233 2004 235by either a male or female Some extrapair copulationbox This was deduced by observing an extrapairbelow Further a single nest built in an alternativeof these copulation attempts Agai ID: 856778
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1 233 (2004) 234NEW ZEALAND JOURNAL OF E
233 (2004) 234NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 28, NO. 2, 2004attempted to defend their paternity by mate guardingendemic to New Zealand. They are sexually dimorphic,females (Armstrong and Ewen, 2001). The male has a1996). In comparison, the relatively dull female isdistinctive white wing bar (Craig, 1985). Hihi are cavitynesters and will nest in nestboxes (Rasch, 1989). Females, 1997). Females do all nest buildingprovisioning (Ewen and Armstrong, 2000). Hihi are(6080 m altitude). Forest remnants cover about 50%1997). Due to the lack of mature forest trees withand 1996/97). In both years the adult sex ratio was12:6 in 1996/97, largely as a result of differentialtranslocation. Nine nesting attempts were observed.of wrap-around C-size plastic bands and a numberedcommencement of nest building). Systematic searches30 m radius around each nest site. Observations
2 werebetween nests. This distance was ch
werebetween nests. This distance was chosen based onwithin a 30 m radius of their nesting tree. The observerwould move quietly around the 30 m perimeter. Theseand tracks provided for the large numbers of peoplediscovered to when egg laying was complete. The08:30, 11:0013:00 and 16:0018:00. The observationtime was divided equally among nests. Due to nestingobserve at one time. Therefore, each nest was usuallybehaviour. We recorded all copulation attempts,females. A measure of each extra-pair males presencetime an individual was seen being displaced, attemptingintervening period. Those individuals counted moreor not copulation occurred. As this was not alwayspossible, e.g. when the female was chased from thecopulations. Copulation was considered to occur if a 235 by either a male or female. Some extra-pair copulationbox. This was deduced by observing an extra-pai
3 rbelow). Further, a single nest built i
rbelow). Further, a single nest built in an alternativeof these copulation attempts. Again similar soundsposition) was achieved. We also, therefore, assumedobservation. Because one or both birds in a pair werewas in sight. The frequency of copulations andhour. Displacements of extra-pair males by pairedmales were calculated per paired male per hour. Anyattempts and % paternity within each nest. An average day of day of laying. Therefore, differencesthe five stages listed above. Differences betweenas a random factor in all models. The individualtime, given the small size of the population. Thisin order to normalise the respective data sets. Meansare expressed ± SE unless otherwise stated.to be resisted by the female. Resistance took the formand when they were absent. These extra-pair malesmates were not in their fertile period. Presence ofof egg laying ( )()at le
4 ast 12 days before egg laying. The firs
ast 12 days before egg laying. The first extra-pair = 0.011) (Fig. 1). Copulation attempts continued, 1999). Nests with thehighest rate of EPC attempts had a lower percentage of 236NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 28, NO. 2, 2004, 1996). Paired male hihi spent significantly )(Fig.3). It was also consistent up to 15 days before egglaying for the two pairs observed that long. Males werefemale. In addition, when females were near the nestfemale was present and absent. However, call rates ) = 0.92) and males did not significantly change their = 0.437).reproductive stage. Female was included in all models as a transformed) within broods in relation to attempted EPC ratesover the fertile period of the female. Each point represents anindividual nest. The line was fitted using simple linear when the female was present and absent (Fig.1). The = 5.63, ) = 2.84, = 0.026).
5 observed copulations). Data from the tw
observed copulations). Data from the two femalessampled for extended periods showed that attemptedup to 11 days before egg laying. It is interesting to notefemales solicitations and 34% of a males). Despite the = 1.918; = 0.209). 237 ET AL.: REPRODUCTIVE BEHAVIOUR IN STITCHBIRDS stages). Values are means ± SE, using nests as the units of Relationship between the frequency of extra-paircopulation attempts and within-pair copulation attempts. Eachdirectly. Males spent significantly more time withinregardless of female presence. Territory defense inactive. In addition, only active nest boxes (i.e. those). Such1995). Female hihi on average spent only 33% of theirtime within 30 m of the nest, suggesting nest defensemay be a poor form of paternity assurance. Males mayand Birkhead, 1993; Lifjeld and Marstein, 1994). This, 1987). Females may take1986). Given that female
6 hihi spent such large amountsextra-pair
hihi spent such large amountsextra-pair copulation (either solicited or resisted). Wenest. Attempted EPCs near nest sites also occur in(Alexander, 1974). Emlen and Wrege (1986) suggested constitute a reliable resource, i.e. theyare predictable both in space and time. Males takeforced copulations at the colony. Although hihi are not 238NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 28, NO. 2, 2004periods. Their presence peaked around one to twolaying commenced. A similar peak was observed forextra-pair copulation attempts. Such behaviourspresence and copulations to coincide. The egg issoon after the previous egg is laid (Drachmann 1997). Although copulations occurring during this1996). A peak in female fertility has been reported to, 1995). Timing of extra-pair copulation attempts infrom forcibly copulating with their partners. Such(Ewen and Armstrong, 2000). A paired males
7 , 1999). Despite paired males witnessin
, 1999). Despite paired males witnessing EPCsdid occur. The reasons why males may ignore femaleinvestigation. Males might focus their copulationnest. These results are similar to those found in indigobuntings (Westneat, 1988). Studies of alpine accentorsgive a good measure of paternity. In many species,DNA fingerprinting has revealed the presence of extra-, 1993). The close relationship between EPC andconspicuous. This close relationship suggests thatfrequencies. Additionally, as all observed attemptedresisted copulations may result in paternity. It isof low quality). Whether resistance by females is toof low quality). Whether resistance by females is to()resist to test male quality] is unknown. For thesecontrol these situations and select males as preferredmating behaviour of this species. Given the numerouson the manuscript, and to Ray and Barbara Walter andaddi
8 tional comments. We also thank Ian Jami
tional comments. We also thank Ian Jamieson andthe manuscript. Our research was done with permissionfrom the Department of Conservation. Financial 239 Wide Fund for Nature. Massey University,Armstrong, D.P.; Ewen, J.G. 2001. Testing food 87-92. 1211-1224.Birkhead, T.R. 1995. Sperm competition: evolutionary 755-775. 105-118. 843-848. 306- 51-60. 563-575. 765-771. 302-310. 223-228. Robertson, C.J.R.Davies, N.B.; Hatchwell, B.J.; Robson, T.; Burke, T. 729-745. 191- 321-328. 1394-1397. 240NEW ZEALAND JOURNAL OF ECOLOGY, VOL. 28, NO. 2, 2004 339-350. 789- 271-277.Animal Behaviour 45: 213-229.extra-pair paternity in eastern bluebirds. 1177-1183. Cambridge 170-186. 261- 127-133. 637-642. 27-36. recovery plan Cambell B. (Editor), Animal Behaviour 35: 865-876. 149-Westneat, D.F. 1990. Genetic parentage in the indigo 67-76. 7- Power, D.M. (Editor),Editorial Board member: Ian Jamies