Zoologischer Anzeiger 249 (2010) 21–32 Competing for last place: Mating behaviour in a pill-box crab, Halicarcinus cookii (Brachyura: Hymenosomatidae) Anneke M. van den Brink, Colin L. McLay à School of Biological Sciences, Canterbury University, PB 4800, Christchurch, New Zealand Received 7 August 2009; received in revised form 14 January 2010; accepted 14 January 2010 Corresponding Editor: S. De Grave Abstract The mating strategy of Halicarcinus cookii was investigated to ascertain how males maximised their fitness through mate choice. An intertidal population at Kaikoura, New Zealand, was dominated by mature crabs of both sexes in summer and by immature crabs in the colder months. More than 95% of mature females were ovigerous with early stage and late stage broods found in almost every month, indicating that egg production and larval release is continuous. The operational sex ratio was less than 1 male/female in summer, but often more than 1.0 in the colder months. The gonosomatic index increased along with brood development so that as soon as zoeae were released, the next clutch of eggs was ready to be fertilised. Males searched for receptive females and began pre-copulatory mate guarding without any courtship display. They mated preferentially with late stage or non-ovigerous females: copulation duration was longest for stage 5 females as was post-copulatory guarding (mean 18.3 h). Late stage females were up to 14% of the female population. Mate attraction seems to be the result of an ovarian signal rather than from the developing brood. Manipulation of the sex ratio had effects upon copulation duration and post-copulatory guarding: presence of a rival male increased duration of guarding. Females showed precocious mating in the penultimate instar and were able to lay fertilised eggs after their pubertal moult in the absence of males. H. cookii females have many mates, but males attempt to ensure paternity by preferentially pursuing mature females close to egg laying and by guarding these females after copulation. These behaviours are all elements of a competitive strategy to ensure that a male loses (not wins) the race to copulate because females have a ventral seminal receptacle, giving sperm precedence to the last male to mate. Male mating behaviour is a consequence and evolutionary response to female morphology. & 2010 Elsevier GmbH. All rights reserved. Keywords: Copulation; Egg development; Female choice; Gonosomatic index; Mate attraction; Mate guarding; Mating behaviour; Operational sex ratio; Precocious maturation; Seminal receptacle structure; Sperm competition 1. Introduction Brachyuran reproductive strategies should maximise individual reproductive fitness (Diesel, 1991). For males the aim is to fertilise as many eggs as possible and for females it is to maximise the supply and genetic diversity ARTICLE IN PRESS www.elsevier.de/jcz 0044-5231/$ - see front matter & 2010 Elsevier GmbH. All rights reserved. doi:10.1016/j.jcz.2010.01.002 à Corresponding author. E-mail addresses: Anneke.brink@gmail.com (A. M. van den Brink), colin.mclay@canterbury.ac.nz (C. L. McLay).