Patuxent's American Black Duck Studies from Chesapeake Bay To

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2016 Patuxent’s American Black Duck Studies from Chesapeake Bay to Maine and Beyond Jerry R. Longcore

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Longcore, Jerry R., "Patuxent’s American Black Duck Studies from Chesapeake Bay to Maine and Beyond" (2016). USGS Staff -- Published Research. 1031. https://digitalcommons.unl.edu/usgsstaffpub/1031

This Article is brought to you for free and open access by the US Geological Survey at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in USGS Staff -- ubP lished Research by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. decoys referred to as“call” ducks (Perry, 1984)were used,and p. 10).Baiting of duckswasallowed(Leopold, 1931),live ally with8-or10-gagedouble-barreled shotguns(Day, 1949, ing howmanysittingduckscould bekilledwithoneshot,usu allowed during springmigration, andhunterstook sport insee large dailybaglimits of75birdsinthe1920s.Shootingwas nonexistent orextremelyliberal, with107-dayseasonsand tions governinghuntingofwaterfowlwerehistoricallynearly Chase, 1950),andthroughoutitsrangeinCanada.Regula 2003), inlandthroughouttheMississippiFlyway(Bellrose and gunners alongthe Atlantic Flyway (Wright, 1947;Sullivan, p. 164). This specieshasbeenafavoritetarget ofcoastal sagacious, waryandwildestofallducks”(Kortright,1942, throughout itsrangebywaterfowlers,isregaledas“themost Background from fertilewetlands. ( changes, andthenowdiscreditedhypothesisthatmallard ecology, analysesofbandingandsurveydatapopulation papers thatcoverfourresearchtopics:chemicalcontaminants, Literature citationsareselective,butincluderepresentative essential tothediscussionofblackduckstudiesisincluded. ( influenced researchatPatuxentonthe Americanblackduck and Stateagenciesprivateacademicinstitutionsthat at Patuxent.LiteratureresultingfromworkotherFederal prepared byUSFWSscientistswhiletheywerenotworking administered bytheseagencies.Someofthecitedreportswere the researchentitycurrentlyknownasPatuxentwasformerly Wildlife Service(USFWS)personnelareincludedbecause Biological Survey(BiologicalSurvey)andU.S.Fish (Patuxent), duringits75-yearhistory. ReportsbyBureauof U.S. GeologicalSurvey, Patuxent Wildlife ResearchCenter ture andunpublishedreportswrittenbystaff membersofthe Introduction By JerryR.Longcore Maine Beyond and to Bay Chesapeake Black American Patuxent’s Duck from Studies Anas platyrhynchos Anas rubripes The blackduck,asportinggameduckardentlysought The informationinthischapterdrawsonpublishedlitera , hereafterreferredtoasblackduck)andthatis ) couldcompetitivelyexcludeblackducks - - - - Interior (DOI). and Wildlife Serviceandtransferred totheDepartmentof the of SportFisheriesand Wildlife, laterrenamedtheU.S.Fish coln. The Biological SurveywastheforerunnerofBureau started abird-bandingprogram headedbyFrederickC.Lin Survey, whichwasinitsinfancy in1920(Hawkins,1984), 1913) andbirdmigration(Cooke, 1906). The Biological although theydidpublishonfoodsofwaterfowl(McAtee, tive economiceffects ofavian species onagriculturalcrops, 1984). Scientistsatthattimefocusedonrecordingthenega 1886 andlocatedintheU.S.Departmentof Agriculture (Perry, of EconomicOrnithology, formedbyan Act ofCongressin Biological Survey, which evolvedin1896fromtheDivision ment wasnotaresearchpriorityduringtheearlyyearsof (Hornaday, 1913). way States,andthePacificCoastespeciallyCalifornia areas werealongthe Atlantic Coast,LowerMississippiFly with impunity(Buckingham,1937). The greatmarkethunting killing ofduckstosellinthemarketslarge cities occurred 1992. PhotographbyMatthewC.Perry, U.S.FishandWildlifeService. Captive blackducksatPatuxentWildlifeResearchCenter, Laurel,MD,March Studying theecologyofblackducksandtheirmanage - - - -

Migratory Bird Research, Monitoring, and Management 70 The History of Patuxent: America’s Wildlife Research Story

1930s of life he spends in reaching the breeding ground and producing the annual crop, but in the gauntlet The Patuxent Research Refuge was established in 1936 of gun-fire he faces from southern Canada to the by Executive Order 7514 as part of the U.S. Department of wintering ground and on the wintering ground. Agriculture. In 1939, the Bureau of Fisheries and the Biologi- The gradual increase in hunting pressure together cal Survey were consolidated into one agency and, in 1940, it with the dying off of his favourite winter food, the was transferred to DOI to form the Fish and Wildlife Service eelgrass, and the reduction of winter range caused (FWS). In 1956, the FWS was divided into the Bureau of by the steady building up of the human population Sport Fisheries and Wildlife (BSFW) and the Bureau of Com- with its attendant demand for mosquito-free summer mercial Fisheries, and the FWS became the U.S. Fish and cottages along the Atlantic seaboard, has reduced Wildlife Service. In 1970, the Bureau of Commercial Fisheries the species to the point where it is impossible, in the was transferred back to the Department of Commerce and the east, to find only one duck of any kind in 14 acres BSFW designation was discontinued. of marsh where they were once found in sufficient The name “Patuxent Research Refuge” was changed number that they could be secured with a club.” to “Patuxent Wildlife Research Center” in 1956. Scientists during the earliest years of Patuxent Research Refuge pur- sued work that had been begun in the Biological Survey days, 1950s mainly exploring the mystery of bird migration (Cooke, 1915; Lincoln, 1935), that led to the concept of biological flyways of During this period, Stewart (1958) published distribution birds as espoused by F.C. Lincoln (Hawkins, 1984), and iden- maps for breeding and wintering black duck populations, and tifying foods of waterfowl (Cottam, 1939; Martin and Uhler, Addy (1953) reported on the fall migration of the black duck. 1939). During this time, concern for the future of diminishing In the mid-1950s, the USFWS initiated a series of mid-winter stocks of waterfowl was acknowledged. Earlier, Cooke (1906, surveys in cooperation with States in the Atlantic and Mis- p. 10) had stated, “The principal causes of the diminished sissippi Flyways to inventory waterfowl. These mid-winter numbers of waterfowl have been market hunting, spring shoot- inventory (MWI) data indicated a total black duck population ing, and the destruction of the breeding ground for farming of 500,000 to 600,000, but this number was declining about purposes.” Waterfowlers on Chesapeake Bay during the “days 2 percent annually (Serie, 1997, p. 14). of plenty” shot from the deadly sinkbox in the 1800s; from 1870 to 1875, it was not uncommon for 15,000 ducks to be killed on Chesapeake Bay in a single day (Sullivan, 2003). A report about gunning on the Eastern Shore of Maryland 1960s described the use of corn bait and unplugged guns, the ship- ping of ducks to markets in Baltimore, and the use of live During the 1960s, an evaluation of the role of chemical decoys, but stated that “The activities of the Biological Survey contaminants in the decline of the black duck was initiated by men have been such as to make the natives take precautions” analyzing for pesticides in eggs (Reichel and Addy, 1968) and (National Association of Audubon Societies, 1937). wings (Heath and Prouty, 1967; Heath, 1969). Several contaminants, especially dichlorodiphenyltrichloroethane (DDT) and its metabolites, were detected in eggs and wings, which prompted experimental pen studies in the early 1970s to deter- 1940s mine if and how DDT affected reproduction. Stewart (1962) analyzed 1953–59 MWI data and described waterfowl popula- During the next decade, Ira Gabrielson (1947) sounded a tions, including that of the black duck, in the Upper Chesa- call to address the declining black duck population, stating that peake Bay region. Lucille Stickel edited Stewart’s 208-page the “program should be accompanied by restrictions on shoot- manuscript, and several Patuxent staff members (Francis M. ing sufficient to limit kill to less than the annual number of Uhler, Alexander Martin, Neil Hotchkiss, and Robert Mitchell) ducks put on the wing.” Cottam (1948) addressed the causes assisted in identifying foods of waterfowl sampled in Chesa- of the waterfowl crisis as “destruction of habitat,” “subnormal peake Bay. Chuck Kaczynski and Jake Chamberlain (1968) production,” and “overshooting.” In this period, studies of reported the number of black ducks counted during aerial black ducks by State biologists, especially in Massachusetts, surveys in eastern Canada. John Sincock (1962) estimated the were initiated. Wright (1947, p. 138–139) reported his find- amounts of food consumed by waterfowl, including the black ings on the black duck in eastern Canada in a progress report duck, in Back Bay, Virginia/Currituck Sound, NC. to the Chief Naturalist of Ducks Unlimited and concluded Atlantic Flyway representatives, who were trained biolo- the following: gists, supported black duck research studies, surveys, and “The evidence therefore indicates that all is not well banding projects. In 1967, the Atlantic Flyway Council, Tech- with the black duck of the Atlantic Flyway, and that nical Section, created a Black Duck Committee (Serie, 2002); the trouble is probably not to be found in the part its first action was to organize a Black Duck Symposium in tion. Monitoring oforganochlorine residuesandmercuryin to thehypothesis thatchemicalswereaffecting reproduc was curtailed(Longcoreand Stendell, 1977),addingcredence by DDEpersistedintothenext year, evenafterthedosage cracked shellsinnests.Negative reproductiveeffects caused ing femaleswasdecreasingbecause ofthelosseggswith This findingconfirmedthat the productivityofsomebreed when femaleswereallowedtoincubatetheirownclutches. Samson (1973)reportedafourfoldincreaseinshellcracking collected in1968(Reicheland Addy, 1968).Longcoreand sive shellthinningintheeggsexaminedcomparedtothose egg shells(Longcoreandothers,1971)documentedexten nyldichloroethylene (DDE)onthethicknessofblackduck 1979). Experimentalstudiesoftheeffects ofdichlorodiphe 1973) andasurveyofleadinwingbones(Stendellothers, contaminants inblackduckeggs(LongcoreandMulhern, concentrations ofpolychlorinatedbiphenyl(PCB)andDDT 1970s of thedeclinefornext30years. controversy thatdelayedtheconfirmationofactualcauses have confoundedblackduckpopulationstudiesandfostered decline inthenumberofblackducks.Suchspeculationmay a speculativeviewthatmallardscouldbethecauseof near future. This paperandother, similarreportsputforward although suchadevelopmentwasconsideredunlikelyinthe pear asadistinctentity through hybridization with themallard, was smallerthanthatofthemallard)couldeventuallydisap raised thepossibilitythatblackduck(whosegenepool in fact,wasdelayedforyears.Inaddition,Johnsgard(1967) harvest. This opportunity, however, wasnotembracedand, provided auniqueopportunitytoimplementplancurtail ducks wasaffecting theblackduckpopulation. This consensus can andCanadianpersonnelagreedthattheharvestofblack Canada andtheU.S.”(Wilder, 1968). At thistime, Ameri legal takeofblackducksshouldconsiderthesituationinboth it” (Munro,1968);and“Anyrationalattempttoreducethe used becausetherearenotenoughblackduckstooccupy gists areoftheopinionthatnotallavailablehabitatisbeing hunting mortality”(Loughrey, 1968);“MostCanadianbiolo 1968); “We needmorequantitative informationaboutnon- to bringcontrollablekillinlinewithproduction…”(Wilder, seems obviousthatmeasuresneedtobetakenimmediately ments madeinthesymposiumproceedingsincluded,“…it control ofpredatorsandcompetitors(McGilvrey, 1968).Com berlain, 1968),environmentalpollution(Stickel,and dynamics (Martinsonandothers,1968),aerialsurveys(Cham contributed papersontopicssuchasharvestandpopulation tify theneedsofblackduck.SeveralPatuxentscientists administrators toreviewknowninformationaboutandiden which broughttogether American andCanadianbiologists vided ageneralreviewofblackduckstatusatthesymposium, Chestertown, MD(Barske,1968).C.E. Addy (1968,p.2)pro This decadebroughtadditionalsurveystodocument Patuxent’s AmericanBlackDuckStudiesfromChesapeakeBaytoMaineandBeyond ------Photo byMatthewC.Perry, U.S.FishandWildlifeService. shells, DDEstudy, PatuxentWildlifeResearchCenter, Laurel,MD,spring1972. Jerry Longcore,U.S.FishandWildlifeService,checkingeggsforcracked Research Laboratory, whichhad beenestablishedin1972, the fall,only39returnedtonest thenextspring. black ducksimprintedtothese cylindersandthenreleasedin equipped withapredatorguard. Of169captive-rearedfemale duck femalestoelevatednest cylindersonasupportpost affect thepopulation,McGilvrey(1971) conditionedblack 1976). Becausenestlossofground-nestingblackduckscould study werelatershowntobeinvalid(AndersonandBurnham, survival rate;however, thestatisticalmethodsusedinthis affect hunting mortalityrate,whichinturnaffects theannual harvest surveysandconcludedthathuntingregulations and Stendell, 1978). fed 3partspermillionofmethylmercuryover2years(Finley egg hatchabilityandlowerducklingsurvivalincaptiveducks duck survivalandreproductionwereshowntoincludereduced Heath, 1976; White, 1979),andeffects ofmercuryonblack black duckwingscontinued(HeathandHill,1974; White and Wildlife Service. Center, Laurel,MD,spring1972.PhotobyMatthewC.Perry, U.S.Fishand American blackduckfemaleandbrood,DDEstudy, PatuxentWildlifeResearch After joiningthe PatuxentMigratoryBirdand Habitat Geis andothers(1971)analyzeddatafromseveral 71

Migratory Bird Research, Monitoring, and Management 72 The History of Patuxent: America’s Wildlife Research Story

Reinecke (1979) reported on the important foods, growth, and ducks caused loss of shell thickness and mass (Longcore and development of juvenile Maine black ducks. Hunting mortal- Stendell, 1982), but by 1978, the thickness of black duck ity typically was considered to be compensatory to other forms eggshells had recovered to a pre-1946 mean (Haseltine and of mortality (Nichols and others, 1984), but the concept of a others, 1980). This discovery lessened the probability that threshold of additivity of hunting losses emerged as Anderson chemicals were decreasing productivity and contributing to the and Burnham (1976, p. 41) stressed that “Whatever this point population decline, but monitoring of organochlorine pesticide is, it may be easy to exceed it on the breeding grounds or on residues in black duck wings continued (Cain, 1981; Prouty areas where the birds may be particularly vulnerable (Jessen, and Bunck, 1986; Hall and others, 1989). Heinz and Hasel- 1970). Harvest rates early in the season on adult females and tine (1981) documented that chromium added to the diet of young on breeding and staging areas could be severe.” young black ducks affected their avoidance behavior; simi- In 1976, with an increased commitment to developing an lar effects were determined for cadmium (Heinz and others, understanding of the variables affecting the black duck, Patux- 1983). Differential susceptibility to lead poisoning between ent sent me to Maine to investigate the breeding ecology of the the black duck and the mallard was suggested as a possible species. At the same time, Patuxent biologist Dr. Ronald Kirby cause of declines in the number of black ducks (Chasko and was assigned to investigate aspects of wintering ecology of others, 1984). Rattner and others (1989) refuted the hypothesis black ducks along the Atlantic Coast, focusing on Chesapeake that the black duck was more sensitive to lead poisoning than Bay and New Jersey. Implications about the role of the mal- the mallard by documenting the absence of any difference in lard in the black duck population decline persisted as Johns- mortality between these species on the same lead pellet dosage gard and DiSilvestro (1976) suggested that “.…the relatively and diet. specialized black duck, through increased competition and The effects of acidic deposition on wetland inverte- hybridization with the much more broadly adaptable mallard, brates raised concern that growth and survival of black duck will continue to become an increasingly rarer [sic] component ducklings could be negatively affected. The role of wetland of the North American bird fauna.” It seemed to some of us acidification on captive black ducks was evaluated at Patux- field biologists studying the black duck, however, that “There ent with constructed ponds that were experimentally acidified is always an easy solution for every human problem—neat, by Haramis and Chu (1987) and Rattner and others (1987), plausible and wrong” (Mencken, 1917). whose findings indicated lower invertebrate food production on acidic ponds and possible adverse effects on ducklings. In subsequent field studies, Longcore and others (2006) reported that black duck broods readily used low-pH wetlands with 1980s good survival of ducklings. Kirby (1988) reviewed enhancement of black duck breed- Black duck conservation and management during this ing habitat in the northeastern United States, and Jorde and decade benefited from establishment of a Black Duck Com- others (1989) compiled information on existing tidal and non- mittee by the Atlantic Flyway Council, which was chaired by tidal wetlands of the northern Atlantic States. Results of sev- H.E. Howard Spencer, Jr. (Spencer, 1980). This committee eral studies on breeding ecology and survival of black ducks compiled a Black Duck Management Plan for North America were published by Patuxent scientists and associated students. 1980–2000 with data provided by personnel of Provincial, Longcore and Ringelman (1980) determined variables affect- Federal, and State agencies; organizations; and private citi- ing breeding densities in the Northeast and developed a black zens. Black duck conservation benefited further from formal duck population model through use of computer simulations establishment of the North American Waterfowl Management (Ringelman and Longcore, 1980). Results of telemetry used Plan (NAWMP) (U.S. Fish and Wildlife Service, 1986) and on breeding pairs of black ducks in Maine revealed move- from increased research, including an array of field studies by ments and wetland selection by brood-rearing black ducks several Patuxent scientists. The NAWMP was signed by the (Ringelman and Longcore, 1982a), survival of broods to fledg- governments of the United States and Canada in 1986 (Serie, ing (Ringelman and Longcore, 1982b), habitat types selected 1997), and the plan identified the black duck as a “species of and sizes of home ranges of males and females (Ringelman international concern.” Under the plan, the Black Duck Joint and others, 1982a), nest and brood attentiveness of females Venture (BDJV) was formed and implemented in 1990 to (Ringelman and others, 1982b), and survival of females coordinate data gathering for population surveys, banding, and (Ringelman and Longcore, 1983). Krementz and others (1987) research. A winter population goal was set at 385,000 black determined sources of variation in survival and recovery rates ducks. A technical committee established within the BDJV, in black ducks, wherein more adults than hatch-year ducks composed of American and Canadian biologists, reviewed pro- survived and more adult males than adult females survived. posed survey, banding, and research projects, thereby improv- Survival rates were similar for young of both genders, but ing the quality of data collected. the recovery rate was greater for young males than for young Patuxent continued its research on exposure to con- females. Although recovery rates were time dependent, taminants and their effects on black ducks. A minute amount survival rates were not, which indicates that some variations (3 parts per million, dry weight) of DDE in the diet of black in mortality caused by hunters may be compensated for by (1985) regarding effects ofhunting onsurvivalratesofblack lenged thevalidity ofinferencesmadebyBoyd andHyslop bands arenotreported.Conroy andKrementz(1986)chal may overestimatethereporting ratebecausesomereward the optimumestimatewasaconstant 0.43,althoughthisvalue poral differences inbandreportingrates forblackducks,but Conroy andBlandin(1984)identified geographicandtem of testsfordifferences inannualsurvivalratewere equivocal. of sympatricblackducksandmallardsweresimilarresults Nichols andothers(1987)determinedthatbandrecoveryrates experiments tounderstandblackduckpopulationdynamics. losses, and Anderson andothers (1987)advocatedtheuseof reviewed evidenceforcompensatorymortalityinwaterfowl excessive harvestofblackducks.Nicholsandothers(1984) agement,” andattributedthelong-termpopulationdecline to as “acaseof28yearsfailurein American wildlifeman mately 1.5percentannuallyinthe1970sand 1980s. noted thataveragedeclineinthepopulationwasapproxi reviewed blackduckpopulationstatusandmanagement as adultmalesandjuveniles).RogersPatterson(1984) additive effect) onsurvivalofsomeageorsexclasses(such ferent regulatoryschemesresultedindirecteffects (thatis,an others (1988)reportedthatchangesinharvestrateunderdif the effects ofhuntingonblackducksurvival,Krementzand to annualsurvival(Krementzandothers,1989).Instudiesof other causes.Bodymassinwinterwasnotpositivelyrelated and WildlifeService. of blackducks,AtlanticCity, NJ,1982.PhotobyMatthewC.Perry, U.S.Fish Dan StottsandMikeConroy, U.S.FishandWildlifeService,recordingweight Grandy (1983)referredtomanagementoftheblackduck Patuxent’s AmericanBlackDuckStudiesfromChesapeakeBaytoMaineandBeyond - - - - - areas (15,019 acres)intherelativeabsenceofmallardbroods were decliningsubstantiallystatewideon36indexwetland and foundfewrecords,butnumbersofblackduckbroods mallard interactionsasnotedinliteraturerelatedtoMaine black duck.Longcoreandothers(1987)evaluatedduck- rized informationonthepopulationstatusandharvestof roosted intheleeofislands.Ruschandothers(1989)summa Maine coastduringtheseverewinterof1980–81,whenducks Gibbs (1988)identifiedcriticalhabitatforblackducksonthe adult birds(Diefenbachandothers,1988b).Longcore but nodifferences indistributionpatternsexistedbetween Young blackduckswinterednortheastofyoungmallards, a strongerfidelitytocoastalwinteringsitesthaninlandsites. distributions ofwinteringpopulationsblackducksthathad was ausefulindex.Diefenbachandothers(1988a)identified for theMWIofblackducksandconcludedthatsurvey ducks thathadalowerbodymass. as 0.73forafter-hatch-year ducksand0.60forhatch-year survival ratesforfemaleblackducksalongthe Atlantic Coast ducks. Conroyandothers(1989b)determinedmeanwinter ment strategies intheUnitedStates,andSerie andBailey (1997) informed onpopulationstatusandharvest manage was heldatthebeginningofthis decade.Serieandothers 1990s to belosingsupport. exclude blackducksfromfertilehabitats,however, appeared duck decline. The beliefthatmallardscouldcompetitively to defendtheirpositionontheroleofmallardinblack and decreasingblackducks.” Ankney andothers (1989) tried “cause andeffect” forthehypothesisof“increasingmallards others (1989a),whocommentedthatnoevidenceexistedfor this assertionwerelacking,however, asnotedbyConroyand were declininginsomepartsofitsrange.Datatosupport the expenseofblackduckpopulation,whosenumbers mallards inOntarioandQuebec,Canada,wasincreasingat (table 1). Mallard Black duck [Modified fromLongcoreandothers,1987] 36 index wetlandsinMaine,1956–86. Table 1. Species Conroy andothers(1988)evaluatedtheaerialtransects The secondBlackDuckSymposium (Kehoe,1997) Ankney andothers(1987)impliedthatthenumberof

Numbers ofblackduckandmallardbroodson 1956–65 457 2 1966–76 Years 328 5 1977–86 178 18 - 73 -

Migratory Bird Research, Monitoring, and Management 74 The History of Patuxent: America’s Wildlife Research Story

(1997) discussed implementation of the BDJV. Longcore and Carney (1992) developed keys to identify species of Ringelman (1997) reported that, although the area occupied wings submitted during harvest surveys, which facilitated by surface water increased in a 58-square-mile area in south- estimating harvest of black ducks by hunters. Conroy and central Maine, the number of pairs and broods of black ducks Krementz (1990) reviewed existing evidence that hunting was decreased from 1958–60 to 1978–80. affecting the black duck population and discussed the biologi- In a study of the effect of acid precipitation on the qual- cal basis of compensatory as opposed to additive mortality. ity of invertebrate food eaten by the black duck, Sparling Blandin (1992) determined population characteristics of black (1990) evaluated the effects of dietary aluminum, calcium, and ducks through simulation modeling. Nichols (1991) pre- phosphorus on the growth and survival of captive black ducks sented an in-depth review of science, population ecology, and and mallards. Black ducks seemed more sensitive than mal- management of black ducks and reported that the statistical lards to treatments low in calcium and phosphorus and high in methods used in earlier papers had been inappropriate, thereby aluminum. Effects of these diets on bone and liver characteris- invalidating their conclusions. Clugston and others (1994) tics of these species were similar (Sparling, 1991). Frazer and documented the effect of hunter kills related to habitat use others (1990a, 1990b) evaluated home range, movements, and for immature female black ducks at Escoumins, Quebec, in habitat use of post-fledging black ducks in Maine and New 1991. The sample of radiomarked ducks was divided into three Brunswick. Krementz and others (1991) documented histori- groups on the basis of the percentage of times (that is, telem- cal changes in egg-laying date, clutch size, and nest success of etry locations) recorded in the St. Lawrence Estuary (table 2). black ducks in Chesapeake Bay and compared the productiv- Most hunting took place in the estuary, so most ducks ity of the black duck to that of the mallard, which was similar that avoided the estuary survived. These findings support (Krementz and others, 1992). the concept of additivity of hunting losses on breeding and Black duck breeding ranges have been decreasing across staging areas described by Anderson and Burnham (1976, the Bird Conservation Regions of Boreal Hardwood Transi- p. 41), who concluded the “threshold” of additivity of hunt- tion and the Great Lakes/St. Lawrence Plain throughout the ing losses “may be easy to exceed on the breeding grounds,” second half of the 20th century (Pendleton and Sauer, 1992). whatever that point might be. Kitchens (1994) determined that Krementz and Pendleton (1991) recorded the movements and opening of hunting seasons disrupted use of prime feeding survival of black duck and mallard ducklings on Chesapeake habitats in Missisquoi Bay in Vermont and Quebec, but use Bay with implanted transmitters and found no differences in resumed when hunting seasons closed. Francis and others movements between species, but black duck duckling survival (1998) estimated annual survival during three periods on the rates were greater than mallard survival rates in 1 of 2 years. basis of changes in harvest regulations. Mean survival rate Longcore and others (1998) determined that mean sizes of increased from the first (1950–66) to the second (1967–82) Class II-III broods of black ducks (slightly less than 4 to 4.5 period following initial restrictions on harvest, a finding that is ducklings per brood) equaled or exceeded those of mallards consistent with a model of additivity of hunting mortality. The regardless of habitat type; moreover, black duck females with increase in survival rates following a second round of harvest broods were not competitively excluded from inhabiting fertile restrictions revealed some evidence for an increase in survival wetlands in Maine. The period (late August to mid-December for immature males between the second (1967–82) and third 1985–87) survival rate for post-fledging female black ducks (1983–93) periods. For adults, however, survival increased equipped with transmitters in Maine was 0.593; survival was less than expected if hunting mortality was additive. These 0.694 when losses from hunting were censored (Longcore and researchers concluded that evidence of additive mortality others, 1991). This period estimate multiplied by interval rates existed in at least some age-sex classes of black ducks in all for hunting, winter, and breeding periods produced an annual periods, but that evidence was weaker in the post-1983 period, survival estimate of 0.262, about 12 percent less than the esti- perhaps indicating that harvest was falling below the threshold mate (0.38) made on the basis of analyses of banding data. for additivity.

Table 2. Mortality of radiomarked black ducks relative to the percentage of times (that is, telemetry locations) that radiomarked ducks were in the Saint Lawrence Estuary.

[Modified from Clugston and others, 1994]

Percentage of telemetry Mortality locations recorded in Unknown Shot / Total ducks Natural the estuary cause probably shot Less than 5 2 1 0 / 0 10 35–65 0 1 1 / 0 13 Greater than 95 0 0 10 / 2 15 seems clear, then,how thegeographicpositionof thenorthern location wason thebreedinggroundsandstaging areas.It additive. The timewasearlyinthehuntingseason,and of mortalityseemtodetermine whetherhuntinglossesare breeding andstagingareas. Therefore, thelocationandtiming old” maybeeasilyexceeded for adultfemalesandyoungon caution of Anderson andBurnham(1976)thatthe“thresh ducks, wasexceededinsomeyears,whichsupportedthe threshold ofadditivityforblackducks,especiallyimmature success andproductivitylow. black ducksandfoundthatstormtidespredatorskept nest productivity onSmithIsland,MD,withradiomarkedfemale black duck.Haramisandothers(2002,p. 22)evaluated the BirdsofNorth America seriesaccountforthe American piled pertinenthistoricalandmorerecentliteraturetoprepare 21.4 percentofthebroods.Longcoreandothers(2000b)com duck broods,thanwetlandswithapH>5.51,whichsupported greater percentageofbroods(78.6percent),includingblack invertebrates, hadlarge numbersofInsectaandsupporteda associated withreducednumbersofacid-intolerantmacro however, revealedthatlow-(<5.51)pHwetlands,although taminants. FieldworkinMaine(Longcoreandothers,2006), studies duringthisdecade,littleattentionwasfocusedoncon 2000s beautiful hypothesisbyanuglyfact….”(Huxley, 1870,p. 6). mon sense. The greattragedyofscience[is]theslayinga became evidentthat“Scienceisnothingbutorganized com in nochange. As objectivefieldworkreplacedconjecture,it ters; theremaining21.7percentofencountersresulted displaced bytheblackduckduring15.0percentofencoun black ducksduring63.3percentoftheencounters,butwere contrast, malemallardsthatinitiatedaninteractiondisplaced change occurredduring12.8percentoftheinteractions.In and displacedmallards87.2percentofthetime,whereasno interaction withmalemallardsdidnotloseanyinteractions breeding. They foundthatmaleblackducksinstigatedan interactions ofblackducksandmallardsinthefieldduring in Maine,McAuleyandothers(1998)observedaggressive lards generallywerenotdominantoverblackducks.Later (1996), however, observingcaptiveducks, reported thatmal duck populationinmanypartsofOntario.Hoysakand Ankney was theprimarycauseforlong-termdeclineofblack “competitive exclusion”ofblackducksfromfertilewetlands were unfounded.Merendinoandothers(1993)speculatedthat survival ofblackducksanddeterminedthatthecriticisms to testforapositiverelationbetweenbodymassandannual of Dufourand Ankney (1990)aboutanalyticalmethodsused lards. Krementzandothers(1990)respondedtocriticisms which mallardswereobservedthanonrouteswithoutmal more likelytobedecliningonBreedingBirdSurveyroutes Earlier, Francisandothers(1998)reportedthatthe Although Patuxentscientistscontinuedworkonvarious Sauer andDroege(1997)reportedthatblackduckswere Patuxent’s AmericanBlackDuckStudiesfromChesapeakeBaytoMaineandBeyond ------Brunswick from0.945to0.348canbeattributedmostly this perioddidnotdie. The decreaseinsurvivalrateNew when thehuntingseasonopened(table3). but declinedsharplyformarkedducksinNewBrunswick were similarinthe1-to2-monthperiodbeforehuntingbegan, survival ratesforNewBrunswick(0.945)andMaine(0.986) until November15.Kaplan-Meier(KaplanandMeier, 1958) Maine (Longcoreandothers,1991),withopeningdelayed with anearlyOctober1huntingseasonopening,andonein adjacent studyareas—oneinNewBrunswick(Parker, 1991), of immaturefemaleblackduckswasdeterminedontwo ing andstagingareasmayhaveahighsurvivalrate.Survival These dataindicatethatblackducksarenotshotonbreed ity inthosenorthernstudyareaswasassociatedwithhunting. and stagingareascouldbesevere,as85percentofallmortal of Anderson andBurnham(1976)thatharvestonthebreeding (Longcore andothers,2000a)furthervalidatedthecontention tion. Telemetry datafromNovaScotia,Quebec,and Vermont ing regulationsiscrucialtothefateofblackduckpopula United StatesandtheCanadianprovinceswithrespecttohunt in Canada. (Dickson, 1995)afterretrievedkilldeclinedsubstantially to 1995(15.916.5per100km recovery rateremainedhigh,thecountstabilizedfrom1994 100 square miles(mi to 16.8per100squarekilometers[km in Quebec.Evenafterasharpdeclinenumbers(from27.5 stabilization oftheresultsbreedingblackducksurvey Another exampleofaresponsetoharvestrestrictionswasthe stabilized theMWIforblackduckin Atlantic Flyway. restrictive harvestregulationsbeginningin1984mayhave a “speciesofinternationalconcern”andnotedthatthemore (Perry, 2002).Serie(2002,p.2)discussedtheblackduckas ducks respondifharvestisrestricted. hunter harvest. The nextquestion,then,waswhetherblack November 1–15 October 16–31 October 1–15 Before September30 November 15] Canada, openedOctober1;waterfowlhuntingseasoninMaine for NewBrunswick,Canada;waterfowlhuntingseasonin [Modified fromLongcoreandothers(1991)forMaineParker function ofwaterfowlhuntingseasonopeningdate. black ducksinMaineandNewBrunswick,Canada,asa Table 3. Time interval Most ducksinMainethatwerenotexposedtohunters The thirdBlackDuckSymposiumwasheldin2002 studied

Survival rateofradiomarkedhatching-yearfemale

2 )]) from1990to1993,wheretheband Survival ratein (1985–87) Maine 0.834 0.885 0.965 0.986 Location (yearsstudied) 2 [41.2to42.7per100mi

2 ] [71.2to43.5per New Brunswick, Survival ratein (1987–88) Canada 0.348 0.465 0.500 0.945

2 ]

- 75 - - -

Migratory Bird Research, Monitoring, and Management 76 The History of Patuxent: America’s Wildlife Research Story

In Maine, the Department of Inland Fisheries and Wild- concluded that the proximate cause of the long-term decline life (P.O. Corr, Maine Department of Inland Fisheries and of the black duck population was unlikely to be related to Wildlife, oral commun., 1983) monitored numbers of water- mallard distribution. Link and others (2006) examined black fowl broods, including black ducks, on 34 wetland brood- duck Christmas Bird Count (CBC) data on a regional basis rearing reference areas. During 1980–83, most duck seasons and found decreasing populations in the southern and central were 50 days long, with split seasons in the southern hunting parts of the wintering range, but more stable populations in zone that opened October 1st in the early or late season. The the northeastern parts of the range. In addition, the CBC and black duck daily bag limit was either one or two in 3 of 4 the MWI showed similar patterns of population change at years. In following years (1984–88), the season opening was the scale of the United States, which lends credibility to the usually delayed in the early split season to about October 15th long-term MWI data. Zimpfer and Conroy (2006), in their in the north zone and about November 16th in the south zone. attempt to model production rates in black duck populations, The daily bag limit was either zero or one in all split seasons discovered that they could not include habitat variables as except 1988, when it reverted to two black ducks per day with predictors and that multicollinearity among some predictors no delayed openings in any split season. Numbers of black affected results, which indicated that the predictive ability of duck broods on these 34 reference areas by year are shown in the models was limited. figure 1. Kirby and others (2000) published keys of wings to iden- Delaying opening date, reducing season length, and tify mallard, black duck, and hybrids of these species. Petrie reducing daily bag in this northern state positively affected and others (2000) found no differences in clutch size, nest suc- the number of broods counted in years following protection of cess, hen success, duckling survival, or hen survival between local breeding pairs. Reed and Boyd (1974) documented the black ducks and mallards in New Brunswick, but purported high mortality of local black ducks breeding in the St. Law- that the difference in population status of the two species was rence Estuary during the opening weekend of hunting. Jorde related to differences in breeding propensity arising from and Stotts (2002, p. 31) dissected the Federal and State MWI competition for breeding resources. In contrast, McAuley and data into geographic areas and showed that trends in the num- others (2004) documented in nearby Maine that competitive ber of black ducks varied with geographic region. exclusion of black duck pairs from fertile wetlands by mal- Conroy and others (2002) assembled data on an array lards was unsupported by field observations, wherein 53 of 65 of variables affecting the black duck population and, with (81.5 percent) wetlands visited for 2 hours or more were used synthetic modeling, evaluated the relative importance of those by both black ducks and mallards. Increasing knowledge of variables. Longcore (2002, p. 7) contrasted the effects of black duck ecology and the positive effects of reduced harvest variables in the summer and winter ranges of black ducks and on the black duck population indicated that “In all science, error precedes the truth, and it is better it should go first than last” (Walpole, 1876, p. 128). The emerging facts seemed to indicate that hybridiza- Before harvest restrictions After harvest restrictions tion was not a likely cause of the black duck decline (Morton, 70 1998; Bolen and others, 2002). Furthermore, competitive exclusion was not plausible in light of increasing beaver- 60 created habitat (Longcore and Ringelman, 1980; Seymour and Mitchell, 2006), fewer breeding pairs (Longcore and others, 50 1987), dynamic use of wetlands by both species (McAuley and others, 2004), the fact that the black duck is as aggressive 40 as the mallard in defending territory and females, and the fact that the black duck is not dominated by the mallard (McAu- 30 ley and others, 1998). Past studies also determined that black duck brood females are not excluded from fertile wetlands and 20 black duck brood sizes are not different from those of mallards on fertile or infertile wetlands (Longcore and others, 1998),

Number of American black duck broods 10 and that mortality of black ducks caused by hunters can be additive to natural mortality (Francis and others, 1998). 0 So, if not the mallard, what was causing the black duck 1980 1981 1982 1983 1984 1985 1986 1987 1988 population to decline? Bolen and others (2002) make a case Year that sensitivity (that is, wariness or neophobia) of black ducks toward humans may have contributed to the black duck Figure 1. Number of black duck broods on 34 index wetlands in population decline. Without question, the prime Chesapeake Maine before (1980–83) and after (1984–88) harvest restrictions Bay wintering area for black ducks has been encroached on were applied to protect local breeding pairs. (Data from Maine by humans around the bay, with a 38-percent increase (from Department of Inland Fisheries and Wildlife, Bangor, ME) 2.0 to about 2.8 million) in the human population since 1970 Fish andWildlife Service,Laurel,MD) 1990–2008. (Data fromOfficeofMigratoryBird Management,U.S. Figure 3. Number of breeding black ducks (X 1,000) exceed thethresholdandbeadditivetonaturalmortality. and others(1998),however, reportedthathunterharvestcould additivity, whateverthatthresholdmighthavebeen.Francis we believedthathunterkillneverexceededathresholdof will affect thespringbreedingpopulation.Inotherwords, as compensatory—thatis,noduckshotinfallorlatewinter has not,beenaffected byharvestregulations?” port aconclusionthattheblackduckpopulationeitherhas,or 1968). The keyquestionwas,“Whatevidenceexiststosup reduced tostopthedeclineinblackducknumbers(Barske, biologists andadministratorsagreedthatharvesthadtobe decline intheblackduckpopulation.Evenlate1960s, consensus existedthatexcessiveharvestwasthecauseof (Longcore, 2002).Fromthe1800sto1930sand1940s,a in North America 350 400 450 500 550 600 Population ecologiststypicallyviewedhuntinglosses 1988

Number ofbreedingblackducks inNorthAmerica, 1990 1992 1994 1996 1998 Year Fish andWildlifeService,Laurel,MD;<,lessthan) 1990–2008. (DatafromOfficeofMigratoryBirdManagement,U.S. Figure 2. Number of black ducks (X 1,000) Adjusted r y =-11,077.2+5.78x 2000

harvested in North America Patuxent’s AmericanBlackDuckStudiesfromChesapeakeBaytoMaineandBeyond 150 200 250 300 350 400 450 1988 2002

2 Number ofblackducksharvestedinNorthAmerica, =0.45;p0.0009 1990 2004 Adjusted r y =19,840.8–9.3x 1992 2006 1994 2 2008 =0.77;p<0.0001 1996 - 2010 1998 Fish andWildlife Service,Laurel,MD;<,lessthan) 1990–2008. (Data fromOfficeofMigratoryBird Management,U.S. Figure 4. Year substantially (fig.4). stantially (fig.3)whilethemallardpopulationalsoincreased reduced (fig.2),thenumberofblackduckshasincreasedsub veys initiatedinthe1990sindicatedthatasharvesthasbeen (Longcore andothers,2000b).Breedinggroundpairsur since 1992havereversedthedownwardpopulationtrend breeding groundsincreases. expected untilthenumberofbreedersthatreturntomajor stantial responseinpopulationgrowthprobablycannotbe now breedintheUnitedStates(asopposedtoCanada),a sub Atlantic Flyway(Serie,2002,p.3).Becausefewblackducks the UnitedStatesharvestcouldonlystabilizeMWIin Canada) werenoteffective untilabout1990,thereductionsin Because restrictionsonthebreedinggrounds(mostlyin Number of breeding mallards (X 1,000) in Eastern survey 2000 Restrictions onharvestintheUnitedStatesandCanada 320 340 360 380 400 420 440 460 480 1988 2002

Number ofbreedingmallardsin EasternSurvey, 1990 2004 1992 2006 1994 2008 1996 2010 1998 Year Adjusted r y =19,949+5.18x 2000 2002 2 =0.63;p<0.0001 2004 2006 2008 - 2010 77 - -

Migratory Bird Research, Monitoring, and Management 78 The History of Patuxent: America’s Wildlife Research Story

Future Challenges approach to investigating all relevant variables to improve understanding of black duck ecology and population dynam- The original goal of the Black Duck Management ics. The consistent and persistent advocacy of private citizens Plan for North America, 1980–2000 (Spencer, 1980), was and waterfowl managers was essential to expose conjecture to “…reverse the apparent downward population trend…” and obtain objective data to explain and propose actions to as expressed in the MWI. As indicated by data from the reverse the long-term population decline of the black duck. improved waterfowl breeding pair survey, that goal has been achieved; however, this success resulted largely from reducing harvest by applying restrictions in areas where opportunity References Cited for exceeding the threshold of additivity was small—that is, south of the primary breeding and staging areas. Conjecture Addy, C.E., 1953, Fall migration of the black duck: U.S. Fish about the role of the mallard in the black duck decline was not and Wildlife Service, Special Scientific Report—Wildlife, supported by objective field studies of sympatric populations no. 19, 95 p. of these species. Additive effects of hunting were exposed as the black duck population began to recover following sub- Addy, C.E., 1968, General review, in Barske, Philip, ed., The stantial reductions in harvest. Even after 80 years of research, black duck—Evaluation, management, and research—A an expanding human population, which will increase human symposium: Boston, MA, Atlantic Flyway Council and disturbance and neophobia (Bolen and others, 2002), and Wildlife Management Institute, p. 2–5. energy development across Canada may affect where black ducks can breed or winter, thereby affecting productivity. Anderson, D.R., and Burnham, K.P., 1976, Population ecology For example, some wintering populations of black ducks are of the mallard—VI. The effect of exploitation on survival: shifting northward (Brook and others, 2007), which may affect Washington, D.C., U.S. Fish and Wildlife Service, Resource breeding success or survival, but the outcome is unknown. Publication 128, 66 p. Over the long term (1955–2007) in Maine, size of waterfowl Anderson, D.R., Burnham, K.P., Nichols, J.D., and Conroy, broods, including those of black ducks, seems to be declin- M.J., 1987, The need for experiments to understand popula- ing (Schummer and others, 2011); this decline may indicate tion dynamics of American black ducks: Wildlife Society contaminant effects on egg hatchability or increased duckling Bulletin, v. 15, no. 2, p. 282–284. [Also available at http:// mortality. Changes in brood survey methods, however, may www.jstor.org/stable/3782623.] have affected these results. For the early brood counts, broods of one or two ducklings were considered “incomplete broods” Ankney, C.D., Dennis, D.G., and Bailey, R.C., 1987, Increas- and were not included in calculating average brood size (H.E. ing mallards, decreasing American black ducks—Coinci- Spencer, Jr., Maine Department of Inland Fisheries and Wild- dence or cause and effect?: Journal of Wildlife Manage- life, oral commun., 1983), thus biasing the means higher than ment, v. 51, no. 3, p. 523–529. [Also available at http:// they would have been if broods of all sizes had been included. dx.doi.org/10.2307/3801262.] The next generation of black duck biologists will undoubtedly be vexed by some of the old issues and faced with new chal- Ankney, C.D., Dennis, D.G., and Bailey, R.C., 1989, Increas- lenges to sustain the North American black duck population. ing mallards, decreasing American black ducks—No evidence for cause and effect—A reply: Journal of Wildlife Management, v. 53, no. 4, p. 1072–1075. [Also available at Acknowledgments http://dx.doi.org/10.2307/3809612.] Barske, Philip, ed., 1968, The black duck—Evaluation, I thank Lynda J. Garrett (Patuxent Wildlife Research management and research—A symposium: Boston, MA, Center [Patuxent]) for always finding the reference I needed, Atlantic Flyway Council and Wildlife Management Insti- Brad Allen (Maine Department of Inland Fisheries and Wild- tute, 193 p. life) for supplying harvest regulations for Maine, and Mat- thew Perry and Shannon Beliew (Patuxent) for reviewing and Bellrose, F.C., and Chase, E.B., 1950, Population losses in the editing this manuscript, which also benefited from reviews mallard, black duck, and blue-winged teal: Urbana, IL, Illi- by Daniel McAuley (Patuxent) and Patrick Devers (U.S. Fish nois Natural History Survey, Biological Notes, no. 22, 27 p. and Wildlife Service). My Orono field station colleagues Dan [Also available at http://dx.doi.org/10.5962/bhl.title.15021.] McAuley and David Clugston; University of Maine gradu- Blandin, W.W., 1992, Population characteristics and simula- ate students James Ringelman, Catherine Frazer, and Charles tion modeling of black ducks: Washington, D.C., U.S. Kitchens; and many summer biological technicians contrib- Fish and Wildlife Service, Technical Report Series, no. 11, uted substantially to research on black ducks at the Maine 200 p. [Also available at http://handle.dtic.mil/100.2/ field station. The many and varied publications by the Patux- ADA322806.] ent staff about black ducks attest to the center’s multifaceted Clugston, D.A.,Longcore,J.R.,andMcAuley, D.G.,1994, Chasko, G.G.,Hoehn, T.R., andHowell-Heller, Penelope, Chamberlain, E.B.,1968,Someobservationsonaerialsurveys Carney, S.M.,1992,Species,ageandsexidentificationof Cain, B.W., 1981,Nationwideresiduesoforganochlorine Buckingham, N.,1937, The Mississippi Valley, Brook, R.W., Ross,R.K., Abraham, K.F., Fronczak,D.L., Boyd, H.,andHyslop,C.,1985, Are huntinglosses of Bolen, G.M.,Morton,Eugene,Greenberg, Russell,andDer [Also availableat Canadian JournalofZoology, v. 72,no. 12,p.2100–2104. ducks ( Habitat useandmovementsofpostfledging Americanblack http://dx.doi.org/10.1007/BF01607518 and Toxicology, v. 32,no.1,p.417–428.[Alsoavailableat fed naturalfoods:BulletinofEnvironmentalContamination 1984, Toxicity ofleadshottowildblackducksandmallards agement Institute,p.113–119. 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[Alsoavailableat 1979–80: PesticidesMonitoringJournal,v. 15,no.3, compounds inwingsofadultmallardsandblackducks, Societies, p.10–16. tomorrow?: New York, National Association of Audubon org/10.2193/2007-424 v. 73,no.1,p.98–103.[Alsoavailableat distribution change:Journalof Wildlife Management, and Davies,J.C.,2007,Evidenceforblackduckwinter org/10.1007/978-1-4612-5138-5_15 v. 29,partC,p. 181–195.[Alsoavailableat New York, Springer-Verlag, LectureNotesinStatistics, B.J.T., andNorth,P.M., eds.,Statisticsinornithology: young blackducks( searchable.pdf at USGS/BRD/ITR—2002–0005, p.16–22.[Alsoavailable U.S. GeologicalSurveyInformationand Technical Report Chesapeake Bayhabitats—Proceedingsofasymposium: views, rickson, Scott,2002,Mallardsreplacingblackducks—Two https://www.pwrc.usgs.gov/prodabs/perry/6039_perry_ in Anas rubripes Perry, M.C.,ed.,2002,Blackducksandtheir .] in Barske,Philip,ed., The blackduck— http://dx.doi.org/10.1139/z94-281 Anas rubripes ) intheSt.Lawrenceestuary, Quebec: .] .] http://www.biodiversityli Patuxent’s AmericanBlackDuckStudiesfromChesapeakeBaytoMaineandBeyond ) toohigh?, .] .] http://dx.doi. http://dx.doi. in in http://www4. Ducksfor Morgan, .] .] - - - Conroy, M.J.,Goldsberry, J.R.,Hines,J.E.,andStotts,D.B., Conroy, M.J.,Costanzo,G.R.,andStotts,D.B.,1989b, Conroy, M.J.,andBlandin, W.W., 1984,Geographicand Conroy, M.J.,Barnes,G.G.,Bethke,R.W., andNudds, T.D., Cottam, C.M.,1939,FoodhabitsofNorth American diving Cooke, W.W., 1915,Birdmigration: Washington, D.C.,U.S. Cooke, W.W., 1906,DistributionandmigrationofNorth Conroy, M.J.,Miller, M.W., andHines,J.E.,2002,Iden Conroy, M.J.,andKrementz,D.G.,1990, A reviewofthe Conroy, M.J.,andKrementz,D.G.,1986,Incorrectinfer v. 52,no.4,p.694–703.[Alsoavailableat American blackducks:Journalof Wildlife Management, 1988, Evaluationofaerialtransectsurveysforwintering org/10.2307/3801314 53, no.1,p.99–109.[Alsoavailableat the Atlantic coast:Journalof Wildlife Management,v. 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