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

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

Eudoxid Muggiaea atlantica Holoplanktonic ASEXUAL and SEXUAL reproduction RAPID development Polygastric Gametes Planula Calyconula

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) Frequency Pre Post 1980s 1980s Blackett et al (2015)

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) ECOLOGICAL IMPACTS Greve (1994), Krsinic & Njire (2001) M. atlantica Weeks 500 ind. m 3 Data source: Greve (1994) The 1989 German Bight Invasion of Muggiaea atlantica

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) ECOLOGICAL IMPACTS Greve (1994), Krsinic & Njire (2001) M. atlantica Copepods + 0 Predation Weeks Data source: Greve (1994) The 1989 German Bight Invasion of Muggiaea atlantica

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) ECOLOGICAL IMPACTS Greve (1994), Krsinic & Njire (2001) M. atlantica Copepods Chlorophyll + 0 + 0 Predation Grazing Weeks Data source: Greve (1994) The 1989 German Bight Invasion of Muggiaea atlantica

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) ECOLOGICAL IMPACTS Greve (1994), Krsinic & Njire (2001) Phosphate Chlorophyll Copepods M. atlantica + 0 + 0 + 0 Predation Grazing Nutrient uptake Weeks Data source: Greve (1994) The 1989 German Bight Invasion of Muggiaea atlantica

EXPANDING DISTRIBUTION Licandro et al (2012), Malej et al (2013), Blackett et al (2015) ECOLOGICAL IMPACTS Greve (1994), Krsinic & Njire (2001) AQUACULTURE IMPACTS Fosså et al 2003, Baxter et al 2011 Mortality

Sampling locations Loch Ewe (2003 2013) Scottish Continental Shelf Stonehaven (1999 2013) L5 (1930 2013) L4 (2009 2013) Western English Channel 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) Factor2 Source Available habitat Habitat suitability Optimal Dispersal Medium Sink Critical 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 (immigration) GRADUAL (reproduction) Abundance Abundance 1 Months 12 1 Months 12

Local production Blackett et al (2014) GRADUAL (reproduction) Abundance 1 Months 12 Significant TROUGH LOCAL PRODUCTION Significant PEAK Kendall (1976), Ibañez (1982)

12 Polygastric 10 Month 8 6 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 English Channel N

12 Polygastric 10 Month 8 6 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 English Channel N

12 ~80% RANDOM 10 Month 8 6 4 2 Abundance 1930 1932 1934 1936 1938 Local production 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 Year 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Blackett et al (2014) Reproduction = recruitment Western English Channel N

12 ~80% RANDOM ~70% LOCAL PRODUCTION 10 Month 8 6 4 2 Abundance 1930 1932 1934 1936 1938 Local production 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 Year 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Blackett et al (2014) Reproduction = recruitment Western English Channel N

12 ~80% RANDOM ~70% LOCAL PRODUCTION 10 Month 8 6 4 2 Abundance 1930 1932 1934 1936 1938 Local production 1947 1949 1957 1959 1965 1867 1969 1971 1973 1975 Year 1977 1979 1981 1983 1985 1987 1997 1999 2005 2007 2009 2011 2013 Blackett et al (2014) Reproduction = recruitment Life history? Persist indefinitely? Western English Channel N

+ Local production vs. temperature Association 0 p = < 0.05 8 10 12 14 16 18 SST ( C) Blackett et al (2014) Reproduction = recruitment Life history? Persist indefinitely? Western English Channel N

Association + 0 Local production vs. temperature p = < 0.05 15 16 C 9 10 C 8 10 12 14 16 18 SST ( C) Blackett et al (2014) Reproduction = recruitment Life history Persist indefinitely? Western English Channel N

Spring/autumn Winter Eudoxid Polygastric Polygastric Carré & Carré (1991) Reproduction = recruitment Life history Persist indefinitely? Western English Channel N

Eudoxid Polygastric Abundance + 0 Eudoxid vs. polygastric abundance 1 Weeks 52 Blackett et al (2015) Reproduction = recruitment Life history Persist indefinitely? Western English Channel N

Eudoxid Polygastric Abundance 10 C (p = <0.05) + 0 Eudoxid vs. polygastric abundance 1 Weeks 52 Blackett et al (2015) Reproduction = recruitment Life history Persist indefinitely Western English Channel N

Reproduction = recruitment Life history Persist indefinitely SOURCE N

? Scottish Continental Shelf? SOURCE

Scottish Continental Shelf 12 Polygastric 10 Month 8 6 4 2 2003 2005 2007 2009 2011 2013 Year Abundance Blackett et al (submitted) Month 12 10 8 6 4 2 Polygastric Abundance 1999 2001 2003 2005 2007 2009 2011 2013 Year Blackett et al (submitted)

Reproduction = recruitment Scottish Continental Shelf 12 Polygastric 10 Month 8 6 4 2 2003 2005 2007 2009 2011 2013 Year Abundance Blackett et al (submitted) Month 12 10 8 6 4 2 Polygastric Abundance 1999 2001 2003 2005 2007 2009 2011 2013 Year Blackett et al (submitted)

Reproduction = recruitment Life history? Persist indefinitely? 13 15 C < 11 / 8 C Scottish Continental Shelf < 8 / 6 C Abundance vs. temperature Abundance vs. temperature + + p = < 0.05 p = < 0.05 Association 0 Association 0 7 8 9 10 11 12 13 14 15 SST ( C) Blackett et al (submitted) 5 7 10 12 13 15 SST ( C) Blackett et et al al (submitted)

Reproduction = recruitment Life history Persist indefinitely Scottish Continental Shelf Immigration? SOURCE

European Slope Current Poleward flow Current Warm, saline water Disperses plankton Slope European SOURCE

Standard anomalies 3 2 1 0-1 -2-3 M. atlantica vs. Slope Current 2003 2004 2005 2006 Blackett et al (submitted) 2007 Year 2008 2009 r = 0.74 p = < 0.05 2010 2011 2012 European Current Slope Slope Current sampling stations Xu et al 2015 SOURCE

Muggiaea atlantica?? Current Increased immigration Improved conditions European Slope Slope Current SOURCE

Partial linear regression Peres-Neto et al (2006) Explanatory variables Source Demographics (SST) Sink Immigration (Slope Current) Dependent variable Muggiaea atlantica variation

Partial linear regression Peres-Neto et al (2006) Explanatory variables Source Demographics (SST) Sink Immigration (Slope Current) Dependent variable Muggiaea atlantica variation

Partial linear regression Peres-Neto et al (2006) Source Sink Explanatory variables Demographics (SST) Intersect Immigration (Slope Current) Dependent variable Muggiaea atlantica variation

% of explained variation in annual abundance Scottish Continental Shelf Blackett et al (submitted) SST & Slope Current 62% Slope p = < 0.05 European Current SOURCE

% of explained variation in annual abundance SST 33% Slope Current 49% Current Slope Scottish Continental Shelf Blackett et al (submitted) p = < 0.05 European SOURCE

Total variation (100%) Scottish Continental Shelf SST 13% Intersect 20% Slope Current 29% Current Slope Residual 38% p = < 0.05 European Blackett et al (submitted) SOURCE

Current SINK Scottish Continental Shelf Slope European SOURCE

Current SINK Scottish Continental Shelf 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 PC1 SAL. 2 1 3 12 4 11 5 10 6 9 7 TEMP. 8 PREY PC 2

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

Inter-annual environmental variability Sink years Source years PC 2 PC1