Zoological Journal of the Linnean Society, 2008, 152, 635–653. With 10 figures
Proposal of new genus for Asterocheres mucronipes
Stock, 1960 (Copepoda, Siphonostomatoida,
Asterocheridae), an associate of the scleractinian coral
Astroides calycularis (Pallas, 1766) in the Strait of
Gibraltar
Ma EUGENIA BANDERA1 and RONY HUYS
FLS2*
1
Biodiversidad y Ecología de Invertebrados Marinos, Departamento de Fisiología y Zoología,
Facultad de Biología. Universidad de Sevilla. Reina Mercedes, 6, 41012-Sevilla, Spain
2
Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK
Received 2 October 2006; accepted for publication 12 June 2007
A new genus of Asterocheridae (Copepoda: Siphonostomatoida) Stockmyzon gen. nov. is proposed for Asterocheres
mucronipes Stock, 1960, and a new, previously misidentified, species Stockmyzon crassus sp. nov. from sponge
washings in Mauritius. Stockmyzon gen. nov. can be differentiated from other asterocherid genera by the
annulated mandibular stylet, the atrophied maxillulary palp with large modified lateral seta, the presence of
beak-shaped processes on the endopods of legs 1, and the transformation of the outer spine on the first exopodal
segment of leg 4 into a seta. Stockmyzon mucronipes comb. nov. is the fourth copepod known to utilize the
hermatypic coral Astroides calycularis (Pallas, 1766) as its host in the Strait of Gibraltar. A reinterpretation of the
original description of Asterocheres stimulans Giesbrecht, 1897 from Naples revealed that it was based on an
amalgam of two diffent species, the male being conspecific with S. mucronipes; the illustrated female is formally
designated here as the lectotype of A. stimulans. The current symbiotic relationship between S. mucronipes and the
gorgonian Eunicella singularis (Esper, 1794) along the French mediterranean coast is reviewed in the light of
potential host switching, following the extinction of A. calycularis in the north-western Mediterranean, north of
40°N, during the late Sicilian regression (Rissian age), about 238 000–225 000 years ago. © 2008 The Linnean
Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653.
ADDITIONAL KEYWORDS: glaciation – scanning electron microscopy – Scleractinia – symbiosis – taxonomy.
INTRODUCTION
Siphonostomatoid copepods are almost exclusively
symbiotic, and utilize a wide range of invertebrate
and vertebrate hosts all around the world (Gotto,
1979; Ho, 1982; Humes, 1993; Humes, 1996; Ivanenko
& Smurov, 1997; Kim, 1998; Boxshall & Halsey,
2004). Those that live as external or internal symbionts of marine invertebrates primarily utilize
sponges, cnidarians, echinoderms, bryozoans, molluscs, and ascidians, but for many members of the
families Asterocheridae and Artotrogidae the hosts
*Corresponding author. E-mail: rjh@nhm.ac.uk
are still unknown. Substantial gaps remain in our
knowledge of symbiotic copepods, even in areas where
marine invertebrates have been the subject of comprehensive investigation. For example, in European
waters siphonostomatoids are most commonly reported from sponges, but documented associations
with scleractinian corals are scarce. This conceivably
reflects sampling bias rather than host-phylum preference. Five years ago, an ongoing programme on the
biology of the hermatypic scleractinian Astroides calycularis (Pallas, 1766) was initiated around Tarifa
Island (Strait of Gibraltar), where it represents the
most important macrobenthic organism in shallow
waters. Although there were no previous records of
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
635
636
M. E. BANDERA and R. HUYS
copepods inhabiting A. calycularis, this coral species
turned out to be the host of a variety of symbiotic
copepods, reinforcing Humes’s (1994) hypothesis that
virtually any marine macroinvertebrate is a potential
host to copepods. Recently, Conradi, Bandera &
López-González (2006) described two new species,
Doridicola helmuti (Rhynchomolgidae) and Asterocheres astroidicola (Asterocheridae), and listed the
first record of Acontiophorus scutatus (Brady & Robertson, 1873) from this coral host. Here, we describe
another new asterocherid that exhibits similarities
with members of the genus Asterocheres, but also
displays some important differences. Comparison of
the nearly 70 species currently assigned to the genus
revealed that one species, Asterocheres mucronipes
Stock, 1960, was morphologically very similar to the
specimens recovered from the Astroides colonies.
Stock’s (1960) description was based on four females
obtained from washings of the gorgonian Eunicella
verrucosa (Pallas, 1766), found at a depth of 30 m
near Cap Béar along the French mediterranean coast
(Roussillon). In a later paper, Stock (1966) emended
the description based on 19 females in washings of
an orange sponge, possibly a species of Oscarella
Vosmaer, 1884, from Mauritius. Stock (1966) also
corrected the identification of the Roussillon host to
Eunicella stricta (Bertoloni, 1810), but the latter is
now generally regarded as a junior synonym of the
white seafan Eunicella singularis (Esper, 1794)
(cf. Weinberg, 1976, 1978). A re-examination of Stock’s
(1960) type material of Asterocheres mucronipes in
the Zoological Museum of Amsterdam proved that the
specimens from Tarifa were conspecific with the Roussillon population. Contrary to Stock’s (1966) opinion,
the material from Mauritius differed significantly
from both Mediterranean populations, justifying the
proposal of a new species. In this paper we establish
a new genus, Stockmyzon, to accommodate the
type species Stockmyzon mucronipes (Stock, 1960)
comb. nov., and a new species Stockmyzon crassus
sp. nov. is proposed for Stock’s (1966) specimens from
Mauritius; detailed descriptions of both species are
presented.
MATERIAL AND METHODS
Colonies of A. calycularis were individually collected
by SCUBA diving at Tarifa Island, and were immediately isolated in separate plastic bags containing a
solution of 8–10% formaldehyde in seawater. Symbiotic fauna was obtained by pouring the wash water
through a 100-mm net. Copepods were extracted from
the filtrate and preserved in 70% ethanol.
Selected specimens were dissected in lactic acid and
examined as temporary mounts in lactophenol. For
scanning electron microscopy, a specimen of each
species was dehydrated in a graded series of ethanol,
critical-point dried, mounted on stubs, coated with
a gold–palladium alloy, and examined in a Phillips XL30 SEM. All figures were drawn with the aid of
a camera lucida mounted on a Zeiss Axioskop differential interference contrast microscope. All appendage segments and setation elements are named and
numbered using the terminology introduced by Huys
& Boxshall (1991).
Material from Tarifa was deposited in the Zoological Museum of Amsterdam (ZMA), and in the collection of the research team Biodiversidad y Ecología de
Invertebrados Marinos of the University of Seville
(BEIM).
SYSTEMATICS
ORDER SIPHONOSTOMATOIDA BURMEISTER, 1835
FAMILY ASTEROCHERIDAE GIESBRECHT, 1899
GENUS STOCKMYZON GEN. NOV.
Diagnosis: Asterocheridae. Body: cyclopiform, comprising dorsoventrally flattened prosome and cylindrical urosome. Siphon of medium size, reaching to or
slightly beyond rear margin of cephalothorax. Sexual
dimorphism present in prosome width, urosomal segmentation, antennules, maxillipeds, and leg 6.
Urosome: four-segmented in female; five-segmented
in male. Antennule: 20-segmented in female, with
large aesthetasc on segment 18; 18-segmented in
male, with large aesthetasc on segment 17 and
geniculation located between segments 16 and 17.
Antenna: with large one-segmented exopod and threesegmented endopod with terminal claw. Mandibular
palp: two-segmented, second segment with two
plumose setae; stylet with annulation in middle part
and denticulate margin subapically. Maxillule:
bilobed, with a rectangular praecoxal endite, and
atrophied palp bearing large characteristically
plumose seta and two or three accessory setae.
Maxilla: two-segmented, with aesthetasc-like tubular
extension on praecoxal portion of syncoxa, and a
claw-like basis recurved towards the apex. Maxilliped: comprising short syncoxa, long basis, and threesegmented endopod; male basis with spinous process
close to syncoxa–basis joint. Legs 1–4: biramous with
three-segmented rami; middle and distal endopodal
segments with beak-shaped spiniform processes.
Outer element on proximal exopodal segment of leg 4:
setiform. Leg 5: with protopod incorporated into
somite (represented by dorsal surface seta) and onesegmented exopod bearing three setae.
Etymology: The genus is named in honour of the
late Prof. Jan Hendrik Stock (Zoölogisch Museum,
Amsterdam) who described its type species. The
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
Greek suffix -myzon (muz ω′ ), meaning to suck, is commonly used in the formation of siphonostomatoid
generic names, and refers to the sucking oral cone or
siphon. Gender: male.
Type species: Asterocheres mucronipes Stock, 1960
= Stockmyzon mucronipes (Stock, 1960) comb. nov.
Other species: Stockmyzon crassus sp. nov.
Remarks
Stock (1960) placed his new species A. mucronipes in
Asterocheres, but expressed some reservations about
his generic assignment. Although he recognized a
superficial similarity in the enlarged prosome with
some other Asterocheres species, such as Asterocheres
lilljeborgi Boeck, 1859 and Asterocheres ovalis Sewell,
1949, certain other characters exhibited by A. mucronipes were considered more significant and potentially of ‘valeur générique’. In particular, Stock (1960)
mentioned the characteristic endopodal spinous processes on legs 1–4, the unusual armature of leg 4
(proximal exopod segment with outer seta), the ‘biarticulated’ mandibular stylet, and the presence of only
two setae (instead of four) on the maxillulary palp,
one of which being enlarged (‘aspect gonflé’). Stock
also noted that the male of Asterocheres stimulans
Giesbrecht, 1897 has similar spinous processes on
legs 1–4 (Giesbrecht, 1899: plate 3). He also claimed
that the maxillulary palp of Asterocheres canui
Giesbrecht, 1897 [= A. lilljeborgi sensu Canu (1892);
cf. Giesbrecht (1897): 11)] displays a transitionary
state, between the typical Asterocheres condition and
that in A. mucronipes, having retained the typical
number of four terminal setae, with one of them being
gonflate. Based on these observations, Stock (1960)
maintained a tentative assignment of A. mucronipes
to Asterocheres was warranted. However, our reinterpretation of Giesbrecht’s (1899) illustrations of male
A. stimulans revealed that it is conspecific with
A. mucronipes (see the Discussion), and comparison of
Canu’s (1892) figure of the maxillule showed it to be
quite different from the A. mucronipes condition, but
remarkably similar to that of other typical Asterocheres species, such as Asterocheres reginae Boxshall
& Huys, 1994 (Boxshall & Huys, 1994: fig. 3F). The
palp in A. canui is not atrophied, and the lateral seta
is not enlarged, excessively plumose, or typically
recurved and concealed under the gnathobasal endite,
as it is in A. mucronipes.
Although A. mucronipes resembles species of
Asterocheres in several aspects, such as the long,
multisegmented antennule, the antenna with onesegmented exopod and three-segmented endopod, the
segmentation of the maxilla and maxilliped, and the
637
one-segmented leg 5 bearing three setae, it differs in
a number of characters, warranting the proposal of a
new genus.
The mandible of Stockmyzon has a two-segmented
palp, which is shared by over two-thirds of the species
of Asterocheres; however, none of these exhibits the
distinctly annulated stylet. This character is regarded
here as an autapomorphy of the new genus. In some
asterocherids the mandibular stylet shows a thinning
of the cuticle halfway along its length, but never a
strong annulation. Johnsson (1998) illustrated a long
‘segmented’ stylet in his description of Asterocheres
crenulatus Johnsson, 1998, but a re-examination of
a female paratype (NHM reg. no. 1997.185) revealed
this to be an observational error, possibly as a result
of excessive squashing during the mounting process.
The bilobate maxillule of Stockmyzon is unique in
its marked size disparity between the outer (palp)
and the inner lobe (gnathobase). The palp is atrophied and has two or three small setae, in addition to
a large, densely plumose lateral seta. Furthermore,
within the Asterocheridae a somewhat similar condition is only found in Acontiophorus Brady, 1880
(e.g. Kim & Je, 2000), but this genus represents a
completely different lineage in the family, deviating
from all others in the morphology of the antennule,
antenna, and mandible. The primitive leg 5 and
swimming leg armature formula also indicate a very
basal position in the Asterocheridae.
The spine and seta formula of the swimming legs in
the new genus is similar to that of Asterocheres,
except for leg 4, which has an outer seta on the first
exopodal segment in Stockmyzon, instead of an outer
spine. The transformation of this element into a seta
is a unique apomorphy within the Asterocheridae. A
similar transformation on the basis of leg 1 (seta
replaced by spine) in A. crenulatus and Asterocheres
spinopaulos Johnsson, 1998 [and three other species
described by Johnsson (1998)] was recently considered potential justification for their removal to a
distinct genus (Kim, 2004b).
Perhaps the most conspicuous feature of Stockmyzon is the presence of large beak-shaped spiniform
processes on the endopodal segments of legs 1–4. This
character has been recorded in some genera previously allocated to the Coralliomyzontidae (e.g. Humes
& Stock, 1991; Humes, 1997), which coincidently also
utilize scleractinian corals as hosts. Within the speciose genus Asterocheres, currently encompassing 67
valid species (Kim, 2004a, b, 2005; Bandera, Conradi
& López-González, 2005; Bispo, Johnsson & Neves,
2006; Conradi et al., 2006), only Asterocheres tubiporae Kim, 2004 exhibits similar modifications on leg 1.
In every other aspect this species is a typical representative of the genus Asterocheres, and consequently
the spinous processes on the leg-1 endopod are likely
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
638
M. E. BANDERA and R. HUYS
to have resulted from convergence. Also note that Kim
(2004b) erroneously described and illustrated the
female antennule of A. tubiporae as 22-segmented; no
other extant siphonostomatoid has more than 21 segments. Comparison with A. reginae (Boxshall & Huys,
1994: fig. 3A–E) suggests Kim (2004b) had inadvertently intercalated a supernumerary segment between
the spine-bearing segment XIV and the aesthetascbearing segment XXI.
Stockmyzon is related to a group of genera that
exhibit a tubular extension of the opening of the
maxillary gland. Although some reports have suggested that this character may be widely distributed
within the Asterocheridae (Boxshall & Huys, 1994), it
has thus far been found only in Asterocheres (e.g. Ho,
1984; Boxshall & Huys, 1994; Ivanenko & Smurov,
1997; Ivanenko, 1997; Kim, 2004a, b), Inermocheres
Boxshall, 1990 and Sinopontius Boxshall, 1990 (Boxshall, 1990), Dermatomyzon Claus, 1889 (Ivanenko &
Ferrari, 2003), and, to a lesser extent, Laperocheres
Ivanenko, 1998.
STOCKMYZON MUCRONIPES
(STOCK, 1960) COMB. NOV.
Synonyms: Asterocheres mucronipes Stock, 1960;
Asterocheres stimulans Giesbrecht, 1897 (씹 only; see
the Discussion).
Original description: Stock (1960: 224–228, figs 4, 5).
Type locality: France, Roussillon; off Cap Béar (near
Banyuls-sur-Mer); washings of E. singularis collected
at a depth of 30 m.
Material examined: (a) Holotype female and one
paratype female (deposited in ZMA) from type locality; collected by J.H. Stock, 17 June 1959; (b) five
females and nine males (deposited in ZMA) associated
with A. calycularis off Tarifa Island (southern Spain)
at a depth of 10–20 m; collected September 1999, by
SCUBA diving; (c) additional specimens from Tarifa
Island deposited in BEIM.
Description
Female: Body (Fig. 1A–B): cyclopiform, consisting of
dorsoventrally flattened prosome and cylindrical
urosome. Total length from anterior margin of
rostrum to posterior margin of caudal rami, 603 mm;
maximum width, 465 mm measured at 4/5 length of
cephalothorax. Prosome: comprising cephalothorax
(fully incorporating first pedigerous somite) and three
free pedigerous somites. Cephalothorax (Fig. 1B)
with posterolateral angles produced into backwardly
directed processes. Rostrum completely fused to
cephalothorax, forming triangular ventrally deflected
lobe. Somites bearing legs 2–3, broad; epimeral areas
with posterolateral angles rounded (leg 2) or pointed
(leg 3) (Fig. 1B). Somite bearing leg 4: much smaller
and narrower than preceding ones. Dorsal cephalothoracic shield and free pedigerous somites ornamented with numerous integumental pores and
sensilla.
Urosome: four-segmented, comprising leg-5-bearing
somite, genital double somite, and two free abdominal
somites. Except for leg-5-bearing somite, all other
urosomites ornamented with large, flattened epicuticular scales, arranged in irregular overlapping
pattern dorsally (Fig. 1D) and ventrally (not shown
in Fig. 1C); scales occasionally with incised distal
margin. Posterior hyaline frills of urosomites with
serrate free margins (Fig. 1C). Leg-5-bearing somite:
narrow, largely concealed under pleurotergite of leg4-bearing somite. Genital double somite (Fig. 1C–D):
laterally produced; about 1.65 times wider than long;
paired genital apertures bipartite, each comprising
lateroventral copulatory pore and dorsolateral gonopore (oviduct opening); lateral margins with setular
tufts in distal third (posterior to genital apertures).
Caudal rami (Fig. 1C–D): about as long as wide
(measured along outer margin); trapezoid with inner
margin much shorter than outer one; entirely covered
by overlapping epicuticular scales; armed with six
setae; seta I absent, setae II–VII all arranged around
posterior margin, with setae II and VII slightly displaced onto dorsal surface.
Antennule (Fig. 1E–G): 20-segmented, about
250-mm long, lengths of segments (measured along
posterior nonsetiferous margin) 16 (30-mm along anterior margin), 7, 8, 8, 7, 7, 6, 11, 11, 1, 7, 13, 16, 14, 14,
13, 13, 13, 8, and 14 mm, respectively. Segmental
fusion pattern as follows (Roman numerals indicating
ancestral segments): 1(I), 2(II), 3(III), 4(IV), 5(V),
6(VI), 7(VII), 8(VIII), 9(IX–XII), 10(XIII), 11(XIV),
12(XV), 13(XVI), 14(XVII), 15(XVIII), 16(XIX),
17(XX), 18(XXI), 19(XXII–XXIII), 20(XXIV–XXVIII).
Segments 1–8, each with two setae, one of which is
plumose; segment 9, with seven setae and a small
spine; segments 10–11, each with one seta and one
small spine at anterodistal corner; segments 12–17,
each with two setae; segment 18, with two setae plus
an aesthetasc; segment 19, with one anterior, one
ventral, and one posterior seta; segment 20, with ten
setae (Fig. 1G). Segment 10(XIII): reduced, forming
incomplete sclerite, partly overlapped by distal
expansion of compound segment 9(IX–XII) (Fig. 1F).
Antenna (Fig. 2A–B): biramous. Coxa unarmed,
with few spinules. Basis unarmed, with fine spinule
rows as shown in Figure 2A. Exopod: one-segmented,
slender, about 2/5 length of proximal endopod
segment; with one small lateral seta and two terminal
setae. Endopod: three-segmented; proximal segment
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
639
Figure 1. Stockmyzon mucronipes (Stock, 1960) comb. nov. (female). A, habitus, dorsal; B, habitus, lateral; C,
urosome (excluding leg-5-bearing somite), ventral; D, urosome, dorsal; E, antennule, ventral; F, detail of antennulary
segments IX–XII, XIII, and XIV; G, detail of antennulary segments XXI, XXII–XXIII, and XXIV–XXVIII.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
640
M. E. BANDERA and R. HUYS
Figure 2. Stockmyzon mucronipes (Stock, 1960) comb. nov. (female). A, antenna; B, detail of second and third
endopodal segments of antenna; C, mandible; D, maxillule, dorsal (posterior); E, detail of praecoxal gnathobase of
maxillule, dorsal (posterior); F, maxilla; G, maxilliped.
elongated, ornamented with lateral and distal rows of
fine spinules, as illustrated; middle segment produced
distally on medial side, but articulating with distal
segment proximally on lateral side (Fig. 2B), bearing
one distal seta; distal segment with large distal claw,
one well-developed pinnate seta, and two short,
smooth setae; outer margin of distal segment with
few coarse spinules and surface with long setules.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
Siphon (Figs 3B, D): long and slender, reaching
nearly to the posterior margin of the intercoxal sclerite of leg 2 (Fig. 1B).
Mandible (Fig. 2C): comprising stylet-like gnathobase and slender two-segmented palp. Proximal
segment of palp: longest, ornamented with rows of
spinules; distal segment minute, with two plumose,
unequal apical setae. Stylet located in oral cone,
formed by anterior labrum and posterior labium
(Fig. 3D). Stylet: with annulation (not a genuine
articulation) at about halfway along its length; basal
part relatively more chitinized, distal part flexible
with denticulate margin subapically (Fig. 3D).
Maxillule (Figs 2D–E, 3E): bilobed; praecoxal gnathobase (inner lobe) distinctly larger than palp (outer
lobe). Praecoxal endite: rectangular, ornamented with
long setules proximally and spinules distally on the
lateral margin, and with a row of long setules medially (Fig. 2E); armed with one short and four long but
unequal setae, latter ornamented with short spinules
proximally and setules distally. Palp strongly
reduced, atrophied, with one elongate strongly
plumose (Fig. 3E) and three shorter pinnate setae.
Maxilla (Fig. 2F): two-segmented, but with partial
transverse surface suture on syncoxa (proximal
segment), possibly marking the plane of the praecoxa–
coxa fusion; praecoxal portion bearing flaccid
aesthetasc-like element medially, representing tubular
extension of external opening of maxillary gland; coxal
portion unarmed, but ornamented with a row of
spinules medially. Basis: claw-like, more or less
straight, but recurved towards the apex; armed with
two vestigial setae in middle third; distal inner margin
of claw provided with a double row of minute spinules.
Maxilliped (Figs 2G, 4C): five-segmented, comprising short syncoxa, long basis, and three-segmented
endopod. Syncoxa: with one short seta distally. Basis:
with a row of spinules on distal outer margin. First
endopodal segment: bearing two short distal setae;
second endopodal segment compound, partial suture
marking original separation of two ancestral segments, with (0,1) armature formula; third endopodal
segment bearing recurved terminal claw plus additional apical seta. Distal margin of claw provided with
rows of minute spinules; apex with pore (Fig. 4C).
Swimming legs 1–4 (Fig. 5A–D): biramous, with
three-segmented protopods (praecoxa not shown in
Fig. 5A–D, but see Fig. 4A for complete protopod) and
three-segmented rami. Intercoxal sclerite present in
legs 1–4, ornamented with patches of spinules in
legs 1–2.
See Table 1 for the spine and seta formula.
Coxae ornamented with spinule rows around outer
margin; inner coxal seta short and bare in leg 1, long
and plumose in legs 2–3, and absent in leg 4. Bases of
P1–P3: with spinules around inner margin; outer seta
Table 1. Spine and seta formula
mucronipes (Stock, 1960) comb. nov.
Leg 1
Leg 2
Leg 3
Leg 4
of
Stockmyzon
Coxa Basis Exopod
Endopod
0–1
0–1
0–1
0–0
0–1;
0–1;
0–1;
0–1;
1–1
1–0
1–0
1–0
I-1; I-1; III,2,2
I-1; I-1; III,I+1,3
I-1; I-1; III,I+1,3
1–1; I-1; III,I+1,3
641
0–2;
0–2;
0–2;
0–2;
1,2,3
1,2,3
1,1+I,3
1,1+I,2
plumose in leg 2, but smooth in other legs, and
extremely long in leg 1. Outer spines of exopodal
segments in legs 2–4 bilaterally serrate; in leg 1,
smooth with subapical tubular extension. Lateral
margins of exopodal segments: with minute serrations or spinular rows; those of endopodal segments
with rows of setules. Middle and distal endopodal
segments in legs 1–4: with a beak-shaped spiniform
process distally (Fig. 4B). Outer element on proximal
exopodal segment of leg 4: setiform instead of spiniform (as in legs 1–3).
Fifth leg (Fig. 1D): with protopod incorporated into
somite; outer basal seta displaced to laterodorsal
surface. Free segment (exopod): elongate-oval, with
three smooth setae distally; outer and inner margins
with spinules.
Sixth leg represented by paired opercular plates
closing off gonopores on genital double somite; each
armed with one plumose seta and one spiniform
element.
Male: Mean body length, 463 mm (450–480 mm), and
greatest width, 323 mm (320–430 mm) (N = 3). Sexual
dimorphism present in prosome width, urosomal
segmentation, antennules, maxillipeds, and leg 6.
Prosome (Fig. 6A): broader than in female, about 1.05
times wider than long. Urosome (Fig. 6B): fivesegmented, comprising leg-5-bearing somite, genital
somite, and three free abdominal somites. Dorsal
surface of genital somite, and dorsal and ventral
surfaces of free abdominal somites, ornamented with
large, epicuticular scales arranged in an irregular
overlapping pattern; scales occasionally with serrate
distal margin. Posterior margin of urosomites: ornamented with hyaline frills with serrate free margins.
Genital somite about 1.4 times wider than long.
Antennule (Figs 3C, 6D–F): 18-segmented, geniculate with geniculation positioned between segments
16(XIX–XX) and 17(XXI–XXIII). Segmental fusion
pattern as follows (Roman numerals indicating ancestral segments): 1(I), 2(II), 3(III), 4(IV), 5(V), 6(VI),
7(VII), 8(VIII), 9(IX–XII), 10(XIII), 11(XIV), 12(XV),
13(XVI), 14(XVII), 15(XVIII), 16(XIX–XX), 17(XXI–
XXIII), 18(XXIV–XXVIII). Segments 1–8, each with
two setae; segment 9, with eight setae; segment 10,
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
642
M. E. BANDERA and R. HUYS
Figure 3. Stockmyzon mucronipes (Stock, 1960) comb. nov. SEM micrographs. A, rostral area (female); B, oral cone
(female); C, antennulary segments XVIII and XIX–XX (male); D, apical part of labrum and stylet-like gnathobases of
mandibles (female); E, large plumose seta on maxillulary palp (female).
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
643
Figure 4. Stockmyzon mucronipes (Stock, 1960) comb. nov. SEM micrographs (female). A, protopodal segmentation of
leg 1, anterior; B, spinous process on proximal endopod segment of leg 4; C, tip of maxillipedal claw.
with one seta and one small spine; segments 11–15,
each with two setae; segment 16, with four setae;
segment 17, with three setae plus an aesthetasc; segment 18 with nine setae. Segment 10 reduced, partly
covered by distal expansion of compound segment 9
(Fig. 6E). Proximal seta on ancestral segments XVIII–
XX: rudimentary (Figs 3C, 6D).
Maxilliped (Fig. 6C): indistinctly six-segmented;
comprising short syncoxa, long basis, and indistinctly
four-segmented endopod. Syncoxa with one short seta
distally, incompletely separated from basis. Basis: with
one small tooth-like process along medial margin near
syncoxa–basis joint; with spinules along outer margin.
First endopodal segment not completely separated
from basis; with two setae and a few spinules near the
distal margin. Second endopodal segment: with one
terminal seta. Third endopodal segment: compound,
showing membranous insert marking plane of fusion
between ancestral segments 3–4; with recurved terminal claw plus short accessory apical seta.
Fifth legs (Fig. 6B): not markedly different from
those of female.
Sixth legs (Fig. 6B): represented by opercula closing
off genital apertures; each with two smooth setae.
Remarks
Comparison with Stock’s (1960) text and illustrations
revealed a number of discrepancies, which may be
attributed to imperfect dissection and/or observation:
(1) Stock (1960) described the female antennule as
19-segmented and stated that segments 18–19 were
indistinctly separated; re-examination showed that
the minute tenth segment (XIII) was overlooked by
Stock, and that the terminal segments are divided by
a clear articulation; (2) the antennary exopod has not
two, but three elements; Stock missed the lateral
exopodal seta, as well as the two smaller setae on the
distal endopod segment; (3) the mandibular palp is
not indistinctly two-segmented, as stated by Stock; (4)
Stock’s illustration of the maxillule shows four terminal setae on the praecoxal endite (the shorter one was
overlooked), and only two instead of four on the palp;
(5) the maxilla has an aesthetasc-like extension on
the proximal part of the syncoxa, which was not
illustrated by Stock [nor in any other asterocherid
descriptions prior to Ho (1984); cf. Asterocheres aesthetes Ho, 1984); (6) Stock described and illustrated
the maxillipedal endopod as distinctly threesegmented, but his segment boundaries do not coincide with the pattern we observed – his proximal
segment is a composite of the genuine first segment
and the proximal half of the middle segment (compare
Fig. 2G), whereas his middle segment corresponds to
only the distal half of that segment – this reinterpretation explains the difference between our endopodal
setal formula [2, (0,1), 1+claw] and Stock’s [1, 1,
1+claw]; (7) Stock overlooked the inner basal seta on
P1, and erroneously illustrated the outer basal seta
as plumose; (8) the epicuticular scales on the urosomited were not illustrated in Stock’s description.
Slight morphological variations occur between the
Tarifa specimens and the Roussillon population. In
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
644
M. E. BANDERA and R. HUYS
Figure 5. Stockmyzon mucronipes (Stock, 1960) comb. nov. (female). A, leg 3, anterior; B, leg 4, anterior; C, leg 2,
anterior; D, leg 1, anterior.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
645
Figure 6. Stockmyzon mucronipes (Stock, 1960) comb. nov. (male). A, habitus, dorsal; B, urosome, ventral; C,
maxilliped; D, antennule; E, detail of antennulary segments VIII, IX–XII, XIII, XIV, and XV; F, detail of antennulary
segments XX–XXII and XXIII–XXVIII.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
646
M. E. BANDERA and R. HUYS
the Roussillon population: (1) the oral cone is slightly
shorter, reaching only as far as the bases of leg 1; (2)
the proximal outer process on the middle endopod
segment of leg 1 is markedly shorter, whereas the
apical process and outer seta on the distal endopod
segment are distinctly longer; (3) the apical spine and
seta on the distal endopod segment of leg 3 are longer
than the segment (but shorter in the Tarifa population); (4) the caudal rami are slightly longer; and (5)
the body length is smaller (551–589 mm).
STOCKMYZON
CRASSUS SP. NOV.
Synonym: Asterocheres mucronipes Stock, 1960 sensu
Stock (1966).
Original description: Stock (1966: 146–147, fig. 1a–c).
Type locality: Mauritius, Chenal du Trou d’Eau
Douce; associated with “small flabby orange sponges”,
without skeleton (Oscarella sp.), in small “grottos” in
the reef at 6–10 m depth’.
Material examined: Holotype female and 16 paratype
females (originally identified as A. mucronipes)
(ZMA Co. 100.955) from type locality; collected by
J.H. Stock, 7 February 1964.
Description: Restricted to differences with the type
species.
Female: Body (Fig. 7A): cyclopiform, consisting of
dorsoventrally flattened prosome and cylindrical
urosome. Total length measured from rostral margin
to posterior margin of caudal rami, 652 mm [564–664,
N = 6 according to Stock (1966)]; maximum width,
440 mm measured at 4/5 length of cephalothorax.
Ratio of length to width of prosome: 1.08 : 1. Ratio of
length of prosome to that of urosome: 2.5 : 1. Genital
double somite and free abdominal somites: covered
with large epicuticular scales arranged in an overlapping pattern (Fig. 8E); scales larger than in S. mucronipes. Somite bearing leg 5 (Fig. 7B): wider than
long, with some spinules around bases of outer basal
setae. Genital double somite (Fig. 7B–C): narrower
and less laterally produced than in S. mucronipes,
about 1.2 times wider than long; with lateral postgenital setular tufts.
Caudal rami (Figs 7B–C, 8E): slightly longer than
wide (measured along outer margin), ornamented
with epicuticular scales all over. Caudal setae IV–V:
distinctly swollen in proximal half.
Antennule (Fig. 7D): 20-segmented, about 312-mm
long; segmental fusion pattern as in S. mucronipes.
Segments 1–8, each with two setae; segment 9, with
six setae and a small spine; segment 10, with two
setae; segment 11, with one seta and one small spine;
segments 12–17, each with two setae; segment 18,
with two setae plus an aesthetasc; segment 19,
with three setae; segment 20, with nine setae.
Segment 10(XIII), reduced, forming incomplete sclerite partly overlapped by distal expansion of compound
segment 9(IX–XII). All setae smooth.
Antenna (Fig. 7E): biramous, 186-mm long. Coxa and
basis: without spinule rows. Exopod: as in S. mucronipes. Endopod: three-segmented; proximal segment
with spinular ornamentation as illustrated; middle
segment with one smooth seta; distal segment with one
naked seta and one distal claw, and with few spinules
along margin and long setules on anterior surface.
Siphon (Fig. 8A): long and slender, reaching to
intercoxal sclerite of leg 1.
Mandible (Fig. 9A): comprising stylet-like gnathobase and slender two-segmented palp. Proximal
segment of palp unarmed; distal segment with two
plumose, unequally long, apical setae. Stylet: as in
S. mucronipes.
Maxillule (Figs 8C, 9B): bilobed. Praecoxal gnathobase four times longer than palp; ornamented with a
row of long spinules distally, and a row of shorter
spinules laterally; armed with five distal setae (one of
them smooth and short). Palp strongly reduced; with
one elongate, strongly plumose seta, and two short
pinnate setae.
Maxilla (Figs 8B, 9C): essentially as in S. mucronipes, but coxal part of syncoxa without surface
spinule row. Vestigial element on claw-like basis: not
discernible.
Maxilliped (Fig. 9D): as in S. mucronipes, but basis
and endopod relatively more slender; endopod
segments 1–2 separated by suture on anterior side
only.
Swimming legs 1–4 (Figs 8D, 10A–D): intercoxal
sclerite present in legs 1–4, ornamented with patches
of spinules in legs 1–3. Spine and seta formula: as for
S. mucronipes. Coxae ornamented with spinule rows
laterally, as illustrated. Middle and distal endopodal
segments in legs 1–4: with a beak-shaped spiniform
process distally (e.g. Fig. 8D). Leg 1 differs from that
of S. mucronipes in the following characteristics:
outer basal seta shorter; inner coxal seta pinnate
instead of bare; proximal outer spinous process on
middle endopod segment shorter; outer seta of distal
endopod segment extending just beyond distal
spinous process. Legs-2–3 inner coxal seta and outer
basal seta: much shorter than in S. mucronipes.
Proximal inner seta of middle endopod segment of
leg 4: much longer than in S. mucronipes and
approaching the length of the distal inner seta.
Fifth leg (Fig. 7B): as in S. mucronipes except for
lateral exopodal seta being distinctly shorter. Sixth
legs (Fig. 7B): represented by paired opercular plates
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
647
Figure 7. Stockmyzon crassus (Stock, 1966) sp. nov. (female). A, habitus, dorsal; B, urosome, dorsal; C, urosome
(excluding leg-5-bearing somite), ventral; D, antennule, ventral; E, antenna.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
648
M. E. BANDERA and R. HUYS
Figure 8. Stockmyzon crassus (Stock, 1966) sp. nov. SEM micrographs (female). A, oral cone; B, maxilla; C, maxillulary palp; D, P4 endopod, anterior; E, surface scales on urosomites, ventral.
closing off gonopores on genital double somite; armed
with one short, smooth seta, and one spiniform
element.
Male: Unknown.
Etymology: The specific name is derived from the
Latin crassus, meaning thick, and refers to the
swollen caudal setae.
Remarks
Stock’s (1966) redescription of A. mucronipes from
Mauritian sponges is concise and limited to illustrations of the urosome, leg 4, and the maxillule. Stock
confirmed several similarities with the Mediterranean type population, such as the annulated struc-
ture of the mandibular stylet and the presence of
beak-shaped processes on the swimming legs;
however, he also claimed that the maxillule differed
slightly in the shape of the ‘outer ramus’ or palp,
being gonflate basally, and distinctly narrower distally (Stock (1966: fig. 1b). This could not be confirmed in the Mauritian material or in the types of
S. mucronipes (Stock doubted his original observation) (Figs 2D, 9B); instead, our re-examination
revealed that Stock (1966) had overlooked a seta on
both the maxillulary endite and palp. His illustration
of leg 4 also shows minor discrepancies with our
Figure 10D, such as proportional length differences in
the outer seta of the proximal exopodal segment and
the inner proximal seta of the middle endopodal
segment, and the apparent slenderness of the outer
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
649
Figure 9. Stockmyzon crassus (Stock, 1966) sp. nov. (female). A, mandible; B, maxillule; C, maxilla; D, maxilliped,
posterior.
exopodal spines (as a result of omitting the membranous flanges). Stock (1966) illustrated the swollen
caudal ramus setae IV–V, but did not remark on this
character in the text.
DISCUSSION
SPECIES
DISCRIMINATION
Stockmyzon mucronipes and S. crassus are morphologically very similar in most aspects, but the latter
can be distinguished from the former by the following
suite of characters: (1) epicuticular scales on genital
double somite and free abdominal somites, larger; (2)
genital double somite, narrower and less laterally
produced (width : length ratio 1.2 vs. 1.7 in
S. mucronipes); (3) caudal ramus setae IV–V distinctly swollen in proximal half; (4) all antennulary
setae smooth; (5) siphon slightly shorter, reaching to
intercoxal sclerite of leg 1; (6) maxillulary palp with
one elongate, strongly plumose seta, and only two
short pinnate setae; (7) coxal part of maxillary
syncoxa without surface spinule row; (8) maxilliped
basis and endopod relatively more slender; (9) leg-1
outer basal seta shorter, and inner coxal seta pinnate
instead of bare; proximal outer spinous process on
middle endopod segment, shorter; outer seta of distal
endopod segment extending just beyond distal
spinous process; (10) legs-2–3 inner coxal seta and
outer basal seta, much shorter; (11) proximal inner
seta of middle endopod segment of leg 4, much longer,
approaching the length of the distal inner seta; and
(12) leg-5 lateral exopodal seta, distinctly shorter.
TAXONOMIC
ASTEROCHERES
GIESBRECHT, 1897
POSITION OF
STIMULANS
Giesbrecht (1897) named three new species of Asterocheres, all of which were collected in the Gulf of
Naples: Asterocheres dentatus, Asterocheres parvus,
and A. stimulans. Although no illustrations or formal
diagnoses were given, Giesbrecht did cite the new
species in his identification key. Such a citation can be
considered the equivalent of a differential diagnosis,
and is sufficient to make the new species names
available. Illustrated descriptions were given in a
subsequent report on the Asterocheridae of the Gulf of
Naples (Giesbrecht, 1899). Stock (1960) recognized
the similarity in swimming leg morphology between
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
650
M. E. BANDERA and R. HUYS
Figure 10. Stockmyzon crassus (Stock, 1966) sp. nov. (female). A, leg 1, anterior; B, leg 2, anterior; C, leg 3, anterior;
D, leg 4, anterior.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
A. mucronipes and the male of A. stimulans, in particular the beak-shaped processes on the endopods.
Examination of Giesbrecht’s detailed illustrations of
both sexes of A. stimulans casts severe doubts on
their conspecificity. His figures of the male include the
habitus, antennule, maxilliped (note the position of
the spinous process on the basis), leg 1, and endopods
of legs 2–4 (note the short proximal inner seta on the
middle endopod segment of leg 4), all of which
conform exactly to S. mucronipes. The only exception
is the illustration of the mouth cone area, which
shows a slightly longer siphon (extending to the intercoxal sclerite of leg 3), and only three small setae on
the maxillulary palp. The first difference is probably
caused by excessive squashing of the specimen, which
results in a slightly posterior displacement of the
siphon in the foreshortened in situ view (also note the
distorted position of the mandibular palp). The significant aspect in the second difference is the absence
of the large, gonflate seta. This seta is typically medially directed, and could easily be concealed by the
praecoxal gnathobase and oral cone in a squashed
preparation (compare Figs 2D and 3E). The palp
(even though Giesbrecht’s illustration is small) shows
the atrophied facies that is characteristic for Stockmyzon. The description of the female of A. stimulans
is concise, and includes figures of the habitus, antennule, maxillule, maxilliped, and urosome. The antennule is 20-segmented, as in S. mucronipes, but
the segmental homologies are different. In female
A. stimulans there are three segments distal to the
aesthetasc-bearing segment XXI, and the vestigial
segment XIII represents the ninth segment; in S. mucronipes only two segments are expressed distal to the
ancestral segment XXI, and segment XIII is homologous with the tenth segment. The maxillule bears no
resemblance to that in S. mucronipes, being similar to
the typical Asterocheres condition. The female maxilliped is atypical in that it differs significantly from
the male in its general slenderness and the length of
the endopodal claw; such sexual dimorphism is
extremely rare among asterocherids and makes the
conspecificity of the two sexes highly questionable. We
strongly believe that Giesbrecht (1899) based his
description of A. stimulans on an amalgam of two
different species. No holotype was designated by Giesbrecht and the original type series no longer exists
(Kölmel, 1980; confirmed during a visit to the Stazione Zoologica in Naples by RH, October 2003). In
order to preserve the stability of nomenclature, we
designate the female specimen illustrated by Giesbrecht (1899: plate 3; figs 1, 3, 6, 7, 12, 14) (as the
lectotype of A. stimulans (ICZN Art. 74.4). The male
illustrated in Giesbrecht (1899: figs 2, 4, 5, 8–11, 13)
is considered here as conspecific with S. mucronipes.
Although its host was unknown to Giesbrecht (1897,
651
1899), we suspect that it was A. calycularis; Giesbrecht’s material was collected in the vicinity of Naples,
which virtually coincides with the northernmost limit
of distribution of this host in Italy (Zibrowius, 1995).
The identity of Canu’s (1898) record of A. stimulans
from algal washings in Maisy (Normandy coast)
requires confirmation. The only female collected
shows a 21-segmented antennule, but the similarity
in siphon size, shape of the genital double somite, and
length of the caudal ramus indicates that conspecificity cannot be ruled out.
HOST
SWITCHING IN THE NORTH-WESTERN
MEDITERRANEAN?
Astroides calycularis is an azooxanthellate dendrophylliid colonial coral, typically inhabiting shallow
waters down to a depth of about 30 m, and preferring
shaded places and strong water movement (Zibrowius, 1980, 1995). It is protected by the Convention on
International Trade in Endangered Species of Wild
Fauna and Flora (CITES), and since 1999 has been
listed by the Spanish Government as a vulnerable
species in the National Catalogue of Endangered
Species (Catálogo Nacional de Especies Amenazadas)
(Anonymous, 1999). Astroides calycularis is essentially endemic to the south-western Mediterranean,
with a few outliers beyond the Straits of Gibraltar in
the west and the Straits of Sicily in the east. Its
distribution is presently limited at 37°38′N (Cape
Palos) on the coast of Spain and 40°48′N (Gulf of
Gaeta) on the coast of Italy (Zibrowius, 1980, 1983).
More recent research based on fossil evidence has
demonstrated that A. calycularis lived on the mediterranean coast of France at 43°42′-43°44′N during
part of the Pleistocene, taking advantage of the
slightly higher surface water temperatures than
those prevailing now in the northern Mediterranean
(Zibrowius, 1995). Field experiments with colonies
transplanted from Italy showed that present-day temperatures allow short-term survival, but fail to
sustain successful reproduction. Stock (1960) found
S. mucronipes in washings of the gorgonian E. singularis in the Banyuls-sur-Mer area, which is south of
the northernmost limit of distribution of A. calycularis during the Upper Sicilian and Upper Tyrrhenian
(interstages of the Riss and Würm glaciations). We
postulate that the symbiotic association between
A. calycularis and S. mucronipes was already established before the Pleistocene and in the entire former
distribution range of the host, i.e. the western mediterranean basin. When climatic conditions changed
during the late Sicilian regression (Rissian age),
about 238 000–225 000 years ago, the drop in temperature caused the extinction of A. calycularis along
the French Mediterranean coast, and the northern
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
652
M. E. BANDERA and R. HUYS
coastal waters of Spain and Italy, but did not necessarily wipe out the symbionts that depended on it.
Switching to suitable hosts that reside in deeper
waters, and are less susceptible to ambient temperature changes, offers a solution for symbionts that are
at the risk of extinction. It appears that S. mucronipes maintained its presence in the north-western
Mediterranean by switching to alternative hosts, such
as the gorgonian E. singularis; however, the authenticity of Stock’s (1960) record requires confirmation by
additional sampling over a wider geographical scale
before this hypothesis can be supported. Stock (1966)
himself failed to collect additional S. mucronipes
specimens from E. singularis in subsequent years,
and therefore suggested the real host may well be a
sponge, as in the case of S. crassus.
ACKNOWLEDGEMENTS
EB acknowledges financial support for a visit to
the Natural History Museum (London) under the
EC-funded TMR Bioresource Large-Scale Facility
Programme. Special thanks are due to Dr Mercedes
Conradi for assistance throughout the course of this
study and to Dr Platvoet, curator at the Zoological
Museum of Amsterdam, for arranging the loan of
Stock’s material. We would also like to thank Isabel
Alfonso for logistical support in collecting the samples
of Astroides calycularis at Tarifa.
REFERENCES
Anonymous. 1999. Ministerio de Medio Ambiente. 13807.
Orden de 9 de junio de 1999 por la que se incluyen en el
Catálogo Nacional de Especies Amenazadas determinadas
especies de cetáceos, de invertebrados marinos y de flora y
por la que otras especies se excluyen o cambian de categoría.
Boletín Oficial del Estado 148: 23921–23922.
Bandera ME, Conradi ME, López-González PJ. 2005.
Asterocheres hirsutus, a new species of parasitic copepod
(Siphonostomatoida: Asterocheridae) associated with an
Antarctic hexactinellid sponge. Helgoland Marine Research
59: 315–322.
Bispo R, Johnsson R, Neves E. 2006. A new species
of Asterocheres (Copepoda, Siphonostomatoida, Asterocheridae) associated to Placospongia cristata Boury-Esnault
(Porifera) in Bahia State, Brazil. Zootaxa 1351: 23–34.
Boxshall GA. 1990. Siphonostome copepods associated with
sponges from Hong Kong. In: Morton BS, ed. Proceedings of
the Second International Marine Biological Workshop. The
Marine Flora and Fauna of Hong Kong and Southern
China. Hong Kong: Hong Kong University Press, 523–547.
Boxshall GA, Halsey HS. 2004. An introduction to copepod
diversity. London: The Ray Society.
Boxshall GA, Huys R. 1994. Asterocheres reginae, a new
species of parasitic copepod (Siphonostomatoida: Astero-
cheridae) from a sponge in Belize. Systematic Parasitology
27: 19–33.
Canu E. 1892. Les Copépodes du Boulonnais. Morphologie,
embryologie, taxonomie. Travaux de l’Institut Zoologique de
Lille et de la Station Maritime de Wimereux 6: 1–292.
Canu E. 1898. Note sur les Copépodes et les Ostracodes
marins des côtes de Normandie. In: de Kerville H. Gadeau,
ed. Recherches sur les faunes marine et maritime de la
Normandie, 2e voyage. Bulletin de la Société des Amis des
Sciences naturelles de Rouen 33: 389–422.
Conradi ME, Bandera E, López-González PJ. 2006. The
copepods associated with the coral Astroides calycularis
(Scleractinia, Dendrophylliidae) in the Strait of Gibraltar.
Journal of Natural History 40: 739–757.
Giesbrecht W. 1897. System der Ascomyzontiden, einer semiparasitischen Copepoden-Familie. Zoologischer Anzeiger 20:
9–14.
Giesbrecht W. 1899. Die Asterocheriden des Golfes von
Neapel und der angrenzenden Meeres-Abschnitte. Fauna
und Flora des Golfes von Neapel 25: i–vi, 1–217.
Gotto RV. 1979. The association of copepods with marine
invertebrates. Advances in Marine Biology 16: 1–109.
Ho J-s. 1982. Copepoda associated with echinoderms of the
Sea of Japan. Report of the Sado Marine Biological Station,
Niigata University 12: 33–61.
Ho J-s. 1984. Copepoda associated with sponges, cnidarians
and tunicates of the Sea of Japan. Report of the Sado
Marine Biological Station, Niigata University 14: 23–62.
Humes AG. 1993. Copepoda associated with gorgonaceans
(Cnidaria) in the Indo-Pacific. Bulletin of Marine Science 53:
1078–1098.
Humes AG. 1994. How many copepods?. In: Ferrari FD,
Bradley BP, eds. Ecology and Morphology of Copepods.
Proceedings of the Fifth International Conference on
Copepoda. Hydrobiologia 292/293: 1–7.
Humes AG. 1996. Siphonostomatoid copepods (Asterocheridae) associated with the sponge Dysidea in the
Moluccas. Systematic Parasitology 35: 157–177.
Humes AG. 1997. Copepoda (Siphonostomatoida) associated
with the fungiid coral Parahalomitra in the southwestern
Pacific. Journal of Natural History 31: 57–68.
Humes AG, Stock JH. 1991. Coralliomyzontidae, fam. n.
(Copepoda: Siphonostomatoida), associated with scleractinian corals in Madagascar. Bulletin Zoölogisch Museum, Universiteit van Amsterdam 13: 17–24.
Huys R, Boxshall GA. 1991. Copepod evolution. London: The
Ray Society.
Ivanenko VN. 1997. Redescription of Asterocheres simulans
(Copepoda, Siphonostomatoida, Asterocheridae) – a symbiont of Suberites domuncula ficus (Spongia) from the White
Sea. Comments on the taxonomy and ecology. Zoologicheskii
Zhurnal 76: 1118–1130.
Ivanenko VN. 1998. Laperocheres koorius, a new genus and
species (Copepoda: Siphonostomatoida: Asterocheridae),
associated with the sponge Amphimedon in Australia. Proceedings of the Biological Society of Washington 111: 263–271.
Ivanenko VN, Ferrari FD. 2003. Redescription of adults
and description of copepodid development of Dermatomyzon
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653
NEW GENUS OF ASTEROCHERIDAE
nigripes (Brady & Robertson, 1876) and of Asterocheres
lilljeborgi Boeck, 1859 (Copepoda: Siphonostomatoida:
Asterocheridae). Proceedings of the Biological Society of
Washington 116: 661–691.
Ivanenko VN, Smurov AV. 1997. Asterocheres flustrae n. sp.
(Copepoda: Siphonostomatoida: Asterocheridae) associated
with Flustra foliacea L. (Bryozoa) from the White Sea.
Systematic Parasitology 38: 111–130.
Johnsson R. 1998. Six new species of the genus Asterocheres
(Copepoda; Siphonostomatoida) associated with sponges in
Brazil. Nauplius 6: 61–99.
Kim I-H. 1998. Pulicitrogus compressus gen. nov., sp. nov.
(Copepoda, Siphonostomatoida, Artotrogidae) associated
with an ascidian in the Sea of Japan. Journal of Marine
Systems 15: 255–260.
Kim I-H. 2004a. New species of copepods (Crustacea) associated with marine invertebrates from the Pacific coast of
Panama. Korean Journal of Biological Sciences 8: 165–
186.
Kim I-H. 2004b. Two new species of siphonostomatoid copepods (Crustacea) associated with the stoloniferan coral
Tubipora musica (Linnaeus) from Madagascar. Korean
Journal of Biological Sciences 8: 187–196.
Kim I-H. 2005. Two new species of copepods (Crustacea)
associated with the sponge Phyllospongia foliascens (Pallas)
from the Moluccas. Korean Journal of Biological Sciences 9:
229–238.
Kim I-H, Je J-G. 2000. Two new species of Asterocheridae
653
(Copepoda, Siphonostomatoida) from Korea. Korean Journal
of Biological Sciences 4: 305–314.
Kölmel R. 1980. Provisional inventory of the dissected slide
material of Wilhelm Giesbrecht held at the Stazione
Zoologica in Naples. Zoologisches Museum der ChristianAlbrechts-Universität zu Kiel. Arbeitsblätter 1: i + 1–21.
Stock JH. 1960. Sur quelques Copépodes associés aux
invertébrés des côtes du Roussillon. Crustaceana 1: 218–257.
Stock JH. 1966. Cyclopoida siphonostoma from Mauritius
(Crustacea, Copepoda). Beaufortia 13: 145–194.
Weinberg S. 1976. Revision of the common Octocorallia of the
Mediterranean circalittoral. I. Gorgonacea. Beaufortia 24:
63–104.
Weinberg S. 1978. Revision of the common Octocorallia of the
Mediterranean circalittoral. III. Stolonifera. Beaufortia 27:
139–176.
Zibrowius H. 1980. Les Scléractiniaires de la Méditerranée
et de l’Atlantique nord-oriental. Mémoires de l’Institut
Océanographique, Monaco 11: 1–284.
Zibrowius H. 1983. Nouvelles données sur la distribution
de quelques scléractiniaires ‘méditerranéens’ à l’Est et à
l’Ouest du détroit de Gibraltar. Rapports et Procès-Verbaux
des Réunions – Commission internationale pour l’Exploration
scientifique de la Mer Méditerranée 28: 307–309.
Zibrowius H. 1995. The ‘southern’ Astroides calycularis in
the Pleistocene of the northern Mediterranean – an indicator of climatic changes (Cnidaria, Scleractinia). Geobios 28:
9–16.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 635–653