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Symbiotic polychaetes from Nhatrang Bay, Vietnam

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Abstract

Shallow-water polychaete fauna associated with benthic invertebrates in Nhatrang Bay, Vietnam was studied based on extensive sampling surveys performed from 1985 to 2012. As a result, 24 polychaete species belonging to four families were found. A check list of all these species including synonymies, list of material examined, extended diagnosis, taxonomic and ecological notes, and distribution is here presented. Most reports include in situ colour photos of representative specimens. The vast majority of symbionts belong to Polynoidae (13 species), followed by Syllidae (5 species), Hesionidae (5 species), and Spionidae (1 species), which were associated with echinoderms (53.8%), cnidarians (23.0%), sponges (11.5%), crustaceans (7.7%), and other polychaetes (3.8%). The number of symbiotic polychaetes occurring in Vietnamese coastal waters is here increased from 10 up to 26 species. The Vietnamese fauna of symbiotic polychaetes differs considerably from the Australian one. However, further studies on poorly investigated or even overlooked hosts (e.g. sponges, octocorals, tube dwelling polychaetes, echinoids and ophiuroids) will certainly lead to an increase in our knowledge on the diversity of symbiotic polychaetes of Vietnam.
Benthic fauna of the Bay of Nhatrang, Southern Vietnam. Vol. 2. Moscow: KMK Scien-
tic Press Ltd. 2012. 491 p.
The book contains 9 chapters describing different groups of marine invertebrates: symbiotic
polychaetes; mollusks – chitons, gastropods of the family Eulimidae, nudibranchs; pontoniin
shrimps; commercial and mangrove species of crabs; sea stars and crinoids. In the result of the
processing of the samples, collected in the Bay of Nhatrang, 474 species of invertebrates are
described, nearly half of them (218 species) were found in Vietnam for the rst time. Eight new
for science species are described. For each species the localities and general distribution data,
synonymy and for many species diagnoses and descriptions are provided. The book is illus-
trated by 62 line drawings text gures and 70 plates with 502 original color photographs of live
specimens taken in nature or in aquaria immediately after collecting.This volume signicantly
increases the knowledge not only about the fauna of the Bay, but of Vietnam in general.
Ed i to rs -i n -c hi ef : Prof. T.A. Britayev, Acad. D.S. Pavlov
Edit o r s: Yu.I. Kantor, T.I. Antokhina
List of reviewers
Dr. Z. Duris, Prof., Department of Biology University of Ostrava Chittussiho, Ostrava Czech Re-
public.
Dr. E. González, El Colegio de la Frontera Sur, Departamento de Ecología Acuática, Chetumal,
Mexico.
Dr. D.J.W. Lane, Ass. Prof., Department of Biology, University Brunei Darussalam, Brunei Darus-
salam.
Dr. D. Martin, Centre d’Estudis Avançats de Blanes (CEAB–CSIC), Blanes (Girona), Catalunya,
Spain.
Dr. C. Messing, Prof., Nova Southeastern University Oceanographic Center, Dania, USA.
Dr. R. Naderloo, Research Institute & Nature Museum of Senckenberg, Germany.
Dr. V. Neumann, Research Institute & Nature Museum of Senckenberg, Germany.
Dr. H. Saito, National Museum of Nature And Science, Tokyo, Japan.
Dr. B.I. Sirenko, Prof., Institute of Zoology, Russian Academy of Sciences, St.-Petersburg, Russia.
Recommended citation:
The whole volume – Britayev T.A., Pavlov D.S. (Eds.). 2012. Benthic fauna of the Bay of
Nhatrang, Southern Vietnam. Vol. 2. Moscow, KMK, 491 pp.
The chapter – Britayev T.A., Antokhina T.I. 2012. Symbiotic polychaetes from Nhatrang
Bay, Vietnam. In Britayev T.A., Pavlov D.S. (Eds.). Benthic fauna of the Bay of Nhatrang,
Southern Vietnam. Vol. 2. Moscow, KMK: 11–54.
ISBN 978-5-87317-860-5
11
CHAPTER 1
Symbiotic polychaetes from Nhatrang Bay, Vietnam
T.A. Britayev1, T.I. Antokhina1
ABSTRACT. Shallow-water polychaete fauna associated with benthic invertebrates in Nha-
trang Bay, Vietnam was studied based on extensive sampling surveys performed from 1985
to 2012. As a result, 24 polychaete species belonging to four families were found. A check
list of all these species including synonymies, list of material examined, extended diagnosis,
taxonomic and ecological notes, and distribution is here presented. Most reports include in situ
colour photos of representative specimens. The vast majority of symbionts belong to Poly-
noidae (13 species), followed by Syllidae (5 species), Hesionidae (5 species), and Spionidae
(1 species), which were associated with echinoderms (53.8%), cnidarians (23.0%), sponges
(11.5%), crustaceans (7.7%), and other polychaetes (3.8%). The number of symbiotic poly-
chaetes occurring in Vietnamese coastal waters is here increased from 10 up to 26 species. The
Vietnamese fauna of symbiotic polychaetes differs considerably from the Australian one. How-
ever, further studies on poorly investigated or even overlooked hosts (e.g. sponges, octocorals,
tube dwelling polychaetes, echinoids and ophiuroids) will certainly lead to an increase in our
knowledge on the diversity of symbiotic polychaetes of Vietnam.
The establishment of specialized symbiotic associations involving polychaetes is a
rather common phenomenon in the World Oceans. According to the most recent review
[Martin, Britayev 1998] there are ca. 370 symbiotic species of polychaetes, either ob-
ligate or facultative. Although their world distribution patterns have not been formally
analyzed, it seems likely that there may be a latitudinal gradient, with a relatively low
diversity in cold and temperate waters that tends to increase towards tropical areas. Con-
versely, the symbiotic polychaete fauna is relatively well known in the cold and temper-
ate European waters, as well as along both coasts of North America [Barel, Kramers
1977; Pettibone 1993], while tropical areas are still poorly investigated. There are no
publications focusing on these organisms from any particular tropical area, with a few
exceptions [e.g. Gibbs 1969; Britayev et al. 1999]. In turn, most data on symbiotic spe-
cies are usually scattered in faunistic, taxonomic, or, more rarely, in ecological papers
[e.g. Devaney 1967; Hanley 1989; Britayev, Zamyschliak 1996; Barnich et al. 2004],
and the Vietnamese fauna is not an exception.
Various polychaete families have been previously studied in Vietnam [e.g. Fauvel
1939; Uschakov 1972; Fauchald 1968; Gallardo 1967; Strelzov 1968, 1972]. However,
only ten symbiotic species are known to date. Most of them belong to the Polynoidae
(8 species), while the Syllidae and the Spionidae were represented by only one species,
respectively [Fauvel 1939; Gallardo 1967; Radashevsky 1996; Britayev, Zamyschliak
1996; Britayev, Fauchald 2005; Britayev, Martin 2005; Martin et al. 2009; Antokhina,
1 A.N. Severtsov Institute of Ecology and Evolution, 33 Leninskij prospect, Moscow, Russia,
119071, britayev@yandex.ru
12
chapter 1
Britayev 2012]. Just to compare, a single paper on the Solomon Islands polychaete ecol-
ogy reported twelve symbiotic polychaetes [Gibbs 1969].
The main object of this paper is, thus, present an extended summary of the results
of our studies on the symbiotic polychaete fauna from Nhatrang Bay, based on the ex-
tensive sampling surveys performed from 1985 to 2008. A check list of all these species
including synonymies, list of material examined, extended diagnosis, taxonomic and
ecological notes, and distribution is here presented. Most reports include in situ colour
photos of representative specimens. In addition, we have also included all available pub-
lished records on symbiotic polychaetes found in Vietnamese coastal waters.
Material and methods
All specimens were collected in Nhatrang Bay (Vietnam) during a series of short-term
expeditions carried out from 1985 to 1990 and from 2002 to 2008, which includes an
additional survey at Phu Quoc Island in 2008 (Figs. 1, 2). All hosts (viz. sponges, sclerac-
tinian corals, octocorals, polychaetes, decapods, starshes, crinoids, sea urchins) were col-
lected by hand either at the intertidal area or by scuba-diving to depths of 35 m. Each host
specimen was placed in a zip-lock plastic pocket while still underwater, to keep the pos-
sible symbionts together with its host. On board, the presence of symbionts was carefully
checked by eye, and all large, noticeable polychaetes were collected by pincers, relaxed
in an isotonic magnesium chloride solution, and xed. The additional symbionts were ex-
tracted by gently washing the hosts with a 10% alcohol/marine water solution through
a 100 μm mesh. The species associated with the starshes Linckia laevigata (Linnaeus,
1758), Protoreaster nodosus (Linnaeus, 1758) and, especially, Culcita novaeguineae Mül-
ler et Troschel, 1842 may be deeply hidden in the ambulacral grooves. In these cases,
each starsh was placed into a closed plastic bag for 30–40 minutes to reduce the oxygen
concentration, which forced the symbionts to leave their shelter making them accessible
for collecting. The symbionts were preserved in 70% alcohol.
Photos of living animals were usually made, either in native conditions or in an
aquarium, with Nikon D300, Nikon D700 or Sea&Sea DX–1g cameras. The preserved
specimens were photographed under a Leika MZ6 stereomicroscope linked to a Leika
DFC295 digital camera. Scanning electron Microscope (SEM) micrographs were made
after critical point drying and coating with 300 Å of gold, at the Laboratory of Electronic
Microscopy, A.N. Severtzov Institute of Ecology and Evolution of Russian Academy of
Sciences (SIEE RAS). The specimens are deposited in the Laboratory of Ecology and
Morphology of Marine Invertebrates of the SIEE RAS.
The references listed in the synonymies are restricted to the original descriptions, as
well as to relevant reviews or major faunistic works, which were used for comparison
and identication. Only rare species include a full synonymies list. In “Specimens ex-
amined”, all records are listed in chronological order. In “Measurements”, the following
abbreviations are used: cs complete specimen, af anterior fragment, mf median
fragment, pf – posterior fragment. Length of worms (L) was measured from the frontal
prostomial margin to the posterior pygidial end. Width was measured ventrally at the
widest body section (usually around chaetigers 8 to 15), without parapodia (WW), with
13
t.a. britayev, t.i. antokhina
parapodia, but without chaetae (WP), and with parapodia and chaetae (WC), except for
Gastrolepidia clavigera Schmarda, 1861 whose ventral lamellae hided the ventral bases
of parapodia. In this case, WW was measured dorsally. We consider WW as the most
consistent measurement, but we have included the other two just to allow comparisons
with previously published data
The diversity of symbiotic polychaetes of Vietnam has been compared with that of the
coastal waters of Australia, the number of Australian species was estimated on the basis of
the modern faunistic and taxonomic papers included references on symbiotic polychaetes
[Blake, Kudenov 1978; Hanley 1984, 1989; Hanley, Burke 1989, 1990; Pettibone 1989;
Glasby, Watson 2001; Glasby et al. 2012; Lattig et al. 2010]. Australian fauna was chosen
for comparison as predominately tropical and relatively well studied one.
Results
Family POLYNOIDAE Kinberg, 1856
Asterophilia culcitae Britayev et Fauchald, 2005
(Pl. 1 A–E)
Asterophilia culcitae Britayev, Fauchald 2005: 17, gs. 2– 4.
Material examined. Nhatrang Bay: Tre Island, 4 m, on Culcita novaeguineae,
coll. Britayev, 23.04.2004 2 cs; Mieu Island, 5 m, on C. novaeguineae, coll. Marin,
14.05.2004 2 cs; 11 m, on Echinaster luzonicus, coll. Savinkin, 24.06.2004 1 cs;
7 m, on Choriaster granulatus, coll. Savinkin, 31.10.2005 – 1 cs; Mot Island, 7 m, on C.
novaeguineae, coll. Antokhina, 18.04.2006 – 1 cs and 1 af; 5 m, on Linckia laevigata,
coll. Antokhina, 16.04.2008 2 cs; Dung Island, 13 m, on Protoreaster nodosus, coll.
Antokhina, 26.04.2006 – 1 cs; 9 m, on Euretaster insignis, coll. Antokhina, 10.05.2006 –
1 cs; Mun Island, 5 m, on Fromia monilis, coll. Antokhina, 03.11.2007 – 1 cs.
Diagnosis. Body attened, with up to 48 segments. Two dorsal transverse ciliary
bands on each segment. Fifteen pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17,
19, 21, 23, 26, 29, 32. Posterior segments cirrigerous, without elytra. Elytra with rounded
macrotubercles, distinct frontal pockets, and micropopillae. Prostomium with rounded
cephalic lobes; lateral antennae inserted terminoventrally. Antennae, tentacular and dor-
sal cirri subdistally inated, with liform tips. Two pairs of eyes, anterior one at widest
part of prostomium, posterior one near hind margin of prostomium. Notopodia small,
with conical acicular, digitiform lobes. Neuropodia with long subtriangular prechaetal
lobes and shorter postchaetal lobes with rounded tips. Notochaetae very few, with short
marginal serration and notched tips. Middle and lower neurochaetae with serrated edge
and without semilunar pocket.
Measurements. With up to 48 chaetigers, L up to 23 mm, WW up to 2.3 mm, WP –
6.3 mm, WC – 2.2 mm.
Coloration. Varying in accordance of host color pattern: white-pink (Pl. 1 E), brown,
red (Pl. 1 A–D) or transparent (for juveniles or males, Pl. 1 B), pale on tips of dorsal cirri
and pockets of elytra.
14
chapter 1
Taxonomic notes. Asterophylia culcitae differs from Asterophylia carlae Hanley,
1989 in the presence of distinct frontal pockets and micropapillae on elytra, and in hav-
ing neurochaetae with serrated edge and without semilunar pocket.
Ecological notes. Found at 4–13 m on the starshes C. novaeguineae, Choriaster
granulatus Lütken, 1869, Echinaster luzonicus (Gray, 1840), Euretaster insignis (Slad-
en, 1882), Fromia monilis (Perrier, 1869), Linckia laevigata, Protoreaster nodosus
(Echinodermata: Asteroidea). Recorded previously as symbiont of starshes (viz. C. no-
vaeguineae, Linckia laevigata, P. nodosus) and unstalked crinoids [Britayev, Fauchald
2005]. The starshes C. granulatus, E. luzonicus, E. insignis and F. monilis are reported
as new hosts for A. culcitae.
Distribution in Nhatrang Bay. Tre, Mieu, Tam, Mot, Mun, Dung Islands.
General distribution. Vietnam, Nhatrang Bay.
Australaugeneria michaelseni Pettibone, 1969
Australaugeneria michaelseni Pettibone 1969: 22–25, gs. 11, 12.
Material examined. Nhatrang Bay: Mot Island, 10–12 m, on Dendronephthya
sp. (D4), coll. Savinkin, 07.06.2006 af (juv.); Nok Island, 15–17 m, boulders, on
Dendronephthya sp. (No 6), coll. Britayev, 02.05.2007 – af and pf, male; Mun Island,
10–26 m, on Dendronephthya sp. (998/12), coll. Savinkin, 27.10.2008 af, mf, pf,
female.
Diagnosis. Body attened, tapering posteriorly, with up to 37 segments. Fifteen
pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32. Pos-
terior segments cirrigerous without elytra. Elytra large, soft, translucent, smooth.
Prostomium with rounded cephalic lobes, no eyes visible; lateral antennae inserted
terminoventrally. Dorsal cirri with long, cylindrical cirrophores and very long styles.
Notopodia with inated basal lobes and digitiform acicular lobes. Neuropodia long,
with subequal prechaetal and postchaetal rounded lobes. Prechaetal neuropodial
lobes of segments 2 and 3 enlarged, hoodlike. Notochaetae as stout as or stouter
than stoutest neurochaetae, smooth, with entire tips. Neurochaetae few in number, of
two types: upper ones slender, with long spinuos region and blunt tips; lower ones,
stout with short spinuous region and hooked tips. Neuropodial hooks in segments 2,
3 and 4.
Measurements. With up to 37 chaetigers, L up to 8.7 mm, WW up to 1.0 mm,
WP – 2.7 mm; juvenile af, 16 chaetigers, L – 3 mm, WW – 0.5 mm, WP 1.3 mm,
WC – 1.4 mm.
Coloration. Unknown for living animals. Whitish semitransparent in alcohol.
Taxonomic notes. Our specimens differ from description of Pettibone [1969] in the
presence of subterminal tooth in lower neurochaetae. Australaugeneria michaelseni dif-
fers from Australaugeneria rutilans (Grube, 1878) in having a slightly stouter body, in
notochaetae as stout as or stouter than stoutest neurochaetae, smooth, with entire tips,
instead of notochaetae slender than the stoutest neurochaetae, dentate along convex bor-
der, with bid tips.
15
t.a. britayev, t.i. antokhina
Ecological notes. Found at 10–26 m on Dendronephthya sp. (Octocorallia: Alcyo-
nacea).
Distribution in Nhatrang Bay. Nok and Mun Islands.
General distribution. The species and genus represent a new record for the Viet-
namese coast. Known also from South-West Australia [Pettibone 1969].
Australaugeneria rutilans (Grube, 1878)
(Pl. 1 F)
Polynoe rutilans Grube 1878: 37, pl. 2, g. 5.
Australaugeneria rutilans: Pettibone 1969: 22, g. 10; Wehe 2006: 56 –58, g. 5.
Material examined. Nhatrang Bay: Mot Island, 10–12 m, on Dendronephthya sp.
(D8), coll. Savinkin, 07.06.2006 – 1 adult cs, 2 juv.: cs and af; on Dendronephthya sp.
(No 4) – 1 adult cs; on Dendronephthya sp. (No 2) – 1 adult cs; on Dendronephthya sp.
(No 5) – 4 juv., cs; on Dendronephthya sp. (No 6) – 3 adults, cs and 2 af; on Dendrone-
phthya sp. (No 7) – 2 adults, af; Nok Island, 15–17 m, boulders, on Dendronephthya sp.
(No 1), coll. Britayev, 02.05.2007 – 1 cs.
Diagnosis. Body attened, tapering posteriorly, with up to 38 segments. Fifteen pairs
of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32. Elytra large,
soft, translucent, smooth. Posterior segments cirrigerous without elytra. Prostomial lobes
rounded without cephalic peaks, two pairs of small pointed eyes; lateral antennae in-
serted terminoventrally. Dorsal cirri with long, cylindrical cirrophores and very long
styles. Notopodia with short conical lobes. Neuropodia long with subequal prechaetal
and postchaetal rounded lobes. Prechaetal neuropodial lobes of segments 2 and 3 en-
larged, hoodlike. Notochaetae more slender than stoutest neurochaetae, dentate along
convex border, with bid tips. Neurochaetae few in number, of two types: upper ones
slender, with long spinuous region and blunt tips; lower ones, stout with short spinuous
region and hooked tips, in some cases with minute subdistal tooth. Neuropodial hooks in
segments 2, 3 and 4, while in juvenile in segments 2 and 3 only.
Measurements. With up to 38 chaetigers, L up to 10.5 mm, WW up to 0.9 mm,
WP –2.3 mm, WC 2.8 mm; juveniles 9–21 chaetigers, L 0.8–2.3 mm, WW 0.2–
0.3 mm, WP – 0.5–0.9 mm.
Coloration. Semitransparent, with a reddish double longitudinal band along mid–
dorsum, reddish stripes at the dorsal bases of parapodia, and a reddish band along the
antero-lateral margin of elytra (Pl. 1 F).
Taxonomic notes. The juvenile has modied hooked chaetae only in segments 2 and 3.
Ecological notes. Found at 10 –17 m on alcyonarians Dendronephthya spp. (Octo-
corallia: Alcyonacea). Previously reported in association with the alcyonarians Xenia sp.
and Dendronephthya sp. [Pettibone 1969; Wehe 2006].
Distribution in Nhatrang Bay. Mot and Mun Islands.
General distribution. This is the rst record A. rutilans for the Vietnamese coast.
Known also from Red Sea, Persian Gulf [Wehe 2006], Philippines, South-West Australia
[Pettibone 1969].
16
chapter 1
Gastrolepidia clavigera Schmarda, 1861
(Pl. 2 A–D)
Gastrolepidia clavigera Schmarda 1861: 159, pl. 36, g. 316; Uschakov 1982: 119, pl. 37, gs
8–12, pl. 38, gs. 5–6; Hanley 1989: 17, gs 8 A–H, 9 A–F, 10 A–G; 1993: 305; Britayev et al.
1999: 366– 68, gs 19–23; Barnich et al. 2004: 302–303; Wehe 2006: 59, g. 6 a–h.
Material examined. Nhatrang Bay: Tre Island, 1–2 m, on Stichopus chloronotus,
coll. Britayev, 07.12.1985 – 5 cs (♂), 3 cs (♀); Tam Island, 3–5 m, on Holothuria atra,
coll. Britayev, 07.11.1985 3 cs (♂), 2 cs (♀); Mun Island, 7 m, on Pearsonothuria
graeffei, coll. Britayev, 21.10.2005 1 af; Nok Island, 7–10 m, on Holothuria edulis,
29.05.08, only photo; Mot Island, 3–4 m, on Synapta maculata, 18.04.08, only photo.
Diagnosis. Body attened dorso-ventrally, tapering anteriorly and posteriorly. Up
to 31 pairs of elytra on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, 35, 36, 38, 40,
42, then varying between specimens. Elytra large, soft, smooth without papillae. Prosto-
mium wider than long, without cephalic peaks. Ceratophores of lateral antennae inserted
terminoventrally, converging midventrally. Median antenna, tentacular and dorsal cirri
with bulbous, clavate style abruptly terminating into slender terminal lum. Palps stout,
tapering gradually. Two pairs of large eyes on posterior half of prostomium. Notopodia
small, tapering to acicular processes. Neuropodia with well developed lobes, prechaetal
one longer, subtriangular, postchaetal rounded. Conspicuous ventral lamellae at base
of parapodia. Notochaetae short to long, curved, with rows of spines and notched tips.
Neurochaetae longer, falcate, with rows of spines below hooded, notched tips in upper
ones and slightly curved, unidentate tips in middle and lower ones.
Measurements. With up to 57 chaetigers, L up to 27,9 mm, WW up to 3,7 mm, WP
7 mm, WC – 9,3 mm.
Coloration. Varying in accordance to host color pattern: semitransparent, white
(Pl. 2 B, C), brown (Pl. A, D) or black.
Taxonomic remarks. Our specimens t well with the description by Hanley [1989].
Gastrolepidia clavigera can be confused with Asterophilia culcitae, but may be easily
distinguished by the number of elytrigerous segments (more than 15), and the conspicu-
ous ventral lamellae at base of parapodia.
Ecological notes. Found at 1–10 m on the holothurians Holothuria atra Jaeger, 1833,
H. edulis Lesson, 1830, Pearsonothuria graeffei (Semper, 1868), Stichopus chloronotus
Brandt, 1835 and Synapta maculata (Chamisso et Eysenhardt, 1821) (Echinodermata:
Holothuroidea). Recorded previously as symbiont of holothurians Actinopyga echinites
(Jaeger, 1833), A. mauritana (Quoy et Gaimard, 1834), Actinopyga sp., Bohadschia
argus Jaeger, 1833, H. atra, H. edulis, H. fuscopunctata Jaeger, 1833, H. hilla Les-
son, 1830, H. leucospilota (Brandt, 1835), H. nobilis (Selenka, 1867), H. scabra Jaeger,
1833, P. graeffei, S. chloronotus, S. herrmanni Semper, 1868, S. horrens Selenka, 1867,
S. mollis (Hutton, 1872), Thelenota ananas (Jaeger, 1833), T. anax Clark, 1921 (Echino-
dermata: Holothuroidea) [Martin, Britayev 1998]. Holothuria S. maculata is reported as
new host for G. clavigera.
Distribution in Nhatrang Bay. Tre, Tam, Mot, Mun, Nok Islands.
General distribution. Socotra Archipelago, Madagascar, Sri Lanka, South China
17
t.a. britayev, t.i. antokhina
Sea, Papua New Guinea, Bismarck Sea, Australia, Western Pacic Islands [Hanley 1989;
Britayev et al. 1999; Barnich et al. 2004; Wehe 2006].
Heteralentia ptycholepis (Grube, 1878)
Polynoe ptycholepis Grube 1878: 39, 40, pl. 2, g. 6.
Halosydna oculata Treadwell 1926: 8–10, gs. 9–13.
Heteralentia ptycholepis: Hanley, Burke 1991: 5861, Fig. 18 (synonymy); Imajima 1997: 70–3,
gs. 32–33; Barnich et al. 2004: 293–294.
Material examined. Nhatrang Bay, Nok Island, 10 –15 m, on Phanogenia gracilis,
coll. Marin, 20.10.2006 – 1 cs in two fragments; Mun Island, 20–25 m, on P. gracilis,
coll. Marin, 10.2006 – 1 cs (lost).
Diagnosis. Body attened, tapering posteriorly, with up 43 chaetigers. Two transverse
ciliary bands on dorsum of each segment. With 16–18 pairs of elytra on chaetigers 2, 4, 5,
7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32, 35, 38, 39; in smaller specimens, two last pairs on
chaetigers 35–36. Four posterior chaetigers without elytra. Elytra large, folded, completely
covering worm’s dorsum, with three spined microtubecles, Prostomium with rounded lobes,
two times wider than long, three thin antennae with liform tips, median one longer than lat-
erals; two massive palps with abruptly tapering tips and longitudinal dorsal ridge; two pairs of
eyes, anterior pair in the widest part of prostomium, larger than posterior one; posterior pair at
the hind margin of prostomium, surrounded by small brown spots. Tentacular segments with
two long tentacular cirri, ventral one longer than dorsal. Buccal segment with large nuchal
fold covering posterior part of prostomium. Notopodia as a small triangle on dorsal base of
neuropodia. Neuropodia with long triangular prechaetal lobes with digitiform acicular ap-
pendages, and shorter, rounded postchaetal lobes. Notochaetae straight with unidentate or
slightly notched tips, thinner than neurochaetae. Neurochaetae of two types: supraacicular
neurochaetae thin, with slightly expended and nely serrated distal part, and bidentated tips;
subacicular neurochaetae slightly thicker than notochaetae, increased in width downward,
shorter, with more prominent serration, in the lowest part of the bundle with unidentated tips.
All neurochaetae with unusual serration along outer distal edge.
Measurements. With up to 43 chaetigers, L up to 25 mm, WW 2.2 mm, WP
4.7 mm, WC – 7.2 mm.
Coloration. Preserved specimen light brown with poorly distinct transverse dorsal
bands in anterior body.
Taxonomic notes. Our specimens belong to the form with 18 pairs of elytra. In gen-
eral, it agrees with the re-description of the holotype from Philippines [Hanley, Burke
1991]. However, there are several small differences: notched notochaetae instead of all
notochaetae with entire tips (holotype) and lower neurochaetae unidentate instead of all
neurochaetae bidentate (holotype).
Ecological notes. Found at 10–25 m on crinoid Phanogenia gracilis (Hartlaub,
1890) (Echinodermata: Crinoidea). This is the rst record of the species in association
with other animals.
Distribution in Nhatrang Bay. Nok and Mun Islands.
18
chapter 1
General distribution. This is the rst record of H. ptycholepis for the Vietnamese
coast. Gulf of Oman, Mauritius, Hainan, Hong Kong, Philippines, Japan, Papua New
Guinea, Chestereld Islands, Samoa [Hanley, Burke 1991; Barnich et al. 2004].
Hololepidella laingensis Britayev, Doignon et Eeckhaut, 1999
(Pls. 2 E, F; 3 A)
Hololepidella laingensis Britayev, Doignon, Eeckhaut 1999: 360–363, gs. 2–12.
Material examined. Nhatrang Bay: Mieu Island, 5–6 m, on Culcita novaeguineae,
coll. Marin, 28.04.2004 – 5 cs; 5–7 m, on C. novaeguineae, coll. Marin, 12.05.2004
1 cs; 5–7 m, on C. novaeguineae, coll. Marin, 14.05.2004 – 6 cs; Tam Island, 5–6 m,
on C. novaeguineae, coll. Marin, 20.05.2004 – 10 cs and 1 af; Mot Island, 5–6 m, on C.
novaeguineae, coll. Marin, 21.05.2004 5 cs and 6 af; 4 m, on C. novaeguineae, coll.
Antokhina, 06.04.2006 – 1 af; 10 m, on Cenometra bella, coll. Savinkin, 17.11.2006 –
1 af; Tre Island, st. 4, 3–5 m, on C. novaeguineae, coll. Marin, 23.05.2004 – 1 af; 7 m, on
Acanthaster planci, coll. Antokhina, 08.05.2006 – 1 cs; Mun Island, on C. novaeguineae,
5–6 m, coll. Marin, 26.05.2004 – 7cs and 5 af; Nok Island, 10 m, on Dichrometra agel-
lata, coll. Mekhova, 26.05.2007 5 af; 10 m, on Liparometra regalis, coll. Mekhova,
26.05.2007 – 1 af; 10 m, on L. regalis, coll. Mekhova, 26.05.2007 – 1 cs and 1 af; 8 m, on
Lamprometra palmata, coll. Mekhova, 16.05.2007 – 4 af; Dung Island, 12 m, on Linckia
laevigata, coll. Antokhina, 24.04.2006 – 1 cs.
Diagnosis. Body short attened, tapering posteriorly, with up to 40 chaetigers. On
the dorsum of each segment one transverse ciliary band. With 13–18 pairs of elytra on
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 31, 34, 36, 38, large, covering
dorsal surface completely, soft, translucent, with group of conical microtubercles on an-
terior part. Prostomium wider than long, with cephalic peaks or with slight indication of
peaks, two pairs of dark brown eyes, anterior pair larger, in some specimens eyes nearly
invisible due to camouage with dark pigmentation; lateral antennae arising anteriorly,
slightly ventral to median antenna. Tentaculophores with 1–3 aciculae in each. Notopo-
dia small, rounded with projecting acicular lobes. Neuropodia with conical prechaetal
acicular lobes and shorter rounded or triangular postchaetal lobes. Notochaetae stouter
than neurochaetae, relatively few, slightly curved, with blunt entire or slightly notched
tips usually 10–12 teeth along convex margin, most of neurochaetae slightly bidentate,
while 1–3 subacicular ones with entire tips.
Measurements. With up to 40 chaetigers, L up to 17.5 mm, WW up to 1.7 mm, WP
2.9 mm, WC – 7.1 mm.
Coloration. Variable, from light brown to dark brown (Pl. 2 E, F), sometimes black.
Taxonomic notes. This species belongs to of the short-bodied Hololepidella. It dif-
fers from H. millari in having a distinctive dark pigmentation, a relatively shorter body,
less elytra, and unidentate neurochaetae.
Ecological notes. Found at 3–12 m on starshes (viz. Acanthaster planci (Lin-
naeus, 1758), C. novaeguineae, L. laevigata) and crinoids (viz. Cenometra bella (Hart-
laub, 1890), Dichrometra agellata (Müller, 1841), Lamprometra palmata (Müller,
19
t.a. britayev, t.i. antokhina
1841), Liparometra regalis (Carpenter, 1888)). Previously recorded as symbiont of
the crinoids Capillaster multiradiatus (Linnaeus, 1758), Dichrometra agellata, L.
palmata, Himerometra robustipinna (Carpenter, 1881), Oxycomanthus bennetti (Mül-
ler, 1841), Comaster multidus (Müller, 1841) in Papua New Guinea [Britayev et al.
1999]. The starshes A. planci, C. novaeguineae and L. laevigata are reported as new
hosts for the species.
Distribution in Nhatrang Bay. Mieu, Tam, Mot, Mun, Dung, Nok Islands.
General distribution. Vietnam, Nhatrang Bay. Known also from Papua New Guinea
[Britayev et al. 1999].
Hololepidella millari Britayev, Doignon et Eeckhaut, 1999
(Pl. 3 B–D)
Hololepidella millari Britayev, Doignon, Eeckhaut 1999: 363–365, gs. 13–14.
Material examined. Nhatrang Bay: Mieu Island, 5–6 m, on Acanthaster planci, coll.
Marin, 14.05.2004 – 2 af; Mot Island, 7–10 m, on Choriaster granulatus, coll, Antokhi-
na, 15.04.2006 – 1 af; Tre Island, 10 m, on Nardoa frianti, coll, Antokhina, 08.05.2006 –
1 cs; Mot Island, 5–6 m, on Culcita novaeguineae, coll.Marin, 28.05.2004 – 1 cs and
1 af; 7 m, on C. novaeguineae, coll. Antokhina, 10.04.2006 – 1 cs; Dung Island, 10 m,
on Halityle regularis, coll. Antokhina, 24.04.2006 – 1 cs; 13 m, on Protoreaster nodo-
sus, coll. Antokhina, 26.04.2006 1 cs; 9 m, on Euretaster insignis, coll. Antokhina,
10.05.2006 – 1 cs.
Diagnosis. Body elongated with more than 55 chaetigers, without prominent pig-
mentation. One transverse ciliary band on dorsum of each segment. Up to 30 and more
pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 31, 34, 36,
thereafter varying between specimens, usually on alternating segments to end of body,
but often asymmetrical. Elytra large, covering dorsal surface completely, soft, translu-
cent, smooth except for some microtubercles on anterior part. Prostomium slightly wider
than long, without distinct cephalic peaks, with two pairs of eyes, anterior pair slightly
larger; lateral antennae very short, with swollen ceratophores. Tentaculophores with one
aciculae. Notopodia small, with projecting acicular lobes; neuropodia with digitiform
prechaetal acicular lobes and shorter triangular postchaetal lobes. Notochaetae about
same width as neurochaetae, relatively few, slightly curved; superior notochaetae with
entire tips, inferior ones with bidentate or split tips and 18–24 rows of spines along the
convex edge, all neurochaetae with bidentate hooked tips.
Measurements. With up to 75 chaetigers, L up to 34 mm, WW up to 1.5 mm, WP –
2.51 mm, WC – 2.98 mm.
Coloration. White or esh-colored without dark pigmentation (Pl. 3 C, D).
Taxonomic notes. This species belongs to the long-bodied Hololepidella.
Ecological notes. Found at 5–13 m on the starshes Acanthaster planci, Choriaster
granulatus, Culcita novaeguineae, Euretaster insignis, Halityle regularis Fisher, 1913,
Nardoa frianti Koehler, 1910 and Protoreaster nodosus. Previously recorded as symbi-
ont of feather stars, Comanthus alternans (Carpenter, 1881) (Echinodermata: Crinoidea)
20
chapter 1
and starshes, Linckia guildingi Gray, 1840 and L. laevigata (Echinodermata: Asteroi-
dea), in Papua New Guinea [Britayev et al. 1999]. Starshes A. planci, C. granulatus,
C. novaeguineae, E. insignis, H. regularis, N. frianti and P. nodosus are reported as new
hosts for the species.
Distribution in Nhatrang Bay. Mieu, Tam, Mot, Mun, Dung Islands.
General distribution. Vietnam, Nhatrang Bay. Known also from Papua New Guinea
[Britayev et al. 1999].
Hololepidella sp.
(Pl. 3 E, F)
Material examined. Nhatrang Bay: Mot Island, 3–6 m, on Galaxea astreata, coll. Marin,
16.04.2009 – cs and af; Mun Island, 6 m, on G. astreata, coll. Marin, 06.10.2003 – 1 cs with
inverted pharynx; Tam Island, 4–7 m, on G. astreata, coll. Marin, 10.10.2003 – 1 cs.
Diagnosis. Body elongated with more than 80 chaetigers, without prominent pig-
mentation. On the dorsum of each segment one transverse ciliary band. Up to 40 and
more pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 31, 34,
36, continuing on alternate segments towards the end of body, but sometimes on two
neighboring segments. Elytra large, covering dorsal surface completely, soft, translu-
cent, smooth except of anterior part with numerous conical microtubercles. Prostomium
as wide as long, hexagonal in outline, with distinct cephalic peaks and two pairs of eyes,
anterior pair slightly larger; lateral antennae very short, with swollen ceratophores. Ten-
taculophores with aciculae and up to 3 chaetae in each. Notopodia small, rounded with
projecting acicular lobes. Neuropodia with conical prechaetal acicular lobes and shorter
rounded postchaetal lobes. Notochaetae numerous (25–29) stouter than neurochaetae,
slightly curved, with entire tips, and 6–9 rows of spines along the convex edge nearly
reaching their tips. Neurochaetae numerous (30–32), all with slight serration and blunt
(rounded) notched tips.
Measurements. With up to 82 chaetigers, L up to 22 mm, WW up to 1.1 mm,
WP – 2.2 mm, WC – 3.2 mm.
Coloration. Without distinctive pigment pattern; light brown or eshly white alive,
white in alcohol (Pl. 3 E, F).
Taxonomic notes. This is likely a new species. It differs from the other long-bodied
species, H. millari, in having distinctive cephalic peaks in prostomial lobes, 1–3 chaetae
in tentaculophores, numerous noto- and neurochaetae, and notochaetae with entire tips
and 6–9 rows of spines along convex edge, instead of notochaetae with notched tips and
18–24 rows of spines along convex edge (in H. millari).
Ecological notes. Found at 4–10 m on the scleractinian coral Galaxea astreata La-
marck, 1816 (Hexacorallia: Scleractinia). This is the rst record of a scaleworm in as-
sociation with scleractinian corals.
Distribution in Nhatrang Bay. Mot, Mun, Tam and Tre Islands.
General distribution. Nhatrang Bay, Vietnam.
21
t.a. britayev, t.i. antokhina
Ophthalmonoe pettiboneae Petersen et Britayev, 1997
(Pl. 4 A, B)
Ophthalmonoe pettiboneae Petersen, Britayev 1997: 266 –272, gs 1–3; Britayev, Martin 2005:
4091–4096, gs 7–8.
Material examined. Nhatrang Bay: Mun Island, 15 m, sand, in tubes of Chaetop-
terus sp., coll. Marin, 16.06.2004 2 specimens (af and pf) and 2 mf; 11 m, sand, in
tubes of Chaetopterus sp., coll. Savinkin 1 cs; western point of Tre Island, 8–12 m,
sandy silt, in tubes of Chaetopterus sp., coll. Marin, 29.06.2004 – 2 cs.
Diagnosis. Body attened, relatively short and wide, truncate anteriorly and gradu-
ally tapering posteriorly. With 19 pairs of elytra on chaetigers 2, 4, 5, 7, 9, 11, 13,
15, 17, 19, 21, 23, 26, 29, 32, 35, 38, 41, 42. Last two to four segments cirrigerous,
without elytra. Elytra relatively large, overlapping antero-posteriorly, basically oval,
but markedly irregular in outline, folded along external margin, delicate, smooth ex-
cept for some scarce irregularly distributed micropapillae. Prostomium slightly wider
than long, without cephalic peaks, with two pairs of large dark eyes, with anterior pair
slightly larger than posterior one, directed anteriorly, both pairs with conspicuous oval
lenses; lateral antennae inserted slightly ventral to median antenna, short, cylindrical,
ceratophores short, converging mid-ventrally. Tentaculophores relatively short, thick,
achaetous, but with acicula. Dorsal cirri alternating in length along body. Notopodia,
small, conical; neuropodia longer and wider than notopodia, with row of cilia along
ventral side, prechaetal lobe elongated, nearly digitiform, substantially longer than
triangular postchaetal lobe. Notochaetae arranged in a fan-shaped bundle orientated
nearly horizontally, numerous, from 70 to more than 150 per bundle, slender than
neurochaetae, long, capillaries, smooth. Neurochaetae numerous, from more than 80
to more than 240 per bundle, stout, with tips hooded, blunt and nearly rounded, with
rows of narrow petaloid spines.
Measurements. Body with 44–46 chaetigers, L – 19–27 mm, WW – 2.6 –3.8 mm,
WP – 9.0–9.8 mm, WC – 11.2–12.3 mm.
Coloration. The body is basically light brownish and has a conspicuous trans-
verse dark brown or black band crossing each segment. On segment 5, there is an
additional characteristic oval, mid-dorsal, black spot posterior to the transverse band
(Pl. 4 A, B). The longest dorsal cirri have more intense dark pigmentation than the
smaller ones. The ventral body surface is always non-pigmented [Britayev, Martin
2005].
Taxonomic notes. Our specimens agree completely with description provided by
Britayev and Martin [2005].
Ecological notes. Found at 8–15 m in tubes of the polychaete Chaetopterus sp.
(Polychaeta: Chaetopteridae).
Distribution in Nhatrang Bay. Mun and Tre Islands.
General distribution. Vietnam [Britayev, Martin 2005], coasts of Ambon Island,
Indonesia, Banda Sea [Petersen, Britayev 1997].
22
chapter 1
Paradyte crinoidicola (Potts, 1910)
(Pl. 4 C, D)
Polynoe crinoidicola Potts 1910: 337, pl. 18, g. 10, pl. 21, gs 39– 41.
Adyte crinoidicola: Hanley 1984: 88–90, g. 1.
Scalisetosus longicirrus: Fishelson, Rullier 1969: 54–55 [not Schmarda, 1861].
Paradyte crinoidicola: Pettibone 1969: 13 –16, g. 7; Tchesunov et al. 1989: 169–172; Hanley,
Burke 1991: 13–15, g. 1; Britayev et al. 1999: 369–371, gs. 24–29; Barnich et al. 2004, 312;
Wehe 2006: 154 –156, g. 41.
Material examined. Nhatrang Bay: Tre Island, 6 m, on Dichrometra agellata, coll.
Mekhova, 01.06.2007 4 cs and 4 af ; 6 m, on Stephanometra indica, coll. Mekhova,
07.05.2007 – 6 af; 8 m, on Comatella nigra, coll. Mekhova, 21.05.2007 – 2 cs and
4 af; 8 m, on Comanthus parvicirrus, coll. Mekhova, 21.05.2007 – 10 cs and 8 af; 8 m,
on Colobometra perspinosa, coll. Mekhova, 21.05.2007 7 cs and 9 af; Tam Island,
6 m, on Lamprometra palmata, coll. Mekhova, 01.06.2007 – 2 cs and 3 af; Mot Island,
10 m, on Cenometra bella, coll. Mekhova, 17.11.2006 – 1 pf; 8 m, on Comaster nobilis,
coll. Mekhova, 30.05.2007 – 118 cs and 24 af; 7 m, on Stephanometra tenuipinna, coll.
Mekhova, 30.05.2007 – 12 cs and 3 af; Mun Island, 10 –12 m, on Amphimetra ensifera,
04.05.2007 – 1 cs and 2 af; 10 –12 m, on C. bella, coll. Mekhova, 04.05.2007 – 14 cs and
4 af; 10 – 12 m, on Comanthus alternans, coll. Mekhova, 04.05.2007 – 9 cs and 21 af;
6 m, on C. gisleni, coll. Mekhova, 04.05.2007 – 60 cs and 18 af; 6 m, on Himerometra
robustipinna, coll. Mekhova, 04.05.2007 – 14 cs and 2 af; 10–12 m, on Amphimetra tes-
sellata, coll. Mekhova, 10.05.2007 – 1 af; Nok Island, 12 m, on Oxycomanthus bennetti,
coll. Mekhova, 25.04.2007 6 af; 10–12 m, on Phanogenia gracilis, coll. Mekhova,
02.05.2007 1 af; 10 m, on Comatella stelligera, coll. Mekhova, 26.05.2007 3 cs;
10 m, on Liparometra regalis, coll. Mekhova, 26.05.2007 – 3 af; 68 m, on Oxycoman-
thus pinguis, coll. Mekhova, 29.05.2008 – 5 af.
Diagnosis. Body attened, elongate-oval, tapered posteriorly, with up to 38 segments.
Two transverse ciliary bands on dorsum of each segment, connecting cirrophore or elytro-
phore bases. Fifteen pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26,
29 and 32. Posterior segments cirrigerous without elytra. Elytra delicate, smooth, without
tubercles. Prostomium wider than long, without frontal peaks, with two pairs of eyes, cera-
tophores of lateral antennae inserted ventrally. Tentacular segment without chaetae. Second
(buccal) segment with small nuchal fold covering posterior prostomial region. Dorsal cirri
with cylindrical cirrophores, bulbous basally, with long tapered styles extending beyond
neurochaetae. Notopodia small triangle; neuropodia with long conical prechaetal lobes and
shorter rounded postchaetal lobes. Notochaetae stouter than neurochaetae, sabre-like, with
notched tips and 1–5 widely spaced spines. Neurochaetae of two kinds, both with basal
semilunar pockets, supraacicular group slender, slightly curved, with long spinous region
and bidentate tips, subacicular group stouter, with tips entire and falcate.
Measurements. With up to 38 chaetigers, L up to 11,6 mm, WW up to 1,1 mm, WP
3,2 mm, WC – 3,4 mm.
Coloration. Usually in accordance with host color, as described in detail by Britayev et
al. [1999]. There are two main pigmentation patterns. Specimens with uniform pigmenta-
tion varying from light pinkish to dark brown, nearly black (Pl. 4 C) and specimens with
23
t.a. britayev, t.i. antokhina
different pigment patterns: (A) light body with a longitudinal dark band covering prostomi-
um and middle of the dorsal surface up to bases of parapodia, ventral surface without pig-
ment; (B) pigment predominates, light areas restricted to elytrophores, to dorsal tubercles
and prostomium, ventral surface with pigmented areas at base of parapodia; (C) similar to
B but without pigment on ventral side. Pigmentation of elytra in these three types usually
uniform, in accordance with body color, sometimes lighter or with dark or light spots.
Taxonomic notes. This is a very variable species in terms of color pattern, body propor-
tions, and relative elytra size. It clearly differs from P. levis in having a distinctive pigmen-
tation, notochaetae with notched tips and 1–5 spines instead of 0–2 spines (in P. levis), and
supraacicular neurochaetae with bidentate tips instead of faintly notched tips (in P. levis).
Ecological notes. Found at 6–12 m on the crinoids Amphimetra ensifera (A.H. Clark,
1909), A. tessellata (Müller, 1841), Cenometra bella, Colobometra perspinosa Carpenter
1881, Comanthus gisleni Rowe, Hoggett, Birtles et Vail, 1986, C. parvicirrus (Müller,
1841), Comatella nigra (Carpenter, 1888), C. stelligera (Carpenter, 1888), Comaster no-
bilis (Carpenter, 1884), Himerometra robustipinna, Lamprometra palmata, Liparometra
regalis, Oxycomanthus pinguis (A.H. Clark, 1909), Phanogenia gracilis, Stephanometra
indica (Smith, 1876), S. tenuipinna (Hartlaub, 1890) (Echinodermata: Crinoidea). This is
one of the most common and widespread symbionts of Indo-West Pacic feather stars.
Found previously on crinoids Capillaster multiradiatus, Colobometra perspinosa, Co-
manthus alternans, C. briareus (Bell, 1882), C. parvicirrus, Comatula purpurea (Müller,
1843), Comaster schlegelii (Carpenter, 1881), C. multidus, Comatella stelligera, Dichro-
metra agellata, Himerometra robustispina, Lamprometra palmata, Oligometra carpen-
teri (Bell, 1884), O. serripinna (Carpenter, 1881), Oxycomanthus bennetti, Petasometra
helianthoides Clark, 1912, Stephanometra indica, Tropiometra carinata (Lamarck, 1816)
[Hanley 1984, 1991; Tchesunov et al. 1989; Britayev et al. 1999]. Feather stars A. en-
sifera, A. tessellata, Cenometra bella, Comanthus gisleni, C. parvicirrus, Comatella nigra,
Comaster nobilis, Lamprometra palmata, Liparometra regalis, O. pinguis, P. gracilis, S.
tenuipinna are reported as new hosts for P. crinoidicola.
Distribution in Nhatrang Bay. Tre, Tam, Mot, Mun, Nok Islands.
General distribution. This is the rst record for the Vietnamese coast. Widely dis-
tributed in the tropical Indo-West Pacic: Red Sea, South Africa, Maldives, Malay Ar-
chipelago, Hong Kong, South China Sea, Papua New Guinea, Bismarck Sea, Australia,
Japan, Caroline, Marshall, and Solomon Islands [Hanley 1984; Hanley, Burke 1991;
Britayev et al. 1999; Barnich et al. 2004; Wehe 2006].
Paradyte levis (Marenzeller, 1902)
(Pl. 4 E, F)
Scalisetosus levis Marenzeller 1902: 575–576, pl. 3: gs 12, 12 A; Fauvel 1933: 38–40, g. 1;
Hartman 1974: 207–209.
Paradyte levis: Imajima 1997: 6–7, gs 3, 4; Wehe 2006: 156–158, g. 42.
Material examined. Nhatrang Bay: Nok Island, 23 m, on Dendronephthya sp., (995–2),
coll. Savinkin, 17.06.2004 – 1 cs with everted pharynx; 18–20 m, on Dendronephthya sp.,
24
chapter 1
coll. Savinkin, 29.05.2006 2 cs, 8 af, 5 mf; 15–17 m, boulders, on Dendronephthya sp.,
(No1, No 6), coll. Britayev, 02.05.2007 – 1 cs; Mot Island, 10–12 m, on Dendronephthya sp.
(D2, D4, D6, D9, D10), coll. Savinkin, 07.06.2006 – 5 cs, 3 af; 10 m, on Dendronephthya sp.,
coll. Savinkin, 17.11.2006 – 12 cs, 45 mf, 13 pf; Mun Island, 18–26 m, on Dendronephthya
sp. (998/12), coll. Savinkin, 27.10.2008 – 3 cs, 4 af (6 juv.).
Diagnosis. Body attened dorsoventrally, elongate, with up to 39 chaetigers. Two trans-
verse ciliary bands on dorsum of each segment, connecting the bases of cirrophores or ely-
trophores. Fifteen pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29
and 32. Posterior segments cirrigerous without elytra. Elytra large, rounded to oval, smooth,
semitransparent, with few, very small, scattered sensory papillae. Prostomium rounded to
hexagonal, about as wide as long, with cephalic peaks, ceratophores of lateral antennae in-
serted ventrally. Tentacular segment without chaetae. Dorsal cirri with cylindrical cirrophore
and long, smooth style gradually tapering and extending beyond neurochaetae. Notopodia of
moderate size, subtriangular, with acicular process; neuropodia with longer, more triangu-
lar, prechaetal, acicular lobes and shorter, rounded, postchaetal lobes. Notochaetae arranged
tuft-like, stouter than neurochaetae, sabre-like, with blunt, unidentate tips, usually without
spines or with one spine, rarely with two spines on convex border. Neurochaetae thinner, of
two kinds: supraacicular chaetae more slender, with subdistal semilunar pockets and numer-
ous rows of spines below slightly hooked and faintly notched tips; subacicular stouter, with
semilunar pockets and fewer rows of spines below more strongly hooked unidentate tips.
Measurements. With up to 39 chaetigers, L up to 18 mm, WW up to 1.5 mm, WP –
3.9 mm, WC – 4.5 mm.
Coloration. Whitish, semitransparent (Pl. 4 E, F).
Taxonomic notes. Our specimens agree well with description provided by Wehe [2006],
however they have notochaetae usually with 0 –1, and rarely 2 lateral spines, while Wehe
indicated up to 3 spines in some anterior segments. One of the specimens collected at Mun Is-
land (998/12) has an additional pair of elytra on 35th chaetiger covering posterior segments.
Ecological notes. Found at 18–26 m on the octocoral Dendronephthya sp. (Octocoral-
lia: Alcyonacea). Previously recorded on muddy sand, amongst sponges and corals, in tubes
of the polychaete Euniphysa tubifex (Crossland, 1904), and on Dendronephthya sp. [Okuda
1950; Imajima 1997; Wehe 2006].
Distribution in Nhatrang Bay. Mot, Mun, Nok Islands.
General distribution. This is the rst record of P. levis for the Vietnamese coasts.
Known also from the Red Sea, Suez Bay, Arabian Sea, Oman Gulf, Japan [Marenzeller
1902; Imajima 1997; Wehe 2006].
Pottsiscalisetosus praelongus (Marenzeller, 1902)
(Pl. 5 A, B)
Scalisetosus praelongus Marenzeller 1902: 75, pl. 3, g. 11; Imajima, Hartman 1964: 39.
Pottsiscalisetosus praelongus: Pettibone 1969: 9, f ig. 9; Barnich et al. 2004: 316–317.
Material examined. Nhatrang Bay: Tre Island, 23.5 m, on Luidia maculata, coll.
Antokhina, 02.05.2006 – 2 cs; 20 m, on L. maculata, coll. Antokhina, 06.05.2006
25
t.a. britayev, t.i. antokhina
1 cs; food-market of Nha Trang city, on Archaster angulatus, coll. Antokhina, May
2006 – 2 cs and 5 af.
Diagnosis. Body with 50 to 110 or more segments. On dorsum of each segment two
transverse ciliary bands. Elytra numerous 36 or more pairs arranged on segments 2, 4,
5, 7, alternate segments to 23, 26, 29, 32, 33, 36, 38, 40, 42, continuing on alternate seg-
ments to end of body. Elytra semitransparent, smooth. Prostomium without cephalic peaks;
lateral antennae inserted terminoventrally. Tentaculophores without chaetae. Notopodia
with short conical acicular lobes. Neuropodia long with prechaetal lobes truncate, without
supra-acicular process; postchaetal lobes shorter, rounded. Notochaetae few, slender, with
ne rows of spines and blunt tips. Neurochaetae stouter, falcate, with subdistal semi-lunar
pockets, ne rows of distal spines, and minutely bi- or unidentate tips.
Measurements. With up to 73 or more chaetigers, L up to 23.8 mm, WW up to
1.4 mm, WP – 3.8 mm, WC – 5.1 mm.
Coloration. White-grey or white-yellow (Pl. 5 A).
Taxonomic notes. Our specimens completely match the original description by Pet-
tibone [1969].
Ecological notes. Found at 20–23 m deep on the starshes Luidia maculata Müller
et Troschel, 1842 and Archaster angulatus Müller et Troschel, 1842 (Echinodermata:
Asteroidea). This is the rst report of these starsh species as a host.
Distribution in Nhatrang Bay. Tre Island, Dam Bay.
General distribution. Vietnam, Nhatrang Bay. Known also from the Indian Ocean,
Hainan, Japan [Pettibone 1969; Barnich et al. 2004].
Uncopolynoe corallicola Hartmann-Schröder, 1960
(Pl. 5 C, D)
Uncopolynoe corallicola Hartmann-Schröder 1960: 70–71, pl. 1: gs. 1, 6, pl. 3: gs 2–5;Amoureux
et al. 1978: 72.; Wehe 2006: 167–168, g. 46.
Material examined. Nhatrang Bay, Mun Island, 18–26 m, rock and sand, on Ne-
phthyidae gen. sp., coll. Savinkin, 27.10.2008 – 1 cs, female.
Diagnosis. Body small, fusiform, with up to 67 segments. Elytra up to 33 pairs on
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32, 34, 36, 38, 39, 41, 42, 44, 46,
48, 49, 51, 53, 55, 57, 59, 61, 63, 65; oval, soft, translucent, smooth. Prostomium broadly
oval, without cephalic peaks, with two pairs of eyes and three antennae; median antenna
with large ceratophore and long style, lateral antennae without ceratophore and small
papilla-shaped styles. Notopodia and notochaetae absent. Neuropodia stout, conical; on
segments 2 to 4 with distinct, rounded pre- and postchaetal lobes, enclosing bases of neu-
rochaetae; following neuropodia bluntly rounded without neuropodial lobes.Neuropodia
on segments 2, 3 and 4 with hooked-like neurochaetae, following segments usually with
two chaetae, one bidentate and one clavate in each neuropodium.
Measurements. Sixty seven chaetigers, L – 5.6 mm, WW – 0.4 mm, WP – 0.95 mm.
Coloration. Whitish, semitransparent (Pl. 5 C, D).
Taxonomic notes. Our specimen agrees in general with the descriptions of Hart-
26
chapter 1
mann-Schröder [1960] and Wehe [2006]. However, all previously described specimens
(including the types) were incomplete, so the total number of both segments and elytra,
as well as the elytra distribution, remained unknown till now. Wehe [2006] reported “at
least 40 segments” and “at least 12 pairs…” of elytra. The nding of complete specimen
in Vietnam conrms that, despite its small size, U. corallicola is a multisegmental worm
with very small segments and numerous elytra on neighboring segments (38,39… 41,
42… 48,49), which is an unusual feature for scale worms.
Ecological notes. Found at 18–26 m crawling on dendronephthyid coral (Octocoral-
lia: Alcyonacea) (Pl. 5 C).
Distribution in Nhatrang Bay. Mun Island.
General distribution. This is the rst record of genus and species for Vietnam, as
well as for the Pacic Ocean. Nothern Red Sea, Gubal Island [Wehe 2006].
Family SYLLIDAE Kinberg, 1856
Alcyonosyllis phili Glasby et Watson, 2001
(Pls. 5 E, F; 6 A, B)
Alcyonosyllis phili Glasby, Watson 2001: 45–49, gs. 1–5; Glasby, Aguado 2009: 60.
Material examined. South China Sea, Nhatrang Bay, Nok Island, 9 m, on Nephthy-
idae gen. sp. (IV–1), coll. Britayev, 12.05.2006 – 1 cs, 1 af; 9 m, on Nephthyidae gen. sp.
(IV–2), coll. Britayev, 12.05.2006 – 1 cs; Phu Quoc Island, 10–12 m, on Nephthyidae
gen. sp., coll. Savinkin, 12.10.2008 1 cs; 10–12 m, on Carijoa sp., coll. Savinkin,
16.10.2008 – 1 cs; 10–15 m, on Dendronephthya sp., coll. Savinkin, 16.10.2008 – 1 cs,
1 af; 10–12 m, on undescribed gorgonian species, coll. Savinkin, 16.10.2008 – 2 cs.
Diagnosis. Body long, robust, similar in width throughout, except for slight taper-
ing at head and tail ends. Prostomium sub-quadrangular, about twice as wide as long.
Two pairs dark laterally-positioned eyes, similar in size. Median antenna about twice
longer and thicker than lateral ones, inserted on midposterior prostomium. Palps large,
slightly longer than prostomium length, separated all along their length. Pharynx with
anterior tooth, occupying rst 4–5 chaetigers, anterior rim chitinized, with smooth
border and 10 small pharyngeal papillae. Proventricle extending through 6 segments
(from 4–5 to 9–10) with about 40 muscle cell rows. Tentacular segment projecting
slightly mid-anteriorly to cover nuchal organs, with two pairs of cirri; dorsal pair
slightly thicker and twice length of ventral pair. Dorsal cirri of two distinct forms:
dorsally-displaced larger cirri with distinct cirrophore, on chaetigers 1, 4, 6, 9, 11 and
thereafter on odd chaetigers; unarticulated, inated, tapering to a point; held erect and
curved over dorsum. Smaller, laterally-directed dorsal cirri arising closer to parapodial
lobes than larger cirri, on chaetigers 2,3,5,7,8, 10 and thereafter on even chaetigers (Pl.
5 E, F). Ventral cirri smooth, approximately as longer as parapodia. Each parapodium
bearing 2–5 unarticulated hooked chaetae (rarely 1) ventral to aciculae: thicker chaeta
more hooked and with very slight subdistal boss; thinner chaeta with more promi-
27
t.a. britayev, t.i. antokhina
nent V-shaped subdistal boss and often with minute tooth immediately below hook.
Aciculae of two types: 3–6 straight, blunt-tipped and 1–2 slender, with attenuated tip,
extending beyond others. Pygidium small, with one pair of large unarticulated cirri,
similar to enlarged dorsal cirri.
Measurements. With up to 191 or more chaetigers, L up to 72 mm, WP – 0.65 mm.
Coloration. Living specimens usually vividly colored with brown dorsal bands on
chaetigers 2, 3, 5, 7, 8, 10 and thereafter on all even ones, blue to purple pigment spot at
base of antennae and tentacular and dorsal cirri, and dark tips of antennae and tentacular
and dorsal cirri (Pl. 5 E, F). The pigmentation intensity varies among specimens, occa-
sionally nearly absent. Pigment pattern fades rapidly after xation.
Taxonomic notes. Our specimens agree well with the original description by Glasby and
Watson [2001], but slightly differ in the absence of a subdistal tooth in the hooked chaetae,
and having a higher number of chaetae (3–5 vs. 2–3) and aciculae (up to 6 vs. up to 5).
Ecological notes. Found at 10–15 m on the soft corals Nephthyidae gen. sp., Carijoa
sp., Dendronephthya sp. (Octocorallia: Alcyonacea). Previously recorded as symbiont of
Dendronephthya sp., Melithaea sp., an undescribed genus and species of Nephtheidae
(Octocorallia: Alcyonacea) [Glasby, Watson 2001], and gorgonians [Glasby, Aguado
2009]. The Darwin Harbor specimens were collected from reef at sediments; all were
relatively small (< 10 mm long), which suggests that young forms may be free-living
[Glasby, Aguado 2009]. The alcyonarian Carijoa sp. is reported as a new host for the
species.
Distribution in Nhatrang Bay. Nok Island.
General distribution. The species and genus represent a new record for the Viet-
namese coasts, including the Phu Quoc Islands. Known also from the northwestern,
northern and eastern coasts of Australia, southern Papua New Guinea, Ashmore Reef
(Timor Sea), Panglao Island (Philippines) [Glasby, Watson 2001; Glasby, Aguado 2009].
The Vietnamese specimens are the northern- and western-most record for the species.
Haplosyllides aberrans (Fauvel, 1939)
(Pl. 6 C, D)
Syllis (Haplosyllis) aberrans Fauvel 1939: 290, 291, g. 3.
Haplosyllides oridana: Uebelacker 1982: 585 (partim); San Martin et al. 1997: 366–378 (partim).
Haplosyllides sp.: Martin et al. 2008: 280, g. 1
Haplosyllides aberrans: Martin et al. 2009: 647–650, gs. 1, 2 E–H, 3, 4, 5 D–F, 7 A–E.
Material examined. Nhatrang Bay: Cauda, collected by M.C. Dawydoff – 1 cs;
Mun Island, 2–3 m deep, attached to Platycaris latirostris, collected by I. Marin,
06.10.2003. – 1 cs.
Diagnosis. Body short, widest at proventricular region, gradually tapering poste-
riorly. Prostomium oval, with one pair of small, brown, posterolateral eyes. Median
antennae attached medially at hind margin of prostomium, long, about three times as
longer as lateral antennae. Lateral antennae inserted anterolaterally to eyes. Palps sim-
ilar in length to prostomium, fused at bases, broadly attached to prostomium. Pharynx
28
chapter 1
similar in length to proventricle, with an anterior pharyngeal tooth, a crown of 10 –12
distal papillae, and a distal ring of cilia. Proventricle short, extending from chaetiger
2 to the end of chaetiger 4, with 31 muscle cell rows. Dorsal tentacular cirri three
times as longer as ventral ones. Antennae, tentacular and dorsal cirri smooth, non-
articulated, slender, tapering to liform tips. Parapodia and cirri with oval elongated
granules roughly distributed in longitudinal parallel rows in dorsal cirri; dorsal cirri
alternating long and short according to following arrangement: chaetiger 1 long (L), 2
and 3 short (S), 4 L, 5 and 6 S, 7 L, then alternating 8 S, 9 L, 10 S, 11 L, 12 S, 13 L,
and 14 S, then 15 S, 16 L , 17 S, and 18 L. Ventral cirri cirriform, with attenuated tips,
smooth, slightly shorter than or equal to parapodial lobes in length. With 2–3 thick
simple chaetae in each parapodium, one usually thicker, all with tridentate curved tips
and sub-distal constrictions with 1–2 aciculae per parapodium, straight, with truncated
tip, similar in width to thicker chaetae.
Measurements. With 20.3 (±4.9) chaetigers, L 3.7 (±1.4) mm, width – 0.50 (±0.15) mm.
Coloration. Semitransparent
Taxonomic notes. The species was recently redescribed by Martin et al. [2009].
Ecological notes. Found at 2–3 m attached to the pontoniin shrimp Platycaris latiro-
stris Holthuis, 1952 (Decapoda: Pontoniinae) (Pl. 6 C, D), an obligatory associate of the
scleractinian coral Galaxea astreata (Hexacorallia: Scleractinia).
Distribution in Nhatrang Bay. Cauda and Mung Island.
General distribution. Nhatrang Bay and Spratly Archipelago, Vietnam, South China
Sea; NE coast of Sumba and Pulu Tongkil, Sulu archipelago, Indonesia [Fauvel 1939;
Martin et al. 2008, 2009].
Inermosyllis sp.
(Pl. 6 E, F)
Material examined. Nhatrang Bay: Tre Island, 23.5 m, on Luidia maculata, coll. An-
tokhina, 02.05.2006 – 1 af; 20 m, on Luidia maculata, coll. Antokhina, 06.05.2006 –1 af;
food-market of Nha Trang city, on Archaster angulatus, coll. Antokhina, May 2006 – 2 af.
Diagnosis. Body long, cylindrical, widest at proventricle level, gradually tapering
posteriorly. Prostomiun pentagonal, eyes not visible when preserved (Pl. 6 F). Median
antenna 1.5 as long as lateral ones (23 and 12–13 articles, respectively), inserted on
midposterior prostomium, lateral antennae on anterior margin. Palps nearly equal or
shorter than prostomium, fused at their bases but separated all long their length, with-
out sensory cilia. Nuchal organs not visible. Pharynx with a crown of 10 large papillae
and an inner ring of cilia, teeth not visible. Peristomium well dened. Proventricle
long, cylindrical, extending through 6 segments (8–14), with about 62 muscular cell-
rows. Dorsal tentacular cirri longer than ventral ones, with 13–20 and 7–14 articles, re-
spectively. Dorsal cirri slender, similar to tentacular ones, with 14–21 articles, without
noticeable alternating in length. Ventral cirri short, reaching to the tips of parapodial
lobes. Each parapodium with 4–5 aciculae and 5–7 bidentate chaetae with proximal
tooth shorter than distal one.
29
t.a. britayev, t.i. antokhina
Measurements. With up to 164 segments or more, L up to 26.8 mm, WW up to
0.7 mm, WP – 1.3 mm, WC – 1.5 mm.
Coloration. Uniform orange; yellow-whitish when preserved.
Taxonomic remarks. This species closely related I. pseudohaploides San Martin et al.,
2008 from Australia, but differs easely in long body with numerous (usually more than 100)
segments, bidentate blade and elongation in distal part of the shaft.
Ecological notes. Found at 20–23.5 m on Luidia maculata, Archaster angulatus
(Echinodermata: Asteroidea). Usually one worm was found per one host. This is the
second record of an association between syllids and starshes after that reported by
Imajima [1966].
Distribution in Nhatrang Bay. Tre Island, Dam Bay.
General distribution. Vietnam, Nhatrang Bay.
Haplosyllis sp. 1
(Pl. 7 A)
Material examined. Nhatrang Bay: Mun Island (west), 10 m, in Callyspongia sp.
(sponge No 4), coll. Zhadan, 07.10.2011 – 1 cs.
Diagnosis. Body long, cylindrical, wide at proventricle level, gradually tapering to-
wards posterior end. Prostomium pentagonal, with two pairs black eyes. Median antenna
1.5 as long as lateral ones (31 and 23–24 articles, respectively), inserted on middle of
prostomium, lateral antennae on anterior margin. Palps longer than prostomium, broadly
triangular, fused at their bases but separated all long their length. Pharynx funnel-shaped,
with anterior teeth, through 8–10 segments. Proventricle long cylindrical, extending
through 8–9 segments, with 6667 muscular cell-rows. Dorsal tentacular cirri longer
then ventral ones with 36–37 and 19–20 articles, respectively. Dorsal cirri similar to
tentacular cirri, longer in pharyngeal area, alternating in length: rst – long (39 articles),
second short (21 article), third and fourth longer (29 and 50 articles), fth – shorter (33
articles), sixth – long (46 articles), then becaming shorter and slightly alternating in
length. Ventral cirri short, reaching to the tips of parapodial lobes. Chaetae all bidentate,
2–3 per parapodium, anterior ones with conspicuous denticles on upper side of main
fang. Aciculae 4–5 per parapodium, with straight tips.
Measurements. With 65 segments, L – 18 mm, WW – 0.8 mm, WP – 1.2 mm.
Coloration. Orange-brown when preserved (Pl. 7 A).
Taxonomic remarks. This species differs from the other sponge inhabitant Hap-
losyllis sp. 2 in body size, number of aciculae (4–5 instead 2), and proventricle ex-
tending through 8–9 (instead of 5–6) segments with 66–67 (instead of 42) muscular
cell-rows.
Ecological notes. Found at 10 m inside the sponge Callyspongia sp. (Porifera: De-
mospongia).
Distribution in Nhatrang Bay. Mun Island.
General distribution. Vietnam, Nhatrang Bay.
30
chapter 1
Haplosyllis sp. 2
(Pl. 7 B–D)
Material examined. Nhatrang Bay: Mun Island, 10 m, in unidentied sponge N 9,
coll. Zhadan, 14.10.2011 – more than 120 cs and about 20 af.
Diagnosis. Body short, cylindrical, widest at proventricle level, gradually tapering
towards posterior end. Prostomiun rounded, with two pairs of black eyes. Median an-
tenna 1.5 as long as lateral ones (16 and 9 articles, respectively), inserted on middle of
prostomium, lateral antennae on anterior margin. Palps long, broadly triangular, fused at
their bases but separated all long their length. Pharynx extending through 6–7 segments,
slightly thinner than proventricle, with a crown of 10 large papillae and one anterior
tooth. Proventricle long cylindrical, extending through 5–6 segments, with 42 muscular
cell-rows. Dorsal tentacular cirri longer then ventral ones with 17–20 and 8–11 articles,
respectively. Dorsal cirri similar to tentacular ones, longer in pharyngeal area, alternating
in length: rst – long (19–23 articles), second short (5–8 articles), third and fourth slight-
ly longer (10–16 and 12–23 articles), fth short (7 articles), then becoming shorter,
slightly alternating in length. Ventral cirri short, reaching to the tips of parapodial lobes.
Chaetae two per parapodium, bidentate; small chaetae with proximal tooth smaller than
distal one, longer in large chaetae, with conspicuous denticles on upper side of main
fang. Aciculae 2 per parapodium of two types, one with almost 90º bent tip; other with
straight, slightly pointed tip.
Measurements. With up 40 chaetigers, L up to 7.4 mm, WW up to 0.5 mm, WP
0.6 mm, proventricle 1.1 mm long, 0.3 mm wide.
Coloration. Light brown when preserved.
Taxonomic remarks. The species needs further SEM observations to provide precise
character descriptions. However, it differs from the other species found in the area in
having a small size, relatively short proventricle, and only two aciculae per parapodium
of two types.
Ecological notes. Found at 10 m inside an unidentied sponge (Porifera: Demospon-
gia). (Pl. 7 B).
Distribution in Nhatrang Bay. Mun Island.
General distribution. Vietnam, Nhatrang Bay.
Family HESIONIDAE Grube, 1850
Ophiodromus sp. 1
(Pl. 7 E, F)
Material examined. Nhatrang Bay: Tre Island, 23.5 m, on Luidia maculata, coll. An-
tokhina, 02.05.2006 – 5 cs and 7 af; 21 m, on L. maculata, coll. Antokhina, 27.05.2006
7 af; food-market of Nha Trang city, on Archaster angulatus, coll. Antokhina, May
2006 – 4 cs and 7 af.
Diagnosis. Body elongated, with tapered posterior end. Prostomium elliptical, wider
than longer, with three antennae, paired palps, and two pairs of eyes. Median antennal
31
t.a. britayev, t.i. antokhina
furrow absent (Pl. 7 F). Cirrophores of dorsal cirri long, cirrostyles short. Notopodial lobes
and notochaetae absent. Each notopodium and neuropodium with a single aciculum. Pre-
chetal and postchaetal neuropodial lobes triangular, postchaetal one slightly shorter.
Measurements. With up to 55 segments. L up to 11 mm, WW up to 0.6 mm, WP –
1.7 mm, WC – 2.3 mm.
Coloration. Unknown for living animals. Whitish semitransparent in alcohol.
Taxonomic remarks. There are six species of Ophiodromus associated with echi-
noderms. Among them, O. exuosus (Delle Chiaje, 1827) and O. obscurus (Verrill,
1873), were found in Atlantic Ocean, while O. angustifrons (Grube, 1878), O. bunbuku
Uchida, 2004, O. puggettensis (Johnson, 1901) and O. berrisfordi (Day, 1967) occur in
the Pacic. Among the Pacic species, only O. puggetensis has notochaetae. Our species
resemble O. angustifrons, O. berrisfordi and O. bunbuku in the absence of notochaetae,
but differs in the absence of median antennal furrow.
Ecological notes. Found at 21–23 m on the starshes Luidia maculata and Archaster
angulatus (Echinodermata: Asteroidea).
Distribution in Nhatrang Bay. Tre Island, Dam Bay.
General distribution. Nhatrang Bay, Vietnam. This is the rst record of the hesionid
species in association with starshes in Vietnam.
Ophiodromus sp. 2
(Pl. 8 A, B)
Material examined. Nhatrang Bay: Tre Island, 21 m, on Luidia maculata, coll. An-
tokhina, 27.05.2006 1 af; food-market of Nha Trang city, on Archaster angulatus, coll.
Antokhina, May 2006 – 1 cs and 3 af; Tre Island (Dam Bay), 0.1 m, on Archaster typicus,
coll. Britayev, 02.04.2010 – 2 cs.
Diagnosis. Body elongated, with tapered anterior and posterior ends. Prostomium
rounded rectangular, wider than longer, with three antennae, paired palps, and two pairs of
eyes. Median antennal furrow present (Pl. 8 B). Cirrophores of dorsal cirri long, cirrostyles
short. Notopodial lobes absent. Each notopodium and neuropodium with a single aciculum.
Single forked notochaetae present in a few segments. Neuropodial prechaetal lobe with a
triangular projection, postchaetal lobe rounded.
Measurements. With up to 56 segments. L up to 13 mm, WW up to 0.5 mm, WP
1.3 mm, WC – 1.6 mm.
Coloration. Unknown for living animals. Whitish semitransparent in alcohol.
Taxonomic notes. This species is similar to O. pugettensis in the presence of notochae-
tae, but differs in having only one notochaeta per segment in a few segments, and in the pres-
ence of one notopodial acicula in each segment, instead of several notochaetae per bundle in
all segments and 1–5 notopodial aciculae per segment.
Ecological notes. Found at 0.1–21 m on the starshes Luidia maculata, Archaster angu-
latus, and Archaster typicus Müller et Troschel, 1840.
Distribution in Nhatrang Bay. Tre Island, Dam Bay.
General distribution. Nhatrang Bay, Vietnam.
32
chapter 1
Ophiodromus sp. 3
(Pl. 8 C, D)
Material examined. Nhatrang Bay, Mun Island, 2–4 m, on Clypeaster cf. reticu-
lates, coll. Marin, 16.10.2010 – 1 cs and 1 af.
Diagnosis. Body elongated, with tapered anterior and posterior ends. Prostomium
rounded rectangular, with three antennae, paired palps, and two pairs of eyes. Median
antennal furrow absent (Pl. D). Cirrophores of dorsal cirri short, cirrostyles long. Noto-
podial lobes absent. Notopodium with two aciculae, and four forked, slightly serrated
notochaetae. Neuropodium with a single aciculum; neuropodial prechaetal lobe with a
digitiform projection; postchaetal lobe sub-triangular.
Measurements. With up to 48 segments. L up to 8,8 mm, WW up to 0,8 mm, WP –
2,4 mm, WC – 2,8 mm.
Coloration. Unknown for living animals. Whitish semitransparent in alcohol.
Taxonomic notes. This species is similar to O. pugettensis in the presence of noto-
chaetae, but differs in the absence of median antennal furrow and in the presence of a
conspicuous constriction on cirrophores of dorsal cirri. The ciliation on notopodial cir-
rophores mentioned by Uchida [2004] in O. pugettensis was not observed in our speci-
mens. It may also be easily distinguished from the other Pacic sea urchin-associated
species, O. bunbuku, in the presence of notochaetae.
Ecological notes. Found at 2–4 m on the sea urchin Clypeaster cf. reticulates (Lin-
naeus, 1758) (Echinodermata: Echinoidea: Clypeasteridae).
Distribution in Nhatrang Bay. Mun Island.
General distribution. Nhatrang Bay, Vietnam. This is the rst record of the hesionid
species in association with sea urchins in Vietnam.
Ophiodromus sp. 4
(Pls. 8 E, F; 9 A, B)
Material examined. Nhatrang Bay, Mun Island, 2–4 m, on Rhynobrissus hemiaster-
oides, coll. Marin, 18.10.2010 – photo only; Tre Island, Dam Bay, 5 m, sand, on Rhyno-
brissus sp., coll. Bratova, 15.04.2012 – 3 cs.
Diagnosis. Body elongated, with tapered anterior and posterior ends. Prostomium
rounded rectangular, with three antennae, paired palps, and two pairs of eyes. Median
antennal furrow present. Elevated and slightly prolonged dorsal cirri on segments 5, 8,
10, 12, 15, 17, and thereafter on every second segment. Cirrophores of dorsal cirri and
cirrostyles long. Notopodial lobes absent. Notopodium with two aciculae, and without
notochaetae. Neuropodium with a single aciculum; neuropodial prechaetal lobe with a
digitiform projection; postchaetal lobe rounded.
Measurements. With 61 chaetigers, L up to 13 mm, WW up to 1.2 mm, WP – 2.0 mm,
WC – 3.0 mm.
Coloration. Uniformly brown, with transverse white bands across dorsal side of seg-
ments 12, 17, 21, 25, 29, 33, 37, 41, 45, 49, irregularly distributed white spots on the
dorsum of following segments, and continuous white pigmentation on the posterior three
33
t.a. britayev, t.i. antokhina
segments (Pl. 8 E, F). Preserved specimens without pigmentation except small black
spots at bases of parapodia dorsally.
Taxonomic notes. This species differs from the other echinoid associated hesionid,
Ophiodromus sp. 3, in the presence of a median antennal furrow on prostomium, and
absence of notochaetae. It is similar with Ophiodromus sp. 2, having median antennal
furrow, but differs in absence of notochaetae.
Ecological notes. Found at 2–5 m on the sea urchins Rhynobrissus hemiasteroides A.
Agassiz, 1879 and Rhynobrissus sp. (Echinodermata: Echinoidea: Brissidae).
Distribution in Vietnam. Nhatrang Bay, Mun and Tre Islands.
General distribution. Nhatrang Bay, Vietnam.
Parahesione sp.
(Pl. 9 C, D)
Material examined. Nhatrang Bay, River Be estuary, intertidal, sandy silt, in bur-
rows of Upogebia sp., coll. Marin, 18.04.2004 – 1 cs in four fragments.
Diagnosis. Body oval, roundly tapering to anterior and posterior ends. Prostomium
rectangular, wider than longer, with two antennae, two biarticulated palps, and two pairs
of small eyes. Enlarged dorsal cirri on segments 1–5, chaetae absent from segments
1–4. Cirrophores of tentacular segments basally fused. Cirrophores starting at segment 6
(rst with noto- and neuropodia), large, attened, partially covering subsequent segment.
Dorsal cirri with indistinct articulation. All segments except 1–5 with notopodial lobes
and notochaetae. Notopodia at base of cirrophore, small, conical, with prechaetal and
postchaetal lobes; with acicula and numerous (more than 30) capillary chaetae, slightly
serrated. Neuropodial prechaetal lobe elongated, with rounded tip and an unusual digiti-
form projection on postero-dorsal part; with thick acicula and compound chaetae. Neu-
rochaetal blades unidentate, prolonged teeth absent; transparent shaft with spiral internal
structure. Pigidium with two very long anal cirri.
Measurements. With 52 segments. L 19.5 mm, WW 1.5 mm, WP – 4.4 mm,
WC – 5.0 mm.
Coloration. Uniformly bright red when alive, yellowish in alcohol (Pl. 9 C, D)
Taxonomic notes. This species differs signicantly from the unique species of Para-
hesione also inhabiting Upogebia burrows, P. luteola (Webster, 1879), in having eight
pairs of tentacular cirri (instead of six in P. luteola), attened dorsal cirropphores, and a
digitiform dorsal appendage on the neuropodium. The species is preliminary referred to
Parahesione. However, the above-mentioned differences seem to indicate that it could
be placed in a different, new monotypic genus.
Ecological notes. Found in an intertidal zone of a polluted river estuary, among the
rare surviving mangroves, inside a burrow of the decapod Upogebia sp. (Decapoda:
Upogebiidae). This is the second nding of a hesionid inside upogebiid burrows.
Distribution in Nhatrang Bay. River Be estuary.
General distribution. South China Sea, Nhatrang. This is the rst nding of Para-
hesione in the Pacic Ocean.
34
chapter 1
Family SPIONIDAE Grube, 1850
Polydorella dawydof Radashevsky, 1996
(Pl. 9 E, F)
Polydorella dawydof Radashevsky 1996: 686–691, gs. 2–3; Williams 2004:
1348–13–52, gs. 6 –7.
Material examined. Nhatrang Bay: Tre Island 7–10 m, on the sponge Chalinula
nematifera, coll. Savinkin, 15.05.2003 – photo only; Dung Island, 7–12 m, on the sponge
Niphates olemda, coll. Savinkin, 16.05.2003 photo only; Dam Bay, 4.7 m, on the
sponge Haliclona sp., coll. Britayev, 01.05.2012 – 12 specimens.
Diagnosis. Based on Radashevsky [1996]. Small spionid with 14–15 segments. Pro-
stomium bid; caruncle short, reaching anterior margin or middle of segment 2; two
round, white eyes observable in life only. Palps extending posteriorly for 8–12 segments.
Segment 1 without notopodial lobes or notochaetae; with neuropodial lobe, without neu-
rochaetae. Unilimbate capillary neurochaetae of segments 2– 4, 6 and 7 arranged in two
rows; up to six bidentate hooded hooks begin on segment 8, not accompanied by capil-
laries; hooks with approximately right angle between main fang and shaft, with constric-
tion on shaft. Segment 5 slightly modied, about the same size as segments 4 and 6, with
dorsal and ventral fascicles of unilimbate capillary neurochaetae. Modied spines of two
types: rst type, up to 9, distally enlarged, with denticulate edge; second type up to 6,
simple, acicular, with subapical shelf on convex side. Single pair of branchiae on seg-
ment 7. Dorsal ciliary bands from segment 2, continuing to terminal segments. Pygidium
cylindrical, slightly tapering to distal end.
Measurements. With 14–15 segments, 1.5–2.5 mm in length, 0.3–0.4 mm in width.
Coloration. The pigmentation is usually absent but, occasionally, some specimens
may show black-pigmented grains.
Ecological notes. Builds small mud tubes on the surface of the sponges Clathria
(Thalysias) erecta (Thiele, 1899), Chalinula nematifera (de Laubenfels, 1954) (Pl. 9F),
Amphimedon sp., Callyspongia sp., Mycale sp., Niphates olemda (de Laubenfels, 1954)
(Pl. 9E), Petrosia sp., Polymastia sp. and Suberites sp. (Porifera: Demospongia). Recorded
in other areas as symbiont with Xestospongia testudinaria (Lamarck, 1815), Xestospongia
sp., Chalinula sp., Haliclona sp., Niphates sp., Petrosia sp. [Radashevsky 1996; Williams
2004]. Likely combining deposit and suspension feeding, they could benet their host
sponges by keeping their surfaces free of debris [Williams 2004]. It forms aggregations
on the sponge surfaces, with densities of up to 130 individuals cm–2 [Radashevsky 1996].
Distribution in Nhatrang Bay. Tam, Tre Islands.
General distribution. South-Eastern coast of Vietnam; Philippines, Puerto Galera;
Red Sea, Egypt, Hurghada [Radashevsky 1996; Williams 2004].
Discussion
Our studies have increased the number of symbiotic polychaetes occurring in the
coastal waters of Vietnam from 10 up to 26 species (Table 1). The mention of Haplosyllis
35
t.a. britayev, t.i. antokhina
Table 1. Symbiotic polychaetes of Vietnam arranged in alphabetic order. Species reported from Vietnam for the
rst time marked by asterisk*, species reported for the rst time as hosts marked by double asterisks**
Symbiont Host Distribution References
1Asterophilia culcitae Britayev et
Fauchald, 2005
Starshes:
Choriaster granulatus**
Culcita novaeguineae
Echinaster luzonicus**
Euretaster insignis**
Fromia monilis**
Linckia laevigata
Protoreaster nodosus
Unstalked crinoids
Coast of Vietnam, Nhatrang Bay Britayev, Fauchald 2005; this report
2Alcyonosyllis phili
Glasby et Watson, 2001*
Alcyonarian:
Carijoa sp.**
Dendronephthya sp.
Melithaea sp.
Nephthyidae gen. sp.
Northern and eastern coasts of
Australia, Timor Sea, southern
Papua New Guinea, Philippines,
Vietnam.
Glasby, Watson 2001; Glasby, Aguado
2009; this report
3Australaugeneria michaelseni
Pettibone, 1969*
Alcyonarian:
Dendronephthya sp. South-West Australia, Vietnam. Pettibone 1969; this report
4Australaugeneria rutilans
Grube, 1878*
Alcyonarian:
Dendronephthya sp.
Xenia sp.
Red Sea, Persian Gulf, Vietnam,
Philippines, South-West Australia
Pettibone 1969; Wehe 2006;
this report
5Gastrolepidia clavigera
Schmarda, 1861
Holothurians:
Actinopyga echinites
Actinopyga mauritana
Actinopyga sp.
Bohadschia argus
Holothuria atra
Holothuria edulis
Holothuria fuscopunctata
Holothuria hilla
Holothuria leucospilota
Socotra Archipelago, Madagascar,
Sri Lanka, South China Sea,
Papua New Guinea, Bismarck Sea,
Australia, Western Pacic Islands
Uschakov 1982; Hanley 1989; Britayev
et al. 1999; Barnich et al. 2004; Wehe
2006, this report
36
chapter 1
Symbiont Host Distribution References
Holothuria nobilis
Holothuria scabra
Pearsonothuria graeffei
Stichopus chloronotus
Stichopus hermeni
Stichopus horrens
Stichopus mollis
Synapta maculata**
Thelenota ananas
Thelenota anax
6Gaudichaudius cimex
(Quatrefages, 1866)
Anomuran decapods of Diogenes
genus
Arabian Sea, west and east coast
of India, Bay of Bengal, Burma,
Malacca Strait, Indochina, South
China Sea, Hainan Is., Yellow Sea.
Fauvel 1939; Uschakov 1982; Pettibone
1986; Barnich et al. 2004
7Haplosyllides aberrans
Fauvel, 1939
Pontoniin shrimp
Platycaris latirostris
Nhatrang Bay and Spratly
Archipelago, Vietnam, South China
Sea; Sulu archipelago, Indonesia
Martin et al. 2009; this report
8Inermosyllis sp.*
Starshes:
Archaster angulatus
Luidia maculata
Nhatrang Bay, Vietnam This report
9Haplosyllis sp. 1 Sponge Callyspongia sp. Nhatrang Bay, Vietnam This report
10 Haplosyllis sp. 2 Unidentied sponge Nhatrang Bay, Vietnam This report
11 Heteralentia ptycholepis
(Grube, 1878)*Crinoid Phanogenia gracilis**
Gulf of Oman, Mauritius, Vietnam,
Hainan, Hong Kong, Philippines,
Japan, Papua New Guinea,
Chestereld Islands, Samoa
Hanley, Burke 1991; Barnich et al.
2004; this report
12 Hololepidella laingensis
Britayev, Doignon, Eeckhaut, 1999
Starshes:
Acanthaster planci**
Culcita novaeguineae**
Linckia laevigata**
Crinoids:
Papua New Guinea, Coast of
Vietnam
Britayev et al. 1999; Antokhina,
Britayev 2012; this report
37
t.a. britayev, t.i. antokhina
Symbiont Host Distribution References
Capillaster multiradiatus
Cenometra bella
Comaster multidus
Dichrometra agellata
Himerometra robustipinna
Lamprometra palmata
Liparometra regalis
Oxycomanthus bennetti
13 Hololepidella millari
Britayev, Doignon, Eeckhaut, 1999
Starshes:
Acanthaster planci**
Choriaster granulatus** Culcita
novaeguineae**
Euretaster insignis**
Halityle regularis**
Nardoa frianti**
Protoreaster nodosus**
Linckia laevigata
Linckia guildingi
Crinoid:
Comanthus alternans
Papua New Guinea, Coast of
Vietnam Britayev et al. 1999; Antokhina,
Britayev 2012; this report
14 Hololepidella sp.*Scleractinian coral Galaxea
astreata Vietnam, Nhatrang Bay This report
15 Ophiodromus sp. 1*
Starshes:
Archaster angulatus
Luidia maculata
Vietnam, Nhatrang Bay This report
16 Ophiodromus sp. 2*
Starshes:
Archaster angulatus
Archaster typicus
Luidia maculata
Vietnam, Nhatrang Bay This report
17 Ophiodromus sp. 3*Sea urchin Clypeaster cf.
reticulatus Vietnam, Nhatrang Bay This report
18 Ophiodromus sp. 4*
Sea urchins:
Rhynobrissus hemiasteroides
Salmacis sp.
This report
38
chapter 1
Symbiont Host Distribution References
19 Ophthalmonoe pettiboneae
Petersen et Britayev, 1997 Polychaete Chaetopterus sp. Indonesia, Banda Sea; Vietnam,
Nhatrang Bay
Petersen, Britayev 1997; Britayev,
Martin 2005; this report
20 Paradyte crinoidicola
(Potts, 1910)*
Crinoids:
Amphimetra ensifera**
Amphimetra tessellata**
Capillaster multiradiatus
Cenometra bella**
Colobometra perspinosa
Comanthus alternans
Comanthus briareus
Comanthus gisleni**
Comanthus parvicirrus**
Comaster multidus
Comaster nobilis**
Comaster schlegelii
Comatella nigra**
Comatella stelligera
Comatula purpurea
Dichrometra agellata
Himerometra robustipinna
Lamprometra palmata**
Liparometra regalis**
Oligometra carpenteri
Oligometra serripinna
Oxycomanthus bennetti
Oxycomanthus pinguis**
Phanogenia gracilis**
Petasometra helianthoides
Stephanometra indica
Stephanometra tenuipinna**
Tropiometra carinata
Widely distributed in the Indo-West
Pacic
Hanley 1984; Pettibone 1969; Wehe
2006; Barnich et al. 2004; Britayev et al.
1999; this report
21 Paradyte levis*
Alcyonarian:
Dendronephthya sp.
In tubes of polychaete Euniphysa
tubifex
Red Sea, Suez Bay, Arabian Sea,
Oman Gulf, Vietnam, Japan
Marenzeller 1902; Imajima 1997; Wehe
2006; this report
22 Parahesione sp. In burrow of decapod
Upogebia sp. Vietnam, Nhatrang Bay This report
39
t.a. britayev, t.i. antokhina
Symbiont Host Distribution References
23 Pararctonoella aphthalma
(Gallardo, 1968) On ophiuoid Vietnam, Nhatrang Bay Gallardo 1967; Pettibone 1996
24 Pottsiscalisetosus praelongus
(Marenzeller, 1902)
Starshes:
Archaster angulatus**
Luidia maculata**
Indian Ocean, Vietnam, Hainan,
Japan
Pettibone 1969; Barnich et al. 2004;
Antokhina, Britayev 2012; this report
25 Polydorella dawydof
Radashevsky, 1996
Sponges:
Amphimedon sp.
Callyspongia sp.
Chalinula nematifera
Chalinula sp.
Clathria (Thalysias) erecta
Haliclona sp.
Mycale sp.
Niphates olemda
Niphates sp.
Petrosia sp.
Polymastia sp.
Rhaphidophlus erectus
Suberites sp.
Xetospongia testudinaria
Xetospongia sp.
South-Eastern coast of Vietnam,
Philippines: Puerto Galera, Red
Sea: Egypt, Hurghada
Radashevsky 1996; Williams 2004;
this report
26 Uncopolynoe corallicola
Hartmann-Schröder, 1960*Alcyonarian coral Red Sea: Gulf of Suez, Vietnam:
Nhatrang Bay
Hartmann-Schröder 1960; Wehe 2006;
this report
40
chapter 1
spongicola by Fauvel [1939], has not been here considered, since this corresponds to a
recently revised species-complex [Lattig, Martin 2009] and so it is not possible to attribute
Fauvel’s report to any recently described species without a re-examination of its material.
About 360 polychaete species were reported from Vietnam coasts [Gallardo 1967],
and this number did not increase substantially until now. Accordingly, about 7.8% of
the known species are symbionts, this being a much higher percentage than that calcu-
lated for the World Oceans by Martin, Britayev [1998], i.e. 2.3%, and even higher than
the 5.5% of the Solomon Islands [Gibbs 1971]. The vast majority of symbionts belong
to Polynoidae (14 species), followed by Syllidae (5 species) and Hesionidae (5 spe-
cies). This agrees well with the general family distribution among symbiotic polychaetes
[Martin, Britayev 1998]. In a similar way, most species were associated with echino-
derms (53.8%), followed by cnidarians (23.0%), sponges (11.5%), crustaceans (7.7%),
and other polychaetes (3.8%) (Fig. 10 A).
The symbiotic polychaete fauna from tropical Australian coastal waters includes
23 species. Likely this underestimates the real diversity, but roughly reects the cur-
rent state of the art. Accordingly, there are only ve common species in Vietnam and
Australia: Australaugeneria michaelseni, A. rutilans, Alcionosyllis phili, Gastrolepidia
clavigera, Paradyte crinoidicola. This low similarity may be caused by real differences
in faunal composition, but certainly there must be an important component dealing with
the applied expertise and sampling methods.
Most Nhatrang Bay species were associated with echinoderm and cnidarian hosts
(Fig. 10 A), while the Australian symbionts were much more regularly distributed among
host taxa (Fig. 10 B). There were as much as three time more echinoderm hosts in Nha-
trang Bay than in the Australian coasts, while there were substantially less sponge and,
particularly polychaete, associated species in our samples. This is certainly a sampling
biass, as our main sampling efforts were addressed to holothurians, starshes, feather
stars and octocorals, while sponges and, especially, tubes and burrows of polychaetes
and other sedentary animals were less thoroughly studied. For example, we have not
found the Lepidasthenia species that are typical inhabitantns of tubes and burrows of
tube-dwelling polychaetes, sipunculans and balanoglossids [Gibbs 1969], while ve spe-
cies of this genus were recorded in the Gulf of Tonkin (northern Vietnam) but without
mentioning whether they were symbionts or not [Uschakov 1982].
Accordingly, we may tentatively conclude that the diversity of symbiotic polychaetes
in the Bay of Nhatrang is relatively high, but also we are certain that the list of symbiotic
species we have compiled is rather far from their real diversity. Further studies of dif-
ferent geographical regions, as well as of poorly investigated or overlooked hosts (e.g.,
sponges, tube dwelling polychaetes, echinoids, ophiuroids), will increase substantially
the diversity of symbiotic polychaetes of Vietnam.
Acknowledgements
We would like to thank the directors of the Coastal Branch of the Russian-Vietnam-
ese Tropical Centre, Dr. V.K. Nezdoly, N.L. Filichev, Dr. Tran Cong Huan and Trang
Thanh Kuang for the logistics of the different expeditions that made possible our work
41
t.a. britayev, t.i. antokhina
in Vietnam. We are grateful to O.V. Savinkin and Dr. I. Marin for eld assistance and
underwater photographs. Special thanks to Dr. D. Martin, for careful revision of the
manuscript and useful editorial advices. We are also indebted to Mrs. A. Zalota for her
help with linguistic corrections. The study was supported by the Russian Foundation for
Basic Research, under grants 12–05–00239–а and 12–04–33267_mol_a_ved.
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Uschakov P.V. 1972. New records of the Eulepethes hamifer (Grube) (Polychaeta, fam.
Eulepethidae Chamberlin). Issledovaniya fauny morei, 10 (18): 329–333 [In Rus-
sian].
Uschakov P.V. 1982. Fauna of the USSR. 2(1): Polychaetes. Polychaetes of the Suborder
Aphroditiformia of the Arctic Ocean and the Northwestern Part of the Pacic, Fami-
lies Aphroditidae and Polynoidae. [In Russian]. NAUKA, Academy of Sciences of
the USSR, Zoological Institute, Moscow, 272 pp.
Wehe T. 2006. Revision of the scale worms (Polychaeta: Aphroditoidea) occurring in
the seas surrounding the Arabian Peninsula. Part I: Polynoidae. Fauna of Arabia, 22:
23–197.
Williams J.D. 2004. Reproduction and morphology of Polydorella (Polychaeta: Spi-
onidae), including the description of a new species from the Philippines. Journal of
Natural History, 38: 1339–1358.
45
t.a. britayev, t.i. antokhina
Симбиотические полихеты залива Нячанг, Вьетнам
Т.А. Бритаев1, Т.И. Антохина1
РЕЗЮМЕ. В работе представлен обзор фауны мелководных симбиотических полихет за-
лива Нячанг (Вьетнам). В результате исследований, выполненных в период с 1985 по 2012
гг., обнаружено 24 вида полихет из 4 семейств. Таким образом, число видов симбиоти-
ческих полихет, известных для прибрежных вод Вьетнама, возросло с 10 до 26. Приве-
ден список видов полихет, для каждого из которых, указаны синонимия, материал, рас-
ширенный диагноз, таксономические и экологические замечания, распространение. Для
большинства видов полихет даны цветные фотографии in situ. Большая часть симбион-
тов относится к семейству Polynoidae (13 видов), остальные – к семействам Syllidae (5 ви-
дов), Hesionidae (5 видов) и Spionidae (1 вид). Хозяевами для полихет являлись иглокожие
(53.8%), кишечнополостные (23.0%), губки (11.5%), ракообразные (7.7%), а также другие
виды полихет (3.8%). Показано, что фауна симбиотических полихет Вьетнама значительно
отличается от австралийской. В то же время, дальнейшие исследования таких групп хозя-
ев, как губки, восьмилучевые кораллы, морские ежи, офиуры и др. могут значительно рас-
ширить наши знания, о разнообразии симбиотических полихет Вьетнама.
1 Институт Проблем Экологии и Эволюции им. А.Н. Северцова, Российская Академия Наук,
Ленинский проспект, 33, Москва, Россия, 119071, britayev@yandex.ru
55
t.a. britayev, t.i. antokhina
Plate 10. Distribution of symbiotic polychaetes among main host taxa in the coastal waters of Vietnam
(a) and Australia (b).
... Cn Candidella helminophora (Nutting, 1908) (Nutting 1908, Cairns 2009 Gorekia crassicirris (Willey, 1902) E Abatus (Pseudabatus) nimrodi (Koehler, 1911) (Miranda & Brasil 2014) Cn Madrepora oculata (Miranda & Brasil 2014) Harmothoe imbricata (Linnaeus, 1767) Ec Lissomyema exilii (Anker et al. 2005) Harmothoe ruthae Miranda & Brasil, 2014 Cn Enallopsammia rostrata (Miranda & Brasil 2014) Cn Solenosmilia variabilis (Miranda & Brasil 2014) Cn Lophelia pertusa (Miranda & Brasil 2014) Cn Madrepora oculata (Miranda & Brasil 2014) Cn Errinia sp. (Miranda & Brasil 2014) Harmothoe vesiculosa Ditlevsen, 1917 Cn Lophelia pertusa (Linnaeus, 1758) (Fiege & Barnich 2009) Cn Madrepora oculata (Fiege & Barnich 2009) Harmothoe oculinarum (Storm, 1879) P Eunice norvegica (Linnaeus, 1767) (Sato et al. 2001, Hong et al. 2017 Hesperonoe japonensis Hong, Lee & Sato, 2017 D Upogebia major (Hong et al. 2017) D Austinogebia narutensis (Sakai, 1986) (Hong et al. 2017) Hesperonoe laevis Hartman, 1961Ec Listriolobus pelodes Fisher, 1946(Anker et al. 2005) Heteralentia ptycholepis (Grube, 1878) Cr Phanogenia gracilis (Hartlaub, 1893) (Britayev & Antokhina 2012) Hololepidella boninensis Nishi & Tachikawa, 1998 O Ophiocoma sp. (Nishi & Tachikawa 1998) Hololepidella laingensis Britayev, Doignon & Eeckahaut, 1999 Cr Amphimetra tessellata (Müller, 1841) (Mekhova & Britayev 2015) Cr Comatula solaris Lamarck, 1816(Mekhova & Britayev 2015 Cr Capillaster gracilicirrus AH Clark, 1912(Mekhova & Britayev 2015 Cr Zygometra comata AH Clark, 1911(Mekhova & Britayev 2015 A Acanthaster planci (Linnaeus, 1758) (Britayev & Antokhina 2012) A Culcita novaeguineae (Britayev & Antokhina 2012) A Linckia laevigata (Britayev & Antokhina 2012) Cr Capillaster multiradiatus (Linnaeus, 1758) (Britayev et al. 1999) Cr Dichometra flagellata (Britayev et al. 1999) Cr Lamprometra palmata (Müller, 1841) (Britayev et al. 1999) Cr Himerometra robustipinna (Carpenter, 1881) (Britayev et al. 1999 Cr Anneissia bennetti (Müller, 1841) (Britayev et al. 1999) Cr Comaster sp. ...
... (Miranda & Brasil 2014) Harmothoe vesiculosa Ditlevsen, 1917 Cn Lophelia pertusa (Linnaeus, 1758) (Fiege & Barnich 2009) Cn Madrepora oculata (Fiege & Barnich 2009) Harmothoe oculinarum (Storm, 1879) P Eunice norvegica (Linnaeus, 1767) (Sato et al. 2001, Hong et al. 2017 Hesperonoe japonensis Hong, Lee & Sato, 2017 D Upogebia major (Hong et al. 2017) D Austinogebia narutensis (Sakai, 1986) (Hong et al. 2017) Hesperonoe laevis Hartman, 1961Ec Listriolobus pelodes Fisher, 1946(Anker et al. 2005) Heteralentia ptycholepis (Grube, 1878) Cr Phanogenia gracilis (Hartlaub, 1893) (Britayev & Antokhina 2012) Hololepidella boninensis Nishi & Tachikawa, 1998 O Ophiocoma sp. (Nishi & Tachikawa 1998) Hololepidella laingensis Britayev, Doignon & Eeckahaut, 1999 Cr Amphimetra tessellata (Müller, 1841) (Mekhova & Britayev 2015) Cr Comatula solaris Lamarck, 1816(Mekhova & Britayev 2015 Cr Capillaster gracilicirrus AH Clark, 1912(Mekhova & Britayev 2015 Cr Zygometra comata AH Clark, 1911(Mekhova & Britayev 2015 A Acanthaster planci (Linnaeus, 1758) (Britayev & Antokhina 2012) A Culcita novaeguineae (Britayev & Antokhina 2012) A Linckia laevigata (Britayev & Antokhina 2012) Cr Capillaster multiradiatus (Linnaeus, 1758) (Britayev et al. 1999) Cr Dichometra flagellata (Britayev et al. 1999) Cr Lamprometra palmata (Müller, 1841) (Britayev et al. 1999) Cr Himerometra robustipinna (Carpenter, 1881) (Britayev et al. 1999 Cr Anneissia bennetti (Müller, 1841) (Britayev et al. 1999) Cr Comaster sp. (Britayev et al. 1999) Cr Comaster multifidus (Müller, 1841) (Britayev et al. 1999) Hololepidella millari Britayev, Doignon & Eeckahaut, 1999 A Acanthaster planci (Britayev & Antokhina 2012) A Choriaster granulatus (Britayev & (Britayev et al. 1999) A Linckia laevigata (Britayev et al. 1999) A Linckia guildingi (Britayev et al. 1999) Hololepidella nigropunctata (Horst, 1915) O Ophiothrix (Acanthophiothrix) purpurea (Britayev et al. 1999) Hololepidella sp. ...
... (Miranda & Brasil 2014) Harmothoe vesiculosa Ditlevsen, 1917 Cn Lophelia pertusa (Linnaeus, 1758) (Fiege & Barnich 2009) Cn Madrepora oculata (Fiege & Barnich 2009) Harmothoe oculinarum (Storm, 1879) P Eunice norvegica (Linnaeus, 1767) (Sato et al. 2001, Hong et al. 2017 Hesperonoe japonensis Hong, Lee & Sato, 2017 D Upogebia major (Hong et al. 2017) D Austinogebia narutensis (Sakai, 1986) (Hong et al. 2017) Hesperonoe laevis Hartman, 1961Ec Listriolobus pelodes Fisher, 1946(Anker et al. 2005) Heteralentia ptycholepis (Grube, 1878) Cr Phanogenia gracilis (Hartlaub, 1893) (Britayev & Antokhina 2012) Hololepidella boninensis Nishi & Tachikawa, 1998 O Ophiocoma sp. (Nishi & Tachikawa 1998) Hololepidella laingensis Britayev, Doignon & Eeckahaut, 1999 Cr Amphimetra tessellata (Müller, 1841) (Mekhova & Britayev 2015) Cr Comatula solaris Lamarck, 1816(Mekhova & Britayev 2015 Cr Capillaster gracilicirrus AH Clark, 1912(Mekhova & Britayev 2015 Cr Zygometra comata AH Clark, 1911(Mekhova & Britayev 2015 A Acanthaster planci (Linnaeus, 1758) (Britayev & Antokhina 2012) A Culcita novaeguineae (Britayev & Antokhina 2012) A Linckia laevigata (Britayev & Antokhina 2012) Cr Capillaster multiradiatus (Linnaeus, 1758) (Britayev et al. 1999) Cr Dichometra flagellata (Britayev et al. 1999) Cr Lamprometra palmata (Müller, 1841) (Britayev et al. 1999) Cr Himerometra robustipinna (Carpenter, 1881) (Britayev et al. 1999 Cr Anneissia bennetti (Müller, 1841) (Britayev et al. 1999) Cr Comaster sp. (Britayev et al. 1999) Cr Comaster multifidus (Müller, 1841) (Britayev et al. 1999) Hololepidella millari Britayev, Doignon & Eeckahaut, 1999 A Acanthaster planci (Britayev & Antokhina 2012) A Choriaster granulatus (Britayev & (Britayev et al. 1999) A Linckia laevigata (Britayev et al. 1999) A Linckia guildingi (Britayev et al. 1999) Hololepidella nigropunctata (Horst, 1915) O Ophiothrix (Acanthophiothrix) purpurea (Britayev et al. 1999) Hololepidella sp. ...
... The Polychaeta (including myzostomids) is one of the most diverse taxa of crinoid symbionts (Britayev and Mekhova 2011). Most polychaetes associated with the crinoid belong to the order Myzostomida, with about 170 species, (Eeckhaut and Lanterbecq 2005), while only seven species belong to another taxonomic group, family Polynoidae or scaleworms (order Phyllodocida) (Martin and Britayev 1998;Britayev et al. 1999;Britayev and Antokhina 2012). Martin and Britayev (1998) reported that the scaleworm Paradyte crinoidicola (Potts 1910) is the most common and numerous species among crinoid symbionts. ...
... Martin and Britayev (1998) reported that the scaleworm Paradyte crinoidicola (Potts 1910) is the most common and numerous species among crinoid symbionts. This symbiont inhabits on more than 30 species of shallow-water crinoids hosts, with relatively high (14 to 62%) infestation prevalence when each infested host harboured from 1 to 7 polychaetes, and multiple infestations with 2 or 3 polychaetes per host are common (Zmarzly 1984;Britayev and Antokhina 2012;Britayev and Mekhova 2014). ...
... The species was described from Maldive Islands (Potts 1915) and was reported later in many localities all over the tropical Indo-West Pacific (Horst 1917;Okuda 1936;Hanley 1984;Fishelson 1974;Zmarzly 1985;Tchesunov et al. 1989;Morton and Mladenov 1992;Hanley and Burke 1991;Britayev et al. 1999;Barnich et al. 2004;Wehe 2006;Deheyn et al. 2006;Britayev and Antokhina 2012;Britayev et al. 2016;Virgili et al. 2020). Surprisingly, among that localities the coast of Indian subcontinent is absent. ...
Article
Full-text available
Polychaetes symbiotic with echinoderms (sea stars, brittle stars, feather stars, basket stars, sea cucumbers, and sea urchins) possess remarkable diversification in the Indo-Pacific. In the present study, one species of polychaetes symbiotic with crinoids are recorded based on collections from selected Islands Kadmat, Amini, Suheli, Kavaratti, and Agatti of Lakshadweep Archipelago. Paradyte crinoidicola were recorded for the first time from the Indian subcontinent. The present study reports the occurrence of a symbiotic scaleworm, P. crinoidicola, from Lakshadweep. This is the first record of the species from the Indian subcontinent we recommend further exploration of the sub-shallow coral reef areas of Lakshadweep.
... Moreover, concerning L. squamatus, data from OBIS do not reflect its real distribution, as most Pacific records, derived from Uschakov [381], are no included. Symbiotic species also seem to have very wide distributions across tropical Indo-Pacific waters, such as Hololepidella nigropunctata (Horst, 1915) [409] or Paradyte crinoidicola (Potts, 1910) [410][411][412]. However, at least the first seems to be a species-complex (Britayev and Fiege, unpublished data). ...
... In addition, these widely polyxenous species often show specialized morphological adaptations, like the hooked ventral chaetae of P. crinoidicola [435] or the ventral sucker-like lobes of G. clavigera [426,436]. The degree of polyxeny of some species is still not well-known and for instance, Asterophilia culcitae Britayev & Fauchald, 2005 [383], was always reported as exclusive associate of asteroids until found on crinoids [412] and holothurians [437]. ...
Article
Full-text available
Phyllodocida is a clade of errantiate annelids characterized by having ventral sensory palps, anterior enlarged cirri, axial muscular proboscis, compound chaetae (if present) with a single ligament, and of lacking dorsolateral folds. Members of most families date back to the Carboniferous, although the earliest fossil was dated from the Devonian. Phyllodocida holds 27 well-established and morphologically homogenous clades ranked as families, gathering more than 4600 currently accepted nominal species. Among them, Syllidae and Polynoidae are the most specious polychaete groups. Species of Phyllodocida are mainly found in the marine benthos, although a few inhabit freshwater, terrestrial and planktonic environments, and occur from intertidal to deep waters in all oceans. In this review, we (1) explore the current knowledge on species diversity trends (based on traditional species concept and molecular data), phylogeny, ecology, and geographic distribution for the whole group, (2) try to identify the main knowledge gaps, and (3) focus on selected families: Alciopidae, Goniadidae, Glyceridae, Iospilidae, Lopadorrhynchidae, Polynoidae, Pontodoridae, Nephtyidae, Sphaerodoridae, Syllidae, Tomopteridae, Typhloscolecidae, and Yndolaciidae. The highest species richness is concentrated in European, North American, and Australian continental shelves (reflecting a strong sampling bias). While most data come from shallow coastal and surface environments most world oceans are clearly under-studied. The overall trends indicate that new descriptions are constantly added through time and that less than 10% of the known species have molecular barcode information available.
... Asterophilia consists of two species: A. carlae Hanley, 1989 andA. culcitae Britayev &Fauchald, 2005, the former occurring from Fiji to Southeast Asia (Hanley 1989) and the latter from Southeast Asia to Japan (Britayev & Antokhina 2012, Britayev & Fauchald 2005, Okinawa General Bureau 2013. Both species are well-known symbionts of a wide range of Asteroidea, although they have recently been found with an unidentified crinoid (Britayev & Antokhina 2012, Martin & Britayev 1998, Britayev & Fauchald 2005. ...
... culcitae Britayev &Fauchald, 2005, the former occurring from Fiji to Southeast Asia (Hanley 1989) and the latter from Southeast Asia to Japan (Britayev & Antokhina 2012, Britayev & Fauchald 2005, Okinawa General Bureau 2013. Both species are well-known symbionts of a wide range of Asteroidea, although they have recently been found with an unidentified crinoid (Britayev & Antokhina 2012, Martin & Britayev 1998, Britayev & Fauchald 2005. The whitish or yellowish spots on the elytra, the whitish inflated tips of the antenna, and the tentacular and dorsal cirri have been suggested to mimic the tube foot of the host asteroids to avoid being detected by predators (Britayev & Fauchald 2005, Hanley 1989. ...
Article
Full-text available
The Polynoidae, commonly known as "scale-worms" due to the scale-like elytra on the dorsal surface, contains many species living in symbioses with other invertebrates. Most of these symbionts are host-specific, but some have a wide range of hosts. The genus Asterophilia includes two species living in shallow subtropical to tropical waters in the Pacific Ocean as ectosymbionts of asteroids and, more rarely, crinoids. Here, we recorded Asterophilia culcitae from asteroid hosts (Culcita novaeguineae, Linckia laevigata, L. guildingi, and Leiaster leachi) and, for the first time, from holothurian hosts [Stichopus chloronotus, Holothuria atra, H. (Stauropora) pervicax, and Bohadschia argus] along warm Japanese Pacific coasts. The cytochrome c oxidase subunit I sequences of the specimens from holothurians did not differ significantly from those on asteroid hosts, proving that A. culcitae has a wide host range across three different echinoderm classes: asteroids, holothurians, and crinoids. The general body color of A. culcitae was constantly reddish (female) or whitish (male), regardless of the host body color, although a previous study suggested that it differs in accordance with the host body color. However, we found that one individual from a holothurian host showed a different color pattern: A. culcitae typically shows three whitish or yellowish elytral mounds that have been suggested to mimic the tube foot of the asteroid hosts, whereas one individual from S. chloronotus had reddish brown translucent mounds, which we suggest might be cryptic on its holothurian host.
... Most of them are free living and inhabit almost every benthic ecosystem, from littoral environments down to bathyal and abyssal depths, and play a key role in benthic communities as they take part in the decay of organic sediment matter or in the transfer of nutrients to the water column (Hutchings 1998). Further, about 370 polychaete species exhibit a symbiotic lifestyle, either obligate or facultative (Britayev and Antokhina 2012). ...
... However, the new species can be distinguished of it by the lack of constriction in its notopodial cirrophores, and by the papilliform instead of long, triangular projection on prechaetal lobe. Although most hesionids are free-living organisms that can be found as carnivores in consolidated or soft substrates, including the interstitial environment, at least 14 species have developed symbiotic life strategies (Martin and Britayev 1998;Britayev and Antokhina 2012;Britayev et al. 2013;Chim et al. 2013). Usually, symbiont hesionids act as facultative commensals of echinoderms (Rizzo and Salazar-Vallejo 2014), but in some cases the relationship is highly host-specific (Britayev et al. 2013). ...
Article
In 2004, Uchida published a partial revision of the family Hesionidae in Japan, where four new genera were proposed and 13 species newly described. However, both the holotype fixations and their depository numbers were omitted in the original article, rendering the names unavailable after Article 16.4 of the International Code of Zoological Nomenclature. In this work, the names are made available by clarifying the depository of the types. We provide diagnoses for the four new genera (Synsyllidia Uchida, Uncopodarke Uchida, Parahesiocaeca Uchida, and Ichthyohesione Uchida) and 13 new species (Synsyllidia alternata Uchida, Oxydromus brevipodius Uchida, O. bunbuku Uchida, O. constrictus Uchida, O. fauveli Uchida, O. longifundus Uchida, O. okudai Uchida, O. parapallidus Uchida, Heteropodarke kiiensis Uchida, Uncopodarke intermedia Uchida, Microphthalmus itoi Uchida, Parahesiocaeca japonica Uchida, and Ichthyohesione gorgasiae Uchida). A key to species of Hesionidae recorded to date from Japan is also presented.
... Marine annelids are typically considered free living, although a large number of species are involved in more or less close symbiotic associations (mainly ectosymbiotic) with other annelids, sponges, cnidarians, molluscs, crustaceans, echinoderms and fishes (e.g. Martin & Britayev, 1998Rouse & Pleijel, 2001;Britayev & Antokhina, 2012;Andrews et al., 2015;Hernández-Alcántara et al., 2015;Molodtsova et al., 2016;Igawa et al., 2017;Goto et al., 2016Goto et al., , 2017. Less commonly, marine annelids are also found as endoparasites of other animals, such as the recently described Proceraea exoryxae Martin, Nygren & Cruz-Rivera, 2017 tunnelling within the tunic of the ascidian Phallusia nigra Savigny, 1816 or the holopelagic Alciopina parasitica Claparède & Pancer, 1867 reported in the gastrovascular canals of the ctenophore Hormiphora plumosa M. Sars, 1859 (Claparède & Panceri, 1869;Fauvel, 1923; see Clark (1956) and Martin & Britayev (1998) for detailed revisions). ...
Article
A new genus of ectoparasitic marine annelids living on ctenophores, Ctenophoricola gen. nov., is described and its feeding behaviour, reproduction and developmental stages are discussed. Its unusual morphology challenged its placement within the known marine families. However, analyses of mitochondrial and nuclear sequence data showed the new genus as member of the Alciopini, a group of holopelagic annelids included within the Phyllodocidae. Ctenophoricola masanorii sp. nov. from Japan and Ctenophoricola rousei sp. nov. from the Canary Islands (Spain) are described. A third species from the Gulf of California is not formally described because the specimens are in poor condition. The new genus is characterized by having: 1) two distinctive body regions, the anterior with reduced parapodia lacking chaetae, and the posterior with long parapodia and chaetae and 2) a pair of large, elongate lensed eyes. These eyes are here described using histology and 3D reconstruction based on a Californian specimen. The two new species mainly differ in colour pattern, shape of parapodia, number of chaetae and body ciliation.
... Thus, it was not unexpected that five out of the six species of Haplosyllis living in association with the four study sponge species were also undescribed. The Vietnam area seems to be rich in symbiotic polychaetes, according to the numerous species of Haplosyllis present there (10,37), although this can be related to the large number of studies carried out in this area. We recorded the presence of host-specific spicules in representative samples from all sponge-associated worms. ...
Article
Full-text available
The symbiotic lifestyle represents a fundamental cryptic contribution to the diversity of marine ecosystems. Sponges are ideal targets to improve understanding the symbiotic relationships from evolutionary and ecological points of view, because they are the most ancient metazoans on earth, are ubiquitous in the marine benthos, and establish complex symbiosis with both prokaryotes and animals, which in turn also harbor their own bacterial communities. Here, we study the microbiomes of sponge-polychaete associations and confirm that polychaetes feed on their host sponges. The study worms select and enrich part of the sponge microbiome to shape their own species-specific bacterial communities. Moreover, worm microbiome diversity runs parallel to that of its food host sponge. Considering our results on symbiotic polychaetes and previous studies on fishes and mammals, diet appears to be an important source of bacteria for animals to shape their species-specific microbiomes.
... In addition, this is the first report on a The total number of hosts and symbionts and indices of infestation are marked in bold The first record of symbiosis between a palaeonemertean (Nemertea) and echinoderms (Echinodermata) symbiotic relationship between nemerteans and echinoids, and the third reported symbiotic association between nemerteans and echinoderms. This report substantially improves our knowledge on the diversity of echinod symbionts in Vietnamese coastal waters that was limited until now to the single paper on the symbiotic fauna of regular sea urchin Salmacis bicolor L. Agassiz (Agassiz & Desor, 1846) (Britayev et al. 2013), and taxonomic papers describing symbiotic polychaetes and copepods associated with several heart and sea urchins (Britayev and Antokhina 2012;Maran et al. 2017). ...
Article
Full-text available
The associations between Cephalotrichella echinicola Chernyshev et al., 2019, and the heart urchin Metallia spp. described here represent the first report on the symbiotic relationship between palaeonemerteans and echinoderms. In addition, it is the first report of a symbiotic relationship between nemerteans and heart urchins, and the third known association between nemerteans and echinoderms. Numerous specimens of C. echinicola were found associated with heart urchins Metallia spatagus and M. sternalis in four localities of the Bay of Nhatrang, Vietnam, South China Sea, at 6–10 m water depth buried in the coarse sand to a sediment depth of 10–20 cm. The infestation prevalence and mean intensity were higher in M. spatagus,than in M. sternalis, 77% and 68%, and 10 and 3 nemertean individuals per heart urchin host, respectively. Each infested host harbored from 1 to 49 nemertean individuals, and multiple infestations with three or more individuals per host were very common. Based on the high level of prevalence, absence of the nemertean on the other sand-dwelling heart urchins in the area, the complete absence of the nemerteans in the sandy bottom of the bay and in the sediment surrounded infested host individuals, and no reaction by the host to the nemertean’s movement on the host’s surface we consider this a symbiotic association.
Article
Extreme sexual size dimorphism is one of the most striking phenomena in evolutionary biology. While the origin has been well discussed and some causes have been suggested, the evolutionary history remains unclear. We found a new species of deep‐sea scale worm (Annelida: Polynoidae) inside gastropod shells, either empty or occupied by hermit crabs collected at 140–306 m in depth, Mie Prefecture, Japan. This highly specialized habitat, together with the fact that it has never been found free‐living, led us to consider the scale worm to be an obligate symbiont. The species is characterized by males being dwarf, with their minute bodies (ca. one‐fourth the length of females) always riding on the dorsal side of females, being thus the first case of extreme sexual size dimorphism in scale worms. Based on a detailed morphological, histological, and molecular phylogenetic approach we are here describing a new species, Eunoe issunboushi sp. nov.
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Polynoid scale-worms have been found living as commensals with deep-water antipatharians (commonly known as black corals) in the Potiguar Basin, off Rio Grande do Norte State, Northeastern Brazil. In this paper two polychaete species and four black corals species are redescribed. Benhamipolynoe cf. antipathicola and Parahololepidella cf. greeffi, and the black coral Stylopathes adinocrada Opresko, 2006 are recorded for the Southwestern Atlantic. Benhamipolynoe cf. antipathicola was first described from off New Zealand and the Malay Archipelago, as symbiont with the black coral Stylopathes tenuispina (Silberfeld, 1909). It was later reported for the North Atlantic, off Florida, associated with Stylopathes columnaris (Duchassaing, 1870). In our study, B. cf. antipathicola was found in association with the black coral S. adinocrada. Parahololepidella cf. greeffi was first described as a free-living from shallow waters off São Tomé and Cabo Verde Islands, West Africa, and later reported as symbiont with the black coral Tanacetipathes cf. spinescens in the same location. Our data expand both the geographical distribution and the host range of this species which is reported for the first time as symbiont with Tanacetipathes barbadensis (Brook, 1889), T. tanacetum (Pourtalès, 1880) and T. thamnea (Warner, 1981) in Brazil. The aim of this study is to discuss commensal associations between two species of scale-worm polynoids and black corals found in the Southwestern Atlantic, and also reporting their global distribution. Finally, we provided an updated list of the commensal polynoids and their black coral hosts.
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