Reproduction in crabs: strategies, invasiveness and environmental influences thereon

This thesis provides insights into the interconnectedness of crab reproductive biology, the selective forces leading to their development, the possible links to invasiveness and the influences of environmental factors thereon. The empirical data collected and presented in this thesis can be used to compare different crab species and make predictions about the effect of climate change on their population dynamics and invasiveness. Two crab species in particular are examined in this thesis, Halicarcinus cookii and Hemigrapsus takanoi , which share similar size and habitat preferences, but have different reproductive strategies and statuses as endemic and invasive. The potential effects of increased water temperature due to climate change on the reproduction and eventual population changes in the species are investigated as well as the possible links reproduction and temperature have with the invasiveness of a species. Halicarcinus cookii is an endemic crab to the coasts of New Zealand and is not known anywhere else. The species shows determinate growth, hard shell mating, continuous brood production and ventral seminal receptacles. After the final moult the female produces numerous offspring limited only by sperm availability. With a hard shell the female also avoids mortality resulting from the regular vulnerable soft shell stage. Individuals mature over a range of sizes but do not continue growing after their pubertal moult. With such a terminal moult, brood size is limited by female size. Hemigrapsus takanoi is native to the north west Pacific, but has been introduced and is very successful in Europe. The species shows indeterminate growth, hard-shell mating, a defined breeding season and ventral seminal receptacles. With indeterminate growth they continue moulting and growing throughout their adult life. After their pubertal moult, these species can mate throughout the year and produce 2-3 broods between each moult. They are not limited in growth or regeneration of limbs and can safely hide from predators during the vulnerable soft-shell inter-moult period rather than mating which exposes them to predators. Despite their different reproductive strategies, broods of both species showed a similar reaction to increased water temperature in that the duration of development of the brood decreased as temperature increased. Extrapolating the results to a climate change scenario, it is suggested that with a temperature rise of 2°C H. cookii could produce one extra brood of over 1000 offspring per female life time, potentially leading to a 10-15% increase in fecundity and possible population growth. As H. takanoi does not show continuous brood production, predicting the effect of temperature rise is more difficult, but evidence suggests that fecundity is also likely to increase in this species with an increase in water temperature. Temperature increase may also lead to a change in invasiveness of a species. If areas currently below the optimum temperature for a species become warmer, it is possible that a species may spread to the new locations. Hemigrapsus takanoi may spread further north in Europe than it’s current distribution (assuming it is limited by temperature). Furthermore, if temperatures increase the rate of reproduction in a non-indigenous species, they may become more invasive in their present location. The colonisation of a new habitat will involve new interactions, such as predation and competition, with species not previously encountered. The interactions of the two invasive crab species H. takanoi and Hemigrapsus sanguineus with the native Carcinus maenas in the delta waters of SW Netherlands was also investigated in this thesis. Whereas C. maenas was the most common shore crab in these waters, its numbers have declined on the soft sediment substrates during the last 20 years. As the two invasive crab species were first recorded in the Dutch delta in 1999, they could not have initiated the decline of the native C. maenas. However, within a few years H. takanoi completely dominated the intertidal hard substrate environments; the same environments on which juvenile C. maenas depend. On soft sediment substrate the native and invasive crab species are presently more or less equally abundant. Nowadays H. takanoi appears to be a fierce interference competitor or predator for small C. maenas specimens by expelling them from their shelters. However, due to the habitat generalist nature of C. maenas, it is unlikely that the Hemigrapsus species will cause it’s local extinction. More likely is that they will learn to live together. The objective to provide new information about a rarely studied species (Halicarcinus cookii) was fulfilled in this thesis and the information can be used as bases for comparison for future research. The hypothesis that temperature has no effect on the reproductive rate of crabs was rejected as both study species showed similar increases in brood development rate with increased temperatures. This suggests that global temperature rises may increase the reproductive rate of wider crab populations. The hypothesis that the arrival, presence and effect of Hemigrapsus takanoi in the Dutch delta waters has had no effect on the native green crab Carcinus maenas was complicated by the fluctuations and the decrease in C. maenas numbers prior to the arrival of H. takanoi. It was concluded that while H. takanoi did not cause the initial decrease in the C. maenas population, it did take advantage of it and now dominates niches previously occupied by juvenile C. maenas where size dependent competition and/or predation on juvenile C. maenas occurs.