bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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*21/>)-*)29(C/$0-& *-)$29& 2A& 2.-& 9"+& 3"1& "11(B-9& V"#/%2(7%&
/%-& "11(B*/$2.& 3"1& )25(1/& /2& $.C)-"1$.7& /%-& Gh=?& ".9&
1(**2)/-9&5+&251-)0"/$2.1XF&".9&"7-&/2&)-*)29(C/$0-&B"/()$/+&
3"1& "11(B-9& /2& 5-& .-7#$7$5#-& $.& /%-& C"1-& 2A& /%-& 1-"& ()C%$.1&
5-C"(1-& 2A& /%-$)& )-#"/$0-#+& #2.7& #$0-1& `fMc4& e2)& /%-& 1%")U& ".9&
"##$7"/2)F&/%-&"7-&2A&)-*)29(C/$0-&1-.-1C-.C-&3"1&"11(B-9&/2&
5-& /%-& "0-)"7-& #$A-1*".& 2A& /%-& 2)7".$1BF& C2.1$1/-./& 3$/%&!
*)-0$2(1&/%-2)-/$C"#&".9&-B*$)$C"#&32)U&`ffc4&@.&"99$/$2.F&"7-&
"/& )-*)29(C/$0-& B"/()$/+& 3"1& (1-9& A)2B& -$/%-)& 2)& 52/%&
7-.9-)1& 3%-.& C2.1$9-)$.7& "##& B"BB"#1n".9& 1-*")"/-9& 5+&
7-.9-)&3%-.&C2.1$9-)$.7&B"BB"#1&V9(-&/2&9"/"&#$B$/"/$2.1X4&
?$.C-&.2/&-6*#$C$/F&A2)&1-"&()C%$.1F&#$//-)&1$T-&".9&5$)/%&1$T-&3"1&
-1/$B"/-9&5+&/%-&.(B5-)&".9&1$T-&2A&VA-B"#-X&7"B-/-14&?$.C-&
()C%$.1& ")-& -6/-)."#& A-)/$#$T-)1F& /%2(7%F& 3-& "11(B-& /%"/& /%$1&
)-#-"1-&$1&/%-&B21/&1/)-11A(#&*")/&2A&Y5$)/%$.7Z&".9&3$##&1(AA$C-4&
R$/%$.&B"BB"#1F&.2&2.-&A"C/2)&"CC2(./-9&A2)&/%-&B"j2)$/+&2A&
*21/>)-*)29(C/$0-& #$A-1*".F& *-)%"*1& $.9$C"/$.7& /%"/& 1-0-)"#&
9$AA-)-./&*")"B-/-)1&C2./)$5(/-&/2&/%-&9-/-)B$."/$2.&2A&*21/>
)-*)29(C/$0-& #$A-1*".& V?5&&'(%(,/2'& @27'(" AX4& @.& "99$/$2.F&
%(B".1& V9"/"& A)2B& /%-& %(./-)>7"/%-)-)>#$U-& W"9T"& ".9&
op(.7X&3-)-&.2/&2(/#$-)1&C2B*")-9&/2&2/%-)&B"BB"#14&
(
0#"#(%,"%1"
&
D"/"/"5#-&a&V!"BB"#1X
D"/"/"5#-&b&VO##&O.$B"#1X&&
&
&
;2&"11-11&3%-/%-)&/%-)-&$1&"&C2.1-)0-9&9-/-)B$."./&2A&*21/>
)-*)29(C/$0-& #$A-& 1*".& VGh=?X& "C)211& 1*-C$-1F& 3-& 7"/%-)-9&
$.A2)B"/$2.&"52(/&#$A-&%$1/2)+&A-"/()-1&2.&"&0")$-/+&2A&".$B"#&
/"6"& Vgf& 1*-C$-1X& 52/%& 3$/%$.& ".9& 2(/1$9-& 5"22".*"F&
$.C#(9$.7& 1-"& ()C%$.1F& 1"#B2.F& 3&%,%4'*."- 2#."$%/",+#&F& ".9&
1*-C$-1& 2A& )-*/$#-1& ".9& 5$)91& V?5&&'(%(,/2'" @27'(" AB"
?5&&'(%(,/2'" @27'(" CXF& A2)& 3%$C%& 3-& C2(#9& 25/"$.&
$.A2)B"/$2.& 2.& )-*)29(C/$0-& 1-.-1C-.C-F& -474F& #$A-& 1*".F& "7-&
"/&)-*)29(C/$0-&B"/()$/+F&"9(#/&1$T-F&*)27-.+&1$T-&"/&5$)/%F&".9&
"0-)"7-& #$//-)& 1$T-& V1--& ?(**#-B-./"#& $.A2)B"/$2.& A2)& "##&
"0"$#"5#-&0")$"5#-1X4&R-&/%-.&C2B*")-9&/%-1-&A-"/()-1&/2&/%-&
*21/>)-*)29(C/$0-&#$A-&1*".&)"/$2&VD$-5+("AEP&
PRLS =
maximum lifespan " age at last birth
&
maximum lifespan " age of female reproductive maturity
W23-0-)F& "& 1()*)$1$.7#+& 1$B*#-& B-/)$CF& /%-& )"/$2& 2A& 2AA1*)$.7&
1$T-&/2&B2/%-)&1$T-F&2)&Y]AA1*)$.7&)"/$2FZ&C2))-#"/-9&3-##&3$/%&
Gh=?& "C)211& /%-& #")7-)& 1-/& 2A& 1*-C$-14& ;%-& B-"1()-& 3"1&
C2B*(/-9&$.&/32&3"+1&9-*-.9$.7&2.&9"/"&"0"$#"5#-P&
Offspring Ratio =
offspring weight at birth
offspring length 3 &
OR
adult weight
adult length 3
;%-& (."9j(1/-9& 2AA1*)$.7& )"/$2& C2(#9& -6*#"$.& "**)26$B"/-#+&
[M\&2A&/%-&0")$".C-&2A&*21/>)-*)29(C/$0-&#$A-1*".&Vhq&r&K4[[ai&
*& r& K4KKKa[NX& VD$-5+(" AFX4& ;%"/& $1F& #")7-)& 2AA1*)$.7& 3$/%&
)-1*-C/& /2& /%-& B2/%-)& $1& "112C$"/-9& 3$/%& #2.7-)& *21/>
)-*)29(C/$0-&*-)$29&2A&/%-&B2/%-)4&VI2/-&/%"/&3%$#-&#$//-)&1$T-&
%"1&".&$.0-)1-&".9&%$7%#+&C2))-#"/-9&Vhq&r&K4LfgX&)-#"/$2.1%$*&
3$/%&Gh=?&VD$-5+("AGXF&$/&$1&"#12&C2))-#"/-9&3$/%&".9&$.0-)1-#+&
)-#"/-9& /2& 2AA1*)$.7& )"/$2& Vhq& r& K4[LaXF& ".9& /%(1& C"..2/& 5-&
9$1/$.7($1%-9&A)2B&2AA1*)$.7&)"/$2&VD$-5+("AHX4&
@.&2)9-)&/2&"//-B*/&/2&1-*")"/-&/%-&-AA-C/1&2A&*%+#27-.+&A)2B&
/"62.2B$C& "9"*/"/$2.1F& B-/%291& 2(/#$.-9& *)-0$2(1#+& 3-)-&
(/$#$T-9& `f[FfLc4& ?*-C$-1>#-0-#& 9"/"& "B2.7& B"BB"#1& 3-)-&
"0-)"7-9&/2&/%-&A"B$#+&#-0-#F&)-C"*$/(#"/$.7&1$B$#")&)-1(#/1&".9&
1(77-1/$.7&/%-&*)-1-.C-&2A&"9"*/"/$2.4&8)"*%$.7&/%-&)-1$9("#1&
2A& 1-*")"/-& 2AA1*)$.7& ".9& "9(#/& 3-$7%/& )-7)-11$2.1& "7"$.1/&
Gh=?& 1-*")"/-9& /%-& "0"$#"5#-& *#"C-./"#& B"BB"#& A)2B& /%-&
1$.7#-& B")1(*$"#& 9"/"& *2$./4& O1& A2)& /%-& .2.>B"BB"#1F& $.&
3
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
!"#$%"&'&!()*%+&,&*)-*)$./&0-)1$2.&,33345$2)6$042)7&
2)9-)& /2& C"*/()-& B2)-& 9"/"& *2$./1F& 3-& B-"1()-9& 2AA1*)$.7&
)"/$2& /%)2(7%& /32& B-/%291& V2(/#$.-9& $.&!-/%291X4& W23-0-)F&
/%-& .2.>B"BB"#1& ")-& 1-#-C/-9& A)2B& "& 3$9-& 0")$-/+& 2A& /"6"F&
".9& -0-.& 3%-.& 12B-& 2A& /%-& C#21-#+>)-#"/-9& 1*-C$-1& ")-&
"0-)"7-9& V-474F& 1-"& ()C%$.1& ".9& 1"#B2.XF& /%-& )-1(#/1& 92& .2/&
C%".7-F& #-11-.$.7& /%-& C2.C-).& "52(/& /%-1-& 9$AA-)-.C-1&
-B-)7$.7& A)2B& *%+#27-.+& V"#/%2(7%& B-/%291& /2& 1-*")"/-&
*%+#27-.+&".9&"9"*/"/$2.&")-&(."0"$#"5#-&$.&/%$1&C"1-X4&
A
Development
FERTILIZATION
Birth
Proportion
Reproductive
Proportion
Reproductive
or Alive or Alive
]()&C2B*")$12.&2A&2AA1*)$.7&)"/$2&/2&Gh=?&"#12&)2(7%#+&7)2(*1&
2)7".$1B1&
5+& )-*)29(C/$0-& 1/)"/-7+4& :#(1/-)-9& $.& /%-& #23&
A
2AA1*)$.7& )"/$2s#23& Gh=?& ")-"& 2A& /%-& 7)"*%& ")-& )-9& 1-"&
()C%$.1F& *()*#-& 1-"& ()C%$.1F& ".9& 1"#B2.& 1*-C$-1n"##&
2)7".$1B1& /%"/& )-#-"1-& (.A-)/$#$T-9& 7"B-/-1& $./2& /%-&
-.0$)2.B-./& VD$-5+(" AFX4& ;%-& OA)$C".& ?U$.UF& 3&%,%4'*."2#."$%/",+#&F& ?C%22#& ?%")UF& ".9& O##$7"/2)& A2)B& ".2/%-)&
7)2(*F&*)29(C$.7&A-)/$#$T-9&-B5)+21&/%"/&")-&)-#-"1-9&$./2&/%-&
-.0$)2.B-./&3$/%&.2&*")-./"#&C")-&VD$-5+("AFX4&e$."##+F&5$)91&
".9&B"BB"#1n".$B"#1&/%"/&C")-&A2)&/%-$)&+2(.7n"**-")&/2&
A2)B& ".2/%-)& C#(1/-)& "/& /%-& %$7%& 2AA1*)$.7& )"/$2s%$7%& Gh=?&
)-7$2.&2A&/%-&7)"*%&VD$-5+("AFE4&
B
Reproductive Period
Adulthood
Reproductive Period
Fertilization
Development
Adulthood
Gestation Weaning Reproductive
Maturity
FERTILIZATION
Birth
Fertilization
Gestation Weaning Reproductive
MaturityDocumented Lifespan
Maximum
Reproductive
Senescence
Reproductive
Senescence
PRLS
PRLS
DEATH
DEATH
Maximum Documented Lifespan
C
C
D$-5+("AI"J)#/.+(&+)354/$6("'$*("#&2,"4)++('2/(#":$/0")**#&+$,-"+2/$)"
4
B
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
!"#$%"&'&!()*%+&,&*)-*)$./&0-)1$2.&,33345$2)6$042)7&
R%-.& #22U$.7& "/& /%-& 9$0-)1$/+& 2A& ".$B"#1F& $/& "**-")1& /%"/&
/%21-&3$/%&/%-&#")7-1/&2AA1*)$.7&)"/$21&%"0-&/%-.7-1/&*21/>
)-*)29(C/$0-&#$0-1F&".9&0$C-&0-)1"4&;%$1&)-1(#/&$1&C2(./-)&/2&/%-&
.2/$2.& /%"/& #")7-)& 2AA1*)$.7& 9-*#-/-& )-12()C-1F& )-1(#/$.7& $.& "&
1%2)/-)& *21/>)-*)29(C/$0-& #$A-& 1*".4& @.9--9F& 2()& )-1(#/1&
1(77-1/& /%"/& /%-)-& $1& .2/& "& 9$)-C/& Y/)"9-2AAZ& 5-/3--.&
)-*)29(C/$0-&".9&*21/>)-*)29(C/$0-&#$A-&`Nc4&@.1/-"9F&3-&*21$/&
/%"/&/%-&2AA1*)$.7&)"/$2&B"+&5-&"&*)26+&A2)&1/)-11&2A&C%$#95$)/%&
2)& *)27-.+& *)29(C/$2.4& ;%-)-& $1& #$U-#+& "& *2$./& "/& 3%$C%& ".&
2)7".$1B& C"..2/& 9-02/-& "9-^("/-& -.-)7+& 2)& ^("#$/+&
B"$./-.".C-& /2& )-*)29(C/$2.& 5(/& 1/$##& %"1& /%-& .-C-11")+&
1/)-.7/%& ".9& *%+1$C"#& $./-7)$/+& /2& #$0-4& O#/%2(7%& $.& /%-&
"51-.C-& 2A& *")-./"#& C")-F& .-$/%-)& *21$/$0-& .2)& .-7"/$0-&
1-#-C/$2.&"C/1&2.&Gh=?F&/%-1-&C2))-#"/$2.1&1(77-1/&"&)-"12.&A2)&
/%-& *")/$C(#")& #-.7/%& 2A& Gh=?& `Mc4& @.& -11-.C-F& Gh=?& $.& B".+&
1*-C$-1&$1&1$B*#+&".&(.1-#-C/-9&)-1$9("#&2A&#$A-&`aMFfgF[KcF&5(/&
/%-& )-"12.& A2)& /%$1& )-1$9("#& %"1& .2/& 5--.& /-1/-9& *)-0$2(1#+4&
R-&1(77-1/F&5"1-9&2.&/%-&C2))-#"/$2.1&A2(.9&%-)-F&/%"/&B2)-&
1/)-11A(#&2)&*%+1$C"##+&9-B".9$.7&A2)B1&2A&)-*)29(C/$2.&B"+&
)-^($)-& 7)-"/-)& 1/)-.7/%& ".9& $./-7)$/+& $.& /%-& 12B"& /2&
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$./-7)$/+&$.&/%-&*21/>)-*)29(C/$0-&*-)$29&".9&C2))-1*2.9$.7#+&
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5-& A2(.9& 3%-.& A-B"#-1& ")-& *(1%-9& /2& )-*)29(C-& 5-+2.9& "&
/+*$C"#&/$B-P&$.&%(B".1F&/%-&B21/&C2BB2.&C"(1-&2A&9-"/%&2A&
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5
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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time (days of adulthood)
time (days of adulthood)
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matricide fraction
fraction reproductive
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time (days of adulthood)
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% reproductive/alive
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time (days of adulthood)
time (days of adulthood)
time (days of adulthood)
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6
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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7
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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11
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
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bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
bioRxiv preprint first posted online Apr. 15, 2016; doi: http://dx.doi.org/10.1101/048835. The copyright holder for this preprint (which was not
peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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