Local variation in sex ratios in three species of wintering ducks: the need for large-scale sampling
DOI:
https://doi.org/10.51812/of.160744Keywords:
wildfowl, sex ratio, wintering, habitat selection, Gulf of GdańskAbstract
This study investigates the hypothesis that site characteristics and flock size influence the sex ratios of wintering ducks by examining three species: the Mallard (Anas platyrhynchos), the Goldeneye (Bucephala clangula), and the Long-tailed Duck (Clangula hyemalis) in the Gulf of Gdańsk, southern Baltic. Birds were counted in mid-January along approximately 160 km of shoreline in the western part of the Gulf of Gdańsk and the estuarine sections of the Vistula River on the southern Baltic coast. Only flocks in which the sex of all individuals was identified were included in the analysis. The sex ratios of the three study species varied across the five habitat types within the study area but did not vary with species-specific flock size. In Mallards the lowest proportion of males was recorded along the open seacoast and municipal beach, suggesting that these areas are primarily occupied by paired birds. The higher proportion of males at other sites is likely due to the shift of unpaired individuals to these locations, where the overall number of this species is the highest. In the Goldeneye the highest percentage of females was observed in the most sheltered areas, while the lowest percentage, alongside low individual numbers, occurred along the open seacoast. These patterns suggest that Goldeneye density and sex ratio are influenced by female tolerance to adverse weather conditions, as this species shows the greatest size difference between males and females among the species studied. Long-tailed Ducks exhibited a more uniform sex ratio across the study area, likely due to their frequent movements to access areas with abundant food resources. This study highlights spatial variations in duck sex ratios within a single water body and underscores the need for large scale sampling to obtain representative sex ratio estimates.
References
Alexander, W.C. 1987: Aggressive behavior of wintering diving ducks (Aythyini). — Wilson Bulletin 99: 38–49
Andrulewicz, E., Kruk-Dowgiałło, L. & Osowiecki, A. 2004: An expert judgement approach to designating ecosystem typology and assessing the health of Gulf of Gdansk. — In Managing the Baltic Sea. Coastline Reports 2: 53–61 (ed. Schernewski, G. & Löser, N.). EUCC, Rostock.
Ashcroft, R.E. 1976: A function of the pairbond in the Common Eider. — Wildfowl 27: 101–105.
Avilova, K.V., Eremkin, G.S. 2001: Waterfowl wintering in Moscow (1985-1999): dependence on air temperatures and the prosperity of the human population. — Acta Ornithologica 36: 65–71.
Badur, J. & Cieślikiewicz, W. 2018: Spatial variability of long-term trends in significant wave height over the Gulf of Gdansk using System Identification techniques. — Oceanological and Hydrobiological Studies 47: 190 –201. https://doi.org/10.1515/ohs-2018-0018.
Bellebaum, J., & Mädlow, W. 2015: Survival explains sex ratio in an introduced Mandarin Duck Aix galericulata population. — Ardea 103: 183–187. https://doi.org/10.5253/arde.v103i2.a7.
Bennett, J.W. & Bolen, E.G. 1978: Stress response in wintering Green-winged Teal. — Journal of Wildlife Management 42: 87–86. https://doi.org/10.2307/3800692
Blums, P. & Mednis, A. 1996: Secondary sex ratio in Anatinae. — Auk 113, 505–511. https://doi.org/10.2307/4088920
Błeńska, M. & Osowiecki, A. 2015: Biotic typology of Polish marine areas based on bottom macrofauna communities. — Bulletin of the Maritime Institute in Gdańsk 30: 167–173. https://doi.org/10.5604/12307424.1186449
Bossema, I. & Roemers, E. 1985: Mating strategy, including mate choice in Mallards. — Ardea 73: 147–157.
Bourget, D., Savard, J.-P.L. & Guillemette, M. 2007: Distribution, diet and dive behavior of Barrow’s and Common Goldeneyes during spring and autumn in the St. Lawrence Estuary. — Waterbirds 30: 230–240. https://doi.org/10.1675/1524-4695(2007)30[230:ddadbo]2.0.co;2
Brides, K., Wood, K.A., Hearn, R.D. & Fijen, T.P.M. 2017: Changes in the sex ratio of the common pochard Aythya ferina in Europe and North Africa. — Wildfowl 67: 100–112.
Campbell, L.H. 1977: Local variations in the proportion of adult males in flocks of Goldeneye wintering in the Firth of Forth. — Wildfowl 28: 77–80.
Carbone, C., Owen, M. 1995: Differential migration of the sexes of Pochard Aythya ferina: results from a European survey. — Wildfowl 46: 99–108.
Cervencl, A., Troost, K., Dijkman, E., de Jong, M., Smit, C.J., Leopold, M.F. & Ens, B.J. 2015: Distribution of wintering Common Eider Somateria mollissima in the Dutch Wadden Sea in relation to available food stocks. — Marine Biology 162: 153–168. https://doi.org/10.1007/s00227-014-2594-4
Choudhury, S., Black, J.M. 1991: Testing the behavioural dominance and dispersal hypothesis in Pochard. — Ornis Scandinavica 22: 155–159. https://doi.org/10.2307/3676546
Cieślikiewicz, W. & Cupiał, A. 2024: Long-term statistics of atmospheric conditions over the Baltic Sea and meteorological features related to wind wave extremes in the Gulf of Gdańsk. — Oceanologia 66: 180–195. https://doi.org/10.1016/j.oceano.2023.10.002
Cieślikiewicz, W., Dudkowska, A., Gic-Grusza, G. & Jędrasik, J. 2017: Extreme bottom velocities induced by wind wave and currents in the Gulf of Gdańsk. — Ocean Dynamics 67: 1461–1480. https://doi.org/10.1016/10.1007/s10236-017-1098-4
Combs, D.L. & Fredrickson, L.H. 1990: Foods used by male mallards wintering in southeastern Missouri. — Journal of Wildlife Management 60: 603–610. https://doi.org/10.2307/3802078
Cramp, S., Simmons, K.E.L. (ed.) 1977: Handbook of the birds of Europe the Middle East and North Africa. Vol. 1. — Oxford University Press, Oxford.
Custer, C.M. & Custer, T.W. 1996: Food habits of diving ducks in the Great Lakes after the Zebra Mussel invasions. — Journal of Field Ornithology 67: 86–99. https://www.jstor.org/stable/4514086
Dabbert, C.B. & Martin, T.E. 2000: Diet of mallards wintering in Greentree Reservoirs in southeastern Arkansas. — Journal of Field Ornithology 71: 423–428. https://doi.org/10.1648/0273-8570-71.3.423
Delnicki, D. & Reinecke, K.J. 1986: Mid-winter food use and body weights of Mallards and Wood Ducks in Mississippi. — Journal of Wildlife Management 50: 43–51. https://doi.org/10.2307/3801486
Donald, P.F. 2007: Adult sex ratios in wild bird populations. — Ibis 149: 671–692. https://doi.org/10.1111/j.1474-919x.2007.00724.x
Dormann, C. F., Elith, J., Bacher, S., Buchmann, C., Carl, G., Carré, G., García Marquéz, J. R., Gruber, B., Lafourcade, B., Leitão, P. J., Münkemüller, T., Mcclean, C., Osborne, P. E., Reineking, B., Schröder, B., Skidmore, A. K., Zurell, D., & Lautenbach, S. 2013: Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. — Ecography 36: 27–46. https://doi.org/10.1111/j.1600-0587.2012.07348.x
Duncan, K. & Marquiss, M. 1993: The sex/age ratio, diving behaviour and habitat use of Goldeneye Bucephala clangula wintering in northeast Scotland. — Wildfowl 44: 111–120.
Dunn, P.K. & Smyth, G.K. 2018: Generalized Linear Models with examples in R. — Springer New York. https://doi.org/10.1007/978-1-4419-0118-7
Dyrcz, C. 2017: Analysis of ice conditions in the Baltic Sea and in the Puck Bay. — Scientific Journal of Polish Naval Academy 58: 13–41. https://doi.org/10.5604/01.3001.0010.6581
Gic-Grusza, G., Kryla-Staszewska, L., Urbański, J., Warzocha, J. & Węsławski, J.M. (ed.) 2009: Atlas of Polish marine area bottom habitats: Environmental valorization of marine habitats. — Broker-Innowacji, Gdynia.
Grieves, L.A., Hing, S., Tabh, J. & Quinn, J.S. 2024: Offspring sex ratio in a communal breeding bird is male-biased when pre-breeding rainfall is low. — Journal of Avian Biology e03262. https://doi.org/10.1111/jav.03262
Evans, D.M. & Day, K.R. 2001: Migration patterns and sex ratios of diving ducks wintering in Northern Ireland with specific reference to Lough Neagh. — Ringing & Migration 20: 358–363. https://doi.org/10.1080/03078698.2001.9674263
Even, JG., Clarke, R.H., Moysey, E., Boulton, R.L., Crozier, R.H. & Clarke, M.F. 2001: Primary sex ratio bias in an endangered cooperatively breeding bird, the black-eared miner, and its implications for conservation. — Biological Conservation 101: 137–145. https://doi.org/10.1016/S0006-3207(01)00022-2
Figley, W. K. & VanDruff, L.W. 1982: The ecology of urban Mallards. — Wildlife Monographs 81: 3–39.
Forni, P., Morkūnas, J. & Daunys, D. 2022: Response of Long-tailed Duck (Clangula hyemalis) to the change in the main prey availability in its Baltic wintering ground. — Animals 12: 355. https://doi.org/10.3390/ani12030355
Fox, A.D. & Cristensen, T. K. 2018: Could falling female sex ratios among first-winter northwest European duck populations contribute to skewed adult sex ratios and overall population declines? — Ibis 160: 929–935. https://doi.org/10.1111/ibi.12649
Frew, R.T., Brides, K., Clare, T., MacLean, L., Rigby, D., Tomlinson, C.G. & Wood, K.A. 2018: Temporal changes in the sex ratio of the Common Pochard Aythya ferina compared to four other duck species at Martin Mere, Lancashire, UK. — Wildfowl 68: 140–154.
Furness, L.J. & Speakman, J.R. 2008: Energetics and longevity in birds. Age 30: 75–87. https://doi.org/10.1007/s11357-008-9054-3
Goodburn, S.F. 1984: Mate guarding in the Mallard Anas platyrhynchos. — Ornis Scandinavica 15: 261–265. https://doi.org/10.2307/3675935
Hair, J.F., Anderson, R.E., Tatham, R.L. & Black, W.C. 1995: Multivariate data analysis. — Prentice-Hall, Englewood Cliffs, NJ.
Hepp, G.R. & Hair, J.D. 1984: Dominance in wintering waterfowl (Anatini): effects on distribution of sexes. — Condor 86: 251–257. https://doi.org/10.2307/1366992
Herra, T. & Wiktor, K. 1985: Composition and distribution of bottom fauna in coastal zone of the Gulf of Gdańsk proper. — Studia i Materialy Oceanologiczne 46: 114–142.
Homolková, M., Musil, P., Pavón-Jordán, D., Gajdošová, D., Musilová, Z., Neužilová, Š., Zouhar, J. 2024. Changes in the adult sex ratio of six duck species breeding populations over two decades. — Avian Research 15: 100187. https://doi.org/10.1016/j.avrs.2024.100187
Jaatinen, K., Lehikoinen, A. & Lank, DB. 2010: Female-biased sex ratios and the proportion of cryptic male morphs of migrant juvenile Ruffs (Philomachus pugnax) in Finland. — Ornis Fennica 87: 125–134. https://doi.org/10.51812/of.133751
Jamieson, S.E., Robertson, G.J. & Grant Gilchrist, H. 2001: Autumn and winter diet of Long-tailed Duck in the Belcher Islands, Nunavut, Canada. — Waterbirds 24: 129–132. https://doi.org/10.2307/1522253
Janas, U., Burska, D., Kendzierska, H., Pryputniewicz-Flis, D. & Łukawska-Matuszewska, K. 2019: Importance of benthic macrofauna and coastal biotopes for ecosystem functioning – Oxygen and nutrient fluxes in the coastal zone. — Estuarine, Coastal and Shelf Science 225: 106238. https://doi.org/10.1016/j.ecss.2019.05.020
Johnson, W.P. & Rohwer, F.C. 1998: Pairing chronology and agonistic behaviors of wintering Green-winged Teal and Mallards. — Wilson Bulletin 110: 311–315. http://www.jstor.org/stable/4163954
Jonsson, J.E. & Gardarsson, A. 2001: Pair formation in relation to climate: mallard, Eurasian wigeon and Eurasian teal wintering in Iceland. — Wildfowl 52: 55–68.
Jorde, D.G., Krapu, G.L., Crawford, R.D. & Hay, M.A. 1984: Effects of weather on habitat selection and behavior of Mallards wintering in Nebraska. — Condor 86: 258–265. https://doi.org/10.2307/1366993
Jónsson, J.E. 2010: Sex ratios of Eurasian wigeon, mallard and common eider in Iceland. — Náttúrufræðingurinn 79: 118–124. (in Icelandic with English summary)
Kaminski, R.M. & Prince, H.H. 1981: Dabbling duck activity and foraging responses to aquatic macroinvertebrates. — Auk 98: 115–126. https://doi.org/10.2307/3807868
Kendeigh S.C. 1970: Energy requirements for existence in relation to size of bird. — Condor 72: 60–65. https://doi.org/10.2307/1366475
Kestenholtz, M. 1990: Verteilungsmuster von Stock-, Reiher- und Tafelente, Gänsesäger und Blässhuhn im Winterhalbjahr am Sempachersee. — Ornithologische Beobachter 87: 131–145.
Kosztolányi, A., Barta, Z., Kupper, C. & Székely, T. 2011: Persistence of an extreme male-biased adult sex ratio in a natural population of polyandrous bird. — Journal of Evolutionary Biology 24: 1842–1846. https://doi.org/10.1111/j.1420-9101.2011.02305.x
Korzeniewski, K. 1993: The Puck Bay. — Foundation for the Development of the University of Gdańsk, Gdańsk. (in Polish)
Krishnan, G., Devaraj, C., Silpa, M.V. & Sejian, V. 2023: Thermoregulation in Birds. In Textbook of veterinary physiology (ed. Das, P.K., Sejian, V., Mukherjee, J. & Banerjee, D.). Springer, Singapore. https://doi.org/10.1007/978-981-19-9410-4_29
Kruk-Dowgiałło, L. 1998: Phytobenthos as an indicator of the state of environment of the Gulf of Gdańsk. — Oceanological Studies 4: 105–121.
Larsson, K. 2022: Age and sex ratios in the declining West Siberian/North European population of Long-tailed Duck wintering in the Baltic Sea: Implications for conservation. — Ornis Fennica 99: 117–131. https://doi.org/10.51812/of.113681
Lee, A.M., Sæther, B.E. & Engen, S. 2011: Demographic stochasticity, allee effects, and extinction: the influence of mating system and sex ratio. — American Naturalist 177: 301–313. https://doi.org/10.1086/658344
Lehikoinen, A., Christensen, T.K., Öst, M., Kilpi, M., Saurola, P. & Vattulainen, A. 2008: Large-scale change in the sex ratio of a declining eider Somateria mollissima population. — Wildlife Biology 14: 288–301. https://doi.org/10.29.81/0909-6396(2008)14[288:lcitsr]2.0.co;2
Lenth, R.V. 2021: emmeans: Estimated Marginal Means, aka Least-Squares Means. R package version 1.6.1. — https://cran.r-project.org/web/packages/emmeans/index.html
Loring, P.H., Paton, P.W.C., McWilliams, S.R., McKinney, R.A. & Oviatt, C.A. 2013: Densities of wintering scoters in relation to benthic prey assemblages in a North Atlantic estuary. — Waterbirds 36, 144–155. https://doi.org/10.1675/063.036.0204
Lüdecke, D., Ben-Shachar, M., Patil, I., Waggoner, P. & Makowski, D. 2021: performance: An R Package for Assessment, Comparison and Testing of Statistical Models. — Journal of Open Source Software 6: 3139. https://doi.org/10.21105/joss.03139
Majewski, A. (ed.) 1990: Gulf of Gdańsk. — Wydawnictwa Geologiczne, Warszawa. (in Polish)
McCullagh, P. & Nelder, J.A. 1989: Generalized Linear Models. — Chapman and Hall, London. https://doi.org/10.1007/978-1-4899-3242-6
Meissner, W., Ciopcińska, K. 2007: Behaviour of Mute Swans Cygnus olor wintering at a municipal beach in Gdynia, Poland. — Ornis Svecica 17: 148–153. https://doi.org/10.34080/os.v17.22685
Meissner, W. & Krupa, R. 2017: Sex-related differences in autumn migration timing of adult common sandpipers Actitis hypoleucos (Linnaeus, 1758) (Charadriiformes: Scolopacidae). — European Zoological Journal 84: 136–140. https://doi.org/10.1080/11250003.2016.1278474
Meissner, W. & Markowska, K. 2009: Influence of low temperatures on behaviour of mallards (Anas platyrhynchos L.). — Polish Journal of Ecology 57: 799–803.
Meissner, W., Rowiński, P., Polakowski, M., Wilniewczyc, P. & Marchowski, D. 2015: Impact of temperature on the number of mallards, Anas platyrhynchos, wintering in cities. — North-Western Journal of Zoology 11: 213–218.
Meissner, W. & Witkowska, M. 2023: The effect of the temperature on local differences in the sex ratio of Mallards Anas platyrhynchos wintering in an urban habitat. — Acta Oecologica 119: 103900. https://doi.org/10.1016/j.actao.2023.103900
Miernik, N.A., Janas, U. & Kendzierska, H. 2023: Role of macrofaunal communities in the Vistula River plume, the Baltic Sea—Bioturbation and bioirrigation potential. — Biology 12: 147. https://doi.org/10.3390/biology12020147
Munday, C. & Rose, P. 2022: Environmental and social influences on the behaviour of free-living Mandarin Ducks in Richmond Park. — Animals 12: 2554. https:// doi.org/10.3390/ani12192554.
Nichols, J.D., Haramis G.M., 1980: Sex-specific differences in winter distribution patterns of Canvasback, — Condor 82: 406–416. https://doi.org/10.2307/1367565
Nilsson, L. 1970a: Local and seasonal variation in sex-ratios of diving ducks in South Sweden during the nonbreeding season. — Ornis Scandinavica 1: 115–128. https://doi.org/10.2307/3676029
Nilsson, L. 1970b: Food seeking activity of south Swedish diving ducks in the non-breeding season. — Oikos 21: 145–154. https://doi.org/10.2307/3543670
Nilsson. L. 1976: Sex-ratios of Swedish Mallard during the non-breeding season. — Wildfowl 27: 91–94.
Owen, M. & Black, J.M. 1990: Waterfowl Ecology. — Blackie and Son, Ltd., Glasgow.
Owen, M. & Dix, M. 1986: Sex ratios in some common British wintering ducks. — Wildfowl 37: 104–112.
Pap, K., Nagy, L., Balogh, C., G.-Tóth, L. & Liker, A. 2013: Environmental factors shaping the distribution of common wintering waterbirds in a lake ecosystem with developed shoreline. — Hydrobiologia 716: 163–176. https://doi.org/10.1007/s10750-013-1560-3
Pattenden, R.K. & Boag, D.A. 1989: Skewed sex ratio in a northern wintering population of mallards. — Canadian Journal of Zoology 67: 1084–1087. https://doi.org/10.1139/z89-152
Pounder, B. 1976: Wintering flocks of Goldeneyes at sewage outfalls in the Tay Estuary. — Bird Study 23: 121–131. https://doi.org/10.1080/00063657609476490
Pöysä, H., Linkola, P., Paasivaara, A. 2019. Breeding sex ratios in two declining diving duck species: between‑year variation and changes over six decades. — Journal of Ornithology 160:1015–1023. https://doi.org/10.1007/s10336-019-01682-7
Rodway, M.S. 2006: Have winter spacing patterns of harlequin ducks been partially shaped by sexual selection? — Waterbirds 29: 415–426. https://doi.org/10.1675/1524-4695(2006)29[415:hwspoh]2.0.co;2
Rodway, M.S. 2007. Timing of pairing in waterfowl II: Testing the hypotheses with Harlequin Ducks. — Waterbirds 30: 506–520. https://doi.org/10.1675/1524-4695(2007)030[0506:topiwi]2.0.co;2
Rodway, M.S., Regehr, H.M., Boyd, W.S. & Iverson, S.A. 2015: Age and sex ratios of sea ducks wintering in the Strait of Georgia, British Columbia: Implications for monitoring. — Marine Ornithology 43: 141–150.
Rohwer, F.C. & Anderson, M.C. 1988: Female biased philopatry, monogamy, and the timing of pair formation in migratory waterfowl. — Current Ornithology 5: 187–221. https://doi.org/10.1007/978-1-4615-6787-5_4
Ruben, J. 1996: Evolution of endothermy in mammals, birds and their ancestors. — In Animals and temperature: phenotypic and evolutionary adaptation (ed. Johnston, I.A. & Bennett, A.F.): 347–376. Cambridge University Press, Cambridge.
Sayler, R.D. & Afton, A.D. 1981: Ecological aspects of Common Goldeneyes Bucephala clangula wintering on the upper Mississippi River. — Ornis Scandinavica 12: 99–108. https://doi.org/10.2307/3676033
Schummer, M.L., Anthony, A.B., Kleespies, S.M., Ankenman, G., Ligouri, M., Bleau, A., Droke, J., Cohen, J., Kowalski, K., Morlock, F. & Eckler, J. 2020: Aggression and behavioural dominance in wintering mallard Anas platyrhynchos and American black Duck A. rubripes. — Wildfowl 70: 167–178.
Serrano-Davies, E., Traba, J., Arroyo, B., Mougeot, F., Cuscó, F., Mañosa, S., Bota, G., Faria, N., Villers, A., Casas, F., Attie, C., Devoucoux, P., Bretagnolle, V. & Morales, M.B. 2022: Biased adult sex ratios in Western Europe populations of Little Bustard Tetrax tetrax as a potential warning signal of unbalanced mortalities. — Bird Conservation International 33: e40. https://doi.org/10.1017/S0959270922000430
Skov, H., Heinänen, S., Žydelis, R., Bellebaum, J., Bzoma, S., Dagys, M., Durinck, J., Garthe, S., Grishanov, G., Hario, M., Kieckbusch, J.J., Kube, J., Kuresoo, A., Larsson, K., Luigujoe, L., Meissner, W., Nehls, H.W., Nilsson, L., Petersen, I.K., Roos, M.M., Pihl, S., Sonntag, N., Stock, A., Stipniece, A., Wahl, J. 2011: Waterbird populations and pressures in the Baltic Sea. — Nordic Council of Ministers. Copenhagen.
Skov, H. & Kube, J. 1996: Habitat selection, feeding characteristics, and food consumption of long-tailed ducks, Clangula hyemalis, in the southern Baltic Sea. — In: The ecology of macrozoobenthos and sea ducks in the Pomeranian Bay (ed. Kube, J.). Meereswissenschaftliche Berichte 18: 83–100.
Sokołowski, A., Jankowska, E., Balazy, P. & Jędruch, A. 2021: Distribution and extent of benthic habitats in Puck Bay (Gulf of Gdańsk, southern Baltic Sea). — Oceanologia 63: 301–320. https://doi.org/10.1016/j.oceano.2021.03.001
Söderholm, S. 2003: Könsfördelning hos övervintrande brunand Aythya ferina i Erstaviken, Sörmland. — Ornis Svecica 13: 17–23. https://doi.org/10.34080/os.v13.22816
Székely, T., Liker, A., Freckleton, R.P., Fichtel, C. & Kappeler, P.M. 2014: Sex-biased survival predicts adult sex ratio variation in wild birds. — Proceedings of the Royal Society B: Biological Sciences 281: 20140342. https://doi.org/10.1098/rspb.2014.0342
Wetlands International 2015: Waterbird Population Estimates. — http://wpe.wetlands.org/
Wood, K.A., Brides, K., Durham, M.E., Hearn, R.D. 2021: Adults have more male-biased sex ratios than first-winter juveniles in wintering duck populations. — Avian Research 12: 51. https://doi.org/10.1186/s40657-021-00286-1
Woolington, D.W. 1993: Sex ratios of canvasbacks wintering in Louisiana. — Journal of Wildlife Management 57: 751–758. https://doi.org/10.2307/3809075
Zar, J.H. 2010: Biostatistical analysis. — Prentice Hall, Upper Saddle River, New Jersey.
Downloads
Additional Files
Published
Issue
Section
License
Copyright (c) 2022 Włodzimierz Meissner, Dorota Kozakiewicz

This work is licensed under a Creative Commons Attribution 4.0 International License.


