Seasonal Abundance of the Exotic Predatory Cladoceran, Bythotrephes Cederstroemi, in Western Lake Erie
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J. Great Lakes Res. 14(4):479-488 Internat. Assoc. Great Lakes Res., 1988 SEASONAL ABUNDANCE OF THE EXOTIC PREDATORY CLADOCERAN, BYTHOTREPHES CEDERSTROEMI, IN WESTERN LAKE ERIE David J. Berg and David W. Garton Department of Zoology The Ohio State University 1735 Neil Avenue Columbus, Ohio 43210 ABSTRACT. The predaceous cladoceran, Bythotrephes cederstroemi, has recently invaded the Great Lakes and basic information on its seasonal abundance and population demographics is lacking. Accordingly, we examined seasonal variation in abundance, length —weight relationships, fecundity, ontogeny of lateral spine production, sex ratio, and incidence of resting vs. subitaneous eggs. Seasonal variation in these parameters was compared with water temperature variation and the presence of a potential competitor, Leptodora kindti. Specimens for these analyses were collected with nocturnal surface tows in the western basin of Lake Erie during summer and fall, 1987. Bythotrephes did not appear until 23 July, had its maximum abundance (111 individuals/m3) at the surface on the night of 9 October, and persisted at lower numbers in samples through the last sampling date, 14 December. The correlation of body weight with tail spine length varies seasonally, with individuals weighing less in the summer than in the fall. Neonates possess one pair of lateral spines on the tail spine and add a pair during each of the first two molts, enabling determination of instar distribution. Females produce one to 12 subitaneous eggs per brood, with males and resting eggs occurring rarely. Bythotrephes abundance was negatively correlated with water temperature and Leptodora abundance, suggesting that Leptodora abundance or temperature, or both, affect the success of Bythotrephes in the western basin of Lake Erie. ADDITIONAL INDEX WORDS: Zooplankton, water temperature. INTRODUCTION sions in Bur et al. 1986 and Evans 1988) and much of The discovery of the predaceous Europoean cla- the ecological literature pertaining to the genus doceran Bythotrephes cederstroemi in Lake Huron in Bythotrephes is concerned with B. longimanus (de December 1984, Lake Ontario in September and Lake Bernardi and Giussani 1975, Guma'a 1978) or does Erie in October 1985 (Bur et al. Lange and Cap 1986) not identify which species was investigated has attracted considerable interest from Great Lakes (Mordukhai-Boltovskaia 1956, 1957, 1958, 1960). researchers. Bytho trephes cederstroemi is a large Now reported in all of the Great Lakes, B. ceder- (maximum size greater than 10 mm) cercopagid stroemi's presence provides an opportunity to study characterized by a long caudal spine that often makes the effects on communities of invasion by novel up over 70% of the total length of the organism (Fig. species. However, before this can be done, some of 1). This spine features one to three pairs of lateral the basic demographic and life history characteristics spines, with the number of pairs of lateral spines on of the invading species must be understood. the caudal spine indicating age, since neonates have The North American native cladoceran which one pair of lateral spines and gain an additional pair Bythotrephes most resembles is Leptodora kindti. in each of the next two molts (Ishreyet 1930). The Both species feed on small cladocerans, copepods, caudal spine also has an s-bend which distinguishes and rotifers and consume prey by grinding it exter- the species from its congener, B. longimanus, which nally and then sucking it into the mouth (Mordukhai- possesses a straight caudal spine. The distinctness of Boltovskaia 1958, 1960; Monakov 1972). the two species is still questionable (see discus- Bythotrephes and Leptodora both exhibit 479 480 BERG and GARTON FIG. 1. Female Bythotrephes cederstroemi illustrating length parameters measured. pronounced diel vertical migration, being almost population in a new community, an invading species completely absent from surface waters during the must do more than simply appear in the ecosystem. day (Andrews 1948, Lehman 1987). Throughout Following its initial "inoculation" in the environment, the summer, populations of both species show a high the species must be capable of utilizing an proportion of females, with reproduction occurring unoccupied niche or competing successfully for an parthenogenetically. Males are distinguished from occupied niche. Edmondson and Litt (1982) showed females by the presence of a hook on the proximal that several species of Daphnia were constantly being end of the last segment of the endopodite of the first introduced into Lake Washington, Washington, from leg (Mordukhai-Boltovskoi 1967). The proportion of upstream sources, but only two species were males increases during the fall (Mordukhai- successful in establishing permanent populations in Boltovskaia 1956) and sexual resting eggs are the lake. The remaining species were constantly formed. This represents the overwintering stage introduced but did not form permanent populations. (Andrew and Herzig 1984). Unlike Leptodora, the Once a species has successfully invaded a sexual egg of Bythotrephes does not develop into a community, it may have important and often nauplius larva. In order to successfully establish a unanticipated effects on the invaded community. 481 BYTHOTREPHES SEASONAL ABUNDANCE The introduction of Leptodora into Kansas lakes was characteristics obtained during attempts at laboratory correlated with population declines of Bosmina culture of Bythotrephes. longirostris (Prophet 1982). Stocking of another predaceous plankter, opossum shrimp (Mysis relicta), in North American and European lakes caused MATERIALS AND METHODS dramatic changes in zooplankton assemblages. Detrimental changes in sport fisheries following Seasonal Abundance Sampling Mysis introduction were observed in Lake Tahoe, California-Nevada, and Lakes Selbusjoen and Our earliest sample of Bythotrephes from the central Stugusjoen, Norway (Richards et al. 1975, Threlkeld basin of Lake Erie (near Cleveland, Ohio) was et al. 1980, Langeland 1981). Assessing the impact of collected from downrigger cables of fishermen in July the appearance of Bythotrephes in the Great Lakes 1986, and was preserved in alcohol. A western basin requires determination of its effects on the plankton sample was collected on 14 July 1986, from a water and fish populations of these lakes. However, the inlet at the F. T. Stone Laboratory at Put-in-Bay, basic life history and demographic characteristics Ohio. This and all subsequent samples were preserved needed to determine the effects of the invasion of this in a sugar-formaldehyde solution. species cannot be obtained from completed studies of Cladoceran abundances were estimated using surface the European source population. Most of the tows taken at night. Bythotrephes and Leptodora European literature on Bythotrephes is taxonomic or migrate vertically, concentrating at the water surface zoogeographic (Ishrayet 1930, 1939; Mordukhai- after dark. Surface towing with a 0.75 m net allows Boltovskoi 1967, 1968) or is made up of short, estimation of cladoceran numbers in the upper 0.75 m limited investigations (Mordukhai-Boltovskaia 1956, of the water column. It does not provide an estimate 1957, 1958, 1960). For example, it is known that of abundance below this depth. However, because Bythotrephes feeds upon copepods, cladocer-ans, and Bythotrephes and Leptodora are such strong vertical rotifers (Mordukhai-Boltovskaia 1960, Monakov migrators, we feel this method does provide a 1972) but consumption rates have never been reasonable estimate of relative cladoceran abundances reported. Demographic characteristics of over time and when compared between species. Bythotrephes populations in European lakes do not appear to have been well studied. An accurate A qualitative surface tow was taken on the night of 18 determination of the effects of the appearance of October 1986 in the channel between South Bass and Bythotrephes in the Great Lakes can only be made Middle Bass islands (Fig. 2) using a 1-meter diameter when environmental tolerances, habitat preferences, ichthyoplankton net (mesh size = 0.5 mm). An and basic life history traits of this species are known. additional qualitative surface tow was taken on the Seasonal abundance correlated with environmental night of 1 May 1987. Beginning on the night of 18 parameters provides an indication of the June 1987, quantitative surface tows were taken using environmental factors that are important to the a 0.75-m diameter plankton net (mesh size = 0.5 mm) survival of a population. Positive correlation of equipped with a General Oceanics mechanical Bythotrephes and Leptodora abundances would flowmeter. Samples were taken approximately indicate potential competition between the species weekly (weather permitting), 45 minutes to 1 hour while negative correlation in population size between after sunset (except for 29 July), through 18 the two species could indicate temporal segregation September 1987. Samples were also taken on 9, 15, and lack of competition. Knowledge of life history 22 October, 2, 12, and 23 November 1987. All traits such as sex ratio, age distribution, fecundity, samples were taken in the channel between South and length — dry weight regressions provides Bass and Middle Bass islands (depth approximately information of use in examining the population 10 m), with the exception of the 29 July sample, dynamics of the exotic species. As a foundation for which was taken at 2330 hours, between Green and further studies that