See discussions, stats, and author profiles for this publication
at: http://www.researchgate.net/publication/271072766Geographic and
seasonal distribution of alittle-known Brazilian endemic
rail(Aramides mangle) inferred from ocurrencerecords and ecological
niche modelingARTICLEinTHE WILSON JOURNAL OF ORNITHOLOGY DECEMBER
2014Impact Factor: 0.57 DOI:
10.1676/13-165.1CITATION1DOWNLOADS31VIEWS1094 AUTHORS:Rafael Sobral
MarcondesLouisiana State University6 PUBLICATIONS 2 CITATIONS SEE
PROFILEGlaucia Del-RioLouisiana State University6 PUBLICATIONS 1
CITATION SEE PROFILEMarco Antonio RgoLouisiana State University16
PUBLICATIONS 13 CITATIONS SEE PROFILELus Fbio SilveiraUniversity of
So Paulo196 PUBLICATIONS 362 CITATIONS SEE PROFILEAvailable from:
Lus Fbio SilveiraRetrieved on: 06 August 2015BioOne sees
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research.Geographic and seasonal distribution of a little-known
Brazilian endemicrail (Aramides mangle) inferred from ocurrence
records and ecologicalniche modelingAuthor(s): Rafael Sobral
Marcondes, Glaucia Del-Rio, Marco Antonio Rego, and Lus
FbioSilveiraSource: The Wilson Journal of Ornithology,
126(4):663-672. 2014.Published By: The Wilson Ornithological
SocietyDOI: http://dx.doi.org/10.1676/13-165.1URL:
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holder.GEOGRAPHICANDSEASONALDISTRIBUTIONOFALITTLE-KNOWNBRAZILIANENDEMICRAIL(ARAMIDESMANGLE)INFERREDFROMOCURRENCERECORDSANDECOLOGICALNICHEMODELINGRAFAELSOBRALMARCONDES,1,2,3,4GLAUCIADEL-RIO,1,2MARCOANTONIOREGO,1ANDLUISFABIOSILVEIRA1ABSTRACT.Regional,
intratropical avian migrations have rarelybeen studied. Here, we
employ an occurrencerecords reviewandecologicalnichemodeling
toolstotestthehypothesis thatanunderstudied
Neotropicalbird,theLittleWood-Rail (Aramides mangle, Rallidae),
seasonally migrates between the humid Atlantic coast and the dry
Caatinga biomeof interior northeastern Brazil. We divided records
geographically between coastal and inland, and temporally between
wet/breeding and dry/non-breeding seasons. Coastal records peak
when inland records are fewest and vice-versa, andindependence
between season and region in which records were made was
statistically rejected. However, ecological nichemodeling shows
that coastal regions are suitable habitats for A. mangle
year-round, and models built with records from eachseason were
considered statistically equivalent. It seems that this species
neither performs erratic, unpredictable
movementsnortypicalavianto-and-fromigration.
Instead,itundergoesperiodicalexpansionofitsrangeandecologicalnichetoincludetheCaatinga,whereitbreeds,inadditiontothecoast.Itmightbecounterintuitivethataspeciescanoccupytwoseeminglysodisparatehabitats,
but railsingeneral areknowntobeveryadaptableandhavewideecological
niches.Further study is needed in order to understand the exact
nature of this species movements and the life-cycle of
individualbirds. But given that most studies of avian movements
have focused on temperate species, it is likely that common
modelsof avian migratory behavior will not easily apply to A.
mangle nor to other Neotropical species. Received 7 October
2013.Accepted11May2014.Keywords: Caatinga,intratropicalmigration,
Rallidae,range.Migrationisoneofthemostremarkableofallavianbehaviors.
Themostemblematicmigratorymovementsarelatitudinal
long-distanceseasonalmigrations frombreeding to wintering
groundsand back (Cox 1985, Berthold 2002, Newton2008). However,
theyrepresent onlyanextremeinacontinuumof
variationinbirdmovements,which also includes everyday routine
movements,dispersive, irruptive, and nomadic behaviors(Dingle 1996,
Berthold 2002, Newton 2008).Nevertheless, most studies of bird
movementshave focused on these classic to-and-frointercontinental
migrations, especially in thetemperate regions of the Northern
Hemisphere.Regional, non-latitudinal, intra-tropical
avianmigrationsintheNeotropicsareamongtheleaststudied forms of bird
movement (Levey and Stiles1992, Chesser 1994, Jahn et al. 2006,
Sekercioglu2010,Davenportetal. 2012).Ideal approaches for studying
bird migrationincludeyear-roundsurveys of specificlocalities(Remsen
and Parker 1990) or tracking ofindividuals through their migratory
route (e.g.,Davenport et al. 2012).
Thedataandtechniquesnecessaryforthesekindsofstudies,however,arenonexistent
or unavailable for most Neotropicalspeciesandresearchers. Inthat
case, analysisofseasonalvariationingeographicdistributionmayprovide
circumstantial evidence for migration(RemsenandParker1990),
anapproachsuccess-fully employed by several authors
studyingNeotropical migrant birds (e.g., Remsen
andParker1990,MarantzandRemsen1991,Remsen2001, MachadoandSilveira
2010, Lopes et al.2011).Ecologicalnichemodels(ENMs)arepowerfultools
for
statisticallytestingputativedifferencesbetweenbreedingandwinteringrangesobtainedfrom
distributional and seasonal data (e.g.,Nakazawaet al. 2004).
ENMscanbeuseful forclarifying species distribution, for
conservationefforts and investments and other studies
ofevolutionaryandecological nature(e.g., Loiselleet al. 2003, Kozak
et al. 2008, Phillips et al. 2009).1SecaodeAves,Museu
deZoologiadaUniversidadedeSaoPaulo.CaixaPostal42494,SaoPaulo,SP,04218-970,Brazil.2Pos-Graduacao,
Departamento de Zoologia, Instituto deBiociencias, Universidade de
Sao Paulo. Caixa Postal11461,SaoPaulo,SP,05422-970, Brazil.3Current
address: MuseumofNatural Sciences, Louisi-ana State University. 119
Foster Hall, Baton
Rouge,Louisiana70803,USA.4Correspondingauthor;e-mail:[email protected](4):663672,2014663Theyareproducedbycombiningspeciesoccur-rencedatawithbroad-scaleenvironmental
layersin a Geographic Information System (GIS)environment and
allowquantification of nichevariationat
anunprecedentedscale(Warrenet al.2008). In the context of migration
studies,occurrencemodels basedondatafromdifferentseasons can,
unlike simple point locality maps, becompared and tested for their
differences using anobjective and quantitative statistical
framework(Warren et al. 2008, McCormack et al. 2010,Gibbons et al.
2011, Marini et al. 2012). ENManalyses have been successfully
employed tostudyseasonal variationintherangeof
putativemigratoryspecies,suchasSwainsonsFlycatcher(Myiarchus
swainsoni, Joseph and Stockwell2000), Black-fronted Ground-Tyrant
(Muscisax-icolafrontalis, Gibbons et al. 2011), andCock-tailed
Tyrant (Alectrurus tricolor, Marini et al.2012). Furthermore, they
allowtesting whethermigrant birds are seasonally switching
theirniches(nicheswitchers), ortrackingthesamesuitable conditions
(niche followers; Naka-zawa et al. 2004). There are various
advantages inproducing predicted distributions and
graphicrepresentations of habitat suitability, but
statisticaltestsareessential tointerpret
thesignificanceofthesepatterns(Warrenetal. 2008).Rails (Rallidae)
aresmall- tomedium-bodied,weak-flying,
cosmopolitanbirdsfoundmainlyinaquatic and semi-aquatic habitats,
such as marshes,mangroves, and margins of rivers and
lakes.Migratory, sporadic, and erratic movements arecommonin
thisfamily (Ripley 1977, RemsenandParker1990, Sick1997,
Taylor1998).
TheLittleWood-Rail(Aramidesmangle)isamongthemostpoorly-known
Neotropical rails. It is usuallyconsideredexclusivelyor mainlya
coastal bird,occurringinmangroves andadjacent habitats
inmostoftheBraziliancoast(Pinto1938,Hellmayrand Conover 1942,
Ripley 1977, Taylor 1996, Sick1997, Taylor 1998, van Perlo 2009).
However,someinlandrecords, mainlyinthedryandveryseasonal Caatinga
biome of northeastern Brazil(Prado 2003, Leal et al. 2005), have
also beenreported, without muchdetail (e.g.,
Pinto1938,HellmayrandConover1942,Sick1997).Littleisknownaboutthedistribution,
habitatpreferences,conservation status, breeding, and feeding
habits ofthisrarelyobservedspecies(Ripley1977, Taylor1996,Taylor
1998),whichiscurrentlyconsideredData Deficient in the Brazilian
list of threatenedfauna (Machado et al. 2005).The inland occurrence
of A. mangle in theCaatinga of northeastern Brazil was first
de-scribedinsomedetailbyLimaetal.(2005),whounexpectedlyobservedit
nestinginthemunici-pality of Jeremoabo (9u 519 S, 38u 389 W), state
ofBahia, .200kmfromthecoast.Furtherinforma-tion on its occurrence
in the Caatinga wasprovided by Redies (2010), who recorded it
inItapaje(3u 489 S, 39u 289 W), state of Ceara, 60 kmfrom the
nearest coast. Even though no nests weredirectlyseen, observationof
half-grownindivid-uals and territorial behavior indicated that
thespecies was reproducing in the area (Redies2010). Both Redies
(2010) and Lima et al.s(2005)
observationsweremadeduringtherainyseason.These records contradict
the traditional viewthat A.
mangleisamangroveorswampy-habitatspecialist, but it
iscurrentlyunknownwhetheritis a permanent inland resident. Redies
(2010)suggestedthatit mightperformmigratorymove-ments between the
Caatinga and more humidhabitats, such as coastal mangroves or
inlandswamps. Here, we reviewthe geographical andseasonal
distribution of A. mangle, based onpoint-localityrecords,
toexaminethehypothesisthat A. mangle performs inlandcoastal
migration.Inaddition, wetest ifENMsbasedoninlandorcoastal records
represent equivalent climaticconditions. Suchanapproachprovides
informa-tiononthespeciespotentialdistributionineachseason, andsheds
light onwhether thespeciesmovements are because of its ability to
trackspecificclimaticconditionswithinitsrange, thatis, if its
ecological niche undergoes
seasonalchange.METHODSRecordsofAramidesmangle.Wecompiledatotal of
99records of A. mangle, obtainedfromseveral sources: 1) specimens
fromsevenorni-thological collections in Brazil, Europe, and
NorthAmerica (21 records; Table1); 2) photographs,videos or sound
recordings (45 records) depositedintheonlinedatabases Wiki Aves
(WA; www.wikiaves.com.br), Xeno-Canto(XC; www.xeno-canto.org) and
Internet Bird Collection (IBC;ibc.lynxeds.com);
3)awideliteraturereview(37records);
and4)unpublishedfieldrecordsbytheauthors or fromtrustedobservers
(five records;seeAcknowledgments).Thesumofrecordsfromeach of the
different sources is .99 because somerecords were obtained through
more than one664 THEWILSONJOURNALOFORNITHOLOGY
NVol.126,No.4,December2014source. Acompletelist
ofrecordsandassociatedinformation is available upon request
fromthecorresponding author. Five additional museumcollections
(Table1) were visited or consulted,but only held specimens of A.
mangle withoutpreciselocalityinformation,whichthuscouldnotbeusedfor
our analyses. Wecarefullycheckedeach record, especially those
obtained fromtheonline databases, and discarded those
whoseidentification as A. mangle was doubtful or
clearlymistaken,andifnecessarycontactedtheirauthorsto confirmthe
circumstances under which theywere made. Data were collected until
26 April2013.Records in the same locality and month(including the
same month in different years)were condensed into a single record.
For eachrecordwerecordedinformationonlocalityand,when available,
date and reproductive activity,which we defined as observation of
an active nest,eggs,and/orchicks.Weclassifiedpoint-localitiesinto
coastal and inland. Coastal recordswere defined as those made
,10kmfromthecoast and/or associated with typically
coastalenvironments, such as beaches, dunes, mangroves,estuaries
andrestingas. All others were consid-ered inland records. Based on
the observedseasonal variation in distribution (see Results),we
divided records into two seasons: wet/breeding(FebJul) and
dry/non-breeding (AugJan). Wechose this temporal division in order
to include
inthewet/breedingseasontheperiod(approximate-lyfromFebMay)whenthenumberofCaatingarecords
is greater (see below), as well as
themonthswhenreproductiveactivitywasnotedinthatregion(MayJul).AssociationbetweenSeasonal
andGeographi-cal Distributions.Toqualitativelyanalyzesea-sonal
variation in the distribution, we builtseparate distributionmaps
for eachseason, andplottedagainst themonththenumber of
inlandandcoastal records, aswell asthedistancefromthe coast for
each record. If A. mangle indeedmigrates, we expect that there will
be seasonalconcentration of records in some areas of themaps
andplots (Lopes et al. 2011). Inorder tostatisticallytest
theassociationbetweenseasonal(wet/breedingor dry/non-breeding)
andregional(coastal or inland) distributionof A. mangle, webuilt
atwo-waycontingencytable,
withthetotalrecordsforeachclass,andperformedaPearsonschi-squaredtest,anditslikelihoodalternative,G-test,
totest the null hypothesis that seasonandregionwerenot
associatedwitheachother.
WeusedtheStatspackageforchi-squareanalysisandtheDeducer packagefor
G-test (Fellows2012) in Rsoftware V. 3.0.2 (RDevelopmentCore Team
2012). A rejection of the nullhypothesis would indicate that there
is anassociation between season and region andprovide support for
the idea that A. mangleperformsseasonalmovements.TABLE1.
Locationandacronymsofornithologicalcollectionsconsultedinthisstudy.CollectionsthatheldspecimensofAramidesmanglewithcompletelocalitydata:FieldMuseumofNaturalHistory
FMNH Chicago,USAMuseudeBiologiaProf.MelloLeitao MBML
SantaTeresa,BrazilMuseudeZoologiadaUniversidadedeSaoPaulo MZUSP
SaoPaulo,BrazilMuseuNacionaldoRiodeJaneiro MNRJ RiodeJaneiro,
BrazilMuseuParaenseEm lioGoeldi MPEG Belem,
BrazilMuseumNationaldHistoireNaturelle MNHN
Paris,FranceMuseumofComparativeZoology,HarvardUniversity MCZ
Cambridge,USACollectionsthatonlyheldspecimensofAramidesmanglewithnoorveryimpreciselocalitydata:AcademyofNaturalSciences
ANSP Philadelphia, USAAmericanMuseumofNaturalHistory AMNH
NewYork,USAMuseumfurNaturkunde ZMB
Berlin,GermanyNationalMuseumofNaturalHistory USNM
WashingtonD.C.,USANaturalHistoryMuseum BMNH
Tring,UKCollectionsthatheldnospecimensofAramidesmangle:CarnegieMuseumofNaturalHistory
CMNH Pittsburgh,USALouisianaStateUniversityMuseumofNaturalSciences
LSUMNS BatonRouge,USAMuseudeHistoriaNaturaldoCapaodaImbuia MHNCI
Curitiba,BrazilYalePeabodyMuseum YPM NewHaven,USAMarcondesetal.
NMOVEMENTSOFALITTLE-KNOWNNEOTROPICALRAIL
665EcologicalNicheModeling.Weobtainedtwodistribution models for A.
mangle using themaximumentropyalgorithm, implementedintheMaxent
3.3.3e software (Phillips et al. 2006). Thismethod estimates the
environmental suitabilityfor a species, based on a list of
georeferencedoccurrencelocalitiesandaseriesofenvironmen-tal
variables used as predictors. To
investigatepossibleseasonalmovements, wecreatedanichemodel for
thedry/non-breedingseason, andonefor the wet/breeding season. The
models weredeveloped using one topographic (altitude) andnine
climatic (temperature and precipitation)variables extracted fromthe
WorldClimdatabase,at a 2.5arc-seconds (,5 km) resolution (Hijmanset
al. 2005; www.worldclim.org). We did notinclude vegetation layers
because these have
beenshowntobesignificantlycorrelatedwithclimate(Schultz and Halpert
1993, Wang et al. 2001, Ichiiet al. 2002). To create a subset of
minimallycorrelated climatic variables, we constructed
apair-wisecorrelationmatrixandchosethosewithcoefficients of
correlation,0.85. The variablesused for the maximumentropy analysis
were:Annual Mean Temperature (BIO 1); Isothermality(BIO 3);
Temperature Seasonality (BIO 4);MinimumTemperature of Coldest Month
(BIO6); Temperature Annual Range (BIO7); MeanTemperature of Driest
Quarter (BIO9); MeanTemperature of Coldest Quarter (BIO
11);PrecipitationSeasonality(BIO15)andPrecipita-tion of the Warmest
Quarter (BIO 18). Toevaluatethe ENMs performance, we
employedtheareaunderthecurve(AUC)approach, whichhas been accepted
as the standard measure forassessing the accuracy of species
distributionmodels (Jimenez-Valverde 2012). AUC values.0.75
indicatemodels potentiallyuseful(Elithetal. 2006). We expect that,
if A. mangle indeedperforms seasonal movements, in
occurrenceprobability maps built from ENMs for eachseason,
regionswithhighoccurrenceprobabilityin the wet/breeding season will
have a lowerprobability in the dry/non-breeding season,
andviceversa.Wecomparedthelogisticmodeloutputsusinganicheequivalencytest
(Warrenet al. 2008)onENMTools 1.3software (Warren et al. 2009).This
test creates pseudoreplicated datasets byrandomly partitioning the
pooled sets of totaloccurrences. For each pseudoreplicate,
nichemodels arecreated,andthesearecomparedusingthe similarity
measures I and Schoeners D(Warrenetal.
2008).Werepeatedthisprocedure100 times, creating a null
distribution of thesesimilarityindices.
Weassessedwhetherdistribu-tion models for both seasons differ
statistically bycomparingobserved I andDvalues with thosecontained
in the null distribution. A one-tailed testallowedustotest
whetherobservedvaluesweregreater than expected by chance alone. A
rejectionofthenullhypothesisthatnichesforeachseasonare equivalent
would support the idea that A.mangle migrates. However, the
opposite result,equivalency of niches, would not
necessarilyrefutethishypothesis, becausethespeciescouldbe a niche
follower. In any case, the results of
thistestwillhelptoassessthebiologicalsignificanceofthespeciesmovements(orlackthereof).RESULTSGeographic
Distribution of Aramides mangle.The records compiled come from70
differentlocalities(48coastal and22inland). TheyshowthatA.
mangleisdistributedalmostcontinuouslyin the coast fromAnjos (0u 299
S, 48u 369 W,in Marajo Island, Para), south to Paranagua(25u 359 S,
48u 309 W, in Parana). There is onesingle record outside Brazil,
fromthe coast ofFrench Guiana (Ingels et al. 2011), almost 800
kmnorthwest of Anjos, withnorecordsinbetween.Among inland
localities, all except four are
withintheCaatinga.Thespeciesiswidelydistributedinthenorthernpartofthisbiome,fromCearasouthtonorthernBahia.
Inlandrecordsdonotseemtobeassociatedspecificallytomarshesoranyotherkindofaquaticorsemi-aquatichabitats.Seasonal
Distribution of Aramides mangle.Weobtained84records
withdate-relatedinfor-mation that we could use to analyze
seasonalvariation in the distribution. A. mangle occursyear-roundat
thecoast andinlandinall monthsexcept October toDecember (Figs.13).
Inlandrecords are more numerous fromFebruary toApril, whencoastal
records are fewest (Fig. 2).All except
onerecord.100kmfromthecoastweremadebetweenFebruaryandJune(Fig.
3).Conversely, the coastal records peak between JulyandOctober,
whentheinlandrecordsarefewest.ReproductiveactivitywasrecordedbetweenMayand
July in the Caatinga (Lima et al. 2005, Redies2010), andinOctober
andNovember inIguabaGrande, in the coast of Rio de Janeiro
state(several photos onWiki Aves, e.g., WA73905and WA 80181, and A.
Navegantes, pers. comm.).666 THEWILSONJOURNALOFORNITHOLOGY
NVol.126,No.4,December2014Thenull hypothesisthat
seasonandregioninwhich records were made are independent
wasrejected(x215 4.58,P 5 0.031;G 5 5.91,P 50.010), supporting the
alternative hypothesis ofdifferential occupation of space in time
by A.mangle.Ecological Niche Modeling.Figure4 showsthe niche
suitability for wet/breeding and dry/non-breeding seasons based on
records of A. mangle foreachseason. TheobtainedAUCvalues for
wet/breedinganddry/non-breedingwererespectively0.960 and 0.930,
pointing to a high performance ofboth models (Elith et al.
2006).Coastal regionsofBrazil appear as suitableareas for A.
mangleinboth seasons. The coastal swath of suitable habitatis wider
in the northeast-facing segment of theSouthAmericancoast
(betweenthestatesofRioGrande do Norte and Amapa). In the
wet/breedingseason, there is an expansion of the suitable
habitatintotheinterior of thecontinent innortheasternBrazil, to
include considerable portions of Caatingain almost the entire
states of Para ba, Alagoas,Sergipe, andCeara.In the equivalency
test results, the I nulldistribution percentiles values were
I50.91I950.97;andtheDrandomdistribution,D50.69D950.83. Niche
similarity indices generated for ourmodels wereI 50.92andD50.73,
thereforefallingwithinthe595percentilesof eachdistri-bution. Based
on these results, we cannot reject thenullhypothesis of equivalency
between niches.DISCUSSIONWe show, contraryto previous
references(e.g.,Pinto1938, HellmayrandConover1942, RipleyFIG. 1.
Distribution of Aramides mangle in the dry/non-breeding season
(AugJan; left) and in the wet/breeding season(FebJul;right). White
symbols denote observation of reproductive activity.The
Caatingabiomeisshown indarker gray.FIG. 2. Number of inland
(continuous line withtriangles)andcoastal
(dashedlinewithcircles)recordsofAramidesmanglethroughtheyear.
Opensymbolsdenoteoccurrenceofreproductiveactivity.Marcondesetal.
NMOVEMENTSOFALITTLE-KNOWNNEOTROPICALRAIL 6671977, Taylor1996,
Sick1997, Taylor1998,
vanPerlo2009)implyingthatA.mangleismainlyorexclusively a coastal
bird, that the inland, dryCaatinga of northeastern Brazil is a
regular part ofthis species range. However, it is not a
permanentresident there. Instead, it seems that thespeciesinhabits
more humid coastal regions of Brazilyear-round and periodically
expands its geograph-ical distributionandecological
nichetoincludethe Caatinga as well, in spite of apparent
disparitybetween the habitats available in these
tworegions.Followingseveral authors whoexaminedsea-sonal
variationof species distributions
basedonreviewofrecords(e.g.,RemsenandParker1990,Marantz and Remsen
1991, Remsen 2001,MachadoandSilveira2010, Lopes et al. 2011),the
clear concentration of Caatinga
recordsbetweenFebruaryandJune(rainyseasoninthisregion)isstrongindirect
evidencethat therangeof A. mangle is seasonally variable. This is
furthersupportedbythestatistical rejectionof thenullhypothesis that
season and region are independentvariables. Thisseasonal
concentrationof recordsargues against theideathat this species
occur-FIG. 3. Distance (km) from the nearest coast of recordsof
Aramides mangle through the year. Open symbols denotereproductive
activity. For this figure only, all
coastalrecordswereconsideredasbeing0kmfromthecoast.FIG. 4. Habitat
suitability for Aramides mangle, asinferred through ecological
nichemodeling basedon records fromthedry/non-breedingseason(AugJan;
left) andfromthewet/breedingseason(FebJul; right). Redindicates
highestsuitability;blueindicateslowestsuitability.668
THEWILSONJOURNALOFORNITHOLOGY
NVol.126,No.4,December2014renceintheCaatingacouldberesultofsporadic,irruptive,
or nomadic movements. Such
move-mentsarecommonintheRallidae(Ripley1977,Sick 1997, Taylor
1998), and Redies (2010)impliedthat thiscouldbethecaseofA.
mangle,butthesemovementsarespatiallyandtemporallyunpredictable
(Berthold 2002, Newton 2008)unlikethepatternobservedhere.
Ourresultsalsodonot provide clear support for the hypothesisthat A.
mangleperformsclassical avianseasonalto-and-fro migration,
becauseinthis type ofmigration all, or most, birds of a
speciesperiodically move from resting to breedinggrounds
andback(Dingle1996, Berthold2002,Newton 2008). In contrast, even
though A. mangleapparentlyvacatestheCaatingafromOctobertoNovember,
it maintains presence on the coastyear-round.Our results (Figs. 13)
show that the geograph-ical range of this species is not switched
from oneseason to another but seems to undergo periodicalexpansion
and retraction. This is evident fromENManalyses.
Thehabitatsuitabilitymapsbuiltfromwet/breeding season records point
to anexpansion of the predicted niche toward theCaatinga,
incomparisontothemodel built fromdry/non-breeding season data
(Fig.4). Coastalregions with high suitability in the
dry/non-breedingseasonretainthehighsuitabilityinthewet/breedingseason,
contrarytowhat wouldbeexpectedifA.mangleperformedclassicto-and-fro
migration and completely switched itsdistribution and niche
seasonally. Furthermore,thehypothesis that niches
fromeachseasonareequivalent couldnot berejected, andA. manglethus
fits the concept of niche follower,proposedbyNakazawaet al. (2004).
Inspiteofnotable seasonal variation in its geographicalrange, our
results suggest that A.
mangleeffec-tivelytracksasingleclimateregimethroughthecourseof
theyear. It isperhapscounterintuitivethat such different
environments as the dryCaatinga andmangroves
alongthehumidcoastcouldbeconsideredpart of thesameecologicalniche.
It shouldbetakenintoaccount, however,that rails aregeneralist
birds. Most
areomnivo-rousandplasticintheirhabitatrequirements;anyrail is prone
to be found in atypical or (seemingly)suboptimal habitats (Taylor
1996, 1998; Sick1997).Insuchcontext, ourdemonstrationthatA.mangle
is able to occupy, at least seasonally, suchapparently disparate
habitats is not that
surprising;awideecologicalnicheistypicalforrails.Eventhoughcoastal
recordsof A. manglearemore than twice as numerous as Caatinga
records,twoofthreeknowninstancesofbreedingactivityof A. mangle are
fromthe Caatinga. Redies(2010)observationsofA. mangleinItapaje
weremadebetweenJanuaryandAugust, whichcoin-cideswiththerainiest
monthsintheregion, andLima et al.s (2005) observation of one active
nestin Jeremoabo was in May, the second rainiestmonth there
(precipitation data from Grieser2006), consistent withthefact that
avianrepro-ductionintheCaatingaisstronglydependent
ontherainyseason(Santos2004,Roosetal. 2006).The third record of
reproduction is from the muni-cipalityof IguabaGrande(22u 509 S,
42u 119 W),inthecoast of thestateof RiodeJaneiro, inaregion known
as Regiao dos Lagos. This region isa peculiar enclave,
climaticallyandfloristicallydistinct from most of southeastern
Brazil, having aCaatinga-likedryclimateandxeromorphicveg-etation
(Ururahy et al. 1987, Coe et al 2007), thusbreeding activity there
may be because ofconditionssimilar totheCaatinga. It
ispossiblethatA. manglealsobreedsinother, moretypical(humid),
coastal localitiesoftheBraziliancoast,but it should be taken into
account that the coast ismuchmorepopulousandmoreintenselyvisitedby
birdwatchers and ornithologists than theCaatinga. IfA.
manglebredregularlythere, it
islikelythattherewouldberecordsofthisactivity.Intheirabsence,itseemsthattheCaatingaisthemainreproductivegroundsofA.
mangle.AlthoughA. mangle cannot be consideredtoundertake regular
to-and-fro migrations, aspecimen(MNRJ 46057) fromItatiaia
NationalPark (22u 269 S, 44u 369 W) provides evidencethat,
likeintypicallymigratoryspecies, individ-uals of A. mangle prepare
inadvance for theirmovements. Thisspecimenwassalvagedafter
itflewintoawallduringnight time(Sick1997andlabel data). Many
normally diurnal species,includingrails, migrateduringthenight
andareattractedtoandcollideagainst lightedbuildings(Dingle1996,
Sick1997, Taylor 1998, Berthold2002), so it seems probable that
this bird wasperforming migratory flight. Furthermore, it
islabeledtohavelotsof fat. Extensivedeposi-tion of fat is one of
the main features of migratorydisposition,
thegeneticallydeterminedconditionthat results inthebehavior
andphysiologyof abirdthat ispreparingtomigrate(Berthold2002,Newton
2008). Thus, even though the movementsof A. mangle
apparentlyrepresent not a typicalMarcondesetal.
NMOVEMENTSOFALITTLE-KNOWNNEOTROPICALRAIL
669caseofto-and-fromigrationbut ratheracaseof seasonal variation of
range, it seems thatindividuals of this species may behave
likeindividualsoftypically(to-and-fro)migratoryspeciesintheirflightactivityandpreparationformigration.Becauseour
records aredispersedover manyyears, from1903 to2013, and most
individualyears haveonlyoneor afewrecords
(theyearwiththemostrecords, 2009, has15records), wecannot be
completely certain that the
rangeexpansionandassociatedmovementsoccureveryyear. It could be the
case that these inlandmovements occurin some yearsbut not in
others,inresponsetovariationinenvironmental condi-tions. Wecannot
discardthat possibility, but theabove mentioned specimen with high
body fatseems to indicate that birds prepare in advance fortheir
movements, and thus that there is somepredictability in them.
Furthermore, if, as itseems, A. mangle breeds exclusively or
mainlyin Caatinga or Caatinga-like environments, themovements must
be annual in order for it toreproduce.If A. mangle indeed
periodically expands itsrangeandnicheintotheCaatinga,
thequestionarises: whydosomeindividualsmoveinlandbythe end of the
dry/non-breeding season whileothers remain on the coast? A
possibility is that bythe endof the reproductive season,
newly-bornjuvenile birds move from the Caatinga to the
coastandstaytherebeyondtheir first year, until theyare sexually
mature to breed. This behavior,known as retarded or deferred return
(Dingle1996, Berthold 2002, Newton 2008), is known
forseveralraptors, seabirds, andshorebirds(Newton2008) but
undescribed for rails. It would beconsistent with the lack of
observations
ofbreedingactivityinthecoast.Anotherpossibilitywouldbepartial
migration, acommonphenom-enon in which some adult individuals of
onepopulationmigrate,whileothersdonot(Berthold2002, Newton2008).
Insuchsituation,
however,theindividualsthatdonotmigratebreedanyway,in the resting
grounds, which does not seem
tobethecasewithA.mangle.Definitelyansweringthesequestionsisnot
possiblewiththeavailabledataandwill requiredetailedstudyof
thelife-cycleofindividualbirds.Our studyof A.
mangleshowsanunexpectedpatternof periodical
expansionandretractionofrange to include two very different
habitats,combinedwithsomeaspectsof typical to-and-fro migration.
This species consequently doesnot easily fit into any common model
of migratorybehavior,thestudiesofwhichhavebeenfocusedonlatitudinal,
long-distancemigrationofspeciesfromtemperate habitats in the
Northern Hemi-sphere(Chesser1994, Sekercioglu2010,
Daven-portetal.2012).ThiswilllikelyalsobethecaseofmanymorespeciesofNeotropical
birds, oncetheyhavetheirmigrationsstudied.We were able, through a
combination of areviewof occurrence localities and ENMs,
todrawinferences about seasonal variation inthedistribution and
ecological niche of A.
mangle.Eventhoughmodelsfromdifferentseasonswereconsidered
statistically equivalent, ENMtoolswerehelpful
inexplainingthesignificanceofA.mangle movements, inasmuch as our
results argueinfavor of thehypothesis that this species is
aniche-tracker anddemonstrate that it undergoesperiodical
expansionandretractionof its niche.Given the absence of systematic,
long-termlocality-basedsurveys of variationinbirdpres-ence, or of
single-individual trackingdata(e.g.,throughbandingor GPS),
wefollowGibbonsetal.(2011)inconsideringourapproachusefulasacomplementary
tool for investigating the little-studied patterns of bird
movements in theNeotropics. We highlight that such approachrelies
heavily on records of occurrence, andtherefore encourage
birdwatchers andornitholo-gistsaliketoalwaysdocument
andpublishtheirfieldrecords, especiallyof
little-knownspecies,andmakethempubliclyavailablethroughdepo-sition
of digital vouchers in an appropriatedatabase, or whenever
possible,
asspecimensinascientificcollection.ACKNOWLEDGMENTSWearedeeplyindebtedtoall
thecollectors, photogra-phers, recordists andobservers
(ornithologists as well asbirdwatchers)
whosefieldworkmadethisstudypossible.We are very thankful to Gustavo
A. Bravo for hisencouragement, as well as for discussions
andcommentson earlier versions of the manuscript. Natalia
MundimTorres and twoanonymous referees also commentedonearlier
versions. We also thank the curators and staff of thephysical and
digital collections consulted or visited
forwelcomingusandsharingthematerialordataundertheircare; Vagner
Cavarzere and Thiago V. V. Costa
forsearchingorcheckingspecimensincollectionsnotvisitedby the
authors; Erica C. P. de Assis for helping withreferencesabout
theCaatinga; andBrunoLima, AmandaNavegantes, Rafael
BessaandEricGallardoforprovidingobservations or additional
information on records of A.mangle. LFS received a grant, and RSM
was supported by a670 THEWILSONJOURNALOFORNITHOLOGY
NVol.126,No.4,December2014scholarship(130272/2011-6)
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