Observations of recruitment and colonization by tunicates and associated invertebrates using giant one-meter² recruitment plates at Woods Hole, Massachusetts

Large recruitment plates measuring 1 × 1 m were deployed over an 18-month period from September 2013 to March 2015 for the purpose of documenting recruitment and colonization processes of marine invertebrate species at Woods Hole, Massachusetts. Each side of two plates was subdivided into 16 subareas (25 × 25 cm), and an observational strategy was developed whereby, at approximately two-week intervals, a different subarea was cleaned. Using this approach, we were able to photographically document species recruitment and growth interactions. Water temperature records from the site show that steady warming and cooling between 3 and 20° C changed at a mean rate of 0.2 ° C d^-1. However, temperature changes during the coolest and warmest parts of the temperature cycle were highly variable. In 2014, between the first and last occurrence of 0° C, temperatures were ≤0° C 15 percent of the time, but in 2015 temperatures were ≤0° C 93 percent of the time. In 2014, between the first and last occurrence of 21° C, temperatures were ≥21° C 88 percent of the time, and this warm period correlated with the disappearance of the hydroid Ectopleura crocea, the solitary tunicates Ascidiella aspersa and Ciona intestinalis, and the 2013 generation of Botrylloides violaceus. In Woods Hole, large plates provided enough space to accommodate both fast- and slow-colonizing species, resulting in the establishment of a diverse assemblage that was observed over a long time period. The most successful colonizing species had relatively long reproductive and recruitment periods, grew rapidly, repelled settlement onto their surfaces by larvae of any species, defended themselves against overgrowth by any species, overwintered, and lived a long time. Of the three dominant species observed in this study, the colonial tunicates Didemnum vexillum and Botrylloides violaceus had these qualities; the encrusting colonial bryozoan Schizoporella unicornis had all but one, it grew more slowly than the others. Barnacles constituted the only biological substrate that was effectively colonized by other species, both by larval recruitment and overgrowth. In Woods Hole, after a substrate had become fully colonized, there was very little opportunity for new recruitment or colony growth until new substrate opened after the death of colonies and individuals and the disappearance of biogenic structures such as amphipod tubes. An understanding of colonization processes utilized by invasive species allows prediction of their potential effects on ecosystems in areas where they are not yet present.