Using an Otolith Growth Chronology as an Archive of Environmental Change Shannon Davis-Foust and Ronald Bruch Sagittal otoliths were embedded in epoxy resin and sections were cut through the core to a thickness of 0.25-0.48 mm with a low-speed diamond blade saw (Fig. 2). Sections were selected based on the clarity of increments in the images to develop the growth chronology. Edge measurements on the otolith sections were not used because the growth for the year of capture is incomplete. The growth chronology index was developed using a mixed-effects linear model (Weisberg et al. 2010) in the program R. Mean daily and monthly air temperatures from 1948 to 2008 were obtained from the National Oceanographic and Atmospheric Administration (NOAA) Satellite and Information Service archives, which were measured near the west side of Lake Winnebago at Oshkosh Wittman Airport (43°59'N / -88°33'W). Growing degree days were calculated annually from high and low daily temperatures using a series of nine threshold (base) temperatures ranging from 0°-18.3°C. Correlations were examined between growth chronologies and climatic data using Pearson’s correlation coefficient (r). The chronology was also entered into the KNMI climate explorer and correlated against gridded, monthly-resolved air temperature, precipitation, and sea level pressure data. -4 chironomid larva 2009 2007 2005 2003 2001 1999 1997 1995 1992 1990 2008 2004 2000 1996 > 400 mm TL Year 2008 Fig. 6 180-362 mm TL 2004 There were a total of 22,373 increment measurements from 1632 fish; ages ranged from 2-58 and years 1948-2008 were included. The high frequency fluctuations of the drum growth chronologies were positively correlated to air temperatures. CDD base 15.6°C provided the strongest correlation with the master chronology (CDD base 15.6°C: r=0.70) (Fig. 3). 12 10 8 6 4 2 0 -2 -4 -6 -8 -10 -12 2000 Results Growth chronologies were compared between small (n=1632) and large (n=461) drum subsets (Fig. 6). Paired Student’s t tests with Bonferroni’s adjustment (α=0.0021) showed that year effects were consistently significantly different between small and large drum every year after 2000, which was shortly after the first report of zebra mussels in Lake Winnebago in 1998. Relative weights (Wr’s) were consistently greater for large drum post zebra mussel establishment versus pre zebra mussel establishment (Fig. 7). 1996 The objectives of this study are to: (1) establish correlations between drum growth chronology and environmental factors, (2) characterize drum diet in relation to body length, and (3) determine whether drum body condition has changed with the establishment of zebra mussels. The third step would provide solid evidence that zebra mussels supply a nutrient-rich food source for molluscivorous fish and therefore directly affect higher trophic positions. 1992 zebra mussel 1964 Objectives 1988 1 0.8 0.6 0.4 0.2 0 Total length (10 mm class) 1960 Fig. 1 1984 Chironomid larvae Zebra mussels Chironomid larvae predicted Zebra mussels predicted 1956 1966-present: Lake sturgeon abundance recovering from low levels (Bruch 2008) 1980 The stomach contents of 987 freshwater drum between 160-640 mm TL were examined, and a distinct body length-related ontogenetic diet shift was observed from chironomid larvae (160-362 mm TL, classified as small drum) to zebra mussels (≥400 mm TL, classified as large drum) (Fig 5). 1952 1955-1966: State intensive rough fish removal period (Staggs and Otis 1996) Relative weight (Wr) values were calculated to determine if body condition of freshwater drum corresponded to growth rates. To search for changes in Wr trends over time in regards to zebra mussel establishment, October trawl captured drum from 1986 through 1999, which was prior to zebra mussel establishment, were compared to 2001 to 2008, post zebra mussel establishment. Comparisons between Wr’s were made between the time periods using ANCOVA where Wr’s are a function of length. error bars = ± 1 s.e. Conclusions • The growth chronology developed from otoliths of freshwater drum in Lake Winnebago, WI, reflects both ambient temperatures (as high frequency fluctuations) and food resource availability (as a low frequency fluctuation). • The correlation of growth rates with food resource availability is supported by the body length-related ontogenetic diet shift from chironomid larvae to zebra mussels and improved body condition of large drum corresponding to the diet shift. • Had the growth chronologies not differed between size classes that had been defined by a diet shift, then another variable such as long-term climate trends could have been implicated as the factor in controlling the recent increase in growth rates. • Zebra mussels can provide enough nutrition to directly enhance drum growth rates, whereas most studies have reported direct effects of zebra mussels only at lower trophic levels. Fig. 4 1948 1936-1990: State rough fish removal (emphasizing drum) (Staggs and Otis 1996) Fig. 2 Year References Year effect 2000-present: Improved water clarity (Bruch 2008) 1998-present: Zebra mussels present (WDNR memo, unpublished) error bars = ± 1 s.e. The drum growth chronology was positively correlated on a regional scale to summer temperatures in the Midwest and negatively correlated to sea surface temperatures in the tropical Pacific (Fig. 4; maps from KNMI Explorer). There were no other significant correlations with climatic data. Fig. 5 1984-present: Best Management Practices reduce nutrient loading (WDNR 2004) 1970-present: Rusty crayfish present (Hobbs III and Jass 1988) 1976 Year 1969: Severe and prolonged flooding event (WDNR 1993) 1900-present: Carp present (WDNR memo, unpublished) 1972 1968 1964 1960 1956 1952 1948 Fig. 3 Fig. 7 1988 60 1960: Severe and prolonged flooding event (WDNR 1993) 1973: Severe and prolonged flooding event (WDNR 1993) 90 70 1992 1960-2000: Reduced water clarity (WDNR, unpublished anecdotal reports) -2 -6 Freshwater drum stomach contents were qualitatively examined (volume was not measured) from 2006-2009. Contents were identified for the presence or absence of the taxonomic levels: Amphipoda, Isopoda, Chironomus spp. larvae, Chironomus spp. - adults, Oligochaeta, Hirudinea, Gastropoda, Bivalvia clams, Bivalvia - Dreissena sp., crayfish, fish, Limnephilidae, Ephemeroptera, microcrustaceans - general, or Mollusca - unknown. In particular, microcrustaceans were all microscopic arthropods, which were likely to be either Cyclops sp., ostracods, Leptodora sp. or Daphnia sp., as described in the diet of Lake Winnebago freshwater drum by (Priegel 1967). For the diet selectivity analysis, 10 mm total length (TL) classes were constructed and proportions of diet items selected were calculated. To determine the mean length at which drum began to consume or stopped consuming major food items, the proportion of drum selecting each food item was calculated and plotted as a function of length. A double logistic model was fitted to the proportion selecting particular food items. 180- 362 mm TL > 400 mm TL 80 1988 1960-2000: High nutrient loading (WDNR, unpublished data) 0 1984 1960-2000: Decline of macrophytes (WDNR, unpublished anecdotal reports) 100 1980 1937-1960: Replacement of floating bog by other macrophytes (WDNR 1993) 2 1976 1850-1937: Large scale formation and disintegration of floating bog (WDNR 1993) 110 1972 1850-present: Water levels maintained at unnaturally high levels (WDNR 1993) 120 4 1968 1850 1875 1900 1925 1950 1975 2000 CDD 15.6°C 160 200 240 280 320 360 400 440 480 520 560 600 640 Growth chronologies have been correlated to temperature for freshwater drum (Aplodinotus grunniens) in Minnesota (Pereira et al. 1995), but not for any other environmental factor, including food availability. Drum are a dominant part of the aquatic community of the shallow Lake Winnebago System. Since European settlement began in central Wisconsin in the early 1800’s, the Lake Winnebago system has faced an onslaught of anthropogenic pressures including loss of thousands of acres of emergent wetlands, hypereutrophication, six decades of rough fish removal, and invasions of non-native species such as common carp, rusty crayfish, and zebra mussels (Fig. 1). Master Proportion It is well-established that growth chronologies developed from relatively stable anatomical structures reflect ecosystem structure and function and can serve as valuable tools to interpret fish community health and regulating factors. Many field studies have established a link between fish otolith growth and temperature, but the effect of resource abundance on incremental growth, while intuitive, has not been reported for fish in a natural environment. Defining relationships between incremental growth and resource abundance is more ambiguous partly because obtaining data to create indices can be intensive, while climatic indices are usually readily available. As a consequence, fish growth chronologies are often hypothesized to be indirectly related to resource availability, such as from eutrophication related to fluctuating water levels or nutrient loading or population density changes related to fluctuating water levels or exploitation. 6 1986 Most of the freshwater drum samples were collected by Lake Winnebago assessment trawling from 1984-2009. Additional samples were collected from fishing tournaments in 2003, 2006, 2009; boom electroshocking and angling in 2009; and from bridge construction blasting in 2007. Total length to the nearest tenth of an inch (mm in 2009) and weight to the nearest tenth of a pound were recorded, and sagittal otoliths were collected for age determination. From 20062009 sex and maturity were determined by the appearance of the gonads. Results (cont.) 1984 The identification of environmental parameters that control biological growth rates is vital to understanding ecological interactions within a community. Growth rates are controlled by the interplay of endogenous and exogenous variables; endogenous variables include age, maturation, and individual growth rates; and exogenous variables include effects from the biotic and abiotic environment (e.g. temperature, photoperiod, precipitation, dissolved oxygen). The summation of these variables constitutes the growth of an individual within a given time frame, and the mean relative contribution of variables influencing total annual growth can be mathematically partitioned to provide information on regulation of growth of an organism or serve as an archive of environmental change. Results (cont.) Wr Methods Year effect Introduction error bars = ± 1 s.e. Bruch, R.M. 2008. Modeling the population dynamics and sustainability of lake sturgeon in the Winnebago system, Wisconsin. Dissertation. University of Wisconsin - Milwaukee. Casselman, J.M. 1987. Determination of age and growth. pp. 209–242. In: S. Gill (ed.) The Biology of Fish Growth, Academic Press, London. Davis-Foust, S.L., R.M. Bruch, S.E. Campana, R.P. Olynyk and J. Janssen. 2009. Age validation of freshwater drum using bomb radiocarbon. Transactions of the American Fisheries Society 138: 385-396. Hobbs III, H., and J. P. Jass. 1988, The crayfishes and shrimp of Wisconsin (Cambaridae, Palaemonidae). Milwaukee Public Museum Milwaukee, WI (USA). KNMI (Koninklijk Nederlands Meterologisch Instituut) Climate Explorer. (n.d.). Retrieved from http://climexp.knmi.nl/ Pereira, D. L., C. Bingham, G. R. Spangler, Y. Cohen, D. J. Conner, and P. K. Cunningham. 1995. Growth and recruitment of freshwater drum (Aplodinotus grunniens) as related to long-term temperature patterns. Canadian special publication of fisheries and aquatic sciences 121:617-629. Staggs, M. D., and K. J. Otis. 1996. Factors affecting first-year growth of fishes in Lake Winnebago, Wisconsin. North American Journal of Fisheries Management 16(3):608-618. Weisberg, S., G. Spangler, and L. Richmond. 2010. Mixed effects models for fish growth. Canadian Journal of Fisheries and Aquatic Sciences 67(2):269-277 . Wisconsin Department of Natural Resources. 1993. Aquatic Macrophyte Ecology in the Upper Winnebago Pool Lakes, Wisconsin. Wisconsin Department of Natural Resources. 2004. Water quality in the Lake Winnebago pool. Wisconsin Department of Natural Resources, Publication FH-229-04, Madison. Acknowledgements This project would not have been possible without the WDNR Fisheries Management crew members who have assisted with obtaining drum samples.