Population Ecology

, Volume 59, Issue 2, pp 131–137 | Cite as

Influence of weather and phenotypic characteristics on pregnancy rates of female roe deer in central Italy

Original article


In many ungulates, female fecundity is influenced by weather, population density and body condition. Based on a six-year survey of the reproductive tracts of adult female roe deer (Capreolus capreolus) harvested in the province of Pisa (Tuscany, central Italy), we evaluated the influence of weather, population density and individual characteristics on pregnancy rates. Eighty-five percent of females were pregnant, with a median litter size of two (range one—three). We found that pregnancy rate was positively correlated with summer rainfall and body mass of females, whereas early-winter conditions, spring rainfall, the age of females and density-dependent factors did not appear to influence pregnancy rate. These results reflect the particular seasonal variation in the abundance and quality of resources in Mediterranean habitats and show that heavier females (high-quality mothers) are more productive than lighter females.


Body mass Capreolus capreolus Hunting bags Mediterranean environment Summer rainfall 


  1. Aanes R, Andersen R (1996) The effects of sex, time of birth, and habitat on the vulnerability of roe deer fawns to red fox predation. Can J Zool 74:1857–1865CrossRefGoogle Scholar
  2. Aiello V, Lovari S, Bocci A (2013) Ranging behaviour and reproductive rate in the threatened population of roe deer in Gargano, South Italy. Ital J Zool 80:614–619CrossRefGoogle Scholar
  3. Akaike H (1973) Information theory as an extension of the maximum likelihood principle. In: Petrov BN, Csaki F (eds) Second international symposium on information theory. Akademiai Kiado, Budapest, pp 267–281Google Scholar
  4. Andersen R, Linnell JDC (2000) Irruptive potential in roe deer: density-dependent effects on body mass and fertility. J Wildl Manage 64:698–706CrossRefGoogle Scholar
  5. Andersen R, Gaillard J-M, Linnell JDC, Duncan P (2000) Factors affecting maternal care in an income breeder, the European roe deer. J Anim Ecol 69:672–682CrossRefGoogle Scholar
  6. Anderson DR, Burnham KP, Thompson WL (2000) Null hypothesis testing: problems, prevalence and an alternative. J Wildl Manage 64:912–923CrossRefGoogle Scholar
  7. Barbieri MM, Berger JO (2004) Optimal predictive model selection. Ann Stat 32:870–897CrossRefGoogle Scholar
  8. Bartoń K (2013) MuMIn: multi-model inference. Model selection and model averaging based on information criteria (AICc and alike). R package version 1.9.13Google Scholar
  9. Bates D, Maechler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1–48CrossRefGoogle Scholar
  10. Bekenov AB, Grachev IuA, Milner-Gulland EJ (1998) The ecology and management of the Saiga antelope in Kazakhstan. Mammal Rev 28:1–52CrossRefGoogle Scholar
  11. Bender LC, Weisenberger ME (2005) Precipitation, density, and population dynamics of desert bighorn sheep on San Andres National wildlife refuge, New Mexico. Wildl Soc Bull 33:956–964CrossRefGoogle Scholar
  12. Bonenfant C, Gaillard J-M, Coulson T, Festa-Bianchet M, Loison A, Garel M, Loe LE, Blanchard P, Pettorelli N, Owen-Smith N, Du Toit J, Duncan P (2009) Empirical evidence of density-dependence in populations of large herbivores. Adv Ecol Res 41:314–357Google Scholar
  13. Bongi P, Ciuti S, Grignolio S, Del Frate M, Simi S, Gandelli D, Apollonio M (2008) Anti-predator behaviour, space use and habitat selection in female roe deer during the fawning season in a wolf area. J Zool 276:242–251CrossRefGoogle Scholar
  14. Borkowski J, Palmer SCF, Borowski Z (2011) Drive counts as a method of estimating ungulate density in forests: mission impossible? Acta Theriol 56:239–253CrossRefPubMedPubMedCentralGoogle Scholar
  15. Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information–theoretic approach, 2nd edn. Springer, New YorkGoogle Scholar
  16. Camuffo D, Bertolin C, Diodato N, Barriendos M, Dominguez-Castro F, Cocheo C, della Valle A, Garnier E, Alcoforado MJ (2010) The western Mediterranean climate: how will it respond to global warming? Clim Change 100:137–142CrossRefGoogle Scholar
  17. Carvalho P, Nogueira AJA, Soares AMVM, Fonseca C (2008) Ranging behaviour of translocated roe deer in a Mediterranean habitat: seasonal and altitudinal influences on home range size and patterns of range use. Mammalia 72:89–94CrossRefGoogle Scholar
  18. Cotrufo MF, Alberti G, Inglima I, Marjanović H, LeCain D, Zaldei A, Peressotti A, Miglietta F (2011) Decreased summer drought affects plant productivity and soil carbon dynamics in a Mediterranean woodland. Biogeosciences 8:2729–2739CrossRefGoogle Scholar
  19. Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré J, García Marquéz JR, Gruber B, Lafourcade B, Leitão PJ, Münkemüller T, McClean C, Osborne PE, Reineking B, Schröder B, Skidmore AK, Zurell D, Lautenbach S (2013) Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography 36:27–46CrossRefGoogle Scholar
  20. Duncan C, Chauvenet ALM, McRae LM, Pettorelli N (2012) Predicting the future impact of droughts on ungulate populations in arid and semi-arid environments. PLoS One 7(12):e51490CrossRefPubMedPubMedCentralGoogle Scholar
  21. Epps CW, McCullough DR, Wehausen JD, Bleich VC, Rechel JL (2004) Effects of climate change on population persistence of desert-dwelling mountain sheep in California. Conserv Biol 18:102–113CrossRefGoogle Scholar
  22. Evans JP (2009) 21st century climate change in the Middle East. Clim Change 92:417–432CrossRefGoogle Scholar
  23. Fernández-Llario P, Mateos-Quesada P (2005) Influence of rainfall on the breeding biology of wild boar (Sus scrofa) in a Mediterranean ecosystem. Folia Zool 54:240–248Google Scholar
  24. Ferretti F, Bertoldi G, Sforzi A, Fattorini L (2011) Roe and fallow deer: are they compatible neighbours? Eur J Wildlife Res 57:775–783CrossRefGoogle Scholar
  25. Flajšman K, Jelenko I, Poličnik H, Pokorny B (2013) Reproductive potential of roe deer (Capreolus capreolus L.): review of the most important influential factors. Acta Silvae et Ligni 102:1–20CrossRefGoogle Scholar
  26. Focardi S, Raganella Pelliccioni E, Petrucco R, Toso S (2002) Spatial patterns and density dependence in the dynamics of a roe deer (Capreolus capreolus) population in central Italy. Oecologia 130:411–419CrossRefPubMedGoogle Scholar
  27. Focardi S, Aragno P, Montanaro P, Riga F (2006) Inter-specific competition from fallow deer Dama dama reduces habitat quality for the Italian roe deer Capreolus capreolus italicus. Ecography 29:407–417CrossRefGoogle Scholar
  28. Freckleton RP, Watkinson AR, Green RE, Sutherland WJ (2006) Census error and the detection of density dependence. J Anim Ecol 75:837–851CrossRefPubMedGoogle Scholar
  29. Gaillard J-M, Festa-Bianchet M, Yoccoz NG, Loison A, Toïgo C (2000a) Temporal variation in fitness components and population dynamics of large herbivores. Annu Rev Ecol Syst 31:367–393CrossRefGoogle Scholar
  30. Gaillard J-M, Festa-Bianchet M, Delorme D, Jorgenson J (2000b) Body mass and individual fitness in female ungulates: bigger is not always better. Proc R Soc Lond B 267:471–477CrossRefGoogle Scholar
  31. Geographica Srl (2012) Ambito Territoriale di Caccia Pisa occidentale 15. Distretti di caccia di selezione: consuntivo stagione venatoria 2011/2012 (in Italian)Google Scholar
  32. Golodets C, Sternberg M, Kigel J, Boeken B, Henkin Z, Seligman NG, Ungar ED (2013) From desert to Mediterranean rangelands: will increasing drought and inter-annual rainfall variability affect herbaceous annual primary productivity? Clim Change 119:785–798CrossRefGoogle Scholar
  33. Hamel S, Côté SD, Gaillard J-M, Festa-Bianchet M (2009) Individual variation in reproductive costs of reproduction: high-quality females always do better. J Anim Ecol 78:143–151CrossRefPubMedGoogle Scholar
  34. Häsler H, Senn J (2012) Ungulate browsing on European silver fir Abies alba: the role of occasions, food shortage and diet preferences. Wildlife Biol 18:67–74CrossRefGoogle Scholar
  35. Hewison AJM (1996) Variation in the fecundity of roe deer in Britain: effects of age and body weight. Acta Theriol 41:187–198CrossRefGoogle Scholar
  36. Hewison AJM, Gaillard J-M (2001) Phenotypic quality and senescence affect different components of reproductive output in roe deer. J Anim Ecol 70:600–608CrossRefGoogle Scholar
  37. Hewison AJM, Angibault JM, Boutin J, Bideau E, Vincent JP, Sempéré A (1996) Annual variation in body composition of roe deer (Capreolus capreolus) in moderate environmental conditions. Can J Zool 74:245–253CrossRefGoogle Scholar
  38. Hewison AJM, Gaillard J-M, Kjellander P, Toïgo C, Liberg O, Delorme D (2005) Big mothers invest more in daughters—reversed sex allocation in a weakly polygynous mammal. Ecol Lett 8:430–437CrossRefGoogle Scholar
  39. Hosmer DW, Lemeshow S (2000) Applied logistic regression. John Wiley and Sons, New YorkCrossRefGoogle Scholar
  40. Imperio S, Focardi S, Santini G, Provenzale A (2012) Population dynamics in a guild of four Mediterranean ungulates: density-dependence, environmental effects and inter-specific interactions. Oikos 121:1613–1626CrossRefGoogle Scholar
  41. Joly DO, Messier F (2004) Testing hypotheses of bison population decline (1970–1999) in Wood Buffalo National Park: synergism between exotic disease and predation. Can J Zool 82:1165–1176CrossRefGoogle Scholar
  42. Jönsson KI (1997) Capital and income breeding as alternative tactics of resources use in reproduction. Oikos 78:57–66CrossRefGoogle Scholar
  43. Koons DN, Terletzky P, Adler PB, Wolfe ML, Ranglack D, Howe FP, Hersey K, Paskett W, Du Toit JT (2012) Climate and density-dependent drivers of recruitment in plains bison. J Mammal 93:475–481CrossRefGoogle Scholar
  44. Lawrence RK, Demarais S, Relyea RA, Haskell SP, Ballard WB, Clark TL (2004) Desert mule deer survival in southwest Texas. J Wildl Manage 68:561–569CrossRefGoogle Scholar
  45. Lele SR, Keim JL, Solymos P (2016) ResourceSelection: resource selection (probability) functions for use-availability data. R package version 0.2–6Google Scholar
  46. Linnell JDC, Nijhuis P, Teurlings I, Andersen R (1999) Selection of bed-sites by roe deer Capreolus capreolus fawns in a boreal forest landscape. Wildlife Biol 5:225–231Google Scholar
  47. Loveridge AJ, Hunt JE, Murindagomo F, Macdonald DW (2006) Influence of drought on predation of elephant (Loxodonta africana) calves by lions (Panthera leo) in an African wooded savannah. J Zool 270:523–530CrossRefGoogle Scholar
  48. Lowe VPW, Thompson-Schwab JDD (2003) Using cohort analysis to reconstruct the size and structure of deer populations in forestry with special reference to roe deer (Capreolus capreolus L.). Forestry 76:437–447CrossRefGoogle Scholar
  49. Mateos-Quesada P, Carranza J (2000) Reproductive patterns of roe deer in Central Spain. Etología 8:17–20Google Scholar
  50. Mazzarone V, Dall’Antonia P, Scarselli D, Varuzza P, Viviani V, Genghi MA, Menconi R, Vecchio G, Giuttari G, Fichera F, Bazzichi L, Luccarini S, Capaccioli A, Mauri L (2012) Proposte per la gestione delle specie faunistiche. In: Mazzarone V, Poli A (eds) Piano Faunistico Venatorio Pisa 2012–15. Pisa, pp 57–144 (in Italian)Google Scholar
  51. Meriggi A, Sotti F, Lamberti P, Gilio N (2008) A review of the methods for monitoring roe deer European populations with particular reference to Italy. Hystrix Ital J Mammal 19:103–120Google Scholar
  52. Minder I (2012) Local and seasonal variations of roe deer diet in relation to food resource availability in a Mediterranean environment. Eur J Wildlife Res 58:215–225CrossRefGoogle Scholar
  53. Miranda JD, Armas C, Padilla FM, Pugnaire FI (2011) Climatic change and rainfall patterns: effects on semi-arid plant communities of the Iberian Southeast. J Arid Environ 75:1302–1309CrossRefGoogle Scholar
  54. Miyashita T, Suzuki M, Ando D, Fujita G, Ochiai K, Asada M (2008) Forest edge creates small-scale variation in reproductive rate of sika deer. Popul Ecol 50:111–120CrossRefGoogle Scholar
  55. Mucci N, Mattucci F, Randi E (2012) Conservation of threatened local gene pools: landscape genetics of the Italian roe deer (Capreolus c. italicus) populations. Evol Ecol Res 14:897–920Google Scholar
  56. Mysterud A, Østbye E (1999) Cover as a habitat element for temperate ungulates: effects on habitat selection and demography. Wildl Soc Bull 27:385–394Google Scholar
  57. Mysterud A, Bjørnsen BH, Østbye E (1997) Effects of snow depth on food and habitat selection by roe deer Capreolus capreolus in an altitudinal gradient in south-central Norway. Wildl Biol 3:27–33Google Scholar
  58. Nilsen EB, Linnell JDC, Andersen R (2004) Individual access to preferred habitat affects fitness components in female roe deer Capreolus capreolus. J Anim Ecol 73:44–50CrossRefGoogle Scholar
  59. Noss AJ, Cuéllar Salidas RL, Ayala Crespo J (2006) Drive counts for grey brocket deer Mazama gouazoupira in the Bolivian Chaco. Mammalia 70:64–69CrossRefGoogle Scholar
  60. Post E (2005) Large-scale spatial gradients in herbivore population dynamics. Ecology 86:2320–2328CrossRefGoogle Scholar
  61. R Development Core Team (2015) R: A language and environment for statistical computing. R foundation for statistical computing, Vienna, Austria. http://www.R-project.org
  62. Rodriguez-Hidalgo P, Gortázar C, Tortosa FS, Rodriguez-Vigal C, Fierro Y, Vicente J (2010) Effects of density, climate, and supplementary forage on body mass and pregnancy rates of female red deer in Spain. Oecologia 164:389–398CrossRefPubMedGoogle Scholar
  63. Sæther B-E (1997) Environmental stochasticity and population dynamics of large herbivores: a search for mechanisms. Trends Ecol Evol 12:143–149CrossRefPubMedGoogle Scholar
  64. Sempéré AJ, Sokolov VE, Danilkin AA (1996) Capreolus capreolus. Mammal Species 538:1–9CrossRefGoogle Scholar
  65. Tellería JL, Virgós E (1997) Distribution of an increasing roe deer population in a fragmented Mediterranean landscape. Ecography 20:247–252CrossRefGoogle Scholar
  66. Toïgo C, Gaillard J-M, Van Laere G, Hewison M, Morellet N (2006) How does environmental variation influence body mass, body size, and body condition? Roe deer as a case study. Ecography 29:301–308CrossRefGoogle Scholar
  67. Torres RT, Virgós E, Santos J, Linnell JDC, Fonseca C (2012) Habitat use by sympatric red and roe deer in a Mediterranean ecosystem. Anim Biol 62:351–366CrossRefGoogle Scholar
  68. Vanpé C, Gaillard J-M, Morellet N, Kjellander P, Liberg O, Delorme D, Hewison AJM (2009) Age-specific variation in male breeding success of a territorial ungulate species, the European roe deer. J Mammal 90:661–665CrossRefGoogle Scholar
  69. Wallach AD, Inbar M, Scantlebury M, Speakman JR, Shanas U (2007) Water requirements as a bottleneck in the reintroduction of European roe deer to the southern edge of its range. Can J Zool 85:1182–1192CrossRefGoogle Scholar
  70. Zuur AF, Ieno EN, Walker NJ, Saveliev AA, Smith GM (2009) Mixed effects models and extensions in ecology with R. Springer, New YorkCrossRefGoogle Scholar

Copyright information

© The Society of Population Ecology and Springer Japan 2017

Authors and Affiliations

  1. 1.Department of Earth and Environmental SciencesUniversity of PaviaPaviaItaly
  2. 2.Geographica srlPisaItaly

Personalised recommendations