Life cycle and seasonal dynamics of Cucullanus cirratus O.F. Müller, 1777 (Nematoda, Ascaridida, Seuratoidea, Cucullanidae) in Atlantic cod, Gadus morhua L.

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Abstract
Mature specimens of Cucullanus cirratus O.F. Müller, 1777 (Cucullanidae) were obtained from the pyloric caeca and intestine of Atlantic cod, Gadus morhua L., from Danish waters. Eggs embryonate in seawater. Third-stage larvae about 400 µm long, with amphids and dereids, hatch from the egg. Experimental studies indicated that third-stage larvae were infective to calanoid and cyclopoid copepods and sand gobies, Pomatoschistus minutus (Pisces, Gobiidae). Larvae entered the haemocoel of copepods but did not grow. In gobies, the third-stage larvae entered the intestinal mucosa and grew to 800 µm in length within 6 months. They were not encapsulated. Experimental infections of cod (8-30 cm long) showed that free-living third-stage larvae are not infective, whereas >700 µm long third-stage larvae from gobies survived in the cod. Third-stage larvae 700-1200 µm long occur in the stomach mucosa, where they develop and moult to fourth-stage larvae. The fourth-stage larvae then migrate to the pyloric caeca and anterior part of the intestine, where they moult and develop to the mature adult stage. No developmental stage became encapsulated. Naturally infected cod (>20 cm total length) harboured moulting third-stage larvae and <2 mm long fourth-stage larvae 2 months post capture. Naturally infected 4- to 5-month-old codlings (8-10 cm total length) harboured 2-3 mm long fourth-stage larvae only, indicating that they acquired the third-stage larvae as planktivorous fry only a few centimetres long. Cucullanus cirratus may have a life-cycle that involves copepod transport hosts and fish (gobies or cod fry) intermediate hosts. Postcyclic development occurs in gadoids when an infected cod is consumed by another cod (cannibalism). Examinations of 350 (8-78 cm total length) naturally infected cod showed that group 1 and older cod are infected throughout the year, with maximum prevalence of third-stage larvae in spring and summer. The greatest prevalence of gravid worms was observed in autumn.