Seasonal changes of the introduced Caulerpa racemosa var. cylindracea (Caulerpales, Chlorophyta) at the northwest limit of its Mediterranean range

A study of the meadows of the invasive Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman et Boudouresque was carried out over one year at Marseilles (Provence, France) where the alga is thriving, probably since 1994, in the cold waters of the north western Mediterranean Sea. At an early phase of colonisation, the C. racemosa meadow is characterized by a patchy distribution pattern. Several years are necessary to obtain a dense and continuous meadow. In one area colonized for more than 4 years, C. racemosa has developed a continuous meadow with wide seasonal variations. Maximum development was reached in autumn (biomass: 82 ± 3 g DW m−2; length of stolons: 1162 ± 86 m m−2; number of apices: 8360 ± 405 m−2; number of erect axes: 20955 ± 1499 m−2) and the minimum from winter to early spring (respectively, 0.3 ± 0.1 g DW m−2; 3 ± 1 m m−2; 220 ± 55 apices m−2; 35 ± 15 erect axes m−2). Seasonal variations in the growth rate were highly significant. The season of high growth lasted from June to October. The apical growth rate of a stolon reached a maximum of 7.5 ± 0.3 mm day−1 in early October, then began to decrease significantly from the end of October to December, before becoming nearly nil from January to early May. Annual net production rate expressed in terms of stolon length and biomass was estimated as 5801 m m−2 a−1 and 612 g DW m−2 a−1, respectively. During the growth period, the turnover rate of the C. racemosa stolons was estimated at from 25 to 46 days. The growth rate was closely correlated to the seawater temperature (R2 = 0.83), whereas no significant correlation was found between growth and irradiance. During the growth period, a decrease in temperature rapidly affects the growth rate, which soon recovers its earlier level when the temperature rises again. In winter, the growth rate decreased rapidly with the seasonal drop in the seawater temperature. Grazing by fish (Sarpa salpa and Boops boops) can also affect the growth rate from September to December by consumption of the erect axes and stolon apices, enhancing the ramification of stolons. Seasonal changes at Marseilles are much sharper than those reported for warmer Mediterranean localities (French Riviera, Italy, Croatia): in winter and early spring C. racemosa meadows decreased and locally disappeared, leaving a barren substrate. C. racemosa survives the lower winter seawater temperatures of the north-western Mediterranean Sea probably in the form of zygotes and/or small fragments (rhizoids, stolons, propagules).