Biology and Ecology of Muggiaea Atlantica in the Northeast Atlantic Mike Blackett Eudoxid

Biology and ecology of Muggiaea atlantica in the northeast Atlantic Mike Blackett Eudoxid

Gametes Muggiaea atlantica •Holoplanktonic •ASEXUAL and SEXUAL

reproduction Polygastric •RAPID development

Planula

Calyconula ION Frequency UT ) TRIB G DIS NDIN EXPA Licandro et al (2012), Malej et al (2013), Blackett et al (2015

Pre 1980s Post 1980s

Blackett et al (2015) 500 ind. m–3 TION IBU ) TR atlantica M. G DIS NDIN EXPA Licandro et al (2012), Malej et al (2013), Blackett et al (2015

IMPACTS

ECOLOGICALGreve (1994), Krsinic & Njire (2001)

Weeks (1994) The 1989 German Greve Bight Invasion of Muggiaea atlantica Data source: TION IBU ) ISTR atlantica M. NG D Predation ANDI EXP + Licandro et al (2012), Malej et al (2013), Blackett et al (2015 0 –

IMPACTS Copepods

ECOLOGICALGreve (1994), Krsinic & Njire (2001)

IMPACTS Copepods Grazing ECOLOGICAL Greve (1994), Krsinic & Njire (2001) + 0 – Chlorophyll

IMPACTS Copepods Grazing ECOLOGICAL Greve (1994), Krsinic & Njire (2001) + 0 – Chlorophyll Nutrient uptake + 0 – Phosphate

Weeks (1994) The 1989 German Greve Bight Invasion of Muggiaea atlantica Data source: !KTUELLETEMA

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Mortality NULJVVNLSµ*LIWHQIUDGHQQHHUPHUHQQJDQJHUVn IMPACTS VWHUN VRP JLIWHQ IUD NREUDVODQJHU /LNHYHO HU NRQWDNW PHG µSRUWXJLVLVNHNULJVVNLSµVMHOGHQG¡GHOLJIRUPHQQHVNHU'HWWH VN\OGHVWUROLJDWQHVOHFHOOHQHHUPLNURVNRSLVNVPnLIRUKROG Baxter et al 2011 WLONREUDVODQJHQVJLIWNMHUWOHU'HUIRUYLOJLIWPHQJGHQVMHOGHQ Y UHVWRUQRNWLOnGUHSHHQVWRUÀVNHOOHUHWPHQQHVNHPHQHU AQUACULTUREFosså et al 2003, IXOOWXWWLOVWUHNNHOLJWLOnJLVPHUWHRJVnUGDQQHOVH &IGUR 0HQXQQWDJHOVHQHÀQQHV1 U$XVWUDOLDRJ,QGRQHVLDNDQGH 3KADER PÍ LAKS ETTER KONTAKT MED KOLONIMANETEN -UGGIAEA VRPEDGHUVW¡WHSnHQOLWHQPDQHWNDOWµVM¡YHSVµVRPKDUHQ ATLANTICA&ISKENlKKBLDNINGERISLIMHINNENEMELLOMTENNENEOG STORESKADERISLIMHINNENEPÍINNSIDENAVGJELLELOKKOGPÍGJELLE JLIWVWHUNQRNWLOnJLKMHUWHVWDQVRJG¡GKRVPHQQHVNHULO¡SHW BUEN&OTO&REDDY*ENSEN &ISKEHELSEOG-ILJ!3 DYInPLQXWWHU ,ESIONS ON SALMON BURNED BY THE SIPHONOPHORE -UGGIAEA ATLANTICA 4HElSHESSUFFEREDFROMMUCOSALHEMORRHAGESBETWEENTHETEETHAND $QGUHPDQHWHUVRPJODVVPDQHWHQKDUJLIWVWRIIHUVRPYDQOLJ PROMINENTDAMAGETOTHEINSIDEOFTHEGILLOPERCULUMANDONTHEGILL YLVLNNHYLUNHUSnPHQQHVNHUPHQVRPIRUWHUG¡GHOLJHSn BOWS HNVHPSHOYLVVLOGHODUYHU

(!6%43-),* Sampling locations

Scottish Continental Loch Ewe Shelf (2003–2013) Stonehaven (1999–2013)

L5 (1930–2013) L4 (2009–2013)

Western EnglishChannel

N Historical distribution

Scottish Shelf • Exceptionally rare Totton & Fraser (1955)

Western English Channel • Transient • ‘Indicator’ species Southward et al (1995)

N Contemporary distribution…?

Persistence or transience?

N Source–sink dynamics Pulliam (2000) 2 Available habitat Habitat suitability Factor Source Optimal

Medium Dispersal

Critical

Sink Unsuitable

Factor1

Helaouët et al (2013) Source Sink

Production = recruitment Production ≠ recruitment

∞ Life history ∝Life history

Persist indeﬁnitely Require immigration

Pulliam (2000) Local production Blackett et al (2014)

RANDOM GRADUAL (immigration) (reproduction) Abundance Abundance

1 Months 12 1 Months 12 Local production Blackett et al (2014)

GRADUAL (reproduction) Abundance

1 Months 12

Signiﬁcant Signiﬁcant LOCAL PRODUCTION TROUGH PEAK

Kendall (1976), Ibañez (1982) Polygastric 12 10 8 6

Month 4 2

Abundance

1930 1932 1934 1936 1938 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Year Blackett et al (2014)

Western EnglishChannel

N Polygastric 12 10 8 6

Month 4 2

Abundance

1930 1932 1934 1936 1938 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Year Blackett et al (2014)

Reproduction = recruitment?

Western EnglishChannel

N ~80% RANDOM 12 10 8 6

Month 4 2

Abundance

1930 1932 1934 1936 1938 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Local production Year Blackett et al (2014)

Reproduction = recruitment

Western EnglishChannel

N ~80% RANDOM ~70% LOCAL PRODUCTION 12 10 8 6

Month 4 2

Abundance

1930 1932 1934 1936 1938 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Local production Year Blackett et al (2014)

Reproduction = recruitment

Western EnglishChannel

N ~80% RANDOM ~70% LOCAL PRODUCTION 12 10 8 6

Month 4 2

Abundance

1930 1932 1934 1936 1938 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Local production Year Blackett et al (2014)

Reproduction = recruitment

Life history? Western ∞ EnglishChannel Persist indeﬁnitely? N Local production vs. temperature +

p = < 0.05

0 Association

–

8 10 12 14 16 18 SST (°C) Blackett et al (2014)

Reproduction = recruitment

Life history? Western ∞ EnglishChannel Persist indeﬁnitely? N Local production vs. temperature + 15–16 °C p = < 0.05

0 Association

9–10 °C –

8 10 12 14 16 18 SST (°C) Blackett et al (2014)

Reproduction = recruitment

Life history Western ∞ EnglishChannel Persist indeﬁnitely? N Spring/autumn Winter

Eudoxid

Polygastric Polygastric

Carré & Carré (1991)

Reproduction = recruitment

Life history Western ∞ EnglishChannel Persist indeﬁnitely? N + Eudoxid 0 polygastric vs.

Polygastric – abundance Abundance Eudoxid 1 Weeks 52 Blackett et al (2015)

Reproduction = recruitment

Life history Western ∞ EnglishChannel Persist indeﬁnitely? N 10 °C (p = <0.05)

+ Eudoxid 0 polygastric vs.

Polygastric – abundance Abundance Eudoxid 1 Weeks 52 Blackett et al (2015)

Reproduction = recruitment

Life history Western ∞ EnglishChannel Persist indeﬁnitely N Reproduction = recruitment ∞ Life history SOURCE Persist indeﬁnitely N Scottish Continental ? Shelf ?

SOURCE Scottish Continental Shelf

Polygastric 12 10 8 6

Month 4 Polygastric 2 12

Abundance 10 8

2003 2005 2007 2009 2011 2013 Year 6

Month Blackett et al (submitted) 4 2

Abundance

1999 2001 2003 2005 2007 2009 2011 2013 Year Blackett et al (submitted) Reproduction = recruitment

Scottish Continental Shelf

Polygastric 12 10 8 6

Month 4 Polygastric 2 12

Abundance 10 8

2003 2005 2007 2009 2011 2013 Year 6

Month Blackett et al (submitted) 4 2

Abundance

1999 2001 2003 2005 2007 2009 2011 2013 Year Blackett et al (submitted) Reproduction = recruitment ∞ Life history? 13–15 °C Scottish Persist indeﬁnitely? Continental < 11 / 8 °C Shelf

< 8 / 6 °C

Abundance vs. temperature Abundance vs. temperature + + p = < 0.05 p = < 0.05

0 0 Association Association

– –

7 8 9 10 11 12 13 14 15 5 7 10 12 13 15 SST (°C) SST (°C)

Blackett et al (submitted) BlackettBlackett et et al al (submitted) (submitted) Reproduction = recruitment ∞ Life history Scottish Persist indeﬁnitely Continental Shelf

Immigration?

SOURCE European Slope Current

• Poleward ﬂow

• Warm, saline water Current • Disperses plankton Slope

European

SOURCE M. atlantica vs. Slope Current 3 r = 0.74 2 p = < 0.05 1 0 -1 -2

Standardanomalies -3 Current

2003 2005 2007

2004 2006 2008 2009 2011

2010 2012 Year Slope Blackett et al (submitted)

European

Slope Current sampling stations SOURCE Xu et al 2015 Muggiaea atlantica

? ? Current

Slope

European Increasedimmigration Improvedconditions

Slope Current SOURCE Partial linear regression Peres-Neto et al (2006)

Source Sink

Explanatory Demographics Immigration variables (SST) (Slope Current)

Dependent Muggiaea atlantica variable variation Partial linear regression Peres-Neto et al (2006)

Source Sink

Explanatory Demographics Immigration variables (SST) (Slope Current)

Dependent Muggiaea atlantica variable variation Partial linear regression Peres-Neto et al (2006)

Source Sink

Explanatory Demographics Immigration variables

(SST) Intersect (Slope Current)

Dependent Muggiaea atlantica variable variation % of explained variation in annual abundance Scottish Continental Shelf SST & Current Slope Current 62% Slope

pEuropean = < 0.05 Blackett et al (submitted)

SOURCE % of explained variation in annual abundance Scottish Continental Shelf Slope

SST Current Current 33% 49% Slope

pEuropean = < 0.05 Blackett et al (submitted)

SOURCE Total variation (100%)

Scottish Continental Shelf Slope

SST Intersect Current Current 13% 20% 29% Slope

Residual 38% p = < 0.05 European Blackett et al (submitted)

SOURCE Scottish Continental Shelf

SINK Current

Slope

European

SOURCE Scottish Continental Shelf SINK

Current SINK?

Slope

European

SOURCE Summary

Muggiaea atlantica has expanded its northern distribution in the northeast Atlantic

• Established a resident population in the Western English Channel

• Established transient populations on the Scottish shelf

• Complex interaction of habitat-speciﬁc demography and current driven dispersal Acknowledgements Photos: © Dennis Riek References

Blackett M, Licandro P, Coombs SH and Lucas CH (2014) Long-term variability of the siphonophores Muggiaea atlantica and M. kochi in the Western English Channel. Progress in Oceanography. 128, 1-14.

Blackett M, Lucas CH, Harmer RA and Licandro P (2015) Population ecology of Muggiaea atlantica (Cnidaria, Siphonophora) in the Western English Channel. Marine Ecolology Progress Series. 535, 129–144.

Blackett M (2015) Biology and ecology of Muggiaea atlantica in the northeast Atlantic. PhD Thesis. University of Southampton.

Blackett M, Lucas CH, Cook K and Licandro P (Submitted) Blooms of the siphonophore Muggiaea atlantica in Scottish coastal waters: source or sink? Journal of Plankton Research. Average annual cycle (compromise)

M. atlantica

1 TEMP. 12 11 10 2 9 8 PC1 7 SAL. 3 6 4 5 PREY PC 2 PC Individual annual cycles (trajectories)

SAL. 1

12 M. atlantica 11 2 10 TEMP. 9 PC1 8

7 3 6 5 PREY 4 PC 2 PC Inter-annual environmental variability

Sink years

Source years PC 2 PC PC1