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Bloom Events of Hydromedusae Bloom events of hydromedusae Blackfordia virginica, an introduced hydromedusa species Jinho Chae1, 3, ByeongHo Kim1, Changgyun Yu1, Gunhee Sung1, Yong Kim, Jang-Seu Ki2, Wonduk Yoon3 1Marine Environmental Research & Information Laboratory (MERIL), Gunpo 15850, Korea 2Department of Life Science, Sangmyung University, Seoul 03016, Korea 3Human and Marine Ecosystem Research Laboratory (HuMER), Gunpo 15850, Korea Hydromedusa blooms The most widespread and diverse representatives of gelatinous plankton Often neglected in plankton ecology ▪ be inconspicuous; escape direct observation ▪ seasonal occurrence; standard plankton sampling techniques easily overlook the blooms ▪ under-reporting (Hamner & Dawson 2009) Bloom examples and their impacts ▪ Predation of fish larvae and eggs by Aequorea victoria (Purcell 1989) ▪ Feeding of Moerisia lyonsi (introduced, Purcell et al 1999) ▪ Reviews on hydromedusae (Mills 2001) ▪ Blooms with upwelling events in Bay of Panama (Miglietta & Collin 2008) ▪ Collapse of zooplankton stocks during Liriope tetraphylla blooms (Yilmaz 2014) Feeding behavior ▪ Diversity of swimming behavior related to feeding and space utilization (Mills 1981) ▪ In situ feeding behavior of 8 co-occurring hydromedusae (Colin et al 2003) Predation of Nemopsis bachei in biological thin layers (Frost et al 2010) Invasions (and domination) ▪ Blackfordia virginica, Maeotias marginata, and Moerisia sp. (for example, Nowaczyk et al 2016 ) World distribution of Blackfordia 1 Mayer (1910) 2 Thiel (1935) 3 Richards (1938) 4 Logvinenko (1959) 5 Kramp (1961) 6 Cronin et al. (1962) 7 Chen-tsu and Chin (1962) 8 Naumov (1969) 9 Ferrante (1970) 10 Santhakumari and Dos santos(1970) 2,8,17,21,22,24 11 Denayer (1973) 11 12 Calder and Hester (1978) 13,32 4 17,18,23, 13 Moore (1987) 27,30,33 1,3,6,8,9,12 14 Bouillon et al(1988) 15 15 Dai et al. (1991) 34,36 7 22 16 Patrick (1994) 19 17 Mills and Sommer (1995) 12 5,13 This study 20 18 Cohen and Carlton (1995) 14 19 Santhakumari et al. (1997) 10 20 Alvarez (1999) 21 Ruiz et al. (2000) 22 Shardin (2000) 13,35 23 Mills and Rees (2000) 24 Gomiou et al. (2002) 16 25 Alvarez-silva s et al.(2003) 28,29,31 26 Buecher et al. (2005) 27 Wonham and Carlton (2005) Blackfordia virginica (Mayer,1910) 28 Genzano et al.(2006) 29 Nogueira and Oliveira (2006) Polyps were found 30 Schroeter (2008) Blackfordia polytentaculata (Hsu & Chin,1962) 31 Bardi and Marques (2009) 32 Chicharo et al. (2009) Blackfordia manhattensis (Mayer,1910) 33 Chang (2009) 34 Wintzer et al. (2011) Blackfordia sp. 35 Freire et al.(2013) 36 Wintzer et al. (2013) - Blackfordia virginica & B. manhattensis are identical (Moore 1987). - B. polytentaculata in South China Sea (Hsu & Chin 1962) was the first and last record of the species. - Broad thermal & salinity tolerances allow it to be easily transported in the ballast water (Kramp 1958); non-native species, marine pest (Molnar et al. 2008, Hayes & Sliwa 2003, Hayes et al. 2005, McDonald et al. 2015, Genzano et al. 2006 and etc.). - Low genetic diversity suggests that it is an introduced species (Harrison 2010, Genelle et al. 2012, Harrison et al. 2013, Meek et al. 2013). - Abnormally large impacts on zooplankton communities (Marques et al. 2015) Studies on Blackfordia virginica 1. Taxonomy ▪ morphological ▪ molecular 2. Population dynamics in Sihwa Lake ▪ Spatio-temporal variation of the density ▪ Individual growths (bell diameter) ▪ Optimal TS condition 3. Impact on zooplankton community (2018) ▪ In situ feeding rate (from gut contents in 24h) ▪ Stable isotope ratio analysis on zooplankton (prey) & Blackfordia virginica (predator) ▪ Phytoplankton composition in studied area ▪ Zooplankton composition in studied area ▪ Zooplankton & jellyfish carbon amount ▪ Bell diameter (bell shrinkage concerned to zooplankton (prey) depletion) 4. Is the species dispersed in other estuaries? ▪ Daecheon-cheon, Sumgingang, Kumgang, Nakdonggang and others in 2018 (?) Morphological taxonomy First record of Blackfordia virginica in Korean waters in 2013 Morphological taxonomy Sihwa 2013 Genzano et al. 2006 First record of Blackfordia virginica in Korean waters in 2013 Mills and Sommer 1995 Blackfordia virginica in Sihwa Lake Sihwa 2013 Molecular taxonomy 18S rDNA NJ tree Clustering in 100 HM194811-Chrysaora melanaster 28S rDNA of 28S rDNA AY039208-Aurelia aurita Out groups 93 EU27601-Aurelia sp.1 Blackfordia EU247811-Microhydrula limopsicola 98 FJ897543-Craspedacusta sowerbyi virginica100 AY920780-Chrysaora melanaster 96 AY920755-Limnocnida tanganyicae AY920754-Aglauropsis aeora EU27601-Aurelia sp 100 60 AY920752-Monobrachium parasiticum EU272545-Anthohebella parasitica 73 EU247814-Olindias sambaquiensis FJ550431-Hebella venusta 73 AY789778-Stegella lobata 98 99 AZ3-1Blackfordia virginica (Korean) AY920757-Melicertum octocostatum 66 AY789779-Billardia subrufa AY920800-Blackfordia virginica 27 FJ550535-Staurodiscus gotoi AY920799-Aequorea victoria 96 78 EU272619-Halecium muricatum 15 EU305528-Rhacostoma atlantica FJ550524-Symplectoscyphus turgidus 6 100 FJ550516-Nemertesia antennina 30 FJ550427-Eucheilota menoni 46 98 HM357627-Plumularia strictocarpa FJ550444-Eucheilota maculata Z92899Selaginopsis cornigera 98 55 EU305503-Thuiaria thuja 64 FJ550455-Eutima curva EU272600-Abietinaria filicula FJ550372-Calycella syringa 98 FJ550557-Diphasia fallax 51 35 FJ550451-Melicertum octocostatum 52 EU272602-Amphisbetia minima FJ550572-Sertularia moluccana FJ550403-Sertularella mediterranea 31 FJ550573-Cnidoscyphus marginatus 83 FJ550433-Halopteris carinata 96 FJ550575-Antennella secundaria 100 EU272583-Plumularia setacea 74 EU305497-Monostaechas quadridens 99 FJ550576-Halopteris carinata 74 FJ550450-Aglaophenia elongata FJ550581-Kirchenpaueria similis 92 53 28 FJ550442-Gymnangium gracilicaule 65 61 FJ550597-Cladocarpus integer 52 FJ550568-Hydrodendron mirabile FJ550453-Cladocarpus integer 0.01 92 EU272601-Aglaophenia tubiformis 77 FJ550429-Sertularia moluccana FJ550591-Lytocarpia sp. 65 FJ550418-Amphisbetia operculata FJ550586-Macrorhynchia sibogae 64 89 FJ550416-Hydrallmania falcata EU272603-Anthohebella parasitica 84 99 AZ3-1-Blackfordia virginica (Korean) 65 FJ550414-Diphasia fallax 99 AF358078-Blackfordia virginica EU305501-Rhacostoma atlantica 76 FJ550567-Hydranthea margarica FJ550587-Eucheilota maculata 80 FJ550600-Eutima gegenbauri FJ550594-Opercularella lacerata FJ550519-Calycella syringa 100% 18S rRNA FJ550596-Campanulina panicula 90 FJ550531-Tiaropsis multicirrata similarity 98 EU272611-Clytia noliformis FJ550589-Obelia bidentata 59 FJ550547-Gonothyraea loveni 88 FJ550590-Laomedea calceolifera AY789773-Lovenella gracilis AY789737-Orthopyxis integra FJ550549-Silicularia rosea 0.01 99 AY789740-Bonneviella regia 65 AY789739-Campanularia volubilis 60 AY789738-Rhizocaulus verticillatus Blackfordia virginica judged by DNA Morphology of polyps A A 0.5mm B Chae 2004 (in Sihwa) Mills & Rees 2000 Polyps of Blackfordia virginica. A: The polyps attached on an abandoned fish-net in Sihwa Lake in 2004. B: A polyp (& a young medusa) photograph form Mills and Rees (2000). Distribution of B. virginica in the East Asia sea ▪ Samples from Sihwa and Jiolong River showed 100 % similarity in ITS rDNA sequencing. ▪ Gene variation in rDNA IGS region, mtCO1, or 16S rDNA need to be examined (Genelle et al 2013). This study See Mayer (1910), Moore (2009), Toyokawa (2015) and etc. for morphological taxonomy. Toyokawa (2015) Hsu and Chin (1962) Occurrence in the East Asia region. Population dynamics of B. virginica in Sihwa Lake 2000 35 B.virginica 14000 Temperature(oC) ) 1500 ) -3 copepods 30 12000 -3 C) o 1000 10000 25 500 8000 (indiv. m (indiv. 200 20 6000 150 4000 Temperature ( 100 15 2000 copepodsm (indiv. B. virginicaB. 50 0 10 0 . .. 4 Jul 2 Oct 2 Jun 12 Jul 4 Sep 27 Apr 19 Jun 28 Jul 5 Sep 5 May 30 Jun 16 Jul 8 Sep 29 Jun 22 May 16 Aug 20 May 12 Aug 10 Aug 31 Mar 12 Sep 2013 2014 2015 2016 Density variation (2013 – 2016). 100 100 100 1400 80 4 Jul 80 20 May 80 2 Jun 1200 60 60 60 40 40 40 -3 1000 20 20 20 800 0 0 0 100 100 100 indiv.m 600 80 12 Jul 80 19 Jun 80 30 Jun 400 60 60 60 200 40 40 40 20 20 20 0 0 0 0 ~18 20 22 24 26 28 30 32 ~18 20 22 24 26 28 30 32 ~18 20 22 24 26 28 30 32 100 100 100 Salinity(PSU) 80 16 Aug 80 28 Jul 80 15 Jul 60 60 60 1400 40 40 40 1200 20 20 20 -3 Percent frequency Percent 1000 0 0 0 100 100 100 800 80 4 Sep 80 13 Aug 80 10 Aug indiv.m 600 60 60 60 40 40 40 400 20 20 20 200 0 0 0 0 <1 1 2 3 4 5 6 7 8 9 10 11 12 13 <1 1 2 3 4 5 6 7 8 9 10 11 12 13 <1 1 2 3 4 5 6 7 8 9 10 11 12 13 ~20 21 22 23 24 25 26 27 ~20 21 22 23 24 25 26 27 ~20 21 22 23 24 25 26 27 O Bell diameter(mm) Temperature( C) Salinity, temperature range and the jellyfish density (2013 – Bell diameter growth (2013 – 2015). 2015). Studies on Blackfordia virginica 1. Taxonomy ▪ morphological ▪ molecular 2. Population dynamics in Sihwa Lake ▪ Spatio-temporal variation of the density ▪ Individual growths (bell diameter) ▪ Optimal TS condition 3. Impact on zooplankton community (2018) ▪ In situ feeding rate (from gut contents in 24h) ▪ Stable isotope ratio analysis on zooplankton (prey) & Blackfordia virginica (predator) ▪ Phytoplankton composition in studied area ▪ Zooplankton composition in studied area ▪ Zooplankton & jellyfish carbon amount ▪ Bell diameter (bell shrinkage concerned to zooplankton (prey) depletion) 4. Is the species being dispersed in other estuaries? ▪ Daecheon-cheon, Sumgingang, Kumgang, Nakdonggang and others in 2018 (?) Other high density hydromedusae, Sarsia tubulosa Turritopsis nutricula, Rathkea octopunctata and Sarsia tubulosa Rathkea octopunctata © Hye Eun Lee © Jinho Chae Turritopsis dohrnii s.l. © Jinho Chae Molecular taxonomy of high density hydromedusae, Turritopsis nutricula, Rathkea octopunctata and Sarsia tubulosa (b) phylogeny using 18S rDNA of Turritopsis dohrnii (sys. T. nutricula?) phylogeny using 28S rDNA of Turritopsis dohrnii (sys. T. nutricula?) Extreme bloom-event of Rathkea octopunctata © Hye Eun Lee, NIFS Rathkea octopunctata Bloom event of R. octopunctata in Saemangum in Apr 2016. Extremely high density (c.a. 100 indiv. m-3 and 607.8 indiv.
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