How Artificial Are Artificial Substrata for Periphyton?

Author(s): Antonella Cattaneo and Marie Christine Amireault Reviewed work(s): Source: Journal of the North American Benthological Society, Vol. 11, No. 2 (Jun., 1992), pp. 244-256 Published by: Society for Freshwater Science Stable URL: http://www.jstor.org/stable/1467389 . Accessed: 06/03/2013 19:53
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J. N. Am. Benthol. Soc., 1992, 11(2):244-256 ? 1992 by The North American Benthological Society

How artificial are artificial substrata for periphyton?

ANTONELLA CATTANEO AND MARIE CHRISTINE AMIREAULT Departement de Sciences Biologiques,Universite de Montreal, C.P. 6126, succursaleA, Montreal, Quebec,Canada H3C 3J7 Abstract. Artificial substrataare increasingly used to study periphyton, but their ability to reproduce naturalsubstrataremains controversial.Although many studies have made contemporaneous comparisonsof periphyton assemblageson artificialand natural substrataat one or a few sites, no broadlybasedcomparisonexists.Wethereforesurveyedthe literatureto establishconditions under which artificialsubstratasatisfactorilymimic both the quantity and the quality of natural periphyton assemblages.In general, epilithon was underestimatedby the artificialsubstrata; epiphyton was overestimated,but less severely. These trends were significantly affectedby the time available for colonization of the artificialsubstratabefore sampling, site trophy, ambient temperature,and whether the study was conductedin a lake or in running water.Neither the composition of the substratumnor its orientationappearedimportant.Naturaldiatomassemblageswere usually well simulated by those on artificialsubstrata,whereas both epilithic and epiphytic green and Since artificialsubstrata bluegreen algae were severely underrepresentedon the artificialsubstrata. often misrepresentboth the quantity and the quality of naturalperiphyton, they should be used with more caution, especially in intersite and interseasonsurveys.
Key words: periphyton, artificial substrata, epiphyton, epilithon, stream, lake.

In streams and the littoral zones of lakes, algae grow on substrata that differ in origin, composition, size, and orientation. This heterogeneity and the associated variation in quantitative removal of algae from different substrata have always hindered quantitative study of the periphyton. Because of these problems, many researchers have elected to study periphyton assemblages that develop on artificial, introduced substrata, which simplify the natural complexity by providing uniform colonization time, material, texture, and size. They can be easily manipulated and they simplify both detachment of the assemblage and determination of the sample area. Since Hentschel (1916) first suspended glass slides in a lake, periphyton workers have experimented with many different materials and anchoring devices. These techniques have been reviewed various times (Sladeckova 1962, Wetzel 1965, Austin et al. 1981, Aloi 1990). Comparative studies of the assemblages on natural and artificial substrata are not lacking; yet, the question of whether artificial substrata satisfactorily mimic natural ones remains open, because the results of these comparisons are contradictory. This contradiction has two probable sources: authors evaluate their results differently in relation to the goals of their study, and the performance of substrata may depend on the environment studied and methods used.

The objective of this paper is to test the capacity of artificial substrata to substitute for natural ones in both quantitative and qualitative studies of periphyton. We apply consistent criteria to data in the literature to determine if performance varies as a function of 1) nature of the natural substratum (stone or plant); 2) environmental conditions (lake or stream, temperature, site trophy); 3) method (colonization time, position and orientation of substrata, material). By identifying general patterns with respect to these common factors, this exercise could suggest under which conditions and for what purposes artificial substrata are useful and, in contrast, when they could lead to seriously flawed conclusions. Methods To investigate the frequency with which periphyton investigators use artificial substrata, we systematically screened 16 years (1975-1990) of 20 journals where periphyton studies are often reported: Annales de Limnologie,Aquatic BotAustralianJournalof any, Archivfur Hydrobiologie, Marine and FreshwaterResearch,BritishPhycological Journal,CanadianJournalof Botany, Canadian Journal of Fisheries and Aquatic Sciences, Ecology, Freshwater Biology, Hydrobiologia, Internationale Revue der gesamten Hydrobiologie,Journal of the 244

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North American Benthological Society, Journal of Phycology, Limnology and Oceanography,Naturaliste Canadien,New Zealand Journalof Marine and FreshwaterResearch,Oecologia,Oikos, Transactions of the American Microscopical Society, and Verhandlungen der Internationalen Vereinigung fur Theoretischeund Angewandte Limnologie. In addition, our search for articles in which artificial and natural substrata were compared was extended through a computer search (Dialog Information Services, Inc., Palo Alto, California). We found 27 articles from which quantitative comparisons could be gleaned (Table 1); and when all the measurements taken at different dates, different sites, or with different techniques were extracted, these articles provided 343 points of comparison. The variables used to describe periphyton quantitatively were chlorophyll a, ash-free dry mass, biovolume, cell density, organic carbon, and production. When more than one variable was used in the same study, we used the data of only one to avoid overemphasizing these results in the analysis. For each comparison, we calculated the ratio of the value measured on the natural substratum to that on the artificial one. In addition, 34 articles allowed 245 comparisons of assemblage composition (Table 2). These taxonomic comparisons usually expressed the data as percentage composition. To judge substratum performance in this context, we used an arbitrary binary choice, whereby "good agreement" (as opposed to "poor agreement") was scored when the percentage of the dominant taxonomic groups differed less than twofold. If the SIMI similarity index, whose value ranges between 0 and 1, was used instead of the percentage composition, we took a "good performance" to be one where the index had a value greater than 0.5 (Sullivan 1975). In a few instances, authors stated clearly that there was marked difference in dominant taxa between substrata, and we accepted that judgement as indicating "poor performance". For each quantitative or taxonomic comparison, we recorded the natural substratum (plant or rock), colonization time, the position, orientation, and composition of the artificial substratum, and environmental characteristics, when available. When date, but not temperature, was provided, we estimated temperature from published values for sites with comparable climate (Wernstedt 1972). The trophic status

TABLE1. Articles containing quantitative comparisons between periphyton assemblages on natural and artificial substrata. Also listed are the number of comparisons in each article (No.) and the variable used in the comparisons: organic carbon (Carbon), chlorophyll a (Chl a), algal density (Density), primary production (Production), and ash-free dry weight (AFDW).

Reference Allen 1971 Biggs 1988b Bowker and Denny 1978 Burkholder and Wetzel 1990 Capblancq 1973 Cattaneo 1990 Cattaneo and Kalff 1978 Cattaneo and Kalff 1979 Coleman and Dahm 1990 Corning et al. 1989 Fairchild and Everett 1988 Foerster and Schlichting 1965 Fontaine and Nigh 1983 Gale et al. 1979 Lamberti and Resh 1985 Lay and Ward 1987 Leland and Carter 1985 Liaw and MacCrimmon 1978 Loeb 1981 Lowe and Gale 1980 Mason and Bryant 1975 McConnell and Sigler 1959 Millie and Lowe 1983 Murphy 1984 Rosemarin and Gelin 1978 Shamess et al. 1985 Stock and Ward 1989 Tuchman and Stevenson 1980

No. 2 136 11 11

Variable Production Chl a Chl a Chl a Chl a Chl a Chl a Chl a Chl a Chl a Chl a Density Chl a AFDW Chl a Chl a Carbon Chl a Carbon Density

1
12 8 19 2 5 1 4 33 12 8 2 11

10
2 35 10 4 7 1 54 32 6 2

AFDW Chl a Density

AFDW Chl a Density Production Density

of most sites could be estimated only roughly. Orthophosphate was the most commonly available indicator of trophy. Unfortunately the meaning of this parameter is uncertain (Rigler 1966) especially in rivers (Peters 1981). We therefore classified the environments only into two broad categories, giving a value of 0 ("clean") to waters where P-PO4 < 10 ,ag/L or when the water was clearly described as draining a pristine mountain basin, and a value of 1 ("rich") to waters with P-PO4 > 10 ,g/L or draining farmland or densely populated areas. Simple and multiple regressions were used to assess if the quantitative performance of the artificial substrata, measured as the natural:

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TABLE2. References that allowed comparisons of the taxonomic composition of periphyton assemblages growing on natural and artificial substrata and number (No.) of such comparisons that could be gleaned from each article.

pared visually by box-and-whisker plots (Tukey

1977) and by analysis types after correction of variance to detect diffor the effects of coloni-

ferences in performance among substratum zation time, temperature, and trophy. Results Basic patterns in the use of artificialsubstrata The number of papers dealing with periphyton has increased steadily over the last 15
yr (Fig. 1A). In the mid-1970s, artificial substrata

Reference Backhaus 1967 Besch et al. 1972 Biggs 1988a Bowker and Denny 1978 Burkholder and Wetzel 1989 Capblancq 1973 Castenholz 1960 Cattaneo et al. 1975 Cattaneo and Kalff 1978 Corning et al. 1989 Elber and Schanz 1990 Eloranta 1982 Evans and Stockner 1972 Fairchild and Everett 1988 Foerster and Schlichting 1965 Fontaine and Nigh 1983 Goldborough and Robinson 1985 Hoagland and Peterson 1990 Lay and Ward 1987 Leland and Carter 1984 Loeb 1981 Lowe and Gale 1980 Mason and Bryant 1975 Millie and Lowe 1983 Shamess et al. 1985 Shortreed et al. 1984 Shortreed and Stockner 1983 Siver 1977 Stock and Ward 1989 Stockner and Armstrong 1971 Tippet 1970 Tuchman and Blinn 1979 Tuchman and Stevenson 1980 Young 1945

No. 6 2 3 2 3 1 4 9 7 5 2 2 1 1 1 33 3 1 2 1 2 28 1 39 32 4 5 7 6 1 4 24 2 3

were used in -40% of these publications, but more recently, they appear in ~60% of periphyton studies (Fig. 1B). The popularity cial substrata is clearly on the rise. use of different substrata. of artifi-

Figure 2 summarizes temporal changes in the

Traditionally, glass

60
X0
a

(A) 50 -

40 -

CL

20

aa ~year

80

X0 40 0

(B)

artificial substratum ratio, was related to environment and method. For these analyses, natural:artificial substratum ratio, time of colonization, and temperature were logarithmically

transformed to stabilize variance and linearize response. Habitat (lake vs. stream), trophy (clean vs. rich), and orientation (horizontal vs. vertical) were introduced as "dummy" variables (Gujarati 1978). Residual values (observed minus predicted values) of these multiple regres-

en o

1975

1980

1985

1990

year
FIG. 1. Number of periphyton papers published from 1975-1990 in selected journals (A), and percentage of these papers in which artificial substrata were used (B).

sions for substrata of different composition (glass, plastic, tile, introduced rock) were com-

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ARTIFICIAL SUBSTRATA FOR PERIPHYTON

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1975- 1979

glass (25.0%)

stones (11.4%)

(43.2?/)

1980- 1984
others(7.4%) nutr.diff.(2.9%) 3X% tiles (10.3%).

glass (26.5%)

Resh 1983) and nutrients (e.g., Fairchild and Lowe 1984). In most studies, substrata are left in the water for 2-4 wk before sampling (Fig. 3). This colonization time is logistically convenient and it avoids long exposures and the chance of loss to vandalism or spates. Moreover, such a time interval is suggested as ideal because it allows the periphyton to develop its maximum biomass, but avoids subsequent sloughing (APHA 1989). The choice of position and orientation of substrata seems to depend partly on habitat type. In streams, substrata are more often positioned horizontally (60%)on the stream bed (74%)rather than suspended vertically. In lakes, substrata are positioned either on the bottom (34 studies) or suspended in the water column (33 studies), but a vertical orientation seems preferred (79% of studies). Quantitative comparisonsof biomass The ratios of the biomass observed on stones (epilithon) to that on artificial substrata tend to be > 1 (median 2.35). The values on natural substrata are more than twice those on artificial substrata in 68% of the comparisons (Fig. 4A). Since periphyton estimates are highly variable (Weitzel et al. 1979, Morin and Cattaneo 1992),

stones (20.6%)' plastic(32.4%)

1985- 1990
s (9.6%), others (21.7%)

nutr.diff.(13.9%)

tiles (20.9%)

plastic(22.6%)
u,
:3
U4

stones (11.3%) 2. Percentage of different materialsused in periphyton studies published in 20 journals.

FIG.

')

o
Q0

E
slides were most common (Sladeckova 1962). Slides are still widely used, but other materials have also become popular, and plastics in many different forms (plexiglass, polyethylene sheets and tapes, plastic aquarium plants, etc.) are now the favored substratum. Precleaned river or lake stones or slabs cut from rocks have also been used frequently in the last 10 yr. The increased use of tile and nutrient-diffusing pottery is associated with the use of these materials in experimental studies of grazers (e.g., Lamberti and z

< 8 8-14

15-30

61-120 31-60 > 120

Days of colonization
FIG.3. Time of colonization of artificialsubstrata used in periphyton studies published between 1975 and 1990 in 20 journals.

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(A)
20

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These devices underestimate epilithon quantity more severely when colonization time is less than 60 d (Fig. 5). On the other hand, the agree15 ment between values measured on artificial u, C (t) 0 substrata and on plants is instead the closest 0 a when short colonization times are considered 10 a U, (Fig. 6). The tendency of artificial substrata to 0 overestimate the biomass on plants increases with colonization time so that the most severe overestimation is found on substrata colonized longer than 120 d (Fig. 6). I.L. To test if other factors besides colonization 64 >128 1 4 16 <0.008 0.016 0.06 0.25 c time affected the performance of artificial subsubstratum Stone/ Artificial (1) strata, we used a multiple regression in which temperature, trophy, habitat, and substratum orientation were added to predict the ratio of biomass on natural to that on artificial substrata. (B) The effect of substratum position could not be tested since all epilithon comparisons, except one, involved artificial substrata on the bottom, whereas artificial substrata compared with plants 0 were always suspended. The ratio of periphyton on stones to that on n 0 comparable artificial substrata decreases with the time of colonization, trophy, and temperature. There is a significant difference between streams and lakes. Since the average ratio in o these comparisons exceeds 1, indicating that 4 16 64 >128 1 <0.008 0.016 0.06 0.25 on natural substrata is higher than on biomass substratum Plant / Artificial artificial ones, this regression suggests im4. Frequencydistributionof the ratio of the FIG. proved performance at warmer, richer sites, in periphyticbiomass measuredon naturalsubstratato that measuredon nearby artificialsubstratain com- streams, and after long periods of colonization, because the ratio declines towards unity in these parisonsinvolving assemblageson stones (A) and on situations. Orientation (vertical or horizontal) (B). plants does not affect the ratio significantly (Table 3). The low explained variation may reflect the large two-fold differences are not always significant. However, even if only four-fold differences are variability of the data, the crude estimations of considered robust, disagreement between as- temperature and trophy, and the importance of semblages on stones and artificial substrata is other untested variables. The ratio of periphyton on plants to that on still observed 40% of the time. In epiphytic assemblages, the ratios of the artificial substrata also declines with time of values on natural and artificial substrata tend colonization, trophy, and temperature (Table 3). Since in this case the median ratio is less than to be <1 (median 0.77). Two-fold differences are observed in 46% of the comparisons and 1, the regression suggests that the tendency of artificial substrata to overestimate biomass on four-fold differences only in 25%(Fig. 4B). Since the distribution of the ratio of the biomass on plants would increase with time, trophy, and natural to that on artificial substrata is signifitemperature. Neither the effects of habitat nor those of orientation could be assessed in the different for and for cantly epilithon epiphyton < test, p epiphyton data set, because only one study was (Kolmogorov-Smirnov two-sample 0.001), the two assemblages will be treated sepperformed in a river and because all comparisons involved vertically oriented artificial subarately in the following analyses. strata. The time of colonization is important in regartificial substrata. of the By examining the residuals around equations performance ulating
> Stone Artificial Stone > Artificial

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ARTIFICIAL SUBSTRATA FOR PERIPHYTON

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61 - 120 days

< 15 days
1o-

Art. > Stone

Stone > Art.

U) c

8-

cn

6-

n 0

() .O 4-

2*

CD 01 U)

U)

0 O (D U) .0 0

O)

<0.008

0.016

0.06 0.25

16

64

>128

Stone / Artificial

c0

U)

o
n O

<0.008 0.016

0.06

0.25

16

64

>128

Stone / Artificial
FIG. 5. Frequency distribution of the ratio of periphytic biomass measured on natural stones to that measuredon nearby artificialsubstrataobserved when the latter were colonized for different times before sampling.

1 and 2 (Table 3) we could test if, after correction for the other variables, the composition of artificial substrata affects their performance. Visual inspection of box-and-whisker plots showing the median, the 25- and 75-percentile values, and the range of the residuals for each substratum type (Fig. 7) suggested similar residuals.

Analysis of variance of these residuals confirmed that the four types of material (glass, plastic, tile, introduced stone) considered for the comparisons with epilithon were not significantly different (p = 0.21). However, comparisons with epiphyton revealed a slight but significant difference (p = 0.05) between glass

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250

A. CATTANEOAND M. C. AMIREAULT < 8 days

8

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11

Art. > Plant

Plant > Art.

C, 0 16 cn

(C)

0 ()
C,)

6-

4-

.0 2

I n 0

U1)

C,)

c, c 0 c o) U1)

n 0

16 - 30 days
10-

U) 8-

0 o 6-

CO 0

c.

u) CO

._

4-

2-

0 <0.008

0.016

0.06

0.25

16

64

>128

<0.008

0.016

0.06

0.25

16

64

>128

Plant / Artificial

Plant/ Artificial

FIG.6. Frequency distribution of the ratio of the periphytic biomass measured on plants to that measured on artificial substrata observed when the latter were colonized for different times before sampling.

and plastic, the only two materials that could be tested for this subset. Overestimation of biomass on plants tended to be less severe for plastic than for glass substrata. Taxonomiccomparisons Good agreement in the total taxonomic composition of the assemblages was seen in 82% of

the comparisons between natural epilithon and artificial substrata. This agreement is repeated in comparisons of the diatoms, which are often the dominant group in periphyton and sometimes the only one analysed by researchers. Green and bluegreen algae tend instead to be underestimated by artificial substrata: the distribution of these taxa was adequately reproduced in less than half of the comparisons (47%

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TABLE 3. Multiple regression analyses of the ratios of periphyton biomass measured on stones (B) and artificialsubstrata(Ba) or on plants (Bp)and artificialsubstrata.The independent variables tested were: colonization time (LogTIME; ?C),trophy (TRO; poor = 0; rich = 1), habitat(HAB; days), temperature(Log TEMP; lake = 0; stream = 1), and substratumorientation (ORI;horizontal = 0; vertical = 1). Neither ORI nor HAB could be tested in Equation2. Partialcoefficientsof determination(R2),and probabilitylevels of each variable are listed. Both equations are highly significant (p < 0.0001). The adjusted R2, the F value, and the sample

size (n) are presented for each equation. Epilithon (equation 1)

Log B/Ba = 2.85 partial R2

Epiphyton (equation 2)
Log Bp/Ba= 2.29 partial R2

-0.56 Log TIME - 1.04 Log TEMP - 0.24 TRO -0.30 HAB
- 0.04 ORI Adjusted R2= 0.44

0.16*** 0.16*** 0.07** 0.05***

- NS (F = 34,

- 0.60 Log TIME - 1.50 Log TEMP - 0.59 TRO

Adjusted R2 = 0.19

0.11** 0.03* 0.05**

(F = 8,

n = 167)

n =87)

NS not significant, * p < 0.05, ** p < 0.01, *** p < 0.001.

for the greens and 38%for the bluegreens; Table 4). The proportion of successful representations of greens and bluegreens on artificial substrata improves somewhat after 60 d of colonization (67% and 40%, respectively). Similar patterns are observed when the taxonomic composition of assemblages on plants and on artificial substrata are compared. However, the performance of artificial substrata is slightly worse in these comparisons than in those involving the epilithon (Table 4). While the composition of the whole assemblages and of the diatoms are adequately reproduced (agreement 61% and 60% of the time respectively), misrepresentation of green and bluegreen algae becomes particularly severe (agreement 38%and 24% of the time, respectively). In contrast to the comparisons with epilithon, this misrepresentation is even larger when only colonization times longer than 60 d are considered (agreement 28% of the time for greens and 11% for bluegreens).

Discussion Stones, especially in lakes, are stable substrata on which periphyton can accumulate over long periods, even years. It takes time for an introduced substratum to reach the same extent of colonization and this time will depend on the productivity of the site. Factors that accelerate growth would tend to decrease the colonization time necessary to reach a biomass as high as

that on the stones. The regression identified three such factors: temperature, trophy, and current. Most studies involve short colonization times that would be most appropriate in rich and warm environments. Typical colonization times of 2-4 wk (Fig. 3) appear to give differences larger than two-fold in 70% of the cases and larger than four-fold in 42% of the measurements. Longer colonization times should therefore be used, but these longer experiments are often impractical because they may require placement of the substrata in the previous season, and because longer exposures increase the chances of losing the substrata to either vandalism or spates. Because temperature and trophy affect the ability of artificial substrata to reproduce the standing crop on natural stones, surveys involving different sites and seasons using a same short colonization time could yield seriously biased results. Authors comparing substrata colonized for different times along seasonal (Low and Gale 1980) or trophic gradients (Biggs 1988b) have already warned about this problem, with apparently little effect because constant, short colonization times are still recommended in intersite comparisons (APHA 1989). Unlike stones, plants provide an ephemeral substratum. Artificial substrata mimicking plants are usually introduced at the onset of the seasonal growth of the macrophyte, so that their colonization time is similar to that of the natural substratum. This synchrony probably explains why assemblages on plants and on artificial sub-

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Epilithon- equation 1
Glass Plastic Tile Introduced stone

-Ii
!

I I I I
I
I
. . . I? .

n =64
n=38

n=8
n =57
I

-0.4

0.0

0.4

0.8

1.2

1.6

Residual

Epiphyton- equation 2
Glass Plastic

II
1
-1.4

n= 15 ?. n = 72

1 I 1
-0.7

I
1 1.4

0.0 0.7 Residual

2.1

FIG. 7. Box-and-whisker plots of the residualsof equation 1 (top panel) and equation 2 (lower panel) from Table 3, observed for the different materials.The median value for each substratumtype is markedby the
central vertical line; 25- and 75-percentile values form the ends of the box; whiskers delimit the range of the observations except for extreme values defined as those that lie beyond 1.5 x the box length. These extremes are represented by points (Tukey 1977).

strata generally agree better than those on stones. However, as the time of colonization and, concomitantly, the age of the plant increases, artificial substrata increasingly overestimate epiphyte biomass (Fig. 6). This overestimation could be caused by a number of mechanisms. It may partly reflect the negative effect on epiphyte biomass of allelopathic compounds released by the plant (Burkholder and Wetzel 1990). Alternately, grazers, particularly leaf-miners, could prefer the natural substratum and deplete epiphytes there. For example, higher chironomid densities have been observed on Typhastems than on nearby glass rods (Mason and Bryant 1975). Finally, the surface of an aging plant can also differ increasingly from an artificial surface in hard water lakes and streams because precipitated carbonate can influence the algal assemblages (Cattaneo and Kalff 1978). Taxonomic differences, at least for

green and bluegreen algae also tend to increase with time (Table 4). Equation 2 suggests that the overestimation of epiphyte biomass by artificial substrata should increase in eutrophic sites and at high temperature, i.e., in more productive situations. This is contrary to the hypothesis of Eminson and Moss (1980) that host type would have its greatest influence in unfertile lakes. We found no significant effect of substratum orientation. However, in the literature, when the two orientations are measured together, horizontal substrata generally yield higher biomass in lakes (Newcombe 1950, Castenholz 1960, Cattaneo 1990) while in rivers the amount is similar or slightly higher on the vertical ones (King and Ball 1966, Liaw and MacCrimmon 1978, Meier et al. 1983). Our analyses suggests that choice of material is not crucial. Preferential colonization of a sub-

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4. Percentageof observationswhere the taxonomiccompositionof assemblageson stones or plants TABLE agreed well with those on nearby artificialsubstrataafter different colonization times (time). Comparisons were based on the whole assemblage and on diatoms, green algae, and bluegreen algae. The number of comparisonson which the percentageswere calculatedis listed in parentheses. Time (days) <15 16-30 31-60 >60 Total <15 16-30 31-60 >60 Total Whole %(No.) 0 (7) 91(43) 83 (18) 94 (16) 82(84) 36 (36) 67 (46) 59 (22) 75 (53) 62(157) Diatoms %(No.) 14 (7) 88(42) 82(17) 92 (13) 81(79) 57 (21) 69 (36) 42(19) 64(47) 61(123) Greens %(No.) 29 (7) 44(16) 67 (6) 67 (3) 47 (32) 25 (4) 78 (9) 18 (11) 33(18) 38 (42) Bluegreens %(No.) 0 (7) 47 (15) 57 (7) 40 (5) 38 (34) 0 (15) 68 (19) 20 (5) 7 (14) 28 (53)

Stones vs. artificialsubstrata

Plants vs. artificialsubstrata

stratum of a particular material, texture, or color has sometimes been reported (Nielsen et al. 1984, Rosemarin and Gelin 1978), but these results are often inconsistent (Lowe and Gale 1980, Shamess et al. 1985). Our tests, based on a very variable data set, are not very powerful so that significant effects of composition and orientation may be undetectable in such a general comparison. Nevertheless, our results indicate that these variations are less important than those introduced by length or colonization, trophy, and temperature. These factors, rather than the type of substratum, should be considered first in experimental design. In many studies, the quality of the assemblage is more important than the quantity. Artificial substrata reproduce both epilithic and epiphytic diatom assemblages well. Close resemblance was observed, for example, in the classical work of Castenholz (1960) and is the basis of the diatometer (Patrick et al. 1954) widely used for monitoring river pollution. On the other hand, green algae and especially bluegreens are underestimated on many artificial substrata. This difference could be related to the slower growth of the latter forms, for diatoms tend to be the first colonizers (Stock and Ward 1989). The consequences of such substratum effects could be serious. In grazing experiments using artificial substrata, the invertebrates could encounter an algal assemblage different in taxonomy and possibly in palatability and size

when compared with the natural prey community. In surveys of the effects of pollution or acidification, artificial substrata could underestimate the growth of the nuisance green filamentous algae usually associated with these perturbations. In many cases artificial substrata are too artificial, as Wetzel warned in 1983. Nevertheless, artificial substrata have been increasingly used, likely reflecting the popularity of grazer and nutrient manipulations often carried out in artificial streams or limnocorrals. The easy handling of such substrata makes them invaluable in these studies and in those where such processes as immigration, colonization, and growth rates are measured (e.g., Stevenson 1984). By providing a uniform substratum for periphytic growth, artificial substrata are often intended to increase the precision of the measurements (Tuchmann and Stevenson 1980, Meier et al. 1983) but this claim is still controversial (Weitzel et al. 1979, Lowe and Gale 1980, Morin and Cattaneo 1992). Besides, the uniformity of artificial substrata may obscure the ecological role of the variety of substratum sizes and textures that typify different sites (Tett et al. 1978). Artificial substrata are used more often than their performance warrants. Their possible distortion of natural periphyton assemblages should be critically evaluated in terms of the conditions and goals of each experiment. More frequent use of natural substrata may involve

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some methodological problems but this effort might be rewarded by more realistic answers to problems in rivers and lakes. Acknowledgements This work was funded by an operating grant from the Natural Science and Engineering Research Council of Canada to AC and a team grant from FCAR (Formation de Chercheurs et l'Aide a la Recherche) to the Equipe des Eaux douces. MCA was funded through the Challenge Program for student summer employment of the Canada Ministry of State for Youth. Robert Peters provided encouragement and advice. The comments of Rosemary Mackay, Gary Lamberti, and an anonymous reviewer helped in the revision of a previous version. This paper is contribution No. 383 of the Groupe d'Ecologie des Eaux douces de l'Universite de Montreal.

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Received: 12 September 1991 Accepted: 13 February 1992

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