Long-Term Population Trends of Colonial Wading Birds in the Southern United States: the Impact of Crayfish Aquaculture on Louisiana Populations
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The Auk 112(3):613-632, 1995 LONG-TERM POPULATION TRENDS OF COLONIAL WADING BIRDS IN THE SOUTHERN UNITED STATES: THE IMPACT OF CRAYFISH AQUACULTURE ON LOUISIANA POPULATIONS BRUCE E. FLEURY AND THOMAS W. SHERRY Departmentof Ecology,Evolution, and Organismal Biology, Tulane University, New Orleans, Louisiana 70118, USA Al•sTl•C•r.--Long-termpopulation dynamics of colonialwading birds (Ciconiiformes) were examinedusing data from Audubon ChristmasBird Counts(CBC, 1949-1988)and Breeding Bird Surveys(BBS, 1966-1989). Winter populationsof Louisianawading birds increased dramaticallyover the 40-yearperiod, with the sharpestincreases occurring during the last 20 years.Several species populations grew exponentiallyfrom 1968to 1988.High overall positivecovariance was found in the abundanceof the variousspecies over time, and cluster analysisshowed that the specieswith similardietary requirements and foraginghabits cov- ariedmost strongly and positivelywith eachother. Trend analysis of CBCand BBSdata from 1966-1989and 1980-1989showed a closecorrespondence between CBC and BBStrends in Louisiana.Most speciesincreased in Louisianaat the sametime as they declined in Florida andTexas. Several factors might explain increases in populationsof wadingbirds in Louisiana, includinglong-term recovery from the effectsof humanexploitation, expansion of breeding populationsin more northern states,changes in weather,recovery from DDT and similar pesticides,and regionalmovements due to habitatloss in other coastalstates. These hypoth- esesare not mutuallyexclusive and meritfurther study. However, based on severalinferential lines of evidence,increased acreage devoted to crayfish(Procambarus) aquaculture in Louisiana appearsto be the mostsignificant factor explaining observed population trends in the state. First,populations of colonialwading-bird species that usecrayfish were correlated positively with the wild crayfishharvest in Louisiana,and even morestrongly with commercialcrayfish pond acreage.Second, the regularity with which theseponds are managedprovides a more predictableforaging habitatthan do correspondingnatural areas.Third, the use of crayfish pondsby wading birds peaksduring pond drawdowns,which may increasereproductive successby concentratingprey availableto wading birds during their nestingseason. Fourth, those speciesof wading birds that specializeon crayfishshowed the greatestpopulation increasesand the strongestcorrelation with crayfishpond acreage.These findings have importantimplications for conservationand managementof Louisiana'swading-bird pop- ulations. Received16 May 1994, accepted5 September1994. ALTHOUGH WE LACK reliable nineteenth cen- and inland (Ogden 1978, Ryder 1978). How- tury data on colonial wading-bird populations ever, wading-bird increaseshave not occurred (order Ciconiiformes), accounts from pioneer in all regions. Whereas populations have in- naturalists like Audubon document enormous creasedand rangesexpanded in the past30 years coloniesthroughout the southernswamps and in someregions of the United Statesfor species estuaries(Bent 1926). By the end of the nine- suchas Snowy Egrets,Great Blue Herons, and teenth century, plume hunting had reduced Cattle Egrets(Bock and Lepthien 1976, Ryder many speciesto near extinction. Legal protec- 1978, McCrimmon 1982, Larson 1982; Table 1), tion began in 1900 with the Lacey Act, which regionaldeclines characterize other species, such prohibited interstate commerce in illegally as the Reddish Egret and Roseate Spoonbill killed animals,followed by the Migratory Bird (Powellet al. 1989,but seeOgden 1991). Florida, Treaty in 1916. The establishmentof wildlife for example, despite having a high historical refuges also helped protect wading-bird colo- populationdensity of most speciesof colonial nies (Parnell et al. 1988). wading birds (Root 1988), has witnessed re- Since their protection from plume hunters, duced wading-bird populations due to pro- many speciesof colonial wading birds have longeddrought, estuarine degradation, and hu- steadily increasedtheir North American pop- man-induced hydrological changes in and ulations, and expanded their rangesnorthward around the Everglades (Bancroft et al. 1988, 613 614 FLœUR¾AND SHERRY [Auk, Vol. 112 Bancroft 1989). Ogden (1994) estimated that TABLE1. Speciesof Ciconiiformesregularly seenon populationsof five speciesof wading birds in 343 Louisiana Christmas Bird Counts 1949-1988, and averagenumber observedper count. the Florida Evergladesdeclined by 93% be- tween 1931-1946 and 1982-1989. Such conflict- Aver- ing trends in populationsof colonial wading age per birds may even indicatelarge-scale shifts of re- Species count gional population centers. American Bittern (Botauruslentiginosus) 1.5 This diversity of population trends among Least Bittern (Ixobrychusexilis) 0.1 regions and speciesindicates the need for in- Great Blue Heron (Ardeaherodias) 41.3 creasedmonitoring, and for in-depth analyses Great Egret (Casmerodiusalbus) 122.8 Snowy Egret (Egrettathula) 118.3 at national and regional scales.This need is par- Little Blue Heron (E. caerulea) 81.3 ticularly apparentin Louisiana,where wading- Tricolored Heron (E. tricolor) 50.4 bird populationtrends have been poorly doc- ReddishEgret (E. rufescens) 0.2 mented in the past,with breeding-colonydata Cattle Egret (Bubulcusibis) 100.9 Green-backed Heron (Butoridesstriatus) 1.1 only availablefor 1977,1984, and 1990(Portnoy Black-crownedNight-Heron (Nycticoraxnyc- 1977, 1978, Keller et al. 1984, Martin and Lester ticorax) 13.0 1990, R. P. Martin 1991), and where wetland Yellow-crowned Night-Heron (N. violaceus) 0.3 acreageis more abundant than in any other White Ibis (Eudocimusalbus) 256.5 state. Glossyand White-facedibises (Plegadis falci- nellusand P. chihi) 432.1 We review population trends of 15 speciesof RoseateSpoonbill (Ajaia ajaja) 21.2 colonial wading birds in Louisianaover the past 40 years, using data extracted from Audubon ChristmasBird Counts (CBC). We compareand contrastthese trends with state, regional, and involves: tropical-fish farms in Florida; catfish nationaltrends from BreedingBird Survey(BBS) farms in California, Arkansas,and Mississippi; data. We use published data to discussbriefly and fish farms in Europe (Draulans 1987b,Stick- and evaluateseveral such factors, namely long- ley and Andrews 1989,Boyd 1991, Cezilly 1992, term recovery from the effects of the plume Stickleyet al. 1992).Wading birds currently are trade and other human exploitation, increased treatedas agriculturalpests under the Depart- winter populationsdue to expandedranges of ment of Agriculture'sAnimal DamageControl migratorywading birds in more northern states, program. Depredation permits have been is- effectsof weather, recoveryfrom heavy pesti- sued in several states for lethal control of sev- cide use, competition, predation, habitat loss, eral migratory bird species(including herons, and land-use changes in Louisiana due to the egrets, pelicans, and cormorants)that feed at growth of crayfish (Procambarus)aquaculture catfish farms and other aquaculture facilities. (note: in Louisiana, the common name "craw- Increasedpopulations of White Ibisesand Yel- fish" is used instead of "crayfish"). low-crowned Night-Herons, which feed heavi- Continueddeclines in the quality and quan- ly on crayfish,are of particularconcern to Lou- tity of foraging habitat in natural wetlands due isiana wildlife biologists and farmers. Sound to drainage, pollution, and other human influ- managementpolicy for colonial wading birds enceshighlight the importanceof understand- in Louisianawill require detailed understand- ing how and why birds use artificial wetlands ing of the relationship between wading-bird suchas flooded fields or aquacultureponds. The populationsand crayfishaquaculture, towards exponentialgrowth of commercialcrayfish pond which our analysesof wading bird population acreagein Louisianamay have directly contrib- dynamicsare an important first step. uted to increasesin Louisianawading-bird pop- ulationsby creatingnew high-quality foraging habitats.Maddock and Baxter(1991) suggested METHODS that food limitation is a critical factor in egret CBC.--We used data from the annual Audubon reproductive success. ChristmasBird Count (CBC) to analyze winter cicon- The useof aquacultureponds by wadingbirds iiform populationtrends in Louisiana.CBCs are held alsoraises important legal and conservationis- on a singleday between15 Decemberand 5 January. sues, both locally and internationally. The All birds seen within a 24.1-km (15-mi) diameter cir- problem of bird predationin aquacultureponds cle are tabulated.In addition,observer effort is gauged July1995] CrayfishAquaculture and Wading-bird Populations 615 of birds countedas stronglyas would the addition of new countcircles in wetland areas(Raynor 1975, Mor- rison and Slack 1977, Bock and Root 1981). Therefore, we calculated the total number of birds observed di- vided by the number of countsheld in that year. There are several methodologicalproblems with CBC data. CBC data are not truly random samples, sinceregions of high speciesdensity are favored in the selection of count areas (Drennan 1981). CBC counts also are subject to error due to year-to-year differencesin weather and timing of the count (Falk 1979, Smith 1979, Rollfinke and Yahner 1990), and due to observer-reliabilityerrors and volunteer turn- 0 1949 1959 1969 1979 1989 over (Stewart 1954, Confer et al. 1979, Arbib 1981, Butcher1990). CBC data are most meaningful when they covermany years and includea broadgeograph-