Limnology (2001) 2:185191

The Japanese Society of Limnology 2001

RESEARCH PAPER

Samuel Aikins Eisuke Kikuchi

Water current velocity as an environmental factor regulating the distribution of amphipod species in Gamo lagoon, Japan

Received: October 10, 2000 / Accepted: August 17, 2001

Abstract The role of current velocity as an environmental factor affecting the distribution of amphipod species was investigated in the brackish Gamo lagoon, Japan. Two gammaridean amphipod species, Eogammarus possjeticus and Melita setiagella, were found inhabiting stations of different water current with same substrates. E. possjeticus and M. setiagella selected substrates at low and high water current, respectively. The lifestyles and anatomical features of these two free-living amphipods contributed to their adaptation to habitats with different water currents. The results of eld and experimental observations suggest the existence of some type of interspecic competition between the two species. The presence of E. possjeticus caused a displacement of the fundamental niche of M. setiagella. Key words Amphipods Water current velocity Articial substrates Niche shift Brackish lagoon

Introduction
An ecological community, an assemblage of interacting species, is structured by various interrelationships linking the individual species. The relationships multiply with the increase in species richness of a community and many linkages are subtle or ephemeral. Species diversity in a community is closely related to habitat stability and several different explanations have been put forward for the degree of diversity. According to the stability-time hypothesis of Sanders (1968), in a habitat where physiological stress is low, biotically accommodated communities may evolve.

S. Aikins (*) Biological Institute, Graduate School of Science, Tohoku University, Sendai 980-8578, Japan Tel. 81-022-217-7561; Fax 81-022-217-7560 e-mail: aikins@mail.cc.tohoku.ac.jp E. Kikuchi Center for Northeast Asian Studies, Tohoku University, Sendai, Japan

Under greater physical stress, resulting from either an increase in physical uctuations or unfavorable physical conditions, communities may become predominantly physically controlled, resulting in decreased species diversity. Environmental factors such as temperature, salinity, sediment type (substratum composition), light intensity, food availability, space, and wave action/exposure (Moore 1973; Fenwick 1976; Van Dolah 1978) have been reported to inuence the distribution of aquatic organisms in their ecological communities. Dommasnes (1968) studied the effect of water movement on algal fauna, especially the abundance of some isopods and amphipods inhabiting Corallina ofcinalis; he concluded that water movement affects the fauna both indirectly through its effects on algal growth and directly through mechanical drag. Hiscock (1983) reported that tidal streams, wave action, and residual currents are the most important environmental factors that determine the community present in a particular location, and thus water movement is both an important mechanical environmental factor and a carrier for other environmental factors such as dissolved gases and silt. Moore (1973) concluded that the amount of detritus deposited within the kelp holdfast was the most important factor inuencing the fauna and that the direct effect of water movement may be of minor importance within the kelp holdfast. In this study, water current velocity was considered to be a major environmental regulator, triggering an interaction between two amphipod species in a brackish lagoon. Naturally high current velocity exerts stressful conditions through mechanical drag on most free-living species and this may lead to survival of the ttest in favorable habitats. Well-adapted species may drive away or force other species to occupy new habitats. In a study of habitat selection by amphipods, the two free-living species Eogammarus possjeticus Tzvetkova and Melita setiagella Yamato were found to inhabit algal species including Gracilaria vermiculophylla (Ohmi) Papenfuss and Polysiphonia fragilis Suringar at different locations in Gamo lagoon (Aikins and Kikuchi 2001). Among prevailing environmental factors, water current velocity differed

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Fig. 2. Water current velocity at ve stations in Gamo lagoon during ood and ebb tides and at low water level. Bars, SD, n 5. St., station

low tide, sandy mud ats appear at the central part and outside the levee, while water remains in the innermost part of the lagoon (Kikuchi et al. 1980). Field experiments Field experiments were carried out in Gamo lagoon to determine the inuence of water current velocity on the distribution of amphipod species. A digital current meter (UC-3, Tamaya, Tokyo, Japan) was used to measure current velocities during ood tide, ebb tide, and low water level. Readings were taken at 100-m intervals, starting from the levee of Gamo lagoon to 500 m away from it (Fig. 2). Based on differences in water current velocities and the path of water movement, ve stations (1 to 5) were selected in the lagoon (Fig. 1). Two types of articial substrate, cylindrical brushes and circular concrete plates, were set up for the eld experiments at all ve stations in September 1998. The cylindrical brushes (Melogy, cleaning brushes for bath pipes, Azuma Kogyo, Hamamatsu, Japan), 5.0 cm diameter and 10.5 cm thick, was made from polypropylene strings (length of string 2.4 cm); the circular concrete plates had a diameter of 8.5 cm and thickness of 1.5 cm. At each station, two lines 50 cm apart were set parallel to each other. Each line was marked by two wooden poles 1 m apart, which were connected with a nylon rope. Using nylon string, the articial substrates were hung on the rope at intervals of 20 cm (Fig. 3). Four substrates consisting of two brushes and two concrete plates were set on each line, making four replicates of each substrate at each station. All substrates were submerged even at low water level. Substrates were sampled after 14 days of submergence. Labeled sampling polyethylene bags were used to wrap the hanging substrates and then removed from the rope. Samples were immediately xed and preserved in 4% neutralized formaldehyde for further treatment in the laboratory. All substrates were vigorously washed several times to remove organisms and attached materials by pouring super-

Fig. 1. Map of Gamo lagoon and experimental stations (15) for the distribution study

signicantly between these locations. Moreover, a eld experiment suggested that one of the main factors strongly related to the distribution of amphipod species, especially E. possjeticus and M. setiagella, was water current velocity (Aikins and Kikuchi 2001). The experiments were carried out in the eld in Gamo lagoon and in the laboratory to elucidate the role of water current in the distribution of amphipod species.

Materials and methods

Study site Field investigations and experiments were carried out in an estuarine lagoon at Gamo (Fig. 1). This is a small brackish lagoon located at the mouth of the Nanakita River running through the northern part of Sendai City, northeastern Japan (3815N, 14100E), with a total area of about 10 ha. A stone levee with three rectangular entry channels forms the gateway through which lagoon water is exchangeable with tidal movement. Salinity varies from 5 to 30 at the center of Gamo lagoon due to tides (Kurihara et al. 1997). The average minimum and maximum daily water temperatures were highest during July and August at 25C and 30C, respectively, and lowest during January and February at 3C and 6C, respectively (Kurihara et al. 1997). During

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Fig. 3. Set-up of articial substrates used in eld experiments

natant solution through a 0.25-mm mesh sieve. Washing and sieving were continued until no organisms appeared in the solution. Invertebrates were then sorted with the aid of a hand lens and preserved in 70% ethanol. Macrofauna were identied to species level and counted under a stereoscopic microscope. Laboratory experiment In the laboratory, a water current chamber was constructed to investigate the role of current velocity in the distribution of amphipod species. The water current chamber was composed of a transparent tank 200 cm 15 cm 20 cm in which water was circulated to generate current (Fig. 4). Two water pumps, (SL-101 (N), Terada, Yamatotakada, Japan), were placed at one end of the tank. These pumps were connected by water hoses with diameters of 25 and 15 mm to the other end of the tank, which is referred to as the source side. Two-thirds of the tank depth was lled with water that circulated within the tank to generate a water current. Bricks 20 cm 5 cm 12 cm were placed in the transparent tank to enhance the water current. Using the digital current meter, current velocities were measured 20, 40, 60, 80, 100, and 120 cm from the mouths of the water hoses. To generate different water current velocities at the same sites, the two water hoses were either combined or used singly. When the hoses were combined, the diameters were 25/25, 25/15, or 15/15 mm, and when single hoses 25 or 15 mm in diameter were used only one pump was employed, as shown in Fig. 5. To reect the range of water current velocities in Gamo lagoon, two-hose combinations were selected and used throughout the laboratory experiments. Brush substrates were set up at sites of known current velocities, because among the seven articial substrates used to analyze habitat selection by amphipods, it was the articial substrate preferred by the amphipods in Gamo lagoon (Aikins and Kikuchi 2001). A net of mesh size of 1 mm2 was used to separate the main stream from the pumps. E. possjeticus and M. setiagella collected from the lagoon were used for the laboratory experiments. Two types of experiments including pure and mixed treatments were carried out in the laboratory. Pure treat-

Fig. 4. Water current chamber to investigate the effects of water current velocity on the distribution of two amphipod species. Upper and lower diagrams show side and top views, respectively

Fig. 5. Water current velocities in the water current chamber using different hose sizes (diameter in mm), either singly or in combinations. Distance was measured from the mouth of the hoses (see Fig. 4 for experimental set-up)

ment consisted of a single species inoculated in isolation at the source. For mixed treatment, E. possjeticus (body length 1012 mm) and M. setiagella (69 mm) were inoculated together into the main stream. Twenty-ve adult individuals were inoculated during pure treatment experiments, and 20 adult individuals of each species were used for mixed treatment experiments. A control experiment was carried out by inoculating both species of amphipod under zero current velocity. The number of animals of each species on brush substrates was recorded at 30, 60, and 120 min postinoculation, as well as at 10 min in the control experiment.

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Results
Water current velocity Water current velocity (cm/s) in Gamo lagoon is shown in Fig. 2. The fastest water current velocity was recorded during ood tides, and the slowest at low water level. During the ood tide, the highest velocity of about 52 cm/s was recorded 100 m from the levee. The velocity decreased to about 45 cm/s at 200 m. After 200 m, there was a drastic decrease in velocity through 300 to 400 m to less than 10 cm/ s at 500 m. At ebb tide, a current velocity of about 30 cm/s was recorded 100 m from the levee. This decreased steadily to about 10 cm/s at 400 m and to less than 10 cm/s at 500 m. At low tide, water movement was at its lowest at all stations recording velocities below 10 cm/s. Density of amphipods on articial substrates Densities of amphipods in terms of number of individuals per square meter on each articial substrate are summarized in Table 1. Four species of amphipods, two free-living (E. possjeticus and M. setiagella) and two tube-dwelling species (Corophium uenoi Stephensen and Grandidierella japonica Stephensen) were found on the substrates. These species were generally found abundantly on algae (G. vermiculophylla, Enteromorpha prolifera Muller, and P. fragilis) in Gamo lagoon. Both free-living forms and tube dwellers selected and inhabited brush substrates rather than concrete ones. The concrete substrate was mainly selected by tube dwellers. C. uenoi, E. possjeticus, and G. japonica recorded maximum densities at station 4, while the highest density of M. setiagella was recorded at station 2, irrespective of substrate. Of the free-living species, M. setiagella dominated sites with stronger currents and E. possjeticus those with weaker currents. The brush substrate at stations 1 and 2 with cur-

rent velocities of a maximum 52 cm/s (ood tides) had high densities of M. setiagella, and stations 4 and 5 with a maximum velocity of 20 cm/s recorded high densities of E. possjeticus (Table 1). The same trend was also exhibited on the concrete plates; M. setiagella was conned to sites with stronger currents (stations 2 and 3), while E. possjeticus was conned to sites with weaker currents (stations 4 and 5). Laboratory experiments Current velocity in chamber Current velocities (cm/s) at 20, 40, 60, 80, 100, and 120 cm in the water current chamber are shown in Fig. 5. Combinations of two water hoses (25/25 and 25/15 mm in diameter) recorded faster current velocities as compared to single hoses 25 and 15 mm diameter at some sites. The combination of 25/ 25 mm hoses recorded the fastest velocities at all sites. A velocity of 85 cm/s was recorded 20 cm from the mouth of hose with this combination. Between 40 and 120 cm, current velocities ranged from 77 to 25 cm/s and 50 to 12 cm/s for hose diameter combinations of 25/25 mm and 25/15 mm, respectively. Using single hoses 25 and 15 mm in diameter, current velocities between 120 and 20 cm were almost the same for each site, ranging from about 70 to 8 cm/s. Comparing water current velocities in Gamo lagoon (Fig. 2) with those in the experimental chamber, hose diameter combinations of 25/15 mm and sites between 40 and 120 cm were used for the laboratory experiments. Distribution of amphipods in water current chamber The number of E. possjeticus decreased with velocity in a similar way at three measurement times (Fig. 6), and about 60% of the inoculated individuals (25 adults) were found at sites with velocities of less than 16 cm/s. The maximum number of M. setiagella was recorded at sites with medium current velocity (approximately 25 cm/s) at all measurement

Table 1. Mean densities per surface area (no. m2 ) with standard deviation of amphipods inhabiting brush and concrete plate substrates in the eld stations Species Station 1 Brush Tube dwellers Corophium Grandidierella Free-living forms Eogammarus Melita Concrete plate Tube dwellers Corophium Grandidierella Free-living forms Eogammarus Melita 2 3 4 5

1364 141 451 159 0 1071 334

2822 20 1320 446 501 111 2782 534

2883 128 962 113 1660 412 697 111

4390 297 1475 193 4062 644 515 74

3331 158 0 3966 476 0

200 23 100 36 0 0

821 57 0 0 112 59

1069 151 481 124 100 32 100 32

1099 168 827 221 486 129 0

762 145 0 223 126 0

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Fig. 6. Distribution pattern of E. possjeticus at 30, 60, and 120 min after inoculation into the water current chamber. Bars, SD, n 5

Fig. 7. Distribution pattern of M. setifagella at 30, 60, and 120 min after inoculation into the water current chamber. Bars, SD, n 5

times (Fig. 7). More M. setiagella individuals (16% of inoculated animals) were recorded at sites with velocities of more than 30 cm/s than E. possjeticus individuals (11%). However, the number of individuals (45%) recorded at sites with low current velocity of less than 20 cm/s exceeded that (16%) recorded at sites with high current velocity of more than 34 cm/s. As mentioned above, distribution trends were the same irrespective of time for both E. possjeticus and M. setiagella when inoculated in isolation. Inoculation of both species together (20 adult individuals per species) resulted in a pattern similar to that found in the eld (Fig. 8). As many as eight M. setiagella individuals were found on a brush substrate at 60 cm with current velocity of 34 cm/s at 30 min. At 60 and 120 min, the maximum of seven individuals occurred at 80 cm and 60 cm with velocity of 24 cm/s. At least four individuals of M. setiagella were recorded at 40 cm with velocity of 53 cm/s at all measurement times. The fewest M. setiagella were found at 100 cm or more with velocity of less than 16 cm/s.

At least nine E. possjeticus individuals were recorded at the 120-cm site with velocity of 15 cm/s at all measurement times. This number decreased markedly to the minimum at sites with velocities of more than 25 cm/s at all readings. Sites with rapid current between 30 and 55 cm/s were thus dominated by M. setiagella, with more E. possjeticus recorded at sites with slower current of 20 cm/s or less. Recordings at different times did not show a change in the pattern of distribution.

Control experiment The results of the control experiment are shown in Fig. 9. Twenty adult individuals of each species were inoculated in a mixed treatment. The number of individuals of each species found swimming in open space or clinging to brush substrates were counted at regular time intervals. Seventeen E. possjeticus individuals were found swimming 30 min

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after inoculation. However, the number decreased to 10, and a maximum of six individuals were clinging to the brush substrate after 120 min. More individuals of this species were found swimming at all recording times compared with those clinging to the brush substrate. A maximum of 18 M. setiagella individuals were clinging to substrates 10 min after inoculation. Meanwhile, the maximum number of swimming individuals was three 30 min after inoculation. Most M. setiagella clung to substrates rather than swimming in open space. In this experiment, animals found under or behind the bricks used as packing were not counted.

Discussion
Other researchers, such as Hagerman (1966), Dommasnes (1968), and Moore (1973), have studied how water current velocity affects the distribution of aquatic organisms. They found that animals with strong grasping appendages, such as amphipods, often dominate in algal fauna at exposed sites (high water current). Hagerman (1966) reported that they were able to do this through their ability to hold rmly to algae. In Gamo lagoon, the distribution of E. possjeticus and M. setiagella may be inuenced by current velocity. Hiscock (1983) reported the relations between water movement and other environmental factors such as dissolved gas transport and siltation. Silt is high in sites with low current velocity as compared with high current sites in Gamo lagoon (Kanaya, personal communication). Silt deposition increases with decreased water movement and may inuence the abundance of some species. However, species with special adaptations can cope with siltation and survive in sites with high silt deposits. Concerning dissolved oxygen as an environmental factor, the lagoon is well aerated due to its shallow nature, with seasonal variations occurring in sites with low water movement (Kurihara et al. 1997). In Gamo lagoon, G. vermiculophylla that occurs mainly at lower current sites tends to be dominated by E. possjeticus, while P. fragilis attached to dead oyster shells at more rapid current sites are dominated by M. setiagella (Aikins and Kikuchi 2001). The structure of the brush substrate used in the experiments was similar to that of G. vermiculophylla and P. fragilis but not to that of the concrete plate substrate. Results from both eld and laboratory experiments suggest that amphipod species are able to inhabit areas of high current velocity (Table 1 and Fig. 8), as described by early workers. However, not all amphipod species found during this investigation had the capability to inhabit very high current velocity sites. Even though amphipods generally can hold rmly to their substrates with their strong grasping appendages, not all species have the ability to grasp substrates for long periods. The type of substrate is important since smooth, hard substrates (such as concrete plates on which low amphipod densities were recorded, Table 1) subject amphipods to greater risk of being washed away, even if they have the ability to cling.

Fig. 8. Distribution pattern of M. setiagella ( lled circles) and E. possjeticus (open squares) at 30, 60, and 120 min after mixed inoculation into the water current chamber. Bars, SD, n 5

Fig. 9. States of condition of M. setiagella (circles) and E. possjeticus (squares) chamber. Black and white symbols indicate the swimming and clinging state, respectively. Bars, SD, n 5

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E. possjeticus and M. setiagella selected substrates at sites with different water current velocities (Table 1). Both species are free-living and equipped with strong appendages. High densities of E. possjeticus were found at sites with low water current velocity and of M. setiagella at stations with high water current velocity when both species were inoculated in the laboratory experiment (Fig. 8), as also demonstrated in the eld (Table 1). Field observations showed that during feeding E. possjeticus do not grasp lamentous substrates rmly. When the water current becomes much stronger, they will not be able to cling to their substrates for a long period and thus may be washed away. In the laboratory experiments, under zero current velocity conditions, E. possjeticus swam more frequently than M. setiagella (Fig. 9). They rarely clung to brush substrates for a long period. This trend was observed with water current as well: E. possjeticus clung to their substrates for only a short time as compared with M. setiagella. The appendages of both species are similar but vary slightly, and they have strong gnathopods and pereopods, but the smaller M. setiagella have larger pereopods than E. possjeticus. Functionally, these pereopods are used for clinging and grasping (Smith and Carlton 1975). M. setiagella were not easily separated from their substrates during sorting. Under zero current velocity conditions, M. setiagella clung strongly to substrates. This characteristic behavior of M. setiagella could be an adaptive feature contributing to its ability to select substrates even under harsh conditions of high water current velocity. In a eld experiment to develop amphipod communities, M. setiagella selected most of the introduced substrates, while E. possjeticus selected mainly brush-type substrates (Aikins and Kikuchi 2001). If these free-living amphipods basically use their pereopods for clinging, then that may be one reason why E. possjeticus clings only to lamentous substrates and M. setiagella to various types of substrate. The results of the pure experiment with M. setiagella (Fig. 7) show that more individuals selected low- and medium-velocity current sites rather than high-velocity sites. Due to the dragging effect of high water current velocity, it may be stressful irrespective of species but especially for free-living organisms. In the eld experiment, the densities of most species tended to be lower at the highest-velocity current station (station 1) than at lower-velocity current stations (stations 3 or 4). The fact that M. setiagella selected fast current zones in the presence of E. possjeticus suggests some type of interspecic competition. Comparing the pure and mixed treatment results, while E. possjeticus maintained its fundamental niche, a displacement of M. setiagella from its fundamental niche was observed. Thus fundamental and realized niches (as proposed by

Hutchinson 1965) were the same for E. possjeticus but different for M. setiagella. Further experiments are needed to conrm the nature of this interaction. Even though silt deposition has been reported to inuence the distribution of amphipod species, the results of laboratory experiments suggest that water current velocity is a major important environmental factor affecting the distribution of E. possjeticus and M. setiagella in Gamo lagoon. The interactions and differences in behavior between E. possjeticus and M. setiagella may also be a contributing factor to their occurrence at slow and fast water current velocities, respectively. Thus both environmental and biological factors affect the population distribution of these amphipod species in Gamo lagoon.
Acknowledgments We are grateful to Drs. M. Nishihira and S. Shikano for their advice and contribution to this manuscript. Special thanks are due to Mrs. L. Aikins for reading the proof of this manuscript, and our profound gratitude is expressed to the students of the Animal Ecology Laboratory of Tohoku University for their assistance in eld sampling.

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