PHYSICAL-CHEMICAL AND BIOLOGICAL CHARACTERIZATION OF SMALL 
STREAMS FOLLOWING INTENSIVE FOREST MANAGEMENT PRACTICES 
IN THE COASTAL PLAIN OF ALABAMA 
 
 
Except where reference is made to the work of others, the work described in this thesis is 
my own or was done in collaboration with my advisory committee. This thesis does not 
include proprietary or classified information. 
 
 
 
Sergio S. Ruiz-C?rdova 
 
 
 
Certificate of Approval: 
 
 
 
Elliot C. Webber, Co-Chair 
Research Fellow 
Fisheries and Allied Aqua-
cultures  
 
 
 
 
 David R. Bayne, Co-Chair 
Professor 
Fisheries and Allied Aqua-
cultures 
Jerry Michael 
Affiliate Associate Professor
USDA Forest Service 
 George Flowers 
Interim Dean  
Graduate School 
 
 
PHYSICAL-CHEMICAL AND BIOLOGICAL CHARACTERIZATION OF SMALL 
STREAMS FOLLOWING INTENSIVE FOREST MANAGEMENT PRACTICES 
IN THE COASTAL PLAIN OF ALABAMA 
 
Sergio S. Ruiz-C?rdova  
 
 
 
 
A thesis 
 
Submitted to 
 
the Graduate Faculty of 
 
Auburn University 
 
in Partial Fulfillment of the 
 
Requirements for the 
 
Degree of 
 
Master of Science 
 
 
 
 
 
 
 
 
Auburn, Alabama 
December 17, 2007
 
iii
PHYSICAL-CHEMICAL AND BIOLOGICAL CHARACTERIZATION OF SMALL 
STREAMS FOLLOWING INTENSIVE FOREST MANAGEMENT PRACTICES  
IN THE COASTAL PLAIN OF ALABAMA 
 
 
 
Sergio S. Ruiz-C?rdova 
 
 
Permission is granted to Auburn University to make copies of this thesis at its discretion, 
upon the request of individuals or institutions and at their expense.  The author reserves 
all publication rights. 
 
 
 
 
 Signature of Author 
 Date of Graduation 
 
 
  
 
 
iv
VITA 
Sergio Sigfredo Ruiz C?rdova, son of Laura C?rdova Corzantes and Jos? R?mulo 
Ruiz Araujo, was born May 21, 1959 in Santa Cruz El Chol, Baja Verapaz, Guatemala.  
He  attended Public Schools in Guatemala and graduated as an elementary school teacher 
from the Escuela Normal Rural No. 4 in 1976.  In January 1978 he entered the Center for 
Marine Studies and Aquaculture (CEMA), branch of the San Carlos University of      
Guatemala, and received the diploma of Aquaculture Technician in December 1980.  In 
January 1983 he enrolled in the School of Marine Sciences of the Autonomous Univer-
sity of Baja California Sur (UABCS) in Mexico and graduated with honors in June 1989 
as a Marine Biologist.  On January 1987 he married Mary Evelyn, daughter of Herbert E. 
and Carol S. Hawk and procreated two beautiful daughters Edith Alexandra and Judith 
Carolina.  Sergio has been working with Alabama Water Watch and Global Water Watch 
since joining Auburn University in the spring of 1993. 
 
v
THESIS ABSTRACT 
PHYSICAL-CHEMICAL AND BIOLOGICAL CHARACTERIZATION OF SMALL 
STREAMS FOLLOWING INTENSIVE FOREST MANAGEMENT PRACTICES 
IN THE COASTAL PLAIN OF ALABAMA 
Sergio S. Ruiz-C?rdova 
 
Master of Science, 17 December 2007 
(B.S., Autonomous University of Baja California Sur (Mexico), 1989) 
 
126 Typed Pages 
Directed by E. Cliff Webber and David R. Bayne 
 
 This study measured changes in five first-order streams (S 1 through S 5) follow-
ing management techniques including clearcut logging and chemical (S 2 and S 3) and 
mechanical site preparation (S 4 and S 5).  The herbicides used in this study included a 
mixture of Imazapyr and Glyphosate. Streams in two watersheds (S 3 and S 5) were left 
with 35-foot (11 m) wide streamside management zones (SMZ) while S 2 and S 4 had no 
SMZ.  The fifth stream (S 1) draining an undisturbed watershed of similar size was used 
as a control. During each phase of the study (predisturbance, harvest and site preparation) 
periphyton, macroinvertebrates and physicochemical data were collected at least two 
times per season from August 1993 to December 1995.  Statistical analyses utilized ran-
domized intervention analysis (RIA).   Pre-disturbance phase data showed similar sea-
sonal variation among all streams regarding water temperature, dissolved oxygen and al-
gal biomass, except for S 3 that was strongly influenced by springs. Water temperature
 
vi
in clearcuts with SMZ was not significantly different from the control while those     
without SMZ increased significantly following harvest.  Among biological communities, 
when compared to the control stream, algal biomass (as chlorophyll a) from periphyton 
showed the greatest change and significantly increased in all streams after harvest,      
particularly in those with no SMZ. These changes probably resulted from the reduced 
canopy cover and increased sunlight reaching the streams because nitrogen and phospho-
rous changed little following harvest or site preparation.  Following site preparation, 
chlorophyll a values remained high compared to the pre-disturbance phase. The herbicide 
had no apparent detrimental effect on periphyton biomass. Macroinvertebrate population 
densities were highly variable during the study period.  Following harvest, macroinverte-
brates in S 2 and S 4 with no SMZ had greater increases in density than that measured in 
the streams with SMZ.  However, only in S 4 was this difference significant. This density 
difference probably reflected the increase in algal biomass in S 2 and S 4.  After site 
preparation macroinvertebrate densities were unaffected when compared with the control 
using RIA. Taxa richness and diversity (i.e. both Shannon-Weaver and EPT) were not af-
fected during harvest or site preparation according to RIA.  Even with a 35-foot SMZ, 
timber harvest as practiced in the southeastern USA increased algal biomass as a result of 
additional light reaching the streams, especially streams with no SMZ. However, the 
presence of SMZ seemed to mitigate any dramatic changes to macroinvertebrate commu-
nities.  Chemical and mechanical site preparation techniques did not significantly affect 
periphyton biomass and macroinvertebrate communities during this study. Timber har-
vest without SMZ appeared to be the management practice that most affected the stream 
biota. 
 
vii
 
 
ACKNOWLEDGMENTS 
The author gratefully acknowledge the funding provided to conduct this research 
by the U. S. Forest Service, SRS, RWU-4105 Environmental Fate and Impacts Team and 
the National Council for Air and Stream Improvement (NCASI).  The Auburn University 
Lower Coastal Plain Substation in Camden, Alabama, the School of Forestry and the   
Department of Fisheries and Allied Aquacultures Aquatic Ecology and Rivers and Reser-
voirs Laboratories provided other funding and support.  
The author would like to express his gratitude to Dr. E. Cliff Webber, Dr. David 
Bayne and Dr. Jerry Michael for their assistance and heaps of patience during this re-
search. Many thanks are also due to family members close and away: Evelyn, Ale, and 
Caro, my brother Carlos and all others back home for their support during the course of 
this long journey.  Also thanks to all friends including Chris, Paul, Esau and family and 
Omar who helped me in many, many ways.  
   
 
viii
 
 
 
Style manual or journal used  
Standard Format Guide from the Auburn University Graduate School and 
The CSE Manual for Authors, Editors, and Publishers 
 
 
   
Computer software used  
  Microsoft Word 2000 for Windows  
 
 
 
ix
 
 
 
 
 
TABLE OF CONTENTS  
 
LIST OF TABLES???????????????????????????..xi 
LIST OF FIGURES??????????????????????????..xiii 
I.  INTRODUCTION.......................................................................................................... 1 
A.  Clearcut Harvest and Site Preparation....................................................................... 1 
B.  Streamside Management Zones ................................................................................. 6 
C.  Herbicides, Silviculture and Aquatic Biota ............................................................... 7 
II.  MATERIALS AND METHODS................................................................................ 11 
A.  Study Site................................................................................................................. 11 
B.  Experimental Design................................................................................................ 12 
C.  Field Measurements................................................................................................. 14 
D.  Laboratory Procedures............................................................................................. 16 
1. Substrate Composition and Benthic Organic Matter........................................ 16 
2. Periphyton ........................................................................................................ 17 
3. Macroinvertebrate Communities...................................................................... 18 
4. Nutrients and Herbicide Analyses.................................................................... 19 
5. Statistical Analysis ........................................................................................... 20 
III.  RESULTS AND DISCUSSION................................................................................ 21 
A.  Canopy Cover .......................................................................................................... 21 
B.  Water Temperature .................................................................................................. 23 
C.  Herbicide Fate.......................................................................................................... 30 
D.  Nitrogen and Phosphorous....................................................................................... 32 
E.  Streambed Substrate................................................................................................. 35 
F.  Benthic Organic Matter............................................................................................ 37 
G.  Periphyton Chlorophyll ........................................................................................... 40 
H.  Macroinvertebrate Communities ............................................................................. 49 
1.  Macroinvertebrate Density.................................................................................... 50 
2.  Effects of Clearcut and Site Preparation on Macroinvertebrate Density .............. 56 
 
x
3.  Taxa Richness, Species Diversity and Community Structure............................... 60 
I.  Summary ................................................................................................................... 68 
LITERATURE CITED ..................................................................................................... 71 
APPENDIX I .................................................................................................................... 83 
Table 1.  Composite list of taxa, tolerance value (TV) and functional feeding group 
(FFG) when known, presence at a sampling site (+) and total number of taxa per site 
collected in Camden, Alabama from March 1994 through December 1995................. 83 
APPENDIX II ................................................................................................................... 88 
Table 1.   Total fauna, number of taxa, and mean number of organisms/m?         
collected at each station in the Camden Study on 18 March 1994................................ 88 
Table 2.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 20 June 1994................................... 91 
Table 3.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 14 September 1994. ........................ 95 
Table 4.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 20 February 1995............................ 98 
Table 5.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 13 June 1995................................. 101 
Table 6.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 16 August 1995............................. 104 
Table 7.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 12 October 1995. .......................... 107 
Table 8.   Total fauna, number of taxa, and mean number of organisms/m?        
collected at each station in the Camden Study on 11 December 1995........................ 110 
 
xi
 
 
 
 
 
LIST OF TABLES 
Table----------------------------------------------------------------------------------------------- Page 
 
Table 1. Experimental design and watershed characteristics of the study area..................13 
Table 2.  Total precipitation and mean daily water discharge from five watersheds in 
Camden, Wilcox County, Alabama during 1995.................................................24 
Table 3.  Weekly mean total nitrate-nitrogen and total phosphate-phosphorous (mg/L) 
in study streams in Camden, AL between October 1994 and November 
1995......................................................................................................................32 
Table 4.  Mean benthic organic matter (g AFDM/m?) in the five streams in Camden, 
AL, during 1995 prior to disturbance, after timber harvest and after site 
preparation within the clearcut area.....................................................................39 
Table 5. Minimum, maximum and mean chlorophyll a (mg/m?) before disturbance 
(n=42) and after timber harvest (n=12), at nine stations in five streams in 
Camden, Alabama during 1993-1995.  a = within clearcut, b=below 
clearcut.................................................................................................................41 
Table 6. Mean chlorophyll a (mg/m?) before site preparation (11/93, 12/93,10/94, 
12/94, n=12)  and after site preparation (10/95, 12/95, n=6) and percentage 
increase from stage to stage for the reference stream and the sites within 
each clearcut watershed. ......................................................................................48 
Table 7.  Major insect orders and mean number of macroinvertebrates collected from 
core samples at sites in Camden, Alabama between August 1993 and 
December 1995. ...................................................................................................50 
Table 8.  Mean EPT index before harvest, after harvest and after site preparation at 
five streams in Camden Alabama from March 1994 to December 1995. 
Dash means no sampling collected due to drought..............................................62 
Table 9.  Mean Shannon-Weaver diversity index (H?) during pre-disturbance, post 
harvest and post site preparation in five streams near Camden Alabama from 
March 1994 through December 1995. .................................................................63 
Table 10.  Percentages of the total macroinvertebrate fauna by functional feeding 
group collected before and after harvest and after site preparation, inside the 
clearcut areas in five stream in Camden, Alabama during March 1994 to 
December 1995.  . ................................................................................................65 
 
xii
Table 11.  Percent composition of the most common macroinvertebrate taxa in the 
five dominant functional feeding groups (FFG) collected inside the clearcut 
areas of four streams in Camden, AL. before disturbance (PreDist), post 
harvest (PostHarv) and post site preparation (PostSite). .....................................66 
 
 
xiii
 
 
LIST OF FIGURES 
Figure------------------------------------------------------------------------------------------------ Page 
 
Figure 1.  Diagram of the five watersheds and streams of the Camden Study...................11 
Figure 2.  Mean percentage canopy cover from February 1994 through December 
1995 inside the clearcut areas of five streams in the Camden study. ..................21 
Figure 3.  Water temperatures for the five streams (S1...S5) in Wilcox County, 
Alabama.. .............................................................................................................24 
Figure 4.  Water temperatures and RIA analysis for the reference stream S 1 and S 2 
and S 4 with no SMZ.  . .......................................................................................26 
Figure 5.  Water temperatures and RIA analysis for the reference stream S 1 and S 3 
and S 5 with an SMZ.  . .......................................................................................27 
Figure 6.  Imazapyr concentrations (mg/L) measured in S 2 and S 3 after chemical 
site preparation in Camden, Alabama, 1995........................................................31 
Figure 7.  Mean substrate composition from five streams at Camden, Alabama during 
1995 prior to disturbance, after timber harvest and after site preparation...........36 
Figure 8.  Benthic organic matter composition (as g/m? of AFDM) in five streams in 
Camden, Alabama during 1995. ..........................................................................38 
Figure 9.  Chlorophyll a (mg/m?) concentrations in S 2 (No SMZ) inside (clear bar) 
and below (dark bar) the clearcut and in S 1 (line) in Camden, Alabama 
from August 1993 to December 1995..................................................................42 
Figure 10.  Chlorophyll a (mg/m?) concentrations in S 3 (SMZ) inside (clear bar) and 
below (dark bar) the clearcut and in S 1 (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................42 
Figure 11.  Chlorophyll a (mg/m?) concentrations in S 4 (No SMZ) inside (clear bar) 
and below (dark bar) the clearcut and in S 1 (line) in Camden, Alabama 
from August 1993 to December 1995..................................................................43 
Figure 12.  Chlorophyll a (mg/m?) concentrations in S 3 (SMZ) inside (clear bar) and 
below (dark bar) the clearcut and in S 1 (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................43 
 
xiv
Figure 13.  Mean chlorophyll a measured in a reference stream S 1 and RIA analysis 
of two experimental streams S 2 and S 4 with no SMZ in Camden, Alabama....44 
Figure 14.  Mean chlorophyll a measured in a reference stream S 1 and RIA analysis 
two experimental streams S 3 and S 5 with an SMZ in Camden, Alabama.  . ....45 
Figure 15.  Mean total macroinvertebrate density (No./ m?) in S 2 ? no SMZ and 
Chemically Site Prepared - inside (dark bar) and below (white bar) the 
clearcut and in the Reference Stream (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................51 
Figure 16.  Mean total macroinvertebrate density (No./ m?) in S 3 ?SMZ and 
Chemically Site Prepared - inside (dark bar) and below (white bar) the 
clearcut and in the Reference Stream (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................51 
Figure 17.  Mean total macroinvertebrate density (No./ m?) in S 4 ? no SMZ and 
Mechanically Site Prepared - inside (dark bar) and below (white bar) the 
clearcut and in the Reference Stream (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................52 
Figure 18.  Mean total macroinvertebrate density (No./ m?) in S 5 ?SMZ and 
Mechanically Site Prepared -  inside (dark bar) and below (white bar) the 
clearcut and in the Reference Stream (line) in Camden, Alabama from 
August 1993 to December 1995. .........................................................................52 
Figure 19.  Mean density of Chironomidae (dark bars) and Other macroinvertebrates 
(clear bars) per m? at S 1 and within the clearcut area at S 2 - S 5 in 
Camden, Alabama from August 1993 to December 1995...................................55 
Figure 20.  Mean density of total macroinvertebrates measured and RIA analysis in    
S 1 (reference stream) and within the clearcut of S 2 and S 4 with no SMZ.  ....57 
Figure 21.  Mean density of total macroinvertebrates measured and RIA analysis in    
S 1 (reference stream) and within the clearcut of S 3 and S 5 with an SMZ.......58 
Figure 22.  Mean taxa richness from selected dates for macroinvertebrates collected 
in S 1 (reference stream) and within the clearcut of the experimental 
streams. ................................................................................................................61 
 
 
 
1
 
 
 
 
 
 
I.  INTRODUCTION 
This study was designed to detect certain physicochemical and biological changes 
in Coastal Plain first-order streams flowing through small watersheds exposed to tradi-
tional southeastern forestry techniques.  The forestry practices utilized during harvest   
included use of stream management zones (SMZ) versus no SMZ; and following harvest, 
mechanical versus chemical site preparation prior to planting pine seedlings.  To assess 
differences between these practices, physicochemical variables were measured in each 
stream along with changes in periphyton and macroinvertebrate communities.  
A.  Clearcut Harvest and Site Preparation 
In the southeastern United States (US) intensive silviculture has contributed to 
forestry becoming one of the regions leading industries.   Loblolly pine (Pinus taeda L.) 
plantations are the predominant type of silviculture practiced in the Southeast (Rosson 
1995).  Several approaches are used in intensive forestry but two, in particular, are of 
concern as potential sources of contaminants to streams.  These two practices are clearcut 
harvest and site preparation prior to planting pine seedlings.  Clearcutting creates areas of 
bare soil that, when exposed to rain, may result in erosion and sedimentation to local   
water bodies.  Site preparation can also result in erosion and depending on the method 
used, may contribute runoff of pesticides to streams and lakes according to a 1993 study 
by the United States Environmental Protection Agency (US EPA). 
 
2
Clearcutting generally involves removal of all trees larger than four inches in     
diameter (Bryce et al. 1989).  Following harvest, the main purpose of site preparation is 
to reduce competition for light, moisture and nutrients between newly planted pine seed-
lings and unwanted vegetation.  In addition, site preparation facilitates planting and en-
hances growth and survival of new trees. 
Site preparation methods are categorized as follows: mechanical, chemical, and 
burning (Roth 1987).  Mechanical preparation involves the use of heavy machinery, such 
as bulldozers with special attachments that directly remove competing vegetation, clears 
debris and incorporates organic matter into the soil.  Consequently, large areas of bare 
soil are often exposed to rain and wind action.  Chemical site preparation utilizes herbi-
cides to reduce undesirable vegetation.  Use of herbicides has become increasingly popu-
lar in recent years especially on steep slopes, because there is little soil disturbance. 
Chemical preparation is also less expensive than mechanical techniques (Roth 1987).  
The most commonly used herbicides in forestry in the Southeastern US are Accord?,   
active ingredient (ai) glyphosate; Arsenal AC? and Chopper?, (ai) imazapyr; Oust?, 
(ai) sulfometuron; Velpar L?, (ai) hexazinone; and Garlon 4?, (ai) triclopyr (Personal 
communication Dr. Bruce Zutter, School of Forestry, Auburn University, 1998).  These 
herbicides are applied alone or as a mixture of two or more chemicals, depending on the 
plant species.  Burning as a method of site preparation may be used exclusively, or as a 
supplement to either chemical or mechanical techniques (Roth 1987).  Aerial applications 
of incendiary mixtures are the most common technique used in burning. 
Harmful effects of these herbicides may persist in the watershed.  Half-life of  
glyphosate was reported in a range from 2 to 174 days in soils and from 3.5 to 70 days in 
 
3
water (SERA 2003).  Michael and Neary (1991) reported half-life for imazapyr in      
Alabama soils ranging from 19 to 34 days.  Fowlkes et al. (2003) reported a half-life of    
3.2 to 3.4 days for imazapyr in microcosms. 
In numerous biological studies of stream periphyton communities increases in  
primary production occurred after logging (Hansmann and Phinney 1973, Gregory 1980, 
Murphy and Hall 1981, Lowe et al.1986, Murphy et al.1986).  In streams without SMZ, 
Murphy and Hall (1981) found significantly higher periphyton densities from watersheds 
harvested 5 to 17 years earlier compared to undisturbed sites.  Borg et al. (1988) also 
found a significant increase in algal blooms in streams from logged sites without SMZ,  
compared to streams with 100-m wide SMZ.  However, Shortreed and Stockner (1983) 
concluded that physicochemical factors that were modified as a result of logging had    
little effect on periphyton communities because phosphorous levels were not generally 
affected.  Kosinski and Merkle (1984) reported that glyphosate significantly inhibited 
alga photosynthesis.  In contrast, Austin et al. (1991) suggested that glyphosate served as 
a nutritional source of phosphorous and stimulated increased periphyton growth in       
experimental streams.  Imazapyr has been found to be more toxic to rooted and floating 
macrophytes than to other aquatic organisms, including algae (Roshon et al. 1999).   
Logging operations during harvest also include disturbances such as road con-
struction, log decks, skid trails and other activities that contribute to erosion and sedimen-
tation in local streams (Bryce et al. 1989).  Protecting stream quality from activities  
associated with intensive silviculture has gained increased attention by state and federal 
agencies.  For example, the Alabama Department of Environmental Management 
(ADEM) and the US EPA require that forest operations be conducted in such a way that 
 
4
stream quality is not impaired (US EPA 1993; ADEM 1992).  State agencies recommend 
the use of best management practices (BMPs) to maintain and protect the physical, 
chemical and biological integrity of waters (AFC 1993). 
Clearcut harvest and site preparation have been shown to disrupt forest ecosys-
tems (Noel et al. 1986; Campbell and Doeg 1989; Davies and Nelson 1994).  Streams 
flowing through these disturbed ecosystems typically undergo five types of modifica-
tions.  These include changes in discharge, water temperature, turbidity and sedimenta-
tion, dissolved nutrients and allochthonous organic detritus (Lynch et al. 1977). 
Stream discharge often increases following canopy removal because water inter-
ception and transpiration on the watershed are severely reduced when trees are subtracted 
(Hornbeck et al. 1970, Aubertin and Patric 1974, Miller 1984).  Significantly higher    
water yields have been reported from clearcut watersheds without SMZ (Beasley and 
Granillo 1988).  Brozka et al. (1982) reported a 95% increase in flow volume in streams 
the first year after clearcutting but such effects have persisted even two years after      
harvest (Miller 1984).  Abdul-Rahim and Harding (1992) concluded that SMZ help ame-
liorate hydrological impacts of logging on streamflow.  They found that peak streamflow 
in clearcut areas without an SMZ increased 58% compared to a clearcut with an SMZ. 
Water temperature in streams usually increases after a clearcut because of addi-
tional sunlight reaching the water (Burton and Likens 1973; Rishel et al.1982).  After a 
timber harvest in West Virginia, mean summer temperatures increased 4?C with maxi-
mum temperatures increases greater than 9?C; while mean winter temperatures decreased 
by 2?C (Lee and Samuel 1976).  SMZ mitigate these temperature fluctuations by provid-
ing cover to the stream (Rishel et al. 1982; Belt and O?Laughlin 1994).  Davies and  
 
5
Nelson (1994) found that stream temperatures increased 10% where SMZ widths were 
less than 10-m, while in streams with SMZ of 10-30 and 30-50-m, water temperatures 
were not significantly different from undisturbed streams. 
Erosion and sedimentation is another common problem associated with clearcut-
ting.  Sediment yields in streams were found to increase after harvest and mechanical site 
preparation (Tebo 1955; Cordone and Kelly 1961; Brown and Krygier 1971; Bormann   
et al. 1974; Patric et al. 1984; McClurkin et al. 1985, Platts et al. 1989).  Harvesting tech-
niques also influence sediment losses from clearcut sites (McClurkin et al. 1985).  A 
study conducted in the Coastal Plain of Arkansas compared sediment movement in 
streams from chemically and mechanically site prepared watersheds versus undisturbed 
ones.  Mean annual sediment losses on mechanically site prepared watersheds (264 
kg/ha) during the first post-treatment year were significantly higher than the mean of       
4 kg/ha from undisturbed controls (Beasley et al. 1986; Beasley and Granillo 1988).    
Michael et al. (2000) suggested from research conducted in the upper Coastal Plain of 
Alabama that using herbicides properly during site preparation in combination with SMZ 
have the potential to reduce sediments reaching the streams. 
Studies on nutrient levels in streams following clearcut have revealed differing  
results.  Some studies have reported increases in nutrients to be negligible, or of short  
duration, following clearcuts (Aubertin and Patric 1974, Patric 1980, Martin et al. 1984); 
however, Brozka et al. (1982) found a 274% increase in nitrate (NO
3
-N) concentration in 
streams following clearcut.  Smith et al. (1988) found that chemical site preparation also 
resulted in significant increased concentrations of nitrate in streams in Maine.  Manual 
applications of hexazinone pellets to a Piedmont watershed caused NO
3
-N to exceed 
 
6
normal levels by two orders of magnitude 2-yr after treatment (Neary et al. 1986).  Me-
chanical site preparation caused a 5-yr increase in nitrate concentration in groundwater in 
Finland (Kubin 1995).  Blackburn and Wood (1990) found significant differences in     
nitrogen and phosphorous in streams with SMZ 6-20 m wide and mechanically site     
prepared, compared to undisturbed streams.  Patric (1980) suggested that a 20-m wide 
SMZ minimized water quality effects, while in treatments without an SMZ nitrate levels 
increased five-fold during the first year after disturbance.  Lowe et al. (1986) suggested 
that light was the factor limiting algal accumulation in forested watersheds after experi-
mental additions of nutrients to streams. 
Clearcuts also alter the quantity, quality and timing of allochthonous organic mat-
ter introduced to streams. Gurtz and Wallace (1984) found significantly higher amounts 
of woody material in streams draining logged watersheds compared to undisturbed ones. 
B.  Streamside Management Zones 
One of the primary forestry BMPs suggested by states to reduce runoff of soil and 
chemicals to streams and lakes include leaving riparian forests or streamside management 
zones (SMZ) along these water bodies to protect aquatic ecosystems and water quality 
(Michael 2004).  SMZ are also referred as ?buffer strips? (Belt et al. 1992).  In Alabama 
the suggested minimum standard width of an SMZ is 11-m (35 feet) from a definable 
bank (Alabama Forestry Commission 1993) and 15-m (50 feet) minimum if wildlife pro-
tection is a major objective. Also, SMZ width should increase 6-m (20 feet) for each 10% 
increase in slope (Brinker 1989).  
Riparian forests are vegetative zones along streams that serve as complex ecosys-
tems.  These zones provide allochthonous organic materials (leaves and woody debris) 
 
7
that help maintain the biological productivity and diversity of streams (Gurtz et al. 1980; 
Murphy et al. 1981; Webster and Waide 1982; Webster et al. 1983; O'Hop et al. 1984; 
Andrus et al. 1988; Carlson et al. 1990).  Logs, branches and leaves also create habitat 
for stream organisms (Sweeney 1993).  SMZ are useful in mitigating or controlling non-
point source pollution by capturing sediment from surface runoff (Brown and Krygier 
1971, Brown 1971; Newbold et al. 1980; Hawkins et al. 1982; Gurtz and Wallace 1984; 
Welsch 1991; Comerford et al. 1992).  SMZ are also effective in removing nutrients (Hill 
1996; Johnson et al. 1996), reducing the amount of herbicide reaching streams (Michael 
2004) and providing shade to cool the waterbody, thus helping maintain light and tem-
perature conditions for stream biota.  
C.  Herbicides, Silviculture and Aquatic Biota 
Herbicides used in site preparation have been shown to contaminate ground and 
surface water and can be toxic to aquatic invertebrates and fish (Swadener 1993; Cox 
1996, 1998).  However, most toxicity studies have been conducted in the laboratory un-
der controlled climatic conditions (Kosinski and Merkle 1984, Austin et al. 1991) or in 
experimental stream channels through which flow is carefully controlled (Kreutzweiser  
et al. 1992, Schneider et al. 1995).  The organisms used in these studies were exposed to 
constant levels of the herbicides during established time periods.  Field studies do not 
provide this type of exposure for organisms.   Limited field studies were found that tested 
the toxicity to aquatic biota of the herbicides typically used in intensive silviculture prac-
tices (Mayack et al. 1982, Michael et al. 1999, Fowlkes et al. 2003).  Michael et al. 
(1999) concluded that benthic macroinvertebrates in Piedmont streams of the  
 
8
Southeastern United States appear insensitive to hexazinone at the exposures observed in 
their study.  
Accord? (a. i. imazapyr) and Arsenal AC? (a. i. glyphosate) were the herbicides 
used for chemical site preparation in this study. Fowlkes et al. (2003) found that a      
concentration of imazapyr 100 times the expected from typical application rates did not 
affect in situ microcosm macroinvertebrate communities.  In addition to herbicide toxic-
ity, concerns exist that 50% or more of the herbicides used in site preparation such as  
glyphosate include inert ingredients that may be more toxic than the active ingredient 
(SERA 2003).  Formulations of Accord? and Arsenal AC? contain more than 50% inert 
ingredients (USDA Forest Service 1995a, 1995b).   
Accord? and Arsenal AC? in concentrations typically used in forestry are con-
sidered practically non-toxic to freshwater fish and aquatic invertebrates (USDA Forest 
Service 1995a, 1995b); even though concentrations of glyphosate between 3 and 25 ppm 
were lethal for the crustacean Daphnia pulex  (Folmar et al. 1979, Servizi et al. 1987, 
Hessen et al. 1994).  In addition, Hartman and Martin (1984) found that glyphosate con-
centrations of more than 1-mg/L significantly reduced D. pulex populations, mainly due 
to mortality of immature stages.  Another aquatic invertebrate, the red swamp crawfish 
Procambarus clarkii, showed a 96-h LD50 when exposed to 47.3 mg/L of glyphosate 
(Holck and Meek 1987). 
Forestry herbicides also affect aquatic vegetation.  Glyphosate concentrations 
greater than 9 mg/L resulted in 50% photosynthetic inhibition of periphytic algae in small 
ponds (Goldsborough and Brown 1988).  Peterson et al. (1994) in a study of periphyton 
communities, reported inhibition of carbon uptake in diatoms in the presence of  
 
9
glyphosate.  The green alga, Selenastrum capricornutum, exhibited growth inhibition   
after 96-h exposure at concentrations of 5,300-5,500 mg/L of imazapyr as well as with 
2,600 mg/L of glyphosate (Thomas et al. 1990).   
Considering the natural undisturbed state present in the Camden watersheds at the 
beginning of this study, macroinvertebrate communities were expected to resemble that 
described by the River Continuum Concept (RCC) for small-undisturbed headwater 
streams. The RCC describes a continuous gradient in the distribution of organic matter 
and macroinvertebrate functional groups from headwater to mouth (Vannote et al. 1980). 
 According to the RCC, small headwater streams are narrow and generally well shaded 
by the riparian canopy.  Thus insufficient light may reach the streambed to promote algal 
growth and nutrients may be low.  Since the stream does not produce enough energy to 
supply the demands of organisms, allochthonous energy becomes critical and enters the 
stream in the form of coarse particulate organic matter (CPOM) from the terrestrial envi-
ronment.  In headwater streams, leaf-shredding macroinvertebrates often constitute a 
large portion of the macroinvertebrate population, thus shredders and collectors are     
usually abundant. Grazers, macroinvertebrates that scrape algae from rock surfaces, are 
often low in abundance in headwater streams because insufficient sunlight reaches the 
channel thus resulting in a scarcity of algae.  Predators should be common throughout the 
river system and can be found in headwaters, in smaller proportion relative to organisms 
of other feeding types. 
Increased numbers of aquatic insects have been found following clearcuts (New-
bold et al.1980, Murphy et al.1981, Carlson et al.1990) even 5 to 10 years after harvest, 
as well as changes in macroinvertebrate diversity and biomass (Murphy et al.1981,  
 
10
Duncan and Brusven 1985).  Apparently, changes in the insect populations were a result 
of shifts in the food base after the disturbance.  Davies and Nelson (1994) found an 80% 
decrease in macroinvertebrate abundance in logged sites with SMZ smaller than 30 m, 
and Newbold et al. (1980) found higher densities of tolerant macroinvertebrate taxa and 
lower diversities in reaches without SMZ.  SMZ provided protection from intensive     
silvicultural practices because macroinvertebrate communities in California streams with 
SMZ wider than 30-m could not be distinguished from those of undisturbed reference 
streams in terms of diversity, similarity or density (Newbold et al. 1980).  In addition, 
Boschung and O?Neil (1981) concluded that the aquatic macroinvertebrate community 
structure in an east Alabama stream was not affected following clearcuts as prescribed by 
the US Forest Services standards including SMZ. 
The purpose of my study was to quantify changes in stream quality associated 
with intensive forestry practices used in pine regeneration.  Water quality, periphytic    
algae and benthic macroinvertebrate communities were examined in five first-order 
streams prior to disturbance. Changes in these variables were measured during clearcut 
harvest and site preparation to determine any physicochemical and biological impacts of 
the disturbance.  Changes were examined in streams with an SMZ and in streams without 
an SMZ.  In addition, chemical versus mechanical site preparation techniques were com-
pared for differences in impacts to the biota. 
 
11
 
 
 
 
 
II.  MATERIALS AND METHODS 
A.  Study Site 
This study was conducted between 1993 and 1995 at the Alabama Agricultural 
Experiment Station, Lower Coastal Plain Substation, located in Wilcox County about    
2.5 km north of the city of Camden, Alabama.  Five small, topographically well-defined 
watersheds (W1?W5) were identified, each drained by a perennial first-order stream     
(Figure 1).  
 
Figure 1.  Diagram of the five watersheds and streams of the Camden Study, in Wilcox County, 
Alabama.  Solid lines are the streams, dotted lines are the watershed boundaries, circles are the 
approximate location of sampling sites and each H represents a flume placed at the lower bound-
ary of each clearcut.  Sampling sites were located upstream and downstream from flumes in ex-
perimental streams. 
 
12
Smithdale and Bama soils, derived from marine and fluvial sediments eroded 
from the Appalachian and Piedmont plateaus, are characteristic of these lower Coastal 
Plain areas.  These soils have loamy subsoil and a sandy loam surface layer with slopes 
less than 5% (Brannon 1998).  Sweetgums (Liquidambar styraciflua), eastern hop horn-
beam (Ostrya virginiana), elms (Ulmus spp.), hickories (Carya spp.) and maples (Acer 
spp.) dominated the tree populations on the watersheds. Loblolly pines (Pinus taeda) 
dominated the basal areas with scattered oaks (Quercus spp), magnolias (Magnolia spp.) 
and poplars (Liriodendron tulipifera) among others (Marshall 1999).  The streams were 
unnamed tributaries of the Alabama River, and William Dannelly Reservoir, and will be 
referred as S1, S2, S3, S4 and S5.  All streams were about 1-m wide with small reaches 
approaching 2 to 3-m in width.  Water depth ranged from less than 5-cm where the 
streambed was wide, to about 30-cm in pool areas where the channel was narrow.  Each 
stream was typical of small-undisturbed Coastal Plain streams as described by Smock 
(1988).  Trees grew to the banks of each stream and the forest canopy was dense.  Al-
lochthonous materials of leaves and other vegetative debris were the main energy inputs 
to the streams as expected based on the RCC.  Stream habitats consisted mainly of shal-
low pools, runs of loose shifting sand and riffles of mostly gravel, sand and some cobble. 
 Steep-sided mud banks were typical along most of the streams. 
B.  Experimental Design 
Studies prior to disturbance were initiated in August 1993.  By December 1994 
each stream had been gaged with an H-flume placed near the downstream edge of the  
clearcut in each watershed (Figure 1).  In S1 the flume was placed just upstream from     
confluence with S2.  Flumes were equipped with air and water temperature probes, Keller 
 
13
PSI pressure transducers, Campbell CR10 data loggers and ISCO 3700 automatic water 
samplers.  Stream velocity, total discharge, water temperature and nutrients data were  
obtained from these automatic samplers.  Rain gauges were placed in the middle of each 
watershed to record precipitation.   Scientists from the USDA Forest Service installed the 
flumes, collected the physicochemical data and conducted the herbicide analysis. 
Intensive forestry practices were applied to four of the watersheds (W2, W3, W4 
and W5).  The fifth watershed (W1) was used as a reference and left undisturbed.  Two 
different experimental treatments were applied to the watersheds receiving clearcuts.  
Two streams were left with a 35-foot (11-m) wide SMZ while two streams had no SMZ.  
In addition, as preparation before planting pine seedlings, two watersheds received 
chemical site treatment and two watersheds were mechanically site prepared (Table 1).   
A single sampling station (1c) was located in the reference stream upstream of the 
flume.  Two stations were established in each of the other four streams; one inside the 
clearcut area above the flume (2a, 3a, 4a and 5a), and one downstream from the flume 
(2b, 3b, 4b and 5b).  Each station included a stream reach of about 100-m.  Streams were 
sampled twice per season from August 1993 to December 1995. 
 
Table 1. Experimental design and watershed characteristics of the study area. 
Watershed Total Area (ha) Clearcut (ha) SMZ Site Preparation 
1 16.6 N/A N/A N/A 
2 20.2 14.4 No Chemical 
3 18.6 14.4 Yes Chemical
4 50.2 14.8 No Mechanical 
5 26.2 13.4 Yes Mechanical 
 
14
During February and March 1995 a commercial logger harvested timber from 
W2, W3, W4 and W5 using a circular saw head-feller buncher, rubber-tired skidders and 
a deck loader.   Between 17 and 21 August 1995, W4 and W5 were mechanically site 
prepared using crawler tractors with a root rake.  Small trees and shrubs less than 4-inch 
in diameter were sheared and pushed into windrows.  On 30 August 1995, W2 and W3 
were chemically treated with a mixture of Arsenal AC?, a.i. imazapyr [2-[4,5-dihydro-4-
methyl-4-(1-methylethyl)-5-oxo-1H-imidazol-2-yl]-3-pyridinecarboxylic acid] and Ac-
cord?, a.i. glyphosate [N-(phosphonomethyl)glycine], in the form of its isopropylamine 
salt.  Labeled rates of Arsenal AC?, Accord? and a surfactant were mixed to provide an 
equivalent of 1.12-kg/ha of imazapyr, 3.36-kg/ha of glyphosate and 0.28-kg/ha of non-
ionic surfactant.  The herbicide mixture was applied by helicopter using a microfoil or 
TVB spray system to help ensure accurate application and minimum drift.  On 21 No-
vember 1995, a prescribed burn was conducted on the four harvested watersheds using an 
incendiary mix dropped from a helicopter with an aerial drip torch, and on the ground us-
ing hand held drip torches.  Each clearcut was planted with loblolly pine in January 1996. 
C.  Field Measurements 
In addition to physicochemical measurements collected from water at each flume, 
from August 1993 to December 1995 the following data were collected.  Water tempera-
ture and dissolved oxygen (DO) were measured in situ at each station using a model 51B 
Yellow Spring Ohio (YSI) Dissolved Oxygen meter.  Canopy cover was measured as a 
percentage using a spherical densiometer (Lemmon 1957).  Streams were sampled for  
periphyton and macroinvertebrates at least one week after heavy rains to allow those  
 
15
communities to recover from any scouring effects that might have resulted from the      
increased discharge.  Quantitative samples of periphyton and benthic macroinvertebrates 
were collected using PVC core samplers developed in the Auburn University Laboratory 
by Dr Cliff Webber and me.  Cores have one end beveled to allow easy insertion into the 
stream bottom and their small size makes them appropriate to obtain a quantitative       
periphyton sample easier than using a dredge or Ponar sampler.  Three periphyton sam-
ples were collected from randomly selected riffles at each station by inserting a 50-mm 
inner diameter core sampler (area = 0.002 m?) into the substrate to a depth of about 2-cm. 
 While firmly holding the corer in place, an acrylic plate was inserted under the PVC 
sampler to prevent loss of the sample.  The sample was lifted from the streambed and 
placed inside a labeled plastic Ziplock? bag.  Each sample was stored in the dark on ice 
for transport to the laboratory.  
Three macroinvertebrate samples from riffles and three from pool areas were col-
lected at each station by inserting a 77-mm inner diameter core sampler (area = 0.005 m?) 
into the substrate to a depth of about 10-cm.  An acrylic plate was inserted under the PVC 
sampler to prevent loss of the sample as it was lifted from the streambed and placed into 
a galvanized bucket.  Riffle and pool habitats were randomly selected from the stream 
reach at each station.  During the predisturbance phase of the study macroinvertebrates 
were separated in the field from sediment and debris by elutriation.  Six times water was 
added to the bucket, vigorously swirled and the water and organisms poured through a 
U.S. Standard No. 60 sieve (250-?m mesh).  Samples were stored in ice chests for trans-
port to the laboratory where they were refrigerated overnight at 4?C.  Estimates of the 
substrate composition at each station were added to the sampling protocol during the   
 
16
experimental phase of the study.  This included one date prior to clearcutting and all     
remaining dates.   
In 1995, two of the three macroinvertebrate cores from riffles and two from pools 
were placed directly in plastic bags and returned to the laboratory for elutriation and 
analysis of substrate composition.  Macroinvertebrates were elutriated in the field from 
the third sample from each habitat type.  Sampling in 1995 included two dates before 
harvest, four dates between harvest and site preparation, and two dates after site prepara-
tion. 
D.  Laboratory Procedures 
1. Substrate Composition and Benthic Organic Matter 
After macroinvertebrates were elutriated from the samples, visual estimates were 
made of the percent composition of particles from the remaining substrate.   The substrate 
material was evenly spread over the bottom of the bucket and estimates were made of the 
proportions of silt (< 0.06 mm), sand (0.06-2.0 mm), gravel (2.0-16 mm), pebble (16-64 
mm) and cobble (> 65 mm).  In addition, benthic organic matter (BOM) was measured 
from each sample. Three categories of BOM were measured using methods described by 
Golladay et al. (1989): woody debris (WD), particles 1-cm? and larger; large benthic or-
ganic matter (LBOM), particles retained by a No.18 sieve (1-mm mesh); and fine  benthic 
organic matter (FBOM), the material retained by a No.60 sieve (250-?m mesh).  Samples 
were poured through a No.18 sieve stacked onto a No.60 sieve.  Leaves, seeds and twigs 
(WD) were manually separated and placed in a plastic container with the station identifi-
cation.  The materials retained by the No.18 and No.60 sieves were separated into indi-
vidual containers labeled LBOM and FBOM, respectively. 
 
17
Samples of BOM were dried to a constant weight in an oven at 103?C - 105?C for 
1.0 hour and dry weight was determined using an analytical balance.  Samples were then 
incinerated in a muffle furnace at 500?C for 60 minutes.  Ash was rewet with distilled 
water after it cooled in a desiccator, and then brought to a constant weight at 103 to 105?C 
to estimate the ash free dry mass (AFDM).  Dry weight and AFDM were calculated as a 
percentage of the sample for WD, LBOM and FBOM following Standard Methods (APHA 
1995).  Mean BOM was estimated for each station. 
2. Periphyton 
Pigment extraction and sample handling was conducted in subdued light to avoid 
degradation of chlorophyll.  In a refrigerator samples were allowed to settle overnight to 
concentrate algae by sedimentation. The following morning excess water was siphoned 
from each bag and 30 ml of 90% alkaline acetone was added.  Samples were refrigerated 
for 24-h at 4?C in the dark and vigorously shaken every two hours to disrupt algal cells 
and facilitate pigment extraction.  After 24-h the chlorophyll extract was poured into 
tubes and centrifuged for 20 minutes at 500 g and 4?C to remove sediment and debris.  
Optical density (OD) of each sample was measured at 630, 647 and 750 nanometers 
using a Beckman DU-50 Series Spectrophotometer.  The chlorophyll content of 
periphyton was determined to estimate algae biomass following the trichromatic method 
(APHA 1995).  Chlorophyll a concentration was determined for each replicate using the 
formula: 
Ca = 11.85*(OD664) - 1.54*(OD647) - 0.08*(OD630) 
 
18
where 11.85, 1.54, and 0.08 are correction factors, and OD630, OD647, and OD664 are 
the corrected optical densities (with a 4-cm light path) at the respective wavelength.  The 
amount of pigment per unit area was calculated using the formula: 
mg chlorophyll a/m? = (Ca * 0.03)/0.002 
where Ca is the chlorophyll a concentration in the extract; 0.03 is the volume of the 
extract in liters, and 0.002 is the surface area of the core sampler in m?. 
3. Macroinvertebrate Communities 
Within 24 hours of collection, macroinvertebrate samples were preserved in 5 to 
10% formalin for several days.  Samples were then washed through a No. 60 sieve and 
rinsed with water to remove the formalin.  Remaining debris and organisms were stored 
in 70% ethyl alcohol.  Macroinvertebrates from all samples were sorted and counted 
using a stereomicroscope.  Macroinvertebrates were identified to the lowest practical 
level (usually genus) using standard taxonomic keys (Merritt and Cummins 1996; 
Wiggins 1996; Edmunds et al. 1977; Bednarik and McCafferty 1979; Stewart and Stark 
1988; and Wiederholm 1983).  Macroinvertebrates were sorted into two categories, the 
?Chironomidae?, midges in the Order Diptera, and ?Others?, the other categories 
included all other insects besides the midges and the miscellaneous invertebrates from 
each sample.   
When known, macroinvertebrates were also assigned to a functional feeding 
group (FFG) based on classification by Merritt and Cummins (1996).  The following 
categories were included: filtering collectors (FC), gathering collectors (GC), scrapers 
(SC), shredders (SH), predators (P) and piercers (PI).  Densities for each core sample 
were expanded to number of organisms per m?.  The total number of taxa (taxa richness) 
 
19
was calculated for each sample as well as the EPT index, based on the number of taxa 
within the orders Ephemeroptera, Plecoptera and Trichoptera (Plafkin et al. 1989).  
Streams with a high EPT richness are considered in better condition than those with 
lower EPT richness since aquatic stages of these organisms are more sensitive to 
pollution. The diversity of macroinvertebrate communities was calculated using the 
Shannon-Weaver index (H?) (May 1975).  The formula used was: 
H = ?
=
?
ppii
T
i
S
ln
1
 
where p
i 
is the proportion of individuals in the ith species, p
i
 = N
i
/N
T
.   
Densities for each core sample were compiled for ?Chironomidae? and ?others? 
for all seventeen sampling dates between August 1993 and December 1995 and expanded 
to number of organisms per m?.  Macroinvertebrates from pool and riffle habitats were 
combined for analysis of total densities.  Analysis of taxa richness, community structure 
and diversity was conducted for selected dates that included four dates before harvest, 
two dates between harvest and site preparation and two dates after site preparation.   
4. Nutrients and Herbicide Analyses 
Nitrogen, phosphorous and herbicide concentrations from each watershed were 
analyzed by the USDA-FS Southern Forest Experiment Station at Auburn University 
using the water samples collected in the flumes from December 1994 to December 1995. 
 Ten-ml aliquots were processed using a Dionex? ion chromatograph to quantitatively 
determine concentrations of nitrates (NO
3
) and phosphates (PO
4
).  Off-site movement of 
imazapyr from the clearcut to streamflow was determined from water samples by high 
performance liquid chromatography (HPLC).  Most methods for analyzing glyphosate, 
 
20
like the EPA method 547, are very expensive, time consuming and require specialized 
equipment. Therefore, no tests were conducted to analyze this herbicide. 
5. Statistical Analysis 
Calculations for each replicate were averaged to conduct the statistical analysis.  
These variables were used to evaluate macroinvertebrate differences among the sites, 
before and after harvest, and before and after site preparation.  Values for the replicates 
from each station were averaged and analyzed statistically.  The nature and scale of this 
study qualified it as a whole-ecosystem experiment and replicating whole ecosystems is 
seldom possible.  A statistical method designed to address such experiments is 
Randomized Intervention Analysis (RIA).  This method detects changes in a 
?manipulated ecosystem? relative to an undisturbed reference ecosystem (Carpenter et al. 
1989).  RIA indicates whether a change has occurred or not, however it does not 
demonstrate that the disturbance was the cause of the change.  Parallel observations from 
the reference and manipulated streams were paired in time spanning periods before and 
after disturbance. A computer program designed for RIA (Carpenter et al. 1989) analyzed 
all measurement means.  When data did not fit the RIA program, a Tukey's test was used 
to statistically compare differences among means with a probability ? = 0.05 for all tests 
of significance.  Chlorophyll a and macroinvertebrate data values were logarithmically 
transformed [log
10
 (X
i
 + 1)] in order to stabilize variance (Zar 1984).  The biological and 
physicochemical measurements collected between August 1993 and February 1995 
provided the pre-disturbance database that was compared against the same variables 
measured during post-harvest (March to August 1995) and post-site preparation 
(September to December 1995).  
 
21
 III.  RESULTS AND DISCUSSION 
A.  Canopy Cover 
Watersheds in this study had not been harvested for many years so at the begin-
ning of this study all streams had a dense canopy consisting of a diverse riparian flora.  
Canopy cover prior to harvest was close to 100% in all streams, decreasing in the fall and 
winter to about 60% as hardwoods shed their foliage (Figure 2). 
 
<---Site 
Preparation
<--Harvest
0
25
50
75
100
Jan-94 Mar-94
May-94
Jul-94
Sep-94 Nov-94
Jan-95 Mar-95
May-95
Jul-95
Sep-95 Nov-95
Canopy Cover (%)
S 1 S 2 S 3
S 4 S 5
Figure 2.  Mean percentage canopy cover from February 1994 through December 1995 in the  
reference stream and inside the clearcut area of four experimental streams in the Camden study. 
 
22
After harvest the percent canopy cover in S 4 with no SMZ was reduced to about 
10% within the clearcut areas, significantly less than that in the two watersheds with an 
SMZ (S 3 and S 5) and the reference stream (S 1).  In S 2 (No SMZ) the decrease in can-
opy cover was gradual because steep slopes near the stream banks prevented mechanical 
harvest of many trees.  Thus, no significant differences in canopy cover were found      
between S 2 and S 1 in the weeks following harvest. Several weeks later trees greater 
than four inches in diameter were manually harvested from the S 2 watershed allowing a 
complete clearcut similar to that on W 4.  Stream 4 had no steep banks so trees greater 
than four inches in diameter were harvested to the edge of the stream.  After harvest, can-
opy cover in S 3 and S 5 with SMZ ranged between 3 and 15% less than that measured in 
S 1, but significant differences were not detected among the three watersheds. These data 
suggested that an ~11-m SMZ was sufficient to mitigate increases in solar radiation 
reaching low order streams flowing through a clearcut.  During the following spring and 
summer after harvest, vegetation within the clearcuts began growing in each watershed.  
Shrubs, grasses and vines were present by mid-summer on the watershed of each stream.  
Chemical site preparation within the harvested area on W 2 and W 3 killed all 
forms of vegetation in August 1995.  Riparian vegetation along the stream banks of S 2 
was eliminated creating significant differences in canopy cover between S 2 and S 1 that 
persisted through the end of the study.  Riparian vegetation within the SMZ along S 3  
remained undisturbed following the herbicide application.  Mechanical site preparation 
within the clearcuts on W 4 and W 5 did not significantly affect percent canopy cover.  
New grown riparian vegetation along S 4 was not disrupted by the treatment, and S 5 was 
 
23
protected by the SMZ.  However it will probably take years for the canopy cover along    
S 4 to return to conditions similar to those present prior to harvest. 
After harvest, from June to August 1995, water tables in the watersheds dropped 
following a long period of high air temperatures (mean =34?C) and little precipitation 
(e.g. 30-mm in June).  Because of the small size of these watersheds, no surface flow was 
evident in S 1 during this time period. Similar conditions occurred in S1 during Septem-
ber 1994 but for a shorter period of time.  Climatic conditions plus evapotranspiration by 
trees on the undisturbed watershed apparently led to the dry streambed.  Flow was       
present in all of the other streams throughout the study.  Values for water temperature in 
S 1 were generated for the dates when the stream was dry.  Multiple regressions of air 
and soil temperatures on mean water temperature from the site were used to calculate 
these values (r?=0.9, n= 51) (Stoneman and Jones 1996, Livingstone and Lotter 1998).  
Linear regression of water temperatures measured in situ on each sampling date and those 
measured through the flumes were also highly correlated (r?=0.9, n=17). 
B.  Water Temperature and Dissolved Oxygen 
Solar radiation was not measured in this study.  However, because the small      
nature of the watersheds it was expected that the angle of the sun had similar effect on 
each watershed and each stream, therefore differences may be the effect of treatments.  In 
all streams except S 3 water temperatures during the pre-disturbance period ranged from 
a low of 4?C in the winter to a high of 25?C in the summer (Figure 3).  The range of    
water temperatures in S 3 during this period was not as wide as that in the other streams.  
The lowest temperature recorded in S 3 was 15?C during the winter and the highest was   
20.5?C measured in the summer.  The narrow range of seasonal temperatures in S 3 was 
 
24
apparently the result of underground springs in the upper reaches of the watershed.     
Discharge data from the five streams provided further evidence that S 3 was spring fed.  
Daily average discharge in S 3 was three times greater than that in the other streams (Ta-
ble 2).  
Total precipitation measured at the substation near all watersheds in 1993, 1994 
and 1995 was 1,504, 1,272 and 1,478 mm respectively (Unpublished Data, Alabama    
Experiment Lower Coastal Plain Substation, Camden, AL).  However, 41% of the rain 
 
Site ---> 
Preparation
Harvest--->
0
5
10
15
20
25
30
35
Aug-93
Oct-93
Dec-93
Feb-94 Apr-94 Jun-94
Aug-94
Oct-94
Dec-94
Feb-95 Apr-95 Jun-95
Aug-95
Oct-95
Dec-95
Water Temperature (?C)
S 1 S 2 S 3
S 4 S 5
 
Figure 3.  Water temperatures for the five streams (S1...S5) in Wilcox County, Alabama. Values 
were measured at the upper station in each stream on each date. 
 
 
 
Table 2.  Total precipitation and mean daily water discharge from five 
watersheds in Camden, Wilcox County, Alabama during 1995. 
Watershed Total Rainfall (mm) Mean Daily Discharge (m
3
) 
1 1360 24.4 
2 1327 31.7 
3 1375 95.4 
4 1229 21.4 
5 1211 22.8 
 
 
25
measured in 1995 was collected in the last three months of the year and included the     
effects of Hurricane Opal.   
Water temperatures measured in S 1 were similar to those measured during the 
same period prior to disturbance.  After harvest between March and August maximum 
temperatures in S 2 and S 4 with no SMZ were significantly higher (p<0.05) than values 
measured in S 1 (Figure 4).  Temperature increases in S 2 were smaller than those        
occurring in S 4, probably because harvest on W 2 was not as complete as that on W 4.  
Mechanical harvest was impossible on the steep banks of S 2 therefore manual removal 
of the trees was required but not until late July 1995.  Riparian trees on the steep banks of 
  S 2 provided some shading to the stream. This data certainly reflected the importance of 
the SMZ in reducing temperature changes in streams of clearcut watersheds. 
After harvest, water temperatures in S 5 with an SMZ was not modified signifi-
cantly  (p>0.05) from predisturbance conditions indicating that the riparian vegetation 
was sufficient to prevent temperature changes (Figure 5).  The significant difference    
observed between S 1 and S 3 after harvest was probably due to the fact that S 3 is 
springfed and its temperature remained relatively constant year-around.  The negative  
difference observed between S 3 and S 1 reflected the fact that temperatures in S 3 did 
not increase during the summer as they did in S 1.  The shift in mean temperature in S 1 
was from around 10? to 30?C, while changes in S 3 averaged only 17? to 19?C because of 
the spring influence.  
When riparian vegetation is logged, water temperatures in streams usually in-
crease during summer and may decline in winter (Graynoth 1979).  In this experiment the 
 
26
 
Figure 4.  Water temperatures for the reference stream S 1 and S 2 and S 4 with no SMZ.  The in-
ter-ecosystem differences between each experimental stream and the reference plus the mean in-
ter-watershed differences are shown for the three phases of the study, pre-disturbance (D-Pre), 
post-harvest (D-Har) and post-site preparation (D-Site).  Each period, post-harvest and post-site 
preparation was analyzed against the pre-disturbance period. 
 
27
  
Figure 5.  Water temperatures for the reference stream S 1 and S 3 and S 5 with an SMZ.  The in-
ter-ecosystem differences between each experimental stream and the reference plus the mean in-
ter-watershed differences are shown for the three phases of the study, pre-disturbance (D-Pre), 
post-harvest (D-Har) and post-site preparation (D-Site).  Each period, post-harvest and post-site 
preparation was analyzed against the pre-disturbance period. 
 
28
most conspicuous effect was increased water temperatures observed during the summer 
because no sampling was conduced after December 1995.  Similar temperature effects 
have been reported from studies conducted in other parts of the USA.  A difference of     
7?C in monthly maximum temperature between clearcut and reference streams was found 
in New England (Noel et al. 1986).  Lee and Samuel (1976) observed a mean temperature 
increase exceeding 4?C but maximum increases exceeding as much as 9?C in a similar 
study in West Virginia.  In a coastal stream in Oregon maximum monthly temperature 
was 7.8?C higher than those recorded before logging, while maximum temperatures     
observed after logging was 29.4?C compared to 13.8?C prior to the clearcut (Hansmann 
and Phinney, 1973).  Also, from an Oregon stream, Brown and Krygier (1970) reported 
an increase of about 8
o
C after harvesting its small watershed. 
After site preparation, water temperatures in the streams from the chemically 
treated watersheds W 2 and W 3 were significantly different from the reference only in    
S 2 (Figures 4 and 5).  This difference was probably related to the treatment effects of the 
herbicide on the riparian vegetation grown following harvest.  However, in S 3 water 
temperatures varied little from pre-disturbance periods because of the spring influence.  
After mechanical site preparation stream temperatures were not significantly different in 
S 4 compared to S 1.  The movement of debris (windrowing) during mechanical site 
preparation on W 4 did not affect the new growth of riparian vegetation.  Following site 
preparation on W 4 dense growths of weeds and small shrubs remained on the banks all 
along the channel of S 4.   Shade provided by this new growth of shrubs apparently       
reduced temperature differences between S 4 and S 1 (Figure 4).  In S 5 the SMZ          
apparently was wide enough to prevent stream temperatures from increasing significantly 
 
29
during the fall.  In addition, deciduous hardwoods dominated riparian vegetation along    
S 5 while 56% of trees in W 1 were pines (Marshall 1999).  The loss of leaves as fall    
approached probably allowed more solar radiation to reach S 5 causing higher tempera-
tures than those measured in S 1 but not statistically different. 
Burton and Likens (1973) found that water temperatures were constant in a 625-m 
stream section prior to harvest; but after a clearcut conditions were altered producing a 
stream section with zones of rapid cooling and warming water.  Similar situations were 
observed during this study when the loss of canopy cover influenced water temperatures 
in the streams.  These changes were more conspicuous within the clearcut areas than 
downstream.  For example in S 4 temperatures taken 300-m downstream from the        
clearcut during mid-summer were 7?C cooler than that measured inside the clearcut area. 
Burton and Likens (1973) also suggested that small headwater streams have the capacity 
to recover quickly and return to normal temperature.  During this study only small non-
significant increases in water temperature were observed below the clearcut area in all 
streams.  Moreover, because of the natural aspect of these small watersheds, the position 
of the sun affected all watersheds in a similar manner.  Temperature differences are  
indeed affected by the angle of the sun and the direction toward which a slope faces, but 
these factors could not be addressed during this study. 
Dissolved oxygen (DO) prior to disturbance was similar in all streams ranging 
from 6.2 ppm in S 4 to 13.6 ppm in S 1.  Lower DO values were recorded during low 
stream flow periods in the summer.  After harvest DO ranged from a low of 5.8 ppm in   
S 5 to a high of 9.6 ppm in S 4.  DO was significantly modified (p<0.05) in S 5 with an 
SMZ after harvest. Low flow conditions, lower periphyton biomass and higher fine  
 
30
benthic organic matter present could be responsible in part of this measurement.  No sig-
nificant differences (p>0.05) in dissolved oxygen were observed among all streams after 
site preparation. 
C.  Herbicide Fate 
Aerial application of the herbicide resulted in little Imazapyr detected in S 2 and   
S 3 on the day of application.  As a result of direct flight over S 2, the 24-hr average Ima-
zapyr concentration measured the day of treatment was 0.127 mg/L in S 2 and   essen-
tially almost zero in S 3 (Figure 6).  The minute concentrations of Imazapyr detected in   
S 3 on that day strongly suggested the mitigating effects of the SMZ.  This mitigating ef-
fect remained evident in S 3 during normal rain events over the next several weeks.  
However, normal rain events recorded during this period did result in runoff of Imazapyr 
to S 2.  Four weeks following application concentrations of Imazapyr in S 2 had declined 
to zero. 
Measurable levels of Imazapyr were not detected in water samples until the fifth 
week after treatment in S 3 (0.028 mg/L) and probably occurred then because of the     
increased surface runoff of water caused by heavy rains following Hurricane Opal on  
October 3-4, 1995.  The heavy rains from Hurricane Opal also resulted in another peak of 
Imazapyr in S 2 (0.045 mg/L).  Following Hurricane Opal, further surface runoff of   
Imazapyr into S 2 and S 3 was essentially non-existent.  In fact, no herbicide was de-
tected in water samples from S 2 or S 3 two months after the treatment.  Between 96 and 
99% of the total Imazapyr offsite movement to S 2 and S 3 occurred during Hurricane 
Opal.  
 
 
31
Figure 6.  Daily average concentrations of imazapyr (mg/L) calculated over 24-hr periods, meas-
ured in S 2 and S 3 after chemical site preparation in Camden, Alabama, 1995. 
 
Imazapyr was found to be lethal to fish and aquatic invertebrates only in concen-
trations greater than 100 mg/L (SERA 2004).  The highest concentrations measured in 
this study were a fraction of those considered lethal to aquatic biota. 
No analyses were conducted for detection of glyphosate, thus it is not known how 
much glyphosate was in streamflow during and after the application.  Glyphosate resi-
dues are not directly amenable to gas chromatography or most detection systems that can 
be used for HPLC (Personal communication, Dr. Jerry Michael, USFS, 1998).  In addi-
tion, methods for analyzing glyphosate like the EPA method 547 are time consuming and 
specialized equipment is required.  Besides, glyphosate is strongly adsorbed to soil, re-
maining in the upper soil layers and has a low propensity for leaching (Gerritse et al. 
 
32
1996; Morillo et al. 2000).  Finally, considering the proportions of the herbicides applied 
in this study, if some glyphosate reached the experimental streams it is likely that it was 
in concentrations below the levels harmful to aquatic fauna (Henry et al. 1994). 
D.  Nitrogen and Phosphorous 
Nutrient content in the watershed soil usually determines the concentrations in the 
stream flowing through that watershed.  Nutrient analyses of soils from the Camden     
watersheds before clearcut indicated concentrations of N, P and K typical for the Coastal 
Plain province (Personal communication, D. Marshall, 1998).  Stream measurements of 
total nitrate-nitrogen (NO
3
-N) and total phosphate-phosphorus (PO
4
-P) taken before    
harvest were variable but were relatively high compared to other small streams. Total    
NO
3
-N concentration was 1.5 mg/L, or higher in 29% of the samples and mean values in 
S 1 and S2 were an order of magnitude higher than in S3, S4 and S5 (Table 3).  
Higher values of NO
3
-N in S 1 and S 2 may have been related to runoff from 
cropland and pastures located on the headwaters of these watersheds.  As the summer 
months approached dense vegetation in W 1 apparently reduced nitrogen, accounting in 
 
Table 3.  Weekly mean total nitrate-nitrogen and total phosphate-phosphorous (mg/L) in 
study streams in Camden, AL between October 1994 and November 1995. 
Pre Disturbance Post Harvest Post Site Preparation 
 
Stream / treatment 
N P N P N P 
S1-Undisturbed Reference 0.508 0.03 0.096 0.00 0.003 0.00 
S2-NO SMZ, Chemical  0.577 0.02 0.232 0.01 0.318 0.00 
S3-SMZ, Chemical 0.017 0.02 0.090 0.00 0.021 0.01 
S4-NO SMZ, Mechanical 0.013 0.68 0.077 0.10 0.036 0.03 
S5-SMZ, Mechanical 0.003 0.00 0.179 0.04 0.092 0.12 
 
 
33
part, for its decline in S 1 to about half of fall-winter values.  NO
3
-N in S 1 declined 
throughout this study.  
After harvest, NO
3
-N concentration in S 2 declined to about one third of pre-
disturbance values.  However, in S 3 and S 4 nitrogen concentrations increased about five 
times and in S 5 the increase was almost sixty times that measured prior to disturbance.  
The nitrogen decrease in S 2 (and S1) suggested depletion by vegetation in each water-
shed as values approached those measured in the other streams. In S 2 the NO
3
-N de-
crease was also probably related to the uptake by the riparian vegetation left along the 
steep banks of the streams that were not harvested until late July 1995.  In addition,    
considerable growth of shrubs and vines developed by the time all trees were harvested in 
W 2.  This vegetation apparently used nitrogen before it could reach S 2, as in the undis-
turbed stream.  Increases in NO
3
-N in S 2 at the end of the study probably reflected the 
lack of vegetation following chemical treatment.  Nitrogen increases in S 3, S 4 and S 5 
after harvest probably resulted from the lack of vegetation in those watersheds to take it 
up. 
After site preparation the SMZ may have played a filtering role for NO
3
-N      
concentrations in S3 where levels declined almost 80% from post harvest values.  Nitrate-
nitrogen in S 3 following site preparation was similar to levels measured prior to distur-
bance.  Compared to post-harvest values, NO
3
-N concentrations in both S 4 and S 5     
decreased 60% following mechanical treatment.  However, even with no SMZ, S 4 had 
vegetation growing along the stream banks that mechanical site preparation did not      
disturb.  Among all streams, nitrogen concentrations in S 4 after site preparation were the 
closest to pre-disturbance values.  Vegetation in the SMZ along S 5 probably played a 
 
34
role in the use of nitrogen before it reached this stream.  Nevertheless, nitrate-nitrogen in 
S 5 following mechanical site preparation was three orders of magnitude higher than  
pre-disturbance values.  Measurements taken at the end of the experiment in December 
1995 revealed NO
3
-N values in S 1 (.023 mg/L) and S 3 (0.657 mg/L) comparable to  
values reported in a study by Haefner and Wallace (1981).  Water samples from S 4 and 
S 5 had lower levels of nitrogen than those in S 1 and S 3, but S 2 had some of the     
highest measurements observed during this study with an average of 4.9 mg/L in Decem-
ber 1995.  
Total PO
4
-P concentrations before harvest were similar in streams S 1, S 2, S 3 
and S 5 with values of 0.03 mg/L or smaller (Table 3).  However, concentrations in S 4 
averaged 0.68 mg/L.   PO
4
-P concentrations remained low through the study at all 
streams and declined in S 4 to similar levels.  In addition, an unexplained slight increase 
in PO
4
-P concentration was observed in S 5 during the study.  However, these changes 
seem to have been not related to the watershed treatments.  Water analyses of samples 
collected from the Camden streams sites 2 years after site preparation revealed NO
3
-N 
and PO
4
-P in concentrations <1 mg/L (Unpublished Data, AU Rivers and Reservoirs 
Laboratory). These values were comparable to those typically found in lower order 
streams (Swank and Douglas 1975, Cuffney et al. 1984). 
Greater nutrient concentrations in streams have been found in different parts of 
the USA up to 3 years after site preparation when compared to undisturbed reference 
sites; even when the clearcuts left an SMZ (Martin and Pierce 1980, Feller and Kimmins 
1984, Blackburn and Wood 1990).  However, a study from the lower Coastal Plain of 
 
35
Georgia found no significant differences of NO
3
-N in groundwater regardless of the    
forest management technique used (Hubbard and Lowrance 1997).    
E.  Streambed Substrate 
Before harvest, the substrate in all streams was similar at all stations with gravel 
and sand making up 71% to 86% of the streambed (Figure 7).  Gravel ranged from a low 
of 33% in S 2 to a high of 52% in S1.  Sand was lowest in S5 and highest in S 3.  Pebbles 
composed about 14% at all sites except S 2 where it was 28%.  Cobble and silt provided 
no more than 10 % of the total substrate in all streams except in S 3 where silt was 15%.   
After harvest gravel and sand still comprised the greater percentage (72-98%) of 
substrate material at all streams including S 1; but the proportion of gravel to sand was 
slightly higher compared to pre-disturbance conditions except in S 3 (Figure 7).  Pebbles 
were reduced in all streams to a small percentage and silt increased reaching a high of 
20% in S 4.  Percentage of sand increased in S 3 after harvest going from a 29% to a 
51%.  Field observations revealed the existence of several gullies inside the SMZ          
facilitating the movement of sediments into S 3.  In addition, Marshall (1999) found    
significantly greater erosion after harvest in all clearcut watersheds compared to pre-
disturbance conditions. Substrate composition after harvest at stations 300-m below the 
clearcuts showed a seven percent increase in sand in treatments without an SMZ but no 
change was observed in the treatments with an SMZ.  
During the fall of 1995 stream substrate was more evenly composed in all streams 
with evident presence of sand and cobble (Figure 7).  The shift in substrate composition 
in the streams during this period may have not been related to site preparation treatments 
because excess sand and cobble was also observed in the reference stream with the      
 
36
undisturbed watershed. A possible explanation for this incidence is the passing of Hurri-
cane Opal in October 1995.  Opal deposited 210 mm of rain in a 48-h period and resulted 
in heavy surface run-off in the area.  Stream channels after Opal had extra sand and/or 
silt, noticeable while walking along the stream banks. 
 
 
Percent Substrate Composition
 
  
Figure 7.  Mean substrate composition from five streams at Camden, Alabama during 1995 prior 
to disturbance, after timber harvest and after site preparation. 
   
 
 
37
Compared to predisturbance conditions, the percentage of silt also increased after 
site preparation at stations below the clearcuts but the chemically prepared sites showed 
smaller increases in silt than those mechanically prepared.  In general substrate  
composition below clearcuts did not change markedly during 1995.  Thus, the dominance 
of gravel indicated good substrate for periphyton and macroinvertebrate communities.  
Sediment movement can be dramatic in small Coastal Plain streams and heavy rains    
following site preparation resulted in large amounts of sediments moved, thus uncovering 
more cobble in stream channels. No statistical analysis to support changes in substrate 
composition related to the logging operation were conducted due to the lack of seasonal 
sampling before clearcut. 
F.  Benthic Organic Matter 
Total benthic organic matter (BOM) prior to harvest was similar in all streams 
with fine benthic organic matter (FBOM) comprising 86 to 96% of the total, large benthic 
organic matter (LBOM) comprised 4 to 13% and woody debris (WD) provided only 3% 
or less (Figure 8).  Similar findings were reported from small streams in North Carolina 
(Golladay et al. 1989) although they also found sites where LBOM was dominant. 
Mean total organic matter, measured in 1995 before harvest, as ash free dry mass 
(AFDM) was variable but within ranges found in streams of similar order.  For instance, 
of the five Camden streams, S 2 had the smaller amount of organic matter at 1,329 g 
AFDM/m? (Table 4). These values were similar to that reported in Carpenter Branch,  
North Carolina of 1,135 g AFDM/m? by Golladay et al. (1989).  Total BOM in S 4 was 
1,832 g/m? essentially the same as the 1,831 g/m? found in a comparable stream at     
Sawmill Branch in North Carolina; among all sites the highest BOM value of 2,759 g/m? 
 
38
recorded in S 3 (Table 4) was still within the range identified by Golladay et al. (1989) 
for small streams.  The higher BOM found in S 3 probably reflected the influence of 
springs that maintained more constant water levels in this stream. 
 
Benthic Organic Matter (g/m? as
 AFDM)
 
  
Figure 8.  Benthic organic matter composition (as g/m? of AFDM) in five streams in Camden, 
Alabama before timber harvest (top), after harvest (middle) and after site preparation (bottom) 
during 1995 (FBOM = Fine Benthic Organic Matter, LBOM = Large Benthic Organic Matter,  
WD = Woody Debris). 
 
 
 
39
After harvest BOM decreased in all streams that received a clearcut, except for    
S 5 (Table 4).  BOM decreased more in S 2 and S 4 (streams left with no SMZ) than in   
S 3 and S 5 with an SMZ.  BOM actually increased in S 5 following the clearcut harvest. 
BOM decrease was also measured from the samples collected at S 1.  Therefore, BOM 
decreases may have been related to the time of the year when this sampling occurred 
(Spring to mid-Summer).  In addition, lack of rains during this period may have also     
influenced low BOM measurements.  BOM increases in S 5 may have been related to the 
fact that the dominance of deciduous trees over pines was greater in W 5 than in the other 
watersheds (Marshall 1999).  
 Also, the reach of S 5 sampled for BOM was a long low gradient section and   
organic matter may have had greater accumulation in this stream channel than in the 
other experimental streams.  The decline in BOM in S 3 following harvest was probably 
related to the clearcut.  There was no good explanation for the small decrease in BOM in 
S 4 because all of the trees 4 inches in diameter, or greater, were removed to the edge of 
the stream bank in this watershed. 
 
Table 4.  Mean benthic organic matter (g AFDM/m?) in the five streams in Camden, AL, 
during 1995 prior to disturbance, after timber harvest and after site preparation within the 
clearcut area. 
Station Treatment Pre Disturbance Post Harvest Post Site Preparation 
   
S 1 Undisturbed Reference 1,687   688 2,172 
S 2 NO SMZ, Chemical  1,329   972 1,185 
S 3 SMZ, Chemical 2,759 2,456 1,977 
S 4 NO SMZ, Mechanical 1,832 1,589 1,779 
S 5 SMZ, Mechanical 1,440 2,874 3,788 
     
 
 
 
40
Following site preparation, BOM increased in all streams from that measured  
during the post harvest period except at S 3 (Table 4).  BOM increases were, in large part, 
natural occurrences since sampling took place during autumn months.  In addition, high 
BOM after site preparation could be related to heavy rains (e.g. Hurricane Opal)  
following dry periods that flushed more organic matter into the stream channels.  The 
dramatic changes in BOM in S 1 may have been a natural variation in this undisturbed 
watershed.  BOM declines in S 3 after site preparation could be related to less organic 
matter input due to the clearcut in W 3 plus the higher presence of evergreen magnolia 
trees and the SMZ.  Conversely, the herbicide applied to W 2 and the mechanical prepa-
ration on W 4 and W 5 apparently resulted in a great deal of dead vegetation on these  
watersheds.  These activities contributed to more BOM runoff into the streams.  
G.  Periphyton Chlorophyll 
Except for S 3, chlorophyll a values in all streams were similar before harvest 
with mean values ranging from 0.3 to 1.7-mg/m? (Table 5).   Chlorophyll a values in S 3 
averaged over four times higher than that in the other streams perhaps because of the 
spring-influenced more stable flow and temperatures year around.  Low values of chloro-
phyll a per m? are common in low order streams with a dense canopy because little solar 
radiation reaches the stream bottom (Lowe et al. 1986).  For example, DeNicola et al. 
(1992) found only 0.01 to 0.33 mg/m? in small streams in eastern Nebraska, and Mulhol-
land and Rosemond (1992) reported 0.07 to 0.61 mg/m? in deciduous-forest streams in 
eastern Tennessee.  
Periphyton communities responded rapidly to changes in water temperature in 
these small, low gradient streams.  During a two-week period in February 1995 prior to 
 
41
harvest (Figure 9, 10, 11, 12) a warming spell occurred with maximum air temperatures 
reaching 21?C.  Historic winter maximum temperatures in the Camden area average 8?C 
and minimum of 2?C.  Soil temperatures reached highs of 17?C and lows of 13?C.  Water 
temperatures during this period reached 17?C.  The reduced canopy from leaf fall plus 
sunny days produced water temperatures that apparently contributed to unusually high 
chlorophyll a values in all streams. 
 
Table 5. Minimum, maximum and mean chlorophyll a (mg/m?) before disturbance 
(n=42) and after timber harvest (n=12), at nine stations in five streams in Camden,     
Alabama during 1993-1995.   *Value estimated from one sampling date (n=3) because   
of lack of flow during other sampling dates after harvest.  a = within clearcut, b=below   
clearcut. 
 Pre Disturbance  (Aug 1993 ? Feb 1995)  Post Harvest  (Mar 1995 ? Aug 1995) 
 Min Max Mean  Min Max Mean 
        
S 1  0.0   2.8 1.2  3.3*    3.3*   3.3* 
S 2a 0.2   8.1 1.7  4.1 28.0 16.3 
S 2b 0.5   4.2 1.6  2.1 12.2  8.7 
S 3a 1.2 14.5 7.4  7.5 64.4 37.1 
S 3b 3.0 13.2 7.5  9.1 82.2 38.0 
S 4a 0.0   3.6 0.9  0.4 99.6 44.8 
S 4b 0.0   4.6 0.9  0.0 31.4 14.7 
S 5a 0.0   0.9 0.3  0.2 17.8 10.6 
S 5b 0.2   4.0 1.2  1.9 19.6 11.9 
   
After harvest, mean chlorophyll a measurements from the two sites with no SMZ, 
S 2 and S 4, were significantly higher (P<0.05) than that measured in the reference 
stream (Figures 9, 11 and 13). These increases in chlorophyll a in S 2 and S 4 apparently 
resulted from the reduced canopy cover and subsequent increased sunlight reaching the 
stream bottom, and/or the availability of more nutrients.  For example, nitrogen and 
phosphorous values were relatively high in S 2 and S 4; S 2 registered the higher NO
3
-N  
 
42
 
Figure 9.  Chlorophyll a (mg/m?) concentrations in S 2 (No SMZ) inside (clear bar) and below (dark 
bar) the clearcut and in S 1 (line) in Camden, Alabama from August 1993 to December 1995. 
 
 
Figure 10.  Chlorophyll a (mg/m?) concentrations in S 3 (SMZ) inside (clear bar) and below (dark 
bar) the clearcut and in S 1 (line) in Camden, Alabama from August 1993 to December 1995.   
 
43
 
Figure 11.  Chlorophyll a (mg/m?) concentrations in S 4 (No SMZ) inside (clear bar) and below 
(dark bar) the clearcut and in S 1 (line) in Camden, Alabama from August 1993 to December 1995. 
 
 
Figure 12.  Chlorophyll a (mg/m?) concentrations in S 3 (SMZ) inside (clear bar) and below (dark 
bar) the clearcut and in S 1 (line) in Camden, Alabama from August 1993 to December 1995.   
 
44
Chlorophyll a 
(mg/L
-1
)
 
I
n
t
e
r-E
co
sy
st
em 
D
i
ffe
r
e
nc
es
 
Chlorophyll a 
(mg/L
-1
) 
I
n
t
e
r-E
co
sy
st
em 
D
i
ffe
r
e
nc
es
 
 
Figure 13.  Mean chlorophyll a measured in a reference stream S 1 and two experimental streams 
S 2 and S 4 with no SMZ in Camden, Alabama.  The inter-ecosystem differences between each 
experimental stream and the reference plus the mean inter-watershed difference are shown for the 
three phases of the study pre-disturbance (D-Pre), post-harvest (D-Pre) and post-site preparation 
(D-Site).  Each period, post-harvest and post-site preparation was analyzed against the pre-
disturbance period. 
 
 
45
Chlorophyll a 
(mg/L
-1
)
 
I
n
t
e
r-E
co
sy
st
em 
D
i
ffe
r
e
nc
es
 
Chlorophyll a 
(mg/L
-1
) 
I
n
t
e
r-E
co
sy
st
em 
D
i
ffe
r
e
nc
es
 
 
 
Figure 14.  Mean chlorophyll a measured in a reference stream S 1 and two experimental streams 
S 3 and S 5 with an SMZ in Camden, Alabama.  The inter-ecosystem differences between each 
experimental stream and the reference plus the mean inter-watershed difference are shown for the 
three phases of the study pre-disturbance (D-Pre), post-harvest (D-Pre) and post-site preparation 
(D-Site).  Each period, post-harvest and post-site preparation was analyzed against the pre-
disturbance period. 
 
 
46
concentration and S 4 registered the higher PO4-P values during this period of time.     
Periphyton biomass values from S 2 within the clearcut were five times higher than those 
measured in   S 1, while values from S4 were 13 times higher (Table 5). 
The largest amount of chlorophyll a, about 100 mg/m? was measured at S 4a (no 
SMZ) in August 1995.  By this time large masses of filamentous algae were observed 
covering almost the entire channel of S 4. Following harvest the chlorophyll a measure-
ments in both S 3 and S 5 were significantly higher (P<0.05) than that measured in the 
reference stream S 1 (Figure 10, 12 and 14). Chlorophyll a increases in the stream 
reaches downstream from the clearcuts were smaller than those occurring within the   
clearcut, but still ranged from five to almost twenty times higher than measurements     
recorded before harvest. 
These increases in chlorophyll a after harvest were higher statistically significant 
(P<0.05) from values recorded prior to harvest (Figure 14).  Chlorophyll a values at S 4b 
were about one eighth that of measurements at S 4a indicating that the dense canopy     
reduced light to the stream bottom (Table 5).  Increases of chlorophyll a after timber   
harvest have been reported previously but usually of smaller magnitude than those   
measured in this study.  For example, Murphy and Hall (1981) reported greater chloro-
phyll a accumulation (up to 36.6 mg/m?) in 5-17 year old clearcut sections of streams in 
Oregon, compared to old growth forested sections.  In Oregon, Gregory (1980) reported 
stream reaches inside clearcut areas had about twice the annual primary productivity of 
undisturbed streams.   
Periphyton cell densities up to six times higher were found in clearcuts compared 
to undisturbed references in 1
st
 and 2
nd
 order streams in New England (Noel et al. 1986). 
 
47
Lowe et al. (1986) found chlorophyll a concentrations up to 20 times higher in 2
nd
 order 
streams from timber-harvested watersheds in North Carolina compared to reference sites. 
Those studies were conducted more than 2 years after harvest, but the difference in pri-
mary productivity was associated with the clearcut disturbance. 
Algal production is by nature highly variable in small streams; however, distur-
bance effects in streams often cause changes in algal biomass.  The increases in chloro-
phyll a observed in all the experimental streams in this study were undoubtedly a conse-
quence of the clearcut disturbance.   
After harvest, the reduced canopy cover allowed more sunlight to reach the 
stream channels and the increased water temperatures had a positive effect on the         
periphyton growth.  Chlorophyll a values from dense periphyton communities is usually 
correlated with canopy cover (De Nicola et al. 1992) and is, in general, greater in streams 
with an open canopy than in streams with a closed canopy (Lowe et al. 1986, Feminella 
et al. 1989, Corkum 1996).  In addition, temperature is well known as a major controlling 
physical factor for organisms and is very important in determining both the community 
structure and the temporal succession of organisms (Hynes 1972).  The February 1995 
warming trend was an example of the rapid positive effect of temperature on periphyton 
communities. 
The availability of nutrients has often been pinpointed as a limiting factor for    
periphyton production in small headwater ecosystems (Stockner and Shortreed 1978, Pe-
terson et al. 1983, Grimm and Fisher 1986, Hart and Robinson 1990).  Increased nutrients 
such as potassium and nitrate-nitrogen through runoff following clearcuts were docu-
mented by Lynch et al. (1985) in central Pennsylvania.  Nutrient concentrations 
 
48
above the typical range for these elements were measured at all streams in this study, 
therefore augmenting the possibilities for increases in periphyton biomass. 
Mean chlorophyll a measured in the reference stream (S 1) during the autumn of 
1993, 1994 and 1995 showed no differences between seasons.  Following site preparation 
in the fall of 1995, within the treated watersheds, chlorophyll a values were three to 
eighteen times higher than those measured during similar time periods before disturbance 
(Table 6).  However, RIA detected significant differences only at stations S 2a and S 5a 
where differences in autumn values of mean chlorophyll a/m? before and after site prepa-
ration were larger. 
Table 6. Mean chlorophyll a (mg/m?) before site preparation (11/93, 12/93,10/94, 
12/94, n=12)  and after site preparation (10/95, 12/95, n=6) and percentage 
increase from stage to stage for the reference stream and the sites within each 
clearcut watershed. 
Station Treatment Pre Disturbance Post Site Preparation Percentage Increase 
    
S 1c Undisturbed Reference 1.3 1.3 0 
S 2a NO SMZ, Chemical 0.8 9.3 1063 
S 3a SMZ, Chemical 6.5 16.7 157 
S 4a NO SMZ, Mechanical 1.2 6.6 450 
S 5a SMZ, Mechanical 0.2 3.6 1700 
     
 
Changes in periphyton occur naturally with seasons in temperate latitudes increas-
ing in summer and decreasing during fall-winter (Rosemond, 1994; Rosemond, et al. 
2000).  Site preparation usually takes place in late summer in the Southeastern USA, 
hence the effects of this type of disturbance on stream communities has to be separated 
from natural seasonal changes.  Based on the chlorophyll a data from autumn 1994 and 
1995 (Figures 9, 10, 11, 12) periphyton communities apparently died back in all streams 
during the fall of 1994.  However, mean chlorophyll a values in 1995 in S 2 and S 3    
 
49
remained higher than those measured in S 4 and S 5.  The herbicide mixture apparently 
had no detrimental effect on periphyton communities during this research.  In addition, 
the stream with an SMZ had smaller decreases in chlorophyll a measurements than that 
without an SMZ.  Percentage increase of chlorophyll a suggested that chemical treatment 
created less disturbance than mechanical site preparation, and that the SMZ mitigated the 
disturbance (Table 6). 
Mechanical site preparation apparently had the greater impact on periphyton 
communities but the treatment with an SMZ had greater change than the one without an 
SMZ.  Evidently, the SMZ width was not adequate to prevent increases in periphyton 
biomass.  In a study testing different SMZ widths, Davies and Nelson (1994) found that 
periphyton significantly increased only in SMZ smaller than 30 m.  The SMZ in this 
study were only ~11-m wide. 
H.  Macroinvertebrate Communities 
A total of 178 taxa were identified from all streams during this study (Appendix 
I).  Aquatic insects in the order Diptera, mostly Chironomidae (midges), comprised 56% 
of the total fauna.  Mayflies in the order Ephemeroptera comprised 11% of the fauna, and 
the remaining insect orders each had less than 10%.  Miscellaneous other aquatic inverte-
brates included mostly amphipods, copepods, crayfish, water mites, nematodes, and     
turbellarians.   
For the eight dates with detailed identification, the mean density for each site   
appears in Table 7.  These data cover predisturbance, post harvest and post site prepara-
tion time periods.   Mean numbers for the reference stream S 1 represent fewer dates    
because the stream was dry in September 1994, prior to harvest, and between May and 
 
50
September 1995 during the post harvest period.  Dipterans in Table 7 were mostly taxa in 
the family Chironomidae.  More sensitive taxa in the Ephemeroptera, Plecoptera and 
Trichoptera (EPT) orders were well represented at all stations. 
 
Table 7.  Major insect orders and mean number of macroinvertebrates collected from 
core samples at sites in Camden, Alabama from August 1993 to December 1995. 
    Stations    
Order 1 C 2 A 2 B 3 A 3 B 4 A 4 B 5 A 5 B 
Aquatic Insects 
         
Coleoptera      52      379     324      52     151 470 131  88  83 
Diptera 646  3,624 3,715 359 344 2,241 1,039  1,159  1,364 
Ephemeroptera 53  486 432 145 134 579 300  324  411 
Megaloptera 0  2 0 1 0 0 0 0 0 
Odonata 1  9 12 4 1 3 3  5  4 
Plecoptera 257  100 191 16 78 106 52  40  86 
Trichoptera 37  149 95 131 180 127 173  71  169 
     
Other Aquatic Invertebrates 
       
Gastropoda 0 13 39 48 43 56 48 1  2 
Oligochaeta 183  225 117 162 122 253 128  55  175 
Pelecypoda 2  1 1 1 0 0 0 1  0 
Miscellaneous 160  881 1,029 60 113 294 219  64  198 
Total 
  
1,410  
  
5,876 
  
5,968 
  
982 
  
1,173 
  
4,142 
  
2,108  
  
1,813  
  
2,496 
 
 
 
1.  Macroinvertebrate Density 
Mean total macroinvertebrate density was determined for all seventeen dates during the 
study from both pool and riffle habitats combined (Figure 15 through 18).  During the 
pre-disturbance period, pool inhabitants among all streams comprised 22 to 40% of the 
total macroinvertebrate density, while riffle inhabitants comprised 60 to 78%. 
 
51
 
S 2 vs S 1
<-----   
131,451 
Harvest
<-----
Site -----> 
Preparation
0
25,000
50,000
75,000
100,000
Aug-93 Nov-93
Mar-94
May-94
Jun-94
Jul-94
Sep-94
Oct-94
Dec-94
Jan-95 Feb-95 Mar-95 Apr-95 Jun-95
Jul-95
Aug-95 Sep-95
Oct-95
Dec-95
No. Macroinvertebrates/m?
 
Figure 15.  Mean total macroinvertebrate density (No./ m?) in S 2 ? no SMZ and Chemically Site 
Prepared - inside (dark bar) and below (white bar) the clearcut and in the Reference Stream (line) 
in Camden, Alabama from August 1993 to December 1995.  
 
 
S 3 vs S 1
Harvest
<-----
Site -----> 
Preparation 
0
25,000
50,000
75,000
100,000
Aug-93 Nov-93
Mar-94
May-94
Jun-94
Jul-94
Sep-94
Oct-94
Dec-94
Jan-95 Feb-95 Mar-95 Apr-95 Jun-95
Jul-95
Aug-95 Sep-95
Oct-95
Dec-95
No. Macroinvertebrates/m?
 
Figure 16.  Mean total macroinvertebrate density (No./ m?) in S 3 ?SMZ and Chemically Site 
Prepared - inside (dark bar) and below (white bar) the clearcut and in the Reference Stream (line) 
in Camden, Alabama from August 1993 to December 1995. 
 
 
 
 
 
 
52
S 4 vs S 1
Harvest
----->
Site ----> 
Preparation
0
25,000
50,000
75,000
100,000
Aug-93 Nov-93
Mar-94
May-94
Jun-94
Jul-94
Sep-94
Oct-94
Dec-94
Jan-95 Feb-95 Mar-95 Apr-95 Jun-95
Jul-95
Aug-95 Sep-95
Oct-95
Dec-95
No. Macroinvertebrates/m?
 
 
Figure 17.  Mean total macroinvertebrate density (No./ m?) in S 4 ? no SMZ and Mechanically 
Site Prepared - inside (dark bar) and below (white bar) the clearcut and in the Reference Stream 
(line) in Camden, Alabama from August 1993 to December 1995.  
 
 
S 5 vs S 1
Harvest
<-----
Site -----> 
Preparation
0
25,000
50,000
75,000
100,000
Aug-93 Nov-93
Mar-94
May-94
Jun-94
Jul-94
Sep-94
Oct-94
Dec-94
Jan-95 Feb-95 Mar-95 Apr-95 Jun-95
Jul-95
Aug-95 Sep-95
Oct-95
Dec-95
No. Macroinvertebrates/m?
 
Figure 18.  Mean total macroinvertebrate density (No./ m?) in S 5 ?SMZ and Mechanically Site 
Prepared -  inside (dark bar) and below (white bar) the clearcut and in the Reference Stream (line) 
in Camden, Alabama from August 1993 to December 1995.  
 
 
 
 
53
In general, among all streams during the pre disturbance period, riffle habitats 
were more densely populated by macroinvertebrates than the pool habitats.  Others stud-
ies have reported that riffles tend to support higher macroinvertebrate densities than pools 
(Gordon et al. 1992).    However, macroinvertebrate densities from pools and riffles in    
S 3 differed little.  S 3 was spring fed and water levels remained relatively constant with 
little change in temperature throughout the study period.  Water temperature in S 3       
remained between 15 and 20?C throughout the study.  The lower water temperatures in   
S 3 may have contributed to higher dissolved oxygen levels in pools so that macroinver-
tebrate densities were similar to those found in riffles.   
Among all streams, macroinvertebrate density was usually lower in winter, in-
creasing through spring and summer to reach maximum densities in fall (Figures 15 
through 18).  Similar results have been reported in other studies in Southeastern US 
streams (Cowell et al. 2004).  Although S 1 was dry during part of the predisturbance 
phase, on at least six sampling dates macroinvertebrate density was higher in the refer-
ence stream than in S 3, S 4 or S 5.   Run-off from crops growing in the headwaters of  
W 1 and W 2 may have contributed nitrates (Table 3) to S 1 and S 2 resulting in higher 
macroinvertebrate densities compared to S 3, S 4 and S 5. Seasonal density variations of 
smaller magnitude were observed in S 3 probably because of the lack of water level and 
temperature fluctuations during the year, as springs fed the stream providing more stable 
conditions.  
Except for S 3, mean density on most dates for sites within the clearcut and those 
below differed little during the predisturbance period.  Also, mean density in S 1 on most 
dates was similar to that in each of the other streams except S 3. An ANOVA test for 
 
54
samples taken during the predisturbance period revealed no significant (p < 0.05) differ-
ences between macroinvertebrate densities at stations within the clearcut and below.   
Following harvest, in the streams with no SMZ, S 2 and S 4, higher macroinver-
tebrate densities were found inside the clearcut areas than below, however only in S 4  
(Figure 15 and 17) were the differences significant (p < 0.05) based on an ANOVA.  This 
difference was probably related to the lack of riparian vegetation that was removed to the 
edge of the stream bank in S 4 during harvest.  Thus, much of the channel in S 4 within 
the clearcut was exposed to direct sunlight because all but 10% of the riparian vegetation 
was removed during harvest (Figure 2).  Along S 2 the percent canopy cover was reduced 
gradually over a period of several weeks because of the difficulty of removing trees on 
the steep banks of the stream.  However, by June 1995 large masses of filamentous algae 
were observed in the clearcut areas of both streams, although much more in S 4.   
After harvest, in the streams with an SMZ, S 3 and S 5 (Figure 16 and 18), an 
ANOVA revealed no significant (p < 0.05) differences for total density between the    
clearcut areas and below.  Also, little filamentous algae was observed within the clearcut 
area of S 3 and S 5 post harvest because the canopy cover was still 85 to 90%.  Total 
macroinvertebrate density in S 1 during this period was relatively low on the one date 
when sampling was possible.  
To illustrate the importance of the Chironomidae (midge larvae) in the benthic 
fauna, the density of the midges compared with the ?Others? category across all phases of 
the study appears in Figure 19.  During the entire study, Chironomidae density exhibited 
similar trends to that of the total macroinvertebrate population in all streams.  Midges 
usually comprised a major portion of the total macroinvertebrate community. 
 
55
 
Figure 19.  Mean density of Chironomidae (dark bars) and Other macroinvertebrates (clear bars) 
per m? at S 1 and within the clearcut area at S 2 - S 5 in Camden, Alabama from August 1993 to 
December 1995.   
 
 
56
Population densities of benthic macroinvertebrates in streams are often highly 
variable (Corkum 1991; Downes et al. 2000) and some densities recorded during this 
study were noticeably higher than those found in other studies.  One possible explanation 
was the small mesh size used in this study (250-?m) compared to that often used by other 
authors (350 to 600-?m).  Nevertheless, Corkum (1991) reported similar densities from 
large streams when using 250-?m mesh sieves.  Little published data was found on 
macroinvertebrate densities from 1
st
-order Coastal Plain streams undergoing timber     
harvest. Using a 350-?m mesh portable invertebrate box sampler, Boschung and O?Neil 
(1981) reported density of 700 organisms/m? before clearcut and 1,700/m? after harvest in 
streams in the Piedmont region of Alabama. 
2.  Effects of Clearcut and Site Preparation on Macroinvertebrate Density 
Following harvest, densities inside the clearcut area from S 4 and S 5 were sig-
nificantly higher (p<0.05) than those found in the reference stream S 1, whereas densities 
in S 2 and S 3 were not statistically different from S 1 based on RIA (Figures 20 and 21). 
 One reason for the differences in density between S 4 and S 1 was related to the sharp 
increases in density (June, July and August 1995) in S 4 compared to predisturbance val-
ues.  S 4 was one of the treatments with no SMZ.  In S 2, the other stream with no SMZ, 
macroinvertebrate densities post harvest differed little compared to those measured dur-
ing the predisturbance period (see also Figure 15).  Therefore, based on RIA, no signifi-
cant differences were detected between S 2 and S 1 (Figure 20).   Also, as mentioned ear-
lier, unlike riparian vegetation in S 4, canopy cover was reduced gradually in S 2 because 
of the difficulty of removing trees along the banks of the stream.  
 
57
 
Figure 20.  Mean density of total macroinvertebrates measured in S 1(reference stream) and 
within the clearcut of S 2 and S 4 with no SMZ.  Also shown are the inter-ecosystem differences 
between each experimental stream and the reference plus the mean inter-watershed differences 
are shown for the three phases of the study pre-disturbance (D-Pre), post-harvest (D-Pre) and 
post-site preparation (D-Site).  Each period, post-harvest and post-site preparation, was analyzed 
against the pre-disturbance period. 
 
58
 
Figure 21.  Mean density of total macroinvertebrates measured in S 1 (reference stream) and 
within the clearcut of S 3 and S 5 with an SMZ.  Also shown are the inter-ecosystem differences 
between each experimental stream and the reference plus the mean inter-watershed differences 
are shown for the three phases of the study pre-disturbance (D-Pre), post-harvest (D-Pre) and 
post-site preparation (D-Site).  Each period, post-harvest and post-site preparation, was analyzed 
against the pre-disturbance period.   
 
59
In S 5 with an SMZ, mean densities after harvest differed little from those meas-
ured during the pre-disturbance phase.  It was not clear why RIA detected significant   
differences between S 5 and S 1.  However the reason may be that on 7 of 10 dates during 
the predisturbance phase, mean densities in S 1 were higher than that measured in S 5. 
The lack of significant differences between S 3 and S 1 was probably related to 
the small variability in density observed within the clearcut in S 3 during the post harvest 
phase compared to the predisturbance period.  However, the trend was evident that sites 
in S 4 and S 2 with no SMZ had greater increases in macroinvertebrate density than S 3 
and S 5 with an SMZ.  Corkum (1991) found that benthic macroinvertebrates density and 
biomass were often greater in open unshaded areas than in reaches with overhanging 
canopies, thus reflecting differences in abundance and production of benthic producers.   
When compared with undisturbed streams, Carlson et al. (1990) reported higher 
densities of benthic macroinvertebrates from clearcut areas in Oregon 5 years after      
harvest.  Murphy and Hall (1981) found similar results 10 years after harvest in low order 
streams in Oregon.  Greater benthic densities were found in low order California streams 
with logged watersheds without an SMZ compared to undisturbed controls (Newbold et 
al. 1980).  These authors suggested that logging affected macroinvertebrate densities    
because of a shift in the food base from detritus to algal production.   In my study, based 
on mean chlorophyll a values (Table 5), higher algal biomass in S 4 and S 2 compared to 
S 1 apparently supported the higher benthic macroinvertebrate densities found in those 
streams after harvest.   
Following site preparation, the mean density of macroinvertebrates from each  
watershed receiving a clearcut was not significantly (P>0.05) different from that found in 
 
60
the reference stream S 1 (Figure 20 and 21).  The lack of significant differences among 
treatments suggested that chemical and mechanical site preparation had little effect on 
macroinvertebrate communities in these streams. Similar effects of imazapyr (Fowlkes   
et al. 2003) and glyphosate (Tu et al. 2001) on benthic macroinvertebrates have been    
reported in other studies.  
3.  Taxa Richness, Species Diversity and Community Structure 
All streams had similar taxa richness (TR) during the pre disturbance period with 
the mean number of taxa collected at a single stream ranging from a high of 86 in S 1  
during June 1994 to a low of 33 in S 3 during February 1995 (Figure 22).  RIA revealed 
no significant differences in mean taxa richness among all streams during this period.  
Across all streams, Chironomidae was the most diverse family represented with 76 total 
taxa and providing, in general, about 44 % of the taxa collected from core samples.  Taxa 
occurring in greatest numbers during the pre-disturbance period were midges of the genus 
Tanytarsus spp. and Rheotanytarsus spp., the ephemeropteran Baetis spp. the coleopteran 
Psephenus herricki and oligochaetes.  Both midges are filtering collectors that filter fine 
particulate BOM from the water column.  Baetis is a gathering collector that feeds on 
BOM in the substrate.  Psephenus herricki is a grazer that feeds by scrapping algae and 
BOM from the substrate.  Oligochaetes are also gathering collectors that feed on BOM 
(Merritt and Cummins 1996).  
After harvest, taxa richness ranged from a mean low of 37 in S 3 to a maximum of 
86 in S 2 (Figure 22).  The reference stream was dry during most sampling dates sched-
uled in the post harvest period.  The single sample collected from the reference stream 
during this period was not selected for taxa identification.  Therefore, no post- harvest 
 
61
Harvest --->
<---    Site   
Preparation
0
20
40
60
80
100
Mar-94 Jun-94 Sep-94 Dec-94 Mar-95 Jun-95 Sep-95 Dec-95
Taxa Richness
S 1 S 2 S 3 S 4 S 5
 
Figure 22.  Mean taxa richness from selected dates for macroinvertebrates collected in S 1 (refer-
ence stream) and within the clearcut of the four experimental streams. 
 
 
comparisons with S 1 were possible. Compared to previous summer, taxa richness after 
harvest was slightly lower inside the clearcuts of all experimental streams except S 2. 
Although macroinvertebrates were not identified for the one date sampled in S 1 
post harvest, RIA generated values based on predisturbance samples.  However, no      
statistical differences in taxa richness were detected among all streams post harvest. Even 
though chlorophyll a concentrations increased in all streams receiving clearcuts (Figures 
9 through 12), this change in algal biomass resulted in no increase in taxa richness. 
Following site preparation, RIA detected no statistical differences in benthic taxa 
richness among the experimental streams compared to S 1.  Based on these results, both 
chemical and mechanical site preparation techniques appeared to have no influence on 
taxa richness. Fowlkes et al. (2003) experimenting with in situ microcosms in north   
Florida, found that imazapyr had little effect on macroinvertebrate taxa richness and 
abundance. 
 
62
Richness for the more sensitive taxa in the orders Ephemeroptera, Plecoptera and 
Trichoptera (EPT) was similar among all streams during the pre disturbance period with 
S 1 having the highest richness (Table 8).  EPT richness values in the range of 12 to 14 
like those found in this study are typical in undisturbed Coastal Plain streams (Personal 
observation, E. C. Webber 2005).  During all phases of this study no significant differ-
ences in EPT richness were found among streams. 
 
Table 8.  Mean EPT index before harvest, after harvest and after site preparation at 
five streams in Camden Alabama from March 1994 to December 1995. Dash means 
no samples were collected due to drought. 
Station Pre-Disturbance Post-harvest Post-Site Preparation 
S 1c  13 --- 
8 
S 2a 12 14 
11 
S 2b 13 10 
12 
S 3a 11 8 
8 
S 3b 9 7 
6 
S 4a 12 11 
11 
S 4b 11 11 
8 
S 5a 9 7 
6 
S 5b 11 12 
5 
 
 
Imazapyr was present in the chemically treated streams S 2 from August 27 to 
October 6 1995 and briefly in S 3 (Figure 6).  The small changes in the EPT index in both 
streams seemed to suggest natural variation instead of effects from the herbicide mixture 
on these more sensitive taxa.  Kaller and Hartman (2004) found that EPT taxa richness 
significantly decreased when fine silt particles increased in the stream substrate. Except 
for S 1, more silt and sediment were observed in all the streams after both harvest and site 
preparation; hence, the possibility existed that small changes in EPT richness may have 
occurred following the disturbances.  However, based on the data collected in this study, 
 
63
changes in the EPT index appeared to reflect natural variability rather than treatment    
effects.  
Macroinvertebrate diversity was similar among all streams before harvest with 
Shannon -Weaver Diversity values ranging from a low of 3.03 in S 2 to a high of 3.79 in 
S 4 (Table 9).  Diversity values measured during the pre disturbance phase of this study 
were greater than 3.0, and 3.0 is considered typical for undisturbed and unstressed 
streams (Wilhm and Dorris 1968).  A Tukey?s test found no significant differences         
(p > 0.05) in species diversity among all streams during the pre disturbance period. After 
harvest, diversity in the experimental streams changed little from that measured prior to 
harvest. 
 
Table 9.  Mean Shannon-Weaver diversity index (H?) during pre-disturbance, post har-
vest and post site preparation in five streams near Camden Alabama from March 1994 
through December 1995. 
Station Pre-Disturbance Post-harvest Post-Site Preparation 
S 1c 3.71 --- 
3.09 
S 2a 3.03 3.50 
3.46 
S 2b 3.17 3.22 
3.05 
S 3a 3.07 3.02 
2.90 
S 3b 3.35 2.91 
2.82 
S 4a 3.25 3.25 
3.19 
S 4b 3.79 3.26 
2.95 
S 5a 3.24 3.19 
2.99 
S 5b 3.25 3.38 
3.25 
 
 
 
Diversity in S 1 was not measured during this period because the single sample 
collected was not selected for taxa identification during the random sampling.  However, 
no significant differences (P>0.05) were detected among the streams during the post   
 
64
harvest period.  Harvest apparently had no effects on macroinvertebrate species diversity. 
In east central Alabama, Boschung and O?Neil (1981) found no differences in diversity 
among streams subjected to clearcut harvest.  Diversity values in their study were similar 
to values found in this study. Also, after site preparation no significant differences were 
detected among streams, or when compared to S 1.   
Among all streams, benthic community structure throughout this study consisted 
predominantly of five functional feeding groups including collectors, both gathering 
(GC) and filtering (FC), predators (P), shredders (SH) and scrapers (SC).  During each 
phase of the study the two groups of collectors comprised 46 to 65 % of the total fauna, 
followed by predators with 7 to 27 %  of the fauna, scrapers with 12 % or less of the total 
fauna and shredders with 17% or less of the fauna (Table 10).  These findings differed 
slightly from the River Continuum Concept (RCC), which assumes that about one third 
of the total macroinvertebrate community in headwater streams usually consists of   
shredders (Vannote et al. 1980).  Even though heavy canopy cover was present along all 
streams prior to harvest; shredders comprised a relatively small percentage of the total 
fauna during this phase and throughout the study in S 1.  
Among these five functional feeding groups seven taxa tended to dominate in all 
streams throughout the study.  Gathering-collectors included species of Baetis and oli-
gochaetes.  Filtering-collectors were usually dominated by the dipterans Tanytarsus spp. 
and Rheotanytarsus spp.  Another group of small dipterans, the Ceratopogonidae were 
the dominant predators identified in samples.  These small midges probably preyed on 
tiny invertebrates such as water mites and crustaceans common in the streams. 
 
 
65
Table 10.  Percentages of the total macroinvertebrate fauna by functional feeding group 
collected before and after harvest and after site preparation, inside the clearcut areas in 
five stream in Camden, Alabama during March 1994 to December 1995.  Dash indicates 
no collection due to drought. (FC= Filtering-Collectors, GC= Gatherers-Collectors,      
P= Predators, SC= Scrapers-Piercers, SH= Shredders). 
 Pre Disturbance (n=4) Post Harvest (n=2) Post Site Preparation (n=2) 
FC GC P SC SH FC GC P SC SH FC GC P SC SH 
S 1c 30 31 7 1 2 -- -- -- -- -- 10 40 13 0  2 
S 2a 49 16 10 1 1 17 41 13  1 1 13 37  7 2   0  
S 3a 12 43 7 6 7 13 42 12 12 10  9 50  8 1 15 
S 4a 18 33 19 4 2 33 25 9  2 1 13 47 12 2  8 
S 5a 23 40 15 1 6 21 25 27  8 1  7 39 24 4 17 
 
 
Another group of dipterans, species of Polypedilum were the dominant shredders. 
 So in the streams, shredders, like the predators, were not represented by larger macroin-
vertebrates such as certain stoneflies and caddisflies, but by minute midges.  In fact,   
species of Polypedilum also feed occasionally as gathering-collectors.  Coleopterans were 
the dominant scrapers consisting of the species Psephenus herricki.  Other taxa occasion-
ally occurred in high numbers in selected streams along with these seven, however no 
trends were evident with regard to treatment effects.  
The seven taxa and their percent composition of the total fauna during each phase 
of the study appear in Table 11 for sites within the clearcut.  These taxa were also com-
mon in S 1 although data are not shown because the stream was dry during much of the 
post harvest phase.  
After harvest small changes in community structure were observed in all streams, 
except S 1 that had no data for comparable dates (Tables 10 and 11).  Scrapers increased 
slightly in S 2, decreased in S 4 and increased considerably in S 3 and S 5 in the forms of 
the caddisfly Glossosoma spp., the mayfly Habrophlebiodes spp. and the snail Elimia 
 
66
Table 11.  Percent composition of the most common macroinvertebrate taxa in the five 
dominant functional feeding groups (FFG) collected inside the clearcut areas of four 
streams in Camden, AL. before disturbance (PreDist), post harvest (PostHarv) and post 
site preparation (PostSite). 
 
   S 2 a    S 3 a  
FFG / Taxa PreDist PostHarv PostSite  PreDist PostHarv PostSite
 n=4 n=2 n=2  n=4 n=2 n=2 
Gatherer-collectors        
     Baetis spp. 2 10 14  8 11 11 
     Oligochaetes 1 5 11  12 21 22 
Filterer-collectors        
     Tanytarsus spp. 39 10 1  <1 6 <1 
     Rheotanytarsus spp. 9 4 <1  3 <1 3 
Predators       
     Ceratopogonidae 4 3 3  4 9 5 
Scrapers      
     Psephenus herricki 5 7 3  <1 <1 2 
     Habrophlebiodes spp. 1 1 2  <1 <1 <1 
Shredders       
     Polypedilum spp. <1 3 8  7 10 15 
         
   S 4 a    S 5 a  
FFG / Taxa PreDist PostHarv PostSite  PreDist PostHarv PostSite
 n=4 n=2 n=2  n=4 n=2 n=2 
Gatherer-collectors        
     Baetis spp. 8 8 3  11 16 19 
     Oligochaetes 7 1 18  2 2 10 
Filterer-collectors        
     Tanytarsus spp. 3 25 3  5 13 2 
     Rheotanytarsus spp. 5 <1 2  12 5 3 
Predators       
     Ceratopogonidae 11 1 10  7 5 15 
Scrapers      
     Psephenus herricki 9 12 2  <1 5 <1 
     Habrophlebiodes spp. 4  2 1   1 8  4 
Shredders      
     Polypedilum spp. 3 5 10  5 1 17 
         
 
 
 
67
spp.  Increases in scrapers in S 2 and S 4 were expected because of the treatment effect of 
no SMZ. However, increases in S 3 and S 5 with an SMZ may only reflect seasonal   
variability.  The lack of canopy cover and more sunlight reaching these streams allowed 
more algae growth, although significantly higher (P<0.05) chlorophyll a values were 
found in all experimental streams after harvest.  This may explain why S 5 also had      
increases in scrapers during this period.  
Shredders represented by midges of the Chironomidae, Polypedilum spp.,          
increased slightly in S 2 and S 4 with no SMZ probably because of the large masses of 
filamentous algae observed in these two streams.  These midges are known to feed on  
filamentous algae (Henriques-Oliveira et al. 2003).  The increase of shredders in S 2 and 
S 4 was the opposite effect of that expected in these streams with no SMZ. 
    Slight increases of the mayfly Baetis spp. (GC) in S 2, S 3 and    S 5 may have 
been related to the increased algal communities in all streams.  However, it was not clear 
why this increase was not observed in S 4.  Wallace and Gurtz (1986) identified           
significant increases in diatom consumption by the mayfly Baetis spp. in North Carolina 
streams after clearcut.  However, these changes in scrapers and other groups appeared to 
be more related to natural variability than harvest effects.    
After site preparation the benthic fauna in all streams continued to be dominated 
by collectors and overall the community structure was similar to that observed in the   
reference stream (Table 10).  Macroinvertebrate communities resembled those found  
during the pre disturbance sampling although shredders constituted a significant          
percentage of the total fauna in S 3, S 4 and S 5.  Overall, considering the macroinverte-
brate communities found in the fall of 1995 in all experimental streams and comparing 
 
68
them to S 1, neither chemical nor mechanical site preparation had a major influence in 
changes of macroinvertebrate community structure. 
I.  Summary 
The experimental streams in the Camden study presented interesting changes both 
after harvest and after site preparation.  Only natural conditions affected the reference 
stream with an undisturbed watershed during the entire study; therefore, changes in  
physicochemical conditions and benthic communities in S 1 apparently reflected only 
natural variations due to seasons.  Drought conditions in the reference stream weakened 
the possibility of comparing with treated streams and analyzing results after harvest.    
After harvest there were increases in sediments observed at all streams except S 1, and 
small decreases in BOM in all streams except S 5.  Higher water temperatures were 
measured in the streams without SMZ after harvest. Shifts in macroinvertebrate commu-
nities observed in the experimental streams may have reflected changes in the watersheds 
because of the clearcuts. As more sunlight reached the streams, it allowed larger primary 
producers (e.g. filamentous benthic algae) to flourish and consequently provide more 
food for primary consumers such as scrapers.  However, less percentage of scrapers was 
found in the streams without SMZ.    After site preparation sediment increases were     
observed, mainly at those sites mechanically prepared and BOM increased at all streams 
except S 3. However, some of these increases were result of the increased runoff associ-
ated with Hurricane Opal.  Algal communities increased after site preparation suggesting  
no detrimental effect of the imazapyr and glyphosate on primary producers in the 
streams; and macroinvertebrate communities were apparently not affected by either 
chemical or mechanical treatment.  Macroinvertebrate diversity and taxa richness did not 
 
69
significantly change in any stream during the study.  Overall, mechanical site preparation 
in clearcuts without SMZ appeared to be the method that most affected the biota.  Also, 
the herbicides imazapyr and glyphosate used in the concentrations applied in treatments 
with an SMZ had no harmful impacts on aquatic biota.   
The structure and function of stream communities is affected by food availability 
and habitat structure (Vannote et al. 1980).  During the Camden study no major changes 
in community structure were observed in the experimental streams and RIA did not      
detect significant differences.  The few sample dates available after site preparation could 
be one of the weaknesses when using this statistical approach.  Future studies should be 
designed with sampling conducted at least one year after disturbances (harvest or site 
preparation) in order to examine seasonal changes before and after site preparation.  This 
could reduce data interpretation problems when comparing seasonally dependant       
variables such as periphyton and macroinvertebrates.  Macroinvertebrate communities 
may recover within a few months to several years after disruption (Heckman 1983, 
Molles 1985) but some disturbances can produce long-term changes in habitats that      
recovery will not occur until the natural habitat is restored (Wallace 1990).  Furthermore, 
the recovery will depend on the spatial scale of the disturbance, position within the 
stream network, timing in relation to the life history stages of the organisms and their 
dispersal abilities (Wallace 1990). 
The use of SMZ helped mitigate some impacts to aquatic biota from the 
disturbances associated with clearcut harvest.  The 11-m SMZ used in this study was 
wide enough to avoid any significant water temperature changes among streams.  The  
11-m SMZ also greatly reduced herbicide runoff and/or leaching to S 3.  However, the 
 
70
11-m SMZ did not prevent significant increases in algal biomass in S 3 and S 5.  
Therefore, timber harvest using wider riparian zones should be tested.  For example, 
Kiffney et al. (2003) and Newbold et al. (1980) concluded that 30-m SMZ are needed to 
reduce biotic changes associated with clearcuts.  Moreover, the stream management 
zones used in this study were limited to the main channel leaving smaller tributaries 
completely exposed within the clearcuts, which allowed more runoff and consequently, 
more sediment to reach the streams.  
 
71
 
 
 
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83
 
 
 
APPENDIX I 
Table 1.  Composite list of taxa, tolerance value (TV) and functional feeding group (FFG) 
when known, presence at a sampling site (+) and total number of taxa per site collected in 
Camden, Alabama from March 1994 through December 1995. 
AQUATIC INSECTS TV FFG 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta   + + +   + + + + 
Coleoptera       + +    
  Dryopidae          
 Helichus  spp. adults 5 GC   +       
      Elmidae  5 GC + + + + + + + + + 
 Ancyronyx  varigatus   6 GC    + + +   + 
 Dubiraphia  spp.  6 SC      +  + + 
 Elmidae adults 5 SC      +  +  
 Macronychus  glabratus   4 SH        +  
 Optioservus  spp.  4 SC  + + + +  +   
 Optioservus  spp. adults 4 GC  +  + +     
 Oulimnius  latiusculus   4 SC + + + + + + +  + 
 Oulimnius  latiusculus  adults 4 GC +         
 Promoresia  spp.  4 SC       +   
 Stenelmis spp. adults 7 GC + + + + +  +  + 
      Psephenidae          
 Psephenus  herricki   4 SC + + + + + + + + + 
 Psephenus  herricki  adults 4   +        
      Ptilodactylidae          
 Anchytarsus  bicolor   5 SH + + + + +  + + + 
Collembola 3 GC + + + +    +  
  Isotomidae     + + + + + + + 
      Sminthuridae 3 GC + + +  + +  +  
Diptera   + + + + + + + + + 
 Diptera adults   + + + + + + + + + 
 Diptera pupae   + + + + + + + + + 
      Ceratopogonidae 6 P + + + + + + + + + 
 Atrichopogon  spp. 6       +    
 Bezzia/Palpomyia  gp. 6 P + + + + + + + + + 
  Chaoboridae 8 P      +    
  Chironomidae 7 GC + + +  + + + + + 
 Ablabesmyia  spp. 8 P     + + +  + 
 Brillia spp. 5 SH +         
 Chaetocladius  spp. 6 GC + +      +  
 Chironomie 7 GC     +     
 
84
Appendix I. Table 1.  Continued? 
AQUATIC INSECTS TV FFG 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera            
  Chironomidae            
 Chironomini 8 GC  + + + + + + + + 
 Cladotanytarsus  spp. 7 FC + +    +    
 Corynoneura  spp. 7 GC + + + + + + + + + 
 Cricotopus bicinctus 7 GC  + +  + +    
 Cricotopus spp. 7 GC + + +       
 Cricotopus  tremulus  gp. 7 GC + +        
 Cricotopus/Orthocladius 7 GC + + + + +  +   
 Cryptochironomus spp. 8 P  + + +    +  
 Cryptotendipes  spp. 6 GC      +    
 Demicryptochironomus  cuneatus 8 GC + + +       
 Dicrotendipes  spp. 8 GC      + +   
 Diplocladius  cultriger 8 GC +  +  +  +   
 Djalmabatista  spp. 3 P     + +    
 Eukiefferiella  claripennis  gp. 8 GC +   + + +  + + 
 Glyptotendipes spp. 10 FC        +  
 Harnischia spp. 8 GC  +     +   
 Heterotrissocladius  macidus  gp. 8 GC  +        
 Hydrobaenus  pilipes  gp. 8 GC    +      
 Krenopelopia  spp. 6 P  + +    + + + 
 Krenosmittia spp. 6 GC + + + + + + + + + 
 Labrundinia spp. 7 P         + 
 Larsia  spp. 6 P + + + + + + + + + 
 Micropsectra  spp. 7 GC   +  + + + + + 
 Microtendipes  pedellus  gp. 6 FC      +    
 Monopelopia spp. 6 P  + +  + + + + + 
 nocladius  spp. 3 GC  +     +   
 tarsia  spp. 8 P  + + +  +  +  
 Nilotanypus spp. 6 P + + + + + + + + + 
 Nilothauma spp. 2 GC  +        
 Nimbocera pinderi 6   +        
 Orthocladiie 6 GC + + + + + +  +  
 Parachaetocladius spp. 6 GC + + + + + + +   
 Parachironomus spp. 10 P +         
 Paracladopelma spp. 7      +     
 Paracricotopus spp. 6 GC      +    
 Parakiefferiella  spp. 6 GC        + + 
 Parakiefferiella  triquetra 4 GC +      +   
 Paralauterborniella nigrohalteralis 8 GC    +   + +  
 Parameri spp. 6 P +  +  + +  +  
 Parametriocnemus spp. 5 GC + + + + + + + + + 
 Parasmittia spp. 6  +         
 Paratanytarsus  spp. 6       +    
 Paratendipes  spp. 8 GC   +   + + + + 
 Pentaneura  spp. 5 GC      +    
 
85
Appendix I. Table 1.  Continued? 
AQUATIC INSECTS TV FFG 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera            
  Chironomidae            
 Polypedilum  (P.) convictum 7 SH + + + + + + + + + 
 Polypedilum  (P.) fallax 7 SH + + + + +     
 Polypedilum  (P.) illinoense 6 SH +  +  + + + + + 
 Polypedilum  (T.) halterale 7 SH  + + + + + + + + 
 Polypedilum  (T.) scalaenum  gp. 7 SH + + + + + + + + + 
 Polypedilum  spp. 6 SH + +       + 
 Potthastia  longima  gp. 4 GC     +     
 Procladius spp. 9 P      +    
 Pseudorthocladius spp. 0 GC   +  +     
 Psilometriocnemus spp. 6 GC  +        
 Rheocricotopus  spp. 6 GC + +      +  
 Rheotanytarsus  spp. 6 FC + + + + + + + + + 
 Robackia  demeijerei 8 GC + + +  +   + + 
 Saetheria spp. 4 GC      +    
 Stempelli  spp. 4 GC + + +    +   
 Stempellinella spp. 4 GC + + + +  + + + + 
 Sublettea coffmanni 6 FC         + 
 Tanypodie 6 P + + +  + + +  + 
 Tanypodie pupae 6   +        
 Tanytarsini 6 FC        +  
 Tanytarsus  spp. 7 FC + + + + + + + + + 
 Thienemannia  spp. 6  +    +  + +  
 Thienemanniella spp. 6 GC + + + + + + + + + 
 Thienemannimyia complex 6 P + + + +  + + + + 
 Tribelos spp. 5 GC +         
 Tvetenia  bavarica  gp. 5 GC + + + + + + + + + 
 Zavrelimyia  spp. 8 P   +   + +  + 
      Culicidae 8 FC     + + + + + 
 Culex  spp. 8 FC + +  +  + + + + 
      Dixidae          
 Dixa  spp. 1 GC     +     
      Empididae          
 Hemerodromia spp. 6 P + + + + + +  + + 
      Psychodidae          
 Pericoma/Telmatoscopus 10 GC  +        
      Simuliidae 6 FC     +     
 Simulium  spp. 6 FC + + + + + + + + + 
      Tanyderidae          
 Protoplasa  fitchii   +   + +     
      Tipulidae 3 SH + + +    +   
 Hexatoma  spp. 4 P + + + + + + + + + 
 Pseudolimnophila spp. 2 P + + + + + + + + + 
 Tipula  spp. 4 SH +  +  +   + + 
 Tipulidae pupae 3      +     
 
86
Appendix I. Table 1.  Continued? 
AQUATIC INSECTS TV FFG 1C 2A 2B 3A 3B 4A 4B 5A 5B
Ephemeroptera   + + +   +    
      Baetidae 4 GC      + +  + 
 Baetis  spp. 6 GC + + + + + + + + + 
      Caenidae          
 Caenis  spp. 7 GC + + + + + + + +  
      Ephemeridae          
 Hexagenia  spp. 6 GC  +    + +  + 
      Heptageniidae          
 Stenonema spp. 5 SC  + +   + +  + 
      Leptophlebiidae          
 Habrophlebia  vibrans 2 GC      +    
 Habrophlebiodes  spp. 2 SC + + + +  + + + + 
Hemiptera  P  +  +      
  Mesoveliidae          
 Mesovelia  spp.  P       +   
      Veliidae  PI   +       
 Rhagovelia spp.  PI     +     
Lepidoptera   + +   +   +  
Megaloptera          
  Corydalidae          
 Nigronia spp. 2 P  +  +      
Odonat  P       +   
  Aeshnidae          
 Boyeria  spp. 3 P  +        
      Calopterygidae          
 Calopteryx  spp. 5 P   + +   + +  
      Cordulegastridae          
 Cordulegaster  spp. 3 P        + + 
      Gomphidae 1 P  + + + + + + + + 
 Erpetogomphus  desigtus 5 P  + + +      
 Progomphus spp. 5 P      +    
 Stylogomphus  albistylus 0 P +  +       
Plecoptera   + + + + + + + + + 
      Chloroperlidae    
 Alloperla spp. 0 GC + + + +  + + + + 
  Leuctridae 0 SH +         
  Nemouridae 2 SH      +    
 Amphinemura spp. 3 SH + + +   + + + + 
      Perlidae 1 P +        + 
 Acroneuria  spp. 1 P  +        
 Eccoptura  xanthenes 1 P + + + + +  +   
Trichoptera    + + + + + + + + 
  Calmoceratidae          
 Anisocentropus  pyraloides 3 SH +  +       
            
            
 
87
Appendix I. Table 1.  Continued? 
AQUATIC INSECTS TV FFG 1C 2A 2B 3A 3B 4A 4B 5A 5B
Trichoptera            
      Glossosomatidae          
 Glossosoma spp. 0 SC + + + + + + + + + 
      Hydropsychidae 4 FC + + + + + + + + + 
 Cheumatopsyche  spp. 6 FC  + +  + + + + + 
 Diplectrona  spp. 4 FC + + + + + + + + + 
 Hydropsyche  spp. 7 FC + + + + + + + + + 
 Macrostemum  carolina 4 FC  +  +    + + 
 Macrostemum  spp. 4 FC     +  +   
      Hydroptilidae 4 PI  +    +    
 Hydroptila spp. 6 PI      +    
      Limnephilidae 4 SH         + 
 Neophylax spp. 3 SC        +  
      Molannidae 6 SC  +        
 Molanna  spp. 6 SC +  + +      
      Philopotamidae    
 Chimarra  spp. 4 FC  + + +      
      Rhyacophilidae          
 Rhyacophila spp. 4 P      +    
Amphipoda    
      Gammaridae 4 GC + +  + +     
Copepoda   + + + + + + + +  
  Calanoida   + + + + + + + + + 
      Harpacticoida   +  +  + + + +  
Decapoda 6 SH      +    
  Cambaridae 5 GC   +       
Gastropoda    + + + + + + + + 
  Pleuroceridae          
 Elimia spp. 5 SC  +  + + + +  + 
Hydracarina 5  + + + + + + + + + 
Nematoda 5  + + + + + + + + + 
Oligochaeta          
      Lumbriculidae 8 GC +       +  
  Oligochaeta 10 GC + + + + + + + + + 
Ostracoda    +  +  + + + + 
Pelcypoda  FC +   +      
  Corbiculidae          
 Corbicula fluminea 4 FC  + +     +  
Turbelaria          
      Plariidae 4  + + +   + + +  
     
Total number of taxa per sampling site 88 103 92 74 83 96 85 85 77
 
88
 
 
 
APPENDIX II 
Table 1.   Total fauna, number of taxa, and mean number of organisms/m? collected at 
each station in the Camden Study on 18 March 1994. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 00100 0 000
Coleoptera   
      Elmidae 00136 1 033
 Optioservus  spp.  00116 0 100
 Oulimnius  latiusculus   91002 3 000
 Stenelmis spp. Adults 00111 0 102
 Elmidae adults 00000 1 000
      Psephenidae   
 Psephenus  herricki   1 2 19 0 0 0 2 0 2
      Ptilodactylidae   
 Anchytarsus  bicolor   20143 0 010
Collembola   
      Isotomidae 00200 0 000
      Sminthuridae 20200 0 000
Diptera 1 2 12 1 1 2 2 2 1
 Diptera adults 30110 0 100
 Diptera pupae 23500 0 010
      Ceratopogonidae 0 14 69 7 0 10 0 1 1
 Bezzia/Palpomyia  gp. 6 13 0 4 3 18 2 5 10
      Chironomidae 00000 0 001
 Brillia spp. 10000 0 000
 Chaetocladius  spp. 02000 0 020
 Corynoneura  spp. 7 14 22 0 0 12 2 3 2
 Cricotopus  tremulus  gp. 10000 0 000
 Cricotopus spp. 30400 0 000
 Cricotopus/Orthocladius 51400 0 000
 Cryptochironomus spp. 02000 0 000
 Demicryptochironomus  cuneatus 30000 0 000
 Diplocladius  cultriger 10000 0 000
 Eukiefferiella  claripennis  gp. 00000 0 001
 Krenopelopia  spp. 04000 0 000
 Krenosmittia spp. 00403 24 041
 Larsia  spp. 20200 6 100
 Micropsectra  spp. 00000 0 010
 Nilotanypus spp. 22000 0 100
 Parachironomus spp. 10000 0 000
 
89
Appendix II. Table 1.  Continued? 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Paracladopelma spp. 00001 0 000
 Parakiefferiella  triquetra 00000 0 100
 Parametriocnemus spp. 10002 2 011
 Paratendipes  spp. 00000 0 001
 Pentaneura  spp. 00000 2 000
 Polypedilum  (P.) convictum 10 1 12 0 6 3 0 0 0
 Polypedilum  (P.) fallax 00200 0 000
 Polypedilum  (P.) illinoense 00000 1 000
 Polypedilum  (T.) halterale 0 0 22 11 3 1 0 0 7
 Polypedilum  (T.) scalaenum  gp. 12270 0 0013
 Polypedilum  spp. 01000 0 000
 Rheocricotopus  spp. 33000 0 000
 Rheotanytarsus  spp. 95800 0 1034
 Robackia  demeijerei 02009 0 010
 Stempellinella spp. 02000 0 000
 Tanytarsus  spp. 19 93 246 0 1 0 0 0 0
 Thienemanniella spp. 00045 3 110
 Thienemannimyia complex 04000 0 000
 Tvetenia  bavarica  gp. 7 4 20 8 14 57 3 11 2
 Orthocladiie 30020 1 000
 Tanypodie 02000 0 000
      Culicidae   
 Culex  spp. 41000 0 000
 Culicidae 00001 0 000
      Simuliidae   
 Simulium  spp. 243810 128 9146
      Tipulidae   
 Hexatoma  spp. 11634 0 3 5 2 3 4
 Pseudolimnophila spp. 00200 1 001
 Tipula  spp. 10000 0 002
 Tipulidae pupae 00001 0 000
Ephemeroptera   
      Baetidae 00000 2 000
 Baetis  spp. 196818 2 040
      Caenidae   
 Caenis  spp. 00002 0 000
      Leptophlebiidae   
 Habrophlebiodes  spp. 02100 1 100
Lepidoptera 01000 0 000
Odonata   
      Gomphidae 00210 0 000
   
   
   
 
90
Appendix II. Table 1.  Continued? 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Plecoptera 116 21 94 2 30 50 14 11 3
      Chloroperlidae   
 Alloperla spp. 27 5 14 0 0 6 14 3 3
      Nemouridae   
 Amphinemura spp. 40200 2 136
      Perlidae 80000 0 000
 Eccoptura  xanthenes 21012 0 000
Trichoptera 000418 0 000
      Glossosomatidae   
 Glossosoma spp. 11108 1 100
      Hydropsychidae 60011 1 013
 Cheumatopsyche  spp. 00100 0 000
 Hydropsyche  spp. 20000 0 100
Molannidae   
 Molanna  spp. 10400 0 000
      Rhyacophilidae   
 Rhyacophila spp. 00000 1 000
OTHER AQUATIC INVERTEBRATES   
Copepoda 00501 0 000
      Calanoida 54700 0 000
      Harpacticoida 10102 0 110
Amphipoda   
      Gammaridae 30000 0 000
Gastropoda 00300 0 000
      Pleuroceridae   
 Elimia spp. 00020 0 000
Hydracarina 35 27 76 1 5 1 3 0 7
Nematoda 96485 6 203
Oligochaeta 5 6 16 8 7 9 4 0 16
Turbellaria   
      Plariidae 00000 2 000
   
Total number of taxa per station 
46 37 43 24 31 33 25 22 27
Total number of organisms per station 379 279 746 84 160 365 72 77 136
Mean number of organisms per m? 27128 19970 53396 6012 11452 26126 5154 5511 9734
 
 
91
Appendix II. Table 2.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 20 June 1994. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 00000 1 001
Coleoptera   
      Elmidae 104612 3 4159
 Ancyronyx  varigatus   00010 0 000
 Optioservus  spp.  00009 0 000
 Optioservus  spp. adults 00021 0 000
 Oulimnius  latiusculus   30012 2 800
 Stenelmis spp. adults 12110 0 000
      Psephenidae   
 Psephenus  herricki   4 142 48 1 0 1 1 0 1
 Psephenus  herricki  adults 04000 0 000
      Ptilodactylidae   
 Anchytarsus  bicolor   43518 0 000
Collembola   
      Isotomidae 00004 3 313
Diptera 20201 0 014
 Diptera adults 00004 2 240
 Diptera pupae 07101 3 875
      Ceratopogonidae 15 17 9 2 6 1 2 6 1
 Atrichopogon  spp. 00000 1 000
 Bezzia/Palpomyia  gp. 00012 6 2423
      Chaoboridae 00000 1 000
      Chironomidae 30000 1 000
 Ablabesmyia  spp. 00001 2 907
 Chaetocladius  spp. 80000 0 000
 Chironomini 00000 0 3100
 Cladotanytarsus  spp. 10000 0 000
 Corynoneura  spp. 41210 0 0 6 42921
 Cricotopus spp. 00300 0 000
 Cricotopus/Orthocladius 00002 0 000
 Demicryptochironomus  cuneatus 14 22 4 0 0 0 0 0 0
 Dicrotendipes  spp. 00000 0 200
 Diplocladius  cultriger 10001 0 000
 Harnischia spp. 00000 0 200
 Hydrobaenus  pilipes  gp. 00010 0 000
 Krenosmittia spp. 0 0 17 0 0 15 19 7 24
 Larsia  spp. 07020 5 29446
 Micropsectra  spp. 00000 0 004
 nocladius  spp. 00000 0 200
 tarsia  spp. 02010 1 020
 Nilotanypus spp. 0 34 0 0 0 5 13 6 25
 Orthocladiie 10200 0 000
 
 
 
  
 
92
Appendix II. Table 2.  Continued? 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Parachaetocladius spp. 12428 0 1 2 0 0 0
 Paracricotopus spp. 00000 1 000
 Parakiefferiella  spp. 00000 0 0210
 Parakiefferiella  triquetra 00000 0 200
 Paralauterborniella nigrohalteralis 00000 0 320
 Parameri spp. 00101 3 000
 Parametriocnemus spp. 17 10 0 6 2 14 26 22 20
 Parasmittia spp. 10000 0 000
 Paratendipes  spp. 00000 2 200
 Polypedilum  (P.) convictum 7 10 13 1 0 2 0 0 12
 Polypedilum  (P.) fallax 40912 0 000
 Polypedilum  (P.) illinoense 10400 0 2133
 Polypedilum  (T.) halterale 00030 1 023
 Polypedilum  (T.) scalaenum  gp. 5 0 12 3 1 3 10 6 4
 Polypedilum  spp. 10000 0 000
 Rheocricotopus  spp. 00000 0 030
 Rheotanytarsus  spp. 18 36 13 0 0 4 2 8 29
 Robackia  demeijerei 5 10 4 0 1 0 0 2 0
 Stempelli  spp. 10200 0 000
 Stempellinella spp. 00020 6 13410
 Tanypodie 10204 7 500
 Tanytarsus  spp. 90 557 59 1 0 22 43 20 41
 Thienemannia  spp. 10001 0 220
 Thienemanniella spp. 00200 0 033
 Thienemannimyia complex 0 0 13 1 0 16 12 5 26
 Tribelos spp. 10000 0 000
 Tvetenia  bavarica  gp. 10013 2 34716
 Zavrelimyia  spp. 00400 1 601
      Culicidae   
 Culex  spp. 20000 3 330
      Empididae   
 Hemerodromia spp. 30302 1 025
      Simuliidae   
 Simulium  spp. 01001 1 074
      Tanyderidae   
 Protoplasa  fitchii 10001 0 000
      Tipulidae 06000 0 000
 Hexatoma  spp. 21412 2 2 3 4 3 4
 Pseudolimnophila spp. 98131 11 011
 Tipula  spp. 00001 0 001
Ephemeroptera 10000 0 000
      Baetidae   
 Baetis  spp. 3 24 7 9 6 31 20 47 123
 
93
Appendix II. Table 2.  Continued? 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Ephemeroptera   
      Caenidae   
 Caenis  spp. 2 11 1 27 2 3 11 0 0
      Ephemeridae   
 Hexagenia  spp. 00000 0 001
      Heptageniidae   
 Stenonema spp. 00300 0 000
      Leptophlebiidae   
 Habrophlebiodes  spp. 19400 6 1934
Hemiptera 00010 0 000
Lepidoptera 00001 0 000
Megaloptera   
      Corydalidae   
 Nigronia spp. 02010 0 000
Odonata   
      Aeshnidae   
 Boyeria  spp. 02000 0 000
      Calopterygidae   
 Calopteryx  spp. 00010 0 020
      Gomphidae 04100 0 113
Plecoptera 139413 1 013
      Nemouridae   
 Amphinemura spp. 00000 0 010
      Perlidae   
 Eccoptura  xanthenes 12030 0 000
Trichoptera 0 3 4 25 17 8 17 10 53
      Calamoceratidae   
 Anisocentropus  pyraloides 10100 0 000
      Glossosomatidae   
 Glossosoma spp. 24012 0 000
      Hydropsychidae 44010 0 3100
 Cheumatopsyche  spp. 00000 0 001
 Diplectrona  spp. 11 40 3 7 2 5 12 24 22
 Hydropsyche  spp. 00000 0 020
 Macrostemum  carolina 00000 0 010
 Macrostemum  spp. 00001 0 000
      Limnephilidae   
 Neophylax spp. 00000 0 010
      Molannidae 02000 0 000
      Philopotamidae   
 Chimarra  spp. 04000 0 000
OTHER AQUATIC INVERTEBRATES   
Amphipoda   
      Gammaridae 02020 0 000
 
94
Appendix II. Table 2.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Copepoda 001001 0 10
      Calanoida 000300 12 12
      Harpacticoida 500012 3 10
Decapoda   
      Cambaridae 001000 0 00
Gastropoda 0 0 5 8 20 0 1 0 0
      Pleuroceridae   
 Elimia spp. 0101000 1 00
Hydracarina 2 127 20 3 9 11 19 2 8
Nematoda 714 4 315 0 5 311
Oligochaeta 63 17 13 38 41 27 20 15 19
      Lumbriculidae 600000 0 10
Ostracoda 000100 5 00
Pelecypoda 200000 0 00
Turbellaria   
      Planariidae 14 18 6 0 0 3 0 0 0
    
Total number of taxa per station 
52 42 45 42 42 49 48 51 44
Total number of organisms per station 381 1231 368 184 188 262 450 368 607
Mean number of organisms per m? 27271 88111 26340 13170 13456 18753 32210 26340 43447
 
95
Appendix II. Table 3.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 14 September 1994. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 0 0000 1 00
Coleoptera 0 0010 0 00
      Elmidae 0 16766 0 04
 Optioservus  spp.  0 0060 0 00
 Oulimnius  latiusculus   0 0002 3 01
 Stenelmis spp. adults 1 2010 1 04
      Psephenidae   
 Psephenus  herricki   15 42 1 2 116 36 3 27
      Ptilodactylidae   
 Anchytarsus  bicolor   594210 2 00
Collembola 1 0100 0 00
      Isotomidae 0 0002 10 00
      Sminthuridae 0 0010 0 00
Diptera 6 3020 0 14
 Diptera adults 0 3200 4 14
 Diptera pupae 2 7003 2 67
      Ceratopogonidae 3 6000 4 11
 Bezzia/Palpomyia  gp. 48 38 0 2 32 32 2 17
      Chironomidae 0 0000 2 14
 Chironomie 0 0060 0 00
 Chironomini 0 0100 1 00
 Corynoneura  spp. 78 122 0 0 18 4 18 7
 Cricotopus/Orthocladius 0 0010 0 00
 Demicryptochironomus  cuneatus 13 12 0 0 0 0 0 0
 Diplocladius  cultriger 0 12000 2 00
 Krenopelopia  spp. 12 8 0 0 0 4 2 31
 Krenosmittia spp. 2 0 0 0 22 11 0 0
 Larsia  spp. 4 20 0 0 13 11 1 8
 Micropsectra  spp. 0 0000 2 00
 tarsia  spp. 2 0000 0 00
 Nilotanypus spp. 0 28 1 0 21 7 34 29
 Nilothauma spp. 2 0000 0 00
 Parachaetocladius spp. 0 24008 1 00
 Parakiefferiella  triquetra 0 0000 2 00
 Parametriocnemus spp. 2 0102 3 03
 Paratanytarsus  spp. 0 0003 0 00
 Polypedilum  (P.) convictum 12 59 2 0 25 1 0 0
 Polypedilum  (P.) fallax 2 0000 0 00
 Polypedilum  (P.) illinoense 0 0003 2 00
 Polypedilum  (T.) halterale 0 0510 0 03
 Polypedilum  (T.) scalaenum  gp. 2 20807 3 719
 Polypedilum  spp. 0 0000 0 03
 Psilometriocnemus spp. 2 0000 0 00
 
 
  
 
96
Appendix II. Table 3.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Rheotanytarsus  spp. 210 131 14 15 73 17 79 61
 Robackia  demeijerei 0 0 0 0 0 0 42 0
 Stempelli  spp. 5 0000 1 00
 Stempellinella spp. 0 0 0 0 0 25 10 33
 Sublettea coffmanni 0 0000 0 04
 Tanypodie 0 0000 1 00
 Tanypodie pupae 2 0000 0 00
 Tanytarsus  spp. 418 718 1 3 20 21 23 59
 Thienemanniella spp. 14 0 1 1 7 0 9 3
 Thienemannimyia complex 0 32002 1 40
 Tvetenia  bavarica  gp. 0 0 21 15 3 0 0 0
      Culicidae 0 0001 1 01
      Empididae   
 Hemerodromia spp. 0 1022 0 00
      Psychodidae   
 Pericoma/Telmatoscopus 1 0000 0 00
      Simuliidae 0 0020 0 00
 Simulium  spp. 3 4030 0 24
      Tipulidae 0 5000 0 00
 Hexatoma  spp. 10 15 1 0 6 16 3 21
Ephemeroptera   
      Baetidae   
 Baetis  spp. 42 35 16 23 90 34 51 107
      Caenidae   
 Caenis  spp. 6 01470 4 00
      Ephemeridae   
 Hexagenia  spp. 0 0008 2 00
      Heptageniidae   
 Stenonema spp. 0 7000 0 04
      Leptophlebiidae   
 Habrophlebiodes  spp. 2 12 0 0 51 42 7 24
Odonata   
      Aeshnidae   
 Boyeria  spp. 1 0000 0 00
      Cordulegastridae   
 Cordulegaster  spp. 0 0000 0 10
      Gomphidae 0 2010 0 10
 Erpetogomphus  desigtus 0 0100 0 00
      Odonata 0 0000 1 00
Plecoptera 12 19 3 40 10 4 1 25
 
 
97
Appendix II. Table 3.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Trichoptera 2 0 13 7 16 6 5 12
      Glossosomatidae   
 Glossosoma spp. 0 0010 0 01
      Hydropsychidae   
 Cheumatopsyche  spp. 0 0040 32 01
 Diplectrona  spp. 0 6 26 16 11 12 4 6
 Hydropsyche  spp. 0 0010 0 09
      Molannidae   
 Molanna  spp. 0 0100 0 00
      Philopotamidae   
 Chimarra  spp. 0 6200 0 00
OTHER AQUATIC INVERTEBRATES   
Amphipoda   
      Gammaridae 1 0000 0 00
Copepoda 1 0102 21 00
Gastropoda 0 8102 10 00
      Pleuroceridae   
 Elimia spp. 0 0802 3 00
Hydracarina 184 200 0 12 40 55 14 79
Nematoda 34 11 6 41 2 7 8 18
Oligochaeta 0 5 815 2 0 012
Ostracoda 1 0000 2 00
Pelecypoda 0 0100 0 00
Turbellaria   
      Planariidae 1 3000 0 00
    
Total number of taxa per station 
40 37 30 31 36 48 29 38
Total number of organisms per station 1164 1651 172 259 633 469 341 660
Mean number of organisms per m? 83315 118173 12311 18538 45308 33570 24408 47241
 
98
Appendix II. Table 4.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 20 February 1995. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Coleoptera   
      Elmidae 812000 0 00
 Dubiraphia  spp.  000000 0 11
 Optioservus  spp.  0 0 3 0 23 0 1 0 0
 Oulimnius  latiusculus   002114 2 02
 Oulimnius  latiusculus  adults 200000 0 00
 Stenelmis spp. adults 000100 0 00
      Psephenidae   
 Psephenus  herricki   1 0 34 2 0 19 20 0 4
      Ptilodactylidae   
 Anchytarsus  bicolor   100010 0 00
Diptera 100002 2 00
 Diptera adults 002000 1 10
 Diptera pupae 214100 0 00
      Ceratopogonidae 0 7 0 0 4 6 1 44 1
 Bezzia/Palpomyia  gp. 92422 3 390 16 121
      Chironomidae 100000 7 00
 Cladotanytarsus  spp. 050000 0 00
 Corynoneura  spp. 8 4 61 0 0 1 3 0 2
 Cricotopus spp. 005000 0 00
 Cricotopus/Orthocladius 202110 2 00
 Cryptochironomus spp. 000100 0 00
 Diplocladius  cultriger 100000 0 00
 Eukiefferiella  claripennis  gp. 000010 0 00
 Harnischia spp. 050000 0 00
 Krenosmittia spp. 050003 0 00
 Larsia  spp. 000000 0 12
 Micropsectra  spp. 001000 0 00
 Monopelopia spp. 000003 4 010
 nocladius  spp. 000000 2 00
 Nilotanypus spp. 300101 21 513
 Nimbocera pinderi 050000 0 00
 Orthocladiie 000010 0 00
 Parachaetocladius spp. 101011 0 00
 Parachironomus spp. 100000 0 00
 Parakiefferiella  triquetra 100000 0 00
 Parameri spp. 100000 0 00
 Parametriocnemus spp. 3 2 24 2 0 4 28 7 12
 Polypedilum  (P.) convictum 3 4 15 0 1 1 7 0 0
 Polypedilum  (P.) illinoense 005000 0 10
 Polypedilum  (T.) halterale 0 0 13 0 3 0 0 17 1
 Polypedilum  (T.) scalaenum  gp. 252021 8 96
 Rheotanytarsus  spp. 21 9 6 0 3 1 23 23 47
 
 
  
 
99
Appendix II. Table 4.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Robackia  demeijerei 010000 0 10
 Stempelli  spp. 0540000 0 00
 Stempellinella spp. 2 0 0 0 0 10 11 0 5
 Tanypodie 000000 0 01
 Tanytarsus  spp. 66 89 378 2 1 4 10 6 4
 Thienemanniella spp. 7 5 10 8 6 1 6 2 1
 Thienemannimyia complex 000000 4 04
 Tvetenia  bavarica  gp. 9 1 25 10 8 1 11 6 6
      Culicidae 000000 0 10
      Empididae   
 Hemerodromia spp. 002080 0 00
      Simuliidae   
 Simulium  spp. 41 7 18 2 3 0 1 2 3
      Tipulidae 110000 0 00
 Hexatoma  spp. 2188015 0 32
 Pseudolimnophila spp. 110000 1 02
 Tipula  spp. 000000 0 11
Ephemeroptera   
      Baetidae   
 Baetis  spp. 12221830 0 48
      Caenidae   
 Caenis  spp. 000131 5 00
      Ephemeridae   
 Hexagenia  spp. 010000 0 00
      Leptophlebiidae   
 Habrophlebia  vibrans 000002 0 00
 Habrophlebiodes  spp. 007100 24 11
Hemiptera 010000 0 00
Odonata   
      Cordulegastridae   
 Cordulegaster  spp. 000000 0 01
Plecoptera 200002 3 927
      Chloroperlidae   
 Alloperla spp. 19 13 32 2 0 7 9 2 4
      Nemouridae   
 Amphinemura spp. 610000 0 61
      Perlidae   
 Eccoptura  xanthenes 000010 0 00
 
 
 
100
Appendix II. Table 4.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Trichoptera 002520 0 00
      Glossosomatidae   
 Glossosoma spp. 3 0 2 9 20 0 0 0 0
      Hydropsychidae 002000 1 03
 Cheumatopsyche  spp. 000001 6 02
 Diplectrona  spp. 002220 0 00
 Hydropsyche  spp. 000110 0 01
Copepoda 510000 0 00
Gastropoda 001001 10 10
      Pleuroceridae   
 Elimia spp. 000202 4 00
Hydracarina 002110 5 02
Nematoda 433425 1 20
Oligochaeta 51 13 21 10 15 73 10 2 84
Pelecypoda   00
      Corbiculidae   
 Corbicula fluminea 010000 0 
Turbellaria   00
      Planariidae 314000 0 
   
Total number of taxa per station 
37 32 36 25 29 28 35 27 34
Total number of organisms per station 306 291 725 91 122 262 270 159 285
Mean number of organisms per m? 21903 20829 51893 6513 8732 18753 19326 11381 20399
 
101
Appendix II. Table 5.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 13 June 1995. 
AQUATIC INSECTS 1 C 2A 2B 3A 3B 4A 4B 5A 5B
Coleoptera   
      Dryopidae   
 Helichus  spp. adults 01000 0 00
      Elmidae 10 4 0 4 37 2 27 10
 Ancyronyx  varigatus   00002 0 00
 Elmidae adults 00000 0 10
 Optioservus  spp.  01020 0 00
 Optioservus  spp. adults 10010 0 00
      Psephenidae   
 Psephenus  herricki   21 29 0 0 151 17 28 0
 Psephenus  herricki  adults 10000 0 00
      Ptilodactylidae   
 Anchytarsus  bicolor   10100 0 11
Collembola 00000 0 10
      Isotomidae 00001 0 00
      Sminthuridae 00000 0 10
Diptera 30000 2 720
 Diptera adults 01201 1 21
 Diptera pupae 9 9 1 0 10 2 5 4
      Ceratopogonidae 21 55 0 2 18 49 26 13
 Bezzia/Palpomyia  gp. 60500 0 015
      Chironomidae 07010 0 03
 Ablabesmyia  spp. 00009 0 00
 Chironomini 04215 0 30
 Corynoneura  spp. 95 68 0 0 0 11 6 32
 Cricotopus bicinctus 0 0 0 0 10 0 0 0
 Cricotopus/Orthocladius 80000 0 00
 Cryptochironomus spp. 00200 0 00
 Demicryptochironomus  cuneatus 18 38 0 0 0 0 0 0
 Djalmabatista  spp. 00010 0 00
 Glyptotendipes spp. 00000 0 30
 Heterotrissocladius  macidus  gp. 90000 0 00
 Krenosmittia spp. 00000 4 1 7
 Larsia  spp. 37 24 0 1 37 10 12 30
 Micropsectra  spp. 00010 0 00
 Monopelopia spp. 04000 0 00
 Nilotanypus spp. 16 8 0 0 0 6 42 28
 Parachaetocladius spp. 070310 3 00
 Parameri spp. 00000 0 30
 Parametriocnemus spp. 28 0 1 1 19 0 5 7
 Paratendipes  spp. 0 0 0 0 10 0 3 0
 Polypedilum  (P.) convictum 50 0 0 0 38 2 4 3
 Polypedilum  (P.) illinoense 00010 0 00
 
 
  
 
102
Appendix II. Table 5.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Polypedilum  (T.) halterale 417110 0 00
 Polypedilum  (T.) scalaenum  gp. 017115 1 81
 Potthastia  longima  gp. 00010 0 00
 Pseudorthocladius spp. 08000 0 00
 Rheocricotopus  spp. 80000 0 00
 Rheotanytarsus  spp. 89 77 0 2 19 4 29 40
 Saetheria spp. 00009 0 00
 Stempellinella spp. 04000 0 04
 Tanypodie 80000 2 03
 Tanytarsus  spp. 198 175 11 2 426 25 82 47
 Thienemanniella spp. 25 0 3 0 110 0 0 3
 Thienemannimyia complex 29 7 0 0 15 10 27 16
 Tvetenia  bavarica  gp. 00030 0 00
 Zavrelimyia  spp. 00000 0 03
      Culicidae   
 Culex  spp. 00107 0 02
      Empididae   
 Hemerodromia spp. 00008 0 12
      Simuliidae   
 Simulium  spp. 18 0 2 13 0 0 0 2
      Tipulidae 15 14 2 0 19 12 14 5
 Pseudolimnophila spp. 17 3 0 0 0 0 1 7
Ephemeroptera   
      Baetidae 13 0 0 0 0 0 0 0
 Baetis  spp. 118 20 8 8 123 22 40 31
      Caenidae   
 Caenis  spp. 21 23 1 10 3 3 0 0
      Heptageniidae   
 Stenonema spp. 21002 1 01
      Leptophlebiidae   
 Habrophlebiodes  spp. 5 18 0 0 35 49 26 23
Odonata   
      Calopterygidae   
 Calopteryx  spp. 01000 1 00
      Gomphidae 01001 0 00
 Erpetogomphus  desigtus 11000 0 00
Plecoptera 610100 3 10
      Nemouridae   
 Amphinemura spp. 40000 0 00
      Perlidae   
 Acroneuria  spp. 10000 0 00
 
 
 
103
Appendix II. Table 5.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Trichoptera 00196 1 22
      Glossosomatidae   
 Glossosoma spp. 00860 0 11
      Hydropsychidae 0 2 0 13 10 0 0 0
 Cheumatopsyche  spp. 310000 0 00
 Diplectrona  spp. 21 10 6 11 34 12 10 37
 Hydropsyche  spp. 40000 0 02
 Macrostemum  carolina 30100 0 00
      Limnephilidae 00000 0 01
      Philopotamidae   
 Chimarra  spp. 10000 0 00
OTHER AQUATIC INVERTEBRATES 948 742 64 97 1180 255 433 407
Amphipoda   
      Gammaridae 20010 0 00
Copepoda 0 2 0 0 40 4 4 0
Decapoda 00002 0 00
Gastropoda 6 8 0 5 28 4 0 0
      Pleuroceridae   
 Elimia spp. 007170 8 00
Hydracarina 135 138 0 3 34 30 7 27
Nematoda 05402 0 14
Oligochaeta 10 15 22 6 14 8 8 3
Ostracoda 1 0 0 0 16 0 3 0
   
Total number of taxa per station 
46 40 25 30 38 31 38 38
Total number of organisms per station 1102 910 97 129 1316 309 456 441
Mean number of organisms per m? 78878 65135 6946 9233 94195 22117 32639 31565
 
104
Appendix II. Table 6.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 16 August 1995. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 00002 0 00
Coleoptera 00006 0 00
      Elmidae 91192 7 14
 Ancyronyx  varigatus   00000 0 03
 Macronychus  glabratus   00000 0 10
 Optioservus  spp.  00050 0 00
 Stenelmis spp. adults 02000 0 00
      Psephenidae   
 Psephenus  herricki   131 34 0 3 84 1 2 3
      Ptilodactylidae   
 Anchytarsus  bicolor   0 23 2 12 0 1 0 0
Collembola 04000 0 00
      Isotomidae 00100 0 00
      Sminthuridae 20003 0 00
Diptera 00100 2 42
 Diptera adults 30001 0 00
 Diptera pupae 18 7 2 1 4 7 4 5
      Ceratopogonidae 32776 8 95
 Bezzia/Palpomyia  gp. 36 60 8 1 10 32 6 6
      Chironomidae 02000 1 24
 Ablabesmyia  spp. 00013 0 00
 Chironomini 54000 0 00
 Cladotanytarsus  spp. 0 0 0 0 24 0 0 0
 Corynoneura  spp. 118000 3 35
 Cricotopus bicinctus 6 0 0 0 14 0 0 0
 Cricotopus spp. 50000 0 00
 Cryptochironomus spp. 50000 0 00
 Cryptotendipes  spp. 00003 0 00
 Demicryptochironomus  cuneatus 64 8 0 0 0 0 0 0
 Dicrotendipes  spp. 0 0 0 0 27 0 0 0
 Djalmabatista  spp. 00003 0 00
 Krenopelopia  spp. 10000 0 23
 Krenosmittia spp. 12 2 0 0 0 32 4 13
 Larsia  spp. 29 8 0 0 35 0 1 0
 Microtendipes  pedellus  gp. 00004 0 00
 Monopelopia spp. 8 25 0 1 10 8 6 3
 nocladius  spp. 50000 0 00
 tarsia  spp. 12 5 0 0 0 0 0 0
 Nilotanypus spp. 613010 4 87
 Orthocladiie 00000 0 10
 Parachaetocladius spp. 17 0 0 0 0 1 0 0
 Paralauterborniella nigrohalteralis 00100 0 00
 Parametriocnemus spp. 00000 1 00
 
 
  
 
105
Appendix II. Table 6.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Diptera   
      Chironomidae   
 Polypedilum  (P.) convictum 14 5 0 0 55 8 1 0
 Polypedilum  (P.) illinoense 00003 0 00
 Polypedilum  (T.) halterale 12 9 11 0 3 1 0 0
 Polypedilum  (T.) scalaenum  gp. 0 0 7 0 11 7 1 1
 Procladius spp. 00003 0 00
 Rheotanytarsus  spp. 222210 1 26
 Robackia  demeijerei 00010 0 00
 Stempellinella spp. 10003 0 11
 Tanytarsini 00000 0 10
 Tanytarsus  spp. 37 186 2 0 80 5 2 3
 Thienemanniella spp. 276 10 2 1 28 1 0 0
 Thienemannimyia complex 25 6 0 0 0 1 2 0
 Tvetenia  bavarica  gp. 00220 0 00
      Culicidae 00002 0 10
      Dixidae   
 Dixa  spp. 00010 0 00
      Empididae   
 Hemerodromia spp. 00110 0 00
      Simuliidae   
 Simulium  spp. 00130 0 00
      Tipulidae   
 Hexatoma  spp. 20 17 1 0 12 7 8 4
 Pseudolimnophila spp. 00000 0 14
 Tipula  spp. 05000 0 00
Ephemeroptera 10000 0 00
      Baetidae 00000 0 01
 Baetis  spp. 97 120 16 13 43 6 65 41
      Caenidae   
 Caenis  spp. 1 4 5 2 55 1 1 0
      Heptageniidae   
 Stenonema spp. 16000 0 00
      Leptophlebiidae   
 Habrophlebiodes  spp. 13 46 0 0 0 23 21 11
Hemiptera   
      Mesoveliidae   
 Mesovelia  spp. 00000 1 00
      Veliidae 04000 0 00
Lepidoptera 00000 0 10
Odonata   
      Gomphidae 00000 0 10
 Erpetogomphus  desigtus 03000 0 00
 Stylogomphus  albistylus 01000 0 00
 
 
 
106
Appendix II. Table 6.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Plecoptera 40028 0 00
      Perlidae 00000 0 01
 Eccoptura xanthenes 00000 1 00
Trichoptera 09020 0 10
      Glossosomatidae   
 Glossosoma spp. 00130 0 00
      Hydropsychidae 30000 1 00
 Cheumatopsyche  spp. 00000 3 03
 Diplectrona  spp. 191148 8 45
 Hydropsyche  spp. 01004 0 00
 Macrostemum  carolina 00000 0 01
      Hydroptilidae 10002 0 00
OTHER AQUATIC INVERTEBRATES   
Copepoda 0 0 1 2 38 0 0 0
      Calanoida 40008 0 02
Gastropoda 0 10 0 0 10 1 0 0
      Pleuroceridae   
 Elimia spp. 00900 2 00
Hydracarina 136 346 0 5 18 16 5 25
Nematoda 12 18 4 0 0 0 2 1
Oligochaeta 108 0 22 11 14 48 6 22
Ostracoda 4 0 2 0 25 0 1 1
Turbellaria 134000 0 00
    
Total number of taxa per station 
44 39 26 26 40 34 35 31
Total number of organisms per station 1152 1089 113 105 674 250 181 196
Mean number of organisms per m? 82457 77947 8088 7516 48243 17894 12955 14029
 
107
Appendix II. Table 7.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 12 October 1995. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 200001 0 10
Coleoptera   
      Elmidae 436023 1 12
 Ancyronyx  varigatus   000062 0 00
 Macronychus  glabratus   000000 0 10
 Oulimnius  latiusculus   000003 3 00
 Promoresia  spp.  000000 5 00
      Psephenidae   
 Psephenus  herricki   01119 1 0 7 12 2 0
      Ptilodactylidae   
 Anchytarsus  bicolor   002030 0 00
Diptera 300010 1 12
 Diptera adults 000000 0 01
 Diptera pupae 300030 1 01
      Ceratopogonidae 17443205 0 78
 Bezzia/Palpomyia  gp. 15 2 16 1 0 17 16 11 4
      Chironomidae 010001 0 01
 Chironomini 000000 1 11
 Cladotanytarsus  spp. 020003 0 00
 Corynoneura  spp. 722141 2 05
 Cricotopus bicinctus 0110004 0 00
 Demicryptochironomus  cuneatus 014000 0 00
 Diplocladius  cultriger 000000 1 00
 Krenopelopia  spp. 000000 1 00
 Krenosmittia spp. 5 0 1 0 0 1 10 10 7
 Labrundinia spp. 000000 0 01
 Larsia  spp. 003000 1 00
 Monopelopia spp. 045001 3 01
 Nilotanypus spp. 302010 2 13
 Orthocladiie 010000 0 00
 Parachaetocladius spp. 001000 0 00
 Parameri spp. 000000 0 50
 Parametriocnemus spp. 000000 1 00
 Paratendipes  spp. 000001 1 10
 Polypedilum  (P.) convictum 0 23 11 0 0 10 18 0 0
 Polypedilum  (T.) halterale 000721 2 10
 Polypedilum  (T.) scalaenum  gp. 000200 0 20
 Rheotanytarsus  spp. 1 43231 2 69
 Robackia  demeijerei 000000 0 01
 Tanytarsus  spp. 1 2 1 0 0 15 42 4 3
 Thienemanniella spp. 040973 0 00
 Thienemannimyia complex 000000 0 10
 Tvetenia  bavarica  gp. 000240 0 00
 
 
  
 
108
Appendix II. Table 7.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
      Simuliidae 000010 0 00
 Simulium  spp. 010000 0 00
      Tanyderidae 000100 0 00
      Tipulidae 010000 4 00
 Hexatoma  spp. 356120 4 83
 Pseudolimnophila spp. 000000 0 10
Ephemeroptera 011001 0 00
      Baetidae   
 Baetis  spp. 11 31 75 5 21 4 12 24 16
      Caenidae   
 Caenis  spp. 1 2 1 1 16 83 9 0 0
      Heptageniidae   
 Stenonema spp. 001000 0 00
      Leptophlebiidae   
 Habrophlebiodes  spp. 1 6 4 0 0 3 4 10 1
Hemiptera   
      Veliidae   
 Rhagovelia spp. 000010 0 00
Lepidoptera 100000 0 00
Odonata   
      Gomphidae   
 Erpetogomphus  desigtus 010000 0 00
 Stylogomphus  albistylus 100000 0 00
Plecoptera 1331000 3 03
      Perlidae   
 Eccoptura  xanthenes 001000 0 00
Trichoptera 000105 0 20
      Hydropsychidae 001403 25 00
 Cheumatopsyche  spp. 020001 0 00
 Diplectrona  spp. 067250 0 00
 Hydropsyche  spp. 044130 3 00
 Macrostemum  spp. 000030 1 00
OTHER AQUATIC INVERTEBRATES   
Copepoda 000010 0 10
      Harpacticoida 000000 0 10
Gastropoda 051002 0 01
      Pleuroceridae   
 Elimia spp. 000119 3 01
 
 
109
Appendix II. Table 7.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Hydracarina 25 52 74 0 4 10 19 1 4
Nematoda 854311 4 41
Oligochaeta 10 23 11 20 11 36 30 3 10
Pelecypoda   
      Corbiculidae   
 Corbicula fluminea 001000 0 10
Turbellaria   
      Planariidae 21723 0 0 1 0 0 0
   
Total number of taxa per station 
22 32 32 20 25 32 34 28 25
Total number of organisms per station 147 240 296 68 126 239 247 112 90
Mean number of organisms per m? 10522 17178 21187 4867 9019 17107 17679 8017 6442
 
110
Appendix II. Table 8.   Total fauna, number of taxa, and mean number of organisms/m? 
collected at each station in the Camden Study on 11 December 1995. 
AQUATIC INSECTS 1C 2A 2B 3A 3B 4A 4B 5A 5B
Insecta 020000 0 00
Coleoptera   
      Elmidae 000004 0 00
 Ancyronyx  varigatus   000100 0 00
 Dubiraphia  spp.  000001 0 00
 Optioservus  spp.  012210 0 00
 Oulimnius  latiusculus   231305 0 00
 Stenelmis spp. adults 010010 0 00
      Psephenidae   
 Psephenus  herricki   065305 0 00
      Ptilodactylidae   
 Anchytarsus  bicolor   001500 0 00
Collembola 1400000 0 00
Diptera 220015 2 10
 Diptera adults 000100 0 10
 Diptera pupae 010110 0 00
      Ceratopogonidae 100540 1 92
 Bezzia/Palpomyia  gp. 32015 5 043 0 7 5
      Chironomidae   
 Chironomini 000000 0 10
 Cladotanytarsus  spp. 000003 0 00
 Corynoneura  spp. 1491560 1 23
 Cricotopus bicinctus 041011 0 00
 Cricotopus  tremulus  gp. 020000 0 00
 Cryptochironomus spp. 021000 0 10
 Demicryptochironomus  cuneatus 0233000 0 00
 Eukiefferiella  claripennis  gp. 100101 0 10
 Krenosmittia spp. 000405 1 21
 Larsia  spp. 072103 0 26
 Micropsectra  spp. 000002 0 00
 Orthocladiie 010001 0 00
 Parametriocnemus spp. 100001 0 10
 Paratendipes  spp. 001000 1 01
 Polypedilum  (P.) convictum 3314201 1 12
 Polypedilum  (T.) halterale 0 0 1 14 0 0 0 25 0
 Polypedilum  (T.) scalaenum  gp. 0 1 0 12 0 51 5 11 10
 Polypedilum  spp. 000000 0 01
 Pseudorthocladius spp. 000010 0 00
 Rheocricotopus  spp. 020000 0 10
 Rheotanytarsus  spp. 0125013 0 24
 Tanypodie 002000 0 00
 Tanytarsus  spp. 271206 2 02
 Thienemanniella spp. 042953 3 13
 
 
  
 
111
Appendix II. Table 8.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
        Chironomidae   
 Thienemannimyia complex 120000 0 00
 Tvetenia  bavarica  gp. 220977 0 10
       Empididae 
 Hemerodromia spp. 01011
 
0 
 
0 0 0
      Simuliidae   
 Simulium spp. 15 18 10 1 18 30 1 0 1
      Tipulidae 002000 0 00
 Hexatoma spp. 013105 1 31
 Pseudolimnophila spp. 000001 0 00
 Tipula spp. 200010 0 00
Ephemeroptera 001000 0 00
      Baetidae 000000 1 00
 Baetis spp. 26424211012 4 2012
      Caenidae   
 Caenis spp. 0000018 3 00
      Leptophlebiidae   
 Habrophlebiodes spp. 050100 0 02
Odonata   
      Gomphidae 000100 0 00
 Progomphus spp. 000002 0 00
Plecoptera 21 9 6 3 0 19 0 2 10
      Chloroperlidae   
 Alloperla spp. 2198000 0 00
      Leuctridae 100000 0 00
      Nemouridae 000001 0 00
 Amphinemura spp. 300000 0 00
Trichoptera 031302 0 01
      Glossosomatidae   
 Glossosoma spp. 000010 0 00
      Hydropsychidae 000000 0 22
 Cheumatopsyche spp. 070003 0 10
 Diplectrona spp. 600000 0 00
 Hydropsyche spp. 0287472 0 00
      Hydroptilidae   
 Hydroptila spp. 000003 0 00
      Philopotamidae   
 Chimarra spp. 020000 0 00
 
 
112
Appendix II. Table 8.  Continued? 
OTHER AQUATIC INVERTEBRATES 1C 2A 2B 3A 3B 4A 4B 5A 5B
Copepoda   
      Calanoida 000002 0 00
Gastropoda 013000 0 00
      Pleuroceridae   
 Elimia spp. 000000 1 00
Hydracarina 14 58 28 2 0 14 1 0 1
Nematoda 18 22 8 11 2 7 1 0 2
Oligochaeta 48 48 36 34 16 78 8 20 9
Turbellaria   
      Planariidae 071001 3 10
    
Total number of taxa per station 
23 39 31 32 18 37 19 25 22
Total number of organisms per station 197 417 183 173 84 391 41 119 81
Mean number of organisms per m? 14101 29848 13099 12383 6012 27987 2935 8518 5798