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 sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academicinstitutions, research libraries, and research funders in the common goal of maximizing access to critical 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: http://www.bioone.org/doi/full/10.1676/13-165.1BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological,ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170journals and books published by nonprofit societies, associations, museums, institutions, and presses.Your use of this PDF, the BioOne Web site, and all posted and associated content indicates youracceptance of BioOnes Terms of Use, available at www.bioone.org/page/terms_of_use.Usage of BioOne content is strictly limited to personal, educational, and non-commercial use.Commercial inquiries or rights and permissions requests should be directed to the individual publisher ascopyright 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:rmarco3@lsu.eduTheWilsonJournalofOrnithology126(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. 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