In addition to exploring the consequences of intra-specific variation, further work on inter-specific variation in penises would be valuable. For example, morphological variation between Coenobita species includes penises bent at various angles [ 9 , 24 ], which in theory could adaptively permit males of some species to simultaneous mate and fight e.
Worldwide, approximately 20 Coenobita species exist [ 22 ], and while all are known to use remodelled shells [ 13 ], no study has quantified the relative level of shell remodelling across these species or mapped any more than the three species examined here onto a molecular phylogeny.
A robust phylogeny of Coenobita , alongside precise measures of shell remodelling across all these species, could enable further tests of the evolution of penis size and its relation to private property, as well as to social evolution more broadly [ 18 , 20 , 30 ]. Intriguingly, distant relatives in Paguridae separate from Coenobitidae and Diogenidae also independently evolved sexual tubes [ 9 , 31 ]; though the functions of these tubes, many of which are bendable rather than permanently hardened, remain mysterious due to a lack of observations on how they are used in nature.
This hypothesis may also inform general tests of private property that shed light on an opposite trend: the evolution of small relative penis size. Phenotypic investments in sexual structures e. Such investments, by trading off against penis size, may create alternative axes of property defence: weapon size versus penis size [ 32 , 33 ]. In the present study, the giant coconut crab Birgus latro had a meagre relative penis size, among the very smallest of all species measured; this species instead puts extreme investment into powerful claws [ 23 ], which enable predation and defence of an extensive non-portable territory and burrow system [ 20 , 35 ].
Broader comparative analyses across taxa could thus explore such trade-offs by examining relative weapon and penis size in relation to private property value. All measurements were made of preserved museum specimens no live animals were used. The entire dataset is available online in the electronic supplementary material. I declare that I have no competing interests.
For guidance on the phylogenetically controlled analyses, I am indebted to Susanne Shultz. For kindly providing housing near the Smithsonian during my study, I thank Jane Stanley. And for valuable feedback, I thank two anonymous reviewers as well as the audience of a fall seminar in the Department of Invertebrate Zoology at the Smithsonian Institution and the audience of a spring conference talk at the International Crustacean Congress in Washington DC.
Females release eggs from these gonopores, which are flat holes and which lack any extension or structure in all species measured. Login to your account.
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Enter your email address below and we will send you your username. Open Access. View PDF. Laidre Mark E. Private parts for private property: evolution of penis size with more valuable, easily stolen shells 6 R. Section Supplemental Material. Open Access Research articles.
Mark E. Abstract Evolution has generated enormous diversity in animal genitalia. Download figure Open in new tab Download powerPoint. References 1.
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The Law of the Body | Emory University School of Law | Atlanta, GA
Crustacean Biol. Oikos , Evidence from natural and field induced shell exchanges. Article 8. A transfer of special co-ownership by agreement shall not be effective unless it has been registered in accordance with the legal provisions and regulations regarding registration. Article 9. When a Cambodian co-owner of a private unit has transferred ownership of a private unit to a foreigner, the special co-owner shall acquire an ownership right to the private unit and only the right to use and enjoy the common areas.
In case the co-owner of the private unit is a foreigner and has transferred the private unit to a Cambodian, the Cambodian co-owner shall acquire an ownership right in the private unit and undivided ownership of the common areas. Article Successors of special co-owners shall receive the rights and obligations of the deceased in accordance with the laws and regulations in effect.
In case the inheritance is unoccupied because there is no successor or the successor abandons it, or if the inheritance is not designated to any person in accordance with the legal provisions, such inheritance shall become the property of the State.
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Article In the event that, after this law becomes effective, any foreigner receives the transfer of or purchases any private unit of a co-owned building in violation of Article 6 of this law, the transaction shall be null and void and the parties to the transaction shall make a restitution. Part 1. Rights and obligations of special co-owners concerning private units. Article 12 Special co-owners of private units shall have the right to freely use, enjoy, and dispose of the ownership of their private units to the extent determined by law. Article Special co-owners of private units cannot use their private units to hurt, infringe, or hinder the use, or to disturb the livelihood, of the other co-owners of private units.
Special co-owners of private units shall not take any action that could harm the maintenance of the co-owned building or take any other action against the common interests of the co-owners of private units in relation to the management or use of the co-owned building. Special co-owners of private units of co-owned buildings shall have the same obligations and burdens as Cambodian co-owners.
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Part 2. Rights and obligations of special co-owners concerning common areas. Common areas shall be under the regime of special undivided ownership of all co-owners of private units. Special co-owners shall have only the right to use and enjoy the common areas. Special undivided ownership and other marginal rights recognized by law shall not be subject to division, disposal, and forced sale separate from the private unit.
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