J. Field Ornithol.

, 68(2):287-295

THE IMPORTANCE OF THE CARIBBEAN COASTAL

WETLANDS OF NICARAGUA AND HONDURAS TO CENTRAL
AMERICAN POPULATIONS OF WATERBIRDS AND JABIRU
STORKS (JABIRU MYCTERIA )
PETER C. FREDERICK
Departmentof WiLtlifeEcolog
3, and Conservation
P.O. Box 110430
Universityof Florida
Gainesville, Florida 32611-0430 USA

JORGECORREASANDOVAL
El Colegiode la b¾onteraSur
Aptdo. Postal 424 Chetumal
77000 Quintana Roo,Mexico

CHARLESLUTHIN 1
ProjectLighthawk
P.O. Box 8163
Santa Fe, New Mexico 87504-8163 USA

MARILYN SPALDING
Departmentof Pathobiolog
O,
Universityof Florida
Gainesville, Florida 32611 USA

Abstract.--During the middle of the dry season (February-March), we performed low-

altitude aerial strip-censusesto estimate densitiesof waterbirds in the Atlantic coastalwet-
lands of Nicaragua and Honduras (Miskito Coast and La Mosquitia, respectively),
areaswhichare poorlyknownornithologically.
We found12.9and 5.5 waterbirds/km"
in
294.6and281.5km2surveyedin Nicaragua
andHonduras,respectively.
GreatEgrets(Ardea
alba) were most common and widespread,representing53% (Nicaragua) and 46% (Hon-
duras) of the sightings.Wood Storks (Mycteriaamericana)and Egv'etta
herons were the next
two most abundant groups. Breeding colonies of Wood Storks, RoseateSpoonbills (Ajaia
ajaja), and Great Egretswere found in both areas, most nestscontaining eggsand small
chicks.We document a previouslyunrecorded population of Jabiru Storks (Jabiru mycteria)
breedingin both areas,with densities
of 0.05 and 0.16 birds/km'2,respectively.
The wet-
lands of both countries appear to support a large proportion of the breeding JaNms in
Central America and should be given high priority for conservation.

LA IMPORTANCIA DE LOS ANEGADOS COSTEROS DEL CARIBE DE NICARAGUA Y

HONDURAS PARA LAS POBLACIONES DE CENTRO AMI•RICA DE AVES
VADEADORAS Y PARAJABIRU MYCTERIA
Sinopsis.--Llevamosa cabo censosa6reosde banda a poca altitud para estimar las densi-
dadesde avesvadeadorasen los anegadosde la costadel AtNntico de Nicaragua y de Hon-

• Current
address:
TheAldoLeopold
Shack
Foundation,
E-12919Levee
Road,Baraboo,
•scon-
sin 53913 USA.

287
288] P. c. Frederich
etal. j. Field
Ornithol.
Spring 1997

duras (Costa Miskito y La Mosquita, respectivamente),/treaspoco conocidasornitol6gica-

mente, a mediadosde la temporadaseca(febrero a marzo). Hallamos 12.9 y 5.5 avesvadea-
doras/km
2 en los 294.6km2 y 281.5Km2examinados
respectivamente.
Lasavesm•s co-
munesy mejor distribuidasfueron Ardea alba, representandoun 53% (Nicaragua) y un
46% (Honduras) de las observaciones. Las otrasdos especiesm/rsabundantesfueron Myc-
teria americanay Egrettathula. Se hallaron coloniasde nidos de Mycteriaamericana,Ajaja
ajaja y Egrettathula en ambas/treas,la mayoria de los nidos con huevosy pichones.Docu-
mentamosuna poblaci6n previamenteignorada de Jabiru mycteria,con densidadesde 0.05
aves/km
2 y 0.16aves/km
• reproduci6ndose
en ambas/treas.
Laszonasanegadas
de ambos
paisesparecen sosteneruna gran proporci6n de losJabiru mycteriaanidando en la Am6rica
Central, y se deberia dar prioridad a su preservaci6n.

The wetlandsof the Atlantic coastalregionsof Honduras (La Mosqui-

tia) and Nicaragua (Miskito coast) are someof the most extensivein Cen-
tral America, but remain poorly known ornithologically (Fig. 1). The
coastal wetlands and some of the more common waterbirds are described
briefly in the IWRB Directory of Neotropical Wetlands (Scott and Car-
bonnel 1986), and Camacho (1983) suggestedthe possibleimportance
of the area forJabiru Storks.Otherwise,the avifaunaof the area is known
in print from one early collection (Huber 1932), Monroe's (1968) survey
of easternHonduras,and Howell's (1971) studyof easternNicaragua.The
avifauna of the region may also be surmisedfrom severalreports from
southeastern Nicaragua(Richmond1893,Sclaterand Salvin1867,Sclater
1873, Will 1991), and Costa Rica (Stiles and Skutch 1989).
In this paper, we report on densitiesof large waterbirds in these wet-
lands during the dry season,as determined through low-altitude aerial
surveys, and comparethesedensitiesto thoseobtainedin studiesof other
New World tropical and subtropicalwetlands.We found that these rela-
tively unimpacted wetlandshost high densitiesof ciconiiform birds and
are of regional importancefor Jabiru Storks.We believe this effort is the
first quantitativesurveyof waterbirdsin the La Mosquitia/Miskito Coast
ecosystem.

METHODS

We performed low-altitude strip-censuses of waterbirds in the Atlantic

coastwetlands of the Miskito Goast of Nicaragua (March 1992) and La
Mosquitiain Honduras (February 1994). We counted birds systematically
on both sidesof a low-flyingaircraft, using standardmethodologyfor sur-
veys of wildlife in open-country savannas(Bancroft and Sawicki 1995,
Bjorkand Powell1994,Hoffman et al. 1990,Norton-Griffiths1975). If the
altitude is known, and observers on both sides of the aircraft restrict their
countsto demarcatedstripsof ground on either sideof the plane (outlined
with marks on the windows and struts), then the area in which animals have
been counted can be calculated (Norton-Griffiths 1975). Densities can
then be extrapolatedto total population countsif the habitat is uniform
or if the countscan be stratifiedby habitat type. We chosenot to do the
latter, sinceadequatevegetationmapswere not availableto us.
Flyingor standingbirdswere counted only if they occurredwithin these
visuallydefined strips.Ground positionswere obtained using a global po-
Vol.6S,No.2 Wetland
BirdsofHonduras
andNicaragua [289

Punta Patuca

0 20 40 km

Puarto Lempira

Honduras
Cabo Gracias
a Dios

Nicaragua

Puerto Cabezas

Haulover

FIGURE1. Map of coastalregionsof northeasternNicaraguaand southeasternHonduras,

showinglarge townsand locationsof strip-transect
aerial surveys(dashedlines).

sitioningsystemwith an accuracy<100 m. Bird speciesand numberswere

recorded onto paper forms by the observers,with positionscalled out
upon demandby a third observer.All participantscommunicatedby head-
set. Data were recorded separatelyfor each straight-linetransect(Fig. 1);
length of each transectwas calculatedusing the startingand ending po-
sitions of the transect.
We concentratedour surveysin areasof wetland that were not heavily
forestedand that had standingsurfacewater. We did not attempt to count
birds in the large open salinelagoonscharacteristicof the region, which
are generally >1 m in depth. Our searchfor wetlandsby definition was
290] •'. C. Frederick
et al. J.Field
Ornithol.
Spring 1997

biased towardssparsemangrove swampsand wet prairies, and tended to

exclude hardwood forest, pine savannas,and gallery forest. While we feel
our transectscovered representativewetland typeswith surfacewater at
the time of surveys,our studyshouldnot be construedashavingsampled
wetland habitat typesin proportion to their availabilities.Transectswere
generallyoriented east-westto make effectsof sun glare equal on both
sidesof the plane. The only exception to this rule were wetlands occur-
ring in thin stripsimmediately adjacent to beaches,where transectswere
conductedparallel to the long axis of wetlands.
SnowyEgrets (Egrettathula) and immature Little Blue Herons (Egretta
caerulea)are both white-plumagedand difficult to distinguishat the alti-
tudes we used. We therefore categorizedboth as "smallwhite herons,"
(Bancroft and Sawicki 1995, Hoffman et al. 1990). Similarly, adult Little
Blue Herons and Tricolored Herons (Egrettatricolor)are difficult to dis-
tinguish when viewed from above, and were both categorizedas "small
dark herons."
M.G.S. and P.C.F. were observerson surveyflights in the Miskito Coast
wetlandsof Nicaragua between the Rio Coco and Wounta on 3, 4, 8, and
9 Mar. 1992. Surveyflights were flown at 108 m altitude and 166 km/h
groundspeedin a Cessna206, counting birds in stripsof approximately
271 m on either sideof the plane.
P.C.F. and J.S.C. were observerson surveyflights in Honduras between
15 and 19 Feb. 1994, between the Rio Coco on the south and Cabo Ca-
maron in the north, in a centerline-thrusttwin-enginedCessna137 Sky-
master at 61 m altitude and 166 km/h groundspeed.Transectstotalled
697 km; usinga calculatedstripwidth of 206 m, we estimatedthat birds
were counted within a total area of 282 km '• in the Honduran wetlands.
RESULTS

Survey.
results.--Wefound3786waterbirds in the 295kin"surveyed in
Nicaragua,and1547in the282kin"surveyed in Honduras,yieldingover-
all densities
of 12.9and 5.5 birds/km
e, respectively
(Table1). In both
surveys,the most common specieswas the Great Egret, accountingfor
53% of the total in Nicaragua,and 46% of the total in Honduras.In Nica-
ragua,Wood Storkswere secondin abundance(17% of birds seen), small
dark heronscomprised10%, and smallwhite herons 7%. In Honduras,
small white herons (largelySnowyEgrets) comprisednearly 15% of the
total. The numbers of small dark herons are likely to be under-estimated
due to poor detectabilityof thesedark birds. White Ibises (Eudocimusal-
bus),a dominant speciesin southeasternUnited Statesand Venezuelan
surveys,were rarely seen in Nicaraguaand Honduras. RoseateSpoonbills
were seen in small numbers during both surveys.
Densitiesin Nicaragua were almost twice those of Honduras. Our den-
sitiesfrom the two countries were in the middle of the range of estimates
for centralFlorida(2.3-164.0birds/km",Frederickand McGehee1994),
and the central (6.6-106.89birds/km",Bancroftand Sawicki1995) and
coastal(17.5-24birds/km",Bjorkand Powell1993) Everglades. In all
Vol.68,No.2 Wetland
BirdsofHonduras
andNicaragua [291

TABLE1. Summaryof waterbirdsurveysin coastalHondurasand Nicaraguaduring the dry

season.

Honduras Nicaragua
Number Number Proportion Number Number Proportion
Species seen per km"of total seen per km'• of total
Anhinga anhinga 2 0.01 0.001 0 0 0
Anas discors 20 0.07 0 0 0 0
Cairina moschata 28 0.10 0.018 130 0.44 0.034
Dendrocygna
spp. 275 0.98 0.178 69 0.23 0.018
Unidentified ducks 4 0.01 0.003 0 0 0
Small white herons 226 0.80 0.146 273 0.93 0.072
Ardea herodias 29 0.10 0.019 126 0.43 0.033
Ardea alba 712 2.53 0.460 2011 6.83 0.531
Small dark herons 62 0.22 0.040 372 1.26 0.098
Ajaia ajaja 24 0.09 0.016 47 0.16 0.012
Eudocimus albus 63 0.22 0.040 59 0.20 0.016
Jabiru mycteria 13 0.05 0.008 47 0.16 0.012
Mycteriaamericana 89 0.32 0.058 652 2.21 0.172
Total, all species 1547 5.50 1.00 3786 12.85 1.00

cases, these densities have been standardized for the amount of wetted
marsh area surveyed (areas without standing surfacewater excluded).
Perhapsthe most strikingfinding wasthe number of Jabiru Storksthat
we counted(47 in Nicaragua,13 in Honduras),yieldingestimateddensi-
ties of 0.16 and 0.05/km2, respectively. If Jabirusseenjust outsidethe
transects are included,we sawa total of 74 in 545 km of flyingin Nicara-
gua, and 24 in 697 km of flyingin Honduras.Jabiruswere typicallysoli-
tary and in pairs, though a singleflock of four wasseen.
Breedingactivity.wOur surveyscoveredonly a small fraction of the po-
tential wetland nesting sitesin each study area. In Honduras, we located
two breeding aggregationsof long-leggedwading birds on islandswithin
or on the edges of Laguna Kohunta and a very small lagoon northwest of
Laguna Apalca. At the Laguna Kohunta colony we counted 165 Wood
Storknestswith eggsor chicks,and 17-20 RoseateSpoonbillnests,prob-
ably with eggs.At the much larger Apalca Northwestcolony,we counted
1520 Wood Stork nestswith eggsor chicks,155 RoseateSpoonbill nests,
and approximately 150 Great Egret nests. SnowyEgrets may also have
been nestingat this latter colony,but we were unable to confirm this with-
out ground visits.Birds in these colonies appeared to be in incubation
and early chick rearing activities,which would place dates of initiation in
late December (Wood Storks) to mid-January(RoseateSpoonbills).
During the Nicaraguan surveysin 1992, we also found small colonies
of Wood Storks(160 pairs) and RoseateSpoonbills(24 pairs). The Wood
Stork nestscontainedsmallyoung and eggs,and the RoseateSpoonbills
appeared to be incubatingeggs.Timing of nestingfor both thesespecies
in Honduras and Nicaragua appears quite similar to that observedin
Texas, Florida, and the Yucatan (Mien 1942, Bjork and Powell 1994, Cor-
rea 1992, Lopez Ornat and Ramo 1992).
292] P. C. Frederick
etal. J. Field Ornithol.
Spring 1997

In Nicaragua, we found four Jabiru nests,two of which were inside our

transects.Two additionalJabiru nestswere found incidentallyduring 6 h
of manatee surveyflights,for a total of six nestsfound in the area. Of the
sixJabiru nestswe found, one had large young,three had medium sized
young, one was apparently incubating, and one was standingon a com-
plete nest platform with no eggsor young in the nest.
In Honduras, we found a total of four Jabiru Stork nests (three with
chicks 1-2-mo old, one without eggsor chicks), two of which fell within
our surveystrips, and two of which were seen while in transit between
surveytransectsand Puerto Lempira.
DISCUSSION

The densityof waterbirdsin Honduraswasabout half that estimatedin

Nicaragua(Table 1). This wastrue for all of the waterbirdspeciesindi-
vidually,with exceptionsonly for whistling ducks (Dendrocygna spp.),
Blue-winged Teal (Arias discors),White Ibises, and small white herons.
Densitiesof Jabiru Storksin Honduras were lessthan 33% thosein Nica-
ragua, thoseof Wood Storkslessthan 15%, and thoseof MuscovyDucks
(Cairina moschata)less than 25%.
Differencesin densitieswere unlikely to have been due to differing
methodologies,becausethe surveyflights in both locationswere per-
formed at similaraltitudes,usingthe sametechniquesfor estimatingstrip
widths,orienting flight transects,counting,recording,and analysis.One
of the two observersin Honduras (PCF) was also an observer in the Nica-
raguan surveys.The variety of habitat and water depths traversed,and
wide geographicaldistribution of transectswasquite similar in both sur-
vey efforts.
Nonetheless,it is not clear that the surveyresultsin the two areasare
due to inherentlydifferent carryingcapacitiesof the two landscapes. The
surveys were performedin different yearswith quite differentweatherpat-
terns, and even at slightlydifferent times of year (late Februaryin Hon-
duras,early March in Nicaragua). Interviewswith residentsin Nicaragua
in 1992 suggestedthat weather and surfacewater conditionswere normal
for that time of year, with the low rainfall typicalof the dry season.Simi-
lar interviewsin Hondurasin 1994 werejust the opposite.Fishermen,lo-
cal pilots, church workers,and other residentsseemedto agree that the
winter of 1994 had been abnormallywet, and that a typicaldry season
had not becomeestablished.Sincewaterbirddensitiesin other large sub-
tropical and tropical wetland regionsin the Yucatan (Lopez Ornat and
Ramo 1992), and the Everglades(Bancroft and Sawicki1995, Hoffman et
al. 1990) are usuallymuch lower during wet conditions,it seemslikely
that weather pattern alone could explain the apparent differences
betweenNicaraguanand Honduran surveys.
We alsosawmore Blue-wingedTeal in the Honduran surveysthan were
counted in Nicaragua.This is likely due to samplingerror, becauseBlue-
wingedTeal are usuallyencounteredin flocksand the total number seen
in Honduraswasrelativelysmall (20 individuals).In Nicaragua,we in fact
Vol.68,No.9 Wetland
BirdsofHonduras
andNicaragua [293

did see severallarge flocks of teal in the lagoons,but did not count them
becausethey did not fall within the surveystrips.
This surveyconfirms the general speciesabundance pattern found in
wetlands throughout Central America, with geographic and numeric
dominance by Great Egrets and other herons, and much smaller num-
bers of Wood Storks, White Ibises, and other tactile feeders (Correa 1992,
Frederickand Spalding1992, Ogden et al. 1988). This pattern is distinct
from that found in the wetlands of both the southeastern United States
and the northeastern coast and interior wet plains of South America,
where ibisestend to dominate the wading bird biomass(Bancroft and
Sawicki 1995, Bildstein 1993, Kushlan and White 1977, Morales and Pa-
checo 1986, Spaans1990).
This studyconfirms that Jabiru Storksbreed in substantialnumbers in
the study area, contraMonroe (1968). Cottea and Luthin (1988) found
0.06JabiruStorks/km
9 in the Usamacinta
Delta wetlandsof Mexico,
about half the density found in Nicaragua, and nearly the same den-
sity as Honduras. Villareal (1996) found between 0.44 and 4.8 indi-
viduals/km
9 in MataRedondoLagoonin CostaRicabetween
January
and April of 1993. Similarly, Brooks (1991, in Hancock et al. 1992)
reported4.38individuals/kin
9 in aggregations
in the Chacoregionof
Paraguay. Gonzalez (1993) and Thomas (1985) found 0.028 and 2.0
nestingpairs/km
9, respectively,
in the Venezuelan
Llanos.The den-
sitiesof adults we saw feeding on the marsh surface are suggestiveof
0.03 and 0.08 pairs/kin
9. Thusdensities
of nestingpairsin the La
Mosquitia/Miskito Coast region may be similar to those in the Llanos of
Venezuela.
Correa and Luthin (1988) suggestedthat the Jabiru Stork population
of Central America may follow seasonalmovements among Mexico,
Belize,and Guatemala.The very high concentrationsfound by Ogden et
al. (1988), Villareal (1996), and Cottea and Luthin (1988) suggestthat
large seasonalinfluxes do occur in various parts of Central America.
Hancock et al. (1992) estimatedthe entire Central American population
of Jabirus to be no more than 250 individuals.The 98 individualswe saw
in our incomplete surveyssuggeststhat the Atlantic coastwetlands of
Nicaragua and Honduras host a large portion of the Central American
population.

ACKNOWLEDGMENTS

This work wassupportedby Lighthawk, Inc. (Honduras and Nicaragua studies)and the
CaribbeanConservationCorporation (Nicaragua),with grantsfrom the U.S. Agencyfor In-
ternationalDevelopment,the Liz Claiborne and Art Ortenberg Foundations,and the Mo-
riah Fund. Our work wassupportedlogisticallyby the staff and resourcesof IRENA (New
name, MARENA, the Nicaraguan Ministry of Natural Resources)and MIKUPIA in Nicara-
gua, and by Lighthawk'sLa Mosquitiaprojectand MOPAWI in Honduras.We thank B. Row-
ell for assistance
with photos and data recording in Honduras. We are indebted to B. Stith
and L. Pearlsteinfor help with GIS applicationsassociatedwith this project. This isJournal
SeriesR-05370of the FloridaAgriculturalExperimentStation.
294] P. c. Frederick
et al. J.Field
Ornithol.
Spring 1997

LITERATURE CITED

ALLEN,R.P. 1942. The RoseateSpoonbill. National Audubon Society,New York, New York.
B•kNCROFT, G. T., XNDR.J. S^•cgI. 1995. The distribution and abundance of wading birds
relativeto hydrologicpatternsin the Water ConservationAreasof the Everglades.Final
report to the South Florida Water Management District. West Palm Beach, Florida.
BILDSTEIN, K. L. 1993. White Ibis, wetland wanderer. Smithsonian Institution Press, Wash-
ington, D.C.
BJORK,R. B., 2•NDG. V. N. Pox&•I.L. 1994. Relationshipsbetween hydrologicalconditions
and quality and quantity of foraging habitat for RoseateSpoonbillsand other wading
birds in the C-111 basin. EvergladesNational Park, Homestead, Florida.
C•XM•CHO, M. G. 1983. Notes on aquatic birds of Nicaragua,Jabiru mycteria.Inst. Nicarag.
Rec. Nat. Ambient. 14 pp.
CORRF&,J. 1992. Statusof aquaticbirds in the coastalwetlandsof the Yucatanpeninsula.
M.Sc. thesis,Universityof Newcastleupon Tyne, England. 110 pp.
, ANDC. S. LUTHIN. 1988. Proposalfor the conservationof theJabiru Stork in South-
eastMexico. Pp. 607-615, in Memoria del Simposiosobrela Ecologiay Conservacion
del Delta de los Rios Usamacintay Grijalva.Instituto Nacional de Investigacionsobre
RecursosBioticos,Division Regional, Tabasco,Mexico.
GONZALEZ, J. A. 1993. Contribucion al estudio de la ecologia de las ciguenos (Fam. Cico-
niidae) en los Llanos de Venezuela. PhD dissertation,Universidad Complutense de
Madrid, Madrid, Spain.
FREDERICK, P. C., ANDS. M. MCGEHEE.1994. Wading bird use of wastewaterwetlandsin
central Florida, USA. Colonial Waterbirds 17:50-59.
•, ANDM. G. SPALDING.1992. Factorsaffecting reproductivesuccessof wading birds
(Ciconiformes)in the Evergladesecosystem.Pp. 569-691, in S. Davisand J. C. Ogden,
eds. Everglades:the Ecosystemand its Restoration. St. Lucie Press,Del Ray Beach,
Florida.
HANCOCK, J. A., J. A. KUSHLAN,
ANDM.P. I•HL. 1992. Storks,Ibises,and Spoonbillsof the
World. AcademicPress,San Diego, California.
HOFFMAN, W., G. T. BANCROFT, ANDR. j. s^wicKi. 1990. Wading bird populationsand dis-
tributionsin the Water ConservationAreasof the Everglades:1985-1988. SouthFlorida
Water Management District, West Palm Beach, Florida.
HOWELL,
T. R. 1971. An ecologicalstudyof the birdsof the lowlandpine savannaand
adjacent rain forest in northeastern Nicaragua. Living Bird 10:185-242.
HUBER,W. 1932. Birds collectedin northeasternNicaragua in 1922. Proc. Acad. Nat. Sci-
ence of Philadelphia84:205-249.
J•LKS,H. 1991. The distribution of foraging wading birds in relation to the physicaland
biological characteristicsof freshwater wetlands in southwesternFlorida. M.Sc. thesis,
University of Florida, Gainesville,Florida.
KUSHLAN, J. A., ANDD. A. WHITE. 1977. Nesting wading bird populationsin southern
Florida. Fla. Sci. 40:65-72.
LOPEZORNATA., ANDC. RAMO. 1992. Colonial waterbird populations in the Sian Ka'an
biospherereserve (Quintana Roo, Mexico). Wilson Bulletin 104:501-515.
MONROE,B. L., JR. 1968. A distributionalsurveyof the birds of Honduras.Ornithological
MonographsNo. 7, American OrnithologistsUnion, Washington,D.C.
MORALES,G., ANDJ. PACHECO.1986. Effectsof diking of a Venezuelan savannaon avian
habitat, speciesdiversity,energyflow, and mineral flow through wading birds. Colonial
Waterbirds 9:236-242.
NORTON-GRIFFITHS,
n. 1975. Countinganimals.Handbookno. 1. Nairobi, Kenya:African
Wildlife Leadership Foundation.
O(;•)EN,J. C., C. E. KNO•)ER,AN•)A. SPRt'STJR.1988. Colonial waterbird populationsin the
Usamacinta delta, Mexico. Pp. 565-605, in Ecologia de los Rios Usamacintay Grijalva.
Instituto Nacional de los Investigacionessobre RecursosBioticos,Tabasco,Mexico.
RICHMOND,C. W. 1893. Notes on a collection from eastern Nicaragua and the Rio Frio,
CostaRica, with a descriptionof a supposednew Trogon. Proc.Nat. Mus. 16:479-532.
S(;LATER, P. L. 1873. Additionsto the list of birdsof Nicaragua.Ibis 3:372-373.
vol.68,No.9 Wetland
BirdsofHonduras
andNicaragua [295

--, ANDO. SALkqN.

1867. Listof birdscollectedfrom BlewfieldsRiver,MosquitoCoast.
Proc. Zool. Soc. Lond., 278-280.
SCOTT,D. h., ANDM. CARBONEI.I.. 1986. A directory of neotropicalwetlands.International
WaterfowlResearchBureau, Slimbridge,United Kingdom.
SP&XNS,
A. L. 1990. Problemsin assessing trendsin breedingpopulationsof ScarletIbises
and other ciconiiform birds. Pp. 1-6, in P. C. Frederick, L. G. Morales, A. L. Spaans,
and C. S. Luthin, eds. The Scarlet Ibis: Status, conservation and recent research. Inter-
national Waterfowland WetlandsResearchBureau SpecialPublication4/11,Slimbridge,
United Kingdom.
STILES, F. G., ANDA. F. SKUTCH.1989. A guide to the birds of CostaRica. Cornell Univer-
sit), Press,Ithaca, New York.
THOM.kS, B.T. 1985. Coexistence and behavior differences amont the three western hemi-
spherestorks.Pp. 921-931, in P. A. Buckley,M. S. Foster,E. S. Morton, R. S. Ridgley,
and F. G. Buckley,eds. Neotropical Ornithology, OrnithologicalMonograph No. 36,
American Ornithologists'Union, Washington,D.C.
VILL•RE^•,J. 1996. Estadoactual, ecologiatrofica y uso de habitat delJabiru (Jabflu myc-
teria) en tres humedalesde la cuencabaja del Tempisque,CostaRica. MS thesis,Uni-
versidad Nacional, Herealia, Costa Rica.
WII,L, T. 1991. Birds of a severelyhurricane-damagedAtlantic coastrain forest in Nicara-
gua. Biotropica23:497-507.
Received14 Mar. 1996; accepted28 Aug. 1996.