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REVUE SUISSE DE ZOOLOGIE

SWISS JOURNAL OF ZOOLOGY

g 3
kl GENEVE JUIN 2006 ISSN 0035 - 418 X

REVUE SUISSE DE ZOOLOGIE

TOME 113—FASCICULE 2

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

DANIELLE DECROUEZ
Directrice du Muséum 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: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.

Administration

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1211 GENÈVE 6

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REVUE SUISSE DE ZOOLOGIE 113 (2): 229-238; juin 2006

Descriptions of three new Platypalpus Macquart from Guangdong,
China (Diptera, Hybotidae, Tachydromiinae)

Ding YANG! , Bernhard MERZ? & Patrick GROOTAERT*

l Department of Entomology, China Agricultural University, Beijing 100094, China.
E-mail: dingyang@cau.edu.cn

2 Key Lab of Insect Evolution & Environmental Changes, Capital Normal University,
Beijing 100037, China.

3 Département d'Entomologie, Muséum d'histoire naturelle, C. P. 6434,
CH-1211 Genève 6, Switzerland.
E-mail: bernhard.merz@ville-ge.ch

4 Department of Entomology, Royal Belgian Institute of Natural Sciences,
Vautierstraat 29, 1000 Brussels, Belgium.
E-mail: Patrick.Grootaert@naturalsciences.be

Descriptions of three new Platypalpus Macquart from Guangdong,
China (Diptera, Hybotidae, Tachydromiinae). - Platypalpus Macquart is
recorded for the first time from Guangdong province. Three new species are
described as new to science: P. convergens sp. n., P. guangdongensis sp. n.,
P. zhangae sp. n. A key to the species of the genus from Guangdong is
presented.

Keywords: Hybotidae - Tachydromiinae - Platypalpus - Guangdong - new
species.

INTRODUCTION

The genus Platypalpus Macquart belongs to the subfamily Tachydromiinae of
the family Hybotidae. It is easily identified by the following combination of characters:
eyes narrowly separated on frons and face, mid leg raptorial, and anal cell present
(Chväla, 1975; Grootaert & Chväla, 1992). The genus is distributed worldwide. Most
species have been recorded from the Palaearctic realm, particularly from Europe where
some 250 species are known (Chvala & Kovalev, 1989; Grootaert & Chväla, 1992).
Smith (1975) reported 31 species from the Oriental Region. However since then, 28
species have been added to the fauna of China alone by Yang & Merz (2005), Yang,
D. & Yang, C. (1989, 1990, 1992, 1997), Yang, C. & Yang, D. (1992), Saigusa & Yang
(2002), and Yang, An & Gao (2002). So far 36 species are known to occur in China.
The major reference dealing with Platypalpus are as following: Brunetti (1920), Frey
(1943), Smith (1965), Chväla (1975, 1989), Grootaert (1984), and Grootaert & Chväla
(1992)

Manuscript accepted 15.06.2005

230 D. YANG ET AL.

Guangdong with a tropical or subtropical climate belongs to the South China
Region of the Oriental realm north of Hongkong. The genus Platypalpus is here
recorded for the first time from Guangdong. Three new species are described and
illustrated. In addition two unnamed species, known from females alone, were found
during the sampling campain of Yang & Grootaert in 2003 in Guangdong. More
species are expected to occur in this province since the sampling was done in the
beginning of the flight period of Platypalpus. A key to the three described species from
Guangdong is presented.

MATERIAL AND METHODS

The specimens were collected at the end of May by Yang, Zhang & Grootaert
respectively. The type material is deposited in the following collections:

CAU Entomological Museum of the China Agricultural University, Beijing, China
MHNG Muséum d'histoire naturelle, Genève, Switzerland
RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium

In order to allow an easy comparison with descriptions of various authors, the
morphological terminology follows generally Grootaert & Chväla (1992), except for
crossveins which are written in capitals. Terminology of antenna follow Stuckenberg
(1999).

The following abbreviations for bristles are used: acr-acrostichial, ad-
anterodorsal, av-anteroventral, dc-dorsocentral, h-humeral, oc-ocellar, npl-notopleural,
pd-posterodorsal, presc — prescutellar, psa-postalar, pv-posteroventral, sa-supraalar sc-
scutellar, vt-vertical.

TAXONOMIC PART
KEY TO THE DESCRIBED SPECIES OF PLATYPALPUS FROM GUANGDONG

The following key should help to separate quickly the three new species.
Because of insufficient knowledge of the Platypalpus fauna of China a large numer of
additional species may be expected to occur in this province.

1 Mhoraxtblack# AM TEL AT DD PR TRIO AE 2
- ihoraxsyellow sees di Sse BIL. II LEO guangdongensis sp. n.
2 R4,5 and M parallel apically; arista as long as postpedicel . . . . . zhangae sp. n.

R4,5 and M convergent apically; arista much longer than postpedicel
MERO SIAMO: SILA PT MSA OI ERE convergens Sp. n.

Platypalpus convergens sp. n. Figs 1-5

MATERIAL: Holotype d, CHINA: Guangdong: Ruyuan, Nanling National Nature
Reserve, (23.20N/115.23E), Shumuyuan, 500 m, 26. II. 2003, leg. L. Zhang (CAU). Paratypes:
1 d 12 2 ©, same data as holotype (CAU & MNHG). 2 88,7 9 9, forest near entrance of the
Nanling National Nature Reserve, 26.III.2003, leg. P. Grootaert (sample 23008, RBINS). The
type locality is a forest situated about 40 km north of Ruyuan (24.44N/113.18E).

ETYMOLOGY: The specific name refers to R4,5 and M which are convergent
apically.

PLATYPALPUS FROM GUANGDONG 231

PS

RS
ww
ag!
um
M mm
m num
1 WAN OT

Fics 1-5

Platypalpus convergens sp. n, male. 1, wing; 2, tip of mid tibia; 3, genitalia, dorsal view; 4, left
epandrial lamella; 5, right epandrial lamella.

DIAGNOSIS: Mainly grey-microtrichose species; postpedicel yellow, 2.8 times as
long as wide, arista over twice as long; 1 pair of vt; hairs on mesonotum very short and
sparse; mid femur with a row of rather short pv; fore tibia thickened with shorter dor-
sal and very long, dense black ventral bristles in apical half; mid tibia with very long
apical spur (Fig. 2); fore basitarsus with long posterior and pv bristles; Ry,5 and M
convergent apically (Fig. 1).

DESCRIPTION:

Male. Body length 3.9-4.3 mm, wing length 4.8-4.9 mm.

Head black, grey microtrichose. Eyes narrowly separated on face; frons wider
than face, parallel-sided, as wide as pedicel. Postoculars hairs black above, very long
and whitish below. Ocellar tubercle with 1 pair of short oc and 2 minute posterior hairs;
1 pair of short vt close together. Antenna with scape and basal half of pedicel dark
brown, remainder yellow; arista dark brown; pedicel with a circle of black apical hairs;
postpedicel long conical, 2.8 times as long as wide, with black hairs except base bare;

232 D. YANG ET AL.

arista rather long, 2.6 times as long as postpedicel, short black pubescent. Proboscis
almost as long as heigth of head, dark brown with blackish hairs; clypeus grey
microtrichose; palpus yellow with 4 pale hairs.

Thorax black grey microtrichose. Pleura microtrichose but sternopleura with a
large polished patch in middle. Hairs on thorax pale, bristles black; hairs on mesono-
tum very short and sparse; humeri well developed with several very short pale hairs,
but without strong h; 1 npl (anterior npl absent), acr biseriate, dc as long as acr, biserial
anteriorly and uniserial in posterior half, 1 psa, 1 distinct presc in front of scutellum;
scutellum with 2 pairs of sc, basal pair as long as 2/5 of apical pair.

Legs mainly yellow, including all coxae; fore femur basally with blackish
antero-ventral spot; mid femur with a blackish antero-ventral stripe in basal half and
with a brown spot ventrally near apex; fore tibia brown especially anteriorly and ven-
trally, posteriorly dark yellow. Mid tibia anteriorly and ventrally dark brown; apex of
hind tibia dark brown; fore tarsomeres 1 and 5, mid and hind tarsomeres 3-5 dark
brown. Hairs and bristles on legs black except for white av bristles on fore femur. Fore
femur 1.8 times and mid femur 2.3 times as wide as hind femur; fore femur with a row
of long av and a row of pv bristles (4 additional hairs forming an second outer row in
the basal quarter); mid femur with two rows of short black spinules ventrally (longer
in basal quarter) and one row of about 10 black pv bristles, which are half as long as
femur is deep. Fore tibia distinctly thickened, with shorter dorsal bristles which are
about as long as tibia is thick, and very long ventral bristles on apical two thirds; ante-
riorly almost along entire length covered with dense, golden hairs; mid tibia with a row
of short weak black ventral hairs, and with a very long, pointed apical spur bearing an
apical hair (Fig. 2). Fore basitarsus with rows of long, black posterior and pv bristles.

Wing hyaline; veins dark brown, R445 and M distinctly convergent apically
(Fig. 1). Basal cells equally long. Anal vein distinct. Vein closing anal cell slightly S-
shaped. Squama dark yellow with pale hairs. Halter yellow.

Abdomen subshining black, thinly grey microtrichose; hypopygium black.
Hairs on abdomen short, pale. Male genitalia (Figs 3-5): Left epandrial lamella rather
wide basally with distinct lateral process bearing very long marginal bristles; right
epandrial lamella short with an apical row of long bristles, its apical projection (sur-
stylus) short and finger-like; left cercus somewhat acute apically; right cercus short,
obtuse apically.

Female. Body length 3.2-5.0 mm, wing length 4.4-4.5 mm. Similar to male, but
legs almost entirely yellow without brown areas, fore tibia weakly thickened, without
the very long black ventral hairs on tip, but being still longer haired than usual in
Platypalpus.

DISTRIBUTION: China (Guangdong).

REMARKS: The new species is very peculiar. It is easily separated from other
known species of the genus occurring in China by the fore tibia distinctly thickened
with conspicuous dorsal and very long ventral bristles, and the long setulose fore
basitarsus. P. convergens belongs to the P. pallidiventris-cursitans species-group of
Chväla (1989).

PLATYPALPUS FROM GUANGDONG 233

Platypalpus guangdongensis sp. n. Figs 6-9

MATERIAL: Holotype d, CHINA: Guangdong: Yingde, Shimentai National Forest Park,
28. III. 2003, leg. D. Yang (CAU). Paratype: 1 9, same data as holotype (CAU). The type
locality is a forest situated 15 km north of Yingde (24.09N/113.25E).

ETYMOLOGY: The specific name refers to the type locality.

DrAGNosis: Mainly yellow species with black head, scutellum and mediotergite
(postnotum); mesonotum subshiny; 2 pairs of vt; antenna yellow; postpedicel short
oval; mid femur without posteroventral bristles; mid tibia with a long apical spur ven-
trally (Fig. 6).

DESCRIPTION:

Male. Body length 2.5 mm, wing length 4.0 mm.

Head black, grey microtrichose. Eyes narrowly separated on face; face narrower
than frons. Hairs and bristles on head pale. Ocellar tubercle with 1 pair of oc and 4 very
short posterior hairs; 2 vt. Antenna yellow; scape without dorsal bristles; pedicel with
a circle of pale apical hairs; postpedicel short oval, 1.1 times as long as wide, with
some apical hairs; arista very long, 3.9 times as long as postpedicel, blackish, short
pubescent. Proboscis almost as long as heigth of head, brownish yellow with brown
hairs; palpus yellow with 3 long pale hairs.

Thorax yellow, thinly grey microtrichose; mesonotum subshining; scutellum
black, mediotergite (postnotum) blackish; polished spot on sternopleuron absent. Hairs
on thorax pale, bristles brownish yellow; 1 short hair-like h curved inwards, 2 npl
(anterior npl short and hair-like), acr 4-seriate, dc pale, as long as acr, 1 long psa, 1
short weak presc in front of scutellum, scutellum with 2 pairs of sc (basal pair very
short, one third as long as apical pair).

Legs yellow, but tarsomere 5 of all legs brown. Hairs on legs pale. Fore femur
1.2 times and mid femur 1.8 times as wide as hind femur; mid femur with two rows of
short black ventral spinules (av very short, pv slightly longer than av bristles, but ven-
tral bristles on basal 1/4 long and brownish yellow), without row of strong pv bristles;
mid tibia with a row of blackish ventral bristles and finger-like apical spur (Fig. 6).

Wing hyaline; veins brownish yellow, R,,; and M parallel apically. Basal cells
equal, crossveins contiguous. Squama yellow with pale hairs. Halter yellow.

Abdomen brownish yellow, pale grey microtrichose; sternites dark yellow,
hypopygium brownish. Hairs on abdomen pale. Male genitalia (Figs 7-9): Left epan-
drial lamella rather narrow in dorsal view, its apical projection (surstylus) large and
subtriangular with long apical bristles; right epandrial lamella slightly widening dor-
sally, its apical projection (surstylus) rather small truncate apically; left cercus rather
long, spine-like apically; right cercus short with acute apex.

Female. Body length 2.0 mm, wing length 2.8 mm. Similar to male, but
abdomen entirely dark yellow.

DISTRIBUTION: China (Guangdong).

REMARKS: The new species is very similar to Platypalpus guangxiensis Yang &
Yang from Guangxi, Sichuan and Hubei, but can be separated from the latter by the

234 D. YANG ET AL.

Fics 6-9

Platypalpus guangdongensis sp. n, male. 6, tip of mid tibia; 7, right epandrial lamella; 8, geni-
talia, dorsal view; 9, left epandrial lamella.

blackish mediotergite and short thick apical projection of the right epandrial lamella.
In P. guangxiensis, the mediotergite is yellow, and the apical projection of the right
epandrial lamella is long and spine-like (Yang & Yang, 1992). The unusual combina-
tion of characters does not allow to place this species in one of the species-groups
defined by Chväla (1989).

Platypalpus zhangae sp. n. Figs 10-13

MATERIAL: Holotype d, CHINA: Guangdong: Ruyuan, Nanling National Nature
Reserve, (23.20N/115.23E), Shumuyuan, 500 m, 26. II. 2003, leg. Lili Zhang (CAU). Paratype:
1 d. Guangdong, Ruyuan, forest near entrance of the Nanling National Nature Reserve,
26.111.2003, leg. P. Grootaert (sample 23008, RBINS). The type locality is a forest situated about
40 km north of Ruyuan (24.44N/113.18E).

PLATYPALPUS FROM GUANGDONG 235

EryMoLOGY: The species is named after the collector of the holotype.

DrAGNosis: Mainly black species with shining black mesonotum; 1 pair of vt;
postpedicel much elongated, 3.8 times longer than wide; arista rather short, at most as
long as postpedicel (Fig. 10); costa with a black stigma; mid femur without pv bristles;
mid tibia black without apical spur.

DESCRIPTION:

Male. Body length 3.0 mm, wing length 3.2 mm.

Head black, grey microtrichose. Eyes narrowly separated on face; frons wider
than face. Hairs on head pale, bristles brownish yellow. Ocellar tubercle with 1 pair of
oc and 4 posterior hairs slightly shorter than oc; 1 pair of short vt. Antenna (Fig. 10)
blackish with black scape; pedicel with a circle of black apical hairs; postpedicel long
conical, 3.8 times longer than wide, with blackish hairs except base bare; arista rather
short, as long as postpedicel, blackish, short pubescent. Proboscis almost as long as
heigth of head, dark brown with blackish hairs; palpus rather large, ovoid, yellow with
3 minute blackish hairs.

Thorax black, but humeri and prothorax paler (at least in alcohol preserved
specimens); mesonotum polished, sparsely haired but laterally with dense hairs;
pleuron subshiny, polished spot on sternopleuron not discernible. Hairs on thorax pale,
bristles brownish yellow; humerus with 6 hairs including 1 rather long hair-like h
curved inward; 1 npl (anterior npl absent), acr rather long, biseriate, diverging, dc
uniseriate and shorter than acr, 1 psa, 1 short presc in front of scutellum; scutellum very
short densely pubescent, with 2 pairs of sc, basal pair 2/3 as long as apical pair.

Legs dark brownish yellow; all coxae and trochanters yellow; mid tibia black,
hind tibia dark brown; tarsomere 5 dark brown. Hairs and bristles on legs pale. Fore
femur 1.2 times and mid femur 1.4 times as wide as hind femur; fore femur with a row
of av and a row of short pv bristles, which are as long as femur is wide; mid femur with
two rows of black ventral spinules (av short and pv rather long), but without row of
strong pv bristles; mid tibia with a row of short black ventral bristles, and without
apical spur.

Wing nearly hyaline; veins dark brown, C with a black stigma; R4,5 and M
parallel apically. Basal cells equally long. Vein closing anal cell recurrent (brown on
basal half, tip indistinct). Anal vein weakly indicated. Anal lobe of wing narrow.
Squama yellow with pale hairs. Halter pale yellow.

Abdomen blackish, grey microtrichose; hypopygium black. Hairs on abdomen
pale, dense, long at sides. Male genitalia (Figs 11-13): Left and right epandrial lamellae
rather broadly fused basally (best seen in dorsal view); left epandrial lamella rather
narrow in dorsal view, its apical projection (surstylus) rather large and not distinctly
separated from basal plate; right epandrial lamella rather wide in dorsal view, its apical
projection (surstylus) short and wide with a finger-like apico-lateral process; left and
right cerci basally with a medial process and apically finger-like.

Female. Unknown.

DISTRIBUTION: China (Guangdong).

REMARKS: The new species is somewhat similar to Platypalpus pallipilosus
Saigusa & Yang from Henan, but can be separated from the latter by the long post-

236 D. YANG ET AL.

Fics 10-13

Platypalpus zhangae sp. n, male. 10, antenna; 11, left epandrial lamella;12, genitalia, dorsal
view; 13, right epandrial lamella.

pedicel (3.8 times longer than wide) with the rather short arista (as long as the post-
pedicel). In P. pallipilosus, the postpedicel is shorter (2.1 times longer than wide) with
a longer arista (1.9 times longer than the postpedicel). Judging from the wing structure,
the shining mesonotum, the rather thin mid femur and the long antenna P zhangae
belongs to the P. longicornis species-group. However, instead of two pairs of vertical
bristles usually present in this species-group there is only one pair developed in the
new species.

PLATYPALPUS FROM GUANGDONG 2371

CONCLUSIONS

Guangdong belongs to the South China region of the Oriental realm with a sub-
tropical and tropical climate. Platypalpus is apparantly a more temperate genus which
is less well represented in hot, tropical parts of the world. Therefore it is not surprising
that up to now only 5 species of Platypalpus are known to occur in Guangdong, three
of which are described in this paper and two remain unnamed. Nevertheless, taking
into account the insufficient collecting effort, the rather early collecting time and the
difficulties to find suitable habitats, it is highly possible that more species will even-
tually be found in this province of China.

ACKNOWLEDGEMENTS

Our sincere thanks are due to Mrs Lili Zhang and Mrs Mengging Wang (China
Agricultural University, Beijing) for their help during the study. The research was
funded by the National Natural Science Foundation of China (No. 30070100_No.
30225009).

REFERENCES

BRUNETTI, E. 1920. Diptera Brachycera Vol. 1. The Fauna of British India, including Ceylon &
Burma. London, 401 pp.

CHVALA, M. 1975. The Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. I.
Fauna Entomologica Scandinavica 3: 1-336. Scandinavian Science Press, Kopenhagen.

CHvALA, M. 1989. Monograph of northern and central European species of Platypalpus
(Diptera, Hybotidae), with data on the occurrence in Czechoslovakia. Acta Universitatis
Carolinae - Biologica 32: 209-376.

CHVALA, M. & KOVALEV, V. G. 1989. Family Hybotidae (pp. 174-227). In: Soös, A. & Papp, L.
(eds). Catalogue of Palaearctic Diptera. Vol. 6. Akadémiai Kiadò, Budapest, 435 pp.

Frey, R. 1943. Übersicht der paläarktischen Arten der Gattung Platypalpus Macq. (=Coryneta
Meig.). (Dipt., Empididae). Notulae Entomologicae 23: 1-19.

GROOTAERT, P. 1984. Two new species of Platypalpus Macquart (Diptera, Empidoidea, Hybo-
tidae) from Papua New Guinea. Bulletin et Annales de la Société Royale d’Entomologie
de Belgique 120: 283-290.

GROOTAERT, P. & CHVALA, M. 1992. Monograph of the genus Platypalpus (Diptera: Empidoidea,
Hybotidae) of the Mediterranean region and the Canary Islands. Acta Universitatis
Carolinae - Biologica 36: 3-226.

SAIGUSA, T. & YANG, D. 2002. Empididae (Diptera) from Funiu Mountains, Henan, China (1).
Studia Dipterologica 9 (2): 519-543.

SMITH, K. G. V. 1965. Diptera from Nepal: Empididae. Bulletin of the British Museum (Natural
History), Entomology 17 (2): 61-112.

SMITH, K. G. V. 1975. Family Empididae (pp. 185-211). In: DELFINADO, M. D. & HARDY, D. E.
(eds). A Catalog of the Diptera of the Oriental Region. Vol. 2. The University Press of
Hawaii, Honolulu, 459 pp.

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

YANG, C. & YANG, D. 1992. Three new species of Empididae from Guangxi — Diptera:
Brachycera. Journal of the Guangxi Academy of Sciences 8 (1): 44-48.

YANG, D., AN, S. & Gao, C. 2002. New species of Empididae from Henan (Diptera) (pp. 30-38).
In: SHEN, X. & SHI, Z. (eds). The Fauna and Taxonomy of Insects in Henan. Vol. 5.
China Agricultural Scientech Press, Beijing, 453 pp.

238 D. YANG ET AL.

YANG, D. & MERZ, B. (2005). Review of the species of Platypalpus Macquart from Guangxi,
China (Diptera, Hybotidae, Tachydromiinae). Revue suisse de zoologie 112(4): 849-857.

YANG, D. & YANG, C. 1989. The dance flies of Xizang (II) (Diptera: Empididae). Acta Agri-
culturae Universitatis Pekinensis 15 (4): 415-424.

YANG, D. & YANG, C. 1990. Eleven new species of the subfamily Tachydrominae from Yunnan
(Diptera: Empididae). Zoological Research 11 (1): 63-72.

YANG, D. & YANG, C. 1992. Diptera: Empididae (pp. 1089-1097). In: CHEN, S. (ed.). Insects of
the Hengduan Mountains Region. Vol. 2. Science Press, Beijing, 1547 pp.

YANG, D. & YANG, C. 1997. Diptera: Empididae (pp. 1469-1476). In: YANG, X. (ed.). Insects of
the Three Gorge Reservoir Area of Yangtze River. Chongging Publishing House,
Chongqing, 1847 pp.

REVUE SUISSE DE ZOOLOGIE 113 (2): 239-246; juin 2006

New species and records of the genus Basanus Lacordaire
(Insecta: Coleoptera: Tenebrionidae)*

Wolfgang SCHAWALLER
Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany.
E-mail: schawaller.smns @naturkundemuseum-bw.de

New species and records of the genus Basanus Lacordaire (Insecta:
Coleoptera: Tenebrionidae). - Newly collected specimens of the genus
Basanus Lacordaire, 1859 from southeastern Asia are presented, including
new distributional data and new species: Basanus halmahericus sp. n.
(Moluccas: Halmahera), Basanus luzonicus sp. n. (Philippines: Luzon),
Basanus poringicus sp. n. (Borneo: Sabah). For Basanus philippinensis
Gebien, 1925 lectotype and paralectotypes are designated.

Keywords: Tenebrionidae - Basanus - new species - distribution - figures.

INTRODUCTION

The genus Basanus Lacordaire, 1859 was revised by Gebien (1925), a few
further species were described by Schawaller (1995) and by Masumoto & Merkl
(2003). The congeners are distributed in the Palaearctic Far East (including the Ussuri
Region, Korea, Japan and Taiwan), continental southeastern Asia, the Himalayas, the
Philippines, the Sunda Islands, Moluccas and New Guinea. In this contribution, accu-
mulated specimens from different collections are presented, including new taxa and
new distributional data. Two additional new species are available but are not described
because of the lack of males.

The generic and specific characters of Basanus have been discussed by
Schawaller (1995). Unfortunately, the shape of the aedeagus within this genus is rela-
tively simple and not distinctly modified between the species (compare Figs 1-5). The
generic limits of Basanus Lacordaire, 1859, Spiloscapha Bates, 1873 and Scaphidema
Redtenbacher, 1849 are still unclear. Doyen et al. (1989) listed some adult and larval
synapomorphic characters and included Basanus in the subtribe Scaphidemini within
the tribe Diaperini.

ACRONYMS OF DEPOSITORIES

CHBM Collection Prof. Dr Hans Bremer, Zoologische Staatssammlung, Miinchen
CRGT Collection Dr Roland Grimm, Tübingen

CSBC Collection Stanisiav Beévar, Ceské Budéjovice

HNHM Hungarian Natural History Museum, Budapest (Dr Otto Merkl)

MHNG Muséum d’histoire naturelle, Genève (Dr Giulio Cuccodoro)

NHMB Naturhistorisches Museum, Basel (Dr Michel Brancucci)

NHMB-F Naturhistorisches Museum, Basel, collection G. Frey (Dr Eva Sprecher)
SMNS Staatliches Museum für Naturkunde, Stuttgart (author)

SMTD Staatliches Museum für Tierkunde, Dresden (Olaf Jäger)

* Contributions to Tenebronidae, no. 55. - For no. 54 see Zootaxa, special issue Martens, 2006.
Manuscript accepted 17.11.2005

240 W. SCHAWALLER

THE SPECIES STUDIED

Basanus amamianus Chujo, 1966 Fig. 6

MATERIAL: Japan, Ryukyu Islands, Okinawa, Yona, Kunigami, 18.-20.II1.1991, leg.
T. Ueno, 1 ex. SMNS.

DISTRIBUTION: Japan, Ryukyu Islands (type locality).

Basanus erotyloides Lewis, 1891

MATERIAL: China, Shaanxi Prov., Taibai Shan above Houzhenzi, 1300-1700 m, 9.VI.-
3.VII.1998, leg. J. Martens & P. Jäger, 1 ex. SMNS. — Japan, Ehime Pref., Mt. Takanawasan,
7.V.1997, leg. H. Kan, 2 ex. SMNS. — Japan, Kyushu, Ohita Pref., Yoshibu, 19.VIII.1995, leg.
T. Ueno, 2 ex. SMNS. — Annam, Phuc-Son, holotype of syn. annamitus Gebien, 1925, SMTD.

DISTRIBUTION: Japan (type locality), China, Indochina.

Basanus fruhstorferi Gebien, 1940 Fig. 7

MATERIAL: S Celebes (Sulawesi), Lompa-Battau, 3000 ft., III.1896, leg. H. Fruhstorfer,
holotype NHMB-F (labelled as type), 1 paratype SMTD (labelled as cotype).

DISTRIBUTION: Sulawesi (type locality).

Basanus halmahericus sp. n. Figs 1, 8

HOLOTYPE (d): Moluccas (=Maluku), Halmahera, Sidangoli, Batu Putih, 100 m,
23.X1.1999, leg. A. Riedel, SMNS.

PARATYPES: Moluccas, Halmahera, Ibu, Desa Nanas, Gunung (= Mt.) Gamkonora, 100-
1000 m, 27.XI.1999, leg. A. Riedel, 2 ex. SMNS. — Moluccas, Halmahera, Buli, Maba, 20-
200 m, 6.-7.XI.1999, leg. A. Riedel, 1 ex. SMNS.

ETYMOLOGY: Named after the island Halmahera where the type series was col-
lected.

DESCRIPTION: Shape and colour pattern of pronotum and elytra see Fig. 8. Head,
pronotum and scutellum black with greenish metallic shine; elytron with identical
metallic shine and with a bigger round yellow spot before the middle and a smaller
yellow round spot before the tip; antenna completely black, legs light ferrugineous
with darker tibiae; ventral side black, ventrites red ferrugineous. Head with denser and
coarser punctation than on pronotum. Pronotum with distinct punctures, distance of the
punctures 4-8 times longer than the diameter; anterior margin completely bordered,
basal margin unbordered. Elytron with 8 rows of punctures, rows indistinct in the
anterior part, third row with about 25 punctures; intervals flat, with scattered punctures;
lateral margin to be seen nearly on its total length. Aedeagus see Fig. 1. Body length
7.2-8.0 mm.

DIAGNOsIs: To be recognized by the metallic dorsal side with small and round
elytral spots and by the shape of the aedeagus. A similar colour pattern and also a
metallic surface possess Basanus hellus Gebien, 1925 from the Philippines, but this
species is distinctly smaller (body length 4.2 mm) and the body is highly convex, so
that the lateral margin of the elytra can be seen in dorsal view only near the shoulders,
whereas in Basanus halmahericus sp. n. the lateral margin can be seen nearly on its
total length.

NEW SPECIES AND RECORDS OF BASANUS 241

IM

Fics 1-5
Aedeagus of Basanus species in dorsal view. 1: B. halmahericus sp. n., holotype, SMNS. 2:
B. javanus, aberant non-type, W Malaysia, CSBC. 3: B. luzonicus sp. n., holotype, SMNS. 4:
B. philippinensis, lectotype, NHMB-F. 5: B. poringicus sp. n., holotype, MHNG.

1 mm

Basanus javanus Chevrolat, 1878 Figs 2, 9

MATERIAL: S Sulawesi, Tanah Toraja, Pulu Pulu, 1700 m, 13.-16.VIII.1990, leg.
A. Riedel, 5 ex. SMNS. — Sulawesi, Kotamobagu, Modoinding, Gunung (= Mt.) Ambang, 1100-
1450 m, 6.XII.1999, leg. A. Riedel, 1 ex. SMNS. — SE Burma, 5.X1.1989, leg. S. Stahnke, 1 ex.
SMNS. — NE Laos, Hua Phan Prov., Ban Saluei, Phu Phan Mt, 1500-2000 m, 26.IV.-11.V.2001,
leg. J. Bezdek, 2 ex. SMNS. — W Malaysia, Island Tioman, Kampong Tekek, 9.III.1998, leg. L.
Dembicky & P. Pacholätko, 1 ex. NHMB. — W Malaysia, Perak, Maxwell Hill above Taiping,
900-1000 m, 12.-16.1.1995, leg. S. Beèvaf, 1 ex. CSBC. — N Thailand, W Chiang Mai, Doi
Suthep Pui NP, 30.V.1999, leg. R. Grimm, 1 ex. CRGT.

REMARKS: The single male from W Malaysia/Maxwell Hill possesses a some-
what different colour pattern with the anterior elytral spots round (Fig. 9) and not band-
ed (Schawaller 1995: fig. 5) as usually present in this species. The other diagnostic
characters (thick antennae, ventral side black, shape of the aedeagus: Fig. 2) coincide.

DISTRIBUTION: Java (type locality), Borneo, Sulawesi (new record), Myanmar,
Laos, Vietnam, W Malaysia (new record), Thailand (new record).

Basanus longior Gebien, 1925 Fig. 10

MATERIAL: Sumatra, 1 ex. NHMB-F. — W Sumatra, Payakumbuh, Harau Valley, 9.-
29.X.1991, leg. A. Riedel, 1 ex. SMNS. - Borneo, Sabah, Mt. Kinabalu NP, Poring Hot Springs,
520 m, 13.V.1987, leg. A. Smetana, 1 ex. SMNS (duplicate from MHNG).

242 W. SCHAWALLER

REMARKS: The pronotum varies from completely black up to black with lighter
margins.
DISTRIBUTION: Sumatra (type locality), Borneo (new record).

Basanus luzonicus sp. n. Figs 3, 11

HOLOTYPE (d ): Philippines, N Luzon, Mountain Prov., Bontoc Region, NW Barlig, 2000
m, 9.IV.2000, leg. L. Dembicky, SMNS.

PARATYPES: Philippines, Luzon, Mt. Polis, leg. Boettcher, 12 ex. SMTD, 2 ex. SMNS,
2 ex. NHMB-F.

ETYMoLoGY: Named after the Philippine Island Luzon, where the type series

was collected.

DESCRIPTION: Shape and colour pattern of pronotum and elytra see Fig. 11.
Head, pronotum and scutellum dark ferrugineous without metallic shine, lateral margin
of pronotum somewhat lighter; elytron also dark ferrugineous and with a bigger round
yellow spot before the middle and a smaller yellow round spot before the tip; antenna
completely dark ferrugienous with the tip of the last antennomere somewhat lighter,
legs ferrugineous; ventral side including ventrites dark ferrugineous. Head with denser
and coarser punctation than on pronotum. Pronotum with distinct punctures, distance
of the punctures 4-8 times longer than the diameter; anterior margin completely
bordered, basal margin unbordered. Elytron with 8 rows of punctures, rows indistinct
in the anterior part, third row with about 30 punctures; intervals flat, with scattered
punctures; lateral margin to be seen in dorsal view nearly on its total length. Aedeagus
see Fig. 3. Body length 5.2-6.2 mm.

DIAGNOSIS: To be recognized by the smaller body size, by the elytral colour pat-
tern, by the indistinct elytral rows and by the shape of the aedeagus. Basanus luzoni-
cus sp. n. is quite similar to Basanus philippinensis Gebien, 1925 also from the
Philippines, but in this species the elytral colour pattern is different (anterior spot trans-
verse, band-like and with sutural interruption, Fig. 12), the punctural rows on the elytra
are dense and distinct and the aedeagus is slightly different (compare Figs 3-4).

REMARKS: This species-group includes also 2 additional new species from
Borneo and Sumatra, which are listed below, but which are not described because of
the lack of males. Both have a similar dorsal colour pattern (Figs 16-17) as in Basanus
luzonicus sp. n. (Fig. 11), but a different dorsal punctation and a different structure of
the elytral intervals.

Basanus philippinensis Gebien, 1925 Figs 4, 12

MATERIAL: Philippines, leg. Semper, 1 4 cotype NHMB-E designated herewith as lec-
totype. — Philippines, Luzon, Malinao, Tayabas, leg. Baker, 1 9 cotype NHMB-F, 1 cotype
SMTD, both designated herewith as paralectotypes. — Philippines, Luzon, Mt. Makiling, leg.
Baker, 1 cotype SMTD, designated herewith as paralectotype.

REMARKS: The lectotype and paralectotypes are designated in order to preserve
stability of the nomenclature in this group, according to the Article 74.7.3 of the
International Code of Zoological Nomenclature (1999).

DISTRIBUTION: Philippines (type localities).

NEW SPECIES AND RECORDS OF BASANUS 243

Fics 6-10

Dorsal view of Basanus species. 6: B. amamianus, non-type, Okinawa, SMNS. 7: B. fruhstor-
feri, paratype, SMTD. 8: B. halmahericus sp. n., holotype, SMNS. 9: B. javanus, aberrant non-
type, W Malaysia, CSBC. 10: B. longior, non-type, Sumatra, SMNS.

244 W. SCHAWALLER

Basanus poringicus sp. n. Figs 5, 13

HOLOTYPE (d): Borneo, Sabah, Mt. Kinabalu NP, Poring Hot Springs, 500 m, 8.V.1987,
leg. D. Burckhardt & I. Lòbl, MHNG.
PARATYPE: Same data as holotype, 1 ex. SMNS.

ETYMOLOGY: Named after the type locality Poring.

DESCRIPTION: Shape and colour pattern of pronotum and elytra see Fig. 13.
Head red ferrugineous, pronotum red ferrugineous with 2 darker longitudinal spots,
scutellum red ferrugineous; elytron darker ferrugineous and with a large, yellow spot
in the anterior part; antenna dark ferrugineous with the 3 basal segments somewhat
lighter and coloured like the head, legs completely ferrugineous; ventral side including
ventrites red ferrugineous. Head with denser and coarser punctation than on pronotum.
Pronotum with distinct punctures, distance of the punctures 2-5 times longer than the
diameter, additionally with scattered and distinctly bigger punctures; anterior margin
completely bordered, basal margin unbordered. Elytron with 8 rows of punctures, third
row with about 50 punctures; intervals convex, with a few scattered punctures; lateral
margin to be seen nearly on its total length. Aedeagus see Fig. 5. Body length
5.8-6.0 mm.

DIAGNOSIS: Basanus poringicus sp. n. can be recognized by the dorsal colour
pattern (Fig. 13), by convex elytral intervals and by the shape of the aedeagus. The
colour pattern combined with convex elytral intervals are quite unusual within the
genus and do not occur in any other congeners.

Basanus soppongensis Schawaller, 1995
MATERIAL: Vietnam, Nam Cat NP, 4.-11.V.1994, leg. J. Zacharda, 1 ex. CHBM.
DISTRIBUTION: Thailand (type locality), Vietnam (new record).

Basanus sulawesicus (Schawaller, 1997) Fig. 14

MATERIAL: C Sulawesi, Palu, Palolo, Lindu NP, 25.-27.VIII.1990, leg. A. Riedel, 4 holo-
type SMNS.

REMARKS: Transferred from the genus Spiloscapha Bates, 1873 to the genus
Basanus Lacordaire, 1859 by Schawaller (2004).

DISTRIBUTION: Sulawesi (type locality).

Basanus sumatranus Gebien, 1925 Figs
MATERIAL: Sumatra, Solok, © holotype NHMB-F (labelled as type)
REMARKS: Gebien (1925) named this species for the first time in the key on page
145 sumatrensis and later in the description on page 147 and in the legend of the plate

sumatranus, the type specimen is also labelled as sumatranus. Thus, sumatrensis in the
key is obviously a printing error and I fix here the name sumatranus for this species.

DISTRIBUTION: Sumatra (type locality).

Basanus sp. A © Fig. 16

MATERIAL: Borneo, Sabah, Mt. Kinabalu NP, Headquarters, 1560 m, 3.-13.VIIL.1988,
leg. A. Smetana, 1 9 MHNG.

NEW SPECIES AND RECORDS OF BASANUS 245

Fics 11-17

Dorsal view of Basanus species. 11: B. luzonicus sp. n., holotype, SMNS. 12: B. philippinensis,
lectotype, NHMB-F. 13: B. poringicus sp. n., holotype, MHNG. 14: B. sulawesicus, holotype,
SMNS. 15: B. sumatranus, holotype, NHMB-F. 16: Basanus sp. A, 2, Borneo, MHNG. 17:
Basanus sp. B, 9, Sumatra, SMNS.

246 W. SCHAWALLER

REMARKS: To be recognized by the small body size (5.0 mm), by the dorsal
colour pattern (Fig. 16), by the ferrugineous ventral side, and by a fine but dense punc-
tation on the elytra without distinct punctural rows and with absolutely flat intervals.
The specimen presents a new species, but I avoid to describe it because of the lack of
males.

Basanus sp. B 9 Fig. 17

MATERIAL: Sumatra, Bengkulu, 20 km S Muko Muko, 20 m, 16.VII.1981, leg. D. Erber,
1 9 SMNS.

REMARKS: To be recognized by the small body size (5.5 mm), by the dorsal
colour pattern (Fig. 17), by the ferrugineous ventral side, and by distinct punctural
rows on the elytra with slightly convex intervals. The specimen presents a further new
species, but I also avoid to describe it because of the lack of males.

ACKNOWLEDGEMENTS

Thanks are due to all colleagues and friends loaning material from their
collections, they are compiled in the list of the depositories. The hospitality of Dr Otté
Merkl (Budapest) and Dr Eva Sprecher (Basel) during my visits is greatly appreciated.
Johannes Reibnitz (Stuttgart) kindly prepared again the photographs and arranged
them on plates.

REFERENCES

DOYEN, J. T., MATTHEWS, E. G. & LAWRENCE, J. F. 1989. Classification and annotated checklist

of the Australian genera of Tenebrionidae (Coleoptera). Invertebrate Taxonomy 3:
229-260.

GEBIEN, H. 1925. Die Tenebrioniden (Coleoptera) des Indo-Malayischen Gebietes, unter
Beriicksichtigung der benachbarten Faunen, IV. Die Gattungen Phloeopsidius, Dysantes,
Basanus, und Diaperis. Philippine Journal of Science 27: 131-157.

ICZN, 1999. International Code of Zoological Nomenclature, 4th Edition. xxix + 306 pp.
International Trust for Zoological Nomenclature, c/o The Natural History Museum,
London.

MASUMOTO, K. & MERKL, O. 2003. A new Spiloscapha and a new Basanus from Taiwan
(Coleoptera. Tenebrionidae. Scaphidemini). Entomological Review of Japan 58:
165-171.

SCHAWALLER, W. 1995. The genus Basanus Lacordaire (Coleoptera. Tenebrionidae) in conti-
nental south-eastern Asia. Acta Zoologica Academiae Scientiarum Hungaricae 41:
327-334.

SCHAWALLER, W. 2004. New species and records of the genus Spiloscapha Bates (Coleoptera.
Tenebrionidae) from the Oriental and Papuan Regions. Stuttgarter Beiträge zur
Naturkunde (A) 661: 1-10.

REVUE SUISSE DE ZOOLOGIE 113 (2): 247-261; juin 2006

Further considerations regarding the status of
Grosphus madagascariensis (Gervais) and Grosphus hirtus
Kraepelin, and description of a new species (Scorpiones, Buthidae)

Wilson R. LOURENCO! & Steven M. GOODMAN?

1 Département de Systématique et Evolution, USM 0602, Section Arthropodes
(Arachnologie), Muséum national d’Histoire naturelle, CP 053, 61 rue Buffon,
75005 Paris, France. E-mail: arachne@mnhn.fr

2 Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago,
Illinois 60605, USA, and WWE, BP 738, Antananarivo (101), Madagascar.

E-mail: sgoodman @fieldmuseum.org

Further considerations regarding the status of Grosphus madagas-
cariensis (Gervais) and Grosphus hirtus Kraepelin, and description of a
new species (Scorpiones, Buthidae). - New considerations regarding the
species Grosphus madagascariensis (Gervais, 1843), type species of the
genus Grosphus Simon, 1880, and Grosphus hirtus Kraepelin, 1900 are pro-
posed, and both species are redescribed. One new species, Grosphus
goudoti sp. n., is described from the Province d’Antsiranana, Forét de
Bobankora, E of Daraina on the northern range of Madagascar. With this
description the total number of known species in this endemic genus is now
15. Some details are presented on the ecological settings of the sites where
the described specimens were collected. A revised key to the species of
Grosphus is given.

Keywords: Scorpiones - Buthidae - Grosphus - new species - taxonomy -
Madagascar.

INTRODUCTION

As already discussed in recent papers (Lourenço, 2003; Lourengo et al., 2004),
the first Grosphus species to be described was Scorpio (Androctonus) madagas-
cariensis Gervais, 1843 = G. madagascariensis (Gervais, 1843). Kraepelin (1900)
contributed to the study of this genus and described several new species, including G.
hirtus, a species morphologically similar to G. madagascariensis. In his monograph on
the scorpions of Madagascar, Fage (1929) characterized both species. After Fage’s
(1929) monograph several new species of Grosphus were described (e. g. Lourenço,
2003; Lourengo et al., 2004), and precise diagnoses have been proposed for G. mada-
gascariensis and G. hirtus (see Lourenço, 1996).

The taxonomy of Grosphus is based mainly on two characters: the pattern of
coloration and the morphology of the basal middle lamellae of the female pectines.

Manuscript accepted 03.05.2005

248 W. R. LOURENÇO & S. M. GOODMAN

This last character has been considered by scorpion taxonomists to be of species
specific significance and possess little intraspecific variation. However, more detailed
investigations have showed that in some cases closely related species have a similar
basal middle lamellae morphology (Lourengo, 2003; Lourengo & Goodman, 2003;
Lourengo et al., 2004). On the basis of this character some populations which have
been attributed to widely distributed species, such as G. madagascariensis, G. lim-
batus, and G. bistriatus, remained undescribed until recently (Lourengo, 2003;
Lourenco et al., 2004). Neverless, problems of faulty species identification remains
possible in morphologically similar taxa, particularly those named in the early stages
of the taxonomy of this genus. This is certainly the case for G. madagascariensis and
G. hirtus.

Grosphus madagascariensis was described on the basis of one single specimen
from Madagascar, without any indication of its precise collection locality. The type
specimen, deposited in the Museum in Paris, has an associated original label with the
distinct handwriting of Gervais reading “S. goudotii”. A subsequent label written by
Kraepelin is more precise and reads S. goudotii (Androctonus), Madagascar, Goudot.
The name goudotii was not used in the original publication (Gervais, 1843) and the
original label only seems to indicate an intention of Gervais to dedicate this species to
the collector Jules Goudot. In Museum collections registration the following remark
was written by Vachon: Androctonus goudotti Gervais, type = Grosphus madagas-
cariensis (Gervais) 1844. The type specimen, presumably a male, is so poorly pre-
served that only metasomal characters can be observed. The illustrations given by
Gervais (1843, 1844) are, however, clear enough to confirm the identity of this
specimen as the type of G. madagascariensis. G. hirtus was described from a precise
locality, Makaraingo in the central region of Madagascar, however, the type specimen
is a juvenile female.

It is known that Jules Goudot travelled in the regions between the east coast, the
region of Tamatave and Sainte-Marie Island. At the time of his journeys, other parts of
Madagascar were inaccessible (forbitten) zones to foreign people (see Dorr, 1997 for
details of Goudot’s journeys). The region of Tamatave (Toamasina) was, and still is,
one of the major harbours in Madagascar, a natural point of entry to the island. The
Province of Toamasina was visited by Jules Goudot on several occasions while he
stayed on Madagascar, so the specimen on which the description of G. madagas-
cariensis was based was most probably collected somewhere in the eastern coastal
region of the island.

Although G. madagascariensis and G. hirtus have overlapping areas of distri-
bution, they inhabit different habitats. The first species is common in humid forests,
whereas the second lives in more dry deciduous forests (Fage, 1929). Herein we
redescribe both G. madagascariensis and G. hirtus based newly collected material
from their typical habitats. One new species, closely associated to the G. madagas-
cariensis/G. hirtus group, is described from a distinct forest habitat in northeastern
Madagascar, the Forêt de Bobankora, E of Daraina in the Province d’ Antsiranana.

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 249

TAXONOMIC TREATMENT
BUTHIDAE C.L. Koch, 1837

Grosphus Simon, 1880
Grosphus madagascariensis (Gervais, 1843) Figs 1-4, 9-12

Scorpio (Androctonus) madagascariensis Gervais, 1843: 129.
Scorpio (Androctonus) madagascariensis Gervais, 1844: 213.
Grosphus madagascariensis Simon, 1880: 377.

Grosphus madagascariensis Kraepelin, 1899: 33.

Grosphus madagascariensis Kraepelin, 1900: 12.

Grosphus madagascariensis Fage, 1929: 642.

Grosphus madagascariensis Lourenço, 1996: 9.

MATERIAL: Madagascar, Province de Toamasina, Forét de Plateau de Makira, Forét de
Vohitaly, site F, 5 km SE village Anjiahely, 15°26’ 58” S, 49°32’06” E, 540-680 m, 28/XII/2002,
pitfall vial marked ‘pitfall Vohitaly’ (V. Andrianjakarivelo), 1 male, 1 female (MHNG).

REDESCRIPTION: (based on male and female, measurements in Table I)
Scorpion of medium size, with a total length of 45-60 mm. General coloration

reddish brown to dark brown. Carapace and tergites brownish, in females darker than
in males; eyes surrounded by black pigment. Metasoma: all segments reddish brown,

TABLE I. Comparative morphometric values (in mm) of the male and female of Grosphus mada-
gascariensis and G. hirtus examined and of the male holotype of G. goudoti sp. n. and G. simoni.

G. madagascariensis —G. hirtus G. goudoti Sp.n. G. simoni

M F M F M M
Total length 45.4 52.5 38.9 48.8 59.8 54.2
Carapace:
- length ID 6.2 48 6.1 HD. 6.4
- anterior width 4.0 44 34 43 3.3 4.8
- posterior width 6.6 7.6 5.8 6.9 U) 6.9
Metasomal segment I:
- length 33 4.1 2.8 3:3 SQ 4.5
- width 31 37 3.1 39) 4.1 3.4
Metasomal segment V:
- length 6.5 6.8 Sw Oxo 8.4 all
- width 3.1 32 27, a 3.4 32
- depth 3.0 3.3 2.8 359 3.4 353,
Vesicle:
- width 2] 3.0 2 4100035 959 2.8
- depth 2.6 2.9 ZINIO] 33 279
Pedipalp:
- femur length 5.4 5.6 4.2 DI eo 6.2
- femur width 17 1.9 1.4 1.8 2.2 1.9
- patella length 6.2 6.8 49 6.1 8.4 To!
- patella width 25 2.6 Dal ZT, 2.9 fl
- chela length 10.3 10.8 8.3 OF 14.3 11.9
- chela width 2.8 255 25 25 3.9 351
- chela depth 235 24 25 2.4 35) 2.8

Movable finger:
- length SS) 6.6 46 6.0 8.2 6.8

250 W. R. LOURENCO & S. M. GOODMAN

Fics 1-4

Grosphus madagascariensis. 1-2. Male from Toamasina, dorsal and ventral aspects. 3-4. Female,
idem.

with segments IV and V slightly darker; carinae blackish brown. Telson reddish brown;
aculeus with reddish base and reddish brown tip. Venter: coxapophysis, sternum,
genital operculum and pectines yellowish; sternites greenish yellow. Chelicerae

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 251

yellowish, with dark variegated pigmentation; fingers yellowish, with the teeth reddish.
Pedipalps reddish, with vestigial variegated spots; legs yellowish in males and reddish
in females, with diffused fuscous spots.

Morphology. Carapace moderately granular, more intensely so in females;
anterior margin almost straight, with a very discrete median concavity. All carinae
moderate to weak; furrows moderately to weakly developed. Median ocular tubercle
anterior to centre of carapace; median eyes separated by a little more than one ocular
diameter. Three pairs of lateral eyes. Sternum between sub-triangular and sub-penta-
gonal in shape. Mesosoma: tergites with minute moderately intense granulation.
Median carina moderately to weakly developed on all tergites. Tergite VII penta-
carinate. Venter: genital operculum consisting of two semi-oval plates. Pectines:
pectinal teeth count 20-20 in male and 16-15 in female; basal middle lamella not
dilated in males, strongly dilated in females. Sternites smooth, with weakly elongated
stigmata. Metasoma: all segments longer than wide, with carinae strongly marked;
segments I to II with 10 carinae, crenulate; segment IV with 8 carinae, crenulate.
Segment V with 5 carinae. Dorsal carinae on segments II to IV with at least one
posterior spinoid granule strongly developed. Intercarinal spaces weakly granular in
males, more intensely granular in females. Telson with granules on latero-ventral and
ventral surfaces; its dorsal surface smooth; aculeus weakly curved and shorter than the
vesicle; subaculear tooth weak to vestigial. Cheliceral dentition characteristic of the
family Buthidae (Vachon, 1963); two distinct small basal teeth present on the movable
finger; ventral aspect of both fingers and of manus densely set with long setae.
Pedipalps: femur pentacarinate; patella with dorsointernal and ventrointernal carinae
and with several spinoid granules on the internal face; chela smooth, with vestigial
carinae. Fixed and movable fingers with 11/13 oblique rows of granules.
Trichobothriotaxy; orthobothriotaxy A-a (Vachon, 1974, 1975). Legs: tarsus with
numerous short thin setae ventrally. Tibial spurs present on legs II and IV; pedal spurs
present on legs I to IV; all spurs strong.

EcoLoGY: The specimens of Grosphus madagascariensis examined where col-
lected in a distinct humid lowland forest of eastern Madagascar. The portion of the hu-
mid forest is close to the Masoala Peninsula, which receives on average almost 6 m of
rainfall per year (Kremen, 2003).

In the Makira Forest, the site where the material was collected, the following
species of scorpions were collected by V. Andrianjakarivelo with pit-fall devices dur-
ing an inventory of small mammals of this region: Grosphus madagascariensis,
Grosphus simoni Lourengo et al., 2004 and Tityobuthus baroni (Pocock, 1890).

Grosphus hirtus Kraepelin, 1900 Figs 5-8, 13-15

Grosphus hirtus Kraepelin, 1900: 15.
Grosphus hirtus Fage, 1929: 645.
Grosphus hirtus Lourengo, 1996: 11.

MATERIAL: The adult specimens were collected in: Province of Mahajanga,
Ankarafantsika Reserve, Forest station of Ampijoroa, Jardin Botanique A (pitfall traps), 16°18’S,
46°48’E, 24-27/11/2001 (G. Garcia Herrero), 1 male, 2 females, 36 juveniles. Juveniles were ob-
tained in the laboratory during the study of the life cycle of the species (all in MHNG).

252 W. R. LOURENÇO & S. M. GOODMAN

FIGS 5-8
Grosphus hirtus. 5-6. Male from Ankarafantsika, dorsal and ventral aspects. 7-8. Female, idem.

REDESCRIPTION: (based on male and females, measurements in Table I)

Scorpion of medium size with a total length of 40-50 mm. General coloration
yellowish to reddish yellow, with variegated brownish spots over body and ap-
pendages. Carapace yellowish, with spots more marked on anterior and lateral edges;

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 253

Fics 9-15

9-12. Grosphus madagascariensis. 9. Metasomal segments III-V and telson of male holotype,
lateral aspect. 10. Metasomal segments IV-V and telson of male from Toamasina, lateral aspect.
11-12. Palpal patella and femur of male from Toamasina, dorsal aspect. 13-15. Grosphus hirtus,
male from Ankarafantsika. 13. Metasomal segments III-V and telson, lateral aspect. 14-15.
Palpal patella and femur, dorsal aspect.

254 W. R. LOURENÇO & S. M. GOODMAN

eyes surrounded by black pigment. Mesosoma: yellowish, with confluent dark zones
on the posterior edge of tergites. Metasoma: segments I to III yellowish; IV reddish
yellow; V reddish; all segments with variegated dark pigmentation; dorsal aspect of
segments I to IV each with a triangular spot. Telson reddish, without spots on carinae;
aculeus with reddish base and dark reddish tip. Venter: coxapophysis, sternum, genital
operculum and pectines yellowish; sternites dark yellow, with V to VII intensely
spotted. Chelicerae yellowish, with dark variegated pigmentation throughout; fingers
dark brown, with reddish teeth. Pedipalps: reddish yellow with variegated brownish
spots. Legs yellowish, with diffused brownish spots.

Morphology. Carapace weakly granular in males, moderately granular in
females; anterior margin almost straight, with a weak median concavity. All carinae
and furrows weakly developed. Median ocular tubercle anterior to centre of carapace;
median eyes separated by one ocular diameter. Three pairs of lateral eyes. Sternum
sub-triangular in shape. Mesosoma: tergites with thin but intense granulation. Median
carina moderately developed on all tergites. Tergite VII pentacarinate. Venter: genital
operculum consisting of two subtriangular plates. Pectines: pectinal tooth count mostly
19-19 in males and 15-15 in females; basal middle lamellae not dilated in males,
strongly dilated in females. Sternites smooth, with weakly elongated stigmata; VII with
two vestigial carinae. Metasoma: segment I wider than long; segments I to III with 10
carinae, crenulate. Segment IV with 8 carinae, crenulate. Segment V with 5 carinae,
crenulate. Dorsal carinae on segments II to IV each with one strong posterior spinoid
granule. Intercarinal spaces strongly granular. Telson with a strong granulation on
latero-ventral and ventral surfaces; its dorsal surface smooth; aculeus weakly curved
and shorter than the vesicle; subaculear tooth vestigial. Cheliceral dentition charac-
teristic of the family Buthidae (Vachon, 1963); two distinct basal teeth present on the
movable finger; ventral aspect of both fingers and of manus densely set with long setae.
Pedipalps: femur pentacarinate; patella with a dorsointernal carina and with several
spinoid granules on the internal aspect; chela smooth, without carinae. Fixed and
movable fingers with 11/12 oblique rows of granules. Trichobothriotaxy; orthobothrio-
taxy A-a (Vachon, 1974, 1975). Legs: tarsus with numerous short thin setae ventrally.
Tibial spurs present on legs III and IV; pedal spurs present on legs I to IV; all spurs
moderately to strongly developed.

EcoLoGY: The most common native forest formation of the Ankarafantsika
region is a dry deciduous forest that occurs on a sandy substrate (Ramangason, 1988).
The principal geological formation in the vicinity of the Ampijoroa Forest Station,
which includes the Jardin Botanique A, is a plateau that rises to an elevation of 310-
340 m above sea level, and this is most likely the zone where the Grosphus hirtus
specimens treated in this paper were collected. Several different plants occurring in this
habitat have varied adaptations to resist desiccation during the pronounced dry season,
and different vertebrate species undergo hibernation or torpor to survive this period.

Annual precipitation in the Ankarafantsika area ranges from 1000-1500 mm,
most of which falls between the months of November to April (Nicoll & Langrand,
1989). The month of January experiences the heaviest rainfall, with slightly less than
50% of the annual total. A very pronounced dry and cool period occurs between the
months of May and September. During this latter period little to no rain falls and the

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 255

forest experiences a distinct drought. Monthly mean temperatures range from 17° to
35° C, and the annual average temperature is 26°C.

The scorpion fauna of the Forest Station of Ampijoroa, part of the Parc National
d’ Ankarafantsika, is composed of Grosphus hirtus, G. ankarafantsika Lourenço, 2003,
G. garciai Lourenço, 2001, Tityobuthus dastychi Lourengo, 1997, and Opisthacanthus
madagascariensis Kraepelin, 1894.

Grosphus goudoti sp. n. Figs 16-24

MATERIAL: Madagascar, Province d’Antsiranana, Forêt de Bobankora, versant ouest, site
N° 2, 11 km E of Daraina (13°13.414’S, 49°45.586’E), 350-550 m (M. Raheriarisena & H. A.
Rakotondravony), X/2002-III/2003, male holotype, deposited in the Muséum d’histoire na-
turelle, Genève.

ETYMOLOGY: This patronym is in honor of Jules Goudot, collector of the first
Grosphus species in Madagascar.

DIAGNOSIS: Scorpions of medium size, with a total length of 60 mm. General
coloration reddish brown to dark brown. Certain morphological characters indicate that
G. goudoti sp. n. is close to the G. madagascariensis/G. hirtus group, and in particular
to G. simoni Lourengo, Goodman & Ramilijaona, 2004 but it can be readily distin-
guished from the last species by the following characters: (i) a much darker coloration
of carapace and tergites, with the presence of one inverted black triangle in the anteri-
or zone of the carapace, stretching from the lateral eyes to the median eyes; (ii) the
teeth of pectines longer than in G. simoni; (iii) dorsal carinae of metasomal segments
II to IV without any posterior spinoid granules.

DESCRIPTION: (based on male holotype, measurements in Table I)

Coloration. Basically reddish brown to dark brown. Prosoma: carapace reddish
brown with one inverted black triangle in the anterior zone of the carapace, stretching
from the lateral eyes to the median eyes; eyes surrounded by black pigment.
Mesosoma: reddish brown, with dark strips on posterior margins of tergites.
Metasoma: segments I to III reddish brown; IV-V dark brown; all segments with some
vestigial dark pigmentation on carinae. Telson reddish brown, with dark zones on
granulations; aculeus with reddish brown base and dark brown tip. Venter: coxapo-
physis, sternum, genital operculum and pectines yellowish; sternites yellow with
greenish zones; VII dark brown. Chelicerae reddish yellow, with dark variegated
pigmentation on the entire surface; fingers reddish brown; teeth reddish. Pedipalps:
reddish to reddish brown; chela fingers dark brown. Legs yellowish, with very diffused
variegated spots.

Morphology. Carapace covered with a thin but intense granulation; anterior
margin almost straight, with a very weak median concavity. All carinae weak; furrows
moderately developed. Median ocular tubercle anterior to centre of carapace; median
eyes separated by one ocular diameter. Three pairs of lateral eyes. Sternum sub-trian-
gular in shape. Mesosoma: tergites with thin but intense granulation. Median carina
moderately developed on all tergites. Tergite VII pentacarinate. Venter: genital oper-
culum consisting of two semi-oval plates. Pectines: pectinal tooth count 19-19, basal
middle lamellae of each pecten not dilated in males. Sternites smooth, with moderate-

256 W. R. LOURENÇO & S. M. GOODMAN

Fics 16-17
Grosphus goudoti sp. n. Male holotype, dorsal and ventral aspects.

ly elongated spiracles; VII with four weak carinae. Metasoma: segments I and II with
10 carinae, moderately crenulate; segments III and IV with 8 carinae, weakly crenu-
late. Segment V with 5 carinae. Dorsal carinae on segments II to IV without any pos-
terior spinoid granules. Intercarinal spaces moderately to weakly granular. Telson mod-
erately granular over latero-ventral and ventral surfaces; its dorsal surface smooth;
aculeus moderately curved and shorter than the vesicle; subaculear tooth vestigial.
Cheliceral dentition characteristic of the family Buthidae (Vachon, 1963); two distinct
basal teeth present on the movable finger; ventral aspect of both fingers and of manus
densely set with long setae. Pedipalps: femur pentacarinate; patella with dorsointernal
and ventralinternal carinae and with some strong spinoid granules on the internal
aspect; chela smooth, without carinae. Fixed and movable fingers with 11/12 oblique
rows of granules. Trichobothriotaxy; orthobothriotaxy A-a (Vachon, 1974, 1975).
Legs: tarsus with numerous short thin setae ventrally. Tibial spurs present on legs III
and IV; pedal spurs present on legs I to IV; all spurs strong.
Female unknown.

EcoLocy: The site where the holotype of Grosphus goudouti sp. n. was
collected is in a forested area forming an exceptional zone of ecotonal transition in the
central portion of northern Madagascar. This is known as the Daraina Forest, named
after the largest local village. This area is biotically very complex and the remaining
forests are greatly reduced in size and fragmented. In a distance of a few kilometres
there are transitions between dry deciduous forest formations prevailing in the west
and humid forests prevailing in the east. Elevation, aspect, soil type, and distance to the
sea are important parameters associated with these transitions. The site where the holo-
type of G. goudoti sp. n. was collected is composed of dry deciduous lowland forest

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS SY)

Fics 18-24

Grosphus goudoti sp. n. Male holotype. 18. Chelicera, dorsal aspect. 19. Metasomal segments
III-V and telson, lateral aspect. 20-24. Trichobothrial pattern of pedipalp. 20-21. Chela, dorso-
external and ventral aspects. 22-23. Patella, dorsal and external aspects. 24. Femur, dorsal aspect.

258 W. R. LOURENCO & S. M. GOODMAN

Fic. 25

Map of Madagascar, with indication of the sites were Grosphus madagascariensis and G. hirtus
have been collected, and the locality of the new Grosphus species. G. madagascariensis (black
star), G. hirtus (black circle), and G. goudoti sp. n. (black circle with white star).

that is relatively intact, with little leaf litter, relatively open understory, which rests on
reddish lateritic soils.

Other species of scorpions known to occur in the forests of the Daraina area are
Heteroscorpion magnus Lourenco & Goodman, 2002, Tityobuthus darainensis
Lourenço & Goodman, 2002, and Grosphus darainensis Lourengo, Goodman &
Ramilijaona, 2004 (Lourenco and Goodman, 2002; Lourenco et al., 2004).

KEY TO THE SPECIES OF THE GENUS GROSPHUS

1 Pectines with a maximum of 21 teeth... .. .22¢ au see 2 ee 2
= Pectines; with more than, 22:teeth vu: COM ae ee eee 7
2 Coloration yellowish to reddish brown; variegated brownish pigmen-

tation present. 32.3 eagle TRO SA so eee 3

- Coloration from yellowish to reddish brown or dark brown; variegated
brownish pigmentation absent (ite ee eee "i

Do

un

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 259

Coloration reddish brown; adult body length 30 mm ............ G. garciai
Coloration yellowish; adult body length 40-50 mm .............. G. hirtus
Metasomal segments II-IV with one or more spinoid posterior granules..... I
Metasomal segments I-IV without any spinoid granule . .... G. goudoti sp. n.
Metasomal segments II-IV with one spinoid posterior granule ............ 6
Coloration reddish yellow; dorsal carinae of metasomal segments II-IV

with 2 to 6 strong spinoid posterior granules............. Grosphus simoni
Coloration yellowish; dorsal carinae of metasomal segments II-IV with

one small spinoid posterior granule ................ Grosphus darainensis
Coloration reddish to dark brown; metasomal segments II-IV with one

strong. spinoid. posterior eranuler . |. jee ee G. madagascariensis
Coloration blackish or reddish brown to yellowish; pectines with 30 to

40) teeth:sadult-bodyclenethumore than 90 mme ann ad er Re 8
Coloration reddish brown to yellowish, never blackish; adult body

lensthvless:than!9 Olmm atto Te RE Pe Samson EN HEN 9
Coloration blackish throughout." iO G. grandidieri
Coloration'reddish:brown'to yellowish’. +. 2:2... fs. se ae ae: G. ankarana
Mesosoma with homogenous coloration, reddish brown or yellowish...... 10
Mesosoma with a blackish longitudinal median band, or with two

blackishiongitudinalilateralibands. se a nr ea. 15

Adult total length more than 65 mm; mesosoma reddish brown . G. flavopiceus
Adult total length less than 60 mm; mesosoma dark yellow; metasomal

segment V and telson with or without blackish spots .................. 11
Metasomal segments and telson without blackish spots ................ 12
Metasomal segments and telson totally blackish or with blackish spots . .... 13
Metasomal segments yellowish, with moderately to strongly developed

CALIMAC (ON MUONI RA G. intertidalis

Metasomal segments reddish yellow; rounded and with vestigial carinae
Bad. MENT uz Anti MANUEL pri di OST MEER Grosphus mahafaliensis

Metasomal segment V and telson with blackish spots .......... G. annulatus
Metasomal segment V and telson totally blackish ..................... 14
Telson with aculeus of the same length or shorter than the vesicle..... G. feti
Helson:withaculeus lonsenthan the vesicle Rote e G. olgae

Mesosoma with a wide blackish longitudinal median band; basal middle
lamellae of female pectines three times longer than wide at their base
and'coverne the tour proximal teeth sa E ete tere G. limbatus
Mesosoma with two narrow blackish longitudinal lateral bands.......... 16
Carapace without a blackish triangular spot; basal middle lamellae of
female pectines weakly curved, widening in proximal half and covering
thestwo,proximaliteether 4... cy rtl G. bistriatus
Carapace with a blackish triangular spot; basal middle lamellae of the
female pectines curved and constantly narrowing from the base to apex,
cCovennorthe four proximaliteethe sara E Roe eae ie G. ankarafantsika

260 W. R. LOURENCO & S. M. GOODMAN

ACKNOWLEDGEMENTS

We are grateful to Mr CI. Ratton and Mrs Florence Marteau, Natural History
Museum, Geneva for the preparation of the photos, and the arrangements of the plates,
respectively. A portion of the material used in this study was obtained during field
missions financed by the Volkswagen Foundation and Conservation International.

REFERENCES

DORR, L. J. 1997. Plant collectors in Madagascar and the Comoro Islands. Kew: Royal Botanic
Gardens, XLVI + 524 pp.

FAGE, L. 1929. Les Scorpions de Madagascar. Faune des Colonies françaises 3. Société
d’Editions Géographiques Maritimes Coloniales, Paris, pp. 637-694.

GERVAIS, P. 1843. Les principaux résultats d’un travail sur la famille des Scorpions. Société
philomatique de Paris. Extraits des procès-verbaux des séances 5 (7): 129-131.

GERVAIS, P. 1844. Remarques sur la famille des Scorpions. Archives du Muséum d’Histoire
Naturelle, Paris 4: 201-240.

KRAEPELIN, K. 1899. Scorpiones und Pedipalpi. /n: F. DAHL (ed). Das Tierreich. Herausgegeben
von der Deutschen zoologischen Gesellschaft. 8 (Arachnoidea). R. Friedländer und
Sohn Verlag, Berlin, 265 pp.

KRAEPELIN, K. 1900. Ueber einige neue Gliederspinnen. Abhandlungen aus dem Gebiete der
Naturwissenschaften. Herausgegeben vom naturwissenschaftlichen Verein in Hamburg
16, 1 (4): 1-17.

KREMEN, C. 2003. The Masoala Peninsula (pp. 1459-1466). In: GOODMAN, S. M. & BENSTEAD,
J. P. (eds). The natural history of Madagascar. The University of Chicago Press, Chicago.

LOURENÇO, W. R. 1996. Scorpions. Faune de Madagascar. Museum national d’Histoire naturelle,
Paris, 102 pp.

LOURENCO, W. R. 2003. New taxonomic considerations on some species of the genus Grosphus
Simon, with description of a new species (Scorpiones, Buthidae). Revue suisse de
Zoologie 110 (1): 141-154.

LOURENÇO, W. R. & GOODMAN, S. M. 2002. Scorpions from the Daraina region of northeastern

Madagascar with special reference to the family Heteroscorpionidae Kraepelin, 1905.
Revista Ibérica de Aracnologia 6: 53-68.

LOURENÇO, W. R. & GOODMAN, S. M. 2003. Description of a new species of Grosphus Simon
(Scorpiones, Buthidae) from the Ankarana Massif, Madagascar. Revista Ibérica de
Aracnologia 7: 19-28.

LOURENÇO, W. R., GOODMAN, S. M. & RAMILIJAONA, O. 2004. Three new species of Grosphus
Simon from Madagascar (Scorpiones, Buthidae). Revista Ibérica de Aracnologia 9:
225-234.

NICOLL, M. E. & LANGRAND, O. 1989. Madagascar. Revue de la conservation et des aires pro-
tégées. WWE, Gland, Switzerland, XVII + 374 pp.

RAMANGASON, G. S. 1988. Flore et végétation de la Forêt d’ Ampijoroa (pp. 130-137). In:
RAKOTOVAO, L., BARRE, V. & SAYER, J. (eds). L’équilibre des écosystèmes forestiers à
Madagascar. IUCN, Gland.

SIMON, E. 1880. Etudes arachnologiques, 12° Mémorie (1). XVIII. Descriptions de genres et
espèces de l’ordre des Scorpiones. Annales de la Société entomologique de France, sér.
5, 10: 377-398.

VACHON, M. 1963. De l’utilité, en systématique, d’une nomenclature des dents des chélicères

chez les Scorpions. Bulletin du Muséum national d'Histoire naturelle, Paris, 2e sér., 35
(2): 161-166.

STATUS OF GROSPHUS MADAGASCARIENSIS AND GROSPHUS HIRTUS 261

VACHON, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de
Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux
et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum national
d Histoire naturelle, Paris, 3e sér., n° 140, Zool. 104: 857-958.

VACHON, M. 1975. Sur l’utilisation de la trichobothriotaxie du bras des pédipalpes des Scorpions

(Arachnides) dans le classement des genres de la famille des Buthidae Simon. Comptes
Rendus des Séances de l'Académie des Sciences 281 (D): 1597-1599.

units Sole Slate i

ehr

n Mic. LR ori

REVUE SUISSE DE ZOOLOGIE 113 (2): 263-268; juin 2006

Vitronura mascula, a new species of Neanurinae
(Collembola: Neanuridae) from northern Vietnam, with a key
to the species of the genus

Adrian SMOLIS! & Louis DEHARVENG?

! Zoological Institute, University of Wroctaw, Przybyszewskiego 63/77,
51-148 Wroctaw, Poland. E-mail: adek@biol.uni.wroc.pl

2 Museum National d’Histoire Naturelle, Laboratoire d’Entomologie, FRE2695 du
CNRS, 45 rue Buffon, F-75005 Paris, France. E-mail: deharven@mnhn.fr

Vitronura mascula, a new species of Neanurinae (Collembola:
Neanuridae) from northern Vietnam, with a key to the species of the
genus. - A new species of the genus Vitronura Yosii, 1954 sensu Cassagnau,
1986 from northern Vietnam is described and illustrated. An identification
key for all the known species of the genus is given.

Keywords: Collembola - Neanuridae - taxonomy - Vietnam.

INTRODUCTION

Yosii (1969) established the subgenus Vitronura in the genus Neanura
MacGillivray, 1893 and designated Neanura mandarina Yosii, 1954 from Japan as its
type species. Cassagnau (1980) raised Vitronura to generic status and later (1983) clas-
sified the taxon to a lineage of blasconurian Neanurinae. At present the genus is
included in the tribe Paleonurini (Cassagnau, 1989). The presence of separate tubercles
Fr and An on dorsal side of the head distinguish Vitronura from all genera of
Paleonurini. As presently defined, the genus comprises 14 species, occurring mainly in
Southeast and Eastern Asia.

Till now only one cosmopolitan species, V. giselae (Gisin, 1950), was recorded
from Vietnam (Nguyen, 1995). Examination of the material of Collembola from
northern Vietnam revealed one new species of the genus. The present paper contains
its description and an updated key to all members of the genus.

TERMINOLOGY

The terminology and layout of the tables used in this paper follow Deharveng
(1983), Deharveng & Weiner (1984) and Greenslade & Deharveng (1990), and the
following abbreviations are used:

GENERAL MORPHOLOGY: abd. — abdomen, ant. — antenna, Cx — coxa, Fe — femur,
Scx2 - subcoxa 2, T — tibiotarsus, th. — thorax, Tr — trochanter, VT — ventral tube.

Manuscript accepted 10.03.2005

264 A. SMOLIS & L. DEHARVENG

GROUPS OF CHAETAE: Ag — antegenital, An. — anal, Fu — furcal, Ve — ventroex-
ternal, Vi — ventrointernal, V1 — ventrolateral.

TUBERCLES: An — antennal, Fr — frontal, Cl — clypeal, De — dorsoexternal, Di —
dorsointernal, DI — dorsolateral, L — lateral, Oc — ocular, So — subocular.

TYPES OF CHAETAE: MI — long macrochaeta, Mc — short macrochaeta, me —
mesochaeta, mi — microchaeta, ms — s-microchaeta, S — or s — chaeta sensualis or sen-
sillum, or — organite of antenna IV, i — ordinary chaeta on antenna IV, mou — thin cylin-
drical sensilla on ant. IV (,,soies mousses”), x — labial papilla x.

DESCRIPTION OF THE NEW SPECIES

Vitronura mascula sp. n.

TYPE MATERIAL: Holotype, adult male on slide; northern Vietnam, Tam Dao
National Park, the top of Tam Dao mount (1300 m. a. s. 1.), 08.04.1997, mosses and
lichens on rhododendron branches, leg. R. J. Pomorski. Paratype, 1 subadult male, the
same data as holotype. Holotype is preserved in collection of the Department of
Biodiversity and Evolutionary Taxonomy, Wroctaw University, Poland; paratype
housed in the collection of the Muséum d’histoire naturelle in Geneva.

ETYMOLOGY: Among known members of the genus, only males of the new
species have male (Latin word "masculus") ventral organ.

DIAGNOSIS: Vitronura mascula sp. n. differs from other Vitronura species in the
presence of male ventral organ and strong reduction of dorsal chaetotaxy on head
(absence of chaetae: O, C, D, E, Oca, Ocp, Di2, De2). In addition the new species is
characterised by ogival labrum, claw without inner denticle and separate tubercles Di
on head and abd. V.

DESCRIPTION: Body length (without antennae): holotype - 1.05 mm, paratype -
1.1 mm. Habitus typical of Paleonurini (Cassagnau, 1989). Tertiary granulation well
developed. Colour of the body in alcohol white. 2+2 unpigmented eyes (Fig. 1).

Types of dorsal ordinary chaetae: macrochaetae MI thickened, moderately long,
narrowly sheathed, the distal three-fourth densely furnished with short pointed scale-
like (teeth) which are extremely dense at the tip (Fig. 4), macrochaetae Mc with similar
morphology, microchaetae very short, smooth, without teeth.

Head. Buccal cone slightly elongate. Labrum pointed, with ventral scleri-
fications ogival as in Figs 2, 3. Labrum chaetotaxy 0 / 2, 4. Chaetotaxy of labium as in
Fig. 3 and Tab. la. Maxilla styliform, mandible thin and tridentate. Chaetotaxy of
antennae as in Tab. la. Apical bulb distinct, trilobed. S-chaetae subequal, long and
moderately thickened. Chaetotaxy of head as in Fig. 1 and in Tab. 1a. Chaeta O absent.

Thorax, abdomen, legs. Chaetotaxy of th. and abd. as in Figs 1, 8 and in Tab. 1b.
Tubercles Di on th. I-III with 2 chaetae. Male ventral organ fully developed only in
adult male with ductus ejaculatorius (in subadult poorly visible), built of thickened and
slightly forked chaetae on abdominal sterna III, IV, V and VI (Fig. 8). Chaetae of
ventral organ in subadult with similar morphology as microchaetae (not thickened and
not forked), but with distinctly greater chaetopor (Fig. 6). Tubercles L on abd. IV and

VITRONURA MUSCULA A NEW SPECIES FROM NORTHERN VIETNAM 265

Table 1. Chaetotaxy of Vitronura mascula sp. n.
a) Cephalic chaetotaxy

Tubercle Number of chaetae Types of chaetae Names of chaetae
CI 4 MI F
mi G
An 1 MI B
Fr 2 MI A
Oc 1 MI Ocm
Di 1 MI Dil
De 1 MI Del
(DI+L+So) 9 MI, Mc, mi impossible to recognise

Number of other cephalic chaetae: Vi, 5; Ve, 7; labrum, 0 / 2, 4; labium, 9, 0x; ant. I, 7; ant. II,
11; ant. III, 17 + 5s; ant. IV, 8S +1 + or + 12mou.

b) Postcephalic chaetotaxy

Terga Legs

Di De DI IG Scx2 Cx Tr Fe T
th.I 1 1(2) 1 = 0 3 6 13 18
th. II 2) 2+s 2+stms 3 2 7 6 12 18
th. II 2 3+s 3+s 3 2 8 6 11 17

Sterna

abd. I 2 2+s 2 3 VT: 4
abd. II À 2+s 2 3 Ve: 2 Vel- absent
abd. III 2 2+s 2 3 Ve: 3-4 Fu:4 me Omi
abd. IV D, 2+s 2, 5 Ve: 8-9 VI: 4
abd. V 1(2) 5+s 3 Ag: 3
abd. VI U Ve:14 An.: 2mi

V respectively with 5 and 3 chaetae (Fig. 5). Cryptopygy absent. Chaetotaxy of legs as
in Tab. 1b. Claw relatively long, without inner tooth (Fig. 7).

REMARKS: The new species appears to be the closest to V. tubercula Lee & Kim,
1990 from Taiwan, because both species have similar morphology of macrochaetae
and claws without teeth. Virronura mascula sp. n. differs from the mentioned species
in separate tubercles Di on head (in tubercula fused), number of chaetae Di of th. II-III
(in mascula: 2, in tubercula: 3) and number of chaetae De of abd. I-III (in mascula:
2+s, in tubercula: 3+s).

The genus Vitronura was first keyed by Cassagnau & Deharveng (1981).
Changes in definition of the genus (Cassagnau, 1986) and four species described after
that date make it necessary to elaborate an updated key. The key includes among others
species Vitronura giselae, V. mandarina and V. dentata, which were formally
synomynyzed by Yoshii (1995). This author considered V. giselae and V. mandarina
as synonyms of rosea, a species very succinctly described from Paris by Gervais in
1842; two species of yellow to pink color exist there, i.e. Vitronura “giselae” Gisin and

266 A. SMOLIS & L. DEHARVENG

ESS

OE E

SEE
MES A
COPS

st ;
ERA, IS
a : È
LINE ED,
SPAR & CPN, De)
\ ER
A \ ;

Fics 1-8

Vitronura mascula sp. n.: 1 - habitus and dorsal chaetotaxy (holotype), 2 - chaetotaxy of labrum,
3 - chaetotaxy of labium and group Vi, 4 - tubercle De of abd. IV, 5 - chaeta of male ventral organ
(adult male), 6 - chaeta of male ventral organ (subadult male), 7 - tibiotarsus and claw of legs III
(lateral view), 8 - chaetotaxy of abdominal sterna II-VI.

VITRONURA MUSCULA A NEW SPECIES FROM NORTHERN VIETNAM 267

Monobella grassei Denis, and there is no conclusive evidence that rosea correspond to
one rather than the other of them. We therefore continue to consider Anoura rosea
Gervais, 1842 as species inquirenda. As differential characters between giselae and
mandarina are not discussed by Yoshii, the synonymy he proposed cannot be accepted
at the moment. For the same reason, the synonymy between dentata Deharveng &
Weiner, 1983 and pygmaea Yosii, 1954 proposed in the same paper (Yoshii, 1995)
cannot be retained. Differential characters between these species are given in the key.

KEY TO THE SPECIES OF THE GENUS VITRONURA

1 HHubercle Oc'on head\witn lKor2’chaetaern.N. tae ee ee ep, 2
- hubercleOcionhead withis chactae een Re ea 10
2 tubercle Oconihead: wath Wchacta nga es Rane V. mascula sp. n.
- hubercleiOc oncshead wathi2 chactae acer eee Ne 3
3 Tubercles Di on head fused along midline . . .. V. tubercula Lee & Kim, 1990
- ilubercles Dion headiseparate tiie). nr... is eae RON LET 4
4 Chacta! © onthead, present ee. i. ty. LI SRL, er: se U
- @haetal@:onkhead.absent Hrn I ae. AE 5
5 Tubercles Di on abd. V fused along midline ........... V. sinica Yosii, 1976
- MU DELCIESDITONTADAAVESE PATATA LIE Re TRE 6
6 Tubercle De of th. V with 5 chaetae ............... V. luzonica Yosii, 1976
- Tubercle De of th. V with 4 chaetae V. gressitti Cassagnau & Deharveng, 1981
7 Tubercle Dirot ths with 2chaetae ss ose eines ek V. latior (Rusek, 1967)
- JiuberclesDirofithplwithylschacta, ge jee RS ee aici ee 8
8 ClawawathtinnerGenticle ROME AMENER opt) CREATE ORO REA Ur 9

@lawzwithoufinnerdentiele . 2... cc. V. mandarina (Yosii, 1954)
9 Tubercle Di of abd. V with 2 chaetae........... V. namhaiensis (Lee, 1974)
- diubercle Divot abd: V with 3:chaetaell.. are. V. pygmaea (Yosii, 1954)
10 Tubercle De on head with 2 chaetae.............. V. joanna (Coates, 1968)
- lubercleDeroniheadiwith}3YCha6t2e: Pr OR NAME A III I e 11
11 Tubercles Dionheidiused'alonsmidhine e 2a... 12
- Hubercies Dionfheadiseparate ne ln. m OT 13
12 Chaeta Olon headipresent ann. nn. V. macgillvrayi (Denis, 1933)
- Chaeta OloniÉad absent inn V. singaporiensis (Yossi, 1956)
13 Claw with inner denticle ............ V. dentata Deharveng & Weiner, 1984
- @lawawithoutiinnerdentiele RR I 14
14 abrutis AIN sival'E fe em. aes V. acuta Deharveng & Weiner, 1984
- TE ADEUMENONZO IVA EEE V. giselae (Gisin, 1950)
ACKNOWLEDGEMENTS

We are greatly indebted to Prof. Romuald J. Pomorski for the loan of material
of V. mascula. Specimens of the species were collected during faunistic investigations
of North Vietnam, sponsored by Museum and Institute of Zoology PAS (Warsaw) and
Wroctaw University (project number 1018/W/IZ/97). The studies of this material were
financed by the University of Wroctaw (project number 2020/W/1Z/2001).

268 A. SMOLIS & L. DEHARVENG

REFERENCES

CASSAGNAU, P. 1980. Nouveaux critères pour un redécoupage phylogénétique des Collemboles
Neanurinae (s. Massoud 1967). In: DALLAI, R. (ed.). Ist International Seminar on
Apterygota, Siena 1978: 115-132.

CASSAGNAU, P. 1983. Un nouveau modèle phylogénétique chez les Collemboles neanurinae.
Nouvelle Revue d’Entomologique 13: 3-27.

CASSAGNAU, P. 1986. Sur l’évolution des Neanurinae paucituberculés à pièces buccales réduites
(Collemboles). Jn: DALLAI, R. (ed.). 2nd International Seminar on Apterygota, Siena:
313-317.

CASSAGNAU, P. 1989. Les Collemboles Neanurinae; éléments pour une synthèse phylogénétique
et biogéographique (pp.171-182). In: DALLAI, R. (ed.). 3rd International Seminar on
Apterygota, Siena: 171-182.

CASSAGNAU, P. & DEHARVENG, L. 1981. Sur le genre Vitronura (Collemboles Neanuridae):
aspect systématique et approche cytogénétique. Bulletin du Museum National d’Histoire
Naturelle, Paris 4 (3): 151-173.

DEHARVENG, L. 1983. Morphologie évolutive des Collemboles Neanurinae en particulier de la
lignée Neanurinae. Travaux du Laboratoire d’Ecobiologie des Arthropodes Edaphiques,
Toulouse 4 (2): 1-63.

DEHARVENG, L. & WEINER, W. 1984. Collemboles de Corée du Nord III - Morulinae et
Neanurinae. Travaux du Laboratoire d’Ecobiologie des Arthropodes Edaphiques,
Toulouse 4 (4): 1-61.

GREENSLADE, P. & DEHARVENG, L. 1990. Australian species of the genus Australonura
(Collembola, Neanuridae). Invertebrate Taxonomy 3: 565-593.

LEE, B-H. & Kim, J-T. 1990. Systematic Studies on Chinese Collembola (Insecta) II. Five New
Species and Two New Records from Taiwan in the Family Neanuridae. The Korean
Journal of Systematic Zoology 6 (2): 235-250.

NGUYEN TRI TIEN, 1995. Species composition and structure of Collembola in North Vietnam. In:
Selected collection of scientific reports on ecology and biological resources. Vietnam
National Center for Natural Science and Technology, Institute of Ecology and Biological
Resources, Hanoi: 578-585.

YosH, R. 1995. Identity of some Japanese Collembola III. Acta Zoologica Asiae Orientalis 3:
50-68.

Yost, R. 1969. Collembola-Arthropleona of the IBP-Station in the Shiga Heights, Central Japan,
I. Bulletin of the National Science Museum Tokyo 12 (3): 531-556.

REVUE SUISSE DE ZOOLOGIE 113 (2): 269-286; juin 2006

The Palaearctic triozids associated with Rubiaceae
(Hemiptera, Psylloidea): a taxonomic re-evaluation of the
Trioza galii Foerster complex

Daniel BURCKHARDT! & Pavel LAUTERER?

! Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland.
E-mail: daniel.burckhardt@unibas.ch

2 Moravian Museum, Department of Entomology, Hviezdoslavova 29a,
CZ-62700 Brno-Slatina, Czech Republic. E-mail: ento.laut@ volny.cz

The Palaearctic triozids associated with Rubiaceae (Hemiptera,
Psylloidea): a taxonomic re-evaluation of the Trioza galii Foerster
complex. - The Palaearctic Trioza galii complex is revised to contain 7. coc-
quempoti sp. n., T. drosopouli sp. n., T. galii Foerster and 7. velutina
Foerster stat. rev., with following new synonymies: 7. rubiae Baeva and
T. rubicunda Loginova = T. galii and Trioza distincta Flor = T. velutina,
respectively. 7. galii f. spinogalii Sulc and T. galii f. aspinovelutina Sulc are
considered to be of infrasubspecific rank and thus unavailable. Trioza
velutina var. thoracica Flor is an available name regarded as nomen
dubium. Lectotypes are designated for Trioza galii and T. velutina. Adults
and last instar larvae (except 7: velutina) are diagnosed and illustrated.
Identification keys are provided. All four species are associated with
Rubiaceae on which they induce galls. Confirmed hosts of 7: galii are
Galium spp. and Asperula cynanchica, literature records also suggest
Sherardia arvensis and Rubia spp. T. cocquempoti and T. drosopouli
develop on Rubia spp. T. velutina is probably associated with Galium spp.
but hard evidence is not available. The 7: galii complex is diagnosed using
adult and larval charcters. The definition is similar to Conci's (1992)
concept of Spanioza but excludes S. tamaninii. S. tamaninii is morpho-
logically intermediate between the 7: galii complex and the 7. centranthi
complex (associated with Valerianaceae) which may be sister groups. The
synonymy of Spanioza with Trioza is confirmed, and the new combination
Trioza tamaninii is proposed.

Keywords: Hemiptera - Psylloidea -Triozidae - taxonomy - new taxa -
Rubiaceae - Palaearctic.

INTRODUCTION

Species of jumping plant-lice are generally well-defined by their adult and

larval morphology as well as their host plants. In particular, the male terminalia are
routinely used for species identification. The Palaearctic Trioza galii Foerster, as

Manuscript accepted 02.12.2005

270 D. BURCKHARDT & P. LAUTERER

currently perceived, is a notable exception as the male genital morphology exceeds the
usual variability encountered in other Trioza species.

Foerster (1848) described both 7: galii and 7. velutina from German and Irish
localities; some material of 7. galii was collected on Galium verum. The descriptions
contain colour and forewing characters but are not diagnostic. Hardy (1853) recorded
deformations on Galium aparine which he attributed to "Psylla" velutina. He said that
velutina appears not to differ from "Psylla" galii. Flor (1861a), providing detailed
descriptions and diagnoses, treated 7. galii and T. velutina as distinct species, pointing
out differences in the male paramere. He also assigned a single female specimen from
Southern France with a lighter thorax and clear forewing colour, to the variety Trioza
velutina var. thoracica. Again on the basis of a single female, he described Trioza
distincta from Thuringia, differing from 7. velutina in the slightly different forewing
shape (Flor, 1861b). Flor's more restricted species concept was confirmed by Löw
(1882) who examined some of Foerster's and Flor's types. Lethierry (1874), Oshanin
(1907) and Aulmann (1913) followed Löw, whereas Scott (1876) and Edwards (1896)
adopted Hardy's view, treating 7. galii and velutina as synonyms. Sulc (1913) pointed
out that his earlier description (Sulc, 1910) of 7. galii corresponded with the types of
T. velutina and having examined types of 7. galii, T. velutina and T. distincta, plus other
material, reached the same conclusion as Hardy (1853). Thus, he treated 7. velutina, T.
distincta and questionably var. thoracica, the type of which appears to be lost, as
synonyms of 7. galii. Based on presence or absence of surface spinules as well as
colour and shape of the forewing, he defined four forms: galii forma typica, f. spino-
galii Sulc, f. velutina and f. aspinovelutina Sulc. Boselli (1930) found many specimens
in Italy on Rubia peregrina which he referred to Spanioza galii aspinovelutina. He
gave detailed descriptions and illustrations of the adults, larvae and eggs and provided
information on their biology. The male paramere of Boselli's material, however, is
more massive than those described and illustrated by Sulc (1910, 1913). 7. galii, in this
broad definition, has been reported from all over the Palaearctic, including Japan and
Taiwan (Aulmann, 1913; Klimaszewski, 1973; Gegechkori & Loginova, 1990;
Ossiannilsson, 1992). According to these authors, the species is associated with
Galium, Sherardia and Rubia spp. (Rubiaceae), where it induces characteristic defor-
mations on the buds, shoots and leaves, which can be above or below ground (e.g.
Houard, 1909; Docters van Leeuwen, 1937).

Adults are regularly encountered by sweeping herbaceous vegetation, although
rarely on the host plants. Larval material, by contrast, is generally difficult to find. The
first larval description referred to 7 galii is by Kieffer (1889) who examined material
from Sherardia arvensis but his description is not diagnostic. The dorsum of the larva
is said to be dark brown. A more detailed larval description referred to T: galii is that
by Boselli (1930). White & Hodkinson (1981) used Boselli's description in their hand-
book of the British species, and Ossiannilsson (1992) in the Fauna Scandinavica
described Italian rather than Scandinavian specimens.

Three additional Palaearctic species are reported from Rubiaceae. Baeva (1972)
described Trioza rubiae from Turkmenia based on a single series collected on Rubia
florida, which she related to Trioza foersteri Meyer-Diir and Trioza rotundata Flor.
Loginova (1978) added Trioza rubicunda, again based on a single series, collected in

PALAEARCTIC TRIOZIDS ON RUBIACEAE ZIA

Kazakhstan on Galium sp. According to Loginova 7. rubicunda is closest to T. galii f.
velutina from which it differs in the reduced surface spinules and a broader paramere.
T. rubiae and T. rubicunda have not been subsequently reported. Conci (1992)
described Spanioza tamaninii from Italian and Slovakian specimens, one of which was
collected on Galium anisophyllon. He suggested that S. tamaninii is closely related to
galii and that its host may be Galium.

Trioza galii was selected by Enderlein (1926) as type species of Spanioza, a
genus which he erected for triozids with a short Rs vein in the forewing. The artificial
nature of this concept was pointed out by Tuthill (1943) who synonymised it with
Trioza. Conci (1992) considered Spanioza a valid genus and redefined it to include
Trioza galii and its "formae", Trioza rubiae Baeva, Trioza rubicunda Loginova and
Spanioza tamaninii Conci. This was not followed by Burckhardt & Couturier (1994)
who treated the Trioza galii group as part of the large artificial genus Trioza.

Over the last few years collections from the Mediterranean as well as Central
and Western Europe yielded material including larvae and host records which permit a
re-evaluation of the taxonomy of the Trioza galii complex. Wagner & Franz (1961)
suggested that 7. galii in the broad definition by Sulc (1913) is a species complex, and
Conci (1992) wrote that Trioza galii and its "formae" require revision. The present
paper provides this revision and re-examines the validity of Spanioza.

MATERIAL AND METHODS

Material has been examined or is mentioned from following institutions:

BMNH _ Natural History Museum, London

DEZA Dipartimento di Entomologia e Zoologia Agraria dell'Universita, Portici
EAUT Institute of Zoology and Botany, Estonian Agricultural University, Tartu
MHNG Muséum d'histoire naturelle, Geneva

MMBC . Moravské Muzeum, Brno

MNHN Museum national d'histoire naturelle, Paris

NHMB Naturhistorisches Museum, Basel

NHMV Naturhistorisches Museum, Vienna

SMTD Staatliches Museum fiir Tierkunde, Dresden

ZISP Zoological Institute, St. Petersburg

ZMHB Museum fiir Naturkunde der Humboldt-Universitàt, Berlin

ZSSM Zoologische Staatssammlung, Munich.

The morphological terminology mainly follows Ossiannilsson (1992).
Drawings were prepared from cleared specimens mounted permanently in Euparal or
Canada Balsam or temporarily in glycerine. Measurements were taken from slide
mounted specimens, except for the total body length which is taken from dry mounted
specimens measured from the apex of head to the apex of forewing when folded over

the body.
RESULTS

The Trioza galii Foerster, 1848 species complex

DESCRIPTION
Adult. Coloration mostly dark brown to black, in 7. cocquempoti and
T. drosopouli thorax yellowish, brown or dark dorsally, sometimes with longitudinal

272 D. BURCKHARDT & P. LAUTERER

stripes; area around wing insertion yellowish or brown; intersegmental membranes
reddish; antennal segment 3, tibiae and tarsi whitish. Forewing almost colourless or
with yellowish or light brown tinge; veins yellowish or light brown, vein C+Sc often
brown or dark brown in basal part. Young specimens lighter.

Head slightly narrower than mesonotum, weakly inclined from longitudinal
body axis, at a 45° angle. Vertex shiny, weakly produced anteriorly on either side of
mid-line. Genal processes variable in size and shape, conical, subacute, ranging from
one to two thirds vertex length. Antenna 10-segmented, segment 3 longer than seg-
ments 4 and 5 together; segment 10 bearing one long and one very short terminal seta
respectively (Fig. 21). Clypeus short, pyriform. Thorax weakly curved dorsally.
Metacoxa with horn-shaped, subacute meracanthus; metatibia weakly widened basally
and apically, with some minute basal spines and 1+3 sclerotised apical spurs; metaba-
sitarsus without sclerotised spurs. Forewing (Figs 1-4) transparent, variable in size,
shape and colour, elongate, strongly pointed, angular or bluntly angular apically; vein
Rs short, bifurcation of vein M slightly or strongly distal to line linking apices of veins
Rs and Cuj,. Wing membrane transparent; apart from wing base usually lacking sur-
face spinules, with the exception of 7. velutina where spinules are always present,
forming more or less extended fields. Radular spinules forming narrow stripes in cells
mj, my, and cu. Hindwing membranous, about two thirds forewing length. Costal
setae ungrouped; R+M+Cu, indistinctly trifurcating. Abdominal tergite 3 (and occa-
sionally 4) in male and tergite 4 (and occasionally 5) in female bearing a row of lateral
setae. Male proctiger (Figs 5, 8, 11, 16) 1-segmented, weakly produced or lobed
posteriorly, covered in moderately long setae apically and posteriorly. Subgenital plate
subglobular, with a few long setae laterally and ventrally. Paramere (Figs 6, 9, 12, 13,
17) shorter than proctiger, massive or lamellar; sparsely covered in long setae on outer
face apically and along hind margin; inner face covered in long thin setae mainly
basally, apically and along hind margin, with a group of shorter, stout setae along fore
margin in basal half. Proximal portion of aedeagus strongly curved at base, almost
straight apically; distal portion (Figs 7, 10, 14, 15, 18) relatively short with large apical
dilatation which varies from almost rectangular to distinctly hooked. Sclerotised end
tube of ductus ejaculatorius short, weakly curved. Female proctiger (Figs 23-26) short
with relatively large circumanal ring. Subgenital plate short.

Fifth instar larva (Fig. 28). Coloration varying from uniformly yellow or light
ochreous to yellow with more or less extended pattern on head, thorax, wing buds and
abdomen to completely dark. Body oval, bearing a single row of densely spaced, lateral
truncate sectasetae. Antenna 6-segmented. Head broadly rounded anteriorly. Legs with
short claws and small, fan-shaped tarsal arolium. Humeral lobes blunt, relatively short.
Circumanal ring small, with a single row of pores. Dorsum of body lacking macro-
scopic setosity. Tarsal arolium transversely oval, lacking pedicel, with unguitractor
(Figs 29-31). Claws relatively small.

Egg. Spindle-shaped, longer than three times its width, with short basal pedicel
and no apical filament. The egg is laid perpendicularly on the leaf margins.

PALAEARCTIC TRIOZIDS ON RUBIACEAE 279

Fics 1-4
Forewing of Trioza spp. 1 — T. cocquempoti; 2 — T. drosopouli; 3 — T. galii; 4 — T. velutina.

DISCUSSION

The 7. galii complex, as defined above, constitutes a small, probably mono-
phyletic group within the large artificial genus Trioza. It is restricted to the Palaearctic.
It may be most closely related to the 7. centranthi complex (7. centranthi (Vallot),
Trioza nana Gegechkori and 7. valerianae Gegechkori), associated with Valerianaceae.
Adult 7. galii and T. velutina resemble T. centranthi (Vallot) in morphology, size and
colour. The distribution of surface spinules on the forewing is variable in 7. centranthi.
They may form relatively extended fields in all cells except for c+sc or they may be
restricted to cell cup. In 7. centranthi vein Rs of the forewing tends to be relatively long
and sinuous, and the bifurcation of vein M lies near the line joining the apices of veins
Rs and Cu] ,; in 7. galii and T. velutina in contrast vein Rs is shorter and almost straight
or concave; the bifurcation of vein M is more distant to the line joining the apices of
veins Rs and Cuj,. T. centranthi bears terminal setae on antennal segment 10, with the
shorter seta about half as long as the longer one (Fig. 22), in the 7. galii complex the
shorter seta is only about a quarter to a third as long as the longer (Fig. 21). The 7. cen-
tranthi complex is characterised by the irregularly triangular paramere, which some-
times bears a large antero-apical lobe (absent in the 7. galii complex), and by the apical
dilatation of the distal portion of the aedeagus, which bears small lateral spines (7. cen-
tranthi, Fig. 20) (absent in the 7. galii complex, Figs 7, 10, 14, 15, 18). The female sub-
genital plate of 7. centranthi (Fig. 27) has, on either side, a group of long setae near the
dorsal margin, which are longer than the other hairs (hairs laterally, ventrally and
terminally are of about the same length in the 7. galii complex, Figs 23-26).

The present concept of the 7 galii complex differs in two major details from
that of Conci (1992). In addition to 7. galii, T. rubiae and T. rubicunda, Conci included
Spanioza tamaninii in the group. S. tamaninii has so far been recorded from a few spe-
cimens from Italy and Slovakia only. It differs from the 7. galii complex and the 7. cen-
tranthi complex in having antennae which have light segments 3-8 rather than dark
segments with a strongly contrasting light segment 3. The paramere shape, the termi-

274 D. BURCKHARDT & P. LAUTERER

Fics 5-10

Male genital structures of Trioza spp. 5-7 — T. cocquempoti; 8-10 — T. drosopouli. 5, 8 —
Terminalia, in profile, scale a; 6, 9 — paramere, inner surface, scale b; 7, 10 — distal portion of
aedeagus, scale b.

nal antennal setae and probably also the lateral setosity on the female subgenital plate
are as in species of the 7. centranthi complex. Apart from the antennal colour S.
tamaninii differs from the 7. centranthi complex in the apex of the aedeagus (Fig. 19)
lacking lateral spines (cf. Fig. 20 for 7. centranthi). Incidentally Conci's drawing of the
aedeagus appears to be artefactual. Larval material is necessary to determine the
phylogenetic relationship of S. tamaninii.

PALAEARCTIC TRIOZIDS ON RUBIACEAE 275

Fics 11-20

Male genital structures of Trioza spp. 11-15 — T. galii; 16-18 — T. velutina; 19 — T. tamaninii; 20
— T. centranthi. 11, 16 — Terminalia, in profile, scale a; 12, 13, 17 — paramere, inner surface, scale
b; 14, 15, 18-20 — distal portion of aedeagus, scale b.

Conci (1992) resurrected the genus Spanioza for the 7. galii complex and
S. tamaninii. He neither provided convincing autapomorphies for Spanioza nor did he
define a sister group. This makes his Spanioza highly artificial and without phyloge-
netic significance. We follow Burckhardt & Couturier (1994) who treated Spanioza as
a synonym of Trioza. Here we formally transfer Spanioza tamaninii to Trioza as T.
tamaninii (Conci) comb. n.

TAXONOMIC TREATMENT

Key for the identification of the species of the Trioza galii complex

Adults

1 Antenna longer than 1.1 mm. Male proctiger strongly produced posteri-
orly (Figs 5, 8). Paramere (Figs 6, 9) massive, blunt apically. Apex of
distal portion of aedeagus subrectangular or reniform (Figs 7, 10).
Ventral and posterior margin of female subgenital plate, in profile (in
cleared specimens), forming an angle > 90° (Figs 23, 24)............... 2

276 D. BURCKHARDT & P. LAUTERER

- Antenna shorter than 1.1 mm. Male proctiger tubular, weakly produced
posteriorly (Figs 11, 16). Paramere (Figs 12, 13, 17) slender, with apical
digitiform process. Apex of distal portion of aedeagus hooked (Figs 14,

15, 18). Ventral and posterior margin of female subgenital plate, in pro-
file (in cleared specimens), forming an angle < 90° (Figs 25, 26).......... 3

2) Male proctiger with narrow posterior lobe (Fig. 5). Paramere (Fig. 6), in
profile, with narrow angular bulge above the middle along the fore
margin; apex angular. Distal portion of aedeagus (Fig. 7) with reniform
apical dilatation. Female terminalia as in Fig. 23. Algeria, France, Italy,
NIOLOCEOHSPAM ee PEN ee A e A re eae ete T. cocquempoti Sp. n.

- Male proctiger with large posterior lobe (Fig. 8). Paramere (Fig. 9), in
profile, with broad bulge in apical third along the fore margin; apex
broadly rounded. Distal portion of aedeagus (Fig. 10) with subrectan-
gular apical dilatation. Female terminalia as in Fig. 24. Greece
PA RS EN EEE U A nee T. drosopouli sp. n.

3 Forewing (Fig. 3) membrane lacking surface spinules except for base of
cell cuy. Paramere robust, fore margin irregularly curved in apical half
(Figs 12, 13). Apex of distal portion of aedeagus with large apical hook
(Figs 14, 15). Female proctiger with relatively long and slender apical
process, subgenital plate, in profile, truncate apically (Fig. 25)....... T. galii

- Forewing (Fig. 4) membrane with surface spinules present in all cells
forming more or less extended fields. Paramere slender, evenly curved
(Fig. 17). Apex of distal portion of aedeagus with relatively short apical
hook (Fig. 18). Female proctiger with relatively short massive apical
process, subgenital plate, in profile, bearing a small point apically
(Bie ADO) ie DE Ne à © T. velutina

Fifth instar larvae
(larvae of 7. velutina unknown)

1 Body dimensions small, body length < 1.8 mm, see also table2...... T. galii
- Body dimensions large, body length > 1.8 mm, see also table 2........... 2
2 Marginal sectasetae on forewing pads over 100, and on hindwing pads
Oven om WestMediterrancant'! Mer Sea Be T. cocquempoti sp. n.
- Marginal sectasetae on forewing pads less than 100, and on hindwing
padsgessithansliS Greece ee Ve ee eee T. drosopouli sp. n.
Trioza cocquempoti sp. n. Figs) 5 7923529

Spanioza galii aspinovelutina sensu Boselli, 1930: 14, nec Sulc, 1913.
Trioza galii sensu White & Hodkinson, 1982: 43, nec Foerster, 1848.

MATERIAL EXAMINED: Holotype d, France: Gard, 5 km W Pont-St.-Esprit, 22.x.1988,
Galium sp. (D. Burckhardt) (MHNG, dry mounted).

Paratypes. Algeria: 1 ®, Atlas of Blida Chréa, Les Glacières, 1100 m, 3.v.1988
(D. Burckhardt) (MHNG, dry mounted); 1 ©, Grande Kabylie, Oued Isser, bridge N24,
21.v.1988 (D. Burckhardt) (MHNG, dry mounted). - France: 2 2, Corsica, Luri, Col de Sainte
Lucie, 24.iv.1992 (W. Della Giustina) (NHMB, dry mounted); 1 ©, Gard, same data as holotype;
1 dg, 1 ©, Var, Roquebrune-sur-Argens, Les Sauterons, 15-20.iv.1990 (C. Lienhard) (NHMB,
dry mounted); 1 &, 2 2, Indre et Loire, La Roche-Clermaut, Malaise trap, 11.viii.1989 (C.

PALAEARCTIC TRIOZIDS ON RUBIACEAE DATEN,

Fics 21-27

Trioza spp. 21, 25 - T. galii; 22, 27- T. centranthi, 23 — T. cocquempoti; 24 — T. drosopouli;
26 — T. velutina. 21, 22 — Antennal segment 10, scale a; 23-27 — female terminalia, in profile,
scale b.

Cocquempot) (MMBC, dry mounted and stored in glycerine); 1 à , Indre et Loire, Roquebrunne-
sur-Argens, 15-20.iv.1990 (C. Cocquempot) (MMBC, stored in glycerine); 2 ©, Tarn, Albi,
Fargues, 30.v.1933, Crataegus (A. Perrier) (MNHN, dry mounted); 1 9, Tarn, Albi, 7.vi.,
Crataegus (A. Perrier) (MNHN, dry mounted); 1 9, Albi, Castelnau-de-Lévi, 8.iv., Rhus (A.
Perrier) (MNHN, dry mounted); 1 2, Albi, Creyssence, 14.vi., Buxus (A. Perrier) (MNHN, dry
mounted); 1 dg, Vaucluse, La Roque-Alric, near Beaumes-de-Venise, 300 m, 23.iv.1984 (C.
Lienhard) (MHNG, slide mounted); 2 4, Vaucluse, near Vacqueyraz, river Ouvèze, 60 m,
26.iv.1984 (C. Lienhard) (MHNG, dry mounted); 6 4, 7 9, without locality data (MNHN, dry
mounted). - Italy: 1 4, 5 2, 2 parasitised larvae, Portici, 25.v.1928, Rubia (F. B. Boselli)
(DEZA, dry mounted); ca. 100 larvae and exuviae, 21.v.1928, Rubia (F. B. Boselli) (DEZA,
alcohol); 10 &, 9 ©, 45 larvae and exuviae, Portici, Parco Gussone, v-vi.2004 (P. A. Pedata)
(DEZA, alcohol; NHMB, slide mounted); 1 4, Basilicata, Potenza, PR Gallipoli Cognato, near
Monte Croccia, 810-910 m, 1-4.v.2001, Malaise trap 2 (Springate et al.) (DEZA, alcohol); 1 à,
Basilicata, Matera, PR Gravina di Matera, 15 km E of Masseria San Francesco, 16-19.111.2002,

278 D. BURCKHARDT & P. LAUTERER

290-310 m, Malaise trap (Springate et al.) (DEZA, alcohol); 2 2, same but 27-30.iv.2001, 290-
370 m; 2 d,2 ©, same but PR Gravina di Matera, 2.5 km W of road to Ginosa, 28-30.iv.2001;
1 2, same but PR Gravina di Matera, 1.5 km W of road to Ginosa, 25-28.ix.2001; 1 6,
Campania, Napoli, Parco Gussone, 55 m, 1-4.1v.2002, Malaise trap (Springate et al.) (DEZA,
MMBC, NHMB, alcohol, dry mounted); 2 4, 4 2, same but 55-80 m, 17-20.v.2001; 3 2, same
but Caserta, WWF San Silvestro, 270-300 m, 13-16.v. 2001. - Morocco: 1 ? without wings,
Tanger (MMBC, Sulc collection). - Spain: 1 6, Zaragoza Province, Monegros Region, Pina de
Ebro, 360 m, 2.vi.1990 (J. Blasco-Zumeta) (MHNG, dry mounted).

Material excluded from type series. France: 1 adult without abdomen, Vaucluse, La
Roque-Alric, 350 m, 27.iv.1984 (C. Lienhard) (MHNG, dry mounted).

Diacnosis: Adult. Total body length 2.8-3.3 mm. Colour of thorax yellow,
sometimes brown or dark brown dorsally, or with brown longitudinal stripes on meso-
praescutum and mesoscutum. Genal processes 0.6 times as long as vertex along mid-
line, subacute apically. Forewing (Fig. 1) elongate, pointed apically; bifurcation of vein
M distinctly distal to line connecting apices of veins Rs and Cu] „. Forewing membrane
lacking surface spinules except at wing base. Hindwing two thirds of forewing length.
Male proctiger (Fig. 5) relatively massive, strongly produced posteriorly. Paramere
(Fig. 6) massive, truncate apically, in profile, with narrow angular bulge in apical third
along the fore margin; apex angular. Distal portion of aedeagus (Fig. 7) with reniform
apical dilatation. Female proctiger (Fig. 23) relatively short, pointed apically; ventral
and posterior margin of female subgenital plate, in profile (in cleared specimens),
forming an angle > 90°. Measurements as in Table 1.

Fifth instar larva described by Boselli (1930) as Spanioza galii aspinovelutina.
With following numbers of marginal sectasetae (one side only): head 42-46, forewing
pad 109-113, hindwing pad 15-18, caudal plate 108-122. Tarsal arolium transversely
oval (Fig. 29). Measurements as in Table 2.

ETYMOLOGY: The species is dedicated to Christian Cocquempot, one of the col-
lectors of the new species.

DISTRIBUTION: Algeria, France, Italy, Morocco, Spain.

BIOLOGY. Host plant. Boselli (1930) collected eggs, larvae and adults exclu-
sively on Rubia peregrina. The larvae induce a strong curling of the leaves. Boselli
could not find any specimens on Galium. The holotype & and a female paratype from
France have been taken on Galium sp. but no larvae were found. At the moment there
is no evidence that Galium is a host plant.

According to Boselli (1930) larvae of 7. cocquempoti aestivate and hibernate in
the galls on the host, which also has a summer and winter dormancy. In spring and
autumn when the host produces new flush, the adults emerge and, a few days later after
copulation, the females lay their eggs. The eggs are laid perpendicularly on the leaf
margins or stems. Depending on the season the egg development takes two (spring
generation) to over four weeks (autumn generation). If the conditions in spring are
particularly favorable, three generations per year are possible.

COMMENTS: The description of Spanioza galii aspinovelutina by Boselli (1930)
concerns 7. cocquempoti, which was also confirmed by the examination of Boselli's
material. For the validity of the name aspinovelutina see comments to 7. galii. The
larval description by White & Hodkinson (1982), tentatively referred to 7. galii, is
based on Boselli's (1930) description and, hence, refers to 7. cocquempoti.

PALAEARCTIC TRIOZIDS ON RUBIACEAE 279

V\ 0.3 mm

Fics 28-31
Trioza spp. 28, 31 — 7. galii; 29 — T. cocquempoti; 30 — T. drosopouli. 28 — Last instar larva, left
dorsal face, right ventral face (scale a), with details of marginal sectasetae and tarsal apex (scale
b); 29-31 — tarsal arolium, scale b.

280 D. BURCKHARDT & P. LAUTERER

Trioza drosopouli sp. n. Figs 2, 8-10, 24, 30

MATERIAL EXAMINED: Holotype d, Greece: Attiki, Tatoi, NW Athens, 250 m, 1.v.1995,
Rubia tinctorum (P. Lauterer) (NHMB, dry mounted).

Paratypes. Greece: 16 0,43 2, 1 exuvia, Attiki, same data as holotype (MMBC, 14 6,
38 © dry mounted, 1 4,2 9, 1 exuvia, stored in glycerine; NHMB, 2 2 © dry mounted, 1 6, 1
2 stored in glycerine); 4 4, 12 2, same data but 200 m, 7.v.1995, clearing and Pinus wood
(MMBC, 3 6, 11 2 dry mounted; NHMB, 1 d, slide mounted, 1 2 dry mounted); 2 6,3 2,
Attiki, Parintha Mountains, Agfa Trias Parinthos, 1200 m, 28.v.1995, Rubia tinctorum in wood
undergrowth (P. Lauterer) (MMBC, 1 4,2 2 dry mounted; NHMB, 1 d, 1 © dry mounted); 1
?, Akarnania, Gavrolimni, 120 m, 17.11.1982 (C. Lienhard) (MHNG, dry mounted); 1 9,
Arkadia, E Peloponissos, 4 km E of Agios Petros, 900 m, 5.v.1995 (P. Lauterer) (MMBC, dry
mounted); 1 exuvia, Messinfa, Messene, between Kalamata and Pilos, 16.v.1979 (D. Burckhardt)
(MHNG, slide mounted).

Material not included in type series. Greece: 1 4, 11 2, damaged, Attiki, same data as
holotype (MMBC); 1 very teneral 9, Messinia, Messene, between Kalamata and Pilos,
16.v.1979 (D. Burckhardt) (MHNG, slide mounted).

DIAGNOSIS: Adult. Total body length 2.8-3.4 mm. Colour of thorax yellow or
ochreous; mesopraescutum dark brown or black, somtimes with a median longitudinal
lighter stripe; mesoscutum ochreous to light brown, sometimes with dark brown longi-
tudinal stripes, generally lighter than mesopraescutum. Genal processes 0.8 times as
long as vertex along mid-line, subacute apically. Forewing (Fig. 2) elongate, pointed
apically; bifurcation of vein M distinctly distal to line connecting apices of veins Rs
and Cuj,. Forewing membrane lacking surface spinules except at wing base.
Hindwing two thirds of forewing length. Male proctiger (Fig. 8) massive, strongly pro-
duced posteriorly. Paramere (Fig. 9) massive, blunt apically, in profile, with large
angular bulge in apical third along the fore margin; apex rounded. Distal portion of
aedeagus (Fig. 10) with angular apical dilatation. Female proctiger (Fig. 24) relatively
short, pointed apically; ventral and posterior margin of female subgenital plate, in
profile (in cleared specimens), forming an angle > 90°. Measurements as in Table 1.

Fifth instar larva. With following numbers of marginal sectasetae (one side
only): head 39-44, forewing pad 94-99, hindwing pad 11-12, caudal plate 97-110.
Tarsal arolium oval (Fig. 30). Measurements as in Table 2.

ETYMOLOGY: The species is dedicated to our friend Sakis Drosopoulos who
organised the collecting trip of Pavel Lauterer to Greece and assisted in the field.

DISTRIBUTION: Greece (Akarnania, Arkadia, Attiki, Messinia).

BIOLOGY: A larval skin and adults, including teneral specimens, have been
collected on Rubia tinctorum which is a likely host. A few teneral adults were collected
on 1 May and followed by mature adults up to 28 May, suggesting that the bionomics
may be similar to that of 7. cocquempoti.

COMMENTS: No reliable characters could be found for separating the females of
T. drosopouli and T. cocquempoti.

Trioza galii Foerster Figs 3, 11-1521 25281

Trioza galii Foerster, 1848: 87; Sulc, 1913: 38, p. p. Lectotype d, Germany: Nordrhein-
Westfalen, near Aachen (A. Foerster) (MMBC), here designated, examined.
Psylla galii; Hardy, 1853: 3876.

PALAEARCTIC TRIOZIDS ON RUBIACEAE 281

Trioza galii Foerster f. aspinovelutina Sulc, 1913: 44. Name unavailable, articles 45.6.1 and
45.6.4, ICZN, 1999.

Spanioza galii; Enderlein, 1926: 400; Conci, 1992: 258.

Trioza rubiae Baeva, 1972: 63. Holotype d, Turkmenistan: north-western part of Turkmen-
Khorasan Mountains, around Yaradzha, 21.iv.1971, Rubia florida (V. Baeva,
O. Rudenko) (ZISP, dry mounted), examined. Syn. n.

Trioza rubicunda Loginova, 1978: 111. Holotype ¢, Kazakhstan: Chimkent District, Aksu-
Dzhibagly Reservation, Talass Mountain Range, 80 km E Chimkent, 10.vi.1966, Galium
sp., (M. M. Loginova) (ZISP, dry mounted), examined. Syn. n.

Spanioza rubiae; Conci, 1992: 258.

Spanioza rubicunda; Conci, 1992: 258.

MATERIAL EXAMINED: Material was examined from following countries: Algeria;
Armenia; Austria: Oberösterreich, Steiermark; Bulgaria; Czech Republic, Bohemia, Moravia;
Cyprus; France: Finistère, Haute-Garonne, Hautes-Pyrénées, Hérault, Indre et Loire, Landes,
Maine-et-Loire, Nord, Pas-de-Calais, Tarn, Var; Germany: Baden-Württemberg, Berlin,
Brandenburg, Hessen (including paralectotype d, Ems, leg. C.H.G. von Heyden), Nordrhein-
Westfalen (including d lectotype and paralectotypes 6 & and 1 adult without abdomen, Aachen),
Sachsen; Greece: Atikî, Corfu, Crete, Cyclades, Evia; Israel; Italy: Alto Adige, Puglia; Jordan;
Kazakhstan (including d holotype of Trioza rubicunda); Kyrgyzstan; Malta; Slovakia; Slovenia;
Spain: Caceres, Zaragoza, Canaries; Switzerland: Aargau, Basel-Land, Genève, Luzern,
Neuchatel, Vaud, Valais; Turkey: Aydin; Tunisia; Turkmenistan (including & holotype of Trioza
rubiae); United Kingdom: England, Scotland, Wales (BMNH, MHNG, MMBC, MNHN,
NHMB, NHMV, SMTD, ZISP, ZMHB, ZSSM).

DIaGNOSIS: Adult. Total body length 2.3-3.2 mm. Colour of thorax usually dark
brown or almost black with exception of area around wing insertion which is
yellowish. Younger specimens with light colour more expanded and dark areas brown
rather than black. Genal processes 0.3-0.7 times as long as vertex along mid-line, blunt
to subacute apically. Forewing (Fig. 3) varying from relatively short and wide to elon-
gate, angular apically; bifurcation of vein M on or slightly distal to line connecting
apices of veins Rs and Cuj,. Forewing membrane lacking surface spinules except at
wing base. Hindwing three quarters or four fifth of forewing length. Male proctiger
(Fig. 11) relatively slender, weakly produced posteriorly. Paramere (Figs 12, 13)
lamellar, with an antero-basal bulge and subparallel margins in the middle third, apical
third abruptly narrowed to posteriorly directed apex. Distal portion of aedeagus (Figs
14, 15) with large hook-shaped apical dilatation. Female proctiger (Fig. 25) with rela-
tively short thick apical process; subgenital plate, in profile, bearing small point
apically. Measurements as in Table 1.

Fifth instar larva (Fig. 28). With following numbers of marginal sectasetae (one
side only): head 34-37, forewing pad 88-110, hindwing pad 11-12, caudal plate 98-101.
Tarsal arolium oval (Fig. 31). Measurements as in Table 2.

DISTRIBUTION: 7. galii is widely distributed in the West Palaearctic and Central
Asia. We have not seen any specimens from the East Palaearctic where the species
probably also occurs. It has been reported from all over the Palaearctic including Japan
and Taiwan (Aulmann, 1913; Klimaszewski, 1973; Gegechkori & Loginova, 1990;
Ossiannilsson, 1992) but some of the records may concern other species of the 7. galii
complex.

BIOLOGY: 7. galii is reported to form galls on the buds, stems and leaves of
Galium, Rubia and Sherardia spp. We have examined larvae and galls on following

D. BURCKHARDT & P. LAUTERER

282

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PALAEARCTIC TRIOZIDS ON RUBIACEAE 283

plants: Galium album, G. aparine, G. palustre and Asperula cynanchica. The species
overwinters as adult. The adults of the new generation appear in June or July. With the
available information it is impossible to deduce the number of yearly generations of 7.
galii.

COMMENTS: The type series of Trioza galii, originally deposited in the NHMV
and now in the MMBC, contains one series from Aachen and one from Ems, leg. von
Heyden. The former is homogeneous, but the latter is mixed, including 1 & of 7. galii
and 3 d of 7. velutina. As Foerster's (1848) description of 7. galii is not diagnostic, and
for stabilising the nomenclature, a lectotype is selected from the series from Aachen.
This accords with Flor's (1861a) and Low's (1882) concept of 7. galii.

Sulc (1913) admitted that Trioza galii in his broad definition, which included 7.
distincta and T. velutina as synonyms, exhibited a surprisingly large variation in size
and shape of the genal processes, in body and forewing colour, in forewing shape,
venation and spinulation as well as in genital morphology. He interpreted differences
in the male terminalia mostly as artefacts. For the other characters he suggested that
extremes are linked by intermediates. Based on forewing characters he subdivided 7.
galii into four forms for two of which he introduced the names f. spinogalii and f.
aspinovelutina, in addition to galii forma typica and f. velutina. According to para-
graph 45.6.4 of the ICZN (1999) a name introduced before 1961 as "variety" or "form"
has to be considered subspecific, unless its author also expressly gave it infrasub-
specific rank, or the content of the work unambiguously reveals that the name was
proposed for an infrasubspecific entity, in which case it is infrasubspecific. The latter
applies to the two names as Sulc mentioned that these narrowly defined forms are
linked by intermediates and, thus, implying that they are artificial groupings for
convenience rather than diagnosable entities such as subspecies or species. As infra-
subspecific names they are not available. Boselli (1930) used the name Spanioza galii
apinovelutina for Italian specimens of 7. cocquempoti, thus misinterpreting Sulc's
taxon, which represents 7. galii judging from Sulc's figures of Austrian specimens.
Boselli did not raise the name spinogalii to subspecies or species rank, and the name
does not become available through Boselli (1930).

The examination of the male holotypes of 7. rubiae and T. rubicunda showed
that they are conspecific with 7. galii, with which they are synonymised.

In Trioza chenopodii Reuter the forewing shape is variable as in 7. galii.
Lauterer (1982) demonstrated for the former, that this variation is seasonal, induced by
day length. We found no evidence in species of the 7. galii complex for seasonal
differences as in 7. chenopodii.

Trioza velutina Foerster, stat. rev. Figs 4, 16-18, 26

Trioza velutina Foerster, 1848: 87. Lectotype d, Germany: Rheinland-Pfalz, near Boppard
(A. Foerster) (MMBC), here designated, examined.

Psylla velutina; Hardy, 1853: 3876.

Trioza disticta Flor, 1861b: 401. Holotype 9, Germany: Thiiringen, Ruhla, mid June (G. Flor)
(MMBC), examined. Syn. n. à

Trioza galii sensu Sulc, 1910: 16, nec Foerster, 1848; Sulc, 1913: 45. Misidentification.

Trioza galii, Sulc, 1913: 38, p. p. |

Trioza galii Foerster f. spinogalii Sulc, 1913: 44. Name unavailable, articles 45.6.1 and 45.6.4,
ICZN, 1999.

284 D. BURCKHARDT & P. LAUTERER

MATERIAL EXAMINED: Material was examined from following countries: Algeria;
Armenia; Austria: Niederösterreich, Oberösterreich, Steiermark; Croatia; Czech Republic;
France: Ain, Haute-Garonne, Haut-Rhin, Nord, Pyrénées Orientales, Tarn; Germany: Baden-
Württemberg, Bayern, Berlin, Brandenburg, Nordrhein-Westfalen, Rheinland-Pfalz (4 lectotype
of Trioza velutina, paralectotypes 1 d,2 ©, Boppard), Sachsen, Thüringen (including 2 holo-
type of Trioza distincta); Greece; Hungary; Italy: Alto Adige, Veneto; Kazakhstan; Mongolia;
Russia: European part, Altay, Dagestan, Irkutsk District; Slovakia; Spain: Andalusia, Canaries;
Switzerland: Aargau, Bern, Basel-Land, Basel-Stadt, Graubiinden, Jura, Schaffhausen, Schwyz,
Vaud; Turkey: Kars; Turkmenistan; Ukraine; United Kingdom: England, Scotland (BMNH, MH-
NG, MMBC, MNHN, NHMB, NHMV, SMTD, ZISP, ZMHB). The following material from the
Flor collection (EAUT) was examined: 1 d, no 1090 on yellow label, 1 2, no 8670 on red la-
bel. Both specimens were identified by Flor as Trioza velutina. There is no list referring to these
particular numbers (O. Kurina, pers. comm.). Their provenience is therefore unknown.

Diacnosıs: Adult. Total body length 2.0-2.8 mm. Colour of thorax usually dark
brown or almost black with exception of area around wing insertion which is
yellowish. Younger specimens with light colour more expanded and dark areas greyish
rather than black. Genal processes 0.5-0.9 times as long as vertex along mid-line, sub-
acute apically. Forewing (Fig. 4) relatively short and broad, angular apically; bifur-
cation of vein M near or slightly distal to line connecting apices of veins Rs and Cuj,.
Surface spinules present in all cells of forewing, forming more or less expanded fields.
Hindwing three quarters of forewing length. Male proctiger (Fig. 16) relatively slender,
weakly produced posteriorly. Paramere (Fig. 17) slender, lamellar, evenly tapering to
apex, blunt apically. Distal portion of aedeagus (Fig. 18) with relatively short apical
hook. Female proctiger (Fig. 26) with relatively short, massive apical process, pointed
apically; subgenital plate, in profile, bearing small point apically. Measurements as in
Table 1.

Fifth instar larva unknown.

DISTRIBUTION: Probably widely distributed in the Palaearctic region. There are
no reliable literature records as the species has been previously confused with 7. galii.

BIOLOGY: Unknown, probably also on Galium spp. on which adults have been
collected.

COMMENTS: For the validity of the name f. spinogalii Sulc see comments on
T. galii.

Trioza velutina Foerster var. thoracica Flor, nomen dubium

Trioza velutina Foerster var. thoracica Flor, 1861a: 514. Holotype 9, France: Bouche du Rhône,
Gémenos near Marseille (G. Flor), not examined, probably lost (Sulc, 1913: 44).

COMMENTS: Flor (1861a) described Trioza velutina var. thoracica for a single ©
from Southern France. According to paragraph 45.6.4 of the ICZN (1999) the name
thoracica has to be regarded as subspecific and as such becomes available. Flor's
description is not diagnostic. Furthermore the holotype could not be traced in the
collections of the EAUT (O. Kurina, pers. comm.) and NHMV (H. Zettel, pers. comm.)
and appears to be lost (Sulc, 1913: 44). For this reason we regard thoracica as nomen
dubium.

PALAEARCTIC TRIOZIDS ON RUBIACEAE 285

ACKNOWLEDGEMENTS

For the loan or gift material we are grateful to Javier Blaso-Zumeta (Pina de
Ebro), Christian Cocquempot (Montpellier), William Della Guistina (Versailles),
Jürgen Deckert (ZMHB), Sakis Drosopoulos (Athens), Rainer Emmrich (SMTD),
David Hollis (BMNH), Izya Kerzhner (ZISP), Tanja Kothe (ZSSM), Olavi Kurina
(EAUT), David Ouvrard (MNHN), Paolo A. Pedata (DEZA), H. Zettel (NHMV). We
thank Ian D. Hodkinson (Liverpool) for his comments on the manuscripts.

REFERENCES

AULMANN, G. 1913. Psyllidarum Catalogus. W. Junk, Berlin, 92 pp.

BAEVA, V.G. 1972. New species of psyllids (Homoptera, Psylloidea) from Turkmenia. Doklady
Akademii nauk Tadzhikistoi SSR 15: 61-63. (In Russian).

BosELLI, F.B. 1930. Studii sugli psyllidi (Homoptera: Psyllidae o Chermidae) IV. Biologia e
sviluppo della Spanioza galii aspinovelutina (Sulc). Bollettino del Laboratorio di
Zoologia generale e agraria del R. Istituto Superiore agrario di Portici 23: 13-27.

BURCKHARDT, D. & COUTURIER, G. 1994. The plant-louse Leuronota calycophylli sp. n.
(Homoptera, Psylloidea), a pest on the timber species Calycophyllum spruceanum
(Rubiaceae) in Peru. Bulletin of Entomological Research 84: 307-312.

Concı, C. 1992. Spanioza tamaninii sp. n., from Trentino (NE Italy) (Homoptera Psylloidea).
Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di
Milano 132 (1991): 257-264.

DOCTERS VAN LEEUWEN, W.M. 1937. New and noteworthy zoocecidia from the Netherlands.
Marcellia 29: 73-86.

EDWARDS, J. 1896. The Hemiptera-Homoptera (Cicadina and Psyllina) of the British Isles. L.
Reeve and Co., London, 271 pp.

ENDERLEIN, G. 1926. Psyllidologica VII. Entomologische Mitteilungen 15: 397-401.

FLOR, G. 1861a. Die Rhynchoten Livlands, II. Abtheilung Sternorrhyncha Am. et S., 2. Zunft
Phytophthires Burm. Dorpat, pp. 436-546.

FLOR, G. 1861b. Zur Kenntnis der Rhynchoten. Bulletin de la Société Impériale des Naturalistes
de Moscou 34: 331-422.

FOERSTER, A. 1848. Uebersicht der Gattungen und Arten in der Familie der Psylloden.
Verhandlungen des naturhistorischen Vereins der preussischen Rheinlande 3: 65-98,

GEGECHKORI, A.M. & Locinova, M.M. 1990. The psyllids (Homoptera, Psylloidea) of the USSR
(an annotated check list). Akademiya Nauk Gruzinskoy SSR, Tbilisi, 162 pp. (In
Russian).

HARDY, J. 1853. On some excrescences, etc., on plants occasioned by mites. Proceedings of the
Berwickshire Naturalists’ Club 3 (3): 111-113. Reprinted in Zoologist 11: 3875-3877.

HOUARD, C. 1909. Les zoocécidies des plantes d’ Europe et du Bassin de la Méditerranée, volume
2. Librairie scientifique A. Hermann et Fils, Paris, pp. 573-1247.

ICZN International Comission on Zoological Nomenclature 1999. International Code of Zoolo-
gical Nomenclature, 4th edition. International Trust for Zoological Nomenclature,
London, 306 pp.

KIEFFER, J.J. 1889. Mittheilungen über Psylliden. Entomologische Nachrichten 14: 222-224.
KLIMASZEWSKI, S.M. 1973. The jumping plant lice or psyllids (Homoptera Psylloidea) of the
Palaearctic: an annotated check list. Annales zoologici, Warszawa 30: 155-286.
LAUTERER, P. 1982. New data on the occurrence, bionomics and taxonomy of some
Czechoslovakian Psylloidea (Homoptera). Acta Musei Moraviae, Scientiae naturales 67:

133-162.

LETHIERRY, L. 1874. Catalogue des Hémiptères du Département du Nord, 2nd edition.

Imprimerie L. Danel, Lille, 108 pp., 2 plates.

286 D. BURCKHARDT & P. LAUTERER

Locinova, M.M. 1978. New species of psyllid (Homoptera, Psylloidea). Trudy Zoologicheskogo
Instituta 61: 30-123. (In Russian).

Low, F. 1882. Revision der paläarktischen Psylloden in Hinsicht auf Systematik und Synonymie.
Verhandlungen der zoologisch-botanischen Gesellschaft Wien 32: 227-254.

OsHANIN, B. 1907. Verzeichnis der palaearktischen Hemipteren mit besonderer Berücksich-
tigung ihrer Verteilung im Russischen Reiche, volume 2 Homoptera, part 2. Druckerei
der Kaiserlichen Akademie der Wissenschaften, St. Petersburg, pp. 193-384.

OSSIANNILSSON, F. 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna
Entomologica Scandinavica 26, 346 pp.

SCOTT, J. 1876. Monograph of the British species belonging to the Hemiptera-Homoptera, family
Psyllidae; together with the description of a genus which may be expected to occur in
Britain. Transactions of the entomological Society 4: 525-569, plates 8-9.

SULC, K. 1910. Monographia generis Trioza Foerster, pars I. Sitzungsberichte der königlich-böh-
mischen Gesellschaft für Wissenschaften Il Klasse 1-34, plates 1-10.

Suc, K. 1913. Monographia generis Trioza Foerster, pars IV. Sitzungsberichte der königlich-
böhmischen Gesellschaft für Wissenschaften Il Klasse 1-48, plates 34-48.

TUTHILL, L.D. 1943. The psyllids of America north of Mexico (Psyllidae: Homoptera). Jowa
State College Journal of Science 17: 443-660.

WAGNER, W. & FRANZ, H. 1961. Uberfamilie Sternorrhyncha (Psylloidea). In: FRANZ, H. (ed.).
Die Nordostalpen im Spiegel ihrer Landtierwelt, vol. 2. Innsbruck, pp. 158-179.
WHITE, IM. & Hopkinson, I.D. 1982. Psylloidea (Nymphal Stages) Hemiptera, Homoptera.

Handbooks for the Identification of British Insects 2(5b), 50 pp.

REVUE SUISSE DE ZOOLOGIE 113 (2): 287-290; juin 2006

A new species of Diathrausta Lederer, 1863 from Africa
(Lepidoptera, Pyraloidea, Crambidae, Spilomelinae)

Koen MAES

Present address: Royal Museum for Central Africa, Leuvensesteenweg 13,
B-1380 Tervuren, Belgium.

Address: AgroBioSys Intl., Kl.Smetledestraat 192, B-9230 Wetteren, Belgium.
Email: kvmaes@belgacom.net

A new species of Diathrausta Lederer, 1863 from Africa (Lepidoptera,
Pyraloidea, Crambidae, Spilomelinae). - Diathrausta semilunalis sp. n. is
described from Southern Africa. The adult, genitalia, and tympanal organs
are illustrated. Its placement in Diathrausta is discussed. Cangetta fulviceps
(Hampson, 1917) comb. n. is excluded from the genus Diathrausta.

Keywords: Diathrausta semilunalis sp. n., Africa

INTRODUCTION

Lederer (1863) erected the genus Diathrausta for his new species Diathrausta
profundalis from “Amboina” [Indonesia].

Nine species were later added by Hampson (1903, 1917), Dyar (1913) and
Druce (1899) mostly from the Old and New World tropics.

Munroe (1956) reviewed the North American species of Diathrausta, reco-
gnizing two species and three subspecies. He is the first to illustrate the male genitalia
of some species in this genus. In total there are now 18 species in this genus (personal
database).

Only one species, Diatrausta fulviceps Hampson, 1917 (Malawi), has been
described from Africa. Presumably, it was placed in Diathrausta based on external
similarities. I dissected specimens of D. fulviceps from Kenya, Tanzania and the
Democratic Republic of Congo and compared the genitalia with those of Diathrausta
reconditalis (Walker, 1859) and D. harlequinalis amaura Munroe, 1956 as illustrated
in Munroe (l.c.). The male genitalia of D. fulviceps have a simple uncus broadly fused
with the tegumen and dorsally with some simple setae, this is in sharp contrast with the
narrow slender uncus of true Diathrausta species. The valva of D. fulviceps are broad
at their base with the tegumen and narrowing near the apex, not ear-shaped as in true
Diathrausta species. D. fulviceps has also a large fibula in the median part of the valva.
The saccus is very large in D. fulviceps and almost absent in the true Diathrausta
species. D. fulviceps does not belong in Diathrausta and is tentatively placed in the
genus Cangetta because of the resemblance in structure of the fibula in the male geni-
talia. It differs with true Cangetta species in the structure of the uncus, which is simple

Manuscript accepted 27.07.2005

288 K. MAES

in Cangetta fulviceps comb. n., but strongly modified, bilobed and carrying long setae
in most true Cangetta species.

During the curation of collections in various museums a striking undescribed
species from southern Africa came to my attention. The male genitalia have the same
basic structure (simple slender uncus, ear-shaped valva, fibula near base of valva) as
found in Diathrausta species as illustrated by Munroe (1956). This new species is
therefore placed in the genus Diathrausta. Pictures of the male and female genitalia are
given, this is the first time the female genitalia of this genus is illustrated.

Abbreviations used:

ABSRC AgroBioSys Reference Collection, Wetteren, Belgium.
MHNG Muséum d’histoire naturelle, Genève, Switzerland.
RMCA Royal Museum for Central Africa, Tervuren, Belgium.
TMP _ Transvaal Museum, Pretoria, South Africa.

Diathrausta semilunalis sp. n. Pl. 1, 2 A-D

TYPE MATERIAL: Holotype d: [SOUTH AFRICA] Karkloof, Natal, 15.Jan.[19]17
A.J.T.Janse, K.Maes Gen.prep.nr.d 20719 (TMP); 10 Paratypes: 19: [SOUTH AFRICA]
Marieps Mnt., Dec.1925, G.van Son, K.Maes Gen.prep.nr. 220738 (TMP); 19: [SOUTH
AFRICA] Mahuba’s KIf., 1050m, 14.1.[19]25, A.J.T.Janse (abdomen lost) (TMP); 1 à : [SOUTH
AFRICA] Mahuba’s KIf., 1050m, 16.1.[19]25, A.J.T.Janse (ABSRC); 19: [SOUTH AFRICA]
Barberton, 25.1.1911, A.J.T.Janse (TMP); 19: [SOUTH AFRICA] Karkloof, Natal , 21.1.[19]17
(ABSRC); 19: [SOUTH AFRICA] Rietvlei,1.4.[19]18, Coll.Janse (TMP); 16: [SOUTH
AFRICA] Sabie, Tvl., 30.iv.-5.v.1977, J.H.Potgieter,c.s. (TMP); 16: [DEMOCRATIC
REPUBLIC OF CONGO] Elisabethville, iv-v.1952, Ch.Seydel (RMCA); 1 2: [DEMOCRATIC
REPUBLIC OF CONGO] Elisabethville, xii.1936, Ch.Seydel, K.Maes Gen.prep.nr. 9 14305
(RMCA); 14: [DEMOCRATIC REPUBLIC OF CONGO] Ht.Katanga, Midingi, 1.vii.[19]30,
J.Romieux (MHNG).

DISTRIBUTION: South Africa, Democratic Republic of the Congo.

ETYMOLOGY: The name refers to the half-moon shaped white spot between the
upper angle of the cell in the fore wing and the costa.

DIAGNOSIS: Fore wings triangular with a half-moon shaped white or cream-
white spot between the upper angle of the cell and the costa.

DESCRIPTION:

Head: frons rounded; labial palps porrect, length about twice diameter of
compound eye, basal part of first segment white or creamy-white, otherwise brown;
maxillary palps long, brown.

Thorax and abdomen: dark brown like ground colour of fore wings; legs brown,
spurs 0,2,4; abdomen dark brown.

Wings: fore wings triangular; ante- and post-medial lines black, almost conti-
nuous for entire length, orbicular and reniform stigma dark; costa black near wing ba-
se; half-moon shaped white or creamy-white spot near upper angle of cell, on outer si-
de delimited by black postmedial line; apex brown as rest of wing, fringe on fore and
hind wings with black and white or creamy-white parts; hind wings lighter brown or
black as fore wings with narrow white or creamy-white postmedial line; wing pattern

A NEW SPECIES OF DIATHRAUSTA 289

PLATE 1
Adult moth: Diathrausta semilunalis sp.n. Holotype (black bar = 1 cm)

PLATE 2

Genitalia and tympanal organs Diathrausta semilunalis sp.n. A: male genitalia. B: aedeagus of
male genitalia. C: female genitalia. D: tympanal organs.

290 K. MAES

more or less evident or largely reduced in dark specimens; frenulum simple in males,
double in females; retinaculum a series of elongated scales near Cu-stem of fore wing;
wingspan: 13-16mm.

Tympanal organs (PI. 2 D): partly invaginated with broad fornix tympani.

Male genitalia (PI. 2 AB): uncus a single large lobe, dorsally covered with short
setae; tegumen almost rectangular, vinculum U-shaped, saccus very small; valva with
strong costa, more membranous near apex and with a few short setae; spatula-shaped
fibula medially at base of valva; juxta shield-shaped; tubular aedeagus with a single
spine-shaped cornutus.

Female genitalia (PI. 2 C): papillae anales with short and long setae; apophyses
posteriores and anteriores of about same length; sinus vaginalis invaginated at VIIth
sternite, forming circular membranous area; calyx-shaped, well-sclerotized ostium
bursae; ductus bursae membranous and rather short; corpus bursae bulbous with plate-
shaped signum carrying straight transversal ridge; appendix bursae lacking.

LIFE CYCLE: unknown.

ACKNOWLEDGEMENTS

I wish to thank Dr. M. Kruger and Mrs. B. Dombrowsky from the Transvaal
Museum in Pretoria, Dr. Bernard Landry from the Muséum d’histoire naturelle in
Geneva, and Dr. Marc De Meyer and Dr. Ugo Dall’ Asta from the Royal Museum for
Central Africa for putting material of their collections at my disposal.

This paper results from research finalized during a project funded by DWTC
(Federale Diensten voor Wetenschappelijke, Technische en Culturele Aangelegen-
heden), Belgium.

LITERATURE

DRUCE, H. 1899. Biologia Centrali-Americana Insecta. Lepidoptera-Heterocera vol.ii :537-569.

Dyar, H. G. 1913. A note on Diathrausta nerinalis Walker (Lepidoptera, Pyralidae). Insecutor
Inscitiae Menstruus 1: 100-102.

HAMPSON, G. F. 1903. The moths of India. Supplementary Papers to the Volumes in " The Fauna
of Bristish India." Series ii. Part x. Journal of the Bombay Natural History Society xv(2):
206-226, 1 pl.

Hampson, G. F 1917. Descriptions of New Pyralidae of the Subfamilies Hydrocampinae,
Scoparianae &. The Annals and Magazine of Natural History (8)20: 201-216.

LEDERER, J. 1863. Beitrag zur Kenntniss der Pyralidinen. Wiener Entomologische Monatschrift
7(8): 243-280; (10): 331-378; (11): 379-506, 18pl.

MUNROE, E. 1956. The North American species of Diathrausta Lederer (Lepidoptera: Pyralidae).
The Canadian Entomologist 88: 579-583, 7 figs.

REVUE SUISSE DE ZOOLOGIE 113 (2): 291-296; juin 2006

Redescription of Ceratophysella lawrencei (Gisin, 1963)
(Collembola: Hypogastruridae)

Dariusz SKARZYNSKI
Zoological Institute, Wroctaw University, Przybyszewskiego 63/77, 51-148 Wroctaw,
Poland. E-mail: hypogast@biol.uni.wroc.pl

Redescription of Ceratophysella lawrencei (Gisin, 1963) (Collembola:
Hypogastruridae). - Ceratophysella lawrencei (Gisin, 1963) is redescribed
based on material from Switzerland (types), Austria, Italy and Poland. A
lectotype is designated. Notes on morphology of the closely related species
Ceratophysella neomeridionalis (Nosek & Cervek, 1970) are given.

Keywords: Collembola - Hypogastruridae - Ceratophysella lawrencei -
Ceratophysella neomeridionalis - taxonomy - Europe.

INTRODUCTION

During a faunistic survey in the Tatra National Park (S Poland) numerous
Ceratophysella specimens with 2 + 2 spine-like setae on the head were found. Iden-
tification led to C. lawrencei with some doubts. This species is insufficiently known
and available literature does not allow to recognize it unambiguously (Thibaud et al.,
2004). Therefore, a redescription is presented here.

Gisin (1949) briefly described Hypogastrura gibbosa (Bagnall, 1940) with 2 + 2
spine-like setae on the head from the Swiss Alps. Later in “Collembolenfauna Europas”
(Gisin, 1960) he repeated the short species diagnosis and noticed that the identity of
British and Swiss specimens of H. gibbosa is doubtful. His suspicion turned out to be
true when Lawrence (1962) redescribed this species based on Bagnall’s types from
Great Britain. He stated that H. gibbosa sensu Gisin 1949, 1960 with cephalic spines
refers to a different species. As a consequence Gisin (1963) established a new species,
H. lawrencei by giving a reference to previous descriptions (Gisin, 1949, 1960).

Examination of Gisin’s material allowed to redescribe this species using a set of
modern diagnostic characters (Fjellberg, 1984, 1998/1999; Babenko er al., 1994;
Skarzyfiski, 2004). Moreover, a lectotype was designated.

Ceratophysella lawrencei (Gisin, 1963)
Hypogastrura lawrencei Gisin, 1963: 97.
Hypogastrura gibbosa (Bagnall, 1940): Gisin 1949, 1960 nec Bagnall, 1940.

TYPE MATERIAL: Lectotype - male on slide (formerly in alcohol), by present designation;
He 242a: Grisons, Swiss National Park, Plan Possa (above II Fuorn), S slope, Pineto-Caricetum
humilis, 1900 m a. s. 1., 13. V. 1945, soil sample (1-4 cm) (in the collection of the Muséum d’his-
toire naturelle in Geneva), paralectotypes - 5 specimens of unidentified sex on slides (one of
them formerly in alcohol), same data as lectotype.

Manuscript accepted 22.02.2005

292 D. SKARZYNSKI

OTHER MATERIAL EXAMINED: 3 specimens of unidentified sex on slide; He 422: Grisons,
Swiss National Park, II Fuorn, S slope, on the way to Plan Posa, Mugeto-Ericetum caricetosum
humilis, 1850 m a. s. 1., on mushrooms, 22-24.VIII. 1950. 2 specimens of unidentified sex on
slide; Törne, [illegible inscription - probably one of collecting sites of A. cf. gibbosa in Austria
mentioned in Törne (1958)], 27.1.1951 (the material is preserved in the collection of the
Muséum d’histoire naturelle in Geneva). 1 female, 3 juveniles on slide; moss on calcareous rocks
(ca. 2000 m a. s. 1.), neighborhood of Cortina d’ Ampezzo (Dolomites, Alps, N Italy), 20.VI-
II.1993, leg. D. Skarzyfiski. 12 males, 10 females, 8 juveniles on slides; moss and mountain
grasslands with mushrooms on calcareous rocks: deep ravine Kraköw (ca. 1100 m a. s. D),
entrance to the cave Mylna (ca. 1100 m a. s. 1.), N slope of the Gtadkie Uptaziafiskie (ca. 1600
m a.s.l.), W slope of the Chuda Turnia (ca. 1800 m a. s. 1.) (Tatra Mountains, Carpathians, S
Poland), 18.VI.2004, 19.VII.2004, leg. D. Skarzyñski (the material is preserved at the
Department of Biodiversity and Evolutionary Taxonomy, Wroctaw University, Poland).

DIAGNOsIs: This species can be recognized by the following set of characters:
fields of coarse granulation on head, thoracic and abdominal terga present, dorsal
chaetotaxy of the C. armata-type, setae long, thick and serrated, setae d, and oc, on
head spine-like, setae p; on abdominal tergum IV present, antennal segment IV with
ca. 25 modified ventral sensilla, eversible sac between antennal segments III-IV
present, head of maxilla and labium of the C. armata-type, outer lobe with 1 sublobal
hair, empodial appendage with long apical filament reaching distinctly beyond inner
tooth of claws, dens with uniform, fine granulation and 7 setae (2-4 inner setae
modified), mucro boat-like with large lateral lamella, anal spines large, curved, situated
on high basal papillae.

REDESCRIPTION: Body length 0.9-1.6 mm (lectotype, male 0.95 mm). Colour of
the body dark greyish-violet (in freshly collected specimens from the Tatra Mts,
greyish-brown in alcoholic lectotype and paralectotype), legs and ventral part of the
body paler, eyes fields black, anal spines yellow.

Integument with fine granules and fields of coarse granulation on head (large
uniform field covering almost whole dorsal side excluding bases of antennae), thoracic
terga II-III (two subaxial fields of medium size), abdominal terga I-III (small fields
around macrochaetae p>), abdominal tergum IV (one axial field of medium size and
two lateral large ones), abdominal terga V-VI (large fields covering almost whole
surface) (Figs 1-3). Fields made of rather irregularly distributed granules, which are
broad, moderately high and sometimes bent in different directions (Figs 1-3). On
abdominal tergum V coarse granules are usually loosely distributed from the line
extended between setae a, to the line extended between setae p,, behind this line
granules are slightly smaller and densely packed. 8-10 granules between setae pj
(Fig. 3).

Dorsal chaetotaxy as in Figs 1-3. Microchaetae and macrochaetae (especially on
last abdominal terga) long, thick and serrated (Figs 1-3). Body sensilla (s) short, thin
and smooth. Setae d, and oc, on head spine-like in unreproductive adults (Fig. 1), in
juveniles like normal or thickened macrochaetae, rarely spine-like. Setae a, on thoracic
tergum II like mesochaetae or macrochaetae. Setae pj and pz on abdominal tergum IV
macrochaetae and microchaetae respectively, setae p; present. Subcoxae I, II, III with
1, 2, 3 setae respectively. Microsensillum (ms) on thoracic tergum II present.

Antennal segment IV with simple apical vesicle, subapical organite, micro-
sensillum, 7 cylindrical sensilla in typical arrangement and ca. 25 short, thick, curved

REDESCRIPTION OF CERATOPHYSELLA LAWRENCEI 293

and flattened at tips sensilla in ventral file (Fig. 5). Antennal III-organ with two long
(lateral) and two short (internal) curved sensilla. Microsensillum on antennal segment
III present. Eversible sac between antennal segments III-IV present (Fig. 5). Antennal
segment I with 7 setae.

Ocelli 8+8, G and H slightly smaller than others (Fig. 1). Postantennal organ
about twice as large as single ocellus, with 4 lobes of which the anterior pair is dis-
tinctly larger than the posterior (Fig. 4). Large accessory boss partly surrounded by
posterior lobes of postantennal organ (Fig. 4).

Labrum with 5, 5, 4 setae and 4 prelabrals. Head of maxilla of the C. armata-
type (see Fjellberg, 1984). Outer lobe with 1 sublobal hair. Labium of the C. armata-
type (see Fjellberg, 1998/1999).

Tibiotarsi I, II, II with 19, 19, 18 setae respectively, clavate setae absent. Claws
with inner tooth and pair of lateral teeth (Fig. 6). Empodial appendage with broad basal
lamella and long apical filament reaching beyond inner tooth of claws (Fig. 6), ratio:
empodial filament/ inner edge of claws = 0.7-1 (0.5-0.7 in the Tatra Mts population).

Ventral tube with 4+4 setae.

Furca fully developed. Dens with uniform, fine granulation and 7 setae (2-4
inner setae modified) (Fig. 8). Mucro boat-like, with large lateral lamella (Figs 7, 8).
Ratio: dens/mucro = ca. 2. Retinaculum with 4+4 teeth.

Anal spines large, curved, situated on high basal papillae (Fig. 3). Ratio: anal
spines/inner edge of claws III = ca. 1.5.

DISCUSSION: C. lawrencei resembles these species of the C. armata-group which
have integument with fields of coarse granulation. It distinctly differs from C. granu-
lata Stach, 1949 and C. scotica (Carpenter & Evans, 1899) in having spine-like setae
on head (v. absent) and from C. silvatica (Rusek, 1964) in their number (C. lawrence:
2 + 2, v. 1 + 1). This species is also easy to distinguish from C. franzi (Butschek &
Gisin, 1949) by different arrangement of cephalic spine-like setae (C. lawrencei: d,
and oc», v. d, and sds).

Undoubtedly C. lawrencei is closely related to C. neomeridionalis (Nosek &
Cervek, 1970) — a species described from the Slovenian Dinaric Mountains (Nosek &
Cervek, 1967, 1970) and also known from the Polish Carpathians (Weiner, 1981,
Smolis er al., 2001, Skarzyñski & Smolis, 2002, Smolis & Skarzyfiski, 2003). They
differ in the structure of empodial appendage (C. lawrencei: basal lamella broad, apical
filament long reaching beyond inner tooth of claws, v. basal lamella small, apical
filament short reaching distinctly below inner tooth of claws) and arrangement of setae
on abdominal tergum IV (C. lawrencei: setae p3 present, v. setae p3 absent).

Nosek & Cervek (1967) considered chaetotaxy of the last abdominal terga of C.
neomeridionalis characteristic for this species. However this opinion seems to be a
result of erroneous identification of some setae. It is evident after examination of
paratype from the collection of the Muséum d’histoire naturelle in Geneva (Nanos,
7.X1.64, leg. S. Cervek) and specimens collected in the Polish Carpathians that they
have body sensilla on abdominal tergum IV in typical position ps (Fig. 9) (p7 in
original description, see Nosek & Cervek, 1967: 247, Fig. 8), setae p, on abdominal
tergum V as strong macrochaetae, setae p, and py as short microchaetae and long body

294 D. SKARZYNSKI

0.01 mm \ \

CH
Ca
CS SN

Fics 1-9

Ceratophysella lawrencei (1-8): 1, chaetotaxy and granulation of head; 2, chaetotaxy and distri-
bution of coarse granulation fields on thoracic tergum II; 3, chaetotaxy and distribution of coarse
granulation fields on abdominal terga III-VI; 4, postantennal organ, accessory boss and neigh-
bour ocelli; 5, ventral side of antennal segments III-IV; 6, claw of legs I; 7, mucro; 8, dens and
mucro, lateral view. (1-4, 6: lectotype; 5, 7, 8: specimen from Tatra Mts). Ceratophysella
neomeridionalis: 9, chaetotaxy and distribution of coarse granulation fields on abdominal terga
IV-V (paratype). Abbreviations in text.

REDESCRIPTION OF CERATOPHYSELLA LAWRENCEI 295

sensilla (Fig. 9) (in original description p, is short microchaeta, p, is long microchaeta,
sensillum is short and py is macrochaeta, see Nosek & Cervek, 1967: 247, Fig. 8).

DISTRIBUTION: C. lawrencei was recorded from the Swiss, Austrian and Italian
Alps, the Appenines (Gisin, 1949, 1960, 1963; Törne, 1958; Christian, 1987; Dallai er
al., 1995) and for the first time from the Polish Tatra Mts (Carpathians).

ECOLOGICAL REMARKS: C. lawrencei seems to prefer different microhabitats
(soil, forest litter, moss tufts, mountain grasslands, mushrooms) associated with cal-
careous rocks in the mountains at an altitude 900-2000 m a. s. 1.

ACKNOWLEDGEMENTS

I wish to express my sincere thanks to Dr Charles Lienhard from the Muséum
d’histoire naturelle in Geneva for supplying Gisin’s material of C. lawrencei and a
paratype of C. neomeridionalis. I am also grateful to the management of the Tatra
National Park for their help with my investigations. The study was sponsored by the
Wroctaw University (grant 2020/W/IZ/2004).

REFERENCES

BABENKO, A. B., CHERNOVA, N. M., POTAPOV, M. B., STEBAEVA, S. K. 1994. Collembola of
Russia and adjacent countries: Family Hypogastruridae. Nauka, Moscow, 336 pp.

CHRISTIAN, E. 1987. Catalogus Faunae Austriae. XIa, U. Kl.: Collembola (Springschwänze).
Verlag der Osterreichischen Akademie der Wissenschaften, Wien, 80 pp.

DALLAI, R., MALATESTA, E., RAMELLINI, P. 1995. Apterygota: Collembola, Protura, Micro-
coryphia e Zygentoma (=Thysanura s.1.), Diplura. Jn: MINELLI, A., RUFFO, S., LA POSTA,
S. (eds). Checklist delle specie della fauna italiana, 33. Calderini, Bologna, 25 pp.

FIELLBERG, A. 1984. Maxillary structures in Hypogastruridae (Collembola). Annales de la
Société Royale Zoologique de Belgique 114: 89-99.

FIELLBERG, A. 1998/1999. The Labial Palp in Collembola. Zoologischer Anzeiger 237: 309-330.

GISIN, H. 1949. Notes sur les Collemboles avec description de quatorze espèces et d’un genre
nouveaux. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 22:
385-400.

GISIN, H. 1960. Collembolenfauna Europas. Muséum d’histoire naturelle, Genève, 312 pp.

GIsIN, H. 1963. Collemboles d’Europe V. Revue suisse de Zoologie 70: 77-101.

LAWRENCE, P. N. 1962. A review of Bagnall’s Hypogastrura types (Collembola). Entomologist's
Gazette13: 132-151.

NOSEK, J. & CERVEK, S. 1967. A new species of springtail from the mediterranean region
Hypogastrura (Ceratophysella) meridionalis sp. n. (Collembola: Hypogastruridae). Acta
Societatis Zoologicae Bohemoslovenicae 31: 245-249.

NOSEK, J. & CERVEK, S. 1970. Beseitigung eines Homonyms in der Gattung Hypogastrura
(Insecta: Collembola). Revue suisse de Zoologie 77: 207.

SKARZYNSKI, D. 2004. Taxonomic position of Ceratophysella granulata STACH, 1949 and
Ceratophysella silvatica (RUSEK, 1964) in the light of morphological and laboratory
hybridisation studies. Acta Zoologica Cracoviensia 47: 147-153.

SKARZYNSKI, D. & SMOLIS, A. 2002. Notes on Willemia virae Kaprus’ and Anurida carpatica
Babenko, two springtails new for the Polish fauna, with remarks on other Collembola
collected in caves of the Beskid Niski Mountains (Polish Carpathians). Polskie Pismo
Entomologiczne 71: 301-306.

SMOLIS, A., SKARZYNSKI, D. & POMORSKI, R. J. 2001. Nowe i rzadkie w faunie Polski gatunki
skoczogonkéw (Collembola). Przeglad Zoologiczny 45: 227-230 [in Polish].

296 D. SKARZYNSKI

SMOLIS, A. & SKARZYNSKI, D. 2003. Springtails (Collembola) of the “Przetom Jasiotki” reserve
in the Beskid Niski Mountains (Polish Carpathians). Fragmenta Faunistica 46: 121-129.

THIBAUD, J.-M., SCHULZ, H.-J. & GAMA, M. M. da. 2004. Synopses on Palearctic Collembola.
Hypogastruridae. Vol. 4, Abhandlungen und Berichte des Naturkundemuseums Görlitz
75(2): 1-287.

TÖRNE, E. von 1958. Faunistische Befunde einer Untersuchung des Collembolenbesatzes im
Excursionsgebiet von Innsbruck. Acta Zoologica Cracoviensia 2: 637-680.

WEINER, W. M. 1981. Collembola of the Pieniny National Park in Poland. Acta Zoologica
Cracoviensia 25: 417-500.

REVUE SUISSE DE ZOOLOGIE 113 (2): 297-303; juin 2006

Description of Ceratophysella robustiseta sp. n. from greenhouses in
England, with notes on synonymy of C. postantennalis Yosii, 1966
and taxonomic status of C. morula Deharveng & Bourgeois, 1991
(Collembola: Hypogastruridae)

Dariusz SKARZYNSKI! & Adrian SMOLIS!
! Zoological Institute, Wroctaw University, Przybyszewskiego 63/77, 51-148 Wroctaw,
Poland. E-mail: hypogast @biol.uni.wroc.pl, adek@biol.uni.wroc.pl

Description of Ceratophysella robustiseta sp. n. from greenhouses in
England, with notes on synonymy of C. postantennalis Yosii, 1966 and
taxonomic status of C. morula Deharveng & Bourgeois, 1991
(Collembola: Hypogastruridae). - Ceratophysella robustiseta sp. n. from
greenhouses in Kew Gardens (London, England) is described. Notes on
morphology and taxonomic status of a related species Ceratophysella
morula Deharveng & Bourgeois, 1991 are given. Ceratophysella postan-
tennalis Yosii, 1966 is synonymized with Hypogastrura nepalica Yosii,
1966, syn. n.

Keywords: Collembola - Hypogastruridae - Ceratophysella - taxonomy.

INTRODUCTION

The large genus Ceratophysella Borner, 1932 comprises three species with a
moruliform postantennal organ: C. mosquensis (Becker, 1905) from Europe, C. postan-
tennalis Yosii, 1966 from Nepal, and C. morula Deharveng & Bourgeois, 1991 from
Thailand. The first species mentioned is well known and widely distributed (Babenko
et al., 1994) while the two latter ones are known from the type locality (C. morula) or
some localities only (C. postantennalis). During visit in Kew Gardens greenhouses in
London several specimens which generally resemble C. postantennalis and C. morula
were found. A comparison with the type material of these species made it possible to
ascertain that the English specimens represented a new species, C. morula requires
taxonomic comments and C. postantennalis is synonymous with Hypogastrura nepal-
ica Yosii, 1966.

RESULTS AND DISCUSSION

Hypogastrura nepalica Yosii, 1966
Ceratophysella postantennalis Yosii, 1966 syn. n.

MATERIAL EXAMINED: Holotype - specimen in alcohol, Maedane Karka, East Nepal,
10.VI. 1963, leg. Yasuda; paratypes - 6 specimens on slides (formerly in alcohol), same data as
holotype. All material deposited in the Muséum d’histoire naturelle in Geneva.

Manuscript accepted 14.11.2005

298 D. SKARZYNSKI & A. SMOLIS

Holotype of Ceratophysella postantennalis - male on slide (formerly in alcohol), Nepal,
Maedane Karka, 10-12.VI.1963, leg. Yasuda, det. Yosii; paratypes - 9 specimens on slides (for-
merly in alcohol), Nepal, Maedane Karka, 10.VI.1963, m-2, leg. Yasuda, det. Yosii 1983; 2 spec-
imens on slides (formerly in alcohol), Nepal, Maedane Karka, 12.VI.1963, m-6, leg. Yasuda, det.
Yosii 1983. All material deposited in the Muséum d’histoire naturelle in Geneva.

REMARKS

C. postantennalis was described from Maedane Karka (the Rolwaling Himal,
East Nepal) on the basis of specimens with distinct morphological features: moruli-
form postantennal organ, short setae, small mucro without lamellae and the presence
of a median hump on abdominal tergum V (Yosii, 1966). Bourgeois & Cassagnau
(1972) stated an opinion that the taxonomic status of C. postantennalis should be
revised considering the low diagnostic value (possible epitokous and ecomorphic
changes) of the discriminant features used by Yosii.

Examination of the types of C. postantennalis showed that the holotype and two
paratypes were conspecific with Hypogastrura nepalica Yosii, 1966. This synonymy
was confirmed by examination of the H. nepalica holotype and paratypes. It was
interesting that all the studied H. nepalica specimens had a median hump on abdominal
tergum V similar to this of C. postantennalis in fig. 2A of Yosii (1966). Such a struc-
ture was not mentioned in the original description of H. nepalica (see Yosii, 1966).
Among other paratypes of C. postantennalis one adult in bad condition (i.e. without
head), 6 juveniles (probably of the first instar) of an unidentified Ceratophysella (den-
ticulata-group) species and 2 unidentified juveniles of the family Hypogastruridae
were found. Some of the Ceratophysella juveniles had indistinctly folded postantennal
organ, long setae, boat-like mucro and median hump on abdominal tergum V absent.

All the data mentioned suggest that Yosii made the C. postantennalis des-
cription on the basis of H. nepalica and the unidentified Ceratophysella specimens.
Since the holotype of C. postantennalis is H. nepalica and the taxonomic status of this
last species is sufficiently clear, C. postantennalis is synonymized here with A.
nepalica.

Ceratophysella morula Deharveng & Bourgeois, 1991

Ceratophysella morula Deharveng & Bourgeois, 1991: 308

MATERIAL EXAMINED: Holotype - juvenile male on slide, Thailand, Chiang Mai province,
Doi Chiang Dao, 1500 m a. s. 1., humus at entrance of a shaft, 21.XII.1980, sample THA31, leg.
Deharveng (collection of the Muséum National d’Histoire Naturelle in Paris).

REMARKS

The status of C. morula was questioned by Babenko et al. (1994). These authors
emphasized the low diagnostic value of the characters which distinguish C. morula
from the related species C. mosquensis: the number of finger-like papillae in the
postantennal organ, the ratio dens/mucro and the presence/absence of setae a’, on the
abdominal tergum V. They also suggested that the original description was inaccurate
since 1 + 1 v setae on the head and 2 sublobal hairs on the maxillary outer lobe are
unique features in the denticulata-group. Examination of the holotype of C. morula
confirmed some critical comments of Babenko er al. (1994), in fact 2 + 2 v setae and
only 1 sublobal hair were visible. Moreover, some other differences between fig. 11 of

DESCRIPTION OF CERATOPHYSELLA ROBUSTISETA SP. N. 299

Deharveng & Bourgeois (1991) and the real chaetotaxy were noted, for instance the
presence of setae p; on thoracic tergum II and different lengths of numerous body setae
and sensilla. Despite all this, C. morula seems to be a good species. From among three
subtle characters used by Deharveng & Bourgeois (1991) to distinguish C. morula
from C. mosquensis, the presence/absence of setae a’, on abdominal tergum V seems
to be reliable. This hypothesis is supported by the results of the study on the taxonomic
status of C. denticulata (Bagnall, 1941) and C. engadinensis (Gisin, 1949) (Skarzyfiski,
2004). In order to establish the diagnostic value of the remaining characters a study on
extensive topotypic material of this species is neccessary. The type material of C.
morula - two juvenile males (holotype - 0.85 mm and 4 + 5 setae on the genital plate)
is not suitable for this purpose. Taking this note into consideration one can cautiously
expect that the shape of ventral sensilla on antennal segment IV could be a useful
feature in diagnostics of the mentioned species (see Tab. 1). C. morula is also similar
to C. robustiseta sp. n. from which it differs clearly in numerous characters presented
in Tab. 1.

It is possible that the two populations from Mingbo and Yaral (the Khumbu
Himal, East Nepal) recognized as C. postantennalis by Yosii (1971) refer in reality to
C. morula. Examination of respective material from the collection of the Muséum
d’histoire naturelle in Geneva (3 adult specimens on slide made by Yosii, Nepal,
Mingbo, 299, 3.VI.1961, leg. Janetschek; 12 adult specimens on slides formerly in
alcohol, Nepal, Mingbo, 4800 m., 299, 28.V.- 3.VI.1961, leg. Janetschek, det. Yosii; 2
adult specimens on slides formerly in alcohol, Nepal, Yaral, 4100 m., 262, IV.-V.1961,
leg. Janetschek, det. Yosii) showed that these specimens generally fit the description of
C. morula. Nevertheless they clearly differ in the length of setae, C. morula has shorter
ones. In spite of this it would be premature to regard these Nepalic populations as a
new species until more information on the morphological variability of C. morula is
available.

Ceratophysella robustiseta sp. n. Figs 1-8

TYPE MATERIAL: Holotype - female on slide, surface of wet mosses near small
waterfalls in greenhouses with tropical plants in Kew Gardens (the Evolution House
and Princess of Wales Conservatory, London, England), 16. VI. 2003, leg. A. Smolis;
paratypes - 12 females, 4 males and 2 juveniles on slides, same data as holotype (type
material preserved in the collection of the Department of Biodiversity and
Evolutionary Taxonomy, Wroctaw University, Poland; one paratype female housed in
the collection of the Muséum d’histoire naturelle in Geneva).

ETYMOLOGY: The new species is named after the robust setae on the dorsal side
of the body.

DrAGNosis: This species is distinguished from other members of the denticula-
ta-group by the following combination of characters: eversible sac between antennal
segments III-IV absent, antennal segment IV with 10-15 only slightly modified sensilla
in ventral file, postantennal organ from almost smooth to weakly moruliform, chaeto-
taxy with setae a’, on abdominal terga I-III and V present, dorsal setae thick abruptly
pointed at tips and serrated, empodial appendage with relatively narrow basal lamella

D. SKARZYNSKI & A. SMOLIS

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DESCRIPTION OF CERATOPHYSELLA ROBUSTISETA SP. N. 301

0.1 mm

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Ceratophysella robustiseta sp. n.: 1, postantennal organ and ocelli; 2-4, postantennal organ;
5, chaetotaxy of thoracic tergum II; 6, chaetotaxy of abdominal terga III-VI; 7, dens and mucro;
8, tibiotarsus and claw III.

302 D. SKARZYNSKI & A. SMOLIS

and apical filament reaching about 3/4 of inner edge of claw, furca shortened with very
delicate cuticular skeleton, dens with 6-7 unmodified setae, basal macrochaeta longer
than dens and mucro small with low lateral lamella.

DESCRIPTION: Body length of males 1.1-1.3 mm, females 1.1-1.5 mm. Body
colour greyish-black, paler ventrally. Granulation almost uniform, coarser on last
abdominal terga, 6-12 granules between setae p, on abdominal tergum V. Dorsal
chaetotaxy of thorax and abdomen as in Figs 5-6. Chaetotaxy of head typical for the
genus. Dorsal macrochaetae and microchaetae relatively short, thick, abruptly pointed
at tips and clearly serrated (Figs 1, 5). Tip of dorsal setae usually with light external
layer (Fig. 1). Body sensilla short (ratio p3/p4(s) and pg/m7(s) on thoracic tergum II =
1.3-1.5 and 1.6-2 respectively), fine and smooth. Setae m, on thoracic terga II-III
absent, mg present. Setae az on thoracic tergum II distinctly longer than a, and a).
Microsensillum (ms) on thoracic tergum II present. One additional seta present outside
lateral sensillum my on thoracic tergum III (see Deharveng & Bourgeois 1991: 309,
Fig. 11). Setae a’, on abdominal terga I-III usually present. Setae p, on abdominal
tergum IV microchaetae, p, macrochaetae. Abdominal tergum V with 4 + 4 a-setae
inside the two ps macrochaetae (a’, present). Subcoxae I, II, III with 1, 3(4), 3(4) setae
respectively.

Antennal segment IV with simple apical vesicle, subapical organite, microsen-
sillum, 6-7 cylindrical, subequal sensilla in typical arrangement (sensilla d sometimes
absent) and about 10-15 thin, slightly curved and blunt-tipped sensilla in ventral file.
Antennal HI-organ with two long (lateral) and two short (internal) curved sensilla.
Microsensillum on antennal segment III present. Eversible sac between antennal
segments III-IV absent. Antennal segment I with 7 setae.

Ocelli 8 + 8. Postantennal organ about twice as long as diameter of ocellus B,
variable in shape. Four primary lobes of postantennal organ with more or less
numerous finger-like papillae (Figs 1-3), sometimes without distinct papillae (Fig. 4).
Accessory boss invisible.

Labrum with 5, 5, 4 setae and 4 rounded distal papillae. Four prelabral setae.
Head of maxilla as in C. denticulata (see Fjellberg, 1984). Maxillary outer lobe with
1 sublobal hair. Labium as in C. denticulata (see Fjellberg, 1998/1999), with 6 pro-
ximal setae.

Tibiotarsi I, II, IH with 19, 19, 18 setae respectively, clavate tenent hairs absent
(Fig. 8). Claws with distinct inner tooth and two pairs of indistinct lateral teeth (Fig.
8). Empodial appendage with relatively narrow basal lamella and apical filament
reaching about 3/4 of inner edge of claw (Fig. 8). Ventral tube with 4 + 4 setae.

Furca shortened (ratio dens + mucro/inner edge of claws III = ca 1.5), cuticular
skeleton of furca very delicate, sometimes invisible. Dens with uniform granulation
and 6-7 unmodified dorsal setae (basal macrochaeta longer than dens) (Fig. 7). Mucro
small, boat-like with low lamellae (Fig. 7). Retinaculum with 4 + 4 teeth.

Anal spines long (about 1.2 as long as claws III), slightly curved, situated on
basal papillae (Fig 6). Colour of anal spines more or less yellowish.

DISCUSSION: C. robustiseta sp. n. resembles species with a moruliform post-
antennal organ such as C. morula and C. mosquensis from which it differs in the

DESCRIPTION OF CERATOPHYSELLA ROBUSTISETA SP. N. 303

characters summarized in Tab. 1. Moreover a set of characteristic features: the absence
of eversible sac between antennal segments III-IV, the presence of only slightly modi-
fied sensilla in ventral file of antennal segment IV, thick dorsal setae, furca shortened
with very delicate cuticular skeleton, dens with long basal macrochaeta and small
mucro place this species near C. michalinae Skarzyfiski, 2005. Nevertheless it can be
easily separated from this species by the following characters (see Skarzyfiski, 2005):
postantennal organ from almost smooth to weakly moruliform (v. smooth), setae a’, on
abdominal tergum V present (v. absent), empodial appendage with relatively narrow
basal lamella and apical filament reaching about 3/4 of inner edge of claw (v. broad
basal lamella and apical filament reaching 1/2 of inner edge of claw), 6-7 dental setae
(v. 4-7).

ACKNOWLEDGEMENTS

We wish to express our sincere thanks to Dr Charles Lienhard for loans of
Yosii’s material of C. postantennalis and H. nepalica and to Dr Louis Deharveng for
supplying holotype of C. morula.

REFERENCES

BABENKO, A. B., CHERNOVA, N. M., POTAPOV, M. B. & STEBAEVA, S. K. 1994. Collembola of
Russia and adjacent countries: Family Hypogastruridae. Nauka, Moscow, 336 pp.

BOURGEOIS, A. & CASSAGNAU, P. 1972. La différenciation du type Ceratophysellien chez les
Collemboles Hypogastruridae. Nouvelle Revue d’Entomologie 2: 271-291.

DEHARVENG, L. & BOURGEOIS, A. 1991. Two new species of Ceratophysella (Collembola:
Hypogastruridae) from Thailand. Acta Zoologica Cracoviensia 34: 303-310.

FJELLBERG, A. 1984. Maxillary structures in Hypogastruridae (Collembola). Annales de la
Société Royale Zoologique de Belgique 114: 89-99.

FJELLBERG, A. 1999. The Labial Palp in Collembola. Zoologischer Anzeiger 237: 309-330.

SKARZYNSKI, D. 2004. Taxonomic status of Ceratophysella denticulata (Bagnall, 1941), C. en-
gadinensis (Gisin, 1949) and C. stercoraria Stach, 1963 (Collembola: Hypogastruridae)
in the light of laboratory hybridisation studies. Insect Systematics and Evolution 35:
277-284.

SKARZYNSKI, D. 2005. Ceratophysella michalinae, a new species from Poland (Collembola:
Hypogastruridae). Genus 16: 1-5.

Yosu, R. 1966. Collembola of Himalaya. Journal of the College of Arts and Sciences Chiba
University 4: 461-531.

Yosa, R. 1971. Collembola of Khumbu Himal. /In: Khumbu Himal.

n dat

Br,
1) AL
on

REVUE SUISSE DE ZOOLOGIE 113 (2): 305; juin 2006

Nomenclatural note on the genus Nans (Ostariophysi, Characidae)

Juan Marcos MIRANDE!, Gastén AGUILERA! &

Marfa de las Mercedes AZPELICUETA?

! Fundaciön Miguel Lillo, Miguel Lillo 251, 4000 Tucumän, Argentina.
E-mail: memirande @tucbbs.com.ar

2 Divisién Zoologia Vertebrados, Facultad de Ciencias Naturales y Museo, Paseo del
Bosque, 1900 La Plata, Argentina. E-mail: azpeli@museo.fcnym.unlp.edu.ar

Nomenclatural note on the genus Nans (Ostariophysi, Characidae). -
The generic name Nans Mirande, Aguilera & Azpelicueta, 2004 is pre-
occupied by an annelid. The monotypic fish genus Nans is therefore
renamed to Nantis.

Keywords: Characiformes - Nans - Nantis - homonymy - new name.

Nantis nom. nov.

TYPE SPECIES: Nans indefessus Mirande, Aguilera & Azpelicueta, 2004.

ETYMOLOGY: Nantis is deriving from the latin verb nare (to swim) (artificial
noun). Gender: masculine.

Nans indefessus Mirande, Aguilera & Azpelicueta, 2004 was described as a new
genus and species from the upper Rio Bermejo basin, Northwestern Argentina.
According to Neave (1940) the generic name Nans is preoccupied by an annelid
(Chamberlin, 1919: 125). In order to clear this homonymy we propose Nantis nom.
nov. as a new substitute name for the genus Nans Mirande, Aguilera & Azpelicueta,
2004.

ACKNOWLEDGEMENTS
We thank John K. Page who called our attention to this homonymy.

REFERENCES

CHAMBERLIN, R. V. 1919. The Annelida Polychaeta. Memoirs of the Museum of Comparative
Zoology, Harvard University 48: 1-514.

MIRANDE, J. M., AGUILERA, G. A. & AZPELICUETA, M. M. 2004. A new genus and species of small
characid (Ostariophysi, Characidae) from the upper rio Bermejo basin, northwestern
Argentina. Revue suisse de Zoologie 111(4): 715-728.

NEAVE, S. A. 1940. Nomenclator Zoologicus. A list of the names of genera and subgenera in
zoology from the tenth edition of Linnaeus 1758 to the end of 1935. Volume III M-P. The
Zoological Society of London.

Manuscript accepted 04.11.2005

mei
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yoda,

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Sri
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REVUE SUISSE DE ZOOLOGIE 113 (2): 307-323; juin 2006

Taygete sphecophila (Meyrick) (Lepidoptera; Autostichidae):
redescription of the adult, description of the larva and pupa, and
impact on Polistes wasps (Hymenoptera; Vespidae) nests in the
Galapagos Islands

Bernard LANDRY (BL)!, David ADAMSKI (DA)?, Patrick SCHMITZ (PS)3,
Christine E. PARENT (CEP)* & Lazaro ROQUE-ALBELO (LR)
1 Muséum d'histoire naturelle, C.P. 6434, 1211 Genève 6, Switzerland.
E-mail: bernard.landry @ ville-ge.ch
2 Department of Entomology, National Museum of Natural History, P.O. Box 37012,
NHB - E523, Smithsonian Institution, Washington, DC, 20013-7012, USA.
E-mail: dadamski@sel.barc.usda.gov
3 Muséum d'histoire naturelle, C.P. 6434, 1211 Genève 6, Switzerland.
E-mail: patrick.schmitz@ville-ge.ch
4Behavioural Ecology Research Group, Department of Biological Sciences,
Simon Fraser University, Burnaby, British-Colombia V5A 1S6, Canada.
E-mail: cparent@sfu.ca
5 Department of Entomology, Charles Darwin Research Station, Casilla 17-01-3891,
Quito, Ecuador; Biodiversity and Ecological Processes Research Group, Cardiff
School of Biosciences, Cardiff University, PO Box 915, Cardiff CF10 3TL, United
Kingdom. E-mail: lazaro@fcdarwin.ec.org

Taygete sphecophila (Meyrick) (Lepidoptera; Autostichidae): redescrip-
tion of the adult, description of the larva and pupa, and impact on
Polistes wasps (Hymenoptera; Vespidae) nests in the Galapagos Islands.
- Taygete sphecophila (Meyrick) (Lepidoptera; Autostichidae) is reported
on the Galapagos Islands. The morphology of the moth, larva, and pupa are
described and illustrated in details. Part of the mitochondrial DNA was
sequenced and made available on GenBank. The incidence of predation by
I. sphecophila on nests of Polistes versicolor Olivier (Hymenoptera;
Vespidae) was measured in four different vegetation zones of Floreana and
Santa Cruz Islands. The percentages of infested nests varied greatly (from
13.9% to 66.7% on Floreana and from 20.0 to 100% on Santa Cruz) and no
clear ecological trends could be ascertained.

Keywords: Micro moths - Autostichidae - Taygete - Polistes - Galapagos
Islands - mitochondrial DNA - larval predation - morphology - ecology.

INTRODUCTION

Taygete was described by Chambers (1873) to accommodate Evagora diffi-
cilisella Chambers, 1872 (Nye & Fletcher, 1991). The latter name proved to be a
synonym of 7. attributella (Walker, 1864). The genus appears to be restricted to the

Manuscript accepted 06.12.2005

308 B. LANDRY ET AL.

New World. Becker (1984) lists 13 names in this genus for the Neotropical fauna while
Hodges (1983) lists six species for the North American fauna, including five that are
stated to be misplaced in this genus. BL's examination of the type specimens of the
Neotropical species at the Natural History Museum, London, points to the possibility
that only 7. sphecophila (Meyrick, 1936) is congeneric with 7. attributella in this re-
gion. However, the types of Epithectis consociata Meyrick, E. notospila Meyrick, and
E. altivola Meyrick have lost their abdomen and cannot be assigned to genus, and the
type of E. lasciva Walsingham, deposited in the USNM, Washington, could not be
found.

Taygete Chambers was considered to belong to the Gelechiidae until Landry
(2002) moved it to the Autostichidae, Symmocinae sensu Hodges (1998). Taygete
sphecophila was described from three specimens bred in Trinidad from "bottom of
cells of the Hymenopteron Polistes canadensis" (Meyrick, 1936). The moth and male
genitalia were later illustrated with black and white photography by Clarke (1969). On
the Galapagos Islands moths of 7. sphecophila were first collected in 1989 by BL, but
the species probably arrived earlier within nests of Polistes versicolor Olivier
(Vespidae).

The purposes of this paper are to redescribe and illustrate the moth of 7. sphe-
cophila, to describe and illustrate the larva and pupa, to present part of its mito-
chondrial DNA, and to report on a few aspects of its biology, particularly with regard
to the incidence of damage to P. versicolor nests by larvae.

MATERIAL AND METHODS

Moths of 7. sphecophila were first collected at night with a mercury vapor light
set in front of a white sheet and powered by a small generator, and with an ultra-violet
lamp powered by a battery. Other adult specimens were reared from contained nests of
Polistes versicolor. Immature stages were found by dissecting Polistes nests and by
exposing them to the sun, which causes larvae to exit nests and run away from them
(Fig. 2).

Specimens are deposited in the Charles Darwin Research Station (CDRS),
Santa Cruz, Galapagos, Ecuador; the Canadian national Collection of Insects (CNC),
Ottawa, Ontario, Canada; the United States National Museum of Natural History,
Washington, D.C., U.S.A. (USNM), and the Muséum d'histoire naturelle (MHNG),
Geneva, Switzerland.

For the study of specimens using electron microscopy, larvae and pupae were
first rinsed several times in water, cleaned in 10% EtOH with a camel hairbrush, and
then dehydrated in EtOH as follows: 10% EtOH for 15 minutes, 20% for 15 minutes,
40% for 15 minutes, 70% for 1/2 hour, 90% for 1/2 hour, and 100% for 1/2 hour each
in two separate baths. After dehydration, specimens were critical-point dried using a
Tousimis critical point dryer, mounted on stubs, and coated with gold-palladium (40-
60%) using a Cressington sputter coater. The ultrastructure of the larvae and pupa was
studied with an Amray scanning electron microscope.

Gross morphological observations and measurements of the larvae and pupae
were made using a dissecting microscope (reflected light) with a calibrated micrometer.

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 309

Fics 1-2

1, Taygete sphecophila, female; 2, part of an abandoned nest of Polistes versicolor exposed to
the sun with at least 8 larvae of Taygete sphecophila exiting from it.

Maps of the larval chaetotaxy were initially drawn using a WILD dissecting micro-
scope with a camera lucida attachment. Terminology for chaetotaxy follows Stehr
(1987).

310 B. LANDRY ET AL.

In order to certify that the larvae corresponded to the adults found we sequenced
a fragment of the mitochondrial gene Cytochrome oxidase I (COI) of both. Whole
genomic DNA was extracted using the Nucleospin kit (Macherey-Nagel). The COI
gene was amplified by PCR with two primers: k698 (5’-TACAATTTATCGCC-
TAAACTTCAGCC-3’), and Pat2 (5’-TCCATTACATATAATCTGCCATATTAG-3’).
The thermal profile started with an initial denaturation at 95°C for 5 min, followed by
35 cycles at 94°C for 30 s, 47°C for 30 s, and 72°C for 1 min 30 s, and a final step at
72°C for 10 min. The purified PCR product was sequenced in both directions using
fluorescent dye terminators in an ABI 377 automated sequencer. The sequence is
available from GenBank (Accession No. DQ309437).

In order to determine the distribution and the density of Taygete sphecophila as
predator on Polistes versicolor nests, several study sites were selected in four of the
vegetation zones of Santa Cruz and Floreana Islands. In each vegetation zone a series
of quadrats of 10 m x 10 m were made at random, and the number of active and
inactive nests of Polistes versicolor were counted. The delimitation of vegetation zones
was based on vegetation composition (Wiggins & Porter, 1971). Nests were found by
visually searching the study sites. In addition, nests found in and near Puerto Ayora, a
small town located on the littoral and arid zones on the south coast of Santa Cruz
Island, were included in the study. The presence of 7. sphecophila in Polistes nests was
determined by the presence of little holes on the back of the nests (Fig. 2) and distinc-
tive breaches on the capped cells normally occupied by wasp pupae. In 1999, nests of
Polistes versicolor were monitored weekly in the area of Puerto Ayora, and nests that
were abandoned after being infested by 7. sphecophila were collected during that
period of time. Some adults of 7. sphecophila that emerged from these nests were
preserved dry for taxonomic identification. The ecological observations were made
between April and August 1999, February and April 2002 and 2003 on Santa Cruz
Island, and between April and August 1999 on Floreana Island. To test for ecological
or insular trends in the frequency of parasitism of P. versicolor nests by 7. sphecophila,
we performed a G-test for goodness of fit (Sokal & Rohlf, 1995) on each island dataset
using the proportion of P. versicolor nests in a given zone to infer the expected
frequency of parasitism by 7. sphecophila.

TAXONOMIC TREATMENT

Taygete sphecophila (Meyrick)

Epithectis sphecophila Meyrick, 1936: 624; Gaede, 1937: 113; Clarke, 1955: 290; Clarke, 1969:
63, pl. 31 figs 4-4b; Makino, 1985: 25; Yamane, 1996: 85.
Taygete sphecophila (Meyrick); Becker, 1984: 47; Landry, 1999: 68; Landry, 2002: 818-819.

MATERIAL EXAMINED FOR MORPHOLOGICAL WORK: Moths (13 specimens from the
Galapagos Islands, Ecuador): SANTA CRUZ: 1 2, C[harles] D[arwin] R[esearch] S[tation], arid
zone, 19.1.1989, M[ercury] V[apor] L[amp] (B. Landry); 4 9 (two dissected with genitalia on
slides CNC MIC 4586 & BL 1196, the latter with right wings on slide BL 1313), CDRS, arid
zone, 3.11.1989, MVL (B. Landry); 2 & (one dissected, slide BL 1126), Barranco, ex larva en
nido Polistes versicolor, 8.11.1999 (L. Roque, No. 99.20); 1 9, NNW Bella Vista, GPS: 225 m
elev., S 00° 41.293”, W 090° 19.665’, 18.11.2005, u[Itra] v[iolet] I[ight] (B. Landry, P. Schmitz);
1 © (dissected, slide BL 1195), 2 km W Bella Vista, 27.11.1989, MVL (B. Landry); 1 ©, casa L.
Roque-Albelo & V. Cruz, GPS: 137 m elev., S 00° 42.595’, W 090° 19.196’, 27.11.2005, uvl (B.

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 311

FIG. 3

Taygete sphecophila, male genitalia (sizes not proportionate). 3a, dorsal view of valvae + vin-
culum + juxta and ventral view of tegumen + uncus + gnathos detached on right side and spread
on left side, phallus removed, setae shown on right side only; 3b, side view of phallus with vesica

everted; 3c, dorsal view of phallus, vesica inverted, scale = 0.1 mm; 3d, lateral view of whole
genitalia.

312 B. LANDRY ET AL.

Landry); 2 3 (dissected, slides BL 1208 & 1209), émergé d’un nid de Polistes, 1999 (C. Parent).
SAN CRISTOBAL: 1 à , antiguo botadero, ca. 4 km SE P[uer]to Baquerizo, GPS: 169 m elev.,
S 00° 54.800’, W 089° 34.574’, 22.11.2005, uvl (B. Landry).

Larvae (166 specimens) and pupae (10 specimens) collected on Santa Cruz by P.
Schmitz in 2004 and 2005.

DIAGNOSIS: The presence in males of this species of a corematal organ at the
base of the abdomen (Fig. 4) and a trifurcated uncus (Fig. 3a) are excellent diagnostic
features with regards to the rest of the Galapagos fauna. Males of Galagete Landry are
the only other Galapagos moths to share a corematal organ, but their uncus is made of
a single projection. In females the shape of segment VII (Fig. 5), especially dorsally,
will separate 7. sphecophila from any other species in the Galapagos and probably the
rest of its range. On the archipelago, some species of Galagete Landry (2002) or
Gelechiidae may appear superficially similar, especially because they share a similarly
shaped hindwing, a similar wingspan, upturned labial palpi, and scales on the proboscis
basally, but the forewing markings of 7. sphecophila (Fig. 1) are unique among these
groups.

REDESCRIPTION: General appearance of moth greyish brown with dark brown
markings on forewing (Fig. 1); scales usually dark brown at their base and paler
apically. Head scales longer laterally and directed medially and ventrally, except on
occiput, directed medially and dorsally. Ocellus and chaetosema absent. Labial palpus
gently curving upward, darker brown laterally than medially, with white rings of scales
mostly at apex of segments; segments II and III shorter together than segment I.
Antenna mostly greyish brown, darker brown toward base; flagellomeres in both sexes
simple and with erect scales ventrally from about middle of flagellum. Thorax conco-
lorous with head, sometimes darker brown at base. Foreleg mostly dark brown, with
beige scales at apex of tarsomere I and on all of tarsomere V. Midleg mostly dark
brown laterally, with paler scales at apex of tarsomeres I and II, and on all of tarsomere
V, uniformly beige medially on femur and tibia, also with short tuft of dark brown
scales dorsally on basal half of tibia. Hindleg paler than other legs, with some dark
brown laterally on femur and tibia, mostly dark brown on tibial spines and at base of
tarsomeres I-IV, also with tuft of long dirty white scales on dorsal margin of tibia.
Wingspan: 7.5-9.0 mm. Forewing mostly greyish brown, with three dark brown trian-
gular markings on costa, largest marking at base, reaching inner margin, smallest sub-
medially situated, barely reaching cell, third marking large, reaching middle of wing;
with dark brown scaling also at apex and as 1-3 small patches of 10 scales or less below
postmedian costal marking; also with variable amounts of yellowish-orange to rusty-
brown scales usually within basal dark brown marking, below postmedian marking,
and toward apex; fringe dark brown at apex, more greyish brown elsewhere. Hindwing
greyish brown without markings, with concolorous fringe. Wing venation (based on
slide BL 1313, female) (Fig. 6): Forewing Sc to about 2/5 wing length; R1 from about
middle of cell; R2 and R3 separate, both from before upper angle of cell; R4, RS, and
MI from upper angle of cell, connected, R4 and RS directed toward costa before apex,
M1 directed toward outer margin below apex; M2 and M3 separate, M2 from lower
angle of cell, M3 from shortly before lower angle of cell; CuA1 and CuA2 separate,
both from shortly before lower angle of cell; CuP absent; cell a little more than half

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 313

Fics 4-5

Taygete sphecophila. 4, ventral view of first abdominal segment; 5, ventral view of female
genitalia, setae shown on right side only.

wing length; A1 and A2 joined at about 1/5 their lengths. Female forewing retinaculum
consisting of anteriorly directed scales at base of cubital stem and posteriorly directed
scales at base of Sc. Hindwing Sc closely following costa, reaching it at about 3/5 wing
length; Rs connected with M1 after upper angle of cell, Rs reaching costa at about 4/5

314 B. LANDRY ET AL.

wing length, M1 directed toward apex; M2 from slightly above lower angle of cell,
reaching outer margin below middle; M3 and CuAl connected for about 1/2 their
lengths after lower angle of cell, M3 to tornus, CuAl to inner margin shortly before
tornus; CuA2 from about 2/3 cell to inner margin at 7/10; CuP and anal veins indis-
tinct; apex distinctly produced; outer margin distinctly concave; female frenulum with
2 acanthae. Abdomen dorsally mostly dark greyish brown, with dirty white scales at
apex of all segments except last; ventrally dark brown on each side of large dirty white
band except for last segment, mostly concolorous, greyish brown; male first abdominal
segment (Fig. 4) ventrally with an invaginated pouch containing a membranous
structure bearing scales (see Note below).

Male genitalia (Fig. 3). Uncus moderately long, with pair of fixed lateral,
pointed and gently tapering glabrous projections; also with movable median projec-
tion, slightly longer than lateral projections, enlarged at apex and bifid, with each end
bulbous and setose, also slightly setose at base laterally. Gnathos a long curved rod
pointing posteriorly, apically more heavily sclerotized, tapered, glabrous, and rounded.
Tegumen broad medially, with moderately narrow pedunculi. Valva with unsclerotized
setose cucullus, tapering, rounded apically, with slightly sclerotized setose ridge at
base on inner side, also with medium sized apodemes directed anteriorly from base of
costa; sacculus with pair of short, narrow, setose, and apically rounded projections,
dorsal projection curved and directed dorsally, ventral one straight and directed
posteriorly. Vinculum narrow, slightly projected anteriorly and upturned. Juxta poorly
developed, small, better sclerotized at posterior edge around phallus. Phallus (= aede-
agus of authors, but see Kristensen, 2003) narrow, with shaft flattened dorsoventrally
beyond middle, better sclerotized on left side in narrow band, slightly upturned
apically; coecum penis medium-sized with pair of very small peduncles laterally;
vesica with minute scobination.

Female genitalia (Fig. 5). Papillae anales large and long, moderately setose,
sclerotized dorsally and laterally at base. Posterior apophyses slightly curved apically,
slightly longer than papillae. Tergum VIII well sclerotized, with few long setae espe-
cially on margin, with deep rounded concavity in middle apically; middle of concavity
with posteriorly directed projection variable in length and bearing two setae. Anterior
apophyses straight, slightly enlarged apically, about as long as papillae. Sternum VIII
with apical margin bell shaped, well sclerotized, with few long setae mostly posteriorly
along margin and midventrally. Intersegmental membrane between sternites VII and
VII slightly sclerotized on each side of midventral line and with pair of short projec-
tions inside body at apical margin. Ostium bursae in middle of sternite VIII, ventrally
protected by slightly protruding crescent of sclerotization. Ductus bursae short,
gradually enlarging, basal half well sclerotized, distal half spiculose and with wrinkles
patterned like brood cells in bee hive. Corpus bursae slightly longer than wide, spicu-
lose, with one large, spiny, curved, and pointed cornutus; latter set in small sclerotized
patch with pair of bumps on each side of its base.

DESCRIPTION OF THE LARVA AND PUPA: Larva. (Figs 7-17): Length 5.0-8.2 mm (n
= 72), < 5.0 mm (n = 94). Body pale gray, textured with microconvolutions; head
capsule amber; prothoracic shield amber, gradually darkening posteriorly; pinacula
pale brown; anal plate pale amber; setae with widened, circular, and slightly raised

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 315

FIG. 6
Wing venation of Taygete sphecophila.

sockets. Head (Figs 7-10, 17): hypognathous, textured with slightly raised, confluent,
polygonal ridges except on area between adfrontal sclerites (Figs 7-8); adfrontal scle-
rites widened distally, frontal setae about equal in length, AF2 above apex of frons,
AFI below; F1 slightly closer to AF1 than to C1; C2 at least 2 1/2 times longer than
C1; clypeus with 6 pairs of setae, 3 pairs on medial half, 3 on distal half; mandible
angular (Fig. 17), shallowly notched subapically forming small apical dentition,
bearing pair of subequal setae on outer surface near condyle, and with 1 large dentition
on inner surface; sensilla types and arrangement on antenna (Fig. 9) and on maxillary
palpi (Fig. 10) similar to those of other Gelechioidea studied by Adamski & Brown
(1987), Adamski (1999), Adamski & Pellmyr (2003), Landry & Adamski (2004), and
Wagner et al. (2004), and other Lepidoptera studied by Adamski & Brown (2001),
Albert (1980), Avé (1981), Grimes & Neunzig (1986a, b), and Schoonhoven & Dethier
(1966). Three stemmata in genal area, 1 approximate pair above antenna, and 1 stemma
below antenna; substemmatal setae about equal in length, arranged as in Fig. 8; S3 and
S1 elongate and about equal in length, S2 short; S3 lateroventral to S2, S2 approximate
to stemma 3, and S1 approximate to stemma 5 (stemmata 1, 2, and 6 absent); A-group
setae above gena, mesal to L1; P1 dorsolateral to AF2, P2 dorsomesal to P1. Thorax
(Figs 11, 14): T1 with L-group trisetose, on large pinaculum extending beneath and
posterior of spiracle; setae anterior to spiracle; L1 approximate and posteroventral to
L2, about 2 1/2 times lengths of L2 and L3; SV-group setae on anterior part of elon-
gate pinaculum; SV1 about 1/3 longer than SV2; coxae nearly touching, VIs very
approximate (not shown); segments of leg textured with slightly elongate ridges, many
produced distally into hairlike spines, claw single (Fig. 11); shield with SD1 slightly
posterior to and about 1/3 longer than XD2 and XD1; XD2, XD1, D1, and SD2 about

316 B. LANDRY ET AL.

Fics 7-13

Scanning electron micrographs of larva of Taygete sphecophila. 7, Frontolateral view of head
capsule, scale = 100 u; 8, Ventrolateral view of head capsule, scale = 100 u; 9, Sensilla of
antenna: 1 = sensilla basiconica, 2 = sensillum chaetica, 3 = sensillum styloconicum, 4 = sen-
sillum trichodeum, scale = 10 u; 10, Sensilla of maxillary palpus: A2 = sensillum styloconicum,
Al, A3, MI-2, L1-3 = sensilla basiconica, SD = sensillum digitiform, scale = 10 u; 11, Distal
portion of left prothoracic leg showing claw, scale = 10 u; 12, Left proleg on A4, scale = 100 u;
13, Anal plate of A10, scale = 100 u.

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 317

equal in lengths, XD2 about twice distance from XD1 than from SD1; DI in straight
line with XDI, slightly posterior to SD2 and D2; D2 about same length as SDI, in
straight line with SD2. T2-T3 (Fig. 14): D2 about 2 times length of D1, both on small
pinaculum; SD1 about 2 times length of SD2, both on small pinaculum; L1 about 1/3
longer than L2, both on small pinaculum, L3 slightly shorter than L2, posterior to or in
vertical line with SVI; MVI on anterior margin between T2-T3, slightly above SV1
(hard to see); Vis on T2-T3 about equal distance apart, at least 4 times distance
between V1s on Tl. Abdomen (Figs 12, 13, 15, 16): A1-A2 (Fig. 15): D2 and DI equal
in lengths or D2 slightly longer, MDI on anterior part of segment anteroventral to DI;
SD1 above spiracle, about 1/3 longer than D2, with minute SD2 (anterior part of pina-
culum); small opening on ventroposterior margin of pinaculum bearing SD1 and SD2;
spiracle on Al slightly larger than those on A2-A7; L1 2 times length of L2, both on
same pinaculum, slightly anterior of spiracle; L3 about same length as L2, anterior to,
in vertical line with, or posterior to D2; SV-group bisetose on Al, trisetose on A2, on
same pinaculum; V1s equal distance apart (not shown). A3-A10 (Figs 12, 13, 16): A3-
A6 with 4 pairs of protuberant prolegs, crochets biordinal, in circle (Fig. 12); setae as
above; A7 as above except, SV-group bisetose and on same pinaculum; A8 as above
except with spiracle slightly larger than on previous segments and SV-group unisetose;
A9 with D2 about 2-2 1/2 times longer than DI; DI anterior to D2 and SDI, equi-
distant to both setae; SD1 about same length as D1; L-group setae slightly anterior to
D1; L1 about 3 times length of L2, on same pinaculum; L3 slightly longer than L2;
SVI slightly shorter than L1; V1s as previous segments; A10 (Figs 13, 16): anal plate
with SD2 and SDI equal in lengths, about twice length of D2; DI slightly shorter than
D2; crochets of proleg biordinal, in semicircle, gradually shortened mesally and
laterally.

Pupa. (Figs 18-21): Length 3.6-4.6 (n = 10): amber, smooth, spiracles protu-
berant; all dorsal setae apically hooked except long seta associated with axillary
tubercle (Figs 19-20). Sclerites of antennae annulated, widely separated anteriorly,
gradually convergent from beyond basal 1/3 of sclerites of maxillae, fused for short
distance beyond distal apices of sclerites of maxillae, gradually divergent posteriorly,
exposing distal part of sclerites of hindlegs; sclerites of midleg not fused distally;
paired nodular scars of prolegs on A5-A6 (Fig. 18); A6-A10 fused, rotating as unit;
cremaster dorsolaterally flattened, trapezoidal basally, extending posterolaterally into 2
slightly divergent and elongate spine-like processes (Fig. 21).

DISTRIBUTION AND PHENOLOGY: The species was described from Trinidad
(Meyrick, 1936) and never mentioned from anywhere else subsequently. In the
Galapagos Islands it has been found on Floreana (from the littoral to the humid zones),
San Cristobal (in the arid zone), and Santa Cruz (from the littoral to the humid zones).
In the Galapagos we have collected live moths of this species in January, February,
March, April, September, November, and December.

NOTES: Preliminary phylogenetic analyses, both morphological and molecular,
support the placement of Taygete sphecophila within Autostichidae (PS, unpublished
data). For example, Kaila’s (2004) matrix was reanalyzed with 7. sphecophila data, and
the species clusters in Kaila’s autostichid assemblage with Galagete Landry.

318 B. LANDRY ET AL.

L3

[DIANAMARQUES

SVI

Fics 14-17

Larva of Taygete sphecophila. 14, Chaetotaxy of thorax; 15, Chaetotaxy of abdominal segments
1-2; 16, Chaetotaxy of abdominal segments 6-10; 17, Mandible.

A comparison of a 1283 base pairs fragment (consisting of most of the COI gene
except the first 254 base pairs) sequenced for a larva and an adult of Taygete spheco-
phila showed no substitution, which clearly indicates conspecificity.

The larva has only three stemmata, a condition that is highly unusual in
Gelechioidea and that may be due to the unique host relationship.

It proved impossible to evaginate the ventro-abdominal pouch (Fig. 4) in
several male specimens of this species. However, BL was able to evaginate this core-
matal organ from a specimen of Taygete attributella (Walker). The organ consists of a
narrow membranous tube, almost as long as the abdomen, on which narrow scales are
connected all around. The membrane of the tube is very thin and the tube collapsed as
soon as specimens were transferred to lactic acid for temporary storage. An illustration
of this structure for the closely related Galagete turritella Landry is provided by
Landry (2002: Figs 17, 18).

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 319

maxillary palp
maxilla

at
| | | femur of foreleg

foreleg
JETS midleg
: Al N
antenna
| = Sl,

forewing

A4
hindleg

[ “yo A5

N ze A6

x [ see A7
Li | LE — A8 - A10 5
18 19 X A 21 à

20 N

Fics 18-21

Pupa of Taygete sphecophila. 18, Ventral view; 19, Dorsal view; 20, Segments 8-10; 21, Lateral
view.

ECOLOGICAL STUDY

PATTERNS OF PREDATION

Although egg-laying was never observed, it is possible that the female moths
lay their eggs within the pupal cells of P. versicolor through numerous small holes of
1-2 mm in diameter that we observed on the back of the nests. In a sample of 25
P. versicolor nests, the number of 7. sphecophila moths found per nest varied between
3 and 13. However, 42 T. sphecophila larvae were recovered by PS from a rather small
nest collected on Santa Cruz in 2004. The food source needed for the development of
the moth’s larvae are the wasps’ pupae which are defenseless because of their isolation
in their capped cells. When ready to emerge from the wasp's cell, the moth makes a
distinctive breach through the cap covering the top of the cell.

DISTRIBUTION OF INFESTED NESTS

The level of 7. sphecophila infestation could only be assessed for nests of
P. versicolor that were abandoned. A total of 103 such nests were found on the different
study sites on Santa Cruz Island between 1999 and 2003, and 141 nests on Floreana
Island in 1999. The percentages of nests that presented signs of predation by 7. sphe-
cophila are given in Table 1, along with the vegetation zones in which they were found.

320 B. LANDRY ET AL.

TABLE 1. Percentage of Polistes versicolor nests found in four different vegetation zones of Santa
Cruz and Floreana Islands presenting signs of Taygete sphecophila predation. Number of nests
per sample are in parentheses.

Vegetation Zone % of nests with 7. sphecophila predation

Santa Cruz Island Floreana Island
Littoral 3555 (n=!) 40.0 (n=5)
Arid 43.0 (n=79) 13.9 (n=101)
Transition 20.0 (n=5) 66.7 (n=6)
Humid 100.0 (n=2) S157 =29))

On Santa Cruz island the arid zone was the area of highest abundance of nests.
This result is similar to that obtained by Roque-Albelo & Causton (1999) for abun-
dance of adult foragers. The percentages of infestation varied between zones (Table 1).
However, very few nests were collected in the littoral, transition, and humid zones.
Nests of P. versicolor were again more common in the arid zone of Floreana. However,
only 13.9% of them were infested by 7: sphecophila in this zone. In contrast to Santa
Cruz, on Floreana nests also were abundant in the humid zone, where 51.7 % of them
were infested.

The results of the G-test for goodness of fit allow us to test for ecological trend
in nest infestation according to vegetation zonation. The proportion of parasitism in
Polistes versicolor nests in the four vegetation zones on Santa Cruz Island does not
show deviation from the expected (based on the proportion of P. versicolor nests found
in each vegetation zone; G = 4.806, df = 3, P > 0.05). However, the situation on
Floreana Island appears different as P. versicolor nests found in the arid zone are
infested by 7. sphecophila less than expected, and nests found in the transition and
humid zones are infested more than expected (G = 15.482, df = 3, P< 0.01).

DISCUSSION

Different factors, including climatic conditions, infestation by nest scavengers
and parasitoids, and predation affect the wasp colony cycle (Yamane, 1996). Across its
range of distribution, from Costa Rica to Southern Argentina, P. versicolor seems to
prefer dry forest habitats (Richards, 1978). Data from previous studies suggest that in
the Galapagos the wasps are more abundant in the arid zone of the islands (Roque-
Albelo & Causton, 1999; Lasso, 1997). This preference in distribution could be asso-
ciated with climatic conditions (Parent, 2000). In the Galapagos the higher zones of the
islands are cooler and receive more rainfall than lower zones, particularly on the
southern slopes, and this factor probably affects nest development. Collection data of
T. sphecophila suggest a similar pattern of distribution. Most moth specimens were
collected in the dryer zones of the islands suggesting a close correlation with nest
abundance.

On Santa Cruz Island the occurrence of 7. sphecophila in different vegetation
zones is a reflection of the frequency of P. versicolor nests. However, 7. sphecophila
seems to be more abundant than expected in the transition and humid zones of Floreana
Island and less frequent in the arid zone. Therefore, 7: sphecophila’s occurrence on
Floreana Island is not strictly a reflection of the abundance of P. versicolor nests,

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 321

suggesting that other ecological or climatic factors might influence its distribution. It
is not clear why there is such a difference between Floreana and Santa Cruz islands,
but one possible hypothesis is that 7. sphecophila has colonized these two islands at
different points in time, so that populations on one of the island have had more time to
adapt to the island’s ecological and climatic context.

Polistes nests, as in many other social wasps, are scavenged and parasitized by
various insects including more than 11 moth species from four families (Makino,
1985). Only Taygete sphecophila was found in the Galapagos, where the species
apparently prefers to attack large nests, and all infested nests collected were large
enough to presume that they were in an advanced stage of the reproductive phase. If
predation by 7. sphecophila is restricted to this stage of the wasp colonies the proba-
bilities for this moth species to be an effective agent of biological control are reduced.
These results support the idea of Miyano (1980) that parasitic and scavenging
Lepidoptera reduce notably the colony’s productivity but are not thought to be a direct
cause of colony failure. However, the possibility to use 7. sphecophila as a biological
control agent against P. versicolor needs to be evaluated.

We believe that the first individuals of Taygete sphecophila probably arrived
within a nest of P. versicolor built on some human-made structure that would have
traveled by boat from the continent. It is actually quite possible that both animals
arrived together on the Galapagos. The wasp was first detected in 1988 on Floreana,
and is thought to have arrived with a shipment of bananas (Abedrabbo, 1991), but
Eduardo Vilema, resident of Santa Cruz, says that he first saw a nest of Polistes versi-
color near Bella Vista, on Santa Cruz, in 1984 or 1985 (pers. comm. to BL in 2004).
And we think it unlikely that the wasps came on banana regimes as they are not known
to build their nests there.

ACKNOWLEDGEMENTS

BL would like to thank the authorities of the CDRS and Galapagos National
Park Service (GNPS) for providing authorizations and logistical support for sampling
micro moths in 1989, 1992, 2002, 2004, and 2005. The first two expeditions were
organized by Stewart B. Peck, Carleton University, Ottawa, and benefited from
financial support of the National Sciences and Engineering Council of Canada.
Professor Peck, as well as Bradley J. Sinclair and several others were excellent field
companions. BL is also grateful to Jean-Frangois Landry, Agriculture & Agri-Food
Canada, Ottawa, for providing lab space, equipment and technical support in 1999-
2000. We are thankful to curator Kevin Tuck of the Natural History Museum, London,
for his kind help to BL in London and for providing information on references
regarding 7. sphecophila and on specimens of other species. Financial support for BL’s
investigations in London in 2000 were provided by the Galapagos Conservation Trust,
United Kingdom, and the Charles Darwin Foundation.

PS would like to thank the CDRS and GNPS for allowing field work and
providing logistical support in the Galapagos in 2004 and 2005. Research funds to PS
were provided by the A. Lombard Foundation, the E. & L. Schmidheiny Foundation,
the Swiss Academy of Sciences, and the Department of cultural affairs of the city of
Geneva.

322 B. LANDRY ET AL.

CEP would like to thank the CDRS and GNPS for providing logistical support
and required permits to conduct field work in Galapagos in 1999. Research funds to
CEP were provided by CIDA, the Canadian International Development Agency.

DA thanks Diana Marques, scientific illustrator, for the illustrations (Figs
14-21) of the larva and pupa and the digital production of the plates, and Scott D.
Whittaker, Manager of the Scanning Electron Microscopy Laboratory, Smithsonian
Institution, Washington, DC, for the help with the preparation of the larval specimens
for study.

LR thanks Henry Herrera of CDRS for his help in the field and in the lab.

Finally, we are grateful to Lauri Kaila and Klaus Sattler for reviewing the
manuscript.

REFERENCES

ABEDRABBO, S. 1991. Nueva avispa introducida en las islas. Carta Informativa 31: 4.

ADAMSKI, D. 1999. Blastobasis graminea, new species (Lepidoptera: Gelechioidea: Coleo-
phoridae: Blastobasinae): A stem borer of sugar cane in Colombia and Venezuela.
Proceedings of the Entomological Society of Washington 101: 164-174.

ADAMSKI, D. & BROWN, R. L. 1987. A new Nearctic Glyphidocera, with descriptions of all stages
(Lepidoptera: Blastobasidae: Symmocinae). Proceedings of the Entomological Society
of Washington 89: 329-343.

ADAMSKI, D. & BROWN, J. W. 2001. Systematic review of the Ecdytolopha group of genera
(Lepidoptera: Tortricidae: Grapholitini). Entomologica Scandinavica, Suppl. 58: 1-86.

ADAMSKI, D. & PELLMYR, O. 2003. Immature stages and redescription of Blastobasis yuccae-
colella Dietz (Lepidoptera: Gelechioidea: Coleophoridae: Blastobasini), with notes on
its biology. Proceedings of the Entomological Society of Washington 105: 388-396.

ALBERT, P. J. 1980. Morphology and innervation of mouthpart sensilla in larvae of the spruce
budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). Canadian
Journal of Zoology 58: 842-851.

AVÉ, D. A. 1981. Induction of changes in the gustatory response by individual secondary plant
compounds in larvae of Heliothis zea (Boddie) (Lepidoptera, Noctuidae). Ph.D. disser-
tation, Department of Entomology, Mississippi State University, 89 pp., 4 tables, 37 figs.

BECKER, V. O. 1984. Gelechiidae (pp. 44-53). In: HEPPNER, J. B. (ed.). Atlas of Neotropical
Lepidoptera, Checklist: Part 1, Micropterigoidea - Immoidea. Dr W. Junk Publishers,
The Hague, xxvii + 112 pp.

CHAMBERS, V. T. 1873. Micro-Lepidoptera. The Canadian Entomologist 5: 221-232.

CLARKE, J. F. G. 1955. Catalogue of the Type Specimens of Microlepidoptera in the British
Museum (Natural History) described by Edward Meyrick. Volume 1. Trustees of the
British Museum, London, vii + 332 pp.

CLARKE, J. F. G. 1969. Catalogue of the Type Specimens of Microlepidoptera in the British
Museum (Natural History) described by Edward Meyrick. Volume 7. Trustees of the
British Museum (Natural History), London, 531 pages.

GAEDE, M. 1937. Familia: Gelechiidae (pp. 1-630). Jn: WAGNER, H., STRAND, E. & BRYK, F.
(eds). Lepidopterorum Catalogus, part 79, Vol. IV. Dr. W. Junk, Berlin.

GRIMES, L. R. & NEUNZIG, H. H. 1986a. Morphological survey of the maxillae in last-stage
larvae of the suborder Ditrysia (Lepidoptera): Palpi. Annals of the Entomological Society
of America 79: 491-509.

GRIMES, L. R. & NEUNZIG, H. H. 1986b. Morphological survey of the maxillae in last-stage
larvae of the suborder Ditrysia (Lepidoptera): Mesal lobes (Laciniogaleae). Annals of the
Entomological Society of America 79: 510-526.

TAYGETE SPHECOPHILA IN THE GALAPAGOS ISLANDS 323

HopGEs, R. W. 1983. Gelechiidae (pp. 19-25). In: HODGES, R. W. et al. (eds). Check List of the
Lepidoptera of America North of Mexico. E. W. Classey Ltd. and the Wedge Entomo-
logical Research Foundation, London, xxiv + 284 pp.

HoDGEs, R. W. 1998. The Gelechioidea (pp. 131-158). Jn: KRISTENSEN, N. P. (ed.). Handbook of
Zoology, Lepidoptera, Moths and Butterflies, Vol. 1: Evolution, Systematics, and Bio-
geography. Walter de Gruyter, Berlin & New York, x + 491 pp.

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

KRISTENSEN, N. P. 2003. Skeleton and muscles: adults (pp. 39-131). Jn: KRISTENSEN, N. P. (ed.).
Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol. 2: Morphology, Physio-
logy, and Development. Walter de Gruyter, Berlin & New York, x + 564 pp.

LANDRY, B. 1999. Quatre premières mentions de Lépidoptères pour la province de Québec
(Opostegidae, Gelechiidae et Tortricidae). Fabreries 24: 66-72.

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, J.-F. & ADAMSKI, D. 2004. A new species of Nanodacna Clarke (Lepidoptera:
Elachistidae: Agonoxeninae) feeding on the seeds of Austrocedrus chilensis (Cupres-
saceae) in andean Argentina. Journal of the Lepidopterists’ Society 58: 100-113.

Lasso, T. L. 1997. Ecologia e impacto de la avispa introducida (Polistes versicolor, Vespidae-
Hymenoptera) en las Islas Floreana y Santa Cruz, Galapagos Ecuador. B.Sc. Thesis,
Universidad Central del Ecuador, Quito, 211 pp.

MAKINO, S. 1985. List of parasitoides [sic] of Polistine wasps. Sphecos 10: 19-25.

MEYRICK, E. 1936. Exotic Microlepidoptera 4: 609-642.

MIYANO, S. 1980 Life tables of colonies of workers in a paper wasp, Polistes chinensis anten-
nalis, in central Japan (Hymenoptera: Vespidae). Research in Population Ecology 22:
69-88.

NyE, I. W. B. & FLETCHER, D. S. 1991. The generic names of moths of the World. Volume 6.
Microlepidoptera. Natural History Museum Publications, London, xxix + 368 pp.

PARENT, C. 2000. Life cycle and ecological impact of Polistes versicolor versicolor (Olivier)
(Hymenoptera: Vespidae), an introduced predatory wasp on the Galapagos Islands,
Ecuador. M.Sc. Thesis, Carleton University, Ottawa, Canada, 139 pp.

RICHARDS, O. W. 1978. The social wasps of the Americas excluding the Vespinae. Trustees of the
British Museum (Natural History), London, Publication No. 785, vii + 580 pp. + 4 pls.

ROQUE-ALBELO, L. & CAUSTON, C. 1999. Monitoreo de avispas introducidas en las Islas
Galapagos. Charles Darwin Research Station report, 12 pp.

SCHOONHOVEN, L. M. & DETHIER, V. G. 1966. Sensory aspects of host-plant discrimination of
Lepidopterous larvae. Archives néerlandaises de zoologie 16: 497-530.

SOKAL, R. R. & ROHLF, F. J. 1995. Biometry (3rd ed.). W. H. Freeman and Company, New York,
887 pp.

STEHR, F. 1987. Cosmopterigidae (Gelechioidea) (pp. 391-392). In: STEHR, F. (ed.). Immature in-
sects, Volume 1. Kendall/Hunt Publishing Company, Dubuque, Iowa, 754 pp.

WAGNER, D. L., ADAMSKI, D. & BROWN, R. L. 2004. A new species of Mompha Hbn. from
Mississippi (Lepidoptera: Gelechioidea). Proceedings of the Entomological Society of
Washington 106: 1-18.

WIGGINS, I. L. & PORTER, D. M. 1971. Flora of the Galapagos Islands. Stanford University Press,
Stanford, xx + 998 pp.

YAMANE S. 1996. Ecological factors influencing the colony cycle of Polistes wasps (pp. 75-97).
In: TURILLAZZY, S. & WEST-EBERHARD, M. J. (eds). Natural History and evolution of
paper-wasps. Oxford University Press, xiv + 400 pp.

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REVUE SUISSE DE ZOOLOGIE 113 (2): 325-410; juin 2006

The European athecate hydroids and their medusae
(Hydrozoa, Cnidaria): Capitata Part 1

Peter SCHUCHERT
Muséum 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): Capitata Part 1. - This study reviews all European hydrozoan
species belonging to the capitate families Acaulidae, Boreohydridae, Can-
delabridae, Cladocorynidae, Cladonematidae, Margelopsidae, Pennariidae,
Protohydridae, and Tricyclusidae. Updated diagnoses for the families and
genera are provided and existing taxonomic problems solved or at least out-
lined. Candelabrum verrucosum Bonnevie, 1898 is regarded as a valid
species and redescribed based on a new record from Greenland. Although
Spadix purpurea Gosse, 1853 may be a senior synonym of Candelabrum
cocksii (Cocks, 1854), the latter is regarded as the valid name, this because
the former name has not been used after 1899, while the latter has been
widely used [ICZN article 23.9.1.1]. Likewise, two senior synonyms of
Eleutheria claparedii Hartlaub, 1898 are declared as invalid as they have
never been used since their original introduction by Haeckel.

Keywords: Marine invertebrates - Cnidaria - Hydrozoa - Anthoathecata -
Capitata - descriptions - revision - taxonomy.

INTRODUCTION

This publication is the second in a series of taxonomic revisions of the
European Anthoathecata (=Anthomedusae, Athecata) which was begun with the
families Oceanidae and Pachycordylidae (Schuchert, 2004). Although it was initially
planned to continue with families of the suborder Filifera, some essential material
could not be obtained to complete them in time. Therefore, this second paper presents
some families of the suborder Capitata.

Following the suggestions of fellow colleagues, the geographic scope of the
fauna under investigation has been somewhat extended to match the territory covered
by the European Register of Marine Species (Costello et al., 2001). The new scope
now also includes the Azores, Iceland, eastern Greenland, Jan Mayen, the Barents Sea,
Svalbard, and Franz Joseph Land. The extension of the geographic scope does not add
many species. A single species needs to be added to the families treated in the first
paper of this series, namely Similomerona nematophora (Antsulevich, 1986). This
species is already mentioned and diagnosed in Schuchert (2004), but its type locality
was erroneously given as "Kurile Islands". Dr Antsulevich informed me that the

Manuscript accepted 19.10.2005

326 P. SCHUCHERT

species was described from material originating from the archipelago of Franz Joseph
Land, thus falling within the limits of the newly adopted coverage. Furthermore,
although the paper was authored by Antsulevich & Polteva (1986), the authorship of
the species name is due to Dr Antsulevich alone. Although the genus name Merona is
an artificial word, its ending is characteristic for the feminine gender, thus requiring a
change of the specific epithet to nematophora.

The current paper treats several families of the suborder Capitata. The families
reviewed here were chosen somewhat arbitrarily and do not form a natural unit, though
some are evidently closely related. The selection had to be based on the material
available for study. The order of the families as given below has no phylogenetic
significance and is only thought to group somewhat similar forms together. For a
phylogenetic analysis of these families see Petersen (1990). Molecular phylogenies of
some families of Capitata can be found in Collins et al. (2005a, and b).

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 by 16S
DNA sequence information. The methods to obtain 16S DNA sequences are described
in Schuchert (2005). All sequences have been submitted to the EMBL database under
the accession numbers: AM088481, AM088482, AM088483, AM088484, AM088485.
The origin and identity of the material used to obtain 16S sequence data are given for
each species in the section "Material examined". Some sequences have been deter-
mined by other laboratories using material described here (accession numbers
AY920796, 18S AY920758, AY787879, AY920762, AY512539). Morphological
characters of the Cladonematidae were used for a phylogenetic analysis using the
parsimony criterion and the program PAUP* (Swofford, 2001). Bootstrap replicates
were performed 1000 times. Only a subset of all Cladonematidae was used. The
members of the genus Cladonema are all very similar and well represented by C. ra-
diatum. For Staurocladia, only those species with known life cycle were used. As out-
group taxa, three members of the Corynidae were used (comp. Schuchert, 2001b).
Table 1 gives the species names and the characters used and their states.

ABBREVIATIONS

BMNH The Natural History Museum, London, England

MHNG Muséum d’histoire naturelle de Genève, Switzerland

ICZN International Code of Zoological Nomenclature

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

CI Consistency index
RI Retention Index
HI Homoplasy Index

TAXONOMIC PART
FAMILY ACAULIDAE FRASER, 1924

DIAGNOSIS: Solitary hydroids, body divided into basal and distal part. Basal part
conical to cylindrical, covered or not by gelatinous perisarc which may form anchoring

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 327

filaments. Upper part with few indistinctly capitate tentacles or many capitate
tentacles, lowest whorl of capitate tentacles may transform into thick filiform tentacles.
Gonophores fixed sporosacs in the lower or middle part of the hydranth. Sometimes
asexual reproduction by transverse fission.

REMARKS: The name Acaulidae was proposed by Fraser (1924) and this spelling
was used by all subsequent authors. According to the ICZN [article 29.3.1.1] there is
thus no need to change it to the formally correct Acaulididae.

According to Bouillon (1985), this family contains two genera: Acaulis
Stimpson, 1854 and Acauloides Bouillon, 1965. Petersen (1990) also included the
genus Boreohydra Westblad, 1937 in the Acaulidae, while Bouillon (1985) kept
Boreohydra in a separate family Boreohydridae, this together with the genus Proto-
hydra. A third genus was later added to the Acaulidae by Thomas et al. (1995):
Cryptohydra. The discovery of Cryptohydra narrowed the gap of the Acaulidae to the
genus Protohydra — and thus Boreohydridae — to a degree that any subdivision now
appears entirely arbitrary. The proximity of both families is further emphasized if the
nematocyst warts on the body of Boreohydra are regarded as reduced capitate tentacles
(Calder, 1974). It is thus tempting to fuse both families, but this should only happen in
the framework of a broader analysis and discussion and the phylogenetic relationships.
The Acaulidae and Boreohydridae have also many similarities with the Candelabridae
and Tricyclusidae, and the demarcations are equally arbitrary. Because a phylogenetic
analysis based on morphological characters alone appears not so promising (see
Petersen, 1990), it is therefore preferable to wait with a revision until a thorough
molecular analysis provides the necessary robust phylogeny. Meanwhile, the classifi-
cation according to Bouillon (1985) is here used in order to maintain nomenclatural
stability.

Through monotypy, the genus Acauloides Bouillon, 1965 is unfortunately based
on Acauloides ammisatum, an insufficiently known species. Acauloides is thus essen-
tially distinguished from Acaulis through the absence of filiform tentacles. The filiform
tentacles of Acaulis are formed relatively late in development through the transfor-
mation of capitate tentacles (Berrill, 1952). Using this ontogenetic argument and also
outgroup comparisons to the Tricyclusidae and Candelabridae, it is evident that filiform
tentacles of Acaulis are apomorphic. The absence of filiform tentacles in the genus
Acauloides is thus a plesiomorphic trait and unsuitable to reveal a monophyletic group.
The genus Acauloides could nevertheless be valid, as in Acauloides ilonae the
gonophores develop in the upper axils of the tentacles, which is clearly an apomorphy
for this genus. Because Acauloides ilonae is perhaps a synonym of A. ammisatum, it is
thus recommendable to continue to use the genus Acauloides in order to maintain
nomenclatural stability.

KEY TO GENERA:

la hydranth small, very elongated, all tentacles indistinctly capitate

PITTI E CIO CIT IAA CI DONO Cryptohydra (not in European fauna)
lb Majority Omentacles distincthy capitate wey RS ON IE 2
2a mature hydranth with a whorl of thick filiform tentacles ................... Acaulis

2b allMentacles capitate "PIRATES REN OMR SIE Acauloides

328 P. SCHUCHERT

Genus Acaulis Stimpson, 1854

TYPE SPECIES: Acaulis primarius Stimpson, 1854.

SYNONYMS: Blastothela Verrill, 1878: 374; ? Myriocnida Stechow, 1909 (invalid nomen
nudum).

DIAGNOSIS: Hydroid connected to substrate by modified hydrocaulus secreting
a gelatinous sheath or forming anchoring filaments; hydranth fusiform; capitate tenta-
cles in one oral whorl and more scattered below, solid, with chordoid gastrodermis; be-
low capitate tentacles one whorl of filiform, long, stout, aboral tentacles developing by
transformation of capitate tentacles; gonophores fixed, carried singly or in clusters on
short pedicels.

REMARKS: Stechow (1909) introduced the genus name Myriocnida for a hydroid
described in Fewkes (1890) as Acaulis (without specific epithet). Because it was not
based on a nominal species, the genus is not valid. It remains unclear to which species
Fewkes' material belonged (he described it from notes made by someone else). It could
have been be either Acaulis or more likely an incomplete Candelabrum species.

Acaulis primarius Stimpson, 1854 Fig. 1

Acaulis primarius Stimpson, 1854: 10, pl. 1 fig. 4; Allman, 1872: 378; Sars, 1874: 123, pl. 5 figs
14-20; Will, 1913: 57, pl. 26; Scheuring, 1922: 167, pl. 5 fig. 1; Berrill, 1952: 17, fig. 6;
Rees, 1957: 466, fig. 13; Bouillon, 1971: 342, pl. 4; ? Verwoort, 1985: 269, plate 1;
Schuchert, 2001a: 35, fig. 22A-D.

Acaulis primaris — Naumov, 1969: 243, fig. 112 [incorrect subsequent spelling].

MATERIAL EXAMINED: Material from Iceland and Greenland as given in Schuchert
(2001a).

DIAGNOSIS: Acaulidae with filiform tentacles in mature animals, sporosacs not
in axils of tentacles, base without attachment filaments.

DESCRIPTION: Solitary hydroids, body thickly fusiform to cylindrical, divided
into basal part (approx. 1/5 of height, variable) and upper part. Both parts separated by
one whorl of five to eight thick tapering tentacles covered evenly with nematocysts.
Basal part conical to cylindrical, surrounded by thick jelly covered by adhering
detritus. Upper part of hydranth roughly cylindrical with up to 50 scattered capitate
tentacles; hypostome rounded. Gonophores on lower part of region with capitate
tentacles but not associated with the tentacles. Gonophores sessile sporosacs with
spadix but without radial or ring canals. Nematocysts: stenoteles, microbasic
heteronemes (euryteles or mastigophores), and desmonemes.

DIMENSIONS: Mature polyps are 5 to 10 mm in height, extended up to 2 cm
(Westblad, 1947), gonophores ca. 0.5-0.6 mm. Nematocysts: stenoteles, (18-21)x(14-
17) mm; heteronemes, (16-20)x(7-10.5) mm; desmonemes, (12-15.5)x(8.5-10.5) mm.

DEVELOPMENT: Young hydranths have no filiform tentacles, only capitate ones.
During growth, the lowest capitate tentacles become elongated and thicker, and the
nematocyte free regions of the tentacles are subsequently covered by nematocytes (see
Fig. 1A-B) (Berrill, 1952). The development of the gonophores can begin before the
filiform tentacles have completed their transformation.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 329

Fic. 1

Acaulis primarius Stimpson, 1854; after Schuchert (2001a), Icelandic material. A) Juvenile,
scale bar 0.5 mm, also valid for sections B-C. B) Intermediate sized specimen with beginning
transformation of lowest tentacles. C) Mature female, filiform tentacles fully developed. D)

Nematocysts, as seen in preserved material: stenotele, microbasic heteroneme, desmoneme,
scale bar 10 um.

ADDITIONAL DATA: Westblad (1947) describes and depicts both the gonophore
development and gametogenesis. Westblad (1947) also depicts the nematocysts and de-
scribes their distribution.

330 P. SCHUCHERT

BioLoGY: Acaulis primarius lives usually partially embedded in sediment
bottoms at depths of down to 350 m, usually not above 20 m depth (Berrill, 1952;
Schuchert, 2001a).

DISTRIBUTION: Northern Atlantic, Arctic to boreal regions; New England, New
Brunswick, north-west coast of Norway, Barents Sea, White Sea (Scheuring, 1922;
Fraser, 1944; Naumov, 1969), North Sea, Sweden (Jäderholm, 1909; Westblad, 1937;
Bouillon et al., 1995), Baltic Sea (Will, 1913; Schônborn et al., 1993), Iceland
(Schuchert, 2001a), western Greenland (Schuchert, 2001a). Perhaps also Bay of Biscay
(Vervoort, 1985, identification uncertain). Type locality: Grand Manan, Bay of Fundy,
Canada.

REMARKS: Stimpson (1854) described Acaulis primarius based on two sets of
animals collected at different times. The first animal he found and depicted corres-
ponds exactly to our current concept of Acaulis primarius. The second set he found
later, attached to red-algae (it is unclear in what depth, he states 5-15 f., which could
be feet or fathoms, both interpretations can be found in the literature, but the latter
being more probable). In the second set of specimens the tentacles were much longer,
the hydranths resembled a corynid and they reportedly produced medusae. Allman
(1872), Will (1913), and Fraser (1924) noticed this difference and assumed that the
second set of Stimpson’s material belonged to another species than Acaulis primarius.
Allman (1872) restricted the scope of Acaulis to the animal depicted in Stimpson
(1854), thus he de facto selected a lectotype and Acaulis primarius is thus well defined.

Genus Acauloides Bouillon, 1965

TYPE SPECIES: Acauloides ammisatum Bouillon, 1965, by monotypy.
SYNONYMS: ? Psammocoryne Monniot, 1962 (invalid nomen nudum).

DIAGNOSIS: Hydroid attached to substrate by modified hydrocaulus, secreting a
gelatinous sheath or mucous film; capitate tentacles in one oral whorl and more
scattered below, solid, with chordoid gastrodermis; without filiform tentacles;
gonophores in upper axils of tentacles, asexual reproduction through transverse fission.

Acauloides ammisatum Bouillon, 1965 Fig. 2

? Psammocoryne. — Monniot, 1962: 274, fig. 14b [nomen nudum].
Acauloides ammisatum Bouillon, 1965: 54; Bouillon, 1971: 335, figs IV8-11, V-VI; Bouillon
et al., 2004: 86, fig. 47H.

MATERIAL EXAMINED: None, type material could not be located.

Diacnosis: Acaulidae with capitate tentacles only, adults 0.6-2 mm, 10-25
tentacles, peduncle without gelatinous tube, only mucous secretion at its end for
attachment to sand grains.

DESCRIPTION: Solitary hydroids, thick fusiform shape, short peduncle of about
1/3 of total height, peduncle without gelatinous tube but at end mucous secretion by
which the animal attaches itself to sand grains. Tentacles all capitate, 10-25, one oral
whorl of 4-6 short tentacles, remaining scattered below on body, those in middle of
body longest, others very short to almost lacking a stalk. Nematocysts: stenoteles;

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 331

FIG. 2

Acauloides ammisatum Bouillon, 1965; modified after Bouillon (1971), no scale given. A)
Polyp. B) Stenotele, larger mastigophore, smaller mastigophore, discharged mastigophore.

desmonemes; holotrichous microbasic mastigophores, when discharged shaft longer
than capsule. Vegetative and sexual reproduction not known.

DIMENSIONS: Polyps 0.6-2 mm. Nematocysts: stenoteles 17x14 um, desmo-
nemes 6x10 um, microbasic mastigophores (16-18)x(6-8)um.

ADDITIONAL DATA: Bouillon (1971) provides further details on the histology and
cytology of this species.

DISTRIBUTION: Roscoff, Brittany; ? Banyuls-sur-Mer (Mediterranean). Type
locality: Roscoff, English Channel.

REMARKS: Acauloides ammisatum Bouillon, 1965 strongly resembles the vege-
tative reproduction stage of A. ilonae (Brinckmann-Voss, 1966) and the two are
possibly conspecific. There are some minor size differences, with A. ammisatum being
smaller and having slightly larger nematocysts. Another formal difference is the
absence of a gelatinous tube in A. ammisatum, for which only a mucous cover has been
described. However, such a mucus cover might be only the initial stage of a gelatinous
tube, a type of cover which is only seen in fully grown animals. There is also a possible
ecological difference: while A. ammisatum occurs on sand, A. ilonae has been found
on mud only. More data — especially on mature animals from Roscoff — are needed
before a final more definite conclusion is possible. Meanwhile, both nominal species
should be kept separate.

Bouillon (1971) also attributed to this species some animals from Banyuls
(Mediterranean) depicted in Monniot (1962). Monniot found her animal in coarse sand
and identified it as Psammocoryne. This name is not a valid genus as it was not asso-

332 P. SCHUCHERT

ciated with a valid nominal species. Furthermore, Monniot's hydroid could easily also
be referred to A. ilonae and it is therefore also somewhat unclear whether A. ammisa-
tum also occurs in the Mediterranean (this in case that they are not conspecific).

Acauloides ilonae (Brinckmann-Voss, 1966) Fig. 3
Acaulis ilonae Brinckmann-Voss, 1966: 292, figs 1-10; Brinckmann-Voss, 1970: 44, text-fig. 51,
pl. 1 figs 4-6.

Acauloides ilonae — Bouillon et al., 2004: 86, fig. 471.

MATERIAL EXAMINED: Syntypes (labelled holotype), BMNH 1963.12.9.1, as Acaulis ilon-
ae, two specimens, both broken. - BMNH 1974.11.21, as Acaulis ilonae, Naples, 20.10.1960-
29.11.1961, leg. Brinckmann-Voss, one female, pedicel broken off.

DIAGNOSIS: Acaulidae with capitate tentacles only, sexually reproducing adults
5-10 mm, about 2 mm in phase of vegetative reproduction; sporosacs in upper axils of
tentacles, peduncle of adult hydranths in gelatinous sheath.

DESCRIPTION (after Brinckmann-Voss, 1966; 1970; and own observations):
Mature hydroid spindle shaped, divided into two parts of roughly the same length: an
upper tentacle-bearing part and a basal part without tentacles called peduncle (or
"foot"). Peduncle tapering, surrounded by a gelatinous sheath which adheres to mud
particles or cultivation vessels, gastrodermis highly vacuolated. Tentacles all capitate,
solid, up to 60, one oral whorl of four to five tentacles and a basal whorl comprising
the same number of tentacles, between them a number of scattered tentacles, often in
groups of two to three around a sporosac, tentacle length unequal, longest ones more
proximal, diameter of capitula also variable. Gonophores are sporosacs without radial
canals, developing always in upper axils of tentacles in middle region of tentaculate
part, oldest sporosacs in middle, younger ones proximal and distal to them. New ten-
tacles often formed close to the gonophores. Colour of the animals pink to orange, with
a number of white dots on surface, very old specimens have a brownish colour, nema-
tocysts: stenoteles, microbasic euryteles and desmonemes. Fertilized eggs developing
a capsule and going through an encysted resting stage.

Polyps in vegetative phase of reproduction small, with 29-35 tentacles, attached
to substratum, multiplying by binary fission in region of tentacles.

DIMENSIONS: Sexually reproducing animals up to 10 mm, polyps in phase of
vegetative multiplication up to 2 mm. Diameter of eggs 0.15 mm. Nematocysts:
desmonemes (8.4-12.6)x(7.6-9.2)um; stenoteles (18.5-22.7)x(16.0-19.3)um; microba-
sic euryteles (11.8-14.3)x(3.4-7.6)um (Brinckmann-Voss, 1966).

DISTRIBUTION: Naples, questionably also Roscoff, Brittany, although the latter
records may refer to A. ammisatum. Type locality: Gulf of Pozzuoli, Naples, Italy, 45 m.

BIOLoGy (after Brinckmann-Voss, 1966; 1970): Acauloides ilonae is able to
creep sluggishly on the substrate. The animals are gonochoristic; female and male hy-
droids continue to spawn their gametes several times for more than a year, the gametes
differentiate continuously. Eggs are usually shed during the night and early morning
hours, fertilisation takes place in the seawater. The embryo does not swim and attaches
itself to the ground and transforms into a flattened body which is covered with thin,
transparent cuticula. The cyst remains attached for at least one month, after which a

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 333

FIG. 3

Acauloides ilonae (Brinckmann-Voss, 1966), modified after Brinckmann-Voss (1966), scale bar
1 mm. A) Sexually reproducing polyp. B) Vegetatively reproducing polyp just after fission, with
aboral and oral parts.

small polyp without tentacles hatches very slowly (it takes 2-3 days). Hydroids hatch
from cysts only if kept in water of 13°C, at 20°C the hatching rate is poor. The young
polyp grows to a size of about 2 mm and then starts the vegetative reproduction phase
by transverse fission. At 20°C divisions take place every 10-45 days and if kept at 20°
the fission continues, the polyps do not exceed 2-3 mm, and they do not develop
gonophores. Polyps that had undergone divisions and are brought to 13° do not divide
anymore, they begin to grow in length, get more tentacles and develop gonophores in
the axils of the tentacles, thus starting the sexual phase. Individuals that had hatched
from cysts and are cultured at 13° undergo one or two rounds of divisions before they
develop gonophores. Sexual maturity is not reached without having first undergone
asexual division.

The hydroid was found on surface of mud in depths of 20 to 80 m. It occurred
in abundance around Naples from November to April. It was not found from July to
October. It is thus a winter species.

REMARKS: Brinckmann-Voss (1966) described her material based on animals
from Naples, but also mentioned that Bertil Swedmark had found it at Roscoff too (via
personal communication by J. Rees). Because it appears that the latter material was

334 P. SCHUCHERT

more likely Acauloides ammisatum (see remarks in Bouillon, 1971: 335), the occur-
rence of A. ilonae outside the Mediterranean is uncertain (if the two indeed prove to be
distinct species, see remarks under A. ammisatum).

FAMILY BOREOHYDRIDAE WESTBLAD, 1947
TYPE GENUS: Boreohydra Westblad, 1937.

DIAGNOSIS: Hydroids solitary, small, with one whorl of reduced tentacles,
capitate or not, located in the oral or median part of body; perisarc covering of base
filmy or absent; gametes in body wall.

REMARKS: Representing likely a simplified form, the affinities of the genus
Boreohydra with its sole species B. simplex remains controversial. Westblad (1947)
considered it related to the Tubulariidae, but sufficiently distinct to deserve being
placed in a separate family. Rees (1957) could not agree and placed it in the
Corymorphidae. Calder (1974) thought that if the nematocyst clusters on the body
should prove to be nothing but reduced capitate tentacles, then the genus should belong
to the Candelabridae. Bouillon (1985) considered the family Boreohydridae as valid
and added also the genus Psammohydra to it, this because of the resemblance of their
cnidome. Petersen (1990) placed Boreohydra among the Acaulidae, but gives no
arguments for this and he does not discuss Psammohydra. As already stated for the
family Acaulidae, the problem is not resolvable by traditional approaches and, pending
a molecular analysis, the family Boreohydridae is here retained for the time being.

KEY TO THE GENERA:

la hydranth with oral capitate tentacles and nematocyst buttons on body ..... Boreohydra
lb filiform tentacles in middle of body: hypostome proboscis-like........ Psammohydra

Genus Boreohydra Westblad, 1937

TYPE SPECIES: Boreohydra simplex Westblad, 1937.

DIAGNOSIS: Solitary hydroids of small size, living buried in the sediment; caulus
covered by filmy perisarc; one whorl of capitate tentacles near mouth and numerous
scattered nematocyst clusters on hydranth body. Gametes in epidermis at junction of
hydranth body and caulus; asexual reproduction by transverse fission.

REMARKS: This is currently a monotypic genus.

Boreohydra simplex Westblad, 1937 Fig. 4

Boreohydra simplex Westblad, 1937: 1, figs 1-4; Westblad, 1947: 1-13, figs 1-4, pls 1-3; Nyholm,
1951b: 531, text fig., pl. 1 figs 1-4; Westblad, 1953: 351, figs 1-2; Prévot, 1959: 97,
pl. 1 fig. 2; Calder, 1974: 1666, fig. 1; Bozhenova et al., 1989: 11, fig.; Petersen, 1990:
148; Schuchert, 2001a: 36, fig. 23A-B; Bouillon er al., 2004: 86, fig. 47L.

MATERIAL EXAMINED: See Schuchert (200142).
DIAGNOSIS: Small, solitary, mud-dwelling hydroids with three to four capitate
oral tentacles and scattered wart-like nematocyst clusters on hydranth body.

DESCRIPTION: Solitary hydroid, composed of hydranth body and conical
peduncle. Hydranth spindle-shaped, hypostome short and rounded, surrounded by

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 335

Fic. 4

Boreohydra simplex Westblad, 1937; Northern Greenland, Pearyland, after Schuchert (2001a).
A) polyp, scale bar 0.2 mm. B) stenotele, desmoneme, isorhiza, scale bar 10 um

three to four short, stubby tentacles, only slightly capitate. On hydranth body 50-60
scattered nematocyst clusters, variable in size, 3-15 capsules per cluster, clusters slight-
ly elevated and wart-like. Hydranth body at base tapering and turning into peduncle,
peduncle conical, constantly tapering into fine tip at its end, covered by wrinkled
perisarc. Gastrodermis with three or four longitudinal folds. Nematocysts: stenoteles;
desmonemes, discharged with four coils; isorhizas.

DIMENSIONS: Hydranth body height 0.8-1.5 mm, diameter about 0.3 mm;
peduncle 0.4-1 mm long. Nematocysts: stenoteles (17-19)x(16)um; desmonemes,
(16-17)x(9-11)um; isorhizas (17-20)x(4.5-5.5)um.

BIOLOGY: Fairly common in mud bottoms characterized by the 'Brissopsis
lyrifera - Amphiura chiajei' community (Petersen, 1913; Jones, 1950) at depths of
around 40 m (Westblad, 1953). The depth range varies from a few metres to more than
600 m (Christiansen, 1972). The animal is able to move quite rapidly on the surface of
mud (Hult, 1941). The latter author also described how it burrows into the mud. The
oral tentacles are instrumental for this process. The food seems to be mainly composed
of small nematodes (Westblad, 1947).

ADDITIONAL DATA: (Westblad, 1947; Nyholm, 1951b) The polyp multiplies by
transverse fission and produces gonophore-like outgrowths without germ-cells. Eggs
were found in the epidermis at the junction of the body and peduncle. Westblad (1947)
gives further details on the variability: of 60 animals, 47 had three tentacles, the others
four. The number of tentacles and the number of gastrodermal folds is not closely
correlated. Prévot (1959) depicts a longitudinal section. Bozhenova et al., (1989)
depict the nematocysts and provide measurements.

336 P. SCHUCHERT

DISTRIBUTION: Bipolar, northern and southern Atlantic Ocean in temperate to
Arctic waters, probably quite frequent, but often overlooked. It was particularly often
recorded along the Atlantic coast of Scandinavia. The southern limit for Europe is the
English Channel. Recorded from the White Sea, Norway to Sweden, Great Britain,
Iceland, North Greenland, North-Eastern Canada, North Greenland, South Georgia
(Hult, 1941; Westblad, 1953; Christiansen, 1972; Calder, 1974; Bozhenova et al.,
1989; Schuchert, 2001a). Type locality: Tromsg and Ramfjord, Norway.

REMARKS: Westblad (1947) supplemented the first description by further
anatomical and ecological details. He observed buds that he interpreted as gonophores,
however he could not find germ cells in them. Later, Nyholm (1951) observed eggs in
the epidermis and concluded that the germ cells of this species are not collected in
sporosacs, but remain in the epidermis like in Hydra. Petersen (1990) interpreted the
gonophore-like outgrowths observed by Westblad (1947) as incipient polyp buds.
More work is needed to reveal the nature of these buds.

Genus Psammohydra Schulz, 1950b
TYPE SPECIES: Psammohydra nana Schulz, 1950b.

DIAGNOSIS: Solitary hydroids of very small size, living attached to sand grains;
one whorl of filiform tentacles in middle region of hydranth; hypostome proboscis-like,
with terminal swelling. Asexual reproduction through fission.

REMARKS: This is currently a monotypic genus. The sexual reproduction is in-
sufficiently known.

Psammohydra nanna Schulz, 1950 Fig. 5

Psammohydra nanna Schulz, 1950b: 122, figs 1-9; Riedl, 1970: 153, pl. 43; Clausen, 1971: 2,
fig. 1; Clausen & von Salvini-Plawen, 1986: 34, fig. 3; Thiel, 1988: 267, fig. 19.1d;
Bouillon ef al., 2004: 87, fig. 48A.

MATERIAL EXAMINED: None, the type material could not be located.

DIAGNOSIS: Very small solitary hydroid living attached to sand grains, body skit-
tle- to spindle shaped, one whorl of filiform tentacles in about middle of body.

DESCRIPTION: Tiny solitary hydroids, living attached to sand-grains; body shape
and size very variable, usually skittle- or spindle-shaped, with flat base when attached,
slightly above middle a single whorl of four (rarely 3-5) tentacles; hydranth body
above tentacles proboscis-like, with terminal swelling. Proboscis movable and exten-
sible, mouth terminal but invisible when not used. Tentacles short, straight, contractile,
filiform, nematocysts evenly distributed, gastrodermis chordoid. Sexual reproduction
insufficiently known, reportedly only one egg is produced. Colour: sand-grey. Nemato-
cysts: stenoteles, desmonemes, atrichous isorhizas.

DIMENSIONS: Total body size 0.28-0.4 mm, but very variable, can contract or
expand its body about two times. Nematocysts: stenoteles (6-7)x(5-6)um, isorhizas
(5.5-6.8)x(2.3-3)um; desmonemes (3.5-5.5)x(2.3-3.5)um (Schulz, 1950b).

BIOLOGY: Vegetative reproduction takes place by transverse fission. The animal
can move by creeping like a freshwater Hydra. It is a member of the meiofauna and

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 337

FIG. 5

Psammohydra nanna Schulz, 1950, redrawn from Schulz (1950b), no scale given. A-C) Animals
on sand-grains in various states of contraction. D) Vegetative reproduction by transverse fission.

occurs in fine sand in shallow waters of a few metres. Schulz (1950b) characterizes the
community where it was found as "Turbanella hyalina" community (Remane, 1933).
It can tolerate reduced salinity down to 8%o (Schönborn et al., 1993).

ADDITIONAL DATA: Of 46 examined animals, 39 had four tentacles, four had
three tentacles, and three five (Schulz, 1950b). Schulz (1950b) also describes the
histology of this animal. The gastrodermal epithelium is high in the lower part of the
animals, almost obliterating the lumen, but it is low above the tentacles and thus leaves
a distinct cavity. The proboscis contains only stenoteles and isorhizas. The desmo-
nemes are confined to the tentacles. The proboscis is very extensible and can bend to
all sides. It acts like a central tentacle and it is used for locomotion and food capture.
Swedmark (1959) mentions without further comment that Psammohydra produces a
single egg only.

DISTRIBUTION: Western Baltic Sea (Schulz, 1950b), English Channel (Teissier,
1965); western Mediterranean (Swedmark, 1956), Adriatic Sea (Salvini Plawen, 1966).

REMARKS: The taxonomic position of this simplified animal is unclear (see also
remarks for family Acaulidae and Boreohydridae).

FAMILY PROTOHYDRIDAE Allman, 1888
TYPE GENUS: Protohydra Greeff, 1869.

DIAGNOsIs: Solitary, small, elongated hydroids usually living in brackish-
waters; without tentacles and gonophores, pedal disc formed by epidermal cells only;

338 P. SCHUCHERT

gametes bulging into gastric cavity, nematocysts: stenoteles and isorhizas, evenly dis-
tributed over body.

Genus Protohydra Greeff, 1869
TYPE SPECIES: Protohydra leuckarti Greeff, 1869.
DiaGNOSIS: With the characters of the family.

REMARKS: Due to their — likely secondary — very simple morphology, the Proto-
hydridae are difficult to classify. Petersen (1990) treated the family as Capitata incerta
sedis. The cnidome is quite interesting as it lacks desmonemes. This could, however be
due to the absence of tentacles. In Psammohydra nanna, a similarly reduced form, the
desmonemes are associated with the tentacles. The family currently comprises two
species: Protohydra leuckarti Greeff, 1869 and Protohydra psamathe Omer-Cooper,
1964. Protohydra caulleryi Dawydoff, 1930, characterized by budding or frustules, is
likely a polyp belonging to the Oliandiasidae (Weill, 1935; Schulz, 1952).

Protohydra leuckarti Greeff, 1869 Fig. 6

Protohydra leuckarti Greeff, 1869: 37, pls 4-5; Luther, 1923: 1, figs 1-11; Koller, 1927: 97, figs
1-2; Westblad, 1930a: 1-4; Westblad, 1930b: 1-13, figs 1-3; Weill, 1934: 448; Weill,
1935: 83, fig. 5; Westblad, 1935: 152, figs 1-4; Vervoort, 1946: 47, fig. 1; Schulz, 1950a:
53; Nyholm, 1951a: 529, pl. 1; Naumov, 1969: 593, fig. 439; Clausen, 1971: 1.

MATERIAL EXAMINED: ZMUV, Denmark, Mariagerfjord, Ajstrupbugt, 0.5 m, 31 July
1955, 3 specimens, det. Kramp. — IRSN, two samples from Roscoff, collected by J. Bouillon in
May 1964 (many specimens) and August 1961 (few specimens).

DIAGNOSIS: Solitary, small, elongated hydroids without tentacles and gono-
phores, gametes bulging into gastric cavity; vegetative reproduction usually by trans-
verse fission or rarely by lateral buds; nematocysts: stenoteles and isorhizas.

DESCRIPTION: Solitary, brackish-water hydroids, usually elongate spindle- to
club-shaped when relaxed, spherical when contracted. Tentacles absent, at aboral end
a small epidermal attachment disc for temporary attachment, no perisarc, nematocysts
evenly distributed in epidermis, not concentrated around mouth. Vegetative repro-
duction by transverse fission, rarely by buds (Schulz, 1952). Gonophores absent,
gametes differentiate from epidermal cells and proliferate into gastric cavity where the
gonad remains attached along one side of the body wall, gonochoristic, females pro-
duce one egg only which is expelled by perforation of the body wall. Nematocysts
(Luther, 1923; Weill, 1934; Schulz, 1950b): stenoteles and basitrichous isorhizas,
desmonemes absent.

DIMENSIONS: Adults maximally contracted to a sphere 0.4 mm, expanded
2-3 mm (Greeff, 1869). Eggs ca. 0.25 x 0.14 mm (Westblad, 1930b). Nematocysts
(Luther, 1923): stenoteles 12-17 um long, isorhizas (7)x(3-3.5)um,

ADDITIONAL DATA: There are pigment granules of variable size in the gastro-
dermis. The colour of the animals depends on the food items and is either colourless or
various shades of red (Greeff, 1869; Madsen, 1939). Westblad (1935) showed that the
germ cells originate from epidermal cells that proliferate into the gastric cavity. The
animals are gonochoristic and females produce a single egg. The egg is expelled by a

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 339

Fic. 6

Protohydra leuckarti Greeff, 1869. A-D) Animal in various states of contraction, oral end up-
wards, scale bar approximately 0.2 mm, redrawn from Greeff (1869). E) Schematic longitudinal
section of male animal, the gonads (stippled) bulge into the stomach lumen, scale bar 0.1 mm,
redrawn from Westblad (1935).

rupture of the body wall and the animal dies afterwards. Westblad (1935) also made
some observations that hint at a possible copulation. This is rather exceptional for
hydrozoans and needs reconfirmation. Schulz (1952) observed that besides the usual
transverse fission, rarely some animals produce lateral polyp buds. Further biological
data are also provided by Muus (1966) and Wehling (1978). Some information on the
histology — e. g. the body wall, the glue cells, and gametogenesis — are provided by
Luther (1923) and Westblad (1935).

BIOLOGY: A coastal, euryhaline species occurring in brackish waters of wave
protected habitats, often in closed bays, estuaries, and lagoons. It is absent from
agitated waters. The depth range is from a few cm to about 20 m and the reported
salinity range 0.38-30%o (Westblad, 1930a; Madsen, 1939; Schulz, 1950a; Schönborn
et al., 1993, Barnes, 1994). Various bottom types have been reported, but there seems
to be a preference for fine sand or mud with much detritus. They can reach very high
densities, reaching from 1000 individuals per square metre in winter to 50'000 in

340 P. SCHUCHERT

summer, or even exceptionally 200'000/m? (Heip, 1971). Rarely it also occurs on
algae. It can be found all year round, but reaches maximum densities in summer when
it also gets sexually mature. Protohydra leuckarti is an important predator in its biotope
and it regulates the density of other animals (Heip, 1971; Heip & Smol, 1976). The
animal is able to creep and to burrow. It lives on a variety of food items, especially
nematodes, copepods, ostracods, chironomid larvae (Schulz, 1950a). It is preyed upon
by e. g. nudibranchs (Evertsen ef al., 2004).

DISTRIBUTION: Circumglobal in temperate brackish waters of the northern hemi-
sphere. The northern limit in Europe is southern Norway (Oslofjord: Christiansen,
1972) and southern Finland (Helsinki: Schneider, 1927). It has been reported from the
Atlantic coast of Sweden (Westblad, 1930a); the Baltic Sea (Schneider, 1927;
Westblad, 1935; Koller, 1927; Nyholm, 1951; Schulz, 1950a); Denmark (Madsen,
1939; Muus, 1966; Rasmussen, 1973); German part of the North Sea (Schulz, 1950a);
Holland (Boaden, 1976); Belgium (Greeff, 1869; Heip, 1971); Southern England
(Baker, 1913; Hickson, 1920); Brittany (Teissier, 1965); Bay of Biscay (Nyholm,
1951a); in the Mediterranean it was found in brackish water lakes of southern France
(Nyholm, 1951a). It has also been found in the Black Sea (Valkanov, 1947; Marcoci,
1956). Outside Europe it has been recorded on the east coast of North America
(Ruebush, 1939), in the north-eastern Pacific (Wieser, 1958), the north-western Pacific
(Naumov, 1969), and the Aral Sea (Maier, 1974). Type locality: Ostende, in mud
among oyster cultures.

FAMILY CANDELABRIDAE STECHOW, 1921
TYPE GENUS: Candelabrum de Blainville, 1830: 284.
SYNONYMS: Myriothelidae Hincks, 1868; Symplectaneidae Fraser, 1941.

DIAGNOSIS: Large, worm-like hydroids, solitary or forming small pseudo-
colonies through connected aggregates, attached to substrate but without stolons;
hydranth elongated, cylindrical to club-shaped; with numerous scattered, hollow or
parenchymatic capitate tentacles, tentacles simple or compound, if compound then
with adnate basal parts. Hydrocaulus with or without perisarc, with tentacle- or root-
like attachment-processes. Gonophores fixed sporosacs, developing either directly on
hydranth body or on club-shaped blastostyles.

REMARKS: Stechow (1921: 248) pointed out that the genus Candelabrum de
Blainville, 1830 has priority over Myriothela Sars, 1850. Consequently, he then also
changed the family name from Myriothelidae Hincks, 1868 to Candelabridae Stechow,
1921. According to the ICZN path ed. 1999, 40.2] such a name change is valid if it was
made before 1961 and if the new name has been widely used. The name Candelabridae
has only recently come into usage (e. g. Bouillon, 1985; Schuchert, 1996; Hewitt &
Goddard, 2001), while other used Myriothelidae (e. g. Calder, 1972; Millard, 1975;
Petersen, 1990). I therefore suggest that for the sake of nomenclatural stability,
Candelabridae should be used from now on. This name matches the genus name and it
is also used in several large electronic databases. If in future Candelabridae should not
become the prevailing name, then a ruling of the International Commission on
Zoological Nomenclature must be requested.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 341

The family currently comprises the following genera: Candelabrum de
Blainville, 1830; Monocoryne Broch, 1910, and Fabulosus Stepanjants, 1990.

KEY TO THE GENERA:

la Capitateytentaelestsimpler RR aes A IS IR E ERO SEE 2
lb capitate, tentacles COmpound tyre yas. ISEE RA Monocoryne
2a Sporosacs/bormeonyblastostyles FE as EEE ie sete cue ER Candelabrum
2b sporosacs borne singly on hydranth body......... Fabulosus (not in European fauna)

Genus Candelabrum de Blainville, 1830

TYPE SPECIES: Lucernaria phrygia Fabricius, 1780 = C. phrygium (Fabricius, 1780).

SYNONYMS (after Segonzac & Vervoort, 1995): Arum Vigurs, 1850; Myriothela M. Sars,
1850; Spadix Gosse, 1853a; Acandela Stechow, 1920.

DraGNOSIS: Solitary hydroids or loose aggregates comprising few polyps that
may have a common perisarc base. Hydranth long, cylindrical or club-shaped, upper
part with numerous simple capitate tentacles; at base a foot region, with or without
perisarc sheath, with attachment processes that are either covered with perisarc or
naked and may have a perisarc disc at their end; gonophores are fixed sporosacs borne
on blastostyles developing in region between tentacles and foot. With or without
clasper tentacles that hold developing embryos. Development direct, leading to young
polyps without a planula stage.

REMARKS: Segonzac & Vervoort (1995) recently revised the genus Cande-
labrum, outlining its synonymy and taxonomic history.

KEY TO THE NORTH-ATLANTIC CANDELABRUM SPECIES:

la Sporosacsawithout nematocyst buttons. it EME RE oan. ae eee 2
lb SpOLrosacsawithmematocystDUMONS ys TER I C. verrucosum
2a foot large, covered by perisarc sheath, hermaphroditie ee nn 3)
2b foot without perisarc sheath, without clasper tentacles, dioecious, deep water or

ATCUCHORME AE ARS ANT RTE EN EI LIL AURA Oman ET Le: QE TA C. phrygium
3a boreal shallow water form, with clasper tentacles holding developing embryos

HO ee ce Ps dec ce me s ane AO SA N. ISO E C. cocksii
3b deep water form, without clasper tentacles . .... C. serpentarii (not in European fauna)
Candelabrum cocksii (Cocks, 1854) Fig. 7

Arum Cocksii Vigurs, 1850: 90, nomen nudum.

Spadix purpurea Gosse, 1853a: 126; Cocks, 1853: 365.

Spadix cocksii — Gosse, 1853c: 386.

Arum cocksii Cocks, 1854: 34, pl. 3, fig. 7-12.

Myriothela phrygia — Hincks, 1868: 77, pl. 12 fig. 3; Allman, 1874: 317; Hardy, 1891: 505, pls
36-37; Hartlaub, 1916: 110, figs 38-39.

[not Candelabrum phrygium (Fabricius, 1780]

Myriothela — Allman, 1876: 549, pls 55-58 [only named Myriothela phrygia in plates].

Arum cocksi — Rees, 1957: 487, figs 37 & 39A-B; Prévot, 1959: 97, pl. 1 fig. 1.

Myriothela cocksi — G. O. Sars, 1874: 130; Bonnevie, 1899: 36; Billard, 1921: 12, fig. 1; Weill,
1934: 373; Manton, 1941: 143, figs la-b, 2.

Candelabrum cocksii — Segonzac & Vervoort, 1995: 37, fig. 2c-d, table 1.
MATERIAL EXAMINED: MHNG INVE 36299, France, Roscoff, near Ile Verte, 0 m,

17 September 2004, 3 specimens, fertile, examined alive, one blastostyle used for serial histo-

logical sections, confirming the presence of two male sporosacs and several female ones. —
MHNG INVE 29591, France, Roscoff, near Ile Verte, 0 m, 30 March 1998, 2 specimens,

342 P. SCHUCHERT

Fic. 7

Candelabrum cocksii (Cocks, 1854); A, after photographs of living animals, C-F, after preserved
material. A) Entire polyp, semi-extended, capitate tentacles contracted, scale bar approximately
5 mm. B) Blastostyle with sporosacs and capitula (stippled), scale bar 1 mm. C) Tip of blas-
tostyle with branched end, same scale as B. C) Embryo held by three clasper tentacles, scale bar
1 mm. E) Attached base of hydranth, perisarc dark, note two small perisarc discs in lighter
region, scale bar 3 mm. F) Nematocysts of mature polyp: large and small desmonemes, stenotele,
microbasic eurytele, scale bar 10 um.

attachment part lacking, fertile, 28S sequence of this material in GenBank under accession
number AY920796, 18S AY920758, 16S AY787879. - MHNG INVE 35726, France, Roscoff,
Tyaozon, 6 May 1910, coll. M. Bedot. - BMNH 1980.3.1.1, misidentified as Candelabrum phry-
gium, England, Devon, Salcombe, Castle Rock, 19 Feb 1980, R. L. Manuel, 3 well preserved
specimens, eggs held by claspers present, base in irregular perisarc.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 343

DiaGNOSIS: Candelabrum species with clasper tentacles that hold embryos, her-
maphroditic, base long and sheathed in perisarc envelope that is adnately attached to
the substratum.

DESCRIPTION: Solitary, relatively large, firmly attached, worm-like polyps,
divided into three regions: foot, blastostyle region in middle, and trunk.

Foot large, about 1/4 to 1/3 of whole animal, bent horizontally so that attach-
ment to substrate is adnate, basal region encased in firm, brown perisarc, distorted and
gnarled to fit irregularities of the substratum and provided with short, lobed or finger-
like projections with flat ends adhering to the solid substratum; the perisarc gradually
thins out distally; on thinner perisarc region there can be a few thick, sharply demar-
cated perisarc discs used for attachment, but without being associated with notable
protrusions of the hydranth wall; further distal some short finger-like protrusions of the
hydranth wall can occur.

Blastostyle region of about the same length as foot, provided with numerous
(>20-40), widely spaced blastostyles bearing the gonophores and with thin clasper
tentacles holding embryos. Blastostyles contractile, hollow and lumen in connection
with gastric cavity, stick- to club-shaped (swollen distally), sometimes branched in
distal region, distal region provided with nematocyst clusters, these in hemispherical
bumps or stubby capitate tentacles with short thick pedicels, quite irregular in
appearance; proximal 2/3 of blastostyle with irregularly scattered gonophores, different
developmental stages mixed. Gonophores are fixed sporosacs without radial canal
system (cryptomedusoid type); males and females occurring on same blastostyle, the
animals are thus simultanous hermaphrodites. Sporosacs spherical, white, female ones
larger than male ones, females initially with numerous small oogonia but ultimately
only one egg matures. Mature or fertilized eggs leave sporosac but are then grasped by
two to five clasper tentacles. Clasper tentacles thin, straight, of variable length, ori-
ginating in pairs or more at bases of blastostyles or also independently of them, without
nematocysts, terminal region sucker-like, enlarged, and attached to the envelope of
developing eggs.

Trunk region comprising about half of the animal, capable of great expansion
and strong contraction, elongate club-shaped with largest diameter following the
blastostyle region, evenly and entirely covered by hundreds of imbricate, indistinctly
capitate tentacles; capitate tentacles contractile, hollow, pedicel short in material taken
out of the sea (even when anaesthetized, but can extend considerably in undisturbed
specimens and become distinctly capitate), capitulum ovoid, diameter not much larger
than the contracted pedicel.

The foot and blastostyle parts are less contractile than the trunk. In animals
which have been preserved without relaxation, the trunk region is thus only as large or
smaller than either the foot and blastostyle region. The long foot is characteristic for
this species (comp. Figs 7 and 8-9).

Nematocysts of mature animals: two types of desmonemes, stenoteles, micro-
basic euryteles. In young polyps, there are also isorhizas (Weill, 1934); they may also
be found rarely in adults (own observations).

Colours: living animals entire body, blastostyles, and sporosacs white, capitula
of tentacles of trunk purple, perisarc dark amber-brown.

344 P. SCHUCHERT

DIMENSIONS: Mature body size variable and difficult to establish due to great
contractibility, ranging from a few cm to 12 cm, reportedly also more, usually 2-3 cm
when contracted; blastostyles up to 4 mm but contractile; capitula of trunk tentacles
diameter about 0.2 mm, expanded tentacles up to 2 mm long (Allman, 1874); clasper
tentacles 1-2 mm long; diameter of fertilized eggs held by claspers 0.7-0.8 mm, female
sporosacs reach same diameter; male sporosacs smaller, 0.30-0.42 mm (Segonzac &
Vervoort 1995). Nematocysts of mature animal (preserved material, see Fig. 7F): large
desmonemes (12-14)x(8.5-9.5)um; small desmonemes (7.5-9)x(5-6)um; stenoteles
(11-11.5)x(8-9)um; microbasic euryteles (18-20)x(5.5-6)um, ratio of everted shaft to
capsule length around 0.9; presumed isorhiza 13.5x5 um.

BroLocy: Along the coasts of Brittany, fertile animals have been documented
from January to September (Teissier, 1965; Castric-Fey, 1970, own data), but likely
some animals are reproductive all year round. Asexual reproduction may take place by
budding small polyps at the junction of the foot and blastostyle region (Hardy, 1891;
Hartlaub, 1916). This asexual budding takes place before the onset of sexual repro-
duction in early spring (Hardy, 1891).

The animals occur at the spring tide low-water-mark, but records down to 17 m
are known (Castric-Fey, 1970; Segonzac & Vervoort, 1995). Sars (1874) gives a depth
of 110-146 metres for his record from the Aalesund, which is unusually deep for this
animal. Intertidally, the species occurs attached on the underside of large boulders.
They can also be attached to holdfasts of laminarians (Castric-Fey, 1970). Billard
(1921) made some preliminary observations on their feeding biology. With their much
extensible trunk they search their surroundings for small benthic amphipods. It takes
about five to six hours to digest one of them. The animals are also able kill shrimps of
up to 2 cm size.

The embryonic development takes place in the embryonic envelope that is held
by the clasper tentacles. The animal can thus be considered an actively brooding
species. The clasper tentacles only attach to fertilized eggs, as only these form the nec-
essary embryonic envelope (Beigel-Heuwinkel, 1988). The development results in a
young polyp with 17-22 tentacles (Billard, 1921). The primary tentacles of the young
polyp are only transitory and are replaced by permanent tentacles. More details on the
development are given in Allman (1876, as Myriothela), Billard (1921), Benoît (1923a,
1925), and van der Vyver (1968).

ADDITIONAL DATA: The gastrodermis has numerous folds and villi (cylindrical
projections), but has no compartmentalization as in the Tubulariidae (Allman, 1876).
The mesogloea is massive and contains thick fibres (Beigel-Heuwinkel, 1982a). It is
also involved in the attachment to the substratum (Manton, 1941). Histological details
can be found in Allman (1876, as M. phrygia) and Hardy (1891, as M. phrygia). The
tentacles are hollow. Prévot (1959) shows them as closed off to the gastric lumen by a
mesogloeal lamella, but Allman (1876, pl. 56, fig. 2) states that they are open. An
examination of some hand-made cross-sections of the available material showed that
there are indeed small openings, but this should be corroborated by more reliable
evidence obtained by serial histological sections. The strong contractibility of the
tentacles may be regulated by the hydrostatic pressure of the water in the lumen of the

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 345

tentacles. If so, an opening to the stomach seems more understandable. The initial
tentacles formed while the embryo is still in the egg capsule develop towards the
gastric lumen and are only everted at a later stage (Allman, 1876). The primary
tentacles must thus have a basal opening. The primary tentacles are quite long and
replaced after hatching by shorter tentacles (Allman, 1876).

The sporosacs and their development have been examined by Allman (1876, as
M. phrygia), Korotneff (1888), Hardy (1891, as M. phrygia), Benoît (1923b), and
Beigel (1976). Manton (1941) studied the foot and the clasper tentacles. The clasper
tentacles were also investigated by Beigel-Heuwinkel (1988). She used light and
electron microscopic data to show that the cells at the tip secrete a substance that acts
to glue the claspers to the embryonic envelope. Unfertilized eggs do not form an
embryonic envelope and are thus not held by the claspers.

Results of regeneration experiments were reported by Billard (1921) and
Beigel-Heuwinkel (1982b). Regenerating upper halves of a polyp form no perisarc
sheath, but anchoring tentacles as in C. phrygium.

DISTRIBUTION: North-Eastern Atlantic, absent from the North Sea, Baltic Sea,
Mediterranean and Black Sea. The northernmost record is Norway (Aalesund,
G. O. Sars, 1874; somewhat doubtful, needs reconfirmation), common in the western
English Channel coast of Great Britain and France, also recorded from the Scilly
Islands, Isle of Man, western England, southern Brittany, Galicia, Bay of Cadiz (south-
ernmost record) (Segonzac & Vervoort, 1995, Medel & Löpez-Gonzälez, 1996, this
study). Type locality: Gyllyngvase, (Gwyllyn-vase in Cocks, 1850), Falmouth,
Cornwall, United Kingdom.

REMARKS: Candelabrum cocksii has usually been attributed to Vigurs (1849),
but that is incorrect. The binomen Arum cocksii was introduced in a paper by Cocks
that almost certainly appeared in 1850 and not 1849 (Cornelius, 1977). Cocks
attributed the name to Vigurs without providing a description. It is thus an invalid name
(nomen nudum). Furthermore, it seems that Cocks initially did not recognize it as a
hydrozoan as he placed it among the Sipuncula. Gosse (1853a: 126) then published the
name Spadix purpurea, of which Cocks (1853: 365) acknowledged in the same journal
that it is identical to his Arum cocksii. Cocks (1853) used Spadix purpurea as species
name followed by Arum cocksii in brackets to formalize the synonymy. A suitable
description and figures of Arum cocksi followed shortly afterwards in Cocks (1854),
which made the name formally available for the first time [ICZN, 1999, 4th ed., 50.1]
[the publication date of Cocks’s paper is somewhat unclear, it could be 1853 or 1854;
according to Cornelius (1977) it was likely 1854]. Gosse (1853c: 386) acknowledged
the synonymy and he apparently ceded his species designation to Cocks by the
footnote ".....I gladly recognise, however, the superior claim [of Cocks] of the specific
appellation, which pays a deserved compliment to an excellent naturalist."

Although Spadix purpurea Gosse, 1853 is likely a senior synonym of Arum
cocksii Cocks, 1854, the combination Candelabrum cocksii (Cocks, 1854) must be
taken as valid, this because to my knowledge Gosse's name has not been used as valid
after 1899 [ICZN, 1999, 4th ed., 23.9.1.1], while C. cocksii has been used regularly (see
Vervoort & Segonzac, 1995).

346 P. SCHUCHERT

Candelabrum cocksii (Cocks, 1854) has unfortunately been confounded with C.
phrygium (Fabricius, 1780) by Hincks (1868), Allman (1876), and many subsequent
authors relying on Hincks (see Vervoort & Segonzac (1995) for synonymy and
misidentifications). Recently, Cornelius (1977) again maintained that both are conspe-
cific. Cornelius based his claim by referring to other authors, e. g. Stechow (1923) and
Teissier (1965). These two authors, however, do not claim such a synonymy, and
Stechow even advocates a separation at the genus level. The opinion of Cornelius is
also not shared by Segonzac & Vervoort (1995) as well as by the present author. Both
species are clearly separable. Candelabrum cocksii and C. phrygium differ in the foot
morphology (long, perisarc covered foot versus short, naked one); clasper tentacles
(presence versus absence), and the sexual reproduction (hermaphroditic versus gono-
choristic).

The tentacles of the trunk are usually short and only indistinctly capitate, this in
preserved material as well as in living material taken into the laboratory. The tentacle
pedicels are very contractile and undisturbed animals have quite long tentacles (2 mm;
Allman, 1874). This has also been observed for the closely related species C. serpen-
tarii (see figs 3A-B in Segonzac & Vervoort, 1995). Candelabrum serpentarii
Segonzac & Vervoort, 1995 is only known from deep waters of the central Atlantic.
This species is also monoecious, but lacks clasper tentacles and its eggs are four to five
times larger. The only Candelabrum species that also has clasper tentacles is the Pacific
Candelabrum fritchmanii Hewitt & Goddard, 2001. This species can form colony-like
aggregates, a unique feature within this genus.

Candelabrum phrygium (Fabricius, 1780) Fig. 8

Lucernaria phrygia Fabricius, 1780: 343.

Myriothela arctica M. Sars, 1850: 14.

Myriothela phrygia — Sars, 1877: 23, pl. 2 figs 29-36; Bonnevie, 1899: 35, pl. 4 figs 5-6;
Jäderholm, 1908: 9, pl. 1 fig. 7; Broch, 1916: 19, fig. C, pl. 1 figs 3 & 8; Rees, 1957: 486,
fig. 36; in part Naumov, 1969: 261, not figures; Calder, 1972: 222, pl. 1 fig. 5.

? Myriothela gigantea Bonnevie, 1898a: 490, pl. 27 figs 46-47; Bonnevie, 1899: 38, pl. 4 fig. 1.

? Myriothela minuta Bonnevie, 1898a: 489, pl. 27 fig. 44: Bonnevie, 1899: 37, pl. 3 fig. 6a-b, pl.
4 fig. 4.

? Myriothela mitra Bonnevie, 1898a: 489, pl. 27 fig. 43; Bonnevie, 1899: 38, pl. 3 fig. 6c-e, pl.
4 fig. 3.

Candelabrum phrygium — in part Cornelius 1977: 521 [excl. synonymy]; Segonzac & Vervoort,
1995: 45, figs 2e-f, 3E-F, table 1 [some references do not refer to this species];
Schuchert, 2001a: 37, fig. 24.

MATERIAL EXAMINED: Re-examined material mentioned in Schuchert (2001a), ZMUC,
Greenland, no exact locality and date known, collected by Liithken, identified by P. Kramp, two
specimens, both broken into two parts, one obvious female on branching bryozoan (Cellariidae),
other animal on red algae, previously identified as male; one blastostyle of both specimens was
used for serial histological sections, both specimens proved to have female sporosacs only,
though the tissue preservation is not good. — All suitable material found in the BMNH that is la-
belled as C. phrygium turned out to be C. cocksii.

DIAGNOSIS: Candelabrum species without clasper tentacles, dioecious, basal
foot short or absent, straight and not sheathed in perisarc envelope, attached to sub-
stratum by tentacle-like filaments usually ending in perisarc discs.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 347

Fic. 8

Candelabrum phrygium (Fabricius, 1780), after preserved material from Greenland. A) Entire
polyp, scale bar 2 mm. B-C) Blastostyles, scale bar 0.5 mm.

DESCRIPTION (after Sars, 1877; Segonzac & Vervoort, 1995; own data): Polyp
solitary, worm-like, size very variable as able to expand and contract enormously.
Hydranth shape also variable, usually cylindrical to conical, subdivided into a distal
tentaculate region (trunk), followed by a blastostyle region and sometimes also a foot
zone.

Foot zone either very short in relation to other parts or absent, not curved,
adhering to substratum by several tentacle-like attachment filaments, each of them with
terminal region sucker-like and enlarged, attached to a perisarc disc that adheres to sub-
stratum, perisarc discs sometimes absent.

Blastostyle region about 1/4 to 1/3 of contracted polyp, beset by many (>20)
club-shaped blastostyles. At distal end of blastostyles four to six nematocyst clusters in
wart-like tubercles. Blastostyles bear sessile sporosacs, the two sexes on different
polyps, sporosacs without nematocyst tubercles. Male sporosacs spherical, without
radial canals, up to 50 per blastostyle at different developmental stages. Females with
two to six sporosacs per blastostyle, maximally two mature, others in development,
spherical, no radial canals, initially many eggs but presumably only one egg per
sporosac attains maturity, fertilized in situ and developing into a young polyp, hence
viviparous.

348 P. SCHUCHERT

Trunk region comprising majority of hydranth length, with numerous (>200)
capitate tentacles, these hollow, extensible, capitula ovoid.

Colours: cream-white. Nematocysts: stenoteles; ? haplonemes; desmonemes of
two size classes.

DIMENSIONS: Height of contracted specimens 2-6 cm, expanded up to 30-40 cm
(Bonnevie, 1899; Broch, 1916), width a few mm. Diameters of male sporosacs 0.4-0.8
mm when mature, females 0.9-1.4 mm. Capitate tentacles with stalk 0.3-0.5 mm,
capitula 0.17-0.25 mm (Segonzac & Vervoort, 1995). Nematocysts (Segonzac &
Vervoort, 1995): stenoteles (10.6-11.5)x(8.2-9.8)um; ? haplonemes (19.7-20.5)x(8.2-
9.9)um; desmonemes of two size classes (12.5-13.0)x(9.0-9.8) um and (8.2-9.0)x(6.4-
6.6) um.

BIoLOGY: Occurs usually at considerable depths of several hundreds of meters
down to 2195 m (Bonnevie, 1899), but in the high Arctic it as been found as shallow
as 13 m (Jäderholm, 1908). The polyps live permanently attached to solid substrata like
rock, bivalves, hydroids, bryozoans, and algae. The animals are viviparous and lack a
planula phase (Sars, 1877; Schuchert, 2001a); the newly released polyp is spherical
and has 20-30 capitate tentacles. The tentacles formed while the embryos is still in the
egg capsule develop inverted into the gastric lumen, but evert before hatching.

DISTRIBUTION: An Arctic, deepwater species penetrating into boreal regions, in
European waters reaching as far south as the Trondheimfjord and the Wyville-
Thomson-Ridge between Scotland and The Faeroes (Broch, 1903; Broch, 1916).
Segonzac & Vervoort (1995) report a find from the Mid-Atlantic Ridge south-west of
the Azores in a depth of 1622 m, wich is the southernmost record of this species. It has
also been recorded in northern Norway (Sars, 1877; Bonnevie 1899), Jan Mayen
(Broch, 1916), Russian Arctic Seas (Jäderholm, 1908; Naumov, 1969), northern Pacific
(Naumov, 1969). Also known from Iceland (Broch, 1916), Greenland (Schuchert,
2001a), Canada (Calder, 1972). (Note that numerous other records under this name
from costal regions of the NE Atlantic refer in fact to Candelabrum cocksii). Type
locality: Greenland.

REMARKS: The synonymy of this species has been worked out quite thoroughly
(Sars, 1877; Bonnevie, 1899; Segonzac & Vervoort, 1995). Myriothela arctica Sars,
1850 was synonymized by Sars himself after he had re-examinated Fabricius’ type
material (Sars, 1877). The boreal shallow-water species Candelabrum cocksii has often
been synonymized with this species, but this is not tenable (see under C. cocksii). Some
of Bonnevie’s Candelabrum species (C. minutum, C. mitra, C. giganteum) are not well
characterized and might also belong to this species (Rees, 1956). They are here listed
as questionable synonyms (see also Segonzac & Vervoort, 1995). The material on
which these three species were based, already fragmentary when examined by
Bonnevie (1899), is now in such a bad condition that it is virtually useless (Rees,
1956). Contrary to the view of Rees (1956), Candelabrum verrucosum (Bonnevie,
1898) is well characterized and it is re-described below.

Candelabrum phrygium is portrayed as being dioecious, but for preserved
material it is often difficult or impossible to determine the sex. This is easy only for

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 349

well advanced female sporosacs as they contain young polyps. For a reliable sex de-
termination, serial histological sections must be made. One blastostyle of each of the
two specimens examined for this study was thus used to make serial sections. Both
animals had female sporosacs only. One specimen did not have sporosacs with
advanced embryos and was thus initially mistaken for a male (Schuchert, 2001a). This
shows that sex identifications of preserved material are often unreliable. More inves-
tigations using histological sections of entire blastostyles are desirable to confirm that
C. phrygium is always dioecious (comp. C. verrucosum).

The material examined in this study and also by other authors (Sars, 1877) had
attachment tentacles that were cemented to the substratum by distinct perisarc discs
that adhere tightly to the substratum and cannot be removed without destroying them.
Dislodged hydranths lack them regularly. Sometimes they are quite thin and incon-
spicuous and if attached on rock they might be hardly visible. This could explain that
some investigators did not find them (e. g. Segonzac & Vervoort, 1995), but is well
possible that they can also be absent.

Candelabrum verrucosum (Bonnevie, 1898) Fig. 9
Myriothela verrucosa Bonnevie, 1898a: 468, pl. 27 fig. 45; Bonnevie, 1899: 37, pl 4, fig. 2.
Candelabrum verrucosum — Segonzac & Vervoort, 1995: 53.

MATERIAL EXAMINED: ZMUC, Kap Farvel expedition station 148, 60.07°N 43.20°W
(Greenland), 50 m, 27 August 1966, from rocky bottom, fertile female with sporosacs contain-
ing embryos, tissues somewhat shrunken. Two blastostyles were used to make serial histological
sections, confirming the presence of male and female sporosacs.

DIaGNOSIS: Candelabrum species with sporosacs bearing scattered nematocyst
buttons on their surface, monoecious, no clasper tentacles; foot straight, not covered by
perisarc, attached via attachment filamants; size 1-4 cm.

DESCRIPTION (after examined specimen): Solitary polyp, 1.1 cm, cylindrical,
divided into three regions: distal tentaculate region (trunk), followed by a blastostyle
region and a foot zone.

Foot zone about 1/6 of animal, straight, with several (>10) tentacle-like attach-
ment filaments of variable length, some of them with enlarged, sucker-like ends; one
filament with terminal perisarc disc, those of other filaments presumably lost.

Blastostyle region taking up about half of the polyp, bearing many (>20) club-
shaped blastostyles, clasper tentacles absent. Nematocyst clusters in wart-like tubercles
at distal end of blastostyles. Blastostyles bear several (up to 8) sessile sporosacs.
Sporosacs with up to eight distinct nematocyst buttons on surface, no radial canals or
ring canals present. Largest sporosacs contain a single young polyp, animal thus
viviparous. Besides female sporosacs, there is also one male sporosac per blastostyle,
animals thus simultanous hermaphrodites.

Trunk region about 1/3 of hydranth length, covered by numerous (>100) capi-
tate tentacles, their capitula ovoid.

Nematocysts (not well preserved): stenoteles, large desmonemes, ? hetero-
nemes.

DIMENSIONS: Hydranth can reach 4 cm (Bonnevie, 1898a), sporosacs up to
0.9 mm, blastostyles up to 2.2 mm, diameter of capitula 0.2 mm.

350 P. SCHUCHERT

Fic. 9

Candelabrum verrucosum (Bonnevie, 1898), after preserved material from Greenland. A) Whole
animal, the blastostyle region covers slightly more than half of the animal, note the nematocyst
buttons on the sporosacs, scale bar 2 mm.

ADDITIONAL DATA: The tentacles formed while the embryos is still in the egg
capsule develop inverted into the gastric lumen, but evert before hatching.

DISTRIBUTION: Northern Norway, Southern Greenland (this study). Type
locality: Norway, Hammerfest (Bonnevie, 1899).

REMARKS: Candelabrum verrucosum (Bonnevie, 1898) has hitherto been
known from one specimen described by Bonnevie (1898a, 1899). Some authors
doubted its validity and thought it might belong to C. phrygium (e. g. Rees, 1956). The
species is characterized by sporosacs that bear scattered buttons of nematocyst clusters.
This is quite a unique feature and has never been observed for C. phrygium. I am
convinced that C. verrucosum is a valid species, distinct from C. phrygium, and that
the material described above belongs to it. The characteristic nematocyst buttons on the
sporosacs are quite conspicuous in material examined with a dissecting microcope.
There are a few discrepancies from the original material described by Bonnevie
(1898a): there are more than two sporosacs per blastostyle, the blastostyle region is
larger, and the animal is smaller. As these traits show considerable intraspecific
variation in other Candelabrum species, they are here also considered as such.

Candelabrum verrucosum is easily distinguished from C. phrygium by the
nematocyst buttons on the sporosacs, but it differs additionally by being monoecious.
Otherwise, both species are very similar. Only two blastostyles could be used to make
serial histological sections. Both blastostyles had female sporosacs of all develop-

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 351

mental stages (some containing almost fully formed young polyps) and a single male
sporosac filled with spermatids. The male sporosacs are smaller than the female ones.

Genus Monocoryne Broch, 1910
TYPE SPECIES: Coryne gigantea Bonnevie, 1898b.
SYNONYMS: Symplectanea Fraser, 1941 (see Rees, 1958).

DIAGNOSIS: Hydroid solitary or a few polyps in loose aggregates that may have
a common perisarc base. Hydranth long, cylindrical, divided into tentaculate body and
foot region. Foot covered by thin perisarc, with root-like attachment processes.
Hydranth body with scattered capitate tentacles, at least some of them compound
tentacles, i.e. branched tentacles with a common epidermis at the base, bases of the
side-branches adnate to the side of the main tentacle for some distance, all ends capi-
tate. Gonophores fixed sporosacs developing directly on hydranth body, associated or
not with tentacles.

REMARKS: For more details consult Rees (1956, 1958), Petersen (1990), or
Stepanjants et al. (2003). The European fauna comprises one species only. Stepanjants
et al. (2003) give an overview on all species.

Monocoryne gigantea (Bonnevie, 1898) Fig. 10

Coryne gigantea Bonnevie, 1898b: 4, pl. 1 fig. 1.
Monocoryne gigantea — Broch, 1910: 138; Broch, 1916: 12, pl. 1 fig. 1; Johannesen, 1924: 1,
figs 1-7, pl. 1-2; Rees, 1956: 117, figs 1-2; Rees, 1957: 488, fig. 38; Calder, 1972: 222,
pl. 1 fig. 4; Antsulevich, 1988: 931, fig'd; Petersen, 1990: 203; Stepanjants et al., 2003:
100, figs 1A-F, 6.1.
DIAGNOSIS: Monocoryne species up to 15 mm; compound tentacles with two to
four capitate ends, usually three; hermaphroditic, sporosacs in upper axils of
compound tentacles.

DESCRIPTION: (after Bonnevie, 1898b; Johannesen, 1924; Rees, 1956;
Stepanjants ef al., 2003) Vermiform hydranths, solitary or clustered into loosely joined
aggregates, attached laterally to substratum by curved proximal end; divisible into foot
(caulus) zone and tentaculate part. Foot roughly 1/2 of total length, covered by close
fitting, soft, thin perisarc, in lower half of foot several large, distinct anchoring
filaments with widened distal end. Tentaculate part also about 1/2 of length, but very
extensile and active in life, all tentacles distinctly capitate with spherical capitula, very
extensible, around mouth about eight simple capitate tentacles, below them many
(< 30) scattered compound tentacles and also some simple ones. Compound tentacles
usually with three, sometimes two or four capitate ends, with middle branch(es)
thickest and longest, the side-branches originate near its base and are adnate for some
distance before they become free, the fused part forming a plate-like base with a
common epidermis. Gonophores sessile sporosacs arising in the upper axils of the
compound tentacles, sporosacs oblong, without radial canals; male, female and herma-
phroditic sporosacs can be produced by the same animal (Broch, 1916; Johannesen,
1924). Nematocysts: stenoteles, microbasic mastigophores, desmonemes, microbasic
euryteles.

352 P. SCHUCHERT

Fic. 10

Monocoryne gigantea (Bonnevie, 1898), modified from Rees (1956). A) Whole polyp, foot with
perisarc stippled dark. B) Compound tentacle. C) Side view of sporosac in axil of compound
tentacle.

DIMENSIONS (Rees, 1956; Stepanjants et al., 2003; consult these references for
additional measurements): Fertile polyps 11-15 mm (preserved material), diameter of
hydranth body 1.4-1.7 mm, length of gonophores 0.6-0.9 mm, diameter 0.4-0.5 mm.
Nematocysts: stenoteles (14-18)x(12-18) um; microbasic mastigophores (19-24)x(8-
11) um; desmonemes 10x8 um, microbasic euryteles (22.5-25)x(12-12.5) um.

BIOLOGY: A rare species, occurs usually in waters of 100 m depth and more, but
in the high Arctic archipelago Franz Joseph Land it was found in only 16-20 m
(Antsulevich, 1988). Known substrata (Swenander, 1904) are a Tubularia spec. and
polychaete tubes, both attached on shells of the bivalve Lima excavata.

DISTRIBUTION: Arctic species, with its southern limit in the Trondheimsfjord
(Johannesen, 1924). It has been recorded from northern Norway (Bonnevie, 1898b),

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 353

Franz Joseph Land (Antsulevich, 1988), North-Eastern Canada (Calder, 1972). Type
locality: Hammerfest, Norway.

REMARKS: Rees (1958) synonymized the genera Symplectanea and Mono-
coryne, but kept M. bracteata provisionally distinct from M. gigantea because of its
larger size and the greater number of capitate heads per compound tentacles, but he
suggested that more material might show it to be synonymous with M. gigantea. It
could be that Monocoryne bracteata is dioecious, thus differing from the monoecious
M. gigantea. The only other named species of the genus is Monocoryne minor Millard,
1966. It is smaller and its sporosacs develop independently of the tentacles.

FAMILY TRICYCLUSIDAE KRAMP, 1949

DraGNOSIS: Solitary, usually benthic hydroids, having a conical to pear-shaped
hydranth and tapering pedicel ending in a small attachment disc. Pedicel covered in a
loose, filmy or gelatinous perisarc. Tentacles in three whorls, one oral whorl, one in
middle of hydranth body, and one near base of body. Oral tentacles capitate, other
tentacles also capitate but bearing additional nematocyst clusters. Vegetative budding
of hydranths below proximal tentacles. Gonophores develop above proximal set of
tentacles and remain fixed. Male gonophores medusoid, with radial canals and circular
canal. Female ones without canal system. Cnidome: Stenoteles, desmonemes, and
heteronemes.

Genus Tricyclusa Stechow, 1919
TYPE SPECIES: Tiarella singularis Schulze, 1876.
SYNONYM: Tiarella Schulze, 1876.

DIAGNOSIS: As for the family.

REMARKS: The genus Tricyclusa is currently monotypic, containing only
Tricyclusa singularis (Schulze, 1876). Schulze (1876) originally proposed the genus
name Tiarella for this species. Because this name is preoccupied for a gastropod and
also other taxa, Stechow (1919: 6) proposed the new name Tricyclusa. It is a charac-
teristic genus and it poses no taxonomic problems.

Tricyclusa singularis (Schulze, 1876) Fig. 11

Tiarella singularis Schulze, 1876: 415, pls 29-30; Bedot, 1911: 209, pl. 11 fig. 2.

Tricyclusa singularis — Stechow, 1919: 6; Rees, 1941: 133, fig. 3; Rees, 1957: 462, 505, figs 6
& 51B; Picard, 1957: 10; Vevers, 1959: 506, figs 1-6; Teissier, 1965: 10; Bouillon, 1974:
142; Petersen, 1990: 146, fig. 15; Bouillon et al., 2004: 103, fig. 55D.

Margelopsis stylostoma Hartlaub, 1903: 28, fig. 2; Hartlaub, 1907: 91, fig. 87; Bedot, 1911: 211;
Rees, 1941: 133.

MATERIAL EXAMINED: BMNH 1957.6.26.1-10, Roscoff intertidal, leg. Cantacuzène, on
red algae, numerous well preserved specimens. - BMNH 1956.11.7.7-13, Baie de Morlaix, 23
June 1955, coll. W.J. Rees, on Chorda filum, several contracted specimens.

DIAGNOSIS: As for the family.

DESCRIPTION (after literature and observed material): Hydroid solitary, usually
attached to algae, sometimes freely floating, divided into hydranth body and pedicel,

354 P. SCHUCHERT

Fic. 11

Tricyclusa singularis (Schulze, 1876), after preserved material. A) BMNH 1957.6.26.1-10,
Roscoff, polyp with long tentacles, scale bar 0.2 mm. B) polyp with short tentacles, after
Petersen (1990), scale bar 0.5 mm. C) Optical section of a female sporosac, modified after Bedot
(1911), scale bar 0.2 mm. D) Optical section of a male medusoid after it has liberated its
gametes, modified after Schulze (1876), scale bar 0.05 mm

both of about the same length. Pedicel in a conical, loose, filmy or gelatinous periderm
cup, wrinkled, with adhering detritus particles. Pedicel tapering proximally, ending at
base in a small attachment disc. Hydranth pear-shaped to conical, broadest near lower
end, with three distinctly separated sets of tentacles, tentacle gastrodermis chordoid.
Oral tentacles in one whorl, four to five in number, short, capitate, directed upwards;

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 355

middle whorl of tentacles approximately in middle of body, usually six in number,
directed obliquely upwards, with terminal nematocyst knob and with or without addi-
tional one to two knobs in distal half of tentacle; basal tentacles 10 to 14, usually 12,
in two closely approximated whorls, alternately pointing up- and downwards, with
terminal knob and additionally one or more proximal knobs, these either clasping or
encircling tentacle. Underneath lowest tentacles vegetative polyp buds, one to six in
number, budded polyps are released as small, fully formed hydranths but have a short
pedicel, newly released polyps may already have incipient gonophores. Gonophores
develop immediately above basal whorl of tentacles, spherical, remaining attached to
hydroid. Male gonophores with four radial canals and ring canal, thus of medusoid
type. Female gonophores sporosacs without canal system, with one large egg. Nemato-
cysts: Stenoteles, desmonemes, microbasic euryteles.

DIMENSIONS: Total height around 2 mm, hydranth about 1 mm high, hydranth
base diameter 0.4-0.6 mm. Female sporosacs after Bedot (1911) about 0.6 mm, male
medusoids after Schulze (1876) about 0.2 mm.

ADDITIONAL DATA: The length of the tentacles is variable, and this is not only
due to contraction. Young individuals and polyps found in the plankton have very long
and thin tentacles that get shorter once the animal attaches itself (Bedot, 1911). Young
animals and those from the plankton also have a short pedicel.

DISTRIBUTION: Adriatic Sea, Northern Brittany, western Ireland. Type locality:
Adriatic Sea, Trieste, Bay of Muggia (Italy), on Cystoseira.

BioLoGY: The polyps live attached to various macroalgae and Zostera plants
near the low-water-mark (Rees, 1941, Teissier; 1965). Occasionally, detached polyps
can be found in the plankton, but this is not the usual mode of life (Bedot, 1911). In
Brittany it occurs mainly from May to July, although some animals were also found
during March and April (Bedot, 1911; Teissier, 1965). In the Mediterranean it was
found in April (Schulze, 1876). The fertile period for Brittany is from May to June
(Bedot, 1911; Teissier, 1965). It is a rare species, but when present, it can occur in large
numbers (Bedot, 1911).

REMARKS: Tricyclusa singularis (Schulze, 1876) is a characteristic and un-
problematic species. Hartlaub (1903, 1907) described a similar species from the
plankton of Roscoff which he named Margelopsis stylostoma, differing mainly only in
the short pedicel. Even Hartlaub (1903, 1907) suspected that it was only a planktonic
form of Tricyclusa singularis. It was later synonymized with Tricyclusa singularis by
Bedot (1911) and Rees (1941).

Although easy to identify and despite its presence near the water surface, this
animal has only rarely been reported. After its discovery, it has never been found again
in the Mediterranean Sea. It seems that it is only occasionally present and quite
seasonal. Most regularly it has been seen at Roscoff (Brittany). It is particularly
interesting to note that it has never been reported from the other side of the English
Channel, despite England being one of the best investigated regions. Perhaps it prefers
warmer waters, an assumption also underlined by its occurrence during the summer
months. Probably it survives unfavourable conditions through a sexually produced
resting stage.

356 P. SCHUCHERT

FAMILY MARGELOPSIDAE UCHIDA, 1927
TYPE GENUS: Margelopsis Hartlaub, 1897.
SYNONYM: Pelagohydridae Dendy, 1902.

DIAGNOSIS: Hydroid solitary, pelagic; hydrocaulus absent or reduced to a small
process; hydranth body vasiform, tentacles filiform to moniliform, arranged into two
separate sets, oral tentacles in one or several whorls; aboral tentacles either in two to
three alternating whorls or numerous and scattered over most of body. Gonophores free
medusae, medusa buds develop among tentacles or above aboral tentacles.

Medusa manubrium with simple mouth; gonads surrounding manubrium
entirely; four radial canals; tentacles solid, generally moniliform, two or more tentacles
per marginal bulb, in some genera tentacles also issuing at different levels on exum-
brella; without ocelli.

REMARKS: With its clustered marginal tentacles in the medusae and the pelagic
polyps, this is a distinct and characteristic family of the Capitata. It comprises the
genera Climacocodon Uchida, 1924, Margelopsis Hartlaub, 1897, and Pelagohydra
Dendy, 1902 (Petersen, 1990). The well established and frequently used name
Margelopsidae Uchida, 1927 is threatened by the senior synonym Pelagohydridae
Dendy, 1902. The latter name has not been used and for the sake of nomenclatural
stability it is preferable to continue to use Margelopsidae. According to the ICZN
[article 29.3.1.1] there is no need to change it to the formally correct Margelopsididae.

KEY TO GENERA:

la medusa! with; several tentacle pairs On\exumbrella 7. bee ee eeeee

PES RE EN A a RN ces ae Climacocodon (not in European fauna)
lb medusa'tentacles‘'in’four’groups'alone bell'Margin e RENON 2
2a polyp with caulus rudiment, aboral tentacles in two to three close whorls . . Margelopsis
2b polyp without caulus rudiment, aboral tentacles scattered .......................

BORE E REI RO CD ME tete Pelagohydra (not in European fauna)

Margelopsis Hartlaub, 1897
TYPE SPECIES: Margelopsis haeckelii Hartlaub, 1897 by monotypy.

DIAGNOSIS: Hydroid with hydrocaulus rudiment, without parenchymatic spe-
cializations of the gastrodermis; tentacles indistinctly moniliform; oral tentacles in one
whorl, aboral ones in two to three whorls, medusae buds above aboral tentacles.

Medusa with four perradial tentacular bulbs on bell margin, each with two to six
solid tentacles.

REMARKS: There are two species of Margelopsis in the European fauna, name-
ly Margelopsis haeckelii and M. hartlaubii. See the diagnoses for ways to distinguish
them.

Margelopsis haeckelii Hartlaub, 1897 Figs 12-13
Margelopsis Haeckelii Hartlaub, 1897: 482, pl. 16b figs 12-18; Hartlaub, 1899: 219, figs 1-3
[hydroid].

Margelopsis haeckeli — Hartlaub, 1907: 89, 91, figs 84-86; Müller, 1908: 43, pl. 4 figs 12-17, pl.
5 figs 18-19; Mayer, 1910: 80, fig. 38; Leloup, 1930: 97, fig.; Kramp, 1930: 12; Kramp,
1937: 32, fig. 10; Thiel, 1938: 294; Leloup, 1946: 1; Russell, 1953: 95, figs. 41A-C,

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 357

Fic. 12

Margelopsis haeckelii Hartlaub, 1897; schematic pictures derived from various preserved sam-
ples. Mature medusa with subitaneous eggs and one young polyp, scale bar 0.3 mm. B) Oral
view of a medusa, tentacles clipped, same scale as A.

42A-B; Werner, 1954: 124, figs 1-9; Werner, 1955: 1-30, figs 1-9; Prévot, 1959: 104, pl.
3 fig. 11; Kramp, 1959: 92, fig. 47; Kramp, 1961: 49; Russell, 1970: 234; Bouillon, 1974:
143.

MATERIAL EXAMINED: BMNH 1967.5.25.1-8, Helgoland, 12 May to 24 July 1958,
medusae and polyps. - BMNH 1967.5.25.6, 2 medusae from Sylt, 10 July 1958, with resting
eggs. — Zoological Museum Hamburg, about 50 mature medusae, collected in plankton by
B. Werner, 24 July 1958, with subitaneous and resting eggs. — Zoological Museum Hamburg,
several polyps from List, collected 27 June 1961, cultivated by B. Werner, with medusae buds.

DIAGNOSIS: Margelopsis polyp with vasiform body, stalk rudiment, two well-
separated sets of tentacles, medusae buds in one whorl oralward of aboral tentacles.
Medusae of up to 2 mm high, umbrella bell-shaped, moderately thick, apical canal
present, 3-6 tentacles per bulb, manubrium base with large vacuolated cells, usually
only females present, eggs of two types: subitaneous eggs developing directly into
polyp stage, diapausing eggs developing first into encysted resting stage.

358 P. SCHUCHERT

Fic. 13

Margelopsis haeckelii Hartlaub, 1897: schematic picture derived from various preserved sam-
ples. Polyp with medusae buds, oral side above, scale bar 0.2 mm.

DESCRIPTION (after examined material and literature): Newly released polyps
actinula-like, ovoid, with short tentacles, tentacles numbers slightly lower than in ful-
ly mature animals.

Mature polyp planktonic, body vase-shaped, resembling a detached tubularian
hydranth, at aboral end a short stalk rudiment with a central depression. Tentacles in
two well-separated sets, rather stiff, with nematocysts concentrated in rings and a small
terminal knob, base of tentacles free of nematocysts. Aboral tentacles slightly longer
than oral ones, 12-15 in two closely approximated whorls, alternately pointing
obliquely up- and downward. Oral tentacles around short conical hypostome, five to
eight in number. Medusae buds borne on body of hydranth just oralward of the aboral
tentacles, sometimes groups of medusa buds on a short stalk. Medusae develop without
envelope, the tentacles are free and not tucked into the subumbrella as in most other
hydromedusae.

Umbrella of very young medusae relatively wider than in adult, apical jelly thin,
nematocysts evenly scattered over exumbrella; broad umbilical canal, stomach length
less than half the height of subumbrellar cavity, a circle of nematocysts surrounds the
mouth-opening, without gonads. Four radial canals and ring-canal narrow. Four
perradial marginal bulbs each with two to three tentacles. Tentacles covered with
scattered clusters of nematocysts, but base of tentacles without nematocysts. Umbilical
canal and marginal bulbs with black pigment. Umbrella becomes higher with further

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 359

growth. In a specimen 1.5 mm high, apical jelly considerably thicker than sides of
umbrella, many exumbrellar nematocysts, umbilical canal has narrowed, stomach
considerably longer. Upper third of stomach without gonads, with large gastrodermal
cells covered by thin layer of epidermis. Gonad covering lower two- thirds of the
stomach thin. Three to four tentacles irregularly placed on each marginal bulb, with
irregularly scattered nematocysts which appear as ring-like bands when tentacle
contracted. Upper basal third of stomach clear and transparent in contrast to opaque
brownish grey appearance of lower part.

Adult medusa with bell-shaped umbrella, slightly higher than wide, without
apical process or with small apical process; with scattered exumbrellar nematocysts;
jelly moderately thick, thicker at apex, velum moderately broad to narrow. Stomach
cylindrical, length 2/3 to 1/1 of subumbrellar height; basal portion with large trans-
parent gastrodermal cells; apical canal regularly present (rest of umbilical canal);
mouth simple, circular, margin armed with nematocysts. Four radial canals and ring
canal narrow. Gonad surrounding stomach, leaving upper third free. Eggs amoeboid,
embryos developing attached to manubrium on pedicel, depending on season either
into young polyp or encysted resting stage. Four perradial rounded marginal bulbs,
each with four to seven somewhat stiff tentacles, usually irregularly radiating, with
nematocysts concentrated in rings and a small terminal knob, thus nearly moniliform,
gastrodermal cells chordoid. No ocelli. Colour of stomach dark grey with dark brown
pigment granules; marginal bulbs brown. Nematocysts desmonemes, basitrichous
haplonemes, microbasic euryteles, stenoteles.

DIMENSIONS: Adult medusa up to 2 mm high, subitaneous egg production starts
at a bell diameter of 1-1.5 mm and with 3-5 tentacles per bulb, resting eggs are
produced by full sized animals only; subitaneous eggs 0.12-0.13 mm, resting eggs
0.18-0.21 mm; newly released medusa 0.5 mm; newly released polyp resulting from
subitaneous eggs 0.3-0.4 mm, adult polyps 1-2 mm (Werner, 1954), aboral tentacles as
long as hydranth or slightly longer.

BIOLOGY: Polyps and medusae are usually present in the plankton from June to
September, but they have also been seen earlier. They are thought to remain floating
through water currents, as they sink in still water. Their numbers fluctuate drastically
from year to year. Werner (1954, 1955) made detailed investigations on the life histo-
ry and development of this species. Male medusae appear to be extremely rare; Werner
(1954) observed only one hermaphrodite in 250 medusae, the others being all female.
The immature eggs are amoeboid and grow by engulfing other eggs (Miiller, 1908).
The eggs develop parthenogenetically (Werner, 1956). Two different types of eggs are
produced: smaller subitaneous eggs and larger resting eggs. Both egg types start their
development attached to the manubrium by a small stalk. The total production of subi-
taneous eggs depends on the food availablity of the medusa and varies in number from
a few eggs to 30 or 40. For the most part, only two to three egg cells mature and emerge
at the same time. Often a medusa bears 20-30 eggs and embryos on the manubrium, of
different ages and different developmental stages. The subitaneous eggs are produced
earlier when water temperatures are between 7 and 15°C. They develop directly within
four to ten days into a small polyp that is then released from the medusa. Later in the

360 P. SCHUCHERT

year, when water temperatures exceed 15°C, the production of subitaneous eggs stops
and resting eggs are produced. These are larger than the subitaneous eggs and possess
nematocysts on their surface. Generally, only one or two of this type are formed at the
same time and usually not more than six of them remain attached to the manubrium.
The initial development takes place when the egg is attached to the manubrium and
reaches a 'stereo-blastula' stage (a thin layer of ectodermal cells surrounds a mass of
yolky endodermal cells). At this stage development stops, the egg detaches and sinks
to the bottom, where it may attach or not as a lens-shaped cyst. The nematocyst layer
helps in the attachment process. The embryo forms a periderm capsule with a charac-
teristic polygonal pattern. The animal overwinters then as a cyst and after 6-9 months
a small polyp hatches in the next spring (Werner, 1984). The newly released polyp lives
in the plankton where it grows to full size and then produces medusae.

It seems that only few resting stages survive the winter, as initially there are
only very few hydroids in the plankton. However, through the following medusa pro-
duction and the polyps resulting from their subitaneous eggs, the population density
can grow very rapidly (Werner, 1955).

DISTRIBUTION: Southern North Sea and Irish Sea. Helgoland (Hartlaub, 1897;
1899; Werner, 1955); Norfolk coast (Hamond, 1964), Solway Firth (Russell, 1970),
Weser, Elbe, and Ems Estuary (Kiihl, 1962, 1967; 1971), Belgium (Kramp, 1930;
Leloup, 1947). Type locality: Helgoland.

ADDITIONAL DATA: The medusae swim very energetically and jerkily. The short
marginal tentacles are held out rather stiffly (Hartlaub, 1907).

Hartlaub (1907) observed that the polyps normally do not swim actively and
sink slowly to the bottom of the vessel. When sinking, the aboral pole is on top. The
polyps are very sensitive to temperature fluctuations. These observations could not en-
tirely be confirmed by Werner (1955), who observed that the polyps do not regularly
sink with the oral pole downward, and he thinks that the polyp in the free water does
not necessarily orient itself with the aboral pole uppermost.

The aboral stalk rudiment of the polyp has a depression lined with a high
epithelium with cylindrical cells, which Hartlaub (1899) considered reminiscent of the
pneumatophore of the Physophorae. Margelopsis haeckelii has been seen as a model
representing an intermediate stage in the evolution of the Siphonophorae (Totton,
1965). This view was contested by Werner (1955), who observed that this organ is used
by the hydroid to attach itself temporarily, acting like a sucker organ. The cells of the
depression secrete a mucus that is often infested with detritus particles. The aboral
stalk rudiment is thus clearly homologous to the corresponding organ by which the
tubularian actinula larva attaches itself, and ultimately also the stalk of the mature
hydranth.

When the medusa switches from the production of subitaneous- to resting eggs,
there may be a period when there are no conspicuous eggs on the manubrium. Such
medusae can easily be mistaken for mature males, although they do not have any
spermatids (Werner, 1955).

Prévot (1959) depicts a longitudinal section of the polyp. The stalk carrying the
medusae buds (blastostyle) is hollow and communicates with the stomach. In the polyp
material examined for this study, one bifid oral tentacle was seen.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 361

REMARKS: The original spelling in Hartlaub (1897) is Margelopsis haeckelii,
which must be retained, despite that Hartlaub (1907) used haeckeli, a spelling then
used by all subsequent authors (the specific epithet haeckelii is the genitive form of the
latinized name Haeckelius).

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 the genus Nemopsis (Bougainvilliidae). 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, only differing in the medusae
buds that are dispersed between the two whorls of tentacles. McCrady (1859: figs 4-6)
also describes and depicts young medusae stages released by this polyp. As A. Agassiz
(1862) and Hartlaub (1899) have already pointed out, McCrady’s medusae from the
plankton (McCrady, 1859: figs 1-3) were actually Nemopsis bachei L. Agassiz, 1849.
Therefore, Hartlaub (1899) restricted the name Margelopsis gibbesii to the polyp
described by McCrady. Although the first revisor was A. Agassiz (1862) who
synonymized the name N. gibbesii with N. bachei, Hartlaub’s proposal was followed
by Mayer (1910) when he attributed some Margelopsis medusae from North Carolina
to M. gibbesii. McCrady’s description did not include mature medusae and the allo-
cation of Mayer appears unfounded, but there is a reasonably good chance that Mayer’s
identifications were correct, as there are no other Margelopsis medusae known from
the region.

Thiel (1938) found a Margelopsis medusa in the Southern Ocean, which he
assigned to M. gibbesii and he claimed that M. gibbesii, M. haeckelii and M. hartlaubii
are conspecific, a possibility already suggested by Mayer (1910: 80). However, this has
not gained acceptance and Kramp (1959) distinguished the medusae of Margelopsis
gibbesii from M. haeckelii by the presence of an apical canal, the thicker jelly and the
viviparity in the latter species. The scattered medusae buds in the polyp of M. gibbesii
can also be added. According to Mayer (1910), Margelopsis gibbesii produces male
medusae, which indicates that both species differ perhaps also significantly in their
life-histories. However, as Werner (1955) cautioned, female medusae that switch from
the production of subitaneous- to resting eggs can easily be mistaken for males. More
biological details on M. gibbesii must be known before both species can be compared
and meanwhile it seems appropriate to keep M. gibbesii separate from M. haeckelii.

Besides M. haeckelii, there is also another Margelopsis species occurring in the
European fauna, namely M. hartlaubii. The latter species is known only from the
medusa phase occurring in deep waters of Norway, thus contrasting with M. haeckelii
which is a shallow water species. Margelopsis hartlaubii can be distinguished by its
lower tentacle number (2-3 per bulb placed beside each other), the thicker jelly, the ab-
sence of an apical canal, and the brick-red manubrium.

Margelopsis hartlaubii Browne, 1903 Fig. 14

Margelopsis hartlaubii Browne, 1903: 10, pl. 1 fig. 2, pl. 3 fig. 3; Mayer, 1910: 82, fig. 40;
Kramp & Damas, 1925: 252, fig. 4; Kramp, 1959: 91, fig. 49; Kramp 1961: 50.
MATERIAL EXAMINED: Zoological Museum Bergen, No 36451, 15 July 1925, Haggernes,

Herdlafjord, 200-400 m, 2 medusae, damaged before fixation. — Zoological Museum Bergen, No

362 P. SCHUCHERT

Fic. 14

Margelopsis hartlaubii Browne, 1903. A) Redrawn from Browne, scale bar 1 mm (1903). B)
Modified after Kramp & Damas (1925).

26825, Herlôfjord, September 1908 (sic), 1 medusa, fertile female, depicted in by Kramp &
Damas (1925). — Zoological Museum Bergen, No 26826, ca. 300 m, September 1908, 2 medusae
in bad condition, identified by Kramp & Damas (1925).

DIAGNOSIS: Margelopsis medusa measuring 2-4 mm, umbrella spherical to bell-
shaped, jelly relatively thick, no apical canal, with two to three moniliform tentacles
per bulb, gonads much bulging, likely no brooding of eggs.

DESCRIPTION (after Kramp & Damas, 1925): Polyp stage unknown. Medusa
umbrella spherical, 3-4 mm for mature animals, exumbrella with scattered nemato-
cysts. Mesogloea thick, especially at apex, its consistency very soft. The voluminous
manubrium takes up a large part of the subumbrellar cavity, its base quadrangular and
not covered by gonad; gonad occupies the middle region, shaped like a cube or barrel;
oral portion of manubrium conical to cylindrical and mouth margin provided with ring
of nematocysts. Four radial canals are narrow and transparent. Four tentacular bulbs
thick, triangular, without ocelli, with two or three tentacles with a nematocyst-free
base, while the distal region is beaded and end with a terminal knob. Velum broad.
Mesogloea and the radial canals perfectly transparent and uncoloured. Stomach,
marginal bulbs, and tentacular nematocyst clusters contain a brick-red pigment
(observed in living animals).

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 363

OWN OBSERVATIONS: Basal, gonad-free region of the manubrium very short;
large vacuolated cells could not be seen in the examined preserved material; top of the
manubrium is flat, apical canal thus absent; gonad covers most of the manubrium and
leaves only short regions uncovered at both ends. Gonad encircles the manubrium
without interruption. Tentacle bulbs prominent, their epidermal portion bulging, a
gastrodermal chamber present. One sample (26825) mature or almost mature female,
numerous eggs are clearly visible within gonad, but no eggs or embryos attached to
manubrium as in M. haeckelii. Some mature medusae could be mature males. Tentacles
distinctly moniliform, apparently quite short, usually two per bulb. Tentacles contain
only desmonemes and microbasic mastigophores, stenoteles not observed; small
stenoteles seem to be present around mouth opening. Undischarged desmonemes (ca.
15x 10 um) have a thread with characteristic rope structure. Mastigophores have a
spherical capsule of about 10 um diameter.

BioLoGy: Occurs in 200-400 m depth, recorded from April to September, but
only few records are available.

DISTRIBUTION: Norwegian fjords. No type locality was specified, the original
material came from Osterfjord and Herlgfjord, 0-400 m.

REMARKS: Margelopsis hartlaubii Browne, 1903 is a very rare medusa, known
only from a few specimens. Its polyp stage is unknown. The species has been syno-
nymized with Margelopsis gibbesii and M. hartlaubii by Thiel (1938), a proposal that
has not gained acceptance (see remarks under M. haeckelii). It is actually quite distinct
from M. haeckelii: about two times larger, it has a thick mesogloea, it has only two to
three tentacles per bulb, and there is no apical canal. Furthermore, the available
material suggests that there is no brooding in this species and males may exist.

Nevertheless, more data on this species are needed, e.g. detailed nematocyst
data and information on its polyp stage should be obtained in order to confirm the
taxonomic position of this species.

FAMILY PENNARIIDAE MCCRADY, 1859
SYNONYM: Halocordylidae Stechow, 1921.

DIAGNOSIS: Hydroid colony pinnate, occasionally bushy, stem monosiphonic,
giving rise alternately from opposite sides to two series of hydrocladia; hydrocaulus
and hydrocladia with terminal hydranths (monopodial); hydranths on short pedicels on
upper side of the hydrocladia; hydranths pear-shaped; tentacles of two types: in distal
half of hydranth more or less capitate tentacles in one oral whorl and more in indistinct
whorls below, on lower part of hydranth one aboral whorl of semifiliform to slightly
capitate aboral tentacles; gonophores developing above aboral tentacles, eumedusoids,
liberated or not.

Medusa a simple eumedusoid; manubrium not extending beyond umbrella mar-
gin; gonads completely surrounding manubrium; four radial canals; four permanently
rudimentary tentacles, usually reduced to mere bulbs, without ocelli.

REMARKS: The family contains only the genus Pennaria Goldfuss, 1820.

364 P. SCHUCHERT

Genus Pennaria Goldfuss, 1820
TYPE SPECIES: Pennaria disticha Goldfuss, 1820.
SYNONYMS: Globiceps Ayres, 1854; Eucoryne Leidy, 1855; Halocordyle Allman, 1872.

Diacnosıs: With the characteristics of the family.

REMARKS: There are several medusa-based species (Kramp 1959, 1968). They
are mostly indeterminate and some of them do not belong to the genus Pennaria at all
(Petersen, 1990). For the synonymy and validity of the name Pennaria see Calder
(1988) and Gibbons & Ryland (1989). Only Pennaria disticha is relevant for the
European fauna.

Pennaria disticha Goldfuss, 1820 lata ils

Pennaria disticha Goldfuss, 1820: 89; Mayer, 1910: 24, fig. 1A-E; Brinckmann-Voss, 1970: 40,

text-figs 43, 45-50; Gibbons & Ryland, 1989: 387, fig. 5 [taxonomy]; Schuchert, 1996:

142, fig. 85a-c; Watson, 1999: 16, fig. 10A-I; Bouillon et al., 2004: 103, fig. SSA-C.
Pennaria Cavolinii Ehrenberg, 1834: 297; Allman, 1872: 364, fig 80.

Pennaria cavolini — Weismann, 1883: 121, pl. 17 figs 1-5, pl. 18.
Halocordyle disticha — Rees & Thursfield, 1965: 4; Millard, 1975: 41, figs 16C-G; Hirohito,

1977: 2, fig. 1, pls 1-3; Garcia-Corrales & Aguirre, 1985: 85, figs 1-3 [synonymy]; Morri

& Boero, 1986: 31, fig. 11; Wedler & Larson, 1986: 73, fig. 4C; Calder, 1988: 57, figs

43-45 [complete synonymy]; Hirohito, 1988: 28, figs 9a-d, pl. 1 fig. C; Ostman et al.

1991: 607, figs 1-18; da Silveira & Migotto, 1991: 437, fig. 1.

MATERIAL EXAMINED: MHNG INVE29809, Mallorca, Cala Murada, coll. P. Schuchert 24
August, 2000 fertile, depth 1 m, 16S sequence AM088481, 18S sequence AY920762. - MHNG
INVE 36918, Naples, coll. 1892, few hydranths left. - MHNG INVE 36919, Naples, coll. 1900,
fertile. - MHNG INVE 36920, origin unknown, likely Mediterranean, fertile. - MHNG INVE
35468, USA, North Carolina, Morehead City, marine anchorage, fertile, collected 7 October
2000 by Dr Alberto Lindner. - BMNH 1964.8.7.5 Pennaria tiarella, slide preparation, Woods
Hole, Mass. USA, fertile, hydrocladia bearing a single hydranth, stenoteles max. 25 um, thus
larger than in Weill (1934) and within the range observed for P. disticha (Hirohito, 1977). —
Honduras, Utila, Blue Bayou Beach, coll. F. Sinniger 13 February 2004, depth 1 m, hydranths
more delicate than European forms. — Thailand, Andaman Sea, Koh Phi Phi, Ao Nui, 16 April
2000, 10-20 m, collected by Dr A. Faucci. - New Zealand, Hauraki Gulf, Devonport, 26 July
2002, one medusa from plankton. — See also Schuchert (1996, 2003).

DIAGNOSIS: Feather-like branched hydroid colony (pinnate), firm perisarc,
gonophore medusoid, released or not, with four bulbs, no ocelli.

DESCRIPTION: Hydroid colonies forming branching, feather-like shoots, arising
from thick creeping, ramified stolons. Growth monopodial with hydranths on all ends.
Main axis thick, often curved, monosiphonic, perisarc with smooth stretches and
annulated stretches in more or less regular intervals, regularly so distal to insertion of
hydrocladia (side-branches) and hydranth pedicels. Hydrocladia of one side form an
angle of about 120-140 degrees with those from other side, curved, longest hydrocladia
usually found at about 1/3 to 1/2 stem length (measured from base), hydrocladia are
about half as thick as stem, they either bear hydranths on pedicels (ramuli) or
secondary hydrocladia with hydranths again on pedicels. Pedicels (ramuli) of
hydranths originate on upper side of hydrocladia, all approximately of same length and
evenly spaced, with annulation at their base or throughout, younger ones without annu-
lation. Hydranths spindle- to pear shaped, hypostome dome-shaped. Tentacles of two
types: an aboral whorl of about 12-14 long, filiform to slightly capitate tentacles and

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 365

Fic. 15

Pennaria disticha Goldfuss, 1820, after Mediterranean material. A) Colony, modified after
Mayer (1910), actual size. B) Part of branch with two hydranths, one with an almost mature
medusoid, scale bar 0.5 mm. C) Female medusoid that has already spawned some eggs, modi-
fied after Garcia-Corrales & Aguirre (1985), scale bar 0.2 mm.

366 P. SCHUCHERT

up to 16 short capitate tentacles distributed in two to three indistinct whorls on distal
half of hydranth. Knobs of capitate tentacles, except of oral ones, can be reduced to
mere nematocyst cap, thus being almost filiform. Nematocysts of aboral tentacles
mainly on aboral side and terminally (= semifiliform according to Petersen, 1990),
aboral epidermis of filiform tentacles about three times thicker than on oral side,
gastrodermal cells chordoid.

Gonophores oblong medusoids arising on short pedicels just above whorl of
long filiform tentacles. Gonads encircling manubrium, four radial canals and circular
canal present, with four marginal bulbs, with small velum, without ocelli, tentacles
normally absent, rarely some rudiments present. Marginal bulbs without a concen-
tration of nematocysts. Colonies gonochoristic, female medusoids with four to six
eggs. Gonophores can be released as short-lived medusoids, or they spawn while still
attached to hydroid.

Nematocysts of polyp: stenoteles of three to four size classes, microbasic
mastigophores with inclusion body, microbasic heteronemes without inclusion body,
basitrichous isorhizas, desmonemes. Medusoids with stenoteles only.

Colour of stem dark brown to black, hydrocladia brown, fading distally to clear.

DIMENSIONS: Fertile colonies usually around 8-15 cm high, but may excep-
tionally reach 40 cm. Hydranths 0.9-1.5 mm high; medusoids 0.7-1.1 mm. More
measurements are given in Garcfa-Corrales & Aguirre (1985). Nematocyst sizes can be
found in Millard (1975), Hirohito (1977), Garcia-Corrales & Aguirre (1985), Calder
(1988), da Siveira & Migotto (1991), Ostmann et al. (1991), Schuchert (1996), Watson
(1999). Some of these authors also figure them.

BroLOGY: Occurs usually in shallow waters of a few meters depth along rocky
coasts with some wave action. In the Mediterranean, the colonies are active from
spring to fall, the colonies overwinter as stolons (Brinckmann-Voss, 1970; Morri &
Boero, 1986). In more tropical waters they can also be present all year round.
Spawning is induced by a reduction of the light intensity (sunset) (Baker, 1936;
Brinckmann-Voss, 1970; Calder, 1988; Genzano & Kubota, 2003). The eggs develop
in the open water.

Sometimes, the colony form can vary quite drastically, the normal pinnate form
can intergrade with a more bushy form (da Silveira & Migotto, 1991). Also Calder
(1988) found that the colony form varies depending on wave exposure: colonies from
sheltered places were more gracile, internodes of both branches and stem were long
and slender, branches and ramuli elongate, while specimens from wave-swept ledges
were small and compact, internodes of caulus and branches were thicker and shorter
and the branches and ramuli relatively shunted. However, the annulation did not differ
significantly.

Prey capture and nematocyst function were investigated by e. g. Clark & Cook
(1986); Östman er al. (1991), and Kem & Östman (1992). The feeding biology was
examined by Pardy et al. (1968, as Pennaria tiarella).

This species has been the subject of a considerable number of experimental and
developmental studies (sometimes under the synonym Pennaria cavolinii). Rege-
neration studies have been made by Cerfontaine (1902), Gast & Godeeski (1903),

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 367

Brinckmann-Voss (1970), Tardent (1963, 1965). Hydranth development is described
and depicted in Berrill (1952). Gonophore development is documented by Weismann
(1883), Berrill (1952), and Garcia-Corrales & Aguirre (1985). Gametogenesis was
examined by Weismann (1883). Initially, there are many oogonia in female
gonophores, but only a few grow to full size and the others are phagocytosed.

There are many reports on the early development and ultrastructure based on
populations from the USA, either named Halocordyle disticha or Pennaria tiarella
(e. g. Hargitt, 1899, 1900, 1904; Cowden 1964, 1965a, 1965b; Summers & Haynes,
1969; Summers, 1970, 1976; Lesh-Laurie, 1976; Thomas et al., 1987; Martin, 1980,
1987, 1988a, 1988b, 1990, 1991, 1992a, 1992b, 2000; Martin & Archer, 1986a, 1986b,
1997; Martin & Thomas, 1977, 1981a, 1981b, 1983; Hotchkiss et al., 1984; Brumwell
& Martin, 1996).

DISTRIBUTION: Circumglobal in warm temperate to tropical waters. The occur-
rence at the Azores (Cornelius, 1992) and the Strait of Gibraltar (Medel & Lépez-
Gonzalez, 1996) seem to mark the northern limit for the eastern Atlantic. In the North-
Eastern Atlantic also known from the Cape Verde Islands (Rees & Thursfield, 1965),
Madeira (Wirtz & Debelius, 2003). Widespread in the western Mediterranean (e. g.
Brinckmann-Voss, 1970; Garcia-Corrales & Aguirre, 1985; Bouillon et al., 2004) and
eastern Mediterranean (e. g. Vervoort, 1993). Often recorded from the western Atlantic
(e. g. Vervoort, 1968; Hirohito, 1977; Wedler & Larson, 1986; Calder, 1988; Migotto,
1996). The distribution further includes the Red Sea (Hirohito, 1977; Vervoort, 1993),
Indian Ocean (e. g. Ritchie, 1910; Jarvis, 1922; Stechow, 1925; Mammen, 1963;
Millard & Bouillon, 1973; Millard, 1975; Watson, 1999), Malayan Archipelago (Pictet,
1893; Schuchert, 2003), North-Eastern Pacific (Hargitt, 1927; Hirohito, 1977; 1988);
south-western Pacific (Schuchert, 1996; Watson, 1999), central Pacific (Cooke, 1977;
Gibbons & Ryland, 1989; Kirkendale & Calder, 2003) and western Pacific (Fraser,
1938; 1948; Calder et al., 2003). Type locality: Gulf of Naples, Mediterranean.

REMARKS: Pennaria disticha is a conspicuous animal and quite well known. For
the European fauna, there are no serious taxonomic problems. Its morphological
variability was certainly the main reason for its complicated taxonomic history. The
complete synonymy is not given here, as this has been done by Calder (1988), Hirohito
(1977), Garcia-Corrales & Aguirre (1985), and Gibbons & Ryland (1989).

Most Pennaria species based on the hydroid phase were synonymized with
P. disticha, with the prominent exception of P. wilsoni (Hirohito, 1988; Watson, 1999).

Pennaria tiarella (Ayres, 1854), a species originally described from Long
Island (New York), was already considered to be closely related to P. disticha by Mayer
(1910). Most subsequent authors (e. g. Fraser, 1944), however, kept it separate. Also
Brinckmann-Voss (1970) held it distinct from P. disticha on account of the variable
length of the ramuli. Later authors (see above) included also P. tiarella in the
synonymy of P. disticha. Weill (1937) reported a comparatively small size for the
stenoteles of P. tiarella, but an examination of material from the same locality (BMNH
1964.8.7.5, Woods Hole) did not confirm this and the stenoteles have a maximal size
that lies within the range found in other populations of P. disticha (see Hirohito, 1977).

The embryonic development to the planula larva has been examined in great
detail by American students using animals from the USA (identified as P. tiarella or P.

368 P. SCHUCHERT

disticha). Surprisingly, only few such investigations have been made using
Mediterranean animals and there is an unresolved discrepancy. Cavolini (as cited in
Weismann, 1883) observed that the fertilized eggs become encapsulated and over-
winter as resting stages. This has never been observed again and perhaps Cavolini’s
observations can be attributed to the simple observation tools available at his time.
Whatever, this should be re-investigated.

FAMILY CLADOCORYNIDAE ALLMAN, 1872

DIAGNOSIS: Hydroid colonial, simple stolonal or sparingly branched, hydranth
club-shaped, one whorl of moniliform or capitate oral tentacles, aboral tentacles
moniliform or branched capitate, scattered or in several whorls; with nematocysts on
body wall arranged in conspicuous pads or scattered around the base of oral and aboral
tentacles; gonophores carried singly or on short, branched pedicels on lower or middle
part of hydranth; gonophores developing into free medusae or fixed sporosacs.

Medusa with two opposite perradial tentacles possessing stalked cnidophores,
bell margin with four bulbs, of which two are without tentacles; above each atenta-
culate marginal bulb an exumbrellar pad containing macrobasic euryteles; tentaculate
marginal bulbs large, without nematocyst pads; gonads interradial on manubrium.

REMARKS: The concept and relationships of the family Cladocorynidae are
outlined by Petersen (1990). It comprises the genera Cladocoryne Rotch, 1871 and
Pteroclava Weill, 1931.

KEY TO THE GENERA:

la polyps with capitate oral tentacles; gonophores are fixed sporosacs . ..... Cladocoryne
lb polyp with moniliform oral tentacles; gonophores are free medusae ................
RR a SONO peas eee Pteroclava (not in European fauna)

Genus Cladocoryne Rotch, 1871

TYPE SPECIES: Cladocoryne floccosa Rotch, 1871 by monotypy.

SYNONYMS: Polycoryne Graeffe, 1883a; Cladocorynopsis Mammen, 1963; Lobocoryne
Mammen, 1963.

DIAGNOSIS: Cladocorynidae with long hydocauli, these unbranched or sparingly
branched, covered by perisarc. Hydranth club-shaped, with oral whorl of capitate
tentacles, one to four whorls of branched-capitate aboral tentacles; on hydranth body
usually nematocyst patches containing macrobasic euryteles; gonophores fixed
sporosacs between or over aboral tentacles.

Cladocoryne floccosa Rotch, 1871 Fig. 16

Cladocoryne floccosa Rotch, 1871: 228; Du Plessis, 1880: 176, pl. 9; Allman, 1872: 380, fig. 82;
Kühn, 1910: 69, fig. D, pl. 6 figs 28-34, pl. 7 figs 35-36; Kühn, 1913: 184, figs 28-36,
pls 6-7; Philbert, 1936: 1, figs 1-8; Weill, 1937: 1, figs 1-5; Brinckmann-Voss, 1970: 69,
figs 80-82; Millard & Bouillon, 1974: 11, fig. 1D-E; Millard, 1975: 65, figs 21A-B;
Bouillon et al., 1987: 297, figs 1 & 5-6; Hirohito, 1988: 52, figs 16 b-f; Migotto, 1996:
17, fig. 4a-b; Schuchert, 1996: 97, fig. 57a-c; Watson, 1999: 11, fig. 7A-D; Pefia Cantero
& Garcia Carrascosa, 2002: 46, fig. 9e; Calder et al., 2003: 1178, fig. 3; Bouillon et al.,
2004: 110, fig. 57G.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 369

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Cladocoryne floccosa Rotch, 1871; A-D after preserved material from the Mediterranean; E-F,
after Kühn (1910). A) Hydroid with sporosacs, scale bar 0.5 mm. B) Slightly compressed
microscopic preparation of a hydranth, note position of eurytele clusters (arrows), scale bar 0.2
mm. C) Part of tentacles of aboral set, note chordoid gastrodermis, scale bar 0.1 mm. D)
Macrobasic eurytele, scale bar 40 um. E) Histological longitudinal section of an almost mature
female sporosac. F) Longitudinal section of a mature male sporosac.

370 P. SCHUCHERT

Hydra corynaria Bosc, 1797.

Cladocoryne pelagica Allman, 1876: 255, pl. 10 figs 6-7.

Polycoryne helleri Graeffe, 1883a: 202, pl. figs 1-5; Graeffe, 1883b: 320, synonym.
Cladocoryne simplex Perrier, 1886: 81, fig. 3.

Cladocoryne floccosa var. sargassensis Hargitt, 1909: 369, figs 1-2.

Cladocoryne sargassensis — Kingsley, 1910: 19, fig. 81.

not Cladocoryne floccosa — Rees & Thursfield, 1965: 46 [= C. haddoni Kirkpatrick, 1890].

MATERIAL EXAMINED: Cala Murada, Mallorca, Spain, June 1997, on rock in 0.5-2 m
depth, fertile, examined alive. - MHNG INVE 29808, Cala Murada, Mallorca, 1 m, 13 August
2000, on Peysonnellia, examined alive, infertile. - BMNH 1974.11.21.38, Italy, Naples, 20 April
1967, material of Brinckmann-Voss (1970), not well preserved (maceration by formalin). - MH-
NG INVE 36913 Naples, Cap Misene 23.01.1895, young sporosacs present, on various sub-
strata. - MHNG INVE 36912, Villefranche-sur-Mer, infertile.

DIAGNOSIS: Cladocoryne species with three to four whorls of aboral tentacles,
patches of macrobasic euryteles among oral and aboral tentacles, gonophores without
radial canals, mature females with one to two eggs.

DESCRIPTION (after own material and literature): Hydroid with unbranched stem
or rarely branched once, arising from adhering, ramified stolons. Stems covered by
perisarc, smooth or with annulated stretches and becoming thinner distally to terminate
below hydranth. Hydranth cylindrical to club-shaped, dome-shaped hypostome; with
tentacles of two types in two well-separated sets: a single whorl of four to six short
capitate oral tentacles, and 12-22 aboral tentacles in three to four alternating whorls,
whorls can be indistinct. Aboral tentacles branched, side-branches short and capitate,
in two lateral rows and one median row on upper side; lateral secondary tentacles up
to seven per row, median row with zero to two. All side-branches and end of main
branch with a terminal spherical nematocyst cluster. Sometimes some secondary
tentacles without stalk and reduced to mere nematocyst clusters. Gastrodermis of
tentacles chordoid. Hydranth with nematocyst patches or pads on body wall between
oral tentacles and lowest aboral tentacles. Colours: hydranth light reddish brown, oral
region with conspicuous white pigment.

Gonophores on short pedicels, two to eight borne on the hydranth among or
above the upper aboral tentacles, spherical to oblong, remaining fixed as sporosacs
without radial canals or circular canal (cryptomedusoid type), females usually with one
egg only, colour white.

Nematocysts: stenoteles and macrobasic euryteles.

DIMENSIONS: Colony height a few mm to 12 mm; caulus diameter 0.15 mm;
hydranth height 1-1.2 mm, diameter 0.3 mm; length aboral tentacles tentacle 0.7-
1.3 mm; sporosac size: 0.3-0.4 mm. Nematocyst sizes are given in Philbert (1936);
Weill (1937); Millard & Bouillon (1974); Millard (1975); Migotto (1996); Schuchert
(1996); Watson (1999).

BioLoGY: Cladocoryne floccosa has been reported from depths ranging from 0
to 50 m. It has often been observed on Sargassum weeds, but also occurs on many other
solid substrata like algae, hydroids, sponges, and rocks. In the Mediterranean, it can be
found from January to autumn, fertile colonies have been observed in January and from
end of April to July (Brinckmann-Voss, 1970; Boero & Fresi, 1986; own observations).

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 371

ADDITIONAL DATA: Fertile animals often reduce their tentacles, sometimes
leading to tentacle-less hydranths (reproductive exhaustion). Kiihn (1910) describes
the formation of the gonophores and the gametogenesis. The mature female sporosacs
usually contain one egg only. Initially there are many eggs, but during the maturation
one of them phagocytoses all the others. The embryonic development takes place in the
sporosac. Philbert (1936) depicts the metamorphosis of the larva to the primary polyp.
The nematocysts are depicted in Weill (1937). A longitudinal section of a hydranth is
depicted Prévot (1959) and in Bouillon et al. (1987, fertile hydranth). Pefia Cantero &
Garcia Carrascosa (2002) found branched stems that had up to four hydranths.

DISTRIBUTION: Circumglobal in temperate to tropical waters. Along the
European coasts known from north-west of Ireland (Allman, 1876, as C. pelagica), the
English Channel (Rotch, 1871; Philbert, 1936; Teissier, 1965), the Iberian Peninsula
(Da Cunha, 1944; Medel & Löpez-Gonzälez, 1996), western Mediterranean (Du
Plessis, 1880; Motz-Kossowska, 1905; Brinckmann-Voss, 1970; Pefia Cantero &
Garcia Carrascosa, 2002; and more), eastern Mediterranean (Picard, 1958; Morri &
Bianchi, 1999). Also known from the western Atlantic (Hargitt, 1909 as Cladocoryne
floccosa var. sargassensis; Migotto, 1996); the Indian Ocean (Millard & Bouillon,
1974; Watson, 1999); the western Pacific (Hirohito, 1988; Schuchert, 1996); the
eastern Pacific (Lees, 1968; Calder et al., 2003). Type locality: on stones at low tide at
Herm, Guernsey, Channel Islands, United Kingdom.

REMARKS: This is a characteristic species that cannot be confounded with any
other hydroids of the European fauna. The synonymy is quite well established.
Polycoryne helleri Graeffe, 1883a was recognized as a synonym by Graeffe (1883b)
himself. Cladocoryne pelagica Allman, 1876 was described based on a fertile colony
found on floating Sargassum seaweed northwest of Ireland. Allman kept it distinct on
account of its smaller size and the annulated base of the pedicel. These traits are no
longer considered diagnostic and in his revision, Philbert (1936) regarded it as a syn-
onym of C. floccosa, likewise also C. simplex Perrier, 1886, and C. haddoni
Kirkpatrick, 1890. Most subsequent authors took up Philbert's conclusion, but Bouillon
et al. (1987) re-described C. haddoni based on material from Papua New Guinea and
found sufficient differences to C. floccosa to permit a reliable identification, even of
immature specimens. Cladocoryne haddoni was therefore recognized as a distinct
species. Cladocoryne pelagica, C. simplex and C. sargassensis are very likely conspe-
cific with C. floccosa, although the original descriptions are usually not detailed
enough. A re-examination of the type material of C. pelagica and a detailed exami-
nation of Cladocoryne hydroids on Sargassum would therefore be helpful. Calder ef al.
(2003) discovered another likely synonym of C. floccosa, namely Hydra corynaria
Bosc, 1797, a Cladocoryne species that was also originally described based on material
growing on Sargassum. Because Bosc's name has not been used since its original
description Calder et al. (2003), based on provisions of the ICZN, invalidated Bosc's
name and to declared Cladocoryne floccosa Rotch, 1871 a protected name.

FAMILY CLADONEMATIDAE GEGENBAUR, 1856

SYNONYMS: Cladonemiden Gegenbaur, 1856; emended to Cladonematidae by Poche
(1914: 70); Stauriidae Hincks, 1868; Dendronemidae Haeckel, 1879; Eleutheriidae Stechow,
1923.

372 P. SCHUCHERT

REFERENCES: Giinther (1903), Hartlaub (1887), Stechow (1923), Bouillon (1985), Calder
(1988), Petersen (1990), Bouillon & Boero (2000).

DIAGNOSIS: Hydroid colony stolonal or rarely sparingly branched, arising from
creeping stolons. Hydranth spindle-shaped, one whorl of four to ten solid oral capitate
tentacles, with or without aboral whorl of filiform tentacles; mouth opening into a pre-
oral chamber formed by epidermal gland cells; medusa buds not enclosed in periderm
film, carried singly or in clusters at base of hydranth or above filiform tentacles if they
are present.

Medusa adapted to benthic mode of life, some species still able to swim freely;
with or without a thickened continuous or broken ring of nematocysts around umbrel-
lar margin, with or without brood-chamber above manubrium; manubrium cylindrical,
with or without protruding pouches; mouth either simple, or with oral nematocyst
clusters, or with ramified oral tentacles; gonads either completely surrounding manu-
brium, or on manubrial pouches, or in special brood-chamber above manubrium, in
latter case also sometimes additionally in subumbrellar epidermis; number of radial
canals variable but usually more than four, some may be branched, final number of
canals entering circular canal usually corresponding or slightly exceeding the number
of marginal tentacles; marginal tentacles hollow, branched, with branches bearing
nematocyst clusters and branches having adhesive organs; with abaxial ocelli on
tentacle base.

REMARKS: As some Cladonematidae can be cultivated quite easily, they have
become the subject of numerous developmental and molecular studies. They thus
became quite well known even to non-specialists. The European Cladonematidae
species present few taxonomic problems and the species are mostly well known,
though the genus Dendronema remains problematic. However, taxonomic problems
persist at the genus level.

The family Cladonematidae has been kept separate from the Eleutheriidae
Stechow, 1923 by a number of authors (for more details see Calder, 1988). The latter
family comprised the genera Staurocladia and Eleutheria. The separation of both
families is ambiguous and not necessary and the proposal of Petersen (1990) to unite
them is here also adopted. A good example in favour of this is Staurocladia portmanni
Brinckmann, 1964, which is perfectly intermediate between Staurocladia and
Cladonema, differing basically only in the branching of the tentacles. The genera
Cladonema, Staurocladia, and Eleutheria are linked by a number of synapomorphic
characters, such as the preoral chamber in the hydroid, the branched medusa tentacles
with adhesive ends, and the number of radial canals (Petersen, 1990).

In current usage (Kramp, 1961; Bouillon, 1985; Bouillon & Boero, 2000), the
family comprises four genera: Cladonema, Dendronema, Staurocladia, and Eleutheria.
Dendronema is somewhat problematic and the distinction of the three remaining
genera relies principally only on the morphology of the capitate tentacles of the
medusa. Cladonema has tentacles that are branched more than once and its umbrella is
not reduced. In Staurocladia, the tentacles are only bifid and the upper branch bears
additional nematocyst clusters besides the terminal one. Eleutheria was diagnosed as
having bifid tentacles with a single terminal nematocyst cluster, and additionally a

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 373

brood-chamber above the manubrium. There remain, however, problems with this clas-
sification, especially so for the scope of Staurocladia.

Hartlaub (1917) established the genus Staurocladia solely to separate
Eleutheria vallentini, E. claparedii, and others from Eleutheria dichotoma. He found
this necessary to account for some of the peculiarities of E. dichotoma: the brood-
chamber, the hermaphroditism, and the gonostyle in the polyp phase. Hartlaub there-
fore used the combination Staurocladia claparedii (Hartlaub, 1889), as this species has
no brood-chamber (Miiller, 1911; Lengerich, 1923a). Lengerich (1923a) used only a
single genus, Eleutheria, for all Cladonematidae. Although this would resolve the
problem of paraphyletic genera, it creates unwanted name changes for widely known
species. Probably unaware of Hartlaub’s publication, Gilchrist (1919) had similar
thoughts and suggested the genus name Cnidonema for the Eleutheria species of the
southern hemisphere. He also thought that it might be necessary to introduce a new
genus for E. claparedii if this species should have no brood-chamber. Browne &
Kramp (1939) and Kramp (1959, 1961, 1968) took up Hartlaub’s diagnosis, but
modified the definition so that Staurocladia was differentiated from Eleutheria by the
number of nematocyst clusters per tentacle (Eleutheria one, Staurocladia more than
one) and the absence or presence of a brood-chamber. Browne & Kramp (1939) placed
E. claparedii in the genus Eleutheria, because they were probably unaware that it has
no brood-chamber. Brinckmann-Voss (1970) became aware of this discrepancy and
changed the genus diagnosis of Eleutheria accordingly to "with or without brood-
chamber above manubrium". The genera Staurocladia and Eleutheria were hence
distinguished only on account of the number of nematocyst clusters. However, later
authors reverted to Kramp's (1961) diagnosis (e. g. Bouillon, 1985; Petersen, 1990;
Bouillon & Boero, 2000), but were inconsistent in placing E. claparedii in the genus
Eleutheria.

Distinguishing a genus solely on the number of tentacular nematocyst clusters
appears quite ambiguous and very prone to give polyphyletic assemblages. It is thus
quite unlikely that the present genera correspond to a monophyletic groups.
Unfortunately, there are few useful characters for a phylogenetic analysis of the
members of the Cladonematidae. Synapomorphies that can be used to find species
groupings are given in table 1. There are more apomorphies, but either they are
autapomorphies (gonostyle, very extensible hydranth, brood-chamber, centripetal
canals, radial canals with processes, apical cavity), or only examined for one or few
species (desmonemes in polyp stage, heteronemes in polyp stage). Some characters are
not really usable (lateral or aboral position of nematocyst clusters).

Using these characters for a cladistic analysis of representative members of the
family was not very helpful (table 1, figure 17). There are several unresolved poly-
tomies, but it is worth mentioning that Staurocladia is evidently a polyphyletic group.
In this case, it seems nevertheless prudent not to use these results to redefine the gen-
era because taxonomic stability should be given more value than a classification based
on a poorly resolved cladogram. Perhaps a thorough molecular analysis can provide the
necessary data, but until then, the diagnoses for Eleutheria and Staurocladia as given
in Brinckmann-Voss (1970) should be used.

P. SCHUCHERT

374

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THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 375

Dipurena reesi

Dipurena strangulata
Cladosarsia minima
Staurocladia portmanni
Staurocladia wellingtoni
Cladonema radiatum
Dendronema stylodendron
Eleutheria dichotoma

Eleutheria claparedii

Staurocladia vallentini

Fic. 17
Genera of the Cladonematidae: Phylogenetic analysis of morphological characters using
maximum parsimony: Strict consensus tree of the 17 trees with minimal length (18 steps,
HI=0.28, CI=0.72, RI=0.81). The consensus tree of the bootstrap analysis was identical, the
percentage of node support is given in boxes. Some selected synapomorphies are also given
(numbers along branches, see table 1).

KEY TO THE CLADONEMATIDAE:

la medusatentacles branched more than once Re N n 2
1b MEdUSAENACIESIDrANChE ONCE ms rs ne. OR N OOO TO 3
2a Oral nematocyst knobs of medusa simple. M i. Cladonema
2b oral nematocyst knobs of medusa branched ........................ Dendronema
3a medusa tentacles with a single nematocyst knob ...................... Eleutheria
3b medusa tentacles with a several nematocyst knobs ................... Staurocladia

Genus Cladonema Dujardin, 1843

TYPE SPECIES: Cladonema radiatum Dujardin, 1843 by monotypy.
SYNONYMS: Stauridia Forbes, 1848; Stauridium Krohn, 1853b.

REFERENCES: Kramp (1961); Bouillon (1985); Calder (1988); Petersen (1990); Bouillon
& Boero (2000).

DraGNOSIS: Hydroid mostly stolonal, occasionally branched, one oral whorl of
four to five capitate tentacles, with or without filiform tentacles, medusa buds borne
singly on hydranth body immediately above filiform tentacles or in similar position
when these are absent. Medusae adapted for swimming and adhering to surfaces, with
bell-shaped umbrella; manubrium cylindrical, in adults with pouches, gonads without
perradial separations, mouth short, unbranched lips bearing four to six spherical
nematocyst clusters; without apical chamber above manubrium; radial canals bifur-
cated or simple, final number of canals entering circular canal usually of same number

376 P. SCHUCHERT

as marginal tentacles; marginal tentacles branching, each with one to six proximal
branches ending in an adhesive organ and one to ten more distal branches with nema-
tocyst clusters; abaxial ocelli with lens.

Cladonema radiatum Dujardin, 1843 Fig. 18

Cladonema radiatum Dujardin, 1843: 1134; Dujardin, 1845: 271, pl. 14 figs C1-C7, pl. 15 figs
C8-C19; Krohn, 1853b: 420, pl. 13; Keferstein & Ehlers, 1861: 85, pl. 13 fig. 5; Van
Beneden, 1867: 139, pl. 12; Hincks, 1868: 62, text fig. 5, pl. 11; Allman, 1871-1872: 216,
pl. 17; Hincks, 1872: 391, pl. 21, fig. 6; Hartlaub, 1887: 266, 651; Billard, 1905: 500;
Hartlaub, 1907: 132, figs 123-126; Müller, 1908: 30, pl. 3 figs 1-2; Mayer, 1910: 99, figs
53-55; Lengerich, 1923a: 331, figs A-S; Weill, 1937: 443, fig. 1; Russell, 1953: 105, figs
49-51; Kramp, 1959: 96, fig. 55; Brinckmann & Petersen, 1960: 388, fig. 2; Kramp,
1961: 57; Kramp, 1968: 22, fig. 52; Naumov, 1969: 245, figs 113, 117A-H; Brinckmann-
Voss, 1970: 76, figs 88-89, pl. 5 figs 1-2; Bouillon, 1971: 333, figs 4.5-7; Millard &
Bouillon, 1973: 12, fig. 1A; Rees, 1979: 300; Calder, 1988: 67, fig. 50; Hirohito, 1988:
41, fig. 12a-b; Schuchert, 1996: 131, fig. 80a-d; Migotto, 1996: 23, fig. 4e-g; Bouillon er
al., 2004: 88, fig. 48B-D.

Coryne stauridia Gosse, 1853d: 257, pl. 16 figs 1-5.

Syncoryne stauridium Krohn, 1853a: 137.

Cladonema Gegenbauri Haeckel, 1879: 109.

Cladonema Krohnii Haeckel, 1879: 109.

Cladonema Dujardinii Haeckel, 1879: 109.

Cladonema Allmani Haeckel, 1879: 109.

Stauridium cladonema Haeckel, 1879: 109.

Cladonema perkinsii Mayer, 1904: 18. pl. 4 fig. 35; Mayer, 1910: 101, pl. 9 fig. 1.

Cladonema mayeri Perkins, 1906: 118; Mayer, 1910: 101, pl. 9 figs 2-3.

Stauridia radiatum — Mayer, 1910: 100.

Eleutheria radiata — Lengerich, 1922: 210, fig. 1; Lengerich, 1923a: 313, figs G-S.

? Cladonema novae-zelandiae Ralph, 1953: 72, fig. 20.

MATERIAL: Aquarium culture of polyp stage originating from the Mediterranean,

18 January 1991, life-cycle observed to mature medusa. - MHNG INVE 29909, Roscoff,
Brittany, polyps on holdfast of laminarian, collected 1 June 2001, cultured at room temperature,
medusae buds developed, very cold-sensitive, died at <14°, grew well at 22-30°C; 16 S sequence
accession number AY512539. — Three medusae, Bay of Portoferraio, Island of Elba, Italy, col-
lected July 2005 by dragging a plankton net over a Posidonia meadow, depth 2-4 m; colour of
whole medusa light brown, 16 S sequence accession numbers of two different individuals
AM088482 and AM088483. — MHNG INVE 37640, 19 February 2004, polyps on "living
stones" from tropical aquarium, origin unknown, medusae cultivated to near maturity, 16 S se-
quence accession number AM088484, appeared identical to other C. radiatum medusae except
that proximal parts of radial canals were white, also on manubrium longitudinal stripes of white
pigment; this material belongs thus perhaps to a separate species of tropical origin, a fact also
suggested by the deviating 16S sequence.

DIAGNOSIS: Hydroid with one oral whorl of capitate tentacles, each with up to
ten gastrodermal cells, with one aboral whorl with usually four filiform tentacles.
Medusa tentacles branched several times, two and more adhesive branches in fully
mature animals, two and more capitate branches; manubrium shorter than bell cavity,
mouth with four to five spherical nematocyst clusters, gonads on manubrial pouches
and on manubrium, radial canals bifurcated or not.

DESCRIPTION: Hydroid colonies stolonal or rarely branched, arising from
creeping, ramified stolons, hydranths on pedicel of variable length, but usually longer
than hydranth. Perisarc smooth, terminating shortly below filiform tentacles.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 377

Fic. 18

Cladonema radiatum Dujardin, 1843. A-C, after living material from Brittany; D, after photos
of living Mediterranean medusae. A) Polyp with medusa bud, scale bar 0.2 mm. B) Lateral view
of newly released medusa in optical section, same scale as A. C) Oral view of newly released
medusa. D) Mature male medusa, scale bar ca. 1 mm.

378 P. SCHUCHERT

Hydranths with one oral whorl of four to five capitate tentacles, each tentacle with
seven to ten chordoid gastrodermal cells. Below capitate tentacles one whorl of four
(occasionally five) slender, filiform tentacles with a slight terminal swelling, without
nematocysts. Hypostome with an epidermal preoral chamber. Medusae buds naked,
arising above filiform tentacles. Nematocysts: stenoteles and mastigophores.

Newly released medusa with eight to ten bifid tentacles, upper branch with a
terminal nematocyst cluster and a few oral-aboral clusters (Fig. 18).

Adult medusa with bell-shaped umbrella, slightly higher than wide, jelly
moderately thin, sometimes with a slight apical projection, velum rather broad. Medusa
able to swim freely but mostly remains attached, does not move by crawling.
Manubrium spindle-shaped, shorter than or as long as bell cavity, with five (sometimes
four or six) perradial pouch-like protuberances in its middle region, protuberances
rather shallow, size variable. The gonads completely surround the upper two thirds of
manubrium inclusive the pouches. Mouth with four to five bulbous nematocyst
clusters. Radial canals may bifurcate close to the manubrium and eight to ten canals
reach the circular canal, branching pattern variable even in individuals from the same
colony. Number of marginal tentacles corresponds to the number of radial canals. Each
tentacle base with an abaxial ocellus. Marginal tentacles branched, with elongated
thickened base and a long main branch from the underside of which grow one to six
short tentacles acting as adhesive organs, the ends of the adhesive tentacles have a dis-
tinct spherical swelling at their end; distal to the adhesive branches the main branch
gives off up to five side-branches bearing nematocyst clusters alternating in aboral and
oral position and one larger terminal cluster, proximal side-branches originate orally,
more distal ones issuing laterally from the main branch. Colour of stomach and mar-
ginal tentacles red, bright-red or brown; ocelli black or deep crimson. Nematocysts:
stenoteles, desmonemes.

DIMENSIONS: Hydroid colonies up to 25 mm, hydranth about 1 mm; newly
released medusa 0.7-1.0 mm, adult medusa up to 3 mm high. Nematocysts see Weill
(1936), Brinckmann & Petersen (1960), Schuchert (1996).

BroLocy: For Mediterranean populations, Brinckmann- Voss (1970) reports that
the polyp colonies remain small in nature (2-3 hydranths), they were found in shaded
placed on algae and other hard substrata. From end of June to the end of July, the
medusae were caught in large numbers in Posidonia beds, later they are rare or absent.
The development of the medusae-buds is quick, at 18°C it takes only 6-7 days until the
medusa is set free. Also Motz-Kossowska (1905) observed the medusae only from June
to August, rarely also in October.

ADDITIONAL DATA: The polyps of Cladonema radiatum are easy to keep in lab-
oratory. Dujardin (1843) kept them for several years, Lengerich (1923a) reports a
colony that was kept for more than 30 years in an aquarium in Berlin. Also the medusa
is relatively easy to keep and reaches maturity in culture vessels. It also turns up regu-
larly in aquariums. It is thus an ideal experimental organism and therefore there is a
large number of developmental and histological investigations which cannot be re-
viewed here in detail.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 379

The morphology, histology, and ultrastructure was dealt with in detail by e. g.
Miiller (1908); Lengerich (1923a; 1923b), Brien (1942), Bouillon (1968), Bouillon &
Houvenaghel (1970), Tardent & Stôssel (1971), Weber & Tardent (1978), Weber (1980,
1981a, 1981b), Bouillon et al. (1988), Falugi et al. (1994).

The nematocysts and their biology were investigated by: Weill (1936), Brien
(1942), Achermann & Tardent (1973), and Bouillon (1971).

The development and medusae budding was investigated by e. g.: Pasteels
(1939, 1941), Brien (1941, 1942), and Bodo (1970).

The behaviour of the medusa is described by: Allman (1872), Browne (1900),
Billard (1905), Lengerich (1923a), and Russell (1953, summary). It is important to
note in this context that the medusa does not crawl like Eleutheria dichotoma, but
changes place by swimming. The adhesive tentacles are thus only used for clinging to
a surface. There is also a differences in the ultrastructure of the adhesive tentacles
(Bouillon, 1968).

The medusae can be hermaphroditic, but usually this is rare (Hartlaub, 1887;
Miiller, 1908; Bouillon & Houvenaghel, 1970). Bouillon & Houvenaghel (1970) found
only two hermaphroditic animals among fifty medusae, while others observed none
(Lengerich, 1923a). They can be successive hermaphrodites beginning either as
females or males (Hartlaub, 1887).

DISTRIBUTION: North-Eastern Atlantic: Norway (Christiansen, 1972), Great
Britain and Ireland (Hincks, 1868; Allman, 1872; Russell, 1953), Sweden (Segerstedt,
1889; Aurivillius, 1898; Kramp, 1935); Denmark (Kramp, 1935); Helgoland (Richters,
1908); Holland (Vervoort, 1946), Belgium (Leloup, 1947), Atlantic coast of France
(Dujardin, 1843; Billard, 1905; Teissier, 1965), Atlantic coast of the Iberian Peninsula
(Medel & Löpez-Gonzälez, 1996). Mediterranean (numerous records, e. 2.
Brinckmann-Voss (1970), Boero & Fresi (1986), Daly Yahia er al. (2003, Tunisia),
Bouillon er al. (2004)). Present in the Black Sea (Thiel, 1935). Also known from Brazil
(Migotto, 1996), Bermuda (Calder, 1988), Indian Ocean (Millard & Bouillon, 1973), ?
New Zealand (Schuchert, 1996), Japan (Hirohito, 1988; introduced?). Type localities:
Dujardin (1843) described material from the Mediterranean, St. Malo, and Lorient
(France).

REMARKS: The medusa of Cladonema radiatum is quite variable with respect to
the number and branching patterns of the radial canals. Also the number of adhesive
tentacles, manubrial pouches, and oral nematocyst clusters is variable in natural
populations. Even monoclonal animals show some degree of variation.

Weill (1937) examined 106 specimens collected at Bermuda and these encom-
passed the characteristics of C. radiatum, C. perkensii, and C. mayeri. Weill concludes
that the three nominal species were no more than varieties of one rather variable
species, as Mayer (1910) had suggested earlier. This view has been adopted by most
subsequent authors (e. g. Russell, 1953; Kramp, 1959; Kramp, 1961).

There are several nominal Cladonema species known, but sometimes the
differences to C. radiatum are unclear or minimal (Kramp, 1968). Rees (1979) and
Stepanjants et al. (1993) review all the differences between the different nominal
species. Stepanjants et al. (1993) concluded that there were only two valid species,
namely C. radiatum and C. californicum.

380 P. SCHUCHERT

The medusae of Cladonema californicum Hyman, 1947 was held distinct from
all other described Cladonema in having only one adhesive tentacle branch and one to
two nematocyst bearing branches, and a manubrium that protrudes from the velar
opening (Hyman, 1957; Rees, 1979). According to Hyman (1957), the gonads cover
the manubrium on the pouches and the rest of the manubrium, this in contrast to the di-
agnosis in Kramp (1968). The polyp appears indistinguishable from C. radiatum.

The polyps of the Californian Cladonema myersi Rees, 1949 lack filiform ten-
tacles, the medusa lacks branching radial canals, and the gonad covers the entire
manubrium (Naumov, 1969). Likewise, the polyps of the Japanese Cladonema uchidai
Hirai, 1958 also lack filiform tentacles.

Naumov (1955) described Cladonema pacificum, which he later (1960, 1969)
synonymized with C. myersi. This was contested by Hirohito (1988). Rees (1979) and
Hirohito (1988) think that C. uchidai belongs to Cladonema pacificum. This species is
distinguished from C. radiatum by a gonad that covers almost the whole manubrium,
and not only the upper 2/3 (compare Bouillon et al., 1988: fig. 1).

Cladonema novaezelandiae Ralph, 1953 has been synonymized with C. radia-
tum by Schuchert (1996). However the polyp stage of the New Zealandic Cladonema
remains unknown and it may turn out to be different from C. radiatum. Its synonymy
with C. radiatum is therefore not sufficiently established.

Although it is evident that many nominal Cladonema species are currently not
objectively distinguishable, it is still likely that several distinct biological species have
been included in C. radiatum. The 16S sequences of the Cladonema sample found in
aquarium with tropical material (see above in material examined) differs in more than
25% of its positions from the Atlantic and Mediterranean sequences. Such a difference
represents more likely interspecific variation (comp. Schuchert, 2005). The differences
among the examined European sequences are less than 4%.

The hydroid of Cladonema radiatum is rather difficult to distinguish from
hydroids of some Corynidae (Schuchert, 2001b). Brinckmann & Petersen (1960) found
that the number of gastrodermal cells in the capitate tentacles offers a reliable tool to
discriminate Cladonema radiatum and D. reesi: while C. radiatum has 10 or less cells,
D. reesi has 13-18 (see also Schuchert, 2001b). Additionally, the macrobasic
heteroneme is characteristic for Cladonema radiatum. Also the hydroid of Stauro-
cladia portmanni is very similar. Brinckmann-Voss (1970) found that S$. portmanni
polyps have regularly six filiform tentacles, while C. radiatum has usually four, some-
times five.

Genus Eleutheria Quatrefages, 1842

TYPE SPECIES: Eleutheria dichotoma Quatrefages, 1842, by monotypy.

SYNONYMS: Clavatella Hincks, 1861; Herpusa Schmidt, 1869.

REFERENCES: Kramp (1961); Bouillon (1985); Petersen (1990); Bouillon & Boero
(2000).

DrAGNosis: Hydroid with an oral whorl of up to ten capitate tentacles, without
aboral filiform tentacles; medusae buds carried in clusters on short blastostyles (gono-
style) or singly at base of hydranth. Medusae adapted for crawling, unable to swim;
umbrella with or without continuous ring of nematocysts along margin; with four or

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 381

more simple and short radial canals; manubrium with broad base, conical, with simple
circular mouth; with or without brood-chamber above base of stomach; gonads either
on manubrium, or in brood-chamber from where they can also extend to the epidermis
of the subumbrella; 6-14 bifurcated marginal tentacles, one upper branch with a single
terminal nematocyst knob, one lower branch with adhesive organ: asexual repro-
duction through medusa budding present.

REMARKS: The problems concerning the distinction of Eleutheria and
Staurocladia have been discussed above under remarks at the family level.

KEY TO THE EUROPEAN ELEUTHERIA MEDUSAE:

la medusa buds on outside, with brood-chamber ............... Eleutheria dichotoma
lb medusa buds in subumbrella, no brood-chamber .............. Eleutheria claparedii
Eleutheria dichotoma Quatrefages, 1842 Figs 19A-C, 20

Eleutheria dichotoma Quatrefages, 1842a: 270, pl. 8; Quatrefages, 1842b: 168; de Filippi, 1866:
375, pl. 1; Hartlaub, 1886: 706, text. fig.; Hartlaub, 1907: 127, figs 119-120; Mayer,
1910: 94, figs 46-48; Miiller, 1908: 34, pl. 3 figs 3-7, pl. 4; Lengerich, 1923a: 359, figs
R'-Z', D2; Drzewina & Bohn, 1913: 49, figs 26-37; Weill, 1936: 816; Russell, 1953: 110,
figs 52-53; Kramp, 1961: 59; Naumov, 1969: 249, figs 118-121; Brinckmann-Voss,
1970: 79, figs 90-92; Bouillon, 1994: 146, fig. 55; Bouillon, et al. 2004: 88, fig. 48B-D.

Clavatella prolifera Hincks, 1861: 73, pls 7-8; Hincks, 1868: 73, pl. 12 fig. 2; Allman, 1872: 384,
pl. 18.

Herpusa ulvae O. Schmidt, 1869: 993, fig.

Eleutheria tetranema Haeckel, 1879: 106.

Eleutheria pentanema Haeckel, 1879: 106.

Eleutheria hexanema Haeckel, 1879: 106.

Eleutheria anisonema Haeckel, 1879: 106.

Eleutheria heptanema Haeckel, 1879: 107.

Eleutheria octonema Haeckel, 1879: 107.

Eleutheria dichocnida Haeckel, 1879: 107.

Eleutheria cnidobdella Haeckel, 1879: 107.

Eleutheria dichobdella Haeckel, 1879: 107.

Eleutheria heteroclada Haeckel, 1879: 107.

Cladonema prolifera — Motz-Kossowska, 1905: 47.

Eleutheria krohni Krumbach, 1907: 453.

Eleutheria gemmipara Du Plessis, 1909: 376.

not Eleutheria dichotoma — Claparède, 1863: 4, pl. 1 figs 4-10 [= E. claparedii].

not Eleutheria dichotoma — Spagnolini, 1876: 312 [= E. claparedii].

MATERIAL: MHNG INVE 34228, Mediterranean, Banyuls-sur-Mer, île Grosse, 2 May
2002, 0 m, > 10 medusae on Ulva lactuca, with eggs/planulae in brood-chamber, examined alive,
development to primary polyp followed, DNA sample deposited, 16S sequence accession
number AM088485. - MHNG INVE36841 living polyp colony obtained from B. Schierwater,
collected 1995, Spain, Tossa de Mar, La Fosca, medusae buds developed after a few days of
cultivation.

DIAGNOSIS: Eleutheria medusa with brood-chamber, medusae buds outside on
bell rim or exumbrella, normally six tentacles, tentacles bifurcated once in middle,
trunk and branches of about the same length, often hermaphroditic, nettle ring thick.

DESCRIPTION: Hydroid stolonal, stolons creeping, branched, perisarc smooth;
hydranths with very short pedicels, sometimes almost sessile, pedicel covered by thin
periderm, sometimes widened like a funnel. Hydranth body cylindrical, very extensile,
fully extended with very thin body and swollen terminal region, one oral whorl of four

382 P. SCHUCHERT

to eight capitate tentacles, 16-18 gastrodermal cells per tentacle. Filiform tentacles
absent. Medusae buds in lowest part of hydranth, either singly or in clusters borne on
short gonostyle (tubular outgrowth of body wall, up to four per hydranth), gonostyles
branching or not, carrying up to four medusae buds, buds not protected by periderm
covering. Preoral cavity visible in living animals. Colour: gastrodermis white or pink,
depending on food.

Medusa umbrella somewhat flattened-hemispherical, with its oral surface more
or less six-sided and with well-developed thickened marginal ring packed with nema-
tocysts. Velum broad, usually closing umbrella and opened only when feeding.
Manubrium broadly conical, filling almost the entire subumbrella, extensible beyond
umbrella, mouth simple and without nematocyst knobs. Gonads in specialized cavity
or brood-chamber situated above the manubrium, often male and female gametes
present, female gonads in lower epithelium of brood-chamber, male gonads in upper
epithelium, and also on subumbrella, embryos develop in brood-chamber, up to seven
planulae can be present. Radial canals very short, usually six in number. Ring canal
relatively large, widening at junctions with radial canals. Tentacles hollow proximally,
chordoid distally, mostly six or five present, but rarely also other numbers possible,
bifurcated at about middle, upper branch only with one terminal nematocyst cluster,
lower branch terminating in adhesive organ used to cling to the substratum. One
abaxial ocellus on base of each tentacle. Medusae budding from bell margin and lower
part of exumbrella regularly present, also in reproductive animals. Colours: body olive-
cream, ocelli red to brown. Nematocysts: polyp with stenoteles of two size classes;
medusa with stenoteles and desmonemes.

DIMENSIONS: Polyp 1-6 mm and more, very contractile; tentacular capitula di-
ameter 0.25-0.30 mm. Medusa bell diameter 0.4-0.8 mm, tentacle length 1.8 mm. Egg
size 47-93 um (Tardent, 1978). Stenoteles of polyp: (30-33.5)x(19-21)um and (16.5-
18.5)x(9-11)um. Stenoteles of medusa: (12-24)x(7-16)um, apparently not falling into
distinct, different size classes. Desmonemes of medusa: (7.5-9.5)x(4-5)um.

DISTRIBUTION: Present along all coasts of Europe and the Mediterranean, rarer
in the southern North Sea, perhaps absent in the Baltic Sea. Also known from the Black
Sea (Thiel, 1935), Madeira (Friedrich, 1961), and the Caribbean (Spracklin, 1982). The
northernmost occurrence seems to be near Bergen in Norway (Kramp & Damas, 1925).
Records for the Swedish west coast are given by Hartlaub (1886), for the British Isles
by e. g. Hincks (1861, 1868), Allman (1872), for France by e. g. Drzewina & Bohn
(1913) and Teissier (1965), for the Atlantic coast of Spain by Medel & Löpez-Gonzälez
(1996). The Mediterranean records are summarized in Brinckmann-Voss (1970). Type
locality: Chausey Isles, English Channel.

BroLocy: The polyp has only rarely been found in nature. Hincks (1868)
observed that in England medusae budding takes place during summer and autumn.
The medusa lives attached to various seaweeds, preferably on Ulva species, but also on
Cystoseira, Gelidium and other substrata. It lives in shallow depths, also frequently in
tide pools. Brinckmann-Voss (1970) found the medusa also in deeper waters (20 m).
Sexual reproduction in the English Channel was observed in May to June and October
to November (Teissier, 1965).

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 383

ADDITIONAL DATA: Eleutheria dichotoma is easy to keep in small containers and
it has therefore been a favourite subject for a number of studies. Hartlaub (1886) ex-
amined the formation of the brood-chamber in histological sections. The subumbrellar
epidermis forms canal-shaped cavities that penetrate the region above the stomach. The
fusion of these canals in the centre then creates the brood-chamber. The brood-chamber
is connected to the subumbrella by interradial openings (thus usually six). The gastro-
dermis of the manubrium becomes completely separated from the umbrella. The eggs
mature in the lower epithelium of the chamber, while spermatids develop in the upper
epithelium. However, only 12% of Hartlaub’s animals were hermaphroditic, the others
were all female. Miiller (1908) supplemented the former observations with more de-
tails. He found that the brood-chamber develops very early in development, even while
the medusa is still attached. The fraction of hermaphroditic animals he found was
higher, namely 35% of the fertile animals. Miiller (1908) also reported that the sper-
matogonia can also be produced in small patches of the subumbrellar epidermis.
Krumbach (1907) made many observations on the living animal and he describes its
movements. He found that the velum tightly closes the umbrella. This he considered
might reduce desiccation in case the animal is exposed to the air. Drzewina & Bohn
(1913) studied the variability of the tentacle numbers in animals collected from nature.
They found the following distribution among 39 animals: 1x8 tentacles, 1x7 tentacles,
30x6 tentacles, 7x5 tentacles. They also made regeneration experiments, observed
anomalies, and investigated the influence of cyanide. The budding of medusae from the
medusa bell includes also part of the radial canal, thus the medusa buds are of ento-
and ectodermal origin (Hartlaub, 1886; Lengerich, 1923b). The development of the
medusa bud includes also the formation of an entocodon. The development of the
gametes takes place in the brood-chamber and the planulae are released by a rupture of
the umbrella. The latter heals again after the release (Lengerich, 1923a). The histology
of the medusa has been examined by several authors, e. g. Hartlaub (1886), Lengerich
(1923a), Bouillon (1968), van de Vyver & Bouillon (1969), and Bouillon (1994: fig.
55). Bouillon (1968) also examined the histochemistry and ultrastructure of the
adhesive organs. The haploid chromosome number is 6 (Makino, 1951; cited in
Tardent, 1978). Hauenschild (1956) observed that the ability to produce gametes can
irreversibly be lost in some clones, this either naturally or induced artificially. Hadrys
et al. (1990) examined the feeding behaviour of the medusa. Ender (1997) analyzed the
population structure using mitochondrial DNA sequences. Further experimental,
developmental and structural details are given by Weill (1936), Hauenschild (1957a,
b), Weiler-Stolt (1960), Schierwater (1989), Schierwater & Havenschild (1990),
Schierwater et al. (1991, 1992), Kuhn et al. (1996), Schierwater & Hadrys (1998).

REMARKS: The synonymy is given by Bedot (1912, 1916, 1918), Mayer (1910),
Lengerich (1923a), Kramp (1961), and Russell (1953). In its medusa phase, Eleutheria
dichotoma is very characteristic and among the European medusae it can only be
confounded with E. claparedii. The external medusae buds and the tentacles that are
bifurcating in the middle make E. dichotoma immediately recognizable. In addition, E.
dichotoma has usually five to six tentacles, while most E. claparedii have eight. Their
polyps appear very similar and are perhaps not distinguishable.

384 P. SCHUCHERT

Eleutheria claparedii Hartlaub, 1889 Figs 19D & 21

Eleutheria dichotoma — Claparède, 1863: 4, pl. 1 figs 4-10; Spagnolini, 1876: 312.

[not Eleutheria dichotoma Quatrefages, 1842].

Eleutheria heteronema Haeckel, 1879: 106.

Eleutheria diplonema Haeckel, 1879: 106.

Eleutheria claparedii Hartlaub, 1889: 665; Mayer, 1910: 95, fig. 49; Brinckmann-Voss, 1970:
80, figs 93-95.

Eleutheria claparedei — Hartlaub, 1907: 129, fig. 128; Müller, 1911: 159, pl. 3 fig. 1; Drzewina
& Bohn, 1913: 49, figs 26-37; Lengerich, 1922: 211, fig. 2; Lengerich, 1923a: 336, figs
V-Z, A1-F1; Browne & Kramp, 1939: 274; Kramp, 1961: 59; Bouillon et al., 2004: 88,
fig. 48E-F.

Staurocladia claparedei — Hartlaub, 1917: 401.

MATERIAL: Roscoff, France, September 2004, one medusa in plankton, mature male with
medusae buds; one immature medusa with medusae buds on Cystoseira spec., developed eggs
after 6 weeks of cultivation. 16S sequence of male medusa accession number AM088486.

DIAGNOSIS: Eleutheria medusa without brood-chamber, medusae buds within
subumbrella, usually eight tentacles, tentacles bifurcated once in distal region and
trunk thus clearly longer than branches, gonochoristic, nettle ring not conspicuous.

DESCRIPTION: Sedentary medusa clinging to seaweeds, flat, umbrella slightly
shallower than a hemisphere, without jelly, usually eight to nine tentacles (range 5-10),
not in phase with radial canals, tentacles relatively long when expanded, held hori-
zontal, bifurcated at 1/4 to 1/6 from end, the two branches of equal length, upper
branch ending in a capitulum, capitulum flat-button-shaped to oval, lower tentacle
branch ends in adhesive pad, not much enlarged, used to cling to the substrate. One
abaxial ocellus at base of each tentacle, colour red. Stomach large, filling nearly entire
subumbrella, conical; four to six short radial canals; ring canal thick, underlying nettle
ring inconspicuous but present. Velum conical, broad, usually closed and usually only
opened for feeding, releasing medusae, gametes or excrements. Gonochoristic, gonads
develop in stomach epidermis, without brood-chamber but embryos develop within
subumbrella. Asexual multiplication by medusae budding frequent, buds arise from
epidermis over radial canal in the subumbrellar cavity. Budding usually stops when
gonads develop. Colours: Manubrium and medusae buds orange, rest whitish-cream or
transparent. Nematocysts: stenoteles and desmonemes.

Polyp only known from cultivation experiments, colonies stolonal, swollen
head, one whorl of 4-5 capitate tentacles, no filiform tentacles, further development un-
known.

DIMENSIONS: Medusa bell diameter 0.4-0.5 mm, tentacle length up to 1 mm,
tentacle capitula diameter 90-110 um. Claparède (1863) reported an egg size of 0.18-
0.3 mm, spawned eggs observed in this study were 0.13 mm in diameter. Nematocysts
of medusa: stenoteles, (11-27)x(8-19)um, ratio of length to width = 1.5; desmonemes
(9-11.5)x(4.5-6)um, r=2.0. Polyp 0.7 mm high, tentacles 0.25 mm long (Drzewina &
Bohn, 1913).

DISTRIBUTION: French Channel coast, Mediterranean (Naples). Type locality:
Tahitou near St. Vaast la Hogue (Normandy, France).

BioLoGY: The medusa was usually found clinging on macroalgae like Ulva
spec. and Fucus spec. in the intertidal zone. In the English Channel, it is most frequent

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE

Fic. 19

A) Eleutheria dichotoma Quatrefages, 1842, cultivated polyp with medusa bud. B) as in A,
enlarged medusa bud on gonostyle. C) Eleutheria dichotoma, mature medusa, an egg can be faint-
ly seen. D) Eleutheria claparedii Hartlaub, 1889, medusa with two buds in the subumbrella.

386 P. SCHUCHERT

in autumn and winter and disappears almost completely in spring (Teissier, 1965).
Asexual production occurs all year round but especially during spring time, animals
with mature gonads were observed from May to June and September to November.

ADDITIONAL DATA: The number of tentacles seems not strictly fixed genetically,
as budded medusae may have other tentacle numbers than the mother medusa
(Drzwina & Bohn, 1913).

Drzewina & Bohn (1913) examined more than 100 medusae and found that
asexual medusae-budding stopped with commencing gonad maturation. One male
animal observed for this study, however, had medusae buds and mature gonads. The
female animal stopped budding before gonad development.

REMARKS: Eleutheria claparedii (original spelling) was named by Hartlaub
(1889) by referring to a description of Claparède (1863) of an animal the latter had
erroneously identified as E. dichotoma. Hartlaub (1889) states that he also found this
species at Naples. He did not designate type material nor a type locality. As Hartlaub
mainly referred to the description of Claparède (1863) based on material from Tahitou
near St. Vaast la Hogue (Normandy, France), and as the major other descriptions of this
species came from this population (Drzwina & Bohn, 1913; Miiller, 1911), this loca-
lity is herewith selected as type locality. Hartlaub (1907) introduced the incorrect
spelling E. claparedei, adopted by most subsequent authors. According to the ICZN
[1999, 4th ed.; articles 31.1.1 and 31.1.3], the epithet can be formed in both versions,
but as the spelling in the original publication was Eleutheria claparedii, this spelling
must be used.

Before Hartlaub (1889), Haeckel (1879) had already introduced two new names
for Claparède's animals: Eleutheria heteronema and Eleutheria diplonema. Although
Haeckel's names are senior synonyms, they have never been used as valid names.
Article 23.9.1.1 of the ICZN [1999, 4th ed.] is therefore invoked to declare them as
invalid and Eleutheria claparedii Hartlaub, 1889 the valid name of this species.

Eleutheria claparedii was investigated in detail by Drzewina & Bohn (1913)
and Lengerich (1923a).

The polyp has not yet been identified in the sea and only the young polyp with-
out medusae buds is known from cultivation experiments. Drzewina & Bohn (1913)
found them indistinguishable from young polyps of E. dichotoma.

Genus Staurocladia Hartlaub, 1917

TYPE SPECIES: Eleutheria vallentini Browne, 1902, by designation of Hartlaub (1917).

SYNONYMS: Wandelia Bedot, 1908; Cnidonema Gilchrist, 1919.

REFERENCES: Hartlaub (1917); Gilchrist (1919); Browne & Kramp (1939); Kramp
(1961); Bouillon (1985); Petersen (1990); Bouillon & Boero (2000).

DIaGNOSIS: Hydroid stolonal or sessile, hydranths with an oral whorl of capitate
tentacles, with or without aboral filiform tentacles; medusa buds borne singly on hy-
dranth body immediately above filiform tentacles or in similar position when these are
absent. Medusa adapted to a crawling and walking mode of life, exceptionally still able
to swim; without brood-chamber above manubrium; with or without continuous or
interrupted nematocyst ring along margin; gonads around manubrium or developed on

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 387

We

Fic. 20

Eleutheria dichotoma Quatrefages, 1842. A) Medusa with developing embryos in brood-
chamber, after living material, scale bar 0.5 mm. B) Polyp stage obtained from medusa shown
in figure A, scale bar 0.5 mm. C) Polyp with medusae buds, modified after Hauenschild (1956).
D) Schematic organization of the medusa in a vertical section, right half perradial, left half
interradial; modified after Lengerich (1922). Abbreviations: bc brood-chamber with developing
embryos, ct upper branch of tentacle with nematocyst knob (capitulum), e developing eggs, gd
gastrodermis, m mouth, nr nettle-ring, oc ocellus, rc radial canal, s stomach, sf lower branch of
tentacle with adhesive organ, sp male gonad, v velum.

388 P. SCHUCHERT

Fic. 21

Eleutheria claparedii Hartlaub, 1889. A) after living animal from Roscoff, note the two medusa
buds within the subumbrella (stippled darker), scale bar 0.2 mm. B) Young hydranth, after
Drzewina & Bohn (1913).

manubrial protrusions; with six to eleven radial canals, some bifurcating shortly distal
to manubrium; mouth circular with or without nematocyst knobs; with up to 60 mar-
ginal tentacles, dichotomous, upper branch with several nematocyst clusters, lower
with adhesive organ; often asexual reproduction by medusa budding or by fission; with
abaxial ocelli at tentacle base.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 389

REMARKS: The problems concerning the distinction of Eleutheria and
Staurocladia have been discussed above under "Remarks" at the family level. There is
only one species in the European fauna.

Staurocladia portmanni Brinckmann, 1964 Fig. 22
Staurocladia portmanni Brinckmann, 1964: 693, figs 1-10; Brinckmann-Voss, 1970: 82, figs 96-

98, pl. 6 figs 1-3; Bouillon et al., 2004: 90, fig. 48I-J.

MATERIAL EXAMINED: Staurocladia portmanni, syntypes BMNH 1963.12.10.2, Ischia,
Naples, 2 mature medusae. — Staurocladia portmanni syntype BMNH 1963.12.10.1, Italy, Gulf
of Sorento, Vico Equense, polyp colony.

DiaGNOSIS: Hydroid with one oral whorl of capitate tentacles and an aboral
whorl of six filiform tentacles, short perisarc covered pedicel, medusae budded directly
from hydranth body above filiform tentacles. Medusa clinging and creeping on sub-
stratum, but also able to swim freely, five to ten branched radial canals, each sending
a protuberance into mesogloea, 18-24 tentacles, bifid, nematocyst clusters terminal and
on oral and aboral sides of upper branch, plus an additional lateral pair near branching
point; five interradial manubrial pouches; gonad surrounds entire manubrium; mouth
with five nematocyst clusters.

DESCRIPTION (after Brinckmann, 1964; Brinckmann-Voss, 1970; Bouillon,
1966; and own observations): Hydroid forming small, stolonal colonies. Stolons
ensheathed by perisarc which extends up to the short pedicel; hydranth with three to
five capitate tentacles in an oral whorl and six filiform tentacles at base of hydranth
body, number of filiform tentacles constant; usually one, rarely two medusae buds
above filiform tentacles. Preoral cavity present. Nematocysts: stenoteles. Colour: pink-
orange. Medusa-bud development very slow, taking up to 50 days.

Newly liberated medusa 0.6-0.7 mm in diameter; seven to nine tentacles, bifid,
oral branch with terminal adhesive end, upper branch short and with one terminal
nematocyst cluster, sometimes also an additional aboral one, two lateral nematocyst
clusters on common trunk near bifurcation always present; five to ten radial canals,
number not increasing with further development, protuberances small.

Adult medusa with bell-shaped umbrella, wider than high. Manubrium not
extending beyond velum, with five protruding stomach-pouches in middle, mouth with
five spherical nematocyst clusters. Gonads surround manubrium and stomach pouches
without perradial interruptions, gonads get separated into five sections only where
radial canals are adnate to top of the manubrium; gonochoristic, no medusa budding.
Five radial canals originating near centre of top of manubrium, then sandwiched
between subumbrella and broad base of manubrium for a third of their length, some
branching dichotomously so that up to ten radial canals reach the circular canal, at
upper third of each radial canal an upright, finger-like protuberance that penetrates into
mesogloea; radial canals widen bulb-like before joining circular canal. Tentacles 16-25
in number, hollow, lumen communicates with circular canal; each tentacle bifurcated
at about the middle; lower branch with terminal adhesive pad; upper branch with
terminal nematocyst cluster (capitulum), plus two to five aboral, and two to four aboral
nematocyst clusters; common trunk bears additionally two lateral nematocyst clusters
near branching point. Each tentacle contains one basal, abaxial ocellus. Border

390 P. SCHUCHERT

FIG. 22

Staurocladia portmanni Brinckmann, 1964, modified after Brinckmann-Voss (1964), above: ma-
ture medusa, below: polyp with medusa bud. For size see text.

between exumbrella and velum is thickened with nematocysts. Colours: gastrodermis
of stomach, circular canal, and tentacles red-orange; upper part of radial canals and
protuberances contain a white pigment, same pigment is also found in the distal part of
radial canals where they widen and join circular canal. Nematocysts: stenoteles,
desmonemes, and a large, unidentified capsule.

DIMENSIONS: Hydranth body 1.5-2 mm in height, Adult medusa 4-6 mm in
diameter, bell-height 2.5-5 mm.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 391

DISTRIBUTION: Only known from the region of Naples (Tyrrhenian Sea,
Mediterranean). The material used for the original description came from the Gulf of
Sorrento (polyp stage) and the island of Ischia (medusa). Under type locality,
Brinckmann (1964) gives only the registration number BMNH 1963.12.10.1. Accor-
ding to the label of this specimen in the BMNH, it came from Vico Equense in the Gulf
of Sorrento and this locality must therefore be assumed to be the type locality.

BioLocy: The medusae of S. portmanni are found in Posidonia oceanica beds
in depths of 15-35 m from end of May to begin of July. The hydroid was found on algae
like Udothea sp. and Halimeda tuna in a depth of 30 m. The medusa development is
extraordinarily slow, it takes up to 50 days from the first bud to the liberation and
another 40-60 to sexual maturity.

The medusa of S. portmanni is able to creep as well as to swim freely. More
details on its behaviour are given in Brinckmann-Voss (1970).

Genus Dendronema Haeckel, 1879

TYPE SPECIES: Dendronema stylodendron Haeckel, 1879 by monotypy.
REFERENCES: Haeckel (1879); Kramp (1961); Bouillon & Boero (2000).

DIAGNOsIs: Cladonematidae with branched oral tentacles; umbrella bell-shaped,
with an apical cavity above manubrium.

REMARKS: A problematic genus containing a single, doubtful species, see dis-
cussion below.

Dendronema stylodendron Haeckel, 1879 Fig. 23
Dendronema stylodendron Haeckel, 1879: 110, pl. 7 fig. 8; Lengerich, 1922: 210; Kramp, 1955:
307; Kramp, 1959: 96; Kramp, 1961: 58.
MATERIAL: None examined, likely no type material exists.

DIAGNOSIS: As for genus.

DEscRIPTION (after Haeckel, 1879): Umbrella bell shaped, height 9 mm,
diameter 6 mm; with pointed, conical apical process, nearly twice as high as broad.
Manubrium spindle-shaped, continued into mesogloea as apical cavity, in proximal
half with four egg-shaped gonads, mouth with four dichotomously branched oral
tentacles (50-60 ends) reaching down to velum level. Four radial canals that divide
close to manubrium into two branches. Eight tentacles, each with an ocellus that has a
lens, each tentacle divided into two principal branches, of which the adaxial one
branches two times and bears four adhesive ends, the abaxial branch divides dichoto-
mously several times and bears many nematocyst knobs. Colours: manubrium and
apical cavity reddish-yellow; tentacles, radial- and circular canals red; ocelli black.

DISTRIBUTION: Known from original description only, type locality: Lanzarote,
Canary Islands.

REMARKS: Besides its first description, Dendronema stylodendron has never
been reported again and some authors doubted its existence, e. g. Lengerich (1923a)
lists it as a synonym of Cladonema radiatum. Kramp (1955, 1961) thought that this is

392 P. SCHUCHERT

Fic. 23
Dendronema stylodendron, after Haeckel 1879.

unlikely, but nevertheless classified it as doubtful (Kramp, 1959). Haeckel based his
description of on a living specimen he collected at Lanzarote. The animal is remark-
ably large for this family and his drawings are quite detailed. So, it seems quite
unlikely that Haeckel confounded it with the much smaller Cladonema radiatum.
However, the species remains somewhat doubtful until it can be found again.

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. I also wish to express my sincere thanks to the museums of Copenhagen,
Bergen, Bruxelles, and Hamburg which provided valuable loans of material. Many
thanks are also due to Dr B. Schierwater who provided a culture of E. dichotoma, Dr
M. Stierwald who provided photos of Cladonema radiatum, and Dr W. Vervoort who
gave me valuable taxonomic advice on the validity of C. cocksii. Mrs J. Pralong kind-
ly made the serial histological sections. I would also like to thank Dr F. Boero, Dr D.
Calder, and Dr A. Marques who commented on initial versions of the manuscript.
However, this is not meant to delegate my responsability for the content and all short-
comings and errors are mine alone.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 393

REFERENCES

ACHERMANN, J. & TARDENT, P. 1973. Untersuchungen zum Problem des Nematocyten-
Nachschubs bei Cladonema radiatum (Hydrozoa). Revue suisse de Zoologie 80: 704-
712%

Acassız, L. 1862. Contributions to the Natural History of the United States of America. Vol. IV.
Little Brown, Boston, pp. 1-380, pls 1-19.

ALLMAN, G. J. 1871-1872. A monograph of the gymnoblastic or tubularian hydroids. Conclusion
of Part I, and Part II, containing descriptions of the genera and species of Gymnoblastea.
Ray Society, London, pp. 155-450, 23 plates.

ALLMAN, G. J. 1874. Notes on the structure and development of Myriothela phrygia. Annals and
Magazine of Natural History (4)14: 317-321.

ALLMAN, G. J. 1876. On the structure and development of Myriothela. Philosophical
Transactions of the Royal Society of London (B)165: 549-575, pls 55-58.

ALLMAN, G. J. 1888. Report on the Hydroida dredged by H. M. S. Challenger during the years
1873-76. Part IL.- The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and
Thalamophora. The Voyage of H. M. S. Challenger, Zoology 23: 1-90.

ANTSULEVICH, A. E. 1988. The first finding of hydroids of the genus Monocoryne (Hydrozoa,
Myriothelidae) in the waters of the USSR. Zoologicheskii Zhurnal 67: 931-933.
ANTSULEVICH, A. E. & POLTEVA, D. G. 1986. Hydroids of the genus Rhizogeton (Athecata,

Clavidae) in fauna of the USSR. Zoologicheskii Zhurnal 65: 965-972.

AURIVILLIUS, C. W. S. 1898. Vergleichende Thiergeographische Untersuchungen über die
Plankton-Fauna des Skageraks in den Jahren 1893-97. Kongliga Svenska Vetenskaps-
Akademiens Handlingar 30: 1-427.

AYRES, W. O. 1854. A description of a new species of polyp from Long Island, allied to
Tubularia, under the name Globiceps tiarella Ayres. Proceedings of the Boston Society
of Natural History 4: 193-196.

BAKER, E. G. S. 1936. Photoperiodicity in the spawning reaction of Pennaria tiarella McCr.
Proceedings of the Indian Academy of Science 45: 251-252.

BAKER, R. J. 1913. Protohydra Leuckartii in tidal pools of River Tavy. Transactions of the
Plymouth District Field Council 1: 23.

BARNES, R. S. K. 1994. The brackish-water fauna of northwestern Europe: an identification
guide to brackish-water habitats, ecology and macrofauna for field workers, naturalists
and students. Cambridge University Press, Cambridge, pp. XVI + 287.

BEDOT, M. 1901. Matériaux pour servir à l'histoire des hydroides. 1re période. Revue suisse de
Zoologie 9: 379-515.

BEDOT, M. 1908. Sur un animal pélagique de la région antarctique. Expédition antarctique
Française (1903-1905), Spongaires et Coelentérés 3: 1-7, plate.

BEDOT, M. 1911. Notes sur les hydroïdes de Roscoff. Archives de Zoologie Expérimentale et
Générale 6: 201-228.

BEDOT, M. 1912. Matériaux pour servir à l’histoire des hydroïdes. 4e période (1872-1880).
Revue suisse de Zoologie 20: 213-469.

BEDOT, M. 1916. Matériaux pour servir à l’histoire des hydroïdes. 5e période (1881-1890).
Revue suisse de Zoologie 24: 1-349.

BEDOT, M. 1918. Matériaux pour servir à l’histoire des hydroïdes. 6e période (1891-1900).
Revue suisse de Zoologie 26, supplement: 1-376.

BEIGEL, U. 1976. Gonophorenentwicklung der Hydroide Myriothela. Cahiers de Biologie
Marine 17: 119-129, pls 1-4.

BEIGEL-HEUWINKEL, U. 1984. Zur Eientwicklung bei Myriothela cocksi (Hydrozoa, Athecata).
Verhandlungen der Deutschen Zoologischen Gesellschaft 77: 273.

BEIGEL-HEUWINKEL, U. 1982a. Helical fibrils in the mesoglea of a hydropolyp. Tissue & Cell 14:
225-230.

394 P. SCHUCHERT

BEIGEL-HEUWINKEL, U. 1982b. Experiments and observations on regeneration in Myriothela
cocksi. Acta Zoologica 63: 199-210.

BEIGEL-HEUWINKEL, U. 1988. Brutpflege bei den marinen Hydroidpolypen Myriothela cocksi
(Vigurs), mit Hilfe spezieller Tentakel. Acta biologica Benrodis 1: 56-67.

BENOÎT, P. 1923a. L’ovogenèse et la segmentation de Myriothela Cocksi (Vigurs). Comptes ren-
dus hebdomadaires des Séances de l’Académie des Sciences, Paris 176: 1836-1838.

BENOÎT, P. 1923b. Le gonophore hermaphrodite de Myriothela cocksi (Vigurs). Comptes rendus
hebdomadaires des séances et mémoires de la Société de biologie et de ses filiales et as-
sociées 89: 507-510.

BENOÎT, P. 1925. L’ovogénèse et les premiers stades du développement chez la Myriothèle et
chez la Tubularie. Archives de Zoologie Expérimentale et Générale 64: 85-326.
BERRILL, N. J. 1952. Growth and form in gymnoblastic hydroids. II. Sexual and seasonal repro-
duction in Rathkea. III. Hydranth and gonophore development in Pennaria and Acaulis.

IV. Relative growth in Eudendrium. Journal of Morphology 90: 1-32.

BILLARD, A. 1905. Note complémentaire sur les hydroides de la Baie de la Hougue. Bulletin du
Muséum national d’histoire naturelle, Paris 11: 500-502.

BILLARD, A. 1921. Note sur la biologie et la régénération de la Myriothèle. Bulletin de la Société
Zologique de la France 46: 12-17.

BOADEN, P. J. S. 1976. Soft meiofauna of sand from the delta region of the Rhine, Meuse and
Scheldt. Netherlands Journal of Sea Research 10: 461-471.

Bobo, F. 1970. Etude du développement embryonnaire de l’anthoméduse Cladonema radiatum
Dujardin (Athécate, Capitée, Cladonematidée). Annales d’embryologie et de mor-
phogenèse 3: 309-313.

Boero, F. & FRESI, E. 1986. Zonation and evolution of a rocky bottom hydroid community.
Marine Ecology 7: 123-150.

BOERO, F. 1981. Systematics and ecology of the hydroid population of two Posidonia oceanica
Meadows. Marine Ecology 2: 181-197.

BONNEVIE, K. 1898a. Zur Systematik der Hydroiden. Zeitschrift fiir Wissenschaftliche Zoologie
63: 465-495, plates 25-27.

BONNEVIE, C. 1898b. Neue norwegische Hydroiden. Bergens Museum Àrbok 5: 1-16, plates 1-2.

BONNEVIE, K. 1899. Hydroida. Norske Nordhavs-Expedition 1876-1878, Zoologi 26: 1-104, pls
1-8, map.

Bosc, L. A. G. 1797. Description des objets nouveaux d’histoire naturelle trouvés dans une tra-
versée de Bordeaux à Charles-Town. Bulletin des Sciences, par la Société philomatique
de Paris 1(2): 9-10.

BOUILLON, J. 1965. In: Tessier (1965).

BOUILLON, J. 1966. Les cellules glandulaires des Hydroides et Hydroméduses. Leur structure et
la nature de leurs sécrétions. Cahiers de Biologie Marine 7: 157-205.

BOUILLON, J. 1968. Sur la structure des tentacules adhésifs des Cladonematidae et Eleutheriidae
(Anthomedusae). Pubblicazioni della Stazione Zoologica di Napoli 36: 471-504.
BOUILLON, J. 1966. Les cellules glandulaires des Hydroides et Hydroméduses. Leur structure et

la nature de leurs sécrétions. Cahiers de Biologie Marine 7: 157-205.

BOUILLON, J. 1971. Sur quelques hydroides de Roscoff. Cahiers de Biologie Marine 12: 323-
364.

BOUILLON, J. 1974. Description de Teissiera milleporoides, nouveau genre et nouvelle espèce de
Zancleidae des Seychelles (Hydrozoaires; Athecates-Anthoméduses), avec une révision
des Hydroides "Pteronematoidea". Cahiers de Biologie Marine 15: 113-154.

BOUILLON, J. 1985. Essai de classification des hydropolypes-hydroméduses (Hydrozoa-
Cnidaria). Indo-Malayan Zoology 2: 29-243.

BOUILLON, J. 1994. Les Hydrozoaires (pp. 29-416). In: P. Grassé & J. DOUMENG (eds). Traité de
Zoologie. Masson, Paris.

BOUILLON, J., BOERO, F. & SEGHERS, G. 1987. Redescription of Cladocoryne haddoni
Kirkpatrick and a proposed phylogeny of the superfamily Zancleoidea (Anthomedusae,
Hydrozoa, Cnidaria). Indo-Malayan Zoology 4: 279-292.

BOUILLON, J., BoERO, F. & SEGHERS, G. 1988. Notes additionnelles sur les Hydroméduses de la
mer de Bismarck (Hydrozoa-Cnidaria) 2. Indo-Malayan Zoology 5: 87-99.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 395

BOUILLON, J., MASSIN, C. & KRESEVIC, R. 1995. Hydroidomedusae of the Belgian Royal Society
of Natural Sciences. Institut Royal Des Sciences Naturelles de Belgique Documents de
Travail 78: 1-106.

BOUILLON, J. & HOUVENAGHEL, G. 1970. Histophysiologie et la digestion chez Cladonema
radiatum, Dujardin, 1843. Pubblicazioni della Stazione Zoologica di Napoli 38: 71-108.

BOUILLON, J. & BOERO, F. 2000. Synopsis of the families and genera of the Hydromedusae of the
world, with a list of the worldwide species. Thalassia Salentina 24: 47-296.

BOUILLON, J., MEDEL, M. D., PAGES, F., GILI, J. M., BOERO, B. & GRAVILI, C. 2004. Fauna of the
Mediterranean Hydrozoa. Scientia Marina 68(Suppl. 2): 1-448.

BOZHENOVA, O. V., STEPANJANTS, S. D. & SHEREMETEVSKY, A. M. 1989. The first finding of the
meiobenthic Cnidaria Boreohydra simplex (Hydrozoa, Athecata) in the White Sea.
Zoologicheskii Zhurnal 68: 11-16.

BRIEN, P. 1941. Remarques au suject des conceptions relatives à l’existence et à la pérennité,
chez les Hydroides, d’une réserve embryonnaire et d’une lignée germinale, à propos du
bourgeonnement et des potentialités de l’ectoderme de "Cladonema radiatum" (Duj.).
Annales de la Société royale zoologique de Belgique 72: 37-62.

BRIEN, P. 1942. Étude sur deux hydroides gymnoblastiques Cladonema radiatum (Duj.) et Clava
squamata (O.F. Miiller) (Origine des cellules blastogénétiques, sexuelles, des cnido-
blastes et des cellules glandulaires. Le polype, la méduse, le gonophore). Mémoires de
l’Académie royale de Belgique, Classe des sciences 20: 1-116.

BRINCKMANN, A. 1964. Observations on the biology and development of Staurocladia portmanni
sp. n. (Anthomedusae, Eleutheridae). Canadian Journal of Zoology 42: 693-706, pls
1-2.

BRINCKMANN, A. 1966. The morphology and development of Acaulis ilonae sp. nov. (order
Anthomedusae-Athecatae, Fam. Acaulidae). Canadian Journal of Zoology 44: 291-301.

BRINCKMANN-Voss, A. 1970. Anthomedusae/Athecata (Hydrozoa, Cnidaria) of the Mediter-
ranean. Part I. Capitata. Fauna e Flora Golfo di Napoli 39: 1-96, pls 1-11.

BRINCKMANN, A. & PETERSEN, K. W. 1960. On some distinguishing characters of Dipurena reesi
Vannucci 1956 and Cladonema radiatum Dujardin 1843. Pubblicazioni della Stazione
Zoologica di Napoli 31: 386-392.

BROCH, H. 1903. Die vom norwegischen Fischereidampfer "Michael Sars" in den Jahren 1900-
1902 in dem Nordmeer gesammelten Hydroiden. Bergens Museums Aarbog (1903) 9:
1-14, pls 1-4, table.

BROCH, H. 1910. Die Hydroiden der Arktischen Meere. Fauna Arctica 5: 127-248, plates 2-4.

BROCH, H. 1916. Hydroida. (Part I). Danish Ingolf Expedition 5: 1-66.

BROWNE, E. T. 1900. Report on the Medusae. The fauna and flora of Valencia Harbour on the
west coast of Ireland. Proceedings of the Royal Irish Academy (3)5: 694-736, pls 20-21.

BROWNE, E. T. 1902. A preliminary report on Hydromedusae from the Falkland islands. Annals
and Magazine of natural History (7) 9: 272-284.

BROWNE, E. T. 1903. Report on some medusae from Norway and Spitzbergen. Bergens Museum
Aarbog 4: 1-36.

BROWNE, E. T., & KRAMP, P. L. 1939. Hydromedusae from the Falkland Islands. Discovery
Reports 18: 265-322.

BRUMWELL, G. B. & MARTIN, V. J. 1996. Ultrastructural localization of RFamide-like peptides
in neuronal dense-cored vesicles of a cnidarian planula larva. Invertebrate Biology 115:
13-19.

CALDER, D. R. 1974. The mud-dwelling hydrozoan Boreohydra simplex in the western North
Atlantic. Journal of the Fisheries Research Board of Canada 31: 1666-1667, pl. 1.

CALDER, D. R. 1988. Shallow-water hydroids of Bermuda. The Athecatae. Royal Ontario
Museum Life Sciences Contributions 148: 1-107.

CALDER, D. R., MALLINSON, J. J., COLLINS, K. & HICKMAN, C. P. 2003. Additions to the hydroids
(Cnidaria) of the Galapagos, with a list of species reported from the islands. Journal of
Natural History 37: 1173-1218.

396 P. SCHUCHERT

CASTRIC-FEY, A. 1970. Sur quelques hydraires de 1° Archipel de Glénan (Sud-Finistère). Vie et
Milieu 21: 1-23.

CERFONTAINE, P. 1902. Recherches expérimentales sur la régénération et l’hétéromorphose chez
Astroides calycularis et Pennaria cavolinii. Archives de Biologie, Paris 19: 245-315.

CHRISTIANSEN, B. O. 1972. The hydroid fauna of the Oslo Fjord in Norway. Norwegian Journal
of Zoology 20: 279-310.

CLAPARÈDE, R. E. 1863. Beobachtungen über Anatomie und Entwicklungsgeschichte wirbelloser
Thiere an der Kiiste von Normandie angestellt. Wilhelm Engelmann, Leipzig, pp. 120, 18
plates.

CLARK, S. D. & Cook, C. B. 1986. Inhibition of nematocyst discharge during feeding in the colo-
nial hydroid Halocordyle disticha (=Pennaria tiarella): the role of previous prey-killing.
Biological Bulletin 171: 405-416.

CLAUSEN, C. 1971. Interstitial Cnidaria: present status of their systematics and ecology.
Smithsonian Contributions to Zoology 76: 1-8.

CLAUSEN, C. & VON SALVINI-PLAWEN, L. 1986. Cnidaria (pp. 33-42). In: BOTOSANEANU, L. (ed).
Stygofauna Mundi, A faunistic, distributional, and ecological synthesis of the worldfau-
na inhabiting subterranean waters (including the marine interstitial). Brill/Dr. W.
Backhuys, Leiden, pp. i-vi, 1-740.

Cocks, W. P. 1850. Contributions to the fauna of Falmouth. Annual Report of the Royal
Cornwall Polytechnic Society 17: 38-103. [publication year 1850 according to Cornelius
(1977)]

Cocks, W. P. 1853. Spadix purpurea, Gosse. Annals and Magazine of Natural History (2) 12:
365.

Cocks, W. P. 1854. Contributions to the Falmouth Fauna. Annual Report of the Royal Cornwall
Polytechnic Society 20: 1-68. [publication year 1854 according to Cornelius (1977)]

COLLINS, A.G., WINKELMANN, S. & SCHIERWATER, B. 2005. An assessment of partial mitochon-
drial 16S rDNA sequences as indicators of Corynidae (Hydrozoa, Anthoathecata) phy-
logeny. Zoologica Scripta 34, 91-99.

COLLINS, A. G., SCHUCHERT, P., MARQUES, A. C., JANKOWSKI, T., MEDINA, M. & SCHIERWATER,
B. 2005. Cnidarian Phylogeny and Character Evolution Clarified by New Large and
Small Subunit rDNA Data and an Assessment of the Utility of Phylogenetic Mixture
Models. Systematic Biology, in press.

COOKE, W. J. 1977. Order Hydroida. Bernice P. Bishop Museum Special Publication 64: 71-104.

CORNELIUS, P. F. S. 1977. On the nomenclature of the hydroid, Candelabrum phrygium
(Fabricius, 1780) (= Myriothela phrygia, Arum cocksi). Journal of the Marine Biological
Association of the U. K. 57: 521-524.

CORNELIUS, P. F. S. 1992. The Azores hydroid fauna and its origin, with discussion of rafting and
medusa supression. Arquipelago Ciencias da Natureza 10: 75-99.

COSTELLO, M. J., EMBLOW, C. & WHITE R. (eds) 2001. European Register of Marine Species. A
check-list of marine species in Europe and a bibliography of guides to their
identification. Patrimoines naturels 50, 1-463. Available online at:
http://www.MarBEF.org/data/erms.php.

COWDEN, R. R. 1964. A cytochemical study of gonophore and oocyte development in Pennaria
tiarella. Acta embryologiae et morphologiae experimentalis 7: 167-179.

COWDEN, R. R. 1965a. A cytological and cytochemical study of hydranths of the hydroid coel-
enterate, Pennaria tiarella. Zeitschrift für Zellforschung und mikroskopische Anatomie
65: 869-883.

COWDEN, R. R. 1965b. Cytochemical studies of embryonic development to metamorphosis in the
gymnoblastic hydroid, Pennaria tiarella. Acta embryologiae et morphologiae experi-
mentalis 8: 221-231.

DA CUNHA, A. X. 1944. Hidropélipos das costas de Portugal. Memorias e Estudos de Museu
Zoologico da Universidade de Coimbra 161: 1-101.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 397

DA SILVEIRA, F. L. & MIGOTTO, A., E, 1991. The variation of Halocordyle disticha (Cnidaria,
Athecata) from the Brazilian coast: an environmental indicator species? Hydrobiologia
216/217: 437-422.

DALY YAHIA, M. N., Goy, J. & DALY YAHIA-KEFI, O. 2003. Distribution et écologie des Méduses
(Cnidaria) du golfe de Tunis (Mediterranée sud occidentale). Distribution and ecology of
Medusae and Scyphomedusae (Cnidaria) in Tunis Gulf (SW Mediterranean).
Oceanologica Acta 26: 645-655.

DAWYDOFF, C. N. 1930. Protohydra caulleryi, nov. sp., des eaux indochinoises. Archives de
Zoologie Expérimentale et Générale, notes et revue 70: 55-57.

BLAINVILLE, H. M. DE 1830. Zoophytes (pp. 326). In: CUVIER, F. (ed.). Dictionnaire des sciences
naturelles. Volume 60. Levrault and Le Normant, Strasbourg & Paris. 72 volumes.

DE FILIPPI, F. 1866. Sopra due Idrozoi del Mediterraneo. Memorie della Reale Accademia delle
Scienze di Torino, Scienze fisiche e matematiche Serie 2 vol. 23: 375-385.

DENDY, A. C. 1902. On a free-swimming hydroid, Pelagohydra mirabilis n. gen. et sp. The quar-
terly journal of microscopic science 46: 1-24, plates 1-2.

DRZEWINA, A. & BOHN, G. 1913. Observations biologiques sur Eleutheria dichotoma et E. cla-
paredei. Archives de Zoologie Expérimentale et Générale 53: 15-59.

Du PLEssis, G. 1880. Observations sur la Cladocoryne floconneuse (Cladocoryne floccosa,
Rotch.). Bulletin de la Société vaudoise des Sciences naturelles (2)17: 119-144, pl. 10.

Du PLESSIS, G. 1909. Note sur l’élevage des Eleutheries de la Méditerranée au moyen de l’iso-
lement. Revue suisse de Zoologie 17: 371-377.

DUJARDIN, F. 1843. Observations sur un nouveau genre de médusaires provenant de la méta-
morphose des Syncorynes. Comptes rendus hebdomadaires des Séances de l’Académie
des Sciences, Paris 16: 1132-1136.

DUJARDIN, F. 1845. Mémoire sur le dévelopment des Méduses et des Polypes Hydraires. Annales
des Sciences Naturelles, Paris (3), Zoologie 4: 257-280.

EHRENBERG, C. G. 1834. Beiträge zur physiologischen Kenntnis der Corallenthiere im allgemei-
nen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen
Systematik derselben. Abhandlungen der Königlichen Akademie der Wissenschaften,
Berlin 1: 225-380.

ENDER, A. 1997 Untersuchungen zur Evolutionsgenetik des athekaten Hydrozoons Eleutheria
dichotoma (Quatrefages 1842). PhD Thesis Fachbereich Biologie, Johan Wolfgang
Goethe Universität Frankfurt, 103 pp.

ESCHSCHOLTZ, F. 1829. System der Acalephen. Eine ausführliche Beschreibung aller medusenar-
tigen Strahltiere. Ferdinand Dümmler, Berlin, 190 pp.

EVERTSEN, J., BAKKEN, T. & GREEN, S. 2004. Rediscovery of Tenellia adspersa (Nudibranchia)
from the Finnish archipelago. Sarsia 89: 362-365.

FABRICIUS, O. 1780. Fauna Groenlandica. G. Rothe, Hafniae and Lipsiae, pp. 1-452, plate 1.

FALUGI, C., MORRI, C., BOUILLON, J., & BOERO, F. 1994. Localization of some neurotransmitters
during development in hydroidomedusae. Tissue & Cell 26: 523-538.

FEWKES, J. W. 1890. A zoölogical reconnoissance in Grand Manan. American Naturalist 24:
432-438.

FORBES, E. 1848. A monograph of the British naked-eyed medusae: with figures of all the
species. Ray Society, London, pp. 104, 13 plates.

FRASER, C. M. 1924. Acaulis primarius Stimpson. Transactions of the Royal Society of Canada,
Sec. V. 173: 165-173.

FRASER, C. M. 1938. Hydroids of the 1932, 1933, 1935 and 1938 Allan Hancock Pacific
Expeditions. Allan Hancock Pacific Expeditions 4: 129-153.

FRASER, C. M. 1941. New species of hydroids, mostly from the Atlantic Ocean, in the United
States National Museum. Proceedings of the United States National Museum 91: 77-89.

FRASER, C. M. 1944. Hydroids of the Atlantic coast of North America. The University of Toronto
Press, Toronto, pp. 1-451, pls 1-94.

398 P. SCHUCHERT

FRASER, C. M. 1948. Hydroids of the Allan Hancock Pacific Expeditions since March, 1938.
Allan Hancock Pacific Expeditions 4: 179-343.

FRIEDRICH, H. 1961. Die Hydromeduse Eleutheria dichotoma Quatrefages im Litoral der
Atlantischen Insel Madeira. Bocagiana 8: 1.

GARCIA CORRALES, P. & AGUIRRE, A. 1985. La especie Halocordyle disticha (Goldfuss, 1820), y
sus sinonimias. Boletin del Instituto Espafiol de Oceanografia 2: 85-96.

GAST, R. & GODLEWSKI, E. 1903. Die Regulationserscheinungen bei Pennaria cavolinii. Archiv
für Entwicklungsmechanik der Organismen 16: 76-116.

GEGENBAUR, C. 1856. Versuch eines Systems der Medusen, mit Beschreibung neuer oder wenig
gekannter Formen; zugleich ein Beitrag zur Kenntnis der Fauna des Mittelmeeres.
Zeitschrift für wissenschaftliche Zoologie, Leipzig 8: 202-273, pls 7-9.

GENZANO, G. N. & KUBOTA, S. 2003. Synchronous mass release of mature medusae from the
hydroid Halocordyle disticha (Hydrozoa, Halocordylidae) and experimental induction of
different timing by light changes. Publications of the Seto Marine Biological Laboratory
39: 221-228.

GIBBONS, M. J. & RYLAND, J. S. 1989. Intertidal and shallow water hydroids from Fiji. 1.
Athecata to Sertulariidae. Memoirs of the Queensland Museum 27: 377-432.

GILCHRIST, J. D. F 1919. On a species of the crawling medusa, Eleutheria, from the Cape of
Good Hope (Cnidonema capensis g. et sp. n.) and the southern Eleutheridae. Quarterly
Journal of microscopic Science 63: 509-529.

GOLDFUSS, G. A. 1820. Handbuch der Zoologie. Erste Abteilung. Schrag, Nürnberg, pp. 1-696.

Gosse, P. H. 1853a. Notes on some new or little-known marine animals. Annals and Magazine
of Natural History (2)12: 124-128.

GosseE, P. H. 1853b. Notes on some new or little-known marine animals. Annals and Magazine
of Natural History (2)12: 153-159.

Gosse, P. H. 1853c. Notes on some new or little-known marine animals. Annals and Magazine
of Natural History (2)12: 384-386.

GossE, P. H. 1853d. A naturalist’s rambles on the Devonshire coast. John van Voorst, London,
pp. xvi+451, 28 plates.

GRAEFFE, E. 1883a. Ueber Polycoryne Helleri eine neue Corynidengattung. Bolletino della
Societa Adriatica di Scienze Naturali in Trieste 8: 202-205.

GRAEFFE, E. 1883b. Nachtrag zu dem Aufsatz über Polycoryne Helleri. Bolletino della Società
Adriatica di Scienze Naturali in Trieste 8: 320.

GREEFF, R. 1869. Protohydra Leuckarti. Eine marine Stammform der Coelenteraten. Zeitschrift
fiir wissenschaftliche Zoologie 20: 37-54, pl. 4-5.

GUNTHER, R. T. 1903. On the structure and affinities of Mnestra parasites Krohn, with a revision
of the classification of the Cladonemidae. Mitteilungen aus der Zoologischen Station zu
Neapel 16: 35-62.

Haprys, H., SCHIERWATER, B. & MROWKA, W. 1990. The feeding behaviour of a semi-sessile
hydromedusa and how it is affected by the mode of reproduction. Animal Behaviour 40:
935-944.

HAECKEL, E. 1879. Das System der Medusen. Erster Teil einer Monographie der Medusen.
Denkschriften der Medicinisch-Naturwissenschaftlichen Gesellschaft zu Jena 1:
XX+360 pp., 20 plates.

HAMOND, R. 1964. Further notes on the Hydrozoa of the Norfolk coast. Annals and Magazine of
Natural History 13: 659-670.

Harpy, W. B. 1891. On some points in the histology and development of Myriothela phrygia.
Quarterly journal of microscopical Science n. ser. 32: 506-537.

HARGITT, C. W. 1899. Early development of Pennaria tiarella. Science, N.Y. n. ser. 9: 368-369.

HARGITT, C. W. 1900. A contribution to the natural history and development of Pennaria tiarella
McCr. American Naturalist 34: 387-414, pls 1-4.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 399

HARGITT, C. W. 1904. The early development of Pennaria tiarella McCr. Archiv für Ent-
wicklungsmechanik der Organismen 18: 453-488.

HARGITT, G. T. 1909. Maturation, fertilization, and segmentation of Pennaria tiarella (Ayres)
and of Tubularia crocea (Ag.). Bulletin of the Museum of Comparative Zoölogy /
Harvard University 53: 159-212.

HARGITT, C. W. 1927. Some hydroids of South China. Bulletin of the Museum of Comparative
Zoölogy of Harvard College 67: 491-520.

HARTLAUB, C. 1886. Ueber den Bau der Eleutheria Quatref. Zoologischer Anzeiger 9: 706-711.

HARTLAUB, C. 1887. Zur Kenntnis der Cladonemiden. Zweite vorläufige Mittheilung. Zoo-
logischer Anzeiger 10: 651-658.

HARTLAUB, C. 1889. Ueber die Claparéde’sche "Eleutheria". Zoologischer Anzeiger 12: 665-
Oils

HARTLAUB, C. 1897. Die Hydromedusen Helgolands. Wissenschaftliche Meeresuntersuchungen
2: 449-536, pls 14-23.
HARTLAUB, C. 1899. Zur Kenntnis der Gattungen Margelopsis und Nemopsis. Nachrichten von

der Gesellschaft der Wissenschaften zu Göttingen. Mathematisch-physikalische Klasse
1899: 219-224.

HARTLAUB, C. 1903. Dendy, A., On a free-swimming hydroid, Pelagohydra mirabilis n. g., n. sp.
Zoologisches Zentralblatt 10: 27-34.

HARTLAUB, C. 1907. XII Craspedote Medusen. Teil 1, Lief. 1. Codoniden und Cladonemiden.
Nordisches Plankton 6: 1-135.

HARTLAUB, C. 1916. Ueber das Altern einer Kolonie von Syncoryne und damit verbundene
Knospungen am Hydrathenkôpfchen. Helgoländer wissenschaftliche Meeresunter-
suchungen 11: 91-125.

HARTLAUB, C. 1917. Craspedote Medusen. Teil 1, Lief. 4, Williadae. Nordisches Plankton 6:
365-479.

HAUENSCHILD, C. 1956. Experimentelle Untersuchungen über die Entstehung asexueller Klone
bei der Hydromeduse Eleutheria dichotoma. Zeitschrift für Naturforschung 11: 394-402.

HAUENSCHILD, C. 1957a. Ergänzende Mitteilung über die asexuellen Medusenklone bei
Eleutheria dichotoma. Zeitschrift für Naturforschung 12B: 412-413.

HAUENSCHILD, C. 1957b. Versuche über die Wanderung der Nesselzellen bei der Meduse
Eleutheria dichotoma. Zeitschrift für Naturforschung 12B: 472-477.

Herp, C. 1971. The succesion of benthic micrometazoans in a brackish water habitat. Biologisch
Jaarboek 39: 191-196.

HEP, C. & SMOL, N. 1976. On the importance of Protohydra leuckarti as a predator of meio-
benthic populations (pp. 285-296). In: PERSOONE, G. & JASPERS, E. (eds). Proceedings of
the 10th European Marine Biology Symposium, volume 2, Population in dynamics of
marine organisms in relation with nutrient cycling in shallow waters. Universa Press,
Wetteren, Belgium, 710 pp.

HEWITT, C. L. & GODDARD, J. H. R. 2001. A new species of large and highly contractile hydroid
in the genus Candelabrum (Hydrozoa: Anthoathecatae) from southern Oregon, U.S.A.
Canadian Journal of Zoology 79: 2280-2288.

HICKSON, S. J. 1920. On the occurrence of Protohydra in England. Quarterly Journal of Micro-
scopic Science 64: 419-424.

HINCKS, T. 1861. On Clavatella, a new genus of corynoid polypes, and its reproduction. Annals
and Magazine of Natural History (3)1: 73-81.

HINCKS, T. 1868. A history of the British hydroid zoophytes. John van Voorst, London. Volume
1, pp. lxvii +338 ; volume 2, 67 plates.

HINCKS, T. 1872b. Contribution to the History of the Hydroida. Annals and Magazine of Natural
History ser. 4 10: 385-395, pl. 20.

HIRAI, E. 1958. On the species of Cladonema radiatum var. mayeri Perkins. Bulletin of the
Marine Biological Station of Asamushi, Tohoku University 9: 23-25.

400 P. SCHUCHERT

HIROHITO, EMPEROR OF JAPAN 1977. Five hydroid species from the Gulf of Aqaba, Red Sea.
Biological Laboratory Imperial Household, Tokyo, pp. 1-26, pls 1-3.

HIROHITO, EMPEROR OF JAPAN 1988. The hydroids of Sagami Bay collected by His Majesty the
Emperor of Japan. Biological Laboratory of the Imperial Household, Tokyo, pp. 179 , 4
plates.

HOTCHKISS, A. E., MARTIN, V. J. & APKARIAN, R. P. 1984. A scanning electron microscopic sur-
face and cryofracture study of development in the planulae of the hydrozoan, Pennaria
tiarella. Scanning Electron Microscopy 2: 717-727.

Hutt, J. 1941. Ueber einige für die Westküste Schwedens neue niedere Evertebraten. Zoologiska
Bidrag fran Uppsala 20: 455-460.

Hyman, L. H. 1947. Two new Hydromedusae from the Californian coast. Transactions of the
American microscopical Society 66: 262-268.

JADERHOLM, E. 1908. Die Hydroiden des sibirischen Eismeeres, gesammelt von der Russischen
Polar-Expedition 1900-1903. Mémoires de l’Académie des Sciences de St.-Petersbourg
18: 1-26, plates.

JADERHOLM, E. 1909. Northern and arctic invertebrates in the collection of the Swedish state mu-
seum (Riksmuseum). IV Hydroiden. Bihang till Kungliga Svenska Vetenskaps-akade-
miens Handlingar 45: 1-24.

JARVIS, F. E. 1922. The hydroids from the Chagos, Seychelles and other islands and from the
coasts of British East Africa and Zanzibar. Transaction of the Linnean Society of London,
Zoology 18: 331-360, pls 24-26.

JOHANNESEN, G. 1924. Monocoryne gigantea (Bonnevie) Broch. Skrifter Norske Videnskaps-
akademi Matematisk Naturvidenskapelig Klasse 1923(18): 1-9, pl. 1-2.

Jones, N. S. 1950. Marine botton communities. Biological reviews and biological proceedings
of the Cambridge Philosophical Society 25: 283-313.

KEFERSTEIN, W. & EHLERS, E. 1861. Zoologische Beitrige gesammelt in Winter 1859-1860 in
Neapel und Messina. Leipzig, pp. vili + 112, 15 plates.

KEM, W. R. & OSTMAN, C. 1992. Occurrence of a hemolytic protein toxin in nematocysts of a
marine benthic hydroid, Halocordyle disticha (Goldfuss). Scientia Marina 56: 193-203.

KINGSLEY, J. S. 1910. A synopsis of the fixed hydroids of New England. Tufts College Studies
(Scientific Series) 3: 13-38.

KIRKENDALE, L. & CALDER, D. R. 2003. Hydroids (Cnidaria: Hydrozoa) from Guam and the
Commonwealth of the Northern Marianas Islands (CNMI). Micronesica 36: 159-188.

KIRKPATRICK, R. 1890. Reports on the zoological collections made in Torres Straits by Professor
A. C. Haddon, 1888-1889. Hydroida and Polyzoa. Proceedings of the Royal Dublin
Society, n. s. 6: 603-626.

KOLLER, G. 1927. Ueber die geschlechtliche Fortpflanzung der Protohydra leuckarti Greeff.
Zoologischer Anzeiger 73: 97-100.

KOROTNEFF, A. DE 1888. Contribution a l’étude des Hydraires. Archives de Zoologie Expéri-
mentale et Générale 6: 21-36, pls 1-2.

KRAMP, P. L. 1930. Hydromedusae collected in the south-western part of the North Sea and in
the eastern part of the Channel in 1903-14. Mémoires du Musée royal d'histoire naturelle
de Belgique 45: 1-45.

KRAMP, P. L. 1935. Polypdyr (Coelenterata) I. Ferskvandspolypper og Goplepolypper. Danmarks
Fauna 41: 1-207.

KRAMP, P. L. 1937. Polypdyr (Coelenterata), II. Gopler. Danmarks Fauna 43: 1-223.

KRAMP, P. L. 1949. Origin of the hydroid family Corymorphidae. Videnskabelige Meddelelser fra
Dansk naturhistorisk Forening i Kobenhavn 111: 183-215.

KRAMP, P. L. 1955. The medusae of the tropical West Coast of Africa. Atlantide Report 3: 239-
324, pls 1-3.

KRAMP, P. L. 1959. The Hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report
46: 1-283.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 401

KRAMP, P. L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological
Association of the U. K. 40: 1-469.

KRAMP, P. L. 1968. The Hydromedusae of the Pacific and Indian Oceans. Sections II and III.
Dana Report 72: 1-200.

KRAMP, P. L. & DAMAS, D. 1925. Les Méduses de la Norvège. Introduction et partie spéciale.
Videnskabelige meddelelser fra Dansk naturhistorik Forening 80: 217-323.

KROHN, A. 1853a. Ueber einige niedere Thiere. Archiv für Anatomie, Physiologie und wissen-
schaftliche Medicin 1853: 137-141.

KROHN, A. 1853b. Ueber die Brut des Cladonema radiatum und deren Entwickelung zum
Stauridium. Archiv fiir Anatomie, Physiologie und wissenschaftliche Medicin 1853:
420-426.

KRUMBACH, T. 1907. Beiträge zur Kenntnis der Meduse Eleutheria (Clavatella) aus dem Golfe
von Triest. PhD Thesis, printed by Wilhelm Gottlieb Korn, Breslau, pp. 47.

KÜHL, H. 1962. Die Hydromedusen der Elbmiindung. Abhandlungen und Verhandlungen des
Naturwissenschaftlichen Vereins in Hamburg 6: 209-232.

KUHL, H. 1967. Die Hydromedusen der Emsmündung. Veröffentlichungen des Instituts für
Meeresforschung in Bremerhaven 10: 239-246.

KUHL, H. 1971. Die Hydromedusen der Wesermiindung. Vie Milieu, suppl. 22: 803-810.

KUHN, A. 1910. Die Entwicklung der Geschlechtsindividuen der Hydromedusen. Studien zur
Ontogenese und Phylogenese der Hydroiden. II. Zoologische Jahrbiicher. Abteilung fiir
Anatomie und Ontogenie der Tiere 30: 43-174.

KUHN, A. 1913. Entwicklungsgeschichte und Verwandschaftsbeziehungen der Hydrozoen. I.
Teil: Die Hydroiden. Ergebnisse und Fortschritte der Zoologie 4: 1-284.

KUHN, K., STREIT, B. & SCHIERWATER, B. 1996. Homeobox genes in the cnidarian Eleutheria di-
chotoma: evolutionary implications for the origin of Antennapedia-class (HOM/hox)
genes. Molecular Phylogenetics and Evolution 6: 30-38.

LEES, D. C. 1968. The addition of the hydroid Cladocoryne floccosa to the California marine
fauna. Bulletin of the Southern California Academy of Sciences 67: 59-64.

Ley, J. 1855. Contributions towards a knowledge of the marine invertebrate fauna, of the
coasts of Rhode Island and New Jersey. Journal of the Academy of Natural Sciences of
Philadelphia, 2nd series 3: 135-152.

LELOUP, E. 1930. A propos de l’hydraire Margelopsis haeckeli Hartlaub. Annales de la Société
royale zoologique de Belgique 60: 97-100.

LELOUP, E. 1946. Margelopsis haeckeli Hartlaub, 1897, forme indicatrice du plancton au large
de la côte belge. Contribution à l’étude de la faune belge, XV. Bulletin du Musée royal
d'histoire naturelle de Belgique 22: 1-3.

LELOUP, E. 1947. Les Coelentérés de la faune Belge. Leur bibliographie et leur distribution.
Mémoires du Musée royal d'histoire naturelle de Belgique 107: 1-73.

LENGERICH, H. 1922. Zur systematische Auffassung der Eleutheriiden. Zoologischer Anzeiger
54: 209-215.

LENGERICH, H. 1923a. Vergleichende Morphologie der Eleutheriiden. Beiträge zur Kenntnis der
Eleutheriiden II. Zoologische Jahrbücher, Abteilung für Anatomie und Ontogenie der
Tiere 44: 311-388.

LENGERICH, H. 1923b. Ektodermale Medusenknospung bei Eleutheria radiata. Verhandlungen
der deutschen zoologischen Gesellschaft 28: 64-65.

LESH-LAURIE, G. E. 1976. Stolon vs. hydranth determination in Pennaria tiarella planulae: a role
for DNA synthesis (pp. 365-375). In: G.O. MACKIE (ed.). Coelenterate Ecology and
Behavior. Plenum Press, New York & London, XIII, 744 pp.

LUTHER, A. 1923. Ueber das Vorkommen von Protohydra leuckarti Greeff bei Tvärminne nebst
komplettierenden Bemerkungen über den Bau dieses Polypen. Acta Societatis pro fauna
et flora Fennica 52: 1-23.

MADSEN, H. 1939. Protohydra leuckarti Greeff, neu für Dänemark. Videnskabelige Meddelelser
fra Dansk naturhistorisk Forening i Kobenhavn 103: 551-557.

402 P. SCHUCHERT

MAIER, E. M 1974. Coelenterata. In: MORDUKHAI BOLTOVSKOGO, ED. [Ed.] [Atlas of the Aral
Sea invertebrates.]. Pishchevaya Promyshlennost, Moscow.

MAMMEN, T. A. 1963. On a collection of hydroids from South India. I. Suborder Athecata.
Journal of the Marine biological Association of India 5: 27-61.

MANTON, S. M. 1941. On the hydrorhiza and claspers of the hydroid Myriothela cocksi (Vigurs).
Journal of the Marine Biological Association of the U. K. 25: 143-150.

MARCOCI, G. 1956. Protohydra leuckarti Greeff in apele teritoriale romineti ale Mrii Negre.
Protohydra leuckarti Greeff dans les eaux roumaines de le Mer Noire. Buletinul de cerc-
etari piscicole 15: 95-98.

MARTIN, V. J. 1980. Nerve elements in the planula of the hydrozoan Pennaria tiarella. Journal
of Morphology 166: 27-36.

MARTIN, V. J. 1987. A morphological examination of gastrulation in a marine athecate hydro-
zoan. Biological Bulletin 173: 324-334.

MARTIN, V. J. 1988a. Development of nerve cells in hydrozoan planulae: 1. Differentiation of
ganglionic cells. Biological Bulletin 174: 319-329.

MARTIN, V. J. 1988b. Development of nerve cells in hydrozoan planulae: 2. Examination of sen-
sory cell differentiation using electron microscopy and immunocytochemistry.
Biological Bulletin 175: 65-78.

MARTIN, V. J. 1990. Development of nerve cells in hydrozoan planulae: 3. Some interstitial cells
traverse the ganglionic pathway in the endoderm. Biological Bulletin 178: 10-20.

MARTIN, V. J. 1991. Differentiation of the interstitial cell line in hydrozoan planulae 1.
Repopulation of epithelial planulae. Hydrobiologia

MARTIN, V. J. 1992a. Characterization of a RFamide-positive subset of ganglionic cells in the
hydrozoan planular nerve net. Cell And Tissue Research 269: 431-438.

MARTIN, V. J. 1992b. Characterization of a RFamide-positive subset of ganglionic cells in the
hydrozoan planular nerve net. Cell And Tissue Research 269: 431-438.

MARTIN, V. J. 2000. Reorganization of the nervous system during metamorphosis of a hydrozoan
planula. Invertebrate Biology 119: 243-253.

MARTIN, V. J., & ARCHER, W. E. 1986a. A scanning electron microscopic study of embryonic
development of a marine hydrozoan. Biological Bulletin of the Marine Biological
Laboratory, Woods Hole 171: 116-125.

MARTIN, V. J., & ARCHER, W. E. 1986b. Migration of interstitial cells and their derivatives in a
hydrozoan planula. Developmental Biology 116: 486-496.

MARTIN, V. J., & ARCHER, W. E. 1997. Stages of larval development and stem cell population
changes during metamorphosis of a hydrozoan planula. Biological Bulletin 192: 41-52.

MARTIN, V. J., & THOMAS, M. B. 1977. A fine structural study of embryonic and larval develop-
ment in the gymnoblastic hydroid Pennaria tiarella. Biological Bulletin of the Marine
Biological Laboratory, Woods Hole 153: 198-218.

MARTIN, V. J., & THOMAS, M. B. 1981a. Elimination of the interstitial cells in the planula larva
of the marine hydrozoan Pennaria tiarella. Journal of Experimental Zoology 217:
303-323.

MARTIN, V. J., & THOMAS, M. B. 1981b. The origin of the nervous system in Pennaria tiarella,
as revealed by treatment with colchicine. Biological Bulletin 160: 303-310.

MARTIN, V. J., & THOMAS, M. B. 1983. Establishment and maintenance of morphological
polarity in epithelial planulae. Transactions of the American Microscopical Society 102:
18-24.

MARTIN, V. J. & THOMAS, M. B. 1977. A fine structural study of embryonic and larval develop-
ment in the gymnoblastic hydroid Pennaria tiarella. Biological Bulletin of the Marine
Biological Laboratory, Woods Hole 153: 198-218.

Mayer, A. G. 1904. Medusae of the Bahamas. Memoirs of Natural Sciences / Museum of the
Brooklyn Institute of Arts and Sciences 1: 1-33, pls 1-7.

MAYER, A. G. 1910. Medusae of the world. Hydromedusae, Vols. I & II. Scyphomedusae, Vol
III. Carnegie Institution, Washington, pp. 735, plates 1-76.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 403

MCCRADY, J. 1859. Gymnopthalmata of Charleston Harbor. Proceedings of the Elliott Society of
Natural History 1: 103-221.

MEDEL, D., & LOPEZ GONZALEZ, P. J. 1996. Updated catalogue of hydrozoans of the Iberian
Peninsula and Balearic Islands, with remarks on zoogeography and affinities. Scientia
Marina 60: 183-209.

MIGOTTO, A. E. 1996. Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of Sao
Sebastiao, Brazil, including a checklist of Brazilian hydroids. Zoologische Ver-
handelingen, Leiden 306: 1-125.

MILLARD, N. A. H. 1966. The Hydrozoa of the south and west coasts of South Africa. Part III.
The Gymnoblastea and small families of the Calyptoblastea. Annals of the South African
Museum 48: 427-487, pl. 1.

MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Annals of the South
African Museum 68: 1-513.

MILLARD, N. A. H. & BOUILLON, J. 1973. Hydroids from the Seychelles (Coelenterata). Annales
du Musée Royal de l’Afrique Centrale, Sciences Zoologiques 206: 1-106, pls 1-5.

MILLARD, N. A. H. & BOUILLON, J. 1974. A collection of hydroids from Mocambique, East
Africa. Annals of the South African Museum 65: 1-40.

MonNIOT, F. 1962. Recherches sur les graviers à Amphioxus de la région de Banyuls-sur-Mer.
Vie et Milieu 13: 231-322.

MORRI, C. & BIANCHI, C. N. 1999. Hydroids (Cnidaria: Hydrozoa) from the Aegean Sea, mostly
epiphytic on algae. Cahiers de Biologie Marine 40: 283-291.

MORRI, C. & BoERO, F. 1986. Volume 7. Hydroids. Office d’Etudes Marines et Atmosphériques,

Bruxelles, 91 pp.

Motz-Kossowska, S. 1905. Contribution à la connaissance des hydraires de la Méditerranée

occidentale. I. Hydraires gymnoblastiques. Archives de Zoologie Expérimentale et

Générale (4)3: 39-98, pl. 3.

MULLER, H. 1908. Untersuchungen tiber Eibildung bei Cladonemiden und Codoniden. Zeit-

schrift fiir wissenschaftliche Zoologie 89: 28-80, pls 3-5.

MULLER, H. 1911. Beiträge zur Kenntnis der Gattung Eleutheria. Archiv fiir Naturgeschichte 77,

suppl. 1: 159-169, pl. 3.

Muus, K. 1966. Notes on the biology of Protohydra leuckarti Greef (Hydroidea, Protohydridae).
Ophelia 3: 141-150.

NAUMOV, D. V. 1955a. Tip Kishetsnopolostnie-Coelenterata (pp. 51-68, pls. 7-9). In: USHAKOV,
P.V. et al. (eds). Atlas bespozvonochnikh dal'nevostochnikh morei SSSR. Akademii Nauk
SSSR, Moscow & Leningrad, 244 pp., 66 leaves of pls.

NAUMOV, D. V. 1969. Hydroids and Hydromedusae of the USSR. Israel Program for scientific
translation, Jerusalem, 463 pp., 30 plates.

NYHOLM, K. G. 1951a. A contribution to the study of the sexual phase of Protohydra leuckarti.
Arkiv for Zoologi 2: 529-530, plate.

NYHOLM, K. G. 1951b. Egg cells in the ectoderm of Boreohydra simplex. Arkiv for Zoologi 2:
531-533.

OsTMAN, C., PIRAINO, S. & Kem, W. 1991. Nematocysts of the Mediterranean hydroid Halo-
cordyle disticha. Hydrobiologia 216-217: 607-613.

OMER-COOPER, J. 1964. On Protohydra psamathe n. sp. from South Africa. Journal of the
Linnean Society, London, Zoology 45: 145-150.

PARDY, R. L. & LENHOFF, H. M. 1968. The feeding biology of the gymnoblastic hydroid,
Pennaria tiarella. Journal of experimental Zoology 168: 197-202.

PASTEELS, J. 1939. Variations et gradient axial chez l’hydroméduse Cladonema radiatum.
Annales de la Société royale zoologique de Belgique 69: 311-325.

PASTEELS, J. 1941. Régénération de l’hydranthe et de la méduse chez Cladonema radiatum.
Annales de la Société royale zoologique de Belgique 72: 63-73.

PENA CANTERO, A. L. & GARCIA CARRASCOSA, A. M. 2002. The benthic hydroid fauna of the
Chafarinas Islands (Alboran Sea, western Mediterranean). Zoologische Verhandelingen
337: 1-180.

404 P. SCHUCHERT

PERKINS, H. F. 1902. Notes on the anatomy and histology of a new form of Cladonema from the
Bahamas. Johns Hopkins University Circulars 21: 25-27, pl. 1.

PERRIER, E. 1886. Les explorations sous-marines. Hachette, Paris, pp. 352.

PETERSEN, C. G. J. 1913. Valuation of the sea. II The animal communities of the sea bottom and
their importance for marine zoogeography. Report for the Danish Biological Station 21:
1-42.

PETERSEN, K. W. 1990. Evolution and taxonomy in capitate hydroids and medusae (Cnidaria:
Hydrozoa). Zoological Journal of the Linnean Society 100: 101-231.

PHILBERT, M. 1936. Etudes sur Cladocoryne floccosa Rotch. Bulletin de l’Institut océano-
graphique de Monaco 708: 1-16.

PICARD, J. 1957. Etude sur les hydroïdes de la superfamille Pteronematoidea, 1. Généralités.
Bulletin de l’Institut océanographique de Monaco 1106: 1-12.

PICARD, J. 1958. Notes sur une collection d’hydroides provenant des côtes méditerranéennes
d’Israel. Bulletin of the Sea Fisheries Research Station, Israel 15: 1-3.

PICTET, C. 1893. Etude sur les hydraires de la Baie d’Amboine. Revue suisse de Zoologie 1:
1-64.

POCHE, F. 1914. Das System der Coelenterata. Archive für Naturgeschichte 80: 47-128.

PRÉVOT, E. 1959. Morphologie et évolution des structures tentaculaires chez les hydraires gym-
noblastes Capitata. Recueil des Travaux de la station marine d’Endoume 29: 91-128.

QUATREFAGES, A. DE 1842a. Mémoire sur l’Eleutherie dichotome (Eleutheria dichotoma nob.),
nouveau genre de Rayonnés, voisin des Hydres. Annales des Sciences naturelles (2)18:
270-288, plate 8.

QUATREFAGES, A. DE 1842b. Mémoire sur l'Eleutherie dichotome (Eleutheria dichotoma A. de
Q.). Nouveau genre de Rayonnés, voisin des Hydres. Comptes rendus hebdomadaires
des Séances de l’Académie des Sciences, Paris 15: 168-170.

RALPH, P. M. 1953. A guide to the athecate (Gymnoblastic) hydroids and medusae of New
Zealand. Tuatara 5: 59-75.

RASMUSSEN, E. 1973. Systematics and ecology of the Isefjord marine fauna (Denmark). With a
survey of the eelgrass (Zostera) vegetations and its communities. Ophelia 11: 1-495.

REES, W. J. 1941. Notes on British and Norwegian hydroids and medusae. Journal of the Marine
Biological Association of the United Kingdom 25: 129-141.

REES, W. J. 1949. On Cladonema myersi, a new species of hydroid from the Californian coast.
Proceedings of the Zoological Society of London 119: 861-865.

REES, W. J. 1956. A revision of some northern gymnoblastic hydroids in the Zoological Museum,
Oslo. Nytt Magasin for Zoologi 4: 109-120.

REES, W. J. 1957. Evolutionary trends in the classification of capitate hydroids and medusae.
Bulletin of the British Museum (Natural History). Zoology series 4: 455-534.

REES, W. J. 1958. The status of the hydroid Symplectanea bracteata Fraser. Nytt magasin för
Zoologi S: 17-19.

REES, W. J. & THURSFIELD, S. 1965. The hydroid collections of James Ritchie. Proceedings of
the Royal Society of Edinburgh (B) 69: 34-220.

REES, J. T. 1979. The polyp and immature medusa stages of Cladonema californicum, Hyman,
1947 (Anthomedusae: Cladonemidae) with biological notes and a discussion of the taxo-
nomy of the genus Cladonema. Journal of Natural History 13: 295-302.

RICHTERS, F. 1908. Cladonema radiatum bei Helgoland. Zoologischer Anzeiger 33: 687.

RIEDL, R. 1970. Fauna und Flora der Adria, ed. 2: 1-640, col. pls 1-8, pls 1-235, figs. Verlag Paul
Parey, Hamburg & Berlin.

RITCHIE, J. 1910. The marine fauna of the Mergui Archipelago, lower Burma etc.- the Hydroids.
Proceedings of the Zoological Society of London 1910: 799-825, plates.

RoTcH, W. D. 1871. On a new genus and species of Hydroid Zoophytes (Cladocoryne floccosa).
Annals and Magazine of Natural History (4) 7: 227-228.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 405

RUEBUSH, T. K. 1939. The occurrence of the two rare genera, Protohydra and Protodrilus on the
East coast of North America. Science, N.Y. 90: 617-618.

RUSSELL, F. S. 1953. The medusae of the British Isles. Cambridge University Press, London, pp.
530, 35 pls.

SALVINI-PLAWEN, L. v. 1966. Zur Kenntnis der Cnidaria des nordadriatischen Mesopsammon.
Veröffentlichungen des Instituts für Meeresforschung in Bremerhaven, Sonderband 2:
165-186.

SARS, G. O. 1874. Bidrag til Kundskaben om Norges Hydroider. Forhandelinger i Videnskabs-
Selskabet i Kristiana (1873): 91-150, pls 2-5.

SARS, M. 1850. Beretning om en i sommeren 1849 foretagen zoologisk reise i Lofoten og
Finmarken. Christina, 91 pp.

SARS, M. 1874. In: SARS, G. O. 1874.

SARS, M. 1877. Nye og mindre bekjendte Coelenterater. - New and little known Coelenterates
(pp. 32, pls 1-2). In: J. KOREN & D. C. DANIELSSEN (eds). Fauna littoralis Norvegiae
Bergen.

SCHEURING, L. 1922. Die Hydroiden. Untersuchungsfahrt des Reichsforschungsdampfer
"Poseidon" in das Barentsmeer im Juni and Juli 1913. Wissenschaftliche Meeresunter-
suchungen. Abt. Helgoland. 13: 157-179.

SCHIERWATER, B. 1989. Allometric changes during growth and reproduction in Eleutheria di-
chotoma (Hydrozoa, Athecata) and the problem of estimating body size in a microscop-
ic animal. Journal of Morphology 200: 255-267.

SCHIERWATER, B. & HADRYS, H. 1998. Environmental factors and metagenesis in the hydroid
Eleutheria dichotoma. Invertebrate Reproduction and Development 34: 139-148.
SCHIERWATER, B. & HAVENSCHILD, C. 1990. The position and consequences of a vegetative mode
of reproduction in the life cycle of a hydromedusa and an oligochaete worm. Advances

In Invertebrate Reproduction 5: 37-42.

SCHIERWATER, B., MURTHA, M., DICK, M., RUDDLE, F. H. & Buss, L. W. 1991. Homeoboxes in
cnidarians. Journal of Experimental Zoology 260: 413-416.

SCHIERWATER, B., PIEKOS, B. & Buss, L. W. 1992. Hydroid stolonal contractions mediated by
contractile vacuoles. Journal of Experimental Biology 162: 1-21.

SCHMIDT, O. 1869. Leben der Krebse, Würmer und ungegliederte Wirbellosen Thiere. /n:
BREHM, A.E. Illustrirtes Thierleben. Hildburghausen. 6 volumes.

SCHNEIDER, G. 1927. Neue Fundorte fiir Protohydra leuckarti Greeff bei der zoologischen
Station Tvärminne. Memoranda Societatis pro Fauna Flora Fennica 2: 8-10.

SCHÖNBORN, C., ARNDT, E. A. & GOSSELCK, F. 1993. Bestimmungsschlussel der benthischen
Hydrozoen der Ostsee. Mitteilungen aus dem Zoologischen Museum in Berlin 69:
201-253.

SCHUCHERT, P. 1996. The marine fauna of New Zealand: athecate hydroids and their medusae
(Cnidaria: Hydrozoa). New Zealand Oceanographic Institute Memoir 106: 1-159.

SCHUCHERT, P. 2000. Hydrozoa (Cnidaria) of Iceland collected by the BIOICE programme.
Sarsia 85: 411-438.

SCHUCHERT, P. 2001a. Hydroids of Greenland and Iceland (Cnidaria, Hydrozoa). Meddelelser om
Grönland, Bioscience 53: 1-184.

SCHUCHERT, P. 2001b. Survey of the family Corynidae (Cnidaria, Hydrozoa). Revue suisse de
Zoologie 108: 739-878.

SCHUCHERT, P. 2003. Hydroids (Cnidaria, Hydrozoa) of the Danish expedition to the Kei Islands.
Steenstrupia 27: 137-256.

SCHUCHERT, P. 2004. Revision of the European athecate hydroids and their medusae (Hydrozoa,
Cnidaria): Families Oceanidae and Pachycordylidae. Revue suisse de Zoologie 111:
315-369.

SCHUCHERT, P. 2005. Species boundaries in the hydrozoan genus Coryne. Molecular Phylo-
genetics and Evolution 36: 194-199.

406 P. SCHUCHERT

SCHULZ, E. 1950a. Zur Oekologie von Protohydra leuckarti Greef. (Studien an Hydrozoa, L.).
Kieler Meeresforschung 7: 53-57.

SCHULZ, E. 1950b. Psammohydra nanna, ein neues solitàres Hydrozoon in der westlichen
Beltsee. (Studien an Hydrozoa, II). Kieler Meeresforschung 7: 122-137.

SCHULZ, E. 1952. Ungeschlechtliche Fortpflanzung durch Knospung bei Protohydra leuckarti
Greef (Studien an Hydrozoa, III). Kieler Meeresforschung 7: 68-69.

ScHULZE, F. E. 1876. Tiarella singularis, ein neuer Hydroidpolyp. Zeitschrift für wissen-
schaftliche Zoologie 27: 403-416.

SEGONZAC, M. & VERVOORT, W. 1995. First record of the genus Candelabrum (Cnidaria,
Hydrozoa, Athecata) from the Mid-Atlantic Ridge: a description of a new species and a
review of the genus. Bulletin du Museum national d'histoire naturelle. Section A,
Zoologie, biologie et écologie animales 17: 31-63.

SEGERSTEDT, M. 1889. Bidrag till Kännedomen om Hydoid-Faunan vif Sveriges Vestkust.
Bihang till Kungliga Svenska Vetenskaps-akademiens Handlingar 14(4): 1-28, plate.

SPAGNOLINI, A. 1876. Catalogo sistematico degli Acalefi del Mediterraneo. Sifonofori e Meduse
Craspedote. Atti della Società italiana di scienze naturali 19: 291-333.

SPRACKLIN, B. W. 1982. Hydroidea (Cnidaria: Hydrozoa) from Carrie Bow Cay, Belize.
Smithsonian Contribution to Marine Science 12: 239-251.

STECHOW, E. 1909. Hydroidpolypen der japanischen Ostkueste. I. Teil: Athecata und Plumu-
laridae. In: DOFLEIN, F. (ed.). Beiträge zur Naturgeschichte Ostasiens. Abhandlungen der
Mathematisch-Phyikalische Klasse der Königlichen Bayerischen Akademie der
Wissenschaften, Supplement Band 1: 1-111, plates 1-7.

STECHOW, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer
Gebiete, nebst Angaben über einige Kirchenpauer’sche Typen von Plumulariden.
Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere
42: 1-172.

STECHOW, E. 1920. Neue Ergebnisse auf dem Gebiete der Hydroidenforschung. Sitzungsberichte
der Gesellschaft für Morphologie und Physiologie in München 31: 9-45.

STECHOW, E. 1921. Neue Genera und Species von Hydrozoen und anderen Evertebraten. Archiv
für Naturgeschichte 87: 248-265.

STECHOW, E. 1923. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer
Gebiete. II. Teil. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und
Biologie der Tiere 47: 29-270.

STECHOW, E. 1925. Hydroiden von West- und Südwestaustralien nach den Sammlungen von
Prof. Dr. Michaelsen und Prof. Dr. Hartmeyer. Zoologische Jahrbücher. Abteilung für
Systematik, Oekologie und Geographie der Tiere 50: 191-270.

STEPANJANTS, S. D., SHEIKO, O. V. & NAPARA, T. O. 1990. Fabulosus kurilensis gen. et sp. n.
(Hydrozoa, Chidaria [Cnidaria]), a new nydroid [hydroid] polyp of the Kuril Islands
shelf. Trudy Zoologicheskogo Instituta 218: 5-17.

STEPANJANTS, S. D., OVANESYAN, I. G. & SIRENKO, B. I. 1993. Cladonematidae representative
(Hydrozoa, Athecata, Capitata) in Saint-Petersburg aquariums. Cladonema revision
[Russian with English summary]. /ssledovaniya Fauny Morei 45: 90-105.

STEPANJANTS, S. D., CHRISTIANSEN, B. O., SVOBODA, A. & ANOKHIN, B. A. 2003. The genus
Monocoryne (Hydrozoa, Capitata): peculiarities of morphology, species composition,
biology and distribution. Sarsia 88: 97-106.

STIMPSON, W. 1854. Synopsis of the marine Invertebrata of Grand Manan, or the region about
the mouth of the Bay of Fundy, New Brunswick. Smithsonian Contributions to
Knowledge 6: 1-67, pls 1-3.

SUMMERS, R. G. 1970. The fine structure of the spermatozoa of Pennaria tiarella (Coelenterata).
Journal of Morphology 131: 117-130.

SUMMERS, R. G., & HAYNES, J. F. 1969. The ontogeny of interstitial cells in Pennaria tiarella.
Journal of Morphology 129: 81-87.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 407

SUMMERS, R. S. 1976. An ultrastructural investigation of the striated subumbrellar structure of
the Anthomedusan, Pennaria tiarella. Journal of Morphology 149: 507-525.

SUMMERS, R. G. & HAYNES, J. F. 1969. The ontogeny of interstitial cells in Pennaria tiarella.
Journal of Morphology 129: 81-87.

SWEDMARK, B. 1956. Étude de la microfaune des sables marins de la région de Marseille.
Archives de Zoologie expérimentale et génerale, 93, notes et revue 2: 70-95.

SWEDMARK, B. 1959. On the biology of sexual reproduction of the interstitial fauna of marine
sand. Proceedings of the XV International Congress of Zoology, 327-330.

SWENANDER, G. 1904. Ueber die Athecaten Hydroiden des Drontheimsfjordes. Kongelige Norske
Videnskabers Selskab Skrifter 1903: 1-16, pl. 1.

SWOFFORD, D. L. 2001. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods).
Version 4. Sinauer Associates, Sunderland, Massachusetts.

TARDENT, P. 1963. Regeneration in the Hydrozoa. Biological Reviews 38: 293-333.

TARDENT, P. 1965. Developmental aspects of regeneration in Coelenterates (pp. 71-88). In:
KIORTIS, V. & TRAMPUSCH, H.A.L. (eds). Regeneration in Animals and related problems.
North-Holland Press, Amsterdam.

TARDENT, P. 1978. Coelenterata, Cnidaria (pp. 71-415). In: SEIDEL, F. (ed). Morphogenese der
Tiere. Gustav Fischer, Stuttgart.

TARDENT, P. & STÔSSEL, F. 1971. Die Mechanoreceptoren der Polypen von Coryne pintneri,
Sarsia reesi und Cladonema radiatum (Athecata, Capitata). Revue suisse de Zoologie 78:
680-688.

TEISSIER, G. 1965. Inventaire de la faune marine de Roscoff. Cnidaires-Cténaires. Travaux de la
Station Biologique de Roscoff 16: 1-53.

THIEL, M. E. 1938. Die Leptolinae der "Meteor" Expedition in systematischer Betrachtung
(I. Anthomedusae). Zoologischer Anzeiger 121: 289-303.

THIEL, M. E. 1935. Zur Kenntnis der Hydromedusenfauna des Schwarzen Meeres. Zoologischer
Anzeiger 111: 161-174.

THIEL, H. 1988. Cnidaria (pp. 266-272). In: HIGGINS, R. P. & THIEL, H. (eds). Introduction to the
study of meiofauna. Smithsonian Institution Press, Washington, USA.

THOMAS, M. B., EDWARDS, N. C. & NORRIS, T. A. 1987. Gastrulation in Halocordyle disticha
(Hydrozoa, Athecata). International Journal of Invertebrate Reproduction And
Development 12: 91-102.

THOMAS, M. B., Epwarps, N. C. & HIGGINS, R. P. 1995. Cryptohydra thieli n. gen., n. sp.: a
meiofaunal marine hydroid (Hydroida, Athecata, Capitata). Invertebrate Biology 114:
107-118.

Totton, A. K. 1965. A synopsis of the Siphonophora. British Museum (Natural History),
London, 230 pp.

UCHIDA, T. 1924. On a new pelagic Hydroid, Climacocodon ikarii n. gen. n. sp. Japanese
Journal of Zoology 1: 59-65.

UCHIDA, T. 1927. Studies on Japanese hydromedusae. I. Anthomedusae. Journal of the Faculty
of Science, Imperial University of Tokyo, Section IV. Zoology 1: 145-241, plates 10-11.

VALKANOV, A. 1947. Protohydra leuckarti Greeff ot Cherno more. Protohydra leuckarti Greeff
vom Schwarzen Meer. Trudove ot Chernomorska Biologichna Stantsiya (Arbeiten aus
der Biologischen Meeresstation am Schwarzen Meer in Varna) 13: 31-35.

VAN BENEDEN, P. J. 1867. Recherches sur la faune littorale de Belgique (polypes). Mémoires de
l'Académie Royale des Sciences et Belles-Lettres de Belgique 36: 1-207.

VEVERS, H. G. 1959. Birth of a Tricyclusa. New Scientist 6: 506-507.

VAN DE VYVER, G. 1968. Etude du développement embryonnaire des hydraires athécates (Gym-
noblastiques) à gonophores. 2. Formes à actinulas. Archives de Biologie 79: 327-363.

VAN DE VYVER, G. & BOUILLON J. 1969. Etude du développement embryonnaire et de l’histo-
genèse de Eleutheria dichotoma (De Quatrefages) (Anthoméduse Eleutheriidae).
Annales d’embryologie et de morphogenèse 2: 317-327.

408 P. SCHUCHERT

VERRILL, A. E. 1878. Notice of recent additions to the marine fauna of the eastern coast of North
America. No. 2. Brief contributions to zoology from the Museum of Yale College. No.
39. American Journal of Science (3)16: 371-378.

VERVOORT, W. 1946. Hydrozoa (C 1) A. Hydropolypen. Fauna van Nederland 14: 1-336.

VERVOORT, W. 1968. Report on a collection of Hydroida from the Caribbean region, including an
annotated checklist of Caribbean hydroids. Zoologische Verhandelingen, Leiden 92:
1-124.

VERVOORT, W. 1993. Report on hydroids (Hydrozoa, Cnidaria) in the collection of the Zoological
Museum, University of Tel-Aviv, Israel. Zoologische Mededelingen 67: 537-565.
VERWOORT, W. V. 1985. Deep-water hydroids (pp. 267-297). In: LAUBIER, L. & MONNIOT, C.
(eds). Peuplements profonds du golfe de Gascogne: Campagnes BIOGAS. [Deep sea
populations from the Gulf of Gascogne: BIOGAS expeditions.] Ifremer, Brest, 629 pp.

[incorrect spelling of name]

VIGURS 1849/1850. In: Cocks, W. P. Contributions to the fauna of Falmouth. Report of the Royal
Cornwall Polytechnic Society 17: 90. [publication date unclear, likely 1850, see
Cornelius (1977)]

WATSON, J. E. 1999. Hydroids (Hydrozoa: Anthoathecata) from the Beagle Gulf and Darwin
Harbour, northern Australia. The Beagle, Records of the Museums and Art Galleries of
the Northern Territory 15: 1-21.

WEBER, C. 1980. Structural organization, ontogeny and regeneration of the eye of Cladonema
radiatum (Anthomedusae) (pp. 347-352). In: TARDENT, P. & TARDENT, R. (eds).
Developmental and cellular biology of coelenterates. Proceedings of the 4th Inter-
national Coelenterate Conference, Interlaken, Switzerland. Elsevier/Biomedical Press,
Amsterdam, New York & Oxford, pp. 499.

WEBER, C. 1981a. Lens of the hydromedusan Cladonema studied by SDSgel-Electrophoresis
and immunofluorescent technique. Journal of Experimental Zoology 217: 15-21.
WEBER, C. 1981b. Structure, histochemistry, ontogenetic development, and regeneration of the
ocellus of Cladonema radiatum Dujardin (Cnidaria, Hydrozoa, Anthomedusae). Journal

of Morphology 167: 313-331.

WEBER, C. & TARDENT, P. 1978. Zur Entwicklung des Linsenauges von Cladonema radiatum
Duj. (Hydrozoa, Anthomedusae). Revue suisse de Zoologie 85: 762-767.

WEDLER, E. & LARSON, R. 1986. Athecate hydroids from Puerto Rico and the Virgin Islands.
Studies on Neotropical Fauna and Environment 21: 69-101.

WEHLING, W. 1978. Observations on the reproductive biology of Protohydra leuckarti.
American Zoologist 18: 643.

WEILER-STOLT, B. 1960. Ueber die Bedeutung der interstitiellen Zellen für die Entwicklung und
Fortpflanzung mariner Hydroiden. Wilhelm Roux' Archiv 152: 398-455.

WEILL, R. 1931. Le genre Preroclava n. gen., l’interprétation systématique des Ptéronemidae
(Hydraires) et la valeur taxonomique du cnidome. Comptes rendus hebdomadaires des
Séances de l’Académie des Sciences, Paris 192(2): 60-62.

WEILL, R. 1934. Contribution à l’étude des Cnidaires et de leurs nématocystes. II. Valeur taxo-
nomique du cnidôme. Travaux de la Station zoologique de Wimereux 11: 349-701.

WEILL, R. 1935. Halammohydra Remane, genre d’hydrozoaires aberrant. Son cnidome et sa
position taxonomique: Affinités avec les Siphonophores Chondrophorides et avec Proto-
hydra Leuckarti Greeff. Bulletin de la Société Zoologique de France 60: 73-87.

WEILL, R. 1936. Le cnidòme des Cladonémides Eleutheria dichotoma et Cladonema radiatum,
son cycle et son interprétation. Comptes rendus hebdomadaires des Séances de l’Aca-
démie des Sciences, Paris 203: 816-818.

WEILL, R. 1937. Existence d’un monocnidome dans le médusoide d’un polype (Pennaria? tiarel-
la McCr) à tétracnidome. Comptes rendus hebdomadaires des Séances de l’Académie
des Sciences, Paris 204: 1749-1752.

WEISMANN, A. 1883. Die Entstehung der Sexualzellen bei den Hydromedusen. Zugleich ein
Beitrag zur Kenntnis des Baues und der Lebensgeschichte dieser Gruppe. Gustav
Fischer, Jena, pp. 295.

THE EUROPEAN ATHECATE HYDROIDS AND THEIR MEDUSAE 409

WERNER, B. 1954. Ueber die Fortpflanzung der Anthomeduse Margelopsis haeckeli Hartlaub
durch Subitan- und Dauereier und die Abhängigkeit ihrer Bildung von äusseren Fak-
toren. Verhandlungen der Deutschen Zoologischen Gesellschaft in Tiibingen 1954:
124-133.

WERNER, B. 1955. On the development and reproduction of the Anthomedusan Margelopsis
haeckeli Hartlaub. Annals of the New York Academy of Sciences 62: 1-30.
WERNER, B. 1956. Der zytologische Nachweis der parthogenetischem Entwicklung bei der
Anthomeduse Margelopsis haeckeli Hartlaub. Naturwissenschaften 43: 541-542.
WESTBLAD, E. 1930a. Protohydra Leuckarti an der Swedischen Küste entdeckt. Arkiv för
Zoologi 21: 1-4.

WESTBLAD, E. 1930b. Ueber die Geschlechtsorgane und die systematische Bedeutung von Proto-
hydra Leuckarti Greeff. Arkiv for Zoologi 21: 1-13.

WESTBLAD, E. 1935. Neue Beobachtungen über Protohydra. Zoologischer Anzeiger 111:
152-158.

WESTBLAD, E. 1937. Boreohydra simplex n. gen., n. sp., ein Solitàrpolyp von der norwegischen
Kiiste. Arkiv for Zoologi 29: 1-6.

WESTBLAD, E. 1947. Notes on hydroids. Arkiv för Zoologi 39: 1-23.

WESTBLAD, E. 1953. Boreohydra simplex Westblad, a "bipolar" hydroid. Arkiv for Zoologi, new
series 4: 351-354.

WIESER, W. 1958. Occurrence of Protohydra leuckarti in Puget Sound. Pacific Science 12:
106-108.

WILL, L. 1913. Acaulis primarius Stimpson, ein neuer Ostsee Bewohner. Sitzungsberichte und
Abhandlungen der naturforschenden Gesellschaft zu Rostock, neue Serie 5: 57-62.

WIRTZ, P. & DEBELIUS, H. 2003. Mediterranean and Atlantic Invertebrate Guide. ConchBooks,
Hackenheim, Germany, pp. 1-305.

INDEX

Acandela, 341

Acaulidae, 326

Acaulis, 328

Acauloides, 330

Allmani, Cladonema, 376
ammisatum, Acauloides, 330
anisonema, Eleutheria, 381
Arum, 341

Blastothela, 328
Boreohydra, 334
Boreohydridae, 334
californicum, Cladonema, 379
Candelabridae, 340
Candelabrum, 341
Cladocoryne, 368
Cladocorynidae, 368
Cladocorynopsis, 368
Cladonema, 375
cladonema, Stauridium, 376
Cladonematidae, 371
claparedei, Eleutheria, 384
claparedii, Eleutheria, 384
Clavatella, 380
Climacocodon, 356
cnidobdella, Eleutheria, 381
Cnidonema, 373

cocksii, Candelabrum, 341
corynaria, Hydra, 370
Corynidae, 380
Dendronema, 391
Dendronemidae, 371
dichobdella, Eleutheria, 381
dichocnida, Eleutheria, 381
dichotoma, Eleutheria, 381
diplonema, Eleutheria, 384
disticha, Pennaria, 364
Dujardinii, Cladonema, 376
Eleutheria, 380
Eleutheriidae, 371

Eucoryne, 364

floccosa, Cladocoryne, 368
fritchmanii, Candelabrum, 346
Gegenbauri, Cladonema, 376
gemmipara, Eleutheria, 381
gigantea, Monocoryne, 381
Globiceps, 364

haeckelii, Margelopsis, 356
Halocordyle, 364
Halocordyle disticha, 364, 367
Halocordylidae, 363
hartlaubii, Margelopsis, 361
helleri, Polycoryne, 370

410

heptanema, Eleutheria, 381
Herpusa, 380

heteroclada, Eleutheria, 381
heteronema, Eleutheria, 384
hexanema, Eleutheria, 381
Hydra corynaria, 370

ilonae, Acauloides, 332
krohni, Eleutheria, 381
Krohnii, Cladonema, 376
leuckarti, Protohydra, 338
Lobocoryne, 368
Margelopsidae, 356
Margelopsis, 356

mayeri, Cladonema, 376
minor, Monocoryne, 353
Monocoryne, 351

myersi, Cladonema, 380
Myriocnida, 328

Myriothela, 340
Myriothelidae, 340

nanna, Psammohydra, 336
nematophora, Similomerona, 325
novae-zelandiae, Cladonema, 376
novaezelandiae, Cladonema, 380
octonema, Eleutheria, 381
pacificum, Cladonema, 380
pelagica, Cladocoryne, 370
Pelagohydra, 356
Pelagohydridae, 356
Pennaria, 364

Pennariidae, 363

pentanema, Eleutheria, 381
perkinsii, Cladonema, 376
phrygium, Candelabrum, 346
Polycoryne, 368

P. SCHUCHERT

portmanni, Staurocladia, 389
primarius, Acaulis, 328
prolifera, Clavatella, 381
Protohydra, 338
Protohydridae, 337
Psammocoryne, 330
Psammohydra, 336

purpurea, Spadix, 341
radiatum, Cladonema, 375
sargassensis, Cladocoryne, 370
serpentarii, Candelabrum, 341
Similomerona, 325

simplex, Boreohydra, 334
simplex, Cladocoryne, 370
singularis, Tricyclusa, 353
Spadix, 341

Stauridia, 375

stauridia, Coryne, 376
Stauridium, 375

stauridium, Syncoryne, 376
Stauriidae, 371

Staurocladia, 372, 386
stylodendron, Dendronema, 391
stylostoma, Margelopsis, 353
Symplectanea, 351
Symplectaneidae, 340
tetranema, Eleutheria, 381
Tiarella, 353

Tricyclusa, 353

Tricyclusidae, 353

uchidai, Cladonema, 380
ulvae, Herpusa, 381

vallentini, Eleutheria, 386
verrucosum, Candelabrum, 349
Wandelia, 386

REVUE SUISSE DE ZOOLOGIE 113 (2): 411-419; juin 2006

Identification biométrique des deux espèces sympatriques de souris
Mus musculus domesticus et Mus spretus en Kabylie du Djurdjura
(Algérie)

Nora KHAMMES!, Sovan LEK? & Stéphane AULAGNIER3

l Faculté des Sciences Biologiques et Agronomiques, Université Mouloud Mammeri,
route d’Hassnaoua, 15000 Tizi-Ouzou, Algérie. E-mail: n_khammes @hotmail.com

2 Laboratoire Dynamique de la Biodiversité, Université Paul Sabatier, 118 route de
Narbonne, F-31062 Toulouse cedex 4, France. E-mail: lek@cict.fr

3 Comportement et Ecologie de la Faune Sauvage, B.P. 52627, F-31326 Castanet-
Tolosan cedex, France. E-mail: aulagnie @toulouse.inra.fr

Biometrical identification of the sympatric mouse species Mus muscu-
lus domesticus and Mus spretus in Kabylie du Djurdjura (Algeria). -
This work aimed to validate biometric identification criteria (body and
cranial measurements) for the two sympatric mouse species living in
Algeria: the House mouse (Mus musculus domesticus) and the Algerian
mouse (M. spretus). Trapped in three localities of Kabylie du Djurdjura,
mice were first identified according to morphological criteria and second
computing stepwise discriminant analyses and leave-one-out discriminant
analyses on sets of the most relevant variables. Body measurements re-
vealed more efficient than cranial measurements to discriminate the species.
According to the original diagnosis of M. spretus, the Algerian mouse’s tail
was shorter than the House mouse’s tail. The ratio ‘tail length over head and
body length’ was highly discriminant (95% of correct classification); the
score was still better (97%) when the diameter of the tail was included in the
discriminant analysis. On the skull, the zygomatic ratio ‘width of the dorsal
ramus of the zygomatic arch over width of the zygomatic arch’ was the only
discriminant variable (90.4% of the correct classification). No identification
was reliable with mandible measurements. Relevant discriminant values
were then compared with values obtained from European samples.

Keywords: Algeria - biometrics - skull - discriminant analysis - Mus spretus
- Mus musculus domesticus.

INTRODUCTION

En 1846 déjà, Loche identifie deux espèces de souris en Algérie: Mus musculus
et Mus reboudi, mais il s’est avéré par la suite que ces deux formes étaient conspéci-
fiques. Puis, toujours en Algérie, Lataste (1883) décrit Mus spretus comme une
nouvelle espéce, caractérisée par une oreille elliptique, une queue environ moitié plus

Manuscript accepted 28.11.2005

412 N. KHAMMES, S. LEK & S. AULAGNIER

courte que le corps et un talon supplémentaire en avant de la première molaire. Mais
plus tard (Lataste, 1887), il exprime des doutes à propos de cette différence spécifique,
aussi par la suite spretus a souvent été considérée comme une sous-espèce d’une autre
souris à queue courte Mus spicilegus Petenyi, 1882 (e.g. Miller, 1912; Cabrera, 1914,
1932) ou de Mus musculus Linnaeus, 1766 (e.g. Schwarz & Schwarz, 1943; Ellerman
& Morrison-Scott, 1951).

Le statut spécifique de Mus spretus Lataste, 1883 a été rétabli lorsque vers la fin
des années 70 deux espèces sympatriques de souris Mus musculus domesticus Rutty,
1772 et Mus spretus ont été différenciées génétiquement dans le sud de la France
(Britton et al., 1976, Britton & Thaller, 1978). Deux interrogations étroitement liées se
sont immédiatement imposées: comment les identifier? où les trouver? Très rapi-
dement, en étudiant la morphométrie externe et crânienne des animaux identifiés sur la
base de leur polymorphisme enzymatique, des critères d’identification ont été proposés
ou confirmés (Darviche, 1978; Marshall & Sage, 1981; Orsini, 1982; Darviche &
Orsini, 1982). Parallèlement, l’examen des spécimens conservés en collection a permis
de préciser l’aire de répartition de l’espèce en Afrique du Nord, dans la péninsule
Ibérique et en France (Saint Girons & Thouy, 1978; Palomo et al., 1983). Puis des
travaux sur des populations naturelles et expérimentales ont permis d’appréhender les
facteurs écologiques et comportementaux à l’origine de la sympatrie, voire de la
syntopie de la Souris à queue courte avec le morphe (brevirostris) ou écotype sauvage
de la Souris domestique, caractéristique de la région méditerranéenne (Orsini et al.,
1982, Cassaing, 1982, 1984; Cassaing & Croset, 1985).

De récentes synthèses ont fait le point sur l’identification des souris euro-
péennes (Gerasimov et al., 1990; Macholän, 1996a, 1996b) ou françaises (Orsini et al.,
2001), mais aucune étude approfondie n’a été consacrée aux souris d’ Afrique du Nord,
alors que la plus grande diversité génétique locale des Mus spretus (Boursot et al.,
1985) peut laisser envisager une variabilité morphologique particulière.

C’est ce travail de calibration des critères d’identification morphométriques
(biométrie externe et crânienne) des deux souris présentes en Algérie qui a été entre-
pris à partir d’un échantillon conséquent en provenance de trois stations de Kabylie du
Djurdjura.

MATÉRIEL ET MÉTHODES

LOCALISATION DES STATIONS D'ÉTUDE

Station I: village de Boukhalfa à 8 km de Tizi-Ouzou (36°42’ N, 4°2’ E;
altitude: 500 m). Cette station est classée dans l’étage bioclimatique subhumide à hiver
chaud. Située à proximité d’un verger, la végétation y est composée d’une strate
arbustive haute à Genista tricuspidata et Calycotome spinosa, et une strate arbustive
basse à Rubus sp. et Cistus salvifolius.

Station 2: village d’Attouche (commune de Makouda) à 25 km de Tizi-Ouzou
(36°47’N, 4°3’ E; altitude: 470 m). Cette station est classée dans |’ étage bioclimatique
subhumide à hiver doux. Délimitée par de grands rochers, cette station s’étend sur deux
parcelles, l’une comprend essentiellement une strate herbacée et des affleurements
rocheux, l’autre est occupée pour les grandes cultures céréalières, maraîchères et
d’arbres fruitiers.

BIOMÉTRIE DISCRIMINANTE DES SOURIS D’ ALGERIE 413

Station 3: village de Bou-Ilfane, au nord-est d’Azazga, à environ 45 km de
Tizi-Ouzou (36°45’ N, 4°27’ E; altitude : 500 m). Cette station est classée dans l’étage
bioclimatique subhumide à variante tempérée. C’est un maquis, formation dense issue
de la dégradation de la forêt de Chêne liège Quercus suber, qui comprend principa-
lement une strate arbustive constituée de Myrtus communis, Genista tricuspidata,
Cistus monspeliensis, Erica arborea et Calycotome spinosa.

ANIMAUX

Le nombre total d’individus capturés au moyen de surfaces engluées (Tripathi
et al., 1994) est de 138 souris des deux espèces pour 1200 nuits-pièges (longueur totale
des lignes de pièges: 6000 mètres). En retirant les juvéniles (usure dentaire nulle,
Keller, 1974; Palomo ef al., 1981) dont la croissance n’est pas terminée, le nombre
d’animaux étudiés est de 101, soit 33 + 20 + 2 Mus musculus domesticus et 28 + 12 +
6 Mus spretus (stations 1 + 2 + 3). Les deux espèces ont été identifiées en fonction du
nombre de plis palatins (avant la préparation ostéologique du crâne): Mus spretus
possède six rangées de plis palatins alors que Mus musculus domesticus en possède
sept (Darviche, 1978; Orsini, 1982).

MESURES

Les animaux ont été mesurés en utilisant un pied à coulisse au 1/10 mm pour
les mensurations corporelles, au 1/100 mm pour les mensurations crâniennes et mandi-
bulaires, et une loupe binoculaire munie d’un micromètre au 1/100 mm pour les men-
surations dentaires.

Les mensurations corporelles ont été prises sur l’animal maintenu à plat sur le
ventre après sa capture: longueur de la tête et du corps (T+C), longueur de la queue
(Q), longueur du pied postérieur (Pp), longueur de l’oreille (Or) et diamètre de la queue
(DQ); le rapport Q / T+C a été calculé.

Vingt-six variables crâniennes et dentaires inspirées des travaux de Sans-Coma
et al. (1979), Thorpe et al. (1982), Darviche & Orsini (1982), Davis (1983), Palomo et
al. (1983) et Lyalyukhina er al. (1991) ont été relevées: longueur du crâne (Lcr),
longueur et largeur du nasal (Ln, In), largeur bizygomatique (1bz), largeur interorbitaire
(io), largeur bisquamosale (Isq), largeur du foramen occipital (focc), longueur et
largeur de la bulle tympanique (Lbul, Ibul), hauteur postérieure du crâne (hcr),
longueur du foramen palatin (fopal), largeur du palais (Ipal), longueur incisivo-palatine
(de l’avant des incisives à l’arrière du palais) (Lipa), largeur du ramus dorsal de
l’arcade zygomatique (Iraz), largeur de l’arcade zygomatique (laz), coefficient zygo-
matique (coz = lraz/laz), longueur de l’orbite (Lorb), largeur du crâne (ler), longueur
du diastème supérieur (diasup), longueur de la rangée molaire supérieure (rms),
longueur de la rangée molaire inférieure (rmi), hauteur de la mandibule (hm), longueur
du diastème inférieur (diainf), longueur de la mandibule (Md), longueur et largeur de
la première molaire inférieure (Lm1, Iml).

ANALYSE STATISTIQUE

Outre des statistiques descriptives et tests de Student, l’identification des deux
espèces a été réalisée au moyen d’une analyse discriminante pas à pas qui sélectionne
successivement la (puis les) variable(s) la (les) plus discriminante(s), complétée par
une analyse discriminante avec validation croisée qui détermine l’affectation de

414 N. KHAMMES, S. LEK & S. AULAGNIER

chaque individu à partir de l’apprentissage réalisé sur tous les autres. Les calculs ont
été effectués avec le logiciel SPSS Professional Statistics 6.1 (Norusis, 1994).

RÉSULTATS

BIOMÉTRIE CORPORELLE

Toutes les variables corporelles, à l’exception de la longueur du pied postérieur,
différencient très significativement les deux espèces (Tableau 1), tout en restant faible-
ment corrélées. La Souris d’Afrique du Nord a une queue plus courte que la longueur
tête plus corps contrairement à la Souris domestique qui a une queue plus longue que
la longueur tête plus corps. Le rapport Q/T+C, qui permet classiquement de caractéri-
ser chacune des deux espèces varie de 0,48 à 0,99 chez Mus spretus et de 0,93 à 1,31
chez Mus musculus domesticus. De fait, les mesures de la queue permettent objective-
ment de déterminer la quasi totalité des individus. Toutefois la première analyse

TABLEAU 1. Mensurations des deux espèces de souris, Mus musculus domesticus et Mus spretus
en Kabylie du Djurdjura (Algérie) (n: effectif de l’échantillon; min: minimum; moy: moyenne;
max: maximum; Et: écart-type; t: test de Student; ddl: degrés de liberté; s: seuil de signification
du test de Student).

Mus musculus domesticus Mus spretus
N min moy Max Et n min moy max Et t ddl S
T+C 55 50,0 65,81 86,0 10,93 46 60,0 76,77 98,0 11,43 612,54 99 0,000
Q 55 52,5 71,20 88,5 11,91 46 44,6 57,83 70,5 7,46 584,60 99 0,000
Pp SSIS OMIS SSP 20/10 751508 46 14,8 16,38 19,6 1,56 2,97 99 0,098
Or SMES 135 MR 1S:0 077 46) 10/0) 125768 M1555) 153 6,06 99 0,000
DQ Sp PS 37, 0,40 AGIO 108 1252024 8,42 99 0,000
OVC RSS: TOS 7153 0,08 AG) O04 VOM 1059) ON 7,11 99 0,000
Ler SoS Ole LOS 2190") 1,74 387 17,38" 20387 22.907122 9,36 74 0,000
Ln 305257627 27752102 9:00) 20,74 45 5,85 7,67 92 091 2,93 93 0,000
In SO) 222295 2588) 027, 45 1,54 2,57 3,60 0,48 1,51 93 0,074
lbz 30:59 2939 LOOK. 0:53 41 849 9,47 10,10 0,40 1,1572771020523
io 44 299 3,76 4,89 0,33 46), 2.9977 3,79 74:96) 0/42 1573) 88) 03528
Isq 38 7,42 859 943 0,53 38) 8,297 7951172958357 10:44 1,22 74 0,360
focc 40 3,63 4,54 5,71 0,60 38 348 447 5,58 0,63 1,49 76 0,063
Lbul 382 25/63 98a 332) 053 29 2599 398 5,1 an 0:60 1,25 65 0,304
Ibul 33, 214" 3/07 4520550 2052 30,002 O74 OZ 1,03 65 0,879
her 38 6,33 7,46 834 0,48 33,657, ask) sa OS! 1530) A1M025SI
fopal 47 3,10 3,96 6,16 0,61 45 3,12 4,78 65 0,84 2,79 90 0,000
Ipal AGW 2:00) 7 2570) 379s) 032 46 2,21 2,80 3,96 0,34 2,28 90 0,000
Lipa 46 6,81 8,88 12,10 0,97 45) 18:01) 958115122048 3,90 89 0,000
lraz 47 022 041 0,84 0,11 46 0:27, 0/57. .0:3172032 11,98 91 0,000
laz 47 048 0,84 1,80 0,24 46 0,36 0,76 1,06 0,14 6,15 91 0,000
coz 47 0,33 0,49 0,4 0,07 46) 0:52) OS 41.067 098 8,65 91 0,000
Lorb SOS OS NC 27072 CIO SO 42 4,64 6,75 7,66 0,61 1:66 2779520215
ler 46 4,55 6,59 937 0,96 467 3:96 16583) 957021534 5,46 90 0,000
diasup 47 3,55 4,87 6,03 0,60 454.327 SI 6260000559, 1,09 90 0,667
rms 49 2,69 3,14 3,90 0,20 461 02:99 3518) 33455 10:08 2.601 7370072
rmi SON 252) e290) 3:03) 10:09 46) 2:055 2:90) 50720073 3,58 99 0,140
hm 33:43 472) 596) 10:66 A312 NA GS SSD nO 69 1,83, 92) 107162
diainf SO 2 ASS 0780572460, 20/45 46722257 STI SANS IT 0149 1.1979950372
Md 558 0501812 0 OI 46 8,85 10,68 12,2 0,77 3,33 97 01032
Lml SS IE 3114870108 46 0,88 1,36 1,68 0,17 1340 99 0,000

Iml SIMO!) 06) 1.20.4, 0:06 46 0,96 1,04 1,12 0,04 7,14 99 0,841

BIOMETRIE DISCRIMINANTE DES SOURIS D’ALGERIE 415

discriminante retient trois variables corporelles dans l’ordre suivant: diamètre de la
queue, rapport Q/T+C et longueur de l’oreille avec un résultat remarquable tant en
apprentissage qu’en validation croisée (Tableau 2). Avec le seul rapport Q/T+C, la
discrimination des deux espèces est très bonne, elle devient excellente en rajoutant le
diamètre de la queue (seul un individu est mal classé en apprentissage, contre trois en
validation croisée).

BIOMÉTRIE CRÂNIENNE

La majorité des variables crâniennes ne sont pas significativement différentes
d’une espèce à l’autre. Sont significativement différentes: la longueur du crâne, la
longueur du nasal, la longueur du foramen palatin, la largeur du palais, la longueur
incisivo-palatine, la largeur du ramus dorsal de l’arcade zygomatique, la largeur de
l’arcade zygomatique (et le coefficient zygomatique), la largeur du crâne, la longueur
de la mandibule et la longueur de la première molaire inférieure.

Les variables crâniennes apparaissent relativement corrélées entre elles, à
l’exception des rangées molaires supérieure (rms) et inférieure (rmi), des largeurs
interobitaire (io) et bizygomatique (bz), de la longueur de la première molaire
inférieure (Lm1), seulement corrélée à la largeur de cette même dent (bm1, r=0,59).
Les variables les plus corrélées sont la longueur du crâne (Lcr) et la longueur incisivo-
palatine (Lipa, r=0,66), et surtout la largeur du ramus dorsal de l’arcade zygomatique
(Iraz) et le coefficient zygomatique (coz, r= 0,78).

L'analyse discriminante pas-à-pas (90,4% de bon classement en apprentissage
et en validation croisée) retient une seule variable: le coefficient zygomatique (coz).
L’exclusion du coefficient zygomatique de l’analyse conduit à un modèle moins
performant qui repose encore sur les mesures de l’arcade zygomatique avec pour
complément la largeur bizygomatique et la longueur incisivo-palatine (Tableau 2). Fort
heureusement l’arcade zygomatique est relativement robuste et pourra être mesurée sur
les crânes issus de pelotes de réjection de rapaces, notamment de Chouette effraie (7yto
alba), de Hibou moyen-duc (Asio otus) ou de Chouette hulotte (Strix aluco).

Les deux analyses discriminantes effectuées sur les mensurations de la mandi-
bule ne peuvent être retenues, les pourcentages de classement, tant en apprentissage
qu’en validation croisée, étant trop faibles (Tableau 2). Il apparaît donc impossible
d'identifier avec une incertitude raisonnable des mandibules isolées édentées, telles
qu’on les trouve souvent dans les pelotes de réjection de rapaces ou bien dans les fè-
ces de carnivores.

DISCUSSION

Il ressort de ce travail que la biométrie corporelle apparaît plus performante que
la biométrie cränienne pour discriminer les deux espèces de souris présentes en
Kabylie du Djurdjura. Conformément à la description originale de Mus spretus, cette
souris possède une queue plus courte que Mus musculus domesticus (57,83 + 7,46 mm
vs 71,20 + 11,91 mm), le rapport de la longueur de la queue sur la longueur tête plus
corps permet de séparer nettement les deux espèces. Compris entre 0,77 et 0,99 chez
la souris à queue courte, il est de 0,93 à 1,31 chez la souris domestique. De plus la
queue de Mus spretus est toujours plus fine que celle de M. m. domesticus. La combi-

416 N. KHAMMES, S. LEK & S. AULAGNIER

TABLEAU 2. Identification des deux espèces de souris en Kabylie du Djurdjura: résultats des ana-
lyses discriminantes sur les variables de morphologie externe et crânienne (pourcentages de clas-
sement en apprentissage (% Appr.) et en validation croisée (% Valid.), et principales fonctions
discriminantes, A: Mus musculus domesticus, B: Mus spretus; si A est supérieur à B l’animal est
Mus musculus domesticus, si B est supérieur à A l’animal est Mus spretus).

Variables % Appr. % Valid. Fonctions discriminantes

Biométrie corporelle

DQ, Q/T+C, Or 99,0 93,0 A = 16,220 DQ + 112,007 Q / T+ C + 11,864 Or — 164,693
B = 11,641 DQ + 83,084 Q / T + C + 10,667 Or — 111,387
Q/T+C 95,0 95,0 A = 78,781 Q/T+C - 43,686
B = 54,119 Q/T+C - 20,982
Q/T+C, DQ 99,0 97,0 A = 15,702 DQ + 83,633 Q / T+C -68,249

B = 11,175 DQ + 57.572 Q / T+C — 33.423

Biometrie cränienne
COZ 90,4 90,4 A = 48,247 COZ -12,559
B = 72,904 COZ - 27,787

lbz, lraz, laz, Lipa 92,2 88,3
Lml, Iml 71,3 70,3
Md, hm 62,0 60,9

DQ: diamètre de la queue; Q / T+C: rapport de la longueur de la queue (Q) sur la longueur de la
tête et du corps (T+C); Or: longueur de l’oreille; COZ (coefficient zygomatique): rapport de la
largeur du ramus dorsal de l’arcade zygomatique (Iraz) sur la largeur de l’arcade zygomatique
(laz); Ibz: largeur bizygomatique; Lipa: longueur incisivo-palatine (de l’avant des incisives à
l’arrière du palais); Lm1-lm1: longueur et largeur de la première molaire inférieure; Md: lon-
gueur de la mandibule; hm: hauteur de la mandibule.

naison des deux variables fournit les meilleures fonctions dicriminantes. Ces résultats
sont conformes aux données publiées sur les spécimens d’Europe (Orsini et al., 1982;
Darviche & Orsini 1982; Palomo et al., 1981), avec toutefois un intervalle différent
pour le rapport de la longueur de la queue sur la longueur téte plus corps (0,6-0,8 pour
Mus spretus, 0,75-1,05 pour Mus musculus domesticus). Cette différence est liée à la
longueur tête plus corps, significativement plus faible chez les souris domestiques
d’ Algérie (65,95 + 11,54 mm contre 76,52 + 11,72 mm pour les souris à queue courte),
alors qu’en Europe Orsini et al. (2001) rapportent des longueurs très proches (75,44 +
1,83 mm et 75,86 + 2,00 mm respectivement). Orsini et al. (2001) évoquent par contre
la longueur plus réduite des membres de Mus spretus, ce qui renforce l’impression de
petite taille chez cette espèce. En Algérie la différence n’est pas significative.

Sur le crâne, le coefficient zygomatique, rapport de la largeur du ramus dorsal
de l’arcade zygomatique sur la largeur de l’arcade zygomatique, est la meilleure
variable pour la discrimination des deux espèces sympatrique (0,33 à 0,64 chez Mus
musculus domesticus, 0,60 à 1,06 chez Mus spretus). Pour les populations européennes
il était déjà considéré comme discriminant par Darviche & Orsini (1982) et Orsini et
al. (1983), avec cependant une valeur seuil de 0,70 (0,79 +0,14 chez Mus spretus et
0,50 +0,08 chez Mus musculus domesticus (Orsini et al., 2001). Macholän (1996b) a

BIOMETRIE DISCRIMINANTE DES SOURIS D’ALGERIE 417

en outre relevé une hauteur du crâne plus élevée chez Mus musculus domesticus, alors
que cette mensuration n’est pas significativement différente chez les souris algériennes.
Enfin, la longueur et la largeur de la première molaire inférieure, qui permettent de
séparer les deux espèces de souris selon ce méme auteur, ne fournissent pas une
discrimination satisfaisante pour les spécimens algériens, en dépit de la longueur
significativement supérieure observée chez Mus spretus. Il reste que pour les mandi-
bules, le dessin de la surface d’usure de cette dent constitue un critère d’identification
supplémentaire, quoique parfois délicat (Orsini, 1979).

CONCLUSION

Hormis quelques légères différences pour certaines variables (longueur téte plus
corps notamment) qu’il faudra confirmer sur des échantillons en provenance d’autres
régions, les deux espèces de souris de Kabylie du Djurdjura peuvent être distinguées
sur la base des critères reconnus en Europe : rapport de la longueur de la queue et de
la longueur tête plus corps en association avec le diamètre de la queue pour la bio-
métrie corporelle et le coefficient zygomatique pour la biométrie crânienne. La fiabi-
lité des identifications permet d’envisager des études sur l’écologie et la biologie des
populations des deux espèces, qui semblent particulièrement inféodées aux activités
anthropiques en Afrique du Nord (Aulagnier & Thévenot, 1986; Khidas, 1993; Khidas
et al., 1999). L'identification des proies des rapaces s’en trouve également facilitée, les
différents caractères morphologiques proposés (synthèse dans Macholän, 1996b et
Orsini et al., 2001) n'étant pas toujours accessibles.

REMERCIEMENTS

Ce travail a été réalisé dans le cadre de l’accord — programme franco-algérien
99MDU423.

BIBLIOGRAPHIE

AULAGNIER, S. & THÉVENOT, M. 1986. Catalogue des Mammifères sauvages du Maroc. Travaux
de l’Institut Scientifique, Rabat 41: 1-163.

BOURSOT, P., JACQUART, T., BONHOMME, F., BRITTON-DAVIDIAN, J. & THALER, L. 1985.
Différenciation géographique du génome mitochondrial chez Mus spretus Lataste.
Comptes Rendus de l’Académie des Sciences de Paris, Série IMI, 301: 161-166.

BRITTON, J., PASTEUR, N. & THALER, L. 1976. Les souris du midi de la France: caractérisation
génétique de deux groupes de populations sympatriques. Comptes Rendus de l’Académie
des Sciences de Paris, Série III, 283: 515-518.

BRITTON, J. & THALER, L. 1978. Evidence for the presence of two sympatric species of mice (ge-
nus Mus L.) in southern France based on biochemical genetics. Biochemical Genetics 16:
214-225.

CABRERA, A. 1914. Fauna ibérica. Mamiferos. Museo nacional de Ciencias naturales, Madrid,
441 pp.

CABRERA, A. 1932. Los Mamiferos de Marruecos. Trabajos del Museo nacional de Ciencias
naturales, Seria Zoologia 57: 1-361.

CASSAING, J. 1982. Les populations sauvages de souris du Midi de la France (Mus domesticus et
Mus spretus) : Approche étho-écologique et conséquences évolutives. Thèse 3€ Cycle
(Evolution et Biosystématique), Université des Sciences et Techniques du Languedoc,
Montpellier, 158 pp.

418 N. KHAMMES, S. LEK & S. AULAGNIER

CASSAING, J. 1984. Interactions intra et inter spécifiques chez les souris sauvages du midi de la
France, Mus musculus domesticus et Mus spretus: conséquences sur la compétition entre
les deux espèces. Biology of Behaviour 9: 281-293.

CASSAING, J. & CROSET, H. 1985. Organisation spatiale, compétition et dynamique des popu-
lations sauvages de souris (Mus spretus Lataste et Mus musculus domesticus Rutty) du
Midi de la France. Zeitschrift für Säugetierkunde 50: 271-284.

Davis, S.J.M. 1983. Morphometric variation of populations of house mice Mus domesticus in
Britain and Faroe. Journal of Zoology, London 199: 521-534.

DARVICHE, D. 1978. Approche morphologique et biométrique de la biosystématique à la lumière
de la génétique biochimique des populations. Application aux genres Mus et Apodemus
(Mammalia, Rodentia). Thèse 3€ Cycle (Evolution et Biosystématique), Université des
Sciences et Techniques du Languedoc, Montpellier, 191 pp.

DARVICHE, D. & ORSINI, P. 1982. Critères de différenciation morphologique et biométrique de
deux espèces de souris sympatriques: Mus spretus et Mus musculus domesticus.
Mammalia 46: 205-217.

ELLERMAN, J.R. & MORRISON-SCOTT, T.C.S. 1951. Checklist of Palearctic and Indian Mammals
1758 to 1946. British Museum (Natural History), London, 810 pp.

GERASIMOV, S., NIKOLOV, H., MIHAILOVA, V., AUFFRAY, J.C. & BONHOMME, F. 1990. Morpho-
metric stepwise discrimant analysis of the five genetically determined European taxa of
the genus Mus. Biological Journal of the Linnean Society 41: 47-64.

KELLER, A. 1974. Détermination de l’âge de Mus musculus Linné par l’usure de la dentition.
Revue suisse de Zoologie 81: 839-844.

KHIDAS, K. 1993. Distribution des rongeurs en Kabylie du Djurdjura (Algérie). Mammalia 57:
207-212.

KHIDAS, K., KHAMMES, N. & KHELLOUFI, S. 1999. Répartition spatiale et sélection de l’habitat
chez le mulot sylvestre (Apodemus sylvaticus Linnaeus, 1758) et la souris sauvage (Mus
spretus Lataste, 1883) en Kabylie du Djurdjura (Algérie). Sciences & Technologie 12:
59-64.

LATASTE, F. 1883. Note sur les souris d’ Algérie et description d’une espèce nouvelle (Mus spre-
tus). Actes de la Société Linnéenne de Bordeaux 37: 13-35.

LATASTE, F. 1887. Catalogue critique des mammifères apélagiques de Tunisie. In: Exploration
scientifique de la Tunisie. Imprimerie nationale, Paris, XV + 42 pp.

LOCHE, V. 1846. Sciences physiques, Zoologie. Histoire naturelle des mammifères. In:
Exploration scientifique de l’Algérie pendant les années 1840, 1841, 1842. Imprimerie
royale, Paris, 123 pp.

LYALYYUKHINA, S., KOTENKOVA, E., WALKOWA, W. & ADAMCZYK, K. 1991. Comparison of cran-
iological parameters in Mus musculus musculus Linnaeus, 1758 and Mus hortulanus
Nordmann, 1840. Acta Theriologica 36: 95-107.

MACHOLAN, M. 1996a. Morphometric analysis of European house mice. Acta Theriologica 41:
255-275:

MACHOLAN, M. 1996b. Key to European house mice (Mus). Folia Zoologica, 45: 209-217.

MARSHALL, J. T. & SAGE, R.D. 1981. Taxonomy of the house mouse. Symposium of the
Zoological Society of London 47: 15-25.

MILLER, G.S. 1912. Catalogue of the mammals of the Western Europe (Europe exclusive of
Russia) in the collection of the British Museum. British Museum (Natural History),
London, 1019 pp.

Norusıs, M.J. 1994. SPSS: SPSS Professional Statistics 6. S.P.S.S. Inc., Chicago, 385 pp.

ORSINI, P. 1979. Notes sur les souris de Provence. Annales de la Société de Sciences naturelles
et d’Archéologie de Toulon et du Var 31: 168-173.

ORSINI, P. 1982. Facteurs régissant la répartition des souris en Europe: intérét du modèle souris
pour une approche des processus évolutifs. Thèse 3€ Cycle (Evolution et Biosysté-
matique), Université des Sciences et Techniques du Languedoc, Montpellier, 134 pp.

BIOMÉTRIE DISCRIMINANTE DES SOURIS D’ALGÉRIE 419

ORSINI, P., CASSAING, J., DUPLANTIER, J.M. & CROSET, H. 1982. Premières données sur l’écologie
des populations naturelles de souris, Mus spretus Lataste et Mus musculus domesticus
Rutty dans le Midi de la France. Terre et Vie 36: 321-335.

ORSINI, P., FAUGIER, C. & BUTET, A. 2001. Identification des Insectivores et Rongeurs de France:
les espèces jumelles de souris, Mus musculus domesticus et Mus spretus. Arvicola 13:
9-11.

PALOMO, L.J., ESPANA, M., LOPEZ-FUSTER, M.-J., GOZALBEZ, J. & SANS-COMA, V. 1981. Sobre la
variabilidad fenética y morfométrica de Mus spretus Lataste, 1883 en la Peninsula
Ibérica. Miscellania Zoologica 7: 171-192.

SANS-COMA, V., LOPEZ-FUSTER, M. J. & GOSALBEZ, J. 1979. Über die Hausmans, Mus musculus
Linné; 1758, auf der Insel Meda Crossa Katalonien, Spanien. Säugetierkundliche
Mitteilungen, 27: 107-113.

SAINT GIRONS, M.C. & THOUY, P. 1978. Fluctuations dans les populations de souris Mus spretus
Lataste 1883, en région méditerranéenne. Bulletin d’Ecologie 9: 211-218.

SCHWARZ, E. & SCHWARZ, H.K. 1943. The wild and commensal stocks of the house mouse, Mus
musculus Linnaeus. Journal of Mammalogy 24: 59-72.

THORPE, R.S., CORTI, M. & CAPANNA, E. 1982. Morphometric divergence of Robertsonian
populations/species of Mus: A multivariate analysis of size and shape. Experientia 38:
920-923.

TRIPATHI, R.S., MATHUR, M., JAIN, A.P. & PATEL, N. 1994. Relative efficacy of glue and other
traps for commensal rodent management. Annals of arid Zone 33: 143-145.

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REVUE SUISSE DE ZOOLOGIE 113 (2): 421-436; juin 2006

A review of Capoeta tinca, with descriptions of two new species
from Turkey (Teleostei: Cyprinidae)

Davut TURAN!, Maurice KOTTELAT?, F. Güler EKMEKCP &

H. Oguz IMAMOGLU!

1 Karadeniz Technical University, Faculty of Fisheries and Aquatic Sciences,
53100 Rize, Turkey. E-mail: davutturan61 @hotmail.com

2 Corresponding author, Route da la Baroche 12, CH-2952 Cornol, Switzerland
(permanent address) and Raffles Museum of Biodiversity Research,
National University of Singapore, Kent Ridge, Singapore 119260.
E-mail: mkottelat @ dplanet.ch

3 Department of Biology, Faculty of Sciences, Hacettepe University, Beytepe Campus,
06532 Ankara, Turkey. E-mail: gulere@hacettepe.edu.tr

A review of Capoeta tinca, with descriptions of two new species from
Turkey (Teleostei: Cyprinidae). - Fishes previously referred to Capoeta
tinca in Turkey and Georgia belong to three species: C. tinca in rivers
draining to the Marmara Sea, C. baliki, new species, in rivers draining to the
southwestern Black Sea, and C. banarescui, new species, in the Coruh River
drainage. Capoeta banarescui is distinguished by missing sexual
dimorphism in the mouth shape (present in the two other species) and fewer
and larger scales. Capoeta baliki is distinguished by its more slender body
and caudal peduncle, and blunter head.

Keywords: New species - Cyprinidae - Capoeta - Coruh River - Sakarya
River - Kızılırmak River - Anatolia.

INTRODUCTION

Heckel (1843) described Scaphiodon tinca from "Brussa in Natolien", the
present city of Bursa, in the Nilüfer drainage, a stream flowing to the Sea of Marmara
in Turkey. Giinther (1868), Steindachner (1897), Berg (1949), and Kosswig & Battalgil
(1943) reported this species as Varichorhinus tinca from Bursa, Ankara, Esksehir,
Sakarya and Trabzon. Karaman (1969) revised the genus Varicorhinus and placed V.
tinca in the genus Capoeta. Since, several authors have reported C. tinca from central
and northern Anatolia (e.g., Kuru, 1975; Balık, 1979; Erk’akan, 1981; Kutrup, 1994).
Although these authors examined different populations, all recorded somewhat similar
lateral line scale and and gill raker counts.

Banarescu & Herzig-Straschil (1999) redescribed C. tinca and mentioned
considerable differences between specimens from Coruh drainage (northeastern
Anatolia) and those from western and central Anatolia. We have compared the different
populations which have been referred to C. tinca and conclude that they are in fact
three species. The Coruh and Sakarya populations are described as two new species, C.
banarescui and C. baliki, respectively.

Manuscript accepted 28.11.2005

422 D. TURAN ET AL.

MATERIAL AND METHODS

Fish were caught with pulsed DC electrofishing equipment. Material is depo-
sited in: ESFM, Museum of the Faculty of Fisheries, Ege University, Izmir; FFR,
Zoology Museum of the Faculty of Fisheries, Karadeniz Technical University, Rize;
and CMK, the collection of the second author. Additional material is preserved in the
collection of the third author. Measurements were taken with digital calipers (0.1 mm
accuracy). Counts and measurements follow Hubbs & Lagler (1947) except as follows.
Head width,: distance between anterior margin of eye; head width): distance between
posterior margin of eyes; snout width: measured at level of nostril. Lateral line scale
count includes scales on the base of the caudal fin. Vertebae count includes the four
Weberian vertebrae and the hypural complex. The last two branched anal and dorsal-
fin rays articulating on the same pterygiophore are counted as one.

Capoeta tinca (Heckel, 1843) Fig. 1

Capoeta tinca Heckel, 1843: 1021 (type locality: "Brussa in Natolien"; lectotype: Natur-
historisches Museum Wien 55931:1, designated by Banarescu & Herzig-Straschil, in
Banarescu, 1999: 413).

MATERIAL EXAMINED: FFR 717, 2, 145-157 mm SL; Turkey: Bursa: Nilifer River,
40°15'N 28°55'E; D. Turan & S. G. Kirankaya, 15 September 2004. - FFR 718, 23, 116-201 mm;
CMK 18538, 10, 113-179 mm SL; Turkey: Balikesir: Koca River, Manyas, 40°05'N 28°02'E; D.
Turan & S. G. Kirankaya, 14 September 2004.

DIAGNOSIS: Capoeta tinca is distinguished from the other species of the genus
by the combination of the following characters: two pairs of barbels; snout rounded:
69-80 lateral line scales; 14-17 scales rows between lateral line and dorsal-fin origin,
12-14 between lateral line and anal-fin origin; 19-23 gill rakers on the first gill arch;
lower jaw slightly arched in males, straight in females; head length 23.3-26.7% SL;
depth of caudal peduncle 10.8-13.4% SL; head width at posterior margin of eye 49.3-
56.5% HL; snout depth at nostril 30.0-41.1% HL; length of anterior barbel 8.1-14.1%
HL and posterior barbel 11.5-19.3% HL.

DESCRIPTION: See Figure 1 for general appearance and Tables 1-2 for morpho-
metric and meristic data. Dorsal head profile convex. Snout rounded, blunt, triangular
in ventral view, depth slightly smaller than width at nostrils. Mouth wide, shape
sexually dimorphic, arched in male, straight in female (Fig. 2). Rostral fold well
developed, partly hiding upper lip. Upper and lower lips adnate to jaws, lower jaw
covered with horny sheath. No tubercles on head. Anterior barbel not reaching corner
of mouth. Posterior barbel reaching about middle of eye. Predorsal profile of body
convex. Body high and weakly compressed.

Dorsal fin with 3 or 4 simple and 8 branched rays, outer margin slightly emar-
ginate, origin slightly in front of vertical through pelvic-fin origin, last simple ray
moderately ossified, proximal two thirds rigid, and with 24-28 serrae on posterior
margin (Fig. 3a). Pectoral fin with 18-20 branched rays. Pelvic fin with 1 simple and
8-9 branched rays. Anal fin with 3 simple and 5 branched rays, outer margin convex.
Caudal fin long and deeply forked. Gill rakers, 6-7 + 1 + 12-15 = 19-23 on outer side
of first arch, number increasing with size (19, in 6 specimens about 125 mm SL; 20, in
8 specimens about 136 mm SL; 21, in 4 specimens about 145 mm SL; 22, in 5 spe-

REVIEW OF CAPOETA TINCA 423

Fic. 1
Capoeta tinca, FFR 718, 129 mm SL; Turkey: Koca River.

FIG. 2
Capoeta tinca, FFR 718, female, 129 mm SL (left) and male, 136 mm SL (rigth).

cimens about 160 mm SL; 23, in 2 specimens about 200 mm SL). 69-80 lateral line
scales, 14-17 between dorsal-fin origin and lateral line and 9-11 between anal-fin origin
and lateral line. 44-46 (modally 45) total vertebrae.

SEXUAL DIMORPHISM: Males collected in September have no tubercles on side
and tip of snout, and on cheeks. Mouth slightly arched in male, straight in female.

COLORATION: Live and formalin preserved specimens dark brown on back and
flanks, yellowish white on belly. Each scale margined by a band of black pigments,
forming a regular reticulated pattern. Dorsal and caudal fins grey; pectoral, pelvic and
anal fins whitish.

D. TURAN ET AL.

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D. TURAN ET AL.

Last simple dorsal-fin ray of: a, Capoeta tinca, FFR 718, 139 mm SL, female; b, C. banarescui,
FFR 712, 144 mm SL, female; and c, C. baliki, FFR 713, 148 mm SL, female.

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FIG. 4

Distribution of Capoeta tinca, C. banarescui, C. baliki and C. antalyensis in Turkey.

DISTRIBUTION: Capoeta tinca is known from the rivers draining to the southern
shore of the Marmara Sea (Fig. 4).

HABITAT AND BIOLOGY: Capoeta tinca inhabits swift flowing water, with cobbles
and pebbles bottom. Alburnoides bipunctatus, Barbus cf. oligolepis, Barbus sp.
Chalcalburnus chalcoides, Rhodeus sericeus, Squalius cf. orientalis, and Vimba vimba

have been collected together with C. tinca.

REVIEW OF CAPOETA TINCA 427

Capoeta banarescui sp. n. Fig. 5

Capoeta tinca (non Heckel, 1843): Derjugin, 1899 : 155 (in part; Coruh River, Georgia); Berg,
1914: 0168 (Olty-tchai [Oltu Cay1], Coruh drainage); Karaman, 1969: 37 (in part; Coruh
drainage: Tortum Reservoir).

Varicorhinus tinca (non Heckel, 1843): Berg, 1914: 554 (Coruh and Rion drainages), 1949: 684
(in part; Coruh and Rion drainage, Georgia); Elanidze, 1983: 117 (Coruh, Georgia).

HoLoTYPE: ESFM-PISI/2004-072, 177 mm SL; Turkey: Artvin: Tortum District: Coruh
drainage, stream Tortum, 100 km north of Erzurum; 40°34'N 41°36'E; D. Turan, F. G. Ekmekgi,
H. O. Imamoglu, O. Serdar & S. G. Kirankaya, 19 July 2004.

Paratypes. ESFM-PISI/2004-073, 4, 166-201 mm SL; FFR 712, 16, 85-232 mm SL;
CMK 18474, 5, 135-193 mm SL; same data as holotype. - FFR 711, 9, 163-231 mm SL; CMK
18540, 9, 121-193 mm SL; Turkey: Artvin: Coruh drainage, Bulamk stream, Savsat, 30 km east
of Artvin, 41°34'N 42°14'E; D. Turan, F. G. Ekmekgi, H. O. Imamoglu, O. Serdar & S. G.
Kirankaya, 19 June 2004. - FFR 720, 3, 92-125 mm SL; CMK 18549, 1, 145 mm SL; Turkey:
Cavuslu, Borçka, 41°21'N 41°42'E; D. Turan, 13 October 2004.

DIAGNOSIS: Capoeta banarescui is distinguished from the other species of the
genus by the combination of the following characters: two pairs of barbels; snout
pointed; 64-77 lateral line scales; 12-14 rows of scales between lateral line and dorsal-
fin origin, 9-11 between lateral line and anal-fin origin; 12-16 gill rakers on first gill
arch; shape of lower jaw not sexually dimorphic; head length 22.2-25.8% SL; depth of
caudal peduncle 9.8-11.7% SL; head width at posterior margin of eye 49.4-58.2% HL;
snout depth at nostril 29.7-35.1%HL; length of anterior barbel 12.4-20.8% HL and
posterior barbel 18.4-28.8% HL.

DESCRIPTION: See Figure 5 for general appearance and Tables 1-2 for morpho-
metric and meristic data. Dorsal head profile convex. Snout pointed, rounded in ventral
view, depth smaller than width at nostrils. Mouth large and slightly arched, shape not
sexually dimorphic (Fig. 6). Rostral fold well developed, partly hiding upper lip.
Middle part of upper lip thick, thinner at corners. Upper and lower lips adnate to jaws,
lower jaw covered with horny sheath. Tubercles on lower half of body smaller, denser
and larger on lower half of caudal peduncle. A row of large tubercles along branched
anal-fin rays. Anterior barbel reaching to below anterior margin of eye, beyond corner
of mouth. Posterior barbel reaching beyond middle of eye. Predorsal profile of body
convex. Body slightly compressed laterally.

Dorsal fin with 3 or 4 simple and 7-9 (modally 8) branched rays, outer margin
straight or slightly emarginate, origin markedly in front of vertical through pelvic-fin
origin, last simple dorsal ray weakly ossified, flexible and with 12-20 serrae on posterior
margin (Fig. 3b). Pectoral fin with 17-19 branched rays. Pelvic fin with 1 simple and
9-10 branched rays. Anal fin with 3 simple and 5 branched rays, outer margin convex.
Caudal fin long and deeply forked. Gill rakers large and rounded, 3-5 + 1 + 8-10 = 12-
16 on outer side of first arch, number increasing with size (12, in 2 specimens about 80
mm SL; 13, in 2 specimens about 100 mm SL; 14, in 4 specimens 94-135 mm SL; 15,
in 11 specimens 125-154 mm SL; 16, in 6 specimens 143-215 mm SL, including holo-
type). 64-77 lateral line scales, 12-14 scale rows between dorsal-fin origin and lateral
line and 8-9 between anal-fin origin and lateral line. 45-46 (modally 45) total vertebrae.

SEXUAL DIMORPHISM: Males collected in July have well developed tubercles on
side and tip of snout, and on cheeks.

428 D. TURAN ET AL.

FIG. 5

Capoeta banarescui, holotype, ESFM-PISI/2004-072, 177 mm SL; Turkey: Coruh drainage:
Tortum.

FIG. 6
Capoeta banarescui, FFR 712, female, 192 mm SL (left) and male, 178 mm SL (right).

COLORATION: Formalin preserved specimens dark brown on back, flank brown,
belly yellowish. Each scale margined by a band of black pigments, forming a regular
reticulated pattern. Dorsal, caudal and pectoral fins brown; pelvic and anal fins whitish.
In life: back and upper flank brownish grey, belly whitish to yellow.

DISTRIBUTION: Capoeta banarescui is presently known only from the Coruh
(Tchorok) River drainage (Fig. 4). The Coruh originates from the Kagkar Mountains in
Turkey, but its lowermost course is in Georgia and it flows to the Black Sea at Batumi.

HABITAT AND BIOLOGY: Capoeta banarescui is known from swift flowing water,
with cobbles and pebbles bottom. In June 2004, the temperature was 15-16°C,
dissolved oxygen 8.23 mg rl, pH 7.19, and conductivity 125 mS. Alburnoides bipunc-
tatus, Barbus tauricus and Oxynoemacheilus sp. were collected together with C.

REVIEW OF CAPOETA TINCA 429

banarescui. Solak (1982) and Yıldırım & Aras (2000) report that C. banarescui spawns
between May and July in Coruh. Males collected in May and June have tubercles on
the snout and the body. Capoeta banarescui feeds mainly on phytoplankton, zoo-
plankton, some invertebrate, algae and other aquatic plants. In stream Oltu, C. ba-
narescui reaches 400 mm (fork length), 838 g and 12 years; it reaches sexual maturity
at 2-3 years (Yıldırım & Aras, 2000).

ETYMOLOGY: The new species is named for Petru Banarescu, in appreciation for
his contributions to the knowledge of the Turkish fish fauna.

Capoeta baliki sp. n. Fig. 7
Varicorhinus tinca (non Heckel, 1843): Kosswig & Battalgil, 1942: 56 (Ankara).
Capoeta tinca (non Heckel, 1843): Erk’akan, 1981: 144 (Sakarya River).

HOLOTYPE: ESFM-PISI/2004-74, 202 mm SL; Turkey: Ankara: Sakarya River: Kızılca-
hamam Stream, K>z>cahamam, 60 km west of Ankara, 40°29'N 32°39'E; D. Turan & M. Turan,
15 April 2004.

PARATYPES: ESFM-PISI/2004-75, 4, 140-190 mm SL; FFR 713, 5, 121-219 mm SL;
CMK 18541, 10, 128-188 mm SL; same data as holotype. - FFR 714, 5, 151-209 mm SL;
Turkey: Ankara: Sakarya River, Ova Stream, Kazan, 50 km west of Ankara, 40°11'N 32°39'E;
D. Turan & M. Turan, 15 April 2004. - FFR 715, 5, 121-183 mm SL; same data, 16 June 2004.
- FFR 716, 10, 168-217 mm SL; Turkey: Sivas: Kızılırmak River, Delice Stream; F. G. Ekmekei
& S. G. Kirankaya, 22 November 2002.

DIAGNOSIS: Capoeta baliki is distinguished from the other species of the genus
by the combination of the following characters: two pairs of barbels; snout bluntly
rounded; 72-86 lateral line scales; 14-17 scales rows between lateral line and dorsal-
fin origin, 10-11 (modally 10) between lateral line and anal-fin origin; 16-22 gill rakers
on the first gill arch; lower jaw slightly arched in males, straight in females; head
length 21.9-24.8% SL; depth of caudal peduncle 9.5-12.3% SL; head width at posterior
margin of eye 55.6-63.5% HL; snout depth at nostril 33.1-41.6% HL; length of anterior
barbel 9.8-18.7% HL and posterior barbel 14.7-25.5% HL.

DESCRIPTION: See Figure 7 for general appearance and Tables 1-2 for morpho-
metric and meristic data. Dorsal body profile straight. Snout bluntly rounded, rounded
in ventral view, depth slightly smaller than width at nostrils. Mouth wide, shape
sexually dimorphic, moderately arched in male, straight in female (Fig. 8). Rostral fold
weakly developed, partly hiding upper lip. Upper and lower lips adnate to jaws, lower
jaw covered with horny sheath. In males collected in April, tubercles on side and tip of
snout. Anterior barbel reaching base of posterior barbel. Posterior barbel reaching to
below anterior margin of eye. Predorsal profile of body only slightly convex. Body not
compressed laterally.

Dorsal fin with 3 or 4 simple and 8-9 (modally 8) branched rays, outer margin
sligthly emarginate, origin in front of vertical through pelvic-fin origin, last simple
dorsal ray weakly ossified, rigid on about two thirds of its length and with 17-23 serrae
on posterior margin (Fig. 3c). Pectoral fin with 17-20 branched rays. Pelvic fin with 1
simple and 9-10 branched rays. Anal fin with 3 simple and 5 branched rays, outer
margin convex. Caudal fin long and deeply forked. Gill rakers 5-7 + 1 + 10-14 = 16-
22 on outer side of first arch, number increasing with size (16, in 2 specimens about
116 mm SL; 17, in 5 specimens about 134 mm SL; 18, in 4 specimens about 155 mm

430 D. TURAN ET AL.

FIG. 7

Capoeta baliki, holotype, ESFM-PISI/2004-74, 202 mm SL; Turkey: Sakarya drainage: Kızılc-
ahamam.

FIG. 8
Capoeta baliki, FFR 714, female, 176 mm SL (left) and male, 164 mm SL (right).

SL; 19-22, in 24 specimens 173-213 mm SL, including holotype). 72-86 lateral line
scales, 14-17 between dorsal-fin origin and lateral line and 10-11 between anal-fin ori-
gin and lateral line. 43-44 (modally 44) total vertebrae.

REVIEW OF CAPOETA TINCA 431

FIG. 9
Capoeta antalyensis, FFR 719, 196 mm SL; Turkey: Gökdere stream.

Fic. 10
Capoeta antalyensis, FFR 719, female, 149 mm SL (left) and male, 181 mm SL (right).

SEXUAL DIMORPHISM: Males with breeding tubercles on snout. Mouth slightly
arched in male, straight in female.

COLORATION: Formalin preserved specimens dark brown on back, flank brown,
belly yellowish brown. Each scale margined by a band of black pigments, forming a
regular reticulated pattern. Dorsal, pectoral and caudal fins brown, pelvic and anal fins
light brown. In life: back and upper flank brown, belly yellow; caudal, pectoral and
pelvic fins dark brown, anal and dorsal fins pale brown.

432 D. TURAN ET AL.

DISTRIBUTION: Capoeta baliki is presently known from the Sakarya and
Kızılırmak river drainages, including in lakes and reservoirs (Fig. 4). The Sakarya
River originates from western central Anatolia and it enters the Black Sea at Sakarya.
Kızılırmak River flows from eastern central Anatolia and and enters the Black Sea at
Samsun.

HABITAT AND BIOLOGY: Capoeta baliki inhabits slowly flowing water, with
cobbles and pebbles substrate. It is also found in lakes and reservoirs, for example
Sariyar Reservoir on Sakarya River (Ekmekçi, 1996) and Gelingüllü Reservoir in
Kızılırmak basin (Ekmekçi & Kırankaya, 2004). Alburnoides bipunctatus, Barbus sp.
Capoeta sieboldi, Chalcalburnus chalcoides, Squalius cf. orientalis, Barbatula
angorae, Oxynoemacheilus cf. banarescui were collected together with C. baliki.
Males collected in June and July have tubercules on the snout. The spawning period in
central Anatolia is in May and June. Sexual maturity is reached 2 years for for males
and 3 years for females (Ekmekgi, 1996; Ekmekçi & Özeren, 2002). Capoeta baliki
reaches 428 mm (fork length), 1178 g and 10 years (Yılmaz, 1994).

ETYMOLOGY: Named for Süleyman Balik, for his contributions to the knowledge
of the Turkish fish fauna.

DISCUSSION

Species of Capoeta have (and still are) often placed in the genus Varicorhinus,
together with a number of very different cyprinids from Africa, South and Southeast
Asia. The type species of Varicorhinus is an African fish and the name is correctly used
only for African species. The genus Capoeta was last revised by Karaman (1969) who
recognized seven valid species: C. tinca, C. fusca, C. pestai, C. buhseri, C. capoeta, C.
trutta and C. barroisi. Banarescu (1999) redescribed some of the species recorded from
Turkey and noted problems with the identification of some populations. Capoeta ba-
narescui and C. baliki are immediately distinguished from most other species of
Capoeta in having two pairs of barbels, a character shared only with C. tinca and C.
antalyensis.

Varicorhinus tinca was described by Heckel (1843) from Bursa (40°15'N
28°55'E), in the Nilüfer drainage. The Nilüfer is a short coastal stream in northwestern
Anatolia, draining to the Sea of Marmara. Banarescu & Herzig-Straschil (in Banarescu,
1999) comment that the distribution of C. tinca is disjunct and includes most rivers
draining to the Sea of Marmara and the Black Sea basins between the Nilüfer and the
Sakarya drainages in Anatolia on the one hand and the Tchorok [Coruh] drainage in
western Transcausia on the other hand. They mention that the Transcaucasian popu-
lation does not show variability (which relates with its small range and presence in a
single drainage). They report considerable differences between the Transcaucasian and
the Anatolian populations. These differences include the number of scale in the lateral
line, which they record as 67-80 in the Coruh population and 72-87 in the western and
Central Anatolian populations (but they do not provide separate values for the different
drainages in this second area). Their data on the Coruh population (their Transcaucian
population) is based on at least 6 specimens from Tortum Reservoir examined by them
and 22 specimens from an unknown location from Elanidze (1983). It is not known if

REVIEW OF CAPOETA TINCA 433

the data were obtained by the same methods. Banarescu & Herzig-Straschil concluded
that the two goups of populations represent different subspecies but they did not name
the Coruh one. Examination of our material confirms that they are distinct. They satis-
fy the criteria of species under the Evolutionary Species Concept (ESC) as they are di-
agnosable and constitute a distinct lineage (Mayden, 2002; Kottelat, 1997).

Our comparison of material from the Coruh, Kızılırmak, Sakarya, Koca and
Nilüfer rivers shows that the Coruh material (C. banarescui) is immediately distin-
guished from all others by the absence of sexual dimorphism in the shape of the mouth.
In C. banarescui, the mouth is regularly arched in both sexes (Fig. 6), while in the other
species the male has a small, arched mouth, with the edge of the lower jaw rounded,
while the female has a broad, straight mouth, with the edge of the lower jaw sharp (Figs
2, 8). Capoeta banarescui also has fewer scale rows between the lateral line and the
dorsal-fin origin (12-14, mean 12.8, vs. 14-17, mean 15.7 in C. tinca, 14-17, mean 14.9
in C. baliki) and the anal-fin origin (8-9, mean 8.1, vs. 9-11, mean 9.5 in C. tinca,
10-11 mean 10.1 in C. baliki), and fewer serrae along the posterior margin of the last
simple dorsal-fin ray (12-20, mean 16.0, vs. 24-28, mean 26.6 in C. tinca, 17-23, mean
19.5, in C. baliki).

Comparison of the material from the Marmara Sea basin (Niliifer and Koca
drainages) and the southern Black Sea Basin (Sakarya and Kizilirmak drainages) also
shows that they are specifically distinct. The type locality of C. tinca is Nilüfer River
and the species from the Marmara basin retains the name C. tinca.

Capoeta banarescui is further distinguished from C. tinca by its more pointed
snout (vs. blunt and rounded). It further differs from C. tinca in having fewer gill rakers
on the first gill arch (12-16, mean 14.7, vs. 19-23, mean 20.6), fewer lateral line scales
(64-77, mean 70.8, vs. 69-80, mean 74.9), a somewhat more slender caudal peduncle
(depth 9.8-11.7% SL, mean 10.7, vs. 10.8-13.4, mean 11.7), a smaller pelvic-anal
distance (18.5-23.6% SL, mean 21.5, vs. 20.9-26.2, mean 23.6), a smaller snout depth
at level of nostrils (29.7-35.1% HL, mean 32.7, vs. 30.0-41.1, mean 34.1), longer
anterior (12.4-20.8% HL, mean 16.9, vs. 8.1-14.1, mean 10.6) and posterior barbels
(18.4-28.8% HL, mean 21.9, vs. 11.5-19.3, mean 15.2), and a wider mouth (29.5-
37.9% HL, mean 34.3, vs. 27.4-34.2, mean 30.6).

Capoeta banarescui is further distinguished from C. baliki by its more pointed
snout (vs. blunt and rounded), in having fewer gill rakers on the first gill arch (12-16,
mean 14.7, vs. 16-22, mean 19.3), fewer lateral line scales (64-77, mean 70.8, vs.
72-86, mean 78.4), more vertebrae (45-46, vs. 43-44), a somewhat smaller pelvic-anal
distance (18.5-23.6% SL, mean 21.5, vs. 20.5-25.4, mean 23.3), a narrower head (at
posterior margin of eye (49.4-58.2% SL, mean 53.8, vs. 55.6-63.6, mean 59.2), and a
smaller snout depth at level of nostril (29.7-35.1% HL, mean 32.7, vs. 33.1-41.6, mean
3712):

Capoeta baliki is distinguished from C. tinca by having fewer serrae along the
posterior margin of the last simple dorsal-fin ray (17-23, mean 19.5, vs. 24-28, mean
26.6), modally fewer scale rows between the lateral line and the dorsal-fin origin (14,
vs. 16), fewer vertebrae (43-44, modally 44, vs. 44-46, modally 45), the head shorter
(length 21.8-24.5% SL, mean 23.6, vs. 23.3-26.7, mean 24.9) and broader (width at
posterior margin of eye 55.6-63.5% HL, mean 59.2, vs. 49.3-56.5, mean 53.1), a some-

434 D. TURAN ET AL.

what more slender caudal peduncle (depth 9.5-12.2% SL, mean 10.8, vs. 10.8-13.4,
mean 11.7), and a much blunter snout (compare Figures 1 and 7).

Varicorhinus antalyensis (Fig. 9) was described by Battalgil (1944) from the
area of Antalya (on the southwestern coast of Turkey). Among other characters, it is
diagnosed by having two pairs of barbels. Karaman (1969) treated V. antalyensis as a
synonym of Hemigrammocapoeta kemali Hanko (1924). Erk’akan & Kuru (1983)
collected Capoeta specimens in Aksu and Kôprü streams near Antalya which they
identified as V antalyensis. They compared them with A. kemali and concluded that
they are not conspecific and that C. antalyensis is a valid species. We examined
25 specimens from Gökdere stream (37°24'N 31°11'E) near Antalya which we identify
as C. antalyensis. They are immediately distinguished from C. banarescui, C. baliki
and C. tinca in having fewer lateral line scales (51-57), fewer scales in tranverse line
(10-12/7) and no serration along posterior margin of last simple dorsal-fin ray. In
C. antalyensis too, the shape of the mouth is sexually dimorphic; the male has a small,
arched mouth, with the edge of the lower jaw rounded, while the female has straight
mouth (Fig. 10).

COMPARISON MATERIAL

Capoeta antalyensis: FFR 719, 10, 78-236 mm SL; CMK 18522, 6, 108-184
mm SL; Turkey: Antalya: Gökdere Stream, 40°29'N 32°39'E; D. Turan, Z. Turan & S.
Engin, 9 September 2004.

ACKNOWLEDGEMENTS

We are pleased to thank Serife Gülsün Kirankaya and Osman Serdar for their
help in the field and Semih Engin for figures.

REFERENCES

BALIK, S. 1979. Bat Anadolu tatlisu balıklarının taksonomik ve ekolojik özellikleri üzerine
arastırmalar [Taxonomical and ecological investigations upon freshwater fishes of
Western Anatolia]. Ege Üniversitesi Fen Fakiiltesi Ilmi Raporlar Serisi 236: 1-69 [In
Turkish].

BANARESCU, P. M. 1999. The freshwater fishes of Europe. 5. Cyprinidae 2. Part I. Rhodeus to
Capoeta. Aula, Wiesbaden, 426 pp.

BATTALGIL, F. 1944. Nouveaux poissons des eaux douces de la Turquie. Istanbul Universitesi
Fen Fakültesi Mecmuasi, Seri B, Tabiî Ilimler 9: 126-133.

BERG, L. S. 1912-14. [Faune de Russie et des pays limitrophes. Poissons (Marsipobranchii et
Pisces). Vol. 3. Ostariophysi]. /zdatelstvo Akademii Nauk, St-Petersburg, 1 (1912):
1-336, 2 pls, 2 (1914): 337-704, 4 pls. [In Russian].

BERG, L. S. 1948-49. [Freshwater fishes of the U.S.S.R. and adjacent countries]. /zdatelstvo
Akademii Nauk SSSR, Moskva & Leningrad, vol. 1 (1948), vols 2-3 (1949). [In Russian;
translation: Israel Program for Scientific Translations, Jerusalem, 1965].

DERIUGIN, K. M. 1899. On the ichthyofauna of southwest Transcaucasia. Ezhegodnik Zoo-
logicheskogo Muzeya Imperatorskoi Akademii Nauk [Annuaire du Musée Zoologique de
l’Académie Impériale des Sciences de St-Pétersbourg] 1899: 148-171 [not seen, from
Berg, 1949].

EKMEKCI, F. G. 1996. [Some of the growth and reproduction properties of Capoeta tinca (Heckel,
1843) in Sariyar Dam Lake (Ankara)]. Turkish Journal of Zoology 20: 117-126 [in
Turkish].

REVIEW OF CAPOETA TINCA 435

EKMEKGI, F. G. & KiRANKAYA, S. G. 2004. Determination of variation in fish growth during
reservoir ontogeny: a case study of the mirror carp population in Gelingüllü Dam Lake
(Yozgat,Turkey). Turkish Journal of Veterinary and Animal Sciences 28: 1129-1135.

EKMEKCI, G. & ÖZEREN, C. 2003. Reproductivite biology of C. tinca in Gelingüllü Reservoir,
Turkey. Folia Zoologica 52: 323-328.

ELANIDZE, R. F. 1983. Ichthyofauna in the rivers and lakes of Georgia. Metzniereva, Tbilisi, 139
pp. [not seen; from Banarescu, 1999].

ERK’AKAN, F 1981. Sakarya havzası balıklarının (Pisces) sistematigi ve biyo-ekolojik iliskileri
üzerine arastırmalar [Studies on systematics and bioecology of fishes (Pisces) of Sakarya
Basin]. PhD Thesis, Hacettepe Üniversitesi, Ankara, 144 pp. [in Turkish].

ERK’AKAN, F. & KURU, M. 1983. Re-discussion of systematical status of Varicorhinus antayen-
sis Battalgil, 1944. Hacettepe Bulletin of Natural Sciences and Engineering 12: 49-65.

GÜNTHER, A. 1868. Catalogue of the fishes in the British Museum. Vol. 7. British Museum,
London, XX +512 pp.

HANKO, B. 1924. Fische aus Klein-Asien. Annales Historico-naturales Musei Nationalis
Hungarici 21: 137-158, pl. 3.

HECKEL, J. J. 1843. Ichthyologie (pp. 991-1099). In: RUSSEGGER, J. Reisen in Europa, Asien und
Afrika mit besonderer Rücksicht auf die naturwissenschaftlichen Verhältnisse der
betreffenden Länder, unternommen in den Jahren 1835 bis 1841. Erster Band. Reise in
Griechenland, Unteregypten, im nôrdlichen Syrien und südôstlichen Kleinasien.
Schweizerbart, Stuttgart, Teil 2: 472-1102.

HuBBs, C. L. & & LAGLER, K. F. 1947. Fishes of the Great Lakes region. Cranbrook Institute of
Science, Bulletin 26: i-xi, 1-186.

KARAMAN, M. S. 1969. Siisswasserfische der Tiirkei. 7. Teil. Revision Kleinasiatischen und
Vorderesiatischen Arten des Genus Capoeta (Varicorhinus, partim). Mitteilungen aus
dem Hamburgischen Zoologischen Museum und Institut 66: 17-54, pls 1-6.

KosswIG, C. & BATTALGIL, F. 1942. Beiträge zur Türkischen Faunengeshichte I. Süsswasser-
flische. Türk Fizikî ve Tabiî Ilimler Sosyetesi Yillik Bildirigleri ve Arsivi [Compte Rendu
Annuel de la Société Turque des Sciences Physiques et Naturelles] 8: 32-60.

KOTTELAT, M. 1997. European freshwater fishes. An heuristic checklist of the freshwater fishes
of Europe (exlusive of former USSR), with an introduction for non-systematists and
comments on nomenclature and conservation. Biologia (Bratislava) 52 (Suppl. 5):
1-271.

Kuru, M. 1975. Dicle-Fırat, Kura-Aras, Van Gölü ve Karadeniz havzası tatlısularında yasayan
baliklarin (Pisces) sistematik ve zoocografik yönden incelenmesi [Systematics and zoo-
geographical study of fishes (Pisces) living in Tigris-Euphrates, Kura-Arax, Van Lake
and Blacksea basins]. Thesis, Atatiirk Universitesi, Fen Fakiiltesi, Erzurum, 181 pp. [in
Turkish].

KUTRUP, B. 1994. Trabzon yöresindeki tatlisu balıklarının taksonomik ve ekolojik yönden ince-
lenmesi [Taxonomic and ecological study of freshwater fish in Trabzon region]. PAD
Thesis, Karadeniz Teknik Üniversitesi, Fen Bilimleri Enstitüsü, Trabzon, 64 pp. [in
Turkish].

MAYDEN, R. L. 2002. On biological species, species concepts and individuation in the natural
world. Fish and Fisheries 3: 171-196.

SOLAK, K. 1982. Coruh ve Aras havzasında yasayan siraz balılıklarının (Capoeta sp.) biyoloji ve
ekolojileri üzerinde arastırmalar [Studies on biology and ecology of siraz fishes
(Capoeta sp.) living in Coruh and Arax basins]. Thesis, Atatürk Üniversitesi, Temel
Bilimler ve Yabancı Diller Yüksek Okulu, Erzurum, 135 pp. [in Turkish].

STEINDACHNER, F. 1897. Bericht über die von Dr. Escherich in der Umgebung von Angora
gesammelten Fische and Reptilien. Denkschriften der Kaiserlichen Akademie der
Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Classe, Abt. 1, 40: 685-
699, 3 pls.

YILDIRIM, A. & ARAS, S. 2000. Some reproduction characteristics of Capoeta tinca (Heckel,
1843) living in the Oltu Stream of Coruh Basin. Turkish Journal of Zoology 24: 95-102.

436 D. TURAN ET AL.

YILMAZ, M. 1994. Kapulukaya Baraj Gölü (Kırıkkale) ’nde yasayan sazan (Cyprinus carpio L.,
1758) ve in baligi (Capoeta tinca Heckel, 1843) ’nin biyo-ekolojik özellikleri
[Bioecological properties of carp sazan (Cyprinus carpio L., 1758) and (Capoeta tinca
Heckel, 1843) living in Kapulukaya Reservoir (Kırıkkale)]. PhD Thesis, Gazi Univer-
sitesi Fen Bilimleri Enstitusü, Ankara, 290 pp. [in Turkish].

REVUE SUISSE DE ZOOLOGIE

Tome 113 — Fascicule 2

Ding YANG, Bernhard MERZ & Patrick GROOTAERT. Descriptions of three
new Platypalpus Macquart from Guangdong, China (Diptera,
Elybotidae*TFachydiomMimMo) Re IA

Wolfgang SCHAWALLER. New species and records of the genus Basanus
Lacordaire (Insecta: Coleoptera: Tenebrionidae) .................

Wilson R. LOURENCO & Steven M. GOODMAN. Further considerations
regarding the status of Grosphus madagascariensis (Gervais) and
Grosphus hirtus Kraepelin, and description of a new species
(Scorpiones,Buthidae). cr tru Rn re eek

Adrian SMOLIS & Louis DEHARVENG. Vitronura mascula, a new species of
Neanurinae (Collembola: Neanuridae) from northern Vietnam, with a
LEYAtOMNEYSPECIESTORINE LENUS ann m RR e e

Daniel BURCKHARDT & Pavel LAUTERER. The Palaearctic triozids associated
with Rubiaceae (Hemiptera, Psylloidea): a taxonomic re-evaluation of
thesziıozargaliinEoerster complexe). IN

Koen MAES. A new species of Diathrausta Lederer, 1863 from Africa
(Lepidoptera, Pyraloidea, Crambidae, Spilomelinae) ..............

Dariusz SKARZYNSKI. Redescription of Ceratophysella lawrencei (Gisin,
I963)(Collembola#Hypogastruridae) Same KR EEE

Dariusz SKARZYNSKI & Adrian SMOLIS. Description of Ceratophysella
robustiseta sp. n. from greenhouses in England, with notes on syno-
nymy of C. postantennalis Yosii, 1966 and taxonomic status of C.
morula Deharveng & Bourgeois, 1991 (Collembola: Hypogastru-
pi da) pere an Te Rn

Juan Marcos MIRANDE, Gastôn AGUILERA & Maria de las Mercedes
AZPELICUETA. Nomenclatural note on the genus Nans (Ostariophysi,
Gharacidae Per ee e E E VE CS RR OPEN ENO CIAO

Bernard LANDRY, David ADAMSKI, Patrick SCHMITZ, Christine E. PARENT
& Lazaro ROQUE-ALBELO. Taygete sphecophila (Meyrick) (Lepido-
ptera; Autostichidae): redescription of the adult, description of the
larva and pupa, and impact on Polistes wasps (Hymenoptera;
Vespidae) nests in the Galapagos Islands .....................

Peter SCHUCHERT. The European athecate hydroids and their medusae
(Hiydrozoay Enidania)a@apitata Part le nn

Nora KHAMMES, Sovan LEK & Stéphane AULAGNIER. Identification bio-
métrique des deux espèces sympatriques de souris Mus musculus
domesticus et Mus spretus en Kabylie du Djurdjura (Algérie) . .....

Davut TURAN, Maurice KOTTELAT, F. Güler EKMEKCI & H. Oguz
IMAMOGLU. A review of Capoeta tinca, with descriptions of two new
species;iromelurkey/(deleostei:Gyprini dae) i serie.) Cat e

Pages

229-238

239-246

247-261

263-268

269-286

287-290

291-296

297-303

305

307-323

325-410

411-419

421-436

REVUE SUISSE DE ZOOLOGIE

Volume 113 — Number 2

Ding YANG, Bernhard MERZ & Patrick GROOTAERT. Descriptions of three
new Platypalpus Macquart from Guangdong, China (Diptera,
iy botidaemachydromiinae) ILE E
Wolfgang SCHAWALLER. New species and records of the genus Basanus
Lacordaire (Insecta: Coleoptera: Tenebrionidae) .................
Wilson R. LOURENCO & Steven M. GOODMAN. Further considerations
regarding the status of Grosphus madagascariensis (Gervais) and
Grosphus hirtus Kraepelin, and description of a new species
(ScorpionesmButhidae) sis sane wis ae we ae ore 5 CRE E
Adrian SMOLIS & Louis DEHARVENG. Vitronura mascula, a new species of
Neanurinae (Collembola: Neanuridae) from northern Vietnam, with a
keyktoithezspecies.ofitheigenus.. 4%. 2... en eee ee
Daniel BURCKHARDT & Pavel LAUTERER. The Palaearctic triozids associated
with Rubiaceae (Hemiptera, Psylloidea): a taxonomic re-evaluation of
themiozascalimEoerster complex... RENE SERRE
Koen Mags. A new species of Diathrausta Lederer, 1863 from Africa
(Lepidoptera, Pyraloidea, Crambidae, Spilomelinae) ..............
Dariusz SKARZYNSKI. Redescription of Ceratophysella lawrencei (Gisin,
I1963)(Collembola:Hypogastruridae)) MALE aes SERRE
Dariusz SKARZYNSKI & Adrian SMOLIS. Description of Ceratophysella
robustiseta sp. n. from greenhouses in England, with notes on syno-
nymy of C. postantennalis Yosii, 1966 and taxonomic status of C.
morula Deharveng & Bourgeois, 1991 (Collembola: Hypogastru-
MIO) RE RL RI
Juan Marcos MIRANDE, Gastön AGUILERA & Maria de las Mercedes
AZPELICUETA. Nomenclatural note on the genus Nans (Ostariophysi,
Characidace) erate as sus oA se oie iI I
Bernard LANDRY, David ADAMSKI, Patrick SCHMITZ, Christine E. PARENT
& Lazaro ROQUE-ALBELO. Taygete sphecophila (Meyrick) (Lepido-
ptera; Autostichidae): redescription of the adult, description of the
larva and pupa, and impact on Polistes wasps (Hymenoptera;
Vespidae) nests in the Galapagos Islands? 2m. EEE
Peter SCHUCHERT. The European athecate hydroids and their medusae
(Hiydrozoa, (Cnidaria): Capitata Part 1.3.55 2. eee
Nora KHAMMES, Sovan LEK & Stéphane AULAGNIER. Biometrical identi-
fication of the sympatric mouse species Mus musculus domesticus
and Mus spretus in Kabylie du Djurdura (Algeria) ..............
Davut TURAN, Maurice KOTTELAT, F. Güler Ekmekci & H. Oguz
IMAMOGLU. A review of Capoeta tinca, with descriptions of two new
Species from, Turkey, (Teleostei: Cyprinidae) ERRORE

Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX

Pages

229-238

239-246

247-261

263-268

269-286

287-290

291-296

297-303

305

307-323

325-410

411-419

421-436

Volume 113 - Number 2 - 2006
Revue suisse de Zoologie: Instructions to Authors

The Revue suisse de Zoologie publishes papers by members of the Swiss Zoological Society and scientific
results based on the collections of the Muséum d’histoire naturelle, Geneva. Submission of a manuscript implies
that it has been approved by all named authors, that it reports their unpublished work and that it is not being
considered for publication elsewhere. A financial contribution may be asked from the authors for the impression of
colour plates and large manuscripts. All papers are refereed by experts.

In order to facilitate publication and avoid delays authors should follow the Instructions to Authors and refer to a
current number of R.S.Z. for acceptable style and format. Papers may be written in French, German, Italian and
English. Authors not writing in their native language should pay particular attention to the linguistic quality of the text.

Manuscripts must be typed or printed (high quality printing, if possible by a laser-printer), on one side only
and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins
must be at least 25 mm wide. Authors must submit three paper copies (print-outs), including tables and figures, in
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the revised version of the accepted manuscript.

We encourage authors to submit the revised final text on a disk (3,5”), or on a CD-R, using MS-WORD or a
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which should be formatted in italics (bold italics in taxa headings) and authors’ names in the list of references (not
in other parts of the text!), which should be formatted in SMALL CAPITALS. LARGE CAPITALS may be used for
main chapter headings and SMALL CAPITALS for subordinate headings. Footnotes and cross-references to specific
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Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, full
name(s) and surname(s) of author(s), and full address(es) including e-mail address(es) if possible.

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should sum-
marise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed by up
to 10 keywords, separated by hyphens, which are suitable for indexing. Some of the terms used in the title may be
omitted from the list of keywords in favour of significant terms not mentioned in the title.

Introduction. A short introduction to the background and the reasons for the work.

Material and methods. Sufficient experimental details must be given to enable other workers to repeat the
work. The full binominal name should be given for all organisms. The International Code of Zoological Nomen-
clature must be strictly followed. Cite the authors of species on their first mention.

Results. These should be concise and should not include methods or discussion. Text, tables and figures should
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp.
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by
synonyms, material examined and distribution, description and comments. All material examined should be listed in
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex
symbols should be used rather than “male” and “female” (text file: $ = 6, £= 2).

Discussion. This should not be excessive and should not repeat results nor contain new information, but
should emphasize the significance and relevance of the results reported.

References. The author-date system (name-year system) must be used for the citation of references in the text,
e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown
et al. (1995) or (Brown et al., 1995; White et al., 1996) should be used. In the text authors’ names have to be
written in standard type face. However, in the list of references they should be formatted in SMALL CAPITALS (see
below). The list of references must include all publications cited in the text and only these. References must be
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. Thèse, Genève, 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, XII + 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. If possible, 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
electronically scanned figures are submitted on disk or CD-R (never in MS-Powerpoint format !), this should be clearly
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should be respected: scanned line drawings: 600 dpi; photographs: 300 dpi.

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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@mhn.ville-ge.ch
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

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