Wilson Bull, 115(4), 2003, pp. 431-437 POPULATION RECOVERY OF THE EASTERN BROWN PELICAN FOLLOWING ITS EXTIRPATION IN LOUISIANA G. O. HOLM, Jr.,1’4 T J. HESS, Jr.,^ D. JUSTIC,i L. McNEASE,^ R. G. LINSCOMBE,^ AND S. A. NESBITU — ABSTRACT. We report on the exponential growth of the Eastern Brown Pelican (Pelecanus occidentalis carolinensis) in Louisiana, following its extirpation in 1963, and its subsequent reintroduction (1968-1980). This population growth pattern is remarkable since the Brown Pelican exhibits low fecundity, a long life span, and is considered largely nonmigratory in the GulfofMexico. To understand the regional changes in breeding numbers of Brown Pelicans in the northern Gulf ofMexico, we investigated the long term trends ofthe other subpopulations in Florida and Texas. The Florida subpopulation of Brown Pelicans has exhibited a stable-limit cycle, but within the last decade the number ofnesting birds has declined while the numberofnesting colonies hassteadilyincreased. ThenumberofBrownPelicansinTexasnowexceedsestimatespriortothetimepesticides caused reproductive failure of the subpopulation. The Louisiana subpopulation now equals or exceeds its his- torical (pre-pesticide) numbers. Although local recruitment can account for the exponential growth of Brown Pelicans in Louisiana, we believe that dispersal from colonies outside of Louisiana may have augmented the growth ofits nesting population. Received 11 February 2003, accepted 15 October2003. Prior to the 1930s, estimates of the Loui- 1985), the extirpation of Louisiana’s subpop- siana subpopulation ofthe Eastern Brown Pel- ulation was attributed to direct bird mortality ican {Pelecanus occidentalis carolinensis) (Blus and Joanen 1975) and more recently to ranged from 12,000-85,000 (Lowery 1955, a reduction in prey abundance resulting from 1974; King et al. 1977; Wilkinson et al. 1994), endrin contamination (Biglane and LaFleur with a conservative estimate of 50,000 indi- 1967; Cohen 2000, 2001). The rapid disap- viduals for Texas and Louisiana combined pearance of the Brown Pelican (Norman and (Schreiber 1980). Extirpation of the Brown Purrington 1970) coincided with endrin use Pelican in Louisiana had occurred by 1963 and production discharges (1958-1964) and (Williams and Martin 1968), and the last re- massive fish kills (1963-1964) caused by en- corded nesting effort of the native population drin contamination (Mount and Putnicki was observed in 1961 (Van Tets 1965). Dur- 1966). However, it was not until 1975, when ing this same period, a severe decline in the a severe die-off of pelicans (after reintroduc- Texas subpopulation occurred, and King et al. tion) in Louisiana occurred, that the extreme (1977) estimated that by 1961 <50 adults re- sensitivity ofpelicans to endrin contamination mained in Texas. Despite the extreme losses was documented (Blus et al. 1979). in Texas and Louisiana, the number ofBrown In 1968, the Louisiana Dept, of Wildlife Pelicans nesting in Florida remained generally and Fisheries and the Florida Fish and Wild- stable (Williams and Martin 1968, Kushlan life Conservation Commission began reintro- and Frohring 1985, Wilkinson et al. 1994). duction of the Brown Pelican into coastal A While reproductive failure from organo- Louisiana. total of 1,276 Florida nestlings chloride pesticide contamination caused the were transplanted from six colonies in Florida decline ofpelicans in Texas (King et al. 1977, to three sites in Louisiana between 1968 and 1980 (for detailed transplanting information see Nesbitt et al. 1978, McNease et al. 1984). ' Coastal Ecology Inst, and Dept, of Oceanography The first nesting attempts oftransplanted birds and Coastal Sciences, Louisiana State Univ., Baton occurred during the spring of 1971, and at that Rouge, LA 70803, USA. time, the birds were almost 3 years old (Wil- ^Louisiana Dept, ofWildlife and Fisheries, Rocke- liams and Joanen 1974). During the spring of feller Wildlife Refuge, Grand Chenier, LA 70643, 1975, endrin pollution caused a die-off of USA. ^Florida Fish and Wildlife Conservation Comm., about 40% of a standing population of 400- Wildlife Research Lab., 4005 South Main St., Gaines- 450 Brown Pelicans (Winn 1975, Nesbitt et ville, FL 32601-9099, USA. al. 1978, Blus et al. 1979). Since then, the '’Corresponding author; e-mail: [email protected] Louisiana population has continued to grow. 431 432 THE WILSON BULLETIN • Vol. 115, No. 4, December2003 The objective of our study was to investi- gate the long term growth trend ofthe Eastern Brown Pelican in the GulfofMexico, with an emphasis on the nesting birds in Louisiana, and compare these numbers to those of Flor- ida and Texas. Given that the Louisiana sub- population has exhibited an exponential in- crease of nesting birds, we investigated the potential for local recruitment versus long dis- tance emigration and dispersal from surround- ing subpopulations in augmenting Louisiana’s population growth. METHODS We used breeding Brown Pelican census data from Louisiana, Texas, and Florida, the states comprising the largest subpopulations of breeding pelicans along the U.S. Gulf of Mexico. These data were based on nest counts (Florida and Louisiana) or breeding pairs (Texas); hereafter, nest numbers are used as a common unit of measure. In Louisiana, peli- can nests and nestlings were counted yearly (except 1991) with a mean of four flights (ro- tor or fixed-wing) per year (1993-2001). The nesting data were collected by the Louisiana Dept, ofWildlife and Fisheries (Hess and Lin- scombe 2001). Surveys prior to 1993 usually involved a combination of aerial and ground census techniques, as summarized by Mc- Nease et al. 1992. In Texas, breeding pairs were estimated as described in the Texas Co- Date lonial Bird Survey (Lange 1993). Nesting pair EIG. 1. The Brown Pelican breeding populations data from Texas were provided by the U.S. in Louisiana and Texas have grown exponentially Fish and Wildlife Service (Houston, Texas). while declines occurred in Elorida. The Florida Fish and Wildlife Conservation Commission collected Florida nesting data. Nest counts in Florida were conducted during RESULTS tpewaok nzeosnteisn:g pAetrliaondtsicanadnsdumGmualrfizoefd bMeetxwieceon icaLnosuiisniaLnoau.i—siTahnea phoapsulgatrioownnofexnpeosnteinntgiaplelly- coasts. from 1971-1999, with slower rates of growth = WN^ee''u)setdo caanlceuxlpaotneentthiealingtrrionwsitchgerqouwattihonrat{eNs, o2c0c0u1r,ritnhge dpuorpiunlgati1o9n99p-e2a0k0e1d (aFtig.16,1)4.05Dunreisntgs of nesting subpopulations in Louisiana and (Fig. 1). A best-fit exponential model describ- Texas (Manugistics, Inc. 1995). In the equa- ing population growth from 1971-2001 sug- tion above, N, is the estimated numberofnests gested an intrinsic growth rate (r) = 0.25 for a given year, is the initial number of - 0.97). nests at time zero, r is the intrinsic growth The overall (1971-2001) mean nestling rate, and t is the time in years. Number of production per active nest was 1.66. Peak nests and time served as dependent and in- nestling production occurred in 2001 with dependent variables, respectively. 34,641 young produced; at this time the high- ; Holm et al. • PELICAN POPULATION RECOVERY 433 Date FIG. 3. InFlorida,theAtlanticBrownPelicansub- populationremainedrelativelystablefrom 1990-2001, but there was a decrease in the Gulf coast subpopu- lation. dated on a 12-ha dredge spoil island (Baptiste Collette or Plover Island, near the mouth of the Mississippi River). This immense, concen- trated shift comprised more than one-half of the total nesting population in Louisiana. Interannual fluctuations in the Louisiana subpopulation occurred during the following three time intervals: 1989-1990, 1995-1996, and 1999-2000 (Fig. 1). Decreases in nest numbers during the spring of 1990 and 1996 correspond to years that had protracted freez- ing temperatures during the preceding winters. During 1999 and 2000, a 50-100 year local drought coinciding with low Mississippi River stages resulted in high estuarine salinity across the Louisiana coast. 1965 1970 1975 1980 1985 1990 1995 2000 2005 — Florida Florida nesting birds have exhib- . Date ited cyclic changes in nest numbers from 1968 FIG. 2. While the number of Brown Pelican col- onward (Fig. 1). Following a general pattern onies increased along Florida’s Gulfcoast (a mean of of increasing growth from 1968-1990, state- one colony every two years since 1968), the number wide pelican nest numbers have declined over ofnesting birds decreased (see Fig. 1). The numberof the past decade, and the number of nests nesting colonies in Texas and Louisiana has remained (6,432) in 2001 fell below the 1968-1990 low despite the immense increase in the number of mean of approximately 8,000 nests. nesting birds. The decline in the number of nests in Flor- ida has occurred most noticeably among the est yearly mean numberofnestlings per active Gulf coast colonies, while colonies located nest (2.11) was recorded for Louisiana. The along the Atlantic coast have remained rela- peak number of nesting colonies (11) in Lou- tively stable (Fig. 3). Concomitant with the isiana occurred during the 2000 breeding sea- decline in nest numbers on Florida’s Gulf son (Fig. 2). An interannual increase in nest coast, there was a linear increase in the num- production (2,639) occurred from 2000-200 ber of colonies, and a mean (1968-2001) in- 1 concomitantly, four nesting colonies were crease of—one colony every two years (Fig. 2). abandoned (Fig. 1, Fig. 2). In 2000, a new Texas. A peak nesting population of3,373 colony ofapproximately 10,000 nests was ini- breeding pairs among a total ofseven colonies 434 THE WILSON BULLETIN • Vol. 115, No. 4, December2003 was recorded for Texas in 2001 (Fig. 1). The does indicate that pelicans in Florida are seek- mean number of nesting pairs from 1997- ing either new nesting habitat or improved 2000 was 2,839. The Texas population has forage conditions, or they are avoiding distur- grown exponentially with an intrinsic growth bance. Schreiber (1976) believed that food rate (r) = 0.22 {R^ = 0.94, Fig. 1). Fluctua- availability was the main reason for move- tions in the number of nesting pairs in Texas ment of nonbreeding Brown Pelicans. Adja- are similar to those observed in Louisiana cent to Florida, Alabama’s Gaillard Island with noticeable interannual peaks anddeclines nesting colony has grown rapidly; there were occurring during 1989-1990, 1995-1996, and four nests in 1983 (the first account ofnesting 1999-2000 (Fig. 1). pelicans recorded in the state), and the popu- lation peaked in 1999 with approximately DISCUSSION 5,200 nests (R. B. Clay pers. comm.). From Since its extirpation and reintroduction, the early band returns. Wood et al. (1995) con- Eastern Brown Pelican in Louisiana has pro- cluded that the Alabama colony (Gaillard Is- vided a remarkable example of exponential land) was initiated by birds from the panhan- growth of a nonmigratory population (Palmer dle of Florida. — 1962) of long-lived (>20 years) birds. Newly Dispersal from natal colonies. In a long introduced species, with rapid maturation and term analysis of Brown Pelican band returns, high annual fecundity, commonly exhibit ex- Schreiber and Mock (1988) concluded that ponential population growth. For example, Florida’s west coast (Gulf of Mexico) birds Bock and Lepthien (1976a, 1976b) found that comprised a distinct subpopulation from that populations of Cattle Egrets (Bubiilcus ibis; r of Louisiana’s. In their study, 82% of band = 0.21, 1956-1971) and House Finches {Car- returns were from Florida’s Gulf coast, indi- = podacus mexicanus, r 0.23, 1962-1971) cating a largely resident population. Other grew exponentially after their introductions band returns were from the Keys (8%), Cuba into regions ofthe United States. Interestingly, (5%), and Alabama (1%), but none were re- the Louisiana pelican subpopulation has ported from Louisiana. In a later study on pel- grown at intrinsic rates similar to that of “r- ican dispersal. Wood et al. (1995) found, con- selected” bird species. This raises the ques- versely, that banded birds from a natal colony tion of whether local recruitment can account on the Florida panhandle moved both east- for the rapid increase in the Brown Pelican ward and westward along the Gulf coast, and nesting population in Louisiana. movement into Louisiana was documented Based on a simple exponential model, local (though only one band was recovered in Lou- recruitment could explain the observed isiana). growth of the Brown Pelican population in Long range dispersal ofpelicans throughout Louisiana, provided that the estimated growth the Gulf of Mexico has been reported histor- rate is 0.25/year. However, since this rate is as ically (Mason 1945, Schreiber and Mock high as those observed in r-selected species, 1988, Sykes and Langridge 1991, Johnsgard we suggest an alternative explanation, i.e., 1993, Wood et al. 1995). As of 2001, banded that immigration into Louisiana has augment- pelicans from Louisiana have been reported ed this growth. This immigration may have from every Gulf coast state, and also from involved subadult pelicans, produced else- Cuba, the Yucatan Peninsula, Belize, and where, joining the Louisiana nesting popula- Guatemala (J. Harris pers. comm.). These re- tion. This hypothesis also would explain the cent Louisiana band returns confirm earlier protracted decline in the Florida Gulf coast findings (Mason 1945, Schreiber 1976) that population concurrent with the rapid increase Brown Pelicans, especially juveniles, move in the Louisiana breeding population. randomly and for long distances. A distinct The ability to detect emigration into Loui- pattern ofdispersal from Louisiana’s colonies, siana from banded birds has been constrained however, is not discernable at this time. by the remoteness of pelican breeding and Whether breeding birds are shifting toward feeding areas in Louisiana. However, the the northernmost portions ofthe GulfofMex- steady increase of nesting colonies on Flori- ico remains debatable. Nonetheless, this pat- da’s Gulfcoast, while nest numbers decreased. tern is conceivable given the northward ex- Holm et al • PELICAN POPULATION RECOVERY 435 pansion ofbreeding colonies along the Atlan- Louisiana has grown only 4.8%. Thus, habitat tic coast ofthe U.S. Since 1980, first breeding fragmentation by coastal land development records for Brown Pelicans have been report- and the probability of nesting colony distur- ed for Georgia, Virginia, Maryland, Delaware, bance is ofgreaterconcern in Florida andTex- and New Jersey (see Wilkinson et al. 1994). as than in Louisiana. — Warm temperatures may facilitate a northward Habitatstability. The greatestfuture prob- dispersal pattern, and the warmest decade lem for Brown Pelicans in Louisiana is the since instrumentation began (1861) occurred stability of natural nesting habitat. Barrier is- 1990-2000 (Houghton et al. 2001). An inter- lands in Louisiana are the aging relics ofMis- decadal comparison between the 1980s and sissippi River deltas, and the rapid erosion of mid-1990s showed a notable reduction in pel- these islands has been largely irreversible, as ican numbers (>70% or 1,300 individuals) in there is little “new” sand or coarse-grained Puerto Rico and the U.S. Virgin Islands (Col- material entering the system. This has resulted lazo et al. 1998). The authors did not find ev- in the need to replenish, with dredge material, idence of reproductive failure or nesting and historically successful natural nesting islands roosting habitat changes. — in Louisiana. The use of created islands Limitations on population growth. Nelson (dredge material) by nesting pelicans has been (1979) proposed that populations of pelicani- encouraging in Louisiana and elsewhere form species are controlled mostly by envi- (Wood et al. 1995). High rates of island frag- ronmental factors (climate and food availabil- mentation in Louisiana conceivably could fa- ity). Fledging success of the California sub- cilitate the establishment ofnew colonies that species (P. o. californicus) was dependent on have reduced predator activity (Kushlan and the local abundance of northern anchovy {En- Frohring 1985). The fact that within one year graulis sp.; Anderson et al. 1982). Along the more than 60% of the nesting population ini- Pacific coastlines ofNorth and South America tiated a colony on a relatively small dredge the importance of food abundance on seabird spoil island reflects this species plasticity to mortality is well understood, especially as it changes in habitat. is related to El Nino-Southem Oscillation The mean number of nests per colony in events (Duffy et al. 1988). In the GulfofMex- ico, relationships between food supply and Louisiana in 2001 (2,344 nests per colony) is an order of magnitude higher than that of breeding success have not been widely stud- Florida’s statewide density (161 nests per col- ied. Poor food availability, however, has been ony). The exponential increase in nests we ob- implicated in reproductive failures in southern Florida near the Keys (Kushlan and Frohring served in Louisiana contrasted with the dis- 1985). Moreover, there was a strong, positive proportionate number ofnew nesting colonies association between statewide mullet (Mugil suggests that a sustained amount of nesting spp.) landings and Brown Pelican nesting ef- habitat, rather than food availability, will be a fort in Florida during the period 1968-1992 future concern in Louisiana. In Florida, the (Nesbitt et al. 1996). Whether food supply has decrease in the number of nesting pairs along influenced decreasing nest numbers along the Gulf coast, with no decline in the number Florida’s Gulf coast is unknown. of colonies, is the reverse of what is being Other density independent forces such as seen in Louisiana. In Florida, the food base, human disturbance (Jehl 1973, Anderson and rather than the availability of suitable nesting Keith 1980, Anderson 1988) inhibit successful sites, may limit the population. breeding at local levels and may influence col- In general, the growth ofthe subpopulations ony site selection or abandonment. There are of Brown Pelicans in Louisiana, Texas, and significant differences, for example, in human Alabama, during periods when declines were population growth in the coastal areas among observed along the Gulf coast of Florida and Louisiana, Florida, and Texas (Barrett 2001). Puerto Rico and in the Virgin Islands, indi- From 1990 to 2000, the human population on rectly suggests that Brown Pelicans may have Florida’s Gulf coast has increased 20.8%, and the ability to shift to habitats where suitable Texas’ population has grown 18.7%. In con- nesting and forage conditions exist outside of trast, the population of coastal parishes in intense human disturbance. 436 THE WILSON BULLETIN • Vol. 115, No. 4, December2003 ACKNOWLEDGMENTS Proceedings of the 19th Ornithological Congress (H. Ouellet, Ed.). Univ. ofOttawa Press, Ottawa, We thank E. Erfling (USFWS Houston, Texas) for Ontario, Canada. providing Brown Pelican data from Texas. R. B. Clay Hess,T.J.,Jr. andJ. Linscombe. 2001. BrownPelican (AlabamaDept, ofConservation andNatural Resourc- coastal survey, site observations, and summary. es) provided data from the Gaillard Island colony in Louisiana Dept, of Wildlife and Fisheries, Grand Alabama. J. Harris (USFWS Slidell, Louisiana) dis- Chenier, Louisiana. cussed banding efforts, returns, andotherobservations Houghton, J. T, Y. Ding, D. J. Griggs, M. Noguer, olefttBeraonwdnCPhealnidcealneunresctoilnogniacetsiviintyLoautistihaenBaa.ptWiesteapCporle-- P2.00J1..VCalnimadteercLhianndgeen,20a01n:d tDh.e sXciiaenotsifuic(Ebdass.i)s.. ciatethesuggestionsofP. J. Mockandtwoanonymous Cambridge Univ. Press, Cambridge, United King- reviewers that improved the manuscript. Lastly, we dom. thank all those who worked hard to bring the Brown Jehl, j. R., Jr. 1973. Studies ofadecliningpopulation Pelican back to Louisiana. of Brown Pelicans in northwestern Baja, Califor- LITERATURE CITED nia. Condor 75:69-79. JoHNSGARD, P. A. 1993. Cormorants, darters and peli- Anderson, D. W. 1988. Dose-responserelationshipbe- cans ofthe world. Smithsonian Inst. Press, Wash- tween human disturbance and Brown Pelican ington, D.C. breeding success. Wildl. Soc. Bull. 16:339-345. King, K. A., D. R. Blankinship,E. 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