<<

Use the following type of citation: North-western Journal of Zoology 2021: e212601

Paper Submitted to The North-Western Journal of Zoology

1 *Handling editor: Kornélia Kurucz

2 *Manuscript Domain: Ornithology

3 *Manuscript code: NwJZ_20_OR_02

4 *Submission date: 29_03_2020

5 *Revised: 28_12_2020

6 *Accepted / Rejected: 07_01_2021

7 *No. of words (without abstract, acknowledgement, references, tables, captions): 8 (papers under 700 words are not accepted)

9 *Editors only: 10

11 Title of the paper: Nest-site selection by deserti in deserts of southern Iran

12 Running head : Nest-site selection by desert

13 Authors: Davood PAKNIAT, Mohsen AHMADI, Gilda SHAHNASERI, Mahmoud-Reza

14 HEMAMI

15 Key Words: Arid environment, ground-nesting , nest microclimate, nest structure, nest

16 entrance orientation

17 No. of Tables: 2

18 No. of Figures: 2 North-western Journal of Zoology 19 No. of Files: 1 Accepted paper until proofing 20 Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-2

21 {NEST-SITE SELECTION BY AMMOMANES DESERTI OF SOUTHERN IRAN}

22 {Davood, PAKNIAT} 1, {Mohsen, AHMADI} 1, {Gilda, SHAHNASERI} 1 and {Mahmoud-

23 Reza HEMAMI} 1

24 1. Department of Natural Resources, Isfahan University of Technology, Isfahan, Iran

25 * Corresponding authors name and email address: Mahmoud-Reza HEMAMI,

26 [email protected]

27 Abstract. We evaluated the effect of habitat characteristics on nest-site selection of desert

28 lark (Ammomanes deserti) in southern Iran. Habitat variables were measured within a 1- and a

29 4-m radius circle plots around 20 active nests and 20 randomly selected control locations. All

30 the detected nests were located beside a big stone with a maximum weight of 6.5 kg. The

31 percentage cover of <50mm sized gravel and non-vegetated areas within both plot sizes as

32 well as the percentage cover of ≥50mm sized gravel in 4-m radius plots were the most

33 important variables affecting nest-site selection by desert lark. About 40% of the nests were

34 built on flat plains, 30% on eastern domains, 15% on southeastern domains, and 10% on

35 southern domains. The majority of the nests’ entrances were oriented toward the east (50%)

36 and the south (25%) directions to reduce fluctuations in the nests’ microclimate from adverse

37 wind and direct exposure to sunlight. Our results revealed that desert lark’s nest-sites are

38 mainly characterisedNorth-western by the availability of bigJournal stones, a ppropriatenessof Zoology of geographical aspect, Accepted paper until proofing 39 the internal slope of the stone, and nest entrance orientation. Environmental characteristics

40 contributing to stabilising nest microclimate are the most important variables influencing the

41 pattern of nest-site selection by desert lark.

42 Key Words: Arid environment, ground-nesting birds, nest microclimate, nest structure, nest

43 entrance orientation

44 Running title: Nest-site selection by desert lark

45 Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-3

46 Introduction

47 The assessment of nest-site selection is important due to the impact of habitat variables on

48 the reproductive success of birds. From an evolutionary perspective, selecting habitat and

49 nest site should bring about increasing reproduction success and survival rate of birds (Smith

50 et al 2003; Hosseini-Moosavi et al 2017).

51 The exact location and the shape and the size of ’s nest have evolved to provide a

52 suitable micro-environment with regard to temperature and humidity, a secure environment

53 for egg-laying, and hatchlings (Hansell 2000; Ar & Sidis 2002; Ardia et al. 2006; Burton

54 2006), and in some birds signal the phenotypic quality of the builder (Mainwaring et al.

55 2014).

56 Microclimate variables such as wind, rain and solar radiation can influence the nest site

57 selection behaviour of birds (Hartman and Oring 2003; Mezquida 2004; Burton 2006; Schaaf

58 et al. 2018b). Nest entrance orientation is one of the most salient determinants of the nest’s

59 microenvironment (Rauter et al. 2002; Burton 2006). Recent studies have shown that nest

60 entrance orientation is largely influenced by latitude, solar radiation (Burton 2006, Schaaf et

61 al. 2018a, Carroll et al 2020), nest-site vegetation (Schaaf 2020a, Hoekman et al. 2002),

62 topography (Porneluzi 2011), and prevailing winds (Long et al. 2009). The nest’s

63 microenvironment/microclimateNorth-western has a significant Journal impact of onZoology the cost of reproduction and Accepted paper until proofing 64 offspring development. For instance, nest’s temperature affects the viability of eggs (Webb

65 & King 1983; Cook et al. 2003, Heenan 2013), energy balance during incubation (Vleck

66 1981), fledging success and clutch sizes (Boer 2018) and nestling development (Starck &

67 Ricklefs 1998; Ardia 2005). Decreasing nest temperature fluctuations through suitable

68 selection of nest entrance position can significantly increase reproductive success in harsh

69 environments with low vegetation cover (With & Webb 1993; Hoekman et al. 2002;

70 Hartman & Oring 2003, Carroll et al 2020). Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-4

71 Habitat and nest-site selection occur through a hierarchical process (Kotliar & Wiens 1990,

72 Jones 2001, Harris et al 2020). The finest level is referred to as nest patch: characteristics of

73 the habitat patch enclosing the nest (Martin & Roper 1988). Studies by Petersen and Best

74 (1985) on Sage Sparrow (Amphispiza belli), and Martin and Roper (1988) on Hermit Thrush

75 (Catharus guttatus) suggested that nest patch lies within a radius of 5 m (area = 78.5 m2)

76 from the nest site. In other studies, however, nest patch size was determined based on the

77 changes in the condition of variables around nest sites (Petit et al. 1988; Holway 1991;

78 Knopf & Sedgwick 1992; With 1994). Although building a nest made of gravel consumes

79 plenty of time and energy (Afik et al. 1991; Shkedy & Safriel 1992), the nest would be

80 prevented from slipping down slopes (Richardson 1965), protected from wind and rain

81 damage (Ferguson-Lees 1960), camouflaged (Etchécopar & Hüe 1967) and structurally

82 sstabilised (Afik et al. 1991, Aznar et al 2016).

83 Desert lark (Ammomanes deserti) is found in arid steppes and semi-desert regions (Vaurie

84 1951) of 37 countries extending from West Asia and the Middle East to North Africa

85 (Birdlife International 2016). The species breeding season is from early March to late June.

86 They build cup-like nests on the ground near big stones and braid a rampart enclosing the

87 nest (Orr 1970) (Fig. 1). Desert lark is a common species of semi-desert steppes of central

88 and southern Iran andNorth-western represents a typical behaviourJournal of of nest Zoology selection by a wide spectrum of Accepted paper until proofing 89 ground-nesting species in such ecosystems. Accordingly studying patterns of nest-site

90 selection by this species provides more insights into the breeding behaviour of desert birds.

91 We evaluated the effect of habitat characteristics on the nest-site selection of desert lark

92 (Ammomanes deserti) in southern Iran. We hypothesised that nest-site selection by desert lark

93 is not random and the species select sites that provide optimal microclimate for chicks’

94 development. We also expected that the species further regulates the nest’s microclimate by

95 nest architecture and construction materials. Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-5

96 Material and Methods

97 Study area

98 This study was carried out in an area of approximately 4000 ha in rugged steppes of Zarrin

99 Dasht City, southern Iran (25" 2 28 to 40" 7 28 north latitude and 15" 33 54 to 19" 43

100 54 east longitude). The majority of the landscape is dominated by dry climates.

101 Field surveys and data collection were performed from 21st March to 15th May 2015 at the

102 peak of the species breeding period. Weather parameters at the time of sampling were

103 obtained from Fars Province synoptic weather station (http://www.farsmet.ir; 28 21 54" N,

104 54 25 44" E). During field investigations, minimum and maximum temperatures were 9.8° C

105 and 33.4° C, respectively (mean of 13.8° C). Strong prevailing wind often occurs during the

106 second half of the day. Nearly 35% of the wind pattern was towards the west and southwest.

107 The wind was calm, less than 0.51 meters per second, on 44% of the days (Fig. 2). The most

108 frequent plant species in the region are Artemisia spp., Fagonia spp., Convolvulus spp.,

109 Astragalus spp., and Salsola species.

110 Data collection

111 To find desert lark’s nests, thirty transects with 500 m long, and 800 m distance in between

112 were systematically surveyed. In total, 20 desert lark’s nests were found. Data collected for North-western Journal of Zoology 113 each nest consisted of geographicalAccepted position paper (latitude until and proofinglongitude), elevation (m.a.s.l.), time 114 and date, the number, weight, and size of pebbles used in fencing the nest, and internal slope

115 of the rock beside the nest. We generated a similar number of random points (control

116 locations) within the area surveyed for the desert lark nests in ArcMap 10.3. Subsequently,

117 these random points were located in the field and checked for the absence of the species nests;

118 no random point was located in vicinity of the detected nests. Two circle plots with radii of 1

119 and 4 m were established around each of the detected nests (occupied plot) and the random

120 points (unoccupied plot). An array of habitat variables was measured in each of these plots Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-6

121 including the percent slope, geographical aspect, the nest entrance orientation, the percentage

122 cover of vegetation, the percentage cover of ground components (≤ 50 mm and > 50 mm sized

123 gravel), and the percentage of bare soil. All variables were measured by one observer to

124 minimise the risk of errors.

125 Data analysis

126 Mann-Whitney test was used, to compare the mean of the variables measured in the nest

127 presence and absence plots of Ammomanes deserti. The effect of plot size in determining the

128 effect of habitat variables was evaluated using the Wilcoxon test. We used Chi-square test to

129 address the hypothesis that the orientation of domains and nest entrances were not randomly

130 selected. Data analysis was performed using SPSS version 21.

131

132 Results

133 The number of pebbles used to fence the nest entrance was found to range between 53 and

134 292 (mean of 119.85 ± 56.87 SD). The internal slope of the main overhanging stones

135 shadowing the nests ranged between 30 and 70 degrees with a mean of 47.55± 12.20 SD.

136 The nests were located between 828 and 900 m.a.s.l. with a mean of 875.9 ± 18.4 SD. Table 1

137 gives a brief descriptionNorth-western of the size of gravels Journal used to fence of theZoology nest entrance. Accepted paper until proofing

138 The results of the Mann-Whitney test showed that the percentage gravel with the size ≤ 50

139 mm and the percentage of bare soil within both plots as well as the percentage gravel with the

140 size <50 mm in 4 m-radius plots were significantly different between the occupied and

141 unoccupied plots (Table 2). Furthermore, mean percentage gravel within the 1 m-radius plots

142 was significantly greater than those found in 4 m-radius plots (Wilcoxon Z = -2.140, p =

143 0.032) indicating that desert ’ nests are located in places with higher amount of gravels Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-7

144 compared to the immediate adjacent sites. However, comparison of bare soil between the two

145 small and large plots did not show a significant difference (Wilcoxon Z = -1.027, p = 0.304).

146 The results of the Chi-square test showed that the geographical aspect of the nesting area and

147 the nest entrance orientation were significantly different from those of the randomly selected

148 pseudo-absence points (p < 0.001). Forty percent of the nests were located on flat plains, 30%

149 on eastern domains, 15% on southeastern domains and 10% on southern domains. Moreover,

150 the majority of the nests’ entrances was oriented towards the east (50%) and south (25%)

151 directions (Fig. 2).

152 Discussion

153 Desert lark’s nesting locations had low vegetation covers (canopy cover of less than 10%), but

154 larks did not build nests in completely non-vegetated areas. They also tended to select the

155 immediate margin of big stones and sometimes small holes on the lateral side of rocks as the

156 nesting locations. Such a propensity in nest-site selection is to protect the nest and its contents

157 from the direct heat of sunlight in regions where few shadow areas are available. Similar

158 studies have also revealed that the availability of large stones is of high importance for the

159 nest-site selection of desert lark (Vaurie 1951; Orr 1970). Our results showed that desert lark

160 could carry pebbles North-westernwith an average size of Journal 450 grams andof Zoologythickness-edge of less than 3 mm 161 to the nest location. Accepted paper until proofing

162 Before noon, wind speed is often below 0.1 m s-1, and then the wind speed increases with

163 increasing temperature. Nearly 35% of proportionally strong winds moved from the west,

164 south, and southwest. It seems that building nests on the ground in the immediate eastern

165 margin of big stones are strategies to avoid the damaging effect of wind. In addition, the

166 majority of the desert lark’s nests were situated on the east, southeast, and or south-facing

167 slopes, which shelter the nests from south-westerly and southerly warm winds and the hot

168 afternoon sun. In contrast, nests receive and absorb the maximum direct solar radiation at the Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-8

169 beginning of a day when the mean minimum temperature is about 9.8° C. These findings are

170 consistent with the results of previous studies on open-cup ground-nesting species

171 (Mainwaring et al. 2014 and references within it). The internal slope of the rock beside the

172 nest provides shade during the middle part of the day. Moreover, building a rampant by a

173 combination of small pebbles and twigs of plants such as Hammada salicornica creates a

174 porous structure, which reduces thermal fluctuations within the nests. Other studies have also

175 outlined that desert lark alleviates the nest temperature by enclosing the nest through a rim of

176 pebbles (Orr 1970; Afik et al. 1991). It is documented that a high percentage of the nest

177 entrance selected by middle-latitude ground-nesting bird species are easterly-oriented (Burton

178 2007). This strategy is effective in protecting nests from harmful effects of sunlight irradiated

179 from the west (Afik et al. 1991; Lloyd & Martin 2004; Burton 2006; Long et al. 2009),

180 reducing water needs (Tieleman et al. 1999) and absorbing the maximum thermal energy of

181 sunlight in the cool morning time (Richardson 1965; Orr 1970; Burton 2006, Schaaf et al

182 2020b).

183 In conclusion, desert lark selects rocky areas with vegetation cover of less than 10 percent as

184 the nesting area and refuse to build nests on non-vegetated bare soils. The presence of big

185 stones with a weight of less than 6.5 kg, the internal slope of the stone, and appropriate

186 geographical aspectNorth-western are important factors Journal in nest-site of Zoology selection by desert lark. The Accepted paper until proofing 187 geographical aspect of the nesting site and the nest entrance orientation are selected to

188 alleviate the effect of direct solar radiation (Schaaf 2020b) , and winds (Mezquida 2004)

189 moved from west and southwest, and in turn, to stabilise microclimate condition inside the

190 nest.

191 Acknowledgement

192 We are grateful to Zarindasht Department of Environment for assisting us during the

193 fieldwork. The authors declare no competing interests. Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-9

194 References

195 Afik, D., Ward, D., Shkedy, Y. (1991): A test of the self-incubation hypothesis for desert 196 birds that build a rampart of stones in front of their nests. Journal of Thermal Biology 197 16: 255–260.

198 Ar, A., Sidis, Y. (2002): Nest microclimate during incubation. In “Avian Incubation: 199 Behaviour, Environment, and Evolution”.(Ed. DC Deeming.) pp. 143–160.

200 Ardia, D.R. (2005): Cross-fostering reveals an effect of spleen size and nest temperatures on 201 immune responses in nestling European starlings. Oecologia 145: 327–334.

202 Ardia, D. R., Pérez, J. H., Clotfelter, E. D. (2006): Nest box orientation affects internal 203 temperature and nest site selection by Tree Swallows. In Journal of Field Ornithology 204 (Vol. 77, Issue 3, pp. 339–344). Blackwell Publishing Inc.

205 Aznar, F. J., Ibáñez-Agulleiro, M. (2016). The function of stones in nest building: the case of 206 Black Wheatear (Oenanthe leucura) revisited. Avian Biology Research 9: 3-12.

207 BirdLife International 2016. The IUCN Red List of threatened species. BirdLife International 208 Cambridge, UK.

209 Boer, J. (2018): The effect of nest box temperature on the breeding success of Pied 210 Flycatchers (Ficedula hypoleuca) in northern Sweden. M.Sc. dissertation, Swedish 211 University of Agricultural Sciences.

212 Burton, N.H.K. (2006): Nest orientation and hatching success in the tree pipit Anthus trivialis. 213 Journal of Avian Biology 37: 312–317.

214 Burton, N. H. K. (2007): Intraspecific latitudinal variation in nest orientation among ground- 215 nesting : a study using published data. The Condor 109: 441-446.

216 Carroll, R.L., Davis, C.A., Fuhlendorf, S.D., Elmore, R.D., Carroll, J.M. (2020): Orientation 217 affects nest North-westerntemperature of ground-nesting Journal birds. of The Zoology Wilson Journal of Ornithology 218 132:83–90. Accepted paper until proofing 219 Cook, M.I., Beissinger, S.R., Toranzos, G.A., Rodriguez, R.A., Arendt, W.J. (2003): Trans- 220 shell infection by pathogenic micro-organisms reduces the shelf life of non-incubated 221 bird’s eggs: A constraint on the onset of incubation? Proceedings of the Royal Society B: 222 Biological Sciences 270: 2233–2240.

223 Etchécopar, R. D., Hüe, F. (1967): The birds of North Africa, from the Canary Islands to the 224 Red Sea. Oliver & Boyd.

225 Ferguson-Lees, I. J. (1960): Studies of less familiar birds: black wheatear. British Birds 53: 226 553–558.

227 Hansell, M. (2000): Bird nests and construction behaviour. Cambridge University Press. Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-10

228 Harris, J., Smith, L., & McMurry, S. (2020): A multiscale analysis of Gray Vireo (Vireo 229 vicinior) nest-site selection in central New Mexico. Avian Conservation and Ecology 230 15(1):12. 231 Hartman, C.A., Oring, L. W. (2003): Orientation and microclimate of Horned Lark nests: the 232 importance of shade. The Condor 105: 158-163. 233 234 Heenan, C. B. (2013): An overview of the factors influencing the morphology and thermal 235 properties of avian nests. Avian Biology Research 6: 104-118.

236 Hoekman, S.T., Ball, I.J., Fondell, T.E. (2002): Grassland birds orient nests relative to nearby 237 vegetation. Wilson Bulletin 114: 450–456.

238 Holway, D.A. (1999): Nest-site selection and the importance of nest concealment in the 239 black-throated blue warbler. NCASI Technical Bulletin 93: 240–241.

240 Hosseini-Moosavi, S.M., Barati, A., Hemami, M.R., Karimpour, R. (2017): Nest-site 241 selection and breeding success of the semi-desert bird, Grey Hypocolius Hypocolius 242 ampelinus, in relation to plant structure of Ziziphus nummularia. Avian Biology 243 Research, 10: 181-189. 244 Jones, J. (2001): Habitat selection studies in avian ecology: a critical review. The auk, 118: 245 557-562.

246 Knopf, F.L., Sedgwick, J.A. (1992): An Experimental Study of Nest-Site Selection by Yellow 247 Warblers. The Condor 94: 734–742.

248 Kotliar, N.B., Wiens, J.A. (1990): Multiple scales of patchiness and patch structure: a 249 hierarchical framework for the study of heterogeneity. Oikos, 59: 253-260.

250 Lloyd, J. D., Martin, T. E. (2004): Nest-site preference and maternal effects on offspring 251 growth. Behavioral Ecology, 15(5): 816–823.

252 Long, A. M., Jensen, W. E., With, K. A. (2009): Orientation of Grasshopper Sparrow and 253 Eastern Meadowlark Nests in Relation to Wind Direction. The Condor, 111(2): 395–399. North-western Journal of Zoology 254 Mainwaring, M. C., Hartley, I. R., Lambrechts, M. M., Deeming, D. C. (2014): The design 255 and function of birds’Accepted nests. Ecology paper and Evolution, until 4(20):proofing 3909–3928.

256 Martin, T.E., Roper, J.J. (1988): Nest Predation and Nest-Site Selection of a Western 257 Population of the Hermit Thrush. The Condor 90: 51–57.

258 Mezquida, E.T. (2004): Nest site selection and nesting success of five species of passerines in 259 a South American open Prosopis woodland. Journal of Ornithology 145: 16-22. 260 261 Mezquida, E.T. (2004): Patrones de orientación de los nidos de Passeriformes en una zona 262 árida del centro-oeste de Argentina. Ornitología Neotropical 15: 145-153.

263 Orr, Y. (1970): Temperature Measurements at the Nest of the Desert Lark (Ammomanes 264 deserti deserti): The Condor 72: 476–478. Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-11

265 Petersen, K.L., Best, L.B. (1985): Nest-Site Selection by Sage Sparrows. The Condor 87: 266 217-221.

267 Petit, K.E., Petit, D.R., Petit, L.J. (1988): On measuring vegetation characteristics in bird 268 territories: nest sites vs. perch sites and the effect of plot size. American Midland 269 Naturalist, 119: 209-215.

270 Porneluzi, P., Van Horn, M.A., & Donovan, T.M. (2011): Ovenbird (Seiurus aurocapilla), 271 version 2.0. The birds of North America. Edited by A.F. Poole. Cornell Lab of 272 Ornithology, Ithaca, NY, 10.

273 Rauter, C.M., Reyer, H.U., Bollmann, K. (2002): Selection through predation, snowfall and 274 microclimate on nest-site preferences in the Water Pipit Anthus spinoletta. Ibis 144: 275 433–444.

276 Richardson, F. (1965): Breeding and Feeding Habits of the Black Wheatear Oenanthe 277 Leucura in Southern Spain. Ibis 107: 1–16.

278 Schaaf, A.A. (2020a). Orientación de nidos de hornero (Furnarius rufus): Efectos de la 279 vegetación, el viento y la radiación solar en el noroeste de la Argentina. Ecología 280 Austral 30: 146-150. 281 282 Schaaf, A.A. (2020b). The effect of climate on nest orientation in the Rufous Hornero 283 Furnarius rufus. Bird Study 67: 168-172. 284 Schaaf, A. A., García, C. G., Puechagut, P. B., Silvetti, L. E., Tallei, E., Ortis, F., Quaglia, A. 285 I. (2018a). Effect of geographical latitude and sun exposure on Rufous Hornero 286 (Furnarius rufus) nest orientation. Journal of Ornithology 159: 967-974. 287 288 Schaaf, A.A., Rojas, T.N., Diaz, A.E., Peluc, S.I. (2018b). Patterns of nest orientation in the 289 golden billed saltator (Saltator aurantiirostris) in central Argentina.

290 Shkedy, Y., Safriel, U.N. (1992): Niche breadth of two lark species in the desert and the size 291 of their geographical ranges. Ornis Scandinavica 23: 89–95. North-western Journal of Zoology 292 Smith, P.A. (2003). Factors affecting nest site selection and reproductive success of tundra 293 nesting shorebirds (PhDAccepted dissertation, paper University until of British proofing Columbia).

294 Starck, J.M., Ricklefs, R.E. (1998): Avian growth and development: evolution within the 295 altricial-precocial spectrum (Issue 8): Oxford University Press, New York.

296 Tieleman, B.I., Williams, J.B., Michaeli, G., Pinshow, B. (1999): The role of the nasal 297 passages in the water economy of crested larks and desert larks. Physiological and 298 Biochemical Zoology 72: 219–226.

299 Vaurie, C. (1951): A study of Asiatic larks. Bulletin of the American Museum of Natural 300 History 97: 431–526.

301 Vleck, C.M. (1981): Energetic cost of incubation in the zebra finch. The Condor 83: 229-237. Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-12

302 Webb, D.R., King, J.R. (1983): An analysis of the heat budgets of the eggs and nest of the 303 white-crowned sparrow, Zonotrichia leucophrys, in relation to parental attentiveness. 304 Physiological Zoology 56: 493–505.

305 With, K.A. (1994): The hazards of nesting near shrubs for a grassland bird, the McCown’s 306 longspur. The Condor 96: 1009–1019.

307 With, K.A., Webb, D.R. (1993): Microclimate of ground nests: the relative importance of 308 radiative cover and wind breaks for three grassland species. The Condor 95: 401-413.

309

310 Table captions

311 Table 1. Range and mean ± SD values of measurements of gravels used to fence the entrance

312 of the desert lark’s nest

313 Table 2. Range and mean ± SD values of variables measured in 1 m- and 4 m-radius occupied and

314 unoccupied plots and the results of the Mann-Whitney test.

315

316 Figure captions

317 Figure 1. Nests of desert lark built by small pebbles and twigs. The nests were shadowed at

318 12:00 AM (A) and 16:00 PM (B) on 15th October 2014

319 Figure 2. The geographical aspect of the nesting area in the nest presence (A) and absence

320 locations (B), the nest entrance orientation built by desert lark (C) and the wind

321 rose of the study areaNorth-western (D). P denotes the percentage Journal of domains of Zoology with no geographical aspect Accepted paper until proofing 322 (slope of less than 10%).

323

324

325

326

327

328 Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-13

329

330 Tables:

331 Table 1. Variable

Length Width Overall Weight Maximum Minimum (mm) (mm) thickness (gr) thickness of thickness of (mm) edge (mm) edge (mm)

Range 11-25 4-15 2-7 0.4-1.5 2-6 0.5-4

Mean±SD 16.3±3.7 9.8±2.7 4.7±1.2 0.8±0.3 4.5±1.3 1.5±0.9

Variable The heaviest pebble used to fence the nest entrance

Length Width Overall Weight Maximum Minimum (mm) (mm) thickness (gram) thickness of thickness of (mm) edge (mm) edge (mm)

Range 20-45 12-35 5-15 3.7-12.1 4-15 0.5-3

Mean±SD 35.1±6.5 23.4±5.7 8.5±2.8 6.7±2.2 8.2±3.1 1.8±0. 7

332

333

334 Table 2.

Variable Plot Mean ± SD Man-Whitney size Presence Absence Z p North-western(m2) Journal of Zoology Gravel (≤50 mm) Accepted3.14 %43.63 paper±25.74 until%14.68 proofing±18 -4.071 0.001 50.27 %34.73±17.27 %18.79±16.61 -2.922 0.003 Gravel (<50 mm) 3.14 %26.36±16.65 %25.84±31.15 -1.305 0.192 50.27 %29.21±16.18 %19.47±18.66 -2.088 0.037 Bare ground 3.14 %25.05±15.69 %56.21±33.81 -2.703 0.007 50.27 %29.21±17.90 %54.21±25.22 -3.083 0.002 Vegetation 3.14 %4.95±4.74 %3.26±4.21 -1.318 0.187 50.27 %6.84±4.89 %7.53±7.26 -0.133 0.894 Slope %29.52±34.08 %11.89±14.62 -0.742 0.458 335

336 Use the following type of citation: North-western Journal of Zoology 2021: e212601

nwjz-14

337 338

339 Figure 1. 340

North-western Journal of Zoology Accepted paper until proofing

341 342 Figure 2. 343