UNIVERSITY OF AMSTERDAM. René W.R.J. Dekker. the Netherlands. Abstract. of moult is more or less. the reason is obvious, but Emu, Dromaius, and

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1 Beuforti INSTITUTE OF TAXONOMIC ZOOLOGY (ZOOLOGICAL MUSEUM) UNIVERSITY OF AMSTERDAM Vol. 40, no. 6 December 19, 1990 Moult wing til-fers in Ostrich, Struthio cmelus Tim G. Brom & René W.R.J. Dekker Institute Txonomic Zoology (Zoôlogisch Museum), University Amsterdm, P.O. Box 4766, 1009 AT Amsterdm, Nerls Abstrct Structure moult wing til full-grown Ostrich, Struthio cmelus, re described. In wing, t lest three fer genertions could be recognized. The pttern moult is more or less symmetricl in both wings sequence fer replcement is not rom. The til consisted 93 fers two different genertions. Moult in til proceeds symmetriclly seems to be fully irregulr. The phylogenetic significnce se findings is discussed. INTRODUCTION reson is obvious, but Emu, Dromius, especilly Rhe, Rhe, Ostrich, Struthio, Differences in ptterns wing til moult in possess sufficient wing fers to llow for different vin tx hve received much ttention (Stresemnn, 1963, 1967; Stresemnn & Stresemnn, 1966, references rein), lredy from erly observtions by Heinroth (1898) Beebe (1914) re is consensus mong systemtists tht phylogenetic component must be involved. However, phylogenetic interprettion phenetic clusters in which se studies hve resulted, remins uncler. Most studies on moult flight-fers hve been purely descriptive study on ir moult ptterns. Although mny chrcters in rtite birds be linked with my secondry loss bility flight, re is no reson to ssume beforeh tht y hve lso lost ncestrl pttern wing moult. Little informtion for wild ostrich is vilble (e.g. Crmp & Simmons, 1977:41). Duerden (1911) observed tht "in wild ostrich only few wing plumes re growing t ny one time insted full number s in domesticted bird, where growth is significnce different ptterns hs been regulted rtificilly" lthough " ssessed in functionl ecologicl terms only nturl order ccording to which vrious (e.g. Wttel, 1985; Noordhuis, 1989). Moult in rtite birds hs been studied. hrdly plumes pper hs not yet been determined, it is well estblished tht fers towrds both With respect to species with smll number ends wing develop in dvnce those in remiges (Kiwi, Apteryx ; Cssowry, Csurius), middle." Noordhuis (1989), however, 103

2 Fig. 1. Left ssumed for from wing, showing remiges (seen irregulr pttern n Struthio. In primry moult Cssowry, Kiwi, ll bove; til-fers no be cn Emu, generl only Struthio prts, hs dt refer bird hd confiscted September curled loose-vned neir structure been exmined In this nor fers. its moult Apprently, study, pttern hs moult structure Ostrich, Struthio Study skins tion birds For one re not ded freshly cmelus cnnot n is L., this kind 104 not investigtion lished s wing (pi) wing specimen 1758, re in cptured Amsterdm 1988 died Zoo re eight (Ntur counted wild, ws in Airport months lter in Artis from Mgistr). (si) outwrd, inwrd, innermost to moult turn it is cler tht for fter ir centre were djcent removl ll wing-coverts. living purpose. RESULTS ptterns, to freshly Structure moult wing lrge series single specimen, strting point Remiges exmined outermost des- vilble, this report, bsed til-fers from outwrd. or upper while be hled for this recogni- ccurte n secondries from specimens fers, compelled Since re stuffed investigtion refore ded birds. esily exct or suitble for individul rrow. full-grown femle Ostrich. t Primries coverts Ostrich by fr. so til-fers in cribed. to been Amsterdm (tl) indicted METHODS hinder consisting long, til P1 distinguished fering removed). MATERIAL AND The mong been All Rhe hve coverts is on pub- furr studies. In & 2): (p9), wing, 16 primries eight metcrpls four middigitls were present (pl-p8), one (pl0-pl3) (Figs. 1 ddigitl three

3 sheet Fig. 2. Moult primries ir coverts. Not bene must be sheth. predigitls (pl4-pl6). The lul consisted secondry-coverts were structurlly incomplete: four smll fers. in left wing, under wing-coverts sio The exct number secondries ws more difficult to estblish, but t lest 20 were sl5-s20 were lcking, in right wing those sl7-s20. counted (Fig. 3). The under surfce wing In left wing, 26 newly replced fers were growing (Figs. 2 & 3): six primries, two ws bre except for single row under wingcoverts. Ech primry ws provided with one fers lul, seven secondries, seven upper one under wing-covert, which ly distlly to primry y belong to (oriented towrds These series becme interrupted towrds outer primries: pi4 in left wing pl4 pi6 in right wing hd wing-tip). no upper wing-covert, in both wings pi5 pl6 hd no under In wing-covert. both wings, upper secondry-coverts formed upper primry-coverts, three upper secondrycoverts, one under secondry-covert. In totl number right wing, ws 30: eight primries, one fer lul, nine secondries, six four upper primry-coverts, upper one under secondry-coverts, primry-covert, one under In secondry-covert. both wings, primries t lest three different complete series 20 fers, but under genertions could be recognized. 105

4 Fig. 3. Moult secondries ir coverts. Structure moult til ech remex. The regulr implnttion The til consisted mny loosely structured fers, which grded into body-fers 4 (Figs. & At both sides 5). pygostyle, eight rows seven fers were present, but s some se rows were structurlly incomplete (not due to moult), totl number fers ws 93; 45 in left hlf 48 in right Unlike condition in (Fig. 5). flying birds, only both centrl verticl rows were ttched to pygostyle, ll ors were loosely implnted in skin, into subcutl ft projecting lyer. Two fers could be distinguished: genertions 40 old 53 new fers. All freshly moulted ones still hd (remnnts ) sheths, 33 se in were growth (figs. 5). secondry coverts demonstrtes tht Struthio is eutxic (cf. Steiner, 1918). Structurl between left symmetry right wing (Figs. 2 & 3) one individul bird is not in exceptionl this since Duerden species, (1920) not only reported similr differences in but even in number wing-coverts, remiges. The ptterns moult re more or less symmetricl in both wings. The suggestion tht fer replcement proceeds t rom cn refore be excluded. The primries might be moultedfrom severl foci, but serilly descendnt moult cnnot be excluded. Our observtions do not support Duerden's (1920) sttement tht outer remiges re moulted prior CONCLUSIONS AND DISCUSSION to those in centre wing. The replcement secondries seems to strt from Structure moult wing Our observtions on structure wing fully gree with those Wry (1887: fig. 2) refore we conclude tht reports number primries or thn 16 (Jeffries, 1881; Furbringer, 1888) re erroneous. The number secondries lest with (t 20) corresponds erlier reports (e.g., Wry, 1887). severl foci. The direction in which moult proceeds cnnot be determined in single specimen. The upper wing-coverts seem to strt moulting lter thn ir corresponding The remiges. under wing-coverts strt lter thn upper coverts. Both rows wingcoverts re not shed with ir toger remiges, but re more likely moulted in groups Similr to wings neognthous birds, (see e.g., upper wing-coverts p9-pl3 in Fig. one upper one under covert ccompny 2)- 106

5 Fig. 4. Til, seen from below. Fig. 5. Structure moult til. 107

6 Structure moult til primry-moult (scendnt, trnsilient, from The til full-grown Ostrich consists focus t p4 or p6, etc., see Stresemnn & Stresemnn, 1966) cn n be considered to be eight rows fers The totl seven (Fig. 5). number 93 rectrices is considerbly higher thn to reported by Stresemnn ( :741). It is difficult to scertin which series til- pomorphic cn be used to define monophyletic groups mong ACKNOWLEDGEMENTS extnt birds. fers is homologous with til in flying The Zoo Ntur Artis Mgistr kindly pro- birds. The number rows t both sides pygostyle corresponds with eight pirs vided Ostrich. Assistnce in dissecting bird ws given by Hns vn Brwijk. Kees til-fers found in chick Struthio Roselr Jn Wttel commented on In (Steiner, 1946). embryo biseril follicles is rrngement found, from which 16 rectrices with ir upper coverts will grow, while underside til is bre (Steiner, mnuscript. The investigtions were in prt Foundtion for Fundmentl supported by Biologicl Reserch which is sub- (BION), sidized by Nerls Orgniztion for 1946). This might indicte tht fers in Scientific Reserch (NWO). most ventrl row represent homologues norml til-fers tht six dorsl REFERENCES BEEBE, C.W., Preliminry phesnt studies Zoologic, 15, (1): CRACRAFT, J. & D.P. MINDELL, The erly history rows consist modified til-coverts. Alterntively, one or both centrl verticl rows til-fers, which re ttched to pygostyle, my be homologous with tilfers in flying birds. This could be determined by mrking til-fers in newly htched chicks following ir development into dult til. modern birds: comprison moleculr morphologicl evidence: In: B. Fernholm, K. Bremer & H. Jôrnvll, eds.: The hierrchy life: molecules morphology in phylogenetic nlysis (Excerpt Medic, Amsterdm, New York, Oxford). Although til moult ws most ctive in CRAMP, S. & K.E.L. SIMMONS, (eds.) The birds centrl rows fers, it seems most likely tht sequence fer replcement is fully irregulr. No symmetry exists between both til-hlves. The left side consists considerbly western Plerctic. Vol. 1 (Oxford University Press, Oxford, London, New York). DUERDEN, J.E., Experiments with Ostriches-XVI. The plumges Ostrich. Agricult. J. Union S. Afr., 1 (1): more newly replced fers thn growing, Methods degenertion in Ostrich. J. right side. Genetics, 9: FURBRINGF.R, M., Untersuchungen In cse furr studies will confirm tht primry moult in Ostrich is serilly descendnt, this will hve phylogenetic implictions. Since tinmous, glliforms, nseriforms ll show descendnt mode or vrition on this scheme ("Stffelmuser" in tinmous some simultneous in glliforms, nerly this is nseriforms), pttern most probble chrcter-stte t bsl node vin tree (see Crcrft & Mindell, 1989). It cn refore be excluded tht simultneous moult is plesiomorphic condition s ws suggested by Verheyen (1958). Or ptterns zur Morphologie und Systemtik der Vogel, zugleich ein Beitrg Antomie zur der Stiitz- und Bewegungsorgne. 1. Specieller Theil: i-xlix, 1-834; 2. Allgemeiner Theil: , Tf. I-XXX (Vn Holkem, Amsterdm). HEINROTH, O., Uber den Verluf des Schwingenund Schwnzmuser der Vôgel. Sitz. Ber. Gesellsch. nturf. Freunde, 1898: JEFFRIES, J.A., On number primries in birds. Bull. Nutt. Orn. CI., 6 (3): NOORDHUIS, R., Ptronen in slgpenrui: oecysiologische npssingen. Limos, 62 (1): STEINER, H., Ds Problem der Disttxie des Vogelflùgels. Jen. Zeitschr. Nturwiss., 55, (N.F. 48) Heft 2/3: Tf

7 ,,, Zur Pterylose des friknischen Strusses, VERHEYEN, R., A propos de l mue des rémiges Struthio cmelus L. Act. Soc. Helv. Sci. nt., 126: primires. Gerfut, 48 (2): STRESEMANN, E., In: W. Kukenthl, WATTEL, J., The relevnce moult studies for txonomy. Act XVIII Congr. Int. Orn. (Moscow): ed.: Hbuch der Zoologie. 7, (2) (de Gruyter & Leipzig). & Co, Berlin WRAY, R.S., On some points in morphology Txonomic significnce wing molt. Proc. wings birds. Proc. zool. Soc. London, 1887: XIII Int. Orn. Congr. (Ithc): Inheritnce dpttion in moult. Proc. XIV Int. Orn. Congr. (Oxford): STRESEMANN, E. & V. STRESEMANN, Die Muser der Vôgel. J. Orn., 109: (Sonderheft). Received: August 29, Institute Txonomic Zoology (Zoôlogisch Museum), University Amsterdm, P.O. Box 4766, 1009 AT Amsterdm, Nerls 109