Thomas W. Therriault

Molecular resolution of the family Dreissenidae (Mollusca: Bivalvia) with emphasis on Ponto-Caspian species, including first report of Mytilopsis leucophaeata in the Black Sea basin.

Considerable uncertainty exists in determination of the phylogeny among extant members of the Dreissenidae, especially those inhabiting the Ponto-Caspian basin, as multiple systematic revisions based on morphological characteristics have failed to resolve relationships within this group of bivalves. In this study we use DNA sequence analyses of two mitochondrial gene fragments, 16S rRNA and cytochrome c oxidase subunit I (COI), to determine phylogenetic relationships among Dreissena rostriformis, D. bugensis, D. polymorpha, D. stankovici, Congeria kusceri, and Mytilopsis leucophaeata. Dreissena stankovici was determined to represent a sister taxa to D. polymorpha and both are more closely related to other extant Dreissena species than Congeria or Mytilopsis. Sequence divergence between D. rostriformis and D. bugensis was relatively low (0.3-0.4%), suggesting that these two taxa constitute a single species. However, environmental differences suggest two races of D. rostriformis, a brackish water race (rostriformis) and a freshwater race (bugensis). Spread of bugensis-type individuals into habitats in the Caspian Sea that are occupied by rostriformis-type individuals may create novel hybridization opportunities. Species-specific molecular markers also were developed in this study since significant intraspecific variation in morphological features complicates dreissenid identification. Using two gene fragments (nuclear 28S and 16S), we identified restriction fragment length polymorphisms (RFLPs) that distinguish among D. rostriformis/bugensis, D. polymorpha, and D. stankovici and revealed the presence of a cryptic invader to the Black Sea basin, Mytilopsis leucophaeata. This is the first report of this North American native in southern Europe.

History of aquatic invertebrate invasions in the Caspian Sea

Incorporation of the fossil record and molecular markers into studies of biological invasions provides new historical perspectives on the incidence of natural and human-mediated invasions of nonindigenous species (NIS). Palaeontological, phylogeographic, and molecular evidence suggests that the natural, multiple colonizations of the Caspian basin via transient connections with the Black Sea and other basins played an important role in shaping the diversity of Caspian fauna. Geographically isolated, conspecific Ponto-Caspian lineages that currently inhabit fragmented habitats in the Ponto-Caspian region show limited genetic divergence, implying geologically recent episodes of gene flow between populations during the Pliocene to Pleistocene. Several molluscan lineages in the Caspian Sea may have descended from Lake Pannon stock before the Late Miocene isolation of the Caspian depression, about 5.8 million years ago. Anthropogenic activities during the 20th century were responsible for a 1800-fold increase in the rate of establishment of new aquatic species in the Caspian Sea compared to the preceding two million years of natural colonization. The observed success of NIS invasions during the 20th century was due primarily to human-mediated transport mechanisms, which were dominated by shipping activities (44%). Human-mediated species transfer has been strongly asymmetrical, toward the Volga Delta and Caspian Sea from or through Black and Azov Seas. Global and regional trade, particularly that mediated by commercial ships, provides dispersal opportunities for nonindigenous invertebrates, indicating that future invasions in the Caspian Sea are anticipated.

Range expansion of quagga mussels Dreissena rostriformis bugensis in the Volga River and Caspian Sea basin

In 1992, we discovered populations of the nonindigenous quagga mussel Dreissena rostriformis bugensis in the middle reaches of the Volga River. The same species was found in samples collected between 1994 and 1997 in the Volga delta and in shallow areas of the Northern Caspian Sea. D. r. bugensis always co-occurred with its more widespread congener, the zebra mussel D. polymorpha (Pallas 1771). The quagga mussels contribution to total Dreissena abundance increased over time in the middle Volga reservoirs and Volga River delta. D. r. bugensis was common in the Volga portion of Rybinsk Reservoir during 1997 and, by 2000, it was in Uglich, Rybinsk and Gorky Reservoirs on the Upper Volga River. D. r. bugensis was neither found in Ivankov Reservoir, nor in terminal sections of the Volga-Baltic corridor including the eastern Gulf of Finland. Presently, all but the northern-most regions of the Volga River have been colonized by D. r. bugensis. We hypothesize that its introduction into the Volga River and Caspian basin occurred no later than the late 1980s via commercial shipping that utilized the Volga-Don waterway to navigate between the source Black-Azov Sea region and recipient areas on the Volga River. Larval drift likely contributed to establishment of populations at downstream sites, while human-mediated vectors may be responsible for introductions to upstream locations on the Volga River. We anticipate continued northward dispersal in conjunction with shipping activities.

The Caspian-Volga-Baltic Invasion Corridor

The north-south transfer of species in the Volga River basin is not new, but the scale and nature of invasions changed along the Volga-Baltic corridor following transformation of the Volga River from a riverine environment to one of a series of cascading reservoirs. Southward penetration of northern species was facilitated by the formation of a cold-water hypolimnion in the Volga reservoirs. Following reservoir impoundment, 106 invasive species have been found in the Volga River basin, a process that occurred over two different time periods. The first period of invasions occurred between 1940 and 1970, and involved many northern species (77% of total species) moving downstream by passive dispersal. The second period of invasions is still on going and involves invasions by many Ponto-Caspian species (51 % of total species) while new invasions by northern species has decreased substantially (7% of total species). The proportion of exotic species (i.e., invaders originating from basins not adjacent to the Volga basin) increased from 7% during the first period to 41% during the second period. Since the late 1970s, water temperatures in the Volga basin have continued to increase and it is postulated that many invasions during the second period are related to global climatic change.

Range Expansion of the Exotic Zooplankter Cercopagis pengoi (Ostroumov) into Western Lake Erie and Muskegon Lake

Previously reported from Lakes Ontario and Michigan, the nonindigenous zooplankter Cercopagis pengoi was found for the first time in western Lake Erie, the Detroit River, and Muskegon Lake, Michigan, during summer 2001. A native of the Ponto-Caspian region, C. pengoi is currently expanding its range in North America. Analysis of mitochondrial gene ND5 sequences confirmed that the Lake Erie haplotype is identical to that reported previously from Lakes Ontario and Michigan and the Finger Lakes, New York. These findings support the hypothesis that C. pengois range expansion in the Great Lakes likely resulted from inter-lake transfer of ballast water, rather than from additional introductions from European locations. Pleasure-craft traffic operating between Lake Michigan and Muskegon Lake is likely responsible for this inland transfer of Cercopagis, a trend that likely will increase due to human activities.

Evaluation of a new screening-level risk assessment tool applied to non-indigenous marine invertebrates in Canadian coastal waters

Screening-level risk assessment tools for non-indigenous species are useful to inform management and policy decisions. While a number of tools have been developed to evaluate the risk of introductions in terrestrial and freshwater environments, only one tool is available for marine invertebrates [Marine Invertebrate Invasiveness Scoring Kit (MI-ISK)] and it has not been calibrated or evaluated previously. This paper introduces the Canadian Marine Invasive Screening Tool (CMIST), a tool based on a series of questions related to the sequence of events in the invasion process. We assessed the performance of both CMIST and MI-ISK using 60 species/ecosystem combinations of non-indigenous marine invertebrate species known to have been introduced to three Canadian marine ecoregions. Assessment scores were compared to the results of an expert opinion survey about the level of ecological risk posed by these species in these ecoregions. In addition, a simple way to quantify uncertainty and compute confidence limits surrounding expert evaluations and CMIST scores is presented; the technique could be incorporated into other existing tools. The risk scores returned by the two tools generally were similar, had comparable levels of inter-assessor variability, and correlated well with the results of the expert opinion survey for two of the three ecoregions. In the third ecoregion, we believe the lack of highly problematic species and greater uncertainty (linked to less reliable information) contributed to the weaker correlation. For all ecoregions, CMIST scores were more strongly correlated with expert opinion scores when compared to MI-ISK scores. Finally, we demonstrate how CMIST can be used for new incursions; for each ecoregion we scored 15 additional species not yet introduced and compared these scores with those of species already present. Overall CMIST is a promising screening-level tool; it distinguishes between elements of invasion risk (likelihood and impact), has fewer questions to score than similar tools, and could easily be adapted for other taxa.