Verh. Internat, Verein. Limno!.

Zooplankton community structure: the role of dispersal

John E. Havel and Jennifer Stelzleni-Schwent

Introduction

Zooplankton present an interesting paradox with respect to dispersal. Most freshwater zooplankton have the capability to form resting eggs, which can tolerate freezing, drying, and digestion by fish and birds (DODSON & FREY 1991). Furthermore, the cladocerans and rotifers reproduce by parthenogenesis, suggesting the capability of single hatchlings to establish populations following dispersal over land. Early experiments suggested that passive dispersal of freshwater organisms should work well, at least over short distances. (MAGUIRE 1963), with the movements of waterfowl a likely vector (PROCTOR & MALONE 1965). However, numerous allozyme studies indicate that cladocerans, copepods and ostracods show strong genetic divergence among even closely neighboring populations (BOILEAU et al. 1992), indicating that successful colonization is probably rarer than was previously recognized.

The movements of man provide new mechanisms for dispersing freshwater organisms over long distances (CARLTON 1992). The large volumes of ballast water of large ships can transmit a wide variety of exotic species from one continent to another (CARLTON & GELLER 1993), and the transcontinental shipment of exotic fish may introduce exotic zooplankton into inland reseryOirs(HAVEL & HEBERT 1993). The bait buckets, bilges and live wells of recreational fishing boats provide additional vectors for moving freshwater species to new watersheds (CARLTON 1992, JOHNSON & PADILLA 1996). Boats have been previously linked to the dispersal of exotic weeds GOHNSTONE et al. 1985) and zebra mussels GOHNSON & CARLTON 1996).

The exotic cladoceran Daphnia lumholtzi provides a good example of a zooplankter probably dispersed by boats. D. lumholtzi was first discovered in North America in the early.1990s (SORENSEN & STERNER 1992) and rapidly spread across the southern and midwestern United States (HAVEL & HEBERT 1993). D. lumholtzi has been detected in a wide variety of

reservoirs, where the is a mid-summer domi-

nant (HAVEL et al. and has also invaded

rivers (THORP et al. 1996)

riparian wetlands. Yearly surveys of over 100 Missouri reservoirs indicated that D. lumholtzi rapidly increased its range and, over a 4-year period, increased its prevalence from 6% to 26% of the reservoirs (HAVEL, unpublished data). The appearance of new populations in different drainage basins up to 170 km from the nearest source lake suggested that recreational fishing boats are important in the transport of D. lumholtzi and other zooplankton into new lakes.

In the following paper, we present evidence from boater surveys and survivorship. experiments which support the boater hypothesis for long-distance transport of zooplankton among lakes.

Materials and methods

Boater surveys

We surveyed recreational fishermen as they arrived at 14 boat ramps on three Missouri reservoirs between 05:00 and 09:00, June-July 1996. Each fisherman was interviewed and asked to contribute a live well sample. A total of 65 fishermen were approached, of whom 62 consented to being surveyed. Live wells were sampled with a turkey baster (large pipette), volume measured, then returned to the lab on ice. Each sample was examined at 25x magnification on the same day as collection. All zooplankton were counted, tallied as dead or alive, and identified (EDMONDSON 1959, DODSON 1994, HEBERT 1995).

Live well survivorship experiments

We examined the survivorship of D. lumholtzi and the native Daphnia parvula in the live wells of stored boats. A single clone of each Daphnia species was cultured in the laboratory in 53-flm filtered lake water, with 105 cells/mL Ankistrodesmus falcatus as food. After rinsing the live wells and adding media, Daphnia were introduced, the wells covered, and the Daphnia removed 1-7 days later and checked for viability.

Experiments 1 (May 1997) and 2 (july) used 32 new Tracker boats, loaned by a local boat dealer, and in the sun on the dealer's parking lot. Experi-

$ 1.25

J. E. Havel & J. Stelzleni-Schwent, A long-distance dispersal hypothesis

ment 1 was designed to compare the effect of food level on survivorship of the two species. Low food levels had filtered lake water alone and high food levels were supplemented with Ankistrodesmus (10' cells/mL). Twenty Daphnia were introduced into each boat along with 100 mL of medium. Each species/food level combination was placed in eight different boats, and survivorship checked at 1, 2, 3 and 7 days. We terminated one set of four replicates after 2 days and the other set after 7 days, with 1- and 3- day endpoints checked in the field and 2- and 7 -day endpoints checked in the lab. Since some reproduction occurred during the experiment, we counted only adults. The air temperature ranged from 17 to 30°C. During the experiment, several boats were sold, so some replicates were lost from the experiment.

Experiment 2 used the same two species, at a constant high food level in a paired comparison design. For each of 20 boats, 10 juveniles of each species were introduced together into 100 mL of media, and survivorship assessed after 3 days. During this experiment, the air temperature ranged from 28 to 32°C. One of the boats was sold during the experiment.

Results

Boater movements and other behaviors

Although the majority of fishermen usually visited the same lake, 25% of all the surveyed fishermen planned to visit a different lake on their next fishing trip, and 69% regularly transported their boats to different lakes. These lakes cover a wide area, including at least 19 lakes in eight states, many of which contain D. lumboltzi (Table 1). As an example, the reported movements of the 40 boaters visiting Stockton Lake are shown in Fig. 1. The respondents frequently use their boats, with 31 % (n = 62) having fished in the last 3 days. Although all the fishermen drained their live wells, only 20% regularly flushed the wells after fishing and only one used any method of sterilization (bleach).

Live well zooplankton survey

Of the 47 boats with live wells, 29 had wells containing water, with a volume averaging 208 mL (range 5-700 mL). (A typical live well has a capacity of 40-50 L.) The remaining 18 boats had live wells which were dry. Thirty eight percent of live wells with water contained invertebrate taxa (Fig. 2), including at least 12 species

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Fig. 1. Lakes last visited and lakes where fishermen entering Stockton Lake, Missouri, plan to go next. Lakes are all in Missouri, except where indicated.

. Numbers on arrows indicate the percentage of boats coming from or going to a specific lake. Survey based upon interviews with 40 fishermen, June-July • 1996

Taxa in Live Well Samples

40 35

~ 30 :5 25 .~ 20 !!!

~ 15

'# 10

n = 47 boats

_alive

5 o

I i

e

e

Fig. 2. Taxa in samples from boat live well samples, collected June-July 1996. Eighteen of the 47 live wells were dry.

of cladocerans and copepods plus chironmids later reared from mud in the lab (Table 2). Live copepods were common in the live wells, as were the cladocerans Bosmina and Chydorus. All individuals of the seven Daphnia species were dead and no ephippia were encountered in the live wells, although Daphnia ephippia have been previously observed (M. EISENBACHER, per-

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Environmental heterogeneity in limnology

Table 1. Lakes commonly visited by fishermen surveyed in June and July of 1996. Numbers indicate number of fishermen at present location who regularly visit the lake in column one. Numbers do not add to totals, as some fishermen visited multiple lakes. Total fishermen surveyed: 62.

Present Day Fishing Location

Pomme de Terre Stockton Table RoCk Total

Arkansas:

3

Beaver" Norfolk" Ouachita

Georgia Lakes
Indiana Lakes
Kansas:
Clinton"
Missouri:
Bull Shoals* 3 6
Mark Twain*
Niangua River
Lake of the Ozarks* 1 1 3
Pomme de Terre" 8 4
Sac River 1
Stockton" 3 36 4
Table Rock* 2 13 14
Taneycomo 1
Truman" 4 2 3
Wappapelo*
Nebraska Lakes
Oklahoma: -"
Grand* 4
Texas:
Fork 1
Total lake visits 19 68 41 128
Total fishermen surveyed 8 40 14 62
*Lakes with Daphnia lumholtzi previously detected (HAVEL et al. unpublished). sonaI communication). In contrast to other studies OOHNSTONE et aI. 1985), none of the 62 boats we observed had vegetation attached to the boat or trailer.

Live well survivorship experiments

Results from experiment 1 are shown in Table 3. Survivorship of both species was high for the first 3 days, then declined to 0% by day 7. Survivorship of D. paruula was significantly greater than that for D. lumholtzi, at least for the first 2 days of the experiment. Two-way ANOVA for each of the first 3 days of the experiment indicated no significant effect of food level on survi-

vorship, except for day 1 (F1,9 = 5.29, P = 0.047).

Survivorship of both species during experiment 2 was poor, with only 6% of D. parvula and 0% D. lumboltzi surviving to day 3 (Table 3). Over this period, the weather was sunny, windy and warm (maximum air temperature 32°C). On day 3, the volume of the live wells had diminished to a smaIl volume (median 10 mL, range 0-60 mL). The water temperature in these wells ranged from 35-40 DC, above the upper thermal limit for both species (BOECKMANN & HAVEL, unpublished data).

J. E. Havel & J. Stelzleni-Schwent, A long-distance dispersal hypothesis

Table 2. Organisms found in live wells, pooled from
42 boats (nc, too numerous to be counted).
Number Alive?
Identified
Branchiura 2
Cladocera
Bosmina cf .. longirostris 55 +
Chydoridae 16 +
Daphnia ambigua 16
Daphnia exilis
Daphnia galeata mendotae 12 ;;'
Daphnia laeois 2
Daphnia lumholtzi 1 - ~.;~.
Daphnia parvula 3
Daphnia retrocurua 2
Copepoda
Acanthocyclops robustus 277 +
Diaptomus reighardi 1 +
Diaptomus sp. 7 +
Mesocyclops edax 2 +
Nauplii 2 +
Unidentified calanoids 1
Unidentified cyclopoids 2 +
Insecta
Chironomid larvae (hatchlings) 255 +
Mosquito larvae nc +
Unidentified insects nc +
Nematodes nc +
Oligochaetes nc +
Protozoans nc +
Rotifers nc + Discussion

Although waterfowl have long been considered significant vectors for the overland dispersal of freshwater invertebrates, both population subdivision (BOILEAU et al. 1992) and demographic analyses QOHNSON & PADILLA 1996) suggest they have limited importance for many crustaceans and molluscs. Furthermore, the late-summer abundance of D. lumholtzi is poorly matched with the seasonality of waterfowl, which generally spend much of this period nesting and feeding QOHNSON & CARLTON 1996). A more-likely explanation of the rapid saltatory dispersal of D. lumholtzi is by the movements of boaters.

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Table 3. Zooplankton survivorship in 1997 live well experiments. For experiment 1, different groups of boats were checked after 1- and 2-day and 3- and 7- day endpoints. P-values are from hypotheses tests for differences between species: two-way ANOVA (food level and species), at 9 df (experiment 1), or from a paired t-test, at 18 df (experiment 2).

Survivorship (%)
Experiment Duration D. lumholtz; D.parvula P-value
(days)
1 1 72 96 <0.001
2 43 72 0.02
3 45 44 0.97
7 0 0
2 3 0 6 0.10 In the current study, the majority of recreational boaters regularly transport their boats to different lakes and nearly a third fish frequently (within 3 days), a time period during which zooplankton survival is high (Table 3). The fact that few boaters drained and flushed their live wells suggests that they could readily move zooplankton among lakes. Of the fishermen surveyed, 74% store their boats protected from the sun, which should enhance conditions for survivorship of the zooplankton. However, the experiments suggest that, over longer periods or in extremely hot weather, the zooplankton in live wells of unprotected boats would suffer high mortality. Thus dispersal seems unlikely between far away lakes during the hottest part of the summer.

Consistent with these data, a large proportion of live wells contained living copepods and cladocerans (Fig. 2). None of the Daphnia were recovered alive, suggesting that these species were less tolerant of conditions in the live wells than were the other taxa. However, because of their parthenogenetic mode of reproduction, even rare survivors or ephippia could be a sufficient innoculum for colonization.

The movements of boaters thus provide a means for long-distance dispersal of zooplankton. SAUNDERS (1993) proposed that the disjunctive distribution of Eurytemora affinis may be linked to boaters. Similarly, the rapid dispersal of D. lumholtzi among reservoirs in different drainages may follow recreational boaters

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Environmental heterogeneity in limnology

or the long distance movements of professional bass fishermen.

Acknowledgements

We thank the numerous students who assisted with our pre-dawn trips to fishing ramps. ALLISON HICKMAN also helped with the 1997 survivorship experiments and DESIREE BETHUNE identified the copepods. MATI EISENBACHER'S insight into bass fishing helped us to better design the surveys of boaters. We are grateful to BILL KOSTELLO of Bass Pro Shops, Inc., for giving us permission to use their new Tracker boats in the survivorship experiments. J. STELZLENI-SCHWENT was supported by an NSF Research Experiences for Undergraduates grant (DEB 96-41390) to J. HAVEL.

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Authors' addresses:

J. E. HAVEL, J. STELZLENI-SCHWENT, Department of Biology, Southwest Missouri State University, Springfield, Missouri, 65804-0095, USA.