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University of Groningen What is natural? Haydar, Deniz IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2010 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Haydar, D. (2010). What is natural? The scale and consequences of marine bioinvasions in the North Atlantic Ocean. Groningen: s.n. Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date: 09-06-2019

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Page 1: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

University of Groningen

What is natural?Haydar, Deniz

IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite fromit. Please check the document version below.

Document VersionPublisher's PDF, also known as Version of record

Publication date:2010

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):Haydar, D. (2010). What is natural? The scale and consequences of marine bioinvasions in the NorthAtlantic Ocean. Groningen: s.n.

CopyrightOther than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of theauthor(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons).

Take-down policyIf you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediatelyand investigate your claim.

Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons thenumber of authors shown on this cover page is limited to 10 maximum.

Download date: 09-06-2019

Page 2: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Appendix II to Chapter 4: The scale of cryptogenesis, in:

Deniz Haydar (2010) What is natural? The scale and consequences of marine bioinvasions in the North Atlantic Ocean.

Dissertation, University of Groningen, 184 pp.

ISBN 978-90-367-4396-9

ISBN 978-90-367-4395-2

available at http://dissertations.ub.rug.nl/faculties/science/2010/

Appendix II Hydrozoa species in the North Atlantic ocean, excluding disjunct amphi-Atlantic species (see table 4.3).

species distribution category depth habitat dispersal comments references

Anthoathecata

Acaulidae

Acaulis primarius

Stimpson, 1854

Greenland, Iceland, White Sea, Norway,

North Sea, Kattegat, Baltic, Bay of Fundy to

Cape Cod, New Brunswick, Bay of Biscay?

AMPHI 20-350m partially embedded in

sediment

P 1, 2, 3, 4, 5,

6, 7

Acauloides ammisatum

Bouillon, 1965

Roscoff, Mediterranean? EU shallow on sand possibly conspecific with Mediterranean

A. ilonae; Mediterranean record

uncertain.

1, 2, 5

Boreohydridae

Boreohydra simplex

Westblad, 1937

Bipolar, N and S Atlantic Ocean. White Sea,

Greenland, Iceland, Norway to Sweden,

British Isles, English Channel, NE Canada,

South Georgia

AMPHI few m->600m (moving) on and burrowing

in mud

not in Mediterranean. Probably more

widespread, but misidentified as worm or

overlooked.

1, 2, 3, 5

Psammohydra nanna

Schulz, 1950

Kattegat, Kiel, English Channel, W

Mediterranean, Adriatic Sea

EU few meters in fine sand, tolerates

reduced salinity, can move

by creeping

P 1, 2, 5

Bougainvilliidae

Bougainvillia britannica

(Forbes, 1841)

S Norway, British Isles, W of Iceland. EU coastal gastropod, bivalve shells M Mediterranean records probably

erroneous (2), One record from Maine

(9), and from China, Alaska, probably

erroneous too

2, 8, 9

Bougainvillia carolinensis

(McCrady, 1859)

Arctic S to Caribbean AM intertidal -

shallow

docks, ships M fouling 4, 11

Bougainvillia macloviana

Lesson, 1830

Falklands, South Africa, Kerguelen, New

Zealand. Introduced to North Sea

EU coastal fouling M hydroid not observed in North Sea.

probably introduced on ships as hydroid;

direct observation of hydroid on ship by 9

2, 9

Bougainvillia muscoides

(Sars, 1846)

Norway, Sweden, British Isles, one record

from Chile

EU 4-200m hydroids M Chile record confirmed by 16S data 2, 8, 9

Page 3: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Bougainvillia principis

(Steenstrup, 1850)

Arctic S to English Channel, Iceland,

Greenland, New England, NE Pacific

AMPHI shallow, 30m stones M 2, 9

Bougainvillia pyramidata

(Forbes & Goodsir, 1853)

British Isles, Sweden EU 30-80m, mainly

near deeper

water

ascidians, hydroids M 2, 8, 9

Bougainvillia superciliaris

(Agassiz, 1849)

Arctic S to North Sea, Cape Hatteras, possibly

San Francisco

AMPHI shallow gastropod shells, mussels M 4, 6, 9

Dicoryne conferta

(Alder, 1856)

Iceland, Barents Sea to Gibraltar,

Mediterranean, South Africa, NE Canada S

to Cape Cod

AMPHI 5-300m gastropod shells, spider

crabs

P South Africa: on Agulhas bank, False

Bay to Mossel bay. 12 does not mention

that it might be introduced.

2, 3, 4, 8, 9,

10, 12

Dicoryne conybearei

(Allman, 1846)

S British Isles S to Iberian Peninsula,

Mediterranean, Japan and New Zealand

EU coastal gastropod shells P cryptogenic. sparsely recorded around

British Isles. Recorded from Japan and

New Zealand? Could be only variety of

D. conferta

2, 8, 9, 10

Garveia brevis

(Fraser, 1918)

Arctic S to Cape Cod AM 18-24m Tubularia, piles P fouling 4

Garveia cerulea

(Clarke, 1882)

Arctic, Cape Cod to Cape Hatteras AM intertidal -

shallow

coralline algae, piles, ships P fouling. possibly conspecific with G.

franciscana

4, 11, 47

Garveia nutans

Wright, 1859

Shetlands to Plymouth, Bay of Biscay,

Gibraltar, Mediterranean, Belgium, France

EU lower shore -

30m (580m)

stone, rock, hydroids,

seaweed, various hard

substrata, in strong tidal

currents

P 2, 8, 9

Koellikerina fasciculata

(Péron & Lesueur, 1810)

Mediterranean, Black Sea, Red Sea, Cape

Verdes, Scotland and Norway

EU H:20-270m, M:

shallow

polychaetes, urchin spines M probably not reproducing in Scotland and

Norway

2, 9

Lizzia blondina

Forbes, 1848

Arctic, Norway to Mediterranean, NW

Atlantic, Iceland, Faroes, W Africa, Florida,

New Zealand

AMPHI nearshore H: unknown M hydroid of this species unknown.

Hydractinia minuta included

2, 4, 9, 10

Lizzia elisabethae

Haeckel, 1879

Jersey, English Channel (type locality) EU M 9

Pachycordyle michaeli

(Berrill, 1948)

Maine to Chesapeake Bay AM shallow wooden floats, Fucus P doubtful species. closely resembles P.

navis, status should be re-evaluated

2, 9, 13

Rhizorhagium album

Rees, 1938

Cornwall EU shore rock pool on filamentous

algae

P doubtful species, possibly subspecies of

R. arenosum

2, 8, 9

Rhizorhagium arenosum

(Alder, 1862)

North Sea coasts British Isles, SW Ireland EU intertidal algae, stones, other

substrates

P 2, 9

Rhizorhagium roseum

Sars, 1874

Arctic circumpolar, S to Scotland, NE

Canada, W Canada, Alaska

AMPHI 15-200m hydroids P 2, 3, 4, 8, 9

Page 4: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Candelabridae

Candelabrum cocksii

(Cocks, 1854)

NE Atlantic, absent from North Sea, Baltic,

Mediterranean and Black Sea. English

Channel, Brittany, Galicia, Bay of Cadiz

EU lower shore-

17m (110-

146m)

under stones, Laminaria

holdfasts

other Deeper records from Aalesund (Norway)

doubtful. C. phrygium from deeper water

regarded as separate species.

Brooding embryos

2, 5, 8, 10

Candelabrum minutum

(Bonnevie, 1898)

N Norway (Tromso) ARCT P likely synonym of C. phrygium 1, 2, 46

Candelabrum phrygium

(Fabricius, 1780)

Arctic, British Isles, off Iberian Peninsula,

Mid Atlantic Ridge, Bay of Fundy to Cape

Cod

AMPHI 13m in high

Arctic - 2195m

on rock, bivalves,

hydroids, bryozoans, algae

other submerges in S part of range, shallow

records from temperate waters are C.

cocksii. Blastothela rosea in 4 is this

species.

viviparous

1, 2, 3, 4, 5,

6, 14

Candelabrum verrucosum

(Bonnevie, 1898)

N Norway, S Greenland ARCT 50m rocky bottom other viviparous 1, 2, 5

Monocoryne gigantea

(Bonnevie, 1898)

Arctic, S to N Norway, NE Canada ARCT 16-20m high

Arctic - >100m

on Tubularia and

polychaete tubes on Lima

excavata shells

P 1, 2, 5, 14

Cladonematidae

Eleutheria claparedii

Hartlaub, 1889

French coast English Channel, Mediterranean EU intertidal medusa clinging on

macroalgae like Ulva and

Fucus, polyp not recorded

in nature

M 1, 2, 5

Corymorphidae

Branchiocerianthus

norvegicus

Brattström, 1957

Bergen EU shallow? 1, 2

Branchiocerianthus

reniformis

Broch, 1918

Davis Strait ARCT 1

Corymorpha appelloefi

Bonnevie, 1901

Bergen EU shallow? P 1, 2

Corymorpha glacialis

Sars, 1859

Arctic. N Norway, Iceland, Greenland,

Barents Sea, White Sea, Kara Sea,

Spitsbergen, Nova Zembla, Davis Strait

ARCT shallow? P 1, 2, 3

Corymorpha groenlandica

(Allman, 1876)

Arctic, S to between Scotland and Faroes ARCT,

EU

usually deeper

than 100m,

1253m

attached to Lophelia P deepwater species? 1, 2, 3, 7

Corymorpha nana

Alder, 1857

Britain, NE Finnmark EU shallow? 1, 2

Page 5: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Corymorpha pendula

Agassiz, 1862

Arctic S to Rhode Island AM 20-275m M 4, 6, 7

Corymorpha sarsii

Steenstrup, 1854

Arctic. Lofoten, NE Finnmark ARCT shallow? 1, 2

Gymnogonos crassicornis

Bonnevie, 1898

Lofoten EU shallow? 1, 2

Plotocnide borealis

Wagner, 1885

Baltic, Oslo, Scotland, Arctic EU M not in Mediterranean 1, 2

Plotocnide incerta

(Linko, 1900)

White Sea ARCT 1

Corynidae

Coryne eximia

Gaertner, 1774

Norway S to Galicia, E coast N America S to

Nova Scotia, Alaska S to California, South

Africa, Chile, New Zealand, W Australia.

Mediterranean, Greenland, Iceland, Brazil,

Papua: need confirmation

AMPHI lower shore -

25m

rocks, floats, laminaria,

mussels, algae

M 1, 2, 3, 4, 8,

10, 15

Coryne filiformis

(Rees, 1936)

British Isles EU shallow on Eunicella verrucosa(=

Sea fan), Pecten shells

P similar to C. pintneri 1, 15

Coryne fucicola

(de Filippi, 1866)

Roscoff, Mediterranean EU shallow algae P recent rediscovery 16

Coryne hincksii

Bonnevie, 1898

Arctic. Norway, Greenland, Canada, Russia ARCT P 1, 2, 14, 15

Coryne muscoides

(Linnaeus, 1761)

British Isles N to Orkneys, Sweden, France,

Iberian Peninsula, Mediterranean, Indo-

Pacific?

EU 0 - 2m algae, lithophytes P Mediterranean Coryne might be different

species

1, 2, 8, 10,

15

Coryne pintneri

Schneider, 1898

Mediterranean, Brittany, Scotland EU 1-30m Posidonia, red algae, rock P similar to C. filiformis 1, 15

Coryne producta

(Wright, 1858)

White Sea, Iceland S to Bay of Biscay, not in

Mediterranean, not in Greenland, not in W

Atlantic

EU just subtidal hydroids, shells, solid

objects

M not in W Atlantic 1, 2, 3, 10,

15, 47

Coryne prolifera

(Forbes, 1848)

British Isles, France, Mediterranean? EU M polyp stage unknown 1, 2, 15

Coryne pusilla

Gaertner, 1774

cosmopolitan. Greenland, Iceland, N Norway

to Cape Verdes, Mediterranean, NE Canada,

South Africa, Kerguelen, Seychelles, Korea,

Japan, New Zealand

AMPHI intertidal -

shallow

brown algae, man made

floats, ships

P ship fouling. rafting on algae 1, 2, 3, 8,

10, 11, 14,

15, 17

Page 6: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Coryne vanbenedeni

(Hincks, 1869)

Ostende (Belgium) EU shallow on crab P doubtful species, probably

misidentification, only known from type

locality and never observed again

1, 2, 15

Dipurena gemmifera

(Forbes, 1848)

Bergen, British Isles S to NW Spain,

Mediterranean

EU M polyp stage unknown 1, 2, 10, 15

Dipurena ophiogaster

Haeckel, 1879

British Isles to Mediterranean, Skagerrak, Sri

Lanka, Japan, Palau Islands, Papua New

Guinea, Pacific Mexico, Chile, New Zealand.

circumtropical

EU shallow algae, barnacles, rocks M 1, 10, 15

Dipurena reesi

Vannucci, 1956

Bay of Biscay, Mediterranean, Brazil,

California

EU 1-50m variety of solid substrates M circumtropical? 2, 10, 15

Dipurena simulans

Bouillon, 1965

Roscoff, Brittany. EU shallow in and on sponge Haliclona

simulans

M probably more wide-spread 1, 15

Dipurena spongicola

Anger, 1972

Germany, Sweden (Baltic) EU 9-10m within canal system sponge

Halichondria panicea

M 1, 15

Dipurena strangulata

McCrady, 1859

Cape Cod to Florida, Puerto Rico, Gulf of

Guinea

AM shallow in sponge Microciona

prolifera

M tolerates reduced salinity 15

Sarsia barentsi

Linko, 1905

Barents Sea ARCT 1, 2

Sarsia densa

Hartlaub, 1897

North Sea EU shallow hard substrates M might be S .tubulosa 1, 15

Sarsia lovenii

(Sars, 1846)

Arctic to boreal Atlantic, S to North Sea,

Baltic Sea, N New England

AMPHI 0 - 200m rocks, stones, weeds, solid

objects

P tolerant of reduced salinity 2, 3, 8, 15

Sarsia piriformis

Edwards, 1983

Scotland EU 15-20m on clinker M only known from type locality 1, 2, 15

Sarsia princeps

(Haeckel, 1879)

Arctic circumpolar, S to Newfoundland ARCT 15-25m shells M 1, 2, 15

Sarsia striata

Edwards, 1983

Scotland EU 15-20m on clinker M 1, 2, 15

Sarsia tubulosa

(Sars, 1835)

Circumpolar, coastal boreal and Arctic.

Boreal Atlantic S to English Channel, Iberian

Peninsula, S to Caribbean. Alaska S to San

Francisco Bay, Japan.

AMPHI P: near low

water, M: 20-

100m

rocks, stones, weeds M near cosmopolitan, may be species

complex. tolerant of reduced salinity.

rafting inferred from distribution

1, 2, 3, 6, 7,

8, 10, 13, 14,

15, 17, 18

Sarsiella dinema

Hartlaub, 1907

Normandy, Mediterranean EU M only medusa known, doubtful species 1, 2, 46

Eucodoniidae

Page 7: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Eucodonium brownei

Hartlaub, 1907

Gibraltar, Mediterranean. Atlantic? EU hydroid

unknown

M 1, 2, 10, 19

Eudendriidae

Eudendrium annulatum

Norman, 1864

North Atlantic, boreal to Arctic. S to English

Channel (beached material)

AMPHI shallow 2, 4, 6

Eudendrium arbusculum

Wright, 1859

Spitsbergen, Greenland, Canada, NW Europe,

Mediterranean

AMPHI 2, 3, 9, 14

Eudendrium cingulatum

Stimpson, 1854

Bay of Fundy to Cape Cod AM shallow 4

Eudendrium dispar

Agassiz, 1862

Arctic to Cape Hatteras AM shallow 4, 6

Eudendrium glomeratum

Picard, 1951

circumtropical. Plymouth, S Devon, Ireland,

NE Spain, Mediterranean

EU 5-40m biological concretions P 2, 8, 10, 20

Eudendrium insigne

Hincks, 1861

Casco Bay, Maine AM shallow 4

Eudendrium islandicum

Schuchert, 2000

Iceland, Greenland ARCT 40-996m very likely synonym of D. album - which

has a disjunct distribution, adding these

records makes the distribution amphi-

Atlantic

2, 3

Eudendrium racemosum

(Cavolini, 1785)

NW Spain, Arctic? Mediterranean, Indo

Pacific?

EU P 2, 10

Eudendrium rameum

(Pallas, 1766)

Arctic S to Mediterranean, S to Cape Cod AMPHI 5-100m various substrata P widely distributed. 2, 4, 8, 10,

14, 21

Eudendrium ramosum

(Linnaeus, 1758)

Iceland S to Mediterranean, Arctic S to

Caribbean, Australia, South Africa

AMPHI 5-200m,

sometimes

deeper

various substrata, ships P not all records are reliable. ship fouling 2, 3, 4, 6, 8,

10, 11

Eudendrium vaginatum

Allman, 1863

Arctic circumpolar, S to N British Isles,

Norway, S to Cape Hatteras, Japan

AMPHI shallow 2, 3, 4

Euphysidae

Euphysa aurata

Forbes, 1848

Arctic, NW Europe, Baltic, Bay of Biscay,

Mediterranean, Massachusetts Bay,

Argentina?

AMPHI shallow on mud M disjunct or amphi-Atlantic? infrequently

recorded because it occurs on mud,

probably amphi-Atlantic

1, 2, 3, 4, 10,

14

Euphysa farcta

(Miles, 1937)

Maine to Cape Cod AM shallow M possibly synonym of E. aurata., doubtful

species

7

Euphysa flammea

(Linko, 1905)

Arctic ARCT not in Mediterranean 1, 2

Page 8: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Euphysa obvoluta

(Kramp, 1933)

E Greenland ARCT 1, 2

Euphysa peregrina

(Murbach, 1899)

Cape Cod S to Cape Hatteras AM 4, 47

Euphysa tentaculata

Linko, 1905

Arctic. Kola Peninsula, Davis Strait, Baltic,

Kattegat

ARCT 1, 2

Hydractiniidae

Clava multicornis

(Forsskal, 1775)

Arctic. S to Iberian Peninsula, Mediterranean.

S to Cape Hatteras, San Francisco Bay

AMPHI intertidal -

163m

Ascophyllum, algae, on and

under stones, rock pools,

shells

P tolerant of reduced salinity and emersion.

Seasonal dormancy. Introduced to San

Francisco Bay. Rafting on algae.

Dispersal by shipping,.

1, 2, 3, 4, 6,

8, 10, 17, 22,

26

Hydractinia allmani

Bonnevie, 1898

Arctic ARCT 40-250m,

extremes: 3-

1500m

gastropod shells M 3

Hydractinia americana

(Edwards, 1972)

Bay of Fundy S to Florida AM intertidal - 50m gastropod shells, hermit

crab shells, bivalves,

barnacles, larger

crustaceans, stones

M used to be included in H. carnea, now

recognized as separate species

6

Hydractinia areolata

(Alder, 1862)

British Isles, Belgium, NW Spain,

Mediterranean

EU M 2, 10

Hydractinia arge

(Clarke, 1882)

Cape Cod S to South Carolina AM shallow gastropod shells, algae,

rock, sponges

M 4, 7, 13, 23

Hydractinia carica

Bergh, 1887

Arctic ARCT 0-120m Buccinum shells P 3, 4, 14

Hydractinia carnea

Sars, 1846

N Norway S to British Isles, North Sea,

Belgium, The Netherlands (Zuiderzee?),

Iberian Peninsula, Mediterranean

EU intertidal - 50m gastropod shells, hermit

crab shells, bivalves,

barnacles, larger

crustaceans, stones

M 2, 3, 8, 10

Hydractinia claviformis

(Bouillon, 1965)

France, NW Spain EU shallow algae, sponges, rocks P 2, 10, 23

Hydractinia echinata

Fleming, 1823

Arctic to Morocco, Mediterranean EU shore - 30m hermit crab gastropod

shells, sometimes other

solid substrata

P not on NE American coast 2, 3, 8, 10,

24

Hydractinia hooperii

(Sigerfoos, 1899)

Cape Cod S to Caribbean AM shallow hermit crab shells M possibly synonym of H. arge 4, 7, 23

Hydractinia monocarpa

Allman, 1876

Arctic. Newfoundland, Spitsbergen, probably

also Greenland, Iceland

ARCT shallow gastropod shells P 3, 14

Page 9: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Hydractinia polyclina

Agassiz, 1862

Maine to Massachusetts AM shallow subtidal on hermit crab shells,

particularly Pagurus

acadianus

one of three American species that were

all ascribed to H. echinata. Agassiz

separated European H. echinata and the

Amerian species, but they were

considered conspecific by later workers.

Breeding experiments have shown that

they are separate species.

24

Hydractinia sarsii

Steenstrup, 1850

Greenland, Iceland, Faroes, Bergen ARCT shallow? P 3

Hydractinia serrata

Kramp, 1943

Greenland, Barents Sea ARCT 20-165m P 3

Hydractinia

symbiolongicarpus

Buss & Yund, 1989

Freeport, Maine to Connecticut AM shallow subtidal on hermit crab shells,

particularly Pagurus

longicarpus

one of three American species that were

all ascribed to H. echinata

24

Hydractinia

symbiopollicaris

Buss & Yund, 1989

Woods Hole, Connecticut AM shallow subtidal on hermit crab shells,

particualrly Pagurus

pollicaris

one of three American species that were

all ascribed to H. echinata

24

Hydractinia valens

Fraser, 1941

Maine AM shallow P 4

Margelopsidae

Margelopsis haeckelii

Hartlaub, 1897

S North Sea, Irish Sea, Helgoland, Weser,

Elbe and Ems estuary, Belgium, E Britain

EU planktonic planktonic M both polyp and medusa planktonic.

overwintering cyst stage

1, 2, 5

Moerisiidae

Moerisia lyonsi

Boulenger, 1908

Black Sea, Middle East, Chesapeake Bay S to

Louisiana, San Francisco Bay

AM shallow brackish water species introduced from Black Sea to Chesapeake

Bay and San Francisco Bay

25, 26, 27

Oceaniidae

Corydendrium dispar

Kramp, 1935

Sweden, Norway, Faroes EU 26-184m shells, hydroids M 2, 22

Oceania armata

Kölliker, 1853

Spain, Portugal, Mediterranean, Canaries,

Azores, Cape Verdes, West Indies, Japan,

New Zealand, Tasman Sea

EU shallow - 200m unknown M polyp never found in nature 2, 22

Pachycordyle navis

(Millard, 1959)

South Africa, Baltic, S England, The

Netherlands, Denmark, Black Sea

EU shallow algae, wood, iron

constructs, Mytilus, other

solid substrata, ships

P =Clavopsella quadranularia, Thieliana

navis. described from ship hull in Table

Bay. introduced. origin could be Black

Sea

12, 22, 39

Rhizogeton fusiformis

Agassiz, 1862

Bay of Fundy to Cape Cod AM intertidal in rock pools P is not R. nudus 4, 22

Page 10: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Rhizogeton nudus

(Broch, 1909)

Spitsbergen, Norway, Greenland, Iceland,

British Isles, NE Canada

AMPHI 0-100m on Mytilus, barnacles,

hydroids, bryozoans,

macroalgae

P records from South Africa, Argentina and

tropical Indian Ocean are probably

separate species

1, 3,14, 22

Similomerona

nematophora

(Antsulevich, 1986)

Arctic. Franz Joseph Land ARCT shallow 5, 22

Turritopsis nutricula

McCrady, 1857

Massachusetts to Caribbean AM shallow on algae, ships P “immortal", reverts back to polyp stage

after sexual maturity. Regarded as

cosmopolitan, but actually four separate

spp. ship fouling.

4, 10, 11, 13,

28, 29

Turritopsis polycirrha

(Keferstein, 1862)

S North Sea, English Channel EU shallow shells, pebbles M has been synonimized with American T.

nutricula, but is a separate species. polyp

phase not well known.

2, 22

Pandeidae

Catablema multicirratum

Kishinouye, 1910

NW Atlantic, N Pacific AM M 9

Catablema vesicarium

(Agassiz, 1862)

Arctic. N Norway, Iceland, Greenland, S to

Cape Cod

ARCT shallow M hydroid stage unknown 6, 9

Halitholus cirratus

Hartlaub, 1914

Arctic, Baltic EU shallow - 100m on several small bivalves,

Nuculidae, Astartidae.

M tolerates brackish waters. =Perigonimus

yoldiaearcticae

2, 3, 9

Halitholus pauper

Hartlaub, 1914

Arctic. Iceland, Greenland, Canada, British

Columbia, Kamchatka, N Japan, New Zealand

ARCT shallow M hydroid stage unknown 9

Hydrichthys mirus

Fewkes, 1887

Cape Cod S to Cape Hatteras AM shallow on fish M 4

Hydrichthys sarcotretis

(Jungersen, 1912)

NW Atlantic, Cape Cod Bay, Europe AMPHI? shallow-deep parasitic on copepods on

redfish (Templeman, 1973)

2

Leuckartiara nobilis

Hartlaub, 1914

Ireland, Scotland, Rockall, Iceland,

Newfoundland, Japan, British Columbia

AMPHI oceanic M hydroid unknown 9

Leuckartiara octona

(Fleming, 1823)

near cosmopolitan. P: England, South Africa.

M: Lofoten, Iceland S to Portugal, W Africa,

Labrador to C Hatteras, Mediterranean, W

Africa, Tristan da Cunha, India, Malayan

Archipelago, NE Australia, Low Archipelago,

China, Japan, Chile, Vancouver, New Zealand

AMPHI 10-400m gastropod shells, spider

crabs, hydroids,

invertebrates, ships

M near cosmopolitan, ship fouling 2, 3, 4, 6, 8,

10, 11

Neoturris breviconis

(Murbach & Shearer,

1902)

circumpolar S to N North Sea, S Iceland,

British Columbia, Japan

ARCT shallow M Atlantic population needs to be

reinvestigated, uncertain status of species

9

Page 11: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Neoturris pileata

(Forsskal, 1775)

M: Arctic S to Iberian Peninsula,

Mediterranean, SW Africa. H: Scotland,

Norway, Sweden

EU H: 40-180m H: exclusively on Nucula,

adapted to live buried in

the mud

M can be confused with Leuckartiara

nobilis

2, 9, 10

Pennariidae

Pennaria disticha

(Goldfuss, 1820)

Massachusetts to Caribbean AM shallow ships =Halocordyle disticha

ship fouling.

rafting inferred from distribution

13, 17

Proboscidactylidae

Proboscidactyla stellata

(Forbes, 1846)

Norway, British Isles, NW France EU shallow sabellid tubes M hydroid stage seldom recorded 2, 8

Ptilicodiidae

Thecocodium brieni

Bouillon, 1967

W Scotland, Roscoff EU shallow unknown 1, 2

Thecocodium penicillatum

Jarms, 1987

NE Atlantic EU shallow 1, 2, 19

Rathkeidae

Rathkea octopunctata

(Sars, 1835)

Arctic S to Gibraltar, Mediterranean, Black

Sea, S to New England, S to British

Columbia, Australia, New Zealand

AMPHI H: shallow

pools; M:

coastal

H: oyster and mussel

shells, M: lagoons,

brackish waters, survives

in polluted waters

M medusae able to reproduce asexually by

budding. New Zealand medusae could be

separate species based on morphology

and genetic data

9

Sphaerocorynidae

Sphaerocoryne agassizii

(McCrady, 1859)

Massachusetts S to S Carolina AM shallow Sargassum, shells, serpulid

tubes, pebbles, wood,

always with encrusting

bryozoa

M often confused with Zanclea gemmosa 4, 13

Stylasteridae

Stenohelia maderensis

(Johnson, 1862)

Iberian Peninsula EU P 10, 2

Tricyclusidae

Tricyclusa singularis

(Schulze, 1876)

W Ireland, N Brittany, Adriatic Sea EU near lower

shore

attached to Zostera and

macroalgae, sometimes

detached polyps or young

individuals in the plankton,

with very long and thin

tentacles until they attach

P possibly extinct in Mediterranean.

rafting?

Associated with Zostera, sometimes in

plankton

5

Page 12: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Tubulariidae

Ectopleura americana

Petersen, 1990

Long Island Sound AM “First and last seen on a ship hull in 1879

in Long Island Sound”

30, 47

Ectopleura larynx

(Ellis & Solander, 1786)

near cosmopolitan.Arctic, S to Iberian

Peninsula, S to Long Island Sound,

Mediterranean, New Zealand, Mid Atlantic

Ridge.

AMPHI lower shore -

>100m, usually

<35m, but also

845m

hydroids, solid substrata,

ships

P tolerant of slightly reduced salinity

ship fouling

1, 2, 3, 4, 6,

8, 10, 11, 31

Hybocodon prolifer

Agassiz, 1862

M: northern boreal circumpolar, S to Bay of

Biscay, S to Chesapeake Bay, S to Puget

Sound, New Zealand. H: NE American coast,

Plymouth, Iceland, New Zealand

AMPHI lower shore-

sublittoral

little known, sponge M disjunct, present in northern hemisphere

and New Zealand

2, 3, 4, 8,

10, 32

Tubularia acadiae

Petersen, 1990

NW Atlantic AM no other information found on this

species

47

Tubularia asymmetrica

Bonnevie, 1898

Norway EU M 1

Tubularia indivisa

Linnaeus, 1758

Arctic, Norway, Iceland, Faroes, British Isles,

S to Iberian Peninsula, W Africa,

Mediterranean, S to Cape Hatteras, N Pacific

AMPHI intertidal -

300m

various solid substrata,

ships

M seasonal dormancy, ship fouling 1, 2, 3, 4, 6,

8, 10, 11

Tubularia regalis

Boeck, 1860

Arctic. Norway, Spitsbergen, Nova Zembla,

Shetland-Faroe channel, NE Greenland, N

Canada

ARCT 1, 2, 3, 14

Zancleidae

Zanclea gemmosa

McCrady, 1859

Cape Cod to Florida AM shallow Sargassum, shells, serpulid

tubes, pebbles, wood,

always with encrusting

bryozoa

M rafting on algae 2, 4

Zanclea sessilis

(Gosse, 1853)

British Isles EU 1, 2

Leptothecata

Aequoreidae

Aequorea albida

Agassiz, 1862

NW Atlantic? AM validity uncertain 6, 48

Aequorea vitrina

Gosse, 1853

British Isles, Belgium, Helgoland, Denmark EU probably coastal M 32

Rhacostoma atlanticum

Agasssiz, 1850

Cobscook Bay, Gulf of Mexico, Nova Scotia

to Brazil

AM M validity uncertain 6, 48

Page 13: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Aglaopheniidae

Aglaophenia acacia

Allman, 1833

British Isles, Brittany to NW Spain, Azores,

Canaries, Mediterranean

EU 8-823m hard substrata P possibly further N in Europe, deepest

record from Azores. NW Atlantic records

misidentifications (47)

2, 8, 32

Aglaophenia

kirchenpaueri

(Heller, 1868)

Iberian Peninsula to Morocco, Mediterranean,

Cape Verdes

EU shallow, some

deep records

rock , corraline algae,

brown algae, gorgonians

P probably overlooked

not reported rafting

2, 8, 10, 32,

33

Aglaophenia octodonta

(Heller, 1868)

Iberian Peninsula, Mediterranean EU shallow P 2, 10

Aglaophenia parvula

Bale, 1882

British Isles, Iberian Peninsula, Brittany,

Morocco, South Africa, Australia, S Indian

Ocean, St Paul Island

EU shallow mostly sponges, once

Mytilus, barnacles, algae

P disjunct distribution? Often confused with

A. pluma. not reported rafting

2, 8, 10, 32,

33

Aglaophenia picardi

Svoboda, 1979

Iberian Peninsula, Mediterranean EU P 2, 10

Aglaophenia pluma

(Linnaeus, 1767)

cosmopolitan. British Isles, N France, Iberian

Peninsula, Morocco, Mediterranean

EU intertidal - 20m brown algae, rock , gravel,

ships

P cosmopolitan, but identification is

difficult, might be a species complex.

Rafting on algae inferred from

distribution, ship fouling

2, 8, 10, 11,

17, 32, 33

Aglaophenia tubiformis

Marktanner-Turneretscher,

1890

N Brittany southwards EU shallow algae, rocks P very similar to A. pluma 2, 10, 32

Aglaophenia tubulifera

(Hincks, 1861)

Scotland S to Cape Verdes, Azores, Morocco,

Strait of Gibraltar, Guinea

EU 10->80m,

100m, 1200m

rocks, boulders, pebbles in

moderate current

P 2, 8, 32

Cladocarpus boucheti

Ramil & Vervoort, 1992

Gibraltar EU deep? P Cladocarpus is primarily, but not

exclusively a deep water genus

2, 10, 47

Cladocarpus flexilis

Verrill, 1885

Cape Cod S to Cape Hatteras AM 50-285m P mostly deeper waters 4

Cladocarpus septatus

Nutting, 1900

Cape Cod S to Cape Hatteras AM 87-787m P mostly deeper waters 4

Gymnangium montagui

(Billard, 1912)

Ireland, W Scotland S to Morocco, twice from

South Africa

EU 60-80m algae, shells, rock P 2, 8, 10, 32

Lytocarpia myriophyllum

(Linnaeus, 1758)

Arctic S to Guinea Bissau, Liberia; S to New

England, SE Falklands, Magellan Strait;

widely reported from Indo-Pacific: Chile,

Borneo, Japan

AMPHI 30m-1600m gravel to silt P different subspecies may be involved.

Records from warm waters of the Indo-

Pacific are highly suspect (47)

2, 3, 4, 6, 8,

10, 32

Streptocaulus dollfusi

Billard, 1924

Gibraltar EU deep? P 2, 10

Streptocaulus pectiniferus

Allman, 1883

Iceland, Azores, Canaries, Portugal, Gibraltar,

Mediterranean

EU 92-1646m P 2, 3, 10

Page 14: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Blackfordiidae

Blackfordia manhattensis

Mayer, 1910

NW Atlantic AM shallow M 47, 48

Bonneviellidae

Bonneviella grandis

(Allman, 1876)

Arctic. W Greenland, Barents Sea, Bering

Sea, Sea of Okhotsk, Japan

ARCT 25-800m P 3

Campanulariidae

Campanularia crenata

Allman, 1876

Arctic Atlantic and Pacific. W and S

Greenland, not Iceland, Iberian Peninsula,

Mediterranean

ARCT shallow - deeper on Halecium, ascidans M 2, 3, 4, 10,

17, 34

Campanularia

groenlandica

Levinsen, 1893

Arctic. N Canada S to Cape Cod, Greenland,

Iceland, Faroes, N Norway, Spitsbergen,

Barents Sea, White Sea, Kara Sea, Laptev

Sea, Sea of Okhotsk, Sea of Japan, Bering

Sea, Alaska

ARCT shallow - deep? on other hydroids P 3, 4, 6, 34

Campanularia volubilis

(Linnaeus, 1758)

Arctic, Iceland, Greenland, S to British Isles,

English Channel, Mediterranean, N Spain,

once Mauritania, Canada, New England, S to

Cape Hatteras, N Russia, Sea of Okhotsk, Sea

of Japan, Bering Sea, Alaska, Washington,

California

AMPHI 25-650m hydroids P probably more widespread but

overlooked, rafting on algae

2, 3, 4, 6, 8,

10, 17, 32,

33, 34

Clytia gracilis

(Sars, 1850)

Possibly near cosmopolitan. Barents Sea,

Iceland, S to Morocco, Mediterranean, Arctic

S to Caribbean, Tierra del Fuego, Brazil,

Alaska S to Vancouver Island, South Africa,

India, Mergui Archipelago. Pelagic form

found along W Africa, North Sea

AMPHI 1- 1443m sand, man made hard

substrata, algae, Sabella

tubes, ascidians, coral,

floating substrata,

planktonic

M not all records reliable, identification

difficult. rafting

2, 3, 4, 7, 10,

17, 32, 33

Clytia hemisphaerica

(Linnaeus, 1767)

near cosmopolitan. Iceland, Bergen S, at least

to Iberian Peninsula, Arctic S to Caribbean

AMPHI intertidal -

150m

M: coastal plankton; H:

wide variety of substrata,

probably more common on

algae than C. gracilis, on

fish and their crustacean

ectoparasites, ships

M near cosmopolitan, but identification

difficult, tolerates brackish water. Rafting

on Sargassum. Ship fouling

2, 3, 4, 6, 7,

8, 10, 11, 13,

17, 32, 33,

34

Clytia islandica

(Kramp, 1919)

N North Sea, Shetlands, Faroes, Iceland,

British Isles

EU M hydroid unknown, possibly more wide-

spread

2, 32

Clytia kincaidi

(Nutting, 1899)

Chesapeake Bay to Caribbean AM shallow wood debris, oyster shells M 4, 13

Clytia linearis

(Thornely, 1899)

circumtropical. Mediterranean, Spanish

Atlantic coast, possibly further N

EU shallow rocky shores M introduced. Lessepsian migrant, now very

abundant in Mediterranean. Has a

possible resting stage. rafting inferred

from distribution

2, 10, 17, 35,

36

Clytia longicyatha

(Allman, 1877)

Cape Cod to Caribbean AM shallow - deep on Sargassum, sponges M rafting on Sargassum 4, 17

Page 15: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Clytia noliformis

(McCrady, 1859)

W Atlantic AM shallow on sargassum warm water species, occurs further N on

floating Sargassum

47

Gonothyraea hyalina

Hincks, 1866

Europe, Sargasso Sea EU shallow - deep P subspecies of G. loveni?

rafting on plastics

8, 17

Gonothyraea loveni

(Allman, 1859)

Arctic S to South Carolina, S to Morocco,

Mediterranean, Baltic, New Zealand,

Australia, Cape Town docks (South Africa)

AMPHI intertidal - 30m,

200m

rock and stones, algae,

hydroids, mollusk shells,

seagrasses, ships

P cryptogenic. tolerant of brackish water.,

seasonal dormancy. candidate for ship-

aided dispersal, see South African record.

rafting on plastics. ship fouling

2, 3, 4, 7, 8,

10, 11, 13,

17, 32, 33,

34

Laomedea amphora

(Agassiz, 1862)

Arctic S to Cape Hatteras AM shallow fouling, ships P ship fouling 4, 11, 34

Laomedea angulata

Hincks, 1861

British Isles, S to NW Spain, including The

Netherlands, Mediterranean

EU intertidal - 8m seagrass (Zostera,

Posidonia), ships

P Disappeared from British Isles with

disappearance of eelgrass, nowhere

outside Europe. Records outside Europe

unreliable, not seen on American coast.

rafting on plastics, ship fouling

2, 4, 10, 11,

17, 32, 33

Laomedea exigua

Sars, 1857

British Isles, Belgium, Bergen (Norway) EU 0-100m, true

limits unknown

tunicates, hydroids,

zoophytes, under stones

P not reported for 120 years, recently

redescribed

2, 32

Laomedea flexuosa

Alder, 1857

Arctic S to Mauritania, Ghana, S to Cape

Hatteras, Caribbean.

AMPHI intertidal - 40m,

100m

rock and stones, fucoid

algae, man-made hard

substrata, mud, hydroids,

mollusc shells, crustaceans,

seagrasses. Rafting.

P rafting inferred, fouling 2, 3, 4, 6, 8,

10, 17, 31,

32, 33

Laomedea neglecta

Alder, 1856

Arctic. Iceland, British Isles, Denmark S to

Mediterranean, S to Cape Hatteras

AMPHI intertidal - 50m beneath stones, on shell

gravel, stones, rock, oyster

shells, in muddy conditions

P tolerant of low salinity, used to be present

in the Zuiderzee (The Netherlands)

2, 3, 4, 32

Laomedea

pseudodichotoma

Vervoort, 1959

Bay of Biscay, Gibraltar EU deep? P 2, 10, 38

Obelia fimbriata

(Dalyell, 1848)

English Channel, Mediterranean EU M 2

Obelia geniculata

(Linnaeus, 1758)

near cosmopolitan. Arctic S to Iberian

Peninsula, Mediterranean, S to Caribbean, S

to Kerguelen, S Georgia, Macquarie Island,

New Zealand

AMPHI intertidal -

100m,

commonly 20m

fouling, ships, algae,

Laminaria, Fucus, dogfish,

parisitic copepods on fish,

rafting.

M near cosmopolitan, also in brackish

waters. possibly introduced to

Massachusetts. Rare or absent in tropical

waters. rafting. ship fouling

2, 3, 4, 7, 10,

11, 17, 32,

33, 34, 37

Obelia longissima

(Pallas, 1766)

near cosmopolitan. Arctic S to Iberian

Peninsula, Mediterranean, S to Caribbean,

Indo Pacific?

AMPHI intertidal - 75m fouling, plants, inert

substrata, ships, rock, sand,

rafting

M produces resting stages. distribution Indo-

Pacific unclear, records under other

names probably refer to this species.

Confusion with O. dichotoma recently

resolved.

rafting. ship fouling.

2, 3, 4, 8, 10,

11, 17, 32,

33, 34, 39,

44

Obelia plicata

Hincks, 1868

Hudson Bay ARCT lower shore M 4, 34, 47, 48

Page 16: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Obelia racemosa

Fraser, 1941

Arctic Canada ARCT 90-120m M 4

Orthopyxis integra

(MacGillivray, 1842)

near cosmopolitan; all oceans tropics to Arctic AMPHI intertidal - edge

shelf

rock and stones, algae,

hydroids, mollusk shells,

floating substrates,

seagrasses. Rafting

M near cosmopolitan, not in brackish waters.

rafting inferred

2, 3, 4, 8, 10,

17, 32, 33,

34

Rhizocaulus verticillatus

(Linnaeus, 1758)

Arctic S to Brittany, Bay of Biscay, Black Sea

(not in Mediterranean); S to Long Island

Sound, S to California, Sea of Japan

AMPHI 50-200m,

extremes: 15-

680m

sandy and rocky grounds,

on algae, pebbles, shells,

ships

P ship fouling 2, 3, 4, 7, 8,

11, 32, 33,

34

Campanulinidae

Calycella hispida

(Nutting, 1896)

S England, N Norfolk, offshore EU intertidal -

offshore

P doubtful species 8, 32

Calycella syringa

(Linnaeus, 1767)

near cosmopolitan. Arctic S to Mediterranean,

S to Long Island Sound, Japan, California

AMPHI 10-1500m hydroids, algae, mussels,

bryozoa, barnacles,

floating substrata, common

on stranded material

P

2, 3, 4, 7, 8,

10, 32, 33,

34, 47

Cuspidella costata

Hincks, 1868

Arctic Canada, Cape Cod to Cape Hatteras AM shallow 4

Cuspidella humilis

(Hincks, 1866)

Arctic. Europe, Bay of Biscay,

Mediterranean, Cape Cod to Florida

AMPHI shallow bryozoa, pilings, ships M doubtful species, is probably Staurophora

mertensi. Cuspidella species are only

distinctive in their medusa phase, difficult

to identify. ship fouling

2, 3, 4, 10,

11, 34

Cuspidella procumbens

Kramp, 1911

Greenland, NE Canada ARCT 3, 34

Cuspidella sp.

? intertidal-

considerable

depth

various substrata M not included in analyses, species complex 2, 8, 27, 40

Egmundella grimaldii

Leloup, 1940

Bay of Biscay, Mediterranean EU P 2, 10

Lafoeina maxima

Levinsen, 1893

Arctic to sub-Arctic, possibly S to British

Isles, S to Cape Cod, Sea of Japan

AMPHI shallow-deep P = L. vilaevelebiti , Keratosum

complexum,. seasonal dormancy

3, 4, 32, 34

Lafoeina tenuis

Sars, 1874

E Greenland, N Norway, Barents Sea, S to

Skagerrak, Roscoff, Bay of Biscay, Madeira,

Azores. Mediterranean.

EU coastal - 610m hydroids, bryozoans,

Sargassum, oil rig, fouling

M rafting inferred from distribution, fouling 3, 10, 17, 32,

33, 41

Opercularella panicula

(Sars, 1874)

Atlantic, Pacific and Indian Oceans, including

the Mediterranean

AMPHI 30-2100m hydroids, mollusk shells,

worm tubes, coral, other

cnidarians, brachiopods

M cold water species that submerges in S

part of its range

2, 3, 10, 32,

33

Page 17: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Tetrapoma

quadridentatum

(Hincks, 1874)

Arctic circumpolar. Greenland, Canada,

Spitsbergen, Barents Sea, White Sea, Kara

Sea, Sea of Okhotsk, New Siberian Islands,

Chile

ARCT shallow-deeper 2, 3, 4, 34

Dipleurosomatidae

Dipleurosoma typicum

Boeck, 1866

Probably boreal circumpolar. H: only from

The Clyde. M: British Isles, Faroes, Norway,

Newfoundland, SW Canada, NW America,

Japan.

ARCT M hydroid stage not well known, only

recorded once

32

Eirenidae

Eirene viridula

(Péron & Lesueur, 1810)

M: British Isles, Denmark, N France, Bay of

Biscay, Mediterranean, W Africa, Djibouti,

Kuwait, Amazon, Bismarck Sea, Papua New

Guinea

EU coastal? M hydroid stage never observed in nature 2, 10, 32,

Eutima gracilis

(Forbes & Goodsir, 1851)

N North Sea, Skagerrak, Kattegat S to

Mediterranean. Twice from China?

EU M: coastal

plankton, H:

unknown

M hydroid not well known. Recorded from

China?

2, 10, 32

Eutonina indicans

(Romanes, 1876)

Iceland, mid Norway S to S North Sea, Baltic,

Alaska S to California, Kamchatka, Japan,

India. H: only known from California

EU M: coastal

plankton, H:

shallow

Zostera, crab, rock M hydroid not known from Europe 3, 32, 33

Helgicirrha schulzei

Hartlaub, 1909

Denmark, British Isles, S to Roscoff (France),

Portugal to Mediterranean, Congo, SW Africa

EU M: coastal

plankton, H:

coastal - deep?

mud M 32

Tima bairdii

(Johnston, 1833)

North Sea, but possibly wider range EU M: coastal

plankton,

possibly open

ocean as well

unknown M extremely long medusa stage (up to 7

months). North American T. formosa

possibly conspecific

32

Eucheilotidae

Eucheilota maculata

Hartlaub, 1894

M: S North Sea, Baltic, English Channel,

once off SW India and once off Argentina. H:

Helgoland, The Netherlands, Belgium, Iberian

Peninsula, Mediterranean

EU M: coastal

plankton, H: 4-

15m

crab, ascidians, algae,

hydroids

M 10, 32

Haleciidae

Page 18: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Halecium arcticum

Ronowicz & Schuchert,

2007

Spitsbergen, NE Canada ARCT 5-40m laminarians, hydroids,

ascidians, barnacles, rock

P 42

Halecium beanii

(Johnston, 1838)

near cosmopolitan. Arctic S to Iberian

peninsula, S to Cape Hatteras

AMPHI 5-100m,

sometimes

deeper (430m)

hydroids, shells, rocky

substrates, ships

P near cosmopolitan. H.scutum included in

this species. not reported rafting. ship

fouling

2, 3, 4, 6, 8,

10, 11, 32,

33

Halecium corrugatum

Nutting, 1899

Arctic ARCT P 2, 4

Halecium curvicaule

Lorenz, 1886

Arctic. Greenland, Canada, Iceland, Barents

Sea, White Sea, Kara Sea, Laptev Sea

ARCT 15-500m P 2, 3, 4, 34

Halecium diminutivum

Fraser, 1940

Bay of Fundy S to Cape Hatteras AM shallow P 4

Halecium groenlandicum

Kramp, 1911

Arctic. Greenland, E Canada, N Russia,

Bering Sea, Sea of Okhotsk, Kuriles

ARCT 50-100m hard bottoms P 2, 3, 34

Halecium halecinum

(Linnaeus, 1758)

Cosmopolitan. Arctic S to South Africa, S to

Chesapeake Bay, Mediterranean, Alaska,

Puget Sound, California, Sea of Okhotsk,

Japan, Moluccas

AMPHI subtidal - shelf

edge,

sometimes

deeper

stones, shells, other hard

substrates, ships

P not reported rafting, ship fouling 2, 3, 4, 6, 7,

8, 10, 11, 32,

33, 34,

Halecium labrosum

Alder, 1859

Arctic to S to Galicia, Azores, Mediterranean;

S to Cape Hatteras, Alaska, Japan

AMPHI 5-200m epizoic, perhaps inanimate

substrata

P 2, 3, 4, 8, 10,

32, 34

Halecium liouvillei

Billard, 1934

NW Spain, Mediterranean EU P 2, 10

Halecium macrocephalum

Allman, 1877

Cape Cod S to Florida AM shallow-deep wharf piling P fouling 4

Halecium mediterraneum

Weissman, 1883

Iberian Peninsula, Mediterranean.

cosmopolitan?

EU P 10, 2

Halecium minutum

Broch, 1903

Arctic. Nova Scotia, Greenland, Iceland,

Norway, Murmansk coast, Bering Sea, S to

Cape Cod

ARCT 15-450m P H. corrugatum might be synonym 2, 3, 4, 34

Halecium muricatum

(Ellis & Solander, 1786)

Arctic S to English Channel, Bay of Biscay, S

to Cape Hatteras, Alaska, Siberia

AMPHI 10-1350m rocks, shells, algae P 2, 3, 4, 6, 8,

10, 32, 33,

34

Halecium plumosum

Hincks, 1868

British Isles EU P questionable synonym of H. sessile 8, 32

Halecium pusillum

(Sars, 1857)

NW Spain, Mediterranean. tropical Atlantic? EU P planktonic propagule: short length of

colony that is buoyant

2, 10, 33

Page 19: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Halecium sessile

Norman, 1867

near cosmopolitan. S to Cape Cod, Senegal,

Mediterranean, Indo-West Pacific, Australia,

New Zealand

AMPHI intertidal - edge

shelf (430m)

Sabellaria tubes, algae,

other substrata

P taxonomic and identification difficulties 2, 3, 4, 6, 8,

10, 38

Halecium sibogae

Billard, 1929

Gibraltar EU P Synonym of Zygophylax sibogae Billard,

1918?

2, 10

Halecium speciosum

Nutting, 1901

Arctic ARCT P 34

Halecium textum

Kramp, 1911

Greenland, NW Canada, W Iceland, Faroes.

Possibly North Sea, Sweden, Norway, British

Isles

AMPHI 46-207m on other hydroids P H. undulatum records from the North Sea

area could also be this species

3, 45

Halecium undulatum

Billard, 1921

N Norway S to off France, Canadian E coast,

Arctic

AMPHI inshore - 220m hydroids, bryozoans, whelk

egg capsules, spider crab,

algae, swimming

vertebrates, ship hull

P probably more widespread than reported,

due to confusion with H. tenellum

hull fouling, not reported rafting

32, 33, 34

Hydranthea margarica

(Hincks, 1863)

Shetlands S to Bay of Biscay, Mediterranean,

Seychelles

EU offshore rock, Laminaria,

bryozoans, mollusks,

hydroids

M "undoubtedly overlooked"

not reported rafting

2, 8, 10, 32,

33

Hydrodendron mirabile

(Hincks, 1866)

British Isles, NW France, Iberian Peninsula,

Cape Verdes, South Africa, mid S Atlantic

Ocean, W Indies, SW Indian Ocean, New

Zealand. Circumtropical?

EU intertidal - 65m algae, especially

Laminaria, hydroids,

bryozoans

P 2, 10, 32, 33

Halopterididae

Antennella siliquosa

(Hincks, 1877)

Bay of Biscay EU P 2, 10

Halopteris diaphana

(Heller, 1868)

Iberian Peninsula, Mediterranean EU P 2, 10, 17

Halopteris liechtensternii

Marktanner-Turneretscher,

1890

Gibraltar, Mediterranean EU P 2, 10

Halopteris tenella

(Verrill, 1874)

Massachusetts to Caribbean AM shallow among ascidians, on docks,

piles

P seasonal dormancy, fouling 4, 13, 43

Kirchenpaueriidae

Kirchenpaueria

bonnevieae

(Billard, 1906)

Norway, Faroes, Shetland, Iceland, Bay of

Biscay, Morocco, Mediterranean, South

Africa, Oman, Zanzibar, New Zealand, Japan

EU shallow on other hydroids P 2, 3, 10

Page 20: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Kirchenpaueria pinnata

(Linnaeus, 1758)

Iceland, Faroes, Trondheim S to Morocco,

Mediterranean, South Africa, especially SW

Cape, Japan

EU intertidal -

100m, 350m

rock and stones, mollusks,

crustaceans, algae,

eelgrass, artificial wooden

structures

P introduced to South Africa by shipping.

rather variable species, several nominal

species are considered synonymous,

discussion ongoing. rafting inferred, ship

fouling

2, 3, 8, 17,

32, 33

Kirchenpaueria similis

(Hincks, 1861)

N Europe to Mediterranean, Azores EU intertidal -

100m

wide variety plant, animal

and inert substrata;

mollusks, crustaceans,

algae, eelgrass, artificial

wooden structures

P doubtful species, considered conspecific

with K. pinnata by many authors. rafting

inferred

17, 32

Lafoeidae

Acryptolaria crassicaulis

(Allman, 1888)

Iberian Peninsula. Cosmopolitan? EU P 2, 10

Acryptolaria triserialis

(Fraser, 1913)

Boreal Atlantic Canada AM 36m P 4

Anthohebella parasitica

(Ciamician, 1880)

Iberian Peninsula, Mediterranean EU parasite? M 2, 10

Bedotella armata

(Pictet & Bedot, 1900)

Iberian Peninsula, Mediterranean.

cosmopolitan?

EU P 2, 10

Cryptolaria pectinata

(Allman, 1888)

British Isles, Iberian Peninsula,

Mediterranean, tropical Atlantic?

EU P 2, 10

Filellum serpens

(Hassall, 1848)

cosmopolitan, tropics to polar seas AMPHI sublittoral to

edge of shelf

epizoic, hydroids, shells,

inert substrata, ships

P rafting inferred, ship fouling 2, 3, 8, 10,

11, 17, 32,

33, 34

Filellum serratum

(Clarke, 1879)

Iceland, NW Atlantic, Iberian Peninsula,

Mediterranean, Indian Ocean, New Zealand,

Chile, South Africa

AMPHI 5-900m P rafting inferred 2, 3, 10, 17

Grammaria abietina

(Sars, 1850)

circumpolar, S to North Sea, S to New

England, Azores. Not in English Channel,

Crozet islands, Falklands, Tierra del Fuego,

Patagonia, Argentina

AMPHI 10-1500m silty and rocky substrates P 2, 3, 4, 8, 32,

33, 34

Grammaria borealis

(Levinsen, 1893)

North Atlantic, Nova Scotia, Labrador, W

Greenland, Iceland, Barents Sea, Kara Sea,

Japan

ARCT 36-314m 3, 34

Grammaria gracilis

Stimpson, 1854

Arctic S to Cape Cod, Cape Hatteras S to

Florida

AM shallow Laminaria P 4

Grammaria immersa

Nutting, 1901

Arctic. Greenland, Bering Sea, Faroes,

Spitsbergen, Iceland, Japan

ARCT shallow - deep 2, 3, 34

Hebella scandens

(Bale, 1888)

Iberian Peninsula, Mediterranean.

cosmopolitan?

EU M 2, 10

Page 21: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Lafoea dumosa

(Fleming, 1820)

near cosmopolitan. Iceland, Spitsbergen,

Faroes, British Isles, Iberian Peninsula, North

Sea, Brittany, France offshore,

Mediterranean, Arctic to Caribbean

AMPHI subtidal - deep

(2078m,

3940m)

rock and stones, algae,

hydroids

P L. gracillima and L. fruticosa included

rafting

2, 3, 4, 6, 7,

8, 10, 17, 32,

33, 34

Scandia gigas

(Pieper, 1828)

Bay of Biscay, Mediterranean EU P 2, 10

Zygophylax biarmata

Billard, 1905

Iberian Peninsula, Mediterranean EU P cosmopolitan? 2, 10

Zygophylax brownei

Billard, 1924

Iberian Peninsula, Mediterranean EU P 2, 10

Zygophylax crassitheca

(Fraser, 1941)

Bay of Fundy to Cape Cod AM 30-327m P 4, 48

Zygophylax elegantula

Leloup, 1940

Bay of Biscay EU P possibly synonym of Z. levinseni 2, 10

Zygophylax levinseni

(Saemundsson, 1911)

Bay of Biscay EU P 2, 10

Zygophylax sibogae

Billard, 1918

Bay of Biscay, Indo Pacific? EU P 2, 10

Laodiceidae

Laodicea undulata

(Forbes & Goodsir, 1853)

cosmopolitan. Iceland S to South Africa,

Tierra del Fuego, Mediterranean, Black Sea,

Adriatic Sea, China Sea

EU coastal - shelf M 2, 10, 32

Modeeri rotunda

(Quoy & Gaimard, 1827)

Boreal to tropical Atlantic and Indo-Pacific,

Barents Sea S to Patagonia, W Africa and off

Antarctica

AMPHI M: deep sea

oceanic, H: 30-

1214m

hydroids M 2, 3, 4, 8, 10,

32

Staurophora mertensii

Brandt, 1838

Bipolar, Arctic S to North Sea, S to Cape

Cod, Japan, Alaska, Falklands, Orkneys

AMPHI M: 0-150m, H:

coastal

M 2, 6, 32

Lovenellidae

Lovenella grandis

Nutting, 1901

Cape Cod S to Cape Hatteras AM 4

Lovenella producta

(Sars, 1874)

Arctic S to Bergen; Gulf of Maine; NW

America

AMPHI 6-2000m hydroids, ascidians, worm

tubes, coral

3, 4, 32, 33

Mitrocomium cirratum

Haeckel, 1879

Bay of Biscay, Mediterranean. tropical

Atlantic?

EU M 2, 10

Page 22: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Malagazziidae

Octophialucium

funerarium

(Quoy & Gaimard, 1827)

Bay of Biscay, Mediterranean EU M 2, 10

Mitrocomidae

Cosmetira pilosella

(Forbes, 1848)

NE Atlantic from Scotland, Bergen S to

Iberian Peninsula.

EU M: planktonic,

coastal to

offshore, H: ?

hard inert substrates M hydroid poorly known, medusa has been

found in the central Atlantic ocean, close

to the Azores

2, 10, 32

Halopsis ocellata

Agassiz, 1863

Arctic S to North Sea, Cape Cod. Falkland

Islands

AMPHI deep? M 32

Mitrocomella brownei

(Kramp, 1930)

British Isles, North Sea, Brittany EU M: coastal

plankton

sea fan Eunicella M hydroid stage not well known 32

Mitrocomella

polydiademata

(Romanes, 1876)

Arctic S to North Sea, British Isles, Kattegat;

S to Gulf of Maine, Washington State

AMPHI coastal M 15

Phialellidae

Opercularella lacerata

(Johnston, 1847)

White Sea, Greenland, Iceland S to

Mediterranean, Baltic Sea, S to Cape

Hatteras, Indo-Pacific?

AMPHI sublittoral - 20,

50m

kelp, mussels, ascidians,

bryozoans, hydroids, other

substrates, ships

P cryptogenic. tolerant of reduced salinity,

not reported rafting, ship fouling

2, 3, 4, 10,

11, 32, 33,

34, 44, 49

Phialella quadrata

(Forbes, 1848)

Sweden, Denmark, British Isles, The

Netherlands, Belgium, N France,

Mediterranean, Sea of Japan, New Zealand,

Chile, Morocco, Gulf of Guinea

EU M: coastal

plankton, H:

intertidal -

shallow

various substrata M 2, 8, 32

Plumulariidae

Monotheca obliqua

(Johnston, 1847)

near cosmopolitan in temperate to tropical

waters, not in NW Atlantic

EU intertidal - 15m weed, sponges, in pools,

floating Sargassum in mid

Atlantic

M rafting on algae 2, 8, 10, 17,

32, 43, 47

Monotheca pulchella

Bale, 1882

Gibraltar, Mediterranean EU deep? P 2, 10

Nemertesia americana

(Nutting, 1900)

Maine to Florida AM 38-680,

offshore

P 4, 6

Nemertesia antennina

(Linnaeus, 1758)

Arctic S to South Africa, Mediterranean,

Azores, Madeira, S to Florida, not Caribbean,

Japan, New Zealand?

AMPHI inshore - deeper

water (~400m)

shells, spider crabs, sandy

bottoms, auto-epizootic

P 2, 3, 4, 8, 10,

32, 34, 38

Nemertesia falcicula

Ramil & Vervoort, 1992

Gibraltar, NE Atlantic, Mediterranean EU deep? P 2, 10

Page 23: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Nemertesia norvegica

(Sars, 1874)

S Iceland, Mid Norway, British Isles, North

Sea, Faroes, Bay of Biscay, Mediterranean

EU 80-800m,

offshore

unknown P probably widespread but unrecorded 3, 32

Nemertesia ramosa

Lamouroux, 1816

Arctic S to NW Africa, Mediterranean,

Azores, Madeira, Canaries, South Africa,

Indian Ocean?

EU 30m- 1182m shells, spider crabs, sandy

bottoms

P records from Indian Ocean probably

distinct species

2, 3, 8, 10,

32, 38

Nemertesia rugosa

(Nutting, 1900)

Nantucket, Massachusetts AM 80m 4, 47

Nemertesia

ventriculiformis

(Marktanner-

Turneretscher, 1890)

Bay of Biscay, Gibraltar, Mediterranean EU P 2, 10

Plumularia floridana

Nutting, 1900

North Carolina to Caribbean AM shallow - deep rafting on Sargassum P warm water species, but occurs N to

offshore Virginia on floating Sargassum

2, 4, 13, 17

Polyplumaria flabellata

Sars, 1874

S Iceland, Norway S to Congo, Azores, Mid

Atlantic Ridge, Gibraltar

EU 30->1378m fouling P probably widespread, but records are few

and patchy

2, 3, 8, 10,

31, 32, 38

Polyplumaria gracillima

(Sars, 1873)

Arctic, Canada S to New England, Barents

Sea, Norway, Iceland, Skagerrak, Kattegat

AMPHI 40-700m dead coral, stones, sponge P 2, 3, 4, 6, 32

Pseudoplumaria marocana

(Billard, 1930)

Gibraltar, Mediterranean EU P 2, 10

Schizotricha frutescens

(Ellis & Solander, 1786)

Iceland, Faroes S to Morocco, Mediterranean.

single records from South Africa and

Kerguelen need confirmation

EU 20-90m rocks and stones

exclusively

P 2, 3, 8, 10,

32, 33

Sertulariidae

Abietinaria abietina

(Linnaeus, 1758)

Arctic S to Madeira, Iberian Peninsula,

Mediterranean, S to Cape Hatteras, S to

California and Japan, Madagascar

AMPHI 10m - 630m shells and stones and

similar substrates on sandy

bottoms

P 2, 3, 4, 8, 10,

32, 33, 34

Abietinaria filicula

(Ellis & Solander, 1786)

Arctic S to British Isles, S to Cape Cod,

higher latitudes NW America

AMPHI 5-656m, mostly

shallower than

50m

P European range seems to have retracted

North recently

2, 8, 32, 3, 4,

34

Abietinaria fusca

(Johnston, 1847)

Barents Sea, Kola Peninsula, Norway, British

Isles, Faroes, Iceland

EU 40-200m hydroids, stones P 2, 3, 8, 32

Abietinaria kincaidi

(Nutting, 1901)

Arctic Canada ARCT 0-15m P 4

Abietinaria pulchra

(Nutting, 1904)

Arctic. Vancouver Island, Bering Sea, Sea of

Okhotsk, Chukchi Sea, E Siberian Sea,

Laptev Sea, Kara Sea, White Sea, Barents

Sea, Spitsbergen, W Greenland, N Canada

ARCT 123-772m P 2, 3, 4, 34

Page 24: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Abietinaria thuiarioides

(Clark, 1877)

Arctic. N Canada, Iceland. Alaska, Bering

Sea, Sea of Okhotsk, Japan Sea, Barents Sea,

Kara Sea Laptev Sea, Chukchi Sea

ARCT P 3, 4

Abietinaria turgida

(Clark, 1877)

Arctic Canada ARCT 2-46m P 4, 34

Amphisbetia operculata

(Linnaeus, 1758)

Shetlands, British Isles, S to Africa,

Mediterranean, Australia, New Zealand,

Argentina, Patagonia, Chile, Java, Pacific

coast N America

EU intertidal -

>70m

brown algae, Laminaria M not in W Atlantic.

rafting inferred from distribution

2, 8, 10, 32

Diphasia attenuata

(Hincks, 1866)

Jan Mayen Island, North Sea, British Isles, S

to Morocco, possibly Azores, Mediterranean,

New Caledonia deep water

EU sublittoral -

1470m

hydroids P possibly conspecific or confused with D.

rosacea

2, 3, 8, 10,

32

Diphasia delagei

Billard, 1912

English Channel, NW France, W of Strait of

Gibraltar

EU 60-1250m hydroids, stones, shells P 10, 2, 32

Diphasia fallax

(Johnston, 1847)

Arctic S to North Sea, S to deep waters off

South Carolina

AMPHI 20-250m hydroids P 2, 3, 4, 8

Diphasia margareta

(Hassall, 1841)

Iberian Peninsula, Mediterranean EU P 2, 10

Diphasia nigra

(Pallas, 1766)

English Channel, S England, N France, Bay

of Biscay

EU 60m, usually

>80m

mussels, other substrata P deeper waters 2, 8, 10, 32

Diphasia pinaster

sensu Hincks, 1868

Arctic Canada, Iceland, Bergen S to Morocco,

Azores, Cape Verdes, Guinea,

Mediterranean?

AMPHI 75-1318m bryozoans, coral, stones,

other substrata

P 2, 3, 4, 8, 10,

32

Diphasia rosacea

(Linnaeus, 1758)

Arctic S to Gibraltar, S to Massachusetts Bay AMPHI intertidal - 80m Laminaria zone on range

of substrata

P possibly conspecific with D. attenuata 2, 8, 32, 3,

10, 4, 34

Dynamena pumila

(Linnaeus, 1758)

Arctic to Iberian Peninsula, S Labrador to

New Engand. Baltic, Mediterranean?

AMPHI intertidal - 5m,

occasionally

deeper

algae, rocks, seagrasses,

sheltered to exposed

shores, rafting

P tolerant of reduced salinity. Seasonal

dormancy. S records on American coast

are highly doubtful

Rafting on algae

2, 3, 4, 7, 8,

17, 32, 33,

34

Dynamena quadridentata

(Ellis & Solander, 1786)

Cape Cod to Caribbean, Australia AM shallow on floating Sargassum P Warm water species. Not in Europe (2)

rafting on Sargassum

2, 4, 17

Hydrallmania falcata

(Linnaeus, 1758)

Arctic S to Bay of Biscay, Long Island

Sound, N Pacific? Deep water New Caledonia

AMPHI 10-1102m hard substrata on sandy

bottoms

P Closely related species, perhaps

conspecific, present on Pacific coast of

Canada and California.

not reported rafting

2, 3, 4, 6, 7,

8, 10, 32, 33

Salacia desmoides

(Torrey, 1902)

Iberian Peninsula, Mediterranean, US Pacific

coast, Indo Pacific?

EU P 2, 10

Sertularella cylindritheca

(Allman, 1888)

Iberian Peninsula, Mediterranean EU P tropical Atlantic 2, 10

Page 25: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Sertularella ellisii

(DesHayes & Milne-

Edwards, 1863)

Iberian Peninsula, English Channel,

Mediterranean

EU P 2, 10

Sertularella gaudichaudi

(Lamouroux, 1824)

Spitsbergen to Cape Verdes, Canadian Arctic,

South Africa, California, Australasia, Indian

Ocean, Argentina

AMPHI intertidal -

shallow

P confused taxonomy, distribution

uncertain

rafting inferred

2, 4, 17, 32,

34

Sertularella gigantea

Mereschowsky, 1878

Arctic S to Cape Hatteras AM 30-140m P 4, 48

Sertularella mediterranea

Hartlaub, 1901

Arctic, British Isles, Iberian Peninsula,

Mediterranean

EU intertidal,

coastal

inert substrata P 2, 8, 10

Sertularella polyzonias

(Linnaeus, 1758)

Arctic S to Angola, South Africa, S to

Georgia, Alaska, Bering Sea, Japan,

Kerguelen

AMPHI intertidal - 50m,

sometimes

300m

shells, algae, rock,

hydroids

P possibly conspecific or confused with S.

gayi. Intertidal colonies may remain

infertile. South African endemic

subspecies

2, 3, 4, 6, 7,

8, 10, 32, 33,

34

Sertularella rugosa

(Linnaeus, 1758)

Arctic S to W France, S to Cape Cod, New

York, Alaska to Washington State, California

AMPHI intertidal-263m bryozoans, hydroids, algae,

man made hard substrata

P fouling 2, 3, 4, 6, 8,

32, 33

Sertularella tenella

(Alder, 1857)

Near cosmopolitan. Arctic S to W France,

Bay of Biscay, Angola, S to Caribbean, S to

Japan, S to California.

AMPHI 10-150m, ca.

1000m

hydroids P Mediterranean records doubtful 2, 3, 4, 8, 10,

32, 34

Sertularia argentea

Linnaeus, 1758

Arctic S to Bay of Biscay, S to Cape Hatteras,

S to California

AMPHI shallow pebbles, shells on sandy,

muddy bottoms, algae

P possibly conspecific with S. cupressina,

but shallow, might be two forms of the

same species

not reported rafting

3, 4, 6, 32,

33, 34

Sertularia cupressina

Linnaeus, 1758

Arctic S to Bay of Biscay, S to New Jersey, S

to California

AMPHI intertidal -

100m,

sometimes

deeper

pebbles, on sandy bottoms,

low shore pools

P seasonal dormancy. possibly conspecific

with S. argentea, but deeper, might be

two forms of the same species

not reported rafting

2, 3, 4, 6, 7,

8, 10, 13, 32,

33, 34

Sertularia dalli

(Nutting, 1904)

Canadian Arctic ARCT P 4, 33, 48

Sertularia fabricii

Levinsen, 1893

Arctic. N Canada to Cape Hatteras,

Greenland, Iceland, Jan Mayen, Kara Sea,

Alaska, Puget Sound

AM 0-293m P possibly conspecific with S. argentea and

S. robusta

2, 3, 4, 34

Sertularia latiuscula

Stimpson, 1854

Arctic S to Virginia AM shallow Laminaria P 4, 6, 7

Sertularia plumosa

(Clark, 1877)

Arctic ARCT shallow P 2, 34

Sertularia robusta

(Clark, 1877)

Arctic S to Cape Hatteras, W Europe AMPHI 0-100m P 2, 4, 34

Sertularia schmidti

Kudelin, 1914

Arctic. White Sea, NE Canada, Greenland ARCT shallow-deeper P 2, 3, 34

Page 26: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Sertularia similis

Clark, 1877

Arctic. N Canada, Iceland, Sea of Okhotsk,

Sea of Japan, Chuchki Sea, Bering Sea,

Alaska. S to New England

ARCT 0-130m P 3, 4, 6, 34

Sertularia tenera

Sars, 1874

Arctic S to Faroes, SW Norway, Kattegat,

Canada, Bering Strait, Alaska, Sea of Japan. S

to Cape Cod

AMPHI sublittoral -

edge of shelf,

<400m

stones, shells P not well known 2, 3, 4, 32,

34

Sertularia (Pericladium)

mirabilis

(Verrill, 1873)

Arctic. N Canada, Greenland, Iceland,

Spitsbergen, Barents Sea, White Sea, Kara

Sea, Sea of Japan, Bering Sea, Alaska

ARCT 15-110m P 3, 4, 34

Symplectoscyphus

pinnatus

(Clark, 1877)

N Canada, Jan Mayen, Sea of Okhotsk,

Bering Sea, Alaska

ARCT shallow P rare species 34

Symplectoscyphus

tricuspidatus

(Alder, 1856)

Circumpolar and widespread in Arctic to

boreal Atlantic and Pacific. S to Scotland,

Tromso, S to New Jersey

AMPHI coastal - >200m animals, Mytilus, not well

known

P 2, 3, 4, 6, 7,

8, 32, 33, 34

Tamarisca tamarisca

(Linnaeus, 1758)

Arctic S to off Brittany, Bay of Biscay, S to

New England. Greenland, Iceland

AMPHI 10m - edge

shelf, typically

>80m off

Brittany

P 2, 4, 6, 8, 10,

32

Thuiaria alternitheca

Levinsen, 1893

Arctic. Davis Strait, Canada, Greenland,

Iceland, Sea of Okhotsk

ARCT shallow - deep? P Synonymized with T. articulata 2, 3, 4, 34

Thuiaria arctica

(Bonnevie, 1899)

Arctic. Spitsbergen, Iceland, Bear Island,

Kola Peninsula

ARCT 38-150m P doubtful species 3

Thuiaria articulata

(Pallas, 1766)

Arctic circumpolar to northern-temperate, S to

Skagerrak, Kattegat, Brittany, Bay of Biscay,

British Isles. S to Cape Hatteras. Cooler

waters S hemisphere

AMPHI 18-300m,

usually 50-

200m

stones, shells P submerges in southern part of range 2, 3, 4, 6, 8,

10, 32, 34

Thuiaria carica

Levinsen, 1893

Arctic. N Russia, Spitsbergen, Faroes,

Iceland, Hudson Bay, British Columbia

ARCT 20-1102m P 3, 4, 34

Thuiaria hartlaubi

(Nutting, 1904)

Arctic? San Juan Archipelago, Kurile Islands,

Bering Sea, Tartar Strait, Kola Peninsula,

Iceland

ARCT 79m P doubtful species, possibly conspecific

with T. arctica and T. cylindrica

3

Thuiaria laxa

Allman, 1874

Arctic. Labrador, Greenland, Iceland, Faroes,

Shetland, Spitsbergen, N Russia, Sea of

Japan, Sea of Okhotsk. S to New England

ARCT 27-92m P 3, 4, 34

Thuiaria sachalini

Kudelin, 1914

Arctic. Sea of Okhotsk, Sea of Japan, Kurile

Islands, W Greenland

ARCT P doubtful speies, possibly a form of T.

alternitheca

3

Page 27: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

Thuiaria thuja

(Linnaeus, 1758)

Arctic S to Portugal, S to New England, S to

Sea of Okhotsk, S to Washington State,

Mediterranean

AMPHI 2-800m shells and similar substrata P often washed up, sometimes from far.

Mediterranean records uncertain.

submerges in southern part of range

2, 3, 4, 8, 10,

32, 34

Syntheciidae

Synthecium evansi

(Ellis & Solander, 1786)

Gibraltar, Mediterranean EU P 2, 10

Tiarannidae

Stegolaria geniculata

(Allman, 1888)

Bay of Biscay. circumtropical? EU P 2, 10

Stegopoma giganteum

Ramil & Vervoort, 1992

Strait of Gibraltar EU M deep-water species? 10, 2

Stegopoma plicatile

(Sars, 1863)

Arctic S to Vancouver Island, S to Florida,

New Zealand, Tasmania, Antarctica, far S

America, Philippines

AM 15-1940m muddy bottoms, rock and

stones, hydroids

M 2, 3, 4, 33,

34, 38

Tiaropsidae

Tiaropsis multicirrata

(Sars, 1835)

Arctic S to Brittany, S to New England; N

Pacific

AMPHI M: shallow

coastal

plankton, upper

9m, H: shallow

Buccinum, hydroids, algae M hydroid stage not well known 6, 32, 33

Limnomedusae

Olindiidae

Monobrachium

parasiticum

Mereschkowsky, 1877

Arctic. N Russia, Sea of Okhotsk, Sea of

Japan, Spitsbergen, W Greenland, S to Nova

Scotia, W Canada, California (doubtful). Not

in Iceland. Iberian Peninsula, Mediterranean

AMPHI shallow (4m) parasitic on bivalve hosts

(Macoma calcaria)

M 2, 3, 4, 10

Dispersal: for species that lack a medusa phase, the planula larva is the dispersive stage, which is indicated by P, dispersal with a

medusa is indicated with M. EU= restricted to European coasts, AM=restricted to Northeast American coasts, ARCT=only in Arctic

or sub-Arctic waters, AMPHI=with a continuous amphi-Atlantic distribution. Habitat: M=medusa, P=polyp. References: 1: (Hansson

1998), 2: (Costello et al. 2004), 3: (Schuchert 2001b), 4: (Fraser 1944), 5: (Schuchert 2006), 6: (Trott 2004), 7: (Calder 1975), 8:

(Hayward & Ryland 1990), 9: (Schuchert 2007), 10: (Medel & López-González 1996), 11: (Woods Hole Oceanographic Institution

1952), 12: (Millard 1975), 13: (Calder 1990), 14: (Calder 1972), 15: (Schuchert 2001a), 16: (Schuchert 2005a), 17: (Thiel & Gutow

2005), 18: (Watling & Maurer 1972), 19: (Bouillon & Boero 2000), 20: (Marques et al. 2000), 21: (Lilly et al. 2002), 22: (Schuchert

Page 28: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

2004), 23: (Bouillon et al. 1997), 24: (Buss & Yund 1989), 25: (Ma & Purcell 2005), 26: (Carlton 1979), 27: (Calder & Burrell 1969),

28: (Piraino et al. 1996), 29: (Miglietta et al. 2007), 30: (Carlton 1998), 31: (Herpin 1935), 32: (Cornelius 1995), 33: (Cornelius

1992), 34: (Calder 1970), 35: (Bouillon et al. 2004), 36: (Lindner & Migotto 2002), 37: (Govindarajan et al. 2005), 38: (Vervoort

1966), 39: (Berman et al. 1992), 40: (Strathmann 1990), 41: (Giulio et al. 2000), 42: (Ronowicz & Schuchert 2007), 43: (Leclère et al.

2007), 44: (Carlton 2003), 45: (Schuchert 2005b), 46: (Schuchert pers. comm.), 47: (Calder pers. comm.), 48: (Integrated Taxonomic

Information System), 49: (Carlton pers. comm.).

Page 29: University of Groningen What is natural? Haydar, Deniz · What is natural? Haydar, Deniz ... Dipurena strangulata McCrady, 1859 Cape Cod to Florida, Puerto Rico, Gulf of Guinea AM

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