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                                                                                 PB82-2C-6156
                                                                           1:PA  600/3-82-OS2
                                                                           Sentcmber 1982
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     DISTRIBUTION AUB ABUNDANCE OF WATERFOWL ANT)
  ^SUBMERGED AQUATIC VEGETATION IN CHESAPEAKE BAY
 -
                           by
               Robert E. Munro              ~ ',
                     and
               Matthew C.  Perry             ^'•1-v
        U.S. Fish and Wildlife Service
Migratory Bird and Habitat Research  Laboratory
            Laurel, Maryland  20811
     Interagency Agreement Number EPA-78-D-X0391
                    Project Officer

                    William A.  Cook
                 Chesapeake Bay Program
         U.S.  Environmental Protection Agency
                       Region III
                 6th and  Walnut Streets
           Philadelphia, Pennsylvania 19106
          OFFICE OF  RESEARCH  AND  DEVELOPMENT
         U.S.  ENVIRONMENTAL PROTECTION AGENCY
               WASHINGTON,  D.C.   20460
                                                            - - .L ,'i^ i »t
                                                           FA  15107
   , IL
                                              NATIONAL  TECHNICAL
                                             INFORMATION SERVICE
                                                 US DtCARlMfXl Of CO«M!RC(
                                                   SPtlNSflllP. »» "161
                                                       EPA Report Collection
                                                    Information Resource Center
                                                         US EPA Region 3
                                                      Philadelphia, PA 19107

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Abstract
Distribution and abundance of waterfowl and submerged aquatic vegetation in
Chesapeake Bay
Munro, Robert E.; Perry, Matthew C.
Fish and Wildlife Service, Laurel, MD. Migratory Bird and Habitat Research
Lab.; Environmental Protection Agency, Philadelphia, PA. Chesapeake Bay Program.
U.S. Environmental Protection Agency ;
1982
EPA 600/3-82-082; EPA/CBP-TR-001S
PB82-266156
11525886
Aquatic plants; Birds; Ecology, Chesapeake Bay; Surveys; Distribution(Property);
Abundance; Food habits; Periodic variations; Animal migrations; Ducks; Statistical
analysis; Trends; Maryland; Virginia; Waterfowl; Habitats; Swans; Geese
Waterfowl-Chesapeake Bay (Md. and Va.) ; Birds-Chesapeake Bay (Md. and Va.) ;
Aquatic plants-Chesapeake Bay (Md. and Va.) ; Chesapeake Bay (Md. and Va.)
xi, 180 p. ; 28cm.
LIBRARY CALL NUMBER LOCATION
EJAD EPA 600/3-82-082 Region 3
Library/Philadelphia,
PA
NTIS PB82-266156 Most EPA libraries have a NTIS
fiche copy filed under the call
number shown. Check with
individual libraries about
paper copy.
Waterfowl species populations in Maryland and Virginia portions of Chesapeake Bay were
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examined during long-term and current (1972-1980) periods to identify trends in
distribution and abundance. Comparisons were also made between State and Atlantic
Flyway populations and waterfowl species distributions among survey areas. Distribution
and abundance of submerged aquatic vegetation among waterfowl survey areas in Md.
were summarized for 7 plant species during 9 years (1971-1979). These data (SAV
species combined) were used to test the hypothesis that annual variation in area
populations of waterfowl was related to variation in the abundance of SAV, following an
adjustment for annual variation in the general abundance of waterfowl. There were few
statistically significant relationships between distribution and abundance of waterfowl and
SAV.
"EPA 600/3-82-082" ; available from NTIS as "PB82-266156."
Perry, Matthew C.
United States. Environmental Protection Agency. Chesapeake Bay Program. ; United
States. Environmental Protection Agency. Chesapeake Bay Program.
Washington, D.C. :
1982.
PC A09/MF A01
LIBRARY Date Modified
EJA 19961025
m
19961015161115
OCLC/T
ENG
OCLC
MERGE
01 149cam 2200265la 45020
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                                                                                    -*• ,'x- """ ' '•'
                                   TECHNICAL REPORT DATA
  3iaC^T NO
   RPA 600/3-82-082
,3. 3.£C:P' = NT S ACCESSIO'»M
;  PB82	266156
  Distribution & Abundance of Waterfowl and Submerged
  Aquatic Vegetation  in  the Chesapeake Bay
  September  1P82
16. ("SflFOOMIMG ORGANIZATION CODE
  Robert ^E. Munro and Matthew C.  Perry
,3 "HFGHMING ORGANIZATION fl£aCST NO

I CBP-TR-001S
9 =>£»e3RMING 0^3 A\:ZATi C^ VA'.tS ANO A3CPESS
  U. S. Fish & Wildlife Service, Migratory Bird and
  Habitat Research Laboratory, Laurel,  Maryland 20811
1*0. aROG3AM £L=M£NT NO.
i 2BA703	
;; 1. CONTRACT GRANT NO.
                                                            EPA 78-D-X0391
 12. SPC\SOPING AGENCY \AWE A.MC AOOHESS
  Region III, U. S. Environmental  Protection Agency
  Chesapeake Bay Program, 2083 West Street
  Annapolis, Maryland 21401
113. TV?E OF 3=PQRT AND "EBIOO COVSBEO
i Project Report  8/1/78-12/31/80
 14. SPONSORING AGENCY COCE
                                 .\OTES
 io. A8STSACT waterfowl species populations in Maryland & Virginia portions of Chesapeake
 Hay were examined during  long-term  and rurrent (1972-1980) periods to identify  trends  in
 distribution and abundance1.  Comparisons were also made between State and Atlantic  Fly-
 way populations & waterfowl species "iistributions among survey areas. Distribution  &
 abundance of SAV among waterfowl  survey areas in Md. were summarized for 7 plant  species
 during 9 years (1971-1979). These data (SAV species combined) were used to test  the
 hypothesis that annual variation  in area populations of waterfowl was related to  varia-
 tion in the abundance of  SAV,  following an adjustment for annual variation in the genera
 abundance of waterfowl.The distribution and abundance of SAV species declined in  Md.
 waters during the 1970's.  There  were few statistically significant relationships betwee i
 distribution and abunc.ance of  waterfowl and SAV.  But there was an implied biological
 relationship, since t.ie most important waterfowl wintering areas were also among  the
 most abundantly vegetated areas.  This report was submitted in partial fulfillment of
 Interagency Agreement No. EPA-78-D-X0391 by the U. S. Fish and Wildlife Service,
 Migratory Bird and habitat Research Laboratory under the sponsorship of thf Chesapeake
 Bay Program, U. S. £nviron.nental  Protection Agency.
                                <6Y .VCRDS ANO DOCUMENT ANALYSIS
                                             |t>..OENTIF' =
                                                          P£-. ENOED THPVS  ...  Cr.SAT' ; 'SlJ.OtCUp
      Unrestricted distribution
 SEC^PI'Y C-AS3
Unclassified
                                                                Rtport.
                                                                        ill NC C-
                 193
                                             120 SEC'JRlT^- CLJ.SS
                                               Unclassified
EPA FO'-n i220-l I9-73I

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                                                      DISCLAIMER
                         This  report has been reviewed  by the Chesapeake Bay Program, U.S.
                     Environmental Protection Agency,  and  approved- for publication.   Approval
                     does not  signify that the contents  necessarily reflect  the views and policies
                     of the  U.S.  Environmental Protection  Agency,  nor does mention of trade  names
                     or commercial products constitute endorsement or recommendation for use.
,*"-•

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V '
                                                    DISCLAIMER




                          This  report has been  reviewed by the Environmental Research




                     Laboratory - Narragan-sett,  Chesapeake Bay Program of the U.S.




                     Environmental  Protection Agency, and approved for publication.  Approval




                     does not signify that the  contents necessarily reflect the views and




                     policies of the U.S. Environmental Protection Agency, nor does mention




                     of  trade names or commercial products constitute e-'orsement or




                     recommendation for use.

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I
                                                  ABSTRACT




                       Waterfowl species populations in Maryland  and Virginia  portions of




                  Chesapeake Bay were examined during long-term and  current  (1972-80)




                  periods to identify trends in distribution and  abundance.  Comparisons




                  were also made between State and Atlantic Flyway populations,  and




                  waterfowl species distributions amonr, survey areas.




                       Distribution and abundance of submerged aquatic vegetation  (SAV)




                  among waterfowl survey areas in Maryland were summarized for seven plant




                  species during nine years (1971-79).   These data  (SAV species  combined)




                  were used to test the hypothesis that annual variation in  area




                  populations of waterfowl was related  to variation  in the abundance of




                  SAV, following an adjustment for annual variation  in the general




                  abundance of waterfowl.




                       Waterfowl food habits were summarized and  analyzed when possible.




                  These dat<  provided additional evidence on relationships between




                  waterfowl and SAV,  and indications of changing  food preferences  within




                  some species.




                       Recent Maryland waterfowl populations (with one exception)




                  generally declined, while those in Virginia increased or remained




                  stable.  Species dispersion among areas was typically stable.  The most




                  important waterfowl wintering ureas in Chesapeake  Bay have been  and




                  continue to be the Chester River, Eastern Bay,  and the lower Choptank




                  River.




                       The distribution and abundance of all SAV  species declined  in




                  Maryland waters during the 1970's. During that period, SAV  was  nost




                  abundant in areas that included the lower Choptank River,  Manokin and




                  Annemessex Rivers,  Chester River, and Eastern Bay.



                                                       iv

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                         There were few statistically significant relationships  between




                    distribution and abundance of waterfowl  and SAV.   But  there  was  an




                    implied biological relationship,  since the  most  important  waterfowl




                    wintering areas were also anong the most abundantly vegetated  areas.




                         Waterfowl species adapted differently  to the  decline  of SAV during




                    the 1970's.  Whistling swans began feeding  in agricultural fields, while




                    others shifted food preferences from SAV to invertebrates.  However,  the




                    most common adaptation was to winter elsewhere in  the  Atlantic Flyway.




                    Due to the northern location of Chesapeake  Bay,  through which  many




                    waterfowl annually migrate,  current populations  of some species  are




                    believed to be temporarily low.  When exposed to increased distribution




                    and abundance of submerged aquatic vegetation, we  anticipate that




                    waterfowl species with preferences for SAV  and  associated  invertebrates




                    will respond positively.




                         This report was submitted in partial fulfillment  of Interagency




                    Agreement No. EPA-78-D-XO^jl by the U.S.  Fish and  Wildlife Service,




                    Migratory Bird and Habitat Research Laboratory under the sponsorship  of




                    the Chesapeake Bay Program of the U.S. Environmental Protection  Agency.




                    This report covers the period August 1,  1978,  to December  31,1980, and




                    work was completed as of  February 15,  1981.

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          \
\
                                CONTENTS
Foreward.
                                                                     ill
Abstract	   iv




Acknowledgment	vlii




Executive.- Summary	   Ix




     1.  Introduction 	  1




     2.  Conclusions	4




     3.  Materials and Methods	7




           Sources and limitations of data	7




                Waterfowl food habits 	 .....  7




                Waterfowl populations 	  9




                Submerged aquatic vegetation	13




           Statistical analyses 	 14




     4.  Results and Discussion	16




           Waterfowl food habits	16




           Distribution and abundance of SAV	25




           Chesapeake Bay waterfowl populations in percpective- ... 31




                Maryland	.31




                Virginia	40




           Effects of SAV on waterfowl	45




                 Swans and geese.	46




                 Black ducks and mallards	48




                 Diving ducks 	 48




           Impacts of waterfowl on SAV	59




References	60




Appendices




  A-  Scientific and common names of waterfowl food items 	 62





                                    vi

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                        B.  Food itens of waterfowl during the period 1890-1959 	  71




                        C.  Food items of waterfowl during the period 1960-1969	107




                        D.  Food items of watertowl during the period 1970-1979	128




                        E.  Frequency occurrence of SAV by species during 1971-1979  •  .  •  .152




                        F.  Summary of Maryland waterfowl populations by species and  area  .159




                        G.  Summary of Virginia waterfowl populations by species and  area  .172
                                                          vii

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                            ACKNOWLEDGMENTS




     We are indebted to B. K. Williams (primary responsibility), P. H.




'•oissler, L. M. Moyer, and E. D. Heiniein for statistical consultation.




'.e also thank V. G. Perros, L. D. Eastman, C. J. Ellis, P. Longabucco,




P. L. Miller, and J. M. Wilcox for technical assistance.  Many people




assisted in food habits analyses useJ in this report and their input is




appreciated.  Of special importance were C. K. Rawls for his work during




the 1960's, and F. M. Uhler whose contributions have spanned decades.




     V. D. Stotts (Maryland Wildlife Administration) offered advice and




expeitise on Maryland waterfowl populations, as well as access to




original data from periodic surveys of submerged aquatic vegetation.  To




the many pilots and biologists who participated in surveys of waterfowl




or vegetation in Chesapeake Bay, we express our gratitude.  H. F.




Percival and B. S. Brun provided valuable administrative expertise.  We




are finally indebted to G. M. Haramis, C. F. Kimball, and J. D. Nichols




for their critical review of this report.
                                   viii

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                           EXEGITTIVE SUMMARY


     The Chesapeake Bay Is the largest estuarlne system along the


Atlantic coast of North America, and the most important wintering area


in  the Atlantic Fl'-way for more than 1.5 million waterfowl (including


ducks, geese, and swans).  Historically (1956-), the Maryland portion of


the Bay has been the principal Flyway wintering area for Canada geese


(Branta canadensis), whistling swans (Cygnus colnmbianus columbianus),


and canvasbacks (Aythya valisineria), and a primary wintering area for


ruddy ducks (Oxyura jamaicensts). common goldeneyes (Bucephala clangula


anericana), redheads (A_. americana), black ducks (Anas rubripes), and


mallards (A_. platyrhynchos).  In addition to wintering other popul-


ations, the Bay provides a resting and feeding place for countless birds


that migrate to and fron wintering areas farther south.  The Pay is


therefore vital as a wintering area and as a link in a chain of resting


areas.


     The traditional importance of Chesapeake Bay to vaterfowl was due


in  large measure to vast beds of submerged aquatic vegetation, which


included species such as widgeongrass (Ruppia maritima), wildcelery


(Vallisneria americana), and sago pondweed (Potamogfcton pectinatus).


But during the 1970's those waterfowl food resources declined, which in


large measure prompted this work.  The main objectives of this report


are to document historic and current distribution, abundance, and food


preferences of waterfowl that winter in Chesapeake Bay, and examine


current relationships between waterfowl and the distribution and


abundance of SAV.


     Linear regression and analysis of varirnce techniques were used to


detect trends in distribution and abundance of waterfowl species


                                    ix

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populations that winter in Maryland and Virginia, and to examine


waterfowl-SAV relationships with simple, linear models.  W?terfowl food


habits were analyzed where possible with a ranking procedure, which


compared historic and current usage of importa"*- lood items.


     Maryland and Virginia waterfowl populations differed considerably


in species diversity and numbers of birds.  The smaller Virginia


populations were also more stable than those in Maryland.  Hovever,


distributions of species populations amorg survey areas in both States


were very consistent over years.


     Distribution and abundance of SAV in Maryland waters declined


luring the 1970's, and particularly frors 1971 to '973.  Of the seven SAV


species considered, all declined; however, widgeongrass remained


relatively abundant and widely distributed.  SAV was isost abundant in


the lower Choptank River, Manokin and Annemessex fivers area, Chester


River, and Eastern Bay during the 1971-79 period.


     The common response by waterfowl species with known preferences


for SAV (e.g., redheads, American wigeon [Anas americana], and pintails


(A. acuta]) was to winter elsewhere in the Atlantic Flyway during the


1970's.  Others (whistling swans and ccnvasbacks) changed food


preferences.  The wintering Canada goose population in Maryland, which


now requires only open water for protection and agricultural fields for


food, continued a long-term increasa in numbers.


     The distribution (dispersion) and abundance of waterfowl populations


among Maryland survey areas generally reflected the distribution and


abundance of SAV, although few statistically significant relationships


were found.  The areas most important <:o wintering waterfowl (Chester


River, Eastern Bay, and the lower Choptank River) prior to and during

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the 1970's were also among the most abundantly vegetated areas during

the 1970's.  It follows from this general relationship that waterfowl

populations have had no adverse or long-term impact on distribution and

abundance of  submerged aquatic vegetation.

     The currently low diversity of waterfowl species wintering in

Maryland is believed to be temporary, and limited  by the distribution

and abundance of SAV.  Since formerly abundant species of waterfowl

still migrate annually through the Bay area, a positive and rapid

response to an improvement in distribution and abundance of submerged

aquatic vegetation may be anticipated.
                                  xi

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                               SECTION 1


                              INTRODUCTION


     Chesapeake Bay, the largest estuarine system in North America,


contains a diverse collection of many plant and animal communities.


Waterfowl (Anatidae) are among the mrst provainent animals during fall


and winter.  Of 45 species of ducks, geese, and swans that are native to


North America (Bellrose 1976), about 30 regularly migrate through or


winter in the Bay area (Stewart 1962).  Although a few species such as


the black duck (Anas rubripes) and the mallard (j\. platvrhynchos) breed


in Bay marshes, most waterfowl ^re present in the area only from October


through March.  The use and importance of Chesapeake Pay to waterfowl


are therefore greatest during this period.


     Historically, the Bay has been the most important wintering area


along the Atlantic coast for a number of waterfowl species, such as the


whistling swan (Cygnus columhianus colunbianus), Canada goose (Branta


canadensis), and the canvasback (Aythya valisineria).  Other populations


have been far leas abundant.  Early migrants such as blue-winged teal


(Anas discors) spend little tine in the Bay before departing in


mid-October for wintering areas in South America.  Some winter


residents, due to their habitat preferences, are not obvious to the


casual observer.  For example, the sea ducks or scoters (Melanitta


spp.), as their colloquial names imply, occupy deep (>3 m), open-water


habitats generally beyond the littoral zone.  Others, such as the


Atlantic brant (Branta bcrnicla hrota), are usually found only in ocean


bay habitats (e.g.,  Sinepuxent Bay, Maryland).


     Ducks are often referred to as c-ither surface-feeding or puddle


ducks, as opposed to divir.f> or bay ducks.  Puddle ducks feed by dabbling

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at the water's surface or tipping up in shallow water, even though they




are capable of diving beneath the water's surface.  They typically




prefer shallow marshes and shoal riverine habitats, and as a group they




consume more plant naterial than do diving ducks.  The diving ducks all




share two physical characteristics that facilitate diving, their primary




method of feeding:  their wings are relatively snail and their legs are




set farther back on their body.  As a consequence, divers need open




water across which to run prior to flight, which prevents them from




feeding in dense marshes or agricultural fields.  Diving ducks are




usually found in exposed, deeper waters (1-5 ia), where vegetation is




less available.




     Subaerged aquatic vegetation (SAV) is an important food resource




for many waterfowl species (Cottam 1939: Martin and Uhler 1951; Yocom




1951; Stewart 1962; Bellrose 1976).  In his account of the canvasback,




Bent (1923:195) reported ". .  .  it [wildcelery (Vallisneria




araericana)] grows nost abundantly in the Chesapeake Bay region and is




supposed to be the chief attraction for the vast nursber of canvashacks-




and other ducks which resort to these waters in winter .  •   •".  The




traditional Importance of the Bay to many waterfowl populations provides




indirect evidence of a long-term stability in the Bay's aquatic plant




and animal food rasources.  But waterfowl populations and the food




resources upon which they depend are not stable in the short-term.




There have been changes in waterfowl distribution and abundance in the




Bay during the last 25 years, as well as more recent changes in




distribution and abundance of SAV.   The purpose of this study is to




examine relationships between waterfowl and SAV resources of the Bay.




     Specific objectives include an examination of 1) waterfowl food

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                     habits  in  the Bay  area,  2) distribution and  abundance  of  SAV,  3)  winter-


                     ing waterfowl populations, 4)  relationships  between waterfowl  and the


                     distribution and abundance of  ?AV,  and 5)  impacts of waterfowl on SAV.

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                               SECTION 2




                              CC XLUSIONS




     Historically, Maryland waterfowo. populations were more diverse in




species and more numerous thai, thosi in Virginia, where populations were




generally more stable.  Distributions (dispersions) of species




populations among survey areas in both States were remarkably stable.




The most important wintering areas for waterfowl in Chesapeake Bay are




the Chester River, Eastern Bay, and the lower Choptank River.




     Significant changes occurred in the Bay's submerged aquatic plant




resources during the 1970's.  SAV abundance declined considerably from




1971 to 1973, and fluctuated thereafter.  Of the seven plants examined,




widgeongrass (Ruppia maritima) was the most widely distributed (number




of areas) and abundant (highest frequency occurrence)  species of SAV in




Maryland.  However, all species declined during the 1971-79 period.  SAV




in Maryland was most abundant In the lower Choptank River,  Manokin and




Annemessex Rivers area, Chester River, and Eastern Bay.




     Species of waterfowl adapted differently to these conditions.




Increased numbers of whistling swans began feeding in  agricultural




fields during the early 1970's, even though this behavior was well




established 10-15 years earlier on their northward nigrations.




Responses among ducks were affected by physical differences between the




two general categories examined.  Puddle or dabbline ducks, which




normally feed by tipping up in shallow water, are capable of feeding in




agricultural fields and dense marsh habitats.  Diving  or bay ducks have




evolved relatively shorter wings and other physical characteristics that




facilitate swimming beneath the water's surface where  they  typically




feed.  These birds, essentially incapable of feeding on land, could

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change their food preferences only within the estuarine environment.  Of




course, both puddle and diving ducks could winter elsewhere within or




outside of the Bay area.




     Maryland populations of the four puddle duck species examined were




considerably less abundant during the 1970's when collared to long-term




levels.  Pintail (Anas acuta) and American wigeon (A. americana)




populations, whl^h have known preferencas for SAV, are now essentially




absent from Maryland wintering areas.  Black duck and mallard




populations, which showed relationships to the distribution and




abundance of SAV, also decreased.  The current level of Maryland puddle




duck populations, as a group, is about one-third of the former level.




     Among diving ducks, canvasbacks and redheads (Avthya anericana)




were most affected by the recent decline in SAV.  Although species of




SAV were principal food items of canvasbacks prior to the 1970's, they




were essentially the only foods of redheads-  Canvasback food




preferences completely changed during the i970's to a diet that




consisted mainly of Baltic macoma clans (Maccraa balthica) and other




invertebrates.  Also during that netiod, Maryland wintered a decreasing




percentage of the Atlantic Flyway ^tvasback population.  Although




significant changes In the Flyway distribution of redheads occurred




prior to the 1970's, the most important difference was a consistently




low number of these birds throughout the 1970's.  The redhead population




in Maryland is now only a remnant of past populations.




     On the positive side, the Canada goose population in Maryland




continued to expand.  These birds began foraging on land during the mld-




1950's when the mechanization of agriculture Increased the availability




of waste (unharvested) cereal grains.  This population averaged about

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265,000 geese between 1956 and 1971, although it increased at a rate of




23,000 birds/year.  During recent years Maryland wintered more than




500,000 Canada geese.  The future of this population rests in




agricultural fields.




     Waterfowl populations have had no long-term impact on distribution




and abundance of SAV.  Prior to the 1970's, waterfowl populations were




greatest in areas such as Chester River, Eastern Bay, and the lower




Choptank River.  During the 1970's, SAV was most abundant in these areas




despite a history of u; •> by abundant waterfowl populations.




     The relatively low diversity of waterfowl species currently




wintering in Maryland, although nitigated somewhat by tremendous




populations of Canada geese, is cause for concern.  Hovever, the outlook




is optimistic due in large measure to the northern location of




Chesapeake Bay with respect to other important wintering areas.  Major




population segments of most Flyuay species annually stop in the Bay area




while on migration to and from wintering areas farther south.  These




populations are therefore exposed to Eay conditions on an annual basis.




We anticipate they would soon respond positively to an improvement in




distribution and abundance of submerged aquatic vegetation.

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                                                  SECTION 3




                                            MATERIALS AND METHODS




                   SOURCES AND LIMITATIONS OF  DATA




                   Waterfowl Food  Habits




                        Food habits of Chesapeake  Bay waterfowl  were summarized  by Stewart




                   (1962).  His original data,  collected mostly  during  the  1950's,  were




                   obtained and supplemented with  additional  samples for  some  species.




                   Although methods and specific areas of collection were identifiable for




                   most samples, these potentially important  sources of variation were




                   disregarded  to  increase sample  si.Tes  and provide  a Bay-wide description




                   of  food items used  by waterfowl.   Other  sources of variation, such as




                   the  organ from which foods were obtained and  identified  (Swanson and




                   Bartonek 1970),  temporal  or long-term  changes  (Perry and IThler '976),




                   and  differences  between species (Stewart 1962) were included.




                        Stewart (1962:2)  determined  the  total volume of food contents for




                   an organ sample  by volumetric displacement, and then visually estimated




                   the  proportion of each  food item.  Samples from other collections




                   differed  only in that volumes for each food itero were separately




                  measured, or that organ  (source) was not recorded.  Scientific and




                  common names of Bay waterfowl food items are shown in Appendix Table A-l




                       Various measures t.ave been used to describe the  food habits of




                  animals  (e.g., Martin et al. 1946; Korschgen 1969; Swnnson et  al. 1974).




                  The latter authors demonstrated  that the aggregate percent method was




                  favored over the aggregate volume method.  Briefly, if  ve have J food




                  items and I birds,  the aggregate percentage is the proportion  of  the jth




                  food item in the  ith bird averaged over all samples.  This procedure




                  gives equal weight  to each bird  in the sample,  and is not affected by

-------
variation in individual sample volumes (Swanson et al. 1974:303); this




is particularly appropriate here since most samples were opportun-




istically obtained.  In addition to aggregate percentages, we also




determined the frequency occurrence, of each food iten in the sample.




     Since we have no measures of availability for many food items used




by Bay waterfowl, we must exercisfi caution when discussing preference,




selection, or importance.  We are consequently limited in the questions




or hypotheses we may examine with these drta.  Johnson (1980) discussed




these problems and proposed a procedure for comparing ranks of usage and




availability measurements.  Johnson's method basically compares two




nultivariate normal vectors of means (ranks), and is sufficiently




general for our purposes.  Here we are interested in the hypothesis of




no difference between two samples in utilization of important food




items.  Our application of Johnson's method therefore compares two




levels of usage rather than usage and availability.




     In a preliminary series of analyses we w^re interested in whether




or not samples of food items identified in two different organs were




sufficiently similar to allow combination.  In practice, we identified a




small list (i.e., the number of items was small compared to available




sample sizes) of items that were Important (frequently utilized) in




samples from organ A.  We then determined the rank of each food item on




the list with respect to the other listed items for each individual in




both organ samples A and B.  Mean ranks of usage for each food item




vithin each category (organ) were ther computed and compared through a




multivarlate analysis of variance.  We used this approach for all




comparative food habits analyses.

-------
I




I




I
Waterfowl Populations




     The U.S. Fish and Wildlife Service (USFWS) in cooperation with the




States conducts annual surveys of wintering waterfowl during January,




when population distributions are nost stable and perhaps most




concentrated.  According to Benning et al.  (1978:1), "This survey




primarily monitors annual changes in waterfowl distribution on the




wintering grounds within the United States, and indicates annual




population trends".  Although survey results are affected by such




factors as weathsr, personnel changes, and  errors of omission (birds not




found) and commission (birds found but counted inaccurately), they




constitute the only long-term source of information on the distribution




and abundance of wintering waterfowl.  Most states have developed




systens of survey areas which generally encenrpass rivers, bays, or large




expanses of marsh habitat.  Survey area boundaries were stable in




Maryland (Figure 1) by 1956 and in Virginia (Figure 2) by 1961.




Waterfowl surveys in the Potomac River, ocean bays, and inland




reservoirs and swamps were not included in  analyses of species




distributions among sarvey areas, since comparable data on submerged




aquatic vegetation were not available.  Some waterfowl species (e.g.,




Atlantic brant, greater snow goose [Anser caerulescens atlantica]).




whose populations were generally restricted to those habitats, were




consequently omitted.  An index to waterfowl survey areas used in this




study is provided in Table 1.




     The Atlantic Flyway, which encompasses Atlantic coastal States from




Maine to Florida, is one of four U.S. flyways used in the management of




waterfcwl populations.  United States and Atlantic Flyway populations




can be used as measures of general abundance or trend that may affect

-------
Figure 1.   Maryland waterfowl survey areas used in this report.
                                   10

-------
Figure 2.   Virginia waterfowl survey areas used in this report.
                                 11

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IT' '
i 1
Chesapeake Bay numbers of ducks, geese, or swans.  We can also use these




data  to identify species that have a tradition of wintering in the Bay.




Since waterfowl populations and habitats in Maryland and Virginia




portions of the Bay are considerably different, we will consider




populations in these areas separately.




     We used mid-winter survey results prior to 1972 as measures of




Ion3-tenn distribution and abundance of United States, Atlantic Flyway,




and Maryland and Virginia wintering waterfowl populations.  We first




identified w?terfowl species whose Bay populations averaged five percent




or more of Atlantic Flyway populations prior to 1972.  Species that did




not meet this criterion were subsequently omitted.  For the remaining




species we then examined population segments (survey areas, Flyway, and




U.S.) for trends in distribution cr abundance prior to the early 1970's.




We used survey results during the years 1972-80 to describe current Bay




waterfowl resources in a similar manner.  Comparisons could then be made




between waterfowl populations during the long-term and current periods,




and between current waterfowl and SAV distribution and abundance.




Submerged Aquatic Vegetation




     Stevenson and Confer (1978) summarized available information on the




distribution and abundance of submerged aquatic vegetation (SAV) in




Chesapeake Bay.  Bayley e* al. (1978) Ascribed long-term (1958-1975)




changes in SAV at the head of the Bay (Susquehanna Flats) , ana Orth et




al. (1979) documented the distribution and abundance of SAV in Virginia




waters during tiie 1970's (1971,1974,  and 1978).  Kerwin et al. (1976)




initiated an extensive (Maryland) and long-term (1971-) survey of f.AV




with which the distribution and abundance of waterfowl could be




compared.  Kerwin's study, which evolved into a cooperative effort






                                    13

-------
I
I
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 I

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 I
              between  the USFWS Migratory Bird and Habitat Research Laboratory  (MBHRL)


              and  the  Wildlife Administration of Maryland Department of Natural


              Resources  (DNR), was based on a stratified random sampling (about 640


              stations)  of shoal water habitats (<2.44 m at mean low water) in


              Maryland (excluding the Potomac River).  To monitor annual trends in

ir
f              distribution and abundance of SAV, established stations were sampled

I
£,             once each  summer for presence/absence and percent composition of  SAV.

*•.
              Procedures are  further described in Kerwln et al. (1976).


                    Since waterfowl survey area boundaries did not correspond to those


              of the SAV survey, we examined latitude-longitude coordinates of sample


              stations in the SAV survey to resolve boundary discrepancies.  We were


              then able  «-o measure (planimeter) the extent of shoal water habitat in


              24 waterfowl survey areas and summarize SAV survey results accordingly.


              We estimated hectares of SAV in an area by multiplying the size of the


              area by  the proportion of stations sampled that were vegetated.  These


              estimates  of SAV are biased upwards due to the inherent assumption of


              complete coverage (dense bed) at a vegetated station.  Use of these


              estimates  to represent distribution and abundance of SAV during winter


              is another limitation.


              STATISTICAL ANALYSES


                    Analyses of variance and linear regression used in this report


              followed the Statistical Analysis System user's guide (Barr et al.


              1976).   We generally used the simplest (fewest parameters) linear model


              that adequately described the relationship between a dependent and one


              or more  independent variables.  Where appropriate, area-specific effects


               ere included if they were significant (p < 0.05).  In models with more


              than one independent variable, we used sums of squares and corresponding



                                                  14

-------
r.
».
                  F statistics that reflected adjustments for all other variables in the
                  model.
                       Given annual estimates of SA^T (summer in year t-1) and waterfowl
                  (January in year t) in a number of survey areas, we were then able to
                  formulate the following linear model:
                                   n    = A + H  (SAV    ,) + C  (ATL ),
                                    a,t        a     a,t-1     a     t
                  where n    is the predicted number of birds in survey area a, A is an
                         a, t
                  overall intercept term, B  is the area-specific effect of SAV, SAV
                                           a                                        a,t—i
                  is the number of hectares of SAV in area a during the previous summer,
                  C  is the area-speoific effect of general species abundance, and ATL  is
                   3                                                                  t
                  the Atlantic Fiyway population during the current year t.  The intercept
                  term was included to determine if a significant portion of an area's
                  population was independent of variation in SAV or general species
                  abundance.  The Atlantic Fiyway population was included to measure the
                  extent to which changes in an area's population merely reflected annual
                  variation in the size of the geo&r.a4. population.  If an effect was not
                  significant (p > 0.05), it was removed and a reduced model was then
                  considered for that species.
                                                      15

-------
I
y            •                   \
                                                    SECTION 4




                                              RESULTS AND DISCUSSION




                      WATERFOWL  FOOD  HABITS




                           A study  of waterfowl  food habits is a  study of variability,  some of




                      which (e.g.,  areas  and methods of collection)  is unavoidable.  More  than




                      3,000 food habits records  accumulated by Stewart (1962), Rawls (in




                      prep.)i  and data from  the  files  (MBHRL, Laurel, Maryland)  included about




                      700  different food  items  (Appendix A).  These  data were  tabulated by




                      waterfowl  species,  tine period (pre-1960, 1960's, and the  1970's  in




                      Appendices B, C, and D), and  organ source (gizzard, gullet, or unknown).




                           Preliminary analyses  using Johnson's (1980) methodology indicated




                      there were substantial differences in usage (aggregate percentages)  of




                      food items in samples  from different organs.   Since this variation




                      persisted  across a  number  of  waterfowl species, we restricted food




                      habits comparisons  to  those species frr which  sufficient gizzard  samples




                      were available. We also were unable to statistically compare 1960's




                      food habits data (Rawls, In prep.) with those  from earlier or later




                      periods, since  we lacked data from individual  samples.




                           A gross  comparison of the incidence of plant material in samples of




                      Bay  waterfowl (Table 2) revealed only a few differences across three




                      time periods.   Based on aggregate percentages  described earlier, usage




                      of plant material by canvasbacits decreased  fron 71 to about 9 percent.




                      A smaller  decrease  was also suggested for the  common goldeneye




                      (Buccphala clangula americatia).  Since other apparent changes were




                      nlnor,  the incidence of plant material in samples of Chesapeake Bay




                      waterfowl  has not considerably changed during  the 1960's or 1970's.




                           There were exceptions to the general importance of vegetation to






                                                          16

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10 
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puddle ducks and animal material to diving ducks.  The diet of the black




duck, a common puddle duck in the Bay, included perhaps 25 percent




animal material whereas the redhead, a diving duck, fed almost




exclusively on vegetation.




     There were only a few species for which adequate samples were




available from the pre-1960 (historic) and 1970's (current) periods.  We




were able to compare usage of important food items within periods for




two puddle ducks (black duck and mallard) and two diving ducks




(canvasback and lesser scaup [Vythya affinis]).  For each of these




species we listed the most important food items found in samples of




these birds during the historic period (Appendix B).  We then compared




relative importance of listed food items during historic and current




periods by testing the following null hypothesis:  usage of historically




important food items by the species has not changed.




     Food items of historic importance to black ducks ranped from the




intertidal salt marsh snail (Melampus bidentatus) to emergent marsh




plants like twig-rush (Cladium mariscoides) and dotted smartweed




(Polygonum punctatum) of fresh coastal areas (Table 3).  Widgeongrass




was the only SAV species considered important to black ducks during that




period: utilization of Widgeongrass, based on relative importance of the




other food items examined, has not recently changed-  Although the




overall comparison indicated significant differences, recent changes in




relative importance were restricted to emergent marsh plants.




     Unlike black ducks, mallards are much more oriented to plant




material including emergent plants of marshes,  corn (Zea mays) of




agricultural origin,  and Widgeongrass of shallow waters (Table A).  The




relative importance of Widgeongrass was again similar (as in black





                                     18

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                    ducks) during both periods, even though the overall comparison suggested




                    recent changes for other items.  Th«'. significantly higher rank of corn




                    during the current, period suggests an increase in the incidence of field




                    feeding by mallards.




                         Although canvasbacks have always eaten invertebrates such as




                    gastropod and pelecypod mollusks, they are best known as feeders on SAV,




                    particularly wildcelery (Table 5).  Of three SAV species compared, two




                    decreased in relative inportance during the current period.  Decreases




                    were also found in current use of some invertebrates, even though there




                    was a substantial increase in the importance of the Baltic macoraa clam.




                    Small size classes (<25 mm) of these clams have recently become very




                    important, as they were found in 93 percent of 273 gizzard samples




                    (Appendix Table D-3).  Extensive use of these clams by individual




                    canvasbacks was also indicated since the aggregate percentage, which




                    measures percentage by volume, averaged 84 percent.




                         Samples from lesser scaup, another diving duck, were similarly




                    analyzed (Table 6).  Current uses (ranks)  of widgeongrass, redhp^d grass




                    (Potamogeton perfoliatus), and wildcelery, all of which were




                    historically important SAV" species, are relatively similar.  Gastropods




                    contributed most to the overall difference in relative importance among




                    the food itens examined.




                         Since current use was compared to historic use, any items that are




                    important now, but not historically, are not included here in comparative




                    food habits analyses.  Although diversity of food items currently




                    available is undoubtedly less, tnere have been relatively recent




                    additions to the Bay's flora and fauna.  Springer and Stewart (1959)




                    reported that by the late 1950's Eurasian  watermilfoil (Myr1ophyT_1un>






                                                        21

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spicatum) had become well established on the Susquehanna Flats at the


head of Chesapeake Bay, an area famous for waterfowl concentrations


during fall months.  Prior to 1960, milfoil was found only in samples of


ruddy ducks (Oxyura lamaicensis, Appendix Table B-2), green-winged teal


(Anas crecca, Appendix Table B-12), and gadwalls (_A- strepera, Appendix


Table B-J4).  During the 1960's, however, Rawls (in prep.) found milfoil


in  13 species of waterfowl (Appendix C) including American wigeon, for


which milfoil was very important (Appendix Table C-9).  Milfoil is


currently not utilized by waterfowl except in small amounts by a few


individuals.


     The brackish water clam (Rangia cuneata) is another recent member


of  the Bay's community (Pfitzenmeyer and Drobek 1964).  This clam was


not found in any samples of Bay waterfowl collected prior to 1960, but


it  has become increasingly important to diving ducks through the 1960's


and into the current period.


     Based on summaries of waterfowl food habits of 12 species, we


conclude that except for the canvasback there have been few important


changes in the incidence of plant material in samples of Chesapeake Bay


waterfowl.  Although current use of historically important food items


was significantly different within each of four waterfowl species


examined, changes in relative importance of invertebrates and emergent


marsh plants contributed substantially to those differences.  We


conclude further that canvasbacks are adaptable diving duel's, since


clams have become very important Bay food resources for these birds.


This adaptation was most likely caused by recent declines in


distribution and abundance of SAV.


     Whistling swans apparently adapted to changing Bay conditions tn



                                      24
                                                                          1 in iiim

-------
I
                  another manner.   Prior to 1970  few swans  fed  in  agricultural  fields




                  while wintering  in Maryland,  even though  this behavior was  well




                  established by swans while migrating  during spring.  Then during  the




                  late winter of 1969-70 significant numbers of swans began feeding in




                  agricultural fields (principally  corn and other  cereal grains) following




                  six weeks of freezing weather (V.  D.  Stotts,  personal communication),




                  which froze over much of the  shoal watei'  habitat in the  upper Bay.




                  Although this provided the initial stimulus,  more and more  swans  began




                  feeding in fields early (November) during subsequent winters  when SAV




                  was known to have declined.  Field feeding by swans wintering in




                  Maryland and other Flyway wintering areas has persisted  to  date (Munro,




                  in prep.).




                  DISTRIBUTION AND ABUNDANCE OF SAV




                       We first considered SAV  without  regard to species (Table 7).  SAV




                  declined considerably during  the  early 1970's.   From a high of 28-7




                  percent during 1971, frequency  occurrence of  vegetated sample stations




                  in Maryland waters declined to  21.0 percent the  following summer  (after




                  Tropical Storm Agnes).  Frequency occurrence  declined further during




                  1973 to 10.5 percent and has  essentially  fluctuated between 8-15  percent




                  thereafter.  Orth et al. (1979),  in their study  of the distribution and




                  abundance of SAV in Virginia  waters,  also found  a precipitous decline in




                  SAV during the 1970's.  Similarly, little SAV was found  during the late




                  1970's in the Potomac River (Carter and Haramis  1980), largest of the




                  Bay's numerous estuaries.




                       Based on nine-year averages,  the lower Choptank River  was the most




                  important area in the Chesapeake  Bay  for  SAV  during the  1970's.   One of




                  the largest areas, the lower  Choptank has consistently produced the most






                                                       25

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Frequency occurrence of SAV av
determined the sequanco of surv
26

-------
SAV except during 1975 when abundance was extremely low.  Another large




area, which includes the Manokin, Big and Little Annemessex Rivers, and




a portior1 of Pocomoke Sound, ranked second.  Vegetation in this area




declined after  1974 and was not found at all during the 1979 sampling.




Among the most  stable areas in the B«y for SA/ are the Chester River and




Eastern Bay.




     The exposed bays^ores, such as those along ^ent County and Kent




Island, are among the least vegetated shoal water haDitats.  Similarly,




sample stations along the Calvert County Bayshore (not listed) have not




yielded SAV during the 1970's, according to tills survey.




     Another ove-view of SAV distribution and abundance is provided in




Table 8, where  frequency occurrence is based on the presence of




vegetation in measurable amounts (see Kerwln et al. 1976).  Ranks of




survey ar»as, bastrd on average SAV abundance, are similar to those shown




in Table 7, although the distribution of SAV (the number of areas




listed) is reduced-  Frequency occurrence of SAV during the nine-year




period followed the same downward trend, but at a reduced scale-




     Widgeongrass was the nost abundant and widely distributed species




of SAV during all nine years (Appendix Table E-l).  Found in almost all




Bay tributaries due to its wide saiinity tolerance, Widgeongrass was the




most important food item for American wigeon historically (Appendix




Table B-10) as well as during the 1960's (Appendix Table C-9).  This




species of SAV, which was also the most important food item for black




ducks, is currently used to some extent by all waterfowl species




examined except for the green-winged teal.




     Widgeongrass is obviously very important to Bay waterfowl, but it




may also comprise nearly 50 percent of all the SAV in " aryland waters;






                                    27

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X* WOO— Gl ro— OO.C retflOI-V
oci-ci-<3uc«:i- xx a: > u or L. re L c
— o i- mm oxoi — o cH-aoo!
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Frequency occurrence of SAV avt
determined tho sequence cf survc
28


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it certainly is the predominant species of SAV in Maryland's raesohaline


(5-18 ppt) waters.  Orth et al. (1979) found that eelgrass (Zostera


farina) and widgeongrass were the most abundant SAV species in


mesohaline and polyhaline (18-30 ppt) waters of Virginia.


     Eelgrass was the .second most abundant species during most years


(Appendix Table E-2).  Eelgrass was found in waters of higher salinity


(>7 ppt), but as far north as Eastern Bay.  Abundance of eelgrass,  like


other SAV species, declined considerably from 1972 to 1973.  Although


this species was found at 65 sample stations in Maryland waters during


1971, it was essentially absent (1 station) during 1979.


     Eelgrass was not an important food resource for Bay waterfowl


although it was found in small amounts in 11 of 16 species examined


(Appendix B).  This species was the principal food of the Atlantic  brant


prior to the wasting disease of the early 1930's, when eelgrass and


brant populations plummeted (Lincoln 1950).  These birds, subsequently


used sea lettuce (Ulva lactuca) as their principal food while wintering


in Atlantic coastal areas.  Sea lettuce, a marine alga found commonly in


ocean bays and occasionally in Maryland waters (>7 ppt), is still the


primary food of wintering brant (Penkala l?/5).


     Redhead grass was not an important waterfowl food, although small


amounts of this item were found in 15 of 16 species examined (Appendix


B).  The distribution of redhead grass was restricted to only seven


survey areas (Appendix Table E-3).  Abundance of this species is


greatest in the Chester River and Eastern Bay,  both of which are among


the most important waterfowl wintering areas.


     Sago pondweed (Potamogeton pectlnatus), historically important to


canvasbacks (Appendix Table B-5) and redheads (Appendix Table B-9), is



                                     29

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 >

               than redhead grass.  Unlike other GAV species discussed previously, sago

 i
 [              pondweed was infrequently found when this survey began in 1971-  It was

. I
 I              also uncommon during subsequent years.

 I
 ?                   Eurasian watermilfoil was a very abundant species during the early


 |              1960's (Bayley et al.  1978).  Although known to have displaced other


               SAV species during its peak in 1963 (Elser 1967), it was an uncommon


               plant during the 1970's (Appendix Table E-5).  Milfoil was occasionally


               an important foi>d item for waterfowl (Appendix C), but perhaps only when


               preferred foods were relatively scarce.


                    Horned pondweed (Zannichellia palustris) is another SAV species


               that was uncommon in late summer during the 1970's (Appendix Table F.-61,


               and infrequently used by a number of waterfowl species.  The distri-


               bution and abundance of this SAV species were the most erratic of those


               examined.  It was essentially not recorded during four of the nine


               surveys.  This may have been caused by variation between years in the


               timing of surveys, since horned pondweed apparently dies back during


               summer (Stevenson and Confer 1978; Carter and Haramis 19PO;  L. J.


               Hindman, personal communication).


                    Wildcelery, considered an excellent waterfowl food (Martin and


               Uhler 1951), was found in the fresher reaches of only a few areas in the


               Chesapeake Bay (Appendix Table E-7).  Of the seven areas in which


               wildcelery was found, only four yielded this valuable plant after 1971.


               Based on frequency occurrence, wildcelery must now be considered an


               uncommon plant in shoal waters of the Bay. Although historically


               important to many waterfowl species (Appendix R), wildcelery is now


               infrequently found in samples from these birds (Appendix D).
    Ij                                                    30

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     We conclude that important waterfowl food plants, some of which

were very common during the late 1960's and 1971, became much less

abundant in both Maryland and Virginia waters particularly by 1973.

Frequency occurrence, as a measure of SAV abundance in Maryland,

indicated a downward trend in submerged aquatic plants as a group and

for most species.  The lower Choptank River, Manokin and Annemessex

Rivers, Chester River, and Eastern Pay were consistently among the most

important areas for a number of SAV species.

CHESAPEAKE BAY WATERFOWL POPULATIONS IN PERSPECTIVE

     Numbers of waterfowl observea during January surveys were used to

measure species abundance.  Distribution of a species was first examined

by comparing the State population to the Atlantic Flyway population,

which was also compared to the United States total.  Species

distribution and abundance among waterfowl survey areas were then

examined.  We also examined the density of birds (per unit of shoal

water habitat) in Maryland waterfowl survey areas.  In all cases we

tested a null hypothesis of no change in distribution/abundance of a

species.

     We used survey results during a previous long-term period (1956-71

in Maryland, 1961-71 in Virginia) to test for trends in distribution and

abundance of waterfowl.  These results provided a proper perspective

with which current (1972-80) waterfowl populations could be compared.

Maryland

     The Maryland portion of Chesapeake Bay has been the most important

wintering area in the Atlantic Flyway for waterfowl in general.   There

is no other area al^ng the entire Atlantic coast that was as important

to so many birds of a variety of species.  This was particularly true


                                     31

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for whistling swans, Canada geese, and canvasbacks whose Maryland

populations comprised over 50 percent of Atlantic Flyway totals (Table

9).  Maryland was also an important wintering area (>15 percent of

Flyway population) for ruddy ducks, buffleheads (Bucephala albeola),

goldeneyes, redheads, black ducks, and mallards during the long-term

period.

     The whistling swan population was one of the few that increased

during the 1956-71 period.  Numbers of swans increased significantly at

the State, Flyway, and U.S. levels.  Current populations showed no

further change in number or percentage of the Flyway population that

wintered in Maryland.  Routes of migration and wintering area site

selection of these birds are among the most traditional of all

waterfowl.  The life-long bond between swan pairs and the maintenance of

the fanily unit throughout the young's first year foster development of

these traditions.

     We examined survey area swan populations during the 1956-71 period

(Appendix Table F-l) and found a percentage distribution that was

essentially stable.  The three most important areas for swans were the

lower Choptank River, the Chester River, and Eastern Bay. Swan densities

in these areas increased significantly, and mean densities were araonR

the highest in tlie Bay.

     The Canada goose population in the Atlantic Flyway changed

dramatically during the 1956-71 period (Table 9).  This population

averaged over 500,000 geese and increased at about 25,000 birds/year,

while Maryland's population averaged over 250,000 and increased at a

similar rate.  The importance of Maryland (i.e., the percentage of the

Flyway population) increased at 2 percent/year.  The goose population


                                    32

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stabilized during the current period but at a much higher level.


Maryland now winters more than 500,000 Canada geese.   This increase,


which began during the 1950's, was caused by a wintering range expansion


of geese into agricultural lands particularly on Maryland's Eastern


Shore.


     The distribution of geese in Maryland was stable, since only a few


minor (<1 percent) changes were detected during the long-term period


(Appendix Table F-2).  Unlike other populations examined, densities of


birds increased in almost all areas-  The Chester River, where goose


populations averaged 25 percent of the State total, was the most


important area, although the highest density (>14 birds/ha) was found  ir


the Sassafras River population.


     We found minor changes in long-terra and current  Maryland


populations of ruddy ducks, scoters, buffleheads, goldeneyes, and scaup,


all of which are diving ducks and not primarily vegetarian in food


preferences-  Species distributions within Maryland (Appendix F) were


generally stable, and densities were relatively low (except for scoters


along Ca'vert County's Bayshore (Appendix Table F-4).  Important areas


varied, although the lower Choptank Fiver and bayshore areas were


frequently prominent.


     Mere than half the Atlantic Flyway canvashack population wintered


in Maryland during the 1956-71 period.  During the current period, the


Flyway population increased in number but decreased as a proportion of


the U. S. total.  Although the current Maryland population showed no


trend, it averaged about 51,000 birds which was considerably less than


the long-term average of 87,000 canvasbacks.  The percentage of the


current Flyway population that wintered in Maryland showed a definite



                                    38

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downward trend of about -3 percent/year, although the canvasback is




still the most numerous duck in Maryland.  Survey area populations of




canvasbacks (Appendix Table F-7) were stable in density and percent




distribution.  The three most important areas were the lower Choptank




River, Calvert County Bayshore, and the Patuxent River.




     The Maryland redhead population averaged about 38,000 birds prior




to 1972 with no detectable trend.  However, there was a significant




decrease (-2 percent/yjsr) in the proportion of the Flyway population




found in Maryland.  Current redhead populations, although stable,




averaged cnly ebout 6,000 birds.  This formerly abundant diving duck is




now found only in small numbers in Maryland.  Area redhead populations




(Appendix Table F-Q) were largest in the lower Choptr.nk River, although




density there trended downward significantly during the 1
-------
                      (Appendix Table F-ll), and pintails (Appendix Table F-13K  oxcept  for


                      Smith Island which was the most important pintail area.   Within-State


                      distribution of wigeon changed more than any other species  population


                      examined.  The Chester River became less important to wintering wigeon
                                *

                      at a rate of -1.3 percent/year during the 1956-7! period-


                           The Maryland mallard population currently averaged  about  26,000


                      birds which is less than the long-term average of 42,000 birds. On a


                      percentage basis, Maryland was only slightly less important to current


                      Flyway mallard populations (about 13 percent) than during the  long-term


                      (17 percent).  Mallards, which are closely related to black ducks, were


                      also most abundant in the Chester River (Appendix Table  F-12).


                      Virginia


                           Virginia populations of whistling f;wans and Canada  geese  (Table


                      10), although stable during the long-term (1961-71),  both increased


                      significantly in number and as a percentage of Atlantic  Flyway total


                      during the current period (1972-80).  Most swans in Virginia wintered in


                      areas such as ocean bays- which are beyond the scope  of  this study


                      (Appendix Table G-l).  The ir.ost important area for Canada geese was the


                      Presquile National Wildlife Refuge (NWR) on the James River, where the


                      number of geese increased significantly during the 1961-71  period

                      (Appendix Table G-2).  However, the distribution of geese in other

                      Virginia survey areas was essentially stable.


                           The ruddy duck population in Virginia averaged about 16,000 birds


                      or 30 percent of the Flyway population,  with no apparent trends during


                      the 1961-71  period (Table 10).  This population recently (1972-80)


                      averaged only 6,400 birds, an average that was stable but lower than the


                      previous long-term level.  The distribution of ruddy  ducks  in  Virginia



                                                          40
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was the most restricted of all species populations examined.  More than

half of the State population wintered in the lower Rappahannock River

area (Appendix Table C-3).

     Populations of scoters and buffleheads, although stable in previou.3

and current periods, both Increased during the latter period; canvasback

and wigeon populations, on the other hand, remained unchanged hut at low

levels (Table 10).  The lower Rappahannock River was the most Important

canvasback area within a stable State distribution (Appendix Table G-6),

whereas the lower James River was most important to wigeon (Appendix

Table G-7).

     The black duck population decreased significantly in Virginia and

the Atlantic Flyway prior to 1971, and then stabilized during the

current period.  On a proportional basis,  Virginia became more important

to Flyway black ducks at a rate of 1.4 percent/year from 1972 to 1980.

The Pamunkey River was the most important  black duck area, although a

significant downward trend in numbers of black ducks was indicated for

that area (Appendix Table G-3).

     The Virginia mallard population, which previously averaged 8,000

birds, recently increased to about 20,000  birds or 9 percent of the

Flyway total.  As with the black duck, the Pamunkey River was the most

Important area (Appendix Table G-9).

     We conclude that Maryland waterfowl populations prior to the 1970's

were considerably larger and more diverse  than those in Virginia.  While

many duck populations in Maryland decreased during the 1970's, those in

Virginia Increased or remained stable.  Maryland populations of Canada

geese and whistling swans, in contrast to  ducks, greatly increased

(geese) or remained stable (swans) .

                                    44

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                     EFFECTS OF SAV ON WATERFOVfL

                          We have deraonscrated that some Chesapeake  Bay waterfowl species

 |                  feed principally on vegetation (including SAV), while others do not or

                     §at least not to an appreciable extent.   We have also shown that, while

                     waterfowl species distributions among survey areas were  generally  stable
 1  I
   I'                  prior to the 1970's, there have been significant  changes in numbers of

   [                  birds at the State level and/or in the  importance of Maryland and  Virginia

 JB                  to Atlantic Flyway populations.  We are now able  to examine the

   |                  relationship between the distribution and abundance of waterfowl and  the
   i
   fj                  distribution and abundance of SAV in Maryland during the 1972-80 period.
   1
   |                       Although some SAV species are more important than others as

I                     waterfowl food resources, all were combined in  the analyses presented
   |
   j                  here.  As in previous analyses of waterfowl distributions,  birds
""' \
                     •surveyed in upland, inland reservoir, and ocean bay habitats were
   I
   I                 excluded.  Of the 13 waterfowl species  whose Maryland populations  were

 \B                  previously examined, 4 were excluded from consideration  here due to the

                     scarcity of their numbers during the 1972-80 period.  These are the

                     ruddy duck, redhead, wigeon, and pintail.  Maryland populations of the

                     latter three species, which feed extensively on SAV, are currently at

                     their lowest recorded levels.

                          The remaining nine populations were each run through the simple

  •                  linear model described earlier (see STATISTICAL ANALYSES),  which

    |                 examined the hypothesis that annual variation in  survey  area populations

  Jj*                 is related to annual variation in the distribution and abundance of SAV
    i
  * S                 and the general abundance of the species.  Appropriateness  of the  full

  I'
  •j                 model was not the same for the nine species populations.  We therefore
    . v
                     examined a number of alternative or reduced models,  based on preliminary
  I

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                results.  For example, if an analysis indicated that SAV had no

                measurable effect on numbers of sea ducks,  a more appropriate or


                efficient model would not include an SAV term.   For each speries, we


                removed non-significant (p > 0.05) parameters and reran the analysis.


                The analyses presented here are therefore the simplest, as they include


                only significant effects.


                Swans and Geese

  r                   Whistling swans and Canada geese in Maryland provide an interesting

                comparison.  Both are large, long-lived birds with stable or increasing


  •              populations.  Maryland is the most important wintering area in the

                Atlantic Flyway for both populations.  Although both feed in

                agricultural fields, swans have done so to  an appreciable extent only


                during the 1970's.


                     Our analysis of survey area swan populations (Table 11) indicates


  i              there is a measurable effect of SAV abundance on numbers of swans in a
  i

  i              few areas, all of which were important wintering areas during the
  i

  ;              previous long-term period (Appendix Table F-l).  For example, the effect
  t
                of SAV on swan numbers in Eastern Bay was significant (1.2 birds/ha of

                SAV).  We also found a significant intercept (1,753 birds), which


                measures the level of a survey area's population that is independent of

  ;              variation in either SAV or the general abundance of swans.  The Wye

  \              River swan population was exceptional due to the rather large, negative

  .r              relationship with SAV abundance, which was  caused by large numbers of

  ;              swans when SAV was uncommon.  This area has recently become an important


  \              loafing area for an increasing number of field-feeding swans.

                     In contrast to swans, there is no relationship (p > 0.05) between


  \              distribution and abundance of Canada geese  and  SAV in Maryland (Table


                                                     46


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12).  Numbers of geese in survey areas are entirely explained by

variation in the Atlantic Flyway population.  In fact, populations of

geese on Maryland's Eastern shore may be responsible for much of the

variation in the Atlantic Flyway total.  Open water areas are now used

mostly for loafing and protection from predators, as geese have become

essentially dependant on surrounding agricultural lands for food.

Black Ducks and Mallards

     These species have been and continue to be the most common and

widely distributed puddle ducks in Maryland, although current

populations of both are considerably less than long-term levels.  We

found that black duck numbers in the Chester River were significantly

related to the abundance of SAV (about 3 birds/ha of SAV, Table 13).

This area was also the most important wintering area for black ducks

during the 1956-71 period (Appendix Table F-l1).  Black ducks in the

Honga River and Bloodsworth-South Marsh Islands areas also varied with

SAV, but at lesser rates (about 0.6 birds/ha of SAV).  Populations in a

number of other areas varied with the general abundance of the species-

     Mallard numbers in the Chester River were also related to the.

abundance of SAV (about 2.7 birds/ha of SAV, Table 14).  However, bird

numbers in other areas were not similarly affected.  Although mallards

are found in all survey areas, most of the variation in-population size

was related to the general abundance of mallards in a few areas, except

in the Chester River.

Diving Ducks

     Although plant material comprised only 25-30 percent of the food

habits of buffleheads, we found that numbers of these birds in the

Manokin, Anneme^sex Rivers and Eastern Bay areas were related to the


                                    48

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abundance of SAV (Table 15).  The relationship (0.3 birds/ha of SAV) was

less pronounced than in the puddle ducks examined, but more important

than general species abundance in these areas.  Population variation in

other important bufflehead areas, such as the lower Choptank and Little

Chcptank Rivers, simply reflected changes in general abundance.

     We demonstrated earlier that the food habits of canvasbacks in

Chesapeake Bay changed during the 1970's nore than any other species

examined.  Canvasbacks now feed extensively on small (<25 mm) Baltic

macoma clams and other invertebrates and infrequently on SAV.  When we

analyzed current distribution and abundance of canvasbacks, we found no

relationship with distribution and abundance of SAV (Table 16).  We did

find a significant intercept (1,910 birds), which is common to all

areas, that estimates a population level independent of SAV or species

abundance.  If the area-specific effect of general abundance is not

significant, :hen that area's population is estimated by the intercept.

However if it is significant, the product of the area's parameter

estimate ana the Flyway population in a given year is added to the

intercept.  Estimated canvasback populations in areas listed under

"Flyway x Survey area" in Table 16 are therefore greater or less than

1,910 birds.

     Maryland populations of scaup and common golJeneyes also were not

related to the distribution and abundance of SAV.   Interpretation of our

analysis of scaup populations (Table 17) is complicated somewhat by the

large, negative intercept (-2,551 birds).  This was primarily caused by

extrapolating beyond the range of the data.   Although area populations

should be small during years when scaup are relatively scarce,  a

straight line fit yielded a negative intercept.  However,  a model that


                                    52

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includes an intercept (if significant) better explains the variation in


area populations than does one without an intercept.  Scaup populations


in areas not listed in Table 17 are essentially very low, or at least


not affected by general species abundance.


     Although our analysis of common goldeneye populations led to the


same conclusion (i.e., no SAV relationship), interpretation of results


shown in Table 18 is simpler.  Goldeneyes are uncommon in most survey


areas, but relatively common and responsive to general abundance in


areas such as Eastern Bay and the lower Choptank River-


     We examined scoter or sea duck populations in Maryland witli the


same set of models, since these birds contrast greatly with -tho.i


waterfowl in food and habitat preferences.  As anticipated, we found no


relationship between distribution snd abundance of SAV and scoters.


Small populations of these birds are widely scattered around the Bay


(significant intercept), and concentrated in only a few areas such as


the lower Choptank and Chester Rivers (Table 19).


     We conclude that populations of wigeon, pintails, and redheads, all


predominantly vegetarian in food preferences, declined to very low


numbers in Maryland during the 1970's as a direct result of the SAV


decline.  Although swan populations showed a response to the level of


SAV in a few important areas, black ducks and mallards were generally


more affected.  Diving ducks as a group showed little relationship if


any to the distribution and abundance of SAV.


     Jhe distribution of waterfowl populations among survey areas


indicates that a few areas are consistently among the most important to


the various species examined.  The Chester River, Eastern Bay, and lower


Choptank River, which are among the four most abundantly vegetated


                                    56

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areas, are equally important as waterfowl wintering areas.  We interpret


this as additional evidence of a relationship between the distribution


of waterfowl and SAV.


IMPACTS OF WATERFOWL ON SAV


     Prior to this point we have attempted to clarify relationships


between SAV as food resources and waterfowl us consumers.  Our objective


here is to evaluate the possibility that Chesapeake Bay waterfowl


populations are in any way responsible for recent declines in


distribution and abundance of submerged aquatic vegetation.


     We are reminded, first of all, that waterfowl wintered in
                   Chesapeake  Bay long before  recorded history.  Populations of these
birds, which have consumed SAV in the Bay for centuries without adverse


impact, can be examined only during the last 25-30 years.  Maryland


waterfowl populations with one exception were larger during the early


1950's when SAV was certainly more abundant as well.  The Canada goose


population was the exception, as it began to increase during the


mid-1950's when the mechanization of agriculture increased the


availability of unharvested cereal grains.


     We showed that distributions of waterfowl species populations in


Maryland were generally stable prior to the 1970's.  We showed further


that three areas, the Chester River, Eastern Bay, and the lower Choptan'c


River, consistently wintered the highest proportions of Maryland's


waterfowl populations during both long-term and current periods.  We


conclude that 1) these populations most likely reflect greater abundance


of SAV in these areas compared to other areas, and 2) since these areas


continue to winter most of the waterfowl and produce most of the SAV,


these birds cannot be responsible for recent declines in their food


resources.
                                    59

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                               REFERENCES

Barr, A. J., J. H. Goodnight, J. P. Sell, and J. T. Helwig.  1976.  A
     user's guide to SAS 76.  SAS Inst. Inc.. Raleigh, N.C. 329 pp.

Bayley, S., V. D. Stotts, P. F. Springer, and J. Steenis.  1978.
     Changes in submerged aquatic macrophyte populations at the head of
     the Chesapeake Bay, 1958-1975.  Estuaries 1(3): 171-182.

Bellrose, F. C. 1976.  Ducks, geese and swans of North America.
     Stackpols Books, Harrisburg, Pa. 543 pp.

Benning, D. S., S. L. Rhoades, L. D. Schroeder, and M. M. Smith, eds.
     1978.  Waterfowl status report, 1974.  U.S. Fish Wildl. Serv.
     Spec. Sci. Rep.-Wildl. 211.  98 pp.

Bent, A. C. 1923.  Life histories of North American wild fowl.  Order:
     Anseres (Part I).  U.S. Natl. Mus. Bull. 126.  Washington, D.C.
     244 pp.

Carter, V., and G. M. Haramis.  1980.  Distribution and abundance of
     submersed aquatic vegetation in the  idal Potonac River -
     implications for waterfowl.  Atl. N, > iralist 33:14-19.

Cottam, C.  1939-  Food habits of North American diving ducks.  U.S.
     Dept. Agric. Tech. Bull. 643.  i40 p, .

Elser, H. J.  1967.  Status of aquatic weed  problems in tidewater
     Maryland, spring 1967.  Manatee project. Md. Dept. Ches. Bay
     Affairs, Annapolis.  (11 pp. mimeo)

Johnson, D. Ti. I960.  The comparison of usage and availability
     measurements for evaluating resource preference.  Ecology
Kerwin, J. A., R. E. Munro, and W. W. A. Peterson.  1976.  Distribution
     and abundance of aquatic vegetation in the upper Chesapeake Bay,
     1971-1974, p. 393-400 jln J. Davis, coord.  The effects of tropical
     storm Agnes on the Chesapeake Bay estuarine system.  Chesapeake
     Res. Cons. Publ. No. 54, The Johns Hopkins Univ. Press,
     Baltimore, Md. 639 pp.

Korschgen, L. J. 1969.  Procedures for food-habits analyses, p. 233-250
     in R. H. Giles, Jr., ed.  Wildlife management techniques.  Wildlife
     Society, Washington, D.C. 623 pp.

Lincoln, F. C. 1950.  The American brant - living bird or museum piece.
     Audubon Mag. 52:282-287.

Martlr-. A. C. and F. M. Uhler.  1951.  Food of ga*e ducks in the United
     States and (,anada.  Res. Rep. 30, U.S. Fish Wildl. Serv.  308 pp.

Martin, A. C., R. H. Gensch, and C. P. Brown.  1946.  Alternative
     methods in upland gamebird food analysis.  J. Wildl. Manage.
           3-12.

                                   60

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  f
  I
  I
  I
  I
   I
Munro, R. E. (in prep.)-  Field feeding by whistling swans in Maryland.
     Proc Second Int. Swan Symp.

Orth, R. J., K. A. Moore, and H. H. Gordon.  1979.  Distribution and
     abundance of submerged aquatic vegetation in the lower Chesapeake
     Bay, Virginia.  EPA Report No. 600/8-79-029/SAV1.  199 pp.

Penkala, J. M. 1975.  Winter food habits and body weight of Atlantic
     Brant.  Trans. N.E. Fish Wildl. Conf. 32:151-169.

Perry, M. C., and F. M. Uhler.  1976.  Availability and utilization of
     Canvasback food organisms in Chesapeake Bay.  Presented at Spring
     Meeting of the Atlantic Estuarine Research Society, Rehoboth Beach,
     Del. (25 pp. mitneo)

Pfitzenmeyer, H. T., and K. G. Drobek.  1964.  The occurrence of the
     brackish water clam, jtangia cuneata, in the Potomac River,
     Maryland.  Ches. Sci. 5(4):209-215.

Rawls, C. K. (in prep.).  Food habits of waterfowl in the upper
     Chesapeake Bay, Maryland.  Univ. of Maryland Center for
     Environmental and Estuarine Studies.  140 pp.

Springer, P. F., and R. E. Stewart.  1959.  Condition of waterfowl
     feeding grounds on the Susquehanna Flats during the fall of 1959
     with notes on the invasion of a serious pest plant.  U.S.  Fish
     Wildl. Serv., Patuxent Wildl. Res. Ctr., Laurel, Md.  (7 pp.
     mimeo)

Stevenson, J. C., and N. M. Confer.  1978.  Summary of available
     information on Chesapeake Bay submerged vegetation.  Final report
     to FWS/OBS-78/66.  335 pp.

Stewart, R. E. 1962.  Waterfowl populations in the upper Chesapeake Bay.
     U.S. Fish Wildl. Serv. Spec. Sci. Rep.-Wildl. 65.  208 pp.

Swanson, G. A., and J. C. Bartonek.  1970.  Bias associated with food
     analysis in gizzards of blue-winged teal.  J. Wildl. Manage-
     34(4):739-746.

Swanson, G. A., G. L. Krapu, J. C. Bartonek, J. R. Serie, and D. H.
     Johnson.  1974.  Advantages in mathematically weighting waterfowl
     food habits data.  J. Wildl. Manage. 38(2):302-307.

Yocom, C. F. 1951.  Waterfowl and their food plants in Washington.
     Univ. of Washington Press, Seattle, Wash.  272 pp.
    .

'1
                                                       61

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