VERTEBRATE RESPONSE TO A TIDAL
MARSH RESTORATION IN
HUMBOLDT BAY, CALIFORNIA
by
Sandra L. Jacobson
A Thesis
Presented to
The Faculty of Humboldt State University
In Partial Fulfillment
of the Requirements for the Degree
Master of Science
December, 1986 VERTEBRATE RESPONSE TO A TIDAL
MARSH RESTORATION IN
HUMBOLDT BAY, CALIFORNIA
by
Sandra L. Jacobson
Approved by the Master's Thesis Committee
Paul F. Springer
Stanley W. Harris
Director, Natural Resources Graduate Program Date
86/W-59/05/22 Natural Resources Graduate Program Number
Approved by the Dean of Graduate Studies
Alba M. Gillespie
ABSTRACT
A 5.5-ha abandoned log pond (originally salt marsh) at the end of Park Street, Eureka, California was chosen as the off-site mitigation area for the destruction of 6.8 ha of wildlife habitat during the construction of the Woodley Island marina. Passive saltmarsh restoration was attempted by breaching a dike separating the log pond from Freshwater Slough (an estuary of Humboldt Bay) in December 1980, thus allowing the periodic tidal intrusion of salt water into 3.8 ha of the area. This study measured the response of amphibians, reptiles, birds, and mammals to the change in vegetation and other habitat conditions brought about by the reestablishment of tidal action.
Previously, in August 1979 an interior dike had been constructed within a portion of the mitigation area to enhance an existing 0.7-ha freshwater marsh, and the effect of this change on the vertebrate life also was appraised.
Frogs disappeared and snakes declined in numbers within the tidal portion of the mitigation area after the dike was breached.
Savannah Sparrows (Passerculus sandwichensis) were the most common birds
in the grassland of the dried log pond before breaching, and declined after breaching. Several species of shorebirds and the Snowy Egret
(Egretta thula) increased their use of the tidal portion of the mitigation area after breaching. Small mammals such as California Vole
(Microtus californicus), Vagrant Shrew (Sorex vagrans), and Western
Harvest Mouse (Reithrodontomys megalotis) were common before breaching,
iii iv but declined in numbers in the first month after breaching. Use of the area by the most common large mammal, the Domestic Dog (Canis familiaris), increased after breaching because people used the flooded marsh to train retrievers.
Within the freshwater marsh five species of birds showed significant increases in numbers during subsequent corresponding seasons, whereas one species showed a significant decrease in numbers.
The success of the restoration after breaching was not fully known at the end of 1.5 years of study, but the trend was towards a renewed, vigorous salt marsh. In addition, the interior dike construction enhanced the freshwater marsh. In the interim the tidal and freshwater marshes provided valuable foraging areas for many species of wetland birds, and the freshwater marsh also served as a nesting area for certain bird species. ACKNOWLEDGEMENTS
I wish to acknowledge the great deal of help provided to me throughout this project.
Financial support was provided by the U. S. Fish and Wildlife
Service through a contract with the Humboldt State University
Foundation, which, in turn, administered the study through the Wildlife
Research Field Station.
Special thanks go to Dr. Paul Springer, whose advice and expert professional help went a long way toward making the project a success.
I especially acknowledge his editorial help; without it I would have missed an important step in my education.
My other committee members, Drs. Stan Harris and John Sawyer, also contributed their time and expertise for my benefit. I appreciate not only the professional help but also the moral support.
Dr. Dave Craigie helped me with the statistical analyses and interpretations. Pat Collins introduced me to the SPSS analysis and helped me to learn the seemingly thousands of details needed to successfully work with a computer.
Dana Base, John Sterling and John Kelly helped with the field
work on those days I was unable to work. Jan Kastler expertly typed the
thesis, and Laura Montagna prepared the figures.
The final person instrumental to the success of the project
contributed neither professional expertise nor field time. He did contribute love, encouragement, moral support, and gas for the many vi trips I took from my summer job sites to the study sites. He is my husband, Chris, and without his contributions I would have had a much more difficult, if not impossible, task in finishing the project. So to
Chris I address my deepest appreciation and thanks. TABLE OF CONTENTS
Page
ABSTRACT iii
ACKNOWLEDGEMENTS
LIST OF TABLES xi
LIST OF FIGURES xii
INTRODUCTION 1
STUDY AREAS 6
METHODS 13
Amphibians and Reptiles 13
Birds 13
Mammals 19
RESULTS 23
Amphibians and Reptiles 23
Birds 25
Saltmarsh Breeding Bird Counts 27
Saltmarsh Year-round Bird Counts 30
Snowy Egret 30
Black-shouldered Kite 32
Shorebirds 32
Vaux's Swift and Swallows 41
Marsh Wren 43
Emberizids 48
vii viii
TABLE OF CONTENTS (continued) Page
Western Meadowlark 53
American Goldfinch 55
Freshwater Marsh Breeding and Year-round Bird Counts . . 55
Mammals 61
Salt Marsh 61
Freshwater Marsh 64
DISCUSSION 66
Salt Marsh 66
Amphibians and Reptiles 66
Birds 66
Snowy Egret 67
Black-shouldered Kite 68
Shorebirds 68
Vaux's Swift and Swallows 69
Marsh Wren 70
Emberizids 70
Western Meadowlark 72
American Goldfinch 73
Mammals 73
Freshwater Marsh 75
Amphibians and Reptiles 75
Birds 75
Mammals 77
CONCLUSIONS 78
MANAGEMENT RECOMMENDATIONS 80
REFERENCES CITED 82 ix
TABLE OF CONTENTS (continued) Page
APPENDIXES
A. Common and Scientific Names of Amphibians and Reptiles Mentioned in the Text or Tables 88
B. Common and Scientific Names of Birds Mentioned in the Text, Tables or Appendixes 89
C. Common and Scientific Names of Mammals Mentioned in the Text or Tables 94
D. Common and Scientific Names of Plants Mentioned in the Text 95
E. Mean Number (and Standard Error) of Birds per Visit in Salt Marsh at the Mitigation Area, Eureka, California by Season 97
F. Mean Number (and Standard Error) of Birds per Visit in Salt Marsh at the Control Area, Eureka, California by Season 106
G. Mean Number (and Standard Error) of Birds per Visit in Freshwater Marsh at the Mitigation Area, Eureka, California by Season 112
H. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1979 at the Mitigation Area, Eureka, California 120
I. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1980 at the Mitigation Area, Eureka, California 121
J. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1981 at the Mitigation Area, Eureka, California 122
K. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1982 at the Mitigation Area, Eureka, California 123
L. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1980 at the Control Area, Eureka, California 124
M. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1981 at the Control Area, Eureka, California 125 x
TABLE OF CONTENTS (continued) Page
APPENDIXES (continued)
N. Approximate Territorial Boundaries of Birds during the Breeding Bird Counts in 1982 at the Control Area, Eureka, California 126 LIST OF TABLES
Table Page
1 Breeding Bird Count Dates at the Mitigation and Control Areas, Eureka, California 15
2 Number and Frequency of Occurrence of Amphibians and Reptiles Observed at the Mitigation Area, Eureka, California before and after Dike Breaching, 1979-1982 24
3 Species of Birds with Significant Differences in Numbers between at Least 2 Years at a Given Season in Salt Marsh at the Mitigation and/or Control Areas, Eureka, California, 1979-1982 . . . . 26
4 Numbers of Breeding Bird Territories during Breeding Bird Counts in Salt Marsh at the Mitigation and Control Areas, Eureka, California, 1979-1982 28
5 Species of Birds with Significant Differences in Numbers between at Least 2 Years at a Given Season in Freshwater Marsh at the Mitigation Area, Eureka, California, 1979-1982 57
6 Species of Birds with Significant Differences (P<0.05) in Numbers in Fall in Freshwater Marsh at the Mitigation Area, Eureka, California, 1980-1981. . 60
7 Number of Small Mammals Caught and the Calculated Density (No. animals/ha), Eureka, California, 1979-1981 62
8 Number and Frequency of Occurrence of Large Mammal Sign Observed at the Mitigation Area before and after Dike Breaching, Eureka, California, 1979-1982 63
9 Number and Frequency of Occurrence of Large Mammal Sign Observed in Salt Marsh at the Control Area before and after Dike Breaching, Eureka, California, 1980-1982 65
xi
LIST OF FIGURES
Figure Page
1 Location of Woodley Island Marina Site, Park Street Mitigation Area, and Bay Street Control Area on Humboldt Bay, Eureka, California 3
2 Habitat Types in 1979-80 before Dike Breaching and Bird Count Route in 1979 before Dike Construction at the Park Street Mitigation Area, Eureka, California 7
3 Habitat Types in 1981 after Dike Breaching and Bird Count Route in 1979-1982 after Dike Construction at the Park Street Mitigation Area, Eureka, California 9
4 Habitat Types and Bird Count Route at Bay Street Control Area, Eureka, California, 1980-1982 11
5 Small Mammal Trapping Grid Locations at the Mitigation Area, Eureka, California, in 1979, 1980 and 1981 20
6 Small Mammal Trapping Grid Locations at the Control Area, Eureka, California, in 1980 and 1981 21
7 Mean Number of Snowy Egrets in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 31
8 Mean Number of Black-shouldered Kites in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 . 33
9 Mean Number of Shorebirds in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 34
10 Mean Number of Killdeer in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 36
11 Mean Number of Greater Yellowlegs in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 37
xii xiii
Figure Page
12 Mean Number of Least Sandpipers in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 38
13 Mean Number of Common Snipe in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 39
14 Mean Number of Vaux's Swifts in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 42
15 Mean Number of Northern Rough-winged Swallows in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 44
16 Mean Number of Cliff Swallows in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 45
17 Mean Number of Barn Swallows in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 46
18 Mean Number of Marsh Wrens in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 47
19 Mean Number of Savannah Sparrows in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 49
20 Mean Number of Song Sparrows in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 52
21 Mean Number of Western Meadowlarks in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 54
22 Mean Number of American Goldfinches in Salt Marsh at the Mitigation and Control Areas, Eureka, California, June 1979 to June 1982 56 INTRODUCTION
Along with many other bays, Humboldt Bay in northern coastal
California has had a drastic reduction in the acreage of surrounding salt marsh in the last hundred years caused by diking, filling, and industrial development (Monroe et al. 1973, Shapiro et al. 1980). In recent years, however, preservation and restoration of salt marshes for their wildlife and human values have been emphasized (Monroe et al.
1973, Daiber 1977). Unfortunately, saltmarsh restoration, especially with emphasis on fish and wildlife resources, is a relatively new concept (LaRoe 1979), and techniques are not yet well developed.
Recently, however, guidelines for restoration based on experience to date have been set forth (Harvey et al. 1983, Josselyn and Buchholz
1984, Zedler 1984).
In order to offset the loss of or damage to wildlife resources from proposed water-resource development programs, the Fish and Wildlife
Coordination Act of 1934 as amended (U.S. Code 1982) requires the preparation of proposed measures for "mitigating or compensating" for such damage, but does not define the terms. The President's Council on
Environmental Quality (1983) defined the term "mitigation" in the
National Environmental Policy Act (U.S. Code 1976) regulations to include among other measures the "compensating for the impact by replacing or providing substitute resources or environments." The U. S.
Fish and Wildlife Service (USFWS) (1981) has adopted this definition and has recommended various mitigation means and measures including
1 2
compensating for impacts by conducting habitat construction activities
to fully restore or rehabilitate previously altered habitat. In this
context the USFWS has defined compensation as "full replacement of
project-induced losses to fish and wildlife resources." The California
Coastal Act of 1976 (California Public Resources Code 1976) requires
mitigation in certain cases of coastal land use development, especially
where the degradation of wetlands is concerned. Claycomb (1983)
contains further discussion of pertinent legislation.
The construction of the Woodley Island marina in Humboldt Bay
(Figure 1) by the Humboldt Harbor, Recreation and Conservation District
in 1979-81 destroyed 1.2 ha of salt marsh, 2.4 ha of mudflat, and 3.2 ha
of riparian and upland habitat. As part of the permit from the
California Coastal Commission, in which "mitigation" of this loss was
required, the District purchased, in 1979, a 7.9-ha property at the end
of Park Street, Eureka, California. It encompassed a 5.5-ha abandoned
log pond, formerly a tidal salt marsh, and was to be restored to its
original condition by breaching a portion of the surrounding dike. In
addition, an interior dike was to be constructed to enhance an existing
freshwater marsh. From June 1979 to July 1982 this study was conducted
to determine the success of the restoration program. In order to
provide a standard against which the degree of success could be
measured, a nearby undiked salt marsh at the end of Bay Street was
selected as a control area.
Prior to initiation of this project, no saltmarsh restoration
projects had been attempted in Humboldt Bay. In August 1980, dike
breaching along the lower part of Elk River, an estuary of mid Humboldt
Bay, commenced. Preliminary findings indicated that restoration of the 3 Area on Humboldt Bay, Eureka, California. Area on Humboldt Bay, Eureka, Figure 1. Location of Woodley Island Marina Site, Park Street Mitigation Area, and Bay Street Control Island Marina Site, Park Street Mitigation Area, Figure 1. Location of Woodley 4 saltmarsh vegetation and its associated bird and mammal life was
proceeding at a relatively slow rate because the high elevation of the area did not permit complete tidal flooding (Stopher et al. 1981, Base
1982b, Caltrans 1982).
Another saltmarsh restoration project was undertaken at the
Bracut marsh in North Humboldt (Arcata) Bay in 1981 through excavating fill, breaching the surrounding dike, and planting marsh vegetation
(Josselyn and Buchholz 1982). Although no study of the project has been
conducted, general observation indicates that restoration is proceeding
slowly (Paul Springer, Wildlife Department, Humboldt State University,
Arcata, CA 95521). In September 1983 a third saltmarsh restoration
project was initiated by installation of a tide gate on a degraded marsh
along Elk River, 0.4 km northwest of the previous project there.
Results to date show little change in the vegetation (Gail Newton, P. 0.
Box 234, Arcata, CA 95521) because the tide gate was not opened until
November 1984.
Similarly, the results of saltmarsh restoration in other areas
along the Pacific Coast are not always directly comparable because the ecological factors at the marshes are quite different (Macdonald 1977).
In San Francisco Bay and south the halophyte growing at the lowest elevation is California Cordgrass (Spartina foliosa) (Macdonald 1977), and, particularly in southern California where precipitation is low, soil moisture is provided primarily by seawater (Zedler 1982). In
Humboldt Bay, precipitation is heavy and Pickleweed (Salicornia
virginica) is the halophyte found at the lowest elevation. North of
Humboldt Bay, increasingly heavier precipitation permits tidal marsh 5 vegetation to become established at lower elevations in relation to mean sea level (Eilers 1975, Jefferson 1975).
In a study in the Salmon River estuary of Oregon, Mitchell
(1981) found that salt marsh was being restored successfully during the
2 years following breaching of a dike; however, the restored marsh would not be the same functionally and compositionally as the marsh prior to diking because too many changes had occurred that changed the potential of the site. She did not address the restoration as it affected the fauna of the site.
The primary objective of this study was to determine the success of saltmarsh restoration at the mitigation area at Park Street based on
(1) utilization of the area by amphibians, reptiles, birds and mammals before and after breaching of the dike, and (2) comparison with conditions at a nearby control salt marsh at Bay Street. A secondary objective was to determine the changes caused by construction of an interior dike within the mitigation area to enhance an existing freshwater marsh. STUDY AREAS
A detailed description of both the restoration site at Park
Street and the control marsh at the foot of Bay Street is provided by
Claycomb (1983). Following is a summary. Plant names primarily follow
Munz and Keck (1973) or Mason (1957).
The abandoned log pond portion of the Park Street property
(Figure 2), hereafter referred to as the mitigation area, is located
along Freshwater Slough at the north side at the end of Park Street
(40°47'N, 124°07'W, Arcata South Quadrangle, USGS). Three distinct
habitats were present on the mitigation area prior to dike breaching:
grassland and remnant salt marsh, freshwater marsh, and dike. The central area (3.8 ha) became tidal when the north dike was breached on
18 December 1980. Grassland (3.2 ha) within the central area consisted of Hairgrass (Deschampsia caespitosa) and escaped cultivated species
including Beard Grass (Polypogon monspeliensis), Italian Ryegrass
(Lolium multiflorum), and Velvet Grass (Holcus lanatus). Also occurring
were nongrass species including Bird's-foot Trefoil (Lotus
formosissimus), Brass-buttons (Cotula coronopifolia), Saltbush (Atriplex
patula), Curly Dock (Rumex crispus), and Salt Rush (Juncus lesueurii).
About 0.6 ha of the central area was remnant salt marsh consisting of
Saltgrass (Distichlis spicata), Pickleweed, Cordgrass (Recent studies by
Spicher [1984] indicate that the species growing at Humboldt Bay is not
California Cordgrass but, instead, Spartina densiflora, which was
introduced from South America), Arrow-grass (Triglochin maritimum), and
6 7
Figure 2. Habitat Types in 1979-80 before Dike Breaching and Bird Count Route in 1979 before Dike Construction at the Park Street Mitigation Area, Eureka, California. 8
Ditch-grass (Ruppia maritime). At the beginning of the study in June
1979 a freshwater marsh formed by springs was located in the southwest corner of the central area. Typical freshwater marsh plants such as
Common Cat-tail (Typha latifolia) graded into the remainder of the central area and also grew thickly along the drainage ditch from the freshwater marsh to the north end of the central area. In August 1979 an interior dike was constructed and a tide gate installed to separate the freshwater marsh from the rest of the central area and to prevent inflow of tidal water after breaching of the north dike.
In the year after dike breaching (Figure 3) the central grassland became increasingly similar to typical Humboldt Bay salt marsh characterized by Pickleweed, Saltgrass, and Cordgrass. Plants present that were not adapted to saline conditions declined, and the areas where those species had occurred were covered with decaying vegetation for much of the remaining study period. Ponded areas occurred in the northwestern, northeastern, and southwestern corners of the original grassland where no drainage systems had yet developed
(Claycomb 1983).
The freshwater marsh (0.7 ha) at the southwest corner of the study area increased in depth from about 0.2 m to 1.2 m after construction of the interior dike. The marsh supported typical freshwater marsh plants including Common Cat-tail, Marsh Pennywort
(Hydrocotyle ranunculoides), Willow (Salix hookeriana), Water Celery
(Oenanthe sarmentosa), and Red Alder (Alnus oregana) and a brackish water plant, Salt Rush, that continued to survive (Claycomb 1983).
Dikes occupied 1.0 ha and prevented tidal flow into the small freshwater marsh on the mitigation area, and, before breaching, the 9
Figure 3. Habitat Types in 1981 after Dike Breaching and Bird Count Route in 1979-1982 after Dike Construction at the Park Street Mitigation Area, Eureka, California. 10 central restoration area. Some of the existing dikes were reinforced in
August 1979 when the dike that protects the freshwater marsh was constructed. This resulted in the loss of a well-developed patch of
California Blackberries (Rubus vitifolius) along the northern edge of the mitigation area. The newly built and reinforced dikes were fertilized and seeded with Soft Chess (Bromus mollis), Tall Fescue
(Festuca arundinacea), Red Alder, and California Blackberry soon after construction, but germination was poor (W. Woodroof, California Coastal
Commission, Eureka, CA 95501). The principal vegetation developing on the new dikes were several species of grasses including Hairgrass,
Velvet Grass, Italian Ryegrass, and Beard Grass and non-grass species such as Bird's-foot Trefoil. Vegetation on the undisturbed dikes consisted primarily of Coyote Brush (Baccharis pilularis), Wax-myrtle
(Myrica californica), and California Blackberry. A small patch of salt marsh consisting of Cordgrass and Pickleweed bordered the eastern dike along Freshwater Slough (Claycomb 1983). Although this was not strictly dike habitat, it was most practical to include this and the narrow stringers of typical saltmarsh vegetation immediately bordering the slough with the dike habitat during the bird counts.
Freshwater Slough (1.3 ha), an estuarine branch of Eureka
Slough, bordered the mitigation area on the east. Hillside (1.1 ha) made up the remainder of the 7.9-ha tract purchased by the Humboldt
Harbor, Recreation and Conservation District.
The undiked marsh at Bay Street (Figure 4), hereafter referred to as the control area, was about 1.5 km west of the mitigation site, closer to the bay (40°48'N, 124°08'W, Eureka Quadrangle, USGS), and occupied approximately 6.2 ha. It was a kidney-shaped tract bordered by 11 Figure 4. Habitat Types and Bird Count Route at Bay Street Control Area, Eureka, California, 1980-1982. Bird Count Route at Bay Street Control Area, Eureka, Figure 4. Habitat Types and 12
First Slough and Eureka Slough on the north; Third Slough on the east; uplands, additional marsh, and a low dike on the south; and a recreational vehicle park on the west. Eureka Slough is a major estuary of Humboldt Bay; the others are branch estuaries. Another small estuary, Second Slough, bisected the area.
The control area had three distinct habitats. The largest was salt marsh (5.1 ha). Pickleweed, Saltgrass, and Cordgrass were dominant
species, and Salt Rush, Saltbush, and Gum-plant (Grindelia stricta) were
secondary species. Salt marsh was used as the standard to appraise the
success of restoration at the mitigation area after breaching.
Dikes occupied 0.4 ha. The dike parallel to Eureka Slough was dominated by Coyote Brush and Hairgrass. The higher ground separating
the recreational vehicle park from the marsh also was classed as dike
because of the similarity in function and vegetation. In addition to
the previously mentioned species, it also supported Argentine Pampas-
grass (Cortaderia selloana) and Willow. The low dike west of the foot
of Bay Street was occupied predominantly by Hairgrass and bare ground.
The last habitat at the control area was tidal slough (0.7 ha).
It included the channels of Second and Third Sloughs up to the permanent
vegetation line but excluded all of Eureka Slough and First Slough.
The control area experienced no habitat changes during the
study. METHODS
Amphibians and Reptiles
No systematic sampling was performed for amphibians and reptiles. Any species observed on the regular bird count was noted, along with its location. Amphibian and reptile names follow Stebbins
(1966).
Birds
Two general techniques were used to count birds: mapping territories from late May to early July during the breeding season (Hall
1964) and mapping the location of individuals during both the breeding and non-breeding season to determine a mean population index, following the method described by Kolb (1965) for winter bird counts. The spacing between adjacent count routes within an area was established to allow detectability of all species but far enough apart to allow the count to be completed within 2 hours' time. At the mitigation area this route
(Figures 2 and 3) was generally a series of north-south transects approximately 70 m apart (45 m to 90 m). At the control area (Figure 4) a circuitous route ranging from 2 m to about 70 m between adjacent parts of the route was required to cover all portions of the site adequately.
Analysis for the summer season also included those visits made during the breeding bird counts.
13 14
For both methods, birds flying over the area but not obviously
feeding within it were not counted; however, birds such as swallows or
raptors which foraged or hunted over the area were counted. Swallows
and Vaux's Swifts (Chaetura vauxi) presented a difficult counting
problem due to their mobility. A 5-minute index (Sorensen 1977) was
attempted during summer 1979 and abandoned after a study of the data
showed obvious inaccuracies. An estimate of the average number of
individuals during a count yielded a more intuitively satisfying result.
However, error was probably higher in the late summer when more
individuals were present. Although swifts and swallows used all
habitats, they were arbitrarily recorded as occurring in the grassland
(before breaching) or salt marsh (after breaching) of the mitigation
area and the salt marsh of the control area for practical reasons.
Because too few samples were taken during the summer of 1979, this
season was excluded from analysis of results for swallows, but their
numbers are included in the graphs.
Counts were conducted from June 1979 through June 1982 at the
mitigation area and from June 1980 through June 1982 at the control area
(Table 1). Between July and November 1979 the number of individuals was
tallied but their location was not mapped, so this period is excluded in
the statistical analyses. Generally, eight counts were considered
adequate to delineate the territories of all breeding pairs. One
breeding bird count (BBC) per year was conducted at each site, for a
total of four at the mitigation area (1979-1982) and three at the
control area (1980-1982). 15
Table 1. Breeding Bird Count Dates at the Mitigation and Control Areas, Eureka, California.