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Long-term patch Long-term patch dynamics in the dynamics in the community shaped by community shaped by bivalves, barnacles, bivalves, barnacles, ascidians and red ascidians and red algae: multiple algae: multiple foundation species in foundation species in the White Sea shallow the White Sea shallow subtidal subtidal
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Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Mar 26, 2015

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Page 1: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Long-term patch Long-term patch dynamics in the dynamics in the

community shaped by community shaped by bivalves, barnacles, bivalves, barnacles,

ascidians and red algae: ascidians and red algae: multiple foundation multiple foundation

species in the White Sea species in the White Sea shallow subtidalshallow subtidal

Page 2: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

EugeneEugene Yakovis Yakovis Anna ArtemievaAnna Artemieva

MichaelMichael FokinFokin Marina Varfolomeeva Marina Varfolomeeva

Natalia ShunatovaNatalia Shunatova

St.-Petersburg StateUniversity, Russia

Page 3: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Facilitation by foundation species shapes Facilitation by foundation species shapes many terrestrial and benthic communitiesmany terrestrial and benthic communities

Communities with multiple functionally differentCommunities with multiple functionally differentfoundation species are poorly studiedfoundation species are poorly studied

Page 4: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Locations: 12 m deep Site 1 and 15 m deep Site 2Locations: 12 m deep Site 1 and 15 m deep Site 2

Site 165° 01.2’N35° 39.7’E

The WhiteThe WhiteSeaSea

Site 265° 00.7’N35° 41.7’E

Page 5: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Epibenthic patches (EPs) on the unstructured sedimentEpibenthic patches (EPs) on the unstructured sediment

Page 6: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Mean density of EPs Mean density of EPs (m–2): 21±2 (Site 1), 6±1 (Site 2)

Page 7: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Species diversity in and around EPsSpecies diversity in and around EPs

epibenthic patches

H’ = 2.28±0.04(24±1 cm2 patches)

119 mobile species(72 polychaetes)

111 sessile species(64 bryozans)

unstructured sediment

H’=2.62±0.02 (55 cm2 cores)

101 mobile species(78 polychaetes)

Page 8: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Biomass in and around EPsBiomass in and around EPs

epibenthic patches

wet weightof macrobenthic

organisms745 g/m2

unstructured sediment

wet weightof macrobenthic

organisms91 g/m2

Page 9: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Overall species diversityOverall species diversity

H’ = 3.36±0.06

250 speciesin total

(16 x 1 m2 frame)

117±7 species per 1 m2 frame

Page 10: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Epibenthic patches by primary substrate typeEpibenthic patches by primary substrate type

Page 11: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Primary substrate types by total areaPrimary substrate types by total area

Page 12: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Total weight of epibenthic organisms within the patchesTotal weight of epibenthic organisms within the patchesbased on different substrate typebased on different substrate type

Page 13: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Live Live Serripes groenlandicusSerripes groenlandicus stays under the sediment stays under the sediment surface and never has sessile organisms attachedsurface and never has sessile organisms attached

Page 14: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Sessile organisms can utilize a shell only Sessile organisms can utilize a shell only when a clam dieswhen a clam dies

Page 15: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Sessile organisms can utilize a shell only Sessile organisms can utilize a shell only when a clam dieswhen a clam dies

Site 1: 6.7±2.9 live individualsof Serripes groenlandicus

per m2

Site 1: 1.8±0.4 unfouled valvesof Serripes groenlandicus

per m2 on the sediment surface

Page 16: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Most sessile organisms live on secondary biogenicMost sessile organisms live on secondary biogenicsubstrates (% individuals)substrates (% individuals)

Page 17: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Principal secondary space providers are barnacles andPrincipal secondary space providers are barnacles andtheir empty shells, ascidians and red algaetheir empty shells, ascidians and red algae

Page 18: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

These are 15 top frequent associations between sessileThese are 15 top frequent associations between sessileorganisms and substrates … organisms and substrates …

Page 19: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

… … and these are top 50and these are top 50

Page 20: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Most live barnacles Most live barnacles Balanus crenatus Balanus crenatus are found on are found on primary substrate and conspecificsprimary substrate and conspecifics

Page 21: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Most ascidians Most ascidians Styela spp.Styela spp. are found on barnacles and are found on barnacles andtheir empty shellstheir empty shells

Page 22: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

As a result, epibenthic patches with following structureAs a result, epibenthic patches with following structuretypes are frequent:types are frequent:

Page 23: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Some of them look like thisSome of them look like this

Page 24: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

The observed variation in structure of epibenthic The observed variation in structure of epibenthic patchespatches::

(i)(i) results from patch dynamicsresults from patch dynamics or or

(ii)(ii) is is just a product of spatial just a product of spatial heterogeneity and variable recruitmentheterogeneity and variable recruitment

??

predictions from predictions from (ii)(ii)::

>> the structure of patches would not depend much on the structure of patches would not depend much on their agetheir age

>> the age-dependent variation of structure would not the age-dependent variation of structure would not match one observed in EPsmatch one observed in EPs

Page 25: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Number of initially empty Number of initially empty SerripesSerripes shells exposed shells exposed

Exposure (yrs)

1998 1999 2000 2003 2006 2007 2008 Total

1 2 8 10 12 12 44

2 13 13

3 10 10

4 8 4 12

5 12 4 16

6 9 9

7 10 10

8 12 12

9 10 10

10 18 18

Installed in (yr)

S = 34.4 ± 0.8 cm2

Page 26: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

1-2 yrs1-2 yrs

Serripesshell

Heteranomiasquamula

Balanuscrenatus

live

Balanuscrenatus

dead

PoriferaSpirorbide and serpulide worms

other bryozoans

Dendrobeania fruticosaEscharella sp.

Scrupocellaria arcticaStomachetosella cruenta

red algaebryozoans

Page 27: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

3-4 yrs3-4 yrs

Serripesshell

Heteranomiasquamula

Balanuscrenatus

live

Balanuscrenatus

dead

PoriferaDendrobeania fruticosaSchizomavella lineata

Stomachetosella cruenta

bryozoansPorifera

red algae

red algae

Page 28: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

5-6 yrs5-6 yrs

Serripesshell

Balanuscrenatus

live

Balanuscrenatus red algae

red algaebryozoans

Porifera

other bryozoans

Verruca stromiaDendrobeania fruticosaSchizomavella lineata

HeteranomiasquamulaStyela spp.

dead

Page 29: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

7-8 yrs7-8 yrs

Serripesshell

Balanuscrenatus

live

Balanuscrenatus red algae

red algaehydrozoansbryozoans

Verruca stromiaDendrobeania fruticosa

Escharella spp.other bryozoans

dead

HeteranomiasquamulaStyela spp.

Page 30: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

9-10 yrs9-10 yrs

Serripesshell

Balanuscrenatus

live

Balanuscrenatus red algae

red algae

Verruca stromiaHeteranomia squamulaDendrobeania fruticosaSchizomavella lineata

Escharella spp.

Styela coriaceaStyela rustica

dead

Page 31: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Relative areas of principal substrates: Relative areas of principal substrates: shellsshells, , live barnacleslive barnacles, , dead barnaclesdead barnacles, , ascidiansascidians and and

red algaered algae by exposure term (shell area as 1) by exposure term (shell area as 1)

0

2

4

6

8

1 0

1 2

1 4

1-2 yrs1-2 yrs 3-4 yrs3-4 yrs 5-6 yrs5-6 yrs 7-8 yrs7-8 yrs 9-10 yrs9-10 yrs

Page 32: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

Relative abundance of sessile organisms (% ind.) on Relative abundance of sessile organisms (% ind.) on shellsshells, , live barnacleslive barnacles, , dead barnaclesdead barnacles, , ascidiansascidians,,

red algaered algae and and otherother substrates by exposure term substrates by exposure term

1-2 yrs1-2 yrs 3-4 yrs3-4 yrs 5-6 yrs5-6 yrs 7-8 yrs7-8 yrs 9-10 yrs9-10 yrs

Page 33: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

MeanMean LogE Shannon diversity of epibenthic assemblages LogE Shannon diversity of epibenthic assemblageson on shellsshells, , live barnacleslive barnacles, , dead barnaclesdead barnacles, , ascidiansascidians,,

red algaered algae and and otherother substrates by exposure term substrates by exposure term

1-2 yrs1-2 yrs 3-4 yrs3-4 yrs 5-6 yrs5-6 yrs 7-8 yrs7-8 yrs 9-10 yrs9-10 yrs

Page 34: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversity (stress=0.08)Shannon diversity (stress=0.08)

w/o substrate

on Serripes shells

on other substrates

on live snails

Page 35: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity1-2 yrs1-2 yrs

Page 36: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity3-4 yrs3-4 yrs

Page 37: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity5-6 yrs5-6 yrs

Page 38: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity7-8 yrs7-8 yrs

Page 39: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity9-10 yrs9-10 yrs

Page 40: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

nMDS of natural and experimental EPs on weights of nMDS of natural and experimental EPs on weights of ascidians, live and dead barnacles, red algae and ascidians, live and dead barnacles, red algae and

Shannon diversityShannon diversity1-10 yrs1-10 yrs

1-21-23-43-45-65-67-87-8

9-109-10

Page 41: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

The observed variation in structure of epibenthic The observed variation in structure of epibenthic patchespatches::

(i)(i) results from patch dynamicsresults from patch dynamics or or

(ii)(ii) is is just a product of spatial just a product of spatial heterogeneity and variable recruitmentheterogeneity and variable recruitment

??

predictions from predictions from (ii)(ii)::

>> the structure of patches would not depend much on the structure of patches would not depend much on their agetheir age

>> the age-dependent variation of structure would not the age-dependent variation of structure would not match one observed in EPsmatch one observed in EPs

Page 42: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

The observed variation in structure of epibenthic The observed variation in structure of epibenthic patchespatches::

(i)(i) results from patch dynamicsresults from patch dynamics or or

(ii)(ii) is is just a product of spatial just a product of spatial heterogeneity and variable recruitmentheterogeneity and variable recruitment

??

predictions from predictions from (ii)(ii)::

>> the structure of patches would not depend much on the structure of patches would not depend much on their agetheir age

>> the age-dependent variation of structure would not the age-dependent variation of structure would not match one observed in EPsmatch one observed in EPs

Page 43: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

The observed variation in structure of epibenthic The observed variation in structure of epibenthic patchespatches::

(i)(i) results from patch dynamicsresults from patch dynamics or or

(ii)(ii) is is just a product of spatial just a product of spatial heterogeneity and variable recruitmentheterogeneity and variable recruitment

??

predictions from predictions from (ii)(ii)::

>> the structure of patches would not depend much on the structure of patches would not depend much on their agetheir age

>> the age-dependent variation of structure would not the age-dependent variation of structure would not match one observed in EPsmatch one observed in EPs

Page 44: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

The observed variation in structure of epibenthic The observed variation in structure of epibenthic patchespatches::

(i)(i) results from patch dynamicsresults from patch dynamics or or

(ii)(ii) is is just a product of spatial just a product of spatial heterogeneity and variable recruitmentheterogeneity and variable recruitment

??

predictions from predictions from (ii)(ii)::

>> the structure of patches would not depend much on the structure of patches would not depend much on their agetheir age

>> the age-dependent variation of structure would not the age-dependent variation of structure would not match one observed in EPsmatch one observed in EPs

Page 45: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

CCrreeddiittss

Funding: RFBR grants Funding: RFBR grants № № 02-04-50020-a, 05-04-48927-a, 05-04-02-04-50020-a, 05-04-48927-a, 05-04-63041-k, 06-04-63077-k, 06-04-58536-z, 06-04-58537-z, 07-04-63041-k, 06-04-63077-k, 06-04-58536-z, 06-04-58537-z, 07-04-10075-k, 07-04-08366-z, 08-04-01373-a, 08-04-10109-k, 09-04-10075-k, 07-04-08366-z, 08-04-01373-a, 08-04-10109-k, 09-04-

10092-k, 10-04-08011-z10092-k, 10-04-08011-z

Alexander TcherenkovAlexander TcherenkovNadezhda TcherenkovaNadezhda Tcherenkova

Alexey GrishankovAlexey GrishankovAlexandra YakovisAlexandra Yakovis

Dmitry TomanovskyDmitry Tomanovsky

Page 46: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

BBoonnuuss ttrraacckk

Page 47: Long-term patch dynamics in the community shaped by bivalves, barnacles, ascidians and red algae: multiple foundation species in the White Sea shallow.

a shellexposed for

9 yrs