The timing of moult in males and females of the monomorphic Pale-winged nabouroup Adrian J. F. K. Craig, Martine Hausberger, Bo Bonnevie, Laurence Henry

To cite this version:

Adrian J. F. K. Craig, Martine Hausberger, Bo Bonnevie, Laurence Henry. The timing of moult in males and females of the monomorphic Pale-winged Starling Onychognathus nabouroup. African Zoology, Taylor & Francis, 2015, 50 (1), pp.69–71. ￿10.1080/15627020.2015.1021548￿. ￿hal-01150388￿

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The timing of moult in males and females of the monomorphic Pale-winged Starling Onychognathus nabouroup

Adrian JFK Craig1,4 , Martine Hausberger2, Bo Bonnevie3 and Laurence Henry2

1 Department of Zoology and Entomology, Rhodes University, Grahamstown, South Africa 2 UMR CNRS 6552 Ethologie animale et humaine, Université de Rennes-1, Rennes, France 3 Information Technology Division, Rhodes University, Grahamstown, South Africa * Corresponding author, email: [email protected]

Pale-winged Onychognathus nabouroup inhabit the arid western interior of southern Africa and moult– breeding overlap may occur. We collected field data in two successive years on the moult of individual , whose sex was confirmed by genetic techniques. Small samples revealed a non-significant tendency for the moult of females in the early stages of wing moult to be more advanced than that of males in both years, but also clear evidence that the starting date of moult differed in the two years. In this species the moult schedule may thus be variable at both the individual and the population levels.

Keywords: moult, Onychognathus , sexual dimorphism, southern Africa, starling

As one of the major events in the annual cycle of all birds, individuals were subsequently sexed. From 1 to 7 November the timing of the complete moult should be adjusted to 2012 an additional 24 adult birds were captured, all of which the requirements of the individual , as well as being were sexed, and two birds ringed and sexed in the previous subject to selection at the species level (Stresemann and year were recaptured. Birds were caught in baited clap Stresemann 1966; Barta et al . 2006). Recent studies of traps, individually colour-ringed for behavioural observa- South African birds have shown that the timing of tions, and released at the capture site. None had brood moult often varies regionally relative to the timing of rainfall patches, and it appeared that nesting had not yet started in in relation to the breeding season (Hulley et al. 2004; either year. Standard measurements (wing, tail, tarsus and Craig et al . 2010). In addition, in some sexually dimorphic culmen length in millimetres, body mass in grams) were species the post-breeding moult starts earlier in males than taken, and primary wing-moult was scored according to the in females (Bonnevie and Oschadleus 2010). This could system whereby 0 = an old feather, 5 = a fully grown new be ascribed to reduced male involvement in the care of the feather, with 1–4 representing intermediate growth stages young in such species (Verner and Willson 1969; Payne (Ginn and Melville 1983). These scores were converted to 1984; Oschadleus and Osborne 2005). ‘percentage feather mass grown’ for analysis in the moult Of the 11 members of the genus Onychognathus, only model of Underhill and Zucchini (1988) as written for R (Erni two species are not sexually dimorphic in plumage. Male et al . 2013). Sex was ascertained by DNA sexing (Labofarm, and female Pale-winged Starlings O. nabouroup are France) from samples of 5–10 small feathers from each bird not distinguishable on plumage characters (Craig and (Han et al . 2009). Feare 2009), although they can be separated in the hand One female bird had interrupted wing moult, with the with some confidence by a combination of morphological two inner primaries new, the rest old, and no growing measurements (LH et al. unpublished data). Earlier work feathers, although there were some body feathers moulting. has shown that moult and breeding in this arid-zone species This is the first record of interrupted primary moult in this can overlap at both the individual and the population level species (Craig 2012), and there is no apparent explana- (Craig 2012). This raises the question as to whether males tion for its occurrence in this bird. Although the sample and females differ in the timing of their moult, which could sizes were relatively small, in both years the stage of moult reflect different investments in parental care. Therefore we as represented by the outermost growing primary remix investigated the stage of primary moult of wild Pale-winged showed a normal distribution, with wing moult in female Starlings, which were later sexed by genetic methods. birds rather more advanced than in males (Figure 1). The The study was carried out in Augrabies Falls National data were then examined statistically (using the Welch Park, Northern Cape, South Africa, (28°59′ S, 20°33′ E) two-sample t-test), comparing mean percentage new over two consecutive years. From 25 to 28 October 2011, feather mass grown in males and females in each year. a sample of 58 adult birds was captured and 42 of these In 2011 females were on average more advanced in their

female (January), whereas three females had more more be females to advanced more inthan male birds. had f tendency a at hints also This [1]). females February [2], three (J counterparts male their whereas than moult wing advanced (January), female correspond the than moult primary advanced more had [1]); November [1], March [2], (January birds both pri same the reached had moult cases four in moult: eight ‘pairs’, these Of locality. same the at date the on collected were Starling Pale-winged female mal a a which in instances 21 are there 2012), (Craig P dacd bt hs a fr rm en statistically ( cant being from far was this was but moult advanced, female that indication an also was there a sgiiaty oe dacd n 01 hn n 201 in than 2011 two-sample in (Welch advanced more significantly was we Y November. 1 after were captured birds all 2012 in data w October, 2011 of week In last the during years. mostly collected the between difference icant feather mass grown; for males, males, for grown; mass feather ol, n te ifrne prahd infcne a ( level 5% significance approached difference the and moult, P to outwards proceeds and 7 P1 at to starts 1 Moult 2012. and 2011 October 28 to 25 from Africa, South 1: Figure igd trig hnld t urbe Fls Nationa Falls Augrabies at handled Starlings winged showsthe outermost growing primary eachin case NUMBER OF BIRDS = Among the museum specimens examined previously previously examined specimens museum the Among However, for both males and females there was a sig a was there females and males both for However, 1 1 2 3 4 5 6 7 8 1 1 2 3 4 5 0.009). 0.009). None P1 P2 P3 P4 P5 P5 P4 P3 P2 P1 None N = Stage of early primary moult of male and female Pal female and male of moult primary early of Stage 16 males, 16males, 10females, N = 17 males, 25 females, females, 25 males, 17 OUTERMOST GROWING PRIMARY t -test, comparing mean percentage percentage mean comparing -test, P = P 0.503). 0.503). = P 0.018; for females females for 0.018; = 0.064). For 2011 For 0.064). Female Male 9; the 9; or moult in moult or November a wing had one male one et moult et signifi- signifi- mary in mary hereas anuary Park, l

and e 2011 2012 same more more x t the the t -axis nif- ing re re e- 2 2

hra tre ae wr nt olig Ti my als may This moulting. less synchronous imply m moult inharsh environments. not wing were of males stages three different whereas very in Starlings Somali fem three handled (2004) Neumann and Gedeon period, Socotra Starling Starling Socotra species occur, the Starling Somali the occur, species two Socotra, of Oschadleus island arid and the On (Bonnevie 2010). Cape Western the of region m r winter wing the in Starlings of Red-winged female and male duration or timing the in Moult–breeding difference cant n is 2009). there but species, Feare this in occurs also and overlap (Craig Starl Africa Pale-winged the South with sympatric locally is but habi humid more favours species, dimorphic sexually n eta Aaoi, ono e al et Johnson Amazonia, central in penicillatus penicillatus Honeyeater White-plumed the in but moult, co metabolic the measured directly have studies Few b and moulting overlap to likely more were duration long a and rates growth feather slower with a species et (Morales individuals al found et such Buchmann have 2007; in others survival breedi 2013), and improved Hau moulting and overlap (Echeverry-Galvis that birds passerine to the moult period significantly reduced the energeti the reduced significantly period moult the et Cag n Fae 09. oee, h pro of period the However, 2009). Feare and (Craig i chicks nest feed sexes both but incubate, females only Afr this starlingsother of in rainfall. localAs on depe probably markedly, vary can breeding of timing whereregularphenomenon2012),(Craig a is overlap breeding arid-zone bird. breeding opport this in basis daily on a replacement feather has been has fundedbythe CNRS(France) andthe Nation bi the ringing and catching in assistance their for an Hulley Pat Smith, Diane Zwieten, van Arjen Ford, als We France. Loudeac, in Labofarm labo at staff the the on of depended birds the of sexing The ation. staf parkand thewe to NationalgratefulPark, are A at birds ring and capture to permission for Parks Acknowledgements and moulting strategies in this arid-zone species. the of flexibility the confirms seasons successive o were moult of timing the in differencesthat fact cyc moult the of feather stages different of at occur rate may ment the in changes that possible is it peri moulting entire the from information more need al We 1999). Hemborg 1997; Nilsson and Svensson (cf. both knownsexuallyin dimorphic monomorphicand sp mo annual the of timingsubsequent the in and care, in sexes the between differences but ex this, biological for tion any offer yet cannot We species. earl mal in this than start females may in rapidly moult more curr progress wing Our and/or primary obs.). that pers. suggest MH data (February; nest the of out fledgl feeding seen been have samples) genetic from determined;females(colour-ringedonly date to bir accu been not has adults the on dependence juvenile lhuh oe il tde ugs uvvl cos survival suggest studies field some Although The Red-winged Starling Starling Red-winged The For Pale-winged Starlings it appears that moult–bree that appears it Starlings Pale-winged For Hoye and Buttemer (2011) found that extending that found (2011) Buttemer and Hoye — We are indebted to South African National African South to indebted are We — O. frater. frater. O. . . 2009). In understorey passerines understorey In 2009). During the same three-week three-week same the During ncontu morio Onychognathus . . O. blythii blythii O. 21) on that found (2012) Onychognathus f for their cooper-their for f Lichenostomus rds. This project This rds. ican radiationican ugrabies Falls ugrabies d Mary Hulley Mary d o thank Mike thank o alResearch ratory skills skills ratory bserved in bserved unistically c costs of costs c ds, sexed ds, parental o signifi- o breeding and the the and replace- replace- er moult er reeding. le. The le. ult, are ult,

oult in in oult d as od, plana- plana- as theas ing in in ing ainfall ndent sts of sts rately the n ecies es in in es oult, oult, tats, ding ings a , ent ent ale ng ng ier ier so ts o l .

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