Analysis of a human-mediated microbioinvasion: the global spread of the benthic foraminifer <i>Trochammina hadai</i> Uchio, 1962

Abstract. A non-indigenous species (NIS) of benthic foraminifera was first identified in a core collected in 1993 in San Francisco Bay, California, USA, and subsequently identified as Trochammina hadai Uchio, 1962. Archived samples and literature reviews were used to determine that the species, which is native to Asia, arrived in San Francisco Bay between the early 1960s and 1983. Through molecular analyses of specimens, archived samples and literature reviews from 1930–1983, and site surveys of harbors and estuaries along the western North American seaboard in 1994–2024, in total more than 2500 samples, we documented the presence of T. hadai at 73 locations in the USA and four in Canada. Trochammina hadai has also been recovered at nine sites in Sweden, two in France, three in Brazil, and two locations at one site in Australia. The rapid temporal and geographic spread of the NIS T. hadai in a non-native location is illustrated by a time series from 1930 to 2024 in San Francisco Bay. Between 1980 and 1986, the species' range expanded from low abundance (1.5 %) at a single site to cover nearly the entire South Bay with &gt; 70 % abundance at some locations. By 1995 and continuing into 2010, the species expanded its range into the central and northern portions of San Francisco Bay, commonly with abundances of &gt; 30 % and sometimes exceeding 70 %. This expansion may predate 1995, but a lack of samples makes it difficult to be more precise. Unfortunately, two Pb-210 and Cs-137-dated cores (BC01 and BC02) recovered from northern South Bay and Central Bay did not clarify this point, but additional cores may. Trochammina hadai is an infaunal opportunist that thrives in polluted locations. We surmise the species was introduced along the west coast of the USA in Puget Sound between 1902 and the 1920s, with cultivated oysters and oyster larvae and associated plant matter and residual sediment. This probably also happened in some areas of France, Sweden, and Brazil, where Japanese oysters were introduced in 1966, 1970, and 1975, respectively. After World War II, commercial shipping expanded dramatically and, with it, the release of ballast water and sediment in receiving ports, which introduced NIS worldwide. This primary vector of introduction occurred in large industrial harbors in several countries, sometimes followed by secondary introductions in small industrial centers and marinas by mud attached to the anchors and anchor chains of smaller boats.

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