The effectiveness of quadrats for measuring vascular plant

diversity

Abstract.--Quadrats are widely used for measuring characteristics of vascular plant communities. It is well
recognized that quadrat size affects measurements of frequency and cover. The ability of quadrats of varying sizes to
adequately measure diversity has not been established. An exhaustive search of sixteen 40 by 40 m old field plots
documented presence or absence of 88 vascular plant species (mean = 58.8 per plot). Against these benchmarks,
measurements of species richness were obtained from 50 samples in each plot and compared among three different
sized quadrats (0.2 by 0.5 m, 0.5 by 0.5 m and 0.5 by 1.0 in). These quadrats differed in the number of species
measured, with an average of 33.6, 38.7 and 41.4 species respectively (P<0.001; Least Significant Difference test
critical difference = 0.7 species). Quadrats are relatively ineffective for documenting the actual number of vascular
plant species present on old field plots (at least 30% of species undetected). However, under the intensity of
sampling used in this study (n=50) a difference of less than one species per plot was detected at P <0.05. Thus,
quadrat sampling regardless of quadrat size can provide a precise index of vascular plant species richness.

Recent controlled field studies provide evidence suggesting that plant diversity contributes to ecosystem resilience,
and that this diversity is related to global nitrogen and carbon cycling (Berendse 1990; Inouye & Tilman 1988;
Tilman 1987; 1988; 1996; Wedin & Tilman 1996). Investigation of these effects in uncontrolled field conditions will
require an excellent understanding of the capabilities of field sampling techniques to reliably measure diversity.
Thus, knowledge of technique effectiveness is not only important for research planning (e.g., Cook & Stubbendieck
1986), but more importantly is essential regarding diversity data interpretation and management recommendations.

Numerous studies have investigated vegetation measurement techniques (e.g., Canfield 1941; Cook & Stubbendieck
1986; Daubenmire 1968; Etchberger & Krausman 1997; Higgins et al. 1996; Stohlgren et al. 1998). Regarding
quadrat sampling, Bormann (1953) demonstrated that many small quadrats achieve greater precision for a given
sample size than larger quadrats sampling an equal area. Generally, long thin quadrats are considered superior
because they cross more vegetation patches (Daubenmire 1968; Krebs 1989). However, these guidelines refer to
measurement of cover and frequency. The relationship of these guidelines to measurement of diversity is not
immediately clear.

Richness estimates were derived and compared among three different sized quadrats (0.2 by 0.5, 0.5 by 0.5 and 0.5
by 1.0 m) in sixteen 40 by 40 m plots within an old field in south-central Oklahoma during June and July 1998. The
influence of quadrat size on detection frequency for 88 plant species that varied markedly in abundance was
investigated.

METHODS

Each of 16 study plots measured 40 by 40 m. Vegetation was sampled using three different sized quadrats (0.2 by
0.5 m [Daubenmire], 0.5 by 0.5 m and 0.5 by 1.0 in). Additionally, plots were exhaustively searched with the
purpose of cataloging presence and absence of all vascular plant species. Each quadrat size was sampled at 50
positions within each plot. Positions within plots were determined by stepping-off a random number of steps
between positions (between 1 and 10), along parallel transects that were randomly spaced. At each position, the
observer dropped the quadrat without looking at the ground. Known richness (determined from exhaustive search)
was compared to richness determined by each sized quadrat. Additionally, frequency of detection for each plant
species was compared among the three different sized quadrats. Analyses included one-way analysis of variance for
randomized blocks with SPSS (SPSS Inc. 1997).

RESULTS

Quadrats underestimated richness by 30-43% (P<0.001) (Figure 1). With few exceptions (17 of 71 species), quadrat
size did not affect frequency estimates for species that occurred in fewer than 10% of quadrats, whereas (except for
Buchloe dactyloides and Aristida oligantha) quadrat size did affect frequency estimates for all species that occurred
in greater than 10% of quadrats (n =17) (Figure 2).

DISCUSSION AND CONCLUSIONS

At first glance, it is reasonable to conclude that a failure to detect 30-43% of species present indicates that quadrats
up to 0.5 [m.sup.2] are unable to effectively document vascular plant richness in old fields. This reasoning is further
supported by the recognition that comparative samples were taken from 40 by 40 m old field plots that are not
considered to be highly diverse. However, the experimental design was able to detect a difference of less than 1
species (0.7) among sampling methods (LSD test). This is the smallest biologically meaningful difference possible.
Therefore, quadrat sampling when conducted with sufficient intensity (n =50 samples per 40 by 40 m plot in this
study) can provide a reliable index of species richness.

It is not surprising that larger quadrats detected more species than did smaller quadrats. It is more important to note
that quadrat size has a limited effect on detection frequency for species that occur in fewer than 10% of quadrats
(Figure 2), and that measurements of 54 of 88 species were unaffected by quadrat size. Thus, whereas quadrats have
significant limitations regarding measurement of individual species (Stohigren et al. 1998; this study), these data
support the use of quadrats for providing comparative estimates of vascular plant diversity (richness).

ACKNOWLEDGMENTS

We thank N. Watts and J. Williams (United States Environmental Protection Agency [EPA]) and D. Engle
(Oklahoma State University) for their support and advice. Although the research described in this article has been
funded wholly by EPA it has not been subjected to Agency review. Therefore, it does not necessarily reflect the
views of the Agency.

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Cook, C. W. & J. Stubbendieck. 1986. Range research: basic problems and techniques. Society for Range
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Daubenmire, R. 1968. Plant communities; a textbook of plant synecology. Harper & Row, Publisher, New York,
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