Beyond  Life+me  Reproduc+ve  Success  The  dynamics  of  posthumous  reproduc+on  in  Trinidad  guppies  

Andrés  López-­‐Sepulcre  

Collaborators  

General Protocol

•! 3 scales sampled, stored

dry in tubes

•! DNA isolation (enough for

~50 PCRs)

•! Genotype at ~12

microsatellite loci (more available)

•! Parentage assignment

via CERVUS, PAPA, PASOS…

Swanne  Gordon  Univ.  Jyväskylä  Finland  

Paul  Bentzen  &  Ian  Paterson  Dalhousie  Univ.    

Canada  

David  Reznick  Univ.  California,  Riverside  

USA  

Sperm  Storage  in  Animals  

•  Common  in  many  species  •  Can  lead  to  posthumous  reproduc+on  •  Its  dynamics  in  nature  largely  unknown  •  Evolu+onary  significance?  

Trinidadian  guppy,  Poecilia  re*culata  

•  Two  dis+nct  phenotypes  associated  to  preda+on  and  resources  

•  Differences  in  mul+ple  traits  and  ecological  effects  •  Several  independent  evolu+onary  events    •  Rapid  evolu+on  aSer  transloca+on  

Reznick et al. 1990. Nature

Experimental  evolu+on  in  the  wild  

Guppy  Mark  Recapture  

•  All  individuals  >14mm  individually  marked  •  Followed  monthly  •  Gene+c  samples  for  pedigree  reconstruc+on  

Closed canopy

h2 = 0.30

Capture probabilities

Male  Size  at  Maturity  

Evolu+on  of  male  size  

•  Common  garden  

Source  Popula+on  

Derived  

Survival  

Standard Length (mm)

Sur

viva

l pro

babi

lity

DEADALIVE

DEADALIVE

Reproduc+on  •  Larger  individuals  reproduce  less  

 …  but  we  had  only  counted  the  live  ones  •  Guppy  females  store  male  sperm  

•  Hot  to  count  in  the  dead?        

Prop

or+o

n  of  re

cruits  sired  

Standard  Length  (mm)  

Juvenile (J)

Mature not reproducing

(M)

Mature reproducing

(R)

Dead reproductive

(S)

!J1" µ( )

!M1" #( )

!R

!S

! Jµ1"#

()

!J µ"

!M"

!1"#

M

()$

1!" R

()#

Conception

3 m

onth

s

!R"R#R

!S"S#S

State  Space  Models  

•  Define  model  that  defines  unobserved  biology  

Model  includes  transi+on    between  states  

Mature  Not  Reproduc+ve  

Mature  Reproduc+ve  

Dead  Reproduc+ve  Juvenile  

State  Space  Models  

•  Overlay  an  observa+on  model  •  Data  is  a  result  of  the  observa+on  model  condi+onal  on  the  biology  model  

•  We  can  es+mate  ‘invisible’  biological  parameters  

Alive   Alive  

Not  Rep.  Observed  

Not  Rep  Not  Obs  

Alive   Alive   Alive   Zombie   Zombie   Dead  

Not  Rep.  Not  Obs.  

Reproduced  Observed  

Reproduded  Not  Obs  

Not  Rep  Not  Obs  

Not  Rep.  Not  Obs.  

Results  

●

● ●

●

● ●●

●

01

23

4Months reproductive

1 2 3 4 5 6 7 8 9 10

Live males(a)

●

●

●

● ●●

●●

●

●

● ●01

23

4

Months dead1 2 3 4 5 6 7 8 9 10

Dead males(b)

Indi

vidu

al c

ontri

butio

n to

pop

ulat

ion

recr

uitm

ent (

%)

● ● ● ● ●● ●

●

●●

01

23

4

Months reproductive1 2 3 4 5 6 7 8 9 10

Females(c)

Individu

al  con

tribu+

on  to

 recruitm

ent  

…whatever  

Results  

2 4 6 8 10

050

100

150

Months since introduction

Num

ber o

f mal

es

Dead reproductives (S)Reproductive adults (R)Non−reproductive adults (M)Juveniles (J)

●

●●

●

●

●

●

●

●

●

2 4 6 8 100.

000.

050.

100.

150.

200.

250.

30

Months since introduction

Prop

ortio

n of

new

recr

uits

sired

by

dead

mal

es

Does  size  macer?  

DEADALIVE

DEADALIVE

Prop

or+o

n  of  re

cruits  sired  

Standard  Length  (mm)  

Prop

or+o

n  of  re

cruits  sired  

Standard  Length  (mm)  

Thoughts  

•  How  does  storage  affect  natural  selec+on?  – Maintenance  of  diversity  –  ‘Bet-­‐hedging’  

Thoughts  

•  How  does  storage  affect  sexual  selec+on?  – Opera+onal  Sex  Ra+o  – Male-­‐male  compe++on  

Carotenoid Melanistic

HP

Source

HP

Source Canopy Open

Canopy Open

Pro

po

rtio

n s

ho

win

g c

olo

r

Thanks!        

Sensi+vity  &  Elas+city  Analysis  

Focal  Experiment    • Individual  based  data  • Monthly  • Phenotype  (pictures)  • Pedigree  and  gene+c  samples  • >15,000  ind,  >10  genera+ons  • ~90%  recapture  rates  • Ecosystem  data  • Physical  variables  • Bimonthly  es+mate  of  compartments  • Annual  15N  drip    

Mesocosms  

• Ecological  effects  

Laboratory  

• Common  garden  phenotyping  

Structure  of  the  study  

Paul Bentzen, Ian Paterson (Univ. Dalhousie)

Open canopy

h2 = 0.16

Mature  male  size  

Reproduc+on  DEAD ALIVE

DEAD ALIVE

Random effects: Groups Name Variance Std.Dev. FishID (Intercept) 2.09198 1.44637 Sampling (Intercept) 0.59734 0.77288 Number of obs: 661, groups: FishID, 101; fSampling, 12Fixed effects: Estimate Std. Error z value Pr(>|z|) (Intercept) -7.97496 4.07333 -1.958 0.0502 .SL 0.01998 0.22781 0.088 0.9301 aliveFALSE 1.26180 3.23465 0.390 0.6965 SL:aliveFALSE 0.02183 0.18019 0.121 0.9036

Random effects: Groups Name Variance Std.Dev. FishID (Intercept) 1.06576 1.03236 fSampling (Intercept) 0.45851 0.67713 Number of obs: 737, groups: FishID, 107; fSampling, 12Fixed effects: Estimate Std. Error z value Pr(>|z|) (Intercept) 2.1571 3.4374 0.628 0.5303 SL -0.5340 0.2004 -2.665 0.0077 ** aliveFALSE -16.7792 3.6527 -4.594 4.35e-06 ***SL:aliveFALSE 1.0186 0.2124 4.796 1.62e-06 ***

Gupp

ies  (HP

)  Co

ntrol  

Natural    resource  levels  

Gupp

ies  (HP

)  Co

ntrol  

High  resources  (canopy  thinned)  

Gupp

ies  (LP)  

Control  

Gupp

ies  (?)  

Control  

evolu+on  

x2  

Experimental  Evolu+on