BIRD-BANDING A JouR,',-^LoF ORNITttOLOGICALINVESTIGATION

VoL. III J:-xxuaRY,1932 No. 1

STUDIES OF S\VALLOWS BY THE BANDING METHOD •

By DR. SEINOSUKEUCHIDA Ornithologist to the Department of Anhnal Industry, Ministry of Agriculture and Forestry, ,

Ix view of the essentialneed of thoroughknowledge of the habitsof lnigratorybirds, the Japanesegovernment, which pays especialattention to the protectionof wild birds,has been making minute investigationsand collectingdata on migratory birds by entrustingmore than eighty schools,experiment Stations, mete- orologicalobservatories, and lighthouse-keepers,•vith the col- lection of the neededinformation for the sol!utionof this important problem. Moreover, since 1924 the governmentitself hasbeen carrying on bird-banding'-experifnkfitswith a view of contributingtoward the solutionof the sameproblem. The fol- lo•vinglist showsthe numberand speciesof birdsbanded by the Japanesegovernment since that time: . In this report about swallowsI will describein detail the resultsof the bandingexperiments referred to in the abovelist: There are three speciesof swall.owsmigrating to Japan: HMt•do rustica•uttural,•s. (Scopoli), ChimneySwallow; Hir- u;tdodaurica nipatensis (Hodgson), MosqueSwallow; t-Iiru.ndo. javmtica (Sparta), Liu-kiu Swallow. Of the abovethree species,the Li'u.ZkiuSwallow inhabitsonly Loo ChooArchipelago, and thus'is excludedfrom this report. As for the two otherspecie, .(see_map) the former is the com- moner in Japan, being distributedall over the country, whereas the latter principallyinhabits the westernpart-of the country. both nesting on the--cavesor ceilingsof human dwellingsand godowns.

XThe tabulated data of returns of banded swallows in my experiments I have already published in Japanese in a paper in Ch•j4 Ch•sa H•koku, Vol. 6. I have• therefore omitted them in this article, i• order to avoid complexity, confining myself t• de•rib. lug only the results of the work. UCfflD.%Stttdy of Swallo•c,s Bird-Banding January Vol.1952nI UCH•DA,Study of Swallows [3

The bandsused for swallowsin Japan,shown in Fig. 1 are usuallyof aluminumand are 12 mm. long and 5 mm. wide, weighingabout 6 centigrams. In Japantwo methodsof bandingare in practicaluse: one ro captureand band the birds while sleeping at night.and the other to captureand bandthem in the daytimeby nettingat the en- tranceand exit of their nestinghouses as they go in and out. To ascertainthe sexeswhen bandingsome skill is needed.so far as the ChimneySwallow is concerned.for the distinctionis rather difficult. In the main, however. the principal points of discrimination are as below: (1) The fentalegenerally has a slightlyshorter tail thanthe male. The lengthof tail: female,70-85 ram., 77 min. on the average;male, 81-100 nun., 88 mm. on the average. (2) The cloacaof the femaleis largeand oblong,while that of the maleis smalland slightly projected. (3) As/or the colorsof the undertail-coverts, forehead, chin, and throat,those of the maleare brighterand morebeautiful than thoseof the female.However, the distinctionis but faint in young birds. (4) The strengthof twittering uotes. (5) It is the femaleonly that sitson the eggs. Let me now show the banding resultsin the form of returns obtainedfor four years,1927-1930. The two speciesof swallows bandedduring the above period totalled2136, of which 124 returned.The localitieswhere they were bandedwere the seven prefectures of , , , Toyama, Hyogo, . , and . " The bandingof swallowsis exclusivelydone at their nesting- places,and their returnswere notedat the localitieswhere they were banded.They never came back to nest in localitiesother than thosewhere they were banded.Accordingly, it is to be un- derstoodthat the returns are only thosebirds which returned to the localitiesof bandingor nesting.The following table shows the resultsof bandingswallows by speciesfor the periodabove mentionedfor eachprefecture by years:

FIG. I, ALUMIIVUl•œBANDS FOR SW;J. LOWS USED BY THE JAPANESE GOVERNMENT 4] UcH[r)3.,Study' of Swallows JanuaryBird-Banding

1927

Species Prefecture Banded Adults Sexes Nestlings Returns Sexes

H. rustics Aomorl 3 0 J

,9• H. rustics Saitama 20 17 3

H. rusticst Hyogo 26 2 •1• 24

H. daurica Hiroshima 24 20

H. rustics Fukuoka 20 7 13 H. rustics 42 8 ; 34

TOTAL ' 135, Adults 38, 13 males, 10 females Nestlings 97.

1928

H. rustics Aomori 47 i 18 , 29',16•2 H. rustics Akita 9 [ 16

Saitama H. rustics Toyams 10'•5C.Xestling Hyogo •o•3 > y.J.G. 223, 23 H. dauriea Ad. 4/ Hiroshima --•9' 22 •2• Fukuoka Adult

qnitial letters of the two species banded.

TOTAL 568, Adults 173, 88 males, 85 females. 14 Nestlings H.D.N. Nestlings 381, 13 males, 16 females. Returns 17 15 Ad. H. rustics, 6c•, 7(2, 1 Ad. c• it. daurica. Vol. III UC•DA, Study of Swallows 1932 [5

1929

Species PrefectureBanded Adults Sexes Nestlings Returns Sexes

H- rustica Aomori 86 16 70

1 H. rustica Akita 61 18 43 Adult

Saltams 156 42 114 I Adults53 \25 9 1 H. rustica Toyurea 30 12 18 Adult

H. rustics Hyogo 194 79 '•41 9 115 f23c' H. daurica 106 47 .24 9 49 '6• H. rustlea Hiro.qhima 26 13 •7 9 13 Adults

Fukuoka 79 18 61

TOTAL 738, Adult H. rustics 168, 79•, 101•2, Nestlings 434. Adult H. daurica 47, 23cf, 24'9. Nestlings 59. Returns 63. Adults 63, 34cf, 29 9 ß

1930 '

6 H. rustics Akita 160 52 108 Adults Remarks

20 Saitama 186 96 90 Ads. 19 Nestlings 139

H. rustics Toyama 20 14 6 Adults

2o',29 H. rustics Hyoga 250 60 190 7 H.J.G. 2 adult, 1o'.1 9 - 4 Ads. 3 H.D.N. and I nestling

H. daurica Hiroshima 30 8 5 Ads.

H. rustics Fukuoka •9 15 32 2

H. daurica

TOTAL 697, Adult H. rustics 239, 810', 104•. Nestlings 426. Adult H. dauriea 10, 6c•, 4•. Nestlings 22. Returns 44.41 H. rustlea. 40 adults, 12o', 22 9, 1 nestling. 3 H. daurica, 2 adalts, lc•, 1 •. I nestling. Studyof Swallows JanuaryBird-Banding

SUMMARY Total Birds Banded, 2136 Aomori Prefecture Adult H. rustica 136. 34 (19 malesand 15 females) and 102 nestlings. Akita Prefecture Adult H. rustica 238. 71 (1 female) and 167 nest- lings. Saimma Prefecture Adult H. rustica 550. 259 (121 males and' 138 fe- males) and 291 nestlings. Adult H. rustica 70. 29 (14 males and 15 iemales) and 41 nestlings. Hyogo Prefecture Adult H. rustica 867. 164 (77 malesand 87 iemales) and 529 nestlings. Adult FI. daurica 59 (31 malesand 28 females) and 115 nestlings. HiroshimaPrefecture Adult FI. rustica 85. 42 (18 malesand 17 icmales) and 43 nestlings. Fukuoka Prefecture Adult H. rustica 190. 43 (7 males and 26 females) •ud 14&nestliugs. Adult H. daurica 2 (1 male and 1 iemalc). Returns,124. Of these,120 were H. rusticaadults, comprising 52 males and 57 females. Four were H. daurica, 3 adults and 1 nestling. The percentageof returnsof adult birdsas shownin the fore- goingtable, is as follows:

Percentageof Returns of Adult Banded Swallows Banded One YearA#o Banded TwoYears,4#o 1928 1929 1930 Average 1929 1930.dvcra9e Akita 100 33 67 Saitama 59 49 24 44 12 9 11 Toyama 33 33 33 I-Iyogo 33 5 13 Hiroshima 57 36 31 41 5 5 Fukuoka 13 11 8 Total Average 34 8

Accordingto the abovetable, the percentageof returnsaver- ages34 in six prefectures.But as returnsin Hyogoand Fukuoka Prefectureshave been excepted for the reasonthat the investiga- tion has beeninadequate, the averagepercentage in the other four prefectures--Akita, Saitama, Toyama, and Hiroshima-- amounts to 46. In other words, about one half of the adult swallowsbanded return to the neighbourhoodof their nesting placesof thepreceding year. The death-rateof swallowsis gen- erallyconsidered exceedingly high because of theirlong-distance migration.We may safelyconclude, however, that mostof them are,as a rule,in thehabit of returningto theirbreeding places of the precedingyear in view of the resultsobtained as shown in the above table. Vol. II! UCHm.x,Study o[ S'rcallo'•'s 1952 [7

• Hitundorus•icagu•uralis Aomor[ v• Hitundodaurioanipalensis

N

•AP SHOWINGDISTRIBUTION OF Two SPECIESOF SWALLOW• I,N- JAPAN 8] UcarD^,oCtudy ofSwallows JanuaryBird-Banding

When we consider the localities where the banded swallows returned,we find that almostall of them return to placeswithin the limits of the villages or towns where they lived during the preceding3'ear, the majority returning even to the samehouses where they nestedthe precedingyear. In fact. out of the 124 examplesof returnedbirds. thosewhich returnedto nesting- housesother than the ones occupied'the precedingyear were only 31. T•vo of them nestednext door to their nesting-houses of the precedingyear. There were 8 examplesof swallows whichreturned to the samevillage or town for threeconsecutive years,and 3 of them even nestedat the same housesfor three consecutiveyears. With regard to the 31 examplesreturning to housesother than their old onesof the preceding5-ear. when ob- servedmore carefully, it has been found that most of them. •n spiteof havingreturned to the samehouses once. lost their mates or werecompelled to chooseother houses,owing to the repair.or someother condition of their old houses.It may easily be seen that swallwstend stronglyto return not only to the samelocali- ties but alsoto the:samehouses occupied during the preceding )-ear. It is a very interestingquestion why so few nestlingswallows return to their old nests.My bandingwork so far has resulted in a very smalluumber of suchreturns, consisting only of the fol- lowing three examples:

No. 36413 Chimney Swallow. One of the 114 nestling swallows banded at Shichiri-mura. Saltarea Prefecture. August 9, 1929. returned to Noda-mura. an adjoining village in the sameprefecture, May 28, 1930. No. E-50 Chimney Swallow. One of the 24 nestlingswallows banded at Uozumi-mura. I-Iyogo Pre- fecture,July 29, 1927. returnedto the samevillage, May 30. 1928. No. 48391 Mosque Swallow. One of the 59 nestling swallows banded at Chigusa-mura. Hyogo Prefecture,June 28, 1929, returned to the samevillage. July 17. 1930.

That returning nestlingsare so remarkablyscarce may be explainedby the fact that they seek new breediugplaces the followingyear without returningto their old homes.or to the extremelyhigh death-rateof swallo•vsduring the first 3-ear.Be the answerwhat it may, after repeatedbanding experiments, one finds that very few swallow nestlingsreturn to their old breedingplaces the succeedingnesting season, as indicatedby the following facts: In various localitiesWhere bandingexperi- mentshave been made in Japanthe numberof s•vallowsbreed- Vol.1932III ing there>'ear after year shows no noticeahleincrease or de- crease.Generally speaking, almost the samentunber of nests is yearlylotrod at thevarious houses. Now, as thenumber of returningparent swallows is 46 percent, as already mentioned, it is onh' uatnral to conclude that the number of returning nestlin• is the remaining54 per cent.According to investiga- tionsmade by Mr. TominosukeNibe (CI•6jlt Ch•saH6koklt, Vol. 2; pp.55, 62 and63,) a pairof parentswallows hatch 7.01 nestlingsas an averagebrood; in otherwords, one swallow gen- erates3.5 nestlings.Now, putting the percentage of thereturned nestlingsat 54 per centof that of parentbirds, we cometo the conclusionthat thesenestlings which return to theirold breeding -placesconaprise only 13.1per cent,it beingpresumed that the remaining87 per centhave died or soughtnew breedingplaces in other localities. Swalloxvswhich have returned for txvo consecutiveyears are very few. In SaltareaPrefecture only two retnrnedby 1929out of those banded in 1927; nine returned in 1930 out of those bandedin 1928; in Hiroshima Prefecture only one in 1929 out of thosebanded in 1927; and only one in 1930 consecutively for two yearsout of thosebanded in 1928•thirteen in all. There has beenno return for three consecutiveyears so far. It shouldbe noted,however, that bandinghas been carried out for only four years,and the numberof birds bandedhas not beenvery large.Although birds returning for threeconsecutive years mav likely be discoveredin future, it is consideredthat suchcases will occurvery seldom,if at all. In view of the aboveobservations, it may safelybe concluded that whereasthe death-rateof swallowsseenas to be rather high until they reachthe ageof a year, the percentageof returnsthe followingyear is highestin caseof adult birds,reaching about half' of all the birds banded; that the number of returns of three- year-old swallowsis lessthan for two-year-oldbirds; and that returnsof four-year-oldsare exceedinglyrare. From thesecon- siderationsit may be presumedthat the averagelife of a swal- low is under four years. Let us now look into the questionof permanentmating in the caseof the swallow, Whether in the East or in the West, various popularviews in connectionwith the questionof bird-partner- ship haveprevailed since times of yore. Most of the authorsof these traditions seemto consider birds in general to be quite chastesuch as the traditionwidely believedin Japanregarding the Mandar/n Duck. Nevertheless,'according to observations- madeby S. P. Baldwin,a pioneerin researchesin referenceto 10] Ucam^,Study of Swallows JanuaryBird-Banding the mating of birds• the House Wren in the United Statesof America, contrary to popular conjecture, COlnlnonlychanges its mateevery bi'eeding season. examples of continuingrela- tionswith the samemate for two breedingseasons being found exceptional.In the caseof otherspecies Mrs. M. M. Nice writes in her paper,"Do Birds UsuallyChange Mates For The Secoud Brood?"," that "in the caseof sevenpairs of [North Alnerican passerincbirds] there was a shifting of matesili one season, whilein twenty pairsof elerenspecies there was no change." As to the permanenceof the mating relationamong Japanese swallowsan examhaationof sixty-four pairs which were handed showedthat tweuty-six pairs returned to the salne nest-houses in the samemating relationas in the preceding3'ear: there were no examplesof the same mates returning to nest-housesother than thoseoccupied by the samepair the previousyear: those returningto thesame nest-houses asin theprecedin• )-ear with different mates were six males, nine females, and three others of unknownsex; and, finally, thosereturning to nestsother than the onesused the precedingyear and changingtheir mates,num- bered eight males, eleven females,and one of unknown sex. If the new mates of the swallows in the above number are carefully examined. it is found that all of them are not banded, or were bandedlater than in the year in which their old mates were banded.In other words,there is nothingto prove that ex- changesof mates were made betweensxvallows which were bandedin the sameyear. Therefore it may be supposedthat they have soughtnew matesbecause their olcl matesdied or did not return to their old b'reedingplaces. The fact that all thosewho did not changetheir matesreturned to the samenest-houses as in the previousyear, not to other houses,as pointed out above,is also noteworthy. As for those which changedtheir mates, eighteen of them returned to the same nest-housesoccupied in the preceding year,and twenty to housesother than in the precedingyear. \Vith referenceto the latter, it is consideredthat they had most likely been forced to abandontheir old nest-houses,being tempted by their new mates to the fatters' old nest-houses. : Is there any recognizabledifference between the male and the female in attachment to their old nest-houses? It has been provedthat out of thoseswallows which have changedtheir matesfrom thoseof the precedingyear when they were banded,

•S. •P. Baldwin: The •{arrlage Relation of the l•ouse Wren, Troglodytc.* edo• •do•, ,'hal:, Vol. XXXVIII, pp. 227.244, 1921. 2Bird.Bandlng, Vol. I; pp. 70-72, 1930. Vol.1932III UC•In&Study o/Swaltores [l 1 thosewhich returned to the samehouses as in the previousyear were five males and nine feinales, whereas those which returned to housesother than in the previousyear, that is, thosewhich may be consideredto havebeen lured by their new mates.were eight malesand elevenfemales. It is interestingto note that. in the casesboth of thosewhich returned to theirprevious honses andof thosewhich returned to otherhouses than in the preced- ing year, the numberof felnaleswas. larger than that of the males.Judging from this restilt.it may be considered1)r•ved that the degree of attachlnentof swallowsto their old llest- honsesdoes not necessarilyclepend upon the sex. _Asmay be inferred froin the above-enulneratedfacts. the swallowhas a persistenthoming habit; not merelydoes it re- turn preciselyto the localitywhere it nestedin the previous year. but it is usual that the bird returns even to the same i•est-houseas in thepreceding year. Consequently, a swallow onght to meet the same nmte in the same nest-houseas in the previousyear, and naturallyboth of themnest and breed there, renewingtheir inarriagerelation. However,in caseof eitherthe male or thefemale having died, the survivormay be compelledto seeka new mate,because of beingunable to findits preyionsmate, in spiteof returningto its old nest-house. Of the male and female swallows which were banded after their sexeswere ascertained,there are only five birds which nestedfor threeconsecutive years: three of themchanged their matesin thethird year,while the remainingtwo continuedwith the samepartners for threeconsecutive years. Therehas so far beenonly one instance ofa bandedJapanese swallowbeing recaptured in its hibernatingt place. The record is importantbecause it is uniqueso far asJapan is concerned. Placeof banding,Notomura Hiki-•ln SaitamaPrefecture, (Themiddle part of themainland of Japan). Date of banding, July1, 1926.Place where it wasrecaptured, Lagna State. Luzon, philippineIslands__ Date oœ recapture, October 26, 1926.

•In usingthe word "hibernating," Dr. Uchidadoubtless has in mindo•r English word "wintering." In his discussionof thetwo species o[ Japaneseswallows he appearsto treatboth of themas a singlespecies in writingo• his returns.This, however, does not invali- dateany of his conclusionssince the numberof •*ipalen.ri.rreturns is negligible($ out of 124), the conclusionshaving to do mainlywith gutturalis.--Ei>rroa.