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T H E 	
   E C O L O G Y 	
   A N D 	
   E V O L U T I O N 	
   O F 	
  




HEROEN VERBRUGGEN, UNIVERSITY                            OF    MELBOURNE
Five	
  slides	
  about	
  taxonomy	
  
What	
  we	
  think	
  we	
  know	
  about	
  species	
  diversity	
  but	
  don’t.	
  
                                        	
  

          Pa<erns	
  of	
  species	
  diversity	
  
            Diversifica@on:	
  specia@on	
  and	
  ex@nc@on.	
  
                                  	
  

  Geographic	
  pa<erns	
  of	
  specia@on	
  
                 How	
  biodiversity	
  got	
  to	
  where	
  it	
  is.	
  
                                     	
  

 Ecological	
  aspects	
  of	
  diversifica@on	
  
       How	
  diversifica@on	
  relates	
  to	
  the	
  thermal	
  niche.	
  
                                   	
  
                                   	
  
                                   	
  
Popula@on	
  structure	
  of	
  Por$eria	
  




                                                              Payo	
  et	
  al.	
  –	
  under	
  review	
  
Picture credits: Rob Anderson & John Huisman, via AlgaeBase
Sampling	
  locali@es:	
  Philippines	
  




                                    Payo	
  et	
  al.	
  –	
  under	
  review	
  
Not	
  one	
  but	
  21	
  species	
  




                     265	
  cox	
  2-­‐3	
  spacer	
  sequences	
  
       logLGMYC	
  	
  =	
  462.92	
  >	
  logLnul	
  =	
  449.25,	
  p	
  <	
  0.001	
  
                  21	
  (19-­‐24)	
  GMYC	
  clusters	
  


                                                                                            Payo	
  et	
  al.	
  –	
  under	
  review	
  
Global	
  diversity?	
  




                           Payo	
  et	
  al.	
  -­‐	
  unpublished	
  
Consequences	
  


Taxonomists	
  in	
  state	
  of	
  paralysis	
  

             Conserva@on…	
  

  More	
  work	
  for	
  me	
  	
  	
  @$#N*!ˇ	
  
Biodiversity	
  
       Pa<erns	
  of	
  species	
  diversity	
  
         Diversifica@on:	
  specia@on	
  and	
  ex@nc@on	
  
                                       	
  
Geographic	
  pa<erns	
  of	
  specia@on	
  
            How	
  did	
  biodiversity	
  get	
  where	
  it	
  is?	
  
                                       	
  
Ecological	
  aspects	
  of	
  diversifica@on	
  
 How	
  does	
  diversifica@on	
  relate	
  to	
  the	
  thermal	
  niche?	
  
How	
  we	
  do	
  what	
  we	
  do…	
  
       Species	
  delimita@on	
  
         Trust	
  no-­‐one	
  but	
  Mr.	
  Sanger	
  
                              	
  

     Phylogene@c	
  hypothesis	
  
                Mul@ple	
  markers	
  
             Diversifica@on	
  pa<erns	
  
                        	
  

      Evolu@onary	
  inference	
  
                Geographic	
  data	
  
              Macroecological	
  data	
  
              Evolu@onary	
  models	
  
Model	
  systems	
  




Halimeda	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  Dictyota	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  	
  Codium	
  
Geographic	
  pa<erns	
  
 of	
  diversifica@on	
  
Dictyotales	
  
Dictyotales:	
  236	
  species;	
  2038	
  specimens	
  
 	
  What	
  we	
  want	
  to	
  know	
  	
  	
  
                                                                                              	
  
              What	
  are	
  the	
  dominant	
  geographic	
  modes	
  of	
  specia@on?	
  
                                                            Where	
  did	
  taxa	
  originate?	
  	
  
                                      How	
  did	
  they	
  spread	
  around	
  the	
  world?	
  


What	
  we	
  have	
  to	
  work	
  with	
  
    	
  
    DNA	
  barcodes	
  
         species	
  boundaries	
  
         where	
  species	
  are	
  distributed	
  
    mul@gene	
  alignment	
  
         phylogene@c	
  tree	
  
         with	
  calibra@on	
  à	
  chronogram	
  
Models	
  of	
  evolu@onary	
  change	
  
                                                 Dispersal	
  -­‐	
  Ex@nc@on	
  -­‐	
  Cladogenesis	
  model	
  




Ree	
  &	
  Smith	
  (2008)	
  Syst.	
  Biol.	
  57:	
  4-­‐14	
  	
  	
  —	
  	
  	
  Ree	
  et	
  al.	
  (2005)	
  Evolu$on	
  59:	
  2299-­‐2311	
  
Basic	
  marine	
  DEC	
  model	
  




Spalding	
  et	
  al.	
  (2009)	
  BioScience	
  57:	
  573-­‐583	
  
Where	
  did	
  it	
  come	
  from?	
  




De	
  Clerck	
  et	
  al.	
  –	
  unpublished	
  
Newhousia imbricata
Zonaria sp1
Zonaria crenata
  Zonaria diesingiana
Zonaria tournefortii
Zonaria angustata
Zonaria spiralis
Zonaria subarticulata
Lobophora papenfussii
Lobophora sp12
Lobophora sp9
Lobophora sp10
Lobophora sp2
    Lobophora variegata
Lobophora sp7
Zonaria diesingiana2
Lobophora sp3
  Lobophora sp1
Lobophora sp4
Lobophora sp5
Homoeostrichus canaliculatus
Homoeostrichus olsenii
Exallosorus harveyanus
Homoeostrichus sinclairii
Homoeostrichus sp1
Homoeostrichus sp2
Homoeostrichus flabellatus
Homoeostrichus formosana
Zonaria sp2
Distromium didymothrix
Distromium sp1
Distromium skottsbergii
Distromium sp2
Distromium decumbens2
Distromium decumbens4
Distromium decumbens3
Distromium decumbens1
Padina melemele7
Padina melemele6
Padina melemele8
Padina melemele2
Padina melemele3
Padina melemele9
Padina melemele1
Padina melemele5
Padina melemele4
Padina moffittiana
Padina moffittiana2
Padina stipitata
Padina sp26
Padina sp29
Padina sp9
Padina sp10
Padina sp11
Padina sp24
Padina arborescens
Padina sp6
Padina sp25
  Padina sp5
Padina crassa
Padina crassa3
Padina gymnospora5
Padina sp8
Padina sp7
Padina gymnospora4
Padina gymnospora1
Padina gymnospora2
  Padina gymnospora3
Padina pavonica2
Padina sp21
Padina sp20
Padina sp28
Padina crassa2
Padina sp22
  Padina sp3
  Padina sp2
  Padina minor
Padina boryana4
Padina boryana3
Padina boryana2
  Padina boryana
Padina sp15
Padina somalensis
  Padina australis
Padina boergesenii
Padina pavonica
Padina thivyi
Padina ryukyuana
Padina sp4
    Padina sp1
Padina fasciata
Padina sanctaecrucis1
Padina sanctaecrucis2
  Padina japonica
Padina sp23
Padina undulata
Padina jonesii
Padina terricolor
Padina okinawaensis
Padina sp27
Taonia atomaria
Dictyopteris divaricata
Spatoglossum pacificum
Dictyopteris undulata
Dictyopteris muelleri
Spatoglossum stipitatum
Spatoglossum asperum
Spatoglossum sp
Spatoglossum crassum
Spatoglossum solierii
Lobospira bicuspidata
Dictyopteris sp1
  Dictyopteris delicatula1
Dictyopteris delicatula2
Dictyopteris sp2
Dictyopteris delicatula3
  Dictyopteris australis
Dictyopteris polypodioides
Dictyopteris longifolia
Dictyopteris latiuscula
Dictyopteris prolifera2
Dictyopteris prolifera1
Dictyopteris crassinervia
  Stypopodium flabelliforme2
Stypopodium flabelliforme
Stypopodium flabelliforme1
Stypopodium hawaiiensis
Stypopodium australasicum
Stypopodium multipartitum
  Stypopodium schimperi
Stypopodium sp1
Stypopodium zonale
Stypopodium sp2
Dictyotopsis propagulifera
Stoechospermum polypodioides
Scoresbyella profunda
Rugulopteryx radicans
Rugulopteryx suhrii
Rugulopteryx okamurae
  Rugulopteryx marginatus
Canistrocarpus crispatus
    Canistrocarpus cervicornis
Dilophus robustus2
Dilophus robustus1
Dilophus sp1
Dilophus gunnianus
Dilophus robustus3
Dilophus fastigiatus
  Dictyota ceylanica5
Dictyota humifusa
  Dictyota pfaffii
Dictyota canaliculata1
Dictyota sp15
Dictyota spiralis
  Dictyota canariensis
  Dictyota implexa
Dictyota linearis
  Dictyota sp20
Dictyota flabellata
Dictyota crenulata
Dictyota sp19
Dictyota pinnatifida
Dictyota concrescens
Dictyota cymatophila
Dictyota mertensii2
Dictyota caribaea
Dictyota pleiacantha2
Dictyota pleiacantha1
Dictyota jamaicensis
  Dictyota sandvicensis2
Dictyota sandvicensis3
Dictyota sandvicensis1
Dictyota nigricans
Dictyota binghamiae
Dictyota kunthii
Dictyota sp29
Dictyota sp25
  Dictyota dichotoma1
Dictyota sp1
Dictyota polyclada1
Dictyota polyclada2
Dictyota sp30
Dictyota paniculata
Dictyota polyclada3
Dictyota adnata
Dictyota sp16
  Dictyota sp22
Dictyota sp12
Dictyota mangrovestolon
Dictyota sp18
Dictyota sp17
Dictyota bartayresiana
Dictyota canaliculata2
Dictyota sp26
Dictyota ceylanica2
Dictyota ceylanica1
Dictyota ceylanica3
Dictyota sp6
Dictyota sp5
  Dictyota fasciola
Dictyota mediterranea
Dictyota friabilis1
Dictyota dichotoma2
Dictyota pardalis
    Dictyota ceylanica4
Dictyota hamifera
Dictyota sp11
Dictyota sp9
Dictyota sp10
Dictyota sp24
Dictyota acutiloba1
Dictyota acutiloba2
Dictyota grossedentata
Dictyota dhofarensis
Dictyota naevosa
Dictyota coriacea
Dictyota plectens
    Dictyota ciliolata
Dictyota rigida4
  Dictyota rigida2
  Dictyota rigida3
Dictyota rigida1
  Dictyota intermedia
Dictyota stolonifera
    Dictyota liturata
Dictyota mertensii1
Dictyota sp23
Dictyota dichotoma3
Dictyota diemensis
Dictyota cyanoloma
Dictyota alternifida
Dictyota sp21
Dictyota koreana
  Dictyota sp7
Dictyota sp3
How	
  did	
  it	
  move	
  around?	
  
                      Tropics	
  à	
  North	
                   Tropics	
  à	
  South	
     South	
  à	
  North	
  




                                                                                                                         dispersal	
  rate	
  
                            Ma




De	
  Clerck	
  et	
  al.	
  –	
  unpublished	
  
Cenozoic:	
  gradual	
  drop	
  in	
  temperature	
  




Zachos	
  et	
  al.	
  (2008)	
  Nature	
  451:	
  279-­‐283	
  
Ecological	
  correlates	
  
  of	
  diversifica@on	
  
Halimeda	
  –	
  Species	
  diversity	
  map	
  




Map	
  courtesy	
  of	
  Tom	
  Schils	
  
Macroecological	
  affini@es	
  




                                                                                            SST




Verbruggen	
  et	
  al.	
  (2009)	
  Global	
  Ecol.	
  Biogeogr.	
  18:	
  393-­‐405	
  
SST	
  affini@es	
  modeled	
  along	
  tree	
  

                                           C1	
                                             C2	
     C3	
     C4	
     C5	
  




Verbruggen	
  et	
  al.	
  (2009)	
  Global	
  Ecol.	
  Biogeogr.	
  18:	
  393-­‐405	
  
Can	
  this	
  explain	
  geographical	
  
    pa<erns	
  of	
  diversity?	
  
The	
  notorious	
  LDG	
  

                                                                                                                                                                low



                                                                                                                                                                high




                                    Decreasing	
  species	
  diversity	
  from	
  tropics	
  to	
  poles	
  
                                              Many,	
  many,	
  MANY	
  explana@ons	
  proposed	
  	
  
                                               Evolu@onary	
  perspec@ve:	
  species	
  turnover	
  


Ti<ensor	
  et	
  al.	
  (2010)	
  Nature	
  466:	
  1098-­‐1107	
  	
  	
  —	
  	
  	
  Jablonski	
  et	
  al.	
  (2006)	
  Science	
  314:	
  102-­‐106	
  
…but	
  for	
  seaweeds	
  




                                                 Bimodal	
  la@tudinal	
  diversity	
  pa<ern	
  
                                              Highest	
  diversity	
  in	
  temperate	
  regions	
  
                                        Do	
  the	
  same	
  evolu@onary	
  processes	
  hold?	
  

Kerswell	
  (2006)	
  Ecology	
  10:	
  2479-­‐2488	
  
Codium	
  -­‐	
  Species	
  diversity	
  map	
  




Map	
  courtesy	
  of	
  Tom	
  Schils	
  
Codium	
  -­‐	
  SST	
  affini)es	
  along	
  tree	
  
     C1	
        C2	
                             C3	
  




                                        l    112	
  species	
  
                                        l    3	
  genes	
  
                                        l    ML	
  –	
  BI	
  
                                        l    )me-­‐calibrated	
  
                                        l    evolu)on	
  of	
  SST	
  
                                              preference	
  inferred	
  
                                 	
  

                                                           Verbruggen	
  et	
  al.	
  -­‐	
  unpublished	
  
Codium	
  -­‐	
  SST	
  affini@es	
  along	
  tree	
  
     C1	
              C2	
                                C3	
  




              Are	
  speciaIon	
  rates          	
  
              higher	
  in	
  the	
  tropics?    l 

                                        3	
  genes	
  
                                                 l 
                                                      	
  
                                        112	
  species	
  

                                                 l    ML	
  –	
  BI	
  
                                                 l    @me-­‐calibrated	
  
                                                 l    evolu@on	
  of	
  SST	
  
                                                       preference	
  inferred	
  
                                          	
  

                                                                    Verbruggen	
  et	
  al.	
  -­‐	
  unpublished	
  
Model	
  of	
  diversifica@on	
  dynamics	
  


  Diversifica@on	
  func@on	
  of	
  trait	
  of	
  interest	
  
             (in	
  our	
  case,	
  SST)	
  



               " λ %
           log $      ' = β SST + α
               # 1− λ &



                                                              Verbruggen	
  et	
  al.	
  -­‐	
  unpublished	
  
Codium	
  -­‐	
  Diversifica@on	
  and	
  SST	
  
  C1	
         C2	
                        C3	
  




                        β = 0.18 ± 0.026




                                                    Verbruggen	
  et	
  al.	
  -­‐	
  unpublished	
  
Codium	
  -­‐	
  SST	
  affini)es	
  along	
  tree	
  
     C1	
        C2	
                             C3	
  




                                        l    112	
  species	
  
                                        l    3	
  genes	
  
                                        l    ML	
  –	
  BI	
  
                                        l    )me-­‐calibrated	
  
                                        l    evolu)on	
  of	
  SST	
  
                                              preference	
  inferred	
  
                                 	
  

                                                           Verbruggen	
  et	
  al.	
  -­‐	
  unpublished	
  
β	
  =	
  0.18	
  ±	
  0.026	
  




                     β	
  =	
  0.018	
  ±	
  0.021	
  




                     Verbruggen	
  et	
  al.	
  –	
  unpublished—	
  	
  	
  Tyberghein	
  et	
  al.	
  –	
  unpublished	
  
Diversifica@on	
  is	
  func@on	
  
   of	
  niche	
  evolvability	
  




                                     Tyberghein	
  et	
  al.	
  –	
  unpublished	
  
Parapatric	
  specia@on	
  along	
  
la@tudinal	
  temperature	
  gradient	
  




                                    Tyberghein	
  et	
  al.	
  –	
  unpublished	
  
Where	
  does	
  this	
  
evolvability	
  come	
  from?	
  
Halimeda	
  –	
  microhabitat	
  evolu@on	
  




Exposed	
  microhabitat	
                     Sheltered	
  microhabitat	
  




                                                   Marcelino	
  et	
  al.	
  –	
  unpublished	
  
SST	
  rate	
               SST	
  rate	
  depends	
  
constant	
                    on	
  exposure	
  




                  vs.	
  




                                        Marcelino	
  et	
  al.	
  –	
  unpublished	
  
Microhabitat	
  specializa@ons	
  as	
  
                exapta@ons	
  for	
  macroecological	
  shiws	
  
                                      7.6	
  
frequency	
  




                                                                   Lineages	
  from	
  
                                                                   exposed	
  habitat	
  
                                                                   4.3×	
  faster	
  




                                  AIC1rate	
  –	
  AIC2rates	
  
                                                                    Marcelino	
  et	
  al.	
  –	
  unpublished	
  
Evolu@on	
  &	
  diversity	
  
       	
     	
  SpeciaIon	
  
                                   allopatric	
  	
  vs.	
  	
  within	
  region	
  
                                  niche	
  shiws	
  	
  vs.	
  	
  conserva@sm	
  
	
  
       	
     	
  DistribuIons	
  
                                  tropics	
  source:	
  OTT	
  confirmed	
  
                                               North	
  sink	
  
	
  
       	
     	
  DiversificaIon	
  
                               Codium:	
  SST	
  +	
  historical	
  effect	
  
                              Halimeda	
  &	
  Dictyota:	
  evolvability	
  
Lennert	
  Tyberghein	
          Vanessa	
  Marcelino	
             Sofie	
  D’hondt	
  




Dioli	
  Ann	
  Payo	
         Olivier	
  De	
  Clerck	
      Frederik	
  Leliaert	
  
Collaborators:	
                    	
                                  	
  
Ma<	
  Ashworth	
                   Chris	
  Maggs	
                    Satoshi	
  Shimada	
  
Eric	
  Coppejans	
                 Antoine	
  N'Yeurt	
                Thomas	
  Silberfeld	
  
Chris	
  Drake	
                    Fa@ma	
  Oliveira	
                 Heather	
  Spalding	
  
Nick	
  Eloot	
                     Mariana	
  Oliveira	
               Frederique	
  Steen	
  
Wiebe	
  Kooistra	
                 Klaas	
  Pauly	
                    Ana	
  Tronholm	
  
Diane	
  Li<ler	
                   Gary	
  Saunders	
                  Christophe	
  Viera	
  
Mark	
  Li<ler	
                    Thomas	
  Sauvage	
                 Brian	
  Wysor	
  
Steve	
  LoDuca	
                   Tom	
  Schils	
                     Rick	
  Zechman	
  

Collectors:	
               	
                         	
                       	
  
Rob	
  Anderson	
           Stefano	
  Draisma	
       Chris	
  Lane	
          Willem	
  P.v.Reine	
  
Elizabeth	
  Bandeira	
     Cindy	
  Fernandez	
       Line	
  Le	
  Gall	
     Craig	
  Schneider	
  
John	
  Bolton	
            Suzanne	
  Fredericq	
     Diane	
  Li<ler	
        Gary	
  Saunders	
  
Barre<	
  Brooks	
          Wilson	
  Freshwater	
     Mark	
  Li<ler	
         Thomas	
  Sauvage	
  
Francis	
  Bunker	
         Fred	
  Gurgel	
           Lydiane	
  Ma|o	
        Tom	
  Schils	
  
Olivier	
  Dargent	
        John	
  Huisman	
          Frederic	
  Mineur	
     Heather	
  Spalding	
  
Kya<	
  Dixon	
             Lisa	
  Kirkendale	
       Klaas	
  Pauly	
         John	
  West	
  
Rainbo	
  Dixon	
           Gerry	
  Kraw	
            Claude	
  Payri	
        …	
  many	
  others	
  …	
  

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The Ecology and Evolution of Seaweed Diversification

  • 1. T H E   E C O L O G Y   A N D   E V O L U T I O N   O F   HEROEN VERBRUGGEN, UNIVERSITY OF MELBOURNE
  • 2. Five  slides  about  taxonomy   What  we  think  we  know  about  species  diversity  but  don’t.     Pa<erns  of  species  diversity   Diversifica@on:  specia@on  and  ex@nc@on.     Geographic  pa<erns  of  specia@on   How  biodiversity  got  to  where  it  is.     Ecological  aspects  of  diversifica@on   How  diversifica@on  relates  to  the  thermal  niche.        
  • 3. Popula@on  structure  of  Por$eria   Payo  et  al.  –  under  review   Picture credits: Rob Anderson & John Huisman, via AlgaeBase
  • 4. Sampling  locali@es:  Philippines   Payo  et  al.  –  under  review  
  • 5. Not  one  but  21  species   265  cox  2-­‐3  spacer  sequences   logLGMYC    =  462.92  >  logLnul  =  449.25,  p  <  0.001   21  (19-­‐24)  GMYC  clusters   Payo  et  al.  –  under  review  
  • 6. Global  diversity?   Payo  et  al.  -­‐  unpublished  
  • 7. Consequences   Taxonomists  in  state  of  paralysis   Conserva@on…   More  work  for  me      @$#N*!ˇ  
  • 8. Biodiversity   Pa<erns  of  species  diversity   Diversifica@on:  specia@on  and  ex@nc@on     Geographic  pa<erns  of  specia@on   How  did  biodiversity  get  where  it  is?     Ecological  aspects  of  diversifica@on   How  does  diversifica@on  relate  to  the  thermal  niche?  
  • 9. How  we  do  what  we  do…   Species  delimita@on   Trust  no-­‐one  but  Mr.  Sanger     Phylogene@c  hypothesis   Mul@ple  markers   Diversifica@on  pa<erns     Evolu@onary  inference   Geographic  data   Macroecological  data   Evolu@onary  models  
  • 10.
  • 11.
  • 12.
  • 13. Model  systems   Halimeda                        Dictyota                                                                    Codium  
  • 14. Geographic  pa<erns   of  diversifica@on  
  • 16. Dictyotales:  236  species;  2038  specimens  
  • 17.    What  we  want  to  know         What  are  the  dominant  geographic  modes  of  specia@on?   Where  did  taxa  originate?     How  did  they  spread  around  the  world?   What  we  have  to  work  with     DNA  barcodes   species  boundaries   where  species  are  distributed   mul@gene  alignment   phylogene@c  tree   with  calibra@on  à  chronogram  
  • 18. Models  of  evolu@onary  change   Dispersal  -­‐  Ex@nc@on  -­‐  Cladogenesis  model   Ree  &  Smith  (2008)  Syst.  Biol.  57:  4-­‐14      —      Ree  et  al.  (2005)  Evolu$on  59:  2299-­‐2311  
  • 19. Basic  marine  DEC  model   Spalding  et  al.  (2009)  BioScience  57:  573-­‐583  
  • 20. Where  did  it  come  from?   De  Clerck  et  al.  –  unpublished  
  • 21. Newhousia imbricata Zonaria sp1 Zonaria crenata Zonaria diesingiana Zonaria tournefortii Zonaria angustata Zonaria spiralis Zonaria subarticulata Lobophora papenfussii Lobophora sp12 Lobophora sp9 Lobophora sp10 Lobophora sp2 Lobophora variegata Lobophora sp7 Zonaria diesingiana2 Lobophora sp3 Lobophora sp1 Lobophora sp4 Lobophora sp5 Homoeostrichus canaliculatus Homoeostrichus olsenii Exallosorus harveyanus Homoeostrichus sinclairii Homoeostrichus sp1 Homoeostrichus sp2 Homoeostrichus flabellatus Homoeostrichus formosana Zonaria sp2 Distromium didymothrix Distromium sp1 Distromium skottsbergii Distromium sp2 Distromium decumbens2 Distromium decumbens4 Distromium decumbens3 Distromium decumbens1 Padina melemele7 Padina melemele6 Padina melemele8 Padina melemele2 Padina melemele3 Padina melemele9 Padina melemele1 Padina melemele5 Padina melemele4 Padina moffittiana Padina moffittiana2 Padina stipitata Padina sp26 Padina sp29 Padina sp9 Padina sp10 Padina sp11 Padina sp24 Padina arborescens Padina sp6 Padina sp25 Padina sp5 Padina crassa Padina crassa3 Padina gymnospora5 Padina sp8 Padina sp7 Padina gymnospora4 Padina gymnospora1 Padina gymnospora2 Padina gymnospora3 Padina pavonica2 Padina sp21 Padina sp20 Padina sp28 Padina crassa2 Padina sp22 Padina sp3 Padina sp2 Padina minor Padina boryana4 Padina boryana3 Padina boryana2 Padina boryana Padina sp15 Padina somalensis Padina australis Padina boergesenii Padina pavonica Padina thivyi Padina ryukyuana Padina sp4 Padina sp1 Padina fasciata Padina sanctaecrucis1 Padina sanctaecrucis2 Padina japonica Padina sp23 Padina undulata Padina jonesii Padina terricolor Padina okinawaensis Padina sp27 Taonia atomaria Dictyopteris divaricata Spatoglossum pacificum Dictyopteris undulata Dictyopteris muelleri Spatoglossum stipitatum Spatoglossum asperum Spatoglossum sp Spatoglossum crassum Spatoglossum solierii Lobospira bicuspidata Dictyopteris sp1 Dictyopteris delicatula1 Dictyopteris delicatula2 Dictyopteris sp2 Dictyopteris delicatula3 Dictyopteris australis Dictyopteris polypodioides Dictyopteris longifolia Dictyopteris latiuscula Dictyopteris prolifera2 Dictyopteris prolifera1 Dictyopteris crassinervia Stypopodium flabelliforme2 Stypopodium flabelliforme Stypopodium flabelliforme1 Stypopodium hawaiiensis Stypopodium australasicum Stypopodium multipartitum Stypopodium schimperi Stypopodium sp1 Stypopodium zonale Stypopodium sp2 Dictyotopsis propagulifera Stoechospermum polypodioides Scoresbyella profunda Rugulopteryx radicans Rugulopteryx suhrii Rugulopteryx okamurae Rugulopteryx marginatus Canistrocarpus crispatus Canistrocarpus cervicornis Dilophus robustus2 Dilophus robustus1 Dilophus sp1 Dilophus gunnianus Dilophus robustus3 Dilophus fastigiatus Dictyota ceylanica5 Dictyota humifusa Dictyota pfaffii Dictyota canaliculata1 Dictyota sp15 Dictyota spiralis Dictyota canariensis Dictyota implexa Dictyota linearis Dictyota sp20 Dictyota flabellata Dictyota crenulata Dictyota sp19 Dictyota pinnatifida Dictyota concrescens Dictyota cymatophila Dictyota mertensii2 Dictyota caribaea Dictyota pleiacantha2 Dictyota pleiacantha1 Dictyota jamaicensis Dictyota sandvicensis2 Dictyota sandvicensis3 Dictyota sandvicensis1 Dictyota nigricans Dictyota binghamiae Dictyota kunthii Dictyota sp29 Dictyota sp25 Dictyota dichotoma1 Dictyota sp1 Dictyota polyclada1 Dictyota polyclada2 Dictyota sp30 Dictyota paniculata Dictyota polyclada3 Dictyota adnata Dictyota sp16 Dictyota sp22 Dictyota sp12 Dictyota mangrovestolon Dictyota sp18 Dictyota sp17 Dictyota bartayresiana Dictyota canaliculata2 Dictyota sp26 Dictyota ceylanica2 Dictyota ceylanica1 Dictyota ceylanica3 Dictyota sp6 Dictyota sp5 Dictyota fasciola Dictyota mediterranea Dictyota friabilis1 Dictyota dichotoma2 Dictyota pardalis Dictyota ceylanica4 Dictyota hamifera Dictyota sp11 Dictyota sp9 Dictyota sp10 Dictyota sp24 Dictyota acutiloba1 Dictyota acutiloba2 Dictyota grossedentata Dictyota dhofarensis Dictyota naevosa Dictyota coriacea Dictyota plectens Dictyota ciliolata Dictyota rigida4 Dictyota rigida2 Dictyota rigida3 Dictyota rigida1 Dictyota intermedia Dictyota stolonifera Dictyota liturata Dictyota mertensii1 Dictyota sp23 Dictyota dichotoma3 Dictyota diemensis Dictyota cyanoloma Dictyota alternifida Dictyota sp21 Dictyota koreana Dictyota sp7 Dictyota sp3
  • 22. How  did  it  move  around?   Tropics  à  North   Tropics  à  South   South  à  North   dispersal  rate   Ma De  Clerck  et  al.  –  unpublished  
  • 23. Cenozoic:  gradual  drop  in  temperature   Zachos  et  al.  (2008)  Nature  451:  279-­‐283  
  • 24. Ecological  correlates   of  diversifica@on  
  • 25. Halimeda  –  Species  diversity  map   Map  courtesy  of  Tom  Schils  
  • 26. Macroecological  affini@es   SST Verbruggen  et  al.  (2009)  Global  Ecol.  Biogeogr.  18:  393-­‐405  
  • 27. SST  affini@es  modeled  along  tree   C1   C2   C3   C4   C5   Verbruggen  et  al.  (2009)  Global  Ecol.  Biogeogr.  18:  393-­‐405  
  • 28. Can  this  explain  geographical   pa<erns  of  diversity?  
  • 29. The  notorious  LDG   low high Decreasing  species  diversity  from  tropics  to  poles   Many,  many,  MANY  explana@ons  proposed     Evolu@onary  perspec@ve:  species  turnover   Ti<ensor  et  al.  (2010)  Nature  466:  1098-­‐1107      —      Jablonski  et  al.  (2006)  Science  314:  102-­‐106  
  • 30. …but  for  seaweeds   Bimodal  la@tudinal  diversity  pa<ern   Highest  diversity  in  temperate  regions   Do  the  same  evolu@onary  processes  hold?   Kerswell  (2006)  Ecology  10:  2479-­‐2488  
  • 31. Codium  -­‐  Species  diversity  map   Map  courtesy  of  Tom  Schils  
  • 32. Codium  -­‐  SST  affini)es  along  tree   C1   C2   C3   l  112  species   l  3  genes   l  ML  –  BI   l  )me-­‐calibrated   l  evolu)on  of  SST   preference  inferred     Verbruggen  et  al.  -­‐  unpublished  
  • 33. Codium  -­‐  SST  affini@es  along  tree   C1   C2   C3   Are  speciaIon  rates   higher  in  the  tropics? l  3  genes   l    112  species   l  ML  –  BI   l  @me-­‐calibrated   l  evolu@on  of  SST   preference  inferred     Verbruggen  et  al.  -­‐  unpublished  
  • 34. Model  of  diversifica@on  dynamics   Diversifica@on  func@on  of  trait  of  interest   (in  our  case,  SST)   " λ % log $ ' = β SST + α # 1− λ & Verbruggen  et  al.  -­‐  unpublished  
  • 35. Codium  -­‐  Diversifica@on  and  SST   C1   C2   C3   β = 0.18 ± 0.026 Verbruggen  et  al.  -­‐  unpublished  
  • 36. Codium  -­‐  SST  affini)es  along  tree   C1   C2   C3   l  112  species   l  3  genes   l  ML  –  BI   l  )me-­‐calibrated   l  evolu)on  of  SST   preference  inferred     Verbruggen  et  al.  -­‐  unpublished  
  • 37. β  =  0.18  ±  0.026   β  =  0.018  ±  0.021   Verbruggen  et  al.  –  unpublished—      Tyberghein  et  al.  –  unpublished  
  • 38. Diversifica@on  is  func@on   of  niche  evolvability   Tyberghein  et  al.  –  unpublished  
  • 39. Parapatric  specia@on  along   la@tudinal  temperature  gradient   Tyberghein  et  al.  –  unpublished  
  • 40. Where  does  this   evolvability  come  from?  
  • 41. Halimeda  –  microhabitat  evolu@on   Exposed  microhabitat   Sheltered  microhabitat   Marcelino  et  al.  –  unpublished  
  • 42. SST  rate   SST  rate  depends   constant   on  exposure   vs.   Marcelino  et  al.  –  unpublished  
  • 43. Microhabitat  specializa@ons  as   exapta@ons  for  macroecological  shiws   7.6   frequency   Lineages  from   exposed  habitat   4.3×  faster   AIC1rate  –  AIC2rates   Marcelino  et  al.  –  unpublished  
  • 44. Evolu@on  &  diversity      SpeciaIon   allopatric    vs.    within  region   niche  shiws    vs.    conserva@sm        DistribuIons   tropics  source:  OTT  confirmed   North  sink        DiversificaIon   Codium:  SST  +  historical  effect   Halimeda  &  Dictyota:  evolvability  
  • 45. Lennert  Tyberghein   Vanessa  Marcelino   Sofie  D’hondt   Dioli  Ann  Payo   Olivier  De  Clerck   Frederik  Leliaert  
  • 46. Collaborators:       Ma<  Ashworth   Chris  Maggs   Satoshi  Shimada   Eric  Coppejans   Antoine  N'Yeurt   Thomas  Silberfeld   Chris  Drake   Fa@ma  Oliveira   Heather  Spalding   Nick  Eloot   Mariana  Oliveira   Frederique  Steen   Wiebe  Kooistra   Klaas  Pauly   Ana  Tronholm   Diane  Li<ler   Gary  Saunders   Christophe  Viera   Mark  Li<ler   Thomas  Sauvage   Brian  Wysor   Steve  LoDuca   Tom  Schils   Rick  Zechman   Collectors:         Rob  Anderson   Stefano  Draisma   Chris  Lane   Willem  P.v.Reine   Elizabeth  Bandeira   Cindy  Fernandez   Line  Le  Gall   Craig  Schneider   John  Bolton   Suzanne  Fredericq   Diane  Li<ler   Gary  Saunders   Barre<  Brooks   Wilson  Freshwater   Mark  Li<ler   Thomas  Sauvage   Francis  Bunker   Fred  Gurgel   Lydiane  Ma|o   Tom  Schils   Olivier  Dargent   John  Huisman   Frederic  Mineur   Heather  Spalding   Kya<  Dixon   Lisa  Kirkendale   Klaas  Pauly   John  West   Rainbo  Dixon   Gerry  Kraw   Claude  Payri   …  many  others  …