Aziz Asian a, Mustafa Yavuz a & Ali Erdogan a a Department of Biology, Faculty of Arts and Sciences,

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1 This article was downloaded by: [Akdeniz Universitesi] On: 30 December 2014, At: 03:56 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Israel Journal of Zoology Publication details, including instructions for authors and subscription information: A COMPARATIVE STUDY OF THE BREEDING ECOLOGY OF THE HOUSE SPARROW (PASSER DOMESTICUS L.): TIMING OF BREEDING AND BREEDING SUCCESS Aziz Asian a, Mustafa Yavuz a & Ali Erdogan a a Department of Biology, Faculty of Arts and Sciences, Akdeniz University, Antalya, 07058, Turkey Published online: 15 Mar To cite this article: Aziz Asian, Mustafa Yavuz & Ali Erdogan (2005) A COMPARATIVE STUDY OF THE BREEDING ECOLOGY OF THE HOUSE SPARROW (PASSER DOMESTICUS L.): TIMING OF BREEDING AND BREEDING SUCCESS, Israel Journal of Zoology, 51:4, To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the Content ) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden.

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3 ISRAEL JOURNAL OF ZOOLOGY, Vol. 51, 2005, pp A COMPARATIVE STUDY OF THE BREEDING ECOLOGY OF THE HOUSE SPARROW (PASSER DOMESTICUS L.): TIMING OF BREEDING AND BREEDING SUCCESS Aziz Aslan,* Mustafa Yavuz, and Ali Erdoğan Department of Biology, Faculty of Arts and Sciences, Akdeniz University, Antalya 07058, Turkey ABSTRACT Breeding activity of the house sparrow (Passer domesticus) occurred between March and August; up to four successful clutches per pair were found in each breeding season. Timing of breeding activities and breeding patterns were compared for the years 2001 to Of the 656, 760, and 600 (total 2016) eggs laid in the three years, 334 (50.91%), 426 (56.05%), and 362 (60.33%) (total 55.65%) hatched; 224, 276, and 209 (total 709) nestlings fledged, giving a breeding success in terms of eggs laid of 34.2%, 36.1%, and 34.8% (total 35.2%); 67.1%, 64.8%, and 57.6% (63.2%), respectively, in terms of the eggs hatched. The mean number of fledglings per clutch was 2.70 ± 0.13, 2.42 ± 0.09, and 2.36 ± 0.11 (2.49 ± 0.06). The mean nestling weight at hatching and before fledgling (day 15) was 1.81 g ± 0.40 (n = 59) and g ± 0.30 (n = 94), respectively. These data provide a base against which future population changes can be monitored. INTRODUCTION The house sparrow (Passer domesticus L.) is common in Europe, North Africa, Asia, and in other parts of the world to which it has been introduced (viz., America, southern Africa, Australia, New Zealand, and some oceanic islands) (Summers-Smith, 1980; Böhner et al., 2003). Occurring in built-up areas, it is a very familiar bird species. In the last century, rapid industrialization and environmental changes have threatened the existence of many bird species (Kiziroğlu, 1987; Donald et al., 1994; Fuller et al., 1995; Chamberlain and Crick, 1999). However, the house sparrow, one of the most common birds in human-altered habitats, has been least affected by the environmental changes. With increases in the built-up habitat, numbers have increased over time. Howeever, this trend has been changing and recently there have been reports of house sparrow declines in many areas, especially in Europe (Summers-Smith, 1999; Bezzel, 2001; Crick et al., 2002; Böhner et al., 2003). The decline has been attributed to lack of food, predation, decreased availability of nesting sites and, hence, a reduction in breeding succcess (Mitschke et al., 2000; Siriwardena et al., 2002). In addition, the decline could also *Author to whom correspondence should be addressed. aaslan@akdeniz.edu.tr Accepted March 2006.

4 362 A. aslan et al. Isr. J. Zool. be the result of decreased survival rate. No decline of urban sparrows has been detected as yet in Turkey, and data on the current breeding success should provide a base against which future changes can be monitored. In the last two decades, uncontrolled tourism and agricultural activities in the Meditterranean region of Turkey have resulted in the reduction of natural habitat (Erdoğan et al., 2002; Aslan and Erdoğan, 2004). Urbanization and reduction of natural habitat have had a negative impact on the breeding success of many bird species and their populattions. This study was carried out to determine: (1) the timing of breeding activities, (2) reproduction success, and (3) population trends. MATERIALS and METHODS Antalya is located in the Mediterranean coastal region of Turkey at latitude N and longitude E. The city is situated in the bay of the same name at an average altitude of 51 m a.s.l. The study area, the regional Department of Forestry site, covers 2.25 hectaares and is located at the city center (Fig. 1). The site consists of office buildings, living quarters, play areas, soccer field, and arboretum, and is surrounded by trees, providing nesting and feeding opportunities for many songbird species. Sixty-two wooden nest-boxes, cm with a 3.5-cm entrance hole and 1.5 cm wall thickness, were hung on trees in the study site. Each nest-box, painted green Fig. 1. Geographical location of study area.

5 Vol. 51, 2005 breeding ecology of the house sparrow 363 and numbered with black paint, was monitored for three breeding seasons; each year the contents of the nest-boxes from previous years were emptied in February. During the breeding season, nests were checked twice a day (between and ) to determine the date of nest building, date of laying of the first egg, and duration of incubation and nestling periods. The date of building the nest was taken as the first day nest material was added to the nest box, the date of laying was the date of the first egg, and the date of fledging was the date the last chick left the nest (Erdoğan, 1989; Kaçar, 2001). Incubation period was taken as the time between laying the last egg of the clutch and hatching of the first egg, and the nestling period as the time between hatching and fledging of the first nestling (Singer and Yom-Tov, 1988; Hole et al., 2002; Pampus et al., 2005). In order to determine the number of clutches during the 2003 breeding season, 24 adult females were ringed with a unique combination of two or three colored plastic spiral rings (white, red, green, and blue). The adults were trapped with a mist net and released after being ringed (Erdoğan et al., 2003; Aslan et al., 2004). Rings were placed on both left and right legs of the females ( The nest-boxes, both those used by ringed and by other females, were checked twice a day to determine the breeding parameters (number of clutch, eggs laid, hatched and unhatched eggs, dead nestlings, fledglings, and duration of incubation and nestling periods). Statistical analysis Analyses to determine whether or not breeding parameters differed significantly between years and clutches were tested using ANOVA. Multiple Comparison Analysis (Duncan test) was also used to determine differences and similarities between years and clutches, and these results are shown in Tables 3 and 4. Groups represented by the same letter are statistically similar while the others are statistically different. Those repressented by more than one letter are similar to groups represented by the same letters. Effects of climatic factors (data were taken from Antalya Meteorology Station and presented in Appendix 1) on breeding activities and parameters and the strength of the relationship between breeding parameters was tested using the partial correlation test. The daily growth rate of nestlings was expressed by pearson correlation and a linear reggression graph was generated. To determine the population trends, the mean clutch size (CS), number of hatched (H) and unhatched (UH) eggs, and number of dead nestlings (DN) and fledglings (FL) were used to generate Pearson Correlation and linear regresssion graphs. Breeding success (BS) was determined using two different likelihood ratio tests. In the first method, breeding success (BS 1 = 100 FL/TEL) is defined as the ratio of the total number of fledglings to the total number of eggs laid and in the second, (BS 2 = 100 FL/H) as the ratio of the total number of fledglings to the total number of eggs hatched (Deckert, 1969; Kiziroğlu, 1981; Singer and Yom-Tov, 1988). Chi-square test was used to determine whether or not the difference in percentage yearly breeding success was statistically significant, and to compare the ratios of total number of hatched and unhatched eggs (H/UH), dead nestlings, and fledglings (DN/FL). A significance level of p < 0.05 was used for the statistical tests. For each test, degrees

6 364 A. aslan et al. Isr. J. Zool. of freedom (df) and significance level were reported. Data were analyzed using SPSS for Windows standard version (SPSS inc., ). RESULTS Timing of breeding activities and ringing results The mean for the date of the start of nest building was April 7, April 1, and March 28 in 2001, 2002, and 2003, respectively; for the date of the first egg was May 13, May 17, and May 19 with the start of incubation May 19, May 21, and May 25; and for the date of fledging June 21, June 13, and June 18. Mean, minimum, and maximum breeding activity dates for clutches and years are shown in Appendix 2. Nineteen (79.1%) of the 24 ringed females used the nest-boxes. Clutches of ringed females ratios and the three-year total ratios were compared; no difference was found (Table 1). Assuming that the clutches in each nest-box were laid by the same pair, the ratios in 2001, 2002, and 2003 were compared with three-year period ratios; no difference was found (Table 2). As seen in Table 2, the ratios of second and third clutches were not statistically differeent between years, whereas the ratio of fourth clutches was. When all nesting attempts were considered together, there was no significant difference in clutch ratios between years (χ 2 = ; df: 6; p > 0.40). Duration of Incubation and Nestling Period The mean incubation periods were 9.8 ± 0.20, 10.1 ± 0.17, 9.1 ± 0.20, and 9.7 ± 0.11 days; differences in the mean incubation period between clutches were statistically signifiicant in 2001, 2002, 2003, and the three-year period, respectively (p < 0.05) (Table 3). Differences in the mean incubation period between years were statistically significant (F 2, 318 = 7.57; p < 0.001); mean incubation periods in 2001 and 2002 were similar, while the incubation period in 2003 had the lowest mean value (Table 4). The mean nestling periods were 15.4 ± 0.18, 15.0 ± 0.17, 15.3 ± 0.17, and 15.2 ± 0.10 days and differences in the mean nestling periods between clutches in 2001 were not statistically significant (p > 0.05), but were in 2002, 2003, and the three-year period (p < 0.05) (Table 3). Diffferences in the mean nestling period between years were not statistically significant (F 2, 279 = 1.27; p > 0.28) (Table 4). Clutch Size The mean clutch sizes were 5.25 ± 0.12, 5.00 ± 0.11, 5.17 ± 0.12, and 5.13 ± 0.07 in 2001, 2002, 2003, and the three-year period, respectively. Differences in the mean clutch sizes between nesting attempts were statistically significant (p < 0.05) (Appendix 3), however, they were not significant between years (F 2,390 = 1.259; p > 0.29) (Table 4). There was a strong positive correlation between CS and H (r = 0.47; p < 0.009), FL (r = 0.59; p < 0.001), and UH (r = 0.74; p < ) in 2001; H (r = 0.77; p < ), DN (r = 0.40; p < 0.001), FL (r = 0.49; p < ), and UH (r = 0.62; p < ) in 2002; H

7 Vol. 51, 2005 breeding ecology of the house sparrow 365 Table 1 Comparison between nesting attempts, ratios of ringed pairs, and three-year period Nesting 2003 Ratio χ 2 Total Ratio χ 2 Attempt (n) (%) (N) (%) 1st nd ; df: 1; p > ; 3rd ; df: 1; p > df: 3; 4th ; df: 1; p > p > 0.90 n Clutch number of ringed females in 2003, N: three-year clutch numbers. Table 2 Comparison of nesting attempts between years and three-year period Nesting 2001 Ratio 2002 Ratio 2003 Ratio χ 2 Total Ratio χ 2 Attempt (n) (%) (n) (%) (n) (%) (N) (%) 1st ; 2nd ; df:2; df: 6; p > rd ; df:2; p > p > th ; df:2; p > 0.03 (r = 0.70; p < ) and DC (r = 0.63; p < ) in 2003; H (r = 0.70; p < ), FL (r = 0.49; p < ), and UH (r = 0.54; p < ), but only a weak correlation between CS and DN (r = 0.36; p < ) in the three-year period (Fig. 2). Hatched Eggs (= Nestlings) A total of 334 (50.9%), 426 (56.1%), 362 (60.3%), and 1122 (55.7%) eggs hatched; the mean number of nestlings per clutch was 3.59 ± 0.14, 3.41 ± 0.11, 3.66 ± 0.14, and 3.54 ± 0.07 in 2001, 2002, 2003, and the three-year period, respectively. Differences in the mean number of eggs hatched per clutch between nesting attempts in 2001, 2002, 2003, and the three-year period were statistically significant (p < 0.05) (Appendix 3), however, differences between years were not (F 2,314 = 1.098; p > 0.34) (Table 4). There was a strong positive correlation between H and FL (r = 0.73; p < ) in 2001; DN (r = 0.29; p > 0.12) and FL (r = 0.67; p < ) in 2002; DN (r = 0.46, p < 0.005) and FL (r = 0.73; p < ) in 2003; DN (r = 0.49, p < ) and FL ( r = 0.72; p < ) in the three-year period. In contrast, there was a negative correlation between H and UH (r = 0.49; p < 0.003) in 2003 and in the three-year period (r = 0.23; p < 0.008) (Fig. 3).

8 366 A. aslan et al. Isr. J. Zool. Table 3 Differences between duration of incubation and nestling period based on clutches D Year clutches ANOVA 1st 2nd 3rd 4th Total Incubation Period ± 0.39 b 8.90 ± 0.33 a 9.89 ± 0.25 ab 9.33 ± 0.56 ab 9.79 ± 0.20 F 3, 93 = (n: 34) (n: 30) (n: 27) (n: 6) (n: 97) p < ± 0.29 b 9.64 ± 0.28 ab 9.26 ± 0.32 a ± 0.54 ab ± 0.17 F 3, 121 = (n: 48) (n: 39) (n: 27) (n: 11) (n: 125) p < ± 0.37 b 8.71 ± 0.29 a 7.95 ± 0.27 a 8.71 ± 0.36 a 9.05 ± 0.20 F 3, 95 = (n: 37) (n: 34) (n: 21) (n: 7) (n: 99) p < Total ± 0.20 b 9.12 ± 0.18 a 9.12 ± 0.19 a 9.62 ± 0.33 a 9.66 ± 0.11 F 3, 317 = (n: 119) (n: 103) (n: 75) (n: 24) (n: 321) p < Nestling Period ± 0.27 a ± 0.40 a ± 0.30 a ± 0.49 a ± 0.18 F 3, 79 = (n: 26) (n: 25) (n: 26) (n: 6) (n: 83) p > ± 0.22 c ± 0.28 b ± 0.37 ab ± 0.54 a ± 0.17 F 3, 109 = (n: 44) (n: 34) (n: 24) (n: 11) (n: 113) p < ± 0.25 b ± 0.30 ab ± 0.31 ab ± 0.60 a ± 0.17 F 3, 82 = (n: 36) (n: 27) (n: 17) (n: 6) (n: 86) p < Total ± 0.14 c ± 0.18 b ± 0.20 b ± 0.32 a ± 0.10 F 3, 278 = (n: 106) (n: 86) (n: 67) (n: 23) (n: 282) p < D Duration of ıncubation and nestling period; n number of clutches

9 Vol. 51, 2005 breeding ecology of the house sparrow 367 Table 4 Differences between breeding parameters and duration of incubation and nestling period based on years BP ANOVA (Mean±SE) (Mean±SE) (Mean±SE) CS 5.25 ± 0.12 a 5.00 ± 0.11 a 5.17 ± 0.12 a F 2, 390 = 1.259; p > H 3.59 ± 0.14 a 3.41 ± 0.11 a 3.66 ± 0.14 a F 2, 314 = 1.098; p > DN 2.08 ± 0.16 ab 1.72 ± 0.10 a 2.19 ± 0.14 b F 2, 207 = 3.964; p < FL 2.70 ± 0.13 b 2.42 ± 0.09 ab 2.36 ± 0.11 a F 2, 277 = 2.482; p > UH 3.07 ± 0.18 b 2.39 ± 0.13 a 2.53 ± 0.18 a F 2, 336 = 5.269; p < **IP 9.79 ± 0.20 b ± 0.17 b 9.05 ± 0.20 a F 2, 318 = 7.569; p < **NP ± 0.18 a ± 0.17 a ± 0.17 a F 2, 279 = 1.268; p > BP breeding parameters; CS clutch size; H number of hatched eggs; DN number of dead nestlings; FL number of fledglings; IP duration of incubation period; NP duration of nestlling period. Fig. 2. Correlation between clutch size (CS) and numbers of hatched eggs (H), dead nestlings (DN), fledglings (FL), and unhatched eggs (UH).

10 368 A. aslan et al. Isr. J. Zool. Fig. 3. Correlation between hatched eggs (H) and number of dead nestlings (DN), fledglings (FL), and unhatched eggs (UH). Dead Nestlings A total number of 110 (32.9%), 150 (35.2%), 153 (42.3%), and 413 (36.8%) nestllings died, the mean number per clutch being 2.08 ± 0.16, 1.72 ± 0.10, 2.19 ± 0.14, and 1.97 ± 0.08 in 2001, 2002, 2003, and the three-year period, respectively. Differences in the mean number of dead nestlings per clutch between nesting attempts in each year and the three-year period were not significant (p > 0.05) (Appendix 3), though they were between years (F 2,207 = 3.964; p < 0.02) (Table 4). There was a strong negative correlation between DN and FL (r = 0.45; p < 0.013) and UH (r= 0.45; p < 0.013) in 2001; FL (r = 0.45; p < 0.013) in 2002; FL (r = 0.25; p < 0.004) in the three-year period (Fig. 4). No statistically significant correlation was found in Fledglings (= Brood size) A total number of 224 (67.1%), 276 (64.8%), 209 (57.7%), and 709 (63.2%) chicks flew with a mean number of fledglings per clutch of 2.70 ± 0.13, 2.42 ± 0.09, 2.36 ± 0.11, and 2.49 ± 0.06 in 2001, 2002, 2003, and the three-year period, respectively. Differences in the mean number of fledglings per clutch between nesting attempts in 2001 were not significant (p > 0.05), but were in 2002, 2003, and the three-year period (p < 0.05) (Apppendix 3). According to ANOVA, difference in the mean number of fledglings between years was not statistically significant (F 2, 277 = 2.482; p > 0.085), but in the Duncan test, 2001 had the highest number of fledglings and 2003 had the lowest number of fledgllings (Table 4). There was no statistically significant correlation between FL and UH in 2001 and 2002, but a negative correlation was found between FL and UH (r = 0.63; p < ) in 2003 and in the three-year period (r = 0.17; p < 0.044).

11 Vol. 51, 2005 breeding ecology of the house sparrow 369 Fig. 4. Correlation between number of dead nestlings (DN) and numbers of fledglings (FL) and unhatched eggs (UH). Fig. 5. Correlation between number of fledglings (FL) and number of unhatched eggs (UH).

12 370 A. aslan et al. Isr. J. Zool. Unhatched Eggs There were 322 (49.1%), 334 (44.0%), 238 (39.7%), and 894 (44.4%) unhatched eggs, with the mean number of unhatched eggs per clutch 3.07 ± 0.18, 2.39 ± 0.13, 2.53 ± 0.18, and 2.64 ± 0.09 in 2001, 2002, 2003, and the three-year period, respecttively. Differences in the mean number of unhatched eggs per clutch between nesting attempts in each year were not statistically significant (p > 0.05), but were in the threeyear period (p < 0.05) (Appendix 3). Differences in the mean number of unhatched eggs between years were statistically significant (F 2,336 = 5.269; p < 0.006); 2001 had the highest and 2003 had the lowest number of unhatched eggs (Table 4). Nestling growth Mean nestling weights at hatching and before fledgling (day 15) were 1.81 g ± 0.40 (n = 59) and g ± 0.30 (n = 94), respectively. Mean fledgling weight was 70.5% of mean adult weight (29.9 ± 0.53, n = 42). Nestling growth rate between day 5 and day 17 after hatching is shown in Fig. 6. The correlation coefficient between age in days and nestling weight was r = 0.82, p < (N = 1173). Effects of climatic factors (temperature, rainfall, and humidity) In the three-year period, there were significant negative correlations between ambieent temperature and CS (r = 0.82, p < 0.002), H (r = 0.80, p < 0.003), UH (r = 0.75, p < 0.008), IP (r = 0.60, p < 0.052), and NP (r = 0.87, p < 0.001). In contrast, there were positive correlations between rainfall and CS (r = 0.64, p < 0.032), H (r = 0.65, Fig. 6. Nestling growth rate from day 5 to day 17 in the three-year period.

13 Vol. 51, 2005 breeding ecology of the house sparrow 371 p < 0.032), UH (r = 0.77, p < 0.006), IP (r = 0.60, p < 0.052), and NP (r = 0.78, p < 0.005), and between humidity and IP (r = 0.69, p < 0.020). There were significant negattive correlations between humidity and date of nest building (r = 0.63, p < 0.039), and between rainfall and both date of laying first egg (r = 0.88, p < ) and fledging (r = 0.84, p < 0.001). In contrast, there were significant positive correlations between ambient temperature and both date of laying first egg (r = 0.99, p < ) and fledging (r = 0.98, p < ) Breeding Success Breeding success (BS 1 ) in 2001, 2002, 2003, and in the three-year period was 34.2%, 36.1%, 34.8%, and 35.2%; differences in breeding success between years were not statistically significant (χ 2 = , df: 2; p > 0.90). Breeding success according to the second definition (BS 2 ) was 67.1%, 64.8%, 57.6%, and 63.2%, in 2001, 2002, 2003, and the three-year period, respectively. Differences in success between years were not statistically significant (χ 2 = , df: 2; p > 0.40) (Table 5). Differences in breeding success between years based on H/UH and DN/FL, the ratio of total number of eggs hatched to total number of eggs that failed to hatch (χ 2 = 8.02; df: 2; p < 0.05) and between total number of dead nestlings to total number of fledglings (χ 2 = 6.46; df: 2; p < 0.05) were not statistically significant (Table 6). Table 5 Differences between breeding success ratios based on years and three-year period Breeding 2001 Ratio 2002 Ratio 2003 Ratio Total Ratio χ 2 success (n) (%) (n) (%) (n) (%) (N) (%) BS , df: 2; p > 0.90 BS , df: 2; p > 0.40 n total number of laid and hatched eggs in each year; N total number in three-year period. Table 6 Significance between breeding parameters based on years Parameters 2001 Ratio 2002 Ratio 2003 Ratio Total Ratio χ 2 (n) (%) (n) (%) (n) (%) (N) (%) H ; df: 2; UH p < 0.05 DN ; df: 2; FL p < 0.05 n total number of hatched (H) and unhatched eggs(uh), dead nestlings (DN), and fledglings (FL), in each year; N total number in three-year period.

14 372 A. aslan et al. Isr. J. Zool. DISCUSSION Haartmann (1969), Alatalo (1975), Erdoğan and Kiziroğlu (1990), Sıkı (1992), and Hole et al. (2002) reported the number of clutches as 2 3 per season. However, house sparrows had four successful breeding attempts in Israel and Iraq (Al-Dabbagh and Jiad, 1988; Singer and Yom-Tov, 1988). The 24 pairs in this study in which the females were ringed laid four clutches, and the number of successful attempts by these pairs in 2003 was similar in each of the years and the three-year period. This result is important since it shows that the species laid 4 successful clutches in our study area. The length of the breeding period was reported as months (Seel, 1969; Hyla, 1971; Alatalo, 1975; Hole et al., 2002), but was 6 months in this study, the same as that in Israel and Iraq (Al- Dabbagh and Jiad, 1988; Singer and Yom-Tov, 1988). According to Erdoğan (1989), the date of the first egg was May 14 for the first breeding attempt, June 12 for the second, and July 20 for the third. There were three nesting attempts in England between May and July (Hole et al., 2002). In this study the first clutches were laid in April, the second in May, the third in June, and the fourth in July, and the date of nesting attempts did not vary from year to year (see Appendix 2). Duration of incubation and nestling period for the house sparrow generally varies between 9 12 and days, respectively (Fitter and Richardson, 1963; Haartmann, 1969; Alatalo, 1975; Erdoğan, 1989; Sıkı, 1992); those obtained in this study were the same as those found in previous studies. It has been reported that the mean length of these periods declines with an increase in ambient temperature (Fitter and Richardson, 1963; Seel, 1968b; Dawson, 1972; Novotny, 1970; Yom-Tov and Ar, 1980; Kiziroğlu, 1981; Erdoğan, 1989; Sıkı, 1992). In the present study, the length of the incubation perriod varied between nesting attempts and years, but nestling period length did not. The mean clutch size was 5.13 eggs in this study population (36 N), which is similar to the clutch size of 5.05 eggs in Israel (32 N) (Singer and Yom-Tov, 1988), in Kansas (39 N) and Alberta, Canada (51 N) (Murphy, 1978b), but greater than the <5 eggs in Iraq (33 N) (Al-Dabbah and Jiad, 1988), in most of central Europe (Dyer et al., 1977), and in Britain (Summers-Smith, 1963; Seel, 1968b). Mean egg numbers in the first and second clutches in this study were 5.42 and 5.45; other studies had 4.4 and 4.7 eggs (Erdoğan, 1989) and 4.57 and 4.85 eggs (Balát, 1974). According to Dyer et al. (1977), clutch size tended to increase from south to north in both eastern and western hemispheres. Weather conditions such as air temperature are known to influence food availability (Perrins and Birkhead, 1983); differences between nesting attempts and the higher mean values of clutch size per year found in this study could be related to availaability of food. Hatching ratio in the three-year period was 55.7%; this is lower than in Israel (70%; Singer and Yom-Tov, 1988), Iraq (72%; Al-Dabbah and Jiad, 1988), in New Zealand, India, Europe, and North America (73%; Dyer et al., 1977), but the number hatched per clutch (3.54) was similar to that in Israel, 3.90, and Iraq, Furthermore, the mean values of nestlings per clutch were also lower than those found by Erdoğan (1989), Kiziroğlu et al. (1987), Balát (1974), and Haartmann (1969). In all years, the number of

15 Vol. 51, 2005 breeding ecology of the house sparrow 373 nestlings decreased from the first to the fourth clutch. Mortality rate and mean number of dead per clutch in this study were 36.8% and 1.97 in the three-year period; this is higher than in Israel, 25.4% and 0.9, and Iraq, 19.5% and 1.55 (Al-Dabbah and Jiad, 1988; Singer and Yom-Tov, 1988), but lower than in Izmir, Turkey, 46.8% (Sıkı, 1992). According to Erdoğan (1989), the number of fledglings was the same as the number hatched (in Ankara); however, there were dead nestlings during all nesting attempts and years in this study. Variability of environmental conditions and predation level in each year could be the reason for the difference in the number of dead nestling between years. The number of fledglings is the most important factor in determining breeding succcess or change in direction of population size (Deckert, 1969; Kiziroğlu, 1981). The mean number of fledglings per clutch and fledgling success in the present study were 2.49 and 63.2% in the three-year period, which is similar to the results reported in Iraq (2.19, 58.5%) by Al-Dabbah and Jiad (1988) and in Israel (2.64, 74.6%) by Singer and Yom-Tov (1988), but lower than reported in Europe by Seel (1969). The mean numbers of fledglings from first and second clutches in previous studies were 4.29 and 5.03, respectively (Erdoğan, 1989), and 4.5 each year (Kiziroğlu et al., 1987), higher than those found in this study (see Table 4). It is a matter of concern if the mean number of fledglings is declining compared to previous studies in Turkey and Europe. The mean percentage and number of unhatched eggs per clutch in the present study (44.4%, 2.64) were higher than those found both in Israel (29.9%, 1.51) and Iraq (28.0%, 1.34) (Al-Dabbah and Jiad, 1988; Singer and Yom-Tov, 1988). Mean proportion of unhhatched eggs in first and second clutches were found to be 17.9% and 7.1%, respectively, by Erdoğan (1989), and 7.84% annually by Sıkı (1992), which is lower than our results. Differences in the ratios of unhatched eggs between years were due to infertility and desertion (eggs containing dead embryos). A strong positive correlation between the mean of CS and H, DN, FL, and UH in this study means that a positive or negative change in the clutch size will affect the other parrameters as well as breeding success. A positive correlation between eggs hatched, dead nestlings, and fledglings was observed on a yearly and a three-year basis. An increase or decrease in the hatched eggs affects the number of dead nestlings and fledglings. In contrast, no statistically significant correlation between hatched and unhatched eggs was observed in a year or between years since these parameters are independent of each other. There was a negative correlation between dead nestlings and fledglings since an increase or decrease in one parameter will result in a similar change in the other. The weight at fledging is known to influence post-fledging survival (Lack, 1966). Murphy (1978 b) showed that post-fledging survival of young house sparrows was corrrelated with their weight at fledging. Mean nestling weight at hatching in this study was higher, but fledgling weight (day 15) was lower than those reported by Singer and Yom- Tov (1988) and Erdoğan (1989). In addition, mean weight on the day before fledging was 70% of adult weight, and this is lower than the values of 90% given by Singer and Yom-Tov (1988) and 84% by Erdoğan (1989). Post-fledging survival was not followed in this study; thus, the effect of fledgling weight on the survival rate is unknown, but in

16 374 A. aslan et al. Isr. J. Zool. spite of this, it can be concluded that the lower breeding success in this study might be correlated with the lower nestling and fledgling weight. Ecological factors like food availability, weather, and predator pressure are known to influence life history and reproductive success (Lack, 1968). Seel (1968a) showed that air temperature was correlated with the start of breeding in England, and Murphy (1978a) also confirmed this conclusion in house sparrow populations in North America. Climatic factors (temperature, rainfall, and humidity) affected both the initiation of the breeding season and breeding patterns in this study. It can be argued that the breeding activities and patterns of the species were affected by variations in climatic factors within and between years. Breeding success ratios in this study based on BS 1 are lower than the average of 42% for 13 populations calculated from Appendix 3.7 in Kendeigh and Pinowski (1977), 52.3% in Israel by Singer and Yom-Tov (1988), and in the first two nesting attempts, namely 82.1% and 92.9%, in Ankara, Turkey (Erdoğan, 1989). According to the second definition (BS 2 ), breeding success is lower than the average of 100% by Erdoğan (1989) and 78.9% by Balát (1974), but higher than 53% by Sıkı (1992) and 35% around Oxfford, England, by Seel (1969). As seen, the breeding success based on both methods in our study was lower. However, in this species there is much inter-pair, inter-site, and inter-year variation in clutch sizes and breeding success (Murphy, 1978a,b; McGillivray, 1983) and it is difficult to draw overall conclusions from results obtained during a threeyear study. In conclusion, the results obtained in the present study when compared to previous studies point to one main conclusion: the breeding performance of the house sparrow population in the study area remained the same in the three year period. In conttrast to other studies in Turkey, it is worth noting that the breeding success of the house sparrow population is declining despite the increase in the number of breeding attempts. In addition, in light of the post-fledging population (1 year period) decline suggested by some researchers (Summers-Smith, 1999; Freeman and Crick, 2002; Hole et al., 2002; Siriwardena et al., 2002; Böhner et al., 2003), the population of the house sparrow in the study area and in Turkey must be studied and monitored for a longer period to obtain conclusive results. ACKNOWLEDGMENTS This study was supported by the Akdeniz University Research Fund. We thank D. Summmers-Smith, Y. Yom-Tov, and J. Pinovski for valuable comments on the manuscript. We also wish to thank numerous students who helped collect the data and, for correcting the manuscript, Orhan Ünal for the earlier versions, and D. Summers-Smith for the final version of the manuscript. REFERENCES Alatalo, R On the breeding biology of the House Sparrow at Oulu. Lint. 10: 1 7. Al-Dabbagh, K.Y., Jiad, J.H The breeding biology of the House Sparrow in central Iraq.

17 Vol. 51, 2005 breeding ecology of the house sparrow 375 International Studies on Sparrows 15: Aslan, A., Erdoğan, A The distribution of the white-spectacled bulbul (Pycnonotus xanthoppygos) and influential factors on its distribution in Turkey. 1st International Euroasian Ornitthology Congress, 8 11 April 2004, Antalya, Turkey. Abstract book, 11 pp. (unpublished). Aslan, A., Erdoğan, A., Albayrak, T., Tunç, M.R Birds of Antalya and ringing studies. Nature and Man 2: Balát, F Gelegegrosse und Brutverlust des Haussperlings, Passer domesticus L. İn Mitttelmahren. Zool. Listy. 23: Bezzel, E Does only sparrow survive? Birds in the exploited landscape Journal Für Ornithologie 142: Böhner, J., Schulz, W., Witt, K Bestand und Abundanz des Houssperlings (Passer domestticus) in Berlin Berl. Ornithol. Ber. 13: Chamberlain, D.E., Crick, H.Q.P Population declines and reproductive performance of skylarks (Alauda arvensis) in different regions and habitats of the United Kingdom. Ibis 141: Crick, H.Q.P., Robinson, R.A., Appleton, G.F., Clark, N.A., Rickard, A.D., eds Investiggation into the causes of the decline of starlings and House sparrows in Great Britain. BTO Research Report. 290 pp. Dawson, D.G The breeding ecology of house sparrows. Ph.D. thesis, Univ. Oxford, Oxfford. Deckert, G Zur Ethologie und Okologie des Haussperlings. Beitraege zur Vogelkunde. 15:1 84. Donald, P. F., Wilson, J.D., Stepherd, M The decline of the corn bunting in Britain. British Birds 87: Dyer, M.I., Pinowski, J., Pinowska, B Population dynamics. In: Pinowski, J., Kendeigh, S.C., eds. Granivorous birds in ecosystems. Cambridge Univ. Press, London, pp Erdoğan, A Ankara/Beytepe Serçe Populasyonları (Passer domesticus L. ve Passer monttanus L.) ile İlgili Biyolojik Çalışmalar. Yüksek Lisans tezi, G.Ü. Fen Bilimleri Enst., Ankara, 97 pp. Erdoğan, A., Kiziroğlu, İ Ankara/Beytepe Serçe Populasyonları (Passer domesticus L. ve Passer montanus L., Passeridae; Aves) Kuluçka Biyolojisi Üzerine Çalışmalar. X. Ulusal Biyoloji Kongresi, Kongre kitabı, Erzurum, pp Erdoğan, A., Albayrak, T., Sert, H., Aslan, A., Tunç, M.R Boğazkent Kocagöl ve Çevresi Kuş Envanteri Hazırlama Projesi Sonuç Raporu, Antalya, 254 pp. Erdoğan, A., Sert, H., Vohwinkel, R., Prünte, W., Albayrak, T., Aslan, A., Tunç, M.R Manavgat/Titreyengöl Kuş Halkalama Çalışmaları. Doǧa ve Insan (Nature and Man) 1: (in Turkish). Fitter, R.S.R., Richardson, A.R The pocket guide to nests and eggs. London, 172 pp. Freeman, S.N., Crick, H.Q.P Population dynamics of House Sparrows Passer domesticcus breeding in Britain: an integrated analysis. In: Crick, H.Q.P., Robinson, R.A., Appleton, G.F., Clark, N.A., Rickard, A.D., eds. Investigation into the causes of the decline of Starlings and House Sparrows in Great Britain. BTO Research Report No 290, DEFRA, Bristol, pp Fuller, R.J., Gregory, R.D., Gibbons, D.W., Marchant, J.H., Wilson, J.D., Carter, N Populattion declines and range contractions among lowland farmland birds in Britain. Conservation Biology 9: Haartmann, L.V The nesting habits of Finnish birds I. Passeriformes. Com. Biological

18 376 A. aslan et al. Isr. J. Zool. Vol. 32: Hole, D.G., Whittingham, M.J., Bradbury, R.B., Wilson, J.D Comparative breeding ecologgy of the house sparrow Passer domesticus before and during population decline in Britain. In: Crick, H.Q.P., Robinson, R.A., Appleton, G.F., Clark, N.A., Rickard, A.D., eds. Investigation into the causes of the decline of starlings and House sparrows in Great Britain. BTO Research Report No 290, DEFRA, Bristol, pp Hyla, W Boabachtungen an der Houssperlings population (Passer domesticus) einer Grossttadt. Charadrius 7: Kaçar, S Antalya Araştırma Ormanlarındaki (BÜK-Lütfi Büyükyıldırım ve Elmalı Seddir) Bahçe Kızılkuyruğu (Phoenicurus phoenicurus L.,) Populasyonları Üzerine Biyolojik Araştırmalar. Yüksek Lisans Tezi, Akdeniz Üniv., Fen Bilimleri Enst., Antalya, 100 pp. Kendeigh, S.C., Pinowski, J Introduction. In: Pinowski, J., Kendeigh, S.C., eds. Granivorrous birds in ecosystems. Cambridge Univ. Press, London, pp Kiziroğlu, İ Ankara Beynam Ormanı ndaki baştankara, Parus L., cinsi (Aves) türlerinin biyoloji, ekoloji ve davranışları ile ilgili araştırmalar. TÜBİTAK, TBAG-371, 216 pp. Kiziroğlu, İ Sayıları Giderek Azalan Yırtıcı Kuşlarımız. Bilim ve Teknik 222: 1 4. Kiziroğlu, İ., Şişli, N., Alp, Ü Zur Interspezifischen Konkurrenz in der Besiedlung von Nistkasten bei Ankara/Türkei, mit Angaben zur Brutbiologie verschiedener Höhlenbrüter- Arten. Die Wogelwelt 198: Lack, D Population studies of birds. Clarendon Press, Oxford. Lack, D Ecological adaptations for breeding in birds. Chapman and Hall, London. McGillivray, W.B Intraseasonal reproductive costs for the house sparrow (Passer domestticus). Auk 101: Mitschke, A., Rathjen, H., Baumung, S House Sparrows in Hamburg: population, habitat choice and threats. Hamburg State Ornithological Protection Station. Hamburger Avifauna Beitr., 30. Murphy, E.C. 1978a. Breeding ecology of the house sparrow: spatial variation. Condor 80: Murphy, E.C. 1978b. Seasonal variation in reproductive output of the house sparrow: The determmination of clutch size. Ecology 59: Novotny, I Breeding bionomy, growth and development of young house sparrow (Passer domesticus (L) 1758). Acta Scientiarum Naturalium Academiae Scientiarum Bohemoslovacae (Brno) 4: Pampus, M., Schmidt, K-H., Wiltschko, W Pair bond and breeding success in Blue Tits Parus caeruleus and great tits Parus major. Ibis 147: Perrins, C.M., Birkhead, T.R Avian ecology. Blackie, Glasgow. Seel, D.C. 1968a. Breeding season of the house sparrow and tree sparrow Passer spp. at Oxford. Ibis 110: Seel, D.C. 1968b. Clutch size, incubation and hatching success in House Sparrow and tree sparrow Passer spp. at Oxford. Ibis 110: Seel, D.C Food, feeding rates and body temperatures in the nestling House S1parrow (Passer domesticus) at Oxford. Ibis 111: Sıkı, M Ev Serçesi (Passer domesticus L.) nin Üreme Biyolojisi Üzerine Araştırmalar. Doğa. Turkish Journal of Zoology 16: Singer, R., Yom-Tov, Y The breeding biology of the House Sparrow Passer domesticus in Israel. Ornis Scandinavica 19: Siriwardena, G.M., Robinson, R.A., Crick, H.Q.P Status and population trends of the House

19 Vol. 51, 2005 breeding ecology of the house sparrow 377 Sparrow Passer domesticus in Great Britain. In: Crick, H.Q.P., Robinson, R.A., Appleton, G.F., Clark, N.A., Rickard, A.D., eds. Investigation into the causes of the decline of starlings and House sparrows in Great Britain. BTO Research Report No 290, DEFRA, Bristol, pp Summers-Smith, D The House Sparrow. Collins, London. Summers-Smith, D House Sparrows down coal mines. British Birds 73: Summers-Smith, D Current status of the House Sparrow in Britain. British Wildlife. 10: Yom-Tov, Y., Ar, A On the breeding ecology of the Dead Sea Sparrow, Passer moabiticus. Israel Journal Zoology 29:

20 378 A. aslan et al. Isr. J. Zool. Appendix 1 Mean monthly temperature, rainfall, and humidity in the study area Climatic Year Months Factors J F Mr Ap My Jn Jy Ag Sp O N D Temp. ( C) Rain (mm) Humidity (%)

21 Vol. 51, 2005 breeding ecology of the house sparrow 379 Appendix 2 Timing of breeding activities of the house sparrow in the study area TBA Year Clutches Total 1st 2nd 3rd 4th Mean date of bulding nest n: n: 53 Min: Min: Max: Max: n: n: 57 Min: Min: Max: Max: n: n: 50 Min: Min: Max: Max: Mean date of first egg n: n: n: n: n: 127 Min: Min: Min: Min: Min: Max: Mx: Max: Max: Max: n: n: n: n: n: 152 Min: Min: Min: Min: Min: Max: Max: Max: Max: Max: n: n: n: n: n: 117 Min: Min: Min: Min: Min: Max: Max: Max: Max: Max: Mean date of leaving nest n: n: n: n: n: 82 Min: Min: Min: Min: Min: Max: Max: Max: Max: Max: n: n: n: n: n: 115 Min: Min: Min: Min: Min: Max: Max: Max: Max: Max: n: n: n: n: n: 86 Min: Min: Min: Min: Min: Max: Max: Max: Max: Max: TBA: timing of breeding activities, n: number of clutches

22 380 A. aslan et al. Isr. J. Zool. Appendix 3 Significance between breeding parameters based on nesting attempts Yr. Clutch clutch size Hatched Dead nestlings Fledglings Unhatched *n Mean ± SE n Mean ± SE n Mean ± SE n Mean ± SE n Mean ± SE st ± 0.18 b ± 0.26ab ± 0.28 a ± 0.24 a ± 0.29 a 2nd ± 0.16 b ± 0.22 b ± 0.30 a ± 0.22 a ± 0.31 a 3rd ± 0.27 b ± 0.23ab ± 0.24 a ± 0.22 a ± 0.28 a 4th ± 0.48 a ± 0.48 a ± 0.00 a ± 0.50 a ± 0.58 a Total ± ± ± ± ± 0.18 ANOVA F = F = F = F = F = p < p > p > p > p > st ± 0.15 c ± 0.18 b ± 0.15 a ± 0.16 b ± 0.22 a 2nd ± 0.21 c ± 0.15 b ± 0.19 a ± 0.17 b ± 0.26 a 3rd ± 0.23 b ± 0.22 a ± 0.19 a ± 0.13 a ± 0.21 a 4th ± 0.27 a ± 0.24 a ± 0.30 a ± 0.18 a ± 0.25 a Total ± ± ± ± ± 0.13 ANOVA F = F = F = F = F = p < p < p > p < p > st ± 0.22 b ± 0.25 a ± 0.22 a ± 0.19bc ± 0.37 b 2 nd ± 0.19 b ± 0.27 a ± 0.27 a ± 0.20 c ± 0.28ab 3 rd ± 0.19 a ± 0.21 a ± 0.25 a ± 0.16ab ± 0.27ab 4 th ± 0.20 a ± 0.44 a ± 0.54 a ± 0.25 a ± 0.33 a Total ± ± ± ± ± 0.18 ANOVA F = F = F = F = F = p < p > p > p < p > years 1 st ± 0.11 c ± 0.13 b ± 0.12 a ± 0.11 b ± 0.17 b 2 nd ± 0.11 c ± 0.13 b ± 0.15 a ± 0.11 b ± 0.16 b 3 rd ± 0.14 b ± 0.13 a ± 0.13 a ± 0.11 a ± 0.14 b 4 th ± 0.19 a ± 0.20 a ± 0.27 a ± 0.18 a ± 0.18 a Total ± ± ± ± ± 0.09 ANOVA F = F = F = F = F = p < p < p > p < p < *n: number of clutches.