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INSECT COMMUNITIES AMONG
APPALACHIAN MOUNTAIN STREAM BRYOPHYTES r

Thesis for the Degree of Ph. D..
MICHIGAN STATE UNIVERSITY
JANICE M. GLIME
1968 '

 

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THESIS 3 1293 00994 2883

Michigan State
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This is to certify that the

thesis entitled

Insect Communities Among Appalachian Mountain
Stream Bryophytes.

presented by

Janice M. Glime

has been accepted towards fulfillment
of the requirements for

Ph.D, degree in Botany

Major professor

 

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ABSTRACT
Insncr commutes more APPALACHIAN MOUNTAIN s'rmu sarorarrms
By Janice in. sun

The primary purpose of this study was to survey the insect
fauna of Appalachian Mountain stream bryophytes. From this informa-
tion, certain implications of community relationships, adaptations, and
uses appeared.

Streams were sampled by hand grabs at arbitrary times and at
varying frequencies. Dry weight is the base used for quantifying the
data.

Among the 28 streams studied in Pennsylvania, Maryland, and

Virginia, three bryophyte-based streams are apparent: Eontinalis, the

 

Hygroamblystegium group, and Scapania. -A Fontinalis stream is generally

 

larger and has a continuous flow of water sufficient to submerge the moss

year-round. Probably due to its larger size, Fontinalis houses the

 

larger of the bryo-insects, but smaller ones occur here too. The gygrgf
amblystegium group comprises streams where several species of bryophytes
appear similar and make similar mats. Both their insect faunas and

the narrow, shallow streams they occupy are similar. These mats

provide homes for small insects. Scapania streams were only repre-
sented by two, but among all the Scapania collections the insects were
small. In Toliver Run, Scapania exhibited more Species and individ

duals of insects than Fontinalis in that same stream.

 

The most important bryo-insects, numberwise, appear to be Diptera
(Chironomidae and Simulidae), while Ephemeroptera, Plecoptera, and
Trichoptera are of secondary importance. But even these secondary
orders may exhibit disproportionately high numbers in individual streams
or during certain seasons.

As indicated by the seasonal trends in sizes, kinds, and numbers

of insects, one use of the bryophyte appears to be that of a nursery--

a substratum where hatchlings develop in protected chambers with a
flowing food supply. Other insects living there are tiny even until
they emerge from the water.. This adaptation of small size is often
accompanied by such adaptations as lateral compression, covered gills
or lack of gills, lack of appendages, or hooks for attachment.

As a result of this study, 150 insect taxa have been named from
bryophyte habitats, while only about 70 are common enough to be con-
sidered true bryo-insects. Among these, most collections have about 15-
20 species. Two of the caddis fly larvae appear to represent new genera_

of the families Hydroptilidae and érachycentridae.

INSECT COMMUNITIES
AMONG

APPALACHIAN MOUNTAIN STREAM BRYOPHYTES

By

Janice M. Glime

A THESIS

Submi t ted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Deparument of Botany and Plant Pathology
1968

gin/70

ACKNOWLEDGMENTS

I wish to express my sincere gratitude to Dr. William B. Drew,
my advisor in the Botany Department of Michigan State University, for
encouragement and counsel during the research and for constant devo-
tion of his time during the first writing of the theses. For enthusi-
asm and encouragement, Drs. Gordon C. Guyer of entomology and T.

Wayne Porter of zoology were a frequent source of revitalization,
while Drs. John E. Cantlon and Gerald w. Prescott of botany and Robert
C. Ball of fisheries and wildlife were willing to guide the research
by suggesting methods and sources of information. I especially wish
to thank Dr. Cantlon for guidance in completing the thesis during Dr.
Drew's absence.

Withoutithe technical assistance of numerous experts, I would
have been unable to complete this work. Sincere appreciation for
prompt assistance in identification of insects goes to: Dr. Oliver
S. Flint, Smithsonian Institution (Trichoptera); Dr. T. Wayne Porter,
Michigan State University (Microvclia); Dr. Lewis Berner, Florida
State University (Ephemerella); Mr. Richard J. Snider, Michigan State
University (Collembola); Dr. George w. Byers, Kansas State University
(Tipulidae); Dr. Glenn Wiggins, Royal Ontario Museum (Brachycentridae
and Hydroptilidae); Dr. Allen Knight and Mr. Dennis Heiman, Kellogg
Biological Station (Plec0ptera); Dr. Douglas M. Davies, McMaster
University, Canada (Simulidae), Mr. Julian P. Donahue, Michigan State
University (some Diptera); and Mr. Ronald willson, Michigan State
University (Coleoptera); while the bryOphytes were promptly determined

by Dr. Ronald A. Pursell, Pennsylvania State University (Fissidensh

ii

Dr. Clyde F. Reed, Baltimore; Dr. Harold Robinson, Smithsonian Museum;
Dr. Howard Crum, University of Michigan; Dr. Winona Welch, DePauw
University (Fontinalis).

Accurate assistance by technicians expedited the completion of
the research: Laurence Breslow, Stephen Gordon, Brian Dalrymple,
Robert Dye, William.Anderson, and Arlene Jim, while Patricia Clancy
volunteered her time to check the data. Miss Jim was the principal
technician, and it was largely through her efficient work that the
research could be completed so quickly.

During the field phase of the research, I welcomed the help of
my parents and Judith Wilburn, who transported me on numerous collect-
ing trips.

While writing the thesis, I appreciated suggestions of Dr. Kenneth
Cummins of Pymatuning Laboratory and Dr. Thomas Waters of the Univer-
sity of Minnesota, while Dr. Herman Struck of the Michigan State

University English Department criticized the structure of the writing.

iii

TABLE OF CONTENTS

 

 

Page

INTRODUCTION AND LITERATURE REVIEW 1
METHODS AND PROCEDURES 6
Choice of Site and Sampling Procedure 6
Preservation and Sorting ll
Counting Chamber . 12
Stream Data 13
RESULTS AND DISCUSSION 21
Fontinalis dalecarlica Streams 28
Muddy Creek 30

Neds Run » 34

Mud Run ‘ 34

Swamp Run, a Mud Run tributary 35

Pohopoco Creek tributary 36
Hygroamblystegium fluviatile Streams 37
Sideling Hill Creek tributary 4O

Sinking Creek 40

Mountain Lake tributary to Sinking Creek 41

Rock Castle Creek 42

Johns Creek tributary 42

Goose Creek 42

Hoyes Run and Ginseng Run 43

Piney Creek tributary 44

Deep Creek tributary 4S

Gramlich Run tributary 46

Little Bennett Creek 47

iv

Seneca Creek tributaries
Elk Creek
Dingman's Creek
Saw Creek

Sciaromium lescurii Streams
Pidcock Creek
Toliver Run tributary

Fissidens bryoides

 

Hygrohypnum
Scapania undulata Streams
Toliver Run
Laurel Run and Hoch Run
Insect Communities

Comparison of insect fauna of various aquatic
bryOphytes

Comparison of streams
Insect communities by seasons
Insect Biology and Ecology
Coleoptera
Collembola
Diptera
EphemerOptera
Plecoptera
TrichOptera
Insect Adaptations to BryoPhyte Life
Dorsal-ventral flattening

Lateral compression

Page
47
48
49
SO
51
51
52
52
53
S4
55
S6

69

7O

77

84
100
100
106
106
109
111
112
117
117

117

Page

Enlargement of adhesive surfaces 118
Small body 118
Attachments 119
Weighting ll9
Reduction of swimming hairs 119
Reapiration 120
Generalized feeding 120
CONCLUSIONS 121
TABULAR SUMMARY AND CONCLUSIONS 126
LITERATURE CITED 128

APPENDICES 133

vi

LIST OF TABLES

Table

I TAXONOMIC LIST OF BRYOFHYTES

II~ TABULAR SUMMARY OF STREAM CHARACTERISTICS
III COMMUNITY COEFFICIENTS BY BRYOFHYTES, MARCH
Iv COMMUNITY COEFFICIENTS BY BRYOFMYTES, MAY

v COMMUNITY COEFFICIENTS BY BRYOPHYTES, JUNE
VI COMMUNITY COEFFICIENTS BY BRYOPHYTES, SUMMER
VII COMMUNITY COEFFICIENTS BY STREAMS, MARCH
VIII COMMUNITY COEFFICIENTS BY STREAMS, MAY

IX COMMUNITY COEFFICIENTS BY STREAMS, JUNE

x COMMUNITY COEFFICIENTS BY STREAMS, SUMMER

XI TAXONOMIC LIST OF INSECTS AND ADAPTATIONS

vii

Page

22

71
71
71
72
78
79
80
81

101

LIST OF FIGURES

Figure Page
1 Comparison of Number of Species against Number of Grams of
Bryophyte 9
2 Increase in Number of Species with Addiion of Individuals,
Listed in the Order Collected 67
3 Relative Abundance in the Five Collecting Seasons 86

viii

LIST OF PLATES

Plate

1 MAP OF STREAM LOCATIONS

2 AVERAGE WARM SEASON PRECIPITATION

3 AVERAGE ANNUAL PRECIPITATION

4 AVERAGE JANUARY TEMPERATURE

AVERAGE JULY TEMPERATURE

Ln

AVERAGE NUMBER OF DAYS WITHOUT KILLING FROST

0‘s

7 FONTINALIS DALECARLICA WITH ATTACHED INSECTS

 

8 PENNSYLVANIA STREAMS

9 SAN CREEK AND DINGMAN'S CREEK

10 MARYLAND STREAMS: ALLEOANY AND CARRETT COUNTIES
11 MARYLAND STREAMS: MONTGOMERY COUNTY

12 VIRSINIA STREAMS

13 MARCH, TOLIVER RUN: RELATIVE NUMBER OF INSECTS IN
THREE SPECIES OF BRYOPHYTES

14 MARCH: RELATIVE NUMBERS OF INSECTS PER DRAM DRY WEITUT OF
BRYOPHYTE

15 MAY: RELATIVE NUMBERS OF INSECTS PER CRAM DRY WEIGHT OF
BRYOPHYTE

16 JUNE: RELATIVE NUMBERS OF INSECTS PER CRAM DRY WEISHT OF
BRYOPHYTE

l7 MID-SUMMER: RELATIVE NUMBERS OF INSECTS PER GRAM DRY
WEIGHT OF BRYOPHYTE

18 DECEMBER: RELATIVE NUMBERS OF INSECTS PER GRAM DRY WEITMT
OF BRYOPHYTE

ix

Page
14
16
17
18

19

31
S7
S9
61

63

75

89

91

93

95

97

Table

XII

XIII

XIV

XV

XVI

LIST OF APPENDICES

MARCH COUNTS OF INSECTS PER GRAM DRY WEIGHT OF
BRYOPHYTE

MAY COUNTS OF INSECTS PER GRAM DRY WEIGHT OF
BRYOPHYTE

JUNE COUNTS OF INSECTS PER GRAM DRY WEIGHT OF
BRYOPHYTE

SUMMER COUNTS OF INSECTS PER GRAM DRY WEIGHT OF
BRYOPHYTE

DECEMBER COUNTS OF INSECTS PER GRAM DRY WEIGHT OF
BRYOPHYTE

Page

135

150

15h

162

180

INTRODUCTION AND LITERATURE REVIEW

Streams, bryophytes, insects: all these have been studied by
many authors, but few have studiedthe interrelationships of the
three in depth. (Thienemann, 1912; Carpenter, 1928; Percival and White-
head, 1929, 1930; Illies, 1952; and Minckley, 1963 all included mosses
in their discussion of stream surveys.) Moreover, even fewer workers
have attempted to determine the relationships among insect communities
associated with bryOphytes in different streams or among several species
of bryophytes. Only Frost (1942) compared the fauna of mosses in an
acid and an alkaline stream, but she did not separate the moss species
in her analysis. Thus, the present study appears to be the first attempt
to compare the bryophyte fauna of streams of the Appalachian Mountains
in the Deciduous Forest Formation (Braun, 1964).

As the first study of its kind in the Appalachian area, this study
is an attempt to determine some of the natural history relationships
existing in the bryophyte-insect communities of flowing water in 28 mid-
dle Appalachian streams. Among the possible aspects for study, several
basic ones were chosen: 1) determination of common bryophyte taxa;

2) observation of ecological aspects of the streams where these bryo-
phytes occur; 3) observation of types of substrata which these bryophytes
provide (e. g. mat, streamer) for insects; 4) determination of taxa

of insects to be found among these bryophytes; 5) observation of any
apparent adaptation of insects to the bryophyte habitat; 6) consider-
ation of possible uses of the bryophyte by the insects; 7) determination
of the most frequent insects; 8) observation of recurrent arrays of
insect species; 9) determination of aspects which appear to warrant

detailed further study.

For the purposes of this study, the bryophytes themselves were the
only substratum of the stream to be considered. Since bryophyte
rhizoids do not appreciably penetrate the rock, the plants themselves
form the observable physical boundaries. But even in so simple a
system, different habitat zones can be detected, e. g., basal, surface,
and mat, and the boundaries of these zones blend in problematic transi-
tional areas. In this study, the entire moss stand is considered as
the community, and these habitat zones are recognized as habitats within
the community, but they are not considered individually in the analysis.
Because insects may pass easily from the surface zone to open water and
back again, the bryo-community fauna is herein defined as those organ-
isms which remain with the bryophyte when it is collected.

In addition to having the advantage of physical and biological
boundaries, this community study is one of the few attempts (according
to Whittaker, 1962) to use invertebrate-plant relations as a means of
defining a community, but it neglects other vital members of the a>mmu-
nity such as plankton, epiphytes, and other arthrOpods. In Whittaker's
coverage of literature on community studies, he states that marine and
littoral communities have been based on dominant animals (Shelford and
Towler, 1925; Newcombe, 1935; Clements and Shelford, 1939), but "Char-
acterization of terrestrial biomes by invertebrate animals has been
scarcely attempted." These terrestrial biomes are usually delimited
by their plant composition (Shelford, 1963; Braun-Blanquet, 1965) and
dominance (Braun, 1964; Shelford, 1963; Oosting, 1956), while inverte-
brates are usually ignored. Ross (1963) is an exception. By comparing
the distribution of aquatic insects with terrestrial plant biomes, he

found that certain insect genera had the same distribution pattern as

3

dominant plants of the biome, e. g. Agg£.saccharum Marsh., and that for
several caddisfly genera the distribution coincided with a terrestrial
biome. This he found was particularly true of smaller streams: that
factor influence (leaf fall, runoff, shade, temperature, rainfall) is
inversely related to stream size; therefore, larger streams are more
similar to those of other biomes; smaller ones are unique. Further-
more, biota of small streams are more restricted in their-composition.
Aside from the biome treatment of terrestrial systems, stream
biologists attempt to describe stream affinities on the basis of
physical, chemical, and biological similarities (Ricker, 1934; Van-
Deusen, 1953; Frost, 1942). To compare members of a single stream
system, Barrel (1966) correlated species diversity indices with stream
order (Horton, 1945; Leopold, 1962. Stream order refers to the
number of tributary junctions.) in the Otter Creek system, northcentral
Oklahoma. Furthermore; he found that physico-chemical conditions of
Otter Creek were closely related to stream order. That location of
a stream in the drainage basin is important in regulation of community
structure is evidenced by a third order adventitious stream which
flows directly into the sixth order stream of the Otter Creek system.
Contrary to expectation, this third order stream exhibits greater
similarity to higher order streams than to other third order streams.
To provide a common system of classification for plant and animal
communities, Klugh (1923) listed 30 associations, each of which he
further dissected into systases and cenoses. He is his own best
critic of the system when he states that "lines have to be drawn where
no hard and fast lines exist, and it must be borne in mind that the

ecotone...is usually a blending and not a sharp line.” According to

his system, streams would be in the Spring Association or Stream
Association, and these are further divided on the basis of flow,
position in stream (plankton, surface, bottom), and bottom type. By
his classification the stream bryophyte could be in the Stream Associ-
ation as an Emophyte Cenosis of a Slow-flowing Stream (Tachydromile
Systasis), or in the Spring Association as part of the Rapids Cenosis.
On the other hand, some workers (Ricker, 1934; Percival and Whitehead,
1929) refer to a mossregion of a stream, encompassing all of these
socies at once.

The literature abounds with faunal studies of stream communities.
In many studies fish have been used to describe the regions of the
stream (Ricker, 1934; Huet, 1949; VanDeusen, 1953; Kuehne, 1962).
Others have used upper, middle, and lower reaches and described the
invertebrate species within these (Berg, 1948; Wilburn, 1964). But
because of the difficulty of delimiting any one community, few studies
actually compare fauna among different streams. Further complicating
the problem of comparison are tremendous variations of physical and
chemical attributes and the necessity for different sampling methods
in different streams. Because the present study uses a biological
boundary rather than a physical one, it is possible that the require-
ment of bryophyte presence leads to greater homogeneity than one kind
of physical or chemical stream type. I expected that any given species
of bryophyte would itself be limited to certain streams and regions
within these streams by certain chemical and physical factors, and
thus encompass communities with more uniform abiotic conditions. When
Frost (1942) compared an acid and an alkaline stream, she found

Fontinalis squamosa Hedw. to be the dominant moss in the acid stream

 

(90 per cent of the bryophytes by weight), while Fontinalis antipyre-

5

£223 and Eurynchium_riparioides were the dominant mosses in the
alkaline one (51.3 per cent and 42.4 per cent of the bryophytes
respectively). Not only mosses differed between the chemically dif-
ferent streams; invertebrates were represented by different species

as well. In the present study bryophyte patches were not found in

all streams nor in all situations in the streams of occurrenCe.
Furthermore, the bryophyte species differed in different situations.

I therefore expect the biological heterogeneity throughout patches

of any given bryophyte species might be lower than throughout a single

physically delhmited habitat type.

METHODS AND PROCEDURES
Choice of Site and Sampling Procedure

To compare bryo-communities in a wide variety of streams,
collections were made in three states in the Appalachian Mountains:
Pennsylvania, Maryland, and Virginia (Plate 1). Here, any stream en-
countered could be included if it had submersed bryophytes in flowing
water. As it was soon discovered, mountain and high elevations were
most likely to have bryOphyte regions in their streams, so for
expediency, most collecting sites were selected in these areas. Areas
of apparent pollution or disturbance were eliminated. Most of the
streams could be sampled only once (primarily March and summer, as
noted later), but year-round collections (March, May, June, July,
August, and December) were made in Garrett County, Maryland, where
small, medium, and large streams permit a wide variety of habitats.

In any comparative community study, it is desirable to have a
uniform sample size. To accomplish this, previous workers collected
a specified area CMinckley, 1963) or attempted to estimate a uniform
weight in the field (Frost, 1942). Later, Frost's weight samples
were reduced to the specified weight (200 gms. wet weight) in the lab.
But in the present study collecting in many streams and many seasons
made it impractical to choose a uniform sample size: 1) In winter,
a large sample of bryOphytes was never available. 2) Some streams
have abundant growth while others have little. 3) Some bryophytes,
such as Fontinalis, were usually in large quantities, but others,

such as Scapania, were frequently scarce. 4) Volumetric and area

samples were impractical because of the irregular surface of the
substratum and the varying thickness of the bryophyte mat.

0n the basis of trials with various sampling techniques and
equipment, the method of hand sampling was chosen. By the simplest
possible means, the collector merely scrapes off a "handful" of
bryophyte with his fingers. (These samples range from .5 to 12
grams, where the low weights usually are Scapgnia and the Hygroambly-
stegium group, while higher ones are for Fontinalis. The area sampled
ranges from 20 to 100 square centimeters, occasionally reaching 200

square centimeters for Fontinalis.) To obtain a quantitative measure

 

of the amount of bryOphytes present, every sample is weighed on a
torsion balance after removal of insects and air drying. To compare
the dry weight with the wet weight used by Frost (1942), four samples
each of three bryophytes were also weighed wet with the following
factors of wet x dry weight: Fontinalis 2.9-4.7, mean 3.8; §capania
4.4-6.7, mean 5.2; Hygrgamblystegium.group 3.8-8.5, mean 5.8. The
overall mean is 4.9. The wet weight was obtained for this study by
holding the bryOphyte out of the water until it seemed to have stOpped
dripping, then weighing it.

There are several sources of error which might result from this
handful method: 1) insects, especially the more active swimmers,
may be lost as the sample is removed; 2) the handful is variable in
cross section and height,and increase in sample size by weight may
not correspond to an increase in number of insect individuals; 3)
the number of species of insects may be related to the sample size;
4) equal weights of different species of bryOphytes may not be com-

parable in terms of substratum availability for insects.

To provide a rough estimate of the loss of insects during sampling,
a screen was placed immediately downstream from a Fontinalis clump
in Ginseng Run and a handful of moss was collected. The number of
insects reaching the screen was less than .6 per cent of the number
remaining in the collection ( Baetis; 2 simulids); repetition produced
the same results. Furthermore, it is possible that these insects were
dislodged from adjacent mosses in the clump rather than from the sample.
Because of its very loose, open nature, Fontinalis was thought to be
more likely to lose insects during collections than the other bryophytes.
Thus, an estimate of less than 1 per cent loss may not be unreasonable.
Of course, for the surface zone component alone the percentage loss
would be much higher.

By increasing sample size, intuitively one expects to increase
the number of individuals correspondingly (Arrhenius, 1921). Since
many other variables were influencing numbers of insect individuals
and species present in a bryOphyte sample, and the bryophyte samples
themselves were scarcely unifonm in bottom area sampled or bryophyte
surface area or bryophyte weight, it is difficult to show the effect
of sample size on numbers of individuals. But when many samples are
combined and presented graphically, general patterns of stream differ-
ences, abundance differences among the various insects, and associations
between particular bryophytes and particular insects can be inferred.

There is no way to adequately determine the relationship of the
number of insect species to the size of the handful sample, for numbers
of insect species could also be related to basal area, bryOphyte sur-
face area, volume, or a combination of these. If weight is accepted
as proportional to surface area, it serves as one measure of size

relationship. In Fig. 1, we see that there:is the expected increase

 

 

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Number of Grams of Bryophyte

11

in number of insect species with increase in weight (determined from

several samples of one species in one stream on the same date). But

it is possible that basal area, and likewise volume, may reflect upon
the quantity and kind of food passing through the bryOphyte mat, and

thus influence the number of species, as well as the number of indi-

viduals.

That different species of bryophytes might have different weight
to available substratum ratios is a difficult problem to resolve.
Even if we consider weight as a measure of biomass available to a
food chain, we have the problem of camparing quality of food. How-
ever, if weight is assumed to be correlated with bryophyte surface
area, we know many reasons why the relationship might be very complex.
Factors such as tissue density, total plant surface area, leaf to
stem ratio, plant form, and nature of growth (mat, trailing, etc.)
contribute to the bryophyte influence on the insect community. Of
the parameters cited above, weight is easiest to measure, and dry
weight is more replicable. In spite of its limitations, air dry
weight is the measure of the bryOphyte biomass present in each hand-

ful in this study.

Preservation and Sorting

At the time of collection, wet bryOphytes were placed in jars
(usually baby food jars) without addition of stream water, labelled
inside with pencil on cards. and numbered. Upon return
to the lab, 95 per cent alcohol was added to all collections.
Duplicates of the label information and collection data were kept in
a bound notebook, while a separate bound notebook was used to record

stream descriptions and observations in the field.

12

The large bulk of material to be sorted necessitated the use of
technicians who removed the insects from the bryophytes and placed
the insects in 2-dram vials with 70 per cent alcohol for later resort-
ing, identification, and counting.' Collections sorted by the techni-
cians were checked until the technician's effectiveness seemed to be
about the same as my own; later the technician's work was occasionally
spot checked. Nevertheless, I sorted 95 per cent of the material for
this study myself, while my main technician sorted 90 per cent of the
remainder.

All sorting except in the counting chamber was done with a

dissecting microsc0pe at 10X magnification.
Counting Chamber

When their numbers are sufficiently high, Chironomidae require
special sorting techniques. In such instances a counting chamber per-
mits counting at 20X magnification. The chamber consists of a petri
dish with straight lines scratched .25 inches apart on the bottom. By
placing the insects in the lid and anchoring the lined bottom inside
the lid with paper clips, one can move the petri dish back and forth
for counting without disturbing the insects' positions. Comparison
between a hand-count and chamber count of the same sample showed less
than five per cent lower count by chamber counting.

Even the latter method is more time-consuming than seems justified
because the Chironomidae are nearly always highest in abundance among
the insects. Because insects settle toward the bottom of the jar,
representative subsampling is impossible. Consequently, the alter-

native procedure for the extremely abundant samples is to sample a

13
few chironomids, but leave the bulk of them on the bryophyte without
counting them. This reduces sorting time by 50 to 70 per cent. When
chironomids are low in numbers or are not obviously the most abundant,
- they are all removed from the bryOphyte and aaunted with a hand counter,

as are all other insects.
Stream Data

Because so many streams are included in this study, and only a
few sampled throughout the year, no specific attempt was made to link
stream chemical characteristics to the bryophyte-insect associations.
U. S. Weather Bureau Climatological Data (Climate and Man, 1941) pro-
vide the temperatures and growing seasons (Plates 2-6), while U. 3.
Geological Survey maps provide the elevations and rock types, except
where other references are cited. It is possible to obtain an average
elevation gradient for the stream by using maps to measure distance
from.most distant source to the collection site and dividing this into
the difference in elevations. Stream order is determined by the number

of tributary junctions (Leopold, 1962).

10.
ll.
12.

13.

14

PLATE 1. MAP OF STREAM LOCATIONS
Saw Creek and Dingman's Creek
Mud Run and tributaries
PohOpoco Creek
Elk Creek
Pidcock Creek
Piney Creek
Gramlich Run
Sideling Hill Creek tributary
Youghiogheny River system
Little Bennett Creek and Seneca Creek tributaries
Sinking Creek and Johns Creek
Mountain Lake tributary to Sinking Creek

Rock Castle Creek and Goose Creek

15

PLATE 1

 

 

 

 

 

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RESULTS AND DISCUSSION

In the upper reaches of the Appalachian Mountains, cool, rapid
streams find their origin where springs, surface runoff, and ground
water insure a continuous flow throughout the year. Because of
these factors, coupled with the shading effect of deciduous forest
trees, the area is an ideal one for stream bryophytes. ‘Yet there is
sufficient variation in stream size, gradient, chemistry, and substrate
to permit a comparative study. Plates 2-6 show USGS climatological
data. Table I lists the species of bryOphytes, while Table II gives
a tabular summary of some of the physical data and bryOphytes for
each stream.

The streams contributing to this study are in Pennsylvania,
Maryland, and Virginia. For purposes of ecological comparison, these
can be grouped in many ways: geography, stream order, elevation,
climate, rock type, chemistry, dominant organisms. Because this study
is concerned with bryophyte communities, the streams are herein grouped
by dominant bryophyte (used here as the bryOphyte composing over 50
per cent of the bryophyte cover as observed at the collecting sites).
Thus, four bryophytes appear to be dominant bryOphytes in the streams
chosen: Fontinalis dalecarlica, acapania undulata, ficiaromiug legcurii,
and the flygroamblystegium fluviagilg group. The latter is a group of
species in which E, fluviatile is usually present with one or more
other bryophytes, and the insect species composition for these is
quite similar, as well as the streams they occupy. Furthermore, the
bryophytes of this group all form a similar mat, varying from rough
to smooth, but all compact.

21

22
TABLE I
TAXONOMIC LIST OF BRYOPHYTES
Mosses*
Amblystegium varium (Hedw.)

Brachythecium plumosum (Hedw.) BSG

 

Brachythecium rivulare BSG
Bryhnia novae-angliae (Sull. & Lesq.) Grout

Eurynchium riparioides (Hedw.) Rich.

 

Fissidens bryoides Hedw.

 

Fissidens cf. minutulus Sull.

Fontinalis antipyretica Hedw. var. gigantea (Sull.) Sull.
Fontinalis dalecarlica Schimp. 25 B80

Fontinalis flaccida Ren. & Card.

Grimmia alpicola Hedw. var. rivularis (Brid.) Broth.
Hygroamblystegium fluviatile (Hedw.) Loeske

fiygroamblystegium fluviatile (Hedw.) Loeske var. orthocladon
(P. Beauv.) Crum, Steere & Anderson

wroamblystem £9335 (Hedw.) Jenn.
Hygrohypnum luridum (Hedw.) Jenn.
Hygrohypnum ocraceum.(Turner) Loeske

Leskea cf. gracilescens Hedw.

Sciaromium lescurii (Sull.) Broth.
gematgphyllum carolinianum (C. Mull.) Britt.
SematOphyllum marylandicum (C. Mull.) Britt.

Thuidium delicatulum (Hedw.) BSG

 

*Names and authors are according to Crum, Steere, and Anderson, 1965.

23

Liverworts
Conocephalum conicum (L.) Dum.
Frullania sp.
Marsupella gphacelata (Gieseke) Dum.
Riccardia sinuata (Dicks.) Trev.
Scapania nemorosa (L.) Dum.
Scapania undulata (L.) Dum.

Plagiochila?

 

24

 

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26

The method of grouping bryophytes by form or type of mat is not
unique. European bryologists (Gimingham and Birse, 1957) have
attempted this procedure by describing life form, primarily based on
terrestrial bryOphytes. By their system, four life forms applicable
herein have been described: rough mats (Mr)--Brachythecium rivulare;
smooth mats (Ms)--§urynchium riparioides; short turfs (t)-dFissidens
bryoides; tall turfs (Te)--Plagiochila asplenioides.

Because Brachythecium rivulare and ggrynchium riparioides should
be separated by the above system (Gimingham and Birse, 1957), they
present a problem in this study, wherein they have been included in
the same group. Jovet (1932), in his work with French streams, com-
ments that the leaves of g, rivulare are a beautiful, fresh bronze,
and their arrangement recalls that of EurynchIQQDripagigides, whereas
the singular Brachythacium rivulare on the rocks of la Petite Cascade
has julaceous branches simulating g, riparioides, while other leaves
resemble Amblystegium rigarium. Because of this variability of g,
riparioides and the others includedeith it in this study, I believe
they might more practically be included in one group as "mats." How-
ever, I have separated one other group (§ciaromium lescurii) because
it can generally be named as a dominant rather than as a co-dominant.
§ciaromium forms a more open mat and was detritus-covered in shallow,
slow streams included in this study; its insect community appears
sufficiently different to warrant a separate consideration.

In this study, the "short turf" (Eissidens bryoides) occurs at
the bases of other plants, so that it does not warrant its own desig-
nation as a stream type. The “tall turf," considered by Gimingham

and Birse (1957) to resemble Plagiochila asplenioides, could include

27

the leafy liverworts of this study, especially Scapania undulata.
These turfs form an open mat more like a loose sponge than a true
mat.

The truly aquatic Fontinalis remains to be classified. This
moss does not form a compact mat, but rather dangles, or nearly floats,
in the moving water. I have given this type the singular designation
of "streamer," a term that could only apply to a long, dangling aquatic.

When Gimingham and Birse discuss life forms of their bryOphytes,
they propose that certain forms relate to the environmental conditions.
For example, Eurynchium riparioides (Ms) grows lowest on the rock
(where it is most moist), has a closely appressed, prostrate shoot
system, and is thus able to withstand the scouring effect of rushing
water while the moss is submersed during most of the year; a smooth
mat offers the least resistance to water and is correlated with plants,
such as g. riparioides, which adhere most strongly. Further support
of this idea is evidenced by Jovet's (1932) work, where he found g.
riparioides to be rheOphilic, occurring in rapid chutes, especially
the spillways of ponds, while he termed Brachythecium rivulare par-

tially rheophilic, replacing g, riparioides in the dripping, but not

 

torrential, part of a waterfall. However, Jovet (1932) and Watson
(1919) both point out the ubiquitous nature of g, riparioides by its
occurrence in very slow as well as very fast waters.

The ecological amplitude of such bryOphytes as Eurynchium lends
criticism to schemes like that of Lorentz or Gams (iEDVerdoorn, 1932),
who classify by habitat. Lorentz divides his Aquaticae into:
Paludosae (marshy), Pontanae (flowing water), Irroratae (moist areas--

the dew plants), Natantes (floating and submerged), and Pluctuantes

28

(floating). Thus, Eugynchium would fall in several of these: Palu-
dosae, Fontanae, Fluctuantes, and possibly Irroratae.

Game (1932) uses Nereidia for constantly submerged and Amphi-
nereidia for amphibious forms. By his system, the Nereidia would
encompass the Fontinalaceae and the Hygrohypnion federation: flyggg-
hypnum spp., Eurynchium riparioides, and maritime species of Sciaromium.
Meanwhile, Eygrohypnum palustre is Amphinereidic. Cams, in 1953,
warns of the use of only a generic designation for a federation "Les
noms des unions, federations, etc., deraient etre choisis de faqon a
exclure toute confusions. Des designations trop abregies sont a
eviter, p. ex...Rhynchostegion (pour Rhynchostegietum riparioides a
Platyhypnetum rusciformis, non pas Rhynchostegietum muralis)..."
Because Rhynchostegigg,riparioides is treated in this paper as £2525-
ghigg_riparioides, I would further prefer not to use generic names
alone until our system of classification is more stable. Thus, instead
of a genus, Gama discusses the gydro-Martinelligg by designating the
Scapanietum undulatae formation by species, as I have done in this

paper.
Fontinalis dalecarlica Streams (Fontinaletum dalecarlicae)

Fontinalis dalecarlica, the most nearly ubiquitous of the bryo-
phytes, occurs in first, second, and third order streams, in 0-3 foot
streams to the widest included in the study (40-60 feet wide), in
depths of about 5 inches to depths of 2-3 feet, in shaded or sunny
areas, in rapids and falls or in pools. And the Fontinalis dominant
streams are the second greatest in number, comprising five of the 28

streams; in addition to these there are several Scapania dominant

29

streams in which dense Fontinalis beds occur. However, the latter
will all be treated as Scapania streams.

Due to the ubiquitous nature of E, dalecarlica, it is impossible
to characterize its stream type. Rather, we can state that these
are streams with large, flowing mats of "streamers." Perhaps the only
characterization one can suggest is the ability of Fontinalis to
occupy the larger, deeper streams, such as Mud Run and Muddy Creek,
where the other bryophytes occur only on the edge or near the water-
air interface on emergent rocks. Moreover, it does not occur in the
narrow, shallow streams where it would surely be out of water part of
the year. Studies by Irmscher (1912) show that other species of Egg:
tinalis (E, antipyretica and g, sguamosa) die after one week of air
drying.

With its long, dangling branches, 2, dalecarlica gives more the
appearance of higher plants than shy other bryophyte studied. Because
its mat is loose and flexible, large insects may occur here, although
they never occur in mats of other bryophytes of the same stream. For
example, only in Fontinalis could I find third year Pteronarcys naiads
--the one-inch long stonefly. But the large plant affords less pro-
tection from the current because turbulence produces a whipping motion
of the plants, preventing the existence within the bryophyte stand
of a water mass that is unaffected by stream turbulence. It also
suggests less protection from predators, assuming that these larger
animals can get into the depths of a Fontinalis clumpmore easily than
into the closely-meshed mats of other bryophytes.

Fontinalis often harbors a diverse insect fauna, but in other

collections may have very few individuals and species. It appears

30
to provide a satisfactory home for Chironomidae, where these tiny
midges often nestle in the axil between the stem and leaf or attach
a tubular sand case to the stem or leaf backs. Like the chironomids,
simulid larvae such as the Simulium tuberogum complex attach them-
selves in leaf axils with only their heads visible. It is possible
that these positions provide safety from drif“ting while enabling the
algal-feeding Simulium (Cummins, pers. comm.) to catch what passes
by. Frequently the branching respiratory filaments of simulid pupae
(Prosimulium hirtipes complex) extend from the axial net cases (Plate
7), or at other times the larvae have used a leaf with a thin net to
make the pupal case, replacing the usual stiff case used on the stems
of these plants. In streams like Toliver and Pohopoco, the caddis
Diplectrona modesta extends nets from branch to branch, catching the
passing detritus and plankton, while the larva reposes in a net and
sand case near the base of the plant. Apparently the most likely
moss for finding micro-caddis (HydrOptilidae), Fontinalis frequently
is decorated with the attached cases of Hydrogtila and ngethira
(Plate 7), while a brachycentrid larva (new genus) not only attaches
the anterior end of its case to the plant, but also uses Fontinalis
leaves, with other bryophyte leaves, in case building (Plate 7).
Muddy Creek
Plate 10, Fig. 1. adapted from USGS Sang Run quadrangle

*Garrett Co., Md.: Swallow Falls State Park, above falls
#(1965z5-4, 7-9, 8-25, 12-25; 1966:3-22, 6-11)

Muddy Creek is one of several Youghiogheny tributaries studied.
The others are Neds Run, Piney Creek, Ginseng Run, Hoyes Run, Toliver
*Location of stream

#(year:month-day)

31

 

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32

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33

Run, and Deep Creek tributary. Among all the streams studied, the
Youghiogheny tributaries flow through the area with the most snowfall
and the coldest, wettest weather in Maryland, where temperatures
seldom rise above freezing from December to February (Dept. Geol.,
Mines, & Water Res., 1954). These extremes contribute to a unique
segment of Maryland flora in which northern hardwoods contrast with
white pines and hemlock stands. Garrett County is a rolling plateau
interrupted by deeply cut stream valleys and northeastwardly oriented
ridges which parallel the northeastward trend of the anticlines and
synclines. The northward flowing Youghiogheny and Casselman Rivers
drain the Ohio River basin in the county, while the Savage River flows
south to the Potomac River., Contributing to local t0pography are
tributaries which gouge out east-west oriented gaps in the ridges.

It is these tributaries, located in elevations of 2200 feet to 2600
feet, bounding over boulders or sliding over shales, which provide
the year-round collections of this study.

As the name implies, the headwater region of Muddy Creek is "muddy"
in color, but it is clear at the collection site. Originating in
Cranesville Pine Swamp, Muddy Creek flows over Pocono, Greenbrier,
and Mauch Chunk shales before reaching the Pottsville conglomerate
of the main collection site. At the collection site in the 100 feet
above the falls, Muddy Creek is the widest stream studied. Like Mud
Run (also a Fontinalis stream), it is in open sun and is bordered by
rhododendron and virgin hemlock forest. In this area, the raging
torrents of snowmelt waters slow to peaceful moderation in summer,
passing over solid bedrock before plunging over thirty-foot falls in

Swallow Falls Park. It is in this 100 feet above the falls that

34

Eonginalis dalecarlica forms large, streaming clumps. Although

located on the bedrock just above the falls, Fontinalis occurs on

the sides of submersed boulders less than 100 feet farther upstream.
No other bryOphyte could be found in the collecting area above the
falls except during high water periods. But on August 25, 1965, the.
water had receded sufficiently to collect gygrohypnum luridum from the
falls.

Below the falls .4 miles, Muddy Creek joins the Youghiogheny
River. This lower, sunny and deeper section of the stream is depau-
perate of bryophytes, but one boulder supported Fontinalis and Sci;
aromium lescurii at about 100 feet above the river junction.
reuse
Plate 10, Fig. 1. adapted from USGS Sang Run quadrangle

Garrett Co., Md.: east of Cranesville Pine Swamp by culvert
(1965:5-4)

Neds Run drains the western side of Piney Mountain to join
Muddy Creek in Cranesville Swamp. At its source it drains the
soft Mauch Chunk shales, but it crosses the Greenbrier limestone at
the collection site .6 miles below. Since it is narrow and flows
through tall, mixed hardwood forest, where its borders are lined with

Caltha palustris L., little direct sunlight reaches its surface.

 

Although this stream exhibits primarily sandy bottom, a few small
boulders occur near its mouth, and here streamers of Fontinalis

dalecarlica undulate in the moderately flowing water.

 

Mud Run
Plate 8, Fig. 1. adapted from USGS Stoddardsville quadrangle

Carbon Co., Pa.; Hickory Run State Park, downstream from bridge, Pa. 903
(1965:7-29)

35

Mud Run finds its origin in Pimple Hill of the northern Pocono
Mountains and gathers waters from such first order tributaries as
Swamp Run, Laurel Run, and Bach Run, all of which drain Lake Mountain
on the western side of the Pocono Mountains. Flowing westward, Mud
Run reaches the Lehigh River, then the Delaware to the Atlantic.
Shaded by a forest of northern hardwoods and hemlocks with a rhodo-
dendron understory, these streams all flow over Mississippian Pocono
sandstones and exhibit rocky bottoms at their collection sites.
However, the physical appearance of the tributaries is quite different
from that of Mud Run.

In general, the forest-shaded Mud Run drains the same soils as
Swamp Run, but from the source to the collection site, it traverses
only Pocono sandstones. Mud Run is one of the largest streams studied.
Because of its width, full sunlight reaches much of the stream surface
during part of the day. Perhaps due to its combination of greater
depth, clear, cool water, and sunlight, the stream has the most lush
growth of Eontinalig of any stream in the study. Here, the moderately
fast waters flow over the Fontinalis-covered rocky bottom while emer-
gent boulders frequently support growths of Scapania undulata just
beneath the water surface.

Swamp Run,,a Mud Run tributary
Plate 8, Fig. 1. adapted from USGS Stoddardsville quadrangle

Carbon Co., Pa.: Hickory Run State Park, south of Pa. 534
(1965:7-29)

Through its upper reaches on Lake Mountain, Swamp Run drains peat
and muck of the Klinesville soils, while near the collection area
its moderately flowing waters erode Papakating silty clay loam, a wet,

poorly drained soil with mucky surface formed under hemlock, spruce,

36
rhododendron, and Sphagnum (Carbon Co. Soil Survey, 1962). In a
sunny stretch streaming glyceria fluitans (L.) R. Br. nearly chokes
the stream above the collection site, but at the edge of a shaded,
swampy section, bordered with Veratrum viride Ait., bryOphytes are the
only vegetation. Whereas Fontinalis occurs on the bottom rocks and
in swirling waters of rapid gradient changes, leafy liverworts occur
on submersed edges of emergent rocks. Because of the preponderance
of the liverwort Scapania undulata in Hoch Run and Laurel Run, these
streams are treated with the Scapania group. But Swamp Run, where
Fontinalis dalecarlica spots the rocky bottom, has only sparse growths
of the leafy liverworts Plagiochila? and Frullania sp., while the
thallose liverwort Riccardia cf. sinuata is an occasional constituent.
Accompanying this unusual combination of liverworts is the aquatic

lichen Hydrothyria venosa Russell, while Fissidens bryoides nestles

 

among the bases. Apparently because the stream is rugged, moderately
fast, and narrow, its heavy load of ice, water, or eroded materials
causes Fontinalis to lose its leaves.

Pohopoco Creek tributary

Plate 8, Fig. 3. adapted from USGS Windgap quadrangle

Monroe Co., Pa.: parallels west of Pa. 115, 2 mi. north of Effort
(l965):7-29)

Pohopoco Creek drains the red sandstones of the Catskill forma-
tion in the southern Pocono Mountains through the stream's multiple
sources before joining the waters of Mud Run in the Lehigh River.
Shaded by northern hardwoods and rhododendrons, the stream slides
around emergent boulders with their clumps of Sciaromium lescurii and

their trailing Fontinalis in slow waters, while occasional polsters

37

of Eurynchium riparioides occur in the rapids on small rocks. Nestled
at the bases of these bryOphytes are tiny plants of Fissidens bryoides.
Evidencing a rough life, the Fontinalis here is stripped of most of
its leaves and displays several old capsules. Watson (1963) states

that capsules of E, antipyretica may be found more often on plants

 

exposed to air in dry weather. However, Elssmann (1923) states that
plants kept in the laboratory produced capsules without being out of
the water in whole or in part. He could not say if the same is true

in nature.

Hygroamblystegium fluviatile Streams

(Hygroamblystegietum fluviatilae)

By far the most frequent species studied, g, fluviatile appears
to avoid segregation. Because of its habit of occurring in streams
with other bryophytes on the same or different rocks, it can be con-
sidered as the dominant indicator of a group which includes one or
more other species: Eurynchium riparioides, Hygrgamblystegium tenax,
Amblystegium varium, Brachythecium plumosum, Brachythecium rivulage,

and less commonly Grimmia alpicola var. rivularis and Leskea cf. gra-

 

cilescens. In addition to these, the ubiquitous Fontinalis dalecarlica
frequently appears. If the latter is dominant, the stream is treated
as a Fontinalis stream. When Scapania undulata appears in small
quantities in these streams, the categorization is more difficult.
While the gygroamblystegium group frequently has a fauna similar to
that of Fontinalis in the same stream, Scapania undulata has the most

' unique fauna of the major bryophytes studied. Consequently, streams

containing both Scapania and Hygroamblystegium will be discussed here

 

38

if the latter is dominant, but of necessity will again be mentioned

in the Scapania group. In his study of Brachythecium plumosum, Gaume

 

(1928) mentions that B, plumosum is characteristic of the Rhacomitrium
aciculare and Scapania undulata group. But this is not true of the
streams in this study.

While Hygroamblystegium fluviatile and its group members (not

 

Fontinalis and Scapania) do not all look the same, they form similar
relatively dense mats on rocks. Apparently their habitat requirements
are about the same, as evidenced by their frequent coficcurrence in the
same mat. So it can be expected that their insect fauna may be sim-
ilar unless the particular species of bryophyte exerts an influence

,on the insects. The data indicate that this influence is not apparent;
thus, grouping seems justifiable. Another justification for grouping
is the extreme variability of several of the moss Species so that their
gametOphyte characters overlap. For example, g, fluviatile, E, 53235,
and Amblystegium varium are difficult to separate; Brachythecium
plumosum, B. rivulare, and Eurynchium riparioides overlap.

If we consider all the streams treated here as Hygroamblystegium
streams, we find that they are all narrow and shallow, usually less
than eight feet wide and only inches deep (Table II). On the other
hand, this does not imply that these bryophytes occur here exclusively.
On the contrary, the 60-foot wide downstream area of Muddy Creek has
fl. fluviatile, but not as a dominant. Rather, it occurs in the larger
streams at the water surface on boulders. Thus all of the occurrences
noted are at the water-air interface. Furthermore, any attempt to
correlate these bryOphytes with stream order proves futile because

they occur in first, second, and third order streams. However, this

39

compares with the observation of Harrel (1966) whose third order
stream was more similar to the sixth order stream than another of the
third order. In the present study, it appears that the Hygroambly-
stggigm_group occupies first and second order streams and those third
order ones which have areas similar to the lower orders.

While comments on H, fluviatile hold true for stream type of
the other mosses of this association, further comment is needed on
specific occupance of the streams. For instance, H, fluviatile and
Grimmia alpicola var. rivularis were not found on vertical rocks of
waterfalls, while Brachythecium plumosum and B, rivulare, along with
their counterpart, Eurynchium :iparioides, were in rapid falls or on
midstream rocks, as already reported in 1919 by Watson. In both of
these habitats one might find the lichen Dermatocarpon aquaticum (Weis.)
Zahlbr. intermingled with the mosses.

Throughout the year one can find clumps of stems with naked costae
due to the scouring of g, fluviatile, while g, riparioides shows the
effects by fraying. It is usual to find the lower leaves torn away
on these aquatic bryophytes (Watson, 1919).

Among the branches and leaves of flyggoamblystegium and its consorts,
the tiny chironomid larvae may hide, but are less frequent in the leaf
axils than in those of Fontinalis, probably because the leaves are
smaller and do not hide the larvae as well. Even more rarely, a larva
incorporates a Eygroamblystegium leaf into a loose case. Occasionally
one can find a branch of the moss attached to the vegetable case of
the caddis Micrasema (Brachycentridae), while the tiniest of these
larvae attach sand cases weakly to the leaves and stems. Older larvae,
prepupae, and pupae frequently attach the front, back, or both ends of

the case to the moss. Another brachycentrid (new genus) occurs among

40

H, fluviatile and Eurynchium riparioides mats, utilizing these mater-

 

ials for its case. These cases are especially interesting because

the larva may incorporate only the costae of Hygroamblystegium fluvi-

ggilg, leaving dangling ends of costae, or it may make a smooth case

with the lamina of the leaf included.

Sideliggiflill Creek tributary

Plate 10, Fig. 4. adapted from Map of Allegany Co., Md. G. S. 1905

Allegany Co., Md.: parallel to U. S. 40, .6 mi. west of Sideling Hill
Creek

(1965:3-29)

While its own tributaries exhibit rapid drops, this third order
tributary is nearly level and only moderately fast as it courses along
the eastern side of the scrub pine-hardwood Town Hill Mountain. Where-
as it originates in Hampshire shales, it traverses Jennings shales
through open sun before it enters Sideling Hill Creek to be carried
to the Potomac River. In the collection area it cuts a shallow chan-
nel over a bottom of sand, pebbles, and soil. Along the banks, rocks
covered with Brachythecium plumosum are submerged by high waters of
spring. The paucity of insects in these spring collections is proba-
bly due to the temporary nature of the habitat.

Sinking:Creek

Plate 12, Fig. 2. adapted from A. S. Caster. 1958. Mt. Lake Biol.
Station & Vicinity

Craig Co., Va.: Newport Recreation Center, Newport.
(1965:3-18)

With tributaries arising at elevations of 24-2500 feet in Cooper
Ridge Dolomite limestone springs on the northern side of Sinking Creek
Mountain, Sinking Creek itself meanders through the valley below and

finally enters the New River, which empties into the Ohio by way of

41

the Kanawha River. Its bottom rocks are often carpeted with thin mats

of Amblystegium varium, Hygroamblystegium fluviatile, and Brachythecium

 

 

plumosum, but more commonly with Hygroamblystegium tenax. Along the

 

banks Conocephalum conicum, growing on tree roots, finds itself in the

 

high springtime waters. According to Gimingham and Birse, 1957, Gono-

cephalum conicum will succumb in moisture; it also adheres weakly and

 

would probably be eroded by running water. But this thallose mat was
entangled by roots of grasses and other vascular plants, so it appeared
relatively safe from erosion.

Mountain Lake tributary to Sinking:Creek

Plate 12, Fig. 3. adapted from A. S. Caster. 1958. Mt. Lake Biol.
Station & Vicinity

Giles Co., Va.: north and west of road to Mountain Lake on Salt Pond
Mountain, east of U. S. 460
(1965:3-18)

This first order Mountain Lake Park tributary, one of the smallest
streams studied, is intermittent in its upper reaches of Salt Pond
Mountain (Caster, 1958). In March it trickled through the collecting
site, where moss-covered rocks were wet primarily because of spray
as the water coursed the rugged channel. On the north it is shaded
by a mixed hardwood forest, but it embraces the afternoon sun from
the south where the road to the biological station replaces the trees.
On the rocks in the splash and submersed are mats of Brachythecium
rivulare, Exggoamblystegium fluviatile, and H, fluviatile var. 25522-
cladon. However, attesting to the temporary nature of the stream, the

insects are few, consisting primarily of Diptera, especially chiron-

omids and Pericoma (Psychodidae) larvae.

42

Rock Castle Creek
Plate 12, Fig. 1. adapted from USGS Floyd quadrangle

Patrick Co., Va.: Va. 710, lst bridge, 2 mi. north of 4ON; Woolwine
(1965:7-21)

Rock Castle Creek originates in southern Sugarloaf and Rocky Knob
Mountain, coursing through a mixed hardwood forest to eventually reach
the Sycamore River. FrOm there this third order stream travels Smith
River, Dans River, Jackson River, and Pee Dee River before entering
the Atlantic at Winyan Bay. In the collection area, most of the slip-
pery, black boulders are carpeted with the riverweed Podostemum

ceratgphyllum Michx. However, a mat of Eurynchium riparioides covers

 

 

rocks in the shallow water of rapids.
Johns Creek tributa;y_

Plate 12, Fig. 5. adapted from A. S. Caster. 1958. Mt. Lake Biol.
Station & Vicinity

Graig Co., Va.: just before Giles Co. on rt. 632
(1965:7-14)

This first order tributary originates in Giles County and flows
northeastward to Craig Creek, draining the north side of Johns Creek
Mountain, and finally reaches the Chesapeake Bay through the James
River system. In a series of step falls, where the stream bounds over
limestone rock (Woodson, 1957), dense mats of Hygroamblystegium fluvi-
gtilg grow. Only on some of the larger submersed boulders can one
find the streamers of Fontinalis dalecarlica.

Goose Creek
Plate 12, Fig. 4. adapted from USGS Elliston quadrangle

Floyd Co., Va.: near and of road south of Piedmont
(1965:3-22)

43

The third order Goose Creek cuts through the Blue Ridge Mountain,
leaving large boulders in the stream bed before emptying into the
Roanoke River. This hemlock-hardwood shaded stream shears across the
thin mats of Hygroamblystegium fluviatile which cling to the midstream
boulders, while other plants dangle into the water from the stream-
bank. While g, fluviatile occurs both on the edge and in the middle,
Euryncbium riparioides, Brachythecium rivulare, and Brachythecium
plumosum occur in only one collection each out of 13.
Hgyes Run and Ginsengikun (Sang Run)
Plate 10, Fig. 1. adapted from USGS Sang Run quadrangle
Garrett Co., Md.: Hoyes--east of Hoyes Run (town); Ginseng-~east of

Sang Run (town), 4.6 mi. west of 219 at McHenry; and near rt. 21

(Hgyggf.l965:8-25; 1966:3-22, 6-11; Ginseng, 1965:5-4, 7-9)

Because of their proximity and similarity, the second order Hoyes
I Run and Ginseng Run (Sang Run) warrant simultaneous consideration.
Both meander in and out of mixed hardwood forests and pastures; both
have moderately fast flow; both are narrow and shallow in the collec-
tion area. Whereas both their collection sites are located at 2040
feet, Hoyes Run drains northern Marsh Hill, while Ginseng Run drains
southern Ginseng Hill. Although their mouths to the Youghiogheny
River are only 2.5 miles apart, Hoyes Run empties across Mauch Chunk
shales while Ginseng finishes in Greenbrier limestone. Furthermore,
Hoyes originates in Pocono sandstone while Ginseng originates in the
Catskill formation, but, at the upstream collection site, Ginseng Run
crosses the Pocono sandstones.

At its collecting site less than a mile from the mouth, a shoulder

of shale provides the substratum for Ginseng Run bryOphytes, where

Hygroamblystegium fluviatile and Eurynchium riparioides grow intermin-

 

 

44

gled. On one mud-covered rock a few branches of Thuidium delicatulum
and Scapania undulata were mixed with these. In the Open water,
branches of Fontinalig_da1ecarligg_extend from the submersed rocks.
During July, one can find vigorous growths of Fissidens bryoides at
the bases of the other bryOphytes. This usually minute moss aroused
the curiosity of Dr. R. A. Pursell (pers. comm.) because of its large,
robust form.

At theupstream site of Ginseng, only Eurynchium riparioides and
Hygroamblystegium fluviatile were collected, but no collections were
taken here in July when the other bryOphytes were found down stream.

Unfortunately, the Ginseng Run collection dates did not coincide
with those of Hoyes Run. On three dates of sampling, Hoyes Run dis-
played only Hygroamblystegium fluviatile and Eurynchium giparioides
growing in large dense carpets on the submersed sloping bedrock, with
a tiny Fissidens, possibly F, pusillus, occurring rarely among their
bases.

Piggy Creek tributary

Plate 10, Fig. 2. adapted from USGS Frostburg and Avilton quadrangles

Garrett Co., Md.: tributary entering south side of Frostburg reservoir,
100 feet from reservoir by bridge of dirt road; reached from U. S. 40

(1965:5-5)

This first order tributary is located in the northeastern part
of Garrett County and flows northward into the Frostburg Reservoir
and finally to the Youghiogheny River through Casselman River in
Pennsylvania. It originates in Hampshire rock, and flows over Jennings
shales and sandstone in the collecting area, where its slow-moving

waters result in a heavy deposit of sand and mud on the few bryOphytes:

Hygroamblystegium fluviatile, Eurynchium riparioides, and Fontinalis

45
dalecarlica, with Fissidens bryoides at its base.
Deep Creek tributagy
Plate 10, Fig. 1. adapted from USGS Garrett Co. Atlas
Garrett Co., Md.: south of Swallow Falls Road by private mailbox
356, just above Deep Creek Lake
(1965:5-4; 1966:3-22, 6-11)

This first order tributary soon dumps into the man-made Deep
Creek Lake that empties into Deep Creek, which finally joins the waters
of the other Garrett County streams in the Youghiogheny River. My
observations of the stream in 1965 indicate that it is intermittent
at the collection site in dry years. Its moderate to rapid flow in
early May gave way to a highly vegetated alley with an irregular trickle
and small pools during the summer dry period. In March, 1966, it was
again active. Characterized by large boulders, the stream stumbles
through numerous riffles, rapids, and waterfalls in the less than
thirty-foot long collection area. This portion of the stream is rapid
and clear in March, probably swollen due to snowmelt runoff. Although
situated at the collection site on soft, nearly neutral Mauch Chunk
shales, and draining primarily the Greenbrier limestone formation in
its upper reaches, this tributary is shaded by the acidOphilous rhodo-
dendron and a hemlock-hardwood forest. Carpeting its banks and emer-
gent rocks are numerous species of bryOphytes. Whereas streamers of

Fontinalis dalecarlica extend from the edges of submersed rocks,

 

Scapania undulata, Eu_rynchium riparioides, Hygrohypnum luridum, aid Hygro—

amblystegium fluviatile form clumps or polsters on basketball-sized

 

rocks, especially where these rocks cause a change in the stream level.
At the base of any of the mosses, one can find the tiny Fissidens br -

oides. In March still other bryophytes can be found on rocks in the

46

spray caused by small waterfalls. Because of such a variety of bryo-
phytes, this stream best fits with the Hygroamblystegium fluviatile
group.

Gramlich Run tributary,

Plate 10, Fig. 3. adapted from USGS Cumberland quadrangle

Allegany Co» Md.: parallels Gramlich Road, LaVale; east of U. S. 40
(1964:12-25; 1965:3-28, 5-3, 6-20, 7-10, 8-24, 12-24; 1966:3-23)

Possessing the third steepest elevation gradient of the 28 streams
--400 feet per mile, the Gramlich tributary tumbles from its source in
Piney Mountain over Pottsville conglomerate, Mauch Chunk, Greenbrier,
Pocono, and Hamilton formations (Allegany Co. Atlas, 1900), before
reaching the Jennings shales at the collecting site. Later it empties
into Braddock Run, to Wills Creek, to the Potomac River.

Waterfalls, upturned shale ledges, small pools, and boulders
provide a variety of possibilities for growth of bryophytes in this
second order Gramlich Run tributary (so named in this study because
it parallels Gramlich Road). This bubbling stream provided the only
collection of Fontinalis flaccida, which came from a clump of leaf
drift in a small pool, while streamers of B, dalecarlica extend from
long, leping submersed bedrock. In the 3-foot waterfall, Eurynchium
riparioides, Brachythecium rivulare, and Brachythecium plumosum cling
to the vertical rocks, while B. riparioides forms mounds at the base
of the falls as well. In summer, the bryophytes are scoured away from
the main channel of cascading water, but on each side of this channel
a film of water streams down the rocks behind such mosses as B. ripari-
EEQEE and B. rivulare, keeping them constantly moist and influencing
their insect fauna. Many of the same mosses also occur on the boulders

and upturned shale ledges in the stream: B, rivulare, Grimmia alpicola

47

var. rivularis, Hygroamblystegium fluviatile, Brachythecigm plumosum,
and Eurynchiumggiparioides with Fissidens bryoideg at its base.

Because the stream passes through a good timber stand of northern
hardwoods, it may be influenced by tannic acids resulting from up-
stream logging. However, the logging road diverges from the tribu-
tary about a mile upstream, and all bgging to date occurs above this
point.
Little Bennett Creek
. Plate 11, Fig. 2. adapted from USGS Damascus quadrangle

Montgomery Co., Md.: west of Oak Drive (old rt. 27), Damascus
(1964:6-3; 1965:3-26, 6-29, 8-20)

This first order tributary cuts its way across the Ijamsville
phyllite through a mixed hardwood scrub pine forest. Here, matted
bryophytes carpet the irregular surfaces of the scattered milky quartz
rocks. Like the streams near Watkins Road, this one diaplays such
bryOphytes as Hygroamblystegium fluviatile, Brachythecium rivulare,

Sciaromium lescurii, and Thuidium delicatulum. The fern moss, Thu-

 

nggm, is from intermittent waters of a spring side channel, while
the others can be found in the main stream as well. Inversion of the
rocks is suggested where these mosses occur on the undersides of the
rocks in the swampy intermittent region.

Although this gurgling Piedmont stream was clear at the outset,
a logging operation soon curtailed the study and at present a log road
fords the stream in the collection area.
Seneca Creek tributaries
Plate 11, Fig. 1. adapted from USGS Gaithersburg quadrangle
Montgomery Co., Md.: south of Watkins Road (Md. 604), east of Md. 27;

by residence of Burroughs and M. A. Bell
(1965:3-27)

48

The smallest streams of the study are these two on the south side
of Md. 604. Because they have no name, they shall be referred to as
Burroughs and Bell, after the names of the nearest property owners.

The first order Burroughs stream flows through a cow pasture,
thus being exposed to an obvious source of pollution. However, the
water at the collection site is clear and is upstream from the
fenced pasture area, so it is probably not polluted. While the owner
reports that the stream always has water in it, this flow is not fast,
even in spring. On the other hand, the Bell stream is definitely in-
termittent, as determined by Mrs. Ellis G. Glime and the Bell's (pers.
comm.) in October, 1967. Originating just below Watkins Road, the
stream barely moves to cross the nearly level open woods to the col-
lecting site less than 100 feet from the road. Apparently this first
order stream is the result of spring rain runoff and snowmelt.

Both streams traverse the Ijamsville phyllite formation under a
canopy of mixed hardwoods. Their sandy bottoms have scattered rocks
which support occasional small moss clumps including Hygroamblystegium
fluviatile in the Burroughs stream and Bryhnia novae-angliae on wood
in the Bell stream. The latter is included here for convenience as
this is the only collection of this hydrOphilic species and it is most

similar to the Hygroamblystegium group. By the classification of

 

Gimingham and Birse, 1957, it is a rough mat.
Elk Creek

Plate 8, Fig. 4. adapted from USGS Centre Hall quadrangle and Lesley,
1885. Centre Co.

Centre Co., Pa.: parallels Pa. 445 just north of Milheim
(1965:7-28)

49

Elk Creek, a warm, sunny, moderately fast stream, cuts a channel
varying in depth from inches in mossy areas to nearly two feet where
large beds of Ranunculus longirostris Godr. grow. Its third order
collection site is in an area of Hudson and Utica formation and gray
Oneida conglomerate sandstone. Arising in Brush Valley slate and
sandstone and draining Brush, Big, and Nittany Mountains, it courses
to the Milheim Valley through an area of limestone. It may be the
limestone which causes the cloudy appearance of the collecting site
waters. On large, slaping boulders embracing the width of the stream,
flygroamblystegium fluviatile is entangled by filaments of green algae
and nets made by Hydropsyche; Eurygchium riparioides appears occasion-
ally.

Béggman's Creek
Plate 9. adapted from White. 1882. Pike & Monroe Co.

Monroe Co., Pa.: west of U. S. 209, near bridge (upstream)
(1965:7-30)

Dingman's Creek passes from its source in Silver Lake over Ham-
ilton sandstone, dropping over a series of waterfalls with their lime-
stone coral, to cross a series of massive boulders at the collection
site before joining the Delaware River about a mile below. At the
partly shaded site, these boulders are suggestive of a tilted water-
fall, creating a rapid drop. Consequently, the rushing waters bind
the bryOphytes to the rocks as one tries to lift them from the stream.
The dark gray, flinty Marcellus limestone shales are extremely smooth
and slippery when wet and make one wonder how such large masses of
Eurynchium riparioides are affixed. Round pot holes in the rocks suggest

the past fury of whirling pebbles which carved their basins. In a 2-3

50

foot deep pool, Fontinalis dalecarlica grows on the sandy bottom, while
other submersed Fontinalis plants undulate from rocks in the rapids.
Saw Creek
Plate 9. adapted from White. 1882. Pike & Monroe Co.
Monroe Co., Pa.: west of U. S. 209, near concession stand for Winona
Five Falls

(1965:7-30)

At the collecting site, the moderately fast third order Saw
Creek gives little suggestion of its numerous waterfalls upstream.
Saw Creek drains the eastern half of Porter Township through Catskill,
Chemung, and Genessee formations, dumping its waters into the Delaware
River. Where Saw Creek passes the concession stand, this hemlock-
hardwood shaded stream courses over sand and rocks overlying Hamilton
shales. Here, a short first order tributary apparently arises from
a spring to cut a 2-3 foot channel to Saw Creek.' It is in this trib-
utary that the unusual moss Fontinalis antipyretica var. gigantea
occurs. With its large cup-shaped leaves, this variety provides a
unique cover for midges to hide and reveal only their heads. Other-
wise, the moss does not provide a protective mat, but rather a loose
group of streamers suggestive of some of the vascular hydrOphytes.

Also in the tributary are the vascular hydrophyte Elatine, rock-
dwelling clumps of Aplozia riparia? (possibly a new record for Penn-

sylvania), and streamers of Fontinalis dalecarlica. However, in the

 

main channel, the bryOphyte mats consist of mostly Eurynchium ripar-

 

ioides, while Hygrohypnum ocraceum mats and Fontigalis dalecarlica

streamers are rare.

51

Sciaromium lescurii Streams

Because of its occurrence in several Bygroamblystegium fluviatile
streams, its similarity of growth form, and its somewhat similar insect
grouping, this moss might best be considered with the B, fluviatile
group. As already mentioned, it carpets rocks in the Little Bennett
Creek tributary and accompanies Fontinalis on a downstream rock in
Muddy Creek, but its occurrence in Pidcock Creek and the Toliver trib-
utary as dominants are the only such cases in this study.

Except for a low species diversity, the Sciaromium lescurii insect

 

communities are little different from those of the Hygroamblystegium

 

fluviatile group, and their insects make about the same use of the
moss. The most notable find is a blind specimen of the springtail

Hydroisotoma schafferi in Little Bennett Creek. (Since the species

 

diversity is so low, this moss is not included in the polygonal graphs.)
Pidcock Creek
Plate 8, Fig. 2. adapted fromIUSGS Lambertville quadrangle

Bucks Co., Pa.: Bowmans Hill, west of Pa. 32
(1965:7-4)

The first order Pidcock Creek meanders down the north side of
Bowmans Hill, gathering its upstream waters intermittently in the New-
ark metamorphic rock. Before entering the Delaware River it passes
the collection site where it crosses the Newark igneous and diabase
rocks. Shaded by a mixed hardwood forest, the collection area courses
slowly over sandy, pebbly bottom. On occasional small rocks one can
find mats of silty Sciaromium, while Eurynchium riparioides is even
less common. Occasionally Fissidens hryoides exists among the bases

of these plants.

52

Toliver Run tributary (Tolliver)*
Plate 10, Fig. 1. adapted from USGS Oakland quadrangle

Garrett Co., Md.: ,Swallow Falls Park, above road near entrance.
(1965:5-4)

Near the collecting site in Toliver Run, this first order tribu-
tary contrasts with Toliver by drapping more than 140 feet per mile,
but it passes here at a slower pace than downstream Toliver. Like
Toliver, its fast headwaters drain from the Allegheny formation of
eastern Snaggy Mountain to the rock of Pottsville conglomerate before
the stream enters Toliver Run, which draps rapidly to the Youghiogheny
River. Although it is narrow and shallow, it remains cool under a
hemlock-rhododendron shade. Perhaps it is the size of the stream,
perhaps it is the upstream substrata, perhaps it is only chance. But

the silt-covered Sciaromium lescurii grows in this stream less than

 

100 feet from the mainstream, where it appears to be totally absent.

 

In this more slowly moving stretch of the tributary, Sciaromium lescurii
is a dominant, but it is not known why it appears to be absent from

the portion of Toliver Run examined.
Fissidens bryoides

In general, the Fissidens of these Appalachian streams is B,
bryoides. This is a species complex first described from a land plant
and later subdivided into many varieties. It is distinguished on the
basis of its marginal and costal cells and the number of laminate cells
between the tip of the costa and the border. In this study, the speci-

mens of Fissidens do not precisely fit the number of terminal lamina

 

*Toliver is the spelling found on USGS maps, but the sign in the
park has Tolliver.

53

cells for.B. bryoides, but approximate this species more closely than
any other. Dr. R. A. Pursell (pers. comm.) is in agreement with the
identification.

Any collections of B. bryoides were accidental. The moss is
usually less than 1 cm. long and appeared in this study at the bases
of other mosses. Because of its small size it was probably frequently
overlooked. Since the scape of this study focuses upon insect communi-
ties in or upon bryophyte mats, such small growths would be unimportant
because they are not dense and provide little more substratum than a
bare rock. Consequently, the insect arrays would likely be similar

to those on submersed rock.
Hygrohypnum

Hygrohygnum appears to be a relatively uncommon moss in the regions
studied, but this may be due to the difficulty in collecting its
habitat. For example, the occurrence of H, lggiggg in the falls of
Muddy Creek has already been mentioned, wherein the plants could only
be reached when the water receded during the summer drought. On the
other hand, B, ocraceum occurs in Saw Creek on small rocks and riffles,
but isztill in rapid water. Like the Hygroamblystegium fluviatilg
group, Hygrohypnum forms a relatively dense mat and consequently pro-
vides only small internal spaces. This contrasts with the flowing
strands of Fontinalis and the loose, spacious mats of Scapania.

On Bygrohypnum ocraceum mats in Saw Creek, the tiny, square cases
of the new brachycentrid larva were attached, head upstream, to branch-
es and leaves of the moss. In some instances the moss leaves were

cut in linear pieces and cemented together to form part or all of the

larval case. However, on Eurynchigm giparioides in the same stream,

54

only the brachycentrids Micrasema (sp. 2) and Brachycentrus nr.
numerosus appeared, without the new genus.

Limnophora larvae and pupae, almost exclusive inhabitants of
Hygrohypnum luridum, frequent rapid water of falls where this moss
is profuse, but the larvae occur among other mosses and other regions

of the streams in lower numbers.
Scapania undulata Streams (Scapanietum undulatae)

One of the least common bryophytes studied, Scapania occurred
only in six of the streams and dominated only three. Any generaliza-
tion of its eXpected habitat from such meagre information is premature.
Watson (1919) states that it seems purely a matter of chance whether
Scapania undulata or Nardia compressa (Hook.) Grey is dominant on any
given patch of rock when they occur together in a Pennine stream (Eng-
land), while Gams (in Verdoorn, 1932) contends that it occurs in acid
springs and brooks, often correlated with the Hygrohypnion federation.
Where it occurred in these Appalachian streams, Fontinalis dalecarlica
was always present in the same stream, but not always Hygrohypnum.

In this study Scapania is the only leafy liverwort dominant and
is unique in having two rows of leaves folded so that a small lobe
lies on top. This fold provides a shelter for small insects, while
the nearly flat surface provides a substratum for the attachment of
simulid pupal cases.

§capania provides case-building materials for several caddisflies.
Older larvae of Lepidostoma incorporate portions of the leaf into their
cases, along with flecks of bark and other vegetable matter. But it

is the larva of the microcaddis Paleagapetus celsus which ”prefers"

 

55

liverworts for its case. In 1962, Flint reported the first record

of the larvae of this species, describing their liverwort case of
§capania nemorosa (L.) Dum. (normally a terrestrial species). The
present study appears to be the first record of cases made of Scapania
undulata and glagiochila asplenioides? in several streams where g,
nemorosa was unavailable. Other microcaddis such as Hydrogtila and
Oxyethira use the liverwort for attachment of larval, pre-pupal, and
pupal cases. Even the free-living fixdroPsyche (HydrOpsychidae) uses
§capania leaves in his retreat near the base of the plant.

In May, numerous pupae of the Prosimuligg.hirtipes complex were
attached to the curled tips of Scapania undulata, dotting both upper
and lower surfaces with their thin net cocoons, while in June pupae
of the Simulium tuberosum group had attached cellulose cases arranged
with no apparent pattern. -Even the chironomids made thin cases for
their pupae to lodge between the upper and lower leaves, while one
pupa occupied the empty Scapania case of Paleagapetus celsus. Larvae and
eggs of these chironomid midges can be found between upper and lower
leaves. Other Diptera, such as the tipulid Limonia sp., lay their
eggs on the leaves, so that at times numerous larvae can be found there.
But large larvae of Limonia are scarce, and it is likely that they
move from this protected habitat to other parts of the stream. Mean-
while, the tipulid larva Dolichopeza americana, which is normally a
terrestrial larva (Byers, pers. comm.), appeared in Scapania mats of
Toliver Falls in December and March.

Toliver Run Egglliver)
Plate 10, Fig. 1. adapted from USGS Oakland quadrangle

Garrett Co., Md.: Swallow Falls Park, west of Md. 219
(1965:3-29, 5-4, 6-19, 7-9, 8-25, 12-25; 1966:3-22, 6-11)

56

Toliver Run begins its course in the Allegheny formation on the
eastern slopes of Snaggy Mountain. Perhaps the most beautiful of the
study streams, this hemlock-shaded Youghiogheny tributary courses over
sloping Pottsville conglomerate, drops over step falls. and splashes
into a pool as it plunges over a 4-foot drOp in the collecting area.

Here, Scapania undulata, with occasional Fissidens bryoides at its

 

base, forms a solid mat across the slaping shales, crescents of the
falls, and anywhere the water is very rapid. In slightly slower waters
and pools, detritus-covered Fontinalis dalecarlica forms patches on
the rocky surface, while the sandy bottom farther upstream (above the
road) supports patches of Scapania. However, perhaps in response to
changing water levels, the two bryophytes occasionally grow side by
side in apparently the same amount of flow. One such case is where
the two hang in the drip area of the 4-foot falls while the summer
drought has directed the main flow to a low spot near the edge of the
stream. This situation provides the data for comparison of physical
features of Fontinalis and Scapania, as discussed later.

Laurel Run and Koch Run

Plate 8, Fig. 1.. adapted from USGS Stoddardsville quadrangle

Carbon Co., Pa.: Hickory Run State Park, above and below Pa. 503
(1965:7-29) ‘

As already discussed, these streams are of the Mud Run stream
system and support growths of Fontinalis dalecarlica and Scapania 22f
dualta. Because of their floral and physical similarities, they should
be considered together: both are shallow, sandy and rocky-bottomed
first order tributaries. Laurel Run emerges from the densely shaded

swamp hardwood forest to a sunny, willow-bordered area before again

 

57

PLATE 8. PENNSYLVANIA STREAMS

l. Mud Run System
2. Pidcock Creek
3. PohOpoco Creek

4. Elk Creek

----- Access road
-/- Stream channel
——--Intermittent stream

0 Collection site

58

a ,g .
a. "‘ " d
r: Q. : Dix‘
. 4» 5 "
a: “ g f
5 :
g :0... lf’k’ ‘\
Pa. .0. .0... .0 ea I
534 , n-- ," 'ash“° "’ 8
.0. ; .. 27
a .o -. Mud Run a 3
find Run . -, 3‘ w
0. . .0 h
0. .. H u.
65 v k a i
a -' a -O
O. 1 w
o'.‘ I g
s . 1:
o -"‘ '3
0
O S h
3 ’ o
" l
5’ “x E?
D- : .8
,‘ O
A.
.. 3
a . t
L- 0..
o
6 o a
O a
Q- s
o O
3 ° 2
p o

 

Pa. .1.9g/..‘ 0
. Elk Creek

...
.0
0‘.
. .....O O

a’4
“>“ i b
3 3
O a 4
n o .o .
x-r ' m1.
#0. C33: 1 l 2 3
«3’ 5“
m3
. .2
. A.

 

59

PLATE 9. SAW CREEK AND DINGMAN'S CREEK

.--.'Access road
’\~“Stream channel
-‘\~Intermittent stream

a Collection site

 

6O
1 N
4. Nichecronk Pond

/ Di
0'4.
’ I ‘ ’11:

1 a
Si ver Lake . cybek

.
g
.-

~'.ooe

. ud Pond

Q... 0.

Twelve Mile
Pond ' ' .

‘

.- 4»
. do
w
.2 ' $
0 £9
0 .' 9
s ' e‘
5‘; '93" S“
(I) '0'.

3 Q L g 4#_}mi.

6l

PLATE 10. MARYIAND STREAMS: ALLEGANY AND GARRETT COUNTIES
FIGS.

1. Youghiogheny River tributaries

2. Piney Creek

3. Gramlich Run

4. Sideling Hill Creek tributary

°"--Access road
’\~—Stream channel
,- -/Intermittent stream

0 Collection site

.3, ,. Ginseng Run

:0. 0: $99
{made 3 9’9
'~.Run 3 E “9%
..0' H 1’" H
z. a: 3. Deep Creek
' ' -. -. Mud. .' Lake
>~ ."
‘ . I: :
... Q s
;

Swallow Falls Rd .

8'
A;
. ....
. “E."
hi0

o r.. 00' '0
. 92- :”‘m - .........
....c': 1 D 're_ to u. s. 219
" ., ".3 '. } rib 168
to...” .'o.'

 
   

Piney. Creek

2

u. s. 40
.0 y"

ribu ‘ - 06,)
‘5:
$4»
Q‘s ((2
a 64

9 1 2l 9nd. .

 

63

PLATE 11. MARYIAND STREAMS: MONTGOMERY COUNTY

FIGS.

1. Seneca Creek tributaries

2. Little Bennett Creek

c.'.Access road
/-/Stream channel
,~--Intermittent stream

a Collection site

64

to Woodfield

 

Damascus

 

9 .1 23m.

65

PLATE 12. VIRGINIA STREAMS
FIGS.
1. Rock Castle Creek
2. Sinking Creek
3. Mountain Lake tributary to Sinking Creek
4. Goose Creek

5. Johns Creek

-'v-.Access road
/*\/ Stream channel
"~« Intermittent stream

a Collection site

 

 

66

  

S nking Creek

 

I
New ort
P 2
I
I
I
Mountain Lake
Biological Piedm-nt¢.c“
Station . .g '0‘... ...:O a;
0 .0 o ... ....0000 $
x .9
0“ -
3 to $9
u. s. 40 3
l‘ ‘- ....
\ a O
\\ Q‘e’flfl
Sink ng Creek O '
so"... 14
a .1 z 9 3m.

 

 

 

67

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68

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69

entering the woods around the collecting site. In this area, the
sandy bottom is dotted with a few Scapania-covered rocks while [29:
tinalis extends from others. Meanwhile, Hoch Run emerges from a man-
made wooden dam before converging into a narrow, rocky channel under-
lying a rhododendron thicket. 0n the logs of the dam and the wet rocks
in the spray at the base, thick polsters of Scapania undulata grow,
while the thicket shades rocks mantled with sand-filled Scapania mats

and streamers of Fontinalis on the edges of rocks in riffles.

Insect Communities

Richness, as used by McIntosh (1967), is a measure of the number
of species present, while species diversity includes number of species
and number of individuals. Although richness is inadequate for a
modern balanced comparison of communities, it is both simple and use-
ful. For such comparison of communities simple formulae are used,

a common one being an adaptation of Sorensen's K (i5 Looman and Camp-
bell, 1960), where K - AZ%_§, wherein A - total number of species in
community 1, B - total number of species in community 2, c . number

of species common to both 1 and 2. In this study, A and B represent
numbers of insect species in two communities to be compared. These
communities may occur in different bryOphytes or in different streams.
The highest coefficients indicate the most similarity, while the low
ones indicate dissimilarity. Streams exhibiting very low coefficients
with other streams frequently do so because the low-coefficient streams
,contain very few species of insects.

In Fig. 2 one can see the relationship between number of indivi-

duals of insects sampled and number of species added through chrono-

70

logical sampling. Except for a seasonal influx, the curve on a semi-
1°310 scale approximates a straight line. From this, one could assume
that the number of insect species to be found among these bryOphyte
species in the middle Appalachians would increase if more samples
were taken, suggesting that the implications of community structure
could change if more samples were taken. Inadequate knowledge of
the number of aquatic insect species occurring in this region prevents
us from.making a reasonable estimate of the number of bryo-insects
to expect.
Comparison of insect fauna of various aquatic bryoPhytes

Certain insects appear as obvious accidentals among the bryophytes
(aphids, thrips), while others adapted for a different aquatic
habitat are so infrequent as to be deemed accidental to the bryOphytes
(Blephariceridae, E eorus . To eliminate the effects of these acci-
dentals, only species appearing more frequently than a minimum number
of times were considered: March, May, and June-~more than 2 collections;
summer--more than 4 collections. (The December collections were not
compared in this way because they represented only two streams.) These
arbitrary minimal values were selected because all of the eliminated
insects were non-specific for a particular bryophyte species and were
represented by one or few specimens in a collection. (The 4 summer
collections of Paleagapetus celsus were considered because of their
specificity for Scapania.) While the method is unprecedented, the
results within this study are comparable with each other.

To obtain the community coefficients, the number of insect species
was based on the total presence list for each bryOphyte species.

Tables III-VI give the bryophyte community coefficients, with the

71

TABLE III

COMMUNITY COEFFICIENTS BY BRYOPHYTES, MARCH

 

 

 

 

 

 

 

 

 

 

 

.5 ' .3
n S; 3%
o o
n. :0 :n
Fontinalis 1.00
gcapania .80 1.00
Hygroamblystegium gr. .63 .68 1.00
TABLE IV

COMMUNITY COEFFICIENTS BY BRYOPHYTES, MAY

Eont.’
Scap.

3

 

 

 

 

 

Fontinalis 1.00

Scapania .76 1.00

Hygroamblystegium gr. .90 .73 1.00
TABLE v

COMMUNITY COEFFICIENTS BY BRYOPHYTES, JUNE

 

34
GO
0

[A m =1

_._---,,.__.._.. “I...

f ont.

Esasiaelis 1.00
.539 Basie. . 7 8 1 . oo
Hygrgamblystegium gr. .71 .63 1.00

 

... ~—- .—~ ..——_.._- n-..- ......

 

...—. “flu-...”... m

- .~ VCfl"..-

72
TABLE VI

COMMUNITY COEFFICIENTS BY BRYOPHYTES, SUMMER

 

 

Hygr
Soap.
Sci.

 

 

 

k.
Fontinalis 1.00
Hygroamblystegium gr. .84 1.00
Scapania ' .84 .80 1.00

Sciaromium . .62 .72 ~ .65 1.00

 

 

73

bryOphytes arranged by their similarities. It appears that Fontinalis

 

and Scapania harbor many of the same insects, with Hygroamblystegium

close to these in all but the March collections. Only the Sciaromium

 

collections in summer appear to be notably different from collections
of the other bryOphytes, and even these differences are slight enough
to be attributed to possible sampling error. It is immediately obvious
that these bryophyte coefficients are higher than the stream coeffi-
cients. Such higher coefficients should be expected because, although
they represent small portions of many streams, part of the range of
conditions is eliminated, as reflected by the absence of the other
bryophyte species and the presence of the species considered. This

is but another reminder to us of the complexity of the stream environ-
ment in its relation to the biological response.

Differences in species composition of the insect fauna remaining
in these hand-collected samples of aquatic bryophytes can be attributed
to a number of causes: 1) Chance or sampling error. 2) Season of
collection. 3) Differences between streams; these are both present
physical conditions and recovery time since last scouring by ice, flood,
forest fire runoff, pollution, etc. 4) Differences within the stream
in substratum, current, water depth, light, and seasonal fluctuations
in these. 5) Differences imposed by the bryophytes themselves in
terms of current, sedimentation, light, and physical difference.

If one wishes to examine attribute number 5, then same of the
preceding can be eliminated or reduced by finding places where the
bryOphyte faunas to be examined are adjacent to one another in the
same stream in as homogeneous an area as possible and sample both on

the same dates. While it is not possible to eliminate the different

74

bryOphyte effects on current, sedimentation, light, etc., one can
ascribe, with adequate, unbiased sampling, the differences to the
bryOphytes per se or to the microenvironmental conditions imposed
by or occupied by each bryoPhyte species.

To compare the insect fauna in two closely adjacent.bry0phyte
species, Fontinalis dalecarlica and Scapania were studied in the
Toliver Falls collection site. As mentioned previously, both

Fontinalis and Scapania occurred in Toliver Run in dripping water

 

away from the main flow of the falls. In this instance (July 29) one
handful of Scapania harbored six times as many total insects per gram

as one handful of Fontinalis, while Fontinalis harbored twice as many

 

individuals of RhyacOphila but had fewer of everything else. On the
other hand, few abundant Scapania insects were excluded on Fontinalis:
Simulium spp. averaged 418 on Scapania and none on Fontinalis while
Leuctra dropped from 42 to .7 per gram. One could assume the differ-
ence in total numbers lies in the great amount of internal chambering
in Scapania, whereas Fontinalis provides little more protection than
a handful of strings. But these higher counts may well be related to
the fact that Fontinalis has a heavy axis and is generally a heavier
plant than Scapania.

When comparing all the Scapania (7 collections), Fontinalis (4),
and SematoPhyllum marylandicum (2) (an occasional species on the
edge of Toliver Run) collections for March (Plate 13), there are again
differences both in kind and number of insects. Certainly the

Sematophyllum insect faunal community is much less diverse, exhibiting

 

only three of the top 20 species of insects and having counts an

order of magnitude lower. On the other hand, Scapania is the most

75

PLATE 13. MARCH, TOLIVER RUN: RELATIVE NUMBER OF INSECTS IN THREE
SPECIES OF BRYOPHYTES. EACH LINE REPRESENTS A DIFFERENT HANDFUL, BUT

ALL ARE ON THE SAME COLLECTION DATE. SCALE IS LOG 0F NUMBERS OF

10
INDIVIDUALS PER GRAM OF BRYOPHYTE.

Fontinalis dalecarlica

 

 

——‘“‘Scapania undulata

 

"""" Sematophyllum marylandicum

 

..

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11.133:

77

diverse by having four species (Ehyacgphila carolina group, Paleaga-
peggg celsus, Peltoperla sp., and Promoresia elegggg) which are
totally absent from both Fontinalis and Sematophyllum clumps. Further,
Scapania has higher counts per gram.dry weight of all insects in near-
ly every collection. Although these three bryophytes can be expected
to produce different physical influences, the presence also reflects
habitat differences such as falls, interfalls, and edge areas, respec -
thufly. It is possible that such habitat differences explain the mutual
exclusion of the Simulium tuberosum group and the gilled Nemoura, for
both of these occur in Fontinalis and Scapania, but never in the
same collection.
Comparison of streams

A glance at Tables VII-X shows that streams range from very simi-
lar (cammunity coefficient of .86) to very different (.07). Those
with the highest coefficients with other streams are generally those
with the most species of insects, and the lowest coefficient series
coincide with streams having the fewest species. For ease of com-
parison, the items in the tables have been grouped by dominant bryo-
phyte. Any attempt to order these streams according to their coeffi-
cients results in a multidimensional chaos due to the complicated
interrelationships. But, the overall picture, when partitioned into
subsets, shows a high similarity within a bryOphyte species, even though

an occasional interspecies pair may also be highly correlated.

 

Small coefficients and a small number of species are exhibited
by Little Bennett Creek and Pidcock Creek. These small, sluggish
streams would stand out, even to the casual observer, as being quite

different from the other cool, rocky, mountain streams. In March the

‘11 lullll .
.I‘I

To
De
Ro
Hy
Gr
Sd
Si
Sth
LB
WBu

WBe

To
De
Ro
HY
Gr

Sd

78
TABLE VII

COMMUNITY COEFFICIENTS BY STREAMS, MARCH

U
50)
a 2: ass as E a s as
1.00
.85 1.00

.54 .59 .58 .62 1.00

.25 .43 .44 .48 .62 1.00

.34 .32 .46 .50 .56 .52 1.00

.22 .17 .26 .29 .33~ .27 .44 1.00

.46 .39 .41 .44 .63 .35 .46 .44 1.00

.22 .26 .26 .21 .42 .27 .33 .40 .44 1.00

.08 .10 .07 .08 .19 .17 .13 .29 .27 .57 1.00

.33 .31 .41 .32 .37 .56 .38 .31 .29 .15 .20 1.00

Toliver Run Sth Mountain Lake trib. to Sinking Creek
Deep Creek tributary 31 Sinking Creek

Goose Creek LB Little Bennett Creek

Hoyes Run WBu Seneca Creek, Burroughs trib.
Gramlich Run WBe Seneca Creek, Bell trib.

Sideling Hill Creek My Muddy Creek

 

1| lulu I 1.6!! {Ila}: t {Ill-Ill...

Gr.
Gi
Tot
De
To
1’?
Ne

My

LB
Gr
61
Tot
De
To
Py

Ne

79

TABLE VIII

COMMUNITY COEFFICIENTS BY STREAMS, MAY

LB
Gr‘
G1

.51 .53 1.00

.52 .62 .72

.44 .44 .57
.42 .59 .71
.44 .57 .51

.25 .42 .48

Little Bennett Creek

Gramlich Run

Ginseng Run

Tot

1.00

.53

.59

a o :x
a: pa 9.
1.00
.78 1.00
.44 .65 1.00

Ne
My

.40 .48 .55 1.00

.26 .37 .30

Toliver Run tributary

Deep Creek tributary

Toliver Run
Piney Creek
Neds Run

Muddy Creek

.25 1.00

80

TABLE IX

COMMUNITY COEFFICIENTS BY STREAMS, JUNE

Cr 1.00

LB .60 1.00

De .50 .66 1.00

To .56 .32 .70 1.00

My .56 .28 .56 .56 1.00

Hy .50 .35 .50 .50 .65 1.00

Gr Gramlich Run

LB Little Bennett Creek
De Deep Creek tributary
To Toliver Run

My Muddy Creek

Hy Hoyes Run

 

 

81

 

 

 

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83

equally sluggish Seneca Creek tributaries show similar dissimilarity
when compared with most of the other streams. Perhaps this is be-
cause they, too, have few species of insects.

As mentioned above, the streams with the greatest number of insect
species have the highest coefficients, even with the streams supporting
a low number of insect species. This may be due to the relatively
small total number of species (150), so that any stream with many spe-
cies nearly covers the species repertoire for a particular season.
Thus, it appears as though conditions which restrict the bryOphytes
can differ in several directions which, in turn, differentially influ-
ence the various insects.

In general, we might state that the similarity among streams
with the same dominant bryOphyte is greater than their similarity with
streams having other dominants. Toliver Run and Deep Creek tributary
are exceptions. In the three seasons sampled, the Toliver Run and
Deep Creek tributary pair always have the highest coefficient. Both
of these streams have Fontinalis and Scapania, but Toliver has Scapan-
ig_as a dominant, while Deep Creek has Hygroamblystegium as its dom-
inant. Since both streams are at a similar elevation in Garrett County,
Maryland, this may be one of the major factors influencing their
similarity. But it has already been pointed out that 39322315 is
relatively uncommon. Its presence in a stream may indicate a selective
set of characteristics which influence the insects as well as Scapania,
thus accounting for the insect faunal similarity of these two proximal
streams. In this same connection, we should note the similarity of
the three Scapania streams in the summer collections: Toliver Run,
Laurel Run, and Hoch Run. But it is also noteworthy that Hoch Run

is most similar to Saw Creek. This connection is difficult to ex-

84

plain, for the two streams occur at different elevations in different
counties in Pennsylvania and do not give a strikingly similar appearance,
wherein Saw Creek is much wider than Hoch Run. Nevertheless, a glance
at the righthand coefficient of each stream set will point out that
most of the streams are most similar to others havingthe same dom-
inant bryophyte, and certain clumps of similar streams might be in-
ferred from the table. As these coefficients have no confidence
limits, it is dangerous to rely heavily on many of the slight differ-
ences indicated, and I will not attempt to offer any explanation for
the clumps.
Insect communities by season;

The relative abundances and species relationships of insects
with the bryOphyte species can be inferred by the use of polygonal
graphs. Hutchinson (1940) and Davidson.(l947) have used polygonal
graphs to show environmental parameters and taxonomic characters,
respectively. In the present study, the graph is adapted to show
community relationships. The parameters are the insect species, which
are shown as mean number per gram by a log10 scale on each radius.
These insects have been arranged from high to low frequency (based
on Tables XII-XVI in the Appendix), with the most frequent on the
right side of the graph and least frequent on the left. Before
obtaining the means, the collection data were organized by bryophyte
species. For each bryophyte species, means of insect counts were
first computed for that species in each stream. Then, the mean of
these was computed to obtain an overall mean for the bryophyte among

the stream types. The Hygroamblystegium group required a third mean:

 

after each bryOphyte mean was obtained, these were averaged to obtain

85

the final mean. (Explanation of the Hygroamblygtegipm group is found
in the bryOphyte section.) 1

Based on the polygonal graphs, certain communities become appar-
ent in the five seasonal groups. Figure 3 compares the seasonal as-
pects of the more abundant ones.

In‘March (Plate 14), the chironomids dominate Fontinalis 9212-

33111;; and the Hygroamblygtegium group, while the Prosimulium hirtipes

 

group larvae are most abundant on Scapania undulata. Fluctuations
in abundance occur among most of the other insects, but certain dif-

ferences are striking: l) The Ephemerella invaria group occurs

 

exclusively with the Hygroamblystegium group. 2) Fontinalis is

 

marked by the absence of Pericoma, Bezzia, Empidae, and Dasyhelea,

 

all of which are small Diptera with poor or no adaptation for cling-

ing, and E, dalecarlica lacks the blimp-shaped Peltoperla, possibly

 

for the same reason. 3) The caddisfly RhyacOphila carolina group
is much more abundant with Scapania than with Hygroamblystegium.and

it is absent on‘g. dalecarlica; the caddisflies Cheumatopsyche and

 

 

HydrOptila are absent in g. undulata communities. Since the sampling

 

for this survey was not designed to provide a rigorous test of this
kind of correlation, no confidence intervals can be provided for these
absences, nor do I have any conjectures on the possible reasons for
these apparent relationships.

By‘ng (Plate 15), the numbers have greatly increased for most
insects, as seen in Fig. 3, and now chironomids on g. undulata show
an order of magnitude lower count than for the other bryOphytes. In
fact, during this period, the insect fauna is poorer in total species

and numbers in S. undulata turf, when compared to the other two bryo-

86

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unsw=¢rxaan
unsavoon he: noun:

 

 

 

 

 

 

 

 

 

 

J

 

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.0a

 

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30H

 

 

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fib0~

 

 

Number of Insects per Gram of BryOphyte

88

phyte groups. This might be expected from S, undulata's limited
distribution and the small total number of collections (6). It should
also be pointed out that if one aquatic insect has-a very high restric-
tion to g, undulata and is at the same time very abundant, it may be
responsible for keepingthe other insects low in this bryophyte in
various direct ways.

Beginning in May and becoming very obviOus in gggg_(Plate 16)
is the scarcity of Promoresia elegans in S, undulata, while Baetis
is totally absent in the June collections of S, undulata. On the
other hand, there are much larger Leuctra and Simulium tuberosum group
populations in Scapania than in the other bryophytes, while the
emerging Prosimulium hirtipes group is nearly absent in all three.
(Note that the chironomids have been omitted on this graph to permit
comparison of more selective species.)

Summer (Plate 17) shows little major difference among the bryo-
phytes except for the absence of large insects in S, undulata:

Togpperla, Pteronarcys spp., Cheumatopsyche. Furthermore, Pteronarcys

 

biloba on F, dalecarlica and Pteronarcys proteus among the Hygroambly-
stegium group are mutually exclusive. At this time, extremely high
counts of midges are obtained, reaching as much as 2500 per gram in
one Scapania collection.

In December (Plate 18), the comparison is really one between
Muddy Creek and Toliver Run. Only 17 taxa were present, 9 on 2. £215-
carlica and 12 on g. undulata. Thus, only 4 occurred on both: the

Prosimulium hirtipes group, Chironomidae, Empidae, and HydrOpeyche.

 

These differences in taxa are probably due 0 multiple stream differ-

ences as well as bryOphyte differences.

89

PLATE 14. MARCH: RELATIVE NUMBERS OF INSECTS PER GRAM DRY WEIGHT
OF BRYOPHYTE
-——-—Hygroamblystegium fluviatile group

‘—-——4Fontinalis dalecarlica

 

...... Scapania undulata

The scale is log10 of insects per gram of bryOphyte.

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91

PLATE 15. MAY: RELATIVE NUMBER OF INSECTS PER GRAN DRY WEIGHT
0F BRYOPHYTE
————~flygroamblysteg;um fluviatile group

Fontinalis dalecarlica

 

----- Scapania undulata

The scale is 10810 of insects per gram of bryOphyte.

 

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93

PIATE 16. JUNE: RELATIVE NUMBERS OF INSECTS PER GRAN DRY WEIGHT

OF BRYOPHYTE

——-—~Hygroamb1ystegium fluviatile group

Fontinalis dalecarlica

 

----- Scapania undulata

The scale is 10g10 of insects per gram of bry0phyte.

9h

.aa

 

 

 

 

 

 

 

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mwuwMOuaom Ilommmvdmuq
wmwuawunuv
mmssodanmodom
amawmmal mmmumwmmm
aumuuuu; mmwmmW¢
fiiflfl:fiwfl@&m
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gamma I-mowfiuwm
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MflwzummMNmm QMHMu

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9S

l‘h‘tl‘l L7. T~§ID~SUHFIEiRz REIATIVE I'JUf'iBl'jRS 0F INSECTS PER GRAN DRY WEIGHT

OF BRYOPHYT '2'}

.. -———--—7. "“ ‘f "L I" ' ‘ V '1 w ‘ |\ - ' ' ‘ v
‘- s)"1“;QCZ‘V‘UJ1:3Lf‘LILIcn flu «’Lt'tl 1.1.; 810.1};

.lAL‘

 

—- --——~ --}7;:.‘;ti.nali s (m lecar HCZ

"" .ficc’ianxa undulata

 

The scale is 10810 of insects 1~r gram ur bryophyte.

96

.au

 

 

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N .a. v nan qummmmmmmm
I
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alfiouuouadm oo 1 3060 m

 

 

 

 

 

 

 

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97

PLATE 18. DECEMBER: RELATIVE NUMBERS 0F INSECTS PER GRAN DRY WEIGHT

OF BRYOPHYTE

—~w—~fontinalis dalecarlica

 

"Scapania undulata

 

98

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osozamouueauno

99

Seasonal pulses, as would be expected, are evidenced by high
numbers of individuals and species in the mid-summer months, while
a low point is reached in the March collections. A comparison of
Toliver Run collections indicates that counts per gram of Chironomi-
dae are higher in December than in March or May, but are nearly equal
in June, while summer counts far surpass all other seasons. Mean-

while, the Prosimulium group and Leuctra also drop from December to

 

March. This study was not designed to measure or explain these dif-
ferences, so we cannot determine whether the higher counts in
December are influenced by such things as the life cycles of the species,
natural reduction of numbers, predation, or a seasonal preference
for the bryOphyte. Macan and Worthington, 1951, mention the rolling
stone that gathers no moss. Perhaps the high December counts of
insects are due to a migration to the stable, moss-covered rocks during
a time when ice and heavy snowmelt loads in Toliver Run could dislodge
insects from other areas. Furthermore, riffles, with their moss-covered
rocks, are usually the last part of the stream to freeze, thus providing
the most likely spot for an insect to receive flowing water and food.
”Masses and liverworts often form extensive awards where the substratum
is rocky or stony, and they profoundly influence the fauna by providing
a foothold for animals which otherwise would be swept away by the
current." (Macan and Worthington, 1951)

From the seasonal data, one can approximate the bryophyte-insect

faunal communities. (Refer to Fig. 3 for the seasonal distribution.)

 

 

 

Bryophytes in general: Hygroamblystegium group:
Chironomidae Ephemerella spp.
Prosimulium hirtipes group Eteronarcys proteus

 

 

Simulium tuberosum group Pericoma

Brypphytes in general (cont.):

Isoperla bilineata
Nemoura spp.
Dolqphiloides distinctus
Dasyhelea spp.

Bezzia spp.
Fontinalis dalecarlica

Micrasema spp.
Pteronarcyggbiloba
Cheumatopsyche
Chimarra aterrima
Promoresia elegans
Baetis spp.

100

Hygroamblysgggiumggroup (cont.):

Rhyacophila invaria
Micrasema spp.
Promoresia elegans
Peltoperla sp.
Baetis spp.
Empididae

Scapania undulata

Paleagapetus celsus
Rhyacophila carolina
ghyacophila invaria
thioservus
Empididae

Insect Biology and Ecology

(Table XI 'summarizes the habitat and adaptations.)

Coleo tera beetles

Among the beetles listed, only the Elmidae, the riffle beetles
(larvae and adults), provide a significant part of the fauna. (The
same is true in the European studies by Carpenter, 1927; Percival
and Whitehead, 1929; and Frost, 1942.) In march their numbers were
low except in Hayes Run, where Promoresia elegans larvae represented
the largest fraction of the papulation, even higer than the midge

larvae of the Chironomidae. In May, Promoresia still was less important

 

except for dominating the Muddy Creek bryo-fauna. .By June, the num-

bers of Promoresia had increased an appreciable extent in both Hayes

 

Run and Muddy Creek, while remaining relatively unimportant in the
Scapania-dominant Toliver. But in summer, its frequency was second
only to the Chironomidae, reaching high counts among nearly every
Then, with the onset of winter, only two

species of bryOphyte.

adults appeared among the Muddy Creek and Toliver collections.

101

TABLE XI

TAXONOMIC LIST OF INSECTS AND ADAPTATIONS

Coleoptera

Dryopidae
Helichus sp.

Dytiscidae
Ilybius sp.
gydr0porus sp.

Elmidae .
Dubiraphia sp. 1
Microcylloepus sp. 1
thioservus sp. 1'
thioservus sp. 2
Promoresia elegans LeConte
Stenelmis crenata (Say)

Gyrinidae
Dineutus sp.

Haliplidae

Brychius sp.

Haliplus sp.
Hydraenidae

Limnebius sp.
HydrOphilidae
Enochrus sp.
Tr0pisternus sp.
Psephenidae
EctoEaria ?
Collembola
Brachystomellidae
Odontella lamellifera (Axelson)
Entomobryidae

 

 

 

 

 

 

 

 

Entomobrya griseoolivata (Packard)

Orches e 1 la glinquefascia ta (Bour let)
HypOgastruridae

flypogastrura armatus (Nicolet)

Schotella glasgowi (Folsom)
Isotomidae

Hydroisotoma schafferi (Krausbauer)

Isotoma violacea Tullberg

Isotoma viridis Bourlet

Isotomurus_palustris (Muller)

 

 

 

 

 

lateral compression

x x x x x x x x
small body

X

XX
XX

><><><><
XNXX

no ventral adhesion

attachment
weight

xxxxxx xx

:4

XXXXXX

no swimming hairs

X
X

XXX><><><
X><><><><><

X
X

XXXX
XXXX

cover gills or none

X

BryOphytes

102

Neanuridae

Pseudachorutes lunatus Folsom
Onychiuridae

Onychiurus subtenius Folsom
Sminthuridae

Sminturides aguaticus (Bourlet)
Tomoceridae

Tomocerus flavescens (Tullberg)

 

 

 

 

Diptera

Blephariceridae
Blepharicera sp.
Chironomidae
Dixidae
Dixa sp.
Dolichopodidae
_flydr0phorus sp.
Empididae
Empid sp. 1
Empid sp. 2
Empid sp. 3
Hemerodromia cf. rogatoris Coquillett
Hemerodromia cf. seguyi Vaillant
Heleidae
Alluaudomyia sp.
Atrichgpogon sp.
Bezzia sp. 1
Bezzia spp.

Dasyhelea sp. 1

Dasyhelea sp. 2
Muscidae

Limnophora sp.
Psychodidae
Pericoma sp.
Rhagionidae
Atherix variegata Walker
Simulidae
Cnephia mutata (Malloch)
Prosimulium hirtipes group (Fries)
Prosimulium magnum.Dyar & Shannon
Prosimulium.mixtum Syme & Davies

 

 

 

 

 

 

 

 

 

 

lateral compression

small body

XXXXXX
XXXXXX
XXXXXX

X

Prosimulium rhizophorum Stone & Jamnback

 

Simulium nr. gauldingi Stone

Simulium impar Davies, Peterson & Wood

 

” Simulium_pgrnassum Malloch
Simulium tuberosum (Lundstrom)
Simulium venustum Say

 

 

 

X
X

XXXXX

X
X

X
X
X

XXXXXXXXXX
XXXXXXXXXX
XXXXXXXXXX

no ventral adhesion

attachment

weight

X

X

XXXXX

X

XXXXX

no swimming hairs

y,

X
X

XXXXX

X X XXXXXX
X

X

X
X

XXXXXXXXXX
XXXXXXXXXX

cover gills or none

XXXXX
"1'1!

XXXXXX
"Iii/JIM

U)

BryOphytes

Him

in

103

Simulium verecundum Stone & Jamnback
Simulium vittatum Zetterstedt

 

 

Tabanidae
Thaumaleidae

Thaumalea sp.

Tipulidae

Antocha sp.

DolichoPeza americana Needham
Hexatoma nr. longicornis (Walker)
Hexatoma nr. pinosa (Osten Sacken)
Limnthila nr. macrocera (Say)
Limnophila sp.

Molophilus sp.?

Ormosia sp.?

Tipula collaris Say

Tipula sp. 1

Tipula sp. 2

Tipula sp. 3

Tipula sp. 4

Tipula sp. 5

Tipula sp. 6

 

 

 

 

 

lateral compression

small body

XX
X
X

no ventral adhesion

attachment
weight

X

X

X

XXXXXXXXXXXXXXX

no swimming hairs

XXX

X
X

XXXXXXXXXXXXXXX
XXXXXXXXXXXXXXX

cover gills or none

XX

U'JUJ

BryOphytes

Maggot?
EphemerOptera

Baetidae
Ameletus sp.
Baetis spp.
Baetisca callosa Traver
Baetisca carolina Traver

Caenidae
Caenis sp.

Ephemerellidae
Ephemerella alleghcniensis Traver
Ephemerella attenuata McDunnough
Ephemerella catawba Traver
Ephemerella deficiens Morgan
Ephemerella funeralis McDunnough
Ephemerella nr. invaria (Walker)
Ephemerella serratoides McDunnough
Ephemerella subvaria McDunnough
Ephemerella temporalis McDunnough

Heptageniidae
Epeorus sp.

Leptophlebiidae
Paraleptophlebia spp.

 

 

 

 

 

 

 

 

 

 

XXXX
XXXX
XXXX

X
X
X

XXXXXXXXX
XXXXXXXXX
XXXXXXXXX
XXXXXXXXX

X
X

104

Siphlonuridae
Isonychia sp.

hemiptera

 

 

 

 

 

 

 

 

 

 

Veliidae
Microvelia spp.
MegaIOptera
Corydalidae
Nigronia sp.
Sialidae
Sialis sp.
Odonata
Cordulegasteridae
Cordulegaster sp.?‘
Gomphidae
. Gomphus sp.?
Octogomphus sp. ?
Plecoptera
Capniidae
Allocapnia sp.
ISOperlidae
Isogerla bilineata (Say)
Isoperla duplicate (Banks)
Leuctridae
Leuctra
Nemouridae
Nemoura sinuata Wu
Nemoura vallicularia Wu
Nemoura venosa Banks
Taeniopteryx sp.
PeltOperlidae
Peltoperla sp.
Perlidae

Acroneuria sp.
Perlesta placida (Hagen)
Phasgangphora capitata (Pictet)
Paragnetina sp.

Pteronarcidae
Pteronarcys biloba Newman
Pteronarcys proteus Newman

Trichoptera

Brachycentridae
Brachycentrid gen. 1
Brachycentrus nr. numerosus (Say)

 

 

 

 

 

 

lateral compression

small body

X

X
X

X
X

XXXX

XXXX

X
X

X
X

no ventral adhesion

X

XXXX

XX

(D
c
m o
u c
-H
m n
.c 0
now
u Cir-l
l: w-‘v-l
w E.“
E 00
U-H
0.: 3 u
m mam m
u-H >
u m o o
m 3 c o_
x x
x x x
x
x x x
x x x F
x x x
x x x F
x x x F
x x x F
x x x F
x x x H
x x x F
x x
x x
x x F
x x

XX
XX
25‘"!

x x x x H
x x x x H

BryOphytes

U)

U}
:m:

105

Brachycentrus sp.

Micrasema sp. 1

Micrasema sp. 2

Micrasema sp. 3
HydrOpsychidae

CheumatOpsyche spp.

Diplectrona modesta Banks

Hydr0psyche spp.

Parapsyche apicalis (Banks)
Hydroptilidae

gydroptila sp.

HydrOptilid gen. 1 (gen. nov.)

Ithytrichia sp.?

Mayatrichia sp.

Neotrichia sp._

Oxyethira
Baleagapetus celsus Ross

Iascobia sp.
Lepidostomatidae
Lepidostoma americana (Banks)
Lepidostoma sp.
Limnephilidae
Neophylax concinnus McLachlan
NeoPhylax consimilis Betten
NeoPhylax oligius Ross
Platycentropus sp.
chnopsyche luculenta (Betten)
gycnopsyche cf. scabripennis (Rambur)
Odontoceridae
Psilotreta sp. 1
Psilotreta sp. 2
Philopotamidao
Chimarra sterrima Hagen
Dolophiloides distinctus (Walker)
Psychomyiidae
Polycentropus,sp.
Rhyac0philidae
Glossosoma sp.
Rhyac0phila nr. carolina Banks
Rhyac0phila fuscula (Walker)
Rhyac0phila nr. invaria (Walker)
Rhyac0phila minors Banks
Rhyac0phila torva Hagen
Rhyac0phila sp. 2 (see Flint, 1962)
RhyacoghiIa sp. 5 (see Flint, 1962)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

lateral compression

small body

XXXXXXXX

XXXX
XXXX

XXXXXXXX
XXXXXXXX
XXXXXXXX

XX
XX

no ventral adhesion

attachment
weight

XXXX
XXXX

X XXXXXX

X

XX

X

XXXXXXXX

XXXX

XXXX

X
X

XXXXXX
XXXXXX

X
X
X

X
X

XXXXXX

X
X

X

xix x x

x x x x ‘no swimming hairs

XXXX

XXXX

XXXXXX

X
X

XXXXXXXX

XXXXXXXX

X

XXXXXXXX

cover gills or none

U3

CD’TIUJ

I

BryOphytes

I:

F H

106

Collembola (Springtails)
Although the Collembola frequent the surface of ponds and lakes

and inhabit the cushions of terrestrial bryOphytes, they are relative-
ly unimportant among the aquatic bryOphytes of rapid waters. While
their tiny size fits one characteristic for bryo-fauna, their mode

of locomotion--a spring on the abdomen--wou1d be difficult to maneuver
inside the mat, and it would lead to an ocean-ward trip if used out-
side. Only the common Isotomurus Balustris appeared in more than two
collections.

Whereas their role in the bryophytes of mountain streams seems
unimportant, one collembolan distribution is interesting, for 955227
ggll§,guinguefasciata from Toliver Run was previously unknown in
North America (Richard Snider, pers. comm.). And gygroisotoma
gghafferi from Little Bennett Creek is blind-~an atypical form.

Di tera lies

"The Chironomidae provide by far the largest numbers of insect
larvae and reach their greatest development among thick mossy growths
such as occur at Harewood Bridge, Pool Weir and on mossy stones at
Grassington and similar places." (Percival and Whitehead, 1930)

Certainly the chironomids are no exception in this study, where
they reach counts as high as 2500 per gram dry weight of moss in
summer and appear as the most frequent insect in every season, although
in some of the March collecuons they were surpassed in numbers by
other insects: Prosimulium spp. and Ephemerella spp. They reach
their highest numbers in the mats and turfs, not the Fontinalis stream-
ers. Because of the loose nature of Fontinalis dalecarlica, many

of these larvae are of the case-building type or inhabit the protected

107

leaf bases, while the highest numbers of individuals occur in mats
and turfs of other bryOphytes. In his discussion of adaptations,
Muttkowski (1929) points out that insects of specialized habitats
(such as these) usually have generalized feeding habits. The
Chironomidae, as a family, are detritus-feeders, herbivores, and
carnivores. .Among the moss inhabitants, the common Hydraeninae are
phytOphagous and detritus-feeders. Percival and Whitehead (1929)
report that Chironomidae feed on diatoms, the major constituent of
the bryOphyte community flora (author, unpub. data).

Second to the Chironomidae, one of the species of Simulidae
(black flies) usually outnumbers the other insects: in March, the

Prosimulium hirtipes group; in May, the Simulium tuberosum complex;

 

in June the Simulium tuberosum complex is third most frequent with

 

the stonefly Leuctra being more frequent but often less abundant,

and in mid-summer, the aiggligg,ggbgggggmbcomplex is surpassed in
frequency by Promoresia elegans; in December the Prosimulium.hirtipes
group again becomes second most frequent. The Prosimulium group was
not encountered after June 19. Other investigators (Davies, Peterson,
& Wood, 1962) have indicated that adults of this group emerge April
through June. Perhaps their absence in the present study indicates
they have all either emerged or died. These same investigators indi-
cate that Simulium tuberosum adults emerge in late May throughout the
summer, and indeed in the present study they were encountered in the
last samples which were taken in August, but were not encountered in
December. In the present study of the middle Appalachians, the ro-
simulium group over-wintered as larvae and even by late March a few

pupae had developed. Meanwhile, Simulium spp. apparently overwinter

108
as eggs; a few tiny larvae appear in March, but their peak abundance
is in May, at which time very few pupae are formed.

The Prosimulium hirtipes larvae from Toliver Run, June 11, 1966,

 

and muddy Creek, March 22, 1966, are close to Prosimulium mixtgg_8yme
& Davies, (Davies, pers. comm.) and probably represent the first
record of this species in the middle Appalachians. (They were first
named in 1958 from Ontario specimens.) Others of the Prosimulium
hirtipes group include larvae from Toliver Run (Dec. 25, 1965) which
are close to Prosimulium saltus (Davies, pers. comm.). Prosimulium

rhizophorum (not included in the Ontario studies) and Prosimulium

 

magnum both occur in Gramlich Run.

Another recently described species, Simulium impar Davies, Peter-

 

son, & Wood (1962), occurs in Toliver Run (pupae: June 11, 1966)

and Little Bennett Creek (larvae: May 8, 1966). Other species
contributing possible Maryland records include Simulium vittatum
(pupae: Seneca Creek tributary at Burroughs' and Bell's, March 27,
1965); Simulium venustum (larvae: Toliver Run, June 19, July 9, 1965;
June 11, 1966); Simulium parnassum.(larvae: Toliver Run, June 19,

1965); Simulium verecundum (larvae: muddy Creek, June 11, 1966);

 

Simulium nr. furculatum or gouldingi (larvae: Saw Creek, July 30,

 

1965). A Simulium (Eusimulium) represents a possible new species

 

(larvae: Saw Creek, July 30, 1965).

Although feeding habits of Simulidae were not observed in the
present study, Percival and Whitehead (1929) report that the Simulium
reptans gut contains moss, diatoms, and other algae. Its method of
filter feeding suggests that the moss entered as detrital fragments
and may not have been digested. The author has seen larvae of another

midge-like dipteran in the Rhyphidae eat moss, but the fragments passed

109

through the digestive tract intact, with only the epiphytes being
digested. (During a conversation with Dr. Kenneth Cummins, we

discussed cases where the animal is adapted to eat foods he is unadapted
to digest, e. g. Smirnov, 1962. I suspect that many moss feeders
including §imnlium fit this category.)

Although Percival and Whitehead (1930) report the Tipulid
fiexatggg sp. in numbers approaching the chironomids, this genus was
rarely represented in the Appalachian collections and was absent in
Frost's streams (1942). Another tipulid, Dolichogeza americana, is
known from temporarily submerged habitats with ficapania nemorosa
(Byers, pers. comm.). But in this study, the larvae occurred among
Scapania undulata in Toliver Falls--an area normally under water
except for dry summers.

While empids and Atherix variegata are occasional constituents,
the only other Diptera of importance are Psychodidae larvae: Pericoma
sp. (similar to g. canescens Meig. and g, cognata Eaton). These
larvae frequent smaller streams in the tight mats of the Hygroambly-
stegium group and are almost completely absent among Fontinalis,
Scapania, and gygrohypnmm, where they do not have much protection

from drifting (Eontinalis), or they would be in rapid water (Scapania

and fixgrohypnmm).
EphemerOEtera (Mayflies) '

Whereas some Ephemerella naiads of this study apparently spend

 

their entire nymphal lives in and around mosses, Baetis spp. can
only be found in early stages, even during March and July when their
maximal sizes occur (Waters, 1966). That Baetis is typically a bottom

form frequenting stream drift (Waters, 1961) explains its absence

110

from the moss. Meanwhile, Paraleptophlebia is a free swimmer, frequent-

 

ly clinging to the surface of the mat in any stage, never abundant,
but possibly lost during sampling.

Typically a clinging form, springtime Ephemerella spp. reach
extremely high numbers in several gygroamblystegium streams, but are
rare in Scapania and Fontinalis streams (Plates 14 & 15). If any
group feeds on mosses, it is probably these herbivores; while the

Ephemerella of Linesville Creek feed on algae and detritus (Cummins,

 

unpub. data), Ephemerella ignita composes most of its diet from moss

 

(Percival and Whitehead, 1929). This hypothesis needs to be substanti-

ated by feeding aufwuch-free moss to Ephemerella to determine if the

 

moss is truely the food source.

Several collections of Ephemerella extend the known range of the

 

species:

g. allegheniensis north to Pennsylvania (Saw Creek, 7-29, 65).
Allen and Edmunds, 1962

E: catawba north to Magyland (Ginseng Run, 7-9, 65), Hoyes Run
(6-11, 65). Allen and Edmunds, 1965

Many of the collections of Ephemerella may be state records,
based on the recent examinations of the genus by Allen, Berner, and

Edmunds:

g. funeralis in Maryland (Little Bennett Creek, 3-25, 65; Toliver
Run, 6-19, 65; 7-9, 65; Ginseng Run, 7-9, 63; in Pennsylvania
(Laurel Run, 7-29, 65; Hoch Run, 7-29, 65); in Virginia (Sink-
ing Creek, 3-18, 65). Allen and Edmunds, 1963b

E. attenuata inm mEyland (Muddy Creek, 7-9, 65). Allen and
Edmunds, 1961

.E. cornutells in Maryland (Ginseng Run, not in moss, 7-9, 65).
Allen and Edmunds, 1962

E. serratoides in Magyland (Muddy Creek, 7-9, 65); in Pennsylvania
(Dingman' s Creek, 7-30, 65, Elk Creek, 7-28, 65; Hoch Run, 7-29,
65; PohOpoco Creek, 7-29, 65). Allen and Edmunds, l963e

 

II n
I {KL
. I. .II‘ I
I. Null. 1

111

g. subvaria (not verified) in Maryland (Toliver Run, 6-11, 65).
Allen and Edmunds, 1965

E, temporalis in Magyland (Hoyes Run, 8-25, 65; Little Bennett
Creek, 8-20, 65; Muddy Creek, 6-11, 66; Neds Run, 5-4, 65;
Toliver Run, 5-4, 65; 6-11, 66); in Pennsylvania (Laurel Run,
7-29, 65; Mud Run, 7-29, 65); in Virginia (Goose Creek, 3-22,
65). Allen and Edmunds, 1963b

Ephemerella invaria might be added for all three states, but
these naiads have not been positively separated from E. dorothea.

Although not dominant bryo-insects, Baetisca carolina from.Goose
Creek (3-22, 65) extends its range northward, while the appearance
of Baetisca callosa is a new record for Virginia. (Berner, 1955)
PlecoPtera (Stoneflies)

The taxonomy of this group is extremely difficult because the
moss inhabitants are quite small, and taxonomic characters are fre-
quently indistinguishable on the early instars. Among the nursery
species is Taeniopteryx, which never appears in older stages, while
Isoperla bilineata, Nemoura 322235, E, vallicularia, and E, sinuata
can be found in all stages. However, the minuteness and large num-
bers of early instar Nemoura made it impractical to separate the
two gilled forms (E, venosa and g. sinuata).

As a generic group, Nemoura shows a slight seasonal increase
from March to mid—summer, with the lowest numbers in December. Like-

wise, Mackereth (1957) shows that Nemoura cambrica Steph. in Westmore-

 

land exhibits a pulse in mid—sumner.
The tiny Leuctra reaches a peak in June, but is present year-

round, especially in Scapania undulata turfs of Toliver Run, where

 

it exhibits a secondary peak in December. In the study of a stony
stream,‘Mackereth shows that seasonal peaks in numbers of Leuctra

differ with the species, so that some species of nymphs are most

 

 

III III llalil

112

abundant in summer, while others are most abundant in winter. The
presence of two peaks of abundance in the present study may indicate
the presence of two or more species. The appearance of an adult
Leuctra in one collection suggests that the naiad may use the moss

for emergence. That Nemouridae and Leuctra are among the most common
nymphs of mosses has been shown by EurOpean workers: Carpenter, 1927;
Frost, 1942; Illies, 1952. But in EurOpe, Amphinemoura and 25252:
nemoura may replace Nemoura.

Peltoperla reaches its highest numbers in Hayes Run, but in
general it occurs in the mats and increases from March through the
summer. Pteronarcys is uncommon, as could be expected of so large
an insect, but it can be seen clinging to Fontinalis. On one
occasion, Pteronarcys biloba was removed and replaced on the stream-
ing Fontinalis dalecarlica. If facing upstream, he entered the
streamers; if facing downstream, he clung quietly for several minutes
before he was removed and replaced facing upstream, whereupon he
immediately entered the streamers. Pteronarcys proteus, a smooth-
bodied, smaller nymph, occurs in gygroamblystegium group mats, but
not among streamers of Fontinalis.

While several Perlidae were found in Fontinalis beds, these
were never abundant.

Trichqptera (Caddisflies)

Within aquatic moss vegetation dwell several caddisfly larvae
heretofore undescribed. That these new genera should be found here
is not surprising: the habitat is a neglected one in this country;
the larvae are tiny; the hydrOptilid larva is caseless and looks like
an elmid larva, so that it tends to be rejected in the field by

caddis-seekers.

113

The hydrOptilid larva apparently is not indigenous to mosses,
for Cummins (pers. comm.) describes a larva matching this one but
occurring in the riffles of Linesville Creek (see Cummins, Coffman,
and Roff, 1966). Nevertheless, it occurs among bryophytes in
Sinking Creek (3-18), Mountain Lake tributary to Sinking Creek (3-18),
upstream Ginseng Run (5-4), Mud Run (7-29), Swamp Run (7-29), Laurel
Run (7-29), Hoch Run (7-29), and Saw Creek (7-30), but shows no
preference for any particular bryophyte species. Wiggins and Flint
(pers. comm.) feel that it is unlike any hydroptilid larva known to
them, but it must be reared to determine its adult relatives. This
larva is small, nearly round in cross section, and bears two poster-
ior hooks; these adaptations would enable it to move easily among
the bryophytes.

The other new genus is a brachycentrid, intermediate between

Brachycentrus and Micrasema. Like Brachycentrus, it builds a square

 

case, utilizing bryOphyte leaves and moss costae, but it lacks the
tibial spur of Brachycentrus. On the other hand, the larva resembles
Micrasema except for the case, which is round for the known Micrasema.
Flint and Wiggins (pers. comm.) hape that it is the undescribed larva
of Adicrophelps hitcocki Flint, known from.Appalachian adults. The
new larvae were first collected among moss mats of Pennsylvania
streams: Elk Creek (7-28, 65); Swamp Run (7-29, 65); Hoch Run (7-29,
65) under the falls and on the wood of the dam, nestled among Scapania
undulata leaves; but they only reached abundance in Saw Creek (7-30,
65), where they occupied mats in mutual exclusion of Micrasema and
Brachycentrus when the latter two occurred in Eurynchium_riparioides

mats. On August 25, 1965, they appeared in collections from Hoyes

114

Run in Maryland. In all these collections, the larvae were tiny
(4-5 mm.) and were thus suited to maneuverability among mosses
because of their size.

Since the abundance and kind of caddisflies vary greatly among
streams and bryophytes, no one group appears more important than
another. As mentioned earlier, Micrasema, Brachycentrus, and the
hydroptilids attach their cases to the moss; Paleagapetus celsus
utilizes Scapania undulata in its case. This latter hydrOptilid larva
was first described by Flint in 1962 from Tennessee and North Carolina
larvae. Its occurrence in the Deep Creek tributary (larvae: 5-4,

65; 6-11, 66), Toliver Run (larvae: 5-4, 8-25, 12-25, 65; 3-22, 6-11,
66; pupae: 6-19, 65; 6-11, 66), and Swamp Run (larvae: 7-29, 65)
apparently record for the first time its northern range. While

the larvae of this study build their cases predominantly from

Scapania undulata (not S, nemorosa like Flint's specimens), they
wander from the Scapania turf to appear occasionally among Fontinalis.
Their only occurrence in a non-§capania stream is with another

species of leafy liverwort in Swamp Run. Flint (1962a) states

that larvae from Tennessee and North Carolina were collected May

19 to June 9 and adults June 7 to July 1; the remainder of the life
cycle is unknown. That pupae appeared only in June for two years

of this study indicates this as their principal pupal season in the
Garrett County, Maryland area. Furthermore, the reappearance of larvae
in late July suggests a short pupal and adult stage unless this insect
has a two-year life cycle. But Pennak (1953) points out that it is
thought that caddisflies may have either one or two generations per

year. Thus, it seems illogical to suggest that so tiny a creature

should take two years to develop.

115

If any group of caddisflies is to be considered most frequent,
it must be Rhyac0philidae. Again, the bryophyte provides a nursery,
particularly for ghyacOphila torva and the B, invaria group. Adapted
for free movement, the case-less Rhyacophila larva has large, free,
posterior hooks and no gills. Attesting to the carnivorous habit
of the larvae (Percival and Whitehead, 1929), one ghyaCOthla nr.
carolina larva was preserved with a chironomid larva still in its
mouth. Like the TrichOptera in general, the distribution of
rhyacophilids in the middle Appalachians is poorly known, and the
following apparent state records occur in this study (see Flint,
1962b):

5, fuscula in Magyland (Ginseng Run, 5-4, 65; Hoyes Run, 8-25,

65; Muddy Creek, 8-25, 65; in Pennsylvania (Laurel Run, 7-29,

65; Mud Run, 7-29, 65; Elk Creek, 7-28, 65; Pohopoco Creek,
7-29, 65).

 

R, tgxxa,in ugrylgni,(noyes Run, 8-25, 65; 3-22, 6-11, 66;
Toliver Run, 3-29, 6-19, 7-9, 8-25, 12-25, 65; Deep Creek
tributary, 5-4, 65; Gramlich Run, 6-20, 65; Ginseng, 7-9, 65;
Muddy Creek, 8-25, 63; in Pennsylvania (Dingman's Creek,
7-30, 65; Mud Run, 7-29, 65; Laurel Run, 7-29, 65; Hoch Run,
7-29, 65; Elk Creek, 7-28, 65).

g, minors in flggyland (Toliver Run, 5-4, 65).

3, sp. 2 in Magyland (Noyes Run, 3-22, 66)--see Flint, 1962b.

3, sp. 5 in nagyland (Toliver Run, 6-19, 65)--see Flint, 1962b.

While none of these records extends the known range, they fill some
rather obvious gaps in our collecting records.

In addition to the rhyacophilids, Parapgychg apigalis (Hydro-

psychidae) appears to contribute a state record for Magyland:

Deep Creek tributary (5-4, 65; 6-11, 66); Neds Run (5-4, 65)--see
Flint, 1961. The other hydropsychids, except Diplectrona modesta,
could not be determined to species, although several Hydrogsychg

species are present. Larvae of Hydropsyche and CheumatOpsyche

 

 

116

frequently decorate the moss mat with their nets. These variable
feeders are known to eat moss (Percival and Whitehead, 1929),

algae and animals (Cummins, unpub. data), and one fiydrogsychg larva
in this study was eating a Baetis sp. naiad when he was preserved.

Another insect in the bryOphyte nursery, the Lepidostoma larva
seldom occurs in mosses when it passes its early sand case stage,
possibly due to the difficulty of maneuvering a bulky case in the
bryophyte mat. For this same reason, the large limnephilid Pycnopsyche
spp. are rare, but occasionally occur among Fontinalis streamers.

But the small limnephilid Neophylax spp. are more frequent among mats,
although still not abundant.

Philopotamids may reach high abundance, wherein Dolophiloides
distinctus appears in the mats and turfs, while the less frequent
Chimarra sterrima seems to be confined to Fontinalis dalecarlica
streamers of larger streams.

Other Orders

While OrthOptera, Hymenoptera, and Neuroptera were totally absent,
early instars of the other aquatic orders were occasional constituents.
Tiny nymphs of Hemiptera, such as Microvelia, occur in shallow waters
while Little Bennett Creek had nearly mature nymphs of this genus.
Early spring found several first instar Odonata--both damsel and
dragonflies-~in.mats of liverworts or mosses, but older instars never
appeared. (Large damselfly naiads occur frequently in the large

Fontinalis clumps of the Red Cedar River, Ingham.Co., Michigan--this

 

author, unpub. data. In these same Red Cedar Fontinalis beds one

 

can find a Nymphalidae caterpillar.) Occasionally young larvae of

Megaloptera (Cgrdulegaster, Nigronia, and Sialis) appear, but these

 

are not regular bryo-community constituents.

117

Insect Adaptations to BryOphyte Life

While the morphological adaptations of insects dwelling among
aquatic bryOphyte vegetation include many that are common among other
mountain stream insects, additional restrictions are placed on the
bryOphyte fauna because of the confined space within these vegetation
mats. Steinmann (1907) lists mountain stream adaptations (in.
Muttkowski, 1929): 1) dorso-ventral flattening; 2) enlargement
of adhesive surfaces; 3) small body compass, tendency to dwarfing;
4) attacmments--temporary and permanent; 5) by weighting (assumed
to mean ballast accumulation); 6) reduction of swimming hairs;

7) respiration--no surface breathers. Among these, several are
appropriate to bryOphyte inhabitants, while I have added the additional
characteristics of generalized feeding (Muttkowski, 1929) and lateral
compression.

Dorsal-ventral flattening

Absence of dorsal-ventral flattening exhibited by virtually all
bryo-insects is an adaptation to clambering about among the small
intra-bryOphyte spaces. That is, such mountain stream insects as
the flat Heptageniidae or Psephenidae would be inhibited in movement
by their broad, but flat, bodies.

Lateral compression

Contrary to dorsal-ventral flattening, lateral compression pro-
vides a stream-lined body for ease in movement. While Baetis dwells
in mosses only during young stages, it, like larvae of the Hydrop-
tilidae and Elmidae, has easy movement by being laterally compressed

and small.

 

 

 

118

Enlargement of adhesive surfaces

Any ventral enlargement would be useless to these insects living
among tiny branches of bryOphytes, and are consequently entirely
absent. Thus, we do not find the rapid water, rock-dwelling Blephari-
ceridae and Psephenidae among the bryophytes.
Small body

Insects dwelling in the bryOphytes cover may be early instars
that are there for only a transient period of their life cycle or
they may be permanent residents. There is some reason to accept
that small size has had selective advantage in both cases by permit-
ting ease of movement among the small internal chambers of bryophyte
vegetation. This appears especially true for the mayfly Baetis spp.,
which is free-ranging and a relatively weak swimmer (waters, 1962),
whose early instars increase to about 10 per gram among mosses in
the summer, but are virtually absent in later stages when they can
be found on the stream bed. Other early instar inhabitants include
the cranefly larva Limonia sp., the stonefly Taeniopteryx, the

caddisflies Lepidostoma and Neaphylax, and occasional Odonata,

 

Hemiptera, and Megaloptera.

Not only is this a nursery for several groups, but the lifelong
inhabitants in general are small. The most abundant insects,
chironomid larvae, accompany the tiny Empididae and heleid midges.
Five genera of microcaddis dwell among bryophytes, while among the
beetles, some of the smallest elmids (Pzgmgxggig_elgggn§ and ngig-
sgryus spp.) frequently are among the most numerous. Only on Fontin-
,glig can one find the larger elmid, Stenelmis crenata, or the larger

perlid stoneflies, whereas on the mat-forming bryophytes smaller

stoneflies such as IsOperla bilineata, Nemoura, and Peltoperla are

 

 

 

119

common. Only Pteronarcy§_and Pycngpgyche can be considered large,

 

 

and PycnOpsyche is rare, while the large Pteronarcys biloba is, like

 

 

Stenelmis, restricted to Fontinalis dalecarlica.

Its small size is the only obvious adaptation of the new brachy-
centrid larva, which fashions the bryophyte into a case.
Attachments

Whereas the ventral suckers of the mayfly Epeorus are useless,
terminal suckers on simulid larvae enable them to remain on bryOphyte
surfaces under torrential flow, while a secreted thread permits them
to change position without being swept away. Meanwhile, they attach
their pupae permanently to the axes and leaves of bryOphytes.

Caddisflies frequently attach their cases to the plants: Brachy-
ggnnzn§_spp., Micrasema spp., Neotrichia spp., Hydrgptila spp.,
Oglethira sp.?, HydrOpsychidae (a thin net retreat which may incor-
porate the bryOphyte), Philopotamidae (a tubular net case enmeshed
with the bryOphyte).

Other insects have hooks which permit climbing and clinging:

 

Rhyac0phila, elmid larvae and adults, Plecoptera, Ephemerella, Philo-

potamidae, Chironomidae, and the new hydrOptilid larva.

Weighting
Like Muttkowski's example, HydrOpsychidae build nets in bryOphytes,
while other TrichOptera are weighted down by heavy cases.
Reduction of swimming_hairs
Except for a few occasional invaders, swimming hairs are
entirely absent on bryophyte insects. Not only are they unnecessary,

but they would probably be a detriment to climbing about.

120

Respiration
Like the mountain stream insects, few surface breathers live
here. Rather, the insects frequently have covered gills (elmid larvae;

Ephemerella-~with platelike cover gills and covered fibrillar portion),

 

have streamlined gills (Paraleptophlebia), or have no gills (Isogerla,

Leuctra, Rhyac0phila, Brachycentridae, Hydroptilidae). These

 

 

adaptations prevent the wear and tear of abrasion as the insect climbs
about the bryophyte. The only example of profuse gills is on the
hydrOpsychids, which live a relatively quiet life in a retreat, trap-
ping their food with a net.
Generalized feeding

Because of the restrictions of the bryOphyte habitat, those
insects with ability to eat whatever is available would have a selec-
tive advantage. Thus, we find adaptations to trap the plankton food
supply as it flushes through the bryophyte system: net-building by
Hydropsychidae; feathery food-trapping appendages on the Simulidae.
Other insects are able to feed on aufwuchs of the bryophytes, util-
izing the bryophyte as a filter for detritus and living organisms.
I even suspect the hydropsychids of feeding on aufwuchs, usingthe
moss in place of a net, for I often found the number of larvae far
surpassing the number of nets. As already mentioned, some insects
probably eat the moss but digest only the aufwuchs. (In this study,
I observed no direct evidence of insects eating mosses.) An abun-
dance of aufwuchs is frequently present, especially the diatoms of

the genera Fragillaria and Cocconeis, while the desmids of the

 

genera Closterium and Cosmarium are less common (author, unpub. data).

 

Evidence of carnivory has already been cited, wherein tiny Baetis

spp. and Chironomidae feed the larger stoneflies and caddisflies.

CONCLUSIONS

A survey was made of the insect fauna associated with the stream
bryophytes of the central Appalachian Mountain region. This study
was planned at the outset as a survey type, and sampling was done
by hand grabs in ways that were not compatable with statistical
treatment ofthe data. Streams were sampled at arbitrary times and
at varying frequencies. With our primitive state of knowledge of
these associations, an extensive first approximation was thought
more in order than a detailed local study.

Among the 28 streams studied, three bryophyte-based stream types

are apparent: Fontinalis, the Hygroamblystegium group, and Scapania.

 

 

A Fontinalis stream is generally larger and has a continuous

 

flow of water sufficient to submerge the moss year-round. Because

of its loose nature, Fontinalis has herein been designated a streamer,

 

where one can find the larger of the bryo-insects: Stenelmis crenata,

Pteronarcys biloba, and Perlidae. But the smaller insects occur here

 

too, and the greatest variety of insect species is found by comparing

al the Fontinalis communities. Nevertheless, when Fontinalis

 

communities of a non-Fontinalis stream are studied, one finds greater

 

variety in insect species among the other bryOphytes of that stream,
as indicated specifically by Toliver Run. Thus, it appears that only

the large (dominant) beds of Fontinalis achieve a great variety of

 

insect species.

The Hygroamblystegium group comprises streams where several

 

bryOphytes appear similar and make similar mats. Indeed, their insect
faunas are not especially different, nor are the generally narrow,

121

122

shallow streams they occupy, so these streams can best be studied

as a group. These mats provide the homes for many small insects such
as Chironomidae, Simulidae, Elmidae, gigrasema, Peltoperla, and the
new brachycentrid larvae. Like several other caddis larvae, the new
brachycentrid larva constructs its case from the moss blades and
costae, but the hydrOptilid is caseless. Because of the compact

internal structure of the gygroamblystegium group mosses, only small

 

insects frequent them, but the insect variety is usually greater than
that found among other bryophytes of the same stream, including
Fontinalis.

Scapania streams can hardly be described on the basis of the
two Scapania-dominant ones of this study. But a comparison of Scapania
with Fontinalis and Sematqphyllum in Toliver Run suggests that g,
undulata might harbor an insect variety greater than that of other
bryophytes, while the number of individuals-per gram is also consider-
ably higher. Even in adjacent collections of Scapania and Fontinalis
in a dripping waterfall, the latter exhibits fewer individuals, while
the folded leaves of Scapania provide a shelter for numerous small
insects, protecting them from being dislodged by the flowing water.

Larvae of the Rhyacqphila carolina group are especially noticeable

 

in Scapania, while they are almost completely absent among other
bryophytes. Here also is the larva of Paleagapetus celsus with its
§capania case. -

Among the bryophytes in general, the most important group of
insects, numberwise, appears to be the Diptera, especially Chirono-
midae and Simulidae, while Ephemeroptera, Plecoptera, and Trichoptera

are of secondary importance. But even these secondary orders may

123

exhibit disproportionately high numbers in individual streams or
during certain seasons, for example, Plecoptera (Leuctra) in May or
Ephemeroptera (Ephemerella) in March.

Seasonal distributions might be inferred from the data on Toli-
ver Run and Muddy Creek, with support from other streams having only
one collecting date. In general, sumer (July and August) collections
show the highest numbers of individuals and species, while numbers
show a drop in December and reach a low in March and Hay collections.
By June, the numbers appear to be climbing before reaching their peak
again in the summer. Exceptions to these patterns are such seasonal

insects as Prosimulium spp., which show a peak in December but are

 

absent in mid-summer.

As indicated by the seasonal trends in sizes, kinds, and numbers
of insects, one use of the bryOphyte appears to be that of a nursery--
a substratum with protective chambers in which tiny insects are
harbored and provided with a source of food supplied by the flowing
water. Within the mats, protection against predators is effected
by the difficulty any large organism would encounter in reaching the
inner chambers, although some bryOphyte inhabitants (PlecOptera and
Trichoptera) prey on smaller insect inhabitants, as noted in this
study.

While the bryOphyte provides a nursery for some young insects,
it also provides a permanent aquatic home for other tiny species.
Adaptations to bryophyte living reflect the difficulty of occupying
a small space: necessity for compactness. Not only are these insects
small; they also may exhibit lateral compression and lack of gills

and appendages, while posterior hooks (Rhyacophilg) and suction cups

124

(Simulidae) protect against being swept away, whereas others use
nets (Hydropsychidae) and head fans (Simulidae) to filter food from
the constantly flowing supply. These food filters are located near
the flowing water of the bryOphyte surface, while the internal cham-
bers of the bryOphytes simulate a pool environment.

Because the bryophyte provides its own peculiar conditions for
the insects, which are manifest in the afore-mentioned adaptations,
the total number of species appears to be somewhat low-~150 total
for the 28 streams studied. From this low species number, even fewer
insects appear to be common ones among the bryOphytes. 0n the basis
of frequency, I would consider only about 70 to be true members of
the bryo-insect community in these streams, while most collections
have about 15-20 species, never exceeding 33.

The comparison of bryophyte communities appears to be an exciting
Opportunity for comparing a particular stream region in a number of
different streams. Based on the low number of insect species found
among these streams of the Appalachians, we might expect a higher
degree of homogeneity than for most habitat choices which have no
biological criteria. As a result of the implications of this study,
we can formulate certain basic questions concerning the relationships
apparent here:

Why do insect arrays in two species of bryophytes in the same stream
differ?

Are the differences related to the morphometry of the bryOphyte or
to outside influences affecting both the bryOphyte and the insects?
What common factors cause species arrays to occur?

How do food species relate to the kinds of insect species?

125

Is the winter increase in numbers of individuals of certain taxa due
to a migration to the bryophyte, to drift caught by the bryOphyte,
or to other factors?
Are the bryOphyte-insect communities continuous or discontinuous with
other stream communities?

These are only a beginning of the questions which need to be
answered. The problem now remains to find a satisfactory sampling
procedure and begin work on why certain relationships appeared in

this study.

l.

TABULAR SUMMARY AND CONCLUSIONS

A general survey was made of the insect fauna associated with the
stream bryophytes of the central Appalachian Mountain region (Penn-
sylvania, Maryland, and Virginia). Because this study was intended

as a survey, and sampling was done by hand grabs, the data are not
amenable to statistical analysis.

Twenty-eight streams from the middle Appalachian Mountains were
surveyed. Samples were taken from Garrett County, Maryland, streams
in March, May, June, summer, and December to show some aspects of

the seasonal picture. Other streams were sampled only once.

Certain insects are relatively constant members of the collective
bryo-community. A few insects appear to relate specifically to certahi
bryOphyte species. ,

These relationships may coincide with case-building materials, type

of protection, microhabitat occupied by the bryOphyte, or adaptation
of the insect for mobility and stability in the bryOphyte.

Certain bryOphytes coincide with certain stream types. BryOphyte

to stream relationships may depend on width, depth, speed of flow,
permanence, or habitats within the stream. A total of 25 bryOphyte
species was observed in this study.

The number of species of insects collected in aquatic bryOphyte
vegetation in streams of the middle Appalachians is relatively low and
ranges from 1 to 33 in an individual stream with 150 in all.

The number of insect individuals per gram (dry weight of bryophyte)
varies from .3 to 2887 in this study. The Chironomidae are usually
most abundant, while other abundant insects include Promoresia elegans,

l26

8.

10.

ll.

13.

14.

127

the Simulium tuberosum group, the Prosimulium hirtipes group,

 

 

Leuctra, Nemoura spp., lsoperla bilineata, Ephemerella spp.,
Baetis, Micrasema spp., and Pericoma.

Presence of certain bryophytes might be usable as an indicator
of the probable presence of certain kinds of insects.

The similarity of insect bryo-communities is high, if compared
by bryOphyte species groups.

The insect bryo-communities of streams with the same bryOphyte
dominant usually have high community coefficients,‘but other
factors may cause a greater similarity with a stream having
different bryophytes.

In the bryOphyte samples certain insects appear to co-occur,
while others appear to be mutually exclusive, but the study was
not designed to verify this.

Some insects occur with mosses only in young stages.

The known ranges of some insects are increased, while other
insects are new state records.

These collections have brought to light some apparently formerly
unknown larvae of insects. These are Trichoptera in the Brachy-

centridae and Hydroptilidae.

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APPENDICES

133

134

In tables XII-XVI the following symbols represent the stream names,
while the Roman numerals are the collection numbers.

De Deep Creek tributary

Di Dingman's Creek

El Elk Creek

2i Iinseng Run
20 Goose Creek
3r Gramlich Run

Hk Hoch Run

Hy Hoyes Run

Jo Johns Creek

La Laurel Run

LB Little Bennett Creek

Md Mud Run

My Muddy Creek

Mt Mountain Lake tributary to Sinking Creek
Ne Neds Run

Pi Pidcock Creek

Po Pohopoco Creek

Py Piney Creek

Ro Rock Castle Creek

Sa Saw Creek

Sd Sideling Mill Creek tributary

Si Sinking Creek

Sw Swamp Run

To Toliver Run

Tot Toliver Run tributary

WBe Seneca Creek tributary at Bell's

WBu Seneca Creek tributary at Burroughs'

 

-————“_ #--

13S
MARCH COUNTS 0F INSECTS PER GRAM DRY WEIGHT OF BRYOPHYTE

TABLE XII .

 

 

 

 

 

 

 

 

 

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assassins”. 76813 1 7 629852 0
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Bezzia group 2 L

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Egppinalie
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HoCCCXV 1.2 1.2 137.2
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.9
.8
.3 1.7
.7
“.8

E

Diplectrona.modosta L

 

.1

' 3.5

2.9

Maggot L

 

 

 

.1
s 3 A
E a a
O O
z: “i *4 ‘° 8
-» 5 dl :3 g
0 In. H H H
saw .2 2 z: '3
W o e s
a 3 g: s 3
|
Is #1 .4} £? 32
.3
.03 _
3-5
1.0

.2

figgroambly—
ategium
fluviatile
SBuLI
SBuLII
SthXXVIII
SthXXX
SthXXXI
Av.
£25221!!!“ -
mega-g
tuna:
s1xxv
SiIIVI
SiXXIII

 

 

Anhlyategua
zgrigg

-—4—.

812111

 

13.919.933.12!
riparioides
GrLXIX
BoXXXII
Av.
BrachythQQLUB.
plumosum
305(va 11 "I"
GrLVII
GrLIX
SdLXXY
SdLXXVI
Av.
Bracgytheciun

 

Tipula collaris L

 

rivulare
GrLIV
SthXXVII

RoXXXVII
Av.

Brzhnia

291292951192

SBoLIII

 

 

F:

m

(0

+3 0

H

F: r4

0

o 0
04

° 3

d 0

«4' H

N +3

8| °

pm $3

.3

.1 .1 .02

.2

“.5

Diplectrona modesta L

 

.2

1&2

Maggot L

.2

1.9

 

Palegggpetus celsus L

 

 

 

r-J A
"I g]
.. 53. s
.3
3 a 3 8'
S :3 :1 73*
'84 '84 84 a
8 8 2 o
5 S 3. 8
.4 r3 a: 04
.1 .02
2.2
ms
2.3
1.2
.2
.3
.08 _ .03
.2
.7
.2 .1

 

2
I4
d
U 0
H A O
:3 ... :3
H d
H 0 O
'O ‘3!
o In d1
0
G a ‘
,1 ya V‘
31 3 ‘°
.4 0 3‘
E4 m m

 

 

9210919
aloicola var. rivularis
SdLXIY1I

Ezszghypgaa
191192!
Doccxcx
DeCCXCII
DoCCXCVI

Undetermined

DeCCXCV

Sciaromium
1pscur11
LBXLVIII 3.9

Samatophyllun
Egrylandicum

ToLXXIII

TOLXXIV

 

 

Undetermined

ToCCCVI

§£§p§915
29421112
ToLXII 1.7
T000011
ToCCCIII .6
T000017
ToCCCVII
TOCCCVIII
T000011
ToCCCX
ToCCCXIII
DeCCXCIV
Av.

P‘#‘
O O
td-o

Undetermined
GrLXII

113'

 

,4
d
.p
.1
o
'0
O
a
S
8 ya
a
o .9
o o
H
a g
c:
.6
1.1
1.3
.05

 

Paleagapetus celsus L

Lhmnophorua L

 

 

Limnophila L

Hydroptilid gen. 1 L

Prosimulium

um L

 

Epntipalts

delgqarlige
ToLXX
Tocccv
TOCCCXI
ToCCCXII
Doccxc
DeCCXCVII
MyCCXCVIII
MyCCXCIX
Hy000
MyCCCI

Av.
Fonting110_

1199210;
GrLV

slime-.212-
etesiee
fluvigtilo
RnXLIV
RoXLII
BoXXXIII
BoXLIII
ROXLV
ROXXXVI
RoxXXIX
BoXL
R0XIXIV
RoXXXV
HoCCGXIV
HoCCCXY
HoCCCXVI
HoCCCXVII
HOCCCXVIII
DeCCXCIII
GrLXIII
SiXXIV
LBXLVI

 

 

 

A
a! 54
...4
h d
s a
C! O
0H «9
d d
.4 H
:3 2
3 3+
0 0
g: d
a :3
.1
.02
5-9
.5
1.2
1.2

Lepidostoma L

.02

Atherix variegata L

 

.1

1 L

Dan helea sp.

1.5

2.6

 

 

:3
2'
Adi-1
o 43- .
.c: d
o P.
0%:
Dd w a
o a o
.3 s :2
g; E! E?
.9
.9
.2
.5 .
1.5
1.0
.3
.3
.5 -5
20.3
1.0

 

Cneghia mutata L

.1

Cnephia.nutata P

 

.02

EXEFOEBbl -
atssiye.
1111.11.01: 11:

3mm:

SBuLII

SthXXYIII

SthXXX

SthXXXI
Av.

EIEEQaQDlz-
ategium
tenax

sxxxv

SiXXVI

s1xx111

Agfilxategium
x0112;

SiXXII

EEEXHEhlE!
riparioides
GrLXIX
RoXXXII
Av.
Breehxthggm
21999322
Roxxxvrll
GrLVII
GrLIX
sanxxv
SdLXIVI
Av.
2209331229100
leniezg
GrLIV
SthXXVII
Roxxxvxl
Av.
£21001;

---.-

SBQLIII

 

Rhyacophila invaria L

2.3
1.2

1.5

3-5

1.2

 

Egyacophila torva L

.2

115

 

 

.4
I!
.p
d
6.0

1—1 0
Ir.

E E

.p

I: H

o «r.

“U h

H 0

e a

.3 4

001 005

.2

.1

.2

.03

.3

.6 1.3

.3 .L’

 

 

.4 :3
H
o A g 1-4
9. 0
m .3 ~u u
a. o a E
o p. a u
r! a o o
0 O. m 0
.d o a o
h u .4
g 'd .3 0:
n a? z: 8
.1 C2 .1 .1
2.2
2.8
-3
.2
1.9
.6
.7
.6 .6
.4 .2

Cngphia mutate L

 

 

Cnephia mutate P

 

Rhyecqphila torva L

 

Rhyecophilg invaria L

Lepidostgg§,L

 

9210239
ajnieola var. rivularis

SdLXXVII '*"”""

32522212222
1uridum

DeCGXCI

DQCCXCII 0.3

DeCCXCVI

Und e te rmi ned
.DoCCXCV
Sciaromium ‘

leggurii
LBXLVIII 2,1

 

§eee§22§lllae

merylendicum
TOLXXIII
ToLIIV

Undetermined

 

T000071

§geeeeie uedulete

TOLXX1 1.3

T000011

T0000111v .6

T00001V

ToCCCVII

ToCCCVIII

ToCCCIX 1.7

T0000! .9

T0000X111

DoCCXCIV 1
Av.

IO.
U'IOU‘

Undeterm‘ned

GrLKII

Atherix verie eta L

 

1h6

e helee sp. 1 L

Hgdroggycho L

 

Micrasema (tiny L)

Cnoghie mutate L

 

Qgtiosorvus L

 

9.1.1.111. mutate 1'

Pteronarcys pgoteue N

 

antinelig

gelocarlicg
TOL11
T00007
T000011
TOCCCXII
D00010
D00010711
MyCCXCVIII
My001011
MyCCC
My0001

Av.
Zontigglip

flaccida
GrL7

3ygroenbll-
stegiun

fluviatilg
301L17

RoXLII

30111111

301L111

301L7

3011171

3011111

301L

3011117

301117

30000117 2.1
3000017

30000171 .9
300001711
3000017111 3.5
DoCCXCIII

GrL1111

$11117

L31L71

cebri enniu L

cnooa h

.1

.02

Baetis app. 3

.8

11.7

 

Pycnopeyche scebripennia L

 

Eteronercys groteue N

Ersroamblz-
staging
Zlnviatlle

SBuLI

SBuLII

$1Mt117111

Sth111

Sth1111
Av. .2 .00

Elggggmbl -
sipsige
1291;

51117

$11171

31xx111

Anblyutggiun
Eerlye
811111

 

Eurynchium
riparioides

GrL111

3011111

 

 

ggechlthecium
2129222!

301117111

GrLVII

GrLIX

$03117

SdL1171

 

bresbxshegige
rivuleae
GrLIV
$1Mt11711
30111711

Bryhnia
epvegtegglleg
SBoLIII

N

D.
0c

etil 0

£l'

.1

1h8

 

.4
"I
...-Q
2: 5'
0|
m D.
3' 1:
..a ..
8i 8
ml
fl 0
a ‘5’.
£1 55
u’ a
'3, 0
AJ 3?

 

92.111.191.41
algicola var. rivularis
SdL11711

516?.9111233!
1111‘. 151.1111!
D000101
DoCCXCII
DoCCXCVI

Undetermined
D000107

510.133.911.199.
1.999.111" 1. _1
LBXLVIII

§ge§9mzllum
merylendicun

TOL11111

TOL1117

 

Undetermined
T000071

fieapeeie
yedeleie

TOL111 .3
T000011
ToCCCIII
T000017
ToCCCVII
ToCCCVIII
T000011
ToCCCX
ToCCCXIII
DeCCXCIV

Av. .02

Undetermined
GrLXII

PP- N

3.8.9113. a

11.9

150
MAY COUNTS OF INSECTS PER GRAM DRY WEI§HT 0F BRYOPHYTE

TABLE XIII.

 

a £823; £80.:

a macaw
daudpnd dHHoucacnmw

 

m Noamaoo
831.2 53.5340.

 

A Noamaoo
oomwamwm.aaHH:-«uoum

 

 

a wmwodoa

21mmmammwa.umdqqu
.Hon«.wwmdamm

m Hoamaoo
an». eagle. £5 8.0.. H25 m

A Moamioo
3%

m cavaaoaouano

A oacwaoqoufino

 

dalecarlica

D0071

 

Fontinali-
T007113
T0011

MyGXII

2.5

3001111
PyCXVI
Av.

2.0

2.5

3.7 21.8

37.
16.

 

fluviatile

ate ium
Py01711
D007
01017

—-...-

31 roemblz-

7376
o o o o
2.462

-4
“8
.4.“ A”

.U
. z;

1
I
1

b.019.n.7>0
o 0 o o o 0

9016.67
1

1
1 0
O 0

121005

c o o o

9 5
O O

1
0

2

5

10.0

1%.“

10.6

010117
GrL11117
Av.

AVKJQRU
0000
1.va<J<J

b.274nw

10
munw1f1

RVRVULU
0000

11..

 

riparioides

 

Eurynchium

Av.
Brachythecium

GrL1117
GrL11171

Gr1011

6.3
7.9

1.0

1.7

1.0

NC

—=—.—

 

 

_ygoeum

GrCII

lggcurii

91
T0t011

Av.
§capanie

undulate
D0017

Av.

 

§91ezom inn
LBLxxu
To cvxu
T007111

LBLXXX

151

‘—

 

 

Promoresia elegane L

Promoresia elegans A
Athe rix variegata L
EgyacOphila invaria L

 

 

 

Cnenhia mutata L

 

Cnephia mutata P

 

Peltoperla H
Lepidostoma L

 

Pericoma L

 

Fontinalis
dalecarlica
DocVI 2

ToCVIIB
ToCIX
MyCXII
NO0X111 . .
PyCXVI
Av. .6 .

Hzgroamblz-

ategium
fluviat110

PyCXVII .
DO0V
GiCXV
GiCXIV .
GrLXXXIV
Av. .
Eurynchium
gigggioidgg
GrLXXXY
GrLKXXVI 1.7
GrXCIX .5
Av. .7
Brachythecium
glgmoeun
GrCII ' .7

 

0 0 0
U‘Ufl
N O\ NUH

29.9 .7

6.1- .1 .

 

.7 .7 19.6

7
.1 .1 .6 1.5
1 1.3 .2 3.6

NH
HU‘Q

O

\O

\I
O
5.0
O
\J
U
0
hr)

602

 

 

§9}§;92122.
1:999:11
TotCXI .9 5.9 .6 1.0 .3
LBLXXIX 1.0
LBLXXX .4
Av. .3 .3 2.0 .2 .3 .2
§992§91§
gagglgie.
nocxv 3.6 .3 1.3
ToCVIIA .5 .1 .2 .1
ToCVIII .2 .2 8.3 .2
Av. .9 .2 .08 .05 .u 2.8 .1

NWkoaJN
OOHQ

501110.8r0up 2 L

let
(nor-o

Fontinalis
gagggarlica
DeCVI
ToCVIIB
ToCIX
MyCXII
NO0X111
PyCXVI
AV .
Hygtoamblzr
gtegiun
fluviatilg
PyCXVII
DO0V
GiCXV
G10XIV
GrLXXIV
Av.
‘Egzxgphigg
119.9319 149...!
GrLXXXV
GrLXXXVI
GrXCIX
Av.
Ergchythqgiys
21299990
GrCII

 

Sgiargmign
lagggzil
TotCXI
LBLKXIX
LBEXXX
Av.

309211.212

......—.-_— ._..

TOCVIIA
ToCVIII
Av.

 

2:
d
fl
‘4
A a
N '3
O
0 A v4
0* H
w H H
a: 3
0 94
H a Q
o 3
u: 'U
I"! 0
g 94 %
an a z
.8
.8
.2 1.0

.014 .2 .2

1.1»
.2
1.3
11.1
2.3 .5 .3
.9
-3
15.6
.6
.2
2.3
.6

Neophylax concinnus L

 

152

Ontiosorvua L

‘

 

.03

Paleagapetus celsus L

 

.03

 

 

 

A
d
.p
I
0
'6
2
0-2 Ila H 2
.3 .53 6°. 6
q-a ad 0 H
+3 09 43
9. O4 6 O
O O «4 0
36 a s '6'
6' a :3 al
.3
1.1
.3
.2
.06 .15
.9 .3
.2
.1 .2
.4
.3 .09 .1
.3
.a
.1

 

II. I 1"!

19:13.1. n91.1-
legggrlica

DO0V1 ‘
ToCVIIB
ToCIX
MyCXII
NeCXIII
PyCXVI
Av.

flzargwblz-
stagiun
fluviatile

Cheumatopaycho L

 

PyGXVII
D007
GiCXV
GiCXIV
‘GrLXXXIV
Av.

131.131.159.113;
leggioidao

GrLXXXV

GrLXXXVI

GrXCIX
Av.

Erachythoag

 

211122929;

GrCII

5010100101
10003211

TthXI
LBLXXIX
LBEXXX
Av.
Scanggig

00001010

D001V
ToCVIIA

TOCVIII
Av.

 

.2

.04

oaervus sp. 2 A

Optioaervus 0p. 1 A

 

Opti

153

 

 

'2:
d
..4
,o
0
H
E
_O
4..
9.
0
H
a!
H
d
a.
.2
.02
.1
.2
.7
.05 .1
2.2
.6
.2
.05

 

A
a!
Z
O
'” 6a
a (v
H
fl 0
'5.“ g
OE
8'33
5'8 3
an IE
.3
.7
.2
1.2

06
06
.2
.5 .3
.2 '3 003
.8
-5
.2 .3
.3

.15

TAPLE XIV.
..J
g
'0
1%
a
S
E
c)
Eonténalip
' dalecarliga
ToCXXVI 220.9
TOCXIVIII 101.0
ToCCCXXXVIII 23.5
TOCCCXXXIX 31.7
ToCCCXL 79.2
TOCCCXLI 51.3
MyCCCXLV 92.1
MyCCCXLVI 9.6
MyCCCXLVII 16.2
MyCCCXLVIII 59.8
MyCCCXLIX 13.0
EyCCCL 28.6
MyCCCLI 36.5
dyCCCLII 46.9
DeCCCLIII 190.9
DeCCCLV 50.0
Av. 73.3
fiygroamblzf
' etegium
fluviatjlg
HOCCCXXXV 107.0
HoCCCXXXVI 211.2
HoCCCXXXVII 96.9
Av. 151.7
Eurynchiug
"ripnrioides
GrCXXX 122.8
GrCXXXII 1282.1
AV. 702.9
Sciaromium
loacuni}
LbCXL 176.2

Chironomidae P

9.5)

1...!
Com #7
O O C

NQGDOb-‘i-‘C'J‘x

JUNE COUNTS OF INSECTS MIR

complex L

Ejag-33.2101um tpbflrosum

N
"

TN A)
O O

h)

H
00::

I\ 1*
1‘--

~4§UN DRY \«IEI'li—i'l‘ OF
753 A
a} ((5: w
6? '2' 2'
2’ a! 3
01 0 14 PS
.3! :4 w: oi
53:04 a." .

49‘ C 0.": dl
{is 1‘! ...-t
9'0 ,0! u’
:lr4 win} 0
.0 h u. i h.
r-i: a :1 “l O:
:‘Q 9} a:
.5;0 5; 2}
)1 73! 0-4!

. 6
1.5

1.5 .L‘V
.9 3.7
.9 1.8
.8 6. 5 1.5
.5 2.8
16. 3
1.6 19.3
4.3
.2 6.5
9.0
2.1 7.0
2.0 4.1

10.5

13.8
.0 0.7 3.8
06 5. (J 2401‘
1(‘J90 2
.7 1.7 99.9
.4 2. 5 109’). l

v) u
DR 1‘

H

H

h)

p
O O
'11 um L‘kn

CFOHH

N

Ox C\
C C

\O ‘J‘~

\] C:-

\1

r‘
J .

L1 ‘\

fi<firo

KP-

\

xrn'w
1‘11113;

80p.

3911§

{NI

...:
0
‘CUQI‘JC’

H
N OW‘JKDUKQ

u)
0
L‘ ‘31

0
3'6

(3

\014-0
U

N.
Q.

“

10000rla b111neqta N

H
U‘ikhkh

and C\

7‘.)
14

11.2
3 0 (3

0.8
6.6

155

z

 

damoGAHAp «AhomomH

 

z .mmw unpomm

 

waxwoam wawouoaoym

 

unawoao wmmwhoeoum

Ill
1.,"

duadnam w wwocob

'Oll'lllllllull

.AonA 0000802

m NoAmaoo
as. 0.393 83 Adam

 

A KoAmaoo

'9‘!

 

 

.mwmouop » mdAAsaAm

z wuposwA

m owc«50cogfigo

A mwdwaocouano

3.6

7
O

I.» 4 0
0 O
1

115.4

0000

13 2

3J32/m61u50.6b.
law/.ZB/Amljon
2312 113
/O.17b359.
8%an5uhw

2.4 2h.

970u51a173
. 0 0 o 0 0

87
37.0
1.5.1
153.6
18.7
116.2

unifilgfig

Scapania
ToCXXI
ToCXXII
ToCXXIII
ToCXXIV
ToCXXV
ToCXXVII
ToCCCXLII
ToCCCXLIII
ToCCCXLIV
DacchIV
D0000LV1

.#

8.9 19.h

.6 95.7 81.6

Av.

 

I'IIII
[‘

156

_1«__2£§

3gg hirti
lex L

U.

r-‘-
' E

_l 0
:33: g‘

 

Egliegeilé N
1 L
ggpphila carolina L

Empid sp.
2;:

r0

C

04

§9n§1n§l1§

galecar119§_
ToCXXVI .6 .6
ToCXXVIII
ToCCCXXXVIII
ToCCCXXXIi
ToCCCXL
ToCCCXLI
MyCCCXLV
MyCCCLXVI
MyCCCXLVII
MyCCCKLVIII
NyCCCXLIX
MyCCCL
MyCCCLI
MyCCCLII
DeCCCLIII 7 7
DeCCCLV 3.1

AV. 1 8
fiygroam 513-

staging

ff 111V 1:,1t1]:§_
HoCCCXXXV 2} 2
HoCCCXXXVI 13.9 .5
HoCCCXXXVII 17 0

Av. 1b 0
Eurynchium

riparioides
GrCXXX 1,0
GrCXXXII .9

Av. 1.0
Sciaromium

lescurii
LBCXL 2.u

—_

2 A

_PE§

-1-.-

heryx N

group N

L

._.._.._- ...—.--—-~—_.___. ._____.‘_..__

comulex P

 

fl

Ontioseyyus ea.

__‘.--

iésgéggfi
Leoidostoma L

_h

ngsimulium hirti
themerella invaria

..—

MWHH
\JO‘inJP-‘C‘

P‘F‘OD
GIH‘C’

157

lrll 11 ll"

< N .mw md>wouofiamm

 

A dumweaud> Rapunpd

 

z Hmwoamowcome

z mfioum
aduapnd «Aaouoacnmw

 

m NOAmaoo
womauhds anAAdEAmoum

 

A NoAmaoo
momfigufis asfiflsaauomm.

 

 

A mmAAouwo aAAnmooahmm

q H .6. camam

 

z .Huomoaflom

-.

Seamania

--~.

 

undulata
ToCXXI

ToCXXII

h.8

.4

.4

1.0

1902056
.0900.
25231.“

1981
000.
12 1
1 23
O O.
1 16
I I
IV I
mxmm
cm...
0000
TTTT

ToCCCXLII

ToCCCXLIII
ToCCCXLIV
DeCCCLIV
DeCCCLVI

1.2

1.2

Av.

p. 2 L

Micrasema e

Eontinallg
dalecarlisé
ToCXXVI
ToCXXVIII
ToCCCXXXVIII
ToCCCXXXIX
ToCCCXL
ToCCCXLI
MyCCCXLV 1.9
MyCCCXLVI
MyCCCXLVII 3.8
MyCCCXLVIII
MyCCCXLIx 2.1
MyCCCL
MyCCCLI .8
MyCCCLII .7
DeCCCLIII
DeCCCLV
Av. .4
8mm 93191!"
91.05323
{122162112
Hocccxxxv
HoCCCXXXVI
Hocccxxxv11
Av.
Enrxnphiug
ripazyqugeg
. GrCXXX
GrCXXXII
Av.
§ciaggmtyg
19922111
LBCXL

.2

O O
...:

158

 

 

A
o
:1
p
0
C2
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p
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group 2 L

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3.0
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4.8

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quicoma L

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R

 

Stenelmis crenata A

 

159

 

 

A wbuou wAAnmmudmmm

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MyCCCLII 22.u
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9226192.

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21932121199
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sp.
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NC
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161

 

 

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162

SUMMER COUNTS OF INSECTS PER CRAM'DRY WEIGHT OF BRYOPHYTE

TARLE XV.

A. .6...
muasohomouuhm

 

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A A .6. 6.69m

 

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4 mQQMMAo «Amonoaoum

 

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m eddAfioaouwno

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1. 5 1. 309.2 2 .51 3... 158 Am :.J 5 9”
O O o O. O O. 00 O
1. 7. .4 1J7.9.1J 7. .1,6 0.1. 6.1.1. Am 0. o.
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1.9J9. 9. .u.1.9. 1.1J 90.1.1 1.9.6.1.9.9. OJQJ ,0
1. 9. 1. (J
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163

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A onohumondwm

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1
22
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23
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032
227
503
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153.2
493-5

ToCCXLII
ToCCXLIII
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376.0
317.0

337.7
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3 55 ..J 6 8331 55
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2.9 95.0 9.5 20.0 .03 5.6 2.1 11.8

306.0

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3321222129
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GiCL 158-3
GiCLI 45-4
GiCLIIA 61*-8
GiCKLV 125-7
GiCXLVI 144-“
HyCCL “62.1
EyCCLII Big-6
3&ng” IA 207.9
DiCCXXIV NC
DiCCXIx no
Diccxx no
DiCCXXI NC
DiCCXXII no
SaCCXVIII NC
Av. NC
Sciaromium
lescurii
1200x111
PoCCXI NC
PichI 190.5
PicXLII 296.9
21011111 211.1
PicXLIV 103.7
Av. 100.?
Hygroggpnnn
luridum
MyCCXXXII 287.1
Myccxxxxll 222.4
r0 pnun
00:30:22
SaCCXVII NC
Fontinalis
antipyrotica
var. gigantea
SaCCXV NO
Undetermined
SaCCXIII NC
achCVII no

Chironomidae P

H
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N

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110.8
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2.9

57.7

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2.8

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1.5

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25.2
62.1
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510

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166

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25.4

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group N

 

 

 

Bezzia group 2 L

W...-

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GicLIII
JoCLXXII
ToCLX
ToCCXLVI .3
SwCXCVI .u .b .4 1.1 .4
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MuCXCI .5?
LaCC .2 * .5
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HkCCII .2
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MyCCXXXV .2 5.
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MyCCXXIVII .
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GiCLIIA 2 2
GiCXLV 1 u
31QXLVI 11.1
EIyCC L 2 3
HyCCLII .8?
PoCCIX .7
thLXXXVIA
DiCCXXIV
DiCCXIX
DiCCXX
DiCCXXI
DiCCXXII
SaCCXVIII
1v. 1.2
§913£99£29
lescurii
LBCCXXIX .9
PoCCXI
P10XLI
PicXLII
P10XLIII
P10XLIV
AV 0
Eysrobygyge
luxiéma
MyCCXXXII
MyCCXXXIII

Hygrohypnum
2232292!
SaCCXVII

 

...—”w-—

aim/Hus:
var. giganggg
SaCCXV

Undetermined
SaCCXIII
SwCXCVII

1

.

fiqzzié 83-

Paraleptophlgbia spp. N

 

H
N99.
\JCh Oanxo \ntoah

waxoooom

1.6

2.1

\ICI)
O\'\)

Diplectrona modesta L

 

\n

2.3

170

group N

 

‘Eghemerella invaria

Empid sp. 2

Hydroptila L

.05

O O
U"

 

 

n
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1.8
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1.9
1.0

2.3

EgéLti na 1131
-ileicar gal.
G10 I B

 

G1CLIII
JoCLXXII
foCLX
ToCCXLVI
SHCXCVI
SVCXCVIII
DiCCXXIII
MuCXCV
MuCXCIII
MuCXCI.
LaCC
LaCCI
HkCCIV
HkCCII
SaCCXVI
SaCCXIV
PoCCX
PoCCVII
PoCCVIII
MyCCXXXIV
MyCCXXXV
MyCCXXXVI

Myccxxxv1I
MyccxxxvxII

MyCCXXXIX

MyCLIV

MyCLV

MyCLVI

MyCLVII

MyCLVIII
Av.

171

 

 

 

 

 

..q
0—! C6
<4 :1
O H
A a: :1 H
.3 14 o o
N a! no 3.
c: o a!
. 0 £1 '6 A D. U
94 H O H
w o :x. h 'l as
I” 49 d «j r4
«3 en .0. :3 3.. L. «H
a «4 o 0 o o .c:
0 E +2 o .C: .2 D.
a: '3 . .2 2+ 3 s
9.. r1 5 o :2! c: d
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«4 0 .21 H t
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.2
2.5P 17.5
18.5
1.1
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4.0
.2
19.b 1.1
1108
.u .1
.2 .1 .1 .1
.1 2.2
09 .6 02
09 07
1.0 1.0
1.9 3.8 3.2
1.0 .2
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2.2
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Rhyac0phila fuscula L

c."

o
\A’

um

.1

 

Ephemerella attenuata N

O\OH

_t2xg 111623 N

Pteggna

-———_-_

g 0
rnv

Micraeama up. 2 L

wag
12.311.11.312

ToCLXII

ToCLIX

ToCLXI

ToCCXLI

ToCCXLII

ToCCXLIII

ToCCXLIV

ToCCXLN

HkCCV

HkCCVI

HkCCIII

LaCXCIX

MuGXCIV

MuCXCII

MuCXC
AN.

Hygroamb1z§
5225135.
fluviatile

HyCCLIII 1.1

HyCCXLVII

HyCCXLVIII

HyCCXLIX

ByCCLI 4.8

EICLKXXVII

EICLXXXVIII

EICLXXIVIB .6

GiCXLVII

GicXLIX

Av. .5

 

172

 

 

H
4 0
3r .. 8
a no
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11.4
1.5
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.02 .3 1.“

hora: L

 

 

0-1
3
I:
8
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‘6 E
D ‘5. g
g 8 3
:3] § 5'
3.7
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5-3
-5
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.05 1.3

fuscglg L

 

”02:11.19

"1
f

Ephemere11§ attenuata _

 

 

Pteronarcys biloba N

 

173

 

Stenqhgig crenata A
Cheumatquyche L
Brachycentrid gen. 1

bu"

 

Micrasema sp. 2 L

EBFXFEEiEE.
ripgzigiéeg

GiCXLVIII 1.4

GiCL .5

G1CLT .JL

GicLlIA

G1CXLV

ciCXLVI

HyCCL .8

HyCCLII

PoCCIX

RkCLXXXVIA 18.6

DiCCXXIv

D1CCXIX

DiCCXX

DiCCXXI

DiCCXXII

SaCCXVIII

AV.

0\

0x2»
C‘I—‘OO\NC'¢

Sciaromium

lggguzll
LBCCXXIX
PoCCXI 1.5
.RicXLI
PifiXLII 3.1
PicXLIII 3.7
PicXLIV

Av. .3 .8

@1529112229
lgriqwg
MyCCXXXII
MyCCXXXIII 1,1

fiygrohypnum
agracggg
SaCCXVII 6,4

Eggyinalis
anttgyretica
var- 8158.11.19.99
Saccxv 11.4

Undetermined
SaCCXIII 4.1

SWCXCVII 1.5

L

tha
one

 

...A._ ._ __

acouhila carolina L

Limnoggggg: P
P

.2
o
.0
.2 .l
7.4
2.5
.2
2.1

211292 N-

C28

a ——-———-.‘——-—m- H_~ __._-.——

Pterqnar

“..A ——

fi‘

Rhyacophila fuscula L
xphemerella attenuata R

mam-“-

 

o tmahsi
0.153%

GiCLIII
JoCLXXII
ToCLX
ToCCXLVI
SwCXCVI
SwCXCVIII
DiCCXXIII
MuCXCV
MuCXCIII
MuCXCI
LaCC
LaCCI
HkCCIV
HkCCII
SaCCXVI
SaCCXIV
POCCX
PoCCYII
PoCCVIII
MyCCXXXIV
HyCCXXXV
MyCCXXXVI

MyCCXXXVII
MyCCXXXVIII
MyCCXXXIX

MyCLIV
My CLV
MyCLVI
MyCLVII
MyCLVIII
Av.

ethira L

O

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2 °: 3
9. a «H
p: z: z:
2.2
-3
18.7
2.5
2.8

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1.2L
.01

 

Pteronarcys proteue N

§9apa91§
undulata
ToCLKII
ToCLIX
ToCLXI
ToCCXLI
ToCCXLII
ToCCXLIII
ToCCXLIV
TO 5‘. CX 11V
HkCCV
HkCCVI
RkCCIII
LaCXCIX
MuCXCIV
MuCXCII
MuCXC

Av.
Bygroambl-x-

staging

§1n34a§112
HyCCLIII
HyCCXLVII
HyCCALVIII
HyCCXLIX
HyCCLI
nchxxxvzt
ElCLXXXVIII
EICLXXXVIB
GiCXLVII
GicXTJX

Av.

thira L

Oxye

 

4:
H
d.
m
a. a
:3
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HI 0
.2? a;
+9? 0.
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>4 9'
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ol 0
3.8
. 5
.5
.2
.3
1.1
1.8
.3 .2
2.1
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175

themere1la catawba N

9112 L

1.8

Wuhemeralla deficiens N

.

 

Yemoura spp.N

13.0

SP

“...—-——_.

Microvelia

 

oroteua 9

L_.—.-__

raphia A
rcye

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a.
O:
H. ‘4’
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3' 8|
Q 04
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.1
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1.7
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1.4

176

PP- N

 

ethira P

 

Nemoura s

 

Ephemerella deficiens N

themerella catawba N

 

O ethira L
Qggioservus sp. 1 A

 

9m
Dixa L

Eprxnchium
ziggzioiéeg

GicXLVIII 1.9
GicL 8.3
GiCLI
GicLIIA .4
GiCXLV
GicXLVI
HyCCL .8
HyCCLII 2.8 13.8
PoCCIX
EkCLXXXVIA
DiCCXXIV 3
DiCCXIX 2,
DiCCXX ' a
DiCCXKI
DiCCXXII 3
SaCCXVIII 10,

AV. .2 .3 1.3 2

figierggigg
12599211
LBCCXXIX
PoCCXI
PicXLI
PicXLII
P1GXLIII
PicXLIV 1.2 11.1
Av. .1 .9
1.13522121an
lgridum
MyCCXXXII
MyCCXXXIII

Hygrogxonum
QQEQQQEQ
SaCCXVII 2.1

 

Fontinalis

ggtigxretica

var-.filfiflfififié
SaCCXV .5

Undetermined
SaCCXIII
SwCXCVII

 

P.

 

Ziezgygrcva gzoteus N

Egpézegéie‘fi

Microvelia s

 

UU

KAN

177

L
Chimarra aterrimg L

Hydroptilid gen. 1 I-

 

?aleaggpotfis celsus P
Micrasema sp. 1 L
Paragnetina sp. N

 

Palggggpetus celsus L

 

BhlecOphila invaria L

Neotrichia P

 

Caenis N
Neotrichi

—-——_—_

Fontgnalis

Gigirigagliga

GicLIII .3

JOCLXXII

ToCLX

ToCCXLVI 1- 0

SVCXCVI

SVCXCVIII

DiCCXXIII

MuCXCV

MuCXCIII

MuCXCI

Lacc

LaCCI

chcxv 1- 3

HKCCII .2

SaCCXVI

SaCCXIV

P0001

PoCCVIII 1 9

MyCCXXXIV

Myccxxxv 3-
1

 

:-
E»

10.1

MyCCXXXVI

Myccmvu

MyCCXXXVII I . 1 -

MyCCXXXIX 1-

MyCLIV

MyCLV .

MyCLVI

MyCLVII 1.

MyCLVIII 2.
AV. 0 03 o

2.6

1.0

.2 .5

5
1
3 .2 .2 .
04 .02 .4 .05 .2 .07 .1 .06

 

Bhyacophila invaria L

Scagania
mm

ToCLXII

ToCLIX

ToCLXI .5
ToCCXLI

ToCCXLII

ToCCXLIII .5
ToCCXLIV

To COIL? . 2
HkCCV .3
HkCCVI

HkCCIII

LaCXCIX

MuCXCIV

MuCXCII

MuCXC

A7. .151

Hygroamblx-
stegium
fluviatile

HyCCLIII

HyCCXLVII

HyCCXLYIII

HyCCXLIX

HyCCLI

EICLXIXVII

EICLXXXVIII

ElCLXXXVIB

GiCXLVII

GiCILIX
Av.

 

Caenis H

N trichia L

Neotrichia P

178

 

Paleggapetus celsus L

.1

 

 

 

II.
n A A
3 .4 :2 3 '”
r4 .5 -
o H o :1
° . 0 z a
o D. 0
:3 a g u 1:!
u y d "‘
0 or-4 H
p. a +3 d «a
‘5 .. 2’: “ ‘6
3' d ad 3 6'
o u h
'3 .2 g :2 2
n. a: 3‘3 0 z?
.3 .5
.6
2.1
01 03
.1

.01

179

etus 931sue L

.-.—.— ._ ———————

Rhyacophila invaria L
Neotrichia L

 

Neotrichia P

 

 

909.919 N
Paleagap

$122013:
riparioides
GiCXLVIII
GiCL
GicLI
GiCLIIA
GtCXLV
GiCKLVI
HyCCL
EyCCLII
PoCCIX
BkCLXXXVIA 2.9
DiCCXXIV
DiCCXIX
DiCCXX
DiCCXXI
D1CCKXII
SaCCXVIlI

 

Sciaromium

 

LBCCXXIX
PoCCXI
PicXLI
P10XLII
PiCXLIII
PicXLIV

Hygrohypnum
111.1: 143.12

MyCCXXXII

{yCCXZXIII

 

varo 10129111,:
QQLéQQQQ
SaCCXVII

Fontinalis
ent 1.917;: 12123
var. 'iggntea
SaCCXV 2,4

 

Undetermined
SaCCXIII .5
SwCXCVII .8

a sp. 1 L

Hydroytilid gen. 1 L

Chimarra atgtriga L

Paleagapetus celsus P
Paragnetina sp. N

 

 

Micrasem

1'" nll I." 'III. I
i

DEC; BER COUNTS OF INSECTS PER GRAN DRY WEIGHT OF BRYOPHYTE

TABLE XVI.

v 8883016 Ejfiazbmdié

q 3335? Efffi535§fififi

u £135§35ffi3§$

u EfEETfiSftifii'Eifififiéfi

0—53535fiffi 51353551

. _-...._... .._‘

1 5533536 §EBE§56161q

 

q oqoxidoqemnéfifi

1 a duel? éfiififi

Eififiifi 9 556652
'Iant'fiihomou

 

q oqofiidOJpfifi
q 839591JBA xtzad?§
q snares anodefieoteg

n azqofifii

 

 

q Buttoxeo attqdooekqx

1 I 'd8 916mm

1 ondmoo
sod;q:;q unttnmxsoag

 

q captuouoxtqo

2.2
1.1

2.2
1.1

3.2
1.6

5.9
3.0

8
9

3
1

219.2
53-2
136.2

_———-

dalecarlica

MyEbIKXKIII

MyCCLXXXIV

ugdulatg

ToCCLKXXV

Av.
Scanania

Fontinalis

1.3

4.8?

7.6

NM
Mr“!

1.2

1.2

61.7 10.1

134.8 70.8
7506

ToCCLXXXVI
ToCCLXXXYII

1.0

1.5

85.5 40.0
88.3 20.2
89.1 28.2

ToCCLXXXVIII
Av.

ToCCLXXXIX

ST V.

MICHIGAN n E UNI LIBRARIES
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312 2883

9300994

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