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The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...
found in the
penis A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males d ...
of many
placental mammal Placental mammals ( infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguish ...
s. It is absent from the
human penis The human penis is an external male intromittent organ that additionally serves as the urinary duct. The main parts are the root (radix); the body (corpus); and the epithelium of the penis including the shaft skin and the foreskin (prepuce) ...
, but present in the penises of some primates, such as the
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...
and chimpanzee. The os penis arises from primordial cells within soft tissues of the penis, and its formation is largely under the influence of androgens. The bone is located above the male urethra, and it aids
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
by maintaining sufficient stiffness during sexual penetration. The homologue to the baculum in female mammals is known as the baubellum or '' os clitoridis'' (also ''os clitoris''), a bone in the
clitoris The clitoris ( or ) is a female sex organ present in mammals, ostriches and a limited number of other animals. In humans, the visible portion – the glans – is at the front junction of the labia minora (inner lips), above the o ...
.


Etymology

The word ''baculum'' meant "stick" or "staff" in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
and originated from el, βάκλον, ''baklon'' "stick".


Function

The baculum is used for copulation and varies in size and shape by
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. Its evolution may be influenced by
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
, and its characteristics are sometimes used to differentiate between similar species. A bone in the penis allows a male to mate for a long time with a female, which can be a distinct advantage in some mating strategies. The length of the baculum may be related to the duration of copulation in some species. In carnivorans and primates, the length of the baculum appears to be influenced by postcopulatory sexual selection. In some bat species, the baculum can also protect the
urethra The urethra (from Greek οὐρήθρα – ''ourḗthrā'') is a tube that connects the urinary bladder to the urinary meatus for the removal of urine from the body of both females and males. In human females and other primates, the urethra c ...
from compression.


Presence in mammals

Mammals having a penile bone (in males) and a clitoral bone (in females) include various eutherians: * Order
Primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...
s, although not in
loris Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae (sometimes spelled Lorisinae) in the family Lorisidae. ''Loris'' is one genus in this subfamily and includes the slender lorises, ''Nycticebus'' is the genus con ...
es,
humans Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
, spider monkeys, or woolly monkeys * Order
Rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are roden ...
ia (rodents), though not in the related order
Lagomorpha The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (hares and rabbits) and the Ochotonidae ( pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λαγ ...
(rabbits, hares, etc.) * Order
Eulipotyphla Eulipotyphla (, which means "truly fat and blind") is an order of mammals suggested by molecular methods of phylogenetic reconstruction, which includes the laurasiatherian members of the now-invalid polyphyletic order Lipotyphla, but not the afr ...
(insectivores, including shrews and hedgehogs) * Order
Carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
(including members of many well-known families, such as ursids (bears),
canids Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamily, ...
(dogs),
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammal, marine mammals. They comprise the extant taxon, extant family (biology ...
s (
walrus The walrus (''Odobenus rosmarus'') is a large flippered marine mammal with a discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. The walrus is the only living species in the f ...
es, seals, sea lions), procyonids (raccoons etc.), mustelids (otters, weasels, skunks and others)). The baculum is usually longer in the Canoidea than in the Feloidea, although fossas have long bacula and giant pandas have short bacula. * Order Chiroptera (bats). It is absent in humans,
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraff ...
s (hoofed mammals),
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
s,
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their bra ...
s (
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monot ...
,
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and t ...
),
marsupials Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a ...
, lagomorphs,
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...
s,
binturong The binturong (''Arctictis binturong'') (, ), also known as the bearcat, is a viverrid native to South and Southeast Asia. It is uncommon in much of its range, and has been assessed as Vulnerable on the IUCN Red List because of a declining po ...
s, sirenians, and
cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
s (whales, dolphins, and porpoises), among others. Evidence suggests that the baculum was independently evolved 9 times and lost in 10 separate lineages. The baculum is an exclusive characteristic of placentals and closely related eutherians, being absent in other mammal clades, and it has been speculated to be derived from the epipubic bones more widely spread across mammals, but notoriously absent in placentals. Among the primates, marmosets, weighing around , have a baculum measuring around , while the tiny
galago Galagos , also known as bush babies, or ''nagapies'' (meaning "night monkeys" in Afrikaans), are small nocturnal primates native to continental, sub-Sahara Africa, and make up the family Galagidae (also sometimes called Galagonidae). They are ...
has one around long. The great apes, despite their size, tend to have very small penis bones, and humans are the only ones to have lost them altogether. In some mammalian species, such as badgers and raccoons (''Procyon lotor''), the baculum can be used to determine relative age. If a raccoon's baculum tip is made up of uncalcified cartilage, has a porous base, is less than in mass, and measures less than long, then the baculum belongs to a juvenile.


Absence in humans

Unlike most other primates, humans lack an os penis or os clitoris, but the bone is present, although much reduced, among the
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the e ...
s. In many ape species, it is a relatively insignificant structure. Cases of
human penis The human penis is an external male intromittent organ that additionally serves as the urinary duct. The main parts are the root (radix); the body (corpus); and the epithelium of the penis including the shaft skin and the foreskin (prepuce) ...
ossification following trauma have been reported, and one case was reported of a congenital os penis surgically removed from a 5-year-old boy, who also had other developmental abnormalities, including a cleft scrotum. Clellan S. Ford and Frank A. Beach in '' Patterns of Sexual Behavior'' (1951), p. 30 say, "Both gorillas and chimpanzees possess a penile bone. In the latter species, the os penis is located in the lower part of the organ and measures approximately three-quarters of an inch in length."''Patterns of Sexual Behavior'' Clellan S. Ford and Frank A. Beach, published by Harper & Row, New York in 1951. In humans, the rigidity of the
erection An erection (clinically: penile erection or penile tumescence) is a physiological phenomenon in which the penis becomes firm, engorged, and enlarged. Penile erection is the result of a complex interaction of psychological, neural, vascular, ...
is provided entirely through blood pressure in the corpora cavernosa. An "artificial baculum" or penile implant is sometimes used to treat erectile dysfunction in humans. In ''
The Selfish Gene ''The Selfish Gene'' is a 1976 book on evolution by the ethologist Richard Dawkins, in which the author builds upon the principal theory of George C. Williams's '' Adaptation and Natural Selection'' (1966). Dawkins uses the term "selfish g ...
'',
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An at ...
proposed honest advertising as the evolutionary explanation for the loss of the baculum. The hypothesis states that if erection failure is a sensitive early warning of ill health (physical or mental), females could have gauged the health of a potential mate based on his ability to achieve erection without the support of a baculum. The tactile stimulation hypothesis proposes that the loss of the baculum in humans is linked to the female choice for tactile stimulation: a boneless penis would be more flexible, facilitating a larger range of copulatory positions and whole body movement, giving females greater general physical stimulation. The mating system shift hypothesis proposes that the shift towards monogamy as the dominant reproductive strategy may have reduced the intensity of copulatory and post-copulatory sexual selection, and made the baculum obsolete. Humans "evolved a mating system in which the male tended to accompany a particular female all the time to try to ensure paternity of her children" which allows for frequent matings of short duration. Observation suggests that primates with a baculum only infrequently encounter females, but engage in longer periods of copulation that the baculum makes possible, thereby maximizing their chances of fathering the female's offspring. Human females exhibit
concealed ovulation Concealed ovulation or hidden œstrus in a species is the lack of any perceptible change in an adult female (for instance, a change in appearance or scent) when she is fertile and near ovulation. Some examples of perceptible changes are swelling an ...
, also known as hidden estrus, meaning it is almost impossible to tell when the female is fertile, so frequent matings would be necessary to ensure paternity. Strengths and weaknesses of these hypotheses were revised in a 2021 study, which also proposed an alternative hypothesis: that conspecific aggression, in combination with the development of self-awareness, may have played a role in the loss. If the presence of a baculum exacerbated the prevalence and severity of penile injuries resulting from blunt trauma to a flaccid penis, increasing ability to foresee the consequences of their actions would also enable hominins to realise that these injuries are a useful tool in male-male competition. This behavioural innovation, planned conspecific aggression with the goal of temporary exclusion of competitors from the breeding pool, would create an environment in which a genetic mutation for a penis without a baculum (or with an unossified baculum) would strongly increase the fitness of the mutant phenotype. Along with the hominin propensity for social learning and cultural transmission, this hypothetical scenario may explain why this phenotype became fixed in all human populations.Jakovlić, Ivan (2021) “The Missing Human Baculum: A Victim of Conspecific Aggression and Budding Self-Awareness?” Mammal Review. https://doi.org/10.1111/mam.12237 An alternative view is that its loss in humans is an example of
neoteny Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compar ...
during human evolution; late-stage fetal chimpanzees lack a baculum. (page 134), cited by:

"In humans, neoteny is manifested in the resemblance of many physiological features of a human to a late-stage foetal chimpanzee. These foetal characteristics include hair on the head, a globular skull, ear shape, vertical plane face, absence of penal bone (baculum) in foetal male chimpanzees, the vagina pointing forward in foetal ape, the presence of hymen in neonate ape, and the structure of the foot. 'These and many other features', Bednarik says, 'define the anatomical relationship between ape and man as the latter's neoteny


Cultural significance

The existence of the baculum is unlikely to have escaped the notice of
pastoralist Pastoralist may refer to: * Pastoralism, raising livestock on natural pastures * Pastoral farming, settled farmers who grow crops to feed their livestock * People who keep or raise sheep, sheep farming Sheep farming or sheep husbandry is the r ...
and hunter-gatherer cultures. It has been argued that the "rib" (
Hebrew Hebrew (; ; ) is a Northwest Semitic language of the Afroasiatic language family. Historically, it is one of the spoken languages of the Israelites and their longest-surviving descendants, the Jews and Samaritans. It was largely preserved ...
צֵלׇע ''ṣēlā, also translated "flank" or "side") in the story of
Adam and Eve Adam and Eve, according to the creation myth of the Abrahamic religions, were the first man and woman. They are central to the belief that humanity is in essence a single family, with everyone descended from a single pair of original ancestors ...
is actually a mistranslation of a Biblical Hebrew euphemism for baculum, and that its removal from Adam in the
Book of Genesis The Book of Genesis (from Greek ; Hebrew: בְּרֵאשִׁית ''Bəreʾšīt'', "In hebeginning") is the first book of the Hebrew Bible and the Christian Old Testament. Its Hebrew name is the same as its first word, ( "In the beginning" ...
is a creation story to explain this absence (as well as the presence of the perineal raphe – as a resultant "scar") in humans. In hoodoo, the folk magic of the
American South The Southern United States (sometimes Dixie, also referred to as the Southern States, the American South, the Southland, or simply the South) is a geographic and cultural region of the United States of America. It is between the Atlantic Ocean ...
, the raccoon baculum is sometimes worn as an amulet for love or luck.


Oosik

''Oosik'' is a term used in Native Alaska cultures to describe the bacula of
walrus The walrus (''Odobenus rosmarus'') is a large flippered marine mammal with a discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. The walrus is the only living species in the f ...
es, seals,
sea lion Sea lions are pinnipeds characterized by external ear flaps, long foreflippers, the ability to walk on all fours, short and thick hair, and a big chest and belly. Together with the fur seals, they make up the family Otariidae, eared seals. ...
s and polar bears. Sometimes as long as , fossilized bacula are often polished and used as a handle for knives and other tools. The ''oosik'' is a polished and sometimes carved baculum of these large northern carnivores. ''Oosiks'' are also sold as tourist souvenirs. In 2007, a fossilized penis bone from an extinct species of walrus, believed by the seller to be the largest in existence, was sold for $8,000. The late
United States Congressman The United States House of Representatives, often referred to as the House of Representatives, the U.S. House, or simply the House, is the lower chamber of the United States Congress, with the Senate being the upper chamber. Together the ...
for
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S ...
,
Don Young Donald Edwin Young (June 9, 1933 – March 18, 2022) was an American politician from the state of Alaska. At the time of his death, he was the longest-serving Republican in congressional history, having been the U.S. representative for fo ...
, was known for possessing an 18 inch walrus ''oosik'', and once brandished it like a sword during a congressional hearing.


See also

* Penile spines * Mammal penis *'' Argumentum ad baculum -'' Latin expression describing an argument based on the use of force.


References


Further reading

* *


External links

* * The San Diego Zoo's Conservation and research for endangered species projects
'What is the significance of the ''baculum'' in animals?'

On the evolution of the mammalian baculum: vaginal friction, prolonged intromission or induced ovulation?

The structure of the penis with the associated baculum in the male greater cane rat (''Thryonomys swinderianus'')
*{{cite news, last1=Panciroli, first1=Elsa, title=How do you sex a fossil? {{! Elsa Panciroli, url=https://www.theguardian.com/science/2018/jan/24/cloacae-sex-arms-and-penis-bones-the-tricky-art-of-fossil-sexing, access-date=24 January 2018, work=The Guardian, date=24 January 2018, language=en Mammal anatomy Reproduction in mammals Andrology Penis Mammal penis Articles containing video clips