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Alterations of Lake Michigan Benthic Communities by the Invasive
Colonial Hydroid, Cordylophora caspia: Effects on Fish Prey

Project Account Number: R/HE-41-06
PRINCIPAL INVESTIGATOR: Nadine C. Folino-Rorem
INITIATION DATE: March 1, 2006
COMPLETION DATE: February 29, 2008
AFFILIATION: Wheaton College

Nadine C. Folino-Rorem
Wheaton College
Wheaton, IL 60187
Phone: 630-752-7038
nadine.c.folino-rorem@wheaton.edu

Objectives
Our primary objective is to determine the impact of the invasive, Ponto-Caspian colonial hydroid, Cordylophora caspia on benthic invertebrate communities in southern Lake Michigan. We propose to address two fundamental questions related to the presence of Cordylophora in southern Lake Michigan: 1) What is the predatory role of this hydroid in Lake Michigan macrobenthic communities?, and 2) Does the filamentous structure of this hydroid enhance macroinvertebrate abundances in zebra mussel colonies?

More specifically, we will determine how this benthic invertebrate influences prey availability for fish. We propose a project examining three objectives involving field and laboratory research to assess the effects of Cordylophora on benthic invertebrates in southwestern Lake Michigan: 1) document specific locations and diets of Cordylophora (specifically at sites where it is associated with zebra mussel colonies); 2) characterize feeding preferences of Cordylophora on common Lake Michigan benthic invertebrates using laboratory feeding experiments; 3) compare species composition and abundances of invertebrates present in mussel colonies with and without Cordylophora to assess the impact of this hydroid as a predator that decreases fish food availability and/or as a filamentous substratum that enhances invertebrate densities in freshwater benthic invertebrate communities.

Methodology
Objective 1: Assessing Cordylophora locations and diets
Zebra mussel colonies will be assessed at numerous locations in southwestern Lake Michigan for the selection of sites where Cordylophora co-occurs with zebra mussels. At two to three of these sites, rocks with coexisting zebra mussels and hydroids will be collected from two depths at each site because non-mussel invertebrate community structure has been found to differ depending on water depth. Samples of comparable rocks/mussel colonies with Cordylophora will be collected at < 10m and > 15 m. During Year 1 of funding, we will use SCUBA to conduct bi-monthly field sampling in June, July and August of 2006. Ten rocks with zebra mussels and associated Cordylophora will be sampled from each depth for a total of 20 rocks at each site; rocks with mussels will be placed into individual Ziploc® bags. The samples will be preserved by adding 95% ethanol to the bags upon returning to the surface; samples will be returned to the laboratory for sorting and identification of invertebrates. From each sample, 200 Cordylophora hydranths (minimum) will be dissected to assess prey items consumed by the hydroid. Comparisons of hydroid hydranth contents and benthic invertebrates from mussel colonies will allow us to determine to what extent hydroid diets reflect fish prey availability at each site.

Objective 2: Hydroid Feeding Preferences in the Lab
Assessing the preference for different benthic prey will allow us to address the hydroid's preference for and ability to consume prey that also serve as fish food. In order to assess feeding preferences, live hydroids from Lake Michigan sites sampled in Objective 1 will be transported to the Folino-Rorem laboratory at Wheaton College. Facilities at Wheaton for culturing hydroids are well established and are ideal to assess feeding preferences. We will use various benthic invertebrate prey types found in hydranths of hydroids collected in Objective 1 (2-3 chironomid species, amphipods, and isopods). These prey items are common in Lake Michigan and are important prey of benthivorous fish. Individual hydranths or feeding polyps of Cordylophora will be cultured on microscope slides in the laboratory aquaria with Lake Michigan water according to a successfully established protocol. Hydranths will be starved 24 hours before each trial, and we will control for prey size of a given prey type (± 0.5 mm). Two aspects of feeding will be assessed: 1) is the prey item caught by the hydranth's tentacles, and 2) is the prey item consumed by the hydranth. One must consider how prey density may create a confounding effect in predation experiments. Because Cordylophora is a passive predator (contact with prey is necessary for predation to occur), prey will be kept in suspension using a bubbler to facilitate encounters between hydranth tentacles and prey or prey will be delivered to the hydranth using a pipette. All possible pair wise combinations of prey will be administered to at least 20 individual hydranths (n = 20). Lastly, we will expose a given hydranth to prey (prey combinations) for 30 mins (Folino-Rorem laboratory: this is ample time for prey capture and the initiation of consumption of prey). If a given hydranth does not capture and/or consume a prey item, it will be fed brine shrimp, Artemia (the laboratory food item for Cordylophora) to ensure feeding.

Objective 3: Cordylophora effects on non-mussel invertebrates in mussel colonies
To further identify the effect of Cordylophora predation on benthic invertebrate communities, we will address how the presence and absence of hydroid colonies affects benthic invertebrate community composition in zebra mussel colonies. We will establish mussel colonies with and without hydroids. The mussel colonies will be deployed at the two to three locations at a single depth designated in Objective 1 in Lake Michigan. To control for mussel colony size, we will establish zebra mussels on 10 X 10 cm unglazed ceramic tiles attached to bricks. For each site, we will establish 8 replicates of mussels that will be void of the hydroid Cordylophora (non-hydroid treatments), whereas 8 replicates will have the hydroid (hydroid treatment). We will deploy mussel colonies at the beginning or middle of June through July and August when colonization rates of most macroinvertebrates are high (M. Berg, pers. obs.). The colonies will be assessed weekly to bi-monthly to prevent hydroid colonization on non-hydroid treatments; hydroids will be removed if present on non-hydroid treatments. After allowing 2-3 months for invertebrate colonization, the colonies and associated non-mussel invertebrates will be collected and analyzed for macroinvertebrate differences between treatments.

Rationale
The ecological ramifications for this research are significant because a largely unstudied Ponto-Caspian invasive species is being investigated. A better and more thorough understanding of the ecological role of the invasive hydroid, Cordylophora caspia, and its association with zebra mussels in Lake Michigan will help elucidate the potential development of a Ponto-Caspian food web. Our findings will be important because few scientists have assessed the ecological impacts of this invasive hydroid on benthic invertebrate species composition and abundances in southwestern Lake Michigan and the ultimate effects on food-web dynamics. Our results will clarify the indirect effects of yet another invasive species on bottom-dwelling fish in the Great Lakes. If this hydroid is an important predator on macroinvertebrates that are common prey for fish, then this hydroid could have localized effects on fish prey with ramifications throughout benthic aquatic food webs in the Great Lakes.

The information obtained from this study will be presented to colleagues at conferences (North American Benthological Society, International Invasive Species Conference) and to those at interested organizations such as the GLIN (Great Lakes Information Network), Great Lakes Commission, Illinois Natural History Survey and others that would benefit from information on the distribution and impact of Cordylophora on fish prey availability in Lake Michigan. Two manuscripts on various aspects of the proposed research will be written and submitted to peer-reviewed journals (Journal of North American Benthological Society, Journal of Great Lakes Research, Biological Invasions and/or Freshwater Biology).