humans, lived in shelters, controlled fire, wore clothing, and are considered the first to bury their dead and to have symbolic ritual (Smithsonian 2014b). No earlier human species has been show to practice such symbolic and sophisticated behavior (Smithsonian 2014b). This practice of intentional burial is one reason given for the finding of so many Neanderthal fossils, including skeletons.

Neanderthal Tool Maker

Neanderthal (Middle Paleolithic) archaeological sites show a different, smaller toolkit than those that have been found in Upper Paleolithic sites, which were perhaps occupied by the modern humans that superseded them. Fossil evidence indicating who may have made the tools found in Early Upper Paleolithic sites is inconclusive.

The characteristic style of stone tools in the Middle Paleolithic is called the Mousterian culture, after a prominent archaeological site where the tools were first found. They typically used the Levallois technique. Mousterian tools were often produced using soft hammer percussion, with hammers made of materials like bones, antlers, and wood, rather than hard hammer percussion, using stone hammers. This technique involves the detaching of flake tools from a prepared stone core in such a way that allowed flakes of predetermined shape to be removed and prepared into tools from a single suitable stone (Smithsonian 2014b). The Mousterian stone tool technology differs from the "core tool" traditions such as the Archeulean technology of Homo erectus, which involved working from a stone that was chipped down to the tool form by removal of flakes from the surface (Smithsonian 2014b). Near the end of the time of the Neanderthals, they utilized the Châtelperronian tool style, which is considered more advanced than that of the Mousterian. They either invented the Châtelperronian themselves or borrowed elements from the incoming modern humans who are thought to have created the Aurignacian style.

The Mousterian flake and simple biface industry that characterize the Middle Paleolithic, wherever found with human remains, are found with Neanderthals, and wherever Aurignacian style is found with remains, those remains are of modern humans (West 1996). The Neanderthal (Mousterian) toolkits consisted of sophisticated stone-flakes, task-specific hand axes, and spears. Many of these tools were very sharp. There is also good evidence that they used a lot of wood, although such artifacts would likely not have been preserved (Henig 2000). Neanderthals were capable of building dugout boats since the Middle Paleolithic. Mousterian stone tools discovered on the southern Ionian Greek islands suggests that Neanderthals were sailing the Mediterranean Sea as early as 110,000 years BP.

Middle Paleolithic industries in Siberia (dated to 70,000 to 40,000 years ago) are distinctly Levallois and Mousterian, reduction technologies are uniform, and assemblages consist of scrapers, denticulates, notches, knives, and retouched Levallois flakes and points. There is no evidence of bone, antler, or ivory technology, or of art or personal adornment (Goebel 1999).

While Neanderthals had weapons, no projectile weapons have yet been found. They had spears, in the sense of a long wooden shaft with a spearhead firmly attached to it, but these were not spears specifically crafted for flight (such as a javelin). However, a number of 400,000 year old wooden projectile spears were found at Schöningen in northern Germany. These are thought to have been made by one of Neanderthal's ancestors, either Homo erectus or Homo heidelbergensis. Generally, projectile weapons are more commonly associated with H. sapiens. The lack of projectile weaponry is an indication of different sustenance methods, rather than inferior technology or abilities. The situation is identical to that of native New Zealand Maoris—modern Homo sapiens who also rarely threw objects, but used spears and clubs instead (Schwimmer 1961).

Although much has been made of the Neanderthal's burial of their dead, their burials were less elaborate than those of anatomically modern humans. The interpretation of the Shanidar IV burials as including flowers, and therefore being a form of ritual burial (Solecki 1975), has been questioned (Sommer 1999). On the other hand, five of the six flower pollens found with fossil Shanidar IV are known to have had traditional medical uses, even among relatively contemporary populations. In some cases Neanderthal burials include grave goods, such as bison and auroch bones, tools, and the pigment ochre.

Neanderthals performed a sophisticated set of tasks normally associated with humans alone. For example, they constructed complex shelters, controlled fire, and skinned animals. Particularly intriguing is a hollowed-out bear femur that contains holes that may have been deliberately bored into it. This bone was found in western Slovenia in 1995, near a Mousterian fireplace, but its significance is still a matter of dispute. Some paleoanthropologists have postulated that it might have been a flute, while some others have expressed that it is natural bone modified by bears.

Language

The issue of whether Neanderthals had a complex language is unsettled, but there are morphological suggestions that such was possible. There is some circumstantial evidence for thinking that Neanderthals had language with words and some kind of syntax; some of their tool-making and hunting tactics would have been difficult to learn and execute without it. A recent extraction of DNA from Neanderthal bones indicates that Neanderthals had the same version of the FOXP2 gene as modern humans. This gene is known to play a role in human language (Wade 2007a).

The idea that Neanderthals lacked complex language was once widespread (Lieberman and Crelin 1971), despite concerns about the accuracy of reconstructions of the Neanderthal vocal tract, until 1983, when a Neanderthal hyoid bone was found at the Kebara Cave in Israel. The hyoid is a small bone that connects the musculature of the tongue and the larynx, and by bracing these structures against each other, allows a wider range of tongue and laryngeal movements than would otherwise be possible. Therefore, it seems to imply the presence of the anatomical conditions for speech to occur. The bone that was found is virtually identical to that of modern humans (Arensburg et al. 1989).

Furthermore, the morphology of the outer and middle ear of Neanderthal ancestors, Homo heidelbergensis, found in Spain, suggests they had an auditory sensitivity similar to modern humans and very different from chimpanzees. Therefore, they were not only able to produce a wide range of sounds, they were also able to differentiate between these sounds (Martinez et al. 2004).

Aside from the morphological evidence above, neurological evidence for potential speech in neanderthalensis exists in the form of the hypoglossal canal. The canal of Neanderthals is the same size or larger than in modern humans, which are significantly larger than the canal of australopithecines and modern chimpanzees. The canal carries the hypoglossal nerve, which supplies the muscles of the tongue with motor coordination. Researchers indicate that this evidence suggests that neanderthalensis had vocal capabilities similar to, or possibly exceeding that of, modern humans (Kay et al. 1998). However, a research team from the University of California, Berkeley, led by David DeGusta, suggests that the size of the hypoglossal canal is not an indicator of speech. His team's research, which shows no correlation between canal size and speech potential, shows there are a number of extant (living) non-human primates and fossilized australopithecines that have equal or larger hypoglossal canal.

Many people believe that even without the hyoid bone evidence, tools as advanced as those of the Mousterian Era, attributed to Neanderthals, could not have been developed without cognitive skills capable of encompassing some form of spoken language.

Many myths surround the reconstruction of the Neanderthal vocal tract and the quality of Neanderthal speech. The popular view that the Neanderthals had a high larynx and therefore could not have produced the range of vowels supposedly essential for human speech is based on a disputed reconstruction of the vocal tract from the available fossil evidence, and a debatable interpretation of the acoustic characteristics of the reconstructed vocal tract. A larynx position as low as that found for modern human females may have been present in adult male Neanderthals. Furthermore, the vocal tract is a plastic thing, and larynx movement is possible in many mammals. Finally, the suggestion that the vowels /i, a, u/ are essential for human language (and that if Neanderthals lacked them, they could not have evolved a human-like language) ignores the absence of one of these vowels in very many human languages, and the occurrence of "vertical vowel systems" which lack both /i/ and /u/.

More doubtful suggestions about Neanderthal speech suggest that it would have been nasalized either because the tongue was high in the throat (for which there is no universally accepted evidence) or because the Neanderthals had large nasal cavities. Nasalization depends on neither of these things, but on whether or not the soft palate is lowered during speech. Nasalization is therefore controllable, and scientists do not know whether Neanderthal speech was nasalized or not. Comments on the lower intelligibility of nasalized speech ignore the fact that many varieties of English habitually have nasalized vowels, particularly low vowels, with no apparent effect on intelligibility.

Another anatomical difference between Neanderthals and modern humans is the former's general lack of a mental protuberance (the point at the tip of the chin). This may be relevant to speech, as the mentalis muscle contributes to moving the lower lip and is used to articulate a bilabial click. While some Neanderthal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans (Schwartz and Tattersall 2000). In contrast, some Neanderthal individuals show inferior lateral mental tubercles (little bumps at the side of the chin).

Steven Mithen (2006) speculates that the Neanderthals may have had an elaborate proto-linguistic system of communication that was more musical than modern human language, and that pre-dated the separation of language and music into two separate modes of cognition. He called this hypothetical lingual system 'hmmmmm' because it would be Holistic, manipulative, multi-modal, musical and mimetic (Mithen 2006).

The fate of the Neanderthals

Neanderthals became extinct in Europe approximately 30,000 years ago. There is recently discovered fossil and stone-tool evidence that suggests Neanderthals may have still been in existence 24,000 to 28,000 years ago, at which time they they disappeared from the fossil record and were replaced in Europe by modern Homo sapiens (Rincon 2006, Mcilroy 2006, Klein 2003, Smithsonian 2014b).

The Neanderthals began to be displaced around 45,000 years ago by modern humans (Homo sapiens), as the Cro-Magnon people appeared in Europe. Despite this, populations of Neanderthals apparently held on for thousands of years in regional pockets, such as modern-day Croatia and the Iberian and Crimean peninsulas. The last known population lived around a cave system on the remote south-facing coast of Gibraltar, from 30,000 to 24,000 years ago.

There is considerable debate about whether Cro-Magnon people accelerated the demise of the Neanderthals. Timing suggests a causal relation between the appearance of Homo sapiens in Europe and the decline of Homo neanderthalensis.

In some areas of the Middle East and the Iberian peninsula, Neanderthals did, in fact, apparently co-exist side by side with populations of anatomically modern Homo sapiens for roughly 10,000 years. There is also evidence that it is in these areas where the last of the Neanderthals died out and that during this period the last remnants of this species had begun to adopt—or perhaps independently innovate—some aspects of the Châtelperronian (Upper Paleolithic) tool case, which is usually exclusively associated with anatomically modern Homo sapiens.

There are various scenarios for the extinction of Neanderthals.

Extinction scenarios

Extinction caused by modern humans

Jared Diamond has suggested a scenario of violent conflict, comparable to the genocides suffered by indigenous peoples in recent human history.

Another possibility paralleling colonialist history would be a greater susceptibility to pathogens introduced by Cro-Magnon man on the part of the Neanderthals. Although Diamond and others have specifically mentioned Cro-Magnon diseases as a threat to Neanderthals, this aspect of the analogy with the contacts between colonizers and indigenous peoples in recent history can be misleading. The distinction arises because Cro-Magnons and Neanderthals are both believed to have lived a nomadic lifestyle, whereas those genocides of the colonial era, in which differential disease susceptibility was most significant, resulted from the contact between colonists with a long history of agriculture and nomadic hunter-gatherer peoples.

On the other hand, many pre-European contact Native Americans were not nomadic, but agriculturalists, such as Mayans, Iroquois, and Cherokee, and this still did not protect them from the epidemics brought by Europeans (notably, smallpox). One theory is that because they usually lacked large domesticated animal agriculture, such as cattle or pigs in close contact with people, they did not develop resistance to species-jumping diseases like Europeans had. Furthermore, the nomadic Eurasian populations, such as the Mongols, did not get wiped out by the diseases of the agriculturalist societies they invaded and took over, like China and eastern Europe.

There are also gradual extinction scenarios due to competition from modern humans that are proposed to account for the decline of Neanderthal population. For example, according to Banks et al. (2008), even a slight competitive advantage on the part of modern humans could account for Neanderthals' replacement by anatomically modern humans on a timescale of 10,000–20,000 years. Competitive disadvantages due to anatomical differences, tools, and division of labor are among the issues proposed. For example, in 2006, anthropologists Steven L. Kuhn and Mary C. Stiner of the University of Arizona proposed proposed an explanation based on division of labor (Wade 2006b). In an article titled "What's a Mother to Do? The Division of Labor among Neanderthals and Modern Humans in Eurasia," Kuhn and Stiner theorize that Neanderthals did not have a division of labor between the sexes (2006). Both male and female Neanderthals participated in the single main occupation of hunting the big game that flourished in Europe during the ice age, like bison, deer, gazelles, and wild horses. This contrasted with humans who were better able to use the resources of the environment because of a division of labor with the women going after small game and gathering plant foods. In addition, because big game hunting was so dangerous, this made humans, at least females, more resilient. Pat Shipman, from Pennsylvania State University, argues that the domestication of the dog gave modern humans an advantage when hunting (Shipman 2012).

The problem with a gradual extinction scenario lies in the resolution of dating methods. There have been claims for young Neanderthal sites, younger than 30,000 years old (Finlayson et al. 2006). Even claims for interstratification of Neanderthal and modern human remains have been advanced (Gravina et al. 2005). So the fact that Neanderthals and modern humans coexisted at least for some time seems certain. However, because of difficulties in calibrating the C14 dates, the duration of this period is uncertain (Mellars 2006a).

Assimilation with modern humans

There have been claims both that Neanderthals assimilated with modern human beings and that they did not assimilate.

It is possible that the Neanderthals, with their small numbers, could have been absorbed by the much larger populations of modern Homo sapiens. In November 2006, a paper was published in the United States journal Proceedings of the National Academy of Sciences, in which a team of European researchers suggest that Neanderthals and humans interbred, citing distinct human and Neanderthal features in a 30,000 year-old fossil found in Romania. Co-author Erik Trinkaus from Washington University explains, "Closely related species of mammals freely interbreed, produce fertile viable offspring and blend populations. Extinction through absorption is a common phenomenon" (Hayes 2006). Genetic studies, such as that of Green et al. (2010), does suggest some interbreeding, although the contribution of Neanderthals to human DNA is small.

Unable to adapt

Shows the pattern of temperature and ice volume changes associated with recent glacials and interglacials

European populations of H. neanderthalensis were adapted for a cold environment. One view on their extinction is that they may have had problems adapting to a warming environment. The problem with this idea is that the glacial period of our ice age ended about 10,000 years ago, while the Neanderthals went extinct about 24,000 years ago.

Another possibility has to do with the loss of the Neanderthal's primary hunting territory: forests. It is speculated that their hunting methods (stabbing prey with spears rather than throwing the spears) and lack of mobility could have placed them at a disadvantage when the forests were replaced by flat lands. It is also suggested that they mainly ate meat, and thus were less adaptable. Homo sapiens, which hunted large prey but did not depend on them for survival, may have indirectly contributed to their extinction this way.

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