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A pheromone bouquet controls the reproductive behaviour of the male shore crab, Carcinus maenas

Fletcher, Nichola; Terschak, John A.; Bartels-Hardege, Helga D.; Bublitz, Ralf; Schirrmacher, Paula; Hardege, Jorg D.


Nichola Fletcher

John A. Terschak

Helga D. Bartels-Hardege

Ralf Bublitz

Paula Schirrmacher

Jorg D. Hardege


The reproduction of many brachyuran crustaceans involves the formation of mating pairs often around the time of the female moult with attraction of a sexual partner and mating behaviour controlled by sex pheromones. In shore crabs, Carcinus maenas, females produce sex pheromones that are released in the urine. High Performance Liquid Chromatography analysis (HPLC) of female urine shows that the pheromone, identified as the nucleotide uridine diphosphate (UDP), elutes as an unresolved peak with structurally related nucleotides. We examined female urine samples over the moult cycle and detected UDP as well as uridine triphosphate (UTP). Bioassays were conducted to establish the possibility of a blend of nucleotides forming a sex pheromone bouquet in C. maenas. Whilst UDP induced the male mate guarding behaviour (cradling), a mixture of the two nucleotides at a ratio of 4:1 UDP:UTP elicited an even stronger mating response than either UDP or UTP individually. The urine concentration and composition of these nucleotides changes over the moult period pre and post ecdysis, providing evidence that a pheromone bouquet composition is not always constant. The change of the bouquet is related to the physiological state of the sender, here the moult cycle. Our study unravels the functionality of reaction-specific molecules in a pheromone bouquet. Whilst UDP is the mating signal, UTP acts as an attractant and combined they maximise the reproductive response. The use of bouquets provides species-specificity, potentially enabling reproductive isolation of sympatric species, and contains valuable information on the physiological state of the sender.


Fletcher, N., Terschak, J. A., Bartels-Hardege, H. D., Bublitz, R., Schirrmacher, P., & Hardege, J. D. (2022). A pheromone bouquet controls the reproductive behaviour of the male shore crab, Carcinus maenas. Aquatic ecology, 56, 419–427.

Journal Article Type Article
Acceptance Date Dec 4, 2021
Online Publication Date Dec 24, 2021
Publication Date 2022-06
Deposit Date Aug 10, 2022
Publicly Available Date Aug 10, 2022
Journal Aquatic Ecology
Print ISSN 1386-2588
Electronic ISSN 1573-5125
Publisher Springer Verlag
Peer Reviewed Peer Reviewed
Volume 56
Pages 419–427
Public URL


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