An integrated assessment of the continued spread and potential impacts of the colonial ascidian, Didemnum sp. A, in U.S. waters

Didemnum sp. A is a colonial ascidian or “sea squirt” of unknown geographic origin. Colonies of Didemnum sp. A were first documented in U.S. waters in 1993 at Damariscotta River, Maine and San Francisco Bay, California. An alarming number of colonies have since been found at several locations in New England and along the West Coast of the contiguous continental United States. Originally believed to be restricted to artificial structures in nearshore habitats, such as ports and marinas, colonies of Didemnum sp. A have also been discovered on a gravel-pavement habitat on Georges Bank at depths of 40-65m. The wide distribution of Didemnum sp. A, the presence of colonies on an important offshore fishing ground, and the negative economic impacts that other species of noninidigenous ascidians have had on aquaculture operations have raised concerns about the potential impacts of Didemnum sp. A. We reviewed the available information on the biology and ecology of Didemnum sp. A and potentially closely related species to examine the environmental and socioeconomic factors that may have influenced the introduction, establishment and spread of Didemnum sp. A in U.S. waters, the potential impacts of this colonial ascidian on other organisms, aquaculture, and marine fisheries, and the possibility that it will spread to other U.S. waters. In addition, we present and discuss potential management objectives for minimizing the impacts and spread of Didemnum sp. A. Concern over the potential for Didemnum sp. A to become invasive stems from ecological traits that it shares with other invasive species, including the ability to overgrow benthic organisms, high reproductive and population growth rates, ability to spread by colony fragmentation, tolerance to a wide range of environmental conditions, apparent scarcity of predators, and the ability to survive in human dominated habitats. At relatively small spatial scales, species of Didemnum and other nonindigenous ascidians have been shown to alter the abundance and composition of benthic assemblages. In addition, the Canadian aquaculture industry has reported that heavy infestations of nonindigenous ascidians result in increased handling and processing costs. Offshore fisheries may also suffer where high densities of Didemnum sp. A may alter the access of commercially important fish species to critical spawning grounds, prey items, and refugia. Because colonial ascidian larvae remain viable for only 12–24hrs, the introduction and spread of Didemnum sp. A across large distances is thought to be predominantly human mediated; hull fouling, aquaculture, and ballast water. Recent studies suggest that colony growth rates decline when temperatures exceed 21 ºC for 7 consecutive days. Similarly, water temperatures above 8 to 10 ºC are necessary for colony growth; however, colonies can survive extended periods of time below this temperature threshold as an unidentified overwintering form. A qualitative analysis of monthly mean nearshore water temperatures suggest that new colonies of Didemnum will continue to be found in the Northeast U.S., California Current, and Gulf of Alaska LMEs. In contrast, water temperatures become less favorable for colony establishment in subarctic, subtropical, and tropical areas to the north and south of Didemnum’s current distribution in cool temperate habitats. We recommend that the Aquatic Nuisance Species Task Force serve as the central management authority to coordinate State and Federal management activities. Five objectives for a Didemnum sp. A management and control program focusing on preventing the spread of Didemnum sp. A to new areas and limiting the impacts of existing populations are discussed. Given the difficulty of eradicating large populations of Didemnum sp. A, developing strategies for limiting the access of Didemnum sp. A to transport vectors and locating newly established colonies are emphasized. (PDF contains 70 pages)

Impact of the invasive colonial tunicate Didemnum vexillum on the recruitment of the bay scallop (Argopecten irradians irradians) and implications for recruitment of the sea scallop (Placopecten magellanicus) on Georges Bank

The invasive colonial tunicate Didemnum vexillum has become widespread in New England waters, colonizing large areas of shell-gravel bottom on Georges Bank including commercial sea scallop (Placopecten magellanicus) grounds. Didemnum vexillum colonies are also fouling coastal shellfish aquaculture gear which increases maintenance costs and may affect shellfish growth rates. We hypothesized that D. vexillum will continue to spread and may affect shellfish larval settlement and survival. We conducted a laboratory experiment to assess interactions between larval bay scallops (Argopectin irradians irradians) and D. vexillum. We found that larval bay scallops avoid settling on D. vexillum colonies, possibly deterred by the low pH of the tunicate’s surface tissue. The results of this study suggest that widespread colonization of substrata by D. vexillum could affect scallop recruitment by reducing the area of quality habitats available for settlement. We propose that the bay scallop can serve as a surrogate for the sea scallop in estimating the negative impact D. vexillum could have on the recruitment of sea scallops on Georges Bank.

Spatial competition induces the mobilization of morula cells in the colonial ascidian Didemnum perlucidum (Tunicata: Didemnidae)

The effects of spatial competition among colonial marine organisms are often evident in the contact zones between colonies. These effects are especially pronounced when the interaction results in overgrowth or necrosis of one of the competitors. Ascidians, one of the dominant taxonomic groups in subtidal sessile communities, have specialized morula cells that provide a defense against microbial infections. Injuries resulting from interspecific competitive interactions might also act as a stimulus for this defensive mechanism. Therefore, we expected to see the recruitment of morula cells in tissues near competitor contact zones. To test the hypothesis that spatial competition elicits this immune response, we placed colonies of the ascidian Didemnum perlucidum from southeastern Brazil in four different types of competitive situations: (1) overgrowth of the competitor, (2) stand-off interactions, (3) overgrowth by the competitor, and (4) free of competitors. Our results indicate that competitive interactions increase the population of morula cells in contact zones, as more cells were observed in interactions that resulted in the overgrowth of individuals of D. perlucidum, and fewer cells were observed in colonies that were free of competitors. We identified the defensive function of the morula cells by showing the presence of the enzyme phenoloxidase within its vacuoles. Phenoloxidase is a widespread enzyme among animals and plants, and is frequently used in defense by synthesizing toxic quinones from polyphenol substrates. This is the first study to document the presence of morula cells in didemnid ascidians and the mobilization of these cells by spatial competition by heterospecifics, and one of the first studies to identify phenoloxidase activity in morula cells.

Effects of competition on sexual and clonal reproduction of a tunicate: the importance of competitor identity

Individual fitness and the structure of marine communities are strongly affected by spatial competition. Among the most common space holders are the colonial ascidians, which have the ability to monopolize large areas of hard substrate, overgrowing most other competitors. The effects of competition on colony growth and on gonad production of the ascidian Didemnum perlucidum were studied in southeastern Brazil by experimentally removing surrounding competitors. Colonies of D, perlucidum competing for space exhibited a growth rate 9 times less than that of colonies that were competitor free. Among the colonies subject to competition, growth rates were unrelated to the percentage of colony border that was free of competitors. However, the identity of the competitor was important in the outcome of border contacts. At the beginning of the experiment, most border encounters of D. perlucidum were with solitary organisms, which in most cases were overgrown. These were progressively replaced by colonial ascidians and bryozoans, resulting mostly in stand-off interactions. Besides reducing asexual growth, spatial competition also affected female gonad production. Colonies free of competitors had a significantly higher proportion of zooids with ovaries. Thus, our findings show that spatial competition reduces both ascidian colony size and gonad production.

Succession of benthic hard-bottom communities in the shallow sublitoral of Comau Fjord, Chile

Succession was already studied over decades. The present thesis investigated the succession on hard substrate at two different study sites within the fjord Comau, Chile. Nine plates were installed at both sites (mouth of fjord and inner fjord) and photographed over three years. Additionally the natural community was recorded and a ground truthing was carried out to verify the analyzed species. Respectively at both sites over 50 different species were identified. Abundance data decreased with only one exception continuously, whereas the percentage cover increased. But the communities on the recruitment plates do still not reach the community structure of the natural environment. The present data showed that the hard-bottom succession in the fjord Comau is best described by the TOLERANCE MODEL (Connell & Slatyer, 1977). An important species of the natural community is the stony coral Desmophyllum dianthus, which normally (outside the fjord) grows beneath 1000 m water depth. The results of this work indicate that the mature community is not reached after 36 months.