A Biotic Index for the Southeastern United States: Derivation and List of Tolerance Values,

with Criteria for Assigning Water-Quality Ratings

Author(s): David R. Lenat
Source: Journal of the North American Benthological Society, Vol. 12, No. 3 (Sep., 1993), pp.
279-290
Published by: Society for Freshwater Science
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J. N. Am. Benthol. Soc., 1993, 12(3):279-290
? 1993 by The North American Benthological Society

A biotic index for the southeastern United States: derivation and

list of tolerance values, with criteria for assigning
water-quality ratings
DAVID R. LENAT
North CarolinaDivision of EnvironmentalManagement, Water Quality Section,
4401 Reedy Creek Road, Raleigh, North Carolina 27607 USA

Abstract. North Carolina'sDivision of EnvironmentalManagementhas a large numberof semi-

quantitative stream macroinvertebratecollections that have been assigned water-qualityratings.
These semiquantitativedata use abundancevalues of Rare= 1, Common = 3, and Abundant = 10,
allowing the calculationof mean abundance(range = 0-10) for each taxon acrossfive water-quality
classes. This informationwas used to derive tolerancevalues and classificationcriteriafor a south-
eastern biotic index. Classificationcriteria were adjusted for both season and ecoregion, but no
correctionswere requiredfor streamsize. Tolerancevalues are listed for >500 North Carolinataxa,
and are comparedwith a similar Wisconsin data base.
Keywords: biotic index, benthic macroinvertebrates,streams,water quality,pollution, biological
monitoring, North Carolina.

Benthic macroinvertebrates are often collect-
ed to help evaluate water quality and/or habitat
quality. This task involves collection of repre-
sentative samples, accurate taxonomy, and some
system to convert invertebrate data into water-
quality ratings. North American stream ecolo-
gists have struggled to establish such ratings
for over 50 years, and a bewildering array of
methods have been proposed (Cairns and Pratt
1993). Currently, more than one type of data
summary (metric) is often used to summarize
invertebrate data, with a final evaluation of wa-
ter quality based on several independent meth-
ods. Many of the proposed metrics, however,
are applicable only to a particular type of col-
lection, a single habitat, or a single geographic
area.
The most widely used metrics are taxa rich-
ness and "biotic indices". Taxa richness is as-
sumed to be inversely related to the degree of
stress, whereas biotic indices attempt to sum-
marize information on the tolerance of the mac-
roinvertebrate community. The North Carolina
(NC) Division of Environmental Management
(DEM) uses taxa richness of the most intolerant
invertebrate groups (Ephemeroptera + Plecop-
tera + Trichoptera, or "EPT")and a biotic index
similar to that of Hilsenhoff (1987). Both metrics
are weighted equally in assigning water-quality
ratings (NC DEM 1992). We have invested sub-
stantial efforts in testing and evaluating both
methods, both in deriving criteria and in mak-
ing adjustments for the effects of ecoregions,
279
stream size, and season. The EPT taxa richness
criteria have been in use for many years, but
we have only recently derived biotic index cri-
teria. This paper describes the method used to
produce the North Carolina Biotic Index (NCBI)
and gives tolerance values for taxa in south-
eastern USA. The emphasis was on the objective
derivation of tolerance values, because other
investigators have already demonstrated the
utility and accuracy of a biotic index (Chutter
1972, Hilsenhoff 1982, 1987, 1988, Narf et al.
1984, Jones et al. 1981).
The first North Carolina biotic index was es-
tablished in 1980, using "expert opinions" to
assign tolerance values for each taxon. Trial val-
ues were assigned by NC DEM biologists, and
adjusted following discussion at a meeting of
benthic ecologists from North Carolina, South
Carolina and Georgia. This index (with poten-
tial values ranging from 0-5) was modeled on
that of Hilsenhoff (1977), which was, in turn,
derived from Chutter's (1972) index:
Sum TV,N,
Total N
where TV, is the tolerance value of the ith taxa,
N, is the abundance of the ith taxa, and Total
N is the number of individuals in the sample.
Total N may be based on either actual densities
(No./m2, etc.) or abundance categories (see
Methods).
Ideally, such an index would be used to target
specific kinds of stress. For example, a species

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280
might be sensitive to metal toxicity, but tolerant
of low dissolved oxygen. Rather than a single
index, an array of indices would be used to
assess water quality and/or habitat quality. Un-
fortunately, this level of information for species
tolerance is not available. Dr. Hilsenhoff tar-
geted his index at organic pollution, although
it is used by other investigators to look at many
other kinds of stress. The NCBI is intended for
examination of the general level of pollution,
regardless of source.
The utility and accuracy of such an index is
only as good as the system used to derive the
tolerance values. The use of expert opinion val-
ues proved to be unreliable in North Carolina,
as biotic index ratings frequently conflicted with
ratings from EPT taxa richness values or ratings
based on professional judgment. Unbiased as-
signment of tolerance values requires a large
number of invertebrate collections that have
already received some kind of water-quality rat-
ing. To my knowledge, only Hilsenhoff (1987)
has used such a method to derive tolerance val-
ues. Other lists of invertebrate tolerance in use
within the United States are modifications of
Hilsenhoff's list with "expert opinion" modi-
fications for taxa not found in Wisconsin. North
Carolina biologists saw a need for indepen-
dently deriving tolerance values for southeast-
ern species, as well as deriving criteria for biotic
index values that could be adjusted for differ-
ences in ecoregion, season, and stream size.
Methods
The NC DEM has a data set of >2000 mac-
roinvertebrate stream samples collected during
water-quality surveys between 1983 and 1992.
These samples have been assigned to five water-
quality categories (bioclassifications): Poor (n =
299), Fair (n = 392), Good-Fair (n = 481), Good
(n = 492), and Excellent (n = 424); they include
both standardized qualitative samples (Lenat
1988) and collections limited to Ephemeroptera,
Plecoptera, and Trichoptera (Eaton and Lenat
1991). Classification is based on EPT taxa rich-
ness and an earlier version of the NCBI.
A variety of information is associated with
each sample, including ecoregion, date, and
stream size. The abundance information for each
taxon is semiquantitative, being tabulated as ei-
ther Rare (1-2/sample), Common (3-9/sample)
or Abundant (>10/sample), and it is coded in
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D. R. LENAT [Volume 12
the computer system, respectively, as 1, 3 or 10.
Calculations of mean abundance for this study
also included zero values, since I used all sam-
ples in a particular water-quality category, not
just samples where this taxon occurred. Similar
abundance categories are used with the Chan-
dler Score, a European biotic index (Chandler
1970).
The primary measurement used to compute
the tolerance value for any taxon was the av-
erage abundance (range = 0-10) in each water-
quality category. I computed how far each taxon
(starting from Excellent) extended into the areas
of poorer water quality. The abundance values
(by water-quality category) were converted into
cumulative percentiles and graphed vs. the wa-
ter-quality score (1 = Excellent, 2 = Good, 3 =
Good-Fair, 4 = Fair, 5 = Poor). I tried a 50th
percentile, a 75th percentile, and a 90th per-
centile value; the 75th percentile produced the
greatest separation of intolerant and tolerant
species (Table 1), while still leaving the facul-
tative species with values near the midrange.
Calculations of such percentiles usually re-
quired the use of simple linear interpolation
between categories. For example, the mean
abundance data (number per sample, by water-
quality class) for Cricotopusbicinctus in Table 1
is: Excellent (1): 1.0, Good (2): 2.4, Good-Fair
(3): 3.1, Fair (4): 4.0, and Poor (5): 3.7. Convert-
ing these abundance values to a cumulative per-
centage produces: 7.0, 23.9, 45.8, 73.9, 100.0. This
indicates that the 75th percentile for the water-
quality score lies between Fair (4) and Poor (5)
and yields by interpolation a value of 4.04.
This procedure produced a range of prelim-
inary tolerance values generally between 1.0
and 4.5, but the desired range (to be comparable
with Hilsenhoff 1987) was 0-10. If the 1-4.5
range is graphed vs. the 0-10 range (with 1 and
4.5 being the x and y intercepts), a conversion
formula for tolerance value (TV) is derived:
Final TV = 2(1.43 x Preliminary TV - 1.43).
The 75th percentile technique was best suited
for taxa with substantial amounts of data. If only
a few data points were available, then the 75th
percentile value often was too high. Review of
our data base suggested that the 75th percentile
was suitable for taxa with at least 25 observa-
tions (out of the 2000 samples). For taxa with
only 10-24 collections, a 50th percentile was
used, and the old "expert opinion" values were
1993] A BIOTIC INDEX FOR SOUTHEASTERN USA 281

TABLE1. Average abundance values (range = 0-10) by water-quality rating (five categories) of represen-
tative taxa, and North Carolina tolerance values (TV) calculated from this information. "+" = <0.1, N = No.
of records for taxon.

Water quality and score

Excel-
lent Good G-F Fair Poor
1 2 3 4 5 N NC TV

Group 1: Intolerant
Drunella wayah 0.4 0.1 + + - 51 0.0
Rhithrogenaspp. 0.7 0.2 0.1 - 155 0.4
Chimarraspp. 2.7 2.9 1.4 0.4 + 650 2.8
Micrasema wataga 0.8 0.8 0.5 0.1 + 280 3.2
Goera spp. 0.6 0.2 + + - 172 0.4
Brachycentrusspinae 0.8 0.1 + + + 73 0.0
Pteronarcys dorsata 0.8 0.7 0.1 + 186 1.8
Acroneuriaabnormis 6.2 4.6 1.7 0.4 + 1031 2.2
Mean TV (?1 SD) 1.4 (+1.3)
Group 2: Facultative
Stenonema modestum 6.6 8.1 8.2 6.9 1.1 1660 5.8
Ephemerellacatawba gr. 3.8 1.9 1.7 0.7 0.4 550 4.0
Eurylophellatemporalis 1.6 1.5 1.2 0.6 0.2 518 4.6
Cheumatopsychespp. 5.5 6.9 7.1 6.6 2.5 1680 6.6
Hydropsyche venularis 1.8 2.7 2.4 1.5 0.3 656 5.4
Perlesta placida 1.5 1.9 1.8 1.1 + 524 5.0
Ancyronyx variegata 1.2 2.3 2.8 2.5 1.0 726 7.0
Polypedilum convictum 2.3 3.1 3.0 1.8 0.4 622 5.2
Mean TV (?1 SD) 5.4 (?1.0)
Group 3: Tolerant
Cricotopusbicinctus 1.0 2.4 3.1 4.0 3.7 744 8.8
C. tremulus gr. 0.3 0.6 1.1 1.3 1.4 279 9.0
Chironomusspp. 0.6 1.0 1.5 3.1 5.0 711 9.8
Polypedilum illinoense 1.4 2.8 2.9 3.9 5.2 932 9.2
Physella spp. 0.8 2.0 2.3 3.5 3.8 825 9.0
Argia spp. 1.7 2.7 3.3 4.0 4.1 914 8.6
Limnodrilushoffmeisteri 0.6 0.9 1.3 2.3 4.0 561 9.8
Asellus spp. 0.5 0.6 1.2 1.4 2.0 390 9.4
Mean TV (?1 SD) 9.2 (?0.4)

used for taxa with <10 records. This system is
designed to be compatible with existing biotic
indices currently in use for the southeastern
United States.
Average biotic index values were calculated
for all standardized qualitative samples, and
analysis of variance methods (two-way and
three-way ANOVA) were used to examine the
effects of three ecoregions (Mountain, Pied-
mont, Coastal Plain), four seasons, four stream
width categories (1-2 m, 3-7 m, 8-20 m, >20
m) and five water-quality ratings (Excellent,
Good, Good-Fair, Fair, Poor). Seasons were de-
fined as Summer: June-August; Fall: Septem-
ber-November; Winter: December-February;
and Spring: March-May.
For the ANOVA calculations, the water-qual-
ity rating again used a score from 1-5. There
were few differences between biotic index val-
ues for flowing streams in the Piedmont and
Coastal Plain ecoregions, so these two areas were
combined in subsequent calculations. After
making seasonal corrections (see Results), the
prior water-quality ratings were then used to
derive classification criteria for the NCBI. For
initial tests of tolerance values, 24 taxa were
chosen that could be roughly divided into In-
tolerant, Facultative, or Tolerant categories.

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282 D. R. LENAT [Volume 12

TABLE2. Seasonally correcteda average NCBI by for each ecoregion in North Carolina. I made
ecoregion and water-quality rating, with ANOVA re- tables for biotic index values vs. season for each
sults. of the ecoregions, and attempted simple addi-
tive corrections that would equalize the average
Water-quality rating NCBI values for all seasons. Fall, winter, and
Excel- Good- spring values were corrected to the average
Ecoregion lent Good Fair Fair Poor summer value. The following system proved
Mountain 3.78 4.58 5.52 6.32 7.80 satisfactory. For the Mountain ecoregion, 0.4
Piedmont/Coastal 5.02 5.65 6.26 7.09 8.32 was added to the fall (September-November)
biotic index, and 0.5 was added to the spring/
Two-way ANOVA winter (December-May) biotic index. For the
Mean
Source df square F-value Piedmont/Coastal Plain ecoregions, 0.2 was
added to the spring/winter biotic index. After
Ecoregion 2 88.5 307.8
4 making these corrections to the NCBI, season
Water-quality score 265.8 924.0
could be eliminated as a significant variable
Ecoregion x score 8 2.4 8.5
Residual 1467 0.2 - when the effects of season and water-quality
class were examined within specific ecoregions
a For mountain streams: 0.5 added to
winter/spring (p = 0.29 for Mountain ecoregion, p = 0.36 for
collections, 0.4 added to fall collections. For Pied- Piedmont/Coastal Plain).
mont/Coastal streams: 0.2 added to winter/spring
collections. Three-way ANOVA examining the effects of
stream width, ecoregion, and water-quality class
on NCBI values (seasonally corrected) indicated
These a priori choices were based on profes- that all three factors significantly affected biotic
sional judgment and were intended to be non- index values (p = 0.001), but that width effects
controversial, but I included taxa known to be were very small (F = 6.8), relative to Ecoregion
tolerant of both organic and toxic materials. (F = 228.4) and Water-quality Score (F = 831.4).
Criteria for each water-quality class were then
derived using mean values (Table 2) as the mid-
Results
point for each class. This procedure showed that
For the 24 taxa chosen for initial testing, the NCBI values for Piedmont/Coastal streams were
distribution of average abundance values across almost one unit higher than for Mountain
five water-quality categories always produced streams of comparable water quality (Table 3).
unimodal patterns, and confirmed the initial
assignment of these taxa as intolerant, faculta- Discussion
tive, and tolerant (Table 1). The calculation of
tolerance values (using the 75th percentile) pro- North Carolina had a data set of 2000+ stream
duced low values for the intolerant group (mean macroinvertebrate samples which were divided
= 1.4), high values for tolerant taxa (mean = into five water-quality ratings. This data set was
9.2), and intermediate values for the facultative used to derive preliminary tolerance values for
group (mean = 5.4). over 500 taxa. A regression equation then was
Appendix 1 is a complete list of tolerance
values for North Carolina stream invertebrates.
TABLE3. Criteriafor North CarolinaBiotic Index
Nomenclature generally follows that of Brig- after seasonal corrections.
ham et al. (1982). Comparable values from a
Wisconsin data base (Hilsenhoff 1987) also have
been listed. Some taxa have not been included Water Ecoregion
in this list, either because we have insufficient quality Piedmont/
class Mountain Coastal
data to calculate a tolerance value (<10 collec-
Excellent <4.18 <5.24
tions), or because a taxon can only be identified
with unpublished keys for particular North Good 4.17-5.09 5.25-5.95
Good-Fair 5.10-5.91 5.96-6.67
Carolina taxa. Fair 5.92-7.05 6.68-7.70
Two-way ANOVA showed that differences in Poor >7.05 >7.71
season significantly affected biotic index values

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1993] A BIOTIC INDEX FOR SOUTHEASTERN USA
used to stretch the initial range of water-quality
scores (1-4.5) to a 0-10 range (NCBI). I then
tested for the effects of season, stream size, and
ecoregion on NCBI. Simple additive seasonal
corrections were derived, correcting spring, fall
and winter values to the mean summer NCBI
values. Stream size did not have a large effect
on NCBI values (relative to ecoregion and water
quality), but Piedmont/Coastal streams had
NCBI values about one unit higher than Moun-
tain streams (Table 2).
Hilsenhoff (1987) used similar procedures to
derive tolerance values for Wisconsin streams
from a data set of 2000+ invertebrate collec-
tions. His initial tolerance values were based on
expert opinion, but were later modified based
on the mean biotic index value for sites where
any given taxon was collected. This procedure
produced a range of values from 1.4-3.9 on a
possible scale of 0-5 (W. L. Hilsenhoff, Univer-
sity of Wisconsin, personal communication).
These numbers were then "stretched" to cover
the entire range of a 0-10 scale. Dr. Hilsenhoff's
values, however, were based on means, while
the NC values used a 75th percentile figure.
Furthermore, the Hilsenhoff Biotic Index (HBI)
was only intended to monitor organic pollu-
tion, while the NCBI measures any stress that
either lowers EPT taxa richness or promotes the
development of tolerant species.
The North Carolina tolerance values func-
tioned well to separate examples of intolerant,
facultative, and tolerant taxa (24 taxa: Table 1).
Comparisons of these tolerance values with Hil-
senhoff's (1987) data were complicated by a
number of problems. Sometimes no comparable
values could be found (3 taxa), sometimes com-
parisons could only be made by using different
species in the same genus (2 taxa), and some-
times tolerance values were listed at a species
level in one data set, but at the generic level in
the other data set (9 taxa). Using the most com-
parable information from the Wisconsin data
set, mean tolerance values for the Intolerant,
Facultative and Tolerant groups in Table 1 were
0.6, 4.1, and 6.1. Although the trend for the
mean tolerance values is similar to the North
Carolina data (Table 1), the values for the Tol-
erant examples showed considerable difference:
6.1 vs. 9.2.
Some species had North Carolina populations
that were rated as more tolerant than compa-
rable Wisconsin populations. For example, the
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283
mayfly flies Stenonema modestumand species in
the Ephemerellacatawbagroup are fairly tolerant,
abundant, and widespread in North Carolina.
Wisconsin data, however, had suggested that
these are very intolerant species. It is possible
that these "populations" actually are different
species that are not separable with current tax-
onomic keys. These problems illustrate the per-
ils of using a biotic index outside its intended
geographic range.
Mean tolerance values for the major taxonom-
ic groups were similar for the North Carolina
and Wisconsin data sets (Table 4). In particular,
both the tolerance values and the rankings for
Ephemeroptera, Plecoptera and Trichoptera
were very close. Chironomidae (as a group) were
ranked somewhat differently for the two data
sets, but comparable taxa were often similar in
their tolerance values. The greatest between-
region differences occurred for Megaloptera and
Odonata.
Some of the differences between North Car-
olina and Wisconsin tolerance values may re-
flect differences in collection methods. The Wis-
consin samples were limited to riffle samples,
whereas North Carolina uses multiple-habitat
collection methods. More slow-water species
(especially Odonata) would occur in the mul-
tiple-habitat collections, and possibly affect the
assignment of tolerance values.
Both our data and Hilsenhoff's data showed
that biotic index values vary with season. Hil-
senhoff's values are standardized for spring and
fall collections, with a subtraction of 0.5 during
summer months (Hilsenhoff 1988). North Car-
olina data, however, are standardized for sum-
mer collections, with some corrections for other
periods of the year. For mountain streams, I
have suggested adding 0.4-0.5 during non-
summer months, similar to the corrections pro-
posed for Wisconsin streams. During periods of
expected change (late spring, fall), stream rat-
ings should be made with caution.
Biotic index values also appear to vary with
ecoregion. This may be related to differences in
stream temperature, although other variables
also may affect between-ecoregion differences.
North Carolina mountain streams and Hilsen-
hoff's Wisconsin streams had similar criteria
ranges; both areas support cold-water fish spe-
cies (especially trout). If the Wisconsin data are
seasonally corrected for summer collections
(subtract 0.5, Hilsenhoff 1988), then the NC
284 D. R. LENAT [Volume 12

TABLE4. Comparisons of tolerance values (TV), by taxonomic group (ranked by Mean TV), for North
Carolina and Wisconsin (Hilsenhoff 1987). N = No. of records/group.

North Carolina Wisconsin

Differencea
Mean Mean
Group Rank N TV Rank N TV N Mean

Plecoptera 1 62 1.8 1 39 1.3 20 +1.1

Trichoptera 2 102 2.3 2 87 2.9 48 <0.1
Ephemeroptera 3 94 2.7 3 72 3.3 49 +0.5
Diptera: Chironomidae 4 150 5.7 8 71 6.3 71 +0.3
Coleoptera 4 29 5.7 6 28 4.8 9 -0.2
Mollusca 6 17 6.1 Not included - -
Megaloptera 7 4 6.2 4 5 3.6 3 +2.9
Diptera: miscellaneous 8 30 6.4 5 48 4.3 19 +1.6
Odonata 9 33 6.9 7 35 5.4 18 +2.2
Crustacea 10 11 7.5 9 6 7.3 Not comparable
Oligochaeta 11 23 8.2 Not included

a North Carolina TV minus Wisconsin

TV, applied only to comparable taxa and not derived from Mean TV
values.

mountain criteria and the Wisconsin criteria be- Literature Cited

come even more similar, rarely differing by > 0.1.
NC Piedmont and Coastal Plain streams are lim-
ited to warm-water fisheries, and NCBI values
for these ecoregions (for streams with similar
water-quality ratings) were much higher rela-
tive to the mountain ecoregion.
Biotic index values were not strongly affected
by stream width, a pattern very different from
EPT taxa richness values (NC DEM, unpub-
lished data). For this reason, biotic indices are
more reliable than taxa richness when ratings
are assigned to smaller streams.
North Carolina streams have been rated by
the Division of Environmental Management us-
ing EPT taxa richness. The development of a
biotic index allows for a second method of rat-
ing. Our current system produces a final rating
that receives equal weight from both methods.
Acknowledgements
The data used to derive this biotic index are
based on the work of many ecologists in the
NC DEM, including Trish MacPherson, David
Penrose, Larry Eaton, Neil Medlin, and Ferne
Winborne. I am grateful to these individuals
for constructive comments during the devel-
opment of the North Carolina biotic index. I
thank Ken Eagleson for developing computer
programs that generated the tolerance values.
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for the biotic index. Great Lakes Entomologist 21: NARF, R. P., E. L. LANGE, AND R. C. WILDMAN. 1984.
9-13. Statistical procedures for applying Hilsenhoff's
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APPENDIX 1. Tolerance values for North Carolina stream macroinvertebrates, with comparable values for
Wisconsin taxa (Hilsenhoff 1987). * = based on less than 25 collections for North Carolina data or less than
5 collections for Wisconsin data. ** = comparisons based on differing levels of taxonomy. NC taxonomy
according to Brigham et al. (1982).

Taxon NCBI WIBI Taxon NCBI WIBI

Ephemeroptera Ephemerellaberneri 0.0*

Acentrella amplus 3.6 Ephemerellacatawba (gr) 4.0 1*
Acerpenna pygmaeus 3.7 4 Ephemerellahispida 0.6
Ameletus lineatus 2.1 0* Ephemerellainvaria (gr) 2.2 1
Attenella attenuata 2.6 3* Ephemerellaneedhami 0.0* 2
Baetis ephippiatus 3.9 Ephemerellarotunda 2.8
Baetis flavistriga 7.2 4 Ephemerellaseptentrionalis 2.0
Baetis frondalis 8.0 5* Ephoronleukon 1.5 2*
Baetis intercalaris 5.8 6 Eurylophellabicolor 5.1 1
Baetis pluto 4.8 Eurylophellacoxalis 2.6
Baetis propinquus 6.2 6* Eurylophellafuneralis 2.3 0*
Baetis tricaudatus 1.8 Eurylophellatemporalis 4.6 5
Baetiscaberneri 0.6* Eurylophellaverisimilis 0.3*
Baetisca carolina 3.6 5** Habrophlebiavibrans 0.0
Baetiscagibbera 1.4 Heptageniajulia 0.5*
Brachycercusspp. 3.5 3* Heptagenia marginalis 2.5
Caenis spp. 7.6 7 Heptagenia pulla 2.3
Callibaetisspp. 9.3 9 Heptagenia spp. 2.8 3**
Centroptilumspp. 6.3 2* Heterocloeonspp. 3.6 2*
Cinygmula subaequalis 0.0 Hexagenia spp. 4.7 6
Cloeon spp. 7.4 4* Isonychia spp. 3.8 2
Dannella lita 0.0* 4* Leptohyphesspp. 2.0*
Dannella simplex 3.9 2* Leptophlebiaspp. 6.4 4
Drunella allegheniensis 1.3 Leucrocutaaphrodite 2.5 1**
Drunella conestee 0.0 Leucrocutaspp. 0.0 1**
Drunella cornutella 0.0 0 Litobrancharecurvata 0.0* 6*
Drunella lata 0.1 Neoephemerapurpurea 2.1
Drunella tuberculata 0.2 Paracleodesspp. 8.7
Drunella walkeri 1.0 Paraleptophlebiaspp. 1.2 1
Drunella wayah 0.0 Potamanthusspp. 1.6 4
Epeorusdispar 1.0 Pseudocloeonspp. 4.4 4**
Epeoruspleuralis 2.0 Rhithrogenaamica 0.0*
Epeorusrubidus 1.4 Rhithrogenaexilis 0.0*
Epeorusspp. 1.2 0** Rhithrogenafuscifrons 0.0*
Ephemeraguttalata 0.0 Rhithrogenaspp. 0.4 0
Ephemeraspp. 2.2 1** Serratellacarolina 0.0*

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286 D. R. LENAT [Volume 12

APPENDIX 1. Continued.

Taxon NCBI WIBI Taxon NCBI WIBI

Serratella deficiens 2.7 2 Malirekus hastatus 1.4

Serratellaserrata 2.7 Neoperla spp. 1.6 1*
Serratellaserratoides 1.5 Paracapniaangulata 0.2 1
Siphlonurusspp. 2.6 7 3.5
Paragnetinafumosa
Siphloplectonspp. 3.1 2*
Paragnetinaichusa 0.0
Stenacron carolina 1.7 1.7
Paragnetina immarginata
Stenacron interpunctatum 7.1 7
Stenacronpallidum Paragnetinakansensis 2.0
2.9
Perlesta placida 4.9
Stenonema carlsoni 2.1
Perlesta spp. (not perlesta) 0.0
Stenonemafemoratum 7.5 5
Stenonema integrum 5.5 4* Perlinella drymo 0.0 1*
Stenonema ithaca 4.1 Prostoia spp. 6.1 2**

Stenonema lenati 2.3 Pteronarcysdorsata 1.8

0
Stenonema mediopunctatum 1.7* 3 Pteronarcysspp. 1.7 0
Stenonema meririvulanum 0.3 Remenus bilobatus 0.3
Stenonema modestum 5.8 1 Shipsa rotunda 0.3* 2
Stenonemapudicum 2.1 Strophopteryxspp. 2.5 3**
Stenonema terminatum 4.5 4 Suwallia spp. 0.0*
Stenonema vicarium 1.0 2 Sweltsa spp. 0.0
Tricorythodesspp. 5.4 4 Taeniopteryxburksi 5.8
Taeniopteryxmetequi 1.4
Plecoptera 6.3 2
Taeniopteryxspp.
Acroneuriaabnormis 2.2 0** 1.4
Talloperlaspp.
Acroneuriaarenosa 2.2
Yugus arinus 0.0*
Acroneuriacarolinensis 0.0
Yugus bulbosus 0.0
Acroneurialycorias 1.5
Acroneuriamela 0.9 Trichoptera
Agnetina annulipes 0.0* Agapetus spp. 0.0 0
Agnetina flavescens 0.0* Agarodes libalis 0.0* 3*
Allocapnia spp. 2.8 3 Anisocentropuspyraloides 0.8
Alloperla spp. 1.4 0* Apatania spp. 0.6
Amphinemuraspp. 3.4 3** Arctopsyche irrorata 0.0
Beloneuriaspp. 0.0 Brachycentrusappalachia 1.1
Clioperlaclio 4.8 1 Brachycentruschelatus 0.0*
Cultus decisus 1.6 Brachycentruslateralis 0.4 1*
Diploperla spp. 2.0* Brachycentrusnigrosoma 2.2
Diploperla duplicata 2.7 Brachycentrusnumerosus 1.8 1
Diploperla morgani 1.5 Brachycentrusspinae 0.0
Eccopturaxanthenes 4.1 Brachycentrusspp. 2.2
Haploperlabrevis 1.3 1 Ceraclea ancylus 2.5 3*
Helopicus bogaloosa 0.0* Ceraclea nr excisa 0.0*
Helopicus subvarians 0.8 Ceracleaflava 0.0*
Isogenoideshansoni 0.0* Ceraclea maculata 6.4
Isoperlabilineata 5.5 Ceraclea transversa 2.7
Isoperla dicala 2.2 2 Ceratopsychealhedra 0.0* 3
Isoperla holochlora 0.0 Ceratopsychebifida 1.0
Isoperla namata (gr) 1.8 Ceratopsychebronta 2.7 5
Isoperla nr holochlora 0.0 Ceratopsychemacleodi 0.9
Isoperla nr slossonae 1.3 Ceratopsychemorosa 3.2 2
Isoperla orata 0.0 Ceratopsycheslossonae 0.0 4
Isoperla similis 0.7 Ceratopsychesparna 3.2 1
Isoperla slossonae 2.6 Ceratopsycheventura 0.0
Isoperla transmarina(gr) 5.6 0 Cheumatopsychespp. 6.6 5
Leuctraspp. 0.7 0 Chimarraspp. 2.8 4**

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1993] A BIOTIC INDEX FOR SOUTHEASTERN USA 287

APPENDIX 1. Continued.

Taxon NCBI WIBI Taxon NCBI WIBI

Cyrnellus fraternus 7.4 8* Psychomyiaflavida 3.3 2

Diplectrona modesta 2.2 0* Psychomyia nomada 2.0
Dolophilodesspp. 1.0 0** Ptilostomis spp. 6.7
Fattigia pele 1.1 Pycnopsychegentilis 0.8
Glossosomaspp. 1.5 00 Pycnopsycheguttifer 2.7
Goera spp. 0.3 0** Pycnopsyche lepida 2.5
Helicopsyche borealis 0.0 3 Pycnopsyche scabripennis 4.0
Heteroplectronamericanum 2.9 Pycnopsyche spp. 2.3 4
Hydatophylax argus 2.3 2 Rhyacophilaacutiloba 0.0
Hydropsyche betteni 8.1 6 Rhyacophilaatrata 0.0
Hydropsyche decalda 4.1 Rhyacophilacarolina 0.0
Hydropsyche demora 1.8 Rhyacophilafuscula 2.0 O*
0*
Hydropsyche hageni 0.0* Rhyacophilaledra 3.4
Hydropsyche incommoda 5.0 7 Rhyacophilaminor 0.0*
Hydropsyche phalerata 3.7 1* Rhyacophilanigrita 0.0
Hydropsyche rossi 4.9 Rhyacophilatorva 1.8
Hydropsyche scalaris 3.0 2 Rhyacophilavuphipes 0.0
Hydropsyche venularis 5.3 Setodes spp. 0.9 2
Hydroptila spp. 6.2 6 Triaenodesabus 4.3
Ironoquiapunctatissima 7.3 3* Triaenodesinjusta 2.2
Lepidostomaspp. 1.0 1 Triaenodestardus 4.7 6**
Leucotrichiapictipes 4.3 2 Wormaldiaspp. 0.4 0"*

Lype diversa 4.3 2

Macrostemumspp. 3.6 3** Coleoptera
Matrioptila jeanae 0.0* Anchytarsus bicolor 3.8
Micrasema bennetti 0.0 Ancyronyx variegatus 6.9 6
Micrasema burksi 0.0* Berosusspp. 8.6
Micrasema charonis 0.3 Copelatusspp. 9.1*
Micrasema rickeri 0.0 Coptotomusspp. 9.0
Micrasema rusticum 0.0* Dineutes spp. 5.5
Micrasema wataga 3.2 2 Dubiraphiaspp. 6.4 6
Molanna blenda 3.9 6** Ectoprianervosa 4.3 5
Mystacides sepulchralus 3.5 4 Enochrusspp. 8.5
Nectopsyche candida 3.8 Falloporusspp. 2.8
Nectopsyche exquisita 4.2 3** Gyrinus spp. 6.3
Nectopsyche pavida 4.2 Helichus spp. 5.4 5
Neophylax concinnus 1.2 Helophorusspp. 7.9
Neophylax mitchelli 0.0 Hydrocanthusspp. 6.9*
Neophylax oligius 2.6 Hydroporusmellitus 1.8
Neophylax ornatus 1.6* Hydroporusspp. 8.9
Neophylax spp. 1.6 3 Laccobiusspp. 8.0
Neureclipsis spp. 4.4 7 Laccophilusspp. 10.0
Nyctiophylax moestus 2.6 Macronychusglabratus 4.7 4
Nyctiophylax spp. 0.9 5 Microcylloepuspusillus 2.1 3*
Oecetis spl (stick case) 4.7 Optioservusspp. 2.7 4
Oecetis sp2 (stick case) 4.1 Oulimnius latiusculus 1.8
Oecetis spp. 5.7 8 Peltodytes spp. 8.5
Orthotrichiaspp. 7.2 Promoresiaelegans 2.2
Parapsyche cardis 0.0 Promoresiatardella 0.0
Phylocentropusspp. 5.6 4 Psephenus herricki 2.5 4
Polycentropusspp. 3.5 6 Sperchopsistessellatus 6.5
Protoptila spp. 2.8 1 Stenelmis spp. 5.4 5
Psilotreta spp. 0.0 0* Tropisternusspp. 9.8

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288 D. R. LENAT [Volume 12

APPENDIX 1. Continued.

Taxon NCBI WIBI Taxon NCBI WIBI

Megaloptera Coelotanypusspp. 6.2

Corydaluscornutus 5.6 6 Conchapelopiagroup 8.7 6
Nigronia fasciatus 6.2 Corynoneuraspp. 6.2 7
Nigronia serricornis 5.5 0 Cricotopusspp. 7
Sialis spp. 7.5 4 Cricotopusbicinctus(C/O spl) 8.7
Cricotopusinfuscatus gr
Odonata (CIO sp5) 9.0
Cricotopusvieriensis gr
Argis spp. 8.7 8**
(CIO sp46) 4.8
Basiaeschnajanata 7.7 6
Cricotopusvaripes gr (C/O sp6) 8.1
Boyeriagrafiana 6.3
Cricotopussylvestris gr
Boyeria vinosa 6.3 2
(C/O sp9) 10.0
Calopteryxspp. 8.3 5** Orthocladiusobumbratusgr
Cordulegasterspp. 6.1 3**
(C/O splO) 8.8
Dromogomphusspp. 6.3 Orthocladiusrobacki
Enallagmaspp. 9.0 9** 7.2
(C/O spl2)
Epicorduliaspp. 5.6 Orthocladiusspp. 6
Erythemissimplicicollis 7.7* Orthocladius(Euorthocladius)
Gomphusspiniceps 4.9
type III (C/O spl3) 6.3
Gomphusspp. 6.2 5* Orthocladius(E.) sp (C/O sp3) 9.2
Hagenius brevistylus 4.0 1* Orthocladiusnigritus gr
Hetaerina spp. 6.2 6 0.9*
(C/O sp47)
Ischnura spp. 9.4 9** Orthocladiusclarkei gr
Lanthus spp. 2.7 5.8
(C/O sp54)
Libellulaspp. 9.8 9** Orthocladiusdorenus gr
Macromia spp. 6.7 2** 6.7 *
(C/O sp7)
Nasiaeschna pentacantha 8.0
Cryptochironomusblarina gr 8.0 8**
Neurocorduliamolesta 3.3* 5*
Cryptochironomusfulvus gr 6.7 8**
Neurocorduliaobsoleta 5.4 0*
Cryptotendipesspp. 6.1 6
Neurocorduliaspp. 5.8
Demicryptochironomusspp. 2.1
Neurocorduliavirginiensis 1.6 Diamesa spp. 7.7 8*
Ophiogomphusspp. 6.2 1
Dicrotendipesmodestus 9.2 5
Pachydiplax longipennis 9.6
Dicrotendipesneomodestus 8.3
Perithemisspp. 10.0
Dicrotendipesnervosus I10.0
Plathemis lydia 10.0 8* I10.0
Dicrotendipessimpsoni
Progomphusobscurus 8.7
Dicrotendipesspp. 7.9
Somatochloraspp. 8.9 1* 7.7 8
Diplocladius cultriger
Stylogomphusalbistylus 4.8 0* Endochironomusnigricans 7.5* 8*
Sympetrumspp. 7.3 10*
Eukiefferiellaspp.
Tetragoneuriaspp. 8.5 8
Eukiefferiellaclaripennisgr
(E. spll) 5.7
Diptera: Chironomidae Eukiefferiellabrehmigr
Ablabesmyiamallochi 7.6 8** (E. spl2) 3.7
Ablabesmyiaparajanta/janta 7.1 Eukiefferiellagracei gr
Ablabesmyiapeleensis 4.6* (E. spl4) 2.7*
Apsectrotanypusjohnsoni 0.0* Eukiefferielladevonica gr
Brillia spp. 5.2 5 (E. sp2) 2.6
Brundiniellaeumorpha 3.8 Eukiefferiellabrevicalcargr
Cardiocladiusspp. 6.2 5 (E. sp6) 1.7
Chironomusspp. 9.8 10 Genus nr Nanocladius B 6.7
Cladopelmaspp. 2.5* 9* Glyptotendipesspp. 8.5 10
Cladotanytarsusspp. 3.7 7 Goeldichironomusholoprasinatus 10.0
Clinotanypuspinguis 9.1 8 Harnischiaspp. 7.5 8
Coelotanypusconcinnus 7.7 Heleniella spp. 0.0

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1993] A BIOTIC INDEX FOR SOUTHEASTERN USA 289

APPENDIX 1. Continued.

Taxon NCBI WIBI Taxon NCBI WIBI

Heterotrissocladiusspp. 5.4 0* Rheosmittianr delicatula 7.1

Hydrobaenusspp. 9.6 8 Rheotanytarsusspp. 6.4 6
Keifferulusdux 10.0 10* Robackiaclaviger 2.4
Labrundiniapilosella 6.0 7** Robackiademeijerei 4.3
Labrundiniavirescens 4.5* Saetheriatylus 8.1 4**
Larsiaspp. 8.3 6 Stelechomyiaperpulchra 4.6
Lopescladiusspp. 2.2 4* Stempellinaspp. 2.0* 2
Micropsectraspp. 1.4 7 Stempellinellaspp. 5.3 4
Microtendipesspp. 6.2 7 Stenochironomusspp. 6.4 5
Nanocladius downesi 2.6 Stictochironomusspp. 6.7 9
Nanocladius spp. 7.2 3* Sublettea coffmani 1.7
Natarsia spp. 10.0 8 Symposiocladiuslignicola 5.4
Nilotanypus spp. 4.0 6 Sympotthastiaspp. 5.7 2
Nilothauma spp. 5.5 2 Synorthocladiusspp. 4.7 2*
Odontomesafulva 5.9 4 Tanypus spp. 9.6 10
Pagastia sp. 2.2 1 Tanytarsusspp. 6.7 6
-
Pagastiella ostansa 2.6* Thienemaniellaspp. 6.0 6
Parachaetocladiusspp. 0.0 Tribelosspp. 6.6 5
Parachironomusmonochromus 7.9 Tvetenia spp. 5
Parachironomusspp. 9.2 10 Tvetenia bavaricagr (E. spl) 4.0
Paracladopelmaspp. 6.4 7* Tvetenia discoloripesgr (E. sp3) 3.9
Paracladopelmanereis 1.8* Unniella multivirga 0.0*
Paracladopelmaundine 5.2 Xenochironomusxenolabis 7.0 0**
Parakiefferiellacf. triqueta 5.6 Xylotopus par 6.6 2*
Parakiefferiellaspp. 5.9 Zavrelia spp. 2.7*
Paralauterborniellanigrohalteralis 4.8 8 Zavrelimyiaspp. 9.3 8
Paramerinaspp. 2.8
Parametriocnemuslundbecki 3.7 5** Diptera: Miscellaneous
Paratanytarsusspp. 7.7 6 Anopheles spp. 9.1
Paratendipesspp. 5.3 8 Antocha spp. 4.6 3
Pentaneura spp. 4.6 6 Atherix lantha 2.1 2
Phaenopsectraflavipes 8.5 Atrichopogonspp. 6.8
Phaenopsectrasp2 6.2 7 Blephariceraspp. 0.2 0**
Phaenopsectrafuscicorne Chaoboruspunctipennis 8.5 8**
(P. sp3) 6.0 Chrysopsspp. 7.3 6
Phaenopsectraspp. 6.8 7 Cnephia mutata 4.0* 5
Polypedilum angulum 5.6 Culex spp. 10.0
Polypedilum aviceps 4.0 Culicoidesspp. 6.5 10
Polypedilum convictum 5.3 Dicranota spp. 0.0 3
Polypedilumfallax 6.7 Dixa spp. 2.8
Polypedilum halterale 7.2 Dolichopodidae 9.7 4
Polypedilum illinoense 9.2 Empididae 8.1 6
Polypedilum scalaenum 8.7 Eristalisspp. 10.0 10
Potthastia gaedi 2.0 Hexatoma spp. 4.7 2
Potthastia longimanus 7.4 Limnophoraspp. 7.0*
Procladius spp. 9.3 9 Limoniaspp. 10.0 6
Prodiamesaolivacea 7.9 3** Palpomyia (complex) 6.9 6
Psectrocladiusspp. 3.8 8* Polymeda/ ormosiaspp. 6.5
Psectrotanypusdyari 10.0 10 Prosimuliummixtum 3.3 3
Pseudochironomusspp. 4.2 5 Prosimuliumspp. 2.6
Pseudorthocladiusspp. 0.0 O* Protoplasafitchii 5.0
Rheocricotopusrobacki(R. spl) 7.7 6** Pseudolimnophilaspp. 7.3 2
Rheocricotopustuberculatus Psychoda spp. 9.9*
(R. sp2) 6.8 * Simuliumspp. 4.4

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290 D. R. LENAT [Volume 12

APPENDIX 1. Continued.

Taxon NCBI WIBI Taxon NCBI WIBI

Simuliumvenustum 7.4 5 Cambarusspp. 8.1

Simuliumvittatum 8.7 7 Crangonyxserratus 8.1*
Tabanusspp. 9.7 5* Crangonyxspp. 8.0
Tipula spp. 7.7 4 Gammarusfasciatus 6.9
Hyallela azteca 7.9 8
Oligochaeta Lirceusspp. 7.7
Aulodrilus limnobius 5.2* Orconectesspp. 2.7
Aulodriluspigueti 4.7* Palaemonetespaludosus 6.7
Branchiurasowerbyi 8.4 Procambarusspp. 9.5
Cambarincolidae 6.2
Dero spp. 10.0 Mollusca
Enchytraeidae 10.0 Amnicola spp. 4.8
Haber speciosus 2.8 Campelomadecisum 6.7
Ilyodrilus templetoni 9.4 Elimia spp. 2.5
Isochaetidescurvisetosus 7.18 Ferrissiaspp. 6.9
Isochaetidesfreyi 7.6 Helisoma anceps 6.5
Limnodriluscervix 10.0 Laevapexfuscus 7.3
Limnodrilushoffmeisteri 9.8 Leptoxisspp. 1.6
Limnodrilusspp. 9.6 Menetus dilatus 8.4
Limnodrilusudekemianus 9.7* Physella spp. 9.1
Lumbriculidae 7.3 Pseudosuccineacolumella 7.2
Nais spp. 9.1 Somatogyrusspp. 6.5
Opisthopora (earthworm gr) 9.1 Stagnicola spp. 8.0
Peloscolex spp. 8.8 Corbiculafluminea 6.3
Pristina spp. 9.9 Elliptio complanata 5.4
Slavina appendiculata 7.1 Elliptio lanceolata 1.9*
Spirospermanikolskyi 7.7 Pisidium spp. 6.8
Stylaria lacustris 8.5 Sphaeriumspp. 7.7
Tubifextubifex 10.0 Other
Hirudinea Belastomaspp. 9.8
Batracobdellaphalera 7.1 Ranatra spp. 7.5
Erpobdella/ mooreobdellaspp. 7.8 Corixidae 9.0
Helobdella elongata 9.9 Climaciaareolaris 6.5
Helobdella stagnalis 6.7 Curaforemanii 5.1
Helobdella triserialis 8.9 Dugesia tigrina 7.5
Mooreobdellatetragon 9.7 Hydracarina 5.7
Placobdellapapillifera 9.0 Petrophilasp. 1.8 5
Placobdellaparasitica 6.6 Prostomagraecens 6.6
Crustacea
Asellus spp. 9.4 8
Astacidae 7.2

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