Explaining individual variation in patterns of mass loss in breeding birds
Tóm tắt
Studies of birds have a disproportionate representation in the literature on life-history evolution, because of the (apparent) ease with which the costs and benefits can be quantified and manipulated. During reproduction, birds frequently show a highly conserved pattern of mass change and changes in mass loss during breeding have been widely considered to be a valid short-term measure of the costs of reproduction. Experimental manipulations of the breeding attempts of birds usually argue that the presence of a response shows that a cost of reproduction exists, but there is little consensus as to how the size of these costs can be measured. We model this mass loss by considering how a parent can maximise its lifetime reproductive success, using a theoretical framework that is particularly suited to modelling parental care in altricial birds. If lifetime reproductive success is taken to be the sum of a parent's current and future reproductive success, we show that the exact forms of these components will influence the optimal amount of mass a parent should lose. In particular, we demonstrate that the shape of the relationship between parental investment and chick survival will lead to differing degrees of investment between parents of different initial qualities: parents with initially high levels of energy reserves could conceivably invested a lesser, similar or greater amount of resources than parents with initially low reserves, and these initially 'heavy' parents could potentially end up being lighter than the initially 'lighter' individuals. We argue that it is difficult to make predictions about the dependence of a parent's final mass on its initial mass, and therefore mass loss should only be used as a short-term measure of the costs of reproduction with caution. The model demonstrates that we require a better understanding of the relationship between mass loss and both current and future reproductive success of the parent, before predictions about mass loss can be made and tested. We discuss steps that could be taken to increase the accuracy of our predictions.
Tài liệu tham khảo
Baldwin SP, Kendeigh SC: Variations in the weight of birds. Auk. 1938, 55: 416-467.
Rands SA, Houston AI, Cuthill IC: Measurement of mass change in breeding birds: a bibliography, and a discussion of measurement techniques. Ring Migrat.
Moreno J: Strategies of mass change in breeding birds. Biol J Linn Soc. 1989, 37: 297-310.
Ricklefs RE: Energetics of reproduction in birds. Avian energetics. Edited by: Paynter RA, jr. 1974, Cambridge: Nuttall Ornithological Club, 152-297.
Murton RK, Westwood NJ: Avian breeding cycles. 1977, Oxford: Oxford University Press
Freed LA: Loss of mass in breeding wrens: stress or adaptation?. Ecology. 1981, 62: 1179-1186. 10.2307/1937282.
Bryant DM: Energy expenditure and body mass changes as measures of reproductive costs in birds. Funct Ecol. 1988, 2: 23-34.
Hillström L: Body mass reduction during reproduction in the pied flycatcher Ficedula hypoleuca: physiological stress or adaptation for lowered costs of transport?. Funct Ecol. 1995, 9: 807-817.
Nur N: The cost of reproduction in birds: an examination of the evidence. J Anim Ecol. 1988, 76: 155-168.
Norberg RÅ: Temporary weight decrease in breeding birds may result in more fledged young. Am Nat. 1981, 118: 838-850. 10.1086/283874.
Nur N: The consequences of brood size for breeding blue tits I. Adult survival, weight change and the cost of reproduction. J Anim Ecol. 1984, 53: 479-496.
Cavitt JF, Thompson CF: Mass loss in breeding house wrens: effects of food supplements. Ecology. 1997, 78: 2512-2523. 10.2307/2265910.
Johnsen I, Erikstad KE, Sæther B-E: Regulation of parental investment in a long-lived seabird, the puffin Fratercula arctica: an experiment. Oikos. 1994, 71: 273-278.
Velando A, Alonso-Alvarez C: Differential body condition regulation by males and females in response to experimental manipulations of brood size and parental effort in the blue-footed booby. J Anim Ecol. 2003, 72: 846-856. 10.1046/j.1365-2656.2003.00756.x.
Johnston RD: The effect of direct supplementary feeding of nestlings on weight loss in female great tits Parus major. Ibis. 1993, 135: 311-314.
Rands SA: Dynamic reserve allocation in breeding birds. PhD thesis. 2000, University of Bristol, School of Biological Sciences
Williams GC: Natural selection, the costs of reproduction, and a refinement of Lack's principle. Am Nat. 1966, 100: 687-690. 10.1086/282461.
Witter MS, Cuthill IC: The ecological costs of avian fat storage. Phil Trans R Soc Lond B. 1993, 340: 73-92.
Cuthill IC, Houston AI: Managing time and energy. Behavioural ecology: an evolutionary approach. Edited by: Krebs JR, Davies NB. 1997, Oxford: Blackwell Science, 97-120. fourth
McNamara JM, Houston AI: The value of fat reserves and the tradeoff between starvation and predation. Acta Biotheor. 1990, 38: 37-61. 10.1007/BF00047272.
Rands SA, Cuthill IC: Separating the effects of predation risk and interrupted foraging upon mass changes in the blue tit Parus caeruleus. Proc R Soc Lond B. 2001, 268: 1783-1790. 10.1098/rspb.2001.1653.
Houston AI: The efficiency of mass loss in breeding birds. Proc R Soc Lond B. 1993, 254: 221-225.
McNamara JM, Houston AI: Risk-sensitive foraging: a review of the theory. Bull Math Biol. 1992, 54: 355-378. 10.1016/S0092-8240(05)80031-X.
Houston AI, McNamara JM: Models of adaptive behaviour: an approach based on state. 1999, Cambridge: Cambridge University Press
Källander H, Smith HG: Manipulation of the brood size of pied flycatchers. Population biology of passerine birds: an integrated approach. Edited by: Blondel J, Gosler A, Lebreton J-D, McCleery R. 1990, Berlin: Springer-Verlag, 257-268.
Slagsvold T, Johansen MA: Mass loss in female pied flycatchers Ficedula hypoleuca during late incubation: supplementation fails to support the reproductive stress hypothesis. Ardea. 1998, 86: 203-211.
Cichoń M, Merilä J, Hillström L, Wiggins D: Mass-dependent mass loss in breeding birds: getting the null hypothesis right. Oikos. 1999, 87: 191-194.
Gebhardt-Henrich SG: When heavier birds lose more mass during breeding: statistical artefact or biologically meaningful?. J Avian Biol. 2000, 31: 245-246. 10.1034/j.1600-048X.2000.310216.x.
Ruf T: Mass-dependent mass loss: how to get the null hypothesis wrong. Oikos. 2000, 90: 413-416. 10.1034/j.1600-0706.2000.900223.x.
Kelly C, Price TD: Correcting for regression to the mean in behavior and ecology. Am Nat. 2005, 166: 700-707. 10.1086/497402.
Ricklefs RE: Weight recession in nestling birds. Auk. 1968, 85: 30-35.
Winkler DW: A general model for parental care. Am Nat. 1987, 130: 526-543. 10.1086/284729.
Ekman J, Rosander B: Survival enhancement through food sharing: a means for parental control of natal dispersal. Theor Popul Biol. 1992, 42: 117-129. 10.1016/0040-5809(92)90008-H.
Mock DW, Parker GA: The evolution of sibling rivalry. 1997, Oxford: Oxford University Press
Gebhardt-Henrich SG, Richner H: Causes of growth variation and its consequences for fitness. Avian growth and development: evolution within the altricial-precocial spectrum. Edited by: Starck JM, Ricklefs RE. 1998, New York: Oxford University Press, 324-339.
Schew WA, Ricklefs RE: Developmental plasticity. Avian growth and development: evolution within the altricial-precocial spectrum. Edited by: Starck JM, Ricklefs RE. 1998, New York: Oxford University Press, 288-304.
Mangel M, Munch SB: A life-history perspective on short- and long-term consequences of compensatory growth. Am Nat. 2005, 166: E155-E176. 10.1086/444439.
Lack D: Ecological adaptations for breeding in birds. 1968, London: Methuen
Ricklefs RE, Starck JM, Konarzewski M: Internal constraints on growth in birds. Avian growth and development: evolution within the altricial-precocial spectrum. Edited by: Starck JM, Ricklefs RE. 1998, New York: Oxford University Press, 266-287.
Case TJ: On the evolution and adaptive significance of postnatal growth rates in the terrestrial vertebrates. Q Rev Biol. 1978, 55: 243-282. 10.1086/410622.
Winkler DW, Adler FR: Dynamic state variable models for parental care: I. A submodel for the growth of the chicks of passerine birds. J Avian Biol. 1996, 27: 343-353.
Merilä J, Wiggins DA: Mass loss in breeding blue tits: the role of energetic stress. J Anim Ecol. 1997, 66: 452-460.
Lessells CM: The cost of reproduction: do experimental manipulations measure the edge of the options set?. Etología. 1993, 3: 95-111.
Monaghan P, Nager RG: Why don't birds lay more eggs?. Trends Ecol Evol. 1997, 12: 270-274. 10.1016/S0169-5347(97)01094-X.
Hanssen SA, Erikstad KE, Johnsen V, Bustnes JO: Differential investment and costs during avian incubation determined by individual quality: an experimental study of the common eider (Somateria mollissima). Proc R Soc Lond B. 2003, 270: 531-537. 10.1098/rspb.2002.2262.
Drent RH, Daan S: The prudent parent: energetic adjustments in avian breeding. Ardea. 1980, 68: 225-252.
Hanssen SA, Folstad I, Erikstad KE: Reduced immunocompetence and cost of reproduction in common eiders. Oecologia. 2003, 136: 457-464. 10.1007/s00442-003-1282-8.
Tieleman BI, Williams JB, Ricklefs RE: Nest attentiveness and egg temperature do not explain the variation in incubation periods in tropical birds. Funct Ecol. 2004, 18: 571-577. 10.1111/j.0269-8463.2004.00882.x.
Clark CW, Mangel M: Dynamic state variable models in ecology: methods and applications. 2000, New York: Oxford University Press