Plumularian hydroids (Cnidaria: Hydrozoa) from
the Strait of Gibraltar and nearby areas
M . D . M e d e l & W . Vervoort
Medel, M . D . & W . Vervoort. Plumularian hydroids (Cnidaria: Hydrozoa) from the Strait of Gibraltar
and nearby areas.
Zool. Verh. Leiden 300,17.vii.1995; 1-72, figs 1-28.—ISSN 0024-1652/ISBN 90-73239-41-9.
M.D. Medel, Laboratorio de Biologia Marina (Zoologia), Departamento de Fisiologia y Biologia animal, Facultad de Biologia, Universidad de Sevilla, Espana (Spain).
W. Vervoort, National Museum of Natural History, Leiden, Netherlands.
Key words: Cnidaria; Hydrozoa; Hydroida; Plumularioidea; Strait of Gibraltar; relationships.
Twenty-four species of plumularian hydroids (Cnidaria: Hydrozoa) from the Strait of Gibraltar area
are described and figured, including material from the coasts of southern Spain (between El Portil
near the Spanish-Portuguese border and Adra, Mediterranean coast of Granada, c. 3°W) and from
Ceuta in northern Africa. The geographical distribution of all species is briefly discussed and where
relevant additional notes are added.
Contents
Introduction
4
L i s t of localities a n d species collected
5
Taxonomie review
7
S u p e r f a m i l y P l u m u l a r i o i d e a H i n c k s , 1868
7
F a m i l y A g l a o p h e n i i d a e B r o c h , 1918
G e n u s Aglaophenia L a m o u r o u x , 1812
7
7
Aglaophenia acacia A l l m a n , 1883
7
Aglaophenia kirchenpaueri (Heller, 1868)
Aglaophenia octodonta (Heller, 1868)
Aglaophenia párvula Bale, 1882
9
11
13
Aglaophenia picardi S v o b o d a , 1979
15
Aglaophenia pluma (Linnaeus, 1758)
17
Aglaophenia tubiformis Marktanner-Turneretscher, 1890
20
Aglaophenia tubulifera ( H i n c k s , 1861)
22
G e n u s Streptocaulus A l l m a n . 1883
Streptocaulus dollfusi ( B i l l a r d , 1924)
G e n u s Gymnangium H i n c k s , 1874
Gymnangium montagui ( B i l l a r d , 1912)
G e n u s Lytocarpia K i r c h e n p a u e r , 1872
Lytocarpia myriophyllum (Linnaeus, 1758)
F a m i l y H a l o p t e r i d i d a e M i l l a r d , 1962
G e n u s Antennella A l l m a n , 1877
Antennella siliquosa ( H i n c k s , 1877)
Antennella secundaria ( G m e l i n , 1791)
G e n u s Halopteris A l l m a n , 1877
24
24
28
28
30
30
32
32
32
35
37
4
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Halopteris diaphana (Heller, 1868)
37
Halopteris liechtensterni (Marktanner-Turneretscher, 1890)
F a m i l y K i r c h e n p a u e r i i d a e M i l l a r d , 1962
G e n u s Kirchenpaueria Jickeli, 1883
Kirchenpaueria pinnata (Linnaeus, 1758)
G e n u s Ventromma Stechow, 1923
39
41
41
41
45
Ventromma halecioides ( A l d e r , 1859)
F a m i l y P l u m u l a r i i d a e H i n c k s , 1868
G e n u s Nemertesia L a m o u r o u x , 1812
45
48
48
Nemertesia ramosa (Lamarck, 1816)
48
Nemertesia antennina (Linnaeus, 1758)
50
Nemertesia irregularis ( Q u e i c h , 1885)
G e n u s Plumularia L a m a r c k , 1816
Plumularia setacea (Linnaeus, 1758)
G e n u s Monotheca N u t t i n g , 1900
Monotheca pulchella (Bale, 1882)
G e n u s Polyplumaria G . O . Sars, 1874
PolyplumariaflabellataG . O . Sars, 1874
G e n u s Pseudoplumaria R a m i l & Vervoort, 1992
Pseudoplumaria marocana ( B i l l a r d , 1930)
52
56
56
58
58
61
61
63
63
Introduction
T h i s s t u d y of the p l u m u l a r i a n h y d r o i d s (families A g l a o p h e n i i d a e , H a l o p t e r i d a e ,
K i r c h e n p a u e r i i d a e a n d P l u m u l a r i i d a e ) f r o m the Strait of G i b r a l t a r is p a r t of a taxon o m i e i n v e s t i g a t i o n of the h y d r o i d f a u n a of that area that has been c a r r i e d o u t over
the last years. T h o u g h R a m i l & V e r v o o r t (1992) s t u d i e d the deeper w a t e r h y d r o i d
f a u n a of b o t h entrances to the Strait of Gibraltar, the m o r e s u p e r f i c i a l waters b o r d e r i n g the coastal area f r o m that g e o g r a p h i c a l l y m o s t interesting z o n e are still l a r g e l y
u n s t u d i e d as far as the h y d r o i d f a u n a is concerned. F o r the present s t u d y m a t e r i a l
has b e e n o b t a i n e d f r o m the coastal area of s o u t h e r n S p a i n b e t w e e n E l P o r t i l near the
S p a n i s h - P o r t u g u e s e b o r d e r a n d A d r a , M e d i t e r r a n e a n coast of G r a n a d a , c. 3°W; a d d i t i o n a l m a t e r i a l f r o m the coastal r e g i o n of C e u t a i n N o r t h A f r i c a has been i n c l u d e d .
C o l l e c t i n g of the m a t e r i a l has m a i n l y been d o n e b y S C U B A d i v i n g (0-40 meters
depth). A d d i t i o n a l m a t e r i a l resulted f r o m océanographie cruises c a r r i e d out d u r i n g
the " F a u n a I " e x p e d i t i o n s of the project " F a u n a Ibérica" (Iberian Fauna), PB87-0397.
T h i s m a t e r i a l originates f r o m deeper waters, d o w n to 1250 m d e p t h .
T h e m a t e r i a l d e s c r i b e d has been d e p o s i t e d i n the collections of the ' L a b o r a t o r i o
de B i o l o g i a M a r i n a ( Z o o l o g i a ) ' of the U n i v e r s i t y of S e v i l l a , S p a i n . A representative
c o l l e c t i o n has been p l a c e d i n the collections of the N a t i o n a l M u s e u m of N a t u r a l H i s tory (Nationaal Natuurhistorisch M u s e u m , formerly Rijksmuseum v a n Natuurlijke
H i s t o r i e ) , L e i d e n , the N e t h e r l a n d s ; this m a t e r i a l is i n d i c a t e d b y the registration n u m ber ( R M N H C o e l . a n d s l i d e numbers).
I n the descriptions 'thecate' means ' p r o v i d e d w i t h a h y d r o t h e c a ' or 'hydrothecate';
'athecate' indicates a c o n d i t i o n w i t h o u t hydrotheca, or 'ahydrothecate'.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. L eiden 300 (1995)
5
List of the localities and species collected
Station/locality
coordinates
depth
Coastal stations
CI.
El Portil
37°12.40'N 07°07.50'W
0­1 m
Aglaophenia pluma (Linnaeus, 1758); Halopt eris diaphana (Heller, 1868); Kirchenpaueria pinnat a
(Linnaeus, 1758); Ventromma halecioides (Alder, 1859).
C2.
Santa María del Mar
36°11.00'N 06°02.00'W
0­2 m
Aglaophenia oct odont a (Heller, 1868); Kirchenpaueria pinnat a (L innaeus, 1758); Plumularia set acea
(Linnaeus, 1758); Monotheca pulchella (Bale, 1882).
C3.
El Chato
36°31.00'N 06°16.00'W
0­1 m
Kirchenpaueria pinnata (L innaeus, 1758)
C4.
Cabo Roche
36°17.60'N 06°08.00'W
0m
Aglaophenia oct odont a (Heller, 1868); A. pluma (L innaeus, 1758); Gymnangium mont agui (Billard,
1912); Kirchenpaueria pinnata (L innaeus, 1758).
C5.
Canos de Meca
36°11.00'N 06°01.00'W
0m
Aglaophenia oct odont a (Heller, 1868); A. párvula (Bale, 1882); Kirchenpaueria pinnat a (L innaeus,
1758); Plumularia setacea (L innaeus, 1758); Monotheca pulchella (Baie, 1882).
C6.
Torre de la Plata
36°05.50'N 05°50.00'W
2­10 m
Aglaophenia picardi Svoboda, 1979.
C7.
Isla de Tarifa
36°48.00'N 05°36.00'W
0­45 m
Aglaophenia kirchenpaueri (Heller, 1868); A. octodonta (Heller, 1868); A. pluma (L innaeus, 1758);
Gymnangium mont agui (Billard, 1912); Antennella siliquosa (Hincks, 1877); A. secundaria (Gmelin,
1791); Halopteris diaphana (Heller, 1868); Kirchenpaueria pinnata (L innaeus, 1758); H liecht enst erni
(Marktanner­Turneretscher, 1890); Plumularia setacea (L innaeus, 1758).
C8.
Punta Paloma
36°04.00 N 05°44.00'W
8m
Aglaophenia kirchenpaueri (Heller, 1868).
C9.
Punta Carnero
36°04.36'N 05°25.36'W
0­25 m
Aglaophenia kirchenpaueri (Heller, 1868); Antennella secundaria (Gmelin, 1791).
CIO. L a Ballenera­Getares(C)
36°04.54 N 05°25.36'W
3­15 m
Aglaophenia kirchenpaueri (Heller, 1868); A. pluma (Linnaeus, 1758).
C I L San Garcia
36°06.35'N 05°25.36'W
5­28 m
Aglaophenia kirchenpaueri (Heller, 1868); A. pluma (L innaeus, 1758); Gymnangium mont agui (Bil­
lard, 1912); Antennella siliquosa (Hincks, 1877); A. secundaria (Gmelin, 1791); Halopt eris liecht en­
sterni (Marktanner­Turneretscher, 1890); Nemertesia ramosa (L amarck, 1816); Ν. antennina (L in­
naeus, 1758); Pseudoplumaria marocana (Billard, 1930)
C12. ElSaladillo
36°07.26 N 05°25.75 W
5m
Aglaophenia kirchenpaueri (Heller, 1868); A. pluma (L innaeus, 1758).
,
,
/
C13.
C14.
C15.
C16.
C17.
Térmica
,
36°10.50'N 05°25.00'W
9­30 m
Aglaophenia pluma (L innaeus, 1758); Nemertesia irregularis (Queich, 1885).
CE.P.S.A.
36°10.60'N 05°25.80'W
12­25 m
Kirchenpaueria pinnata (L innaeus, 1758).
Crinavis
36°10.10'N 05°23.70'W
5­35 m
Aglaophenia acacia Allman, 1883; A. pluma (Linnaeus, 1758); Plumularia setacea (L innaeus, 1758)
San Felipe
36°09.45'W 05°21.48'W
2­9 m
Aglaophenia picardi (Svoboda, 1979); A. pluma (L innaeus, 1758); Kirchenpaueria pinnata (L innaeus,
1758).
Puerto de Gibraltar
36°08.90'N 05°21.60'W
18­28 m
Aglaophenia kirchenpaueri (Heller, 1868).
C18. L aAtunara
36°10.54'N 05°20.00'W
Halopteris liechtensterni (Marktanner­Turneretscher, 1890).
C19. Torre Carbonera
36°14.60'N 05°17.00'W
Aglaophenia pluma (L innaeus, 1758).
40 m
6m
6
Medel & Vervoort. Plumularian hydroids. Zool. Verh. L eiden 300 (1995)
C20. L a Hacienda
36°14.30 N 05°18.60'W
2­3 m
Aglaophenia octodonta (Heller, 1868); A. pluma (L innaeus, 1758); Halopteris diaphana (Heller, 1868);
Plumularia setacea (L innaeus, 1758).
C21. L aHerradura
36°43.90'N 03°43.70'W
14 m
,
Aglaophenia tubiformis Marktanner­Turneretscher, 1890
C22*. Punta Almina
35°54.00'N 05°16.20'W
20­25 m
Aglaophenia kirchenpaueri (Heller, 1868); A. pluma (L innaeus, 1758); A. tubulifera (Hincks, 1861);
Gymnangium mont agui (Billard, 1912); Antennella siliquosa (Hincks, 1877); A. secundaria (Gmelin,
1791); Halopteris liechtensterni (Marktanner­Turneretscher, 1890).
C24*. Punta Saudino
35°54.35'N 05°17.95'W
38 m
Aglaophenia kirchenpaueri (Heller, 1868); Antennella siliquosa (Hincks, 1877); A. secundaria (Gme­
lin, 1791).
C25*. Aguja del Campo
35°54.20'N 05°19.65 W
20 m
Aglaophenia t ubiformis Marktanner­Turneretscher, 1890; Halopt eris liecht enst erni (Marktanner­
Turneretscher, 1890); Kirchenpaueria pinnata (L innaeus, 1758).
C26*. Punta del Desnarigado
35°53.50'N 05°16.20'W
10 m
/
Antennella siliquosa (Hincks, 1877); A. secundaria (Gmelin, 1791).
C27*. ElPineo
3 5 ° 5 2 . 8 r N 05°19.65 W
,
8­12 m
Aglaophenia t ubiformis Marktanner­Turneretscher, 1890
C28*. Benzu
35°55.10'N 05°23.25'W
25 m
Aglaophenia kirchenpaueri (Heller, 1868); A. pluma (Linnaeus, 1758); Α. t ubiformis Marktanner­
Turneretscher, 1890; Antennella siliquosa (Hincks, 1877); A. secundaria (Gmelin, 1791).
C29.
Punta Europa
C30.
Kirchenpaueria pinnata (L innaeus, 1758); Plumularia setacea (L innaeus, 1758)
Isla Cristina
37°11.86'N 07°11.16'W
8­12 m
Aglaophenia pluma (L innaeus, 1758)
36°06.00'N 05°21.00'W
5­18 m
Campaign F A U N A I
Fl.
Golfo de Huelva
36°51.62'N 07°10.27 W
,
Nemertesia antennina (L innaeus, 1758).
F4.
Málaga
36°40.34'N 04°11.55'W
Streptocaulus dollfusi (Billard, 1924).
F5.
Vêlez Málaga
36°41.93'N 04°04.88 W
Nemertesia irregularis (Queich, 1885).
F8. L a Herradura (Granada)
36°41.06'N 03°48.36'W
/
F13.
F38.
F57.
F58.
F63.
190 m
60 m
67­68 m
238­291 m
Nemertesia irregularis (Queich, 1885).
Cabo Sagra
36°40.89 N 03°22.59 W
62 m
Aglaophenia tubulifera (Hincks, 1861); Nemertesia ramosa (L amarck, 1816).
opposite Estepona
36°18.25'N 05°13.25'W
60­62 m
Nemertesia irregularis (Queich, 1885).
Trafalgar
36°04.84'N 06°01.00'W
76­80 m
Lytocarpia myriophyllum (L innaeus, 1758); Nemertesia ramosa (L amarck, 1816).
Trafalgar
36°08.60'N 06°01.20'W
33­34 m
Nemertesia ramosa (L amarck, 1816).
Trafalgar­Tarifa
36°03.13'N 05°50.58'W
97­118 m
Lytocarpia myriophyllum (L innaeus, 1758); PolyplumariaflabellataG.O. Sars, 1874
,
,
Campaign AR.SA/92
A6.
opposite Chiclana
36°19.70 N 06°29.90 W
84 m
A13.
Streptocaulus dollfusi (Billard, 1924)
Golfo de Cadiz
36°58.50'N 08°00.00'W
Nemertesia ramosa (Lamarck, 1816)
84 m
,
[N.B.: L ocalities in North Africa are marked by asterisk*]
,
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
7
Taxonomie review
Superfamily Plumularioidea Hincks, 1868
Family Aglaopheniidae Broch, 1918
Genus Aglaophenia Lamouroux, 1812
Aglaophenia acacia Allman, 1883
(fig. 1)
Aglaophenia acacia Allman, 1883: 38, pl. 12 figs 1-4; Svoboda, 1979: 79-82, figs 12d, 13d, 14b, 15d(l-5),
16d; Svoboda & Cornelius, 1991:14-16, figs 1,17a-b, 20a-b, 21a-b.
Aglaophenia elongata; Van Gemerden-Hoogeveen, 1965: 79-80, fig. 44.
Material— C15: (21.Í.1989), numerous hydrocauli with corbulae (RMNH Coei. no. 27144, three slides
no. 2562).
Measurements (in mm)
Maximum height of hydrocaulus
146
Internodes of hydrocaulus, length
0.649-0.807
idem, diameter
Hydrocladial internodes, length
0.157-0.342
0.412-0.526
idem, diameter
0.070-0.096
Hydrotheca, length
0.324-0.403
idem, diameter at rim
Length free part adcauline hydrothecal wall
Corbula, number of ribs
idem, length
idem, diameter
0.149-0.245
0.131-0.192
6-8
2.00-3.55
0.925-1.295
D e s c r i p t i o n . — C o l o n y y e l l o w i s h , c o m p o s e d of m o n o s i p h o n i c h y d r o c a u l i , beari n g alternately a r r a n g e d h y d r o c l a d i a , each h y d r o c a u l u s w i t h a basal athecate z o n e
f o l l o w e d b y a prosegment, separated b y o b l i q u e nodes as are also present b e t w e e n
r e m a i n i n g internodes f o r m i n g the axis: those internodes b e a r i n g three nematothecae
a n d a n abortive h y d r o t h e c a . Some of the internodes bear r a m i f i c a t i o n s , n o r m a l l y
there are t w o at about the same l e v e l of the internode, directed left a n d r i g h t a n d
f o r m i n g a n acute angle w i t h m a i n axis that is s l i g h t l y c u r v e d b a c k w a r d s (fig. l a , b).
S e c o n d a r y h y d r o c a u l i w i t h s o m e of the basal internodes w i t h o u t h y d r o c l a d i a , b u t
w i t h c o r m i d i a a l m o s t s i m i l a r to those of the h y d r o c l a d i a ; h o w e v e r , the m e d i a n
nematotheca is p l a c e d o n the l o w e r part of the internode, w i t h o u t contact w i t h the
a b c a u l i n e h y d r o t h e c a l w a l l . Basalmost c o r m i d i u m w i t h t w o m e d i a n nematothecae,
one basal, one i n contact w i t h the abcauline h y d r o t h e c a l w a l l . H y d r o c l a d i a i n s e r t i n g
o n latero-distal a p o p h y s e s of a x i a l internodes a n d alternately t u r n e d left o r right,
each b r o k e n u p into internodes (cormidia) separated b y straight n o d e s ; perisarc
transparent, coenosarc colourless. H y d r o c l a d i a l c o r m i d i a each w i t h s l i g h t l y elongate d h y d r o t h e c a ; h y d r o t h e c a l r i m w i t h n i n e r o u n d e d , s h a l l o w cusps a n d a m o d e r a t e l y
d e v e l o p e d a d c a u l i n e s e p t u m i n basal t h i r d of h y d r o t h e c a l i n t e r i o r (fig. l c ) . M e d i a n
nematotheca short, r e a c h i n g half l e n g t h of h y d r o t h e c a l abcauline w a l l ; free p a r t c.
o n e - t h i r d of total nematothecal l e n g t h , o p e n i n g gutter-shaped. L a t e r a l nematothecae
Fig. 1. Aglaophenia acacia Allman, 1883. a, colony; b, point of branching of main axis; c, two cormidia;
d, (male) corbula. Scales: a, 1 cm; b, 0.5 mm; c, 0.15 mm; d, 1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
9
s u r p a s s i n g h y d r o t h e c a l r i m , s l i g h t l y s w o l l e n ; r i m w i t h deep a d c a u l i n e sinuosity.
M a l e corbulae o n l y have b e e n f o u n d . Costae c o m p l e t e l y f u s e d ; n o free costae
h a v e b e e n o b s e r v e d (fig. I d ) .
S u b s t r a t u m . — M a t e r i a l referred to above f o u n d detached. A c c o r d i n g to S v o b o d a
& C o r n e l i u s (1991) to be f o u n d o n r o c k y s u b s t r a t u m .
D i s t r i b u t i o n . — T r o p i c a l A t l a n t i c (Boero & B o u i l l o n , 1993). D i s t r i b u t e d i n the
n o r t h e r n A t l a n t i c f r o m the coasts of the U n i t e d States ( N o r t h C a r o l i n a , Tortugas,
F l o r i d a , C a r i b b e a n ) (Svoboda & C o r n e l i u s , 1991) to the C a n a r y Islands, the A z o r e s
a n d the coasts of Great B r i t a i n a n d Ireland. A l s o o b s e r v e d i n the M e d i t e r r a n e a n .
D e p t h range 12-822 m (Svoboda & C o r n e l i u s , 1991).
Aglaophenia kirchenpaueri (Heller, 1868)
(fig-2)
Plumularia Kirchenpaueri Heller, 1860: 40, 82, pi. 2 fig. 4.
Aglaophenia kirchenpaueri; Marktanner-Turneretscher, 1890: 263, pi. 7 figs 9,22; Svoboda, 1979: 87-90, figs
12g, 13g, 14c, 15g(l-2), 16g, pi. 5 fig. f; Svoboda & Cornelius, 1991:20-22, figs 4,17c-d, 20d, 21c-d.
Aglaophenia septifera Broch, 1912: 66; Broch, 1913: 6-7, fig. 8.
Material.— C7: (9.VÜ.1990), 14 m, two colonies with corbulae; (10.vii.1990), 25 m, one colony without
corbulae; (10.ix.1990), 25 m, ten colonies with corbulae.— C8: (20.viii.1991), 8 m, two colonies with corbulae.— C9: (19.viii.1991), 0-25 m, five colonies without corbulae —C10: (26.vii.1990), 1-3 m, several colonies.— C l l : (14.vii.1991), 3-5 m, nine colonies with corbulae; (30.vii.1991), 3 m, 40 colonies with corbulae (RMNH Coel. no. 27155, slide no. 2563); (29.X.1944), 6-9 m, many colonies with corbulae.— C12:
(20.vii.1993), 5 m, six colonies with corbulae.— C17: (14.vii.1993), 10-18 m, several colonies with corbulae.— C22: (14.V.1986), 25 m, two colonies with corbulae; (13.V.1986), 35 m, one colony with corbulae —
C24: (6.V.1986), 38 m, four colonies with corbulae — C28: (lO.v.1986), 20 m, four colonies with corbulae.
Measurements (in mm)
Maximum height of hydrocaulus
57
Internodes of hydrocaulus, length
0.18-0.46
idem, diameter
Hydrocladial internodes, length
0.10-0.27
0.31-0.40
idem, diameter
0.11-0.14
Hydrotheca, length
0.28-0.39
idem, diameter at rim
0.21-0.29
idem, length free part adcauline wall
0.08-0.18
Corbula, number of ribs
5-11
idem, length
1.71-3.96
idem, diameter
0.93-1.40
D e s c r i p t i o n . — H y d r o r h i z a reticulate, g i v i n g rise to m o n o s i p h o n i c , y e l l o w i s h ,
u n b r a n c h e d h y d r o c a u l i . Basal p a r t of axis athecate, f o l l o w e d b y u p to three p r o s e g ments, separated b y o b l i q u e nodes a n d p r o v i d e d each w i t h a frontal nematotheca.
Rest of axis c o m p o s e d of h y d r o c l a d i a b e a r i n g internodes, separated b y o b l i q u e
n o d e s a n d each b e a r i n g a latero-distal a p o p h y s i s , three nematothecae, of w h i c h one
m e d i a n a n d t w o latero-distal, a n d a n abortive h y d r o t h e c a o r p s e u d o n e m a t o t h e c a .
H y d r o c l a d i a , i n s e r t i n g o n apophyses, alternately t u r n e d left a n d r i g h t (fig. 2a).
H y d r o c l a d i a also y e l l o w i s h , b r o k e n u p into c o r m i d i a b y means of transverse
Fig. 2. Aglaophenia kirchenpaueri (Heller, 1868). a, colony; b, three cormidia; c, (male) corbula. Scales: a,
1 cm; b, 0.2 mm; c, 0.5 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
11
nodes, each w i t h one h y d r o t h e c a a n d three nematothecae: one m e d i a n , t w o lateral.
H y d r o t h e c a f a i r l y s h a l l o w , w i t h b i g abcauline s e p t u m almost transversing interior of
h y d r o t h e c a ; a d c a u l i n e w a l l c u r v e d (fig. 2b). H y d r o t h e c a l r i m w i t h n i n e cusps, of
w h i c h m e d i a n i s the largest. First p a i r o n b o t h sides distinct, r e m a i n i n g m a r g i n a l
cusps g r a d i n g t o w a r d s axis of c o r m i d i u m . M e d i a n nematotheca n o t s u r p a s s i n g
h y d r o t h e c a l r i m , adnate f o r about t w o - t h i r d s of its length, o p e n i n g
L a t e r a l nematothecae projecting
s l i g h t l y above h y d r o t h e c a l
c u r v e d b a c k w a r d s , w i t h gutter-shaped
gutter-shaped.
r i m ; distal portion
aperture.
B o t h m a l e a n d female corbulae have b e e n observed. M a l e corbulae p a r t l y o p e n
a n d w i t h o u t free costae (fig. 2c); female corbulae u s u a l l y closed a n d w i t h free costa.
S u b s t r a t u m . — F o u n d associated w i t h algae a n d o n h a r d substrata l i k e Balanus,
B r y o z o a a n d stones. A l s o f o u n d o n G o r g o n a r i a a n d calcareous algae (Svoboda &
C o r n e l i u s , 1991).
D i s t r i b u t i o n . — M e d i t e r r a n e a n a n d L u s i t a n i a n species ( R a m i l & V e r v o o r t , 1992a).
R e c o r d e d f r o m the B r i t i s h coasts, M o r o c c o , C a p e Verde Islands a n d the A d r i a t i c Sea
(Patriti, 1970; S v o b o d a & C o r n e l i u s , 1991). Frequent i n the littoral zone. M a x i m u m
d e p t h r e c o r d 373 m ( R a m i l & Vervoort, 1992a).
Aglaophenia octodonta (Heller, 1868)
(fig. 3)
Plumularia octodonta Heller, 1868: 40,82, pi. 2 fig. 3.
Aglaophenia helleri Marktanner-Turneretscher, 1890: 271, pi. 7 figs 3,13-16 (including the variety).
Aglaopheniafilicula;Kühn, 1909:452, fig. Ua, pi. 21 figs 64-69, pi. 22 figs 40-74.
Aglaophenia adriática Babié, 1911: 541, figs 1-2.
Aglaophenia pluma var. helleri; Bedot, 1919: 265.
Aglaophenia octodonta; Svoboda, 1979,1979: 65-70, figs 12a, 13a, 15a, 16a, pi. 5c; Svoboda & Cornelius,
1991: 23-25, fig. 6.
Material.— C2: (19.viii.1989), intertidal zone, various colonies without corbulae; (3.iii.l991), intertidal
zone, various colonies with corbulae ( R M N H Coel. no. 27116, two slides no. 2564).— C4: (23.vii.1989),
tidal pool, several colonies with corbulae; (12.xi.1989), tidal pool, 10 colonies without corbulae;
(9.ÏX.1990), from material cast ashore, numerous colonies with corbulae; (17.vii.1993), tidal pool, several colonies with corbulae.— C5: (ll.ii.1990), 0 m, various colonies with corbulae — C7: (18.V.1990), 0
m, several colonies with corbulae; (17.ii.1991), 3-4 m, various colonies without corbulae.— C20:
(3.Ü.1991), 2-3 m, several colonies with corbulae.
Measurements (in mm)
Maximum height of hydrocaulus
40
Internodes of hydrocaulus, length
0.245-0.394
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrothecae, total length
0.122-0.219
0.228-0.333
0.060-0.087
0.24-0.31
idem, diameter at rim
0.14-0.17
idem, length of free part abcauline wall
0.06-0.10
Corbula, number of costae
7-8
idem, total length
1.35-2.35
idem, diameter
0.80-1.80
Fig. 3. Aglaophenia octodonta (Heller, 1868). a, two colonies; b, basal part of axis; c, three cormidia; d, (female) corbula. Scales: a, 1 cm; b, c, 0.15 mm; d; 0.5 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
13
D e s c r i p t i o n . — H y d r o r h i z a reticulate, g i v i n g rise to m o n o s i p h o n i c , y e l l o w h y d r o c a u l i that n o r m a l l y are u n b r a n c h e d (fig. 3a). Basal part of h y d r o c a u l u s athecate, of
v a r i e d l e n g t h a n d w i t h several transverse nodes, f o l l o w e d b y 1-3 prosegments separated b y o b l i q u e nodes (fig. 3b). Rest of h y d r o c a u l u s c o m p o s e d of h y d r o c l a d i a beari n g internodes separated b y o b l i q u e nodes a n d b e a r i n g a latero-distal a p o p h y s i s ,
three nematothecae a n d a n abortive h y d r o t h e c a (pseudonematotheca).
H y d r o c l a d i a inserted o n a p o p h y s e s a n d alternately directed left or r i g h t (fig. 3a),
transparent o r y e l l o w i s h , c o m p o s e d of c o r m i d i a separated b y transverse nodes.
H y d r o t h e c a n a r r o w i n g b a s a l l y a n d thus of t r i a n g u l a r shape. H y d r o t h e c a l r i m w i t h 9
s t r o n g cusps, of w h i c h u n p a i r e d abcauline t o o t h is occasionally c u r v e d interiorly.
W e l l d e v e l o p e d a d c a u l i n e s e p t u m present i n l o w e r t h i r d of h y d r o t h e c a l c a v i t y (fig.
3c). M e d i a n nematotheca adnate for t w o - t h i r d s of its length, c o v e r i n g large p o r t i o n
of a b c a u l i n e h y d r o t h e c a l w a l l : o n l y a s m a l l part remains free. O p e n i n g of m e d i a n
nematotheca deep gutter-shaped; e n d of nematotheca a l m o s t at l e v e l of h y d r o t h e c a l
r i m . L a t e r a l nematothecae s m a l l , a p i c a l part s w o l l e n , r e a c h i n g h y d r o t h e c a l r i m ; aperture gutter-shaped.
B o t h m a l e a n d female corbulae h a v e been observed. M a l e corbulae w i t h o u t free
costa a n d u s u a l l y p a r t l y o p e n ; female corbulae u s u a l l y f u l l y closed, w i t h one w e l l
d e v e l o p e d free costa (fig. 3d).
S u b s t r a t u m . — O n shells of Mytilus, o n Balanus, stones a n d algae (e.g. Sargassum).
A l s o r e c o r d e d f r o m sponges, Cystoseira spec, a n d Posidonia spec. ( S v o b o d a & C o r n e l i us, 1991), f r o m A s c i d i a ( R a m i l , 1988) a n d f r o m Vermetus s p e c , calcareous algae a n d
f r o m Zostera spec. (Garcia-Carrascosa, 1981).
D i s t r i b u t i o n . — N o r t h e a s t e r n A t l a n t i c a n d M e d i t e r r a n e a n . I n the A t l a n t i c occurr i n g f r o m the E n g l i s h coasts s o u t h w a r d u n t i l Senegal ( S v o b o d a & C o r n e l i u s , 1991).
M a x i m u m d e p t h 30 m (Stechow, 1919), b u t m u c h m o r e c o m m o n i n s h a l l o w waters
(Roca M a r t i n e z , 1986).
Aglaophenia párvula Bale, 1882
(fig-4)
Aglaophenia párvula Bale, 1882: 35-36, pi. 14 fig. 3; Patriti, 1970: 49, fig. 67; Svoboda & Cornelius, 1991:
25-29, figs 7-9, lOa-e, 13h, 18, 22-23.
Aglaophenia heterodonta Jäderholm, 1904, 296-297, pl. 13 fig. 10, pl. 14 fig. 1.
Aglaophenia pluma var. párvula; Millard, 1957: 239-240, fig. 15D-F.
Aglaophenia dichotoma var. heterodonta; Patriti, 1970:48, fig. 66.
Material.— C5: (22.xii.1991), 0 m, various colonies without corbulae.
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrothecae, total length
idem, diameter at rim
idem, length free part abcauline wall
17
0.84 -0.921
0.175-0.192
0.219-0.236
0.061-0.070
0.192-0.263
0.175-0.184
0.061-0.078
Fig. 4. Aglaophenia párvula Bale, 1882. a, colony; b, basal part of stem with prosegment and first hydrocladia; c, three cormidia. Scales: a, 1 cm; b, c, 0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
15
D e s c r i p t i o n . — H y d r o r h i z a reticulate, g i v i n g rise to m o n o s i p h o n i c , i r r e g u l a r l y
b r a n c h e d , y e l l o w to b r o w n h y d r o c a u l i (fig. 4a). Basal part of h y d r o c a u l u s athecate,
w i t h i n o l d e r stems some transverse a n d one o b l i q u e n o d e , f o l l o w e d b y a p r o s e g m e n t l i m i t e d b y quite o b l i q u e nodes (fig. 4b). A f t e r this there is a n o r m a l sequence of
h y d r o c l a d i a b e a r i n g internodes each w i t h a latero-distal a p o p h y s i s , three nematothecae
a n d o n e abortive
hydrotheca
(pseudonematotheca).
Some
h y d r o c a u l i are
b r a n c h e d ; the s e c o n d a r y axes each h a v e a prosegment.
H y d r o c l a d i a s u p p o r t e d b y the apophyses, alternately directed left o r right, a n d
y e l l o w i s h , b r o k e n u p i n t o c o r m i d i a separated b y straight septa. H y d r o t h e c a e f a i r l y
deep, n a r r o w i n g b a s a l l y a n d consequently t r i a n g u l a r ; r i m w i t h n i n e quite u n e q u a l
m a r g i n a l teeth (fig. 4c). M e d i a n , abcauline tooth flexed interiorly, almost h i d d e n b y
first p a i r of cusps; t h i r d p a i r w i t h b i f i d apex, d e v e l o p m e n t of i n c i s i o n v a r i e d , s o m e times f a i r l y deep ( g i v i n g the i m p r e s s i o n of 11 teeth b e i n g present), sometimes scarcel y i n d i c a t e d . Basal t h i r d of interior of h y d r o t h e c a w i t h distinct, w e l l d e v e l o p e d septum,
a l m o s t r e a c h i n g opposite h y d r o t h e c a l w a l l . L a t e r a l nematothecae short, n o t
reaching hydrothecal r i m .
N o corbulae h a v e been f o u n d .
S u b s t r a t u m . — O n Cacospongia spec. A c c o r d i n g to S v o b o d a a n d C o r n e l i u s (1991)
this species occurs frequently o n sponges, t h o u g h other substrata have also b e e n
r e c o r d e d (algae, Mytilus shells, C i r r i p e d i a , etc.).
D i s t r i b u t i o n . — Eastern A t l a n t i c , f r o m the coasts of S o u t h A f r i c a n o r t h w a r d t i l l
the B r i t i s h coasts. N o t k n o w n f r o m the M e d i t e r r a n e a n . A l s o c o m m o n o n s o u t h e r n
A u s t r a l i a n coasts (Svoboda & C o r n e l i u s , 1991; G i l i , V e r v o o r t & Pages, 1989). D e p t h
records f r o m 0 meters ( R a m i l , 1988) to 350 m ( G i l i , V e r v o o r t & Pages, 1989).
Aglaophenia picardi S v o b o d a , 1979
(fig. 5)
Aglaophenia picardi Svoboda, 1979: 70-74, figs 12b, 13b, 15b, 16b; Svoboda & Cornelius, 1991: 29-30, fig.
11.
Material.— C6: (22.ix.1991), 6 m, 18 colonies with corbulae (RMNH Coel. no. 27177, slide no. 2565).—
C16: (18.vii.1991), 3-5 m, 15 colonies with corbulae.
Measurements (in mm)
Maximum height of hydrocaulus
28
Internode of hydrocaulus, length
idem, diameter
0.271-0.412
Hydrocladial internode, length
0.280-0.342
0.087-0.175
idem, diameter
0.052-0.078
Hydrotheca, total length
0.245-0.280
idem, diameter at rim
0.14-0.16
idem, length free part abcauline wall
0.105-0.49
Corbula, number of costae
idem, total length
idem, diameter
3-7
1.26-1.8
0.62-0.8
Fig. 5. Aglaophenia picardi Svoboda, 1979. a, three colonies springing from same stolon; b, basal part of
axis; c, three cormidia; d, (male) corbula. Scales: a, 1 cm; b, 0.5 mm; c, 0.15 mm; d, 0.2 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
17
D e s c r i p t i o n . — H y d r o r h i z a s l i g h t l y reticulate, b e a r i n g u n b r a n c h e d , b r o w n , m o n o s i p h o n i c h y d r o c a u l i (fig. 5a). Basal part of axis athecate, occasionally w i t h i r r e g u l a r l y
o c c u r r i n g transverse nodes; there are n o prosegments (fig. 5b). Rest of axis c o m p o s e d
of h y d r o c l a d i a b e a r i n g internodes that have a latero-distal a p o p h y s i s , three n e m a tothecae a n d a r e d u c e d h y d r o t h e c a (pseudonematotheca);
they are separated b y
o b l i q u e nodes.
H y d r o c l a d i a b o r n o n the apophyses, alternately directed left a n d right, c o m p o s e d of c o r m i d i a w i t h transparent perisarc separated b y transverse nodes. H y d r o theca f a i r l y deep, r i m w i t h n i n e distinct teeth, p a r t i c u l a r l y o n frontal side of h y d r o theca u s u a l l y s h a r p l y p o i n t e d , a n d separated b y r o u n d e d e m b a y m e n t s
(fig. 5c).
A d c a u l i n e s e p t u m i n l o w e r t h i r d of h y d r o t h e c a present t h o u g h a p p a r e n t l y occasiona l l y r e d u c e d . A d n a t e part of m e d i a n nematotheca c o v e r i n g s l i g h t l y m o r e t h a n half
l e n g t h of abcauline h y d r o t h e c a l w a l l ; free p o r t i o n short, n o t r e a c h i n g h y d r o t h e c a l
r i m , o p e n i n g gutter-shaped. L a t e r a l nematothecae also w i t h gutter-shaped aperture,
just r e a c h i n g h y d r o t h e c a l r i m .
C o r b u l a e inspected a p p a r e n t l y m a l e , b e i n g i n majority o p e n , i.e. w i t h large o p e n ings b e t w e e n the pairs of opposite costae (fig. 5d). T h e female corbulae are closed
w i t h the p r o x i m a l p a i r of ribs almost free.
S u b s t r a t u m . — O n h a r d concretions, C i r r i p e d i a , etc. A l s o f o u n d o n Mytilus, o n
sponges, o n r e d algae a n d occasionally o n Posidonia oceânica (Svoboda & C o r n e l i u s ,
1991).
D i s t r i b u t i o n . — N o r t h e a s t e r n A t l a n t i c , i n the M e d i t e r r a n e a n a n d off Senegal; also
o b s e r v e d i n the B l a c k Sea (Isasi U r d a n g a r i n , 1985; S v o b o d a & C o r n e l i u s , 1991).
T h o u g h u s u a l l y f o u n d b e t w e e n 3 a n d 10 m , it also lives at 30 m d e p t h ( S v o b o d a &
C o r n e l i u s , 1991).
Aglaophenia pluma (Linnaeus, 1758)
(fig- 6)
Sertularia pluma Linnaeus, 1758: 811.
Aglaophenia pluma; Svoboda, 1979: 98-102, figs 15j(l-4); Svoboda & Cornelius, 1991: 30-34. figs lOf, 12,
13a-g, 19a-b, 24a-b.
Aglaophenia pluma p.p. Hincks, 1868: 286-288, fig. 37, pi. 63 fig. 1
Plumularia cristata Lamarck, 1816:125
Aglaophenia cristata; McCrady, 1859: 202-203.
Material.— C l : (9.XÜ.1988), intertidal zone, numerous colonies with corbulae; (5.iii.l989), intertidal
zone, numerous colonies with corbulae.— C4: (20.X.1998), in material cast ashore, numerous colonies
with corbulae; (5.VÜ.1992), in fishing nets, several colonies with corbulae.— C7: (8.viii.l993), 15 m,
several colonies with corbulae.— C10: (6.viii.l991), 12 m, one colony with corbulae.— C l l :
(17.vii.1991), 5 m, one colony with corbulae.— C12: (20.vii.1993), 5 m, six colonies with corbulae —
C13: (27.Í.1992), 8 m, numerous colonies with corbulae (RMNH Coel. no. 27188, three slides no.
2566).— C15: (3.VÜ.1990), 15 m, ten colonies with corbulae.— C16: (27.vii.1991), 6 m, 20 colonies with
corbulae.— C19: (21.ix.1985), 6 m, two fragments without corbulae.— C20: (3.Ü.1991), 2-3 m, various
colonies with corbulae.— C22: (14.V.1986), 25 m, numerous colonies with corbulae.— C28: (lO.v.1986),
25 m, several colonies with corbulae.— C29: (14.vii.1993), 5-18 m, various colonies with corbulae.—
C30: (3.VÜ.1990), 8 m, numerous colonies with corbulae.
18
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrotheca, total length
idem, diameter at rim
idem, length free part abcauline wall
120
0.447-0.552
0.184-0.271
0.280-0.426
0.061-0.122
0.29 -0.35
0.15 -0.23
0.080-0.122
Corbula, number of costae
4-7
idem, total length
1.511-2.311
0.638-1.298
idem, diameter
D e s c r i p t i o n . — H y d r o c a u l i m o n o s i p h o n i c , unbranched or d i c h o t o m o u s l y branched,
r i s i n g f r o m h y d r o r h i z a l n e t w o r k u s u a l l y o n algae, but also o n fixed objects (fig. 6a).
H y d r o c a u l i w i t h basal athecate p o r t i o n , f o l l o w e d b y one or t w o prosegments; rest of
h y d r o c a u l u s m a d e u p of internodes w i t h latero-distal apophysis, three nematothecae
a n d a n abortive hydrotheca (pseudonematotheca). Prosegments a n d internodes separated b y oblique nodes.
H y d r o c l a d i a i n s e r t i n g o n a p o p h y s e s a n d alternately d i r e c t e d left a n d r i g h t , c o m p o s e d of u s u a l l y w h i t i s h c o r m i d i a separated b y transverse nodes. H y d r o t h e c a e f a i r l y
deep, p l a n e of r i m o b l i q u e a n d i n c l i n e d d o w n w a r d s , r i m w i t h n i n e distinct, m o r e o r
less e q u a l teeth (fig. 6b). Intrathecal a d c a u l i n e s e p t u m i n l o w e r t h i r d of h y d r o t h e c a l
c a v i t y u s u a l l y w e l l d e v e l o p e d . M e d i a n nematotheca c o v e r i n g at least h a l f the abcaul i n e h y d r o t h e c a l w a l l , short, not r e a c h i n g h y d r o t h e c a l r i m , w i t h gutter-shaped o p e n i n g ; f o r a m e n o p e n i n g into h y d r o t h e c a l c a v i t y u s u a l l y distinct. L a t e r a l nematothecae
r e a c h i n g l e v e l of h y d r o t h e c a l cusps, w i t h deep e m b a y m e n t o n a x i a l side.
C o r b u l a e often y e l l o w i s h , m a l e as w e l l as female corbulae often w i t h free costa.
M a l e corbulae closed w i t h slit l i k e o p e n i n g s b e t w e e n the costae; female corbulae
w i t h f u s e d costae a n d s m a l l e r slits (fig. 6c).
S u b s t r a t u m . — O n r o c k y b o t t o m , o n algae a n d sponges. A c c o r d i n g to S v o b o d a &
C o r n e l i u s (1991) c o m m o n o n the b r o w n alga Halidrys siliquosa, b u t also o n Posidonia,
Sargassum, Laminaria, Cystoseira a n d Gelidium. A l s o f o u n d o n shells, other h y d r o i d s ,
C r u s t a c e a a n d calcareous concretions.
D i s t r i b u t i o n . — C o s m o p o l i t a n ( R a m i l , 1988; A l v a r e z C l a u d i o , 1993; Boero &
B o u i l l o n , 1993). F r o m i n t e r t i d a l z o n e d o w n to 100 m d e p t h ( A l v a r e z C l a u d i o , 1993).
D i s c u s s i o n . — A s S v o b o d a & C o r n e l i u s (1991) p o i n t out, this species has n o spec i a l d i a g n o s t i c characters. Because of this a n d its frequent occurrence, the differences
w i t h other species of Aglaophenia are not a l w a y s quite clear. M a n y f o r m s a n d v a r i e ties h a v e b e e n d e s c r i b e d i n the course of time (Bedot, 1919; M i l l a r d , 1975), b a s e d o n
m o d e of r a m i f i c a t i o n , l e n g t h of the h y d r o t h e c a , d e v e l o p m e n t of m a r g i n a l cusps a n d
l e n g t h of m e d i a n nematotheca. A c c o r d i n g to S v o b o d a (1979) the absence of z o o x a n thellae i n A. pluma d i s t i n g u i s h e s this species f r o m the related A. tubiformis, w h e r e
z o o x a n t h e l l a e are present. A c c o r d i n g to several authors (Roca M a r t i n e z , 1986; A l v a rez C l a u d i o , 1993) this character is insufficient a n d n o t absolutely d i s c r i m i n a t o r y
b e t w e e n the t w o species w h i l s t G i l i (1986) m e n t i o n s observations o n l i v i n g A. pluma
w i t h zooxanthellae. S v o b o d a & C o r n e l i u s (1991) p o i n t o u t that the absence (or
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Fig. 6. Aglaophenia pluma (Linnaeus, 1758). a, colony; b, three cormidia; c, (female) corbula. Scales:
cm; b, 0.2 mm; c, 0.7 mm.
20
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
i m p o s s i b i l i t y to demonstrate the presence) of zooxanthellae i n p r e s e r v e d m a t e r i a l
cannot be c o n s i d e r e d c o n c l u s i v e evidence, n o r is it i n s u c h cases w h e r e the tissue has
b e c o m e lost. T h i s also a p p l i e s to m a t e r i a l treated w i t h acids. S v o b o d a & C o r n e l i u s
(1991: 46-47, tab. I) present a c o m b i n a t i o n of characters to d i s c r i m i n a t e b e t w e e n the
t w o species, b u t some of those characters s h o w overlaps i n o u r m a t e r i a l .
C o m p a r i n g o u r m a t e r i a l w i t h zooxanthellae w i t h colonies w i t h o u t w e observe
that:
1, the i n d e x l e n g t h versus diameter of the h y d r o t h e c a i n general terms does n o t
c o r r e s p o n d to that g i v e n b y S v o b o d a a n d C o r n e l i u s (1.3-1.5 i n A. pluma a n d 1.5-2 i n
A. tubiformis);
2, the distance b e t w e e n the h y d r o c l a d i a is different (shorter i n A. pluma t h a n i n A.
tubiformis), n o r does the h o m o g e n e i t y of the m a r g i n a l cusps fit (unequal i n A. pluma,
e q u a l i n A. tubiformis). A s far as the n u m b e r of prosegments is concerned: one is the
m o s t frequent n u m b e r i n b o t h species, t h o u g h t w o h a v e occasionally b e e n c o u n t e d
i n A. tubiformis a n d u p to f o u r i n A. pluma (1-3 i n A. pluma a n d 1 i n A. tubiformis
a c c o r d i n g to S v o b o d a & C o r n e l i u s , 1991). T h e c o r b u l a i n A pluma u s u a l l y has n o free
costa; s u c h a costa is u s u a l l y present i n A. tubiformis, t h o u g h w i t h exceptions. A relat i v e l y constant feature is the difference i n c o l o u r : i n A. pluma the d a r k b r o w n c o l o u r
of the perisarc of the h y d r o c a u l u s contrasts w i t h the y e l l o w i s h c o l o u r o r transparenc y of that of the h y d r o c l a d i a b o t h i n l i v i n g a n d freshly preserved material. T h e coenosarc is w h i t e to y e l l o w . I n A. tubiformis h y d r o c a u l u s a n d h y d r o c l a d i a are y e l l o w i s h .
T h e coenosarc is y e l l o w i s h i n freshly p r e s e r v e d m a t e r i a l a n d greenish i n l i v i n g colonies. D i c h o t o m o u s r a m i f i c a t i o n occurs i n b o t h species. T h i s r a m i f i c a t i o n , a c c o r d i n g
to S v o b o d a (1976,1979) d e p e n d s u p o n the w a t e r temperature, w h i c h i n b o t h species
has a d e c i d e d i n f l u e n c e o n c o l o n y d e v e l o p m e n t . I n o u r o p i n i o n d i s t i n c t i o n b e t w e e n
b o t h species rests w i t h presence o r absence of zooxanthellae a n d c o l o r a t i o n , w h i l e
presence or absence of a free c o r b u l a r i b or n u m b e r of prosegments are of s e c o n d a r y
importance.
Aglaophenia tubiformis Marktanner-Turneretscher, 1980
(%· 7)
Aglaophenia tubiformis Marktanner-Turneretscher, 1890: 269-270, pi. 7 figs 4,5, 6,17 (including variety);
Svoboda, 1979: 90-95, figs 12h, 13h, 14d, 15h, 16h, pis 5a, b, 6, 8, 9a(l,2); Svoboda & Cornelius,
1991: 34-36, figs 14, 25a (complete synonymy).
Material.— C21: (27.vi.1993), 1-5 m, numerous hydrocauli with corbulae (RMNH Coel. no. 27119, two
slides no. 2567).— C25: (5.V.1986), 20 m, 20 colonies without corbulae.— C27: (7.V.1986), 8-10 m, 15
colonies with corbulae.— C28: (9.V.1986), 0-3 m, six fragments with corbulae; (12.V.1986), 12 m, three
colonies without corbulae; (8.v.1986), 10 m, several colonies with corbulae.
Measurements (in mm)
Maximum height of hydrocauli
Internodes of hydrocaulus, length
idem, diameter
114
0.405-0.745
0.184-0.315
Hydrocladial internodes, length
0.359-0.596
idem, diameter
Hydrotheca, total length
0.070-0.140
0.30-0.40
Fig. 7. Aglaophenia tubiformis Marktanner-Turneretscher, 1890. a, colony; b, three cormidia; c, (female)
corbula. Scales: a, 1 cm; b, 0.15 mm; c, 1 mm.
22
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
idem, diameter at
rim
0.15-0.22
length of free part adcauline wall
Corbula, number of
0.131-0.184
ribs
7-11
total length
2.71-3.91
diameter
0.97-1.40
D e s c r i p t i o n . — C o l o n i e s w i t h reticulate h y d r o r h i z a f r o m w h i c h rise m o n o s i p h o n ic, d i c h o t o m o u s l y b r a n c h e d or u n b r a n c h e d , y e l l o w i s h h y d r o c a u l i (fig. 7a). B a s a l p o r t i o n of h y d r o c a u l u s athecate, w i t h some transverse nodes, f o l l o w e d b y one (or rarely
two) prosegments separated b y o b l i q u e node(s); the rest of the h y d r o c a u l u s b e i n g
m a d e u p of internodes separated b y transverse nodes a n d b e a r i n g the u s u a l l y laterod i s t a l a p o p h y s i s , three nematotheca a n d a r e d u c e d h y d r o t h e c a (pseudonematotheca). T h e a p o p h y s e s s u p p o r t y e l l o w i s h h y d r o c l a d i a alternately directed left o r r i g h t
a n d b r o k e n u p into c o r m i d i a b y means of straight nodes. C o r m i d i u m m u c h as i n A.
pluma, w i t h a deep h y d r o t h e c a w i t h m o d e r a t e l y d e v e l o p e d a d c a u l i n e s e p t u m i n
l o w e r t h i r d of h y d r o t h e c a l cavity; r i m of h y d r o t h e c a w i t h n i n e m o r e o r less e q u a l
cusps (fig. 7b). M e d i a n nematotheca c o v e r i n g half l e n g t h of abcauline h y d r o t h e c a l
w a l l or s l i g h t l y m o r e ; free p o r t i o n p o i n t i n g a w a y f r o m h y d r o t h e c a , of v a r i e d l e n g t h
a n d occasionally as l o n g as free part abcauline w a l l ; aperture gutter-shaped. L a t e r a l
nematothecae r e a c h i n g l e v e l of m a r g i n a l cusps or s l i g h t l y s u r p a s s i n g . A l l coenosarc
f i l l e d w i t h b r o w n i s h zooxanthellae.
C o r b u l a e y e l l o w i s h , m a l e w i t h slits b e t w e e n ribs; female gonothecae w i t h a l m o s t
c o m p l e t e l y f u s e d ribs, o n l y m i n o r slits a n d a (basal) free costa (fig. 7c).
S u b s t r a t u m . — T h e present specimens w e r e collected o n algae; also f o u n d o n C y s toseira, Sargassum a n d Halidrys as w e l l as o n r o c k y a n d s a n d y b o t t o m s ( R a m i l , 1988;
S v o b o d a & C o r n e l i u s , 1991).
D i s t r i b u t i o n . — N o r t h e a s t e r n A t l a n t i c f r o m M o r o c c o n o r t h w a r d t i l l the B r i t i s h
coasts; also i n the M e d i t e r r a n e a n (Svoboda & C o r n e l i u s , 1991). F o r d i s c u s s i o n see A.
pluma.
Aglaophenia tubulifera ( H i n c k s , 1861)
(fig. 8)
Plumularia tubulifera Hincks, 1861: 256, pi. 7 figs 1-2.
Aglaophenia tubulifera; Hincks, 1868: 288-289, pi. 63 fig. 2, pi. 64 fig. 3; Svoboda, 1979: 86-87, figs 12f,
13f, 15f(l-4), 16f, pi. 5 figs g-i; Svoboda & Cornelius, 1991: 36-38, figs 15-16,19c-d, 24c-d.
Material— C22: (14.V.1986), 25 m, three fragments without corbulae; (4.V.1986), 40-42 m, 5 colonies
with corbulae (RMNH Coel. no. 27120, slide no. 2568).— F13: (ll.vii.1989), several fragments with corbulae.
Measurements (in mm)
Maximum height of hydrocaulus
50
Internodes of hydrocaulus, length
0.245-0.570
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrotheca, length
idem, diameter at rim
0.219-0.429
0.298-0.377
0.087-0.131
0.263-0.324
0.140-0.175
Fig. 8. Aglaophenia tubulifera (Hincks, 1861). a, colony; b, basal part of axis; c, three cormidia; d, (male)
corbula. Scales: a, 1 cm; b, 0.2 mm; c, 0.3 mm; d, 1 mm.
24
Medel «Sc Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
idem, length of free part abcauline wall
Corbula, number of ribs
idem, total length
idem, diameter
0.114-0.157
7-8
2.57-2.70
1.0-1.2
Description.— Colony composed
of m o n o s i p h o n i c ,
y e l l o w or b r o w n , u n -
b r a n c h e d h y d r o c a u l i r i s i n g f r o m a creeping, occasionally b r a n c h e d h y d r o r h i z a (fig.
8a). A x i s b^sally w i t h athecate p o r t i o n , occasionally w i t h s o m e straight nodes, f o l l o w e d b y a single prosegment w i t h o b l i q u e nodes (fig. 8b). Rest of axis f o r m e d b y
short internodes separated b y straight septa each w i t h a latero- d i s t a l a p o p h y s i s ,
three nematothecae a n d a r e d u c e d h y d r o t h e c a (pseudonematotheca).
Hydrocladia,
s u p p o r t e d b y the apophyses, are alternately directed left o r r i g h t (fig. 8a) a n d are
closely p a c k e d ; each b e i n g c o m p o s e d of a large n u m b e r of c o r m i d i a separated b y
transverse nodes. H y d r o t h e c a o n l y m o d e r a t e l y deep, w i t h w e e k l y d e v e l o p e d a d c a u l i n e s e p t u m i n the l o w e r t h i r d of the h y d r o t h e c a l cavity; h y d r o t h e c a l r i m w i t h 9 m a r g i n a l cusps of a l m o s t e q u a l size (fig. 8c). L a t e r a l a n d m e d i a n nematothecae w i t h circ u l a r r i m w i t h o u t e m b a y m e n t . M e d i a n nematotheca c o v e r i n g c. half the abcauline
h y d r o t h e c a l w a l l or s l i g h t l y more, directed a w a y f r o m h y d r o t h e c a ; free part of v a r i e d
length, u s u a l l y m u c h longer t h a n free part abcauline w a l l (fig. 8c). L e n g t h of free
p o r t i o n of m e d i a n h y d r o t h e c a v a r i e d i n the h y d r o c l a d i u m a n d over the w h o l e colon y ; the longest b e i n g f o u n d i n the d i s t a l part. D i s t a l t h i r d of free part m e d i a n n e m a totheca w i t h a n n u l a r t h i c k e n i n g . L a t e r a l nematothecae projecting s o m e
distance
above h y d r o t h e c a l r i m .
M a l e c o r b u l a w i t h free costae (fig. 8d); female corbulae m o r e closed b y d i s t a l
extension of costae; free ribs occasionally present i n b o t h sexes.
S u b s t r a t u m . — D e t a c h e d h y d r o c a u l i o n l y h a v e so far been s t u d i e d . T h e species
has b e e n o b s e r v e d o n rocks, stones, algae, shells, other h y d r o i d s a n d tubes of p o l y chaetes ( H i n c k s , 1868; R a m i l , 1988; C o r n e l i u s & R y l a n d , 1990).
D i s t r i b u t i o n . — N o r t h e a s t e r n A t l a n t i c , f r o m the coasts of the B r i t i s h Isles s o u t h w a r d u n t i l G u i n e a Bissau. A l s o cited f r o m the M e d i t e r r a n e a n , the A l b o r a n Sea a n d
eastern M o r o c c o ( G i l i , V e r v o o r t & Pages, 1989; T e m p l a d o et a l . , 1986; S v o b o d a & C o r n e l i u s , 1991). M a x i m u m d e p t h 1200 m ( R a m i l & Vervoort, 1992a).
Genus Streptocaulus Allman, 1883
Streptocaulus dollfusi ( B i l l a r d , 1924)
(figs 9,10)
Cladocarpus dollfusi Billard, 1924: 87-89, fig. 1; Billard, 1934: 229-230, figs 3-4.
Cladocarpus ventricosus; Vervoort, 1959: 300-302, fig. 49.
Material — A6: (U.x.1992), 84 m, several stems, one with single phylactocarp ( R M N H Coel. no. 27121,
slide no. 2569).— F4: (10.vii.1989), 60 m, one fragment.
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length
400
1.00-1.05
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrothecae, total length
idem, diameter at rim
Gonotheca, length
25
0.15-0.16
0.55-0.61
0.08-0.10
0.38-0.45
0.17-0.20
idem, diameter
Phylactocarp, length
0.68
0.19
1.50
D e s c r i p t i o n . — H y d r o c a u l i l o n g , p o l y s i p h o n i c , y e l l o w a n d rarely i r r e g u l a r l y
b r a n c h e d . O n l y the central tube of h y d r o c a u l u s b e a r i n g h y d r o c l a d i a (fig. 9a). T h i s
central axis is i n d i s t i n c t l y d i v i d e d into internodes b y means of straight septa, at
times
scarcely v i s i b l e . E a c h internode w i t h a r o w a several (frontal) nematothecae
a n d a lateral a p o p h y s i s s u p p o r t i n g the h y d r o c l a d i u m . A x i a l nematothecae w i t h t w o
apertures, one t e r m i n a l a n d one basal at the a d c a u l i n e w a l l . N u m b e r of nematothecae per i n t e r n o d e v a r i a b l e , u s u a l l y three to five. O n e of these nematothecae, of s m a l l er size, i n the a x i l of the a p o p h y s i s , occasionally s u c h a s m a l l nematotheca
also
occurs besides the a p o p h y s i s . H y d r o c l a d i a alternately directed left o r r i g h t (fig. 9a);
some internodes w i t h o u t h y d r o c l a d i a . Internodes of h y d r o c l a d i a separated b y slightl y o b l i q u e nodes, a l l are thecate a n d beside the (large) h y d r o t h e c a also s u p p o r t three
nematothecae: one m e d i a n a n d t w o laterals. Interior of i n t e r n o d e w i t h a n u m b e r of
septa (rings): o n e at the b o t t o m of the m e d i a n nematotheca
a n d f i v e b e h i n d the
h y d r o t h e c a , of w h i c h the m o r e s u p e r i o r at the base of the lateral nematothecae. O c c a s i o n a l l y basal s e p t u m (ring) d o u b l e d , b r i n g i n g the total u p to seven. H y d r o t h e c a
large, a d c a u l i n e w a l l f u l l y adnate, w i t h i n t e r n a l s e p t u m at c o n v e x i t y i n l o w e r t h i r d
of a d c a u l i n e w a l l ; above a n d b e l o w attachment of s e p t u m h y d r o t h e c a l w a l l concave.
Intrathecal s e p t u m projecting as far as m i d d l e of hydrothecal cavity, c u r v i n g u p w a r d s .
A b c a u l i n e b o r d e r of h y d r o t h e c a convex i n l o w e r half, straight i n u p p e r half. H y d r o thecal r i m w i t h distinct abcauline, m e d i a n c u s p a n d a w e a k l y d e v e l o p e d , r o u n d e d
lateral cusp o n each side, sometimes v i s i b l e as a mere s i n u o s i t y (fig. 9b). M e d i a n nematotheca free f r o m h y d r o t h e c a l abcauline w a l l , short, w i t h circular r i m a n d large
f o r a m e n o p e n i n g into internode. Laterals t u b i f o r m , projecting above h y d r o t h e c a l
border, w i t h circular r i m a n d a d d i t i o n a l aperture i n a d c a u l i n e w a l l o p e n i n g into
h y d r o t h e c a l cavity.
O n l y a single p h y l a c t o c a r p is present; it is s l i g h t l y d a m a g e d a n d c o m p o s e d of
eight segments, each w i t h a p a i r of o p p o s i t e nematothecae, a l m o s t as the lateral nematothecae b u t w i t h bigger aperture. A single gonotheca occurs o n the basal segment,
i n s e r t i n g b e t w e e n the p a i r of nematothecae. G o n o t h e c a o v a l , c u r v e d a p i c a l l y a n d
there w i t h o v a l aperture o n the i n n e r side (fig. 10).
R e m a r k s . — Because of the structure of the p h y l a c t o c a r p this species s h o u l d b e
r e m o v e d to the genus Streptocaulus A l l m a n , 1883 ( R a m i l & V e r v o o r t , 1992b).
S u b s t r a t u m . — T h e c o l o n y s t u d i e d w a s detached.
D i s t r i b u t i o n . — A t l a n t i c coasts of M o r o c c o , Sierra L e o n e a n d Senegal ( B i l l a r d ,
1924, 1934; V e r v o o r t , 1959; P a t r i t i , 1970). It is considered to be a boreal species b y
Boero & B o u i l l o n (1993). R e c o r d e d depths v a r y b e t w e e n 70 a n d 380 m ( B i l l a r d , 1934).
Fig. 9. Streptocaulus dollfusi (Billard, 1924). a, part of main axis with hydrocladia; b, two internodes
with hydrothecae and nematothecae. Scales: a, 0.1 cm; b, 0.1 mm.
Fig. 10. Streptocalus dollfusi (Billard, 1924), phyllactocarp with single gonotheca. Scale: 0.2 mm.
28
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Genus Gymnangium Hincks, 1874
Gymnangium montagui ( B i l l a r d , 1912)
(fig- H )
Aglaophenia pennatula; Hincks, 1868: 292-294, fig. 33, pl. 63 fig. 3.
Halicornaria allmanii var. Marktanner-Turneretscher, 1890: 277-278, pl. 6 fig. 23.
Halicornaria pennatula; Ritchie, 1910: 225.
Halicornaria montagui Billard, 1912: 473, figs 6, 8.
Gymnangium montagui; Millard, 1975: 446, fig. 134F.
Material.— C4 (5.VÜ.1992), from fisherman, several colonies, no gonothecae.— C7: (23.xii.1984), 4 colonies, no gonothecae; (23.xii.1984), 6 colonies, no gonothecae; (9.v.1993), 8 m, numerous colonies with
gonothecae ( R M N H Coel. no. 27122, slide no. 2570).— C l l : (21.vii.1991), 20 m, 14 colonies, no gonothecae— C22: (14.V.1986), 25 m, 9 colonies, no gonothecae; (19.V.1986), 25 m, 10 colonies, no gonothecae.
Measurements (in mm)
Maximum height of colonies
Internodes of hydrocaulus, length
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrothecae, total length
120
0.517-0.684
0.245-0.535
0.324-0.412
0.175-0.324
0.166-0.289
idem, diameter at rim
0.175-0.385
length of free part abcauline wall
0.166-0.289
Gonotheca, length
idem, diameter
number of gonothecae in one row
0.425-0.75
0.4-0.5
19-35
D e s c r i p t i o n . — Erect, m o n o s i p h o n i c colonies r i s i n g f r o m a thick h y d r o r h i z a ;
c o l o u r l i g h t or d a r k y e l l o w , d e p e n d i n g u p o n age of c o l o n y (fig. 11a). E x c e p t f o r basal
part, h y d r o c a u l u s (axis) d i v i d e d into short internodes separated b y s l i g h t l y o b l i q u e
nodes. E a c h a x i a l internode w i t h t w o short apophyses, one latero-basal a n d o n e later o - d i s t a l , p o i n t i n g alternately r i g h t or left a n d s u p p o r t i n g alternate h y d r o c l a d i a 8-10
m m l o n g . There are t w o c u p s - s h a p e d nematothecae at the base of each a p o p h y s i s , a n
u p p e r a n d a l o w e r ; those of the basal a n d the d i s t a l a p o p h y s e s of consecutive internodes f o r m i n g t w o i n d i s t i n c t r o w s o n frontal aspect of axis (fig. l i b ) .
H y d r o c l a d i a b r o k e n u p into short internodes separated b y a l m o s t transverse
septa; each i n t e r n o d e w i t h one h y d r o t h e c a a n d three nematothecae. H y d r o t h e c a s p a cious, c u p - s h a p e d , w i t h t w o - t h i r d s of concave a d c a u l i n e w a l l adnate a n d a distinct,
s l i g h t l y c u r v e d abcauline internal s e p t u m at half l e n g t h abcauline w a l l , d i v i d i n g that
w a l l i n t o t w o c o n v e x p o r t i o n s . H y d r o t h e c a l aperture i n c l i n e d d o w n w a r d , m a k i n g a n
angle of 45° w i t h axis of internode; r i m s i n u o u s , occasionally w i t h i n d i c a t i o n of latera l c u s p o n each side (fig. 11c). P a i r of f l a n k i n g nematothecae also c u p - s h a p e d ; abcaul i n e w a l l c u r v e d , m u c h l o n g e r t h a n totally adnate a d c a u l i n e w a l l . M e d i a n n e m a t o theca m u c h elongated, c u r v e d , s u r p a s s i n g h y d r o t h e c a l b o r d e r considerably, free part
at least half l e n g t h of adnate part or m o r e , w i t h gutter-shaped o p e n i n g a l o n g w h o l e
free l e n g t h .
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
29
Fig. 11. Gymnangium montagui (Billard, 1912). a, colony; b, part of main axis with four hydrocladia; c,
three internodes with hydrothecae and nematothecae; d, gonotheca. Scales: a, 2 mm; b, 0.2 mm; c, d,
0.1 mm.
G o n o t h e c a e i n t w o r o w s o n f r o n t a l aspect of colony, each attached to h y d r o c l a d i a l a p o p h y s i s , c o l o u r w h i t e , i n contrast w i t h y e l l o w - b r o w n of rest of colony. Shape
c o n i c a l , w i t h truncate top a n d there w i t h c o n c a v i t y ; n a r r o w i n g p r o x i m a l l y at attachm e n t to i n t e r n o d e , o v a l o n cross section (fig. l i d ) .
S u b s t r a t u m . — O n r o c k y escarpments, also o n sponges, algae, B i v a l v i a a n d other
h y d r o i d s ( R a m i l , 1988; C o r n e l i u s , 1990; Isasi U r d a n g a r i n , 1986).
D i s t r i b u t i o n . — Restricted to the Eastern A t l a n t i c , w h e r e i t is f o u n d f r o m t h e
coasts of G r e a t B r i t a i n s o u t h w a r d t i l l S o u t h A f r i c a ( R a m i l , 1988; C o r n e l i u s & R y l a n d ,
1990; M i l l a r d , 1975). U n t i l 20 m d e p t h .
30
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Genus Lytocarpia Kirchenpauer, 1872
Lytocarpia myriophyllum (Linnaeus, 1758)
(fig-12)
Sertularia myriophyllum Linnaeus, 1758: 810.
Aglaophenia myriophyllum) Hincks, 1868: 290-292, pl. 64 fig. 2.
Lytocarpus myriophyllum; Marktanner-Turneretscher, 1890: 277, pl. 7 figs 10-11.
Thecocarpus myriophyllum; Billard, 1906: 226-227.
Lytocarpia myriophyllum; Stechow, 1923: 246.
Aglaophenia radicellata G.O. Sars, 1874: 9-10, pl. 2 figs 1-6.
Material— F57: (20.vii.l989), 76-80 m, 10 colonies without corbulae ( R M N H Coei. no. 27123, slide no.
2571).— F63: (20.vii.1989), 97-118 m, single colony without corbula.
Measurements (in mm)
Maximum height of hydrocaulus
150
Internodes of hydrocaulus, length
0.30-0.42
idem, diameter
Hydrocladial internodes, length
idem, diameter
Hydrothecae, total length
0.18-0.21
0.42-0.52
0.13-0.22
0.35-0.41
idem, diameter at rim
0.20-0.23
idem, length free part abcauline wall
0.21-0.32
D e s c r i p t i o n . — C o l o n y attached i n s u b s t r a t u m ( u s u a l l y s a n d bottom) b y m e a n s of
thick b u n d l e of h y d r o r h i z a l fibres l e a v i n g the h y d r o c a u l u s (axis) over a considerable
distance (fig. 12a). H y d r o c a u l u s p o l y s i p h o n i c , y e l l o w to b r o w n , w i t h u p to f o u r w e l l
m a r k e d , o b l i q u e nodes, d i s t r i b u t e d over n o n - h y d r o c l a d i a b e a r i n g l o w e r p a r t of
h y d r o c a u l u s (fig. 12b). A x i s c o m p o s e d of frontal tube a n d m a n y p a r a l l e l s e c o n d a r y
tubes, a l l w i t h a l o n g i t u d i n a l r o w of s m a l l c u p - s h a p e d nematothecae; o n the f r o n t a l
tube insert the h y d r o c l a d i a , almost w i t h o u t v i s i b l e a p o p h y s i s ; they are alternately
directed left or r i g h t a n d basally m o r e w i d e l y spaced that d i s t a l l y (fig. 12c). N o r m a l l y there i s one nematotheca b e t w e e n t w o successive a p o p h y s e s ; i n a d d i t i o n there i s
one besides a n d one b e h i n d the a p o p h y s i s ; those nematothecae w i t h d e e p l y s c o o p e d
adcauline wall.
H y d r o c l a d i a d i v i d e d into internodes b y means of transverse septa, each intern o d e w i t h a large h y d r o t h e c a a n d three nematothecae, one m e d i a n a n d t w o laterals
(fig. 12d). H y d r o t h e c a of characteristic shape b u t of v a r i e d d e p t h , w i t h a d c a u l i n e
w a l l a l m o s t c o m p l e t e l y adnate a n d r o u n d e d b o t t o m . F r o m the c u r v e d e n d of the
a d c a u l i n e w a l l a transverse s e p t u m p o i n t s into the h y d r o t h e c a l c a v i t y w i t h o u t closi n g i t completely. H y d r o t h e c a l b o r d e r almost p e r p e n d i c u l a r to i n t e r n o d a l axis o r
s l i g h t l y i n c l i n e d d o w n w a r d s ; r i m w i t h a n u m b e r of r o u n d e d cusps of v a r i e d d e v e l o p m e n t , a l m o s t d i s a p p e a r i n g laterally. There is a l w a y s a p r o m i n e n t m e d i a n c u s p o n
the a b c a u l i n e side (fig. 12e). M e d i a n nematotheca short, a l m o s t c o m p l e t e l y adnate
a n d w i t h short free part w i t h circular aperture. L a t e r a l nematothecae s l i g h t l y surp a s s i n g h y d r o t h e c a l r i m ; a d c a u l i n e w a l l d e e p l y s c o o p e d out. There is a n u m b e r of
septa (rings) i n the i n t e r n o d e of v a r i e d d e v e l o p m e n t ; i n o u r specimens there are six:
Fig. 12. Lytocarpia mynophyllum (Linnaeus, 1758). a, colony; b, oblique node in basal part of axis; c, part
of axis showing frontal tube with hydrocladia; d, three internodes with hydrothecae and nematothecae; e, single hydrocladial internode in lateral view. Scales: a, 1.5 cm; b, 3 mm; c, d, 0.5 mm; e, 0.2 mm.
32
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
one a b o v e base of l a t e r a l n e m a t o t h e c a e , t w o b e h i n d a d n a t e a d c a u l i n e h y d r o t h e c a l
w a l l , t w o at c u r v e d e n d of a b c a u l i n e h y d r o t h e c a l w a l l a n d o n e at base of m e d i a n
nematotheca.
A l l c o l o n i e s o b s e r v e d w e r e sterile; the species i s k n o w n to h a v e a large, o p e n corb u l a ; each of the costae w i t h a b a s a l h y d r o t h e c a .
S u b s t r a t u m . — O n s u b l i t t o r a l s a n d y b o t t o m s , o f t e n i n great n u m b e r s ( C o r n e l i u s
& R y l a n d , 1990).
D i s t r i b u t i o n . — A t l a n t i c species, d i s t r i b u t e d f r o m the A r c t i c s o u t h w a r d t i l l G u i n ea B i s s a u ; also i n the I n d o - P a c i f i c ( G i l i , V e r v o o r t & Pages, 1989; R a m i l & V e r v o o r t ,
1992a). B o e r o & B o u i l l o n (1993) refer to the species as b e i n g c o s m o p o l i t a n . T h e b a t h y m e t r i c a l r a n g e is f r o m 35 m ( G i l i , V e r v o o r t & Pages, 1989) d o w n to 1592 m ( R a m i l &
V e r v o o r t , 1992a).
F a m i l y H a l o p t e r i d i d a e M i l l a r d , 1962
G e n u s Antennella A l l m a n , 1877
Antennella siliquosa ( H i n c k s , 1877)
(fig. 13)
Plumularia siliquosa Hincks, 1877:148, pi. 12 figs 2-6.
? Antenella simplex Bedot, 1914: 84-86, pl. 5 figs 2-5.
? Antenella diaphana f. siliquosa; Broch, 1933: 26-27.
Antennella diaphana f. siliquosa; Vervoort, 1959: 286-289, fig. 43.
Halopteris diaphana diaphana p.p. Vervoort, 1968: 58-61, fig. 27c.
Halopteris diaphana f. siliquosa; García-Corrales, Aguirre Inchaurbe & Gonzalez Mora, 1978: 45-46, fig.
19; Ramil & Vervoort, 1992a: 148-149, fig. 38a.
Antenella siliquosa; Patriti, 1970: 58, fig. 82.
Antennella glutinosa; García-Corrales, Aguirre Inchaurbe & Gonzalez Mora, 1978: 40-42, fig. 17.
Material— C7: (10.ix.1990), 20-25 m, 60 colonies with gonothecae.— C l l : (31.vii.1991), 27 m, ten colonies without gonothecae.— C22: (13.V.1986), 35 m, numerous colonies with gonothecae ( R M N H Coel.
No. 27124, slide no. 2572); (14.V.1986), 25 m, various colonies with gonothecae.— C24: (6.V.1986), 38 m,
numerous colonies without gonothecae.— C26: (8.V.1986), 10 m, numerous colonies without gonothecae.— C28: (10.V.1986), 25 m, numerous colonies with gonothecae.
Measurements (in mm)
Maximum height of hydrocladia (stems)
45
Hydrocladial thecate internodes, length
0.29-0.43
Hydrocladial athecate internodes, length
0.44-0.70
idem, diameter
Hydrothecae, total length
idem, diameter at rim
0.08-0.14
0.19-0.34
0.16-0.34
Male gonothecae, length
0.575-0.696
idem, diameter
0.242-0.303
Female gonotheca, length
idem, diameter
1.515-1.66
0.363-0.66
D e s c r i p t i o n . — U n b r a n c h e d h y d r o c l a d i a r i s i n g f r o m creeping h y d r o r h i z a (fig.
13a); p e r i s a r c transparent; coenosarc w h i t i s h . H y d r o c l a d i a w i t h athecate b a s a l p a r t of
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
33
v a r i e d l e n g t h h a v i n g several transverse nodes a n d some frontal nematothecae. Rest
of h y d r o c l a d i u m w i t h o b l i q u e a n d transverse nodes separating athecate a n d thecate
internodes i n regular alternation (fig. 13b); first athecate internode w i t h basal transverse a n d t e r m i n a l o b l i q u e s e p t u m . A l l athecate internodes w i t h t w o nematothecae
i n one r o w . Those nematothecae scarcely m o v a b l e , short a n d f i r m l y attached, b i t h a l a m i c a n d w i t h a w i d e a n d deep e m b a y m e n t i n the a d c a u l i n e w a l l .
Thecate internodes w i t h c y l i n d r i c a l h y d r o t h e c a , one m e d i a n a n d t w o lateral
nematothecae; there is n o (reduced) nematotheca b e h i n d the a d c a u l i n e h y d r o t h e c a l
w a l l . W a l l s of h y d r o t h e c a p a r a l l e l , almost i m p e r c e p t i b l y everted, about half of
a d c a u l i n e w a l l free; r i m circular, s m o o t h (fig. 13b). M e d i a n (basal) nematotheca short
a n d strong, i m m o v a b l e , w i t h a d c a u l i n e w a l l d e e p l y s c o o p e d out. L a t e r a l nematothecae p l a c e d o n short a p o p h y s e s of s l i g h t l y v a r i e d l e n g t h (fig. 13c); they are short, d o
n o t reach the h y d r o t h e c a l r i m a n d are b i t h a l a m i c , w i t h s w o l l e n a p i c a l c h a m b e r a n d
d e e p l y s c o o p e d o u t a d c a u l i n e w a l l . O n the abcauline side there is a n i n f e r i o r e m a r g i nation.
F e m a l e a n d m a l e gonothecae f o u n d o n same h y d r o c l a d i u m . F e m a l e gonothecae
m o r e or less c y l i n d r i c a l , w i t h a short p e d i c e l i n s e r t i n g just u n d e r the h y d r o t h e c a at
the b a s a l internodes. There are u p to f o u r nematothecae o n the base; t o p w i t h w e l l
m a r k e d , c i r c u l a r o p e r c u l u m (fig. 13d). M a l e gonothecae shorter, p y r i f o r m , w i t h t w o
basal nematothecae a n d i n s e r t i n g o n the d i s t a l h y d r o c l a d i a l internodes; there is also
a n a p i c a l , c i r c u l a r o p e r c u l u m (fig. 13e).
S u b s t r a t u m . — F o u n d o n v a r i o u s types of h a r d substrata, o n B r y o z o a , C i r r i p e d i a ,
A s c i d i a , G o r g o n a r i a a n d algae. A l s o f o u n d o n w o r m tubes, o n sponges a n d o n other
h y d r o i d s (Vervoort, 1959; R a m i l & V e r v o o r t , 1992a).
D i s t r i b u t i o n . — M e d i t e r r a n e a n , Strait of Gibraltar, A t l a n t i c coast of M o r o c c o ,
I v o r y coast ( R a m i l & V e r v o o r t , 1992a). A c c o r d i n g to Boero & B o u i l l o n (1993) a M e d i t e r r a n e a n - A t l a n t i c species. D e p t h d i s t r i b u t i o n u n t i l 144 m d e p t h ( R a m i l & V e r v o o r t ,
1992a).
D i s c u s s i o n . — T h i s species has b e e n c o n s i d e r e d a n o n - p i n n a t e f o r m of Halopteris
diaphana (Heller, 1868): Halopteris diaphana f. siliquosa ( H i n c k s , 1877). A s Halopteris diaphana is also represented i n o u r c o l l e c t i o n w e h a v e been able to c o m p a r e the differences a n d o n account of this c o m p a r i s o n w e reached the c o n c l u s i o n that b o t h ' f o r m s '
are separable o n the specific l e v e l . I n the n o n - p i n n a t e f o r m , described above, the
nematothecae differ c o n s i d e r a b l y f r o m those of the p i n n a t e f o r m . T h e laterals, i n the
n o n - p i n n a t e f o r m , h a v e a g l o b u l a r a p i c a l chamber w i t h a deep e m b a y m e n t o n the
a d c a u l i n e side a n d a less deep one o n the abcauline side; they u s u a l l y d o n o t reach
the h y d r o t h e c a l border. I n the m e d i a n nematothecae the a d c a u l i n e w a l l of the basal
c h a m b e r is a l m o s t c o m p l e t e l y f u s e d w i t h the internode, r e n d e r i n g that nematotheca
f u l l y i m m o b i l e ; the a p i c a l chamber is d e e p l y s c o o p e d o n the a d c a u l i n e side. I n the
p i n n a t e f o r m , to be described presently as Halopteris diaphana, the laterals are c o n i c a l
a n d the e m b a y m e n t o n the a d c a u l i n e side, t h o u g h present, is s h a l l o w ; there is n o n e
o n the abcauline side; the m e d i a n nematotheca is m o v a b l e . T h e n o n - p i n n a t e f o r m
agrees w i t h the d e f i n i t i o n for Antennella A l l m a n , 1877 g i v e n b y M i l l a r d (1962, 1975),
the h y d r o c l a d i a r i s i n g directly f r o m the h y d r o r h i z a , t h o u g h a n occasional b r a n c h i n g
m a y occur; it is r e c o r d e d here as Antennella siliquosa ( H i n c k s , 1877). I n Halopteris A l l m a n , 1877, a p i n n a t e c o l o n y w i t h cauline hydrothecae is the n o r m a l , a d u l t c o n d i t i o n ,
Fig. 13. Antennella siliquosa (Hincks, 1877). a, bundle of colonies; b, thecate and athecate internodes; c,
lateral nematotheca on short apophysis; d, female gonotheca; e, maie gonotheca. Scales: a, 1 cm; b-e,
0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
35
t h o u g h d e v e l o p i n g colonies m a y have a transient Antennella stage.
A s i n d i c a t e d i n the d e s c r i p t i o n the l e n g t h of the a p o p h y s e s s u p p o r t i n g the lateral
nematothecae is v a r i a b l e . García-Corrales et a l . (1978) have r e c o r d e d specimens w i t h
quite short a p o p h y s e s as Halopteris glutinosa ( L a m o u r o u x , 1816). I n o u r o p i n i o n these
specimens, c o m p o s e d of h y d r o c l a d i a r i s i n g directly f r o m the stolon, b e l o n g i n Antennella siliquosa.
G a r c i a - C a r r a s c o s a (1981) separates A. siliquosa a n d H. diaphana b y the m o r p h o l o g y of t h e gonothecae a n d their n u m b e r of nematothecae. W e h a v e n o t o b s e r v e d
gonothecae i n o u r specimens of H. diaphana, b u t the n u m b e r of nematothecae o n the
gonotheca i n A. siliquosa m a y be m o r e than t w o , the n u m b e r m a i n t a i n e d f o r that species b y G a r c i a - C a r r a s c o s a .
M o r e o v e r , Garcia-Carrascosa
states that
the female
gonothecae i n A. siliquosa are g l o b u l a r a n d those of H. diaphana elongated. I n o u r A.
siliquosa the m a l e gonothecae are g l o b u l a r a n d the female elongated. G i v e n the
r e c o r d e d differences a n d the absence of gonothecae i n o u r m a t e r i a l of H. diaphana w e
r e f r a i n f r o m attaching diagnostic v a l u e to the shape of those gonothecae.
Antennella secundaria ( G m e l i n , 1791)
(fig. 14)
Sertularia secundaria Gmelin, 1791: 3856.
Aglaophenia secundaria; Lamouroux, 1824:19.
Plumularia secundaria; De Blainville, 1830:442; Carus, 1884:18; Pictet & Bedot, 1900: 27-28, pl. 6 fig. 7.
Anysocalyx secundarius; Costa, 1842:12.
Schizotricha secundaria; Bedot, 1921a: 23; Blackburn, 1942:108.
Antenella secundaria; Patriti, 1970: 57, fig. 81A-B.
Antennella secundaria; Millard, 1975: 332-334, fig. 10F-L; Ramil & Vervoort, 1992a: 143-145, fig. 37a-d.
Polyplumaria secundaria; Picard, 1955:189.
Plumularia dubiaformis Mulder & Trebilcock, 1911:119, pi. 2 fig. 7.
Material— C7: (08.viii.1993, 15 m, several colonies with gonothecae — C9: (25.ix.1991), 3 m, several
colonies with gonothecae.— C l l (03.vii.1991), 3 m, various colonies with gonothecae.— C22:
(13.V.1986), 35 m, 20 colonies with gonothecae (RMNH Coel. no. 27125, slide no. 2573).— C24:
(06.V.1986), 38 m, 15 colonies without gonothecae.— C26: (08.V.1986), 10 m, several colonies with
gonothecae.— C28: (10.v.1986), 25 m, several fragments without gonothecae.
Measurements (in mm)
Maximum height of hydrocladia
Thecate hydrocladial internodes, length
Athecate hydrocladial internodes, length
idem, diameter at node
Hydrothecae, length abcauline wall
idem, diameter at rim
Gonothecae, length
idem, maximum diameter
20
0.35-0.55
0.30-0.48
0.06-0.10
0.16-0.30
0.21-0.28
0.49-0.80
0.22-0.31
D e s c r i p t i o n . — C o l o n i e s c o m p o s e d of creeping s t o l o n ( h y d r o r h i z a ) f r o m w h i c h
rise the, n o r m a l l y u n b r a n c h e d , h y d r o c l a d i a w i t h r e d d i s h coenosarc (fig. 14a). Basal
p a r t of h y d r o c l a d i u m athecate, c o m p o s e d of a v a r i e d n u m b e r of internodes o n l y
b e a r i n g nematothecae a n d separated b y transverse nodes. Rest of h y d r o c l a d i a w i t h
Fig. 14. Antennella secundaria (Gmelin, 1791). a, bundle of colonies; b, thecate internode; c, part of axis
with female gonothecae; d, maie gonotheca. Scales: a, 1 cm; b, c, d, 0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
37
r e g u l a r l y alternating o b l i q u e a n d straight nodes, f o r m i n g a sequence of athecate a n d
thecate internodes, also alternately a r r a n g e d (fig. 14c). A t h e c a t e internodes b a s a l l y
w i t h straight, a p i c a l l y w i t h o b l i q u e n o d e a n d t w o nematothecae i n a f r o n t a l r o w ;
those nematothecae t w o c h a m b e r e d a n d m o v a b l e , a d c a u l i n e w a l l s l i g h t l y i n v a g i n a t e d . Thecate internodes b e a r i n g one h y d r o t h e c a a n d f o u r nematothecae: one o n basal
part of internode, a f l a n k i n g p a i r o n distinct a p o p h y s e s a n d a f o u r t h , s m a l l n e m a totheca
b e h i n d the a d c a u l i n e h y d r o t h e c a l w a l l . H y d r o t h e c a c y l i n d r i c a l , half of
a d c a u l i n e w a l l adnate, r i m circular, n o n everted, p e r p e n d i c u l a r to h y d r o t h e c a l l o n g
axis. Basal nematotheca c u r v e d ; a p i c a l chamber w i t h a d c a u l i n e e m b a y m e n t . D i s t a l
nematotheca of about same shape b u t smaller. L a t e r a l nematothecae w i t h n a r r o w e d
basal part, a p i c a l chamber c u p - s h a p e d , r i m m o r e o r less circular, w i t h s h a l l o w
a d c a u l i n e e m b a y m e n t (fig. 14b).
F e m a l e a n d m a l e gonothecae f o u n d o n separate h y d r o c l a d i a , t h o u g h s p r i n g i n g
f r o m same stolon. F e m a l e gonothecae elongated p y r i f o r m , w i t h c i r c u l a r l i d a n d t w o
o p p o s i t e nematothecae o n the n a r r o w e d basal p o r t i o n (fig. 14c). T h e m a l e gonothecae are c o n i c a l w i t h flattened t o p , also presenting a circular l i d , a n d a p a i r of o p p o site nematothecae at the base (fig. 14d). A l l nematothecae o n gonothecae are as the
laterals.
S u b s t r a t u m . — O n algae, sponges, tubes of P o l y c h a e t a a n d o n B r y o z o a .
D i s t r i b u t i o n . — C o s m o p o l i t a n species w i t h a preference f o r w a r m e r w a t e r s ( C o r n e l i u s & R y l a n d , 1990; R a m i l & V e r v o o r t , 1992a; Boero & B o u i l l o n , 1993); m a x i m u m
r e c o r d e d d e p t h 1250 m (Bedot, 1921b; A l v a r e z C l a u d i o , 1993).
Genus Haloptens A l l m a n , 1877
Halopteris diaphana (Heller, 1868)
(fig. 15)
Anisocalyx diaphanus Heller, 1868: 42, 82, pi. 2 fig. 5.
Plumularia diaphana; Bennitt, 1922: 254-255.
Schizotricha diaphana; Leloup, 1932:163-164.
Antennella diaphana f. typica Broch, 1933: 24.
Thecocaulus diaphanus; Vannucci-Mendes, 1946: 576, pi. 5 figs 46-47.
Antennella diaphana diaphana; Van Gemerden-Hoogeveen, 1965:49-54, figs 23-28.
Plumularia tenella Verrill, 1873: 731.
Schizotricha tenella; Nutting, 1900: 80, pi. 4 figs. 4-5.
Plumularia alternata Nutting, 1900: 62, pi. 4 figs 1-2.
Material.— C l : (15.X.1989), 0 m, four colonies without gonothecae — C7: (08.viii.1993), 15 m, several
colonies without gonothecae.— C l l : (29.x.1994), 6-9 m, five colonies without gonothecae.— C20:
(03.ii.1990), 3 m (RMNH Coel no. 27126, all in slide no. 2574), four colonies without gonothecae.
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length thecate internode
idem, athecate internodes
idem, diameter at node
Hydrocladial internode, length of thecate internode
10
0.22-0.78
0.27-0.55
0.06-0.10
0.24-0.49
38
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
idem, length athecate internode
0.17-0.29
idem, diameter at node
0.04-0.05
Hydrotheca, total length
0.12-0.15
idem, diameter at rim
0.12-0.15
idem, length abcauline wall
0.12-0.15
Description.—
Colony
with
erect,
monosiphonic,
unbranched,
transparent
h y d r o c a u l i , r i s i n g f r o m c r e e p i n g s t o l o n (fig. 15a). B a s a l p a r t of s t e m athecate, c o m p o s e d of v a r i e d n u m b e r of i n t e r n o d e s s e p a r a t e d b y transverse septa; there i s a f r o n tal r o w of nematothecae. D i s t a l p a r t of s t e m w i t h r e g u l a r s u c c e s s i o n of t r a n s v e r s e
Fig. 15. Haloptens diaphana (Heller, 1868). a, two colonies springing from creeping stolon; b, part of
axis with two hydrocladia; c, hydrocladial thecate internode in slightly oblique lateral view. Scales: a,
0.5 cm; b, c, 0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
39
a n d o b l i q u e septa, separating a regular sequence of athecate a n d thecate internodes.
A t h e c a t e internodes w i t h basally a straight a n d d i s t a l l y a n o b l i q u e n o d e ; one o r t w o
nematothecae are a r r a n g e d i n one r o w of the f r o n t a l surface (two is the n o r m a l c o n d i t i o n ) . T h e thecate internodes, w i t h b a s a l l y a n o b l i q u e a n d d i s t a l l y a straight sept u m , h a v e the h y d r o t h e c a at about half its l e n g t h , a n d three nematothecae, o n e
m e d i a n at the basal part of the internode a n d t w o f l a n k i n g nematothecae p l a c e d o n a
m i n o r e l e v a t i o n besides the h y d r o t h e c a . A l l nematothecae s i m i l a r , c o n i c a l , b i t h a l a m ic, w i t h distinct perisarcal r i n g l i m i t i n g the a p i c a l chamber. A d c a u l i n e w a l l of a l l
nematothecae w i t h a d c a u l i n e e m b a y m e n t ; the laterals just reach the h y d r o t h e c a l b o r der. There is n o (reduced) nematothecae b e h i n d the a d c a u l i n e h y d r o t h e c a l w a l l o r o n
the d i s t a l part of the thecate internode. H y d r o t h e c a e directed o b l i q u e l y f o r w a r d ,
a l m o s t c y l i n d r i c a l , w a l l s s l i g h t l y w i d e n i n g , w i t h half of a d c a u l i n e w a l l adnate, r i m
not everted, circular; p l a n e of aperture p e r p e n d i c u l a r to h y d r o t h e c a l l e n g t h axis.
H y d r o c l a d i a i n s e r t i n g o n apophyses of the thecate stem internodes, each a p o p h y s i s
just u n d e r the i n s e r t i o n of the lateral nematothecae of the a x i a l hydrothecae. T h e first
t w o h y d r o c l a d i a f o r m a p a i r ; the r e m a i n i n g are alternately a r r a n g e d a n d directed left
a n d r i g h t (fig. 15b). A r r a n g e m e n t of internodes, hydrothecae a n d nematothecae of
h y d r o c l a d i a as o n the stem (fig. 15c), b u t the athecate internodes u s u a l l y h a v e a s i n gle nematotheca.
N o gonothecae h a v e been observed.
S u b s t r a t u m . — Present material collected f r o m h a r d substratum (rocks). A l s o f o u n d
o n other h y d r o i d s (Halocordyle disticha) (Vervoort, 1959).
D i s t r i b u t i o n . — C i r c u m t r o p i c (Vervoort, 1959; Boero & B o u i l l o n , 1993).
Halopteris liechtensterni (Marktanner-Turneretscher, 1890)
(fig. 16)
Plumularia liechtensterni Marktanner-Turneretscher, 1890: 257-258, pi. 6 figs 2, 2a; Motz-Kossowska,
1908: LV, figs 1-3.
Polyplumaria liechtensterni; Picard, 1955:188; Rossi, 1961: 78.
Halopteris liechtensterni; Boero & Bouillon, 1993: 264.
Material— C7: (05.vii.1991), 3 m, several colonies, no gonothecae.— C l l : (30.vii.1991), 3 m, 12 colonies with gonothecae.— C18: (01.iii.1985), 20-25 m, several colonies, no gonothecae.— C22: (13.V.1986),
35 m, several colonies, no gonothecae.— C25: (05.V.1986), 20 m, several colonies, no gonothecae,
Measurements (in mm)
Maximum height of hydrocaulus
30
Internodes of hydrocaulus, length
0.48-0.64
idem, diameter
Hydrocladial internodes, length of thecate internodes
0.08-0.17
0.24-0.31
idem, length of athecate internodes
0.15-0.24
idem, diameter at nodes
0.06-0.08
Hydrotheca, total length
0.22-0.24
idem, diameter at rim
0.20-0.22
length of abcauline wall
0.17-0.20
Gonotheca, length
idem, maximum diameter
1.00-1.13
0.33-0.40
40
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Fig. 16. Halopteris liechtensterni (Marktanner-Turneretscher, 1890). a, two colonies; b, part of axis with
hydrocladia and gonothecae; c, flanking nematotheca; d, one of pair of reduced nematothecae behind
lateral nematothecae; e, infracalycine nematotheca; f, nematotheca on athecate hydrocladial internode;
g, stem internode with hydrotheca and nematothecae; h. three internodes of hydrocladium, two athecate and one thecate. Scales: a, 2.88 mm; b, 0,66 mm; c-f, 0.15 mm; g, h, 0.35 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
41
D e s c r i p t i o n . — C o l o n y c o m p o s e d of erect, m o n o s i p h o n i c , u n b r a n c h e d stems, risi n g f r o m a reticulate h y d r o r h i z a (fig. 16a). Basal part of s t e m athecate, c o m p o s e d of
u p to s e v e n internodes separated b y transverse septa, each of w h i c h m a y bear a p a i r
of nematothecae. Rest of s t e m c o m p o s e d of thecate internodes separated b y o b l i q u e
septa, each w i t h a h y d r o t h e c a a n d seven o r eight nematothecae
(fig. 16b). A x i a l
h y d r o t h e c a e c u p - s h a p e d , w i t h straight, s l i g h t l y d i v e r g i n g w a l l s ; r i m circular, n o t
everted (fig. 16g). N e m a t o t h e c a e b i t h a l a m i c , i n p r i n c i p a l c o n i c a l , b u t v a r i e d i n size
a n d shape a c c o r d i n g to p o s i t i o n . E a c h a x i a l i n t e r n o d e w i t h one m e d i a n i n f e r i o r n e matotheca, t w o p a i r s of laterals a n d t w o o r three nematothecae o n the d i s t a l p a r t of
each i n t e r n o d e . M e d i a n i n f e r i o r (fig. 16e) a n d m e d i a n d i s t a l nematothecae
of
r e d u c e d size, o n l y m o r e o r less m o v a b l e , w i t h deep e m b a y m e n t of the a d c a u l i n e
w a l l . D i s t a l nematothecae a r r a n g e d i n t w o f r o n t a l r o w s ; three is the n u m b e r u s u a l l y
f o u n d o n d i s t a l internodes. O f the f l a n k i n g nematothecae o n e p a i r is f o u n d inserted
o n a p o p h y s e s o n b o t h sides of the h y d r o t h e c a ; they are c o n i c a l , w i t h d i s t i n c t l y separated d i s t a l c h a m b e r w i t h deep a d c a u l i n e e m b a y m e n t a n d r e a c h i n g the h y d r o t h e c a l
r i m (fig. 16c, g). S e c o n d p a i r quite s m a l l , f o u n d i n the a x i l of a p o p h y s i s a n d intern o d e (fig. 16d). H y d r o c l a d i a i n s e r t i n g o n a p o p h y s e s besides a x i a l hydrothecae, altern a t e l y d i r e c t e d left a n d right; s o m e of the basal h y d r o c l a d i a m a y be b r a n c h e d . First
i n t e r n o d e of h y d r o c l a d i u m short, separated b y straight nodes, athecate. F o l l o w i n g
i n t e r n o d e athecate, w i t h straight basal a n d o b l i q u e d i s t a l n o d e a n d b e a r i n g o n e o r
t w o f r o n t a l nematothecae. Rest of h y d r o c l a d i u m c o m p o s e d of regular sequence of
thecate a n d athecate internodes (fig. 16h). Thecate internodes w i t h o b l i q u e basal a n d
transverse d i s t a l n o d e , one h y d r o t h e c a (as o n s t e m internodes), one m e d i a n i n f e r i o r
nematotheca a n d t w o p a i r s of f l a n k i n g nematothecae. A t h e c a t e internodes u s u a l l y
w i t h a s i n g l e f r o n t a l nematotheca (fig. 16f), t h o u g h t w o h a v e o c c a s i o n a l l y b e e n
o b s e r v e d o n basal internodes.
F e m a l e gonothecae large, c y l i n d r i c a l a n d a p i c a l l y truncate, w i t h c i r c u l a r l i d ;
basal p a r t n a r r o w e d , w i t h t w o o r r a r e l y three nematothecae, i n s e r t i n g b y means of a
short p e d i c e l besides a x i a l or h y d r o c l a d i a l hydrothecae (fig. 16b). M a l e gonothecae
smaller, p y r i f o r m , w i t h t w o nematothecae.
S u b s t r a t u m . — Sponges, algae a n d h a r d s u b s t r a t u m (rocks); also f o u n d o n b a l a n i d s a n d tubes of P o l y c h a e t a (Roca M a r t i n e z , 1986, as Halopteris sp.).
D i s t r i b u t i o n . — C o n s i d e r e d e n d e m i c f o r the M e d i t e r r a n e a n (Rossi, 1961; Boero &
B o u i l l o n , 1993); it h a s b e e n cited f r o m M e d i t e r r a n e a n coasts of M o r o c c o , France,
Italy, a n d f r o m the Baleares (Rossi, 1961; R o c a M a r t i n e z , 1986). R e c o r d e d depths
b e t w e e n 0 a n d 40 m (Rossi, 1961; R o c a M a r t i n e z , 1986).
Family Kirchenpaueriidae M i l l a r d , 1962
Genus Kirchenpaueria Jickeli, 1883
Kirchenpaueria pinnata (Linnaeus, 1758)
(figs 17,18)
Sertularia pinnata Linnaeus, 1758: 813.
Plumularia pinnata; Hincks, 1868: 295-296, pi. 65 fig. 1.
Kirchenpaueria pinnata; Bedot, 1916: 645; Roca & Moreno, 1987: 20, figs 12, 39; Ramil & Vervoort, 1992a:
158-161, fig. 41a-c.
42
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Plumularia echinulata Lamarck, 1816:126; Hincks, 1868: 302-303, pl. 65 fig. 2.
Plumularia similis Hincks, 1859:128; Hincks, 1868: 303-304, pl. 65 fig. 3.
Kirchenpaueria similis; Roca & Moreno, 1987a: 20, figs 13,40.
Plumularia elegantula G.O. Sars, 1874:103-104, pl. 3 figs 9-14.
Material.— C l : (25.v.1989), 0 m, numerous colonies with gonothecae; (02.vi.l989), 0 m, numerous colonies with gonothecae; (21.ii.1993), 0 m, numerous colonies with gonothecae ( R M N H Coel. no. 27127,
slide no. 2575).— C2 (03.iii.1991), 0 m, numerous colonies with gonothecae.— C3 (12.iv.1990), 0 m, ten
colonies without gonothecae.— C4 (08.Í.1989), 0 m, numerous colonies with gonothecae.— C5:
(ll.ii.1990), numerous colonies with gonothecae.— C7: (18.V.1990), 20 colonies with gonothecae.—
C i l : (29.X.1994), 6-9 m, 10 colonies without gonothecae.— C14: (15.vii.1993), 3-20 m, three colonies
with gonothecae.— C16: (23.xi.1991), 2-5 m, 15 colonies without gonothecae — C25: (5.V.1986), 20 m,
four colonies without gonothecae.— C29: (14.vii.1993), 10-18 m, ten colonies without gonothecae.
Measurements (in mm)
Maximum height hydrocaulus
Internodes of hydrocaulus, length
idem, diameter
Hydrocladial internodes, length of thecate internodes
idem, length of athecate internodes
idem, diameter at node
Hydrotheca, length abcauline wall
idem, diameter at rim
32
0.219-0.524
0.048-0.109
0.256-0.414
0.085-0.256
0.048-0.060
0.060-0.109
0.100-0.158
D e s c r i p t i o n . — C o l o n y c o m p o s e d of erect, m o n o s i p h o n i c , u n b r a n c h e d o r occas i o n a l l y b r a n c h e d h y d r o c a u l i r i s i n g f r o m a n i r r e g u l a r l y reticulate h y d r o r h i z a (fig.
17a). Basal p a r t of each axis n o t b e a r i n g h y d r o c l a d i a , c o m p o s e d of v a r i e d n u m b e r of
i n t e r n o d e s separated b y straight septa. Rest of axis m a d e u p of internodes w i t h a p i cal a p o p h y s i s , separated b y straight nodes, the a p o p h y s e s a n d h y d r o c l a d i a alternatel y d i r e c t e d left a n d right. A ' n a k e d ' n e m a t o p h o r e occurs o n the a d c a u l i n e s i d e of
each a p o p h y s i s a n d a s e c o n d is f o u n d o n the a x i a l internode just above the a p o p h y sis (fig. 17b). F i r s t i n t e r n o d e of h y d r o c l a d i u m short, athecate, w i t h straight b a s a l a n d
o b l i q u e d i s t a l n o d e ; occasionally t w o or three s u c h basal internodes are present. Rest
of h y d r o c l a d i u m m a d e u p of thecate internodes (fig. 18a, d ) , o r of a regular (fig. 18b)
or i r r e g u l a r (fig. 18c) sequence of thecate a n d athecate internodes, separated b y
straight septa. Thecate internodes w i t h o n e h y d r o t h e c a a n d t w o n e m a t o p h o r e s ;
hydrotheca fairly low, cup-shaped, walls gradually w i d e n i n g , occasionally slightly
c o n v e x ; a d c a u l i n e w a l l almost f u l l y adnate. N e m a t o p h o r e s i n one r o w o n front of
i n t e r n o d e , o n e c o m p l e t e l y ' n u d e ' n e m a t o p h o r e b e h i n d a d c a u l i n e w a l l of h y d r o t h e c a ,
the s e c o n d at the h y d r o t h e c a l base, w i t h m i n o r frontal p e r i d e r m a l s h i e l d (fig. 17b).
G o n o t h e c a e i n s e r t i n g o n h y d r o r h i z a o r i n a r o w o n the axis. G o n o t h e c a e b a r r e l s h a p e d , w i t h short p e d i c e l , h e x a g o n a l i n cross section a n d w i t h s i x l o n g i t u d i n a l c a r i nae, b e a r i n g d i g i t i f o r m p r o l o n g a t i o n s of w i d e l y v a r i e d d e v e l o p m e n t : l o n g i n s o m e
gonothecae (fig. 17c, c'), absent i n others (fig. 17d, d'). I n one c o l o n y the longest
a p p e n d a g e s are u s u a l l y f o u n d o n the female gonothecae.
V a r i a b i l i t y . — There is considerable v a r i a t i o n i n the b u i l d of the h y d r o c l a d i a :
s o m e h a v e a regular sequence of thecate internodes (fig. 18a, d ) , i n others there is a
r e g u l a r o r i r r e g u l a r sequence of thecate a n d athecate internodes (fig. 18b, c). There is
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
43
Fig. 17. Kirchenpaueria pinnata (Linnaeus, 1758). a, four colonies springing from creeping stolon; b, part of
axis with basal part of hydrocladium; c, c', female and male gonothecae with long spines; d, d', female
and male gonothecae with spines of reduced length. Scales: a, 2 cm; b, 0.1 mm; c, c', d, d', 0.5 mm.
Fig. 18. Kirchenpaueria pinnata (Linnaeus, 1758). a-d, hydrocladia from various colonies to show variation in structure. Scalea: a, 0.1 mm; b-d, 0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
45
also considerable difference i n l e n g t h of the athecate internodes: short i n t e r n o d e s fre­
q u e n t l y o c c u r r i n g i n colonies w i t h i r r e g u l a r alternation of thecate a n d athecate inter­
n o d e s , w h i l e l o n g (athecate) internodes p r e d o m i n a t e i n colonies w i t h a r e g u l a r
sequence (fig. 18b, c). T h e v a r i a t i o n i n d e v e l o p m e n t of the g o n o t h e c a l ' s p i n e s ' h a s
a l r e a d y b e e n i n d i c a t e d (fig. 17c, c ' a n d 17d, d').
S u b s t r a t u m . — O n algae, sponges, B r y o z o a , stones a n d r o c k y b o t t o m s .
Also
f o u n d o n other h y d r o i d s , o n polychaete tubes a n d o n shells ( H i n c k s , 1868; L e l o u p ,
1952; C h a s B r i n q u e z & R o d r i g u e z Babio, 1977; A l v a r e z C l a u d i o , 1993).
D i s t r i b u t i o n . — Kirchenp aueria p innata is m a i n l y a t r o p i c a l A t l a n t i c species (Boero
& B o u i l l o n , 1993) that is quite c o m m o n i n the M e d i t e r r a n e a n ( R a m i l & V e r v o o r t ,
1992a; R o c a M a r t i n e z , 1986; R o c a & M o r e n o , 1987b).
D i s c u s s i o n . — G i v e n the considerable v a r i a b i l i t y of the species as discussed above,
it is scarcely a s t o n i s h i n g that a n u m b e r of species is r e c o g n i z e d b y s o m e authors
(Bedot, 1916; D a C u n h a , 1944; Garcia­Carrascosa, 1981; Roca & M o r e n o , 1987; C o r n e l i u s
& R y l a n d , 1990), b a s i n g the d i s t i n g u i s h i n g characters o n presence or absence of athe­
cate internodes a n d their regular alternation w i t h hydrothecate internodes, the pres­
ence o r absence of the d i s t a l nematophore (on the thecate internode), the n u m b e r of
h y d r o c l a d i a s u p p o r t e d b y each a x i a l i n t e r n o d e a n d the d e v e l o p m e n t o f the g o n o ­
thecal armature [K p innata (Linnaeus, 1758), Κ echinulata (Lamarck, 1816), Κ similis
( H i n c k s , 1859), a n d Κ elegantula ( G . O . S ars, 1874)]. O f those species o n l y Κ pinnata
a n d Κ similis are accepted b y m o d e r n authors as C o r n e l i u s (1990, 1992) a n d R o c a &
M o r e n o (1987), t h o u g h not w i t h the same characteristics! O u r m a t e r i a l is character­
i z e d b y the constant presence of a single h y d r o c l a d i u m p e r a x i a l internode, a l l h a v e a
n e m a t o p h o r e b e h i n d the h y d r o t h e c a a n d o n the same c o l o n y b o t h h e a v i l y ' s p i n e d '
a n d ' n u d e ' gonothecae occur. A s far as the last character is concerned, w e h a v e
o b s e r v e d , as d i d M i l l a r d (1975), that s p i n u l a t i o n is dependent u p o n age a n d sex, the
m a l e gonothecae h a v i n g m o r e o r less d e v e l o p e d spines o r b e i n g totally ' n u d e ' . O u r
m a t e r i a l demonstrates v a r i a t i o n i n the structure of h y d r o c l a d i a :
a, short athecate internodes, often w i t h scarcely v i s i b l e nodes, that occur i r r e g u ­
l a r l y ; b . n o athecate internodes present; c, athecate internodes r e g u l a r l y a l t e r n a t i n g
w i t h thecate internodes a n d of a l m o s t the same l e n g t h .
C o l o n i e s of the last category u s u a l l y are finer a n d m o r e slender t h a n the others,
t h o u g h this does not appear f r o m the measurements. W i t h r e g a r d to the great v a r i ­
a b i l i t y i n the above n a m e d characters w e d o not believe those to d i s c r i m i n a t e at the
specific l e v e l .
Genus Ventromma Stechow, 1923
Ventromma halecioides ( A l d e r , 1859)
(fig. 19)
Plumularia halecioides Alder, 1859:353, pi. 12.
Kirchenpaueria halecioides; Bedot, 1923: 227,232; Billard, 1926: 98­99.
Ventromma halecioides; García­Corrales, Aguirre Inchaurbe & Gonzalez Mora, 1978:51­53, fig. 23.
Anisocalyx bifrons Heller, 1868:43, 82, pi. 2 fig. 6.
Plumularia inermis Nutting, 1900: 62­63, pi. 5 figs 1, 2, 2a; Fraser, 1912: 382, fig. 50; Bennitt, 1922: 255­
256, figs 3,4.
46
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Antennularia pinnata; Bennitt, 1922: 252-254, fig. 2 (not Antennularia pinnata Nutting, 1900: 71-72, pl. 10
figs 5, 6)
Plumularia irregularis Millard, 1958: 210-212, fig. 13A-C.
Kirchenpaueria irregularis; Millard & Bouillon, 1974: 9.
Material.— C l : (21.V.1989), 0 m, numerous colonies with gonothecae; (02.vi.1989), numerous colonies
with gonothecae; (18.viii.1989), numerous colonies with gonothecae (RMNH Coel. no. 27128, two
slides no. 2576).
Measurements (in mm)
Maximum height of hydrocaulus
39
Internodes of hydrocaulus, length
0.330-0.535
idem, diameter
0.070-0.184
Hydrocladial internodes, length thecate internodes
idem, diameter
0.359-0.622
0.035-0.087
Hydrothecae, total length
0.070-0.096
idem, diameter at rim
0.096-0.122
idem, length of abcauline wall
0.131-0.201
Gonothecae, length
idem, diameter
0.91-1.46
0.511-0.822
D e s c r i p t i o n . — Erect h y d r o c a u l i r i s i n g f r o m a c r e e p i n g s t o l o n , w e a k l y p o l y s i p h o n i c at the base, b r o w n a n d transparent i n the y o u n g e r parts, w i t h o c c a s i o n a l l y
s o m e r a m i f i c a t i o n s (fig. 19a). W i t h the e x c e p t i o n of the basal p a r t each h y d r o c a u l u s
is c o m p o s e d of internodes separated b y straight nodes, each i n t e r n o d e b e a r i n g one
lateral a p o p h y s i s , alternately directed r i g h t o r left, perisarc t h i n a n d transparent.
E a c h a p o p h y s i s has a ' n u d e ' n e m a t o p h o r e near the a x i l ; the i n t e r n o d e , above the
i n s e r t i o n of the a p o p h y s i s , has a laterally directed, b i t h a l a m i c b u t i m m o b i l e , c u p s h a p e d nematotheca (fig. 19b). H y d r o c l a d i a w i t h t h i n a n d h y a l i n e perisarc, alternatel y d i r e c t e d left a n d right, each b e g i n n i n g w i t h a short athecate i n t e r n o d e f o l l o w e d b y
a regular succession of thecate a n d (shorter) athecate internodes separated b y
straight nodes. Thecate internodes each w i t h c u p - s h a p e d h y d r o t h e c a a n d t w o n e m a tothecae; a l l nematothecae
of a h y d r o c l a d i u m f r o n t a l l y directed a n d i n o n e l i n e .
H y d r o t h e c a o n d i s t a l half of internode, abcauline w a l l s l i g h t l y c o n v e x ; a b o u t onet h i r d to o n e - f o u r t h of a d c a u l i n e w a l l free. O n e of nematothecae some distance b e l o w
base of h y d r o t h e c a , the second just above h y d r o t h e c a l r i m ; b o t h nematothecae s m a l l ,
c u p - s h a p e d , i n d i s t i n c t l y b i t h a l a m i c a n d i m m o b i l e (fig. 19b).
G o n o t h e c a e i n s e r t i n g o n front of axis, m o r e o r less c y l i n d r i c a l , w i t h t h i c k p e r i d e r m , truncate at the apex a n d there w i t h circular l i d ; base n a r r o w e d t o w a r d s short
p e d u n c l e . M a t u r e gonotheca w i t h c. 10 transverse, r o u n d e d ribs (fig. 19c).
S u b s t r a t u m . — O n sponges, B r y o z o a , o n rocks a n d o n fine s a n d . A l s o o n algae,
p h a n é r o g a m e s a n d o n other h y d r o i d s (Garcia-Carrascosa, 1981; G i l i , 1986; R a m i l ,
1988).
D i s t r i b u t i o n . — C o s m o p o l i t a n ( R a m i l , 1988; Boero & B o u i l l o n , 1993). A c c o r d i n g
to C o r n e l i u s & R y l a n d (1990) p r i m a r i l y restricted to boreal a n d t r o p i c a l waters.
D e p t h f r o m 0 to 20 m ( G i l i , 1986; C o r n e l i u s & R y l a n d , 1990).
Fig. 19. Ventromma halecioides (Aider, 1859). a, two colonies; b, monosiphonic part of axis with hydrocladium; c, mature gonotheca. Scales: a, 1 cm; b, 0.1 mm; c, 0.5 mm.
48
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Family Plumulariidae Hincks, 1868
Genus Nemertesia Lamouroux, 1812
Nemertesia ramosa ( L a m a r c k , 1816)
(figs 20,23a)
Antennularia ramosa Lamarck, 1816:123.
Nemertesia ramosa Lamouroux, 1816:164.
Cymodocea ramosa; Lamouroux, 1816: 212, pl. 7 figs la, Α.
Aglaophenia gracilis Lamouroux, 1816:171.
Nemertesia Janini Lamouroux, 1816:163, pi. 4 figs 3a, B, C.
Cymodocea comata Lamouroux, 1821:15, pi. 67, figs 12,13.
Sertularia seticomis Hogg, 1827: 33.
Antennularia Janini; De Blainville, 1830: 451.
Antennularia arborescens Hassal, 1843: 111.
Sertularia (Antennularia) ramosa; Dalyell, 1847: 202, pl. 39 figs 7­19, pl. 40.
Heteropyxsis ramosa; Kirchenpauer, 1876: 29,38, pl. 2 figs 22,22a, pl. 3 fig. 22.
Plumularia gracilis Kirchenpauer, 1876: 30.
Antennularia simplex Allman, 1877: 34, pi. 21 figs 1­2.
Antennularia ramulosa Merejkowsky, 1882: 280.
Antennularia profunda Queich, 1885a: 10, pi. 2 fig. 5.
Antennularia racemosa Fauvel, 1895: LXVII.
Antennularia variabilis Broch, 1903:10­11, pl. 4 figs 22­25.
Material.— C l l : (22.vii.1990), 5­28 m, two stems without gonothecae ( R M N H Coel. no. 27129, slide
no. 2577).— F13: (ll.vii.1989), 62 m, one stem without gonothecae.— F57: (20.vii.1989), 76­80 m, sever­
al fragments with gonothecae ( R M N H Coel. no. 27137, slide no. 2585).— F58: (20.vii.1989), 97­118 m,
several fragments with gonothecae.— A13: (15.X.1992), single stem without gonothecae.
Measurements (in mm)
Maximum height of hydrocaulus
200
Internodes of hydrocaulus, length
0.66­0.67
idem, diameter at node
0.15­0.28
Hydrocladial internodes, length of thecate internodes
idem, diameter at node
0.525­0.677
0.084­0.135
Hydrotheca, length of abcauline wall
0.084­0.135
idem, diameter at rim
0.118­0.152
Lateral nematothecae, length
0.075­0.100
Gonotheca, total length
0.625­0.700
idem, diameter
0.325­0.400
Description.— Colony with
formed b y many
thick,
strongly
intertwined hydrorhizal matting,
r a m i f i e d tubes, the p o l y s i p h o n i c y e l l o w i s h ­ b r o w n s t e m s r i s i n g
f r o m this h y d r o r h i z a l m a s s (fig. 23a). S tems t h i c k , b r a n c h e d , w i t h t h i c k p e r i s a r c , o n e
of c o m p o s i n g tubes of greater d i a m e t e r a n d this tubes i n h i g h e r , w e a k l y p o l y s i p h o n ­
ic a n d m o n o s i p h o n i c p a r t s of c o l o n y g i v e s rise to (verticils of) a p o p h y s e s
bearing
h y d r o c l a d i a ; this c e n t r a l t u b e h a s little m a r k e d transverse n o d e s . T h e s e n o d e s h a v e
one o r m o r e f r e q u e n t l y t w o v e r t i c i l s of a p o p h y s e s ; coenosarc c o m p o s e d of v a r i o u s
tubes (canaliculated).
A p o p h y s e s i n y o u n g e s t p a r t s i n o p p o s i t e p a i r s (fig. 20a), i n
o l d e r p a r t s i n v e r t i c i l s of three o r four, a r r a n g e m e n t s u c h that a p o p h y s e s of s u c c e e d ­
Fig. 20. Nemertesia ramosa (Lamarck, 1816). a, upper (young) part of stem with opposite apophyses; b,
apophysis, 'mamelon' invisible; c, hydrocladial internode; d, female gonotheca; e, male gonotheca.
Scales: a, 1 mm; b, c, 0.1 mm; d, e, 0.5 mm.
50
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
i n g p a i r s alternate, r e s u l t i n g i n t w i c e the n u m b e r of l o n g i t u d i n a l r o w s of h y d r o c l a d i a . A x i a l tubes i r r e g u l a r l y set w i t h nematothecae. A p o p h y s e s ( f i g . 20b) w i t h t h i c k
p e r i d e r m , c o n t r a r y to the t h i n p e r i d e r m of the h y d r o c l a d i a , a n d h a v e a ' m a m e l o n '
a n d a v a r i e d n u m b e r of nematothecae, seven b e i n g the m a x i m u m . H y d r o c l a d i a
transparent o r y e l l o w i s h , c o m p o s e d e x c l u s i v e l y of thecate internodes, separated b y
a l m o s t transverse nodes. E a c h i n t e r n o d e w i t h one h y d r o t h e c a a n d f o u r nematothecae: one basal, t w o laterals a n d one o n the d i s t a l part of the i n t e r n o d e ; a l l u n p a i r e d
nematothecae i n o n e r o w . A l l nematothecae c o n i c a l , b i t h a l a m i c a n d w i t h w e l l
m a r k e d a d c a u l i n e e m b a y m e n t . H y d r o t h e c a f a i r l y s m a l l , c u p - s h a p e d w i t h straight
w a l l s , r i m circular, a l m o s t p e r p e n d i c u l a r to l e n g t h axis of i n t e r n o d e (fig. 20c).
F e m a l e a n d m a l e gonothecae o n l y s l i g h t l y different, o v o i d w i t h o b l i q u e l y t r u n cate t o p , n a r r o w i n g t o w a r d s base a n d w i t h short p e d i c e l . M a l e gonotheca w i t h large
c i r c u l a r aperture, p l a n e s l i g h t l y o b l i q u e (fig. 20e). F e m a l e gonotheca larger, aperture
m o r e or less o v a l , m o r e s t r o n g l y i n c l i n e d t h a n i n m a l e gonotheca (fig. 20d). B o t h
sexes insert o n the a p o p h y s e s .
S u b s t r a t u m . — T h e m a t e r i a l inspected is c o m p o s e d of detached colonies. T h e
species has been f o u n d attached to shells of b i v a l v e s a n d o n m u d d y a n d s a n d y bottoms (Garcia-Carrascosa, 1981).
D i s t r i b u t i o n . — C o s m o p o l i t a n species ( R a m i l , 1988; A l v a r e z C l a u d i o , 1993), o b s e r v e d b e t w e e n 5 a n d 1425 m d e p t h ( G i l i , 1986; Bedot, 1921b; A l v a r e z C l a u d i o , 1993).
Nemertesia antennina (Linnaeus, 1758)
(figs 21,23b)
Sertularia antennina Linnaeus, 1758: 811.
Antennularia antennina; Hincks, 1868: 280-281, pi. 61.
Nemertesia antennina; Bedot, 1917b: 42; Ramil & Vervoort, 1992a: 163-169, figs 42a-r, 43a-h.
Antennularia janini; Marktanner-Turneretscher, 1890: 259, pi. 6 figs 9, 9a [not Nemertesia janini Lamouroux, 1816 = Nemertesia ramosa (Lamarck, 1816)].
Antennularia octoseriata Jäderholm, 1896:15-16, pl. 2 fig. 6.
Antennularia Perrieri var. antennoides Billard, 1904: 217-219.
Nemertesia irregularis p.p. Stechow, 1913: 93-94.
Nemertesia perrieri; Vervoort, 1966:138-139, fig. 40.
Material— C l l : (03.vii.1991), 5-28 m, ten colonies without gonothecae.— C17: (26.viii.1981), 25 m,
numerous colonies without gonothecae (RMNH Coel. no. 27130, slide no. 2578).— F l : (09.vii.1989),
190 m, 3 stems with gonothecae.—F2: (09.vii.1989), 116-118 m, two stems without gonothecae.
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length
idem, diameter
295
0.45-1.50
0.16-0.56
Hydrocladial internodes, length thecate internodes
idem, length athecate internodes
idem, diameter
Hydrotheca, total length
idem, diameter at rim
Lateral nematothecae, length
Gonothecae, total length
idem, maximum diameter
0.29-0.49
0.15-0.29
0.05-0.06
0.06-0.08
0.11-0.13
0.15-0.29
0.52-0.70
0.28-0.30
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
51
Fig. 21. Nemertesia antennina (Linnaeus, 1758). a, insertion of hydrocladium on apophysis; b, part of
hydrocladium with lengthened nematothecae; c, normally developed thecate internode with hydrothecal rim ventrally inclined; d, thecate internode with lengthened nematothecae; e, female gonotheca
inserting on apophysis. Scales: a, b, 0.5 mm; c, d, 0.1 mm; e, 0.3 mm.
52
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Description.— Colonies w i t h
well
d e v e l o p e d mass
of i n t e r t w i n i n g , m u c h
b r a n c h e d , w h i t i s h h y d r o r h i z a l fibres, s p r i n g i n g f r o m basal p o r t i o n of stems a n d
f i r m l y a n c h o r i n g c o l o n y i n s e d i m e n t o r attaching i t to s u b s t r a t u m (fig. 23b). A x i s
f i r m a n d erect, m o n o s i p h o n i c , n o t b r a n c h e d , w i t h thick, y e l l o w i s h to b r o w n perisarc
a n d d i v i d e d into little m a r k e d internodes b y means of i n d i s t i n c t , transverse n o d e s ;
each i n t e r n o d e w i t h one o r t w o verticils of a p o p h y s e s ; i r r e g u l a r l y d i s t r i b u t e d n e m a tothecae m a y occur. Coenosarc of axis c o m p o s e d of several tubes (canaliculated).
A p o p h y s e s u s u a l l y i n verticils of four, those of s u c c e e d i n g verticils alternating, each
w i t h ' m a m e l o n ' o n s u p e r i o r surface a n d u p to seven nematothecae; perisarc f i r m .
H y d r o c l a d i a w i t h t h i n perisarc, c o m p o s e d of a regular succession of athecate a n d
thecate internodes, the latter w i t h a basal o b l i q u e a n d a d i s t a l a l m o s t transverse
n o d e . Basal part of h y d r o c l a d i u m w i t h one o r t w o short, athecate internodes each
w i t h a nematotheca (fig. 21a); those first articles m a y also b e absent i n w h i c h case the
first i n t e r n o d e of the h y d r o c l a d i u m is thecate. Athecate internodes u s u a l l y w i t h a
frontal nematotheca, occasionally w i t h t w o . Thecate internodes w i t h a h y d r o t h e c a
a n d three nematothecae, o n e m e d i a n p r o x i m a l a n d t w o laterals, i n s e r t i n g o n the
i n t e r n o d e just u n d e r the h y d r o t h e c a l r i m (fig. 21c, d ) . A l l nematothecae c o n i c a l ,
b i t h a l a m i c a n d w i t h d i s t i n c t l y separated a p i c a l chamber; a d c a u l i n e w a l l w i t h d i s t i n c t
e m b a y m e n t . L a t e r a l nematothecae i n s o m e colonies greatly elongated (fig. 21b, d ) .
Hydrotheca comparatively shallow, cup-shaped, w i t h diverging walls; r i m circular
t h o u g h s i n u o u s i n s o m e colonies, p r o x i m a l l y i n c l i n e d c o m p a r e d to i n t e r n o d a l l e n g t h
axis (fig. 21c).
B o t h female a n d m a l e gonothecae o b s e r v e d , f o u n d attached to the u p p e r surface
of the a p o p h y s e s . T h e female gonothecae are o v o i d , w i t h a c o n v e x a n d a flattened
w a l l ; the aperture is o v a l a n d p l a c e d o n the u p p e r p a r t of the flattened s i d e ; i t is
c l o s e d b y a n o p e r c u l u m (fig. 21e).
V a r i a b i l i t y . — Some colonies h a v e l o n g a p o p h y s e s a n d l e n g t h e n e d lateral n e m a tothecae o n the thecate internodes (fig. 21b, d).
S u b s t r a t u m . — O n v a r i o u s types of b o t t o m : s a n d , m u d , g r a v e l a n d shells. A l s o
f o u n d o n corals (Roca M a r t i n e z , 1986; A l v a r e z C l a u d i o , 1993).
D i s t r i b u t i o n . — C o s m o p o l i t a n species (Boero & B o u i l l o n , 1993; A l v a r e z C l a u d i o ,
1993).
Nemertesia irregularis ( Q u e i c h , 1885)
(figs 22,23c)
Antennularia irregularis Queich, 1885: 8-9, pl. 2 fig. 4 [Not Antennularia irregularis Fraser, 1938 = Nemertesia fraseri Ramil & Vervoort, 1992a].
Nemertesia irregularis; Ramil & Vervoort, 1992a: 170-173, fig. 48a.
Nemertesia irregularis p.p. Stechow, 1913: 93-94.
Antennularia Perrieri Billard, 1901: 73-74.
Antennularia dendritica Stechow, 1907:195-196.
Nemertesia minor p.p. Stechow, 1923:18.
Material.— C13: (05.vii.1991), 5 m, 16 colonies with gonothecae.— F4: (10.iv.1989), 11 colonies with
gonothecae.— F5: (10.vii.1989), 67-68 m, seven colonies without gonothecae.— F8: (10.vii.1989), 238291 m, ten colonies without gonothecae ( R M N H Coel. no. 27131, slide no. 2579).— F38: (16.vii.1989),
60-62 m, seven colonies with gonothecae.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
53
Measurements (in mm)
Maximum height of hydrocaulus
Internodes of hydrocaulus, length
idem, diameter
250
0.57-1.37
0.41-0.48
0.338-0.491
Hydrocladial internodes, length of thecate internodes
idem, length athecate internodes
idem, diameter at node
0.338-0.474
Hydrotheca, total length
0.084-0.118
idem, diameter at rim
Lateral nematotheca, length
Female gonotheca, total length
idem, maximum diameter
0.050-0.084
0.118-0.152
0.118-0.152
0.92
0.43
D e s c r i p t i o n . — M a t t i n g of h y d r o r h i z a fibres y e l l o w i s h , s p r i n g i n g f r o m basal p o r t i o n of axis a n d r o o t i n g c o l o n y i n s e d i m e n t or attaching it to s u b s t r a t u m . A x i s m o n o s i p h o n i c , y e l l o w ; perisarc thick; coenosarc w i t h m a n y p a r a l l e l tubes (canaliculated).
D i v i s i o n i n t o internodes indistinct, nodes transverse; each i n t e r n o d e w i t h one or t w o
verticils of a p o p h y s e s , n u m b e r of a p o p h y s e s p e r verticils t w o (youngest parts) to
f i v e (older parts), alternating w i t h those of s u c c e e d i n g v e r t i c i l ; each a p o p h y s i s w i t h a
' m a m e i o n ' a n d u p to six nematothecae, a r r a n g e d i n pairs. H y d r o c l a d i a w i t h thecate
a n d athecate internodes; nodes m o r e or less o b l i q u e . F i r s t i n t e r n o d e of h y d r o c l a d i u m
short a n d athecate, w i t h a single nematotheca; this i n t e r n o d e absent i n s o m e c o l o nies, w h e r e the first i n t e r n o d e i s a n o r m a l thecate i n t e r n o d e w i t h h y d r o t h e c a a n d
three nematothecae, one m e d i a n o n p r o x i m a l p a r t a n d a p a i r of f l a n k i n g nematothecae. I n the rest of the h y d r o c l a d i u m there m a y be a regular sequence of thecate internodes, as d e s c r i b e d above, a n d athecate internodes b e a r i n g t w o f r o n t a l nematothecae, these m a y be separated b y a n o d e (fig. 22a). T h e thecate i n t e r n o d e m a y h a v e a n
i n c o m p l e t e s e p t u m (perisarcal ring) u n d e r the i n s e r t i o n of the m e d i a n nematotheca.
H y d r o t h e c a s h a l l o w , c u p - s h a p e d , w i t h d i v e r g i n g w a l l s , the a d c a u l i n e w a l l b e i n g
a l m o s t c o m p l e t e l y adnate; r i m s i n u o u s to circular. A l l nematothecae c o n i c a l , b i t h a l a m i c , w i t h distinct a p i c a l chamber a n d f a i r l y deep a d c a u l i n e e m b a y m e n t (fig. 22b).
O n l y female gonothecae have been o b s e r v e d ; these are o v o i d , w i t h s u p e r i o - latera l , o v a l aperture, n a r r o w i n g p r o x i m a l l y (fig. 22c). T h e y insert o n the a p o p h y s e s .
S u b s t r a t u m . — O n bottoms of m i x e d s a n d a n d m u d a n d o n concretions of p o l y chaete tubes.
D i s t r i b u t i o n . — W i d e l y d i s t r i b u t e d i n the eastern A t l a n t i c , r a n g i n g f r o m the B a y
of Biscay to Senegal ( R a m i l & V e r v o o r t , 1992a); also f o u n d i n Japanese waters (Stechow, 1913).
D i s c u s s i o n . — A c c o r d i n g to R a m i l & V e r v o o r t (1992a) N. irregularis is less v a r i a b l e
t h a n N. antennina a n d differs b y the presence of t w o nematothecae o n the athecate
internodes ( t h o u g h this internode m a y be d i v i d e d ) , b y the absence of nematothecae
o n the axis, a n d b y the occurrence of a single basal i n t e r n o d e p e r h y d r o c l a d i u m ,
b e a r i n g a single nematotheca. The occasional presence of m o r e s u c h internodes is
e x p l a i n e d b y regeneration after d a m a g e .
I n the present m a t e r i a l the lack of v a r i a t i o n is w e l l d e m o n s t r a t e d , b u t b o t h N .
antennina a n d N. irregularis, m a y h a v e h y d r o c l a d i a w i t h t w o basal athecate internodes o r h a v e none. A x i a l nematothecae are absent i n o u r m a t e r i a l of b o t h species,
Fig. 22. Nemertesia irregularis (Queich, 1885). a, apophysis with hydrocladium; b, athecate and thecate
internodes from hydrocladium; c, female gonothecae. Scales: a, c, 0.5 mm; b, 0.1 mm.
Fig. 23. a, Nemertesia ramosa (Lamarck, 1816), colony; b, Nemertesia antennina (Linnaeus, 1758), two
stems; c, Nemertesia irregularis (Queich, 1885), single stem. Scales: a, 1 cm; b, c, 2 cm.
56
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
w h i l e o c c a s i o n a l l y there are t w o nematothecae o n the athecate i n t e r n o d e s
of N .
antennina. H o w e v e r , the u s u a l c o n d i t i o n is o n e n e m a t o t h e c a o n the athecate h y d r o c l a d i a l i n t e r n o d e s i n A. antennina against t w o i n N. irregularis. F o r the m o m e n t b o t h
species are k e p t separate.
Genus Plumulana
Lamarck, 1816
Plumularia setacea ( L i n n a e u s , 1758)
(fig. 24)
Sertularia setacea Linnaeus, 1758: 813.
Aglaophenia setacea; Lamouroux, 1816:171.
Plumularia setacea; Johnston, 1847: 97, pi. 22 figs 3-5; Hincks, 1868: 296- 299, pi. 66 fig. 1; Ramil & Vervoort, 1992a: 191-193, fig. 47f-i.
Aglaophenia Gaimardi Lamouroux, 1824: 611, pi. 95 figs 9,10.
Plumularia tripartita Von Lendenfeld, 1885: 477,626, pl. 12 figs 8-10.
Plumularia multinoda Allman, 1886:157, pi. 27 figs 4-6.
Plumularia lagenifera Allman, 1886:157-158, pi. 26 figs 1-3.
Plumularia túrgida Bale, 1888: 779, 786, pi. 20 figs 12-13.
Plumularia corrugata Nutting, 1900: 64, pi. 6 figs 1-3.
Plumularia palmeri Nutting, 1900: 65, pi. 6 figs 4-5.
Plumularia milleri Nutting, 1905: 934, 951, pi. 5 fig. 1, pi. 12 figs 6-7.
Material— C2 (03.iii.1991), 0 m, 60 colonies with gonothecae; (18.viii.1989), 0 m, numerous colonies
with gonothecae ( R M N H Coel. no. 27132, slide no. 2580.— C5: (11.02.1990), 0 m, abundant colonies
with gonothecae— C7: (07.vi.1992), 5 m, 18 colonies with gonothecae; (08.viii.1993), 15 m, 25 colonies,
some with gonothecae. C26: (08.V.1986), 10 m, 15 colonies with gonothecae.— C15: (26.vii.1991), 3 m,
seven colonies with gonothecae— C20: (03.ii.1991), 0 m, 15 colonies with gonothecae.— C29:
(14.vii.1993), 10-18 m, 11 colonies without gonothecae.
Measurements (in mm)
Maximum height of hydrocaulus
10
Internodes of hydrocaulus, length
0.22-0.48
idem, diameter
Hydrocladial internodes, length athecate internodes
idem, length thecate internodes
diameter at node
Hydrotheca, total length
idem, diameter at rim
0.08-0.16
0.110-0.260
0.240-0.370
0.05-0.07
0.08-0.10
0.10-0.13
Lateral nematothecae, length
0.05-0.06
Gonothecae, total length
0.65-0.85
idem, maximum diameter
0.12-0.35
D e s c r i p t i o n . — Erect, m o n o s i p h o n i c a n d u n b r a n c h e d h y d r o c a u l i r i s i n g f r o m a
c r e e p i n g s t o l o n ( f i g . 24a); c o l o u r b r o w n i n oldest, t r a n s p a r e n t
i n younger
H y d r o c a u l u s d i v i d e d i n t o i n t e r n o d e s separated b y straight n o d e s ; e a c h
parts.
internode,
e x c e p t those at base of s t e m , w i t h a l a t e r o - d i s t a l a p o p h y s i s a n d t w o n e m a t o t h e c a e ,
o n e at the b a s a l p a r t a b o v e the p r e c e d i n g a p o p h y s i s , the s e c o n d a l m o s t i n the a x i l of
the a p o p h y s i s (fig. 24b). H y d r o c l a d i a i n s e r t e d o n a p o p h y s e s , a l t e r n a t e l y d i r e c t e d left
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
57
Fig. 24. Plumularia setacea (Linnaeus, 1758). a, two colonies rising from creeping stolon; b, part of axis
with apophyses, hydrocladium and gonotheca (with collar-shaped apical part); c, thecate and athecate
internodes, d, gonotheca. Scales: a, 1 cm; b-d, 0.1 mm.
a n d r i g h t , c o m p o s e d of regular succession of thecate a n d , shorter, athecate intern o d e s ; the first h y d r o c l a d i a l internode is short a n d athecate, w i t h imperfect s e p t u m
or r i n g (fig. 24b). Thecate internodes w i t h c u p - s h a p e d h y d r o t h e c a a n d three n e m a tothecae: o n e o n basal part of i n t e r n o d e a n d t w o f l a n k i n g the h y d r o t h e c a . There m a y
be a n i n c o m p l e t e s e p t u m o r r i n g u n d e r the basal nematotheca. A d c a u l i n e w a l l of
h y d r o t h e c a a l m o s t c o m p l e t e l y adnate, r i m circular a n d s l i g h t l y tilted d o w n w a r d s .
A t h e c a t e internodes w i t h a single frontal nematotheca a n d t w o rings o r septa: o n e
basal, o n e d i s t a l (fig. 24b, c). A l l nematothecae conical, w i t h distinct a p i c a l c h a m b e r
a n d adcauline embayment.
G o n o t h e c a e elongated o v o i d , n a r r o w i n g b a s a l l y t o w a r d s short p e d u n c l e a n d
inserted o n a p o p h y s e s ; d i s t a l l y w i t h s m a l l , circular aperture (fig. 24b, d ) . S o m e
gonothecae n a r r o w e d a p i c a l l y to f o r m a collar (fig. 24b). N o differences b e t w e e n
m a l e a n d female gonothecae have b e e n observed.
S u b s t r a t u m . — O c c a s i o n a l l y f o u n d o n h a r d , r o c k y surfaces, b u t quite c o m m o n as
58
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
e p i z o i t e of other h y d r o i d s (Aglaophenia tubiformis, A. octodonta). A l s o f o u n d o n algae,
mussels, A s c i d i a , b a l a n i d s , B r y o z o a a n d tubes of P o l y c h a e t a (Roca M a r t i n e z , 1986;
R a m i l & V e r v o o r t , 1992a; A l v a r e z C l a u d i o , 1993).
D i s t r i b u t i o n . — C o s m o p o l i t a n . F o u n d o n b o t h sides of the A t l a n t i c , a l o n g the
w h o l e extent of the coasts. A l s o r e c o r d e d f r o m the M e d i t e r r a n e a n , the Pacific, I n d i a n
a n d A r c t i c Oceans ( M i l l a r d , 1975; C o r n e l i u s & R y l a n d , 1990; Boero & B o u i l l o n , 1993).
A c c o r d i n g to C o r n e l i u s & R y l a n d (1990) preferentially b e t w e e n 0 a n d 20 m , b u t also
f o u n d at 740 m d e p t h (Broch, 1918).
Genus Monotheca Nutting, 1900
Monotheca pulchella (Bale, 1882)
(fig. 25)
Plumularia pulchella Bale, 1882: 42-43, pi. 15 fig. 6; Totton, 1930: 221-222, fig. 58a-d; Blanco, 1973: 73-76,
figs 1-3; Millard, 1975: 398-399, fig. 125C-D.
Monotheca pulchella; Stechow, 1923: 224.
PlumulariaflexuosaBale, 1894:115-116, pi. 5 figs 6-10.
Plumularia (Monotheca)flexuosa;Stechow, 1925: 499.
Plumularia oblica; Patriti, 1970: 58-59, fig. 83.
Plumularia femina García-Corrales, Aguirre Inchaurbe & Gonzalez Mora, 57-61, fig. 26.
Material — C2: (18.viii.1989), 0 m, numerous colonies with gonothecae (RMNH Coel. no. 27133, slide no.
2581); (10.ii.1990), several colonies; (24.vii.1993), 0 m, 15 colonies with gonothecae.— C5: (23.xii.1991), 0
m, several colonies; (ll.ii.1990), several colonies.
Measurements (in mm)
Maximum height of hydrocaulus
7
Internode of hydrocaulus, length
idem, diameter
Hydrocladial internodes, length thecate internode
idem, length of athecate internode
diameter at node
Hydrotheca, total length
0.21-0.27
0.05-0.06
0.19-0.20
0.08-0.09
0.03-0.04
0.12-0.15
idem, diameter at rim
idem, length abcauline wall
Gonotheca, total length
idem, diameter
0.12-0.13
0.13-0.14
0.21-0.27
0.05-0.06
D e s c r i p t i o n . — C o l o n i e s c o m p o s e d of s t r o n g a n d reticulate h y d r o r h i z a , f r o m
w h i c h rise erect, m o n o s i p h o n i c , occasionally s l i g h t l y b r a n c h e d h y d r o c a u l i (fig. 25a).
P e r i d e r m of h y d r o c a u l i a n d h y d r o c l a d i a t h i n , b r o w n i s h to transparent (in the y o u n g est parts); h y d r o c a u l i d i v i d e d i n t o internodes b y means of transverse septa, each
w i t h l a t e r o - t e r m i n a l a p o p h y s i s (fig. 25c). I n a d d i t i o n each i n t e r n o d e w i t h three n e matothecae: one i n the l o w e r half a n d t w o f l a n k i n g the a p o p h y s i s (fig. 25b). Those
nematothecae h a v e the shape of a s l i g h t l y c u r v e d f u n n e l ; they are t w o - c h a m b e r e d ,
m o b i l e ; d i s t a l m a r g i n p r o v i d e d w i t h distinct e m b a y m e n t . H y d r o c l a d i a i n s e r t i n g o n
the a p o p h y s e s a n d alternately d i r e c t e d left a n d right; they l i e m o r e o r less i n the
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
59
Fig. 25. Monotheca pulchella (Bale, 1882). a, three colonies rising from creeping stolon; b, part of axis
with three hydrocladia; c, axial apophysis, nematothecae omitted; d, gonotheca. Scales: a, 3 mm; b, d,
0.1 mm; c, 0.05 mm.
same p l a n e (fig. 25b). E a c h h y d r o c l a d i u m starts w i t h a basal athecate i n t e r n o d e separated b y transverse nodes; i n a d d i t i o n there is a n i n t e r n a l perisarcal r i n g near the
d i s t a l e n d . T h e athecate internode is f o l l o w e d b y a single thecate i n t e r n o d e , b e a r i n g ,
besides the h y d r o t h e c a , a p a i r of f l a n k i n g nematothecae a n d a n u n p a i r e d m e d i a n
nematotheca. H y d r o t h e c a deep campanulate, a d c a u d a l l y c o m p l e t e l y adnate to intern o d e ; m a r g i n straight a n d almost p e r p e n d i c u l a r to h y d r o c l a d i a l l e n g t h axis, s m o o t h
a n d a l m o s t i m p e r c e p t i b l y everted (fig. 25b). The three nematothecae are a l m o s t i d e n -
60
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
tical to those o n internodes of h y d r o c a u l u s ; they are also m o b i l e . Laterals p l a c e d o n
distinct a p o p h y s e s that reach the h y d r o t h e c a l border.
Gonothecae i n s e r t i n g generally o n basal internodes of h y d r o c a u l u s . T h e y are b i g ,
c y l i n d r i c a l w i t h flattened top a n d p r o v i d e d w i t h thick, y e l l o w i s h p e r i d e r m w i t h 8-10
transverse ribs over the w h o l e l e n g t h of the gonotheca (fig. 25d). I n the m a t u r e
gonotheca the truncate top has a c i r c u l a r o p e r c u l u m .
S u b s t r a t u m . — O b s e r v e d o n h a r d a n d soft substrata, the latter b e i n g c o m p o s e d
c h i e f l y of algae.
D i s t r i b u t i o n . — A u s t r a l i a (Bale, 1882, 1894), N e w Z e a l a n d (Totton, 1930; R a l p h ,
1961), T a s m a n i a (according to M i l l a r d , 1975), P e a r s o n I s l a n d (Watson, 1973), S o u t h
A f r i c a ( M i l l a r d , 1975), G o l f o S a n M a t i a s , A r g e n t i n e (Blanco, 1973), Iberian P e n i n s u l a
(Garcia C o r r a l e s et a l . , 1978; R o c a & M o r e n o , 1987). R e c o r d e d f r o m corals a n d shells
of G a s t r o p o d a . O b s e r v e d b y W a t s o n o n r e d algae. The d e p t h d i s t r i b u t i o n is f r o m 0 to
100 m ( M i l l a r d , 1975).
R e m a r k s a n d d i s c u s s i o n . — Totton (1930) stated that the h y d r o r h i z a of M . pulchella bears nematothecae. H e d i d n o t e x p l a i n h i s s y n o n y m y , o n l y says that the t r o p h o s o m a of M.flexuosais smaller t h o u g h v e r y m u c h l i k e that of M. pulchella a n d that the
g o n o s o m e is s l i g h t l y different f r o m that of M . pulchella; M. flexuosa f a l l i n g w e l l w i t h i n
the range of v a r i a t i o n of M. pulchella.
W a t s o n (1973) after the s t u d y of Bale's slides of Plumularia pulchella states that the
species h a s a robust h y d r o c a u l u s , w i t h several septa i n the internodes a n d w i t h o u t
c a u l i n a r nematothecae, w h i l e i n Bale's slides Plumulariaflexuosahas a slender h y d r o c a u l u s w i t h c a u l i n a r nematothecae. A c c o r d i n g to W a t s o n the gonotheca of M . pulchella has n o i n t e r n a l cusps as are present i n M. flexuosa.
A s far as the robustness of the h y d r o c a u l u s is concerned; this character m a y v e r y
w e l l be d e p e n d e n t u p o n the d e v e l o p m e n t of the colony. T h e d e v e l o p m e n t of i n t e r n o d a l septa, a c c o r d i n g to M i l l a r d (1975) is also h i g h l y variable. It r e m a i n s r e m a r k a b l e ,
h o w e v e r , that W a t s o n (1973) d i d n o t observe c a u l i n a r nematothecae i n M . pulchella.
O n the other h a n d , the presence o r absence of i n t e r n a l cusps i n the gonotheca, i f i t is
not d u e to s e x u a l d i m o r p h i s m , is a character to be c o n s i d e r e d , as d i d W a t s o n (1973).
F i n a l l y the authors cited above (Watson, 1973, R a l p h , 1961) state that the gonothecae
of M. pulchella (Bale, 1882) are s m o o t h o r m a y h a v e some i r r e g u l a r ribs ( M i l l a r d ,
1975). T h e gonothecae w e h a v e o b s e r v e d , as w e l l as those of Plumularia femina
d e s c r i b e d b y G a r c i a C o r r a l e s et a l . (1978), h a v e regular transverse ribs o v e r the
w h o l e l e n g t h . W e have o b s e r v e d o n l y female gonothecae. It is possible that s e x u a l
d i m o r p h i s m p l a y s a role a n d that the m a l e gonothecae are s m o o t h ; these h a v e b e e n
d e s c r i b e d w i t h a n d w i t h o u t i n t e r n a l cusps!
I n o u r o p i n i o n t w o distinct species c o u l d be i n v o l v e d , the first w i t h c a u l i n e n e matothecae a n d w i t h a gonothecae w i t h internal cusps a n d a second species w i t h o u t
s u c h c a u l i n e nematothecae a n d l a c k i n g the i n t e r n a l gonothecal cusps, t h o u g h it is
r e m a r k a b l e that M i l l a r d (1975) figures colonies w i t h cauline nematothecae b u t w i t h out cusps i n the gonothecae ( w h i c h are also not m e n t i o n e d i n the description).
C a s t r i c - F e y (1970: 8-12, figs 10-14) d e s c r i b e d specimens of Monotheca f r o m the
G l é n a n Islands i n the n o r t h e r n part of the B a y of Biscay as Monotheca s p . I n her specim e n s the a p o p h y s e s of the a x i a l internodes also bear t w o nematothecae, b u t the
specimens differ f r o m ours b y the stiffness of the h y d r o c a u l u s a n d the considerable
d e v e l o p m e n t of the perisarc, p a r t i c u l a r l y o n the frontal w a l l of the h y d r o t h e c a . These
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
61
characters m a y be greatly i n f l u e n c e d b y t i d a l m o v e m e n t s o r w a v e action. U n f o r t u n a t e l y the specimens w e r e sterile; i n the present context they are d i f f i c u l t to evaluate.
Genus Polyplumaría
G . O . Sars, 1874
PolyplumariaflabellataG . O . Sars, 1874
(fig. 26)
PolyplumariaflabellataG.O. Sars, 1874: 101-102, pi. 2 figs 16-22; Ramil & Vervoort, 1992a: 193-197, fig.
50a-g.
Polyplumaria flabellata p.p. Gili, Vervoort & Pages, 1989: 91-92.
Diplopteron insigne Allman, 1874:479-480, pi. 68 fig. 2.
Polyplumaria pumila Allman, 1883: 31, pi. 4 figs 7-8.
Polyplumaria cantabra Arevalo, 1906: 89-97, pis 15-18.
Polyplumaria Billardi Bedot, 1921b: 14-17, pi 1 figs 4,8-9, pi. 2 figs 12-16, pi. 3 figs 17-18.
Material— F63: (21.vii.1989), one colony without gonothecae (RMNH Coel. no. 27134, slide no. 2582).
Measurements (in mm)
Maximum height of hydrocaulus
50
Internodes of principal axis of hydrocaulus, length
idem, diameter at node
0.27-0.79
0.12
Thecate hydrocladial internodes, length
idem, diameter at node
0.18-0.36
0.048-0.084
Hydrotheca, length abcauline wall
0.120-0.168
idem, length of adcauline wall
0.096-0.144
idem, diameter at rim
0.120-0.144
Lateral nematothecae, length
idem, diameter at
0.084-0.108
rim
0.048-0.060
D e s c r i p t i o n . — H y d r o c a u l i erect, p o l y s i p h o n i c a n d b r a n c h e d , at the base w i t h
some thick stolonal fibres. Branches u s u a l l y p o l y s i p h o n i c over part of their lengths,
a r r a n g e d opposite a n d i n one plane (fig. 26a). P r i n c i p a l tube of stem a n d branches, as
w e l l as secondary tubes w i t h nematothecae. P r i n c i p a l axis of branches i n d i s t i n c t l y
d i v i d e d into internodes, b e a r i n g alternately arranged apophyses s u p p o r t i n g h y d r o c l a d i a directed frontally a n d m a k i n g a n angle of c. 45° w i t h the axis. A p o p h y s e s w i t h
a large ' m a m e i o n ' a n d a p a i r of nematothecae (fig. 26b). H y d r o c l a d i a c o m p o s e d of
thecate internodes separated b y oblique nodes, w i t h the exception of the first intern o d e w h i c h is athecate a n d occasionally f u s e d w i t h the a p o p h y s i s . Thecate i n t e r n o d e
w i t h b i g h y d r o t h e c a , a basal nematotheca, a p a i r of flanking nematothecae
and a
f o u r t h nematotheca above the hydrotheca; this nematotheca m a y occasionally b e
p l a c e d o n a separate internode. H y d r o t h e c a w i t h c. o n e - t h i r d of a d c a u l i n e w a l l
adnate; w a l l s of h y d r o t h e c a g r a d u a l l y w i d e n i n g , m a r g i n s l i g h t l y everted, circular.
L a t e r a l nematothecae just reaching h y d r o t h e c a l r i m or s l i g h t l y shorter; a l l nematothecae i n the c o l o n y conical, w i t h distinct, s l i g h t l y c u p - s h a p e d a p i c a l chamber w i t h deep
e m b a y m e n t of the a d c a u l i n e w a l l (fig. 26c). T h e h y d r o c l a d i u m m a y be b r a n c h e d , the
b r a n c h o r i g i n a t i n g f r o m the first thecate internode; first internode of s e c o n d a r y
h y d r o c l a d i u m w i t h o u t h y d r o t h e c a b u t w i t h a single (apical) nematotheca (fig. 26b).
Fig. 26. Polyplumanaflabellata(G.O. Sars, 1874). a, small colony; b, part of monosiphonic branch with
two hydrocladia, one of which is branched; c, thecate hydrocladial internode with hydrotheca and
nematothecae (only one of laterals shown). Scales: a, 1 cm; b, c, 0.1 mm.
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
63
D i s t r i b u t i o n . — Species of deeper water of the temperate A t l a n t i c (Vervoort, 1966;
G i l i , R a m i l & V e r v o o r t , 1992a).
Genus Pseudoplumaria Ramil & Vervoort, 1992b
Pseudoplumaria marocana ( B i l l a r d , 1930)
(figs 27,28)
Plumularia marocana Billard, 1930: 79, fig. 1; Patriti, 1970: 54-55, fig. 76; Ramil & Vervoort, 1992a: 186191, figs 48b-d, 49a, b.
Pseudoplumaria marocana; Ramil & Vervoort, 1992c: 491-492, fig. 3.
IPolyplumaria Billardi var. deloni Bedot, 1921:17-18, pl. 1 figs 7,10-11; Patriti, 1970: 56, fig. 79.
Polyplumariaflabellatap.p.; Gili, Vervoort & Pages, 1989: 91-92, fig. 19.
Material.— C l l : (31.vii.1991), 27 m, two colonies without gonothecae ( R M N H Coel. no. 27135, slide
no. 2583).
Measurements (in mm)
Maximum height of hydrocaulus
220
Internodes of hydrocaulus, length
0.79-1.41
idem, diameter
Hydrocladial internodes, length of thecate internode
idem, diameter
Hydrothecae, total length
0.11-0.12
0.37-0.45
0.05-0.07
0.16-0.19
idem, diameter at rim
0.16-0.18
idem, length of abcauline wall
0.16-0.19
Lateral nematothecae, length
0.08-0.10
D e s c r i p t i o n . — C o l o n y w i t h thick, p o l y s i p h o n i c , m u c h b r a n c h e d stem r i s i n g f r o m
t h i c k m a t t i n g of h y d r o r h i z a l fibres; b r a n c h i n g a l l i n one p l a n e , w i t h branches o p p o site o r i r r e g u l a r l y d i s t r i b u t e d (fig. 27). A x i a l tubes, as w e l l as those of branches, p a r allel, i n d i s t i n c t l y d i v i d e d into internodes a n d w i t h t w o l o n g i t u d i n a l r o w s of s m a l l
nematothecae. P r i n c i p a l tube f o u n d o n front of colony, w i t h short a p o p h y s e s alternately p o i n t i n g left a n d r i g h t a n d s u p p o r t i n g the (alternate) h y d r o c l a d i a , t w o to six
are f o u n d p e r internode (fig. 28a). O p p o s i t e a p o p h y s i s a large, elevated ' m a m e i o n ' ,
s u r r o u n d e d b y three nematothecae, one above, one beside the ' m a m e l o n ' at the centre of tube a n d one, m o r e d i f f i c u l t to observe, b e h i n d the a p o p h y s i s . H y d r o c l a d i a
w i t h one o r t w o basal athecate internodes w i t h a single nematotheca, f o l l o w e d b y a
regular sequence of thecate internodes separated b y o b l i q u e nodes; the first s e p t u m ,
h o w e v e r , b e i n g transverse (fig. 28a). E a c h thecate internode w i t h b i g , c y l i n d r i c a l
h y d r o t h e c a a n d f o u r nematothecae. W a l l s of h y d r o t h e c a c y l i n d r i c a l , m a r g i n n o n everted. P a i r of f l a n k i n g nematothecae conical, slender, w i t h t h i n r i n g separating
b o t h chambers; a d c a u l i n e w a l l w i t h m i n o r e m b a y m e n t . M e d i a n i n f e r i o r nematotheca
b i t h a l a m i c , short, i m m o b i l e . F o u r t h nematotheca f o u n d i n a x i l b e t w e e n free part
a d c a u l i n e w a l l a n d internode, r e d u c e d to f r o m a p e r i d e r m a l scale (fig. 28b). C o e n o sarc of c o l o n y w i t h m a n y zooxanthellae.
D i s t r i b u t i o n . — A t l a n t i c coast of M o r o c c o a n d Strait of G i b r a l t a r (Patriti, 1970;
R a m i l & V e r v o o r t , 1992a), m a x i m u m d e p t h 1378 m ( R a m i l & V e r v o o r t , 1992a).
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
. 27. Pseudoplumana marocana (Billard, 1930), part of large colony. Scale: 1 cm.
Fig. 28. Pseudoplumaria marocana (Billard, 1930). a, part of colony, frontal view; b, two thecate hydrocladia with hydrothecae and nematothecae. Scales: a, 1 mm; b, 0.1 mm.
66
Medel & Vervoort. Plumularian hydroids. Zool. Verh. Leiden 300 (1995)
Concluding biogeographical remarks
O f the t w e n t y - f o u r species s t u d i e d eight are cosmopolites (Aglaophenia pluma,
Lytocarpia myriophyllum, Antennella secundaria, Kirchenpaueria pinnata, Ventromma halecioides, Nemertesia antennina, N. ramosa a n d Plumularia setacea). N i n e are A t l a n t i c M e d i t e r r a n e a n (Aglaophenia acacia, A. kirchenpaueri, A. octodonta, A. picardi, A. tubiformis, A. tubulifera, Streptocaulus dollfusi, Antennella siliquosa a n d Polyplumaria flabellata).
T h e species i n these t w o g r o u p s are the most a b u n d a n t .
T h e r e m a i n i n g seven species are restricted to the nearby zones of the A t l a n t i c
a n d M e d i t e r r a n e a n . O f these seven species three are c i r c u m t r o p i c a l (Aglaophenia párvula, Halopteris diaphana a n d Nemertesia irregularis). O f the r e m a i n i n g four, Gymnangium montagui has never been f o u n d i n the M e d i t e r r a n e a n p r o p e r ; the B a y of A l g e c i ras a n d P u n t a A l m i n a near C e u t a , f r o m w h e r e the species is n o w r e c o r d e d , are b o t h
u n d e r M e d i t e r r a n e a n influence a n d p r o b a b l y represent l i m i t s of the d i s t r i b u t i o n of
this A t l a n t i c species i n this area. T h e same m a y v e r y w e l l a p p l y to Pseudoplumaria
marocana; this A t l a n t i c species is here recorded f r o m the B a y of A l g e c i r a s . Halopteris
liechtensterni is a p u r e l y M e d i t e r r a n e a n species, penetrating into the Strait of G i b r a l tar. Monotheca pulchella, finally, is here considered as a n I n d o - P a c i f i c species occurr i n g at v a r i o u s places a l o n g the M e d i t e r r a n e a n coast of S p a i n . It m a y h a v e m i g r a t e d
t h r o u g h the Suez C a n a l (Lessepsian m i g r a t i o n ) . The s y n o n y m y of this species, as has
been p o i n t e d o u t above, is b y n o means certain.
Acknowledgements
Research o n the present c o l l e c t i o n has been d o n e p a r t l y i n the L a b o r a t o r i o d e
B i o l o g i a M a r i n a (Zoologia), F a c u l t a d d e B i o l o g i a , U n i v e r s i d a d d e S e v i l l a , S p a i n ,
p a r t l y i n the Vreewijkstraat annex of the N a t i o n a a l 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 en, T h e N e t h e r l a n d s . T h e authors express their gratitude to the authorities of that
M u s e u m a n d to those of the J a n Joost Ter P e l k w i j k f u n d , L e i d e n , for f i n a n c i a l s u p port. W e also w i s h to thank C E P S A , M a n c o m u n i d a d d e Municípios d e l C a m p o d e
G i b r a l t a r F u n d a c i ó n S e v i l l a n a de E l e c t r i c i d a d y E x m o , A y u n t a m i e n t o d e los B a r r i o s ,
for their p a r t i a l f i n a n c i a l a i d for this w o r k .
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Edited: J.C. den Hartog