
Introduction
Sexual dimorphism (genetic/morphological differences between the sexes) is incredibly common throughout primates, and our ancestors were no exception to this. In primates, especially apes, males are typically much larger than females, along with other differences, such as larger maxillary (upper) canines. At its greatest extent, primates can be up to 50% sexually dimorphic. Modern humans are much less sexually dimorphic as compared to our relatives, and we usually are around 15% sexually dimorphic. Sexual dimorphism likely began decreasing in humans around 2 million years ago, though some earlier hominins, such as Ardipithecus ramidus had low levels of sexual dimorphism at 4.4 million years ago.
Sexual dimorphism in later hominins, in the genus Australopithecus, is controversial. Research on the femora of Australopithecus afarensis and Australopithecus africanus suggests that these species had very high levels of sexual dimorphism, similar to what is seen in modern gorillas and orangutans. If this is accurate, female australopiths may have had different locomotor styles (methods of walking) than males due to their sexual differences.
Other research suggests that Australopithecus had little levels of sexual dimorphism, closer to what is seen in modern humans. As it stands right now, it appears that Australopithecus did express some levels of sexual dimorphism, though this is subject to change as new evidence comes about. However, no matter how sexually dimorphic our australopith ancestors were, there is one hominin genus that was clearly very sexually dimorphic, and that is Paranthropus.
What is Paranthropus?
Paranthropus is a unique genus of hominin which lived from 2.5-1.4 million years ago. Members of this genus are also commonly known as the ‘robust australopiths’ due to their odd and robust skeletal and cranial morphology. Paranthropus was originally considered to be in the genus Australopithecus, until it was decided that it was unique enough among other australopiths to be placed within its own genus, though some anthropologists still consider it to be in the Australopithecus genus.
Paranthropus is an off-branching lineage, and is not directly ancestral to modern humans. It likely branched off from Australopithecus afarensis or Australopithecus africanus sometime in the late Pliocene epoch. Paranthropus contains 3 known and recognized species: P. aethiopicus, P. boisei, and P. robustus. The former two lived throughout eastern Africa, while Paranthropus robustus lived in South Africa.
Perhaps its most unique trait was its diet. Paranthropus was seemingly mostly herbivorous. The eastern species had a diet mostly composed of C4 plants, meaning they were mostly consuming savannah grasses, while Paranthropus robustus had a slightly more varied diet, with a mix of C4 and C3 (forest) plants, and likely changed its diet seasonally. This diet was possibly a fall back diet, where the genus adapted for a more accessible diet, not a preferred diet.
Because of this unique diet, Paranthropus had some specialized morphology. This genus possessed a pronounced sagittal crest, (a bony crest on the top of the head), large round zygomatic arches (cheek bones), and incredibly large molars and thick tooth enamel, a condition known as post-canine megadontia.
All these traits gave Paranthropus a very powerful bite, perfect for the tough vegetation it was eating. This gave P. boisei the nickname “nutcracker man”, though analysis of tooth wear on Paranthropus teeth shows it wasn’t eating much tough food. Tougher meats, nuts, and seeds may occasionally have been a part of its diet however.
Perhaps one of the most interesting things about Paranthropus is their high levels of sexual dimorphism, which makes them stand out even among other living primates.
Sexual Dimorphism in Paranthropus
Paranthropus exhibited a great deal of sexual dimorphism. Female individuals were smaller in body size and weight, smaller in skull size, had smaller brains, teeth, and lacked the distinguishing sagittal crest. Male Paranthropus boisei were about 4 ft. 6 in. tall and weighed about 108 lbs while females were about 4 ft. tall and weighed about 75 lbs. Paranthropus robustus was slightly smaller with males being about 3 ft. 10 in. tall and weighing 120 lbs and females being about 3 ft tall and weighing about 90 lbs. Because of a lack of post cranial material from Paranthropus aethiopicus, its height and weight is unknown, but it was likely similar to that of P. boisei.
A Paranthropus molar from Gondolin, South Africa, known as GDA-2 also shows a great deal of sexual dimorphism in P. robustus. The size of the molar falls into the range of the larger P. boisei. This could possibly suggest that P. boisei had made it to southern Africa, or more likely, that P. robustus males had much larger teeth than females. Other teeth, found in Swartkrans, South Africa, are much smaller than what is expected in Paranthropus, suggesting that female individuals had smaller teeth.
Two individuals from P. boisei and P. robustus give great insight into sexual dimorphism in Paranthropus.
One female fossil specimen from Koobi Fora, Kenya, gives a better look into sexual dimorphism in Paranthropus boisei. The cranial specimen, known as KNM-ER-732, lacks a sagittal crest, and has a smaller cranial capacity than male individuals, though not by much. This individual had a cranial capacity of around 500 cc, while other male individuals, such as KNM-ER-406, had a cranial capacity of about 510 cc. The female specimen is also smaller in size overall than male individuals.

The next specimen which exhibits sexual dimorphism was found in Drimolen, South Africa, and belongs to Paranthropus robustus. This specimen, nicknamed Eurydice, formally known as DNH 7, is the most complete specimen of its species. Eurydice is significantly smaller in skull and tooth size compared to male specimens of her species, such as SK 46. Along with that, she also lacked a sagittal crest, showing high levels of sexual dimorphism. Interestingly, Eurydice is one of the only female specimens out of the many P. robustus specimens found at the site, potentially giving insight into the social structure of her species.

Paranthropus Social Structure
Sexual dimorphism often has a big role in the social structure of animals. Gibbons for example show very little sexual dimorphism, resulting in them having monogamous social structures. This means that most gibbons live in small family groups, with one male and one female. This activity is known as pair bonding. Some groups in China and Khao Yai, Thailand contain multiple adults, but for the most part, the low sexual dimorphism in gibbons results in monogamous groups, which is beneficial for both individuals in the family group.
When deciphering the social structures of Paranthropus, looking at modern primates with similar levels of sexual dimorphism can give good insight. For Paranthropus, one primate with similar levels of sexual dimorphism are gorillas.
Gorillas exhibit great levels of sexual dimorphism, with females being about half the size of males. This is similar to what has been observed in fossils of Paranthropus. Most gorillas have what is known as a harem society. This is a group containing one adult head male, several females, and their offspring. The adult male defends the females from foreign males, and intern, has breeding rights to all the females, unless he proves himself to be a poor leader in which case the females typically abandon the male and join a better group.
In some species of gorilla, such as the eastern gorilla (Gorilla beringei), the harem group contains several related males which defend the group. Due to the similar levels of sexual dimorphism, this harem society may be what Paranthropus used. This is further backed up by strontium isotope analyses of the holotype specimen of Paranthropus robustus, TM157c. When analyzing the teeth of this female specimen, anthropologists deduced that she had moved to her location of death before her teeth had fully mineralized, suggesting that female Paranthropus migrated less in their groups, which is behavior found in harem groups of gorillas.
However, there is one major difference between gorillas and Paranthropus, that is their environment. While modern gorillas live in more dense and lush jungles, where they can be protected and hidden by foliage, Paranthropus lived in open dry woodland and savannah habitats. A better modern comparison for Paranthropus social structure would instead be baboons. Baboons exhibit similar levels of sexual dimorphism, but live in more open dry savannah, similar to Paranthropus.
Baboons often live in large social groups with several males, females, and offspring. Typically, baboon groups consist of one primary male, several secondary males, and the females and offspring. This system allows for better protection in the open savannah as there are more males to defend the group. This type of group is known as a multilevel society.

However, this system often leads to lots of intolerance between males. Some baboon species, such as the Hamadryas baboon (Papio hamadryas) are especially aggressive, whereas others, such as Guinea baboons (Papio papio) are less aggressive. This could possibly mean that male Paranthropus were aggressive towards each other, and lived in very large social groups.
Sites throughout South Africa, such as Swartkrans and Kromdraai show an uneven sex ratio in Paranthropus robustus, with 15 male individuals being found and 4 female individuals, though this information may be inaccurate. If it is accurate, it may add further support to the idea that Paranthropus upheld multilevel societies, with many males and few females. It could also suggest a higher mortality rate in male Paranthropus, which would make sense under a harem or multilevel society, as males would be more likely to be kicked out of the group, making them more vulnerable.
Conclusion
Paranthropus, like other hominins and primates in general, exhibited a great amount of sexual dimorphism. Male individuals possessed larger bodies, brains, skulls, and teeth. This high level of sexual dimorphism would have had a large impact on the social structures and behavior. Due to the open woodland/savannah environment they lived in, Paranthropus likely formed large multilevel social groups, with a head male, and lower males and females, similar to what is seen in modern baboons. This system is beneficial for the group as it allows for more group protection. Further research into the sexual dimorphism and behavior of our ancient ancestors and relatives is very important as it can help us understand ourselves and our evolution.
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