Content uploaded by Amanda Foss
Author content
All content in this area was uploaded by Amanda Foss on May 31, 2020
Content may be subject to copyright.
Page 1 of 13
GreenWater Laboratories
Potentially Toxigenic (PTOX) Cyanobacteria List
Authors: Andrew Chapman & Amanda Foss
Updated: February 27, 2020
Document # 200227_PTOX
Some Known and Suspected Toxigenic Cyanobacteria from Around the World
This list includes taxa that have had toxins identified, have been implicated in toxic events in the
field, or have elicited a positive response in laboratory assays. New toxigenic cyanobacteria
species are continually being discovered and data concerning known and suspected toxin
producers continues to be refined and expanded. As a result, this list is by no means meant to be
an exhaustive list of toxin producing cyanobacteria and their toxins.
Page 2 of 13
Species
Cyanotoxin(s)
Produced
Reference
Anagnostidinema amphibium
(previously Geitlerinema
amphibium)
STXs – GTX4
(Borges et al., 2015) - Brazil
Anagnostidinema carotinosum
(previously Geitlerinema
carotinosum)
Microcystins (-LY)
Anatoxin-a
(Aboal, 2017) - Benthic
Anagnostidinema lemmermannii
(previously Geitlerinema
lemmermannii)
STXs – GTX1
(Borges et al., 2015) - Brazil
Anabaenopsis arnoldii
Microcystins
(-RR, -YR, unidentified
variants)
(Mohamed and Al Shehri, 2009)– Saudi Arabia
LC-UV & ELISA
Anabaenopsis milleri
Microcystin(s)
(Lanaras and Cook, 1994) – Extracted from bloom, confirmed
hepatotoxic via mouse bioassay, LC-UV max abs=238-240,
likely microcystin
Anabaena spp. WA102, AL93 &
37
(would be classified as
Dolichospermum due to gas
vesicle presence)
Anatoxin-a
(Brown et al., 2016) – strain WA102; confirmed using
molecular techniques & LC-MS/MS.
(Rantala-Ylinen et al., 2011) – strains 14, 37, 54, 86, 130
confirmed using molecular primers (anaC) & LC-MS/MS. All
described strains are planktonic.
Anabaena lapponica
Cylindrospermopsin
(Spoof et al., 2006)—Finland; CYN confirmed with LC-UV,
LC-MS/MS and TOF
Anabaena cylindrica
(previously A. subcylindrica)
Microcystins (-YR, -LR)
(Mohamed et al., 2006)—Saudi Arabia; benthic mats – isolates
measured using ELISA and LC-UV after mouse bioassay
confirmed hepatotoxicity
Aphanizomenon flos-aquae &
Aph. flos-aquae/klebahnii
Cylindrospermopsin
Anatoxin-a detected by UV/PDA (Rapala et al. (1993) but not
confirmed
Cylindrospermopsin production reported by Preußel et al.,
(2009, 2006) has been contested by Oregon D. Ag. due to AFA
harvesting. Saxitoxin production was originally thought to be
Aph. flos-aquae, but the organism was reclassified to Aph. gracile
(NH) or C. issatschenkoi (China) by W. Carmichael.
Aphanizomenon gracile
Cylindrospermopsin
STXs – GTX1,4,5, STX,
dcSTX, NEO, dcNEO
(Kokociński et al., 2013)—Poland CYN confirmed in isolates
using molecular & toxin analyses (ELISA, LC-UV & MS/MS)
(Pereira et al., 2004)—Portugal strain LMECYA40 isolated and
NEO & STX confirmed LC-FL (post);
US-MA lake sample dominated by Aph. gracile contained
GTX1,4, NEO & STX via LC-FL (pre)
Turkey Isolate (Yilmaz et al., 2018) had dcSTX. dcNEO, NEO,
STX, & 3 unknowns (MS/MS)
Aphanocapsa cumulus
Microcystins
(Domingos et al., 1999)– Brazil – ELISA used to screen, too low
(≤0.3 ng/mg) for LC-UV confirmation; an unidentified
spherical picoplankton was confirmed, possibly A. cumulus
Arthrospira fusiformis
(AB2002/10)
Anatoxin-a
Microcystin-YR
(Ballot et al., 2004)—Kenya isolate; MCs screened using ELISA
& confirmed with LC-UV; ANTX only LC-UV
Blennothrix lyngbyacea
(previously
Hydrocoleum lyngbyaceum)
Anatoxin-a
Homoanatoxin-a
(Méjean et al., 2010)- benthic marine associated with giant
clam poisoning event (ciguatoxins)
Page 3 of 13
Species
Cyanotoxin(s)
Produced
Reference
Chrysosporum bergii
(previously Anabaena bergii)
Cylindrospermopsin
Microcystins?
(Schembri et al., 2001)—Australia CYN production supported
with molecular work & LC-MS/MS
Anecdotal microcystin observation (by Adda ELISA) in bloom
dominated by C. bergii in a Texas Lake – Field Collection
Chrysosporum ovalisporum
(previously Aphanizomenon
ovalisporum)
Cylindrospermopsin
(Yilmaz et al., 2008)—US-FL isolates confirmed using LC-
MS/MS, ELISA and molecular work
(Akcaalan et al., 2014)—Turkey; blooms confirmed LC-MS/MS
(Banker et al., 1997)— Lake Kinneret, identified using UV, MS
& NMR
Cuspidothrix issatschenkoi
(previously Aphanizomenon
issatschenkoi)
Anatoxin-a
STXs – GTX5&6, STX,
NEO, dcSTX
(Wood et al., 2007)— New Zealand ANTX production
confirmed LC-MS/MS (CYN genes present, but no CYN
production)
(Ballot et al., 2010) – Germany ANTX by LC-MS/MS
(Li et al., 2003; Pereira et al., 2000)– reclassification from Aph.
flos-aquae to Aph. issatschenkoi); Both studies for STXs used LC-
FL & LC-MS
Cylindrospermum sp. (Finland)
Anatoxin-a
(Sivonen et al., 1989)
Cylindrospermum stagnale
STXs – NEO, dcSTX, STX,
GTX1
(Borges et al., 2015) - Brazil
Desmonostoc muscorum
(previously Nostoc muscorum)
Aplysiatoxins
Microcystins
(Mynderse et al., 1977)
(Oudra et al., 2009)—Morocco
Dolichospermum circinale
(previously Anabaena circinalis)
Anatoxin-a
Microcystins
STXs – STX, GTX235, C12,
dcSTX, dcGTX3
Unknown cytotoxin
(Beltran and Neilan, 2000; Froscio et al., 2011; Pereyra et al.,
2017; Vezie et al., 1998)
Dolichospermum crassum
(previously Anabaena crassa)
Anatoxin-a(s)
(Becker et al., 2010)—Brazil
Not certain if identification accurate
Dolichospermum flos-aquae
(previously Anabaena flos-
aquae)
Anatoxin-a
Anatoxin-a(s)
Microcystins
(Devlin and Edwards, 1977; Sivonen et al., 1989)
(Matsunaga et al., 1989)
(Harada et al., 1991)
Dolichospermum lemmermannii
(previously Anabaena
lemmermannii)
Anatoxin-a
Anatoxin-a(s)
EPI-CYN/CYN
Microcystins
STXs - STX
(Lepistö et al., 2005)
(Henriksen et al., 1997; Onodera et al., 1997)
(Onodera et al., 1997)—Denmark
Oregon Bloom of D. lemmermannii produced Epi-CYN (dom)
& CYN, re-occurs annually
(Rapala et al., 2005) – STX; not isolated strain
Dolichospermum macrosporum
(previously Anabaena
macrospora)
Anatoxin-a
(Park et al., 1993) - only LC-PDA used, not confirmed
Dolichospermum mendotae
(previously Anabaena mendotae)
Anatoxin-a
Cylindrospermopsin
(Rapala et al., 1993) - only LC-PDA used, not confirmed
(Akcaalan et al., 2014)- Turkey CYN
Dolichospermum planctonicum
(previously Anabaena
planctonica)
Anatoxin-a
(Bruno et al., 1994; Park et al., 1993)
Park et al. study only used LC-UV
Bruno et al. utilized GC-MS for derivatized ATX
(pentafluorobenzyl-ATX)
Dolichospermum spiroides
(previously Anabaena spiroides)
Anatoxin-a
Anatoxin-a(s)
(Park et al., 1993) (only LC-UV)
(Abreu and Ferrão-Filho, 2013) – Thought to be misidentified
in other manuscripts and may be Sphaerospermopsis torques-
reginae
Dolichospermum/
Microcystins
(Fewer et al., 2011)– Finland
Page 4 of 13
Species
Cyanotoxin(s)
Produced
Reference
Anabaena sp.
(Namikoshi et al., 1992b)
(Wang et al., 2012) - Anabaena sp. strain 90 genome, likely D.
circinale
Fischerella sp. strain CENA161
Microcystin-LR
(Fiore et al., 2009)—Brazil
Geitlerinema splendidum
(previously Phormidium
splendidum)
Microcystins
(-LF, -RR)
Pro-inflammatory & anti-
AChE substances
Anatoxin-a
(Aboal et al., 2005)
(Rangel et al., 2014)
(Aboal, 2017) - Benthic
Gloeotrichia echinulata
Microcystins
(Carey et al., 2007)—USA-NH; ELISA only, not confirmed
Gloeotrichia natans
Microcystins
(-RR & -LF)
(Aboal, 2017) - Benthic
Hapalosiphon hibernicus
BZ-3-1
Microcystin-LA
(Prinsep et al., 1992)
Soil sample – Hawaii (MS Thesis 2006 Philmus)
Hassallia sp.
antifungal
(Vestola et al., 2014)
Heteroscytonema cf. crispum
(previously Scytonema crispum)
(benthic)
STXs – GTX1,2,3,4,5
dcGTX23 dcSTX
(Smith et al., 2012)
Iningainema pulvinus
Nodularin
(McGregor and Sendall, 2017) - freshwater
Kamptonema formosum
(previously Phormidium
formosum)
Anatoxin-a,
Homoanatoxin-a
(Hemscheidt et al., 1995)
Leibleinia gracilis (previously
Phormidium gracile)
Hoiamide A,
Debromoaplysiatoxin?
(Pereira et al., 2009) – marine; associated with Moorena
producens
Leptolyngbya sp. CENA103 &
CENA112
Microcystin
(Furtado et al., 2009) – Brazil wastewater isolates, only
measured using ELISA (Beacon) at 0.14-0.31 ppb (ng/mL), no
confirmatory analysis
Leptolyngbya boryana
(previously Plectonema
boryanum)
Microcystins
(Mohamed et al., 2006) – ELISA, Bioassay & HPLC (-LR, -YR)
Limnothrix redekei
Microcystins
(Pineda-Mendoza et al., 2012)—Mexico - (low levels by ELISA
and not confirmed by other techniques
Limnothrix sp. CENA109 &
CENA110 (likely L. redekei
based on genetics)
Microcystin
(Furtado et al., 2009) – Brazil wastewater isolates, only
measured using ELISA (Beacon) at 0.19-0.42 ppb (ng/mL), no
confirmatory analysis
Lyngbya confervoides
Cytotoxins
Lyngbyapeptins
(Williams et al., 2002)—US
Merismopedia sp. CENA106 (M.
cf. tenuissima)
Microcystin
(Furtado et al., 2009) – Brazil wastewater isolate, only
measured using ELISA (Beacon) at 2.17 ppb (ng/mL), no
confirmatory analysis
Microcoleus autumnalis
(previously Phormidium
autumnale)
Anatoxin-a
Homoanatoxin-a
(Heath et al., 2010)
Microcystis aeruginosa
Microcystins
(Le Ai Nguyen et al., 2012)
Microcystis botrys
Microcystins
(Stefanelli et al., 2017)
Microcystis ichthyoblabe
Microcystins
(Sabour et al., 2002)
Microcystis panniformis
Microcystins
(Bittencourt-Oliveira et al., 2005)
Microcystis smithii
Microcystins
(Liu et al., 2011)- China
Microcystis viridis
Microcystins
(Kameyama et al., 2004; Song et al., 1998)
Page 5 of 13
Species
Cyanotoxin(s)
Produced
Reference
Microcystis wesenbergii
Microcystins
(Namikoshi et al., 1992a)
Microcystis sp. (Japan)
Anatoxin-a
(Park et al., 1993)-Only LC-UV used, highly suspect
Microcoleus autumnalis
(previously Phormidium
autumnale)
Anatoxin-a
Homoanatoxin-a
DhATX, DhHTX
Microcystin-LR
(Heath et al., 2014)
(Aboal, 2017) - Benthic
Microseira (Lyngbya) wollei
CYN
DeoxyCYN
STXs - dcSTX, dcGTX23,
LWT1-6
(Seifert et al., 2007)—Australia
(Onodera et al., 1997)—USA-AL
(Foss et al., 2012)—USA-FL
Moorena bouillonii (previously
Lyngbya)
Lyngbyapeptins
(Klein et al., 1999)—New Guinea
Moorena producens (previously
Lyngbya majuscula)
Aplysiatoxins
Lyngbyatoxins
Antillatoxin
Kalkitoxin
Jamaicamide
(Capper et al., 2005; Edwards et al., 2004; Liu and Rein, 2010;
Osborne et al., 2008, 2001; Taylor et al., 2014) - marine
Nodularia sphaerocarpa
Nodularin
(Beattie et al., 2000) - freshwater
Nodularia spumigena
Nodularins
(Mazur-Marzec et al., 2013) - marine
Nostoc sp. 5/96
Scytophycin
(Tomsickova et al., 2014)
Nostoc sp. 152
[DMAdda5] &
[ADMAdda5] MCs
(Sivonen et al., 1992)
Nostoc spp.
Microcystins
(Sivonen et al., 1990)
Nostoc cf. commune
Microcystins
(-LF, -LY)
STXs
(Teneva et al., 2012) - ?
(Aboal, 2017) - Benthic
Nostoc linckia
Microcystins
STXs
(Teneva et al., 2012) – ?
Nostoc paludosum
Microcystins
(La Claire and Manning, 2015)
Nostoc carneum (previously
Nostoc spongiaeforme)
Microcystins (-YR, -LR)
(Mohamed et al., 2006)—Saudi Arabia; benthic isolate
confirmed using mouse bioassay, ELISA & LC-UV
Oscillatoria limosa
Microcystins
(Mohamed, 2008)—Saudi Arabia
Oscillatoria margaritifera
Microcystins
(-RR, -LF, -LY)
(Aboal, 2017) - Benthic
Oscillatoria spp.
Anatoxin-a
(Cadel-Six et al., 2009)
Oscillatoria PCC 6506
Homoanatoxin-a,
Anatoxin-a (99:1)
7-epi-CYN & CYN
(Méjean et al., 2010)—FR: New Caledonia, (Mazmouz et al.,
2010)
Oscillatoria tenuis
Microcystins
(Brittain et al., 2000)—Egypt
oscillatorialean strains PCC
10601, PCC 10702, PCC 10608
(A, B, C in micrograph)
Homoanatoxin-a
Anatoxin-a
(Cadel-Six et al., 2007)
Phormidium spp.
Microcystin
(Izaguirre et al., 2007) – Analyzed by PP1A and HPLC-PDA
and/or MS indicated MC-LR was present
Potamolinea aerugineocaerulea
(previously Phormidium
aerugineo-caeruleum)
Cytotoxic
(Teneva et al., 2003)– freshwater Europe
Phormidium corium
Microcystin
(Mohamed et al., 2006)—Saudi Arabia
Page 6 of 13
Species
Cyanotoxin(s)
Produced
Reference
Phormidium favosum
Anatoxin-a
Homoanatoxin-a
(Gugger et al., 2005)—France (river)
Phormidium nigroviride
(previously Oscillatoria
nigroviridis)
Aplysiatoxins
(Mynderse et al., 1977) - Tropic/subtropic sea water
Phormidium willei
Microcystins
(Krienitz et al., 2003)
Phormidium uncinatum
STXs – GTX1
Microcystins
(-LF, -LY)
Anatoxin-a
(Borges et al., 2015)
(Aboal, 2017) - Benthic
Planktothrix agardhii
Anatoxin-a Microcystins
(Osswald et al., 2007)
(Luukkainen et al., 1993)
Planktothrix isothrix
Microcystins
(Bittencourt-Oliveira et al., 2014)
Planktothrix rubescens
Anatoxin-a
Microcystins (Hty
variants)
(Viaggiu et al., 2004)
(Ernst et al., 2006; Niedermeyer et al., 2014)
Planktothrix sp. FP1
STXs
(Pomati et al., 2000)—Italy- identification based on genetics
which may support novo status; pre- & post-column oxidation
LC-FL & LC-MS, no MSMS
Pseudanabaena galeata
Microcystins
(Oudra et al., 2002)– benthic – LC-UV/PDA & ELISA
Pseudanabaena mucicola
Microcystins
(Oudra et al., 2002) – planktonic - LC-UV/PDA &ELISA
Pseudanabaena limnetica
Microcystins
(Marsalek et al., 2003)—Czech Repub. – not unialgal – LC-
UV/PDA
Pseudocapsa dubia
Microcystins
(-RR, -YR)
(Cantoral Uriza et al., 2017) - Benthic
Radiocystis fernandoi
Microcystins
(Vieira et al., 2003)
Raphidiopsis curvata
Cylindrospermopsin/Deo
xyCYN
(Eaglesham et al., 2003)
Raphidiopsis brookii
STXs – GTX23 & STX
(Yunes et al., 2009)– Sub tropical
Raphidiopsis mediterranea
Anatoxin-a
Homoanatoxin-a
Cylindrospermopsin/Deo
xyCYN
(Hodoki et al., 2013)
(Watanabe et al., 2003)
(McGregor et al., 2011)
Raphidiopsis raciborskii
(previously Cylindrospermopsis
raciborskii)
Anatoxin-a
Homoanatoxin
Cylindrospermopsin
STXs - STX, NEO, GTX
2,3,56, dcNEO
(Vehovszky et al., 2009)— Hungary; only single quad MS used
after + neuronal effect (snail neurons) for ATX/HTX
(Schembri et al., 2001)—Australia – confirmed CYN in isolates
molecular & LC-MS/MS
(Lagos et al., 1999; Miotto et al., 2017)—Brazil – mouse
bioassay, LC-FL & MS on isolates
Rivularia biasolettiana
Microcystins
(-RR, -LR, -LY)
(Aboal et al., 2005)
(Cantoral Uriza et al., 2017)
Rivularia haematites
Microcystin
(Aboal et al., 2005)
Romeria caruaru
Microcystin
(Komárek et al., 2001)
Schizothrix calcicola
Aplysiatoxins
(Mynderse et al., 1977)
Scytonematopsis crustacea
(previously Calothrix crustacea)
Aplysiatoxin
(Mynderse et al., 1977)
Scytonema drilosiphon
Microcystins
(-LY)
(Cantoral Uriza et al., 2017) - Benthic
Page 7 of 13
Species
Cyanotoxin(s)
Produced
Reference
Scytonema ocellatum
Scytophycin
(antineoplastic &
antifungal)
(Ishibashi et al., 1986)
Scytonema pseudohofmannii
Scytophycin
(antineoplastic &
antifungal)
(Ishibashi et al., 1986)
Snowella lacustris
Microcystin
(Sant’Anna and Azevedo, 2000)—Brazil
Sphaerospermopsis
aphanizomenoides (previously
Anabaena and Aphanizomenon)
Anatoxin-a?
Cylindrospermopsin
STXs
Microcystins?
(Bittencourt-Oliveira et al., 2011) – potential CYN & STX
producer – Brazil
(Sabour et al., 2005) – MC production questioned by (Cirés
and Ballot, 2016)
Sphaerospermopsis torques-
reginae (previously Anabaena)
Anatoxin-a(s)
(Dörr et al., 2010) – Originally published as T? Anabaena
spiroides, Anabaena oumiana, changed to Sphaerospermopsis
oumiana and is now Sphaerospermopsis torques-reginae
Stenomitos frigidus (previously
Pseudanabaena frigida)
Microcystin-RR
(Aboal, 2017) - Benthic
Symploca muscorum
Aplysiatoxin
(Mynderse et al., 1977)
Synechococcus sp.
Microcystins
(Carmichael and Li, 2006) - marine
Synechococcus sp. CENA108
Microcystin
(Furtado et al., 2009) – Brazil wastewater isolate, only
measured using ELISA (Beacon) at 0.22 ppb (ng/mL), no
confirmatory analysis
Synechocystis sp.
Microcystins
(Oudra et al., 2002) UV PDA only
Synechocystis sp. PCC 6803
Hemolysins
(Bi et al., 2011)
Synechocystis aquatilis
Microcystin
(Domingos et al., 1999) – Brazil, brackish lagoon isolate
Tolypothrix conglutinata
Scytophycin
(antineoplastic &
antifungal)
(Ishibashi et al., 1986)
Tolypothrix distorta
Microcystins
(Aboal et al., 2005)
Trichodesmium erythraeum
Microcystins
Aplysiatoxins (5 variants)
(Gupta et al., 2014; Ramos et al., 2005)– brackish, marine
Trichodesmium thiebautii
Trichotoxin 2 - neurotoxin
(Schock et al., 2011) – brackish, marine
Trichormus variabilis
(previously Anabaena)
Microcystin (-YR, -LR)
(Mohamed et al., 2006)—Saudi Arabia, benthic mat isolate
toxic via mouse bioassay, MCs by ELISA and LC-UV
Tychonema bourrellyi
Anatoxin-a
(Shams et al., 2015)
Umezakia natans
Cylindrospermopsin
(Harada et al., 1994)
Woronichinia naegeliana
Microcystin
(Bober et al., 2011) – trace MC-LR
Page 8 of 13
References:
Aboal, M., Puig, M.Á., Asencio, A.D., 2005. Production of microcystins in calcareous Mediterranean streams: The Alharabe River,
Segura River basin in south-east Spain. J. Appl. Phycol. 17, 231–243.
Abreu, F.Q. de, Ferrão-Filho, A. da S., 2013. Effects of an Anatoxin-a(s)-Producing Strain of Anabaena spiroides (Cyanobacteria) on the
Survivorship and Somatic Growth of Two Daphnia similis Clones. J. Environ. Prot. (Irvine,. Calif). 04, 12–18.
Akcaalan, R., Köker, L., Oğuz, A., Spoof, L., Meriluoto, J., Albay, M., 2014. First report of cylindrospermopsin production by two
cyanobacteria (Dolichospermum mendotae and Chrysosporum ovalisporum ) in Lake Iznik, Turkey. Toxins (Basel). 6, 3173–3186.
Ballot, A., Fastner, J., Lentz, M., Wiedner, C., 2010. First report of anatoxin-a-producing cyanobacterium Aphanizomenon issatschenkoi in
northeastern Germany. Toxicon 56, 964–971.
Ballot, A., Krienitz, L., Kotut, K., Wiegand, C., Metcalf, J.S., Codd, G. a., Pflugmacher, S., 2004. Cyanobacteria and cyanobacterial toxins
in three alkaline Rift Valley lakes of Kenya - Lakes Bogoria, Nakuru and Elmenteita. J. Plankton Res. 26, 925–935.
Banker, R., Carmeli, S., Hadas, O., Teltsch, B., Porat, R., Sukenik, A., 1997. Identification of Cylindrospermopsin in Aphanizomenon
Ovalisporum (Cyanophyceae) Isolated From Lake Kinneret, Israel1. J. Phycol. 33, 613–616.
Beattie, K.A., Kaya, K., Codd, G.A., 2000. The cyanobacterium Nodularia PCC 7804, of freshwater origin, produces [L-Har2]nodularin.
Phytochemistry 54, 57–61.
Becker, V., Ihara, P., Yunes, J.S., Huszar, V.L.M., 2010. Occurrence of anatoxin-a(s) during a bloom of Anabaena crassa in a water-supply
reservoir in southern Brazil. J. Appl. Phycol. 22, 235–241.
Beltran, E.C., Neilan, B. a., 2000. Geographical segregation of the neurotoxin-producing cyanobacterium Anabaena circinalis. Appl.
Environ. Microbiol. 66, 4468–4474.
Bi, S., Wang, W., Zhao, Y., Ru, S., Liu, Y., 2011. Studies on hemolysis of hemolysin produced by Synechocystis sp. PCC 6803. J. Ocean
Univ. China 10, 362–368.
Bittencourt-Oliveira, M. do C., Piccin-Santos, V., Kujbida, P., Moura, A. do N., 2011. Cylindrospermopsin in Water Supply Reservoirs in
Brazil Determined by Immunochemical and Molecular Methods. J. Water Resour. Prot. 03, 349–355.
Bittencourt-Oliveira, M.D.C., Kujbida, P., Cardozo, K.H.M., Carvalho, V.M., Moura, A.D.N., Colepicolo, P., Pinto, E., 2005. A novel
rhythm of microcystin biosynthesis is described in the cyanobacterium Microcystis panniformis Komarek et al. Biochem. Biophys.
Res. Commun. 326, 687–694.
Bittencourt-Oliveira, M.D.C., Piccin-Santos, V., Moura, A.N., Aragão-Tavares, N.K.C., Cordeiro-Araújo, M.K., 2014. Cyanobacteria,
microcystins and cylindrospermopsin in public drinking supply reservoirs of Brazil. An. Acad. Bras. Cienc. 86, 297–309.
Bober, B., Lechowski, Z., Bialczyk, J., 2011. Determination of some cyanopeptides synthesized by Woronichinia naegeliana
(Chroococcales, Cyanophyceae). Phycol. Res. 59, 286–294.
Borges, H.L.F., Branco, L.H.Z., Martins, M.D., Lima, C.S., Barbosa, P.T., Lira, G.A.S.T., Bittencourt-Oliveira, M.C., Molica, R.J.R., 2015.
Cyanotoxin production and phylogeny of benthic cyanobacterial strains isolated from the northeast of Brazil. Harmful Algae 43,
46–57.
Brittain, S., Mohamed, Z.A., Wang, J., Lehmann, V.K.B., Carmichael, W.W., Rinehart, K.L., 2000. Isolation and characterization of
microcystins from a River Nile strain of Oscillatoria tenuis Agardh ex Gomont. Toxicon 38, 1759–1771.
Brown, N.M., Mueller, R.S., Shepardson, J.W., Landry, Z.C., Morré, J.T., Maier, C.S., Hardy, F.J., Dreher, T.W., 2016. Structural and
functional analysis of the finished genome of the recently isolated toxic Anabaena sp. WA102. BMC Genomics 17, 457.
Bruno, M., Barbini, D.A., Pierdominici, E., Serse, A.P., Ioppolo, A., 1994. Anatoxin-a and a previously unknown toxin in Anabaena
planctonica from blooms found in Lake Mulargia (Italy). Toxicon 32, 369–373.
Cadel-Six, S., Iteman, I., Peyraud-Thomas, C., Mann, S., Ploux, O., Méjean, A., 2009. Identification of a polyketide synthase coding
sequence specific for anatoxin-a-producing Oscillatoria cyanobacteria. Appl. Environ. Microbiol. 75, 4909–4912.
Cadel-Six, S., Peyraud-Thomas, C., Brient, L., De Marsac, N.T., Rippka, R., Méjean, A., 2007. Different genotypes of anatoxin-producing
cyanobacteria coexist in the Tarn River, France. Appl. Environ. Microbiol. 73, 7605–7614.
Cantoral Uriza, E.A., Asencio, A.D., Aboal, M., 2017. Are we underestimating benthic cyanotoxins? extensive sampling results from
Spain. Toxins (Basel). 9.
Page 9 of 13
Capper, A., Tibbetts, I.R., O'Neil, J.M., Shaw, G.R., 2005. The fate of Lyngbya majuscula toxins in three potential consumers. J.
Chem. Ecol. 31, 1595–1606.
Carey, C.C., Haney, J.F., Cottingham, K.L., 2007. First report of microcystin-LR in the cyanobacterium Gloeotrichia echinulata. Environ.
Toxicol. 22, 337–339.
Carmichael, W.W., Li, R., 2006. Cyanobacteria toxins in the Salton Sea. Saline Systems 2, 5.
Cirés, S., Ballot, A., 2016. A review of the phylogeny, ecology and toxin production of bloom-forming Aphanizomenon spp. and related
species within the Nostocales (cyanobacteria). Harmful Algae.
Devlin, J., Edwards, O., 1977. Anatoxin-a, a toxic alkaloid from Anabaena flos-aquae NRC-44h. Can. J. Chem. 1–5.
Domingos, P., Rubim, T.K., Molica, R.J.R., Azevedo, S.M.F.O., Carmichael, W.W., 1999. First report of microcystin production by
picoplanktonic cyanobacteria isolated from a northeast Brazilian drinking water supply. Environ. Toxicol. 14, 31–35.
Dörr, F.A., Rodríguez, V., Molica, R., Henriksen, P., Krock, B., Pinto, E., 2010. Methods for detection of anatoxin-a(s) by liquid
chromatography coupled to electrospray ionization-tandem mass spectrometry. Toxicon 55, 92–99.
Eaglesham, G.K., Li, R., Watanabe, M.M., Brittain, S., Liu, Y., Carmichael, W.W., Shaw, G.R., 2003. First report of the cyanotoxins
cylindrospermopsin and dexycylindrospermopsin from Raphidiopsis curvata (Cyanobacteria). J. Phycol. 37, 1121–1126.
Edwards, D.J., Marquez, B.L., Nogle, L.M., McPhail, K., Goeger, D.E., Roberts, M.A., Gerwick, W.H., 2004. Structure and Biosynthesis of
the Jamaicamides, New Mixed Polyketide-Peptide Neurotoxins from the Marine Cyanobacterium Lyngbya majuscula. Chem. Biol.
11, 817–833.
Ernst, B., Hoeger, S.J., O’Brien, E., Dietrich, D.R., 2006. Oral toxicity of the microcystin-containing cyanobacterium Planktothrix rubescens
in European whitefish (Coregonus lavaretus). Aquat. Toxicol. 79, 31–40.
Fewer, D.P., Halinen, K., Sipari, H., Bernardová, K., Mänttäri, M., Eronen, E., Sivonen, K., 2011. Non-autonomous transposable
elements associated with inactivation of microcystin gene clusters in strains of the genus Anabaena isolated from the Baltic Sea.
Environ. Microbiol. Rep. 3, 189–194.
Fiore, M.F., Genuário, D.B., da Silva, C.S.P., Shishido, T.K., Moraes, L.A.B., Neto, R.C., Silva-Stenico, M.E., 2009. Microcystin production
by a freshwater spring cyanobacterium of the genus Fischerella. Toxicon 53, 754–761.
Foss, A., Phlips, E., Aubel, M., Szabo, N., 2012. Investigation of extraction and analysis techniques for Lyngbya wollei derived Paralytic
Shellfish Toxins. Toxicon 60, 1148–1158.
Froscio, S., Sieburn, K., Lau, H.M., Humpage, A., 2011. Novel cytotoxicity associated with Anabaena circinalis 131C. Toxicon 58, 689–692.
Furtado, A.L.F.F., Calijuri, M.D.C., Lorenzi, A.S., Honda, R.Y., Genuário, D.B., Fiore, M.F., 2009. Morphological and molecular
characterization of cyanobacteria from a Brazilian facultative wastewater stabilization pond and evaluation of microcystin
production. Hydrobiologia 627, 195–209.
Gugger, M., Lenoir, S., Berger, C., Ledreux, A., Druart, J.-C., Humbert, J.-F., Guette, C., Bernard, C., 2005. First report in a river in
France of the benthic cyanobacterium Phormidium favosum producing anatoxin-a associated with dog neurotoxicosis. Toxicon 45,
919–928.
Gupta, D., Kaur, P., Leong, S., Tan, L., Prinsep, M., Chu, J., 2014. Anti-Chikungunya Viral Activities of Aplysiatoxin-Related
Compounds from the Marine Cyanobacterium Trichodesmium erythraeum. Mar. Drugs 12, 115–127.
Harada, K., Ohtani, I., Iwamoto, K., Suzuki, M., Watanabe, M.F., Watanabe, M., Terao, K., 1994. Isolation of cylindrospermopsin from a
cyanobacterium Umezakia natans and its screening method. Toxicon 32, 73–84.
Harada, K.I., Ogawa, K., Kimura, Y., Murata, H., Suzuki, M., Thorn, P.M., Evans, W.R., Carmichael, W.W., 1991. Microcystins from
Anabaena flos-aquae NRC 525-17. Chem. Res. Toxicol. 4, 535–540.
Heath, M.W., Wood, S.A., Barbieri, R.F., Young, R.G., Ryan, K.G., 2014. Effects of nitrogen and phosphorus on anatoxin-a,
homoanatoxin-a, dihydroanatoxin-a and dihydrohomoanatoxin-a production by Phormidium autumnale. Toxicon.
Heath, M.W., Wood, S.A., Ryan, K.G., 2010. Polyphasic assessment of fresh-water benthic mat-forming cyanobacteria isolated from
New Zealand. FEMS Microbiol. Ecol. 73, 95–109.
Hemscheidt, T., Rapala, J., Sivonen, K., Skulberg, O.M., 1995. Biosynthesis os anatoxin-a in Anabaena flos-aquae and homoanatoxin-a in
Oscillatoria formosa. J. Chem. Soc. 13, 1361–1362.
Henriksen, P., Carmichael, W.W., Jisi, A., Moestrup, Ø., 1997. Detection of an anatoxin-a(s)-like anticholinesterase in natural blooms
Page 10 of 13
and cultures of cyanobacteria/blue-green algae from Danish lakes and in the stomach content of poisoned birds. Toxicon 35, 901–
913.
Hodoki, Y., Ohbayashi, K., Kobayashi, Y., Takasu, H., Okuda, N., Nakano, S., 2013. Anatoxin-a-producing Raphidiopsis mediterranea
Skuja var. grandis Hill is one ecotype of non-heterocytous Cuspidothrix issatschenkoi (Usačev) Rajaniemi et al. in Japanese lakes.
Harmful Algae 21, 44–53.
Ishibashi, M., Moore, R.E., Patterson, G.M.L., Xu, C., Clardy, J., 1986. Scytophycins, cytotoxic and antimycotic agents from the
cyanophyte Scytonema pseudohofmanni. J. Org. Chem. 51, 5300–5306.
Izaguirre, G., Jungblut, A.-D.D., Neilan, B.A., 2007. Benthic cyanobacteria (Oscillatoriaceae) that produce microcystin-LR, isolated from
four reservoirs in southern California. Water Res. 41, 492–498.
Kameyama, K., Sugiura, N., Inamori, Y., Maekawa, T., 2004. Characteristics of microcystin production in the cell cycle of Microcystis
viridis. Environ. Toxicol. 19, 20–25.
Klein, D., Braekman, J.-C., Daloze, D., Hoffmann, L., Castillo, G., Demoulin, V., 1999. Lyngbyapeptin A, a modified tetrapeptide from
Lyngbya bouillonii (Cyanophyceae). Tetrahedron Lett. 40, 695–696.
Kokociński, M., Mankiewicz-Boczek, J., Jurczak, T., Spoof, L., Meriluoto, J., Rejmonczyk, E., Hautala, H., Vehniäinen, M., Pawełczyk, J.,
Soininen, J., 2013. Aphanizomenon gracile (Nostocales), a cylindrospermopsin-producing cyanobacterium in Polish lakes. Environ.
Sci. Pollut. Res. 20, 5243–5264.
Komárek, J., Azevedo, M.T.D.P., Domingos, P., Komárková, J., Tichý, M., 2001. Background of the Caruaru tragedy: a case taxonomic
study of toxic cyanobacteria. Arch. Hydrobiol. Suppl. Algol. Stud. 103, 9–29.
Krienitz, L., Ballot, A., Kotut, K., Wiegand, C., Pütz, S., Metcalf, J.S., Codd, G.A., Pflugmacher, S., 2003. Contribution of hot spring
cyanobacteria to the mysterious deaths of Lesser Flamingos at Lake Bogoria, Kenya. FEMS Microbiol. Ecol. 43, 141–148.
La Claire, J.W., Manning, S.R., 2015. Ichthyotoxins, Phycotoxins.
Lagos, N., Onodera, H., Zagatto, P.A., Andrinolo, D., Azevedo, S.M.F.Q., Oshima, Y., 1999. The first evidence of paralytic shellfish
toxins in the freshwater cyanobacterium Cylindrospermopsis raciborskii, isolated from Brazil. Toxicon 37, 1359–1373.
Lanaras, T., Cook, C.M., 1994. Toxin extraction from an Anabaenopsis milleri - Dominated bloom. Sci. Total Environ. 142, 163–169.
Le Ai Nguyen, V., Tanabe, Y., Matsuura, H., Kaya, K., Watanabe, M.M., 2012. Morphological, biochemical and phylogenetic
assessments of water-bloom-forming tropical morphospecies of Microcystis (Chroococcales, Cyanobacteria). Phycol. Res. 60, 208–
222.
Lepistö, L., Rapala, J., Lyra, C., Berg, K.A., Erkomaa, K., Issakainen, J., 2005. Occurrence and toxicity of cyanobacterial blooms
dominated by Anabaena lemmermannii P. Richter and Aphanizomenon spp. in boreal lakes in 2003. Arch. Hydrobiol. Suppl. Algol.
Stud. 117, 315–328.
Li, R., Carmichael, W.W., Pereira, P., 2003. Morphological and 16s rRNA gene evidence for reclassification of the paralytic shellfish
toxin producing Aphanizomenon flos-aquae LMECYA 31 as Aphanizomenon issatschenkoi (Cyanophyceae). J. Phycol. 39, 814–818.
Liu, L., Rein, K.S., 2010. New peptides isolated from Lyngbya species: A review. Mar. Drugs.
Liu, Y., Tan, W., Wu, X., Wu, Z., Yu, G., Li, R., 2011. First report of microcystin production in Microcystis smithii Komorek and
Anagnostidis (Cyanobacteria) from a water bloom in Eastern China. J. Environ. Sci. 23, 102–107.
Luukkainen, R., Sivonen, K., Namikoshi, M., Fardig, M., Rinehart, K.L., Niemela, S.I., 1993. Isolation and identification of eight
microcystins from thirteen Oscillatoria agardhii strains and structure of a new microcystin. Appl. Environ. Microbiol. 59, 2204–
2209.
Marsalek, B., Blaha, L., Babica, P., 2003. Analyses of microcystins in the biomass of Pseudanabaena limnetica collected in Znojmo
reservoir. Czech Phycol. 3, 195–197.
Matsunaga, S., Moore, R.E., Niemczura, W.P., Carmichael, W.W., 1989. Anatoxin-a(s), a potent anticholinesterase from Anabaena flos-
aquae. J. Am. Chem. Soc. 111, 8021–8023.
Mazmouz, R., Chapuis-Hugon, F., Mann, S., Pichon, V., Méjean, A., Ploux, O., 2010. Biosynthesis of cylindrospermopsin and 7-
epicylindrospermopsin in Oscillatoria sp. strain PCC 6506: Identification of the cyr gene cluster and toxin analysis. Appl. Environ.
Microbiol. 76, 4943–4949.
Mazur-Marzec, H., Kaczkowska, M.J., Blaszczyk, A., Akcaalan, R., Spoof, L., Meriluoto, J., 2013. Diversity of peptides produced by
Page 11 of 13
Nodularia spumigena from various geographical regions. Mar. Drugs 11, 1–19.
McGregor, G., Sendall, B., Hunt, L., Eaglesham, G., 2011. Report of the cyanotoxins cylindrospermopsin and deoxy-cylindrospermopsin
from Raphidiopsis mediterranea Skuja (Cyanobacteria/Nostocales). Harmful Algae.
McGregor, G.B., Sendall, B.C., 2017. Iningainema pulvinus gen nov., sp nov. (Cyanobacteria, Scytonemataceae) a new nodularin producer
from Edgbaston Reserve, north-eastern Australia. Harmful Algae 62, 10–19.
Méjean, A., Peyraud-Thomas, C., Kerbrat, A.S., Golubic, S., Pauillac, S., Chinain, M., Laurent, D., 2010. First identification of the
neurotoxin homoanatoxin-a from mats of Hydrocoleum lyngbyaceum (marine cyanobacterium) possibly linked to giant clam
poisoning in New Caledonia. Toxicon 56, 829–835.
Miotto, M.C., Costa, L.D.F., Brentano, D.M., Nader, C., Dos Santos Souza, L., Gressler, P.D., Laudares-Silva, R., Yunes, J.S., Barufi, J.B.,
Rörig, L.R., 2017. Ecophysiological characterization and toxin profile of two strains of Cylindrospermopsis raciborskii isolated from a
subtropical lagoon in Southern Brazil. Hydrobiologia 1–17.
Mohamed, Z. a., Al Shehri, A.M., 2009. Microcystin-producing blooms of Anabaenopsis arnoldi in a potable mountain lake in Saudi
Arabia: Research article. FEMS Microbiol. Ecol. 69, 98–105.
Mohamed, Z.A., 2008. Toxic cyanobacteria and cyanotoxins in public hot springs in Saudi Arabia. Toxicon 51, 17–27.
Mohamed, Z.A., El-Sharouny, H.M., Ali, W.S.M., 2006. Microcystin production in benthic mats of cyanobacteria in the Nile River and
irrigation canals, Egypt. Toxicon 47, 584–590.
Mynderse, J., Moore, R., Kashiwagi, M., Norton, T., 1977. Antileukemia activity in the Osillatoriaceae: isolation of Debromoaplysiatoxin
from Lyngbya. Science (80-. ). 196.
Namikoshi, M., Kenneth, L., Sakai, R., Stotts, R.R., Dahlem, A.M., Beasley, V.R., Carmichael, W.W., Evans, W.R., 1992a. Identification of
12 hepatotoxins from a Homer Lake bloom of the cyanobacteria Microcystis aeruginosa, Microcystis viridis, and Microcystis
wesenbergii: nine new microcystins. J. Or 57, 866–872.
Namikoshi, M., Sivonen, K., Evans, W.R., Carmichael, W.W., Sun, F., Rouhiainen, L., Luukkainen, R., Rinehart, K.L., 1992b. Two new L-
serine variants of microcystins-LR and -RR from Anabaena sp. strains 202 A1 and 202 A2. Toxicon 30, 1457–1464.
Niedermeyer, T.H.J., Schmieder, P., Kurmayer, R., 2014. Isolation of microcystins from the cyanobacterium Planktothrix rubescens strain
No80. Nat. Products Bioprospect. 4, 37–45.
Onodera, H., Oshima, Y., Henriksen, P., Yasumoto, T., 1997. Confirmation of anatoxin-a(s), in the cyanobacterium Anabaena
lemmermannii, as the cause of bird kills in Danish lakes. Toxicon 35, 1645–1648.
Osborne, N., Seawright, A., Shaw, G., 2008. Dermal toxicology of Lyngbya majuscula, from Moreton Bay, Queensland, Australia.
Harmful Algae 7, 584–589.
Osborne, N.J.T., Webb, P.M., Shaw, G.R., 2001. The toxins of Lyngbya majuscula and their human and ecological health effects. Environ.
Int. 27, 381–392.
Osswald, J., Rellán, S., Gago, A., Vasconcelos, V., 2007. Toxicology and detection methods of the alkaloid neurotoxin produced by
cyanobacteria, anatoxin-a. Environ. Int. 33, 1070–1089.
Oudra, B., Dadi-El Andaloussi, M., Vasconcelos, V.M., 2009. Identification and quantification of microcystins from a Nostoc muscorum
bloom occurring in Oukaïmeden River (High-Atlas mountains of Marrakech, Morocco). Environ. Monit. Assess. 149, 437–444.
Oudra, B., Loudiki, M., Vasconcelos, V., Sabour, B., Sbiyyaa, B., Oufdou, K., Mezrioui, N., 2002. Detection and quantification of
microcystins from cyanobacteria strains isolated from reservoirs and ponds in Morocco. Environ. Toxicol. 17, 32–39.
Park, H.-D., Watanabe, M.F., Harada, K.-I., Nagai, H., Suzuki, M., Watanabe, M., Hayashi, H., 1993. Hepatotoxin (microcystin) and
neurotoxin (anatoxin-a) contained in natural blooms and strains of cyanobacteria from Japanese freshwaters. Nat. Toxins 1, 353–
360.
Pereira, A., Cao, Z., Murray, T.F., Gerwick, W.H., 2009. Hoiamide A, a Sodium Channel Activator of Unusual Architecture from a
Consortium of Two Papua New Guinea Cyanobacteria. Chem. Biol. 16, 893–906.
Pereira, P., Li, R., Carmichael, W.W., Dias, E., Franca, S., 2004. Taxonomy and production of paralytic shellfish toxins by the freshwater
cyanobacterium Aphanizomenon gracile LMECYA40. Eur. J. Phycol. 39, 361–368.
Pereira, P., Onodera, H., Andrinolo, D., Franca, S., Ara??jo, F., Lagos, N., Oshima, Y., 2000. Paralytic shellfish toxins in the freshwater
cyanobacterium Aphanizomenon flos-aquae, isolated from Montargil reservoir, Portugal. Toxicon 38, 1689–1702.
Page 12 of 13
Pereyra, J.P.A., D’Agostino, P.M., Mazmouz, R., Woodhouse, J.N., Pickford, R., Jameson, I., Neilan, B.A., 2017. Molecular and
morphological survey of saxitoxin-producing cyanobacterium Dolichospermum circinale (Anabaena circinalis) isolated from
geographically distinct regions of Australia. Toxicon 138, 68–77.
Pineda-Mendoza, R.M., Olvera-Ramírez, R., Martínez-Jerónimo, F., 2012. Microcystins produced by filamentous cyanobacteria in urban
lakes. A case study in Mexico City. Hidrobiologica 22, 290–298.
Pomati, F., Sacchi, S., Rossetti, C., Giovannardi, S., Onodera, H., Oshima, Y., Neilan, B.A., 2000. The freshwater cyanobacterium
Planktothrix sp. FP1 : Molecular identification and detection of paralytic shellfish toxins. J. Phycol. 36, 553–562.
Preußel, K., Stüken, A., Wiedner, C., Chorus, I., Fastner, J., 2006. First report on cylindrospermopsin producing Aphanizomenon flos-
aquae (Cyanobacteria) isolated from two German lakes. Toxicon 47, 156–162.
Preußel, K., Wessel, G., Fastner, J., Chorus, I., 2009. Response of cylindrospermopsin production and release in Aphanizomenon flos-aquae
(Cyanobacteria) to varying light and temperature conditions. Harmful Algae 8, 645–650.
Prinsep, M.R., Caplan, F.R., Moore, R.E., Patterson, G.M.L., E. Honkanen, R., Boynton, A.L., 1992. Microcystin-LA from a blue-green
alga belonging to the Stigonematales. Phytochemistry 31, 1247–1248.
Ramos, A.G., Martel, A., Codd, G.A., Soler, E., Coca, J., Redondo, A., Morrison, L.F., Metcalf, J.S., Ojeda, A., Suárez, S., Petit, M., 2005.
Bloom of the marine diazotrophic cyanobacterium Trichodesmium erythraeum in the Northwest African Upwelling. Mar. Ecol.
Prog. Ser. 301, 303–305.
Rangel, M., Martins, J.C.G., Garcia, A.N., Conserva, G.A.A., Costa-Neves, A., Sant’Anna, C.L., De Carvalho, L.R., 2014. Analysis of the
toxicity and histopathology induced by the oral administration of Pseudanabaena galeata and Geitlerinema splendidum
(Cyanobacteria) extracts to mice. Mar. Drugs 12, 508–524.
Rantala-Ylinen, A., Känä, S., Wang, H., Rouhiainen, L., Wahlsten, M., Rizzi, E., Berg, K., Gugger, M., Sivonen, K., 2011. Anatoxin-a
synthetase gene cluster of the cyanobacterium Anabaena sp. strain 37 and molecular methods to detect potential producers. Appl.
Environ. Microbiol. 77, 7271–7278.
Rapala, J., Robertson, A., Negri, A.P., Berg, K.A., Tuomi, P., Lyra, C., Erkomaa, K., Lahti, K., Hoppu, K., Lepistö, L., 2005. First report of
saxitoxin in Finnish lakes and possible associated effects on human health, in: Environmental Toxicology. pp. 331–340.
Rapala, J., Sivonen, K., Luukkainen, R., Niemela, S.I., 1993. Anatoxin-a concentration in Anabaena and Aphanizomenon under different
environmental conditions and comparison of growth by toxic and non-toxic Anabaena-strains - a laboratory study. Appl. Phycol.
5, 581–591.
Sabour, B., Loudiki, M., Oudra, B., Oubraim, S., Fawzi, B., Fadlaoui, S., Chlaida, M., Vasconcelos, V., 2002. First results on Microcystis
ichthyoblabe Kutz. toxic bloom in the hypertrophic Oued Mellah reservoir (Morocco). Ann. Limnol. J. Limnol.
Sabour, B., Loudiki, M., Oudra, B., Vasconcelos, V., Oubraim, S., Fawzi, B., 2005. Dynamics and toxicity of Anabaena aphanizomenoides
(Cyanobacteria) waterblooms in the shallow brackish Oued Mellah lake (Morocco). Aquat. Ecosyst. Heal. Manag. 8, 95–104.
Sant’Anna, C.L., Azevedo, M.T. De, 2000. Contribution to the knowledge of potentially toxic Cyanobacteria from Brazil. Nov. Hedwigia
71, 359–385.
Schembri, M. a, Neilan, B. a, Saint, C.P., Christopher, P., Saint, C.P., 2001. Identification of genes implicated in toxin production in the
cyanobacterium Cylindrospermopsis raciborskii. Environ. Toxicol. 16, 413–421.
Schock, T.B., Huncik, K., Beauchesne, K.R., Villareal, T.A., Moeller, P.D.R., 2011. Identification of Trichotoxin, a Novel Chlorinated
Compound Associated with the Bloom Forming Cyanobacterium, Trichodesmium thiebautii. Environ. Sci. Technol. 45, 7503–7509.
Seifert, M., McGregor, G., Eaglesham, G., Wickramasinghe, W., Shaw, G., 2007. First evidence for the production of cylindrospermopsin
and deoxy-cylindrospermopsin by the freshwater benthic cyanobacterium, Lyngbya wollei (Farlow ex Gomont) Speziale and Dyck.
Harmful Algae 6, 73–80.
Shams, S., Capelli, C., Cerasino, L., Ballot, A., Dietrich, D.R., Sivonen, K., Salmaso, N., 2015. Anatoxin-a producing Tychonema
(cyanobacteria) in European waterbodies. Water Res. 69, 68–79.
Sivonen, K., Carmichael, W.W., Namikoshi, M., Rinehart, K.L., Dahlem, A.M., Niemelä, S.I., 1990. Isolation and characterization of
hepatotoxic microcystin homologs from the filamentous freshwater cyanobacterium Nostoc sp. strain 152. Appl. Environ.
Microbiol. 56, 2650–2657.
Sivonen, K., Himberg, K., Luukkainen, R., Niemelä, S.I., Poon, G.K., Codd, G.A., 1989. Preliminary characterization of neurotoxic
cyanobacteria blooms and strains from Finland. Toxic. Assess. 4, 339–352.
Page 13 of 13
Sivonen, K., Namikoshi, M., Evans, W.R., Färdig, M., Carmichael, W.W., Rinehart, K.L., 1992. Three new microcystins, cyclic
heptapeptide hepatotoxins, from Nostoc sp. strain 152. Chem. Res. Toxicol. 5, 464–469.
Smith, F.M.J., Wood, S.A., Wilks, T., Kelly, D., Broady, P.A., Williamson, W., Gaw, S., 2012. Survey of Scytonema (cyanobacteria) and
associated saxitoxins in the littoral zone of recreational lakes in Canterbury, New Zealand. Phycologia 51, 542–551.
Song, L., Sano, T., Li, R., Watanabe, M.M., Liu, Y., Kaya, K., 1998. Microcystin production of Microcystis viridis (cyanobacteria) under
different culture conditions. Phycol. Res. 46, 19–23.
Spoof, L., Berg, K.A., Rapala, J., Lahti, K., Lepistö, L., Metcalf, J.S., Codd, G.A., Meriluoto, J., 2006. First observation of
cylindrospermopsin in Anabaena lapponica isolated from the boreal environment (Finland). Environ. Toxicol. 21, 552–560.
Stefanelli, M., Scardala, S., Cabras, P.A., Orrù, A., Vichi, S., Testai, E., Funari, E., Manganelli, M., 2017. Cyanobacterial dynamics and
toxins concentrations in Lake Alto Flumendosa, Sardinia, Italy. Adv. Oceanogr. Limnol. 8.
Taylor, M.S., Stahl-Timmins, W., Redshaw, C.H., Osborne, N.J., 2014. Toxic alkaloids in Lyngbya majuscula and related tropical marine
cyanobacteria. Harmful Algae 31, 1–8.
Teneva, I., Asparuhova, D., Dzhambazov, B., Mladenov, R., Schirmer, K., 2003. The freshwater cyanobacterium Lyngbya aerugineo-
coerulea produces compounds toxic to mice and to mammalian and fish cells. Environ. Toxicol. 18, 9–20.
Teneva, I., Stoyanov, P., Belkinova, D., 2012. Production of cyanobacterial toxins from two Nostoc species (Nostocales) and evaluation of
their cytotoxicity in vitro. J. BioSci. Biotec 1, 33–43.
Tomsickova, J., Ondrej, M., Cerny, J., Hrouzek, P., Kopecky, J., 2014. Analysis and Detection of Scytophycin Variants by HPLC-ESI-MS.
Chem. Nat. Compd. 49, 1170–1171.
Vehovszky, Á., Ács, A., Kovacs, A.W., Szabó, H., Gyori, J., Farkas, A., 2009. Isolated strains of Cylindrospermopsis raciborskii from Lake
Balaton (Hungary) produce anatoxin-a like neurotoxins. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 153, S88.
Vestola, J., Shishido, T.K., Jokela, J., Fewer, D.P., Aitio, O., Permi, P., Wahlsten, M., Wang, H., Rouhiainen, L., Sivonen, K., 2014.
Hassallidins, antifungal glycolipopeptides, are widespread among cyanobacteria and are the end-product of a nonribosomal
pathway. Proc. Natl. Acad. Sci. 111, E1909–E1917.
Vezie, C., Brient, L., Sivonen, K., Bertru, G., Lefeuvre, J.C., Salkinoja-Salonen, M., 1998. Variation of microcystin content of
cyanobacterial blooms and isolated strains in Lake Grand-Lieu (France). Microb. Ecol. 35, 126–135.
Viaggiu, E., Melchiorre, S., Volpi, F., Di Corcia, A., Mancini, R., Garibaldi, L., Crichigno, G., Bruno, M., 2004. Anatoxin-a toxin in the
cyanobacterium Planktothrix rubescens from a fishing pond in northern Italy. Environ. Toxicol. 19, 191–197.
Vieira, J.M. dos S., Azevedo, M.T.D.P., De Oliveira Azevedo, S.M.F., Honda, R.Y., Corrêa, B., 2003. Microcystin production by
Radiocystis fernandoi (Chroococcales, Cyanobacteria) isolated from a drinking water reservoir in the city of Belém, PA, Brazilian
Amazonia region. Toxicon 42, 709–713.
Wang, H., Sivonen, K., Rouhiainen, L., Fewer, D.P., Lyra, C., Rantala-Ylinen, A., Vestola, J., Jokela, J., Rantasärkkä, K., Li, Z., Liu, B.,
2012. Genome-derived insights into the biology of the hepatotoxic bloom-forming cyanobacterium Anabaena sp. strain 90. BMC
Genomics 13, 613.
Watanabe, M.F., Tsujimura, S., Oishi, S., Niki, T., Namikoshi, M., 2003. Isolation and identification of homoanatoxin-a from a toxic
strain of the cyanobacterium Raphidiopsis mediterranea Skuja isolated from Lake Biwa, Japan. Phycologia 42, 364–369.
Williams, P.G., Yoshida, W.Y., Moore, R.E., Paul, V.J., 2002. Isolation and structure determination of obyanamide, a novel cytotoxic
cyclic depsipeptide from the marine cyanobacterium Lyngbya confervoides. J. Nat. Prod. 65, 29–31.
Wood, S.A., Rasmussen, J.P., Holland, P.T., Campbell, R., Crowe, A.L.M., 2007. First report of the cyanotoxin anatoxin-a from
Aphanizomenon issatschenkoi (cyanobacteria). J. Phycol. 43, 356–365.
Yilmaz, M., Foss, A.J., Selwood, A.I., Özen, M., Boundy, M., 2018. Paralytic shellfish toxin producing Aphanizomenon gracile strains
isolated from Lake Iznik, Turkey. Toxicon 148, 132–142.
Yilmaz, M., Phlips, E.J., Szabo, N.J., Badylak, S., 2008. A comparative study of Florida strains of Cylindrospermopsis and Aphanizomenon
for cylindrospermopsin production. Toxicon 51, 130–9.
Yunes, J.S., De La Rocha, S., Giroldo, D., Silveira, S.B. Da, Comin, R., Bicho, M.D.S., Melcher, S.S., Sant'Anna, C.L., Vieira, A.A.H.,
2009. Release of carbohydrates and proteins by a subtropical strain of Raphidiopsis brookii (cyanobacteria) able to produce
saxitoxin at three nitrate concentrations. J. Phycol. 45, 585–591.