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Modified Whittaker Plots as an Assessment and


Monitoring Tool for Vegetation in a Lowland
Tropical Rainforest

Article in Environmental Monitoring and Assessment May 2002


DOI: 10.1023/A:1015264720284 Source: PubMed

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MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND
MONITORING TOOL FOR VEGETATION IN A
LOWLAND TROPICAL RAINFOREST

PATRICK CAMPBELL1 , JAMES COMISKEY1 , ALFONSO ALONSO1 ,


FRANCISCO DALLMEIER1, PERCY NUEZ2 , HAMILTON BELTRAN3 ,
SEVERO BALDEON3 , WILLIAM NAURAY2 , RAFAEL DE LA COLINA2 ,
LUCERO ACURIO2 and SHANA UDVARDY1
1 Smithsonian Institution Monitoring and Assessment of Biodiversity Program, Conservational
Research Center, National Zoological Park, 1100 Jefferson Drive, SW, Suite 3123, Washington, DC,
20560-0705, U.S.A.; 2 Departmento de Botanica, Facultad de Ciencias Biologicas, Universidad San
Antonio Abad del Cusco, Cusco, Peru; 3 Museo de Historia Natural, Universidad Nacional Mayor
de San Marcos, Avenida Arenales 1256, Jesus Maria, Lima, Apartado 140434, Lima, 14 Peru

Abstract. Resource exploitation in lowland tropical forests is increasing and causing loss of biod-
iversity. Effective evaluation and management of the impacts of development on tropical forests
requires appropriate assessment and monitoring tools. We propose the use of 0.1-ha multi-scale,
modified Whittaker plots (MWPs) to assess and monitor vegetation in lowland tropical rainforests.
We established MWPs at 4 sites to: (1) describe and compare composition and structure of the sites
using MWPs, (2) compare these results to those of 1-ha permanent vegetation plots (BDPs), and
(3) evaluate the ability of MWPs to detect changes in populations (statistical power). We recorded
more than 400 species at each site. Species composition among the sites was distinctive, while mean
abundance and basal area was similar. Comparisons between MWPs and BDPs show that they record
similar species composition and abundance and that both perform equally well at detecting rare
species. However, MWPs tend to record more species, and power analysis studies show that MWPs
were more effective at detecting changes in the mean number of species of trees 10 cm in diameter
at breast height (dbh) and in herbaceous plants. Ten MWPs were sufficient to detect a change of 11%
in the mean number of herb species, and they were able to detect a 14% change in the mean number
of species of trees 10 cm dbh. The value of MWPs for assessment and monitoring is discussed,
along with recommendations for improving the sampling design to increase power.

Keywords: assessment, modified Whittaker plots, monitoring, tropical forests

1. Introduction

Resource exploitation by ever increasing human populations is leading to the


highest rates of loss in Earths biodiversity ever recorded (Whitmore, 1997). Threats
to biodiversity stemming from exploitation of tropical forests are particularly dis-
tressing. Tropical lowland forests, which harbor most of the worlds biodiversity
(Lovejoy, 1997), are being cleared at alarming rates. In South America, for ex-
ample, tropical forest cover declined by 23 277 000 hectares (ha) a loss of 2.7%

Environmental Monitoring and Assessment 76: 1941, 2002.


2002 Kluwer Academic Publishers. Printed in the Netherlands.
20 P. CAMPBELL ET AL.

from 1990 through 1995 (FAO, 1997). This decline in forest cover is inevitably
leading to loss of wildlife habitat and, ultimately, loss of biodiversity (Myers,
1986). The need for sound scientific data regarding the distribution, abundance
and dynamics of populations of organisms has never been greater.
Assessment and monitoring programs provide a means to obtain this data, and
adaptive management principles provide the framework within which managers
can make appropriate decisions regarding the use of forest resources (Spellerberg,
1992; Dallmeier and Comiskey, 1998). Under adaptive management principles,
clear research objectives are established. Assessment and monitoring are the means
to evaluate and meet the objectives. The assessment provides baseline data relev-
ant to the site of interest and should consist of field surveys, inventories (includ-
ing identification and classification of species) and literature reviews (Spellerberg,
1991; Dallmeier and Comiskey, 1998). Monitoring is the repeated measuring and
sampling of species over time and comparing the results to the baseline, noting
any deviation from an expected norm (Hellawell, 1991). Thus, assessment and
monitoring act as early warning tools to determine forest conditions, formulate
additional research hypotheses and, most importantly, measure progress toward or
success at meeting objectives. In the end, the evidence exists to adapt management
strategies and actions, continue them or cease altogether (Holling, 1978; Dallmeier
and Comiskey, 1998; Elzinga et al., 1998).
Biologists use a variety of methodologies to assess vegetation, and they have
made attempts to standardize techniques at national and global scales (Dallmeier
and Comiskey, 1996; Ashton, 1998). This allows cross-site comparisons to increase
our understanding of taxonomic distributions (Terborgh and Andresen, 1998) and
trends (Phillips and Miller, in press). Common methods may include simple col-
lections, which provide a checklist of species present, or more systematic methods
such as plots or transects, which provide quantitative information and thus are
valuable components of both assessments and monitoring.
Transects have been used extensively to assess vegetation. They are often es-
tablished along obvious environmental gradients, the point being to describe max-
imum variation over the shortest distance in a minimum amount of time (Kent
and Coker, 1992). Gentry (1982, 1988a, 1995) developed a transect method that
he used to characterize the diversity of vegetation throughout the tropics. Gentrys
approach allows for the collection of quantitative floristic data from a wide range
of sites using consistent protocols, which enables comparisons of tropical forest
biodiversity along latitudinal and rainfall gradients (Gentry, 1982, 1995; Clinebell
et al., 1995). A disadvantage is that most transects miss important patches (or
hotspots) of diversity (Stohlgren et al., 1997) and are not permanent which prevents
the collection of temporal data.
Permanent plots, which allow researchers to follow a population of marked
trees over time, offer the greatest potential as a long-term monitoring tool. A
common method is the 1-hectare (ha) biodiversity plot (BDP; Dallmeier, 1992)
used by the Smithsonian Institutions Monitoring and Assessment of Biodiversity
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 21

Program (SI/MAB). Initially, a BDP provides an assessment of the species present


in a particular habitat type, including an inventory of tree species and a detailed
description of the structure of the forest (Dallmeier and Alonso, 1997; Alonso and
Dallmeier, 1998; Comiskey et al. 2001). These plots are recensused periodically,
lending a dynamic aspect to the survey. By tracking the history of known indi-
viduals, managers can observe changes in the forests structure and composition
through measurement of mortality, recruitment, changes in relative abundance and
growth rates. They can also investigate species replacement patterns and ultimately
make predictions about future forest composition (Hubbell and Foster, 1987).
Biologists have also established a worldwide network of permanent 50 ha plots
(Condit, 1995; Ashton, 1998). Most species are rare in tropical forests (Hubbell and
Foster, 1986), and to obtain valuable information on the dynamics of a wide range
of species, it is necessary to sample larger areas and size classes (Clark and Clark,
1992). Such large sample sizes provide a unique experimental site for examining
theories on the maintenance of species diversity (Hubbell, 1998) and monitoring
community change at the species level in response to climatic fluctuations (Con-
dit et al., 1995). However, these large plots can be labor intensive and costly to
maintain (Hubbell and Foster, 1992).
BDPs and the 50 ha plots are typically established as single sampling units
because of time and cost constraints. The lack of replication means that they fail
to detect spatial variability over larger scales (Condit, 1995), and this makes it
difficult to draw conclusions about the changes in an entire region. Furthermore,
the plots fail to indicate whether what was measured is typical of the surrounding
habitat (Stohlgren et al., 1997).
Modified Whittaker Plots (MWPs) were developed by Stohlgren et al. (1995),
based on the original nested vegetation sampling method developed by R. H. Whit-
taker and described by Shmida (1984). These small, 0.1 ha, multi-scale plots allow
investigators to examine the temporal and spatial aspects of vegetation at a perman-
ently marked site. Shmida (1984) used Whittaker plots to record species richness
data at multiple spatial scales to investigate species accumulation with increasing
area. Stohlgren et al. (1995) modified this design to reduce the autocorrelation of
the nested plots that was inherent in the original design and found that the modified
design was more effective at estimating species/area relationships compared to the
older design. A series of permanent MWPs at an assessment and monitoring site
may provide the temporal and spatial data necessary for an effective monitoring
program.
For any monitoring program to be successful, the methods used must be care-
fully evaluated. Since an objective is to detect changes in population parameters,
we need to test the effectiveness of potential sampling designs and efforts to detect
such change. For this reason, Type II statistical errors are of concern to those
involved in monitoring biodiversity. A Type II error is when a researcher fails
to reject the null hypothesis when, in fact, it is false or when a researcher fails
to detect a change when, in fact, it did occur. Statistical power analysis (Cohen,
22 P. CAMPBELL ET AL.

1977) provides a means to investigate the probability of correctly rejecting a null


hypothesis that is false (Steidl et al., 1997). During the early phases of a monitoring
program, power analysis can provide insight into the effort necessary to achieve
a sufficient level of power or to determine the probability that the effect size of
interest will be detected with a certain sample size. This is known as prospective
power analysis (Peterman, 1990).
In 1996, SI/MAB undertook a study of biodiversity in the Lower Urubamba
region of southeast Peru. At the time, SI/MAB was working jointly with Shell
Prospecting and Development, Peru (SPDP) in a unique venture aimed at achieving
environmentally responsible development of natural gas and condensate resources
(Dallmeier and Alonso, 1997; Alonso and Dallmeier, 1998, 1999). As part of the
assessment of vegetation, we established 11 1-ha BDPs in 3 vegetation types in
the region (Comiskey et al., 2001). We also established sets of 10 MWPs around
each of 4 of the BDPs to evaluate spatial variability in species composition. To
the best of our knowledge, this represents the first published use of MWPs in a
tropical forest ecosystem. In this article, we primarily focus on the use of MWPs
as assessment and monitoring tools based on 3 objectives: (1) describe and compare
composition and structure of the 4 sites using MWPs, (2) compare the results of
MWPs to those of the BDP methodology (for example, number of individuals,
total basal area and number of species) and (3) evaluate the ability of MWPs to
detect changes in populations (statistical power). The project was not designed as
an experiment with hypotheses for testing. Still it allowed us to investigate the
value and uses of MWPs.

2. Study Area

The study was conducted in the Peruvian Amazon, one of the most biologically
diverse regions in the world (Gentry, 1988b, 1990). The study area, approximately
20 30 kilometers (km), is situated at 12 S latitude and 73 W longitude between
Manu National Park and the Apurimac Reserve Zone. The rugged terrain is charac-
terized by a series of plateaus separated by steep sloping hills that vary in elevation
from approximately 300 to 600 meters (m). A detailed description of the study area
can be found in Dallmeier and Alonso (1997).
The dominant vegetation of the area is lowland tropical rainforest. Initially,
4 sites near gas drilling operations were chosen for assessment and monitoring:
San Martin-3 (Sanm-3), Cashiriari-2 (Cash-2), Cashiriari-3 (Cash-3) and Pagoreni
(Pag). Each was located in old-growth terra firme forest (sensu Prance, 1989), a
habitat common in lowland neotropical rainforest. Terra firme forest is an upland
habitat described as having high species richness and an abundance of large-stature
trees (Campbell et al., 1986; Gentry, 1988b; Korning et al., 1991; Valencia et
al., 1994; Comiskey et al., 2001). Species composition was similar among the
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 23

sites and all were dominated by the arborescent palm Iriartea deltoidea. Natural
disturbances were common in the area (Comiskey et al., 2001).
Temperatures varied little across the study region and throughout the year, with
a mean of about 24 C, and relative humidity typically exceeded 80%. Rainfall
averaged between 3000 and 3500 millimeters a year and occurred mostly during
the wet season, October through April. (Alonso and Dallmeier, 1999).

3. Methods

3.1. F IELD METHODS


In 1998, we established 4 permanent 100 100 m BDPs in four nonadjacent units
of mature terra firme forest. The BDPs were subjectively placed in what botanists
viewed as the most homogeneous part of the habitat. Because of rugged terrain,
accessibility was also a consideration. Plots were established according to Dall-
meier (1992). The process included locating, measuring, marking and mapping
all trees with a diameter at breast height (dbh) 4 centimeters (cm); dbh was
measured immediately above the tree buttress when present. Further details along
with comprehensive results for all plots are described in Comiskey et al. (2001).
Voucher specimens for all species recorded in this study have been deposited at
the Museo de Historia Natural, Universidad Nacional Mayor de San Marcos in
Lima, Peru and the Smithsonian Institution, National Museum of Natural History,
Washington, DC.
Surrounding each of the BDPs within the same habitat unit and between 50 and
300 m from the well site, we randomly established 10 0.1-ha MWPs for a total of
40 MWPs. At each well site, 4 trails were cut into the forest extending out from
the well site in the cardinal directions. To identify a plot site, we first randomly
selected a trail, then randomly selected a distance between 50 and 300 m along
the trail, a random direction between 1 and 360 and then a random distance from
the trail in that direction between 0 and 100 m. Plots were oriented with the long
axis in the east-west direction. MWPs could not be located within or overlap the
boundaries of a BDP, they could not be placed on an incline of more than 50 and
they had to be within a similar vegetation type as that for the BDP.
The MWPs (Figure 1) followed the configuration of Stohlgren et al. (1995).
Each consisted of fourteen multi-scale, rectangular, nested subplots of consist-
ent proportional dimensions, which were corrected for slope. All trees 10 cm
dbh (large trees) were measured, marked and identified throughout the 0.1 ha
(50 20 m) MWP (D). In the center 20 5-m subplot (C), all trees 5 cm dbh
(medium trees) were measured, marked and identified. In a similar fashion, all
trees 1 cm dbh (small trees) were measured in the two 5 2-m subplots (B1 and
B2). All individual stems of herbaceous plants (herbs) were identified and counted,
but not marked, in the 10 2 0.5-m subplots (A). For further details see Alonso et
al. (1999).
24 P. CAMPBELL ET AL.

Figure 1. Diagram of modified Whittaker plot and subplot establishment.

3.2. D ESCRIPTIVE AND COMPARATIVE DATA ANALYSIS

For each set of 10 MWPs, total number of individuals, total basal area and total
number of species for each size class were generated. In addition, for each para-
meter the mean values per MWP and standard errors (SE) were estimated. Total and
mean number of individuals and total and mean basal area can also be generated for
each species, but these results are not presented in this manuscript. Means among
the four sets of MWPs were then compared using one-way analysis of variance
(ANOVA).
We used detrended correspondence analysis (DCA; Hill and Gauch, 1980) to
examine the similarity in the species composition of the sampling plots. Species
presence and absence data for each individual MWP and BDP were used in the
analysis. Because of possible effects caused by discrepancies in size between indi-
vidual MWPs and BDPs, we also conducted a DCA using cumulative presence and
absence data for each set of MWPs together with individual BDPs in the analysis.
Species Importance Values (SIV) were used to describe the ten most dominant
species in each BDP and each set of MWPs. The SIV is equal to the sum of the
relative density (total individuals of a species/total number of individuals of all
species), the relative basal area (total basal area of a species/total basal area of all
species) and the relative frequency (number of plots in which a species occurs/total
number of plots sampled) for each species. Relative frequency was equal to 1 in
calculations for the BDPs. Species were then ranked according to SIV, and the
species with the highest SIV was considered the most important in the plot.

3.3. S TATISTICAL POWER ANALYSIS

We used prospective statistical power analysis (Cohen, 1977), to investigate the


effectiveness of MWPs in detecting changes in mean number of species and mean
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 25
TABLE I
Total and mean number of species and standard deviation for 10 0.1-ha modified Whittaker plots
(MWP) by size class in the Lower Urubamba region, Peru (size classes based on diameter at breast
height [dbh], measurements in centimeters, means compared by One-Way Analysis of Variance
[ANOVA])

Site Total Mean (Standard Deviation) per MWP by size class (dbh) Total
10 cm 10 5, <10 1, <5a, b Herbaceousa, c all size
dbh classes

Cashiriari-2 198 43.3 (5.6) 9.1 (3.0) 8.1 (2.0) 53.6 (5.4) 419
Cashiriari-3 183 36.3 (5.1) 6.5 (3.6) 6.2 (3.6) 49.0 (8.0) 403
Pagoreni 234 43.0 (7.6) 7.5 (3.1) 11.6 (4.9) 68.2 (8.2) 509
San Martin-3 192 40.1 (7.0) 8.7 (4.4) 9.0 (4.1) 72.2 (13.4) 435
a p < 0.05 (ANOVA).
b Pairwise comparisons based on Bonferroni ad hoc test: Pagoreni > Cashiriari-3.
c Pairwise comparisons based on Bonferroni ad hoc test: Pagoreni > Cashiriari-2 and Cashiriari-3.
San Martin-3 > Cashiriari-2 and Cashiriari-3.

number of individuals. There are 5 parameters of interest when examining stat-


istical power: sample size, or number of plots (n); effect size, or the degree of
change we want to detect (ES); a measure of within-sample variability ( 2 ); and
the probabilities of Type I () and Type II () errors. In this study, n = 10 in all cases
and variance was estimated from initial study results. We also used statistical power
analysis to investigate the effectiveness of MWPs to detect changes in population
parameter estimates for the single species Iriartea deltoidea in the 10 cm dbh
size class and Philodendron guttiferum in the herb class. I. deltoidea was chosen
because it was the most abundant tree in this class at all sites, while P. guttiferum
was the most common species in the herbaceous layer throughout the Cash-2 site
and thus provided adequate data. Statistical power analyses were done on data from
the Cash-2 site only. We used PC-SIZE: Consultant software version 1.02 (Dallal,
1990) to calculate number of plots necessary to achieve selected effect sizes and to
calculate statistical power based on ES.

4. Results

4.1. S PECIES RICHNESS

The initial assessment of the vegetation in the Lower Urubamba region supports
the premise that the forests of western Amazonia are among the most species rich
on Earth. At each of the 4 sites, MWPs recorded more than 400 species. The most
speciose site was Pag, which recorded 509 total species and 234 species of large
trees in 1 ha of total sampling area (Table I). Mean species richness for large and
26 P. CAMPBELL ET AL.

TABLE II
Total and mean number of individuals and standard deviation for 10 0.1-ha modified Whit-
taker plots (MWP) by size class in the Lower Urubamba region, Peru (size classes based on
diameter at breast height [dbh], measurements in centimeters, means compared by One-Way
Analysis of Variance [ANOVA])

Site Total Mean (Standard Deviation) per MWP by size class (dbh)
10 cm dbh 10 5, <10 1, <5a, b Herbaceousa, c

Cashiriari-2 657 65.7 ( 9.8) 10.3 (3.2) 8.5 (2.7) 119.8 (28.5)
Cashiriari-3 608 60.8 ( 5.6) 7.6 (4.3) 6.7 (3.9) 138.1 (37.3)
Pagoreni 588 58.8 (12.4) 8.7 (3.9) 13.2 (5.5) 193.5 (44.1)
San Martin-3 647 64.7 (14.3) 11.7 (6.1) 9.8 (4.0) 214.2 (54.5)
a p < 0.05 (ANOVA).
b Pairwise comparisons based on Bonferroni ad hoc test: Pagoreni > Cashiriari-3.
c Pairwise comparisons based on Bonferroni ad hoc test: Pagoreni > Cashiriari-2 and
Cashiriari-3. San Martin-3 > Cashiriari-2 and Cashiriari-3.

TABLE III
Total and mean basal area (m2 /ha) and standard deviation for 10 0.1-ha modified Whittaker plots
(MWP) by size class in the Lower Urubamba region, Peru (size classes based on diameter at
breast height [dbh], measurements in centimeters, means compared by One-Way Analysis of
Variance [ANOVA])

Site Total Mean (Standard Deviation) per MWP by size class (dbh)
10 cm dbh 10 5, <10 1, <5a, b

Cashiriari-2 29.7 3.0 (0.6) 0.04 (0.02) 0.004 (0.002)


Cashiriari-3 39.7 4.0 (0.7) 0.03 (0.02) 0.003 (0.002)
Pagoreni 30.1 3.0 (1.4) 0.04 (0.02) 0.006 (0.003)
San Martin-3 33.1 3.3 (1.0) 0.04 (0.02) 0.004 (0.002)
a p < 0.05 (ANOVA).
b Pairwise comparisons based on Bonferroni ad hoc test: Pagoreni > San Martin-3.

medium trees was similar among the 4 terra firme sites, although Pag tended to be
richer in small trees while both Pag and Sanm-3 tended to be richer in herbs (p <
0.05, ANOVA); (Table I).

4.2. F OREST STRUCTURE

The 4 sites were structurally similar as well. Total abundance of large trees for each
set of MWPs (total area per set = 1 ha) ranged from 588 to 657, while total basal
area varied from 29.7 to 39.7 m2 ha1 . There were no differences among sites (p >
0.05, ANOVA) in the mean number of individuals or mean total basal area between
the 2 largest size classes (Tables II and III). However, Pag and Sanm-3 had a greater
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 27
TABLE IV
Summary results for the composition and structure of trees 10 cm diameter at breast height
in 4 1-ha biodiversity plots (BDP), and the total accumulation for corresponding sets of 10
0.1-ha modified Whittaker plots (MWP) in the Lower Urubamba region, Peru (source data for
BDP from Comiskey et al., 2001)

Site # of individuals Total basal area (m2 /ha) # of species


BDP MWP BDP MWP BDP MWP

Cashiriari-2 592 657 34.9 29.7 155 198


Cashiriari-3 564 608 28.6 39.7 159 183
Pagoreni 575 588 27.8 30.1 185 234
San Martin-3 481 647 22.2 33.1 138 192

mean abundance of individuals than the other 2 sites among small trees and herbs
(Tables II and III).
Modified Whittaker Plots and BDPs revealed similar patterns in the structure
of the vegetation. Direct comparisons between the 2 methods were restricted to
large trees (dbh 10 cm) because this is the only category where both methods
collected the same data on the same scale; that is, number of species, number of
individuals and total basal area per 1 ha. In all cases, MWPs recorded more species,
more individuals and greater total basal area for trees of the same size over the
same area than the BDPs (Tables IIV). However, both methods described Pag as
being the most speciose and Cash-2 as having the highest abundance of individuals
(Tables IIV). Results for BDPs are reported in more detail in Comiskey et al.
(2001).

4.3. O RDINATION

Similarity of species composition among all plots based on the results of the DCA
using species presence and absence data is shown in Figures 2 and 3. Plots that
are closer together on the graph are more similar in species composition and vice
versa. When all 40 MWPs and 4 BDPs are analyzed individually (Figure 2), we
can see that each set of 10 MWPs were distinctive and tended to be more similar to
each other than to any of the other sets. And in all cases, except for Pag, the species
composition of the BDPs was closely related to the corresponding set of MWPs.
Because of differences in area and because the environmental gradients crossed
by the 2 plot designs were probably not the same between individual MWPs and
BDPs, there may have been erroneous results. So we also analyzed the data using
the combined species composition for each set of MWPs versus each BDP. In this
case, the DCA again revealed that the species composition of the BDPs was closely
related to the corresponding set of MWPs, except for Pag (Figure 3).
28 P. CAMPBELL ET AL.

Figure 2. Results of a Detrended Correspondence Analysis for 40 0.1-ha modified Whittaker plots
(MWPs) and 4 1-ha biodiversity plots based on species presence/absence data for trees 10 cm,
diameter at breast height, in the Lower Urubamba region, Peru. Filled shapes represent BDPs, open
shapes represent MWPs. Eigenvalue for Axis 1 = 0.43, eigenvalue for Axis 2 = 0.35.

4.4. C OMMONNESS AND RARITY

Species richness was high, and many species were rare. At all sites, 40% or more
of all species among all size classes encountered were found in only 1 MWP, and
fewer than 13% were found in more than 5 MWPs (Figure 4). There was also a
large proportion of species that were represented by a single individual. Among
all size classes, 34% of the species encountered in the MWPs were represented
by a single individual, and nearly half the species 10 cm dbh were singletons
(Table V). A similar pattern was apparent in the BDPs where at least 41% of species
10 cm dbh were represented by a single individual.
Because rarity is common, we would not expect much overlap in species com-
position among sites. Therefore, it may be of more value to examine and compare
only important species (high SIV) at each site (Table VI). The two most important
species in each of the BDPs were always among the most important in the MWPs.
The large palm I. deltoidea was the most important species 10 cm dbh in the
study. MWPs ranked it the most important species at 3 sites and second at Pag,
while BDPs ranked it as most important at all 4 sites. In addition to I. deltoidea,
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 29

Figure 3. Results of a Detrended Correspondence Analysis comparing species presence/absence data


for trees 10 cm, diameter at breast height at 4 sites in the Lower Urubamba region, Peru. At each
site, species composition was recorded in 10 0.1-ha Modified Whittaker Plots (open symbols) and
4 1-ha biodiversity plots (filled symbols). Data from each set of 10 modified Whittaker plots was
summed and total area therefore represents 1 ha. Eigenvalue for Axis 2 = 0.33.

TABLE V
Number of species and proportion of total represented by 1 individual in
each of 4 1-ha biodiversity plots (BDP) and 4 sets of 10 0.1-ha modified
Whittaker plots (MWP) at 4 sites in the Lower Urubamba region, Peru
(in 1-ha plots, all individuals 10 cm diameter at breast height [dbh]; in
MWPs, individuals shown as 10 cm dbh, all individuals counted)

Site BDP MWP


dbh 10 cm dbh 10 cm All size classes

Cashiriari-2 74 0.48 100 0.51 177 0.42


Cashiriari-3 65 0.41 89 0.49 158 0.39
Pagoreni 102 0.55 117 0.50 245 0.48
San Martin-3 67 0.49 93 0.48 147 0.34
30 P. CAMPBELL ET AL.

Figure 4. Proportion of species recorded in a given number of modified Whittaker plots (MWP) at 4
sites in the Lower Urubamba region, Peru. Each site is represented by 10 MWPs.

there were other similarities among MWPs and BDPs. Most of the important spe-
cies at Cash-2 and Pag were the same in the 2 methods, while 4 species were the
same at Sanm-3; however, only 2 of 10 important species were the same in Cash-3.

4.5. P OWER ANALYSIS

Modified Whittaker Plots were most effective in detecting changes in the mean
number of species for large trees and herbs. In our analysis, 10 MWPs would be
sufficient to detect a change of 11% in the mean number of species of herbs, and
they would be able to detect a 14% change in large trees ( = 0.05, = 0.20).
However, it would be more difficult to detect trends in the 2 middle size classes
(Figure 5). Some authors (Elzinga et al., 1998) have suggested increasing (which
will increase power) when Type II errors are of concern; in other words, increasing
our confidence level if we do not want to miss a significant change. By weakening
our criteria for rejecting a Type I error to = 0.10, ten MWPs would be sufficient
to detect a change of 9% in the mean number of species of herbs, and they would
be able to detect an 11% change in large trees. By examining power curves, we
see that the probability of making a Type II error ( = 1 power) when trying to
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 31

TABLE VI
The 10 most important species at each of 4 sites in the Lower Urubamba
region, Peru as determined by modified Whittaker plots (MWP; import-
ance based on Species Importance Values, the sum of relative density
[total individuals of a species/total number of individuals of all species],
relative basal area [total basal area of a species/total basal area of all
species] and relative frequency [number of plots in which a species oc-
curs/total number of plots sampled] for each species; species listed in
descending order of importance; also shown, species importance rank of
the species as determined by 1-ha biodiversity plots [BDP] at each site
[Comiskey et al., 2001])

Site Species Rank in BDPa

Cashiriari-2 Iriartea deltoidea 1


Rinorea guianensis 5
Matisia cordata 2
Gustavia peruviana NR
Inga edulis 35
Calatola venezuelana 3
Sapium marmieri 7
Pentagonia parviflora 8
Guarea macrophylla 10
Cecropia engleriana 39

Cashiriari-3 Iriartea deltoidea 1


Otoba parvifolia 71
Bombacopsis sp1 NR
Pterygota amazonica 20
Pseudolmedia laevigata 24
Poulsenia armata NR
Hevea brasiliensis 13
Pseudolmedia laevis 3
Hyeronima alchorneoides NR
Ficus killipii NR

Pagoreni Matisia cordata 4


Iriartea deltoidea 1
Hevea brasiliensis NR
Guarea kunthiana 9
Hura crepitans NR
Pseudolmedia laevigata 18

a NR = Not recorded in the 1-ha biodiversity plot.


32 P. CAMPBELL ET AL.

TABLE VI
(continued)

Site Species Rank in BDPa

Pagoreni (continued) Guarea pterorhachis 12


Pentagonia parvifolia 2
Socratea salazarii 11
Bauhinia tarapotensis 7

San Martin-3 Iriartea deltoidea 1


Matisia cordata 2
Pentagonia parvifolia 8
Poulsenia armata 3
Otoba parvifolia 19
Lunania parviflora 22
Apeiba membranacea NR
Inga sp1 50
Neea chlorantha 62
Sloanea sp1 26

a NR = Not recorded in the 1-ha biodiversity plot.

detect relatively small changes is low for large trees and herbs, but much higher for
medium and small trees (Figure 6).
Modified Whittaker Plots were slightly less effective in detecting changes in the
mean number of individuals for large trees and herbs. In our analysis, 10 MWPs
would be sufficient to detect a change of 25% in the mean number of individuals
of herbs, and they would be able to detect a 15% change in large trees (Figure 7);
( = 0.05, = 0.20). Again, MWPs were less effective at detecting trends in the
small and medium tree classes. By examining power curves, we see that the prob-
ability of making a Type II error ( = 1 power) when trying to detect relatively
small changes is low for large trees and herbs, but much higher for medium and
small trees (Figure 8).
MWPs were least effective in detecting changes in the abundance of individual
species. The power of MWPs to detect such changes is lower than for the 3 size
classes of trees. For example, the population of I. deltoidea trees (mean = 3.5, sd =
2.1) could change by 60% and that of P. guttiferum (mean = 5.2, sd = 3.5) by 70%
before 10 MWPs would detect the change.
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 33

Figure 5. Number of modified Whittaker plots required to detect various amounts of proportional
change in the mean number of species in each of 4 size classes (classified by diameter at breast
height) present at Cashiriari-2. Effect size is the proportional change in mean number of species
relative to the initial estimate. In this case, = 0.05 and = 0.20.

5. Discussion

There are 2 critical pieces of information that are necessary when establishing
an assessment and monitoring program for vegetation: (1) a comprehensive as-
sessment of the species present and (2) an estimate of spatial variability to make
comparisons. In addition, the program should provide some measure of its ability
to detect potential changes in parameters of interest. Our preliminary evaluation of
the use of MWPs as an assessment and monitoring tool indicate that they are useful
in addressing each of these issues.

5.1. A SSESSMENT

The first step in any long-term monitoring program is the initial assessment. The as-
sessment provides the baseline data that allows scientists to identify components of
the environment in need of monitoring and to set the objectives on which they can
build the monitoring program. However, it is challenging to obtain these baseline
data in tropical forests where species richness is high (Hubbell and Foster, 1983;
Gentry, 1988b) and rarity is the norm (Greig-Smith, 1964; Hubbell and Foster,
34 P. CAMPBELL ET AL.

Figure 6. Power curves for detecting various proportions of change in mean number of species at
Cashiriari-2. Species are classified according to size (diameter at breast height). In this case, =
0.05 and n = 10.

1983). Therefore, a technique that best captures the variability in species richness
is preferred. This is important because to detect change we need to have a measure
of the natural variability in the sample.
Plot shape and distribution have influence on the number of species sampled.
The rectangular shape of the MWPs allows us to detect more of the species present
in an area than other shapes. Rectangular plots sample more species per unit area
than square plots because long sampling units are more likely to cover more di-
verse parts of the sampling area and to include species that tend to occur farther
apart (Podani et al., 1993; Stohlgren, 1995; Condit et al., 1996). Furthermore,
noncontiguous plots tend to detect more species than contiguous or single plots.
This is because plots placed closely together are usually more similar in species
composition than plots spaced further apart (Hubbell and Foster, 1983; Stohlgren
et al., 1997). Replicated, rectangular plots spread randomly throughout the study
area should capture a more representative sample of the variability in the study
area and thus maximize the number of species detected, which is ultimately the
objective of the assessment phase. In all cases, MWPs recorded more species and
more rare species than their associated square BDP.
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 35

Figure 7. Number of modified Whittaker plots required to detect various amounts of proportional
change in the mean number of individuals in each of 4 size classes (classified by diameter at breast
height) present at Cashiriari-2. Effect size is the proportional change in mean number of individuals
relative to the initial estimate. In this case, = 0.05 and = 0.20.

Plot size can also influence the number of species detected and their estim-
ated spatial variance. Plots that are too small will under represent large individu-
als, while measuring small individuals in large plots can be too time consuming
(Kent and Coker, 1994). A nested arrangement, like that used in MWPs, allows
for sampling of all size classes by using plots that are related to the size of the
individuals being sampled and minimizing the effort involved in sampling the plot
(Kent and Coker, 1994; Podani et al., 1993).
The MWPs recorded more individuals over 1 ha than their associated BDPs
in all cases (Table IV). This may be an effect of the size and shape of MWPs,
but we attribute this outcome to either sampling error in the MWPs or the sub-
jective placement of the BDPs. The BDPs tended to be placed on level ground,
whereas the MWPs were established randomly across the hilly terrain. In a study
that examined landscape-scale variation in tropical forest structure, Clark and Clark
(2000) showed that abundance tended to be higher on slopes than on flat ground.
Therefore, the BDPs may be underestimating true abundance. Future studies re-
garding methodologies for monitoring of tropical forests should examine this issue
in more detail.
36 P. CAMPBELL ET AL.

Figure 8. Power curves for detecting various proportions of change in mean number of individuals
af Cashiriari-2. Species are classified according to size (diameter at breast height). In this case, =
0.05 and n = 10.

5.2. M ONITORING

The purpose of monitoring is to detect changes, if any, from the baseline values
acquired during the assessment. Detecting changes requires an estimate of spatial
variation in species composition, which is possible through the use of randomly
distributed and replicated plots. A random presentation reduces bias in the results,
and through increasing the number of replicates, we obtain a more precise estimate
of species composition by decreasing variability in the sample. An estimate of
variation allows for statistical comparisons between sites spatially and comparis-
ons within sites temporally and, ultimately, a measure of change in both species
composition and abundance.
The MWPs recorded similar species composition, numbers of individuals and
total basal area of large trees as did BDPs. The results from this study provide
a clear example as to the benefits of replicated plots for monitoring. If our study
only involved non-replicated BDPs, we would have been tempted to conclude that
abundance was higher in Pag and Cash-2 and that basal area was much lower
in Sanm-3 because there was no measure of variability. But abundance estimates
from the 4 sets of MWPs show that there is no difference in the mean number of
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 37

individuals at these sites. Furthermore, the lower values recorded in Sanm-3 and
Cash-3 BDPs are well within the range of values obtained at the sample of MWPs.

5.3. P OWER ANALYSIS

The objective when monitoring biodiversity is to determine whether a particular


population parameter is increasing or decreasing. As examples, the parameter of
interest may be abundance, canopy cover or species richness. To meet this objec-
tive, managers often ask how many samples are necessary to detect a change and
what is the probability that a change can be detected if it exists. Statistical power
analysis (Cohen, 1977) is a methodology that provides a means by which managers
can answer these questions.
Type II errors are of particular concern. The failure to reject a null hypothesis
when in fact it is false, or the failure to detect a trend in the population parameter of
interest when it exists, could potentially lead to negative and irreversible impacts
on the species of interest. It is often better to be cautious and assume a change has
taken place even if it may not be real. As a result, many researchers have begun to
advocate the use of power analysis in monitoring (Toft and Shea, 1983; Gerrodette,
1987; Steidl et al., 1997; Van Strien et al., 1997).
We examined the power of MWPs using the data from the Cash-2 site. We base
our example on a power of 0.80, a commonly accepted value (Osenberg et al.,
1994; Zielinski and Stauffer, 1996). This means that the probability of correctly
rejecting a null hypothesis that is false is 0.80, or alternatively the probability of a
Type II error is 0.20.
With just 10 MWPs, we were able to detect changes between 10 and 15% in
mean number of species of large trees and herbs. The risk of not detecting was
only 0.20. This has obvious value to those monitoring biodiversity. If the overall
biodiversity in a region is declining, managers will be able to detect this decline at
an early stage and can take management measures to mitigate the decline. MWPs
did not work as well for the middle size classes, perhaps because the smaller
sampling areas do not capture sufficient individuals. However, detecting changes in
the small trees is of critical importance. Change occurs slowly among large mature
trees, and it may take many years to observe a 10% change in this class. But change
occurs rapidly in the smaller size classes because of higher mortality rates (Hubbell
and Foster, 1990; Clark and Clark, 1992). It is important to employ a method that
can detect change with the same ability. The flexibility of MWPs offers potential
solutions, one of which is to add additional MWPs. Increasing the sample size
always leads to a lower probability of both Type I and Type II errors (Cohen, 1977).
In addition, MWPs are quick to establish and survey. In the Lower Urubamba, we
established and surveyed two MWPs per day with a team of 5 experienced people.
Stohlgren et al. (1997) report similar speed with smaller crews. To compare, it
required a crew of 10 people for 10 days to establish a BDP. Thus to double the
sample size of MWPs would have taken us less than 3 weeks.
38 P. CAMPBELL ET AL.

It is not necessary to establish entire MWPs. In our case, it would require 40


plots per site to detect subtle changes (15%) in the medium tree class. Rather than
establishing all of the plots, there are 2 options to obtain a higher statistical power.
The first would be to change the sampling criteria of the subplots. One could record
information on medium trees in the largest subplots; that is, count trees 5 to 10 cm
dbh in the D plots, then reevaluate the results to see if a change in design provides
a more precise estimate. Another solution would be to add additional subplots
outside of the primary MWP. The entire MWP need not be established because
the sample size is already sufficient to detect acceptable changes in the larger size
class. One could add 30 C subplots to reach a sufficient sample size for medium
trees (Figure 5).
Although MWPs show evidence of having high statistical power when used for
detecting overall trends, they do not seem to have the power to detect changes in
single species. In our example, 10 MWPs would not detect a trend in the total
number of I. deltoidea trees until this value changed by 60% or a change in P.
guttiferum herbs until the value changed by 70%. In most cases, this level of power
would be inadequate. Furthermore, both of these species were common throughout
the area. Our ability to detect changes in rare species would be even less. Other
methods should be used when the objective is to monitor for a single species. One
is the plotless method used by Clark and Clark (1992) that aims at monitoring a
representative number of individuals of different sizes for a single species.

6. Conclusions and Recommendations

Stohlgren et al. (1995) suggest that MWPs may be valuable tools for quantifying
and detecting trends in species richness. Our preliminary investigations suggest that
this is true. MWPs have value as both short-term and long-term tools for assess-
ment and monitoring of biodiversity. They are easy to establish and cost effective,
yet they yield abundant information, and their flexibility allows them to be easily
modified to fit any situation. They also detect similar patterns in species abundance
and habitat structure as do standardized BDPs. Ultimately, a monitoring program
would benefit from using a combination of MWPs and larger BDPs. MWPs provide
information on spatial patterns of species, allow for statistical comparisons and can
be used to detect trends in richness over time. BDPs can be used to examine the
structure of forests and population dynamics, and they allow comparisons to other
studies by using standard protocols. We believe that a network of MWPs would
better enable managers to determine placement of large BDPs. By understanding
the spatial distribution of species and vegetation types, managers would be better
equipped to choose monitoring sites that are most representative of the area in
question.
MODIFIED WHITTAKER PLOTS AS AN ASSESSMENT AND MONITORING TOOL 39

Acknowledgements

The authors extend their appreciation to all who helped make this project a positive
experience including the Smithsonian Institution Monitoring and Assessment of
Biodiversity Program. We especially wish to thank the staff of Shell Prospecting
and Development Peru for their assistance, hospitality and enthusiasm and the
reviewers of the manuscript for invaluable comments.

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