The document discusses the baculum, a bone found in the penis of many mammal species. It serves several potential functions, including providing mechanical support during copulation, stimulating the female, and influencing penile morphology. The baculum displays remarkable diversity in shape and size between species. This diversity is believed to have evolved through sexual selection, with females potentially biasing fertilization towards males with bacula that stimulate them most.
The document discusses the baculum, a bone found in the penis of many mammal species. It serves several potential functions, including providing mechanical support during copulation, stimulating the female, and influencing penile morphology. The baculum displays remarkable diversity in shape and size between species. This diversity is believed to have evolved through sexual selection, with females potentially biasing fertilization towards males with bacula that stimulate them most.
The document discusses the baculum, a bone found in the penis of many mammal species. It serves several potential functions, including providing mechanical support during copulation, stimulating the female, and influencing penile morphology. The baculum displays remarkable diversity in shape and size between species. This diversity is believed to have evolved through sexual selection, with females potentially biasing fertilization towards males with bacula that stimulate them most.
copulation. Consistent with a role mechanical support or influence on penile morphology). Interestingly, in mechanical support, certain complementary female structures Paula Stockley primates and carnivore species have been discovered in ground that have prolonged copulation squirrels (Spermophilus spp.), where What is a baculum? The baculum also have a longer baculum. A role the protruding teeth of the baculum (os penis) is a bone found within the in female stimulation also seems apparently interdigitate with folds in penis of certain mammals, including likely, particularly in species where the vagina during copulation. many primates, rodents, bats, copulation is prolonged with multiple carnivores, and some insectivores. intromissions, thrusting or multiple Why are bacula so diverse? The It is an isolated bone, derived from ejaculations. Copulatory stimulation baculum has long been a bone of connective tissue and located at the can have important consequences contention, not least because such distal end of the penis, above the for reproduction as it may facilitate diversity of form has proved difficult to urethra. Lower mineral density and sperm transport and affect female explain. One now largely discounted reduced stiffness of the baculum sexual receptivity, as well as stimulate hypothesis was that the diversity of compared to skeletal bones may help ovulation or initiate pregnancy. bacula is a non-adaptive by-product reduce the risk of fracture under strain The baculum may help stimulate of selection on other traits. Nowadays during copulation, although breakage such responses, either directly it is believed that the baculum and can sometimes occur. (where it extrudes from the tip of penis are subject to direct selection the glans penis) or indirectly (via as a result of their role in copulation. You mentioned primates, do we humans have such a bone? No, human males don’t have a baculum! This is surprising because all other apes and Old World monkeys have one, despite a trend towards reduced size of the baculum among the great apes. As fossil primate bacula are extremely rare, it is unknown when the baculum was lost within the hominid lineage. And despite some speculation relating to upright posture and changing mating strategies, why human males lack a baculum remains enigmatic.
What does it look like? A particularly
striking characteristic of the baculum is its extreme anatomical diversity. Bacula of different species come in a multitude of forms, with variation in their length, thickness, curvature and complexity of shape (Figure 1). The baculum can also be small or large relative to body size, reaching more than 60 cm in the walrus, Odobenus rosmarus. More complex forms may feature bizarre looking teeth or digit-like projections, including components that protrude from the glans penis in certain rodents. Such diversity of form makes the baculum a particularly useful feature for species identification and taxonomy.
What is it for? Although the
baculum was first identified in the seventeenth century, its exact functional significance is still Figure 1. Diversity of bacula. unclear. Multiple functions seem Examples of bacula of North American mammals. Top left: bacula of ground squirrels (Sper- mophilus sp.) with spoon-shaped distal ends and tooth-like projections. Top right: trident plausible: for example, a baculum shaped bacula of rice rats (Oryzomys sp.) and voles (Microtus sp.); illustrations demonstrate may provide mechanical support complexity of bacula shape from different perspectives. Bottom: relatively simple bacula of for the penis, assist in overcoming some large carnivores (two bear species (Ursus) illustrated above a sea lion (Zalophus)). Illus- vaginal resistance or protect the trated by William L. Brudon, reproduced with permission from Burt (1960). Magazine R1033
Historically, much interest focused primates. Also, sexually selected
on the potential role of mammalian traits may show high phenotypic Primer genitalia in promoting reproductive variance relative to non-sexual traits isolation of species. The ‘lock and key and scale positively in relation to hypothesis’, for instance, proposed body size. Indeed, in the muskrat Behavioral that genitalia are radically different between species in order to prevent (Ondatra zibethicus) large males have relatively larger bacula than architecture of the interspecific mating and hybridization. small males and baculum traits cortical sheet A variant of this hypothesis is that (especially width) are particularly bacula of different shapes elicit variable between males. However, different responses from females high levels of variation and positive Rodney J. Douglas during copulation, and that only allometry are not necessarily a and Kevan A.C. Martin species-specific stimulation will lead consequence of sexual selection. to successful reproduction. Baculum morphology has also The effortless ability of vertebrates to been linked to male social status in explore and exploit their environment How did the diverse bacula evolve the bank vole (Myodes glareolus). is strongly correlated with the then? Most recent attention has Dominant males have wider bacula evolution of the most anterior part of focused on the idea that rapid and than subordinates relative to their their nervous system, the forebrain, divergent evolution of male genital body size, which might at least where data from autonomic (visceral), morphology, including the baculum, partly explain the superior success limbic (emotive), and internal and is driven by sexual selection. This of dominant males in sperm peripheral sensors of the external applies particularly to species where competition. world are combined to develop, females mate with multiple males, as decide, and deploy advantageous is very common among mammals. Do females have something similar behaviors. The correlation of There are ways in which the baculum to the baculum? Females do indeed behavioral performance with forebrain could influence sexual selection: have a homologous bone, the expansion suggests that evolution for example, females might bias baubellum or os clitoridis, which is has discovered the developmental fertilisation in favour of males whose present in the clitoris of most if not all means of building vertebrate brains baculum (and penis) stimulates them species with a baculum. It is usually to produce a scalable, special- most during copulation, a process a small bone with the appearance of purpose architecture for efficiently known as ‘cryptic female choice’. an underdeveloped baculum, but it processing and expressing behavior. Also, females may reliably assess can sometimes be relatively large, as In mammals, the exuberant expansion male size or quality based on the in the ring-tailed lemur (Lemur catta) of this forebrain is dominated by the baculum. Being able to assess where the baubellum is nearly half the growth of their cortex — the two- male quality during copulation length of the baculum. Unfortunately, dimensional sheet that is the major could be of particular benefit when even less is known about the source of their intelligent behavior, opportunities for quality assessments baubellum than about the baculum... especially for primates. before mating are limited, such The complexity of the brain is so as when copulation takes place Where can I find out more? overwhelming that at every level underwater or underground. Burt, W.H. (1960). Bacula of North American investigators have been forced to mammals. Misc. Publ. Mus. Zool., Univ. Baculum evolution could also be Michigan 113, 1–75. focus only on particular aspects of influenced by sperm competition. Dixson, A.F. (2012). Primate Sexuality: Comparative its evolution and development, or Studies of the Prosimians, Monkeys, Apes and For example, the baculum might help Human Beings, Second Edition (Oxford: Oxford its structure and function, or the deliver sperm optimally or displace University Press). behavior it generates. However, the ejaculates of rival males. By Eberhard, W.G. (1985). Sexual Selection and over decades all these individual Animal Genitalia, (London: Harvard University supporting prolonged intromission Press). brushstrokes have accumulated after ejaculation, a baculum might Hosken, D.J., and Stockley, P. (2004). Sexual to produce a surprisingly coherent selection and genital evolution. Trends Ecol. also help males to reduce the risk Evol. 19, 87–93. and conceptually simple picture of that females will mate again with Lemaitre, J.-F., Ramm, S.A., Jennings, N., and the interrelationships of evolution, others. Stockley, P. (2012). Genital morphology linked development, and brain organization to social status in the bank vole (Myodes glareolus). Behav. Ecol. Sociobiol. 66, 97–105. to produce what we call here, the Is there evidence for these ideas? Leonard, J. and Cordoba-Aguilar, A. (2010). The ‘behavioral architecture’ of the Evolution of Primary Sexual Characters in There are some indications. For Animals, (Oxford: Oxford University Press). cortical sheet. In this primer we instance, if a larger baculum is Ramm, S.A. (2007). Sexual selection and genital locate our current state of knowledge advantageous in sperm competition, evolution in mammals: A phylogenetic analysis in a single conceptual framework of baculum length. Am. Nat. 169, 360-369. baculum size should be greater Tasikas, D.E., Fairn, E.R., Laurence, S., and that unifies these seemingly among species with more intense Schulte-Hostedde, A.I. (2009). Baculum disparate fields of investigation. variation and allometry in the muskrat (Ondatra sperm competition. The correlation zibethicus): a case for sexual selection. Evol. We show that while the relations varies between groups, as might be Ecol. 23, 223–232. between different parts of the expected if the baculum functions sheet may be complex, they are Mammalian Behaviour and Evolution Group, differently across taxa: more intense not arbitrary, not least because Department of Evolution, Ecology and competition appears to favour a Behaviour, University of Liverpool, Leahurst the actual physical organisation of longer baculum among rodents and Campus, Neston, CH64 7TE, UK. the cortical sheet itself defines a carnivores, but not among bats and E-mail: P.Stockley@liverpool.ac.uk coherent logic by which effective