Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Alan G. Leach; Amanda W. van Dellen; Thomas V. Riecke; James S. Sedinger

doi:10.1111/ibi.12480

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Alan G. Leach
, Amanda W. van Dellen
, Thomas V. Riecke
, James S. Sedinger

W. A. Franke College of Forestry and Conservation

University of Nevada, Reno

Research output: Contribution to journal › Article › peer-review

5 Scopus citations

Abstract

Lack () proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per-capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two-egg clutches to 0.50 for seven-egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three-egg clutches to 27.7 days (95% CI 27.3–28.1) for six-egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five-egg clutch.

Original language	English
Pages (from-to)	588-599
Number of pages	12
Journal	Ibis
Volume	159
Issue number	3
DOIs	https://doi.org/10.1111/ibi.12480
State	Published - Jul 2017

Funding

We thank the following people for assisting with work at Tutakoke River during this study: M. Blom, B. Cheatham, A. Clark, A. Ganick, D. Gibson, E. Huskinson, H. Johnson, D. Messmer, M. Schreiner, H. Singer, L. Snoddy, A. Stewart, E. Willey and N. Yeldell. Studies at the Tutakoke River Brent colony were funded by the U.S. Geological Survey, Alaska Science Center; U.S. Fish and Wildlife Service, Migratory Bird Management Region 7; Ducks Unlimited; The Morro Bay Brant Group; Phil Jebbia (in memory of Marnie Shepherd); and the National Science Foundation (OPP 9214970, DEB 9815383, OPP 9985931, OPP 0196406, DEB 0743152, DEB 1252656). All handling of animals was approved by Institutional Animal Care and Use committees at the University of Alaska Fairbanks and the University of Nevada, Reno (most recent protocol number 00056). Logistical support was provided by staff from the Yukon Delta National Wildlife Refuge. M. Irinaga and L. Gullingsrud of CH2M Hill Polar Services provided additional logistical field support. We are grateful to Todd Arnold and two anonymous reviewers whose comments greatly improved an earlier version of the manuscript. Ruedi Nager suggested using mixed effects models for hatching and egg success.

Funders	Funder number
DEB 1252656, DEB 0743152, OPP 0196406, OPP 9214970, OPP 9985931, DEB 9815383
University of Nevada, Reno	00056
University of Alaska Fairbanks

Keywords

Branta bernicla nigricans
egg survival
hatchability
incubation limitation hypothesis
life-history trade-offs
nesting period
optimal clutch size

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10.1111/ibi.12480

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title = "Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans",

abstract = "Lack () proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per-capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two-egg clutches to 0.50 for seven-egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95\% CI 25.1–25.7) for three-egg clutches to 27.7 days (95\% CI 27.3–28.1) for six-egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five-egg clutch.",

keywords = "Branta bernicla nigricans, egg survival, hatchability, incubation limitation hypothesis, life-history trade-offs, nesting period, optimal clutch size",

author = "Leach, \{Alan G.\} and \{van Dellen\}, \{Amanda W.\} and Riecke, \{Thomas V.\} and Sedinger, \{James S.\}",

note = "Publisher Copyright: {\textcopyright} 2017 British Ornithologists{\textquoteright} Union",

year = "2017",

month = jul,

doi = "10.1111/ibi.12480",

language = "English",

volume = "159",

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T1 - Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

AU - Leach, Alan G.

AU - van Dellen, Amanda W.

AU - Riecke, Thomas V.

AU - Sedinger, James S.

PY - 2017/7

Y1 - 2017/7

N2 - Lack () proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per-capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two-egg clutches to 0.50 for seven-egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three-egg clutches to 27.7 days (95% CI 27.3–28.1) for six-egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five-egg clutch.

AB - Lack () proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per-capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two-egg clutches to 0.50 for seven-egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three-egg clutches to 27.7 days (95% CI 27.3–28.1) for six-egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five-egg clutch.

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KW - egg survival

KW - hatchability

KW - incubation limitation hypothesis

KW - life-history trade-offs

KW - nesting period

KW - optimal clutch size

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U2 - 10.1111/ibi.12480

DO - 10.1111/ibi.12480

M3 - Article

AN - SCOPUS:85019098110

SN - 0019-1019

VL - 159

SP - 588

EP - 599

JO - Ibis

JF - Ibis

IS - 3

ER -

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

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