Ecology of North American Freshwater Fishes. Stephen T. Ross Ph. D.
Читать онлайн книгу.Weighting in Studies of Fish Movement
An appropriate experimental design is critical for assessing fish movement using mark-recapture approaches. This is especially so because the likelihood of capturing a marked fish declines with distance from the point of release, leading to the risk of underestimating longer movements. A robust experimental design is an important issue even with essentially linear stream systems—additional complexity of the aquatic system (e.g., tributary streams, lakes with numerous coves, etc.) further increases the challenge of obtaining reliable data. Following Rodríguez (2002), in quantitative terms, the density of marked fish multiplied by meters away from the region of their release, n(x), is given by
where No is the number of fish originally marked, s is the probability of their surviving to the sampling period, π is the catchability of the fish, and f(x) describes the decline in density as a function of the distance from the release area (referred to as a dispersal function). The key point that this equation makes is that the number of recaptures at any given location must be evaluated relative to the probability of recapture.
Albanese et al. (2003) evaluated the impact of distance weighting on the assessment of movement by three species of southeastern stream fishes. He used a modeling study to illustrate the impact that increasing the number of 50 m sampling sections would have on distance weighting. Fish were considered marked in ten 50 m sections. The modeling approach showed how the zone of uniform sampling (i.e., sampling at or near 100%) changed as the sampling area was increased. If sampling only occurred within the 500 m marking section, then the impact of distance weighting was extreme. The proportion of total possible movements sampled (PSd) was only 100% for sampling within ± 50 m (i.e., the zone of uniform sampling) and declined sharply for movement distances > ± 50 m. Clearly, a study design that only included sampling within the same stream reach used to mark fish would be strongly biased toward short-term movements. With a 1,000 m sampling effort, the PSd values were 100% for fish movements within ± 250 m, and with a 2,000 m sampling effort, PSd values are 100% for fish movements within ± 750 m. Both of these are much more robust designs in terms of understanding movement. However, even with the 2,000 m sampling effort, any fish movements greater than ± 1,200 m would have been undetected.
In the following figure, based on Albanese et al. (2003), the modeling study shows the proportion of total possible movements (PSd) that would be detected for three different sampling designs: one in which the sampling reaches were the same as the marking reaches (500 m), one in which the sampling reaches (1,000 m) were twice that of the 500 m marking reach, and one in which the sampling reaches (2,000 m) were four times that of the 500 m marking reach. Solid lines show PSd values; dashed lines show the sampling lengths. Movement can be either upstream (+) or downstream (−) from the marking section.
The effect of the size of the resampling areas (indicated by dashed lines) on the ability to detect marked fishes, relative to the distances that the fishes move. As determined by a modeling study, the largest resampling area (2,000 m) can detect 100% of fish movements up to 750 m upstream or downstream from the marking site (indicated by shading). The smallest sampling area (500 m and the same as the marking area) could only detect 100% of fishes that moved less than approximately 50 m. Based on Albanese et al. (2003).
In theory, the probabilities of capture from distance weighting could be used to adjust the observed captures of fish (see Albanese et al. 2003). In actuality, outside of the zone of uniform sampling, the numbers of fish captured were so low that adjustments generally were not possible—adjustments are not possible if no fish are captured! The take-home message is that an understanding of distance weighting using PSd values is most useful for the a priori design of the sampling study. The a posteriori application of correction factors generally cannot correct for a poor study design.
In spite of the cautionary words by Gerking (1959) about the problem of experimental design and the presence of fish straying, and the suggestion by Funk (1955) about sedentary and mobile groups, the restricted movement paradigm became entrenched in the literature. One of the first to take issue with the restricted movement paradigm, specifically in regard to salmonids, was Gowan et al. (1994), who pointed out that although most tagging studies captured the majority of fish near the point of release, in 78% of the salmonid studies that they reviewed, over half of the fishes were never seen again after being marked. Whether these fish represented mortalities or fish that simply moved much greater distances is the crux to understanding the level of movement of fishes. Gowan et al. (1994) also suggested that the mobile fraction of fish populations had been downplayed through the use of such deprecating terms as “strays” and argued that more attention needed to be given to the experimental design of fish movement studies and to the underlying mechanisms involved in fish movement. The greater realization of the often high degrees of movement shown by freshwater fishes has had important consequences for the better understanding and management of fish populations (Fausch et al. 2002).
A study of fish movement in a small Ouachita Highlands stream (Arkansas) involved four species of stream fishes (Creek Chub, Semotilus atromaculatus; Blackspotted Topminnow, Fundulus olivaceus; Green Sunfish, Lepomis cyanellus; and Longear Sunfish, L. megalotis) (Smithson and Johnston 1999). The study area was 500 m long and consisted of 10 pools, and the possibility of movement of fishes outside of the study area was determined to be unlikely. Most fishes were recaptured in the same pool where they were initially marked; however, there were differences among the species. Compared to the other three species, Blackspotted Topminnow moved significantly greater distances. For all species, there were some individuals that moved greater distances than others, although there were no apparent morphological correlates associated with greater movement and, in fact, the same individuals switched between static and mobile behaviors over the course of the study. This suggests that individual fish periodically engage in exploratory travel, perhaps assessing habitat quality in areas outside of their home pool.
Field and theoretical approaches were used by Skalski and Gilliam (2000) to examine characteristics of movement of primarily four species (Bluehead Chub, Nocomis leptocephalus; Creek Chub; Redbreast Sunfish, Lepomis auritus; and Rosyside Dace, Clinostomus funduloides) in a small southeastern stream. In contrast to Smithson and Johnston (1999), species showed only weak differences in their degree of movement. However, within a species there was evidence for both “movers” and “stayers.” In addition, the distribution of distances moved tended to be leptokurtic, with more short and long movements and fewer intermediate movements compared to a normal distribution. Relationships between the propensity to move and morphological characteristics were complex and varied among species. Within Bluehead Chub, the probability of movement increased with size for individuals that had slow growth but decreased with size for those having fast growth. In Creek Chubs the probability of movement increased with body size but was not related to growth rate. There was no relationship between body size and the probability of movement in Redbreast Sunfish or Rosyside Dace. Correcting the movement data to account for distance weighting had little effect. However, this does not negate the strong potential effect of distance weighting but emphasizes the importance of an appropriate sampling design that, a priori, deals with the problem of distance weighting—a conclusion also reached by Albanese et al. (2003) (Box 5.2).
FIGURE 5.5. Patterns of migration in fishes. The circles enclose life-history stages using a particular resource. Solid lines indicate regular movement; dashed lines indicate aperiodic movement to refuge areas when there are harsh environmental conditions. Distances (A, B, C) between circles can vary greatly from a few meters to hundreds or thousands of kilometers. Adapted from Harden Jones (1968) and Schlosser (1995).
A general pattern that emerges from these studies of fish movement is that although the majority of fishes are often sedentary, there is often another, albeit smaller, group that undertakes much more extensive movements. Although use of the terms “movers” and “stayers” is descriptively appealing, several studies, including Smithson and Johnston (1999), have shown that the same individuals