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Most barnacles are sessile, simultaneous hermaphrodites that reproduce by copulation. This is achieved through the extension of a muscular penis, famous for being the proportionally largest in the animal kingdom. The penis is a long cylindrical or conical organ, composed of a series of folded rings, allowing it to stretch to great lengths. The penises are covered with chemosensory setae allowing them to seek out receptive neighbors. For many species, the condition of the penis changes seasonally. In the most extreme circumstances, it degenerates and is shed during the first post-mating molt and is re-grown for the next mating season. Barnacle penises have been shown to exhibit phenotypic plasticity in response to many different challenges. When exposed to heavy waves, diameter is increased by thickening both the cuticle and muscles. When mates are far, length increases by adding ringed annulations. Experiments have shown that these plastic traits are modular, capable of changing independently from each other and that they improve mating ability. Alternate strategies to increase reproductive ability by barnacles include the production of dwarf and complemental males, sperm casting and sperm leakage, and aerial copulation. All of these mating strategies may have important implications for the study of reproductive biology, life history, and sex allocation theory., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases toward an asymptote of 50% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed., (© 2012 The Author(s). Evolution© 2012 The Society for the Study of Evolution.)

Acorn barnacles are important model organisms for the study of sex allocation. They are sessile, nonselfing hermaphrodites that copulate with penises that have been suggested to be phenotypically plastic. On wave-exposed shores, Semibalanus balanoides develop penises with relatively greater diameter whereas in wave-protected sites they are thinner. A reciprocal transplant experiment between wave-exposed and protected sites tested whether these exposure-specific morphologies have adaptive value. Mating success was compared over a range of distances to compare the ability of barnacles to reach mates. Barnacles that grew in the wave-protected site and mated in the wave-protected site fertilized more broods at increasing distances than those transplanted to the wave-exposed site. For barnacles that developed in the wave-exposed site, there was no difference in the ability to fertilize neighbors between sites of differing exposure. This study demonstrates the adaptive value of plasticity in penis morphology. The results suggest a trade-off between development of a penis adapted to wave exposure and the ability to fertilize distant mates. Barnacles in different physical environments are limited by different factors, which may limit numbers of potential mates, constrain optimal sex allocation strategies and alter reproductive behavior.