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Numbers of theBULLETINOFTIlEFLORIDASTATE arc published at irregular intervals. Volumes contain about300pages and arc not necessarily completed in anyonccalendar year.OLlVER L. AUSTL'I,JR., Editor \V ALTEn AUFFEKnRnG,M anaglng Editor Consultants for this issue:John W. Aldrich Joseph C. Moore Communications concermng purchaseorexchange ofthepublication and allmanuscripts should be addressed to the Editor of the Bulletin, Florida State Museum, Seagle Building, Gainesville, Flerida. Published 8 May 1964 Price for this issue $1.35


Contributions to theBULLETINOFTIlE lo"LOlUDA STATE ?\lUSE"L'"J.f maybein any field of biology. Manuscripts dealing with nalural history or systematic problems involving the southeastern United States or the Caribbean area are solicitedespecially.Manuscripts shouldbeofmediulll length---.50 to200pages. ExaminationforsuitabilityismadebyanEditorial Board.The BULLETIK isdistributed \vorldwide through institutional subscriptionsandexchanges only. Itisconsidered the respon.sihility ofthe author to distrihutehispaper to all interested individuals. To aidinthis, fiftyare furnished Uleauthor without cost.PREPARATION 01'MAl\:USCRIPT Highly recormncndcd as a guide isthevolume:ConferenceofBiological Editors, Committeeon Form andStyle.1960. Style manual for biological journals.Amer. Inst. BioI. Sci., \Vashington.92p.Manuscripts shouldbetypewritten with double spacing throughout, with amplemargins, and on only one side of the paper. The author should keep acopy;theoriginalanda carbonmustbesubmitted. Tables and legends of should be typed on sheets separate fromthe text. Several legends or tablesmaybeplaced on a single sheet.Illustrations,includingmapsandphotographs,shouldbereferredtoas"figures."Allill\Lstrations are reduced to a maximum of -i-In by7-1/8inches. Size scales,wherever they arc necessary, shouldbeincorporatedintothe figure. References to literature should conform with the style used in recentnumbers oftheBULLETIN.Spelloutin fnll the lilies of non-English serials andplaces of publication.Footnote material shouldbekept to a111lDUHum.However, provide copy for a footnote detailingthetitle, affiliations, andaddressof the author (see recent numbers of the Bffi,LETIN). Manuscripts mustheaccompaniedby a brief and faclual summary (not a mere description)ofthecontents and conclusions, which points outthepresenceofanyneWinfonnation and indicates its relevance. Initlist allneworganisms described and give their ranges; indicate allta.:xonomicchangespro posed. The synopsis, written in full sentences, shouldbeconcise, but completely intelligible in itself without references to the paper, thereby enabling the busyreader to decide morc surely thanhecan from thetitlealone whetherthepapermerits his reading.Thesynopsis willbepublished with the paper.Itdoes notreplace the usual conclusionsorsummary sections. It may also serve as copyforthe abstracting services.Manuscripts and all editorial matters shouldbeaddressed to: i\'lanaging Editor of theBULLETINFlorida State l\iluseum Seagle Building Gainesville, Florida




THETERNSOFTHEDRYTORTUGAS\VILLIAMB.ROBERTSON,JR.' SYIXOPSIS:New information from unpublished sources and from published recordshitherto overlooked pennit,1re-l:valnation of the history of theDryTortllgas and of the terns thatinhabit them.Thegeography and ecology of the 11 keysthat have variously comprised the group sinceitwaSfirst mappedinthe1770's are described and their major changes traced. The recorded occurrences ofthe of terns reported nesting on the keys are analyzed in detail.TheSooty 'fern colony has Huctuated frolll a low ofabont .5,000 ad nits in 1903toareported peakof190,000in 1950; for the past four yearsithas remained steadyatabout 100,000.TbeBrown ,",oddy population, which reached a peak of 35.000in 1919, was redncedby rats toabout400 adults in 1938; itisintheneigbbor hood of 2,000 today. A colony of 1.50 to 450 Roseate Terns has nestedinmost years from 1917tothepresent. About 500 Lea,t Terns nested regularly from 1916 to 1932,thenunaecountably dwindled to a few pairsby1937andshortly aftcrward disappeared. RoyalandSandwich Terns nested abundantlyinthemid-l9tb century,anda colony of Royals may bave existed as late as 1890.Both species arebelievedtohave helm extirpa.kd from the Tortugasbyegging. No verifiable evidence exists for the nestingofthe Common Tern,whichhasbeen reported several times.TbeBlack Noddy, first reported for the continentalUnited States atDryTortugas in 1960, has heen found there each summer since . The author is Park BiologistatEverglades National ParkandFortJefferson National Monument, Homestead, Florida. Manuscriptsubmitted10October1963.-Eu.Robertson, William B., Jr., 1964.Theterns oftbeDryTortugas, Bull. Florida State Mus., vol, 8,nO.1, pp. 1-95.


I (, ( TABLE OFCONTEKTS Introduction Acknowledglllcnts Location and Physiography History and Names ofthe Tortugan KeysSooty Tern .._._0'._RecordofNesting Discussion Brown N otldy Record ofNesting Discussion Roseate Tern Discussion Least Tern . CommonTern Royal and Sandwif.'h Terns _BlackNoddyLiterature Cited 2 3 4617173852 52 58 64 70737,;_ 7879 82 Theterncolonies ofthcDryTortugas, in particularthegreatbreed ing aggregationsoftheSootyTcrn, Stema fuscataLinnacus, and the Brown Noddy,Anous stolidus(Linnaeus), havebeenof interesttoornithologists since Audubon visited them in 1832. Although the areaisremoteanddifficult of access even today, fewbirdcolonies in North America can boast so long a record of observationsorsoextensive a Iitcrahue. Duringtheearly years oftheCarnegie Institution of \Vashington's Tortugas Laboratory, JohnB,\Vatsonandhis co-workersmadeextendedobscrvationsonSooty TcrnsandBrown Noddies (\Vatson, 1907, 1908, 1910;WatsonandLashley, 1915; Lashley, 1915),Theirwork provided nearly allofthedetailed life historydataavailable for these species until recently.Italso included pioneer experimental shldies of behavior, homing,andorientation, as well asanearly in stance oftheuse of metal legbandstomarkbirds. Exceptingtheworkof\Vatson aud his associates,the Iiterahue consists almost entirelyofdescriptions oftheterneryas observed during brief vi.its.J'vIany accounts since 1900 include estimatesofthcnumberof Sooty Terns, Brown Noddies,andotherbreeding spe-


1964ROBERTSON: DRY TORTUGAS TERNS3cies.Lengthier studiesbyNational Park Service personnelin1937 and1938(Beard, 1939)wereconcerned particularly with predation upon Sooty Tern chicksbyMagnificent Frigate-birds,Fregata mag nificensMathcws. Parts ofthehistorical recordweresummarizedbyBartsch(1919),Vinten (1943), Sprunt(1948b),and Moore andDilley(1953).Modern handingattheDryTortugasbeganwiththe activitiesofJack C. Russell in 1936andwas continued annuallythrough1941, principally on outings sponsored bytheFlorida Audubon Society. About 13,300 Sooty Ternsand246 Brown Noddieswerebanded. The bandingswerereported scparatelybyeightormore individuals and no analysis ofthedatawasundertakenuntil rcccntly (Austin,1962MS.).In June 1959,theNational Park Service,FloridaState Museum,andFloridaAudubonSocictybegana cooperative mass-banding study ofthemovementsanddemographyoftheSootyTernpopulation.Attheend of the 1963 seasonnewbandingsofSooty Ternsbyproject cooperators totalled approximately 32,300 adultsand41,900 juveniles.Inthecourseofthis workitbecameevidentthatanumberofthc widely scattered publishcd rcportsandmuch unpublishedinformationhadnotbeentaken into accountbyprevious compilers. Becauseofthis, severalapparentmisinterpretations ofthchistoryofthe colonyhadgained wide currency.Thepresent summary resulted. The names of birds are those oftheCheck-ListofNorth American Birds,American Ornithologists' Union, 1957, except forthechanges rcsulting fromtherecentdiscovery of Anaus tenuirostris(Temminck) at the Dry Tortugas (Robertsonet al., 1961). ACKKOWLEDG:\fEKTs Itispossible to mentionhereonly a fewofthepeoplewhohelpedmeto assemblethedatathispapersummarizes.Thecooperationofthose namedandmany others contributed greatly to this review. I am particularly indebtedtoJoseph C. I\Ioore for permission to refer to the unpublished reports of tern ccnsuseshemadein1953,1954,and1955. C. Russell Mason alsomadeextensive field notcs available to me. Others who contributed unpublisheddataorphoto graphs included Robert P. Allen, H. G. Deignan, JohnR.De'Weesc, Willard E. Dilley, TheodoreR.Greer, David O. Hill, JamesB.Meade, DennisR.Paulson, Roger T. Peterson, S.Robbins, Alexander Sprunt IV, and LouisA.Stimson.


4 HULLETI/\ FLOHIDA STATE MUSEUMVol.8Albert MaIlUcy provided a wcalthof information from his re search on the history ofDryTortugasanduseful adviceonhistorical sources.Ofthose who helpedmeto obtain copies ofrare publica tions, maps, and material from archives,Imust thank in particular LuisR.Arana, CharlesM.Brookfield, C. Gordon Fredine, Lowell Sumner, andC.R.Vinten. CharlesI.Park, Julius F. Stone, Jr., andC.C. Von Paulsen gavc me their recollections ofDryTortugas in the late 1920's and early 1930's, a period for which little written record exists. Recent workatDryTortugas hasdependedgreatly uponthecooperation of National Park Service personnel in theareaparticu larly District Managerand lVlrs. WallaceB.Elms, District Manager JamesA.Olson, and Park Rangers Roy Evenson, CarlS.Christensen, and JamesE.Markette. Finally,IamgratefultoOliverL.Austin, Jr. for assistance in locating references, friendly encouragement,andmany helpful com ments onthemanuscript. LOC-ATION PHYSIOGRAPHYThe Dry Tortugas, the westernmost outliers oftheFlorida Keys,areanareaof shoals with several small, low islands located about 70 miles westbyslightly north of KeyWest(figure1).Theshoals have the shape of a roughly elliptical atoll with its long axis north east-southwest.Theyenclose a lagoonabout10miles in greatest diameter, its center lyingatapproximately24'N, 82052'\V.The10-fathom line closely approachcstheouter' perimeter of the shoals. Depths within the lagoon are mostly 5 to10fathoms. According to Vaughan (1914) the shapeandalignment of the shoalsweredeter mined primarilybycurrents and antedate the present luxuriant growth of reef corals.ThenearestlandistheMarquesas Keys, about 50 miles east.Theislands of theDryTortugas (Vaughan, 1914; Davis, 1942) are madeupof coarse, unconsolidated calcareous sand and larger detrital fragments, chiefly the remains of lime-secreting marine or ganisms. Skeletons of corals predominate. Because ofthestrong currentsandheavy wave action during storms, little fine sediment accumulatesandtheshorelines oftheislands change frequently. Highest elevationsonmost ofthepresent Tortugan islets donotexceed 3 or 4 feet above normalhightides. Except for Garden Key and Loggerhead Key, all are subject to some overflow bystorm tides.


lNeU_1t. DRYTORTUGAS Under S F.tb.," iI-Exilll'l Isl d. For.er hllQ4 S .. lo (I(,utlc.1Mile,) 1/10 C'..,.......-MiddleIi, ,pG.,... -HospitalKe,'f.,.1') .,' Ke," 10'" I o Bird le, LonlKe, C'.,. G?,.-G-.,.*-,.-.>.......".....'...,"LOllotlletdlfJ.---'L... ti.. H.p S.ulhwo.t Dr,Torturu" Gulf .r Mule..l-'IGUHJo: 1. DryTorlugas,\"(.1)' 585"Dry proaches." showing shoul areas and the location of existingandformt'fislands. Based on Coastand Geodetic Sux LO<.'ation offunncrislands from U.S.Coast :lnd Sun'l'Y4713 "TortugasHarborandAp-


6BULLETINFLORIDASTATE HISTORYANDNAMES OF THETORTUGANKEYSVol.8-AccordingtoHerrera's chronicle of the first Florida voyage (Davis, 1935: 21), Juan PoncedeLeonreached the Tortugas 21 June 1513.Theislandshadbeen sighted fromtheeast astheexpedition was rounding the tip oftheFlorida Keys some weeks earlier.Herreraspeaks ofanarchipelago of "eleven rocky islets" named "Las Tortu gas" because many sea turtles were captured there.TheTortugas offered a protected anchorage where sea birds, hlrtles,andseals (presumably tlJe WestIndian Seal,Manachus trapicalis,now ex ceedingly rare if not extinct) couldbetaken to augment a ship's food supply.Itis likelythatthe islands were visited frequently duringthe250 years following their discovery,butlittle record of this period survives.TAnT.E1. .1\.nms OF THETOHTUGA::-.rKEYSo CauldChart1773-75 Booby Kay East KayBush KayKayLogger HeadTurtle Kay Rocky Kay Bird Kay North Kay Sandy KaySouth \VestKay Tatnall Cednery Chart1829 Bird Key Long KeyEast Key Garden Key Sand Key Loggerhead Key Blish Key Middle Key "Iorth Key :-lorth EastKeySouth 'Vest KeyCoast Survey 185,3-54 Bird Key LongKeyEastKey Garden KeySand Key Loggerhead Key Bll"hKey Middle Key North Key Korth EastKey South West KeyChart 471a 1868-75, 1896 Bird KeyLongKeyEastKey Garden Key Sand Key Loggerhead Key Blish Key :\Iiddle Key Chart58.51958 Bush Key East KeyGardenKey Hospital KeyLoggerhead Key Long Key Middle Key l) Blanks indicate that no island existed at the timeofthe survey.Thefirst modern chart, and the earliest I have seenthatgives namestothe individual keys, was based on a survey madeby George Gauld for the British Admiralty in 1773-75 (Gauld, 1790). Gauld's chart appliesthename"DryTortugas"tothe group as a whole and shows10keys; the namesitgives for 6 of these differ from tlJose used later (table1).TheDryTorhlgas were next chartedbyLieu-


1964 nOBERTSOJ\: DRY TORTUGASTERNS7tenants JosiahTatnallandG.R.GedneryfortheUnited States Navy Department in September 1829. A tracingofthischartisinthefilesof CastillodeSan Marcos National 1\lonument, St. Augustine, Florida (C.R.Vinten, in litt.).The1829charthas particular value because it gives areasandelevations for 6 ofthe11 keysthenemerged. Parties fromtheUnited States Coast Survey workedatthe Tortugas in 1853-54 CTortugas Island", Scale 1:31,680;and"Sec tionNo.VI", Scale 1:400,000, in Bache, 1858),andin 1868-75 CT 1410",Scale 1:10,000,inCoast Survey, 1878;Chart471a, "Tortugas HarborandApproaches", Scale 1:40,000, United States CoastandGeodetic Survey, 1896).Thechartofthearea presently in useisCoast and Geodetic Survey 585,"DryTortugas", Scale 1:30,000, first issued in 1922andlast revised in 1958.Table1 showsthekeysofthe Dry Tortugasthatexistedatthetimeofeachoftheabove surveysand the names applied to them onthevarious charts. Two general typesofkeys maybedistinguished intheDryTor tugas, those little morethanbarrensandbanks slightly c1evated above normal tides,andthelarger, higher,andusually morepermanentislands with considerableplantcover.Thefirst group includes Hos pital, Long, Yliddle, North, Northeast,andSouthwest Keys;thesecond, Bird, Bush, East, Garden, and Loggerhead Keys.BirdKeywastheprincipal nestinggroundofSooty TernsattheDryTortugas fromatleast 1832 (Audubon, 1835) andofBrown Noddies from at least 1857 (Wurdemann, 1861) untiltheisland washedawayinthe early 1930's. During periods of military activityatFortJefferson,Bird Key also servedattimes as a hospital site,quarantinesta tion,andcemetery.Thefonnerhospital buildings later housedtheAudubon amI Biological Survey wardens guardingthetern colony. The 1829 survey recordedtheareaof Bird Key as"4acres 2 roods20poles", slightly morethan 41h acres,andtheelevation as "3 feet8 inches" (Vinten,inlitt.).Latercomments on its area, dimensions, and elevation vary widely.Thearea in 1890 wasstatedas"abouteight acres" (Scott, 1904: 278), in 1910-13 as"about6,000 square yards" (WatsonandLashley, 1915: 35)andas "somewhat lessthan5 acres" (Lashley, 1915: 61), in 1915 as "8 acres" (Pearson, 1915: 412), in 1918 as"about6 acres" (AsheandLowe, 1918 andinabout1926as"lessthanfive acres" (England, 1928:14).Dimensions giveninvarious publicationsrangefrom 500 x 250 feetin1904 (Millspaugh,1907:233)to400 x 300 yardsin1907 (Watson, 1908: 191),andthekey is credited with various elevationsupto "6 feetahovemeantidelevel" (WatsonandLashley, 1915). A comparison oftherepresenta-


8BULLETINFLORIDA STATE MUSECM Vol.8lionsofBird Keyonthecharts of different periods suggeststhatmuchofthereportedvariation existed mainly intheeyeoftheobserver.ItiscommonlystatedthatBird Key was destroyedbya hurricane in 1935,theLaborDayhurricanethatdevastatedtheFlorida Keys oftenbeingspecified (Stevenson, 1938; Davis, 1942; Vinten, 1943; Sprunt,1946b, 1948b).Otherauthors citethe"hmricancof 1933" (Robinson, 1940: 3; Peterson, 1950: 318)and"the big hurricane of 1938" (PetersonandFisher, 1955: 142) asthestorm responsible. Many accounts suggestthatthe key was destroyed suddenly. Dilley (1950:67)wrote: "At times changes maybevery sudden, as illustratedbythecomplete disappearance of Bird Keyduringthehurricaneof1935." Stevenson (1938)notedthatBird Keyhadbeeneroding grad uall y for some time beforethe1935 storm,andRobinson (1940), Peterson (1950),andPetersonandFisher (1955)statethatitbeganto "sink" in 1928.ThedisappearanceofBird Key appears tohavebeenanextended process following destructionoftbevegetation, and without immediaterelationtoany ofthestorms mentioned. In 1832thekeyhada thick coverofbushes (Audubon, 1835),andin 1857Wurdemann(1861: 426) describeditas"covered withbaycedar[Surialla maritima]bushes sevenoreight feetinheight interspersedhereandtherewiththecactus."Laterdescriptionsofthevegetationupto 1910 are almost identical to \Vurdemann's.Asearly as 1904, however, some erosionhadbegun. !vfilIspaugh (1907: 23.'3) notedfrom Lan sing's observations: "VVave action fromthenorthwest appears toberapidly erodingthewestern beach,thevegetation ontheshore plain ly showingtheenchroachmcnt."Thesevere hurricane of 15-17October1910 (Tannehill, 1950: 175-176) wasthefirstimportantevent inthedestruction of Bird Key.Ofits effects Lashley (1915: 62-6.'3) wrote: "The Key was formerly overgrown thickly witbbaycedars,butthegreaternumberof these were killedbythe hurricancof1910andonly a few living ccdars remain."In1915-16theeffcctsofthe 1910 storm wcre still evident. Bird Key thenbadonly scattered patches ofbaycedarbushes (Bow man, 1918:124).On10-11 Scptember 1919,anothcrsevere hurri cane passed directly overDryTortugas (Tannehill, 1950: 186-187).Inhis assessment ofthedamagedone on Bird Key, 'Varden T.J.Ashe (1919MS.)wrote: "All vell:etation on island destroyed." Accounts of visitstoBird Key after 1919 (Bartsch, 192.'3, 1931,19,32;England, 1928)tracetherapid erosionofthcdenudedisland.Thelatcr stages arc indicated inthcfollowing commentsbyCharlesI.Park (in litt.):"WhenIwenttherein 1929, Bird Keyhadalready


1964ROBERTSON:DRYTORTl;GAS TERNSstarted to wash away.Thehouse which thefonnerwardenhadoc cupied was considered unsafe so I lived onGardenKeyandcom muted byboattotheotherkeys. ...Eachyear erosion on BirdKeyprogressed until in 1934 there was very little oftheisland above water level."AsofJune1935, Longstreet(1936a:37) stated:"theremains of Bird Key[are]nowerodedto a negligible sandbar." FIGURE 2. Aerialviewlooking west, nryTortugas, January194,5.LongKeyinforeground,HushKey and CaroenKey next rear, and Loggerhead Key inbackground.ThewhilespotontheshoaltotheleftofGardenKeyandslightly above itisa sand bar at thefannerlocation of Bird Key. (Officialphotograph, C. S. Navy.) The1935LaborDay hurricane was a stormofextreme intensitybutsmall diameterthatstruckthecentral Florida Keys (Tannehill,1950).Tnreply to questionsabont stormandthe one of 4-6 November, which wastheonlyotherhurricane intheareain 193.5, Gordon E. DunnoftheUnitedStates \Veather Bureau. Miami,wroteme(inlilt.):"Neither of th"se storms passed very closetoDryTor tugas or to Bird Key, anci itisdonhtfulthateither of these stonns should have primary responsibility for the disappearancc of Bird Key. I would expect thattheir effect011Bird Key wonld have been rela tively minor." After storms in Jannary1940(Felton,1940 :VIS.),a 4O-footsandhar elevated 2 fcet ahove high wateremergedat the former locationofBird Key.Otherintermittent reappearances have ocenrred1I10rerecently (fignre 2). Bush Key,where most of the Sooty Terns and Brown r\oddies have uested in recent years, has an involved history complicatcdbycon fusion of names.The names Bush Key andLong Key have been


10 BULLETIN FLORIDA STATE :\{CSEU:\I Vol. 8appliedatvarious times to each of the two adjacent islands ontheshoal ofFortJefferson (table 1, figures 2 and3).Thenamesincurrent use were established withthe first edition ofChart585(1922),buttheconfusion persisted somewhat longer (viz., Coast Pilot,1936:78).FICURE:3. At'rial vie.wlooling Garden Key andFort Jefferson in ground. Dushhl:y, the ('ast Sl,it, andthe northernmost sand ridgeofLongKeyat upperright.Largetreesin ofFortJeffersonaremainlybutton wOCKb(Cono(;(lrJws erectus), possiblyremnants ofthe origin..'11 stand. Pilingsatthenorth and south extremities of GilruenKey formerly supported thecoaling docks.The center ofBush Keyisathicket uf hay cedar(Surianamaritima)enclosing severalmangrove-fringedponds. Brown :'Juddies nestattheedgesofthis area. Tht:' SootyTerncolony occupiesopenarcasbetweenthehay cedars and the shore. (Official photo!(raph, U.S.Navy, byU.S. "laval Air Station, Key West, 1959.)Nowthe second largest oftheTortugan islets, Bush Key has under gone several cycles of building and erosion. Gauld'schartshows no landinthearea.The1829 survey reported(as"Long Key")anisland withanareaof"5acres 3 roods 22 poles"andanelevation of "2 feet 4 inches" (Arana, in lilt.).By1832 this island (or possibly Long


1964 ROBERTSOI\: DRY TORTtiCAS TERNS11Key)was thickly covered with bushesandlow trees,andAudubon(1835)referred toitas "Noddy Key" because most ofthepopulationofBrown Noddies was nesting there. Maps of 1853-54 (Bache, 1858) show a sizable islandatthepresentlocation of Bush Key.Duringthe military occupation ofFortJefferson inthe1860's,theisland servedasapastureandslaughter grounds for cattleandhogsbroughtinasfood forthegarrison (Holder, 1868: 262; 1943: 321). Shortly after this time, Bush KeyandLong Key are saidtohavebeen, "almost entirely obliteratedbya hurricane" (Holder, 1892:77).About1889Bush Key was abarrensandbank (Coast Pilot, 1889: 40)andChart 471a of 1896 shows only a small area abovehighwater. Scott's(1890)detailed account of Tortugan geography as ofthespring of1890mentions no landatthis location. The historyofBush Key after 1900 is thoroughly bedevilledbyconfusion of names. Significant observationsonTortugangeographyinthis periodweremadebyLansing in 1904 1907), Bow man in 1915-16 (Bowman, 1918),andBartsch in 1917 (Bartsch, 1919). doesnotmention Rush Key. Bowman (1918: 128-129) describes a large, irregular island thathad shmbs about12 years old. Bartsch (1919: 469, 482) refers to Rush Keyas"an elevated coral reef' withthestatement "allthevegetation, in fact, most everything shiftable abovethesea, ha. long sincebeensweptawaybythewaves."These recordsappearto showthat Rush Key didnotexistin1904,builtuprapidly until 1915-16,andthenwas suddenly reduced and dcvcgetated (presumahly hy storms)toproduce the conditions Bartsch found in 1917. Davis (1942: 187-189)andSprunt(1948b:5-6)adopt approximately this interpretation of its history. DavisalsopointsoutthatLansing may have overlooked a small islandin1904because a considerahle quantity of sand was removed from the areain1901-05 for use as fill during the construction of coaling shedsandpiers on G,nden Key (figure3).The recordisopen tothealternative interpretationthat Bush Keyhada history of steady growth from before1900.Close ex amination oftheaccountsofMillspaugh (1907) and Bartsch (1919) strongly suggestthatthese authors, followingthenomenclature of the chartsthencurrent, referredtothe present Bush Key as Long Keyandvice versa. Bartsch (1919: 469), for example, wrote of LongKey:"the northernendconsistsofabarrenrim of c'Oral boulders that curveseastwardandsouthward, to join withthereef fringe of Bush Key." This is a fairly accurate description of present geogra phy withthenamesofthekeys reversed. Bowman (1918) discussed


12 BCLLETDI FLORIOA STATE MUSEUMVol. 8Bush and Long Keys togetherbutappears tohavefollowed present usage in his applicationofthenames."LongKey" is described in Millspaugh's (1907:225)account as, "so low as tobeawashduringheavy weather" and "voidofvegetation." Bartsch(1919)indicatedthatthesouthernpartof"LongKey" supported a sparse vegetationofgrassandbushes in 1917.Theprobable historyofBush Keymaybesummarized as follows. After having existed as a well-vegetated island for 40or50 years,itwas destroyedbya hurricane around 1870. Sandbars soon reappearedatthe site,butas late as 1904theywere smallandhadnopermanent vegetation.Duringthe nextdecadesome plants be came establishedanda series of ridgesandbars developed as shown in Bowman's (1918) sketch. By191.5(Bowman, 1918)or1919 (Davis, 1942) several ofthesandbarshadgrown together tocutoff ponds fromtheocean. Mostofthearea betweenthecoalesced bars grad ually filledanda long sandspitbuiltupfromtheeastendtogivethekey approximately itspresentshape (Davis, 1942; figure3).Bush Key continued tobuildupduringthe1930'sand1940'sandcontainedanestimated 110,000 square yards in 1946 (Sprunt,1946b:5). .\Iore recently some oftheshores have eroded,buttheisland seems tobemoreorless stabilizedatabout 20 acres. East Keyappears on all maps of theareaand, unique amongthepresent islandsoftheDryTortugas,ithasbornethesamenamethroughout its history. Although more stablethanmanyTortugan islets,EastKey has undergone substantial changes iu size and vegeta tion. Gauld'schartshowsitasthesecond largest island ofthegroup. Thisiscorroboratedbythe 1829 survey which recordedanelevation of morethan4 feetandanareaofabout12 acres, second in size only to Loggerhead Key.Duringthelate1800's and early 1900's, East Keymayhavesuffered several periodsofdevegetationanderosion. About 1860 (Holder, 1892)itwas covered with a dense stand ofbaycedarbushesandnumerous mangroves.In1875itwas reportedtobe"partly coveredwith agrowthof cedar" (Coast Sur vey, 1878) and later (Coast Pilot, 1889) was saidtohave"afew bushes on it."Atalmostthesame time, Scott (1890: 3(2) wrote of East Key:"Itisa low, sandy, coral island, covered in parts with stunted bushes,andcontains an area of perhaps eighteen acres." By 1904 littlebutherbaceous growth persisted and i\lillspaugb (1907: 224-225) describer!EastKey as "little more tllan a mere sand bank 280 x 50 feet in area."Hemay, bowever, have been misinformedaboutits size.In1915-16 (Bowman, 1918: 131-132)theisland wassaid tobc"almost entirely


1964 ROBERTSOl'\; DRY TORTUGAS TERNS13covered with vegetation" including '1arge, well-grown bushes,"butnobay cedar. Its dimensions were given asaboutone-third mile long and less than one-sixth mile wide. Davis (1942: 191) found a thicket of baycedaronthe highest sand ridgeandreported the island's dimensionstobeabout1200 x 600 feet.HestatedthatEastKey"has probably grown in sizeandbecome more stabilized inthepast half-century." Sprunt(1948b;17) wrote ofEastKey: "It com prises about 85,000squareyards," indicating continued growth. At present sizable bushes of bay cedar, sea lavender(Tournefortia gnaphalodes),andScaevola plumieriare well distributed overEastKey.Ithas oftenbeensaidthatSooty Terns and Brown Noddies were not knowntohavenested onEastKeyandseveral authors have re marked uponthefailure oftheternstouseso suitable a nesting area. These comments overlook various records of the 19th century.Largebreeding colonies ofbothspecies occupiedEastKey inthe1850's (Wurdemann, 1861; Bryant,1859a).Sooties,atleast, still nested thereaslateas 1890 (Scott, 1890). Continual persecutionbyeggers, mentionedbyevery early writer,mayfinally have driven the ternsfromEastKey. Though awardenwas in residenceatBird Key each nesting season from 1903 on, his surveillanceisnotlikelytobave extendedtotheoutlying islands.Itisof interestthatno terns have bredonEastKey during thepast28years of strict protection.GardenKeyadjoinsthebestprotected anchorage in Tortugan waters andhaslongbeenthe center ofhumanactivity inthearea(Manucy,1943).Most ofthekey is occupied by the immense ruin ofFortJefferson(figures 2 and 3). A lighthouse was built on Garden Key as earlyas1825.Construction ofthefortbeganinDecember1846 and was discontinuedabout20 years later withthework still far from com plete. After use chiefly as a military prison,thepost was abandonedinthe 1870's.Itwas reoccupiedduringand aftertheSpanish-Ameri can \Var and WorldWarI, first as a coaling station, later as a sea plane base and wireless station. Gauld'schartshowsGardenKey with an irregular shoreline and the 1829 survey reported itsareaasabout 7'h acres.Aninteresting map in the files of the U.S. CorpsofEngineers (Bache, 1845MS.)isa detailed topographic survey of Garden Keyasitwas immediately before the construction ofFortJefferson began.Theshapeisrough ly elliptical, highest land elevationsarejust over 5 feet abovemeanlow water, andthecenter of the islandisshown as low and evidently swampy.Theexact scale ofthemapisuncertain. Calculations (by


14BULLETlNFLORIDASTATE Vol. 8---WilliamM.Alexander, Assistant Park Engineer, Everglades National Park) baseduponthescale takcn from a superimposed outline draw ingoftheFort, laidoutontheoriginalmappresumablybyMajor Bache, give alandarea of 8.8 acrcs above high tide line.Thesize ofGardenKey was increasedbyfillingwhentheFort beingbuiltandagainabout1900whenthecoalingstruchueswerebuilt. Davis (1942: 185) gavethearea as 16 acres, of which 5 acres lay outside the walls oftheFort. No terns arc known tohavenestedonGardenKeyuntilrelatively recent years. Detail shownonthe1845map suggests thatthein terior oftheislandmayoriginallyhavebeentoo heavily vegetated to attract nesting Sooties, althoughmuchofitwasapparentlysuit able for Brown Noddies. Anythatmay have nestedthereundoubt edly were displaced soon after 1845. A few pairs of Brown Noddies have nested on pilingsandintheruinsofthenorth coaling dock in anumberof years sinceatleast 19.32 (Bartsch, 1932).In 19.37 (Long street, 1937), 19.38 (Beard, 19.38), and1947 (Sprunt,1947a)large num bersofSooty Ternsnestedalongtheeast sideofGardenKey. A substantialpartoftheBrown Noddy population also nestedtherein 19.37 and 19.38, butnot in 1947 (Sprunt,1948a). Hospital Key,although always a small, shiftingsandbarwith little vegetation, has existed since the earliest surveys oftheDryTor tugas.Thepresent name, which was used as early as 1875 (Coast Survey, 1878) stems fromtheisolation hospital for yellow fever patients built there inthe1860's. Sand Key,anearlier name, rc mained in commonuscuntilthe1940's. Various plantshavebeenrecorded from Hospital Key,buttheislandisso often awash in rough weatherthatnopermanentplantcover has become estab lished. Least Terns nested on Hospital Key in 1907 (Watson, 1907)and 19.37 (c.R. i\Iason, inlitt.)and a colonyofRoseate Terns has occu pied the key in anumberof recent ycars since 19.37 (Mason, in litt.).Spront(1948b:17) suggestedthatSooty Terns might find Hospital Key a suitable nesting area, a prediction fulfilledwhena few Sooties nestedtherein 1957and1959.LoggerheadKeyisthelargest, highest,andmost heavily vegetated oftheTortugasandthesiteofthe150-foot LoggerheadLight(figure4)builtin 1856-60.Thesizeandshapeofthekeyhavebeenremarkably constant.It had an area ofabout .30 acres in 1829 and approximately the same sizeatpresent, erosionofthewest shore


1964ROREHTSOK: DRY TOHTUGASTERKS1.'5 having been balancedbythegrowth of sandspitsatthenortheast and southwest ends. Loggerhead Key has been creditcd with an elevation of 9 feet above mean tide 1907: 235; Davis,1942:179)butit seems likelythatthis estimate isexcessive.The1829 survey gave the elevationas "4 feet 4 inches." Least Terns nestedonthe Loggerhead Key sandspits intermit tently from before 1900 to 1936 (Russell, 1938 4).No other tcrnisknown to have nested on the island. FIGt:I1E 4. Loggcrhe.ad Key abollt HH5. looking sOllthwest fromthenorth tip.In is the(omlt>rsite of the Tortugas Lahoratory, Carnegie tution of Washinb'lon. Inthe cenler of the island. Loggerhead Light. (Official photograph. lJ. S. :'-Ia"y, hy U.S. \/a"al Air Shtion,K,y West.)Long Keyisabarorshoal of reef debris with several dune-like ele vations of broken coral (figure 2).Davis (1942:189)estimatedthatmore than one-third ofthekey was flooded by normalhightides and that the sparse vegetation of herbaceous halophytes and scattered small mangroves covered less than one-third of the area above high tide.Ifallowanceismade forapparentconfusion of namesinthepast (see Bush Key), it appearsthatLong Key has never been greatly different. Gauld'schartof 1773-75 which shows a small islandatthe north end of thebarandbelow itthenotation "Ridge of rocks almostdryand very steep", closely approximates present conditions.


16 IJULLETIK FLOIUDA STATE Vol. 8A few Least Terns oc-'Casionally nested onthehigher sandbanksatthenorthendof Long Keyaslate as 1948 (Sprunt, 1948e). Roseate Terns have nested there from timetotime, most recentlyin1962 (Robertson, 1962). Some SootiesandBrown Noddies probably nested there in 1932and1933 (Bartsch, 1932,1933).In1943 (Budlong, 1944MS.),1952 (MooreandDilley, 1953), 1956 (Robertson, 1956 ]\[s.), andperhaps in other years, many Sooties have tried to nest in rocky spots betweenthedunesandfarther south on Long Key,butbecause even moderate storm waves wash over this section,theattempts arc be lievedtohave been largely unproductive. Middle Keyisshown on Gauld'schartasa fair-sized island, andthemap symbols indicatethatitsupported some vegetationatthattime.In1875 (Coast Survey, 1878) Key was still considerably larger thanitisnowbutwithout established vegetation. r.lore recentlythekey has existed only intermittentlyasa low strip ofbaresand with few or no plants. Several pairs of Least Terns may have nested on :Yliddle Key in 1947 (Sprunt,1948a),anda small colony of Roseates nestedtherein 1953 (DeWeese, 1953MS.),andpossibly also in 1960. Gauld's name fortheisland, "Bird Kay," suggeststhatitwas once a more important nesting locality. North Key, Northeast Key,and Southwest Key all werebarrensand islandsthathadwashed awayby1875 (Coast Survey, 1878). They have shownnotendency to reappear,buttheformer location of Southwest Keyismarked on present charts asbareatlow water. No plants are recorded from any of these keysandno terns are knowntohave nestedonSouthwest Key, which may neverhavebeenmuch morethana high place inthereef. Northeast Key harbored a large colony of RoyalandSandwich Terns inthelate 1850's (Bryant,1859a).Theonly definite referencetonesting on North Key seemstobe Holder's (1892: 155) mention of "a solitary gull's egg" (fromthecontext possibly a SootyTernegg) found onthebaresummit of a sand ridge.Inaddition, Bartsch (1919: 492-493) believedthattheisland-about8 miles northeast of TortugasLighthouse-"asmall sand-bar a few acresinextent, called BoobyIsland"-whereAudubonfound large numbers of some species of Booby, was probably North Key.


1964 ROBERTSO!\: DRYTORTUGAS TER!\S 17SOOTYTERNAppearanccandbehavior combinetomaketheSootyTerna con spicuous bird,andithas usuallybeenthemostabundantspecies:inthe Tortugas terneries.Littlewonder, then,thatthecrowdedandnoisybreeding colonies of Sootieshaveclaimed most oftheattentionofohservcrswhovisitcdtheDryTortugas. Perhaps inevitahly,muchofthecommcnt ontheSootyTernatthe Dry Tortugas has ccntered onthequestion,howmany? Early ornithologists contented themselves withwordpicturesthatsuggest merely large numbers of birds,hutfew 20th century authors have failed toattempta numerical reckoningofthesizcofthecolony. Their figuresrangein quality from guessesmadeafterbriefobserva tion to estimates calculated from measurementsofcolonyareaanddensityofnests.Table2 showswhatI consider the soundest figurcs available for numbers ofadultSooties ineachycarof record from1903to 1956. Population figures for several of the yearshavehadaneventful history inthehands of compilers,andquantitativedatawere foundforanumberof years previouslythoughttobegaps in the record. \iVith these correctionsandadditionsthcbroadoutline of the historyofthccolony seems clear,thoughmany detailsremainobscure. ThcDryTortugas ternery has been called The OldestBird Col ony" (Peterson, 1950) ontheassumptionthatits known history reachesback tothediscovery ofthearea in 1.'5].'3. Itis reasonabletosup posethatthc"other birds" ofHerrrera'sstatemcnt(Davis, 1935), "...therewerekilledmanypelicansandotherbirdsthatamountedtofive thousand...",included Sooty Terns.Theaccounts ofotherearly visitors,suchas John Hawkins (Longstreet,19360),andmuchlater ones, suchasGeorgeGauld(1796), contain similar imprecise allusionstotheabundanceof sea-fowlattheDryTortugas. No cer tain record ofanyternisknownfor the area, however,priortoAudu bon's visit in May 1832.RECORDOFNESTING 1882. Audubon(1835: 263-269) reported SootiesbreedingingreatnumbersonBird KeyandNoddies breeding on Bush Key. His account showsthatbothcolonieswerethenbeing heavily exploitedasa sourceoffood. Besides several referenc's tothekilling ofadultbirdsandthegathering of eggsitincludesthefollowing: At Bird Keywefound apartyof Spanish Eggers from Havannah. Theyhadalready laid in a cargo ofabouteight tons oftheeggs of


18 Bl;LLET1:-J FLORIDASTATE Vol. 8 TAHLE 2. BnEEDlKc.POPt.,-LATIOKS OF SOOTYTERNS ATTHI-:DRYTORTUGAS Year 190:31907 190919lI I912191.31914HI1.5 19161 ll17Illl8Illill1921l19:3.519:36 19:3719:38 19:39 1940 1941 1942 194.1 1944 1945 1946 1947 1948 1949 19.5019.51 1952195:3 19.5419.5.519,",6Number ofAdults.5000 18,8.58 40,000 48,000 48,000 :30,000 97,.500 102,000 60,000 80,000 100,000lIO,OOO80.000 :30,000 40,000 72.000 64,057 70.000 100.000 100.000 65.000 100,000 130,000 101l.00097.20064,270 104,000 120,220 190,876 167,770 76.326 84,.569 88,7767l,l02 90,452 Method Estimate Area xDensity Estimate Estimate EstimateEstimate Estimate EstimateEstimateEstimateEstimateEstimateEstimateEstimateEstimate Area x Den"ity Area x Density DirectCowlLEstimate EstimateEstimateEstimateEstimateEstimateArea xDensity Area xDensityArea x Density Direct Count Area xDensity Area xDensityArea x Density Area xDensityArea x Density Area xDensityArea x OcnsityArea x DensityArea xDensityDirectCountReferenceBurton (inDntchcr,1904)Watson(1908) Peacon (1909 ms.) Peacon(lllllms.) Pcacon (l\J12 ms,)Peacon(191:3 ms.) Peacon (1914 ms.) Asl,,' andBethel (191.5 illS,) Rethel (1916 ms,) Aslw (in Pearson, 1917) Ashe (in Pearson, 1918) Ashe (1919illS.)Park(m5,notes) (1936) DOl' andRussell (19.36) Longstreet(19.37) Beard (19:38)HuhinsOIl(11 ..):39) Hobinson (1\)40) Peterson (19,;0) Budlong (in Vinten, 194.3) Budlong (1943 ms,)Vinten (in lift.)Sprunt(1946a)Spnmt(1946b)Spnmt (Hl47a) Spnmt(l948e)Dilley (H),;O) MooreandDilley (19,;'3) MooreandDilley (1953) andDilley (1953) (19,')4 illS.) (1954 ITlS,) '>[oore (19,;.5 ms.) Robertson (19,;6 ms,)


19M ROHEHTSO:'J:DRYTORTUGAS TER:'JS 19this TernandtheNoddy.Onasking themhowmanytheysupposed they had,theyansweredthattheynevercountcdthem,evenwhile selling them,butdisposedofthematseventy-five centspergallon; andthatoneturntomarketsometimesproducedupwards of two hundred dollars, whileittook only a week to sail backwardsandforwards and collect their cargo. Some eggers,whonowandthencome from Key \V cst, selltheireggsattwelveandahalfcentsthedozen;butwhereverthese eggsarecarried, they must soonbedis posed ofandcaten, fortheybecomeputridin a few weeks." Sprunt(1948b:8)pointsoutthatAudubon's account contains nothing definiteaboutthenumberofterns.Despitethis,laterwriters almostwithoutcxeeptionhavesupposedthatAudubonfound Sooty Terns in far greater numbersthanwereever seen againattheDryTortugas.Thestatement, "both specieswereon their respective breeding-groundsbymillions," has beencited both as evidence of formerabundanceandas typical of Audubon'sbentfor extravagant language. Althoughattributedto Audubon, in factitis only re portedbyhimastheremarkofanofficer oftheMarion,madeastheship approachedDryTortugas and before Audubonhadseen the tern colony. Peterson (1950: 318) used one of the statistics oftheCubaneggtrade cited abovetoobtain an estimate oftheHumber of terns inthecolony in 1832.Hewrote:"Asooty's egg weighsaboutthirtygrams, oraboutfifteen eggs to the pound.Eighttonswouldcometoabout240,000 eggs.Assooties and noddiesnonnallylaybutoneeggthis shows irrefutablythattheconcentration wasfarlargerthanitis now." A repetition of the exercise (PetersonandFisher, 1955:142)arrivedatan estimate ofabout250,000.HadAudubon mentionednootherstatistics, this ingenious reason ing mightindeedbedifficult to dispute.Theeggerswhospoke of an eight-ton cargo, however, also toldAudubonthattheysometimes realized "upwards of twohundreddollars"pertrip to market.Ifthisisinterpretedtohavebeen as much as $250,the250,OOO-cggcargo was soldattenfor a penny. This seems too good a bargain in eggs even for 1832, especially astbeprice in KeyWestis given as "twelveandahalfcentsthedozen."TheSootyTernpopulation can alsobeestimated onthebasis of areturnof $2-50 persuccessful tripandthe statedHavanapriccof "seventy-five centspergallon,"ifthelatteristakentomean fluid egg contents.Worth(1940: 56) calculated the volume of a SootyTernegg as 1.95 cubic inchesorabout118 eggs to thestandardgallon.


20BCLLETINFLORIIJA STATE MUSECM Vol.8At 75 centspergallon a cargo wouldamountto about 334 gal lons, therefore equalling about 39,412 eggs. Someorall of Audubon's statistical informationaboutegging evi dently is inexact. Attempts to derive a population estimate fromanyofthedetailshegives seem unwarranted.Thequestionofthesize ofthecolony in 1832canbeapproachedbyconsidering thenumberof nests Bird Key could accommodatc. Although Sooty Tcrns nest in dense aggregations, a limit of colony compressibility exists. 'Vatson (1908: 200) wroteofSooties ncsting on Bird Key in 1907: "Eachpair...dcfendeda circular territory roughly 14 inches totwofeet in diameter."Ifthesmaller figure is taken to represent maximum densityofncsting observedby"Vatson, then the minimum areaoftbeterritory of a nesting Sooty was 154 square inches andthemaximum densityofnestingabout8.4 nestspersquare yard. Detailed observations ofthedensity of nestingofSooty Terns on Bush Keyweremadein19.53-56on 20to30plots eachof8squareyards distributedthroughoutthepartsoftheisland judged suitable for nesting.Thelargestnumberof eggs laid on aplotwas 56 (7.00squareyard) on one plot ill 1954. Field maps show four instances in which10eggs occurred in areas of onesquareyardwithinthelarger plots.Ineach case, however, somewerelocatedattheedges,andno one square yard area appears to contain10entire territories.Theaveragenumberof eggs persquarc yard for all occupied plotsandthenumberofplotsthatcontained oneormore eggs were: 19.53-3.00 persquare yard (14 plots), 19.54-3.12 (21), 1955-2.1 (26) (Moore 1954I11S.,1955 ms.);and 19.56--2.53 (20) (Robertson, 1956 ms.). Measured nesting densitiesreportedfor other SootyTerncolonies are mostly similar toorlowerthanthose foundonBush Key.Datafor twobreedingseasons on Ascension Island (Ashmole,1963a: 309),for example, show maximum densities (on plotsof25square yards area) of 5.28and5.00 eggspersquare yard; average densities forallplots occupiedof1.95and2.00 eggspersquare yard.The1953-56datafrom Bush Key suggestthat10 nestspersquareyardisaboutthemaximum density breeding Sooties wiII tolerate.Fewcolonies are this crowded, except locally, because vegetationorterrain limitthenumberof ac-ceptable nestsites. Nesting Sooties ordinarily avoid areas with dense shrubberyorheavy herbaceous ground cover. Ashmole(1963a)foundthatnests alsowerefewer on featurelessbareground deficient inthelocal cluesthatenable


1964ROBERTSON: DHYTORTCGAS TERNS21a bird toreturntotheproperegg.TheTortugas ternery, however, lacks extensivebareareas. From the Tatnall-Gednery survey Bird Keyisknowntohavehadan areaofabout 41h acres (21,780squareyards)in1829. Assuming for themomentthatthe SootyTerncolony occupied its entire sur face, an average density of 11.5 nestspersquareyard wouldbenec essary to accommodate 2.50,000 ne"ts. Parts of Bird Key, however, were thickly coveredwithbaycedarbushes inthe19th century. Photographs takenmuchlater,afterthehurricaneof 1910 had greatlyreduced theamountofplantcover(e.g.,Bartsch, 1919:Plate13), show large areas still not availabletonesting Sootiesbecauseofthedense bush growth. Therefore, I thinkitunlikelythatthe maximum breeding populationofSooty Terns on Bird Keymuchexceeded.50,000pairs. Audubon'smannerof reference to his visit to Bird Key suggests thathesawtremendousnumhersofSooty Terns. Alargesuhjectivc element, however, seeminglymustbeallowed in verbal descriptions of first visits to SootyTerncolonies. \Vhen HerbertK.JobsawtheBird Key terneryatits lowestebbin 19D.'3 hewrote (1905:87)oftheSootics: "Therearesuch clouds of themthataccurately to estimate their numbers was impossible ..."This language also (.'()uld betaken to indicategreatabundancewereitnotfortherest oftheRev erend Job's sentence which reads: ...butmyguess of six oreightthousand I thinkcannotbefaroutoftheway."Itseems characteristic ofmodemsto supposethatAudubousaw allbirdconcentrations in their pristine glory.Howevertruethis may havebeenofmanyplaceshevisited,itdoesnotapplyinthecase oftheDryTortugas. Bird Key was adjacent to a fine anchorage, itself adjacenttoamajmshippinglanethathadbeenusedformorethanthree centuries.Thattherewerenoaccuratecharts before Gauld's surveyofthe1770's can scarcely havedeterredmariners from usingTortugasharbor.Audubonwas toldthattheternshadfrequented Tortugas "sincetheoldest wrecker onthatcoast can recollect,"Itis altogether likelythatthetemerywas firstdisturbedonthe dayofits discovery,andas often thereafter as shipsputin to Tortugas inappropriateseason.Themostthatcanbeassumed isthatexploitationuptoabout1832 had been infrequent enough topermitthe Sootiestorearyoung in most years.1840-1902.Althoughmuchofthis period wasmarkedbyintensivehumanactivityattheDryTortugas, therecordoftheterncolonies inthe19thcenturyafter Audubon's visitislimitedtoobservations


22BULLETINFLORIDA STATE MlJSEl;MVol. 8 by Bryant(18.59a),Wurdemann(1861), Maynard (1881),andScott (1890). Comments by thefirstthreeof these authorsarehrief. Scott discussesthc Dry Tortngas iugreaterdetail,butmostofhis in formationaboutternsishearsay,becausethecolonieswerenotactiveatthe dates of his visit, 19 l\Iarch to 10 April 1890.Dataaccompanyinghirdspecimcns fromthe Dry Tortngasinseveral collections showthatother ornithologists may have visitcdthetern colonicsduringthis period,butleft littleornopublishedrecordoftheir observations.Onesuch visit was by A.L.HeermannandJohnKrider,probablyin ,',,fay 1848. Howell (1932:13)mentions this expeditionbutdoesnotinclude the Dry Tortugas amongtheplaces visited.Heermann(1853: 34), however, lists eggsofthe Sooty TernandBrown Noddy from "Tortugas Islands"presented by himtothe()()Iogical collection oftheAcademyof l\'atural Sciences of Philadel phia. At least apartof this matcrial (ANSP Catalogue Nos. 32055, 32060,and32061)isstillinthe collection (Henry M. Stevenson,illlitt.)Thespecimenshearno date,butHeermannisknown tohavevisitedFlorida only once. Inthebook in whieh John Krider sum marized his eareer, including "only those species of birdsofthe United StatesthatIhavemyself collectedandmounted",herefers tothe Sooty Tern as follows (1879: 81) : "Common onthe Keys ofFloridaandtheTortllgas,whereitbreedsin large numbers. I have two specimensin my collection."Thetwomain items tobegleaned from thelater19th eenhrry papers are: SootiesandNoddies then nestedonEast Key aswenasBird Key, andthecolonieswereunderinereasing pressure from eggers. Dr. HenryBryant traveledandcollected extensivelyinFloridain the deeadc 1850-1860,buthisobituaryintheAnllual Report oftheBoston Soeiety of Natural History for 1867 givesnodetails of his work in Florida,andlittle seems toberecorded elsewhere. Allthathasbecnknownofhis visit to Dry Tortugas isthathewasthereon 8 May. Dataonbirdspecimenshecollected now inthe 'vlusellm ofComparativc Zoology (David O. Hill,inlitt.)suggestthatthe year was 1850.Two Sooty Terns (MCZ Catalogue Nos. 42097and42099) carry the dates 10 May and11May, respectivcly,withnoyear; aGreatWhiteHeron (lVlCZl\'o. 42534)hecollectedatSand Key off Key \Vest, however, places Bryant ncartheTortugasOn16 April 1850. His visittilerecanhaveoccurred nolaterthanthe1853 nest ing season, becauseon19 April 1854hedonatedhis collection of birds' eggs from Florida, including eggs ofthe Sooty and Noddy totheBoston SoeietyofNaturalHistory. Hisownaccountofhis visit


1964 ROBERTSON: DRY TORTUGAS TER'JS 2;}(1859a:19-21) statesmerelythathefoundSootiesandNoddies nest ing principally onEastKeyand"in asgreatnumbcrsasatthetimcof Audubon,"andthatRoyalandSandwichTernswerebreeding "in greatnumbers"onNorthcastKey.InapapcronBahamanbirds(1859b:1:34)heremarksthatSootiesandNoddiesoccurthere "in immense numbcrs, asattheTortugas." Gustavus "Vurdemann (1861: 426)describedhis visittotheTor tugasthelastweekofJune1857 in aletterthataccompanieda shipmentofbirdspecimens totheSmithsonian,puhlishedtwo years after hisdeathin1859. "At theTortugasaretwokeys or islands,EastKeyandBirdKey,whichserve as places of resort tothenoddiesandlaying gulls todeposittheireggsandraisctheiryoung.TheyarewatchedcloselyatEastKeybyboatmen,whogathertheeggstocarrythemtoKcy'Vcstfor salc.ButatBird KcythcbirdsareunderspecialprotectionofCaptainD. P. "Voodbury,theofficer inchargeofconstructionofthcfortifications. ...ThekeysarecoveredwithBaycedarbushesseven oreightfeet inheight,interspersedhercandthcrewiththccactus,amongwhichsomeyounglaying gullssoughtrefuge.Theireggs are laid onthesand,whilstthenoddieslayin nestsbuiltfromtwotosix feet fromthegroundofdriedsticksortwigs.Onlyoneeggwasfoundincachnoddie's nest,andabouttwointhelaying gull's.Theireggsaresaidtohavebeentaken some time previous toour visit, andthattheylay usually two or three. Ipickedupseveralfemalelaying gullswithmyhands,andmighthavecaughtnoddies if ] had notbeenencumberedwiththegun, hirds,andeggs. Noyoungnoddieswereseenatthis time, whichwas t.."e lastweekofJune...."Otherspecimens intheNational \!useum (Deignan, in litt.)anda SootyTerninthe I\luseum ofComparativeZoologyhetookthere10June1858 (Hill, in litt.)show'Vurdemannalso visitedtheTortugasthenextyear.e.J. lVlaynard nevervisitedtheDryTortugasinperson.AsHowell(1932:16)notes "In 1874heworkedatCcdarKcys from.Ianuary26 to l\hrch 1.Fromthere, in a smallyachthewentdownthecoast as far asClearwater,butfromthatpointhcwasobligedtoreturnhomeonaccountof illness, leaving his assistantstocompletethetrip,whichtookthemasfarastheTortugas." Thismaypartlyexplaintheseveral geographicalandhistorical inaccuracics in hisaccount(1881: 480): "The Sooty Ternsarenowonlyfoundinanynumbersonthesmall islands which lietothesouthward[sic] of Key "Vest andwhichareknownastheDryTortugas.Heretheybreedon Bird Keywhichisaboutfourmiles [sic] fromFortJcffer son, depositingtheireggs earlyin "lay. Thebirdsareextremely


24 BULLETII\ FLORIDA STATE Vol.8tamewhennesting, insomuchso,thattheymaybekilledwithsticksorevencaughtwiththehand,andtheydeposittheeggs onthenakedsand.Therewerethousands of these birdsonthis littlekeyin 1874,butasthesoldiersofFortJeffersonhadbeeninthehabitof takingtheeggs regularly everyotherday,butfewor no youngwereraiscd.Theofficer whohadcommandofthefort,prohibitedshootingthebirds ontheisland,butthecontinualrobbingoftheeggsmustulti matelydrivetbcSootyTernsfrom thisbreeding ground." Theactualdateof s assistants' visit isindicatedbya SootyTerninthcMuseum ofComparativeZoology (No. 204310, Hill,inlitt.)takenthere25 J\Iay 1874,thoughno collector is named.Theaccountcontains no first-handcommcntontheNoddy,nordoes it mentionEastKey,whichlayapproximately 4 miles fromFortJefferson.Regardingthecommentsoneggingbysoldiers inthearea,theFortJefferson garrisonhadbeenwithdrawn11January1874, leaving only a smalldetailtoguardordnancestoresandamuchreducedconstructioncrewengagedmainly in closingdowntheoperation (Manuey, 1961 Inaddition,CaptainWoodbury,theonlycommandingofficer known tohaveshown an interest in protectingthcterncolony,hadlefttheDryTortugasin 1860anddiedinKeyWestof yellowfever15 August 1864 (Cullum, 1891: 496-497; j\Janucy, 1961 :'>IS.). Itseems likelythatthereportMaynardreceived from his assistants in 1874blendedconsiderablehearsaywiththeiractualobservations. Most of W.E.D. Scott's infonnatiollaboutnestingternswassentto himafterhisreturnfromtheDryTortugasbyDr. F.S.Goodman,whowas stationedattheQuarantineStation onGardenKey. Scott reports (1890: 307) Sooties nesting onEastKeyandBird Key,andNoddies "mainly confined" to Bird Key,buthis commcnts on egging are ofgreatestinterest: "AlloftheGullsandTernsthatbreedattheDryTortugashavebeenmuchdiminished in numbers inthepasttenyears.Ithas alwaysbeenthecustom forsomeofthcboatsengagedin fishingandspongingaboutKeyWestto rcsort to these islandsduringthebreedingscason,andlatelytheirdepredationshavereallymadeavcryappreciabledifferenceinthebirdsthatresortto thisbreedinggronnd. Iamtoldthattheeggshavea commercialvalucas an articlc of foodinthemarkets of Kcy \Vcst,wherebarrelsofbirds' eggs fromtheTortugasarebroughtevery seasonoflateyears." Vinten (1943: 54) suggeststhatsearchoftherecordsofgovernmentagenciesthathadmaintaincd operationsattbeDryTortugasmightreveal additionaldataabouttheternpopulationduringthis


1964ROBERTSO"l,DRYTORTUGASTERI\S25 period. mostof the searchofthevoluminousArmyarchives of Fort Jefferson remains tobeaccomplished,thestudies of historians show these archives doindeed contain informationpertinenttothehistory ofthe ternery. AlhertManucy (in litt.)advisesmethatamong records he examinedherecalls having seen corrcspondcncc relatingtothe visit of Louis Agassiz totheDry in 1858,andthat the Fort JeffersonLetterBooksincludesuch itcms as alcttcrfrom !,[ordecai and Co. to \Voodhuryon 3 May 1859 ship ment of \Voodbury'sbirdspecimens totheSmithsonian Institution. The historical records l\hnucy (1961MS.)studied BirdKeysuffered evenmoredishlrhancethantheauthors of calworkson the DryTortugashaveappreciated. Shortly afterwarbegan in 1861, for instance, concern for the safety ofFortJefferson, still unfinishedandweaklyarmed, led totheappropriationearlyin1862of 8200,000tofortify Bird Key.Thepreliminary survey, in cluding extensive borings todeterminesubsoil structure, was de layedbypersonnel changesandslow deliveryofmaterials,andwas not completed untilthe of 1864.Theprojcctthensecms to have lapsed,hutitcanhardlyhave failed todisrupttheternsattempting tonestduringthesurvey. '.'!anucyalso cites aletterof III July 1865 tothePostCommandantfromEdwardFrost, AssistantEngineerin Charge, complaining of the removalofanumherofhogs "fromtheirranging onLongKey toBiniKey"whichcontained"thescatteredgraves ofmanyUnion SoldierswhohavediedatthisPost thc war."Whetherornotthehogs werereturnedtoLongKey seems tobeunrecorded. MostprohahlytheSooty Terns failed torearyoungattheDryTortngas in mostoftheyears from 1860 totheearly 1870'swhenFortJefferson was heavily garrisoned. This loss ofannualrecruitmentplus anundoubtedlyheavy mortality of adultsmusthavereducedthepopulation rapidly. Little definite informationaboutthetemeryexists fortheyears 1890-190.3. Itmaybepresumedthatthecolony wasraidedregularly by eggers,andthatsome time in thisperiodSooty Ternsnestedfor the last tillle onEastKey. J.\V.Atkins, a well-knownresidentcol lector of Key \Vest. collected specimensnowintheMuseumof Com parative Zoologyat the DryTortugasinIVlay1896,butnootherrecord of histripisknown.A.G.Mayervisited BirdKeyin 1898,buttheonly dahllllpublished(inDutcher,1906)ishis impressionthatSootieswerethenaboutone-third as numerous asathis next visit in 1906.


26 BULLETI]\" FLORIDA STATE MUSEU'.I Vol. 8WiththeoutbreakoftheSpanish-American War,FortJefferson was garrisoned once more from 1898 untilabout1906.In1900theDryTortugasweretransferredtotheNavyDepartmentandcon struction of a coalingdepotatGardenKey began.Sprunt(1948b:9) suggeststhattherenewedmilitary activityatFortJeffersonprobablyputadditional pressureontheterncolony.TheNavyatthecommandlevel wasawareoftheneedtoprotectBird Key, for a letter fromCaptainT. C.TreadwellquotedbyDutcher(1903: 120-121) statesTreadwellordered egging stoppedsoonafterhe assumedcommandoftheU.S.Naval Station, Key \Vest,inJune1901. Unfortunately orders from Key \Vestwerenotaltogetheref fectiveattheDryTortugas, for according toThompson (19OS: 77-78) the terns "suffered very seriously" from cggers in 1902.Thompsonadds,presumablywithreferencetotherecentpastandtobothSootiesand l\' oddies:''Therehavebeen yearswhennota single individual was raised, every egg bavingbecntaken shortlyafteritwas laid." Thanks to \VilliamDutcher'suntiring effortssternermeasures toprotectBird Kcy followed in 1901. TheSecretaryofthe ?\avy issued anorderon 24 Aprilprohibitingthetaking of cggs ordisturbingof ternsatDryTortugas,andin \lay \V.R.Burton wasdctailedthereas a specialwardenrepresentingtheAmerican Ornithologists' Unionwiththepermissionandlogisticalsupportof theNavy(Dutcher, 1904). Burton arrivedatBird Key accompaniedbyH.K.Job19May1903.Themodcrnhistory oftheternerycan fairlybesaid tobeginonthatdate.1903.FourestimatesoftheSootyTernpopulation in 1903arcavailablefromthepublisbedcommentsofthcoriginal observers.Theyarc: "3600"byJobandBurtonmadeberoreJobreturnedto Key \Vest on 22 :'Yray;"at least50(X)"byBurtonin alettertoDutcherdated15 July 190:3, theincreaseaccountedforbybirdsthatbegannestingafterJob'sdeparture;"five to sixthousand"byJobin alettertoDutcher(allthreefigures publishedinDutcher, 19(4); and, "sixoreightthousand" (Job, 1905: 87).Thecontextoftheaccounts sug geststhatthefiguresrefertonumberofadultSootiesratherthannumberofuests,bntnowhereis this clearly stated. Compilershavegiventhe 190:3 population as 3600 (Longstreet,1936u;Vinten, 1943; Sprunt,1947b;Peterson 1950), 6-8000 (Sprunt, 1948b), ancl"about7000 nests" (FisherandLockley, 19.54: 60;PetersonandFisher, 1955: 142).Thefigures,whereidentified,areinall casescreditedto Job. I considerthewarden'sfigure of 5,000,baseduponobservation ofthe


1964 ROBERTSO:\l: DRY TORTUGA.S TER:\IS 27colony throughtheentirenesting season, tohethesoundestestimatc available. I have found norecordofthecondition ofthecolonyin1904and no estimatesofthepopulationfortheseasons of1904through1906.Charles Russell,thewardenin1905,reported"avery success ful season" (Dutcher,1905).Aftervisitingthecolonyin1906A.G. Mayer informedDutcher(Dutcher,19(6)thattheSooticsappearedto bethreetimes as numerous as theywerein1898.1907.JohnB.Watsonbeganhis studies ofthetern colonyin1907and also served asthewardenoftheNational AssociationofAudubonSocieties forthatseason.Inadditionto hisotherwork \Vatsonmadea careful estimateofthenestingpopulationof Sooties.Hedivided the colony into 10 sectionspresumablydistinguishedbyconspicuous featnresofvegetationorterrain. Bydeterminingtheareaandsam plingthedensityof distribution of nestswithineach section,hear rivedatanestimated9,429nests or18,858breedingadults (Watson,1908: 198).1908. summaries ofthechangesinsize oftheTortngasSootyTernpopnlationinelndean estimate of20,000(or 10,000 nests) asthepopulationin1908.All authors who cite anauthoritycreditthis figure to \Vatson who, according totheCarnegieannualreports, was not attheDryTortugasin1908or1909.Theearliest reference I find to itisLashley's(1915: 61)statementthatSootyTernnests to talled "morethan10,000 in1908,"withnomentionofthesonrce of his information. Ihaveomittedthefigure from table 2becauseI can find noauthorityfor it.1909-1916.On6 April1908ExecutiveOrderNo.779of PresidentTheodoreRoosevelt establishcdtheTortugasKeys Reservation for protection of birds nesting iuthearea.Theorderspecifiedthatuse as abirdreservation wasnotto interfere with military uses (under President Polk's ExecutiveOrderof17September1845establishingtheDryTortugas \lilitary Reservation) exceptthatmilitaryuseof Bird Key was prohibited. ProtectionoftheTortngas Keys Reserva tionbecametheresponsibility oftheBurcauof Biological Survey. After1908wardenprotectionattheDryTortugas wassupportedjointlybytheBiological SnrveyandtheNational AssociationofAudubonSocieties. T.J.Ashe of Key West,whowas in generalchargeofbirdprotection activitiesintheFloridaKeysduringmost oftheensuing decade,hiredandsupervisedthemenstationedatBird Key.


28 BlJLLETI:-i FLORIDA STATE Vol.8ThesewereJohn Peacon (1909-1914),LudwigBethel (1915-1916),andWilliamE.Lowe(1917-1919). Warden's reports onthecondition and size oftheterncolony weremadeannually tobothsupporting organizations.Fromtheannual reportstotheBiological Survey I have seen onlythedataenteredinthebirddistribution filenowatPatuxent vViidlife ResearchCentcroftheBureau of Sport Fisheries and Wildlife.Theannual reports totheNational Association of Audubon So cieties for this period are still inthefiles ofthatorganization. Some werepreparedbyAsheandsubmitted in his name; others seem to have beenpreparedbybiswardensatBird Key.Theyinclude esti mates ofthepopulation of Sooties inaUtheyears 1909through1916 except 1910. Watson apparentlypreparedthe1910reportforBird Lore,butitwas not publishedandhasbeenlost. Thisisunfortunate because comments in a later report (Watson, 1912 1\1S.) indicatehemadethe1910countof Sootiesbythcsamemethodheusedin 1907. Previous summariesofthecolony include no mentionofSooty Tcrm in these years,butskip dircctly fromthequestionable 1908 figure to 1917.Inaddition totheannual warden's reports, several published comments for this periodhavebeengenerally overlooked. Ofthepopulation in 1913 WatsonandLashley (1915: 38) wrote: "There areprobablymorethan18,000 (possibly 30,000) sooties on BirdKey."On28 \lay 1915HerbertK. .robandH.R. \lills visited Bird Keytotake motion pictures for tile National Association of Au dubon Societies. Abriefexcerpt published from Job'sreport(Pear son, 1915) givesthenumberof Sootiesas"possibly 75,000." Pearson alsoprepareda longer alticle aboutthis trip (1915MS.),evidently copy forBird-Loretbatwasn't used, which quotesmoreextensively from Job.Itrevealsthatti,e 75,000 population figure was based on area-density calculationsbyMills. Because these calculations con tain obvious inaccuracies impossible to resolve today, 1haveusedthe1915 population estimate fromthewarden'sreportin table 2. 1917. According toWardenT.J.Ashe'sannualreport(Pearson, 1917: 398).....therewereprobably80,000 of these birds [Sooty Terns] nesting ontheisland." This figure hasbeenoverlookedbycompilers, who insteadhavemisquotedthe1917 population of Sooties from Bartsch as "18,000" (Longstreet,19360;FisherandLockley, 1954)or"25,000" (Vinten, 1943;Spnmt,1947b, 1948b). Baltsch's list ofthebirds seenattheDry Toltugas 19-31 July 1917 (1919:471)in cludesunderSootyTern"adult118,000 young27,200."Thefigures are keyed to footnotesthatread: "1 Based upon Doctor \Vatson's een-


1964 ROBERTSON: DRY TORTUGAS TERNS29susof 1908.", and "2An estimate admitting two-fifths as many off springaswehadparents." Bartsch also (1919:473)wroteoftheSooties: ... probably more than 25,000arepresenton Bird Keyatthe close of the breeding season." Apparently Bartschmadeno independent estimate oftheSooty Tern population in 1917; the figure 18,000isan approximation of Watson's total for 1907 (not 1908)andthe "more than 25,000"ismerely 18,000 adults plus Bartsch's arbitrary figure for young oftheyear.1918-1934.None oftheearlier compilations mentions these years. I have seen warden's reports only for 1918and1919. A Federallaweffective 1 July 1919endedtheNational Association of Audubon Societies' participation in the protection of Bird Key (Pearson, 1919). The Biological Survey continued to employ a wardenattheDryTor tugas during the summeratleast through 1930,butno wardens' re ports canbelocated in the files now storedatPatuxent Wildlife Research Center (Robbins, inlitt.),andthe distribution files contain only the warden's estimate oftheSootyTernpopulation in 1929 (Park,MS.notes). Severalpopulararticles published in the 1920's refer in passingtothe number of ternsattheDryTortugas. England(1928:86) men tions a population of 50,000, and a photograph in an articlebyLong ley(1927:66)iscaptioned: "The west shore of Bird Key showing some of the 33,000 birdsthatbreedhere annually." These figures are not consideredbona fidepopulation estimates. Neithercanbeassociated with a definite year, and the 33,000issuspiciously near Bartsch's(1919:471)total of 32,810 for all the birds(19species) he identifiedattheDryTortugas in July 1917. Bartsch visited Bird Key several times duringthe1920's and in August of 1931, 1932, and 1933. Existing records of his trips containnoreference to the total numbers of Sooty Terns.Thebrief published aecounts of the later visits (Bartseh, 1931, 1932, 1933) have great in terest because Bird Key was then eroding rapidly.In1932 Bartsch notedthata few Sooties were nestingonBush [Long] Key.Thefollowing yearhereported (1933: 267)thatmore than halfthepopu lationhadleft Bird Key and "It isbeginning to look as if the major portion would eventually establish itself on Long [Bush1Key." C.C.Von Paulsen of Homestead, Florida, then an officer oftheU.S.Coast Guard, visitedtheDryTortugas frequently in the years 1932 1934.Ashe remembersit(personal communication) a substantialpartof the Sooties nested on Bird Key in 1933 and smaller numbers remainedtherein 1934.


:10 BULLETINFLOHIIJA STATE MUSEl:MYol. 8 Fromthescant information availableitappearsthatthetern colony may havebeenwithoutwardenprotection intheearly 1930's. Charles I. Park, the last Bird Keywardenandnowa resident of Key "Vest, wrotemein aletter14December1959:"AswellasI can remember, I served aswardenintheTortugasareafrom 1929 throughthesummer of 1934, a total of six years."AsG.A.England(1928:14)refers to CharlesParkas the Bird Keywardenduringhis visit there the summer of 1926or1927,apparentlyMr.Parkbegan hissixyearsofservice twoorthreeyears earlierthanherecollects. Others whoknewtheDryTortugas intheearly 1930's do not recall awardenintheareaduringthose years (Julius F. Stone, Jr., Charles M. Brookfield,andC. C, Von Paulsen, personal communications). Absenceofwardenprotection would explaintheapparently well founded rumorsthattheearly depression years saw a vigorous re newalofeggingattheDryTortugas.Itseems likelythatprotection oftheterneryatleast was less vigilant inthenesting seasonsof1931 through 1934, although Sootiesareknown to have succeeded in rear ing many young in some of these years (Bartsch, 1932).TheNationalParkService assumed administrative responsibility for theDryTortugas early in 19.35. Mason (1936:18)mentionsthattheCustodianofFortJeffersonturnedawaymanyboatparties from Key "Vest thatcametogathereggsonBush Key in the spring of 1935. Correspondence in NationalParkService files suggeststhatthe colony was raidedlateinthe1935 season and anumberof young birds taken. ProtectionofthecolonybytheNational Park Service probably wasnotfully effective untilthenesting seasonof1936.From1935 through 1941 oneortwo groups of observers visited theDryTorhlgaseachJuneon trips sponsoredbytheFloridaAudubon Society.Thevisitswerebrief,eachgroupspending from two to five daysattheTortugas. Adultandyoung ternswerebandedin 1937 through 1941,andthepublished accounts of allthetrips, exceptthatof 1941, include estimates ofthenumberof Sooty Terns.In1937and1938manySooties nested alongtheeast side ofGardenKey (figures ,5 and6) as well as on Bush Key. 1935.Thepopulation figure in table 2 is an averageofestimatesbymembers oftheparty (I\Iason, 1936). Somethoughtasmanyas.50,000Sootieswerepresent. 1936. DoeandRussell (1936: 6-7) stateofthe published population estimate:"Itwasthegeneral opinion of thosewhohadbeenonthe


196-1 nOIlEHTSON: DRY TORT1:CAS TERKS 31trip in 1935thatthetern colonyhadincreased one-third." r.Iason(MS. notes) enteredan estimate of48,000in his field noteswiththecommentthatheconsideredit"very conservative" hecausethenest ill/!; colonyofSooties covered a much largerareaon Hush Kcy than it had in 193.5.FlGu, 5. Portion oftheSootyTerncolony ontheeast side ofGardenKey in 193;: (top) June; and, (bottom) August, showing many well-grown juveniles. (Kalional Park Service photographs hy Philip C. Puderer.)


:32 BULLETII\ FLORIDA STATE MUSEUMVol. 81937.Thccolony was said to occupy anareaof 8000 square yards onGardenKeyand4000squareyards on Bush Kcy.Fromthisarcaanda nesting densityof "about six sootiestothesquareyard,"determined fromonesampleplotof 9squareyards in "a typical sec tion"oftheGardenKey colony, Longstreet (1937:8)calculated a total of "72,000[adultSooties] achmllypresentatone time."Though72,000 birdspresentatone time would represent a total of 144,000 breeding adultsbytheusual methods of reckoning, Longstreet (1937:8)continued: "It would seemnotfar wrong to calculatcthenumberofadultsooty ternsattheTorhlgas inJune1937, as approximating 100,000. Thiswouldbea tremendous increasc over any previous estimates,andforthatreason maybcseriously in error. But,atany rate,itisan estimatebasedon actualcountofbirds in a given area, multipliedbythcnumberoftimcsthatareais found inthetotalarcaoccupiedbythebirds." All summaries ofthehistory oftheTorhlgas SootyTernpopulation have citedthe 19.'37 population as 100,000 from this source. Hussell (1938 also "estimatcdthenumbcrof Sooties to excecdlOG,OOO."Otherobservcrsappeartohave considered this estimate too high. YoungandDickinson (1937) bclievedthatBush Keyhadno morethan20,000 Sooties,andMason(MS.notes) rccordedanestimateof75,000 for tI,e totaladultpopulation. Longstreet (1937:7)includes a photograph,takenfromthcterrepleinofFortJefferson, ofthesampleploton whichthefigure for density of ncsting was based.Thepichlfe shows mostoftheSootiesareeithcrincubatingorbrood ing small young,andhencc distributed oneadultperterritory. Be causeoftheangleitisnot possible to tell exactlyhowmanySootiesareon nests withinthe9squareyardplot,butthenumberis20 to 30, certainlynot54. Thus, Longstreet's figure of 6 birdspersquareyardisapparentlybasedon a nesting densityofabout3 nestspersquareyardwith allowance fortheabsentmcmberofeachpair. Accordingly, 72,000 is consideredthesoundest estimate ofthebreeding population ofadultSooties in 1937. 1938. This year Sooty Terns again nested onbothGarden(figure6)andBush Keys,butthecolony dividedmoreequallybetweenthetwo. Considerable effort was devoted to careful measurementsofareas occupiedandnest densities onbothkeys,andtheresulting es timate (Beard, 1938) isundoubtedlyoneofthemore accuratc ofthepopulation figures foradultSooties intheTorhlgas ternery. Di rectcounts of nestsonthe-coaling docks (figure 6a) and in small, irregular patchesofdense vegetation onGardenKey totalled 3950.


r1964 ROBERTSON: DRY TORTCGAS TERNSFlGUlU:6.SootyTerncolony onGardenKeyin1938:(top) Sootks nesting on the north coaling dock; (bottom)anothersectionofthecolony c",rly intheseason. Bush Key in the background in hoth photo.'. (National Park Service photographs by DanielB.Beard.)


.34 BULLETINFLORIDASTATE Vol.8Themainopennesting areas occupied 5442 square yards on Garden Keyand11,097 square yards on Bush Key; a measured sample of 276 square yards onGardenKey yieldedanaverage nesting density of 1.8 nestspersquareyard, which was takenastypical forbothkeys. Nesting density inthemoreheavily vegetated parts oftheBush Key colony was determined as 1.25 nestspersquare yard. Beard's (1938) calculations contain a slight errOr in addition, andthecorrect totalis64,057,not64,058. 1939.Theentire colony of Sooties nested on Bush Key this season.O.B.Taylor (1939 :\IS.) was toldbytheCustodianthatpartofthebirds first settledonGardenKey in early May,butsoon moved acrosstoBush Key. Robinson (1939: 7)thoughttheyabandonedGardenKey because "mostofthecover aroundtheforthadbeencutdown prior tothearrival oftheterns this season."Thoughhespeaks of counting birds on "sample areas," Robinsonprobablyarrivedathis population figurebycalculating from approximationsofthecolony areaandnesting density.Asitisnotcertainthatanyareasweremeasured, thisandhis 1940 figure are considered simple estimates. Vinten (1943) creditsanotherestimate, also of 70,000andperhaps taken from Robinson, to James B. Felton,thenCustodian ofthefort. Taylor (1939 :\[s.) recordedanindcpendentestimateof65,000adultSooties from his observations latcr inJune1939.1940.A sketchofthe colony (Robinson, 1940) showsthatSooties occupied mostofBush Key excepttheeastern sandspit, astheyhadin1939.Theaccompanying text reads: "At firstitdidnotseemthattherewerequite as many sooty ternsaslast year,buta complete tourofthekey revealedthatthercweremorethanweexpected.Thesamemethodwas employed to estimatethenumberofbirds as last year,andourfigures showthattherewere100,000 sooty terns inthecolony." Justhowthis was calculatedhedoesnotsay. 1941.Thepublishedreportofthetrip (Rea, Kyle,andStimson, 1941) included no estimateofthenumberofSooties,butR.T. Peter son, who aceompaniedthesecondofthetwo parties, wrote (1950: 318): "On ourvisit in 1941wehardlydaredestimatethenumberexactly,butitwas well over the 100,000 mark." 1942-1944. Information for these years comes fromtheofficial re ports of Custodian RobertR.Budlong.Ashewasunableto spend much time observingthecolony, his comments on numbers and popu-


r 1964 1l0BERTSOI\: DRY TORTUGAS TEHI\S lation trends mustbeviewed as impressionsratherthancareful esti mates. Military aircraftwereactive in theDryTortugas area duringthisperiod. Budlong (1942 commentsthattheterncolony was frequentlydisturhedbylow-flying planes in 1942.Thereportof theA.a.V.Committee on Bird Protection for 1943 (Allen, 1944: 629) states: "Vnauthorized useofBirdKey[sic],FortJefferson National Monument, as a bombingtargetbyunidentified aircraftlatein 1942 resulted in a firethatburnedall vegetation. Thisandseveral less injurious actsofsimilarnaturehave beenthesubjectofprotests to the several militaryandnaval establishments. Fortunately,thefire occurred outside tile nesting season,buttheisland will notbeusable bytheSooty andNoddyTernsuntil itisrevegctated."Thecomments presumably apply to BushKey.Burning ofthevegetation isnotlikely to have discouraged SootyTernnestingbutitmay wellhaveaffectedtheNoddies.In1942 Budlong (1942 MS.) stated the colonyhaddecreasedaboutone-thirdandestimatedthenumberof Sootiesat60-70,000, allonBush Key. Vinten's (1943)statementofthefigure as 65,000hasbeenfollowed.In1943 Budlong (1943 1\IS.) consideredthepopulationtohave shown a 50percentincrease to"about100,000."In1944theSooties abandonedLongKey,theeastspitof Bush Key,andseveral large areas on BushKeyproper, all used heavily in 1943,butBud long (1944MS,)believedtherewere"asmanyormore Sootiesinthecolony" asin1943.Attheendofthe season Vinten (in litt. to Re gional Director,V,S.FishaudWildlife Service, Atlanta, Ga.) com mented: "About 130,000 birds nested there duringthepastsummer." 1945-1948.Datafor these yearsarequotedfromthereports of Alex ander Sprunt, Jr., whomadeannual trips to the colony inJuneanddeterminedthesize of theadultSooty population eachyearbyan areaXdensity method. \Vhile hepaidcareful attentiontothespace the colony occupied, justhowhemeasuredtheaverage nesting den sities isn't always clear.In1945heappears to have used those de terminedbyBeard (1938),about1.8 nestspersquareyard in open areasand1.25inmore heavily vegetated sections.Theotheryearshedetermined separate nesting densities foreachsection ofthecol onythatappearedto differ materially,buthegives sizes oftheareas sampledandcountsofnests in each only for Bush Keyin1946(1946b:5).Sprunt also describedtheremarkablespreadof vegetation on Bush Key in thisperiodandits effect onthelocationanddensity ofthenesting Soaties.Herecords the spacethecolony occupied on Bush


36BULLETINFLORIDA STATE MCSEUM Vol. II Keyin1945 as.34,000square yards(1946a),in 1946 as 27,200 square yards(1946b) ,andin1947 as about7,000 square yards(1947a).In1947 some of the colony nested on Garden Key again as theydidin 1937 and 1938.In1948 the entire colony again locatedonBush Key; the vegetation was still luxuriant,butthe Sooties dispersed more thinly over anareaof 52,000 square yards(1948c).1949-1956. Population estimates for these years weremadebyper sonnel of Everglades National Park. Willard E. Dilley, then Chief Park Naturalist, workedattheDryTortugas in 1949 and 1950, andheand JosephC.I\Ioore, then Park Biologist, worked together there in 1951. Nloore continued the anuual surveys through 195.5. Imadethe population estimate of 1956 following procedures established by Moore. Results of the surveys of 1949 through 1952 have beenpublished;datafor 1953 through 19156 are from typed reports in the Everglades National Park files. All population estimates were obtained by the usual areaXden sity methods. Those of 1949 through 19151 were based upon separate determinations of area and density of nesting in anumberof sub areas wherethepattemof occupationbynesting Sooties seemedto differ noticeably, essentiallythesame procedure folllowedbySprunt, Beard, and others back to ''Vatson in 1907.Thenumberof sub-areas distinguishedandmeasured separately was: 1949,7;1950, 22; and, 19151,IS. In1952 Moore established 20 marked plots each of 8 square yards distributed throughout the parts of Bush Key consid ered tobeavailable to nesting Sooties.Dataon density of nesting used in calculatingtheSooty Tern populations of 1952 and 1953 were taken from these plots, anddatafor 1954 through 1956weretaken from these plots plus 10 additional plots Moore establishedin1954.In1951,1952, and 1956 numbers of Sooties nested among rough coral rubbleatlow sites on Long Key. Moore and Dilley (Moore,MS.notes) estimated 455adultSooties nesting on Long Key in 1951, and in 1952 Moore (Moore and Dilley, 1953:76)believedabout2000 present, although few yethadeggs. On 26-27 :\Iay 1956, David O. Karraker, my wife,andI counted 2880 Sooty Tern nests with eggs in place on Long Key, and saw about 700 scattered eggs from neststhathad been Hooded (Robertson, 1956 ]\1S.). Allthe Long Kcy nest ings were behindtheschedule of the main colonyandproducedfew or no young.Inreporting Sooty Tern observations from a visit to theDryTor tugas in I\Iay 1953, Fisher (in Peterson and Fisher, 1955: 143) com mented: "My own estimate of thenumberof occupiednests-


1964ROBERTSON: DRY TORTUCAS TERNS 37BO,OOO-wasnotfar off. A ccnsus based on sampleplotcountswhichwasmadetwo wcekslaterbytheParkServicecameupwitha figure of 84,569 sooty nests."Thefigure mentioned was infactanestimate of thenumberof breeding adults (Moore, 1954 thenumberofnests actuallyamountedtobutfewmorethanhalfFisher's estimate. 1957-1963.In1957 Sooty TernswerefirstrecordednestingonHos pital Key. Mr.andMrs. JohnR.DeWeesefoundabout200 nests thereinJunebutwerenot certainthatany youngwerereared(inlitt.).NonenestedonHospital Key in 1958 (RichardWard,inlitt.),buton 15June1959 O.L.Austin, Jr., C.R. andIfoundabout50adultSooties amongthecolony of RoseateTernsthereandlocated about 10 SootyTernnestswitheggs.InthemainterneryofSootiesonBush Key hatching wasatleast 90percentcompleteatthis tline andthelargeryoungwereabouthalf-grown.Inthenesting seasons of 1960-1962 no SootieswereobservedatHospital Key,butin July 1963about8 adultsappearedtobesettled there, again associated with nesting Roseates. No search wasmadefor nests,butthebehavioroftheSooties suggestedthattheywerenesting.Itisofin terestthatall occurrencesofSooties on Hospital Keywerein years when Roseates also nested there, none havingbeennotedintheyears whentheRoseate Ternslocatedelsewhere.WorkattheTortugasin 1959-1963 consisted chieflyofbanding adultandyoung Sooty Terns in large numbers,andnodirectesti matesofthe size ofthecolonywereattempted. ]\ly impression is thatin 1959-1963 the population was intherangeof 70,000 to 100,000 breedingadultSootiesandvaried relatively little fromyeartoyear. Approximately 6,500 to 11,000 young Sootieswerebandedeachyear in 1960-1963 and therecordedmortalityofeggsandsmall young accounted foranadditional 2,000to 6,.500 nesting cfforts annually. Counts of living young andofyoung founddeadmadeeachyearafterbandingwas completed have shown consistently that from one-third to one-quarter of the birdsoftheyearwerebanded.Each year since 1960 asampleof from 7,000 to 8,200adultSootics has beencapturedin mist nets setattheperimeterofthccolony (figure7).Itshouldbepossible to estimatethenumberof adults accurately fromtheproportion ofbandedindividuals occurringinsamples takenlaterthesamebreedingseason. But calculations from May-banded adults in samples of adultsnettedthefollowing July yield population estimates consideredthreetoRvetimcs too high.Twocharacteristics oftheTortugas SootyTernpopulation, strong localization of individuals within the colony and straggling arrival


:18 BULLEn',FLORIUA STATE Vol. 8 and departure, hamper use of mark-recapturedatafor estimating total numbers. Banded adults do not become randomly distributed throughout the colouy,andthesampled population changes in com position from week to week duringthebreeding season.FIGUnE7. Memher.; oftheFlorida field excursion, 13th InternationalOrnitho logical Congress, mist.nettingadult SlMltyTerns onthewestbeachof Bush Key.Inforeground fromthe left, Josias Cunningham (U.K.) and Staffen Ulfstrand(Sweden). The matted t'over is sea purslane' (Sesudum pnrlulacastrum).(PIlOtogra\lh hy A.Schifferli, 1-1 June1962.)DISCUSSIONQnly a limited interpretation oftherecordofthepopulation of SootyTemsattheDryTortugas (table2) can bcundertaken now.Thcpresent discussion aims merely to review the cstimates inthelight of thc species' behavioral characteristicsandtheTortuganenviron mental factorsthatmay havc influenced them over the years. Somc ofthelocal limiting factorsandthedifficulticsofaccurate censusing were recognizedanddiscussedbyearlier writcrs.Thepreliminary results of worknowin progress include some additionalpertinentinformation. Commentsarclimited tothepopulation records from 1903 to datc.Thefactthatall adults inthecolonydonotbegin to nestataboutthe same time has plagued SootyTemcountersatDryTortugas from


1964ROBERTSON: DRY TORTUGAS TERNS39the beginning.Thevarying population reports of 1903 (Dutcher, 19(4) resulted inpartfromthearrival ofmanySootiesafterJobandBurtonhadmadetheir first estimate.Watson(1908: 318) specified that his 1907countwasmade"late inthebrooding season,afterall the eggshadbeenlaid."Laterobservations, particularlybyMoore intheearly 1950's, makeitclearthatsimply todelaythecountuntil all the birds arrive isnota satisfactory solution.Itmaybeuseful to reviewwhatis knownofthearrivalandlanding of Sooty Terns attheDryTortngas. A period of nocturnal swarming overthebreedinggrounds before actual nesting begins ischaracteristic ofthespecies(ct.Ashmole,1963a).TheBrownNoddyalso exhibits this behaviorattheDryTor tugas,anditmaybepartofthepre-breeding activity of all pelagic terns, although observationsonspeciesotherthanthe Sootyarefew. The occurrenceofSooty Terns on regular nocturnal visits inlatewinterwasnotreportedattheDryTortugas untilratherrecently (Vin ten, 1944; Baker, 1944).Ourknowledge ofthephenomenoncoincides closelywiththeperiodofSootyTernoccupancy of Bush KeyandNational Park Service occupancy of adjoiningGardenKey. Park Service personnel stationed onGardenKey havckepta complete record ofthedatestheSoatieswerefirstheardovertheareaatnight and first secntolandon Bush Keybyday annually since 1943. Thctwo eventsaretheoutstanding phenologicalphenomenaof thc, for some,ratherhumdrumyearatthcDryTortugas,andone suspectstheyhave oftenbeenrecordedaswelcome evidence ofthepassage of time.Datesofthefirst night occurrence oftheSooties rangc from 8February(1943 and 1956) to 7 March (1950) withanaveragedateoverthe 20-year period of 20 February.Theaveragedateof the first daytime landingisalmost exactly 2 months later,21April. Typically tllenumberof birds,asjudgedbytheamountofnoise, begins to increase nightly soon afterthefirst report,butfor a montll tosixweeksnoSootiesaretobeseen in tlle vicinitybyday.Theirdaytime whereaboutsduring thisperiodisregardedlocallyassome thingofa mystery,butmyrecords of 24-26February1964 suggestthatthebirdsfrequenttheGulf of Mexicoprobablyatnogreatdis tance. IheardSooties calling asearlyas2030 hours (all times EST) andaslateas0615 hours,both 2.'5 February.Theyseemedtoap proach fromthenorthwestandthelast birds over Garden Keyneardawn seemed todepartinthatdirection. Activity onthenightof25-26Febrnary1964 centeredaboutone mile north-northwest of Rush Key. Observation from aboatin this


40 BCLLETI:-.I FLOHIUA STATE NIUSECM Vol. 8area disclosed several separate flocks, each apparently ofmanythou sands, millingaboutin rapid flight within a few feet ofthesurface ofthewater.Noappreciable movementtowardthe colony site on Bush Keyoccurredfrom 2100to0130 hours. At least a month beforethefirst daytime landing, however, Sooties havebeenreportedland ing on Bush Keyatnight,andoccasional precocious females lay eggs during nocturnal visits asmuchasthree weeks before nesting begins (RobertandStevenson, 1951).Thenight landings presumably cor respondtothegatherings on Ascensiontermed"nightclubs"byAsh mole (l00sa: 301ff.),butnodetailedobservationsareavailable fromtheTortugas. A few days to a fortnight beforethedefinitive landing afewSooties oftenremainon Rush Key wellpastdaybreak, and scattered birds and occasionallargeflocksareseenatsea ncarby. Soonafterthese first daytime sightings Sootieseitherlandatnightandremain ontheislandorbegin landingbyday, usually in early morning.Thenumberthatlandthefirstdayreportedly varies from a fewhundredtomanythousands. Elms (personal communication) estimated 40,000thefirstdayin 1963.Layingbeginsatonce. Commonlyhundredsofbirdsareincubatingbytheafternoon ofthedayof landing.Newflocks arrive nightly foratleast several weeks. :\foore andDilley (195.'3) notedthatthespreadof hatching dates indicated the periodofarrival was greatlymoreprolongedin some years.Thelargerthecolony becomes, themoredifficultarenewarrivals to detect, exceptastheyoccupyentirelynewground.Inthe usualpatternoflanding, successive flocks settle inunedi ately contiguous tothegroundalready occupied. Almost invariablythefirst Sootieslandonthewest sideofRush Key.Fromthis nucleusthecolony builds eastward alongbothshores,thelast birds to come in landing on theeastspitor(occasionally) Long Key.Felton(1941MS.)suggestedthatthustheSooties first settle ontheoldestpartofRush Key, an intriguingideaimpossible to verify.Inatleast two oftheyearswhenSooties nestedonGardenKey (Beard, 1938; Gibbs, 1947MS.)thefirst birdslandedonthenorth coaling docks (figure6).In1938newflocksbuiltsouthwardfromthatpointuntil all ofGardenKey east ofthefort was occupiedbeforeanylandedonthewest side of Bush Key. Over thepast20 seasons first landings of SootiesattheDryTortu gas have become consistently earlier.Theaveragedateoffirst land ings for 1943-52 was 27 April, for 1953-62 it was 14 April.Theland ings of 7 April 1961, 6 April 1962,:3April 1963,and28 March 1964 are the earliestofreliable record. No similartrendcanbeseen


r1964ROBERTSON:DRY TORTL:CAS TERNS41inthe record of first nocturnal visits. Theseareless likely tobere corded accurately,becausetheyapparentlyarebriefhighoverflightsbya few birds inthemiddleofthenight.Onthenightof19-20 January 1964,about3 weeksbeforethethenearliest report, IheardSooties calling overGardenKeyfourtimesbetween2345and0200 hours.Eachtimeoneortwobirdslewover rapidlyandfairly high, the passagemarkedbythreeto five unmistakable "wideawake" calls. The followingnightitwascolderwithhighwinds,andI listened from2200to 0200 hourswithouthearing any Sooties.Onmynext in late February 1964largenumberswerevisitingtheareanightly. Brief observationsofSooties astheyfirstlandedin AprilandMay1963revealed several interesting characteristics ofappearanceandbehavior inthenewly-arrived birds.Despitethefactthattheypre sumably havebeenonthewing almost continuously for aperiodof several monthsorlonger, most individualsarefatandappeartobeinpeak physical condition. Sootiesappeartobeheavieratfirst landingthanatanyothertimeinthebreeding season; anumberof birds at this time weighed 190 to 210 grams, whereas weights exceeding180gramsareunusual later. Ashmole (1963a: 340fI.)reportedthatmost Ascension Sootieshadcompleted moltbythetimetheybeganto assemble inthecolonyatnightandthesameprobablyistrueof the Tortugas population.Inthehandtheplumageof newly-arrived birdsisconspicuously freshandunworn.Theattenuatetipsoftheouterpairof rectrices extendasmuchasSOmOlbeyondthe adjacent pair in delicate streamersthataresoon lost apparentlybyabrasion on land.InJnneand Julytheoutcrrectrices are only 15-20mOllongerthanthenext pair. Tightlypackedroostinginthecolony is a characteristic group behavior for several days immediately afterthelanding. Allthebirds settleatonceandform a ncarly continuous cover over the ground.Thecolonythenappears much more denselytenantedthanit does after incubation begins, when onlyonemember ofthepairisusually presentatthenest. Also typical of thisperiodareflights, presumablypartofthepair formation ritual,inwhich two individuals stay very close together. In these flights two birds leave the ternery, circle to an altitude of several bmdred feet, then change to an exaggeratedlydeepandslow wingbeat, meanwhile giving a callthatapparently is peculiar totheoccasion.Theflightsmayoccur overthecolony or remote from it. They vary indurationfrom a fcw seconds toaboutaminuteandmay consist of onc ritualized flightorofseveral alternatedwithintervals of normal flight. Flights usuallyendabruptlywiththetwo terns


42 Bl'LLETIN FLORIJ)A STATE MUSEUMVol. 8making aheadlongdescentcheckeda fewfeetabovethewater,andthenflying rapidly back tothecolony. Occasional two-bird Hights occurthroughoutthebreedingseason,butthey aremorefrequentand protracted among Sootiesthathaverecently landed,whendozens of flights maybein progress simultaneously. Brown Noddies on Bush Key perform ritualized two-bird Hightsthatclosely resemble those just described fortheSootyTern,andIhaveoccasionally seen similar flights by Royal Terns winteringattheTortugas.Inallthrecspecies a distinct call is associated withtheflightsandoftendrewmyattention tothebirdsengagedin them. Warham's(1956: 89-90)description of a "ButterflyFlight"of Brown NoddiesandBlack NoddiesonPelsart Island,WesternAustralia, seems toapplyequallywell to flights observedattheTortugas,andpresumably allrepresentthe"High Flight" aerial displaythathasbeendescribed formanyterns.Sprunt(1948b: Ill) andseveralotherobservers suspectedthatthedateof sampling in a given yearmightaffecttheestimate of popula tion,hutno means of quantifyingthesuspicion existed until \'1oore markedplots on Bush Key in1952.In 195:3 thefirst Sootieslanded14Aprilandthefirsthatchingwasnoted23 \Iay. \Ioore'snest counts thatyear (19.'54 MS.)showed64,724adultspresent14 \Iay, 81,210on 2:3 and84,569on28 \Iay. Fromthese figureshecalculatedthatanaverage of1832newbirdsenteredtheternery daily from 14 to23May,andtheratedroppedto672daily from 2.'3 to28 \Iay. Nolaterchecksweremadethat season, hutgeneral ob servations ofthecolony suggestthatSooties continue to arrivc andstartnestingthroughmuchofJline in some years. Moore(19.'54:\IS.) suggestedthat,to assurecomparablepopulation estimates from year to year, density of nestingdatashouldbebasedoncounts of nestsmadeone week after first hatching. Thisperhapsisthemost practical solution possible,but the waySooty Terns arrive to nest clearly makesitdifficult to estimatethesize of a colony ac curately except fromrepeatedcounts spaced tosampletheentire season. Population estimatesbasedonnesting densitiesmadeeither muchbeforeormuchafterhatchingbeginsarclikely tobetoo low. FromtheknownpatternintheCommonTernitappearedlikelythatlate-arriving Sooties includetheyoung adultsreturningtothecolony forthefirst time,andthatageatfirstreturnis 3or4 years. Several returns recorded in1937-41,however, seemed to show Sootiesbandedas young oftheyear back intheternerythefirstorsecond year after handing. \Venowbelieve these reports resulted from mistakes inreadingorrcportingbandnumbers.


r 1964ROBERTSON:DRYTORTUCASTERNS43Noreturnsofthe5500juvenilesbandedinJune1959werere corded in handling a total of19,327adultSooties(927returns) in and July1960and1961andinMay1962 (4513, 426returns),butthe198returnsprovidedbyasampleof4190adults taken8-15July1962included11ofthe1959cohort. Samples of adults mist-nettedatthe colonyin1963on8-11April(1125, 140returns),15-19May(4021,685returns),and9-14July(3807, 362returns) contained0,6,and50respectively ofthe1959juveniles.TheJuly1963samplealsoincluded .'3 returns fromthecohort of10,127juvenilesbandedinJuly1960.Itthus appearsthatSooties first return tothenatalcolony late in theirthirdyearandfirstreturnin forcelatein their fourth year.Ourdatashowthatalthough0.2percentofthe1959cohort of young Sooties rcturnedtoBush Key intheirthirdyear, only 0.00 perceutofthe1960 cohortdidso. This is of interest in viewofthe recovery record ofthetwo cohorts since leavingthecolony. For the youngof1959nota single recovery hasbeenreported; for the1960groupwehave13recoveries,6ofwhichwerebirds found dead alongthestorm trackofhurricaneDonnaofSeptember1960.This suggeststhatthe1960cohort sufferedmuchheavier mortality during its extra-Torhlgan years. Late-arriving adults apparently often pioneer in the changcs of colony site. Bartsch(1932: 281)in rcportingthefirst move of Sooties fromtheancestral Bird Key location observedthat30pairs nestingonBush [Long] Key stillhadeggsorsmall young on10-21August1932while most oftheyoung inthemain colonywerealready on the wing. All oftherecorded nestings on Hospital Key and Long Key were wellbehindtheusual scheduleandpresumablywereiniti atedbybirdsthatarrivedlateand failed to find space intheparentcolony. Latc-nesting Sooty TernsattheTortugas seldom succeed in rear ing young, inpartbecausethcyso often nest.inunsuitablc places sllch as the easily Hooded sites on Long Key,andinpartbecause isolated nesting groupsareespccially subject to predation.In1963,for example, wedetectedno significant loss from predation inthemain colony,butpredators(both ratsandCattleEgrets suspectcd) dcstroyedtheeggsofanestimated1,500pairsofSootiesthatsettled ontheeastspitofBush Key (figure .'3) late intheseason.NoSooties hadlandedontheeastspit24Aprilbuton15-19 :'vlay itwas fully occupiedbyincubating birds and othersthathadnot yet laid eggs. Attackbypredators must have occurred soon after, because no Sooties remainedthereon 5 July, and broken eggsthatranged from slightlyincubatedtoaboutreadytohatchwerescattered overtheground.


44BULLETINFLORIDASTATE MUSEUMVoL 8In1963 mostofthe3and4-year-oId adults apparently arrived duringlateMayandJuneandsoughtspaceinthemain colonyratherthanatits edgesoratoutlying sites.About95percentofthereturns forthejuvenile cohorts of 1959 and 1960werelocalized inthesouth western one-quarterofBush Key,thesameareain which chicks werebandedmost heavily in those years.Largesamplesofadults taken farther eastonBush Key (including a sample of 329 from birds then landing ontheeast spit, 16 May) included feworno returns of juve niles bandecl in 1959and1960. Thustheyoung adults returning forthe first time seemed tocentertheir activitiesnearthenatalnest location, eventhoughthatpartofthecolony already was densely occupied. Although strong site tenacity in terns (Austin, 1949) un doubtedly servestomaintaintheestablished colony,itmust also weigh heavily againstthelikelihoodofsuccessfulbreedingbyyounger adults. 'V'e havenoclear evidencethatany Sooties nested in their thirdorfourth years.Wesuspectthatinexperience,latear rival, and site tenacity combine to make successfulbreedingbyyoung adults a rare occurrence,atleast in colonies whereadultmortality lowandspace relatively limited. Stragglingdepartureis as characteristicoftheTortugas Sooties as straggling arrival,butthis aspect of seasonalchangeinthepopu lation has seldombeenmentioned.Earlywriters believedtheterns left in onebodyor within a few days. Thompson's (1903:82) statementthatthe Noddies leave "ingreatflocksandatnight....Theentire exodus consumesbuttwoorthreedays"istypicalofcom ments forbothspecies. Later, Bartsch (1919: 473)quotedreports oftheBird Key wardens to showthatnoticeable mass departures oc curred over a period of 2 to 6 weeks. Morerecentobservations(.'On firm thisanddo not extendtheextreme dates Bartsch mentions, 9 Augustand25 September. Although a decrease in the sizeoftile colonyisseldom obvious before mid-August, several lines of evidence suggest departures beginmuch earlier. Birdsthatdo not renest aftcr failing intheir first breedingattemptprobably begin to leave the terneryinMay.Eggremoval cxpcri mentsbyRidleyandPercy (1958) on Desnoeufs Island, Seychelles,andbyAshmole (1963a) on Ascension showthatSooties arc far less persistent layersthanhasbeensupposed. l'i' a morethan.50percentof those whose first eggs were removed laid a replacement, and re nesting seldom occurred after lossofwell-incubated eggsornewlyhatchedchicks.Thefew observationsatthe Tortugas seemtoagree with these findings, and suggest in additionthatthelikelihood of renesting begins to decline sharply at a relatively carlydatcinthe


r1964ROBERTSON: DRY TORTUGAS TERNS45breeding season. Birds whose first eggsweredestroyedbypredatorsonthe east spitofBush Key in late 1\Iay orJune1963 laid no replace ments there. "Ve found nothing in July to suggestthattheyhadrenested elsewhereonBush Key,nordidthereseem tobeanysub stantialnumberofunemployed adultsaroundthecolony. Appar ently Tortugas Sooties whose first breeding effort ends in failure afteraboutmid-Maytendto leavethecolony soon afterward without renesting.Theearliestdeparturefor which definite evidence exists isthatof anadultbandcdon Bush Key in :\Iay 1960andfounddeadatRuskin, Hillsborough County, Florida,about 215 miles northbya little east oftheTortugas, on25July 1960.Otherbandingdata, however, suggestthatmanyadultSooties leavethecolonybetweenlate Mayandearly July.Largesamplesofadultsweretaken in mist netsinboth J\Jay andJuly, 1960through1963. Extreme datesofthesampling periodswere 15-.31l\1ay and8-17 July,andtheloca tionandmethodofcapturewere virtuallythesame for all samples. The 14,884 adultshandledin July 1960-1963 included 160 (1.08pCI'cent)bandedin May ofthesame year, whereas 12,100handledinMay1961-1963included412 (3.4percent)bandedin May oftheprevious year.Therangeinthevarious samples was: \Jay toJuly repeats, 0.65-1.6percent; May to May returns, 2.8-3.9percent. May-banded adults thus occurredthreetimesmorefrequently in samples ofthefollowing Maythanin samples taken 5 to 6 weekslaterintheyearofbanding.Thesimplest explanation isthatmanyadultspresentin May leave beforethesecond weekofJuly.Thesepresumably in cludebothfrustrated breeders whose ties tothecolony are relaxedbyloss of eggsoryoung and early breeders whose younghaveHedged. A few dozentoseveralhundredSooties usually remain on Bush Key aftertherestaregone. Mostarc jnvenilesandmostaresick, injured,ordeformed.Onlyrarely do any survivethefall Hights of accipitersandfalcons in late SeptemberandOctober. Tortugas Sooties seldomabandonhealthy young, although reports suggest thismayhavehappenedonceortwicewhendeparturewashurriedbysevere storms inlateAugustorearly September.On8-11 September1962about50 young birds remained on Bush Keyduringthe day. Mostwereobviously infirmandseveral died every day.Eachevening 200-300 adults andabout100 strong-Hying young re turnedtothecolony. Alltheyoung were still being fedbyadults andtherelative numbers of youngandadults suggestedthatbothmembersofmost pairswerepresent.


46 BULLETll\ FLORIDASTATE Vol. 8Thecarlydepartureofsome adultsprobablyhas notbeena major sourceoferror in population estimates (table 2).Thecritical event for estimates based onnestcounts isdcpartureof hatchlings fromthenest site, which becomes important even earlier. Ncst densitydataused in calculating populations in 1945-1948werere corded after mid-Juneandthus may considerably underestimatc actual numbers. Nest countsof1951-1956wcremadein late Mayandin severalofthese years large numbers of adults werethoughttohave arrivedandnested afterthecounts.Theknowledgethatyoung Sootiesdonotreturninforce until their fourth year makes the reported increasesof1903 to 1907, 1913 to 1914, 1939 to 1940, 1942 to 1943,and1949 to 1950 highly improbable from Tortugan recruit ment alone. Those of 1938 to 1939, 1943 to 1944, 1947 to 1948,and1955to1956 seem unlikely inthattheyleave little room for mortality intheintervening years.Thereportedpopulations of 1950and1951 standoutas much above other estimates. I have reviewedtheficld records for these yearsandtheerrors, if any, are inthedata,notinthecalculations. Moore and Dilley (1953:79)suggestedthattheunprecedentedly high populations of 1950and1951 "maybeattributabletoseveral years being unusually favorable forweatherandfood." However, evidencethattherelevant years, 1946and1947,weremarkedbyespecially successful rcproduction is wanting. In spiteofthequestions raised above, the reports since 1903 prob ably reprcscnttheactual trends of populationwithfair accuracy.Ingeneral,theBird Key colony of Sooty Tcrns increasedunderprotection toabout80,000 to 100,000 breeding adultsbyaround 1917.Thesevere hurricanesof1910,1915,and1919thatultimatcly caused agreatrcduction inthenumberofpreferred ncst sites available to Brown NoddiesonBird Key probablymadeenough moreareaavail able to Sooties to compensate for thc area lostbyerosion.Inany eventtheSootyTernpopulation apparently maintainedabout tile same lcvel fromc.1917 toc.1930. Disturbancc resulting from renewedegging intheearly 1930's,andprobably also from tile en forced movement of the colony from Bird Key, seemsadequateto account forthereported decrease to about 30,000 adults in 1935. Within a relatively few yearsafter1935thecolony,nowonBush Kcy, again attained approximatelythcsameupperlevelthatithadon Bird Key.Theviewthat60,000to100,000 adults representsthcnonnalSootyTernpopulationofDryTortugasunderprotection (MooreandDilley, 1953) probablyisclose tothemark. Fluctuation within these limits doubtless results in largepartfrom varying success


r1964 ROBERTSON: nllYTORTUCAS TER1\S 47inrearing young because ofycartoyearvariations iuweather,food supply,andpredation,andfrom varying mortalityduringthepopu lation's pelagic phases. PredationisseldomimportantatDryTor tugas, although Magnificent Frigate-birds take fair numbers of young Sooties in some seasons (Beard, 1939; Dilley, 1949MS.)andinstancesofpredationbyrats, cats (Russell, 1938MS.),LaughingGulls, atricillaLinnaeus (Watson and Lashley, 1915: 38),andaGreatWhiteHeron,Ardea occidentalisAudubon(Robertson, 1962),havebeenreported.Itisnotclearwhatfactorsactto settheupperlimit attainablebythe Tortugas SootyTernpopulation, norhowtheyact,butI suggest tentativelythatthelimits maybedeterminedasmuchbythespecies' behaviorpatternasbysuch environmental factors as foodandter ritory.ThequestionofwhetherornotSooties areevercrowdedon Bush Key hasbeendebatedbyauthors tonOconclusion.Itis clearly a strongdeparturefrom normal behavior, however, for Sooties to nest elsewherethanatthecolony site ofthepreviousyearorattheedgeof a mass of Sooties already nesting.ThatTortugas Sooties rather frequentlyhavesettledatnewlocations suggeststhatBush Key hasbeenovercrowdedattimes, howeveritmayhaveappearedtohumanobservers.Theobvious questionthenis,whyhasn'tthecolony spread to nearby islandsthatseem fully as suitable as Bush Key?Thereasou appears tobethattheearlier and more successful breederstendstrongly to settleatthecolony site of the previous year.Presentdataindicatethatwhentl,eprogenyof these birdsreturnto nest,theyseek nesting spacenearthelocationwheretheywerereared. Such apatterntends to maintain a strong nucleusattheexpenseofpossible colony expansion.Theindividualsthatcolonize peripheral or out lying locationsarethose compelled todoso, principallybecausetheyarrive lateatthe colony.Asagroupthesemaytendtobechronic unsuccessful breedersthathave lost site tenacity.Thenewloca tionstheyoccupyarecommonlymuchmore exposedtoweatherand predators,andlate arrival reducesthelikelihood of renestingafterdisturbance. Thus,thepioneeringthatmightleadto establishment at new locationsandan increase inthelocal population is almost always foredoomed to fail.Thesuccessful shift ofthecolony site from Bird Key to Bush Key in the early 1930's wasprobablyfacilitated, once theareaofBird Key wasreducedto a certain point,bylandings on Bush Key early enough in the seasonbylargeenoughnumbersof hirds for successful breed ing.Thebehavior ofthecolony in 1937,1938, and 1947whenthe


48 BULLETI:II FLORIDA STATE :\ICSEUM Vol.8first Sootiesthatarrived settled onGardenKey is less easily explained.Inthese eases, however, unusual conditions apparently existed on Bush Keyatthetime nocturnal swarming began, an infestation of rats inthe1930's (Russell, 1938MS.;Beard, 1938)andlUmsually lux uriant vegetation inthe1940's (Sprunt,1948h).Perhaps these dis turbancesweresufficient toproduceatypical behavior.Itwas formerly believed (Murphy, 1936: 1125-1127)thatall Sooty Terns deserted their nesting areas for a period of timebetweenbreed ing seasons. lV!ore complete information, however, showsthatSooties are present intheneighborhood of some coloniesthroughouttheyear. Ashmole (1963a: 301)states"...therewas no month in which \Videawakes couldnotbeseenorheardfrom Ascension."Thesame appears tme of colonies off Oahu, Hawaii, studiedbyRichardson and Fisher (19,50), and of thoseatWillis Island (Hogan, 192,5) and Raine Island (Warham, 1961), northeastern Australia. Ashmolc and others havedrawna contrastbetweenthecolonieswhereSooties are continuallypresentandlatitudinally more peripheral colonies, suchastheDryTortugas, wheretheyareabsentfor several months oftheyear,butthesupposed differencemaydisappearwithmore study. Excepting birds carried northbyhurricanes, all recoveries (through 1963) of Sooty TernsbandedasadultsattheDryTortugashavebeenwithintheGulf of Mexico, indicatingthatthebreeding population doesnotdisperse widely.TheJanuary records cited above leave NovemberandDecemberastheonly months in which SootieshavenotbeenreportedattheDryTortugas.Thepossibilitythatsome individuals remain within commuting distanceandmake occasional night flights overthecolonythroughouttheoff-season cannotatpresentbeexcluded.TheSooty Ternsofthe Tortugas have oftenbeencited as typical ofthepopulationsthathave a 12-month cycleandbeginbreedingataboutthesame time every year. This appearstrueofrecords from the time of Audubontotheearly 1940's, all of which indicatethatlaying began inlateAprilorearly May. Overthepasttwo decades, however, first eggshavebeenlaidatconsistently earlier dates.Thecycle remains essentially annual,butnesting now begins a fullmonthearlier thanitdidinthe1940's.Thesignificance of this slow ad vance of breedingdateandthefactorsthatmightaccount foritareunknownatpresent. No relationship to a particular moon phase (Chapinand'Wing, 19,59; Ashmole, 1963a: 349)isapparent.Thedateof landing and first eggsin1964 coincided withthefull moon,butthe landingsof1961-1963 occurred 23, 14,and6 days respectively beforethefull moon.Theearlierbreedingof Sooty Terns inthe


r1964ROBERTSON: DRY TORTUGAS TERNS49Tortugasmaybemerely another phenomenon ofthesortthatis commonlyattributedtoa supposedtrendtowardwarmerclimate. Another possibilityisthatearlier breeding is associatedwithanin crease inthesizeofthecolony, althoughclearproof ofanincreaseislacking. FisherandLockley (1954) cite instances formanyspeciesofseabirds showingthatlarger coloniestendtobecomc active earlierinthe scason. RichardsonandFisher (1950) reportedthatSooty Terns on two small islands locatedabout10 milesapartoffthewindwardcoastofOahuhaddistinctly different breeding seasons,thecolony on :'vlokn Mann beginning tonestin November whilethaton l\'lananabeganinApril.TheysuggestedthatMananamighthavebeencolonizcdbybirds from Midway Islandorsomeotherdistant population which breeds inthenorthcrn hemisphere spring.ThcpossibilitythatSooties nesting onMananaarethcoverflow fromthelarger colony on Moku!vlanu,however, does notappcartobeexcludedbytheinformationsofar publishedaboutthese populations. Egg-laying on :'vloku l'vlanu bcgan in Novemberandextended through :'vJareh, whiletheseasononMananaisshown as April throughJune(Richardson and Fisher,1950:304,table4).Thustheseasonon :'vIanana merely continued that of 'vlokn Mauu,ratherthanbeing distinct from it. This char acteristic,thefactthatthepopulation on tvlanana varied greatlyinthe two breeding seasons obscrvcdandthatfew yOllngwererearedineither season,andthefactthatthcMoku :'vlanu colony was re portedly overcrowded all suggest pioncering of a newsitebybirds that arrived late, such as hasbeenobserved several timesattheTortugas. Ashmole (1963a) recently publishcd a highly informativeaccountof tw'O breeding seasons of the Sooty TcrnsofAscension Island.Fromhisobservations of Sootiesandotherspeciesthatbreedtherehe ad vanced (Ashmole, 1963b)thehypothesisthatcompetition for avail able food within foraging rangeofthenesting colony wastheprin cipal factor regulatingthenumbersof tropical seabirds. No single study as intensive as Ashmole's hasbeenmadeattheDryTortugas, but, because ofthelong record of observations availablc,theTortu gas colony ranks asperhapsthebest-known SootyTernpopulation aftcr Ascension. A comparison (table 3) shows striking differences betweenthetwo populations in mortality factorsandbreeding suc cess. Althoughitisoftheorderof one-tenththesize oftheAscension population,theTortugas population appearstohavereared a sub stantially largernumberofyoung in some years.


.50 BULLETI"FLORIDA STATE MUSEl:MVol. 8 Ashmole (1963a) describes heavy mortality of SootyTernchicks, apparently from starvation,andprescntsotherevidence indicatingthatadultshadgreatdifficulty obtainingadequatefood for young, especially inthesecond seasonheobserved.Hecites (1963b:46.5)records similarly consistentwithhis hypothesis of population con trol fromotherSootyTerncolonies and forotherspecies of tropi cal seabirds. Feeding conditions apparently are somuchmore favor able for Sooty Terns intheneighborhoodoftheDryTortugasthatthe likelihood of their numbers being limitedbycompetition for available food seems improbable. Recent observationsattheTor tugassupportWatson's (1908: 192-195) statementsthattheternsdomost of their feeding within a radiusofabout15 miles fromthecolony. Although muchofthe feeding oecurs outsidetheTortugas lagoon,Hocksof Sooties often fish within sightoftlle colony. Fishandsquidsregurgitatedbyadults returning from feeding commonly are intact,asif takenbuta few minutes before. By contrast Aslunole (1963a: 333) foundthatfeeding adults wereabsentfor extended pe riods;hesawno Sooties fishingnearAscension;andthefood remains regurgitatedbytlle birdswercseldom recognizeable.From his sh.dies of the development of young on Ascension, Ashmole concluded (l0003a:0320): "The capacityofyoung \Videawakestosurvive for long periods on relatively little food, while growinghardlyatall,butto accept large quantitiesoffoodwhenitis available, is clearly an adaptationtoanenvironment in whichthefoodsupplyisprecarious." While few detaileddataare available onthedevelopment of young SootiesattheTortugas, general observations indicate tIlat tIleir growthisregular and rapid andthatthe abilitytoflyshortdistances is at tainedat5 to 6 weeks of age, in contrast totheHedgingperiodof 8 weeksorlonger recorded for Ascension (Ashmole, 1963a: 321). incidents of mass starvation of yonng SootieshavebeenreportedattheTortugas. Acknowledgingtheneed for additional critical data,itappearsthatno shortage of food available to SootyTernsexists inthevicinity of theDryTortugas colony; and, therefore,thatcompetition for food duringthcbreedingseason cannotbethefactorthatchecks the in crease ofthepopulation short of the limits of available nesting terrain.Assuggested above, socialfactors-inparticular, site tenacity in young adults, andthetendencyof late arrivals to choose insecurenestsitesnearthecolonyratherthm. more secure sitesata littledistanceappearto operate in a density-dependentmannertolimit growth of population. Ashmole(196.'3b)issurely correct in pointingoutthatcompetition for nesting spacecouldnot regulate total species popu-


r1964 HOBERTSON: DRYTORTUGASTERNS51lations of scabirds effectivelybecause individuals thatfail to find space inonecolony couldgoelsewhere.Howmuchmovementofthissort actually occursinthcSooty Tern is a mootpointat present.Itseems likelythatasynchrony ofbreedingcycleswouldinhibitex change of birdsbetweensomecolonies. \Vhateverthefactors are that ultimately controlthetotalnumberof Sooty Terns, however, observationsattheTorhlgasstrongly suggestthatsocial forcescaneffectivelyregulatenumbersattheleveloftheindividual colony. TAHLE.'3.SOI\O: CHARACTEIUSTICSOF THE SOOTY TEH:"lrOPULATIO:"lS OF ASCEt\SIO::'IJISLANO(t'HOl\1ASIL\IOI.E,196:1a) ANDTHEDTIYTOltTUGASEstimatednumberof Breeding adults spaceavailable Mortalityofadultsatthe eulony Mortality ofeggsandyoung caused bypredation of eggs and young causedby weather resultingfromattacksuponstraychicksbyadult Sootics of}'OlUlgfromstarvation OWT-allhreeding suc ceSs (young Hedged as a '1, of eggs laid) Interval from startofonebreedingperiod to startof the next. Ascension Islandc.750,llOORelatively nnlimited Estimated 1to 3% killed by feral cats;predationproportionally heavier on theear Her hreeders. Heavy (Celts and frigatt' birds)Noinformation Fn.'qucnt, much heavier inSOlliebreeding seasons. Low, eslim.ated at c.10 andperhapsTlotover 2% intwo suc('essive breeding periods. c.9.7 monthsDry Tortugasc.80,OOllLimitedonBush Key, severalotherhabitable islands nearby.Insignificant,virtuallyall losses from birds becomingentangled invegetation. Usually minor(mainly rats), exceptinoutlyingnesting arei.\s. Occasionally heavyatlow sites and \\'hen hurricanesoccurduringthebreedingseason.Probablythe 111

Bl:Ll.ETI:'II FLORIDASTATE \[CSEUM BROWNNODDYRECORD OF NESTINGVol.819th Century.AccountsbynaturalistswhovisitedtheDryTortugas in this period suggestthatBrown Noddieswerenumerousandidenti fy the keys on which they ncstedatdifferent times,butbcyondthatcontribute relatively little tothehistoryofthcTortuganpopulation.Tn1832 alltheNoddies were nesting on Bush Key,but"several thousand" nestsnotin use were seen on Bird Key as well, leading Audubon (1835) to supposethatSooty TernshaddriventhcNoddies fromthclatterislandnotlong before. Bartsch seems to havethoughtthatNoddies persisted in ncstingapartfrom thc Sooties until Bush Key was washed awayaround1870.Hewrites (1919: 482): "Since then [1832]thecolony has been forced to make a complete shiftandthe choicebetweenBirdandLoggerhead Key has fallentotheformer.... \Vurdemann(1861) andBryant(1859a),however, re portedbothspccies nestingonBirdandEastKcy inthe1850's.Theseparation of the nesting are,lS of SootiesandNoddics observed by Audubonmaywellhavcbeen temporary, for hisreportseemstobetheonly recordthatNoddies nestedonBush Keyduringits 19th century emergence. In 1890 (Scott, 1890) most oftheNoddies were nestingonBird Key. Severaloftheearly reports suggestthatNoddies and Sooties werethenaboutequallyabundantattheTortugas. Audubon (1835: 268)wroteoftheNoddies: "They nearly equal innumbertheSooty Terns.... Scott (1890) wasinfonnedthatNoddies weremore com mon than Sooties, and Holder's (1892: 194ff.) account ofanegging sortie to Bird Keyabout1860 suggests tIle same.Laterwritershavetaken such comments to indicatethattheNoddy population oftheDryTortllgas suffered particularly severe reduction. Job, for cxample,rcmarked(1905: 87-88)ofhis obscrvations in 1903: "OftheNoddiestherearehardlya thousand, which is agreat de crease fromthenumbersthatwcre once here."Itappearstomethatthercports ofnearparityin numbers of SootiesandBrown Nod dies are more likely evidencethatthepopulation of Sootieshadbeenmuch reduced.TheBrownNoddyattheDryTorulgas has always nested mainly in bushes. Noreportsuggests otherwise. More spe cifically, its nestingisconfined largely totheedgesofclumpsorthickets ofbaycedar.Fewnests are placed within dense shrubbery. Fromwhatis knownofthevegetationofkeys where theterneryhasbeenlocated,itseems certainthattheSooties, nesting indensemasses ontheground,wouldalwayshavebeenable to reachmuchgreater


1964 ROllEHTSOK: DRY TORTUGAS TERKS numbers thantheNoddiesbeforetheir increase was limitedbyscarcity ofnestsites.Itseems likely also that,inatemerysuch as the DryTortugas, the Noddypopulationmightbeexpected to decline more slowlythanthe Sootiesunderthepressure of sustained egging. EggerspreferredSootyTerneggstothoseofotherterns.Manywriters mention this, Audubon, for example, informing usthateggsoftheSooty are "de licious, inwhateverway cooked ...." Recause Sootiesnestcloser togetherandaremuchmorestrongly territorialthanNoddies, re peateddisturbanceofa mixedternerywould almost certainly re sult in disproportionately high mortality of SootyTernchicks.ThefactthatNoddynestsaremorescatteredandplacedinheaviercoverwould makeitmuchmoredifficult for eggers togatheranentirelaying.Finally, the usual nesting seasonofNoddiesattheDryTor tugas is considerablymoreextended thanthatoftheSooties which, again,wouldmake loss of an entire season'sproductionless likely.1902.Thompson(1903)presentedanexcellentandwell-illustrated life historystudyoftheBrownNoddyas observed on Rird Key in1902.Hisaccountincludestheearliest clearly stated estimate of the size ofthepopulation: "As nearly as canbejudged itrthe Nod dy colony] containsahoutthree thousand individuals."1903.AswiththeSootyTern,several estimates of thenumberof Noddies are available from1903observationshyBurtonandJob. They are"about400" ,lid"at least...600"(Burton inDutcher,1904),and '11ardly a thousand" (Job,1905).All summaries ofthehistory ofthepopulationcitethenumberin190:3as400andcredititto Job.Thewarden's end-of-season figure of600,however, seemsthebest estimate available.Thewarden on Bird Key (in Dutcher,1905)saidthattheternshada successful season in 19m" and r>layer (inDutcher,1906)reportedin1900thatNoddiesOnBird Keyhadincreased since1898butnotso much astheSooties. Nothing elseisknownaboutthecolony of Noddies fortheperiod1903-1907. 1907.Aspartof hisremarkablyvaried investigations on Bird Key the summer of1907,\Vatsonmadethefirst known estimateoftheTortugaspopulationofRrown Noddiesbasedon a direct countoftheirnests.Hepublished two explicit descriptions of hismethodandresults (Watson,1907: 311, 1908: 197).Thelatterreads: "By means of a mechanicalcountingdevice it was found possible actually


BULLETIN FLORIDA STATE MUSEL:M Vol.8 1 to countthetotalnumberof(active) Koddy nests.Thecountgave 603 nests.In somc places,whercthebay-cedarbushesarevery denseandtheareahastohecovered 'dog-fashion' (orattimes even still more primitively), and inotherswherethecactusgrowthisvery luxuriant, error in countingwaseasily possibly. Onaccountof these possibilitiesoferror, I believethat700 nestsisa more repre sentative number. Sinc'e twobirdsoccupyonenest,wehave atotalof 1,400adultnoddiesontheisland."Despite"Vatson'sabundantlyclear exposition alllaterreferences exceptBent (1921: 303)andLongstreet(1936b,butnot1936a)givethe1907populationas "4000". Iv[any, inaddition, cite "1400" asthepopulationin 1908, crediting this figure also to "Vatson,andtheapparentdecreaschas drawn commcnt:e.g., "The noddypopulationtook an unexplaineddropfrom 4,000in1907 to 1,400 in 1908" (Sprunt, 1947b: 215).Twoerrorsareinvolved hcrc.Theyseem to stcm respectively from a mistake in Bartsch's (1919) accountof \Vatson's observations,andfrom misreadingofBartsch,whogivestwofigures forthenumberofadult :\Toddies on Bird Key.The first (1919: 471) occursinatableandreads: "Noddy tern, estimated,adultl4,000."Thenumeral ''1'' refers to a footnoteonthesamepagethatrcads: "Based uponDodoI' \!If atson' s census of 1908".Thefigure, "4,000," appears tobea lapsltY and, aspointedoutunderSootyTern,"VatsonapparentlydidnotworkattheDryTortugas in 1908.Thesecondreference(Bartsch, 1919: 482) givestheeorrectfigurebutattributesittothewrong year: ... \!lfatsou estimatedthepreseneeof 1,400adultbirdsin1908."Threepoints scemclearfromthctangleof mistaken citations:1. \Vatson's, estimateofthepopulation in 1907 was 1400 adults.2.No estimate ofthepopulation of Noddies in 1908 exists. :3. Com pilershaveoften eited \Vatson from Bartseh Ol" fromoneanotherratherthanfrom \Vatson. \Vatson touchesupona problemthathasplaguedmanylaterobservers of NoddiesattheDryTortugasin hiscomment(1907: 311): ...onefeelsthatthereisa vastlygrcaternumber[than1400] present."HeconcludedthatmanyoftheNoddiesatBirdKeywerenon-breeders.1909-1929.Thereports ofwardensstationedonBirdKeytotheNational Association ofAudubonSocieties (through 1919)andtothcBiological Surveyincludeestimates of tilenumberofadultNoddies for all years oftheperiod1909-1919,andfor 1929 (table 4).The


1964 HOBEHTSON: DRY TORTtJGASTERNS estimate for 1910isbased uponanothercountof Noddy nestsbyWatson.Thewarden's estimate for 1918 was published (in Pearson,1918).Howell (1932: 272) summarizes allthereports, presumablyfromthefiles oftheBiological Survey, mentioning specificallythepopulation figuresof1910,1916,and 1929. Stevenson (1938: 307)alsorefers to the 1929 figure.Othersummariesjumpfrom 1908 to1917to 1935. Allwhoinclude 1917 (Vinten,194:3:57; Fisher and Lockley, 1954: 60;et al.)givethepopulationthatyear as "4,000" citing the figure from Bartsch (1919). Bartsch, however,madeno independent estimateofthepopulation in 1917. Curiously, this esti mate of "4,000"isthesame asthatcredited (mistakenly) to \Vatsoninboth1907and1908.TAfiLE4. POPULATIOKS OF DHOW),!NODUlES :\TDRYTORTUGAS Year of MethodReferenceAdultso 1902 .'3000 EstimateThomllson (HJlJ'l)190:} 600 EstimateBurlon(inDllldwr. 19(4) 1907 1206 1\,,t Connt \Vatson(1908a) (BOO) 1909 5000 EstimatePeacon (1909HIS.) HJlO 1710 .Kest Count Asht' (ms. notes)19112000EstimatePeacon(1911 1I1S.) 1912 1500Estimate Pcacon (1912 Ins.) I'll:} 600Estimate Pe;l{'()] I (1!)J3 1115.)1914 2.300 Estimate Peacon (1914IUS.)191.55000EstimateAshe( Il1S.notes)1916 6000Estimate Belhel (1916lllS.) 1U17 10JIOOEstimate Lrl\'ve(H1l7 Ins.) 1918 15.000EstimateAshe(inPearson,1915) 19ID0'3.5.000 EstimateAshe(1919 Ins.) 1922 1600Estimate (ms.notes) 1929 :3000 EstimatePark(IllS.notes) 19:3.38000 Estimale J\,-tason (1936)19:364000 Estimate DoeanelRussell (1936) 19.'37 2000EstimateLongstreet (19.'37)J9.'3S:392 NestCountBeard (41:3) 1939 .'3S0Nest CountRobinson(19:39) 19:}94.34I\\'st Count Taylor(19.'39 ll1s.)1940 ISO Estimate Rohinson(1940) (continued)


.56BULLETINFLORIDASTATE MUSEUM Vol.8TABLE4(continued) Year of MethodReference Adults .._-1940 7050 EstimateFelton (1940 illS.) 1941400EstimateStimson(inlitt.)19411000Estimate Peterson (inVinlen, 194: 1942 4050 Budlong (in Vintt:'n. 194:3) 194.')625Estimak Sprunl(I946a)1946492 CountSprunt(1946b) (05050) 1947202 ConntSprunt (HJ48a) (250)1948282NestCountSprunt (194Sc)(:300) 1949.566 J\esl Count Dilley(l950) (622) 19.'>0 490NestCount andDilley(l!--);),J)(S:>S) 1951 0518Nest Count and Dilley 1195:l) (0570)19.')2 890 Nest COUllt :\-'1oore,ltH)Dilley 1 195:l) (978) 1905:3 842 !\est Cotlllt lvlooIe(19.')4Il1sJ (926) 19.54970 "Jest Connt \Inon' (1 moS. )(1066) 19505 lIOS C:ount \-f(J(Jf(' (HJ.j,j ms.)(1218) 19622180Kesl Count \v. H. <\ncl Betty ----o Inmany years whenthe breedingpopulation of Brown OOdies wasdeby counting nests, observers added anarbitrary figure(commonly10 per cent) to account for nests nut found. Population estimates that include such additions are placed in parenthes('s beneath the figure hased on the aetua1 nest ('Ol1nt. Figures for 1915-1919quotedfromthewarden's reports (table4)contradict all previous commcnts regardingpeakpopulations of Nod diesattheDryTortugas.Evenifthe figures arc substantially discountedto allow for overenthusiasticinterpretationbytheindivid uals directly responsible,itappears likelythattheTortnganNoddycolony reachedby far its highestpopulationduringthis period.1930-1.934,Theonly referenccstothecolony in these yearsthatI have scenarebyBartsch (sec Sooty Tern) amI his comments include


r1964ROBERTSON:OilYTORTUGAS TERNS57 no estimate of population. In 1931 Bartsch (1931: 373) noted: "TIus change of vegetation [destruction of baycedaronBird Keybystorms]hasforcedthenoddy terns tochangefrom a tree-ne;-ting to a sand nesting habit."Thefollowing year (Bartsch, 1932: 281) mostoftheNoddies still nested on Bird Key,butabout 70 nests were found in densc mats of sea purslane (Sesllvirtm portulacastsum) on "Hush [Long] Key," 32 nests inhaycedarbushes on "Long [Bush] Key," and a few nestsOnpilings aroundthecoaling docks onGardenKey, where apparently few young were reared because most of them fell off into the water. Presumably Hartsch's (1933:267)statementthat"morethanhalf'the colonyhadleftHird Key in 1933 referred to Noddies as well as Sooties. 193.5-1962. Thepopulation figures (table4)require little comment. Data for most oftheyearswereobtainedbydireetCOUlltsof nests. In the years for whichindependentestimates are available, ohserv ers wercinclose agreement except in 1939,1940,and1941. Havingnobasis for a decisionbetweenthetwo figures available for eachofthese years, I hav" included hoth.Theentire colony of Noddies FI(;l:U.. : R. BrO\vn Koddi('!' nesting onGardeD Key. 1938: (n.) pair at nest on apile ufsea oats(Uninla/Janiefdata)cut and rakffi before thetern, landed; (b.)adult incubatinghtid on hare ground amidSC'3 purslane. Park Sel"icephoto!!!aph' by Daniel B. Beard.)


58 nULLETll\ FLORIDA STATE \IUSEU\! Vol. 8nested on Bush Key in most years of this period.Departuresfrom this pattern, all involving nesting onGardenKey,werereportedasfollows: 19.36, 6 pairs nested on docksandpilings (DoeandRussell, 1936); 1937, most ofthepopulation nested onGardenKey (Young and Dickinson, 1937; Longstreet, 1937; Hussell, 1938MS.);1938, nest ing was dividedaboutequallybetweenGardenKey (figure8)andRush Key (Beard, 1938); 1939,aboutthe same division as in 1938 (Rob inson, 1939; Taylor, 1939 1947, 9 nests on Garden Key (Sprunt,1948a);and, 1948, 1 unsuccessful nest onGardenKey (Sprunt,1948c). DISCUSSIOI\' Brown Noddies havebeenhandedandrecapturedattheDryTortu gas inmuchsmaller nnmbers thantheSooty Terns, andbandingdata(:ontrihute little toananalysis ofpastpopulation records.Itseems likely, for example,thatBrown Noddies do notretnrntobreedforthefirst time untiltheyreachthreeorfour years of age,butno proof is available. Reasons for believingthata Brown J\'oddy population is likely to decrease more slowlythana SootyTempopulationunderthepressure of egginganddisturbancearegiven above. Similar reasoning fromtheinformation available on nesting dates, nest site preferences, and mortality of Brown ,,"oddies atDryTorhlgashelpstoexplain parts of the population record (table 3), butthefitisa goodbitpoorcrthanfortheSooty.Themain difficulty in estimating populations of Brown Noddies seemstoarise less fromthcobvious influxofnew birdsthanfrom long-delayed nesting startsbybirds already in the colony. This has vexed nest counters morethanthose who undertook to estimatethenumberof adults in the area.Asmentioned above, \Vatsondoubtedthathiscountofncstsin1907 recordedtheentire BrownNoddypopulation. Somelaterobservers also have feltthatthenumberof nests found failcd toaccountfor the adultsonhand. Obser vationsattheDryTorhlgas in 1960-196:3 suggestthatitisusual for nesting startsbyBrown Noddies tobedistributed over aperiodofatleast 10 weeks from April to early July.Itmaybethattheloafing birds \Vatsonandothers considcred tobenon-breedersweremerely late breeders.Toillustrate, 39 (13.1pcrcent) of 298 BrownNoddyneststhatI examined on Bush Key11-1,5July 1962 contained eggs. Allthatweremarked for later checking contained eggsorsmall young on 2 August, and youngthatwerenot more than half-grown on 8-11 September. 1havenoreason to thinkthatany of these nests repre-


1964ROBERTSON: DRY 'I'ORTUGAS.59 sented renesting after failure of earlier attempts,butthepossibility cannotbeexcluded. Dates of \Vatsou's 1907couutof nestsarenotrecorded,butesti mates oftheBrownNoddypopulationin19:39,1945,1946,and1948 through 1955arebasedoncounts of nestsduringorbeforethethirdweek of June.Itis tobesuspectedthattheseunderestimatethebrceding population.Nestcountsneartheendoftheseasonpresentnodifficultybecausewithfewexceptions recentlyusednestsarecasily distinguished fromanynestremnantsthatmaypersist from the year before.Theunproductiveness characteristic of late nestingbytheSooty TcrnattheDryTortugasseemsuntrueoflatenests oftheBrown Noddy. Nodeclineintheattentiveness ofadultBrown Noddies with young inthenestin earlySeptember1962 was apparent. Barring accidents ofweatherorpredation,theyoung seemed likely to ledge successfully. Such accidents, of course,becomemore likely astheseaSOnof hurricanesandhawkmigration advancesattheDryTor tugas. Factorsotherthanhumanpredationbelieved tohaveaffectedtheBrown Noddies intheTortngascolonyatvarious timesarepredation by rats, mortalitycausedbystormsduringthebreedingseason,andstormdamagetobaycedarbushes.Thepopulationrecordsince 1800 reflects to some extent the recorded occurrences ofratinfesta tionsandsevere storms. Informationbelowonhurricaneoccurrenceistaken from contemporary reports and fromDunnand lodiller (1960)andTannehill (19.so). Theyears from 1900through1910hadonebadsummerstorm,16June1906,andan infestation of rats on Bird Keyissaidtohavebccn climinatedby1908 (Dutcher, 1908b; I\Jayer, 1908). I suspectthatThompson's(190:3)estimate for 1902 wasnearthemark,andthatthe1903 Job-Burton estimate was much too low.Exceptforthe190:3figure,agrecmentbetweenthepopulation recordandtherecord ofdisturbanceisreasonably good.Thecolonyappearsatfirst to have increased slowly;thentohavedeclined slightly,andthen oncemoretohavcincreased slowly to theendoftheperiod. Ihaveseen norecordthatratswerepresenton Bird Keyinap preciablenumbersafter1908.Inthedecade1910-1919 BirdKey wasrepeatedlybatteredbyhurricanes.Thefirst of importance, 15-17October1910,didgreatdamagetothebushyvegetation (see p. 8).Earlyhurricanes ofgreatseverity occurred on 13-15 Augustand3and28September1915,and4 July 1916.In1919theDryTortugas


6UBVLLETIN FLORIDASTATE MVSEU\{ Vol. 8werehitsquarelybya hurricaneofextreme intensityon10-11 Sep tember. Records oftheBrownNoddypopulation for this period, allfromwardenreports,showa declinethroughtheseason of 1913,thena meteoric tothe1919 figurc,thehighestevcrreportedfor the colony. This record doesnottally satisfactorilywiththerecordofdisturbance,ifoneassumesthattheTortugas colony is a discrete population allofwhose sun-iving adultsreturntobreedannually.The1910 hurricane was too late intheseason to have causcdmuchdirect mortality,anditsdamagctovegetationcannothave affected breeding successbeforethescason of 1911. Available information (Peacon, 1911 Ashe, MS. notcs) suggcststhatnestingin1911 was normal.For1912and1913thereports indicate abnornlal behavior andgreatdecreases intheuumberof adults whichtheobserversattributedto scarcityofnest sites. Ashe notes) reportedthatNoddieswereseen on Bird Key on 20 j'vlarch 1912, anunusually early date,butby22 May only an cstimated 400hadappeared, althoughthcpopulationlaterincreased toabout1500 (Peacon, 1912MS.).Inanotherreporton the 1912 season \Vatson (1912MS.)advises planting bay cedar bushes on Bird Key "in large quantities."Thereportfor 1913 (Peaeon, 1913 lists onlyGOOadultNoddiesandcommentsOnthegreatdecreaseofthespecies.Ifthcpopulationdataare consid eredatall reliable,thedecreases of W12 and 1913 musthaveresulted fromthefailureofadults toreturntothecolony.Thedecline seems too earlyandtooabruptto result from less successfulbreedingafter 1910. Interpretation ofthepopulation trcmd for the years 1914-19HJ presents evengreaterdifficulty.Itisknown (Bowman, 1918)thatshrubby growth on Bird Keyhadrecovered to a considerable extentby191.5-1916.The1910 stormmayhave greatly increasedtheamountof thicket edge, andhencethenumberof available nest sites,hybreakingupfonnerly solid standsofbaycedar.Ifso, conditions favorable to a rapid increase of Noddies may have existedby1914 or 191.5. Theyear-to-year increase, however,ismuchtoo large toheaccounted for entirely bythesuccessfulbreedingof a discrete Tortugas population. the summer stomlS of 191.5 re portedly caused heavy mortality ofadultand young Noddies on Bird Key (AsheandRethel, 191.5 MS.)andthesameis almost surely true ofthehurricane of 4 July 1916.The1919 stormstrippedRird Keyofvegetation and also killed "many" terns (Ashe, 1919 This storm greatlyreducedthenumberof suitable nest sitps and no recordofrecovery ofthebaycedar


1964ROBERTSON: DRY TORTUGAS TERKS 61growth exists. Unlessthe1919 storm killed mostoftheadults, how ever,thedecrease from 35,000in1919 to 1600 in 1922canbeex plained onlybythefailure ofmanyadults toreturntotheDryTor tugas.Therecords indicatenodecline in numbersovertheyearsofthe enforced movement ofthecolony from Bird Key to Bush Key. The more recent record shows low points inthelate1930'sandthe late 1940's.Thefirst wasattributabletoratpredation.Thecausesoftheseconddownturnaremore obscure,butrecords sug gestthatrats may againhavebeena factor. Noneofthereportsfor1936 mentionstheprescnce of rats,butRussell (1938 whospent the summers of 1936and1937atthcDryTortugas, statesthatrats became common aroundthefort inthefallof1936andthatsomeapparently swamthechanneltoBush Key. Bythesummerof1937rats weresonumerous on Bush Keythatthey couldbescen commonlybydayand"by thousands"atnight. Russell believedtheBrown Noddy nesting season of 1937 a nearly complete failure, with more than 90percentoftheeggsandyoung lost to rats.Otherau thors give substantiallythesame account. I have seen nothing to indicatethatthe rats killedadultBrown Noddies in any numbers, yet only some 400 adultswcrcin the colonythefollowing year.Asthe population figures for these two years are open to little question, either theadultpopulation suffered extra-Tortuganmortalityofa catastrophicnatureormost ofthepopulation eitherbredelsewhere or notatall. Practically no informationisas yet available on colony fidelity in this speciesandthefactorsthatmaymodify it. Whatever its mortality intheextraTortugan phasesofits annual cycle,themortality of Brown Noddiesattheternery is low, certainly much lower than intheSooty Tern. On 11-15 Julyand8-9 Septem ber 1962 JamesB.Meade (in July),mywife,andI countedandburiedall ofthedeadternsandunhatchedeggsthatwecould find on Bush Key and countedandexamined allthenestsofBrown Noddies.Thetotal observed mortalityofBrown Noddies was 2 downy chicksand11larger juveniles, a remarkably low 1.23 percent. Although someofthe young still unfledged in early Septembermaynothavema tured, this suggeststhatunderfavorableconditions-nopredatorsandno summerstorms-BrownNoddy populations can closely approach their maximum possible rateofincrease fortheegg to fledging stage. The Tortugas colony hasbeenlargely free of disturbancebyrats or severe summer storms sincetheearly 1950's,andtheincreaseofBrown Noddies inthepastdecademayapproximatethatpossible for a colonyofits size performing as a discrete reproductive unit.AsintheSooty Tern,thequestionoffactors limitingthepopu-


62BULLETINFLORIDASTATE MUSEUMVol. 8lation of Brown Noddies is debatable.In1936whenthecolony last reached the level ofabout4000adults, unusual numberswerefound nesting ontheground. Several authors (DoeandRussell, 1936; Allen, 1936; Dickinson, 1941) suggestedthatthis behavior re sulted fromthescarcity of nest sites in bushes,butBeard (1938:lO)disagreed stating: "There are more available nesting locations forbothspeciesofbirds[onBush Key]thanwereever present on Bird Key."In1955 Moore (1955 ::\IS.) reportedthehighest total of Noddies since 1937andcommented: ...itwas amply evident afieldthatBush Key has suitableSurianathicket-edge forthenesting of twice thispresentpopulation."TABLE5.BROWN NESTSITES THE \VESTHAl.F OFBUSH .KEY OUTSIDETHEMATNBAY CEDAR THICKETS KestSite Dead Bushes (mostly Suriana) Sea Rocket(Cakile lanceolata) BayCedar (Suriana martima)Bare Ground Spurl(e (Euphorbiabuxifolia) Prickly PearCactus (Gpuntia sp.) SeaLavender (TouTrlefortia gnaphalode.)Sea Oats (Uniow runicnluta) Sea Purslane porlulacasfrum)Numberof 1962196:368761.52720 1.3 IU10IU 25 7 5Total 1:341'36 Close studyofrecords ofthevegetation of Bird Key suggest that, althoughitwas amuchsmaller island,itmay wellhavehadmorebrushyedgeattimesthanBush Key does now, particularly if thesurmisethathurricanes fragmented formerly solid thicketsiscor rect.Asof1962,thepoint ofdoubledpopulationtowhich l\'Ioore referredhadbeenreached. Brown Noddies inrecentyears have used awiderangeofnestsites in additiontothetypicalbaycedar fringes.Table5 shows nest sites usedbyNoddies in 1962and1963 intheareabetweentheshore ofthewesternhalfof Bush Keyandtheouteredgesofthelargebaycedarthicketinthecenteroftheisland where mostoftheNoddies nest (see figure3).Ne'sts in isolated livingbaycedarbushesandsome of those indeadbusheswerebuiltin a typicalmannerandwithin the usual rauge of heights above


WIl4ROBERTSOl\: DRY TORTUGAS TERl\S6'3 ground, roughly 3to10 feet. Most otherNoddynests in this scction were no morethan18 inches fromthcground,rudimentaryin form, and intimatelysurroundedbynesting Sooties.Parent Noddics atnests of this sorttcndedto participate in the panicsthataffected ncarby oftheSootyTemcolony,andthcyoung seemed to bcginleavingthencst carlierthantheyordinarilydonests inbaycedar. oftenasdownychicks.Inbothyears alargeproportion oftheBrown Noddy nests placed at low sites withintheSootyTemcolonyprobablyfailed. Observed pre-fledging mortalityofNoddies was strongly concentrated in this partofthecolony, particularlyneargroundnestsandthoseplacedonlow herbaceous plants suchassea rocket, spurge, and cactus.InJulyofbothyears evidence of nest success inthefonnof adults at tending young birds was conspicuouslyabsentfromthevicinity of many such nests in addition to thosewhereunhatchedeggsordeadyoungwerefound. recordedatthewestern outlying nests in 1963(3eggsand16 chicks)cannotberelated to total mortality because Ididnot cxamine alloftheBrownNoddynests inthecen tral thicket.In1962, however, 77percentof the observed mortality(10of 13 chicks) was associatcd with nests outsidc the main thicket in the wcstern half of Bush Key, which comprised only 13percent(134 of 1065) of the total nests of Brown Noddies.WhcntheBrownNoddypopulation on Bush Kcyishigh, use by Noddiesofnest sitesnot typical for this colony apparently in creases,andsuch nestsarcmuchmore likely to fail thanarethosc placed in livebaycedarbushcs. Shortage of securc nest sites in the immediate vicinityoftheestablished colony thus maytendtosetan upper limit of population size. Ashmole(1936h: 458-4.59) arguesthatcompetition for nest sites seldom regulates population sizc in tropical sea birds because individualsthatfailtogetnestingspacein one colony can usually emigrate to lesscrowdedcoloniesorfoundnewcolonies. Acknowledgingthatsucccssful emigrationmusthave oc currcdmanytimes inthehistory of every wide-spread colonial spe cies,itisclearthatattraction totheknown breeding placc is a po tent countering factor in some sea birds. Potential emigrants seem far more likely to expend their reproductivc cffortatless secure sitesinorneartheexisting colony.Atleast four timesthepresumed overHow populationofBrown NoddiesattheTortugas has used atypi cal sites ontheislandthe colciny was occupyingratherthanmoving into vacantbaycedarthickets onotherislandsnearby-onBird Kcy after the hurricane of UllO andintheearly 1930's; on Bush Key at times of peak numbers in 19.36 andat present.


64 BeLLEn"! FLORIO1\. STATE \fCSEUM Vol.8Indeed,Ashmole's (1962) account oftheBlack Noddies of Boat swain Bird Ishmd, Ascension, suggeststheybehavesimilarlywhenfacedbya shortageofpreferrednest sites, Crowdingtherereportedlyledthemtouse unsuitable ledgesratherthancolonizenewcliffs, Strong attractiontothetraditionalbreedingplace seems often to inhibittheemigrationofsurplus birdsandto determine a sequence of eventsthateffectively regulates colony size,Exceptfor Audubon's fanciful account,theeffects of a concen trationofSootyTernsuponthebehavior,andperhapsthenumbers, of Brown Noddies nesting inthesame area has scarcelybeenconsid ered. Nest sitesofmost oftheNoddies on Bush Keyareringedbynesting Sooties.Disturbancebythe Sootiesmayfigureatleast in directlyinthepoorsuccess of low nests of Noddies withintheSootyTerncolony.Atothertimes presenceofthe Sooties seems tohavefavoredtheNoddies. Russell (1938MS.) notedthatalmosttheonly Noddies whose young survivedtheratplagueof 1937werethose nesting in bushessurroundedbydense concentrations of nesting Sooties. ContactsbetweenNoddiesandSootiesareinfrequentintheTortugas colony.Ofa similar aggregation on PelsartIsland, Western Australia,Warham(1956:89)stated: "...thereseemedtobeno frictionbetweenthetwo species." Nevertheless,thebehavior of Brown NoddiesattheDryTortugasdiffers considerably intheabsence oftheSooties. \Vhere Sootiesarepresentinnumbers, Brown Noddies enteringandleavingtheterncrytendtoHyfairly highandrelatively few are taken in mist netssetontheopen beach es.InSeptember1962 with most of the Sooties gone, Brown Nod dies swooped from perchesonthebushtopsandlefttheislandinlow,rapidHight. This difference in theirbehaviorisreHected inthemist-net catchesof9-10 SeptemberwhenBrown Noddiesweretakenatan averagerate(8pernethour)thatfar exceededanyprevious results for netssetintheopen.ROSEATE TER:-J AudubonreportedRoseate Terns,Sterna dougaUiiMontagu,nesting intheFlorida Keys (Howell, 1932: 264),butheandotherearly ob serversapparentlysawnoneattheDryTortugas.ThefirstwerereportedbyBartsch (1919)wholocated abreedingcolonyofabout100 pairsonBush KeyorLongKeyinJuly 1917. Most reportsofvis itors tothearea since 1917 have included some mention of Roseate Terns.Threepartialsummariesoftherecordshavebeenpublished (Sprunt,1948a,1949, 1951),thelast carryingthelocal historyofthe


1964ROBERTSON:ORYTORTUGAS TERNS 6.5 species throughthebreeding season of 1949. Table 6 showstherecord of breeding occurrence oftheRoseateTernatDryTortugas, 1917-1963.Thereports for anumberoftheyears in this span de serve comment.1917.In initial remarks on his observations of the summcr of 1917, Bartsch (1918: 171) wrote: "...probably 200 common terns formed a rookery ontherough coral shore oftheeastern end of the island [Bush Key]. Their young birds of various agcs couldbeseen at all times."Thclater, morcdctailedreport ofthe1917 breeding scason (Bartsch, 1919) does not mention thc CommonTcrnin text,butdis cusses(p.489) a colony of "ahout 100 pairs"ofRoseate Terns brced ing "on the rough coralandshellstrewn northcastern end of Long Key."Thelegends to Plates 27-32 inthe1919 publication, a seriesofphotographs ofthecolony site andofyoung birds,statethatthepictures show Common Terns,bnta footnotc (p.5(0)corrccts this to read Roseate Tern.Thedowny chick shown in oneofthephoto graphs (Bartsch, 1919: Plate 28a)isclearly a Roseate.Itappears certainthatthese reports refertoa single colony. A RoseateTernspecimen intheU.S.National Ivluseum Hartsch collectedonBush Key 17 1919 doubtless servcd to establish the correct identity. In his writings on the Tortllgas, Bartsch appears on somc occa sions tohavefollowedthenomcnclature ofoldercharts on which application of the names Long Kcy and Hush Key is revcrsed from present usage.Forthis reasonitisimpossibletodetermine conclu sivelywhethertheRoseateTerncolonies of 1917,1921, 1922, and 1932 were located ontheeastern sandspit of Bush Keyoron one ofthericksofcoral fragmentsthatcomprisetheislandnowknown as Long Key. This uncertaintyisoflittle importanee, becausethesites are similarandnot more than a fewhundredyards apart. 1921. Nestswith1, 2,and3 eggs were seen (Bartsch, notes). 1922-1925. According to Bartsch(MS.notes),theternswereassem bledatthecolony site 14 May 1922,buthadnotbeguntonest.For7June1924henoted, "some seen,butcolonynotbreeding."Healso observed numbersofRoseate Terns feedinginthe Tortugas area on 5 September 1923and12-18 August 1925. 1935.Thetotal shown in table 6isa synthesis of the estimatesbymembers oftheAudubon party. Some oftheobserversthoughtthatnomorethan 100 pairsofRoseates were inthecolony (Russell, 1938


66BeLLETI\! FLORIDA STATE \ll:SEL'\1 Vol. 8 Mason (1936andin litt.)reports young 3 or4 daysoldandmany eggs not yet hatched. 1936.Thepublished account (DoeandRussell, 1936:7)states: "Therewereprobablyaboutthe samennmberof roseate terns as last year..., butDoe(MS.notes) recordedthattheRoseateTerncolonynumbered"about400 nests." Nesting wasapparentlyjust beginning, for Mason(MS.notes)wrote"96 nests located, all with 1 or 2 eggs. Noneyethatched." 1937. Reports ofthetrip(YoungandDickinson, 1937; Longstreet, 1937) givenoestimateofthenumberofRoseate Terns seen. This omission misledSprunt(1949, 1951)tostate that nonebredattheDryTortugas in 1937. YoungandDickinson, however, mention (pp. 3-4)thatthey visited a keywhereRoseate Ternswerenestingandthey include (p.6) a photographwiththelegend, "RoseateTernbandedbyC.R. Mason on Sand Key." Banding schedules showthat Mason bandedthreeadultRoseates on Sand (Hospital) Key on25June1937andheadvises (inlitt.)thatasherecallsitthecolony nesting there was slightly smaller thantheones observed on BushKeyin 1935 and1936. 1938. Mason (1938:1)notedthe nesting colony included"betterthan300 birds." HisMS.notes recordthat157 nestswitheggs were counted,andthatthelocationoftheternery,notmentioned inthepublished report, wastheeastern sandspit of Bush Key.1940.Nesting was jnst beginning.Theobservers found 5 nests each containing a single egg. In addition,theFortJefferson Custodian told Robinson (1940: 3)that"quitea number" of Roseate Terns were believed tobenesting on Bird Key,thenre-emerging as a sandbar. 1941.Thebrief report ofthetwo parties of observers who visitedtheDryTortugas inJune1941 (Rea, Kyle,andStimson, 1941) mentions onlythatRoseate TernswerenestingonBush Key. LouisA.Stim son (inlitt.)writesmethatthe firstgroupsawbutone Roseate Tern, in flight overFortJefferson. Roger T. l'eterson, who accompanied the second group, writesme(inlitt.)thathesaw no Roseates,butthattheCustodianofthefort told himtherewas a nesting colonyonthe east end of Bush Key. Individuals whohadseenthecolony in both years toldR.R.Budlong (1942MS.)thatitwasaboutthesame size in 1941andin 1942.Aswiththe1937 report, lack of a definite popllla-


r1\1641l0BERTSO!'i: DRYTORTUGAS TERl\S 67tion figure has resulted inthestatementthatthespecies was absent fromDryTortugas in 1941 (Sprunt, 1949). 1942.Thecolony was located "on the reef between Bush andLongKeys."Thetotal shown seemstohavebeenonly aroughestimate, the author commentingthathewas able to visittheareabutonce, on 2 July,andfound "numerous eggsandyoung birds." 1943. No population figure is given inthereport which merely states "The RoseateTerncolony seemstocontainaboutthesamenumberofbirds as last year." 1947.Sprunt(1948a:29) counted 67 nests onLongKey, 54 on Bush Key,and21 on Hospital Key.Inaddition,about12 young (not in cluded inthetotal) survived from an earlier nesting onLongKey disruptedbyhigh tides. 1948. Sprunt(1948c:14) counted a totalof216 nestswithhatching "about50%complete."Headds: "It isvirtually certainthata few were missed, despite care. A totalof225isvery likely." 1949. Dilley (1950: 68) located 44 active nests on Bush Keyand7onLong Key. An additional 17 nests on Hospital Key (not included in total) are saidtohavebeenabandoncd.1950.Nest locations inthethreeyears were: 19.50, 55 on BushKeyand7 on Hospital Key; 1951, 35 on Hospital Keyand33onLongKey;1952, 136 on Long Keyand58 on Bush Key.Fromobservations later in the summer of 1950, JohnR.DeWeese(inlitt.to Dilley) reported storm tides Hooded alltheRoseateTernnests sothatno young wcre rearedthatyear.1953.An earlier group of 9 nests on Middle Key,theonly Roseate Tern nests present inthearea on 26 lvlay (Moore, 19054 MS.),was de stroyedbyhigh tidesduringa storm 28 l'day. Thelater nesting included79nests on Hospital Key, 26on l\'liddle Key,and IS on Bush Key. 1956. ,lI,Then counted, many ofthenests (88 on LongKeyand14onBush Key)hadincomplete clutchesandanumberoffresh nest serapes without eggs were present.Twoweeks earlier \Iargaret H. Hundleyhadestimated ISO Roseate Terns intheTortugas area (Stevenson, 19.'56: 327).


68 BULLETlK FLORIDA STATE .\IUSEU".{ Vol. 81957-1958.TheI'ecord for these yearsisalmost certainly incom plete. No 1957 observations later than midIay areavailable, and in 1958 no partieular effort was made to locatethecolony. Possibly coloniesofmore normal size developed each ycar. 1959. A hastycountlocated approximately 225 nests in adensematof Sesucium onthehighestridgeof Hospital Key. Hatching was about half completed withthelargest chicks about one week old.Thepartybanded80 chicks.1960.ObserverswhovisitcdtheDry Tortugas in early '.lay saw somc Roseate Terns around the eastendofBush Key,butthecolony apparentlyhadnotbcgun nesting(I.Joel Abramson,inlitt.).Band crs workingthere27-31 May frequently saw a few Roseates fishing inthebightbetwecnBush KeyandLong Key,butfound no nests. Severe squalls prevented visits to anyofthe outlying keysbyeithcr party in May.On11July members of a sc<-'Ond groupofbanders landed on all the keys. No Roseates were nesting atthatdate,buta densely masscd assemblage of ternsandgulls on Middle Key in cludedabout100 individuals of some speciesofwhiteSterna,manyof which were birds of the year.Thebehaviorandunsteady flight of these youngsters indicatedthattheyhadbeenrearedatDryTor tngas, althoughnotnecessarily on 1liddle Key. Opportunity to stndytheadults was brief,andtheobservers,awareoftheuncertainty sur rounding reports of southern nestings oftheCommonTernconcludedthatthe birdswereRoseates.Thesingle juvenilenettedandbandedon Middle Key was so reported. 1961. Oliver L. Austin, Jr.,andWilliam G.Atwaterbanded20 well grown juveniles ontheeast spit of Bush Key 16 July. 1962.TheRoseate Terns first located on several elongate heapsofrough coralfragmentsnearthesouth endofLongKey. A member ofthebandingparty, TIleodoreR.Greer, devoted several daystophotographing (figure9)and observingthecolony from a blind.On27 May he counted 118 nests, 34 containing single eggsand84withtwo-egg clutches.On13Junea field excursion groupofthe13th International Ornithological Congress (Robertson, 1962) foundthecolony site desertedandbrokenegg shells remaining inthenest de pressions. Slight vascularization oftheinner shell membranes indi catedthatpredationhadoccurred early in incubation,andthewaytheshellswerebroken suggestedtheworkofan avian predator.The


r 1964 ROBERTSON: DRYTORTliCASTERI\S 69most likely suspectswerethesome20cattleEgrets,Bubulcus ibis,then frequenting BushandGardenKeys.CattleEgretsattheDryTortugashaveformedsomeunusualfeeding habits.In \llay andJune 1962andMay1963theywerefrequentlyseen to stalkandkill injuredorexhaustedspringmigrantpasserines (mainly Parulids)andtofeeduponsmallbirdsalreadydead. 9. Roseate Tern incubating, southendofLongKey,26 May 1962.The colony lX.'cupiedseveral dune-like elevations of eOTaI nlbhle.The bird pichIred had a darker hill than mostof tilt' adults in the <:oluny, but note that lower manclihle islighter (reddish) at the Ix",'. (Photograph by Theoclore R. Crcer.)Some oftheRoseatesappeartohaverenested onHospitalKey in July.C.R.Masonandothers foundabout ,'50 uests,allwithone egg, thereon16 July.ParkRanger CarlS.Christensen (in litt.) the 10 dayslaterandreportedthathatchinghadbe gun.Atthattime none ofthenestscontainedmorethanoneegg, apparentlythenormal clutch for second nestings of RoseatesattheDry Tortugas. 1963.On17and18 \'lay membersofthebandingpartycounted73nests, each containingoneortwo eggs, andbanded 32 adultRoseates on Hospital Key.On7 July large young from the !vlay nestingwerecongregated onthebeaches and an estimatecl 1.'50 additional adults had arrived andbegunnesting.


70BULLETINFLORIDASTATE MUSEU.\\ Vol. 8 DrscussIOK Commentsonthenesting oftheRoseateTernattheDryTortugas have stressedtheisolationofthecolony and itserraticfluctuations in size from yeartoyear.Spnmt(1951a:14), for example, writes:"Themarked fluctuationofthis tern at Torhlgas seemswithoutex planation as, indeed, does the very fact of its being there!" Analysis of the longerrecordofpopulationnowavailable suggeststhatmuchofthesupposed fluehration results from incomplete data.TheTorhrgan Roseate colony has shifted frequentlybetweentheBush Key-Long Key area oppositeFortJeffersonandtheareaof HospitalandMiddle Keys, several miles northeast oftheFort.Insome yearstheentirebreedingpopulation hasbeenconcentratedon one key; inotheryearstwoorthreeseparate colonies existed (table 6). Observersarcnotlikely tohaveoverlooked Roseate Terns nesting on Bush KeyorLong Key,hutHospitalandMiddle Keysareless easily accessible and neitherwas visitedbythe observerswhoreportedfor 1935,1936,1939, 1940, H141, and 1956,norprobablyfor 1938, 1942. 1943, and 1945. TAHLt: 6. BlU:iliDlI'l;RECORDSOF ROSEATE TERNSAT THE DHY TOHTUGA.S LOt'ation ofNumberofDateTerneryAdults Source 19-31 July"Long Key" 200 Bartsch (1919) 1917 9Junc"BushKey" 200Bartsch(111S.notes)192114 "Bush Key" 200 Bartsch(illS.notes) 1922 Angust, "inthe quarters the usual Bartsch (19:J2) 1932previously...colony"occupied"19-20JuneRush Key 400 .\Iasou (19:36)193,5Longstreet (19:J6a, 1936b) 17-19JuneBush Key 400Doe&Russell (19:16)19:16 23-25JuneHospital Key 25Q-:300:\"[ason (ms.notes) 19:17 20-25JuneBushKey 314 :\'[ason (ms. notes) 1938 21June Bttsh Key 214 Taylor (19:19 TIls.) 1009 about3June Bmh Key 20 Rohinson(I940) 1940 (continued)


r 1964 ROBERTSON:DRY TORTlJGASTERl'\S TABLE6 (continued) 71 Date early June, 1941 2July 1942 1943 18-20June1945 2:3-2.5 June194715-11lJune1941l 13 June 1949 1-5 June19.'>0al May-4June195127 1952 July 1953 27 May 1954 a June195526 May 1956 19 1957 June1955 1:3Inne 1959 7-8 May 1960 7-17July 1961 25-28 May 1962 MayandJuly 19fi3 Location ofTerneryBushKey Long Key Bush KeyBush KeyLong KeyBtL.<;h KeyHospital KeyLongKey Bush Key LongKey(Hospital Key) Bush Key Hospital KeyHuspital Key LongKey I,aug Key Hnsh Key Hospital Key Kl'y Bush Key Hospital Key Bush KeyLongKeyBush KeyIf ospital:Key BushKe)Hospital Key (Bush Key)Middle Key? Bush KeyLougKey Hospital KeyNumber ofAdults ? c. 150c.150170284 450 102 124 1:36 allil2<10:370 204:30J5 450 100-12.5 120 c. 300SourceSec comments Budlong (1942 tns.) Budlong (194a tns.) Sprunt (1946a)Spnmt(1948a)Sprunt(1948e) Dilley (1950) Moor,' &Dilley (1953) &Dilley(195:3)Moore&Dilley (19.5a) DeWeese(195.3IllS.) (1954 ms.) (19.55 ms.) Hobertson (1956 ms.) D,Weese (1957IllS.) \Vad(in litt.)O.L. Amtin, Jr.etal. Abramson(in litt.) Robertson (1961) Greer(in Utt.)C. H. Mason,W.B.Robertson.etal.


72BULLETINFLORIDA STATE Vol.8In addition to low counts rcsulting from incomplete coverage, some countsmadewhilethecolony was forming surely underestimatetheactual breeding population. Roseate TernsattheDryTortugas are often wellbehindtheSooties intheirbreeding schedule, ,md nesting dateshavevaried considerably from ycar to year.Insome years the full complement of breeding adults assemblesbymid-May and laying beginsduringthefirstweekof June.Inotheryearsmanyfirst nests stillhaveincomplete clutchesthelast week of June.Therelationbetwccnthcdateandthesize ofthepopulation re corded is well illustratedbytherecords for 1939whenon 9JuneRob inson (1939:9)estimatcdabout80 Roseate Terns ontheeastendof Bush Keyandfound 13 nests with eggs.On21Juneinthesame L'Olony Taylor (1939MS.)counted 107 nestswitheggs. }''1oore, in 1952 (MooreandDilley, 1953: 78), counted 93 nests on Bush and Long Keys24 IVlay and194 ncststhereon27 May.InlateMay 1953 Moore (1954 ;\IS.) was abletolocate only 9 RoseateTernnestsattheDryTortugas,butbyJuly (DeWeese, 1953MS.)120 nestswerepresentonthreekeys. Checksofthis sort arenotavailable for other years,buttheearlicr counts clearlyhavetendedtobelower.Thelow RoseateTempopulations recorded in 1940, 1950,1951,and1957 all dcrive from countsmadein 'vlay orthefirst week of June.In1946 a search of all of theTortugankeys 16-20Junerevealed no Roseate Terns (Spront, 194&: 1,7).TheCustodian ofFortJeffer son, however,hadreporteda few Roseate Terns intheareaduring 'vlay (Gibbs, 1946MS.).Perhaps none nestedattheDryTortugas in 1946,butitmay alsobethatformationofthecolony was unusu allyretardedthatseason,orthatan early nesting was destroyedbyspring tides or predators. Excepting only 1946, all complete surveysoftheknown nesting keysmadeafter mid-June have located breed ing aggregations of approxinlatcly 150 to 450adultRoscateTems.ThustheTorhlgan Roseate colony appears neither particularly erratic in its breeding nor to undergo numerical fluctuations of unusual mag nihIde.Thedifficultiesofseasonandlocation mentioncd abovemaypossibly account forthefailure of 19thcenturyornithologiststofind Roseate Ternsatthe TorhIgas.Whileitscems likelythatAudubon and others would have investigated allthckeys,itisnotcertainthatany of them did. Scott,andprobablyalso Audubon, Bryant, and 'vlaynard's assistants wereatDryTortugas too early inthespring to find Roseate Terns, assumingthatthc colony existedandfollowed its present seasonal schedule.Inaddition Bryant's tcstimony is renderedequivocalbythepossiblitythatheconfused RoseateandCom-


1964ROBERTSON: DRY TORTUGAS TERNS73mon Terns. Hisstatementthathesaw no Roseate Terns follows his comment(1859a:21):"1foundtheLittleTernandWilson'sTernbreedingatdifferent localities amongthekeysandalongtheshore ofthemainland."AsfortheTortngancolony'sreputedisolation,whileitisindeedremote fromtheremainderoftheRoseate Tern's breedingrangein the United States,itisattheperipheryofthespecies' extensive An tillean-Caribbean range.Intermsofthespecies'worldrange,itisrathertheNorth Atlantic coloniesthatareunusual. MostotherRoseate Terns nestwithinabout30 degreesoftheequator(Voous,1960:151,map196).Thecoloniesthatbrecdfrom VirginiatoNova Scotiaandfrom BritannytoJutlandmayberelict asFisherandLock ley (1954: 87) suggest,butlocal extirpationcannotbedisregarded as a possible causeofpresentbreedingrangediscontinuities.InFlor ida, for example,noRoseateTernnesting colonieshavebeenknown sincethemid-19th century, excepttheoneattheDryTortugas. Many summer sight re<:ords of Roseate Terns inthecentralFloridaKeysandtherecentreportof a small colony "ontheVacaKeys" in which 3 chickswerebandedandseveral nestswithsingle eggs seen on11June1962 (Pace, 1962) suggestthepossible recolonizing of formerbreedingrange. This colony ofabout30 adults was again active inthesummer of 1963 (ChristineA.Bonney, pcrsonal communication).LEAST TFlDl Theeggcollection catalogueoftheU.S.National Museum lists eggs oftheLeast Tern,SteT1laalbifronsPallas, takenatDryTortugas in 1859and1861 (Robbins,inlitt.).Littleelse isknownofits nesting there prior to 1900. Scott (1890: 306) was informedthatitoccurred commonly,butsawnoneduringhis visit in March-April 1890. By 1900 theTortuganpopulatiou ofLeastTernshaddeclined greatly. Thompson (190:3: 83-84) found a colony of 30 pairsonLongKey in 1902andreportedthatformerly populous colonies onLoggerhead Key hadbeen dispersedbyeggers. Watson (1907:31.5-316)reported.50pairsattcmptedto nest 6rst onLoggerheadKeyandthcnon Hos pital Kcy,butpredationanddisturbancepreventedbothcolonies from producing youngandheconsidered the species "ncaring cx tinction" atthe Dry Tortugas.WiththeestablishmentoftheCarnegieTortugasLaboratory,A.G.Mayer,the6rstdircctor, undertooktostopthegathering of tern eggs and to controltheratsthatinfested several ofthekeys. In 1908hewrotethatthesuccess of these efforts couldbeseenin


74 BCLLETf:"i FLORIDASTATE MUSEL'I>I Vol. 8,thethriving coloniesofLeastTernsonbothends ofLoggerheadKeyandonBush Key,thelattersaid tonumberabout1000birds(Dutcher,1908b; \fayer 1908),TherevivalofthecoloniesonLoggerheadKey was short-lived.In1917Bartsch(1919: 487)observedthat "theper sistent efforts of eggers"hadfinally driventhcbirdsawayfromtheisland. Small nestingswerereportedonLoggerheadKeyin1932(Bartsch,1932: 287)andin19:35-1936(Russell,1938MS.),buttherehasbeennosubsequentindicationthattheLeastTernmightrecol onize its originalTortuganbreedingground. Its failure todoso at LoggerheadmaybeexplainedinpartbythefactthatdogsandcatskeptbytheIigbtbouse personnel oftenhavebadfreerangeoftheisland.Incontrast tothecheckeredhistoryoftheLoggerheadcolonies,LeastTerns nestingintheBush Key-Long Key areamaintaineda fairlyconstantpopulationformanyyears. Bartsch(1916, 1917, 1919)reporteda colony of200onBusb Keyin1915,.500therein1916,and500onLongKey in1917.On9June1921thercwere400 500birds onLongKeyandBartsch noted on3June1924 ()ofS. notes)that"several colonies"werebreedingintheTortugas. "Varden Charles1.Park ()ofS. notes) estimated500LeastTerns nestingonLongandBush KeysinJuly1929,and700nesting on Bush Keyin1930.Bartsch(1932: 281)found"theusualbreedingcolonies" active in August1932.ThefirstFloridaAudubonSocietyTortugastrip in1935reported20()Least Terns nestingonthecastspit of Bush Key C..Iason 1936: 18),which suggests the population w>lSsomewh>lt reducedfromthatpresenta few years earlier.Thefollowing yearDocandRussell(1936:7)estimated this colonytonumberonly100birds.In19:37therewere noLeastTernsonBush Key,butabout25 pairs nested intheRoseateTerncolony on Hospital Key (Russell,1938 )01S.) Inthenext four yearsLeastTernswerereportedontheeastspitof Bush KeyandadjacentpartsofLongKey as follows:In1938Mason(1938: 4)found11nests.In1939Robinson(1939:9)reported25adults,andthefollowingyear(1940: 3)"NotmorethanadozenLeastTernsinthecolony."In1941Stimson (inlitt.)reported "a fewLeastTernsnesting."In1946Sprunt(1951: 15)sawemptynest scrapes onbothBushandHospital Keys,butdoesnotmentionhowmanybirdswerepresent.In1947hereported(1948a:30)"lessthana dozen" adultsonBushandseveral scrapeswithouteggson Middle orHospital Key.In1948hereported(1948c:13-14) 12birdsonBush Key,wherethefollowingyearDilley(1950:68) found a singlenestwith eggs.In


1964 ROBERTSON, DRY TORTI.:CAS TERNS1951MooreandDilley (1953: 78) saw two birdsbutfound no nest. Thereafter no LeastswerereportedattheTortugasduringthe brccd ing scason until July 1963, whenaboutfive adultswereseen for sev eral daysaroundGardenandLongKeys.AsmentionedunderRoseate Tern, HospitalKeywasnotvisitedinsome years,andsomeofthevisitsaftcr1935 mayhavebeen too early intheseason to rccordLeastTerns nesting.Therecordis doubtless incomplctc,butccrtainlythespecies no longer breeds reg ularlyattheDryTortugas. No clear explanation oftherapid disappearanceoftheLeastasa breeding speciesattheDryTortugas canbeadvanced. Be tween 1932 and1937 a stableandlong-established population of ap proximately 500 breeding adults on BushKey-LongKey decreasedtoa few birds, withnoevidencethatthecolony suffered disturbanceofany sort.Thedecline of theLeastTemsduringtheyearswhenthe colony of Sooty Terns was bccoming establishedandincreasing on Hush Key suggests the poSSibility of some relationshipbetweenthe two events. Also of possible significance isthegreatincrease of LeastTerncolonies alongtheadjacent coastsofsOllthern Florida since the early 19.'30's. Dredgingalong theInlandWaterwayandfor coastal real-estate developments hascreatedinnumerablc small, sheltered isletsandbarswhich provide ideal nesting sites, perhaps preferable to more exposed islands liketheTortugas. COMMOl\"TERl\" UntilHallman(1961) reported two nests found1Ilthemidst ofthecolony of Least Terns" on a spoil island ill St. Joscph's Bay, Gulf County, inJune1961, observationsatDryTortugasprovidedtheonly generally accepted evidenceofthe nesting of the CommonTemin Florida (Howell, 1932: 263).Infact, theTortugancolony has bcen consideredtheonlyoneintheentire Gulfof [v[exico region (Lowery and Newman, 1954: 530), althoughtheA.O.U. Check-List (1957: 235) mentions breeding coloniesontheeoastofTexas,andStcwart(1962:485)recentlyreportcda possible nestingOntheGulf coast of Mis sissippi.Thefew records of breeding at the Tortugas are not altogether satisfactory.Theyaredocumentedneitherbyspecimensnorphoto graphs,anda strong possibility of confusionwiththe RoseateTemexists.Ashasbeennoted,thefirstreportof nesting CommonTemsinthearea (Bartsch, 1918)provedtobebasedon a misidentification


76 BULLETl1\: FLORIDA STATE MUSEUM Vol. 8of Roseate Ternsandwas later corrected (Bartsch, 1919).Theother reports are summarized below. Bartsch (),1S. notes) reported a colony of75pairs of Common Terns on Bush Key 3June1924andnotedthatthenests contained "1-4 eggsornewlyhatched young." On13 August 1925hereported "quite a colony present"atBushandLongKeys,andin 1932 (Bartsch, 1932: 281) "the usualbreedingcolonies" werc said tobeactive. I havebeenunabletolearn anything moreaboutthese observations.Itistobenoted, however,thatBartsch alsoreportedRoseate TernsatDryTortugasonaboutthesame datesin1924and1925.Theremaining recordsdatefromthebreeding seasons of 1935, 1936, and 1937.Ineachcase Common Ternswerereported nesting with a larger groupofRoseate Terns onthceastspit of Bush Key in 1935and1936andon Hospital Key in 1937. Available information suggests some uncertainty inthemindsoftheobservers concerning identification ofthebirdsasCommonTernsandthenumberof pre sumed Common Terns present.Thenumberin 1935 was reported var iously as 50 birds (Mason, 1936: 18; Longstreet, 193&: 33) "about 75 pair" (Lougstreet,1936b:99),and "100 pair" (Doe, )'1S. notes). Long street(1936a:42) commented: "Mr. Mason andI believethatwefoundthecommon tern breeding on Bush Key. However,wcdidnotcollect any birdsoreggs."ThereportofthcFlorida Audubon Society's Tortugastripof1936 (DoeandRussell, 1936:7)states: ...thecommon terns showed a marked decrease, only a few pairs being noted." Mason advised me(inlitt.)thatonly four birdswereseen on Bush Key in 1936 and tIlat no nestswerelocated. Russell (1938MS.),however, wrote elsewhere: ...in 1935and1936 I estimatcdthesame colony tocontainabout200 birds."Thelatter statement could pertain to observationsmadelaterinthesummer. Published accountsofthe1937trip(Longstreet, 1937; Young and Dickinson, 1937) donotmentiontileCommon Tern,but I-dason (inlitt.)saw a few adultsthatbebelieved were Common Terns amongtheRoseates on Hospital Key. Russell(1938MS.)statesthatCommon Terns nestedonHospital Kcy in 1937 without indicatinghowlargethecolony was. Since 1937,theonlyreportedoccurrenceattheDry Tortugas duringthebreeding season appearstobetwoseen on lvIiddle Keyby andMrs. JohnR.DeWeese, 29 May 1955 (\foore, 1955MS.).Itseems necessary to conclude fromtheabovethatbreedingoftheCommonTernattbeDryTortugasisnotproved.Thedowny youngoftheCommonandRoseate Terns arc easily distinguishable


r1964ROBERTSON, DRY TORTUGAS TERKS 77byanyone familiar with them. Chicks of the presumed Common TernsatTortugas were seenin1924andprobably also in 1935 (Mason, in litt.),butno recordthattheywerecompared critically with Roseate Tern chicks exists.Inaddition, some of the identificationsofadult Common Terns apparently were based uponthebill color, which oftenisunreliable for separating CommonsandRoseates. Greer(inlitt.) advised methatno morethan15or 20 oftheadults in the colony of Roseates on Long Key in May 1962hadentirelydarkbills, thc others havingatleast the basal third of the bill orange-red. The latterisconsidered tobethe nvariant" conditionbyPeterson(1947:plate 37), wIllie Pough (1951: 288) states: "Its billisblack ex cept for a little redatthebase (occasionally more)."TheCommonTernhasbeenreported to nestatanumberof New World localities south of its regular breeding range. Considerable uncertainty surrounds most of these records, however, because of the similarity between Common and Roseate Terns, and because band ing evidence showssubadultCommon Terns often summer in the tropics.AsVoous (1957: 139) notes: "Its nesting in theWestIndian region has been almostasfrequently stated asithas been reject ed..." Bond (1956:58)gave full credence tononeof the numerous reports of breeding in the Bahamasandelsewhere in the'VestIndies. Similardoubtattaches in some degree to most or all ofthealleged nestings on the South Atlantic and Gulf coasts oftheUnited States, including thoseattheDryTortugas. Voous(1957:137-140; 1963) has recently published unquestion able proof of the CommonTernnestingat Cura"ao, Aruba, and Bonaire inthesouthern Caribbean. Recordsthatheassembled sug gestthatthe species has a long historythereas an irregular breederinsolitary pairsorsmall colonies of fewer than 20 adults.Itisalso saidtobreedinthenearby Venezuelan islands of Los Roques and Las Aves (Phelps and Phelps, 1958:111).AsBond (1958:5)states, this proof of southern nesting compels a re-examination oftheearlier reports.Thesignificance of the proved and reported southern nestings of the CommonTernisnotclear,butmost oftherecords seem to conform to apattern-smallnumbers of birds nesting sporadically, often in associationwithlarger colonies of other terns, especially Roseates and Leasts. Band recoveries showthatmany 1-, 2-, and3year-old Common Terns summer intheCaribbean and elsewhere south of the species' usual breeding range. I suggest as a provisional explanationthatsome individuals in these normallysubadultage groups reach sexual maturity in southern latitudesandareoccasiona-


78 B\JLLETI)l FLORIDASTATE Vol. 8ally stimulated tonestwhenthey become associated with terneries of other species. Voous (1960: 128) has commented: "The few recorded breeding placesinthetropics, very limited in extentandaltogether isolated from one another,mustberegardedasrecent colonizationsbybirds left behind after wintering .. ." Aspresently known, however, southern nestings of the CommonTernseem to fit more closelythehypothesisthattheyare anomalous, impennanent,andperhaps re lated totheageoftheindividuals.Itis doubtfulthatthey have sig nificanceasextensions ofthespecies' breeding range.ROYALANDSANDWICHTERNSAudubon found the Royal Tern,Thalasseus maximus (Boddaert), nesting abundantlyattheDryTortugas in May 1832. John Krider (1879: 80), presumably from observationsmadeinthespring of 1848, writes of it: "Veryabundanton Tortugas Island, Florida,andbreeds ontheKeys of Florida."In :Vlay, probably of 1850, Bryant (1859a: 20-21) visited Nortlleast Key,DryTortugas, whereheobserved this speciesandtheSandwich Tern,Thalasseus sandvicemis (Latham), breeding "in great numbers."TheU.S.National Museum contains eggs of the RoyalTerncollectedatDryTortugasbyGustavusWurdemann in 1858,andeggs ofbothspecies collectedbyDr. D. \V. \Vhite hurstandCaptainD.P.Woodbury in 1859 (H.G.Deignan,in litt.).Scott (1890) doesnotmentiontbeSandwich Tern,buthesawsizableRocksof Royal TernsattheDryTortugas in early April 1890,andwastoldthatmany remained there to brecd. These brief comments span the entire rccord of breedingbythese speciesattheTortugas, exceptthata single Royal Tern egg was found on Middle Key in May 1952withnofurtherevidenceofnesting (DeWeese, 1952 In Sprunt's (1962: 84)reportofmy7 November1961 observationofRoyal Terns ...therc appeared tobeseveral times thisnumbernesting on the soutb end of Long Key," nesting is a typographical error for resting. Royal Terns still visittheTortugas regularly, sometimes in large numbers.FortheSand wich Tern anumberof observations exist from thc neighborhood of Key West, including several in summer,butthree sight records, two oftllCmrccent,aretllConly known occurrcncesattheDryTortugas in thiscenhlry(Sprunt, 1962:84).Northeast Key, mentioned asthesite ofthenesting wlony of Royal and Sandwich Terns,hadwashedawayby1875.Thenarra tive of a surveymadeinthatyear states (Coast Survey, 1878): "North


r 1964 ROBERTSOJ\; DRY TORTUGAS TERNS79Key, NortheastKey,and SouthwestKey,as represented on old maps, have no existence now,notbeingbareevenatlow water."Otherislands nearby, such as EastKey,hadareasapparentlysuitable for the species,anditseems doubtfulthatlossofone key could have caused loss ofthecolony. A more likely explanation isthatthein creasingly persistent egging afterabout1880 (Scott, 1890) eventually extirpatedthetemery. Both Thalasseus species nested commonly at anumberofsouthern Florida localities inthe 1800's, butno breed ing colonyofeither species is knowntoexist inthearea today.BUCKNODDYThe Black Noddy, Anons tennirostris(Temminck) was first recordedinthe continental United StatesatBush Key,DryTortugas, 13 July1960(Robertsonetal.,1961),whenone was collectedanda second individual seen.Duringthesummers of 1961, 1962,and1963thespecies was seen repeatedly on Bush Key.Withoneexceptiontheobservations havebeenof single birds, usuallyperchedwith Brown Noddies inthedeadtreeatthesouth shoreofBush Key from which the 1960 specimen was collected. Todateatleast five different indi vidualshavebeenseen,andthespeciesapparentlyis of morethancasual occurrence there.TheBlack Noddy is slightly smaller and darkerthanthe Brown Noddy, its bill is thinner,andits crownpatchis whiter, more sharply defined,andextends farther backOnthenape. Yetthetwo species arc so similar in general appearance and behaviorthatOnecould easilybeoverlooked in a congregationoftheotherunlesstheob serverwereexpectingorwatching for it. Sutherland (1961) describes howhefirst spotted the Black Noddy in 1960, while making pro longed observations on agroupof Brown Noddies inthe"noddytree" on Bush Key to record their calls. Otherwisethespeciesmighteasily have gone undetected,anditisindeed possiblethata few birds may have frequentedtheTortugastemeryunnoticed for many years.In1961thefirstpartyofbanders saw a BlackNoddy daily inthenoddy tree 26-31 May.Thebirdis clearly recognizable in 16-mm color moviesB.G.Hubbardtook 27 May (figure 10). The second banding group also found one Black Noddy on station inthetree7,la,11,and IS July.Thebirdseen in May hadanindistinct slash of lighterbrownacrosstheleft middle coverts, apparently causedbyworn feathers ithadnotmolted.Thehird seen in July lacked this mark and may havebeena different individual. Repeated attempts


80BULLETINFLORIDA STATE MUSEUMVol.8bybothgroups ofbanderstocapturethebirdfailed.Itshowedtheextreme tameness characteristic of noddiesandtolerated ap proachtowithin a few feet,butwasmuchmore agileonthcwing thantheBrown Noddies and easily avoidedbothmistandhandnets.In1962 four partieswitha combined total ofmorethan50 ob servers searched Bush Key for Black Noddies without success 5-6 May, 11-14 May, 25-28 May,and 1,3 June.Thesecond bandingpartyfound oneattheusual roost on 7and11July,andon 13 Julycaughtitin ahandnet.Itwasbanded(683-12000 on right leg, un numbered red plastic on left), weighed (103 grams), measured (wing arc 218 mm, exposed culmen 42 mm), photographcd (figure 12) and released. Its mouth lining,byPalmerandReilly's (1956) color standarcIs,was approximately "scarlet-orange", strikingly different from the "orangc-ycllow" oftheBrown Noddy's mouth. This individual hasnotbeenrcportcdsince.Thepartysawnomore Black Noddiesthrough15 July, and I could find none on 2 August.Onthe eve ning of 9 September, however,mywifeandIcaughta second Black Noddy in a mistnetonthewestbeachamIbandedit(683-11999).Thefirst 1963 bandingpartysaw one Black NoddynearHospital Key 17 May,butcould find none on Bush Key.On6 Julythcsecondpartyfound oneunbandedbirdperchedamong Brown Noddiesatthenorthcoaling dock onGardenKeyandphotographeditfrom a dis tanceofa few feet.On9 July two Black Noddies, neitherbanded,roosted for several hoursatthesame plac'C. Oneofthese differed from all others seenattheTortugas in havingthebackofthepileum duskyratherthanwhite. Presumablyitwas a younger individuaL Anumberof interspecific squabbles for roosting spacewereobserved, in whichthelargerBrown Noddy was usually dominant. Thus oneormoreofatleastfive individual Black Noddies havebeenpresentattheTortugasterneryduringfour successive summers.Theirknown extreme dates of occurrence, 17 May-9 September, span virtuallytheentirebreedingperiod of terns inthearea. Since 1960 we have devoted considerable timc,perhaps50ormore man-hours,tosearching for a possible BlackNoddynest.SofarnoBlack Noddy hasbeenseenata nest, and no nests, eggs,oryounghavebeenfoundthatappearedtodiffer from those oftheBrown Noddy.TheBlackNoddyof July 1962 was several times observed to leave itsperchinthenoddytreeandflydirectly intoanareaofdense brushnearthewestendofthekey. This behavior was suspiciously likethatoftheoff-dutymemberofanincubatingorbrooding pair,butminutesearchofthearea-severalacres of tightly interwoven


1964 ROBERTSON: DRYTORTUGAS TERNS81FrGunE10. Black Noddy (left) and Brown Noddy, BushKey,27 May 1961. (Photograph enlarged from 16 rom moviebyB.G.Hubbard.) FIGURE n.Black Noddy (upper left) and Brown Noddies, BushKey, DryTor tugas, 7 July 1962. (Photograph hy Nagahisa Kuroda.)


82BULLETIK FLORIDASTATE MUSEUM Vol.8old bay cedars growing amid thick beds ofOpuntiacactus-wasim possible inthetime available.Thebirdwebandedsome 7 weeks later in Septemberwenettedon the shore ucar the same area.Thebird was coming into the temery, and while being handled disgorged a rounded food mass about40mm in diameter, the compacted remains of a largenumberof tiny minnows.TheBrown Noddiesatthe time were still fecding a fcw large younginornear the nests. Therefore whilewestrongly suspect and would like to believethatthe Black Noddy has been nestingatDryTortugas,wehave not as yetbeenabletoprove it. FJGlJIU: 12. Close-up of the head of the first Black Koddy bandedatDry Tor tugas, Bush Key, 1-3 July 1962. (Photograph byJamesB.Meade.)LlTERAnmECITEDManydataof interest found during preparation of thispaperhavenotbeen formally published. These are contained in written material of two gcneral kinds.1.Completed manuscripts and official reportsthatbeara definitedatearecitedbydatewith the designation"MS."to in dicatethatthe source isanunpublished manuscript.2.Field notesandother materialnotin completed form are citedas"MS.notes" without date. Copies of most of the manuscripts and manuscript notes are in my possession.Theplace of deposit of the completed manuscript reports and the source ofthemanuscript notesareindi cated in the citations or comments.


1964 ROBERTSON: DRY TORTUGAS TER:"IS 83 Allen,A.A.1944. Report of theA.O.V.Committee onbirdprotection for 194:3. Auk, 61: 622-6:35. Allen, Robert P. 1936. WiththeAudubon wardens. Bird-Lore, 38: 295-297.American Ornithologist,,' U nian.1957. Check-List of North American Birds. PublishedbytheA.O.U.,xiii+691 pp.[111estatement of therange of the SootyTern(p. 2:37), reading in part: "Breeds from Alaeran Reef off Yucatan,theGulfCoast "f Texas, Louisi ana, Florida (the Dry Tortugas, in recent times) ... ,"is in error bothasregards recent breeding at Tortugas and the present existenceofestablishedcolonies in Louisiana and probably also in Texas.]Ashe, T. J. MS. notes. Population estimates of Noddies in 1910 and 1915. [From entriesintheBird Distribution file, Patuxent Wildlife Research Center, noted for mebyChandlerS.Robbins.] 1919 'IS. Warden's report, BirdKey,DryTortugas. Original eopy in files of Research Department, National Audubon Society. Holographic, 4 pp.[Dated6 October 1919. Summary of 1919 nesting season withpopulation estimates. Hurricane damage to vegetation on Bird Key.]Ashe, T. J.,andLudwig Bethel191.5 MS.'Varden's aIlllual report, Bird Key,DryTortugas. National Audubon Society files,NewYork. [PreparedbyAshe from information pro videdbyAssistantWardenBethel. Estimates 107,000 "terns". Reportsheavy mortalityofLents caused byhurricanesof la August and 3 Sep tem]wr 191.5.] Ashe, T. J., and William E. Lowe 1918 MS. Warden'sannualreport, Bird Key,DryTortugas. Original copy in files of Research Department, National Audubon Society. Holo graphic (Ashe), 5 pp.[WrittenbyAshe from information providedbyLowe, his a

84BULLETINFLORIDA STATE MUSEUM Vol. 8l Austin, Oliver L. 1949. Site tenacity, a behavior trait of the Common Tern (Sterna hirundo Linn.). Bird-Banding, 20: 1-39. Austin, Oliver L., Jr. 1962 Comparative demographies of Sterna hirundo and Sterna fuscata. Delivered beforethe13th International Ornithological Congress (Ithaca, 1962). Bache,A.D. 1858. Report of the Superintendent oftheCoast Survey, showing the progress of the survey duringtheyear 1857. Executive Document No. 33,35thCongress,1st Session. Washington. 'Villiam D. Harris, Printer.Bache, Harbnan1845 MS. Topographical survey of Garden Key. Drawer 74-1,filesofU.S.Corps of Engineers, Military Construction Branch, Construction Divi sion,WarDepartment, Washington, D. C. [Map of the island asitwas immediately before construction ofFortJefferson began. I haveseennonotes that may have accompanied the map, nor correspondencerelating to it.] Baker, John H. 1944.Thedirector reports to you. Audubon Magazine, 46: 178-183. Bartsch, Paul 1916. Birds observed ontheFlorida Keysandalongtherailroad ofthemain land from Key Largo to Miami, June 17-July 1, 1915. Carnegie Inst. Washington, Yearbook for 1915 (No.14):197-199. 1917. Birds observedin1916,intheregionofMiami andtheFlorida Keys from May 15toJune 4,andalong the railroad from Key West to Miami onJune24. Carnegie Inst. Washington, Yearbook for 1916 (No. 15): 182-188. 1918. Fifth annual list of birds observed on the Florida Keys. Carnegie Inst. Washington, Yearbook for 1917 (No. 16): 170-173. 1919.Thebirdrookeries oftbeTortugas. Smithsonian Institution, Annual Report for 1917: 469-500, 38 plates. MS. notes. Estimate of Noddy populationsin1922; observations of Common Ternsin1924 and 1925; observations of Roseate Terns in 1921 through 1925; and observations of Least Ternsin1921 and 1924. [From en triesintheBird Distribution file, Patuxent Wildlife Research Center, taken from field notesbyBartsch. Compiled for mebyChandlerS.Robbins.] 1923. Birds of the Florida Keys. Carnegie Inst. Washington, Yearbook for 1922 (No. 21): 165. 1931. Report on Cerion eolonies planted on Florida Keys. CarnegieInst.Washington, Yearbook for 1930-1931 (No. 30): 373-378. 1932.TheBird Rookeries oftheTortugas. Carnegie Inst. Washington, Year book for 1931-1932 (No. 31): 281.


1964 ROBERTSON: DRY TORTUGASTERNS 8.5 1933. Cerion Studies. Carnegie Inst. of \Vashington, Yearbook for 1932 1933 (No. 32): 266-267. Beard, Daniel B. 1938.Tbe1938 count of NoddyandSootyTemsattheDryTortugas. FloridaNaturalist. 12: 7-10. 1938MS.Special report 1938 count of Sooty and Noddy TernsatFortJeffer son National Park Service files. Typewritten, 8pp.[Observa tions of 20-25June1938. Similar tothepublishedreport(Beard, 1938),butineludes comments ontheadvisability of continuingbanding terns and more extended comments onthegreat decreaseofNoddiesnotedin1938.] 1939. Man-o'-war-birdspreyon eastern Sooty Terns. Auk, 56: 327-329. Bent, Arthur Cleveland 1921. Life HistoriesofNorth Americangullsanelterns. Bull. U.S. Nat. Mus. 113: x+345 pp., 93 plates. Bethel, Luelwig1916. :MS. 'Varden's annual report, Bird Key, Dry Tortngas. National AudubonSociety files, New York. Bond, James 1956. Check-Listof Birds oftheWestIndies. ix+214 pp., frontis.TheAcademy of Natural Sciences of Philadelphia. 1958.ThirdSupplement to the Clwek-Iist of Birds oftheWestIndies (1956).11pp. Academy of Natural Sciences of Philadelphia. Bowman,H.H.M.1918. Botanical eeology oftheDryTortugas. Carnegie Inst. of Washington, Papers fromtheTortugas Lahoratory, 12: 109-138. Bryant, Hemy18.59a.[Birds observed in cast Florida, south of St. Augustine]. Proc. Boston Soc. Nat. flist., 7: 5-21. (Read 19 January 1859). 1859b. A list of birds seenattheBahamas, fromJanuary20 to May14,1859,with descriptionsofneworlittle-known species. Proe. Boston Soc.Nat. IIist.7:102-1.34. Budlong, Robert R. 1942MS.Annual wildlifereportFortJefferson National Monument. Na tional Park Service files. Typewritten 3 pp.[Dated4October1942.Comments on nesting terns presumably refer to the 1942 nesting season.]1943MS.Annual report oftheCustodian.FortJefferson National Monu ment. NationalParkService files. [Forthefiscal year ending30June 1943.OfthisreportI have seen onlythematerial on nestingterns that is quotedinletters from Director, National Park Service, toSecretariesofWarandNavy protesting aerial hom bingandmaehine gunning practicenearBush Key.]t944MS.Custodian's monthly narrative report for :\clay 1944. National Park Service files. Typewritten, 3pp.


86 BULLETINFLORIDASTATE Vol. 8 Chapin, James P.,andL. W.Wing1959.TheWideawake calendar, 195:3to 19,;8. Auk, 76: 1,;:3-V'8. Coast Pilot 1889. Atlantic Local Coast Pilot, Sub-Division 22. Straits of Florida, Jupiter Inlet totheDryTortugas. First Edition. U.S.CoastandGeodetic Sur vey. GPO, Washington. viii+91pp. 1936. United States Coast Pilot, Gulf Coast, KeyWesttotheRio Grande.Second Edition. U.S.Dept.ofCommerce, Coast and GeodeticSurvey.GPO, Washington vi+397 pp.Coast Survey 1878. Report oftheSuperintendent of the U.S. Coast Survey duringtheYear 1875. ExecutiveDocumentNo.81,HouseofRepresentatives,44th Congress, 1st Session. GPO, \Vashington. Cullum.G.W.1891. BiographicalRegister ofthe Officers and Graduates of the U.S. I\lilitary Academy. Vol. 1(1802-1840). Davis, John H., Jr,1942.Theecologyof the vegetation and topographyofthesand keys of Florida. Carnegie Inst. 'Vashington, Papers from the Tortugas Lahoratory, 33: 113-19.5.Davis, T. Frederick1935. HistoryofJuan Ponce de Leon's tu Florida. Florida HistoricalQuarterly, 14:70pp. DeWeese, John H.19.5.'3Superintendent's monthly narrativereport for June 1953. l\ational Park ServicemesoTypewritt{'n, 2 pp. [Connt of nests of Ros('ate TernsonMiddle, Hospital.andBush Keys.] Dilley, Williard E.1949 10.15. Reportof the terns of Fort Jefferson National 1\'fonument for the year 1949.Everglades National Parkfiles.Mimeographed)7 pp. 1950.Theterns oftheDryTortugas, June, 1949. Florida NaturalL,t, 23: 67-68. Dickinson,J.C., Jr. 1941. NoddyandSooty Terns nestingonbare ground. Auk, 58: 2059,Doc, Charles EMS.notes. Notes relating lo the Florida AudubunSudetyTortugas trips of1935,1936,and1937 and holographicdraftof Doe's artiele describingthe 1936 trip. [Extracted from field notebooks in the files oftheFlorida Stale MuscumbyOliverI..Austin, Jr.] Doc, Charles E.,andJack C. Russell19:36.Tortugas colonies again visited. Florida Naturalist, 10: 6-8.


r 1964ROBERTSON:DRYTORTUGASTERNS87Dunn, Cordon E., and Banner I. 1960. Atlantic Hurricanes. Louisiana State Oniversity Press. xx +326pp., 87 figures.Dutcher,William190,'3.Reportofthe A.G.tT. committee on the protectionofNorth Am.erkan birds. Auk, 20: 101-159.1904. Report oftheA.D.V.committee on the protection of North Americanbirds fortheyear1903. Auk, 21: 97-208. 1905.AnnualreportoftheNationalAssociationofAudubonSocieties for 1905. State reports, Florida. Bird-Lore, 7::314.1906. Bird Key, Tortugas, Florida. Bird-Lore, 8: 146.19080. The Tortugas Reservatiun. Bird-Lure, 10: 142-143.1908b.Reservation news, Turtllgas, Florida. Bird-Lure, 10: 187. England, Geurge Allan 1928. Bird Key.SaturdayEveningPost, 201: 14-1.5, 8.5-86, 88,IIphutographsoftenIS.Felton, James B.1940 MS. Custodian's monthly narrative report lorJanuary 1940. NationalPark Servi('e Files.Typewritten, a pp.1941 MS. Custudian'smonthlyreportforJuly1941.Typewritten,4 pp. :National ParkServicefiles.[BriefcommentsOilthepatternof landing uf Suoty TernsunBlish Key.] Fisher, James,andR.M. 1.oekley1954. Sea-Birds.Houghtonl\litHin Company, Boston. xvi+320pp., 8eolor plates,40plates,.55 figure,. Gauld, George 1790.Anaccurate Chartof the Tortugas and Florida Keys or Surveyed by GeorgeGauld, iI..:M. in tbeYears 177.3, 4,&5, By OrderOf The RightIlunourable The Lords Commissiuners OfThcAdmiralty, AndnO\\' Published hy pcnllissionofTheir Lordships. London: Pub lishedby\Vm. Faden, Geographer of the King, Charing Cross, 5thApril, 1790. Second Edition: 1820. COITected at HydrographicOffice,Admiraltytu 18,'3,5. [The copy seen was printed much later. It in dudes an inset map

88BULLETINFLORIDASTATE MUSECM Vol. 8pherToHis Majesty, and ToHisRoyal/HighnessThePrince of Wales. / 1796. 28pp.Gibbs, RussellA.1946 MS. Custodian'smonthlynarrativereportforMay1946. National Park Service files.Typewritten, 4 pp. [As of30 May "someRoseate Ternshadaroved."]1947 MS.Custodian's monthlyreportfor April, 1947.Typewritten,3pp.NationalParkService files. [Brief discussionofthe landing ofSooty Terns onGardenKeyin1947.] Hallman, Roy C. 1961.CommonTern(Sterna hirundo) nesting innorthwestFlorida.FloridaNahualist,34: 221-222.Heermann,Adolphus L. 1852-1853.Catalogueoftheoological collectioninthe Academy ofNaturalSciencesofPhiladelphia. Proe. Academy Nat. Sci. Philadelphia, 6: 1-35. Hogan, J. 1925. Bird notes from Willis Island.Emu,24: 266-27.5.Holden[sic],J.B. 1868.TheDryTortugas.Harper'sNewMonthly Magazine, 37: 260-267. Holder, CharlesF.1892. Along theFloridaReef.D.Appleton and Company,NewYork.ix+273 pp., many textfigures. Howell,ArthurH.1932. FloridaBirdLife. Publlihed byFloridaDept.of Gam"and FreshWaterFishandBureauof Biological Surve}", U. S.D.A.CowardMcCann, Inc., Kew York. xxiv+579pp., 58 plates, 72 t.e,,1: figures. JDb,HerbertKeightley 1905.WildWings.HoughtonMifflinandCo. Boston. xxiv+:341pp.160photos. Krider,John1879.Forty Yean; NotesofaFieldOrnithologist. Philadelphia, xi +84pp.Lashley, K.S.1915. Kotes onthenesting activitiesoftheNoddyandSooty Terns.CarnegieInstitutionof'Vashington, Papers from the Tortugas Laboratory, 7:61-83. Longley, WilliamII.1927.Lifeon aCoralReef. National Geographic Magazine, 51: 61-83. Lowe, WilliamE.1917.MS.Warden'sannualreport,Bird Key, DryTortugas.NationalAuduhon Society files,NewYork.


1964 ROBERTSON: DRY TORTUGASTERNS89 Longstreet, R. J. 19360.TheFlorida Audubon Society Tortugas expedition of 1935.PartII, Part III. Florida Naturalist, 9: 22-42. 1936b. Notes fromtheDryTortugas. Auk, 53: 99-100. 1937. Bird population oftheDryTortugas. Florida Naturalist, 11: 7-10. Lowery, George H., Jr.,andRobert J.Newman1954.ThebirdsoftheGulf of Mexico, pp. 519-540.In"Gulf of Mexico Its Origin, Waters,andMarine Life," Fishery Bulletin 89, U.S. FishandWildlife Service. Manucy, Albert 1943.TheGibraltar oftheGulf of Mexico. Florida Historical Quarterly, 21: 303-331. 1961 MS. A constructional history ofFortJefferson 1646-1874. National Park Service Jiles. Typewritten, 242 pp. [Doeumcntcdreportbasedupon searchofthe original records in archives, letters, and otherbasicsources. Construction activities, 1846-65, arc treatedindetailwitha brief resume of concurrent general historyofthearea. Constructionthrough 1874isreported in less dctail. Brief comment onlatereventsthat affected the stmetures to 1935.) Mason,C.Russell MS. notes.Fieldnotes relating totheFlorida Audubon Society trips of 1935 through 1938. 1936.TheFloridaAudubonSociety Tortugas expedition of 1935.Part1.Florida Naturalist, 9: 17-22. 1938. Society conducts fourth Tortugas trip. Florida Naturalist, 12: 1-6. Mayer, Alfred Goldsborough 1908. Tortugas Reservation. Bird-Lore, 10: 229. Maynard, C.].1881. Birds ofEasternNorth America. iv+532 pp.,32plates. Millspaugh, Charles Frederick 1907.Floraofthesandkeys of Florida. Field Columbian Museum, Botanical Series 2 [Publication 118]: 191-243. J osepb C. 1954 MS. 1953 and 1954 censuses of tern colonies ofFortJefferson National MonulIlent. Everglades Kational Park Jiles. Typcwritten, 4 pp. [Summarizes censusesofSooties in 1952, 1953, and 19.54 based upon counts of nestsin20or30sample areas each of8 sq. yds. CensusesofNod dies and Roseateshydirect countofnests. Difficultiesofconsusing becau'ie of the prolonged arrival periodofSuuties.]1955 ).IS. Tern census of Fort Jefferson Kational for 1955. Ever I(lades National Park Jiles. Typewritten, 4 pp.[Dated7June1955. Estimatcs 71,102 Sooties, lowcst sincc 1947.Healthof colony poor, unusual numbers ofdeadadultsandyoung seen. Recorded 1218 Nod diesand436 Roseates bothbydirect count of nests.)


90 BULLETI1\' FLORIDASTATEMUSEUMVol. 8'IS.notes. [Field notesonhis observationsatDryTortugasin1952,1954,and1955.Otherfield notesandcalculationsandseveralvegetationmapsofBush Key, allpertainingtotheearly1950's.] Moore, Joseph C.,andWilliardE.Dilley 1953.Terncounts atDryTortugas 1950,19.51,and1952.FloridaNaturalist,26: 75-81. Robert Cu>hman19.36. Oceanic BirdsofSouth America. Volume 2.TheMacmillanCompany,TbeAmerican MuseumofNatmalI1istory,NewYork. xii+pp.641 124.",text figures 62-80, plates 39-72, 10unnumberedcolored plates.Pace, JerryF.1962.BandingLeastTernsinFlorida.EBBA News. 25: 191-193. Palmer,RalphS.,andE. Reilly, Jr.1956. A concise color standard. 8pp.PublishedbyAmerican Ornithologists'UnionHandbookFun.Park, Charles I. MS. notes. Populations of l\oddies in 1929 andofLeast Ternsin1929and 19:30. [From entries intheBird Distribution file, Patuxent WildlifeResearch Center, apparently based upon Park's warden reportstothe Biological Survey.CompiledformebyChandlerS.Robbins.] Pcacon,John1909 MS. Warden'sannualreport, the islands oftheTortugasGroup(Bird Key). KationalAudubon Society files,NewYork. [Attachedtotbereport arc a sketch map of the islands of the Tortugas and copies of correspondence ht>tween Peacon and T.J.Ashe.] 1911 \Varden's aUllualreport, Bird Key,DryTortugas. National AudubonSociety files, :-lew York.1912 MS. 'Vanlen's:-lllIlllalreport, Bird Key Dry Tortugas. AudubonSociety Illes, NewYork. 191.3MS.'Varden's annual report, Bird Key, DryTortugas, National Audu bonSociety 1\'cw York.1914 :\IS. 'Varden's annual report, Bird Key, DryTortugas. AudubonSoei(,tyfil(,s, New York. [AllinfonnationabouttI,e report< ofJohn Peacon is from notes made formebyRobert P. Allen. I have !'ieennone of the original reports.] Pearson,T.Gilbert1915. ... Iotion-pictures fortheNational Association. Bird-Lore, 17: 410-412. 1915 MS. DryTortnga", Florida. Typewritten, 5pp.Files of Research Deparbnent, 1\ational Audubon Society.[Alongaccountofthe condition of the colonyin1915 mainly as observed byRev.H.K.Job,Dr.H. R. Mills,andCapt.T.J. Ashewhovisited Bird Key 28 May1915. Quotes extensively from Job's reportofthis trip and from the annual reportofAshe. used an area-density technique to esti-


1964 ROBERTSO:'lr: DRY TORTUGAS TERNS91matethepopulation of Sooties. Ashe reportedthatterus su!fered heavy mortality as a resultofthehurrieaues of 13 August and 5 September 1915. This ms. appearstoheeopypreparedbyPearsoufor publication in Bird-Lore, butonlya brief excerpt fromitwaSpublished.] 1917. Report of T. Gilbert Pearson, Secretary AuduhonWardenWork. Bird Lore, 19: 397-402. 1918. Report of T. Gilbert Pearson, Secretary AudubonWardenWork. Bird Lore 20: 459-460. 1919. AudubonwardenworkinFifteenthAnnualreportoftheNationalAssociation of Audubon Societies. Reportof the Secretary. Bird-Lore,21: 401-404. Peterson, Roger Tory 1947. A Field Guide totheBirds.Houghton :'li/llin Co., Boston. xxiv-+ 290pp.,60plates, line drawings.1950. BirdsOverAmerica. Dodd, Meade andCompany,NewYork. xiii+342 pp., 105 photographs. Peterson, Roger Tory, and James Fisher195.5.Wild America. Houghton I>lifHin Company. Boston. xii+434 pages. Phelps. William II.,andWilliamII.Phelps,]r.1958. Listadelas ayes de Venezuela con su distribucion. Tomo II, Parte 1, NoPasscrifonnes. Bol. Soc. Venezolana deCiencia..;;Nat., 90: 1-317.Pough, Richard H. 1951. AuduhonWaterBird Guide. Doubleday and Co.,NewYork. xxviii+352pp.,48 plates, Hum.crousline drawings. Rea, t\,largaret P., Jennie Lynne Kyle, and Louis A. Stimson1941.The1941 Florida Audubon Society ('xpedition to the Dry Torlugas. Florida Naturalist, 15: 16-17. Richardson, Frank,andHarvey I. Fisher 1950. Birds of Moku Manuand Manana Islands offOahu,Hawaii. Auk, 67: 285-306. Rid1<-y, W., andLordR.C.Percy1958.Theexploitationofsea birds inthe Seychelles, Colonial Research Studies :-lo. 25, pp. 1-78.Robert, Henry, and Henry Stevenson1951. :-lotes on birds on the DryTortugas. Florida I\aturalist, 24: 100-10.5, 108. Robertson, WilliamB.,Jr.1956 MS. 19.56 teru eount. Everglades National Park files. Typewritten, 4 pp. 1962. Florida Region. AudubonField Noles 16: 468-473.


92BULLETINFLORIDASTATE MUSEUMVol.8Robertson, William B., Jr., DennisR.Paulson,andC. Russell Mason 1961. AternnewtotheUnited States. Auk, 78: 423-425. Robinson, GeorgeDigby1939. 1939 Tortugas E''1)edition FirstGroup--June5-10. Florida Nat uralist, 13: 7-12. 1940. 1940 Tortugas expedition. Florida Naturalist, 14: 1-6. Russell, Jack C. 1938 MS. Narrative report on wildlife ofFortJeHerson National Monument. National Park Service files. Typewritten, 39pp.[Dated24 February 1938. Report of observations made during the author's employment as a student wildlife technician,10June-15 September 1937, with notes from earlier visits in1935and1936.The first 24 pages deal chiefly with birds; the remainder with marine life, principally mollusks.Discusses all nesting ternsindetail.]Scott,W.E. D. 1890.Onbirds observed attheDryTortugas, Florida, during parts of March and April, 1890. Auk, 7: 301-314. 1904.TheStory of a Bird Lover,xi+372 pp. Macmillan,NewYork. Sprunt, Alexander, Jr. 1946a. Surveyoftheterncolomes of the DryTortugas season of 1945, Flor ida Naturalist, 19: 26-34. 1946b. Population survey ofthetern colomes of the DryTortugasFortJeHer son National Monument 1946. Florida Naturalist, 20: 1-8. 1947a,1948a. Pupulation survey,terncolumes oftheDryTortugas,FortJeHerson National Monument, 1947. Florida Naturalist, 21[Ocl., 1947]: 16-21; 21[]an., 1948]: 2,5-31. 1947b. Blizzard of birds:TheTortugas terns. National Geugraphie Magazine, 91: 213-2:30. 1948b.Theterneolunies of the Dry Tortugas Keys. Auk, 65: 1-19.1948c.Tern coloniesoftheDryTortugas, Fort Jefferson National 1948. Florida Naturalist, 22: 9-16.1949. StatusofRoseate Tern as a breeding species in southern United States.Auk, 66:206-207. 1951. A list ofthebirds uf theDryTortugas Keys 1857-1951. Florida AuduhonSuciety, 27 pp., 5 phutographs. 1962. Birds of theDryTortngas 1857-1961. Flurida Xaturalist, 35: 34-40, 58, 82-85, 129-132.Stevenson, Henry :\:[. 1956. Florida Region. Audubon Field Notes, 10: 325-329.StevensOD, James O.1938.Theterncolomes ofDryTortugas. Bird-Lore, 40: :305-S09. Stewart, James, Jr. 1962. Central Suuthern Rcgiou. Audubon Field Notes, 16: 480-481,485-486.


,1964ROBERTSON: DRY TORTUGAS TERNS 93 Sutherland, Charles Alan 1961. Excursion toanavian island. Florida Naturalist, 34: 7-10. Tannehill, Ivan Ray 1950. Hurricanes (7th Edition). Princeton Univcrsity Press, Princeton. x+304 pp., 135 figures. Taylor, O. B. 1939 MS. Special report on wildlifeFortJefferson National Monument. Na tional Park Service files. Typewritten, 8 pp. [Based on a visit of 19-21 June 1939. Brief commcnts on nesting ternsandsea turtles with adiscussionofthe measures needed to improve protectionofwild..lifeinthe arc a.) Thompson, Joseph 1903. The Tortugas tcrn colony. Bird-Lore, 5: 77-84.Uaited States Coast and Geodetic Survey 1896. Tortugas Harbor and Approaches. W'CGS 471a. Scale 1:40,000. 1958.DryTortugas. CGS 585. Scale 1:30,000. Vaughan, Thomas Wayland 1914. The huilding of the Marquesas and Tortugas atolls and a sketchofthegeologic history of the Florida reef tract. Carnegie Inst. of Wash ington, Papers fromthcTortugas Laboratory, 5: 55-67. Vinten, C.R.1943. The Noddy and SootyTerncolonies oftheDryTortugasFortJefferson National Monument: A summary of the records. Florida Naturalist, 16: 53-61. 1944. Tortugas Note. Florida Naturalist, 17: 71-72.Voous,K.H. 1957.Thebirds of Aruba, Curacao,andBonaire. Studies ontheFaunaofCuracaoandother Caribbean Islands, 8: 1-260. 1960. Atlas of European Birds. Nelson, London. 284 pp. 1963. Tern colonicsinAruba, Curacao and Bonaire, south Caribbean Sea. Proc.XIllIntern. Ornitho!. Congr.: 1214-1216. Warham, John 1956. Observationsonthe birds of Pclsart Island.Emu,56: 83-93. Watson, John B. 1907. Report of JohnB.WatsonontheconditionoftheNoddyandSooty Tcrn colony on Bird Key, Tortugas, Florida. Bird-Lore,9:307-316. 1908.Thebchavior of Noddy and Sooty Tcrns. Carnegie Inst. Washington, Papers fromtheTortugas Laboratory, 2: 187-255. 1910. Further data onthehoming sense of Noddy and Sooty Terns. Science, 32 (n.s.): 470-473.


94BULLETINFLORIDASTATE Vol.8l1912 MS. SpecialreportTortugas Reservation. NationalAudubonSociety files,NewYork. [StatesthathevisitedBird Key too lateinthe seasonto make accurate population estimates (Carnegie Inst. \VashingtoD,Yearbook for 1912 (No. 11):12.5,however, givesthedates of Watson's 1912 workintheareaas 6 June-12 July). Recommends planting of "large quantities" ofbaycedaronBird Keytorepairdamagedone to vegetationbythehurricane of 1910.FromnotesbyRohertP.Allen. I have notseentheoriginal.]Watson,JohnB.,and K. S.Lashley1915. An historicalandexperimental study of homing. Carnegie In

CATALOGUEOFFOSSIL BIRDS Part I (Archaeopterygiformes through Ardeiformes)PIERCEBRODKORB'SYNOPSIS:Thefirst installment of a catalogue ofthefossil birds oftheworMcovers49families in15ordersofbirds,ornearly. half of the orders and aboutone-fourth. ofthefamilies known.Thespeciestreatednumber374, of whieh273 are extinct, and 101 represent living species recorded from fossilorprehistoric sites.Forthepalcospccicsthedataincludecitationoftheoriginaldescription, synonyms, nature and repositoryoftypes, reference to pertinent revisionary papers, and detailed geological and geographic ranges, with bibliographic reference to their occurrence.Majortaxonomic changes include recognitionofthree subclasses, Sauriurae forArchaeopteryx, Odontuholcae fortheHesperornithidae, and Ornithurae for the remaining birds. Three infrac1asses of Ornithurae arc recognized, Dromaeognathae(fortheTinamidae),Ratitae,andCarinatae. Changesinposition include transfer ofthefamily OpisthodactylidaetotheRheiformes, Enaliomithidae totheGaviiformes, and BaptornithidactothePodicipedifonnes. On prioritytheordinal name Ciconiiformes yieldstoArdeifonnes. Prior family names adopted include Emeidae for Anomaloptcrygidac. Occanitidae for I-Iydrobatidae,andPlataleidae for Thrcskiomithidae.Newtaxa proposed are Colymboidinae(newsubfamily, Gaviidae).Cayetanornis(newgenus,Tinamidae),andPalaeeudyptes marplesi(newspecies, Spheniscidae). The misprinted name Pelagodomithidae is emended to Plega4 dornithidae, to conformwiththespelling ofthetype-genus.lTheauthor is ProfessorofBiological Sciences attheUniversityofFlorida, Gainesville. Manuscript received28January1963.-ED.

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180BULLETINFLORIDASTATE MUSEUMTABLEOFCONTENTSVol. 7INTRODUCTION186 CLASS AVES Linnaeus .........................................189 SUBCLASStSAURlURAEHacckcl 189tARCHAEOPTERYGIFORMESFiirbringer 189t ArchaeopterygidaeHuxley...............................189tArchaeopteryxMeyer...........189 SUBCLASStoDONTOHOLCAEStejneger 191 tHEsPERORNITHIFORMES (Fiirbringer) 191j'Hespcrornithidae Marsh 191 tHesperornisMarsh191tConiornisMarsh 192 SUBCLASS ORN1THURAE Haeckel 193 1NFRACLASS DROMAEOGNATHAEHuxley....................193 'l'rNAMIFOllMES (Huxley) 193Tinamic1ac Gray193tTinamisornisRovercto 193tCayetanornisBrodkorb ....................... ......194 tQuerandi(}misRusconi ............... ........... 194NothuraWagler194Neospecies of Tinamidae ....................... ....... 194INFRACLASS RATITAE Merrem 196STRUTHIONIFORMES(Latham)196tEleutherornithidaeWetmore................. ...........196 tEleutherornis 196StruthionidaeVigors.....................................196StruthioLinnaeus 196 NeospeciesofStruthionidae 199RHEIFORMES(Forbes)200 tOpisthodactylidae Amcghino 200 tOp'isthodoctylus Arneghino 200 Rheidae(Bonaparte)200 tH eterorheaRovereto 200RheaBrisson 200 Pt(''Tocnemia Gray "201 Neospecies of Rheidae 201CASUARIIFORMES(Sclater)202Casuariidae Kaup202 Casuarius Brisson202Dromiceiidae Wetmore202VromiceiusVieillot 202 :r\eospecie.s of Dromicciidae203tDromornithidae Fiirbringer "203tDromornisOwen 2Q,3 tGenyornisStirlingandZietz 20:3 tAEPYORNITHTFORMES(Newton)205 t Aepyornithidae(Bonaparte)20.5tEremopezinae Lambrecht 205tEremopezu8Andrews'"" 205

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 181 t Aepyornithinae(Bonaparte)205tStromeriaLamhrecht205tMulleTornisMilne-Edwards and Grandidier ,206tAepyornisGeoffroy .................................206tD'NORN'TH'FORMES(Gadow)208tEmeidae(Bonaparte)208 t Anomalopteryginae (Oliver) 208 tAnomaloptenjxReichenbach 208tMegalapteryxHaast209 tPachyomisLydekker 210tEmcinacBonaparte212t Eme'Us Reichenhach .......................212 t EuryaptenjxHaast 213 tZelornisOliver 214tDinomithidaeBonaparte215tDinornisOwen....................................215APTERYGIFORMES(Haeckel)219 Apterygidae(Gray)219tPseudapteryxLydekker 219NeospeciesofApterygidae ....... .......................219INFRACLASS CARINATAE Merrcm ...220 GAVIlFORMES WetmoreandW.D.Miller....................... 22.0 tEnaliornithidac Fiirhringer 220tEnaliornisSeeley 220tLonchodytidaeBrodkorb,221tLnnchodytesBrodkorb .........221 Gaviidae Allen222tColYIIlhoidinae Brodkorb .............................. 222 tEupterornisLemoine 222tCulymboidesMilne-Edwards.........................222tGaviellinaeWetmore223tGaviellaWetmore.................................. 228 Gaviinae(Allen)223CaviaForster 22.'3 Neospecies of Caviinae224 PODlCll'.IilllFOHMBS (Fiirbringer)226tBaptornithidac American Omithologist.<;' Union226tBaptornisMarsh .................. 226 tNeogaeoNlis Lambrecht.............................226Podicipedidae(Bonaparte)226PodicspsLatham227 tPliodytesBrodkorb 228 Neospeeies of Podieipedidae ............................228 Sharpe.....................................231 Spheniscidae Bonaparte 231tPalaeeudyptinaeSimpson 231tP"lueeudyptesHuxley 231t Pachydypte" Oliver .................................232tArchaeuspheniscusMarPles 232tDuntroonornisMarples 232

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182BULLETINFLORIDASTATEMUSEUMVol. 7 t PlatydyptesMarples 233 tKororaMal1'!cs ............. 2,33 tAnthropodytesSimpson 234 tN otodyptesMarples 234t Anthropornis Wiman234 tOrthopteryx Wilnan234tEosphaeniscusWiman..' ,... 235tDelphinomis ,Viman '.,,, ,.. 235tlchtyapteryxWiman,. 23,1) tArthrodytesAmeghino ..............................235tPalaeosphcniscinaeSimpson................ ...........236 'rPalaeosphenisctts Moreno and Mercerat ',..236tPerispheniscusAmeghino ,.............237 t ParaspheniscusAmeghino,............238tParaptenodytinaeSimpson ,.... 238 tParal'tenodytes Ameghino ....................... 238 t!sotrcmornis Ameghino ............ 239tPscudospheniscw; Ameghino239tNeculusAmeghino.,,........240Spheniscinae Bonaparte .................240N eospecies ofSpheniscinae .............................240 Furbringer 241Diomedeidae(Gray),,....241tGigantornisAndrews241tManuMarples,'"241DiomedeaLinnaeus241 ofDiomedeidae.............................242Procellariidae (Boie),'..............242 Puffinus Brisson242tArgyrodyptes Ameghino ,......................245 tPl<>tmnisMilne-Edwards ....................... 24,5 NeospeciesofProcellariidae ................245Oceanitidae (Salvin) 246Oceanodruma Reichenbach.., ,,.....247NeospeciesofOeeanitidae ..............................247Pelecanoididae (Gray) 247 Neo
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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS 183 Neospedes of Phalacrocoracinae 254 Anhingidae Ridgway 256tPrntoplotusLamhrecht ...........................256AnhingaBrisson 256 Neospecies of Anhingidae .................. 257 Sulidae (Reichenhach) 257Suw Brisson ....................................... 257tMicrosula Wetmore.....259MorosVieillot .....................................259 tPoloeosuZo Howard ..............................260 tM iosuZo L.Miller .......................260 Neospecies of'Sulidae .................................. 261 PHAETHONTESSharpe ...................................... 261 Phaethontidae (Bonaparte)..........................261tProphoetonAndrews 261 Neospecies of Phaethontidae 262tODONTOPTERYCIASpulski ..................................262 tOdontoptcrygidae Lamhrecht 262tOdontopteryxOwen 262 tPseudodontornithidae Lambrecht ......................262tPseudndnntnrnisLambrecht........................... 262tOsteodontornisHoward 263 tPelagornithidae (Fiirbringer) 263 tPe/agnrnisLartet 263tCLADORNITHESWetmore264tCladornithidae Ameghino ................................264tCladornisAmeghino ......................... 264 tCyphornithidae Wetmore.,..............................264tCyphornisCope 264 tPaweochenoides Shufeldt ............................264PELECANISharpe 265 Pelecanidae Vigors 265Pelecanu8Linnaeus265tLiptorni . Ameghino 267 Neospecics of Pelecanidae 267 FREGATAE(Sharpe)268 Fregatidae Garrod 268NeospeciesofFregatidae268AHDEIFORMES(Wagler)269PHOENICOPTERIFtirbringer269tToroligidae Brodkorb 269tGaUornisLamhrecht ................................269tParascaniurnisLambrecht269tTorotixBrodkorb ................................... 270 i'Scaniornithidae Lambrecht 270 i'Sconiornis Dames 270 i'Telmabatidae Howard 270 tTe/mahatesHoward 270 tAgnopteridacLamhrecht 270 tAgnopterosMilne-Edwards 271

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184BULLETINFLORIDASTATE MUSEUM Vol. 7 Phoeni'Copteridae Bonaparte271tElornisAymard 271tTiliomisAmcghino272 Phnenicopterus Linnaeus272 Neospecies ofPhoenicopteridae ..........................273 tPalaelodidae (Stejncger) 274IPalaelodusMilne-Edwards.274 tMegapaloelodus A.H.Miller........................276 PLATALEAl:::: A.Newton.....................................276tPlcgadornithidae(Wetmore)276tPlegadornisWetmore...............................276PlataleidaeBonaparte ....................................277Thrcskiomithinae(Riehmond)277 tIbidopsis Lydekker .................................277 tIlJidopodia Milne-Edwards 277 EudocimusWa!!:ler .....................278 tProtibisAmeghino 278PlegadisKaup 278CarphibisReichenbach 279 Plataleinae Bonaparte .................................. 279 PlutuleaLinnaclls 279Neosl'ccies of Plataleidae 279AnDEAE Wa!!:ler 280 Ardeidae Vigors 280tPraherodiusLydekker 280 tEoceornis Shufeldt 280t Hotauroide. Shufeldt 280tProardeaLambrecht281tGoliathiaLambreeltt 281tArdeacitesHaushalter 281tRotauritesAmmon 281ArdeaLinnaeus 282NycticoraxForster283tPalaeaphayxMcCoy 283Butor/desBlyth ....................................283 of Ardeidae2H4 Coehleariidac Ridgway 286 Scol'idac (Bonaparte) 2&J Balaenicipitidae (Bonaparte) 286CICONIAEBonaparte .......................................286Ciconiidac(Gray)286Ciconiinae Gray 286 t PelargopappusStejne!!:er 287-i'PropelargusLydekker 287 tPaweoephippiorhyndms Lambrecht ............288tCiconiopsisAmeghino ............................... 288 t Amphipelatgus Lydekker ......................... 288 LeptoptilosLesson 289 Ciconia Brisson289tPelargosteonKretzoi 290

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS18,5tProciconiaAmeghino 290 tPalaeopelargus DeVis ...............................291tXenorhynchopsis DeVis 291 Xenorhynchus Bonaparte ...........................291 Mycteriinae American Ornithologists' Union292IbisLacepede292MycteriaLinnaeus 292Ncospccics of Ciconiidae ...............................292

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186BULLETINFLORIDA STATE MUSEUMINTHODUCTIONVol.7Ourknowledge of fossil birds has increased greatly during the thirty-year interval sincethepublication of Lambrecht'sHandbuch der Palaeomithologiein 1933. Re-evaluation oftheclassical forms andthediscovery of new Mesozoic material afford an entirely different perspective of the earliest birds:Wenow believetbatthetheory of the toothed birds was mostly fictitious; we realizethatanumberofmodemorders existed duringtheAge of Reptiles;andwe may even hintata possible polyphyletic origin of the class Aves. Discov ery of more than a dozenncwfamilies gives greaterbreadthanddepthto our understanding oftheevolution of birds. New collecting techniques have resulted inanincrease oftheknown fossil speciesbymore than a thirdandthefilling in of the fossil record of many living spccies.Ittherefore secms time to bringthelist of fossil birdsofthe worldupto date.Theclassification adopted hcre is, with modifications,thatof Wet more (1960, Smithsonian misc. ColI., vol. 139, no. 11, pp. 1-37), which has many advantages overtheseveral other recent attcmpts to clas sify birds. Itsmanyeditions have benefited fromtheauthor's rich experiencewithbothlivingandfossil birds, whereas for somc strange reason other systems totally ignore thc fossil record.Theuse of uni form endings for order-group taxaandtheir formation from valid generic namcs arc useful mnemonics, unfortunately abandonedbycertain other authors. And the recognition of intermediate, non-man datory taxa is helpful in suggesting relationships. In matters of nomenclature Iattemptto follow the InternationalCodeof Zoological NomenclatureadoptedbythcXVInternational Congress of Zoology (London, 1961), not without certain misgivings, however.Thenew edition oftheCode for the first timc setsuprules to cover family-group names,butunfortunately these do not all quite followthesame principlesthatgovern genus-groupandspecies-group names. Exercise oftheCommission's plenary powers in suspendingtherulesatits discretion,theestablishment of different effective dates for an increasingnumberof rules,andthenumerous lists of "nomina conservanda" impose a seemingly greater bibliographicandlegalistic chorethanwouldtherigid application of thelawof priority. Abbreviation of serial publications follows Romer, Wright, Edinger,andVanFrank(1962, Bibliography of fossil vertebrates exclusiveofNorth America, 1509-1927, Geological Society of America, Memoir87).Sources not listed in Homeretal.are given in fullwhenfirst citedandabbreviated thereafter.

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1963BRODKORB: CATALOGUEOFFOSSIL BIRDS187Thecatalogue includes all higher taxa of birds. Familieswithno fossil record are included with appropriate notation, to emphasize gaps in our knowlcdge. Bibliographic reference is givcn to the ap parent first valid publication of names employed for order-groupandfamily-group taxa; suchdataare not otherwise summarized in orni thological works ofthelasthundredyears. Complete principal syn onymiesareincluded onlywherenecessarytosubstantiate change from current usage. Abovethelevel ofthespecies, daggers intheheadings differcntiate paleotaxa from neotaxa. Insofarascompatible with phylogenetic considerations, the arrangement of taxa follows geological sequcnce.Thecatalogue admits paleospecies when validly described on diag nostic parts of the skeleton. Unless later corroboratedbyidentifiable bones, species founded upon indeterminate elements, eggshells, feather impressions, footprints, orotherunsatisfactory evidencearerelegated tothecategory of Incertae Sedis. These willbelistedattheendof the catalogue, as will nominanudaandnon-avian forms originally described as birds.Thecoverageunderthe fossil species includes reference to the original description, synonyms,natureof holotypes and museum where preserved,andreference to certain revisionary work. Distribu tionaldatainclude details of the geological horizonandgeographic range, with bibliographic references to such occurrences. Paleospecies are numbered consecutively within a family. Each family concludeswitha list of its living species known as fossils, with bibliographic citation to their geographic occurrence as fossils. Localitics from prehistoric depOSits(markedwith asterisks) are incorporated insofar as they have come to my attention, although a thorough search oftheanthropological Iiteraturc was not made. Neospeciesarenumberedseparately within each family. Much difference of opinion exists regardingtheboundaries of geo logical time units.Forthis reason I have stressed formationsorother rock units,ratherthanso-called provincial faunal ages.Uponcom pletion of thc catalogue a correlation chartisplanned.TheNational Science Foundation aided preparation of the cata logue throughgrantnumberG19595. HildegardeHowardof Los Angeles was good enough toreadthe bulk ofthemanuscript, and Elliot W. Dawson of Wellington, New Zealand, kindly criticizedthesection on the moas. Throughout this work I have benefited greatly through repeated discussion with Alexander \Vetmore of the United States National Museum. Extensive correspondence with JamesFisherof London has proved most useful. Others who havc been helpful in

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188BULLETINFLORIDA STATE MUSEUMVol.7providing specimens or information include Oscar Arredondo, Ha bana, Cuba;WalterAuffenberg, Bouldcr, Colorado; Oliver L. Austin, Jr., Gainesville, Florida; RipleyP.Bullen, Gainesville; WilliamA.Clemens, Jr., Lawrence, Kansas;WaltcrW. Dalquest, Wichita Falls, Texas; Georges Dementiev, Moscow, USSR;J.C. Dickinson, Jr., Gainesville;A.GordonEdmund,Toronto, Canada; J.S.Erskine, Wolfville, Nova Scotia; Richard Estes, Boston, Massachusetts; W. D. Frankforter, Grand Rapids, Michigan; Joseph T. Gregory, Berkeley, California;J.Hill Hamon, Terre Haute, Indiana; Claude W.Hibbard,Ann Arbor, Michigan; J. Alan Holman, Normal, Illinois; Marie L. Hopkins, Pocatello, Idaho; PhilipS.Humphrey, Washington, D. C.; H.G.Kugler, Basel, Switzerland; John F. Lance, Tucson, Arizona; JohnJ.McCoy, J acksonvil1e, Florida; AldenH.Miller, Berkeley; Loye Miller, Davis, California; Rachel H. Nichols, New York,NewYork;StanleyJ.Olsen, Tallahassee, Florida; Bryan Patterson, Cam bridge, Massachusetts; ClaytonE.Ray, Gainesville; DonaldE.Sav age, Berkeley;J.Arnold Shotwell, Eugene, Orcgon; ElwynL.Simons, New Haven, Connecticut; George Gaylord Simpson, Cambridge, Mas sachusetts; Bob H. Slaughter, Dallas, Texas; H. A.Stirton, Berkelcy; Robert W. Storer, Ann Arbor; DWight W. Taylor, Washington; RichardH. Tedford, lHverside, California; HarrisonB.Tordoff, Ann Arbor; William D. Turnbull, Chicago, Illinois; Robert D. Weigel, Normal;DruidWilson, Washington; JohnA.Wilson, Austin, Texas; Robert W. Wilson, Rapid City, South Dakota; ElizabethS.Wing, Gainesville;andDavidB.Wingate, Hamilton, Bermuda.Thepresent installment covers the orders Archacoptcrygiformcs through Ardeiformes, exclusive of the Ichthyornithiformes, which will be treated latcr.Thetwo or three further installmentsneededto com pletethework will, itishoped, appearatfrequent intervals.Theliterature of avian palcontologyissoscatteredthatitisdifficult to avoid overlooking important papers. Therefore authorsareurgedtoscnd additionsandcorrections for inclusion in a supplement tobepublished on conclusion ofthework.

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196.3BRODKORB:CATALOGUEOFFOSSILBIRDSClass AVESLinnaeus 189Aves Linnacus,System aNaturae, ed. 10, vol. 1,pp.12,78(classis).SubclasstSAUlUURAEHaeckel Sauriurae I1aeckel, 1866, Generelle Morphologic clefOrganislIlen,vol. 2, p. cxxxix(Subklasse).-Sallruri Vogt. 187ll,Hev. sci.(Paris),vol. 17, p.247.-SallruraeStejnegcr, 1885, Stand. nat. Hist., vol. 4, p.21(sub-class).-Saurura Steinmann, 1907,Einfiihrung in die PaHiontologie, cd.2,p.460(Unterklasse).ArchaeornithesGadow,1893,Bronn Klass. Ordn., Vogel,pt.2,pp.86, 299 (Unterclasse) .OrdertARCHAEOPTERYGIFORMESFiirbringcr Saurtuae Huxley, 1867, Pmc. zool. Soc. London,p.418 (ortIer, ex subclassSauriuraeHaeckeI; not based on generic]889, Amer. Natural.,vol.23,p. 869 (superorder).-SaururaSteinmannandDoderlcin,1890,EJe menteder PaHiontologie, p. 668 (Ordnung). OrnitlwpappiStejne!'(er, 1885,Stand. nat. Hist., vol.4,p.21(order;notbased011generic name).Archomithes Furbringer,1888,Untersuchungen zur Morphologie undSystcmatikder Vogel, vol. 2, p.156,5(ordo; not h
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190BULLETINFLORIDASTATE MUSEUM Vol. 7Gryphosaurus "A. Wagner," Marschall, 1873, Nomenc1ator Zoologicus, p. 49 (lapsus) .Griphomis"Owen, 1862, Rev. Nat. Hist., p. 313," H. Woodward, 1962(Dec.),Intellectual Observer, vol. 2, no. 5, p. 317(newnameforGriphosaurus). Gryphomis"Owen, 1862, p. 313," Lambrecht, 1933,HandbuchderPalaeorni thologie,p.-80(Iapsus).ArchaeomisPetronievics, 1917 (Apr.20),Proc. zoo!. Soc. London, pt.1,p. 5footnote(typebyIllonotypyArchaeopteryx siemensiiDames).1.Archaeopteryx lithographicaMeyerArchaeopteryx lithographicaH. von Meyer, 1861(afterSept.30),Neues Jahrb.Min. Geol. Pal.,p.679(typefrom Kohler's cut, feather impression in Berlin Mus., reverse in MunichMus.).Griphosaurus problematieu.A.Wagner, 1861(afterNov.9),Sitz.-Ber. bayer. Akad. Wiss., vol. 2, p. 146(typeskeleton from Ottman's cut, Brit. Mus. no.37001). Gripho.aurtlS longieaudatusOwen. 1862. Rev. Nat. Hist., p. 313(newnamefor Griphosaurus problematicusWagner).Archeopteryx macrura Owen,1864, Philos. Trans. Roy.Soc.London for 1863,vol. 15:3, p.33note,pI.1-4(newnameforGriphornis lungicaudlltusOwen).Archaeopteryx macrouraKleinschmidt,1951,Pmc.X. internal. orn. Congress,p. 631 (emendation or lapsus).Archaeopteryx uweni Petronievics, 1917, Proe. zool. Soc. London,p.5(newname forArcheopteryx macruraOwen).UPPERJURASSIC,PORTLANDIAN(SolnhofenerPlattenkalk).BAVARIA:Kohler'scutin communityquarryatSolnhofen (Meyer, 1861);Ottman'scut(Wagner,1861)andOpitsch'squarry(Heller, 1959,Erlanger geologischc Abhandlungen, va!. 31, p.9)atLangenalthcimerHaardtnearPappenheim.2.ArclUleopteryx siemensiiDamesArchaeopteryx siemensii Dames, 1897 (Aug.9),Sitz.-Bcr. Akad. Wi". Berlin, vol.,38,p. 829[po12 of separate],fig.1-2(typeskeleton fromDorrcut, BerlinMus.).UPPERJURASSIC,PORTLANDIAN(SolnhofenerPlattenkalk).BAVARIA:DorrcutatBlumbergncarEichstatt.1lAspointed outby\Vetrnore(1960,Smithsonian misc:. ColI., vol. 139, no. 11. pp.1-3),the arguments for specific identity with A. lithographica are not wholly convincing.

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDSSubclassfODONTOHOLCAEStejneger191OdvntoholcaeStejneger, 1885, Stand. nat. Hist., vol. 4, p. 27 (subclass, ex orderOdontolcaeMarsh;type Hesperornis Marsh).OrderfHESPERORNITHIFORMES(Fiirbringer)OdontolcaeMarsh, 1875, Amer. Jour. Sci., ser. 3, vol. 10, p. 407(order:typeH esperorni.Marsh).OdontolgaeForhes, 1884, Ibis, ser.5,vol.2,no. 5,p.119(order).OdontornithesForbes, 1884, Ibis, ser. 5, vol. 2, no. 5, p. 119(superorderforArchaeopteryx,Hesperornis,andIchthyornL,). Dromaeopapp/Stejneger, 1885, Stand. nat. Hist., vol. 4, p.27(order;typeH esperornis Marsh).HesperornithesFiirbringer, 1888, Untersuch. Morph. Syst. Vogel, vol. 2,pp.1165, 1438, 1541("gens;"typeHesperornisMarsh) .-HesperornithifoT11l' Sharpe, 1899, Hand-listoftheGeneraandSpeciesofBirds, vol. 1, p. 116(order).-HesperornithomorphiHay,1930, Publ.CarnegieInstn. Washington, no. 390, vol. 2, p. 277(order).OdontognathaeWetmore, 1930, Proc. U.S.nat. Mus., vol. 76, art. 24, p. 1(superorderfor HesperornithiformesandIchthyornithiformes).FamilyfHESPERORNITHIDAEMarshHesperorn/daeMarsh, 1872, Amer. Jour. Sci., ser. 3, vol. 3, p. 363(typeHesper ornis ?-.1arsh).-Hesperornithidae Marsh, 1876, Amer. Jour. Sci.,SCI.3, vol.II,p. 509.-He,perornithoidea Shufeldt, Amcr. Natural., vol. 37, no. 433, p.59(superfamily) .GenustHesperomisMarshHe.perornisMarsh, 1872, Amer. Jour. Sci., ser. 3, vol. 3, p. 57(nomennudum);p. 360(typeby monotypyH esperorni. regaltsMarsh).LestornisMarsh, 1876, Amer. Jour. Sci., ser..3,vol. 11, p.509(typcbymono typyLestornis cra,s,ripes Marsh).HargeriaLucas, 1903, Proc. U.S.nat. Mus., vol. 26, p. 552(typebymonotypyH esperornis gracilisM"rsh).1.H esperomis regalisMarsh IIe"jfU?rnrnis regalis 1872, Amer. Jour. Sci., ser. 3, vol. 3, p. 57(nomennudum);p. 360(lectotypefrom 20 miles castofWallace,partialskeleton, YalePeabodyMus. no. 1200, designatedbyMarsh,1880).UPPER CRETACEOUS, CONIACIAN(Niobraraformation, Smoky Hill chalkmember).KANSAS:Logan County: southbankof Smoky Hill River,20miles cast of Wallace (Marsh, 1872, I.e.); Smoky Hill River,12miles east of Wallace (Marsh, 1880, Odontornithes, p. 195);TwoMile Creek (Wetmore, 1940, Smiths. Misc. Coil., vol.99,nO.4, p.3).2.Hesperornis crassipes (Marsh)Lestornis crass/pes\{arsh, 1876, Amer. Jour. Sci., vol. 11, p. 509(typefroUl western Kansas, incomplete postcranial skeleton, Yale Peahody:\1us. no.1474).

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192BULLETIN FLORIDA STATE MUSEUMVol. 7UPPERCllETACEOUS,CONIACIAN(Niobraraformation, Smoky Hill chalkmember).KANSAS:probablyfrom Smoky Hill RiverinWallaceCounty, asthetypewas collectedbyG.P.Cooperin April1876.3.H esperornis gradlis Marsh Elesperorrtis gt'adlisMc.rsh,1876.Amer. lour. Sci.,!l'{'r.3,vo2. 11, p .510left Yale Peaoody Mus. no. 1478). UPPERCRETACEOUS,CONIACIAN(Niobraraformation, Smoky Hill chalkmember). KANSAS: Smoky Hill River(probablyinWallaceCounty,asthetypewas collectedbyG.P.Cooper in April1876).Genus IConionds MarshConiomisMarsh, 18fl:3 (Jan.),Amer. Jour. Sci., scr. 3, vol. 4.'), no. 265, p. 82.(typebymonotypyC tmiarnis alttlsMarsh).4.Coniornis altusMarsh Goniornis altus1893 (Jan.),Amer. Jour. St'i.,Ser.:'),vol. 45,lID.26,5, p. R2,fig.1-3(typedistal half of righttibiol.""s,Yale Peabolly Mus. no.515). He81Jerorni8montanaShufeldt, 1915(June),Auk, voL 32, no.3,p. 298,pI. 18, fig. 4, 6, 8, 10, 12(type23dvertehra, U.S.Nat. t\Ius. no.8199).'UPPERCRETACEOUS,CAMPANIAN(upperpartofClaggettformation).MONTANA:FergusCounty: 1 mile abovemouthofDogCreek,nearmouth ofJudithRiver. I'rhe two names are based on elements ofcomparable fromthe same hori ;lOll andlucality.ThesuppositionthatthetypelucalityofC. altwi lay nearthe base of the JudithRiver formationratherthanintheunderlyingma rineClaggettfOml(ltionresulted inpartfroUlearly usage oftheterm "Judith River beos" ina general sense toincludeallthelater-named Cretaceousfonnatiuns inthe area.stated thatthetype WAS collected with marine fossils,

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1963 BRODKORB:CATALOCUEOFFOSSILBIRDSSubclassORNITHURAEHaeckel193OrnithuraeHaeckel, 1866, Ccnerelle der Organisrnen, vol.2,p.139 (SubkIasse).Odoutomithe.Marsb, 1873, Amer. Jour. Sci., ser. 3, vol.,5,pp. 161, 162(sobclass; type Ichthyomis Marsb).OdolltormaeStcjoeger, 1885, Stand. nat. Hist., vol. 4, p. 28 (sub-dassforIchthyomis).-OdnntotormaeMenzbier, 1887, Bull. Soc. Natural.]\.108('ou,no.2,p.63(Unterclasse, ex orderOdontutormaeMarsh). Neornithes Gadow, 1893, Bronn Klass. Ordn., Vogel, pt.2, pp. 90, 299 (UntercIasse).1InfraclassDROMAEOGNATHAEHuxley2DromaeugnathaeHuxley, 1867, Pro<:. zool. Soc. London, p. 456(suborder;notbasedongenericname).OrderTINAMIFORMES(Huxley)TinamomorphaeHuxley, 1872, Manual AnatomyVertebratedAnimals, p.2,34(suborder?; type Ti,UZtrlUS Hermann). Crypturi SdaterandSalvin, 1872, Nomenclator Aviulll Neotropicalium, p. 152 (type CrypturusIlliger, a junior synonymofT ina11HlS HermanIl).FamilyTINAMIDAEGrayCrypturidaeBonaparte, 1831, Saggio diunaDistribuzione Metodica degli Animali Vertebrati, p.,53(type Crypttlru. llIiger, a junior synonymof Tinamtts Hermann).TinomidaeG. R. Gray, 1840, ListCencraBirds, p.63(typeTinamusHermann).Gentls RoveretoTinami.ornisRovereto, 1\)14, An. Mus. nae. Buenos Aires, vol. 2.5, p. 161(type Tirwmisomis intermediusRovereto, designatedby Richmond, 1927, Proc. U.S.nat.Mus., vol. 70, no. 2664,p. 3.5; 1'inamisornis pa-rvulus Rovereto was desii(nated byBrodkorb, 1961, Auk, vol. 78, p. 257,in oversight of Richmond's action). Rnveretnrnis Brodkorb, 1961, Auk, vol. 78, no. 2, p. '257 (typeby original designation Tinamis01'nis intermediusRovercto).1.Tinamisornis intermediusRovereto Tinamisornis intermediusRovereto, 1914, An. Mus.nacoBuenos Aires, vol. 25,pp. 161, 16'2, pI. 2.5, fig. 2-2e only(lectotypefrom Monte Hermosa, left humerus, Buenos Aires Mus., designatedbyBrodkorb,1961).UPPERPLIOCENE(MonteHermoso formation).ARGENTINA:ProvoBuenos Aires: Monte Hermoso.JAbout36other subclass nameshavebeen proposed for various groups of living birds.llNew rank.

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194BULLETINFLORlDASTATEMUSEUMVol. 7GenustCayetanornisBrodkorb'2.Cayetanornis parvulus(Rovereto)Tinamisomis pamulusRovereto, An. Mus.nacoBuenos Aires, vol.25,pp. 161162, pI. 25, fig. 3-3c(lectotype righthumems,Buenos Aires Mus., desig nated by Brodkorb,1961).UPPERPLIOCENE(MonteHermusoformation).ARGENTINA:Pro'll.BuenosAires:MonteHermoso.GenustQuerandiornisRusconiQuerandiornisRusconi, 1958, Rev. r.,,1us. Hist. nat. Mendoza, vol. 11, nos. 1-4, p. 157(typebymonotypyQuerandiornis ramaniRusconi).3.Querandiornis romaniRusconiQuerandiornis ramaniRusconi, 19.58, Rev. Mus. Rist. nat. Mendoza, vol. 11,nos. 1-4, p. 157.UPPERPLIOCENE(MonteHermosaformation).ARGENTINA:Pro'll.BuenosAires:MunteHermoso.GenusNothuraWaglerNothuraWagler, 1827, Systema avium, vol.I,folio 19(typeTinamus boraquiraSpix).4.Nothura palttdosaMerceratNothura paludosaMercemt, 1897, An. Soc. cien. argentina, vol.43,p.239(typefemur,LaPlataMus.).UPPERPLEISTOCENE(Pampasformation).ARGENTINA:Pro'll.BuenosAires: Arrecifes.NeospeeiesofTinamidaefromPleistocenesites:1. Tinamus ma;or(Cmelin). BRAZIL: Minas Gcraes: Lapa da Escrivanianear Lagoa Santa(0.Winge, 1887, E Mus. Lund., vol. 1, no. 2, p.16).2.Crypturellu. obsoletus(Temminck). BRAZiL: Minas Geraes:LapadoBahu, Lapa do Capao Sccco, LapaciaEscrivania, Lapa do Marinho, and LapadoTaquaral(Winge, op. cit., p.16).3.Crypturellus noctivagus(Wied). BRAZIL: Lapa da Escrivania (Winge, op. cit., p.16).4.Crypturellus parvirostris(Wagler).BRAZIL:LapadaEscrivania (Winge, Op, cit., p, 17).5.Crypture!!u", tataupa(Temminck). LapadaEscrivania, LapadoMarinho,andLapadoCapao Seeco (Winge, op. cit., p.16).INewgenus.TypcTinamisomis parvulusRovereto. For characters see Brodkorb (1961, Auk, vol. 78, p.257).Namedfor Cayelano Rovereto.

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS '195 6.Rhynchotus rufescens (Temminck).BRAZIL:LapadaEscrivania and LapadaLagoado Sumidouro(Winge, up. cit.,p.18).7.Nothoprocta cinerascens(Burmeister).ARGENTINA:Buenos Aires: Lujan (Ameghino, 1891, Rev. argentina Hist. nat., vol.1,p.446).8.Nothura macu/usa(Temminck). BRAZIL: Lapa da EscrivaniaandLapadaLagoadoSumidouro (Winf:e, op. cit., p. 17). ARGENTINA:Lujan (Amcghino,op. cit., p. 446).9.Nothuraminor (Spix).BRAZIL:Lapa da Escrivania (Winge, op. cit.,p.17).10.Taoniscus nanus(Tcmminck).BRAZIL:Lapa da Escrivania (Winge, ap. cit., p. 17).

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196BULLETINFLORIDASTATEMUSEUMInfraclassRATITAEMerrem'Vol. 7Aves RatitaeMerrem, 1813, Abh. Akad.Wi".Berlin, p. 259.PalaeagnathaePycraft, Trans. zool. Soc. London, vol.1.5,p.149.-PalaegnathaeWetmore, 1930, Proc.U.S.nat. Mus., vol. 76, art. 24, p. 2(superorder).OrderSTRUTHIONIFORMES(Latham)StruthionesLatham,1790, Index pp.xv,662(typeStruthioLinnaeus) .-StruthioniformesFiirbringer, 1888, U nter.sllch. 11orph. Syst.Vogel. vol. 2, pp. 1540, 1565(suhordo).FamilytELEUTHERORNITHIDAEWetmoreEleutherornithidaeWetmore, 1951 (Nov.1),Smithsonian misc. Coli., vol. 117, no. 4, pp. 3, 14(typeEleutherornisSehauh).GenustElelltherornisSchau hEleutherornisSchaub, 1940, Eclogae geol. Helvetiae, vol. 33, no. 2, p. 283(typeby monotypyEleutherornis helvetieusSchaub).1.Elelltherornis helveticusSchaubEleutherornis Sebaub, 1940, Eclogae geol. Helvetiae, vol. 33, no. 2, p. 283,fig.1-4(typepelvis from Bohnerz, Basel Mus. no.Eh781).LOWERMIDDLEEOCENE(Egerkingengamma).SWITZERLAND:13ohnerz.FamilySTIlUTHIONIDAEVigorsStruthianid.aeVigors, 1825, Trans. Linn. Soc. London, vol. 14, pp. 478, 480(typeStruthioLinnaens).GenusStruthioLinnaeusStruthioLinnaeus, 1758, Syst. Nat., ed. 10,vol.1, p. 155(typeStruthio camelusLinnaells,Recent).StruthiolithusBrandt, 1873, Bull. Acad. Sci. St. Petersburg, vol. 18, p. 158 (typebymonotypyStruthiolithus chersonensisBrandt).MegaloseelornisLydekker, 1879, Rec. geol. Surv. India,vol.12, p. 53(typebymonotypyMegaloseelornis sivalensisLydckkcr). Palaeostruthio Burchak-Abramovich, 1953, Akad. Nauk Azerbaidzhanskoi S.S.R., p.81(typebyoriginal desigrnttionPalaeostruthio sternatusBurcbak-Abramovich).PachystruthioKretzoi, 1953, Acta geologica,vol.2, pp. 231-242 (subgenus; typebymonolypyStTuthio (Pachystruthio) pannonicusKretzoi).lNewrank.vVhether theratites form anaturalgroupis still a far fromsettledquestion,anditis likelytoremainunanswereduntil their originscanhetracedin the fossil record. Both ratitcs and carinates could have arisen from a tinamoulike stock.

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1963 BRODKORB:CATALOCUEOFFOSSIL BIRDS1.Struthio asiaticus Milne-Edwards197Struthio palaeindicaFalconer, 1868, Palaeontological Memoirs, vol.1,pp. xxi, 554(nomennudum; 15 clements from Siwalik Hills listed, withBrit. Mus.cat.nos.,from nnpnblished PlateR).Struthio asiaticusMilne-Edwards,187],Oiseanx Foss. France, vol. 2, sheet 74,p.587 (briefdescription; type tarsometatarsus from Siwalik Hills, Brit.Mus.).Megaloscelomis sivalensi. Lydekker, 1879, Rce. geol. Surv. India, vol. 12, p. 56,part(types from Siwalik Hills, tibiotarsusandfibula, Indian Mus., Calcutta, casts in Brit.Mus.).Struthio indie... Bidwel!, 1904, Ibis, scr. 9, vol. 4, p. 760,fig.7(typesfrom Nullas, 7 eggshell fragments, Brit. Mus., Tring Mus., andCalcuttaMus.).LOWERPLIOCENE(Siwalik series).INDIA:UNITED PROVINCES: Siwa lik Hills,probablynearHardwar(Falconer,1868); Nullas on Ken River inBandadistrict (Bidwell, 1904).2.Struthio chersonensis(Brandt)1 Struthiolithus chersunensis Braudt, 1873, Bull. Acad. Sci. St. Petersburg, vol. 18, p. 1.58 (typeegg from Malinovka destroyed, cast said tobein St. Peters burg Acad. Sci.).Struthio karatheodorisForsyth Major, 1888,C. R. Acad. Sci. Paris, vol. 107, p. 1178(typefrom Samos, femur, Barbey colI., Valleyres).Struthio Alexejew, 1916, Animaux fossiles du villageNovoElisavetovka, p. 388,fig.55-56 (types from Novo-Elisavetovka, distal portions of 3 tarsometatarsi, Novorossyisk Univ. nos. 1559-1.561). Struthio braehydaetylusBurehak-Abramovich, 1939, Priroda, no. 5, p. 95; re described 1949, Doklady Akad. Nauk SSSR, vol. 67, no.1,p. 14,fig.1-4 (type from Grcbcniki, skeleton lackingsternulll,wings, and sacrum, Acad.Sci. Moscow). Palaeostruthio.""ernatu", Burchak-Abramovich, 1953, Akad. Nauk Azerbaidzhan skoi SSSR, p. 81, pI. 18,fig.1(typefrom Grebeniki, sternum, Acad. Sci. Moscow no.408/367).LOWERPLIOCENE(Pannonian).GREECE:SamosIsland(ForsythMajor, 1888).UKRAINE:MalinovkancarKherson(Brandt,1873); KuyalnikestuarynearOdessa, Vyshiva (Novo-Pokrovsk),andNovo Elisavetovka (Alexejew, 1916); Grebeniki (Burchak-Abramovieh, 1939).KAZAKSTAN:Pavlodar(Howard,1939, Fortschr. PaLvoL2, p. 313).Probablyreferablehereare specificallyundeterminedrec ords fromMaraghehinIranandfrom Garet-el-Muluk,Egypt(Lambrecht, 1933,Handb.Palaeorn.,pp.103-104).lThat more than one speciesofostrich existed during the early Pliocene hasnotbeenproved.

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198BULLETINFLORIDASTATE MUSEUM3.Struthio wimaniLoweVol. 7Strothio wimaniLowe, 1931, Pal. sinica, ser. C, vol. 6, fa,c. 4, p. 18, pI. 1,fig.1; pI. 2, fig. 2; pI.3;pI.4,fig.1,4(typefrom Locality 30,T'aiChiaKOH,pelvis, Palaeont. Mus.,Upsala).Strothio mongolicusLowe, 1931, Pal. siniea, ser. C, vol. 6, fase. 4, p. 34, pI. 4,fig.5(typeseggshell fragments, Upsala Mus.; locality of type not stated,butfiguredspecimenis from GlanChorea).LOWERPLIOCENE(Hipparionredclays).CHINA:ProvoShansi:TaiChiaKou in Pao-tc Hsicn (Lowe, 1931); Hsiang-ning Hsien(Lambrecht, 1933, Handb. Palaeorn., p. 106).ProvoKansu: Ching YangFu(Lambrecht,1933).LOWERPLIOCENE(Ertemtestage).MONGOLIA:Ertemtc,alanChorea, Tjclgol-Tabool,andDoshen (Lowe, 1931); Choei Tong K'eou, Sjara Ossa Gol, Hong-Tcheng, Shabarakh Ussu, Djadochta,andHungKurek(Lambrecht,1933).4. Struthio pannonicusKretzoiStrothio (Pachystrothio) pannonicusKretzoi, 1953, Acta geologica, vol. 2, pp. 231-242, pI. 1-3(typefrom Kisling, right pedal phalanx 1 of digitIll).LOWERPLEISTOCENE(upperCalabrian).HUNGARY:Transdanubia: Kishlng. 5.Struthio oldawayiLoweStruthio oldawayiLowe, 1933, Ihis, ser. 13, vol. 3,110.4, pp. 652, 654(typefrom Oldaway, pelvisandsacrum, apparentlyinBrit.Mus.).LOWERPLEISTOCENE(Olduvaiseries,bed1).TANGANYIKA:Olduvai(Oldaway).6.Struthio anderssoniLoweStruthio anderssoniLowe, 19:31, Pal. sinica, ser. C, vol. 6, fasL:. 4,p.26,fig.2(typecomplete eggshell fromllonan,Brit. Mus. no.A.130R;femora later recordedbyBouleandTeilharddeChardin, 1928; Howard,1939).UPPERPLEISTOCENE(Samnenseries,Fenhostage).CHINA:ProvoHopeh: YaoKuanChuang; Ching Hsing coal mine; K'ou-An; Chou k'ou-tien.ProvoShantung: Sha-vVa-Tsun.ProvoShansi: Tang-T'aiChuang;Liang-ChiaTan.ProvoHonan: K'iho onWeiRiver; Wu-An Hsien; Chengchow Hsien;Han''Vanh Cheng in110YinHsien; SSu shui Hsien;ChaoKon in Kung Hsien; Ts'ai Chia Chuang, Hsia JunYii,andTungHuangNii Yiian in Hsi An Honein;FcngMing P'o, KuoYiiKou,andYang Shao Tsun in Mien Chih Hsien(Lambrecht,1933,Handb.Palaeom., p.104).

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS 199Neospecies of Struthionidae from Pleistocene and'prehistoricsites:1.StTuthio camelus Linnaeus. ALCERIA: Hassi-el RatmaiaintheGrandErgand Mouilah-Maatalah (Lamhrecht, 1933, Handb. Palaeom.,p.108). AnAnIA: Tuwairifa, Ain Sala, UmOlal Qurun, UmOlTina,Qa'amiyat, Abu Sabbau. Summan Mahadir, and BaniMaaridh(Lowe, 1933, Ihis, p.658).BURYATMONGOL RI!:PUHLIC: Troitzkosavask, Sclenga River, and Chorenchoi (Lambrecht,1933, p. 107).OUTERMONCOLiA:Shabarakh Ussn and Djadochta(Lambrecht,1933, p. 108).

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200BULLETINFLORIDASTATEMUSEUMOrderHHEIFORMES(Forbes)Vol. 7RheaeForbes, 1884(Jan.),Ibis,ser.5, vol. 2, no. 5. p. 119(typeRheaBrisson).RheiformesFiirbringer, 1888, Untersueh. Morph. Syst. Vogel, vol. 2,pp.1540, 1565(subordo).FamilyfOPISTHODACTYLIDAEAmegbino Opi8thodactylidae Ameghino, 1895, Bol. Inst. geog. argentino, vol. 15, p. 81 (typeOpisthodactylus Ameghino). GenusfOpisthodactylusAmeghino1OpisthodactylusAmeghino, 1891(Dec.),Rev. argentina Hist. nat., vol.1,p. 453(typebymOllotypy01'isthodactylus patagonieusAmeghino).1.Opisthodactylus patagonicusAmeghinoOpisthodaetylus patagonicusAmeghino, 1891(Dec.),Rev. argentinallist.nat., vol. 1, p. 453 (type distal portion of tarsometatarsus, Brit.Mus.).LOWEREOCENE(Casamayorformation).ARGENTINA:southernPatagonia.Family Rm:IDAE (Bonaparte)RheinaeBonaparte, 1853, C.R.Aead. Sci. Paris, vol. 37, no. 18, p. 646 (sons famille;typeRhea Brisson). Genus fIIeterorheaHoveretoHeterorheaRovereto, 1914, An. Mus. nae. Buenos Aires, vol. 25, p. 160(typebymonotypy Heterorhea dabbeneiRovereto).1.IIeterorhea dabbeneiRoveretoHeterorhea dabbeneiRovereto, 1914, An. Mus. nae. Buenos Aires, vol. 25, p. 160, pI. 25, fig. 1(typetarsometatarsos, Buenos AiresMus.).UPPERPLIOCENE(MonteHermosoformation).ARGENTINA:Provo Buenos Aires:MonteHermoso. GenusRheaBrissonRheaBrisson, 1760, Ornithologia, vol.1,p.46(typeStruthio americanusLinnacus,Recent).lTransfcrrcd from the Phororhacoidea to the rheasbyPatterson and Kraglievieh(1960,Publ. Mus. municipal Cien. nat.ytrad.Mar del Plata, vol. 1, no.1,p.11)withoutanysupporting evidence.Thecharactersofthedistalendof the tarsometatarsus mentionedbyAmeghino, namely the concavityofthe plantar sur face above the trochleae and the facet fur a hind toe, would preclude its reference to the Rheidae as currently understood.

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GenusPterocnemiaGrayPteroenemiaGray, 1871, Hand-list Birds Brit. Mus., vol. 3, p. 2(typeRhea pennatad'Orbigny,Recent).2.Rhea anchorenensisC.AmeghinoandRusconiRhea americana anchorenenseC. Amcghino and Rusconi, 1932, An.Soc.den. argentina, vol. 114, p. 38,fig.1(typedistal half of tarsometatarsus, Museo La Plata).LOWERPLEISTOCENE(basalEnsefiadan).ARGENTINA:ProvoBuenos Aires: Anchorena.3.Pterocnemia tossilis(Ameghino) Rhea fossilisF.Ameghino, 1882, Catalogue specialdela Section anthropologique et paIeontologiquede laRepublicArgentine, Exposition universelle de 1878, Group second, Classe p.42(typefrom Olivera, incomplete skeleton, MuseoLaPlata nos. 200-228). Rhea pampeanaMoreno and 1891, An. rvIus. La Plata, Pal. arg., vol.1,pp. 27, 70, pI. Hl, fig.1, 3-10, 13; pI. 20,fig.1-4; pI. 21,fig.1-4(sametypeasR.fossilisAmeghino).[?lRhea nanaLydekker, 1894, Knowledge(London),vol. 17, p. 265(typea runt egg of unknown ageandlocality). 201 BRODKORB: CATALOGUEOFFOSSIL BIRDS 1963UPPERPLEISTOCENE(Pampasformation,upperlevel).ARGENTINA:ProvoBuenos Aires: Olivera. Neospecies ofRheidaefrom Pleistocene sites:1.Rhea americana(Linnaeus).BRAZIL::MinasCeraes: LapadeAnna Felicia, I .. apa da Anta no.1,and Lapa da Escrivania no. 1(0.Winge, 1887, E Mus. Lund., vol. 1, no. 2, p.18).ARGENTINA:ProvoBuenos Aires: Lujan (Ameghino, 1891, Rev.argoHist. nat., vol. 1, p.448);Mardel Plata(RheafossilisMorenoandMercerat, 1891; see below). Fossil synonyms of this species (fide Ameghino, 1891) include:RheafossilisMoreno aod Mercerat, 1891,An.Mus.LaPlata, Pal. arg., vol. 1, pp. 28, 71, pl. 19,fig.2, 11, 16; pI. 20,fig.2; pI. 21,fig.6 (types fromMardclPlata, fragmentary right tibiotarsus, fragmentary rightand left tarsometatarsi,Mus.LaPlata nos. 229-2,33);Rhea 8ubpampeanaMorenoandMercerat, 1891, op. cit., pp. 27, 70, pI. 20,fig.22(typerightouter digital trochlea, Mus.LaPlatano.199, said tobefrom Lagnna deVitel,butboth localityand age erroneous according toAmeghino).

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202BULLETINFLORIDASTATE MUSEUM Vol. 7OrderCASU ARIIFO RMES(Sclater)Casuarii ScI"ter, 1880, Ibis, scr. 4, vol. 4, no. 16, Pl'. 410,4Il(order; type Ca.marius Linn.eus).-Casuariiformes Fiirbringer, 1888, Untersuch. Morph. Syst. Vogel, vol. 2, Pl'. 1541, 1565(subordo).FamilyCASUARlIDAEKaup Casuariidae Kaup, 1847, fide Gray, 1870, Hand-list Gen. Sp. Birds, pt. 3, p. 2(typeC asuariusLinnaeus).Genus CasuariusBrisson Ca,suarius1760, Omithologia,voL1, p. 46(type Struthio cusuariusLinnaeus,Recent).1.Casuarius lydekkeriRothschildCasuariu. lydekkeri Rothschild,19I1,Verh.V.internat. ornith. Kongr. Berlin 1910, Pl'. 151, 162(typefrom Wellington Valley, distalpartofrighttibio tarsus, Australian Mus. no.MF1268; cast Brit. Mus. no. A.158 == B.I0394; see A.H. Miller, 1962, Rec. Austral. Mus., vol. 25, p.235).UPPERPLEISTOCENE(cavedeposit).NEWSOUTHWALES:WellingtonValley.FamilyDROMICEIIDAERichmondDramaiinae Gray, 1870, Hand-list Gen. Sp. Birds, pt. 3, Pl'. v, 2 (subfamily;typeDromaiu8Vieillot, 1818, a junior synonymofDromiceiusVieillot).DramaeidaeA.Newton, 1896, Dictionary of Birds, p. 213(type"Dramaeu." Vieillot). Dramiceiidae Richmond, 1908, Proc. U.S.nat. Mus., val. 35, no. 1656, pp. 59S, 6,51(typeDramiceiu. Vicillot).GenusDromiceiusVieillotDrumiceiu. Vicillot, 1816, Analyse nouv. om. ,,!em., p .54 (typeCasuarius novaehallandiae Latham, Recent).1.Dromiceius patricius(DeVis)Dromaius patriciusDeVis, 1888, Proc. Linn. Soc. N.S. Wales, ser. 2, vol. 3,1': 1290, pI. 36,fig.13a-c (types from King's Creek, proximal anddistal ends of right tibiotarsus, left coracoid, prohably in Queensland Mus,). UPPER PLEISTOCENE(Chinchillabeds).QUEENSLAND:King's Creek.UPPERPLEISTOCENE(Katipirisands,Malkunifauna).SOUTHAus-TRALIA:WurdulumankulanearLakeEyre(DcVis, 1906, Ann. QueenslandMus., no.6,p.25).UPPERPLEISTOCENE(cavedeposits).NEWSOUTHWALES:WellingtonValley(Lydekkcr,1891, Cat. Foss. Birds Brit. Mus.,p.352).

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDS 2032.Dromiceius gracilipes(DeVis) Dromaius graci/ipesDeVis, 1892,Proc. binn. Soc.N.S.Wales, ser. 2, vol. 6, p. 445, pI. 23,fig.7(typetarsometatarsus).UPPERPLEISTOCENE(Chinchillabeds).QUEENSLAND.3.Dromiceius minor(Spencer) Dromaeus minorSpencer, 1906,VidoriaNatnralist, vol. 23, p. 140(typepartial skeleton).Dromaeus bassiLegge, 1907,Emu,vol. 6, p. 119.Dromiceius spenceriMathews, 1912, Novit. zool.(London),vol. 18, p. 176 footnote.QUATERNARY.AUSTRALIA:KingIslandinBass Strait. Recently extinct species ofDromiceiidaefromthePleistocene:1.Dromiceius diemenianus(Jennings). AU8cl'RALIA: Kangaroo Island(Lambrecht, 1933,Handb.Palaeorn., p.114).FamilyfDROMORNITHlDAEFiirbringerDromornithidaeFiirbringer, 1888, Untersuch. Morph. Syst. Vogel, vol. 2,pp.1435, 1565(typeDromornisOwen).GenusfDromornisOwenDromornisOwen, 1872,Proc. zool. Soc. London, p. 682(typebymonotypy Dromornis australis Owen).1.Drornornis australisOwenDromornis australisOwen, 1872, Proc. zool. Soc. London, p. 682 (typefromPeak Downs, right femur, Sydney Mus., cast in Brit. Mus.).Dromaius amtrulis Woods, 1883, Proc. Linn. Soc. N.S.Wales, vol. 7, p. 387 (types from PenoIa, 2 tibiae, 2 tarsomctatarsi, colI. Rev.].E.Tenison\Voods, perhaps now in Pcno]a Institute).[?IDinornis queeuslundiae DcVis, 1884, Proc. Roy. Soc. Queensland, vol.1,p. 23, pI. 3-4(typefrom King's Creek, proximalendof femur, Queensland Mus.).UPPERPLEISTOCENE(Chinchillabeds).QUEENSLAND:PeakDowns(Owen,1872);King'sCreek?(DeVis,1884).UPPERPLEISTOCENE. SOUlH AUSTRALIA:PenolainGambierRange(Woods,1883).GenusfGenyornisStirlingandZietz Genyornis Stirling andZietz, 1896, Tmns. Roy. Soc.S.Australia, vol. 20, p. 182(typebymonotypyGenyornis newtoniStirlingandZietz).

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204BULLETINFLORIDA STATE 2.Genyornis newtoniStirlingandZietzVol.7Genyornis newtoniStirling and Zietz, 1896, Trans. Roy. Soc.S.Australia, vol.20, p. 182(typefrom Lake Callabonna, skeleton, South Australian Mus. in Adelaide).UPPERPLEISTOCENE.SOUTHAUSTRALIA:Lake Callabonna (StirlingandZietz, 1896); Normanville, Baldina Creek near Burra, Parroo River,andMount Gambier (Lambrecht,1933,Handb. Palaeorn., p. 117).NEWSOUTHWALES:GorecandCanadian Gold Lead, near Mudgee (Lambrecht, 1933).

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDSOrdertAEPYORNITHIFORMES(Newton)205AepyornithesA.Newton, 1884, Encyclop. brit., ed. 9, vol. 18, p.44(typeAepyornisGeoffroy).-AepiornithesStejneger, 1885, Stand. nat. Hist'l vul.4, p.47.-AepyornithiformesFiirbringcr, 1888, Untersucb. Morph. Syst. Vogel, vol. 2, pp. 1541, 1565(intennediaresubordo).-AepiornithiformesRidgway, 1901, Bull.U.S.nat. Mus., no. 50, pt. 1, p.9.Family tAEPYORNITHIDAE(Bonaparte)EpyornithinaeBonaparte, 1853, C.R.Acad. Sci. Paris, vol. 37, no. 18, p. 643(sous-famille; type "Epyornis" Geoffroy).-AepyornithidaeA. Newton, 1884,Encyclop. brit., ed. 9, vol. 18, p. 44.Subfamily tEREMOPEZTNAELambrechtEremopezinaeLambrecht, 1933,Handb.Palaeon., p. 216(typeEremopezusAndrews).Genus tEremopezusAndrewsEremopezusAndrews, 1904, Proc. zool. Soc. London, p. 163(typebymonotypyErernopezus eocaen-usAndrews).[?]PsammornisAndrews, 1911, Verh. V internaL ornith. Kongr. Berlin 1910, p.169(type by monotypyPsammornis rothschildiAndrews).1.Eremopezus eocaenusAndrews Eremopews eocaenus Andrews, 1904, Proc. zool. Soc. London, p. 163, pI. 5 (type from Birket-cl-Qurun, distalendof tibiotarsus, Brit.Mus.).[?JPsammornis rothschildiAndrews, 1911, Verh. V internat. ornith. Kongr. Berlin 1910, p. 169 (types from eastofTouggourt, eggshell fragments, Brit. Mus. and Tring Mus.).[?IPsammornis lybicusMoltoni, 1928, Ann. Mus. Storia nat. Giacomo Doria, vol. 52,p.399, fig. (types from south ofHatietel-Hueddaandsouth of Ciara huh, eggshell fragments).UPPEREOCENE(Fayumformation, Birket-el-Qurunstage).EGYPT:Fayum:northof Birket-eI-Qurun (Andrews, 1904).EOCENE?LYBI": 27 kilometerssouthofHatietel-Hucddaandsouth of Giarabub? (Moltoni, 1928).ALGEillA:12 miles east of Touggourt, Biskra, Ouargla,EIGolea,andTemacine(Andrews, 1911).ARABIA:Shuqquat aI Khalfat(Lowe,1933, Ibis,p,656). Subfamily tAEPYORNITHINAE(Bonaparte)GenustStrorneriaLambrechtStromeriaLambrecht, 1929, Abh. bayer. Akad. Wiss., Math.-Naturw. Abt.,F.4,p.1(typebymonolypyStromeria fajumensis Lambrecht).

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206BULLETINFLORIDASTATEMUSEUMVol. 72.Stromeria fajumemisLambrechtStromeria faiumensis Lambrecht, 1929, Abh. hayer. Akad. Wiss., Math.-Naturw. Abt., F. 4, p. 1, pI. 2(typefrom Dimeh, distalthirdofrighttarsometatarsus,Munich Mus.).Stromeria fayumensisLambrecht, 1933,Handb.Palaeorn., p. 193.LOWEROLIGOCENE(Fayumformation,Qatramstage).EGYPT:Fayum:northofDimeh.GenustMullerornisMilne-EdwardsandGrandidierMullerornisMilne-Edwards amI Grandidier, 1894, C.R.Acad. Sci. Paris, vol. 118, p.12,5(typeMnllerornis betsi/ei,designatedbyRichmond, 1902, Proc. U.S.Nat. Mus., vol. 24, p.697).FlacourtiaAndrews, 1895, Novit. zool.(London),vol. 2, p. 23(typeMnllerornis rudisMilne-EdwardsandGrandidier).3.Mullerornis betsileiMilne-EdwardsandGrandidierMnllerornis betsileiMilne-EdwardsandGrandidier, 1894, C.R.Aead. SeLParis, vol. 118,p.125 (types tibia,tarsometatarsllS,ParisMus.).QUATERNARY.MADAGASCAR:Antsirab6,incenterof island.4.Mullerornis agilisMilne-EdwardsandGrandidier "gillsMilne-EdwardsandCrandidier, 1894,C.R.Aead. Sci. Paris, vol. 118, p. 125(typetibia, ParisMus.).QUATERNARY.MADAGASCAR:southwestcoastnearMourOlmdava.5.Mullerornis rudis Milne-EdwardsandGrandidierMullerornis rudisMilne-EdwardsandCrandidier, 1894,C.R.Acad. Sci. Paris, vol. 118, pI.26(types tibia, tarsometatarsus, ParisMus.).QUATERNARY.MADAGASCAR:westcoasthetween B6lo andMouroundava.GenustAepyornisGeoffroyAepyornis1.Geoffroy-Saint-lIilaire, 1851(afterJan.27),C. R. Acad. Sci. Paris, vol. 32, no. 4, p. 104(typebymonotypyAepyornis maximnsGeoffroy).AepiomisGeoffroy, 1851, Rev. Mag. Zool.(Paris),ser. 2, vol. 3, p. 52(emendation).Epiornis Miiller and Baldamus, 1851, Naumannia,vo1.1, no. 4,p.48 (emendation).EpyomisBonaparte, 1854, Ann. Sci. nat.(Paris),ser. 4, vol. 1, fase. 3, p. 139 (emendation; Epyornis used as a common namebyGeoffroy, 1851, C. R.,I.e.) .

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS2076.Aepyornis maxirnusGeoffroyAepyornis maximusL Geoffroy-Saint-Hilaire, 1851(afterJan.27),C. R.Aead. Sci. Paris, vol. 32, no. 4,p.104 (types from Masikoru,eggand lower end of left metatarsus, Paris Mus.).Aepyornis modestusMilne-EdwardsandGrandidier, 1869, Ann. Sci. nat.(Paris),ser.5,vol. 7, p. 314(typefrom Amholisatra, ParisMus.).Aepyornis titanAndrews, (Jan.12),Geol. no. 3.';5 =n.s., decade 4, vol.1,no.1,p. 18(typefromltampulu Ve, left tibiotarsus, Brit.Mus.).Aepyornis ingen.Miloe-EdwardsandGrandidier, 1894(afterJan.13),C. R.Acad. Sci. Paris, vol. 118, no. 3,p.124 (types from west coast between Bela and Mouroundava,femllr,tibia, Paris Muti.). QUATERNARY.MADAGASCAR:AmbolisatraorAmboulitsate(MilneEdwardsandGrandidier,1869);YlasikoroorMachicora(MilneEdwardsandGrandidier,1894);Mouroundava,between Belo andMouroundava,andltampulu Ve (Andrews,1894);Lamboharana(Lambrecht,1933,Handh.Palacorn., p.198).7.Aepyornis mediusMilne-EdwardsandGrandidierAepyornis mediu.sMilne-Ed\\!<.uds andGrandidier, 1866,RecherchessurIafanne ornithologiqll etp.inte des nes Mascareigneset oe1'v1adugascar, p. 97, note 2(type,Paris MlIs.).Aepyornis grandidieriRowley, 1867, Proe. zooI. Soc. London, p. (typefrom Cape Sainte Marie, eggshell fragment, colI. AlfredGrandidier).AepyorniscursorMilne-Edwards and Grandidier, 1894 (afterJan. V5), C.R.Acad. Sci. Paris, vol.118,nu. 3,p.124(typetarsomelatarsus, ParisMus., localitynotstaLed).Aepyornis lefl/lls Milne-EdwardsandGrandirlier, (afterJan.15),C.R.Acad. Sci. Paris, vol. 118, no. 3,p.124(typetarsometatarsm;,Paris locality notstated).QUATERNAHY. .\!IADAGASCAR: CapeSainte-Marie;probablybetween Bela andMouronndava(Milne-EdwardsandGrandidier).8. Aepyomis hildebrandli BmckhardtAepyornis hildehrundti Burckhardt, 1883, Pal. Abh., vol. 6, p. 127. pI. 13-16(typetarsometatarslls,BerlinMus.).Aepyornis mulleriMilne-Edwards and Grandidier, 1894(afterJan.15),C.R.Acad. Sci. Paris, vol. 118, no.3,p.124(typefromAntsirabe,nearlycompleteskeleton, ParisMus.).QUATERNARY.MADAGASCAR:Antsirahe.9.Aepyornis gracilisMonnier Ael'yornis gracilis Monnier, 1913, Ann. Pal. (Paris),vol. 8, p. 15, pI. 10(type femur, Paris :Mus.). QUATERNARY.MADAGASCAR.

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208BULLETINFLORIDASTATE Vol. 7OrdertDlNORNITHIFORMES(Gadow)ImmanesA.Newton,1884, Eneyc!op. brit., ed. 9, vol. 18, p. 44.DinornithesGadow, 1893, Bronn Klass. Ordn., Vogel, pt. 2, pp. 105, 299(typeDinornis Owen).-DinornithiformesRidgway, 1901, Bull. U.S.Nat. Mus., no. ,'50, pt. 1, p. 9.FamilytEMETDAE(Bonaparte)EmeinaeBonaparte, 18.54, Ann. Sci. uat.(Paris),vol.1,p. 48(typeEmeusReichenbach) .AnomalopterygidaeOliver, 1930,NewZealand Birds,p.28(typeAnomalopteryxReichenbach) .SubfamilytANOMALOPTERYGINAE(Oliver)AnomalapteryginaeArchey, 1941(May29),Bnll. Auckland Inst.andMus., no.1,pp. 11, 77 (slIb-family).GenustAnomalopterlJxReichenhachAnomaloptet'yxReichenhach, 1852, Avium Systema Naturale,p.xxx(typebymonotypy Dinornis didiformisOwen).GrayaBonaparte,18.56(afterNov.3),C. R. Aead. Sci. Paris, vol. 43, no. 18,p.841(typehypresent designationDinornis dromaeoidesOwen).AnomalomisHutton, 1897(June),Trans. N. Zealand Inst., vol. 29, p. .543 (newname forAnomalopteryxReichenbach).1.AnomalopterlJx anti'luusHuttonAnomalopteryx antiquflsHutton, 1892(May),Trans. N. Zealand Inst., vol. 24, p. 124 (lectotype tibia, Canterbury Mus., designatedbyArchey, 1941, p.29).UPPERMIOCENEorLOWERPLIOCENE.NEWZEALAND:SOUTIIISLAND:GlenitiValleynearTimaru.2.Anomalopteryx didiformis(Owen) Dinornis didiformisOwen, 1844 (June5),Trans. zool. Soc. London, vol. 3,pt. .3, p. 242, pI. 27,fig.3-6(typefrom Poverty Bay, metatarsus, Royal Col lege of Surgeons; cast Brit. Mus. no. 18595).Dinarnis dromaeoidesOwen, 1844(June5),Trans. zool. Soc. London, vol. 3, pt.:3,p. 2S3, pI. 22,fig.1-2; pI. 23,fig.1(typefrom Poverty Bay, femur,Roy. ColI. Surg.DO..16; cast Brit. no. ] Sr;98jcast CanterhuryMus.).Dinornis dromioide.r; Owen,1846 (July), Proc.zoot Soc. London,pf.14,pp.46, 47(emend"tionor lapsus). Dinornis PQrvus Owen, lRR3 (Jan.),Trans. zoo!. Soc. London, vol. 11, pt. 8,no.1,p. 233, pl. SI-.'58 (typefrom Pokororo, skeleton, Brit. Mus. no.A.3).Anomalapteryx fortisHutton, 1893(May),Trans.N.Zealand Inst., vol. 2.'5, p. 9 (lectotype from Glenmark, Canterhury MIlS., designated by Archey,1941).QUATERNARY.NEWZEALAND:NORTHISLAND:PovertyBay(Owen,1844);WaingongoroandTeRangatapu(Lydekker,1891,Cat.Foss,

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1963BRODKORB:CATALOGUEOFFOSSILBIRDS209Birds Brit. Mus.,p.666); Waipu, Akaitio, Karamu, Moawhango, Pa taua, Martinborough, Rotorua, Opito,TeAute,andLyall Bay (Lam brecht, 1933, Handb. Palaeorn., p. 143); Nuhaka, Awamarino,TeAnga, Mangaotaki, and Waikaremoana (Archey, 1941, Bull. Auckland lust. Mus., no.1,p. 137); Tukituki River, Hangatiki, Tahora, Pohue, Coonoor, Makirikiri, Mangaone, Kaiiwi, Wanganui, Lake Kaitoke, and Levin (Oliver, 1955,N.Zealand Birds, cd.2,p. 582); Gisborne, Whan garei, and Coromandel (Scarlett, 1957, Proc. N. Zealand ccol. Soc.,no.4,p.17).SOUTHISAND:Pokororo (Owen, 1883);40miles northofNelson, Waikouaiti, Ruamoa near Oamaru, and Otago (Lydekker,1891,pp. 278, 666); Glenmark (Hutton, 1893); Takaka Hill, Hamil ton Swamp, Waiau, Cheviot, Aorere River, and Kapua (Lambrecht,1933,p.144); Castle Rocks, Collingwood, and Mount Arthur (Archey, 1941); Shag River, Broken River, and Papatowai (Oliver, 1955).Although birds from the South Island average slightly smaller, the dif ferences are too slight to permit subspecific separation. GenustMegalapteryxHaastMegalapteryx Haast, 1886 (Dec.),Trans.zool. Soc.London,vol.12,pt.5,p.161(typebymonotypyMega/apteryx hectoriHaast).Palaeocasuarius Forbes,1892 (May),Trans. N. Zealand Inst., vol.24,p.189(nomennudum).-Forbes,189:3(July),Ibis, ser.6,vol.5,no. 19,p.450 (generic characters;includedspecies P. luzasti andP. velox,bothnominanudaatthispoint).-Rothschild, 1907 (Nov.12),ExtinctBirds, p.219(typebyoriginaldesignationPalaeocasuarius haastiForbes"). 3.Megalapteryx didinus (Owen)DinornisdidinusOwen,1883(Jan.),Trans. zool. Soc.London,vol. 11, pt.8,p.257,pI.59-61(typefromQueenstown,incompleteskeleton, Brit. Mus.nO.A.16).Megalapteryx heetoriHaast,1886(Dec.),Trans.zoo1.Soc.London,vol.12,pt. 5, p.161,pI.30(typefromTakaka,legbones, NelsonMus.).MegalapteryxtenuipesLydekker,1891(Apr.25),Cat. Fossil Birds Brit. Mus., p.2,51,fig.69a(typefromLakeWakatipll,righttibiotarsus, Brit. Mus. no.49990).Palaeocasuarius haastiForbes, 1893(July),Ibis, ser. 6, vol. 5, no. 19,p.451(nomennudum).-Rothschild,1907(Nov.12),ExtinctBirds,p.220(typefrom Manitoto,femur,LiverpoolMus.).PalaeocasuariusveloxForbes, 1893 (July),Ihis, ser.6,vol.5,no. 19,p.451(nomennudum).-Rothschild,1907(Nov.12),ExtinctBirds, p.220(typefrom Manitoto, femur, LiverpoolMus.).PalaeocasuariuselegamRothschild,1907(Nov.12),ExtinctBirds,p.220(typefrom Manitoto, femur, LiverpoolMus.).MegalapteryxhamiltoniRothschild,1907(Nov.12),ExtinctBirds,p.197(typefrom Waingongoro,leftfemur, Brit. Mus. no.32145).

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210BULLETINFLORIDASTATE MUSEUM Vol. 7QUATERNARY.NEWZEALAND:SOUTHISLAND:QueenstownnearLakeWakatipu(Owen,1883); Takaka(Haast,1886); Maniototo (Roths child, 1907); Buller River, Kapua,andNelson(Lambrecht,1933,Handb.Palaeorn., p. 142); Mount Arthur, Inangahua,OldMan Range,andAniseed Valley (Archey, 1941, Bull. Auckland Inst. Mus., no.1,pp. 31,33, 138); D'Urvil!c Island, Pokororo, Glenmark, Cromwell, Manuherikia, Kingston, Papatowai,andTeAnau (Oliver, 1955,N.Zealand Birds, ed.2,p. 583);InangahuaJunction (Scarlett, 1957; Proc.N.Zealand ecol. Soc., no.4,p. 17).NORTHISLAND:Waingongoro (Ly dekker, 1891, Cat. Foss. Birds Brit. Mus., p. 251; doubtful, fide Oliver); Makirikiri (Archey, 1941, p. 35).4.Megalapteryx benhamiArcheyMegalapteryx benhamiArchey. 1941(May29),Bull. Aucklaud Inst. Mus., no.1,pp. 35, 138(typefrom Mount Arthur, femur, AucklandMus.).QUATERNARY.NEWZEALAND:SOUTHISLAND:Mount Arthur (Archey, 1941); Wairangi (Oliver, 1955,N.Zealand Birds, ed.2,p. 585). GenustPachyornisLydekkerPachyornisLydekker, 1891 (Apr.25),Cat. Fossil Birds Brit. Mus., p. 316(typeDinornis elephantopusOwen,byoriginal designation).5.Pachyornis elephantopus(Owen)Dinornis elephantopus Oweu, 1856(July30),Proc. zool. Soc. London, pt. 24,p. ,154 (lectotype from Awamoa, left metatarslls, Brit. Mus. designatedby Archey, 1941, Bull. Auckland Inst. Mus., no. 1, p.36).Dinornis crassusvar.majorHutton, 1875(July),Trans.N.Zealand lnst., vol. 7,pp. 276-278 (lectotypefromHamilton Swamp,metatarslls,designatedbyArchey, 1941, p.38).Pachyornis immanisLydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 343,fig.66B(typefrom South Island, left tarsomctatarsus, Brit. Mus. no. A.168).Euryapteryx lmnderoRus Hntton, 1891 (Nov.), N. Zealand }onr. Sci.,newissue, vol. 1, no. 6, p.249(lectotype from Hamilton Swamp, metatarsus, OtagoMus., designatedbyArchey, 1941, p.36). Pachyorni., rothschildiLydekker, 1892(Apr.),Proc. zool. Soc. London for 1891, no. 33, p. 479 (types from unknown locality, associated right femur, tibiae,metatarsi,TringMus.),Pachyornis inhalJilis Hutton, 1893(May),Trans. N. Zealand Inst., vol. 25, p. 11(typefrom "probably somewhereinCanterbury," incomplete skeleton,Canterbury no. 9.2.23).Pachyornis valgus HnUon, 1893 (}"fay), Trans.N.Zealand Inst., vol. 25, p. 12(types from Enfield, rightandleft tibiae, CanterburyMus.).QUATERNARY.NEWZEALAND:SOUTHISLAND:Awamoa(Owen,1856); Hamilton Swamp(Hutton,1875); RuamoaandGlemnark Swamp (Lydekker, 1891, Cat., p, 321); Kapua, Waitaki River,and

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1963 BRODKORB: CATALOGUE OF FOSSIL BIRDS211StewartIsland(Oliver,1930,N.ZealandBirds, p.51);Waikouaiti,Broken River,Motunau,Riverton Beach,TakakaHill,andShagPoint(Lambrecht,1933,Handb.,Pl"150-151);Enfield(Hutton,1893);Pyramid Valley(Archey,1941,1'.138);Tarakohc,Herbert,andPapatowai (Oliver,1955,N.ZealandBirds, ed.2,p.576).6.Pachyornis pygmaeus(Hutton)Euryllpteryxpygmaeu8Hutton, loS!)1 (Nov,),N.ZealandJour. Sc.'i., newissue, vol.1,no. 6, p.249(lcctotypcsfromTakaka,rightandleftmetatarsi, Nebon designated by Hutton, 1892, Trans.N.Zealand Inst., vol. 24, p.139).QUATERNARY.NEWZEALAND:SOUTHISLAND:Takakatahleland.7.Pachyornis mappiniArcheyPachyomis mappini Archey, 1941 29),Bull. Auckland Inst. Mus., no. 1, 11,fig.4,pI.12,fig.5;pI.15,fig.Ia-c(typefrom Mangaotaki, skeleton, Auckland Mus. no.124).QUATERNAHY. NEWZEALAND: NORTHISLAND:Mangaotaki,\Vaika remoana,DoubtlessBay,Coromandel,Makirikiri,Karamu,andAmo deo Bay(Archey,1941);Waipu,Gisbome,Mangaone,Nuhaka,TeAute, Coonoor,Martinborough,andEketahuna(Oliver,1955,N. Zealand Birds, ed. 2,p.574).8.Pachyornis oweni(Haast)Dinomis oweniHaast, 1885, Proc. zool. Soc. London for 1885, no. 31, p. 482 (nomennudum).-Haast,1886(Dec.),Trans. zool. Soc. London, vol. 12,pt.,5,p.171,pI. :31-32 (typefromPataua nt;ar \Vhangarei,skeleton, Auckland Mus. no. A.M.384).QUATERNAHY. NEWZEAl.AND: NOHTHISLAND:Pataua(Haast,1886);Tom Bolling Bay,DoubtlessBay,Waikawau,andWestmerenearAuckland(Archey,1941,Bull.AucklandInst. Mus., no.1,p.44);Lake Ohia,TeAute,Martinborough,andMakirikiri (Oliver,1955,N.Zealand Birds, cd.2,p.582).9.Pachyornis septentrionalisOliverPachyornis septentrionalisOliver, 1949,"MoasN.ZealandAustralia,p. 61 (typefromPohlle,incompleteskeleton,Dominion Mus. at\Vellington).QUATERNARY.NEWZEALAND:NORTHISLAND:Pohue(Oliver,1949);Doubtless Bay,Whangarei,BayofPlenty,Waikaremoana,TeAute, andMartinborough(Oliver,1955,N.ZealandBirds, ed.2,p.574);CoonoorandWanganlli(Scarlett,1957,Proc. N.Zealandeeol. Soc., no.4,p.17).

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212BULLETINFLORIDASTATE Vol. 710.Pachyomis murihikuOliverPachyornis murihiku Oliver, 1949, Moas N.ZealandAustralia, p.67(typefromGreenhills, skeleton,SouthlandMus.).QUATERNARY.NEWZEALAND:SOUTHISLAND:GreenhillsnearBluff.11.Pachyomis australisOliverPachyornis australis Oliver, 1949, Moas N.ZealandAustralia, p.70(typefromSalisburytableland,skulland neck vertebrae,DominionMus.).QUATERNARY.NEWZEALANU:SOUTHISLAND:Salisbury tableland at headwaters of Takaka Hiver (Oliver, 1949); SouthlandandNelson (Oliver, 1955, N. Zealand Birds, ed.2,p. 575). SubfamilytEMEINAEBonaparteEmeinaeBonaparte,1854, Ann. Sci. nat.(Paris),vol. 1, p.48(typeEmeusReichenbach).'Genus tEmeusHeichenbachEmeusReichenbach,1852, Avium Systema Naturale, p. xxx (type hy monotypyDinornis crassus Owen). Syornis Reichenbach,1852,AviunlSystcmaNaturale,p.xxx(typeDinornis casuarinus Owen).MeionornisHaast,1874(June),Trans. N.ZealandInst., vol. 6, p.426(typeDinornis casuarinus Owen,designatedbyArchey, 1941, Bull.Aucklandlnst. Mus., no. 1, p.45).MesopteryxHutton,1891(Nov.),N.ZealandJour. Sci., newissue, vol. 1, no. 6, p.248(typebymonotypy Dinnrnis huttoniiOwen).12.Emeus crassus(Owen) Dinurnis craSSU8Owen,1846(July),PIoe. zool. Soc.London,pt.14,p.46(lectotypefrom Waikouaiti,nowlost,designatedbyLydekker,1891, Cat., p. 307; casts, Brit. Mus. no. A.186, Aucklaud Mus. no. A.M.298).Dinornis casuarinusOwen,1846(July),Proe. zool. Soc.London,pt.14, p. 47(lectotypefrom Waikollaiti,nowlost,designatedbyLydekker, 1891,p. 2.'57).[?]Dinornis rheides Owen,1851(Jan.1),Trans.zuul. Soc. London,vol.4,pt.1,p. 8(indeterminate?).QUATERNARY.NEWZEALAND:SOUTHISLAND:Waikouaiti (Owell, 1846); Glenmark (Lydekker, 1891, Cat., p. 257); Enfield, Kapua, Hamilton Swamp, Awamoa,Dunedin,andEarnscleugh Cave(Lambrecht, 1933,Handb.,p. 147);PyramidValleyandKiaOra(Archey, 1941, pp. 51, 149); Waitaki, Shag Hiver, Ohai, Papatowai, Greenhills, Hiverton,andWakapatu(Oliver, 1955, N. Zealand Birds, ed.2,p. 577).NORTHISLAND:MartinboroughandTeAutc (according to Archey, 1941, p. 51, thesearetheonly valid records fromtheNorthIsland).Hecorded from Stewart IslandbyLambrecht,butnot con firmedbysubsequentauthors.

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BRODKORB:CATALOGUEOFFOSSILBIRDS 21313.Emeus huttonii(Owen) Olnomis huttoniiOwen, 1879, Extinct Birds N, Zealand, p. 430 (lectotypefromHamilton Swamp, right metatarsus, Otago Mus., designatedbyArchey,1941,p.52), Euryapteryx compactaHutton, 1893(May),Trans. N. Zealand Inst., vol. 25, p.11(typefrom Enfield, tibia,CanterburyMus.).QUATERNARY.NEWZEALAND:SOUTHISLAND:Hamilton Swamp (Owen, 1879); Enfield (Hutton, 1893); Kapua(Hutton,1896, Trans.N.Zealand Inst., vo!.28,p. 636); Glenmark(Hutton,1897, op. cit., vol.29,p. 559); Takaka (Lambrecht, 1933, Handb., p. 148); Waka patu and Pyramid Valley (Archey, 1941, pp.53,140); Broken River, Waireka, Castle Rock, Waikouaiti, Papatowai, andWaipapa(Oliver,1955,p. 559). GenustEuryapteryxHaast Oela Reichenbacll, 18.52, Avium Systema Naturale, p.xxx(typebymonotypyDinornis curtusOwen).PreoccupiedbyCelaMoehring, 1758;CelaOken,1816;Celallliger, 1826.CeleusBonaparte, 1856(afterNov.3),C.R.Acad. Sci. Paris, vol. 43, no. 18, p. 841(newname forCelaReichenbach).Preoccupied byCeleusBoie, 1831. Euryapteryx Haast, 1874(June),Trans. N. Zealand Inst., vol. 6, p. 427 (typeDinorni" gravisOwen, designatedbyArchey, 1941, p.53).14.Eztryapteryx gravis(Owen)Dinomis gravisOwen, 1870(Jan.),Trans. zool. Soc. London, vol. 7, pt. 2,p.141, pI. 14(typefrom Kakanui, skull, call. BaronessA.BurdettCoutts), Emeus gravipesLydekker, 1891 (Apr.25),Cat. Fuss. Birds Brit. Mus., p. 297(type from Kakallui, metatarsus, Brit. :r,,1Ils. no.A.1591).Emeus boothiRothschild, 1907 (Nov.12),Extinct Birds, p. 210(typefrom Shuy River, skull, Brit. Mus.?).Euryapteryx kuranuiOliver, 1930, N. Zealand Birds, p..52(typefrom CastlePoint, skeleton, Cantcrlmry Mus.).QUATERNARY.NEWZEALAND: STEWART ISLAND(Benham, 1910, Trans.N.Zealand lnst., vol. 42, p. 354).SOUTHISLAND:Kakanui River (Owen, 1870); Shuy RiverandShag River (Rothschild, 1907); Mount Arthur, Hiverton, and Pyramid Valley (Archey, 1941, Bull. Auckland Inst. Mus., no.1,pp. 56, 141); Herbert, Earnscleugh Cave, and Wakapatu (Oliver, 1955,N.Zealand Birds, ed.2,p. 578).NORTHISLAND:Castlepoint (Oliver, 1930); Portland IslandandWaikare moana (Archey, 1941, p. 56)TeAute, Hunterville,andNga Rata (Oliver, 1955).15.Euryapteryx geranoides(Owen)Palapteryx geranoidesOwen, 1848 (Apr.13),Proc. zool. Soc. London,pp.1,

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214BULLETINFLORIDASTATE I>.mSEUM Vol. 7 7 (measurements of skull).-Owen,1848 (Apr.22),Trans. zool. Soc. London, vol. 3, pt. 5, p. 361,1'1.54,fig.1-5(typefromTeRangatapu, skull).Dinornis expunctu8Archey, 1927 (Aug.15),Trans. N. Zealand 1nst., vol. 58, p. 152(newname for Pa/
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1963BRODKORB:CATALOGUEOFFOSSILBIRDS215QUATERNARY.NEWZEALAND;NORTHISLAND:WangaehuRivermouth(Hutton,1897);Rangatapu(Lydekker,1891);DoubtlessBayandWaiotapu(Archey,1941,Bull.AucklandInst. MilS.,nO.1,p.141);WanganuiandNapier(Scarlett,1957,Proe. N. Zealand eeo!. Soc., no.4,p.17).19.Zelornis haasti (Rothschild)Emeus haasti Rothschild, 1907(Nov.12),ExtinctBirds,p.210(typefromGlcnmark,skull).Emeus parkeriRothschild, 1907(Nov.12),ExtinctBirds,p.211 (type fromShag Point,skull).QUATERNARY.NEWZEALAND:SOUTHISLAND:GlenmarkandShagPoint (Rothschild,1907); Enfield andRiverton (Oliver,1955,N.Zealand Birds,ed.2, p..580).FamilytDINORNITHIDAEBonaparteDinornithidaeBonaparte,1853(afterOct.31),C.R.Aead. Sci. Paris, vol. 37,. no.18, p. 646(typeDinornisOwen).-DinonlithinaeBonaparte,1853, op. cit.,p. 646 (soUS-f;:lIIlille) .-DinornithnideaeStejneger, 1884, Sci. Rec., vol.2, p.1.5,5. PalapteryginaeBonaparte,1854, Ann. Sci. nat.(Paris),vol. 1,p.48 (type Palapteryx Owen)'-Palapterygidae Haast,1874(June),Trans. N.ZealandInst., vol. 6,p.419.GenusfDinornisOwenDinornisOwen, 1843(July), l'TOC. zool. Soc.London,pt.11,no. 121,p.10(typebymonofypy IJinornis novae-zealandiaeOwen).MegalomisOwen, lR43, Proc. zool. Soc. LondolJ, pt.11,no. 122,p.19(Dinomissubstituted for walluscript name Mega[ornis Oweninpaperreadatpreviousmeeting;preoccupiedhyMegalornisGray,1841),PalapteryxOwen, 1846(July), Provo zooL Soc.London,pt.]4,p. 46(typeDinornis ingensOwen,rlcsibTllatcd byLydekker. 1891.Cat. Foss. Birds Brit. Mus., p,224). Mavia Reichenbach, 1852, Aviumsystcmanaturale,p.xxx(typebymonotypyDinorois ingensOwen).Moo Reichtmbach. 1852, Aviumsystemanaturale,p.xxx(typebyI1l0IlOtypy Dinornzs giganteusOwen). Owenia Gray, lR55, Cat,GeneraSubgeneraBirds,p, 1.52 (type Dinomisstru thoidesOwen; seeBonaparte, 1856,G.R. Acad. Sci. Paris, vol. 43, no. JR, p.841). Tylopteryx Hutton,1891(Nov.),N.ZealandJour. Sci.,newissue, vol. 1,no. B, p.247(typeDinornis gracilisOwen,designatedhyRichmond. 1902, Proe.U.S.nat.Mus., vol. 24, no. 1267, p.120;Dinornis torosus Hutton.designatedbyArchey, 1941, Bnll.Auckland Imt. Mus., no. 1.p,61,inoversightofRichmond'saction).

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216BULLETINFLORIDASTATE MUSEU-'-! Vol. 71.Dinornis novae-zealandUie OwenDinornis novae zealandiaeOwen, 1843(July),Proc. zool. Soc. London, pt. 11,no.121.p.8(ledotypesfrom Poverty Bay, left femur, left metatarsus, RoyalCollege of Surgeons, nos.f12, m3,designatedbyArchey, 1941, p. 64: casts, Brit. Mus. noo. 18588,18590).Dinorni ..gtruthoides Owen, 1844(March),Proe. zool. Soc.Londonfor 1843, pt. 11, no. 129, p. 144(briefdescription).-Owen,1844(June5),Trans. zool. Soc. London, vol. 3, pt. 3, p. 244(typefrom Poverty Bay, metatarsus, Roy. ColI. Surg., no.m3).Dinornis struthioidesLydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 242(cmendation).QUATERNARY.NEWZEALAND:NORTHISLAND:PovertyBay(Owen,1843);Wanganui,Hastings, Doubtlcss Bay, Karamu, Mangaotaki,Waikaremoana,andHaupouri(Archey, 1941,pp. (14,fJ7, 142);'Naipu,Tahora,TeAnte, Martinborongh, Makirikiri,and Pm'cmata (Oliver, 1955, N. Zealand Birds, ed. 2,p.585).2.DinornistoroSllSHuttonDinornis torosusHutton, 1891(Nov.),N. Zealand Jour. SeL,newissue, vol.1,no. 6, p. 247(typefrom Takaka, Auckland Mus. no. A.M.352).Palapteryx plenusHutton,1891(Nov.),N.Zealand Jour. Sci., newissue, vol. 1, no. 6, p. 248 (leetotype from South Island, tibia, selectedbyArchey, 1941, p. 70).Dinomis strenuusHutton,1893(May),Trans. N, Zealand Inst., vol. 25, p. 8 (leetotype from EnEeJd, metatarsus, selectedbyArchey, 1911, p. 70, CanterburyMus. no. 1.14.13).QUATERNARY.NEWZEALAND:SOUTHISLAND:Takaka(Hutton,1891); Enfield(Hutton, 189.'3); MountArthur,Glenmark,Timarn,andHamiltonSwamp(Archey, 1941,pp.70, 143);Kapua(Hutton,1896, Trans. N.ZcalandInst., vol. 28, pp. 6.'34, 642);Broken River,Herbert,Shag River, Castle Rock,TakakaHill,andSlovensCreek(Oliver,1955, N. Zealand Birds, ed.2,pp.585,586);Rahu(Scarlett, 1957, Proc. N. Zealand ecol. Soc., no. 14, p.17).3.Dinornis ingem OwenDinorni" ingens Owen, 1844(June5), Trans. zool. Soc. London, vol. 3,pt.3,p.247(typefrom Poverty Bay, tibiotarslls, Hoy. CoIl. Sllrg. no.t2;cast Brit.Mus.).Dinorni ..graci/i.. Owen, 1855 (Apr.11),Proc. zool. Soc.Londonfor 1854, pt. 22, p. 246 (lectotype fromNorthIsland, metatarsus, Brit. Mus. no. 32272, selectedbyLydekker, 18\)1, Cat., p.248).Dinornis lirmus Hutton, 1891(Nov.),N. ZealandJOUf.SeL,newissue,vol1,no.6,p.247(lectotypes from Poverty Bay, femnr, tibia, metatarsus, colI. ofRev. W. Colenso, selectedbyArchey, 1941, p.68).

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 217QUATERNARY.NEWZEALAND:NORTHISLAND:Poverty Bay(Owen,1844); Karuma, Mangaotaki, Waikarcmoana, Te Aute, Patangata, Kai waka, Hastings,andMakirikiri (Archey, 1941, pp.68,143); Ruakaka, Matapouri, Clevedon,TeKuiti, Moawhango, Coonoor, Martin borough, Karori, Paekakariki,andKaiiwi (Oliver, 1955,N.Zealand Birds, ed.2,p..586).4.Dinornis robustusOwen Dinorni" ingensvar.robustusOwen,1846(July),Proc. zool. Soc. London,pt.14, p. 48(lectotypefrom South Island, metatarsus, Roy. Call. Surg., nowapparcnlly lost, selectedbyArchey, 1941, p.71).Dinornis potensHutton, 1891(Nov.),N. Zealand Jour.Sci"newissue, vol. 1,DO.6,p.247(typesfromHeathcote, femur, tibia, metatarsus, Canterbury]\/us.).QUATEHNAHY. NEWZEALAND: SOUTHISLAND:\Vaikouaiti(Owen,1851, Trans. zoo!.Soc.London,vo!.3,pt. 4, pp. 321, 329); Hamilton Swamp (Hutton, 1875, Trans.N.Zealand Inst., vol.7,p. 279); Heath cote(Hutton,1891); Greymouth(Hutton,1892, Trans.N.Zealand Inst.,vo!.24,p. 113); Kapua and Enfield(Hutton,1896, Trans.N.Zealand Inst.,vo!.28, pp. 633, 645); Castle Rock, Timaru, Glenmark, Knobby Range, TigerHill,andPyramid Valley (Archey, 1941, p. 144); Takaka Hill, Westport, BrokenRiver,Papatowai, Clyde,andD'UrvilleIsland (Oliver, 1955, p. 586).5.Dinornis giganteusOwenDinornis giganteusOwen, 1844 ('>/arch), Proc. zool. Soc.Londonfor 1843,pt.11, no. 129, p. 144(typefrom Poverty Bay, tibia, Roy. Coll. Surg. no. 2170; cast Brit. Mm. no.18588).Dinornis excelsusHutton 18tH(Nov,), N. Zealand Jour.SeL,newissue,vol.1,no. 6, p. 247(lectotypefromTeAute, tibia, selectedbyArchey, 1941, p.69).QUATERNARY.NEWZEALAND:NOHTHISLAND:Poverty Bay(Owen,1844);TeAute(Hutton,1891); Doubtless Bay, Awhitu, Moawhango, Makirikiri, Maungaraki Gorge, and Hawke Bay (Archey, 1941, p. 143); Coononr, Omaranni,andMartinborough (Oliver, 1955, p. 588).6.Dinornis rnaxirnusOwenDinornis maximusOwen, 1867, Proc. zool. Soc.Londonfor 1867, no. 57, p. 891(nomennudum).-Owen,in Haast, 1869(May),Trans. N. Zealand Inst., vol.1, p. 87 (typesfromGlcnmark Swamp,femur,tibia, and part of metatarsus,CanterburyMus.).-Owen,1869(June1),Trans. zool. Soc. London, vol. 6, pt. 8, p. 497, pI. 89-90 (types from Glenmark Swamp, from same individualas Haast's types, left femur, left tibiotarsus, right tarsometatarstls, colI.ofMajorJ. nowsupposedtobeinMadras Mus.butapparentlylost; casts Brit. Mus. no. A.161, Auckland Mus. no. A.M.385).

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218BULLETINFLORIDASTATE MUSEUM Vol. 7Dinornis altusOwen, 1879, Extinet Birds New Zealand, pp. 253, 361, pI. 79, Ilg. 4(typefrom South Island, left metatarsus, Brit. Mus. no. 3.'58:32). DinornisvalidmHutton, 1891(Nov.),N. Zealand Jour. Sci., new issne, vol.1,no. 6, p.247(typefrom Glcnmark Swamp, skeleton, Canterbury Mus.). QUATERNARY. NEW ZEALAND:SOUTHISLAND:Glenmark Swamp(Haast,1869); Kapua, Enfield,andRiverton(Hutton,1896, Trans. N. Zealand Inst., vol.28,pp. 632,646, 652); Pyramid Valley, Broken River, Shag Valley, Waikouaiti,andSumner (Archey,1941,p.144);Raki's Table, Herbert, Awamoa, Seacliff, Colac Bay,andInvercargill (Oliver, 1955, p. 588).7.Dinornis gazellaOliverDinornis gazellaOliver, 1949, MoasN.ZealandandAustralia, p. 1Illl(typefromTeAnte, pelvis, Dominion Mus., \Vellington).QUATERNARY.NEWZEALAND:NORTHISLAND:TeAute (Oliver, 1949); Karamu, Makirikiri,andParemata(Oliver, 1955, p. 585).8.Dinornis herculesOliverDinornis herculesOliver, 1949, N. Zealand and Australia,p.174(typefrom Caonoor, tibia, Dominion 1-fus., \Vcllington).QUATERNARY,NEWZEALAND:NORTHISLAND:Coonoor (Oliver, 1949); Waitomo, Mangaone,TeAute, Makirikiri, Doubtless Bay, Moawhango, Poverty Bay,andAwhitu (Oliver, 1955, p. 588).

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1963 BRODKORB:CATALOCUEOFFOSSIL BIRDS 219 OrderAPTERYGIFORMES(Haeekel)ApterygiaIIaeekcl, 1866, Genere]]e MorphologiederOrganism en, vol. 2, p. 139(typeApteryxShaw).Family APTICRYGIDAE (Gray)ApteryginaeGray, 1840, List Genera Birds, p.63(typeApteryxShaw,Recent).GenustPseudapteryxLydekkerPseudapteryxLydckker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 218(typebymonotypyPseudapteryx gracilis Lydekker).1.Pseudapteryx gracilisLydekkerPseudapteryx gracilis Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 218,fig. .53A(typeleft tarsometatarsus, Brit. Mus. no.32237a).PLEISTOCENE.NEWZEALAND.NeospeciesofApterygidaefromQuaternarysites:1.ApteryxaustralisShaw.NEWZEALAND:SOUTHISLAND:Timarlland Nelson(Lydekker, 1891, p.216);Pyramid Valley (Scarlett, 1955, Ree.CanterburyMus., vol. 6, no.4, p.261).NORTHISLAND:Waingongoro (Lydekker, 1891,p.217).2.Apteryx oweniiGould. N"w ZEALAND:SOUTH I.LAND? (Lydekker, 1891,p.218).NORTHISLAND:Akiteo, Kamao,Opita, Hukanui, Pigeon Bush,andRangatapuPa(Scarlett, 1962, Notomis, vol. 10, p.84).3. Apteryx haasliiPotts.NEWZEALAND:SOUTHISLANU: Nelson (Lydekker71891, p.217).

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220BULLETINFLORIDASTATEMUSEUMInfraclassCARINATAEMerrem'Vol. 7Aves CarifUltae Merrem,1813,Abh. Akad. Wiss. Berlin, p.259.-CarinataeHuxley,1887,Proc. zool. Soc. London, p.418(order).NeognathaePycraft,1900,Trans. zool. Soc. London, vol.15,p.149.OrderGAVIIFOHMESWetmoreandW.D.MillerEnaliornithes Fiirbrinller, 1888,Untersuch. Morph. Syst. Vogel, vol.2,p.1543 ("gens/, typeEnaliornisSeeley).ColymbiformesSharpe,1891,Revicw Rcccnt Attempts to Classify Birds, p.71(order;typeColymbusLinnaeus,Le.loons).-ColymhiGadow,1893,Bronn Klass. Ordn., Vogel, pt.2,pp.76, 121, 299 (Unterordnunll, for loons).GaviiformesWetmoreandW.D. Miller,1926(July),Auk, vol.43,no.3,p.340(typeGaviaForster).FamilytENALTORNITIIIDAEFiirbringerEnaliornithidaeFiirbringer,1888,Untersuch. Morph. Syst. Vogel, vol.2,pp.1152, 1426note,1543, 1.565 (typeEnaliornisSeeley).GenustEnaliornisSeeleyPalaeocolyntus[sic]Seeley,1864,Proc.Cambridgephilos. Soc., vol.1,p.228(nomennudum,titleonly,notext).Pelargonis[sic] Seeley,1864,op. cit., p.228(nomennudum,titleonly).PelagornisSeeley,1866,Ann. Mag. nat. Hist., ser.3,vol.18,p.110(nomenl1udum).-Sceley,1876,Quart. Jour. geol. Soc. London, vol.32,p.497footnote(typebymonotypyPelagornissedgwickiSeeley; name available fromtllis date,butrejectedbySeeleyandpreoccupiedbyPelagornisLartet,1857).EnaliornisSeeley, 1869, Index to the fossil remains of Aves, Ornithosauria, and Reptilia fromthesecondary system of strata arranged in the vVoodwardian Museum of the UniversityofCambridge, p. xvii(nomennudum; the refer ence top.7ofthis work is merely a listofelements, without descriptionorname).-Seeley,1876(afterJunc7),Quart. Juur. geol. Soc. London, vol.32,p.499(name valid from this date; type by present designationEnaliornisbarretttSeeley).PalaeocolymbusSeeley,1876(afterJune7),Quart. Jour. geol. Soc. London, vol.32,p.497footnote(nameavailable from this date,hutrejectedbySeeley;typebymonotypyPalaeocolymbus barrettiSeeley).1.Enaliornis barrettiSeeleyPalaeoeolyntus[sic]BarrettiSeeley,1864,Proc.Cambridgephilos. Soc., vol.1,p.228(nomennudum, titleofarticleonly).Pelagorni., BarrettiSeeley,1866,Ann. Mag. nat. Hist., ser.3,vol.18,p.110(nomennudum).-Seeley,1876(afterJune7),Quart. Jour. geol. Soc. London, vol.32,p.496. Enaliornis BarrettiSeeley,1869,Index AvesvVoodwardianMus., p. xvii(nomennudum).-Seeley,1876(afterJune7),Quart. Jour. geol. Soc. London, vol.32,p.499,pI.26,fig.1-11, 14-27;pI.27,fig. 1-5, 19-25(original description;INewrank.

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1963BRODKORB: CATALOGUEOFFOSSIL BIRDS221 lectotypebypresentdesignation fromnearCambridge,distalendof left tarsometatarsus, call.ofT.Jessou; cast Brit. Mus. no. A.1112).Pakleocolymbus BarretliSeeley, 1876, op. cit., p. 497 footnote.LOWERCRETACEOUS,ALBIAN(UpperGreensand).ENGLAND:Cambridgeshire:probablyncarColdhamCommonorGranehester.2.Enaliornis sedgwickiSeeleyPelarganis[sic]SedgwickiSeeley, 1864, Proc.Cambridgephilos. Soc., vol. 1, p.228(titleofarticleonly). ETUJ1iornis SedgwickiSeeley, 1869, Index Aves WoodwardianMus"p.xvii(nomennudum).-Sceley,1876(afterJune7),Quart. Jour. geol. Soc. Lon don, vol. 32,p.501, pI. 26, fig. 12-13; pI. 27, fig. 6-7, 9-11, 13-18 (originaldescription; lectotype bypresent designation, from near Cambridge,proximal endofright tibiotarsus, Woodwardian Mus.).Pelagomis SedgwickiSeeley, 1876(afterJune7),Quart.Jour. geol. Soc. Lon rlan, vol. .32, p. 497 footnote.LOWERCRETACEOUS,ALBIAN(UpperGreensand).ENGLAND:Cambridgeshire;probablynearColdhamCommonorGranehester.FamilytLONCHODYTIDAEBrodkorbLonchodytidaeBrodkorb, 1963(inpress),Proc.XIIIinternat. ornith. Congr.Ithaca,p.000(typeLonchodytesBrodkorb) .GenustLonchodytesBrodkorbLonchodytesBrodkorb, 1963(inpress),Proe. XIII intern at. ornith. Congr.Ithaca,p.000(typebyoriginal designation LOTUJhodytes estesiBrodkorb).1.Lonchodytes estcsiBrodkorbLonchodytes estesi Brodkorb, 1963(inpress),Proc.XIIIinternat. ornith. Congr.Ithaca,p. 000,fig.1-2(typefromLanceCreek, distalpartofrighttarsometatarsus, Dniv. Calif. Mus. Paleo. no.53954).UPPERCRETACEOUS,MAESTRICHTIAN(Lanceformation).WYOMING:NiobraraCounty;LanceCreek.2.Lonchodytes pterygiusBrodkorb Lonchadytp-s pterygius Brodkorb, 1963(in press), Proc.XIlIinternat. ornith.Congr. Ithaca, p. 000, fig. 3(typefrom Lance Creek, distal part of left carpometacarpus, Univ. Calif. Mus. Palco. no.53961).UPPERCRETACEOUS,MAESTRICHTIAN(Lanceformation). ,\VYOMING: NiobraraCounty:LanceCreek.

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222BULLETINFLORIDASTATEMUSEUMFamilyCAVIIDAEAllenVol. 7Colymhidae"Leach,"Vigors, 1825, Trans. Linn. Soc. London, vol. 14, p. 498(typeColymbusLinnaeus,Le.,loons,incontrastwith Podiceps Latham).ColymhinaeBonaparte,1831, SaggiodiunadistribuzionemctodicadegliAnimaliVertebrati, p. 62 (subfamily for loons).UrinatoridaeRidgway,1887, Man. N.Amer.Birds,pp.4, 6(typeUrinatoreuvier,1800,ajuniorsynonymof Cavia Forster,1788).GaoiidaeJ.A.Allen, 1897(July),Auk, vol. 14, no. 3, p. 312(typeGaviaForster).SubfamilyfCOLYMBOIDINAEBrodkorb1Genus fEupterornisLemoineEupterornisLemoine, 1878, Recherches sur les oiseaux fossiles des terrainstertiaires inerieurs desenvironsde Rehus, vol. 1,p.56(typebymonotypy Eupterornis remensisLemoine).Positiontentative.1.E upterornis remensisLemoine Euptemrnis l'emensisLemoine,1878,cp.cit., pp. 12,56,pI. 5, fig.1-6(typesdistalhalfofleft ulna, phalanx 1ofindex fin!,:er). UPPERPALEOCENE(conglomeratedeCernay).FRANCE:Dept.Marne: Chalons-sur-VeslenearSoissons.CenusfColymboidesMilne-EdwardsColymboidesMilne-Edwards, 1867, Ois. Foss.France,vol.1,pI. .'>4, fig. 1-14: Milne-Edwards, 1868, op. cit., vol. 1, sheet 38, p. 297(typehymonotypyColymhoidesminutusMilne-Edwards).HydrornisMilne-Edwards, 1867, Ois. Foss.France,vol.1,pI. 57, fig. 18-22; Milne-Edwards, 1868, op. cit., vol.1,sheet 46, p. 362(typebymonotypyIIydrornis natatorMilne-Edwards).DyspetornisOberholser, 1905(May13),Smithsonian misc. Call., vol. 48,pt.1, no.1.579, p. 61(new name forHydrornis preoccupiedby Hy
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1963BRODKORB:CATALOGUEOFFOSSILBIRDS2233.Colymboides minutusMilne-Edwards Colymboide" minutusMilne-Edwards,1867,Ois. Foss. France, pI.54,fig.1-14: 1868,vol.I,sheet38,p.297(typesrighthumerus,rightulna,2left femora, ParisMus.).Hydrnrnis natatorMilne-Edwards,1867,Ois. Foss. France, vol.I,pI.57,fig.18-22: 1868,sheet46,p.362(typerighttarsometatarsus, ParisMus.).LOWER (Aquitanian).FRANCE:Dept.Allier:Langy.SubfamilytGAvTELLTNAEWetmore Gaviellinae Wetmore,1940(Jan.2),Jour. Morphol., vol.66,no.I,p.30(typeGavie/ia\Vetmore).GenustGaviellaWetmoreCaviellaWetmore,1940(Jan.2),Jour. Morphol., vol.66,no.I, p. 28(typebyoriginal designationGavia pusillaShufeldt).4.Gaviella pusilla(Shufcldt) Cavia pusillaShufeldt,1915(Feb.),Trans. Connecticut Acad. Arts Sci., vol.19,p.70,pI.13,fig. 106 (typeproximal portiun uf left carpumetacarpus, YalePeabody MIlS. no.864).OLIGOCENE (vVhite Rivergroup).VVYOMTNG:NiobraraCounty:nearLusk. SubfamilyGAVTTNAE(Allen) Gaviinae Wetmore, 1940 (Jan.2),Jour. Morphol., vul.66,nu.I,p.30.CenusCaviaForster' Colymbus Linnaeus,17.58,Syst. Nat.,eel.10,vul.I,p.13.5(typeColymhus arctieusLinnaclls,designated hyCray, 185.5, Cat. Gen. Suhgen. Birds, p. 125,andbyLawrence,18.58,Rept. Expl. Surv.R.R.Pac., vol.9, p. 887).Genericname suppressedbythe International Commission,whenit could notdecidewhichwasthetype species.2Gada Forster, 1788, Enchiridion historiac naturali,p. 38(typP,Colymhus immerBruuuich: see Allen,1907,Bull. Am. Mus. nat. Ilist., vol.23,p.290). IGavia, sp. indet., recorded from Middle Miocene (Calvert formation) nearPluUl Point, Calvert County, Maryland(Wetmore,1941,Ank, vol. ,,,}8, p. ,"}(7). would .seem that Colymbus shouldberestoredasthe generic name of the loons, with the namesoftheeorresponcling higher taxa altered accordingly.Thedesignation ofColumbus arcticusastypeofthe genus long antedates the desig nationofa grebe,Columbus cristatusLinnaeus,byBaird, Brewer, and Ridgway(1884,WaterBirds Amer., vul.2,p.425)andbyHellmayrandConover(1948, Field Mus. PubI., zool. ser., vol.13,pt.I,nu.2,p.18).Theaction ofBrisson(1760,Ornithologia, vol. 6, p.33)in aeliminating" theloons from Columbus appears to have no bearing under the rules as written,insoiteof theurging uf Stejneger(1882,Proc. U. S. nat. Mns., vol. ,"), p.42)andAllen(1897,Auk,voJ.14, p. who were operating under a different code. Salomonsen(1951,Proe. Xintemat.omith.CDngress, pp.149-1.54)hasoutlinedthehisturyof this nomenclatorial controversy. I useCaviahere, with reluctance, in deferenceto the International Commission.

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224BULLETINFLORIDASTATEMUSEUM5.Cavia palaeodytesWetmoreVol. 7 Cavia palaeudytes Wetmore, 1943(June23).Proc.NewEnglandzool. Club, vol. 22, p. 64,fig. 1-2(typefrom Pierce, left coracoid, Mus. Camp. Zool.Harvardno. 2369; cast Brodkorb coIl.).LOWERPLIOCENE(Bone Valley gravel).FLORTDA:Polk County: Pierce (Wetmore, 1943); Brewster (Brodkorb, 1953, Condor, vol. 55,p.212).6.Cavia concinnaWetmore CatJia concinna Wetmore, 1940 (Jan.2),Jour. Morpho!., vol. 66, no.1,p. 2.5, fig.1-4(typefrom Sweetwater Canyon, proximal portionofleft ulna, U.S.Nat. Mus. no.16160).LOWERPLIOCENE(Bone Valley gravel).FLORIDA:Polk County:nearBrewster (Brodkorb, 1953, Condor, vol. 55, p. 211).MIDDLEPLIOCENE(Etchegoin formation).CALIFORNIA:MonteIey County: Sweetwater Canyon east of King City (Wetmore, 1940).MIDDLEPLIOCENE(SanDiego formation).CALIFORNIA:San Diego County: San Diego(Howard,1949, Publ. Carnegie Instn. Washington,nO.584,p.185).7.Cavia howardaeBrodkorbCavia Iwwardae Brodkorb, 1953(July20),Condor, va!. 55, no. 4,p.212, fig;. IB(typefrom San Diego, distal portion of left humerus, Los Angeles Mus. no.2111).MIDDLEPI.IOCENE(SanDiego formation).CALIFORNIA:San Diego County: San Diego. Heported in error from Florida (Wetmore, 1956, Smithsonian misc. Call., vol. 131, no. 5, p.7).8.Cavia portisi (Hegalia) Colymbusportisi Regalia. HJ02, Palaeontogr. italica, va!. 8, p. 231, pI. 27,fig.19-20(typefromOrciano Pisano, cervical vertebra, Roberto colI.ondeposit in Istituto di Studi SupcrioriinFlorence).MIDDLEPLIOCENE(argillemarine). ITALY: provincia di Pisa: Orciano Pisano near VallediFine. Neospecies of Caviidae from Pleistoceneand'prehistoricsites;1.Cavia stel/ata(Pontoppidan).DENMAl\K:Mejlgaard, Havnoe, ErteboeIle,Gudumlund, Klintesoe, Havelse, Soelager,OnmlAa, and >1llKolding Fjord

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1963llRODKORll:CATALOGUEOFFOSSILBIRDS 225(H.Winge, 1903, Vidensk. Meddel. naturh. Foren. Copenhagen, vol. 6, p.91).IRELAND:Shandon cave (Lydekker, 1891, Ibis, p.394).ENGLAND:Mlmdesley(E.T. Newton, 1883, Geol. Mag., p. 97, pI.3).ITALY:GroUa Romanelli and GroUa dei Colombi?(Lambrecht,1933,Handb.Palaeorn., p.731). ALASKA: Lawrence Island (Friedmann, 1934, Jour. \Vashington Acad.SeL,vol.24,p.86);"Amaknak Island and"CapeDenbeigh(Friedmann,1934, op. cit.,pp.231,237);"Kodiak Island(Friedmann,1935, op. cit., vol. 25, p.46);"CapePrince of Wales(Friedmann,1941, op. cit., vol. 31, p.405).CALIFORNIA:Newport Bay(Howard,1958, Condor, vol. 60, p. 1,36);'Emeryville(Howard,1929, Univ. Calif. Publ. Zool., vol. 32, p.326). 2. Cavia arctica(Linnaus). DENMAHK: Fannerup, 1\Iejlgaanl, Ertebuel1e, I\.faglemose, Klintesoe, Soelager, 'O-Borresbjerg, and Fjord(H.Winge,1903, Vidensk. Meddel. naturh. Forcn. Copenhagen, vol. 6, p.91).ALASKA:St.LawrenceIsland(Friedmann,1934, Jour. Washington Acad. Sci., vol. 24, p.86);"Kodiak Island (Friedmann,1935, op. cit., vol. 25, p.46);"DutchHarbor(Friedmann,1937, op. cit., vol. 27, pp. 432,435);'CapePrince of Wales(Friedmann,1941, op. cit., vol. 31, p.405).WASHiNGTON:'PugetSound(L.Miller, 1960, Wilson Bull., vol. 72, p.394).CALIFORNIA:SanPedroand?Newport Bay(Howard,1949, Condor, vol. 51, p.21);'BuenaVistaLake(DeMay,1942, Condor, vol. 44, p.228).3. Cavia immer(Brtinnieh). NOI{WAY: Vardo(Lambrecht,1933,Handb.Palaeorn;, p.731).IRELAND:EdenvaleCave(Lambrecht,1933).ALASKA:"KodiakIsland(Friedmann,1934, Jour. Washington Acad. Sci., vol. 24, p.234); (l.-Little Kiska Island (Friedmann, 1937,ap,cit., vol. 27,p.436); o-Cape PrinceofWales (Friedmann,]941,ap. cit., vol. 31, p.405). \VASHIN(;TON:bPnget Sound(L.Miller, 1960, Wilson Bull., vol. 72, p.394).CALIFORNIA:San Pedro?(L.Miller, 1914, Univ. Calif. Publ. Geol., vol. 8, p.33);DelRey Hills?(Howard,1936, Condor, vol. 38, p.211);Lomita?(Howard,1944, Bull.S.Calif. Aead. Sci., vol. 43, pt. 2, p.75);NewportBay(Howard,1949, Condor, vol. 51, p.21);"Emeryville(Howard,1929, Univ. Calif. Publ. Zool., vol. 32, p.325).NOVASCOTIA: ORear Riverand Timber Island Brook (HalifaxMus.). :r"IARYLAND: bctweenChesapeake BeachandPlum Point(Wetmore,1962, Smithsonian misc.Coil., vo!. 145, no. 2, p.3). FLOI{WA: Lake Monroe (Brudkorb, 1953, Condor, vol. 55, p.214);Rock Spring (Woolfenden, 1959, Wilson Bull., vol. 71, p. 1R.5); Wakulla Spring (Brodkorb coil.); "Big Pinc Key(Wetmore,1935, Auk, vol. 52, p.300);"Good's shellpit (Neill, Gut,andBrodkorb, 1956, Amer. Antiquity, vol. 21, p.388);"GreenMound(Hamon,1959, Auk, vol. 76, p.533);"SummerHaven(Brodkorb, 1960, Auk, vol. 77, p.342).Thesupposedrecords fromDENMARK(Lambrecht,lococit.)referto Cavia arctica.4.Cavia adamsii(Gray).ALASKA:"St. Lawrence Island(Friedmann,1934,Jour.\Vashington Acad. SeL, vol. 24, p. 86);Amaknak Island, (l.-Cape Dcnheigh,and'Kowieruk(Friedmann,1934, op. cit.,pp.231,237);"KodiakIsland(Friedmann,1935, op. cit., vol. 25, p.46);"DutchHarborand"Little Kiska (Friedmann,1937, op.cit"vol. 27, pp. 432, 435-436);"CapePrince of Wales (FriedmanTl, 1941, op. cit., vol. 31, p.405).

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226BULLETINFLORIDASTATE MUSEUM OrderPODICIPEDIFORMES(Fiirbringer)Vo!' 7PodidpitiformesFiirbringer,1888,Untersuch.Morph.Syst. Vogel, vol. 2,pp. 154.3, 156,5(subordo;typePadicepsLatham).-PodicipediformesSharpe,1891, ReviewRecentAttemptsto Classify Birds,p.71(order).-PodicipedesGadow, 189.3, Bronn Klass. Ordn.,Vogel, pt. 2,pp.76, 121,299(Untcrordmmg).FamilytBAPTORNITHIDAEAmerican Ornithologists' UnionBaptomithidaeAmerican Ornithologists' Union, 1910, Check-listNorthAmer. Birds, ed. 3, p. 378(typeBaptomis Genus tBaptornisMarshBaptomisMarsh, 1877, Amer., Jour. Sci., scr. 3, vol. 14, p.86 (type hymonotypy Baptornis advenusMarsh).I.Baptornis advenusMarshBaptomis advenusMarsh, 1877, Amer. Jour. Sci., ser. 3, vol. 14,p.86(typefrom\Vallace Co., juvenile right tarsometatarslis, YalePeabody 110.1465).UPPEIlCRETACEOUS,CONIACIAN(SmokyHillchalkmemberof Niobrara formation). KAl\'SAS: WallaceCounty (:-'1arsh, 1877); Butte Creek in Logan County(Lambrecht,1933,lIandb.Palaeorn., p. 258). Cenus tNeogaeornisLambrechtNeogaeomisLambrecht,1929, Pal. Zeitsehr., vol. 11, p. 121(typebymonotypyNeogaeornis wetzeliLambrecht),2.Neogaeornis u.;etzeli LambrechtNeogaeornis wetzeliLambrecht, 1928, Pal. Zdtsehr., vol. 11, p. 121,fig.1-4(typefromSanVicenteBay, tarsometatarsus, Kiel Univ.Mus.).UPPEHCRETACEOUS,MAESTlUCHTIAN(Quiriquinabeds).CHILE:ProvoConcepcion: west end of San Vicente Bay, Tumbes peninsula(Lambrecht,1929); Cerro del Concjo, Vegas del Gualpen, southeastofSan Vicente in Dept. Talcahllano (Schneider, 1940, Revista Chilena Hist. Nat., p.51).FamilyPODICIPEDIDAE(Bonaparte)PodicepinaBBonaparte,1831, Saggio ui unadistribuzione Illptodica degliAnimaliVertehrali,p.62(type PodiceJls Latham.)-PodiciJlidaeBonaparte, 185.3, C.R. Acad. Sci. Paris, vol. .37, no. 18,p.646.-PodicipedidaeCoues, 1880(Sept. .30), Bull. U.S.geol. geog. Surv.Terr.,vol. 5, no. 4, p.1O;39.-PodiciJlitidaeForhes,1884(Jan.),Ibis, ser.,5,vol. 2, no.5,p.119.-PodiciJletidaeAllen, 1907 (Apr. 1.5), Hull. Amer. Mus. Nat. Hist., vol. 23,p.287.

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1963BRODKORB:CATALOGUEOFFOSSILBlRDSGenusPodicepsLatham227PodicepsLatham,1787,SupplementtotheGeneralSynopsis of Birds, vol. 1,p.294(typeColymbus cristatusLinnaeus).1.Podiceps oligoceanus(Shufeldt)Colymbus oligoceanusShufeldt, 1915(Feh.),Trans.ConnecticutAead. Arts Sci., vol. 19,p.,54(typedistalpartofleftfemur, Yale PeabodyMus. no.983).Wetmore,1937, Proe. California Acad. Sci., ser. 4, vol. 23, no. 13, p. 197,fig.6-7(typerestudied).LOWER MIOCENE (JohnDayformation).OREGON:MalheurCounty: lower reaches of Willow Creek.2.Podiceps pisanus(Portis)Fulica pisanus Portisl 1889, Gli ornitoliti del Valdamosllperioree di alcune
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228BULLETINFLORIDASTATEMUSEUMVul. 75.Podiceps dixiBrodkorbPodieel's dixiBrodkorb, 1963(inpress),Quart. Jour. Florida Acad. Sci., vol. 26, no. 1, p. 000,fig.1-2(typefrom Reddick, proximalpartofrightcarpo metacarpus, Brodkorb no.1113).MIDDLEPLEISTOCENE(Reddickbeds).FLORIDA:Marion County: Dixie Lime Products Company mine, 1 miles south of Reddick. GenustPliodytesBrodkorbPliodytesBrodkorb, 1953(Dec.),Ann.\lag.nat. IIist., ser. 12, vol. 6, p. 953(typebyoriginal designationPliDdytes lanquistiBrudkorb). 6. Pliodytes lanquZstiBordkorbPliodytes lanquistiBrodkorb, 1953(Dee.),Ann. Mag. Nat. Hist., ser. 12, vol. 6 p. 953, fig:(typefrom Brewster,rightcoracoid, Brodkorb no.299).LOWERPLIOCENE(BoneValleygravel).FWRIJ)A:PolkCounty: south of Brewster. Neospecies of Podicipedidae from Pleistoceneand"prehistoric sites:1.Podiceps Tuficollis(Pallas). DEKMAlU(: Erteboelle and Suelager(H.Winge, 1903, Vidensk. Meddel. naturhist. Foren. Copenhagen, vol. 6, p.90).IRELAND:Newhall Caveand Edenvale Cave (Lamhrecht, 1933,Handb.Pal aeorn., p. 7.31). ITALY:Grotta dei Colombi(Lambrecht,19,33).GERMANY:Weimar-Taubaeh(Lambreeht,1933).2.Podiceps dominicus(Linnaeus).BRAZIL: Lapa da Escrivania(0.\Vinge,1887, E Mus.Lund.,vol. 1, no. 2, p.25).3. Podicel's rufopectus (Cray).NEWZEALANV:"PyramidValley Swamp (Searlett, 1955, Rel'. CanterburyMus., vol. 6, p.261).4. Podicepsallritus (Linnaclls).ITALY;GroUa Romanelli, Terra d'Otranto,anrlr;rottadeiColombi(Lambreeht,1933,Handb.Palaeorn., p.731).HUNCARY:Pilisszanto(Lambrecht,191:3, Aquila, vol. 20, p.428).MONGOLIA:Sjam-Osso Gol, Ordos(Lambrecht, 1933, p.731).ALASKA:"Kodiak Island(Friedmann,19:3,5,Jour.WashingtonAcad. Sci., vol. 25, p.46).CALIFORNIA:San Pedro?(Howard,1949, Condor, vol..51,p.21).NOVA Sc.oTIA: "Whynacht(HalifaxMus.).TENNESSEE: hone caves (Shufeldt, 1897, Amer. Natural., vol. 31, p.646). Fr.mUJ)A: Seminole Field (\Vetmore, 1831, Smithsunian misc. ColI., vol.8.5,no.2,p.12);Rock Spring (Woolfenclen, 19.'>8, Wilson Bull., vol.71,p.185).Erroneous records include Fossil Lake, Oregon (Shufeldt, 1892, Jour. Acad. nat. Sci. Philadelphia, vol. 9, p. 396; correctedbyHoward,1946, Publ. Carnegie Insto. Washington, no.5.51,p.148),andItehtuekneeRiver,Florida(Wetmore,1931, p.12),thelatterhasedon a largehumerusof Podilymbus podiceps, formerly Florida Geol. Surv. no. V-4619, now Broclkorb no. 8001.5. Podieel" ea'1JicllS (Hablizl).HUNGARY:Nagyharsany Berg?(Lambrecht,1916, Aquila, vol. 22, p.174).WASHINGTON:"PugetSound(L.Miller, 1960, WiI-

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS229 IIl son Bull., vol. 72, p.394).OREGON:FossilLake(Shufeldt,1892, Jour. Acad. nat. Sci. Philadelphia, vnl. 9, p.396).CALIFORlNA:SanPedro(Howard,1949, Con dor, vol. 51, p.21);"BuenaVistaLake(DeMay,1942, Condor, vol. 44,p.228).NEVADA:SmithCreekCave(Howard,1952, Bull.S.Calif. Acad. Sci., vol..51,pt.2, p..54).KANSAS:Jones Sink (Downs, 19.54, Condor, vol. 56,p.209).Recorded alsofrOln i\1iddle Pliocene Edson beds ofOgallala formatiun,Shermau County, Kansas(Wetmore,1937),Condor, vol. 39,p.40),but need, comparisonwithnewly described forms.6.Poc/ieeps cristatu8 (LinnaCllS).DENMARK: !\/Iejlgaarcl, Havlloe,Krabhesholm, Virksund, E.rteboelle, Maglemose, Klintesoe, Hoensehals, Havclse, Soclager, Aalborg, and Rocshorg Soe (H. 'Vinge, 1903, Viddcnsk. natur hist.FUTCH.Copenhagen, vol. 6, p.90).SWEDEN: ncar()nnarp (Lamhrecht,1933, Hanelb. Palaeorn., p.731). lJ"':LAND: Kesh Cave,EdenvaleCave, Bantiek Cave,andNewhallCave(Lambrecht, ENGLAND:Cam hridgeshire fens(Milne-Edwards,1868, Ihis,p.364).hALY:GrotiaRomanelli(Lamhreeht,1933).7. Poc/ieeps grisegena(Boddaert).DENMARK:Erteboelle(H.Winge, 19Q.1, Vidensk. Meddel. naturhist. Foren.Copenhagen,vol. 6, p.90). CZEc.HOSLO VAKIA:Certova dira (Capek, 1910, Ber. V lnternat. Orn. Kongr. Berlin, p.941).hALY:Grottadei Colombi?(Lambrecht,1933,Handb. l'aJamrn., p.731).ALASKA: bKodiak Island (Friedmann,1934,Jour. \Vashington Acad.Sci., vol. 24,p.234).WASIIINGTON:"PugetSound(L.Miller, 1960, Wilson Bull., vul. 72, p.394).NOVASCOTIA:'Whynaeht(Halifax Mus.). Recorded in error from Fossil Lake, Oregon(Sbufeldt,1892, Jour. Acad. nat. Sci. Philadelphia,vol.9,p.396;see Howard,1946, Publ.Carnegie Instn.\Vashington, no.. 5.51, Pl'.148,190).8. Aechmoplwfus occidentalis(Lawrence). \V ASHIKGTON: PugetSound(L.Miller, 1960, Wilson Bull., vol. 72, p.394).OIlEGON: Fossil Lake(AeehmopllOru8 lueasi L.Miller,Feb.4, 1911, Univ. Calif. Publ. Geol., vol. 6, no. 4,p.83, fig. 1-3; types tarsometatarslls, coracoid, femur,Univ. Calif. Paleo. nos. 1260312605).CALIFORNIA:RodeoandSan Pedro(L. .vlillcr, )912, Univ. Calif. Pnbl. Gcol., vol. 7, Pl'. 112,115); .vianix (Compton,1934, Condor, vol. 36, p.168);DelRey Hills(Howard, 1\)36, Condor, vol. 38, p.211);NewportBay(Howard,1949, Gondor, vol. 51, p.21);'Emeryville(Howard,1929, Dniv. Calif. Pnbl. Zoo!., vol. 32, p.329);"BuenaVistaLake(DeMay,1942, Condor,voL44, p.228),Specimens from Fossil Lake and someofthe Californian localities average large and are perhaps recognizable as a temporal subspecies, AechmophortlS occidentaIisluca."li L. Miller,9. Podilymbuspodiceps (1,innaeus). OR.EGON: Fossil Lake (Podilymbu."l magnusShufeldt,July9, 1913, Bull. Amer. Mus.nat.Hist., vol. 32, art. 6, pp. 136, 1.5.5 inpart,pI. 38,fig.439-440, 449 only; types two left tarsometatarsi, AMNH no.3574).CALIFORNIA'McKittrick(L. .vI iller, 1925, Univ. Calif. Pnbl. Ceol. Sci., vol. 15, p.307);RanchoLaBrea(Howard,1936, Condor, vol. 38, p.34);"BuenaVista Lake(DeMay,1942, Condor, vol. 44,p.228).NEVADA:Smith Creek Cave (Howard, 19.52, Bull.S.Calif. Acad. Sci., vol. 51, pt. 2, p. .54).AlUWNA: "GrandFalls(Hargrave, 19,19, Condor, vol. 41, p.207).TEXAS:Groesbeck Creek (MidwesternUniv.).ARKANSAS: bLake Texarkana(Southern 1fethodist Univ.).FLORIDA:SeminoleFieldand Itchtucknee River(Wetmore, 19.11, Smithsonian misc. Call., vol. 85, no. 2,1'.12);Haile (Broclkorb, 1953, Wilson Bull., vol.6.5,p.49);Reddick (Brodkorb, 1957, JOllr.Palmnt.,

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230BULLETINFLORIDASTATEMUSEUMVol. 7 vol. 31, p. 134);Arredondo(Bmdkorh, 1859, Bull.FloridaStateMus., vol. 4, p.273);RockSpring(Woolfenden,1959, Wilson Bull., vol. 71, p.185);VemBeach, stratum 2(Weigel,1963,Floridageol. Surv. Spec. Publ., no. 10, p.25); Sauta FeRiver (Brodkorb, 19fi3, Auk, vol. 80, p. 115);Jennys Spring,HornshySpring,andLake Monroe (Brodkorhcoll.);Bradenton(Univ.Florida);Bluffton, Good's!'Jhellpit,I1LemonBluff, and -ll-Silver Glen Springs (Neill, Gut, and Brodkorb, 1956, Atner. Antiquity, vol. 21, p.388); South Indian Field(\Veigel,1959, Florida Anthropologist, vol. 12, p.73).PUERTOHleo:"Barrio Canas (Wetmore,1938, Auk, vol. 55, p.53). MEXICO: nearTepexpan(Wetmore,1949, Condor, vol. ,'i1, p. 150).BUAZIL:LapadaEserivania(0.Winge, 1887, E Mus.Lund.,vol. 1, pt. 2, Pl'. 4, 2.5). Specimens from Fossil Lake andsomeof the Floridian localities average large and are perhaps recognizable as a tem poralSllbspccics,Podilymbus podiceps magnusShufeldt.

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196.3 BRODKORB:CATALOGUEOFFOSSILBIRDS 231OrderSPHENISCIFORMESSharpe Spen;"cifurmes Sharpe, 1891, Review of Recent AttemptstoClassify Birds, p. 71(typeSpheniscusBrissun).FamilySPHENISCIDAEBonaparte1SpheniscidaeBonaparte, 1831, Saggio di una distribuzionc metodica degliAnimali Vertehrati, p. 62(typeSpheniscusBrisson).SubfamilytPALAEEUDYPTINAF.SimpsonPalaeeudyptinaeSimpson, 1946 (Aug.8),Bull. Amer. Mus. nat. Hist., vol. 87, p. 69(type Palaeeudyptes Huxley).AnthropornithinaeSimpson, 1946 (Aug.8),Bull. Amer. Mlls. nat. Hist., vol. 87, p. ll9 (typeAnthrnpornisWiman).Genus tPlllllcclldyptes HuxleyPalaeeudyptesHuxley, 1859,Quart.Jour. geol. Soc.London,vol. 1.5, p.670(typebymonotypyPulueeudyptes antarclicusHuxley).1.Plllaeeudyptes marplesi Brodkorb2UPPEREOCENE(Burnsidemarl).NEWZEALAND:SOUTHISLAND:BurnsidenearDunedinin Otago.UPPEREOCENE(Transitional marlmemberof BlanchePointmarls).SOUTHAUSTRALIA:WittonBluff,atsouthendof Christie's Beach,16miles south of Adelaide (Pllilleeudyptesd.antarcticus Simpson,1957,Rec.S.Austr. Mus., vol.13,no.1,p.52,fig.1;S.Austr. Mus. no. P10870).2.Palaeeudyptes antllrctiCIlS HuxleyPalueeudyptes anturcticusHuxley, 1859, Quart. Jour. geol. Soc.London,vol. 15,p.670,fig.1-2(typefrom Kakanui, right tarsometatarsus, Brit. Mus. no.A.I048).LOWEROLIGOCENE(typeapparentlyfrom Kakanui limestone;othersfromtheyoungerMaerewhenuagreensand). NEW ZEALAND:SOUTHISLANll:KakanuinearOamaru(Huxley,1859);Duntroon,Earth-1Afragmentary femurofanunidentified penguin hasbeenrecorded fromtheLower Eocene Heretaungan stage at Gore Bay, Cheviot,NewZealand (Marples,1952, Pal. Bull., N. Zealand geol. Survey no. 20, p.51). species.Typefrom Burnside marl, left tarsometatarsus, Otago Mus. no. C.50.28; associated elements, Otago Mus. nos. C.50.25':'47; referred specimens, Otago Mus.nOs.C.48.73-81. TarsometatarsliS large and stout, with internal euge of shaft strongly concave(inP.antarcticlJstarsometatarsus smaller,withinternal edge nearly straight). Femur likewise large. Humerus short,withshaftsigmoid instead of straight.Ulnasmall. See Marples. 1952, Pal. Bull. N.Zealandgeol. Surv., no. 20,pp.31, 53, 55, 56, pI. 2, fig. 1; pl. 4, fig. 5; pI. 5; fig. 3, 6; pI. 8, fig.1,10,11.

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232BULLETINFLORIDASTATE MUSEUM Vol.'7quakesnearDuntroon,andSeal RocknearBrighton (Marples, 1952, Pal. Bull. N. Zealand geol. Surv., no. 20, p.28).MIDDLEOLIGOCENE(Burnsidegreensand).NEWZEALAND:SOUTHISLAND:BurnsidenearDunedin(Marples, 1952).MIDDLE?OLIGOCENE(Gambierlimestone).SOUTHAUSTRALIA: Pritch. ardBrothers' Quarry, 7J! milesWNWof Mt.Gambier(Palaeeudyptinae, gen.etsp. indct.,A,Simpson, 1957, Rec.S.Austral. Mus., vol. 13, p. 56, fig.3;S.Austral. Mus. no. P 10863); referral tentative. GenustPachydyptesOliver Pachydyptes Oliver, 1930, New Zealand Birds, p. 86(typePachydyptes ponder osus Oliver.)3.Pachydyptes ponderosusOliverPachydyptes ponderosus Oliver, 1930,N.Zealand Birds, p. 86, fig. (typefrom Fortification Hill, humerus, Dominion Mus.atWellington no.1450).UPPEREOCENE(Runanganstage).NEWZEALAND:SOUTHISLAND:FortificationHillnearOamaru(Oliver, 1930); Taylor'squarryat CormacksnearOamaru(Marples, 1952, N.Z.Geol. Surv. Pal. Bull.20,p.37).GenustArchaeospheniscus :\1arples ArchaeospheniscusMarples, 1952(May),Pal. Bull. N. Zealand geol. Surv.,no.20,p. (typebyoriginal designationArchaeospheniscus loweiMarples).4.Archaeospheniscus loweiMarples Archaeospheniscus loweiMarples, 1952(May),Pal. Bull. N. Zealand geol. Surv., no. 20, p. 40, pI. 2,fig.4; pI.4.fig.4(typeincomplete skeleton, Otago Mus. no. C.47.20).LOWER OLIGOCENE (Maerewhenuagreensand).NEWZEALAND:SOUTH ISLAN"D: DuntrooninNorthOtago.5.Archaeospheniscus lopdelliMarplesArchaeospheniscus lopdelliMarples, 1952(May),Pal. Bull. N. Zealandgeol.Surv., no. 20, p. 41, text-fig. 2,pI.3, fig. 9: pI. 4, fig 6; pI. 5, fig. 4; pI. 8.fig.5(typeposteranial skeleton, Otago Mus. no.C.47.2l).LOWEROLIGOCENE(Maercwhenuagreensand).NEWZEALAND:SOUTHISLAND:Duntroon. GenustDuntroonornisMarplesDuntroonomisMarples, 1952(May),Pal. Bull.N.Zealand geol. Surv., no. 20,p.42(typehyoriginal designationDuntroonornis parvusMarples).

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1963 BRODKORB:CATALOGUEOFFOSSIL BIRDS2336. Duntroonornis parousMarplesDuntroonomis parvus Marples, 1952(May),Pal. Bull.N.Zealand geol. SUN., no. 20, p. 42, pI. 8, fig.3-4(typeleft tarsometatarsus, Otago Mus. no. C.47.31).LOWEROLIGOCENE(Maerewhenuagreensand).NEW ZEALAND: SOUTHISLAND:Duntroon. GenustPlatydyptesMarplesPlatydyptesMarples, 1952(May),Pal. Bull.N.Zealand geol. SUN., no. 20, p. 37(typeby original designationPaehydyptes novaezealandiaeOliver).7.Platydyptes novaezealandiae(Oliver)Paehydyptes novaezealcmdiaeOliver. 1930,N.Zealand Birds, p. 86(typesfromOamaru district, humerus, radius, ulna, scapula, 2 vertebrae, Dominion Musr no.1451).LOWEROLIGOCENE(Maerewhenuagreensand).NEW ZEALAND: SOUTHISLAND:Duntroon(Marples,1952, Pal. Bull. N. ZealandgeotSurv.,no20, p. 38; needs confirmation, radiusandulnaonly). LowERi' OLIGOCENE(Wharekurilimestone?).NEWZEALAND:SOUTHISLAND:WaitakiValley?(notOamaruas labeledi', Marples, 1952).MIDDLE?OLIGOCENE(Waitakianstage?).NEWZEALAND:SOUTHISLAND:Oamarudistrict (Oliver, 1930).8.Platydyptes amiesi MarplesPlatydyptes amiai Marples, 1952(May),Pal. Bull.N. ZeahmJ geol. SUN., no. 20, p. 39, pI. 4, fig. 3: pI. 5,fig.5 (types from Hakataramea valley,humerus, radius, Otago no.C.50.61).MIDDLEOLIGOCENE(Waitaldanstage).NEWZEALAND:SOUTHISLAND:Hakatarameavalley inSouthCanterbury(Marples, 1952); White RocksnearDuntroon(Marples, 1952; possiblyDuntroonianage),GenustKororaMarplesKororaMarples, 1952(May),Pal. Bull.N.Zealand geol. Surv., no. 20, p.43(typeby original designation Karora oliveri Marples).9.Korora oliveri MarplesKorora oliveri Marples, 1952(May),Pal. Bull.N.Zealand geol. SUN., no.20p. 43, pl. 8,fig. 7-8 (typetarsometatarsus, Otago Mus. no. G.48.7).MIDDLEOLIGOCENE(\Vaitakianstage).NEW ZEALAND: SOUTHISLAND:lIakatarameavalleyinSouthCanterbury.

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234 BULLETINFLORIDASTATEMUSEUMVol. 7GenustAnthropodytesSimpsonAnthropodytesSimp,on, 1959(July23),Proc. Roy. Soc. Vicloria, vol.71,pt. 2, p. 113 (lype byoriginal designationAnthropodytes gilliSimpson).10. Anthropodytesgilli SimpsonAnthropodytes gilliSimpson, 1959 (July23),Proc. Roy. Soc. Vicloria, vol.71,pt. 2, p. 113,fig.I(typeright humerus, Nat. Mus. of Victoria, no. P171(7).LOWER?MIOCENE(Balcombianstage).AUSTRALIA:westernVictoria:southendofDevil'sDen,oneastbankofGlenelgRiver,northofDartmoor.GenustN otodyptesMarples !'iotodypte,Marples, 1953(JlIne),Scient. ReLlt.Falkland Is. Depeml.Sum, no. 5._ p. 11(type:byt>rigina\('p.sign
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1963 BRODKORB:CATALOGUEOFFOSSIL BIRDS 235LOWER \f JOCENE(SeymourIslandbeds).SEYMOURISLAND.GenustEosphaeniscusWimanEosphaeniscusWiman, 1905, Bull geol. Instn. Upsala, vol. 6, p. 280(typebymonotYIJYEosphaeniscus gunnari Wiman).EospheniscusAmeghino, 1905 (Nov.30),All. Mus.nacoBuenos Aires, vol. 13, pp.97,132,165(emendation).15.Eosphaeniscus gunnariWimanEosphaeniscus gunnariWiman,190,5,Bull. Ceol. lnst. Upsala, vol. 6, p. 280, pI. 12,fig.5(type right tarsometatarsus, UpsalaMus.).LOWERMIOCENE(SeymourIslandbeds).SEYMOUR ISU\.ND. GenustDelphimomisWimanDelphinomisWiman, 1905, Bull. geol. Instn. Upsala, vol. 6, p. 250(typebymonotypy Delphinornis larseniiWiman).16.Delphinornis larseniiWimanDelphinnrnis larseniiWiman, 1905, Bull. ;;;eol. lnstn. Upsala, vol. 6, p. 250, pI. 12.fig.1(typeleft tarsometatarsllS. UpsalaMus.).LOWERMIOCENE(SeymourIslandbeds).SEYMOURISLAND.GenustIchtyopteryxWimanIchtyopteryxWiman, 190.5, Bull. geol. InstIl. Upsala, vol. 6, p. 251(typebymonutypy]chtyopteryxgracilis\Viman).IchthllopteryxLambrecht. 1933,Handb.Palaeom., p. 231(emendation).17.Ichtyopteryx gracilisWimanIchtyopteryx gracilisWiman, 1905, Bull. geol. Instn. Upsala, vol. 6, p. 251, pl.12,fig.4(typedistal partofrighttarSOIlletatarslIs,UpsalaMus.). LOWEn MIOCENE(SeymourIslandbeds).SEYMOURISLAND.GenustArthrodytesAmeghinoArthrodytesAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, p. 143(typebyoriginal designationParaptenodytes grandisAmegbino).AnthrodytesSimpson, 1957 (Apr.30),Rec.S.Australian Mus., vol. 13, no.1,p.68(lapsusor misprinl forArthrodytes; only included species Anthrodytes?andrewsi[Ameghino]).18.Arthrodytes grandis(Ameghino) Puraptenuclytesgran
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236BULLETINFLORIDASTATEMUSEUMVol. 7LOWER MIOCENE (Juliensemember of Patagoniaformation). A!\CENTINA: TeL Santa Cruz: San Julian. SubfamilyfPALAEOSPHENISCINAESimpson Palaeosphenlscinae Simpson, ]946 (Aug. 8),Bull. Amer. Mus.nat.Hist., vol.87,art.1,p.69(typeP"laeospheniscusMorenoandMercerat).Genus i PalaeospheniscusMorenoandMercerat Paloeospheniscus MorenoandMercerat,1891(May),An. Mus.LaPlata,Pal. Arg., vol.1,pp.16, 29 (type PalaeospheniscU8 patagonicus andMercerat,designatedbyAmeghino,1891,Rev.argentinaHist.nat.,\01.1,p.447).Apterodytes Ameghino, 1901,An. Soc. cien.argentina,vol.,'51,p.81(typebymonotypyApterodytes ictus Ameghino). PreocclJpiedby ApterodytesJ. Hermann,1783,Tabl. Affin. An., p.235.PalaeoapterodytesAmeghino,1905(Nov.30),An. Mus. nacoRuenos Aires,vol.13.p.156(ne\vname for Aptemdytes Ameghino,becauseof similarity to"ApterodytaSoP.1786,"i.e.ApteroditaScopoh,1786,Deliciaefloraeetfaunae in>nbricae, pt. 2,p.91). 19.Palaeospheniscus graciliS Ameghino Palaeosphenlscus gracilisAmeghino,1899(July),SinOpsis geoI6gico-paleon tol6gica, Snplemento, p. 9 (type from GuaranitkodePatagonia," right tarsometatarsus, Allleghino colI.).-Ameghino,1905, An. Mus.nacoBuenosAires, vol.13,pp.111,163,pI.2, fig. 9(typefrom golfodeSanJorge, redescribed).Apterodytes ictus Ameghino,1901,An. Soc. cien.argentina,vol.,'51,p.81(typefromGolfodeSan Jorge, proximalhalfofrighthumerus,Ameghinocoil., perhapsnowin Buenos Aires Mus. ictus Ameghino, 190.'5, An. MilS. nacoBuenosAires, vol.13,pp.120, 164,pI.3, fig. 16(typeredeseribed). Palaeospheniscus medianusAmeghino, 1905(Nov.,30), An.:Mus.nae. Btlenos Aires, vol. 13,pp.108,162,pI.I, fig. 6 (type from Trt"lew, right tarsometa tarsus, Museo LaPlata).LOWERMIOCENE(Patagoniaformation, Juliense member)'! ARGENTINA:Ter.Chubut:Gulf of San JorgeandTrelew(Ameghino,1905).20.Palaeospheniscus patagonicus ;\iloreno andMerceratPolaeosphenisctls patagonicusMoreno and Merccrat, 1891(May),An. Mus.LaPlata,Pal. arg., vol.1,pp.16, 31; 1891(Aug.,'5),pI.1,fig.7-9, 12-13,lTypeofP.gracilis attributed to OligoceneDeseadofonnation,butaccordingto Simpson,1946,probablydriftfrom basal partof Patagoniano

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 237 15-16, 21,23, 27;1'1.2, fig. 5 (lectotype from Trelew, left tarsometatarsus,LaPlata Mus. no. 34, designatedbyAmegbino, 1891, Rev. argentina Hist. nat., vol. 1, p.447).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter.Chubut:Trelew.21.Palaeospheniscus menzbieriMorenoandMerceratPalaeosphenisclls menzbieriMoreno and Mercerat, 1891(May),An. Mus.LaPlata, Pal. arg., vol. 1, Pl'. 17, 33; 1891 (Aug.5),1'1.1,fig.3, 5-6, 10-11, 14,17, 22, 24;1'1.2, fig. 6 (lectotype from Ter.Chubut,right tarsometatarsus, Mus. La Plata no. 62, designatedbyAmeghino, 1891, Rev.argoHist.nat., vol. 1, p.447).Ameghino, 1905, An. Mus.nacoBuenos Aires, vol. 13, Pl'. 103, 162,1'1.1,fig.3(typefrom Trelewredescribed).Palaeospheniscus interruptusAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 104,162,1'1.1, fig. 4(typefrom Trelew, right tarsometatarsus, Mus. La Plata).[?JPalaeospheniscus planusAmcghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 109,163,1'1.1,fig.7;1'1.2,fig.7(typefrom GolfodeSanJorge, lefttarsometatarslls,coIl.Ameghino).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:TerChubut:Trelew; GolfodeSan Jorge.22.Palaeospheniscus rothiAmeghinoPalaeospheniscus rothiAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 110, 163,1'1.2,fig.8(typefrom Trelew, left tarsometatarsus,LaPlataMus.).Palaeospheniscus intermediusAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 113,163,1'1.2,fig.10(typefrom GolfodeSan Jorge, left tarsometatarsus, Ameghino coll.).Palaeospheniscus affinisAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 114,163,1'1.2,fig.11(typefrom Trelew, left tarsometatarsus,LaPlataMus.).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter.Chubut:Trelew; GolfodeSan Jorge. GenustPerispheniscusAmeghino Peri,<;pheniscus Ameghino, 1905(Nov.30),An. 1\1118. nacoBuenos Aires, vol.13, p. 117(typeby monotypyPerispheniscus wimaniAmeghino).TreleudytesAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, p. 156(typebymonotypyTreleudytes crassaAmeghino) .23. Perispheniscus robustus(Amcghino)Palaeospheniscus robustusAmcghino, 1895, Bol. Inst. gcog. argentina, vol. 15, p. 588,fig.1(typefrom Trelew, left humerus, Brit.Mus.).

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238BULLETINFLORIDASTATEMUSEUMVol. 7 Perispheniscus wimaniAmeghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, Pl'. 117, 164,1'1.2,fig.14; pI. 3,fig.14-15(typefrom "costasdePatagonia," left tarsometatarsus, La plata Mus.; referred humerus, Ameghinocall. ).Treleudytes crassaAmeghino, 1905 (Nov.30),An. Mus.nacoBucnos Aires, vol. 13, p. 156, text-fig. 4(typefrom Trelew, left tarsometatarsus,LaPlataMus.).LOWERMIOCENE(Patagonia formation, Juliensemember).ARGENTINA:Ter.Chubut:Trelew. Genus fParasphcniscus AmeghinoParaspeniscu.s Ameghino. 1905(Nov.30),An. 1\.1U5. nne. Buenos Aires, vul. 13,p. 115 (type byoriginal designationPalaeospheniscus bergiMorenoandMcrcerat).24.Paraspheniscus bergi(MorenoandMercerat) Palaedsphellis(:us bergl :\torcno and Mt'r<.:erat, 1891(May),An. Mus.La Plata, Pal. arg., vol.1,Pl'.18, 34; 1891 (Aug.5),1'1.1, fill;. 2, 4, 18-20, 25-26;1'1.2, fig. 7-8(lectotypefrom Trelew, left tarsometalarsus, LaPlata Mus., selected by Amell;hino, 1891, Rev. argentina Hist. nat., vol. 1,p.447).Para spheniscus bergi Ameghino, 1905, An. Mus. naco Buenos Aires, vol. 13, pp.115, 163,1'1. 2, fig. 12(typeredescribed).LOWERMIOCENE(Patagonia formation, Juliensemember).ARGENTINA:TerChubut:Trelew.25. Parasphenwcus twreius (Ameghino) Palaeosphenisctls nereiusAmeghino, 1901, An. Soc.den.argentina, vol. p. 81(typefrom GolfodeSan Jorge, left tarsometatarsus, Ameghinocall.). Para,spheniscus l1creius Ameghinu, 1905, An. nacoBuenosAires, vol.13, Pl'. 116, 163,1'1. 2, fig. 13(typereuescribed).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter.Chubut:GolfodeSan Jorge. SubfamilyfPARAPTENODYTINAESimpsonParaptenodytinaeSimpson, 1946 (Aug. Ill, Bull. Amer. Mus. nat. Hist., vol. 87, p.69(typeParaplenodytesAmeghino ).GenusfParaptenodytesAmeghino Paraptenodyte.< Ameghino, 1891(Dec.l),Rev.argentinaHist. nat., vol. 1,p.447(typebymOllotypy Palaeosplteniscus antarcticus Mureno and I\'fercerat) Metafl
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1963 BRODKORB;CATALOCUEOFFOSSIL BIRDS 239 tarsus,LaPlata Mus, nos, 2, 4,6,designated by Ameghino, 1905, p, 139),Paraptenodytes antarlictts [sic] Ameghino, 1891, Rev. argentina Hist. nat.,voL1, p, 447(lapsus),-Paraptenodytes antarcticusAmeghino, 1905, An, Mus, nac, Buenos Aires,voL13,pp,139, 166, pI. 5, fig, 32;pL6,fig,33-34(typesrestudied),LOWERMIOCENE(Patagoniaformation, Juliensemember).Al\GENTINA:TerSanta Cruz: mouth of Rio Santa Cruz. TeL Chubut:south side of Rfo Chubu, opposite Gaiman (Simpson, 1946, Bull. Amer. Mus. nat Rist,vol87,p.9).27.Paraptenodytes andrewsiAmeghinoParaptenodytes andrewsiAmeghino, 1901, An. Soc.den.argentina, vol. 51, p. 81 (types from San Julian, associated right humerus, right curacoid, proximal part of right scapula, Ameghino coIl., now perhaps in Buenos AiresMus.).Arthrodytes andrewsiAmeghino, 1905, An. Mus.nacoBuenos Aires, vol. 13,pp, 146,166, pI. 7, fig, 37; pI. 8, fig, 38-39 (typesrcdescribed), LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter. Santa Cruz: San Julian.28.Paraptenodytes curtusAmeghinoParaptenodytes canus Ameghino, 1901, An. Soc.den.argentina, vol. 51, p. 81 (type nomSan Julian, right tarsometatarslls, Ameghino coll.).-MetancydorniscurtusAmeghino, 1905, An. f\.'1us. nacoBuenos Aires, vol. 13, pp, 129, 16,5, pL4,fig, 2.5-26 (typeredescribed ).LOWER MIOCENE (Patagonia, formation, Tuliensemember).ARGENTINA:Ter. Santa Cruz: San Julian. GenustIsotl'emornisAmeghinolsotremornisAmeghino, 1905(Nov.30),An. :Mus. nae. Buenos Aires, vol. 13, p. 134(typebyoriginal designation lsotremo'rnis nordensk;oldiAmeghino).29.Isotremornis nordenskioeldiAmeghinoIsotremornis nordensk;oldi Amegbino, 1905 (Nov.30),An,Mus, nac, Buenos Aires, vol. 13, pp. 134,165, pL 4,fig.28 [fig, 28aisduplicated,theupperone representing this species]; pL5, fig, 29-31 (holotype from SanJulUm,proximal partofleft tarsomctatarsus, with associated right humerus, distalhalf of left humerus, distal half of left femur, Ameghino call., now perhapsin Buenos Aires the right humerus v."as designatedaslectotypebySimpson, 1946, p. 33,butthisaction appearsinvalid 'inviewofAmeghino'swording).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter. Santa Cruz: San Julian. GenustPscudospheniscusAmeghino Pseudospheniscl.ls Ameghino, 1905(Nov.30),An. nacoBuenos Aire:"!, vol.13, p, 123(typcbyoriginal designationPseudospheniscus interplanusAmeghino).

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240BULLETINFLORIDASTATEMUSEUMVol. 730.Pseudosphensicus inierplanusAmeghinoPseudospheniscus interplanu8 Ameghino, 1905(Nov. 30), An. nacoBuenos Aires, vol. 13,p.123(typedistal part of left tarsometatarsus, from San Julian, Ameghino coll., nowperhapsin Buenos AiresMus.). PseudosphenisCtM planusAmcghino, 1903 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, p. 164. pI. 3, fig. 19(sametype;asfirst reviser 1 select P.interp/anus) .LOWERMIOCENE(Patagoniaformation, Juliensemember).AHGENTINA:Ter. Santa Cruz: San Julian.31.PseudDspheniscusconcavU8Ameghino?Pseudospheniscus COflCat;Us Ameghino, 1905(Nov.30),An. Mus.nacoBuenosAires, vol.J3, p. 124(typefrom San ] lIlian, distalhalfof right tarsometatarsus, Ameghino colI., now perhaps in Buenos AiresMus.).Pseudosphenisctls cunvexusAmeghino,H105(Nov.30),An. r.,.fus. nacoBuenosAires, vol. 13, p. 164, pI. 3, fig. 20(some type: as first reviser 1 sclcct P. concavus).LOWERMIOCENE(Patagoniaformation, Juliensemember).ARGENTINA:Ter. Santa Cruz: San Julian. GenustN eculus ArneghinoNeeulusAmcghino, 1905 (Nov.30),An. Mus.nacoBuenos Aires, vol. 13, p. 127(typeby original designation Neculus rothi Ameghino).32.Neculus rothiAmeghinoNeeulus rothi Amcghino, 1905 (Nov.30),An. Mus.nacoBuenos Aircs, vol. 13, pp. 127, 165, pI. 4,fig.23(typefrom Trelew, distalpartof left tarsometa tarsus,LaPlataMus.). LOWEH MIOCENE(Patagoniaformation, Julicnse member). ARGENTINA:Ter.Chubut:Trelew. SubfamilySPHENISCINAE(Bonaparte) Spheniscidue Bonaparte, 1831, Saggiodiuna distribuzionc metodica dcgli Animali Vertcbrati, p. 62(typeSpheniseusBrisson).No extinct fossil species. N eospecies of Spheniscinae from Pleistocene sites:1.Eudyptes erestatus (J.F. \1ilIer). N>:w ZEALAND:Waikonaili(Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p.196).2. Megadyptes antipodes(Hombron .-mu Jacquinot).NEWZEALAND:Waikouaiti (Lydekker, 1891, p.195).3. Eudyptula minor (J.R.Forster).NEWZEALAND:Waikouaiti(Lydckker, 1891, p.197).

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDSOrderPROCELLARIIFORMESFiirbringer241Proeellariiformes Fiirbrin!/;er, 1888, Untersueh. Morph. Syst. Vogel, vol. 2, p.1544 (intermediate suhordo; typeProcellariaLinnaeus).FamilyDIOMEDEIDAE(Gray)Diomedeinae Gray, 1840, List Genera Birds, ed. 1, p. 78 (subfamily;typeDiomedeaLinnaeus).GenustGigantornisAndrewsGiganturnisAndrews, 1916, Proc. zool. Soc. London, p.,519(typebymonotypyGibantornis eaglesomeiAndrews). Position tentative.1.Gigantornis eaglesomeiAndrews Gigantorniseagles
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242BULLETINFLORIDASTATEMUSEUMVoL74.Diomedea anglicaLydekkerDinmedea anglicaLydekker,1891(Apr.'25),Cat.Foss. Birds Brit. Mus., p. 189, lip;. 42(typesfromRedCragat Foxhall, right larsomelalarsus, phalanx1 of toe IV,IpswichMus.; easls Brit. Mus. no.A.87).LOWERPLIOCENE(BoneValleyformation).FLORIDA:PolkCounty:Pierce(Wetmore,1943, Proc.NewEnglandzool.Club,vol. 22,p.66; identification uncertain, specimennotcomparabletotype).UPPERPLIOCENE(CorallineCrag).ENGLAND:Suffolk: Foxhall(Lydekker,1891, Ibis, p. 395; specimen notcomparablewithtype).LOWERPLEISTOCENE(RedCrag).ENGLAND:Suffolk: Foxhall(Lydekker,1891, Cat., p. 189). Neospecies ofDiomedeidaerecordedfrom Pleistoceneand pre historic sites:1.Diomedea exulansLinnaeus.ENGLAND:llford(Lambrecht,1933,Handb.palaeom., p.732).CUATHAMISLANDS(Lambrecht,1933, p. 273). 2.Diomedea albatrusPallas.JAPAN: Iki Island(Kuroda,1959, Bull. hiogeog. Soc.Japan,vol. 21, p.73). ALASKA:St. LawreDee Island(Friedmann,1934, Jour. Washington Aead. Sci.,voL24, p.87);"AmaknakIslandand"Kodiak Island(Friedmann,1934, op. cit.,pp.231, 2.34); 'Dntch Harbor,"LittleKiska,"AtkaIsland,and"Attn Island(Friedmann,1937, op. cit., voL 27,pp.432-437).OREGON:"Maxwell Point(Wetmore,1928, Condor, voL 30, p.191).CALIFORNIA:DelRey Hills(Howard,1936, Condor,voL38, p.212);NewportBayand?San Pedro(Howard, 1949, Condor,voL51, p.23);"Emeryville(Howard,1929, Univ. Calif. PubL ZooL, vol. 32, p.332).3. Diomedea nigripesAudubon.ALASKA:"Kodiak Island(Friedmann, 193.'5, Jour.WashingtonAcad. Sci., vol. 25, p.46).CALIFORNIA:San Pedro?(L. :\1iller, 1914, Univ. Calif. Publ. CeoL, vol. 8, p.34).4.Diomedea chlororhynchos Cmelin.NEWZEALAND:Waikouaiti(Lydekker,1891,Cat.Foss. Birds Brit. Mus., p.189).FamilyPROCELLARIIDAE(Boic)PrncellaridaeBoie, 1826, Isis von Oken, vol. 19, col.980(typeProcellariaLinnaeus).GenusPufjinusBrissonPufjinusBrisson, 1760, Omithologia, vol.1,p. 56(typebytautonomy Prncel laria pufjinusBriinnich).1.Pufjinus raemdonckii(VanBeneden)Larus raemdonckiiVan Beneden, 1871, Bull. Acad. Sci. Belgique, ser. 2, vol. 32, no.II,p. 258, lig. 1(lectotypefromRupelmonde,distalpartof lofthumerus,designatedbyBrodkorb, 1962, Auk, vol. 79, p.707).MIDDLEOLIGOCENE(Rupeliansand).BELGIUM:EastFlanders:mouthoftheRupel.ProvoAntwerp:Edeghem(VanBeneden,1871).

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDS2432.Puffinus arvernensis Milne-Edwards PuUinusarvetllensis Milne-Edwards, 1871, Ois. Foss. France, vol.2,p.572(nomennudum).-Milne-Edwards,in Shufcldt, Proc. Acad. nat. Sci. Phila delphia, p. SID, p!. 24,fig.1-2 unly(typefrom St.-Gerand-Ie-Puy, left tarsometatarsus, Paris Mus.). LOWERMIOCENE(Aquitanian). FRANCE: Dept.Allier: Saint Gerand-Ie-Puy.Therecord fromtheTortonianatGrive-St.-Alban (Shufeldt, 1896)mustrepresentsomeotherspecies or a mixing of localities. 3. Puffinus micraulaxBrodkorbPuffinus micraula" Brodkorb, 1963(in press), Quart. Jour. Florida Acad. Sci., vo!. 26, no. 2, p. 000,fig.0(typefrom Gainesville, distalpartof left humerus, Univ. Florida no.4872).LOWERMIOCENE(Hawthorneformation).FLORIDA:Alachua County: Gainesville.4.Puffinus aquitanicus(Milne-Edwards)Procellaria aquitanicaMilne-Edwards, 1874, Bib!. It-cole hautes Etudes Paris,sect. sci. nat.. vo!. 11, art. 3, p. 6, pI. 12, fig. 1(typedistalpartofhumerus).MIDDLEMIOCENE(Burdigalian). FI'ANCE: Dept.Gironde:FalunsdeSaucats.5.Puffinus antiquus(Milne-Edwards)Procellaria antiqua Milne-Edwards, 1874, Bib!. Ecole hautes Etudes Paris, sect. sci. nat., vul. 11, art. 3, p. 7(type proximal partofhumerus).MIDDLEMIOCENE(Burdigalian). Fl'ANCE: Dept.Gironde: FalunsdeSaueats.6.Puffinus conradiMarshPuffinu, conradiMarsh, 1870, Amer. Jour. Sci., ser. 2, vol. 49, no. 146, p.212(typedistalpartof left humerus, Acad. Nat. Sci. Phila. no. 13360: cast U.S.Nat. Mus.) .-Shufeldt,1915, Trans. Connecticut Acad. Arts Sci., vol. 19, p. 62, pI. 8,fig.63-64(typerestudied).-Wetmore,1926, Auk, vo!. 43, p. 46.3 (typerestudied).MIDDLE MIOCENE (Calvertformation).MARYLAND:CalvertCounty.7.Puffinus inceptoTWetmorePuffinu"inceptorWetmore, 19.30quly 1.5), Proc. California Acad. Sci., ser. 4, vol. 19, no. 8, p. 86,fig. 1-.3 (typefrom Sharktooth Hill, distalpartof righthumerus, Calif. Acad. Sci. no.5223).MIDDLEMIOCENE(Temblorformation),CALIFORNIA:KernCounty:Sharktooth Hill, 7 milesnortheastof Bakersfield.

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244BULLETINFLORIDASTATEMUSEUMVol. 78.Puffinus priscusL. MillerPuffinus priscusL.Miller, 1961(Oct.3),Condor, vol. 63, no.5,p.399, fig. 1 center(typefrom Sharktooth Hill, distalthirdof left humerus, Univ. Calif. Mus. Paleo. no.58185).MIDDLEMIOCENE(Temblorformation).CALIFORNIA:Kern County: Sharktooth Hill.9.Puffinus mitchelliL. MillerPuffinus mitchelliL.Miller, 1961 (Oct.3),Condor, vol. 63, no. 5, p. 400,fig.1 right(typefrom Sharktooth Hill, distal half of right humerus, Univ. Calif. Mus. Paleo. no. ,58184).MIDDLEMIOCENE(Temblorformation).CALIFORNIA:Kern County: Sharktooth Hill.10.Puffinus diatomicusL. MillerPuffinus diatamicusL. Miller, 1925(Aug.),Puhl. Carnegie Inst. Washington,no. 349, p.Ill,pI.1-2,7a(typefrom near Lompoc, skeleton impression,Uuiv. Calif. Mus. Paleo. no.26541).UPPERMIOCENE(Valmonte diatomitememberof Montereyshale).CALIFORNIA:Los Angeles County: San Pedro breakwater (L. Miller, 1935, Univ. Calif. Los Angeles Publ. bioI. Sci., vol.1,p.74);Lomita(Howard,1955, Los Angeles County Mus., sci. ser. no. 17, p.14);Sherman Oaks(Howard,1962, Condor, vol. 64, p. 512).UPPERMIOCENE(basal1000 feet of Sisquoc formation).CALIFORNIA:Santa Barbara County: Johns Mansville "Celite" mines, 3Jf miles south of Lompoc(L.Miller, 1925).11.Puffinus felthamiHowardPuffinu8 felthamiHoward, 1949 (June 22),Publ. Carnegie Instn. Washington, no. 584, p. 194,pI.2,fig.4, 6(typedistalpartof right humerus, Los Angeles Mus. uo.2037).LOWERPLIOCENE(Repettoformation).CALIFORNIA:Orange County: 3 milesnorthof Corona del Mar.12.Puffinus kanakaffi HowardPuffinus kanakoffiHoward, 1949 (June 22),Publ. Carnegie Instn. Washington, no. 584, pp. 187, 195 note, pI. 2,fig.3, 5(typetarsomet.tarsus, Los Angeles Mus. no.2122).MIDDLEPLIOCENE(San Diego formation).CALIFORNIA:San Diego County: San Diego.

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 245GenusfArgyrodyptesAmeghino Argyrodyptes Amcghino, 1905(Nov.30),An.Mus.nacoBuenos Aires, vol. 13, p. 121(typebyoriginal designationArgyrndyptes microtar,msAmeghino).ArgyrodytesTrouessart, 1906, Rev. crit. Paleozoo!., vol. 10,pp.90, 251 (emendation).13, Argyrorlyytes microtarsusAmeghinoArgyrodyptes microtarsusAmeghino, 1905 (Nov.30),An. Mus.naCoBuenos Aires, vol. 13, pp. 121, 161, pI. 3,fig.17-18 (lectotype bypresent dcsignation distal partofleft tihiotar.'Slts, Ameghino coIl., with associated distal part of right femur,llOW perhllp.o; in Buenos Aires :Mm;,). LOWERMIOCENE(Patagonia formation, JuHensemember),ARGENTINA:Ter. Santa Cruz: Rio SeeoatSan Julian. GenusfP!otornisMilne-EdwardsPlotornisMilne-Edwards, 1878, Bibl. Ecole hautesEtudesParis, sect. sci. not.. vol. 11, .rl. 3,pp.4 .. 5 (typeby monotypy Pl%rni.delfortriiMilne EJw."ds). 14. Plotornis delfortriiMilne-EdwardsPlotamis delfortrii Mihle-Edwards, 1878, Bibl. Ecoleh.utesEtudcs Paris, sect. sci. nat.,voL11,art.3, [Jp.,1-5, pI.11 (types tarsornetatarstls and distal pttrt of hunwTus). MIDDLEMIOCENE(MolasseedeLeognan).FRANCE:Dept. Gironde: Leognan near Bordeaux'! Neospecies of Procetlariidae from PleistoceneandprehistOTic sites:1.Macrunec/es giganteus(Gmelin).NEWZEALAND:Waingongoro(Lydekker,1891, Cat. Foss. Birds Brit. Mm" p.187).2.P!llmarus glacialisNORWAY'Va,do (Lamhrecht, 1933, Handb. PaIRP-OTrJ., p.732).ALASKA: I) St. Lawrence Island (Friedmann, 1934,Jour.Washington Acad, Sci., vol. 24, p.87);"Kodiak Island(Friedmann,1934, op. cit., p.234);"DutchHarhorand Attu Island(Fricdmann,1937, op. cit.,pp. 43,<;, 438). C.rIFOHNIA: San Pedro(L.Miller, 1914, Univ. Calif. Pnb!. Geol., \'OJ. 8, p. 35); Newport Bay (How.,d, 1949, Condor. vol. 51, p. 21).3.PutJinus IClIcunwlas (Temminek).JAPAN:rkiIsland (Kuroda, 1959, Bull. biogeog. Soc. Japan, vol. 21, p.73).4.PutJinus diomedea(Scopoli).PORTUGAL:Crotte de Furinha(Lambrecht,1933, Handb. p.732). GIHRALTAN: Devils Tower (Bate,1928, Jour.Roy. anthrop. Inst., vol. 58,p.104). SAtillINrA: Grottu Pietro TampQn} on TavQ1Fltlmarus, sp. indet., recorded from Middle Miocene Calvert formation atChesapeake Beach, Maryland (Wetmore,1926, Auk, vol. 4.3, p.464).

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246BULLETINFLORIDASTATEMUSEUMVol. 7laraIsland(PufJinus eyermaniShufeldt, 1896, Proc. Acad. nat. Sci. Philadelphia, p. 511,pI.24, fig. 3-4,8;types right tibiotarsus, right tarsometatarsns, nowU.S.Nat. Mus. no.2166).5. PufJinus gravis(O'Reilly).NOVASCOTIA:Reid site (HalifaxMus.).6. Puffinus griseus (Gmelin)'.SARDINIA:GroUa Pietro Tamponi(Regalia, 1897, Avicula, vol. 1, p.165);Monte Giovanni?(Lambrecht,1933,Handb.Palacorn., p.732). ALASKA: 'DutchHarbor,'LittleKiska,and'AttuIsland(Friedmann,1937, Jour. Washington Acad. Sci., vol. 27, pp.435-438).7.PufJinus tenuirostris(Temminck).ALASKA:'St.LawrenceIsland and'AmaknakIsland(Friedmann,1934, Jour. Washington Acad. Sci., vol 24, pp. 87,231);'CapcPrince of Wales(Friedmann,1941, op. cit., vol. 31,p.405).8.PufJinus pufJinus(Briinnich).DENMARK:'OrdrupMose(Oestrelatasp.,H.Winge, 1903, Vidcnsk. Mcddel. naturhist. Foren. Copenhagen, vol. 6, p.92).GmRALTAR:Devils Tower (Bate, 1928, Jour. Roy. anthrop. Inst., vol.. 58, p.104).SAIIDINIA:Grotta Pietro Tamponi?(Lambrecht,1933,Handb.Pal aeorn., p.732).ITALY:Buca dcl Bersagliere(Lamhrecht,1933).BERMUDA:'caves(PufJinus mcgalliShufeldt, Oct. 1916, Ibis, p. 630;typcsternum; Brit. Mus.?).BAHAMAS:'GordonHills on Crooked Island(Wetmore,1938, Auk, vol. 55, p.51).FLORIDA:Melhourne(Wetmore,1931, Smithsonian misc. Coil., vol. 85, no. 2, p.13).CALIFORNIA:SanPedro(L.Miller, 1914, Univ. Calif. Publ. Geol., vol. 8, p.35);Del Rey Hills(Howard,1936, Condor, vol. 38, p.212);Newport Bay?(Howard,1949, Condor, vol. 51, p.21).Includes Puffinus opisthomelasCones.9.Puffinus lherminieriLesson.BERMUDA:'caves(Puffinus pamus Shufeldt,Oct. 1916, Ibis,p.632; types miscellaneous elements; Brit. Mus.?); Rail Cave, (tShearwater Cave, and #Castle Harbour Islands (Brodkorb coIL); Cockroach Island(Wetmore,1962, Smithsonian misc. CoIl., vol. 145, no. 2, p.15).BAHAMAS:'GordonHills on Crooked Island(Wetmore,1938, Auk, vol. 55, p.51).ST.THOMAS:"midden(Wetmore,1918, Proc.U.S.nat. Mus., vol. 54, p.514).ST.CROIX:'Concordia(Wetmore,1937, Jour. Agr. Univ. Pucrto Rico, vol. 21, p.6).BARBUDA:caves(Univ.Florida).ANTIGUA:'MillReefmidden(Univ.Florida).10.Pierodroma cahow(NicholsandMowbray).BERMUDA:'caves(Aestrelata vociferans Shufeldt, Oct. 1916, Ibis, p. 633; practically a nomennudumexcept forgeneric characters; types Brit. Mus.?); vVilkinson Quarry and Cockroach Island(Wetmore,1960, Smithsonian misc. ColI., vol. 145, no. 2, p.16);Rail Cave, Crane Crevice,andWilson's Cave (Brudkorb coll.).BAHAMAS:"GordonHillsonCrooked Island(Wetmore,1938, Auk, vol..55,p.51).11.Pterodroma hasitata(Kuhl).MARTINIQUE:"Paquemar(Wetmore,_1952, Auk, vol. 69, p.460).FamilyOCEANITIDAB(Salvin)OceanitinaeSalvin, 1896, Cat. Birds Brit. Mus., vol. 25, pp. xiv, 343, 358(subfamily; OceanitesKeyserling and Blasius).HydrobatidaeMathews, 1912, Birds Australia, vol. 2, p. 9(typcHydrobatesBoie).Family name preoccupiedbyHydrobatidaeGray, 1869, Hand-list Gen. Sp. Birds, pt. 1, p. 266(typeHydrobataVieillot, 1816, a junior synonym of Cind"s Bechstcin, 1802)andantedatedbyOceanitinaeSalvin.

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDSGenusOceanodromaReichenbach247Oceanodroma,Reichenbach, 1852, Avium systema naturale1p.iv(typebyoriginal designationProcellariafHreataGmelin).1.Oceanodroma: hubbsiL.MillerOceanodroma hubbsi L. Miller, 1951(March27),Condor, vol. 53, no. 2,p.78, fil(. I(typeskull, vertebrae, pelvis, Jeft leg, Univ. Calif. Mus. Paleo. no.39979).UPPERMIOCENE(Capistrano formation).CALIFORNIA:OrangeCounty: 1 mile south of Capistrano Beach. Neospecies of Oceanitidae fromtheQuarternary:1.Oceanodromahombyi(Gray).CHILE:Toeopilla (Strescmann, 1924, Ornith. Monatsber., p.61).FamilyPELECANOIDIDAE(Gray)HalodrominaeBonaparte, 18156, C.R.Acad. Sci. Paris, vol. 37, no. 18, p. 643(typeHalodromaIlliger,18Il,a junior synonym ofPelecanoides Lacepede, 1800).PelecanoidinaeGray, 1871, Hand-list GeneraandSpecies of Birds, pt. 3, pp. x, 102(typePelecarlOides Lacepl:dc). Neospecies of Pelecanoididac fromthePleistocene.1.Pelecanoides gamotii (Lesson). PERU: IslasdeLohosde Afucra(Clarke,1882, Proc. philos. Soc. Glasgow, vol. 1.3, p.573).

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248BULLETINFLORIDASTATEMUSEUMVol. 7OrderPELECANIFORMESSharpePelecaniformesSharpe, 1891, ReviewofRecent Attempts to Classify Birds, p.76(typePelecanusLinnaeus).SuborderSULAESharpe Suu,e Sharpe, 1891, ReviewofRecent AttemptstoClassify Birds, p. 76 (typeSula Brisson). Family j ELOPTERYGIDAELambrechtElopterygidueLambrecht, 1933,HandbuchPalacom., p. 287(typeElopteryxAndrews)GenustElopteryxAndrewsElopteryxAndrews, 1913(May),Geol. Mag., n.s., decade 5, vol. 10, no. 5, p. 195 (typebymonotypyElopteryx nopcsaiAndrews) .1.Elopteryx nopcsaiAndrewsElopteryx nopcsai Andrews, 1913(May),Gool. Mag., n.s.,decade5, vol. 10, no. 5, p. 195,fig.1-2(typefrom Szentpeterfalva, proximalpartof left femur, Brit.Mus.).UPPERCRETACEOUS,limestone).RUMANIA:(=Hateg). ( TransylvanianfreshwaterTransylvania: SzentpeterfalvancarHatszegGenustArgillornisOwenMegaloruis Seeley, 1866, Ann. Mag. nat. Hist., sar. 3, vol. 18, p. 110(typebymonotypyLlthornis emuinus Dowerhank).Preoccupieuby Gray, 1841, List. Gen. Birds, cd. 2, p.85).ArgilloruisOwen, 1878, Quart. Jour. geol. Soc. London, vol. 34, p. 124(typeArgillortlis lungipennis Owen). 2.Argillornis emuinus(Bowerbank) Lithomi" emuinusBowerbank, 1854, Ann. Mag. nat. Hist., ser. 2, vol. 14, p.26.1,fig.(typefrom Sheppcy, tibiotarsus" ==shaft of humerus, Brit. Mus. no.38941).Lithomis emuianusSeeley, 1866, Ann. Mag. nat. Hist., ser. 3, vol. 18, p. 110(Iapsllsoremendation). Argillornls1011gipnmis Owen, 1878, Quart. Jour. geol. Soc. Lonuun, vol. 34,p. 124, pI. 6, fig. 1-3, 7-12, 16(typesfrom Sheppey, fragments of associatedrightandleft humeri, Brit. Mns. nos. A.5-9).Argillornis longipc. "Sharpe," LamlJrecht, 19.33,Handb.Palaeorn., p. 282 (lapsus; spelled correctly in Sharpe, 1899, Hand-list, vol. 1, p.240).LOWEREOCENE(Londonelay).ENGLAND:Kent:SheppeyIsleatmouthofThames(Bowerbank,1854):Eastchureh(Seeley,1866).

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 249MIDDLEEOCENE(Bruxellian).BELGIUM:EtterbeeknearBrussels(Dalla,1909,Ann. N.Y.Acad. Sci., vol.19,no.4,p.HI).Needsrestudy.GenusfEostegaLambrechtEostegaLambrecht,1929,C.R.X Congr. internat. Zool.Budapest1927, p. 1272(typebymonotypyEostega lebedinskyiLambrecht).s.Eostega lehedinskyiLambrecht Eostega lebedinskyiLambrecht, 1929, C.R.X Congr. internat. Zool.Budapest1927, p. 1272,fig.12-13(typefrom Kolozsmonostor, mandible,WienerNaturhistorische Hofsmuseum).MIDDLE EOC'ENE (SteinbruchGrobkalk).HUMANIA:Transylvania:Kolozsmonostornear Kolozsvar (Kluj),intheSiebenbiirgen.FamilyPHALACHOCORACIDAF.(Bonaparte)PhalaeroeoraeeaeBonaparte, 1853, C. R. Acad. Sci. Paris, vol. 37, no. 18, p. 643(typePhalaeroeoroxBrissou).Phalacrocoracinae Bonaparte, 1854, Ann. Sci. nat.(Paris),vol. 1, p. 38. SubfamilyfGRACULAVINAEFiirbringer Graculavinae Fiirhringer, 1888, Untersuch. Morph.,Syst.Vogel, vol. 2, p.1565footnote(type Graeulovus Marsh).GenusfGraculavusMarsh Groeulavu" Marsh, 1872, Amcr. Jour. Sci., scr. 3, vol. 3, p.363(type Grocukwus veloxMarsh,by gen. etsp. nov. convention, andbydesignation of Hay,1902,Bull. U.S.Geol. Surv., no. 179, p..533). Limosads Shufeldt,191.5(Feh.),Trans. Connecticut Acad. Arts Sci., vol. 19, p. 19(substitutenameforGroeulavusMarsh, consideredinappropriate).1.Graculavus veloxMarsh Graeulavus velox Marsh, 1872, Amer. Jour. Sci., ser. 3, vol. 3, p.363(typefrom Hornerstown, proximal endofleft hurneruf::i, Yale Peabody Mus. no. 855).UPPERPALEOCENE(Hornerstownmarl).NEWJERSEY:OceanCounty:Hornerstown.2.Graculavl/.s pumilusMarshGraculavus pumilusMarsh, 1872, Am. Jour. Sci., ser. 3, vol. 3, p.364(lectotypefrom Hornerstown, "distal" [proximal]endof right humerus, YalePeabody :-01us. no. 1209, designatedbyShufeldt, 1915,wheresaid tobefrom Battle Creek, Kansas!).

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250BULLETIN FT"OnIDA STATE MUSEUM Vol. 7UPPERPALEOCENE(Hornerstownmarl).NEWJERSEY:OceanCounty:Hornerstown.SubfamilyPHALACROCORACINAEBonapartePhalaeroeoraeinae Bonaparte, 1854, Ann. Sci. nat.(Paris),vol. 1, p. 38(typcPhalaeroeorax Brisson).GenusfActiornisLydekkerActiorni" Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 56(typebyoriginal designation Actiorni.'angliC1J" Lydekker) .3. Actiornis anglicusLydekkerAetiorni" anglieu" Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus.,p.56,fig.13(typefrom Honlwell, proximalpartof right ulna, Brit. Mus.nO.30328).MIDDLEEOCENE(Hordwellbeds).ENGLAND:Hampshire:Hordwell.GenusPhalacrocoraxBrissonPhalaeroeorax Bri"on, 1760, Ornithologia, vol.1,p.60(typePeleean .... carboLinnaeus).Oligoeorax Lambrecht, 1933, Hamlb. Palaeorn., p. 290(typeGrae"l"s littorali. Milne-Edwards, designatedbyBrodkorb, 1952, Condor, vol. 54, p.175).Mioeorax Lambrccht, 1933,Handb.Palaeorn., p. 291(typePhalaeroeorax femor alis Miller, designatcd by Brodkorb,1952).Paracorax Lambrecht, 1933,Handh.Palaeorn., p. 292 (typePhal.ocroeorax destefanii Regalia, designatedbyBrodkorb, 19.52). A""traloeorax Lamhrecht, 1933,Handb.Palaearn., p. 292(typePholaeroeorax oetustus DeVis, designatedbyBrodkorL,1952).4.Phalacrocorax mediterraneusShufeldtPhalaeroenrax mediterraneus Shnfeldt, 1915(Feb.),Trans. Connecticut Aead. Arts Sci., vol. 19,p.58,pI.15,fig. 1:18 (typefrom Gerry's Ranch, proximalpartof ri!';ht carpometacarpus, Yale Peabody "!us. no. 94.3).LOWERorMIDDLEOLIGOCENE(WhiteRiverformation).COLORADO:WeldCounty:Gerry'sRanchatChalkBluffs,Township11North,Range64West.5.Phalacrocorax littoralis(Milne-Edwards)Graenlus littorali., Milne-Edwards, 1863(afterJune C. R. Acad. Sci. Paris, vol. 56, p. 1222(typefrom Dept. Allier: almost a nomennudum). -Milne-Edwards, 1867, Dis, Foss. France, vol.1,sheet3.3,p. 263, pI. 42,fig.5-12: pI. 43,fig.1-7; pI. 44,fig.1-8(typescomplete left coracoid from in colI. Poirrier; complete righthumemsfromBillyin colI.Milne-Edwards;andfrom Dept. Allier in call. Milne-Edwards proximalpartofleft metatarsus, proximal two-thirds of left ulna, proximalpartof left femur,anddistal eod of left tibia: all fi!';ured hiltthe la>t twonotdescribed),

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1963 BRODKORB: CATALOCUEOFFOSSILBIRDS 251SaintLangy LOWER MIOCENE(Aquitanian).FRANCE:Dept. Allier: Pour9ain andBilly (Milne-Edwards, 1867); Vaumasand(Paris, 1912, Rev. fran9aise Ornith., vol. 4, p. 289).LOWERMIOCENE(Hydrobienkalk?).GERMANY:harbor constructionatFrankfort am Main (Lambrecht, 1933, Handb. Palaeorn., p. 290).6.Phalacrocorax miocaenus(Milne-Edwards)Graculus miocaenusMilne-Edwards, 1867, Gis. Foss. France, vol.I,sheet32,p. 255, pI. 39, fig. 5-18; pI. 40-41; pI. 42,fig.1-4 (types from Langy, lefttarsometatarstis, left tibia, right femur, pelvis, sternum, upper fragmentoffurclllnm, left coracoid, several scapulae, right humerus, right ulna, radius,earpometaearpus, alar digitsII-IandIII,eoll.Milne-Edward.).LOWERMIOCENE(Aquitanian).FRANCE:Dept. Allier:Langy(Milne-Edwards, 1867); Vaumas (Paris, 1912, Rev. Fran9, Orn., vol.4,p. 289); St.-Gerand-le-PuyandMontaigu (Lambrecht, 1933, Handb. Palaeorn., p. 290).7.Phalacrocorax subvolans BrodkorbPhalacrocorax ,,,buolansBrodkorb, 1956 (Sept. 24),Condor, vul. 58, no.5,p.367,fig.1(typefromThomas Farm, proximal part of right humerus, Univ.Florida no.4500).LOWERMIOCENE(1nomasFarmlocal fauna, Hawthorneage).FLOJUDA:Gilchrist County: Thomas Farm, 8 miles north of Bell.8.Phalacrocorax marinavisShufeldtPhalacrocorax mar'inavis Shufeldt, 1915 (Feb.), Trans. Connecticut Acad. Arts Sci., vol. 19, p. 56, pI. 14,fig.114, 116-118, 122 (types from Willow Creek, distal parts uf 2 humeri, distalpartof left tarsometatarsus, proximal halfofright ulna, Yale Peabody Mus. no.936).LOWERMIOCENE(JohnDayformation).OREGON:Malheur County: Willow Creek.9.Phalacrocorax intermedius(Milne-Edwards)Graculus intermediusMilne-Edwards, 1867, Gis. Foss. France, vol. 1, sheet 34" p. 266, pI. 43,fig.8-11(typefrom Orleanais, proximalpartof right humerus,coll. Nouel,nowapparently in Paris Mus.)UPPERMIOCENE(FalunsdeTouraine).France:provoOrleanais.10.Phalacrocorax praecarboAmmonPhalacrocorax praecarbovon Ammon. 1918, Abh. naturwiss. Ver. Regensburg,.vol. 12, p. 28,fig.3 (type from Mayer and Reinhard clayworks,upperendof left coracoid, Mus. Naturw. Vereins zu Regenshurg).

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252BULLETINFLORIDASTATE MUSEUM Vol. 7UPPERMIOCENE(BraunkohlenderOberpfalz).BAVARIA:clayworks ofMayerandReinhard,betweenDeehbettenandPriifeningnearRegensburg.11.Phalacrocorax femoralisL. PhaWCTOcoraxlemuralis L. Miller, 1\)29 (July15),Condor. vol. 31, no. 4, p. 167, fig. 58-59(typefrom Poyer quarry, skeleton impression, Univ. Calif.atLos Angeles, reverse in call. Dr. Frederick Kellogg, Los Angeles).UPPERMIOCENE(Modeloformation).CALIFORNIA:Los AngelesCounty:quarryofDanJ.Poyer,inNW of secti0I!18,Township1 North,Range17 West,nearCalabasas.12.Phalacrocorax wetmoreiBrodkorbPhalaeTOcorax wotmorei Brodkorb, 1955 (Nov.30), Florida Geol. Surv. Rept. Invest., no. 14, p. 12, pI. 3, fig. 10-11(typefrom Brewster, right coracoid, Brodkerb coli. no. PB530).LOWERPLIOCENE(BoneValleygravel).FLORIDA:PolkCounty:BrewsterandPierce (Rrodkorb, 1955).LOWERPLIOCENE(Alachuaclay).FLORIDA:AlachuaCounty:nearNewberry(Brodkorb, 1963, Spec. Pub!.Floridageol. Surv., no. 2,paper4, p.2).13.Phalacrocurax leptopus llrodkorbPhalncrocorax leptopus Brodkorb, 1961 (Nov.7),Quart. Jour. Florida Acad. Sci., vol. 24, no. 3, p. 170,fig.1(typefrom Juntura, proximal half of lefttarsometatarsus, Univ. Ore. Mns. Nat. Hist. no. F-7904). LOWERandMIDDLEPLIOCENE(Junturabeds).OREGON:MalheurCounty:Juntura.14.Phalacrocorax kennelliHoward Phalncrocorax kennelliHoward, 1949(June22),Publ. Carnegie Instn. Wash ington, no. 584, p. 188,p1.3. fig. 7-8(typefrom locality 1080,upperhalf of left coracoid, Los Angeles Mus. no.2127).MIDDLEPLIOCENE(SanDiegoformation).CALIFORNIA:San DiegoCounty:San Diego, locality 1080 onWashingtonBoulevard freeway.15.Phalacrocorax des/cfani negalia PhalacrocoraxdestefaniRegalia, 1902, Palaeontogr. ital., vol. 8, p. 225, pI. 27,fig.4-14(types from Orciano Pisano, cervical vertebra, furculum, coracoid, humerus. ulna, femur, tibiotarsus, tarson1etatarsus, Roberto Lawley col!., ondeposit in GabinettodiGeologia, Istituto di Studi SuperioriatFlorence).

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS 253MIDDLEPLIOCENE(argillemarine), ITALY: ProvinciadiPisa: Orciano Pisano near Valle di Fine.16. Pha1acrocorax idahenbis (Marsh)Graculus idahensisMarsh, 1870, Amer. Jour, Sci., ser. 2, vol. 49, p. 216(typefrom Castle Creek, proximal half of left carpometacarpus, Yale Peabody Mus. no. 527).-Phalacrocorax idahensis Shufeldt, 1915, Trans. Connecticut Acad. Arts Sci., vol. 19, p. 68, pI. 6,fig.44(typerestudied).MIDDLEPLIOCENE(ChalkHills formation).IDAHO:Owyhee County: Castle Creek,about10miles northwest ofGrandView, Referred specimens fromtheLower Pleistocene Glenns Ferry formation intheHagerman lake beds,Idaho(Wetmore, 1933, Smithsonian misc. ColI., vol. 87, no. 20, p.5),andfromtheLower Pliocene Bone Valley gravelatBrewster, Florida (Brodkorb, 1955, Florida geol. Surv. Rept. Invest" no, 14, p.14),are not comparable tothetypeandprobably represent other species.17. Pha1acrocorax macerBrodkorbPhalacrocorax macerRrodkorb, 1958(Oct.31),Wilson Bull., vol. 70, no. 3, p. 237,fig.1(typefrom sec. 28, right carpomctacarpus, Univ. Mich. Mus. Paleo. no. 33918).LoWERPLEISTOCENE(Hagermanlake beds of Glenns Ferry forma tion).IDAHO:TwinFalls County: section 28, Township 7 South, Hange 18 East, opposite Hagerman.18.Phalacrocorax rogersiHowardPholaeroenrax rogersiHoward, 1932(May16),Condor, vol. 34, no. 3, p. 118,fig.19(typefrom Veronica Springs,leftcoracoid, Santa Barbara Mus. no.32.1).LOWERPLEISTOCENE(Santa Barbara formation).CALIFORNIA:Santa Barbara County: Veronica Springs stone quarry.19.Phalacrocorax macropus(Cope)Graeulus maeropW! Cope, 1878, Bull. U,S.geol. geog. Surv. Terrs., vol. 4, no. 2, p. 386 (lectotype tarsometatarsus,Am.Mus. Nat. Hist, no. 3555, selectedbyHoward, 1946, Publ. Carnegie Instn, Washington, no.5,51,p,153).MIDDLEPLEISTOCENE(Fossil Lake formation),OREGON:Lake County: Fossil Lake.Thespecimen reported fromtheLower Miocene Arikaree sandstone of Montana (Shufeldt, 1915, Auk, vol. 32, p. 485)isunidentifiable even to family(d.Wetmore, 1955, Condor, vol. 57,p.371).

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254BULLETINFLORIDASTATEMUSEUMVol. 720.Phalacrocorax pampeanusMorenoandMerceratPhalacrocorux pampeallusMorello alld Mercerat, 1891(May), An.Mus. LaPlata, Pal. arg., vol. 1,pp.19,3,5;1891 (Aug.5),pI. 18, fig. 8(typefrom Lujall, proximalpartofrighthumerus,LaPlata Mus.110.82). UPPER PLEISTOCENE(Pampasformation).ARGENTINA:Provo Buenos Aires: LUjan.21.Phalacrocorax gregoriiDeVisPlwlacrocorax gregoriiDeVis, 1906, Ann. Queensland Mus., no. 6,p.18,pI.,5,fig. 6; pI. 6,fig.3-5; pI. 7, fig. 1-4; pI. 8, fi!':. 1-2(typesfrom variouslocalities near Lake Eyre, premaxilla) fragments of 2 curacoids. 7 humeri, carpometacarpus,7femora,4tibiotarsi, 6 tarsometatarsi, 7pelves).UPPER PLEISTOCENE (Katipiri sands, Malkunifauna).SOUTHAUSTRALIA:\Vankameminna; Malkuni; Kalamurina;Wurdumankula;Wurdulumankula;Mulcani, all on lowerCoopernearLakeEyre.22.Phalacrocorax vetustusDeVisPhalacrocorax 1)elustu8 DeVis, 1906, Ann. Queensland !\'lus., no. 6,p.22,pI.8, fig. 3-7; pI. 9, fig. 1-5, 7(typesfrom localities near Lake Eyre, fragmentsof3 coracoids, 7 humeri, 2 ulnae, 4 carpometacarpi, 2 femora, 2 tibiae, 1 tarsometatarsus) UPPER PLEISTOCENE(Katipiri sands, Malkuni fauna).SOUTIIAUSTRALIA:Malkuni; Kalamurina;Wurdumankula;\Vurdumulankula, allonlower CoopernearLakeEyre. GenusfPliocarboTugarinov Pliocarbo Tugarinov, 1940, Doklady Akad.Nauk S.S.S.H.., vol. 26,110.2, p. 197(typePliocarbo lOllgipesTugarinov).23.Pliocarbo longipesTugarinovPliocarbolOllgil'e" Tugarino\', DokladyAbd.NaukS.S.S.R., vol. 26,110.2,p. 197, fig. 1-2(typefrom Slnhodka,tarsometatarus).LOWERPLIOCENE(Meotianstage). UKRAINE: SlobodkancarOdessa. Neospecies of Phalacroeoracidae from Pleistoceneand"prehistoric sites;1.Phalacrocorax auritus (Lesson).OREGON:DryCreek(L.Miller, 1944, COlldor, vol. 46, p. 27);FossilLake?(Howard,1946, Publ.CarnegieIllstll. Washingtoll, no. 551, p.156). CALH'ORNIA: Sallta MOllica(L. 1925, COlldor, vol. 27, p.145);Sail Pedro (Howard, 1949, COlldor, vol. 51, p.23):MallixLake?(Howard,1955, U.S.geol. Surv. profess.Paper,110.264-J,p.202);"Emeryville(Howard,1929, Ulliv. Calif. Publ. Zool., vol. 32, p.312);

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 255"BuenaVistaLake(DeMay,Condor, vol. 44, p.228).NEVADA:Rattlesnake Hill?(Wetmore,1940, Smithsonian misc. Call., vol. 99, no. 4,p.13); Crypt Cave(Howard,1958, vol. 60, p.412).IUHAO:Hagerman(Wetmore,1933, Smithsonian misc. Coli., vol. 87, no. 20, p.6);TwinFalls County (Brodkorb, 1958, Wilson Bull., vol. 70, p.237).IOWA:"MillCreek(Hamon,1961, Plains Anthropologist, vol. 6, p.209).NovASCOTiA:"Bear River (HalifaxMus.).FLORIDA:Seminole Field, HogCreek near Sarasota. Itchtucknee River, Rock Spring, and Vera Beach (Wetmore, 1931, Smithsonian misc. ColI.,vol. 85, no. 2, p.13);Bradenton(Wetmore,1945, Auk, vol.62, p. 4.59);LakeMonroeandLake Washington (Brodkorbcall.);"Good's shellpit,"LemonBlull,and"Blullton (Neill,Cut,andBrodkorb, 1956, Amer. Antiquity, vol. 21, p.388):"SouthIndianField 1959, Florida Anthropologist, vol. 12, p.73);"GreenMound(Hamon,1959, Auk, vol. 76, p.533);"CastleWindy(Bullenand Sleight, 1959, Rept. Bryant FoundationAmer. Studies,no.1,p.20).2. Phalacrocorax nlivaceus(Humboldt). BltAZIL: Lapa da Escrivania andLapadaLagoadoSnmidouro(0.Winge,1887, E Mus.Lund.,vol. 1, no.2,p.5)..3.Phalacrocorax carbo(Linnaeus). NORWAY: Vardo(Lambreebt,1933,Handb.Palaeorn., p.733).DENMARK:Fannerup,Mejlgaard, Havnoe, Krabbe sholm, Erteboelle,Gudumlund,Maglemose, Klintcsoc, Soelager,"ErlangVig, "Vejleby, *Borrebjerg, Barsmark, "Noerre Broby, "Vimosc, BrogaardsMose,and"OrdrupMosc(H.Winge, 1903, Vidensk. Meddel. natllThist. Foren. Copenhagen, vol. 6, p.99). IRELAKD: NewhallCave(Lambrecht,1933).SCOTLAND: Caithne", (Lydekker, 1891, Ibis, p.388);Cnoc-Sligeach Oransay(Lambrecht,1933).ENGLANn:Grays (Milne-Edwards, 1867, Ois. Foss. France, pI.42);WestRunton(E.T. Newton, 1887, Ceol. Mag., p.147). Devils Tower? (Bate. 1938, Jour. Roy. anthrop. Inst., vol.58,p.104). ITALY' Grotta Romanelli(Lambrecht,1933). ALASKA: "Kodiak Island(Friedmann,1933, Condor, vol. 35, p.30).NOVASCOTIA:'PortJollie (HalifaxMus.),4. Plllt!,acrocoraxpenicillatus (Brandt).CALU'ORNIA:Santa Barbara?(Howard, 1931, Condur, vul. 33, p.30),Newport Bay, San Pedro, andSanta Monica(Howard,1949, Condor, vol..51,pp,21-27);"Emeryville(Howard,1929, Univ. Calif. Publ. Zool., vol. 32, p.312).5. Phalacrocorax pelagicus Pallas. ALASKA: qSt. Lawrence Island,
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256BULLETINFLORIDASTATE MUSEUM Vol. 7 1893, Arch. Antrop. Etnol., vol. 23, p.262);Grotta RomanelliandBuea deI BersagIterc(Lambrecht,1933).FamilyANHINGIDAERidgwayPlotinaeBonaparte, 1831, Saggioeliuna distribuzione rnetodica degli Animali Vertebrati,p.61(typePlatusLinnaeus, 1766, a junior synonym of AnhingaBrisson) Anhingidne Ridgway, 1887, Manual N. Amer. Birds, p.73(typeAnhingaBrisson) .GenusfProtoplotusLambrechtProtoplotusLambrecht, 1930, WeI. Meded. Dienst Mijnb. K Indies, no. 17, p. 15(typeProtoplotus beauforti1 ,ambreeht ).1.Protoplotus beaufortiLamhrechtProtoplotus beaufortiLambrecht, 1930, WeI. Meded. Dienst Mijnb. E. Indies, no. 17, p. 15, text-fig. 1-4, pI. 2-3(typefrom Sipang, skeleton impression, Mus. Mijnbouw Bureauvanden OpoporigsdienstderNeder!. Indischen Regienmg,Randoeng, Java; casts inKg!.Ung. Geo!. Anstalt,Budapest).MIDDLE?EOCENE(freshwaterfishheds),SUMATRA:Sipang.GenusAnhingaBrissonAnhingaBrisson, 1760, Ornithologia, va!. 1, p. 60; vol. 6, p. 476(type Plotus anhingaLinnaeus).2.Anhinga pannonica(Lambrecht) pannonicusLambrecht, 1916, Mitt. Jahrb. ungar. geol. Anst., vol. 24, p. 1, fig. 1,3,5,7 (type's from Tatams, carpometacarpus, cervical vertebra,Kg!. Ung. Geo!. Anstalt,Budapest).LOWERPLIOCENE(Pannonianbeds),HUNGARY:KomitatBihar: Tataros.3.Anhinga parva(DeVis)Plotus parvusDeVis, Proc. Linn. Soc.N.S.Wales, va!. 3, no. 2, p. 1286, pl. 35,fig.10(typefrom River Condamine, left humerus, QueenslandMus.?).UPPERPLEISTOCENE(DarlingDownsbeds).QUEENSLAND:northbankof RiverCondamine,3 miles from Chinchilla.4.Anhinga laticeps(DeVis)Plotus latieepsDeVis, 1906, Ann. Qneensland Mus., no. 6, p. 17, p!. 6,fig.1-2(types from lower Cooper, cranium, pelvis, Queensland Mus.?).UPPERPLEISTOCENE(Katipirisands, Malkunifauna).SOUTHAUSTRALIA:lowerCooperCreek,nearLakeEyre.

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 2575.Anhinga nana(NewtonandGadow)Plotus nanusE. T. NewtonandGadow, 1893, Trans. zool. Soc. London, vol. 131 p.288,pI.34,fig.1-5(typesfromMare aux Songes, humerus, pelvis,tarsometatarsus, CambridgeUniv.).QUATERNARY.MAURITIUS:Mare aux Songes(NewtonandGadow, 1893).QUATERNARY.MADAGASCAR: SiraM (Andrews, 1897, Ibis, p. 358). Neospecies of Anhingidae from Pleistoceneand sites:1.Anhinga anhinga ( Linnaeus).FLORIDA;Melbourne (Wehnore, 1931,Smithsonian misc. Coli., vol. 85, no. 2, p.14);Rock Spring (Woolfcndcn, 1959, Wibon Bull., vol. 71, p.185);Itchtucknee River (McCoy, 1963, Auk, vol.SO,p.000);Lake Monroe (Brodkorb call.);'LemonBluff (Neill,Gut,andBrodkorb, 1956, Amcr. Antiquity, vol. 21, p.388).FamilySULIDAE(H.cichenbach)SularinaeReichenbach, 1849, Avium systcma naturale, p.00(typeSula Bris son).-SulinaeBonaparte, 1853, C.n. Acad. Sci. Paris, vol. 37, no. 18, p. 643.GenusSulaBrissonSulaBrisson, 1760, Ornithologia, vol. 1, p. 60 (typePelecanus piscatorLinnaeus).1. Sula ronzoniMilne-EdwardsMergus ruuzuni Gervais,IS49(IS51?), Mem. Acad. Sci. Lett. Montpellier, sec.sci., vol.1,p.220(nomennudum).Sula ronzoniMilne-Edwards, 1867, Ois. Foss. France, vol. 1, sheet 34, p. 271, pI. 44,fig.9(typefrom Ronzon, pelvis, Mus. Saint PierreatLyon).LOWEROLIGOCENE(mamescalcairesdeRonzon).FRANCE:ProvoAuvergne: RonzonnearPuy-en-Velay.2.Sula arvernensisMilne-EdwardsSula arvernensis Milne-Edwards, 1867, Ois. Fuss. France, vol. 1, sheet 34,p.267, pI. 43,fig.12(typefrom Gannat, pelvis, call.Milne-Edwards).UPPEROLIGOCENE(calcairedeGannat).FRANCE:Dept.Allier'Gannat.3.Sula universitatisBrodkorbSuZauniversitatisBrodkorb, 1963(inpress),Quart. Jour. Florida Acad. Sci., vol. 26, no. 2, p. 000,fig.0(typefrom Gainesville, proximalpartof left carpometacarpus, Brodkorb call. no.8505).LOWERMIOCENE(Hawthorneformation).FLORIDA:AlachuaCounty:Gainesville.

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BULLETINFLORIDASTATEMUSEUMVol. 74. Sula pohliHowardSula pohliHoward,19,58(Aug.15),Los Angeles County Mus. Contr. in Sci.,no.25,p.4,fig.1-2(typefrom Studio City, wings, pectoral girdle, sternum,Los Angeles Mus. no. 2674; reverse Pohl Mus. no.PV68).UPPERMIOCENE(Monterey shale, Valmonte diatomitemember).CALIFORNIA:Los Angeles County: Studio City, Ventura BoulevardbetweenWhitsett AvenueandColdwater Canyon Road.5. Sula willettiL.MillerSula willettiL. Miller, 1925(Aug.),Puhl. Carnegie Instn. Washington,no.349,p.112,pI.3, 8(typefrom Lompoc, skeleton impression, Univ. Calif.Mus. Paleo. no.26542).UPPERMIOCENE(Montereyshale, Valmonte diatomitemember).CALIFORNIA:Los Angeles County: Lomita?(Howard,1958, Los Angeles County Mus. Contr. Sci., no. 25, pp.S,10);Sherman Oaks(Howard,1962, Condor, vol.64,p. 512).UPPERMIOCENE(Sisquoc formation).CALIFORNIA:SantaBarbaraCounty: Johns Manville mines, SJf miles south of Lompoc(L.Miller, 1925) .6.Sula guanoBrodkorbSula guanoBrodkorb, 1955 (Nov.30),Florida Geol. Surv. Rept. Invest., no. 14, p. 9, pI. 1,fig.2, 5; pI. 2,fig.8(typefrom Brewster, left coracoid, Brod korb no.301).LOWERPLIOCENE(Bone Valleygravel).FLORIDA:Polk County: Brewster.7.Sula phosphataBrodkorbSula phosphataBrodkorb, 1955 (Nov.30),Florida Geol. Surv. Rept.Invest., no. 14, p. 11, pI. 1, fig. 3, 6; pI. 2,fig.9(typefrom Brewster, right coracoid, Brodkorb no.302).LOWERPLIOCENE(Bone Valley gravel).FLORIDA:Polk County: Brewster.8. Sula humeralisL.MillerandBowmanSula humeraUs L. MillerandBowman, 19.58(March6),Los Angeles County Mus. Contr.inSci., no. 20, p. 9,fig.2(typefrom San Diego, distalendofright humerus, Univ. Calif. Mus. Paleo. no.45889).MIDDLEPLIOCENE(SanDiego formation).CALIFORNIA:San Diego County: San Diego.

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 2.59 GenustMierosulaWetmoreMicrosuZaWetmorc, 1938 (Jan.14),Proc. U.S.Nat. Mus., vol. 8.5, p. 2.5 (typebyoriginal designationSula adta Wetmore;subgenus).9.Microsulapygmaea(Milne-Edwards)Sula llygmaea Milne-Edwards, 1874, Bihl. Ecole hautes Etudes Paris, sect. sei.nat., vol. 11, art. 3, p. 8, pI. 12, 2(typefrom Lcognan, lefthumerus,Delfortrie coiL).MIDDLEMIOCENE(molassedeLeognan).FRANCE:Dept. Gironde: Leognan.10.Microsula avita(Wetmore)Sula avitaWctmorc,1938(Jan.14),Proc. U.S.nat. Mus., vol. 8.5, p. 22, fig. 2-3(typefrom Plumpoiot, distalpartof humerus, U.S.Nat. Mus. no. 138.54). tvlrDDLE MIOCENE(Calvertformation, zone 10).MARYLAND:Calvert County:nearPlumpoint. GenusMarusVieillotMorusVieillot, 1816, Analyse d'une nouvelle omithologie elementaire, p.63(typePelecanus Linnaeus).11.M orus loxostyla(Cope) 5t1Zaloxostyla Cope, 1870 (Dee.), Trans. Amer. philos. Soc., n.s., vol. 14, p. 236,fig..53(typefrom Calvert Co., left coracoid,Copecoll.).Sula atlanticaShufclclt, l\1l5 (Feb.),Trans. Connccticut Acad. Arts Sci., vol. 19, p. 62, pI. 1.5, fig. 123(typefromNewJersey, left coracoid, YalePeahodyMus. no.937).-Wetmore,1926, Auk, vol. 43, p. 465(typerestudied).MIDDLEMIOCENE(Calvertformation).MARYLAND:Calvert County: Chesapeake Beach(Wehnore,1926, Auk, vol. 43, p. 465).MIDDLEMIOCENE(Kirkwood formation,Ammodonbeds; sec Marsh, 1893, Am. Jour. Sci., p. 412).NEWJERSEY:apparentlyFarmingdalein Monmouth County.12.Morus vagalHl1ldus (Wetmore)Moris vagabundu", Wetmore, 1930(July 1.5), Proc. Calif. Acad. Sci., ser.4,vol. 19, p. 89, fig. 4(typcfrom Sharktooth Hill, distal eml ofrighthumerus,Univ. Calif. Mus. Paleo. no.31062).MIDDLEMIOCENE(Temblorformation).CALIFORNIA:Kern County: Sharktooth Hill, in sec.25,Township28South,Range28East, 7 miles northeast of Bakersfield (Wetmore, 1930); westbranchof Granite

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260BULLETINFLORIDASTATEMUSEUMVol. 7Creek, in sec.28,Township 27 South, Range28East,11miles north of Bakersfield (Compton, 1936, Proc. Calif. Acad. Sci., ser. 4, vol. 23,p.84).13.Morus lornpocanus(L.Millcr)Sula lompocanaL.Miller, 1925(Aug.),Publ. Carnegie Imtn. Washington, no. 349, p. 114, pI. 4,fig.7b, 9(typefrom Lompoc, skeleton impression, Univ. CaIif. Mus. Paleo. no.26544).UPPERMIOCENE(Sisquoc formation).CALIFORNIA:Santa Barhara County: Johns Manville mines, 3)f miles south of Lompoc.14.M orus peninStilaris BrodkorbMorus peninsularis Brodkorb, 1955 (Nov.30),Florida Geol. Surv. Rept.Invest., no. 14, p. 8, pI.1, fig. 1, 4; pI. 2,fig.7(typefrom Brewster, left coracoid, Brodkorb no.148).LOWERPLIOCENE(Bone Valley gravel).FLORIDA:Polk County: Brewster.15.Morus reyanus(Howard)Moris reyanaHoward, 1936 (Sept.15),Condor, vol. 38, no. 5, p. 213(typefrom Del Rey Hills, left coracoid, Los Angeles Mus. no.991).UPPERPLEISTOCENE(Palos verdessand).CALIFORNIA:Los Angeles County:DelRey Hills, 2 miles east-northeast of Playa del Rey(Howard,19,'36).Orangc County: Ncwport Bay(Howard,1949, Condor, vol. 51, pp. 21-29). Genus fPalaeosula HowardPalaeosulaHoward, 1958 (Aul(. 15),Los Angeles County Mus. ContI'.in Sci., no. 25, p. 12(typebyoriginal designationSula stocktoniMiller).16. PalaeoStila stocktoni(L.Miller)Sula stocktoniL.Miller, 1935 12),Univ. Calif. Los Angeles Pub!.bioI. Sci., vol. 1,p.75, fig. 2(typefrom Lomita, \vings, sternum, coracoid,Univ. Calif. Mus. Paleo. no.32105).UPPERMIOCENE(Monterey shale, Valmonte diatomitemember).CALIFORNIA:Los Angeles County: near Lomita;ElSereno(Howard,1958, Los Angeles County Mus. Contr. Sci.,nO.25, pp. 3,12).Genus fMiosula L. MillerMiosulaL. Miller, 192,5 (Aug.),Pub!. Carnegielnstn.Washington,no.349, p. 114(typebymonotypyMiosula mediaMiller).

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1963 BRODKORB;CATALOGUEOFFOSSILBIRDS17. Miosula mediaL.Miller261 Miosnla mediaL.Miller, 192,'5 (Aug.),Publ. Carnegie Insln. Washington, no. 349, p. 114, pI. 5(typefrom Lompoc, skeleton impression, Univ. Calif. Mus. Paleo. no.26543).UPPERMIOCENE(Sisquoc formation).CALIFORNIA:Santa Barbara County: Johns Manville mines, 3J.> miles south of Lompoc.18.Miosula recentior HowardMiasnla recentiorHoward,1949(June22),Publ.Carnegie InstIL Washington, no.,584,p. 190, pl. 2, fig. 1-2(typefrom San Diego, distalpartofright tibio tarsus, Los Angeles no.2117).MIDDLEPLIOCENE(SanDiego formation).CALIFORNIA:San Diego County: San Diego. Neospecies of Sulidae from Pleistoceneand'prchistoricsites:1.Sula daetylatraLesson.ST.CROIX;"Concordia(Wetmore,1937, Jour. Agr. Univ.PuertoRico, vol. 21, p.6).2.Sula snla (Linnaeus). RODRIGUlCZ; 'superficialdeposits(Lydekker,1891, Cat. Foss. Birds Brit. Mus., p. 46). ST.CROIX:'Concordia(Wetmore, HXJ7. Jour. Agr. Univ.PuertoRico, vol. 21, p. 6). 3.S"la leneogaster(Boddaert).BAHAMAS;"CordonHills on Crooked Island(Wetmore,1938, Auk, vol. 55, p.52).ST.THOMAS;"midden(Wctmore,1918, Proc. U. S. nat. Mus., vol. 54, p.514).ST.CROIX:'Concordia(Wetmore,1937, Jour. Agr. Univ. Puerto Rico vol. 21, p.6).4, Mvrus bassanus (Linnacus).NORWAY:Vardo(Lambrecht, 19:1,1, Handb.Palaeorn.,p. 7:33). DENMARK:Fannerllp,Erteboelle, Hesseloe, and'OrdrupMosc(H.Winge, 1903, Vidensk. Mcddel. naturbisl. Foren.Copenhagen, vol. 6, p.100).SCOTLANn; "Caithne" (Lydekker,1891, Ibis, p. :J88);Colonsay, bOransay, Orkney, and (lAndrossan (Lambrecht,1933).IRELAND:'WhitepeakBay (Lamhreeht, 1933).ENGLAND:"Whitburn(Lambrecht,1933).NOVASCOTiA:"Whynacht(HalifaxMus.).FLORIDA:GreenMound(Hamon, 19,'59, Auk, vol. 76, p.533);'CastleWindy(BullenandSleight, 1959, Rept.BryantFound.Amer. Studies, no. 1, p.20);"SummerHaven(Brodkorb, 1960, Auk, 77, p. :142). SnborderPHAETHONTESSharpePhaethontesSharpe, 1891, Review Recent Attempts to Classify Birds, p. 76 (type PhaelhonLinnaeus).FamilyPHAETHONTIDAE(Bonaparte)PhaetonidaeBonaparte, 1853, C.R.Aead. Sci, Paris, vol. 37, no. 18, p. 643 (type"Phaeton"Linnaeus).GenustPl'OphaetonAndrewsProphuetonAndrews, 1899, Proc. zoul. Soc. London, p. 776(typebyIllonotypyProphaetofl 8hrubsvlei Andrews).

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262BULLETINFLORIDASTATEMUSEUMVol. 71.Prophaeton shrubsoleiAndrewsProphaeton shrubsoleiAndrews, 1899, Proc. zool. Soc.London,p.776,text-fig. 1-2, pI. 51, fig. 1-2(typefrom Sheppey, skulJ, pelvis, femur, Brit.Mus.).LOWEREOCENE(Londonclay).ENGLAND:Kent:SheppeyIsleatmouthofThames. Neospecies ofPhaethontidaefrom sites:1.Phaethon lepturusDaudin.RonnJGUEZ: "'superficial deposits (Lambrecht,1933,lIandh.Palaeorn., p.732).BERMUDA:'CockroachIsland(Wetmore, 1902, Smithsunian misc. Call., vol. 145,nU.2, p.17).SubordertODONTOPTERYGIASpulskiOdontopterygiaSpulski, 1910 (Apr.4),Zeitsehr. deutsch. geol. Ges. Monatsber., Abh. 22, nu. 7, p. 521(Ordnung;typeOdontopteryxOwen).-OdontopterygesLambrecht,1933,Handh.Palaeorn.,p.304(sllbordo).-OdontopterygiformesHuward,1957(Feb.1),SantaBarbaraMus. nat. Hist., Bull. Dept. Ceol., ua.I,p. 21(order).FamilytODONTOPTERYGIDAELambrechtOdontopterygidoeLambrecht,1933,Handb.Palaeorn.,pp.ix,304(typeOdantapteryxOwen).GenustOdontopteryxOwenadantapteryxOwen,1873(readJune25),Quart.Jour. geol. Soc.London,vol. 29,pt.I,p. 511(typebymonotypyOdantapteryx toliapicusOwen).OdantornisOwen, 1873(readJune25),Quart. Jour. geol. Soc.London,vol. 29, pt.I,p. 521 footnote(equivalenttoOdantopteryx: "I shouldhavepreferred the term Odontornis formygenus;butit is bespoke for Marsh's suhclass.") .1.Odontopteryx toliapicaOwenOdotltopteryx toliapicusOwen, 1873(readJune25),Quart. Jour. ileal. Soc. London, vol. 29, pt.I,p.5Il,pI. 16-17(typefrom Sheppey, skul1, Brit. Mus. no.44096).LOWEllEOCENE(Londonclay).ENGLAND:Kent:SheppeyIsle.FamilytPSEunoDONTORNITHIDAELambrechtPseudodontornithidaeLambrecht,1933,Handb.Palaeorn., pp.ix,30.5(typePseudodotltornisLambrecht).GenustPseudodontomisLambrechtPseudadontornisLambrecht,1930(Jan.2,5), Ceol. hungarica, ser. pal., fasc. 7, p. I(typebymonotypyOdontopteryx Iongil'Ostris Spulski).

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDS2631.Pseudodontornis longiro;tris (Spulski) aduntapteryx longir"strisSpulski, 1910 (Apr.4),Zeitschr. deutsch. geol. Ges. Monatsher., Abh. 22, no. 7, p. 507,fig.1-7(typefrom unknown locality, skull, Albertus-Magnus Univ., Konigsberg).-Lambrecht,1930, Ceol. hungarica, ser. pal., fase. 7, p. 1, text-fig. 3,6;pI. 1-2(typerestudied).MIOCENE?BRAZILorGERMANY?GenustOsteodontornisHoward O'teocUmtornis 1957(Feb.1),Santa Barbara Mus. nat. Hist., Bull. Dept. Geol.,no. 1,p. 3(typebyoriginal designationOsteodontornis orriHoward).2.Osteodontornis orriHowardOsteoduntuTnis urriHoward,19.57(Feb. 1), Santa Barbara Mus. nat. IIist.,Bull. Dept. Ceo!', no. 1,p.3, fig. 2-8(type fromTepusquet Canyon,incom plete skeleton impression, Santa Barbara no.309).UPPERMIOCENE(Montereyshale, Valmonte diatomitemember).CALIFORNIA:SantaBarbaraCounty:westside ofTepusquetCreek, f1agstunequarryuf C. Antulini&Sons(Howard,1957). Los Angeles County: Sherman Oaks(HowardandWhite, 1962, Los Angeles Co. Mus.,Contr.Sci.,nu. 52, p.3,fig.2-5). FamilytPELAGORNITHIDAE(Fiirhringer) PelagornithinaeFiirbringer, 1888, Untersuch. Morph. Syst. Vi;gel, vol. 2, p. 1565footnote (subfamily; type Pelagurnis Lartet).-Pelagornithidae\Vetmore,1930, Proc. U.S.nat. Mus., vol. 76, p. 2.GenustPelagornisLartetPelagornisLartet, 1857(readApr.6),C.R.Acad. Sci. Paris, vol. 4,1, no. 14, p. 740(typebymonotypyPelagornis miocaenusLartet).1.Pelagornis miocaenllsLartetPelagornis Lartet, 1857 (read Apr.6),C.R.Acad. Sci. Paris, vol.44, no. 14, p. 740 [separate includes a plate, fide Milne-Edwards](typefrom Armagnac, left humerus, ParisrvIlIs.).-\IIiJne-Edwards, 1867, Ois. Foss.France, vol.I,p. 273, pI. 45(typerestudied).PelagornisdelfortriiLamhrecht, 11133, Handb.Palaeorn., p. 282 (nomennudum).MIDDLE YhOCENE (molasse coqnillieremarinedc]'Armagnac). FRAKCE: Dept. Gers; Armagnae(Lartet, 1857). MIDDLEMIOCENE(molassede Leognan). FRANCE:Dept. Gironde: Leognan (Milne-Edwards, 1B74, Bib!. Ecole llautes Etudes,Paris, sec. sci. nat., vo!.11,art.3,p.1).

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264BULLETINFLORIDASTATEMUSEUMSubordertCLADOHNITHESWetmoreVol. 7 Clmlornit"es Wetmore,1960(June23),Smithsonian misc. Call., vol. 139, no. 11, pp. 4, 25 (suborder;type Gladomis Ameghino).FamilytCLADORNITHIDAE(Ameghino)CladornidaeAmeghino, 1895 [separate apparently published 1894], Bol. Inst. geog. argentino, vol. 15, cahiers 11-12, p. 584 [1>5 of separate](typeCla damis Ameghino ).Gladornit"idaeWetmore, 1930, Proc.U.S.nat. Mus" vol. 76, no. 2821, p. 2.GenustCladornisAmeghinoGladornisAmeghino, 1895[l894?],Bol. Inst. i(eog. argentino, vol. 15, cahiers 11-12, p.58.5[86 of separate](typeby monotypy Gladornis pac"ypusAmeghino).1.Cladornis pachypusAmeghinoGladomis pachypusAmcghino, 11>95 [1894?], Bol.Tnst.geog. ari(entino, vol. 15, cahiers 11-12, p. 585 [86 of separate], Iii(. 35(typefromPyrotheriumbeds, distal part of right tarsometatarstls, nmv in Brit. LOWEROLIGOCENE(Deseadoformation).ARGENTINA:Ter. Santa Cruz: Rio Deseado. FamilytCYPHORNITIIIDAE ''Vetmore Gyplwrnit"idaeWetmore, 1928(March15),Canad.Dept.Mines, Ceol. Surv. Bull.,no.49, p. 4(typeGyphornisCope).GenustCyphornisCope GyphornL, magnusCope, 1894(May:31), Jour. Acad. nat. Sci. Philadelphia, ser. 2, vol. 9, p. 449,(typebymonotypy Gyphor"is mag"usCope).1.Cyphornis magnusCopeGyphornis magnusCope, 1894(May .31), Jour. Acad. nat. Sci. Philadelphia, ser. 2, vol. 9, p. 451, pl. 20, Iii(. 11-16(typefromCarmanahPoint, proximalpartof left metatarsus, Can. GeoI. Surv. no. 6.32.3).-Wetmorc, 1928,Canad.Dept.Mines, Gcol. Surv. Bull., no. 49, p. 1, lig. 1(typerestudied).LOWERMIOCENE(CarmanahPointbeds),BRITISHCOLUMBIA:Vancouver Island:CarmanahPoint. Genus tPalaeochenoidesShufeldt Palaeoc"enoides Shufeldt, 1916(Aug.),Geol. Mag., n.s.,decade6, vol. 3, p. .347 (typebymonotypy Pakwoc"enoides mioceanusShufeldt).

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1963 BRODKORB: CATALOGUEOFFOSSILBInDS2.Palaeochenoides mioceanusShufeldt265Palaeoehenoides mioeeanusShufeldt, 1916(Aug.),Geol. Mag., n.s.,decade6, vol. 3, p. 347, pI. 15(typefrom Stano River, distalpartof left femur, U.S.nat.Mus.).-Wetmore,1917, Jour. Geol., vol. 25, no. 6, p. 555,fig.1(typerestudied) .LOWERMIOCENE(Hawthorneformation).SOUTHCAROLINA:Charleston County: near source of Stono River. SuborderPELECANISharpePeleeaniSharpe, 1891, Review Recent Attempts to Classify Birds, p.76(typePelecanusLinnaeus),FamilyPELECANIDAEVigorsPelecanidaeVigors, 1825, Trans. Linn. Soc. London, vol. 14, pp. 498, .'504 (typePelecanusI ,innaeus ).GenusPelecanusLinnaeusPeleeanus Linnaeus, 1758, Syst. Nat., ed. 10, p. 132(typePelecanus unoero talusLinnaeus).1.Pelecanl1s gracilisMilne-EdwardsPelecanus gracilisMilne-Edwards, 1863, C. R. Acad. Sci. Paris, vol. 56, p. 1222(nomen nudum).-11ilnc-Edwards, 1867, Dis. Foss. France, vol. 1, sheet 32,.. p. 350, pI. 38-39 (types fromLabeur,fureulumandupperpartof tarsometatarsus, colI. Poirrier; also from Langy, upper endofhumerus, completefemur, scapula, call. AbbotVandenhecke).LOWERMIOCENE(Aquitanian).FRANCE:Dept. Allier:LabeurnearVanmasandLangy (Milne-Edwards, 1867); Saint-Gerand-le-Puy(Lambrecht,1933, Handb. Palaeorn., p. 277). Dept. Puy-de-Domc: Montaigut (Lambrecht, 1933).2.Pelecanus intermediusFraasPelecanus intermedius O.Fraas, 1870, Jahresh.VeT.Naturk. Wiirttemberg,.vol. 26, p. 281, pI. 13,fig.3-4(typefrom Hahnenberg, skullandmandible,StuttgartMus.).UPPERMIOCENE(0bere Siisswassermolasse).GERMANY:Wiirttem berg:Hahncnbcrg(Fraas, 1870); Steinhaim (Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p.39).3.Pelecanl1s traasiLydekkerPelecanus fraasi Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p.44,fig.lOA(typefrom Klcin-Sorhcim, cranium, Brit. Mus. no.47862).

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266BULLETINFLORIDASTATEMUSEUMVol. 7UPPERMIOCENE( obere Siisswassermolasse).BAVARIA:Klein Sorheim. Wiirtternherg: Lierheim IlearHahnenbcrg(Lydekker,1891).4.P elecanus cautleyiDaviesPelicanus cautleyiDavie" 1880, Gco1. decade2, vol. 7, p. 26(typefrom Siwalik Hills; dbtalend ofleft ulna,Brit. Mus. no. 39740).-Pelecanuscautleyi Lyrlekker, 1884, Pal. indica, scr. 10, vol.3,pI. 4, p. 137,pI.14, fig. 11(typerestudied).LOWERPLIOCENE(Siwalik series).INDIA:UnitedProvinces: Siwalik Hills.5.Pelecanus sivalensis DaviesPelicanus!?) siva/ensisDavies, 1880, Ceol. Mag.,decade2, vol.1,p.26(typefrom Siwalik Hills, distalendofrightulna, Brit. Mus. no.39745).-Pelecanus sivaiensisLydekker, 1890, Ree. geo!. Surv.India,vol.2:3,p.235, Ri(. 2(typerestudied).LOWERPLIOCENE(Siwalik series).INDIA:United Provinces: Siwalik Hills.6.Pelecanus odessanusLambrechtPelicanus odessanU8 tossilisWildhall1, 1886, Schrift. Nellruss. Ges. Naturf. Odessa,vol. 10,Bdlage,p.4,pI.5,fig.[,4(nun-binomial).-Pelecanus odessanusLambrecht, 1933,Handb. l'alaeorn., p. 278 (types from Slobodka, coracoid, tarsomctatarslls) .LOWER PLIOCENE (Meotian). UKRAIN>:: SlobodkanearOdessa.7.Pelecallus haLieus WetmorePelecanus halieus\Vetmore, 193.3(Del'. 27),Smithsonian misc. Cull., vol. 81,no.20,p.3,fig.1-2 (type fromscc. 16, proximalpartofrightradius,U.S. Nat. Mus.nO.12233).LOWERPLETSTOCENE(GlennsFerryformation,HagermanLakeheds).TDAHO:Gooding County:NW of section16,Township7South, Range13East,2miles west ofHagerman.8. Pelecml1ls gralldicepsDeNis PelecllnusgrandicepsDeVis, 1906, Ann. Queensland no. 6.p.18,pI.5,fig. I-a (typefrom lower COOpE'I, left quadrate, left coracoid. di.-dal partoflefttarsometatarsus).UPPERPLEISTOCENE(Katipiri sands, Malkunifauna).SOUTHAUSTRALIA:lowerCooperCreeknearLakeEyre.

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1963 BRODKORB:CATALOGUEOFFOSSILBlRDS9.Pelecanus proavusDeVis267Pelecanus proavusDeVis, 1892, Proe. Linn. Soc. N. S. Wales, ser. 2, vol.6,p. 444, pI. 24,fig.6(typefrom Queensland, proximalpartofcarpometa carpus ).UPPERPLEISTOCENE(DarlingDownsbeds).QUEENSLAND(DeVis,1892).UPPERPLEISTOCENE(Katipirisands, Malkunifauna).SOUTHAUSTRALIA:lower Cooper Creek (DeVis, 1906, Ann.QueenslandMus., no. 6, p.17,pI.5,fig.4-5).GenustLiptornisAmeghinoLiptornisAmeghino, 189.5, Bol. inst. Geog. argentino, vol. 1.5, cahiers 11-12, p. 99(typebymonotypyLiptornis hesternw; Ameghino ).10.Liptornis hesternusAmeghino Liptorni,f!t hesternus Ameghino, 1895, Bol. Inst. geog. argentino, vol. 15, cahicrs11-12, p. 99(typelower cervical vertebra, Brit. Mus.). MIDDLEMIOCENE(SantaCruzformation).ARGENTINA:Tcr.SantaCruz: Cueva. Neospecies of Pelecanidae from Pleistoceneand sites:1.Pelecanus onocrotalu8Linnaeus.ENGLAND:Norfolk (Lydekker, 1891, Ibis,p.387).SWITZERLAND: Sec(Lambrecht,1933,Handb.Palaeom.,p.732).2. Pelecanus crispusBruch.DENMARK:Havnoe(H. vVingc, 1903, Vidensk.Meddel. naturhist. Foren. Copenhagen, vol. 6, p.100).ENGLANV:FeltwellFen(Newton, Proc. zool. Soc. London, p.702);BurntFennearLittleport(Harmer,1898, Geol. Mag., p.418).AZERBAIJAN:BingadancarBaku(Pe/eeanus erispus palaeocrispusSeTebrovsky, 1941, Doklady Akad.NaukS.S.S.R., vol. 33, p.472).3.Peleeanu. erythorhyuclw8 Gmelin.OREGON:Fossil Lake (Howard,1946,Puhl. Carnegie Instn. \Vashington, no.,5,5],p. 153;Shufeldfsearlier records erroneous);DryCreek?(L.Miller, 1944, Condor, vol. 46, p.26).CALlFORNIA:Manix (Compton, 1934, Condor, vol. 36, p.167);"Emeryville(Howard,1929, Univ. Calif. Publ. Zool., vol. 32, p.312);"BuenaVistaLake(DeMay,1942, Condor, vol. 44, p.228).NEVADA:Rattlesnake Hill(Wetmore,1940, Smithsonian misc. ColI., vol. 99, no. 4,p.10).SOUTHDAKOTA: I)Corson County(L.Miller, 1961, Bull.S.Calif. Aead. Sci., vol. 60,pt.3, p. 12.5). KANSAS:ShOTtSCreek (Stettenheim, 19.58, Wilson Bull., vol. 70, p.197).OKLAHOMA:Beaver County (Mengel, 19.52, Auk, vol. 69, p.81).IOWA:"Mill Creek(Hamon,1961, Plains Anthropologist, vol. 6, p.209).ILLINOIS:'Snyderand'Cahokia(Parmalee, 19.58, Auk, vol. 7.5, p.170).4. Pelecanus occidentalisLinnaeus.CALIFORNIA:Carpinteria(DeMay,1941,Publ. Carnegie Instn. Washington, no .530, p.64);'Emeryville(Howard,1929,

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268BULLETINFLORIDASTATE MUSEUMVol. 7 Univ. Calif. Publ. Zool., vol. 32, p. 312); "BuenaVistaLake(DeMay,Condor, vol.44,p.228).rLLJNOIS:'Modoc (Parlllcllee, 1958, Auk, vol. 75, p. 170 [needsconfinnation]).FLORIDA:"CastleWindy(BullenandSleight, 1959, Rept. Bryant Found. Amer. Studies, no. 1. p.20).PUERTORICO: Canas(Wetmore,1938, Auk, vol. 55, p.53).ST.CROIX:'Concordia(Wetmore, 1917, Jour. Agr. Univ.Puerto Rico, vol. 21, p.6).SuborderFREGAT AE(Sharpe)FregatiSharpe, 1891, Review RecentAttemptsto Classify Birds, p.77(suborder; type Fregata and \N. D.Miller, 1926, Auk,\'01.43, uo. 3, p.341).Family FREGAcTIDAcE GarrodFregatidaeCarrod,1874(rcadFeb.3),Pree. zool. Soc. London, p. 117(typeFregata Lacepedc). Neospecies ofFregatidaefromprehistoricsites:1.Fregata magnificens Mathews.ST.THOMAS:"midden(Wetmore,1918, Proc. U. S.nat.Mus., vol. 54, p. 515). AI
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1963 BRODKORB:CATALOGUEOFFOSSILBIRDSOrderAHDEIFORMES(Wagler)269Ardeae\Vagler, 1830, Naturliches SystemderAmphibienmit vorangchcnderClassificationderSaugethiereuudVogel, p. 000 (ordo; typeArdeaLinnaeus).Wagler, 1831, Isis vonOken,Heft4, p.530.-Ardeiforme8Gadow, 1892, Proc. zool. Soc. London, p. 240(order).TantaliWagler, 1830, Nat. Syst. Amphib. Skiugcth. Vogel, p. 000(ordo;typeTantalusLinnaells,1758,asynonymofMycteriaLinnaeus,1758).-Wagler,18.31, Isis von Okcn,Heft4, p. 530.Herodione8Bonaparte, 1853, C. It Acad. Sci. Paris, vol. 37, no. 18, p. 642 (ordo; typeHerodiasBoic, 1822, a junior synonym of Egretta Forster,1817).-Bonaparte, 1854, Ann. Sci. nat.(Paris),p.,37.-HerodiiCope, 1889(Oct.),Amer. Natural., vol. 23, no. 274, p. 871(suborder).Ciconiae Bonaparte, 1854, Ann. Sci. nat.(Paris).p.37 ("tribus," i.e. suborderofIIerodiones; type CiconiaLinnaeus).-CiconiiformesGarrod, 1874, Froc.zool. Soc. London, pp. 117, 122(order).SuborderPHOENICOPTERIFlirbringerPhoenieo"teriFiirhringcr, 1888, Untersuch. Morph. Syst. Vogel, vol. 2, p. 1565 (gens; type Phoenicopterus Linnaeus).-PhoenicopteriformesSharpe, 1891, Review Recent Attempts to Classify Birds, p. 76(order).FamilyfToROTIGIDAEBrodkorbTorotigidaeBrodkorb, 1963 (inpress), Pmc. XIII intcrnat. ornith. Congr. Ithaca,p. 000(type1'orntixBrodkorh).GenusfGallomisLambrechtGallomisLambrecht,1931, Bull. Mus. lIist. nat. Belgique, vol. 7, no. 30, p. 1(typebymonotypyGallornis straeleniLambrecht). Position tentative.1.Gallomis straeleniLambrechtGallornis strae/eni Lambrecht, 1931, Bull. Mus. Hist. nat. Belgique, vol. 7, no. 30, p. 1,fig.1-3(typefrom Auxerre, proximalendof femur, BrusselsMus.).LOWERCRETACEOUS,NEOCOMIAN. FHANCE: Dept.Yonne: Auxerre.GenusfParascaniomisLambrechtPara8caniorni8Lambrecht, 1933, Hanelb. Palacom., p. 335 (typebymonotypyParascaniornis ,nensiOi Lambrecht). Position tentative.2.Parascaniornis stensioiLambrechtPara8eaniorni8 8ten.-noi Lambrecht,1933,Handb.Palaeom., p..335,fig.116(typefrom Iva, vertebra, Mineralogical-Geological Museum,Copenhagen).UPPEHCRETACEOUS,CAMPANIAN(Shellfragmentlimestone).SWEDEN:I vo.

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270BULLETINFLORIDASTATE MUSEU/vIVol. 7GenustTorotixBrodkorbTorotixBrodkorh, 1963(inpress),Proc.XIIIinternat.omith.Con gr.Ithaca,p.000(typehyoriginal desii(nationTo-rolixclemen.i Brodkarh).3.Torotix clemensiBrodkorbl'orotix clemensi BrodkOl'b,1963(in press), Proc. XIII internat. ornith. CangI. Ithaca,p.ODD,fig.4-.5(typeelistal ofrighthumerus,Univ. Calif. Mus. Palco. no.53(58).UPPERCRETACEOUS,MAESTRICHTIAN(Lanceformation).VVYOMING:NiobraraCounty:LanceCreek.FamilytSCANIORNITHIDAELambrechtScaniornithidaeT ,ambre<.:ht, 1933, -Hanelb. Palaeorn.,p.334(typeSccmiurnisDames).GenustScaniornisDamesScalliorni.Dames,1890(readJan.8),Bihangsvcn.kaVet.-Akad.Handl.,vol.16,pt.4,no.I,p.4(typebymanotypyScalliornis lundgreniDames).1.Scaniornis lundgreniDamesScaniornis ZundgreniDames,1890(readJan.8),BihangsvenskaVet.-Abd.Handl.,val.16, pt. 4, no. 1, p. 4, pI.(typesframAnnetorpquarry,righthumerus, coracoid, scapula, Univ.Lund).LOWERPALEOCENE(SaItholmskalk).SWEDEN:AnnetorpquarrynearLimhamn.FamilytTELMABATIDAEHowardTelmahatidaeHoward,19,5,5(March11),Amer. Mus. Navit., no.1710,p.23(typeT elmabatesHoward).Genus tTelmahates Howard Telmabates Howard, 19.55 (MarchII),Amer. :\1us. Novit., no. 1710, p. 3(typehyoriginal designationT elmabates antiquusHoward).1.Telmabates antiquusHowardTelmabates antiquasIIoward, 195.5 (March11),Amer. :\Ius. Navit.,nO.1710,p.3, fig.1-8(typefromCanadonHondo,pastcranialskeleton, Am. Mus.Nat. Hilit. na.3170).LOWEREOCENE(Casamayorformation).ARGENTINA:Ter.Chubut: Caiiadbn HondonearPasoNiemann,southof Rlo ChicodelChubut.FamilytAGNOPTERIDAELambrechtAgnopteridaeLambrecht,1933,Handb.Palaeorn., p. 33:1 (type Agnoptcrus:\1ilne-Edwards) .

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1963 BRODKORB:CATALOGUE OF FOSSIL BIRDS 271GenustAgnoptcrusMilne-EdwardsPtenorn;.Seeley, 1866, Ann, Mag. nat. IIist., ser. 3, vol. 18, p. 109(inadequatedescription andno specific name).AgnoptertlsMilne-Edwards, 1868, Ois. Foss. France, vol. 2, pI. 89, fig. 10-15(typebymonotypy Agnoptews laurillardiMilne-Edwards).-Milne-Edwards,1870, op. cit., vol.2,sheet 11, p. 83.1.Agnortcrus hantonicnsisLydekkerAgnopterus(?) hantoniensi. Lydekker, 1891 (Apr.25),Cat. Fuss. Birds Brit. Mus., p. 96,fig.23(typefrom Hordwcll,rightcoracoid, Brit. Mus. no.30325). UPPEll EOCENE(Hordwellbeds).ENGLAND:Hampshire:Hordwell(Lydekker, 1R91).hIe of \Night: Hempstead?(Seeley, 1866, Ann. Mag. nat. Hist., p. 109).2.Agnortcrus laurillardiMilne-Edwards AgllOl,tews /;lUrillardi Milne-Edwards, 1868, Ois. Foss.France,vol. 2, pI. 89,fig.10-15; 1870, vol. 2, sheet 11, p. 83(typefrom environsdeParis, distalpart of tihiotarsus, ParisMus,).UPPEREOCENE(gypsedeMontmartre). FRAI'CE: Dept.Seine: Montmartre.3.Agnoptcrus turgaicnsisTugarinoY Agnoptcrus turgaiensisTugarinov, 1940, Doklady Akad.NaukS.S.S.R., vul. 26, no. 3, p. 308,fig.2(typefrom Lake Chelkar-Teniz).-AgrlOpterus turgaensis Belyaeva,IH62, Cat. Tertiary Fossil Sites of Land 1famrnals in U.S.S.R.,p.8).UPPEROLIGOCENE(Indricotheriumbeds).KAZAKSTAN:LakeChel kar-Tcniz.FamilyPHOENICOPTERlDAEBonaparte Phoo#cupteridae Bonaparte. 1831, Saggio di una distribuzione metodica ueg]iAnimali Vertebrati,p.59(type Phoenicoptertls Linnaclis).GenustElornisAymard Elotnis Aymard, 1856, Congr. sci. France, vol. 1, p.234(typeElornis littoralisAymard, designatedbyLydckker, L891, Cat. Fuss. Birds Brit. Mus., p.80).HelomisLydekker, 1891, Ibis, ser. 6, vol. 3, p. 3fl6 (emendation).1.Elornis anglicusLydekkerElomis(?)allglicusLydekker, 1891 (Apr. 2,5), Cat. Foss. Birds Brit.Mus"p. 80,fig.22(typefrom Hurdwell, left humerus, Brit. Mus. no.36792).UPPEREOCENE(Hordwellbeds).ENGLAND:Hampshire:Hordwell.

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272 BULLETINFLORIDASTATEMUSEUMVol. 72.Elarnis littamlis AymardElornis littoralisAymard, 1856, Congr. sci. France, vol. 1, pp. 234, 267 (lectotype from Ronzon, humerus, suggested by Lydekker, 1891, Cat., p.80).Elomis antiquusAymard, 1856, Congr. sci. France, vol. 1, p. 234(typesfrom Ron7,onl proximal partof tarsometatarsliS, humerus).LOWER OLIGOCENE (mamescalcairesdeRonzon).FRANCE:Dept.HautLoire: RonzonnearPuy-en-Velay.3.Elarnis grandis AymardElornis grandis Aymard, 1856, Congr. sci. France, vol.1,pp. 234, 267(typefrom Ronzon, proximal partofhumerus).LOWEROLIGOCENE(marnes calcairesdeRonzon).FRANCE:Dept.HautLoire: RonzonnearPuy-cn-Velay. Genus fTiliarnis Ameghino Tiliornis Ameghino,1899(July),Sinopsis geologico-paleonto16gica, Suplemento,p. 9 (type bymonotypyTiliomis senexAmcghino).4.Tiliornis senex AmeghinoTiliornis I)'eTtex Ameghino, lR99 (July),Sinopsis geologico-paleontulogic-a, Suplemcnto,p. 9 (typefrom "Guaranitico de Patagonia," coracoid).LOWEROLiCOCENE(Deseadoformation).ARGENTINA:Patagonia. Genus Phaenicaptertts LinnaeusPhoenicopterus Linnaeus, 1758, Syst. :Nat., cd. 10, vol.1,p.139(typePhoeni copteru.';ruber Linnaeus).5. PlwenicalJteTlls croizeti. GervaisPhoenicopterus emizetiGervais, 1849, Mem. Acad. Sci. Lett. Montpellier, sec. sci., vol. 1, p.220(nomennudum;basedon"FlamantsemblableauPh. P.Gerv., Ois. foss.,p.21").-Gervais, 18,,2, Zool.etPal. Fran<;aises,ed.1,p. 2:33, pI.2, fig. 4-5(typestarsomctatarsus fromGergovieandskullfromClermont-Fl'rral1d).LOWERMIOCENE(Aquitanian).FRANCE:Dept. Puy-en-Dome: Cler mont-Ferrand and Gergovie (Gervais, 1852);ChaptuzatandCournon (Milne-Edwards, 1869, Ois. Foss. France, vol. 2, p.572);PerignatandSanveta:t (Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p.78).Dept. Allier: Chavroches, Gannat, ABets, andLangy(Milne-Edwards);Saint-Gerand-le-Puy(Lydekker).Dept. Somme: Crechy(Lamhrecht,1933,Handb.Palaeorn., p. 344).LOWER MIOC:ENE (H ydrobiensehichten).GERMANY:Prov. Rhein hessen: BudenheimnearMainz(Lambrccht).

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS6.Phoenicopterus floridanusBrodkorb273Phoenicopterus fioridaTlus Brodkorb,1953(June9),Nat. Hist. Misc., no. 124,p.1,fig.1-2(typefrom Brewster, distalpartof right tibiotarsus, Brodkorh no.147).LOWERPLIOCENE(Bone Valley gravel).FLORIDA:Polk County: Brewster.7.Phoenicopterus stockiL. MillerPhoenicopterus stockiL. Miller, 1944(June),Wilson Bull., vol. 56, no. 2, p.77, fig. 1-2(typefrom Rinc-on, distalendoftibiotarsus, Los Angeles Co. Mus.no. C.LT. 324.'1). MIDDLEPLIOCENE(Chihuahuaformation).MEXICO:Chihuahua:Rincondela Concha,nearYepomera, valley of Rio Papigochic.8.Phoenicopterus copeiShufeldtPhoenicopterus copeiShufeldt, 1891(Sept.),Amer. Natural., vol. 2S, no. 297, p.820.-Shufeldt,1892, Jour. Acad. nat. Sci. Philadelphia, vol. 9, p. 410, pI. 15,fig.11, 13; pI. 17,fig.28-29, .38 (types from Fossil Lake, distalendoflefttarsometatarslISandwingphalanx, Am. Mus. no.3485).MIDDLEPLEISTOCENE(Fossil Lake formation).OREGON:Lake County: Fossil Lake (Shufeldt, 1891).UPPERPLEISTOCENE(Manix lakebeds).CALIFORNIA:San Ber nardino County: Manix?(Howard,1955, U.S.geol. Surv. profess. Paper, no. 264-J, p. 202).9.Phoenicopterus rninutusHowardPhoenicopterus minutusHoward, 19.'1S (June8),U. S. geol. Surv. profess. Paper, No. 264-J, p. 202, pI. 50,fig.1-7(typefrom Manix, right tihiotarsus and associated proximal partoftarsometatarslls,LosAngeles Mus. no.244S).U l'PER PLEISTOCENE(Manix lakebeds).CALIFORNIA:San Ber nardino County: Manix.Neospeeies of Phoenicopteridae from Pleistoceneand prehistoric sites:1.Phoenicopterus ruberLinnacus.PUERTORICO: Barrio Canas(Wetmore, 1938, vol. 55, p.53).ST.CRorX:'Concordia(Wetmore,1937, Jour. Agr. Univ. Puerto Rico, vol. 21, p.7).ANTfGUA:'MillReefmidden(Univ.F1orida).2.Phoenicopterus chilensisMolina.ARGENTINA: Lujan (Ameghino, 1891, Rev.argentina Hist. nat., vol. 1, p.44S).

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274BULLETINFLORIDASTATEMUSEUMVol. 7FamilytPALAELODIDAE(Stejneger)PawelodontidaeStejneger, 1885, Stand. nat. Hist., vol. 4, p. 154(typePalaelo dus Milne-Edwards). PalaelodickeFtirbringer, 1888, Untersuch. Morph. Syst. Vogel, vol. 2, p. 1565.Paloelodidae Howard, 1955, Amer. Mus. Novit., no. 1710, p. 22.Genlls tPaklelodus Milne-EdwardsPalaelodusMilne-Edwards, 1863(readJune29),C.R. Acad. Sci. Paris, vol. 56,p.1220(type Palaelodus ambiguus Milne-Edwards, designatedby 1869, Ois. Foss. France,voL2, p.59). PaluelodusMilne-Edwards, 1868, Ois. Foss. France. vol. 1, pI. 82-89: 1869, vol. 2, sheet 8, p..58 (typugraphieal error furPalaelodus?) .-0.Fraas, 1870, Jahresh. Ver. Naturk. Wiirttemberg, vol. 26, p. 285(emendation).1.Palaelodus goliathMilne-EdwardsPaloelodus goliathMilne-Edwards, 1868, Ois. Foss. France, vol. 1, pI. 87: pI. 88,fig.1-3;J870, vol. 2, p. 79(lectotype from Langy, tarsometatarsus, selectedbyLydekker, 1891, Cat. Foss. Birds Brit. Mus., p.95).LOWER MIOCENE (Aquitanian).FRANCE:Dept. Allier: Langy (Milne-Edwards, 1871, vol.2,p. 572); Saint-Gerand-Ie-Puy(Lambrecht,1933.,Handb.Palaeorn., p, 341). Dept. Somme: Crcchy (Lambrecht, 1933). LOWlill MIOCENE(Hydrobienkalk). GEHMANY: Hessen: Budenheim, Kastel Bruch, River HesslerbetweenWiesbadenandMaim:, and Neuer Bruch(Lambrecht,1933).2.Palaelo,zus crassipesMilne-EdwardsPalaelodus crassipesMilne-Edwards, 1863(readJune29),C. R. Acad. Sci. Paris, vol. 56, p. 1221(almosta nomennudum;type from dept.Allier, elementnot specified).-Palueludus cms,'ipes Milne-Edwards, 1868, Ois. Foss. France, vol. 1, pI. 88,fig.4-11; pI. 89,fig.1-5.-'vlilne-Edwards,1870, Ojs. Foss. France, vol. 2, sheet 10,p.77.LOWERMJOCENE(Aquitanian).FRANCE:Dept. Allier:LangyandGannat(Milne-Edwards, 1871, vol.2,p, 572); Saint-Gerand-Ie-Puy [not Ciernat?](Lamhrecht,1933, Handb. Palaeorn., pp. 341, 884). Dept. Puy-de-Dome: MontaiglltIeBlin(Lambrecht,p. 341).3.Palaelodus ambigllus Milne-EdwardsPalaelodus ambiguusMilne-Edwards, 1863(readJune29), C. R Acad. Sci. Paris, vol. 56, p. 1221 (deser.type fromdept. Allier}.-Palneludus ambiguus Milne-Edwards, 1868, Ois. Foss. France, vol. 1, pI. 82-81, pI. 85,fig.1-11; 1869, vol. 2, sheet 8, p.60(typesredescribed).

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1963 BRODKORB, CATALOGUE OFFOSSILBIRDS 275LOWERMIOCENE(Aquitanian).FRANCE:Dept.Allier: Gannat, Billy, Langy, Vaumas, and Saint-Gerand-le-PllY?(Milne-Edwards).Dept. Puy-de-Dome: CournonandChaptuzat(Milne-Edwards);Pont-du-CMteauandPcrignat (Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p. 83). Dept. Somme: Crechy (Lambrecht, 1933,Handb.Palaeorn., p. 341).LOWERM,OCENE(Hydrobienkalk).GERMANY:Hesse: \Veisenau (Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p.83);River Hessler between WiesbadcnandMainz, "Dickkopf" near rvlonsheim, Budcn heim,andKasteler Bruch (Lambrecht, pp. 340, 670).4.Palaelodlls minutusMilne-EdwardsPaloe/ad"s min"Ws Milne-Edwards, 1868, Ois. Foss. France, vol. 1, pI. 86,fig.17-20, 1870, vol. 2, sheet 10, p. 75 (lectotype from Allier, tarsomctatarsus, cull. Milne-Edwards, selectedby Lydekker, 1891, Cat. Fuss. Birds Brit. Mus., p.92).LOWERM,OCENE(Aquitanian).FRANCE:Dept. Allier:Langy(Milne-Edwards, 1871, Ois. Foss. France, vol.2,p. 572); Saint Gerand-Ie-Puy (Paris, 1912, Rev. franc;aise Ornith., vol. 4, p. 291); Chavroches (Lambrecht, 1933, Handb. Palaeom., p. 342). Dept. Somme: Crechy (Lambrecht, p. 667).LOWER MIOC.ENE (Hydrobienkalk).GERMANY:Hesse: Kastel BruchandBudenheim(Lambrecht).5.Palaelodus graci/ipesMilne-EdwardsPalaelodtl8 gracilipes Milne-Edwards, 1863 (readJune29),C.R.Acad. Sci. Paris, vol. 56, p. 1221 (almost anomennudum;typefrom Allier).-Paloelo dus gracilipes Milne-Edwards, 1868, Ois. Foss.France,vol.1,pI. 85,fig.12-16: pI. 86, fig.1-16; 1870, vol. 2, sheet lO, p. 73(typesredescribed).LOWERMIOCENE(Aquitanian).FnANcE:Dept. Allier: Langy, Gannat, Vaumas,andSaint-Gerand-Ie-Puy (Milne-Edwards, 1871, vol.2,p.572).6.Palaelodus steinheirnensisFraasPaluelodus steinheime1lSis O. Fraas, 1870, Jahresh. Vel'. Vaterl. Naturk. \Viirttem herg, vol. 26, p. 285, pI. 7,fig.13(typefrom Steinheim, distalendof left tibiotarsus,StuttgartMus.).UPPER MIOCEKE (obereSiisswassermolasse).GERMANY:Wiirttem herg: Steinheim(Fraas).Records ofP.goliathfrom Goldberg(Lambrecht, p. 670)andP.lImbiguusfrom Goldberg, Spitzberg, Stein heim,andHahnenberg (Lambrecht, pp. 339, 678)maybereferable

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276 BULLETINFLORIDASTATEMUSEUMVol. 7to this large species, whereastherecords ofP.gracilipesfrom Stein heim(Fraas,p. 286) andP. minutus from Goldberg(Lambrecht,pp. 342, 670) suggestthepresence ofanundescribed small species intheUpperMiocene. GenustMegapaloelodusA.H.MillerMegapaluelodusA.H. Miller, 1944(June 22), Univ. Calif. Publ. geol. Sci., vol. 27, no. 4, p. 86(typebymonotypyMegapnloeludus connectensA.H. Miller ),MegapalaeludusWetmore, 1951,Proe.X.internat. ornith. Congr.,pp.58, 66(emendation).7.Megapaloelodus connectensA.II. MillerMegupaloelodus connectensA. H. Miller, 1944(June22),Univ. Calif. Puhl. geol. Sci., vol. 27, no. 4, p. 86,fig.1-2(typefromFlintHill, distalendofright Univ. Calif. Mus. Paleo. no.37367). LOWEll MIOCENE(Rosebud formation).SOUTHDAKOTA:Bennett County:FlintHill,9 milesWSWof Martin (A.H.Miller, 1944).UPPERMIOCENE(Barstow formation).CALIFORNIA:San Bernardino County: Barstow (L. Miller, 1950, Condor, vol. 52, p.69;1952, Condor, vol. 54, p. 296); elementsnotcomparable totypeandmay represent another species.8.Megapaloelodus opsigonusBrodkorbMegapaloelodus opsigonusBrodkorh, 1961 (Nov.7),Quart. Jour. Florida Aead. Sci., vol. 24, no. 3, p. 173,fig.2(typefromJuntura,proximalendof left tarsometatarsus, Univ. Ore. Mus. Nat. Hist., no.F-5459).LOWERPLIOCENE(Junturabeds).OREGON:Malheur County: Jun tura. Suborder PLATALEAE NewtonPlataleaeA. Newton, ] 884, EJleydop. brit., ed. 9, vol.]8,p. 47(typePlataleaLinnaeus).[bidesCoues, 1BB4 (Aprilorlater),Key N. Amer. Birds, ed. 2,pp.ix,648(typeIbisCuvier, a synonymofThreskiornisCray).FamilytPLEGADORNITHIDAE(Wetmore)Pelagoclornithidae[sic] Wetmore, 1962(June26),Smithsonian misc. Call., vol. 145, no. 2, p. 3(typePlegadornis Wetmore,).-Pelagodornithoidea[siclWetmore, 19H2, op. cit., p. ,3 (superfamily).GenustPlegadomisWetmorePlegadornisWetmore,19H2(June26),Smithsonian misc. Call., vol.14,5,no. 2, p. 1(typebyoriginal designation PlegadornL' antecessorWetmore).

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS 2771.Plegadomis antecessorWctmorcPlegadornis antecessor\Vetmore, 1962 (June 26),Smithsonian misc.Col]"vol.145, no. 2, p. 1, fig. 1(typefrom Hewletts farm, distalpartoflefthumerus,U.S.Nat. Mus. no.22820).UPPERCRETACEOUS,SANTONIAN(MoorevilletongueofSelmachalk).ALABAMA:GreeneCounty:Hewlettsfarm, 3 milesnortheastofBoligee.FamilyPLATALETDAEBonaparte Plataleitwe Bonaparte, 1838, Geographicaland comparative list ofthe birdsofEuropeand North America, p.48(subfamilia;typePlatalea Linnaeus). PlataleidaeBonaparte, 18.53, C.R.Acad. Sci. Paris, vol. 37, no. 18, p. 643(familia). IZ,inae Bonaparte, 1853, C. R. Acad. Sci. Paris, vol. 37, no. 18, p.643(snbfamilia;type"lhis,Savig. Cuv. (Threskinrni, Wagl.)",see Bonaparte, 18.';4, Ann. Sci. nat.(Paris),p. :38; Ibisin this sense is preoccupiedby fbiliLal'epede,1799).-Ibididae Coues, 1884(Aprilorlater),Key N. Amer. Birds, ed. 2,Pl'.ix,648.PlegadidaeMathews, 1913(Jan.),Auk, vol. 30, no.1,Pl'.93, 95(typePlegadisKaup).ThreskiornithidaeRichmond, 1917 (Aug.16),Proc. U.S.nat. Mus., vol. 53, no. 2221,Pl'.580,636(typeThreskiornisCray).SubfamilyTHRESKIORNITHINAE(Richmond) IZ,inae Bonaparte, 1853, C. B. Acad. Sci. Paris, vol. 37, no. 18, p.643(type Ibis Savigny, Cuvier, notIbis Lacepedc). EudociminaeBonaparte,18.54,Ann. Sci. nat.(Paris),p.. 38 (typeEudocimusWagler).Threskiornithidae Richmond, 1917 (Aug.16),Pmc.U.S.nat. Mus., vol. 53, no. 2221,Pl'.580, 636 (family;typeThreskiornisGray).-ThreskiomithinaeWetmore and W.D. Miller, 1926, Auk, vol. 43, p.341).GenustIbidopsisLydekker Ibidop.sis Lydekker, 1891 (Apr.25),Cat.Foss. Birds Brit. Mus., p.74(typebyoriginal designation IZ,idop.is hordwelliensisLydckker).1.Ibidopsis hordwelliensisLydekkerIZ,idopsis hordwelliell.sis Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 74, fig. 20(typefrom Hordwell, distalpartofright tibiotarsus, Brit. Mus. no.36793).UPPEREOCENE(Hordwellbeds).ENGLAND:Hampshire:Hordwell.GenustIbidopodiaMilne-EdwardsIbidopodiaMilne-Edwards, 1868, Ois. Foss.France,vol. 1, sheet .59, p. 463(type bymonotypylbidopodia palustrisMilne-Edwards).

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278BULLETINFLORIDASTATEMUSEUMVol. 72.Ibidopodia palustrisMilne-Edwards 1I>idopodia palustris Milne-Edwards, 1868, Ois.Foss.France,vol. 1, sheet 59, 465, pI. 71, fig. 17-21(lectotypecraniumfromLangy,selectedbyLydek ker, 1891, p.74). LOWEll MIOCENE(Aquitanian).FRANCE:Dept. Allier: Langy. GenusEudocimusWagler Eu.dodmus \Vagler, 1832, Isis von Oken, p. 1232(typeScolopax rubra Linnaeus).3.Eudocimus paganus(Milne-Edwards) Ibis paganalVlilne-Edwards, 1868, Ois. Foss.France,vol. 1, sheet 57, p. 450, pI. 69-70, pI. 71, fip;. 1-12(typesnumerous elements fromLangyandSaint Gerand-1c-Puy) .LOWERMIOCENE(Aquitanian).FRANCE:Dept. Allier:Langyand Saint-Gerand-le-Puy(Milne-Edwards).Dcpt. Puy-de-Dome: Mon taigut(Lambrecht, 1\;''33, Handb. Palaeorn., p.331).'GenustProtibisAmeghinoProtibisAmeghino, 1891(Dec.1),Rev. arp;entina lIist. nat., vol.1,p. 445(typehymonutypyProtibis cnemialisAmcghino).4.Protibis cnemialisAmeghinoProtibis cnemialisAmeghino, 18!H (Dec.1),Rev. argentina Hist. nat., vol.1,p.44.5(typefrom MunteObservadon,distal partoftibiotarsus,nowinBrit.Mus.).-Ameghino,1895, Bul. Inst. geog. argentino, vol. 15, p. 98, fig. 42(typeredescribed). 'vfIDDLEMIOCENE (SantaCruzformation).ARGENTiNA:Ter. SantaCruz:Monte Observaci
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1ge,3BRODKORB: CATALOGUEOFFOSSILBIRDS 279LOWERPLEISTOCENE(CitaCanyonbeds),TEXAS:HandallCounty:CitaCanyon,atNewtonHarrell-EddHanch, Genus Carphibis Reichenbach Ca1'11hibis Reichenbach, 1853, Avium systema naturale,p.xiv(type Ibis spini collisJamesun),6,Carphibis condita(DeVis) 1M,!?) conditusDeVis, woe, Ann, Queensland Mus" no, 6, p, 10, pL 2,fig,2(typefrom vVurdulumankula, femur).UPPERPLEISTOCENE(Malkunifauna, Katipirisands).SOUTHAusTRALIA:WurdulumankulanearLakeEyre. SubfamilyPLATALEINAEBonapartePlataleinae Bonaparte, 1838, Grogr. cump. List Birds Enf. N. Amer.,p.48( typePlata leaLinnaeus) .GenusPLataleaLinnaeusPlataleaLinnaeus, 1758, Sy,!. Nat"ed. 10, vol. 1, p. 139(typePlatalea leucorodia Linnaeus).7.Platalea subtenuisDeVisPlatalea subtenuis DeVis, 1892, Pmc. Linn. Soc. N.S. Wales, SCI.2, vol. 6,p. 443, pl. 24,fig,5(typesfrom Queensland, fragmentary femurandtibiotarsus) UPPER PLEISTOCENE(DarlingDownsbeds).QUEENSLAND.Neospecies of Plataleidae from Pleistoceneand"prehistoric sites:LNipponia nippon(Tenuninck).JAPAN:'Iki Island? (Kuroda, 19,59, BulL biogeog. Soc. Japan, vul. 21,p.68, pl.1,fig.D-E).2. Theristicus caudatus(Boddaert).BRAZIL:Lapa da E.scrivania?(Winge,1887,E. MIlS. Lund.,voL1,no. 2, p,29),3,Eudocimus albus(Linnaeus).FLORIDA:SeminoleField(Wetmore,1931, Smithsonian misc, Call., vol. 85, no. 2, p,18);Haile (Brodkurb, 1953, Wilson Bull., vol. 65, p.49);ltehtuckneeRiver (McCuy, 1963, Auk, vol. 80, p.000),BAHAMAS:'GurdonHills un Crooked Island(Wctmorc, 1938, Auk, vol. 55, p.52), PU";RTO RICO:'BarrioCanas(Wetmore,1938, Auk, vol. 55, p.53).4. Eudocimus ru.ber(Linnaeus).VENEZUELA:Hacienda Tocoron? (\Vet more, 1935, Auk, vol. 52, p. ,329), 5. Plegadif; falcinellus(Linnaeus).PUERTORICO: BarrioCarta:s (Wetmore, 19,38, Auk, vol. 55, p.53).6.Plegadis chihi(Vieillot).CALIFORNIA:RanchoLaBrea(L.Miller, 1925, Publ. Carnegie Instn. Washington, no. 349, p.73);'Emeryville(Howard,1929, Univ. Calif. PubL Zool., vul. 32, p.312);'BuenaVista Lake(DeMay,1942, Condor, vol. 44, p.229).

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280 BULLETINFLORWASTATE MUSEUM Vol. 77. Platalea lIlba Scopoli. MADAGASCAn: 'Sirabe (Andrews, 1897, Ibis, 1'. 358).8. Aiuia oja;a (Linnaeus.).CALIFORNlA: Rancho La Brca?(Howard, 1930. Condor, vol. 32, p.84).FLORIDA: Roek Spring (Vvoolfenden, 19,59, Wilson Bull., vol. 71, p. 18,5). Suborder ARDEAEWaglerArdelleWagler, 1831, Isis von Oken, p. ,530 (ordo;typeArdeaLinnaeus).FamilyARilEIDAEVigors Ar
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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS281MIDDLEEOCENE(Bridgerformation).WYOMING:SweetwaterCounty: Spanish John's Meadow. GenusfProardeaLambrechtPrulIrdeaLambrecht, 1933,Handb.Palaeom., p. 311(typebymonotypyArdea amissu Milne-Edwards).4.Proardea amissa(Milne-Edwards)Ardeaarni8sa 1892, C.R.2.Congr.intemat.omith.Budapest,p. 73(typefrom phosphatedeChaux, tarsumetatarsus, ParisMus.).Ardea armissaParis, 1912, Rev. Onlith., vol. 4,p.291(lapsus).UPPEREOCENEorLOWEROLiGOCENE(phosphoritesduQucrcy).FRANCE: Dept. Tarn-ct-Garonne: Chaux. GenusfGoliathiaLambrechtGuliathiaLambrecht,1930(Jan.25),Geol. hungarica, ser. pal., fasc. 7, p. SO (typebymonotypyGaliathia andrewsiLamhrecht).5.Goliathia andrewsiLambrechtGoliathia andreswiLamhrecht,1930(Jan.25),Ceol. hungarica, ser. pal., fase. 7, p. 30,fig.7(typeulna, Brit. Mus. no.A.883).UPPEREOCENEorLOWEROLIGOCENE(Fayumseries).EGYPT;Fayum(exactlocalityunknown).Genusf Ardeadtes HaushalterArdeaeitesHaushalter, 1855, Merkwiirdige fussile Tieriiberreste ausdcrAllgauer Molasse, p. 11(typebymonotypyArdeaeites rnolassicusIlaushalter).6.Ardeacites molassicusHaushalterArdeacites molassicusHaushalter, 18,55, Merkwiirdige fossile Tiertiberreste ausderAllgaucr Molasse, p. 11, pI. 2,fig.1(typefrom Allgau,humems,Muuieh Mus.,nowlost).UPPERMIOCENE(obereMeeresmolasse).BAVARIA:AllgaunearHarbartshofen. GenusfBotauritesAmmonBotauritesvon Ammon, 1918, Abh. Naturw. Vcr. Regenshurg, vol. 12, p.31(typebymonotypy Bataurites avitu,y vonAmmon).7.Botaurites avitusAmmonBotaurites avitusvon Ammon, 1918, Abh. Naturw. Ver. Regensburg, vul.12,p.31, fig. 5-6(typefrom clay works,7thor8thcervical vertebra,Naturw.Verein zu Rcgcnshllrg.)

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282 BULLETINFLORIDA STATE MVSEUM Vol. 7UPPERMIOCENE(BraunkohlenderOberpfalz).GERMANY: Wiirt temberg:clay works of andReinhard,hetweenDechbettenandPriifening.8.Botauritessimi/is(Fraas)Ardell similisO. Fraas. 1870, Jahresh. Vcr. Naturk. \\'tirttembcrg, vol. 26, p. 284.pI.7,fig.14(typefrom Steinhcim, distalendof right tibiotarsus, StuttgartMus.).UPPERMIOCENE(obereSiisswassermolasse).GERMANY:Wiirttemberg: Steinheim. GenusArdeaLinnaeusArdellLinuaeus, 17.58,Sy,t. Nat., eel. 10, vol. 1, p. [41 (typeArdea cinerea Linnaeus) \I.Ardea aurelillnensis Milne-EuwardsArden uurelianensis Milne-Edwards. 1871, Ois.-FOS5.FranCE\ vol. 2, sheet 74.p. .58.5 (typefrom Suevres,humerus).UPPERMIOCENE(falnnsdeTouraine).FRANCE:Dept. Inure-et Loire: Suevres northeast of Tours.10.Ardea perplexa lvlilne-Edwardslirdea perplexaMilne-Edwards, 1868, Ois. Foss. France, vol.1.pI. 96,fig.1.3: 1869, vol. 2, shed 14, p. 108(typefrom Sans all, distalpartofrighthumerus).UPPERMIOCENE(gisement lac,:,stre deSansan).FRANCE:Dept.Gers: Sansan.n.Ardea brunhuberivon AmmonArdea hrunhuberivon Ammon, 1918, Ahh. Naturw. Ver. Regensburg, vol. 12, p. 30,fig.4(typefrom clay works, proximaleJJdof left metacarpus, Naturw.VcreinzuRegensburg),UPPERMIOCENE(BraunkohlenderOberpfalz).GERMANY:Wiirt temberg: clHy works of MayerandReinharu,betweenDechbettenandPriifening.12. Ardea polkensisBrodkorbA"dea /lolkensisBrvdkorb, 1955 (Nov.30), Florida Ceo!. Surv. Rept.Invest., no. 14, p. 17, pl. 4,fig. 13-1.5 (typefrom Brewster, proximalpartofrighttarsoilletatarsus, Brodkorb no.308).LOWER PLIOCENE (BoneValleygravel).FLORIDA:Polk County: Brewster.

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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS28313.Ardea lignitumGiebel Arclea lignitumGiebel, 1860(Sept.),Zeitschr. Naturwiss., vol. 16, no. 9, p. 152, pl.1, fig. 3(typefrom Rippersroda, distalpartof leftfemur).UPPERPLIOCENE(BraunkohlevonRippersroda).GERMANY:Thuringia:Rippersroda.GenusNycticoraxForsterNycticoraxForster, 1817, Syn. Cat. Brit. Birds, p. 59(typeArdea nyctieoraxLinnaeus).14. Nycticorax fidensBrodkorbNycticorax fide"sBrodkurb, 1963(Feb.8),Florida Geol. Surv. Spec. Publ.,no.2,paper4,p.3,pI.1(typefrom McGehee farm. left femur, Univ.Florida no.3285).LOWERPLIOCENE(Alachuaclay).FLORIDA:AlachuaCounty:C.C.McGeheefarm, section 22,Township9 South,Range17East,3.6 milesnorthofNcwbcrry.15.Nycticorax megacephulus(Milne-Edwards)Ardea megacephalaMiloe-Edwards, 1873, Bibl. Eeule hautesEtudesParis, sec. sci. nat., vol. 9, art. 3, p. 8, pI. 14,fig.1-14(typesfromRodriguez).QUATERNARY.RODRIGUEZISLAND.CenusfPalaeophoyxMcCoyPalaenpllOYxMeCuy 1963(inpress),Auk, vol. 80, no.,3,p. 000(type by uriginal designationPalaeophoyx columbianaMcCoy).16.Palaeophoyx columhianaMcCoyPalaeophoyx columbianaMcCoy, 1963(in press) Auk, vol. 80, no. 3, p.000fig. 1(typefrom Itehtuckncc River, right coracoid, Brodkorh no.32),UPPERPLEISTOCENE(ItchtuckneeHiveI'beds).FLORIDA:ColumbiaCounty:ItchtucknceRiver.GenusButoTidesBlythButoridesBlyth, 1852, Cat. Birds Mus. Asiatic Soc., p. 281(typeArdea iavanica Horsfield ).17.Butorides mauritianusGiintherandE.NewtonButorides mauritianusGuntherandE. Newton, 1879, Philo,. Trans. Roy. Soc. London, vol. 168, extra vol., p. 424, pI. 41,fig.a-f (types from Marc attx Songf!s, Cambridge Univ., casts in Brit. Mils.).QUATERNARY.MAUIUTIUSISLAND:Mareanx Songes.

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2S4 BULLETINFLORIDASTATE MUSEUM voL 7NeDspecies ofArcleitlaefrom lJei,toceneand prehistoric sitl's: l.Ardea cinereaLillnaeus. DENMARK: Ertcboelle,Maglcmose, and "'Barsmark (H. Winge, 1903, Vidcnsk. Medde!. naturhist. Foren. Copenh.gen, vol. 6, p.99). IRE"MW: Ballycolton,Ed"lValc,andNewhallcaves (Lambre<'ht, 1933.Handb.Palaeo"l.. p. 734). EN(;LANO: Clevedon Cave and GI.,tonburyce (I"unbrecht, 1933)Z.Aracahe""I;,,->Lmnoeus.OREW""FO"ilI"...(Shufeldt, 1913. .. ll. Amer. Mus. nat.lIi,t.,vol. 32,pp. 1.>, 157). CALlFOR""A: Rancho L,I3rea (L.Miller, 1909, Univ. Calif. Pub!. Geo], vol. .>, p.. 306), McKittrick(L. 1925, Univ. Calif. Pobl. GeoI., va!. 1.>, p.317);Emeryville(Howard,1929, UnIV. Calif. Pub!. Z""I., vo!.32,p. 31:).); "HuenaVistaLake(DeMay,1942, C.-J[ldor, vol. 44,p. 228). FLORIDA: Field, Itchtu<:knl'c River, Melh(Jurt',,Aradentun, and 1)Vl;TQBc-a<..:h stratum J(\Vdmore,un I, S[nLthscmianmbc ColL, vDI.1I5,no.2, p. I\oelr\Woo\!r",len,19:5\).Wiholl BulL. "01. 71,p.185);Good's ,hellpit and "LeIllDII Illuff (Neill, Cu!.l\rodkorh, 1956, Amer. Antiquity,vol. 21,p.388);'SouthInrhanField 1959,FloridaAn thropologist, va!. 12, p. 73);"CastleWindy(Bullen and Sleight, 1959, Rep!. Found.Arner. Studies, no. 1,p.20).ST.CROIX:"Concordia(WetmJsagliefe? (Lambrecht,1933,Handb.Palacorn.,p.734).5.Butorides vire.'iC(JfIS (Linnaclls).C.-\LIFORNIA: RancjJd La Bren.(Howard,1\J16. Condor,vol. :j8,p. 34); McKittrick (DeMay, 1941. Publ. Camegielo (Wcigel,1963, Spec. Pub!. Florid' gco!. Surv., no. 10,p. 25).fj, Casmerodiu8 alhlls (Liunaeus). CA-UFORNIA: Ranc:h<)LaBrea (HnwSmithsonianmisc. con ..vol. 85, no. 2,p.15); Itchtucknel:" River,(McCoy, 19B3, Auk,vo!. 80,p.000);"Hialeah(Laxson, 1953,Florida Anthropologist, vo!. 6,p.98);Good'sshcllpitand"Lemon Illuff (Neill,Gut,andBrodkorb,1956, Amer Antiquity,vol. 11. p.388);SouthIndianField (IVeigel. 1959, FloridaAndH'()pCllogist, voLL2, p. 73); O\'(;ro stratum3 (\Vdgel,196],Spt'L'Yllb\'gl?oL Surv., 1)1).to, p. CUDA:lh'Clego\1onh'TPl,Vf"t-

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1963 BRODKORB:CATALOGUEOFFOSSILBIRDS285more,1928, Amero Mus. Novit., no. 301,p.1). VENEZUBLA: LosTamarilldos(Wetmore,1935, Auk, vol. 52, p.329).7.Ardeola ralloides(Scopoli).ITALY:Buea del Bersagliere (Lambrecht,1933,Handb.Palaeoro., p.734).8.Florida caerulea (Linnaeus).CALIFORNIA:McKittrick(DeMay,1941, Publ. Carnegie Instn. Washington, no. 530, p.35):RanchoLaBrea(Howard,1962, Los AngelesCountyMus. Contr. Sci., no. 58, p.20).FLORIDA:SeminoleField(Wetmore,1931, Smithsonian misc. Call., vol. 8.'5, no. 2, p.16);ItchtuckneeRiver (McCoy, 1963, Auk, vol. 80, p.000).9. Fl011da thula(Molina).FLOIUDA:Bradenton(Wetmore,1931, Smithsonian misc. Call., vol. 85, no. 2, p.15);VeroBeachstratum3(Weigel,1963, Spec. Publ. Florida geol.SUIV.,no. 10, p.25).Tentativerecord from RanchoLaBrea, California(Howard,1936, Condor, vol. 38. p.35),withdrawn(Howard,1962, Los Angeles County Mus. Contr. Sci., no. 58, p.20).10.Egretta garzetta(Linuaeus). ITALY: Bersagliere(Lambrecht,1933,Handb.Palaeorn., p.734).11.Mesophoyx intermedia(Wagler).MADAGASCAR: Sirahe (Andrews, 1897, Ihis, p.358).12. Hydranassa tricolor (MiilIer).FLORIDA:SeminoleField(Wetmore,1931, Smithsonian misc. Call., vol. 85, no. 2, p.15).13. Nyctanassa viol"cea (LinnaCllS).FLORIDA:Seminole Field(\Vetmore,1931, Smithsonian misc. Coil., vol. 85, no. 2,p.16); Beach stratum 3(Larus vera Shufcldt, 1917, Jour. Geol., p. 18,typeleft carpomctacarpus, formerly Florida Geol. Surv. no. V320,nowinU.S.Nat. Mus., cast coli.Brodkorb; see vVetmorc,1931).ST.CROIX: "4Cuncordia (Wetmore, 1937, Jour.Agr. Univ. Puerto Rico, vol. 21, p.7).ST.THOMAS:midden(Wetmore,1918, Proc.U.S.nat. '>lus., vol. 54, p.515).ANTIGUA:MiIl Reef midden(Univ.Florida).14.Nycticurax ny<:ticorax (Linnacus).CALIFORNIA: Ri.lncho La Brea (Howard,1929, Condor, vol. 31, p.252): '>IcKittrick (L.Miller, 193.'5, Condor, vol, 37, p.75):"Buena Vista Lake(DeMay,1942, Condor, vol. 44, p.228).FLORIDA:Bradenton and Itchtucknce River (\Vetmore, 1931, Smithsonian misc.Coil., vol. R,'5, no. 2, p.16);Rock Spring (Woolfenden, 19.59, Wilson Bull., vol.71, p.185).NUEVO LEON: San Jasedta cave(L.Miller, 1943, Univ. Calif. Publ. Zool., vol. 47, p. 1.50). 15. Ixohrychus m-inutus (Linnaeus). ITALY: Buea delBersagliere? ('Lam brecht, 19-33, Handb.Palaeorn., p.734).16.hohrychusexilis(Gmelin).CALIFORNIA:"BnenaVistaLake(DeMay,1942, Condor, vol. 44, p. 228). CUIlA: Banos deCiego Montero(Wetmore,1928, Amer. Mus. Novi!., no. 301, p.2).BRAZIL:LapadaEscrivania(0.Winge, lRR7, E Mus. Lund., vol. 1, no. 2, p.30).17. Botaurtts stellaris(Linnaells).DENi\fARK: lvlaglcmose and 4Rodals !\1ose (H.Winge, ] 903, Vidensk. Meddel. naturhist. Foren. Copenhap;en, vol. 6, p.99).ENGLAND:Cambridgeshire (Milne-Edwards, 1868, Ibis, p.364):Burwell fens, Reach fens, andGlastonbury(Lamhrecht, 1933, Handb.Palacorn., p.734).FUANCE:tourhieres (Milne-Edwards, 1871, Ois. Foss.France,vol. 2, p.601). 18. ButuuftlS lentiginos:us (Rackett).OREGON;Fossil Lake (includesArdeapalnccidentalisShufeldt, 1892, Jour. Acad. nat. Sci. Philadelphia, vol. 9, p. 411,pI.17,fig.31,typedistal partofright tarsometatarslIs,Arner. Mus. Nat. Hist.

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286BULLETINFLORIDASTATEMUSEUM Vol. 7 no. 3483; seeHoward,1946, Publ. Carncgie 1nstn. Washington,nO.551, p.156).CALIFORNIA:RanchoLaBrea(L.Miller, 1921, Condor, vol, 23, p.129);"BnenaVista Lake(DeMay,1942, Condor, vol. 44, p.228).NORTHDAKOTA:"MortonConnty(L.Miller, 1961, Bull.S.Calif. Acad. Sci., vol. 60,pt.3, p.125).IOWA:"Mill Creek(Hamon,1961, Plains Anthropologist, vol. 6, p.209).FLORIDA:SeminoleFieldandHogCreekatSarasota(Wetmore,1931, Smithsonian misc. Coli., vol. 85, no. 2, p.17);Rock Spring (Woolfenden, 1959, Wilson Bull., vol. 71, p.185);Vera Beach(Weigel,1963, Spec. Publ. Florida geol. Surv., no. 10, p.26).FamilyCOCHLEARIIDAERidgwayCancromidaeBonaparte, 1853, C.R.Acad. Sci. Paris, vol. 37, no. 18, p. 643(typeCancromaLinnaeus,1766,a juniorsynonymofCochleariusBrisson,1760).CochleariidaeRidgway, 1887, Manual N. Amer. Birds, p. 122(typeCochleariusBrisson),Nofossil record. FamilySCOPIDAE(Bonaparte)ScopinaeBonaparte, 1853, C.R.Acad. Sci. Paris, vol. 37,nO.18, p. 643 (sonsfamille; type ScopusBrisson).Nofossilrecord,FamilyBALAENICIPITIDAE(Bonaparte)BalaenicepinaeBonaparte, 1853, C. R. Acad. Sci. Paris, vol.37,no. 18, p.643(sous-famille;type Balaenicep. Gould).No fossil record.SuborderCICONIAEBonaparteTantaliWagler, 1831, IsisvonOken, p,530(ordo;typeTantalusLinnaeus, 1758, a synonym ofMyeteriaLinnaeus,1758).CiconiaeBonaparte, 18.54, Ann. Sci. nat.(Paris),p. 37("tribus,"i.e. suborder;typeCiconiaLinnaeus).FamilyCrcoNHDAE(Gray)TantalidaeBonaparte, lR31, Saggio di una distribuzione metodica degli Animali Vertebrati,p.57(typeTantalusLinnaeus, asynonymofMycteriaLinnaeus).CiconiinaeGray, 1840, ListGeneraBirds, p.000(typeCiconiaBrisson),MycteriinaeAmerican Ornithologists' Union, 1908, Auk, vol. 25, no. 3, p. 363 (type MycteriaLinnaeus).SubfamilyCrCONHNAEGrayCiconiinaeGray, 1840, ListGeneraBirds, p. 000(typeCiconiaBrisson).

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1963 BRODKORB: CATALOGUEOFFOSSILBIRDS 287GenustPelargopappusStejneger Pewrgopsis Milne-Edwards, 1868(afterApril), Ois. Foss. France, vol.1,sheet 58, p. 460(typebyoriginal designation Pewrgopsis magnusMilne-Edwards). Preoccupied byPelargopsisGloger, 1841.PelargopappusStejneger, 1885, Stand. nat. Hist., vol. 4, p. 163(newnameforPelargopsisMilne-Edwards).PelargoidesLydekker, 1891, Nature, vol. 45, p. 71(newnameforPelargopsisMilne-Edwards) .PelargodesLydekker, 1892(Apr.1),Proc. zool. Soc. London for 1891, p. 477 (emendation ofPelargoidesLydekker). Pewrgocrex Milne-Edwards, 1893 (July4),Bull. Brit. ornilh. Club, vol.1,p. 54(newnameforPelargopsisMilne-Edwards).1.Pelargopappus stehlini(Gaillard)PelargopsisstehliniGaillard, 1908, Ann. Univ. Lyon, n.s., vol. 1, fase.23,p.82,text-fig. 21, pI.4,fig.5-8(typefrom Quercy, distal end of right tarsometa tarsus, Basel Mus.no.QH.146).UPPEREOCENEorLOWEROLIGOCENE(phosphoritesduQuercy).FRANCE:plateauof Quercy.2.Pelargopappus trouessarti(Gaillard) Pewrgopsis trouessartiGaillard, 1908, Ann. Univ. Lyon, n.s., vol. 1, fasc. 23, p. 84, text-fig. 22, pI. 4,fig.9-12(typefrom Quercy, distalendof left tarso metatarsus, Basel Mus. no.QH.147).UPPEREOCENEorLOWEROLIGOCENE(phosphOritesduQuercy).FRANCE:plateauof Quercy.3.Pelargopappus magnus(Milne-Edwards) Pewrgopsis magnusMilne-Edwards, 1868(afterApril), Ois. Foss. France, vol. 1, sheet 58, p. 460, pI. 72,fig.1-19 (lectotype from Langy, distalpartof tarsometatarsus, Paris Mus.,designatedbyLydekker,1891,Cat. Foss. Birds Brit.Mus., pp.67-68).LOWERM,OCENE(Aquitanian).FRANCE:Dept.Allier:LangyandSaint-Gerand-le-Puy (Milne-Edwards, 1868).Dept.Puy-de-Dome (Lydekker, 1891). GenustPropelargusLydekkerPropelargusLydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 65(typebyoriginal designationPropelarguscayluxensisLydekker).4.Propelargus cayluxensisLydekkerPropelargus cayluxensis Lydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 66,fig.16(typefrom Bach, distalpartof right tarsometatarsus, Brit. Mus. no. A.109).

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BULLETINFLORIDASTATE MUSEUM Vol. 7UPPEREOCENEorLOWEROLIGOCENE(phosphoritesdeBach).FRANCE:Dept.Lot:Bach.5.Propelargus edwardsiLydekkerPropelargus edwardsiLydekker, 1891(Nov.3),Proc. zool. Soc. London, p. 479,fig.3(typesfrom Saint-Gerand-Ie-Pny,rightcoracoid, left metacarpus, Brit. Mus.).LOWERMIOCENE(Aquitanian).FRANCE:Dept. Allier: Saint Gerand-le-Puy.6.Prapelargus olseniBrodkorbPropelargus olseniBrodkorb, 1963(inpress), Quart. Jour. Florida Acad. Sci., vol. 26, no. 2, p. 000, fig. 00(typefrom Tallahassee, left tarsometatarsus, Brodkorb no.8504).LOWERMIOCENE(Hawthorneformation).FLORIDA:LeonCounty: Tallahassee, SwitchyardB,SeaboardAirline Railroad Company. GenustPalaeoephippiorhynchusLambrechtPalaeoephippiorhynchusLambrecht, 1930(Jan.2.5), Geol. hungarica, ser. pal., fasc. 7, p. 18(typeby monotypyPaweoephippiorhynchus dietrichiLambrecht).7.PalaeoephippiOl'hynchlls dietrichiLambrechtPaweoephippiorhynchus dietrichiLambrecht, 1930 (Jan. 25),Geol. hungarica, ser. pal., fasc. 7, p. 18, pI. 3,fig.1-4(typefrom Qasr-el-Qurun, skull, man dible, Naturaliensammlung,Stuttgart).LOWEROLIGOCENE(Fayumseries, fluviomarinebeds).EGYPT:Fayum:Qasr-el-Qurun. GenustCiconiopsisAmeghinoCiconiopsisAmeghino, 1899(July), geo16gico-paleontologica, Suplemento, p. 8(typebymonotypyCiconiopsis antarcticaAmeghino).8.Ciconiopsis antarcticaAmeghinoCiconiopsis antarcticaAmeghino, 1899(July),Sinopsisgeologico-paleonto16gica,Suplemento, p. 8(typefrom "formacion guaranitica,"metacarpus).LOWEROLIGOCENE(Deseadoformation).ARGENTINA:Patagonia. GenustAmphipelargllsLydekkerAmphipelargusLydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus., p. 68(typebyoriginal designationAmphipelargus majoriLydekker).

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196.1BRODKORB: CATALOGUEOFFOSSIL BIRDS 2899.Amphipelargus mujuriLydekkerAmphipelargas ma;ariLydekker, 1891 (Apr.25),Cat. Foss. Birds Brit. Mus.,p.69,fig.18(typefrom Samos, distalendof left tibiotarsus, Brit. Mus. no. A.123).LOWERPLIOCENE(Samosbeds).GREECE:Samos Island, GenusLeptoptilosLessonLeptapti/asLesson, 1831, Trait" d'Ornithologie, livr. 8, p. 583(typeArdea dubioGmelin).10.Leptoptilos falconeri(Davies)Argala falconeriMilne-Edwards, 1868, Ois, Foss. France, val. 1, sheet 56, p. 449 footnote(nomennudum).-Davies,1880, Geol. Mag., decade 2; val. 7, p. 24, pI. 2,fig.4 (lectotype from Siwalik Hills, distalpartof right tibiotarsus, Brit. Yills. no. 39753, desigllaled by Lydekker, 1884, Mem. geol. Surv. India, Palaeontologia indica, ser. 10, val. 3, pt.4,p.139).LOWERPLIOCENE(Siwalik series).INDIA:UnitedProvinces: Siwalik Hills(Davies,1880).Punjab(Lydekker,1884).11.Leptoptilos titanWetmoreLeptoptilos titanWetmore, 1940(Sept.),Jour. Paleont., vol. 14, no. 5, p. 447,fig.1-5(typefromWatoealang,lefttarsometatarsus,MiningandGeologicalSurvey, Dept. Netherlands Indies, no.3313).UPPERPLEISTOCENE(Solo Riverbeds),JAVA:Watoealaug,nearSolo River. GenusCiconiaBrisson Cirania Brisson, 1760. Ornithologia, val.I,p. 48; val. 5, p. 361(typeArdeaciconiaLinnaeus).12.Ciconia gaudryiLambrechtCicania gaadryiLambrecht, 1933,Handb.Palaeorn., p. 323(typefrom Pikermi,humerns, ParisMus.).LOWERPLIOCENE(Pikermiredclay).GREECE:Attica: Pikermi.13.Ciconia tlwltha L.MillerCiconia rnaltha L.Miller, 1910 (Aug.5),Univ. Calif. PlIbi. Geol., vol. 5, no. 30, p. 140,fig.1-7(typefrom RanchoLaBrea, left ta'Sometatarsus, Univ. Calif. Mus. Paleo. no. 11202).labira? weillsiSellards, 1916,EighthAnn. Rept., Florida geol. Surv., p. 146, text-fig. 15c, pI. 26,fig.1-4(typefrom Vero Beach, right humerus, formerly Fla. Geol.SUTY.no. 5961, now in U.S.Nat.MilS.,cast call. Bradkorb).

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290BULLETIN FLORIDA STATE MUSEUMVol. 7MIllDLEPLEISTOCENE(Bruneauformation).IDAHO:OwyheeCounty: Barbour Ranch. 3.8 miles east of Bruneau-MountainHomebridge(L.Miller, 1944, Condor, vo!. 46, p.27).UPPERPLEISTOCENE(Amerioan Falls lakebeds.).IDAHO:PowerCounty: American Falls(Howard,1942, Pub!. Carnegie Instn. Wash ington, no. 530, p. 189).UPPERPLEISTOCENE(tarpits).CALIFORNIA:Los Angeles County: RanchoLaBrea(L.Miller, 1910). Santa Barbara County: Carpinteria(L.Miller, 1931, Univ. Calif. Pub!. geol. Sci., vo!. 20, p. 366). Kern County: McKittrick(L.Miller, 1935, Condor, vol. 37, p.75).UPPERPLEISTOCENE(Manix lakebeds).CALIFORNIA:San Bernar dino County: Manix(Howard,1955, U.S.geol. Surv., profess. Paper,nO.264-J, p. 202).UPPERPLEISTOCENE(ItchtllckneeRiverbeds).FLORIDA:Columbia County: Itchtucknee River(Wetmore,1931, Smithsonian misc. Coli., vol. 85, no.2,p. 17).UPPERPLEISTOCENE(Pamlico formation).FLORIDA:Flagler County: BonTerraFarm, 6" miles south of Marineland(Howard,1942).OrangeCouuty: Rock Spring (Woolfenden, 1959, Wilson Bull., vol.71,p. 18.'5). Brevard County: Melbourne(Wetmore,1931). IndiauRiver County:WinterBeach (Brodkorb coli.); Vero Beach (Sellards, 1916). Pinellas County: "Seminole Field" in St. Petersburg(Wetmore,1931). Sarasota County: Venice(Wetmore,1931);WarmMineral Springs (Brodkorb coIL).UPPERPLEISTOCENE(springs deposits).CUBA:Prav. Santa Clara: BanosdeCiego Montrero(Wetmore,1928, Amer. Mus. Novit., no. 301, p.2).GenusfPelargosteonKretzoi Pelnrgosteon Krelzoi. 1962(Feb.).Aquila, vol. 67-68, p. 169(typeby monotypyPelargosteon tothi Kretzoi).14.Pelargosteon tothiKretzoi Pelnrgosteon tothi Krelzoi, 1962(Feb.),Aquila, vol. 67-68, p. 169(typefrom Betfia no..5,fragmentary sternum, Oradea Mus. no.1899/1).UPPERLOWERPLEISTOCENE(Biharianfauna).RUMANIA:Betfia. GenusfProciconiaAmeghinoProciconiaAmeghino, 1891(Dec.1),Rev. argentina Hist. nat., vol.1,p. 445(typebymonotypyProciconia lydekkeriAmeghino).

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1963 BRODKORB:CATALOGUEOFFOSSILBlRDS29115.Prociconia lydekkeriAmeghinoProciconia lydekkeriAmeghino, 1891(Dec.1),Rev. argentina Rist. nat., vol. 1,p.445(newnamefor"Palaeociconia australis, Moreno," Lydekker, 1891,Cat. Foss. Birds Brit. Mus., p. 65,fig.15; types fromLagoaSanta, distal ends of rightandleft tarsometatarsi, Brit. Mus. nos. 18878,18879).UPPERPLEISTOCENE(caverndeposits).BRAZIL:Minas Geraes:cavenearLagoaSanta.1GenusfPalaeopelargusDeVisPaZaeopelargu8DeVis, 1892, Froc. Linn. Soc. N.S.Wales, ser. 2, vol. 6,p.441 (type byIIlonotypyPulueupeluTgus ,wbilisDeV;"). 16.Palaeopelargus nobilisDeVisPalaeopelargus nobilis DeVis, 1892,Proc. Linn. Soc. N.S.Wales, ser. 2, vol. 6, p. 441, pI. 20,fig.4(typefrom Queensland, distalpartofcarpometacarpus).UPPERPLEISTOCENE(DarlingDownsbeds).QUEENSLAND.GenusfXenorhynchopsisDeVis Xenorhynchopsis DeVis, 1906, Ann. Queensland Mus., no. 6, p. 9(typebypresentdesignationXenorhynchopsis tibialisDeVis).17.Xenorhynchopsis tibialisDeVisXenorhynchopsis tibialisDeVis, 1906, Ann. Queensland Mus., no. 6,p.10,pI.1,fig.6(typesfrom Lower Cooper, distalendsofrightandleft tibiotarsi).UPPERPLEISTOCENE(Katipirisands, Malkunifauna).SOUTHAusTRALIA:lowerCooperCreek, east ofLakeEyre.18.Xenorhynchopsis minorDeVis Xenorhynchopsis minorDeVis, 1906, Ann. Queensland Mus., no. 6, p. 10, pI. 2,fig.1(lectotypebypresentdesignation, fromUnduwampa,distalendofrighttibiotarsus ).UPPERPLEISTOCENE(Katipirisands, Malkunifauna).SOUTHAusTRALIA:UnduwampaandWurdulumankula(DeVis,1906).GenusXenorhynchusBonaparteXenorhynchusBonaparte, 1855, Conspectus generum avium, vol. 2, p. 106(typeMycteria australisShaw).'Referredwithoutsupporting evidence to genusJab/ruRellmayrbyPattersonandKraglievich (1960, Pub!. Mus.Mardel Plata, vol. 1, p. 8, footnote).IfLydekker's figure is accurate, such actionisunwarranted.Ciconia malthaMiller needs comparison with this species.

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292BULLETINFLORIDASTATEMUSEUMVol. 719.Xenorhynchus nanusDeVisXenorhynchus nanus DeVis, 1888, Proc. Linn. Soc. N.S. Wales, vol. 3,p. 1287, pI.35,fig.11(lectotypebypresentdesignation. fromDarlingDowns, distalpartofright tihiotor
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1963 BRODKORB: CATALOGUEOFFOSSIL BIRDS 2933.Euxenura galeata(Molina).ARGENTINA: (l,Llajta-Maiica and 4Las Represasin Santiago del Estero (KraglievichandRusconi, 1931, Physis, vol. 10, p.240).4. Ibis ibis(Linnaens).SARDINIA:bone breccia(Tantalus hresciensis Giebel. 1847, Fauna der Vorwelt. vol. 1,pt.2,pp.28, 40:typeulna; a nomen nudumhere, possibly previously describedbydela MarmoraorKeferstein).5.Ibis leucocephalus(Pennant).INDIA:Karoul district in Madras (Lydekker, 1891, Cat. Foss. Birds Brit. Mus., p. 70,fig.19).6. Mycteria americanaLinnaeus.FLORIDA;Itchtucknee River(McCoy,1963,Auk, vol. 80, p.000);CastleWindy(BullenandSleight, 1959, Rept. Bryant Found. Amer. Studies, no. 1, p.20).VENEZUELA:'LosTamarindos (Wetmore, 1935, Auk, vol. 52, p.329).

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Catalogue of fossil birds Part 1 (Archaeopterygiformes through Ardeiformes)
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00001514/00001
 Material Information
Title: Catalogue of fossil birds Part 1 (Archaeopterygiformes through Ardeiformes)
Alternate title: Bulletin of the Florida State Museum ; volume 7, number 4
Physical Description: Book
Language: English
Creator: Brodkorb, Pierce
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1963
 Record Information
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA2634
ltuf - ACK0908
System ID: UF00001514:00001


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Table of Contents
    Front Cover
        Front page 1
        Front page 2
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
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Full Text







Volume 7

Number 4

Pierce Brodkorb





lished at irregular intervals. Volumes contain about 300 pages and are not nec-
essarily completed in any one calendar year.


Consultants for this issue:
Elliot W. Dawson
Hildegarde Howard
Alexander Wetmore

Communications concerning purchase or exchange of the publication and all man-
uscripts should be addressed to the Editor of the Bulletin, Florida State Museum,
Seagle Building, Gaincsville, Florida.

Price for this issue $1.40

Published 19 June 1963



SYNOPSIS: New information from unpublished sources and from published rec-
ords hitherto overlooked permit a re-evaluation of the history of the Dry Tortugas
and of the terns that inhabit them. The geography and ecology of the 11 keys
that have variously comprised the group since it was first mapped in the 1770's
are described and their major changes traced. The recorded occurrences of the
seven species of terns reported nesting on the keys are analyzed in detail. The
Sooty Tern colony has fluctuated from a low of about 5,000 adults in 1903 to a
reported peak of 190,000 in 1950; for the past four years it has remained steady
at about 100,000. The Brown Noddy population, which reached a peak of 35,000
in 1919, was reduced by rats to about 400 adults in 1938; it is in the neighbor-
hood of 2,000 today. A colony of 150 to 450 Roseate Terns has nested in most
years from 1917 to the present. About 500 Least Terns nested regularly from
1916 to 1932, then unaccountably dwindled to a few pairs by 1937 and shortly
afterward disappeared. Royal and Sandwich Terns nested abundantly in the
mid-19th century, and a colony of Royals may have existed as late as 1890.
Both species are believed to have been extirpated from the Tortugas by egging.
No verifiable evidence exists for the nesting of the Common Tern, which has
been reported several times. The Black Noddy, first reported for the continental
United States at Dry Tortugas in 1960, has been found there each summer since.

'The author is Park
National Monument,

Biologist at Everglades National Park and Fort Jefferson
Homestead, Florida. Manuscript submitted 10 October

Robertson, William B., Jr., 1964.
State Mus., vol. 8, no. 1, pp. 1-95.

The terns of the Dry Tortugas. Bull. Florida


Introduction .
Acknowledgments -----
Location and Physiography
History and Names of the Tortugan Keys
Sooty Tern .....
Record of Nesting -.
Discussion -. ..-..-
Brown Noddy ........
Record of Nesting -
Discussion .........--
Roseate Tern
Discussion ...........
Least Tern ..... --
Common Tern
Royal and Sandwich Terns-- --
Black Noddy ..........
Literature Cited ...-

-------- -- 3

------- 17
........ 17
.---- 38
------.. 52
...... 64
--- -.-.- 73
............ 75
--------- 75
.......... 79


The tern colonies of the Dry Tortugas, in particular the great breed-
ing aggregations of the Sooty Tern, Sterna fuscata Linnaeus, and
the Brown Noddy, Anous stolidus (Linnaeus), have been of interest
to ornithologists since Audubon visited them in 1832. Although the
area is remote and difficult of access even today, few bird colonies
in North America can boast so long a record of observations or so
extensive a literature.
During the early years of the Carnegie Institution of Washington's
Tortugas Laboratory, John B. Watson and his co-workers made ex-
tended observations on Sooty Terns and Brown Noddies (Watson,
1907, 1908, 1910; Watson and Lashley, 1915; Lashley, 1915). Their
work provided nearly all of the detailed life history data available for
these species until recently. It also included pioneer experimental
studies of behavior, homing, and orientation, as well as an early in-
stance of the use of metal leg bands to mark birds.
Excepting the work of Watson and his associates, the literature
consists almost entirely of descriptions of the ternery as observed
during brief visits. Many accounts since 1900 include estimates of
the number of Sooty Terns, Brown Noddies, and other breeding spe-


cies. Lengthier studies by National Park Service personnel in 1937
and 1938 (Beard, 1939) were concerned particularly with predation
upon Sooty Tern chicks by Magnificent Frigate-birds, Fregata mag-
nificens Mathcws. Parts of the historical record were summarized by
Bartsch (1919), Vinten (1943), Sprunt (1948b), and Moore and Dilley
Modern banding at the Dry Tortugas began with the activities
of Jack C. Russell in 1936 and was continued annually through 1941,
principally on outings sponsored by the Florida Audubon Society.
About 13,300 Sooty Terns and 246 Brown Noddies were banded.
The bandings were reported separately by eight or more individuals
and no analysis of the data was undertaken until recently (Austin,
1962 Ms.).
In June 1959, the National Park Service, Florida State Museum,
and Florida Audubon Society began a cooperative mass-banding
study of the movements and demography of the Sooty Tern popula-
tion. At the end of the 1963 season new bandings of Sooty Terns
by project cooperators totalled approximately 32,300 adults and 41,900
juveniles. In the course of this work it became evident that a number
of the widely scattered published reports and much unpublished in-
formation had not been taken into account by previous compilers.
Because of this, several apparent misinterpretations of the history
of the colony had gained wide currency. The present summary
The names of birds are those of the Check-List of North American
Birds, American Ornithologists' Union, 1957, except for the changes
resulting from the recent discovery of Anous tenuirostris (Temminck)
at the Dry Tortugas (Robertson et al., 1961).

It is possible to mention here only a few of the people who helped
me to assemble the data this paper summarizes. The cooperation
of those named and many others contributed greatly to this review.
I am particularly indebted to Joseph C. Moore for permission to
refer to the unpublished reports of tern censuses he made in 1953,
1954, and 1955. C. Russell Mason also made extensive field notes
available to me. Others who contributed unpublished data or photo-
graphs included Robert P. Allen, H. G. Deignan, John R. DeWeese,
Willard E. Dilley, Theodore R. Greer, David O. Hill, James B.
Meade, Dennis R. Paulson, Roger T. Peterson, Cbandler S. Robbins,
Alexander Sprunt IV, and Louis A. Stimson.


Albert Manucy provided a wealth of information from his re-
search on the history of Dry Tortugas and useful advice on historical
sources. Of those who helped me to obtain copies of rare publica-
tions, maps, and material from archives, I must thank in particular
Luis R. Arana, Charles M. Brookfield, C. Gordon Fredine, Lowell
Sumner, and C. R. Vinten. Charles I. Park, Julius F. Stone, Jr., and
C. C. Von Paulsen gave me their recollections of Dry Tortugas in
the late 1920's and early 1930's, a period for which little written
record exists.
Recent work at Dry Tortugas has depended greatly upon the
cooperation of National Park Service personnel in the area particu-
larly District Manager and Mrs. Wallace B. Elms, District Manager
James A. Olson, and Park Rangers Roy Evenson, Carl S. Christensen,
and James E. Markette.
Finally, I am grateful to Oliver L. Austin, Jr. for assistance in
locating references, friendly encouragement, and many helpful com-
ments on the manuscript.

The Dry Tortugas, the westernmost outliers of the Florida Keys, are
an area of shoals with several small, low islands located about 70
miles west by slightly north of Key West (figure 1). The shoals
have the shape of a roughly elliptical atoll with its long axis north-
east-southwest. They enclose a lagoon about 10 miles in greatest
diameter, its center lying at approximately 24040'N, 82052'W. The
10-fathom line closely approaches the outer perimeter of the shoals.
Depths within the lagoon are mostly 5 to 10 fathoms. According to
Vaughan (1914) the shape and alignment of the shoals were deter-
mined primarily by currents and antedate the present luxuriant growth
of reef corals. The nearest land is the Marquesas Keys, about 50
miles east.
The islands of the Dry Tortugas (Vaughan, 1914; Davis, 1942)
are made up of coarse, unconsolidated calcareous sand and larger
detrital fragments, chiefly the remains of lime-secreting marine or-
ganisms. Skeletons of corals predominate. Because of the strong
currents and heavy wave action during storms, little fine sediment
accumulates and the shorelines of the islands change frequently.
Highest elevations on most of the present Tortugan islets do not
exceed 3 or 4 feet above normal high tides. Except for Garden Key
and Loggerhead Key, all are subject to some overflow by storm tides.

Vol. 8







'2 r
-c U

ia s *J

i510 1

in2 i
Wit a






/ o



*^ I





According to Herrera's chronicle of the first Florida voyage (Davis,
1935: 21), Juan Ponce de Leon reached the Tortugas 21 June 1513.
The islands had been sighted from the east as the expedition was
rounding the tip of the Florida Keys some weeks earlier. Herrera
speaks of an archipelago of "eleven rocky islets" named "Las Tortu-
gas" because many sea turtles were captured there. The Tortugas
offered a protected anchorage where sea birds, turtles, and seals
(presumably the West Indian Seal, Monachus tropicalis, now ex-
ceedingly rare if not extinct) could be taken to augment a ship's
food supply. It is likely that the islands were visited frequently
during the 250 years following their discovery, but little record of
this period survives.


Cauld Chart Chart Coast Survey Chart 471a Chart 585
1773-75 1829 1853-54 1868-75, 1896 1958

Booby Kay Bird Key Bird Key Bird Key
Long Key Long Key Long Key Bush Key
East Kay East Key East Key East Key East Key
Bush Kay Garden Key Garden Key Garden Key Garden Key
Middle Kay Sand Key Sand Key Sand Key Hospital Key
Logger Head Loggerhead Loggerhead Loggerhead Loggerhead
Turtle Kay Key Key Key Key
Rocky Kay Bush Key Bush Key Bush Key Long Key
Bird Kay Middle Key Middle Key Middle Key Middle Key
North Kay North Key North Key
Sandy Kay North East North East
Key Key
South West South West South West
Kay Key Key

SBlanks indicate that no island existed at the time of the survey.

The first modern chart, and the earliest I have seen that gives
names to the individual keys, was based on a survey made by George
Gauld for the British Admiralty in 1773-75 (Gauld, 1790). Gauld's
chart applies the name "Dry Tortugas" to the group as a whole and
shows 10 keys; the names it gives for 6 of these differ from those
used later (table 1). The Dry Tortugas were next charted by Lieu-

Vol. 8


tenants Josiah Tatnall and G. R. Gednery for the United States Navy
Department in September 1829. A tracing of this chart is in the
files of Castillo de San Marcos National Monument, St. Augustine,
Florida (C. R. Vinten, in litt.). The 1829 chart has particular value
because it gives areas and elevations for 6 of the 11 keys then
emerged. Parties from the United States Coast Survey worked at
the Tortugas in 1853-54 ("Tortugas Island", Scale 1:31,680; and "Sec-
tion No. VI", Scale 1:400,000, in Bache, 1858), and in 1868-75 ("T-
1410", Scale 1:10,000, in Coast Survey, 1878; Chart 471a, "Tortugas
Harbor and Approaches", Scale 1:40,000, United States Coast and
Geodetic Survey, 1896). The chart of the area presently in use is
Coast and Geodetic Survey 585, "Dry Tortugas", Scale 1:30,000, first
issued in 1922 and last revised in 1958. Table 1 shows the keys of
the Dry Tortugas that existed at the time of each of the above sur-
veys and the names applied to them on the various charts.
Two general types of keys may be distinguished in the Dry Tor-
tugas, those little more than barren sandbanks slightly elevated above
normal tides, and the larger, higher, and usually more permanent
islands with considerable plant cover. The first group includes Hos-
pital, Long, Middle, North, Northeast, and Southwest Keys; the
second, Bird, Bush, East, Garden, and Loggerhead Keys.

Bird Key was the principal nesting ground of Sooty Terns at the Dry
Tortugas from at least 1832 (Audubon, 1835) and of Brown Noddies
from at least 1857 (Wurdemann, 1861) until the island washed away
in the early 1930's. During periods of military activity at Fort Jeffer-
son, Bird Key also served at times as a hospital site, quarantine sta-
tion, and cemetery. The former hospital buildings later housed the
Audubon and Biological Survey wardens guarding the tern colony.
The 1829 survey recorded the area of Bird Key as "4 acres 2 roods
20 poles", slightly more than 41/ acres, and the elevation as "3 feet
8 inches" (Vinten, in litt.). Later comments on its area, dimensions,
and elevation vary widely. The area in 1890 was stated as "about
eight acres" (Scott, 1904: 278), in 1910-13 as "about 6,000 square
yards" (Watson and Lashley, 1915: 35) and as "somewhat less than 5
acres" (Lashley, 1915: 61), in 1915 as "8 acres" (Pearson, 1915: 412),
in 1918 as "about 6 acres" (Ashe and Lowe, 1918 Ms.), and in about
1926 as "less than five acres" (England, 1928: 14). Dimensions given
in various publications range from 500 x 250 feet in 1904 (Millspaugh,
1907: 233) to 400 x 300 yards in 1907 (Watson, 1908: 191), and the
key is credited with various elevations up to "6 feet above mean tide
level" (Watson and Lashley, 1915). A comparison of the representa-



tions of Bird Key on the charts of different periods suggests that much
of the reported variation existed mainly in the eye of the observer.
It is commonly stated that Bird Key was destroyed by a hurricane
in 1935, the Labor Day hurricane that devastated the Florida Keys
often being specified (Stevenson, 1938; Davis, 1942; Vinten, 1943;
Sprunt, 1946b, 1948b). Other authors cite the "hurricane of 1933"
(Robinson, 1940: 3; Peterson, 1950: 318) and "the big hurricane of
1938" (Peterson and Fisher, 1955: 142) as the storm responsible. Many
accounts suggest that the key was destroyed suddenly. Dilley (1950:
67) wrote: "At times changes may be very sudden, as illustrated by
the complete disappearance of Bird Key during the hurricane of
1935." Stevenson (1938) noted that Bird Key had been eroding grad-
ually for some time before the 1935 storm, and Robinson (1940),
Peterson (1950), and Peterson and Fisher (1955) state that it began to
"sink" in 1928.
The disappearance of Bird Key appears to have been an extended
process following destruction of the vegetation, and without immedi-
ate relation to any of the storms mentioned. In 1832 the key had
a thick cover of bushes (Audubon, 1835), and in 1857 Wurdemann
(1861: 426) described it as "covered with bay cedar [Suriana mari-
tima] bushes seven or eight feet in height interspersed here and there
with the cactus." Later descriptions of the vegetation up to 1910
are almost identical to Wurdemann's. As early as 1904, however,
some erosion had begun. Millspaugh (1907: 233) noted from Lan-
sing's observations: "Wave action from the northwest appears to be
rapidly eroding the western beach, the vegetation on the shore plain-
ly showing the encroachmentt"
The severe hurricane of 15-17 October 1910 (Tannehill, 1950:
175-176) was the first important event in the destruction of Bird Key.
Of its effects Lashley (1915: 62-63) wrote: "The Key was formerly
overgrown thickly with bay cedars, but the greater number of these
were killed by the hurricane of 1910 and only a few living cedars
remain." In 1915-16 the effects of the 1910 storm were still evident.
Bird Key then had only scattered patches of bay cedar bushes (Bow-
man, 1918: 124). On 10-11 September 1919, another severe hurri-
cane passed directly over Dry Tortugas (Tannehill, 1950:186-187).
In his assessment of the damage done on Bird Key, Warden T. J.
Ashe (1919 MS.) wrote: "All vegetation on island destroyed."
Accounts of visits to Bird Key after 1919 (Bartsch, 1923, 1931,
1932; England, 1928) trace the rapid erosion of the denuded island.
The later stages are indicated in the following comments by Charles
I. Park (in litt.): "When I went there in 1929, Bird Key had already

Vol. 8


started to wash away. The house which the former warden had oc-
cupied was considered unsafe so I lived on Garden Key and com-
muted by boat to the other keys. ... Each year erosion on Bird
Key progressed until in 1934 there was very little of the island above
water level." As of June 1935, Longstreet (1936a: 37) stated: "the
remains of Bird Key [are] now eroded to a negligible sandbar."

FicURn 2. Aerial view looking west, Dry Tortugas, January 1945. Long Key
in foreground, Bush Key and Garden Key next rear, and Loggerhead Key in
background. The white spot on the shoal to the left of Garden Key and slightly
above it is a sand bar at the former location of Bird Key. (Official photograph,
U. S. Navy.)

The 1935 Labor Day hurricane was a storm of extreme intensity
but small diameter that struck the central Florida Keys (Tannehill,
1950). In reply to questions about this storm and the one of 4-6
November, which was the only other hurricane in the area in 1935.
Gordon E. Dunn of the United States Weather Bureau. Miami. wrote
me (in lift.): "Neither of these storms passed very close to Dry Tor-
tugas or to Bird Key, and it is doubtful that either of these storms
should have primary responsibility for the disappearance of Bird Key.
I would expect that their effect on Bird Key would have been rela-
tively minor." After storms in January 1940 (Felton, 1940 MS.), a
40-foot sandbar elevated 2 feet above high water emerged at the
former location of Bird Key. Other intermittent reappearances have
occurred more recently (figure 2).

Bush Key, where most of the Sooty Terns and Brown Noddies have
nested in recent years, has an involved history complicated by con-
fusion of names. The names Bush Key and Long Key have been


applied at various times to each of the two adjacent islands on the
shoal east of Fort Jefferson (table 1, figures 2 and 3). The names in
current use were established with the first edition of Chart 585
(1922), but the confusion persisted somewhat longer (viz., Coast
Pilot, 1936: 78).

FIGURE 3. Aerial view looking east. Garden Key and Fort Jefferson in fore-
ground. Bush Key, the east spit, and the northernmost sand ridge of Long Key
at upper right. Large trees in the parade of Fort Jefferson are mainly button-
woods (Conocarpus erectus), possibly remnants of the original stand. Pilings at
the north and south extremities of Garden Key formerly supported the coaling
docks. The center of Bush Key is a thicket of bay cedar (Suriana maritime)
enclosing several mangrove-fringed ponds. Brown Noddies nest at the edges of
this area. The Sooty Tern colony occupies open areas between the bay cedars
and the shore. (Official photograph, U. S. Navy, by U. S. Naval Air Station,
Key West, 1959.)

Now the second largest of the Tortugan islets, Bush Key has under-
gone several cycles of building and erosion. Gauld's chart shows
no land in the area. The 1829 survey reported (as "Long Key") an
island with an area of "5 acres 3 roods 22 poles" and an elevation of
"2 feet 4 inches" (Arana, in litt.). By 1832 this island (or possibly Long

Vol. 8


Key) was thickly covered with bushes and low trees, and Audubon
(1835) referred to it as "Noddy Key" because most of the population
of Brown Noddies was nesting there. Maps of 1853-54 (Bache, 1858)
show a sizable island at the present location of Bush Key. During
the military occupation of Fort Jefferson in the 1860's, the island
served as a pasture and slaughter grounds for cattle and hogs brought
in as food for the garrison (Holder, 1868: 262; Manucy, 1943: 321).
Shortly after this time, Bush Key and Long Key are said to have been,
"almost entirely obliterated by a hurricane" (Holder, 1892: 77). About
1889 Bush Key was a barren sandbank (Coast Pilot, 1889: 40) and
Chart 471a of 1896 shows only a small area above high water. Scott's
(1890) detailed account of Tortugan geography as of the spring of
1890 mentions no land at this location.
The history of Bush Key after 1900 is thoroughly bedevilled by
confusion of names. Significant observations on Tortugan geography
in this period were made by Lansing in 1904 (Millspaugh, 1907), Bow-
man in 1915-16 (Bowman, 1918), and Bartsch in 1917 (Bartsch, 1919).
Millspaugh does not mention Bush Key. Bowman (1918: 128-129)
describes a large, irregular island that had shrubs about 12 years old.
Bartsch (1919:469, 482) refers to Bush Key as "an elevated coral
reef" with the statement "all the vegetation, in fact, most everything
shiftable above the sea, has long since been swept away by the
These records appear to show that Bush Key did not exist in
1904, built up rapidly until 1915-16, and then was suddenly reduced
and dcvegetated (presumably by storms) to produce the conditions
Bartsch found in 1917. Davis (1942: 187-189) and Sprunt (1948b:
5-6) adopt approximately this interpretation of its history. Davis
also points out that Lansing may have overlooked a small island in
1904 because a considerable quantity of sand was removed from
the area in 1901-05 for use as fill during the construction of coaling
sheds and piers on Garden Key (figure 3).
The record is open to the alternative interpretation that Bush
Key had a history of steady growth from before 1900. Close ex-
amination of the accounts of Millspaugh (1907) and Bartsch (1919)
strongly suggest that these authors, following the nomenclature of
the charts then current, referred to the present Bush Key as Long
Key and vice versa. Bartsch (1919: 469), for example, wrote of Long
Key: "the northern end consists of a barren rim of coral boulders
that curves eastward and southward, to join with the reef fringe of
Bush Key." This is a fairly accurate description of present geogra-
phy with the names of the keys reversed. Bowman (1918) discussed



Bush and Long Keys together but appears to have followed present
usage in his application of the names. "Long Key" is described in
Millspaugh's (1907: 225) account as, "so low as to be awash during
heavy weather" and "void of vegetation." Bartsch (1919) indicated
that the southern part of "Long Key" supported a sparse vegetation
of grass and bushes in 1917.
The probable history of Bush Key may be summarized as follows.
After having existed as a well-vegetated island for 40 or 50 years,
it was destroyed by a hurricane around 1870. Sandbars soon re-
appeared at the site, but as late as 1904 they were small and had
no permanent vegetation. During the next decade some plants be-
came established and a series of ridges and bars developed as shown
in Bowman's (1918) sketch. By 1915 (Bowman, 1918) or 1919 (Davis,
1942) several of the sandbars had grown together to cut off ponds
from the ocean. Most of the area between the coalesced bars grad-
ually filled and a long sandspit built up from the east end to give
the key approximately its present shape (Davis, 1942; figure 3). Bush
Key continued to build up during the 1930's and 1940's and contained
an estimated 110,000 square yards in 1946 (Sprunt, 1946b: 5). More
recently some of the shores have eroded, but the island seems to be
more or less stabilized at about 20 acres.

East Key appears on all maps of the area and, unique among the
present islands of the Dry Tortugas, it has borne the same name
throughout its history. Although more stable than many Tortugan
islets, East Key has undergone substantial changes in size and vegeta-
tion. Gauld's chart shows it as the second largest island of the
group. This is corroborated by the 1829 survey which recorded an
elevation of more than 4 feet and an area of about 12 acres, second
in size only to Loggerhead Key. During the late 1800's and early
1900's, East Key may have suffered several periods of devegetation
and erosion. About 1860 (Holder, 1892) it was covered with a dense
stand of bay cedar bushes and numerous mangroves. In 1875 it was
reported to be "partly covered with a growth of cedar" (Coast Sur-
vey, 1878) and later (Coast Pilot, 1889) was said to have "a few bushes
on it." At almost the same time, Scott (1890: 302) wrote of East Key:
"It is a low, sandy, coral island, covered in parts with stunted bushes,
and contains an area of perhaps eighteen acres." By 1904 little but
herbaceous growth persisted and Millspaugh (1907: 224-225) described
East Key as "little more than a mere sand bank 280 x 50 feet in area.
He may, however, have been misinformed about its size. In 1915-16
(Bowman, 1918: 131-132) the island was said to be "almost entirely

Vol. 8


covered with vegetation" including 'large, well-grown bushes," but
no bay cedar. Its dimensions were given as about one-third mile
long and less than one-sixth mile wide. Davis (1942: 191) found a
thicket of bay cedar on the highest sand ridge and reported the
island's dimensions to be about 1200 x 600 feet. He stated that East
Key "has probably grown in size and become more stabilized in the
past half-century." Sprunt (1948b: 17) wrote of East Key: "It com-
prises about 85,000 square yards," indicating continued growth. At
present sizable bushes of bay cedar, sea lavender (Tournefortia
gnaphalodes), and Scaevola plumieri are well distributed over East
It has often been said that Sooty Terns and Brown Noddies were
not known to have nested on East Key and several authors have re-
marked upon the failure of the terns to use so suitable a nesting area.
These comments overlook various records of the 19th century. Large
breeding colonies of both species occupied East Key in the 1850's
(Wurdemann, 1861; Bryant, 1859a). Sooties, at least, still nested
there as late as 1890 (Scott, 1890). Continual persecution by eggers,
mentioned by every early writer, may finally have driven the terns
from East Key. Though a warden was in residence at Bird Key
each nesting season from 1903 on, his surveillance is not likely to
have extended to the outlying islands. It is of interest that no terns
have bred on East Key during the past 28 years of strict protection.

Garden Key adjoins the best protected anchorage in Tortugan waters
and has long been the center of human activity in the area (Manucy,
1943). Most of the key is occupied by the immense ruin of Fort Jeffer-
son (figures 2 and 3). A lighthouse was built on Garden Key as early
as 1825. Construction of the fort began in December 1846 and was
discontinued about 20 years later with the work still far from com-
plete. After use chiefly as a military prison, the post was abandoned
in the 1870's. It was reoccupied during and after the Spanish-Ameri-
can War and World War I, first as a coaling station, later as a sea-
plane base and wireless station.
Gauld's chart shows Garden Key with an irregular shoreline and
the 1829 survey reported its area as about 7% acres. An interesting
map in the files of the U.S. Corps of Engineers (Bache, 1845 Ms.)
is a detailed topographic survey of Garden Key as it was immediately
before the construction of Fort Jefferson began. The shape is rough-
ly elliptical, highest land elevations are just over 5 feet above mean
low water, and the center of the island is shown as low and evidently
swampy. The exact scale of the map is uncertain. Calculations (by


William M. Alexander, Assistant Park Engineer, Everglades National
Park) based upon the scale taken from a superimposed outline draw-
ing of the Fort, laid out on the original map presumably by Major
Bache, give a land area of 8.8 acres above high tide line. The size
of Garden Key was increased by filling when the Fort was being built
and again about 1900 when the coaling structures were built. Davis
(1942: 185) gave the area as 16 acres, of which 5 acres lay outside the
walls of the Fort.
No terns are known to have nested on Garden Key until relatively
recent years. Detail shown on the 1845 map suggests that the in-
terior of the island may originally have been too heavily vegetated
to attract nesting Sooties, although much of it was apparently suit-
able for Brown Noddies. Any that may have nested there undoubt-
edly were displaced soon after 1845. A few pairs of Brown Noddies
have nested on pilings and in the ruins of the north coaling dock in
a number of years since at least 1932 (Bartsch, 1932). In 1937 (Long-
street, 1937), 1938 (Beard, 1938), and 1947 (Sprunt, 1947a) large num-
bers of Sooty Terns nested along the east side of Garden Key. A
substantial part of the Brown Noddy population also nested there in
1937 and 1938, but not in 1947 (Sprunt, 1948a).

Hospital Key, although always a small, shifting sandbar with little
vegetation, has existed since the earliest surveys of the Dry Tor-
tugas. The present name, which was used as early as 1875 (Coast
Survey, 1878) stems from the isolation hospital for yellow fever
patients built there in the 1860's. Sand Key, an earlier name, re-
mained in common use until the 1940's. Various plants have been
recorded from Hospital Key, but the island is so often awash in
rough weather that no permanent plant cover has become estab-
Least Terns nested on Hospital Key in 1907 (Watson, 1907) and
1937 (C. R. Mason, in litt.) and a colony of Roseate Terns has occu-
pied the key in a number of recent years since 1937 (Mason, in litt.).
Sprunt (1948b: 17) suggested that Sooty Terns might find Hospital
Key a suitable nesting area, a prediction fulfilled when a few Sooties
nested there in 1957 and 1959.

Loggerhead Key is the largest, highest, and most heavily vegetated
of the Tortugas and the site of the 150-foot Loggerhead Light (fig-
ure 4) built in 1856-60. The size and shape of the key have been
remarkably constant. It had an area of about 30 acres in 1829 and
is approximately the same size at present, erosion of the west shore

Vol. 8


having been balanced by the growth of sandspits at the northeast
and southwest ends. Loggerhead Key has been credited with an
elevation of 9 feet above mean tide (Millspaugh, 1907: 235; Davis,
1942: 179) but it seems likely that this estimate is excessive. The 1829
survey gave the elevation as "4 feet 4 inches."
Least Terns nested on the Loggerhead Key sandspits intermit-
tently from before 1900 to 1936 (Russell. 1938 mrs.: 4). No other tcrn
is known to have nested on the island.

FIGcnE 4. Loggerhead Key about 1915, looking southwest from the north tip.
In the foreground is the former site of the Tortugas Laboratory, Carnegie Insti-
tution of Washington. In the center of the island, Loggerhead Light. (Official
photograph, U. S. Navy, by U. S. Naval Air Station. Key West.)

Long Key is a bar or shoal of reef debris with several dune-like ele-
vations of broken coral (figure 2). Davis (1942: 189) estimated that
more than one-third of the key was flooded by normal high tides and
that the sparse vegetation of herbaceous halophytes and scattered
small mangroves covered less than one-third of the area above high
tide. If allowance is made for apparent confusion of names in the
past (see Bush Key), it appears that Long Key has never been greatly
different. Gauld's chart of 1773-75 which shows a small island at
the north end of the bar and below it the notation "Ridge of rocks
almost dry and very steep", closely approximates present conditions.


A few Least Terns occasionally nested on the higher sandbanks
at the north end of Long Key as late as 1948 (Sprunt, 1948c). Roseate
Terns have nested there from time to time, most recently in 1962
(Robertson, 1962). Some Sooties and Brown Noddies probably nested
there in 1932 and 1933 (Bartsch, 1932, 1933). In 1943 (Budlong, 1944
Ms.), 1952 (Moore and Dilley, 1953), 1956 (Robertson, 1956 rs.), and
perhaps in other years, many Sooties have tried to nest in rocky spots
between the dunes and farther south on Long Key, but because even
moderate storm waves wash over this section, the attempts are be-
lieved to have been largely unproductive.

Middle Key is shown on Gauld's chart as a fair-sized island, and the
map symbols indicate that it supported some vegetation at that time.
In 1875 (Coast Survey, 1878) Middle Key was still considerably larger
than it is now but without established vegetation. More recently the
key has existed only intermittently as a low strip of bare sand with
few or no plants.
Several pairs of Least Terns may have nested on Middle Key in
1947 (Sprunt, 1948a), and a small colony of Roseates nested there in
1953 (DeWeese, 1953 Mis.), and possibly also in 1960. Gauld's name
for the island, "Bird Kay," suggests that it was once a more important
nesting locality.

North Key, Northeast Key, and Southwest Key all were barren sand
islands that had washed away by 1875 (Coast Survey, 1878). They
have shown no tendency to reappear, but the former location of
Southwest Key is marked on present charts as bare at low water. No
plants are recorded from any of these keys and no terns are known
to have nested on Southwest Key, which may never have been much
more than a high place in the reef. Northeast Key harbored a large
colony of Royal and Sandwich Terns in the late 1850's (Bryant,
1859a). The only definite reference to nesting on North Key seems
to be Holder's (1892: 155) mention of "a solitary gull's egg" (from
the context possibly a Sooty Tern egg) found on the bare summit of
a sand ridge. In addition, Bartsch (1919: 492-493) believed that the
island-about 8 miles northeast of Tortugas Lighthouse--"a small
sand-bar a few acres in extent, called Booby Island"-where Audu-
bon found large numbers of some species of Booby, was probably
North Key.

Vol. 8


Appearance and behavior combine to make the Sooty Tern a con-
spicuous bird, and it has usually been the most abundant species in
the Tortugas terneries. Little wonder, then, that the crowded and
noisy breeding colonies of Sooties have claimed most of the attention
of observers who visited the Dry Tortugas.
Perhaps inevitably, much of the comment on the Sooty Tern at
the Dry Tortugas has centered on the question, how many? Early
ornithologists contented themselves with word pictures that suggest
merely large numbers of birds, but few 20th century authors have
failed to attempt a numerical reckoning of the size of the colony.
Their figures range in quality from guesses made after brief observa-
tion to estimates calculated from measurements of colony area and
density of nests. Table 2 shows what I consider the soundest figures
available for numbers of adult Sooties in each year of record from
1903 to 1956. Population figures for several of the years have had
an eventful history in the hands of compilers, and quantitative data
were found for a number of years previously thought to be gaps in
the record. With these corrections and additions the broad outline
of the history of the colony seems clear, though many details remain
The Dry Tortugas ternery has been called "The Oldest Bird Col-
ony" (Peterson, 1950) on the assumption that its known history reach-
es back to the discovery of the area in 1513. It is reasonable to sup-
pose that the "other birds" of Herrrera's statement (Davis, 1935),
"... there were killed many pelicans and other birds that amounted
to five thousand ", included Sooty Terns. The accounts of other
early visitors, such as John Iawkins (Longstreet, 1936a), and much
later ones, such as George Gauld (1796), contain similar imprecise
allusions to the abundance of sea-fowl at the Dry Tortugas. No cer-
tain record of any tern is known for the area, however, prior to Audu-
bon's visit in May 1832.

1832. Audubon (1835: 263-269) reported Sooties breeding in great
numbers on Bird Key and Noddies breeding on Bush Key. His
account shows that both colonies were then being heavily exploited
as a source of food. Besides several references to the killing of adult
birds and the gathering of eggs it includes the following:
"At Bird Key we found a party of Spanish Eggers from Havannah.
They had already laid in a cargo of about eight tons of the eggs of




Year of

1903 5000
1907 18,858
1909 40,000
1911 48,000
1912 48,000
1913 30,000
1914 97,500
1915 102,000
1916 60,000
1917 80,000
1918 100,000
1919 110,000
1929 80,000
1935 30,000
1936 40,000
1937 72.000
1938 64,057

1939 70,000
1940 100.000
1941 100.000
1942 65.000
1943 100,000
1944 130,000
1945 109.000
1946 97.200
1947 64,270

1948 104,000
1949 120,220
1950 190,876
1951 167,770
1952 76.326
1953 84,569
1954 88,776
1955 71,102
1956 90,452



Area x Density
Area x Density
Area x Density
Direct Count
Area x Density
Area x Density
Area x Density
Direct Count
Area x Density
Area x Density
Area x Density
Area x Density
Area x Density
Area x Density
Area x Density
Area x Density
Area x Density
Direct Count


Burton (in Dutcher, 1904)
Watson (1908)
Peacon (1909 ms.)
Peacon (1911 ms.)
Peacon (1912 ms.)
Pcacon (1913 ms.)
Peacon (1914 ms.)
Ashe and Bethel (1915 ms.)
Bethel (1916 ms.)
Ashe (in Pearson, 1917)
Ashe (in Pearson, 1918)
Ashe (1919 ms.)
Park (ins. notes)
Mason (1936)
Doe and Russell (1936)
Longstreet (1937)
Beard (1938)

Ruloinson (1939)
Robinson (1940)
Peterson (1950)
Budlong (in Vinten, 1943)
Budlong (1943 ms.)
Vinten (in litt.)
Sprunt (1946a)
Sprunt (19461b)
Sprunt (1947a)

Sprunt (1948c)
Dilley (1950)
Moore and Dilley (1(
Moore and Dilley (1(
Moore and Dilley (1i
Moore (1954 ms.)
Moore (1954 ins.)
Moore (1955 ms.)
Robertson (1956 ms.)

Vol. 8


this Tern and the Noddy. On asking them how many they supposed
they had, they answered that they never counted them, even while
selling them, but disposed of them at seventy-five cents per gallon;
and that one turn to market sometimes produced upwards of two
hundred dollars, while it took only a week to sail backwards and
forwards and collect their cargo. Some eggers, who now and then
come from Key West, sell their eggs at twelve and a half cents the
dozen; but wherever these eggs are carried, they must soon be dis-
posed of and eaten, for they become putrid in a few weeks."
Sprunt (1948b:8) points out that Audubon's account contains
nothing definite about the number of terns. Despite this, later writers
almost without exception have supposed that Audubon found Sooty
Terns in far greater numbers than were ever seen again at the Dry
Tortugas. The statement, "both species were on their respective
breeding-grounds by millions," has been cited both as evidence of
former abundance and as typical of Audubon's bent for extravagant
language. Although attributed to Audubon, in fact it is only re-
ported by him as the remark of an officer of the Marion, made as the
ship approached Dry Tortugas and before Audubon had seen the
tern colony.
Peterson (1950: 318) used one of the statistics of the Cuban egg
trade cited above to obtain an estimate of the number of terns in the
colony in 1832. He wrote: "A sooty's egg weighs about thirty grams,
or about fifteen eggs to the pound. Eight tons would come to about
240,000 eggs. As sootics and noddies normally lay but one egg this
shows irrefutably that the concentration was far larger than it is
now." A repetition of the exercise (Peterson and Fisher, 1955: 142)
arrived at an estimate of about 250,000.
Had Audubon mentioned no other statistics, this ingenious reason-
ing might indeed be difficult to dispute. The eggers who spoke of
an eight-ton cargo, however, also told Audubon that they sometimes
realized "upwards of two hundred dollars" per trip to market. If
this is interpreted to have been as much as $250, the 250,000-egg
cargo was sold at ten for a penny. This seems too good a bargain
in eggs even for 1832, especially as the price in Key West is given as
"twelve and a half cents the dozen."
The Sooty Tern population can also be estimated on the basis of
a return of $250 per successful trip and the stated Havana price of
"seventy-five cents per gallon," if the latter is taken to mean fluid egg
contents. Worth (1940: 56) calculated the volume of a Sooty Tern
egg as 1.95 cubic inches or about 118 eggs to the standard gallon.


At 75 cents per gallon a $250 cargo would amount to about 334 gal-
lons, therefore equalling about 39,412 eggs.
Some or all of Audubon's statistical information about egging evi-
dently is inexact. Attempts to derive a population estimate from
any of the details he gives seem unwarranted.
The question of the size of the colony in 1832 can be approached
by considering the number of nests Bird Key could accommodate.
Although Sooty Terns nest in dense aggregations, a limit of colony
compressibility exists. Watson (1908: 200) wrote of Sooties nesting
on Bird Key in 1907: "Each pair defended a circular territory
roughly 14 inches to two feet in diameter." If the smaller figure is
taken to represent maximum density of nesting observed by Watson,
then the minimum area of the territory of a nesting Sooty was 154
square inches and the maximum density of nesting about 8.4 nests
per square yard.
Detailed observations of the density of nesting of Sooty Terns on
Bush Key were made in 1953-56 on 20 to 30 plots each of 8 square
yards distributed throughout the parts of the island judged suitable
for nesting. The largest number of eggs laid on a plot was 56 (7.00
square yard) on one plot in 1954. Field maps show four instances
in which 10 eggs occurred in areas of one square yard within the
larger plots. In each case, however, some were located at the edges,
and no one square yard area appears to contain 10 entire territories.
The average number of eggs per square yard for all occupied plots
and the number of plots that contained one or more eggs were:
1953--3.00 per square yard (14 plots), 1954-3.12 (21), 1955---2.1
(26) (Moore 1954 ms., 1955 ms.); and 1956-2.53 (20) (Robertson, 1956
Measured nesting densities reported for other Sooty Tern colonies
are mostly similar to or lower than those found on Bush Key. Data
for two breeding seasons on Ascension Island (Ashmole, 1963a: 309),
for example, show maximum densities (on plots of 25 square yards
area) of 5.28 and 5.00 eggs per square yard; average densities for all
plots occupied of 1.95 and 2.00 eggs per square yard.
The 1953-56 data from Bush Key suggest that 10 nests per square
yard is about the maximum density breeding Sooties will tolerate.
Few colonies are this crowded, except locally, because vegetation
or terrain limit the number of acceptable nest sites. Nesting Sooties
ordinarily avoid areas with dense shrubbery or heavy herbaceous
ground cover. Ashmole (1963a) found that nests also were fewer
on featureless bare ground deficient in the local clues that enable

Vol. 8


a bird to return to the proper egg. The Tortugas ternery, however,
lacks extensive bare areas.
From the Tatnall-Gednery survey Bird Key is known to have had
an area of about 41/2 acres (21,780 square yards) in 1829. Assuming
for the moment that the Sooty Tern colony occupied its entire sur-
face, an average density of 11.5 nests per square yard would be nec-
essary to accommodate 250,000 nests. Parts of Bird Key, however,
were thickly covered with bay cedar bushes in the 19th century.
Photographs taken much later, after the hurricane of 1910 had great-
ly reduced the amount of plant cover (e.g., Bartsch, 1919: Plate 13),
show large areas still not available to nesting Sooties because of the
dense bush growth. Therefore, I think it unlikely that the maximum
breeding population of Sooty Terns on Bird Key much exceeded
50,000 pairs.
Audubon's manner of reference to his visit to Bird Key suggests
that he saw tremendous numbers of Sooty Terns. A large subjective
element, however, seemingly must be allowed in verbal descriptions
of first visits to Sooty Tern colonies. When Herbert K. Job saw the
Bird Key ternery at its lowest ebb in 1903 he wrote (1905: 87) of the
Sooties: "There are such clouds of them that accurately to estimate
their numbers was impossible This language also could be
taken to indicate great abundance were it not for the rest of the Rev-
erend Job's sentence which reads: but my guess of six or eight
thousand I think cannot be far out of the way."
It seems characteristic of moderns to suppose that Audubon saw
all bird concentrations in their pristine glory. However true this may
have been of many places he visited, it does not apply in the case
of the Dry Tortugas. Bird Key was adjacent to a fine anchorage,
itself adjacent to a major shipping lane that had been used for more
than three centuries. That there were no accurate charts before
Gauld's survey of the 1770's can scarcely have deterred mariners
from using Tortugas harbor. Audubon was told that the terns had
frequented Tortugas "since the oldest wrecker on that coast can
recollect." It is altogether likely that the ternery was first disturbed
on the day of its discovery, and as often thereafter as ships put in to
Tortugas in appropriate season. The most that can be assumed is
that exploitation up to about 1832 had been infrequent enough to
permit the Sooties to rear young in most years.

1840-1902. Although much of this period was marked by intensive
human activity at the Dry Tortugas, the record of the tern colonies
in the 19th century after Audubon's visit is limited to observations


by Bryant (1859a), Wurdemann (1861), Maynard (1881), and Scott
(1890). Comments by the first three of these authors are brief. Scott
discusses the Dry Tortugas in greater detail, but most of his in-
formation about terns is hearsay, because the colonies were not
active at the dates of his visit, 19 March to 10 April 1890.
Data accompanying bird specimens from the Dry Tortugas in
several collections show that other ornithologists may have visited the
tern colonies during this period, but left little or no published record
of their observations. One such visit was by A. L. Heermann and
John Krider, probably in May 1848. Howell (1932: 13) mentions this
expedition but does not include the Dry Tortugas among the places
visited. Heermann (1853: 34), however, lists eggs of the Sooty Tern
and Brown Noddy from "Tortugas Islands" presented by him to the
zoological collection of the Academy of Natural Sciences of Philadel-
phia. At least a part of this material (ANSP Catalogue Nos. 32055,
32060, and 32061) is still in the collection (Henry M. Stevenson, in
litt.) The specimens bear no date, but Ileermann is known to have
visited Florida only once. In the book in which John Krider sum-
marized his career, including "only those species of birds of the
United States that I have myself collected and mounted", he refers
to the Sooty Tern as follows (1879: 81) : "Common on the Keys of
Florida and the Tortugas, where it breeds in large numbers. I have
two specimens in my collection."
The two main items to be gleaned from the later 19th century
papers are: Sooties and Noddies then nested on East Key as well
as Bird Key, and the colonies were under increasing pressure from
Dr. Henry Bryant traveled and collected extensively in Florida
in the decade 1850-1860, but his obituary in the Annual Report of the
Boston Society of Natural History for 1867 gives no details of his
work in Florida, and little seems to be recorded elsewhere. All that
has been known of his visit to Dry Tortugas is that he was there on
8 May. Data on bird specimens he collected now in the Museum of
Comparative Zoology (David 0. Hill, in litl.) suggest that the year
was 1850. Two Sooty Terns (MCZ Catalogue Nos. 42097 and 42099)
carry the dates 10 May and 11 May, respectively, with no year; a
Great White Heron (MCZ No. 42534) he collected at Sand Key off
Key \est, however, places Bryant near the Tortugas on 16 April
1850. His visit there can have occurred no later than the 1853 nest-
ing season, because on 19 April 1854 he donated his collection of
birds' eggs from Florida, including eggs of the Sooty and Noddy to
the Boston Society of Natural History. His own account of his visit


(1859a: 19-21) states merely that he found Sooties and Noddies nest-
ing principally on East Key and "in as great numbers as at the time of
Audubon," and that Royal and Sandwich Terns were breeding "in
great numbers" on Northeast Key. In a paper on Bahaman birds
(1859b: 134) he remarks that Sooties and Noddies occur there "in
immense numbers, as at the Tortugas."
Gustavus Wurdemann (1861: 426) described his visit to the Tor-
tugas the last week of June 1857 in a letter that accompanied a ship-
ment of bird specimens to the Smithsonian, published two years after
his death in 1859. "At the Tortugas are two keys or islands, East
Key and Bird Key, which serve as places of resort to the noddies and
laying gulls to deposit their eggs and raise their young. They are
watched closely at East Key by boatmen, who gather the eggs to
carry them to Key West for sale. But at Bird Key the birds are
under special protection of Captain D. P. Woodbury, the officer
in charge of construction of the fortifications .... The keys are
covered with Bay cedar bushes seven or eight feet in height, inter-
spersed here and there with the cactus, among which some young
laying gulls sought refuge. Their eggs are laid on the sand, whilst
the noddies lay in nests built from two to six feet from the ground
of dried sticks or twigs. Only one egg was found in each noddie's
nest, and about two in the laying gull's. Their eggs are said to have
been taken some time previous to our visit, and that they lay usually
two or three. I picked up several female laying gulls with my
hands, and might have caught noddies if I had not been encumbered
with the gun, birds, and eggs. No young noddies were seen at this
time, which was the last week of June. Other specimens in
the National Museum (Deignan, in lift.) and a Sooty Tern in the
Museum of Comparative Zoology he took there 10 June 1858 (Hill,
in lift.) show Wurdemann also visited the Tortugas the next year.
C. J. Maynard never visited the Dry Tortugas in person. As
Howell (1932: 16) notes "In 1874 he worked at Cedar Keys from
January 26 to March 1. From there, in a small yacht he went down
the coast as far as Clearwater, but from that point he was obliged
to return home on account of illness, leaving his assistants to com-
plete the trip, which took them as far as the Tortugas." This may
partly explain the several geographical and historical inaccuracies
in his account (1881: 480): "The Sooty Terns are now only found in
any numbers on the small islands which lie to the southward [sic]
of Key West and which are known as the Dry Tortugas. Here they
breed on Bird Key which is about four miles [sic] from Fort Jeffer-
son, depositing their eggs early in May. The birds are extremely


tame when nesting, insomuch so, that they may be killed with sticks
or even caught with the hand, and they deposit the eggs on the naked
sand. There were thousands of these birds on this little key in 1874,
but as the soldiers of Fort Jefferson had been in the habit of taking
the eggs regularly every other day, but few or no young were raised.
The officer who had command of the fort, prohibited shooting the
birds on the island, but the continual robbing of the eggs must ulti-
mately drive the Sooty Terns from this breeding ground."
The actual date of Maynard's assistants' visit is indicated by a
Sooty Tern in the Museum of Comparative Zoology (No. 204310,
Hill, in litt.) taken there 25 May 1874, though no collector is named.
The account contains no first-hand comment on the Noddy, nor does
it mention East Key, which lay approximately 4 miles from Fort
Jefferson. Regarding the comments on egging by soldiers in the
area, the Fort Jefferson garrison had been withdrawn 11 January
1874, leaving only a small detail to guard ordnance stores and a
much reduced construction crew engaged mainly in closing down
the operation (Manucy, 1961 ms.). In addition, Captain Woodbury,
the only commanding officer known to have shown an interest in
protecting the tern colony, had left the Dry Tortugas in 1860 and
died in Key West of yellow fever 15 August 1864 (Cullum, 1891:
496-497; Manucy, 1961 Mis.). It seems likely that the report Maynard
received from his assistants in 1874 blended considerable hearsay
with their actual observations.
Most of W. E. D. Scott's infonnation about nesting terns was
sent to him after his return from the Dry Tortugas by Dr. F. S.
Goodman, who was stationed at the Quarantine Station on Garden
Key. Scott reports (1890: 307) Sooties nesting on East Key and Bird
Key, and Noddies "mainly confined" to Bird Key, but his comments
on egging are of greatest interest: "All of the Gulls and Terns that
breed at the Dry Tortugas have been much diminished in numbers
in the past ten years. It has always been the custom for some of
the boats engaged in fishing and sponging about Key West to resort
to these islands during the breeding season, and lately their depreda-
tions have really made a very appreciable difference in the birds
that resort to this breeding ground. I am told that the eggs have a
commercial value as an article of food in the markets of Key West,
where barrels of birds' eggs from the Tortugas are brought every
season of late years."
Vinten (1943: 54) suggests that search of the records of govern-
ment agencies that had maintained operations at the Dry Tortugas
might reveal additional data about the tern population during this

Vol. 8


period. Though most of the search of the voluminous Army archives
of Fort Jefferson remains to be accomplished, the studies of historians
show these archives do indeed contain information pertinent to the
history of the ternery. Albert Manucy (in litt.) advises me that among
records he examined he recalls having seen correspondence relating
to the visit of Louis Agassiz to the Dry Tortugas in 1858, and that
the Fort Jefferson Letter Books include such items as a letter from
Mordecai and Co. to Woodbury on 3 May 1859 concerning ship-
ment of Woodbury's bird specimens to the Smithsonian Institution.
The historical records Manucy (1961 MS.) studied suggest Bird
Key suffered even more disturbance than the authors of ornithologi-
cal works on the Dry Tortugas have appreciated. Shortly after war
began in 1861, for instance, concern for the safety of Fort Jefferson,
still unfinished and weakly armed, led to the appropriation early in
1862 of 8200,000 to fortify Bird Key. The preliminary survey, in-
cluding extensive borings to determine subsoil structure, was de-
layed by personnel changes and slow delivery of materials, and was
not completed until the spring of 1864. The project then seems to
have lapsed, but it can hardly have failed to disrupt the terns attempt-
ing to nest during the survey.
Manucy also cites a letter of 18 July 1865 to the Post Commandant
from Edward Frost, Assistant Engineer in Charge, complaining of
the removal of a number of hogs "from their ranging ground on Long
Key to Bird Key" which contained "the scattered graves of many
Union Soldiers who have died at this Post during the war." Whether
or not the hogs were returned to Long Key seems to be unrecorded.
Most probably the Sooty Terns failed to rear young at the Dry Tor-
tugas in most of the years from 1860 to the early 1870's when Fort
Jefferson was heavily garrisoned. This loss of annual recruitment
plus an undoubtedly heavy mortality of adults must have reduced
the population rapidly.
Little definite information about the ternery exists for the years
1890-1903. It may be presumed that the colony was raided regularly
by eggers, and that some time in this period Sooty Terns nested for
the last time on East Key. J. W. Atkins, a well-known resident col-
lector of Key West. collected specimens now in the Museum of Com-
parative Zoology at the Dry Tortugas in May 1896, but no other
record of his trip is known. A. G. Mayer visited Bird Key in 1898,
but the only datum published (in Dutcher, 1906) is his impression
that Sooties were then about one-third as numerous as at his next
visit in 1906.


With the outbreak of the Spanish-American War, Fort Jefferson
was garrisoned once more from 1898 until about 1906. In 1900 the
Dry Tortugas were transferred to the Navy Department and con-
struction of a coaling depot at Garden Key began.
Sprunt (1948b: 9) suggests that the renewed military activity at
Fort Jefferson probably put additional pressure on the tern colony.
The Navy at the command level was aware of the need to protect
Bird Key, for a letter from Captain T. C. Treadwell quoted by Dutch-
er (1903: 120-121) states Treadwell ordered egging stopped soon after
he assumed command of the U. S. Naval Station, Key West, in June
1901. Unfortunately orders from Key West were not altogether ef-
fective at the Dry Tortugas, for according to Thompson (1903: 77-78)
the terns "suffered very seriously" from eggers in 1902. Thompson
adds, presumably with reference to the recent past and to both
Sooties and Noddies: "There have been years when not a single
individual was raised, every egg having been taken shortly after it
was laid."
Thanks to William Dutcher's untiring efforts sterner measures to
protect Bird Key followed in 1903. The Secretary of the Navy issued
an order on 24 April prohibiting the taking of eggs or disturbing of
terns at Dry Tortugas, and in May W. R. Burton was detailed there
as a special warden representing the American Ornithologists' Union
with the permission and logistical support of the Navy (Dutcher,
1904). Burton arrived at Bird Key accompanied by H. K. Job 19 May
1903. The modern history of the ternery can fairly be said to begin
on that date.

1903. Four estimates of the Sooty Tern population in 1903 are avail-
able from the published comments of the original observers. They
are: "3600" by Job and Burton made before Job returned to Key
West on 22 May; "at least 5000" by Burton in a letter to Dutcher
dated 15 July 1903, the increase accounted for by birds that began
nesting after Job's departure; "fi\e to six thousand" by Job in a letter
to Dutcher (all three figures published in Dutcher, 1904); and, "six
or eight thousand" (Job, 1905: 87). The context of the accounts sug-
gests that the figures refer to number of adult Sooties rather than
number of nests, but nowhere is this clearly stated. Compilers have
given the 1903 population as 3600 (Longstreet, 1936a; Vinten, 1943;
Sprunt, 1947b; Peterson 1950), 6-8000 (Sprunt, 1948b), and "about
7000 nests" (Fisher and Lockley, 1954: 60; Peterson and Fisher, 1955:
142). The figures, where identified, are in all cases credited to Job.
I consider the warden's figure of 5,000, based upon observation of the

Vol. 8


colony through the entire nesting season, to be the soundest estimate
I have found no record of the condition of the colony in 1904
and no estimates of the population for the seasons of 1904 through
1906. Charles Russell, the warden in 1905, reported "a very success-
ful season" (Dutcher, 1905). After visiting the colony in 1906 A. G.
Mayer informed Dutcher (Dutcher, 1906) that the Sootics appeared
to be three times as numerous as they were in 1898.

1907. John B. Watson began his studies of the tern colony in 1907
and also served as the warden of the National Association of Audubon
Societies for that season. In addition to his other work Watson made
a careful estimate of the nesting population of Sooties. He divided
the colony into 10 sections presumably distinguished by conspicuous
features of vegetation or terrain. By determining the area and sam-
pling the density of distribution of nests within each section, he ar-
rived at an estimated 9,429 nests or 18,858 breeding adults (Watson,
1908: 198).

1908. Most summaries of the changes in size of the Tortugas Sooty
Tern population include an estimate of 20,000 (or 10,000 nests) as the
population in 1908. All authors who cite an authority credit this
figure to Watson who, according to the Carnegie annual reports, was
not at the Dry Tortugas in 1908 or 1909. The earliest reference I
find to it is Lashley's (1915: 61) statement that Sooty Tern nests to-
talled "more than 10,000 in 1908," with no mention of the source
of his information. I have omitted the figure from table 2 because
I can find no authority for it.

1909-1916. On 6 April 1908 Executive Order No. 779 of President
Theodore Roosevelt established the Tortugas Keys Reservation for
protection of birds nesting in the area. The order specified that use
as a bird reservation was not to interfere with military uses (under
President Polk's Executi\e Order of 17 September 1845 establishing
the Dry Tortugas Military Reservation) except that military use of
Bird Key was prohibited. Protection of the Tortugas Keys Reserva-
tion became the responsibility of the Bureau of Biological Survey.
After 1908 warden protection at the Dry Tortugas was supported
jointly by the Biological Survey and the National Association of Au-
dubon Societies. T. J. Ashe of Key West, who was in general charge
of bird protection activities in the Florida Keys during most of the
ensuing decade, hired and supervised the men stationed at Bird Key.


These were John Peacon (1909-1914), Ludwig Bethel (1915-1916), and
William E. Lowe (1917-1919). Warden's reports on the condition
and size of the tern colony were made annually to both supporting
organizations. From the annual reports to the Biological Survey I
have seen only the data entered in the bird distribution file now at
Patuxent Wildlife Research Center of the Bureau of Sport Fisheries
and Wildlife.
The annual reports to the National Association of Audubon So-
cieties for this period are still in the files of that organization. Some
were prepared by Ashe and submitted in his name; others seem to
have been prepared by his wardens at Bird Key. They include esti-
mates of the population of Sooties in all the years 1909 through 1916
except 1910. Watson apparently prepared the 1910 report for Bird-
Lore, but it was not published and has been lost. This is unfortunate
because comments in a later report (Watson, 1912 uM.) indicate he
made the 1910 count of Sooties by the same method he used in 1907.
Previous summaries of the colony include no mention of Sooty
Terns in these years, but skip directly from the questionable 1908
figure to 1917. In addition to the annual warden's reports, several
published comments for this period have been generally overlooked.
Of the population in 1913 Watson and Lashley (1915: 38) wrote:
"There are probably more than 18,000 (possibly 30,000) sooties on
Bird Key." On 28 May 1915 Herbert K. Job and H. R. Mills visited
Bird Key to take motion pictures for the National Association of Au-
dubon Societies. A brief excerpt published from Job's report (Pear-
son, 1915) gives the number of Sooties as "possibly 75,000." Pearson
also prepared a longer article about this trip (1915 MS.), evidently
copy for Bird-Lore that wasn't used, which quotes more extensively
from Job. It reveals that the 75,000 population figure was based on
area-density calculations by Mills. Because these calculations con-
tain obvious inaccuracies impossible to resolve today, I have used
the 1915 population estimate from the warden's report in table 2.

1917. According to Warden T. J. Ashe's annual report (Pearson,
1917: 398) there were probably 80,000 of these birds [Sooty
Terns] nesting on the island." This figure has been overlooked by
compilers, who instead have misquoted the 1917 population of Sooties
from Bartsch as "18,000" (Longstreet, 1936a; Fisher and Lockley,
1954) or "25,000" (Vinten, 1943; Sprunt, 1947b, 1948b). Bartsch's list
of the birds seen at the Dry Tortugas 19-31 July 1917 (1919: 471) in-
cludes under Sooty Tern "adult 118,000 young 27,200." The figures
are keyed to footnotes that read: "'Based upon Doctor Watson's cen-

Vol. 8


sus of 1908.", and "2An estimate admitting two-fifths as many off-
spring as we had parents." Bartsch also (1919: 473) wrote of the
Sooties: probably more than 25,000 are present on Bird Key
at the close of the breeding season." Apparently Bartsch made no
independent estimate of the Sooty Tern population in 1917; the
figure 18,000 is an approximation of Watson's total for 1907 (not 1908)
and the "more than 25,000" is merely 18,000 adults plus Bartsch's
arbitrary figure for young of the year.

1918-1934. None of the earlier compilations mentions these years.
I have seen warden's reports only for 1918 and 1919. A Federal law
effective 1 July 1919 ended the National Association of Audubon
Societies' participation in the protection of Bird Key (Pearson, 1919).
The Biological Survey continued to employ a warden at the Dry Tor-
tugas during the summer at least through 1930, but no wardens' re-
ports can be located in the files now stored at Patuxent Wildlife
Research Center (Robbins, in litt.), and the distribution files contain
only the warden's estimate of the Sooty Tern population in 1929
(Park, Ms. notes).
Several popular articles published in the 1920's refer in passing
to the number of terns at the Dry Tortugas. England (1928: 86) men-
tions a population of 50,000, and a photograph in an article by Long-
ley (1927: 66) is captioned: "The west shore of Bird Key showing
some of the 33,000 birds that breed here annually." These figures
are not considered bona fide population estimates. Neither can be
associated with a definite year, and the 33,000 is suspiciously near
Bartsch's (1919: 471) total of 32,810 for all the birds (19 species) he
identified at the Dry Tortugas in July 1917.
Bartsch visited Bird Key several times during the 1920's and in
August of 1931, 1932, and 1933. Existing records of his trips contain
no reference to the total numbers of Sooty Terns. The brief published
accounts of the later visits (Bartsch, 1931, 1932, 1933) have great in-
terest because Bird Key was then eroding rapidly. In 1932 Bartsch
noted that a few Sooties were nesting on Bush [Long] Key. The
following year he reported (1933: 267) that more than half the popu-
lation had left Bird Key and "It is beginning to look as if the major
portion would eventually establish itself on Long [Bush] Key." C. C.
Von Paulsen of Homestead, Florida, then an officer of the U. S.
Coast Guard, visited the Dry Tortugas frequently in the years 1932-
1934. As he remembers it (personal communication) a substantial
part of the Sooties nested on Bird Key in 1933 and smaller numbers
remained there in 1934.


From the scant information available it appears that the tern
colony may have been without warden protection in the early 1930's.
Charles I. Park, the last Bird Key warden and now a resident of Key
West, wrote me in a letter 14 December 1959: "As well as I can
remember, I served as warden in the Tortugas area from 1929
through the summer of 1934, a total of six years." As G. A. England
(1928: 14) refers to Charles Park as the Bird Key warden during his
visit there the summer of 1926 or 1927, apparently Mr. Park began
his six years of service two or three years earlier than he recollects.
Others who knew the Dry Tortugas in the early 1930's do not recall
a warden in the area during those years (Julius F. Stone, Jr., Charles
M. Brookfield, and C. C. Von Paulsen, personal communications).
Absence of warden protection would explain the apparently well-
founded rumors that the early depression years saw a vigorous re-
newal of egging at the Dry Tortugas. It seems likely that protection
of the ternery at least was less vigilant in the nesting seasons of 1931
through 1934, although Sooties are known to have succeeded in rear-
ing many young in some of these years (Bartsch, 1932).
The National Park Service assumed administrative responsibility
for the Dry Tortugas early in 1935. Mason (1936: 18) mentions that
the Custodian of Fort Jefferson turned away many boat parties from
Key West that came to gather eggs on Bush Key in the spring of
1935. Correspondence in National Park Service files suggests that
the colony was raided late in the 1935 season and a number of young
birds taken. Protection of the colony by the National Park Service
probably was not fully effective until the nesting season of 1936.

From 1935 through 1941 one or two groups of observers visited the
Dry Tortugas each June on trips sponsored by the Florida Audubon
Society. The visits were brief, each group spending from two to
five days at the Tortugas. Adult and young terns were banded in
1937 through 1941, and the published accounts of all the trips, except
that of 1941, include estimates of the number of Sooty Terns. In
1937 and 1938 many Sooties nested along the east side of Garden
Key (figures 5 and 6) as well as on Bush Key.

1935. The population figure in table 2 is an average of estimates by
members of the party (Mason, 1936). Some thought as many as
50,000 Sooties were present.

1936. Doe and Russell (1936: 6-7) state of the published population
estimate: "It was the general opinion of those who had been on the

Vol. 8


trip in 1935 that the tern colony had increased one-third." Mason
(As. notes) entered an estimate of 48,000 in his field notes with the
comment that he considered it "very conservative" because the nest-
ing colony of Sooties covered a much larger area on Bush Key than
it had in 1935.

&.4 26 WO
*7w', 6

V ~ ~ ~ _'

6.*- *

4 -

.4~ *~ -. *

s -
.' 1, 4.

FIGUHR 5. Portion of the Sooty Tern colony on the east side of Garden Key in
1937: (top) June; and, (bottom) August, showing many well-grown juveniles.
(National Park Service photographs by Philip C. Puderer.)



Q. U -dl) tl d
rWln~ ~tL;. at~
~ e
4. ?1! ii
'I; r .. I:

:~ 1
a_ .ic ~~ ~: c


1937. The colony was said to occupy an area of 8000 square yards
on Garden Key and 4000 square yards on Bush Key. From this area
and a nesting density of "about six sooties to the square yard," de-
termined from one sample plot of 9 square yards in "a typical sec-
tion" of the Garden Key colony, Longstreet (1937: 8) calculated a
total of "72,000 [adult Sooties] actually present at one time." Though
72,000 birds present at one time would represent a total of 144,000
breeding adults by the usual methods of reckoning, Longstreet (1937:
8) continued: "It would seem not far wrong to calculate the number
of adult sooty terns at the Tortugas in June 1937, as approximating
100,000. This would be a tremendous increase over any previous
estimates, and for that reason may be seriously in error. But, at any
rate, it is an estimate based on actual count of birds in a given area,
multiplied by the number of times that area is found in the total area
occupied by the birds." All summaries of the history of the Tortugas
Sooty Tern population have cited the 1937 population as 100,000
from this source. Russell (1938 ts.) also "estimated the number of
Sooties to exceed 100,000."
Other observers appear to have considered this estimate too high.
Young and Dickinson (1937) believed that Bush Key had no more
than 20,000 Sooties, and Mason (us. notes) recorded an estimate of
75,000 for the total adult population. Longstreet (1937: 7) includes
a photograph, taken from the terreplein of Fort Jefferson, of the
sample plot on which the figure for density of nesting was based.
The picture shows most of the Sooties are either incubating or brood-
ing small young, and hence distributed one adult per territory. Be-
cause of the angle it is not possible to tell exactly how many Sooties
are on nests within the 9 square yard plot, but the number is 20 to
30, certainly not 54. Thus, Longstreet's figure of 6 birds per square
yard is apparently based on a nesting density of about 3 nests per
square yard with allowance for the absent member of each pair.
Accordingly, 72,000 is considered the soundest estimate of the breed-
ing population of adult Sooties in 1937.

1938. This year Sooty Terns again nested on both Garden (figure 6)
and Bush Keys, but the colony divided more equally between the
two. Considerable effort was devoted to careful measurements of
areas occupied and nest densities on both keys, and the resulting es-
timate (Beard, 1938) is undoubtedly one of the more accurate of
the population figures for adult Sooties in the Tortugas ternery. Di-
rect counts of nests on the -coaling docks (figure 6a) and in small,
irregular patches of dense vegetation on Garden Key totalled 3950.

Vol. 8



-~ a

1: '4 C
Sit. Lb-

FIGUKE 6. Sooty Tern colony on Garden Key in 1938: (top) Sootics nesting on
the north coaling dock; (bottom) another section of the colony early in the season.
Bush Key in the background in both photos. (National Park Service photographs
by Daniel B. Beard.)

. ,. ^i *^



The main open nesting areas occupied 5442 square yards on Garden
Key and 11,097 square yards on Bush Key; a measured sample of
276 square yards on Garden Key yielded an average nesting density
of 1.8 nests per square yard, which was taken as typical for both keys.
Nesting density in the more heavily vegetated parts of the Bush Key
colony was determined as 1.25 nests per square yard. Beard's (1938)
calculations contain a slight error in addition, and the correct total
is 64,057, not 64,058.

1939. The entire colony of Sooties nested on Bush Key this season.
O. B. Taylor (1939 Mrs.) was told by the Custodian that part of the
birds first settled on Garden Key in early May, but soon moved across
to Bush Key. Robinson (1939: 7) thought they abandoned Garden
Key because "most of the cover around the fort had been cut down
prior to the arrival of the terns this season." Though he speaks of
counting birds on "sample areas," Robinson probably arrived at his
population figure by calculating from approximations of the colony
area and nesting density. As it is not certain that any areas were
measured, this and his 1940 figure are considered simple estimates.
Vinten (1943) credits another estimate, also of 70,000 and perhaps
taken from Robinson, to James B. Felton, then Custodian of the fort.
Taylor (1939 1M.) recorded an independent estimate of 65,000 adult
Sooties from his observations later in June 1939.

1940. A sketch of the colony (Robinson, 1940) shows that Sooties
occupied most of Bush Key except the eastern sandspit, as they had in
1939. The accompanying text reads: "At first it did not seem that
there were quite as many sooty terns as last year, but a complete
tour of the key revealed that there were more than we expected.
The same method was employed to estimate the number of birds as
last year, and our figures show that there were 100,000 sooty terns
in the colony." Just how this was calculated he does not say.

1941. The published report of the trip (Rea, Kyle, and Stimson.
1941) included no estimate of the number of Sooties, but R. T. Peter-
son, who accompanied the second of the two parties, wrote (1950:
318): "On our visit in 1941 we hardly dared estimate the number
exactly, but it was well over the 100,000 mark."

1942-1944. Information for these years comes from the official re-
ports of Custodian Robert R. Budlong. As he was unable to spend
much time observing the colony, his comments on numbers and popu-

Vol. 8


lation trends must be viewed as impressions rather than careful esti-
mates. Military aircraft were active in the Dry Tortugas area during
this period. Budlong (1942 xrs.) comments that the tern colony was
frequently disturbed by low-flying planes in 1942. The report of
the A.O.U. Committee on Bird Protection for 1943 (Allen, 1944: 629)
states: "Unauthorized use of Bird Key [sic], Fort Jefferson National
Monument, as a bombing target by unidentified aircraft late in 1942
resulted in a fire that burned all vegetation. This and several less
injurious acts of similar nature have been the subject of protests to
the several military and naval establishments. Fortunately, the fire
occurred outside the nesting season, but the island will not be usable
by the Sooty and Noddy Terns until it is revegetated." The comments
presumably apply to Bush Key. Burning of the vegetation is not
likely to have discouraged Sooty Tern nesting but it may well have
affected the Noddies.
In 1942 Budlong (1942 Ms.) stated the colony had decreased about
one-third and estimated the number of Sooties at 60-70,000, all on
Bush Key. Vinten's (1943) statement of the figure as 65,000 has been
followed. In 1943 Budlong (1943 MS.) considered the population to
have shown a 50 per cent increase to "about 100,000." In 1944 the
Sooties abandoned Long Key, the east spit of Bush Key, and several
large areas on Bush Key proper, all used heavily in 1943, but Bud-
long (1944 MS.) believed there were "as many or more Sooties in the
colony" as in 1943. At the end of the season Vinten (in litt. to Re-
gional Director, U. S. Fish and Wildlife Service, Atlanta, Ga.) com-
mented: "About 130,000 birds nested there during the past summer."

1945-1948. Data for these years are quoted from the reports of Alex-
ander Sprunt, Jr., who made annual trips to the colony in June and
determined the size of the adult Sooty population each year by an
area X density method. While he paid careful attention to the space
the colony occupied, just how he measured the average nesting den-
sities isn't always clear. In 1945 he appears to have used those de-
termined by Beard (1938), about 1.8 nests per square yard in open
areas and 1.25 in more heavily vegetated sections. The other years
he determined separate nesting densities for each section of the col-
ony that appeared to differ materially, but he gives sizes of the areas
sampled and counts of nests in each only for Bush Key in 1946
(1946b: 5).
Sprunt also described the remarkable spread of vegetation on Bush
Key in this period and its effect on the location and density of the
nesting Sooties. He records the space the colony occupied on Bush


Key in 1945 as 34,000 square yards (1946a), in 1946 as 27,200 square
yards (1946b), and in 1947 as about 7,000 square yards (1947a). In
1947 some of the colony nested on Garden Key again as they did in
1937 and 1938. In 1948 the entire colony again located on Bush Key;
the vegetation was still luxuriant, but the Sooties dispersed more
thinly over an area of 52,000 square yards (1948c).

1949-1956. Population estimates for these years were made by per-
sonnel of Everglades National Park. Willard E. Dilley, then Chief
Park Naturalist, worked at the Dry Tortugas in 1949 and 1950, and
he and Joseph C. Moore, then Park Biologist, worked together there
in 1951. Moore continued the annual surveys through 1955. I made
the population estimate of 1956 following procedures established by
Moore. Results of the surveys of 1949 through 1952 have been pub-
lished; data for 1953 through 1956 are from typed reports in the
Everglades National Park files.
All population estimates were obtained by the usual area X den-
sity methods. Those of 1949 through 1951 were based upon separate
determinations of area and density of nesting in a number of sub-
areas where the pattern of occupation by nesting Sooties seemed to
differ noticeably, essentially the same procedure followed by Sprunt,
Beard, and others back to Watson in 1907. The number of sub-areas
distinguished and measured separately was: 1949, 7; 1950, 22; and,
1951, 15. In 1952 Moore established 20 marked plots each of 8
square yards distributed throughout the parts of Bush Key consid-
ered to be available to nesting Sootics. Data on density of nesting
used in calculating the Sooty Tern populations of 1952 and 1953 were
taken from these plots, and data for 1954 through 1956 were taken
from these plots plus 10 additional plots Moore established in 1954.
In 1951, 1952, and 1956 numbers of Sooties nested among rough
coral rubble at low sites on Long Key. Moore and Dilley (Moore,
is. notes) estimated 455 adult Sooties nesting on Long Key in 1951,
and in 1952 Moore (Moore and Dilley, 1953: 76) believed about 2000
present, although few yet had eggs. On 26-27 May 1956, David O.
Karraker, my wife, and I counted 2880 Sooty Tern nests with eggs
in place on Long Key, and saw about 700 scattered eggs from nests
that had been flooded (Robertson, 1956 MS.). All the Long Key nest-
ings were behind the schedule of the main colony and produced few
or no young.
In reporting Sooty Tern observations from a visit to the Dry Tor-
tugas in May 1953, Fisher (in Peterson and Fisher, 1955: 143) com-
mented: "My own estimate of the number of occupied nests-

Vol. 8


80,000--was not far off. A census based on sample plot counts which
was made two weeks later by the Park Service came up with a figure
of 84,569 sooty nests." The figure mentioned was in fact an estimate
of the number of breeding adults (Moore, 1954 mis.); the number of
nests actually amounted to but few more than half Fisher's estimate.

1957-1963. In 1957 Sooty Terns were first recorded nesting on Hos-
pital Key. Mr. and Mrs. John R. DeWeese found about 200 nests
there in June but were not certain that any young were reared (in
litt.). None nested on Hospital Key in 1958 (Richard Ward, in litt.),
but on 15 June 1959 0. L. Austin, Jr., C. R. Mason, and I found about
50 adult Sooties among the colony of Roseate Terns there and located
about 10 Sooty Tern nests with eggs. In the main ternery of Sooties
on Bush Key hatching was at least 90 percent complete at this time
and the larger young were about half-grown. In the nesting seasons
of 1960-1962 no Sooties were observed at Hospital Key, but in July
1963 about 8 adults appeared to be settled there, again associated
with nesting Roseates. No search was made for nests, but the be-
havior of the Sooties suggested that they were nesting. It is of in-
terest that all occurrences of Sooties on Hospital Key were in years
when Roseates also nested there, none having been noted in the
years when the Roseate Terns located elsewhere.
Work at the Tortugas in 1959-1963 consisted chiefly of banding
adult and young Sooty Terns in large numbers, and no direct esti-
mates of the size of the colony were attempted. My impression is
that in 1959-1963 the population was in the range of 70,000 to 100,000
breeding adult Sooties and varied relatively little from year to year.
Approximately 6,500 to 11,000 young Sooties were banded each year
in 1960-1963 and the recorded mortality of eggs and small young
accounted for an additional 2,000 to 6,500 nesting efforts annually.
Counts of living young and of young found dead made each year
after banding was completed have shown consistently that from
one-third to one-quarter of the birds of the year were banded.
Each year since 1960 a sample of from 7,000 to 8,200 adult Sootics
has been captured in mist nets set at the perimeter of the colony
(figure 7). It should be possible to estimate the number of adults
accurately from the proportion of banded individuals occurring in
samples taken later the same breeding season. But calculations from
May-banded adults in samples of adults netted the following July
yield population estimates considered three to five times too high.
Two characteristics of the Tortugas Sooty Tern population, strong
localization of individuals within the colony and straggling arrival


and departure, hamper use of mark-recapture data for estimating
total numbers. Banded adults do not become randomly distributed
throughout the colony, and the sampled population changes in com-
position from week to week during the breeding season.

-. ": 4. A

dh 1

-. q'. '-

V '- 7$ '.-

Flcunr 7. Members of the Florida field excursion, 13th International Ornitho-
logical Congress, mist-netting adult Sooty Terns on the west beach of Bush Key.
In foreground from the left, Josias Cunningham (U.K.) and Staffen Ulfstrand
(Sweden). The matted ground cover is sea purslane (Sesuviurm portulacastrum).
(Photograph by A. Schifferli. 13 June 1962.)

Only a limited interpretation of the record of the population of Sooty
Terns at the Dry Tortugas (table 2) can be undertaken now. The
present discussion aims merely to review the estimates in the light
of the species' behavioral characteristics and the Tortugan environ-
mental factors that may have influenced them over the years. Some
of the local limiting factors and the difficulties of accurate censusing
were recognized and discussed by earlier writers. The preliminary
results of work now in progress include some additional pertinent
information. Comments are limited to the population records from
1903 to date.
The fact that all adults in the colony do not begin to nest at about
the same time has plagued Sooty Tern counters at Dry Tortugas from


the beginning. The varying population reports of 1903 (Dutcher,
1904) resulted in part from the arrival of many Sooties after Job and
Burton had made their first estimate. Watson (1908: 318) specified
that his 1907 count was made "late in the brooding season, after all
the eggs had been laid." Later observations, particularly by Moore
in the early 1950's, make it clear that simply to delay the count until
all the birds arrive is not a satisfactory solution. It may be useful
to review what is known of the arrival and landing of Sooty Terns
at the Dry Tortugas.
A period of nocturnal swarming over the breeding grounds before
actual nesting begins is characteristic of the species (cf. Ashmole,
1963a). The Brown Noddy also exhibits this behavior at the Dry Tor-
tugas, and it may be part of the pre-breeding activity of all pelagic
terns, although observations on species other than the Sooty are few.
The occurrence of Sooty Terns on regular nocturnal visits in late win-
ter was not reported at the Dry Tortugas until rather recently (Vin-
ten, 1944; Baker, 1944). Our knowledge of the phenomenon coincides
closely with the period of Sooty Tern occupancy of Bush Key and
National Park Service occupancy of adjoining Garden Key.
Park Service personnel stationed on Garden Key have kept a
complete record of the dates the Sooties were first heard over the
area at night and first seen to land on Bush Key by day annually since
1943. The two events are the outstanding phenological phenomena
of the, for some, rather humdrum year at the Dry Tortugas, and one
suspects they have often been recorded as welcome evidence of the
passage of time. Dates of the first night occurrence of the Sooties
range from 8 February (1943 and 1956) to 7 March (1950) with an
average date over the 20-year period of 20 February. The average
date of the first daytime landing is almost exactly 2 months later,
21 April.
Typically the number of birds, as judged by the amount of noise,
begins to increase nightly soon after the first report, but for a month
to six weeks no Sooties are to be seen in the vicinity by day. Their
daytime whereabouts during this period is regarded locally as some-
thing of a mystery, but my records of 24-26 February 1964 suggest
that the birds frequent the Gulf of Mexico probably at no great dis-
tance. I heard Sooties calling as early as 2030 hours (all times EST)
and as late as 0615 hours, both 25 February. They seemed to ap-
proach from the northwest and the last birds over Garden Key near
dawn seemed to depart in that direction.
Activity on the night of 25-26 February 1964 centered about one
mile north-northwest of Bush Key. Observation from a boat in this


area disclosed several separate flocks, each apparently of many thou-
sands, milling about in rapid flight within a few feet of the surface
of the water. No appreciable movement toward the colony site on
Bush Key occurred from 2100 to 0130 hours. At least a month before
the first daytime landing, however, Sooties have been reported land-
ing on Bush Key at night, and occasional precocious females lay eggs
during nocturnal visits as much as three weeks before nesting begins
(Robert and Stevenson, 1951). The night landings presumably cor-
respond to the gatherings on Ascension termed "night clubs" by Ash-
mole (1963a: 301 ff.), but no detailed observations are available from
the Tortugas.
A few days to a fortnight before the definitive landing a few Sooties
often remain on Bush Key well past daybreak, and scattered birds
and occasional large flocks are seen at sea nearby. Soon after these
first daytime sightings Sooties either land at night and remain on
the island or begin lauding by day, usually in early morning. The
number that land the first day reportedly varies from a few hundred
to many thousands. Elms (personal communication) estimated 40,000
the first day in 1963. Laying begins at once. Commonly hundreds
of birds are incubating by the afternoon of the day of landing.
New flocks arrive nightly for at least several weeks. Moore and
Dilley (1953) noted that the spread of hatching dates indicated the
period of arrival was greatly more prolonged in some years. The
larger the colony becomes, the more difficult are new arrivals to
detect, except as they occupy entirely new ground.
In the usual pattern of landing, successive flocks settle immedi-
ately contiguous to the ground already occupied. Almost invariably
the first Sooties land on the west side of Bush Key. From this nucleus
the colony builds eastward along both shores, the last birds to come
in landing on the east spit or (occasionally) Long Key. Felton (1941
MS.) suggested that thus the Sooties first settle on the oldest part of
Bush Key, an intriguing idea impossible to verify. In at least two of
the years when Sooties nested on Garden Key (Beard, 1938; Gibbs,
1947 Nis.) the first birds landed on the north coaling docks (figure 6).
In 1938 new flocks built southward from that point until all of Garden
Key east of the fort was occupied before any landed on the west side
of Bush Key.
Over the past 20 seasons first landings of Sooties at the Dry Tortu-
gas have become consistently earlier. The average date of first land-
ings for 1943-52 was 27 April, for 1953-62 it was 14 April. The land-
ings of 7 April 1961, 6 April 1962, 3 April 1963. and 28 March
1964 are the earliest of reliable record. No similar trend can be seen

Vol. 8


in the record of first nocturnal visits. These are less likely to be re-
corded accurately, because they apparently are brief high overflights
by a few birds in the middle of the night On the night of 19-20
January 1964, about 3 weeks before the then earliest report, I heard
Sooties calling over Garden Key four times between 2345 and 0200
hours. Each time one or two birds flew over rapidly and fairly high,
the passage marked by three to five unmistakable "wideawake" calls.
The following night it was colder with high winds, and I listened from
2200 to 0200 hours without hearing any Sooties. On my next visit in
late February 1964 large numbers were visiting the area nightly.
Brief observations of Sooties as they first landed in April and May
1963 revealed several interesting characteristics of appearance and
behavior in the newly-arrived birds. Despite the fact that they pre-
sumably have been on the wing almost continuously for a period of
several months or longer, most individuals are fat and appear to be
in peak physical condition. Sooties appear to be heavier at first land-
ing than at any other time in the breeding season; a number of birds
at this time weighed 190 to 210 grams, whereas weights exceeding
180 grams are unusual later. Ashmole (1963a: 340 ff.) reported that
most Ascension Sooties had completed molt by the time they began
to assemble in the colony at night and the same probably is true of
the Tortugas population. In the hand the plumage of newly-arrived
birds is conspicuously fresh and unworn. The attenuate tips of the
outer pair of rectrices extend as much as 80 mm beyond the adjacent
pair in delicate streamers that are soon lost apparently by abrasion
on land. In June and July the outer rectrices are only 15-20 mm
longer than the next pair.
Tightly packed roosting in the colony is a characteristic group
behavior for several days immediately after the landing. All the
birds settle at once and form a nearly continuous cover over the
ground. The colony then appears much more densely tenanted than
it does after incubation begins, when only one member of the pair
is usually present at the nest.
Also typical of this period are flights, presumably part of the pair
formation ritual, in which two individuals stay very close together.
In these flights two birds leave the ternery, circle to an altitude of
several Ehundred feet, then change to an exaggeratedly deep and slow
wingbeat, meanwhile giving a call that apparently is peculiar to the
occasion. The flights may occur over the colony or remote from it.
They vary in duration from a few seconds to about a minute and may
consist of one ritualized flight or of several alternated with intervals
of normal flight. Flights usually end abruptly with the two terns


making a headlong descent checked a few feet above the water, and
then flying rapidly back to the colony. Occasional two-bird flights
occur throughout the breeding season, but they are more frequent
and protracted among Sooties that have recently landed, when dozens
of flights may be in progress simultaneously.
Brown Noddies on Bush Key perform ritualized two-bird flights
that closely resemble those just described for the Sooty Tern, and
I have occasionally seen similar flights by Royal Terns wintering at
the Tortugas. In all three species a distinct call is associated with
the flights and often drew my attention to the birds engaged in them.
Warham's (1956: 89-90) description of a "Butterfly Flight" of Brown
Noddies and Black Noddies on Pelsart Island, Western Australia,
seems to apply equally well to flights observed at the Tortugas, and
presumably all represent the "High Flight" aerial display that has
been described for many terns.
Sprunt (1948b: 18) and several other observers suspected that the
date of sampling in a given year might affect the estimate of popula-
tion, but no means of quantifying the suspicion existed until Moore
established marked plots on Bush Key in 1952. In 1953 the first
Sooties landed 14 April and the first hatching was noted 23 May.
Moore's nest counts that year (1954 xus.) showed 64,724 adults present
14 May, 81,210 on 2.3 May, and 84,569 on 28 May. From these figures
he calculated that an average of 1832 new birds entered the ternery
daily from 14 to 23 May, and the rate dropped to 672 daily from 23
to 28 May. No later checks were made that season, but general ob-
servations of the colony suggest that Sooties continue to arrive and
start nesting through much of June in some years.
Moore (1954 xis.) suggested that, to assure comparable population
estimates from year to year, density of nesting data should be based
on counts of nests made one week after first hatching. This perhaps
is the most practical solution possible, but the way Sooty Terns arrive
to nest clearly makes it difficult to estimate the size of a colony ac-
curately except from repeated counts spaced to sample the entire
season. Population estimates based on nesting densities made either
much before or much after hatching begins are likely to be too low.
From the known pattern in the Common Tern it appeared likely
that late-arriving Sooties include the young adults returning to the
colony for the first time, and that age at first return is 3 or 4 years.
Several returns recorded in 1937-41, however, seemed to show Sooties
banded as young of the year back in the ternery the first or second
year after banding. We now believe these reports resulted from
mistakes in reading or reporting band numbers.

Vol. 8


No returns of the 5500 juveniles banded in June 1959 were re-
corded in handling a total of 19,327 adult Sooties (927 returns) in
May and July 1960 and 1961 and in May 1962 (4513, 426 returns), but
the 198 returns provided by a sample of 4190 adults taken 8-15 July
1962 included 11 of the 1959 cohort. Samples of adults mist-netted
at the colony in 1963 on 8-11 April (1125, 140 returns), 15-19 May
(4021, 685 returns), and 9-14 July (3807, 362 returns) contained 0,
6, and 50 respectively of the 1959 juveniles. The July 1963 sample
also included 3 returns from the cohort of 10,127 juveniles banded
in July 1960. It thus appears that Sooties first return to the natal
colony late in their third year and first return in force late in their
fourth year. Our data show that although 0.2 per cent of the 1959
cohort of young Sooties returned to Bush Key in their third year, only
0.03 per cent of the 1960 cohort did so. This is of interest in view
of the recovery record of the two cohorts since leaving the colony.
For the young of 1959 not a single recovery has been reported; for
the 1960 group we have 13 recoveries, 6 of which were birds found
dead along the storm track of hurricane Donna of September 1960.
This suggests that the 1960 cohort suffered much heavier mortality
during its extra-Tortugan years.
Late-arriving adults apparently often pioneer in the changes of
colony site. Bartsch (1932: 281) in reporting the first move of Sooties
from the ancestral Bird Key location observed that 30 pairs nesting
on Bush [Long] Key still had eggs or small young on 10-21 August
1932 while most of the young in the main colony were already on
the wing. All of the recorded nestings on Hospital Key and Long
Key were well behind the usual schedule and presumably were initi-
ated by birds that arrived late and failed to find space in the parent
Late-nesting Sooty Terns at the Tortugas seldom succeed in rear-
ing young, in part because they so often nest in unsuitable places
such as the easily flooded sites on Long Key, and in part because
isolated nesting groups are especially subject to predation. In 1963,
for example, we detected no significant loss from predation in the
main colony, but predators (both rats and Cattle Egrets suspected)
destroyed the eggs of an estimated 1500 pairs of Sooties that settled
on the east spit of Bush Key (figure 3) late in the season. No Sootics
had landed on the east spit 24 April but on 15-19 May it was fully
occupied by incubating birds and others that had not yet laid eggs.
Attack by predators must have occurred soon after, because no
Sooties remained there on 5 July, and broken eggs that ranged from
slightly incubated to about ready to hatch were scattered over the


In 1963 most of the 3 and 4-year-old adults apparently arrived
during late May and June and sought space in the main colony rather
than at its edges or at outlying sites. About 95 per cent of the returns
for the juvenile cohorts of 1959 and 1960 were localized in the south-
western one-quarter of Bush Key, the same area in which chicks were
banded most heavily in those years. Large samples of adults taken
farther east on Bush Key (including a sample of 329 from birds then
landing on the east spit, 16 May) included few or no returns of juve-
niles banded in 1959 and 1960. Thus the young adults returning for
the first time seemed to center their activities near the natal nest
location, even though that part of the colony already was densely
occupied. Although strong site tenacity in terns (Austin, 1949) un-
doubtedly serves to maintain the established colony, it must also
weigh heavily against the likelihood of successful breeding by younger
adults. We have no clear evidence that any Tortugas Sooties nested
in their third or fourth years. We suspect that inexperience, late ar-
rival, and site tenacity combine to make successful breeding by young
adults a rare occurrence, at least in colonies where adult mortality
is low and space relatively limited.
Straggling departure is as characteristic of the Tortugas Sooties
as straggling arrival, but this aspect of seasonal change in the popu-
lation has seldom been mentioned. Early writers believed the terns
left in one body or within a few days. Thompson's (1903: 82) state-
ment that the Noddies leave "in great flocks and at night. ... The
entire exodus consumes but two or three days" is typical of com-
ments for both species. Later, Bartsch (1919: 473) quoted reports of
the Bird Key wardens to show that noticeable mass departures oc-
curred over a period of 2 to 6 weeks. More recent observations con-
firm this and do not extend the extreme dates Bartsch mentions, 9
August and 25 September. Although a decrease in the size of the
colony is seldom obvious before mid-August, several lines of evidence
suggest departures begin much earlier.
Birds that do not renest after failing in their first breeding attempt
probably begin to leave the ternery in May. Egg removal experi-
ments by Ridley and Percy (1958) on Desnoeufs Island, Seychelles,
and by Ashmole (1963a) on Ascension show that Sooties are far less
persistent layers than has been supposed. No more than 50 per cent
of those whose first eggs were removed laid a replacement, and re-
nesting seldom occurred after loss of well-incubated eggs or newly
hatched chicks. The few observations at the Tortugas seem to agree
with these findings, and suggest in addition that the likelihood of
renesting begins to decline sharply at a relatively early date in the

Vol. 8


breeding season. Birds whose first eggs were destroyed by predators
on the east spit of Bush Key in late May or June 1963 laid no replace-
ments there. We found nothing in July to suggest that they had
renested elsewhere on Bush Key, nor did there seem to be any sub-
stantial number of unemployed adults around the colony. Appar-
ently Tortugas Sooties whose first breeding effort ends in failure
after about mid-May tend to leave the colony soon afterward without
The earliest departure for which definite evidence exists is that
of an adult banded on Bush Key in May 1960 and found dead at
Ruskin, Hillsborough County, Florida, about 215 miles north by a
little east of the Tortugas, on 25 July 1960. Other banding data,
however, suggest that many adult Sootics leave the colony between
late May and early July. Large samples of adults were taken in
mist nets in both May and July, 1960 through 1963. Extreme dates
of the sampling periods were 15-31 May and 8-17 July, and the loca-
tion and method of capture were virtually the same for all samples.
The 14,884 adults handled in July 1960-1963 included 160 (1.08 per
cent) banded in May of the same year, whereas 12,100 handled in May
1961-1963 included 412 (3.4 per cent) banded in May of the previous
year. The range in the various samples was: May to July repeats,
0.65-1.6 per cent; May to May returns, 2.8-3.9 per cent. May-banded
adults thus occurred three times more frequently in samples of the
following May than in samples taken 5 to 6 weeks later in the year
of banding. The simplest explanation is that many adults present
in May leave before the second week of July. These presumably in-
clude both frustrated breeders whose ties to the colony are relaxed
by loss of eggs or young and early breeders whose young have fledged.
A few dozen to several hundred Sooties usually remain on Bush
Key after the rest are gone. Most are juveniles and most are sick,
injured, or deformed. Only rarely do any survive the fall flights of
accipiters and falcons in late September and October. Tortugas
Sooties seldom abandon healthy young, although reports suggest
this may have happened once or twice when departure was hurried
by severe storms in late August or early September. On 8-11 Sep-
tember 1962 about 50 young birds remained on Bush Key during
the day. Most were obviously infirm and several died every day.
Each evening 200-300 adults and about 100 strong-flying young re-
turned to the colony. All the young were still being fed by adults
and the relative numbers of young and adults suggested that both
members of most pairs were present.



The early departure of some adults probably has not been a
major source of error in population estimates (table 2). The critical
event for estimates based on nest counts is departure of hatchlings
from the nest site, which becomes important even earlier. Nest
density data used in calculating populations in 1945-1948 were re-
corded after mid-June and thus may considerably underestimate
actual numbers. Nest counts of 1951-1956 were made in late May
and in several of these years large numbers of adults were thought
to have arrived and nested after the counts. The knowledge that
young Sooties do not return in force until their fourth year makes the
reported increases of 1903 to 1907, 1913 to 1914, 1939 to 1940, 1942
to 1943, and 1949 to 1950 highly improbable from Tortugan recruit-
ment alone. Those of 1938 to 1939, 1943 to 1944, 1947 to 1948, and
1955 to 1956 seem unlikely in that they leave little room for mortality
in the intervening years.
The reported populations of 1950 and 1951 stand out as much
above other estimates. I have reviewed the field records for these
years and the errors, if any, are in the data, not in the calculations.
Moore and Dilley (1953: 79) suggested that the unprecedentedly high
populations of 1950 and 1951 "may be attributable to several years
being unusually favorable for weather and food." However, evidence
that the relevant years, 1946 and 1947, were marked by especially
successful reproduction is wanting.
In spite of the questions raised above, the reports since 1903 prob-
ably represent the actual trends of population with fair accuracy.
In general, the Bird Key colony of Sooty Terns increased under
protection to about 80,000 to 100,000 breeding adults by around 1917.
The severe hurricanes of 1910, 1915, and 1919 that ultimately caused
a great reduction in the number of preferred nest sites available to
Brown Noddies on Bird Key probably made enough more area avail-
able to Sooties to compensate for the area lost by erosion. In any
event the Sooty Tern population apparently maintained about the
same level from c. 1917 to c. 1930. Disturbance resulting from re-
newed egging in the early 1930's, and probably also from the en-
forced movement of the colony from Bird Key, seems adequate to
account for the reported decrease to about 30,000 adults in 1935.
Within a relatively few years after 1935 the colony, now on Bush
Key, again attained approximately the same upper level that it had
on Bird Key. The view that 60,000 to 100,000 adults represents the
normal Sooty Tern population of Dry Tortugas under protection
(Moore and Dilley, 1953) probably is close to the mark. Fluctuation
within these limits doubtless results in large part from varying success

Vol. 8


in rearing young because of year to year variations in weather, food
supply, and predation, and from varying mortality during the popu-
lation's pelagic phases. Predation is seldom important at Dry Tor-
tugas, although Magnificent Frigate-birds take fair numbers of young
Sooties in some seasons (Beard, 1939; Dilley, 1949 MS.) and instances
of predation by rats, cats (Russell, 1938 Ms.), Laughing Gulls, Larus
atricilla Linnaeus (Watson and Lashley, 1915: 38), and a Great White
Heron, Ardea occidentalis Audubon (Robcrtson, 1962), have been
It is not clear what factors act to set the upper limit attainable by
the Tortugas Sooty Tern population, nor how they act, but I suggest
tentatively that the limits may be determined as much by the species'
behavior pattern as by such environmental factors as food and ter-
ritory. The question of whether or not Sooties are ever crowded
on Bush Key has been debated by authors to no conclusion. It is
clearly a strong departure from normal behavior, however, for Sooties
to nest elsewhere than at the colony site of the previous year or at
the edge of a mass of Sooties already nesting. That Tortugas Sooties
rather frequently have settled at new locations suggests that Bush
Key has been overcrowded at times, however it may have appeared
to human observers.
The obvious question then is, why hasn't the colony spread to
nearby islands that seem fully as suitable as Bush Key? The reason
appears to be that the earlier and more successful breeders tend
strongly to settle at the colony site of the previous year. Present data
indicate that when the progeny of these birds return to nest, they
seek nesting space near the location where they were reared. Such
a pattern tends to maintain a strong nucleus at the expense of possible
colony expansion. The individuals that colonize peripheral or out-
lying locations are those compelled to do so, principally because they
arrive late at the colony. As a group these may tend to be chronic
unsuccessful breeders that have lost site tenacity. The new loca-
tions they occupy are commonly much more exposed to weather and
predators, and late arrival reduces the likelihood of renesting after
disturbance. Thus, the pioneering that might lead to establishment
at new locations and an increase in the local population is almost
always foredoomed to fail.
The successful shift of the colony site from Bird Key to Bush Key
in the early 1930's was probably facilitated, once the area of Bird Key
was reduced to a certain point, by landings on Bush Key early enough
in the season by large enough numbers of birds for successful breed-
ing. The behavior of the colony in 1937, 1938, and 1947 when the



first Sooties that arrived settled on Garden Key is less easily explained.
In these cases, however, unusual conditions apparently existed on
Bush Key at the time nocturnal swarming began, an infestation of
rats in the 1930's (Russell, 1938 MS.; Beard, 1938) and unusually lux-
uriant vegetation in the 1940's (Sprunt, 1948b). Perhaps these dis-
turbances were sufficient to produce atypical behavior.
It was formerly believed (Murphy, 1936: 1125-1127) that all Sooty
Terns deserted their nesting areas for a period of time between breed-
ing seasons. More complete information, however, shows that Sooties
are present in the neighborhood of some colonies throughout the
year. Ashmole (1963a: 301) states .. there was no month in which
Wideawakes could not be seen or heard from Ascension." The same
appears true of colonies off Oahu, Hawaii, studied by Richardson
and Fisher (1950), and of those at Willis Island (Hogan, 1925) and
Raine Island (Warham, 1961), northeastern Australia. Ashmole and
others have drawn a contrast between the colonies where Sooties are
continually present and latitudinally more peripheral colonies, such
as the Dry Tortugas, where they are absent for several months of
the year, but the supposed difference may disappear with more study.
Excepting birds carried north by hurricanes, all recoveries (through
1963) of Sooty Terns banded as adults at the Dry Tortugas have been
within the Gulf of Mexico, indicating that the breeding population
does not disperse widely. The January records cited above leave
November and December as the only months in which Sooties have
not been reported at the Dry Tortugas. The possibility that some
individuals remain within commuting distance and make occasional
night flights over the colony throughout the off-season cannot at
present be excluded.
The Sooty Terns of the Tortugas have often been cited as typical
of the populations that have a 12-month cycle and begin breeding at
about the same time every year. This appears true of records from
the time of Audubon to the early 1940's, all of which indicate that
laying began in late April or early May. Over the past two decades,
however, first eggs have been laid at consistently earlier dates. The
cycle remains essentially annual, but nesting now begins a full month
earlier than it did in the 1940's. The significance of this slow ad-
vance of breeding date and the factors that might account for it
are unknown at present. No relationship to a particular moon phase
(Chapin and Wing, 1959; Ashmole, 1963a: 349) is apparent. The
date of landing and first eggs in 1964 coincided with the full moon,
but the landings of 1961-1963 occurred 23, 14, and 6 days respectively
before the full moon. The earlier breeding of Sooty Terns in the

Vol. 8


Tortugas may be merely another phenomenon of the sort that is
commonly attributed to a supposed trend toward warmer climate.
Another possibility is that earlier breeding is associated with an in-
crease in the size of the colony, although clear proof of an increase
is lacking. Fisher and Lockley (1954) cite instances for many species
of seabirds showing that larger colonies tend to become active earlier
in the season.
Richardson and Fisher (1950) reported that Sooty Terns on two
small islands located about 10 miles apart off the windward coast
of Oahu had distinctly different breeding seasons, the colony on Moku
Manu beginning to nest in November while that on Manana began
in April. They suggested that Manana might have been colonized
by birds from Midway Island or some other distant population which
breeds in the northern hemisphere spring. The possibility that Sooties
nesting on Manana are the overflow from the larger colony on Moku
Mano, however, does not appear to be excluded by the information
so far published about these populations. Egg-laying on Moku Manu
began in November and extended through March, while the season
on Manana is shown as April through June (Richardson and Fisher,
1950: 304, table 4). Thus the season on Manana merely continued
that of Moku Manu, rather than being distinct from it. This char-
acteristic, the fact that the population on Manana varied greatly in
the two breeding seasons observed and that few young were reared
in either season, and the fact that the Moku Manu colony was re-
portedly overcrowded all suggest pioneering of a new site by birds
that arrived late, such as has been observed several times at the
Ashmole (1963a) recently published a highly informative account
of two breeding seasons of the Sooty Terns of Ascension Island. From
his observations of Sooties and other species that breed there he ad-
vanced (Ashmole, 1963b) the hypothesis that competition for avail-
able food within foraging range of the nesting colony was the prin-
cipal factor regulating the numbers of tropical seabirds. No single
study as intensive as Ashmole's has been made at the Dry Tortugas,
but, because of the long record of observations available, the Tortu-
gas colony ranks as perhaps the best-known Sooty Tern population
after Ascension. A comparison (table 3) shows striking differences
between the two populations in mortality factors and breeding suc-
cess. Although it is of the order of one-tenth the size of the Ascension
population, the Tortugas population appears to have reared a sub-
stantially larger number of young in some years.


Ashmole (1963a) describes heavy mortality of Sooty Tern chicks,
apparently from starvation, and presents other evidence indicating
that adults had great difficulty obtaining adequate food for young,
especially in the second season he observed. He cites (1963b: 465)
records similarly consistent with his hypothesis of population con-
trol from other Sooty Tern colonies and for other species of tropi-
cal seabirds. Feeding conditions apparently are so much more favor-
able for Sooty Terns in the neighborhood of the Dry Tortugas that
the likelihood of their numbers being limited by competition for
available food seems improbable. Recent observations at the Tor-
tugas support Watson's (1908: 192-195) statements that the terns
do most of their feeding within a radius of about 15 miles from the
colony. Although much of the feeding occurs outside the Tortugas
lagoon, flocks of Sooties often fish within sight of the colony. Fish
and squids regurgitated by adults returning from feeding commonly
are intact, as if taken but a few minutes before. By contrast Ashmole
(1963a: 333) found that feeding adults were absent for extended pe-
riods; he saw no Sooties fishing near Ascension; and the food remains
regurgitated by the birds were seldom recognizable. From his
studies of the development of young on Ascension, Ashmole concluded
(1963a: 320): "The capacity of young Wideawakes to survive for long
periods on relatively little food, while growing hardly at all, but to
accept large quantities of food when it is available, is clearly an
adaptation to an environment in which the food supply is precarious."
While few detailed data are available on the development of young
Sooties at the Tortugas, general observations indicate that their growth
is regular and rapid and that the ability to fly short distances is at-
tained at 5 to 6 weeks of age, in contrast to the fledging period of 8
weeks or longer recorded for Ascension (Ashmole, 1963a: 321). No
incidents of mass starvation of young Sooties have been reported at
the Tortugas.
Acknowledging the need for additional critical data, it appears
that no shortage of food available to Sooty Terns exists in the vicinity
of the Dry Tortugas colony; and, therefore, that competition for food
during the breeding season cannot be the factor that checks the in-
crease of the population short of the limits of available nesting terrain.
As suggested above, social factors-in particular, site tenacity in young
adults, and the tendency of late arrivals to choose insecure nest sites
near the colony rather than more secure sites at a little distance-
appear to operate in a density-dependent manner to limit growth of
population. Ashmole (1963b) is surely correct in pointing out that
competition for nesting space could not regulate total species popu-


lations of seabirds effectively because individuals that fail to find
space in one colony could go elsewhere. How much movement of
this sort actually occurs in the Sooty Tern is a moot point at present.
It seems likely that asynchrony of breeding cycles would inhibit ex-
change of birds between some colonies. Whatever the factors are
that ultimately control the total number of Sooty Terns, however,
observations at the Tortugas strongly suggest that social forces can
effectively regulate numbers at the level of the individual colony.


Ascension Island Dry Tortugas

Estimated number of
Breeding adults

Nesting space available

Mortality of adults at
the colony

Mortality of eggs and
young caused by

Mortality of eggs and
young caused by

Mortality resulting from
attacks upon stray
chicks by adult Sooties

Mortality of young from

Over-all breeding suc-
cess (young fledged
as a of eggs laid)

Interval from start of
one breeding period to
start of the next.

c. 750,000

Relatively unlimited

Estimated 1 to 3% killed
by feral cats; predation
proportionally heavier
on the earlier breeders.

Heavy (cats and frigate-

No information


Frequent, much heavier
in some breeding seasons.

Low, estimated at c. 10-
20'% and perhaps not over
2%' in two successive
breeding periods.
c. 9.7 months

c. 80,000

Limited on Bush Key,
several other habitable
islands nearby.

Insignificant, virtually
all losses result from birds
becoming entangled in

Usually minor (mainly
rats), except in outlying
nesting areas.

Occasionally heavy at
low sites and when hur-
ricanes occur during the
breeding season.

Probably the major cause
of mortality of chicks.

Never reported.

Probably seldom below
70V' in years with no
summer hurricanes.

c. 12 months


19th Century. Accounts by naturalists who visited the Dry Tortugas
in this period suggest that Brown Noddies were numerous and identi-
fy the keys on which they nested at different times, but beyond that
contribute relatively little to the history of the Tortugan population.
In 1832 all the Noddies were nesting on Bush Key, but "several
thousand" nests not in use were seen on Bird Key as well, leading
Audubon (1835) to suppose that Sooty Terns had driven the Noddies
from the latter island not long before. Bartsch seems to have thought
that Noddies persisted in nesting apart from the Sooties until Bush
Key was washed away around 1870. He writes (1919: 482): "Since
then [1832] the colony has been forced to make a complete shift
and the choice between Bird and Loggerhead Key has fallen to the
former Wurdemann (1861) and Bryant (1859a), however, re-
ported both species nesting on Bird and East Key in the 1850's. The
separation of the nesting areas of Sooties and Noddies observed by
Audubon may well have been temporary, for his report seems to be
the only record that Noddies nested on Bush Key during its 19th
century emergence. In 1890 (Scott, 1890) most of the Noddies were
nesting on Bird Key.
Several of the early reports suggest that Noddies and Sooties
were then about equally abundant at the Tortugas. Audubon (1835:
268) wrote of the Noddies: "They nearly equal in number the Sooty
Terns ...." Scott (1890) was informed that Noddies were more com-
mon than Sooties, and Holder's (1892: 194ff.) account of an egging
sortie to Bird Key about 1860 suggests the same.
Later writers have taken such comments to indicate that the Noddy
population of the Dry Tortugas suffered particularly severe reduction.
Job, for example, remarked (1905: 87-88) of his observations in 1903:
"Of the Noddies there are hardly a thousand, which is a great de-
crease from the numbers that were once here." It appears to me
that the reports of near parity in numbers of Sooties and Brown Nod-
dies are more likely evidence that the population of Sooties had been
much reduced. The Brown Noddy at the Dry Tortugas has always
nested mainly in bushes. No report suggests otherwise. More spe-
cifically, its nesting is confined largely to the edges of clumps or
thickets of bay cedar. Few nests are placed within dense shrubbery.
From what is known of the vegetation of keys where the ternery has
been located, it seems certain that the Sooties, nesting in dense masses
on the ground, would always have been able to reach much greater


numbers than the Noddies before their increase was limited by
scarcity of nest sites.
It seems likely also that, in a ternery such as the Dry Tortugas,
the Noddy population might be expected to decline more slowly than
the Sooties under the pressure of sustained egging. Eggers preferred
Sooty Tern eggs to those of other terns. Many writers mention this,
Audubon, for example, informing us that eggs of the Sooty are "de-
licious, in whatever way cooked .... Because Sooties nest closer
together and are much more strongly territorial than Noddies, re-
peated disturbance of a mixed ternery would almost certainly re-
sult in disproportionately high mortality of Sooty Tern chicks. The
fact that Noddy nests are more scattered and placed in heavier cover
would make it much more difficult for eggers to gather an entire lay-
ing. Finally, the usual nesting season of Noddies at the Dry Tor-
tugas is considerably more extended than that of the Sooties which,
again, would make loss of an entire season's production less likely.

1902. Thompson (1903) presented an excellent and well-illustrated
life history study of the Brown Noddy as observed on Bird Key in
1902. His account includes the earliest clearly stated estimate of
the size of the population: "As nearly as can be judged it [the Nod-
dy colony] contains about three thousand individuals."

1903. As with the Sooty Tern, several estimates of the number of
Noddies are available from 1903 observations by Burton and Job.
They are "about 400" and "at least 600" (Burton in Dutcher, 1904),
and "hardly a thousand" (Job, 1905). All summaries of the history
of the population cite the number in 1903 as 400 and credit it to
Job. The warden's end-of-season figure of 600, however, seems the
best estimate available.
The warden on Bird Key (in Dutcher, 1905) said that the terns
had a successful season in 1905, and Mayer (in Dutcher, 1906) re-
ported in 1906 that Noddies on Bird Key had increased since 1898
but not so much as the Sooties. Nothing else is known about the
colony of Noddies for the period 1903-1907.

1907. As part of his remarkably varied investigations on Bird Key
the summer of 1907, Watson made the first known estimate of the
Tortugas population of Brown Noddies based on a direct count of
their nests. He published two explicit descriptions of his method
and results (Watson, 1907: 311, 1908: 197). The latter reads: "By
means of a mechanical counting device it was found possible actually


to count the total number of (active) Noddy nests. The count gave
603 nests. In some places, where the bay-cedar bushes are very
dense and the area has to be covered 'dog-fashion' (or at times even
still more primitively), and in others where the cactus growth is very
luxuriant, error in counting was easily possibly. On account of
these possibilities of error, I believe that 700 nests is a more repre-
sentative number. Since two birds occupy one nest, we have a total
of 1,400 adult noddies on the island."
Despite Watson's abundantly clear exposition all later references
except Bent (1921: 303) and Longstreet (1936b, but not 1936a) give
the 1907 population as "4000". Many, in addition, cite "1400" as the
population in 1908, crediting this figure also to Watson, and the
apparent decrease has drawn comment: e. g., "The noddy population
took an unexplained drop from 4,000 in 1907 to 1,400 in 1908" (Sprunt,
1947b: 215).
Two errors are involved here. They seem to stem respectively
from a mistake in Bartsch's (1919) account of Watson's observations,
and from misreading of Bartsch, who gives two figures for the num-
ber of adult Noddies on Bird Key. The first (1919: 471) occurs in a
table and reads: "Noddy tern, estimated, adult' 4,000." The numeral
"1" refers to a footnote on the same page that reads: "Based upon
Doctor Watson's census of 1908". The figure, "4,000," appears to
be a lapses and, as pointed out under Sooty Tern, Watson apparently
did not work at the Dry Tortugas in 1908. The second reference
(Bartsch, 1919: 482) gives the correct figure but attributes it to the
wrong year: Watson estimated the presence of 1,400 adult
birds in 1908."
Three points seem clear from the tangle of mistaken citations:
1. Watson's estimate of the population in 1907 was 1400 adults. 2.
No estimate of the population of Noddies in 1908 exists. 3. Com-
pilers have often cited Watson from Bartsch or from one another
rather than from Watson.
Watson touches upon a problem that has plagued many later
observers of Noddies at the Dry Tortugas in his comment (1907: 311):
" one feels that there is a vastly greater number [than 1400]
present." He concluded that many of the Noddies at Bird Key were

1909-1929. The reports of wardens stationed on Bird Key to the
National Association of Audubon Societies (through 1919) and to the
Biological Survey include estimates of the number of adult Noddies
for all years of the period 1909-1919, and for 1929 (table 4). The


estimate for 1910 is based upon another count of Noddy nests by
Watson. The warden's estimate for 1918 was published (in Pearson,
1918). Howell (1932: 272) summarizes all the reports, presumably
from the files of the Biological Survey, mentioning specifically the
population figures of 1910, 1916, and 1929. Stevenson (1938: 307)
also refers to the 1929 figure. Other summaries jump from 1908 to
1917 to 1935. All who include 1917 (Vinten, 1943: 57; Fisher and
Lockley, 1954; 60; et al.) give the population that year as "4,000"
citing the figure from Bartsch (1919). Bartsch, however, made no
independent estimate of the population in 1917. Curiously, this esti-
mate of "4,000" is the same as that credited (mistakenly) to Watson
in both 1907 and 1908.


Year of Method Reference

1902 3000 Estimate Thompson (1903)
1903 600 Estimate Burton (in Dut(her. 1904)
1907 1206 Nest Count Watson (1908a)
1909 5000 Estimate Peacon (1909 ms.)
1910 1710 Nest Count Ashe (ms. notes)
1911 2000 Estimate Peacon (1911 ins.)
1912 1500 Estimate Peacon (1912 ms.)
1913 600 Estimate Peacon (1913 ms.)
1914 2500 Estimate Peacon (1914 ms.)
1915 5000 Estimate Asle (ms. notes)
1916 6000 Estimate Bethel (1916 ins.)
1917 10.000 Estimate Lowe (1917 ms.)
1918 15.000 Estimate Ashe (in Pearson, 1918)
1919 35,000 Estimate Ashe (1919 ms.)
1922 1600 Estimate Bartsch (ms. notes)
1929 3000 Estimate Park (ms. notes)
1935 10001 Estimate Mason (1936)
1936 4000 Estimate Doe and Russell (1936)
1937 2000 Estimate Longstreet (1937)
1938 392 Nest Count Beard (1938)
1939 380 Nest Count Robinson (1939)
1939 454 Nest Count Taylor (1939 ms.)
1940 180 Estimate Robinson (1940)



TABLE 4 (continued)

Year of Method

1940 750 Estimate
1941 400 Estimate
1941 1000 Estimate
1942 450 Estimate
1945 625 Estimate
1946 492 Nest Count
1947 202 Nest Count
1948 282 Nest Count
1949 566 Nest Count
1950 490 Nest Count
1951 518 Nest Count
1952 890 Nest Count
1953 842 Nest Count
1954 970 Nest Count
1955 1108 Nest Count
1962 2130 Nest Count


Felton (1940 ms.)
Stimson (in litt.)
Peterson (in Vinten,
Budlong (in Vinten.
Sprunt (1946a)
Sprunt (1946b)

Sprunt (1948a)

Sprunt (1948c)

)illey (1950)

Moore and Dilley (195,3)

MN\ore and I)illey (1953)

Moore and Dilley l195-3)

Moore (1954 ins,)

Moore (19.54 ms.)

Moore (1955 ms.)

\\. B. and Bett Robhertson

In many years when the breeding population of Brown Noddies was de-
termined by counting nests, observers added an arbitrary figure (commonly 10
per cent) to account for nests not found. Population estimates that include such
additions are placed in parentheses beneath the figure based on the actual nest

Figures for 1915-1919 quoted from the warden's reports (table 4)
contradict all previous comments regarding peak populations of Nod-
dies at the Dry Tortugas. Even if the figures are substantially dis-
counted to allow for overenthusiastic interpretation by the individ-
uals directly responsible, it appears likely that the Tortugan Noddy
colony reached by far its highest population during this period.

1930-1.934. The only references to the colony in these years that I
have seen are by Bartsch (see Sooty Tern) and his comments include

Vol. 8


no estimate of population. In 1931 Bartsch (1931: 373) noted: 'This
change of vegetation [destruction of bay cedar on Bird Key by storms]
has forced the noddy terns to change from a tree-nesting to a sand-
nesting habit." The following year (Bartsch, 1932: 281) most of the
Noddies still nested on Bird Key, but about 70 nests were found in
dense mats of sea purslane (Scsvinum portulacastsum) on "Bush
[Long] Key," 32 nests in bay cedar bushes on "Long [Bush] Key,"
and a few nests on pilings around the coaling docks on Garden Key,
where apparently few young were reared because most of them fell
off into the water. Presumably Bartsch's (1933: 267) statement that
"more than half" the colony had left Bird Key in 1933 referred to
Noddies as well as Sooties.

1935-1962. The population figures (table 4) require little comment.
Data for most of the years were obtained by direct counts of nests.
In the years for which independent estimates are available, observ-
ers were in close agreement except in 1939, 1940, and 1941. Having
no basis for a decision between the two figures available for each
of these years, I have included both. The entire colony of Noddies

Fict:I: 8. Brown Noddies nesting on Garden Key. 1938: (a.) pair at nest on a
pile of sea oats (Uniola paniculata) cut and raked before the terns landed; (b.)
adult incubating egg laid on barn ground amid sea purslane. (National Park
Service photographs by Daniel B. Beard.)


nested on Bush Key in most years of this period. Departures from
this pattern, all involving nesting on Garden Key, were reported as
follows: 1936, 6 pairs nested on docks and pilings (Doe and Russell,
1936); 1937, most of the population nested on Garden Key (Young
and Dickinson, 1937; Longstreet, 1937; Russell, 1938 Ms.); 1938, nest-
ing was divided about equally between Garden Key (figure 8) and
Bush Key (Beard, 1938); 1939, about the same division as in 1938 (Rob-
inson, 1939; Taylor, 1939 Ms.); 1947, 9 nests on Garden Key (Sprunt,
1948a); and, 1948, 1 unsuccessful nest on Garden Key (Sprunt, 1948c).

Brown Noddies have been handed and recaptured at the Dry Tortu-
gas in much smaller numbers than the Sooty Terns, and banding data
contribute little to an analysis of past population records. It seems
likely, for example, that Brown Noddies do not return to breed for
the first time until they reach three or four years of age, but no proof
is available. Reasons for believing that a Brown Noddy population
is likely to decrease more slowly than a Sooty Tern population under
the pressure of egging and disturbance are given above. Similar
reasoning from the information available on nesting dates, nest site
preferences, and mortality of Brown Noddies at Dry Tortugas helps
to explain parts of the population record (table 3), but the fit is a
good bit poorer than for the Sooty.
The main difficulty in estimating populations of Brown Noddies
seems to arise less from the obvious influx of new birds than from
long-delayed nesting starts by birds already in the colony. This
has vexed nest counters more than those who undertook to estimate
the number of adults in the area. As mentioned above, Watson
doubted that his count of nests in 1907 recorded the entire Brown
Noddy population. Some later observers also have felt that the num-
ber of nests found failed to account for the adults on hand. Obser-
vations at the Dry Tortugas in 1960-1963 suggest that it is usual for
nesting starts by Brown Noddies to be distributed over a period of
at least 10 weeks from April to early July. It may be that the loafing
birds Watson and others considered to be non-breeders were merely
late breeders.
To illustrate, 39 (13.1 per cent) of 298 Brown Noddy nests that
I examined on Bush Key 11-15 July 1962 contained eggs. All that
were marked for later checking contained eggs or small young on 2
August, and young that were not more than half-grown on 8-11
September. 1 have no reason to think that any of these nests repre-

Vol. 8


seated renesting after failure of earlier attempts, but the possibility
cannot be excluded.
Dates of Watson's 1907 count of nests are not recorded, but esti-
mates of the Brown Noddy population in 1939, 1945, 1946, and 1948
through 1955 are based on counts of nests during or before the third
week of June. It is to be suspected that these underestimate the
breeding population. Nest counts near the end of the season present
no difficulty because with few exceptions recently used nests are
easily distinguished from any nest remnants that may persist from
the year before.
The unproductiveness characteristic of late nesting by the Sooty
Tern at the Dry Tortugas seems untrue of late nests of the Brown
Noddy. No decline in the attentiveness of adult Brown Noddies with
young in the nest in early September 1962 was apparent. Barring
accidents of weather or predation, the young seemed likely to fledge
successfully. Such accidents, of course, become more likely as the
season of hurricanes and hawk migration advances at the Dry Tor-
Factors other than human predation believed to have affected the
Brown Noddies in the Tortugas colony at various times are predation
by rats, mortality caused by storms during the breeding season, and
storm damage to bay cedar bushes. The population record since
1900 reflects to some extent the recorded occurrences of rat infesta-
tions and severe storms. Information below on hurricane occurrence
is taken from contemporary reports and from Dunn and Miller (1960)
and Tannehill (1950).
The years from 1900 through 1910 had one bad summer storm,
16 June 1906, and an infestation of rats on Bird Key is said to have
been eliminated by 1908 (Dutcher, 1908b; Mayer, 1908). I suspect
that Thompson's (1903) estimate for 1902 was near the mark, and
that the 1903 Job-Burton estimate was much too low. Except for
the 1903 figure, agreement between the population record and the
record of disturbance is reasonably good. The colony appears at
first to have increased slowly; then to have declined slightly, and
then once more to have increased slowly to the end of the period.
I have seen no record that rats were present on Bird Key in ap-
preciable numbers after 1908. In the decade 1910-1919 Bird Key was
repeatedly battered by hurricanes. The first of importance, 15-17
October 1910, did great damage to the bushy vegetation (see p.
8). Early hurricanes of great severity occurred on 13-15 August and
3 and 28 September 1915, and 4 July 1916. In 1919 the Dry Tortugas


Vol. 8

were hit squarely by a hurricane of extreme intensity on 10-11 Sep-
Records of the Brown Noddy population for this period, all from
warden reports, show a decline through the season of 1913, then a
meteoric rise to the 1919 figure, the highest ever reported for the
colony. This record does not tally satisfactorily with the record of
disturbance, if one assumes that the Tortugas colony is a discrete
population all of whose surviving adults return to breed annually.
The 1910 hurricane was too late in the season to lave caused much
direct mortality, and its damage to vegetation cannot have affected
breeding success before the season of 1911. Available information
(Peacon, 1911 Ms.; Ashe, iM. notes) suggests that nesting in 1911 was
normal. For 1912 and 1913 the reports indicate abnormal behavior
and great decreases in the number of adults which the observers
attributed to scarcity of nest sites. Ashe (Mxs. notes) reported that
Noddies were seen on Bird Key on 20 March 1912, an unusually early
date, but by 22 May only an estimated 400 had appeared, although
the population later increased to about 1500 (Peacon, 1912 Ms.). In
another report on the 1912 season Watson (1912 rM.) advises planting
bay cedar bushes on Bird Key "in large quantities." The report for
1913 (Peacon, 1913 sis.) lists only 6() adult Noddics and comments on
the great decrease of the species. If the population data are consid-
ered at all reliable, the decreases of 1912 and 1913 must have resulted
from the failure of adults to return to the colony. The decline seems
too early and too abrupt to result from less successful breeding after
Interpretation of the population trend for the years 1914-1919
presents even greater difficulty. It is known (Bowman. 1918) that
shrubby growth on Bird Key had recovered to a considerable extent
by 1915-1916. The 1910 storm may have greatly increased the amount
of thicket edge, and hence the number of available nest sites, by
breaking up formerly solid stands of bay cedar. If so, conditions
favorable to a rapid increase of Noddies may have existed by 1914
or 1915. The year-to-year increase, however, is much too large to
be accounted for entirely by the successful breeding of a discrete
Tortugas population. Moreover, the summer storms of 1915 re-
portedly caused heavy mortality of adult and young Noddies on
Bird Key (Ashe and Bethel, 1915 Ms.) and the same is almost surely
true of the hurricane of 4 July 1916.
The 1919 storm stripped Bird Key of vegetation and also killed
"many" terns (Ashe, 1919 Ms.). This storm greatly reduced the num-
ber of suitable nest sites and no record of recovery of the bay cedar


growth exists. Unless the 1919 storm killed most of the adults, how-
ever, the decrease from 35,000 in 1919 to 1600 in 1922 can be ex-
plained only by the failure of many adults to return to the Dry Tor-
tugas. The records indicate no decline in numbers over the years
of the enforced movement of the colony from Bird Key to Bush Key.
The more recent record shows low points in the late 1930's and
the late 1940's. The first was attributable to rat predation. The
causes of the second downturn are more obscure, but records sug-
gest that rats may again have been a factor. None of the reports
for 1936 mentions the presence of rats, but Russell (1938 Ms.) who
spent the summers of 1936 and 1937 at the Dry Tortugas, states that
rats became common around the fort in the fall of 1936 and that
some apparently swam the channel to Bush Key. By the summer of
1937 rats were so numerous on Bush Key that they could be seen
commonly by day and "by thousands" at night. Russell believed the
Brown Noddy nesting season of 1937 a nearly complete failure, with
more than 90 percent of the eggs and young lost to rats. Other au-
thors give substantially the same account. I have seen nothing to
indicate that the rats killed adult Brown Noddies in any numbers,
yet only some 400 adults were in the colony the following year. As
the population figures for these two years are open to little question,
either the adult population suffered extra-Tortugan mortality of a
catastrophic nature or most of the population either bred elsewhere
or not at all. Practically no information is as yet available on colony
fidelity in this species and the factors that may modify it.
Whatever its mortality in the extra-Tortugan phases of its annual
cycle, the mortality of Brown Noddies at the ternery is low, certainly
much lower than in the Sooty Tern. On 11-15 July and 8-9 Septem-
ber 1962 James B. Meade (in July), my wife, and I counted and buried
all of the dead terns and unhatched eggs that we could find on Bush
Key and counted and examined all the nests of Brown Noddies. The
total observed mortality of Brown Noddies was 2 downy chicks and
11 larger juveniles, a remarkably low 1.23 percent. Although some
of the young still unfledged in early September may not have ma-
tured, this suggests that under favorable conditions-no predators and
no summer storms-Brown Noddy populations can closely approach
their maximum possible rate of increase for the egg to fledging stage.
The Tortugas colony has been largely free of disturbance by rats or
severe summer storms since the early 1950's, and the increase of
Brown Noddies in the past decade may approximate that possible
for a colony of its size performing as a discrete reproductive unit.
As in the Sooty Tern, the question of factors limiting the popu-


lation of Brown Noddies is debatable. In 1936 when the colony
last reached the level of about 4000 adults, unusual numbers were
found nesting on the ground. Several authors (Doe and Russell,
1936; Alien, 1936; Dickinson, 1941) suggested that this behavior re-
sulted from the scarcity of nest sites in bushes, but Beard (1938: 10)
disagreed stating: "There are more available nesting locations for
both species of birds [on Bush Key] than were ever present on Bird
Key." In 1955 Moore (1955 Mls.) reported the highest total of Noddies
since 1937 and commented: it was amply evident afield that
Bush Key has suitable Suriana thicket-edge for the nesting of twice
this present population."


Number of Nests
Nest Site 1962 1963

Dead Bushes (mostly Suriana) 68 76
Sea Rocket (Cakile lanceolata) 15 27
Bay Cedar (Suriana martima) 20 13
Bare Ground 10 10
Spurge (Euphorhia buxifolia) 10 2
Prickly Pear Cactus (Opuntia sp.) 5 7
Sea Lavender (Tournefortia gnaphalodes) 5
Sea Oats (Uniola paniculata) 1
Sea Purslane (Sestivhiur portulacastrum) I

Total 134 136

Close study of records of the vegetation of Bird Key suggest that,
although it was a much smaller island, it may well have had more
brushy edge at times than Bush Key does now, particularly if the
surmise that hurricanes fragmented formerly solid thickets is cor-
rect. As of 1962, the point of doubled population to which Moore
referred had been reached. Brown Noddies in recent years have
used a wide range of nest sites in addition to the typical bay cedar
fringes. Table 5 shows nest sites used by Noddies in 1962 and 1963
in the area between the shore of the western half of Bush Key and
the outer edges of the large bay cedar thicket in the center of the
island where most of the Noddies nest (see figure 3). Nests in isolated
living bay cedar bushes and some of those in dead bushes were built
in a typical manner and within the usual range of heights above

Vol. 8


ground, roughly 3 to 10 feet. Most other Noddy nests in this section
were no more than 18 inches from the ground, rudimentary in form,
and intimately surrounded by nesting Sooties. Parent Noddies at
nests of this sort tended to participate in the panics that affected
nearby parts of the Sooty Tern colony, and the young seemed to be-
gin leaving the nest earlier than they ordinarily do nests in bay cedar.
often as downy chicks.
In both years a large proportion of the Brown Noddy nests placed
at low sites within the Sooty Tern colony probably failed. Observed
pre-fledging mortality of Noddies was strongly concentrated in this
part of the colony, particularly near ground nests and those placed
on low herbaceous plants such as sea rocket, spurge, and cactus. In
July of both years evidence of nest success in the form of adults at-
tending young birds was conspicuously absent from the vicinity of
many such nests in addition to those where unhatched eggs or dead
young were found. Mortality recorded at the western outlying nests
in 1963 (3 eggs and 16 chicks) cannot be related to total mortality
because I did not examine all of the Brown Noddy nests in the cen-
tral thicket. In 1962, however, 77 percent of the observed mortality
(10 of 13 chicks) was associated with nests outside the main thicket in
the western half of Bush Key, which comprised only 13 percent (134
of 1065) of the total nests of Brown Noddies.
When the Brown Noddy population on Bush Key is high, use
by Noddies of nest sites not typical for this colony apparently in-
creases, and such nests are much more likely to fail than are those
placed in live bay cedar bushes. Shortage of secure nest sites in the
immediate vicinity of the established colony thus may tend to set an
upper limit of population size. Ashmole (1936b: 458-459) argues that
competition for nest sites seldom regulates population size in tropical
sea birds because individuals that fail to get nesting space in one
colony can usually emigrate to less crowded colonies or found new
colonies. Acknowledging that successful emigration must have oc-
curred many times in the history of every wide-spread colonial spe-
cies, it is clear that attraction to the known breeding place is a po-
tent countering factor in some sea birds. Potential emigrants seem
far more likely to expend their reproductive effort at less secure sites
in or near the existing colony. At least four times the presumed
overflow population of Brown Noddies at the Tortugas has used atypi-
cal sites on the island the colony was occupying rather than moving
into vacant bay cedar thickets on other islands near by-on Bird Key
after the hurricane of 1910 and in the early 1930's; on Bush Key at
times of peak numbers in 1936 and at present.


Indeed, Ashmole's (1962) account of the Black Noddies of Boat-
swain Bird Island, Ascension, suggests they behave similarly when
faced by a shortage of preferred nest sites. Crowding there report-
edly led them to use unsuitable ledges rather than colonize new cliffs.
Strong attraction to the traditional breeding place seems often to
inhibit the emigration of surplus birds and to determine a sequence
of events that effectively regulates colony size.
Except for Audubon's fanciful account, the effects of a concen-
tration of Sooty Terns upon the behavior, and perhaps the numbers,
of Brown Noddies nesting in the same area has scarcely been consid-
ered. Nest sites of most of the Noddies on Bush Key are ringed by
nesting Sooties. Disturbance by the Sooties may figure at least in-
directly in the poor success of low nests of Noddies within the Sooty
Tern colony. At other times presence of the Sooties seems to have
favored the Noddies. Russell (19.38 MS.) noted that almost the only
Noddies whose young survived the rat plague of 1937 were those
nesting in bushes surrounded by dense concentrations of nesting
Sooties. Contacts between Noddies and Sooties are infrequent in
the Tortugas colony. Of a similar aggregation on Pelsart Island,
Western Australia, Warham (1956: 89) stated: there seemed
to be no friction between the two species." Nevertheless, the be-
havior of Brown Noddies at the Dry Tortugas differs considerably in
the absence of the Sooties. Where Sooties are present in numbers,
Brown Noddies entering and leaving the ternery tend to fly fairly
high and relatively few are taken in mist nets set on the open beach-
es. In September 1962 with most of the Sootics gone, Brown Nod-
dies swooped from perches on the bush tops and left the island in
low, rapid flight. This difference in their behavior is reflected in the
mist-net catches of 9-10 September when Brown Noddies were taken
at an average rate (8 per net hour) that far exceeded any previous
results for nets set in the open.


Audubon reported Roseate Terns, Sterna dougallii Montagu, nesting
in the Florida Keys (Howell, 1932: 264), but he and other early ob-
servers apparently saw none at the Dry Tortugas. The first were
reported by Bartsch (1919) who located a breeding colony of about
100 pairs on Bush Key or Long Key in July 1917. Most reports of vis-
itors to the area since 1917 have included some mention of Roseate
Terns. Three partial summaries of the records have been published
(Sprunt, 1948a, 1949, 1951), the last carrying the local history of the

Vol. 8


species through the breeding season of 1949. Table 6 shows the
record of breeding occurrence of the Roseate Tern at Dry Tortugas,
1917-1963. The reports for a number of the years in this span de-
serve comment.

1917. In initial remarks on his observations of the summer of 1917,
Bartsch (1918: 171) wrote: probably 200 common terns formed
a rookery on the rough coral shore of the eastern end of the island
[Bush Key]. Their young birds of various ages could be seen at all
times." The later, more detailed report of the 1917 breeding season
(Bartsch, 1919) does not mention the Common Tern in text, but dis-
cusses (p. 489) a colony of "about 100 pairs" of Roseate Terns breed-
ing "on the rough coral and shellstrewn northeastern end of Long
Key." The legends to Plates 27-32 in the 1919 publication, a series
of photographs of the colony site and of young birds, state that the
pictures show Common Terns, but a footnote (p. 500) corrects this
to read Roseate Tern. The downy chick shown in one of the photo-
graphs (Bartsch, 1919: Plate 28a) is clearly a Roseate. It appears
certain that these reports refer to a single colony. A Roseate Tern
specimen in the U. S. National Museum Bartsch collected on Bush
Key 17 May 1919 doubtless served to establish the correct identity.
In his writings on the Tortugas, Bartsch appears on some occa-
sions to have followed the nomenclature of older charts on which
application of the names Long Key and Bush Key is reversed from
present usage. For this reason it is impossible to determine conclu-
sively whether the Roseate Tern colonies of 1917, 1921, 1922, and
1932 were located on the eastern sandspit of Bush Key or on one of
the ricks of coral fragments that comprise the island now known as
Long Key. This uncertainty is of little importance, because the
sites are similar and not more than a few hundred yards apart.

1921. Nests with 1, 2, and 3 eggs were seen (Bartsch, Ms. notes).

1922-1925. According to Bartsch (MS. notes), the terns were assem-
bled at the colony site 14 May 1922, but had not begun to nest. For
7 June 1924 he noted, "some seen, but colony not breeding." He
also observed numbers of Roseate Terns feeding in the Tortugas area
on 5 September 1923 and 12-18 August 1925.

1935. The total shown in table 6 is a synthesis of the estimates by
members of the Audubon party. Some of the observers thought that
no more than 100 pairs of Roseates were in the colony (Russell, 1938


rs.). Mason (1936 and in litt.) reports young 3 or 4 days old and
many eggs not yet hatched.

1936. The published account (Doe and Russell, 1936:7) states:
"There were probably about the same number of roseate terns as
last year but Doe (MS. notes) recorded that the Roseate Tern
colony numbered "about 400 nests." Nesting was apparently just
beginning, for Mason (MS. notes) wrote "96 nests located, all with 1
or 2 eggs. None yet hatched."

1937. Reports of the trip (Young and Dickinson, 1937; Longstreet,
1937) give no estimate of the number of Roseate Terns seen. This
omission misled Sprunt (1949, 1951) to state that none bred at the
Dry Tortugas in 1937. Young and Dickinson, however, mention (pp.
3-4) that they visited a key where Roseate Terns were nesting and
they include (p.6) a photograph with the legend, "Roseate Tern
banded by C. R. Mason on Sand Key." Banding schedules show
that Mason banded three adult Roseates on Sand (Hospital) Key on
25 June 1937 and he advises (in litt.) that as he recalls it the colony
nesting there was slightly smaller than the ones observed on Bush
Key in 1935 and 1936.

1938. Mason (1938: 1) noted the nesting colony included "better
than 300 birds." His Ms. notes record that 157 nests with eggs were
counted, and that the location of the ternery, not mentioned in the
published report, was the eastern sandspit of Bush Key.

1940. Nesting was just beginning. The observers found 5 nests each
containing a single egg. In addition, the Fort Jefferson Custodian
told Robinson (1940: 3) that "quite a number" of Roseate Terns were
believed to be nesting on Bird Key, then re-emerging as a sandbar.

1941. The brief report of the two parties of observers who visited the
Dry Tortugas in June 1941 (Rea, Kyle, and Stimson, 1941) mentions
only that Roseate Terns were nesting on Bush Key. Louis A. Stim-
son (in litt.) writes me that the first group saw but one Roseate Tern,
in flight over Fort Jefferson. Roger T. Peterson, who accompanied
the second group, writes me (in litt.) that he saw no Roseates, but that
the Custodian of the fort told him there was a nesting colony on the
east end of Bush Key. Individuals who had seen the colony in both
years told R. R. Budlong (1942 MS.) that it was about the same size in
1941 and in 1942. As with the 1937 report, lack of a definite popula-

Vol. 8


tion figure has resulted in the statement that the species was absent
from Dry Tortugas in 1941 (Sprunt, 1949).

1942. The colony was located "on the reef between Bush and Long
Keys." The total shown seems to have been only a rough estimate,
the author commenting that he was able to visit the area but once,
on 2 July, and found "numerous eggs and young birds."

1943. No population figure is given in the report which merely states
"The Roseate Tern colony seems to contain about the same number
of birds as last year."

1947. Sprunt (1948a: 29) counted 67 nests on Long Key, 54 on Bush
Key, and 21 on Hospital Key. In addition, about 12 young (not in-
cluded in the total) survived from an earlier nesting on Long Key
disrupted by high tides.

1948. Sprunt (1948c: 14) counted a total of 216 nests with hatching
"about 50% complete." He adds: "It is virtually certain that a few
were missed, despite care. A total of 225 is very likely."

1949. Dilley (1950: 68) located 44 active nests on Bush Key and 7
on Long Key. An additional 17 nests on Hospital Key (not included
in total) are said to have been abandoned.

1950-1952. Nest locations in the three years were: 1950, 55 on Bush
Key and 7 on Hospital Key; 1951, 35 on Hospital Key and 33 on Long
Key; 1952, 136 on Long Key and 58 on Bush Key. From observations
later in the summer of 1950, John R. De Weese (in litt. to Dilley)
reported storm tides flooded all the Roseate Tern nests so that no
young were reared that year.

1953. An earlier group of 9 nests on Middle Key, the only Roseate
Tern nests present in the area on 26 May (Moore, 1954 MS.), was de-
stroyed by high tides during a storm 28-30 May. The later nesting
included 79 nests on Hospital Key, 26 on Middle Key, and 15 on
Bush Key.

1956. When counted, many of the nests (88 on Long Key and 14
on Bush Key) had incomplete clutches and a number of fresh nest
scrapes without eggs were present. Two weeks earlier Margaret H.
Hundley had estimated 150 Roseate Terns in the Tortugas area
(Stevenson, 1956: 327).



1957-1958. The record for these years is almost certainly incom-
plete. No 1957 observations later than mid-May are available, and
in 1958 no particular effort was made to locate the colony. Possibly
colonies of more normal size developed each year.

1959. A hasty count located approximately 223 nests in a dense mat
of Sesuvium on the highest ridge of Hospital Key. Hatching was
about half completed with the largest chicks about one week old.
The party banded 80 chicks.

1960. Observers who visited the Dry Tortugas in early May saw
some Roseate Terns around the east end of Bush Key, but the colony
apparently had not begun nesting (I. Joel Abramson, in litt.). Band-
ers working there 27-31 May frequently saw a few Roseates fishing
in the bight between Bush Key and Long Key, but found no nests.
Severe squalls prevented visits to any of the outlying keys by either
party in May. On 11 July members of a second group of banders
landed on all the keys. No Roseates were nesting at that date, but a
densely massed assemblage of terns and gulls on Middle Key in-
cluded about 100 individuals of some species of white Sterna, many
of which were birds of the year. The behavior and unsteady flight
of these youngsters indicated that they had been reared at Dry Tor-
tugas, although not necessarily on Middle Key. Opportunity to study
the adults was brief, and the observers, aware of the uncertainty sur-
rounding reports of southern nestings of the Common Tern concluded
that the birds were Roseates. The single juvenile netted and banded
on Middle Key was so reported.

1961. Oliver L. Austin, Jr., and William G. Atwater banded 20 well-
grown juveniles on the east spit of Bush Key 16 July.

1962. The Roseate Terns first located on several elongate heaps of
rough coral fragments near the south end of Long Key. A member
of the banding party, Theodore R. Greer, devoted several days to
photographing (figure 9) and observing the colony from a blind. On
27 May he counted 118 nests, 34 containing single eggs and 84 with
two-egg clutches. On 13 June a field excursion group of the 13th
International Ornithological Congress (Robertson, 1962) found the
colony site deserted and broken egg shells remaining in the nest de-
pressions. Slight vascularization of the inner shell membranes indi-
cated that predation had occurred early in incubation, and the way
the shells were broken suggested the work of an avian predator. The

Vol. 8


most likely suspects were the some 20 cattle Egrets, Bubulcus ibis,
then frequenting Bush and Garden Keys. Cattle Egrets at the Dry
Tortugas have formed some unusual feeding habits. In May and
June 1962 and May 1963 they were frequently seen to stalk and kill
injured or exhausted spring migrant passerines (mainly Parulids) and
to feed upon small birds already dead.

FIcuirE 9. Roseate Tern incubating, south end of Long Key, 26 May 1962.
The colony occupied several dune-like elevations of coral rnbble. The bird pic-
tured had a darker bill than most of the adults in the colony, but note that lower
mandible is lighter (reddish) at the base. (Photograph by Theodore R. Grcer.)

Some of the Roseates appear to have renested on Hospital Key
in July. C. R. Mason and others found about 50 nests, all with one
egg, there on 16 July. Park Ranger Carl S. Christensen (in litt.)
visited the colony 10 days later and reported that hatching had be-
gun. At that time none of the nests contained more than one egg,
apparently the normal clutch for second nestings of Roseates at the
Dry Tortugas.

1963. On 17 and 18 May members of the banding party counted 73
nests, each containing one or two eggs, and banded 32 adult Roseates
on Hospital Key. On 7 July large young from the May nesting were
congregated on the beaches and an estimated 150 additional adults
had arrived and begun nesting.



Comments on the nesting of the Roseate Tern at the Dry Tortugas
have stressed the isolation of the colony and its erratic fluctuations
in size from year to year. Sprunt (1951a: 14), for example, writes:
"The marked fluctuation of this tern at Tortugas seems without ex-
planation as, indeed, does the very fact of its being there!" Analysis
of the longer record of population now available suggests that much
of the supposed fluctuation results from incomplete data.
The Tortugan Roseate colony has shifted frequently between the
Bush Key-Long Key area opposite Fort Jefferson and the area of
Hospital and Middle Keys, several miles northeast of the Fort. In
some years the entire breeding population has been concentrated on
one key; in other years two or three separate colonies existed (table 6).
Observers are not likely to have overlooked Roseate Terns nesting
on Bush Key or Long Key, but Hospital and Middle Keys are less
easily accessible and neither was visited by the observers who re-
ported for 1935, 1936, 1939, 1940, 1941, and 1956, nor probably for
1938, 1942, 1943, and 1945.


Location of Number of
Date Tcrnery Adults Source

19-31 July
9 JuneI
14 May

"Long Key"

"Bush Key"

"Bush Key"

"in the quarters

200 Bartsch (1919)

200 Bartsch (ms. notes)

200 Bartsch (ns. notes)

"the usual
... .colony"

Bartsch (1932)

19-20 June Bush Key 400 Mason
1935 Longstr
17-19 June Bush Key 400 Doe &
23-25 June Hospital Key 25(0300 Mason
20-25 June Bush Key 314 Mason
21 June Bush Key 214 Taylor
about 3 June Bush Key 20 Robinso


eet (1936a, 1936b)
Russell (1936)

(ms. notes)

(ins. notes)

(1939 ms.)

n (1940)

Vol. 8


TABLE 6 (continued)

Location of Number of
Date Ternery Adults Source

early June, Bush Key ? See comments
2 July Long Key c. 150 Budlong (1942 ms.)
1943 Bush Key c. 150 Budlong (1943 ms.)
18-20 June Bush Key 170 Sprunt (1946a)
23-25 June Long Key 284 Sprunt (1948a)
1947 Bush Key
Hospital Key
15-18 June Long Key 450 Sprunt (1948c)
13 June Bush Key 102 Dilley (1950)
1949 Long Key
(Hospital Key)
1-5 June Bush Key 124 Moore & Dilley (1953)
1950 Hospital Key
31 May-4 June Hospital Key 136 Moore & Dilley (1953)
1951 Long Key
27 May Long Key 388 Moore & Dilley (1953)
1952 Hush Key
1 July Hospital Key 240 DeWeese (1953 ns.)
1953 Middle Key
Bush Key
27 May Hospital Key 370 Moore (1954 ms.)
3 June Bush Key 436 Moore (1955 ms.)
26 May Long Key 204 Robertson (1956 ms.)
1956 Bush Key
19 May Hospital Key 30 DeWcese (1957 ms.)
June Bush Key 15 Ward (in lit,)
13 June Hospital Key 450 0. L. Austin, Jr. et al.
7-8 May (Bush Key) 100-125 Abramson (in litt.)
1960 Middle Key?
7-17 July Bush Key 120 Robertson (1961)
25-28 May Long Key 236 Greer (in litt.)
May and July Hospital Key c. 300 C. R. Mason,
1963 W. B. Robertson, et al.


In addition to low counts resulting from incomplete coverage,
some counts made while the colony was forming surely underestimate
the actual breeding population. Roseate Terns at the Dry Tortugas
are often well behind the Sooties in their breeding schedule, and
nesting dates have varied considerably from year to year. In some
years the full complement of breeding adults assembles by mid-May
and laying begins during the first week of June. In other years
many first nests still have incomplete clutches the last week of June.
The relation between the date and the size of the population re-
corded is well illustrated by the records for 1939 when on 9 June Rob-
inson (1939: 9) estimated about 80 Roseate Terns on the east end of
Bush Key and found 13 nests with eggs. On 21 June in the same
colony Taylor (1939 Ms.) counted 107 nests with eggs. Moore, in
1952 (Moore and Dilley, 1953: 78), counted 93 nests on Bush and
Long Keys 24 May and 194 nests there on 27 May. In late May 1953
Moore (1954 rs.) was able to locate only 9 Roseate Tern nests at the
Dry Tortugas, but by July (DeWeese, 1953 Ms.) 120 nests were pres-
ent on three keys. Checks of this sort are not available for other
years, but the earlier counts clearly have tended to be lower. The
low Roseate Tern populations recorded in 1940, 1950, 1951, and 1957
all derive from counts made in May or the first week of June.
In 1946 a search of all of the Tortugan keys 16-20 June revealed
no Roseate Terns (Sprunt, 1946a: 1,7). The Custodian of Fort Jeffer-
son, however, had reported a few Roseate Terns in the area during
May (Gibbs, 1946 is.). Perhaps none nested at the Dry Tortugas
in 1946, but it may also be that formation of the colony was unusu-
ally retarded that season, or that an early nesting was destroyed by
spring tides or predators. Excepting only 1946, all complete surveys
of the known nesting keys made after mid-June have located breed-
ing aggregations of approximately 150 to 450 adult Roseate Terns.
Thus the Tortugan Roseate colony appears neither particularly erratic
in its breeding nor to undergo numerical fluctuations of unusual mag-
The difficulties of season and location mentioned above may
possibly account for the failure of 19th century ornithologists to find
Roseate Terns at the Tortugas. While it seems likely that Audubon
and others would have investigated all the keys, it is not certain
that any of them did. Scott, and probably also Audubon, Bryant,
and Maynard's assistants were at Dry Tortugas too early in the spring
to find Roseate Terns, assuming that the colony existed and followed
its present seasonal schedule. In addition Bryant's testimony is ren-
dered equivocal by the possibility that he confused Roseate and Com-

Vol. 8


mon Terns. His statement that he saw no Roseate Terns follows his
comment (1859a: 21): "I found the Little Tern and Wilson's Tern
breeding at different localities among the keys and along the shore
of the mainland."
As for the Tortugan colony's reputed isolation, while it is indeed
remote from the remainder of the Roseate Tern's breeding range in
the United States, it is at the periphery of the species' extensive An-
tillean-Caribbean range. In terms of the species' world range, it
is rather the North Atlantic colonies that are unusual. Most other
Roseate Terns nest within about 30 degrees of the equator (Voous,
1960: 151, map 196). The colonies that breed from Virginia to Nova
Scotia and from Britanny to Jutland may be relict as Fisher and Lock-
ley (1954: 87) suggest, but local extirpation cannot be disregarded as
a possible cause of present breeding range discontinuities. In Flor-
ida, for example, no Roseate Tern nesting colonies have been known
since the mid-19th century, except the one at the Dry Tortugas.
Many summer sight records of Roseate Terns in the central Florida
Keys and the recent report of a small colony "on the Vaca Keys"
in which 3 chicks were banded and several nests with single eggs
seen on 11 June 1962 (Pace, 1962) suggest the possible recolonizing
of former breeding range. This colony of about 30 adults was again
active in the summer of 1963 (Christine A. Bonney, personal com-
The egg collection catalogue of the U. S. National Museum lists eggs
of the Least Tern, Sterna albifrons Pallas, taken at Dry Tortugas in
1859 and 1861 (Robbins, in litt.). Little else is known of its nesting
there prior to 1900. Scott (1890: 306) was informed that it occurred
commonly, but saw none during his visit in March-April 1890. By
1900 the Tortugan population of Least Terns had declined greatly.
Thompson (1903: 83-84) found a colony of 30 pairs on Long Key in
1902 and reported that formerly populous colonies on Loggerhead
Key had been dispersed by eggers. Watson (1907: 315-316) reported
50 pairs attempted to nest first on Loggerhead Key and then on Hos-
pital Key, but predation and disturbance prevented both colonies
from producing young and he considered the species "nearing ex-
tinction" at the Dry Tortugas.
With the establishment of the Carnegie Tortugas Laboratory,
A. G. Mayer, the first director, undertook to stop the gathering of
tern eggs and to control the rats that infested several of the keys.
In 1908 he wrote that the success of these efforts could be seen in



the thriving colonies of Least Terns on both ends of Loggerhead Key
and on Bush Key, the latter said to number about 1000 birds (Dutcher,
1908b; Mayer 1908). The revival of the colonies on Loggerhead Key
was short-lived. In 1917 Bartsch (1919: 487) observed that "the per-
sistent efforts of eggers" had finally driven the birds away from the
island. Small nestings were reported on Loggerhead Key in 1932
(Bartsch, 1932: 287) and in 1935-1936 (Russell, 1938 iM.), but there
has been no subsequent indication that the Least Tern might recol-
onize its original Tortugan breeding ground. Its failure to do so at
Loggerhead may be explained in part by the fact that dogs and cats
kept by the lighthouse personnel often have had free range of the
In contrast to the checkered history of the Loggerhead colonies,
Least Terns nesting in the Bush Key-Long Key area maintained a
fairly constant population for many years. Bartsch (1916, 1917,
1919) reported a colony of 200 on Bush Key in 1915, 500 there in
1916, and 500 on Long Key in 1917. On 9 June 1921 there were 400-
500 birds on Long Key and Bartsch noted on 3 June 1924 (.Ms. notes)
that "several colonies" were breeding in the Tortugas. Warden
Charles I. Park (Ms. notes) estimated 500 Least Terns nesting on Long
and Bush Keys in July 1929, and 700 nesting on Bush Key in 1930.
Bartsch (1932: 281) found "the usual breeding colonies" active in
August 1932.
The first Florida Audubon Society Tortugas trip in 1935 reported
200 Least Terns nesting on the east spit of Bush Key (Mason 1936:
18), which suggests the population was somewhat reduced from that
present a few years earlier. The following year Doe and Russell
(1936: 7) estimated this colony to number only 100 birds. In 1937
there were no Least Terns on Bush Key, but about 25 pairs nested
in the Roseate Tern colony on Hospital Key (Russell, 1938 Ms.)
In the next four years Least Terns were reported on the east spit
of Bush Key and adjacent parts of Long Key as follows: In 1938
Mason (1938: 4) found 11 nests. In 1939 Robinson (1939: 9) reported
25 adults, and the following year (1940: 3) "Not more than a dozen
Least Terns in the colony." In 1941 Stimson (in litt.) reported "a
few Least Terns nesting."
In 1946 Sprunt (1951: 15) saw empty nest scrapes on both Bush
and Hospital Keys, but does not mention how many birds were
present. In 1947 he reported (1948a: 30) "less than a dozen" adults
on Bush and several scrapes without eggs on Middle or Hospital Key.
In 1948 he reported (1948c: 13-14) 12 birds on Bush Key, where the
following year Dilley (1950: 68) found a single nest with eggs. In

Vol. 8


1951 Moore and Dilley (1953: 78) saw two birds but found no nest.
Thereafter no Leasts were reported at the Tortugas during the breed-
ing season until July 1963, when about five adults were seen for sev-
eral days around Garden and Long Keys.
As mentioned under Roseate Tern, Hospital Key was not visited
in some years, and some of the visits after 1935 may have been too
early in the season to record Least Terns nesting. The record is
doubtless incomplete, but certainly the species no longer breeds reg-
ularly at the Dry Tortugas.
No clear explanation of the rapid disappearance of the Least
as a breeding species at the Dry Tortugas can be advanced. Be-
tween 1932 and 1937 a stable and long-established population of ap-
proximately 500 breeding adults on Bush Key-Long Key decreased
to a few birds, with no evidence that the colony suffered disturbance
of any sort. The decline of the Least Terns during the years when
the colony of Sooty Terns was becoming established and increasing
on Bush Key suggests the possibility of some relationship between
the two events. Also of possible significance is the great increase of
Least Tern colonies along the adjacent coasts of southern Florida
since the early 1930's. Dredging along the Inland Waterway and
for coastal real-estate developments has created innumerable small,
sheltered islets and bars which provide ideal nesting sites, perhaps
preferable to more exposed islands like the Tortugas.

Until Hallman (1961) reported two nests found "in the midst of the
colony of Least Terns" on a spoil island in St. Joseph's Bay, Gulf
County, in June 1961, observations at Dry Tortugas provided the
only generally accepted evidence of the nesting of the Common Tern
in Florida (Howell, 1932: 263). In fact, the Tortugan colony has been
considered the only one in the entire Gulf of Mexico region (Lowery
and Newman, 1954: 530), although the A.O.U. Check-List (1957: 235)
mentions breeding colonies on the coast of Texas, and Stewart (1962:
485) recently reported a possible nesting on the Gulf coast of Mis-
The few records of breeding at the Tortugas are not altogether
satisfactory. They are documented neither by specimens nor photo-
graphs, and a strong possibility of confusion with the Roseate Tern
exists. As has been noted, the first report of nesting Common Terns
in the area (Bartsch, 1918) proved to be based on a misidentification



of Roseate Terns and was later corrected (Bartsch, 1919). The other
reports are summarized below.
Bartsch (-rs. notes) reported a colony of 75 pairs of Common
Terns on Bush Key 3 June 1924 and noted that the nests contained
"1-4 eggs or newly hatched young." On 13 August 1925 he reported
"quite a colony present" at Bush and Long Keys, and in 1932 (Bartsch,
1932: 281) "the usual breeding colonies" were said to be active. I
have been unable to learn anything more about these observations.
It is to be noted, however, that Bartsch also reported Roseate Terns
at Dry Tortugas on about the same dates in 1924 and 1925.
The remaining records date from the breeding seasons of 1935,
1936, and 1937. In each case Common Terns were reported nesting
with a larger group of Roseate Terns on the east spit of Bush Key in
1935 and 1936 and on Hospital Key in 1937. Available information
suggests some uncertainty in the minds of the observers concerning
identification of the birds as Common Terns and the number of pre-
sumed Common Terns present. The number in 1935 was reported var-
iously as 50 birds (Mason, 1936: 18; Longstreet, 1936a: 33) "about 75
pair" (Longstreet, 1936b: 99), and "100 pair" (Doe, mrs. notes). Long-
street (1936a: 42) commented: "Mr. Mason and I believe that we
found the common tern breeding on Bush Key. However, we did
not collect any birds or eggs."
The report of the Florida Audubon Society's Tortugas trip of
1936 (Doe and Russell, 1936: 7) states: the common terns
showed a marked decrease, only a few pairs being noted." Mason
advised me (in litt.) that only four birds were seen on Bush Key in
1936 and that no nests were located. Russell (1938 Ms.), however,
wrote elsewhere: "... in 1935 and 1936 I estimated the same colony
to contain about 200 birds." The latter statement could pertain to
observations made later in the summer.
Published accounts of the 1937 trip (Longstreet, 1937; Young and
Dickinson, 1937) do not mention the Common Tern, but Mason (in
litt.) saw a few adults that he believed were Common Terns among
the Roseates on Hospital Key. Russell (1938 MS.) states that Common
Terns nested on Hospital Key in 1937 without indicating how large
the colony was. Since 1937, the only reported occurrence at the
Dry Tortugas during the breeding season appears to be two seen on
Middle Key by Mr. and Mrs. John R. DeWeese, 29 May 1955 (Moore,
1955 MS.).
It seems necessary to conclude from the above that breeding of
the Common Tern at the Dry Tortugas is not proved. The downy
young of the Common and Roseate Terns are easily distinguishable

Vol. 8


by anyone familiar with them. Chicks of the presumed Common
Terns at Tortugas were seen in 1924 and probably also in 1935 (Ma-
son, in litt.), but no record that they were compared critically with
Roseate Tern chicks exists. In addition, some of the identifications
of adult Common Terns apparently were based upon the bill color,
which often is unreliable for separating Commons and Roseates.
Greer (in litt.) advised me that no more than 15 or 20 of the adults in
the colony of Roseates on Long Key in May 1962 had entirely dark
bills, the others having at least the basal third of the bill orange-red.
The latter is considered to be the "variant" condition by Peterson
(1947: plate 37), while Pough (1951: 288) states: "Its bill is black ex-
cept for a little red at the base (occasionally more)."
The Common Tern has been reported to nest at a number of New
World localities south of its regular breeding range. Considerable
uncertainty surrounds most of these records, however, because of the
similarity between Common and Roseate Terns, and because band-
ing evidence shows subadult Common Terns often summer in the
tropics. As Voous (1957: 139) notes: "Its nesting in the West Indian
region has been almost as frequently stated as it has been reject-
ed .. ." Bond (1956: 58) gave full credence to none of the numerous
reports of breeding in the Bahamas and elsewhere in the West Indies.
Similar doubt attaches in some degree to most or all of the alleged
nestings on the South Atlantic and Gulf coasts of the United States,
including those at the Dry Tortugas.
Voous (1957: 137-140; 1963) has recently published unquestion-
able proof of the Common Tern nesting at Curacao, Aruba, and
Bonaire in the southern Caribbean. Records that he assembled sug-
gest that the species has a long history there as an irregular breeder
in solitary pairs or small colonies of fewer than 20 adults. It is also
said to breed in the nearby Venezuelan islands of Los Roques and
Las Aves (Phelps and Phelps, 1958: 111). As Bond (1958: 5) states,
this proof of southern nesting compels a re-examination of the earlier
The significance of the proved and reported southern nestings
of the Common Tern is not clear, but most of the records seem to
conform to a pattern-small numbers of birds nesting sporadically,
often in association with larger colonies of other terns, especially
Roseates and Leasts. Band recoveries show that many 1-, 2-, and 3-
year-old Common Terns summer in the Caribbean and elsewhere
south of the species' usual breeding range. I suggest as a provisional
explanation that some individuals in these normally subadult age
groups reach sexual maturity in southern latitudes and are occasiona-


ally stimulated to nest when they become associated with terneries
of other species.
Voous (1960: 128) has commented: "The few recorded breeding
places in the tropics, very limited in extent and altogether isolated
from one another, must be regarded as recent colonizations by birds
left behind after wintering As presently known, however,
southern nestings of the Common Tern seem to fit more closely the
hypothesis that they are anomalous, impermanent, and perhaps re-
lated to the age of the individuals. It is doubtful that they have sig-
nificance as extensions of the species' breeding range.

Audubon found the Royal Tern, Thalasseus maximus (Boddaert),
nesting abundantly at the Dry Tortugas in May 1832. John Krider
(1879: 80), presumably from observations made in the spring of 1848,
writes of it: "Very abundant on Tortugas Island, Florida, and breeds
on the Keys of Florida." In May, probably of 1850, Bryant (1859a:
20-21) visited Northeast Key, Dry Tortugas, where he observed this
species and the Sandwich Tern, Thalasseus sandvicensis (Latham),
breeding "in great numbers." The U. S. National Museum contains
eggs of the Royal Tern collected at Dry Tortugas by Gustavus Wurde-
mann in 1858, and eggs of both species collected by Dr. D. W. White-
hurst and Captain D. P. Woodbury in 1859 (H. G. Deignan, in litt.).
Scott (1890) does not mention the Sandwich Tern, but he saw sizable
flocks of Royal Terns at the Dry Tortugas in early April 1890, and
was told that many remained there to breed.
These brief comments span the entire record of breeding by these
species at the Tortugas, except that a single Royal Tern egg was found
on Middle Key in May 1952 with no further evidence of nesting
(DeWeese, 1952 MS.) In Sprunt's (1962: 84) report of my 7 Novem-
ber 1961 observation of Royal Terns there appeared to be
several times this number nesting on the south end of Long Key,"
nesting is a typographical error for resting. Royal Terns still visit
the Tortugas regularly, sometimes in large numbers. For the Sand-
wich Tern a number of observations exist from the neighborhood of
Key West, including several in summer, but three sight records, two
of them recent, are the only known occurrences at the Dry Tortugas
in this century (Sprunt, 1962: 84).
Northeast Key, mentioned as the site of the nesting colony of
Royal and Sandwich Terns, had washed away by 1875. The narra-
tive of a survey made in that year states (Coast Survey, 1878): "North


Key, Northeast Key, and Southwest Key, as represented on old maps,
have no existence now, not being bare even at low water." Other
islands nearby, such as East Key, had areas apparently suitable for
the species, and it seems doubtful that loss of one key could have
caused loss of the colony. A more likely explanation is that the in-
creasingly persistent egging after about 1880 (Scott, 1890) eventually
extirpated the ternery. Both Thalasseus species nested commonly
at a number of southern Florida localities in the 1800's, but no breed-
ing colony of either species is known to exist in the area today.

The Black Noddy, Anous tenuirostris (Temminck) was first recorded
in the continental United States at Bush Key, Dry Tortugas, 13 July
1960 (Robertson et al., 1961), when one was collected and a second
individual seen. During the summers of 1961, 1962, and 1963 the
species was seen repeatedly on Bush Key. With one exception the
observations have been of single birds, usually perched with Brown
Noddies in the dead tree at the south shore of Bush Key from which
the 1960 specimen was collected. To date at least five different indi-
viduals have been seen, and the species apparently is of more than
casual occurrence there.
The Black Noddy is slightly smaller and darker than the Brown
Noddy, its bill is thinner, and its crown patch is whiter, more sharply
defined, and extends farther back on the nape. Yet the two species
are so similar in general appearance and behavior that one could
easily be overlooked in a congregation of the other unless the ob-
server were expecting or watching for it. Sutherland (1961) describes
how he first spotted the Black Noddy in 1960, while making pro-
longed observations on a group of Brown Noddies in the noddyy
tree" on Bush Key to record their calls. Otherwise the species might
easily have gone undetected, and it is indeed possible that a few
birds may have frequented the Tortugas ternery unnoticed for many
In 1961 the first party of banders saw a Black Noddy daily in the
noddy tree 26-31 May. The bird is clearly recognizable in 16-mm
color movies B. G. Hubbard took 27 May (figure 10). The second
banding group also found one Black Noddy on station in the tree
7, 10, 11, and 15 July. The bird seen in May had an indistinct slash
of lighter brown across the left middle coverts, apparently caused by
worn feathers it had not molted. The bird seen in July lacked this
mark and may have been a different individual. Repeated attempts


by both groups of banders to capture the bird failed. It showed
the extreme tameness characteristic of noddies and tolerated ap-
proach to within a few feet, but was much more agile on the wing
than the Brown Noddies and easily avoided both mist and hand nets.
In 1962 four parties with a combined total of more than 50 ob-
servers searched Bush Key for Black Noddies without success 5-6
May, 11-14 May, 25-28 May, and 13 June. The second banding
party found one at the usual roost on 7 and 11 July, and on 13 July
caught it in a hand net. It was banded (683-12000 on right leg, un-
numbered red plastic on left), weighed (103 grams), measured (wing
arc 218 mm, exposed culmen 42 mm), photographed (figure 12) and
released. Its mouth lining, by Palmer and Reilly's (1956) color stand-
ards, was approximately "scarlet-orange", strikingly different from the
"orange-yellow" of the Brown Noddy's mouth. This individual has
not been reported since. The party saw no more Black Noddies
through 15 July, and I could find none on 2 August. On the eve-
ning of 9 September, however, my wife and I caught a second Black
Noddy in a mist net on the west beach and banded it (683-11999).
The first 1963 banding party saw one Black Noddy near Hospital
Key 17 May, but could find none on Bush Key. On 6 July the second
party found one unbanded bird perched among Brown Noddies at the
north coaling dock on Garden Key and photographed it from a dis-
tance of a few feet. On 9 July two Black Noddies, neither banded,
roosted for several hours at the same place. One of these differed
from all others seen at the Tortugas in having the back of the pileum
dusky rather than white. Presumably it was a younger individual.
A number of interspecific squabbles for roosting space were observed,
in which the larger Brown Noddy was usually dominant.
Thus one or more of at least five individual Black Noddies have
been present at the Tortugas ternery during four successive summers.
Their known extreme dates of occurrence, 17 May-9 September, span
virtually the entire breeding period of terns in the area. Since 1960
we have devoted considerable time, perhaps 50 or more man-hours,
to searching for a possible Black Noddy nest. So far no Black Noddy
has been seen at a nest, and no nests, eggs, or young have been found
that appeared to differ from those of the Brown Noddy.
The Black Noddy of July 1962 was several times observed to leave
its perch in the noddy tree and fly directly into an area of dense
brush near the west end of the key. This behavior was suspiciously
like that of the off-duty member of an incubating or brooding pair,
but minute search of the area-several acres of tightly interwoven

Vol. 8


FrcurE 10. Black Noddy (left) and Brown Noddy, Bush Key, 27 May 1961.
(Photograph enlarged from 16 mm movie by B. G. Hubbard.)


FIGURE 11. Black Noddy (upper left) and Brown Noddies, Bush Key, Dry Tor-
tugas, 7 July 1962. (Photograph by Nagahisa Kuroda.)


old bay cedars growing amid thick beds of Opuntia cactus-was im-
possible in the time available.
The bird we banded some 7 weeks later in September we netted
on the shore near the same area. The bird was coming into the
ternery, and while being handled disgorged a rounded food mass
about 40 mm in diameter, the compacted remains of a large number
of tiny minnows. The Brown Noddies at the time were still feeding
a few large young in or near the nests.
Therefore while we strongly suspect and would like to believe
that the Black Noddy has been nesting at Dry Tortugas, we have
not as yet been able to prove it.

Ftriun 12. Close-up of the head of the first Black Noddy banded at Dry Tor-
tugas, Bush Key, 13 July 1962. (Photograph by James B. Meade.)

Many data of interest found during preparation of this paper have not
been formally published. These are contained in written material of
two general kinds. 1. Completed manuscripts and official reports that
bear a definite date are cited by date with the designation "Ms." to in-
dicate that the source is an unpublished manuscript. 2. Field notes
and other material not in completed form are cited as "Ms. notes"
without date. Copies of most of the manuscripts and manuscript
notes are in my possession. The place of deposit of the completed
manuscript reports and the source of the manuscript notes are indi-
cated in the citations or comments.

Vol. 8


Allen, A. A.
1944. Report of the A.O.U. Committee on bird protection for 1943. Auk,
61: 622-635.

Allen, Robert P.
1936. With the Audubon wardens. Bird-Lore, 38: 295-297.

American Ornithologists' Union.
1957. Check-List of North American Birds. Published by the A.O.U., xiii +
691 pp.
[The statement of the range of the Sooty Tern (p. 237), reading in part:
"Breeds from Alacran Reef off Yucatan, the Gulf Coast of Texas, Louisi-
ana, Florida (the Dry Tortugas, irregularly in recent times) ... "
is in error both as regards recent breeding at Tortugas and the present
existence of established colonies in Louisiana and probably also in Texas.]

Ashe, T. J.
MS. notes. Population estimates of Noddies in 1910 and 1915. [From entries
in the Bird Distribution file, Patuxent Wildlife Research Center, noted
for me by Chandler S. Robbins.]
1919 ms. Warden's report, Bird Key, Dry Tortugas. Original copy in files
of Research Department, National Audubon Society. Holographic,
4 pp. [Dated 6 October 1919. Summary of 1919 nesting season with
population estimates. Hurricane damage to vegetation on Bird Key.]

Ashe, T. J., and Ludwig Bethel
1915 Mis. Warden's annual report. Bird Key, Dry Tortugas. National Audu-
bon Society files, New York. [Prepared by Ashe from information pro-
vided by Assistant Warden Bethel. Estimates 107,000 "terns". Reports
heavy mortality of terns caused by hurricanes of 13 August and 3 Sep-
tember 1915.]

Ashe, T. J., and William E. Lowe
1918 MS. Warden's annual report, Bird Key, Dry Tortugas. Original copy
in files of Research Department, National Audubon Society. Ilolo-
graphic (Ashe), 5 pp. [Written by Ashe from information provided by
Lowe, his assistant in charge of Bird Key. Dated 26 September 1918.
Population estimates of Sooties and Noddies. Summary of the highly
successful 1918 nesting season.]

Ashmole, N. P.
1962. The Black Noddy Anous tenuirostris on Ascension Island. Part 1. Gen-
eral biology. Ibis, 103b: 235-273.
1963a. The biology of the Widcwake or Sooty Tern Sterna fuscata on Ascension
Island. Ibis, 103b: 297-364.
1963b. The regulation of numbers of tropical oceanic birds. Ibis. 103b: 458-

Audubon, John James
1835. Ornithological Biography. Volume 3. xvi + 638 pp.



Austin, Oliver L.
1949. Site tenacity, a behavior trait of the Common Tern (Sterna hirundo
Linn.). Bird-Banding, 20: 1-39.

Austin, Oliver L., Jr.
1962 Nls. Comparative demographics of Sterna hirundo and Sterna fuscata.
Delivered before the 13th International Ornithological Congress (Ithaca,
Bache, A. D.
1858. Report of the Superintendent of the Coast Survey, showing the progress
of the survey during the year 1857. Executive Document No. 33, 35th
Congress, 1st Session. Washington. William D. Harris, Printer.

Bache, Hartman
1845 Ms. Topographical survey of Garden Key. Drawer 74-1, files of U. S.
Corps of Engineers, Military Construction Branch, Construction Divi-
sion, War Department, Washington, D. C. [Map of the island as it
was immediately before construction of Fort Jefferson began. I have
seen no notes that may have accompanied the map, nor correspondence
relating to it.]

Baker, John H.
1944. The director reports to you. Audubon Magazine, 46: 178-183.
Bartsch, Paul
1916. Birds observed on the Florida Keys and along the railroad of the main-
land from Key Largo to Miami, June 17-July 1, 1915. Carnegie Inst.
Washington, Yearbook for 1915 (No. 14): 197-199.
1917. Birds observed in 1916, in the region of Miami and the Florida Keys
from May 15 to June 4, and along the railroad from Key West to Miami
on June 24. Carnegie Inst. Washington, Yearbook for 1916 (No. 15):
1918. Fifth annual list of birds observed on the Florida Keys. Carnegie Inst.
Washington, Yearbook for 1917 (No. 16): 170-173.
1919. The bird rookeries of the Tortugas. Smithsonian Institution, Annual
Report for 1917: 469-500, 38 plates.
Ms. notes. Estimate of Noddy populations in 1922; observations of Common
Terns in 1924 and 1925; observations of Roseate Terns in 1921 through
1925; and observations of Least Terns in 1921 and 1924. [From en-
tries in the Bird Distribution file, Patuxent Wildlife Research Center,
taken from field notes by Bartsch. Compiled for me by Chandler S.
1923. Birds of the Florida Keys. Carnegie Inst. Washington, Yearbook for
1922 (No. 21): 165.
1931. Report on Cerion colonies planted on Florida Keys. Carnegie Inst.
Washington, Yearbook for 1930-1931 (No. 30): 373-378.
1932. The Bird Rookeries of the Tortugas. Carnegie Inst. Washington, Year-
book for 1931-1932 (No. 31): 281.

Vol. 8


1933. Cerion Studies. Carnegie Inst. of Washington, Yearbook for 1932-
1933 (No. 32): 266-267.
Beard, Daniel B.
1938. The 1938 count of Noddy and Sooty Terns at the Dry Tortugas. Flor-
ida Naturalist, 12: 7-10.
1938 Ms. Special report 1938 count of Sooty and Noddy Terns at Fort Jeffer-
son N.M. National Park Service files. Typewritten, 8 pp. [Observa-
tions of 20-25 June 1938. Similar to the published report (Beard,
1938), but includes comments on the advisability of continuing band-
ing terns and more extended comments on the great decrease of Nod-
dies noted in 1938.]
1939. Man-o'-war-birds prey on eastern Sooty Trns. Auk, 56: 327-329.
Bent, Arthur Cleveland
1921. Life Histories of North American gulls and terns. Bull. U.S. Nat. Mus.
113: x + 345 pp., 93 plates.
Bethel, Ludwig
1916. Ms. Warden's annual report, Bird Key, Dry Tortugas. National Au-
dubon Society files, New York.
Bond, James
1956. Check-List of Birds of the West Indies. ix + 214 pp., frontis. The
Academy of Natural Sciences of Philadelphia.
1958. Third Supplement to the Check-list of Birds of the West Indies (1956).
11 pp. Academy of Natural Sciences of Philadelphia.
Bowman, H. II. M.
1918. Botanical ecology of the Dry Tortugas. Carnegie Inst. of Washington,
Papers from the Tortugas Laboratory, 12: 109-138.
Bryant, Henry
1859a. [Birds observed in east Florida, south of St. Augustine]. Proc. Boston
Soc. Nat. Hist., 7: 5-21. (Read 19 January 1859).
1859b. A list of birds seen at the Bahamas, from January 20 to May 14, 1859,
with descriptions of new or little-known species. Proc. Boston Soc.
Nat. Iist. 7: 102-134.
Budlong, Robert R.
1942 Ms. Annual wildlife report Fort Jefferson National Monument. Na-
tional Park Service files. Typewritten 3 pp. [Dated 4 October 1942.
Comments on nesting terns presumably refer to the 1942 nesting season.]
1943 MS. Annual report of the Custodian. Fort Jefferson National Monu-
ment. National Park Service files. [For the fiscal year ending 30
June 1943. Of this report I have seen only the material on nesting
terns that is quoted in letters from Director, National Park Service, to
Secretaries of War and Navy protesting aerial bombing and machine-
gunning practice near Bush Key.]
1944 xs. Custodian's monthly narrative report for May 1944. National Park
Service files. Typewritten, 3 pp.


Chapin, James P., and L. W. Wing
1959. The Wideawake calendar, 1953 to 1958. Auk, 76: 153-158.

Coast Pilot
1889. Atlantic Local Coast Pilot, Sub-Division 22. Straits of Florida, Jupiter
Inlet to the Dry Tortugas. First Edition. U. S. Coast and Geodetic Sur-
vey, GPO, Washington. viii + 91 pp.
1936. United States Coast Pilot, Gulf Coast, Key West to the Rio Grande.
Second Edition. U.S. Dept. of Commerce, Coast and Geodetic Survey.
GPO, Washington vi + 397 pp.

Coast Survey
1878. Report of the Superintendent of the U.S. Coast Survey during the
Year 1875. Executive Document No. 81, House of Representatives,
44th Congress, 1st Session. GPO, Washington.

Cullum, G. W.
1891. Biographical Register of the Officers and Graduates of the U.S. Military
Academy. Vol. 1(1802-1840).

Davis, John H., Jr.
1942. The ecology of the vegetation and topography of the sand keys of
Florida. Carnegie Inst. Washington, Papers from the Tortugas Labora-
tory, 33: 113-195.

Davis, T. Frederick
1935. History of Juan Ponce de Leon's voyages to Florida. Florida Historical
Quarterly, 14: 70 pp.

DeWeese, John R.
1953 Ms. Superintendent's monthly narrative report for June 1953. National
Park Service files. Typewritten, 2 pp. [Count of nests of Roseate
Terns on Middle, Hospital, and Bush Keys.]

Dilley, Williard E.
1949 Msi. Report of the terns of Fort Jefferson National Monument for the
year 1949. Everglades National Park files. Mimeographed, 7 pp.
1950. The terns of the Dry Tortugas, June, 1949. Florida Naturalist, 23:

Diclinson, J. C., Jr.
1941. Noddy and Sooty Terns nesting on bare ground. Auk, 58: 259.

Doe, Charles E.
Ms. notes. Notes relating to the Florida Audubon Society Tortugas trips of
1935, 1936, and 1937 and holographic draft of Doe's article describing
the 1936 trip. [Extracted from field notebooks in the files of the
Florida State Museum by Oliver L. Austin, Jr.]

Doe, Charles E., and Jack C. Russell
1936. Tortugas colonies again visited. Florida Naturalist, 10: 6-8.

Vol. 8


Dunn, Gordon E., and Banner I. Miller
1960. Atlantic Hurricanes. Louisiana State University Press. xx + 326
pp., 87 figures.

Dutcher, William
1903. Report of the A.O.U. committee on the protection of North American
birds. Auk, 20: 101-159.
1904. Report of the A.O.U. committee on the protection of North American
birds for the year 1903. Auk, 21: 97-208.
1905. Annual report of the National Association of Audubon Societies for
1905. State reports, Florida. Bird-Lore, 7: 314.
1906. Bird Key, Tortugas, Florida. Bird-Lore, 8: 146.
1908a. The Tortugas Reservation. Bird-Lore, 10: 142-143.
1908b. Reservation news, Tortugas, Florida. Bird-Lore, 10: 187.

England, George Allan
1928. Bird Key. Saturday Evening Post, 201: 14-15, 85-86, 88, 11 photo-
graphs of terns.

Felton, James B.
1940 Ms. Custodian's monthly narrative report for January 1940. National
Park Service Files. Typewritten, 3 pp.
1941 is. Custodian's monthly report for July 1941. Typewritten, 4 pp.
National Park Service files. [Brief comments on the pattern of landing
of Sooty Terns on Bush Key.]

Fisher, James, and R. M. Iocklley
1954. Sea-Birds. Houghton Mifflin Company. Boston. xvi + 320 pp., 8
color plates, 40 plates, 55 figures.

Gauld, George
1790. An accurate Chart of the Tortugas and Florida Keys or Martyrs, Sur-
veyed by George Gauld, A.M. in the Years 1773, 4, & 5, By Order
Of The Right Ilonourable The Lords Commissioners Of The Admiralty,
And now Published by permission of Their Lordships. London: Pub-
lished by Win. Faden, Geographer of the King, Charing Cross, 5th
April, 1790. Second Edition: 1820. Corrected at Hydrographic Office.
Admiralty to 1835. [The copy seen was printed much later. It in-
cludes an inset map "Dry Tortugas Anchorages of Garden Key (Bush
Kay)" taken from the U.S. Coast Survey of 1854, and gives compass
variation for 1874.]
1796. Observations / on the / Florida Kays, Reef and Gulf / with / Direc-
tions for Sailing Along the Kays, / from Jamaica by the Grand Cay-
man and the West End of Cuba: / Also / A Description, with Sailing
Instructions, / of the / Coast of West Florida, / Between the Bay of
Spirit Santo and Cape Sable. / By George Gauld, / To accompany
his charts of those coasts, surveyed and published by Order of the /
Right Ionourable the Lords Commissioners of Admiralty. / .
Entered at Stationers Hall / London: / Printed for W. Faden, Geogra-


pher To His Majesty, and To His Royal / Highness The Prince of
Wales. / 1796. 28 pp.
Gibbs, Russell A.
1946 MS. Custodian's monthly narrative report for May 1946. National Park
Service files. Typewritten, 4 pp. [As of 30 May "some Roseate Terns
had arrived."]
1947 Mis. Custodian's monthly report for April, 1947. Typewritten, 3 pp.
National Park Service files. [Brief discussion of the landing of Sooty
Terns on Garden Key in 1947.]
Hallman, Roy C.
1961. Common Tern (Sterna hirundo) nesting in northwest Florida. Florida
Naturalist, 34: 221-222.
Heernmann, Adolphus L.
1852-1853. Catalogue of the zoological collection in the Academy of Natural
Sciences of Philadelphia. Proc. Academy Nat. Sci. Philadelphia,
6: 1-35.
Hogan, J.
1925. Bird notes from Willis Island. Emu, 24: 266-275.
Holden [sic], J. B.
1868. The Dry Tortugas. Harper's New Monthly Magazine, 37: 260-267.
Holder, Charles F.
1892. Along the Florida Reef. D. Appleton and Company, New York. ix +
273 pp., many text figures.
Howell, Arthur H.
1932. Florida Bird Life. Published by Florida Dept. of Game and Fresh
Water Fish and Bureau of Biological Survey, U. S. D. A. Coward-
McCann, Inc., New York. xxiv + 579 pp., 58 plates, 72 text figures.
Job, Herbert Keightley
1905. Wild Wings. Houghton Mifflin and Co. Boston. xxiv + 341 pp.
160 photos.
Krider, John
1879. Forty Years Notes of a Field Ornithologist. Philadelphia, xi + 84 pp.
Lashley, K. S.
1915. Notes on the nesting activities of the Noddy and Sooty Terns. Carnegie
Institution of Washington, Papers from the Tortugas Laboratory, 7:
Longley, William II.
1927. Life on a Coral Reef. National Geographic Magazine, 51: 61-83.
Lowe, William E.
1917. MS. Warden's annual report, Bird Key, Dry Tortugas. National Au-
dubon Society files, New York.

Vol. 8


Longstreet, R. J.
1936a. The Florida Audubon Society Tortugas expedition of 1935. Part II,
Part III. Florida Naturalist, 9: 22-42.
1936b. Notes from the Dry Tortugas. Auk, 53: 99-100.
1937. Bird population of the Dry Tortugas. Florida Naturalist, 11: 7-10.
Lowery, George H., Jr., and Robert J. Newman
1954. The birds of the Gulf of Mexico, pp. 519-540. In "Gulf of Mexico
Its Origin, Waters, and Marine Life," Fishery Bulletin 89, U.S. Fish
and Wildlife Service.
Manucy, Albert
1943. The Gibraltar of the Gulf of Mexico. Florida Historical Quarterly, 21:
1961 Ms. A constructional history of Fort Jefferson 1846-1874. National
Park Service files. Typewritten, 242 pp. [Documented report based
upon search of the original records in archives, letters, and other basic
sources. Construction activities, 1846-65, are treated in detail with
a brief resume of concurrent general history of the area. Construction
through 1874 is reported in less detail. Brief comment on later events
that affected the structures to 1935.]
Mason, C. Russell
ms. notes. Field notes relating to the Florida Audubon Society trips of 1935
through 1938.
1936. The Florida Audubon Society Tortugas expedition of 1935. Part I.
Florida Naturalist, 9: 17-22.
1938. Society conducts fourth Tortugas trip. Florida Naturalist, 12: 1-6.
Mayer, Alfred Goldsborough
1908. Tortugas Reservation. Bird-Lore, 10: 229.
Maynard, C. J.
1881. Birds of Eastern North America. iv + 532 pp., 32 plates.
Millspaugh, Charles Frederick
1907. Flora of the sand keys of Florida. Field Columbian Museum, Botanical
Series 2 [Publication 118]: 191-243.
Moore, Joseph C.
1954 Ms. 1953 and 1954 censuses of tern colonies of Fort Jefferson National
Monument. Everglades National Park files. Typewritten, 4 pp. [Sum-
marizes censuses of Sooties in 1952, 1953, and 1954 based upon counts
of nests in 20 or 30 sample areas each of 8 sq. yds. Censuses of Nod-
dies and Roseates by direct count of nests. Difficulties of consusing
because of the prolonged arrival period of Sootics.]
1955 .ls. Tern census of Fort Jefferson National Monuments for 1955. Ever-
glades National Park files. Typewritten, 4 pp. [Dated 7 June 1955.
Estimates 71,102 Sootics, lowest since 1947. Health of colony poor,
unusual numbers of dead adults and young seen. Recorded 1218 Nod-
dies and 436 Roseates both by direct count of nests.]


Ms. notes. [Field notes on his observations at Dry Tortugas in 1952, 1954, and
1955. Other field notes and calculations and several vegetation maps
of Bush Key, all pertaining to the early 1950's.]

Moore, Joseph C., and Williard E. Dilley
1953. Tern counts at Dry Tortugas 1950, 1951, and 1952. Florida Natural-
ist, 26: 75-81.

Murphy, Robert Cushman
1936. Oceanic Birds of South America. Volume 2. The Macmillan Company,
The American Museum of Natural History, New York. xii + pp. 641-
1245, text figures 62-80, plates 39-72, 10 unnumbered colored plates.

Pace, Jerry F.
1962. Banding Least Terns in Florida. EBBA News, 25: 191-193.

Palmer, Ralph S., and E. M. Reilly, Jr.
1956. A concise color standard. 8 pp. Published by American Ornithologists'
Union Handbook Fun.

Park, Charles I.
rMS. notes. Populations of Noddies in 1929 and of Least Terns in 1929 and
1930. [From entries in the Bird Distribution file, Patuxent Wildlife
Research Center, apparently based upon Park's warden reports to the
Biological Survey. Compiled for me by Chandler S. Robbins.]

Peacon, John
1909 s, Warden's annual report, the islands of the Tortugas Group (Bird
Key). National Audubon Society files, New York. [Attached to the
report are a sketch map of the islands of the Tortugas and copies of
correspondence between Peacon and T. J. Ashe.]
1911 xis. Warden's annual report, Bird Key, Dry Tortugas. National Audu-
bon Society files, New York.
1912 Mis. Warden's annual report, Bird Key Dry Tortugas. National Audu-
bon Society files, New York.
1913 iMS. Warden's annual report, Bird Key, Dry Tortugas, National Audu-
bon Society files, New York.
1914 Mis. Warden's annual report, Bird Key, Dry Tortugas. National Audu-
bon Society files, New York. [All information about the reports of
John Peacon is from notes made for me by Robert P. Allen. I have
seen none of the original reports.]

Pearson, T. Gilbert
1915. Motion-pictures for the National Association. Bird-Lore, 17: 410-412.
1915 rs. Dry Tortugas, Florida. Typewritten, 5 pp. Files of Research
Department, National Audubon Society. [A long account of the con-
dition of the colony in 1915 mainly as observed by Rev. H. K. Job,
Dr. H. R. Mills, and Capt. T. J. Ashe who visited Bird Key 28 May
1915. Quotes extensively from Job's report of this trip and from the
annual report of Ashe. Mills used an area-density technique to esti-

Vol. 8


mate the population of Sooties. Ashe reported that terns suffered
heavy mortality as a result of the hurricanes of 13 August and 5
September 1915. This ms. appears to he copy prepared by Pearson
for publication in Bird-Lore, but only a brief excerpt from it was pub-
1917. Report of T. Gilbert Pearson, Secretary Audubon Warden Work. Bird-
Lore, 19: 397-402.
1918. Report of T. Gilbert Pearson, Secretary Audubon Warden Work. Bird-
Lore 20: 459-460.
1919. Audubon warden work in Fifteenth Annual report of the National
Association of Audubon Societies. Report of the Secretary. Bird-Lore,
21: 401-404.

Peterson, Roger Tory
1947. A Field Guide to the Birds. Houghton Mifflin Co., Boston. xxiv +
290 pp., 60 plates, numerous line drawings.
1950. Birds Over America. Dodd, Meade and Company, New York. xiii +
342 pp., 105 photographs.

Peterson, Roger Tory, and James Fisher
1955. Wild America. Houghton Mifflin Company, Boston. xii + 434 pages.

Phelps, William 11., and William 11. Phelps, Jr.
1958. Lista de las aves de Venezuela con su distribution. Tomo II, Parte 1,
No Passcrifonnes. Bol. Soc. Venezolana de Ciencias Nat., 90: 1-317.

Pough, Richard H.
1951. Audubon Water Bird Guide. Doubleday and Co., New York. xxviii
+ 352 pp., 48 plates, numerous line drawings.

Rea, Margaret P., Jennie Lynne Kyle, and Louis A. Stimson
1941. The 1941 Florida Audubon Society expedition to the Dry Tortugas.
Florida Naturalist, 15: 16-17.

Richardson, Frank, and Harvey I. Fisher
1950. Birds of Moku Manu and Manana Islands off Oahu, Hawaii. Auk,
67: 285-306.

Ridley, M. W., and Lord R. C. Percy
1958. The exploitation of sea birds in the Seychelles, Colonial Research Studies
No. 25, pp. 1-78.

Robert, Henry, and Henry M. Stevenson
1951. Notes on birds on the Dry Tortugas. Florida Naturalist, 24: 100-105,

Robertson, William B., Jr.
1956 MS. 1956 tern count. Everglades National Park files. Typewritten, 4
1962. Florida Region. Audubon Field Notes 16: 468-473.



Robertson, William B., Jr., Dennis R. Paulson, and C. Russell Mason
1961. A tern new to the United States. Auk, 78: 423-425.
Robinson, George Digby
1939. 1939 Tortugas Expedition First Group-June 5-10. Florida Nat-
uralist, 13: 7-12.
1940. 1940 Tortugas expedition. Florida Naturalist, 14: 1-6.
Russell, Jack C.
1938 Ms. Narrative report on wildlife of Fort Jefferson National Monument.
National Park Service files. Typewritten, 39 pp. [Dated 24 February
1938. Report of observations made during the author's employment
as a student wildlife technician, 10 June-15 September 1937, with
notes from earlier visits in 1935 and 1936. The first 24 pages deal
chiefly with birds; the remainder with marine life, principally mollusks.
Discusses all nesting terns in detail.]
Scott, W. E. D.
1890. On birds observed at the Dry Tortugas, Florida, during parts of March
and April, 1890. Auk, 7: 301-314.
1904. The Story of a Bird Lover, xi + 372 pp. Macmillan, New York.
Sprunt, Alexander, Jr.
1946a. Survey of the tern colonies of the Dry Tortugas season of 1945. Flor-
ida Naturalist, 19: 26-34.
1946b. Population survey of the tern colonies of the Dry Tortugas Fort Jeffer-
son National Monument 1946. Florida Naturalist, 20: 1-8.
1947a, 1948a. Population survey, tern colonies of the Dry Tortugas, Fort
Jefferson National Monunent, 1947. Florida Naturalist, 21[Oct., 1947]:
16-21; 21[Jan., 1948]: 25-31.
1947b. Blizzard of birds: The Tortugas terns. National Geographic Magazine,
91: 213-230.
1948b. The tern colonies of the Dry Tortugas Keys. Auk, 65: 1-19.
1948c. Tern colonies of the Dry Tortugas, Fort Jefferson National Monument-
1948. Florida Naturalist, 22: 9-16.
1949. Status of Roseate Tern as a breeding species in southern United States.
Auk, 66: 206-207.
1951. A list of the birds of the Dry Tortugas Keys 1857-1951. Florida Au-
dubon Society, 27 pp., 5 photographs.
1962. Birds of the Dry Tortugas 1857-1961. Florida Naturalist, 35: 34-40,
58, 82-85, 129-132.
Stevenson, Henry M.
1956. Florida Region. Audubon Field Notes, 10: 325-329.
Stevenson, James O.
1938. The tern colonies of Dry Tortugas. Bird-Lore, 40: 305-309.
Stewart, James, Jr.
1962. Central Southern Region. Audubon Field Notes, 16: 480-481, 485-486.

Vol. 8


Sutherland, Charles Alan
1961. Excursion to an avian island. Florida Naturalist, 34: 7-10.

Tannehill, Ivan Ray
1950. Hurricanes (7th Edition). Princeton University Press, Princeton. x +
804 pp., 135 figures.

Taylor, O. B.
1939 Ats. Special report on wildlife Fort Jefferson National Monument. Na-
tional Park Service files. Typewritten, 8 pp. [Based on a visit of
19-21 June 1939. Brief comments on nesting terns and sea turtles
with a discussion of the measures needed to improve protection of wild-
life in the area.]

Thompson, Joseph
1903. The Tortugas tern colony. Bird-Lore, 5: 77-84.
United States Coast and Geodetic Survey
1896. Tortugas Harbor and Approaches. USCGS 471a. Scale 1:40,000.
1958. Dry Tortugas. CGS 585. Scale 1:30,000.

Vaughan, Thomas Wayland
1914. The building of the Marquesas and Tortugas atolls and a sketch of
the geologic history of the Florida reef tract. Carnegie Inst. of Wash-
ington, Papers from the Tortugas Laboratory, 5: 55-67.

Vinten, C. R.
1943. The Noddy and Sooty Tern colonies of the Dry Tortugas Fort Jefferson
National Monument: A summary of the records. Florida Naturalist,
16: 53-61.
1944. Tortugas Note. Florida Naturalist, 17: 71-72.

Voous, K. H.
1957. The birds of Aruba, Curacao, and Bonaire. Studies on the Fauna of
Curacao and other Caribbean Islands, 8: 1-260.
1960. Atlas of European Birds. Nelson, London. 284 pp.
1963. Tern colonies in Aruba, Curacao and Bonaire, south Caribbean Sea.
Proc. XIII Intern. Ornithol. Congr.: 1214-1216.

Warham, John
1956. Observations on the birds of Pelsart Island. Emu, 56: 83-93.

Watson, John B.
1907. Report of John B. Watson on the condition of the Noddy and Sooty
Tern colony on Bird Key, Tortugas, Florida. Bird-Lore, 9: 307-316.
1908. The behavior of Noddy and Sooty Terns. Carnegie Inst. Washington,
Papers from the Tortugas Laboratory, 2: 187-255.
1910. Further data on the homing sense of Noddy and Sooty Terns. Science,
82 (n.s.): 470-473.


1912 ,s. Special report Tortugas Reservation. National Audubon Society
files, New York. [States that he visited Bird Key too late in the season
to make accurate population estimates (Carnegie Inst. Washington,
Yearbook for 1912 (No. 11): 125, however, gives the dates of Watson's
1912 work in the area as 6 June-12 July). Recommends planting of
"large quantities" of bay cedar on Bird Key to repair damage done
to vegetation by the hurricane of 1910. From notes by Robert P.
Allen. I have not seen the original.]
Watson, John B., and K. S. Lashley
1915. An historical and experimental study of homing. Carnegie Inst. Wash-
ington, Papers from the Tortugas Laboratory, 7: 7-60.

Worth, C. Brooke
1940. Egg volumes and incubation periods. Auk, 57: 44-60.

Wurdemann, Gustavus
1861. Letter Relative to the Obtaining of Specimens of Flamingoes and Other
Birds from Southern Florida. By the late Gustavus Wurdemann. In-
dian Key, Florida. August 27, 1857. Annual Report Smithsonian In-
stitution for 1860 (1861): 426-430.
Young, T. R., Jr., and J. C. Dickinson, Jr.
1937. 1937 Tortugas expedition. Florida Naturalist, 11: 1-6.


Part 1 (Archaeopterygiformes through Ardeiformes)

SYNOPSIS: The first installment of a catalogue of the fossil birds of the world
covers 49 families in 15 orders of birds, or nearly half of the orders and about
one-fourth. of the families known. The species treated number 374, of which
273 are extinct, and 101 represent living species recorded from fossil or
prehistoric sites. For the palcospecics the data include citation of the original
description, synonyms, nature and repository of types, reference to pertinent
revisionary papers, and detailed geological and geographic ranges, with biblio-
graphic reference to their occurrence.
Major taxonomic changes include recognition of three subclasses, Sauriurae for
Archaeopteryx, Odontoholcae for the Hesperornithidae, and Ornithurae for the
remaining birds. Three infraclasses of Ornithurae are recognized, Dromacogna-
thae (for the Tinamidae), Ratitae, and Carinatae.
Changes in position include transfer of the family Opisthodactylidae to the
Rheiformes, Enaliorithidae to the Gaviiformes, and Baptornithidac to the
On priority the ordinal name Ciconiiformes yields to Ardeiformes. Prior family
names adopted include Emeidae for Anomalopterygidac, Occanitidae for Hydro-
hatidae, and Plataleidae for Threskiomithidae.
New taxa proposed are Colymboidinae (new subfamily, Gaviidae), Caye-
tanornis (new genus, Tinamidae), and Palaeeudyptes marplesi (new species,
Spheniscidae). The misprinted name Pelagodornithidae is emended to Plega-
dornithidae, to conform with the spelling of the type-genus.

'The author is Professor of Biological Sciences at the University of Florida,
Gainesville. Manuscript received 28 January 1963.-ED.



INTRODUCTION ............................................... 186
CLASS AVES Linnaeus .......................................... 189
SUBCLASS tSAURIURAE Hacckcl ............................... 189
tARcHAEOPTERYnIFORMES Fiirbringer .......................... 189
tArchaeopterygidae Huxley .............................. 189
tArchaeopteryx Meyer ............................... 189
SUBCLASS TODONTOHOLCAE Stejneger ........................ 191
THEsPEROBNITmFlrOnES (Fiirbringer) .......................... 191
tHesperornithidae Marsh ................................. 191
tHesperornis M arsh ................................. 191
tConiornis Marsh ................................... 192
SUBCLASS ORNITHURAE Haeckel ............................. 193
INFRACLASS DROMAEOGNATHAE Huxley .................... 193
TINAMIFOHMES (Huxley) ..................................... 193
Tinamidac Gray ........................................ 193
tTinamisornis Rovercto .............................. 193
tCayetanornis Brodkorb .............................. 194
tQuerandiornis Rusconi .............................. 194
Nothura W agler ................................... 194
Neospecies of Tinamidae ............................... 194
INFRACLASS RATITAE Merrem ............................... 196
STRUTHIONIFOUMES (Latham) ................................ 196
tEleutherornithidae Wetmore ............................. 196
tEleutherornis Schaub ............................... 196
Struthionidan Vigors ..................................... 196
Struthio Linnaeus .................................. 196
Neospecies of Struthionidae ............................ 199
RHEIFOBMES (Forbes) ........... ........................... 200
tOpisthodactylidae Amcghino ............................ 200
tOpisthodactylus Amegbino .......................... 200
Rheidae (Bonaparte) .................................... 200
tHeterorhea Rovereto ................................ 200
Rhea Brisson ............... ....................... 200
Pterocnemia Gray ................................. 201
Neospecies of Rheidae ................................. 201
CASUARIFORMES (Sclater) ................................... 202
Casuariidae Kaup ....................................... 202
Casuarius Brisson ................................... 202
Dromiceiidae W ctmore ................................ 202
Dromiceius Vieillot .................................. 202
Neospecies of Dromicriidae ............................ 203
tDromornithidae Fiirbringer .............................. 203
tDromornis Owen .................................. 203
tGenyornis Stirling and Zietz ......................... 203
tAEPYORNITHIFORME8 (Newton) ............................. 205
tAepyornithidae (Bonaparte) ............................. 205
tEremopezinae Lambrecht ............................. 205
tEremopezus Andrews ................ ............ 205


tAepyornithinae (Bonaparte) ........................... 205
tStromeria Lambrecht ............................... 205
tMullerornis Milne-Edwards and Grandidier ............. 206
tAepyornis Geoffroy ................................ 206
tDINORNITHIFORMES (Gadow) ................................ 208
tEmeidae (Bonaparte) .................................. 208
tAnomalopteryginae (Oliver) ........................... 208
tAnomalopteryx Reichenbach ......................... 208
tM egalapteryx Haast ................................ 209
tPachyornis Lydekker ............................... 210
tEmcinae Bonaparte .................................. 212
tEmeus Reichenbach .............................. 212
tEuryapteryx Haast ................................. 213
tZelornis Oliver .................................... 214
tDinornithidae Bonaparte ................................ 215
tDinornis Owen .................................... 215
APTERYGIFORMES (Haeckel) .................................. 219
Apterygidae (Gray) ..................................... 219
tPseudapteryx Lydekker ............................. 219
Neospecies of Apterygidae .............................. 219
INFRACLASS CARINATAE Merrem ........................... 220
GAVIIFORMES Wetmore and W. D. Miller ....................... 220
tEnaliornithidae Fiirbringer .............................. 220
tEnaliornis Seeley ............... ................. 220
tLonchodytidae Brodkorb ............................... 221
tLonchodytes Brodkorb .............. ................ 221
Gaviidae Allen ......................................... 222
tColymiboidinae Brodkorb .............................. 222
tEupterornis Lemoine ............................... 222
tColymboides Milne-Edwards ......................... 222
tGaviellinae W etmore ................................. 223
tGaviella W etmore .................................. 223
Gaviinae (Allen) ..................................... 223
Gavia Forstcr ...................................... 223
Neospecies of Gaviinae ................................ 224
PormciPEDVIoHMEs (Fiirbringer) ............................... 226
tBaptornithidac American Ornithologists' Union ............. 226
tBaptornis Marsh ................................... 226
tNeogaeonis Lambrcclit ............................. 226
Podicipedidac (Bonaparte) ............................... 226
Podiceps Latham .................................. 227
tPliodytes Brodkorb ................................. 228
Neospecies of Podicipedidae ............................ 228
SPHENISCIFOUMES Sharpe ..................................... 231
Spheniscidae Bonaparte .................................. 231
tPalaeeudyptinae Simpson ................ ........... 231
tPalueeudyptes Huxley .............................. 231
tPachydyptes Oliver ................................. 232
tArchaeospheniscus Marples .......................... 232
tDuntroonornis Marples ............. ............ 232


tPlatydyptes Marples ................................ 233
tKorora Marples ................................... 233
tAnthropodytes Simpson ............. .. ............. 234
tNotodyptes Marples ................................ 234
tAnthropornis Wiman ............................... 234
torthopteryx Wiman ........... .................... 234
tEosphaeniscus Wiman .............................. 235
tDelphinornis Wiman ............................... 235
tlchtyopteryx Wiman .................. .............. 235
tArthrodytes Ameghino .............................. 235
fPalaeosphcniscinae Simpson ........................... 236
tPalaeospheniscus Moreno and Mercerat ................ 236
tPerispheniscus Ameghino ............................ 237
tParaspheniscus Ameghino ........................... 238
tParaptenodytinae Simpson ............................. 238
tParaptenodytes Ameghino ........................... 238
tlsottenwrnis Ameghino ............................. 239
tPseudospheniscus Ameghino ......................... 239
tNeculus Ameghino ........... .................... 240
Spheniscinae Bonaparte ................................ 240
Neospecies of Spheniscinae ............................. 240
PROCELARtFORMEs Fiirbringer ................... ............ 241
Diomedeidae (Gray) .................................... 241
tGigantornis Andrews ............................... 241
tManu Marples .................................... 241
Dionedea Linnaeus ................................ 241
Neospecies of Diomedeidae ............................ 242
Procellariidae (Boie) ............... ..................... 242
Puffinus Brisson ................................... 242
tArgyrodyptes Ameghino ............................ 245
tPlotornis Milne-Edwards ............................ 245
Neospecies of Procellariidae ............................ 245
Oceanitidae (Salvin) ................................... 246
Oceanodroma Reichcnbach .......................... 247
Neospecies of Oceanitidae .............................. 247
Pelecanoididae (Gray) ................................... 247
Neospecies of Pelecanoididae ............................ 247
PELECANIFORMES Sharpe .................................. .. 248
SULAE Sharpe ............................................ 248
tElopterygidae Lamhrecht ............................... 248
tElopteryx Andrews ................................ 248
\Argillornis Owen .................. .............. 248
tEostega Lannhrecht ................................. 249
Phalacrocoracidae (Bonapartc) ............................ 249
tGraculavinae Fiirbringer .............................. 249
tGraculavus Marsh ................................. 249
Phalacrocoracinae Bonaparte ................... ......... 250
tActiornis Lydekker ................................. 250
Phalacrocorax Brisson ..... ......................... 250
tPliocarbo Tugarinov ................................ 254