Commemorative volume for the 80th birthday of Willem Vervoort in 1997
Further notes on Leptolida (Hydrozoa: Cnidaria) from
Canadian Pacific Waters
A . Brinckmann­Voss & M . N . Arai
Brinckmann­Voss, A. & M . N . Arai. Further notes on Leptolida (Hydrozoa: Cnidaria) from Canadian
Pacific waters.
Zool. Verh. Leiden 323, 31.xii.1998: 37­68, figs 1­12.— ISSN 0024­1652/ISBN 90­73239­68­0.
Anita Brinckmann­Voss, Center for Biodiversity and Conservation Biology, Royal Ontario Museum,
100 Queen's Park, Toronto, O N , Canada M5S 2C6. Mailing address: P.O.B. 653, Sooke, B.C., Canada
V0S 1N0, e­mail: anitab­[email protected].
Mary Needier Arai, Pacific Biological Station, 3190 Hammond Bay Road, Nanaimo, B.C., Canada V9R
5K6, e­mail: [email protected].
Key words: taxonomy; Leptolida; Anthoathecatae; Bythotiaridae; Pandeidae; Trichydridae; Paragotoea;
Paulinum; Euphysa; Tiarannidae.
This paper includes new data on Anthoathecatae and Tiarannidae. Paulinum gen. nov., Paul inum l ineatum spec. nov., and Euphysa vervoorti spec. nov. are described. A single specimen was assigned tenta­
tively to the Trichydridae as incertae sedis gen., incertae sedis spec. nov. A n additional four species
(Calycopsis bigel owi, Merga reesi, Paragotoea bathybia and Modeeria rotunda) are recorded from Canadian
Pacific and adjacent waters for the first time. The species Bythotiara depressa Naumov, 1960 is re­
described and its relationship to other Calycopsidae is discussed. Other data are briefly summarized
to update the Anthomedusae portion of Arai & Brinckmann­Voss (1980:1).
Introduction
T h e inshore f a u n a of the L e p t o l i d a (medusa stage) f r o m Puget S o u n d a n d off Brit­
i s h C o l u m b i a i s f a i r l y w e l l k n o w n ( s u m m a r i z e d i n A r a i & B r i n c k m a n n ­ V o s s , 1980: 1;
M i l l s , 1987: 33). H o w e v e r , there are f e w p u b l i s h e d data o n the deep water f a u n a f r o m
C a n a d i a n a n d adjacent waters of the northeast Pacific O c e a n . Recent collections h a v e
p r o v i d e d specimens of s u c h offshore H y d r o z o a . T h e present p a p e r uses this m a t e r i a l
to u p d a t e the Anthoathecatae a n d T i a r a n n i d a e of o u r p r e v i o u s m o n o g r a p h .
Materials and methods
The majority of the samples w e r e s u p p l i e d b y the g r o u n d f i s h section of the Pacific
B i o l o g i c a l Station, N a n a i m o , B . C . , p a r t i c u l a r l y f r o m cruises J a n u a r y ­ A p r i l 1987 a n d
J a n u a r y ­ A p r i l 1988. A i m
2
T u c k e r s a m p l e r w a s u s e d w i t h three 335 μιη nets w h i c h
w e r e o p e n e d a n d closed at specific depths. Stations o c c u p i e d at this time w e r e i n
three p a r a l l e l r o w s off Pachena P o i n t a n d B a r k l e y S o u n d , west of V a n c o u v e r Island
f r o m inshore to b e y o n d the 1200 m contour l i n e ( A r a i et a l . , 1993: 1). T h e i r earlier,
1980 a n d 1986, samples off the C a n a d i a n west coast w e r e o b t a i n e d b y o b l i q u e b o n g o
t o w s ( F u l t o n et a l . , 1982: 1; A r a i et al., 1993: 2). F o r other samples collected b y o u r ­
selves o r s u p p l i e d b y other w o r k e r s , see the section " M a t e r i a l " of the species accounts
and "acknowledgements".
Specimens w e r e f i x e d i n 5 % f o r m a l d e h y d e s o l u t i o n b u f f e r e d i n s o d i u m borate,
a n d stored i n this s o l u t i o n o r 70% a l c o h o l . P r e l i m i n a r y s o r t i n g w a s done at the Pacific
38
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
B i o l o g i c a l Station, a n d f i n a l sorting at the D e p a r t m e n t of B i o l o g i c a l Sciences, U n i v e r sity of C a l g a r y , C a l g a r y , A l b e r t a .
The material referred to i n this paper w i l l be mostly deposited i n the R o y a l British
C o l u m b i a M u s e u m , Victoria, B.C., C a n a d a . Accession numbers refer to specimens
already deposited; the r e m a i n i n g specimens are still i n our collections for further study.
The p h o t o g r a p h s of Modeeria rotunda were taken b y M . N . A . , a n d a l l other o r i g i n a l
illustrations w e r e done b y A . B . - V .
Abbreviations:
ROM
= R o y a l O n t a r i o M u s e u m , Toronto, O n t . C a n a d a ;
ZMC
= Zoological M u s e u m , Copenhagen, Denmark;
RBCM
= Royal British Columbia Museum,Victoria, B.C. Canada.
ZISTP
= Z o o l o g i c a l Institute St Petersburg, R u s s i a .
L i s t of species
Class H y d r o z o a O w e n , 1843
Subclass L e p t o l i d a H a e c k e l , 1879
O r d e r Anthoathecatae C o r n e l i u s , 1992
Suborder F i l i f e r a Kühn, 1913
S u p e r f a m i l y P a n d e o i d e a H a e c k e l , 1879
F a m i l y Bythotiaridae M a a s , 1905 (= C a l y c o p s i d a e H a r t l a u b , 1913)
G e n u s Bythotiara Günther, 1903
Bythotiara depressa N a u m o v , 1960
Bythotiara huntsmani (Fraser, 1911)
G e n u s Calycopsis Fewkes, 1882
Calycopsis bigelowiVanhöffen,
1911
Calycopsis nematophora B i g e l o w , 1913
F a m i l y H a l i m e d u s i d a e A r a i & B r i n c k m a n n - V o s s , 1980
G e n u s Halimedusa B i g e l o w , 1916
Halimedusa typus B i g e l o w , 1916
F a m i l y Pandeidae H a e c k e l , 1879
G e n u s Amphinema H a e c k e l , 1879
Amphinema platyhedos A r a i & B r i n c k m a n n - V o s s , 1983
G e n u s Geomackiea M i l l s , 1985
Geomackiea zephyrolata M i l l s , 1985
G e n u s Merga H a r t l a u b , 1913
Merga reesi Russell, 1956
G e n u s Pandea Lesson, 1843
Pandea rubra B i g e l o w , 1913
F a m i l y T r i c h y d r i d a e H i n c k s , 1868
G e n u s a n d species incertae sedis
Suborder C a p i t a t a Kühn, 1913
S u p e r f a m i l y Sphaerocorynoidea Prévôt, 1959
F a m i l y Paragotoeidae R a l p h , 1959
G e n u s Paragotoea K r a m p , 1942
Paragotoea bathybia K r a m p , 1942
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
39
F a m i l y incertae sedis
G e n u s Paulinum gen. n o v .
Paulinum lineatum spec. n o v .
S u p e r f a m i l y C o r y m o r p h o i d e a A l l m a n , 1872
F a m i l y C o r y m o r p h i d a e A l l m a n , 1872
G e n u s Euphysa Forbes, 1848
Euphysa vervoorti spec. n o v .
S u p e r f a m i l y C o r y n o i d e a Johnston, 1836
F a m i l y C o r y n i d a e Johnston, 1836
G e n u s Sarsia Lesson, 1843
Sarsia cliffordi B r i n c k m a n n - V o s s , 1989
O r d e r Leptothecatae C o r n e l i u s , 1992
S u p e r f a m i l y L a o d i c e o i d e a L . A g a s s i z , 1862
F a m i l y T i a r a n n i d a e Russell, 1940
G e n u s Modeeria Forbes, 1848
Modeeria rotunda ( Q u o y & G a i m a r d , 1827)
A c c o u n t of species
O r d e r Anthoathecatae C o r n e l i u s , 1992
Suborder F i l i f e r a Kühn, 1913
S u p e r f a m i l y P a n d e o i d e a H a e c k e l , 1879
F a m i l y Bythotiaridae M a a s , 1905 (= C a l y c o p s i d a e H a r t l a u b , 1913)
P a n d e o i d e a medusae w i t h o u t a p i c a l projection; w i t h s i m p l e or crenulate lips;
w i t h or w i t h o u t b r a n c h i n g of r a d i a l canals; w i t h or w i t h o u t centripetal canals; w i t h
four or m o r e h o l l o w m a r g i n a l tentacles w i t h p r o x i m a l part of tentacles adnate to base
of e x u m b r e l l a , rather t h i c k i n most species; w i t h o u t significant t h i c k e n i n g at base of
tentacle ( m a r g i n a l b u l b ) . C n i d o c y s t s often concentrated i n t e r m i n a l s w e l l i n g s of tentacle; w i t h or w i t h o u t r u d i m e n t a r y tentacles; w i t h or w i t h o u t ocelli.
T y p e genus: Bythotiara Günther, 1903.
R e m a r k s . — The F a m i l y Bythotiaridae is greatly i n need of r e v i s i o n , w h i c h is outside the scope of a paper o n species f r o m the Northeast Pacific. H o w e v e r , the distinct i o n b e t w e e n Heterotiara anonyma M a a s , 1905 a n d Bythotiara depressa N a u m o v , 1960 is
clarified. The v a r i a b i l i t y of other species is discussed a n d n e w collecting records are
i n c l u d e d . The f a m i l y n a m e Bythotiaridae M a a s , 1905 has p r i o r i t y over C a l y c o p s i d a e
H a r l a u b , 1913 (not B i g e l o w , 1913 as cited b y several recent papers).
G e n u s Bythotiara Günther, 1903
Bythotiaridae w i t h f o u r s i m p l e , or i r r e g u l a r l y bifurcate, r a d i a l canals, no centripetal canals. G o n a d s s m o o t h or w i t h folds w h e n mature.
T y p e species: Bythotiara murrayi Günther, 1903.
R e m a r k s . — T h i s genus is d i s t i n g u i s h e d f r o m Heterotiara M a a s , 1905 o n l y b y the
f o l d i n g of the mature gonads. A s discussed b e l o w , this is a v e r y variable character. In
a future f a m i l i a l r e v i s i o n these genera s h o u l d therefore p r o b a b l y be m e r g e d .
40
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Bythotiara depressa
(fig. l a , l c )
Heterotiara anonyma Bigelow, 1913: 25 (in part); 1919: 287 (in part); Arai & Brinckmann­Voss, 1980: 69
(in part) (not Heterotiara anonyma Maas, 1905:19, pl. 3).
Bythotiara depressa Naumov, 1960:191; Arai & Brinckmann­Voss, 1980: 64; Bouillon, 1980: 316 (in part).
Material.— Bythotiara depressa: 2 specimens 20.V.1906, 53°05'N 138°31'W, 1 specimen, 3­4.vi.1906, Bow­
ers Bank, Bering Sea, and 1 specimen, 20.vi. 1906,
off Startschkof I sland, E. of Kamchatka, coll. H.B.
Bigelow; 3 specimens, 13.vii.1966 "Vitiaz" Pacific Station No. 5600, 3500­600 m, coll. ZI STP; 3 of the
following 5 specimens deposited: R B C M 998­225, R B C M 998­226, R B C M 998­227: 5 specimens, ii­
iv.1980, 49 30'­54 00'Ν 127°34'­133 48'W, 0­610 m (preliminary record Fulton et al., 1982: 23) 223 spec­
ο
ο
0
imens during ii­iv.1982­1988, at outer shelf stations off the west coast of Vancouver I sland, depths
sampled with closing nets 300­700 m; 1 specimen, 19.vi.1990, 47°57'N 129°05'W, 0­1750 m, coll. B.J.
Burd, R.E. Thomson & G.S. Jamieson. I mmature Bythotiara depressa: R B C M 998­228: 1 specimen,
27.ii.1982, 48°43'N 126°40'W, 0­1180 m; R B C M 998­229:1 specimen, 16.ii.1987, 48°26'N 126°14'W, 300­0
m; Heterotiara anonyma; R B C M 998­230: 3 of 30 specimens deposited; Bay of Hansa, off Laing I sland,
Papua New Guinea, coll. J. Bouillon.
Specific characters.— U m b r e l l a laterally flattened. F o u r u n b r a n c h e d r a d i a l canals.
E i g h t tentacles, each w i t h t e r m i n a l nematocyst cluster thicker t h a n diameter of d i s t a l
part of tentacle, s p h e r i c a l w i t h f e w exceptions.
D e s c r i p t i o n . — U m b r e l l a u p to 20 m m h i g h , w i t h t h i c k mesoglea, especially at
apex, laterally flattened. M a n u b r i u m half l e n g t h of b e l l c a v i t y o r less; g o n a d s w i t h
v a r i e d degree of f o l d i n g . E i g h t tentacles thick at base, b e c o m i n g v e r y t h i n before ter­
m i n a l nematocyst clusters; w i t h o u t secondary tentacles. T e r m i n a l nematocyst clusters
of a d u l t specimens m o s t l y spherical, rarely ovate b u t still a r i s i n g a b r u p t l y f r o m a nar­
r o w p o r t i o n of the tentacle (fig. l a ) .
R e m a r k s . — Bythotiara depressa has frequently been confused w i t h Heterotiara anon­
yma M a a s 1905, o r i g i n a l l y described f r o m off Indonesia, b u t w i d e l y d i s t r i b u t e d i n
w a r m oceans. Based o n the literature descriptions of B. depressa a n d h i s o w n a b u n ­
dant m a t e r i a l of H. anonyma f r o m P a p u a N e w G u i n e a , B o u i l l o n (1980: 316) suggested
that the t w o species m i g h t be conspecific. H o w e v e r , for reasons g i v e n b e l o w they
s h o u l d be k e p t separate.
Bythotiara depressa w a s described b y N a u m o v (1960: 191) o n the basis of 31 speci­
mens f r o m 20 locations i n the n o r t h w e s t Pacific. The type, f r o m 51°53.1'N 161°49.6Έ,
is p r e s e r v e d i n the Z o o l o g i c a l Institute, St Petersburg (Savitskaya, 1977: 135). F u r t h e r
specimens f r o m the K u r i l ­ K a m c h a t k a T r e n c h w e r e e x a m i n e d b y N a u m o v (1971: 10).
W e have e x a m i n e d 232 specimens of B. depressa f r o m the northeast a n d n o r t h w e s t
Pacific ( A r a i & B r i n c k m a n n ­ V o s s , 1980: 64; m a t e r i a l listed above). These have been
c o m p a r e d w i t h 30 specimens of Heterotiara anonyma f r o m P a p u a N e w G u i n e a .
The t w o species resemble one another. E a c h s p e c i m e n has f o u r u n b r a n c h e d r a d i a l
canals a n d tentacles w i t h t e r m i n a l nematocyst thickenings. The tentacle n u m b e r i s
n o r m a l l y eight. A l t h o u g h p l a c e d i n different genera based o n degree of g o n a d f o l d ­
i n g , the species are i n fact v e r y s i m i l a r i n this respect; i n b o t h the gonads m a y be
s m o o t h or s h o w loose f o l d i n g as the m a n u b r i u m contracts.
A d i s t i n g u i s h i n g character is that the t e r m i n a l nematocyst clusters of the tentacles
are u s u a l l y spherical i n Bythotiara depressa, a n d a l w a y s arise a b r u p t l y f r o m a v e r y t h i n
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
41
Fig. 1. Structures in, a. Bythotiara depressa tentacle tip (recent collection), b. Heterotiara anonyma tentacle
tip (South Pacific, courtesy J. Bouillon), scale bar a,b: 0.15 mm, c. Bythotiara depressa margin and tentacle tip, adult, d. same species, juvenile, e. Heterotiara anonyma, scale bar c-e: 0.3 mm.
42
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
p o r t i o n of the tentacle, whereas they are m o r e elongate i n Heterotiara anonyma (see
B o u i l l o n et a l . , 1988: 94) (fig. 1). D e v e l o p i n g tentacles i n y o u n g B. depressa are m o r e
l i k e H. anomyma, b u t i n a d u l t specimens the difference is quite m a r k e d . R a d i a l canals
of B. depressa are twice the thickness of c o m p a r a b l y s i z e d H. anonyma. B. depressa does
not s h o w a n y t h i c k e n i n g at the j u n c t i o n of r a d i a l a n d r i n g canals, whereas H. anonyma
s h o w s a slight t h i c k e n i n g at that p o i n t . The v a c u o l a t e d p o r t i o n of e n d o d e r m a l cells i n
the adnate tentacle base is thicker i n B. depressa t h a n i n H. anonyma (fig. l c & e).
A n o t h e r distinctive character of B. depressa is the laterally compressed u m b r e l l a
described b y N a u m o v (1960: 191) a n d v e r i f i e d b y u s , w h i c h is l a c k i n g o r less d e v e l ­
o p e d i n H. anonyma (see B o u i l l o n , 1980: 316; present w o r k ) .
The above d e s c r i p t i o n of Bythotiara depressa w a s based o n a d u l t specimens. I n
a d d i t i o n w e have e x a m i n e d t w o smaller (4 a n d 4.8 m m h i g h , b o t h 4 m m w i d e ) speci­
mens w h i c h w e have tentatively assigned to the same species. B o t h specimens have
s m o o t h gonads, a n d some of the tentacles have spherical t e r m i n a l clusters, whereas
other shorter tentacles have n o distinct t e r m i n a l cluster b u t o n l y t h i c k e n e d tips (fig.
I d ) . Since s m o o t h gonads are t y p i c a l of d e v e l o p i n g gonads i n the s i m i l a r species
Bythotiara huntsmani ( B r i n c k m a n n ­ V o s s , 1979: 1228), w e assume that gonads of Bytho­
tiara depressa d e v e l o p i n the same w a y , a n d that these smaller specimens are y o u n g
Bythotiara depressa. T h e y are h o w e v e r , d i s t i n g u i s h e d f r o m Bythotiara huntsmani b y the
different n u m b e r a n d structure of the tentacles.
The o r i g i n a l d e s c r i p t i o n of Heterotiara anonyma b y M a a s (1905: 19) w a s based o n
t w o specimens l a c k i n g tentacles, f r o m 0°18'S 1 2 9 ° 1 5 Έ , H a l m a h e r a Sea. T h e t w o s y n ­
types are i n the Z o o l o g i c a l M u s e u m of A m s t e r d a m ( v a n Soest, 1975: 30). I n 1909 t w o
specimens w i t h 11 a n d 12 tentacles respectively w e r e assigned to the species f r o m col­
lections off P e r u (Bigelow, 1909: 216). T h i s i d e n t i f i c a t i o n w a s questioned b y V a n ­
hòffen (1911: 211), w h o reported another s p e c i m e n of H. anonyma f r o m the I n d i a n
O c e a n . I n 1913 B i g e l o w reported H. anonyma f r o m several localities i n the N o r t h
Pacific. W e e x a m i n e d f o u r of B i g e l o w ' s specimens (deposited i n the S m i t h s o n i a n
M u s e u m as Heterotiara anonyma), a n d f o u n d t h e m i d e n t i c a l w i t h Bythotiara depressa
Naumov.
The above incorrect identifications of H. anonyma have resulted i n a v e r y confused
literature to the present d a y . T h e species has been reported i n a v a r i e t y of w a r m
water locations, b u t u s u a l l y w i t h insufficient details to a l l o w a c o n f i r m a t i o n of i d e n ­
tity. I n 1980 w e reported Heterotiara anomyma f r o m the G u l f of A l a s k a based o n
B i g e l o w ' s incorrect records. W e n o w believe that it has never been correctly i d e n t i f i e d
i n the N o r t h e r n Pacific. Since w e h a d n o material at the time, the s p e c i m e n illustrated
( A r a i & B r i n c k m a n n ­ V o s s , 1980: 70) w a s obtained f r o m R.J . L a r s o n ' s collection off
M o n a Island, Puerto R i c o .
A g a i n s t the above b a c k g r o u n d , reports of Bythotiara depressa outside the N o r t h
Pacific m u s t also be treated w i t h scepticism u n t i l the details are k n o w n . P h i l l i p s
(1972: 98) described a single s p e c i m e n f r o m the G u l f of M e x i c o w h i c h w a s v e r y c r u m ­
p l e d , u n l i k e t y p i c a l specimens w i t h t h i c k mesogloea. A l t h o u g h w e have n o t checked
material, w e suggest that the records of Bythotiara depressa f r o m the Y u c a t a n Shelf a n d
M e x i c a n C a r i b b e a n b y Segura­Puertas (Segura­Puertas, 1992: 357; Segura­Puertas, L .
a n d Ordófíez­López, 1994:108) m a y also be referable to Heterotiara anonyma.
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
43
Bythotiara huntsmani (Fraser, 1911)
Crypta huntsmani Fraser, 1911:19, pl. 1.
Endocrypta huntsmani Fraser, 1912: 216; 1914:109, pl. 1; Rees, 1980: 48; (? Briggs & Gardner, 1931:181);
(not Endocrypta huntsmani Trebilcock, 1928: 1 = Bythotiara parasitica (Kirk, 1915)); (not Endocrypta
huntsmani Ralph, 1953: 66 = Bythotiara parasitica (Kirk, 1915)).
Bythotiara huntsmani Brinckmann-Voss, 1979: 1226; Arai & Brinckmann-Voss, 1980: 66; (?Bouillon,
1995a: 224; Bouillon et al. 1995: 8 = Bythotiara parasitica (Kirk, 1915)).
Material.— 1 specimen, 28.iv.1976, Departure Bay, surface, coll: M . N . Arai; R B C M 998-231: 1 specimen, 22.iv.1984, Cumshewa Inlet, Hecate Strait, surface, coll. J. Purcell.
Specific C h a r a c t e r s . — U m b r e l l a deep b e l l - s h a p e d i n a d u l t specimens. F o u r
u n b r a n c h e d r a d i a l canals. E i g h t a d r a d i a l gonads w i t h degree of f o l d i n g v a r y i n g f r o m
rather m a r k e d folds to n e a r l y s m o o t h surfaces. F o u r p e r r a d i a l tentacles w i t h t e r m i n a l
cnidocyst clusters not thicker t h a n diameter of contracted tentacles.
D e s c r i p t i o n of M e d u s a . — U m b r e l l a u p to 7.1 m m h i g h , u p to 6.5 m m w i d e , w i t h
scattered cnidocysts o n e x u m b r e l l a . S t o m a c h o c c u p y i n g about half of s u b u m b r e l l a r
cavity. L i p s f o u r - c o r n e r e d w i t h n u m e r o u s c n i d o c y s t patches. R a d i a l a n d r i n g canals
s m o o t h . W i t h o u t ocelli. M a n u b r i u m , j u n c t i o n of r a d i a l canals a n d r i n g canal, a n d tentacle t i p orange to d a r k b r o w n . G o n a d s of m a l e specimens m i l k w h i t e , female m o r e
transparent. R e m a i n d e r of m e d u s a colourless.
H y d r o i d . — The h y d r o i d has f o u r to five i r r e g u l a r w h o r l s of f i l i f o r m tentacles. Its
m o r p h o l o g y a n d b u d d i n g have been described b y Fraser (1911:19; 1914:109), B r i n c k m a n n - V o s s (1979:1227) a n d Rees (1980: 48).
R e m a r k s . — Bythotiara huntsmani has been o b t a i n e d f r o m ascidians collected i n the
Strait of G e o r g i a a n d off the west coast of V a n c o u v e r Island, B r i t i s h C o l u m b i a a n d
near F r i d a y H a r b o u r , W a s h i n g t o n (Fraser, 1911: 20; 1913: 149; 1914: 110; Rees, 1980:
48; B r i n c k m a n n - V o s s , 1979:1226). M o s t observed m e d u s a e h a v e been raised f r o m the
h y d r o i d s . H o w e v e r M i l l s (1981: 12) collected t w o f r e e - l i v i n g specimens f r o m Saanich
Inlet. O u r present m a t e r i a l i n c l u d e s another s p e c i m e n f r o m the Strait of G e o r g i a area
i n D e p a r t u r e Bay, a n d one f r o m C u m s h e w a Inlet, Hecate Strait.
B y t h o t i a r i d h y d r o i d s l i v i n g i n the p r e b r a n c h i a l z o n e of ascidians h a v e been
described f r o m several locations i n the Pacific. M e d u s a e h a v e been raised f r o m these
h y d r o i d s o r j u v e n i l e medusae, a n d (rarely) adults h a v e also been collected i n the
f i e l d . There is need f o r further research o n life cycles a n d v a r i a b i l i t y to c l a r i f y the
n u m b e r of species present. I n the above s y n o n y m y w e h a v e taken the conservative
a p p r o a c h that Bythotiara stilbosa M i l l s & Rees, 1979 a n d B. parasitica ( K i r k , 1915) are
distinct species ( M i l l s & Rees, 1979: 285; Schuchert, 1996: 23).
Calycopsis F e w k e s , 1882
B y t h o t i a r i d a e w i t h f o u r p r i m a r y r a d i a l canals, either w i t h centripetal canals b l i n d l y e n d i n g , o r j o i n i n g the base of the stomach, o r w i t h secondary canals f o r m i n g
branches of the p r i m a r y canal. Base of stomach b r o a d , often e x t e n d i n g a l o n g r a d i a l
canals f o r m i n g a cross.
T y p e species: Calycopsis typa F e w k e s , 1882
44
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
R e m a r k s . — W e consider the above d e f i n i t i o n of the genus Calycopsis to b e p r o v i sional u n t i l a complete r e v i s i o n of the f a m i l y c a n be m a d e . T h e type species, Calycopsis typa, w a s the first b y t h o t i a r i d described (Fewkes, 1882: 304). A s other species have
been d i s c o v e r e d , a n u m b e r of subsequent authors have attempted to d i s t i n g u i s h the
genus b y the extent of b r a n c h i n g of the r a d i a l canals a n d / o r the presence of centripetal canals. U n f o r t u n a t e l y b r a n c h i n g of several b y t h o t i a r i d s is h i g h l y i r r e g u l a r (e.g.
Bythotiara murrayi (see R u s s e l l , 1953: 217) a n d Calycopsis borchgrevinki (Browne, 1910)
(see M o o r e , 1984: 250)). B l i n d e n d e d centripetal canals c a n be clearly d i s t i n g u i s h e d .
H o w e v e r , those canals w h i c h connect the r i n g canal to the base of the stomach or the
f o u r p r i m a r y r a d i a l canals c a n n o t be differentiated into centripetal o r centrifugal
canals (as noted b y K r a m p , 1959: 27). I n some species o n l y occasional centripetal
canals are observed, a n d these have been interpreted as abberations (eg. B. murrayi
(see K r a m p , 1926: 97)) o r as i n d i c a t i o n of a n centripetal o r i g i n of other canals (eg. Calycopsis nematophora B i g e l o w , 1913 (see below)). T h e genus Sibogita c a n therefore n o t
be clearly d i s t i n g u i s h e d f r o m Calycopsis b y a lack of centripetal canals, at least u n t i l
d e v e l o p m e n t of the canals has been described a n d s h o w n to be centrifugal.
Calycopsis bigelowi (?) Vanhöffen, 1911
(fig-2)
Calycopsis bigelowi Vanhöffen, 1911: 218, fig. 12;
Kramp, 1957: 21; van der Spoel & Bleeker,
1988:181; ?Schuchert, 1996: 26.
Material— RBCM 998-232: 1 specimen, l . i i .
1987, 48°32'N 126°37'W, 500 m.
Specific characters.— F o u r r a d i a l
canals,
four interradial
centripetal
canals a n d eight p e r r a d i a l a n d interrad i a l tentacles w i t h t w o to three smaller
tentacles between t h e m .
Description.— Deep
bell-shaped
e x u m b r e l l a a n d m o r e conical s u b u m brella. E x u m b r e l l a 20 m m h i g h , 14 m m
w i d e , s u b u m b r e l l a 12 m m h i g h . F o u r
r a d i a l canals a n d three v i s i b l e b l i n d
interradial
centripetal
canals (the
f o u r t h q u a d r a n t is b a d l y damaged).
D e e p l y cleaved gonads i n u p p e r part of
m a n u b r i u m , l e a v i n g l o w e r part of
m a n u b r i u m g o n a d free. L a r g e m o u t h
w i t h sparsely f o l d e d lips, w i t h o u t
nematocyst clusters. E i g h t f u l l y d e v e l o p e d tentacles w i t h base adnate to
e x u m b r e l l a . T i p s of tentacles b r o k e n
off. B e t w e e n each of large tentacles one
Fig. 2. Calycopsis bigelowi (recent collection); small
tentacles could not be seen in all quadrants, because
margin was partly damaged. Scale bar: 4 mm.
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
45
to three d e v e l o p i n g tentacles.
R e m a r k s . — A l t h o u g h the tentacle tips of this s p e c i m e n w e r e b r o k e n off, w e c o u l d
assign it tentatively to Calycopsis bigelowi because of the tentacle arrangement a n d a
b l i n d centripetal canal i n each of the three u n d a m a g e d quadrants.
A l t h o u g h rarely recorded, C. bigelowi is w i d e l y d i s t r i b u t e d . It w a s o r i g i n a l l y
described f r o m the G u l f of A d e n (Vanhöffen, 1911: 219). It has since been collected i n
the A t l a n t i c west of the C a p e of G o o d H o p e ( K r a m p , 1957: 21) a n d off Scotland (Fraser, 1974: 13), a n d i n the Pacific i n the M a l a y a n A r c h i p e l a g o ( K r a m p , 1965: 46), i n the
B a n d a a n d A r u Seas (van d e r Spoel & Bleeker, 1988:167) a n d off N e w Z e a l a n d (Schuchert, 1996: 26).
Calycopsis nematophora B i g e l o w , 1913
(fig. 3)
?Sibogita simulans Bigelow, 1909: 213 (in part).
?Calycopsis typa Vanhöffen, 1911: 214, pl. 22 (in part, not Calycopsis typa Fewkes, 1882: 301, pl. 1).
Calycopsis nematophora Bigelow, 1913: 23, pi. 2, 3; 1940: 290; Naumov, 1956: 37; 1960: 211, pi. 29; Renshaw, 1965: 841; Arai & Brinckmann-Voss, 1980: 68; van der Spoel & Bleeker, 1988:167, fig. 15.
Perigonimus nematophora Naumov, 1955: 53, pi. 7.
Material.— RBCM 998-233:1 specimen, 16.iii.1980, 54°00'N 133°48'W, 0-620 m; RBCM 998-234:1 specimen, 19.iv.1980, 53°20*N 133°19'W, 0-610 m; each preliminary record Fulton et al. (1982: 23); 1 specimen, 25.Í.1987, 48°15'N 126°40'W, 500 m; R B C M 998-235: 1 specimen, 15.ii.1987, 48°26'N 126°20'W, 0700 m.
Specific characters.— M o u t h w i t h l a b i a l nematocyst clusters.
D e s c r i p t i o n . — U m b r e l l a u p to 30 m m h i g h , nearly as w i d e as h i g h i n preserved
specimens. M e s o g l e a about 1 / 5 - 1 / 4 diameter of e x u m b r e l l a . U p p e r p o r t i o n of s t o m ach b r o a d , extended o n f o u r p r i m a r y r a d i a l canals f o r m i n g a cross as seen f r o m
above. I n recent collections length of stomach less than half of the s u b u m b r e l l a ,
a l t h o u g h B i g e l o w (1913: 23) reported the length of the m a n u b r i u m as u p to the l e n g t h
of the b e l l cavity. L i p s intensively f o l d e d , p r o v i d e d w i t h o b l o n g cnidocyst clusters of
w h i c h the larger ones are stalked. F o u r p r i m a r y r a d i a l canals thick, f o r m i n g t r u n k s at
their base near the stomach, f o r k i n g into thinner r a d i a l canals a short distance f r o m
the stomach, w i t h a m a x i m u m n u m b e r of 17 canals (16 i n o u r specimens). E a c h canal
b o r d e r e d b y a b a n d of l o n g i t u d i n a l muscles. G o n a d s i n t e r r a d i a l w i t h several r o w s of
slit-like pits (rather t h a n folds as described b y some earlier authors). R a r e l y u p to 60
(33 i n o u r specimens) tentacles of different sizes; 2-8 thick h o l l o w longer tentacles,
w i t h majority of r e m a i n i n g tentacles m u c h shorter a n d s t u m p - l i k e . E a c h tentacle w i t h
a t e r m i n a l cnidocyst cluster (often d a m a g e d i n the longer tentacles). G o n a d s a n d
m a n u b r i u m d a r k r e d to p u r p l e i n l i v i n g specimens, often r e m a i n i n g r e d - b r o w n after
f o r m a l i n preservation.
R e m a r k s . — A l t h o u g h the genus Calycopsis is d e f i n e d p a r t l y as h a v i n g centripetal
canals, C. nematophora m a y have a l l its canals e x t e n d i n g f r o m the r i n g canal to the
four t r u n k - l i k e p r i m a r y r a d i a l canals. T h e o n l y exceptions described i n the literature
are 1 to 3 b l i n d canals connected to the r i n g canal i n 2 of B i g e l o w ' s specimens (Bige l o w 1913: 24) a n d one of R e n s h a w ' s 40 specimens (Renshaw, 1965: 846). I n a l l speci-
46
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Fig. 3. Calycopsis nematophora (collected 1959, Gulf of Alaska), specimen height 17 mm.
mens f r o m o u r collections canals connect the r i n g canal to the p r i m a r y r a d i a l canals,
w i t h o u t a n y b l i n d canals. Nevertheless the canals have been a s s u m e d to be d e v e l o p e d centripetally (Bigelow, 1913: 24), a n d w e have f o l l o w e d p r e v i o u s authors i n
i n c l u d i n g the species i n this genus.
The above specimens are the first collected i n C a n a d i a n waters, a l t h o u g h the species h a d been collected i n adjacent areas. P r e v i o u s records f r o m the N o r t h Pacific
w e r e s u m m a r i z e d b y A r a i & B r i n c k m a n n - V o s s (1980: 69). M o r e recent records f r o m
this area i n c l u d e those of A l v a r i n o (1980: 153), M i l l s (1981: 12), C o o n e y (1981: 960),
C h u c h u k a l o & B a b i c h (1981: 51) a n d v a n d e r Spoel & Bleeker (1988: 167), e x t e n d i n g
the range s o u t h to C a l i f o r n i a a n d the P h i l i p p i n e A r c h i p e l a g o .
F a m i l y H a l i m e d u s i d a e A r a i & B r i n c k m a n n - V o s s , 1980
P a n d e o i d e a m e d u s a e w i t h f o u r r a d i a l canals; w i t h s u b u m b r e l l a p r o t r u d i n g into
stomach g i v i n g a p e d u n c l e - l i k e appearance; m o u t h c r u c i f o r m w i t h r o w of nematocysts; w i t h f o u r p e r r a d i a l tentacles a n d f o u r i n t e r r a d i a l g r o u p s of tentacles, each h o l l o w a n d l a c k i n g adhesive organs; w i t h m a r g i n a l b u l b s w i t h abaxial ocelli.
G e n u s Halimedusa B i g e l o w , 1916
H a l i m e d u s i d a e w i t h characters of the f a m i l y .
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
47
Halimedusa typus B i g e l o w , 1916
Halimedusa typus Bigelow, 1916: 91, pi. 1; Arai & Brinckmann-Voss, 1980:
62.
Specific characters.— W i t h the characters of the genus.
R e m a r k s . — A detailed d e s c r i p t i o n of this species w a s g i v e n i n A r a i & B r i n c k m a n n - V o s s (1980: 62). A t that time the o n l y r e c o r d of this species i n the research area
w a s the o r i g i n a l one of three specimens off A m p h i t r i t e Point, B r i t i s h C o l u m b i a (Bige l o w , 1916: 91). It h a d been r e c o r d e d further south, a n d w e illustrated a s p e c i m e n
f r o m Y a q u i n a Bay, O r e g o n . M o r e recently w e h a v e e x a m i n e d f o u r other specimens
f r o m Masset Inlet, B r i t i s h C o l u m b i a ( A r a i , 1987:110).
F a m i l y P a n d e i d a e H a e c k e l , 1879
P a n d e o i d e a medusae w i t h or w i t h o u t a p i c a l projection. M a n u b r i u m m o r e or less
quadratic w i t h a f o u r cornered m o u t h (except i n Octotiara), w i t h l i p s s i m p l e to c o m p l e x l y f o l d e d . F o u r r a d i a l canals (except i n Octotiara) either s m o o t h or jagged. G o n a d s
interradial or a d r a d i a l o n m a n u b r i u m , rarely e x t e n d i n g to u p p e r part of r a d i a l canals,
s m o o t h or w i t h warts a n d pits, or cleaved or c o m p l e x l y f o l d e d . R a d i a l canals w i t h
s m o o t h w a l l s , or jagged. M a r g i n a l b u l b s w i t h or w i t h o u t ocelli. F i l i f o r m tentacles
w i t h or w i t h o u t tentaculae between t h e m . H y d r o i d s w h e r e k n o w n stolonal or s m a l l
u p r i g h t colonies, w i t h single w h o r l of tentacles i n the majority of species, or rarely
w i t h t w o w h o r l s or scattered tentacles.
T y p e genus: Pandea Lesson, 1837.
R e m a r k s . — Since 1980 t w o n e w species of p a n d e i d s have been described f r o m
this area; Amphinema platyhedos A r a i & B r i n c k m a n n - V o s s , 1983, a n d Geomackiea zephyrolata M i l l s , 1985. W e i n c l u d e brief descriptions of these species here, together w i t h
n e w records of Merga reesi R u s s e l l , 1956 a n d Pandea rubra B i g e l o w , 1913.
G e n u s Amphinema H a e c k e l , 1879
P a n d e i d a e medusae w i t h not m o r e t h a n t w o opposite, p e r r a d i a l tentacles; w i t h
m a r g i n a l warts or tentaculae; w i t h o u t gastric p e d u n c l e ; w i t h or w i t h o u t mesenteries;
m o u t h w i t h f o u r s i m p l e or w a v y l i p s ; gonads a d r a d i a l or i n t e r r a d i a l or e x t e n d i n g
a l o n g r a d i a l canals; w i t h or w i t h o u t ocelli.
T y p e species: Amphinema dinema (Peron & L e s u e u r , 1809).
Amphinema platyhedos A r a i & B r i n c k m a n n - V o s s , 1983
Amphinema platyhedos Arai & Brinckmann-Voss, 1983: 2179; Bouillon, 1985b: 254.
Specific characters.— V e r y b r o a d m a r g i n a l b u l b s , l o n g tentaculae b e t w e e n b u l b s
a n d large p o i n t e d a p i c a l projection.
R e m a r k s . — D e t a i l e d descriptions of three specimens w e r e g i v e n b y A r a i &
B r i n c k m a n n - V o s s (1983: 2179). These specimens h a d d a m a g e d lips, b u t B o u i l l o n
(1985b: 254, f i g . 4) i l l u s t r a t e d a f o u r cornered m o u t h w i t h s l i g h t l y c u r v e d l i p s .
48
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
The three Northeast Pacific specimens w e r e collected f r o m the Strait o f G e o r g i a
a n d B u r k e C h a n n e l ( A r a i & B r i n c k m a n n - V o s s , 1983: 2179). The species has also been
observed f r o m a s u b m a r i n e i n Jervis Inlet ( M a c k i e , 1985: 761). The f o u r t h collected
s p e c i m e n w a s f r o m H a n s a Bay, B i s m a r c k Sea, P a p u a N e w G u i n e a ( B o u i l l o n , 1985b:
254: B o u i l l o n et al., 1986:136). The specimens are d e p o s i t e d i n the C a n a d i a n M u s e u m
of N a t u r e (Frank et a l , 1985: 28), the R o y a l B r i t i s h C o l u m b i a M u s e u m , a n d TInstitut
R o y a l des Sciences N a t u r e l l e s de B e l g i q u e ( B o u i l l o n et al., 1995:14).
G e n u s Geomackiea M i l l s , 1985.
P a n d e i d a e w i t h f o u r p e r r a d i a l tentacles h a v i n g t a p e r i n g conical bases a n d w i t h
f o u r b r o a d , i n t e r r a d i a l bulbs, each r i m m e d b y a cluster of short tentaculae.
T y p e species: Geomackiea zephyrolata M i l l s , 1985.
Geomackiea zephyrolata M i l l s , 1985.
Geomackiea zephyrolata Mills, 1985: 2172.
Specific characters.— W i t h the characters of the genus.
R e m a r k s . — M i l l s (1985: 2172) gave detailed descriptions of b o t h a d u l t a n d j u v e nile specimens. The species has to date o n l y been recorded f r o m the N o r t h e a s t Pacific
area, i n the u p p e r 130 m of Saanich Inlet a n d F r i d a y H a r b o r .
G e n u s Merga H a r t l a u b , 1913
P a n d e i d a e medusae w i t h p e r r a d i a l sides of stomach connected to r a d i a l canals at
least to one t h i r d of stomach length. W i t h gonads not i n folds; w i t h four, eight, o r
m o r e tentacles w i t h or w i t h o u t r u d i m e n t a r y tentacles b e t w e e n t h e m .
T y p e species: Merga violacea ( A g a s s i z & M a y e r , 1899).
R e m a r k s . — Merga treubeli Schuchert, 1996 s h o u l d perhaps b e r e m o v e d f r o m the
genus Merga a n d the f a m i l y P a n d e i d a e , because the structure of the p r o x i m a l part of
the tentacles a n d the absence of m a r g i n a l b u l b s seems m o r e l i k e l y of c a l y c o p s i d t h a n
a p a n d e i d character (Schuchert, 1996: 74, f i g . 44c).
Merga reesi R u s s e l l , 1956
(fig. 4)
Merga reesi Russell, 1956b: 493.
Material.— R B C M 998-236 (slightly damaged): 1 specimen collected 15.ii.1980, 50°30'N
128°36'W,
depth 523 m; preliminary record Fulton et al. (1982: 23).
Specific characters.— M e d u s a e w i t h n o a p i c a l projection, w i t h gonads i n t e r r a d i a l
i n warts a n d grooves; w i t h four tentacles w i t h s w o l l e n p r o x i m a l section.
D e s c r i p t i o n of m e d u s a . — U m b r e l l a b e l l s h a p e d , height u p to 10 m m , w i d t h 8 m m .
Stomach flask s h a p e d w i t h b r o a d base. M o u t h w i t h s l i g h t l y crenulated l i p s a r m e d
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
49
Fig. 4. Merga reesi, a. after Russell (1956), with permission of Cambridge University Press, b. detail of
basal bulb and proximal part of tentacle (recent collection), ra: radial canal, ri: ring canal, t: tentacula,
s: thickened proximal part of perradial tentacle, scale bar: 0.5 mm.
w i t h a b a n d of nematocysts. G o n a d s i n t e r r a d i a l w i t h w a r t s a n d grooves.. R a d i a l
canals attached to s t o m a c h over m o r e t h a n half its length, f o r m i n g mesenteries. R a d i al canals s m o o t h , about t w i c e as w i d e as r i n g canal. F o u r tentacles w i t h s w o l l e n basal
part, w i t h e n l a r g e d e n d o d e r m (fig. 4b), l e a v i n g actual m a r g i n a l b u l b rather s m a l l .
O n e s m a l l tentacula i n each i n t e r r a d i u s . N o ocelli. C o l o u r of stomach a n d gonads
b r o w n r e d , f a d i n g i n a l c o h o l preservation.
H y d r o i d not k n o w n .
50
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
R e m a r k s . — I n the above d e s c r i p t i o n " W a r t s a n d g r o o v e s " is m o r e accurate t h a n
" i r r e g u l a r l y f o l d e d to f o r m n u m e r o u s raised c o r r u g a t i o n s " as described b y R u s s e l l
(1956: 493) i n his text. H i s figure (Russell 1956: 493, f i g . 1) h o w e v e r , s h o w s a g o n a d
w i t h warts a n d grooves, w h i c h concurs w i t h o u r specimen. Merga reesi a n d Amphinema krampi R u s s e l l , 1956, h a v e been p l a c e d i n different genera because of their tentacle
n u m b e r s . H o w e v e r , Merga reesi, M. bulbosa B o u i l l o n , 1980 a n d A. krampi share the
r e d u c e d m a r g i n a l b u l b s , a n d the t h i c k e n e d p r o x i m a l part of tentacles (Russell, 1956a:
371; 1956b: 493; 1958: 81; B o u i l l o n , 1980: 333). Merga reesi a n d A. krampi also have s i m ilar gonads. I n future revisions of the f a m i l y these latter t w o species s h o u l d b e p l a c e d
i n the same genus for structural reasons, rather t h a n b e i n g separated d u e to n u m e r i cal characters s u c h as tentacle n u m b e r s .
The three collections, each of single specimens of this species, have been w i d e l y
d i s t r i b u t e d : at the m o u t h of the E n g l i s h C h a n n e l , (Russell, 1956b: 493); i n the B a y of
Bengal, (Navas, 1971: 8; N a v a s - P e r e i r a & V a n n u c c i , 1991: 45); a n d the Northeast Pacific, present specimen.
G e n u s Pandea Lesson, 1843
P a n d e i d a e medusae w i t h u m b r e l l a w i t h or w i t h o u t conical a p i c a l projection; w i t h
mesenteries;
w i t h gonads f o r m i n g a n irregular n e t w o r k of ridges w i t h pits i n
between.
T y p e species: Pandea conica ( Q u o y & G a i m a r d , 1827).
Pandea rubra B i g e l o w , 1913
(fig. 5)
Pandea rubra Bigelow, 1913: 14, pi. 2; Kramp, 1926: 96, pl. 2; Bigelow, 1938:107; Kramp, 1957: 18; 1965:
41; Russell, 1970: 252; Arai «Sc Brinckmann-Voss, 1980: 61; Bleeker & van der Spoel, 1988: 231, fig.
14.
Material: 1 specimen, 2.iii. 1986, 48°11'N 125°56'W; R B C M 998-237: 2 specimens, 12.ii.1988, 48°11'N
125°56'W, 500-700 m; RBCM 998-238: 1 specimen, 19.vi.1990, 47°57'N 129°05*W, 1-1750 m, coll. B.J.
Burd, R.E. Thomson & C S . Jamieson.
Specific characters.— W i t h o u t e x u m b r e l l a r nematocyst tracts, w i t h o u t a p i c a l p r o cess; r a d i a l canals jagged, m a n u b r i u m deep b r o w n (may fade i n p r e s e r v e d specimens), densely f o l d e d , f r i l l y lips.
D e s c r i p t i o n . — U m b r e l l a b r o a d bell-shaped, u p to 75 m m h i g h a n d w i d e , w i t h soft
w a l l s , n o a p i c a l projection. M a n u b r i u m b a r r e l shaped, w i d e , about half as l o n g as b e l l
cavity, attached to p e r r a d i a l canals for about 4/5 of its length, m o u t h w i t h f r i l l y , d e l i cate a n d c o m p l e x l y f o l d e d lips; gonads close n e t w o r k o n u p p e r 2/3 of m a n u b r i u m ,
l e a v i n g lowest part g o n a d free. R a d i a l canals of present specimens d i s t a l l y b r o a d
w i t h w a v y or jagged outline; r i n g canal s m o o t h , about half the w i d t h of r a d i a l canals.
U p to 32 tentacles w i t h large, n o t laterally depressed b u l b s ; w i t h t h i n e x u m b r e l l a r
s p u r (fig. 5a, b).
R e m a r k s . — Specimens f r o m recent collections w e r e b e t w e e n 5 a n d 30 m m h i g h
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
51
a n d 5 to 28 m m w i d e , w i t h u p to 18 tentacles. D u e to the extremely delicate e x u m b r e l l a specimens are often t o r n (Bigelow,
1913:15) so that precise measurements are
i m p o s s i b l e to achieve. A s a result of the
t o r n e x u m b r e l l a the e x u m b r e l l a r s p u r is
difficult to see i n m a n y specimens a n d has
been o m i t t e d i n illustrations s u c h as those
of A r a i & B r i n c k m a n n - V o s s (1980: 61, f i g .
34c). H o w e v e r , i n o u r recent material w e
c o u l d c o n f i r m K r a m p ' s i l l u s t r a t i o n (1926:
p i . 2, f i g . 15) (fig. 5a & b).
These are the first records of Pandea
rubra f r o m C a n a d i a n waters since the o r i g i n a l d e s c r i p t i o n of B i g e l o w (1913:14), west
Fig. 5. Pandea rubra detail of marginal bulb
with abaxial spur: a. from recent collections,
of C a p e St. James, Q u e e n
Charlotte
b. after Kramp (1926: pl. 2, fig. 15). ex: exumIslands. It has, h o w e v e r , been collected at
brella, sp: abaxial spur.
w i d e l y separated locations i n the N o r t h
Pacific a n d other oceans (see s u m m a r i e s
b y A r a i & B r i n c k m a n n - V o s s , 1980: 62, A l v a r i n o , 1988: 107 a n d N a v a s - P e r e i r a & V a n nucci, 1990: 111).
F a m i l y T r i c h y d r i d a e H i n c k s , 1868
P a n d e o i d e a medusae w i t h o u t e x u m b r e l l a r cnidocyst tracts o r abaxial spurs;
m a n u b r i u m c r u c i f o r m w i t h f o u r p e r r a d i a l s i m p l e to flared l i p s ; gonads s m o o t h , interr a d i a l ; f o u r s m o o t h r a d i a l canals, w i t h or w i t h o u t fine b r a n c h canals connecting to the
r i n g canal; f o u r o r m o r e r o u n d o r elongate m a r g i n a l b u l b s ; tentacles, w h e r e k n o w n ,
s o l i d ; w i t h o u t ocelli. H y d r o i d , w h e r e k n o w n , Trichydra.
T y p e genus: Trichydra W r i g h t , 1858.
R e m a r k s . — The f a m i l y T r i c h y d r i d a e w a s i n t r o d u c e d b y H i n c k s (1868: 215) for the
h y d r o i d Trichydra pudica W r i g h t , 1858. E d w a r d s (1973a: 87) s h o w e d that the m e d u s a
w a s Pochella polynema H a r t l a u b , 1917, w h i c h h a d p r e v i o u s l y been p l a c e d i n the P r o boscidactylidae. In a d d i t i o n E d w a r d s transferred Pochella oligonema K r a m p , 1955, also
w i t h a c r u c i f o r m m a n u b r i u m , to Trichydra. H e p l a c e d the f a m i l y i n the A n t h o m e d u sae. Schuchert (1996: 87) re-examined the type specimens of K r a m p (1955: 270, p l . 2)
a n d p l a c e d Trichydra oligonema i n the genus Fabienna, r e t u r n i n g it p r o v i s i o n a l l y to the
Proboscidactylidae based o n tentacle structure. The f a m i l y T r i c h y d r i d a e became a
m o n o t y p i c f a m i l y of the Anthoathecatae.
Trichydra pudica has been described f r o m this research area ( A r a i & B r i n c k m a n n V o s s , 1980: 75; M i l l s , 1981: 16). A single s p e c i m e n of a s i m i l a r species has recently
been collected, a n d is descibed b e l o w .
T r i c h y d r i d a e incertae sedis gen., incertae sedis spec,
(fig. 6)
Material.— R B C M 998-239:1 specimen, ll.v.1986, 48°22'N 125°35'W, 0-148 m.
52
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Fig. 6. Trichydridae gen. and sp. incertae sedis a.
from above, b. side-view photograph of same
animal (recent collection), scale bars: 1 mm.
D e s c r i p t i o n of the m e d u s a . — U m b r e l l a 3.2 m m h i g h a n d 3 m m w i d e ; m a n u b r i u m
extended into four pouches, f o r m i n g a w i d e p e d u n c l e (fig. 6); m a n u b r i u m f o r m i n g a
neck-like constriction above the f l a r i n g four cornered l i p s ( m a n u b r i u m 1.6 m m at its
w i d e s t u p p e r part, 0.24 m m at its " n e c k " ) ; gonads smooth, c u s h i o n - l i k e , o n i n t e r r a d i a l
sides of stomach w i t h a deep i n t e r r a d i a l cleft; r a d i a l canals s m o o t h ( w i d t h 0.08 m m )
w i t h n o a d d i t i o n a l fine branches observed, r i n g canal h a r d l y v i s i b l e (0.04 m m ) ; f o u r
pear-shaped m a r g i n a l b u l b s (length 0.52 m m , diameter 0.36 m m ) thinner part of each
t a p e r i n g into the tentacles. F o u r p e r r a d i a l tentacles; n o a d d i t i o n a l b u l b s or tentaculae;
tentacles f i l i f o r m , n o v i s i b l e cnidocyst clusters (tentacles damaged). N e m a t o c y s t s n o t
checked because o n l y a single s p e c i m e n w a s available. C o l o u r of gonads a n d e n d o d e r m of m a r g i n a l b u l b s b r i g h t orange; otherwise colourless.
R e m a r k s . — W e have p l a c e d this species p r o v i s i o n a l l y i n the f a m i l y T r i c h y d r i d a e .
It closely resembles a d u l t Trichydra pudica because of the distinctive f l a r i n g of the l i p s
beneath the m a n u b r i u m , as w e l l as the characters listed i n the f a m i l y diagnosis above.
It differs i n the presence of o n l y f o u r tentacles, a n d i n the deep i n t e r r a d i a l clefts of the
gonad.
Suborder C a p i t a t a Kühn, 1913
S u p e r f a m i l y S p h a e r o c o r y n o i d e a Prévôt, 1959
R e m a r k s . — T h i s s u p e r f a m i l y w a s i n t r o d u c e d b y Petersen (1990:130). It unites the
families Sphaerocorynidae, Zancleopsidae, a n d Paragotoeidae. Because Petersen
(1990:101) u s e d a cladisitic a p p r o a c h h i s arrangement of families deviates significantl y f r o m p r e v i o u s arrangements s u c h as Petersen (1979: 105) a n d B o u i l l o n (1985a: 29).
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
53
W e agree w i t h Petersen (1990: 118) that s y n a p o m o r p h i c characters i n the medusae
l i n k the F a m i l y Paragotoeidae w i t h the Sphaerocorynidae.
F a m i l y Paragotoeidae R a l p h , 1959
A n t h o m e d u s a e w i t h at least one stiff tentacle w i t h s o l i d e n d o d e r m a n d a t e r m i n a l
cnidocyst b u l b . W i t h b r o a d m a n u b r i u m w i t h g o n a d e n c i r c l i n g it i n a w i d e g i r d l e .
W i t h four-cornered, rather w i d e m o u t h , w i t h s l i g h t l y f l a r i n g l i p s . W i t h o u t ocelli.
T y p e genus: Paragotoea K r a m p , 1942.
R e m a r k s . — The genus Paragotoea w a s o r i g i n a l l y i n c l u d e d i n the F a m i l y T u b u l a r i i dae b y K r a m p (1942: 29; 1959: 5). It w a s p l a c e d i n its o w n f a m i l y b y R a l p h (1959:176)
b u t most other authors have i n c l u d e d it i n the C o r y m o r p h i d a e ( B r i n c k m a n n - V o s s ,
1970: 22; B o u i l l o n , 1985a: 108; Boero & B o u i l l o n , 1993: 261). It w a s r e t u r n e d to the
F a m i l y Paragotoeidae b y Petersen (1990: 133), w h i c h w e consider justified. Recently
Pages a n d B o u i l l o n (1997:487) m o v e d the genus a g a i n to the C o r y m o r p h i d a e ; h o w e v er, w e still m a i n t a i n that the genus Paragotoea s h o u l d r e m a i n i n a separate f a m i l y
(see b e l o w ) .
G e n u s Paragotoea K r a m p , 1942
W i t h characters of the f a m i l y .
T y p e species: Paragotoea bathybia K r a m p , 1942.
R e m a r k s . — T h e genus Paragotoea w a s established b y K r a m p (1942: 26) f o r P.
bathybia. A s discussed b e l o w some specimens w h i c h have been assigned to P. bathybia
b y subsequent authors s h o u l d p r o b a b l y be referred to other species. I n a d d i t i o n M a r g u l i s (1989:127) a d d e d P. elegans to the genus. Except for some folds o n the e x u m b r e l la of P. elegans, w h i c h m a y be the result of preservation, a n d a longer m a n u b r i u m i n
some of M a r g u l i s ' specimens, the species are s t r i k i n g l y s i m i l a r .
Paragotoea bathybia K r a m p , 1942
(fig-7)
Paragotoea bathybia Kramp, 1942: 26, fig. 1-2; 1959: 5; (not Paragotoea bathybia Ralph, 1959:171 (see discussion below); not Paragotoea bathybia Brinckmann-Voss, 1970: 21, fig. 20 and 21 = undescribed
species; not Paragotoea bathybia Goy, 1972: 972, fig.3 = undescribed species); Pages and Bouillon,
1997: 487, fig.2.
Material.— 1 specimen, 4.iv.l986, 48°19'N 126°27'W, 500-700 m; deposited in Z M C , no number.
Specific characters.— E x u m b r e l l a s m o o t h ; m a n u b r i u m n o t longer than e x u m b r e l la; one tentacle w i t h its t e r m i n a l cnidocyst b u l b not faceted.
D e s c r i p t i o n . — E x u m b r e l l a a p p r o x i m a t e l y as h i g h as w i d e ; s p e c i m e n 1.9 m m h i g h ,
2.1 m m w i d e i n w i d e s t part of e x u m b r e l l a , s l i g h t l y less at m a r g i n . M e s o g l e a t h i n . O n e
tentacle, 0.8 m m l o n g , b r o a d at base, t h i n n i n g to about a t h i r d of its w i d t h d i s t a l l y .
W i t h large, almost spherical t e r m i n a l b u l b , diameter 0.3 m m . F o u r elongate, t r i a n g u lar cnidocyst p a d s of different size c l a s p i n g the m a r g i n of the b e l l , the largest reachi n g a p p r o x i m a t e l y h a l f w a y u p the e x u m b r e l l a . M a n u b r i u m barrel-shaped w i t h w i d e
54
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Fig. 7. Paragotoea bathybia (recent collection); specimen turned inside-out. g: gonad, m: manubrium, x:
exumbrella pad, scale bar: 0.5 mm.
quadratic m o u t h a n d s l i g h t l y f l a r e d l i p s . G o n a d s f o r m i n g a g i r d l e a r o u n d the m a n u b r i u m , w i t h n o i n t e r r a d i a l or p e r r a d i a l d i v i s i o n or pouches. R a d i a l canals 0.05 m m
thick, r i n g canal s l i g h t l y broader. N o mesenteries, (fig. 7).
R e m a r k s . — K r a m p (1942: 26) described this species f r o m a s p e c i m e n f r o m the
N W A t l a n t i c , off S W G r e e n l a n d , a n d c o n f i r m e d the m o r p h o l o g i c a l details w i t h a
s p e c i m e n f r o m off the C a p e of G o o d H o p e w h i c h w a s s l i g h t l y bigger t h a n the type
( K r a m p , 1959: 5). O u r s p e c i m e n agrees w i t h the type, w h i c h w a s also e x a m i n e d .
Because o u r s p e c i m e n w a s i n s i d e - o u t it w a s possible to see the m a n u b r i u m m o r e
clearly.
A n u m b e r of other specimens have been assigned to this species. R a l p h (1959:
171) e x a m i n e d three specimens f r o m west of the E n g l i s h C h a n n e l . O n e of these specimens c o r r e s p o n d e d to K r a m p ' s t w o specimens i n b e a r i n g a single tentacle. The seco n d (her f i g . 1 B, C ) possessed one complete tentacle a n d the s t u m p of a second i n the
p e r r a d i a l p o s i t i o n . She t r i e d to l i n k these w i t h a four-tentacled " a d u l t " specimen,
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
55
f r o m w h i c h , h o w e v e r , the tentacles h a d been b r o k e n off, so that the t y p i c a l t e r m i n a l
cnidocyst b u l b s of Paragotoea c o u l d not be v e r i f i e d . H e r figure d is a reconstruction of
her " a d u l t " s p e c i m e n of figure a, w i t h tentacles a d d e d i n the reconstruction. H e r
a r g u m e n t w a s that the one-tentacled specimens of P. bathybia have rather thick mesoglea o n the tentacle bases, a feature shared w i t h the f o u r tentacle s t u m p s of the
" a d u l t " specimen. H o w e v e r , i n other respects R a l p h ' s four-tentacled "Paragotoea" differs f r o m the P. bathybia described b y K r a m p . N e i t h e r K r a m p ' s type s p e c i m e n n o r h i s
second s p e c i m e n f r o m the S o u t h A t l a n t i c ( K r a m p 1959: 5) has stomach pouches as
s h o w n i n R a l p h ' s specimens, instead gonads encircle the stomach i n " s i m p l e a n n u l a r
g o n a d s " ( K r a m p 1959: 90). N e i t h e r K r a m p ' s t w o specimens n o r o u r s p e c i m e n have
any s i g n of a d d i t i o n a l tentacles as d o R a l p h ' s specimens . W e therefore consider
R a l p h ' s "Paragotoea bathybia" (Ralph,1959: 171) to p r o b a b l y b e l o n g to another species
or e v e n genus. U n f o r t u n a t e l y R a l p h ' s material w a s n o t available to u s .
"Paragotoea bathybia" has also been reported f r o m the M e d i t e r r a n e a n (Brinckm a n n - V o s s , 1970: 21; G o y , 1971: 399; G o y , 1972: 972; B r i n c k m a n n - V o s s , 1987: 133;
D a l l o t et a l . 1988: 197). W h e r e clear descriptions are g i v e n , these specimens differ
f r o m the type of Paragotoea bathybia, as has already been m e n t i o n e d b y B r i n c k m a n n V o s s (1970: 21) a n d M a r g u l i s (1989:129).
A f t e r the present paper w a s r e v i e w e d a n d i n press w e received a c o p y of "Pages,
F. a n d B o u i l l o n J., 1997. A r e d e s c r i p t i o n of Paragotoea bathybia K r a m p 1942 ( H y d r o i d o m e d u s e : C o r y m o r p h i d a e ) w i t h a n e w diagnosis for the genus Paragotoea. Sei. M a r .
61 (4): 487-493". Pages a n d B o u i l l o n (1997: 488) e x a m i n e d n i n e n e w specimens of the
species f r o m the A n t a r c t i c whereas w e h a d e x a m i n e d one n e w s p e c i m e n f r o m the
Pacific a n d the K r a m p (1942: 26) h o l o t y p e . F u l l d i s c u s s i o n is n o t possible here. W e
w o u l d need to examine Pages' a n d B o u i l l o n s specimens i n order to interpret the m o r p h o l o g y of the m a n u b r i u m ; f o r instance the v a c u o l a t e d cells at the base of the
m a n u b r i u m , w h i c h w e r e n o t present i n o u r specimen, nor h a v e they been m e n t i o n e d
b y K r a m p (1942: 26; 1959: 5). W e agree w i t h Pages a n d B o u i l l o n (1997: 492) that the
four-tentacled s p e c i m e n reconstructed b y R a l p h (1959: 173) is n o t Paragotoea bathybia,
h o w e v e r w e also m a i n t a i n that the M e d i t e r r a n e a n f o r m s h o u l d not be i n c l u d e d i n
that species (see above). W e c o n c l u d e that Paragotoea bathybia does n o t b e l o n g i n the
C o r y m o r p h i d a e , b u t i n a separate f a m i l y based o n the characteristics of the restricted
species.
F a m i l y incertae sedis
G e n u s Paulinum gen. n o v .
S p h a e r o c o r y n o i d e a w i t h cone-shaped e x u m b r e l l a . W i d e stomach, w i t h a conical
extension of the base into the mesoglea; m a n u b r i u m n o t t u b u l a r at m o u t h e n d . F o u r
thick r a d i a l canals a n d r i n g canal present; f o u r m a r g i n a l b u l b s w i t h a d a x i a l t h i c k e n ings, at least t w o of w h i c h bear stiff tentacles t e r m i n a t i n g i n a r o u n d cnidocyst b u l b .
T y p e species: Paulinum punctatum (Vanhöffen, 1911)
R e m a r k s . — Several authors have considered Dicodonium H a e c k e l , 1879 a d o u b t f u l
genus ( K r a m p , 1959: 83; B r i n c k m a n n - V o s s , 1970: 47; B o u i l l o n , 1985a: 89; Petersen,
1990: 210). It contains species w i t h t w o w e l l - d e v e l o p e d tentacles. T w o r u d i m e n t a r y
56
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
tentacles m a y be present or l a c k i n g . T h e species,
w h e r e recognizable, m a y b e l o n g to different families
or e v e n suborders. The type species Dicodonium cornutum H a e c k e l , 1879, is p o o r l y described. N e v e r t h e less D. punctatum (fig. 8) clearly differs f r o m the type
i n possessing a v e r y short m a n u b r i u m , f o u r tentacle
b u l b s , a n d tentacles each t e r m i n a t i n g i n a r o u n d
cnidocyst b u l b . It is therefore separated as the type
of a n e w genus.
In
a d d i t i o n to Paulinum
punctatum
the n e w
genus i n c l u d e s P. lineatum spec. n o v . The latter species is v e r y s i m i l a r to P. punctatum,
d i f f e r i n g i n the
arrangement of the mesogloeal inclusions, a n d p r o b ably i n the presence of f o u r tentacles (see b e l o w )
whereas P. punctatum has t w o f u l l y d e v e l o p e d tentacles a n d t w o r u d i m e n t s .
The b r o a d m a n u b r i u m a n d t w o to f o u r stiff tentacles m a y justify the referral of this genus to the
Fig. 8. Paulinum punctatum after
Vanhöffen (1911: 6), slightly modified. Scale bar approximately 0.5
S p h a e r o c o r y n o i d e a . If m o r e specimens of either of
mm (Vanhöffen gave only approx-
the t w o species of Paulinum
imate measurements).
become available, a n d
m o r e m o r p h o l o g i c a l details, as for instance the presence o r absence of stenoteles (see b e l o w ) are then v e r i f i e d , the genus m a y h a v e to b e
m o v e d a c c o r d i n g l y to another s u p e r f a m i l y or e v e n suborder. The a d a x i a l thickenings
o n the m a r g i n a l b u l b s s h o w s a relationship to the Sphaerocorynidae, h o w e v e r i t is
not clear if the cone-shaped extension of the stomach base into the mesoglea is h o m o l ogous w i t h the " a p i c a l c h a m b e r " of the Sphaerocorynidae.
E t y m o l o g y . — The genus w a s n a m e d i n h o n o u r of D r P a u l C o r n e l i u s , The N a t u r a l
H i s t o r y M u s e u m , L o n d o n , w h o has w r i t t e n a n u m b e r of detailed a n d extensive
papers o n the systematics a n d nomenclature of h y d r o z o a n s a n d thus c l a r i f i e d the taxo n o m y of n u m e r o u s families, genera a n d species i n this class.
Paulinum lineatum spec. n o v .
(figs 9 & 10)
Material.— holotype, R B C M 998-151-001: 1 specimen, 16.ii.1980, 52°30'N 131°53'W, off west coast
Moresby Island, B.C., 0-550 m; coll. T.C.Mullin.
Specific C h a r a c t e r s . — L i n e s of i n c l u s i o n s (zooxanthellae?) i n the mesoglea, m a i n l y a l o n g r a d i a l canals.
D e s c r i p t i o n . — M e d u s a deep bell shaped, 2.25 m m h i g h a n d 1.6 m m w i d e (preserved
i n 4% formaldehyde solution). Stomach about one third of subumbrella w i t h a large base,
m o u t h rather w i d e w i t h slightly w a v y lips. Stomach extending into conical apical projection. G o n a d s can be seen as thickenings o n stomach, but as the mesoglea is opaque further details c o u l d not be determined. Mesoglea thin, not more than one tenth of the d i a m eter of the m e d u s a peripherally, about triple that thickness apically. W i t h i n exumbrella
eight irregular adradial lines of r o u n d e d y e l l o w groups of cells, each g r o u p w i t h a darker
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
unstructured
center
(fig.
57
10a).
Four simple rather broad radial
canals, r i n g canal about half the
w i d t h of radial canals. F o u r very
thick, slightly triangular marginal
bulbs, each w i t h an adaxial thickening. O c e l l i absent. T w o adjacent
marginal bulbs bear rather
tentacles w i t h a large
nematocyst
cluster.
stiff
terminal
The
endo-
d e r m of the tentacles seemed h o l l o w i n their p r o x i m a l , solid i n
their distal, parts, t h o u g h it was
difficult to see i n the single specimen. The remaining t w o bulbs
carry o n l y stumps, probably broken off tentacles so that there were
possibly four tentacles. C n i d a e of
the m a r g i n a n d marginal bulbs
are microbasic euryteles; those of
the terminal cluster of tentacular
cnidocysts are atrichous (fig. 10b).
Because w e c o u l d not use the single w h o l e a n i m a l for cnidae identification, the possibility of other
cnidocyst types such as stenoteles
remains open.
Fig. 9. Paulinum lineatum spec. nov. (recent collection),
margin of specimen folded inwards. Scale bar: 0.5 mm.
R e m a r k s . — A l t h o u g h w e h a d o n l y one specimen, its g o o d state of preservation
a l l o w e d us to assess its m o r p h o l o g i c a l characters, so that a d e s c r i p t i o n of a n e w species is justified. Paulinum
lineatum bears a s t r i k i n g resemblance to P. punctatum ( V a n -
höffen) (fig. 8), except that P. punctatum has its mesogleal i n c l u s i o n s m o r e scattered
t h a n i n P. lineatum, a n d p r o b a b l y i n that P. lineatum has four tentacles.
S u p e r f a m i l y C o r y m o r p h o i d e a A l l m a n , 1872
F a m i l y C o r y m o r p h i d a e A l l m a n , 1872
M e d u s a e w i t h s i m p l e , circular m o u t h , one to f o u r h o l l o w tentacles m o s t l y m o n i l i f o r m , or s e l d o m w i t h nematocyst clusters spaced out a l o n g the tentacles, w i t h o u t
ocelli. G o n a d s e n c i r c l i n g m a n u b r i u m u n d i v i d e d . H y d r o i d s , w h e r e k n o w n , w i t h one
or m o r e aboral w h o r l s of m o n i l i f o r m or f i l i f o r m tentacles, a n d one or m o r e o r a l
w h o r l s of m o n i l i f o r m , capitate, or f i l i f o r m tentacles.
T y p e genus: Euphysa Forbes, 1848
G e n u s Euphysa Forbes, 1848
M e d u s a e w i t h o u t a p i c a l canal, w i t h e v e n l y r o u n d e d e x u m b r e l l a ; w i t h one to four
58
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
b
a
Fig. 10. Paulinum l ineatum spec, nov., a. detail of inclusions in mesoglea, see square in fig. 9 for loca­
tion, b. nematocyst from terminal cnidocyst bulb on tentacles. Scale bars each 10 μπι.
tentacles u n e v e n l y d e v e l o p e d b u t a l l of same structure. H y d r o i d s , w h e r e k n o w n , soli­
tary w i t h a n c h o r i n g filaments, w i t h short oral capitate o r m o n i l i f o r m tentacles, a n d
one or m o r e w h o r l s of l o n g aboral m o n i l i f o r m tentacles.
T y p e species: Euphysa aurata Forbes, 1848.
R e m a r k s . — The m e d u s a e Euphysa japonica (Maas, 1909) a n d E. tentaculata L i n k o ,
1905, a n d the h y d r o i d E. ruthae N o r e n b u r g & M o r s e , 1983, h a v e been r e p o r t e d f r o m
the research area ( A r a i & B r i n c k m a n n ­ V o s s , 1980: 6; M i l l s , 1981: 13; N o r e n b u r g &
M o r s e , 1983: 1). A r a i & M a s o n (1982: 8, f i g . 2) collected apart f r o m t y p i c a l E. japonica,
specimens f r o m the Strait of G e o r g i a w i t h d i v e r t i c u l a e o n the r a d i a l canals w h i c h
they suggested to b e l o n g to another species. H o w e v e r , subsequent e x a m i n a t i o n of
over 200 specimens f r o m waters off B r i t i s h C o l u m b i a s h o w a considerable v a r i a t i o n
i n the r a d i a l canals f r o m s m o o t h t h r o u g h rather jagged to s u c h specimens w i t h d i v e r ­
ticulae. I n other respects these specimens cannot be d i s t i n g u i s h e d f r o m E. japonica. I n
a d d i t i o n , specimens of a Euphysa w h i c h d o not possess the m o r p h o l o g i c a l characters
of the already described species of the genus, h a v e been f o u n d i n the research area
since 1980, a n d w e consider t h e m to b e l o n g to a n e w species described b e l o w .
Euphysa vervoorti spec. n o v .
(fig. I D
Material.— paratype ROMI Z B­3012: 1 specimen, 29.vi.1982, 48°42'N 125°03'W, 0­140 m, coll. M . N .
Arai; 1 specimen, 3.iv.l986, 48°36'N 125°8'W, 0­105 m; 1 specimen, 29.iii.1987, off Secretary I sland,
B.C., Juan de Fuca Strait, l m ; holotype R B C M 998­152­001:1 specimen 30.vi.1994, off Secretary Island,
B.C., Juan de Fuca Strait, 1 m, coll. A Brinckmann­Voss.
Specific C h a r a c t e r s . — H i g h , d o m e s h a p e d e x u m b r e l l a , delicate, often d a m a g e d i n
p l a n k t o n collections. M o n i l i f o r m tentacles. M a r g i n a l b u l b s of d i f f e r i n g sizes w i t h
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
59
thick endodermal swelling, a n d w i t h
large, often p r o t r u d i n g abaxial n e m a ­
tocyst pads.
Description of m e d u s a . — U m b r e l ­
la u p to 6 m m h i g h a n d 5 m m w i d e .
M a n u b r i u m slightly shorter than bell
cavity,
gonad
completely
surrounded b y
leaving only
short
part i n
m o u t h region free. L i p s simple, r o u n d .
A d u l t specimens w i t h ripe gonads
(eggs visible) i n specimens 4 m m h i g h
or larger. N o peduncle. O n e very large
marginal bulb, w i t h a large endoder­
m a l chamber and abaxially protruding
nematocyst p a d . Three smaller margi­
nal bulbs of gradually diminishing
size. M o n i l i f o r m tentacles rather thick
proximally, tiiinning distally. Tentacles
of differing sizes even i n fully mature
specimens, however the one originat­
i n g f r o m the largest marginal b u l b
always being the longest. C o l o u r of
m a n u b r i u m a n d e n d o d e r m of margi­
nal bulbs deep orange­red. U m b r e l l a
colourless.
R e m a r k s . — There
seven
other
species
are currently
of the
genus
Euphysa of w h i c h the medusa stage has
been adequately described: Euphysa
aurata Forbes, 1848, E. brevia (Uchida,
1947), E. flammea
(Linko,
1905), E.
japonica (Maas, 1909), Ε. problematica
Schuchert, 1996, Ε. tentaculata L i n k o ,
1905, a n d E. tetrabrachia Bigelow, 1904.
Euphysa
monotentaculata
Zamponi,
Fig. 11. a. Euphysa vervoorti spec, nov., a. type speci­
men (recent collection), b. marginal bulb Euphysa
1983, is too poorly described to be
japonica, c. marginal bulb Euphysa tentacul ata (a, b and
identified. Euphysa brevia, E. problemati­
c in specimens of the same size). Scale bar: 1.1 mm.
ca, a n d E. tetrabrachia differ f r o m the
other species because the tentacle structure is not m o n i l i f o r m . I n fact they m a y be better
r e m o v e d f r o m the genus based o n the tentacle structure a n d some other m o r p h o l o g i ­
cal characters. T h e r e m a i n i n g species a l l h a v e m o n i l i f o r m tentacles as E. vervoorti
does, h o w e v e r they lack the thick e n d o d e r m a l s w e l l i n g s i n the m a r g i n a l b u l b s a n d
the extensive nematocyst p a d s of that species. O f the m o n i l i f o r m species o n l y E. tenta­
culata shares w i t h E. vervoorti the character of h a v i n g tentacles of different l e n g t h
w h e n a d u l t . H o w e v e r , E . tentaculata is d i s t i n g u i s h e d f r o m the latter species b y the
presence of a s m a l l p e d u n c l e , m u c h thinner r a d i a l canals, s m a l l r o u n d e n d o d e r m a l
60
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
s w e l l i n g s i n the m a r g i n a l bulbs, a n d thinner p r o x i m a l parts of the tentacles (compare
f i g . 11a w i t h 11c).
E. japonica a n d E. flammea are not distinguishable i n a d u l t stages (see d i s c u s s i o n
A r a i & B r i n c k m a n n - V o s s , 1980: 7). Euphysa japonica medusae are released r e g u l a r l y
f r o m its h y d r o i d w i t h f o u r equal tentacles ( B r i n c k m a n n - V o s s , u n p u b l i s h e d data). T h i s
species is definitely present i n the reseach area. M i l l s (1981: 13) also collected v e r y
y o u n g medusae w i t h o n l y one tentacle at F r i d a y H a r b o u r a n d i d e n t i f i e d t h e m tentat i v e l y as E. flammea. H o w e v e r , u n t i l the life cycles of E. vervoorti spec. n o v . a n d E. tentaculata are established, it is m o r e probable that the single-tentacled juveniles b e l o n g
to one of these k n o w n l o c a l species t h a n to E. flammea.
E t y m o l o g y . — T h i s species w a s n a m e d i n h o n o u r of Prof. W i m V e r v o o r t , N a t i o naal N a t u u r h i s t o r i s c h M u s e u m , L e i d e n , to w h o m this v o l u m e is dedicated. W e w i s h
to h o n o u r h i m for his extensive a n d erudite publications o n the t a x o n o m y of H y d r o zoa, i n c l u d i n g his b i b l i o g r a p h y of L e p t o l i d a , a n d thank h i m for his assistance a n d
encouragement of m a n y other scholars i n the f i e l d .
S u p e r f a m i l y C o r y n o i d e a Johnston,
F a m i l y C o r y n i d a e Johnston,
1836
1836.
M e d u s a e , w h e n present, w i t h h i g h e r t h a n w i d e u m b r e l l a ; w i t h s i m p l e , circular
m o u t h ; w i t h gonads r i n g like; four r a d i a l canals a n d four e q u a l l y d e v e l o p e d tentacles;
w i t h abaxial ocellus. H y d r o i d colonies erect w i t h stems b r a n c h e d or u n b r a n c h e d ,
w i t h a m o r e or less f i r m perisarc; w i t h w h o r l of capitate tentacles a r o u n d m o u t h , w i t h
a d d i t i o n a l capitate tentacles b e l o w ; w i t h or w i t h o u t tentacle l i k e a d d i t i o n a l structures
(Tardent & Stòssel, 1971: 682).
T y p e genus: Coryne Gaertner, 1774.
R e m a r k s . — There have recently been a n u m b e r of papers o n this f a m i l y , a n d controversy o n the importance of generic characters, the extent of the f a m i l y , a n d o n the
g r o u p i n g of the medusae a n d h y d r o i d s into genera (see C a l d e r , 1988: 63; B r i n c k m a n n - V o s s 1989: 688; Petersen, 1990: 205; B o u i l l o n , 1995b: 305; Schuchert, 1996:119).
Research o n the genus Sarsia of this r e g i o n is o n g o i n g ( M i l l e r , 1982: 153; B r i n c k m a n n - V o s s , 1985: 673) a n d w i l l be further reported elsewhere. O n e n e w species has
been reported since 1980; Sarsia cliffordi B r i n c k m a n n - V o s s , 1989 ( B r i n c k m a n n - V o s s ,
1989: 685).
O r d e r Leptothecatae C o r n e l i u s , 1992
S u p e r f a m i l y L a o d i c e o i d e a L . A g a s s i z , 1862
R e m a r k s . — The d i s t i n g u i s h i n g feature of the medusae i n the s u p e r f a m i l y is the
presence of m a r g i n a l c o r d y l i ( B o u i l l o n , 1985a: 143), a l t h o u g h these structures m a y not
be h o m o l o g o u s i n the different families. A clear d e f i n i t i o n of the m o r e diverse h y d r o i d
stages is d i f f i c u l t .
F a m i l y T i a r a n n i d a e R u s s e l l , 1940
L a o d i c e o i d e a w i t h h e m i s p h e r i c a l to s l i g h t l y
flatter
b e l l ; w i t h large
stomach
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
61
attached to s u b u m b r e l l a a l o n g p e r r a d i a l f u r r o w s . W i t h f o u r s i m p l e r a d i a l canals,
w i t h o u t centripetal canals; w i t h s i m p l e f o l d e d gonads situated either a d r a d i a l l y o n
u p p e r part of stomach o r a l o n g r a d i a l canals o n extended stomach pouches; w i t h o u t
ocelli; w i t h h o l l o w m a r g i n a l tentacles w i t h m o r e o r less conical b u l b s ; w i t h s p i n d l e shaped c o r d y l i w i t h t e r m i n a l nematocysts. H y d r o i d , w h e r e k n o w n , Stegopoma.
T y p e genus: Tiaranna H a r t l a u b , 1914.
R e m a r k s . — T h e f a m i l y T i a r a n n i d a e w a s i n t r o d u c e d b y R u s s e l l (1940: 518) f o r
medusae of the genera Tiaranna ( n o w Modeeria) a n d Chromatonema. The h y d r o i d genera Stegopoma a n d Stegolaria have also been i n c l u d e d (see d i s c u s s i o n b y B o u i l l o n 1985a:
145). M o s t recently the m e d u s a n genus Margalefia
Pages, B o u i l l o n & G i l i , 1991 has
p r o v i s i o n a l l y been a d d e d (Pages et a l , 1991: 92).
G e n u s Modeeria Forbes, 1848
M e d u s a . — T i a r a n n i d a e w i t h gonads r e g u l a r l y a n d laterally f o l d e d o n u p p e r part
of stomach, a d r a d i a l w i t h i n t e r r a d i a l connections i n y o u n g specimens, a d r a d i a l i n
a d u l t medusae w i t h short extension onto r a d i a l canals i n largest specimens.
T y p e species: Modeeria formosa Forbes, 1848.
Modeeria rotunda ( Q u o y & G a i m a r d , 1827)
(fig. 12)
Dianaea rotunda Quoy & Gaimard, 1827:181, pi. 6A, figs. 1-2.
Modeeria formosa Forbes, 1848: 70, pi. 7, fig. 1.
Campanularia fastigiata Alder, 1860: 73-74, pi. 5, fig.l.
Tiara (Tiaranna) rotunda Haeckel, 1879: 57, pi. 3, figs. 9-10.
Stegopoma fastigiatum Levinsen, 1893:180, pl.6, fig.8.
Tiaranna rotunda Hartlaub, 1914: 266-8, figs. 218-219.
Modeeria rotunda Edwards, 1963: 464, fig. 1; 1973: 573, figs. 1-3.
F o r the extensive s y n o n y m y of this species see E d w a r d s (1973b: 588), R a m i l &
V e r v o o r t (1992: 29), C o r n e l i u s (1995:109), a n d H i r o h i t o (1995: 88).
Material of medusae.— BCPM (= RBCM) 976-1126-2: 1 specimen, 24.iii.1976, 52°N 128°W, Mathieson
Channel, off Hecate Strait, 0-319 m, coll. A . Peden; R B C M 998-240: 1 specimen, 27.ii.1982, 48°43'N
126°40'W, 0-1180 m; 5 specimens, 24.i.l987-21.iii.l987, 47 56-48 26'N 126 14'-126°40 W, 500 m.
0
0
o
,
Specific characters.— W i t h the characters of the genus.
D e s c r i p t i o n . — U m b r e l l a h e m i s p h e r i c a l , s l i g h t l y w i d e r t h a n h i g h i n most specimens, i n present material 10-20 m m w i d e a n d 5-17 m m h i g h ; jelly v e r y thick, often
m o r e than one t h i r d of height of b e l l apically, t h i n n i n g o u t p e r i p h e r a l l y . Stomach
shaped l i k e a four s i d e d p y r a m i d i n its u p p e r part; w i t h gonads as folds o n the edges
of the p y r a m i d a l part of the stomach. L o w e r part of the stomach e x t e n d i n g about one
t h i r d into s u b u m b r e l l a r space (fig. 12a) w i t h four cornered m o u t h w i t h s l i g h t l y crenulate m a r g i n s ; r a d i a l canals smooth, entering m i d d l e of m a n u b r i u m w h i l e b e n d i n g
i n w a r d s a n d raised to f o r m deep pockets i n s u b u m b r e l l a (fig. 12b). M a r g i n w i t h rather b r o a d conical b u l b s w i t h u p to 6 m a r g i n a l tentacles p e r quadrant i n specimens
62
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Brinckmann-Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
63
f r o m recent collections. O n e (in most specimens) to 3 s p i n d l e s h a p e d c o r d y l i b e t w e e n
each t w o tentacles (fig. 12c). M a n u b r i u m b r i g h t r e d , f a d i n g i n p r e s e r v e d specimens;
mesoglea transparent except m o r e w h i t i s h a n d thick i n area of s u b u m b r e l l a r pockets.
R e m a r k s . — A l l o u r specimens are medusae. F o r descriptions of the h y d r o i d see
R a m i l & V e r v o o r t (1992: 30), C o r n e l i u s (1995:110), a n d H i r o h i t o (1995: 88).
T h i s is the first r e c o r d of this species f r o m the northeast Pacific O c e a n . H o w e v e r ,
it is w i d e l y recorded f r o m the w o r l d ' s oceans. The h y d r o i d has been r e c o r d e d i n the
northwest Pacific O c e a n f r o m Japan ( H i r o h i t o , 1995: 88) a n d K o r e a (Park, 1988: 62).
A s n o t e d b y E d w a r d s (1973b: 582) Tiaranna sagamina U c h i d a , 1948, w a s described
f r o m a d a m a g e d i m m a t u r e m e d u s a f r o m S a g a m i Bay, Japan ( U c h i d a , 1948: 334); h o w ever, since there are n o c o r d y l i m e n t i o n e d a n d g o n a d s are not d e v e l o p e d , the i d e n t i t y
of this species is d o u b t f u l . ( A l t h o u g h often q u o t e d as 1947, U c h i d a ' s paper w a s actually p u b l i s h e d i n J u l y 1948, w i t h a n attached strip of p a p e r r e c o r d i n g the correct
date).
Acknowledgements
W e are grateful to the m a n y people w h o h a v e h e l p e d us w i t h this w o r k . P r e p a r a t i o n of the present p a p e r w a s carried out p r i m a r i l y b y A . B r i n c k m a n n - V o s s at the
R o y a l O n t a r i o M u s e u m , Toronto, O n t a r i o , a n d she thanks that i n s t i t u t i o n for p r o v i d i n g space a n d facilities. M . N . A r a i is grateful to R.J. B e a m i s h a n d D . N o a k e s for p r o v i s i o n of space a n d facilities at the D e p a r t m e n t of Fisheries a n d Oceans Pacific B i o l o g i cal Station, N a n a i m o , B . C . d u r i n g v a r i o u s p o r t i o n s of this w o r k . W e are grateful to
K . W . Petersen ( Z o o l o g i c a l M u s e u m , C o p e n h a g e n ) for d i s c u s s i o n of some species a n d
to D a l e C a l d e r ( R o y a l O n t a r i o M u s e u m ) for m u c h advice a n d h e l p . W e are i n d e b t e d
to Z . K a b a t a a n d W . v . H e r t z e n b e r g for translations of R u s s i a n literature.
M e m b e r s of the G r o u n d f i s h Section of the Pacific B i o l o g i c a l Station collected the
majority of the f i e l d samples. S o r t i n g of cnidarians f r o m the 1987 a n d 1988 f i e l d s a m ples at P.B.S. w a s carried out b y T. A p p l e t o n a n d J.E. Strong s u p p o r t e d b y a D e p a r t ment of Fisheries a n d Oceans grant to M . N . A r a i . G . A . M c F a r l a n e acted as the Fisheries a n d Oceans l i a i s o n officer for the project. S o r t i n g of 1980 a n d 1986 b o n g o samples
b y K . T h o m p s o n a n d I . M . C h a n w a s s u p p o r t e d b y N a t u r a l Sciences a n d E n g i n e e r i n g
Research C o u n c i l G r a n t A 2 0 0 7 to M . N . A r a i . T h i s grant also s u p p o r t e d subsequent
sorting of the H y d r o z o a a n d p r e l i m i n a r y taxonomie w o r k at the D e p a r t m e n t of B i o l o g i c a l Sciences, U n i v e r s i t y of C a l g a r y , i n c l u d i n g collaboration w i t h A . B r i n c k m a n n Voss.
W e thank J. P u r c e l l , J.C. M a s o n , B. B u r d , a n d the R o y a l B r i t i s h C o l u m b i a M u s e u m
for a l l o w i n g us to examine other samples f r o m the research area. W e thank J. B o u i l l o n
(Institut R o y a l des Sciences N a t u r e l l e s de Belgique, Bruxelles), K . W . Petersen (Zool o g i c a l M u s e u m , Copenhagen), S. Stepanjants ( Z o o l o g i c a l Institute, St. Petersburg),
a n d the S m i t h s o n i a n Institution, W a s h i n g t o n , D . C , for s e n d i n g m a t e r i a l for c o m p a r i son.
Fig. 12. Modeeria rotunda (recent collections) a. manubrium, scale bar: 1.4 mm, b. connection of radial
canal to manubrium forming subumbrellar pocket, arrow shows bend of radial canal towards manubrium, scale bar: 1 mm, c. margin to show cordyli and tentacles, scale bar: 0.5 mm.
64
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
References
Alder, J.G., 1860. Descriptions of a zoophyte and two species of Echinodermata new to Britain.—
Ann. Mag. Nat. Hist. (ser. 3) 5: 73­75.
Alvarino, Α., 1980. The relation between the distribution of zooplankton predators and anchovy lar­
vae.— Calif. Coop. Oceanic Fish. Invest. Rep. 21:150­160.
Alvarino, Α., 1988. Pandea cybeles, a new medusa from the Sargasso Sea (Coelenterata: Anthomedusae:
Pandeidae).— Proc. Biol. Soc. Wash. 101:102­108.
Arai, M . N . , 1987. Population ecology of the hydromedusae of Masset I nlet, British Columbia. In: J.
Bouillon, F. Boero, F. Cicogna, and P.F.S. Cornelius (eds). Modern Trends in the Systematics,
Ecology, and Evolution of Hydroids and Hydromedusae.— Clarendon Press, Oxford: 133­141.
Arai, M . N . and A. Brinckmann­Voss, 1980. Hydromedusae of British Columbia and Puget Sounds­
Can. Bull. Fish. Aquat. Sei. 204:1­192.
Arai, M . N . and A . Brinckmann­Voss, 1983. A new species of Amphinema: Amphinema platyhedos n. sp.
(Cnidaria, Hydrozoa, Pandeidae) from the Canadian West Coast.— Can. J. Zool. 61: 2179­ 2182.
Arai, M . N . and J.C. Mason, 1982. Spring and summer abundance and vertical ditribution of Hydro­
medusae of the central Strait of Georgia, British Columbia.— Syesis 15: 7­15.
Arai, M . N . , G.A. McFarlane, M.W. Saunders, and G . M . Mapstone, 1993. Spring abundance of medu­
sae, ctenophores, and siphonophores off southwest Vancouver I sland: Possible competition or
prédation on sablefish larvae.— Can. Tech. Rep. Fish. Aquat. Sei. 1939:1­37.
Bigelow, H.B., 1904. Medusae from the Maldive Islands.— Bull. Mus. Comp. Zool. Harv. Coll. 39: 245­
267.
Bigelow, H.B., 1909. Reports on the scientific results of the expedition to the eastern tropical Pacific, in
charge of Alexander Agassiz, by the U.S. Fish Commission Steamer "Albatross". XVI . The medu­
sae.— Mem. Mus. Comp. Zool. Harv. Coll. 37:1­243. pis 1­48.
Bigelow, H.B., 1913. Medusae and Siphonophorae collected by the U.S. Fisheries Steamer "Albatross"
in the northwestern Pacific, 1906.— Proc. U.S. Natl. Mus. 44(1946): 1­119, pis 1­6.
Bigelow, H.B., 1916. Halimedusa, a new genus of Anthomedusae.— Trans. R. Soc. Can. Sect. 4. 91­95,
pi i­
Bigelow, H.B., 1919. Hydromedusae, siphonophores and ctenophores of the "Albatross" Philippine
Expedition.— U. S. Nat. Mus. Bull. 1 (5): 279­362, pis 38­43.
Bigelow, H.B., 1938. Plankton of the Bermuda Océanographie Expeditions. VI I I . Medusae taken dur­
ing the years 1929 and 1930.— Zoologica (Ν. Y.) 23: 99­189.
Bigelow, H.B., 1940. Eastern Pacific Expeditions of the New York Zoological Society. XX. Medusae of
the Templeton Crocker and Eastern Pacific Zaca Expeditions, 1936­1938.— Zoologica (N.Y.) 25:
281­ 321.
Bleeker, J. and S. van der Spoel, 1988. Medusae of the Amsterdam M i d North Atlantic Plankton Expe­
ditions (1980­1983) with description of two new species.— Bijdr. Dierkd. 59: 227­258.
Boero, F. and J. Bouillon, 1993. Zoogeography and life cycle patterns of Mediterranean hydromedusae
(Cnidaria).— Biol. J. Linn. Soc. 48: 239­266.
Bouillon, J., 1980. Hydroméduses de la mer de Bismarck (Papouasie, Nouvelle­Guinée). I I I : Antho­
medusae Filifera (Hydrozoa­Cnidaria).— Cah. Biol. Mar. 21: 307­344.
Bouillon, J., 1985a. Essai de classification des Hydropolypes­Hydroméduses (Hydrozoa­Cnidaria).—
Indo­Malayan Zool. 1: 29­243.
Bouillon, J., 1985b. Notes additionnelles sur les Hydroméduses de la mer de Bismarck (Hydrozoa­
Cnidaria).— Indo­Malayan Zool. 2: 245­266.
Bouillon, J., 1995a. Hydromedusae of the New Zealand Océanographie I nstitute (Hydrozoa, Cnida­
ria).— N.Z. J. Zool. 22: 223­238.
Bouillon, J., 1995b. Classe des Hydrozoaires (Hydrozoa Owen, 1843). In: D.Doumenc, (ed.), Traité de
Zoologie Anatomie, Systématique, Biologie III(II) Cnidaires. Cténaires.— Masson, Paris: 29­416.
Bouillon, J., F. Boero, and G . Seghers, 1988. Notes additionelles sur les Hydroméduses de la mer de
Bismarck (Hydrozoa­Cnidaria) II.— Indo­Malayan Zool. 5: 87­99.
Bouillon, J., M . Claereboudt, and G. Seghers, 1986. Hydroméduses de la baie de Hansa (Mer de Bis­
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
65
marck; Papouasie Nouvelle­Guinée): Répartition, conditions climatiques et hydrologiques.—
Indo­Malayan Zool. 3:105­152.
Bouillon, J., C. Massin, and R. Kresevic, 1995. Hydroidomedusae de l'I nstitut royal des Sciences natu­
relles de Belgique.— Doc. Trav. Inst. R. Sei. Nat. Belg. 78:1­106.
Briggs, E.A. and V.E. Gardner, 1931. Hydroida.— Sei. Rep. Great Barrier Reef Exped. 4:181­196.
Brinckmann­Voss, Α., 1970. Anthomedusae/Athecatae
(Hydrozoa, Cnidaria) of the Mediterranean.
Pt. I. Capitata.— Fauna Flora Golfo Napoli 39:1­96, pis 1­11.
Brinckmann­Voss, Α., 1979. The life­cycle of Bythotiara huntsmani (Fraser 1911) (Calycopsidae, Hydro­
zoa, Cnidaria).— Can. J. Zool. 57:1226­1231.
Brinckmann­Voss, Α., 1985. Hydroids and medusae of Sarsia apicula (Murbach and Shearer, 1902) and
Sarsia princeps (Haeckel, 1879) from British Columbia and Puget Sound with an evaluation of
their systematic characters.— Can. J. Zool. 63: 673­681.
Brinckmann­Voss, Α., 1987. Seasonal distribution of hydromedusae (Cnidaria, Hydrozoa) from the
Gulf of Naples and vicinity, with observations on sexual and asexual reproduction in some spe­
cies. I n: J. Bouillon, F. Boero, F. Cicogna, and P.F.S. Cornelius (eds). Modern Trends in the
Systematics, Ecology, and Evolution of Hydroids and Hydromedusae.— Clarendon Press,
Oxford: 133­141.
Brinckmann­Voss, Α., 1989. Sarsia cl iffordi n. sp. (Cnidaria, Hydrozoa, Anthomedusae) from British
Columbia, with distribution records and evaluation of related species.— Can. J. Zool. 67: 685­
691.
Calder, D.R., 1988. Shallow­water hydroids of Bermuda: The Athecatae.— R. Ont. Mus. Life Sei. Con­
trib. 148:1­107.
Chuchukalo, V.l., and A . G . Babich, 1981. Conditions of food stock of Laemonema and Macrurus in win­
ter.— Izv. Tikhookean. Nauchno­Issled. I nst. Rybn. Khoz. Okeanogr. 105: 50­55.
Cooney, R.T., 1981. Bering Sea zooplankton and micronekton communities with emphasis on annual
production. I n: D.W. Hood and J.A. Calder (eds). The Eastern Bering Sea Shelf: Oceanography
and Resources.— N O A A , Seattle: 947­974.
Cornelius, P.F.S., 1995. North­west European Thecate Hydroids and their Medusae. Part 1. Laodicei­
dae to Haleciidae. —Synopses of the British Fauna (New Series), R.S.K. Barnes and J.H. Crothers
(eds) No. 50: i­vii, 1­347.— Shrewsbury.
Dallot, S., J. Goy and C. Carré, 1988. Peuplements de carnivores planctoniques gélatineux et struc­
tures productives en Méditerranée occidentale.— Oceanol. Acta 1988, num. spec: 193­209.
Edwards, C , 1963. On the anthomedusae Tiaranna rotunda and Modeeria formosa.— J. mar. biol. Assoc.
U.K. 43: 457­467.
Edwards, C , 1973a. The hydroid Trichydra pudica and its medusa Pochel l a pol ynema.— J. mar. biol.
Assoc. U.K. 53: 87­92.
Edwards, C , 1973b. The medusa Modeeria rotunda and its hydroid Stegopoma fastigiatum, with a review
of Stegopoma and Stegol aria.— J. mar. biol. Assoc. U.K. 53: 573­600.
Fewkes, J.W., 1882. O n the Acalephae of the east coast of New England.— Bull. Mus. Comp. Zool.
Harv. Coll. 9: 291­310, pl. 1.
Forbes, Ε., 1848. A Monograph of the British Naked­eyed Medusae: with Figures of all the Species, 1­
104, pis 1­13.— Ray Society, London.
Frank, P.G., J.A. Fournier, and J. Madill, 1985. Type specimens of invertebrates (Mollusca and Arthro­
poda excluded) in the National Museum of Natural Sciences National Museums of Canada.—
Syllogeus 60:1­119.
Fraser, C M . , 1911. The hydroids of the west coast of North America with special reference to those of
the Vancouver I sland region — Bull. Lab. Nat. Hist. State Univ. I owa (New Ser.) 28, 6(1): 1­91, pis
1­8.
Fraser, C M . , 1912. Endocrypta huntsmani.— Science (Washington DC) (New Ser.) 35(893): 216.
Fraser, C. M . , 1913. Hydroids from Vancouver I sland.— Bull. Vic. Mem. Mus. Geol. Surv. Can. 1:147­
155.
Fraser, C M . , 1914. Some hydroids of the Vancouver I sland region.— Trans. R. Soc. Can. Ser. 3, 8 Sect.
4: 99­209, pis 1­26.
66
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Fraser, J.H., 1974. The distribution of medusae in the Scottish area.— Proc. R. Soc. Edinb. Sect. Β
(Biol.) 74:1­25.
Fulton, J., M . N . Arai, and J.C. Mason, 1982. Euphausiids, Coelenterates, Ctenophores, and other zoo­
plankton from the Canadian Pacific Coast I chthyoplankton Survey, 1980.— Can. Tech. Rep. Fish.
Aquat. Sei. 1125:1­75.
Goy, J., 1971. Sur la répartition bathymétrique des Hydroméduses en mer de Ligurie.— Rapp. P­v.
Réun. Commn. int. Explor. Sei. Mer Mediterr. 20: 397­400.
Goy, J., 1972. Les Hydroméduses de la mer Ligure.— Bull. Mus. Nat. Hist. Nat. (Zool.) (3) 83: 965­
1008.
Haeckel, E., 1879. Das System der Medusen. Erster Theil einer Monographie der Medusen. System der
Craspedoten: 1­360, pis 1­20.— Denkschr. Med. naturwiss. Ges. Jena 1:1­360.
Hartlaub, C , 1914. Craspedote Medusen I, (3) Familie IV Tiaridae.— Nord. Plankton 17 (XII): 237­363.
Hincks, T, 1868. A history of the British hydroid zoophytes. Vol. 1, i­lxviii, 1­338; Vol. 2, pis 1­67—
Van Voorst, London.
Hirohito, 1995. The Hydroids of Sagami Bay I I Thecata: i­vi, 1­355, pi. 1­13.— Biological Laboratory
Imperial Household, Tokyo.
Kirk, H.B., 1915. On Ascidioclava, a new genus of gymnoblastic hydroids.— Trans. Proc. N . Z . I nst. 47:
146­148. pl. 1.
Kramp, P.L., 1926. Medusae. Part II. Anthomedusae.— Dan. I ngolf Exped. 5:1­102, pis 1­2.
Kramp, P.L., 1942. Medusae. In: The Godthaab Expedition 1928.— Medd. Grönland 81(1): 1­168.
Kramp, P.L., 1955. The medusae of the tropical west coast of Africa.— Atl. Rep. 3: 239­324, pis 1­3.
Kramp, P.L., 1957. Hydromedusae from the Discovery collections.— Discovery Rep. 29: 3­128, pis 1­7.
Kramp, P.L., 1959. The Hydromedusae of the Atlantic Ocean and adjacent waters.— Dana­Rep. Carls­
berg Found. 46:1­283.
Kramp, P.L., 1965. The Hydromedusae of the Pacific and Indian Oceans.— Dana­Rep. Carlsberg
Found. 63:1­162.
Levinsen, G.M.R., 1893. Meduser, Ctenophorer og Hydroider fra Grönlands Vestkyst, tilligemed
Bemaerkninger om Hydroidernes Systematik.— Vidensk. Medd. Dan. Naturhist. Foren. 4: 143­
220.
Linko, Α., 1905. Zoologisch Studien im Barents­Meere.— Zool. Anz. 28: 210­220.
Maas, O., 1905. Die Craspedoten Medusen der Siboga­Expedition.— Siboga Exped. 10:1­84, pis 1­14.
Maas, O. 1909. Japanische Medusen.— Abh. Bayer. Akad. Wiss. Math.— Naturwiss. Abt. 1, Suppl. 8:
1­52.
Mackie, G.O., 1985. Midwater macroplankton of British Columbia studied by submersible PI SCES
IV.—J. Plankton Res. 7: 753­777.
Margulis, P. Ya., 1989. New hydroid jelly­fishes of the Family Tubulariidae (Coelenterata, Hydro­
zoa).— Zool. Zh. 68 (6): 126­130 (Russian with English summary).
Miller, R.L., 1982. Identification of sibling species within the "Sarsia tubul osa complex" at Friday Har­
bor, Washington (Hydrozoa: Anthomedusae).— J. Exp. Mar. Biol. Ecol. 62:153­172.
Mills, C E . , 1981. Seasonal occurrence of planktonic medusae and ctenophores in the San Juan Archi­
pelago (NE Pacific).— Wasmann J. Biol. 39: 6­29.
Mills, C E . , 1985. A new hydrozoan, Geomackiea zephyrol ata gen. nov., sp. nov. (Anthomedusae: Pan­
deidae), from inland marine waters of British Columbia and Washington State.— Can. J. Zool. 63:
2172­ 2175.
Mills, C E . , 1987. Hydromedusae. I n: E. Kozloff. Marine invertebrates of the Pacific Northwest.— Uni­
versity of Washington Press, Seattle: 32­44.
Mills, C E . and J.T. Rees, 1979. Bythotiara stil bosa, new species, (Anthomedusae: Calycopsidae) from
neritic waters in central California.—J. Nat. Hist. 13: 285­293.
Moore, S.J., 1984. Morphological variation in the hydromedusa Cal ycopsis borchgrevinki (Browne, 1910)
(Coelenterata, Hydrozoa).— Cah. Biol. Mar. 25: 245­256.
Naumov, D.V., 1955. Phylum Coelenterata. I n: E.N. Pavlovskii (ed.). A n Atlas of the I nvertebrates of
the Far Eastern Seas of the USSR.— Izdarel'stvo Akademii Nauk SSSR, Moskva, Leningrad: 51­
68, pis 7­11. (Transi, from Russian by Israel Program for Sei. Transi., Jerusalem, 1966).
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
67
Naumov, D.V., 1956. Medusae in the far­eastern waters of the Soviet Union.— Tr. Probl. Temat.
Soveshch. Zool. Inst. Akad. nauk SSSR 6: 33­41 (In Russian).
Naumov, D.V., 1960. Hydroids and Hydromedusae of the USSR. Keys to the fauna of the USSR.—
Zool. I nst. Acad. Sei. USSR 70:1­626 (Transi, from Russian by Israel Program for Sei. Transi., Jeru­
salem, 1969).
Naumov, D.V., 1971. Hydromedusae and Scyphomedusae from the Kurile­Kamchatka Trench.— Tr.
Inst. Okeanol. Akad. Nauk SSSR 92: 9­17.
Navas, D., 1971. New records of hydromedusae from the I ndian Ocean.— Contrib. Avuls. Inst. Ocean.
22:1­33.
Navas­Pereira, D. and M . Vannucci, 1990. Antarctic Hydromedusae and water masses.— Pesqui.
Antart. Bras. 2:101­141.
Navas­Pereira, D. and M . Vannucci, 1991. The hydromedusae and water masses of the I ndian
Ocean.— Bol. Inst. Oceanogr. 39: 25­60.
Norenburg, J.L. and M.P. Morse, 1983. Systematic implications of Euphysa ruthae n. sp. (Athecata:
Corymorphidae), a psammophilic solitary hydroid with unusual morphogenesis.— Trans. A m .
Microsc. Soc. 102:1­17.
Pages, F., J. Bouillon, and J.­M. Gili, 1991. Four new species of Hydromedusae (Cnidaria, Hydrozoa)
from the coast of south­western Africa.— Zool. Scr. 20: 89­98.
Pages, F., and J. Bouillon, 1997. A redesciption of Paragotoea bathybia Kramp 1942 (Hydroidomedusae:
Corymorphidae) with a new diagnosis for the genus Paragotoea. Sei. Mar. 61 (4) : 487­493.
Park, J.H., 1988. Three hydroids (Cnidaria: Hydroida) from Ullûngdo and Chejudo, Korea.— Korean
J. Syst. Zool. 4(1): 57­66.
Petersen, K.W., 1979. Development of coloniality in Hydrozoa. In: G . Larwood and B.R. Rosen (eds).
Biology and Systematics of Colonial Organisms.— Academic Press, London: 105­140.
Petersen, K.W., 1990. Evolution and taxonomy in capitate hydroids and medusae (Cnidaria: Hydro­
zoa).— Zool. J. Linn. Soc. 100:101­231.
Phillips, P.J., 197£. ï h e pelagic cnidaria of the Gulf of Mexico: zoogeography, ecology and systematics.—
PhD dissertation, Texas A. & M . University, 1972:1­212.
Quoy, J.R.C.,/and J.P. Gaimard, 1827. Observations zoologiques faites à bord de Γ Astrolabe, en mai
1826, dans le détroit de Gibraltar.— Ann. Sei. Nat. 10:1­21,172­193,225­239.
Ralph, P.M., 1953. A guide to the athecate (gymnoblastic) hydroids and medusae of New Zealand.—
Tuatara 5: 59­75.
Ralph, P.M., 1959. The status and affinities of the anthomedusan Paragotoea bathybia Kramp, 1942.—
Proc. Zool. Soc. Lond. 133:171­177.
Ramil, F. and W. Vervoort, 1992. Report on the Hydroida collected by the " B A L G IM " expedition in
and around the Strait of Gibralter.— Zool. Verh. (Leiden) 277:1­262.
Rees, J.T., 1980. The symbiotic hydrozoan Endocrypta huntsmani, its ascidian hosts, and its affinities
with calycopsid hydromedusae.— Wasmann J. Biol. 37: 48­54.
Renshaw, R.W., 1965. Distribution and morphology of the medusa, Cal ycopsis nematophora, from the
North Pacific Ocean.—J. Fish. Res. Board Can. 22: 841­847.
Russell, F.S., 1940. On the nematocysts of Hydromedusae I I I .— J. mar. biol. Assoc. U.K. 24: 515­523.
Russell, F.S., 1953. The Medusae of the British Isles: Anthomedusae, Leptomedusae, Limnomedusae,
Trachymedusae and Narcomedusae, i­xiii, 1­530, pis 1­35.— Cambridge University Press, Cam­
bridge.
Russell, F.S., 1956a. O n a new medusa, Amphinema krampi n. sp.— J. mar. biol. Assoc. U.K. 35: 371­373.
Russell, F.S., 1956b. O n two new medusae, Merga reesi n. sp. and Tiaropsidium atl anticum n. sp.— J.
mar. biol. Assoc. U.K. 35: 493­ 498.
Russell, F.S., 1958. Notes on the medusa Amphinema krampi Russell.— J. mar. biol. Assoc. U.K. 37: 81­
84.
Russell, F.S., 1970. The Medusae of the British Isles.II. Pelagic Scyphozoa with a Supplement to the
First Volume on Hydromedusae, i­xi, 1­284, pis 1­15, I s.— Cambridge University Press, Cam­
bridge.
Savitskaja, K.V., 1977. A list of type species of the Hydromedusae and Siphonophora.— Issled. Fauny
68
Brinckmann­Voss & Arai. Further notes on Leptolida. Zool. Verh. Leiden 323 (1998)
Morei 19:135­137.
Schuchert, P., 1996. The marine fauna of New Zealand: Athecate hydroids and their medusae (Cni­
daria: Hydrozoa).— N.Z. Oceanogr. I nst. Mem. 106:1­159.
Segura­Puertas, L., 1992. Medusae (Cnidaria) from the Yucatan Shelf and Mexican Caribbean.— Bull.
Mar. Sei. 51: 353­359.
Segura­Puertas, L. and U . Ordónez­López, 1994. Análisis de la comunidad de medusas (Cnidaria) de
la Region Oriental del Banco de Campeche y el Caribe Mexicano.— Caribb. J. Sei. 30:104­115.
Tardent, P. and F. Stössel, 1971. Die Mechanorezeptoren der Polypen von Coryne pintneri, Sarsia reesi
und Cl adonema radiatum (Athecata, Capitata).— Rev. suisse Zool. 78: 680­688.
Trebilcock, R.E., 1928. Notes on New Zealand Hydroida.— Proc. R. Soc. Vie. 41:1­31, pis. 1­7.
Uchida, T., 1947. Some medusae from the central Pacific.— J. Fac. Sei. Hokkaido Univ. (Ser. VI ), Zool.
9 (3): 297­319.
Uchida, T., 1948. Medusae in the vicinity of Shimoda.— J. Fac. Sei. Hokkaido Univ. (Ser. VI ), Zool. 9
(4): 331­343.
Vanhöffen, Ε., 1911. Die Anthomedusen und Leptomedusen der deutschen Tiefsee­Expedition 1898­
1899.— Deutsche Tiefsee­Expedition 1898­1899.19 (5): 191­233, pl. 22.
van der Spoel, S. and J. Bleeker, 1988. Medusae from the Banda Sea and A r u Sea plankton, collected
during the Snellius II Expeditions, 1984­1985.— Indo­Malayan Zool. 5:161­202.
van Soest, R.W.M., 1975. A catalogue of the coelenterate type specimens of the Zoological Museum of
Amsterdam. I . General introduction, pelagic coelenterates.— Beaufortia 24: 27­35.
Zamponi, M.O., 1983. Nuevas adiciones a la medusofauna de la region subantartica. I. Anthomedusae
y Narcomedusae (Coelenterata: Hydrozoa).— Neotropica 29 (82): 173­181.