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A] 
| rf \LES 


de la 

SOCIETE SUISSE DE ZOOLOGIE 

et du 

MUSEUM D'HISTOIRE NATURELLE 
de la Ville de Genève 


tome 114 
fascicule 2 
2007 


E L 
kl GENEVE JUIN 2007 ISSN 0035 - 418 X 


REVUE SUISSE DE ZOOLOGIE 


SWISS JOURNAL OF ZOOLOGY 


REVUE SUISSE DE ZOOLOGIE 


TOME 114— FASCICULE 2 


Publication subventionnée par: 
ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT) 
VILLE DE GENÈVE 
SOCIÉTÉ SUISSE DE ZOOLOGIE 


DANIELLE DECROUEZ 
Directrice du Museum d’histoire naturelle de Genève 


ALICE CIBOIS, PETER SCHUCHERT 
Chargés de recherche au Muséum d’histoire naturelle de Genève 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités 
suisses. a 

Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, 
systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. 


Administration 


MUSEUM D’HISTOIRE NATURELLE 
1211 GENEVE 6 


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ANNALES 


de la 

SOCIETE SUISSE DE ZOOLOGIE 

et du 

MUSEUM D’HISTOIRE NATURELLE 
de la Ville de Geneve 


tome 114 
fascicule 2 
2007 


REVUE SUISSE DE ZOOLOGIE 


SWISS JOURNAL OF ZOOLOGY 


8 : 
EI GENEVE JUIN 2007 ISSN 0035 - 418 X 


REVUE SUISSE DE ZOOLOGIE 


TOME 114 — FASCICULE 2 


Publication subventionnée par: 
ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) 
VILLE DE GENEVE 
SOCIETE SUISSE DE ZOOLOGIE 


DANIELLE DECROUEZ 
Directrice du Museum d’histoire naturelle de Genève 


ALICE CIBOIS, PETER SCHUCHERT 
Chargés de recherche au Muséum d’histoire naturelle de Genève 


Comité de lecture 


Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités 
suisses. 


Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, 
systématique, faunistique, phylogénie, Evolution, morphologie et anatomie comparée. 


Administration 


MUSEUM D’HISTOIRE NATURELLE 
1211 GENEVE 6 


Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


PRIX DE L’ABONNEMENT: 


SUISSE Fr. 225.— UNION POSTALE Fr. 250.— 


(en francs suisses) 


Les demandes d’abonnement doivent étre adressées 
a la rédaction de la Revue suisse de Zoologie, 
Museum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 


REVUE SUISSE DE ZOOLOGIE 114 (2): 175-184; juin 2007 


Two new species of Chionodes Hübner from Ecuador, 
with a summary of known Galapagos records of Gelechiidae 
(Lepidoptera) 


Patrick SCHMITZ & Bernard LANDRY 
Museum d’histoire naturelle, C. P. 6434, CH-1211, Geneva, Switzerland. 
E-mails: patrick.schmitz@ ville-ge.ch / bernard.landry@ville-ge.ch 


Two new species of Chionodes Hübner from Ecuador, with a summary 
of known Galapagos records of Gelechiidae (Lepidoptera). - Two new 
species of the genus Chionodes Hübner, 1825 (Lepidoptera, Gelechiidae) 
from Ecuador are described and illustrated. Chionodes stefaniae sp. n. 
occurs in the Galapagos on the islands of Floreana, Isabela, Pinta, Pinzon, 
Rabida, San Cristobal, and Santa Cruz, where it is believed to be endemic. 
Chionodes manabiensis sp. n. occurs on the Ecuadorian coast in Machalilla 
National Park. The previous Galapagos records of Aristotelia howardi 
Walsingham, 1909, and Stegasta bosqueella Chambers, 1875 were erro- 
neous. Stegasta zygotoma Meyrick, 1917 is reported from the Galapagos for 
the first time. Altogether, five species records of Gelechiidae are now 
considered valid for the Galapagos. 


Keywords: Lepidoptera - Gelechiidae - new species - Chionodes - endemic 
- Galapagos Islands - Manabi. 


INTRODUCTION 


The Gelechiidae represent one of the largest families of Lepidoptera with more 
than 4,530 species described (Hodges, 1998). Kaila (2004: 322) reported that the 
monophyly of the group is supported by 12 synapomorphies of which one is unique, 
i.e. the presence on the forewing of a row of narrow scales ventrally on vein R in 
females only. Genus Chionodes Hiibner, 1825 is found throughout the Holarctic region 
and in the Neotropics. The largest number of species occurs in North America and in 
the mountain zones of the Palearctic region including the Arctic (Huemer & Sattler, 
1995). An apparent apomorphy for Chionodes is the presence of a caecum on the phal- 
lus (Hodges, 1999: 15). In his revision of the North American species, Hodges (1999: 
20, 24, 25) recognized and characterized six species-groups for the 187 species 
occurring in America North of Mexico and the 21 species in the Neotropical region. 
The two species described here belong to the phalacrus-group which already contains 
five described species in the Neotropics (i.e., south of the U.S.A.): C. argosema 
(Meyrick, 1917) described from the Ecuadorian Andes, C. consona (Meyrick, 1917) 
described from Peru, C. donatella (Walker, 1864) described from Jamaica, C. eburata 


Manuscript accepted 12.02.2007 


176 P. SCHMITZ & B. LANDRY 


(Meyrick, 1917) described from Colombia, and C. phalacrus (Walsingham, 1911) 
described from Mexico. Two undescribed species from Brazil were also recorded to 
belong to this group by Hodges (1999: 165). The known host plants of the group are in 
the Malvaceae: Abutilon, Hibiscus, Malacothamnus, Malvastrum, Sida, Sidalcea, 
Sphaeralcea, and Wissadula (Hodges, 1999). 

As part of a project to document the entire microlepidopteran fauna of the 
Galapagos, dissected specimens of the Gelechiidae species collected by Bernard 
Landry (BL) in the Galapagos were critically examined by him with the help of Dr 
Klaus Sattler at the Natural History Museum, London, England (BMNH) in 2000. 
Among them was a new species of Chionodes which was used as an outgroup to study 
the evolution of the Galapagos endemic genus Galagete Landry (Autostichidae; 
Schmitz er al., submitted). And while searching for potential taxa possibly related to 
Galagete at Machalilla National Park, on the coast of continental Ecuador, north of 
Guayaquil, Patrick Schmitz (PS) came upon another new species of Chionodes that 
proved to be different, yet very similar to the Galapagos species and to a few other 
described species. These two new species are described below. 

So far only four valid species of Gelechiidae have been reported for the 
Galapagos. Schaus (1923) reported Aristotelia howardi Walsingham, 1909, and 
Stegasta bosqueella Chambers, 1875. However, we can report here that these two 
records were erroneous and respectively represent Aristotelia naxia Meyrick, 1926, 
described from the Galapagos, and Stegasta zygotoma Meyrick, 1917, described from 
Colombia, Ecuador, and Peru, and for which Clarke (1969) selected a lectotype from 
Ecuador, Huigra, 4,500 ft. The two erroneous records and A. naxia were subsequently 
listed also by Linsley & Usinger (1966) along with Gelechia protozona Meyrick, 1926 
and G. gnathodoxa Meyrick, 1926, which Landry (2002) transferred to the 
Autostichidae. The other previous gelechiid records are those of the widespread inva- 
sives Sitotroga cerealella (Olivier, 1789) and Tuta absoluta (Meyrick, 1917) in 
Causton et al. (2006). Collections hold at least 12 more Gelechiidae species from the 
Galapagos, and they will be treated in a forthcoming paper. 


MATERIAL AND METHODS 


The 57 specimens forming the basis of this study were collected mostly by BL 
during three expeditions to the Galapagos in 1989, 1992, and 2002. Other specimens 
were collected by both of us during two more expeditions on the archipelago in 2004 
and 2005, and by PS in the Galapagos and in Machalilla National Park in 2006. Seven 
additional specimens come from the collection of the Invertebrates Department of the 
Charles Darwin Research Station, Santa Cruz Island, Galapagos (CDRS). In addition 
to this institution, specimens will be deposited in the Natural History Museum, 
London, England (BMNH), the Canadian National Collection of Insects, Ottawa, 
Canada (CNC), and the Museum d’histoire naturelle, Geneva, Switzerland (MHNG). 

The manner of giving the label data of the holotypes and paratypes is presented 
in Landry (2006) and so are the methods used for specimen collecting, genital pre- 
paration, forewing length measurement, and illustrations. The terminology regarding 
genitalia follows Hodges (1999). 


NEW SPECIES OF CHIONODES FROM ECUADOR 177 


DESCRIPTIONS 


Chionodes stefaniae sp.n. Figs 1, 3-8 


MATERIAL EXAMINED: Holotype d, [1] “ECU[ADOR]., GALAPAGOS/ Isabela, 
V[olcan]. Darwin/ 630 mleters] elev[ation]., 17.v.1992/ M[ercury]V[apour]L[amp], leg/ir]. 
B[ernard]. Landry”. [2] “HOLOTYPE/ Chionodes/ stefaniae/ Schmitz & Landry”. Specimen in 
perfect condition except for small hole in left forewing. Deposited in MHNG. 

Paratypes, Ecuador: 19 d, 29 9, from the Galapagos Islands, collected with an ultra- 
violet light and by B. Landry, unless specified otherwise. — Floreana: 1 9, close to Loberia, 
G[lobal]P[ositioning]S[ystem]: S 01°17.002’, W 90°29 460”, 11.iv.2004 (P. Schmitz); 2 2, Las 
Cuevas, 23.iv.1992, M[ercury]V[apour]L[amp]; 1 4 (dissected, slide MHNG 3204), Zona arida, 
300 mfelt[er]s, Finca Las Palmas, 26.xii.1997, UVL-F.L. (L. Roque). — Isabela: 1 3, V[olcan]. 
Darwin, campamento base, 1.iii.2000, Malaise trap (L. Roque); 1 ©, 1 km.W [of] Puerto 
Villamil, 3.11.1989, MVL; 1 ©, 11 km N Puerto Villamil, 9.iii.1989, MVL; 2 9, 11 km N Pto 
Villamil, 13.111.1989, MVL; 1 d (dissected, slide MHNG 3201), NE slope [Volcan] Alcedo, Los 
Guayabillos camp, GPS: elev[ation]. 869 m, S 00°24.976° W 91°04.617’, 2.1v.2004, 4h00-5h30 
(B. Landry, P. Schmitz); 1 ©, [Volcan] Alcedo, lado NE, 200 m, camp arida alta, 14.1v.2002 
(B. Landry, L. Roque); 2 ©, [Volcan] Alcedo, lado NE, 400 m, camp pega-pega, 15.iv.2002 
(B. Landry, L. Roque); 1 9 (dissected, slide MHNG 3208), [Volcan] Alcedo, lado NE, low arid 
zone, bosq[ue]. palo santo, 18.iv.2002 (B. Landry, L. Roque); 3 d, V. Alcedo, 570 m elev., 
11.x.1998 (L. Roque). Pinta: 1 3 (dissected, slide MHNG 3202), 1 9,200 m elev., 16.11.1992, 
MVL; 1 2, 400 m elev., 17.i.1992, MVL; 1 2, N 00°34.476’, W 90°45.102’, 372 m elev., 
16.111.2006 (P. Schmitz, L. Roque); 1 4, 400 m elev., 18.iii.1992, MVL; 3 9, N 00°34.591’, W 
90°45.137’, 421 m elev., 18.iii.2006 (P. Schmitz, L. Roque). — Pinzon: 1 @ (dissected, slide 
PS028), 01.v.2003 (L. Roque). — Rabida: 1 2 , Tourist trail, 3.iv.1992, MVL. —San Cristobal: 1 
2 (dissected, slide MHNG 3209), antiguo botadero, ca. 4 km SE Pto Baquerizo, GPS: 169 m 
elev., S 00°54.800’, W 089°34.574’, 25.11.2005; 1 d (dissected, slide MHNG 3203), near 
Loberia, GPS: elev. 14 m, S 00°55.149°, W 89°36.897’, 16.111.2004 (B. Landry, P. Schmitz). — 
Santa Cruz: 1 3 , Tortuga Res[erve]., W [of] Santa Rosa, 6.11.1989, MVL; 1 9, NNW [of] Bella 
Vista, GPS: 225 m elev., S 00°41.293’, W 090°19.665’, 18.11.2005 (B. Landry, P. Schmitz); 1 9, 
E[stacion]. Clientifica]. C[harles]. D[arwin]., 7.111.1992; 1 © , transition zone, recently cut road, 
GPS: S 00°42.528’, W 90°18.849’, 12.111.2004 (B. Landry, P. Schmitz); 1 ©, El Barranco, 
ECCD, 13.111.2000, MVL Trap (L. Roque); 1 à , Finca S[teve]. Devine, 17.11.1989, MVL; 1 dg, 
ECCD, El Barranco, S 00°44.291’, W 90°18.107’, 22 m elev., 23. iii.2006 (P. Schmitz); 2 à, 
Finca Vilema, 2 km W [of] Bella Vista, 1.iv.1992, MVL; 1 2, C[harles] D[arwin] R[esearch] 
S[tation], [El] Barranco, 20 m elev., 30.1v.2002; 4 4 (one dissected, slide MNHG 3212), 3 2 
(one dissected, slide MHNG 3207), Los Gemelos, 27.v.1992, MVL; 1 6, Barranco, CDRS, 
23.x.2001 (L. Roque). Deposited in the BMNH, CDRS, CNC, and MHNG. 


ETYMOLOGY: We are pleased to name this species in honour of Stefania Bertoli- 
Schmitz for her love and support to PS through the last seven years. 


DIAGNOSIS: Among the species of Chionodes, C. stefaniae is similar in wing 
pattern to C. argosema, C. donatella, C. manabiensis sp. n., C. mariona (Heinrich, 
1921), and C. petro Hodges, 1999 of the phalacrus-group of Hodges (1999). In male 
genitalia, C. stefaniae differs from C. argosema, C. donatella, C. mariona, and C. petro 
in having a shorter mesial projection of the uncus and short, curved and rather stout 
valval projections as opposed to long and thin projections in C. argosema (Clarke, 
1969), and short and straight projections in the other three species (Hodges, 1999). 
Furthermore, C. stefaniae differs from C. donatella, C. mariona, and C. petro in having 
the male abdominal tergum VIII wider than long versus about twice as long as wide 
(Hodges, 1999: pl. W figs 16, 17, 20), and in the female, there is a modification at the 
posterior margin of tergum VII while modifications occur at the posterior margin of 
abdominal tergum VI and anterior margin of tergum VII in the other species (Hodges, 


178 P. SCHMITZ & B. LANDRY 


‘ 
LI 
f 


1 2 
Fics 1-2 
Holotypes of Chionodes spp. (1) C. stefaniae. (2) C. manabiensis. 


1999: pl. VV figs 11-13). Chionodes stefaniae differs from C. manabiensis in several 
characters mentioned in the Diagnosis and Description of this species, below. 


DESCRIPTION: MALE (n=20) (Figs 1, 3-5): Head off-white with yellowish orange 
scales on forehead. Haustellum dark brown; maxillary palpus off-white, 4-segmented. 
Labial palpus dark brown on first segment; second segment off-white to yellowish 
orange with broad scale brush; third segment off-white to yellowish orange, slender, 
dark brown on distal 1/3 to 1/2. Antennal scape dark brown with white scales ventrally; 
flagellum dark brown. Thorax off-white, tegula and metathorax dark brown. Foreleg 
coxa, tibia, femur and tarsomeres dark brown with white at apices of tarsomeres I and 
V. Midleg and hindleg femora and tibiae dark brown at base, apices 1/3 off-white; 
spurs white; tarsomeres I-V mostly dark brown with white at apex of each segment. 
Wingspan: 9.3-11.0 mm (Holotype: 11.0 mm). Forewing dark brown with pair of 
prominent off-white patches with yellowish orange scales on costal margin at 3/4 and 
on inner margin at end of fold; sometimes with slightly darker brown markings visible 
as two small spots submedially in middle, one above the other, and another spot post- 
medially above second off-white patch; with some scattered off-white scales between 
patches, on termen, and around both small spots; fringe dark brown. Hindwing dark 
greyish brown, fringe pale greyish brown. Abdomen dark brown dorsally, off-white 
ventrally; tergum VIII with more thickly sclerotized anterior margin a broad inverted 
V, with posterior margin broadly rounded; sternum VIII with lateral margins slightly 
convex and with conspicuous striae, anterior margin bearing pair of submesial lobes, 
posterior margin with broad and short rounded lateral lobes (apically folded on Fig. 5). 

Male genitalia (n=4) (Figs 3, 4). Uncus with broadly rounded anterobasal lobes 
with stout setae ventrally and few hairlike setae on apical margins; with small, blunt, 
apicomesial projection. Median hook of gnathos rather long and thin, slender from 
base to apex, upturned and pointed apically. Dorsal connection of tegumen wide; 
pedunculi short and broad, shorter than vinculum. Valva with long and slender sickle- 
like projection of 1/2-1/3 X length of tegumen; with small recurved knob at lateral base 
of each projection; sacculus short, with few setae at base ventrally. Vinculum tapering 
to narrowly rounded saccus. Phallus narrow, with distinct, sclerotized rim around 


NEW SPECIES OF CHIONODES FROM ECUADOR 179 


Fics 3-5 


Male genitalia of Chionodes stefaniae from specimen on slide MHNG 3201. (3) Whole genitalia 
without phallus. (4) Phallus; (5) Abdominal segment VIII. 


opening of ductus ejaculatorius, broadest at this point; slightly upturned at 5/6 of 
length; with long, rodlike caecum of about 1/3 X length of phallus; vesica without 
spines or cornuti. 

FEMALE (n=29): Colour as in male. Antenna slightly thinner than that of male. 
Female wingspan: 8.7-10.9 mm. Tergum VII apically modified, with low, median 
depression associated with thin canal directed proximally. 

Female genitalia (n=5) (Figs 6-8). Papillae anales slightly longer than wide. 
Posterior apophyses long, straight, very slightly enlarged at apex (about 2.4 X length 
of papillae), reaching base of antrum. Anterior apophyses partly fused with antrum, 
free proximal section arising from lateral margin of antrum near base, down curved 
with slightly enlarged apex. Antrum well developed with short, longitudinal sclerotized 
band in dorsal wall; also with heavily sclerotized lateral bands of about half length of 
ductus bursae and fused toward apex, with left band bent toward right one. Ductus 
bursae short, of medium girth (width = 0.2-0.25 X its length), slightly constricted at 
apex of lateral bands of antrum. Inception of ductus seminalis at base of corpus bursae. 
Corpus bursae elongate, widening and rounded at proximal end, with light scobination; 
signum situated posteriad middle of corpus bursae, triangular with heavily sclerotized, 
inwardly directed, large spine arising from each angle, sometimes with small extra 
spine. 


BIOLOGY: The moths were collected at light in February, March, April, May, 
October, and December from sea level to 869 m. 


180 P. SCHMITZ & B. LANDRY 


6 8 
Fics 6-8 


Female genitalia of Chionodes stefaniae from specimen on slide MHNG 3207. (6) Whole 
genitalia. (7) Segment VII. (8) Signum. 


DISTRIBUTION: Currently known only from the Galapagos islands of Floreana, 
Isabela, Pinta, Pinzon, Rabida, San Cristobal, and Santa Cruz; presumed to be endemic 
to the archipelago. 


REMARKS: This species and the following one are apparently more closely 
related to each other than to any of the other Chionodes species on the basis of the short 
mesial projection of the uncus, the curved and rather short valval projections, the wide 
abdominal tergum VIII of the male, and the modified posterior margin of tergum VII 
of the female. 


Chionodes manabiensis sp. n. Figs 2,9-14 


MATERIAL EXAMINED: Holotype d, [1] “ECU[ADOR], Manabi, Parque nacional/ 
Machalilla, Los Frailes/ S 01°29.340°, W 80°46.686/ 40 ml{eters] elev[ation]., u[Itra] v[iolet] 
Ifight], 25.iv.2006/ leg/it]. P[atrick]. Schmitz”. [2] “HOLOTYPE/ Chionodes/ manabiensis/ 
Schmitz & Landry”. Specimen in perfect condition except for small notch in left hindwing. 
Deposited in MHNG. 

Paratypes, Ecuador: 4 d,3 ©, from Manabi, collected at uvl by P. Schmitz. 1 9, Puerto 
Lopez, Hosteria Mandala, S 01°32.955’, W 80°48.617’, 10 m elev., 24.iv.2006; 3 d (one 
dissected, slide MHNG 3187), with same data as holotype; 1 9 (dissected, slide MHNG 3210), 
Parque nacional Machalilla, Agua Blanca, S 01°31.421’, W 80°46.081”, 45 m elev., 26.1v.2006; 


NEW SPECIES OF CHIONODES FROM ECUADOR 181 


Fics 9-11 


Male genitalia of Chionodes manabiensis from specimen on slide MHNG 3187. (9) Whole 
genitalia without phallus. (10) Phallus. (11) Abdominal segment VII. 


1 2, Parque nacional Machalilla, Los Frailes, S 01°29.369’, W 80°46.805’, 45 m elev., 
27.1v.2006; 1 d , Parque nacional Machalilla, Los Frailes, S 01°29.053’, W 80°47.064’, 86 m 
elev., 28.1v.2006. Deposited in the BMNH and MHNG. 


ETYMOLOGY: The name of C. manabiensis is derived from that of the type 
locality. 


DIAGNOSIS: Based on wing pattern, C. manabiensis is impossible to distinguish 
from C. argosema, C. donatella, C. mariona, C. petro, and C. stefaniae of the pha- 
lacrus-group of Hodges (1999), but the same diagnostic characters mentioned under 
the diagnosis for C. stefaniae can be applied to separate C. manabiensis from C. argo- 
sema, C. donatella, C. mariona, and C. petro. Chionodes manabiensis differs from 
C. stefaniae in the slightly darker ground colour of the forewing, in the presence of a 
pair of patches of modified scales on abdominal tergum VIII of the males, in the female 
modification of tergum VII, which is located more medially and is associated with a 
low, elongate crest, and in genital characters as mentioned below. 


DESCRIPTION: MALE (n=5) (Figs 2, 9-11): As C. stefaniae, except forewing and 
abdomen darker; wingspan: 9.0-10.0 mm (Holotype: 10.0 mm); abdominal tergum 
VIII more narrowly rounded apically, with pair of patches of modified scales, with 
anterior margins more rounded; abdominal sternum VIII slightly longer, less striated 
laterally, with lobes of anterior margin less heavily sclerotized. 

Male genitalia (n=1) (Figs 9, 10). As in C. stefaniae, except uncus with more 
heavily sclerotized setae ventrally; tegumen longer than vinculum; long projection of 
valva less strongly bent and broader at base; small recurved lobe at base of projection 
slightly longer; sacculus shorter, with more setae ventrally; vinculum broader, 
especially at apex; phallus more broadly bent upward at 3/4 of length. 


182 P. SCHMITZ & B. LANDRY 


Fics 12-14 


Female genitalia of Chionodes manabiensis from specimen on slide MHNG 3210. (12) Whole 
genitalia. (13) Segment VII. (14) Signum. 


FEMALE (n=3): As in C. stefaniae, except colour as in male of C. manabiensis; 
wingspan: 9.5-11.9 mm; modification of tergum VII located more medially and asso- 
ciated with low, elongate crest. 

Female genitalia (n=1) (Figs 12-14). As in C. stefaniae, except papillae anales 
longer (1.5 X longer than broad); posterior apophyses longer (about 3.4 X length of 
papillae); antrum with sclerotized band in dorsal wall much longer, apically reaching 
beyond margin of sternum VII, proximally reaching apices of lateral bands and more 
thickly sclerotized; lateral bands narrow, tapering gradually, at least 2 X length of those 
of C. stefaniae, with left one bending over right one, free apically. 


BroLocy: The adults were attracted to light in April from sea level to 86 m. 


DISTRIBUTION: Currently known only from Machalilla National Park and the 
adjacent town of Puerto Lopez on the Ecuadorian coast (province of Manabi). 


NEW SPECIES OF CHIONODES FROM ECUADOR 183 


REMARKS: The divergence in a DNA fragment of the Cytochrome Oxidase 
mitochondrial gene (555 base pairs long) between C. stefaniae and C. manabiensis is 
6.7% (GenBank accession numbers EF423724 and EF423725). 


ACKNOWLEDGEMENTS 


We are very thankful to the authorities of the Galapagos National Park and those 
of the Charles Darwin Research Station for allowing fieldwork and for logistical 
support in 1989, 1992, 2004, 2005, and 2006. We thank also Carlos Zambrano Bravo 
for the permission to collect in the Machalilla National Park. We are indebted to Lazaro 
Roque-Albelo of the CDRS for his collecting and rearing efforts, and for the loan of 
material. The comments of Ron Hodges, Lazaro Roque-Albelo, and Klaus Sattler on 
our manuscript were also appreciated. This work was carried out with the financial 
support of the Claraz, Lombard, and Schmidheiny Foundations, the Swiss Academy of 
Sciences, the Baslerstiftung für biologische Forschung, the Departement des affaires 
culturelles of the City of Geneva, the MHNG, and the University of Geneva to PS, and 
S. B. Peck, the Galapagos Conservation Trust, and the MHNG to BL. We are most 
grateful to Klaus Sattler for critical information, and Florence Marteau (MHNG) for 
producing the plates. 


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of the British Museum (Natural History). London, 537 and 531 pp. 

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Zoology, Lepidoptera, Moths and Butterflies, Vol. 1: Evolution, Systematics, and Bio- 
geography. Walter de Gruyter, Berlin & New York, X + 491 pp. 

HODGES, R. W. 1999. Gelechioidea, Gelechiidae (part), Gelechiinae (part-Chionodes). In: 
Dominick, R. B. et al. (eds). The Moths of America North of Mexico, fasc. 7.6. Wedge 
Entomological Research Foundation, Washington, D.C., 339 pp. 

HUEMER, P. & SATTLER, K. 1999. A taxonomic revision of Palaearctic Chionodes (Lepidoptera: 
Gelechiidae). Beitrdge zur Entomologie 45: 3-108. 

KAILA, L. 2004. Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an 
exemplar approach. Cladistics 20: 303-340. 

LANDRY, B. 2002. Galagete, a new genus of Autostichidae representing the first case of an 
extensive radiation of endemic Lepidoptera in the Galapagos Islands. Revue suisse de 
zoologie 109: 813-868. 

LANDRY, B. 2006. The Gracillariidae (Lepidoptera, Gracillarioidea) of the Galapagos Islands, 
Ecuador, with notes on some of their relatives. Revue suisse de zoologie 113: 437-485. 

LINSLEY, E. G. & USINGER, R. L. 1966. Insects of the Galapagos Islands. Proceedings of the 
California Academy of Sciences. 4" Series. 33: 113-196. 


SCHAUS, W. 1923. Galapagos Heterocera with descriptions of new species. Zoologica 5: 22-48. 


184 P. SCHMITZ & B. LANDRY 


APPENDIX 


Resumen. Dos especies nuevas del genero Chionodes Hübner en Ecuador con un resumen 
de las especies de Gelechiidae previamente reportadas para Galapagos (Lepidoptera). - 
Dos especies nuevas del genéro Chionodes Hübner, 1825 (Lepidoptera, Gelechiidae) de Ecuador 
son descritas e illustradas. Chionodes stefaniae sp. n. occurre en las Galapagos sobre las islas de 
Floreana, Isabela, Pinta, Pinzön, Räbida, San Cristöbal y Santa Cruz, donde parece ser endémica. 
Chionodes manabiensis sp. n. occurre en la costa ecuatoriana, Parque Nacional de Machalilla. 
Las menciönas anteriores de Aristotelia howardi Walsingham, 1909 y de Stegasta bosqueella 
Chambers, 1875 para Galapagos eran erröneas. Stegasta zygotoma Meyrick, 1917 es reportada 
en las Galäpagos por la primera vez. En conjunto, cinco menciönas de especies de Gelechiidae 
ahora se consideran validas para las Islas Galapagos. 


REVUE SUISSE DE ZOOLOGIE 114 (2): 185-194; juin 2007 


A new Sapromyza Fallen, 1810, from the Swiss Alps 
(Diptera, Lauxaniidae) 


Bernhard MERZ 

Museum d’histoire naturelle, Departement d’entomologie, C. P. 6434, 
CH-1211 Genève, Switzerland. 

E-mail: bernhard.merz@ville-ge.ch 


A new Sapromyza Fallén, 1810, from the Swiss Alps (Diptera, 
Lauxaniidae). - A new Western Palaearctic Sapromyza with a rather well- 
developed presutural dorsocentral seta is described and illustrated. Its 
phylogenetic position is briefly discussed. Keys to the Western Palaearctic 
genera of the Sapromyza-Lyciella group of genera and to the Western 
Palaearctic species of the Sapromyza obsoleta group are given. 


Keywords: Diptera - Lauxaniidae - Sapromyza - new species - Switzerland. 


INTRODUCTION 


Sapromyza Fallén, 1810, is a heterogeneous genus of Lauxaniidae which is 
widespread in the Old and New World. Some 600 species have been described so far 
as «Sapromyza», and over half of them are still placed in this genus. It belongs to the 
subfamily Lauxaniinae and is defined by negative characters, like the absence of a 
postsutural intra-alar seta, without a well-developed presutural dorsocentral seta, 
without setulae on the anepimeron, a flat face, bare veins, and antennae without parti- 
cularities. Already Stuckenberg (1971, p. 570) pointed out that the species of 
Sapromyza are grouped together through a paraphyletic assemblage of characters and 
their separation from Lyciella Collin, 1948, and other genera is not always possible. 
Lyciella, for instance, differs from Sapromyza by the presence of a strong presutural 
dorsocentral seta and some fine setulae on the anepimeron (see Merz, 2003, for details) 
but some «intermediate» species of Sapromyza with 1-3 (smaller) presutural dorso- 
central setae are known (best seen in the type-species of Sapromyza, S. obsoleta Fallén) 
and some Lyciella lack the fine setulae on the anepimeron (these species are now 
placed in two different genera: Poecilolycia Shewell, 1986, and Pseudolyciella 
Shatalkin, 2000). Stuckenberg (1971) observed also that most Australasian and 
Neotropical «Sapromyza» have a strong presutural dorsocentral seta but he did not 
propose new genera for them. 

Recently, a pair of a «Sapromyza» with a distinct presutural dorsocentral seta 
was found in Switzerland. It is described here as a new species of the S. obsoleta group 
which includes in the Western Palaearctic Region the following species: S. amabilis 
Frey, 1930, S. apicalis Loew, 1847, S. obsoleta Fallen, 1820, S. schnabli Papp, 1987, 


Manuscript accepted 12.02.2007 


186 B. MERZ 


S. setiventris Zetterstedt, 1847, and S. simplicior Hendel, 1908. The group is defined 
by the following characters: body subshining yellow brown (partly grey in S. sim- 
plicior), frons rather strongly shining; apex of postpedicel black; head in profile with 
higher gena than width of postpedicel and wide parafacial; postpedicel dorsally 
concave, about twice as long as wide; abdomen without black spots on tergites. 

The two specimens from Switzerland differ from other Western Palaearctic 
genera superficially resembling Sapromyza (e. g., Aulogastromyia, Cnemacantha, 
Eusapromyza, Lyciella, Mycterella, Neoparoecus, Poecilolycia, Pseudolyciella, Tri- 
cholauxania) in diagnostic characters (Papp & Shatalkin, 1998; Shatalkin, 2000; Merz, 
2003, and see key below). It is therefore reasonable to place this species in Sapromyza 
pending a cladistic analysis of the genera involved in this complex. 


MATERIAL AND METHODS 


The specimens of this study are deposited in the Muséum d’histoire naturelle 
Geneve (MHNG). Terminology follows the first chapter in Papp & Darvas (2000), 
except for the terms of the antennae which are according to Stuckenberg (1999). 


SYSTEMATIC PART 


Sapromyza alpina sp. n. Figs 1-15 


MATERIAL: Holotype d, Switzerland: VS, Jeitzinen, 1550 m, 26.VI.1999, Merz & 
Eggenberger (MHNG). Paratype: 19, Switzerland: VS, Visperterminen, 1400 m, 24.VII.1993, 
leg. B. Merz (MHNG). 

The holotype is double mounted on a minutien pin on a polyporus block. It is in 
moderate condition, with some setae on head and mesonotum broken. 


ETYMOLOGY 
The name is derived from the Alps, the mountains of the type locality. 


DIAGNOSIS 

A yellow brown, rather strongly shining species which is placed despite the 
presence of a presutural (and 3 postsutural) dorsocentral setae in Sapromyza. Apical 
half of postpedicel and palpus black; 4-6 rows of short acrostichal setae present; no 
postsutural intra-alar seta; 2 katepisternal setae; no anepimeral setae or setulae; anterior 
tarsus contrastingly black; hind basitarsus in male ventrally along entire length with a 
brush of black setulae; wing hyaline, without setulae on veins; hind margins of tergites 
in female without longer setae or setulae; terminalia in both sexes characteristic. 


DESCRIPTION MALE 

Wing length: 4.1 mm. 

Head: Colouration - Ground colour yellow brown, with strongly shining frons, 
fronto-orbital plates, and occiput; face paler yellowish, matte; parafacial and gena 
slightly microtrichose, dull; occiput dorsally of neck with a white, more or less shining 
patch; apical half of postpedicel, arista except for basal two segments, and apical half 
of palpus contrastingly black, sharply separated from yellow bases (especially in post- 
pedicel). Structure - In profile 1.1 times higher than long, with strongly developed 
occiput; gena with 2 somewhat irregular rows of black setulae along ventral margin; 


A NEW SAPROMYZA 187 


FiGs 1-5 


Sapromyza alpina sp. n. (1) Head, lateral view (female paratype). (2) Antenna in largest 
extension (female paratype). (3) Thorax, dorsal view (female paratype, setae only drawn on left 
side, setulae on right side). (4) Wing (male holotype, drawn in situ). (5) Hind leg of male 
holotype, anterior view. 


1.2 times higher than width of postpedicel and about one third as high as compound 
eye; the latter about 1.3 times higher than long; parafacial about one third as wide as 
width of postpedicel; frons medially with a slight depression; about 1.7 times wider 
than compound eye and about 1.15 times wider than long (measured between posterior 


188 B. MERZ 


margin of posterior ocelli and lunule); in anterior third with about 15 black setulae; 
face more or less flat. Antenna — Scape shorter than wide, dorsally with few black 
setulae; pedicel about as long as wide, apically with the usual circle of black setulae 
and one stronger seta dorsally; postpedicel about twice as long as wide, dorsally 
slightly concave, tip apico-dorsally smooth; arista short setulose, longest rays about as 
long as diameter of arista at base. Mouthparts — Palpus with short, black, stiff setulae. 
Chaetotaxy — 2 probably reclinate fronto-orbital setae (all broken), 1 short ocellar seta 
reaching base of anterior fronto-orbital seta, inserted distinctly within the triangle of 3 
ocelli; 1 inclinate medial vertical seta; 1 eclinate lateral vertical seta; postocellar seta 
inclinate, short, about as long as longest postocular setae; the latter in one row, and one 
additional row posterior on occiput. 

Thorax: Colouration — Everywhere yellow brown, mesonotum very thinly 
microtrichose and therefore conspicuously subshining; pleura less shining. Structure — 
Mesonotum in lateral view almost flat, in dorsal view about 1.2 times longer than wide. 
Chaetotaxy — Acrostichal setae anterior to transverse suture in about 6 irregular rows, 
posterior to suture in 4 rows; setulae of medial two rows slightly longer than those of 
lateral rows; prescutellar acrostichal seta probably at most as long as presutural dorso- 
central seta (judging from size of alveolus); 1+3 dorsocentral setae, the presutural seta 
distinct, but slightly shorter than anterior postsutural seta; 1 presutural intra-alar seta; 
2 supra-alar setae; 1 postalar seta; no posterior intra-alar seta; 1 postpronotal seta; 2 
notopleural setae, the posterior slightly shorter and weaker; 2 (pairs of) scutellar setae, 
no additional setulae between them; 1 proepisternal seta; 1 anepisternal seta (and some 
black setulae in posterior half); 2 katepisternal setae; anepimeron entirely bare, without 
setulae ventrally of wing base; prosternum with few black setulae. 

Wing (Fig. 4): Surface slightly yellowish, entirely hyaline; all veins bare; black 
setulae on costa ending halfway between R2+3 and R4+5; distance between R-M and 
DM-Cu on M1+2 about twice the length of DM-Cu; last section of M1+2 about 1.5 
times longer than penultimate section (between crossveins). 

Legs (Fig. 5): Yellow brown, but anterior tarsus contrastingly black; last 4 tar- 
sal segments of mid and hind leg increasingly darker towards tip; all tibiae with a short 
black preapical dorsal seta; fore femur without ctenidium; hind femur with a subapical 
dorsal seta; mid tibia with one apical seta ventrally; hind basitarsus ventrally 
throughout length with a dense brush of short black setulae. 

Male abdomen and terminalia (Figs 6-11): Yellow brown, less shining than 
mesonotum; tergites black setulose, without longer setulae at hind margins; sternite 5 
(Fig. 6) posteriorly smoothly concave; epandrium in profile (Fig. 7) slightly higher 
than long; cercus distinctly protruding; surstylus (Fig. 8) free, setulose in ventral half, 
over twice as wide as high, evenly curved, without tooth on inner side; hypandrium 
(Fig. 9) not closed basally, with two acute clasps; pregonite (Fig. 10) of irregular shape, 
apically bilobate, the longer tooth apically slightly hooked; postgonite (Fig. 11) bare, 
consisting of two protuberances of different length; aedeagus as long as surstylus, 
apically with short, fine setulae. 


DESCRIPTION FEMALE 
Very similar to male, with following differences (condition in male in paren- 
theses). 


A NEW SAPROMYZA 189 


Fics 6-11 


Sapromyza alpina sp. n., male holotype. (6) Sternite 5. (7) Genitalia, lateral view. (8) Surstylus 
in largest extension. (9) Genitalia, ventral view. (10) Pregonite in largest extension. (11) 
Postgonite and aedeagus. 


Wing length: 4.0 mm. 

Head (Figs 1-2): Gena with only one distinct row of black setulae (two rows); 
frons about 1.25 times wider than long (1.15 times), in anterior half with about 20 
black setulae (about 15 setulae); 2 reclinate fronto-orbital setae (broken off); ocellar 


190 B. MERZ 


seta ending slightly anterior of alveolus of anterior fronto-orbital seta (ending on level 
of alveolus of anterior fronto-orbital seta); postocellar seta about twice as long as 
longest postocular seta (as long as longest postocular seta). 

Thorax (Fig. 3): Acrostichal setae throughout in 4 more or less regular rows 
(6 rows anterior to transverse suture). 

Legs: Hind basitarsus with a row of short, rather wide-set blackish setulae 
ventrally, which do not form a brush as in the male. 

Female abdomen and terminalia (Figs 12-15): Hind margins of tergites without 
stronger black setae; a small, intercalary pair of sclerotized plates present between 
tergite 7 and tergite 8 (Fig. 12); sternite 8 of complicated structure (Figs 13-14), ven- 
trally an apically pointed, setulose plate reaching tip of sternite 9; this plate connected 
at base with a lateral plate which ends in a small tooth; dorsal of this tooth is a weakly 
sclerotized, setulose protuberance; at base of tooth with darker sclerotization, left and 
right tooth connected by a weakly sclerotized «bridge»; cercus about as long as wide 
at base; 3 spermathecae, with smooth surface, the unpaired egg-shaped, the paired 
spherical. 


BIOLOGY 

Both specimens were swept from vegetation in a dry habitat with shrubs, 
extensively managed meadows and open forests. They were collected between 1400 
and 1600 m near the driest and hottest place in Switzerland. No further data about the 
biology of this species is available. 


REMARKS 

With the discovery of a new species with a short, but distinct presutural dorso- 
central seta the separation of Sapromyza, as it is currently defined for the Palaearctic 
and Nearctic Regions (Stuckenberg, 1971; Papp, 1979; Papp & Shatalkin, 1998; 
Shatalkin, 2000; Merz, 2003) from Lyciella and related genera is even more proble- 
matic. Based on its combination of characters S. alpina is assigned to the Sapromyza 
obsoleta group of species. However, this placement is tentative because no cladistic 
anaylsis or comparative study of terminalia of the genera of the Sapromyza-Lyciella 
group has ever been carried out. 

Sapromyza alpina can be separated from other species of the genus by the 
presence of a rather well-developed presutural dorsocental seta and by the combination 
of the entirely black anterior tarsus in both sexes, the presence of a dorsal preapical seta 
on hind tibia, and in the male by the ventral brush of black setulae on the hind basi- 
tarsus, but the hind tibia is unmodified. In external characters the new species can be 
confused with S. obsoleta which has the same colouration of antenna, palpus and fore 
tarsus, and often carries 1-3 longer setulae anterior of the 3 postsutural dorsocentral 
setae. S. obsoleta differs from the new species by the absence of the dorsal preapical 
seta on hind tibia (present in S. alpina). The male carries a brush of strong black setulae 
apicoventrally on hind tibia, and a curved apical spine anteriorly (no modifications of 
hind tibia in S. alpina). Another superficially similar species is S. schnabli with the 
same colouration of head and thorax, and which has also the fore tarsus darkened (but 
not distinctly black along entire length as in S. alpina). However, the hind basitarsus 
of the male is not modified, and the terminalia of both sexes are very different. In 


A NEW SAPROMYZA 191 


Fics 12-15 


Sapromyza alpina sp. n., female paratype. (12) Last segments (intercalary sclerite dotted). 
(13) Genitalia, lateral view. (14) Same, ventral view. (15) Spermathecae. 


S. schnabli the male surstylus is armed with a tooth on inner surface and the post- 
gonites are short and rounded apically. The female sternite 8 is shorter and has a 
different shape. 


KEY TO WESTERN PALAEARCTIC GENERA OF THE SAPROMYZA-LYCIELLA 
GROUP 


The following key allows a separation of the Western Palaearctic genera of 
Lauxaniinae with the combination of the following characters: Presutural intra-alar 
seta present; postsutural intra-alar seta absent; face straight without tubercle or rim; 
wing without horse-shoe like pattern. 


192 


B. MERZ 


Characters given in parentheses apply to all Western Palaearctic species of the 


couplet, but may also be present in some species of the alternative. 


la 


lb 
2a 


2b 
3a 
3b 
4a 
4b 
Sa 
5b 


6a 


6b 


la 


7b 


8a 


8b 


9a 


9b 


10a 


Either R2+3 or R4+5 with some setulae (always with 1+3 dorsocentral 
setae; at least along DM-Cu with brown infuscation) ................... 2 
Wine vems bare (Figs A)... i Saban ate ER 3 
R2+3 ventrally throughout length setulose; wing hyaline, but with 
brownish infuscation along DM-Cu; R4+5 bare . Tricholauxania Hendel, 1925 
R2+3 bare, but R4+5 in basal half dorsally and ventrally with few setu- 


lae; wing with brown pattern on surface ........ Eusapromyza Malloch, 1923 
Mid tibia apico-ventrally with two setae (body black, 6 rows of acro- 

stichal setae, 1+3 dorsocentral setae) ......... Cnemacantha Macquart, 1835 
Mid tibia‘apico-ventrally with one seta... CRE 4 
Arista thick, covered with dense, white or brown to black rays........... 5 
Aristaslender, rays rather sparse, always dark (Piss2)) 7.2 222.2. ce 6 
Postpedicel at most 2.5 times longer than wide; rays of arista brown to 

HACKER ON AT AA PL Neoparoecus Ozdikmen & Merz, 2006 


Postpedicel at least 3 times longer than wide, rays of arista white 

A Re en leap ey aan ÉTAT Mycterella Kertész, 1912 
Postpedicel almost entirely black. Male: hind basitarsus distinctly 
shorter than mid basitarsus; claws of hind leg asymmetrical: one claw 
almost as long as hind tarsus, the other claw as usual. Female: sternite 7 

and tergite 7 fused, forming a sclerotized oviscape (body yellow brown; 

1+3 dorsocentral setae; anepimeron with 1-3 setulae) 

none den PRATI SD Lie Aulogastromyia Hendel, 1925 
At least basal quarter of postpedicel yellow brown (Fig. 2); male and 
female without these modifieationse RR 2 2 O 7 
Anepimeron with few fine, short setulae below wing base; always with 

1+3 dorsocentral setae (mesonotum yellow brown, unstriped) 
OR I aie thew cuneate Lyciella Collin, 1948 
Anepimeron bare; with or without presutural dorsocentral seta........... 8 
Ocellar setae inserted outside the triangle formed by the 3 ocelli 
(without presutural dorsocentral seta; body dull yellow brown including 
postpedicel and palpus; last 1-3 tergites of abdomen each with a pair of 

black spots laterally. Male: shape of postgonites very complicated, with 


black hooks and protuberances) ............... Sapromyzosoma Lioy, 1864 
Ocellar setae closer together, inserted inside ocellar triangle; combi- 
nation of other characters different AEREA 9 


Head and mesonotum grey, frons with a pair of brown vittae, meso- 
notum grey, with two pairs of brown longitudinal vittae; 1+3 dorso- 
central setae which do not stand on darker spots . . Poecilolycia Shewell, 1986 
Frons without a pair of brown vittae; combination of other characters 
differents RL US PRINT Re 10 
Mesonotum usually grey, setae inserted on dark spots; always with 1+3 
dorsocentral setae; 2-4 rows of acrostichal setae, those of the medial two 
rows almost as long as dorsocentral setae ..... Pseudolyciella Shatalkin, 2000 


10b 


A NEW SAPROMYZA 193 


Mesonotum black, grey, or yellow brown; setae not on dark spots; 
medial two rows of acrostichal setae much shorter than dorsocentral 
setae (Fig. 3); usually without presutural dorsocentral seta, or, if present, 
then mesonotum yellow brown.................. Sapromyza Fallen, 1810 


KEY TO WESTERN PALAEARCTIC SPECIES OF THE SAPROMYZA OBSOLETA 
GROUP 


The following key allows a separation of the Western Palaearctic species of the 


Sapromyza obsoleta group with the combination of the following characters: body 
subshining yellow brown (partly grey in S. simplicior), frons rather strongly shining; 
apex of postpedicel black; head in profile with higher gena than width of postpedicel 
and wide parafacial; postpedicel dorsally concave, about twice as long as wide; 
abdomen without black spots on tergites. 


la 
lb 
2a 


2b 


3a 


3b 


4a 


4b 


5a 


5b 


6a 


6b 


Ralous blackan apieal-halt(Eig. 1)... we es me. er. 2 
Palpus yellow brown, rarely slightly infuscated ........................ 6 
Hind tibia without preapical dorsal seta. Male: hind tibia apicoventrally 

with brush of black setulae and with a curved, black spine at tip 

E OE RE SE All S. obsoleta Fallén, 1820 
All tibiae with a preapical dorsal seta. Male: hind tibia without modifi- 
CAHONSE ER er ln. IE IRE N 3 
Fore basitarsus entirely black; presutural dorsocentral seta present 

(Fig. 3). Male: hind basitarsus ventrally on entire length with a brush of 
SELWASEIESS) ne N EEE ER S. alpina sp.n. 
Fore tarsus yellow to dark brown, at least base of basitarsus and last 

tarsal segment paler (yellow brown to brown); sometimes some longer 
setulae anterior the anteriormost postsutural dorsocentral seta present, 

but they are not distinctly thicker than surrounding setulae. Male: hind 
basitarsusswithouemodinecatons sm nn te AI 4 
More than apical half of postpedical black; border between black tip and 
yellow brown base not sharp. Male: genitalia large, surstyli converging, 
apically pointed (Scandinavia and North of Russia) . . . . S. amabilis Frey, 1930 
At most apical half of postpedicel black; border between black tip and 
yellow brown base sharp. Male: genitalia small, surstylus not pointed 
IPC A A ERRO 3) 
Fore tarsus yellow; less than apical half of postpedicel black. Male: sur- 

stylus without tooth on inner side; both branches of postgonite sharply 
pointed, longer than aedeagus. Female: tergite 3 with some distinctly 
longer setae along posterior margin ................ S. apicalis Loew, 1847 
Fore tarsus brown to dark brown except for base of basitarsus and last 

tarsal segment; postpedical black in about apical half. Male: surstylus 

with a tooth on inner side; two branches of postgonite shorter, apically 

not distinctly pointed. Female: tergite 3 without longer setae along pos- 

LÉO MMA RN S. schnabli Papp, 1987 
Mesonotum light grey; frons with a linear dark patch medially 

RITI O SEO ALSO TIRI RO A S. simplicior Hendel, 1908 
Mesonotum uniformly yellow brown; frons without darker patch 
MEGLAUVER SERIEN E LE S. setiventris Zetterstedt, 1847 


194 B. MERZ 


ACKNOWLEDGEMENTS 


I would like to express my best thanks to L. Papp (Budapest) and A. Shatalkin 
(Moscow) for fruitful discussions, to S. Gaimari (Sacramento) for his comments on an 
earlier version of the manuscript, and to F. Marteau (Genève) for her technical help 
with the illustrations. 


REFERENCES 


MERZ, B. 2003. Einführung in die Familie Lauxaniidae (Diptera, Acalyptrata) mit Angaben zur 
Fauna der Schweiz. Mitteilungen der Entomologischen Gesellschaft Basel 52 (2-3) 
(2002): 29-128. 

Papp, L. 1979. 57. abraval. Korhadéklegyek-Pajzstetiilegyek. Lauxaniidae-Chamaemyiidae. 
Fauna Hungaricae 27: 1-59. 

Papp, L. & DARVAS, B. (eds). 2000. Contributions to a Manual of Palaearctic Diptera. Volume 1. 
Science Herald, Budapest, 978 pp. 

Papp, L. & SHATALKIN, A. I. 1998. 3. 37. Family Lauxaniidae (pp. 383-400). In: Papp, L. & 
DARVAS, B. (eds). Contributions to a Manual of Palaearctic Diptera. Volume 3. Science 
Herald, Budapest, 880 pp. 

SHATALKIN, A. I. 2000. Keys to the palaearctic flies of the family Lauxaniidae (Diptera). 
Zoologicheskie Issledovania 5: 1-102 (in Russian; english translation: SCHACHT, W., 
KURINA, O., MERZ, B. & GAIMARI, S. 2004. Zweiflügler aus Bayern XXIII (Diptera: 
Lauxaniidae, Chamaemyiidae). Entomofauna, Zeitschrift für Entomologie 25 (3): 
41-80). 

STUCKENBERG, B. R. 1971. A review of the Old World genera of Lauxaniidae (Diptera). Annals 
of the Natal Museum 20 (3): 499-610. 

STUCKENBERG, B. R. 1999. Antennal evolution in the Brachycera (Diptera), with a reassessement 
of terminology relating to the flagellum. Studia Dipterologica 6 (1): 33-48. 


REVUE SUISSE DE ZOOLOGIE 114 (2): 195-396; juin 2007 


The European athecate hydroids and their medusae 
(Hydrozoa, Cnidaria): Filifera Part 2 


Peter SCHUCHERT 
Museum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. 
E-mail: Peter.Schuchert@ville-ge.ch 


The European athecate hydroids and their medusae (Hydrozoa, 
Cnidaria): Filifera Part 2. - This study reviews all European species 
belonging to the filiferan families Bougainvilliidae, Cytaeididae, Rath- 
keidae, and Pandeidae. Updated diagnoses for the families and genera are 
provided and taxonomic problems solved or at least outlined. Lizzia blon- 
dina is removed from the Bougainvilliidae and transferred to the Rathkeidae 
based on results from 16S sequence data. Dysmorphosa minuta Mayer, 
1900 is regarded as a synonym of Lizzia blondina; morphological and 
molecular evidence is provided. Cytaeis minima Trinci, 1903, formerly 
included in the Hydractiniidae, is transferred to a new genus Podocory- 
noides and included in the Rathkeidae as Podocorynoides minima (Trinci, 
1903). Perigonimus nanellus Stechow, 1919 belongs to the genus Dicoryne 
and is likely a synonym of Dicoryne conybearei (Allman, 1864). Perigo- 
nimus nudus Stechow, 1919 is recognized as a new synonym of Eirene 
viridula (Péron & Lesueur, 1810). The Mediterranean records of Rhizo- 
rhagium michaeli and R. arenosum are most likely based on misidenti- 
fications of Pachycordyle pusilla (Motz-Kossowska, 1905). Annatiara 
lempersi Bleeker & van der Spoel, 1988 is a synonym of Annatiara affinis 
(Hartlaub, 1914). 


Keywords: Cnidaria - Hydrozoa - Bougainvilliidae - Pandeidae - 
Cytaeididae - Ratkeidae - revision - taxonomy - northeastern Atlantic - 
Mediterranean. 


INTRODUCTION 


This study is the third in a series of taxonomic revisions of the European 
Anthoathecata (=Anthomedusae, Athecata). The previous ones being Schuchert (2004; 
Oceaniidae and Pachycordylidae) and Schuchert (2006; Acaulidae, Boreohydridae, 
Candelabridae, Cladocorynidae, Cladonematidae, Margelopsidae, Pennariidae, Proto- 
hydridae, and Tricyclusidae). The present publication treats the families Bougain- 
villiidae, Cytaeididae, Rathkeidae, and Pandeidae. 


Manuscript accepted 10.02.2007 


196 P. SCHUCHERT 


MATERIAL AND METHODS 


For morphological methods see Schuchert (1996; 2004) or Bouillon et al. 
(2004). Where possible, it was attempted to supplement the species descriptions with 
16S DNA sequence information. This allows a much more precise evaluation of the 
specimens used and will facilitate the work of future revisors. The methods to obtain 
16S DNA sequences are described in Schuchert (2005). All new sequences have been 
submitted to the EMBL database under the accession numbers AM411408- 
AM411423. Some of the sequences mentioned here have already been used in 
Schuchert & Reiswig (2006). Phylogenetic analyses were done as described in 
Schuchert (2005). Additionally, Maximum Likelihood analyses were performed using 
the program PHYML described by Guindon & Gascuel (2003). 


ABBREVIATIONS 


BMNH The Natural History Museum, London, England 

BELUM Ulster Museum, Belfast, Northern Ireland 

MHNG Museum d’histoire naturelle de Genève, Switzerland 

MNHN Musee National d’Histoire Naturelle, Paris, France 

NMNH National Museum of natural History, Washington, USA 
ICZN International Code of Zoological Nomenclature 

IRSN Institut Royal des Sciences Naturelles de Belgique, Bruxelles 
ZMUC Zoological Museum Copenhagen, Denmark 

ZMHC Zoological Museum of Harvard College, USA 

ZSM Zoologische Staatssammlung, Munich, Germany 


TAXONOMIC PART 


FAMILY BOUGAINVILLIDAE LÜTKEN, 1850 

SYNONYMS (see Calder, 1988): Bougainvilleae, Lütken, 1850 (corrected to 
Bougainvilliidae by Allman, 1876). Hippocrenidae McCrady, 1959. — Nemopsidae L. Agassiz, 
1862. — Dicorynidae Allman, 1864a. — Atractylidae Hincks, 1868. — Bimeridae Allman, 1872. — 
Margelidae Haeckel, 1879. — Lizusidae Haeckel, 1879. — Thamnostomidae Haeckel, 1879. — 
Pachycordylini Cockerell, 1911. — Clavopsellidae Thiel, 1962. 

DIAGNOSIS: Hydroids solitary or colonial; hydranths stolonal, branched, or 
rarely sessile, arising from creeping hydrorhiza; erect colonies monopodial with 
terminal hydranths, stems monosiphonic or polysiphonic; perisarc terminating either at 
base of hydranths or forming a pseudohydrotheca; hydranths with one or more whorls 
of filiform distal tentacles, tentacles confined to narrow zone below hypostome; 
gonophores free medusae or fixed sporosacs developing either on hydrocauli, stolons, 
or blastostyles. 

Medusae bell-shaped; mouth circular, with simple or dichotomously branched 
oral tentacles inserted distinctly above mouth rim, ending in nematocyst clusters; four 
radıal canals and circular canal; marginal tentacles solid, contractile, either solitary or 
in clusters, borne on 4, 8, or 16 tentacular bulbs; gonads on manubrium, either forming 
a continuous ring or in adradial, interradial, or perradial position; adaxial ocelli absent 
or present. 


REMARKS: Although the genus Bougainvillia — the core-taxon of the family - is 
relatively well defined and easy to recognize, the family Bougainvilliidae as currently 
classified is difficult to delimit from several other families, notably the Cytaeididae 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 197 


(Calder, 1988). The latter family and the Russellidae Kramp, 1957a possess oral 
tentacles that insert well above the mouth rim, a potential synapomorphy. While the 
medusa based genera are relatively easy to assess, this is much more difficult for the 
genera with gonophores reduced to sporosacs. The hydroids of many Bougainvilliidae 
and Pandeidae are very similar and show considerable intraspecific variation 
(Vannucci, 1960). If not provided with gonophores, they are often not objectively 
distinguishable. Pandeid species that have lost their medusa phase would thus become 
indistinguishable from bougainvilliid hydroids. Although it must be assumed that 
medusa reduction has also occurred in some species of the Pandeidae, they would 
presently be classified among the Bougainvilliidae. The classification of the genera 
without a medusa stage is therefore especially unsatisfactory and must be seen as 
provisional. 

Contrary to my earlier view (Schuchert, 2004), I now think that the genera 
Pachycordyle and Silhouetta are closely related to the Bougainvilliidae and cannot be 
separated unambiguously from them with our present knowledge (comp. remarks 
under B. muscus). The two genera are thus included in the key to the genera of the 
Bougainvilliidae, but the Pachycordyle species are not treated again in the present 
paper (see Schuchert, 2004). As discussed in Schuchert (2004), the genus Millardiana 
Wedler & Larson, 1986 should not be included here. 

The genus Lizzella Haeckel, 1879 is regarded as unrecognizable. Haeckel 
(1879) erected the genus Lizzella for Lizzella octella Haeckel, 1879 (= type species for 
the genus). Lizzella octella is not well characterized and Uchida (1927: 146, 233) 
referred it to a young Spirocodon saltatrix (Tilesius, 1818). This ambiguity renders the 
genus doubtful and useless. The genus Lizella should therefore not be used anymore. 
Also the other species included in this genus by Haeckel, Lizella hyalina (Haeckel, 
1880), is a doubtful species. 

In contradistinction to most taxonomic systems in use, Lizzia blondina Forbes, 
1848 is here thought to be more closely related to the Rathkeidae Russell, 1953 than 
the Bougainvilliidae (see Remarks under family Rathkeidae). 


KEY TO THE GENERA OF THE BOUGAINVILLIIDAE: 


la gonophores sporosacs, medusoids, or adult medusa stage not yet known .......... 8 
1b gonophores a free medusa, mature stages known ............................ 2 
2a tougztentaeular.bulbse ee co ea er a le ee en 3 
2b eighizormore tentacularbulbswe. ke nn ee 6 
3a moreithantone.tentaclespersbulb"2...: Re 2 re OE  CE AE + 
3b onestentaclejpenibulb) SE Pete ar ad ker nee ie ie 5 
4a besides filiform tentacles also club-shaped or capitate ones .............. Nemopsis 
4b aliktentaciestrdenticalim structure PER RE PEER RER Bougainvillia 
Sa oralstentaclesibranched en a IE I ee Thamnostoma 
5b oralitentaclesisimple Saito, Bu ee a Re Nubiella 
6a eight tentacular bulbs, these subdivided by cleft ..................... Chiarella* 
6b tentaculanibulbs not'subdivided) by clefttes45 RR if 
Ta Sıtentacularbulbsiwithlseyeralitentacles 2. 200 wa ea Koellikerina 
7b 4 or 8 tentacular bulbs with a single tentacle ..................... Thamnostoma 
8a hy droidioccunineanireshwaterolicayes EE PR RO EN eka Velkovrhia 
8b hydroid occurring in seawater, rarely in brackish waters ...................... 9 
9a gonophores on special blastostyles, released as swimming sporosacs ....... Dicoryne 


9b gonophoresionhydrocaulivormstolonsree eee OO 10 


198 P. SCHUCHERT 


10a  gonophores a free medusa, adult not yet known ...................... Silhouetta 
10b  gonophores a sporosac or a medusoid (medusa without tentacles).............. 11 
lla  hydranth with pseudohydrotheca, one or more whorls of tentacles .............. 12 
11b  hydranth without pseudohydrotheca, two or more whorls of tentacles . . Pachycordyle° 
12a  pseudohydrotheca sheathes also basal part of tentacles .................. Bimeria 
PE basalipartoftentacles freelofipseudohydrotheca 35. 325 NET ep 13 
Ba oneswhorl'ofitentacles}..... ...%...: tele cie ee ct A AR 14 
IS beemtworormoreswhorlsioftentacles 2.0.2... 2.0.2 ET Parawrightia* 
NA AEC OLOMICS ELECL Eres ee e AS SR IT EI Garveia 
IA be coloniesistolonall.+3%., RR LE DEIR) RR RE PE Rhizorhagium 


* not treated here, as no species recorded in the area under investigation. 
° see Schuchert (2004) 


Genus Bougainvillia Lesson, 1830 


TYPE SPECIES: Bougainvillia macloviana Lesson, 1830, by monotypy. 

SYNONYMS: Hippocrene Brandt, 1835. — Margelis Steenstrup, 1850. — Perigonimus M. 
Sars, 1846. — Atractylis Wright, 1858 [type species = Eudendrium ramosum sensu Van Beneden, 
1844 = Bougainvillia muscus, by designation of Totton, 1930]. — Lizusa Haeckel, 1879. 

DIAGNOSIS: Hydroid colony stolonal or branched, more rarely hydranths sessile; 
perisarc terminating at base of hydranth or extending onto hydranth as pseudo- 
hydrotheca; hydranth cylindrical to spindle-shaped, tentacles in one or two closely 
approximated whorls, tentacle-bases never enveloped by pseudohydrotheca, tentacles 
alternately inclined up- and downward (amphicoronate). Gonophores develop into free 
medusae, arising singly or in clusters from stem, branches or stolons. 

Medusa with four perradial marginal bulbs bearing two or more identical ten- 
tacles, with or without ocelli; four perradial oral tentacles, usually branched and ending 
in nematocyst clusters; gonads interradial or adradial on manubrium, sometimes also 
along basal perradial extensions of the manubrium. 


REFERENCES: Kramp (1926, taxonomic history of genus); Vannucci & Rees 
(1961, synopsis); Edwards (1966, British species); Calder (1988, taxonomic history 
and synonymy). 


KEY TO THE BOUGAINVILLIA HYDROIDS OF THE NE-ATLANTIC AND MEDITERRANEAN: 


la eoloniesterect, branched/severalttimes "Aer eme 2 
lb colonies stolonal (= unbranched), or sparingly branched ...................... 4 
2a if polysiphonic; then all tubes roughly equal... "EU “Ses See eee 3 
2b polysiphonic, central tube much thicker, medusae without ocelli....... B. muscoides 
3a colonyarresulariy branched Lr RR le Sonn CE OE B. muscus 
3b main axis and branches straight, medusa buds grouped on branched stalks 
ee Ne hes Aes Se SR B. pyramidata 
4a hydranths sessile or with very short pedicel .....1.7.......0.02...2 ee 5 
4a hydranths with distinct pedicels)... : 21%... eo nee RER eee 6 
Ja peduncle present in advanced medusa bud ...................... B. superciliaris 
Sb no,peduncleipresentän advanced medusa bud 2245 SER B. principis 
6a very long pedicels, buds in clusters on branched stalks .............. B. britannica 
6b pedicels not particularly long, buds not clustered .................. B. macloviana 


KEY TO THE ADULT BOUGAINVILLIA MEDUSAE OF THE NE-ATLANTIC AND MEDITERRANEAN: 
la gastric peduncle absent or short (much smaller than manubrium) ................ 2 
lb distinct, gastric peduncle’ present 24 MAIL I ee eee Se eee 8 
2a Without medusae ‘budding from manubrium 452.2295 -4 5-6] eee eee 3 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 199 


2b With medusacibuddine fromimanubnume nr ee... 7 
3a MWathoce PRE RER Te ee a + 
3b Withoutioceli ne re AO à RES AGI LISI LEE 6 
4a I-4>*mmbWlessithemtgitentaclestpe bb RR ees B. muscus 
4b >iimmamoresthanisstentaclesiperibUlbXe RER ER 5 
5a oral tentacles with long trunk, crescentic or oval ocelli on tentacles...... B. britannica 
Sb oral tentacles with short trunk ocelli round and on bulbs .............. B. principis 
6a 3-5 mmihiehs-7/stentaclesiperibulb EE nr A er B. muscoides 
6b < Amm 2 -ontentac les per DUI bye ga ea: B. aurantiaca 
7a oral tentacles basal trunk absent, stomach flat ..................... B. platygaster 
7b oral tentacles with long trunks, stomach flask-shaped ................... B. niobe 
8a Gonads also on perradial outgrowths of manubrium, ocelli on bulbs ............. 9 
8b no perradial outgrowths of manubrium, ocelli crescent-shaped on tentacles 

ERM Pee OR ES ER ONE ia RE se B. superciliaris 
9a 4-9 tentacles per bulb, oral tentacles with moderately long basal trunk . . B. pyramidata 
9b 30-65 tentacles per bulb, oral tentacles with short basal trunk ........ B. macloviana 
Bougainvillia muscus (Allman, 1863) Figs 1-3 


Eudendrium ramosum. — Van Beneden, 1844: 56, pl. 4. figs 10-13. 

[not Eudendrium ramosum (Linnaeus, 1758)] 

Medusa octocilia Dalyell, 1847: 70, pl. 11. figs 9-10. — Edwards, 1966: 141, synonym. 

Bougainvillia britannica. — Wright, 1859a: 108-109, pl. 1 figs 1-3. 

[not Bougainvillia britannica (Forbes, 1841)] 

Bougainvillia diplectanos Busch, 1851: 22, pl. 2, fig. 9. — Haeckel, 1879: 80, synomyn. 

Atractylis ramosa. — Wright, 1859a: 108-109, pl. 1 figs 1-3. 

Perigonymus muscus Allman, 1863: 12. 

Podocoryne alderi Hodge, 1863: 82, pl. 2 figs 11-15 [medusa only, not hydroid]. — Calder, 1988: 
27, synonym. 

Corynopsis alderi. — Allman, 1864a: 354 [medusa, not hydroid]. 

Bougainvillia maniculata Haeckel, 1864: 327. — Haeckel, 1879: 89, pl. 5 figs. 4-5. — Mayer, 
1910: 170, fig. 92. — Kramp, 1961: 78. — Vannucci & Rees, 1961: 71. 

Bougainvillia fruticosa Allman, 1864b: 58. — Hincks, 1868: 110. — Allman, 1872: 314, pl. 9 figs 
1-4. 

Bougainvillia ramosa. — Hincks, 1868: 109, pl. 19 fig. 2. — Allman, 1872: 311, pl. 9 figs 5-74. — 
Hartlaub, 1911: 183, figs 162-167. — Broch, 1911: 17, fig. 13. — Brink, 1925a: 726. — 
Brink, 1925b: 126-203, text figs A-C, pls 6-71. — Kramp, 1926: 43. — Russell, 1938b: 
152. — Vervoort, 1946: 135, figs 52a & 53. — Russell, 1953: 153, figs 74A-C, pl. 8 fig. 1, 
pl. 9 figs 4-5 — Kramp, 1959a: 109, fig. 91.—Kramp, 1961: 81.— Vannucci & Rees, 1961: 
82. — Kramp, 1968: 34, fig. 87. — Morri, 1981: 50, fig 15. — Hirohito, 1988: 97, figs 
34 b-f. 

Bougainvillia muscus.— Hincks, 1868: 111. — Allman, 1872: 317, pl. 10 figs 1-3. — Calder, 1988: 
24, figs 19-20. — Schuchert, 1996: 31, fig. 15a-e. — Pefia Cantero & Garcia Carrascosa, 
2002: 22, fig. 3a-c. 

Lizusa octocilia. — Haeckel, 1879: 80. 

Bougainvillia autumnalis Hartlaub, 1897: 465, pl. 15 figs 11-13. — Hartlaub, 1911: 184, 
synonym. — Neppi & Stiasny, 1913: 29, pl. 2, figs 17-21. — Schmidt & Benovic, 1977: 
636. 

in part Bougainvillia flavida Hartlaub, 1897: 456, pl. 14 fig. 5 [female medusa only]. 

Bougainvillia benedenii Bonnevie, 1898a: 484, pl. 26 figs 34-35. — Brink, 1925b: 36, synonym. 
— Rees, 1956b: 113, synonym. 

? Bougainvillia obscura Bonnevie, 1898b: 7, pl. 1 fig. 4. — Hartlaub, 1911: 184. 

Lizusa octociliata. — Aurivillius, 1898: 114 [incorrect subsequent spelling]. 

Bougainvillia Gibbsi Mayer, 1900a: 5, pl. 4 figs 14-15. 

Bougainvillia triestina Hartlaub, 1911: 154, fig. 138. 

Bougainvillia ramosa var. nana Hartlaub, 1911: 189. 


200 P. SCHUCHERT 


Fic. 1 


Bougainvillia muscus (Allman, 1863). (A) Silhouettes of hydroid colonies, showing variability 
from small, monosiphonic (left) to large polysiphonic colony, scale bar 1 cm. (B) Hydranths with 
medusa buds, scale bar 0.5 mm. (C) Newly released medusa, about 0.8 mm high. (D) Lateral and 
oral view of a cultivated, mature medusa with low tentacle numbers and unbranched oral 
tentacles, scale bar 0.2 mm. (E) Manubrium seen from aboral of an older stage than shown in E 
(not same scale). (F-H) Mature medusae from plankton, note variation of manubrium and gonad 
shape, not to scale, bell heights are F=2.5 mm, G=2.2 mm, H= 1.8 mm. A, after preserved 
material; B-E, after living animals; F, modified after Hartlaub (1897); G-H, modified after 
Russell (1953). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 201 


FIG? 
Bougainvillia muscus (Allman, 1863), living colony from Villefranche-sur-Mer, Mediterranean. 


Bougainvillia ramosa var. minima Kramp & Damas, 1925: 254, figs 5-7. 

Bougainvillia autumnalis var. magna Babnik, 1948: 290, fig. 2. — Schmidt & Benovic, 1977: 636. 
Bougainvillia ? ramosa. — Millard, 1975: 97, figs 33EH. 

Bougainvillia ramosa f. musca. — Millard, 1975: 99. 

Bougainvillia ramosa f. fruticosa. — Millard, 1975: 99. 

Bougainvillia ramosa f. ramosa. — Millard, 1975: 99. 

Bougainvillia ramosa f. vanbenedeni. — Millard, 1975: 99. 


202 P. SCHUCHERT 


FIG. 3 


Bougainvillia muscus (Allman, 1863), living colony from Norway, scale bar 5 mm, inset shows 
a higher magnification of a hydranth and medusa buds. This form with its straight stem, the 
clustered medusa buds, and unusually coloured hydranths was first mistaken for B. pyramidata 
(comp. Fig. 9A). Rearing experiments showed it to be B. muscus. 


MATERIAL EXAMINED: MHNG INVE 29410, St. Pol de Leon, Brittany, 20 m; 6 June 2000, 
on Ciona intestinalis, examined alive, colonies 4 cm, polysiphonic, much branched (ramosa- 
fruticosa form), medusa grown to maturity, ocelli red; 16S sequence AY787880. — France, 
Brittany, Roscoff, under l’Estacade (pier of Ile de Batz ferry), on algae; 15 September 2004, 1 
cm colonies, monosiphonic (muscus form), branched; newly released medusae with dark- 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 203 


B. principis, Scotland, AM183128 


B. principis, Norway 
Pachycordyle pusilla, AM183132 


B. muscus Norway, surface, AM411408 


B. muscus, Norway, deep water 


B. muscus, Atlantic, Roscoff, muscus form AM183126 


B. muscus, Atlantic Roscoff, ramosa-fruticosa form, AY787880 
B. muscus, New Zealand, medusa, AM411411 


B. muscus, Mediterranean, Banyuls, M411409 


0.01 B. muscus, Mediterranean, Villefranche, AM411410 


FIG. 4 


16S sequence differences of B. muscus samples represented as a Neighbour-joining tree based 
on sequence divergences (HKY model, indels ignored). The length of the sum of the horizontal 
lines is a measure of the sequence difference. 


crimson-red ocelli, 2 tentacles per marginal bulb, oral tentacles unbranched or branched once, 
Jelly thin, cultivated medusae developed gonads within days, 16S sequence AM183126. - MH- 
NG INVE48732, 48733, 48746, and 48750; Norway, Bergen, near Espeland marine station; 
growing on Mytilus,0.5-1 m depth, June 2006, numerous large, polysiphonic colonies; examined 
alive, some colonies with medusa buds, others developed buds in culture; medusae of one colony 
(MHNG INVE 48746) grown to sexual maturity, marginal bulbs then with 2-3 tentacles and oral 
tentacles branched or not; extracted DNA, 16S sequence AM411408.— Norway, Korsfjord, great 
Skorpa Island; depth 80-160 m, collected by scraping almost vertical wall, 16 June 2006, infer- 
tile colony on bryozoan, used for DNA extraction, 16S sequence identical to AY787880. — 
Aquarium Zoological Institute Basel, 19 January 1991, 13°C, material originating from Banyuls- 
sur-Mer, examined alive, medusae cultivated to sexual maturity. - MHNG INVE32947, 
Mediterranean, marina of Banyuls-sur-Mer, 7 May 2002, 0 m, 1-5 cm colonies, polysiphonic, no 
medusa buds, 16S sequence AM411409. — Mediterranean, Villefranche-sur-Mer 4 April 2005, 
several colonies with medusa buds, on Phallusia mammilata, colonies monosiphonic, up to 2 
cm, 16S sequence AM411410. — Several living medusae collected 6 July 2002 near Motutapu 
Island (Hauraki Gulf, New Zealand), up to five tentacles per marginal bulb, otherwise typical, 
16S sequence AM411411.—- MHNG INVE26550, Naples, collection pre 1900, large colonies. — 
BMNH 1957.3.2.129, Great Britain, Eddystone, 15 September 1897, slide labelled as 
Bougainvillia ramosa, 16 mm colony with buds. -— BMNH 1957.3.2.137, Great Britain, 
Eddystone, 7 December 1897, slide labelled as Bougainvillia ramosa, 40 mm colony with buds. 
— BMNH 1957.3.2.163, Great Britain, Eddystone, 22 August 1898, slide labelled as 
Bougainvillia ramosa, 25 mm colony without buds. - BMNH 1957.3.2.134, Great Britain, 
Eddystone, 19 November 1897, slide labelled as Bougainvillia ramosa, 35 mm colony with 
buds. - BMNH 1985.9.10.22, Great Britain, Millport, Keppel Pier, 2 6 September 1940, labelled 
as Bougainvillia ramosa, 2 medusae in formalin, mature with 2 tentacles per bulb. - ZMUC 
without number, Denmark, July 1947, labelled as Bougainvillia ramosa, juvenile medusae, 
likely released from polyp, det. Kramp. 


DIAGNOSIS: Hydroid colony erect, irregularly branched, mono- or polysiphonic, 
hydranth with 10-18 tentacles, medusa buds in rows on branches. 


204 P. SCHUCHERT 


Medusa 1-4 mm, no distinct peduncle, usually two to four tentacles on each 
marginal bulb, oral tentacles branching zero to two times (rarely three times), ocelli 
round, on marginal bulbs, red or black. 


DESCRIPTION: Hydroid colony varies in growth and form from monosiphonic, 
sparingly branched stems (‘muscus’ form) to tall irregularly and profusely branched 
tree-like forms with fascicled stems (‘ramosa’ and ‘fruticosa’ forms); stolons tubular, 
forming a network; perisarc infested with detritus, corrugated at bases of branches but 
not regularly annulated, thinning out over hydranths, pseudohydrotheca usually present 
in animals from nature, absent in cultivated animals; hydranths terminal on branches 
and branchlets, with 10-18 tentacles, hypostome rounded to conical; gonophores pear- 
shaped, on unbranched stalks of variable length, arising singly, in rows, or in groups 
on the branches just below the hydranths. Especially in distal regions of colony 
regularly stolon-like, naked outgrowths that serve to attach the colony or broken parts 
of it to other substrata (apical stolons, tendrils). Nematocysts: microbasic euryteles and 
desmonemes. Colours: hydranths pink, red, orange, or greyish purple; colour depends 
on food items. 

Medusa at release bell-shaped; jelly uniformly thin, a few nematocysts on 
exumbrella, apical canal may be present, no peduncle; two tentacles and two ocelli on 
each bulb, ocelli red or black; four oral tentacles unbranched. In females, small 
oogonia can be visible on the manubrium, but mostly these are not developed yet. 

Medusa with mature gonads bell-shaped, jelly rather thick, thicker at apex, very 
low gastric peduncle sometimes present; radial canals narrow; bulbs rather small, oval; 
2-6 tentacles per bulb (cultivated animals; when found in plankton usually 3-4); bulbs 
rounded, ocelli round, situated adaxially on bulbs near origin of tentacles; manubrium 
short (about !/2 of subumbrella), conical or in large specimens square- to slightly cross- 
shaped; oral tentacles divided dichotomously two times (rarely 3-4), basal trunks of 
moderate length, length variable; gonads interradial pads, sometimes extending slight- 
ly along radial canals. Nematocysts: microbasic euryteles and desmonemes. Colours: 
marginal bulbs and stomach reddish brown, red or yellowish; sometimes with greenish 
tinge; depends on food items. 


DIMENSIONS: Hydroid colonies with medusa buds variable from a few mm to 12 
cm; hydranth length 0.7-0.9 mm; medusa buds up to 0.5 mm. Medusa at release about 
0.7 mm wide and high. Adult medusa 1-4 mm wide and high. Nematocysts (Medi- 
terranean material, polyps and young medusae): microbasic euryteles (7-8)x(3.5-S)um; 
desmonemes (3-4)x(4-5.5)um. For more nematocyst measurements see Russell 
(1938b), Millard (1975), and Schuchert (1996). 


DISTRIBUTION: A widely distributed species, perhaps also due to human activity. 
Bougainvillia muscus has been recorded in the eastern Atlantic from the Norwegian 
(Bonnevie, 1901; Rees, 1952: Christiansen, 1972) and Swedish coasts (Rees & Rowe, 
1969), Denmark (Rasmussen, 1973), the Baltic Sea (Schönborn er al., 1993); Germany 
(Hartlaub, 1911), Great Britain and Ireland (e. g. Hincks, 1868; Russell, 1953; Ballard 
& Myers, 2000), Holland (Vervoort, 1946; Tulp, 2005), Belgium (Leloup, 1947); 
France (Teissier, 1965; Fey, 1970); Portugal and Spain (Medel & Löpez-Gonzälez, 
1996), Strait of Gibraltar (Pena Cantero & Garcia Carrascosa, 2002), southwards to at 
least Morocco (Patriti, 1970) and Cape Verde Islands (Vanhöffen, 1912). Often also 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 205 


recorded in the Mediterranean (Pefia Cantero & Garcia Carrascosa, 2002), rarely in the 
Black Sea (Valkanov, 1957). The records from Iceland are uncertain, it is absent from 
Greenland (Schuchert, 2001). Also known to occur along the east coast of America 
(Mayer, 1910, as B. autumnalis; Vannucci, 1957; Calder, 1988), northern Pacific 
(Hirohito, 1988), tropical Pacific (Pictet, 1893; Bouillon, 1980), southern Pacific 
(Schuchert, 1996); South Africa (Millard 1975; Gili et al., 1989). Type locality: 
Torquay, Devon, Great Britain, in tide pools near low-water mark. 


BIOLOGY: Bougainvillia muscus hydroids have been found on a wide variety of 
substrata, e. g. seaweeds, hydrozoans, sponges, mollusc shells, polychaete tubes, and 
man made objects (Pefia Cantero & Garcia Carrascosa, 2002). In temperate waters, they 
seem to bud medusae from spring to autumn, planktonic medusa are sometimes also 
present in winter (Hartlaub, 1911). The hydroid of Bougainvillia muscus is a common, 
eurybathic form, found from tidal levels to about 100 m depth (Pefia Cantero & Garcia 
Carrascosa, 2002), but it can also occur in deep waters, e. g. 1193 m (Gili et al., 1989). 
Usually it occurs in fairly wave sheltered conditions, often in slightly reduced salinity 
and also harbours. Considering the abundance of the hydroid along the European coasts, 
the medusa seems to be remarkably scarce in the plankton (Russell, 1953). 


16S SEQUENCE DATA: A phylogenetic analysis of the available Filifera sequences 
always placed the various sequences of B. muscus within a well supported mono- 
phyletic lineage (results not shown, but see Schuchert & Reiswig, 2006). The closest 
relative being surprisingly Pachycordyle pusilla, followed by B. principis. The close 
relationship of P. pusilla hints at major discrepancies with our current macro- 
taxonomic system. 

The available B. muscus sequences are not numerous enough for a serious 
phylogeographic analysis, but they nevertheless allow some interesting preliminary 
comparisons. Figure 4 shows a tree based on distance measures (due to the few shared 
differences other phylogenetic analyses like maximum parsimony make little sense). 
All B. muscus sequences are quite distant from B. principis (>10%, including base 
differences and insertions) and P. pusilla (>6.6%), while they are relatively similar to 
each other (max. 1.5% divergence). There are no obvious, distinct intraspecific 
lineages. The observed intra- and interspecific differences are comparable to those 
observed for some Corynidae (Schuchert, 2005; see also under Lizzia blondina). 
Polysiphonic and monosiphonic colonies from Roscoff — both mature and liberating 
medusae — had identical sequences, confirming earlier conclusions that they belong to 
the same species. The divergence of the two Mediterranean samples is 0.5%, those 
from Norway 0.34%. These values can be taken as a first estimate of the intra-popu- 
lation variation. Higher divergences are observed between the Mediterranean and 
Norwegian samples, reaching values of 1.2%. Comparing the European samples with 
one from New Zealand gives slightly higher divergence values, but the values overlap 
with what is seen for the differences between Atlantic and Mediterranean specimens. 

This all suggests that the examined B. muscus likely belong to the same lineage 
and biological species, confirming its widespread, almost cosmopolitan distribution. 


REMARKS: This species has a complicated synonymy which has been worked 
out by several authors, e.g. Hartlaub (1911), Brink (1925a, 1925b), Russell (1953); 
Vannucci & Rees (1961); Edwards (1964a, 1964b), Calder (1988), to cite only the most 
important. 


206 P. SCHUCHERT 


I agree with Mayer (1910) and Vannucci & Rees (1961) that Bougainvillia 
maniculata is likely a degenerate form referable to B. muscus. The species has only 
been observed by Haeckel, despite its type locality being Villefranche-sur-Mer, one of 
the most thoroughly investigated localities, whose the plankton has been continuously 
observed for the decades (Trégouboff & Rose, 1978; Goy, 1973; Buecher & Gibbons, 
1999). 

Bougainvillia muscus is a very variable, in its polyp as well as in the medusa 
stage. Earlier authors attempted to attribute different forms to different species. In the 
hydroid phase, some authors distinguished a muscus and fruticosus form (see Russell, 
1953; Millard, 1975). The former having small, monosiphonic colonies, the latter 
large, polysiphonic ones. The nominal species B. vanbendeni is characterized by its 
distal branches producing stolon-like tendrils (terminal search stolons). In a thorough 
analysis, Brink (1925a, 1925b) showed that these forms are not objectively separable 
and likely due to environmental influences. 

Likewise for the medusa there are several specific and varietal names available. 
Bougainvillia octocilia (Dalyell, 1847), Bougainvillia ramosa var. nana Hartlaub, 
1911 or Bougainvillia ramosa var. minima Kramp & Damas, 1925 represent mature 
medusa with only 2-3 tentacles per bulb and the oral tentacles are unbranched or 
branched once. Russell (1953: 158) thought it possible that two species might be 
involved, one producing black ocelli, the other red ones. Calder (1988) also considered 
the status of many nominal species as unclear and not resolvable. Here they are 
considered as variants of B. muscus. 

Material from Norway, Roscoff and the Mediterranean was used to cultivate 
medusae to sexual maturity. Some medusae reached a stage with three tentacles per 
bulb, but often developed only to a stage with two tentacles per bulb and the oral 
tentacles remained unbranched. This does not conform with the commonly described 
planktonic adult (see above). It is likely that this growth retardation is due to the sub- 
optimal in-vitro cultivation conditions. Similar results were obtained by Schuchert 
(1996) and (Calder 1988), who assumed that the full size is only achieved under natural 
conditions in the sea, but similar adult medusae have also been observed in the 
plankton (Hartlaub, 1911; Kramp & Damas, 1925). Wright (1859a) was able to 
cultivate the medusa to a stage with six tentacles per bulb. 

Reports of Bougainvillia muscus medusa with eight or more tentacles per bulb 
are likely due to a confusion with other similar species, notably B. pyramidata or B. 
britannica. 

The polyp stages of polysiphonic B. muscus and B. pyramidata are very similar 
and may not always be distinguishable (see under B. pyramidata). 


Bougainvillia britannica (Forbes, 1841) Fig. 5 


Hippocrene britannica Forbes, 1841: 84, pl. 1 figs 2a-c. 

Bougainvillia britannica. — Lesson, 1843: 291. — Forbes, 1848: 62, pl. 12 fig. 1. — Hartlaub, 
1911: 160 & 162, figs 141-151. — Kramp, 1926: 43. — Russell, 1938b: 152, fig.s 23-25. 
— Russell, 1953: 158, figs 75A-B, 77A-B, 78A, 79A, pl. 8 figs 2-3, pl. 9 figs 1-3. — 
Vannucci & Rees, 1961: 61.— Edwards, 1964a: 1, figs 1-2.— Edwards, 1966: 147.— Tulp, 
2005: 47, fig. 1. 

Bougainvillia dinema Greene, 1857: 246, pl. 15 fig. 6a-f. — Edwards, 1966: 141, synonym. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 207 


Fic. 5 


Bougainvillia britannica (Forbes, 1841). (A) Hydroid colony with medusa buds. (B) Newly 
released medusa. (C) Two days old medusa, size ca. 1 mm. (D) Subadult medusa, scale bar 
1 mm. (E) Same as D, oral view, same scale. (F) Marginal bulb of medusa shown in D, note 
oblong ocelli on tentacles, scale bar 0.2 mm. (G) Oral tentacle of medusa shown in D, note long 
basal trunk, same scale as F. (H) Mature medusa, scale bar 1 mm. (I) Aboral view of manubrium 
of medusa shown in H, same scale as in H. A-C, redrawn after Edwards (1964a); D-G, after 


living medusa from Dunstaffnage; H-I, schematic reconstruction based on several preserved 
medusae. 


208 P. SCHUCHERT 


Margelis zygonema Haeckel, 1880: 635. — Edwards, 1966: 141, synonym. 

Atractylis linearis Alder, 1862b: 313, pl. 14 figs 1-3.— Edwards, 1964a: 9, synonym. — Cornelius 
& Garfath, 1980: 278. 

Bougainvillia xantha Hartlaub, 1897: 461, pl. 15 figs 2-3, pl. 16 fig. 3, pl. 16b fig. 19.— Hartlaub, 
1911: 169, synonym. 

Bougainvillia bella Hartlaub, 1897: 470, pl. 15 fig. 7. — Hartlaub, 1911: 162, synonym. 

? Bougainvillia britannica var. coeca Hartlaub, 1911:168. 

in part Bougainvillia flavida Hartlaub, 1897: 456, pl. 14 figs 1-4, 7-10 (non 5-6). — Edwards, 
1964a: 8, synonym. 


MATERIAL EXAMINED: Dunstaffnage Bay, Scotland, May 2004, several subadult medusae 
from surface plankton, examined alive, one used to extract DNA and to determine 16S sequence, 
EMBL/GenBank accession number AM183127. - BMNH 1982.11.30.1; Ireland, Lough Ine, 
Skibberean; collected July 1939. — BMNH 1955.11.23.198-231, 232-288; Great Britain, 
Plymouth; 20.04.1934. - BMNH 1956.1.1.5-7; Great Britain, Clyde; 24.05.1955; W. J. Rees 
collection. - BMNH 1955.11.23.23; Great Britain, Plymouth; 25.05.1934; figured in Russell 
1953, pl. 8 fig. 2. - BMNH 1955.11.23.1-3; Great Britain, Plymouth; 03.05.1935. - BMNH 
1954.11.13.106-114; Ireland, Valentia Island; 10.08.1899; label: Margelis bella. - ZMUC, no 
number; Germany, Helgoland; about 50 medusae, 26 May 1931, det. P. Kramp. 


DIAGNOSIS: Medusa 5-12 mm, jelly thick; no gastric peduncle, basal trunk of 
oral tentacles long, ocelli crescentic or oval, often appearing almost linear in oral view, 
situated adaxially on bases of tentacles, not on bulbs, 12-17 tentacles per bulb, oral 
tentacles branch 4-5 times; polyp stolonal or sparingly branched, long pedicels, 
medusa buds in clusters on branched stalks, young medusa with very narrow sub- 
umbrella. 


DESCRIPTION (after examined material, Russell, 1953, and Edwards, 1966): 
Hydroid colonies forming erect shoots, not much branched, branching at acute angles; 
stems arising from a large-meshed irregular network of tubular stolons; stems rela- 
tively tall; stems and branches monosiphonic; perisarc of stems wrinkled transversely 
to slight degree only, not annulated; perisarc expands around larger hydranths to form 
cup-like pseudohydrotheca; hydranths terminal on stems and branches, 6-10 tentacles 
(max. 14) in a single whorl; gonophores thick-jellied and globular at maturity, in 
clusters on fairly long branched stalks arising from stems and branches, several clusters 
on one stem may occur, buds completely invested in a delicate perisarcal sheath, which 
remains attached to the colony after rupture and the release of the medusa. 

Newly released medusa almost spherical; jelly very thick, exumbrella with 
scattered nematocysts, subumbrellar cavity very small and narrow, thin apical canal; 
one tentacle on each bulb, without ocelli, tentacles retained in subumbrella, four oral 
tentacles unbranched, velum rudimentary. The apical canal disappears rapidly during 
the further growth, the tentacles are extended towards the outside, 

Adult medusa with very thick jelly; no gastric peduncle, the jelly of the um- 
brella overhangs the margin of the subumbrellar cavity and the marginal tentacles 
emerge from channel-shaped depressions in this jelly; radial canals rather broad; bulbs 
triangular to kidney-shaped, about half as wide as intervals, but can be contracted very 
much; up to 30 tentacles on each bulb, usually 12-17, very contractile; ocelli lenticular 
or oval, often appearing almost linear in oral view, situated adaxially on bases of ten- 
tacles and not on bulbs; oral tentacles fairly long, dichotomously divided up to 4-6 
times, with long basal trunks; stomach broad-based, in form of perradial cross. Gonads 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 209 


eight adradial pads, confluent interradially, separated by perradial cleft even in fully 
mature animals. Nematocysts: microbasic euryteles and desmonemes. Colours of 
polyp white, of medusa: bulbs orange to brown-red, ocelli dark purple to black. 


VARIATION: The thickness of the jelly varies considerably, in some specimens 
the apical jelly takes up half of the total bell-height, while in others it is not much 
thicker than the lateral bell-wall. Likewise, the breadth of the radial canals, the size of 
the manubrium, and the size and shape of the marginal bulbs are very variable. 


DIMENSIONS: Stems of polyps up to 15 mm, hydranths 0.3-0.55 mm, gonophores 
0.55 mm in diameter, newly released medusa 0.52 mm wide (Edwards, 1964a). Adult 
medusa 5-8 mm, exceptionally up to 12 mm; egg size 0.14-0.15 mm, planulae 0.36 mm 
(Russell, 1953). Microbasic euryteles (10-11)x(3.5-4)um, desmonemes (6-6.5)x(3.5- 
4)um (Russell, 1938b). More data on dimensions, the development of bell size, 
tentacle numbers, and branching of the oral tentacles can be found in Russell (1953) 
and (Edwards, 1964a). 


DISTRIBUTION: North-eastern Atlantic and North Sea in coastal areas, with the 
English Channel as southern limit and Norway as northern limit. Also present west of 
Iceland (Fraser, 1953). There is also a record from Maine (USA) based on a single 
medusa (Mayer, 1910). Absent from the Baltic Sea, Mediterranean, snd Black Sea. 
There are also a records from China (Chow & Huang, 1958), Papua New Guinea 
(Bouillon, 1980), and Northern Alaska (see Vannucci & Rees, 1961). In view of its 
rather restricted distribution in the cool temperate waters of the NE Atlantic, records 
outside this regions, especially in warm or tropical waters, need reconfirmation by life- 
cycle studies as they could be due to a confusion with other, similar species. No exact 
type locality has been specified, Forbes (1841) had medusae from North of Ireland and 
East of Scotland. 


BIOLOGY: Edwards (1964a) found the polyp on shells of Aporhais pespelecani 
(L.) at depths of 60-80 m on a mud bottom. Colonies collected from end of March to 
mid May had medusa buds. Other authors found the polyp on a variety of bivalve- and 
gastropod-shells. The medusa can be found in the plankton from April to October 
(Russell, 1953; Edwards, 1964a; Ballard & Myers, 1996). The life span of the medusa 
is likely not much more than two months; at day-time, it lives usually in 20-30 m depth 
(Plymouth: Russell, 1953; Lough Hyne: Ballard & Myers, 1996). Some data indicate 
a diel vertical migration (Russell, 1953). 


REMARKS: The synonymy and taxonomic history of this species has been 
worked out by Russell (1953) and Edwards (1964a, 1966). The latter also described the 
life-cycle in detail. The polyp had been previously been known as Atractylis linearis 
Alder, 1862 and as Bougainviliia flavida Hartlaub, 1897. 

The medusa of Bougainvillea britannica can be difficult to distinguish from B. 
principis. The following differences help to differentiate them. In B. britannica the 
jelly is usually much thicker and hence the subumbrellar cavity is smaller than in B. 
principis. The basal attachment of the stomach is usually quite horizontal in B. britan- 
nica whereas in B. principis there is a tendency towards a shallow peduncle. The basal 
trunks of the branches of the oral tentacles are long in B. britannica, while they are 


210 P. SCHUCHERT 


rather short in B. principis, but contraction can obliterate the difference. In B. britan- 
nica the ocelli are oblong and they are situated on the tentacles, while they are more 
rounded in B. principis at the junctions of the tentacles with the bulb (appearing on the 
bulbs in contracted specimens). 

Bougainvillea britannica is also incorrectly listed as occurring in the 
Mediterranean (Bouillon et al., 2004). The record goes back to Thiel (1935) who listed 
it for the Black Sea based on a literature record of the polyp stage of B. muscus 
(recorded as Eudendrium ramosum). Thiel incorrectly assumed that B. britannica is the 
medusa stage of this polyp, a frequent error going back to Wright (1858, 1859a) and 
repeated by several other prominent authors (see Kramp, 1926: 42). 


Bougainvillia principis (Steenstrup, 1850) Fig. 6-7 


Margelis principis Steenstrup, 1850: 35. — Haeckel, 1879: 88, pl. 6 figs 14-16. 

? Medusa duodecilia Dalyell, 1847: 70, pl. 11 figs 11-12. — Edwards, 1966: 138, questionable 
synonym. 

? Hippocrene simplex Forbes & Goodsir, 1853: 313, pl. 10, fig. 6a-e. — Edwards, 1966: 140, 
questionable synonym. 

Bougainvillia fruticosa. - Romanes, 1876: 526. — Haeckel, 1879: 88, synonym. 

[not Bougainvillia fructicosa Allman, 1864, = B. muscus] 

Bougainvillia allmanii Romanes, 1877: 190. — Haeckel, 1879: 88, synonym. 

Nemopsis heteronema Haeckel, 1879: 93, pl. 5 figs 6-9. — Kramp, 1926: 48, synonym. 

In part Margelis principis. — Haeckel, 1879: 88, pl. 6 figs 14-16. — Kramp, 1955a: 154. 

Hippocrene aurea Linko, 1905: 216. — Hartlaub, 1911: 179. 

Bougainvillia superciliaris. — Hartlaub, 1897: 466, pl. 16a figs 5, 8, 12. 

[not Bougainvillia superciliaris (L. Agassız, 1849)] 

Bougainvillia principis. — Mayer, 1910: 160. — Hartlaub, 1911: 177, fig. 158. — Le Danois, 
1914b: 307, figs 5-6. — Kramp, 1926: 48, chart VIII. — Russell, 1953: 164, figs 76A-B, 
78B, 79B.- Kramp, 1959a: 108, fig. 88.— Vannucci & Rees, 1961: 79. — Edwards, 1966: 
129, figs 1-2. — Kramp, 1968: 32, fig. 83. - Naumov, 1969: 216, fig. 85. — Russell, 1970: 
238, figs 4s-5s. — Pages er al., 2006: 371, fig. 5B-E 

in part Bougainvillia superciliaris. — Hartlaub, 1911: 174, figs 154-156 [polyp and young 
medusae]. 


MATERIAL EXAMINED: MHNG INVE35768, Scotland, Dunstaffnage Bay, 7-13 May 2004, 
several mature medusae from surface plankton, examined alive, DNA made of one individual, 
16S sequence AM183128. - MHNG INVE48763; Norway, Raunefjord, 0-35m, June 2006, 2 
living medusae from plankton, DNA made from one specimen, 16S sequence was identical to 
AM183128.— BMNH 1955.11.23.23-33, Great Britain, Plymouth, 12-14 April 1937, few badly 
preserved mature medusae, leg. & det. F.S. Russell. - ZMUC (without number), Denmark, NW 
of Hanstholm, 200 m wire, 27.09.1930, 3 medusae, leg & det P. Kramp. — ZMUC (without 
number), Denmark, NW of Lodbjerg, 60 m wire, 27 September 1930, 3 medusae, leg. & det. P. 
Kramp. 


DrAGNosis: Medusa 7-8 mm, with 20-30 tentacles per bulb, oral tentacles 
branching 5-6 times, basal trunk of oral tentacles short, ocelli round and mostly on bulb 
and not tentacle; hydroid stolonal, sessile or with very short pedicels, small, medusa 
buds on stolons, released without peduncle, with three tentacles and three ocelli on 
each bulb. 


DESCRIPTION (polyp and newly released medusa after Edwards, 1966): Hydroid 
stolonal, arising singly at intervals from fine creeping stolons, which form a very open 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 211 


FIG. 6 


Bougainvillia principis (Steenstrup, 1850). (A) Two polyps, note that the left one has no pedi- 
cel, while the right one has an indistinct one, for dimensions see text. (B) Medusa bud issuing 
from stolon, same scale as A. (C) Newly released medusa, size about 1 mm, note the presence 
of three tentacles per bulb as well as branched oral tentacles. (D) Mature medusa in side view, 
tentacles truncated, scale bar 1 mm. (E) Same animal and scale as in D, oral view. (F) Oral ten- 
tacles of one quadrant, note the very short basal trunk, scale bar 0.5 mm. (G) Marginal bulb, the 
ocelli are at the junctions of the tentacles with the bulb, scale bar 0.5 mm. A-C, modified after 
Edwards (1966); D-E, after living medusa from Dunstaffnage; F-G, after preserved material. 


212 P. SCHUCHERT 


FIG. 7 


Bougainvillia principis (Steenstrup, 1850), photograph of a living medusa, bell height about 
lcm 

network; hydranths have very short stems or are sessile; perisarc of stolons smooth, the 
one investing lower part of hydranth often wrinkled, it thins out distally; hydranths 
spindle-shaped, hypostome short and rounded, 5-8 tentacles in a single whorl; 
gonophores globular, borne singly on very short stalks arising from the stolons. Colour: 
hypostome opaque white, gastrodermis dull pink. 

Newly released medusa deep bell-shaped; jelly thin, thickness uniform, exum- 
brella with numerous nematocysts, four moderately deep interradial longitudinal 
furrows and four slight perradial longitudinal furrows in umbrella; without peduncle; 
three tentacles and three ocelli on each bulb; manubrium tubular, with four oral 
tentacles divided dichotomously 1-2 times, trunk long. Colours: gastrodermis of 
manubrium and bulbs brick-red. 

Adult medusa wider than high or globular; jelly moderately thick, rather thicker 
above than at sides, umbrellar interradial furrows very shallow and perradial ones 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 213 


slight; with or without broad shallow peduncle; four radial canals moderately broad; 
bulbs long, epaulette-shaped, as long or longer than intervals; 20-40 tentacles per bulb; 
round adaxial ocelli at the junctions of the tentacles with the bulb (appearing on the 
bulbs in contracted specimens); oral tentacles rather short, dichotomously divided 5-8 
times, basal trunks very short or absent and thus sometimes like two independent oral 
tentacles per perradial position (Fig. 6F), tips with nematocyst clusters, but not 
swollen; stomach broad-based, in form of perradial cross; gonads in form of eight 
adradial pads, confluent interradially, clearly separated by perradial cleft even in fully 
mature animals, in subadults separated perradially by gonad-free stripe. Colours: 
centre of bulbs with a red to darkish brown or orange pigment that forms radiating 
stripes on the abaxial side of the bulb; ocelli black; gonads orange-pink. : 


DIMENSIONS: Adult medusa usually 5-8 mm wide, maximally 10-11 mm. Newly 
released medusa 1.1 mm wide and 1.3 mm high. Hydranths up to 1.1 mm high, 
diameter 0.18 mm; diameter gonophores 0.63 mm; more measurements are given in 
Edwards (1966). 


VARIATION: Edwards (1966) gives more details on the growth of the medusa and 
the variation of tentacle numbers and the branching of the oral tentacles. 


DISTRIBUTION: A northern boreal medusa occurring in coastal areas from the 
Barents Sea to the English Channel and the North Sea. Also present around Iceland, 
western Greenland and New England (Hartlaub, 1911; Kramp, 1927; Russell, 1953; 
Kramp, 1961; Zelickman, 1972; Pages et al., 2006). Chow & Huang (1958) recorded 
it in the East China Sea. The records for NE Pacific were referred to the very similar 
B. multitentaculata Foerster, 1923 by Arai & Brinckmann-Voss (1980), but later 
changed to B. principis (Cairns et al., 2002). Type locality: Sandvaag, The Faroe 
Islands. 


ADDITIONAL DATA: The ultrastructure of the ocelli is described by Singla (1974). 


BIOLOGY: The polyp was found on the surface of pieces of clinker, coal and 
stone dredged at 30 m depth. Gonophores were observed from March to May. The life 
span of the medusa is about two to three months (Edwards, 1966). The seasonal 
occurrence of the medusa has been summarized or examined by Kramp (1926), Russell 
(1953), Edwards (1966), Ballard & Myers (1996). It is a summer species and can be 
found in the plankton from April to September. In more northern localities it will 
appear later than in the south. In a sea-lough SW of Ireland, Ballard & Myers (1996, 
2000) found it usually in depths of 20-30 m. Although they could not detect any diel 
vertical migration, B. principis might migrate at other places. 


REMARKS: A distinct species, although sometimes it can be difficult to separate 
from B. britannica (see there). Its synonymy and life cycle has been well worked out 
by Edwards (1966). Bougainvillia multitentaculata Foerster, 1923 from the NE Pacific 
resembles very closely B. principis and has been confounded with this species (see 
Arai & Brinckmann-Voss, 1980). Bougainvillia multitentaculata differs from B. prin- 
cipis in having no perradial interruption of the gonads. 


214 P. SCHUCHERT 


Bougainvillia muscoides (Sars, 1846) Fig. 8 


Perigonimus muscoides M. Sars, 1846: 8, pl. 1 figs 19-21. — Jäderholm, 1909: 46, pl. 1 fig. 18. 

Bougainvillia nigritella Forbes, 1848: 63, pl. 12, fig. 2a-f. — Edwards, 1966: 139, synonym. 

Margelis nordgaardii Browne, 1903: 14, pl. 2, fig. 1, pl. 3 figs 5-6. — Mayer, 1910: 168, fig. 91. 
— Kramp & Damas, 1925: 256, figs 11-12.— Kramp, 1926: 43.— Rees, 1938: 2, synonym. 
— Kramp, 1959a: 110, fig. 94. — Kramp, 1961: 79. 

Bougainvillia obscura Bonnevie, 1898b: 7, pl. 1 fig. 4. — Rees, 1956b: 113, synonym. 

Thamnitis sp. Browne, 1905b: 758. — Edwards, 1964b: 741, synonym. 

Bougainvillia muscoides. — Rees, 1938: 2, fig. 1. — Rees, 1956b: 113. — Edwards, 1964b: 730, 
figs 5-6. life cycle. — Rasmussen, 1973: 24, fig. 8. 

Thamnostoma spec. — Russell, 1953: 150, fig. 73A-E. — Edwards, 1964b: 741, synonym. 


MATERIAL EXAMINED: MHNG INVE48760 and INVE48761; Norway, Raunefjord, 
Flesland beacon, 20-100 m, 26 June 2006, on tubes of Sabella spec., hydroid colonies with 
medusa buds, release of medusae observed, newly released medusa without ocelli, orange; DNA 
extracted, 16S sequence AM411412. - MHNG INVE48764; Norway, Fanafjord, 0-150 m, 22 
June 2006; one mature female medusa from plankton, examined alive. -— BMNH 
1965.1.14.39+62; Sweden, Kosterfjord, Sächen Reef; polyp colonies without medusa buds; 
depth 80-100 m; 23 September 1964. — BMNH 1965.1.14.74+75; polyp colony on slides; 
Sweden, Kosterfjord, Sächen Reef; 80-100 m; 23 September 1964. — BMNH 1985.9.12.12 
Norway, Bogngstrémmen; polyp colonies without medusa buds from Ascidia mentula; 14 
August 1937; coll. W. J. Rees, labelled as Perigonimus muscoides. - BMNH 1985.9.11.5; 
Norway, Bogngströmmen; polyp colonies with medusa buds; 18 August 1937; coll. W. J. Rees. 
— BMNH 1969.12.2.93; Norway, Bogngstrémmen; polyp colony on slides, with medusa buds; 6 
September 1937; coll. W. J. Rees. - BMNH 1985.9.12.10; Norway, Herdla and Brattholmen; 
medusae and polyps, medusa from plankton; August 1937; coll. W. J. Rees. -— BMNH 
1949.2.1.41; Norway, Mofjord; 1 May 1903; 3 medusae from plankton; labelled as Bougainvillia 
nordgaardi, leg. E. T: Browne. - BMNH 1949.2.1.42; Norway, Mofjord; 2 May 1903; several 
medusae from plankton; labelled as Margelis nordgaardi, leg. E. T: Browne. — ZSM, Stechow 
collection, 20000639 and 2000641, two slides as Perigonimus muscoides, loc. Capri, most like- 
ly material described in Stechow (1923a). -MHNG INVE38775; Chile, Huinay marine station, 
February 2006; several mature medusae from plankton; leg. H. Galea; 16S sequence AM411413. 


DiaGnosıs: Medusa without ocelli, 3-5 mm high, oral tentacles up to four times 
branched, 3-7 tentacles per bulb, tentacle bases swollen, stomach narrowed at base, 
gastric peduncle short. Hydroid colony erect, branched polysiphonic, with thin 
auxiliary tubes creeping over thick central tube, hydranths on short pedicels, medusa 
buds single. 


DESCRIPTION (Own observations; Kramp & Damas, 1925; Edwards, 1964b): 
Hydroid forming erect, polysiphonic stems arising at intervals from a network of 
tubular stolons; stem fairly thick and not much branched, secondary branches short, 
rather few; stem composed of a thicker central tube covered by up to seven thinner 
auxiliary tubes originating as stolons, towards distal the number of auxiliary tubes gets 
reduced and the distal-most parts are only formed by central tube; perisarc of central 
tube undulated, sometimes multilayered. Hydranths arise from central tube and 
auxiliary tubes, pedicels often very short and hydranths thus close to stem, many 
almost adnate; hydranths of very variable size, terminal ones larger than lateral ones; 
perisarc of polyp pedicel thin, wrinkled and forming cup around proximal part of 
hydranth like a short pseudohydrotheca; hydranth with 8-12 tentacles. Gonophores 
abundant, pear-shaped, arising singly on short stalks on central tube, auxiliary tubes, 
hydranth pedicels, and sometimes the stolons, frequently in rows on auxiliary tubes, 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 


N 
= 
Nn 


FIG. 8 


Bougainvillia muscoides (Sars, 1846). (A) Colony silhouette, scale bar 1 cm. (B) Part of branch 
with polyps and medusa buds, note thick central tube and thinner auxiliary tubes, scale bar 0.5 
mm. (C) Higher magnification of perisarc of central tube, scale bar 50 wm. (D) Newly released 
medusa. (E) Side view of mature medusa, tentacles truncated, scale bar 0.4 mm. (F) Oral view 
of medusa, same scale as E. (G) Tentacle bulb of juvenile medusa in adaxial view, note absence 
of ocelli and basal swelling of tentacle. A-C, E-F, after preserved material from Norway; D & G, 
modified after Edwards (1964b). 


216 P. SCHUCHERT 


covered by filmy perisarc. Colours: polyps orange or reddish, medusa buds red and 
white. 

Newly released medusa with uniformly thin jelly, bell relatively opaque, with 
four shallow perradial longitudinal furrows in exumbrella, exumbrellar nematocysts 
present; no peduncle, manubrium short; no ocelli, only one tentacle on each bulb, 
tentacle bases swollen or not; four oral tentacles unbranched. Colours: manubrium 
reddish, less brilliant than in the buds; tentacle bulbs with reddish brown pigment. 

Adult medusa about as wide as high; bell relatively angular and opaque; jelly 
moderately thick, thicker apically than at sides; exumbrella with four interradial and 
four perradial furrows; rather shallow gastric peduncle present; radial canals 
moderately wide; bulbs rather large, rounded triangular; 3-7 tentacles on each bulb, 
tentacles swollen at base, distal end also thickened, bulbs without ocelli; oral tentacles 
divided dichotomously 4-7 times ending in weak capitations, basal trunks of oral 
tentacles short or of moderate length (1/5 or more of total length). Stomach relative 
small, spanning about !/ of subumbrellar cavity, ovoid, narrowed at base; gonads form 
bulging interradial pads on stomach wall. Colour of manubrium and bulbs red-orange. 


DIMENSIONS (Kramp & Damas, 1925; Edwards, 1964b; Rasmussen, 1973; own 
observations): Mature polyp colonies up to 2-7 cm, central tube 0.25-0.35 mm 
diameter, auxiliary tubes 0.08-0.1 mm in diameter; hydranths up to 1 mm long and 
0.3 mm wide. Medusa buds 0.4 mm high, diameter 0.3 mm. Newly released medusa 
0.55-0.7 mm. Height of adult medusa 3-5 mm. Egg size about 0.2 mm (Norwegian 
medusae). 


VARIATION: In the north of the British Isles, the medusae are about 3 mm in 
height and have up to three tentacles per bulb. In Norway, the medusae reach sizes of 
4-5 mm and can have up to seven tentacles per bulb. In preserved material, the basal 
swelling of the tentacles is not always present. 


BıoLoGY: The polyp is found on Tubularia indivisa, Ascidia mentula, Mytilus, 
Sabella pavonia, Sertularella gayi, in depths of 4-200 m, mostly around 40 m 
(Edwards, 1964b; Rasmussen, 1973). Rasmussen (1973, Denmark) observed newly 
settled hydroids in October-November. Medusa buds in Norwegian, Scottish and 
Danish colonies were observed in June and August (Edwards, 1964b; Christiansen; 
1972; Rasmussen, 1973; own data). The medusa is found in the plankton from June to 
November, mostly in deeper waters (around 100 m) but sometimes also at the surface 
(Kramp & Damas 1925; Edwards, 1964b; own data). The colonies are dioecious. 


DISTRIBUTION: Reliable records of both polyp and medusa phase are restricted 
to a relatively narrow region reaching from Trondheimsfjord in Norway south to 
Scotland (Edwards, 1964b). Also present along the west coast of Sweden (Jäderholm, 
1909; Allwein, 1968; Rees & Rowe, 1969, Christiansen, 1972) and Denmark 
(Rasmussen, 1973). Records from Chile and British Columbia (NE Pacific) were 
thought to be erroneous (Kramp, 1952; Arai & Brinckmann-Voss, 1980). However, 
recently Dr Horia Galea (MHNG INVE38775) has found and documented medusae in 
a Chilean Fjord that are indistinguishable from B. muscoides. A sequence comparison 
of about 590 base-pairs of the 16S gene of the Chilean specimens with polyp material 
from Norway gave only a difference of a single base pair, suggesting that they are 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 217 


conspecific (see also under B. muscus). There are also records from the Mediterranean 
(Stechow, 1923a), Red Sea (Schmidt, 1973a, Papua New Guinea (Bouillon, 1980) and 
New Zealand (Bouillon, 1995). The records from warm or tropical regions are pro- 
bably misidentifications of closely related species (see also Schuchert, 1996). In the 
Mediterranean, the medusa has never been seen, and the presence of the polyp stage in 
the Mediterranean is not certain (see below). Type locality: Mangerfjord, Norway, 
polyp on Ascidia mentula and Tubularia indivisa, 36-55 m. 


REMARKS: Stechow (1923a: 75) reported an infertile colony of Bougainvillia 
muscoides from near Capri in 200 m depth, but he admitted that it was only tentatively 
identified. Although Stechow’s material appears indistinguishable from Atlantic 
material (material re-examined for this study), the presence of B. muscoides in the 
Mediterranan is not certain as long the medusa has not been found or cultivated. 
Stechow also noted the similarity of his material to Bimeria biscayana Browne, 1907 
(here referred to Amphinema bicayana), and it is possible that it belongs to this species 

Due to the absence of ocelli, the medusa of B. muscoides is easily distin- 
guishable from all other Bougainvillia species of the region (note that in formalin fixed 
specimens the pigment of ocelli fades rapidly). However, Goy et al. (1991) described 
a Bougainvillia medusa originating from Lebanon in the eastern Mediterranean. They 
assigned it to Bougainvillia aurantiaca Bouillon, 1980, a species originally described 
from Papua New Guinea. The Lebanese medusa closely resembles B. muscoides, but 
it is distinctly smaller (1.5 mm), has only two tentacles per bulb and its oral tentacles 
branch only two times. More life cycle information is needed for the Lebanese popu- 
lations in order to evaluate their status and to provide more diagnostic characters to 
distinguish it more reliably from B. muscoides. 


Bougainvillia pyramidata (Forbes & Goodsir, 1853) Fig. 9 


Hippocrene pyramidata Forbes & Goodsir, 1853: 312, pl. 10 fig. 4. 

? Medusa ocilia Dalyell, 1847: 64, pl. 11 figs 1-10. — Edwards, 1966: 138, questionable 
synonym. 

in part Bougainvillia flavida Hartlaub, 1897: 456, pl. 14 fig. 6 [male medusa only]. — Hartlaub, 
1911: fig. 168. — Edwards, 1964b: 734, synonym. 

Bougainvillia pyramidata. — Mayer, 1910: 168. — Hartlaub, 1911: 180, fig. 159. — Russell, 1953: 
167, fig. 82A-C. — Kramp, 1959a: 108, fig. 86. — Kramp, 1961. — Edwards, 1964b: 725, 
figs 1-4. — Russell, 1970: 236, fig. 6s. 


MATERIAL EXAMINED: BMNH 1971.5.11.10, Great Britain, Isle of Cumbrae, Millport, 36 
m, 24 September 1970, polyp on Tubularia and reared young medusae, det. C. Edwards. — 
BMNH 1967.12.1.7, Great Britain, Little Cumbrae, off Lighthouse, 82 m, 08 July 1966, polyp 
on Tubularia, hydranths lost, coll. W. J Rees. - BMNH 1966.1.4.13+24, Sweden, Gullmarfjord, 
50 m, 13 October 1964, polyp on ascidian, coll. W. J. Rees. - BMNH 1966.1.4.38, Sweden, 
Gullmarfjord, Skär, 80 m 3 October 1964, polyp colony, det. W. J. Rees. - BMNH 1985.9.10.23, 
Great Britain, Millport, Keppel Pier, one medusa, 30 September 1940, det. W. J. Rees. - BMNH 
1985.9.4.12, Ireland, Kerry county, Valentia Island, 27 July to 5 August 1938, 7 medusae, coll. 
W. J. Rees. 


DrAGNosis: Medusa with cone-shaped gastric peduncle, 4-9 tentacles per bulb, 
oral tentacles 3-4 times branched, with moderately long basal trunk; gonads adradial 
on perradial outgrowths of stomach; ocelli round on bulbs. Hydroid erect, branching, 
polysiphonic, with distinct and straight main axis, medusa buds in groups. 


218 P. SCHUCHERT 


Bougainvillia pyramidata (Forbes & Goodsir, 1853). (A) Colony, size about 4 cm. (B) Hydranth 
with medusa buds, for sizes see text. (C) Part of polysiphonic stem with cluster of medusa buds. 
(D) Newly released medusa, note absence of peduncle, unbranched oral tentacles, two tentacles 
per bulb, scale bar 0.3 mm. (E) Manubrium of juvenile medusa, left aboral view, right lateral 
view. (G) Manubrium of mature, cultivated medusa, left aboral view, right side-view. (H) Mature 
medusa from plankton, side view, scale bar 0.5 mm. (I) Same as in H, oral view. (J) Adaxial view 
of tentacle bulb. A-G, modified from Edwards (1964b); H-I, schematic composite picture from 
several preserved medusae; J, after Russell (1953). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 219 


DESCRIPTION (after own observations; Russell, 1953; Edwards, 1964b): Hydroid 
forming erect stems with a distinct, straight, central primary axis, primary, secondary 
and tertiary branches present. Stolons creeping, tubular, branching and forming mesh- 
work, sometimes polysiphonic, stem and primary branches polysiphonic, branches 
straight and regular; perisarc corrugated at bases of branches, thins out over hydranths 
to pseudohydrotheca; hydranths terminal on the branches and stem, cylindrical and 
quite long, with 6 to 13 tentacles in one whorl; medusa buds egg-shaped, on distinct 
stalks, arising in clusters on the branches just below the hydranths, on stem or on short 
stems arising from the stolons, stalks of medusa buds can be branched. Colours: 
hypostome white, gastrodermis pink. 

Medusa at release subglobular; jelly uniformly thin, exumbrella with numerous 
scattered nematocysts and with four shallow interradial longitudinal furrows; no 
peduncle; two tentacles and two ocelli on each bulb; four oral tentacles unbranched. 

Adult medusa subglobular; jelly thick, especially apically; manubrium on a 
conical peduncle, height of peduncle variable; radial canals narrow; bulbs oval, much 
smaller than space between them; 7-9 tentacles per bulb; ocelli round, situated 
adaxially on bulbs at bases of tentacles; oral tentacles divided dichotomously 3-4 
times. Perradial lobes of the stomach extend along radial canals on peduncle and 
gonads lie in the walls of these lobes and adjacent basal part of stomach walls, 
appearing V-shaped in side-view and cross-shaped in axial view. Colours: marginal 
bulbs reddish brown or yellow; stomach yellowish brown; ocelli black. 


VARIATION: The development of the medusa is described in Russell (1953) and 
Edwards (1964b). The peduncle is only fully formed in mature medusae. 


DIMENSIONS: Hydroid colony up to 4 cm high, hydranths up to 0.66 mm long 
and 0.1 mm wide, gonophores 0.45 mm in diameter (after Edwards, 1964b, for more 
measurements see this publication). Medusa at release about 0.72 mm wide and 
0.63 mm high. Adult medusa 2-5 mm wide and high, exceptionally up to 8 mm. 


DISTRIBUTION: Bougainvillia pyramidata appears to be restricted to the north- 
eastern Atlantic along the western coasts of Scotland, the Irish Sea, the western coast 
of Ireland, and the western coast of Sweden (Russell, 1953; Rees & Rowe, 1969; Boyd 
et al., 1973; Ballard & Myers, 1996). Type locality: Loch Laigh, Mull, Scotland. 


BioLoGY: The polyp was found in depths of 30-80 m on stems of Tubularia 
indivisa and Virgularia mirabilis and on tests of Ascidiella aspersa. Along the Scottish 
coast, the medusa is present in the plankton (Scotland) from June to begin of 
November, with a clear maximum towards the end of August. The medusa seems to be 
associated with moderately deep waters, it is purely neritc, it has been taken commonly 
near the surface and there is no evidence that it lives in deep layers (Edwards, 1964b). 
Ballard and Myers (1996, 2000) observed it in a lough in SW Ireland in low numbers 
from April to November, depths of 10-20 m. No diel vertical migration was observed. 


REMARKS: The hydroid of Bougainvillia pyramidata is prone to be mistaken for 
B. muscus (Edwards, 1964b; Rees & Rowe, 1969). According to Edwards (1964b) 
Bougainvillia pyramidata is distinguished by its erectness and straightness of the 
stems, the straightness of the branches and their regularity of arrangement. It does not 


220 P. SCHUCHERT 


appear tree-like as in large B. muscus colonies, but has a conspicuous, more or less 
straight central main stem running to the top of the colony. Branched pedicels of the 
medusa buds (Fig. 9B-C) are so far only known in B. pyramidata. Although they may 
not always be present, they are likely a diagnostic difference. The tentacle number may 
also be lower with a maximum of 13 (B. muscus 18), and its occurrence seems to be 
restricted in depths below 30 m, while B. muscus can also be found immediately below 
the water surface. Whatsoever, the morphologies of both species — both medusa and 
polyp - intergrade and they are sometimes not objectively separable. Some Norwegian 
colonies (MHNG INVE48746) were initially taken for B. pyramidata as they differed 
markedly from other nearby colonies of typical B. muscus (comp. Figs. 2 and 3). The 
colonies had a straight central axis and clustered medusa buds. They also differed in 
colour from the more typical B. muscus colony. However, the rearing of the medusa 
and the 16S data proved that these colonies were B. muscus. The hydroid stage of both 
species can thus sometimes not be distinguished morphologically, but the medusa 
clearly separates the two species. 


Bougainvillia superciliaris (L. Agassiz, 1849) Fig. 10 


Hippocrene superciliaris L. Agassiz, 1849: 273, pl. 1-3. 

? Hippocrene Bougainvillii. — Brandt, 1835: 29 [pagination of reprint]. — Brandt, 1838: 393, 
pl. 20. 

[not Cyanea Bougainvillii Lesson, 1830 = B. macloviana] 

? Bougainvillia Mertensii L. Agassiz, 1862: 344, new name for Hippocrene Bougainvillei 
Brandt. 

Not Bougainvillia Mertensii. — A. Agassiz, 1865: 152, hydroid. — Allman, 1872: 318. 

Bougainvillia paradoxa Mereschkowsky, 1879: 177, pl. 20 figs 1-5. — Hartlaub, 1911: 176, 
synonym. 

in part Bougainvillia superciliaris. — Mayer, 1910: 162, figs 87-88, 162, pl 17 fig. 1.— Hartlaub, 
1897: 466, pl. 16a figs 5, 8, 12. — Hartlaub, 1911: 171, figs 153 [not polyp and young 
medusae]. 

Bougainvillia superciliaris. — Bigelow, 1909: 305, pl. 31 fig. 2.— Russell, 1953: 169, figs. 83-85. 
— Kramp, 1959a: 108, fig. 87. — Uchida & Nagao, 1960: 249, figs 1-4. — Vannucci & 
Rees, 1961: 85.— Kramp, 1961: 82.— Werner, 1961: 206, figs 1-5. — Edwards, 1966: 148. 
— Kramp, 1968: 32, fig. 80. — Russell, 1970: 236, 243. — Arai & Brinckmann-Voss, 1980: 
SS oreo. 

Not Bougainvillia superciliaris. - Naumov, 1969: 215, figs 83-84. 


MATERIAL EXAMINED: IRSNB, collection IG 27838, collected by J. Bouillon 1970, 
Atlantic Ocean, 4 females and one male. — BMNH 1970.1.4-7 and 12-15, Great Britain, Wales, 
Dale Fort Field Centre, 25-26 March 1963, several medusae collected W. J. Rees. - BMNH 
1931.12.21.16-19, Canada, Hudson Strait, Port Burnwell, collected 1931, medusae. — ZMUC, 
Denmark, Aggersund, 6 May 1929, 6 medusae. - ZMUV, Denmark, Hvalpsund, 21 m depth, 22 
April 1924, 2 medusae. 


DraGNOSIS: Medusa jelly moderately thick, with distinct gastric peduncle, 11-15 
tentacles per bulb, oral tentacles branched 4-5 times, basal trunk very short; crescent- 
shaped ocelli on base of tentacles (not on bulbs). Hydroids small, sessile, medusa buds 
on stolons, peduncle visible in buds, more than two ocelli and tentacles per bulb. 


DESCRIPTION (after Russell, 1953; Werner, 1961; own data): Polyps arising from 
a network of creeping tubular stolons; hydranth small, without distinct pedicel hence 
sessile, body covered by very thin, tightly adhering, membranous perisarc (pseudo- 
hydrotheca), expanded polyp fusiform to cylindrical, rounded hypostome, 7-13 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE DA 


Fic. 10 


Bougainvillia superciliaris (L. Agassiz, 1849). (A) Polyp stage, scale bar 0.4 mm. (B) Advanced 
medusa bud, same scale as A. (C) Newly released medusa, note presence of peduncle, scale bar 
1 mm. (D) Mature medusa, scale bar 1 mm. (E) Marginal bulb seen from oral side, note position 
of crescent shaped ocelli on tentacle bases, scale bar 0.4 mm. A-C, modified after Werner (1961); 
D-E, after preserved material. 


tentacles in one whorl. Gonophores arise singly on short stalks from the stolons, egg- 
or pear-shaped, larger than polyps. Nematocysts: desmonemes and microbasic 
euryteles. 

Newly released medusa bell-shaped; jelly of moderate thickness, thicker api- 
cally than at sides; usually with well developed peduncle; four rather shallow inter- 
radial longitudinal furrows in umbrella, exumbrella with numerous nematocysts in 
groups of 2-4; 5-9 tentacles and about 5 ocelli to each bulb, ocelli on bases of tentacles; 
four oral tentacles divided dichotomously 2-3 times. 


DD?) P. SCHUCHERT 


Adult medusa almost globular; or rather higher than wide; jelly thick, thicker at 
top than at sides; interradial furrows variably developed, umbrella cross-section rather 
quadrangular; well-developed gastric peduncle; radial canals relatively broad; bulbs 
triangular, less wide than intervals; 11-15 (exceptionally up to 22) tentacles per bulb, 
ocelli large, crescentic-round, adaxial, on bases of tentacles; oral tentacles divided 
dichotomously 4-5 times (exceptionally up to 6-8 times), basal trunks short, distal ends 
beset with nematocyst but not distinctly swollen; stomach broad-based, in form of per- 
radial cross; gonads interradial, in mature specimens covering the whole surface of the 
stomach, but separated perradially; eggs without nematocysts, most planulae develop 
enveloped in capsules and attached to gonad by a short stalk (larviparous). Colour of 
stomach reddish brown, marginal bulbs chestnut brown, ocelli black. Nematocysts: 
desmonemes and microbasic euryteles of two size classes. 


VARIATION: The presence of a gastric peduncle in the newly released medusa is 
a characteristic trait of this species, but it can be absent in a minority of individuals, 
mostly due to pathological development (Werner, 1961). The Pacific medusae are 
larger than the Atlantic ones (up to 12 mm; Uchida & Nagao, 1960). 


DIMENSIONS: Polyp stage (Werner, 1961) height up to 1.0 mm, usually 0.5-0.7 
mm; gonophores up to 2.2 mm high, 1.4 mm wide. Newly liberated medusa (Werner, 
1961) 1.9-2.5 mm wide, 1.6-2.5 mm high. Mature medusa bell height 4-9 mm. See 
Werner (1961) for the sizes of nematocysts of hydroid and medusae. 


ADDITIONAL DATA: Linko (1900) examined the histological structure of the 
ocelli. Gerd (1892) describes the early embryology. The diploid chromosome number 
is 30 (Werner, 1961). 


DISTRIBUTION: An Arctic species penetrating into colder boreal regions. 
Originally described from Massachusetts, also present along northern New England, 
Canada, and Western Greenland. In Europe resident on the coasts of Iceland, 
Spitsbergen, Norway, Denmark, Germany, and Holland. In British waters the medusa 
has been reported on the North Sea coasts. Further occurrences are known from the 
Bering Sea, Hokkaido, Yellow Sea, Aleutian Islands, British Columbia, perhaps as far 
south as San Francisco (for references see Hartlaub, 1911; Kramp, 1926; and Arai & 
Brinckmann-Voss, 1980). 


BIOLOGY: In Arctic waters, the medusa of Bougainvillia superciliaris is present 
during summer from May to July or September, in the southern parts of its distribution 
(North Sea, south of Cape Cod) it is only found in early spring, mainly from February 
to April (Kramp, 1926; but see Zelickman, 1972). 

Werner (1961) examined in detail the ecology and development of 
Bougainvillia superciliaris for the population of the North Sea. In culture, the polyp is 
eurytherm (2-15°C) and can even be maintained at room temperature, but medusae 
budding takes place only at low temperatures (5-7 °C). In contradistinction to the 
polyp, the medusa is stenotherm and it dies at temperature above 11-14°C. It takes very 
long for the medusa buds to develop (8 weeks), but the liberated medusa matures 
within four weeks. Females are able to produce eggs in several rounds. The fertilized 
eggs remain attached to the stomach wall and develop there, although some may detach 
and develop in the subumbrella. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 223 


The medusa uses its oral tentacles to capture herring larvae, euphausid nauplii 
and polychaete trochophores in preference to various other crustaceans (see refs in Arai 
& Brinckmann-Voss, 1980). 

Nagao (1964) found the hydroid in nature growing on a shell of a gastropod 
belonging to the genus Neptunea. Orlov (1995) examined the settling behaviour of the 
larvae. 


REMARKS: This species has a complicated taxonomic history partially described 
by Edwards (1966: 136). Also Vannucci & Rees (1961) provide a synonymy. Some 
comments concerning Bougainvillia mertensii L. Agassiz, 1862 are nevertheless 
needed. Brandt (1835, as Hippocrene Bugainvillii) attributed a medusa from the Bering 
Sea to Cyanea Bougainvillii Lesson, 1830, which he supplemented later (1838) by a 
figure. However, Cyanea Bougainvillii Lesson, 1830 is as synonym of Bougainvillia 
macloviana Lesson, 1830. As Brandt’s medusa is evidently not referable to B. maclo- 
viana, L. Agassiz (1862) gave it the new name Bougainvillia mertensii. Haeckel (1879) 
indicated the close similarity of Bougainvillia mertensii to B. superciliaris and 
Vannucci & Rees (1961) listed it as a questionable synonym of B. superciliaris. This 
view is also adopted here. 

The knowledge of the life-cycle of B. superciliaris has also an intricate history. 
Agassiz (1865) associated an erect, branched, 5 cm high Bougainvillia colony from 
Massachusetts to this species. He observed the release of the medusae and followed 
their development. Of the obtained medusa he says (Agassiz, 1865: 155), «it assumes 
all the principal features of the adult, as in Fig. 232, with the exception of the simpler 
character of the tentacles of the actinostome». In other words, the oral tentacles did not 
branch as much as in the medusae found in the plankton. Although Agassiz’s account 
appears plausible, Hartlaub (1911) doubted his results and attributed Agassiz’s hydroid 
to B. muscus. He did so because he believed that he had found the true polyp of B. 
superciliaris, a colony of sessile hydranths found in an aquarium of the Helgoland 
station. Hartlaub also claimed to have followed the development of the medusa until 
its identity was evident. Others continued to iterate Agassiz’s version (e. g. Mayer, 
1910; Fraser, 1937, 1944; Berrill, 1948; Vannucci & Rees, 1961). The life cycle was 
then re-investigated in detail by Uchida & Nagao (1960) for the Japanese population 
and by Werner (1961) for European animals. Both studies started with planulae 
released by the medusa stage and found a sessile hydroid, but did not discuss the 
discrepancy of their and Agassiz’s observations. There were also some discrepancies 
in Werner’s and Hartlaub’s observations, which were resolved by Edwards who real- 
ized that Hartlaub had in fact found the hydroid of B. principis and not B. superciliaris. 

Unfortunately, Agassiz did not depict the medusae he obtained from his rearing 
experiments, so we are unable to decide how reliable his identifications are. Although 
it is more probable that Agassiz indeed mistook another Bougainvillia hydroid for the 
one of B. superciliaris (unlikely B. muscus as the medusae had at least seven tentacles 
per bulb), there remains the possibility that B. superciliaris as currently perceived is 
composed of at least two species differing in their polyp stage only. New life-cycle 
studies on B. superciliaris from Massachusetts should be made to resolve this. 


224 P. SCHUCHERT 


Bougainvillia macloviana Lesson, 1830 Fig. 11 


Bougainvillia macloviana Lesson, 1830: 118, pl. 14 fig. 3.— Mayer, 1910: 160.- Hartlaub, 1911: 
156, fig. 139. — Künne, 1933: 249, fig. 1. — Browne & Kramp, 1939: 284, pl. 14 fig. 6, 
pl 15 figs 7-14. — Russell, 1953: 173, figs 86-88B. — Kramp, 1959a: 108, fig. 85. — 
Kramp, 1961: 78. — Vannucci & Rees, 1961: 69. — Edwards, 1966: 147. — Kramp, 1968: 
32, fig. 78. — Millard, 1975: 96, fig. 33A-C. — O’Sullivan, 1982: 38, fig. 16, map 15. — 
Pages et al., 1992: 3, fig. 1.— Schuchert, 1996: 30, fig. 14a-c. 

Cyanea Bougainvillii Lesson, 1830: 118, pl. 14 fig. 3, simultaneous synonym. 

Hippocrene macloviana. — Haeckel, 1879: 90, pl. 5 figs 1-2. 

Perigonimus maclovianus. — Vanhöffen, 1910: 284, fig. 10a-d. 


MATERIAL EXAMINED: BMNH 1939.6.8.1009-1025, Falkland Islands, collected 1898, 
young, mature medusae, collection Vallentin, leg. E. T. Browne. — See also Schuchert (1996). 


DIAGNosIs: Medusa up to 8-15 mm, jelly moderately thick, large gastric pe- 
duncle present, bell margin quadrangular, 30-65 tentacles per bulb, ocelli on marginal 
bulbs, oral tentacles branching 5-7 times, basal trunk short; stomach forms perradial 
lobes extending upwards onto peduncle and bearing the gonads. Hydroid small, 
stolonal, medusa buds singly on stolons and hydranth pedicels. 


DESCRIPTION (Vanhöffen 1910, Browne & Kramp, 1939; Edwards, 1966; and 
own observations): Hydroid forming stolonal colonies, hydranths arising from tips of 
creeping tubular stolons which form an open network; perisarc much wrinkled, not 
annulated, extending as thin membrane over body of hydranth; up to 16 tentacles; 
medusa-buds pear-shaped, borne singly on short annulated stalks arising from stolons 
and pedicels. 

Newly released medusa bell-shaped; jelly very thin, exumbrella even, but early 
in development deep interradial furrows and shallow adradial ones appear; no 
peduncle; bulbs globular, each with 2-5 tentacles and 2-3 ocelli; ocelli round, on bulbs 
at bases of tentacles; four oral tentacles unbranched or with one division. 

Adult medusa with bell as high as wide or slightly higher than wide, with 
quadrangular cross-section; jelly fairly thick, thickened at apex; deep interradial longi- 
tudinal furrows in umbrella, bell-margin with perradial lobes; large conical peduncle 
that can span 1/2 to 2/3 of the subumbrella. Radial canals moderately broad; tentacle 
bulbs crescentic or V-shaped, about half as wide as intervals; about 30 tentacles per 
bulb in European animals (up to 65, usually 40-55 in Southern Ocean), when more than 
18-20 tentacles then in a double row, sometimes three rows, young and old tentacles 
mixed; ocelli large, round or crescentic, on bulbs near bases of tentacles. Manubrium 
short; conical, with four long narrow perradial lobes running along peduncle; oral 
tentacles divided 5-7 times and with up to 80 ends ending in only slightly swollen 
nematocyst clusters, branching not always dichotomous, basal trunks very short. 
Gonads pad-like on adradii of stomach and along its perradial lobes, separated perra- 
dially and usually also interradially. Eggs covered with euryteles. Nematocysts: 
microbasic euryteles, on tentacles and eggs; desmonemes. Colours: stomach, gonads 
and marginal bulbs bright red, reddish brown, or orange-rose, ocelli red or black. 


DIMENSIONS: Hydroid up to several millimetres high, medusa buds reach 1 mm 
in diameter. Newly released medusa 0.5-0.75 mm high. Gonad maturation starts at a 
size of 2-4 mm (Browne & Kramp, 1939). Adult medusa up to 8.5 mm in the NE 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 225 


Fic. 11 


Bougainvillia macloviana Lesson, 1830. (A) Mature, but not fully grown medusa, scale bar 
2 mm. (B) Oral tentacle of fully grown medusa. (C) Hydroid with medusa bud. (D) Very young 
medusa, height about 0.6 mm. A, after preserved material from the Falkland Islands; B & D; 
modified after Browne & Kramp (1939); C, modified after Vanhöffen (1910). 


Atlantic, 15 mm in Southern Ocean. Nematocysts (Schuchert, 1996): microbasic 
euryteles (7-8)x(3-4)um; desmonemes, (6-7)x(4-4.5)um. 


DISTRIBUTION: Originally a species of the Southern Ocean, it has been known 
since 1895 to be resident in the North Sea and is believed to have been introduced there 
in the hydroid stage on ships. It is rare in European waters, but in its native region it 
can reach very high densities. The medusa was also found off the Isle of Cumbrae, 
Firth of Clyde, and the Mull of Kintyre (Edwards, 1966). The polyp has not been 


226 P. SCHUCHERT 


detected in Europe, it is only known from the Kerguelen Islands and South Africa 
(Millard, 1975). The medusa is also present in the coastal waters of the Kerguelen 
Islands (Vanhöffen, 1910); South Africa and Mozambique (Millard, 1975); Benguela 
Current (Pages er al., 1992); southern tip of New Zealand, Campell and Auckland 
Islands (Schuchert, 1996). Type locality: Soledad Bay, Falkland Islands, South 
Atlantic. 


BIOLOGY: A coastal species. In the South Atlantic, mature medusae are present 
from November to March (austral summer, Browne & Kramp, 1939). In the North Sea, 
they occur from April to June (Russell, 1953). Buecher & Gibbons (2003) examined 
the dial vertical migration and found that Bougainvillia macloviana was largely 
confined to surface waters and did not appear to display any vertical migration. 


OTHER DATA: The development of the medusa is described by Browne & Kramp 
(1939) and Russell (1953). Contrary to most other Bougainvillia species which add 
tentacles only at both sides of the bulbs, in B. macloviana young tentacles also develop 
among the older ones, causing an overcrowding which results in a displacement of the 
tentacles, so that they become arranged in two rows, both of which contain old and new 
tentacles (Browne & Kramp, 1939). According to Künne (1933), the North Sea 
specimens have black ocelli black, orange-rose gonads and manubrium, and a rusty 
brown zig-zag band on each bulb. 


REMARKS: In the present fauna, Bougainvillia macloviana medusae can poten- 
tially only be confounded with B. pyramidata, but mature animals differ considerably 
in size and the number of tentacles per bulb. 

There has been some confusion on the date and name of the first description of 
this species, because Lesson (1830) used the names Cyanea Bougainvillii as heading 
for a species, which he considered incorrect and he named it on the same page 
Bougainvillia macloviana. As explained by Calder (1988), both names are simulta- 
neous synonyms and later Lesson (1836), acting as first reviser, chose B. macloviana 
as name. 

Vanhöffen (1910) observed mature medusae of B. macloviana in high densities 
while his ship was in the Observatory Bay of Kerguelen Island. Later he found that 
polyps had settled on the ship hull, which he then attributed to B. macloviana. 
Although there remains some uncertainty because a direct link of the adult medusae 
and the observed was not established, it is very probable that Vanhöffen’s conclusions 
were correct. 


Bougainvillia platygaster (Haeckel, 1879) Fig. 12 


Hippocrene platygaster Haeckel, 1879: 91. 

Bougainvillia platygaster. — Kramp, 1957a: 9, text. fig. 1, pl. 3 figs 1-6. — Kramp, 1959a: 109, 
fig. 89. — Kramp, 1961: 80. — Vannucci & Rees, 1961: 78. — Kramp, 1968: 34, fig. 86. — 
Winkler, 1982: 30, fig. 3.— Bleeker & van der Spoel, 1988: 230, figs 8-9. — Pages et al., 
1992: 4, fig. 2. — Bouillon, 1995: 227, fig. 1.— Schuchert, 1996: 33, fig 16. — Bouillon er 
al., 2004: 44, fig. 26D. — Pages er al., 2006: 372, fig. SF. 

not Bougainvillia platygaster. — Goy et al., 1991: 108, fig. 20. 


MATERIAL EXAMINED: ZMUC, Neotype designated by Kramp (1957a), det. E. Haeckel, 
25.067°S 27.433°W, off Brazil, collected 1869, 1 medusa, bell torn. - ZMUC, Dana station 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 227 


Fic. 12 


Bougainvillia platygaster (Haeckel, 1879). (A) Medusa, the widely open mouth is likely a 
fixation artefact, scale bar 2 mm. (B) Stomach seen from aboral, note small medusa buds, scale 
bar 1 mm. (C) Immature specimen, aboral view of stomach with medusa buds and polyp 
formations. A, modified from Schuchert (1996); B, after preserved material from W Atlantic; C, 
from Kramp (1959a). 


3547, 11.000°N 77.480°W, western Atlantic, 0-300 m, collected 24 August 1927, 1 medusa. — 
ZMUC, Dana station 3544, 18.417°N 56.050°W, western Atlantic, 14 August 1928, 1 medusa. 
ZMUC, Dana station 3543, 21.833°N 50.200°W, western Atlantic, 12 August 1928, 1 medusa. — 
BMNH 1957.12.29.148-151, Discovery station 680, 22.600°S 30.025°W, tropical western 
Atlantic, 0-260 m, 30 April 1931 several medusae, material mentioned in Kramp (1957). 


DrAGNosis: Medusa 4-12 mm, globular, thick jelly, no peduncle, manubrium 
flat, with medusa buds, oral tentacles divided up to six times, basal trunk absent, up to 
14 tentacles per marginal bulb, with ocelli. 


DESCRIPTION: Medusa with globular umbrella, slightly flattened apex, jelly very 
thick, subumbrella cylindrical, the jelly of the umbrella overhangs the margin of the 
subumbrellar cavity and the marginal tentacles emerge from channel-shaped 
depressions in this jelly; without gastric peduncle. Radial canals rather thin, ending in 
small but broad triangular marginal bulbs with 7 - 14 short tentacles, ocelli crescent- 
shaped with concavity turned outwards. Manubrium very broad and flat, in younger 
specimens more cross-shaped, in older ones more quadrangular, in preserved 
specimens mouth often widely gaping and quadratic. Gonads four interradial flat pads, 


228 P. SCHUCHERT 


not divided interradially. Oral tentacles arising rather close to radial canals and 
branching dichotomously up to six times immediately from their point of origin, thus 
no basal trunk present, each distal end of oral tentacles with a nematocyst cluster. 
Immature animals regularly with medusa buds arising from the adradial stomach walls; 
medusae may be budded in clusters directly from manubrium or from polypoid 
structures, polyps sometimes also with stolons. 

Hydroid only known from vegetative buds on the corners of the medusa 
manubrium, with, short pedicel, 10-12 filiform tentacles, medusa buds on stalks 
immediately below the hydranths or on pedicels from stolons. 


DIMENSIONS: Medusa normally 4-7 mm, maximally 12 mm. 


OTHER DATA: The medusa- and polyp-budding of the medusa is described in 
detail by Kramp (1957a). 


BioLOGY: The depth distribution of the medusa in the North Atlantic is spanning 
the upper 300 m, with concentrations in the 100-200 m interval during daytime and 
0-50 at night, isolated specimens at depths greater than 500 m (Pagès et al., 1992). 
Diurnal vertical migration seems to take place (Winkler, 1982; Bleeker & van der 
Spoel, 1988). The medusa is often parasitized by the narcomedusa Pegantha triloba 
Haeckel, 1879 (Winkler, 1982). 


DISTRIBUTION: Widely distributed in tropical and subtropical waters of the 
Atlantic and Indian Oceans: Canary Islands, Cape Verde Islands, western Medi- 
terranean, Brazil, Sargasso Sea, Benguela Current, South Africa, Bismarck Sea, 
Malayan Archipelago, Fiji Islands, New Zealand. (Pages et al. 1992; Schuchert, 1996; 
Bouillon et al., 2004). It is clearly a species restricted to warmer waters and thus it does 
normally not occur north of the 40° latitude. An exception seems to be the find in the 
Gulf of Maine by Pages er al. (2006). Type locality: Island of Trinidade, Brazil 
(Neotype designated by Kramp, 1957a). 


REMARKS: Although characteristic and regularly encountered in preserved 
material of this species, the widely open mouth is certainly a fixation artefact (Pagès et 
al., 2006). I concur with Bouillon er al. (2004) that the record of B. platygaster for 
Lebanese waters by Goy et al. (1991) is a misidentification. Bouillon et al. (2004) refer 
this record to Bougainvillia niobe Mayer, 1894. The Lebanese record is based on a 
single specimen without tentacles and ocelli. The paucity of the material and dis- 
crepancies in the shape of the manubrium makes it preferable not to assign this material 
to a nominal species. More and better specimens are needed to confirm the presence of 
B. niobe for the Mediterranean. Bougainvillia niobe and B. platygaster both produce 
medusa buds on the manubrium, but B. niobe can easily be distinguished as it has a 
flask-shaped manubrium, its buds not in groups, the oral tentacles have a long trunk, 
there are only about eight tentacles per bulb, and sexually mature animals have eight 
adradial gonads (Kramp, 1959a: 12). 


Bougainvillia aurantiaca Bouillon, 1980 


Bougainvillia aurantiaca Bouillon, 1980: 309, fig. 1. — ? Goy et al., 1991: 107, fig. 19. — ? 
Schuchert, 1996: 28, fig. 11. — Bouillon et al., 2004: 43, fig. 25D. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 229 


DiaGNOsIs: Mature medusa < 2 mm, without ocelli, 2-3 tentacles per bulb, oral 
tentacles with long basal trunks, divided 2-3 times, weak peduncle, large interradial 
gonads. 


DESCRIPTION (Bouillon, 1980): Adult medusa slightly higher than wide; jelly 
moderately thick, thicker apically than at sides; rather shallow gastric peduncle 
present. Radial and circular canal thin, bulbs rather small, hemispherical, without 
ocelli; 2 or rarely 3 tentacles on each bulb; tentacles with swollen distal end; oral ten- 
tacles divided dichotomously 2-3 times and ending in weak capitations, basal trunks of 
oral tentacles long (1/2 or more of total length). Stomach spanning about !/2 of sub- 
umbrellar cavity, conical. Gonads voluminous bulging interradial pads on stomach 
wall. Colours: distal end of tentacles green, inner side of gonads orange. Nematocysts: 
microbasic euryteles and desmonemes. Polyp stage unknown. 


DIMENSIONS (Bouillon, 1980): Height of adult medusa max 1.8 mm high, 
gonads visible even in specimens of 0.5 mm size. 


DISTRIBUTION: Papua New Guinea (Bouillon, 1980), ? Lebanon (Goy et al., 
1991), ? New Zealand (doubtful record, see Schuchert, 1996). Type locality: Laing 
Island, Papua New Guinea. 


REMARKS: This species resembles strongly Bougainvillia muscoides, notably 
through the absence of ocelli. However, it remains much smaller (1.8 mm versus 
3-5 mm), has fewer tentacles (2-3 versus 3-7), and the trunks of the oral tentacles are 
longer. The so far unknown polyp stage might perhaps present more distinguishing 
characters. 

The Mediterranean records of Bougainvillia aurantiaca are based on a single 
medusa from Lebanon reported by Goy et al. (1991). The animal depicted by them has 
no gonads despite it is 1.5 mm high. According to Bouillon (1980), animals of this size 
should have developed gonads. As the life cycle of this species is not known, I think 
that the presence of B. auriantiaca in the Mediterranean is not reliably proven. 


Bougainvillia niobe Mayer, 1894 
Bougainvillia niobe Mayer, 1894: 236, pl. 1 fig. 2. — Mayer, 1910: 166, pl. 18 figs 1-3, text 
fig. 90. — Kramp, 1959a: 110, fig. 93. — Bouillon et al., 2004: 44, fig. 26C. 
? Bougainvillia platygaster. — Goy et al., 1991: 108, fig. 20. 
REMARKS: This species is not present in European waters, but perhaps in the 
eastern Mediterranean, thus still in the region covered by the European Register of 
Marine Species (ERMS, Costello et al., 2001). See comments under B. platygaster. 


Bougainvillia multicilia (Haeckel, 1879) 
Lizusa multicilia Haeckel, 1879: 81, pl. 6 fig. 13. 
Bougainvillia multicilia. - Kramp, 1955b: 249. — Kramp, 1961: 78. — Vannucci & Rees, 1961: 

72. — Bouillon et al., 2004: 44. 

REMARKS: A doubtful species, perhaps based on a misinterpretation. Haeckel 
(1879) erected this species based on a medusa he saw near Algeciras (Gibraltar). The 
combination of unbranched oral tentacles, 10-12 tentacles per marginal bulb, and the 
abaxial ocelli is highly unusual. The species has never been seen again and the verdict 


230 P. SCHUCHERT 


of Kramp (1955b) is still valid: «...one can hardly suppress a doubt about the 
correctness of the description of this medusa.» 


Genus Koellikerina Kramp, 1939 
SYNONYM: Koellikeria L. Agassiz, 1862 [preoccupied by Koellikeria Cobbold, 1860 
(Trematoda)]. 


TYPE SPECIES: Melicerta fasciculata Péron & Lesueur, 1810, by designation of Kramp 
(1939). 


DiAcnosis: Hydroids colonial, stolonal or branching, covered by perisarc 
infested with foreign material; filmy perisarc also envelops hydranth body and basal 
part of tentacles; hydranths with one whorl of filiform tentacles; gonophores arise 
singly from hydranth pedicels, liberated as free medusae. 

Medusa with eight clusters of marginal tentacles, all tentacles similar in 
structure; with or without ocelli; with four dichotomously branching oral tentacles; 
gonads in perradial or adradial position. 


REMARKS: See Kramp (1939) for the taxonomic history. A tabular comparison 
of all species is given in Xu & Huang (2004). 


Koellikerina fasciculata (Péron & Lesueur, 1810) Jara 113) 


Melicerta fasciculata Péron & Lesueur, 1810: 353. 

Rathkea fasciculata. — Mayer, 1910: 179, fig. 94. 

Lizzia koellikeri Gegenbaur, 1854: 176, pl. 2 figs 1-9. 

Perigonimus cidaritis Weismann, 1883: 117. — Motz-Kossowska, 1905: 73, fig. 7. — Brückner, 
1914: 448, figs 1-6, pl. 8 figs 1-2. 

Thamnitis dichotoma Briickner, 1914: 460. 

Thamnostoma russelli Rees, 1938: 22, fig. 7. — Kramp, 1959a: 106, fig. 83. — Petersen & 
Vannucci, 1960: 486, synonym. 

Not Thamnostoma spec. — Russell, 1953: 151, fig. 73C-E [= B. muscoides]. 

Koellikerina fasciculata. — Kramp, 1959a: 112, fig. 98. — Petersen & Vannucci, 1960: 473, figs 
1-14.- Kramp, 1961: 85. — Brinckmann- Voss, 1970: pl. 9, figs 1-3. — Pages et al., 1992: 
5, fig. 3. — Bouillon er al., 2004: 46, fig. 27H-L. 


MATERIAL EXAMINED: One living medusa collected at Villefranche-sur-Mer, 0-70 m, May 
2001, not preserved, used to make DNA, 16 S sequence AM183129. — One living medusa 
collected at Villefranche-sur-Mer 6 April 2005, 0-70 m. — ZSM, Stechow collection, two slides 
with numbers 20000703 and 20000704, as Thamnitis cidaritis, Naples, April 1879, with young 
medusa buds, mentioned in Stechow (1919). - MHNG INVE29033, as Thamnostoma cidaritis, 
Naples, on spines of cidarioid sea-urchin, no medusa buds. - MHNG INVE32788, Naples 1892, 
7 mature medusae. — ZMUC, type material of Thamnostoma russelli. 


DESCRIPTION: Polyp growing on polychaetes and spines of cidaroid sea urchins. 
Stolons creeping, branched, giving off shoots of variable height, shoots with a long thin 
basal part widening distally into hydranth, shoots unbranched or occasionally branched 


Fic. 13 


Koellikerina fasciculata (Péron & Lesueur, 1810). (A) Mature medusa in lateral and oral view, 
scale bar 2 mm. (B) Oral tentacles, scale bar 0.5 mm. (C) Cluster of marginal tentacles seen from 
adaxial side, scale bar 0.5 mm. (D) Silhouette of a colony growing on sea urchin spine, scale bar 
5 mm. (E) Polyps with medusa bud, scale bar 0.5 mm. (F) Newly released medusa, height about 
0.8 mm. A-E, after preserved material; F, modified from Petersen & Vannucci (1960). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 231 


Ck 
IN 
\ DA, 


232 P. SCHUCHERT 


once or twice. Perisarc smooth, occasionally with corrugated stretches, perisarc 
continued as detritus infested gelatinous layer over hydranth and also investing lower 
parts of tentacles, wrinkled on hydranth body. Hydranths quite broad and therefore 
appearing club-like, with 14-18 tentacles in two very closely set whorls, hypostome 
conical. Medusa buds arise below hydranths, never on stolons, not detritus covered, 
pear-shaped, with pedicel. Nematocysts: microbasic euryteles and desmonemes. 

Newly released medusa with four bulbs, each with one tentacle, no ocelli, oral 
tentacles branched once, manubrium height about 1/4 of bell cavity, manubrial peduncle 
absent. Colour: manubrium light orange, with brick-red interradial bands, bulbs 
orange. 

Adult medusa nearly as broad as high, barrel-shaped bell, flatly rounded apex 
and relatively thick walls, velum narrow. Radial canals broad, mostly smooth. Four 
perradial and four interradial bulbs, bulbs broadly triangular and flat, about as broad as 
intermittent free bell margin; each of the eight bulb with up to 19 tentacles, tentacles 
issuing nearly parallel from bulb, getting slightly thicker after origin, then tapering 
again, where thickest with one adaxial ocellus, thus ocelli rather far away from bulb. 
Tentacles with chordoid gastrodermis, developing continuously, the youngest and 
shortest ones being the outermost ones, sometimes being mere stumps only, ocelli of 
youngest tentacles often not formed yet. Manubrium on a short broad peduncle, 
manubrium short, about 1/3 to 1/2 of bell cavity, cruciform cross-section; mouth 
cruciform, without lips, smooth; four perradial sets of oral tentacles, these inserting 
high above mouth level under protruding gonads, oral tentacles branched many times 
and with numerous ends (>100), ends with a concentration of euryteles but only indis- 
tinctly capitate; branching pattern of oral tentacles resembling a tight bush when 
tentacles contracted, oral tentacles can be expanded very much and the ends form a 
cloud in lower part of subumbrella. Oral tentacles with parenchymatic gastrodermis. 
Gonads adradial, U-shaped, lower ends free and overarching bases of oral tentacles; 
gonad partitioned by radiating furrows, thus forming up to ten lappets on each side. 
Nematocysts: microbasic euryteles and two types of desmonemes differing in their 
morphology. Colours: Oral tentacles and gonads intensively red. 


OTHER DATA: Petersen & Vannucci (1960) described and depicted the develop- 
ment of the medusa in great detail. 

Mayer (1910) observed that whenever the medusa is disturbed by a mechani- 
cal shock, the oral suddenly expand to their full extent, forming a network of filaments 
around the mouth. Such a reaction is remarkable, for in common with other medusae 
all other parts of the animal contract when disturbed. 


DIMENSIONS: Polyp pedicels 4-6 mm, basal diameter 0.07 mm, hydranths about 
0.5 mm high (preserved material). Mature medusa 7-13 mm high, egg size 110 um 
(Petersen & Vannucci, 1960). Detailed nematocyst measurements can be found in 
Petersen & Vannucci (1960). 


DISTRIBUTION: Mediterranean; Black Sea; Cape Verde Islands; Red Sea 
(Schmidt, 1973a); polyp and medusae also in Scotland and Norway, but perhaps not 
reproducing there (Petersen & Vannucci, 1960; Fraser, 1973). Type locality: 
Mediterranean (Péron & Lesueur, 1810). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 233 


BroLocy: The medusa lives for seven months or more and it requires tempera- 
tures above 8°C for maturation. The periods with sufficiently warm water temperatures 
in the NE Atlantic are perhaps too short to allow the medusa to mature. In the 
Mediterranean, the medusa is found from autumn to spring (Petersen & Vannucci, 
1960). The medusae are most abundant in late autumn and winter, occasional 
specimens may be found as late as April or May. The medusae can tolerate reduced 
salinities down to 28 ppt. The polyps lives in deeper waters (20-270 m) on the spines 
of sea urchins (Stylocidaris affınis, Dorocidaris papillata) and on the parapodia of 
polychaetes (Aphrodite aculeate, Hermione hystrix). They prefer colder waters and are 
therefore not found above 20 m depth (Petersen & Vannucci, 1960). 


REMARKS: The blood-red colour and the eight tentacle clusters make this 
medusa very conspicuous and it is easy to recognize. The life cycle was revealed by 
Brückner (1914) and Petersen & Vannucci (1960). For the synonymy see also the lat- 
ter publication. 


Genus Thamnostoma Haeckel, 1879 


SYNONYM: Thamnitis Haeckel, 1879. 
TYPE SPECIES: Lizzia dibalia Busch, 1851 [designation by Kramp, 1959a]. 


DIAGNOSIS: Bougainvilliidae medusa with four to eight solitary marginal 
tentacles and with four branched oral tentacles inserting above mouth rim; four 
interradial gonads; with ocelli. Polyp stage not reliably known, similar to genus 
Koellikerina. 


REMARKS: The gender of Thamnostoma is neuter (ICZN 30.1.2). No polyp stage 
of an unambiguously identifiable (mature) Thamnostoma medusa is known. Hirohito 
(1988) described some polyps that released young medusae referable to the genus 
Thamnostoma. However, also Koellikerina fasciculata medusae go through a 
Thamnostoma stage during their development (Petersen & Vannucci, 1960; described 
as a distinct species, Thamnostoma russelli Rees, 1938). 


Thamnostoma dibalium (Busch, 1851) Fig. 14 


Lizzia dibalia Busch, 1851: 23, pl. 1 figs 7-9. 

Thamnostoma dibolia. — Haeckel, 1879: 86. — Neppi & Stiasny, 1913: 27, pl. 2, fig. 15. — 
Bouillon et al,. 2004: 49, Fig. 29D. [incorrect spelling]. 

Thamnostomum dibalium. — Kramp, 1959a: 106, fig. 82. — Kramp, 1961: 91. 


DIAGNOSIS: See genus. 


DESCRIPTION (Busch, 1851; Neppi & Stiasny, 1913): Umbrella bell-shaped, 
somewhat higher than wide; manubrial peduncle short, conical; eight marginal tentacle 
bulbs, each with one tentacle, all tentacles of the same length in mature animals, at 
tentacle base an adaxial adnate spur-like process which bears an ocellus at its end. 
Manubrium cylindrical to ovoid, spanning about half of the bell cavity; four thick oral 
tentacles, two to three times dichotomously branched (the third division incipient 
only). Gonads as four bulging interradial pads along almost the whole length of the 
manubrium, eggs large, conspicuous. Colour: stomach and marginal bulbs dark-brown. 


DIMENSIONS: Bell height 3.5-7 mm, 3-6 mm wide. 


234 P. SCHUCHERT 


Fic. 14 


Thamnostoma dibalum (Busch, 1851). (A-B) Medusae, no scale given. (C) Tentacle seen from 
side, note process with ocellus. (D) Tentacle bulb, base of tentacle and adaxial spur with ocellus 
seen from adaxial side. A, modified after Neppi & Stiasny (1913); B-D, after Busch (1851). 


BIOLOGY: A rare species, observed in September and October. 


DISTRIBUTION: Adriatic Sea (Benovic & Lucic, 1996), Mediterranean coast of 
Spain (Riera er al., 1986). Type locality: Trieste, Adriatic Sea. 


REMARKS: The gender of Thamnostoma is neuter, therefore the specific epithet 
must be dibalium. Haeckel (1879) proposed the name dibolia, which is incorrect (see 
Russell, 1953: 152). According to Neppi & Stiasny (1913), there is only one ocellus 
per tentacle. Haeckel (1879) mentioned two ocelli, one on a process, the other not. 
Busch (1851) describes that there is a pigment in the bulb as well as on a process of 
the tentacles like a stalked eye, but he also explicitly states that he could not decide 
whether both spots were really functional eyes. 


Genus Nemopsis L. Agassız, 1849 
TYPE SPECIES: Nemopsis bachei L. Agassiz, 1849, my monotypy. 


DIAGNOSIS: Hydroids, solitary, small, club-shaped, with pedal disk, hydrocaulus 
not distinctly delimited from hydranth body, large medusa buds in basal region. 
Medusa with four clusters of marginal tentacles, in each cluster a median pair of club- 
shaped or capitate tentacles flanked by a number of simple, filiform tentacles; with 
adaxial ocelli; with four perradial dichotomously branched tentacles; manubrium with 
four radial lobes extending outwards along the radial canals; gonads on these lobes. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 235 


Nemopsis bachei L. Agassiz, 1849 Fig. 15 


Nemopsis bachei L. Agassiz, 1849: 289, fig. 1.— A. Agassiz, 1865: 149, figs. 227-231. — Mayer, 
1910: 173, pl. 17, figs 5-6. — Hartlaub, 1911: 194, fig. 174. — Tiffon, 1956; 550, fig. 1.— 
Kramp, 1959a: 111, fig. 97. - Kramp, 1961: 89. — Kühl, 1962: 221, figs 11, pls 5-6. — 
Thiel, 1969: 71, fig. 3. — Kühl, 1972: 250, figs 18-22. — Denayer, 1973: 290, fig. 7. — 
Dumoulin, 1997: 102, figs 3-6. — Faasse & Ates, 1998: 72, figs 1-2. — Tulp, 2002: 89, 
figs. 1-2, photo 1. 

Nemopsis gibbesii McCrady, 1859: 160, pl. 10, figs 1-3 [medusa only, not hydroid]. 

? Hippocrene crucifera Forbes & Goodsir, 1853. — Edwards, 1966: 143, synonym. 

? Hippocrene cruciata Forbes & Goodsir, 1853 [in legend to figures, incorrect spelling]. 

Nemopsis heteronema Haeckel, 1879: 93, pl. 5 figs 6-9. — Hartlaub, 1911: 197, fig. 174. 

Bougainvillia charcoti Le Danois, 1914a: 15, figs 1-3. — Kramp, 1959a: 111. — Edwards, 1966: 
142, synonym. 

Nemopsis crucifera. — Hartlaub, 1911: 195, fig 173. 

Bougainvillia charkoti. — Hartlaub, 1917: 406 [subsequent incorrect spelling]. 

? not Nemopsis Bachei. — Brooks, 1883: 468 [hydroid]. 


MATERIAL EXAMINED: BMNH 1967.10.20.12-20; USA, North Carolina, Morehead City; 
18.03.1964; 50 mature medusae. - BMNH 1982.11.30.2; Germany, Weser Estuary; 22.02.1961; 
6 mature, badly preserved medusae, up to 1 cm in size. 


DIAGNOsIs: Medusa up to 12 mm, four marginal bulbs, each with 14-34 filiform 
tentacles and a pair of capitate or clavate tentacles in middle; stomach base with large 
perradial basal lobes rendering it cruciform, gonads on these lobes; four oral tentacles 
branched up to eight times. Polyp very small, solitary, club-shaped, indistinct pedicel, 
large pedal disc, one whorl of long tentacles, medusa buds large. 


DESCRIPTION: Polyp stage (after Kiihl, 1962; 1972), small, solitary, with a 
relatively large pad-like attachment disc; hydranth body club-shaped, thick distal end 
turning into thinner pedicel without clear transition. Primary polyps with 2-4 tentacles, 
mature polyps with a single whorl of 8-14 tentacles, these much longer than the size of 
the polyp, often curled up; hydranth body enveloped by detritus. The medusa buds 
develop below the tentacles (likely at junction of hydranth body and pedicel), up to 
four buds in one whorl. 

Newly released medusa spherical, jelly thick, 2-3 tentacles per marginal bulb, 
club-shaped tentacles not formed yet, oral tentacles not branched, no manubrial lobes 
present. 

Adult medusa bell-shaped, higher or as high as wide, jelly very thick, apical 
Jelly about 1/3 of bell-height, top and sides curved, margin lobed through perradial 
embayments, no manubrial peduncle. Four radial canals ending in broad marginal 
bulbs, ring canal thin, bulbs usually contracted to U-shape, outline in oral view 
irregular, each bulb with 8-28 filiform tentacles, ends sometimes swollen in living 
animals, ocelli on bulb at origin of tentacles, in addition to filiform tentacles a median 
pair of short club-shaped to capitate tentacles, in younger animals this pair of shorter 
tentacles more club-shaped (depending on state of contraction) or even absent, ocelli 
of capitate tentacles on adaxial side of tentacles on a swelling and not on bulb. 
Manubrium conical, ending in a mouth with four short lips, base of manubrium with 
very large perradial lobes, lobes extend along the radial canals for up to 5/6 of the 
canals, lobes render stomach large and cruciform; four perradial oral tentacles arising 
well above mouth margin, branching 5-8 times, basal trunk long, termini slightly 


236 P. SCHUCHERT 


Fic. 15 


Nemopsis bachei L. Agassiz, 1849; after preserved material. (A) Medusa, scale bar 1 mm. (B) 
Stomach seen from aboral, same scale as in A. (C) Tentacle bulb with the characteristic capitate 
tentacles. (D) Flattened tentacle bulb from adaxial, note that in living animals the bulb is usually 
contracted to a U-shape, scale bar 0.2 mm. (E) Capitate tentacle in side view, note ocellus on 
basal swelling. 


swollen with concentration of nematocysts. Gonads a thick layer on the manubrium 
and its basal outgrowths, with vertical folds. Colours: Manubrium and gonads milky 
white to yellow. Nematocysts: microbasic euryteles and desmonemes 


VARIATION: Thiel (1969) made a detailed investigation on the correlation of bell- 
size, tentacle number, and branching order of the oral tentacles. The German popu- 
lation may have up to 36 filiform tentacles per bulb (Kiihl, 1962), but the usual range 
is about 14-27 tentacles. The American population seems to have less (14 acc. to 
Mayer, 1910), while the Japanese N. dofleini has about 40 (Kramp, 1968). Hargitt 
(1901) described the variations and aberrations found in the American population. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 237 


DIMENSIONS: Adult medusa around 10 mm, usually not larger than 12 mm, but 
very rarely up to 14-18 mm (Thiel, 1969). Newly hatched polyps about 0.15-0.18 mm 
high, mature polyps about 0.5-0.6 mm, tentacle 0.9 mm long, newly released medusa 
diameter 1.5 mm (Kühl, 1962; 1972). 


DISTRIBUTION: In the western Atlantic occurring from Woods Hole to Florida 
and the northern Gulf of Mexico (Kramp, 1959a; Moore, 1962), in the NE-Atlantic 
occurring from Norway to the Bay of Biscay, usually in estuaries. European records are 
e. g.: Norway (Nemopsis heteronema Haeckel, 1879; Sognefjord); Scotland (Edwards, 
1966; Fraser, 1973); Germany (Kühl, 1962, 1972; Thiel, 1968, 1969); Holland (Faasse, 
& Ates, 1998; Tulp, 2002); Belgium (Dumoulin, 1997); France, Loire Estuary 
(Denayer, 1973); France, Gironde (Tiffon, 1956). Type locality: Nantucket, Vineyard 
Sound, Massachusetts, USA. 


BioLoGY: Nemopsis bachei prefers reduced salinities and high plankton 
densities, it is thus often found in estuaries. Kühl (1972) reports that the medusa occurs 
from May to October in great quantities in the Elbe Estuary. Nemopsis bachei is 
thought to have been introduced several times to Europe through its polyp stage. The 
history of these introductions is analyzed in Thiel (1968). Aspects of the ecology of N. 
bachei were investigated by Purcell & Nemazie (1992) and Purcell et al. (1994). The 
polyp grows very slowly and it takes 9-12 months to attain its full size. Also the 
medusa-bud development is very slow, taking up to two months (Kühl, 1972). 


REMARKS: Among the medusa of the present fauna, Nemopsis bachei is an 
easily identifiable species due to its very long stomach lobes in combination with 
clustered tentacles. The pair of small capitate tentacles makes this species even more 
unique, but these capitate tentacles are often missing or not visible in preserved 
plankton samples. 

The synonymy of this species has been discussed by several authors, e. g. by 
Mayer (1910), Nagao (1964) and Thiel (1969). Current opinion is that there are only 
two valid Nemopsis species, namely N. bachei and N. dofleini Maas, 1909. Nemopsis 
dofleini from Japan has more tentacles per bulb (about 40) and is larger (13 mm) 
(Kramp, 1968). Thiel (1968) showed that the Japanese medusae have an elevated 
tentacle number at all size stages and he interpreted this as a good argument that N. 
dofleini is a separate species. The present author has nevertheless some doubts, as local 
populations of introduced species like Nemopsis medusae often have reduced genetic 
variability due to founder effects and they can be quite distinct from other populations 
because of the absence of intermediate genotypes. Although tentacle numbers seem 
indeed lower in N. bachei, the maximal tentacle number of 36 filiform tentacles per 
bulb of the European population seems not so drastically different from N. dofleini. The 
American medusae seem to have fewer tentacles per bulb (14 after Mayer, 1910). 

In his study on medusae of Charleston Harbor, McCrady (1859) also described 
Nemopsis gibbesii. His description was mainly based on a series of medusae that are 
clearly referable to Nemopsis bachei. But McCrady also found a polyp in the plankton 
which he erroneously associated with this medusa. The polyp closely resembles the 
polyp of Margelopsis haeckelii and Hartlaub (1899) pointed out that McCrady’s 
medusae were actually Nemopsis bachei and the polyp belonged to the genus 


238 P. SCHUCHERT 


Margelopsis. Similar conclusions concerning the medusa were already formulated by 
A. Agassiz (1865) and Brooks (1883). Brooks (1883) seems to have made another error 
when describing the hydroid stage of N. bachei. The hydroid was found at a place 
where N. bachei occurred abundantly. The colony was about 2.5 cm high and branched. 
The hydranths had funnel-shaped hypostomes, 24 tentacles, six to eight gonophores 
developed on the hydranth body. Brooks noted that the hydroid looked in fact much 
more like a Eudendrium than a Bougainvillia. Nevertheless, he claimed to have found 
newly released bougainvilliid medusae which he could keep for a short time until they 
had formed stage with four tentacles per bulb. To me, the presented data are insufficient 
to prove that Brooks indeed observed the medusa of N. bachei as he did not follow the 
medusae to a stage were they are unambiguously identifiable. The hydroid is most 
peculiar and perhaps he had in fact a Eudendrium species that had some epizoic 
Nemopsis hydroids. The hydroid of N. bachei was described by Kühl (1962; 1972) and 
shortly afterwards also the hydroid of N. dofleini became known (Nagao, 1964). Both 
authors found a very small, solitary hydroid and it is thus easily conceivable that 
Brooks had perhaps overlooked these polyps in his sample. 

The hydroids of both nominal species N. bachei and N. dofleini appear very 
similar, again suggesting that both species might in fact be conspecific. As the figures 
in Kühl (1962; 1972) are not suitable for reproduction, the polyp of N. dofleini is shown 
instead (Fig. 16). 

The position of the gonophores in Nemopsis is quite remarkable. The 
gonophores of the Bougainvilliidae and Pandeidae are never on the hydranth body as 
e. g. in the Hydractiniidae and most other hydroids (= plesiomorphic condition). In 
Nemopsis, the gonophore position is not entirely clear, it could be on the hydranth body 
or at the junction of the body and the pedicel. The pedicel is, however, now well dis- 
tinguishable from the hydranth body. This reversal, or near-reversal, of the gonophore 
position is certainly linked to the modification of the polyp to a solitary, almost sessile 
form. 


Genus Nubiella Bouillon, 1980 
TYPE SPECIES: Nubiella mitra Bouillon, 1980. 


DIAGNOSIS: Bougainvilliidae medusae with four unbranched oral tentacles; 
umbrella with four marginal bulbs, each with a single tentacle. 


Nubiella mitra Bouillon, 1980 
Nubiella mitra Bouillon, 1980: 314, fig. 4. — Goy et al., 1991: 109, fig. 22. — Bouillon et al., 

2004: 47, fig. 28G. 

DrAGNosis: Medusa bell with conical apex, apical jelly about 1/3 of total height, 
bell height 1.7 mm, 1.2 mm diameter; no gastric peduncle; manubrium cylindrical, 
spanning more than half of the bell cavity, with four simple capitate oral tentacles 
attached above mouth rim; four marginal bulbs each with one tentacles, without ocelli; 
gonads encircling manubrium; several medusa buds can be present on the upper part 
of the manubrium. 


DISTRIBUTION: Papua New Guinea; eastern Mediterranean. Type locality: Laing 
Island, Papua New Guinea. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 239 


Fic. 16 


Nemopsis dofleini Maas, 1909; modified from Nagao (1964). The polyp stage of N. bachei is 
practically indentical. (A) Polyp stage with two medusa buds, scale bar 0.2 mm. (B) Newly 
released medusa, scale bar 0.5 mm. 


REMARKS: This species, the only one in its genus, has not been recorded in 
European waters. The Mediterranean records are based on two medusae found off 
Lebanon by Goy et al. (1991). In the figure given by Goy et al. (1991), the oral 
tentacles arise at the mouth margin and not above it as it is typical for this species. 


Genus Silhouetta Millard & Bouillon, 1973 
TYPE SPECIES: Silhouetta uvacarpa Millard & Bouillon, 1973. 


DrAGNosis: Colonial hydroids, stolonal or erect, with firm perisarc terminating 
at hydranth base; hydranths broad, ovoid to club-shaped, tentacles filiform, in two or 
more whorls concentrated in a band below hypostome, tentacles of one whorl alter- 
nating with those of adjacent whorls. 

Gonophores free medusae, arising in clusters from stem and branches. Medusae 
at liberation with four simple or dichotomously branched oral tentacles attached above 
mouth rim; four tentacle bulbs, each with a single tentacle; ocelli present. 


REMARKS: The hydranths of Silhouetta appear identical to those of Pachy- 
cordyle napolitana (see Schuchert, 2004). 


Silhouetta uvacarpa Millard & Bouillon, 1973 Fig: 17 


Silhouetta uvacarpa Millard & Bouillon, 1973: 25, figs 3A-D, pls 2-3. — Millard & Bouillon, 

1975: 3. — Calder, 1988: 17, figs 13-14. 

MATERIAL EXAMINED: Holotype, Musée Royal de |’ Afrique centrale, Tervuren, Belgium; 
reg. No 2808; loc. Seychelles, Silhouette, 1966; several colonies, with medusa buds, leg. Millard 
and Bouillon. - BMNH 1990.5.2.7 and 1990.5.2.9, labelled Clavopsella navis, later added 
Silhouetta uvacarpa; Azores Faial, on Mytilus galloprovincialis, 0.3 m depth, 26.07.1989, coll. 
P. F. S. Cornelius, some with medusa buds. 


DIAGNOSIS: As for genus. 


240 P. SCHUCHERT 


Fic. 17 


Silhouetta uvacarpa Millard & Bouillon, 1973; after type material. (A) Terminal branch with 
hydranth and two typical clusters of medusa buds, the lower one almost completely spent, scale 
bar 0.2 mm. (B) Tip of semi-extended tentacle, note moniliform arrangement of nematocyst 
clusters, scale bar 0.05 mm. 


DESCRIPTION: Colonies initially stolonal, later erect, arising from creeping 
hydrorhiza; older colonies can be polysiphonic (Calder, 1988). Perisarc wrinkled in 
certain areas, smooth in others, no annulation, perisarc ends below hydranth body, no 
pseudohydrotheca. Hydranths terminal on stem and branches, large, club-shaped, 
widest diameter in terminal third, evenly tapering towards proximal, hypostome 
conical, with 16-26 tentacles in 2-4 closely alternating whorls immediately below the 
hypostome. The youngest hydranths have only two whorls and new tentacles appear to 
arise proximal of them, thus increasing the number of whorls with age. Extended 
tentacles moniliform. Nematocysts: microbasic euryteles and desmonemes. 

Gonophores free medusae, developing in grape-like clusters on stem or 
branches, 12-20 to a cluster, pedicels long and thin, originating as a common trunk with 
a distal arborescent branching carrying terminal medusa buds. Each bud completely 
surrounded by transparent perisarc. 

Newly released medusa with four marginal bulbs, each with one black ocellus 
and one tentacle; four unbranched oral capitate tentacles arising just above the mouth 
margin. 

Mature medusa unknown. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 241 


Dimensions (Millard & Bouillon, 1973, 1975; Calder, 1988): Colonies reaching 
2.3 cm in height, diameter of branches 0.29 mm; hydranth up to 1.5 mm high and 
0.9 mm wide. Newly released medusa up to 0.3 mm high and 0.23 mm wide. 
Nematocysts see Calder (1988). 


BioLoGY: The hydroid has been found in shallow waters (1-30 m) of tropical 
seas on Pecten shells and sponges. 


DISTRIBUTION: Tropical seas, reported from the Seychelles (Millard & Bouillon, 
1973), Puerto Rico (Wedler & Larson, 1986), Bermudas (Calder, 1988), and Guam 
(Kirkendale & Calder, 2003). Type locality: Île Silhouette, Seychelles Islands. 


REMARKS: Cornelius (1992) briefly mentioned that this species also occurs in 
the region of the Azores. Cornelius’s material could be re-examined for this study and 
it was compared with the type material. The colony is growing on a Mytilus shell and 
there can be no doubt that it is the material Cornelius was referring to. However, the 
specimens could not convince me that this is Silhouetta uvacarpa because they lack 
some of the characteristic features of this species, namely the clustered gonophores on 
arborescent pedicels, the tentacles in several whorls, and the large, club-shaped 
hydranth. There are if fact only few medusa buds and the whole colony resembles more 
a Bougainvillia colony, despite that some hydranths are thicker than usual. It is possible 
that Cornelius had based his identification on the released medusae, but these could not 
be re-examined. 

Considering its wide distribution, Silhouetta uvacarpa could perhaps occur in 
the Azores, but currently its presence there must be taken as unproven. 


Genus Dicoryne Allman, 1859 

TYPE SPECIES: Dicoryne stricta Allman, 1859b [= D. conferta (Alder, 1856)]. 

SYNONYM: Heterocordyle Allman, 1864a [type species Heterocordyle conybearei 
Allman, 1864]. 

Diacnosis: Colonial hydroids, branched or not, perisarc not ensheathing bases 
of tentacles; hydranth with one whorl of filiform tentacles; gonophores on specialized 
blastostyles replacing a hydranth, numerous, released as swimming ciliated sporosacs 
provided with one or two tentacle-like processes at their proximal end (tentaculoids). 


REMARKS: See Ashworth and Ritchie (1915) for a first important revision of the 


genus. 

KEY TO THE SPECIES: 
la sporosacs with mostly 2 tentaculoids, females 2 eggs ............ Dicoryne conferta 
lb sporosacs with mostly 1 tentaculoid, females 1 egg ........... Dicoryne conybearei 
Dicoryne conferta (Alder, 1856) Fig. 18 


Eudendrium confertum Alder, 1856: 354, pl. 12 figs 5-8. — Cornelius & Garfath, 1980: 278. 

Dicoryne stricta Allman, 1859b: 369. — Allman, 1861: 168, synonym. 

Dicoryne flexuosa Sars, 1874: 128, pl. 5 figs 21-26. — Fraser, 1944: 56, pl. 6 fig. 26. 

Dicoryne conferta.— Allman, 1861: 168, fig’d. — Hincks, 1868: 105, pl. 18 fig 1.— Allman, 1872: 
293, pl. 8. — Goette, 1907: 66, pl 6. - Ashworth & Ritchie, 1915: 266, pl. 8 fig. 16. — 
Fraser, 1944: 55, pl. 6 fig. 25. - Naumov, 1969: 218, fig. 86. — Millard, 1975: 101, fig. 
34E-J. — Schuchert, 2001: 24, fig. 13. — Bouillon et al., 2004: 45, fig. 26E-G. 

Bougainvillia conferta. — Broch, 1916: 50, fig. O. 

Perigonimus conferta.— Vervoort, 1946: 144, figs 52b & 56. 


242 P. SCHUCHERT 


a 


SER u. 
mn. 


Fic. 18 
Dicoryne conferta (Alder, 1856). (A) A single stem with hydranths and blastostyles, scale bar 
0.5 mm. (B) Hydranth and blastostyles with sporosacs, scale bar 0.2 mm. (C) Stem perisarc, note 
the internal corrugation, scale bar 0.1 mm. (D) Young female sporosacs with two eggs, scale bar 
0.05 mm. (E) Mature, but still attached sporosacs in side view, the rear tentaculoid is not visible, 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 243 


MATERIAL EXAMINED: MHNG INVE33562; Iceland, 65.35°N 13.61°W, 65 m, 19 July 
1991, large colony on 5 cm hermit crab shell, fertile female, two eggs per sporosac. - MHNG 
INVE27337; Iceland, 65.35°N 13.33°W, 173-177 m, 18 July 1987, fertile male. - BMNH 
1965.1.14.15; Sweden, Kosterfjord, Ramsö, coll. W. J. Rees 1 October 1964, on shell with 
Eupagurus berhardus, luxurious female colony, often two eggs per sporosac, some confirmed 
with 2 tentaculoids. - BMNH 1912.12.21.157; Great Britain, Shetland; Norman collection; on 
gastropod shells used by hermit crabs, female with 2 eggs per sporosac, some 1. - BMNH 
1957.10.24.32; Italy, Naples, off Santa Lucia; coll. W. J. Rees, October 1957, on Aporrhais pe- 
spelicani, female 1-2 eggs per sporosac, not well preserved. - BMMN 1962.11.8.7 and 
1962.11.8.6 (slide); Sweden, Gullmarfjord, Gäsö, Ranna; 20-30 m; on bucciniid shell with her- 
mit crab; coll. 27 August 1962 by W. J. Rees, males and females, among 19 examined female 
sporosacs 18 with two eggs, one with one egg, two tentaculoids were visible. - BMNH 
1964.8.7.17 slide; Sweden, Kristineberg; male colony, 3-4 times branched, sporosacs with 2 ten- 
taculoids. - BMNH 1964.8.7.19-20, slides; North Sea, stems up to 8 mm, branched several 
times, male, at least one sporosac seen with two tentaculoids - ZMUC; Denmark, Frederikshavn, 
Mellum; 8 m, 24 June 1939; fertile. - ZMUC; Denmark, Frederikshavn, Blanetbund; 15 m, 14 
August 1972, fertile. - ZMUC, Denmark, North of Haustholm; 33 m, 26 September 1930, fertile 
female with 2 eggs per sporosacs. - ZMUC, without number; Denmark, Frederikshavn; liberated 
sporosacs from polyp growing on Aporrhais pespelecani. — ZSM, Stechow collection, 
Helgoland, slides 2000523 & 2000524, fertile males, identity not clear. 


DIAGNOSIS: Dicoryne species with predominantly two eggs and two tentaculoids 
per sporosac 


DESCRIPTION: Colonies growing on gastropod shells, stems numerous and 
dense, arising from reticulate, perisarc covered stolons. Individual stems unbranched 
or variably branched with up to 20 hydranths, monosiphonic, height depending on 
environment, all branches ending with hydranths or blastostyles, stems increasing 
slightly in diameter towards distal, often with distinct primary axis and shorter side- 
branches. Perisarc outside wrinkled and infested with detritus, inner wall strongly and 
irregularly corrugated throughout, in terminal region filmy and variably continued or 
not on hydranth body as pseudohydrotheca, usually short and only covering basal part 
of the hydranth. Hydranth fusiform, conical hypostome, 10-16 (usually 10-12) 
tentacles in one or two very closely set whorls. Blastostyles arise either as side 
branches from stems or directly from stolons, their caulus perisarc covered, blastostyle 
spindle-shaped, often lower part bearing the sporosacs thinner, no tentacles, no mouth, 
terminal region densely studded with large euryteles giving a strong refringency and 
thus well visible, lower 2/3 densely covered by numerous sporosacs on short stalks; the 
blastostyle - especially the upper part without sporosacs - can make worm-like move- 
ments, the shape is thus not fixed. Colonies gonochoristic. Perisarc colour brown. 

Gonophores released when mature as swimming sporosacs of the styloid type, 
when attached covered with a very thin perisarc membrane, released sporosacs ovoid, 
covered by cilia, with two tentaculoids at its base directed away from the sporosacs, 
sometimes only one tentaculoid present, the tentaculoids develop only in the final 
maturation phase of the sporosac; female sporosacs predominantly with two eggs, a 


scale bar 0.05 mm. (F) Optical horizontal cross section of sporosac shown in E, note the two 
tentaculoids and the two eggs, same scale as E. (G) Optical section of an attached male 
sporosacs, note the two tentaculoids, same scale as E. (H) Free, swimming sporosacs, note that 
the posterior egg is hidden by the anterior one. A-G, after preserved material from Iceland and 
Sweden; H, after Allman (1861). 


244 P. SCHUCHERT 


minority of sporosacs with one egg or eggs unequally developed, eggs in a plane 
perpendicular to the two tentaculoids. 

Nematocyst: large microbasic euryteles, only on tip of blastostyles; smaller 
microbasic euryteles; desmonemes, only on tentacles of gastrozooids. 


DIMENSIONS: Stems up to 25 mm high, usually less than 12 mm; stem diameter 
0.1-0.12 mm; hydranths 0.4-0.9 mm high, terminal hydranth usually largest, other 
hydranths 0.4-0.6 mm. Sporosacs 0.13-0.15 mm long, diameter 0.10-0.12 mm, ten- 
taculoids 0.10-0.12 mm. Nematocyst: large microbasic euryteles of blastostyle 
(13-16)x(5.5-6.5)um; microbasic euryteles of hydranth (6-8)x(3-5)ym; desmonemes 
3x2.5 um. 


FURTHER DATA: Histological section of the sporosacs and their development are 
show in Goette (1907) and Ashworth & Ritchie (1915). 


BIOLOGY: Grows on gastropod shells, either inhabited by the gastropod or 
hermit crabs, sometimes on spider crabs, usually in depths of 5 to 300 m. In the 
Atlantic, mature sporosacs have been observed from March to October, in the 
Mediterranean in summer and autumn. 


DISTRIBUTION: Widely known in the eastern Atlantic from the Barents Sea to the 
Strait of Gibraltar and likely also western Mediterranean (e. g. Sars, 1874; Hincks, 
1868; Allman, 1872; Motz-Kossowska, 1905; Robson, 1914; Vervoort, 1946; Leloup, 
1947; Naumov, 1969; Vervoort, 1985; Medel & Löpez-Gonzälez, 1996; Bouillon et al., 
2004); also in the North Sea (Hartlaub, 1894; Jäderholm, 1909; Rees & Rowe, 1969 
Christiansen, 1972); Iceland and the Faroes (Broch, 1916; Schuchert, 2001), Atlantic 
Canada to Massachusetts Bay (Calder, 2003), and South Africa (Millard, 1975). Type 
locality: Cullercoats, England. 


REMARKS: Dicoryne conferta and Dicoryne conybearei are relatively easy to 
recognize, even when infertile, due to the characteristic internal perisarc corrugation 
and their epizoic growth on gastropod shells. However these two species are by now 
means easily separable. The table provided by Ashworth and Ritchie (1915) is only 
partially correct and I have not found characteristics of the trophosome that would 
permit distinguishing the two. The same view has already been expressed by Allman 
(1864b) himself when describing D. conybearei. The only reliably observable 
difference is found in the sporosacs. Dicoryne conferta has usually two tentaculoids 
and the female sporosacs have two eggs. Although for a given colony this seems to 
hold true, there is nevertheless some variation known. There may be one or two 
tentaculoids in Dicoryne conybearei (Ashworth & Ritchie, 1915; Schuchert, 1996), 
leaving only the egg number as a diagnostic character. But also this character shows 
some variation in D. conferta. In 18 examined sporosacs of one colony, one had only 
one egg, the others two. Also Millard (1975) observed such a variation for D. conferta. 
The same variation was observed for nominal D. conybearei samples: 2 out of 20 
sporosacs had two eggs instead of one only (see below). This makes it arguable 
whether these two species should be regarded as distinct. They might be just two 
morphotypes of the same biological species, the one of D. conferta apparently more 
prevalent in northern regions. Because the egg number is either predominantly one or 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 245 


two, which correlates rather well with the number of tentaculoids, I prefer to keep them 
separate for the moment. More detailed studies are clearly needed. 

Because many authors based their identification on unreliable differences of the 
trophosome, many records are not trustworthy. Especially the Mediterranean records 
need re-confirmation. 

An important revision of the genus was made by Ashworth & Ritchie (1915). 
Sars (1874) depicted the female sporosacs of his Dicoryne flexuosa and they have 
clearly two eggs. So, this name is obviously a synonym of D. conferta. The differences 
in the trophosome as given in Fraser (1944) are not diagnostic. 


Dicoryne conybearei (Allman, 1864) Rise: 19 


Heterocordyle conybearei Allman, 1864a: 366. — Allman, 1864b: 59, pl. 2. — Hincks, 1868: 107, 
pl. 18 fig. 2. — Allman, 1872: 307, pl. 10 figs 4-7. 

Heterocordyle conybeary — Weismann, 1883: 84, pl. 11 figs 7-9 [subsequent incorrect spelling]. 

Dicoryne conybeari. — Motz-Kossowska, 1905: 76. — Philbert, 1935: 19, fig. 2. - Bouillon et al., 
2004: 45, fig. 17A. [incorrect spellings]. 

Dicoryne conybearei. — Ashworth & Ritchie, 1915: 257, fig. 3, pl. 6. 

? Perigonimus (?) nanellus Stechow, 1919: 14, fig. C. — Rees, 1956a: 342. 

? Heterocordyle conybearei — Neppi, 1921: 21. 

Heterocordile conibeari. — Bavestrello, 1985: 351, fig. 1-a-b, [subsequent incorrect spelling]. 
MATERIAL EXAMINED: ZSM 2000643 & 20000644, holotype material of Perigonimus (?) 

nanellus, 2 slides, from Dorocidaris papillata spines. Blastostyles without gonophores present. 

— MHNG INVE32949, Banyuls-sur-Mer, examined alive, 8 May 2002, on Hinia incrassata, 

fertile female, 16S sequence AM183141.- Ischia, Naples, 17 November 1992, examined alive, 

on Murex spec., fertile female with one egg per sporosac. - ZSM 20000526 and 20000527, as 

Dicoryne conferta, two slides, Stechow collection, Villefranche-sur-Mer, May 1909, fertile 

females; sporosacs with | tentaculoid, one egg per sporosac but rarely (2 in 20) two eggs. — 

IRSNB, Roscoff, coll. J. Bouillon, fertile male on small gastropod shell inhabited by hermit crab, 

almost exclusively stolonal, one tentaculoid per sporosac. - BMNH 1957.10.24.29 as Dicoryne 

conferta; Italy, Naples, off Stazione Zoologica, October 1957, on Cerithium vulgaris shell, coll. 

W. J. Rees, female, one egg per sporosac seen, but not well preserved, thus more likely D. cony- 

bearei. 


DIAGNOSIS: Dicoryne species with predominantly one egg and one tentaculoid 
per sporosac. 


DESCRIPTION: See under Dicoryne conferta, the only differences are given in the 
diagnosis. 


OTHER DATA: Ashworth & Ritchie (1915) provide many histological details of 
the sporosac and its development. Philbert (1935) observed terminal stolons that are 
used for spreading to other substrata. 


DIMENSIONS AND OTHER DATA: Up to 8 hydranths per stem seen, stems up to 
10 mm; hydranths 0.3-0.6 mm, 8-16 tentacles. Sporosacs about 0.12 mm long and 
0.1 mm wide. Nematocysts (living capsules in material from Ischia): gonozooids have 
small microbasic euryteles (9-10)x(5-5.5) um and large microbasic euryteles 
(21-23)x(7.5-8.5)um. Gastrozooids have microbasic euryteles (7.5-8.5)x(3.5)um and 
desmonemes (3.5-4)x(2-3)um. 


BIOLOGY: Grows on gastropod shells either inhabited by the mollusc or hermit 
crabs (Allman, 1864b; Weismann, 1883; Motz-Kossowska, 1905; Russell, 1957; 


246 P. SCHUCHERT 


Fic. 19 


Dicoryne conybearei (Allman, 1864). (A) Stems with hydranths and blastostyles bearing 
sporosacs, scale bar 0.5 mm. (B) Stem perisarc magnified, note corrugation, scale bar 0.05 mm. 
(C) Young female sporosac, scale bar 0.05 mm. (D) Liberated sporosac. All after living material, 
A-C Mediterranean material; D, New Zealand. 


Bavestrello, 1985; Schuchert, 1996). Likely also epizoic on spines of cidaroid sea- 
urchins. In the Mediterranean, mature colonies were observed from March to the end 
of autumn (Weismann, 1883; Motz-Kossowska, 1905; Ashworth & Ritchie, 1915), in 
the British Isles at least from August to October (Allman, 1872; Russell, 1957). 


DISTRIBUTION: North-eastern Atlantic from Ireland and English Channel to 
Iberian Peninsula (e. g. Allman, 1864b; Philbert, 1935; Teissier, 1965; Medel & Löpez- 
Gonzälez, 1996); widespread in the Western Mediterranean (Weismann, 1883; Motz- 
Kossowska, 1905; Bavestrello, 1985). The northernmost record is perhaps Oban, 
Scotland (Hincks, 1868), though this identification is uncertain. Also recorded from 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 247 


Japan (Hirohito, 1988) and New Zealand (Schuchert, 1996). Type locality: Ireland, 
Cork, Glengariff Harbour, on hermit crab shells. 


REMARKS: As discussed above, Dicoryne conybearei could be only a variety of 
D. conferta. The species is diagnosed through its sporosac with one egg, although 
occasionally some sporosacs contain two eggs (observed in 2 of 20 sporosacs of one 
colony). There is usually only one tentaculoid present, but this trait is even more 
variable (Ashworth & Ritchei, 1915; Schuchert, 1996). 

The type material of Perigonimus (?) nanellus Stechow, 1919 was re-examined 
for this study and it became evident that this must be a Dicoryne species. The colony 
has some tentacle free polyps, these are also depicted Stechow (1919), which have a 
cap of large heteronemes at their tips. It is quite obvious that these are identical to 
blastostyles of Dicoryne, but without the sporosacs. These have either been spent or not 
developed yet. Moreover, the perisarc has the same irregularly corrugated structure and 
also the size of the hydranths matches well. However, in the absence of sporosacs it is 
impossible to attribute it either to D. conferta or D. conybearei. Dicoryne conybearei 
appears more plausible, as this is the more frequent of the two in the Mediterranean. 
The only unusual feature of Perigonimus nanellus is its occurrence on sea urchin 
spines. 


Genus Bimeria Wright, 1859 


TYPE SPECIES: Bimeria vestita Wright, 1859b by monotypy. 
SYNONYM: Manicella Allman, 1859a: 51. 


DIAGNOSIS: Colonies stolonal or erect, with perisarc enveloping hydranth and 
extending as a sheath over proximal ends of tentacles. Hydranth ovoid to vasiform; 
hypostome dome-shaped; tentacles in single whorl or in two close whorls. Gonophores 
fixed sporosacs on hydrorhiza and branches. 


REMARKS: See Calder (1988) for taxonomic details concerning this genus. 


Bimeria vestita Wright, 1859 Fig. 20 


Bimeria vestita Wright, 1859b: 109, pl. 8 fig. 4. — Hincks, 1868: 103, pl. 15 fig. 2. — Allman, 
1872: 297, pl. 12 figs 1-3.— Hamond, 1957: 297, figs 3-4. — Millard, 1975: 95, fig. 32C- 
H. — Calder, 1988: 21, figs 17-18, synonymy. — Hirohito, 1988: 94, fig. 33d-f, 34a. — 
Ramil & Vervoort, 1992: 14, bibliography. — Migotto, 1996: 9, figs 2a-b. - Genzano & 
Zamponi, 1999: 63, figs 4-5. — Marques er al., 2000: 322, figs 1-3. — Bouillon et al., 
2004: 42, fig. 25A-C. — Vervoort, 2006: 196, fig. 5.2. 

Eudendrium vestitum Allman, 1888: 3, pl. 1 figs 1 & la. — Marques er al., 2000: 322, synonym. 

Manicella fusca Allman, 1859a: 51. — Hincks, 1868: 104, synonym. — Calder, 1988: 22, 
synonym. 

Bimeria humilis Allman, 1877: 8, pl. 5 figs 3-4. — Marques et al., 2000: 322, synonym. — Fraser, 
1944: 49, fig. 17. 

? Bimeria amoyensis Hargitt, 1927: 492, fig. 1. — Hirohoto, 1988: 96, ? synonym. 

Bimeria vestita f. nana Leloup, 1932: 142, fig. 14. 

Bimeria corynopsis Vanhöffen, 1910: 287, fig. 12. 

Bimeria (? Garveia) humilis. — Wedler & Larson, 1986: 89, fig. 9B. 

not Bimeria vestita. — Annandale, 1907: 141, fig. 3 [= Garveia franciscana (Torrey, 1902)]. 


MATERIAL EXAMINED: Roscoff, 8 June 2000, infertile colonies, all used to extract DNA, 
16S sequence AM183130. - MHNG INVE32957, Banyuls-sur-Mer, 11 May 2002, 2 m, on 
algae, infertile. - MHNG INVE35920 Spain, Alboran Sea, 36.522°N 2.837°E, 107, 12 April 


248 P. SCHUCHERT 


Fic. 20 


Bimeria vestita Wright, 1859. (A) Silhouette of colony, scale bar 5 mm. (B) Part of stem with 
side branch bearing a hydranth, note sheathed tentacles, scale bar 0.2 mm. (C) Optical section of 
stem perisarc with annulated branching point. (D-F) Male sporosacs of different developmental 
stages, last stage is likely mature, note branched spadix, scale bar 0.1 mm. (G-I) Developmental 
stages of female sporosac, last stage likely with planula, scale bar equals 0.1 mm. A-C, after 
preserved material from the Mediterranean; D-F, after preserved material from Devon, England; 
G-I modified after Hirohito (1988). 


2004, leg. H. Zibrowius, infertile, on Lafoea dumosa. — IRSNB, coll. J. Bouillon Roscoff, 
Brittany, 1964, small infertile colony on Abietinaria abietina. - BMNH 1957.3.2.122, England, 
Devon, Start Bay, 20 August 1898, with male sporosacs, growing on Abietinaria abietina, slide 
preparation. - BMNH 1957.3.2.123, Bay of Biscay, 07 March 1916, 137 m, infertile, on 
Sertularella spec., slide preparation. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 249 


DIAGNOSIS: Branching hydroid colony, proximal part of tentacles ensheathed 
with soft perisarc, gonophores sporosacs. 


DESCRIPTION (after examined material and literature): Colonies erect, mono- 
siphonic, arising from creeping tubular stolons. Shoots branched, when fully grown 
with central main axis and shorter side-branched, irregularly pinnate with alternate 
branches, all ends with hydranths; stem and branches covered by firm perisarc, single- 
layered, smooth or annulated over stretches, frequently annulated or spirally corru- 
gated at the origin of side branches, perisarc continued on hydranths as soft pseudo- 
hydrotheca, covering also hypostome and forming tubular sheaths around basal part of 
tentacles, perisarc of hydranth encrusted with detritus and fine sand. Hydranth vasi- 
form, merging below almost imperceptibly into pedicel, pedicels quite long; hypo- 
stome conical; 10-12 tentacles (reportedly up to 16) in one whorl or two very close 
whorls, amphicoronate. Gonophores fixed sporosacs on stem and branches, ovoid to 
oblong, without radial canals, enveloped in loose filmy perisarc infested with detritus; 
spadix in younger sporosacs irregularly branched or with irregular lateral processes, 
female with sporosac with one egg only, planula developing in situ, colonies likely 
gonochoristic. Colour: perisarc yellowish. Nematocysts: microbasic euryteles, and 
desmonemes. 


DIMENSIONS: Stems from a few mm to 25 mm, stem diameter 60-80 um; 
hydranth 0.25-0.5 mm high; sporosac up to 0.35 mm long and 0.2 mm wide (incl. 
perisarc covering); egg size 0.2 mm (Millard, 1975). Microbasic euryteles (6.5-7)x(3- 
3.5) um; desmonemes 3 x 2 um. More measurements are given in Millard (1975), 
Calder (1988), Genzano (1992), and Migotto (1996). 


DISTRIBUTION: Bimeria vestita is reportedly a cosmopolitan species (Ramil & 
Vervoort, 1992), but it is unlikely to occur in cold and Arctic waters. Along the 
European coasts it has been reported from Scotland, western and eastern England, the 
English Channel, the southern North Sea, Bay of Biscay, the Iberian Peninsula, and the 
Mediterranean (e. g. Allman, 1872; Motz-Kossowska, 1905; Browne, 1907; Hamond, 
1957; Teissier, 1965: Fey, 1970; Patriti, 1970; Ramil & Vervoort, 1992; Medel & 
Löpez-Gonzälez, 1996; Faasse, 2003; this study). Scotland seems to be its northern 
limit. It has also been reported from the Black Sea (Manea, 1980); western and 
southern Africa (Leloup, 1937; Buchanan, 1957; Millard, 1975; Vervoort, 2006), 
Argentina (Genzano & Zamponi, 1999); western Atlantic (Fraser, 1944, as Bimeria 
humilis; Calder, 1988; Vervoort, 1968), Indian Ocean (Mammen, 1963; Millard, 1975), 
western Pacific (Leloup, 1932; Hirohito, 1988); eastern Pacific (Fraser, 1938). Type 
locality: Firth of Forth, Scotland. 


BIOLOGY: Occurs frequently on hydroids, but also algae, pebbles and other 
unspecific hard substrata in depths of 0-200 m. Aspects of its ecology in Argentinean 
waters were investigated by Genzano & Zamponi (1999). They found that the colonies 
were present all year round and they produced gonophores in the austral summer. In 
the NE Atlantic, the fertility period is likely also early summer, but here are not enough 
data available. In the Mediterranean (Boero & Freesi, 1986), reproductive colonies oc- 
cur from October to May, they are found frequently on Eudendrium species in 10-20 
m, and the reproduction takes place from October to February. 


250 P. SCHUCHERT 


REMARKS: The synonymy and history of this species have been treated by 
Calder (1988) and Marques er al. (2000a). Hirohito (1988) thought that also Bimeria 
amoyensis Hargitt, 1927 could be a synonym with this species. The type material of 
this species must be re-examined to come to a reliable conclusion. Hirohito (1988) in- 
cluded also Bimeria corynopsis Vanhöffen, 1910, an Antarctic species, in the syn- 
onymy of B. vestita. Although Vanhöffen’s material was infertile and thus not reliably 
identifiable, he kept it apart from Bimeria vestita on account of the swollen tentacle 
tips. The biogeographic differences argue also in favour of the two being separate 
species. 

With its ensheathed tentacles this is a very characteristic species. Koellikerina 
fasciculata, a species with a medusa phase, has a similar hydroid (Fig. 13F). 


Genus Garveia Wright, 1859 


TYPE SPECIES: Garveia nutans Wright, 1859b. 

SYNONYMS: Corythamnium Allman, 1859a [type species Eudendrium bacciferum 
Allman, 1859a = G. nutans]. — Calyptospadix Clarke, 1882 [type species Calyptospadix cerulea 
Clarke, 1882]. — Pruvotella Motz-Kossowska, 1905 [type species Pruvotella grisea Motz- 
Kossowska, 1905]. 


DragnosIs: Colonial hydroids with branched stems, monosiphonic or poly- 
siphonic; hydranth fusiform, hypostome dome-shaped or conical, usually one distal 
whorl of tentacles, rarely two; pseudohydrotheca covering polyp base but not 
ensheathing tentacles; gonophores fixed sporosacs, borne on pedicels on hydrocauli or 
on hydrorhiza. 


REMARKS: The genus Perigonimus M. Sars, 1846 is sometimes still incorrectly 
used as a synonym of Garveia Wright, 1859 or Rhizorhagium M. Sars, 1874. Because 
the type species of Perigonimus — P. muscoides — is currently placed in the genus 
Bougainvillia as B. muscoides, Perigonimus becomes a synonym of Bougainvillia (see 
Rees, 1938). 

The Garveia species are quite heterogeneous and the genus is unlikely a clade. 
But following most recent authors and for the sake of nomenclatural stability, 
Pruvotella grisea Motz-Kossowska, 1905 is here also kept in the genus Garveia, 
despite several features of this species might warrant to place it in a separate genus: the 
double layered perisarc, the somewhat unusual sporosacs with high, vacuolated cells, 
the adnate side-branches, the large heteroneme, and the extraordinarily contractible 
hydranth body and tentacles. See also the discussion under Rhizorhagium M. Sars, 
1874. 


KEY TO THE GARVEIA SPECIES TREATED HERE: 


la mature colonies:monosiphonic .........2...1: = RS SSA SSA 2 
lb mature.colonies.polysiphonic......._ ili 3 
2a several sporosacs clustered on hydranth pedicels, branches not adnate 

PES TA TRS > EE NA ne te oi Ii Garveia franciscana 
2b sporosac large and solitary, branches adnate ...................... Garveia grisea 
3a branches adnate, perisarc double layered, tentacles in several whorls . . . Garveia grisea 
3b branches not adnate, perisarc single layered, one whorl of tentacles.............. 4 
4a hypostome conical, broadly arborescent, deep water species ........ Garveia arborea 
4b hypostome rounded to spherical, colonies elongate, shallow water species 


SITR ERB dei RL EA RES AT A eu) IRR Ee ee UE Garveia nutans 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 251 


Garveia nutans Wright, 1859 Fig. 21 


Garveia nutans Wright, 1859b: 109, pl. 7, fig 5. — Hincks, 1868: 102, pl. 14 fig. 4. — Allman, 
1871: 44, fig. 15B. — Allman, 1872: 295, 4, pl. 12 figs 4-11.- Philbert, 1935: 18.— Fraser, 
1937: 36, pl. 5 fig. 22. — Leloup, 1952: 19, fig. 57. - Ramil & Vervoort, 1992: 15. — 
Bouillon et al., 2004: 46, fig. 27F-G. 

Eudendrium bacciferum Allman, 1859a: 53. — Hincks, 1868: 102, listed as synonym. 


MATERIAL ExAMINED: BMNH 1948.9.8.52; Great Britain, Plymouth; collected 1898, 
infertile. - BMNH 1969.11.29.1-3; Great Britain, Menai Bridge, Church I; 22 May 1963; fertile 
female. - BMNH 1985.9.1.22; Great Britain, Plymouth; May 1936; fertile female, no hydranths. 
— BMNH 1957.3.2.170 slide; Great Britain, Plymouth; 17 October 1898; fertile female. —ZSM, 
as Bimeria nutans, Plymouth, slides 20000453, 20000454, and 20000455, Stechow collection, 
material mentioned in Stechow (1919), fertile male. — The Faroes 62.317°N 06.833°W (BIOFAR 
station 370); 66 m; 25 July 1988, infertile. 

DIAGNOSIS: Hydroid colony with polysiphonic stem, stem tubes more or less 
equal: Hydranths yellow, broad, hypostome very large and often spherical, one whorl 
of tentacles, pseudohydrotheca present, gonophores large sessile sporosacs resembling 
a medusoid, mature without radial or circular canal, spadix present, large subumbrellar 
cavity, females with several eggs. 


DESCRIPTION: Colonies elongate, with conspicuous central main axis and 
thinner, short, straggling side branches, main stem strongly polysiphonic, thinning out 
to monosiphonic, side branches mostly monosiphonic. Stolons creeping, ramified. 
Perisarc usually one-layered, smooth or irregularly corrugated or wrinkled, expanding 
at distally as filmy pseudohydrotheca onto hydranth body, not infesting tentacles. 
Hydranths at end of long branches, rather broad, urn-shaped, reminiscent of the genus 
Eudendrium, hypostome very large, up to 1/3 of total hydranth height, spherical to high 
dome-shaped, one whorl of 10-14 tentacles, these evenly covered by nematocysts, 
gastrodermis chordoid. Gonophores arising from main stem, ovoid, on tapering pedicel 
of about same length, pedicel covered by conical perisarc sheath that gets gradually 
thinner towards distal. Gonophores fixed sporosacs, in younger stages reminiscent of 
medusoids, oval shape, mature without radial canals, circular canal, or velum, with 
spadix projecting into spacious subumbrellar cavity. Females with up to 10 eggs, these 
relatively large, sporosac not covered by perisarc film in older stages, eggs developing 
in situ, thus larviparous. Colours: hydranths and eggs have an intense yellow-orange 
colour, may be red, tentacles yellow. Disturbed hydranths bend themselves towards 
below (see Fig. 21B, lowest hydranth). 


DIMENSIONS: Colony up to 3 cm, polyps up to 0.8 mm high, diameter 0.35 mm. 
Diameter of distal branches 0.16 mm. Sporosacs 0.9 mm long, diameter 0.65 mm. Egg 
size ca. 0.25 mm. 


ADDITIONAL DATA: The Gonophores possess rudimental radial canals during 
their early developmental stages (Allman, 1872). 


BIOLOGY: Occurs at sites with strong or very strong tidal streams, attached to 
stones, bed-rock, hydroids, seaweeds and other hard substrata. Mature February to 
October, but perhaps also in other months. Usually in shallow water of a few metres 
depth (laminarian zone) and even at the low water mark, but there is also a record from 
580 m (Ramil & Vervoort, 1992). 


252 P. SCHUCHERT 


Fic. 21 


Garveia nutans Wright, 1859; after preserved material from Plymouth. (A) Colony silhouette, 
scale bar 5 mm. (B) Part of colony, note pseudohydrothecae on hydranth bodies, scale bar 1 mm. 
(C) Optical section of hypostome, note its spherical shape, scale bar 0.1 mm. (D) Perisarc of 
hydranth pedicel, note transition of corrugated to smooth perisarc, scale bar 0.1 mm. (E) Young 
male sporosac in optical section, scale bar 0.1 mm. (F) Mature male sporosac in optical section, 
note spadix with spermatids (dark) projecting onto spacious cavity (subumbrella), same scale as 
E. (G) Mature female sporosac with developing embryos, scale bar 0.2 mm. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 253 


DISTRIBUTION: Great Britain, The Faroe Islands, English Channel, Belgium, Bay 
of Biscay, Strait of Gibraltar. (Allman, 1872; Browne, 1907; Philbert, 1935; Leloup, 
1952; Hammond, 1957; Russell, 1957; Cabioch, 1976; Ramil & Vervoort, 1992). The 
record from Alaska (Nutting, 1901) is somewhat doubtful. Type locality: Inch Garvie, 
Firth of Forth, Shetland Islands, United Kingdom. 


REMARKS: With its straggling colonies and yellow hydranths having spherical 
hypostomes, Garveia nutans Wright, 1859 can easily bee mistaken for a Eudendrium 
species. However, the pseudohydrotheca and the sporosacs immediately distinguish it 
from the genus Eudendrium. 


Garveia franciscana (Torrey, 1902) Fig. 22 


Bimeria franciscana Torrey, 1902: 28, pl. 1 fig. 4. — Weill, 1934a: 77, fig. 71a-b. — Weill, 1934b: 
386. — Fraser, 1937: 31, fig. 14. 

Bimeria Monidi Billard, 1927: 467, figs 1-2a. — Vervoort, 1964: 125, synonym. 

Bimeria tunicata Fraser, 1943: 76, pl. 15 fig. 2. — Fraser, 1944: 5, fig. 18. — Vervoort, 1964: 125, 
synonym. 

Perigonimus megas Kinne, 1956: 257, figs 1-7. — Vervoort, 1964: 125, synonym. 

in part Cordylophora lacustris — Pennycuik, 1959: 165, pl. 2 figs 4-5. 

[not Cordylophora lacustris Allman, 1844] 

Garveia franciscana. — Vervoort, 1964: 127, figs 1-4. — Morri, 1981: 53, fig. 16, pl. 1 fig. 2. — 
Morri, 1982: 381, figs 1-5, pl. 1. — Barnes, 1994: 62, fig. — Calder & Mayal, 1998: 73. — 
Schönborn er al., 1993: 220, pl. 4 fig. 2. — Bouillon et al., 2004: 45, fig. 27B-D. 


MATERIAL EXAMINED: ZSM, slide 20000536, The Netherlands, Zuiderzee, Vuile Gat, 
WN 77,29 September 1926, ex. Zool. Mus. Amsterdam, det W. Vervoort, fertile female colonies. 
— BMNH 1966.10.8.30 and 37, 39, 40; Nigeria, Port Harcourt; 10 September 1957; fertile, 
several male and female colonies from raft, also 3 slides. - BMNH without registration number; 
Germany, Kiel-Kanal, near Rendsburg, 13 September 1961, male and female colonies; collected 
Hjalmar Thiel, labelled as Perigonimus megas.— BMNH 1981 .11.4.3 alcohol plus slide material, 
as Dicoryne conferta, Alexandria, Egypt, 20 November 1980, leg. A. Shoukr, not well preserved, 
5 cm polysiphonic stems, sporosacs on hydranth pedicels, likely belongs to Garveia franciscana. 


DIAGNOSIS: Occurring in brackish waters, colonies branching, tall, mono- 
siphonic; one whorl of tentacles; several sporosacs on hydranth pedicels, one egg per 
sporosac, larviparous. 


DESCRIPTION: Occurs only in brackish waters, colonies erect, monosiphonic 
throughout, much branched, bushy to irregularly pinnate, branches long; root-like 
hydrorhiza. Branches arise at an angle of about 60°, not adnate. Perisarc one-layered, 
smooth or irregularly corrugated, at origin of side-braches an annulated stretch of 
variable length. Hydranth body covered by thin pseudohydrotheca; hydranth fusiform, 
hypostome conical, 8-12 filiform tentacles in one whorl. Colonies are dioecious, 
sporosacs concentrated on hydranth pedicels, usually several ones present, up to 20 
possible. 

Sporosacs ovoid, enveloped in filmy perisarc, in advanced stages loose. 
Sporosacs without radial or circular canals, no velar opening, spadix present. Female 
sporosacs with usually one egg only, spadix remains below egg, egg develops in situ 
to planula (larviparous). Male sporosacs slightly larger, spadix present, spermatozoa 
develop on spadix. 

Nematocysts: desmonemes and microbasic euryteles. 


254 P. SCHUCHERT 


Fic. 22 


Garveia franciscana (Torrey, 1902). (A-B) Colony silhouettes, scale bar 1 cm. (C) Branch with 
hydranth and female sporosacs, some sporosacs broken off, scale bar 0.2 mm. (D) Branch with 
hydranth and male sporosacs, same scale as C. All after preserved material, A & C from Holland, 
B & D from Nigeria. 


DIMENSIONS: Colony heights 1-20 cm; branch diameters 0.15-0.3 mm; hydranth 
about 0.5 mm high; female sporosacs about 0.3 mm long, diameter 0.2 mm. 
Desmonemes 4 um long, microbasic euryteles 7 um (Morri, 1982). 


BIOLOGY: Tolerates a wide range of salinity, but occurs only in brackish waters 
in very shallow depths down to a few meters, can withstand elevated temperatures of 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 255 


above 30°C, but regresses below 9°C (Crowell & Darnell, 1955). Morri (1982) gives a 
preferred salinity range of 20-30 ppt. Schönborn et al. (1993, Baltic Sea) found it in 
1-18 ppt salinity, mostly below 6 ppt; reproduction was observed from June to 
September if temperatures remained above 14°. The polyps fed on copepods and 
oligochaetes. The hydranths remain smaller in reduced salinities. 


DISTRIBUTION: California; Atlantic coast of America from Brazil up to Virginia; 
east and west coasts of India; Andaman Sea; west coast of Africa; eastern Australia; 
Holland (now extinct), North Sea; Baltic Sea; Mediterranean lagoons (Fraser, 1937; 
Vervoort, 1964; Morri, 1982; Schönborn et al., 1993; Barnes, 1994; Calder & Mayal, 
1998). Type locality: San Francisco, California, intertidal. 


REMARKS: The synonymy of this species has been worked out by Vervoort 
(1964) and his view is here adopted. Garveia franciscana can easily be confounded 
with Pachycordyle navis (Millard, 1959), a species also occurring in brackish waters 
(see Schuchert, 2004 and Fig. 29). The latter, however, has hydranths with tentacles in 
several whorls (though sometimes close set), forms smaller colonies, has often a 
bi-layered perisarc, the sporosacs are not clustered on the hydranth pedicels, and the 
female sporosacs contain several eggs (comp. Fig. 29C). 


Garveia arborea (Browne, 1907) Fig. 23 


Bimeria arborea Browne, 1907: 15, pl. 1 figs 1-3, pl. 2. 
Garveia arborea. — Vervoort, 1985: 273, pl. 2. — Ramil & Vervoort, 1992: 15. 
? Bimeria arborea. — Hirohito, 1988: 94, fig. 331a-c. 


MATERIAL EXAMINED: Holotype, BMNH 1948.10.1.117, as Bimeria arborea, alcohol and 
slide preparations; Bay of Biscay; 752 m; August 1906; fertile. - MHNG INVE36661 (part of 
CARACOLE collection station KGS 11); 53.779°N 13.938°W (west of Ireland), 793 m, 
collected by Usnel corer; date 9 August 2001; on mud/polychaete tube; fertile male. 


DIAGNOSIS: Deep-sea Garveia species with polysiphonic, irregularly arbo- 
rescent colonies, central main tube larger than peripheral auxiliary tubes; hydranths 
with small, conical hypostome, tentacles in one whorl; sporosacs small, not in clusters, 
on stem, branches and rarely on hydranth pedicels and stolons; males without vestige 
of subumbrellar cavity. 


DESCRIPTION: Colonies erect, polysiphonic, branching tree-like, composed of a 
branching, central main tube covered by many thinner, parallel auxiliary tubes, number 
of auxiliary tubes decreasing towards distal until only main tube is left at distal branch 
tips, main tube of branches originates from main tube of stem, perisarc of main tube 
and auxiliary tubes mostly smooth. Stolons tubular, root-like or creeping. Hydranths 
arise from the main tube on stems and branches, with pedicels of variable length, 
perisarc smooth, corrugated or wrinkled, continued over hydranth body as filmy 
pseudohydrotheca terminating below tentacles, detritus infested. Hydranths typical for 
Bougainvilliidae, spindle shaped, slender, with short conical hypostome, one whorl of 
8-12 filiform tentacles, strongly contractible. Gonophores develop on stems and 
branches, issuing from the main tube, occasionally also on hydranth pedicels and even 
stolons. Gonophores are sessile sporosacs of the cryptomedusoid type. Male sporosacs 
ovoid to spherical, tapering proximally into a short pedicel; mature sporosacs with a 


256 P. SCHUCHERT 


Fic. 23 


Garveia arborea (Browne, 1907); after preserved material (MHNG INVE36661). (A) Colony 
silhouette, scale bar equals 5 mm. (B) Hydranth, scale bar equals 0.1 mm. (C) Male sporosac, 
scale bar equals 0.1 mm. (D) Cross-section of a branch, note central primary tube, the dotted 
region around the main tube represents detritus particles, scale bar equals 50 um. 


thickening of distal epidermis; simple central spadix covered by a thick layer of germ 
cells filling cavity entirely, this round mass of gametes with four shallow embayments 
at its base. At least in younger sporosacs four short, pouch-like vestiges of radial 
canals. Female sporosacs unknown. Nematocysts: larger heteronemes (likely micro- 
basic euryteles) on sporosacs and in coenosarc; smaller microbasic euryteles and 
desmonemes abundant on tentacles. 


DIMENSIONS (after Browne, 1907; Vervoort, 1985; own data): Stems 1-3 cm 
(max. 8.75 cm), basal stem diameter up to 3 mm (Vervoort, 1985); main tube 0.1-0.25 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE DS 


mm, auxiliary tubes 0.05-0.1 mm; hydranth pedicels up to 1 mm, hydranth body up to 
0.2-0.25 mm high; sporosacs 0.2-0.3 mm. Larger heteroneme (6.5-8.5)x(3.5-4)um, 
smaller microbasic euryteles (5-6.5)x(2.5-3)um, desmonemes (3.5-4.5)x(2.5-3)um. 


BIOLOGY: A deep water species, known from depths of 420-3100 m. Can grow 
on solid substrata like polychaete tubes worms (this study) or anchored in soft sub- 
strates by a root-like hydrorhiza (Ramil & Vervoort, 1992). 


DISTRIBUTION: Continental margins from west of Ireland to Strait of Gibraltar 
(Browne, 1907; Vervoort, 1985; this study), perhaps also Somalia (Stechow, 1925; as 
B. crassa), South Africa (Millard, 1977; as G. crassa), and Japan (Hirohito, 1988). 
Type locality: Bay of Biscay, 48°7’N 8°13’W, 412 fathom (= 752 m), sand, mud and 
hard ground (Browne, 1907). 


REMARKS: This is a rare, deep water species. Hirohito’s (1988) identifications 
are based on infertile material and should therefore be considered with some reser- 
vations (see also Vervoort, 2006). 

Vervoort (1985) suggested that Garveia crassa (Stechow, 1923b) could be 
conspecific with G. arborea. A re-examination of the type material of Bimeria crassa 
(ZSM 20000436 & 20000437, slides) indicated that this species has different 
gonophores. The gonophores in the slide material are not fully mature and no gametes 
could be seen. They are oblong, usually about two times as long as wide, thus clearly 
different from those of G. arborea. Furthermore, their internal structure appears also 
different. The gastrodermal layers form a cup-like structure and no spadix is present 
(although the lumen of the cup-like structure feigns a spadix). If these differences are 
not due to a sexual dimorphism, then it is unlikely that B. crassa is conspecific with G. 
arborea. 

The presence of stenoteles could not be confirmed in the available material, this 
in contradistinction to Vervoort (1985). 

Concomitantly with Bimeria arborea, Browne (1907) also described the simi- 
lar Bimeria biscayana. The latter species produces medusae and was transferred 
provisionally to the genus Amphinema (see below). Fertile material of both species is 
easily separable, but also infertile material has a different habitus and in B. arborea the 
main tubes of the branches originate from main tubes of the stem. 


Garveia grisea (Motz-Kossowska, 1905) Figs 24-25 


Pruvotella grisea Motz-Kossowska, 1905: 77, fig. 9, pl. 3 figs 10-15. 

Bimeria fragilis Stechow, 1919: 28, fig. E. — Picard, 1958:190, synonym. 

Garveia grisea. — Picard, 1951: 339. — Morri, 1982: 389, fig. — Boero & Freesi, 1986: 140. — 
Morri et al., 1991: 31.— Medel & Löpez-Gonzälez, 1996: 193. — Bouillon et al., 2004: 
46, fig. 27E. 


MATERIAL EXAMINED: Syntype series of Bimeria grisea, ZSM, microslides 20000442, 
20000443, 20000438, 20000439, Stechow, 1919. -MHNG INVE34436, France, Calanque Port 
d’Alon near Bandol, 22 April 2003, 0-2 m depth in shaded places and caves, examined alive, 
several infertile colonies up to 2 cm height, 16S sequence AM183131. — Ligurian Sea, 
Villefranche-sur-Mer 5 April 2005, 6 infertile colonies, some cultivated for 3 weeks, hydranths 
regenerated well although they did not feed on Artemia, no gonophores. 


DiaGNOSIS: Hydroid forming branching colonies, polysiphonic or not, double- 
layered perisarc, branches adnate, hydranths very contractile, with macrobasic 


258 P. SCHUCHERT 


Fic. 24 


Garveia grisea (Motz-Kossowska, 1905). (A) Colony silhouette, note that in shallow waters the 
colony can be stolonal, scale bar 5 mm. (B) Optical section of a contracted hydranth, note the 
collar-like membrane (co) into which the tentacles can retract, the pseudohydrotheca (ph), and 
the double layered perisarc (ps); scale bar 0.1 mm. (C) Strongly contracted cultivated hydranth, 
note large heteronemes on the bulging hydranth-wall, scale bar equals 0.1 mm. (D) 
Undischarged nematocysts, in the upper row the large macrobasic heteronemes in two views, 
lower row microbasic eurytele and desmoneme, scale bar equals 10 um. A-B, after preserved 
material; C-D, after life. 


heteronemes, tentacles can be in two whorls or scattered. Gonophores large styloid 
sporosacs with one egg and thick vacuolated epidermis; larviparous. 


DESCRIPTION: Colonies branching or not, sometimes stem and major branches 
polysiphonic, but stolonal colonies occur also. Stolons creeping, ramified. Side- 
branches adnate for some distance, perisarc in older parts double layered, inner layer 
thicker and undulated, outer layer filmy, the latter continued onto hydranth body as 
pseudohydrotheca, both layers connected by lamellar membranes. Hydranth spindle 
shaped, hypostome dome-shaped, in colonies from nature 7-12 tentacles in two whorls, 
distance of whorls variable, some hydranths with tentacles scattered in upper part of 
hydranth. The hydranth can contract very strongly, so that the tentacles disappear in a 
collar-like epidermal fold (seen in preserved material) or the hydranth gets very short 
and the body-wall forms a bulge. Colour of hydranths: milky-white. 

Gonophores relatively large, developing on stem; fixed sporosacs, oblong 
ovoid, broader end distal; with distinct pedicel. Pedicel covered by firm perisarc, 
gonophore enveloped by filmy perisarc. Structure of sporosac of styloid type, but 
atypical and complex. Epidermis very thick, composed of high, vacuolated cells, below 
these vacuolated cells a sac-shaped inner lining. Mature females contain inside this 
second lining an ovoid egg or planula enveloped by two layers of cells. Young female 
sporosacs with one oocyte in epithelium of inner cell layer. Larviparous. Male 
sporosacs with spermatogenesis in gastrodermal layer. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 259 


Fic. 25 


Garveia grisea (Motz-Kossowska, 1905), after type material of Bimeria fragilis (see page 260), 
scale bar equals 0.1 mm, valid for all figures. (A) Stem with side-branch, note adnate base of 
branch and double layered perisarc. (B) Semi-relaxed hydranth, same scale as A. (C) Young 
female sporosac. (D) Fully grown female sporosac, note the large vacuolated cells. 


260 P. SCHUCHERT 


Nematocysts: (1) microbasic euryteles, shaft discharged about 0.8-0.9 times as 
long as capsule, abundant on tentacles. (2) desmonemes, abundant on tentacles. (3) 
macrobasic heteroneme, on hydranth body, shaft in with 3-4 coils, coiling pattern vari- 
able, not seen discharged. 


Dimensions (Motz-Kossowska, 1905; own observations): Colonies 3 mm in 
shallow waters, up to 3 cm in deeper waters. Diameter of branches 0.1-0.13 mm. 
Hydranth about 0.3 mm. Female sporosacs 0.7 to 1 mm long and 0.4-0.6 mm wide. 
Nematocysts (living material): microbasic euryteles, (8-10)x(4.5-5.5) um, shaft 
discharged about 0.8-0.9 times as long as capsule, abundant on tentacles; desmonemes, 
(4-5)x(2.5-3.0) um, abundant on tentacles; macrobasic heteroneme, on hydranth body, 
shaft in with 3-4 coils, coiling pattern variable, not seen discharged, (19-21)x(8-9.5) 
um. 

ADDITIONAL DATA: Motz-Kossowska (1905) describes the development of the 
female sporosacs. Already Motz-Kossowska (1905) noted that the hydranths of 
Garveia grisea are able to contract very strongly. This contraction can lead to the 
formation of a collar-like epidermal membrane that covers and hides the tentacles (Fig. 
24B). The living material observed for this study confirmed this extraordinary 
contractility. The collar like membrane, however, was not always observed. Its 
presence may depend on the degree of contraction of the hydranth. The hydranths of 
Garveia grisea are very sensitive and they deteriorated very rapidly in freshly collected 
material. It was very difficult to get them relaxed with 7% MgCl, solutions. Despite 
their initial sensitiveness, they regenerated well in culture, though not the pseudo- 
hydrotheca. Surprisingly, in the regenerated hydranths the tentacles grew in one whorl 
and not scattered in a narrow band as was often seen in hydranths collected in the sea. 
If disturbed — e. g. by squeezing a tentacle by tweezers — the hydranth contracts 
strongly so that the body forms a ring-like bulge (Fig. 24C). The large heteroneme 
capsules on the hydranth body are thus brought into an exposed position. It is very 
suggestive to regard this as a defensive reaction. The large heteroneme could not be 
identified more precisely as their discharge could not be induced, this despite numerous 
native capsules were examined. 


DISTRIBUTION: Western Mediterranean*. Type locality: Mediterranean, France, 
Banyuls-sur-Mer. 


BIOLOGY: A winter species, being present from September to May, occurs along 
rocky shores in depths of 0-80 m, reproduction: September to November (Motz- 
Kossowska, 1905), or October-January (Boero & Freesi, 1986). Occurs often on large 
Eudendrium colonies. 


SYNONYMY AND TAXONOMY: Picard (1958) synonymized Bimeria fragilis 
Stechow, 1919 with Garveia grisea without further discussion. A re-examination of the 
type material of Bimeria fragilis Stechow, 1919 confirmed that it is indistinguishable 
from G. grisea in nearly all aspects, except for the contracted hydranths. The type 
material of B. fragilis showed moderately expanded hydranths identical to those 
observed in cultivated material of G. grisea. I suspect that Stechow has either treated 
his material with muscle-relaxing chemicals or he had colonies with regenerated 


* Note added in proof: also West of Ireland, MHNG INVES3151. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 261 


hydranths. Because B. fragilis is otherwise identical to G. grisea, notably sharing the 
same unique sporosacs, the double-layered-perisarc, the colony form, the adnate side- 
branches, and the large supplementary nematocysts, it is here regarded as conspecific 
with G. grisea. 


REMARKS: Garveia grisea is one of the few bougainvilliid hydroids that can al- 
so be identified in the absence of gonophores. The adnate branches, the double-layered 
perisarc, the strongly contractile hydranths, and the large heteroneme make it distinct 
from all other bougainvilliids. The colonies resemble quite Turritopsis dohrnii, with 
which they can easily be confounded. However, the hydranths of the latter species are 
not as contractile, they have their tentacles scattered over most of the hydranth body, 
and they never have a pseudohydrotheca (see Schuchert, 2004). 


Genus Rhizorhagium M. Sars, 1874 


TYPE SPECIES: Rhizorhagium roseum M. Sars, 1874 by monotypy. 

SYNONYMS: Wrightia Allman, 1872 [type species Arractylis arenosa Alder, 1862 by 
monotypy; invalid junior homonym of Wrightia L. Agassiz, 1862]. — Gravelya Totton, 1930 
[type species is Perigonimus antarcticus Hickson & Gravely, 1907 by designation of Totton, 
1930]. — Aselomaris Berrill, 1948 [type species Atractylis arenosa Alder, 1862, designation by 
Calder, 1988]. 

DIAGNOSIS: Hydroids forming stolonal colonies, hydranths on perisarc covered 
hydrocauli, rarely some shoots with one or two side branches; hydranth fusiform; 
hypostome dome-shaped, conical, or nipple-shaped, surrounded by one distal whorl of 
tentacles; pseudohydrotheca covering polyp base but not sheathing tentacles; 
gonophores fixed sporosacs, borne on hydrocauli and/or on stolons. 


REMARKS: Rees (1956b) outlines the taxonomic history of this genus, although 
he erroneously attributed it to a later publication of M. Sars. 

In its prevailing usage (Millard, 1975; Bouillon, 1985), the scope of 
Rhizorhagium is only gradually different from Garveia Wright, 1859 and not well 
separable from the latter. Its scope should therefore be modified, an opinion already put 
forward by Bouillon er al. (2004). The stolonal colony organisation is always a growth 
stage in erect forms and it is likely that it can evolve quite easily from an erect form. 
Stolonal hydroids of the families Bougainvilliidae or Pandeidae that produce a medusa 
stage can also relatively easily evolve into Rhizorhagium through a simple medusa 
reduction. It is therefore very likely that the current concept of Rhizorhagium is poly- 
phyletic. However, due the paucity of morphological characters, only a molecular 
phylogeny can resolve the relationships. As it seems unwise to make not well founded 
provisional changes that might prove obsolete once molecular phylogenies are 
available, and in order to maintain taxonomic stability, the current classification is 
retained here. This compromise was already proposed by Ree (1938: 6), although he 
did not include R. arenosa in this genus. He gave no arguments for this, but likely 
excluded it because it has its gonophores on the hydrocauli and not the stolons. 
Likewise, also Calder (1988) proposed to restrict Rhizorhagium to species with a 
nipple shaped hypostome. 


262 P. SCHUCHERT 


KEY TO THE RHIZORHAGIUM SPECIES TREATED HERE: 


la sonophores. on stolons... u... ns trailer det IAC NE 2 
1b gonophoressonhydrocauli cibo R. arenosum 
2a hypostome nipple-shaped, long hydrocauli .......................... R. roseum 
2b hypostome conical or dome-shaped, hydranths subsessile ................ R. album 
Rhizorhagium roseum M. Sars, 1874 Fig. 26 


? Atractylis coccinea Wright, 1861: 130. 

? Atractylis miniata Wright, 1863: 351.— Rees, 1956a: 341. 

Rhizorhagium roseum M. Sars, 1874 (in G. O. Sars, 1874: 129). — M. Sars, 1877: 28, pl. 2 figs 
37-43. — Jaderholm, 1909: 46, pl. 3 fig. 7. — Broch, 1911: 16, fig. 12. — Broch, 1916: 54. 
fig. R. — Rees, 1938: 8.— Rees, 1956b: 113. — Calder, 1972: 225, pl. 2 fig. 4. — Schuchert, 
2001: 25, fig. 14A-B. 

Perigonimus coccineus. — Hincks, 1871: 77. 

Perigonimus sarsii Bonnevie, 1898b: 6, pl. 1 fig 3.— Rees, 1956b: 113, synonym. 

Garveia groenlandica Levinsen, 1893: 155 .— Kramp, 1911: 263. — Broch, 1916: 55, synonym. 
— Fraser, 1937: 35, fig. 21. 

Wrightia coccinea. — Russell, 1907: 53, fig. 

Perigonimus roseus. — Naumov, 1969: 205, fig. 73. 


MATERIAL EXAMINED: MHNG INVE28734, Greenland, Akuliaruseg, 60.16°N 44.16°W, 
40-60 m, 11 August 1966; male; dense colony on Abietinaria abietina. - MHNG INVE27654, 
Iceland, 66.42°N 23.83°W, 138-152 m, 12 July 1989 (BIOICE station 2530); female colony on 
hydroid. - MHNG INVE33554, Faroes, 62.18°N 04.97°W, 184 m, 07 May 1988 (BIOFAR 
station 165), female colony on hydroid. - BMNH 1967.12.1.3; identified by W. J. Rees as 
Perigonimus coccineus (label); Great Britain, Little Cumbrae, off Lighthouse, 82 m; 8 July 
1966; female. - BMNH 1969.12.2.1966; identified as ? R.roseum by W.J. Rees, is clearly this 
species; on Tubularia; female; origin and date not known. — BMNH 1965.7.21.1; as Perigonimus 
coccineus; Scotland, Bute, Firth of Clyde, Mount Stuart; 20 June 1962; on Tubularia indivisa; 
male. 


DIAGNOSIS: Stolonal hydroid, hydrocauli long and well demarcated from hy- 


dranth, hydranths with pseudohydrotheca, hypostome nipple-shaped, one whorl of ten- 
tacles; sporosacs arising from stolons, on relatively long pedicels. 


DESCRIPTION: Colonies stolonal or stems rarely branched once; stolons 
creeping, ramified to reticulate. Hydrocauli long, enclosed in perisarc that is mostly 
smooth but can have some corrugated regions; perisarc continues distally as thin, 
wrinkled membrane over body of hydranth as a pseudohydrotheca. Hydranth spindle- 
to urn-shaped, diameter about 1/3 of height, appearing relatively broad, 6-12 (normally 
8) filiform tentacles. Hypostome high, thin, characteristically nipple-shaped. 
Gonophores develop from stolons, on pedicels, covered by perisarc, perisarc may be 
loose. Young female sporosacs oblong, later spherical, main body enclosed by 
wrinkled membrane, with a club-shaped spadix on which eggs develop, 2-6 eggs per 
sporosac. The eggs are fertilized in situ and develop within the membrane into planulae 
(larviparous). Male sporosacs oblong, tightly packed with spermatozoids. Colour of 
hydranth: rose to red. Upon disturbance, the hydranth bends back towards the pedicel. 


Dimensions (Rees, 1956b; own observations): Hydrocauli 2-8 mm high 
(according to Broch (1916) up to 12 mm), hydranth 0.4-0.45 mm height, diameter of 
hydranth 0.3-0.35 mm, hydrocaulus diameter 0.12-0.2 mm, length of female sporosac 
0.6-1.2 mm, diameter of gonophores 0.5-0.75 mm, length of sporosac stalk 0.35-0.7 
mm, male sporosacs up to 0.6 mm, diameter 0.45 mm. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 263 


Fic. 26 


Rhizorhagium roseum Sars, 1874, after preserved material, scale bar 0.2 mm. (A) Hydranth with 
pseudohydrotheca and distal part of caulus. (B) Male sporosac. (C) Younger female sporosac 
sporosac. (D) Female sporosac with spadix bearing one egg and two developing embryos. 


DISTRIBUTION: A circumpolar Arctic-boreal species. Its southern limit seems to 
be western Scotland (Firth of Clyde; Russell, 1907; this study). Known from Norway 
(Jäderholm, 1909; Christiansen, 1972), western Sweden (Jäderholm, 1909; Rees & 
Rowe, 1969); the Faroe Islands (this study); western Scotland; Greenland (Schuchert, 
2001); Iceland (Schuchert, 2001); Barents Sea; White Sea (Naumov 1969); north- 
eastern Canada (Calder, 1972); western Canada (Brinckmann-Voss, 1996); Alaska 
(Fraser, 1937). Type locality: Norway, Mangerfjord, Bongnestrgmmen, on Tubularia 
indivsa (Rees, 1956b). 


264 P. SCHUCHERT 


BroLoGY: Found mostly on other hydroids, mainly on Tubularia indivisa and 
Abietinaria abietina at a depth of 15-200 m. Fertile colonies have been observed from 
May to October, but a longer period is possible. 


REMARKS: The synonymy of this species as elaborated by Broch (1916) and 
Rees (1938, 1956b) is adopted here. Arractylis coccinea Wright, 1861, is not 
recognizable, but likely belongs to this species. 

The characteristic hypostome makes this hydroid quite easy to identify, even for 
infertile material. The unique shape of the hypostome was also the reason why Calder 
(1988) excluded from the genus Rhizorhagium all similar species that have a dome- 
shaped hypostome (see also Schuchert, 1996). 

As noted by Rees (1956b), the reaction of the hydranths if disturbed or fixed is 
to bend back towards the stem. This reaction also often seen in the hydroids of the 
genus Amphinema (Schuchert, 1996: 62). 

The size of the sporosacs observed in this study was smaller than those 
observed by Rees (1938, 1956b). 


Rhizorhagium arenosum (Alder, 1862) Fig. 27 


Atractylis arenosa Alder, 1862a: 144. — Alder, 1862b: 313, pl. 13 figs 5-7. — Wright, 1863: 47, 
pl. 4 figs 7-10. — Hincks, 1868: 88, pl. 16 fig 1.— Cornelius & Garfath, 1980: 277. 

Wrightia arenosa. — Allman, 1872: 300. 

Rhizorhagium arenosum. — Bouillon er al., 2004: 49, Fig. 29A. 

? not Aselomaris arenosa. — Pennycuick, 1959: 163, pl. 2 fig. 6. 

Aselomaris arenosa. — Hamond, 1964: 662, fig. 1. 


MATERIAL EXAMINED: MHNG INVE3666, slide; 51.433°N 11.755°E (station: CARA- 
COLE 125-03), Porcupine Seabight, 890 m, collected by submersible robot arm, 3 August 2001, 
male colony, covered by bacteria, epizoic on other hydroid, may belong to this species. 

DIAGNOSIS: Stolonal hydroid, hydrocauli short and not well demarcated from 
hydranth, hydranths with pseudohydrotheca, hypostome dome-shaped to conical, one 
whorl of tentacles, sporosacs on hydrocauli, females with 30 and more eggs, embryos 
develop in acrocyst. 


DESCRIPTION (after literature and own data): Hydroids arising from ramified, 
creeping stolons; stolons covered by perisarc, hydrocauli unbranched, often not 
distinctly delimited from hydranth body which tapers towards base; conical pseudo- 
hydrotheca into which the hydranth can completely withdraw, covered by mud and 
particles. Hydranth spindle-shaped, hypostome conical to dome-shaped, 6-12 tentacles 
in one whorl. Gonophores are sporosacs of the styloid type; one or two sporosacs on 
hydrocaulus, likely also stolons. Female sporosacs oblong (two times as long as wide, 
but ratio may vary), covered by filmy perisarc; mature with many eggs (>30), 
embryonic development takes place in a gelatinous mass (acrocyst) to the planula 
stage, thus larviparous. Male sporosacs similar to female ones. Colours: milky white. 


DIMENSIONS: Hydranth up to 1 mm (estimation), hydrocaulus and pseudo- 
hydrotheca up to 1.4 mm; female sporosac up to 0.5 mm (measured in Hamond, 1964), 
male sporosacs 0.3 mm long, 0.15 mm wide. 


OTHER DATA: Wright (1863) described the maturation of the sporosac 
(Fig. 27E-G). When mature, the thin periderm envelope ruptures and the sporosac 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 265 


Fic. 27 


Rhizorhagium arenosum (Alder, 1862). (A) Expanded hydranth. (B) Contracted hydranth. (C) 
Hydranth with two sporosacs. (D) Hydrocaulus with two young female sporosacs, scale bar 0.5 
mm. (E) Mature female sporosac before extrusion of eggs. (F) Female sporosac in process of 
extruding its eggs. (G) Same as F, eggs are in gelatinous acrocyst. A-B, from Alder (1862b); 
C, from Hincks (1868); D, modified after Hamond (1964), E-G, from Wright (1863). 


266 P. SCHUCHERT 


emerges in a long cylindrical form, it contracts like a medusa bell. Concomitantly, the 
apical part of the sporosac secretes a gelatinous substance and further contractions let 
the sporosac rupture, so that the eggs are pushed into the gelatinous acrocyst where 
they develop into planulae. 


BıoLoGy: Predominantly an intertidal species occurring on algae, stones and 
other substrates. 


DISTRIBUTION: North Sea coast of Britain (Allman, 1872), perhaps also SW 
Ireland (Duerden, 1893; this study). Type locality: Tynemouth and Cullercoats, 
Northumberland, intertidal, on stones and Laminaria holdfasts. 


REMARKS: This is a rather poorly documented species, which may be more 
frequent than reflected by its few records. As remarked by Hamond (1964), infertile 
colonies of this species are indistinguishable from immature Bougainvillia muscus. 

The examined material from deep waters SW of Ireland differs from the 
existing descriptions in that the sporosacs arise from the stolons. In combination with 
its occurrence in deep waters, this identification is therefore somewhat doubtful. 

The Mediterranean record of this species (Lo Bianco, 1909) must be a misiden- 
tification of Pachycordyle pusilla (Motz-Kossowska, 1905) (see Schuchert, 2004). Lo 
Bianco (1909) states: «occurs on Posidonia in shallow waters». 


Rhizorhagium album (Rees, 1938) Fig. 28 


? Eudendrium pudicum Van Beneden, 1867: 116, pl. 8 figs 1-2. 
Rhizorhagium album Rees, 1938: 9, fig. 2a-c. 

DIAGNOSIS: Stolonal hydroid, hydrocauli very short or absent, hydranths with 
pseudohydrotheca, hypostome conical, one whorl of tentacles, styloid sporosacs on 
stolons, ovoid shape. 


DESCRIPTION (after Rees, 1938): Colonies small, stolons creeping, covered by 
thin, irregularly wrinkled perisarc. Hydranths with very short hydrocauli, sometimes 
hydranths subsessile, pedicels covered by wrinkled perisarc continued as thin, filmy 
pseudohydrotheca over hydranth body; hydranths tubular, hypostome conical, 6-10 
filiform tentacles in a single whorl. Colour of polyps white. Gonophores ovoid, 
developing from stolons on relatively long stalks, sporosacs of the styloid type, spadix 
present, only males known. 


DIMENSIONS (after Rees, 1938): Stolon diameter 0.1 mm, hydrocauli maximally 
0.3 mm high, hydranths 0.35-0.55 mm high, diameter 0.1-0.14 mm. Pedicels of 
sporosacs 0.3-1.0 mm, sporosac diameter 0.4 mm. 


BroLocy: The original colony was found on filamentous algae in a rock pool. 
DISTRIBUTION: Known from the type locality only, Newquay (Cornwall, 
England). 


REMARKS: This species is only known from its original description based on 
cultivated material. The principal difference to R. arenosum is the location of the 
sporosacs on the stolons. This could be, however, a cultivation artefact as cultivated 
hydroids can differ from those grown under natural conditions. I strongly suspect that 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 267 


Fic. 28 


Rhizorhagium album (Rees, 1938), modified after Rees (1938), scale bar 0.3 mm. (A) Part of 
cultivated colony. (B) Single hydranth. (C) Sporosac. 


R. album is only a form of R. arenosum. Also the similar Rhizorhagium antarcticum 
(Hickson & Gravely, 1907) can have its sporosacs either on the hydrocauli or the 
stolons (Schuchert, 1996). 


Genus Pachycordyle Weismann, 1883 

TYPE SPECIES: Pachycordyle napolitana Weismann, 1883 by monotypy. 

SYNONYMS: Clavopsella Stechow, 1919; Thieliana Stepanjants et al., 2000 [type species 
Clavopsella quadranularia Thiel, 1962]. 

DragNosIs: Stolonal or branching hydroid colonies, hydranths pedicellate, 
perisarc terminating at base of hydranth. Hydranths club-shaped, hypostome dome- to 
nipple-shaped, tentacles in two or more whorls. Gonophores borne on stems or 
pedicels. Gonophores fixed sporosacs or degenerated medusae lacking mouth, mar- 
ginal tentacles, oral tentacles, radial canals, and ocelli. 


REMARKS: See Schuchert (2004) for a more information on the species 
Pachycordyle napolitana Weismann, 1883, Pachycordyle navis (Millard, 1959), and 
Pachycordyle pusilla (Motz-Kossowska, 1905). 


Pachycordyle michaeli (Berrill, 1948) n. comb. Fig. 30 


Aselomaris michaeli Berrill, 1948: 289, figs 1-4. 

not Clavopsella michaeli. — Piraino, 1992: 241, figs 1-13, [= Pachycordyle pusilla (Motz- 
Kossowska, 1905)]. 

not Rhizorhagium michaeli. — Bouillon et al., 2004: 49, fig. 29B-C. 


MATERIAL EXAMINED: Paratypes BMNH 1985.9.13.9, as Aselomaris michaeli Berrill, 
1948, USA, Boothbay Harbour, Maine, leg. N. Berrill, large colonies, male and females present. 


268 P. SCHUCHERT 


Fic. 29 


Pachycordyle navis (Millard, 1959). (A) Colony silhouettes, scale bar 0.4 mm. (B) Part of branch 
with two male sporosacs, scale bar 0.2 mm. (C) Young female sporosacs, right one as optical 
section, same scale as B. (D) Perisarc of branch, outside is left, note transition from single- to 
double layered perisarc, scale bar 50 um. After preserved material from Widewater, Lancing, 
Sussex, UK, BMNH 1959.10.4.2. 


DESCRIPTION: Colony stolonal or occasionally sparingly branched. Hydrocauli 
and stems covered by perisarc, terminating below hydranth, thus without pseudo- 
hydrotheca. Perisarc mostly corrugated, sometimes smooth. Hydranths club-shaped, 
hypostome high, conical to dome-shaped; tentacles 12-20, in 2-3 closely approxi- 
mated, alternate whorls. Gonophores fixed sporosacs (heteromedusoid type), numerous 
on hydranth pedicels, spherical to ovoid, with conspicuous spadix, stalks distinct and 
usually relatively long, but length variable. Colonies dioecious. Female sporosacs with 
about 6-10 eggs, when mature with nematocysts on surface. Eggs develop into 
planulae in situ. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 269 


Fic. 30 


Pachycordyle michaeli (Berrill, 1948), after preserved paratype material. (A) Colony silhouette, 
height approx. 3 mm. (B) Hydranth and several male sporosacs, scale bar 0.2 mm. (C) Spent 
male sporosac in more proximal region of shoot shown in B, note the very long stalk, same scale 
as B. 


DIMENSIONS: Stems up to 4 mm, hydranths 0.4 mm high. Sporosacs up to 
0.5 mm long and 0.35 mm wide, pedicels sporosacs length very variable 0.07-1.2 mm. 


BIOLOGY: Shallow water species, growing on wooden floats and Fucus. Light 
induces spawning (Berrill, 1948). 


DISTRIBUTION: Presumably only known from region of type locality, Boothbay 
Harbour (Maine, USA). The Mediterranean record likely refers to P. pusilla. 


REMARKS: Piraino (1992) described the nematocyst-covered eggs of a 
Mediterranean bougainvilliid species which he named Clavopsella michaeli (Berrill, 
1948). The hydroid was cultivated in the laboratory and Dr Piraino kindly furnished me 
with some preserved material of this colony, part of which was deposited in the 
MHNG (as INVE38150). This material was compared to the type material of 
Aselomaris michaeli Berrill, 1948 (see above), material of Pachycordyle pusilla (Motz- 
Kossowska, 1905) (MHNG INVE36202; INVE32953, both Mediterranean) and P. 
navis (Millard, 1959) (MHNG INVE31852 and INVE34226, both from Denmark; 


270 P. SCHUCHERT 


BMNH 1959.10.4.2, UK material, Fig. 29). The material of Piraino appears indistin- 
guishable from Pachycordyle pusilla (Motz-Kossowska, 1905) (see review in 
Schuchert, 2004), the only unusual trait being its original substrate. Piraino (1992) 
found his hydroid on the tubes of the polychaete Sabella spallanzani (Gmelin, 1791), 
while Pachycordyle pusilla was so far only Known to occur on seagrasses (Posidonia 
and Cymodocea). It is nevertheless possible that P. pusilla can also grow on other sub- 
strates. Müller (1913, as Pachycordyle fusca) cultivated P. pusilla over long periods 
and he observed that the colonies also spread onto lead blocks. I am therefore con- 
vinced that the hydroid described in Piraino (1992) should be assigned to P. pusilla. 
The higher number of sporosacs and the nematocysts on the eggs cannot be used to 
keep it separate. In the original description of P. pusilla, Motz-Kossowska (1905) men- 
tions that there can be 2-5 sporosacs per hydrocaulus and there are nematocysts on the 
egg surface. 

The re-examination of the type material of Aselomaris michaeli showed that the 
description of Berrill (1948) must be modified somewhat. The hydrocauli bear mostly 
only one hydranth, but rarely some branching occurs. More important is that the tenta- 
cles are not in a single whorl, but in two close-set whorls, sometimes even in three well 
separated whorls. The hydroid resembles very closely Pachycordyle navis (Millard, 
1959) (see Fig. 29) and I suspect that they are conspecific. The very long pedicels of 
the sporosacs are not unique to R. michaeli, similarly long pedicels of the sporosacs 
have also been depicted by Thiel (1962) for Clavopsella quadanularia, a synonym of 
P. navis (Millard, 1975). 

As already mentioned in Schuchert (2004), P. navis resembles also P. pusilla, 
the differences being mainly the longer hydrocauli that can also be branched, the long 
gonophore pedicels, and its preference for reduced salinities. There remains, however, 
the possibility that even P. pusilla is only a form of P. navis growing in fully marine 
environments. Detailed investigation using molecular methods are needed to re- 
evaluate the status of P. pusilla, P. navis, and R. michaeli. Some preliminary results 
(Fig. 4) show that P. pusilla is in fact closely related to some Bougainvillia species, 
indicating that even the inclusion in the genus Pachycordyle is incorrect. 


Genus Velkovrhia MatjaSiC & Sket, 1971 
TYPE SPECIES: Velkovrhia enigmatica MatjaSıC & Sket, 1971. 


DIAGNOsIs: Colonial, stolonal freshwater hydroids occurring in caves. Hydranth 
pedicels covered by irregularly corrugated, multi-layered perisarc, hydranth body urn- 
shaped, without pseudohydrotheca, filiform tentacles in two whorls, hypostome nipple- 
shaped. Gonophores arise on hydranth pedicels, fixed sporosacs, females with 
branched gastrodermal system, larviparous. 


REMARKS: The genera Parawrightia Warren, 1907 and Velkrovhria appear very 
similar. There are some small differences in their sporosacs (Parawrightia has 
sporosacs with radial canals (comp. Warren, 1907; Wedler & Larson, 1986; Velikonja, 
1994). Additionally, Parawrightia has a pseudohydrotheca and is a marine species. 
Although these traits are normally not considered to be important enough to separate 
genera, the genus name Velkrovhria should be retained until it is clear whether its type 
species might not belong to Cordylophora (see below). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 271 


Velkovrhia enigmatica MatjaSi¢ & Sket, 1971 Fig.31 

Velkovrhia enigmatica MatjaSiöE & Sket, 1971: 139, figs 1-4. — KuStor, 1977: 51. — Sket & 
Matjasié, 1977: 263-265, figs 1-2. — Velikonja, 1986: 123, fig. 2. — Velikonja, 1994: 39, 
fig 1. — Holstein, 1995: 99, fig. 52. — Bouillon et al., 2004: 49, fig. 29E. 


MATERIAL EXAMINED: No material was available, but Dr B. Sket kindly provided some 
good colour photographs of living animals. 


DIAGNOSIS: As for genus. 


DESCRIPTION (after literature data): Small stolonal colonies, stolons creeping on 
substrate, perisarc covered. Hydranths on perisarc covered pedicels, perisarc multi- 
layered, irregularly corrugated, terminating below hydranth. Hydranth ufn-shaped, 
with high, nipple-shaped hypostome, up to 10 tentacles in two close set whorls, 
alternating. Gonophores are sessile sporosacs, developing on hydrocauli, spherical to 
ellipsoid, covered by filmy periderm, colonies dioecious; female sporosacs with about 
5 eggs, planulae develop in situ, gastrodermal system (spadix?) branched, adnate to 
wall. Colour milky white, periderm yellowish-brown. Nematocysts: desmonemes and 
microbasic euryteles (KuStor, 1977). 


DIMENSIONS: Hydrocauli 1.3-1.7 mm, hydranths 0.4 mm, hydrocaulus diameter 
0.25 mm. Sporosacs about 0.4 mm, planula size 0.2 mm. 


BIOLOGY: A freshwater species living in carstic caves. The hydranth density is 
dependant on the season, with a summer minimum and a winter maximum (Velikonja, 
1986). 


DISTRIBUTION: Slovenia, Croatia, and Hercegovina. Type locality: Rake 
Passage, Cave of Planina, Slovenia. 


REMARKS: Although Matjaÿié & Sket (1971, in English summary) describe this 
species as having one whorl of tentacles only, their figures show two whorls. Sket & 
Matjasi¢ (1977) describe the hydranth as having two tentacle whorls. The spadix of the 
female sporosac is branched as suggested by Figure 31B and also colour photographs 
of living animals (given as black and white photographs in Sket & Matjaëië, 1977). 

Velkovrhia enigmatica shares many traits with Cordylophora caspia (Pallas, 
1771) (for a review see Schuchert, 2004): the hydranth shape, the female sporosacs 
with the branched spadix, and its occurrence in fresh-water. Kinne (1956; 1957) has 
shown in extensive experiments that Cordylophora caspia changes drastically its mor- 
phology in varying salinities. In freshwater, the colonies remain stolonal, the hydranths 
get small and broad, and the tentacle number is reduced, approaching thus the mor- 
phology of V. enigmatica. But even in freshwater the colonies could form gonophores. 
It could thus be that V. enigmatica is nothing but a subdued form of Cordylophora 
caspia. However, at this stage this stage this is only an assertion that has to be proven 
by either by cultivating V. enigmatica in water with varying salinities, or by 
comparisons of 16S sequence data. 


INDETERMINATE BOUGAINVILLIDAE OR SPECIES REFERABLE TO OTHER FAMILIES 


Atractylis bitentaculata Wright, 1867 
Atractylis bitentaculata Wright, 1867: 45, pl. 1 fig. 5. 
Perigonimus bitentaculatus.— Hincks, 1968: 98, fig. 9. — Rees, 1956a: 338. 


TYPE LOCALITY: Firth of Forth, near Inch Keith, on a Pecten shell. 


272 P. SCHUCHERT 


Fic. 31 


Velkovrhia enigmatica MatjaSiC & Sket, 1971. (A) Two hydranths issuing from a common 
stolon, rear polyp with sporosac, after photograph in Velikonja (1994), some tentacles imaginary 
(not clearly visible in photo). (B-C) Histological sections of female and male sporosacs; em 
developing planula, gd gastrodermis, oc oocyte, sp spermatids; from Velikonja (1986). 


REMARKS: Rees (1956a) thought that this species, the gonophores of which are 
unknown, might belong to the genus Proboscidactyla. Because the members of this 
genus occur exclusively around the openings of the tubes sabellid polychaetes and 
Wright’s species grew on a Pecten shell, this appears less likely. Moreover, the figures 
in Hincks (1868, reproduction of Wright, 1867?) shows polyps with distinct pedicels 
and a conical hypostome. Allman (1872: 329) regarded it as an immature form of an 
indeterminate hydroid. 


Atractylis quadritentaculatus Wright, 1867 
Atractylis quadritentaculatus Wright, 1867: 45, pl. 1 fig. 6. 


REMARKS: An indeterminate species. 
Perigonimus georginae Hadzi, 1913 
Perigonimus georginae Hadzi, 1913: 98, figs 7, 10-11. — Rees, 1956a: 340. 


REMARKS: An indeterminate species, could be Amphinema rugosum or A. 
dinema (Rees, 1956a). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 273 


Perigonimus inflatus Duerden, 1895 
Perigonimus (?) inflatus Duerden, 1895: 329, pl. 14 fig. 4. — Rees, 1956a: 340. 

REMARKS: An indeterminate species, could be a Rhizorhagium species (Rees, 
19562). 


Perigonimus nutans Hincks, 1877 
Perigonimus ? nutans Hincks, 1877: 149, pl. 12 fig. 1. — Rees, 1956a: 342. 

REMARKS: An indeterminate species, could be Amphinema rugosum or A. dine- 
ma (Rees, 1956a). 


Perigonimus decorans Schneider, 1898 
Perigonimus decorans Schneider, 1898: 479. — Rees, 1956a: 340. 

REMARKS: An indeterminate species, it is unclear to which family it belongs 
(Rees, 19562). 


Perigonimus steinachi Jickly, 1883 
Perigonimus steinachi Jickly, 1883: 617, pl. 27 figs 1-9. — Rees, 1956a: 344. 

REMARKS: An indeterminate species, it is unclear to which family it belongs 
(Rees, 1956a). Perhaps referable to Perarella schneideri. 


Eirene viridula (Péron & Lesueur, 1810) 
Oceania viridula Péron & Lesueur, 1810: 346. 
Perigonimus nudus Stechow, 1919: 16, fig. D. new synonym 
Eirene viridula. — Cornelius, 1995: 223, fig. 51. 

REMARKS: The type material of Perigonimus nudus Stechow, 1919 could be ex- 
amined (ZSM, 20000646 and 20000647, 2 slides, loc. Trieste, leg. Maas 1908). The 
material is obviously Eirene viridula (for description see Cornelius, 1995). 


Lizzella hyalina (Van Beneden, 1867) 
Circe hyalina Van Beneden, 1867: 95, pl. 3 fig 14. 
? Lizzella hyalina. — Haeckel, 1880: 634. 
Lizzella hyalina. — Hartlaub, 1911: 151, fig. 137. — Kramp, 1959a: 86, doubtful species. 
REMARKS: Van Beneden (1867) described a medusa from the Belgian coast that 
resembled closely R. octopunctata, but it had eight radial canals and lacked medusa 
buds. He named it Circe hyalina. Haeckel (1880) thought that he had found the same 
species at the coast of Normandy. Because Circe is a preoccupied name for a mollusc 
genus, Haeckel referred it to the genus Lizzella, a genus he had proposed earlier for 
Lizzella octella Haeckel, 1879. However, Haeckel’s (1880) and Van Beneden’s (1867) 
descriptions differ in some important aspects (colour, number of radial canals) that it 
appears improbable that both had observed the same species. Van Beneden (1867) 
described and depicts eight radial canals for his Circe hyalina. This was contested by 
Haeckel (1880) who dismissed the radial canals of Van Beneden as longitudinal 
muscles. This, however, appears unlikely for a bougainvilliid medusa. A quite similar 
medusa but without oral tentacles has been described from Japan, namely Octorathkea 
onoi Uchida, 1927. 
If the description of Van Beneden was accurate, then his species should be 
recognizable. Because the species has never been found again, despite of it originating 


274 P. SCHUCHERT 


in one of the best explored waters the world, it is more likely that Van Beneden had an 
abnormal Rathkea octopunctata, a view already expressed by Russell (1953). 


FAMILY CYTAEIDIDAE L. AGASSIZ, 1862 
TYPE GENUS: Cytaeis Eschscholtz, 1829. 


DIAGNOSIS: Hydroids forming mostly non-polymorphic colonies, rarely poly- 
morphic, arising from reticulate stolons covered by perisarc. Hydranths without 
distinct caulus (sessile), with one whorl of filiform tentacles below conical hypostome. 
Base of hydranths often with a perisarc collar. Gonophores develop from stolons and 
are liberated as medusa, medusoids, or remain fixed as sporosacs. Medusa with simple, 
circular mouth, with four or more non-branching oral tentacles near mouth. Gonads 
develop in interradial position on manubrium or encircle manubrium. With four, rarely 
eight, solid marginal tentacles without lumen. Ocelli lacking. 


REMARKS: Although the original name was Cytaeidae, the correct name 
formation following the provisions of the ICZN is Cytaeididae (Kramp, 1961). The 
latter spelling must be used as it is now the prevalent one. 

Rees (1962) provides a taxonomic review of the hydroids of this family. Also 
Calder (1988) gives important taxonomic details. However, the scope of the family 
remains problematic. See also remarks under the genus Perarella. 


KEY TO THE GENERA OF THE CYTAEIDIDAE: 


la gonophores medusae having 4 or 8 marginal tentacles ........................ 2 
lb gonophores medusoids Of SPOrOSACS ENT LR Ce CCC ee eee Perarella 
2a medusa with erehtmaremal tentacles’ 557) -). hi ijt ee eee ee Paracytaeis 
2b medusaswithfourmarsinalitentacles |. Ei. Wa Re Cytaeis 


Genus Cytaeis Eschscholtz, 1829 

TYPE SPECIES: Cytaeis tetrastyla Eschscholtz, 1829 by monotypy. 

SYNONYMS (after Calder, 1988): Cyractis de Blainville, 1834: 284 [incorrect spelling]. — 
Cyteis Van Beneden, 1867: 18 [incorrect spelling]. — Cytheis Van Beneden, 1867: 18 [incorrect 
spelling]. — Nigritina Haeckel, 1879: 73. — Cytaeidium Haeckel, 1879: 75 [type species Cytaeis 
pusilla Gegenbaur, 1856]. — Cytaesis Bouillon, 1978a: 129 [incorrect spelling]. 

DIAGNOSIS: Hydroids monomorphic, arising from reticulate stolons covered by 
perisarc. Hydranths sessile, with one whorl of filiform tentacles below conical hypos- 
tome; base of hydranths often with a perisarc collar. Gonophores develop from stolons, 
liberated as medusa with four tentacles. 

Medusa with four or more capitate oral tentacles arising well above mouth mar- 
gin, medusa budding from manubrium. With four marginal tentacles. 


REMARKS: Calder (1988) gives the taxonomic history of this genus. There are 
several nominal Cytaeis species, but their status remains unclear (Kramp, 1961; Rees, 
1962; Calder, 1988; Pages et al., 1992; Schuchert, 1996; Bouillon ef al., 2004). 
Bouillon er al. (2004) included in this genus also forms with sporosacs or medusoids. 
As it is not clear that these forms are really members of the Cytaeididae, I did not fol- 
low this. The arguments are discussed under the genus Perarella. 

Currently there is only one accepted species known in the region of inves- 
tigation. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 275 


Cytaeis tetrastyla Eschscholtz, 1829 Fig. 32 


Nigritina atlantica Steenstrup. — Kramp, 1961: 64, name not available. 

Cytaeis tetrastyla Eschscholtz, 1829: 104, pl. 8 fig. 2. — Will, 1844: 67. — Haeckel, 1879: 73. — 
Mayer, 1910: 133. — Hartlaub, 1911: 139, figs 127-129. — Vanhôffen, 1911: 204, fig. 6. — 
Bigelow, 1918: 367. — Kramp, 1953: 263. — Kramp, 1959a: 99, fig. 62, pl. 1 figs 3-6. — 
Kramp, 1965: 9. — Kramp, 1961: 63. — Rees, 1962: 382. — Kramp, 1968: 26, fig. 64. — 
Brinckmann-Voss, 1970: pl. 7 fig. 4. — Goy, 1973: 976. — Bouillon, 1980: 318. — Bleeker 
& van der Spoel, 1988: 230, fig. 5.— van der Spoel & Bleeker, 1988: 168, fig. 4— Goy er 
al., 1991: 106, fig. 13. 

? Bougainvillia mediterranea Busch, 1851: 21, pl. 2 figs, 10-11. — Haeckel, 1879: 73, synonym. 

Cytaeis pusilla Gegenbaur, 1856: 228, pl. 8 fig. 8. — Keferstein & Ehlers, 1861: 84, pl. 13 figs 
8-9. — Haeckel, 1879: 75. — Kramp, 1959a: 99, doubtful synonym. — Kramp 1961: 63. 

Cytaeis macrogaster Haeckel, 1879. pl. 6 fig. 1.- Kramp, 1955a: 153, synonym. 

Cytaeis nigritina Haeckel, 1879: 74, pl. 6 figs 2-5. — Maas, 1904: 8, pl. 1 figs 3-4. — Kramp, 
1955a: 154, synonym. 

? not Cytaeis vulgaris Agassiz & Mayer, 1899: 161, pl. 2 figs 3-5. — Mayer, 1910: 134. — Kramp, 
1968: 27, perhaps valid. 

Cytaeis herdmani Browne, 1905a: 135, pl. 1 fig. 1, pl. 4 fig. 12. — Mayer, 1910: 135. — Browne, 
1916: 177, synonym. 

Cytaeis vulgaris. — Bigelow, 1909: 190, pl. 6 fig. 3, pl. 40 figs 2, 5, pl. 43 fig. 4. 

Cytaeis atlantica. — Mayer, 1910: 133, figs 71-72 [invalid combination]. 

? not Cytaeis pusilla. — Neppi & Stiasny, 1913: 23. 

Cytaeis japonica Uchida, 1927: 215, fig. 39. - Kramp, 1959a: 99, synonym. 

[not Cytaeis japonica. — Komai, 1931, = Cytaeis uchidae Rees, 1962] 

? Cytaeis vulgaris. — Kramp, 1928: 44, fig. 19. 

Cytaeis. spp. — Pages et al., 1992: 5, fig. 4. — Bouillon et al., 2004: 55, fig 33A. — Pages et al., 
2006: Fig. 7G (colour photograph). 

Cytaeis cf. tetrastyla— Schuchert, 1996: 61, fig. 35a-b. 


MATERIAL EXAMINED: BMNH 1970.10.8.55; Brit. Terra Nova Expedition 1910-1913, 
station H28(59); 0° N 25.25°W, equatorial Atlantic; 16 May 1913; 7 medusae. - MNHN1625; 
France, Villefranche-sur-Mer; 03 January 1967; material of Goy (1973); 2 medusae, with and 
without gastric peduncle, immature, 1.4 mm high. 


DIAGNOSIS: See genus diagnosis. 


DESCRIPTION (after own data and various sources): Fully grown medusa with 
bell-shaped umbrella, widest near base, jelly not thick, apical jelly sometimes slightly 
thicker than lateral walls, with or without short peduncle. With four broad radial canals 
and thin circular canal. With four tentacles of about bell length when contracted, thick, 
tapering. Epidermis of tentacle bulbs triangular, flat, attached to exumbrella around 
tentacles (not in juveniles). Manubrium very voluminous, pear-shaped, length up to 4/5 
of bell cavity; manubrium with 8-32 (max. 50) simple, capitate tentacles scattered in a 
band slightly above mouth rim, adnate at origin. Stomach with medusa buds in animals 
without gonads, irregularly scattered, usually more towards top, some animals with 
large number of buds spreading over almost entire manubrium. Gonads interradial 
pads. Colour (Kramp, 1955b, eastern Atlantic): stomach light brown with darker 
coloured perradial edges, central part of tentacle bulb black, epidermis brown or bright 
reddish brown. Younger stages likely not coloured. 


DIMENSIONS: Medusa usually not larger than 2-4 mm (rarely up to 6 mm high 
after Kramp, 1961), 3 mm wide. Nematocyst dimensions are given in Bouillon (1980) 
and Schuchert (1996). 


276 P. SCHUCHERT 


waded 


Cytaeis tetrastyla Eschscholtz, 1829; after preserved material. (A) Fully grown medusa with 
medusa buds from near the Fiji Islands, modified after Schuchert (1996), scale bar 1 mm. Note 
that future investigations might show that this Pacific medusa belong to a separate species (B) 
Cnidome of medusa shown in A, microbasic euryteles, desmonemes, microbasic mastigophores, 
scale bar 10 um. (C) Juvenile medusae from the Ligurian Sea (Mediterranean), note the presence 
of a gastric peduncle, the exumbrella has weak longitudinal furrows, scale bar 0.4 mm. 


Fic. 32 


OTHER DATA: Polyp buds on the manubrium were described by Kramp (1959a). 


BIoLOGY: Prefers depths of 0-100 metres depth (Bigelow, 1909). Bleeker & van 
der Spoel (1988) give a range 0-520 m. 


DISTRIBUTION: Occurs circumglobally in tropical to subtropical seas, partly 
oceanic (Kramp, 1959a; 1965). In the eastern Atlantic it occurs from the Strait of 
Gibraltar to South Africa (Maas, 1904; Vanhöffen, 1911; Kramp, 1955b; Bleeker & van 
der Spoel, 1988; Pages er al., 1992). It is also present in the Mediterranean (Will, 1844; 
Goy, 1973; Brinckmann-Voss, 1970 & 1987; Dallot et al., 1988; Goy et al., 1991; 
Benovic & Lucic, 1996). Type locality: Atlantic Ocean, south of Equator. 


REMARKS: Eschscholtz’s (1829) description and figure of Cytaeis tetrastyla 
clearly permit to recognize the medusa as a Cytaeis species. His medusa was certainly 
based on a juvenile medusa of about 1 mm size and with only eight oral tentacles. This 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 274 


caused some problems when later adult forms were described as distinct species. 
Kramp (1953) gives a good overview on the taxonomic history and situation of this 
species. He thinks that there is only one valid species, but later (Kramp, 1968) he 
apparently changed his mind and kept Cyfaeis vulgaris sensu Agassiz & Mayer, 1899 
distinct (not other records assigned to this species). The taxonomy of Cytaeis tetrastyla 
from the Indian- and Pacific Ocean remains unsatisfactory. Experienced students like 
Vanhöffen, Bigelow, and Kramp, who have seen numerous specimens and from differ- 
ent oceans, tended lump all nominal Cytaeis medusa into C. tetrastyla. Rees (1962) 
working with the hydroid phase found several distinct species and suspected that 
Cytaeis tetrastyla sensu Kramp (1961) represents an aggregate of species. This view 
was also adopted by e. g. Bouillon (1980) and Pages er al. (1992). The latter authors 
and also Bouillon er al. (2004) thought that the Cyfaeis tetrastyla medusae sensu 
Kramp (1961) should not be given a specific name until the taxonomy has been sorted 
out properly. We could indeed be faced with a situation as in many Clytia and Obelia, 
where the species can only be distinguished in the hydroid phase. However, at least for 
the eastern Atlantic, no distinct Cytaeis polyps are known and there is so far no indi- 
cation that there are distinct morphotypes of the medusa phase. The mature form as 
described here has been regularly found in the same region (see under Distribution). 
Cytaeis tetrastyla is partly an oceanic form and it can be expected to have a widespread 
distribution. We can thus be reasonably sure that for the eastern Atlantic, Cytaeis 
tetrastyla is the correct name. 

The situation of the Mediterranean is more difficult to evaluate because only 
few authors documented the fully grown form as described above (e. g. Goy et al., 
1991). The uncertainty mainly relates to the status of Cytaeis pusilla. Gegenbaur 
(1856) described this species based on a juvenile Cytaeis he found near Messina 
(Sicily). It was about 2 mm high and had a gastric peduncle. In his figure he depicted 
the medusa with a peduncle of about the same length as the manubrium itself, but in 
the text he qualifies it as short. Haeckel (1879) used the presence of a peduncle to sep- 
arate this species from all other Cytaeis, placing it even in a separate subgenus 
Cytaeidium. Gegenbaur did not discuss why he regarded C. pusilla as distinct from 
C. tetrastyla, but from his discussion on the genus one can deduce that he erroneously 
assumed that C. tetrastyla has ocelli, while there are none in his C. pusilla. Keferstein 
& Ehlers (1861) found C. pusilla again and described the peduncle as short. Haeckel 
(1879) had again material from Messina with a long peduncle (umbrella 2-3 mm). Goy 
(1973, Ligurian Sea, see Fig. 32C) found both forms, with and without a peduncle, in 
the same catch. I think that not too much weight should be placed on this character as 
it is likely variable and in later growth the peduncle might be obliterated by the mas- 
sive growth of the manubrium. I therefore regard C. pusilla as a questionable synonym 
of C. tetrastyla. The medusa identified by Neppi & Stiasny (1913) as C. pusilla, how- 
ever, could be a different species. Their single medusa from the Adriatic Sea was very 
small, measuring only 0.62 mm in height, but nevertheless it had mature gonads. 

The polyp stage of C. tetrastyla remains unknown. The polyp described by 
Calder (1988) as Cytaeis spec. could belong to C. tetrastyla. 

More developmental and life-cycle studies of various Cytaeis medusae and 
polyps are evidently needed to clarify the situation. Above all, a neotype of Cytaeis 


278 P. SCHUCHERT 


tetrastyla should be designated, this based on a medusa from the tropical Atlantic and 
whose 16S and COI gene sequences are known. 


Genus Perarella Stechow, 1922 
TYPE SPECIES: Perigonimus schneideri Motz-Kossowska, 1905 by monotypy. 


DrAGNoOsis: Hydroids with perisarc-covered, reticulate stolons. Hydranths 
naked, sessile or nearly so, sometimes with a basal collar of perisarc, one whorl of 
filiform tentacles. Gonophores arise from stolons, developing either into medusoids or 
fixed sporosacs. 


REMARKS: Following the idea of Petersen (1990) that genera should not be 
delimited based on the degree of gonophore reduction, Bouillon er al. (2004) 
synonymized Perarella with Cytaeis. This approach is certainly correct in cases where 
ahydroid can be attributed unambiguously to a certain family, as e. g. in the Corynidae 
or Tubulariidae. But in the case of the genus Perarella this is not so. Perarella schnei- 
deri, the type species of Perarella, could as well belong to the Bougainvilliidae or 
Pandeidae. Comparing it with some Rhizorhagium species (see above), one is even 
tempted to synonymize Perarella with this genus. It is therefore advisable to retain the 
genus Perarella until a reliable molecular phylogenetic analysis has revealed its true 
affinities. 


Perarella schneideri (Motz-Kossowska, 1905) Fig. 33 


Perigonimus schneideri Motz-Kossowska, 1905: 72, fig. 61. — Rees, 1956a: 344. 

? Perigonimus steinachi Jickeli, 1883: 617, pl. 27 figs 1-9. — Rees, 1962: 393. 

Stylactella (Hydractinia) else-oswaldae Stechow, 1921: 251. 

Clavopsis schneideri. — Stechow, 1921: 252. 

Stylactella elsae-oswaldae.— Stechow, 1923a: 64, fig. E. — Picard, 1958: 190, synonym. 

Perarella schneideri Stechow, 1922: 145. — Rees, 1962: 393. — Bavestrello et al., 2000: 141, 
figs 1,4-5. 

Cytaeis schneideri. — Bouillon et al., 2004: 55, fig. 321. 


MATERIAL EXAMINED: MHNG INVE49086, Italy, Portofino, coll. June 2006, infertile, on 
Schizoporella longirostris (Hincks) growing on glass bottles, leg. & det. S. Puce, 16S sequence 
AM411414. 

DIAGNOSIS: Hydroid growing on bryozoan, two types of hydranths, larger ones 
with many tentacles and smaller hydranths with few short tentacles; gonophores free 
medusoids released with mature gonads. 


DESCRIPTION (Bavestrello er al., 2000; own data): Hydroid colonies stolonal, 
growing on the cheilostomate bryozoan Schizoporella longirostris (Hincks), stolons 
reticulate, running along the zooid margins of the bryozoan, often embedded in the 
skeleton of the host. Hydranths dimorphic with a larger form with long tentacles and a 
slender form with only 4-5 stubby tentacles; both forms of hydranths without distinctly 
demarcated pedicel, base in a short perisarc tube that usually widens somewhat 
towards distal, perisarc thin, not annulated. Larger hydranths club-shaped, high conical 
hypostome, one whorl of 8-14 long, filiform tentacles. Smaller hydranths thinner, very 
contractile, hypostome short, one whorl of 4-5 short, stubby tentacles. Gonophores 
develop on stolons, with distinct pedicel enclosed in perisarc that widens like a funnel, 
gonophores released as medusoids with fully developed gonads. Free medusoid with 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 279 


nu SS 


ae ER 
SS teen 


\ 


Vi 
ae 


Fic. 33 


Perarella schneideri (Motz-Kossowska, 1905). (A) Hydranth form with long tentacles, scale bar 
0.2 mm. (B) Hydranth type with few short tentacles, same scale as A. (C) Free female medusoid, 
scale bar 0.5 mm. (D) Scanning electron microscopic photo of the skeleton of Schizoporella 


longirostris, scale bar 0.3 mm. A-B, after preserved material; C, modified after Bavestrello er al. 
(2000). 


280 P. SCHUCHERT 


bell-shaped umbrella, slightly higher than wide, jelly thin, four radial canals ending in 
marginal bulbs, ring canal, no tentacles, velum present; manubrium long, non-func- 
tional, reaching almost to velum level, encircled by voluminous gonad filling subum- 
brella cavity, females with 80 or more eggs; only female medusoids swim, males inac- 
tive. Nematocysts: microbasic euryteles and desmonemes. Colours (Motz-Kossowska, 
1905): hydranths red, male gonophores whitish with red gastrovascular canals. 


DIMENSIONS (Bavestrello er al., 2000; Stechow, 1923a; own data): Hydranth 
with long tentacles about 0.7 mm, pedicel (as delimited by perisarc) about 0.2 mm; 
gonophores 0.64 mm high and 0.4 mm wide; medusoids 1.1 mm high; egg size 
80-100 um; for nematocyst sizes see Bavestrello er al. (2000). 


BıoLoGy: The biology of this species is described by Bavestrello er al. (2000). 
The colonies can be found from 1 to more than 100 m depth (Motz-Kossowska, 1905), 
always on Schizoporella longirostris growing on hard substrata like rock or the bivalve 
Pinna nobilis. The hydroid is most easily collected by examining discarded glass 
bottles lying on sandy bottoms. Female medusoids can swim from five days before 
spawning. Male medusoids remain inactive. The two types of hydranths show two 
different trophic strategies. The larger ones feed preferably on meiobenthic organisms 
like nematodes and polychaetes, but also the larvae of the bryozoan host. The slender 
hydranths with few short tentacles in turn engulf the tentacles of the bryozoan host and 
feed on its collected small food particles. The tentacles of the bryozoan seem not to be 
harmed, though. 

Perarella schneideri hydroids show a marked seasonality, being present from 
September through June with a maximum in February. No hydroids can be found 
during summer. The period of medusoid production is between the second half of May 
to the end of June. 


DISTRIBUTION: Endemic to the Mediterranean. Type locality: Motz-Kossowska 
(1905) did not explicitly specify a type locality, but the only locality mentioned is 
Pollensa (island of Mallorca). 


REMARKS: The hydranth size of 15 mm given in the original description is likely 
a misprint for 1.5 mm as it does not match the size of the hydranth shown in figure. 

Picard (1958) listed Stylactella else-oswaldae Stechow, 1921 as a synonym of 
P. schneideri and I concur. In his description, Stechow (1923a) did not mention the 
substrate his colony was growing on. The type specimen of Stylactella else-oswaldae 
is kept by the ZSM and Dr. Ruthensteiner told me that the hydroid is indeed growing 
on an encrusting bryozoan which adheres to a piece of stone. Perigonimus steinachi 
Jickeli, 1883 was described as growing on a bryozoan, but Jickli’s description is 
inadequate and the species remains indeterminate. 

The hydranths of Perarella schneideri have no clearly demarcated hydrocaulus, 
but the basal part in a perisarc tube (Fig. 33A) must be homologized to a hydrocaulus. 


Perarella propagulata Bavestrello, 1987 Fig. 34 


Perarella propagulata Bavestrello, 1987: 19, figs 5.1-5.4. 
Cytaeis propagulata. — Bouillon et al., 2004: 55, fig. 32F-H. 


MATERIAL EXAMINED: Holotype, Museum of natural History of Genoa, C. E. 47716-a; 
Italy, Bay of Paraggi; 21 April 1983, depth 10 m, female; soft tissues quite macerated. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 281 


B 


Fic. 34 


Perarella propagulata Bavestrello, 1987; modified from Bavestrello (1987). (A) Two hydranths 
and a male sporosac, scale bar 0.5 mm. (B) Male sporosacs, scale bar 0.1 mm. (C) Female 
sporosacs, same scale as B. (D) Propagule issuing from stolon, scale bar 0.1 mm. 


DragnosIs: Perarella with sessile hydranths lacking basal perisarc collar; 


sporosacs rather elongate, spindle shaped, no radial canals; propagule formation from 
stolons. 


DESCRIPTION (Bavestrello, 1987): Hydroid living on Hinia incrassata shells, 
colonies stolonal, creeping reticulate stolons forming hydrorhiza, stolons in thin 


282 P. SCHUCHERT 


membranous perisarc. Hydranths monomorphic, without pedicels (sessile), without 
basal perisarc collar, spindle shaped, hypostome conical, 7-15 filiform tentacles in one 
whorl. Gonophores on stolons, without pedicels; sporosacs without radial canals, 
bulbs, and velar opening, very elongate, spadix spanning about two-thirds of the total 
height, spadix disappears after liberation of gametes; colonies gonochoristic, female 
sporosacs with 15-25 eggs. Some colonies produce propagules from stolons, tubes of 
naked coenosarc, surrounded at base by perisarc tube. Nematocysts: microbasic 
euryteles and desmonemes. 


DIMENSIONS: Hydranths approximately 0.6 mm high; sporosacs up to 0.3 mm 
high and 0.1 mm wide; propagules 0.3-0.5 mm; microbasic euryteles 6 x 3 um, 
desmonemes 4.5 x 3 um. 


BIoLoGy (Bavestrello, 1987): This species grows on shells of Hinia incrassata 
(Miiller), either inhabited by the gastropod or by hermit crabs. Only a small fraction 
(1-2%) of the investigated shells had the hydroid. It occurs in depths of 5-10 m. 
Hydranths are present from April to May. Colonies kept in the laboratory released 
numerous propagules which can spread the hydroid to other suitable substrates. 


DISTRIBUTION: Only known from the region near the type locality. Type locality: 
Paraggi Bay, Liguria, Italy, Mediterranean, 10 m. 


Genus Paracytaeis Bouillon, 1978 
TYPE SPECIES: Paracytaeis octona Bouillon, 1878, by monotypy. 


DIAGNOSIS: Cytaeididae with eight marginal tentacles; with four opaque inter- 
radial exumbrellar spots. 


Paracytaeis octona Bouillon, 1978 


Paracytaeis octona Bouillon, 1878: 149, fig. 8. — Bouillon, 1980: 319, fig. 6. — Bouillon et al., 
1988: 95, pl. 1 figs 1-2. — Goy er al., 1991: 106, fig. 14. — Bouillon et al., 2004. 56, 
fig. 33B. 


DIAGNOSIS: See genus diagnosis. 


DESCRIPTION: Umbrella bell shaped, with four opaque interradial exumbrellar 
spots of vacuolated cells; velum narrow; eight marginal tentacles, with large tentacular 
bulbs, adaxial epidermal nematocyst pad clasping bulb. Manubrium conical, half as 
long as umbrellar height, mouth circular with up to 20 oral adnate capitate tentacles; 
with medusa buds in four interradial clusters in upper part of manubrium. Gonads 
interradial, covering manubrium almost entirely. 


DIMENSIONS: Umbrella up to 2.5 wide and 3 mm high. 


DISTRIBUTION: Seychelles, Papua New Guinea, Mediterranean. Type locality: 
Ternay Bay, Mahe Island, Seychelles, Indian Ocean. 


REMARKS: This species has not yet been recorded in European waters. The 
Mediterranean records are based on medusae found in the eastern Mediterranean near 
Lebanon by Goy et al. (1991). The hydroid of this species is unknown. 


FAMILY RATHKEIDAE RUSSELL, 1953 emended 


TYPE GENUS: Rathkea Brandt, 1835. 
SYNONYM: Lizzinae Russell, 1953. new synonym 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 283 


DiAGnosıs: Medusa bell-shaped, manubrium either with four elongated lips 
forming simple or branched oral arms with terminal knobs, or perradial unbranched 
oral tentacles inserted vertically or very oblique at level of mouth margin; immature 
animals with or without orthogonally arranged interradial medusa buds. With four or 
eight radial canals and a circular canal. Usually eight bulbs, interradial ones with more 
than one tentacle, rarely only four bulbs and four tentacles. Ocelli absent. Gonads 
encircle manubrium completely 

Hydroids (where known) colonial, arising from ramified stolons. Hydranths 
monomorphic, without pedicel, with one whorl of filiform tentacles. Medusa buds arise 
from stolons, or more rarely from base of hydranths. 


REMARKS: In contradistinction to most taxonomic systems in current use, Lizzia 
blondina Forbes, 1848 is here thought to be more closely related to the Rathkeidae 
Russell, 1953 than the Bougainvilliidae. This decision is primarily based on the 16S 
phylogeny given in Fig. 39, but there is also a potential synapomorphy shared by Lizzia 
blondina and Rathkea octopunctata (M. Sars, 1835): the highly ordered, orthogonally 
arranged medusae budding from the interradial manubrium wall (Chun, 1896; Berrill, 
1952; Werner, 1956). Furthermore, both species resemble each other so closely 
(symmetry, size, form, manubrial peduncle) that they are easily confounded, although 
these traits are difficult to see as synapomorphies. Despite the polyp of Lizzia remains 
unknown, the genus was usually included in the Bougainvilliidae (Russell, 1953) due 
to the presence of perradial oral tentacles. This view was thereafter adopted in virtually 
all important subsequent works (e. g. Kramp, 1961; Bouillon, 1985; Bouillon & Boero, 
2000b), although Mayer (1910) regarded Lizzia and Rathkea as synonyms (which is 
correct in the formal sense, see below). It was generally asserted that the oral tentacles 
of Lizzia originate above the mouth like in Bougainvillia, which is not true. A closer 
examination shows that the position and attachment of oral tentacles of Lizzia blondina 
are slightly different from those found in the Bougainvilliidae. They become free at the 
level of the lips and not at some distance above the rim. They are also attached very 
obliquely and remain adnate to the manubrium for quite some distance, being 
continued as ridges along the manubrium accompanied by underlying vacuolated 
gastrodermal cells (Fig. 36D). The same situation is also found in Rathkea species 
(Fig. 35E). The same arguments also apply to Hydractinia minuta (Mayer, 1900) and 
H. minima (Trinci, 1903), which will be removed from the Hydractiniidae. Hydractinia 
minuta is regarded as a conspecific with Lizzia blondina and the new genus Podo- 
corynoides is proposed to accommodate Cytaeis minima Trinci, 1903. The diagnosis 
was therefore adapted to fit the new scope of the family. 


KEY TO THE GENERA: 


la four-radialicanal srt alt lerne o ii gta i ur ia 2 
lb eiehtsradialicanalseg to gut e Ra i o a I an a Ao Allorathkea* 
2a obliquely or vertically inserted oral tentacles with a distinct, round pedicel ........ 3 
2b oralFarms are elongated comers’of the mouth.) e e I 46-00. se Rathkea 
3a eight marginal bulbs, perradial ones sometimes with more than one tentacle . . . . Lizzia 
3b foumbulbs each with one tentacleionly, peer yarn. MR RES Podocorynoides 


* not present in area under investigation 


284 P. SCHUCHERT 


Genus Rathkea Brandt, 1838 


TYPE SPECIES: Oceania blumenbachii Rathke, 1835. 
SYNONYMS: Margellium Haeckel, 1879. — Currently also Lizzia Forbes, 1846 [type 
species Cytaeis octopunctata M. Sars, 1835, by monotypy and original designation]. 


DIAGNOsIs: Rathkeidae medusa with four radial canals; eight marginal tentacle 
bulbs, all with two or more tentacles; the four perradial corners of the mouth drawn out 
so as to form oral arms with clusters of nematocysts. Hydroid colonial, small, sessile, 
with long filiform tentacles, medusa buds arise from stolons. 


REMARKS: There is only one species of Rathkea medusa known in the region 
under investigation, but a second Rathkea-like polyp occurs. O’Sullivan (1984) gives 
an overview on all known species. 


Rathkea octopunctata (M. Sars, 1835) Fig. 35 


Cytaeis octopunctata M. Sars, 1835: 28, pl. 6 figs 14a-g. 

Oceania Blumenbachii Rathke, 1835: 21, pl. 1 figs 1-4. 

Lizzia octopunctata. — Forbes, 1848: 64, pl. 12 fig. 6. 

Lizzia grata A. Agassiz, 1862: 100, figs 28-29. 

Rathkia Blumenbachii. — L. Agassiz, 1862: 345 [incorrect spelling]. 

? Circe hyalina Van Beneden, 1867: 95, pl. 3 fig 14. 

Margellium octopunctatum Haeckel, 1879: 95. 

Margellium gratum. — Haeckel, 1879: 95. 

Lizzia shimiko Kishinouye, 1910: 25, pl. 5. 

Rathkea Blumenbachii. — Brandt, 1838: 353. — Neppi & Stiasny, 1913: 34, pl. 2, figs 23. — 
Hartlaub, 1911: 229, figs 196-199. — Hartlaub, 1917: 408. 

Rathkea octopunctata. — Haeckel, 1879: 97. — Mayer, 1910: 177, pl. 20 fig. 11. — Kramp, 1926: 
58, chart X. — Rees & Russell, 1937: 71, figs 7-8. — Russell, 1938b: 153, figs 26-32. — 
Russell, 1953: 137, pl. 7 figs 3-4, text figs 65A-E, 66, 67A-B. — Werner, 1958: 138, figs 
1-13. — Kramp, 1959a: 103, fig. 75. — Kramp, 1961: 72. — Kühl, 1962: 218, figs 7-9. — 
Kramp, 1968: 30, fig. 74. - Naumov, 1969: 212, figs 80-81.— Russell, 1970: 236. — Goy, 
1973: 979. — Arai & Brinckmann-Voss, 1980: 33, fig. 16. — O’Sullivan, 1984: 868, tab. 
2. — Schuchert, 1996: 59, fig. 34a-c. — Bouillon et al., 2004: 77, fig. 44D-H. 


MATERIAL EXAMINED: BMNH 1985.9.2.15; Great Britain, Plymouth; May 1937, coll. 
W.J. Rees; ca. 50 medusae. — France, Brittany, Roscoff; April 1998; few living medusae from 
plankton; not preserved; 16S sequence accession number AM183140. — Roscoff; few living 
medusae, not preserved; 10-20 m; 5 June 2000. — Norway, Raunefjord, living medusae, not pre- 
served; 0-3 m; 14 June 2006; 5 medusae; 16S sequence AM411415. - MHNG INVE 33454; 
New Zealand, Hauraki Gulf, Devonport, Narrow Neck Beach 0-1 m; 2-3 July 2002; several 
medusae, examined alive; 16S sequence AM411416. 

DIAGNOSIS: Four radial canals; eight marginal tentacular bulbs, four perradial 
bulbs each with 3-5 tentacles, four interradial bulbs with 2-3 tentacles; black pigment 


in manubrium and bulbs; oral tentacles bifid, terminal and lateral nematocyst clusters. 


DESCRIPTION (Werner, 1958; own observations): Hydroids arising from 
creeping, reticulate stolons, stolons enclosed in thin perisarc. Hydranths small, uni- 
form, without pedicel, cylindrical to spindle-shaped, base with thin gelatinous perisarc, 
one whorl of 4-6 tentacles, hypostome conical. Tentacles very thin and long. Medusa 
buds arise from stolons or rarely from base of hydranth, pear shaped, larger than 
hydranths, advanced stages with relatively long pedicel. Nematocysts: microbasic 
euryteles, desmonemes. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 285 


Fic. 35 


Rathkea octopunctata (M. Sars, 1835). (A) Hydroid colonies, right one with stolonal medusa 
bud; scale bar 0.5 mm; redrawn from Kühl (1962). (B) Lateral view of sexually mature medusa, 
scale bar 0.4 mm, combined after living and preserved animals. (C) Oral view of animal depicted 
in B. (D) Manubrium with medusa buds of a medusa of the same size as shown in B. (E) 
Enlarged mouth region in perradial view, from Russell (1953). (F-G) Aboral view of younger 
stages with fewer tentacles than shown in B, from Russell (1953). 


286 P. SCHUCHERT 


Primary medusae (newly liberated medusae originating from polyp stage) 
similar to adults, eight bulbs, three tentacles per perradial bulb, 1-2 tentacles per inter- 
radial bulb, manubrium with up to 4 medusa buds, oral lips not branched, terminal 
capitula not much developed. The primary medusae have a more elongate bell than the 
secondary medusae budded from the manubrium of a medusa. 

Mature medusa with globular bell, slightly higher than wide or as wide as high; 
jelly relatively thick, thickened at apex (up to 1/3 of total umbrella height); with gastric 
peduncle of variable length, shorter than manubrium length; dilated velum spanning 
1/2 of radius. Four radial canals and ring canal narrow. Eight marginal tentacular bulbs 
with long filiform tentacles, four perradial bulbs, each with three to five tentacles, four 
interradial bulbs, each with two to three tentacles; bulbs with dark pigment in gastro- 
dermis but no ocelli. Manubrium cylindrical, not reaching beyond umbrella margin in 
full extension; gastrodermis with dark pigment. Mouth with four lips, each divided at 
its extremity into two short arms, both ends with a spherical nematocyst cluster, in fully 
developed animals also a pair of lateral nematocyst knobs on common trunk of oral 
arms; bases of oral arms continued along manubrium for some distance as perradial 
ridges with vacuolated gastrodermal cells. Perradial medusa buds, appearing in regular 
opposite succession, up to 16 buds per manubrium, usually fewer; budding stops with 
the onset of gonad maturation. Gonads completely surrounding stomach, some indi- 
cation of perradial separation. Colour of marginal tentacular bulbs pale yellowish, dark 
yellowish brown or deep brown, appearing black by transmitted light. Colour of 
stomach brownish or black, persists even when fed on carotenoid rich Artemia nauplii. 
Nematocysts: euryteles and desmonemes. 


DIMENSIONS: Hydranth size 0.15-0.5 mm, tentacle length up to 3 mm (Rees & 
Russell, 1937; Werner, 1958; Kühl, 1962). Newly liberated medusa 0.9 mm (Kühl, 
1962). Sexually mature medusa usually 2-3 mm high and about as wide, height of up 
4 mm have been reported. Eggs 0.14 mm in diameter, planulae 0.2-0.24 mm long and 
80-90 um wide (Rees & Russell, 1937). For nematocyst sizes see Russell (1938b) and 
Schuchert (1996). 


ADDITIONAL DATA: The medusa budding process on the manubrium has been 
investigated by several authors (e. g. Chun, 1896; Berrill, 1952; Bouillon, 1962). The 
buds are derived from epidermal cells only. They develop in an ordered succession. 
The medusa buds of the hydroid stage derive as in other species from both epidermal 
and gastrodermal cells (Bouillon & Werner, 1965). The sexes are usually separate but 
hermaphroditic medusae are known to occur (Kulikova, 1957). 


DISTRIBUTION: North-western Europe from Norway to Strait of Gibraltar; 
Mediterranean; Black Sea; Barents Sea and White Sea; Iceland; Greenland; Hudson 
Strait; Newfoundland; New England; Bermuda; Aleutian Islands; Kamchatka; northern 
Japan; British Columbia; Victoria, Australia; New Zealand (Kramp, 1959a; Arai & 
Brinckmann-Voss 1980; O’Sullivan 1984; Schuchert, 1996). Kramp (1926) reviews all 
European records. Type locality: Bergen, Norway. 

BioLoGY: Due to its small size, the polyp has only rarely been found in nature 
(Kühl, 1962; Teissier, 1965; Karlsen, 1980; ? Watson, 1998). Kühl (1962) found it in 
shallow pools on Mytilus shells and stones; Teissier (1965) reports oyster shells as sub- 
strate. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 287 


The medusa occurs in coastal waters, sometimes also in lagoons and brackish 
waters (Kühl, 1962; Barnes, 1994). It may also survive in polluted waters (Beyer, 
1968) and can occur in long lasting mass aggregations (Zelickman et al., 1969). 

Being quite a common medusa, there exist a number of investigations on its 
seasonality (Kramp, 1926; Russell, 1953; Werner, 1958, 1962; Kühl, 1962; 
Rasmussen, 1973; Ballard & Myers, 2000). 

In the North Sea and most of the adjacent areas, the occurrence of the medusa 
is limited to the cold seasons of the year. It appears during the months of October to 
December, and is always to be found in small numbers in the winter plankton. In the 
spring, from April to May, there is a great increase in numbers, then within a brief time 
the medusa disappears almost entirely. In the Mediterranean and the Black Sea, the 
medusa was described as occurring during the spring, whereas it is a distinct summer 
form in the Arctic boreal area. 

The medusa has the ability to reproduce asexually by budding medusae. Two 
different generations of medusae are therefore to be recognized, i.e. a) the primary 
medusae originating from medusa buds of the hydroid, b) the secondary medusae 
produced by the budding process on the manubrium of the primary medusa. The 
secondary medusae are also able to reproduce asexually in the same way and give rise 
to more generations. It is because of the asexual reproduction of the medusae that there 
is a rapid increase medusa numbers during the spring. 

Werner (1956, 1958, 1962) made detailed experiments to demonstrate how the 
temperature influences the budding of medusae from the polyp and medusa stage. The 
life cycle of R. octopunctata is determined by seasonal temperature changes. Rapidly 
falling temperatures in autumn induce the polyps to develop medusae which then 
appear in the winter plankton. The young medusae multiply by budding more medusae 
until rising water temperatures in spring induce the development of gonads and later 
sexual reproduction. The medusae then disappear from the plankton by late summer 
(North Sea). 

Rathkea octopunctata is a nonselective polyphage which has been observed 
feeding on a number of animals including Clytia, Pleurobrachia, rotifers, Cladocora, 
copepods, nauplii, trochophores, crab zoeae and eggs, Sagitta, appendicularians, fish 
larvae (for references see Arai & Brinckmann-Voss, 1980). 

As an abundant species it is an influential consumer of other planktonic 
organisms (Zelickman et al., 1969; Matsakis & Conover, 1991). 


REMARKS: The synonym and taxonomic history of this species is outlined in 
Kramp (1926) and Russell (1953). Kramp (1926) also provided arguments why the 
name R. octopunctata M. Sars, 1835 is to be preferred over R. blumenbachii (Rathke, 
1835). 

The three available 16S sequences of R. octopunctata did not form a mono- 
phyletic group, the sequence from New Zealand being quite distant (Fig. 39). The 
animals from New Zealand differ also slightly in their morphology: the oral tentacles 
are about two times as long and they have a distinct, round trunk inserted at the mouth 
rim. The oral tentacles are paired and they resemble more those of Lizzia octostyla than 
those of Rathkea octopunctata from the Atlantic. It is possible that the New Zealand 
Rathkea medusae belong to a separate species. 


288 P. SCHUCHERT 


Rathkea spec. 


MATERIAL EXAMINED: Living hydroids observed in June 1998, September 2004, and 
September 2006; originating from Ile de Callot, Baie de Morlaix, France, English Channel; 
always embedded in Haliclona simulans (Johnston). 


DIAGNOSIS: Hydroid 0.1-0.2 mm high, Rathkea-like but tentacles shorter, 
predominantly four tentacles, polyp embedded in surface of the sponge Haliclona 
simulans (Johnston), colonial, stolons without perisarc. Nematocysts: heteronemes, 
6 x 2.5 um; desmonemes 4 x 2.5 um. 


REMARKS: This hydroid occurs regularly in the sponge Haliclona simulans 
colonized by the hydroids Dipurena simulans Bouillon, 1965 and D. halterata (Forbes, 
1846). There are usually hundreds of hydranths per sponge. They are extraordinary 
small and possess four tentacles only. The hydroid lives immediately below the sponge 
surface and only the short tentacles protrude beyond the surface. It can withdraw 
completely into the host tissue and apparently it is able to retract its tentacles 
completely into the hydranth body. The polyps are very difficult to observe and I only 
became aware of them when I tried to feed small Artemia nauplii to the Dipurena 
hydroids and the nauplii were efficiently captured by the almost invisible hydroid. The 
Artemia diet rendered the hydroids orange and better observable. Gonophores could 
not be found (June, and September). The shorter tentacles and the host species make it 
unlikely that this is Rathkea octopunctata. It could potentially belong to Lizzia blon- 
dina, a common medusa in the southern English Channel. 


Genus Lizzia Forbes, 1846 
TYPE SPECIES: Currently Cytaeis octopunctata Sars, 1835, see below. 


DraGNOSIS: Medusae with four or eight simple, unbranched perradial oral 
tentacles inserting at level of mouth rim; with manubrial peduncle; eight or excep- 
tionally 16 tentacle bulbs, each with one or more simple tentacles, usually with more 
tentacles on the perradial than the interradial bulbs; interradial medusa buds develop 
on manubrium; gonads enveloping manubrium. 


REMARKS: When Forbes (1846: 286) defined the genus Lizzia he stated 
«.founded for Cytaeis octopunctata of Sars...». This type species designation was 
iterated in Forbes (1848) when he added Lizzia blondina Forbes, 1848 to this genus. 
The type species of Lizzia is thus Rathkea octopunctata (M. Sars, 1835) by original 
designation as well as by monotypy. This leads to the awkward situation that Lizzia 
Forbes, 1846 is actually a junior synonym of Rathkea Brandt, 1838. As the genus 
Lizzia is well known and it has been used consistently for the rather common species 
Lizzia blondina Forbes, 1848, it is in the interest of nomenclatural stability to maintain 
this usage. After the publication of this study, a case to the International Commission 
on Zoological Nomenclature will be made asking them to use their plenary power to 
overrule Forbes’s type species designation and select Lizzia blondina instead. 

The polyps are not known. 


KEY TO THE LIZZIA SPECIES RECORDED FOR EUROPE: 


la four.solitary.oralitentacles..... ilari EE 2 
lb eishtioral’tentacles in four perradial pairs OE Lizzia octostyla 
2a without ocelli, interradial bulbs normally with one tentacle only ...... Lizzia blondina 


2b with adaxial ocelli, interradial bulbs with two tentacles ........... Lizzia elisabethae 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 289 


Lizzia blondina Forbes, 1848 Figs 36-38 


Lizzia blondina Forbes, 1848: 67, pl. 12 fig. 4a-f. — Browne, 1895: 265. — Hartlaub, 1911: 145, 
figs 131-135.— Kramp & Damas, 1925: 266, figs 13-14. — Kramp, 1926: 52, chart IX. — 
Russell, 1938b: 153, figs 33-36. — Russell, 1953: 145, figs 69-71, not 72, pl. 7 figs 1-2, 
pl. 34 figs 5-6. — Kramp, 1959a: 105, fig. 78. — Kramp, 1961: 87. — Brinckmann-Voss, 
1970: plate 8 fig. 2. — Goy, 1973: 980. — Bouillon er al., 2004: 47, fig. 28A-D. 

Lizzia spec. Claparède, 1860: 401, pl. 32, fig 1-2. 

? Dysmorphosa fulgurans A. Agassiz, 1865: 163, figs 259-260. 

Cubaster gemmascens Haeckel, 1879: 76, pl. 6 figs 8-12. — Kramp & Damas, 1925: 266, 
synonym. 

Dysmorphosa minima Haeckel, 1879: 78, pl. 6 fig. 7. — Browne, 1895: 261. — Browne, 1897: 
148. — Hartlaub, 1911: 145, synonym. 

[not Cytaeis minima Trinci, 1903 = Podocorynoides minima (Trinci, 1903)] 

Lizzia claparèdei Haeckel, 1879: 82.— Hartlaub, 1911: 145, synonym. 

? Lizzia minuta McIntosh, 1893: 344. 

Lizzia claparédei. — Chun, 1896: 36, figs 2-4. — Braem, 1908: 795, figs 4-5 

Dysmorphosa minuta Mayer, 1900b: 41, pl. 18 fig. 42. new synonym 

[not Hydractinia minuta Bonnevie, 1898a = H. carica Bergh, 1887] 

Podocoryne minuta. — Mayer, 1910: 140, pl. 14 fig. 1. — Kramp, 1959a: 102, fig. 68. — Kramp, 
1961: 69. — Goy, 1973: 978. — Goy et al., 1991: 107, fig. 18. 

In part Podocoryne minuta. — Neppi & Stiasny, 1913: 24, pl. 2 fig. 13a-b. 

Lizzia fulgurans. — Brinckmann-Voss, 1970: plate 8 fig. 1. 

Podocoryna minuta. — Schuchert, 1996: 50, fig. 28a-b. 

Hydractinia minuta. — Bouillon et al., 2004: 66, fig. 39F-G. 


MATERIAL EXAMINED: BMNH 1985.9.11.10; Ireland, Valentia Harbour; 05 August 1938; 
few medusae. - BMNH 1985.9.12.16; Great Britain, Plymouth; 7 June 1937; >200 mature 
medusae, 1-3 perradial tentacles. — Atlantic, France, Roscoff, 20 September 2006, 5 medusae, 
depth 5-15 m, with 2 tentacles per perradial bulb (stumps when collected), 16 S sequence made 
of 2 medusae gave identical sequence AM411423; some cultivated for 3 weeks, secondary 
medusae mostly with eight tentacles only. - MHNG INVE48742, Norway, Ranefjord, 0-3 m, 
16.06.2006, about 10 fully grown medusae, examined alive, all with 8 tentacles only, cultivated 
for two weeks to sexual maturity, not further tentacles developed, colour changed to orange 
when fed with Artemia nauplii; 16S sequence AM411417. — Mediterranean, France, 
Villefranche-sur-Mer, 7 April 2005, 6 medusae, initially identified as Hydractinia minuta, all 
specimens used to extract DNA, direct sequencing of amplified the 16S gene showed it to be 
polymorphic, the fragments were therefore cloned using standart techniques and then sequenced 
again, AM183124 and AM411419. - ROMIZ B775, as Lizzia fulgurans; Italy, Naples, 28 June 
1961, collected by A. Brinckmann-Voss (mentioned or depicted in Brinckmann-Voss, 1970; 
1987); about 20 medusae. - USNM 58065; as Hydractinia minuta; Belize, Carrie Cay lagoon; 
coll. 1977; 7 medusae; det. R. J. Larson. - USNM 58063; as Hydractinia minuta; Belize, Carrie 
Cay Lagoon; 16 May 1977; 7 medusae; det. R. J. Larson. - USNM 58064; as Hydractinia 
minuta; Belize, Carrie Cay Lagoon; 23 May 1976; 7 medusae; det. R. J. Larson. — New Zealand, 
Wellington Harbour, Seatoun; several medusae, examined alive, from surface plankton collected 
11-12 July 2002, all with 8 tentacles only, therefore identified initially as Hydractinia minuta, 
16S sequence determined from several animals AM411418. 


DIAGNOSIS: Lizzia with four oral tentacles, each perradial bulb with 1-3 tenta- 
cles, four interradial bulbs each with a single tentacle only, rarely two; medusa buds on 
manubrium, with manubrial peduncle; no ocelli. 


DESCRIPTION: Umbrella spherical, sometimes a little higher than wide, usually 
with a shallow, rounded apical process; jelly moderately thick, especially in apical 
region; gastric peduncle present, slim, usually shorter than manubrium. Four radial 
canals and ring canal narrow, radial canals along manubrial peduncle with enlarged 
gastrodermal cells. Eight marginal bulbs, without ocelli, adaxial side with nematocyst 


290 P. SCHUCHERT 


Fic. 36 


Lizzia blondina (Forbes, 1848). A) Diagrammatic reconstruction of sexually mature female 
medusa in side view, scale bar 0.2 mm. (B) Oral view of medusa shown in A. (C) Manubrium of 
medusa before gonad maturation and with medusa buds. (D) Oral region in side view, note the 
origin of the oral tentacles very close to the margin of the manubrium, scale bar 50 um. (E-G) 
Growth stages in aboral view. A-D, after preserved material; E-G, modified from Russell (1953). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 29] 


Fic. 37 


Lizzia blondina (Forbes, 1848), after living material from Norway, lateral and oral view, scale 
bar 0.4 mm. Note that this form with one tentacle per bulb has previously often been named 
Hydractina/Podocoryne minuta or Lizzia fulgurans. 


cushion, four perradial ones each with one to three tentacles, four interradial bulbs with 
a single tentacle, rarely two. Manubrium short, not reaching beyond umbrella margin 
in full extension, cone-shaped, either with broad quadrangular base attached to 
peduncle or perradial corners somewhat extending along peduncle, gastrodermis 
thickened interradially; medusae budding from stomach wall in interradial position, 
medusa budding phase overlaps with gonad maturation phase; four unbranched oral 
tentacles, attached very obliquely and adnate for some distance, lower axils at level of 
mouth-margin, oral tentacles relatively long but contractile, each with one terminal 
nematocyst cluster. Gonads completely surrounding stomach in a form of ring-shaped 
cushion. Colour of marginal tentacle bulbs and stomach pale straw, yellow, or 
yellowish brown, depends on food items. Nematocysts (Russell, 1938b): microbasic 
euryteles and desmonemes. 


DIMENSIONS: Sexually mature medusa height around 1 mm, maximally 2 mm; 
earliest stages of vegetatively produced medusa 0.5-0.7 mm. Eggs measure 0.08- 
0.12 mm in diameter (Chun, 1896). Nematocysts see Schuchert (1996). 


VARIATION: The perradial bulbs can have 1-3 tentacles. Mature medusae with 
three perradial tentacles are quite rare. Medusae budded from a medusa are released 
with one tentacle per bulb, therefore there are regularly also representatives of this 
stage in a population. Also the eight-tentacled medusae can have gonads. The size of 
mature medusae is quite variable. 

The interradial bulbs of Lizzia blondina have mostly only one tentacle, but 
occasionally some of the bulbs may have two tentacles, but never all of them. 


FURTHER DATA: The earliest known stages of medusa have already eight mar- 
ginal bulbs, each with one tentacle; scattered nematocysts on exumbrella; medusa buds 
are present at a very early stage, they are borne in two, or rarely three series and in each 


292 P. SCHUCHERT 


FIG. 38 
Lizzia blondina (Forbes, 1848), living medusa with buds from Roscoff. 


series there are four interradial buds approximately at right angles; the proximal series, 
nearest the base of the stomach, is the first to develop and the buds in each series show 
a graded order of development in that the two oldest buds stand opposite one another. 
The gonads start to ripen towards the termination of asexual reproduction and ripe 
sexual cells may be present while some buds still remain unliberated. 

The histology of the vegetative medusa budding was investigated by Chun 
(1896, as Lizzia claparedei) and Braem (1908). The sexes are separate (Chun, 1896). 


DISTRIBUTION: Lizzia blondina occurs from Norway to the Mediterranean as 
well as in the NW Atlantic (Allwein, 1967; Russell, 1970). It has been recorded from 
Norway as far north as Bergen (Kramp & Damas, 1925; Rees, 1952), around Denmark 
(Hartlaub, 1911; Kramp, 1926; Rasmussen, 1973); North Sea (Hartlaub, 1911); all 
round the British Isles, being scarcest in the eastern end of the English Channel and the 
southern North Sea (Russell, 1953; Fraser, 1973; Russell, 1970); Iceland (northernmost 
record), The Faroes, and Faroe-Shetland Channels (Hartlaub, 1911; Kramp, 1926); 
France (Hartlaub, 1911; Teissier, 1965); Iberian Peninsula from the Bay of Biscay to 
the Mediterranean (Medel & Löpez-Gonzälez, 1996); western Mediterranean 
(Hartlaub, 1911; Goy, 1973). Under the name Podocoryne minuta it has also been 
recorded along the coast of western Africa (Thiel, 1938), Florida (Mayer, 1900b, as 
Dysmorphosa minuta), and New Zealand (Schuchert, 1996). Type locality: Originally 
described from the Shetland Islands (Sound of Brassay and Fitful Head). 


BIOLOGY: Lizzia blondina is a summer species found in nearshore waters. 
Owing to the rapid proliferation by asexual budding this species may at times be 
exceedingly abundant. It occurs in the plankton around the British Isles from March to 
December, but more abundantly in May to October (Russell, 1953; Ballard & Myers, 
2000). In Norway, the medusae appear in June and last until end of October (Kramp & 
Damas, 1925; own data). In the Kattegat, they are present from at least July to 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 293 


November or December (Kramp, 1926). In the Mediterranean, the species can be found 
all year round (Goy, 1973; Riera er al., 1986). 


REMARKS: The life cycle of this otherwise relatively common medusa is not 
known. Kramp (1926) suspected that the polyp lives mainly along the southern and 
western coasts of the British Isles and the distribution in the Danish waters let Kramp 
think that the polyp lives only in areas with soft, clayey bottoms. Perhaps the polyp 
described above as Rathkea spec. is its polyp. 

The synonymy and history of this species is treated by Russell (1953) but new 
data suggest that at least also Dysmorphosa minuta Mayer, 1900 must be added to the 
synonymy. | 

Since the medusae may be found mature at any stage of the tentacle develop- 
ment, the identity of the species was at first not certain and three separate species were 
described (see Fig. 36E-G, Dymorphosa minima one tentacle per bulb, Lizzia clapa- 
redei two perradial tentacles, Lizzia blondina three tentacles per radial bulb). Asexually 
produced Lizzia blondina are always liberated with eight bulbs and eight tentacles and 
will typically develop more perradial tentacles. However, also the eight tentacles stage 
can become sexually mature. As these forms co-occurred with the normal blondina 
form, they could quite easily synonymized (Browne, 1897; Hartlaub, 1911). 

Kramp & Damas (1925) and Kramp (1926), however, also attributed popu- 
lations from Norway and Denmark to this species, despite that the animals of these 
populations always have one perradial tentacle only. Such pure eight-tentacled Lizzia 
populations (Fig. 37) would otherwise be identified as Podocoryne/Hydractinia minuta 
(Mayer, 1900b) or Lizzia fulgurans. Kramp (1961) already suspected that the 
Mediterranean records of Podocoryne minuta by Neppi & Stiasny (1913) were actually 
L. blondina. A careful comparison of Hydractinia minuta material from the 
Mediterranean (see Material examined) confirmed that it is indistinguishable from 
eight-tentacled Lizzia blondina originating from Great Britain. Likewise, eight- 
tentacled Lizzia from Norway were otherwise indistinguishable from Lizzia blondina 
from the English Channel. This similarity was also confirmed by their 16S sequence 
data (Fig. 39), the two sequences differed in a single base pair only. 

The purported (Hartlaub, 1911: 144) difference of the insertion site of the oral 
tentacles of Lizzia blondina and Hydractinia minuta does not exist. The oral tentacles 
in Lizzia insert very close to the mouth margin, they are very oblique (Fig. 36D) and 
filled with chordoid gastrodermal cells. They thus insert lower than usually seen in 
adult Bougainvillia spec. (but also in young Bougainvillia spec. the insertion can be 
quite close to the mouth rim). 

Also material from New Zealand, previously identified by myself (Schuchert, 
1996) as Podocoryne minuta, conforms with the eight-tentacle-stage of Lizzia 
blondina. 

Some preliminary comparisons of 16S sequence data confirm the hypotheses 
outlined above (Fig. 39). Lizzia blondina from the English Channel (specimens with 
two perradial tentacles) is part of a well supported clade that includes also the samples 
referable to the Hydractinia minuta morphotypes. Lizzia blondina from the Channel 
showed only a single base pair difference to the Norwegian specimens. Likewise, 
Lizzia blondina differed in only one or two positions (of 570 compared base pairs) 


294 P. SCHUCHERT 


Leuckartiara octona AM411421 

Hydractinia borealis AY787878 

Hydractinia minuta New Zealand AM411418 
ZO blondina Norway AM411417 
lHydractinia minuta Mediterranean AM411419 


Hydractinia minuta Mediterranean AM183124 
Lizzia blondina Britanny AM411423 


Rathkea octopunctata New Zealand AM411416 


Rathkea octopunctata Britanny AM183140 
Hydractinia minima New Zealand AM411420 
Hydractinia minima Mediterranean AM183125 
Koellikerina fasciculata AM183129 

Bougainvillia muscus AY787880 


Fic. 39 


Molecular phylogeny of some Rathkeidae based on partial 16S sequences and Maximum 
Likelihood analyses using the software PHYML (HKY model, default parameters). The numbers 
near some nodes indicate the percentage of bootstrap support (1000 replicates nonparametric 
bootstrap analysis as implemented in PHYML). Initial analyses included many other 16S 
sequences of a variety of Filifera species (see also Schuchert & Reiswig, 2006), but no signi- 
ficant relationship of the Rathkeidae could be detected. In the analysis shown here, only a few 
selected outgroup taxa were used in order to improve the readability. The topology of the 
Rathkea clade did not change significantly by reducing the number of outgroup taxa. 

The accession numbers of the sequences are given after the species name. In order to better 
visualize the revision made in the text, the initial species identifications are given. Hydractinia 
minuta must be synonymized with Lizzia blondina, Hydractinia minima is referred to 
Podocorynoides minima. 


from Mediterranean medusae initially identified as H. minuta. These differences are 
certainly attributable to intraspecific variation (comp. also Fig. 4 and remarks for B. 
muscus). The Lizzia clade also includes the sequence of H. minuta from New Zealand, 
although it is more distant (14 bp difference). The good support of the Lizzia clade and 
the relative minor sequence differences suggest that H. minuta is nothing but a form of 
Lizzia blondina. 

This raises the question whether also the original descriptions of Dysmorphosa 
fulgurans A. Agassiz, 1865 and Dysmorphosa minuta Mayer, 1900 were actually based 
on L. blondina. There is nothing in Agassiz’s description that would make his D. ful- 
gurans distinguishable from eight-tentacled Lizzia blondina, except for the presumably 
lower number of tentacles in newly liberated medusae. This seems to be an error. 
Agassiz (1865: fig. 260) depicts a manubrium with an advanced bud that has at least 
eight bulbs and tentacles. Because Mayer (1910) mentions that it can have occasionally 
up to 16 bulbs, I somewhat hesitate to synonymize it with L. blondina. It is possible 
that the form with 16 bulbs belongs to a different species. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 295 


Mayer (1900b) kept his Dysmorphosa minuta distinct from Dysmorphosa 
fulgurans A. Agassiz, 1865 based on colour and size differences (bell size of 0.3 mm 
for D. minuta). His Dysmorphosa minuta were not sexually mature and could thus well 
be small, newly liberated medusae. The colour of the bulbs of Lizzia blondina depends 
on food items. 

I tried to locate the type material of both D. fulgurans and D. minuta but could 
neither find them in the HCZH nor the NMNH. The type material of both species is 
likely lost. Medusae identified as Podocoryne minuta originating from Belize, which 
can be considered as belonging to the same biogeographic region as Florida (type loca- 
lity of D. minuta), were indistinguishable from European Lizzia blondina with eight 
tentacles. I therefore think that Dysmorphosa minuta Mayer, 1900b and likely also 
Dysmorphosa fulgurans A. Agassiz, 1865 are nothing but eight-tentacled L. blondina. 

Cytaeis minima Trinci, 1903, which Mayer (1910) considered to be only a 
young stage of D. minuta, is clearly a distinct species that always has four tentacles 
only. 


Lizzia octostyla (Haeckel, 1879) Fig. 40 


Dysmorphosa octostyla Haeckel, 1879: 78, pl. 6 fig. 6. 

Podocoryne octostyla. — Mayer, 1910: 140. 

Lizzia octostyla. — Neppi & Stiasny, 1913: 32, pl. 2, fig 22. — Kramp, 1959a: 106, fig. 81. — 
Kramp, 1961: 88. — Brinckmann-Voss, 1970: plate 8, fig 3. — Bouillon et al., 2004: 47, 
Fig. 28F. 

Koellikerina fasciculata. — Kramp, 1957c: 118, fig. 1. 


MATERIAL EXAMINED: ROMIZ B773, Naples, Mediterranean, 6 medusae collected by A. 
Brinckmann-Voss, 22 February to 24 March 1961, damaged. 

DraGNOSIS: Medusa 0.5-1 mm, low apical apex, with peduncle, eight oral 
tentacles in perradial pairs, medusa buds on manubrium, eight marginal bulbs, 8-12 
tentacles. 


DESCRIPTION: (Neppi & Stiasny, 1913; own observations) Adult medusa 
spherical; jelly thin, shallow apical process; manubrium on a conical peduncle; radial 
canals narrow; eight rather small marginal bulbs, interradial ones slightly smaller, no 
ocelli but dark pigment in bulbs; in younger specimens one tentacle per bulb, in older 
ones perradial bulbs with paired tentacles; tentacles long, very expandable; eight 
unbranched adradial oral tentacles inserting slightly above mouth margin, oral 
tentacles in pairs, ends with distinct capitulum (nematocyst cluster); interradial medusa 
buds on middle of manubrium; gonads as flat pads on manubrium wall, medusae 
budding can continue after gonad maturation. 


DIMENSIONS: Medusa bell 0.4-1.3 mm high and wide, usually around 0.5 mm. 
BIOLOGY: Recorded in the months February, March, and August to October. 


DISTRIBUTION: Mediterranean only; Ligurian Sea (Kramp, 1957d, as juv. 
Koellikerina fasciculata); Tyrrhenian Sea (Brinckmann Voss, 1970); Adriatic Sea 
(Neppi & Stiasny, 1913; Benovic & Lucic, 1996); Ionian Sea (Haeckel, 1879). Type 
locality: Island of Corfu, Greece. 


REMARKS: This is perhaps only a form of Lizzia blondina. 


P. SCHUCHERT 


296 


Fic. 40 
Lizzia octostyla (Haeckel, 1879), schematic reconstruction after preserved material, scale bar 
0.2 mm. (A) Lateral view. (B) Oral view. 


Lizzia elisabethae Haeckel, 1879 Fig. 41 


Lizzia Elisabethae Haeckel, 1879: 83, pl. 6 fig. 12. — Hartlaub, 1911: 149, fig 136. — Russell, 
1953: 150. — Kramp, 1959a: 105, fig. 70. — Kramp, 1961: 88. 


DIAGNOSIS (Kramp, 1959a): Umbrella 6 mm high, 4 mm wide, with thick apex, 
perradial bulbs with four, interradial bulbs with two tentacles; an ocellus at the base of 
each tentacle; medusa buds not observed. 

DISTRIBUTION: Type locality only: Jersey, English Channel. 


REMARKS: Lizzia elisabethae has never been seen again since its original 
description by Haeckel (1879). Its size and ocelli would make it certainly quite distinct, 


Na 


PTT 
he a 
SLR 
SST 
È II x 
ì II 


Fic. 41 
Lizzia elisabethae Haeckel, 1879, after Hartlaub (1911) from Haeckel (1879), originally drawn 
after life, scale bar 2 mm. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 297 


however some doubts on its validity remain, Russell (1953: 153) thought that it might 
have been an abnormal L. blondina while Kramp (1959a, 1961) kept it as valid. 


Genus Podocorynoides n. gen 
TYPE SPECIES: Cytaeis minima Trinci, 1903. 


DraGNOsIS: Medusae with four vertical, unbranched oral tentacles inserting at 
level of mouth rim; with manubrial peduncle; four bulbs, each with one simple 
tentacle; interradial medusa buds on manubrium; gonads enveloping manubrium. 


REMARKS: The 16S sequence data (Fig. 39) indicate that Podocorynoides 
minima (Trinci, 1903) is closely related to Lizzia and Rathkea and it does not belong 
to the Hydractiniidae as usually assumed (Mayer, 1910; Kramp, 1959a, 1961). 
Referring it to the Rathkeidae made it necessary to transfer it to the new genus 
Podocorynoides. The name gives an allusion to its similarity to medusae formerly 
included in the genus Podocoryna. 


Podocorynoides minima (Trinci, 1903) n. comb. Fig. 42 


Cytaeis minima Trinci, 1903: 26, pl. 1 figs 1-30. 

[not Dymorphosa minima Hackel, 1879: 78, pl. 6 fig 7.— Hartlaub, 1911: 145 = Lizzia blondina] 

Podocoryne simplex Kramp, 1928: 45, fig. 20. — Russell, 1953: 136, synonym. — Kramp, 1968: 
28, fig. 70. 

Podocoryne minima. — Russell, 1953: 134, figs 63-64. — Kramp, 1961: 69. — Chow & Huang, 
1958: 176, pl. 1 figs 10-11. — Kramp, 1968: 28, figs 69-70. — Brinckmann-Voss, 1970: 
pl. 7 fig. 1. — Uchida & Sugiura, 1977: 53, figs 1-2. - Goy et al., 1991: 107, fig. 17. 

In part Podocoryne minuta. — Neppi & Stiasny, 1913: 24, pl. 1 fig 12 (not others). 

[not Dysmorphosa minuta Mayer, 1900] 

Podocoryna minima. — Schuchert, 1996: 50, fig. 27. 

Hydractinia minima. — Bouillon et al., 2004: 65, 39D-E. — Chaplygina & Dautova, 2005: 142, 
fig. 1. 


MATERIAL EXAMINED: Mediterranean, Ligurian Sea, Bay of Villefranche-sur-Mer; depth 
0-70 m; 6 April 2005; 6 medusae; no material preserved, DNA made from 4 medusae, 16S 
sequence accession number AM183125.— New Zealand, Hauraki Gulf, Motutapu Island; 0-3 m; 
30 June 2002; plankton; no material preserved, DNA made from several medusae, 16S sequence 
accession number AM411420. 


DIAGNOSIS: See genus diagnosis. 


DESCRIPTION: Medusa with globular bell; apical jelly slightly thickened in fully 
grown animals; relatively long gastric peduncle present (up to 1/3 of bell cavity). With 
four radial canals and ring canal, rather narrow. With four perradial tentacle bulbs and 
tentacles, without ocelli, adaxial epidermis of bulbs thickened and pad-like, gastro- 
dermis with brown pigment, tentacles very contractile. Manubrium length 1/2 of bell 
cavity, cylindrical, dark pigment in interradial gastrodermis, perradial lip margins elon- 
gated to short oral tentacles terminating in a spiny, spherical knob of nematocysts. 
Interradial medusa buds on manubrium with ordered budding sequence. Gonads 
surrounding stomach completely. Nematocysts (Chaplygina & Dautova, 2005): 
microbasic euryteles and desmonemes. 


DIMENSIONS: Bell size 0.3-0.9 mm (Uchida & Sugiura, 1977; own data). For ne- 
matocyst dimensions see Chaplygina & Dautova (2005). 


298 P. SCHUCHERT 


Fic. 42 


Podocorynoides minima (Trinci, 1903) n. comb., mouth gaping, size about 1 mm, after photo- 
graphs and living material from the Mediterranean. 


OTHER DATA: The buds on the manubrium are of epidermal origin (Trinci, 1903). 
The medusae are gonochoristic. The development of the secondary medusa is 
described by Uchida & Sugiura (1977). There are no significant differences in size and 
general structure between medusae with gonads and those without them. Even animals 
with gonads can continue to bud medusae. 


BroLOGY: In the Mediterranean, the medusa is present at least during the months 
March to September (Trinci, 1903; Goy, 1973; Brinckmann-Voss, 1987). 


DISTRIBUTION: Mediterranean (e. g. Trinci, 1903; Goy, 1973; Brinckmann- Voss, 
1987; Goy et al., 1991; Benovic & Lucic, 1996); English Channel (Russell, 1953); 
Brazil (Vannucci, 1957); North Carolina (Allwein, 1967); Red Sea (Schmidt, 1973a); 
New Zealand (Schuchert, 1996); Papua New Guinea (Bouillon, 1980); north-western 
Pacific (Kramp, 1928; Chow and Huang, 1958; Uchida & Sugiura, 1977). Type 
locality: Gulf of Naples, Mediterranean. 


REMARKS: Although they have a distinct, round trunk, the oral tentacles of this 
species resemble much the Podocoryna (=Hydractinia) type and it is quite surprising 
to find this species more closely related to the Rathkeidae than the Hydractiniidae. The 
oral tentacles are mostly held in a nearly vertical position, thus somewhat unlike the 
oblique tentacles of Lizzia blondina. 


FAMILY PANDEIDAE HAECKEL, 1879 


DiaGNOSIS: Hydroid stage normally stolonal, exceptionally branching. 
Hydranths bearing one whorl of filiform tentacles (exceptionally two whorls, scattered 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 299 


tentacles or no tentacles). Perisarc developed to a variable degree, occasionally missing 
completely, but often also enveloping hydranth as a filmy pseudohydrotheca. 
Reproduction is by free medusae (except in some genera of questionable affınity). 
Medusae with or without an apical projection; manubrium large, mounted or not 
on a peduncle; mouth with either four simple, or crenulated, or complexly folded lips. 
With four (rarely eight as in Octotiara) radial canals, often broadened to a band; 
exceptionally centripetal canals can be present; with or without mesenteries. Gonads 
either with a smooth surface or complexly folded; on the manubrium in adradial or 
interradial position; in a few species extending onto the radial canals. Tentacles hollow, 
originating from conical tentacle bulbs tapering gradually into tentacles; tentacles 
never with terminal nematocyst clusters (capitation). Sometimes with rudimentary 
tentacles, cirri-like tentacles, or marginal protuberances. Ocelli present or not. 


REMARKS: The Pandeidae comprise a large number of genera and some need a 
revision. Therefore, the key below relates only to medusa based genera used in this 
study. For a complete key of the Pandeidae genera see Bouillon & Boero (2000). 


KEY TO THE ADULT PANDEIDAE MEDUSAE OF THE NE ATLANTIC AND MEDITERRANEAN: 


la cishtradialicanalspatià TR. AMEN ARR RR SE Octotiara 
lb fourgradialicanalst RO RINCARO el N. 2 
2a bwollarscehperradialitentaelese A. a Re RENNES 5 
2b KOUOISMOTEHENtacles ES EVE LTT ee I TS RS 4 
3a four interradial gonads that extend onto subumbrella and forming a bell like cavity 

IA NN I ESTA ET 0e cp AI cle NN Codonorchis 
3b SONAUS UNIRCADONE VEN EDEN AE ea. era RARES Amphinema 
4a gonads not on folds or in pits, surface smooth or rugose ................... Merga 
4b LONIASOMSIOMACNHOLAS-OLANIPUS CRA TONE 5 
5a manubrium short, perradial stomach corners drawn out into a large cross-like form 

en EN PERI III SSR IOTA TI NI SPERI RIO CIA OI ARTI OT Annatiara 
5b Stomachilargestlask=SMAPEA A Dre Re AR RN 6 
6a with mesenteries (elongated connection of stomach and radial canals)............. 7 
6b IMÉSCMETIÉS ASEM nari wend cada ee US Lan Enr ATELIER II Halitholus 
7a gonads forming a reticulated network of pits. ............02 een. Pandea 
7b gonads with folds only, or folds and pits combined ........................... 8 
8a gonads in long folds that tend to be vertical in interradial region, some pits . Catablema 
8b adradialbrowstol¢conad folds ss cpr. re acti seins MER 9 
9a no or very few pits, gonad folds directed towards perradial, with ocelli . . . Leuckartiara 
9b with many interradial pits, gonad folds directed towards interradial, without ocelli 

ET OEP EC SI RE EN ed ee ANA AL Moe at Neoturris 


Genus Pandea Lesson, 1843 emended 

TYPE SPECIES: Dianaea conica Quoy & Gaimard, 1827, designation by Mayer, 1910. 

SYNONYM: Campaniclava Allman, 1864a: 352 [type species Syncoryne cleodorae 
Gegenbauer, 1854]. 

DiaGNOSIS: Hydroids, where known, forming stolonal colonies with naked 
hydranths on very short pedicels; with filiform tentacles. Dactylozooids may be 
present. Medusa buds arise from stolons. 

Medusa with or without apical projection, with or without longitudinal ridges 
and nematocyst tracks on exumbrella; lips large and much folded; gonads at first in the 
adradii and eventually covering the manubrium, forming a complex reticulated 
network; radial canals band-like; more than eight tentacles, without rudimentary 
marginal tentacles or marginal warts; with or without abaxial ocelli. 


300 P. SCHUCHERT 


KEY TO THE PANDEA SPECIES TREATED HERE: 


la with exumbrellar ribs and nematocyst tracks, gonad network coarse, marginal 
bulbs laterally compressed, ocelli present, subumbrella colourless ..... Pandea conica 
lb without exumbrellar ribs and nematocyst tracks, gonad network close meshed, 
marginal bulbs not much laterally compressed, ocelli absent, manubrium and 
Fadialicanals:deepiredì o it. „u mn AUS See Oe Pandea rubra 
Pandea conica (Quoy & Gaimard, 1827) Figs 43-44 


Dianaea conica Quoy & Gaimard, 1827: 182, pl. 6 figs 3-4. 

Oceania sedecimacostata Kölliker, 1853: 324. 

Syncoryne cleodorae Gegenbauer, 1854: 166, pl. 1 fig 3. — Picard, 1956: 1, synonym. 

Campaniclava cleodorae.— Allman, 1872: 261. 

Tiara reticulata Haeckel, 1879: 60, pl. 3 fig. 11. — Vanhöffen, 1911: 209, synonym. — Hartlaub, 
1914: 340, synonym. 

Tiara pileata var. ampullacea Haeckel, 1879: pl. 3 fig. 7. 

Pandaea conica. — Haeckel, 1879: 54 [incorrect subsequent spelling]. 

Pandea conica. — Mayer, 1910: 118, fig. 63. — Hartlaub, 1914: 339, figs 283-287, synonymy. — 
Uchida, 1927: 214, fig. 38. — Kramp, 1924: 8, fig. 7. — Ranson, 1936: 84, pl. 1 figs 1-6. 
— Russell, 1953: 207, figs 107-110. — Picard, 1956: 1, figs 1-3. — Kramp, 1961: 111. — 
Brinckmann-Voss, 1970: pl. 11 fig. 2. — Russell, 1970: 250, figs 14s-15s. — Winkler, 
1982: 34, fig. 6. — Lalli & Gilmer, 1989: 127, fig. 47b. — Pages et al., 1992: 14, fig. 13. 
— Schuchert, 1996: 71, fig. 42a-c. — Bouillon et al., 2004: 74, fig. 43A-B. 


MATERIAL EXAMINED: MHNG INVE35427, polyp colonies on two specimens of Clio 
cuspidata, Villefranche-sur-Mer, with numerous medusa buds. - MHNG INVE35433, about 7 
medusae from Villefranche-sur-Mer, identified 1911 by C. Hartlaub. - MHNG INVE35434, 
Naples, 2 badly preserved medusae. — Serial sections of Campaniclava cleodorae from the 
Picard collection (kept by Station Marine d’Endoume, Marseille), leg & det. J. Picard, coll. 
1933, Villefranche-sur-Mer France. 

DIAGNOSIS: Pandea medusa with exumbrellar ribs bearing nematocyst tracks, 
marginal tentacles laterally compressed, with ocelli, subumbrella colourless, 16-24 
tentacles. Polyps on Clio cuspidata, polymorphic, feeding zooid almost sessile, two 
whorls of tentacles, with tentaculozooids, medusa buds stolonal, medusae released 
with two tentacles. 


DESCRIPTION: Polyp stage living on the pelagic gastropod Clio cuspidata; 
polyps polymorphic, with gastrozooids and tentaculozooids, arising from creeping 
stolons. Stolons fine, covered by thin perisarc, long straight stretches, not much 
ramified, bifurcations usually coincide with polyp positions. Polyps with a very short 
pedicel, almost sessile, base in membranous perisarc cup, body spindle shaped, 
diameter about 1/3 of height, hypostome high (about 1/5 of total height), conical to 
nipple-shaped, 8-10 filiform tentacles in two closely set whorls, distal tentacles longer, 
gastrodermis chordoid. Tentaculozooids present but infrequent, usually near gastro- 
zooids, about as high as gastrozooids, with slightly swollen tip. Medusa buds arise 
directly from stolons, short pedicel, buds covered with thin perisarc. Nematocysts: 
microbasic euryteles and desmonemes. 

Newly liberated medusa (Picard, 1956) as high as wide, bell-shaped with 
moderately thick jelly. Jelly at apex interrupted by remains of the gonophore stalk. 
Manubrium length 1/2 of bell cavity, conical with four very inconspicuous lips, no 
gonads visible. With four broad, smooth radial canals and circular canal. Two opposite 
larger and two smaller tentacle bulbs present. Only larger bulbs bear a tentacle. Ocelli 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 301 


Fic. 43 


Pandea conica (Quoy & Gaimard, 1827). (A) Part of hydroid colony on Clio cuspidata, with 
gastrozooid, medusa buds on stolons, and tentaculozooid, scale bar 0.3 mm. (B) Newly released 
medusa. A, after preserved material from the Mediterranean; B, redrawn from Picard (1956). 


not present, but a dispersed red pigment present in bulbs. Above bulbs on exumbrella 
patches of tissue with nematocysts that later grow towards top. 

Mature medusa bell-shaped to conical, usually higher than wide, with small 
rounded or conical apical projection covered by opaque epidermis; with longitudinal 
keel-like exumbrellar ribs, corresponding to number of marginal tentacles, crest of ribs 
with a line of thickened epidermis studded with nematocysts (nematocysts tracks) 
originating from marginal tentacle; jelly fairly thick. Four radial canals fairly broad, 
with smooth or jagged outlines, forming mesenteries; ring canal narrower, with smooth 
outlines. Sixteen to 24 (sometimes as many as 45) smooth, marginal tentacles, each 
with a conical, laterally compressed bulb that clasps umbrellar margin, with one 
abaxial ocellus, abaxial spur continued as nematocyst track up to top of bell, no rudi- 
mentary marginal tentacles. Stomach voluminous, almost filling upper half of sub- 
umbrellar cavity, length variable, usually about half of subumbrella cavity in length, 
attached to subumbrella along perradii for about 2/3-4/5 of its length (mesenteries). 
Mouth with four perradial lips with much folded or crenulated margins, beset with 
nematocysts but these not in clusters. Gonads interradial, in fully grown specimens 
covering upper 2/3 of the manubrium wall, not continuous as separated perradially by 
mesenteries, gonad forming coarse-meshed network of ridges with pits between. 
Colour of stomach and gonads, reddish, brownish or yellowish; mouth-lips reddish or 
pink; tentacles milky yellow; ocelli red or reddish brown; subumbrella colourless (in 
part after Russell, 1953). Nematocysts of medusa: microbasic euryteles. 


DIMENSIONS: Mature medusa 10-20 mm high; polyps about 1 mm. For nemato- 
cyst sizes see Picard (1956) and Schuchert (1996). 


OTHER DATA: Ranson (1936) describes the growth and development of the 
medusa (summarized in Russell, 1953). Also Mayer (1910) depicts young medusae. In 
younger stages the gonads have an interradial cleft (= horse-shoe shaped). 


302 P. SCHUCHERT 


Pandea conica (Quoy & Gaimard, 1827); adult medusa, after preserved material from the 
Mediterranean, scale bar 2 mm. 


BıoLoGY: The polyps of this species occur on the shell of the pteropod Clio 
cuspidata (see Picard, 1956; Lalli & Gilmer, 1989). In the Mediterranean, Kramp 
(1924) recorded the medusa from January to September. Bleeker & van der Spoel 
(1988) found the medusa at night in depths of 45-95 m, during the day in 50-300 m. 


DISTRIBUTION: An oceanic medusa recorded from the Mediterranean; the 
Atlantic Ocean from Scotland to Bermuda, South Africa and Patagonia; East Africa; 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 303 


Sri Lanka; Central North Pacific, California; southern Japan; China; Vietnam; 
Philippines; Rarotonga; Papua New Guinea; east of Australia; New Zealand (Ranson, 
1936; Russell, 1953; Kramp, 1968; Bleeker & van der Spoel, 1988; Pages er al. 1992; 
Schuchert, 1996). The polyp phase has been recorded from the Mediterranean, South 
Atlantic and Indian Ocean (Picard, 1956). However, its host Clio tricuspidata has a 
much wider distribution and probably also the polyp stage of Pandea conica. Type 
locality (medusa): Strait of Gibraltar. 


REMARKS: The correlation of the polyp stage, previously known as Campani- 
clava cleodorae (Gegenbauer, 1854), and Pandea conica was established by Picard 
(1956). Picard observed only the newly released medusa and he apparently did not 
cultivate it to older, more distinctive stages. The only character that allow a tentative 
identification of the young medusa were incipient exumbrellar nematocyst tracks. 
Although Campaniclava cleodorae is very likely the polyp stage of Pandea conica, it 
seems desirable that this is proven again by more conclusive data. 

The tentaculozooids observed in the present material have not been reported so 
far. It could be that they are not formed in all colonies, a phenomenon also observed in 
several other hydroids. 

There seems to be a difference in the cnidome between Pandea conica of 
Europe and New Zealand (comp. Schuchert, 1996). The large isorhiza found in animals 
from New Zealand could not be found in Mediterranean medusae (this study and 
Picard, 1956). It could be that the New Zealand Pandea belongs to a separate species. 

There are two other Pandea species in the Atlantic: Pandea rubra Bigelow, 
1913 and Pandea cybeles Alvarino, 1988. Pandea rubra lacks an apical projection and 
the exumbrellar nematocysts. Pandea cybeles from the Sargasso Sea differs more 
gradually from P. conica, mainly in having about twice as many tentacles, a broader 
bell, and a larger manubrium. Alvarino (1988) tabulates more differences of these three 
species. 


Pandea rubra Bigelow, 1913 Fig. 45 


Pandea rubra Bigelow, 1913: 14, pl. 2, figs 1-7. — Kramp, 1926: 96, pl. 2 fig. 15, chart XVII. — 

Bigelow, 1938: 107. — Russell, 1953: 211, figs 11-112. — Kramp, 1961: 112. — Naumov, 

1969: 207, fig. 75. — Kramp, 1968: 51, fig. 135. — Arai & Brinckmann-Voss, 1980: 61, 

fig. 34. — Bleeker & van der Spoel, 1988: 231, fig. 14, map fig. 15. — Brinckmann- Voss 

& Arai, 1998: 50, fig. 5. 

MATERIAL EXAMINED: ZMUC; Discovery station 1889; 55.887°S 32.780°W, South 
Atlantic; 1200-1500 m; 10 March 1937; 1 much damaged medusa. — ZMUC; 54.083°N 
26.133°W, North Atlantic; 0-1000 m; 15 July 1913; 1 fragmented medusa (material of Kramp, 
1926). -— ZMUC; 6.867°N 79.500°E, Indian Ocean; 300-1000 m; 24 Nov. 1929; 1 much damaged 
medusa. — Photograph of living medusa kindly provided by Dr K. Rascoff, medusa observed in 
June 2001 near Oceanographers Canyon (off Massachusetts), depth about 700 metres. 


DIAGNOSIS: Pandea species without exumbrellar nematocyst tracks; gonad 
forming fine-meshed network of pits, no ocelli, manubrium and radial canals deep-red. 


DESCRIPTION (Russell, 1953): Umbrella bell-shaped, as high as or slightly 
higher than wide; with rounded summit without marked apical process; without 
exumbrellar nematocyst tracks; jelly very soft and fragile. Four radial canals, broad, 
with wavy or jagged outlines; ring canal broad, with smooth outlines. Radial canals 


304 P. SCHUCHERT 


A 
I) 
= 
= 
= 
=; 
SI 
Ù 
& 
= 
=; 
= 
3 


Fic. 45 

Pandea rubra Bigelow, 1913. (A) Young, though fertile medusa. (B) Manubrium of a larger 
specimen. (C-D) Lateral views of tentacle bulbs, arrow points at abaxial spur, in C it is partially 
embedded in the jelly. A, after photograph of living animal kindly provided by Dr. K. Raskoff; 
B, from Hartlaub (1914); C-D, redrawn from Kramp (1926) and Brinckmann-Voss & Arai 
(1998). 

narrow proximal, wider distal. Seven to twenty-four marginal tentacles of varying 
sizes, much extensible, hollow, smooth, each with large conical basal bulb, not much 
laterally compressed, with distinct abaxial spur clasping margin of exumbrella. Abaxial 
spurs of bulbs either pigmented or not, may be embedded in the jelly of the exumbrella. 
No rudimentary tentacles. No ocelli. Stomach large, with broad base, about half height 
of subumbrellar cavity in length; attached to subumbrella along perradii for about 1/2 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 305 


or more of its length (mesenteries). Mouth with four lips with much folded and crenu- 
lated margins. Gonads situated interradially on stomach, forming very close-meshed 
irregular network of ridges with pits between. Colour stomach, mouth, gonads, radial 
and circular canals, and marginal tentacles deep brownish red to chocolate brown. In 
older or larger specimens, the red pigment spreads also over the subumbrella so that it 
can get opaque (Bigelow, 1913). 

Polyp stage unknown. 


VARIATION: The abaxial spurs of bulbs can either be pigmented or not, they may 
be embedded in the jelly of the exumbrella (Fig. 45C). A table with sizes and tentacle 
numbers of growth stages is given in Bigelow (1913) and Russell (1953). The smallest 
known specimens measured 5 mm (Brinckmann-Voss & Arai, 1998). The maximal 
number of tentacles so far observed is 32 (Brinckmann-Voss & Arai, 1998), usually 
there are only 12-24. 


DIMENSIONS: Medusa bell height rarely up to 75 mm, usually 30-40 mm. 


DISTRIBUTION: North and north-east Pacific (Bigelow, 1938; Arai & 
Brinckmann-Voss, 1980; Brinckmann-Voss & Arai, 1998); India (Vannucci et al., 
1970); Bermuda (Bigelow, 1938); north-eastern Atlantic (Kramp, 1920; Kramp, 1926; 
Fraser, 1973; Bleeker & van der Spoel, 1988); north-western Atlantic (K. Raskoff, 
pers. comm.); South Atlantic (Kramp, 1957a; Bleeker & van der Spoel, 1988). Type 
locality: Albatross Station 4797, NE Pacific, off south-east coast of Kamtchatka, 
0-548 m. 


BIOLOGY: A deep water species, occurring in all oceans, usually far away from 
the continents, maximal recorded depths 1750 m, usually in depths of several hundred 
meters. The specimens of the Atlantic were taken in July (Russell, 1953), May and July 
near Bermuda; in the autumn near the Azores (Bleeker & van der Spoel, 1988) As deep 
water species it is likely not much seasonal. 


REMARKS: A rare, though distinct and easily recognizable species. The 
mesogloea is very fragile, thus specimens taken with a plankton net are often much 
damaged. 


Genus Amphinema Haeckel, 1879 

TYPE SPECIES: Oceania dinema Péron & Lesueur, 1810, by designation of Rees & Russell 
(1937). 

DragNOsIS: Medusae with two large opposite tentacles; with or without ocelli; 
gastric peduncle absent; mostly with a considerable apical projection; with or without 
marginal small tentaculae or marginal warts; with or without mesenteries; four simple 
oral lips, not or not much crenulated. Gonads on manubrium in adradial or interradial 
position, occasionally extending to the radial canals. 

Hydroid colonies usually stolonal (except for one species of uncertain affinity), 
hydranths with a well developed caulus, caulus longer than hydranth, covered by 
perisarc; hydranths without pseudohydrotheca, spindle-shaped, conical hypostome, 
one whorl of filiform, amphicoronate tentacles. Polyps bend over when disturbed. 
Gonophores arise either from cauli, stolons or both, released as free medusae. 


306 P. SCHUCHERT 


REMARKS: See Rees & Russell (1937), Russell (1953), and Bouillon et al. 
(2000) for important revisions and the taxonomic history of the genus. The last men- 
tioned publication gives also a tabular overview on most Amphinema medusae known 
at that time. Two species have to be added, namely Amphinema tsingtauensis Kao, Li, 
Chang & Li, 1958 and Amphinema cheshirei Gershwin & Zeidler, 2003. The gender of 
the name is neuter (ICZN article 30.1.2.) It is possible that not all endings of the spe- 
cific epithets are correct, but it seems more desirable to maintain taxonomic stability 
and to use the names as given in Kramp (1961). 


KEY TO THE AMPHINEMA MEDUSAE TREATED IN THIS STUDY: 


la without. conspicuous apical chamber ©) ........... 2... ER eee 2 
lb with conspicuous apical chamber above manubrium .................. A. rubrum 
2a gonadsmot.extending;along.radial.canals;.....- trat 22 ET EEE 3 
2b gonads extending along radial'canalsi-. 1.0. a oe A. turrida 
3a red-brown manubrium, gonad surface rugose, cellular strands issuing from 
Tadıialkcanals er... nre te epee dense en das CITE A. krampi 
3b NOU ASFADOVE®. . inno liti ER 4 
4a with ocelli, with aboral processes of manubrium projecting into mesogloea A. bouilloni 
4b NOL AS ADOVE |S net, MIRI eil ra RE 5 
Sa folded gonads, with small tentaculae, tentacle bulbs brownish-yellow . ... A. rugosum 
Sb gonads smooth, no tentaculae, tentacles purple, manubrium green ......... A. dinema 
Amphinema dinema (Péron & Lesueur, 1810) Fig. 46 


Oceania dinema Péron & Lesueur, 1810: 346. — Goy, 1995: 255, plate. 

? Oceania diadema Lamarck, 1817: 506. — Haeckel, 1879: 50, synonym. 

Perigonimus serpens Allman, 1863: 10. — Allman, 1872: 327, pl. 11 figs 7-9. 

Saphenia apicata McCrady, 1859: 129, pl. 8 figs 2-3. 

Stomotoca apicata. — Fewkes, 1881: 152, pl. 2 figs 1, 4, 9. — Rittenhouse, 1910: 333, figs 1-32. 

Saphenia titania Gosse, 1853: 387, pl. 26, figs 7-9. 

Dinematella cavosa Fewkes, 1881: 151, pl. 2 figs 2-3, pl. 4 fig. 3. 

Amphinema titania. — Haeckel, 1879: 50, pl. 4 figs 8-9. 

Amphinema apicatum. — Haeckel, 1879: 50. 

Tiarula coeca Hartlaub, 1914: 261, fig. 215. 

Stomotoca dinema. — Mayer, 1910: 109, pl. 9 figs 8-10, pl. 10 figs 1-4. Neppi & Stiasny, 1913: 
18, pl. 1 fig. 8. 

in part Amphinema dinema. — Hartlaub, 1914: 259, fig. 215 [others = A. rugosum]. 

Amphinema dinema. — Rees & Russell, 1937: 62, figs 1-4. — Russell, 1938b: 153, figs 37-40. — 
Russell, 1953: 180, text-fig. 89, pl. 10, figs 1, 2,4, pl. 11 figs 1, 3.— Kramp, 1959a: 117, 
fig. 109. — Kramp, 1961: 93. — Kramp, 1968: 42, fig. 108. — Goy er al., 1991: 109, fig. 
24. — Schuchert, 1996: 63, fig. 36a-d. — Bouillon er al., 2004: 68, fig. 40B-C. 


MATERIAL EXAMINED: France, Brittany, Roscoff; 10-20 m; 06 June 2000; 1 medusa, ex- 
amined alive, 16S sequence AM183136. — BMNH 1985.9.2.20; Great Britain, Plymouth; 11 
October 1938; 50 medusae from plankton. - BMNH 1985.9.1.16; Great Britain, Plymouth; 25 
June 1937; hydroid colonies on Scalpellum from rearing experiments, leg. W. J. Rees. - BMNH 
1985.9.1.15; Great Britain, Plymouth; 5 July 1937; cultivated hydroid on glass, with medusa 
buds, from rearing experiments of W. J. Rees. - BMNH 1969.12.2.128; Great Britain, Plymouth; 
18 February 1937; hydroid reared from medusa, material of Rees & Russell (1937). 


DIAGNOSIS: Amphinema with simple, not folded gonads on each of the eight 
adradial surfaces of stomach; fourteen to twenty-four small marginal bulbs without 
tentacles; manubrium intensively green, tentacles purple. Hydroid stolonal, medusa 
buds only on stolons. Newly liberated medusa without apical process. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 307 


Fic. 46 


Amphinema dinema (Péron & Lesueur, 1810). (A) Part of hydroid colony with a medusa bud, 
scale bar 0.5 mm. (B) Newly liberated medusa, size about 0.7 mm. (C) Mature medusa in lateral 
and oral view, the gonads are not yet fully developed. A, after preserved material; B, modified 
from Rees & Russell (1938); C, after life. 


DESCRIPTION (Rees & Russell, 1937; Russell, 1953; own observations): Stolonal 
hydroid colonies arising from creeping, ramified stolons; hydrocauli not branched, 
enclosed in perisarc, perisarc at base with or without annulated region, perisarc 
becoming very thin distally and end is difficult to observe, perisarc can be infested with 
detritus. Hydranth spindle shaped with a dome shaped hypostome, tapering below 


308 P. SCHUCHERT 


gradually into hydrocaulus, below hypostome one whorl of 8-10 (max. 14) amphi- 
coronate tentacles. Medusa buds arise from stolons only, ovoid to pear-shaped, 
tapering below into short pedicel, pedicel shorter than bud, bud enclosed in thin 
perisarc membrane. Nematocysts: microbasic euryteles, desmonemes. Colour of 
hydranth brilliant orange-red. 

Newly liberated medusa spherical, jelly thin, no apical process present, 
scattered nematocysts on exumbrella, only two opposite tentacles, rather long, at place 
of other tentacles only small swellings (marginal warts); manubrium cylindrical, about 
1/3 of height of bell cavity; colour of tentacle bulbs and stomach reddish orange, 
developing soon into a vivid emerald green manubrium and violet tentacles. 

Mature medusa bell slightly wider than high, jelly of uniform thickness, with a 
large conical, solid apical process, shape and size of process very variable, process 
height usually depassing bell size, bell top with slight perradial furrows; dilated velum 
spanning 1/2 of radius. With four broad radial canals, no mesenteries, only two 
diametrically opposed tentacles with thick base, hollow, tentacles tapering evenly and 
very long (ten times bell size), additionally with 6-12 marginal warts, but no short 
tentacles. Manubrium intensively green, manubrium length 3/4 of subumbrella to 
reaching slightly beyond velum, cross-shaped in section, mouth with four prominent, 
slightly recurved lips. Eight gonads in adradial pads with smooth surface. Ocelli 
absent. Colours: proximal part of tentacles intensively purple. Nematocysts: only 
microbasic euryteles. 


DIMENSIONS (Rees & Russell, 1937; Russell, 1953; own observations): Hydroid 
total height 1.6-4 mm, hydranth body about 0.8-1.0 mm (transition to hydrocaulus 
gradual), hydrocaulus diameter 0.05-0.10 mm, stolons 0.04-0.08 mm; medusa buds 
ready for liberation 0.30-0.33 mm long and 0.20-0.23 mm wide. Newly liberated 
medusae 0.6-0.7 mm high. Bell size of mature medusa 1-2.3 mm, total height maxi- 
mally up to 6 mm and 4 mm wide; the apical process develops at a bell size of about 
1 mm; eggs 0.15 mm diameter; planulae 0.25 mm long and 0.09 mm wide. For 
nematocyst sizes and shapes see Russell (1938b) and Schuchert (1996). 


DISTRIBUTION: The medusa is widespread, it occurs in the eastern Atlantic from 
the Shetland Islands (Kramp, 1926) to the Gulf of Guinea in western Africa (Kramp, 
1955b) (see also Russell, 1953; Kramp, 1961; Russell, 1970; Fraser, 1973); Western 
Mediterranean (e. g. Neppi & Stiasny, 1913; Goy, 1973; Benovic & Lucic, 1996; 
Bouillon er al., 2004); Eastern Mediterranean (Goy et al., 1991); North America from 
Cape Cod to Florida (Mayer, 1910); Brazil (Vannucci, 1957); Indian Ocean (Ganapati 
& Nagabhushanam, 1958); Tropical Pacific Ocean (Kramp, 1953; Bouillon, 1980); 
New Zealand (Schuchert, 1996). The hydroid is known from British Isles (Allman, 
1872; Rees & Russell, 1937; Russell, 1957); the Atlantic coast of Spain (Medel & 
Löpez-Gonzälez, 1996) and Morocco (Patriti, 1970), New Zealand (Schuchert, 1996); 
Mediterranean (e. g. Motz-Kossowska, 1905). Type locality: English Channel. 


BıoLoGY: The medusa is widespread but usually only few specimens can be 
caught. In the English Channel, the hydroid has been found on stems of Eunicella 
verrucosa and frequently on the stalked barnacle Scalpellum in the months March to 
September (Rees & Russell, 1937; Russell, 1957). In the western Mediterranean, 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 309 


Motz-Kossowska (1905) found the polyp fixed on the bryozoan Cellaria fistulosa on 
muddy bottoms, medusae were produced during winter the months. Brinckmann-Voss 
(1987) reports that medusae are budded from May to August. Boero & Fresi (1986) 
give a depth range of 0.5-20 m, and as substrates algae, sponges and barnacles. The 
medusae mature in the Mediterranean from September onward (Neppi & Stiasny, 
1913). In the English Channel, the medusa appears in the plankton in late summer, 
being most abundant in September and October (Russell, 1953). Lebour (1922) 
describes how the medusa eats other planktonic organisms like Obelia and Eutima 
medusae, Calanus and Sagitta species. Rittenhouse (1910) attempted to cross A. ru- 
gosum and A. dinema without success. 


REMARKS: The life cycle of this species has been investigated by Rittenhouse 
(1910), Rees & Russell (1937), and Schuchert (1996). 

Contrary to the opinion of Rees & Russell (1937), the hydroid pedicel can also 
be annulated at its base (own observations and Russell, 1953). 


Amphinema rugosum (Mayer, 1900) Fig. 47 


Stomotoca rugosa Mayer, 1900a: 4, pl. 2 fig. 5. 

Saphenia dinema. — Forbes, 1848: 25, pl. 2 fig. 4. — Le Danois, 1914a: 18, fig. 5. 

[not Amphinema dinema (Péron & Lesueur, 1810)] 

In part Amphinema titania. — Haeckel, 1879: 50, pl. 4 figs 8-9. 

[not Saphenia titania Gosse, 1853 = Amphinema dinema] 

? Amphinema apicatum. — Brooks, 1883: 711. 

Stomotoca rugosa. — Mayer, 1910: 112, pl. 10 figs 5-6, pl. 11 figs 1-2. — Uchida, 1927: 202, 
fig. 32. 

in part Amphinema dinema. — Hartlaub, 1914: 259, fig. 214, 216-217, not 215. 

Amphinema rugosum. — Rees & Russell, 1937: 67, figs 5-6. — Russell, 1938a: 423. — Russell, 
1953: 183, figs 90A-B, pl. 10 fig. 3, pl. 11 figs 2 & 4. - Kramp, 1959a: 117, fig. 110. — 
Kramp, 1961: 94. — Kramp, 1968: 43, fig. 110. — Wedler & Larson, 1986: 96, fig. 7Aa- 
b. — Migotto, 1996: 15, figs 2f-g. — Schuchert, 1996: 64, fig. 37a-d. — Goy et al., 1991: 
109, fig. 25. — Bouillon et al., 2004: 69, fig. 40E-F. 


MATERIAL EXAMINED: BMNH 1985.9.1.14; Great Britain, Plymouth; 15 May 1937; in- 
fertile hydroid colony on floating cork. - BMNH 1969.12.2.135; Great Britain, Plymouth; June 
1937; slide preparation, part of material of Rees & Russell (of BMNH 1985.9.1.14). - BMNH 
1955.11.23.77-105; Great Britain, Plymouth; 1930-1934; about 15 medusae from plankton; coll. 
F. S. Russell. 

DIAGNOSIS: Amphinema medusa with folded gonads on each of the eight 
adradial surfaces of stomach; besides two large tentacles 14-15 reduced, small ten- 
taculae; manubrium and tentacles brownish-yellow to orange. Hydroid mostly stolonal, 
medusa buds only on stolons and hydrocauli. Newly liberated medusa with apical 
process. 


DESCRIPTION (Rees & Russell, 1937; Russell, 1953; own data): Hydroid stolonal 
or very rarely some hydrocauli branched once or twice, arising from creeping, ramified 
stolons; stolons covered by perisarc. Hydrocauli long, gradually expanding their 
diameter about two times from proximal to distal, without distinct demarcation of 
hydranth body and coenosarc of hydrocaulus; hydrocaulus covered by thin perisarc, at 
junction to stolons with annulated stretch (2-5 rings), upper end distinct, not forming a 
pseudohydrotheca, the perisarc can be infested with detritus. Hydranth cylindrical, 


310 P. SCHUCHERT 


FIG. 47 


Amphinema rugosum (Mayer, 1900). (A) Part of hydroid colony with medusa buds. (B) Newly 
liberated medusa, note presence of small apical process. (C) Mature male medusa in side view, 
scale bar 1 mm. (D) Same as in C, oral view. A-B, modified from Rees & Russell (1937); C-D, 
after preserved material from Plymouth. 


with a dome-shaped hypostome; below hypostome one whorl of 8 to 12 amphicoronate 
tentacles. Medusa buds arise from stolons and from hydrocauli, 1-3 buds per hydro- 
caulus can be present, pedicels of medusa buds shorter than bud height; bud is covered 
by thin perisarc. Nematocysts microbasic euryteles and desmonemes. Colour: 
hydranths and coenosarc possess a bright reddish orange in European specimens, 
perisarc horn coloured. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 311 


Newly liberated medusa spherical, with small apical process and usually the 
remains of an apical canal, exumbrella with scattered nematocysts; manubrium half the 
length of the subumbrellar cavity; two opposite tentacles, with or without two small 
opposite perradial tiny tentacles (tentaculae). Colour of tentacle bulbs reddish-orange, 
with faint tinge of yellow, stomach bright ochreish yellow. 

Adult medusa with bell slightly higher than wide, jelly uniformly thin besides 
top, on top a conical to hemispherical apical process, size variable; top of umbrella 
with slight perradial furrows. With four broad radial canals with jagged and smooth 
margins, normally without mesenteries, sometimes with short mesenteries. Two dia- 
metrically opposed tentacles with thick bases, tapering and very long (ten times bell 
size), bulbs conical, hollow; additionally 14-24 small, reduced tentaculae. Manubrium 
reaching almost to velum, cross-shaped in section, mouth with four prominent, slightly 
recurved lips. Eight gonads in adradial pairs, each with three to four oblique folds 
sloping downwards towards the interradii. The gonad can be fused perradially in the 
more oral part, while there can be an interradial connection at the top. Ocelli not 
present. Nematocysts: microbasic euryteles. Colour of marginal tentacle bulbs and 
stomach rich brownish yellow to orange, with central masses of deep purple-brown 
pigment; cores of two marginal tentacles ochre; marginal tentaculae colourless. 


DIMENSIONS (after Rees & Russell, 1937; Russell, 1953; own data): Total height 
of medusa when fully grown 5-6 mm, usually smaller. Newly liberated medusa 0.42- 
0.65 mm in height. Hydroid stems up to 2.0-3.5 mm (total height), hydranths 0.5-0.75 
mm; stolons 0.05-0.06 mm wide; diameter of hydrocaulus at base 0.10-0.15 mm, 
hydrocaulus distal end 0.2 mm; medusa buds up to 0.35 mm high and 0.25 mm wide, 
pedicels 0.13-0.15 mm. For nematocyst dimensions, see Russell, (1938b), Migotto 
(1996) and (Schuchert (1996). 


FURTHER DATA: Russell (1953) provides tables with details of the medusa 
development. The medusa is prone to abnormalities. The radial canals may be 
connected to the stomach by short mesenteries. It is not possible to distinguish the 
hydroids of A. dinema and A. rugosum if no medusa buds are present. There is no 
significant size difference and the perisarc annulation can also be present in A. dinema. 


BIoLOGY: The medusa of Amphinema rugosum occurs in the plankton at 
Plymouth usually from April to July, it has occasionally been seen as late as October 
(Russell, 1953). In daytime, the medusa lives generally in the deeper water layers 
below 20 m, but may migrate right to the surface at night. In south-western Ireland, 
Ballard & Myers (2000) recorded the medusa from July to September. 

The polyp stage occurs on a variety of hard substrata. In the Mediterranean, 
Boero & Freesi (1986) found it regularly in depths of 0.5-20 m. It was present all year 
round but more abundant during the winter months. It grew on algae, sponges, 
hydroids, bryozoans, and polychaete tubes. 

Rittenhouse (1910) attempted to cross A. rugosum and A. dinema without 
success. 


DISTRIBUTION: The medusa occurs circumglobally, it has been recorded along 
the American coasts from New England to Florida (Mayer, 1910); Caribbean Sea 
(Kramp, 1959a); in the eastern Atlantic from the Shetland Isles and the British Isles to 


312 P. SCHUCHERT 


the Azores (Hartlaub, 1914; Russell, 1953; Russell, 1957; Cornelius, 1992); rarely in 
the North Sea (Allwein, 1968); western and eastern Mediterranean (Brinckmann-Voss, 
1987; Goy et al., 1991); Red Sea (Schmidt, 1973a); Madagascar (Kramp, 1965); Japan 
(Uchida, 1927); New Caledonia, Sumatra, South China Sea (Kramp, 1965); New 
Zealand (Kramp, 1965; Schuchert, 1996); Papua New Guinea (Bouillon, 1980). 

The polyp has been found in the British Isles (Rees & Russell, 1937), Bay of 
Biscay (Medel & Löpez-Gonzälez, 1996); Mediterranean (Boero & Fresi, 1986); 
Puerto Rico (Wedler & Larson, 1986); Brazil (Migotto 1996); and New Zealand 
(Schuchert, 1996). No type locality was specified by Mayer (1900a). He based his 
description on medusae from Newport (Rhode Island), Charleston (South Carolina) 
and the Tortugas (Florida). 


REMARKS: Amphinema rugosum has frequently been confused with A. dinema 
until the life cycle was investigated by Rees & Russell (1937) who provided criteria to 
distinguish it from A. dinema. At least for the European fauna, the medusae and polyps 
with medusa buds are unambiguously distinguishable. In other regions this might be 
not more difficult. 

Migotto (1996) observed a hydroid colony and newly liberated medusae. The 
latter lacked an apical process. It is thus possible that this process is not always present 
in all newly liberated medusae. A similar observation was made by Schuchert (1996). 

There might be more Amphinema species that resemble A. rugosum and A. 
dinema. Rees (2000) describes a hydroid from California that had polyps matching 
Amphinema rugosum, but the immature and adult medusae resembled A. dinema. 

Amphinema rugosum var. shantungensis Chow & Huang, 1958 from Chefoo 
(China) differs from the typical form in lacking marginal tentaculae and being larger. 

Amphinema rugosum var. tsintanensis Kao, Li, Chang & Li, 1958 has ocelli and 
thus deviates significantly from the typical form. This variant was therefore justly 
raised to species level by Xu (1993) as Amphinema tsintanensis Kao, Li, Chang & Li, 
1958. 


Amphinema turrida (Mayer, 1900) Fig. 48 


Dissonema turrida Mayer, 1900b: 44, fig. 3-4. — Mayer, 1910: 116, pl. 10 fig. 1, pl. 22 fig. 1. 
Amphinema turrida. — Bigelow, 1909: 200, pl. 7 fig. 2, pl 40, fig. 6, pl. 43 fig 3, pl. 44 figs 3-4. 
— Kramp, 1959a: 118, fig. 112. — Kramp, 1961: 95. — Kramp, 1968: 42, fig. 106. — Goy, 
1973: 982, fig. 6. — Bouillon, 1980: 322, fig 8. — Bouillon et al., 2004: 69, fig. 40G. 
Stomotoca turrida. — Mayer, 1915: 199, pl. 1 fig. 1. 


MATERIAL EXAMINED: MNHN No. 1638; Bay of Villefranche-sur-Mer, Ligurian Sea, 
France; part of material described by Goy (1973), one much damaged two-tentacled medusa and 
two medusae with four perradial tentacles. 

DIAGNOSIS: Amphinema species with elongated, folded gonads that extend 
along 1/2 of radial canals; two opposite tentacles and many small tentaculae or rudi- 
mentary bulbs, most with ocelli. 


DESCRIPTION (Mayer, 1900b; Bigelow, 1909; Bouillon, 1980): Umbrella bell- 
shaped, somewhat higher than wide; with relatively small apical projection; above 
stomach a clear gastric chamber. Two opposite perradial tentacles, hollow, basal bulbs 
elongated, conical, laterally compressed; between the single pair of tentacles 12-16 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 313 


di 
Uf 
mes 
Fic. 48 


Amphinema turrida (Mayer, 1900), modified after Bouillon (1980), note that this figure 
represents the prevailing Pacific form which has no tentaculae but only rudimentary bulbs; scale 
bar 2 mm. 


small, solid tentaculae (Atlantic and Mediterranean), or up to 28 rudimentary bulbs 
(Pacific); all with red ocelli. Manubrium large, reaching nearly to velum level or 
beyond it, pyriform, mouth cruciform, lips simple, recurved. Gonads as adradial pairs, 
folded, extending along 3/4 of the subumbrella and manubrium thus forming long 
mesenteries. 


DIMENSIONS: Bell height 2-7 mm (Bigelow, 1909; Goy, 1973). 


OTHER Data: Mayer (1910) reports that females have 4-7 eggs in each of the 
four gonads. 


VARIATION: Most Pacific animals lack the tentaculae and have rudimentary 
bulbs only (Bigelow, 1909; Mayer, 1915; Bouillon, 1980). Bigelow (1909) observed 


314 P. SCHUCHERT 


rudimentary bulbs and tentaculae in the same animal. It is thus a variable character. 
Some animals may have four perradial tentacles, although one pair remains small 
(Bigelow, 1909). Goy (1973) observed animals with four equally developed perradial 
tentacles (see below). At least in younger stages, the gonads are connected interradially 
on the manubrium by a thin layer (Bigelow, 1909). 


DISTRIBUTION: Florida and Bahamas (Mayer, 1910); Mediterranean (Goy, 
1973); Torres Strait (Mayer, 1910); Papua New Guinea (Bouillon, 1980); Japan 
(Uchida, 1938a); Pacific Side of Mexico (Bigelow, 1909); Chile (Kramp, 1966). Type 
locality: Tortugas, Florida, USA. 


REMARKS: This species is very rare along the European coasts, being known 
only from four specimens found in the Mediterranean. While the medusa depicted in 
Goy (1973) is evidently A. turrida, other specimens collected by Goy had four equally 
developed perradial tentacles instead of two only. These specimens could be re- 
examined for this study. The gonads of these specimens are smooth and unlike the ones 
in Fig. 48. The gonads are rather large and bulging, enveloping and almost completely 
hiding the rest of the manubrium (see also Codonorchis). It is not clear whether this 
represents only a fixation artefact. New Mediterranean material is needed for a further 
evaluation of the status of this form. 


Amphinema krampi Russell, 1956 Fig. 49 
Amphinema krampi Russell, 1956a: 371, figs 1-2.— Russell, 1958: 81, figs 1-3f.- Kramp, 1959a: 

118, fig. 113. — Kramp, 1961: 93. — Russell, 1970: 245, fig. 8s. 

MATERIAL EXAMINED: Holotype BMNH 1956.1.10.1,47.20°N 07.67°W, west of Brittany, 
0-821m; 21.07.1955; one badly damaged medusa. 

DIAGNOSIS: Medusa oblong; two opposite perradial tentacles, 6-8 small ten- 
taculae; gonads interradial, rugose; manubrium reddish brown and with long 
mesenteries, several fine cellular strands connecting radial canals and exumbrella. 


DESCRIPTION (Russell, 1956a; 1970): Medusa with bell-shaped umbrella, dis- 
tinctly higher than wide, with a small apical process; jelly moderately thick. Four radial 
canals and ring canal moderately broad, with smooth margins; radial canals attached to 
stomach along half of its length, thus forming mesenteries; up to 17 strands of tissue 
running from each radial canal to the umbrella surface. Two opposite perradial 
tentacles with swollen elongated basal bulbs; 6-8 marginal tentaculae with nema- 
tocysts. No ocelli. Manubrium cross-like in section, about two-thirds length of 
umbrella cavity, without peduncle; mouth with four simple perradial lips, not crenu- 
lated. Four cushion-like gonads, one on each interradial wall of stomach, female 
gonads irregularly folded and much corrugated, male ones perhaps smooth. Colour of 
stomach and gonads rich reddish brown, ring canal brownish. Nematocysts: likely two 
sizes of microbasic euryteles. 


DIMENSIONS: Medusa 5-7 mm high, diameter 4 mm. 


DIMENSIONS: See Russell (1958) for more histological details of the tentaculae 
and the tissue strands. 


BIOLOGY: A rare deep water medusa, taken with wire lengths of 800 to 1600 m. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 315 


Fic. 49 


Amphinema krampi Russell, 1956; from Russell (1958). (A) Female medusa, size 7 mm. 
(B) Histological section depicting a cellular strand connecting the radial canal (rc) with the 
exumbrella (ex). 


DISTRIBUTION: Deep waters west of Brittany. Type locality: 47.20°N 07.67°W, 
max. 821 m. 


REMARKS: This species resembles Amphinema rubrum (Kramp, 1957), but lacks 
the characteristic apical chamber. The tissue strands connecting the radial canals and 
the exumbrella have also not been observed in A. rubrum. It is not entirely clear 
whether the male gonads are smooth or also rugose as in the female. In Russell (1956a) 
they are depicted with a smooth surface. 


Amphinema rubrum (Kramp, 1957) Fig. 50 


Merga rubra Kramp, 1957a: 14, pl. 2 fig. 4. 

Amphinema rubra. — Kramp, 1959a: 118, fig. 114. 

Amphinema rubrum. — Kramp, 1961: 94. — Goy, 1973: 982, fig. 5. — Gili et al., 1998: 117. — 
Bouillon et al., 2004: 68, fig. 40D. 


MATERIAL EXAMINED: MNHN No 1737, 43°39.5’N 07° 17.4’E, 7 Aug. and 1 Oct. 1964; 
2 specimens; material of Goy (1973); bell height up to 4.8 mm. 

DIAGNOSIS: Amphinema species with apical process and large apical chamber, 
manubrium red-brown, bell margin with tentaculae, gonads smooth. 


DESCRIPTION (Kramp, 1957a; own data): Umbrella of medusa higher than wide, 
moderately thick wall, with pointed apical projection; broad apical chamber above 
stomach. Two opposite perradial tentacles with large, conical bulbs; no ocelli observed; 
two perradial and four interradial tenon-like tentaculae. Stomach large, round or cruci- 


316 P. SCHUCHERT 


Fic. 50 


Amphinema rubrum (Kramp, 1957a). (A) From Kramp (1959a). (B) Schematic drawing after 
preserved medusae from the Mediterranean, scale bar 1 mm. 


form in section, connected to radial canals by relatively long mesenteries (1/2 of 
manubrium height), mouth likely simple and cruciform. Four smooth gonads com- 
pletely covering adradial-interradial walls of stomach. Stomach deep reddish-brown. 


Dimensions (Kramp, 1957a): Medusa bell up to 7 mm high, 4.5 mm wide. 
BIOLOGY: The medusa occurs in depths below 250 m. 


DISTRIBUTION: Antarctica, Mediterranean. Type locality: 57.60°S 29.908°W, 
500-250 m, North of South Orkney Islands, Antarctica. 


REMARKS: Kramp (1957a) based his description on a preserved, damaged 
specimen that lacked the mouth region. As ocelli are usually rapidly lost in preserved 
material, it is also not known whether they were originally present. The species has 
also been recorded for the Mediterranean by Goy (1973) and Gili et al. (1998). A re- 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 317 


examination of the material of Goy (1973) confirmed her identification. The manu- 
brium is attached below the highest point of the subumbrella, thus creating a trans- 
parent, chamber-like cavity above the upper level of the manubrium (but which is not 
separated from the gastric cavity). 


Amphinema biscayana (Browne, 1907) Fig. 51 


Bimeria biscayana Browne, 1907: 21, fig. 1, pl. 1 figs 4-5. 
Garveia biscayana. — Vervoort, 1985: 277. 
Amphinema biscayana. — Schuchert, 2000: 415, fig. 3. — Schuchert, 2001: 21, fig. 11A-D. 


MATERIAL EXAMINED: Syntypes BMNH 1948.10.1.118, slides and alcohol material; Bay 
of Biscay (Allen’s station XIII); 752 m; August 1906; infertile. - MHNG INVE25967, alcohol 
and slide material; 62.340°N; 16.997°W; Iceland; 2074 m; 27 August 1991; with medusa buds. 
— MHNG INVE36665 (ex CARACOLE 132-10 KGS 15); 55.544°N 15.687°W (Rockall 
Plateau); 865 m; 13 August 2001; infertile. 

DEscRIPTION: Colonies erect, when fully grown stem and branches stout, may 
be covered by silt particles. Very thin stolons forming a dense rooting structure 
(rhizoid) for anchoring in soft substrata. Alternatively, stolons can be creeping on solid 
substrata. Colony highly polysiphonic, branching profusely, distal part monosiphonic. 
Polysiphonic part, composed of a thick-walled axial tube surrounded by thinner, tightly 
packed, parallel auxiliary tubes. Axial tubes of side branches originate from an 
auxiliary tube of main branch and not from axial tube. Only in distal monosiphonic 
parts, the axial tube is branching. Distal monosiphonic outgrowths as thick as axial 
tubes, composed of long main branches with mostly short side branches, all with 
hydranths. Some hydranths on short pedicels also grow out of auxiliary tubes on lower 
polysiphonic parts of colony. Hydranths covered by adhering pseudohydrotheca that 
ends below tentacle whorl, can be infested with fine silt particles. Hydranth cylindrical, 
one whorl of 10-12 filiform tentacles, hypostome conical or dome-shaped. Some 
hydranths bent at base. Gonophores arise singly, far below hydranths in perisarc 
covered zone. Young gonophores pear-shaped, older ones spherical, enclosed in thin 
perisarc membrane. Gonophores develop into medusa. Oldest medusae observed had a 
simple manubrium without appendages, four broad radial canals, two opposite tentacle 
bulbs were strongly developed and large, elongated, conical, presumably without 
ocelli, each large bulb with one long tentacle without desmonemes. Other pair of 
perradial bulbs small or absent. Between larger tentacles (presumably at interradial 
positions) at least two very small tentaculae (totally at least 4), their tips provided with 
desmonemes. Nematocysts of polyp: desmonemes; asymmetric microbasic euryteles. 
Nematocysts of young medusa: like polyp plus an additional egg-shaped heteroneme 
(for figures see Schuchert, 2000). 


DIMENSIONS: Colonies up to 4 cm high, rhizoid length may reach 2 cm; diameter 
of stems up to 3 mm; axial tubes about 0.4 mm, auxiliary tubes 0.1 mm. 


BIoLOGY: A deep water species recorded in depths of 752-2074 m. 


DISTRIBUTION: Bay of Biscay (Browne, 1907); Rockall Plateau (this study); 
south of Iceland (Schuchert, 2000). Type locality: 48.117°N 08.217°W. 


REMARKS: The axial tubes of the side branches are normally produced by a tube 
that grows out of an auxiliary tube of the original branch, and not by a branching axial 


318 P. SCHUCHERT 


FIG. 51 


Amphinema biscayana (Browne, 1907). (A-C) Silhouettes of colony forms, scale bar 1 cm. (D) 
Hydranth with pseudohydrotheca and young medusa bud, scale bar 0.2 mm. (E) Advanced stage 
of medusa bud, same scale as D. (F) Segment of a cross-section of the stem showing central axial 
tube and peripheral auxiliary tubes (not all shown), scale bar 0.2 mm. After preserved material 
from Iceland (A, D-F) and the Rockall Plateau (B-C). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 319 


tube itself. This latter, more common branching pattern, is known from the similar e.g. 
Garveia arborea (Browne, 1907). 

This species cannot yet be allocated to the correct genus, as information on the 
mature medusa is not available. The genus Amphinema used is only a guess, the 
colonies could belong to several other member of the family Pandeidae. Speculating 
on the possible adult medusa of A. biscayana, and considering only pandeid medusae 
known from the North Atlantic having no ocelli and possessing tentaculae, Amphinema 
krampi Russell, 1956a and Merga reesi Russell, 1956b seem to be good candidates. 
Both are deep water medusae first discovered close to the type locality of Amphinema 
biscayana. 


Amphinema bouilloni n. spec. Fig.52 
Codonorchis octaedrus.— Boero et al., 1997: 359, figs 1-3. 
[not Codonorchis octaedrus Haeckel, 1879] 


TYPE MATERIAL: Not examined, according to Bouillon er al. (1997) an adult male medusa 
has been deposited as type specimen at the Institut Royal des Sciences naturelles de Belgique, 
Bruxelles, IG 27838. 

DIAGNOSIS: Amphinema species with ocelli, up to 14 rudimentary bulbs present, 
apical process present, aboral processes of manubrium penetrating into mesogloea; 
gonads interradial on manubrium, horse-shoe shaped, with slight folds. Hydroid 
sessile, medusa buds on stolons, with pedicels. 


DESCRIPTION (Boero er al., 1997): Hydroid colonial, stolons tubular; hydranths 
sessile (without pedicel); hydranth body naked, yellow, white hypostome, single whorl 
of 4-6 filiform tentacles; hydrant contracts longitudinally and not sideways. Medusa 
buds orange, ovoid, arising singly from stolons, on pedicel of variable length. 
Nematocysts: microbasic euryteles and desmonemes. 

Newly released medusa higher than wide, with small apical projection, exum- 
brella with numerous nematocysts; four ribbon-like radial canals and ring canal; with 
four perradial marginal bulbs, each with a red ocellus, one opposite pair of bulbs larger, 
conical, with a short tentacles on each. Manubrium tubular, orange, reaching !/4 of sub- 
umbrellar height, mouth quadratic with four small lips. 

Mature medusa higher than wide, lateral wall relatively straight, with well 
formed apical projection, no gastric peduncle; four radial canals ribbon-like, broad, 
jagged, forming short mesenteries where entering stomach. Margin with 16 bulbs, all 
with abaxial ocelli; one perradial opposite bulb-pair large and tapering into long 
tentacles, other bulbs small, conical, sometimes with a short tentacle rudiment. 
Manubrium relatively small, reaching about 1/, of the height of the subumbrellar 
cavity, with quadratic base, with one big and several small apical processes penetrating 
into apical mesogloea, mouth-region cruciform in section, four simple lips. Gonads on 
interradial wall of manubrium only, separated perradially, large, horse-shoe shaped, 
with some interradial folds. Nematocysts: microbasic euryteles and desmonemes. 


DIMENSIONS (Boero et al., 1997): Polyp 0.25 mm; newly liberated medusa 
0.65 mm wide and 1 mm high; bell height of mature medusa 2.2 mm, apical projection 
0.7 mm. 


TYPE LOCALITY: Mediterranean Sea near Otranto. Apulia. Italy; depth 25 m. 


320 P. SCHUCHERT 


| 
\ 


Fic. 52 


Amphinema bouilloni new species; from Boero et al. (1997), for dimensions see text. 
(A) Hydroid with medusa bud. (B) Newly released medusa. (C) Mature medusa. 


REMARKS: This species is based on material originally attributed to Codonorchis 
octaedrus by Boero et al. (1997). The arguments why I disagree with this identification 
are given in the remarks under the genus Codonorchis. The species is therefore 
described as new. It resembles closely Amphinema australis (Mayer, 1900b) and 
Amphinema tsingtauensis Kao et al., 1958, two other Amphinema species with ocelli. 
Amphinema turrida also has ocelli, but its unique gonads make it quite distinct. 
Amphinema australis differs from A. bouilloni in the following details (Bigelow, 1909; 
Mayer, 1910): no aboral processes of manubrium, only 8 marginal bulbs and not 16, 
the bell-shape is relatively wide and not so high, the gonads are smooth and bulging. 
Amphinema tsingtauensis from China appears even more similar, but differs in the 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 321 


following traits (Kao et al., 1958): no aboral processes of manubrium, the size is 2-3 
times larger, 22 bulbs instead of 16, gonads in marked oblique folds. Admittedly, these 
differences relate to somewhat variable characters that can be considered as not so suit- 
able to distinguish species. However, as long as the life cycle of the A. australis and A. 
isingtauensis remain unknown, a more detailed comparison with A. bouilloni is 
impossible and it is therefore preferable to have it named as a new species. 


Genus Leuckartiara Hartlaub, 1914 

TYPE SPECIES: Geryonia octona Fleming, 1823 [designation by Kramp, 1959a]. 

SYNONYM: Dinema Van Beneden, 1867 [type species Dinema slabberi Van Beneden, 
1867, = L. octona]. 

REFERENCES: Hartlaub (1914: 282), Kramp (1926: 76), Ranson (1936: 70), Rees (1938: 
11); Kramp (1959a: 120), Russell (1953: 187), Schuchert (1996: 68), Pages et al. (1992: tab. 1), 
Bouillon & Boero (2000: 107). 

DIAGNOSIS: Hydroids colonial, arising from creeping stolons; hydrocauli 
covered by perisarc, not or only sparingly branched, stems monosiphonic. Perisarc 
extends onto hydranth body as a more or less gelatinous pseudohydrotheca which does 
not envelop the tentacles. Hydranths with a conical hypostome and one whorl of 
filiform tentacles. Gonophores develop on cauli or stolons, liberated as medusae. 
Medusa mostly with an apical projection of variable size. Manubrium voluminous, 
connected to radial canals by mesenteries. Mouth with extensively folded and crenu- 
lated margin. Gonads on interradial walls of manubrium, bipartite but with broad 
connection in upper half, with with various degrees of folding, no or only few inter- 
radial pits. Radial canal very broad, often jagged. With many tentacles arising from 
elongated bulbs, laterally compressed; small bulbs or growth arrested tentacles often 
present. With or without ocelli. 


REMARKS: See also under Neoturris. Pages et al. (1992) provided a tabular 
review on all known Leuckartiara species. A key to all species is provided by Xu & 
Huang (2004). 

Even for a restricted region like the north-eastern Atlantic, juvenile and subadult 
medusae of the genera Leuckartiara and Neoturris can be very difficult to identify. 
There is also quite a lot of variation in fully mature specimens that renders identi- 
fications not an easy task. 


KEY TO THE LEUCKARTIARA MEDUSA TREATED HERE: 


la more thantoursullysdevelopeditentacles ar... Re ee 2 
1b only four perradial tentacles fully formed, others small; no ocelli ......... L. brownei 
2a MESENtEHES ALON Sxentine:StOMACh EME CE RL LIN RER L. grimaldi 
2b MeEsenterics Shorten thansSlOmac Mes ne Wa NAS O SIINO 3 
3a mature with 1-3 rudimentary bulbs between fully formed tentacles, with abaxial 

SPULS re ie elle lee cua geeks SIRIO DT I L. octona 
3b few rudimentary bulbs but majority of tentacles not fully grown, no abaxial 

SDULS Le TS nn nee rc ee be ere ORS EAC IT L. nobilis 
Leuckartiara octona (Fleming, 1823) Fig. 53 


Geryonia octona Fleming, 1823: 299. 

Oceania pileata. — Ehrenberg, 1837: 190, pl. 8 figs 2-4. 
[not Neoturris pileata (Forsskäl, 1775)] 

? Turris neglecta Lesson, 1843: 284. 


P. SCHUCHERT 


322 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 323 


Oceania episcopalis Forbes, 1848: 27, pl. 2 fig. 1. 

Oceania turrita Forbes, 1848: 28, pl. 2 fig 2. i 

Eudendrium sessile Wright, 1857: 60, pl. 3 figs 16-17. 

Eudendrium pusillum Wright, 1857: 84, pl. 2 figs 8-9. 

Atractylis repens Wright, 1858: 450, pl. 22 figs 4-5. — Wright, 1859a: 108, pl. 1 figs 4-5. 

Atractylis palliata Wright, 1861: 129, pl. 4 fig. 6. 

Perigonimus minutus Allman, 1863: 11. 

Perigonimus vestitus Allman, 1864b: 57. — Allman, 1872: 326, pl. 11 figs 1-3. — Rees, 1956a: 
345, synonym. 

Tiara smaragdina Haeckel, 1864: 336. — Hartlaub, 1914: 294, synonym. 

Dinema slabberi Van Beneden, 1867: 130, pls 9-10. 

? Oceania gaedii Van Beneden, 1867: 95. 

Perigonimus repens. — Hincks, 1868: 90, pl. 16 fig. 2. 

Oceania coronata Allman, 1871: 33, fig. 8. 

Tiara octona. — Haeckel, 1879: 57. 

Perigonimus Jonesii Osborn & Hargitt, 1894: 27, fig. 1-12. — Rees, 1956a: 340, synonym. 

? Perigonimus gelatinosus Duerden, 1895: 327, pl. 14 figs 2-3. — Rees, 1956a: 340, possible 
synonym. 

Perigonimus pugetensis Heath, 1910: 74, figs 1-2. — Rees, 1956a: 343, synonym. 

? Perigonimus napolitanus Hargitt, 1904: 571, pl. 22 fig. 25. 

Tiara papua. — Maas, 1909: 9 pl. 1 fig. 3. 

[not Neoturris papua (Lesson, 1843)] 

Perigonimus vestitus f. radicans Vanhöffen, 1910: 286, fig. 11a-c. 

Turris vesicaria. — Mayer, 1910: 126, pl. 12 figs 2-3, pl. 13 fig. 7. 

[not Catablema vesicarium (A. Agassiz, 1862)] 

Perigonimus repens.— Broch, 1911: 14, fig.11. — Broch, 1916: 52, fig. P.— Vervoort, 1946: 141, 
figs 54-55. 

Leuckartiara octona. — Hartlaub, 1914: 285, figs 238-214, figs 244-253. — Kramp, 1926: 76, pl. 
2 figs 5-7, text fig. 35. — Uchida, 1927: 211, fig. 37. — Ranson, 1936: 75. — Rees, 1938: 
11, fig. 3-5. — Russell, 1938b: 153, figs 41-44. — Russell, 1953: 188, pl. 11 figs 5-6, pl. 
12. fig. 3, pl. 30-31, text figs 91-96. — Kramp, 1959a: 121, fig. 119. — Kramp, 1961: 105. 
— Kramp, 1968: 47, fig. 121. — Millard, 1975: 123, figs 41a-d. — Arai & Brinckmann- 
Voss, 1980: fig. 29. — Hirohito, 1988: 106, fig. 38f-g. — Pages et al., 1992: 10, 12, fig. 11. 
— Ramil & Vervoort, 1992: 21. — Migotto, 1996: 16, fig. 3e-g. — Schuchert, 1996: 68, 
fig. 40a-b. — Bouillon et al., 2004: 70, fig. 41D-F. — Pages et al., 2006: Fig. 7F. 

Leuckartiara octona var. minor Ling, 1937: 353: fig. 2. 


MATERIAL EXAMINED: Mediterranean, Ligurian Sea, Bay of Villefranche-sur-Mer, 70 m; 
4 April 2005; 1 fertile medusa, examined alive, used for DNA extraction, 16S sequence 
AM411421. - MHNG INVE49100, hydroid culture obtained from Prof. G. Plickert, Cologne, 
originally isolated from hermit crab shells from the North Sea, grew well and produced many 
medusa, these cultivated to a size of 3-4 mm, 6-8 tentacles plus 8 bulb rudiments, with abaxial 
spurs, some hydroids preserved and deposited in MHNG; 16S sequence AM411422. - MHNG 
INVE35728, large colony on Aporrhais pespelecani Italy, Naples, Palazzo Donn’Anna; 30 


Fic. 53 


Leuckartiara octona (Fleming, 1823). (A) Hydroid colony with medusa buds; for sizes see text: 
modified from Hincks (1868). (B) Optical section of hydrocaulus with branching point, note 
detritus infested outer perisarc layer (arrow), scale bar 0.1 mm, after preserved material from 
Helgoland. (C) Advance medusa bud, scale bar 0.1 mm; after preserved material from 
Helgoland. (D) Newly liberated medusa, bell height 1.2 mm; modified after Russell (1953). (E) 
Young medusa with beginning gonad formation, bell height 4 mm; modified from Russell 
(1953). (F) Mature medusa after preserved material from Denmark, scale bar 3 mm. (G) 
Magnified section of bell margin with the bases of a pair of normal tentacles flanking three rudi- 
mentary bulbs, the middle one is club-shaped. (H) Lateral view of a tentacle base showing 
abaxial spur (arrow), after preserved material. 


324 P. SCHUCHERT 


March 1911. - ZMUC; Denmark, Frederikshavn, inner side of harbour; 0 m; 27 July 1981; 
mature medusae. - BMNH 1982.11.30.5; Great Britain, Eddystone; 4 June 1925; coll. F. S. 
Russell, numerous mature medusae. - BMNH, slide with no registration number, Loch Striven, 
Clyde, 55 m, 19 September 1931, several polyps with medusa buds. - BELUM Md643, Ireland, 
Donegal, St Johns point, Portnagh Rock; 2 mature medusae. — ZSM, slides 20000635 and 
20000636, as Perigonimus gelatinosus, Duerden, on Pagurus, loc. Helgoland, Stechow 
collection, mentioned in Stechow (1919). 


DIAGNOSIS: Leuckartiara medusa with 8-23 fully developed marginal tentacles, 
one to three rudimentary marginal bulbs between adjacent pairs of tentacles; tentacle 
bulbs with distinct abaxial spur bearing an ocellus; without exumbrellar nematocyst 
tracks, with apical process. Mature 10-15 mm in height. 


DESCRIPTION (Kramp, 1926; Rees, 1938; Russell, 1953; own observations): 
Hydroids arising from creeping, branching to reticulate hydrorhiza. Shoots either 
unbranched or sparingly branched; sister-branches not adnate for some distance; 
hydranth pedicels can be relatively long, but variable and depending on environment, 
sometimes almost subsessile. Perisarc often two-layered with a particle infested outer 
gelatinous layer and a solid inner layer, solid perisarc smooth or variable corrugated, 
transparent or horn coloured with a distinct membranous dilatation reaching to base of 
tentacles (pseudohydrotheca). Hydranths spindle-shaped, with a single whorl of 6-12 
tentacles. Medusa buds borne on short stalks arising from stolons or the hydranth 
pedicels. Advanced medusa buds ovoid, covered by thin perisarc membrane, medusa 
structure well visible, only two tentacles developed. Nematocysts: desmonemes and 
microbasic euryteles (Bouillon, 1985). 

Newly liberated with or without small apical process, with scattered exumbrel- 
lar nematocysts. Stomach cylindrical, about one-third the height of subumbrella cavity 
in length, mouth simple, without lips. Two opposite perradial tentacles and two 
opposite perradial rudimentary marginal bulbs, without ocelli; the marginal tentacle 
bulbs, rudimentary marginal bulbs and stomach are pale straw to ochreish in colour. 
The first ocelli appear on the second perradial pair of tentacles when the bell reaches a 
size of about 2 mm. At the eight tentacle stage, with a height of about 4 mm, the mouth 
is already considerably crenulated and the gonads begin to form on the adradial 
surfaces of the stomach. Sixteen tentacles are reached at about 7-12 mm bell height. 

Mature medusa umbrella bell-shaped, higher than wide, with conical or sphe- 
rical solid apical process varying much in size; jelly thin; velum narrow. Four radial 
canals broad, with smooth or jagged outlines; ring canal broad, with smooth outlines; 
radial canals joined to stomach by long mesenteries. Eight to 23 (usually around 16) 
marginal tentacles, hollow, smooth, with large, laterally compressed, elongated conical 
basal bulbs, clasping umbrella margin and forming pronounced abaxial spurs on 
exumbrellar surface, nematocysts of tentacle concentrated in adaxial clasps. Usually 
one to three rudimentary marginal bulbs between adjacent tentacles (temporarily 
growth arrested), either simply hemispherical buttons or pine-cone shaped appendage, 
not tapering into short tentacle. One small abaxial ocellus on each tentacular and 
rudimentary marginal bulb. Stomach flask shaped, not extending beyond umbrella 
margin in full extension, with broad base; mouth with four crenulated lips. Gonad 
situated interradially on folds of stomach wall, in two tiers, joined by transverse bridge, 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 325 


with several folds directed outwards to each perradius. Nematocysts: microbasic 
euryteles and microbasic mastigophores (Russell, 1938b). Colour of marginal tentacles 
and stomach a deep flesh-pink to crimson or yellowish; ocelli crimson. 


DIMENSIONS: Mature medusa 5-20 mm, usually around 10 mm; medusa when 
first liberated from its hydroid about 1.1 mm in height. Nematocysts of medusa 
(Russell, 1938b): microbasic euryteles (8-10.5)x(3-4.5)um; microbasic mastigophores 
(7)x(3.5-4)um. Hydroid dimensions (Rees, 1938): stolons diameter 0.05-0.07 mm; 
total height of stems 1.5-3.8 mm; hydrocaulus diameter 0.08-0.12 mm; hydranths up to 
0.5-0.7 mm, diameter of hydranths 0.25-0.3 mm; length of medusa buds 0.6 mm, 
diameter of medusa buds 0.45-0.5 mm, stalk of medusa bud 0.1-0.3 mm. 


DISTRIBUTION: A neritic, circumglobal species occurring in subtropical to 
temperate seas. The medusa is common and widely distributed on the north-western 
Furopean coasts (see summaries in Hartlaub, 1914; Kramp, 1926; Russell, 1953; 
Kramp 1959a; Kramp, 1965): coasts of the British Isles; west coast of Norway to 
Lofoten; Skagerrak and Kattegat but not the Baltic Sea; south of Iceland; Rockall; 
France and Iberian Peninsula; western Mediterranean; east coast of North America 
from Newport to Labrador; western Africa (Kramp (1959a); South Atlantic (Pages er 
al.. 1992); Red Sea (Schmidt, 1973a); Indian- and Pacific Ocean (Hartlaub, 1914; 
Kramp, 1965; Schuchert, 1996). Arai & Brinckmann-Voss (1980) do not include the 
medusa in the fauna of British Columbia. The polyp has also been widely reported 
(usually as Perigonimus repens) along coast of the north-eastern Atlantic and 
Mediterranean (Rees, 1938); also South Africa (Millard, 1975), Brazil (Migotto, 1996), 
and the Pacific and Indian Ocean (Rees, 1938). 


ADDITIONAL DATA: For more structural and developmental details see Kramp 
(1926), Rees (1938), and Russell (1953). The shape and size of the apical process is 
variable. The hydroid morphology and dimensions are also variable and depend on 
food and substrate (Rees, 1938). The ultrastructure of the ocelli is described by Singla 
(1974) (species identity perhaps not clear, see Arai & Brinckmann-Voss, 1980). 


BIOLOGY: The hydroid is nearly always living in association with other animals, 
it occurs in association with many species of crustacean and molluscs, in particular also 
on the parapodia of the polychaete Aphrodite (Leloup, 1934; Latham, 1963). Also 
known to occur on fish (Rees, 1938). Depth distribution 10-400 m (Christiansen, 
1972). In Norway, Christiansen (1972) observed medusa budding from May to 
October. Kramp (1930) says that medusa liberation stops in August or somewhat later. 
Robson (1914) records the hydroid as liberating medusae from September to February 
on the Northumberland coast. 

As summarized by Kramp (1926), the medusa is found at the British coasts 
from April or May (young individuals) to October or November. In Danish waters, the 
young individuals appear first in May or June; during autumn L. octona is one of the 
most abundant medusa of the region; in certain years a number of individuals may 
surpass the winter and be observed in the plankton as late as March or April the next 
year. At the coast of Norway, the medusa appears in June and occurs at least until 
November. In the Mediterranean, the medusa is less frequent than in the Atlantic, it has 


326 P. SCHUCHERT 


been observed from February to November (Goy, 1973; Brinckmann-Voss, 1987). The 
depth range of the medusa is about 0-300 m (Kramp, 1965), occasionally also caught 
in deeper waters. 

The medusa makes daily vertical migrations, it is present at the surface only 
during night-time (Buecher & Gibbons, 2003). Feeding habits have been described by 
(Lebour, 1922, 1923). The medusa is highly voracious, eating almost everything given 
to them. 

Vannucci (1960) observed that a female medusa was able to spawn eggs almost 
daily for a period of two months. 


REMARKS: Leuckartiara octona is a common and well known species, but it has 
a complicated taxonomic history that has been worked out by several authors (e. g. 
Hartlaub, 1914; Rees, 1938; Russell, 1953; Rees, 1956a; Kramp, 1961). There is thus 
no need to discuss it here again. The list given above may be not complete as there are 
also several doubtful cases. The species has repeatedly been confounded with other 
similar species like Leuckartiara nobilis, Catablema vesicarium, and Neoturris 
pileata. Only after the thorough revision of Hartlaub (1914), the taxonomy became 
more or less stable. However, as indicated in Arai & Brinckmann-Voss (1980: 56), the 
situation might be more complicated. 

Leuckartiara octona var. minor Ling, 1937 is about 5 mm high and has only 
eight tentacles. It falls well within the range of variation seen in other populations and 
is therefore here considered a synonym. 


Leuckartiara nobilis Hartlaub, 1914 Figs. 54-55 


Leuckartiara nobilis Hartlaub, 1914: 308, figs 257-260. — Kramp, 1926: 83, fig. 36a-e, pl. 2 fig. 
9. — Russell, 1953: 195, pl. 12 fig. 4, text-fig. 97. — Kramp, 1959a: 120, fig. 120. — 
Kramp, 1961: 104. — Kramp, 1968: 47, fig. 123. — Arai & Brinckmann-Voss, 1980: 54, 
fig. 28. — Pages et al., 1992: 12. 


MATERIAL EXAMINED: BMNH 1954.11.13.85; Ireland, Valentia; several medusae 
collected 1898 [by E. Browne ?], could be paratype material. - BMNH 1954.11.13.b3; France, 
Villefranche; 13 March 1913; labelled as ? paratype, leg. & det. C. Hartlaub (comment: material 
is not mentioned by Hartlaub (1914), paratype status unclear. — Scotland, Firth of Lorn, 
Dunstaffnage Bay; 0 m; 7 May 2004; one medusa, identity not entirely clear (see Fig. 55); 14 
mm high, 8 tentacles of varying size, 8 tentacles and 8 rudiments; tentacles without abaxial 
spurs, with ocelli, gonad and manubrium not fully developed, apical process 1/2 of total height, 
16S sequence AM183135. - BMNH 1952.2.20.71; France, Villefranche, Cap Ferrat; 0 m; 15 
April 1949; ca. 16 specimens; collected A. K. Totton. —- ZMUC; 57.050°N 11.333°W, eastern 
Atlantic; 65 m wire; 28 May 1908; Thor Station 12(8), 1 medusa, det. Kramp. - ZMUC Thor 
station 181; 61.567°N 18.750°W, eastern Atlantic; 20 m wire; 10 July 1904; 1 medusa. 


DraGNOSIS: Up to 40 relatively widely spaced tentacles, of these only up to 
8-23 fully developed, others small, in variable stages of development, but with thin, 


pointed ends; only few rudimentary bulbs; bulbs without distinct abaxial spurs, with 
ocelli. 


DESCRIPTION (after Kramp, 1926; Russell, 1953; own data): Umbrella bell- 
shaped, higher than wide, with pointed or rounded solid apical process of variable size; 
jelly thin. Velum narrow. Four radial canals very broad, with irregular jagged outlines; 
ring canals broad with smooth outlines; radial canals joined to stomach by mesenteries. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 327 


> 
3 


Fic. 54 


Leuckartiara nobilis Hartlaub, 1914; after preserved material from the Mediterranean. (A) 
Mature medusa, about 23 mm high. (B) Perradial tentacle in side view, note absence of abaxial 
(ab) spur on bulb and the nematocyst clasps on the abaxial side (ab). (C) Part of bell margin with 
characteristic tentacle arrangement: two fully formed tentacles flank three smaller, not fully 
developed tentacles. 


Up to forty or more marginal tentacles, hollow, smooth, with elongated, conical, 
laterally compressed basal bulbs which clasp the umbrella margin but do not form 
well-developed abaxial spurs; marginal tentacles fairly widely spaced; only about 8-23 
tentacles are fully developed, the intercalated remaining tentacles are much smaller and 
less developed, ending in a pointed tip (unlike club or pine-cone); with only a few 
scattered rudimentary bulbs. One small ocellus on each tentacle bulb, at abaxial side 
close to exumbrella. Tentacles have nematocysts concentrated on adaxial side, forming 
clasps, also gastrodermis has an irregular outline. Stomach flask-shaped, constricted 
near mouth, not extending beyond umbrella margin in full extension, with broad base; 
mouth with much folded lips. Gonads interradial on folds of stomach, bipartite with 
transverse connection in upper half, adradial parts with many folds directed outwards 
to each perradius, there are regularly a few pits in the interradial region of the gonads. 
Colour: stomach, i.e. gonad, orange red; tentacle bases bright yellowish-brown or 
white. 


328 P. SCHUCHERT 


Fic. 55 


Leuckartiara nobilis Hartlaub, 1914, living medusa from Dunstaffnage, Scotland, about 14 mm 
high. This plate illustrates the difficulty to identify subadult Leuckartiara species. The medusa 
has 8 tentacles and adradial rudiments, the typical gonad-folds are clearly visible. It was iden- 
tified as L. nobilis and not L. octona because there are no abaxial tentacle spurs and at this size 
L. octona should have more tentacles. However, there remains some ambiguity. The shape and 
size of the apical process is highly variably in both species. 


DIMENSIONS: Total height of medusa when fully grown 20-27 mm, apical 
process up to 1/5 of total height, in younger specimens up to 1/2. 


VARIATION: Apparently fully grown medusae from the Atlantic can have 12-23 
maximally developed tentacles, while Mediterranean medusae have only 8-12. 


DISTRIBUTION: In the North Atlantic found usually rather far off the coast. 
Recorded from the western Mediterranean, SW of Ireland, Atlantic Ocean west of 
Scotland, Rockall, at Iceland, New Foundland, one exceptional finding at Denmark 
(Hartlaub, 1914; Kramp, 1926; Goy, 1973); also in the Pacific near Japan and British 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 329 


Columbia (Kramp, 1965; Arai & Brinckmann-Voss 1980). Type locality: Not 
designated, Hartlaub (1914) had material from the Ligurian Sea (Monaco, 
Villefranche-sur-Mer) and from western Ireland (Valencia, now spelled Valentia). 


BIOLOGY: A species that seems to prefer oceanic regions. There are only few 
records, these indicate an occurrence during the summer months in the Atlantic and in 
spring for the Mediterranean (Goy, 1973; Brinckmann-Voss, 1987). Fraser (1969) 
describes the feeding behaviour. 


REMARKS: This is a relatively little known species of which the polyp stage 
remains to be discovered. Some Atlantic specimens identified by Hartlaub (1914) and 
Kramp (1926) as L. breviconis appear only gradually different from L. nobilis (see 
discussion under N. breviconis). 

The sample from Scotland (Fig. 55) was rather young and it was not possible to 
identify it with the desirable reliability. Its 16S sequence was quite different (7.9% 
bases) from an unambiguously identified L. octona (AM411421). 


Leuckartiara grimaldii Ranson, 1936 Fig. 56 
Leuckartiara grimaldii Ranson, 1936: 78, pl. 1 figs 7-8. — Kramp, 1961: 104. — Kramp, 1959a: 
120. 


DIAGNOSIS: 16 mm high, 12 mm wide; bell-shaped, thin walls, no apical pro- 
jection. Stomach very large, smooth, filling 2/3 or more of subumbrella, mesenteries 
along entire length of stomach; gonads not observed; mouth folded. Radial canals 
short, broad, slightly jagged. 24 tentacles and 32 small bubs. Presence of ocelli 
unknown. 


DISTRIBUTION: Known from type locality only: 37.467°N 25.117°W (near 
Azores), 0-1000 m. 


Fic. 56 
Leuckartiara grimaldii Ranson, 1936, redrawn from Ranson (1936). 


330 P. SCHUCHERT 


REMARKS: This species is based on a single, damaged specimen. Gonads were 
not observed by Ranson (1936) and the ocelli could not be observed as the material had 
been conserved for more than 30 years when it was described. The inclusion into the 
genus Leuckartiara was based solely on the tentacles which resemble those of L. no- 
bilis or N. breviconis as described by Hartlaub (1914). However, other genera are also 
possible, e. g. Pandea or Merga. Ranson (1936) evokes the possibility that the 
specimen was a senescent animal with spent gametes. The species remains 
insufficiently known. 


Leuckartiara abyssi (G.O. Sars, 1874) Fig. 57 


Perigonimus abyssi G.O. Sars, 1874: 126, pl. 5 figs 27-30. — Broch, 1916: 53, fig. Q. — Rees, 
1956a: 338. — Naumov, 1969: 202, fig. 70. 

Leuckartiara abyssi. — Rees, 1938: 19, fig. 6a-d. — Rees, 1956b: 114. — Edwards, 1965: 446. 
DIAGNOSIS: Stolonal hydroids on bivalves and scaphopods, medusa buds on 

hydrocauli and stolons, with pseudohydrotheca, newly liberated medusa with four 

tentacles. 


DESCRIPTION (Rees, 1938): Hydroid epizoic on bivalves and scaphopods, 
colonies stolonal hence hydrocauli unbranched. Stolons form an open network, colour 
light brown. Perisarc of stems with adhering mud particles; perisarc often irregularly 
wrinkled throughout, but never ringed at the hydrocaulus base, becoming dilated 
around the base of the hydranth to form a pseudohydrotheca, reaching almost to the 
base of the tentacles where it thins out and disappears. Hydranth fusiform to flask- 
shaped, with a well developed conical hypostome and one whorl of six to eight filiform 
tentacles, the long ones held almost vertical and the shorter ones at right angles to the 
hydrocaulus. The limits of the hydranth and the hydrocaulus are not clearly marked, 
the transition being gradual. Gonophores develop on short stalks from the stolons and 
the hydrocauli. 

Newly liberated medusa deeply bell-shaped, no apical process, jelly fairly thin 
and of uniform thickness, scattered nematocysts on exumbrella. Velum present. Four 
radial canals and ring canal fairly broad. With four perradial tentacles with elongated 
hollow conical bulbs; of these two opposite tentacles were better developed than the 
other two; without ocelli. Manubrium short, quadrangular in section, reaching about 
1/3 the height of the umbrella cavity, mouth without lips. Colour of bulbs pale 
yellowish green. 


DIMENSIONS (Rees, 1938; Rees, 1956b): Total height of stems up to 1.1 mm, 
perisarc covered part 0.7 mm, hydranth body 0.35 mm, diameter of hydranth 
0.175 mm, stem diameter 0.05 mm, stolon diameter 0.05-0.07 mm, tentacle length up 
to 1.2 mm, medusa bud 0.45-0.50 mm long, stalk of medusa bud 0.05 mm, newly 
liberated medusa 1.15 mm high and 1.05 mm wide. 


DISTRIBUTION: A northern boreal to Arctic species, but, because of the likely 
confusion with other species, most records outside Norway should be regarded with 
reservation. Recorded from Norway (Sars, 1874; Bonnevie, 1901; Rees, 1938; 
Christiansen, 1972), western Sweden (Jäderholm, 1909); Greenland (Kramp, 1911, 
1914, 1943); Bear Island (Bonnevie, 1899), Spitsbergen (Broch, 1910); Barents Sea 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 33] 


Fic. 57 
Leuckartiara abyssi (G.O. Sars, 1874), from Rees 1938, for dimensions see text. 


(Spassky, 1929). A record from western Ireland by Ritchie (1913) has been regarded as 
doubtful by Edwards (1965). Type locality: Norway, Hvitingsg (= Kvitsgya, island 
near Stavanger), 146-365 m, on Dentalium dentale (Linnaeus), designation by Rees 
(1956b). 


BIOLOGY: The hydroid grows on scaphopods (Dentalium dentale) and bivalve 
shells of the genus Nucula and Nuculana occurring normally in depths of 100-600 m 
(range 20-1600 m). Medusa buds were observed from July to September (Rees, 1938; 
Christiansen, 1972). 


REMARKS: All morphological and biological traits of L. abyssi from Norway 
appear identical to the hydroid of Neoturris pileata described by Edwards (1965) based 
on material from Scotland. Although it is likely that also the Norwegian hydroid 
belongs to this species — the medusa of N. pileata also occurs there — Edwards correctly 
refrained from synonymizing the two species because more life cycle information for 
the Norwegian hydroid is needed. Because many pandeid hydroids are very similar, L. 
abyssi could also correspond to other medusae known from the region, e. g. Neoturris 
breviconis or L. nobilis. 


Leuckartiara brownei Larson & Harbison, 1990 Fig. 58 


Perigonimus sp. Browne, 1910: 16. 
Leuckartiara brownei Larson & Harbison, 1990: 20, fig. 1. — Pagés et al., 1992: 12. — Bouillon 
et al., 2000: 89, fig. 2. — Bouillon et al., 2004: 70, fig. 41B. 
DiAGnosıs: Medusa up to 10 mm, four normal perradial tentacles and up to 28 
much smaller tentacles; no ocelli; gonad on two longitudinal interradial folds, no trans- 
verse connection. 


332 P. SCHUCHERT 


Fic. 58 


Leuckartiara brownei Larson & Harbison, 1990; modified from original publication, bell height 
10 mm. 


DESCRIPTION: Umbrella with a small pointed apical projection of variable 
height; mesogloea moderately thick, velum narrow. Radial canals four, fairly broad, 
mostly smooth outlines. Only the four perradial tentacles are fully formed, tapering, 
not laterally compressed, coiled when contracted; additionally up to 28 short rudimen- 
tary tentacles (= juvenile tentacles), number variable, developing in succession; all 
tentacles without ocelli and abaxial spurs. Manubrium relatively large, more than half 
of the bell cavity height; mesenteries well developed; lips somewhat folded. Gonads 
occupy the interradial areas of the manubrium, each gonad with a pair of longitudinal 
folds adjacent to the interradii forming a continuous interradial groove, additionally a 
few isolated folds in the adradii mostly oriented perradially. Coloration in life: gonads 
orange-brown, tentacles pink-orange. 

The youngest specimens show that the medusa is probably released with two 
tentacles only. 


DIMENSIONS: Umbrella up to 10 mm high and 9 mm wide. 


BIOLOGY: In the Antarctic occurring from the surface to depths of several hun- 
dred meters (Bouillon er al., 2000), this always at sub-zero temperatures (Larson & 
Harbison, 1990). 


DISTRIBUTION: Antarctica, Mediterranean. Type locality: McMurdo Sound, Ross 
Sea, Antarctica. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 333 


REMARKS: It is somewhat surprising to find a cold water medusa in the relatively 
warm waters of the Mediterranean. The Antarctic Leuckartiara brownei medusae occur 
in waters of sub-zero temperatures, while the Mediterranean waters remain always 
warmer than 12°C at all depths (Klein & Roether, 2001). 


Genus Neoturris Hartlaub, 1914 


TYPE SPECIES: Medusa pileata Forsskal, 1775, designation by Kramp (1959a). 

SYNONYMS: Turris Lesson, 1843 for Turris papua Lesson, 1843: 283, name preoccupied. 
— Tiara Lesson, 1843, for Tiaria papalis Lesson, 1843 = Neoturris pileata, name preoccupied. 

DiaGnosis: Hydroids colonial, arising from creeping stolons; hydrocauli 
covered by perisarc, not or only rarely sparingly branched, stems monosiphonic. 
Perisarc extends onto hydranth body as a more or less gelatinous pseudohydrotheca 
which does not envelop the tentacles. Hydranths with a conical hypostome and one 
whorl of filiform tentacles. Gonophores develop on cauli or stolons, enclosed in thin 
perisarc membrane. Gonophores liberated as free medusae. 

Medusa mostly with an apical projection of variable size. Manubrium volu- 
minous, connected to radial canals by mesenteries. Mouth with extensively folded 
margin. Gonads on interradial walls of manubrium, each quadrant with two adradial 
longitudinal rows of transverse folds, interradial region depressed, with numerous 
gonadal pits. Radial canal very broad, often jagged. With many tentacles arising from 
elongated bulbs, laterally compressed, without rudimentary tentacles or marginal 
warts. Without ocelli. 


REMARKS: Following the opinion of Arai & Brinckmann-Voss (1980), this 
genus also includes N. breviconis (Murbach & Shearer, 1902). 

The polyp stage of Neoturris pileata, the only one known, is indistinguishable 
from those found in the genus Leuckartiara. The main differences to Leuckartiara 
medusae are the adradial rows of gonadal folds, typically directed towards interradial, 
flanking a region with pits. Also the absence of ocelli is quite characteristic, although 
Leuckartiara brownei also lacks them. 


KEY TO THE NEOTURRIS MEDUSAE TREATED HERE: 


la eight adradial rows of gonads folds, folds directed towards interradial, many 
Ponadalipues; (S20) TORE se ee ee N. pileata 

lb gonad folds arranged like horseshoe, folds directed towards perradial and aboral, 
relatively few interradial gonadal pits (<20) ....................... N. breviconis 

Neoturris pileata (Forsskal, 1775) Fig. 59-60 


Medusa pileata Forsskal, 1775: 110. — Forsskal, 1776: pl. 33, fig. D. 

Oceania Lesueurii Péron & Lesueur, 1810: 345. — Goy, 1995: 244, plate 
Carybdea pisifera Oken, 1815: 125. 

Oceania pileus de Blainville, 1830: 258. 

Oceania ampullacea Sars, 1835: 22, pl. 6 fig. 8. — Haeckel, 1879: 58, synonym. 
Tiaria papalis Lesson, 1843: 287. — Haeckel, 1879: 58, synonym. 

Turris digitale Forbes, 1846: 286. — Hartlaub, 1914: 324, synonym. 

Turris digitalis. — Forbes, 1848: 21, pl. 3 fig. 1. — Haeckel, 1879: 61, pl. 4 figs 2-3. 
Oceania episcopalis Forbes, 1848: 27, pl. 2 fig. 1. — Haeckel, 1879: 58, synonym. 
Oceania cocciniea Leuckart, 1856: 20, pl. 2 fig. 3— Haeckel, 1879: 58, synonym. 
Oceania constricta Patterson, 1859: 279, figs. 


334 P. SCHUCHERT 


FIG. 59 


Neoturris pileata (Forsskäl, 1775); from Edwards (1965). (A) Typical distribution of the hydroid 
colony on a bivalve. (B) Part of hydroid colony bearing medusa buds), for dimensions see text. 
(C) Newly liberated medusa, height 1.2 mm. (D) Juvenile medusa (2.7 mm) with beginning 
formation of gonadal ridges. (E) Young medusa of about 5 mm height, a small apical process is 
present and more gonad-folds and pits have formed. 


Tiara pileata. — Haeckel, 1879: 58, pl. 3 figs 6-8. 

Turris coeca Hartlaub, 1892: 19, fig. 1.— Hartlaub, 1914: 329, synonym. 
in part Turris pileata. — Mayer, 1910: 123, pl. 12 fig. 4, pl. 13 fig. 6. 
Tiara pileata. — Le Danois, 1914a: 17, fig. 4. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 335 


Fic. 60 


Neoturris pileata (Forsskäl, 1775). (A) Mature medusa from Villefranche-sur-Mer, preserved 
material, scale bar 5 mm. (B) Mature medusa from off Mauritania, preserved material, scale bar 
10 mm. This form lacks an apical process and differs in other details from the typical form shown 
in A. It could belong to a different, unnamed species. (C) Subadult medusa from Scotland, after 
photograph of living specimen, approximately same scale as B. 


Neoturris pileata. — Hartlaub, 1914: 326, figs 270, 273, 274-281. — Kramp, 1926: 92, fig. 37, pl. 
2 figs 13-14, chart XVIII. — Russell, 1953: 203, figs 104-106, pl. 12 fig. 1. — Kramp, 
1959a: 122, fig. 124. — Kramp, 1961: 109. — Edwards, 1965: 461, figs 1-4. — Kramp, 
1968: 50, fig. 131. — Russell 1970: 247, figs 10s-13s— Bouillon et al., 2004: 73, fig. 
42C-E. 


336 P. SCHUCHERT 


MATERIAL EXAMINED: MHNG INVE35522; Mediterranean, Villefranche-sur-Mer; 6 ma- 
ture medusae, 2-3 cm, leg. and det. C. Hartlaub 1911, material mentioned in Hartlaub (1914). — 
MHNG INVE47956, Scotland, Great Cumbrae Island, 0 m, 15 May 1992, 3 subadult medusae, 
colour photographs made before fixation. - ZMUC, Dana station 4007, off Mauritania, 18.367°N 
18.233°W, 600 m wire length, 15 March 1930, 10 medusae 1.5-3 cm, id. P. Kramp. - BELUM 
Md645, Ireland, Donegal, St. Johns Point, Portnagh Rock; one mature medusa. - BMNH 
1985.9.2.25; England, Plymouth; July 1937; 3 medusa from plankton. - BMNH 1985.6.13.1; 
medusa misidentified as Leuckartiara nobilis Great Britain, Northern Ireland, Strangford Lough. 
— BMNH 1965.1.14.1-10; Sweden, Kosterfjord, Ulvillarna; 100 m; 1 Oct 1964; polyps on 
Nucula shells, with medusa buds. - BMNH 1962.11.8.1; Sweden, Gullmarfjord, Gäsö, Ranna; 
20-30 m; 27 August 1962; polyps on Dentalium (Scaphopoda), with medusa buds 


DiAGnosis: Bell higher than wide, height 2-4 cm, no exumbrellar nematocyst 
ridges, with or without apical projection, no apical canal, with about 60-80 tentacles, 
interradial gonad region large and without folds but with many gonadal pits (>20), 
eight adradial rows of gonads folds, folds directed towards interradial; no papillae on 
gonads, radial canals jagged. 


DESCRIPTION (Edwards, 1965; own observations): Hydroid phase forming dense 
colonies on shells of Nucula bivalves. Hydrorhiza a close network of creeping stolons. 
Hydranths stolonal, stems rarely branched once; perisarc of pedicels irregularly or 
spirally corrugated but not annulated, continued over hydranth as filmy pseudo- 
hydrotheca ending below tentacles. Hydranth spindle-shaped, clearly set apart from the 
thin pedicel, with prominent conical hypostome, 6-7 (range 4-9) filiform tentacles in a 
single whorl, longer and erect tentacles alternating with shorter and depressed ones. 
Medusa buds borne on the hydranth pedicels, less frequently on the stolons; pear- 
shaped to spherical, short pedicel, enclosed in filmy perisarc. Mature buds with four 
large tentacle bulbs, each bearing a tentacle. 

Newly released medusa bell-shaped, with scattered exumbrellar nematocysts, 
umbilical canal; manubrium conical, simple, 1/4 to 1/3 of subumbrellar depth; radial 
canals smooth, thick; four perradial tentacles, each with a conical base not laterally 
compressed, of these tentacles two opposite ones well developed and the two others 
less developed, no rudiments of further bulbs, no ocelli. During subsequent growth, 
tentacles are intercalated between existing ones, the manubrium increases its size, after 
a size of 2 mm the mesenteries form, when about 2.7 mm high the gonadal bulges 
commence their development, later followed by the pits, the mouth margin gets frilled. 

Adult medusa bell- to bullet-shaped, higher than wide, with solid apical process 
varying much in size, sometimes lacking; jelly relatively thin but firm. Velum narrow. 
Four broad radial canals, with short lateral diverticles; ring canal broad, with smooth 
outlines. Proximal parts of radial canals joined to stomach my mesenteries for about 
1/3 of their length. Marginal tentacles numerous, usually about sixty, up to ninety 
possible, hollow, with large, laterally compressed, elongated conical basal bulbs 
clasping the umbrella margin but not forming conspicuous abaxial spurs; without 
ocelli; nematocysts concentrated in crescent-shaped clasps, in distal regions confined 
to adaxial side of tentacle. Tentacles bases directed towards below. Developing 
tentacles or bulbs can be present, but usually rare in mature animals. Manubrium flask- 
shaped, of variable length (1/3 to 5/6 of subumbrellar height), with broad base, inten- 
sive red colour, mouth with strongly folded lips. Gonads interradial in two adradial 
series of folds directed towards the interradius (in region of mesenteries only) and a 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 337 


depressed interradial portion with numerous round pits, rendering this region reticu- 
lated. Nematocysts: microbasic euryteles, abundant on tentacles; perhaps also an 
additional, much rarer, smaller capsule. 


DIMENSIONS (Edwards, 1965; own data): Hydroid stems 1.6-2.2 mm, occa- 
sionally up to 4 mm, hydranths 0.5-1.1 mm high, medusa buds up to 0.7 mm long, 
stalks of buds 0.11-0.18 mm. Newly released medusa about 1.2 mm high. Adult 
medusa 2-3.5 cm high, diameter 1.5 cm; microbasic euryteles (7-8)x(3) um. 


VARIATION: Certain characters of the fully-grown medusa show considerable 
variation. The apical process may vary very much in size: it may be a large almost 
spherical mass comprising over a third of the total height of the medusa, or jt may be 
completely absent. The stomach also varies considerably in length, in some reaching 
almost to the umbrella margin while in others it is not even half the height of the sub- 
umbrellar cavity. 


DISTRIBUTION: The medusa is present in the north-eastern Atlantic as far north 
as 64°N, occurring along the Norwegian coast, British Isles, northern North Sea, 
Skagerrak and northern Kattegat, Bay of Biscay, off southern Portugal, western 
Mediterranean; north-west, west and south-west Africa. Also present in the mid- 
Atlantic and at Iceland. Absent in southern parts of the North Sea and the eastern 
English Channel. There is no certain evidence that the species is present in the Pacific 
and Indian Ocean (see also Kramp, 1968; Arai & Brinckmann-Voss, 1980). Edwards 
(1965) reviews all records of the medusa. The polyp stage is known from western 
Scotland, Norway, and Sweden. The preferred host of the polyp phase, Nucula sulcata 
Bronn, 1831 has a similar distribution like the medusa of Neoturris pileata (Edwards, 
1965). Type locality: Mediterranean. 


BroLoGy (Edwards, 1965): The polyps occur exclusively on shells of the 
bivalve genus Nucula, preferably on Nucula sulcata Bronn, 1831, in depths of 40-180 
m. Gonophore production in the Atlantic starts in January to February, apparently at the 
annual minimum of the water temperature and the maximum of oxygen concentration. 
The polyp is adapted to live buried within the mud. Polyps of Leuckartiara octona and 
N. pileata polyps can co-occur on the same shell. In favourable years, fully grown 
medusae appear in the Atlantic in May. The length of life of the medusa appears to be 
3-4 months. The medusae are rare and it is only in occasional seasons that fully grown, 
mature medusae are present, they may be seen in numbers swimming at and near the 
surface, in more northerly waters and in the Skagerrak and Kattegatt the medusae 
persist longer, some to October. In the Mediterranean, the medusae occur earlier than 
in the Atlantic. They are absent during the summer month. 


FURTHER DATA: Krasinska (1914) examined some histological details of the 
medusa. 


REMARKS: The synonymy established by Haeckel (1879), Bedot (1901, 1905), 
Le Danois (1914a), Hartlaub (1914) was here fully adopted. Hartlaub (1914) was able 
to examine some of the type specimens. 

The correct spelling of the author of this species is apparently Forsskal, despite 
in the title of the original publication of 1775 the spelling Forskal was used 
(Cernohorsky et al., 1991). 


338 P. SCHUCHERT 


In the distal portions of the tentacles, the nematocysts are all on one side and 
concentrated in crescent-shaped bulges. This has already been mentioned by Kramp 
(1926) and he states that the clusters lie on the adaxial side. Kramp (1926) maintains 
that the cross-section of the tentacles is triangular, but this was not the case in the 
present material. 

Neoturris pileata differs from N. breviconis in the more slender shape of the 
umbrella, the higher number of interradial pits, and the lower number of tentacles (Arai 
& Brinckmann-Voss, 1980). 

The hydroid resembles much L. octona, but according to Edwards (1965) the 
stems are less branched and shorter, as well as the hydranths and medusa buds are 
larger. The newly liberated medusa offers better traits to distinguish the colonies 
(N. pileata four tentacles, L. octona two tentacles). 


Neoturris breviconis (Murbach & Shearer, 1902) Fig. 61 


Turris breviconis Murbach & Shearer, 1902: 73.— Murbach & Shearer, 1903: 170, pl. 18 figs 1- 
2.- Mayer, 1910: 127. 

Leuckartiara brevicornis. — Hartlaub, 1914: 304, figs 254-256 [subsequent incorrect spelling]. 

Leuckartiara breviconis. — Kramp & Damas, 1925: 280. — Kramp, 1926: 80, pl. 2 fig. 8. — 
Uchida, 1938b: 49, fig. 1. — Russell, 1953: 198, pl. 12 fig. 2. — Kramp, 1959a: 120, 
fig. 121. - Kramp, 1961: 103. — Kramp, 1968: 4, fig. 124. 

Perigonimus breviconis. — Naumov, 1969: 204, fig. 72. 

Neoturris breviconis. — Arai & Brinckmann-Voss 1980: 57, figs 30- 33. 


MATERIAL EXAMINED: MHNG INVE35729, British Columbia, Malcolm Island, Mitchell 
Bay, 5 June 2002, coll. & id. A. Brinckmann-Voss, 1 medusa of about 1 cm, subadult, ca. 80 ten- 
tacles mostly equally developed, gonad folds and pits weakly developed, pits numerous. — 
BMNH 1954.11.13.68; North Sea; 14 March 1913; 1 medusa badly preserved, could also be L. 
nobilis, leg. & det. C. Hartlaub. - ZMUC, Thor station 241(04); 64.583°N 11.750°W; eastern 
Atlantic; 8 August 1904; 5 medusae; material described in Kramp (1926). — ZMUC, no number; 
Iceland, Vestmanaeyjar; 12 July 1904; 5 medusae; material described in Kramp (1926). 


DIAGNOSIS (Atlantic form): Height 2-4 cm, as broad as high, with up to 140 ten- 
tacles, no exumbrellar nematocyst ridges, with shallow apical projection, no apical 
canal, gonad folds arranged like horseshoe, folds directed towards perradial and aboral, 
relatively few interradial gonadal pits (<20), no papillae on gonads; radial canals 
jagged; mesenteries rather short. 


DESCRIPTION (Atlantic form): Medusa bell relatively wide, as broad as high or 
slightly higher, with apical rounded dome-shaped process of variable height, lateral 
jelly relatively thin. Four broad radial canals with much jagged outlines, ring canal 
smooth; radial canals joined to stomach by mesenteries, these relatively short (1/2 or 
less of manubrium length), abaxial side with transverse ridges. Marginal tentacles up 
to 105, one row, many not fully developed, sizes thus very variable, no or very few 
rudimentary bulbs, tentacles hollow, laterally compressed, clasping umbrella but not 
forming abaxial spurs. Manubrium flask-shaped, relatively large (2/3 of bell cavity), 
broad base, mouth with much crenulated lips. Gonads folds, arranged like a horse- 
shoe, folds relatively irregular, directed mostly towards perradial, central interradial 
region somewhat depressed and in large animals with up to 20 gonadal pits and also 
some folds. Polyp stage unknown. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 339 


i 
9 


Fic. 61 


Neoturris breviconis (Murbach & Shearer, 1902). (A) Pacific form, about 20 mm high, from 
Uchida (1938b). (B) Bell margin and tentacles of medusa shown in A. (C) Bell margin with 
tentacles of an Atlantic medusa, modified from Hartlaub (1914). (D) A relative young specimen 
from the North Sea, height about 15 mm, from Hartlaub (1914). (E) Typical gonad of a mature 
Atlantic medusa from east of Iceland, 20-30 mm wide and about 60 tentacles. 


VARIATION: Derived from data and material of Kramp (1926), the following 
approximate correlation of bell diameter and tentacle number can be given for the 
Atlantic form: bell diameter 4 mm 16 tentacles; 5 mm 12-32 tentacles, 7 mm 16-46 
tentacles; 10 mm 40-68 tentacles, 11 mm 38-65 tentacles; 20-30 mm 60-64 tentacles; 
30 mm 89 tentacles. 


340 P. SCHUCHERT 


The Pacific form differs in the following details: tentacle number is higher (up 
to 140; Uchida (1938) had an animal of 15 mm diameter with 120 tentacles; the 
examined specimen was about 10 wide and had around 80 tentacles), the tentacles are 
more equally developed, the bell margin is very densely beset with tentacles so that it 
get crowded and the tentacles may appear in two rows; the lateral mesogloea thicker; 
stomach with gonads distinctly shorter (1/3 of bell cavity), mouth part about as long as 
stomach part; gonad folds short, pitted region larger, with more pits (about twice or 
more). 


DIMENSIONS: Atlantic form 10-40 mm high and up to 30 mm wide; Pacific 
medusa 45 mm high and 35 mm broad. 


OTHER DATA: Kramp (1926) reported that the large tentacles have abaxial trans- 
versal folds in the distal region, contrary to many other Pandeidae which have adaxial 
folds (could not be confirmed for Pacific specimen). Nematocysts: microbasic 
euryteles and mastigophores (Mackie & Mackie, 1963). 


DISTRIBUTION: Currently seen as a circumpolar Arctic to northern boreal species. 
In the eastern Atlantic, the southern limits are the northern North Sea, Firth of Clyde, 
western Ireland, and the south of Iceland (Kramp & Damas, 1925; Kramp, 1926; 
Naumov, 1969; Russell, 1953; Russell, 1970; Zelickmann, 1972; Boyd et al., 1973). In 
the Pacific it occurs as far south as British Columbia and Japan (Uchida, 1938b; Arai & 
Brinckmann-Voss 1980). Type locality: St. Paul Island, Pribyloff Islands, Bering Sea. 


BIOLOGY: A quite rare medusa, recorded in the NE Atlantic during the summer 
months (Russell, 1953; Boyd er al., 1973). 


REMARKS: The examined specimen from the north-eastern Pacific, likely a good 
representative for the typical form, differs somewhat from the Atlantic specimens (see 
Variation). Uchida (1938b) gives a good figure of the Pacific variant and it is here 
reproduced in Fig. 61A. A good colour photograph of a living medusa from the Pacific 
is given in Heeger (1998) and also Arai & Brinckmann-Voss (1980) show a photo- 
graph. It is evident that the Pacific form corresponds much more to a Neoturris than 
the Atlantic specimens and the species is here left in this genus following Arai & 
Brinckmann-Voss (1980). The Atlantic specimens have gonads that resemble more 
those of Leuckartiara species (Fig. 61E). I therefore suspect that the Atlantic N. brevi- 
conis (sensu Hartlaub, 1914; Russell, 1953; Kramp 1959a) belongs to a different 
species than the Pacific N. breviconis. It could even be that the Atlantic form of N. bre- 
viconis is actually L. nobilis with exceptionally high tentacle numbers and much 
developed gonads. It must also be noted that most of the so far described Atlantic 
specimens had been in preservative for years or decennia. As the pigment the ocelli 
fades rapidly in formalin, it is therefore not certain whether they really lacked ocelli 
originally. New investigations using living animals are needed to re-evaluate the status 
of the Atlantic population of N. breviconis. 


Genus Halitholus Hartlaub, 1914 
TYPE SPECIES: Halitholus pauper Hartlaub, 1914, designation by Kramp (1959a). 


DIAGNOSIS: Pandeid medusae with four or more tentacles, large dome-shaped 
apical projection, quadrangular manubrium without mesenteries, gonads adradial with 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 341 


or without horizontal interradial connection, gonad folds directed towards perradial, 
mouth margin moderately folded; with or without ocelli. 

Hydroid stolonal, hydranths on short pedicels, with pseudohydrotheca, hypo- 
stome conical, filiform tentacles, number relatively low (around six); medusa buds on 
stolons. 


REMARKS: The genera Halitholus and Leuckartiara are quite similar, but the ab- 
sence of mesenteries makes Halitholus quite easily separable. 


KEY TO THE HALITHOLUS SPECIES OF THE NORTH ATLANTIC: 


la 30-40 tentacles, gonads without interradial connection, without ocelli . . . . . H. cirratus 

lb 4-8 tentacles, adradial portions of gonads with interradial connection, with ocelli 
3 RN Un i ieh Arster ee ‘H. pauper 

Halitholus cirratus Hartlaub, 1914 Fig. 62 


? Perigonimus yoldia-arcticae Birula, 1897: 88, pl. 10 fig. 3. — Naumov, 1969: 200, fig. 67. 

Perigonimus cirratus Hartlaub, 1914: 274, fig. 234. — Arndt, 1964: 332, figs 1-3. 

Halitholus cirratus Hartlaub, 1914: 274, figs 225-234. — Kramp, 1926: 74, pl. 2 fig. 4, chart XII. 
— Kramp, 1959a: 119, fig. 117. — Kramp, 1961: 101.— Kramp, 1968: 45, fig. 116. 

? Halitholus yoldiaarcticae. — Schuchert, 2001: 22, fig. 12. 


MATERIAL EXAMINED: BMNH 1950.3.1.203; Baltic Sea; 14 March 1913; 12 medusae; 
leg. and det. C. Hartlaub. - ZMUC, Dana Station 2965; Denmark, SW of Bagenkop; 30-80 m 
wire; 21 April 1923; 50 damaged medusae. — ZMUC, Dana Station 3050; Denmark, N of Anholt; 
depth 15 m; 18 May 1923; 7 medusae. — IRSNB; Baltic Sea; 3 damaged medusae; coll. J. 
Bouillon. 


DrAGNosis: Medusa 5-16 mm high, large apical projection; with shallow 
manubrial peduncle; lips only weakly folded; 30-40 tentacles; no ocelli; gonads 
irregular and variable, often in eight vertical rows of folds. Polyp stolonal, on bivalves, 
with pseudohydrotheca, about six tentacles, medusa buds on stolons. 


DESCRIPTION (after Hartlaub, 1914; own data): Polyp stage resembles closely L. 
octona (Figs. 53A; 62B and D), about six tentacles, stems shorter but length variable, 
medusa buds so far only observed on stolons. Newly released medusa with two 
tentacles. 

Mature medusa with wide bell, apical process of variable size and shape, may 
be half the total height of the medusa. Mesogloea of lateral wall relatively thin, 
manubrium on a shallow peduncle. Four radial canals moderately broad, smooth or 
with wavy outlines, not forming mesenteries, at top recurved due to shallow manubrial 
peduncle; ring canal smooth. Marginal tentacles 30-40, rarely up to 50, no rudimentary 
bulbs, bulbs gradually tapering into tentacles, laterally compressed, without abaxial 
spurs, without ocelli, nematocysts concentrated on adaxial side. Manubrium volu- 
minous, cross-shaped in section, reaching almost down to level of velum, mouth region 
relatively short; mouth wide, rim only weakly folded: Gonads very irregular, usually 
in eight adradial series, usually with transverse folds directed towards perradius, 
gonads widely separated perradially; interradial region depressed and wide, rather 
smooth, only occasionally with folds, interradial connecting fold absent, usually no pits 
present, sometimes isolated pits on gonadal folds; sometimes gonads without folds and 
upper ends fused interradially. Colours: Manubrium brick-red. 


342 P. SCHUCHERT 


\ 


Fic. 62 


Halitholus cirratus Hartlaub, 1914. (A) Mature medusa, scale bar 2 mm. (B) Hydroid, size of 
polyps about 2 mm in height. (C) Newly liberated medusa. (D) Hydroid on the bivalve Astarte 
borealis (Schumacher), note the empty envelope of a medusa bud (lower right), scale bar 1 mm. 
A, after preserved material; B-C, from Hartlaub (1914); D, modified after Arndt (1964). 


DIMENSIONS: Mature medusa up to 16 mm high, diameter up to 14 mm, usually 
smaller (10 mm), in Baltic Sea 5-11 mm high. Hydroid up to 2 mm high (Hartlaub, 
1914), hydranth body 0.8-0.8 mm, pedicel 0.2 mm (Arndt, 1964). 


DISTRIBUTION: A costal Arctic species occurring along the northern coasts of 
Eurasia and America, western Greenland; frequent at Spitsbergen (Kramp, 1926; 
Naumov, 1969; Kramp 1961); present also in the Baltic Sea and the Kattegat. Type 
locality: not specified, Hartlaub had material from Svalbard, the Barent Sea, the Baltic 
Sera and the North Sea. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 343 


BIOLOGY: At Spitsbergen, the medusa is common from June to August (Kramp, 
1926). The putative hydroid phase described by Arndt (1964) and Schönborn er al. 
(1993) occurred on shells of Nuculidae and Astartidae and other bivalve shells, down 
to 100 m, it supported reduced salinities down to 8 ppt, medusa budding was observed 
in March and April. The polyps fed on nematodes. 


REMARKS: The putative polyp of this species was first described by Hartlaub 
(1914) based on colonies growing on an isopod collected in the Baltic Sea. Hartlaub 
did not rear the medusa to maturity, but concluded from the absence of any other pan- 
deid medusa in the Baltic Sea that this must be the hydroid of Halitholus cirrata. 
Although Hartlaub’s conclusion is likely correct, there remains an element of uncer- 
tainty and the life cycle of this species should be reinvestigated. The hydroids of many 
pandeids and bougainvilliids are exceedingly similar and offer few or no reliable traits 
to distinguish them securely. The only trait that distinguishes Hartlaub’s polyp from 
similar ones like L. octona is its lower tentacle number (about 6). The polyp was also 
described and depicted by Arndt (1964), but without providing information on the 
medusa stage. Arndt (1964) also found the medusa at the same locality where he had 
collected the hydroid. It is thus very likely that they had the polyp of A. cirratus. 

The hydroid Perigonimus yoldiaarcticae Birula, 1897, growing preferably on 
shells of the Arctic bivalve Yoldiella, is at present not sufficiently described as its life 
cycle is unknown (see also Naumov, 1969; Schuchert, 2001). Birula’s polyp could 
belong to several other Arctic Bougainvilliidae or Pandeidae species. Birula’s polyp 
had also only few tentacles and Naumov (1969) thought that it is conspecific with 
Halitholus cirratus Hartlaub, 1914. Although this appears plausible, Naumov’s claims 
were apparently not based on rearing experiments starting with polyps on Yoldiella 
shells. I therefore hesitate to synonymize A. cirratus and P. yoldiaarctica. 


Halitholus pauper Hartlaub 1914 Fig. 63 


in part Tiaria conifera Levinson, 1893: 144. — Mayer, 1910: 128. 

[not Tiara conifera Haeckel, 1879] 

Halitholus pauper Hartlaub, 1914: 272, figs 223-224. — Kramp, 1926: 71, pl. 2 figs 1-3, chart XI. 
— Uchida, 1933: 128, figs 4-5. — Kramp, 1926: 71 pl. 2 figs 1-3. — Kramp, 1959a: 119, 
fig 116. — Kramp, 1961: 101. — Kramp, 1968: 45, fig 115. — Arai & Brinckmann-Voss, 
1980: 46, fig. 22. — Schuchert, 1996: 67, fig. 39. 


MATERIAL EXAMINED: Holotype, ZMUC, loc. Greenland, Egedesminde; four fully 
formed tentacles, interradial tentacles only rudiments. — IRSN Bruxelles, IG 27.838, loc. 
Greenland, 3 medusae. 


DraGNOSIS: Medusa with 4-8 tentacles, with small ocelli, adradial portions of 
gonads connected interradially by transverse fold. 


DESCRIPTION: Umbrella broad, with low rounded apical projection, no or very 
shallow manubrial peduncle; four radial canals, rather broad, margins smooth or some- 
times partly jagged, entering stomach at its top; ring canal smooth. With four perradial 
tentacles, these with thick base, then tapering, not much laterally compressed, clasping 
bell margin, abaxial ocelli; four interradial tentacles can be present, these often small 
or only as rudimentary bulbs; usually also several rudimentary adradial bulbs. Manu- 
brium massive, reaching 2/3 of subumbrellar height, cross-shaped in section, mouth 


344 P. SCHUCHERT 


Fic. 63 
Halitholus pauper Hartlaub, 1914; after preserved material from Greenland, scale bar 1 mm. 


margin moderately folded; gonads adradial with a distinct transverse interradial 
connection in the upper part giving a horse-shoe shape, folds and openings directed 
towards perradial and upwards. Colours: gonads orange or reddish-orange (Arai & 
Brinckmann-Voss, 1980). 

Polyp stage unknown. 


DIMENSIONS : Medusa 5-12 mm high. 


DISTRIBUTION: Iceland, western and eastern Greenland, Arctic waters of Canada, 
British Columbia, southern Kamchatka, northern Japan, New Zealand (Arai & 
Brinckmann-Voss, 1980; Schuchert, 1996). Type locality: Western Greenland, 
Egedesminde (Aasiaat). 


REMARKS: This species is so far not known to occur along the continental coasts 
of Europe, but it could be present in the Arctic regions of Eurasia. 


Genus Catablema Haeckel, 1879 
TYPE SPECIES: Turris vesicaria A. Agassiz, 1862 [designation by Kramp, 1959a]. 
REFERENCES: Haeckel, 1879; Hartlaub, 1914; Kramp, 1961; Bouillon & Boero, 2000. 
DrAGNosis: Medusa with large apical projection; radial canals broad and 
strongly jagged; mesenteries relatively short; tentacles numerous, with or without rudi- 
mentary tentacle-bulbs between normal tentacles; with abaxial ocelli; manubrium large 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 345 


with broad base; mouth large with much folded rim; gonads in long folds that tend to 
be vertical in the interradial region, gonadal pits may be present on and between the 
folds; gonads separated perradially. 


REMARKS: The gonads on long, oblique to almost vertical stomach folds is the 
main characteristic of this species. To Haeckel (1879) they resembled to a theatre- 
curtain and he created the name Catablema — meaning curtain in Greek — as an allusion 
to this character. The gender is neuter. Only one species is present in area of inves- 
tigation, Catablema vesicarium. 


Catablema vesicarium (A. Agassiz, 1862) Fig. 64 


Turris vesicaria A. Agassiz, 1862: 97.— A. Agassiz, 1865: 164, figs 261-268. 
Catablema campanula Haeckel, 1879: 63. — Maas, 1904: 13, synonym. 
Catablema eurystoma Haeckel, 1879: 63. — Maas, 1904: 13, synonym. — Hartlaub, 1914: 319, 


synonym. 

? Tiara conifera Haeckel, 1879: 59. — Hartlaub, 1914: 284, likely synonym. 

Catablema vesicarium. — Haeckel, 1879: 64. — Maas, 1904: 12, pl. 1 fig. 8, pl. 2 fig. 10. — 
Bigelow, 1909: 304, pl. 30 figs 3-4, pl. 31 fig. 6. — Hartlaub, 1914: 315, figs 263-267. — 
Kramp, 1926: 87, pl. 2 figs 10-11, chart 17. — Kramp, 1959a: 122, fig. 125. — Kramp, 
1961: 96. — Kramp, 1968: 50, fig. 132. 

in part Turris vesicaria. — Mayer, 1910: 126, pl. 12 figs 2-3, not pl. 13. fig 7. 

not Catablema vesicarium var. nodulosa Bigelow, 1913: 17, pl. 1 figs 8-9 [= C. nodulosa 
Bigelow, 1913]. 

in part Perigonimus vesicarius. — Naumov 1969: 202, fig. 69 [some perhaps C. nodulosa]. 


MATERIAL EXAMINED: ZMUC; Greenland, Godthaab Station 50, 69.740°N 57.367°W; 
200 m wire; 13 July 1928; 10 medusae; det. Kramp. — ZMUC; Dana Station 2362, Greenland, 
63.583°N 57.183°W; 50-320 m; 320; 27 June 1925; 5 damaged medusae. 

DrAGNosis: Medusa up to 25 mm wide and 30 mm high, including the large, 
globular apical projection; gonads in long, irregular folds, oblique in lateral parts, 
almost perpendicular in middle part of each gonad, with or without densely reticulate 
surfaces; about 32 tentacles; mesenteries short. 


DESCRIPTION (Hartlaub, 1914; own data): Medusa with rather broad umbrella 
bearing a large globular or dome-shaped apical process, lateral bell walls relatively 
thin. Four radial canals, very broad and with lateral diverticules; ring canal broad, with 
undulated to jagged outlines. Proximal parts of radial canals joined to stomach my 
mesenteries for up to 1/4 of their length, thus relatively short mesenteries. Marginal 
tentacles up to about 32, rarely more, with large conical basal bulbs, laterally 
compressed, clasping the umbrella margin, forming slight abaxial spurs; with red 
ocelli, can be inconspicuous; nematocysts concentrated on adaxial side of tentacle, 
adaxial side in contracted tentacles corrugated; between tentacle pairs often a wart-like 
tentacle-rudiment. Gastrodermal chamber of some tentacle bulbs has finger-like 
protrusion reaching upward into the mesogloea (Fig. 64B). Manubrium flask-shaped, 
voluminous, size variable, sometimes reaching nearly to velum level, with broad base, 
mouth with strongly folded lips. Gonads on interradial wall of stomach, in irregular 
folds, oblique in lateral parts, almost perpendicular in middle part of each gonad, 
giving impression of radiating from a centre; in many larger specimens also with 
numerous pits on and between the folds, producing a reticulate surface. Colour of 
manubrium yellow-orange (after living specimens from Spitsbergen seen by Hartlaub, 
1914). 


346 P. SCHUCHERT 


) 


Catablema vesicarium (A. Agassiz, 1862). (A) Mature medusa, specimen without gonadal pits, 
schematic reconstruction after several preserved medusa from Greenland, scale bar 0.4 mm. (B) 
Enlarged lateral view of tentacle base, note villi of gastrodermal chamber that project into 
mesogloea. (C) Manubrium of young medusa (5 mm high, 11 tentacles) with beginning 
formation of the gonadal folds, from Hartlaub (1914). (D) Manubrium of large specimen that 
also have numerous pits on the gonadal folds; from Hartlaub (1914). 


ne 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 347 


Polyp stage unknown, the smallest known medusae (3.5 mm diameter; Kramp, 
1926) had four large perradial tentacles, four somewhat smaller interradial tentacles, 
eight adradial very short tentacles, and 16 tentacular rudiments. The radial canals were 
already jagged at this stage. 


DIMENSIONS: Medusa up to 25 mm wide and 30 mm high (including the large, 
globular apical projection), size at maturity is very variable (diameters 12-25 mm, 
Kramp, 1926). 


DISTRIBUTION: An Atlantic medusa confined to Arctic regions, rarely penetrating 
into boreal regions. There is a record from northern Norway from the region south of 
The Faroes (Fraser, 1973). It is also known from north of Iceland and at Greenland; 
coast of North America from Cape Cod northward (Hartlaub, 1914; Kramp, 1926; 
1959a; Zelickman, 1972). Pacific records are uncertain (Arai & Brinckmann-Voss, 
1980). Type locality: Nahant, Massachusetts Bay, USA. 


BIOLOGY: A cold water species, usually collected during the summer months 
(Zelickmann, 1972). The polyp stage is unknown. 


REMARKS: The synonymy of this species has been discussed by Maas (1904) 
and Hartlaub (1914). Haeckel placed this species in the genus Catablema and changed 
the ending of original specific epithet vesicaria to vesicarium. This was spelling was 
also used by most subsequent authors. 

Hartlaub (1914: 321) and Kramp (1926: 88) regarded Catablema vesicarium 
var. nodulosa Bigelow, 1913 as a synonym of C. vesicarium. This view was contested 
by Brinckmann-Voss (1980) who raised the variant to full species level (Catablema 
nodulosum Bigelow, 1913). The latter species has only 8-16 tentacles, rarely 32. The 
shape of the gonads cannot be used to distinguish the two species. 

Catablema multicirratum Kishinouye, 1910 from the North Pacific and the 
north-western Atlantic is similar to C. vesicarium, but larger (up to 60 mm high) and it 
has more than 80 tentacles (see Arai & Brinckmann-Voss, 1980). 


Genus Merga Hartlaub, 1914 
TYPE SPECIES: Pandea violacea Agassiz & Mayer, 1899 by designation of Hartlaub 
1914). 
i SYNONYMS: Tiarula Hartlaub, 1914 [type species Tiarula tergestina Neppi & Stiasny, 
1912]. — Mergintha Hartlaub, 1914 [type species Mergintha lobianci Hartlaub, 1914]. 

DIAGNOSIS: Pandeid medusae with smooth or granulate adradial or interradial 
gonads, with mesenteries, stomach with cross-shaped base, manubrium not twisted, 
with simple or faintly folded oral lips; four, eight, or more tentacles, with or without 
rudimentary bulbs or tentaculae, with or without ocelli. 

Hydroids where known colonial, arising from tubular, ramified hydrorhiza, 
cauli slightly branched or not, with or without pseudohydrotheca, when present not 
enveloping tentacles, one whorl of filiform tentacles. Medusa buds arise from stems or 
stolons. 


REMARKS: The genus is discussed by e. g. Hartlaub (1914), Russell (1953), 
Kramp (1959a), Vannucci & Yamada (1959), Picard (1960), Arai & Brinckmann-Voss 
(1989). Hartlaub established this genus to accommodate pandeid medusae that 


348 P. SCHUCHERT 


resemble Leuckartiara or Halitholus species but that have gonads without folds. 
However, some (M. tergestina) can have variably some folds in their gonads, which 
somewhat compromises the usefulness of this genus. The species Merga reesi Russell, 
1956b has gonads with a warty-granulated surface and therefore fits not well into the 
genus. In order to keep this species within Halitholus, Kramp (1959a) modified the 
definition by allowing also granulated gonad surfaces. Arai & Brinckmann-Voss (1989) 
proposed that it should be removed from this genus and united it together with 
Amphinema krampi, a species with similarly structured gonads. Although this appears 
very reasonable, I refrained from doing so as this should be made based on a compre- 
hensive phylogenetic analysis of all Pandeidae genera. 


KEY TO THE EUROPEAN MERGA SPECIES: 


la with distinct apical process, at least some bulbs with abaxial spurs .............. 2 
1b without apical process, no abaxial spurs on marginal bulbs .................... 3) 
2a 4-8 tentacles, up to 8 rudimentary bulbs, mesenteries short........ Merga tergestina 
2b 8-16 tentacles, perradial bulbs with band-shaped ocelli, mesenteries long Merga galleri 
3a gonad rough by warts and groves, red-brown manubrium, with tentaculae . Merga reesi 
3b gonads smooth, no tentaculae . .........12 I I E 4 
4a 4 perradial tentacles, no rudimentary bulbs, no tentaculae, no ocelli . Merga tregoubovi 
4b 8-12 tentacles, 24-36 rudimentary bulbs, all with ocelli ............ Merga violacea 
Merga tergestina (Neppi & Stiasny, 1912) Fig. 65 


Tiaria tergestina Neppi & Stiasny, 1912: 556. — Neppi & Stiasny, 1913: 21, pl. 1 fig. 10. 

Merga tergestina. — Kramp, 1959a: 116, not fig. 107 [= Amphinema dinema]. — Vannucci & 
Yamada, 1959, 320, figs 1-7. — Vannucci, 1960: 393, figs 1-17. — Kramp, 1961: 107. — 
Brinckmann-Voss, 1970: pl. 10, fig. 1. — Goy er. al., 1991: 110, ? fig. 28. — Bouillon er 
al., 2004: 72, fig. 411-J. 

in part Tiarula tergestina. — Hartlaub, 1914: 253, fig. 208, not 209. 


MATERIAL EXAMINED: BMNH 1959.6.8.1-5; Italy, Naples; coll. 1959; some; young 
medusae reared from polyp by M. Vannucci, not well preserved. - ZMUC; Italy, Naples; coll. 5 
June 1959; 5 well preserved medusae, mature; leg. M. Vannucci, likely grown from hydroid. 


DIAGNOSIS: Merga medusa with distinct apical projection, 4-8 tentacles, up to 8 
rudimentary bulbs, with simple ocelli on abaxial spurs, spurs also on interradial ten- 
tacle-bulbs, no tentaculae, mouth margin not much folded, mesenteries short. 


DESCRIPTION (Vannucii & Yamada, 1959; Vannucci, 1960; own data): Polyp 
stage resembles L. octona (Figs 53A, 65B) and several other Pandeidae; colonies 
stolonal, hydrorhiza creeping, reticulate stolons, hydrocaulus of variable length from 
short to almost sessile to relatively long, hydrocaulus covered by thin, corrugated, 
horn-coloured perisarc that may or may not continue as pseudohydrotheca over 
hydranth body; 6-9 tentacles (normally 8); hypostome conical. Medusa buds arise from 
stolons and hydrocauli, often with relatively long pedicel but also short ones occur, bud 
ovoid, enclosed in thin perisarc membrane. Hydranth colourless. Nematocysts: two 
types of microbasic euryteles and desmonemes. 

Newly liberated medusa with apical process, umbrella oval, umbilical canal 
present, radial canals wide, abaxial side irregular, adaxial side smooth, two long per- 
radial tentacles (extended up 25 times bell height), ocelli develop rapidly on abaxial 
spurs of the bulbs. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 349 


Fic. 65 


Merga tergestina (Neppi & Stiasny, 1912). (A) Schematic drawing of a medusa, scale bar 
0.5 mm. (B) Polyp with two medusa buds, scale bar 0.5 mm. (C) Variability of gonad form and 
lip folds; scale bar 0.5 mm, valid for whole series. (D) Variability of apical process and bell- 
shape, scale bars equal 1 mm. A, after preserved material; B, modified from Vannucci & Yamada 
(1959); C-D, from Vannucci (1960). 


350 P. SCHUCHERT 


Mature medusa with bell-shaped umbrella bearing a distinct apical projection of 
variable size and form, lateral walls relatively thin; four radial canals moderately 
broad, mostly smooth, attached to manubrium through short mesenteries (about 1/6 or 
less of manubrium height). With 4-9 long marginal tentacles plus about the same 
number of small intercalated rudimentary bulbs, no tentaculae; tentacle bases laterally 
compressed and forming long abaxial spurs bearing an ocellus at its end, length of 
spurs variable; bulbs taper quite abruptly into thinner part of tentacle; rudimentary 
bulbs small, with or without ocelli. Manubrium large, in fully grown animals reaching 
almost to velum level, cruciform in section, mouth region relatively small, mouth 
margin with a few folds. Four large interradial gonads reaching to perradial, forming 
bulging adradial pads, surface usually smooth, but in very advanced specimens with up 
to five irregular folds (may also depend on state of contraction). Nematocysts: 
microbasic euryteles. Colour of manubrium and bulbs in animals from nature brownish 
(Neppi & Stiasny, 1913). 


DIMENSIONS (Vannucci & Yamada, 1959; Vannucci, 1960): Hydranth 0.24-0.62 
mm high; mature medusa buds 0.5-0.68 mm wide and 0.7-0.8 mm long; newly 
liberated medusa bell height 0.98-1.05 mm, apical process up to 0.3 mm. Mature 
medusa up to 7 mm high and 4 mm wide, more usually 3-4 mm high; egg size 68-84 
um. More data and dimensions of nematocysts are given in Vannucci (1960). 


OTHER DATA: The polyp and young medusa were first described by Vannucci & 
Yamada (1959). The morphological variation of the polyp and medusa has been inves- 
tigated in detail by Vannucci (1960). Hydroids from different substrates differ 
considerably in habitus so that they appear to belong to different species, but in culture 
they become more similar. The hydroids differed in hydranth size, tentacle number, 
perisarc development, length of hydrocaulus, and also nematocysts. This paper also 
reports upon the early embryology. Growth stages of the medusa are also tabulated in 
Neppi & Stiasny (1913). 


BIOLOGY (Vannucci, 1960): The hydroid has been found on spines of the sea 
urchin Stylocidaris affinis (Philippi), regularly on the parapodia of large specimens of 
the polychaete Laetmonice hystrix (Savigny, 1820)(= Hermione hystrix), on shell of 
Murex spec., and on tube of the polychaete Spirorbis sp. Depth range: 15-60 m. The 
medusa lived up to 145 days in vitro. They started to spawn eggs when 14-26 days old 
and continued to do so almost daily for a period of up to 24 days. Eggs of L. octona 
could rarely be fertilized with sperms of M. tergestina and a hybrid planula developed. 
Neppi & Stiasny (1913) found the medusa in the plankton near Trieste from July to 
October, mature animals were present from August onwards. 


DISTRIBUTION: Mediterranean and Papua New Guinea (Bouillon, 1980). The 
only record for the Atlantic (Kramp, 1955b; Gulf of Guinea) has been attributed to 
Ampinema dinema by Vannucci (1960). In the Mediterranean, it has been reported from 
the Adriatic Sea (Neppi & Stiasny, 1913; Benovic & Lucic, 1996; medusa phase), the 
Tyrrhenian Sea (Vannucci & Yamada, 1959; Brinckmann-Voss, 1970; polyp), Ligurian 
Sea (Vannucci, 1960; polyp), and off Lebanon (Goy et al.. 1991; medusa). Type 
locality: Trieste, Mediterranean. 


ni 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 351 


REMARKS: The interclonal and intraclonal variation of the polyp was investigat- 
ed by Vannucci (1960). The morphology of the polyps is considerably influenced by 
the environment. In view of the variation encountered, Vannucci found that it is 
impossible to identify many pandeid and bougainvilliid hydroids down to the genus 
and species level whenever gonophores are absent. 


Merga galleri Brinckmann, 1962 Fig. 66 


Merga galleri Brinckmann, 1962: 1, figs 1-3.— Brinckmann-Voss, 1970: pl. 10 fig. 3.— Bouillon 
et al., 2004: 71, fig. 41G-H. 


MATERIAL EXAMINED: Syntype; ZMUC; Italy, Naples, Mergellina, 6 m; 13 July 1961; 
hydroid on Cerithium vulagatum; soft tissues almost completely macerated. — Syntype; ZMUC; 
one medusa, leg. A. Brinckmann; 27 Sept. 1960. 


DIAGNOSIS: Merga medusa reaching 8 mm, with 16 bulbs and 8-16 tentacles, 
large apical process, only perradial tentacle-bulbs with long abaxial spurs and band- 
shaped ocelli, lips much folded. 


DESCRIPTION (Brinckmann, 1962; own observations): Polyp stage resembles 
Dicoryne spec. (Figs 18A, 19A, 66A) and other moderately branched Bougainvilliidae; 
hydranths stolonal or on sparingly branched shoots with up to four polyps; hydrorhiza 
creeping, reticulate stolons; stolons and shoots covered by thin perisarc, perisarc 
infested with detritus, widening distally into a wrinkled pseudohydrotheca; hydranth 
with conical hypostome, one whorl of four to six filiform tentacles. Medusa buds arise 
on the hydrocauli, ovoid with short pedicel, surrounded by thin perisarc. Nematocysts: 
microbasic euryteles and desmonemes. 

Newly liberated medusa with apical process and two perradial tentacles, ocelli 
on bulbs bearing no tentacles. 

Mature medusa with bell-shaped umbrella, nearly as wide as high, bearing a 
large apical projection; four broad, jagged radial canals, ring canal also jagged on 
upper side, radial canals attached to manubrium via mesenteries of moderate length 
(about 1/3 of manubrium height). Bell margin with 16 large conical bulbs bearing 8-16 
tentacles, lengths unequal; four perradial bulbs with long abaxial spurs and reddish 
ocelli in a semi-circular band, other bulbs without spurs and with dot-like ocelli. 
Manubrium large (1/2 of subumbrella depth) and broad, cruciform in section, mouth 
region separated by a neck, mouth margin strongly folded; gonads in eight bulging 
adradial pads, rarely with an interradial connection, surface smooth or one or two weak 
folds. Nematocysts: microbasic euryteles and desmonemes. 


DIMENSIONS (Brinckmann, 1962): Hydroid up to 1 cm, newly liberated medusa 
0.9-1.2 mm high and 0.9-1.3 mm wide. Adult medusa 9-11.5 mm total height, 3.5-4.5 
mm bell diameter. Dimensions of the nematocysts are also given in Brinckmann 
(1962). 


BroLoGY: The polyp was found on shells of the gastropod Cerithium vulgatum 
Bruguière in a depth of 6-10 m. The medusa is only known from rearing experiments. 
It took 24-28 days to reach sexual maturity (Brinckmann, 1962). 


DISTRIBUTION: Known from type locality only, outside Mergellina Harbour, 
Naples, Italy. 


352 P. SCHUCHERT 


SP 
RTS fa 
Pac? nn È 
ARL EN SERA 
5 GOT IENE à Anne à 
> i 


duro 


hr 


FIG. 66 


Merga galleri Brinckmann, 1962; modified after Brinckmann (1962). (A) Shoot of hydroid 
colony, with medusa buds. (B) Mature medusa, total height about 8 mm. 


Merga reesi Russell, 1956 Fig. 67 


Merga reesi Russell, 1956b: 493, fig. 1. — Kramp, 1959a: 116, fig. 108. — Kramp, 1961: 105. — 
Russell, 1970: 246, fig. 9s. — Brinckmann-Voss & Arai, 1998: 48, fig. 4. 


MATERIAL EXAMINED: Holotype, BMNH 1956.4.14.1; Bay of Biscay 47.20°N 07.67°E, 
821m wire out; coll. 21 July 1955; on medusa, badly damaged and macerated, almost useless. 

DIAGNOSIS: Merga medusa without apical process, four perradial tentacles and 
four interradial tentaculae, red-brown gonad, surface with warts and grooves. 


DESCRIPTION (Russell, 1956b; Brinckmann-Voss & Arai, 1998): Umbrella bell- 
shaped, higher than wide, jelly thin, no apical process; four radial canals smooth, 
attached to manubrium for more than half their length via mesenteries, ring canal 
smooth, narrower than radial canals. Four large perradial tentacles and four small 
interradial tentaculae, without ocelli; perradial tentacles with swollen basal part 
through enlarged gastrodermis, narrowing towards bell margin. Manubrium large, 
spanning more than 3/4 of subumbrella, flask-shaped with broad base; mouth with 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 353 


FIG. 67 
Merga reesi Russell, 1956; from Russell (1970). 


slightly crenulated lips armed with a band of nematocysts. Gonads large, interradial, 
only separated perradially by mesenteries, surface irregularly corrugated by warts and 
grooves. Colour of stomach and gonads dark reddish brown. 

Polyp stage unknown. 


DIMENSIONS (Brinckmann-Voss & Arai, 1998): Bell height up to 10 mm, dia- 
meter up to 8 mm. 


OTHER DATA: Brinckmann-Voss & Arai (1998) depict the tentacle base with its 
enlarged proximal part. 


BIOLOGY: A very rare medusa occurring in depths of several hundred meters. 


DISTRIBUTION: Atlantic Ocean, off Brittany (Russell, 1956b); Indian Ocean, Bay 
of Bengal (Navas, 1971; Navas-Pereira & Vannucci, 1991); north-eastern Pacific 
Ocean (Brinckmann-Voss & Arai, 1998). Type locality: 47.20°N 07.67°E. 


REMARKS: Merga reesi shows many similarities with Amphinema krampi, 
notably both have the same peculiar gonad surface. Although the latter species is cur- 
rently clearly separable on account of its tentacle number, the apical process, and the 
tissue strands in the mesogloea, the possibility that both are actually only forms of the 
same species should not be disregarded entirely. See also comments under Amphinema 
biscayana. 


Merga tregoubovii Picard, 1960 Fig. 68 


Merga tregoubovii Picard, 1960: 333, fig. 1. — Kramp, 1961: 444. — Goy, 1973: 985. — Bouillon, 
1980: 334. — Bouillon et al., 2004: 72, fig. 42A. 


DiAGnosis: Merga without apical process, four perradial tentacles, no rudi- 
mentary bulbs or tentaculae, no ocelli. 


354 P. SCHUCHERT 


FIG. 68 


Merga tregoubovii Picard, 1960; after Picard (1960) based on preserved material, the bell could 
thus be more voluminous in living animals. 


DESCRIPTION (Picard, 1960; Bouillon, 1980): Umbrella of mature medusa rather 
spherical, jelly very thin, without apical process, without manubrial peduncle; four 
moderately broad radial canals, smooth, radial canals attached to manubrium via 
mesenteries that run along about 1/2 of the manubrium height. Bell margin with four 
large conical bulbs bearing four tentacles, no abaxial spurs, no ocelli. Manubrium 
large, quadrangular, reaching almost to velum level, four lips without folds. Gonads 
very voluminous, in eight bulging adradial masses with an interradial connection at the 
top, surface smooth; in very advanced animals the gonads are connected over the entire 
interradial surface. Nematocysts: microbasic euryteles. 


DIMENSIONS (Picard, 1960): The preserved medusae had a height and diameter 
of about 2.5 mm. 


DISTRIBUTION: A very rare medusa, only known from a few specimens from the 
Ligurian Sea (Picard, 1960; Goy, 1973) and more than hundred specimens from Papua 
New Guinea (1980). Type locality: Villefranche-sur-Mer, France, Mediterranean. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 355 


Merga violacea (Agassiz & Mayer, 1899) Fig. 69 


Pandea violacea Agassiz & Mayer, 1899: 160. — Bigelow, 1909: 205, pl. 41, figs 10-11.— Mayer, 
1900b: 34, pl. 1 fig. 1. — Mayer, 1910: 119, pl. 11 fig. 7, pl. 12 fig. 1, text fig. 64. 

Pandea spec. — Lo Bianco, 1903: 217, pl. 7 fig. 1. — Mayer, 1910: 119, synonym. 

Mergintha lobianci Hartlaub, 1914: 250, fig. 205. — Kramp, 1953: 265, synonym. 

Merga violacea. — Hartlaub, 1914: 249, fig. 204. — Menon, 1932: 7, pl. 1 fig. 10. — Kramp, 1953: 
265.— Kramp, 1959a: 116, fig. 106. — Kramp, 1961: 107. — Kramp, 1962: 313.— Kramp, 
1968: 41, fig. 104. — Goy, 1973: 985. — Schmidt, 1973a: 21. — Bouillon et al., 2004: 72, 
fig. 42B. 

? not Merga violacea. — Goy et al., 1991: 111, fig. 28. 


MATERIAL EXAMINED: ZMUC, as Merga lobianci, Thor station 147; 31.583°N 19.150°E; 
southern Mediterranean; 25 July 1910, det. Kramp. — ZMUC, as Merga lobianci, Thor station 
200; 39.300°N 10.183°E; Tyrrhenian Sea; 25 m; 26 Aug. 1910; 2 damaged medusae, det. Kramp. 
— ZMUC, as Merga lobianci, Thor station 142; 35.733°N 15.117°E; southern Mediterranean; 25 
m; 22 July 1910; 1 damaged medusa, det. Kramp. —- ZMUC, as Merga lobianci, Thor station 147; 
31.583°N 19.033°E; southern Mediterranean; 25 m; 25 July 1910; 1 damaged medusa, det. 
Kramp. — ZMUC, as Merga lobianci, Thor station 184; 38.167°N 22.550°E; SE Mediterranean; 
10 m; 17 Aug. 1910; 2 medusae, det. Kramp. — ZMUC, Dana station 3809; 6.367°S 105.200°E; 
Indonesia; 0-100 m; 04 Sept. 1929; 2 medusae. - ZMUC; Vietnam, Nhatrang; 29 June 1936; 1 
medusa, det. Kramp. 


DraGNOSIS: Bell 4-11 mm high, with thick walls and dome-like apex, no apical 
process; four slightly folded lips, mesenteries very long; 8-12 long tentacles and 24-36 
rudimentary tentacles, all with ocelli. 


DESCRIPTION: Medusa umbrella bell-shaped, top relatively flat, walls thick, very 
thick at apex, no gastric peduncle; four radial canals, margins smooth, attached to 
manubrium by long mesenteries (about 1/, of total length); ring canal smooth. With 8- 
12 fully formed tentacles, their bases laterally flattened, without abaxial spurs, thinning 
quite abruptly into round tentacles after some distance; between pairs of fully formed 
tentacles normally three rudimentary marginal bulbs; all tentacle bases and rudi- 
mentary bulbs with a small ocellus. Manubrium 1/2 to 3/4 as long as bell cavity, cross- 
shaped in section; mouth margin slightly crenulated. Gonads large adradial bulges, 
surface smooth, with deep interradial fold but gonad may also cover interradial 
position. Colours: the manubrium and bulbs of the Atlantic forms are light pink to red- 
pink (Mayer, 1910); Menon (1932) observed a light-greenish tint. 

Polyp stage unknown. 


DIMENSIONS : Bell height up to 11 mm, usually less (4-6 mm). 


BIoLoGy: Occurs in tropical to subtropical seas, usually in less than 100 m 
depth, but some Mediterranean specimens (Lo Bianco, 1903) came from 900-1100 m 
depth. According to Mayer (1900), the medusa is common at the Tortugas throughout 
the summer. 


DISTRIBUTION: Mediterranean, Florida, Bahamas, Brazil, Red Sea, India, 
Nicobars, Vietnam, Sunda Strait, NE Australia, Fiji Islands, Papua New Guinea, Pacific 
coast of Mexico Pacific, Chile (Kramp, 1966, 1968; Goy, 1973; Schmidt, 1973a; 
Bouillon, 1980). Type locality: The first description was based on material from the 
Dry Tortugas (Florida) and Suva Harbour (Fiji Islands). 


356 P. SCHUCHERT 


et 


Fic. 69 


Merga violacea (Agassiz & Mayer, 1899); schematic drawing after several preserved 
Mediterranean specimens, scale bar 1 mm. 


REMARKS: In preserved material, the gonad can have some irregular folds. The 
animal depicted in Goy et al. (1991) has an apical process and short mesenteries and 
is unlikely M. violacea. Perhaps the legend is wrong and it should depict M. tergestina. 


Genus Annatiara Russell, 1940 
TYPE SPECIES: Tiaranna affinis Hartlaub, 1914 by monotypy. 


DIAGNosIs: Medusa bell without apical process, with exumbrellar nematocyst 
tracks. With four radial canals. Manubrium short and wide, cruciform through four 
long perradial lobes, mouth rim folded. Gonads on stomach lobes and interradial walls. 
Many hollow marginal tentacles, elongated conical bases, without well developed 
abaxial spurs; alternating with tentacles rudimentary tentacles or bulbs. With abaxial 
ocelli. 

REMARKS: The genus is regarded as monotypic (see below). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 357 


Annatiara affinis (Hartlaub, 1914) Fig. 70 


Tiaranna affinis Hartlaub, 1914: 269, figs 220-221. — Kramp, 1920: 6, pl. 1 fig. 1. — Kramp, 
1926: 68, pl. 1 figs 15-17, map 11. — Ranson, 1934: 436. 

Annatiara affinis. — Russell, 1940: 518. — Russell, 1953: 200, figs 101-103. — Kramp, 1955b: 
251, fig. 3. — Kramp, 1959a: 121, fig. 123. — Kramp, 1961: 96. — Kramp, 1968: 48, 
fig. 125. - O’Sullivan, 1982: 41, fig. 18, map 17. — Bleeker & van der Spoel, 1988: 231, 
figs 10-12. — Schuchert, 1996: 66, fig. 38. 

Annatiara lempersi Bleeker & van der Spoel, 1988: 249, fig. 13. new synonym 


MATERIAL EXAMINED: ZMUC, Thor Station 178, 48.067°N 12.667°W, eastern Atlantic, 
1000 m wire; 2 Sept. 1906; det. Kramp; 2 damaged medusae. — ZMUC, Dana Station 855, 
29.008°N 59.750°W, western Atlantic, 500 m wire; 9 June 1920; det. Kramp; 1 much damaged 
medusa. — ZMUC, 46.967°N 19.100°W, eastern Atlantic, 300 m wire; 21 July 1910; 1 much 
damaged and macerated specimen. 


DIAGNOSIS: As for genus. 


DESCRIPTION (Russell, 1953; own data): Umbrella bell-shaped, about as high as 
wide; Jelly relatively thick, thickest at top; no apical projection: exumbrella with 
numerous meridional nematocyst tracks originating from the tentacle bases. Velum 
narrow. Four straight radial canals issuing at ends of stomach lobes, narrow at first, 
widening somewhat distally with uneven margins, entering by wide orifice into narrow 
ring canal. No centripetal canals. Usually with 24-32 large hollow smooth tentacles (up 
to 44 observed), with elongated laterally compressed conical bases which clasp 
umbrella margin but do not form well-developed abaxial spurs. One very small or rudi- 
mentary tentacle between adjacent large tentacles. One small abaxial ocellus on bases 
of large and rudimentary tentacles. Stomach short, cross-shaped trough four long tri- 
angular perradial lobes, extending along more than half of the subumbrella meridian in 
fully grown animals; walls thin; mouth very large, cruciform, with folded margins. 
Gonads on irregular folds and pits on inter- and adradial walls of stomach, more de- 
veloped adradially, gonad pits bulge into stomach, pits sometimes in rows along stom- 
ach base. Colour of ocelli brown; gonads yellowish or colourless. Polyp stage un- 
known. 


VARIATION: Younger medusae have a square shaped stomach, which later 
develops its cruciform shape. Russell (1953) provides a table correlating bell diameter 
and tentacle number. 


DIMENSIONS: Mature medusa usually around 12 mm high and 14-15 mm wide 
(Kramp, 1961), maximal diameters 23 mm (Ranson, 1934). 


BioLoGY: Occurs in depths of several hundred meters, but apparently not a 
typical deep water species. 


DISTRIBUTION: Rare, usually off-shore in deeper waters. Atlantic Ocean between 
the Faroes and the equator; south-western Africa to South Africa; central Indian Ocean; 
Sri Lanka; east of Australia; New Zealand; Indonesia; Papua New Guinea; north- 
eastern Pacific (Ranson, 1934; Kramp, 1959a, 1961, 1965; Bouillon, 1980; Fraser, 
1973; Bleeker & van der Spoel, 1988; Schuchert, 1996; Wrobel & Mills, 1998). Type 
locality: southwest of Ireland, 48.483°N 13.917°W, 300 m wire out. 


REMARKS: The jelly of this rare species is rather delicate and specimens taken 
by plankton nets are almost invariably severely damaged. 


358 P. SCHUCHERT 


\ 
Li " 


Usd 


Fic. 70 


Annatiara affinis (Hartlaub, 1914). (A) A medusa from the Pacific, bell slightly contracted as in 
repulsion phase, drawn after photograph of living animal kindly provided by Dr D. Wrobel. 
(B) Interradial view of stomach with mature gonads, from Hartlaub (1914). 


Besides reporting details on Annatiara affinis, Bleeker & van der Spoel (1988) 
described also the new species Annatiara lempersi based on a single medusa from the 
mid North-Atlantic. They admit that their medusa was perhaps not fully mature, but 
they distinguished it from A. affinis on account of the lower tentacle number (24) and 
the stomach being square shaped instead of cruciform. It is quite clear that Annatiara 
lempersi is only based on a younger stage of Annatiara affinis. It matches rather 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 359 


perfectly the description of the type material in Hartlaub (1914), who also had animals 
with 24 tentacles and a square-shaped stomach. 

The polyp stage of A. affınis remains unknown. A potential candidate with a 
similar distribution in the north-eastern Atlantic is Hydrichthys sarcotretis. 


Genus Hydrichthys Fewkes, 1887 

TYPE SPECIES: Hydrichthys mirus Fewkes, 1887. 

SYNONYMS: ? Nudiclava Lloyd, 1907 [type species Nudiclava monocanthi Lloyd, 1907]. 
— Ichthyocodium Jungersen, 1913 [type species Ichthyocodium sarcotretis Jungersen, 1913]. 

DIAGNOSIS: Hydroid parasitic on fishes or parasitic copepods of fishes, hydro- 
rhiza reticular or forming a naked, encrusting plate; polyps dimorphic or: not, club- 
shaped or tubular, without tentacles, hydrocaulus indistinct and without perisarc 
covering; gonophores either growing at base of feeding zooids or on special gono- 
styles; gonostyles branched or not. Gonophores develop into medusa or fixed 
sporosacs. Newly liberated medusa with two opposite tentacles only. 


REMARKS: The first complete life cycle of a Hydrichthys hydroid was revealed 
by Larson (1982). He could identify the resulting medusa as Stomotoca pterophylla 
Haeckel, 1879. Because his hydroid showed little host specificity and many other 
Hydrichthys polyps are rather similar, Larson concluded that they will also produce 
medusae referable to the genus Stomotoca L. Agassiz, 1862. Therefore, he suggested 
to synonymize Hydrichthys and Stomotoca. This was contested by Arai (1989) and 
Boero & Bouillon (1989). The latter authors could show that the type species of 
Stomotoca — S. atra L. Agassiz, 1862 — had a polyp stage that was completely different 
from Hydrichthys (free living, with scattered tentacles). Moreover, the polyp of a 
tentatively identified Hydrichthys mirus Fewkes, 1887 — type species of the genus 
Hydrichthys — produced a Leuckartiara-like medusa (Boero er al., 1991). Because 
Stomotoca pterophylla and Stomotoca atra have different polyps, Boero et al. (1991) 
proposed the new genus Larsonia for S. pterophylla. 

More life-cycle information on various other Hydrichthys hydroids could show 
an even larger array of medusa morphologies and some might be referred to known 
medusa based genera. 

Because the Hydrichthys polyps produce medusae belonging to different 
genera, it must be assumed that the Hydrichthys-like polyp evolved several times 
convergently through a simplification and adaptation to the parasitic mode of life. Just 
like for many Bougainvillia hydroids, the Hydrichthys hydroids offer few morpho- 
logical traits to distinguish reliably separate species. The host specificity seems not to 
be very strict neither (Millard, 1975; Larson, 1982). Hydrichthys species identification 
is therefore difficult and assignments to nominal species should generally only be made 
when the material comes from approximately the same geographic region as the type 
specimen. It remains thus not beyond any doubt that the medusa of Hydrichthys mirus 
Fewkes, 1887 corresponds to the medusa described by Boero er al. (1991) based on a 
colony from Papua New Guinea, whereas H. mirus was originally described from New 
England. Further clarification of the genus depends on the rearing and identification of 
mature medusae of each nominal Hydrichthys species based on material from the 
original biogeographic region. 


360 P. SCHUCHERT 


Following Arai (1989), the morphological differences of Hydrichthys 
sarcotretis (Jungersen, 1913) and H. mirus are considered not sufficient to warrant a 
generic separation. Nudiclava Lloyd, 1907 was based on a hydroid which fixed 
sporosacs. The affinities of this genus are thus not clear, but it could be a synonym of 
Hydrichthys. 


KEY TO THE EUROPEAN HYDRICHTHYS SPECIES : 


la on copepods parasitizing fish, medusa buds not on hydrorhiza........ H. sarcotretis 
lb on fins of fish, medusa buds on hydranths and hydrorhiza........... H. cyclothonis 
Hydrichthys sarcotretis (Jungersen, 1913) Fig. 71 


Ichthyocodium sarcotretis Jungersen, 1913a: 1, figs. 5-6, pls. 1. figs 1-6. — Jungersen, 1913b: 
211.—Kramp, 1921: 13, figs 7-8.— Jones, 1966: 173, figs 1-2.— Templeman, 1973: 1355, 
figs 2-5. 
Hydrichthys sarcotretis. — Arai, 1998: 94, figs 1-2. 
MATERIAL EXAMINED: ZMUC, holotype colony; loc. SW of Ireland; coll. 1906. - ZMUC, 
W Greenland; 64.010°N 55.300°W; 8 May 1909; det. W. Jungersen (material not part of 
Jungersen, 1913a but mentioned in Jones, 1966). —- ZMUC; E Greenland; 60.933°N; 45.766°W; 
550 m wire; 22 August 1953; det. P. Kramp. 
DIAGNOSIS: Hydrichthys growing on copepod parasites of deep-sea fishes, 
polyps not dimorphic, medusa buds not on hydrorhiza. 


DESCRIPTION (Jungersen, 1913a; own observations): Hydroid growing on para- 
sitic copepods of fish. Stolons thin and not well visible, without perisarc, forming a 
tight meshwork and often coalescing into a plate-like hydrorhiza. Polyps dense, naked, 
non-reproductive polyps club-shaped but form variable, usually with an indistinctly 
demarcated pedicel widening distally into hydranth body, subterminally a slight neck 
and thus delimiting a head-like hypostome, contraction can obliterate this regiona- 
lization, mouth region without or with very few nematocysts, no tentacles. Medusa 
buds develop at lower part of normal hydranths (pedicels), hydranths can get gradually 
atrophied to mere stumps with ongoing budding, buds in several whorls (3-4), up to 20 
buds can be present. 

Advanced medusa buds are elongate (nearly two times longer than wide); two 
large opposite perradial bulbs, tapering into a short tentacle, abaxial fold in bulbs, other 
pair of bulbs small or not yet developed; many exumbrellar nematocysts; no sex cells 
visible in medusa buds. 


DIMENSIONS (Jungersen, 1913a; Jones, 1966; own data): Non-reproductive 
polyps 1-8 mm, diameter 0.05-0.2 mm; advanced medusa buds 0.4-0.84 mm long. 


ADDITIONAL DATA: Some hydranths can be branched, the gastrodermis has 6-11 
(usually 8) longitudinal ridges (Templeman, 1975). 


BIoLOGY: This hydroid lives on copepods (Sarcotretes scopeli, Sphyrion lumpi) 
parasitizing different fish species (Benthosema glaciale, Sebastes mentella) 
(Jungersen, 1913a; Jones, 1966; Templeman, 1973). The host fish occur in depths of 
several hundred meters, usually over very deep water. Jungersen (1913a) collected the 
hydroid in depths of 100-450 m. 

Microscopic examination of the stomach content showed that the hydroid likely 
feeds on blood and tissues of the host fish (Jones, 1966). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 361 


Fic. 71 


Hydrichthys sarcotretis (Jungersen, 1913). (A) Benthosema glaciale (Reinhardt) with the para- 
sitic copepod Sarcotretes scopeli (Jungersen) (arrow) bearing the hydroid colony; about two 
times magnified. (B) Schematic drawing of hydroid colony (polyp density reduced) with non- 
reproductive hydranth (left) and two hydranths producing medusae, hydranth at right much 
reduced or lost and with a medusa ready to be released, scale bar 0.2 mm. A, from Jungersen 
(1913); B, after preserved material. 


DISTRIBUTION: Subarctic to boreal Atlantic, recorded from The Faroes to 
southwest of Ireland (Jungersen, 1913a); between Greenland and Iceland (Jungersen, 
1913a, 1913b; Jones, 1966); western Greenland (Jungersen, 1913b; this study); 
Atlantic coast of Canada (Templeman, 1973). Type locality: southwest of Ireland. 


REMARKS: The publication date of Jungersen’s work is usually given as 1911, 
but according to the volume cover it was published in 1913. 

Kramp (1921) re-examined Jungersen’s material and he also made histological 
sections. He doubted that the gonophores are liberated as free medusae and he inter- 
preted the infolded tentacles in the subumbrella as thickened gastrodermal tissue. 
Having also examined Jungersen’s and additional material, I think that Kramp was 
wrong, a conclusion also put forward by Damas (1934) and Jones (1966). There were 
clearly some medusa buds with everted tentacles in the examined material (Fig. 71B). 


362 P. SCHUCHERT 


The adult medusa of Hydrichthys sarcotretis (Jungersen, 1913) is unknown. The 
distribution of this species overlaps with two other congeners, H. mirus and H. cyclo- 
thonis. Hydrichthys mirus Fewkes, 1887 was originally described as parasitizing a 
coastal fish of New England. It grows directly on the fish and differs also morpho- 
logically as it has dimorphic polyps. The gonozooids have many side branches bearing 
the medusa buds. Hydrichthys cyclothonis Damas, 1934 occurs immediately south of 
H. sarcotretis, ranging from the Gulf of Biscay to the Morocco and Madeira. It grows 
directly on the fins of the host. Apart from the host organism, the morphological 
differences are very not pronounced: the basal plate digesting the host tissues has a 
wide cavity; the medusa buds occur on the hydranth bases, the basal plate, and on 
blastostyles; the nematocysts on the medusa bud are evenly distributed. The first two 
differences could be linked to the different substrate and the blastostyles seem to be 
reduced hydranths in this case too. The nematocyst distribution on the medusa bud is 
variable in H. sarcotretis (Jones, 1966). As at least some Hydrichthys species can grow 
on copepods and fish simultaneously (McCormick et al., 1967), Hydrichthys cyclo- 
thonis is thus perhaps a form of H. sarcotretis (Jungersen, 1913). 


Hydrichthys cyclothonis Damas, 1934 Fig. 72 
Hydrichthys cyclothonis Damas, 1934: 1, figs 1-4. 


DIAGNOSIS: Hydrichthys growing on fish of the genus Cyclothone, medusa buds 
at base of polyps and on hydrorhiza. 


DESCRIPTION (Damas, 1934): Hydroids growing on epidermis of fish of the 
genus Cyclothone. Hydrorhiza plate-like, with large gastrodermal cavity, without 
perisarc, without processes infiltrating host tissue («haustoria»). Polyps dense, non- 
reproductive polyps club-shaped, widening distally, without distinct pedicel, without 
tentacles, their epidermis with few nematocysts only, mostly around mouth opening. 
Medusa buds develop either at base (pedicels) of feeding zooids, or from hydrorhiza, 
or from blastostyle-like polyps (likely reduced hydranths); numerous medusa buds 
give a grape-like appearance to the colony. 

Advanced medusa buds elongate (nearly two times longer than wide); with two 
large opposite perradial tentacle-bulbs; no sex cells visible; nematocysts of exumbrella 
evenly distributed. 


DIMENSIONS (Damas, 1934): Non-reproductive polyps 0.6 mm high and 0.2 mm 
wide (preserved material); advanced medusa buds 0.64 mm long and 0.25 mm wide. 


BIOLOGY: Grows on fins of Cyclothone species. It digests and destroys the 
tissue, it is thus a true parasite. Damas (1934) caught his fish with nets using cable 
lengths of 300-2120 m. According to Froese & Pauly (2006), Cyclothone alba has a 
depth range of 300-600 m. 


DISTRIBUTION: Gulf of Biscay; off Portugal, between Morocco and Madeira, 
Azores (Damas, 1934) Type locality: Not specified. 


REMARKS: For the differences to H. sarcotretis see under remarks for this 
species. Damas (1934) gives as host species Cyclothone signata (Garman). This 
species is currently seen as restricted to the Pacific Ocean (Froese & Pauly, 2006), but 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 363 


Fic. 72 


Hydrichthys cyclothonis Damas, 1934; figures from Damas (1934). (A) Colony on ventral fin, 
the black dots are the luminescent organs of the fish, scale bar 1 mm. (B) Advanced medusa bud, 
scale bar 0.2 mm. 


was earlier regarded as a cosmopolitan species. It is likely that Damas’s fish belonged 
to the very similar Cyclothone alba Brauer, a nominal species initially described as 
subspecies of C. signata. 

McCormick et al. (1967) examined more than 2000 specimens of Cyclothone 
originating from the north-eastern Pacific, among them C. signata, but they found no 
parasitic hydroids. They found them only on other genera and also on parasitic cope- 
pods. In a few cases the Hydrichthys polyps grew on both the parasitic copepod and 
the fish. 


Genus Octotiara Kramp, 1953 
TYPE SPECIES: Octotiara russelli Kramp, 1953 by monotypy. 


DIAGNOSIS: Pandeid medusa with eight simple radial canals in adults, manubri- 
um reaching far beyond velum level, with eight transversely folded gonads; without 
mesenteries, no ocelli, eight or more tentacles, with or without gastric peduncle. 

Hydroid on bryozoans, colony stolonal, hydrocaulus covered by thin perisarc, 
hydranth with a single whorl of filiform tentacles, medusa buds on stolons. 


REMARKS: The genus is currently monotypic. 


Octotiara russelli Kramp, 1953 Fig. 73 


Octotiara russelli Kramp, 1953: 266, pl. 1 figs 1-3. — Kramp, 1961: 111.-— Kramp, 1965: 38. — 
Kramp, 1968: 44, fig. 114. — Boero & Bouillon, 1989: 1, figs 1-3, 4A. — Bouillon et al., 
2004: 73, fig. 42F-H. 

Octotiara violacea Kramp, 1959b: 234, fig. 8a-c. — Kramp, 1965: 38, synonym. 


MATERIAL EXAMINED: MNHN, No 1645, coll. April 1954; Bay of Villefranche-sur-Mer, 
Ligurian Sea; material of Goy (1973), 1 damaged infertile medusa, bell diameter ca. 2 mm. 


DIAGNOSIS: See genus diagnosis. 


DESCRIPTION (Russell, 1968; Boero & Bouillon, 1989): Hydroid colonial, 
growing on specific cheilostomate bryozoans; colony stolonal, hydrorhiza tubular, 


364 P. SCHUCHERT 


Fic. 73 


Octotiara russelli Kramp, 1953. (A) The bryozoan Steginoporella mandibulata with the hydroid 
of Octotiara russelli, one medusa bud is present. (B) Higher magnification of the hydroid. 
(C) Newly released medusa. (D) Fifteen-day-old medusa, the second set of radial canals has not 
yet reached the ring canal. (E) Mature medusa. A-D, from Boero & Bouillon (1989); E, from 
Kramp (1968). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 365 


embedded in skeleton of host, hydranths arising between bryozoan zoecia; hydrocaulus 
short, covered by thin perisarc; hydranth with a single whorl of 3-4 filiform tentacles, 
rounded hypostome, reddish body. Medusa buds arise singly from stolons. 

Newly liberated medusa roundish to quadratic, exumbrella green; small apical 
projection present in some individuals; four radial canals only. Two long opposite mar- 
ginal tentacles originating from triangular-elongate perradial tentacular bulbs, two 
small atentaculate perradial bulbs; four smaller interradial bulbs; all bulbs reddish. 
Manubrium reddish, cylindrical, reaching 2/3 of height of subumbrellar cavity, mouth 
with four simple lips. 

Umbrella of mature medusa flatter than hemisphere, sometimes with a large 
and broad gastric peduncle; eight radial canals fairly narrow, with smooth edges, in 
younger individuals only the four perradial canals may reach the ring canal, interradial 
ones shorter; circular canal narrow; velum narrow. Usually eight large tentacles but 
occasionally up to 32, additionally 8-64 rudimentary bulbs or tentacles; nematocysts of 
tentacles evenly distributed. Manubrium long, reaching far beyond velum level, along 
its entire length eight deep longitudinal furrows; mouth part and stomach part approxi- 
mately same length, mouth part with eight sharp edges terminating in eight pointed 
lips. Gonads usually beyond level of velum along each of the eight perradial edges of 
the stomach, each with 6-10 deep irregular transverse folds. Nematocysts: microbasic 
euryteles. In living specimens the wall of the stomach between the gonads have a dark, 
almost black violet colour. 


DIMENSIONS: Hydranths about 0.5 mm high; newly liberated medusa 0.5- 
0.7 mm; diameter of mature medusa 7-11 mm. 


BioLocy: The hydroid was found in 10-20 m depth on the Indo-Pacific 
cheilostomate bryozoan Steginoporella mandibulata Harmer. This bryozoan genus is 
not known to occur in the Mediterranean or Atlantic (Costello et al., 2001). If the polyp 
is present in the Mediterranean, then it likely occurs on other bryozoans species. Boero 
& Bouillon (1989) found that the medusa was abundant near the water surface at 
Wuvulu Island, Papua New Guinea. Kramp (1953, 1965) collected it from near the 
surface to more than 400 m depth. 


DISTRIBUTION: Great Barrier Reef (Kramp, 1953); Sri Lanka (Kramp, 1959b); 
Malayan Archipelago and east coast of Africa (Kramp, 1965); Papua New Guinea 
(Bouillon 1980; Boero & Bouillon, 1989); Mediterranean (Goy, 1973). Type locality: 
Papuan Pass, Great Barrier Reef, Australia, depth more than 400 m. 


REMARKS: The only European record of this species was published by Goy 
(1973). She found one mutilated and infertile medusa in the Ligurian Sea 
(Mediterranean) which she attributed to O. violacea. This name has been synonymized 
with O. russelli by Kramp (1965). Goy’s specimen could be re-examined for this study. 
Although the identification may be correct, the state of preservation precludes a 
reliable identification. The presence of this species in the Mediterranean is therefore 
somewhat uncertain. 


Genus Codonorchis Haeckel, 1879 
TYPE SPECIES: Codonorchis octaedrus Haeckel, 1879 by monotypy. 


366 P. SCHUCHERT 


DIAGNOSIS: Pandeidae medusa with only two opposite tentacles; no manubrial 
peduncle; no mesenteries, with ocelli. The gonad are composed of four large flat inter- 
radial leaves that extend from the stomach surface onto the subumbrella, their margins 
are adnate to the radial canals, thus forming a bell-like structure within the subum- 
brella. 


REMARKS: This is a problematic genus based on single specimen of which there 
is no illustration available. Haeckel promised to provide an illustration in a subsequent 
paper, but he never did so. The genus is characterized by a unique form of the gonad. 
This was, however, not made very clear in the synopsis of Kramp (1961). Haeckel 
(1879) gives more details on this genus (translated from German, my additions in 
brackets): «I establish the genus Codonorchis for a peculiar Tiaridae which differs 
from all other genera of this group by its peculiar formation of the gonads. The gonads 
form four large interradial lappets which do not only cover most of the four sides of 
the tetraedric stomach, but they are continued onto the subumbrella where they cover 
its upper half; the thickened margins of the gonads are adnate to the radial canals and 
the gonads occupy completely the space between the proximal [upper] halves of the 
radial canals. By this, the four gonads form a bell-like cavity within which the stomach 
dangles. This peculiar formation arose probably when the eight gonadal ridges, which 
in turn originated through a cleavage of the four original perradial bulges, grew from 
the stomach base along the margins of the radial canals and got a connection through 
interradial lamellae.» 

The name Codonorchis is composed of the Greek words for bell and gonad. The 
interradial leaf-like gonads that extend halfway to the subumbrella are unique and 
should render the genus very distinct, if it is not based on a misinterpretation. The only 
Pandeid species that shows some resemblance to this species is Amphinema turrida 
(Mayer, 1900) (see Fig. 48). Also in this species the gonads extend to the subumbrella, 
but in this species they are adradial to perradial and not interradial. Picard (1960: 336) 
thought that C. octaedrus could actually be A. turrida. Also Bigelow (1909: 200) and 
Hartlaub (1914: 264) thought that Codonorchis octaedrus is related to A. turrida. 

For more problems associated with this genus, see the remarks under C. octae- 
drus. 


Codonorchis octaedrus Haeckel, 1879 
Codonorchis octaedrus Haeckel, 1879: 51. — Picard, 1960: 336. — Kramp, 1961: 98. 
Not Codonorchis octaedrus. — Boero et al., 1997: 359, figs 1-3. 

DESCRIPTION (Haeckel, 1879): Umbrella highly arched, shape quite octaedric 
through longitudinal exumbrellar perradial edges and one equatorial edge, with high 
cylindrical apical process, umbrella almost twice as high as wide. Two opposite 
tentacles, much longer than umbrella, conical bulbs; along bell margin ten bulbs, all 
bulbs with ocelli. Manubrium tetraedric, attached with broad quadratic basis, spanning 
half of the subumbrellar height; mouth-lips simple, short. Gonads four interradial 
lappets which cover the upper half of the subumbrella. Colours: stomach, gonad and 
tentacles yellowish; ocelli brown-red. 


DIMENSIONS: Diameter of bell 2.5 mm, bell-height 4 mm. 
DISTRIBUTION: Type locality only, Croisic (Brittany, France), depth 20 m. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 367 


REMARKS: Although its peculiar gonads and the ocelli should render this species 
distinct, it has never been found again. There exists very likely no type material. It must 
be regarded as a doubtful species 

Although the medusae of the region of the type locality — close to the mouth of 
the Loire — have not been investigated ever since, it is quite unlikely that C. octaedrus 
should be confined to this region. As the medusae of the nearby coasts are otherwise 
relatively well investigated, it is difficult to imagine why it has never been seen again. 
It could well be that Haeckel misinterpreted his specimens or he had shrunken and 
damaged material. As Haeckel repeatedly mistook some diffuse pigments for ocelli, it 
could also be that his medusa did not have true ocelli. Perhaps C. octaedrus was a well 
grown A. rugosum as shown in Fig. 47. Some authors linked O. octaedrus to 
Amphinma turrida (see above); Kramp (1961) considered it a doubtful species. 

Recently, Boero er al. (1997) thought that they have found this species again. 
They described adult medusae reared from a hydroid collected in the Ionian Sea. Their 
medusa shares several features with O. octaedrus, like the bell-shape, the apical 
process, tentacle number, the ocelli, and the bulb rudiments. However, it lacks the 
crucial feature of the genus Codonorchis, the gonad that extends to the subumbrella. I 
therefore hesitate to accept that this was O. octaedrus and consider it to represent a 
new, undescribed Amphinema species. In accordance with Dr. F. Boero and following 
his suggestion, it is therefore here described as the new species Amphinema bouilloni. 
The details are given under the Amphinema species. 

Boero et al. (1997) also designated neotype material for C. octaedrus. This 
designation can be regarded as invalid as it does not met the provision of article 75.3.5 
of the ICZN («evidence that the neotype is consistent with what is known of the former 
name-bearing type from the original description and from other sources...»). It might 
nevertheless be necessary to have a formal ruling of the International Commission on 
Zoological Nomenclature. 


PANDEIDAE INCERTA SEDIS: 


Genus Perigonella Stechow, 1921 
TYPE SPECIES: Perigonimus sulfureus Chun, 1889, by monotypy. 


DiaGNOSIS: Hydroid on shells of pelagic gastropods; colony stolonal, stolons 
tubular, hydranth with very short pedicel, one whorl of very short tentacles. Perhaps 
also dactylozooids. Gonophores borne on hydrorhiza, with pedicels, giving rise to free 
medusae. Advanced buds with four perradial tentacles, manubrium simple. 


REMARKS: According to Stechow (1921), this genus belongs to the 
Hydractiniidae. This is quite unlikely and following Bouillon (1985) it is placed among 
the Pandeidae. It shows many similarities with other Pandeidae, e. g. Pandea. 


Perigonella sulfurea (Chun, 1889) Fig. 74 


Perigonimus sulfureus Chun, 1889: 524. — Steche, 1907: 30, figs. 1-2. 
Perigonella sulfurea. — Stechow, 1921: 250. — Stechow, 1923a: 57. — Rees, 1956a: 344. 


DIAGNOSIS: See genus diagnosis. 


368 P. SCHUCHERT 


FIG. 74 


Perigonella sulfurea (Chun, 1889). (A) Feeding zooid, the constriction in the middle of the 
hydranth body is exceptional; the tentacles are very short, figure from Steche (1907). 
(B) Hydrorhiza with medusa buds, from Steche (1907) after drawings of C. Chun. 


DESCRIPTION: Hydroid colony stolonal, growing on shells of the pteropod 
Cavolinia tridentata (Forsskäl, 1775), hydrorhiza formed by reticulate stolons, covered 
by perisarc. Hydranths thick club-shaped, hypostome short, surrounded by 6-9 thick 
and very short tentacles; hydranth tapers rapidly near base into a pedicel with a hinge- 
joint, likely covered by perisarc. Gonophores are free medusae, developing singly from 
the stolons, pedicel a variable length present. Most advanced medusa buds known with 
four long tentacles and large conical bulbs, exumbrella with nematocysts, manubrium 
without oral tentacles. Colours: Gastrodermis of polyp and manubrium of medusa 
sulphur-yellow in living colonies. 


OTHER DATA: Steche (1907) observed an unusual hydranth with a horizontal 
constriction (Fig. 74A). 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 369 


BioLoGY: This species occurs on shells of Cavolinia tridentata (Forsskäl, 
1775), the original identification of Chun (1889) was corrected by Steche (1907). 
Steche (1907) also mentions the stomach content: molluscan eggs. They are most 
likely the eggs of the host. 


DISTRIBUTION: Only known from the Canary Islands (type locality). 


REMARKS: No type material of P. sulfurea could be located. It is perhaps lost, 
although Steche (1907) could re-examine it. 

This species has perhaps also dactylozooids. They were mentioned by Steche 
(1907) as «conical processes that represent modified zooids with a defensive function». 
These were interpreted by Stechow (1923a) as spines, but more likely they correspond 
to what Kramp (1921) identified as incipient gonophores in Kinetocodium danae, but 
later recognized as dactylozooids. The account of Steche (1907) is somewhat ambi- 
guous concerning the perisarc covering of the hydrocaulus, but it seems that there is 
perisarc on the pedicel. He mentions that the perisarc of the pedicel forms a 
constriction that serves as a hinge-joint. 

The hydranth tentacles were short and knob-like, this also while the colony was 
still alive. Although Steche (1907) interpreted this as the natural condition, it could also 
be a reduction due to mechanical and thermal stress. 

Kramp (1921) thought that this species is perhaps congeneric with his 
Kinetocodium danae. As the latter species produces a medusa with only three tentacles 
this seems not very likely and both genera should be retained until more is known on 
the mature medusae. 

In the absence of more life-cycle information, this species is difficult to classify. 
As the medusa has apparently no oral tentacles, it is best placed among the Pandeidae 
incerta sedis. 


Genus Kinetocodium Kramp, 1921 
TYPE SPECIES: Kinetocodium danae Kramp, 1921. 


DiaGNOSIS: Hydroid colony stolonal, living on pelagic gastropods; polyps poly- 
morphic with gastrozooids and dactylozooids; gastrozooids sessile, simple, with few 
short tentacles below rounded hypostome; dactylozooids filiform, nematocysts evenly 
distributed; gonophores on stolons, with pedicels, liberated as free medusae. 

Juvenile medusae with flask-shaped manubrium, mouth with oral nematocyst 
ring; four radial canals; three bulbs and three marginal tentacles, one perradial bulb 
rudimentary and without tentacle. 


Kinetocodium danae Kramp, 1921 Fig. 75 


Kinetocodium danae Kramp, 1921: 1, figs 1-8, pl. 1. - Kramp, 1957b: 47, figs 1-5. — Lalli & 
Gilmer, 1989: Fig. 8. 


MATERIAL EXAMINED: Holotype, ZMUC, polyps on Diacra trispinosa (Lesueur). 
DIAGNOSIS: See under genus. 


DEscRIPTION (Kramp, 1921, 1957b; own data): Hydroid growing on shells of 
the pelagic gastropod Diacra trispinosa (Lesueur). Stolons reticulate, flattened tubes, 
covered by very thin perisarc, stolons widened where bearing hydranths. Polyps 


370 P. SCHUCHERT 


FIG. 75 


Kinetocodium danae Kramp, 1921; modified from Kramp (1921, 1957), for dimensions see text. 
(A) Colony on the pteropod Diacra trispinosa (Lesueur). (B) Single hydranth, note sharp bend 
near base and short tentacles, which are reportedly a natural feature of this species. (C) Stolon 
with two medusa buds and a dactylozooids. (D) Newly released medusa, note that there are only 
three tentacles. 


dimorphic, gastrozooids and dactylozooids present. Gastrozooids naked, without 
distinct pedicels, long and slender, cylindrical, widened base, often sharply bend 
immediately above base, some polyps nevertheless straight; hypostome large, dome- 
shaped, below it one whorl of up to seven very short oval tentacles, usually four 


a 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 371 


tentacles, some hydranths without tentacles. Dactylozooids about as high as 
gonophores, filiform and very thin, nematocysts evenly distributed, dactylozooids do 
not coil but can contract. Gonophores develop from stolons on pedicels of variable 
length, ovoid, surrounded by a delicate perisarc, developing into free medusae. Colour: 
gastrodermis of hydranths brownish-orange. 

Newly liberated medusa bell-shaped, slightly higher than wide, jelly very thin, 
thickness uniform, exumbrella with nematocysts. Radial canals and ring canal narrow, 
velum well developed; only three perradial bulbs developed and bearing a tentacle, 
fourth rudimentary and without tentacle; tentacle bulbs broad, surrounded by a thick 
transverse band of nematocysts, broad on adaxial, narrow on abaxial side, without 
ocelli; tentacles covered evenly and densely with nematocysts throughout their length, 
gastrodermis parenchymatic; two opposite tentacles of same length (five time the bell 
size when expanded ), third tentacle slightly shorter. Manubrium roundly flask-shaped 
and half as long as the bell cavity, or more elongated and reaching nearly to the velar 
opening, without mesenteries, radial canals enter at the top, mouth quadrangular, 
margin beset with nematocysts, not particularly concentrated perradially, no lips, no 
oral tentacles or oral arms. Colours: gastrodermis of manubrium brightly orange. 

Mature medusa unknown. 


DIMENSIONS (Kramp, 1921; 1957b): Polyps up to 3 mm high, 0.2 mm wide, 
stolons 40 um wide. Medusa buds without pedicels 0.4 mm high and 0.25 mm wide. 
Newly released medusa 0.5 mm high and 0.3-0.4 mm wide. 


OTHER DATA: The bends of polyps are accompanied by a unilateral thickening 
of the mesogloea and gastrodermis. More histological details are given in Kramp 
(1921, 1957b). 


BioLoGY: Kramp (1957b) thinks that the hydroid parasitizes on its host, but he 
could not provide solid evidence for this. 


DISTRIBUTION: A rare species known from the West Indies, mid-Atlantic, 
Azores, Portugal, Madagascar (Kramp, 1957b). Type locality: 20.65° 61.80°W, 300 m 
wire out, between the Bermudas and the West Indies. 


REMARKS: With its three-tentacled medusa Kinetocodium does not fit in any 
family of the Filifera. It may be that this is only a feature of the juvenile medusa and 
the adult has more tentacles. Kramp (1921, 1957b) though that Kinetocodium might be 
a synonym of with Perigonella. Because of the dactylozooids and the manubrium 
mouth with nematocysts, Kramp (1957b) associated the genus Kinetocodium with the 
Hydractiniidae. This seems unlikely as the medusa buds are on the stolons and not on 
the hydranths as in typical Hydractiniidae (Schuchert, 2001). The medusa does like- 
wise not correspond to a Hydractinia species. In order to allow a comparison and with 
Perigonella and Hydrichthys, Kinetocodium is here provisionally placed among the 
Pandeidae as incerta sedis, awaiting more information on its mature medusa. 
Admittedly, the tentacle bulbs with their nematocyst rings are atypical for the 
Pandeidae. The nematocysts could give some important cues to the affinities of 
Kinetocodium, but they could not be examined in the available material (holotype). 


372 P. SCHUCHERT 


ACKNOWLEDGEMENTS 


This study was made possible through a SYNTHESIS grant of the European 
Union that enabled me to examine hydrozoans of the Natural History Museum in 
London. Dr J. Galeron (IFREMER, France) generously provided some deep water 
samples. I also wish to express my sincere thanks to the museums of Copenhagen, 
Bergen, Bruxelles, Munich, Tervuren, and the Smithsonian Institution for providing 
valuable loans of material. Dr K. Raskoff, Dr D. Wrobel, and Dr H. Galea provided 
photographs of living medusae and their help was much appreciated. Many thanks are 
also due to Dr G. Plickert who sent me a culture of Leuckartiara octona. 

I am particularly grateful to Dr Anita Brinckmann-Voss who corrected and 
commented a first draft of this publication. Her comments helped to improve this paper 
significantly. The remaining errors, though, are solely my responsability. 


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Taiwan Strait with description of new species and new combinations. Acta Oceanologica 
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ZELICKMAN, E. A., GELFAND, V. I. & SHIFRIN, M. A. 1969. Growth, reproduction and nutrition of 
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ZELICKMAN, E. A. 1972. Distribution and ecology of the pelagic hydromedusae, siphonophores 
and ctenophores of the Barents Sea, based on perennial plankton collections. Marine 
Biology, Berlin 17: 256-264. 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 393 


INDEX brownei, Leuckartiara 331 
Calyptospadix 250 
abyssi, Leuckartiara 330 Campaniclava 299 
abyssi, Perigonimus 330 campanula, Catablema 345 
affinis, Annatiara 357 Catablema 344 
affinis, Stylocidaris 233 cavosa, Dinematella 306 
album, Rhizorhagium 266 cerulea, Calyptospadix 250 
alderi, Corynopsis 199 charcoti, Bougainvillia 235 
alderi, Podocoryne 199 charkoti, Bougainvillia 235 
allmanii, Bougainvillia 210 cirratus, Halitholus 341 
amoyensis, Bimeria 247 claparedei, Lizzia 289 
Amphinema 305 Clavopsella 267 
ampullacea, Oceania 333 Clavopsellidae 196 
ampullacea, Tiara pileata var. 300 cleodorae, Campaniclava 300 
Annatiara 356 cleodorae, Syncoryne 300 
antarcticus, Perigonimus 261 coccinea, Atractylis 262 
Aphrodite aculeata 325 cocciniea, Oceania 333 
apicata, Saphenia 306 Codonorchis 365 
apicata, Stomotoca 306 coeca, Bougainvillia britannica var. 208 
arborea, Bimeria 255 coeca, Tiarula 306 
arborea, Garveia 255 conferta, Dicoryne 241 
arenosa, Atractylis 264 confertum, Eudendrium 241 
arenosa, Wrightia 264 conica, Dianaea 300 
arenosum, Rhizorhagium 264 conica, Pandea 300 
Aselomaris 261 conifera, Tiara 345 
atlantica, Cytaeis 275 conifera, Tiaria 343 
Atractylidae 196 constricta, Oceania 333 
Atractylis 198 conybearei, Dicoryne 245 
aurantiaca, Bougainvillia 228 conybearei, Heterocordyle 245 
aurea, Hippocrene 210 conybeary, Heterocordyle 245 
autumnalis var. magna 201 corynopsis, Bimeria 247 
autumnalis, Bougainvillia 199 Corythamnium 250 
bacciferum, Eudendrium 251 cruciata, Hippocrene 235 
bella, Bougainvillia 208 crucifera, Hippocrene 235 
benedenii, Bougainvillia 199 cyclothonis, Hydrichthys 362 
Bimeria 247 Cytactis 274 
Bimeridae 196 Cytaeididae 274 
biscayana, Amphinema 317 Cytaeidium 274 
biscayana, Bimeria 317 Cytaeis 274 
bitentaculata, Atractylis 271 Cyteis 274 
bitentaculatus, Perigonimus 271 Cytheis 274 
blondina, Lizzia 289 danae, Kinetocodium 369 
Blumenbachii, Oceania 284 decorans, Perigonimus 273 
Blumenbachii, Rathkea 284 diadema, Oceania 306 
Blumenbachii, Rathkia 284 dibalia, Lizzia 233 
Bougainvilleae 196 dibalium, Thamnostoma 233 
Bougainvillia 198 dichotoma, Thamnitis 230 
Bougainvillii, Cyanea 223 Dicoryne 241 
Bougainvillii, Hippocrene 220 Dicorynidae 196 
Bougainvilliidae 196 digitale, Turris 333 
bouilloni, Amphinema 319 Dinema 321 
breviconis, Neoturris 338 dinema, Amphinema 306 
breviconis, Turris 338 dinema, Bougainvillia 206 
brevicornis, Leuckartiara 338 dinema, Oceania 306 
britannica, Bougainvillia 206 diplectanos, Bougainvillia 199 


britannica, Hippocrene 206 duodecilia, Medusa 210 


394 


elisabethae, Lizzia 296 
else-oswaldae, Stylactella 278 
enigmatica, Velkovrhia 271 
episcopalis, Oceania 323 
eurystoma, Catablema 345 
fasciculata, Koellikerina 230 
fasciculata, Melicerta 230 
flavida, Bougainvillia 199 
flexuosa, Dicoryne 241 
fragilis, Bimeria 257 
franciscana, Bimeria 253 
franciscana, Garveia 253 
fruticosa, Bougainvillia 199 
fulgurans, Dysmorphosa 289 
fusca, Manicella 247 
gaedii, Oceania 323 
galleri, Merga 351 
Garveia 250 

gelatinosus, Perigonimus 323 
gemmascens, Cubaster 289 
georginae, Perigonimus 272 
gibbesii, Nemopsis 235 
Gibbsi, Bougainvillia 199 
glaciale, Benthosema 360 
grata, Lizzia 284 

Gravelya 261 

grimaldii, Leuckartiara 329 
grisea, Garveia 257 

grisea, Pruvotella 257 
groenlandica, Garveia 262 
Halitholus 340 

herdmani, Cytaeis 275 
Heterocordyle 241 
heteronema, Nemopsis 210 
Hippocrene 198 
Hippocrenidae 196 
humilis, Bimeria 247 
hyalina, Circe 273 

hyalina, Lizzella 273 
Hydrichthys 359 

hystrix, Hermione 233 
Ichthyocodium 359 
inflatus, Perigonimus 273 
Japonica, Cytaeis 275 
Jonesii, Perigonimus 323 
Kinetocodium 369 
Koellikerina 230 

krampi, Amphinema 314 
lempersi, Annatiara 357 
Lesueurti, Oceania 333 
Leuckartiara 321 

linearis, Atractylis 208 
Lizusa 198 

Lizusidae 196 

Lizzia 288 

Lizzinae 282 


P. SCHUCHERT 


lobianci, Mergintha 355 

lumpi, Sphyrion 360 
macloviana, Bougainvillia 224 
maclovianus, Perigonimus 224 
macrogaster, Cytaeis 275 
Manicella 247 

maniculata, Bougainvillia 199 
Margelidae 196 

Margelis 198 

Margellium 284 

mediterranea, Bougainvillia 275 
megas, Perigonimus 253 
mentella, Sebastes 360 
Merga 347 

Mergintha 347 

Mertensii, Bougainvillia 220 
michaeli, Aselomaris 267 
michaeli, Clavopsella 267 
michaeli, Pachycordyle 267 
miniata, Atractylis 262 


minima, Bougainvillia ramosa var. 201 


minima, Cytaeis 297 

minima, Dysmorphosa 289 
minima, Podocoryne 297 
minima, Podocorynoides 297 
minuta, Dysmorphosa 289 
minuta, Lizzia 289 

minuta, Podocoryne 289, 293 
minutus, Perigonimus 323 
mitra, Nubiella 238 

Monidi, Bimeria 253 
monocanthi, Nudiclava 359 
multicilia, Bougainvillia 229 
multicilia, Lizusa 229 
muscoides, Bougainvillia 214 
muscus, Bougainvillia 199 
muscus, Perigonymus 199 
nana, Bimeria vestita f. 247 


nana, Bougainvillia ramosa var. 199 


nanellus, Perigonimus 245 
napolitanus, Perigonimus 323 
neglecta, Turris 321 
Nemopsidae 196 
Nemopsis 234 

Nemopsis bachei 235 
Neoturris 333 

nigritella, Bougainvillia 214 
Nigritina 274 

Nigritina atlantica 275 
nigritina, Cytaeis 275 
niobe, Bougainvillia 229 
nobilis, Leuckartiara 326 
nordgaardii, Margelis 214 
Nubiella 228 

Nudiclava 359 

nudus, Perigonimus 273 


THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 


nutans, Garveia 251 

nutans, Perigonimus 273 
obscura, Bougainvillia 199 
ocilia, Medusa 199 
octaedrus, Codonorchis 320, 366 
octocilia, Lizusa 199 
octocilia, Medusa 199 
octona, Geryonia 322 

octona, Leuckartiara 322 
octona, Paracytaeis 282 
octopunctata, Cytaeis 284 
octopunctata, Lizzia 284 
octopunctata, Rathkea 284 
octostyla, Dysmorphosa 295 
octostyla, Lizzia 295 
Octotiara 363 

Pachycordyle 267 
Pachycordylini 196 

palliata, Atractylis 323 
Pandea 299 

Pandeidae 298 

papalis, Tiaria 333 

papillata, Dorocidaris 233 
papua, Tiara 323 

Paracytaeis 282 

paradoxa, Bougainvillia 220 
Parawrightia 198,270 
pauper, Halitholus 343 
Perarella 278 

Perigonella 367 

Perigonimus 198 

pileata, Neoturris 333 
pileata, Oceania 321 

pileus, Oceania 333 

pisifera, Carybdea 333 
platygaster, Bougainvillia 226 
platygaster, Hippocrene 226 
Podocorynoides 297 
principis, Bougainvillia 210 
principis, Margelis 210 
propagulata, Cytaeis 280 
propagulata, Perarella 280 
Pruvotella 250 

pudicum, Eudendrium 266 
pugetensis, Perigonimus 323 
pusilla, Cytaeis 275 

pusilla, Pachycordyle 205, 206, 267 
pusillum, Eudendrium 323 
pyramidata, Bougainvillia 217 
pyramidata, Hippocrene 217 
quadritentaculatus, Atractylis 272 
radicans, Perigonimus vestitus f. 323 
ramosa, Atractylis 199 
ramosa, Bougainvillia 199 
ramosum, Eudendrium 199 
Rathkea 284 


Rathkeidae 282 

reesi, Merga 352 

repens, Atractylis 323 
reticulata, Tiara 300 
Rhizorhagium 261 

roseum, Rhizorhagium 262 
rubra, Merga 315 

rubra, Pandea 203 
rubrum, Amphinema 315 
rugosa, Stomotoca 309 
rugosum, Amphinema 309 
russelli, Octotiara 363 
russelli, Thamnostoma 230 
sarcotretis, Hydrichthys 360 
sarcotretis, Ichthyocodium 360 
sarsii, Perigonimus 262 
schneideri, Perarella 273, 278 
schneideri, Perigonimus 278 
scopeli, Sarcotretes 360 
sedecimacostata, Oceania 300 
serpens, Perigonimus 306 
sessile, Eudendrium 323 
shimiko, Lizzia 284 
Silhouetta 239 

simplex, Hippocrene 210 
simplex, Podocoryne 297 
slabberi, Dinema 323 
smaragdina, Tiara 323 
steinachi, Perigonimus 273 
stricta, Dicoryne 241 
sulfurea, Perigonella 367 
sulfureus, Perigonimus 367 


superciliaris, Bougainvillia 220 


superciliaris, Hippocrene 220 
tergestina, Merga 348 
tetrastyla, Cytaeis 275 
Thamnitis 214,233 
Thamnostoma 233 
Thamnostoma spec. 214 
Thamnostomidae 196 
Thieliana 267 

Tiara 333 

Tiarula 347 

titania, Saphenia 306 
tregoubovii, Merga 353 
triestina, Bougainvillia 199 
triloba, Pegantha 228 
tunicata, Bimeria 253 
turrida, Dissonema 312 
turrida, Amphinema 312 
Turris 333 

turrita, Oceania 323 
uvacarpa, Silhouetta 239 
Velkovrhia 270 
vesicarium, Catablema 345 
vestita, Bimeria 247 


395 


396 


vestitum, Eudendrium 247 
vestitus, Perigonimus 323 
violacea, Merga 355 
violacea, Octotiara 363 
viridula, Eirene 273 


P. SCHUCHERT 


vulgaris, Cytaeis 275 

Wrightia 261 

xantha, Bougainvillia 208 
yoldia-arcticae, Perigonimus 341 
zygonema, Margelis 208 


REVUE SUISSE DE ZOOLOGIE 114 (2): 397-415; juin 2007 


A new species of Lygosoma (Squamata: Sauria: Scincidae) from the 
Central Truong Son, Vietnam, with notes on its molecular 
phylogenetic position 


Thomas ZIEGLER!, Andreas SCHMITZ, Astrid HEIDRICH! 3, Ngoc Thanh VU & 

Quang Truong NGUYEN? 

! AG Zoologischer Garten Köln, Riehler Straße 173, D-50735 Köln, Germany. 
E-mail: tziegler@zoo-koeln.de 

2 Muséum d’histoire naturelle, Department of Herpetology and Ichthyology, C.P. 6434, 
CH-1211 Geneva 6, Switzerland. E-mail: andreas.schmitz@ville-ge.ch 

3 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 
D-53113 Bonn, Germany. E-mail: astrid_heidrich@web.de 

4 Centre for Natural Resources and Environmental Studies (CRES), Vietnam National 
University, Hanoi, University of Science, Faculty of Biology, Department of 
Vertebrate Zoology, Zoological Museum, 334 Nguyen Trai Str., Thanh Xuan, 
Hanoi, Vietnam. E-mail: vnthanh@fpt.vn 

5 Institute of Ecology and Biological Resources (IEBR), Vietnamese Academy of 
Science and Technology, 18 Hoang Quoc Viet St., Hanoi, Vietnam. 
E-mail: truong@iebr.vast.ac.vn 


A new species of Lygosoma (Squamata: Sauria: Scincidae) from the 
central Truong Son, Vietnam, with notes on its molecular phylogenetic 
position. - A new Lygosoma species is described from the Central Truong 
Son (Annamite mountain range) of Quang Binh Province, Vietnam. The 
description is based on a single female specimen, collected during the dry 
season in the karst forest of Phong Nha — Ke Bang National Park. The new 
Lygosoma differs from any other congener by the dorsal scales with pseudo- 
keels in combination with a pair of frontoparietals; a scaly lower eyelid; 
seven supralabials; seven infralabials; 32 midbody scale rows; 66 middorsal 
(paravertebral) scales; smooth ventral scales, arranged in 81 transverse 
rows; 108 smooth, not enlarged median subcaudal scales; the fourth toe with 
14 keeled subdigital lamellae; a reddish brown to brownish black dorsum 
and an orange-yellowish to greyish ventral side in life; as well as greyish 
black edged sutures of anterior supra- and infralabials. The new Lygosoma 
species is the third karst-adapted scincid species that has been described 
from Phong Nha - Ke Bang National Park since 2005. A first molecular 
positioning of the new species within the genus Lygosoma is given as well 
as a key to the Vietnamese Lygosoma species. 


Keywords: Sauria - Scincidae - Lygosoma boehmei sp. n. - taxonomy - 
phylogeny - Central Truong Son - Vietnam. 


Manuscript accepted 10.02.2007 


398 T. ZIEGLER ET AL. 


INTRODUCTION 


During recent herpetofaunal investigations in the Phong Nha — Ke Bang 
National Park, Central Truong Son (Annamite mountain range) of Quang Binh 
Province, in Central Vietnam (e.g., Ziegler & Herrmann, 2000; Ziegler & Le, 2005, 
2006; Ziegler et al., 2004, 2005, 2006) a remarkable scincid lizard was collected. At 
first glance, the specimen appeared with its apparently keeled dorsal scales to be an- 
other representative of Lygosoma carinatum, that was described a decade ago by 
Darevsky & Orlova (1996) based upon two specimens from the “Tau Nguen (sic)” 
(= Tay Nguyen) plateau in Kon Tum Province, Vietnam. However, as is pointed out in 
detail below, our investigations showed the dorsal keels to be pseudo-keels in fact. In 
addition, the single specimen found by us showed a distinctly differing pholidosis, i.e. 
larger body scales, resulting in distinctly lower midbody, middorsal, and ventral scale 
counts compared to L. carinatum, and differed as well from the remaining Lygosoma 
(and Riopa) species listed in Nguyen er al. (2005) for Vietnam. It must be noted that 
there exists quite some confusion concerning the assignment of species to the genera 
Lygosoma Hardwicke & Gray, 1827 and Riopa Gray, 1839, respectively. While Riopa 
was long considered as a valid genus (e.g., Smith, 1937; Mittleman, 1952), Greer 
(1977) carried out a careful morphological revision and as a result has synonymized 
the genus Riopa with Lygosoma. This author gave a missing “close ecological and mor- 
phological continuity among [the genus] own species and sharp ecological and 
morphological discontinuity from its near relatives” as the main reason to place Riopa 
(together with other proposed skink genera) into the synonymy of Lygosoma. But this 
assignment has not been unambiguously followed since. Some more recent authors 
such as Manthey & Grossmann (1997) again list specimens that bear supranasals under 
Riopa, while others either follow Greer (1977) and accept the synonymization of Riopa 
with Lygosoma or simply do not distinguish between both genera (e.g., Honda et al., 
2003). A solution to this problem cannot be provided in the framework of this paper 
and should be treated in a comprehensive phylogenetic approach, preferably based on 
modern molecular analyses. We preliminarily follow in this paper the definition of 
Greer (1977) and consider Riopa as a synonym of Lygosoma. As our specimen was 
neither assignable to any of the other scincid genera listed by Nguyen ef al. (2005) for 
Vietnam (Dasia, Emoia, Eumeces, Leptoseps, Lipinia, Eutropis [see Mausfeld et al., 
2002 and Mausfeld & Schmitz, 2003 on the split of Mabuya sensu lato and the 
resulting nomenclatural changes], Paralipinia, Scincella, Sphenomorphus, Tropido- 
phorus, and Vietnascincus), nor to any of the described Lygosoma (or “Riopa’) species 
from Vietnam or its neighbouring countries (Bourret, unpubl.; Manthey & Grossmann, 
1997: Darevsky & Orlov, 1994, 1997, 2005), we describe it herein as new. 


MOLECULAR METHODS 


To support our morphological results molecular data were collected to identify 
the phylogenetic position of the new species in a general framework of related species 
of the genus Lygosoma (including its supposed synonym Riopa). We sequenced a 
portion of the mitochondrial 16S rRNA gene and compared it with the following 
species, some of which are also present in the key to the Vietnamese species (see 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 399 


below): Lygosoma quadrupes (type species of the genus Lygosoma), L. albopunctatum, 
and Lygosoma sp. (India). Further, we included specimens from several localities of L. 
koratense (2), L. lineolatum (2; used as outgroups), and the widespread L. bowringii 
species-complex (3). For GenBank accession numbers see Table 1. 

DNA was extracted from the tissue samples using QuiAmp tissue extraction kits 
(Quiagen) or a modified Chelex-Protocol (Walsh et al., 1991; Schmitz, 2003). The 
primers 16sar-L (light chain; 5° - CGC CTG TTT ATC AAA AAC AT - 3°) and 16sbr- 
H (heavy chain; 5° - CCG GTC TGA ACT CAG ATC ACG T - 3°) of Palumbi er al. 
(1991) were used to amplify a section of the mitochondrial 16S ribosomal RNA gene. 
PCR cycling procedure followed Schmitz et al. (2005). PCR products were purified 
using Qiaquick purification kits (Qiagen). Sequences (including complimentary 
strands for assuring the accuracy of the sequences) were obtained using an automatic 
sequencer (ABI 377). The obtained sequences (lengths referring to the aligned 
sequences including gaps) comprised 548 bp. Sequences were aligned using ClustalX 
(Thompson et al., 1997; default parameters) and manually checked using the original 
chromatograph data in the program BioEdit (Hall, 1999). We used PAUP* 4.0b10 
(Swofford, 2002) to compute the uncorrected pairwise distances for all sequences. 

We performed maximum parsimony (MP), maximum likelihood (ML), 
Neighbor-joining (NJ) and Bayesian (PP) reconstructions. For ML and Bayesian ana- 
lysis parameters of the model were estimated from the data set using Modeltest 3.7 
(Posada & Crandall, 1998) and MrModeltest 2.2 (Nylander, 2005), respectively. The 
NJ-analysis used the uncorrected ‘p-distances’. Additionally, we used bootstrap ana- 
lyses with 2000 (MP and ML) and 20000 (NJ) pseudoreplicates to evaluate the relative 
branch support in phylogenetic analysis. For the MP analysis, we used the “heuristic 
search” with the “random addition” option of PAUP* (Swofford, 2002) with 10 repli- 
cates, using the TBR (tree bisection-reconnection) branch swapping option. All 
Bayesian analyses were performed with MrBayes, version 3.0b4 (Huelsenbeck & 
Ronquist, 2001). We ran two MCMC analyses for 106 generations each. The initial 
100000 (10%) trees were disregarded as “burn-in”. We consider probabilities of 95% 
or greater to be significantly supported. The exact parameters used for the Bayesian 
analyses followed those described in detail by Reeder (2003). 


RESULTS AND DISCUSSION 


Lygosoma boehmei sp.n. 

HOLOTYPE: Zoologisches Forschungsmuseum Alexander Koenig, ZFMK 86359: female 
(Figs 1-8), from the karst forest of Cha Noi, 350-400 m a.s.l., Phong Nha — Ke Bang National 
Park, Quang Binh Province, Vietnam; collected by Astrid Heidrich and Thomas Ziegler at the 
end of the dry season (21 June) 2006. 

ETYMOLOGY: We name this new species in honor of Professor Dr Wolfgang 
Böhme, vice director of the Zoological Research Museum Alexander Koenig in Bonn, 
head of the vertebrate section and curator for herpetology, in recognition of his out- 
standing contributions not only towards lizard systematics throughout the last three 
decades. 


DIAGNOSIS: The new species can be distinguished from any other Lygosoma / 
Riopa currently known by the following combination of characters: (1) Body elongate 


400 T. ZIEGLER ET AL. 


Fics 1-2 


(1) Female holotype of Lygosoma boehmei sp. n. (ZFMK 86359) in life. (2) Portrait of the 
holotype of Lygosoma boehmei sp. n. (ZFMK 86359) in life. 


(SVL 86.0 mm), the distance between axilla and groin being slightly more than 3 times 
the length of the forelimb; (2) forelimb and hindlimb short, pentadactyl, fingers and 
toes widely separated when adpressed, the distance between them corresponding to the 
length of the hindlimb; (3) dorsum reddish brown in life, brownish grey in preser- 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 401 


TABLE 1. List of samples used for genetic analysis (geographic origin, locality and GenBank ac- 
cession numbers). Acronyms are as follows: KUZ for Herpetological Collection of the 
Department of Zoology, Kyoto University, Japan; MHNG for Muséum d’histoire naturelle, 
Geneva, Switzerland; ZFMK for Zoologisches Forschungsmuseum Alexander Koenig, Bonn, 
Germany 


Species Geographic Locality Voucher Accession 
origin number 

L. albopunctatum India Near Ajur, ZFMK 73430 AY308262 
Peryar-Trivandrum 

L. boehmei n. sp. Vietnam Cha Noi, Phong-Nha ZFMK 86359 EF193650 
Ke Bang NP 

L. bowringii Indonesia Pondok Sari, ZFMK 78822 . AY308263 
Permuteran, Bali 

L. bowringü (I) Thailand Khao Chong KUZ 37884 AB028786 

L. bowringii (II) Thailand Tha Uthen MHNG 2679.72 EF193649 

L. koratense Malaysia locality unknown ZFMK 71715 AY308269 

L. koratense Thailand locality unknown KUZ 27358 AB028817 

L. lineolatum Myanmar Rakhine State, Gwo CAS 206647 AY308270 
Township 

L. lineolatum Myanmar Mandalay Div., Popa CAS 210669 AY308271 
Mountain Park 

L. quadrupes Thailand Bankok KUZ 40033 AB028818 

Lygosoma sp. India Ooty-Bandypur ZFMK 77814 AY308272 


vative, the surfaces of the limbs and tail being brownish black; ventral side yellowish 
beige to greyish below the tail in preservative, with some indistinct dark marbling; in 
life, chin and throat are light orange and the remaining ventral side of the body cream 
to light brownish, turning to greyish at the underside of the tail; sides of body and neck 
with irregular dark flecking; sutures of anterior supra- and infralabials edged by 
greyish black; (4) rostral separated from undivided frontonasal by supranasals (5) pre- 
frontals widely separated; (6) parietals forming a suture behind interparietal, no 
enlarged nuchals; (7) four supraoculars; bordered by six supraciliaries; posteriorly, the 
fourth supraocular is bordered by a small scale (postsupraocular); (8) lower eyelid 
scaly; (9) two loreals; (10) seven supralabials, fifth right below the eye; (11) seven 
infralabials; (12) postmental undivided; (13) first pair of chinshields in broad contact, 
bordered by six scales; (14) ear opening small, ovoid to roundish; (15) 32 midbody 
scale rows; (16) 66 middorsal (paravertebral) scales (from the posterior end of parietals 
to insertion of hindlimb); (17) dorsal and dorsolateral scales notched, appearing as 
three longitudinal (pseudo-)keels on neck, body, and anterior half of tail; (18) ventral 
scales smooth, arranged in 81 transverse rows from first gular row between third pair 
of chinshields to preanals; (19) six slightly enlarged preanals; (20) 108 median sub- 
caudal scales, not enlarged, smooth; (21) fourth toe with 14 keeled subdigital lamellae. 


DESCRIPTION OF HOLOTYPE: Female specimen, for measurements see Table 2. 
Body elongate, the distance between axilla (end of forelimb) and groin (insertion of 
hindlimb) slightly more than 3 times the length of the forelimb. Forelimb and hindlimb 
short, pentadactyl. Fingers and toes widely separated when adpressed, the distance be- 
tween them corresponding to the length of the hindlimb. Head only slightly set-off 
from neck, snout rounded. Rostral wider than high, visible from above. A pair of 


402 T. ZIEGLER ET AL. 


TABLE 2. Measurements (taken by a caliper, in mm) and some scalation features of the female 
holotype of Lygosoma boehmei sp. n. compared to the female holotype of Lygosoma carinatum 
(ZIN 20482) from Kannack, Kon Tum province, Vietnam, studied by us (* = data from Darevsky 
& Orlova, 1996). SVL: snout-vent length (from snout tip to cloaca); TaL: tail length (from cloa- 
ca to tail tip); TL: total length; HL: head length (distance from snout tip to posterior margin of 
interparietal); HW: maximum head width; SL: maximum snout length (from tip to anterior mar- 
gin of the eye); ET: maximum eye to tympanum length (from hind margin of the eye to anterior 
border of tympanum); TW: maximum tympanum width; SFl: maximum snout to forelimb 
length; AG: maximum axilla to groin length; FIL: maximum forelimb length (from body inser- 
tion to beginning of claw of fourth finger); HIL: maximum hindlimb length (from body insertion 
to claw of fourth toe); IL: Infralabials; MB: midbody scale rows; MD: middorsal scale rows 
(from posterior end of parietals to insertion of hindlimb / to centre of thigh / and to posterior 
hindlimb); V: transverse rows of ventral scales (from first gular row subsequent to first pair of 
chinshields to preanals); MSS: median subcaudal scales; LAT: Lamellae beneath fourth toe. 


Lygosoma boehmei sp. n. Lygosoma carinatum 
ZFMK 86359 ZIN 20482 

SVL 86.0 TAO 
TaL 91.0 PTL) 
NL, 177.0 *148.0 
HL 123 10.0 
HW 10.5 8.9 
SL 6.1 SI 
ET 64 5.6 
TW 3) 1.1 
SFI 294 2591: 
AG 47.0 40.3 
FIL 14.7 122 
HIL 19.0 15.0 
IL 7} 6-7 
MB 32 38 
MD 66 / 70 / 72 80 / 82/85 
V 81 92 
MSS 108 115 
LAT 14 16 


supranasals, somewhat wider than long, forming a distinct median suture, touching 
nasals and anterior loreal laterally. Prefrontals widely separated, touching both loreals 
laterally. Frontonasal wider than long, in contact with supranasals, anterior loreals, pre- 
frontals, and frontal. Frontal large, longer than its distance to tip of snout, narrowing 
posteriorly, in contact with frontonasal, prefrontals, first supraciliary, first and second 
supraoculars, and frontoparietals. A pair of frontoparietals, about as wide as long. 
Parietals forming a suture behind the interparietal, that is longer than wide, smaller 
than frontal and narrowing posteriorly. Small transparent spot on interparietal, show- 
ing location of parietal foramen. No distinctly enlarged nuchal scales. Nostril piercing 
nasal, that is in contact with rostral, first supralabial, anterior loreal, and supranasal. 
Two loreals, anterior the higher (bordering nasal, first and second supralabials, 
posterior loreal, prefrontal, frontonasal, and supranasal), posterior the longer. The 
posterior loreal is in contact with the first supraciliary, the prefrontal, the anterior 
loreal, the second and third supralabials, and is bordered behind by two large pre- 
oculars. The upper one of these large preoculars is posteriorly bordered by a somewhat 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 403 


Fics 3-4 


(3) Lateral view of the holotype of Lygosoma boehmei sp. n. (ZFMK 86359) in life. (4) Ventral 
view of the holotype of Lygosoma boehmei sp. n. (ZFMK 86359) in life. 


smaller preocular, that touches the first and second supraciliary above; the lower one 
of the large precoculars is posteriorly bordered by a somewhat smaller, more elongated 
preocular (presubocular), that is in broad contact below with the fourth supralabial and 
further touches the corner of the fifth supralabial. Lower eyelid scaly. Supraoculars 
four, plus one small posterior supraocular (postsupraocular: small scale posterior to the 
supraocular series). Supraoculars laterally bordered each by six supraciliaries. Three 
large postoculars, the upper one bordering the postsupraocular above, the lower one 
(postsubocular) borders the fifth and sixth supralabials below. Ear opening small, ovoid 
to roundish. Seven supralabials, fifth right below the eye. Mental in contact with first 
infralabials and postmental. Undivided postmental in contact with mental, first and 


404 T. ZIEGLER ET AL. 


it n Somme ie ECG 
Cie i mcr lr {i ‘ha 


Fics 5-8 


Lygosoma boehmei sp. n. (ZFMK 86359), preserved holotype. (5) Lateral view of the head. (6) 
Dorsal view. (7) Ventral view. (8) Underside of the left foot. 


second infralabials, and first pair of chinshields. First pair of chinshields in broad 
contact, posteriorly bordered by six scales. Seven infralabials. 32 midbody scale rows. 
66 middorsal (paravertebral) scales from posterior end of parietals to insertion of 
hindlimb (70 to centre of thigh, 72 to posterior hindlimb). Dorsal and dorsolateral 
scales smooth, appearing as three longitudinal (pseudo-)keels on neck, body, and 
anterior half of tail. Although visual inspection and the macro-photograph of the dorsal 
midbody scales of the holotype indicates a keeled appearance of the scale surfaces 
(Fig. 9), the SEM (scanning electron microscopy) pictures revealed that the scales bear 
in fact only pseudo-keels (Fig. 10). Contrary to its naming by Darevsky & Orlova 
(1996), the same is actually true for the holotype of L. carinatum (Fig. 11). 

Dorsal body scales of the holotype of L. boehmei sp. n. as large as ventrals. 
Ventral scales smooth, arranged in 81 transverse rows from first gular row subsequent 
to first pair of chinshields to preanals. Six slightly enlarged preanals. 108 median sub- 
caudal scales (without tail tip), not enlarged, smooth. Fourth toe with 14 keeled 
subdigital lamellae. 

Colour in preservative brownish grey on dorsal and dorsolateral surfaces of 
head and body, the surfaces of the limbs and tail appear brownish black. Especially the 
dorsal scales are distinctly edged by brownish black. Except for the somewhat darker 
tail, the lateral parts of the tail base, body, neck and head become lighter. The light ven- 
tral side is yellowish beige (lightest below head and neck) to greyish below the tail with 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 405 


Fic. 9 


Macro-photograph of the apparently keeled dorsal midbody scales in the preserved holotype of 
Lygosoma boehmei sp. n. (ZFMK 86359). 


some indistinct dark marbling. Sides of body and neck with irregular dark flecking. 
Sutures of anterior supra- and infralabials edged by greyish black. In life, the dorsal 
colour is reddish brown. Chin and throat are light orange and the remaining ventral side 
of the body is cream to light brownish, turning to greyish at the underside of the tail. 


COMPARISONS: The new species is easily distinguishable from the skink species 
listed in the genera Lygosoma and Riopa known to occur in Vietnam (Darevsky & 
Orlova, 1996, Nguyen et al., 2005) by a combination of proportions, coloration and 
scalation features (Table 3). The new species differs: 

from Lygosoma carinatum by having seven infralabials instead of six lower 
labials, 32 midbody scale rows instead of 38-40, and only 66 middorsal (paravertebral) 
scales instead of 81-85 according to Darevsky & Orlova (1996). However, our study 
of the holotype of L. carinatum revealed that the specimen only bears 6 infralabials on 
the right side, but 7 on the left (see Table 2); the holotype of L. carinatum further had 
92 transverse rows of ventrals (instead of 81 in the new species), 115 median subcau- 
dal scales (instead of 108), 16 keeled subdigital lamellae beneath the fourth toe (instead 
of 14), and the holotype of L. carinatum had an enlarged nuchal scale on the left side; 

from L. quadrupes by lacking an extremely elongate body (the length of the 
forelimb is not contained twelve to fourteen times in the distance between the fore- and 
hindlimbs), by the rostral not being in contact with the frontonasal, by having 32 mid- 
body scale rows instead of 24-26, by having 14 instead of 5 lamellae beneath the fourth 


406 T. ZIEGLER ET AL. 


TABLE 3. Overview of supralabial (SuL), midbody scale (MbS), and middorsal (MdS) scale 
counts, as well as presence of dorsal keels or pseudo-keels (DK: +/-) and number of lamellae 
beneath fourth toe (LAT) for the Lygosoma / Riopa species recorded from Vietnam (after 
Bourret, unpubl.; Smith, 1935; Taylor, 1963; Darevsky & Orlova, 1996; Manthey & Grossmann, 
1997, and own data). 


Sul MbS MdS DK LAT 

albopunctatum Ù 26-28 63-72 - 12-15 
angeli ? 30 110-115 - 5) 
boehmei sp. n. U 52 66 + 14 
bowringü 7 26-32 52-58 -/+ 10-15 
carinatum 7 38-40 80-85 + 15-16 
corpulentum 6 36-38 ? - 13-14 
punctatum 7 24-28 62-76 - 11-14 
quadrupes 6-7 24-26 104-121 - 5 


toe, and by lacking the tail being as thick as the body for a considerable part of its 
length (Smith, 1935); 

from L. (‘Riopa’) punctatum by lacking a lower eyelid with an undivided semi- 
transparent disc, smooth dorsal scales, by having 32 midbody scale rows instead of 
24-28, and by lacking dorsal scales with a dark basal spot, usually confluent into four 
to six longitudinal lines down the back in the young (Smith, 1935). 

from L. (‘Riopa’) albopunctatum by having 32 midbody scale rows instead of 
26-28, lacking dark brown or black sides of neck and anterior part of the body, and by 
lacking smooth dorsal scales with a dark spot, forming longitudinal series (Bourret, un- 
publ.; Smith, 1935); 

from L. angeli (as ‘Riopa angeli’ in Bourret, unpubl.; Nguyen et al., 2005) by 
lacking a single frontoparietal, having 32 midbody scale rows instead of 30, having 
pseudo-keeled dorsal scales, 66 middorsal (paravertebral) scales instead of 110-115 
scales down the middle of the back, and by having 14 instead of 5 lamellae beneath the 
fourth toe (Bourret, unpubl.; Darevsky & Orlova, 1996). 

from L. bowringii (as ‘Riopa bowringi’ in Smith, 1935, and as Riopa bowringii 
in Nguyen er al., 2005) in lacking a single frontoparietal and a pair of nuchals, by 
having 66 middorsal (paravertebral) scales instead of 52-58 scales down the middle of 
the back, by lacking smooth dorsal scales with a darker spot forming more or less 
continuous longitudinal lines, and by lacking a dark brown or black dorso-lateral stripe 
of variable thickness, light-edged above (Bourret, unpubl.; Smith, 1935; Darevsky & 
Orlova, 1996); 

from L. corpulentum (as ‘Riopa corpulenta’ in Smith, 1935; Nguyen ef al., 
2005) in having seven supralabials (vs. six in corpulentum), pseudo-keeled dorsals (vs. 
smooth body scales in corpulentum), 32 midbody scale rows instead of 36-38, and in 
coloration and pattern (Lygosoma corpulentum is light yellowish-brown and thickly 
mottled on the back and sides with dark brown) (Smith, 1935; Darevsky & Orlova, 
1996). 

From the Lygosoma / Riopa species of the neighbouring countries, the new 
species differs [e.g. comp. Bourret (unpubl.); Werner (1909); Smith (1935); Taylor 
(1963); Manthey & Grossmann (1997)] by lacking lower eyelids with disc in combi- 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 407 


+ 600 um — 


Fics 10-11 


(10) Lygosoma boehmei sp. n. (ZFMK 86359), preserved holotype. SEM (scanning electron mi- 
croscopy) picture of a dorsal midbody scale; the externally visible scale surface is smooth, show- 
ing the “keels” being in fact pseudo-keels. (11) SEM picture of a dorsal midbody scale of the 
holotype of Lygosoma carinatum (ZIN 20482); the externally visible scale surface is smooth, 
showing the “keels” being in fact pseudo-keels. 


nation with 22 smooth midbody scale rows, only 6-10 lamellae under the fourth toe, 
and more or less developed dorso-lateral line (as in the species anguinum and lineo- 
latum), by lacking a single frontoparietal in combination with a) supranasals fused 
anteriorly with nasals, and 88-98 middorsal scales (as in isodactylum), b) in combi- 
nation with 40-42 midbody scale rows, and 143 middorsal scales (as in haroldyoungi), 
or c) in combination with 28-30 midbody scale rows, 56-60 middorsal scale rows, 78 
slightly enlarged median subcaudal scales, and dorsolateral lines (as in fronto- 
parietale), by lacking a combined nasal-supranasal scale in combination with eight 
supralabials and smooth dorsals (as in koratense), by lacking a midbody scale count of 
26-30 in combination with six infralabials, 55-57 middorsal scales and a more or less 
developed stripe from eye along side of body (as in herberti), and by having different 
midbody scale counts from the Indonesian species bampfyldei (38-40) and opistho- 
rhodum (30), the latter of which has in contrast a light lateral stripe and a blackish 
brown anterior dorsum, which is getting paler backwards and turning to a light reddish 
brown tail. 


DISTRIBUTION: Currently, the new species is only known from its type locality, 
the karst forest of Cha Noi, 350-400 m a.s.l., within Phong Nha - Ke Bang National 
Park in Quang Binh province, central Vietnam (Fig. 12). 


NATURAL HISTORY: The female holotype of Lygosoma boehmei sp. n. was 
discovered at night on the forest ground in a steep primary karst forest area (compare 
Fig. 13). We did not find any water courses in the immediate vicinity, however, the 
discovery took place at the end of the dry season. The skink was found crawling at the 
base of a tree stump in the surrounding of karst rock outcrops (Fig. 14). 

The stomach content of the holotype of Lygosoma boehmei sp. n. contained 
brown earth-like masses, plant and tissue remains which most probably come from an 
earth-worm. 


408 T. ZIEGLER ET AL. 


CHINA 


VIETNAM 


Hanoi 
e 


CAMBODIA 


Fic. 12 


At present, Lygosoma boehmei sp. n. is only known from its type locality in the Phong Nha - Ke 
Bang National Park, Quang Binh Province, Vietnam. 


Lygosoma boehmei sp. n. is the third scincid species that has been described 
from the karst forests of Phong Nha - Ke Bang National Park since 2005 (Darevsky & 
Orlov, 2005; Ziegler et al., 2005); the discovery of the new species took place at the 
locus typicus of the recently discovered water skink species Tropidophorus noggei (see 
Ziegler et al., 2006). 


MOLECULAR PHYLOGENETIC POSITION OF LYGOSOMA BOEHMEI SP. N. 

Since L. carinatum, the supposed closest relative of L. boehmei sp. n. could not 
be included in the molecular data set (the former species is only known from the type 
series), we could not make a direct genetic comparison of all Vietnamese Lygosoma 
species (but compare the morphological discussion). Therefore, we decided to analyze 
the phylogentic position of L. boehmei sp. n. within the general framework of the taxa 
of the genus Lygosoma. 

All molecular analyses produced an almost identical tree topology shown in 
Figure 15; the only difference was found in the MP-tree (not shown) where L. albo- 
punctatum was not directly clustered within the clade containing Lygosoma sp. and the 
3 included members of the L. bowringii species complex. The heuristic search of the 
MP analysis produced 2 most-parsimonious tree (tree length = 194; CI = 0.680; RI = 
0.682; RC = 0.464). The comparison between the different likelihood scores for each 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 409 


Fics 13-14 


(13) The steep karst forests of Cha Noi: habitat of Lygosoma boehmei sp.n. in the Phong Nha - 
Ke Bang National Park. (14) Microhabitat of Lygosoma boehmei sp. n.: the female holotype was 
collected at night crawling at the base of this tree stump. 


410 T. ZIEGLER ET AL. 


Lygosoma lineolatum Myanmar (Rakhine State) 


Lygosoma lineolatum Myanmar (Mandalay Div.) 


Lygosoma boehmei sp. nov. Vietnam 


86/84/94 
Lygosoma quadrupes Thailand 


Lygosoma koratense Thailand 


100/99/100 


Lygosoma koratense Malaysia 


100/100/100 


Lygosoma albopunctatum India 


Lygosoma sp. India 


Lygosoma bowringii (l)Thailand 


Lygosoma bowringii Indonesia (Bali) 


100/99 
/95 
1.00 


Lygosoma bowringii (II) Thailand 


Fic. 15 


Phylogram based on 548 bp of the mitochondrial 16S ribosomal RNA gene sequences. Values 
above the nodes represent bootstrap values in percent for maximum parsimony, neighbor-joining 
and maximum likelihood analyses, respectively; lower values are Bayesian posterior probabi- 
lities. Values below 50% (or 0.50 for the Bayes analyses) not shown. 


model showed that the TRN+I+G model (Tamura & Nei, 1993) was determined to be 
the optimal ML model for data set. This model incorporates unequal base frequencies 
[TA) = 0.3461, mq) = 0.2335, tc) = 0.2448, 1g) = 0.1756], a proportion of invariable 
sites (I = 0.4165), and a gamma distribution shape parameter (a = 0.2513). The optimal 
ML tree had a log-likelihood of -InL = 1766.91104. 

The resulting tree shows three clearly separated groups with the two specimens 
of L. lineolatum genetically well separated from the rest of the included species; this 
was true even when these two OTUs were included as ingroup taxa and a different out- 
group was used. The newly described Lygosoma boehmei sp. n. is consistently placed 


411 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 


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412 T. ZIEGLER ET AL. 


into a second clade, which also includes the type species of the genus Lygosoma (L. 
quadrupes), as well as the two L. koratense specimens from Thailand and Malaysia 
(whose sequences showed a 100% identity despite coming from two different coun- 
tries). This whole clade is strongly supported in all used analyses methods (MP: 86 / 
NJ: 84 / ML: 94 /PP: 1.00) and shows that L. boehmei sp. n. is a true member of the 
genus Lygosoma, regardless of the ongoing discussion on the validity of the genus 
Riopa (see Introduction). Lygosoma boehmei sp.n. stands basal to the other OTUs of 
this clade, but a definitive polarity decision cannot be made with our data set. The ge- 
netic differences between the different species of this clade varied from 6.8%-8.5%. L. 
boehmei sp. n. is also well separated from the members of the third recovered clade, 
which includes all three specimens of the L. bowringii species complex as well as 
Lygosoma sp. and L. albopunctatum (both from India). The genetic differences be- 
tween the taxa of this third clade were similar to those found in the second clade and 
varied from 6.4%-9.5%. Regarding the distances between the two clades one finds that 
they are normally slightly higher than the distances within the two clades (9.4%- 
11.7%), but this is not completely consistent since the genetic distance of one of these 
species (L. boehmei sp. n.) towards the taxa of the third clade is in the range of or even 
slightly lower (8.1%-9.2%) than the maximum inter-clade differences mentioned 
above. Also we found that in some cases the genetic differences between geographi- 
cally close taxa are not necessary lower than those living in far-away habitats (e.g. L. 
boehmei sp. n.-L. lineolatum and L. boehmei sp. n.-L. koratense; Table. 4). This shows 
that a subdivision of the genus Lygosoma would need to be based on a rather complete 
taxon sampling, since our preliminary molecular data already indicate a rather compli- 
cated intrageneric structure within Lygosoma. One further notable result is that (equal 
as shown in our key to the Vietnamese Lygosoma species where the L. bowringii 
species complex is found on two different positions in the key) the genetic analyses al- 
so strongly support that L. bowringii is a species-complex which comprises at least 2 
different taxa. The recovered genetic difference between the two different specimens 
from Thailand is 6.4% and therefore has about same amount of genetic divergence as 
between other Lygosoma species, and the new L. boehmei sp. n. is about equidistant 
with all three included L. bowringii taxa (8.1%-8.6%). 


KEY TO THE VIETNAMESE LYGOSOMA SPECIES 


la Dorsal scales appearing Smooth . .. . li... OSSEE 2 
lb Dorsal scales with keels or pseudo-keels;.> 7. -.t.. ene 7 
2a Silamellac*beneath fourth toe’ MEME RE 2020 EEE 3 
2b 10 15 lamellae beneath fourth toe. E, Ae RAEE 4 
3a 24-26 midbody scale rows, 2.4.5. IH 225 eee quadrupes 
3b S0imidbody:scale TOWSELE au LE ae eee ee angeli 
4a 36-38 midbody scale rows, six supralabials ................. corpulentum 
4b 24-32 midbody scale rows; seven supralabials 00 E eee 9 
Sa S2-9 8umiddorsaliscalese enne Bet bowringii (species complex) 
Sb 62-76: middorsal scales: <4... tenti ESRI ERA 6 


6a lower eyelid scaly (central scales may be enlarged) .......... albopunctatum 


NEW SPECIES OF LYGOSOMA FROM VIETNAM 413 


6b lower eyelid with undivided semitransparent disc ............... punctatum 
7a 26-32 midbody scale rows, 52-66 middorsal scales .................... 8 
7b 38-40 midbody scale rows, 80-85 middorsal scales .............. carinatum 
8a 9258: middorsalsseales® 2.2... 24... 22 20.2.2. 0008 bowringii (species complex) 
8b GOpmiid dorsal: SCALES 4 ita boehmei sp. n. 
ACKNOWLEDGEMENTS 


We thank Dr V. T. Hoang (Centre for Natural Resources and Environmental 
Studies, Vietnam National University, Hanoi), the People’s Committee of Quang Binh, 
and the Phong Nha — Ke Bang National Park directorate (T. H. Nguyen, X. C. Cao, and 
M. T. Luu) for their continuous support as well as for issuing respective permits. The 
National Park staff, especially Dang Ngoc Kien helped again to make fieldwork suc- 
cessful. For their support we further would like to express our thanks to Dr Martina 
Vogt (Cologne Zoo nature conservation project, Phong Nha - Ke Bang) and Bernhard 
Forster (Frankfurt Zoological Society / Cologne Zoo nature conservation project, 
Phong Nha - Ke Bang). We are further grateful to Dr Natalia B. Ananjeva (ZIN, 
Zoological Institute, Russian Academy of Sciences, St. Petersburg) for enabling us to 
examine the holotype of Lygosoma carinatum. We thank Philipp Wagner and Thomas 
Wilms (ZFMK, Bonn) for critically reviewing the manuscript. For the translation of the 
scientific description of Lygosoma carinatum in Russian language we would like to 
express our thanks to our friend and colleague Ho Thu Cuc (Institute of Ecology and 
Biological Resources, Vietnamese Academy of Science and Technology). Philipp 
Wagner (ZFMK, Bonn) kindly prepared the SEM pictures. Our field work was funded 
in large by the Zoological Garden Cologne, the Kölner Kulturstiftung der Kreiss- 
parkasse Köln, and BIOPAT (www.biopat.de). Current lizard research in the Phong 
Nha - Ke Bang region is supported as well by the Alexander-Koenig-Gesellschaft 
(AKG), the Alexander Koenig Stiftung (AKS), the Deutsche Gesellschaft für 
Herpetologie und Terrarienkunde (DGHT), and by the European Union of Aquarium 
Curators (E.U.A.C.). 


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ZIEGLER, T., HERRMANN, H.-W., Vu, N. T., LE, K. Q., NGUYEN, T. H., CAO, X. C., Luu, M. T. & 
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ZIEGLER, T. & LE, K. Q. 2006. A new natricine snake of the genus Amphiesma (Squamata: 
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REVUE SUISSE DE ZOOLOGIE 114 (2): 417-435; juin 2007 


Five new European species of the Rhamphomyia (s. str.) 
albosegmentata group (Diptera: Empididae) 


Miroslav BARTAK 

Czech University of Agriculture, Faculty of Agrobiology, Food and Natural Resources, 
Department of Zoology and Fishery, 165 21 Praha 6 - Suchdol, Czech Republic. 
E-mail: bartak@af.czu.cz 


Five new European species of the Rhamphomyia (s. str.) albosegmentata 
group (Diptera: Empididae). - Rhamphomyia (s. str.) bellinosetosa sp. n. 
(Italy), Rhamphomyia (s. str.) piedmontensis sp. n. (Italy), Rhamphomyia 
(s. str.) ponti sp. n. (Austria), Rhamphomyia (s. str.) saintbaumensis sp. n. 
(France), and Rhamphomyia (s. str.) sellacrinita sp. n. (Italy, Switzerland), 
are described and illustrated. The key to the Palaearctic species of the 
Rhamphomyia (s. str.) albosegmentata group is updated. 


Keywords: Rhamphomyia - new species - Europe - taxonomy - key. 


INTRODUCTION 


The species of the Rhamphomyia (s. str.) albosegmentata group are usually 
medium-sized (wing size from 3.2 to 8.9 mm), with characteristic shape of genitalia 
(cercus [= dorsal genital lamella] simple, without «subcercal process», with finger-like 
projection near base of its upper margin, phallus broadened apically), proepisternal 
depression setose, axillary angle acute, anal vein (A1) complete, prosternum bare, 
costal seta present, haltere dark and hind femur with ventral setae. Females are some- 
times difficult to distinguish from other groups and species complexes of the subgenus 
Rhamphomyia (s. str.). 

Palaearctic species of the group were revised by Bartäk (1981), however, three 
species were described more recently (Bartäk, 1998) and five additional species are 
described here as new to science, and this necessitates revision of the earlier key 
(Bartäk, 1981). 


MATERIAL AND METHODS 


The material studied is deposited in the following collections: 
CNC Canadian National Collection, Ottawa 
CUA Czech University of Agriculture, Prague 
MHNG Muséum d’histoire naturelle, Geneva 
NHM Natural History Museum, London 
SMNS Staatliches Museum für Naturkunde, Stuttgart 
UMO University Museum, Oxford 


Manuscript accepted 24.12.2006 


418 M. BARTAK 


The genitalia were macerated in 10% KOH (24 hours, room temperature) and 
they are stored together with specimens in plastic microvials with glycerine. The mor- 
phological terms used here follow Merz & Haenni (2000) and Sinclair (2000). 
Abbreviations: T11,T21,T3l = length of fore, mid, hind tibia; B11,B21,B31 = length of 
fore, mid, hind basal tarsomere; B1w,B2w,B3w = width of fore, mid, hind basal tar- 
somere; M2/D = length of vein M2: greatest length of discal medial cell (discal cell); 
M3/Db = length of apical: preapical sections of vein CuA1; lw: ww = greatest length 
of wing: greatest width of wing. Ratio of antennal segments = length of first: 2nd: 3rd: 
style (in 0.01 mm scale). 


SYSTEMATIC TREATMENT 


Rhamphomyia (s. str.) bellinosetosa sp. n. Figs 1,2 


HOLOTYPE MALE: Italy: Piedmont, Celle de Bellino, 11.-12.vii.1974, Malaise trap, 
W. Baker leg. (CNC). 


DESCRIPTION 

Male: Eyes holoptic, facets in lower half of eye much smaller than upper ones. 
Frons black, grey pruinose, bare. Ocellar setae black, fine, 2/3 as long as frons, ocellar 
triangle with single pair of additional slightly smaller setulae. Face black, grey 
pruinose, about 0.35 mm broad below and 0.20 mm long, bare. Occiput black, light 
grey pruinose, medium densely black setose, dorsal setae quite strong, postocular row 
incomplete. Antennae black, ratio of antennal segments = 17: 10: 55: 11, both basal 
segments covered with long setae, longest ones about 0.25 mm long. Labrum brownish 
black, polished, subequally long as head is high. Palpus brown, long, densely covered 
with very long bristly setae (the longest 0.50 mm long). Genae narrow and mostly 
polished, clypeus pruinose. Thorax black, light grey pruinose, mesoscutum light grey 
without distinct stripes. All setae and setulae black. Chaetotaxy: about 15 setulae on 
proepisternum; prosternum bare; proepisternal depression with one setula; 20 biserial 
acrostichals; 20-25 irregularly 2-3 serial dorsocentrals ending in 4 strong prescutellars 
(both acrostichals and dorsocentrals rather strong and up to 0.40 mm long); 1-2 
intrahumeral; 1-2 posthumeral; about 20 additional setulae lateral of dorsocentrals in 
presutural area; 1 postpronotal seta and about 12 additional shorter setae; 2-3 noto- 
pleurals (2-4 setulae in front part of notopleura, some of them quite strong); a line of 
4-5 strong setae in supraalar and prealar areas, 1-2 prealar and 6-7 additional setae in 
unusual position between supraalars and dorsocentrals (even between hindmost 
supraalars and hind dorsocentrals); 1 long, 1 shorter and several additional small post- 
alar; 3 pairs of scutellars; laterotergite (metapleura) with black setae. Coxae brownish 
black and pruinose, concolorous with pleura. Legs blackish brown and pruinose, black 
setose. A long seta present in comb at tip of hind tibia. Fore femur (Fig. 2) with sparse 
and irregular rows of anteroventral, posteroventral and posterior setae slightly longer 
than femur is deep, dorsal setulae very short, but several anterodorsal and nearly 
complete row of anterior setae are somewhat longer. Fore tibia (Fig. 2) with row of (8) 
very strong anterodorsal and irregular row of (greater number) similarly strong 
posterodorsal and posterior setae up to 3 times as long as diameter of tibia (up to 
0.45 mm long with many much smaller and finer setulae between them), very fine 


NEW EUROPEAN RHAMPHOMYIA 419 


Fics 1-2 


Rhamphomyia (s. str.) bellinosetosa sp. n. (1) Male terminalia (macerated), lateral view. Scale 
0.1 mm. (2) Male fore femur, tibia and basitarsus, posterior view. Scale 0.5 mm. 


ventral ciliation consists of setulae subequally long as diameter of tibia. Mid femur 
with anteroventral row of setae slightly longer than femur is deep, posteroventral setae 
much shorter and finer, only a single strong posteroventral present on basal third. Mid 
tibia with two rows of anterodorsal and posterodorsal setae (about 6 in each row) 
almost 4 times as long as diameter of tibia and with anteroventral and posteroventral 
setae (3 in each row) twice as long as diameter of tibia. Hind femur with 5 antero- 
ventral, a single long submedian posteroventral seta and 2-3 subapical anterodorsal 
setae at least as long as femur is deep, several fine posterior setae on base of femur 
long, other ciliation short. Hind tibia with anterodorsal and posterodorsal setae (6-8 in 
each row) twice as long as diameter of tibia (small setulae between them are much 
shorter), ventral setulae short. Basal tarsomere of fore leg thin, with several strong 
setae dorsally up to 3 times as long as diameter of this tarsomere and with short ventral 
spines, T1l: B1] = 2.1, BI]: BIw = 5.0; basal tarsomere of mid leg short and thin, short 
setose and with short ventral spines, T21: B21 = 2.8, B21: B2w = 5.0. Basal tarsomere 
of hind leg thin, dorsally with some 6 setae three times as long as diameter of this 
tarsomere and with long ventral spines, T31: B31 = 2.2-2.3, B31: B3w = 7.7. Wing clear 
or slightly brownish, stigma light brown, veins brown, anal vein (A|) complete. Costal 
seta long, axillary angle sharply acute (30°). M2/D = 1.3, M3/Db = 1.8, lw: ww = 3.0. 
Haltere brown, calypter yellow with dark fringes. Abdomen black, light grey pruinose 
(light grey in dorsal view). Setae and setulae all black. Hind marginal setae on sides of 
tergites 2-5 strong and slightly longer than their segments, on segments 6-7 fine and 
slightly shorter than their segments, discal setulae shorter than hind marginals. The first 
sternite with several setulae on sides. Terminalia as in Fig. 1; apical angle of cercus 
slightly obtuse. Length of body: 5.7 mm, wing: 6.1 mm. 
Female: Unknown. 


DIFFERENTIAL DIAGNOSIS: Rhamphomyia (s. str.) bellinosetosa sp. n. is similar to 
R. hirtimana Oldenberg, 1922, however, the latter species has fore tibia with almost 
homogeneously fine dorsal ciliation, more sparsely setose mesoscutum (about 10 setae 


420 M. BARTÄK 


on presutural area of mesoscutum lateral of dorsocentrals), labrum longer than head is 
high and palpi sparsely setose (less than 15 setae on each palpus, whereas more than 
20 in the new species described above). 


DERIVATIO NOMINIS: The species name is derived in part from the type locality 
denomination (bellino-) and -setosa which refers to the strongly setose fore tibia. 


DISTRIBUTION: Italy. 
DATES OF OCCURRENCE: July. 


Rhamphomyia (s. str.) piedmontensis sp. n. Fig. 3 


HOLOTYPE MALE: Italy: Celle de Bellino, Piedmont, July 11-12, 1974, W. Baker - Mal. 
Trap (CNC). 
DESCRIPTION 

Male: Eyes holoptic, facets in lower half of eye smaller than upper ones. Frons 
black, grey pruinose, bare. Ocellar setae black, about as long as upper postocular ciliae, 
ocellar triangle with 4-5 pairs of additional slightly shorter setulae. Face black, grey 
pruinose, about 0.45 mm broad below and 0.30 mm long, bare. Occiput black, grey 
pruinose, rather densely and long black setose. Antenna black, ratio of antennal seg- 
ments = 24: 13: 60: 15, both basal segments covered with very long setae (longest 
about 0.45 mm long). Labrum brownish black, polished, slightly longer than head is 
high. Palpus brown, long, very densely covered with very long bristly setae (longest up 
to 0.60 mm long). Genae narrow and pruinose, clypeus pruinose. Thorax black, dark 
grey or slightly brownish grey pruinose, mesoscutum without stripes. All setae and 
setulae black. Chaetotaxy: about 40 setulae on proepisternum; prosternum bare; about 
15 setulae on proepisternal depression; great number of 4-5 serial acrostichals; great 
number of multiserial dorsocentrals, ending in 3-4 strong prescutellars, spreading 
down sides of mesoscutum in both pre- and postsutural area (whole mesoscutum is 
densely covered with setulae, leaving only very narrow stripes between acrostichals 
and dorsocentrals bare, stripes much narrower than space occupied by acrostichal 
setae, areas just around postalar callus and notopleuron behind notopleural setae bare), 
setae on mesoscutum about 0.40 mm long; intrahumeral and posthumeral setae not dif- 
ferentiated; postpronotal scarcely differentiated from many similar setulae; 4 noto- 
pleurals (many long setulae in front part of notopleuron); supraalar and prealar area 
densely covered with setulae, 2-3 supraalars stronger and longer; 1 long and 1 short 
postalars; 4 pairs of scutellars; laterotergite (metapleura) with black setae. Coxae and 
legs concolorous with pleura, pruinose, black setose. Long seta present in comb at tip 
of hind tibia. Fore femur with rather sparse anteroventral and more dense 
posteroventral and posterior setulae about as long as femur is deep, dorsal setulae 
shorter. Fore tibia with almost equally long but unequally strong setae and setulae 
dorsally only slightly longer than diameter of tibia, ventral setulae very short. Mid 
femur with anterodorsal, anteroventral and posteroventral setae slightly longer than 
femur is deep, other ciliation short. Mid tibia with 4-5 pairs of setae dorsally at most 
twice as long as diameter of tibia, anteroventral and posteroventral setae shorter. Hind 
femur with ventral, anterodorsal and posterior setae slightly longer than femur is deep. 
Hind tibia very slightly flattened, with 7-8 pairs of closely inserted anterodorsal and 
posterodorsal setae slightly longer than diameter of tibia, ventral setulae very short. 


NEW EUROPEAN RHAMPHOMYIA 421 


Basal tarsomere of fore leg thin and short setose, ventral spines not prominent, T11: B11 
= 2.5, Bll: Blw = 7.1, basal tarsomere of mid leg short and thin, with several ventral 
spines, T21: B21 = 2.6, B21: B2w = 5.0. Basal tarsomere of hind leg thin, dorsally with 
several setae more than twice longer than diameter of this tarsomere, ventral spines 
slightly longer than ciliation, T3l: B3l = 2.4, B3l: B3w = 5.7. Wing slightly brownish, 
stigma brown, veins brown, anal vein (A|) complete. Costal seta long, axillary angle 
sharply acute. M2/D = 1.4, M3/Db = 2.4, lw: ww = 3.0. Haltere brownish-black, 
calypter brown with dark fringes. Abdomen black, dark brownish-grey pruinose 
(brown in dorsal view). Setae and setulae black. Hind marginal setae on sides of 
tergites 2-5 1.5 times longer than their segments, on tergites 6-7 shorter, discal setulae 
shorter than hind marginals. Dorsum of tergites with unusually long setae (marginals 
about as long as their segments). The first sternite setulose on sides. Terminalia as in 
Fig. 3. Length of body: 6.0 mm, wing: 6.5 mm. 
Female: Unknown. 


DIFFERENTIAL DIAGNOSIS: Rhamphomyia (s. str.) piedmontensis sp. n. differs 
from all other known species of R. albosegmentata group by multiserial acrostichal 
setae. In spite of female remaining unknown, I believe it could be very easily identified 
according to the following combination of characters: multiserial acrostichals, long 
costal seta, dark haltere, axillary angle sharply acute and complete anal vein. These 
characters are usually not sexually dimorphic and are unique among the Palaearctic 
fauna of Rhamphomyia. 


DERIVATIO NOMINIS. The species is named after the type locality. 
DISTRIBUTION: Italy. 
DATES OF OCCURRENCE: July. 


Rhamphomyia (s. str.) ponti sp. n. Figs 4,5 


HOLOTYPE MALE: Austria: Tyrol, Oetztal, Obergurgl, Gaisbergertal to Hohe Mut, heath, 
2400 - 2600 m, 7.viii.1981, leg. A.C. Pont (NHM). - Paratype. Trogeralm, Glockner, 16.vii.1941, 
“Rhamphomyia hirtimana Loew, Lindner det.”, 1 male (SMNS). 


DESCRIPTION 

Male: Eyes holoptic, facets in lower half of eye much smaller than upper ones. 
Frons black, grey pruinose, bare. Ocellar setae black, fine, 2/3 as long as frons, ocellar 
triangle with 2-3 pairs of additional slightly smaller setulae. Face black, grey pruinose, 
about 0.30 mm broad below and 0.20 mm long, bare. Occiput black, light grey 
pruinose, medium densely and medium long black setose, postocular row incomplete 
and distant from eye margin in middle. Antenna black, ratio of antennal segments = 18: 
11: 55: 14, both basal segments covered with long setae (longest about 0.25 mm long). 
Labrum brownish black, polished, slightly longer than head is high. Palpus brown, 
long, sparsely covered with long bristly setae (longest are 0.40 mm long). Genae 
narrow and mostly polished, clypeus pruinose. Thorax black or brownish black, meso- 
scutum with distinct brownish tinge, without stripes or with three darker stripes under 
rows of acrostichals and dorsocentrals visible only in sharp caudal view. All setae and 
setulae black. Chaetotaxy: 10-15 setulae on proepisternum; prosternum bare; 1-3 
setulae on proepisternal depression; 24-26 biserial acrostichals; 25-30 irregularly 


422 M. BARTÀK 


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Rhamphomyia (s. str.) piedmontensis sp. n. (3) and Rhamphomyia (s. str.) ponti sp. n. (4-5). (3- 
4) Male terminalia, lateral view. Scale 0.1 mm. (5) Male fore femur, tibia and basitarsus, 
posterior view. Scale 0.5 mm. 


triserial dorsocentrals ending in 1-2 strong prescutellars (both acrostichals and 
dorsocentrals moderately strong and about 0.30 mm long); 0-2 intrahumeral; 0-1 
posthumeral; about 10 additional setulae lateral of dorsocentrals in presutural area; 
1 postpronotal scarcely differentiated from 15 setulae; 3-4 notopleurals (3-5 setulae in 
front part of notopleuron); 1 long supraalar, 3-4 prealars and 2-3 other setae in line 
between them; 1 long and 1 short postalars; 2 pairs of strong and 1 pair of fine scu- 
tellars; laterotergite (metapleura) with black setae. Coxae brownish black and pruinose, 
concolorous with pleura. Legs blackish brown and pruinose, black setose. Long seta 
present in comb at tip of hind tibia. Fore femur (Fig. 5) densely covered with irre- 
gularly arranged posteroventral setae and less numerous anteroventrals both at least 
twice as long as femur is deep, also anterior and posterior setae longer than femur is 
deep, dorsal setae subequally long as femur is deep. Fore tibia (Fig. 5) densely covered 
with long setae along dorsal and posterior surfaces (almost 0.70 mm long, i.e. 4-5 times 
as long as diameter of tibia), dense ventral ciliation consists of setulae subequally long 
as diameter of tibia and between these setulae with irregular row of 6-8 ventral setae 
2-3 times as long as diameter of tibia. Mid femur with two rows of ventral setae at least 


NEW EUROPEAN RHAMPHOMYIA 423 


1.5 times as long as femur is deep, other ciliation short. Mid tibia with two irregular 
rows of anterodorsal and posterodorsal setae, 4-5 of them in each row about as long as 
those on fore tibia (0.70 mm), and with two rows (3-4 setae in each row) of unusually 
long anteroventral and posteroventral setae (0.50 mm long). Hind femur with 3-8 
anteroventrals (8 strong in holotype and 3 fine in paratype), single long submedian 
posteroventral and 2 subapical anterodorsal setae at least as long as femur is deep, also 
several fine posterior setae on base of femur long, other ciliation short. Hind tibia with 
anterodorsal and posterodorsal setae (4-8 in each row) up to three times as long as 
diameter of tibia, ventral setulae contrastingly short. Basal tarsomere of fore leg thin, 
densely covered with setae dorsally only slightly shorter than the length of this 
tarsomere, also ventral spines unusually long, longer than diameter of this tarsomere 
(long dorsal setae occur also on second tarsomere), T1l: B11 = 1.9-2.0, Bll: Blw = 
5.0-5.5; basal tarsomere of mid leg short and thin, with several long dorsal setae and 
long ventral spines, T21: B21 = 2.4-2.6, B21: B2w = 5.0-6.1. Basal tarsomere of hind 
leg thin, dorsally with some 6 setae three times as long as diameter of this tarsomere 
and with long ventral spines, T3l: B3l = 2.1-2.2, B3l: B3w = 5.8-6.1. Wing clear, 
stigma light brown, veins brown, anal vein (A|) complete. Costal seta long, axillary 
angle sharply acute (35°). M2/D = 1.3-1.5, M3/Db = 2.1-2.2, lw: ww = 2.8-3.1. Haltere 
brown, calypter yellowish brown with dark fringes. Abdomen black, light grey 
pruinose (light grey in dorsal view). Setae and setulae all black. Hind marginal setae 
on sides of tergites 2-5 strong and slightly longer than their segments, on segments 
6-7 fine and slightly shorter than their segments, discal setulae shorter than hind mar- 
ginals. Dorsum of tergites with short setae in midline but longer just outside this line. 
The first sternite with several setulae on sides. Terminalia as in Fig. 4; apex of cercus 
right-angled. Length of body: 5.7-6.4 mm, wing: 5.9-6.7 mm. 
Female: Unknown. 


DIFFERENTIAL DIAGNOSIS. Rhamphomyia (s. str.) ponti sp. n. is somewhat similar 
to R. ursina Oldenberg, 1915 or R. kreischi Bartäk, 1998 (due to several long setae 
ventrally on fore tibia), however, the new species differs from both in many characters, 
e.g. mid tibia in both R. ursina and R. kreischi covered along dorsal side with numerous 
long setae not forming rows and abdomen is brown viewed from above. In several 
characters this new species is somewhat smilar to R. hirtimana, however, the latter 
species has short setose second fore tarsomere, mesoscutum light bluish-grey without 
brownish tinge and sparse posterodorsal setae on fore femur forming nearly a row. 


DERIVATIO NOMINIS. The species named after the collector of the holotype, the 
leading world specialist on Muscidae, Dr Adrian C. Pont (Oxford). 


DISTRIBUTION: Austria. 
DATES OF OCCURRENCE: July-August. 


Rhamphomyia (s. str.) saintbaumensis sp. n. Figs 6-8 


HOLOTYPE MALE. France: Provence, Saint Baume, 30.?.1945, deposited in UMO. 


DESCRIPTION. 
Male: Eyes holoptic, facets in lower half of eye smaller than upper ones. Frons 
black, grey pruinose, bare. Ocellar setae rather weak, black, about as long as dorsal 


424 M. BARTÀK 


postocular ciliae, ocellar triangle with 2 additional slightly shorter setulae. Face black, 
grey pruinose, about 0.30 mm broad below and equally long, bare. Occiput black, grey 
pruinose, densely and long black setose. Antenna black, both basal segments brown, 
ratio of antennal segments = 19: 11: 50: 14, longest setulae on basal two segments 
about 0.30 mm long. Labrum brown, polished, about as long as head is high. Palpus 
brown, long, densely covered with long bristly setae. Genae narrow and polished, 
clypeus pruinose. Thorax black, grey pruinose, mesoscutum dark brownish grey, with 
somewhat darker stripes below rows of acrostichals and dorsocentrals visible in sharp 
caudal view. All setae and setulae black. Chaetotaxy: proepisternum with about 30 
setulae; prosternum bare; proepisternal depression with 8 setulae; about 35 biserial, 
fine acrostichals; great number of multiserial dorsocentrals, ending in 2 strong pre- 
scutellars, spreading down the sides of mesoscutum in presutural area (both acro- 
stichals and dorsocentrals about 0.30 mm long); intrahumeral seta not differentiated 
from setulae, 1 posthumeral accompanied with several setulae, postpronotal scarcely 
differentiated from many similar setulae; 3 notopleurals (4-5 long setulae in front part 
of notopleuron); 2 supraalars and about 10 setulae in prealar area; 1 long and 1 short 
postalars; 3 pairs of scutellars; laterotergite (metapleura) with black setae. Coxae 
concolorous with pleura, pruinose, black setose. Legs brown, pruinose, black setose. 
Long seta present in comb at tip of hind tibia. Fore femur densely covered with long 
posteroventral and posterodorsal setae longer than femur is deep, anteroventral ones 
sparser and shorter, posteroventrals do not form row. Fore tibia with dense and almost 
homogeneous setae in posterodorsal area more than 3 times as long as diameter of tibia 
(also at least basal two fore tarsomeres with similar setae), ventral setulae very short. 
Mid femur with rows of anteroventral and posteroventral setae slightly longer than 
femur is deep, other ciliation short. Mid tibia with 4-5 long and strong anterodorsal and 
posterodorsal setae up to 4 times as long as diameter of tibia, and 4 almost equally long 
posteroventrals. Hind femur with 4-5 rather fine anteroventral setae about as long as 
femur is deep and with equally long two subapical anterodorsals and 1 posteroventral 
near middle of femur. Hind tibia thin, with 6-7 pairs of anterodorsal and posterodorsal 
setae, longest setae twice as long as diameter of tibia, ventral setulae short. Basal 
tarsomere of fore leg slightly swollen, densely covered with long setae dorsally (also 
at least second tarsomere long setose), T1l: B11 = 2.3, Bll: Bl w = 3.5, basal tarsomere 
of mid leg slightly swollen, covered with rather long spines ventrally, T21: B21 = 3.0, 
B21: B2w = 3.5. Basal tarsomere of hind leg thin and short setose, T31: B31 = 2.3, B31: 
B3w = 6.7. Wing clear to slightly yellowish, stigma brown, veins brown, anal vein 
(A) complete. Costal seta long, axillary angle sharply acute. M2/D = 1.3-1.4, M3/Db 
= 2.0, lw: ww = 3.2. Haltere dark brown, calypter brownish-yellow with dark fringes. 
Abdomen black, light grey pruinose even in dorsal view. Setae and setulae all dark. 
Hind marginal setae on sides of tergites 2-3 slightly shorter than their segments, on 
tergites 4-5 shorter than their segments and very short on tergites 6-7, discal setulae 
shorter than hind marginals. Dorsum of tergites with slightly shorter setulae. The first 
sternite with several setulae on sides. Terminalia as in Figs 6-8; hypandrium slightly 
bifurcated apically; phallus with lateral «wing», apical swelling irregular. Length of 
body 6.4 mm, wing 6.0 mm. 
Female: Unknown. 


NEW EUROPEAN RHAMPHOMYIA 425 


DIFFERENTIAL DIAGNOSIS. Rhamphomyia (s. str.) saintbaumensis sp. n. is allied 
to several species of the R. albosegmentata group (e.g. R. curvinervis Oldenberg, 1915, 
R. tristriolata Nowicki, 1868, R. hirtimana Oldenberg, 1922) sharing with them the 
following characters: densely and long setose fore femur and light grey abdomen. 
However, it differs from all these species in swollen fore basal tarsomere covered 
(together with the following tarsomere) with very long setae dorsally. Moreover, the 
new species has peculiarly shaped phallus (with lateral «wings») and dark and rather 
brownish mesoscutum. 

DERIVATIO NOMINIS: The species is named after the type locality. 


DISTRIBUTION: France. 


DATES OF OCCURRENCE: Unknown (it is impossible to read the month of collect- 
ing which is illegibly hand-written on the label - March?). 


Rhamphomyia (s. str.) sellacrinita sp. n. Fig. 9 


HOLOTYPE MALE. Italy: Passo Sella, forest boundary, 1800 m, 46°26’N, 11°46’E, 
8.viii.1988, leg. M. Bartäk (CUA). - Paratypes. Same data as the holotype, 1 male (CUA); 
Switzerland, VS Visperterminen, ob. Dorf, 1550 m, 27.v111.2001, Merz and Landry leg., 1 male 
(MHNG). 

DESCRIPTION 

Male: Eyes holoptic, facets in lower half of eye much smaller than upper ones. 
Frons black, grey pruinose, bare. Ocellar setae black, very fine, hardly 1/3 as long as 
frons, ocellar triangle with 1-2 pairs of additional nearly equally long setulae. Face 
black, grey pruinose, about 0.18 mm broad below and 0.40 mm long, bare. Occiput 
black, dark grey pruinose, rather sparsely and short black setose (uppermost postocular 
setae only about 0.10 mm long). Antenna black, ratio of antennal segments = 15: 10: 
39: 12, both basal segments covered with medium long setae (longest about 0.20 mm 
long). Labrum brownish black, polished, slightly shorter than head is high. Palpus 
brown, long, sparsely covered with long bristly setae (longest 0.25 mm long). Genae 
very narrow and mostly pruinose, clypeus pruinose. Thorax black or brownish-black, 
mesoscutum deep black and slightly subpolished, without stripes. All setae and setulae 
black. Chaetotaxy: proepisternum with about 10 setulae; prosternum bare; 1-3 setulae 
on proepisternal depression; about 12 narrowly biserial acrostichals; 12-15 irregularly 
2-3 serial dorsocentrals ending in 2-3 strong prescutellars (both acrostichals and dor- 
socentrals medium strong and about 0.20 mm long); 2 intrahumeral; 0-2 posthumeral 
setae (inserted in unusual lateral position, almost in area of notopleuron), 6-8 fine 
additional setulae lateral of dorsocentrals in presutural area; 1-2 postpronotals in 
addition to several shorter setulae; 3 notopleurals (5-8 setulae in front part of noto- 
pleura, some of them quite strong); 2 supraalars and 2-3 prealars; 1 long and 1 short 
postalars; 2 pairs of strong and 0-1 pair of fine scutellars; laterotergite (metapleura) 
with black setae. Coxae brown and pruinose, legs brown and mostly polished, black 
setose. Long seta present in comb at tip of hind tibia. Fore femur with rather sparse 
anteroventral and posteroventral rows of setae about twice as long as femur is deep one 
third before tip of femur but absent in basal half of femur, dorsal setulae very short. 
Fore tibia with almost homogeneous dorsal and posterodorsal ciliation subequally long 
as diameter of tibia, ventral setulae short. Mid femur with very short and fine setulae, 


426 M. BARTAK 


Fics 6-9 


Rhamphomyia (s. str.) saintbaumensis sp. n. (6-8) and Rhamphomyia (s. str.) sellacrinita sp. n. 
(9). (6) Male terminalia (macerated), lateral view. Scale 0.1 mm. (7) Male phallus and hypan- 
drium (macerated), lateral view. Scale 0.1 mm. (8) Male cercus (macerated), lateral view. Scale 
0.1 mm. (9) Male terminalia (macerated), lateral view. Scale 0.1 mm. 


both anteroventrals and posteroventrals at most 1/3 as long as femur is deep. Mid tibia 
with 4-5 very long anterodorsals almost 4 times as long as diameter of tibia (up to 0.50 
mm long), and with 3-4 shorter posterodorsal; anteroventrals and posteroventrals (1-3 
in each row) up to twice as long as diameter of tibia. Hind femur with 8 strong 
anteroventral and one submedian strong posteroventral setae slightly longer than femur 
is deep and with 1-2 subapical anterodorsal and several posterior setae in basal part of 
femur, other ciliation short (setulae between anteroventral setae much shorter than 
strong anteroventrals). Hind tibia with anterodorsal and posterodorsal setae (5-7 in 
each row) twice as long as diameter of tibia, ventral setulae short as well as setulae 
between strong anterodorsals and posterodorsal. Basal tarsomere of fore leg thin and 
short setose, with short ventral spines and with peculiar long, fine and curved ventral 
setulae (these setulae distinct but shorter also on following tarsomeres), T1l: B11 = 
2.3-2.5, Bll: Blw = 4.2-5.0, basal tarsomere of mid leg short and thin, with short 
ventral spines and also with several curved ventral setulae, T21: B21 = 2.8-3.0, B21: 
B2w = 4.5-5.0. Basal tarsomere of hind leg thin, dorsally with 2-3 pairs of setae twice 
longer than diameter of this tarsomere and with short ventral spines, T31: B31 = 2.4-2.5, 


NEW EUROPEAN RHAMPHOMYIA 427 


B3l: B3w = 5.6-5.7. Wing brownish, stigma brown, veins brown, anal vein (Aj) 
complete. Costal seta long, ax angle sharply acute. M2/D = 1.6-1.7, M3/Db = 2.9-3.0, 
lw: ww = 2.9-3.1. Haltere brownish black, calypter brownish yellow with dark fringes. 
Abdomen brownish black, grey pruinose (brown in dorsal view). Setae and setulae all 
black. Hind marginal setae on sides of tergites 2-5 1.5 times as long as their segments, 
on tergites 6-7 very short, discal setulae shorter than hind marginals. Dorsum of 
tergites with short setae. The first sternite with single setula on each side. Terminalia 
as in Fig. 9; epandrium (=lateral genital lamella) oblong-ovate, with only fine setulae 
at dorsal part apically, strong setae occur in middle part; apical angle of cercus slightly 
obtuse. Length of body: 4.4-4.6 mm, wing: 5.2 mm. 
Female: Unknown. 


DIFFERENTIAL DIAGNOSIS: Rhamphomyia (s. str.) sellacrinita sp. n. is most allied 
to R. crinita Becker, 1887 (and also to at least three other still undescribed species) 
sharing with them a single row of posteroventral setae on fore femur confined to apical 
third of femur. However, it differs from all of them in the following combination of 
characters: fore basal tarsomere ventrally with a few peculiar, long and fine, upright 
setulae with bent tip; mesoscutum deep black (as in R. montana Oldenberg, 1915); 
wing brownish and fore tibia with homogeneous ciliation dorsally about as long as 
diameter of tibia. 


DERIVATIO NOMINIS. The species name is composed of: sella- (after type locality) 
and -crinita (stresses the similarity with R. crinita). 


DISTRIBUTION: Italy, Switzerland. 
DATES OF OCCURRENCE: August. 


KEY TO PALAEARCTIC SPECIES OF RHAMPHOMYIA (S. STR.) 
ALBOSEGMENTATA GROUP 


Notes: “additional characters” are useful for distinguishing keyed species from 
still undescribed species or species with which they can be confused or (in the case of 
females) from species of other Rhamphomyia (s. str.) species groups. Abbreviation: 
“M2/D” is not used in the key, instead of it, expressions like “vein M2 1.2-1.3 times as 
long as discal cell” are used. The key should be used together with the previously 
published keys and descriptions of Bartäk (1981, 1982, 1998) for checking illus- 
trations, detailed descriptions and further remarks. 


la Acrostichals multiserial (unknown female probably keys here) 

SAI ORRORE I SR DR R. piedmontensis sp. n. 
lb Acrostichals;bisenale rn U RN pie APM ENE nare ea OA 2 
2a (1) Dorsocentrals almost uniserial and 2-3 times longer and stouter than 

ac. (Additional characters: mesoscutum deep black, without any 

stripes; female with several anteroventral setae on hind tibia at least 

half as long as diameter of tibia; male fore femur with 1-5 strong pos- 

terior setae; male tergites 2-8 and female tergites 2-5 polished, other 

Pattsiofgabdomen;pruinose)/ EER e ee R. anthracina Meigen, 1822 
2b Dorsocentrals 2-4 serial, at most slightly longer and stronger than 

derostichalse.... PRI LOI NALI PIER DREH VI URRA ORTE n. 3 


428 


3a (2) 


3b 


4a (3) 
4b 


Sa (4) 
Sb 
6a (5) 
6b 


Ta (6) 


7b 


8a (7) 


8b 


9a (8) 


9b 


10a (7) 


10b 


M. BARTAK 


Mesoscutum with four longitudinal polished stripes. Female hind tib- 

ia slightly swollen, with anterior surface virtually bare 

NE NEE MUSSTE SEHEN Re ee ER R. luridipennis Nowicki, 1868 
Mesoscutum uniformly pruinose. Female hind tibia with another 


CITA ONY I FA TE ER MR E 4 
Malerei. Ii HE I RE OR SETTE 5 
Female (of R. lautereri, R. saintbaumensis, R. sellacrinita, R. ponti, 
Rerecta and R. bellinosetosa unknown)... OSE 32 
Posteroventral setae on fore femur absent. Mid femur with a group of 
smallksetaersubbasally rem ENDE R. parvicellulata Frey, 1922 
Posteroventral setae on fore femur present. Mid femur without any 
PTOUPIOMSEtad nr. ae le Se ie re 6 
Fore femur with a tuft of long anteroventral setae subapically on 
Gistinc&protuberancems EEE PER ie Re R. armimana Frey, 1922 
Anteroventral setulae on fore femur spread subequally, they do not 
formituftinai la er aller QRS SR ee ORTO 7 


Fore tibia with setae ventrally at least twice as long as tibia is thick. 
(Additional character: basal tarsomere of mid leg with setae dorsally 


atleast 4/5 asılongsas its length). 1. 2000 ce Se 8 
Fore tibia ventrally with homogeneous setulae shorter than diameter 
of.tibiar.. terrena e Re 0 D o io 0 6 © oo c 10 


Mid tibia with setae dorsally arranged in two rows of. Abdomen grey 
when viewed from above. Hind femur with a few setae ventrally about 
asjlons astemurisideepi RI ICI R. ponti sp. n. 
Mid tibia with very long setae both dorsally and ventrally not forming 
rows. Abdomen brown when viewed from above. Hind femur at least 
in basal two thirds with'short setae ventrally te "PORC CEE 9 
Mid femur with long anterodorsal and posterodorsal setae, the longest 
setae at least twice as long as femur is deep. Fore tibia not swollen, 
uniformly setose on posterior surface .......... R. ursina Oldenberg, 1915 
Mid femur with very short anterodorsal, dorsal and posterodorsal 
setulae, about one third as long as femur is deep. Fore tibia swollen, 
with bare pruinose stripe on posterior surface ...... R. kreischi Bartak, 1998 
Posteroventral setae on fore femur form a single row, which may be 
confined to apical half of femur only. Posteroventral (posterior) 
surface of fore femur polished at least above posteroventral setae or 
posteroventrals absent in basal half of femur eee 11 
Posteroventral surface of fore femur covered with setae and setulae 
not forming row or this row is irregular. Posteroventral area of fore 
femur pruinose and simultaneously posteroventrals present in basal 
halfofifemur 2:8: ool SM EEE 15 


lla (10) Posteroventral row of setae on fore femur incomplete, confined to 


11b 


apicaljhalfioffemur iii. 65 085. dee ISEE 12 
Posteroventral row of setae on fore femur complete, these setae are 
almost equally long throughout all length of femur .................. 13 


NEW EUROPEAN RHAMPHOMYIA 429 


12a (11) Basal tarsomere of fore leg ventrally with a few peculiar, long and 


12b 


fine, upright setulae with bent tips. Mesoscutum deep black. Wings 
brownish. (Additional characters: fore tibia with homogeneous 
ciliation dorsally about as long as diameter of tibia; abdominal tergites 
6-7 with very short hind marginals) ................ R. sellacrinita sp. n. 
Basal tarsomere of fore leg with normal setulae ventrally. Meso- 
scutum mostly light or brownish grey. Wing clear. (Additional charac- 
ters: both fore tibia and basitarsus dorsally with heterogeneous setae 
much longer than diameter of these tarsomeres) .. . . R. crinita Becker, 1887 


13a (11) Hind femur slightly bent in anterior view, with ventral «pilosity» 


13b 


continedito basalthalfionly i re oe R. lautereri Bartak, 1981 
Hind femur straight as usual, with ventral «pilosity» throughout its 
length. (If mid tibia with only one ventral row of setae, fore tibia with 
posterodorsal setulae about as long as tibia is thick, abdomen some- 
what subpolished, and apical angle of cercus slightly acute, compare 
RESUDAOLOMICA) SIEDO RIO Ra 14 


14a (13) Anteroventral setae on mid femur 1/3 as long as femur is deep or even 


14b 


shorter. (Additional characters: discal cell short, vein M2 1 .4-1.6 times 

as long as discal cell; abdomen brown viewed from above; epandrium 
Wwithishort: spines nearthetip) ar ar. an: R. montana Oldenberg, 1915 
Anteroventral setae on mid femur nearly as long as femur is deep. 
(Additional characters: posteroventral setae on mid femur subequally 

long as corresponding anteroventral - about as long as femur is high; 
posteroventral seta on hind femur present; proepisternum with normal 
setulae; discal cell slightly elongated; abdomen light grey viewed 

from above). (If discal cell about 1.1 times as long as vein M2, 
compare R. aucta; if vein M2 more than 1.4 times as long as discal 

cell, acrostichals and dorsocentrals more than twice as long as the dis- 

tance between their rows, and apical angle of cercus obtuse, compare 

R. hirtimana; if dorsal setae on mid tibia short, about twice as long as 

tibia is thick, compare R. sanctimauritii) 

RTE NN ANS EEE BRATEN. R. albosegmentata Zetterstedt, 1838 


15a (10) Discal cell elongated and narrow, about as long as vein M2. Apical 


15b 


angle of cercus obtuse. Fore femur short setulose dorsally. Hind femur 
with 1 long and strong posteroventral. Abdomen light grey viewed 
from above. (If discal cell elongated and other characters disagree, 


compare À. serpentata or R. nubigena) ......... R. aucta Oldenberg, 1917 
Discal cell shorter, vein M2 usually more than 1.1 times as long as 
discalcell Othencharactersidisagree a PP PEN eee ee 16 


16a (15) Abdomen grey when viewed from above. In difficult cases simulta- 


neously: basal tarsomere of fore leg above with setae longer than half 
of its length, mid tibia with two ventral rows of setae, discal cell short 
- vein M2 more than 1.3 times as long as discal cell, the longest dorsal 
setae on mid tibia at least three times as long as tibia is thick and face 
alwaysibaret mesta e RSA A EE Le MPS eee i, 17 


430 


16b 


17b 


M. BARTAK 


Abdomen brown when viewed from above. In difficult cases basal tar- 
somere of fore leg above mostly with setae shorter than half of its 
length and: either mid tibia with one ventral row of setae or discal cell 
elongated - vein M2 less than 1.2 times as long as discal cell or face 


with a few setulae or mid tibia with short setae dorsally .............. 23 
17a (16) Hind femur with at most 1-2 fine anteroventral setae shorter than 

femur is deep. Posteroventral setae on hind femur present or absent ..... 18 

Hind femur with 3 or more strong anteroventral setae about as long as 

femur is deep. Posteroventral seta on hind femur always present........ 19 


18a (17) Mesoscutum light grey when viewed from above, often with darker 


18b 


longitudinal stripes between rows of setae. Posteroventral setae on 
mid femur up to twice as long as femur is deep. Abdominal tergites 
4-7 with very short setulae. Costal seta often short and fine. Base of 
wing somewhat milky white ................ R. chionoptera Bezzi, 1904 
Mesoscutum brown when viewed from above. Wings not milky white. 
Posteroventral setae on mid femur shorter than femur is deep. Other 
characterssdisasteekn see ay M: R. crassimana Strobl, 1898 


19a (17) Basal tarsomere of fore leg swollen, 3.5 times as long as broad (and 


19b 


0.21 mm broad), densely covered with long setae dorsally. Also the 

2nd fore tarsomere dorsally with setae longer than this tarsomere. 
Phallus of unusual shape, with lateral «wings». (Additional charac- 

ters: larger species - wing 6.0 mm; scutum dark brownish grey) 

Ae ERR ate Sp eh a ao se ito R. saintbaumensis sp. n. 
Basal tarsomere of fore leg not swollen, more than 4 times as long as 

broad (and usually about 0.15 mm broad), sparsely covered with 
shorter setulae dorsally. The 2nd fore tarsomere dorsally short setulose 

(only preapicals may be longer). Phallus of usual «albosegmentata» 

ESE A PS e Le SRT AN EME LR NE N en > io uno: 20 


20a (19) Apical angle of cercus right to obtuse. Acrostichals and dorsocentrals 


20b 


shorter and stronger, about twice as long as the distance between their 
rows (rarely up to 2.5 times as long). Dorsocentrals sparsely spreading 
down sides of mesoscutum (10-20 setae in presutural area of meso- 
scutumilateral'ofidorsocentrals)£ e er ee ea ee 21 
Apical angle of cercus acute. Acrostichals and dorsocentrals long and 
fine, about three times as long as the distance between their rows and 
setae densely cover even the area lateral from dorsocentrals in front of 
SUGUTE LY OR EINE CRE SR eee 22 


21a (20) Fore tibia with strong anterodorsal, posterodorsal and posterior setae 


21b 


clearly differentiated from nearby setulae. About 20 setae in presutural 
area of mesoscutum lateral of dorsocentrals. Labrum shorter than head 
is high. Palpi densely long setose (more than 20 strong setae on each 
palpus) Hear u Re se se a R. bellinosetosa sp. n. 
Fore tibia with almost homogeneous dorsal ciliation. About 10 setae 
in presutural area of mesoscutum lateral of dorsocentrals. Labrum 
longer than head is high. Palpi sparsely setose (less than 15 strong 


NEW EUROPEAN RHAMPHOMYIA 43] 


setae on each palpus). (If fore tibia with rather short setae dorsally, 
palpi densely setose and apical angle of cercus very obtuse, compare 


RAJANOVERSIS) aA BE ENT 2 R. hirtimana Oldenberg, 1922 
22a (20) Apical angle of cercus about 45° ......... R. curvinervis Oldenberg, 1915 
22b Apical angle of cercus about 80° ........... R. tristriolata Nowicki, 1868 


23a (16) Posterodorsal setae on fore tibia almost homogeneous, about as long 

as diameter of tibia. Discal cell short, vein M2 1.6-1.8 longer than dis- 

cal cell (if discal cell elongated, compare R. sanctimauritii). Apical 

angle of cercus sharply acute. (If nearly right, compare R. sub- 

dolomitica). Both acrostichals and dorsocentrals relatively short, not 

much longer than the distance between their rows 

I ep get Sts. NNER SEW MABE rel pr R. brevipila Oldenberg, 1922 
23b Fore tibia with heterogeneous setae dorsally longer than tibia is thick. 

@thenicharactersidisagree ME Ben RO e 24 
24a (23) Fore femur dorsally and anterodorsally with only short setulae. (Addi- 

tional characters: abdomen light grey in lateral view, pruinose; face 


andsironsibare)r rn. EE, SER co REES ET 25 
24a Fore femur with setulae dorsally or anterodorsally at least as long as 
TCMUMRISIACEP ARES er ete Poe N 211 


25a (24) Mid tibia with short setulae dorsally, at most twice as long as tibia is 
thick. Larger species (wing more than 7.5 mm). (Additional charac- 
ters: apical angle of cercus obtuse; discal cell slightly elongated, vein 
M2 1.1 - 1.3 times as long as discal cell; mesoscutum with distinct 
stripes; posteroventral setae on mid femur strong and about 0.20 mm 


ONE Sa tae R. sanctimauritii Becker, 1887 
25b Mid tibia with longer setulae dorsally, at least three times as long as 
diameter OLE DIARI AR ETS PRET E NEED IE. MIU. (EN ae A OA 26 


26a (25) Apical angle of cercus slightly acute. Discal cell short (vein M2 

usually more than 1.4 times as long as discal cell) 

CERANO TAI MR I R. subdolomitica Bartäk, 1981 
26b Apical angle of cercus very obtuse. Discal cell slightly elongated (vein 

M2 1.2-1.3 times as long as discal cell) ........ R. janovensis Bartäk, 1981 
27a (24) Apical angle of cercus acute. Abdominal tergites pruinose, light grey 

in lateral view. Mid tibia with two ventral rows of setae. Eyes meet on 

frons. Face bare. Discal cell slightly elongated, vein M2 less than 1.2 

times as long as discal cell. (If discal cell short, acrostichals and dor- 

socentrals twice as long as the distance between their rows and only 1 

posteroventral on f3 present, compare R. luridipennis) 

RP tee Peat OE I SANE SRI eS RA nt Et DURE à ADONE I R. serpentata Loew, 1856 
27b Apical angle of cercus right or slightly obtuse. Abdominal tergites at 

least moderately subpolished (if pruinose, follow couplets 13 or 20). 

Face often with a few setulae. Other characters in another combination . . . 28 
28a (27) Basal tarsomere of fore leg dorsally with setae longer than half the 

length of basitarsus. Mid femur with anteroventral and posteroventral 

setae longer than femur is deep. Mid tibia with two complete rows of 


432 M. BARTÀK 


setae ventrally. (Additional character: mesoscutum pruinose, without 


Sthipes) ge eterea alate a 29 
28b Basal tarsomere of fore leg dorsally with setae shorter than half of its 

length. Mid femur with at least anteroventral setae shorter than femur 

is deep. Mid tibia usually with only one ventral row of setae........... 30 
29a (28) Abdominal tergites subpolished, not glossy polished. The 8th tergite 

without “brush” of erect setae dorsally ........... R. lindneri Bartak, 1998 
29b Abdominal tergites glossy polished at least partly. The 8th tergite with 

peculiamabrush? olerectsetae me re ee R. erecta Bartak, 1998 


30a (28) Hind femur with dense and long anteroventral setulae, subequally 
long as strong setae. Tergites 2-7 polished, other parts of abdomen 
pruinose. Acrostichal setae distinctly shorter and finer than dorso- 


centralsprare Verne R. loewi Nowicki, 1868 
30b Hind femur with short and sparse anteroventral setulae in addition to 

much longer and stronger setae. Tergites at most subpolished, with 

distinct microchaetae. Acrostichal and dorsocentral setae subequal ...... Sill 


31a (30) Mesoscutum subpolished, without traceable stripes. Epandrium 
equally rounded at tip. Apical angle of cercus slightly obtuse 
N SIDLEY DL SG Co ALO ER TEE TEE TEE R. discoidalis Becker, 1889 
31b Mesoscutum pruinose, with two distinct darker stripes between rows 
of setae. Epandrium not rounded at tip, but with distinct dorsal 
constriction. Apical angle of cercus exactly right one. (Additional 


characters: abdominal tergites 2-6 pruinose) ..... R. morio Zetterstedt, 1938 
32a (4) Discal cell elongated almost to wing margin, more than 2.5 times as 

longaasivein\ M2 HH. Ma ie ees «cies ee eee 33 
32b Discal cell shorter, at most twice as long as vein M2................ 34 


33a (32) Discal cell of unusual shape, vein bordering its lower side clouded 
Perrys Senta Wai Teil Soa atea ati R. serpentata Loew, 1856 


33b Discal cell of usual shape. Hind femur anteriorly with an irregular row 

of setae nearly as long as femur is deep ........ R. aucta Oldenberg, 1917 
34a (32) Wing partly clouded, at least along apical sections of veins M2 and 

Curette Ro rata LE eset ER A 400.0 0.000 0 35 
34b Wing uniformly coloured, clear to brown. (If the whole posterior 

margin of wings clouded, compare R. discoidalis) .................. 36 
35a (34) Apical section of vein MI clouded ......... R. tristriolata Nowicki, 1868 


35b Apical section of vein M1 without clouding . . R. curvinervis Oldenberg, 1915 
36b (34) Both acrostichals and dorsocentrals shorter than 1/3 of the distance 
between their rows. Hind femur without any strong anterior setae in 


basallhallae an Bau seen venia ii RRS ee 00 0 0 à 0 5 0 o 37 
36b Acrostichals or dorsocentrals longer than 1/3 of the distance between 
their rows or hind femur with strong anterior setae even in basal half . . . . 38 


37a (36) Hind femur somewhat dilated, covered with short and almost pennate 
ciliation both above and beneath in addition to 4-6 strong antero- 
ventral setae. Discal cell not elongated, vein M2 1.3-1.7 times as long 
as discal cell. (Additional characters: mesoscutum dark black; acro- 


NEW EUROPEAN RHAMPHOMYIA 433 


stichals and dorsocentrals stout, spinose). (If acrostichals and dor- 

socentrals fine and hair-like, compare R. brevipila) 

EINE MERO IAT EROI R. montana Oldenberg, 1915 
37b Hind femur not dilated and without subpennate ciliation. Discal cell 

elongated, vein M2 less than 1.2 times as long as discal cell. 

(Additional character: wings brownish, discal cell and the area behind 

it and second basal cell conspicuously lighter than other parts of wing 


MCIMOLANG Ae. ea Or ee RL R. discoidalis Becker, 1889 
38a (36) Basal three fore tarsomeres somewhat dilated, covered with fine, 
SUVeRPILOSIlY ages ne a Pein ah ol R. crassimana Strobl, 1898 
38b Front tarsi shaped and setose as usually. (Additional character: mid 
femuUrstraishtnOtbent)r mr NITRO 39 
39a (38) Discal cell elongated, vein M2 less than 1.2 times as long as discal cell . . 40 
39b Discal cell short, vein M2 more than 1.2 times as long as discal cell. 
(Additional characters: face bare; ocellar setae long and strong, 
usuallyslonger than 2B80r tons)e...... oo a rn: 44 


40a (39) Face with a few long fine setulae. (Additional character: abdomen 
pruinose). (If mesonotum subpolished and without stripes, acro- 
stichals short and posterior margin of wings dark and discal cell much 


paler; compare Ri discoidalis) 2... ...........: R. morio Zetterstedt, 1938 
40b aces Dateien han Mt onan tern I ORTO, 41 
41a (40) Abdomen light grey. (If fore femur pruinose and not at least polished, 

tolowscouplet 44) 2. ere o R. albosegmentata Zetterstedt, 1838 
41b Abdomenibrown,atleast onstersitesn na ee 42 


42a (41) Fore tibia without rows of setae ventrally. Mid femur with setae ven- 
trally 1/3 as long as femur is deep. Abdominal tergites subpolished 
N Or e e ee eR R. lindneri Bartäk, 1998 


42b Fore tibia with distinct rows of ventral setae (sometimes very short). 
Mid femur short setulose ventrally. Abdominal tergites 2-7 polished . .... 43 
43a (42) Both tergites and sternites of abdomen polished, at least sides of 
SEINS Gite wie ee ct Li R. ursina Oldenberg, 1915 


43b Sternites pruinose, at most last segments slightly subpolished. (If fore 
tibia with very short ventral setulae and fore coxa with anterior and 
anterodorsal setulae longer than diameter of fore tibia, compare R. 
LOCI igre ta oe easy Re RICCO Wis PRT Aa dE R. kreischi Bartäk, 1998 
44a (39) Hind femur somewhat dilated, with short and almost subpennate 
ciliation above and sometimes also beneath, in addition to 2-5 strong 
anteroventralsspinesen urteilen MPa A re 45 
44b Hind femur without subpennate ciliation and not at least dilated ........ 46 
45a (44) Many (usually more than 20) setulae lateral of dorsocentrals in pre- 
sutural part of mesoscutum. Both acrostichals and dorsocentrals usu- 
ally shorter than half of the distance between their rows. Mid femur 
with short subpennate posteroventral ciliation. Abdomen grey dusted 
CREDO ai R. brevipila Oldenberg, 1922 


434 M. BARTÀK 


45b Almost no setulae lateral of dorsocentrals. Both acrostichals and 

dorsocentrals usually longer than half of the distance between their 

rows. Posteroventral surface of mid femur sparsely covered with 

simple (not flattened) setulae shorter than half of depth of femur. 

Tergal part of abdomen slightly polished . . . . R. subdolomitica Bartäk, 1981 
46a (44) Marginal setae on abdominal segment 3 about 1/4 as long as this 

segment, following segments almost bare. Costal seta short and fine, 

almost absent. Hind femur almost bare ventrally . R. chionoptera Bezzi, 1904 
46b Abdominal segment 3 with hind marginal setae longer, also the 

following segments with considerable pubescence. Costal seta long. 

Elindifemuriwithwventralisetae ARRE, eee eee 47 
47a (46) Tergites 2-7 polished, sternites pruinose. Mid and hind femora with 

very short setulae and very short ventral spines . . . . R. loewi Nowicki, 1868 
47b Not as above. (Additional character: hind femur with long and strong 

Setae ventrally, ia LL 1 10T N 48 
48a (47) Hind femur with a complete row of strong anterior (or anterodorsal) 

setae. Both acrostichals and dorsocentrals usually shorter than half of 

the distance between their rows. (Additional character: large species, 


Winesmoreithanio 9 MM), a ese ee acer R. sanctimauritii Becker, 1887 
48b Strong anterior or anterodorsal setae absent at least in basal half of 

hind femur. Both acrostichals and dorsocentrals usually longer than 

the distance between theinrowse 5.15 er. u LR ORE 49 


49a (48) Hind femur thickened apically. Mid femur and tibia covered with pe- 

culiar, rather adhered and thick pubescence. (Additional character: 

posteroventral surface of fore femur bare and polished) 

NET EN. MRO RA re re R. parvicellulata Frey, 1922 
49b Hind femur not thickened. Mid leg with ordinary pubescence. (Addi- 

tional characters: both acrostichals and dorsocentrals less than twice 

as long as the distance between their rows; tergites uniformly 

DIUINOSC) EA Tota Re SEITE IA Te SITO IR FITTE, 50 
50a (49) Usually more than 30 acrostichal setae. Hind femur usually with 3 or 

more anterodorsal setae or fore femur with strong anterior seta. 

Lhaltere dark AC" SSI CERN dee tte PSE PRE EEE Sil 
50b Usually less than 30 acrostichal setae. Hind femur with at most 2 

anterodorsal setae and fore femur usually without anterior seta. 

Haltere yellow to dark. (Additional characters: femora pruinose; fore 

femur with distinct posteroventral setulae; scutum usually with more 

than 10 setulae in presutural area beside intrahumeral and post- 

humeral; dorsocentrals multiserial; hind tibia with ordinary ciliation) ....52 
Sla (50) Mid femur with one strong posterior seta in apical third 

RR DEREN Ree eh RE 3 op es cide Bi o R. armimana Frey, 1922 
51b Mid femur without strong posterior seta in apical third 

RE QUE LA SON MC PS RA GAS Ve ENS R. janovensis Bartäk, 1981 
52a (50) Acrostichals rather stout. Haltere pale, often almost yellow. Abdomen 

ereysviewedifromiabove.! etto R. hirtimana Oldenberg, 1922 


NEW EUROPEAN RHAMPHOMYIA 435 


52b Acrostichals fine. Haltere dark. Abdomen brown viewed from above. 
(Additional character: fore femur pruinose). (If acrostichals reach far 
into presutural area, compare R. janovensis)....... R. crinita Becker, 1887 
ACKNOWLEDGEMENTS 


This paper was supported by IRP MSM 6046070901 and by NAZV QH72151. 
I thank Bernhard Merz (Geneva), Adrian Pont (Oxford), Hans-Peter Tschorsnig 
(Stuttgart), and Jeffery Cumming (Ottawa) for providing valuable specimens for this 
study. My special thanks are due to Bernhard Merz for many valuable comments to 
earlier versions of the manuscript. I thank also my wife for drawing the illustrations. 


REFERENCES 


BARTAK, M. 1981. A revision of the Rhamphomyia albosegmentata - group (Diptera, Empi- 
didae), with descriptions of new species. Acta Universitatis Carolinae - Biologica 
1979: 361-407. 


BARTAK, M. 1982. The Czechoslovak species of Rhamphomyia (Diptera, Empididae), with 
description of a new species from Central Europe. Acta Universitatis Carolinae - 
Biologica 1980 (1982): 381-461. 

BARTAK, M. 1998. Rhamphomyia (Diptera: Empididae) from the State Museum of Natural 
History, Stuttgart, with descriptions of new species. Stuttgarter Beiträge zur 
Naturkunde, Serie A 583: 1-26. 

MERz, B. & HAENNI, J.-P. 2000. 1.1. Morphology and terminology of adult Diptera (pp. 21-51). 
In: Papp, L. & DARVAS, B. (eds). Contributions to a Manual of Palaearctic Diptera. 
Volume 1. Science Herald, Budapest, 978 pp. 

SINCLAIR, B. 2000. 1.2. Morphology and terminology of Diptera male terminalia (pp. 53-74). In: 
Papp, L. & DARVAS, B. (eds). Contributions to a Manual of Palaearctic Diptera. Volume 
1. Science Herald, Budapest, 978 pp. 


BIUTERINA SPP. FROM THE IVORY COAST 437 


ERRATUM 


Georgiev, Boyko B. & Mariaux, Jean 2007. Cestodes of the genus Bluterina 
Fuhrmann, 1902 (Cyclophyllidea: Paruterinidae) from passeriform and piciform 
birds in the Ivory Coast, with a key to the species of the genus. Revue suisse de 
Zoologie 114 (1): 141-174. 


Due to a technical problem, the symbol > was rendered incorrectly in the key to the 
species of the publication cited above. The correct key is therefore printed again below. 


KEY TO THE SPECIES OF BIUTERINA 


la. 
lb. 
2a. 


2b. 


3a. 


sb: 


4a. 
4b. 


5a. 
5b. 


6a. 


6b. 


7a. 
7b. 
TE: 
8a. 
8b. 
9a. 
9b. 


10a. 


10b. 


lla. 
11b. 


Parasites of Paradisaeidae in the Australian Region .................... 2 
Barasıtes;of ONE DAS eee see (ates Eee 3 
Testes -30 in number. Genital atrium without distinct dilatory muscles 
ARI LA ee bee B. clavula 
Testes -15 in number. Genital atrium with distinct dilatory muscles 

areas Ba in dello DATE taz ae ete B. mertoni 
Anterior rostellar hooks < 17 long; if anterior rostellar hooks 17 long 
thenstotalmumberoof.rostellar.hooks = 32... .n.0... e TOR rrE 4 
Anterior rostellar hooks > 17 long; if anterior rostellar hooks 17 long or 
slightly longer then total number of rostellar hooks 2 40 ................ 5 
Rostellarihooks 2%in number crs zn ne ita sted ce B. globosa 


Rostellar hooks 30-32 in number 
ee re o AR Sta eS B. motacillabrasiliensis and B. motacillacayanae 


Uterus entirely embedded in paruterine organ in pregravid proglottides ..... 6 
Paruterine organ situated anterior to uterus in pregravid proglottides, not 

entirelyiembeddine it E44 yeasts Seel ve ne Na See 8 
Developing uterus horseshoe-shaped, with posteriorly directed lateral 

CAremies fee Scene en oe es 7 
Developing uterus consisting of two rounded lateral chambers 

connected anteriorly by narrow, branched isthmus ............ B. passerina 
Rostellar hooks 26-30 in number; anterior hooks = 56 long ...... B. cordifera 
Rostellar hooks 56-62 in number; anterior hooks 24-25 long . . . . B. pogoniuli 
Rostellar hooks 40-42 in number; anterior hooks 20 long ....... B. chlorurae 
Anterrogrostellarinooks=> Sanlone kur. der aii ee eee 9 
Anterior rostellarhooks:< 35 long... RR 21 
Numberofrostllanhooks <20 an Bene ers ne 10 
Numberotrostellamhooks= 26868 ANR SE: 11 
Number of rostellar hooks 14; anterior rostellar hooks 56-60 long; cirrus 

SACK AON OMS rte die RL ES ÈS B. erithaci 
Number of rostellar hooks 18-20; anterior rostellar hooks 44-48 long; 

Cirrusisac ~90 long. Fer esse reise B. fallax 
Anterior tostellar hooks 75-83 lone N. nun. ie e ie) ee B. africana 


Antenorrostellawhooks: = 70) longs erring sae a re Se: 12 


438 


12a. 
12b. 
182% 


13b. 


14a. 
14b. 
15a. 
15b. 
16a. 


16b. 


17a. 
17b. 
18a. 


18b. 


19a. 


19b. 


20a. 
20b. 


2la. 
21b. 
22a. 
22b. 
23a. 
23b. 
24a. 


24b. 
25a. 
D0). 
26a. 


26b. 
26c. 


B. B. GEORGIEV & J. MARIAUX 


Anterior rostellar hooks 58-66 long... 13 
Anterionrostellar hooks 38-49 long”... 4. Re 2. 22. a 2 ee 14 
Rostellar hooks 34-36 in number; testes 15-18 in number; cirrus sac 86- 

OS Kay ee IO A016 0:0 0: B. morgani 
Rostellar hooks 40-44 in number; testes 6-9 in number; cirrus sac > 120 

ONE IR DE B. triangula 
Rostellanhooksspine-ltké jets. PEN MEN ORSO B. lansdowni 
Rostellar hooks triangular, of typical shape for Biuterina ............... 15 
Rostellarhooksi<30%mmumber ... "NP 0 0 eee 16 
Rostellarhooks > 35 in number... 2m... CN eee 17 


Rostellar hooks 26 in number, anterior rostellar hooks 43-46 long; paru- 

terine organ wide, occupying almost entire width of median field 
Een white ee LIZA. dll TL DAAD ae B. campanulata 
Rostellar hooks -30 in number; anterior rostellar hooks 41 long; paru- 


terme organuin/middle ofimediantield es anta CERA B. trapezoides 
Anteriomrostellar hooks. <4510ng 7.7.2. MR CT TC CRE 18 
Anteriomrostellamhooksi>45 long TIL 2025) IRR LEE ARE 19 


Gravid strobila consisting of -180 proglottides; cirrus sac crosses poral 
osmoregulatory canals and substantial part of it situated in median field 
DR RE ce ta tette e E men 25 e A RT ISA LORI B. macrancistrota 
Gravid strobila consisting of -50-60 proglottides; cirrus sac in lateral 


Mela e VPE BAUDO IRA 10 SCOUTED MAE AR ERBE B. petroniae 
Posterior rostellar hooks not longer than 55% of length of anterior ros- 
tellan/hooks it a RR, EUR LIRE. B. planirostris 
Posterior rostellar hooks longer than 60% of length of anterior rostellar 

NOOKSER AE DIARIO EU DEI SITR BO) OE ROUEN PIN SEE 20 
Blade of rostellar hooks 4-5 times shorter than handle . . . . . B. diversihamata 
Blade of rostellar hooks slightly longer than handle or of same length 

MRO: SCHNEE SUNT, CINI, IONI SOS, SL: DATE B. sobolevi 
Rostellanhooks ~20 inmumber 2.2.2 2222.22 222. ee B. distincta 
Rostellarhooks >'30in number’. .2...L SRE MIR FE 22 
Antenorstostellarhooks'’>30,0n2F 7 E. RR ARR eee B. reynoldsi 
Anterior rostellar hooks!<30 long 1... 20.0. E Eee 23 
Rostellar hooks >56innumber N... MEN ENT IE er CR 24 
Rostellar hooks’=&52 in number. 27 III SE 27 
Rostellum consists of weak musculature and strongly developed glan- 
AUIATATISSUE RR eme shakers: INA CE SCORE RENE EE B. quelea 
Rostellum consists of strongly developed musculature, with no glandu- 

lar tissue or glandular tissue weakly developed . ...................... 25 
Paruterine organ in pregravid proglottides occupies entire width of 
median held >: ss: pa Hanser B. ugandae 
Paruterine organ distinctly narrower than width of median field .......... 26 
Parasite of Furnariidae in the Neotropics (Brazil) .......... B. trigonacantha 
Parasite of Dicruridae in India. 1 422 S22. = Ae RE B. singhi 


Parasite of Eanndae in the Palacarctici M CON RUES B. collurionis 


27a. 
27b. 
28a. 
28b. 
29a. 
29b. 
30a. 


30b. 


BIUTERINA SPP. FROM THE IVORY COAST 439 


Rostellarhooks 40linnumber tt ac eee ce eee ne eae ees 28 
Rostellar hooks = 48 (possibly, as exception, 42 in B. fuhrmanni)........ 29 
Antenorrostellarhooks27410Ng EE > acca. eis ie ene B. meropina 
Anteniorrostellanhookss<20llongsr 2. 22... 2. 2.0. Ma B. dicruri 
il'estes number=il Zeta eee ARTO B. fuhrmanni 
Mestestn um ber AE at E IE 30 


Rostellar hooks 52 in number; anterior rostellar hooks 23-25 long 
E N PIT IT III B. cylindrica 
Rostellar hooks 48 in number; anterior rostellar hooks 19-22 long 
ME e E RARI A I n li ee goa B. zambiensis 


Lo “Ma Art 


iR 
A 


DTA 
Li few 


2 : i y At OW had Vial. DRAG ue | 
ar Al Hape : f hi (Bac h ‘y . N n 
i se (0 ele ee 2 
B De 4 > 


i u u K | di n M 7 
er ie is. sù ni 


Er 
? 
n° 


| 


REVUE SUISSE DE ZOOLOGIE 


Tome 114 — Fascicule 2 


SCHMITZ, Patrick & LANDRY, Bernard. Two new species of Chionodes 
Hübner from Ecuador, with a summary of known Galapagos records 
onGelechidael(Éepidoptera) s.r ORO GONO 

MERZ, Bernhard. A new Sapromyza Fallen, 1810, from the Swiss Alps 
(Diptera Eauxantidae) ana in RIINA 

SCHUCHERT, Peter. The European athecate hydroids and their medusae 
(Eiydrozoar Cnidaria): Eilitera,Part 22 LL E 

ZIEGLER, Thomas, SCHMITZ, Andreas, HEIDRICH, Astrid, Vu, Ngoc Thanh 
& NGUYEN, Quang Truong. A new species of Lygosoma (Squamata: 
Sauria: Scincidae) from the Central Truong Son, Vietnam, with notes 
onats molecular phylogenetic position: 

BARTAK, Miroslav. Five new European species of the Rhamphomyia 
(s. str.) albosegmentata group (Diptera: Empididae)............. 

Erratum. GEORGIEV, Boyko B. & MARIAUX, Jean 2007. Cestodes of the 
genus Bluterina Fuhrmann, 1902 (Cyclophyllidea: Paruterinidae) 
from passeriform and piciform birds in the Ivory Coast, with a key to 
the species of the genus. Revue suisse de Zoologie 114 (1): 141-174. 


Pages 


175-184 


185-194 


195-396 


397-415 


417-435 


437-439 


REVUE SUISSE DE ZOOLOGIE 


Volume 114 — Number 2 


SCHMITZ, Patrick & LANDRY, Bernard. Two new species of Chionodes 
Hübner from Ecuador, with a summary of known Galapagos records 
ora@elechisdaeillEepidoptera)... .-....... ee EEE 

MERZ, Bernhard. A new Sapromyza Fallen, 1810, from the Swiss Alps 
(Diptera sica uxa n dae). en E 

SCHUCHERT, Peter. The European athecate hydroids and their medusae 
(Elyvdrozoa Cnidaria)=Filifera Part 27,2. E22 RO ee 

ZIEGLER, Thomas, SCHMITZ, Andreas, HEIDRICH, Astrid, Vu, Ngoc Thanh 
& NGUYEN, Quang Truong. A new species of Lygosoma (Squamata: 
Sauria: Scincidae) from the Central Truong Son, Vietnam, with notes 
onsts molecular phylogenetic positions 724s eee LIA 

BARTAK, Miroslav. Five new European species of the Rhamphomyia 
(s. str.) albosegmentata group (Diptera: Empididae) ............. 

Erratum. GEORGIEV, Boyko B. & MARIAUX, Jean 2007. Cestodes of the 
genus Bluterina Fuhrmann, 1902 (Cyclophyllidea: Paruterinidae) 
from passeriform and piciform birds in the Ivory Coast, with a key to 
the species of the genus. Revue suisse de Zoologie 114 (1): 141-174. 


Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX 


Pages 


175-184 


185-194 


195-396 


397-415 


417-435 


437-439 


PUBLICATIONS DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE 


CATALOGUE DES INVERTEBRES DE LA SUISSE, N08 1-17 (1908-1926) ........ serie Fr. 


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REVUE DE PALEOBIOLOGIE ............ RR Echange ou par fascicule Fr. 
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THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND 
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TINOQISS RISE ORIANA ee PR Fr. 


CLASSIFICATION OF THE DIPLOPODA 


RSIERFIOEEMANG 2310 PMO EINE Fr. 


LES OISEAUX NICHEURS DU CANTON DE GENÈVE 
P. GEROUDET, C. GUEX & M. MAIRE 


351 ps nombreuses cartes et figures, 1983 20 he bins sie es sien |) eee RUE Fr. 


CATALOGUE COMMENTE DES TYPES D'ECHINODERMES ACTUELS 
CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
A LA CONNAISSANCE DES ECHINODERMES ACTUELS 


MÉTANGOUX Grp lo SS ae er OA sehen aula Re on ee Fr. 


RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE 
(COTENTIN-BAIE DE SEINE, FRANCE) 


Yo FINET; J. WUEST' & K. MAREDA, 62 p., 1991 - 2. nee. Fr. 


GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: 
SHELLS AND RADULAS 
YAEINETJEWUESTI& KE MAREDAMT OOD PP E Fr. 


O. SCHMIDT SPONGE CATALOGUE 


R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 p., 1992...................... se Er: 


ATLAS DE REPARTITION DES AMPHIBIENS 
ET REPTILES DU CANTON DE GENEVE 


A. KELLER, V. AELLEN & V. MAHNERT, 48 p., 1993 ............................ Fr. 


THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 
A DOCUMENTED FAUNAL LIST 


N=JEINET;2180:P:31995: nce Fr. 


NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 


TECACATIIEZ Op MO SE ee ee ehe Fr. 


PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS 
OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... Fr 


CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) 


(Instrumenta Biodiversitatis I), I. LOBL, xii + 190 p., 1997 ...................... Fr. 


CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) 
DE LA REGION AFROTROPICALE 


(Instrumenta Biodiversitatis II), H. G. DIRICKX, x +187 p., 1998 ..... ............ Fr. 


A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE 
WEST PALAEARCTIC REGION 


(Instrumenta Biodiversitatis III), S. BOWESTEAD, 203 p., 1999 ................... Fr. 


THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 
(Instrumenta Biodiversitatis IV), B. SCHATTI & A. DESVOIGNES, 


EPG RA TI e a ANI Fr. 


PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY 
(Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS, 
ITB TAS Da PAU Pig aco paths RE O TOR ARE RSA Fr. 


REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA 
THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) 
(Instrumenta Biodiversitatis VI), G. SABELLA, CH. BUCKLE, V. BRACHAT 
CCCIBESUCHET VICE 83 PEZZO RE OTO IO Fr. 


PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID FLIES 
(DIPTERA, TEPHRITOIDEA) 
Proceedings of the “3rd Tephritoid Taxonomist’s Meeting, Geneva, 19.-24. July 2004” 


(Instrumenta Biodiversitatis VII). B. MERZ, vi + 274 p., 2006 Fr. 


285.— 


35.— 
35.— 


30.— 


30.— 


ASD 


15.— 


10.— 


. 160.— 


60.— 


70.— 


180.— 


100.— 


100.— 


7 ö ee 


A drain Pitre 


Oe 
2, 3. 1 val 
WE AGLI LAI 


4 
M x 
22 de OI rat pr Dee rw: Br 


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listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated 
for each reference. The title of the paper and the name of the journal must be given in full in the following style: 
PENARD, E. 1888. Recherches sur le Ceratium macroceros. These, Geneve, 43 pp. 

PENARD, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61. 
MERTENS, R. & WERMUTH, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. 
HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: WENZEL, R. L. & TIPTON, V. J. 

(eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XI + 861 pp. 

Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit 
122 x 180 mm, each table on a separate sheet and numbered consecutively. 

Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered 
consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering 
should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain 
from mixing drawings and half tones. Originals of figures (ink drawings, photographs, slides) should be submitted 
together with the revised version of the accepted manuscript. Original drawings will not be returned automatically. 
The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If scanned figures 
are submitted on CD, this should be clearly indicated on the print-out. Scanned line drawings must be saved as TIF files 
in bitmap mode with a resolution of at least 600 dpi. Half tone illustrations and photos must have at least 300 dpi reso- 
lution. 

Legends to figures. These should be typed in numerical order on a separate sheet. 

Proofs. Only page proofs are supplied, and authors may be charged for alterations (other than printer’s errors) if 
they are numerous. 

Offprints. The authors receive a total of 25 offprints free of charge; more copies may be ordered at current prices 
when proofs are returned. 

Correspondence. All correspondence should be addressed to 


Revue suisse de Zoologie, Muséum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. 
Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: danielle.decrouez@ville-ge.ch 
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 


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