'White-Faced' Terns
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164 PAL:•XER,'White.faced' Terns I.[Auk April 'WHITE-FACED' TERNS BY RALPH S. PALMER Plate 7 DxvvEt•sTplumages of the samespecies of ternshave been given new namesso many times that the Marpies have included in their bookon 'SeaTerns' (1934:opp. p. 23) a plate labelled"Comparative Diagram of the 'Spurious'Terns." One of the birds shownis the non-breeding'white-faced' Arctic Tern, Sterna 'portlandica.' While conductinga behaviorstudy of the CommonTern (Sternahitundo) on the SugarloafIslands, at Popham Beach, Maine, the writer has collectedand observedboth breedingand non-breeding'white-faced' examplesof this species.Since the significanceand occurrenceof 'white-faced'plumages in ternsare not generallyunderstood, the writer has undertakento sum up here the available evidenceon plumage sequencein the CommonTern. While most of the factspresented refer specificallyto the CommonTern, they apply equallywell to the Arcticand, in manyrespects, to the Roseateand other terns. The writer wishesto acknowledgethe assistancehe has received from the following institutionsand individuals: the authorities in chargeof the U.S. NationalMuseum and BiologicalSurvey collections, the authoritiesof the American Museum of Natural History, and Mr. Arthur H. Norton of the Portland (Maine) Societyof Natural History for the loan of valuablespecimens. Professor S. A. Asdell,of the Departmentof Animal Breedingat Cornell University,has kindly loanedthe writer many reprintsfrom his filesand has made helpful suggestionspertaining to the endocrineaspect of plumagesequence. EXPLANATION OF PLATE 7 Contmon Terns--The left-hand bird is a juvenal and can be so recognizedby the markingson the back. In other respectsit is much like the next two speci- mens,which are summer non-breedersin fresh eclipse,i.e., 'portlandica,' plumage. The two right-handbirds are old-agebreeders. The nature of the cap and the color of the basesof the bills at oncedifferentiate them from 'portlandica' specimens. Arctic Terns--The left-hand bird is the type of Sterna portlandica Ridgway. It is badly soiled and does not show the cubital wing band very well in the plate. The next two are also summer-eclipseor 'portlandica' birds. The fourth is an old-age breeder of the 'pikei' type. The fifth may possiblyrepresent either a partial assumptionof breeding plumage by an 'immature' bird or a condition approachingold age. THE AuIq VOL. 58 PLATE 7 JUVENAI.AND '•¾}IITE-FACEI)' COhll•ION t'•¾HITE-FACEI)' fiiRCTIC TERNS Vol.1941,58] a PALMER,'Whiteffaced' Terns 165 RELATION OF GONADS TO BREEDING AND F.CLIPSE PLUMAGES As here used,the term 'eclipse'includes all postjuvenalplumages assumedby certain'hooded' or 'capped'Laridae when the gonadsare quiescentor relativelyso. The juvenal stagemight alsobe called eclipse,but it is not socalled because this plumage is easilydistinguish- able from subsequentnon-breeding plumages. This definition of eclipsepertains to the speciesunder discussion, since the physioloõical basisfor arrivingat equivalentplumages in ducksand otherbirds is knownto differfrom that of gullsand terns. Both sexes of the Common, Arctic and Roseate Terns exhibit the followingcharacters in thejuvenal and in the eclipsestates: (1) a light foreheadand a blackishcrown; (2) purewhite under parts; (3) a dusky cubitalband on eachwing; (4) lesshighly colored bills and feet than in breedingplumage. Both sexesof the samethree specieshave the followingcharacters in breedingplumage (with an exceptionwhich will be discussedlater): (I) a black cap,which includesboth foreheadand crown; (2) under partseither •ray or white tinted with pink; (3) no duskycubital band; (4) relativelyhighly coloredbills and feet (in breedingRoseate Terns the bill at the base is bright orange-redand not all black, as is commonlybelieved). The juvenal and eclipseplumaões are assumedduring the time when the gonadsare relatively quiescent. The breeding plumage in at leastpart of the Laridae is apparentlythe one dependentupon the secretionof a sex hormonefor its development. Whereasthis is not the casein many birds, as has been explainedadequately by Domm (1939),I feel certain it is true with the ternswith which we are deal- ing becauseof evidencefrom castrationexperiments on the Black- headed Gull, Larus ridibundus (Van Oordt and Junge, 1930, 1933a, 1933b). This gull hasa plumagesequence and certainsecondary sexual charactersvery similar to thosein Common,Arctic and RoseateTerns. It has a 'hood' on the head, as well as bright-coloredbill and feet, which are presentduring the breedingseason in both sexes. Char- acters of this nature have been called 'ambosexual'characters by Champy (1930)and Champyet al. (1931). If gulls of this species are castratedin winter plumage-eitherfirst winter or succeedingones- the nuptial plumagefails to develop. At the time of the prenuptial molt the birdsagain assume the winter (eclipse)plumage. Further- more, the bill and feet remain in winter condition. Therefore, the assumptionof the nuptial plumage and bright bill and foot color appear to be influencedby a testishormone which cannotbe secreted by castratedbirds. 166 PALMER,'White.laced' Terns L[Auk April While Van Oordt workedon male birds only, a matter commented on by Domm (1939:283), he statedthat he supposedthat ovariectomy would producethe sameresult, sincefemales and maleshave similar plumages. The fact that Common and Arctic Terns of both sexes, that have very small gonadsand do not breed, revert to the eclipse plumageat the time of the springmolt, lendssupport to his statement. As we shall soonsee, Common Terns ordinarily do not breed at an early age, nor do first-yearbirds appear on the breedinggrounds in any numbers. Those which do appear usually have minute non- breedinggonads and are found to be in fresheclipse plumage which they acquiredby a completespring molt. That 'portlandica'birds were non-breedersthat acquiredsuch a plumagein spring was well BREEDING ECLIPSE NATAL Sum. Fall Win. Spr. •u•. Fall Win. SpF. •um. F&11 Win. Spl'. Sum. TEXT-FIe. 1.--Diagrammaticrepresentation of the sequenceof plumagesin early life in the CommonTern. The probablenormal sequenceis shownby the heaviest line. Dotted spacesindicate, in a general way, times of molt. understoodby Dwight (1901: 52). These birds are, in plumage, equivalentto Van Oordt'scastrated gulls. Just how long a tern may revert to the eclipsestage semi-annually is not definitelyknown, but it is fair to assumethat they do so aslong as they stayaway from their breedinggrounds. The matter can only be definitely settled when many banded non-breedingbirds are re- coveredat various times of the year. In general, the sequenceof plumagesof the CommonTern in early life appearsto be thus: (1) natal (acquiredbefore hatching); (2) juvenal; (3) first-wintereclipse; (4) first-summereclipse--'portlandica' (rarely a breedingplumage is acquired instead); (5) second-wintereclipse; (6) second summer, either: a, eclipseagain ('portlandica'),or b, first-nuptialplumage. Many birds require anotheryear beforeattaining their first-nuptial stage. The diagramof plumagesequence shows the probablenormal sequenceup to the fourth summer. It doesnot showduration of molt. Vol.•94• •8] a PALMER,'White-[aced' Terns 167 I havenot calledthe immaturesummer plumages 'nuptial' plumages, as Dwight (1925) has done with gulls, sincethe term is more ap- propriatefor breedingstages. The Common Tern resemblessome of the smaller gulls much more closely,both in time of beginningto breedand in time of assumption of breedingplumage, than many writers have hcrctoœorcrealized. The followingdata will make this clear. YouNG 'WHITE-FACED' TERNS Attainment of breedingage.--No Commonor Arctic Tern appears yet to have bccn recordedwith absolutecertainty as breedingin any form of immature 'portlandica'or eclipseplumage. Coloniesof Cabot'sand SandwichTerns (racesof Thalasseussandvicensis) and Royal Terns (T. maximus)have a fair shareof the breedingpopula- tion showingwhite in the œorchcads.These lighter birds appear to bc first-yearbirds and arc quite definitely known to breed in this plmnagc(H. W. Robinson,1910, 1940). One SandwichTern in "œull winter plumage"has bccn sccn to incubatean egg (Robinson,1940: 150). The Black-headedGull breedsregularly in "immature"plum- age (Kirkman, 1937: 111). Havcrschmidt(1931)mentions the same fact and also statesthat the Herring Gull (L. argentatus)and the LesserBlack-backed Gull (L. fuscus)breed bcœorcattaining the fully adult stage. Tinbergen (1936)reported that immatureHerring Gulls may pair, nest and copulatc,but fail to lay eggs. Many sub-adult Larinac very commonly'play' with nestingmaterial at the season when adults arc breeding. Juvenal CommonTerns oœtcnpick up grassand twigson or near the territory where they wcrc hatched,but the reasonfor their sodoing is not clear. Whereas the Marplcs (1934: 190) stated that Common Terns normally breed when one year of age, the Austins think otherwise. O. L. Austin,Jr., (1999: 130)concluded from his very extensiveband- ing and rctrappingrecords that this speciesdoes not breedat the age of one year, that all do not breed when two yearsold, but that the third year normally sccsthem productive. Later, O. L. Austin, Jr., (1932: 137)reported capturing a singleyear-old Common Tern which was incubatingeggs. Sincenothing was said to indicate otherwise, wc may assumethat it had attained the breedingplumage and that it had a definite connectionwith the eggs. Thus it seemsthat the Common Tern may rarely breed when one year old. From the factsthat year-oldbirds arc rarely rctrappcdat