ThesisPDF Available

Evolutionary strategies behind maternal investment in mammalian reproduction

Authors:
Evolutionary strategies behind
maternal investment in
mammalian reproduction
Tim E.R.G. Huijsmans
Evolutionary strategies behind maternal investment in mammalian reproduction I Tim E.R.G. Huijsmans
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Evolutionary strategies behind maternal
investment in mammalian reproduction
Tim E.R.G. Huijsmans
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Promoters
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© Tim E.R.G. Huijsmans
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Members of the jury
Prof. Dr. Dr. h.c. Ann van Soom, DVM, PhD, Dipl. ECAR, promoter
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Prof. Dr. Katrien Smits, DVM, PhD, promoter
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Dr. Jella Wauters, DVM, PhD, promoter
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Prof. Dr. Thomas B. Hildebrandt, DVM, PhD, Dipl. ECZM, promoter
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Prof. Dr. Ann Martens, DVM, PhD, Dipl. ECVS, chairman
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Prof. Dr. Frank Pasmans, DVM, PhD, Dipl. ECZM, secretary
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Prof. Dr. Bruno Levecke, DVM, PhD
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Prof. Dr. Tom Stout, DVM, PhD, Dipl. ECAR
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Prof. Dr. Ruth Appeltant, DVM, PhD
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Dr. Cyriel Ververs, DVM, PhD, Dipl. ECAR
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Table of contents
List of abbreviations
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List of evolutionary biology definitions
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List of figures
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List of tables
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Chapter 1
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Chapter 2
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Chapter 3
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Chapter 4
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Chapter 5
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Chapter 6
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Chapter 7
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Summary
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Samenvatting!
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Acknowledgements
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Curriculum vitae!
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Bibliography!
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Addendum 1
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1.1
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1.1
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Chapter 1
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General introduction
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5Didelphis virginiana,6!/09!43'!H''&@/4!5Suricata suricatta,!5V4'/9!'4!/1)6!EWEE,)!!
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MONOTREME
Tachyglossus aculeatus
METATHERIAN
Didelphis virginiana
EUTHERIAN
Suricata suricatta
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Monotremata: T.a. = Tachyglossus aculeatus (Short-beaked echidna), O.a. = Ornithorhynchus anatinus (Platypus); Metatheria: S.m. =
Sminthopsis macroura (Stripe faced dunnart), T. v. = Trichosurus vulpecula (Common brushtail possum), D.v. = Didelphis virginiana
(Virginia opossum); Eutheria: M.m. = Mus musculus (mouse); O.c. = Oryctolagus cuniculus (European rabbit), E.m. = Elephantulus
myurus (Eastern rock elephant shrew).
!
1.2.1.1 Monotremata
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1 Trends Desk, 2022. Here’s ‘the oldest & the only elephant to be living beyond 100 years’
https://indianexpress.com/article/trending/trending-in-india/heres-the-oldest-the-only-elephant-to-be-
living-beyond-100-years-7993390/, Visited 21/12/2023
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Chapter 3
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Quantifying maternal investment in
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./>!;.'4(5S9QD!#$"#m!E35Q.D!"OO"m!7.'4(/!94!.=*D!#$$fl*!79!4I85!'9=39>!(/!53e!.=49'/.43K9!
'9M'9553(/!1(>9=5!4(!95431.49!4I9!=3449'!1.55!.4!]9./3/M*!0@9C3aC.==GD!]9!859>!.!531@=9!
=3/9.'! '9M'9553(/!k0RElD! .! =3/9.'! kI949'(5C9>.543Cl!13e9>c9Y9C45! 1(>9=! .CC(8/43/M! )('!
@IG=(M9/943C!3/9'43.! k&RBBlD!.!54./>.'>359>! 1.g('!.e35! '9M'9553(/!k0B,lD! .! 1.g('!.e35!
'9M'9553(/! kB,lD! .! 18=43@=9! =3/9.'! '9M'9553(/! 1(>9=! '9@'959/43/M! ./! 9e49/53(/! ()! 4I9!
1(>9=! 0RE! kB0RElD! ./>! .! 18=43@=9! =3/9.'! '9M'9553(/! '9@'959/43/M! ./! 9e49/53(/! ()! 4I9!
1(>9=! &RBB! kB&RBBl*! W/! 4(@! ()! 4I9! 9Y9C4! ()! 4I9! .>8=4! 1.55! (/! 4I9! =3449'! 1.55! .4!
]9./3/M!C(/53>9'9>! 3/!4I9!?3K.'3.49! 1(>9=5!k0RED! &RBBD!0B,!./>!B,lD!4I9! B0RE!./>!
B&RBB!1(>9=5!.=5(!.CC(8/4!)('!.!@(553?=9!9Y9C4!()!3/K95419/4!>8'.43(/*!:I9!'.43(/.=9!
?9I3/>!4I959!18=43@=9!=3/9.'!'9M'9553(/!1(>9=5!k3*9*!B0RE!./>!B&RBBl!35!4(!.CC(8/4!)('!
! ! !
ON!
!
4I9!@(553?3=34G!4I.4!./!9e49/>9>!1.49'/.=!3/K95419/4!3/!(Y5@'3/M!C(8=>!'958=4!3/!M'9.49'!
(Y5@'3/M!M'(]4I*!:I35!@(553?3=34G!35!/(4!a priori (?K3(85!53/C9!4I9!9Y9C4!()!4I9!3/K95419/4!
>8'.43(/! 1.G! .='9.>G! ?9! .CC(8/49>! )('! 3/! 4I9! 531@=9'! 1(>9=5D! K3.! 4I9! .>8=4! 1.55!
@'9>3C4('D!>89!4(!4I9!.==(194'3C!'9=.43(/5I3@!?94]99/!?(>G!1.55!./>!4I9!@.C9!()!=3)9*!:I9!
'.43(/.=9!?9I3/>!4I9!1(>9=5!&RBB!./>!B&RBB!35!4I.4D!)('!.!M3K9/!.>8=4!1.55D!4I9!=3449'!
1.55!.4!]9./3/M!1.G!>3Y9'!?94]99/!4.e.!>89!4(!4I9!3/d89/C9!()!).C4('5!C(''9=.49>!]34I!
@IG=(M9/G! 58CI! .5! 4I9! M9/943C! 1.Q9c8@! ()! 3/>3K3>8.=5! ('! 4I93'! 9C(=(MG*! :I9'9)('9D!
.CC(8/43/M!)('!4I9! @IG=(M9/943C!>354./C95!?94]99/! 5@9C395!]I3=9!a443/M! 4I9!.==(194'3C!
'9=.43(/5I3@!>9C'9.595!4I9! 31@.C4!()!K.'3.43(/!3/!4I9! >9/534G!()!(?59'K.43(/5! .=(/M!4I9!
1.11.=3./!@IG=(M9/943C!4'99!(/!95431.495!@'(>8C9>!?G!'9M'9553(/!194I(>5*!
79!a449>! 4I9!=3/9.'!'9M'9553(/5! (/!.! 53/M=9!58?594!()!(8'!>.4.! k[!z! _NPl!4I.4D!)('! 9.CI!
5@9C395D!C(/4.3/9>!3/)('1.43(/!.?(84!4I9!=3449'! 1.55!.4!]9./3/MD!4I9!.>8=4!?(>G!1.55D!
4I9!3/K95419/4!>8'.43(/!./>!@IG=(M9/943C!@(5343(/!kJ@I.1!94!.=*D!#$"Ol*!:I9!95431.495!()!
4I9!.==(194'3C!'9=.43(/5I3@!(?4.3/9>!)('!4I9!>3Y9'9/4!'9M'9553(/!194I(>5!.'9!>9@3C49>!3/!
Figure 3.1!./>!5811.'359>!3/!Table 3.1*!
!
N/D1'! L)(*! ?04'&.'248! /09! /11:H'4&#.! 8./1#0$! .:'O#.#'048! :-! 8#I! 9#O'&'04! 1#0'/&! &'$&'88#:0! H:9'18!
Y44'9!4:!43'!8/H'!9/4/8'4!:-!H/HH/1#/0!82'.#'8!5c!f!JLa,)!Ha!&'2&'8'048!43'!9'-/%14!/9%14!H/88!/09!
9!43'!#0G'84H'04!9%&/4#:0)!+11!&'$&'88#:0!H'43:98!1'9!4:!/!$::9!/9h%84H'04!:-!43'!9/4/!5&E!i!W)MXJ6!2!
j!W)WW(,)!c:4'!43/4!#04'&.'248!$#G'0!3'&'!.:&&'82:09!4:!43'!'84#H/4'8!9#&'.41>!$#G'0!D>!H:9'1!Y48)!N3'>!
43%8!.:&&'82:09!4:!43'!1:./4#:0!A3'&'!43'!&'$&'88#:0!1#0'!A:%19!.%4!43'!1:$10R4&/08-:&H'9!>R/I#8)!
Model
Intercept (CI95%)
Scaling ma (CI95%)
Scaling d (CI95%)
0RE!
c$*"Of!kc$*#"fD!c$*"_fl!
$*__P!k$*_f^D!$*_O"l!
c!
&RBB
!
c$*"O#!kc$*^N%D!$*"_Nl
!
!$*P$f!k$*_P#D!$*PNNl
!
c
!
0B,!
c$*"O$!kc$*#"$D!c$*"_$l!
$*_OO!k$*_PfD!$*P"Nl!
c!
B,!
c$*"O#!kc$*#"#D!c$*"_#l!
!$*_O^!k$*_P"D!$*P$Pl!
c!
B0RE!
$*ff%!k$*^$PD!$*P"Ol!
$*Pf_!k$*P%_D!$*PP_l!
c$*NOP!kc$*%_$D!c$*N#_l!
B&RBB!
$*#$%!kc$*#^^D!$*f#Pl!
$*PNf!k$*P$%D!$*P_$l!
c$*"OO!kc$*NN%D!!
c$*$_$"l!
!
O%!
!
J53/M!4I9!53e!a449>!1(>9=5D!4I9!9e@9C49>!=3449'!1.55!.4!]9./3/M!)('!.!M3K9/!5@9C395!C./!
?9! @'9>3C49>! ?.59>! (/! 4I9! .>8=4! 1.55D! ./>! 3/! C.59! ()! 4I9! 18=43@=9! =3/9.'! '9M'9553(/!
1(>9=5D! ?.59>! (/! 4I9! .>8=4! 1.55! ./>! 3/K95419/4! >8'.43(/D! )('! 4I.4! 5@9C395*! ;G!
C(1@.'3/M!4I9!.C48.=! =3449'!1.55! .4!]9./3/M!4(! 4I9!@'9>3C49>!=3449'! 1.55!.4!]9./3/MD! .!
194'3C!C./! ?9!@'(>8C9>!4(! .==(]!)('! 4I9!(?g9C43K9!C(1@.'35(/! ()!1.49'/.=! 3/K95419/4!
?94]99/!5@9C395*!79!@'(@(59!4I9!)(==(]3/M!1.49'/.=!3/K95419/4!194'3C5!?.59>!(/!4I9!
'.43(! ?94]99/! 4I959! Z8./434395`! 𝑀𝐼 =𝑙𝑜𝑔%&"'!()*+),-."//)*-0#((-#/ -1)#2"23
."//)*-0#((-#/-1)#2"23-4*),"$/),-!5-#..'0)/*5D!
]I9'9!MI!35! 4I9! 1.49'/.=!3/K95419/4!k.! 8/34=955! /81?9'!53/C9!•QMŽ×•QMŽ-1z•"Žl*! :I9! 53e!
1.49'/.=! 3/K95419/4! 194'3C5! 549113/M! )'(1! (8'! 53e! =3/9.'! '9M'9553(/! 1(>9=5! .'9!
@'959/49>!3/!Table 3.2*!!
!
N/D1'! L)E*! FG'&G#'A! :-! 43'! H/4'&0/1! #0G'84H'04! H'4&#.8! #0! H/HH/18! /..:&9#0$! 4:! 8#I! /14'&0/4#G'!
&'$&'88#:0!H:9'18!Y44'9!4:!43'!8/H'!9/4/8'4!5c!f!JLa!82'.#'8,)!N3'!#04'&.'24!.:&&'82:098!4:!43'!1:$10!
4&/08-:&H'9!#04'&.'24!#0!N/D1'!L)()!V''!1'$'09!N/D1'!L)(!-:&!H:&'!9'4/#18)!
Model
Metric
0RE!
𝑀𝐼678 =𝑙𝑜𝑔%&
𝑚.1
0.637 × 𝑚#
&9::;"
&RBB!
𝑀𝐼<7== =𝑙𝑜𝑔%&"𝑚.1
0.643 × 𝑚#
&9;&>"
0B,!
𝑀𝐼6=? =𝑙𝑜𝑔%&
𝑚.1
0.645 × 𝑚#&9:@@"
B,!
𝑀𝐼=? =𝑙𝑜𝑔%&"𝑚.1
0.643 × 𝑚#&9:@A"
B0RE!
𝑀𝐼=678 =𝑙𝑜𝑔%&
𝑚.1
4.61 × 𝑚#&9;>: × 𝑑B&9C@; "
B&RBB!
𝑀𝐼=<7== =𝑙𝑜𝑔%&"𝑚.1
1.60 × 𝑚#&9;C> × 𝑑B&9%@@ "
!
:I9!MI!194'3C5!>3'9C4=G!C(''95@(/>!4(!4I9!'953>8.=5!.55(C3.49>!]34I!4I9!'9M'9553(/!=3/95!
5I(]/!3/!Figure 3.1*!F('!?3K.'3.49!1(>9=5!853/M! (/=G!.!53/M=9! @'9>3C4('!k.>8=4!1.55lD! .!
5@9C395!CI.'.C49'359>!?G!MI!K.=89!()!$!35!4I85!.!5@9C395!4I.4!>(95!/(4!>9K3.49!)'(1!4I9!
'9M'9553(/!=3/9! )('! 4I9! .==(194'3C!'9=.43(/5I3@! ?94]99/! 4I9!=3449'! 1.55! .4! ]9./3/MD!4I9!
! ! !
O^!
!
.>8=4!1.55D!./>!3/!C.59!()!B0RE!./>!B&RBB!4I9!3/K95419/4!>8'.43(/*!B('9!M9/9'.==GD!
./!MI K.=89!()!$!C(''95@(/>5!4(!.!5@9C395!)('!]I3CI!4I9!=3449'!1.55!.4!]9./3/M!9Z8.495!4(!
]I.4!35!9e@9C49>!)('!./!.K9'.M9!5@9C395!]34I!4I9!5.19!.>8=4!1.55!./>!k3)!.@@=3C.?=9l!4I9!
5.19! 3/K95419/4! >8'.43(/*! ,! 5@9C395! ]34I! .! @(5343K9! ('! /9M.43K9! MI!K.=89! 4I85!
C(''95@(/>5!4(!.! 5@9C395!)('!]I3CI!4I9! =3449'!1.55!.4! ]9./3/M!35!'95@9C43K9=G!I3MI9'!('!
=(]9'!4I./!@'9>3C49>!)'(1!.!M3K9/!'9M'9553(/!1(>9=*!\(1@843/M!4I9!MI!)('!.==!_NP!5@9C395!
)('!]I3CI!4I9!>9).8=4!.>8=4!1.55D!4I9!3/K95419/4!>8'.43(/D!./>!4I9!@(5343(/!()!./G!M3K9/!
5@9C395!]34I3/! 4I9! @IG=(M9/943C!4'99! ]9'9! Q/(]/!'958=49>! 3/! 5@9C395c=9K9=!@'9>3C43(/5!
'9=.43K9=G! C(/53549/4! .C'(55! 4I9! 53e! '9M'9553(/! 194I(>5*! B(>9=5! /(4! .CC(8/43/M! )('!
@IG=(M9/943C!3/9'43.!'958=49>! 3/! MI! 95431.495!5313=.'! ?94]99/! 9.CI! (4I9'!k.==! '! }! $*O#D!
b8.>9!4954`! F3,2211!z! $*f%#D!@! z!$*^Ol*! <(]9K9'D!4I9! a4!()! 4I9!@IG=(M9/943C! =3/9.'!13e9>!
1(>9=5!k3*9*!1(>9=5!&RBB!./>!B&RBBl!'9K9.=9>!.!54'(/M!@IG=(M9/943C!53M/.=!3/!4I9!>.4.!
4I.4! 35! /(4! C.@48'9>! ?G! 4I9! .==(194'3C! '9=.43(/5I3@5*! A/>99>D! ?(4I! @IG=(M9/943C! =3/9.'!
13e9>! 1(>9=5! a449>! 4I9! >.4.! 53M/3aC./4=G! ?9449'! 4I./! 4I93'! /(/c@IG=(M9/943C!
C(8/49'@.'45!k&RBB!K5!0RE`!R3Q9=3I((>!E.43(!:954!kRE:l!z!ONND!>)!z!ND!@!~!$*$$"m!B&RBB!
K5!B0RE`!RE:!z!P#_D!>)!z!ND!@!~!$*$$"l*!
A/! 4I9! 1(>9=! B&RBBD! 4I9! C(/a>9/C9! 3/49'K.=! )('! 9Y9C4! ()! 3/K95419/4! >8'.43(/! k.5!
'9@'959/49>!?G!4I9!5C.=3/M!C(9Y3C39/4!()!4I9!.==(194'3C!'9=.43(/5I3@!?94]99/!3/K95419/4!
>8'.43(/!./>!4I9!=3449'!1.55!.4!]9./3/Ml!].5!53M/3aC./4=G!/9M.43K9!kTable 3.1l*!:I35!>(95!
/(4!31@=G!4I.4!.!=(/M9'!3/K95419/4!>8'.43(/!19./5!.!=(]9'!1.49'/.=!3/K95419/4*!A/!).C4D!
3)! 3/K95419/4! >8'.43(/! ?9C(195! 4I9! 5(=9! @'9>3C4('! 3/! 4I9! 1(>9=D! 4I9! 9Y9C4! 35! C=9.'=G!
@(5343K9!k5C.=3/M! >!z! "*P#m!\A95%!z! "*f$!c!#*$%m! [!z! _NPl*!:I9! /9M.43K9!9Y9C4! ()!4I9! =(M10!
3/K95419/4!>8'.43(/!3/!4I9!1(>9=!B&RBB!3/549.>!31@=395!4I.4!4I9!=(/M9'!1(4I9'5!3/K954!
3/!4I93'!(Y5@'3/M!4I9!=955!4I9!=3449'!1.55!.4!]9./3/M!3/C'9.595!]34I!.>8=4!1.55*!:(M94I9'D!
4I9! .>8=4! 1.55! ./>! 4I9! 3/K95419/4! >8'.43(/! 4I9'9)('9! @'9>3C49>! 1.49'/.=! 3/K95419/4!
53M/3aC./4=G!?9449'D!=9.>3/M!4(!.!1('9!19./3/M)8=!>9/(13/.4('!)('!4I9!MI!K.=89*!F('!4I959!
'9.5(/5D! ]9! CI(59! 4(! @9')('1! .==! 58?59Z89/4! 549@5! ()! (8'! ./.=G535! 853/M! 4I9! 1(>9=!
B&RBBD! ]I3CI! C.@48'95! ?(4I! 4I9! 9Y9C4! ()! 4I9! @IG=(M9/G! ./>! 4I.4! ()! 4I9! 3/K95419/4!
>8'.43(/*!
!
!
Of!
!
3.3.1.3. Sensitivity of the allometric relationship to adult mass definition
:I9!a'54! 59'395! ()!1(>9=5! ].5! ?.59>!(/! 4I9!>9).8=4!.>8=4! 1.55!k3*9*! C(1@849>! ?G!4I9!
>.4.?.59! @'(K3>9'! •! ,1/3(49! kBGI'K(=>! 94! .=*D! #$"^l! •! .5! q4I9! ?(>G! 1.55! ()! ./! .>8=4!
3/>3K3>8.=! 3/! M'.15rD! /(4! 4.Q3/M! 59e! 3/4(! .CC(8/4l*! 03/C9D! 3/! 1.11.=5D! 4I9! 1.49'/.=!
3/K95419/4! 5I(8=>! ?9! '9=.49>! 4(! 4I9! ?(>G! 1.55! ()! 4I9! )91.=9! 5@9C3aC.==GD! ]9! 859>! .!
58?594!()!4I9!>.4.!)('!]I3CI!95431.495!()!.>8=4!1.55!]9'9!.K.3=.?=9!)('!?(4I!59e95!k[!z!
"$^l!./>!'9a449>!4I9!1(>9=!B&RBB!59@.'.49=G!)('!4I9!4](!@'(e395!)('!.>8=4!1.55!kFigure
3.2l*!:I9!@'9>3C49>!1.49'/.=!3/K95419/4!194'3C5!'91.3/9>!'9=.43K9=G!5313=.'!)('!4I9!5.19!
5@9C395!3''95@9C43K9!()!4I9!4G@9!()!.>8=4!1.55!@'(eG!859>!k9e.C4!73=C(e(/c&'.44!53M/9>c
'./Q!4954D!S!z!"*ffD!@!z!$*$O_$l*!:I9!>3Y9'9/C9!?94]99/!4I9!1.49'/.=!3/K95419/4!194'3C!
C(1@849>!853/M!4I9!>9).8=4!.>8=4!1.55!./>!4I9!1.49'/.=!3/K95419/4!194'3C!C(1@849>!
853/M! 4I9! )91.=9! .>8=4! 1.55! >3Y9'9>! ?G! 1('9! 4I./! $*"! )('! (/=G! "^*#v! ()! 4I9! 5@9C395!
kFigure 3.3l*!,!>3Y9'9/C9!()!$*"!3/!1.49'/.=!3/K95419/4!.CC(8/45! )('!C.*!$*%$!0+!3/!4I9!
K.'3.43(/!3/!4I9!MI 194'3C5*!:I959!'958=45!>91(/54'.49!4I.4!4I9!CI(3C9!()!4I9!.>8=4!1.55!
@'(eG!I.>!=344=9!3/d89/C9!8@(/!3/)9'9/C95!(/!1.49'/.=!3/K95419/4!.C'(55!.!]3>9!'./M9!()!
1.11.=5*!:I35!a/>3/M! g8543a95!(8'!>9C353(/! 4(!./.=G59!>9).8=4! .>8=4!1.55!'.4I9'! 4I./!
)91.=9!.>8=4!1.55D!.5!34!.==(]5!85!4(!?9/9a4!)'(1!.!18CI!=.'M9'!5.1@=9!53S9*!
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-:&!A3#.3!D:43!43'!9'-/%14!/9%14!H/88!/09!43'!-'H/1'!/9%14!H/88!A'&'!/G/#1/D1'!5c!f!(W\!82'.#'8,)!N3'!
8./44'&!21:4!83:A8!43/4!43'! /11:H'4&#.! &'1/4#:083#2! D'4A''0! /9%14! H/88! /09! 1#44'&!H/88! /4! A'/0#0$!
/$'! 9#O'&'9! 1#441'! 9'2'09#0$! :0! 43'! /9%14! H/88! 2&:I>! .:08#9'&'9)! T/&@'&! 2:#048! /&'! 43'! &'8%14! :-!
:G'&1/22#0$!:D8'&G/4#:08! -:&! 9#O'&'04! 82'.#'8)! N3'! 4A:! &'$&'88#:0! 1#0'8! &'2&'8'04! 43'! 2&'9#.4#:08!
84'HH#0$!-&:H!/!H%14#21'!23>1:$'0'4#.!1#0'/&!H#I'9!H:9'1!5Q=_QQ,)!
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"#$%&'! L)L*! V./44'&! 21:4! :-! 43'! H/4'&0/1! #0G'84H'04! H'4&#.! 5MIMPLMM,! #0! H/HH/18! 2&'9#.4'9! %8#0$!
9'-/%14!:&!-'H/1'!/9%14!H/88!5c!f!(W\!82'.#'8,)!=&'9#.4#:08!84'H!-&:H!/!H%14#21'!23>1:$'0'4#.!1#0'/&!
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3.3.2.1 E?ect of mammalian subclasses on maternal investment
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Species
Scientific name
Subclass
Order
MIMPLMM
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Tenrec ecaudatus
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musculus
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L84I9'3.
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L84I9'3.!
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Aepyceros
melampus
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Halichoerus
grypus
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Pan troglodytes
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Lepus europaeus
L84I9'3.!
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Macropus rufus
B94.4I9'3.!
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Lasiorhinus
latifrons
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Tachyglossus
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B(/(4'9c
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?'.3/! 1.55D! kAAAl! 5(C3.=! M'(8@! 53S9D! kA-l! 4'(@I3C! =9K9=D! k-l! >394! ?'9.>4ID! ./>! k-Al! I.?34.4!
?'9.>4I*!:I9!'958=45!()!4I9!a449>!1(>9=!.'9!@'959/49>!3/!Table 4.2*!!
!
N/D1'!X)E*!FG'&G#'A! :-! 43'! :%42%4! 5&'$&'88#:0! .:'O#.#'04!/09! 2RG/1%',! -:&!43'! 8#I!2&'9#.4:&8! %8#0$!/!
H%14#21'!23>1:$'0'4#.!1#0'/&!H#I'9!H:9'1!4:!2&'9#.4!H/4'&0/1!#0G'84H'04!5c!f!(XK,)!
Predictors
Regression coefficient
p-value
B.e3181!=(/M9K34G!
c$*$$"PP!
$*NP%!
;'.3/!1.55!
f*NP9c$^!
$*$f$N!
0(C3.=!M'(8@!53S9!
$*$$"N^!
$*"O"!
:'(@I3C!=9K9=!(1/3K('G!k'9)*!I9'?3K('Gl!
c$*$$##_!
$*O^$!
:'(@I3C!=9K9=!C.'/3K('G!k'9)*!I9'?3K('Gl!
c$*"$^!
$*$NO^!
+394!?'9.>4I!
c$*$$_Nf!
$*_"%!
<.?34.4!?'9.>4I!
$*$"N%!
!$*#""!
!
W/=G!4I9!4'(@I3C! =9K9=!C.'/3K('GD!]34I!I9'?3K('G!.5! .!'9)9'9/C9D!'958=49>! 3/!.!53M/3aC./4!
9Y9C4!(/!1.49'/.=!3/K95419/4*!<(]9K9'D!.5!.!'958=4!()!4I9!=31349>!5.1@=9!53S9!)('!5(C3.=!
M'(8@!53S9!k[!z!"P"lD!4I9!4(4.=!/81?9'!()!5@9C395!3/!4I9!./.=G535!].5!(/=G!"%f*!:I9'9)('9D!
./(4I9'!B&RBB!].5!a449>D!9eC=8>3/M!4I9!@'9>3C4('!5(C3.=!M'(8@!53S9D!'958=43/M!3/!.!=.'M9'!
5.1@=9! 53S9! k[! z! %%Nl*! :I35! 1(>9=! >3>! /(4! '958=4! 3/! .! 53M/3aC./4! 9Y9C4! ()! ./G! ()! 4I9!
@'9>3C4('5*! F8'4I9'1('9D! @IG=(M9/943C! =3/9.'! 13e9>! 1(>9=5! 853/M! .! 53/M=9! @'9>3C4('!
k&RBB5l!.4!.!4319!]9'9!859>D!4(!1.e31359!4I9!5.1@=9!53S9!)('!4I9!3/>3K3>8.=!@'9>3C4('5!
k1.e3181!=(/M9K34G!k[!z!fO#lD!?'.3/!1.55!k[!z!%P_lD!5(C3.=!M'(8@!53S9!k[!z!"P"lD!4'(@I3C!
=9K9=!k[!z!_#OlD!>394!?'9.>4I!k[!z!ffOlD!./>!I.?34.4!?'9.>4I!k[!z!_"NllD!]I3CI!.==!'958=49>!
.M.3/!3/!4I9!.?59/C9!()!53M/3aC./C9*!A/!.CC('>./C9!]34I!4I9!'958=45!@'959/49>!3/!Table
4.2D!/(!C=9.'!9Y9C45!()!4I9!@'9>3C4('5! (/!1.49'/.=!3/K95419/4!.'9!K353?=9!3/!Figure 4.1D!
]I3CI! >9@3C45! 4I9! '9=.43(/5I3@! ?94]99/! 9.CI! ()! 4I9! 53e! @'9>3C4('5! ./>! 4I9! 1.49'/.=!
3/K95419/4!194'3C!.4!4I9!5C.=9!()!4I9!'.]!>.4.*!!
!
"N%!
!
!
!
"#$%&'!X)(*!FG'&G#'A!:-!8#I!2&'9#.4:&8!21:44'9!/$/#084!H/4'&0/1!#0G'84H'04)!N3'!21:48!83:A!H/4'&0/1!
#0G'84H'04!G'&8%8!5+,!H/I#H%H!1:0$'G#4>!5c!f!KME,6!57,!D&/#0!H/88!5c!f!XaJ,6!5;,!8:.#/1!$&:%2!8#Z'!5c!f!
(a(,6!5T,!4&:23#.!1'G'1!5c!f!JEM,6!5<,!9#'4!D&'/943!5c!f!KKM,6!/09!5",!3/D#4/4!D&'/943!5c!f!J(L,)!N3'!1/&$'!
8>HD:18!#0!43'!.'04&'!:-!43'!G'&4#./1!1#0'8!#0!21:48!T6!<6!/09!"!&'2&'8'04!43'!H'9#/0!G/1%'8)!N3'!8H/11'&!
8>HD:18!1:./4'9!&#$34!-&:H!43'!G'&4#./1!1#0'8!#09#./4'!#09#G#9%/1!82'.#'86!A#43!>RG/1%'8!.:&&'82:09#0$!
4:!43'!2&'9#.4'9!H/4'&0/1!#0G'84H'04)!N3'8'!8>HD:18!/&'!82&'/9!:%4!/1:0$!43'!IR/I#8!-:&!43'!2%&2:8'!
:-!G#8%/1#8/4#:0)
!
:I9!.?59/C9!()!53M/3aC./4!9Y9C45!()!4I9!@'9>3C4('5!(/!1.49'/.=!3/K95419/4!3/!4I9!B&RBB!
./>! &RBB5! 31@=395! 4I.4! 1(54! ()! 4I9! >3Y9'9/C95! 3/! 1.49'/.=! 3/K95419/4! ./>! 4I9!
! ! !
"N^!
!
@'9>3C4('5!.'9!5I.@9>!?G! @IG=(M9/G!./>!/(4!?G! >3Y9'9/C95!3/!4I9!@'9>3C4('5!@9'!59*!:(!
4954!4I9!9Y9C4!)'(1!4I9!@'9>3C4('5!(/!.!=(]9'!4.e(/(13C!=9K9=D!3*9*!4I9!4.e(/(13C!('>9'D!
)(==(]3/M!4I9!'.43(/.=9!4I.4!.!>9C'9.59!3/!4.e(/(13C!K.'394G!13MI4!'9K9.=!4I9!9Y9C4!()!4I9!
@'9>3C4('5! (/! 1.49'/.=! 3/K95419/4D! ]9! 59=9C49>! .! 58?594! ()! E(>9/43.! k[! z! #f#D!
'9@'959/43/M!Nfv!()! 4I9!5@9C395!3/!4I9!>.4.594lD!4I9!1(54! 5@9C3(59!1.11.=3./!('>9'D!
9/C(1@.553/M!NOv!()!.==!1.11.=3./!5@9C395!k;8'M3/!94!.=*D!#$"Pl*!:I9!B&RBB!853/M!4I9!
E(>9/43. 58?594!'9K9.=9>!/(!53M/3aC./4!31@.C4!()!./G!()!4I9!53e!@'9>3C4('5!(/!1.49'/.=!
3/K95419/4D!I(]9K9'D!5(19!&RBB5!>3>!kTable 4.3l*!
!
N/D1'!X)L*!FG'&G#'A!:-!43'!:%42%4!5&'$&'88#:0!.:'O#.#'046!2RG/1%'6!/09!c,!-:&!43'!8#I!2&'9#.4:&8!%8#0$!
23>1:$'0'4#.!1#0'/&!H#I'9!H:9'18!5=_QQ8,!4:!2&'9#.4!H/4'&0/1!#0G'84H'04!:0!/!8%D8'4!:-!S:9'04#/!5c!
f!EKE,)!
Predictors
Regression coefficient
p-value
N
B.e3181!=(/M9K34G!
c$*$"$%!
$*$$_P_!
#NP!
;'.3/!1.55!
$*$$#$"!
$*%#O!
"N_!
0(C3.=!M'(8@!53S9!
c$*$$$%f$!
$*Of_!
%"!
:'(@I3C!=9K9=!(1/3K('G!k'9)*!I9'?3K('Gl!
$*$N^O!
$*$_NN!
#f"!
!
:'(@I3C!=9K9=!C.'/3K('G!k'9)*!I9'?3K('Gl!
$*"$^!
$*$P^N!
+394!?'9.>4I!
c$*$N_%!
$*$N#"!
##P!
<.?34.4!?'9.>4I!
$*$$$^^$!
$*ON%!
#^_!
!
:I9!&RBB5!'9K9.=9>!.!53M/3aC./4D!I(]9K9'!51.==!9Y9C4!()!1.e3181!=(/M9K34G!./>!>394!
?'9.>4I!(/!1.49'/.=!3/K95419/4*!B.e3181!=(/M9K34G!./>!>394!?'9.>4I!>9C'9.59>!]34I!./!
3/C'9.59!3/!1.49'/.=!3/K95419/4!kFigure 4.2l*!
\(/53>9'3/M!4I9!4'(@I3C!=9K9=!.5!.!@'9>3C4('!()!4I9!'953>8.=!K.'3./C9!3/!4I9!kBl&RBB!1(>9=!
31@'(K9>! 4I9! M((>/955! ()! a4! ()! 4I9! 1(>9=! ./>! '9K9.=9>! 4I.4! 4I9! '953>8.=! K.'3.43(/! 3/!
1.49'/.=! 3/K95419/4! ?94]99/! 5@9C395! ].5! I3MI9'! )('! C.'/3K('95! k'953>8.=! K.'3./C9! z!
$*$#N_l! 4I./! 34! ].5! )('! 934I9'! I9'?3K('95! k'953>8.=! K.'3./C9! z! $*$"N$l! ('! (1/3K('95!
k'953>8.=!K.'3./C9!z!$*$$O$fl*!
!
"Nf!
!
!
!
"#$%&'!X)E*!FG'&G#'A!:-!8#I!2&'9#.4:&8!21:44'9!/$/#084!H/4'&0/1!#0G'84H'04!%8#0$!/!8%D8'4!:-!S:9'04#/!
5c! f! EKE,)! N3'! 21:48! 83:A! H/4'&0/1! #0G'84H'04!G'&8%8! 5+,! H/I#H%H! 1:0$'G#4>6! 57,! D&/#0! H/886! 5;,!
8:.#/1!$&:%2!8#Z'6!5T,!4&:23#.!1'G'16!5<,!9#'4!D&'/9436!/09!5",!3/D#4/4!D&'/943)!N3'!1/&$'!8>HD:18!#0!43'!
.'04&'! :-! 43'! G'&4#./1! 1#0'8! #0! 21:48! T6! <6! /09! "! &'2&'8'04! 43'! H'9#/0! G/1%'8)! N3'! 8H/11'&! 8>HD:18!
1:./4'9!&#$34! -&:H! 43'! G'&4#./1! 1#0'8! #09#./4'! #09#G#9%/1!82'.#'86!A#43!>RG/1%'8!.:&&'82:09#0$!4:!43'!
2&'9#.4'9! H/4'&0/1! #0G'84H'04)! N3'8'! 8>HD:18! /&'! 82&'/9! :%4! /1:0$! 43'! IR/I#8! -:&! 43'! 2%&2:8'! :-!
G#8%/1#8/4#:0)
!
="=#$%)47))%1-#
:I9!'9M'9553(/!./.=G595! C(/>8C49>!3/!4I35!548>G! .319>!4(!3/K9543M.49!4I9!'9=.43(/5I3@5!
?94]99/!1.49'/.=!3/K95419/4! ./>! 1.e3181! =(/M9K34GD!?'.3/! 1.55D! 5(C3.=! M'(8@!53S9D!
4'(@I3C! =9K9=D! >394! ?'9.>4ID! ./>! I.?34.4! ?'9.>4I! .C'(55! >3Y9'9/4! 4.e.! 4(! 9/I./C9! (8'!
C(1@'9I9/53(/!()!3/K95419/4!54'.49M395!./>!4I93'!9K(=843(/.'G!>9K9=(@19/4!4I'(8MI(84!
! ! !
"N_!
!
1.11.=3./!5@9C395*!H3K9/! 4I9!>(C819/49>! 9Y9C4!()!4I959! 53e!@'9>3C4('5!(/! ./.4(1G!
k+8/?.'D!#$$Om! +8Z89c\(''9.!94!.=*D! #$#"m!GonzálezLagos et al., 2010; 0134I! 94!.=*D!
#$"_m!7.=>1./!94!.=*D!#$#NlD!@IG53(=(MG!k\=844(/c;'(CQD!#$#"m!<()1./D!"OPNm!<(@@9!94!
.=*D!#$#"lD!./>!?9I.K3(8'!kHämäläinen et al, 2022; Henke-von Der Malsburg and Fichtel,
2018; Silk, 2007; Zhu et al., 2023)D!]9!IG@(4I95359>!4I959!@'9>3C4('5!4(!I.K9!./!9Y9C4!
(/! 1.49'/.=! 3/K95419/4*! A/! C(/4'.>3C43(/! 4(! (8'! IG@(4I9595D! 4I9! 54.43543C.=! ./.=G595!
1.e313S3/M! 5.1@=9! 53S9! >3>! /(4! '9K9.=! .! 53M/3aC./4! '9=.43(/5I3@! ?94]99/! 1.49'/.=!
3/K95419/4!./>!./G!()!4I9!53e!@'9>3C4('5!49549>!(/!4I9!=9K9=!()!4I9!4.e(/(13C!C=.55D!3*9*!
4I9! B.11.=3.D! I(]9K9'! 5313=.'! ./.=G595! (/! 4I9! 5C.=9! ()! 4I9! ('>9'D! 853/M! .! 58?594! ()!
5@9C395!)'(1!4I9!?3MM954!1.11.=3./!('>9'D!3*9*!4I9!E(>9/43.D!'9K9.=9>!.!53M/3aC./4!9Y9C4!
()! 1.e3181! =(/M9K34G! ./>! >394! ?'9.>4I! (/! 1.49'/.=! 3/K95419/4*! :I9! .?59/C9! ()!
/(43C9.?=9!C(''9=.43(/5!(/!4I9!=9K9=!()!4I9!4.e(/(13C!C=.55D!58MM954!4I.4!>3Y9'9/C95!3/!
1.49'/.=!3/K95419/4!./>!4I9!@'9>3C4('5!.'9!1(54=G!C.@48'9>!?G!).C4('5!'9d9C49>!?G!4I9!
@IG=(M9/G*!!
B.e3181! =(/M9K34G! 35! 439>! 4(! 4I9! @.C9! ()! =3)9! kj8./! 94! .=*D! #$#Nl*! :I9! 5I('4954! =3K9>!
1.11.=5D!3*9*!C9'4.3/!5@9C395!()!5I'9]!./>!13C9D!I.K9!.!=3)95@./!()!.?(84!.!G9.'D!>8'3/M!
]I3CI!4I9G!.'9!?('/D!?9C(19!59e8.==G!1.48'9D!M3K9!?3'4ID!'.359!(Y5@'3/MD!./>!>39*!:I35!).54!
@.C9!()!=3)9!35!CI.'.C49'359>!?G!.!I9.'4!'.49!()!P$$c"$$$!?9.45!@9'!13/849!k0CI13>4!94!.=*D!
#$"fl*!W/!4I9!(@@(5349!9/>! ()!4I9!5@9C4'81D!4I9!?(]I9.>! ]I.=9!kBalaena mysticetusl!
I.5!.!1.e3181!=(/M9K34G!()!#""!G9.'5!k0931!94!.=*D!#$"%l*!:I9!I9.'4!'.49!()!C9'4.3/!5@9C395!
()!]I.=9!35!Q/(]/!4(!?9!.'(8/>!"^!?9.45!@9'!13/849D!'9@'959/43/M!.!K9'G!5=(]!@.C9!()!=3)9!
kH(=>?(M9/!94!.=*D!#$"Ol*!+95@349!4I9!'./M9!()!4I35!=(/M9K34G!./>!@.C9!()!=3)9!5@9C4'81D!/(!
3/d89/C9!(/!1.49'/.=!3/K95419/4!].5!>949C49>!.4!4I9!=9K9=!()!4I9!4.e(/(13C!C=.55*!W/!
4I9!C(/4'.'GD!3/!4I9!E(>9/43.!1.e3181!=(/M9K34G!I.>!.!53M/3aC./4D!?84!51.==D!31@.C4!(/!
1.49'/.=!3/K95419/4*!:I9!>3'9C43(/!()!4I9!9Y9C4!(/!?(4I!4.e(/(13C!=9K9=5!].5!3/!=3/9!]34I!
4I.4!@'9>3C49>!?G!4I9!>35@(5.?=9!5(1.!4I9('G!()!.M93/M!k<.119'5!94!.=*D!#$"Nl*!:I35!'958=4!
5I(8=>!I(]9K9'!?9! 3/49'@'949>!]34I!C.843(/! M3K9/!4I9!51.==!5=(@9!k95431.49!z!c$*$"$%l!
./>! 4I9! ./43cC(/59'K.43K9! /.48'9! ()! 49545! 853/M! 1.49'/.=! 3/K95419/4! 194'3C5! .5! .!
'95@(/59! K.'3.?=9! k3*9*D! ).=59! @(5343K9! '.495! I3MI9'! 4I./! /(13/.=! =9K9=5m! 599! 194I(>5l*!
H3K9/! 4I.4! '9@'(>8C43K9! 54'.49M395! ?94]99/! 4I9! (@@(5349! 9/>5! ()! 4I35! @.C9! ()! =3)9!
5@9C4'81!.'9!C(1@=949=G!>3Y9'9/4D!4I9!'9M8=.4('G!19CI./3515!)('!3/K95419/4!.'9!=3Q9=G!
!
"NP!
!
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'9=9K./C9!()! 4I9! @'9>3C4('! I.5!4(! ?9! C(/53>9'9>*!,=4I(8MI! 1.e3181! =(/M9K34G! M3K95!.!
M9/9'.=!3>9.!()!4I9!=3)95@./!()!.!5@9C395D!(/=G!.!)9]!3/>3K3>8.=5!'9.CI!1.e3181!=(/M9K34G*!
:I9!1.e3181!=(/M9K34G!.=5(!@'959/45!.!I3MI!5.1@=3/M!K.'3./C9!]I3CI!1.Q95!95431.495!
8/'9=3.?=9! 8/=955! 5.1@=9! 53S95! .'9! =.'M9*! :I9'9)('9D! 4I9! O$v! =(/M9K34G! 194'3C! ('! 4I9!
.K9'.M9! =(/M9K34G! ](8=>! ?9! .! ?9449'! @'(eG! )('! =3)95@./! kE(/M94! ./>! H.3==.'>D! #$#$l*!
J/)('48/.49=GD!(/=G!1.e3181!=(/M9K34G!35!]3>9=G!.K.3=.?=9!3/!>.4.5945*!!
,5!.!59C(/>!@'9>3C4('!)('!1.49'/.=!3/K95419/4D!]9!49549>!?'.3/!1.55*!&'9K3(85!548>395!
@'(@(59>!>3Y9'9/4!@'(e395!)('! 3/49==3M9/C9D!58CI!.5!?'.3/!1.55D!?'.3/! ?(>G!1.55!'.43(D!
./>! 4I9! 9/C9@I.=3S.43(/! Z8(439/4! k,M83''9! 94! .=*D! #$"_m! 69'35(/D! "O__l*! ,=4I(8MI! 58CI!
194'3C5!C(''9=.49!]34I!3/49==3M9/C9D!4I9!.CC8'.CG!()!4I959!194I(>5!35!543==!?93/M!>9?.49>!
kCairό, 2011)*!E9C9/4!'959.'CI!@(3/45!(84!4I.4!4I9!M'(]4I!()!?'.3/!53S9!./>!?(>G!53S9!I.5!
/(4!C(/53549/4=G!)(==(]9>!@.'.==9=!@.4I5!3/!9K(=843(/!k01.9'5!94!.=*D!#$#"l*!:I9'9)('9D!]9!
.CC(8/49>!)('!.==(194'G!./>! @IG=(M9/943C!3/9'43.!?G!853/M! .!@IG=(M9/943C!13e9>!1(>9=!
]34I!?'.3/!1.55!.5!@'(eG!)('!3/49==3M9/C9*!\(/4'.'G!4(!(8'!548>GD!@'9K3(85!'959.'CI!.C'(55!
K9'49?'.495!5I(]9>!4I.4!3/C'9.59>!'9=.43K9!?'.3/!1.55!35!.55(C3.49>!]34I!.!53M/3aC./4=G!
I3MI9'!1.49'/.=! 3/K95419/4! kB8==!94! .=*D! #$#$l*!,=4I(8MI! 4I9! >3'9C43(/!()! 4I9! 9Y9C4!]9!
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Chapter 5
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Review: The role of prolactin in balancing
the parent-offspring conflict!
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Chapter 6
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Elephant gestation: insights into idiopathic
abortions and stillbirth!
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Bibliography
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Conference abstract
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Do what 99%
of the people
don’t do
!
Evolutionary strategies behind
maternal investment in
mammalian reproduction
Tim E.R.G. Huijsmans
Evolutionary strategies behind maternal investment in mammalian reproduction I Tim E.R.G. Huijsmans
Do what 99%
of the people
don’t do
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