Revision of Scale Worms in Arabian Seas

FAUNA OF ARABIA 22: 23–197
Date of publication: 18.12.2006
Revision of the scale worms (Polychaeta:

Aphroditoidea) occurring in the seas surrounding
the Arabian Peninsula. Part I: Polynoidae
Thomas Wehe
A b s t r a c t : The study presented here is the first in a series on the scale worms of the seas surrounding the Arabian Peninsula:
Suez Canal, Red Sea, Gulf of Aden, Arabian Sea, Gulf of Oman, The Gulf. Based on type specimens and additional published and
unpublished specimens, this is the first revision of the Polynoidae that are currently recognised from this area. Detailed descriptions
and figures are provided as well as identification keys to all taxa, and taxonomic problems are discussed. Discussions of the distribu-
tion, faunal affinities, Lessepsian migration and endemism are given in a zoogeographic account.
The occurrence of 44 species belonging to 23 genera and seven subfamilies is confirmed. Twenty-three species represent new
records for one or more of the respective seas. Four new species are described: Harmothoe marerubrum n. sp., Lepidonotus polae n.
sp., Parahalosydnopsis arabica n. sp., Pararctonoella marginopapillata n. sp.; and Uncopolynoinae n. subf. is introduced. Harmothoe
grisea, Lepidasthenia nuda and Lepidonotus impatiens are redescribed. Bouchiria, Iphione reticulata, Lepidonotus impatiens meridionalis
and L. australiensis are considered junior synonyms. Harmothoe vesicudenta is transferred to the genus Malmgreniella. The diagnoses
of 10 genera are emended.
The fauna documented is closely related to that of the Indo-West Pacific with at least 30 species out of 44 in common. Two spe-
cies are likely to represent Lessepsian and two Anti-Lessepsian migrants. For the present, 10 species (23 %) are considered endemic
to the area.
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.Uncopolynoinae �� Pararctonoella marginopapillata � Parahalosydnopsis arabica �
Bouchiria, �� .Lepidonotus impatiens � Lepidasthenia nuda � Harmothoe grisea ��
�� address:
Author’s �� .�� L. australiensis � Iphione reticulata, Lepidonotus impatiens meridionalis
Dr. Thomas
.�� Wehe,)Ruprecht-Karls-Universität,
(�� Heidelberg, Institut für Zoologie,
�� . Malmgreniella ��Abteilung 5: Morphologie
�� Harmothoe & Ökologie,
vesicudentaIm��
Neuenheimer Feld 230, D 69120 Heidelberg, Germany and Forschungsinsitut & Naturmuseum Senckenberg, Senckenberganlage
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25, D 60325 Frankfurt a.M., Germany; email: t.wehe@urz.uni-heidelberg.de
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T. Wehe
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Lepidonotus polae � Harmothoe marerubrum :�� (��) �� .��
.Uncopolynoinae �� Pararctonoella marginopapillata � Parahalosydnopsis arabica �
Bouchiria, �� .Lepidonotus impatiens � Lepidasthenia nuda � Harmothoe grisea ��
�� .�� L. australiensis � Iphione reticulata, Lepidonotus impatiens meridionalis
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Introduction
Aphroditoid polychaetes or scale worms form one of the major taxa within the polychaetes. They
are currently divided into Acoetidae Kinberg 1858, Aphroditidae Kinberg 1856, Eulepethidae
Chamberlin 1919, Pholoidae Kinberg 1857, Polynoidae Kinberg 1856 and Sigalionidae Kinberg
1856 (see e.g. Pettibone 1982, Glasby et al. 2000, Barnich & Fiege 2003). This is generally true
as regards their species and genera richness as well as their frequent occurrence, especially at lower
latitudes, as was reported by Fauvel (1932): “As is usual in tropical seas, the families best repre-
sented are the Aphroditidae [all scale worm taxa sensu Fauvel] and Nereidae ...”; and also by Knox
(1957): “The majority of the species ... [in the Indo-Pacific] ... belong to the families Polynoidae,
Nereidae, and Eunicidae”.
However, even though considerable efforts were made in the past to revise polynoid scale
worms (see below), as this family is the richest in genera within the Polychaeta and is probably
exceeded only by the Syllidae in terms of species richness, the systematics are still confused and in
a state of flux.
In order to contribute to a better understanding of the systematics of this group, the present
work represents the first revision of the Polynoidae covering all the species known to occur in the
seas surrounding the Arabian Peninsula, i.e. the Suez Canal, Red Sea, Gulf of Aden, Arabian Sea,
Gulf of Oman and The Gulf (fig.1). Until recently, no comprehensive information on the poly-
chaetes of the seas surrounding the Arabian Peninsula was available. To fill in this gap and in order
to lay the basis for the intended revision of scale worms presented here, Wehe & Fiege (2002)
compiled an annotated checklist of all the polychaete species reported from this region. All the
specimens reported and listed in Wehe & Fiege (2002) from around the Arabian Peninsula and
available in zoological collections worldwide have been re-examined. This includes a considerable
number of additional and freshly collected specimens which have not been studied or reported
before, in total about 800 specimens.
The species confirmed here are described and figured in detail in the systematic section below,
together with taxonomic remarks where necessary. In addition, identification keys are provided to
all the subfamilies currently recognised worldwide as well as to all the genera and species occurring
in the study area.
Apart from the taxonomic revision of polynoid scale worms, another objective of this work
has been to consider the faunistic affinities among the seas surrounding the Arabian Peninsula as
well as to the adjacent Mediterranean Sea and to the Indo-West Pacific. The zoogeographic section
therefore contains a discussion of the faunal affinities, depth distribution, rate of endemism and
Scale worms occuring in the seas surrounding the Arabian Peninsula. I. Polynoidae 25
Fig. 1: Map of the area investigated showing the various seas surrounding the Arabian Peninsula.
occurrence of Lessepsian migrants. Affinities to the adjacent Mediterranean Sea and to the Indo-
West Pacific, respectively, are also discussed, summarising the current state of knowledge on these
topics.
History of polychaete research in the Arabian region
The specimens examined during this study were obtained during a number of expeditions going
back to Napoleon’s campaign in Egypt over 200 years ago. Some specimens more than 100 years
old have never been examined or published, e.g. specimens from the Red Sea collected by Eduard
Rüppell (1822‑27, 1830‑34) or by the Austrian “Pola”-Expeditions in 1895‑98. Specimens col-
lected more recently by Fiege and the author from remote locations such as the Sanganeb Atoll,
Sudan, or the Socotra Archipelago, Yemen, are also reported for the first time.
In order to provide as much information as possible on the Polynoidae from the seas sur-
rounding the Arabian Peninsula and to highlight possible sources of specimens for future research,
26 T. Wehe
the following information is given on the history and origin of the specimens examined here. This
account therefore gives a brief overview of the expeditions and other research activities carried out
in the Arabian marine region during which collections of polychaetes in general and Polynoidae in
particular were obtained and subsequently reported in important publications. It should be em-
phasised that much more research has been carried out and numerous other references are available
dealing with polychaetes from this region (Wehe & Fiege 2002).
Suez Canal. — In order to study the fauna and flora of the Suez Canal, “The Cambridge
Expedition to the Suez Canal” set out in 1924 (Fox 1926). In addition to other biota, numerous
polychaetes were collected and subsequently described in the papers by Fauvel (1927) and Potts
(1928). Focusing more on the migration of organisms through the Suez Canal, some sampling was
carried out as part of the extensive survey undertaken by the Hebrew University - Smithsonian
Institution Joint Program “Biota of the Red Sea and the Eastern Mediterranean” (Por et al. 1972),
with occasional sampling up to 1973. Within the framework of this survey, samples of polychaetes
were taken from 1967‑1969 by “The Hebrew University Expeditions to the Suez Canal” with
additional sampling carried out by the Norwegian team of H. Brattstrom & J.P. Tassen in 1973.
Some results on the Polychaeta were published by Ben-Eliahu (1972 a, 1972 b), and a monograph
on the Serpulidae of the Suez Canal is currently being prepared by Ben-Eliahu & ten Hove (Ben-
Eliahu, pers. comm.).
Red Sea. — The oldest available descriptions of polychaete species from the Red Sea are those
by Savigny, prepared as part of the outstanding “Description de l’Égypte”. However, the authorship
credit cannot be given to Savigny (1822). A manuscript for this work was probably already finished
by Savigny in 1815 (Fauchald 1992), and both Cuvier (1817) and Lamarck (1818) quoted from
this manuscript. Most of Savigny’s species were thus mentioned for the first time in Lamarck (1818)
and have to be cited as Savigny in Lamarck (1818). The specimens described by Savigny were col-
lected during the “French Military Expedition” to Egypt led by Napoleon Bonaparte in 1798‑1801.
Unfortunately, it appears that most of the polychaete specimens collected and described by Savigny
are now lost. In his remarks on the annelids of the Museum in Paris, Grube (1870) noted that
Savigny’s specimens had never been incorporated into the museum’s collections. Nevertheless, some
specimens have survived, as mentioned in the recently published catalogue of type specimens in the
collections of the Muséum National d’Histoire Naturelle, Paris (Solís-Weiss et al. 2004) (see also
the redescription given here of Lepidonotus impatiens (Savigny in Lamarck, 1818)).
At the time that Savigny’s work was being published, three German explorers travelled to Africa
and the Red Sea. Christian Gottfried Ehrenberg and Friedrich Wilhelm Hemprich travelled through-
out the Orient from Libya to Lebanon between 1820 and 1825, studying marine life and collecting
an enormous number of specimens of all kinds in the Red Sea in the years 1824 and 1825 (Bolling
1976). According to a compilation by Ehrenberg (Bolling 1976), among the total of 34000 speci-
mens and 3987 species collected there were 261 specimens of annelids belonging to 67 species. The
polychaetes were later examined and published by Eduard Grube, mainly in Grube (1869).
Almost at the same time, Eduard Rüppell of the Senckenberg Museum, Frankfurt a.M.,
made his first journey to north-eastern Africa and the Sinai Peninsula from January 1822 to Au-
gust 1827, followed by a second expedition from December 1830 to February 1834 (Mertens
1949). Rüppell made very extensive collections of various animal groups, including marine species
from the Red Sea, which he sent back to the Senckenberg Museum. Descriptions of the marine
invertebrates that he collected were published by Rüppell & Leuckart (1828 [1830]). However,
polychaetes were not considered in these accounts and some polynoid scale worms still available in
the collections of the Senckenberg Museum and The Natural History Museum, London (resulting
from an exchange), are reported here for the first time.
From 1895 to 1896 and 1897 to 1898, the first extensive deep-sea studies were carried out in
the Red Sea by the Austrian vessel S.M.S. “Pola”, led by Franz Steindachner (Schefbeck 1991).
During these expeditions, both oceanographic and biological observations and collections were
made, and Emil von Marenzeller studied the corals, polychaetes and echinoderms. Whilst three
publications on corals appeared, the results on the other two groups were never published (Stagl
et al. 1996). Stagl et al. (1996) also provided a list of polychaete species identified by Marenzeller
taken from his inventory book. As in the case of Rüppell mentioned above, the Polynoidae origi-
nating from the “Pola” expeditions are described here for the first time.
The British zoologist Cyril Crossland began to investigate the fauna and flora of the Sudanese
Red Sea in 1904. From 1905 to 1922 he was head of the Sudanese Pearl Fisheries, and in 1930 he
founded the first Red Sea research station at Ghardaqa (= Hurghada) in Egypt where he stayed
until 1938 (Head 1987, Vine & Schmid 1987). During this long period he made extensive studies
and collections of marine animals. The results on the polychaetes that he collected from this region
were published in e.g. Crossland (1923, 1924) and Monro (1939 b, 1939 c, 1939 d).
During 1927‑1929, Robert Ph. Dollfus travelled through Egypt and made extensive collec-
tions of animals. Specimens of several animal groups were published in a series of papers in the
“Mémoires présentés à l’Institut d’Egypte” (Dollfus 1933). Polychaetes originating mainly from
the Gulf of Suez and the Gulf of Aqaba were described by Fauvel (1933).
Oceanographic studies were carried out in the Red Sea, but mainly in the Northern Indian
Ocean, by the research vessel H.E.M.S. “Mabahiss” during “The John Murray Expedition” (see
also below) from 1933 to 1934, under the scientific leadership of R.B. Seymour Sewell (Sewell
1935, Rice 1986). Monro (1937) examined and described the polychaetes obtained by this expe-
dition.
From 1951 to 1952 polychaete annelids were collected mainly from Abulat Island and the
Farasan Archipelago (Saudi Arabia) by M. Drach, M.M. Cherbonnier and C. Mercier during a
campaign of the “Calypso”, under the command of Jacques Yves Cousteau. The resulting specimens
were examined and described by Fauvel (1955).
During a field study in the vicinity of the Marine Biological Station at Ghardaqa in 1956,
polychaetes were collected by Adolf Remane and Erich Schulz. Descriptions of the species were
prepared by Banse (1959) and Hartmann-Schröder (1960); more general results on ecology were
published by Remane & Schulz (1964). In 1965 and 1967 Volker Storch, a student of Remane’s,
also visited the station in order to study polychaetes. The results of these studies were published in
several papers (Storch 1966, 1967 a, 1967 b, Storch & Niggemann 1967).
Fauvel (1957, 1958) identified and reported numerous polychaete species, which were col-
lected mainly in the Gulf of Aqaba by the Hebrew University of Jerusalem and H. Steinitz.
In 1962 and 1965, two Israeli expeditions to the Southern Red Sea took place within the
multi-national program of the “International Indian Ocean Expedition” (see below), known as “Is-
rael South Red Sea Expeditions”. Collections were made in the Dahlak Archipelago, Eritrea (Oren
1962, Lewinsohn & Fishelson 1967). The Polychaeta obtained were studied by Day (1965) and
Fishelson & Rullier (1969).
A major contribution to the polychaete fauna occurring around the Sinai Peninsula was made
by Amoureux et al. (1978) based on collections made in the Gulf of Suez and the Gulf of Aqaba
between 1968 and 1973 by L. Fishelson, Ch. Lewinsohn, Y. Dafni and M. Goren.
N.M. Ben-Eliahu studied the polychaete cryptofauna from the rims of intertidal vermetid
reefs in the Gulf of Aqaba and compared them with those occurring along the Mediterranean coast
of Israel. The polychaetes from the Gulf of Aqaba were collected from Wadi Kabila in spring 1973.
The faunistic results of this comparative study were published in a series of papers (Ben-Eliahu
28 T. Wehe
1975 a, 1975 b, 1976 a, 1976 b, 1977 a, 1977 b, 1977 c), and ecological aspects were dealt with
in Ben-Eliahu & Safriel (1982), Ben-Eliah et al. (1988), Safriel & Ben-Eliahu (1991) and Ben-
Eliahu (1991 a).
Against the background of intended mining activities in the deep Red Sea, large-scale ecological
studies of the deep-sea benthos were carried out on behalf of the “Saudi Sudanese Commission for
the Exploitation of the Red Sea Resources”. Samples were obtained by the German research vessels
RV “Sonne”, during the cruise MESEDA I (Metallifeorus Sediments, Atlantis-II-Deep) in 1977, and
the RV “Valdivia”, during the second cruise MESEDA II in 1979 (Thiel 1980). The deep-sea ben-
thic polychaetes collected during these expeditions were studied by Rosenfeldt (1989).
From January to September 1987, cruise no. 5 of the German research vessel RV “Meteor”
of the project MINDIK (“Mittelmeer-Indik”) took place, leading the research vessel all the way
through the Red Sea and into the Western Indian Ocean (Nellen et al. 1996 a, 1996 b). The
objectives were the investigation of oceanographic, geological as well as biological features. The
polychaetes collected during this cruise have not yet been described, and specimens belonging to
the family Polynoidae are reported here for the first time.
Within the framework of a joint project on “Comparative Ecological Analysis of Biota and
Habitats in Littoral and Shallow Sublittoral Waters of the Sudanese Red Sea”, studies were carried
out between 1991 and 1993 by the Forschungsinstitut & Naturmuseum Senckenberg and the
Faculty of Marine Science and Fisheries, Port Sudan, focusing on fishes, decapod crustaceans and
polychaetes. Polychaete specimens were mainly collected during the second expedition to the San-
ganeb Atoll and Wingate Reef from September to October 1992 by D. Fiege, and a list of prelimi-
nary identifications was given in an unpublished report (Fiege 1993). The polynoid scale worms
obtained during this expedition are described here.
Gulf of Aden. — Records in the literature of polychaetes from the Gulf of Aden proper are
scarce. However, extensive studies at the beginning of the 20th century focused on the small west-
ernmost part of this region, the Gulf of Tadjoura (Djibouti). In the late 1890s until 1904, the
French scientists F. Jousseaume, M. Coutière, E. Perrier and Ch. Gravier collected specimens from
this region. The results of these studies were published in a fine series of contributions by Gravier
(1899 a, 1899 b, 1900, 1902, 1904 a, 1904 b, 1905 a, 1905 b, 1905 c, 1905 d, 1905 e, 1905 f,
1906 a, 1906 b, 1906 c, 1906 d, 1906 e, 1908).
Pelagic polychaete specimens collected in 1933 and 1934 from the same region by J.-L. Dan-
tan were described by Fauvel (1951 a, 1951 b, 1951 c, 1951 d).
Arabian Sea and Gulf of Oman. — Polychaetes collected in the Arabian Sea and to a lesser extent
from the Gulf of Oman originate mainly from the following studies. Over several decades beginning in
1881, the Royal Indian marine survey ship “Investigator” made soundings in the Arabian Sea as well as
in the Bay of Bengal (Alcock 1902). In the course of numerous cruises, dredgings and faunistic studies
were also carried out. The polychaetes collected during these cruises and held in the Indian Museum/
Zoological Survey of India, Calcutta, were described in a voluminous monograph by Fauvel (1932).
Additional polychaetes from the Arabian Sea and the Gulf of Oman were collected during the
“John Murray Expedition” (1933‑34), which was mentioned above.
An enormous quantity of marine biota was collected during the multi-national “Indian Ocean
Expedition” from 1959 to 1965. 42 vessels from 14 nations were involved in this project, surveying
the whole Indian Ocean including the Gulf of Aden, the Arabian Sea and the Gulf of Oman (Wen-
zel 1978). Some of the polychaete annelids collected were described by Hartman (1974 a, 1974 b).
Since 1998, an international and multi-disciplinary survey has been conducted in the Socotra
Archipelago (Yemen), within the framework of the long-term United Nations Development Pro-
gramme (UNDP)/Global Environment Facility (GEF) project “Conservation and Sustainable Use
of Biodiversity of Socotra Archipelago”. Marine habitat, biodiversity and fisheries surveys have
been carried out under the leadership of the Senckenberg Research Institute (Krupp et al. 2002).
In addition to other marine animals, M. Apel and the present author collected large quantities of
polychaetes in 1999 and 2000. The polynoids represent the first polychaetes to be described from
the Socotra Archipelago, apart from a single freshwater nereid species that was described from Abd
al-Kuri Island by Fiege & Van Damme (2002).
The Gulf. — The first major contribution to the polychaetes occurring in The Gulf was pub-
lished by Fauvel (1911). He studied the collections made in Kuwait, Bahrain and Būshehr (=Bu-
shire) by M.N. Bogoyawlenski in 1902.
The second important work on polychaetes from this region was again published by Fauvel
(1919), this time reporting on specimens collected by M.Ch. Pérez and M.J. Bonnier along the
coasts of the Arabian Peninsula in 1901, but mainly originating from The Gulf.
In 1937 G. Thorson and in 1938 B. Løppenthin collected large numbers of polychaetes dur-
ing fishery investigations carried out by H. Blegvad on behalf of the Iranian government. In con-
trast to the collections from The Gulf mentioned above, specimens were mainly obtained along
the Iranian rather than the Arabian coast, and to a lesser extent also from the Gulf of Oman. The
description of these specimens resulted in the third major contribution to the polychaete fauna of
The Gulf by Wesenberg-Lund (1949).
Mohammad (1970 a, 1970 b, 1970 c, 1971, 1972 a, 1972 b, 1973, 1980) reported on poly-
chaetes collected in more recent times, e.g. by the Japanese research vessel “Umitaka-Maru”.
In recent years, several research projects have been carried out in The Gulf, for three main rea-
sons: (1) to assess the possible consequences of intended oil-drilling programmes, (2) to evaluate the
ecological consequences of the 1991 oil spill during the Gulf War, and (3) to contribute to the estab-
lishment of a marine sanctuary (e.g. Abuzinada & Krupp 1994, Krupp & Jones 1993, Krupp et al.
1996). Unfortunately, the polychaete specimens collected during such monitoring and consultation
programmes have not usually been published in scientific papers but rather in unpublished reports
which have a restricted circulation and are therefore often difficult to access. Studies carried out e.g.
by ARAMCO (Arabian American Oil Company) resulted in voluminous reports by O’donnell
(1981, 1982, 1984, 1986) which are not formally published. The Senckenberg Museum houses a
rather large collection of polychaetes from The Gulf, only some of which have so far been listed in
a project report (Fiege 1992). Once again, the Polynoidae belonging to these collections, as well
as additional specimens which were kindly made available by P. Garwood (Identichaet, Newcastle-
upon-Tyne), D. George (The Natural History Museum, London) and M. Stachowitsch (Institute of
Ecology and Conservation Biology, University of Vienna), are described here for the first time.
The history and current state of the taxonomy of the Polynoidae Kinberg, 1856
The first known Polynoidae were described as belonging to the genus Aphrodita Linnaeus, 1758
(the first scale worm genus to be described), as Aphrodita squamata Linnaeus, 1758 (= Lepidonotus
squamatus) and A. imbricata Linnaeus, 1767 (= Harmothoe imbricata); also as A. cirrhosa (= Gat-
tyana cirrhosa) and A. lepidota (= Harmothoe imbricata) by Pallas (1766), and as A. punctata (=
Polynoe squamata?) and A. cirrata (= Harmothoe imbricata) by Müller (1776). Between the 1770s
and 1853, several more polynoid species were described within the genus Aphrodita.
With Lepidonotus Leach 1816 and Polynoe Savigny in Lamarck 1818, the first new genera that
were later referred to the Polynoidae were introduced. But it was Kinberg (1856) who made the first
major advances, by erecting several further polynoid genera, i.e. Antinoe, Halosydna, Harmothoe,
30 T. Wehe
Hermadion, Iphione, all of which are still valid, and by providing a classification of major scale worm
groups, amongst them the families “Iphionea” and “Polynoina”. According to §36 of the ICZN,
these are the first available family names and the authorship should thus be credited to Kinberg
(1856). Until recently many authors have not realised this and the credit for erecting the family name
Polynoidae has been repeatedly given to Malmgren (1867) by e.g. Hartman (1959), Day (1967 a),
Fauchald (1977), Hanley & Burke (1991 a), Hartmann-Schröder (1996), Fauchald & Rouse
(1997), or to Grube (without date) by Fauvel (1923, 1953), whilst the authorship of Iphioninae was
attributed to Baird, 1865 i.a. by Pettibone (1986 a) and Hanley & Burke (1991 a). However, Muir
(1982) clearly pointed out that the authorship for the Polynoinae and so for the Polynoidae as well
belongs to Kinberg, but it appears that this was not adopted until Barnich & Fiege (2000).
A major taxonomic problem was the designation of the type species for Polynoe, which was
discussed in detail by Muir (1979). According to this author, Polynoe was described by Savigny in
Lamarck (1818) to include six species. Unfortunately, Savigny did not designate a type species and
all of his species were subsequently referred to other genera within the family Polynoidae. In 1822,
Savigny again listed the same six species with an additional seventh one, P. scolopendrina. This species
was accepted by later authors as type species for the genus (e.g. Hartman 1959, Day 1967, Fauchald
1977). However, according to Muir (1982), this was in contravention of §76 h of the ICZN, since
a nominal species which was not included in the original description of the genus cannot be desig-
nated as the type species of that genus. To avoid the further taxonomic confusion which would have
arisen from the strict application of this article, and to fulfil the principle of stability expressed by the
ICZN, the Commission followed a request by Muir (1979) and set aside all prior designations and
designated P. scolopendrina Savigny, 1822 as the type species of the genus (Melville 1984).
Since the time of Kinberg (1856), many new genera and species have been described and
the Polynoidae now comprises by far the greatest number of genera (about 165, see below) of
all the polychaete families and also one of the highest number of species (> 800). However, the
information available in the literature about the number of genera and species in the Polynoidae
varies considerably. Hartman (1959) listed 72 genera and 503 species in her catalogue, Fauchald
(1977) registered 94 genera with 569 species, Pettibone (1982) mentioned about 120 genera and
600 species, and according to Hutchings (2000) there are 165 genera and 748 species. Following
a census carried out for this paper, 167 genera and about 815 species are currently recorded.
Naturally, this large number of taxa also encompasses a wide range of characters and character
states, respectively. Several attempts have therefore been made to establish subgroups within the
family, which has led to 20 subfamilies that are currently recognised, i.e.: Polynoinae Kinberg,
1856, Iphioninae Kinberg, 1856, Lepidonotinae Willey, 1902, Macellicephalinae Hartmann-
Schröder, 1971, Bathyedithinae Pettibone, 1976, Bathymacellinae Pettibone, 1976, Macellicepha-
loidinae Pettibone, 1976, Macelloidinae Pettibone, 1976, Polaruschakovinae Pettibone, 1976,
Admetellinae Uschakov, 1977, Gesiellinae Muir, 1982, Lepidonotopodinae Pettibone, 1983 a,
Branchipolynoinae Pettibone, 1984, Branchiplicatinae Pettibone, 1985 b, Branchinotogluminae
Pettibone, 1985 c, Lepidastheniinae Pettibone, 1989 b, Arctonoinae Hanley, 1989, Acholoinae
Pettibone, 1996 b, Eulagiscinae Pettibone, 1997, Vampiropolynoinae Marcus & Hourdez, 2002. A
further subfamily Uncopolynoinae n. subfam. is added here.
Muir (1982) also suggested reducing the Polyodontidae Augener, 1918 (= Acoetidae Kinberg,
1856) to subfamily level within the Polynoidae and gave several arguments in support of this. How-
ever, his suggestion was not followed by Pettibone (1989 c), who revised the family Acoetidae and
whose opinion has been generally accepted (e.g. Fauchald & Rouse 1997, Hutchings 2000).
The phylogenetic relationships within the Polynoidae are still largely unknown. Early sugges-
tions were put forward by Seidler (1923) and Uschakov (1977), but these ideas are now outdated
since several subfamilies were not included or were subsequently erected and contained many new
genera and species, especially deep-sea forms.
The subfamilial and generic positions of many species are still in a state of flux as is the inclusive-
ness of these taxa, which is in part due to the inconsistent use of characters in distinguishing genera
and subfamilies. Hanley (1989) for instance erected the subfamily Arctonoinae to separate several
genera formerly placed in the Polynoinae which are characterised by the presence of terminoventrally
or ventrally inserted ceratophores of the lateral antennae (see below), tentaculophores without cha-
etae and deeply notched neuropodia. The characters used by Hanley (1989) are subject to dispute
and some of the genera referred to the Arctonoinae by Hanley (1989) or Hartmann-Schröder
(1996), respectively, are retained in the Polynoinae by Barnich & Fiege (2001, 2003), e.g. Adyte
Saint-Joseph, 1899, Malmgreniella Hartman, 1967, Paradyte Pettibone, 1969 a and Subadyte Pet-
tibone, 1969 a, as are the three latter genera and Hololepidella Willey, 1905, in the present paper. On
the other hand, Uncopolynoe Hartmann-Schröder, 1960 was considered to belong to the Polynoinae
(Fauchald 1977), but with the erection of more and new subfamilies and the ongoing refinement
of diagnoses, the genus can no longer be referred to any of the currently accepted subfamilies. It is
therefore referred here to the Uncopolynoinae n. subfam. It is anticipated that further revision of the
Polynoidae may lead to several new subfamilies and/or to the exclusion and elevation of some to new
families. As regards the taxonomic position on the generic level, the genera Harmothoe and Malm-
greniella may serve as a good example where the assignment of species to one or the other genus needs
to be revised, e.g. see Malmgreniella vesicudenta (Hanley & Burke, 1991) in the present paper.
A review of the literature reveals that the same names have often been or are still applied to
scale worm taxa in different senses. For example, Day (1967 a) considered the family Aphroditi-
dae Kinberg 1856 (as Aphroditidae Malmgren, 1867) to include all the scaled polychaete species
and his subfamily Polynoinae is consistent with the family Polynoidae Kinberg, 1856 as used by
e.g. Pettibone (1982), Hanley & Burke (1991 a), Barnich & Fiege (2003). In contrast to Day
(1967 a), however, the Polynoinae (= Harmothoinae in Hanley & Burke (1991 a)) is used only for
a distinct group of species within the Polynoidae Kinberg, 1856 e.g. by Barnich & Fiege (2003)
and Hanley & Burke (1991 a). In the present paper, the Polynoidae is used sensu Pettibone
(1982), Hanley & Burke (1991 a), Fauchald & Rouse (1997) and Barnich & Fiege (2003).
Even though our knowledge of the systematics of the Polynoidae is far from being complete, a
considerable number of revisions of parts of the taxon have been published in the past. Some of the
more important works are: Seidler (1923), Pettibone (1969 a, 1969 c, 1975, 1976, 1977), Muir
(1982), Pettibone (1983 a, 1983 b, 1984, 1985 a, 1985 b, 1985 c, 1985 d, 1986 a), Hanley
(1987), Hanley & Burke (1988), Hanley (1989), Hanley & Burke (1989), Pettibone (1989 b),
Hanley & Burke (1990), Pettibone (1991 a, 1991 b), Hanley & Burke (1991 a, 1991 b),
Hanley (1991, 1993), Pettibone (1993 a, 1993 b, 1995 a, 1995 b, 1996 a, 1996 b, 1997), Bar-
nich et al. (2000), Barnich & Fiege (2000, 2001, 2003, 2004), Barnich et al. (2004).
Several more regional studies also include extensive information, e.g. Horst (1917), Fauvel
(1923, 1953), Day (1967 a), Pettibone (1970), Uschakov (1982), Amaral & Nonato (1982),
Tebble & Chambers (1982), Weston (1984), Ruff (1995), Hartmann-Schröder (1996), Cham-
bers & Muir (1997) and Imajima (1997).
Anatomy and physiology
There is not a great deal of data on the morphology and physiology of polynoid scale worms in the
literature. The most comprehensive information with reference to the original sources is provided
32 T. Wehe
by Hutchings (2000). Storch (1968, 1969) investigated in detail the muscle system of polychae-
tes. As representatives of the Polynoidae, Lepidonotus squamatus, Antinoella sarsi (Malmgren, 1865)
and Iphione muricata were studied. As regards the circulatory system and oxygen transport, blood
pigments are usually absent. Extracellular haemoglobin, however, was reported in Branchipolynoe
by Hourdez & Toulmond (1998). The release of excretory and sexual products in polynoids
through nephridia or mixonephridia, respectively, were studied in more detail by Darboux (1899),
Goodrich (1945) and Daly (1972). The sensory organs of polychaetes including Polynoidae
were reviewed e.g. by Storch & Schlötzer-Schrehardt (1988). Horridge (1963) found up to
1000‑1200 sensory cells on the dorsal cirri of Harmothoe. The elytra investigated by Anton-Erx-
leben (1981) and Storch & Alberti (1995) were found to be equipped with sensory cells as well.
The eyes of Arctonoe vittata for example have been studied (Singla 1975). The cephalic nervous
system and the innervation of head appendages were carefully studied by Orrhage (1991) in Lepi-
donotus squamatus and Harmothoe longisetis (Grube, 1863). As in other polychaete families such
as the Chaetopteridae, some polynoid scale worms are able to exhibit bioluminescence. Extensive
studies on the physiology of this phenomenon were carried out by i.a. Nicol (1953, 1954, 1957 a,
1957 b, 1957 c, 1958), Nicolas (1977), Nicolas et al. (1978, 1981, 1982), Herrera (1979),
Bilbaut (1980 a, 1980 b), Anctil (1987). About 15 species are reported to show bioluminescence,
amongst them for instance Harmothoe impar (Johnston, 1839), Harmothoe imbricata, Malmgren-
iella castanea (McIntosh, 1876), or Acholoe astericola (Delle Chiaje, 1841 a).
Biology and ecology
The Polynoidae are cosmopolitan. They are found in polar regions (e.g. Hartman 1978) as well as
in the tropics and subtropics (i.a. Hanley & Burke 1991 a; this paper) or in boreal regions (e.g.
Pettibone 1963, Hartmann-Schröder 1996, Chambers & Muir 1997). Their vertical distribu-
tion reaches from the intertidal to the deep-sea. Branchinotogluma burkensis Pettibone, 1989 a for
example was found at 3660 m, Bathykermadeca hadalis (Kirkegaard, 1956) at 6720 m (Pettibone
1976), Bathyedithia berkeleyi (Levenstein, 1971) at 7000 m (Pettibone 1976), and Macellicepha-
loides grandicirra Ushakov, 1955 a [1969] even at 9950 (!) m (Pettibone 1976). Our knowledge of
the ecology and biology is sparse, based on the investigation of a very few species.
In general polynoids share a benthic life style. There are only a few pelagic forms such as An-
tinoella sarsi (see Pettibone 1963). However, according to Sarvala (1971), this species is benthic
but performs vertical movements during the night. Støp-Bowitz (1991) reports several pelagic
species. Some of them, however, represent only pelagic larval stages, e.g. Lepidasthenia grimaldii
(Marenzeller, 1892), whilst for others, e.g. Harmothoe hubrechti (McIntosh, 1900), an obligate
pelagic life pattern is only assumed.
Polynoids are generally found in a marine environment. Some species, however, are capable of
tolerating moderate to low salinities and are found in brackish environments, for example Antinoe-
lla sarsi in the north-eastern Baltic Sea at a salinity of 6 ‰ (Sarvala 1971) or Harmothoe charlottae
Hutchings & Murray, 1984 and Paralepidonotus ampulliferus (Grube, 1878) in Australian estuaries
at salinities of about 33‑35 ‰ (Hutchings & Murray 1984).
High abundances of polynoids may occur. Sarvala (1971) reported a maximum of 800‑1300 in-
dividuals of Antinoella sarsi per m2 at the time of settlement, with 35‑50 remaining after about one-
and-a‑half years. On the basis of size distribution and growth rates, the same author concluded that
two years of age is very rarely exceeded in this species. Britaev et al. (2002) report life spans of up to
five years in Lepidonotus squamatus and up to three years in Harmothoe imbricata.
Polynoids are considered to be carnivorous. They prey on small crustaceans such as copepods,
cladocerans, amphipods, but also on echinoderms, polychaetes, gastropods, rotifers, turbellarians,
hydroids or sponges (Darboux 1899, Blegvad 1914, Meunier 1930, Korringa 1951, Sarvala
1971, Fauchald & Jumars 1979, Pettibone 1982, Hartmann-Schröder 1996). Commensal
species may feed on the faecal matter of their hosts (Ajeska & Nybakken 1976). Macginitie &
Macginitie (1968), however, classified four different feeding categories for commensal species.
Algae also play an important role in the nutrition of polynoids according to Rauschenplat (1901).
Food composition seems to be correlated to size and age (Sarvala 1971). Planktotrophic larvae
feed on phytoplankton (Cazaux 1968), and the feeding mechanisms of planktotrophic polynoid
trochophores were investigated by Phillips & Pernet (1996). The role of the head appendages for
capturing prey in adult specimens was studied under laboratory conditions by Daly (1973).
The sexes are separate. Reproduction and development have only been investigated very spo-
radically. Thorson (1936, 1946) gave an extensive account of the larval development of marine in-
vertebrates in general, also providing information on a few polynoid larvae. According to Wilson
(1991), 10 out of 11 species have free-spawning and planktotrophic larvae, whereas only H. im-
bricata carries out brood care. Even earlier, Harmothoe impar, Harmothoe antilopes McIntosh, 1876
and Arctonoella sinagawaensis (Izuka, 1912) (as Gattyana) were reported by Schroeder & Hermans
(1975) to carry out brood care. Investigations into ecology and reproduction were also conducted
in Antinoella sarsi (see Meunier 1930, Sarvala 1971) or Arctonoe Chamberlin, 1920 (see Pernet
2000). However, H. imbricata is the most popular species to be used for observations, e.g. by Daly
(1972, 1973), Garwood (1980), Clark (1988), Watson et al. (2000), and Gaudron & Bentley
(2002). Recently the reproductive biology and population structure of the deep-sea hydrothermal
vent species Branchipolynoe seepensis Pettibone, 1986 b was investigated (Jollivet et al. 2000).
A common phenomenon amongst many polynoids is their commensal life pattern. According
to Martin & Britayev (1998), no fewer 160 species, that is about 20 % of all known species, ex-
hibit this life pattern. Associations have been described with echinoderms (Gibbs 1969, Pettibone
1969 b, Hartmann-Schröder 1981, Hanley 1984, Ruff 1991, Pettibone 1993 a, Britayev &
Zamishlyak 1994, Freeman et al. 1998, Britayev et al. 1999), with sponges (Martin et al. 1992,
Pettibone 1993 b, see also remarks on Thormora Baird, 1865 in the present paper), antipathar-
ians and gorgonians (Hartmann-Schröder 1985, 1989, Pettibone 1991 a, 1991 b), mussel beds
(personal observations in the Baltic Sea), deep-sea mussels (Pettibone 1984, 1986 b), gastropods
(Whorff 1991), nudibranchs (Ajeska & Nybakken 1976), and polyplacophores or limpets (Ger-
ber & Stout 1968).
Britayev & Zamishlyak (1994) provide an extensive description of the biology and the com-
mensal life pattern of the Indo-West Pacific Gastrolepidia clavigera Schmarda, 1861. Aggressive
behaviour, both intra- and interspecific, was observed in Arctonoe pulchra (Johnson, 1897) by Di-
mock (1974) and in Hesperonoe Chamberlin, 1919 by Macginitie & Macginitie (1968).
Polynoidae are also known to inhabit the tubes of other polychaetes such as those of Terebelli-
dae (Hutchings 2000). Petersen & Britayev (1997) reported 13 species to be found commensal-
ly in tubes with Chaetopterus species. Malmgreniella andreapolis (McIntosh, 1874) for example lives
in the tubes of Eupolymnia sp. and Amphitrite edwardsi Quatrefages, 1866 (personal observations,
intertidal near Roscoff, Brittany, France). According to Martin & Britayev (1998), some species
seem to be host-specific. In some taxa such as Capitulatinoe Hanley & Burke, 1989 or Asterophilia
Hanley, 1989 commensalism is obligate, whereas it may vary amongst species within other gen-
era. According to Hanley (1989), almost all members of the Arctonoinae seem to be commensal.
Hartman in Fauchald & Jumars (1979) holds the view that any species without or with only
faint ornamentation on the elytra will be found to be commensal.
34 T. Wehe
Polynoids themselves may serve as hosts for other animals. Epizoic entoprocts (= Kamptozoa)
are often found attached to polychaetes. Arroyo & Benito (2000) reported more than 200 speci-
mens of a Loxosomella species attached mainly between the parapodia and below the elytra of a sin-
gle specimen of Lepidonotus clava. Another group reported to use polynoids as hosts is the parasitic
copepods (Gravier 1912, Stock 1996), e.g. for Lepidonotus cristatus Grube, 1876 (Humes & Ho
1967). During the examination of specimens discussed in the present study, individuals of Herma-
dion africanus Hartman, 1974 a were found to be infested.
Eidonomy and critical discussion of characters
Polynoids are short to long scale worms, with a more or less dorsoventrally flattened body. Seg-
ments may vary from less than 20 to over 200. Some species have very stout bodies; others are
very small and fragile. The largest species currently known is probably Eulagisca gigantea Monro,
1939 a with a length of about 20 cm and a width of 10 cm and with a segment number of only
40 (Hutchings 2000). In terms of segment numbers, very long species belong to i.a. the genera
Acholoe Claparède, 1870, Benhamipolynoe Pettibone, 1970 or Lepidasthenia Malmgren, 1867.
However, these worms have a slender body shape and seldom attain more than 10 cm. Species with
only a very few segments, i.e. 15 or less, are found for example amongst the genera Bathylevenstein-
ia Pettibone, 1976 or Bathykurila Pettibone, 1976. One of the smallest species in terms of its total
length seems to be Gesiella jameensis (Hartmann-Schröder, 1974 b) with a maximum of 10 mm,
whereas looking at the ratio of segment number versus total length, Uncopolynoe corallicola Hart-
mann-Schröder, 1960 appears to be the smallest species, with an anterior fragment measuring only
4.8 mm in length and 0.66 mm in width and with a segment number of 44. However, members
of very abundant and widely distributed polynoid genera such as Harmothoe or Lepidonotus exhibit
average body lengths between 2 and 4 cm with roughly about 25 to 40 segments.
Living polynoids can be very colourful (see e.g. Beesley et al. 2000). Elytra, dorsal cirri and head
appendages have more or less distinct colour patterns, which are most probably characteristic for each
individual species and could facilitate the identification of living specimens (see Hutchings 2000).
Unfortunately, specimens in general lose their colouration during preservation and only residues of the
pigmentation are retained, if any. It would therefore be good to mention colour in the description of
living individuals or freshly collected specimens, but it should not be used as a character in diagnoses
or identification keys since most taxonomic studies rely predominantly on preserved specimens.
The integument is smooth or more or less strongly papillate in some genera (e.g. Hermenia
Grube, 1856, Iphione, Parahalosydnopsis Pettibone, 1977). However, “epidermal papillae” in the
Polynoidae are scored as absent in the phylogenetic analysis of polychaete families by Rouse &
Fauchald (1997). As this is not consistent for all species currently included in the Polynoidae,
the statement of a general absence of epidermal papillae is misleading. One might well consider
different types of epidermal papillae in different polychaete families to be homologous or not, as
explained by Fauchald & Rouse (1997, pp. 82‑83). However, it is not legitimate to neglect the
assumed “non-homologous” papillae occurring in several families such as those in the Polynoidae.
The prostomium is generally bilobed with a more or less distinct anterior notch in which the
median antenna is inserted, if present. The anterior borders of both prostomial lobes are rounded
(e.g. Hololepidella), project as anterior peaks (e.g. Harmothoe) or frontal filaments (e.g. Bathykuri-
la), or extend anteriorly to form the ceratophores for the lateral antennae (e.g. Lepidonotus).
The majority of genera possess three distinct antennae, one median and two lateral ones, in-
serted anteriorly on the prostomium. Medioantenna Imajima, 1997 forms a rare exception, where
Fig. 2: Schematic drawings of different arrangement types of lateral antennae on prostomium. From left to right in dorsal, ventral
and right lateral view: a: terminally; b: ventrally; c: terminoventrally; d: subterminally (a-c: modified acc. to Hanley (1987); d:
original (SMF13200), Alentia gelatinosa).
the median antenna arises dorsally on the prostomium. The antennae usually consist of a basal
ceratophore and a distal style, but again a rare exception is found in Uncopolynoe, where the cerato-
phores are absent.
The position of the three antennae relative to each other and their insertion on the prosto-
mium is one of the most important characters to distinguish between subfamilies and also to dis-
tinguish between genera if more than one type of arrangement occurs within a single subfamily.
The terminology applied to different types of arrangement is inconsistent in the literature and has
been subject to dispute, as discussed in detail in the following account. The differentiation of four
36 T. Wehe
general types of antennal arrangement and the use of four different terms, respectively, is applied
here. In the first type, all three antennae are inserted on the same horizontal level and the cerato-
phores of the lateral antennae protrude as anterior prolongations from the prostomial lobes. This
type of insertion of the lateral antennae is called terminal and a prostomium exhibiting such an
antennal arrangement is often referred to as “lepidonotid”, typically found e.g. in the Lepidonoti-
nae or Lepidastheniinae (Fig. 2 a). In the second type, the ceratophores of the lateral antennae are
inserted on a lower level than the median one and are located below the anterior lobes of the pros-
tomium. This insertion is called ventral and the type of prostomium often “harmothoid”, which is
characteristic for e.g. Polynoinae (Fig. 2 b). In the third type, the arrangement of the lateral cerato-
phores is intermediate between the two former ones in that in dorsal view the ceratophores of the
lateral antennae appear to protrude from the anterior borders of the prostomial lobes on the same
horizontal level as the median ceratophore, but are inserted distinctly below the median antenna as
seen in ventral view, where the bases of the ceratophores join midventrally. A further characteristic
of this type of antennal arrangement is that the lateral ceratophores are usually very wide and the
styles often short and distinctly conical. This arrangement of antennae is called terminoventral
(Fig. 2 c). Subfamilies showing this feature are for instance Arctonoinae or Acholoinae. For a long
time, this type of arrangement has not been recognised and has led to recurrent disputes about the
subfamilial and generic placement of species with this character, e.g. Malmgreniella (see Hanley
1987, Pettibone 1993 a, Hartmann-Schröder, 1996, Barnich & Fiege 2001). This can also be
seen from the various terms applied in the past: terminoventrally, ventrally converging midven-
trally, subterminally or laterally (see Barnich & Fiege, 2001). Hanley (1987) was the first who
tried to elucidate this problem. He compared and discussed the arrangement of the antennae of
several genera in detail and provided very helpful figures (Fig. 2 a‑c), clearly demonstrating the in-
termediate arrangement of the lateral antennae, which he termed subterminally. The present work,
however, follows Pettibone (1993 a) and Barnich & Fiege (2001) in preferring the term termi-
noventrally, since it appears to be more appropriate as was argued by these authors. Furthermore,
the use of the terms terminoventrally and subterminally is inconsistent in the literature and neither
term has been clearly defined and differentiated. On the one hand, both have been used to describe
the same feature, as just mentioned. On the other hand, subterminally has been used to describe
two different types of antennal arrangement, e.g. the type of arrangement found in Malmgreniella
(Hanley 1987) and in Heteralentia Hanley & Burke 1991 (1991 a: 58, remarks section on the
genus; see also the extensive remarks on Heteralentia in the present paper). Nevertheless, it is ex-
plained here that in addition to the terminal and ventral insertion of the lateral ceratophores there
is not one but two intermediate states of arrangement which can be clearly separated. In contrast
to the terminoventral arrangement explained above, the ceratophores of the lateral antennae may
protrude anteriorly as distinct prolongations of the prostomial lobes, not on the same level as in
the former type but clearly more ventrally to the median antenna, in part covered by the median
ceratophore in dorsal view (Fig. 2 d). In the latter type the ceratophores of the lateral antennae
are usually also much more slender, with long slender styles, and confined to the inner part of the
prostomial lobes, whereas terminoventrally inserted ceratophores are usually broad and occupy
more or less the whole width of each lobe. This fourth type of arrangement of lateral antennae
should be referred to as subterminally.
Several other subfamilies do not exhibit any of the above mentioned types of arrangements.
They do, however, include only a minority of genera and species, which are clearly separated by
characters other than the antennal arrangement. We find here, for example, the Uncopolynoinae
n. subfam., where only three small papilla-like antennae without ceratophores are present, the
Iphioninae with only two lateral antennae, and subfamilies with lateral antennae absent, e.g. the
Branchipolynoinae and Macellicephalinae, or subfamilies where the antennae are completely ab-
sent such as the Bathyedithinae or Polaruschakovinae.
Another type of prostomial appendage, the palp, is always present as a pair ventrally inserted
on the prostomium. They are usually much stouter than the antennae. Palps are smooth or papil-
late (e.g. in Thormora). Only one genus, Bouchiria Wesenberg-Lund, 1949, was described without
palps. This was, however, based on a misinterpretation and the genus Bouchiria is synonymised
here (see description of Lepidasthenia vesiculosa (Wesenberg-Lund, 1949) below).
Two pairs of eyes are present in most Polynoidae, although they are not described in some
genera (e.g. Iphionella McIntosh, 1885 or Gesiella Pettibone, 1976) or are reported to be absent in
some species belonging to genera usually known to possess eyes. Most of the time, the first pair of
eyes is situated laterally to dorsolaterally at about the widest part of the prostomium, whereas the
posterior pair is mostly situated dorsally close to the posterior corners of the prostomium in front
of the hind margin. There are some exceptions from this general type of arrangement, e.g. in Har-
mothoe, Lepidonotus or Verrucapelma Hanley & Burke, 1991 a, where the anterior pair of eyes may
be situated more anteroventrally to the cephalic peaks (e.g. H. imbricata) or dorsally to them (e.g.
V. nigricans (Horst, 1915)), or both pairs of eyes are situated laterally on the prostomium (e.g. L.
tenuisetosus (Gravier, 1902)).
In contrast to the well-developed prostomium, the peristomium is presumably reduced to the
lips around the mouth (Fauchald & Rouse 1997, Rouse & Pleijel 2001). The muscular pharynx
is eversible and possesses numerous soft papillae arranged in two semicircles on its terminal edges
and usually two pairs of strongly recurved, chitinous jaws.
According to Fauchald & Rouse (1997) and Hutchings (2000), nuchal organs should be
present in the Polynoidae. In sigalionid scale worms nuchal organs are often obvious as small
rounded lobes between the prostomium and the parapodia of the second segment. However, in
none of the polynoid specimens examined during this study could such structures be observed,
nor was it possible to find any descriptions of nuchal organs in the literature. However, close to the
hind margin of the prostomium on the dorsal surface of the second segment, several structures can
be found (see below) in different taxa of Polynoidae which have been named nuchal fold, lobe, flat,
nuchal nodules, nodes or knobs, respectively. These structures, however, are not to be regarded as
nuchal organs, since they do not fulfil the definition of paired, sensory, ciliated structures on the
dorsal side of the prostomium or peristomium as mentioned e.g. by Storch & Schlötzer-Schre-
hardt (1988) or Fauchald & Rouse (1997). Nuchal organs are therefore regarded as absent in
the Polynoidae. Nevertheless, Fauchald (pers. comm.) has observed structures in a few specimens
wedged between the prostomium and the parapodia of the second segment, which might represent
nuchal organs.
A more or less distinct structure, the facial tubercle, may be present ventrally, between the pos-
terior edge of the prostomium and the frontal edge of the eversible pharynx. According to Faucha-
ld (2001), this may be a structure unique to scale worms. The size and structure of this character
varies considerably between specimens of the same species, which may be due to natural variation
or may even depend on the preservation of the prostomium. Furthermore, it is influenced by the
degree to which the pharynx is everted. This character appears to be of limited value for taxonomic
purposes at generic or species level.
The following first or tentacular segment is curved around the prostomium, but is not vis-
ible dorsally. It carries a pair of tentaculophores, the modified first pair of parapodia, inserted
laterally to the prostomium and projecting anteriorly, each one maintained by a single acicula.
Each tentaculophore bears a dorsal and ventral tentacular cirrus and may be either achaetous or
possessing a single or few chaetae, usually similar to the notochaetae of the following parapodia.
38 T. Wehe
According to Fauchald & Rouse (1997), tentacular cirri are defined as altered dorsal and ventral
cirri of cephalised segments which have maintained a more or less distinct parapodial character.
In other families such as the Phyllodocidae or Nereididae, several pairs may be present. In recent
characterisations of the family Polynoidae, different opinions on the development of this first pair
of parapodia are to be found. According to Fauchald & Rouse (1997), it is biramous with the
neuropodia fused to the lower side of the head. Rouse & Pleijel (2001) state “segment 2 and the
following segments are biramous with notoaciculae and simple notochaetae ... and neuroaciculae
... and simple neurochaetae”, obviously implying another character state than biramous for the
tentacular parapodia. Hutchings (2000) again follows Fauchald & Rouse (1997) in that parapo-
dia are biramous (p. 152) but also states (p. 154) that each tentacular parapodia has a single acicula
and notochaetae may be present or absent. For reasons explained in detail in the discussion of the
parapodial characters below, the tentaculophores are here regarded as uniramous. The term tentac-
ulophores seems to be a more appropriate term than first parapodia to describe the morphological
differences from the following parapodia.
The second or buccal segment is the first one to carry biramous parapodia (see below), the first
pair of elytra, and usually much longer and/or stouter ventral cirri, which are termed buccal cirri.
There is no major difference between the general pattern of the second pair of parapodia and that
of the subsequent ones, despite their often being smaller than the following ones. Sometimes the
buccal segment exhibits some additional features on the dorsum, which are not found on other seg-
ments, such as a nuchal lobe or nuchal nodules as mentioned above, more or less distinct structures
protruding in an anterior direction, sometimes covering the posterior part of the prostomium. The
remaining body dorsum does not usually show any further characters, but e.g. in Iphione there are
a few segments with nodular structures.
Unlike sigalionid scale worms, gills are not a character commonly observed in the Polynoidae.
They are developed in some deep-sea forms, such as Branchipolynoe Pettibone, 1984, where they
have an arborescent appearance and are situated dorsally above the notopodia. Simple digitiform
branchiae on the dorsal tubercles are found in Gaudichaudius Pettibone, 1986 a. The opinion
of Fauchald & Rouse (1997) and Rouse & Fauchald (1997) that gills are generally absent
in Polynoidae is not followed here. The opinion of these authors on gills is contradictory. Even
though structures are explicitly mentioned and reported to be presumably of respiratory function
(Fauchald & Rouse 1997: 112), they are not considered as branchiae, since they do not fulfil the
definition given by Fauchald & Rouse (1997: 82) which requires a closed circulatory system.
They are consequently scored as absent in these authors’ cladistic analysis of polychaete families
(Rouse & Fauchald 1997). In a similar way, the much more obvious respiratory appendages of
the Glyceridae Grube, 1850 were also not considered as branchiae by these authors. Böggemann
(2002), however, in his revision of the family Glyceridae, pointed out that branchiae are primarily
characterised by a surface enlargement and a thin respiratory epithelium (see also Kilias 1993: 15).
In contrast to the Sigalionidae and Glyceridae, the family Pectinariidae was considered to posses
branchiae and the character was scored as present in the cladistic analysis by Rouse & Fauchald
(1997), even though Storch & Alberti (1978) clearly demonstrated that branchiae in Pectinaria
koreni (Malmgren, 1866) only possess blood spaces instead of a closed circulatory system. Although
different respiratory structures amongst polychaetes may not be homologous, there is no reason for
not considering them as functional with respect to gas exchange, i.e. to be true branchiae. For this
reason, the digitiform extensions of the elytrophores and the dorsal tubercles, respectively, are re-
garded as branchiae in the present paper, as they are e.g. by Pettibone (1984, 1986 a).
The number and distribution of the elytra along the body is very different among most of the
genera, but some overall similarities can be observed in the pattern of the arrangement. Even if the
scales are lost, the pattern may be reconstructed by the arrangement of the elytrophores to which
the scales are attached. The first 12 pairs of elytra are found in almost all genera on segments 2,
4, 5, 7, and then on alternating segments up to 23 (genera with fewer segments possess a smaller
number of pairs of elytra, but the same distribution pattern up to the last segment). One exception
from this rule is found in Benhamipolynoe, where the elytral arrangement is variable posterior to
segment 15. Usually, the first differences between genera occur posterior to segment 23. Very often
the following three pairs continue on segments 26, 29 and 32, but they are only rarely found to
continue on alternating segments, e.g. Pseudopolynoe Day, 1962. Deviant singular types of arrange-
ment are found in e.g. Arctonoe, Bathynoe Ditlevsen, 1917 or Capitulatinoe. Many genera possess
a definite number of pairs of elytra, for example 12 in Lepidonotus, Thormora and Hermenia, 15
in Harmothoe and Adyte, 18 in Acanthicolepis Norman in McIntosh, 1900 and Parabathynoe Pet-
tibone, 1990, or 22 in Hyperhalosydna Augener, 1922 a. In genera with more than 15 pairs of
elytra, distribution patterns become increasingly different, although in some genera with numerous
pairs the distribution on the posterior segments is often regular again in that the elytra are found
on alternating (e.g. Antipathipolyeunoa Pettibone, 1991 b) or every third segment (e.g. Acholoe).
Distinct numbers of pairs of elytra are often coupled with a distinct number of segments, and this
is especially the case in some lepidonotid scale worms such as, for instance, Lepidonotus and Eu-
phione McIntosh, 1885, where 12 pairs of elytra occur together with 26 body segments, or in some
Iphioninae such as Iphione or Iphionella with 13 pairs of elytra and 29 segments. There are also a
few long-bodied genera with numerous segments but with only a few pairs of elytra, leaving much
of the posterior body uncovered (e.g. Pseudopolynoe). The number and arrangement of elytra are
very important characters and are currently used to distinguish genera.
The elytra themselves are very variable in terms of size, shape and ornamentation. Generally,
they are attached to a more or less obvious elytrophore. In general, the appearance of the elytra is
not a very useful character for distinguishing between subfamilies or genera, although some trends
can be noticed, such as honeycomb polygonal compartments on the elytra of Iphioninae or almost
smooth elytra in Lepidastheniinae, but not necessarily all the species exhibiting these characters
belong to these two subfamilies. However, as regards identification at the species level, the appear-
ance of elytra is one of the most important characters, and so specimens with all the elytra lost are
indeterminable, especially in very speciose taxa such as Harmothoe or Lepidonotus. If not totally
smooth, numerous structures have evolved on the surface of the elytra and may be present in
various combinations. These include micro- and macrotubercles, various papillae, vesicles, thorns,
spines, mounds, ridges, folds, etc. Again, some of these structures are developed in various ways;
tubercles for example may be smooth or spinous, papillae globular, clavate or digitiform, short or
long. Often elytra are encrusted with sand or other inorganic matter, covering the above-men-
tioned structures. Encrusted elytra are very typically found for instance amongst Iphione species.
The margins of the elytra may be smooth or very often possess short to long digitiform papillae.
Sometimes the elytra are very large and cover the body completely (e.g. Lepidonotus), sometimes
they are small and leave large parts of the dorsum uncovered (e.g. Lepidasthenia), or they are almost
inconspicuous (e.g. Hermenia). They may be very stiff and tightly imbricated (e.g. Iphione) or soft,
folded and not tightly imbricated (e.g. Alentia Malmgren, 1865, Paradyte).
Dorsal cirri instead of elytra are found on non-elytrigerous segments. The homology of these
two structures was proposed long ago and was studied in detail by Duncker (1905). There are cir-
rigerous and elytrigerous segments in the Aphroditoidea, and no dorsal cirri are found on elytriger-
ous segments, whereas other polychaete families have dorsal cirri on each segment (if the parapodia
are not reduced as in Capitellidae, for example). The Phyllodocidae and Alciopidae, however, have
leaf-like structures in the position of the dorsal cirri and elytra, suggesting an intermediate state
40 T. Wehe
and a possible path of development between cirri and elytra. Generally, the dorsal cirri in the Poly-
noidae consist of a distinct, more or less cylindrical or globular cirrophore, and a distal style with
or without subterminal swelling or papillae. Usually the dorsal cirri are situated near to the body,
dorsoposteriorly on the parapodia, in contrast to the more dorsomedially-situated elytrophores. In
the same position as the latter, dorsal tubercles may be present on cirrigerous segments, as more or
less distinct nodular to globular structures. They are often inconspicuous, absent, or rarely form
very prominent structures like the T‑shaped dorsal tubercles in Acholoe or the pseudo-elytra in Eu-
phionella Monro, 1936.
In addition to the fact that the parapodia become smaller towards the end of the body, their
general development is the same all along the body. They possess a larger neuropodium and a
smaller or even vestigial notopodium. In the past several terms have been applied, depending on
the notopodial development, to distinguish distinct types of parapodia within the Polynoidae.
Fauchald & Rouse (1997) discussed the use of the terms uni-, bi-, sesqui- or subbiramous (the
two latter having the same meaning), and concluded that applying one or another term is more
or less subjective and depends on the opinion of the author in question. Indeed, this is true for
many descriptions of polynoid species, where more or less the same types of parapodia are referred
to as biramous or subbiramous by different authors. For example, Seidler (1923: 6) referred to
biramous parapodia in the Lepidonotinae: “An den Seiten finden sich die Parapodien, die immer
in zwei Äste geteilt sind, die Dorsal- und Ventral-Aste”. From his diagnosis on Lepidonotus it be-
comes clear that his view was based on the presence or absence of notochaetae, as he stated further:
“Die Parapodien sind mit Ausnahme weniger Arten zweiästig oder besser gesagt, der Dorsalast
trägt Borsten, ...”. Fauvel (1923) also considered the parapodia of Lepidonotus to be biramous
(p. 45: “parapodes biramés”). According to Day (1967 a), the parapodia of the Aphroditidae (=
all scale worms sensu Day) “... [are] almost always biramous”. In the section in which he explains
the diagnostic characters of the Polynoinae (= Polynoidae sensu the present work), parapodia are
described as follows: “The notopodium may be well developed or be reduced to a mere papilla
with an internal acicula so that the foot is uniramous or sesquiramous”. Unfortunately, in the sub-
sequent account of genera and species, also including Lepidonotus, he makes no further reference
to the particular type of parapodia occurring in different genera. Nevertheless, contrary to Seidler
(1923), the terminology applied by Day (1967 a) is based on the development of the notopodium
rather than on the presence or absence of notochaetae. Pettibone (1963, 1982) regarded the
parapodia as biramous in all scale worms, supported by aciculae. More recently, the parapodia of
the Lepidonotinae in general and of Lepidonotus in particular have been described as biramous or
subbiramous by Hanley & Burke (1991 a) and Imajima (1997), whereas Barnich & Fiege (2003)
referred only to subbiramous parapodia for those taxa.
To avoid such ambiguity, Fauchald & Rouse (1997) preferred to use only the two terms uni-
and biramous parapodia for all polychaete families, and stated that only the Saccocirridae possess
“truly uniramous parapodia”. The present author agrees with the latter authors that there is no
objective criterion for drawing a line between biramous, subbiramous or sesquiramous parapodia,
respectively, but on the other hand Fauchald & Rouse (1997) and Rouse & Fauchald (1997) do
not define which characters constitute a uniramous or a biramous parapodium.
Traditionally, the character states biramous and subbiramous are widely in use to distinguish be-
tween subfamilies or genera in the Polynoidae, see for example Pettibone (1969 a, 1976), Hanley
(1989), Hanley & Burke (1991 a), Barnich & Fiege (2000, 2003). The term subbiramous, how-
ever, merely describes the fact that notopodia are smaller than neuropodia, which is the case in all
Polynoidae, thus leaving it as a subjective decision for any worker at which size of notopodium the
parapodium is regarded as bi- or subbiramous, or even uniramous, as one might be tempted to
state when looking at the vestigial notopodia in Lepidasthenia for instance. Furthermore, different
perceptions of this character not only occur among different authors but are also found within the
work of one and the same author. Comparing for example the figs 12 C (biramous parapodium of
Iphione ovata Kinberg, 1856), 19 B (subbiramous parapodium of Hermenia acantholepis (Grube,
1876)) and 21 C (biramous parapodium of Lepidonotus glaucus (Peters, 1854)) in Hanley & Burke
(1991 a) clearly demonstrates the inconsistent use of the terms bi- and subbiramous.
The idea followed in the present paper is to base the terms applied to different parapo-
dial developments on the presence or absence of aciculae. If no aciculae at all are developed (e.g.
Capitellidae, see Fauchald & Rouse 1997), parapodia should be termed aramous if they are not
completely reduced. In cases where there is only a single acicula or a single set of aciculae (a set is
here regarded as composed of several aciculae but forming a single functional unit separated from
a second one if such a one is present), the parapodia should be termed uniramous as is the case
e.g. for some members of the Lumbrineridae and Oenonidae (see Paxton, 2000 a: fig. 1.60 D;
Paxton, 2000 b: fig. 1.61 D). Finally, if there are two separated aciculae or two separated sets of
aciculae, the parapodia should be regarded as biramous. This character state is found i.a. amongst
members of the Eunicidae (see Fauchald 1992), Nereididae (see Wilson 2000: figs 1.80 E, F) and
all scale worms including the Polynoidae. The parapodia in the Polynoidae are thus considered
here to be biramous. The term subbiramous is considered to be uninformative and ambiguous,
as demonstrated above, and should no longer be used. Instead, notopodia might be described like
other characters as prominent, large, indistinct or small, etc. Of course, these descriptors are not
objective either, but they do not imply a strict categorical scheme into which a gradually occurring
character has to fit.
The notopodia are generally situated anterodorsally with respect to the always larger neuro-
podia. Their size varies from more or less prominent and large (e.g. Harmothoe, Neohololepidella
Pettibone, 1969 c) to small (e.g. Australaugeneria Pettibone, 1969 a, Parahalosydnopsis) to almost
vestigial (e.g. Lepidasthenia, Hermenia). Distinct notopodial lobes are usually rounded or conical,
sometimes with long acicular processes.
The neuropodia may be short and truncate, as is the case in Lepidonotus or Hermenia for in-
stance, or they may be longer and more slender as in e.g. Lepidasthenia, or rarely very long and
slender as in Macellicephaloides Uschakov, 1955 a [1969]. Neuropodia possess two parapodial
lobes, a prechaetal and a postchaetal one, with a bundle or fan of neurochaetae in between. Both
lobes may be of the same or different lengths, but in the latter case the postchaetal lobes are usually
shorter. The neuropodial aciculae are always found within the prechaetal lobe, which often forms a
small to prominent process containing the acicula, e.g. in Harmothoe or Macellicephala McIntosh,
1885. Sometimes the aciculae penetrate the epidermis, as is usually found in Harmothoe. The distal
parts of the neuropodia may be deeply incised (or notched) vertically, forming more obvious pre-
and postchaetal lobes, as in the Lepidastheniinae or Acholoinae.
Apart from these characters, the parapodia are rather uniform and do not have many other
features. Papillae may rarely be present on the distal borders, for example in the genera Iphione,
Halosydnopsis Uschakov & Wu, 1959, Parahalosydnopsis or Lepidonopsis Pettibone, 1977.
Ventral cirri are found on all neuropodia, sometimes inserted more basally, sometimes more
distally. Cirrophores are usually small and often indistinct, and styles are smooth (e.g. Lepidonotus)
or papillate (e.g. Harmothoe).
Nephridial pori are often located on the tips of nephridial papillae, which, if present, are
situated at the posteroventral bases of the parapodia next to the joint to the following segment.
Nephridial papillae are present or absent, even amongst specimens of the same species, which may
be due to different reproduction phases (see below) or simply to preservation artefacts. The devel-
42 T. Wehe
opment of nephridial papillae may also be due to sexual dimorphism as was demonstrated by Van
Dover et al. (1999) in the genera Branchipolynoe and Opisthotrochopodus Pettibone, 1985 c. The
use of nephridial papillae as an identification character is therefore problematic. Some genera like
Paralepidonotus and Gastrolepidia possess ventral lamellae, more or less semilunar lobes or protru-
sions found at the ventral bases of the parapodia next to the posterior joint of each segment.
Contrary to the name giving a Greek root for “Polychaeta”, seta(e) instead of chaeta(e) is very
often used to name the bristles (e.g. Pettibone 1969 a, 1969 c). The present work consequently
uses chaeta(e), in order also to emphasize the different modi of origin between polychaete bristles,
which are borne by single epidermal chaetoblasts, and the cuticle-borne bristles or seta(e) of ar-
thropods, for which the latter term is mostly used.
All chaetae in the Polynoidae are simple, and compound neurochaetae as in the Sigalionidae
are completely absent. However, there is a wide variety of different types of chaetae. Notochaetae
are numerous and arranged in dense, tuft-like or radiating bundles (e.g. Iphione, Hermadion), mod-
erate in number (e.g. Lepidonotus), only very few or even completely absent (e.g. Hyperhalosydna,
Lepidasthenia). They may be more slender or stouter than neurochaetae, rounded or flattened in
cross-section, straight or curved, smooth or spinous, their tips may be blunt, pointed, or capil-
lary, sometimes notched distally. Neurochaetae also vary from numerous to very few, but they are
never completely absent. They are usually arranged vertically in fan- or rack-like bundles. They are
stouter or more slender, but usually longer than the notochaetae. Although also exhibiting a wide
range of character states, the majority of neurochaetae are falcate and are similar in shape in that
they are straight, becoming more or less thickened subdistally and tapering to more or less curved,
pointed or blunt, uni- or bidentate tips. Rows of spines or single spines are usually restricted to the
area between the subdistal swelling and the tip. Clearly different from this type of neurochaetae are
the ones found for example in several members of the Macellicephalinae (see Pettibone 1976), or
in Australaugeneria and Uncopolynoe (see the present paper). With very rare exceptions (e.g. Aus-
tralaugeneria, Uncopolynoe) chaetae are similar on each parapodium along the body, and only slight
gradual changes in their appearance may be found on the anterior- and posteriormost segments.
Along with the elytra, chaetae are the most important character for the identification of species.
However, according to Hutchings (2000), some species may be based only on intraspecific
variations, since the investigations of Hillger & Reish (1970) for example have shown that the
formation of the chaetae may be a reflection of the surrounding water temperature. Depending on
the latter, two different species of Halosydna were able to develop uni- or bidentate neurochaetae,
a very important character which is not only used to separate the two species in question but also
numerous other species in other genera.
The pygidium usually bears two anal cirri similar to the dorsal cirri. They are rarely absent
(e.g. Iphione).
Materials and methods
Area investigated
The area investigated (Fig. 1) comprises the following marine regions situated in an anticlockwise
direction around the Arabian Peninsula: (1) The Suez Canal. — Taking Lessepsian migration (Por
1978) into consideration, it seems legitimate to include records from the Suez Canal as its fauna is
of predominantly Red Sea rather than Mediterranean origin. The Suez Canal thus represents the
northwestern-most part of the whole Arabian marine region. (2) The Red Sea with its two northern
branches, the Gulf of Suez and the Gulf of Aqaba. — According to the International Hydro-
graphic Bureau (1986), the southern delimitation of the Red Sea and the Gulf of Aden is by con-
vention drawn by a line running from Hisn Murād in Southern Yemen (12°40'20"N/43°30'20''E)
south-westwards to Cape Si Ane (= Ras Siyan) (12°28'50"N/43°19'10''E), and so the Straits of Bāb
al Mandab together with the island of Perim belong to the Red Sea. (3) The Gulf of Aden, con-
necting the Red Sea and the Arabian Sea. — The westernmost part forms the Gulf of Tadjoura,
whereas to the east the Gulf of Aden may be delimited by a line joining Cape Fartac (= Ras Fartac)
(15°38'N/52°13’30''E) in Southern Yemen and Cape Asir (= Ras Asir) (11°50'N/51°17'E) in Somalia
(International Hydrographic Bureau 1986). (4) The Arabian Sea, representing the northernmost
part of the Indian Ocean. — This sea might be considered as a deep-sea basin bordered by the south-
ern coastline of the Arabian Peninsula, the western coast of the Indian subcontinent, the Carlsberg
ridge to the southwest and the Maldive ridge to the southeast. According to the International Hy-
drographic Bureau (1986), the lines of delimitation of the Arabian Sea run from Cape Hafun (=
Ras Hafun) (10°26'N/51°25'E) in Somalia to the southernmost part of the Maldives Islands, namely
the Addu Atoll (0°42'S/73°09'30''E), and from the latter westwards along the Maldives Islands and
Lakshadweep up to Devgad Island (India) (14°49'10''N/74°03'30''E) in the north. For the purpose
of compiling an annotated checklist listing all valid polychaete species reported from the Arabian re-
gion, Wehe & Fiege (2002) decided to set the limits of the Arabian Sea in terms of the usage of the
respective geographic vocabulary, that is to say that only publications referring to “Arabian Sea” in the
title or works in which collection sites are explicitly cited as “Arabian” were taken into consideration.
However, for the revision of the scale worms presented here, some additional specimens available
from the waters around Karachi, Pakistan, have been included. (5) The Gulf of Oman, connecting
the Arabian Sea with The Gulf. — The Gulf of Oman might be delimited from The Gulf by a line
drawn from Cape Līmah (= Ras Līmah) (25°56'35''N/56°27'30''E) in Oman to Cape Al Kūh (= Ras
Al Kūh) (25°47'30''N/57°18'E) in Iran, and from the Arabian Sea by a line stretching from Cape Al-
Hadd (= Ras Al-Hadd) (22°32'N/59°47'30''E) in Oman to Cape Jiwani (= Ras Jiwani) (25°01'10''N
/61°44'15''E) in Pakistan (International Hydrographic Bureau 1986). (6) The Gulf proper.
Depositories of specimens
This study is based on all the available published and unpublished specimens which were collected
at various sites throughout the region defined above. Of all the specimens examined during this
study, about 800 could be identified to species and are listed here. Of these specimens about 16 %
belong to the collections of the Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt
a.M., Germany (SMF). All other specimens are deposited in the following institutions (abbrevia-
tions as used in the systematic part):
BMNH The Natural History Museum, London, Great Britain (formerly British Mu-
seum of Natural History)
CEWRD Centre for Environmental and Water Research Institute, King Fahd University
of Petroleum & Minerals, Dhahran, Kingdom of Saudi Arabia
Coll. George The Natural History Museum, London, Great Britain (formerly British Mu-
seum of Natural History)
44 T. Wehe
HUJ The Hebrew University, Zoological Collections, Jerusalem, Israel

IEA Institut d’Écologie Appliquée, Angers, France
IOCAS Institute of Oceanology, Academy of Science, Qingdao, China
LACM-AHF Los Angeles County Museum of Natural History, California, U.S.A. (contain-
ing Allan Hancock Foundation collections)
MNHN Muséum National d’Histoire Naturelle, Paris, France
MPW Museum of Natural History, University of Wrocław, Poland
NHCY Natural History Collection Yemen (temporarily deposited at SMF)
NHMW Naturhistorisches Museum, Vienna, Austria
NMNHT National Museum of Natural History at Tel-Aviv University, Israel
SMNH Swedish Museum of Natural History, Stockholm, Sweden
USNM United States National Museum of Natural History, Smithsonian Institution,
Washington D.C., U.S.A.
ZMB Naturhistorisches Forschungsinstitut, Museum für Naturkunde, Zentralinsti-
tut der Humboldt-Universität zu Berlin, Germany
ZMH Zoologisches Institut und Museum der Universität Hamburg, Germany
ZMUC Zoologisk Museum, University of Copenhagen, Denmark
ZMUM Zoological Museum, University of Moscow, Russia
Outline of species description
In the synonymy given under each species, the references listed deal primarily with Arabian poly-
chaetes. Any additional references listed are restricted to the original descriptions and important
revisions or major faunistic works which have been used for comparison and identification.
In the “specimens examined” sections, records are listed according to their occurrence in the
respective seas in an anticlockwise direction around the Arabian Peninsula.
In the “measurements” section, the following abbreviations are used: cs = complete specimen,
af = anterior fragment, mf = median fragment, pf = posterior fragment. Notation of measurements
are given according to the following standard: cs (26) 40/15/10 [af (8) 15 + pf (18) 25] or af (55)
60/8/5. Numbers in parenthesis refer to the segment number, those separated by diagonal slashes
indicate from left to right the length, the width including and excluding chaetae. For complete
but fragmented specimens, segment number and length of each fragment are given additionally
in square brackets. Length was measured from the anterior margin of the prostomium to the hind
margin of the pygidium. Width was measured on the widest segments, usually in the anterior
body part, from the tips of the chaetae or the lateral margins of neuropodia, respectively. For many
specimens, measurements are only approximations due to curling.
Data on “depth” and “occurrence” within the Arabian region are based on the specimens ex-
amined. Additional information based on the literature may be provided in the “remarks”, “biol-
ogy and ecology”, “depth” and “distribution” sections for each species.
Place names
Sheppard et al. (1992) discussed the problem of naming “The Gulf ”. The name “The Gulf ” is
used throughout this work as given in the latest edition of “The Times Comprehensive Atlas of
the World” (The Times 2003). Names for collecting sites mentioned in the present work are cited
as they appear on the labels in the collection vials or in the relevant references, sometimes supple-
mented by an additional different spelling based on the The Times atlas.
Matrix
Values computed are based on the total number of species as mentioned in the chapter on zoog-
eography and the data matrix given below. Data for the Polynoidae occurring in the Mediterranean
Sea were compiled according to Table 1 in Barnich & Fiege (2003). The Indo-West Pacific data
on Polynoidae were compiled according to Barnich et al. (2004), Hanley (1992, 1993), Hanley
& Burke (1990, 1991 a), Imajima (1997), McIntosh (1885), Pettibone (1969 a, 1977, 1986 a),
and Rullier (1972).
Polynoidae species in common to two areas:
Suez Canal ↔ Red Sea: 5

- ↔ Gulf of Aden: 2
- ↔ Arabian Sea: 3
- ↔ Gulf of Oman: 0
- ↔ The Gulf: 2
- ↔ Mediterranean Sea: 1
- ↔ Indo-West Pacific: 4
Red Sea ↔ Gulf of Aden: 6
- ↔ Arabian Sea: 12
- ↔ The Gulf: 8
Gulf of Aden ↔ Arabian Sea: 5
- ↔ The Gulf: 2
Arabian Sea ↔ Gulf of Oman: 3
- ↔ The Gulf: 5
Gulf of Oman ↔ The Gulf: 2
The Gulf ↔ Mediterranean Sea: 2
Computation of faunal affinity values
Values for the comparison of the faunas of two different areas were computed using
(a) The Second Kulczynski Coefficient,
(b) The Ochiai/Otsuka Coefficient.
46 T. Wehe
The notation used here follows that of Cheetham & Hazel (1969). These coefficients measure the
similarity between two bioassociationed areas and are given by:
C ( N1 + N 2 ) C
(a) (b)
2( N 1 N 2 ) ( N1 N 2 )
with:
C being the number of species in common to both areas compared,
N1 being the total number of species present in the first area,
N2 being the total number of species present in the second area.
Light microscopy, preparation of drawings and photography
Preserved specimens were examined using stereomicroscopes (WILD Heerbrugg M5, Leica WILD
MZ8) and compound microscopes with interference contrast (Leica Diaplan, Olympus BH-2).
Drawings were made using a camera lucida. If not already detached, elytra from the anterior body
regions were removed for drawings of the prostomium. The elytra figured are arranged with the
anterior border directed to the top. In some cases, all elytra were detached, making correct orienta-
tion difficult.
Photographic documentation used a SLR camera (Olympus OM 4 Ti) with macrophoto
lenses of 20, 38 and 90 mm and a digital camera Canon EOS D60.
Abbreviations
Listed below are abbreviations used in the text which are not all in general use in taxonomy and
are not explained elsewhere in this work. Further abbreviations found in the “specimens examined”
sections and not listed below are not introduced in this work but refer to information given on the
original labels or in the references in question.
coll. collected
Coll. Collection
det. determined
JMWS Jubail Marine Wildlife Sanctuary
n.d. no data
spm(s). specimen(s)
Taxonomic account
Polynoidae Kinberg, 1856

D i a g n o s i s : Body short to long, dorsoventrally flattened, subrectangular or rounded in cross-
section. Segments few to numerous, with at least 12 to up to 100 and more segments, mostly
ranging between 20 and 50. Prostomium bilobed, usually with three antennae (sometimes only the
two laterals, only the median or none present), with a pair of ventral palps and two pairs of eyes
(sometimes absent). Median antenna usually in anterior notch of prostomium, lateral antennae with
different arrangement patterns in relation to median antenna and prostomial lobes. Facial tubercle
more or less distinct. Prostomium sometimes with additional characters such as cephalic peaks,
frontal filaments or occipital papilla. First or tentacular segment not visible dorsally, with a pair of
tentaculophores lateral to prostomium and palps. Each tentaculophore with a single acicula, with
or without chaetae and a pair of tentacular cirri. Tentacular cirri of similar length or dorsal ones
longer than ventral ones. Everted pharynx with two semicircles of soft papillae on its distal border
and two pairs of chitinous jaws. Second (buccal) segment with first pair of elytra, usually with buc-
cal cirri longer than following ventral cirri, and with or without dorsal nuchal fold or nodules. Pairs
of elytra few to numerous, six to about 90 pairs; usually ranging between 12 and 20; very often
12 or 15. The arrangement of pairs of elytra up to segment 23 usually as follows: 2, 4, 5, 7, 9, 11,
13, 15, 17, 19, 21, 23, then variable. Elytra large to minute, covering body completely or leaving
middorsum or posterior end uncovered. Surface of elytra variously ornamented with micro- and
macrotubercles, papillae, vesicles, ridges, etc., or completely smooth. Margins with or without fringe
of papillae. Dorsal cirri present on all non-elytrigerous segments, situated posterodorsally on para-
podia, with distinct cirrophores and long styles. With or without small to large dorsal tubercles on
non-elytrigerous segments. Parapodia biramous, each ramus with a single acicula. Notopodia coni-
cal-like, anterodorsal of larger neuropodia. Notopodia may be large and distinct, sometimes small or
even vestigial. Neuropodia with distinct prechaetal and postchaetal lobes, sometimes deeply incised
dorsally and ventrally; prechaetal lobe rounded or pointed, postchaetal lobe usually rounded. Both
rami with or without different acicular processes. Ventral cirri usually small, gradually tapering.
Nephridial papillae on posteroventral bases of parapodia present or absent. Additional characters
such as ventral lamellae on ventral bases or branchiae on dorsal surface of parapodia may be present.
Notochaetae and neurochaetae always simple, of various kinds. Notochaetae arranged more or less
tuft-like, few to numerous, sometimes absent; stouter or more slender than neurochaetae, straight or
curved, rounded or flattened, smooth or spinous, tips notched, blunt, pointed or capillary. Neuro-
chaetae usually longer and straighter than notochaetae, more or less arranged in vertical fans, usually
numerous, subdistally more or less thickened, with rows of spines only distally, tips notched, blunt
or pointed, uni- or bidentate. Pygidium usually with a pair of anal cirri.
Key to the subfamilies of Polynoidae (subfamilies marked with * are treated in this paper)
1 Prostomium without antennae................................................................................... 2

– Prostomium with antennae........................................................................................ 3
2 With rounded palpophores. Pharynx with 9 pairs of papillae, jaws with denticled
bases Bathyedithinae
– Without palpophores. Pharynx with 7 pairs of papillae, jaws without denticled bases
Polaruschakovinae
3 With one (median) antenna....................................................................................... 4
– With two or three antennae....................................................................................... 11
4 With distinct branchiae dorsally on parapodia........................................................... 5
– Without distinct branchiae........................................................................................ 7
5 Prostomium more or less rounded. Median antenna situated dorsally. Branchiae sack-
like Branchiplicatinae
– Prostomium bilobed. Median antenna in anterior notch. Branchiae arborescent........ 6
48 T. Wehe
6 Branchiae with short, more densely arranged terminal filaments. With prominent
frontal filaments. Elytra large, covering dorsum Branchinotogluminae
– Branchiae with long, more loosely arranged terminal filaments. With minute frontal
filaments. Elytra small, not covering dorsum Branchipolynoinae
7 First segment with strongly curved acicular lobes Vampiropolynoinae
– First segment without strongly curved acicular lobes.................................................. 8
8 Elytra ornamented. Elytrophores, dorsal tubercles and bases of both with numerous
ciliated ridges. Distal borders of parapodial rami fimbriated Lepidonotopodinae
– Elytra smooth (if known). No ciliated ridges on elytrophores, dorsal tubercles and
their bases. Distal borders of parapodial rami not fimbriated...................................... 9
9 Neuropodia elongated, deeply incised dorsally and ventrally. Without notochaetae
Macelloidinae
– Neuropodia not deeply incised. With notochaetae..................................................... 10
10 Noto- and neuropodia with long projecting acicular process. 7‑12 pairs of elytra
Macellicephalinae
– Only notopodia with long projecting acicular process. 8 pairs of elytra
Macellicephaloidinae
11 With two (lateral) antennae....................................................................................... 12
– With three (median & lateral) antennae..................................................................... 13
12 With long lateral antennae attached anteriorly or short lateral extensions only. Elytra
reticulated, with hexa- or polygonal honeycomb-like areas Iphioninae*
– With lateral antennae attached posteriorly. Elytra without honeycomb-like areas
Genus Cervilia
13 With antennal scales or with antennal and tentacular sheaths Admetellinae
– Neither antennal scales nor antennal and tentacular sheaths present........................... 14
14 With accessory filamentous organs on dorsal cirrophores Gesiellinae
– Without accessory filamentous organs on dorsal cirrophores...................................... 15
15 Lateral antennae terminally or subterminally.............................................................. 16
– Lateral antennae ventrally or terminoventrally converging midventrally..................... 19
16 Lateral antennae terminally, very small, papilla-like, without ceratophores. Clavate
neurochaetae present Uncopolynoinae*
– Lateral antennae distinct, not papilla-like. Without clavate neurochaetae................... 17
17 Lateral antennae subterminally. Noto- and neuropodia with prominent acicular proc-
ess. Notochaetae numerous Eulagiscinae
– Lateral antennae terminally. Noto- and neuropodia without prominent acicular proc-
ess. Notochaetae moderate, few or absent................................................................... 18
18 Neuropodia not deeply incised dorsally and ventrally. Elytra usually large, covering
middorsum, ornamented. Dorsal tubercles more or less distinct Lepidonotinae*
– Neuropodia deeply incised dorsally and ventrally. Elytra usually small, not covering
middorsum, smooth (rarely with few micropapillae). Dorsal tubercles absent
Lepidastheniinae*
19 Lateral antennae ventrally.......................................................................................... 20
– Lateral antennae terminoventrally.............................................................................. 21
20 18 segments, 9 pairs of elytra. Palps conical, short, not visible dorsally

Bathymacellinae
– More than 18 segments, more than 9 pairs of elytra. Palps long, visible dorsally
Polynoinae*
21 Cephalic peaks absent. Distal border of acicular neuropodial lobes bifid. With or
without chaetae on tentaculophores. Dorsal tubercles bifurcate, bulbous or indis-
tinct Acholoinae*
– With or without cephalic peaks. Distal border of acicular neuropodial lobes rounded
or pointed. Without chaetae on tentaculophores. Dorsal tubercles usually indistinct
Arctonoinae*
Acholoinae Pettibone, 1996

D i a g n o s i s : Body elongate, with numerous segments (more than 100 in Acholoe) or moderate
in number (up to 50 in Intoshella, Arctonoella and Pararctonoella). Prostomium without cephalic
peaks, three antennae present. Large ceratophore of median antenna in anterior notch of prostomi-
um, lateral antennae with distinct ceratophores inserted terminoventrally converging midventrally,
with short styles. Palps stout, conical. Usually two pairs of eyes, variable in arrangement. Tentacu-
lophores with or without chaetae. Second (buccal) segment without nuchal fold. Elytra numerous
pairs (more than 40 in Acholoe) on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, continuing
on every third segment to end of body, or moderate in number with 16 to 23 pairs (Intoshella, Arc-
tonoella and Pararctonoella) on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, then variable in
arrangement. Elytra surface usually smooth, sometimes with few microtubercles or nodular papil-
lae; margin with or without fringing papillae. Dorsal cirri with smooth or papillate styles. Dorsal
tubercles on cirrigerous segments bifurcate (Acholoe), bulbous or inconspicuous. Notopodia small
with acicular process. Neuropodia deeply incised dorsally and ventrally; prechaetal lobe distally
bifid or not, with variably developed supra- and subacicular processes; postchaetal lobe shorter
and rounded. Notochaetae few to numerous, shape variable. Neurochaetae moderate to numerous,
with unidentate tips, shape variable. Pygidium with a pair of anal cirri.
R e m a r k s : Species of Acholoinae are usually associated with asteroids, ophiuroids or glass
sponges.
Key to the genera and species of Acholoinae

1 Segments more than 100. Elytra in numerous pairs. Dorsal tubercles bifurcate. Elytra
surface with microtubercles, margin smooth Acholoe astericola
– Segments less than 50. Elytra 15‑20 pairs. Dorsal tubercles indistinct. Elytra surface
without microtubercles, margin with papillae. Notochaetae spinous
Pararctonoella marginopapillata
Acholoe Claparède, 1870

Acholoe Claparède, 1870. — Mémoires de la Sociéte de Physique et d’ Histoire Naturelle de Genève 20(2): 382.
Acholoe. — Pettibone 1996 b: 634‑635.
Ty p e s p e c i e s : Polynoe astericola Delle Chiaje, 1841 a.
D i a g n o s i s : Body vermiform, very long, tapering posteriorly, segments more than 100. An-
tennae short, smooth. Two pairs of eyes, anterior pair anterolaterally, slightly in front of horizontal
50 T. Wehe
Fig. 3: Acholoe astericola (BMNH 1924.6.18.224). a: anterior end; b: elytron from midbody; c: microtubercles on elytra; d: right
parapodium from segment 35; anterior view; e: same, posterior view; f: notochaeta; g: neurochaetae.
midline, posterior pair in front of hind border. Tentaculophores without chaetae. Buccal segment
with rectangular, bulbous protrusion dorsally. Elytra numerous, more than 40 pairs on segments
2, 4, 5, 7, alternating to 23, 26, 29, continuing on every third segment to end of body. Elytra
large, smooth, with scattered microtubercles, margin without fringing papillae. Dorsal cirri with
long styles, extending beyond neurochaetae. Dorsal tubercles prominent, bifurcated, t‑shaped.
Notopodia very small. Neuropodia with slightly longer, bifid, prechaetal lobe and shorter, rounded
postchaetal lobe. Notochaeta few, short, slender, curved, minutely spinous, with blunt tips. Neuro-
chaeta stouter, falcate, with few spinous rows below blunt tip.
Acholoe astericola (Delle Chiaje, 1841) Fig. 3 a‑g

Nereis squamosa Delle Chiaje, 1825 [1827]. — Memorie sulla storia e notomia degli animali senza vertebre del Regno di Napoli 2:
368, 400, 425 [referred to Polynoe astericola by Delle Chiaje, 1841 a: 106].
Polynoe astericola Delle Chiaje, 1841 a. — Descrizione e notomia degli animali invertebrati della Sicilia citeriore osservati vivi
negli anni 1822‑1830 5: 57, 62, 106.
Polynoe astericola. — Delle Chiaje 1841 b; Descrizione e notomia degli animali invertebrati della Sicilia citeriore osservati vivi
negli anni 1822‑1830 7: pl. 94: fig. 15, pl. 129: fig. 7.
Acholoe astericola. — Pettibone 1996 b, Proceedings of the Biological Society of Washington 109(4): 635‑637, fig. 1; Barnich &
Fiege 2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 559: 78‑80, fig. 41.
Ty p e l o c a l i t y : Naples, Italy, Western Mediterranean Sea.
S p e c i m e n s e x a m i n e d : R e d S e a : 1 spm., 9 September 1904, Coll. Potts, BMNH 1924.6.18.224.
M e a s u r e m e n t s : BMNH 1924.6.18.224: cs (172) 79/6/5 [af (20) 9 + mf (26) 17 + pf (126) 53].
D e s c r i p t i o n : Prostomium rounded to hexagonal (Fig. 3 a). Ceratophore of median an-
tenna large, cylindrical, style broken off. Ceratophores of lateral antennae slightly smaller, styles of
lateral antenna smooth, short, gradually tapering. Right palp stout, smooth, tapering to a pointed
tip, left palp broken off. Pharynx not extended. Facial tubercle distinct. Tentacular cirri all but
one missing, smooth, gradually tapering, slightly longer than lateral antenna. Buccal segment with
rectangular bulbous protrusion dorsally (Fig. 3 a). Buccal cirri only slightly longer than follow-
ing ventral cirri. Elytra 61 pairs, mostly detached. Elytra leaving parts of middorsum uncovered,
very small on anterior body (Fig. 3 a), larger on median and posterior part (Fig. 3 b). Elytra thin,
translucent, rounded to oval; surface mostly smooth with only a few conical microtubercles near
anterior margin (Fig. 3 c). Some elytra with faint brownish pigment. Dorsal cirri with cylindrical
to bottle-shaped cirrophores and smooth, gradually tapering styles. Dorsal tubercles on segment
three nodular, on following segments ciliated, bifurcate or t‑shaped (Fig. 3 a, 3 e), with both lap-
pets becoming distinctly elongated on posterior segments. Parapodia (Fig. 3 d, 3 e) with very
small, conical notopodia. Neuropodial lobes of similar length, prechaetal one only slightly bifid.
Ventral cirri smooth, short, gradually tapering. Notochaetae few, more slender than neurochaetae,
slightly curved, with faint rows of spines and blunt tips (Fig. 3 f ). Neurochaetae few, stouter and
longer than notochaetae, falcate, with several rows of spines subdistally and blunt, unidentate tips
(Fig. 3 g). Pygidium indistinct, anal cirri broken off.
R e m a r k s : The single specimen examined here clearly represents Acholoe astericola. The only
difference between this specimen and those reported in Pettibone (1996 b) and Barnich & Fiege
(2003) is the indistinct bulbous areas dorsally on segments three and four. Pettibone (1996 b)
also reported no microtubercles on the elytra, but these are described by Barnich & Fiege (2003).
According to the latter authors, the species is widely distributed throughout the Mediterranean
Sea and is also known from the North East Atlantic from the English Channel to West Africa.
Unfortunately, the exact location in the Red Sea where the specimen examined here was found is
unknown. The species was also reported from the Red Sea by Amoureux et al. (1978) (see Wehe
& Fiege 2002). However, re-examination of this specimen revealed that it was misidentified and
52 T. Wehe
belongs to Australaugeneria rutilans instead (see the description of that species). The geographic
range of Acholoe astericola raises some doubt as to the origin of the single specimen examined here
and the occurrence of the species in the Red Sea. The specimen was possibly mislabelled. However,
more sampling is necessary to evaluate the occurrence of A. astericola in the Red Sea.
B i o l o g y & e c o l o g y : The species is known as a commensal of asteroids and ophiuroids
(Pettibone 1996 b, Freeman et al. 1998) but also occurs amongst algae and in sea grass beds (Bar-
nich & Fiege 2003).
D e p t h r a n g e : Unknown, but elsewhere reported from the intertidal down to 100 m (Bar-
nich & Fiege 2003).
D i s t r i b u t i o n : Red Sea. First record of the species for the Red Sea. Elsewhere: Western,
Central and Eastern Mediterranean Sea, North East Atlantic from the English Channel to West
Africa (Pettibone 1996 b).
Pararctonoella Pettibone, 1996

Pararctonoella Pettibone, 1996 b. — Proceedings of the Biological Society of Washington 109(4): 639.
Ty p e s p e c i e s : Intoshella indica Day, 1973.
D i a g n o s i s (emended): Body elongate, with up to 48 segments, tapering posteriorly. Antennae
short, smooth or papillate. With or without two pairs of eyes. Tentaculophores with a single chaeta
each. Elytra 15 to 20 or more pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, then variable
in arrangement. Elytra large, surface smooth, margin with or without small fringing papillae. Dorsal
cirri with long styles, extending beyond neurochaetae, with or without papillae. Dorsal tubercles in-
distinct or bulbous. Notopodia small conical. Neuropodia with acicular prechaetal lobe distally bifid,
with a longer, conical, supraacicular and shorter, rounded, subacicular process; postchaetal lobe short-
er, rounded. Notochaetae numerous; slender, smooth capillaries in P. indica (Day, 1973), slender,
spinous capillaries in P. marginopapillata n. sp., or stout, acicular in P. aphthalma (Gallardo, 1968).
Neurochaeta of two kinds: supraacicular ones with a long spinous region and fine tips, subacicular
ones slightly stouter, shorter with only few spines below blunt or pointed tips.
R e m a r k s : P. marginopapillata n. sp. described here casts some doubt on the validity of the
two genera Arctonoella Buzhinskaja, 1967 and Pararctonoella Pettibone, 1996 b, which were sepa-
rated by Pettibone (1996 b) based on the different number of pairs of elytra (16 in Arctonoella and
18‑20 in Pararctonoella) and by the presence of two types of notochaetae in Arctonoella and only
one type in Pararctonoella. However, two different types of notochaetae (slender, smooth capil-
laries and stout, acicular ones, respectively) have also been described in the latter genus but occur
separately in two species. The characters exhibited by a specimen examined here do not fit with the
above mentioned characters, in that there are only 15 pairs of elytra present and the notochaetae
are of one type, but slender, spinous capillaries. Furthermore the elytra posses small fringing papil-
lae, which have not been described before. The characters used by Pettibone (1996 b) to separate
the genera are elsewhere used to distinguish between species, and so it is very difficult to classify P.
marginopapillata n. sp. A re-examination of the respective type specimens is necessary to clarify this
problem. For the time being it was decided to emend the diagnosis of Pararctonoella to include the
new species, ranking the presence of a single type of notochaetae higher than the number of pairs
of elytra, a variable character in many other polynoid genera.
Pararctonoella marginopapillata n. sp. Fig. 4 a‑j

S p e c i m e n s e x a m i n e d : H o l o t y p e : Gulf of Oman, Oman, Mina al Fahl, north of Muscat, ZMUC POL-1359.
M e a s u r e m e n t s : cs (32) 10/4/3.
Fig. 4: Pararctonoella marginopapillata n. sp. (holotype, ZMUC POL-1359). a: anterior end; b: 1st left elytron; c: 2nd right elytron;
d: 6th left elytron; e: fringing papillae; f: left parapodium from segment 12, anterior view; g: same, posterior view; h: notochaeta;
i: upper neurochaeta; j: lower neurochaeta.
54 T. Wehe
D i a g n o s i s : Styles of antennae and cirri papillate. Surface of elytra smooth, margin with
small fringing papillae. Notochaetae slender, spinous. Neurochaetae of two kinds: upper ones more
slender subdistally, with several rows of spines below pointed tips; lower ones stouter subdistally
with spines few in number or absent, and blunt or pointed tips.
D e s c r i p t i o n : Body elongate, tapering posteriorly, subrectangular in cross-section. Prosto-
mium rounded to slightly hexagonal (Fig. 4 a). Median antenna with a large, cylindrical cerato-
phore, style moderate in length, gradually tapering. Ceratophores of lateral antennae as stout as
median ceratophore, with slightly shorter styles. All three styles with minute papillae. Palps stout,
conical, smooth, gradually tapering. Two pairs of rounded eyes, anterior pair dorsolaterally on
more or less widest part of prostomium, posterior pair in front of hind margin. Tentaculophores
with distinct acicular process and one chaeta each. Tentacular cirri longer than antennal styles, pap-
illate, gradually tapering. Facial tubercle distinct. Buccal cirri much longer than following ventral
cirri, shaped as tentacular cirri. 15 pairs of elytra. Elytra soft, translucent, large, covering dorsum
completely; first pair rounded (Fig. 4 b), second pair reniform (Fig. 4 c), following ones more-or-
less oval, slightly notched anteriorly (Fig. 4 d), partially folded frill-like. Surface smooth, partially
with faint, brownish pigmentation. Margin laterally with few minute fringing papillae (Fig. 4 e).
Dorsal cirri on non-elytrigerous segments with a large, cylindrical cirrophore and a very long, grad-
ually tapering, papillate style distinctly extending beyond neurochaetae. Dorsal tubercles absent.
Parapodia (Fig. 4 f, 4 g) with small, subconical notopodia with a short acicular process. Neuropo-
dia elongate, with a bifid, prechaetal, acicular lobe with a long, conical, supraacicular and shorter
rounded, subacicular process. Aciculae penetrating epidermis in notch between processes. Ventral
cirri short, inserted ventromedially, conical, gradually tapering. Notochaetae (Fig. 4 h) numerous,
more slender than neurochaetae, of one type, slightly curved with numerous rows of spines, taper-
ing to capillary tips. Neurochaetae slightly thicker and longer than notochaetae, of two kinds: up-
per ones slightly thickened subdistally, with numerous rows of spines below pointed, unidentate
tips (Fig. 4 i); lower ones shorter, more thickened subdistally, spines only few in number or absent,
with blunt or pointed tips (Fig. 4 j). Noto- and neurochaetae of second segment slightly different:
notochaetae stouter with coarser rows of spines, neurochaetae more slender, with more numerous
rows of spines and gradually tapering fine tips.
R e m a r k s : Pararctonoella marginopapillata n. sp. is separated from other species within the
genus by the presence of spinous, capillary notochaetae and small fringing papillae on the elytra, as
well as the low number of segments and pairs of elytra. See also the remarks on the genus.
B i o l o g y & e c o l o g y : Unknown.
D e p t h r a n g e : Unknown.
D i s t r i b u t i o n : Gulf of Oman, near Muscat.
E t y m o l o g y : The name refers to the papillae present on the lateral margins of the elytra;
Latin margino = to frame; Latin papilla = nipple.
Arctonoinae Hanley, 1989

D i a g n o s i s : Body usually elongate, with few (about 40) to numerous (up to about 180)
segments, flattened dorsoventrally. Prostomium with or without cephalic peaks, with three an-
tennae. Large ceratophore of median antenna in anterior notch of prostomium, lateral antennae
with distinct ceratophores inserted terminoventrally or ventrally, converging midventrally. Palps
stout, conical. Two pairs of eyes usually on posterior half of prostomium. Tentaculophores without
chaetae. Second (buccal) segment with or without nuchal fold. Elytra usually numerous pairs on
segments 2, 4, 5, 7, alternating to 23, then variable in arrangement. Elytra surface mostly smooth,
margin without fringing papillae. Dorsal cirri usually with smooth styles. Dorsal tubercles incon-
spicuous. Notopodia small, sometimes vestigial, with or without acicular process. Neuropodia
deeply incised dorsally and ventrally; prechaetal, acicular lobe rounded to pointed; postchaetal lobe
shorter, rounded. Notochaetae few or absent, shape variable. Neurochaetae few to moderate, with
uni- or bidentate tips, shape variable. Pygidium with a pair of anal cirri.
R e m a r k s : This subfamily was erected by Hanley (1989) to include Gastrolepidia Schmarda,
1861, Adyte, Hololepidella, Bathynoe, Arctonoe, Australaugeneria, Paradyte, Pottsiscalisetosus Pet-
tibone, 1969 a, Subadyte, Neohololepidella, Parahololepidella Pettibone, 1969 c, Minusculisquama
Pettibone, 1983 b, Disconatis Hanley & Burke, 1988, and Asterophilia. Malmgreniella was subse-
quently added by Hartmann-Schröder, 1996. Separation of these genera from the Polynoinae
was mainly based on the terminoventral or ventral insertion of the ceratophores of the lateral
antennae, the absence of chaetae on the tentaculophores, and the presence of subbiramous, deeply
incised neuropodia. As already emphasised by Barnich & Fiege (2001, 2003), a clear distinction
between the Arctonoinae and the Polynoinae is not possible, the assignment of genera is not un-
equivocal, and a revision of the genera in question is necessary to solve this problem.
Comparing character states, the insertion of the lateral ceratophores is not helpful as a distin-
guishing character, since a ventral or terminoventral arrangement occurs in both subfamilies. As
regards deeply incised neuropodia, some genera do not show this character as distinct (e.g. Sub-
adyte), as for example Acholoe or Lepidasthenia. Furthermore, the character state of subbiramous
neuropodia is not useful as has been demonstrated in the general discussion of characters above.
This leaves the main difference between the two subfamilies as the presence or absence of chaetae
on the tentaculophores. However, this character is not unequivocal, since, after the examination
of additional specimens, Hololepidella was referred back to the Polynoinae (as Harmothoinae) by
Hanley & Burke (1991 a) due to presence of chaetae on the tentaculophores, which is confirmed
in this work.
For these reasons, Barnich & Fiege (2001, 2003) are followed here and the specimens belong-
ing to the genera Hololepidella, Malmgreniella, Paradyte, and Subadyte examined during this study
are referred to the Polynoinae, whereas Australaugeneria and Gastrolepidia are retained in the Arc-
tonoinae for the time being.
Key to the genera and species of Arctonoinae
1 Styles of median antenna, tentacular-, dorsal- and anal cirri strongly bulbous. Para-
podia with ventral lamellae. Enlarged neuropodial lobes and hooked neurochaetae
absent Gastrolepidia clavigera
– Styles of appendages tapered. Ventral lamellae absent. Prechaetal neuropodial lobes
on anteriormost segments strongly enlarged, enclosing hooked neurochaetae
Australaugeneria rutilans
Australaugeneria Pettibone, 1969

Australaugeneria Pettibone, 1969 a. — Proceedings of the Biological Society of Washington 82: 20.
Australaugeneria. — Pettibone 1969 d, Proceedings of the Biological Society of Washington 82: 519‑521.
Ty p e s p e c i e s : Polynoe rutilans Grube, 1878.
D i a g n o s i s (emended): Body dorsoventrally flattened, tapering posteriorly, segments about
40. Prostomium margin rounded without cephalic peaks. Ceratophores of lateral antennae slender,
inserted terminoventrally, converging midventrally. Styles of antennae slender. Palps stout, gradually
56 T. Wehe
tapering. With or without two pairs of eyes. Buccal segment without nuchal fold, without or with
few notochaetae, neurochaetae strongly hooked. Elytra 15 pairs on segments 2, 4, 5, 7, alternating
to 23, 26, 29, 32. Elytra soft, translucent, smooth. Dorsal cirri with long, cylindrical cirrophores
and long styles. Notopodia small, with acicular process. Neuropodia with a prechaetal and postcha-
etal lobe of similar length, both bluntly rounded. Prechaetal lobes of segments 2 and 3 more or less
strongly enlarged, hood-like, enclosing hooked chaetae. Notochaetae few, stouter or more slender
than neurochaetae, smooth or dentate, tips unidentate. Neurochaetae few, present as stout hooks in
anteriormost segments, those of following segments falcate, smooth or faintly spinous below more
or less strongly hooked tips, sometimes bidentate, with a small secondary tooth.
R e m a r k s : Of the three described species, A. rutilans and A. pottsi Pettibone, 1969 d have
bidentate notochaetae according to Pettibone (1969 a, 1969 d), whereas A. michaelseni Pettibone,
1969 a has unidentate notochaetae. This opinion is not followed here. The notochaetae of the two
first-mentioned species are only ostensibly bidentate, because of the distalmost of a series of teeth
along the convex notochaetal edge reaching almost to the tip of the chaetae. These teeth are miss-
ing in A. michaelseni, leaving them clearly unidentate. Figure 10 g in Pettibone (1969 a) could
give the impression of bidentate tips, but in the specimens examined here the notochaetal teeth
reach the tip of the chaetae without a gap (see Fig. 5 e), as is the case for A. pottsi (see Pettibone
1969 d: Fig. 1 f ). The bidentate tip of a chaeta should be clearly separated from subdistally located
teeth, spines or serrations (e.g. Harmothoe, Lepidonotus). The notochaetae in Australaugeneria are
here considered to be unidentate and the generic diagnosis is emended for this character.
According to Pettibone (1969 a), the lateral antennae are inserted ventrally. Again, this opin-
ion is not followed here, and the lateral antennae are considered to be inserted terminoventrally.
Australaugeneria rutilans (Grube, 1878) Fig. 5 a‑i

Polynoe rutilans Grube, 1878. — Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8): 37, pl. 2, Fig. 5.
Australaugeneria rutilans. — Pettibone 1969 a, Proceedings of the Biological Society of Washington 82: 22, fig. 10.
Acholoe astericola. — Amoureux et al. 1978, Israel Journal of Zoology 27: 61‑62: fig. 1 [not Delle Chiaje, 1841 a].
Ty p e l o c a l i t y : Lapinig (= Lapinin) Island, Philippines.
S p e c i m e n s e x a m i n e d : S y n t y p e : Philippines, Lapinig Island, Coll. Semper, ZMB 1137. — R e d S e a : Microscope
slide, as Acholoe astericola, Aphroditides, Mer Rouge, 1973, IEA Z5. — T h e G u l f : 3 spms., 28 °N/49°36'50''E, “University of
Miami Expedition”, cruise 4, on octocoral Dendronephthya sp., det. Pettibone, 1983, USNM 080102.
M e a s u r e m e n t s : Syntype ZMB 1137: cs (37) 7/2/1.5 [af (25) + mf (3) + pf (9)]. IEA Z5: af (18) 3/1/0.8. USNM
080102: cs (26) 5/2/1.8; cs (22) 2.5/1/0.9; cs (33) 5/2/1.8.
D i a g n o s i s : Anteriormost prechaetal neuropodial lobes strongly elongate, enclosing strongly
bent hooks. Notochaetae with spines along convex edge.
D e s c r i p t i o n : Body dorsoventrally flattened, tapering posteriorly, short, with about 26 to
37 segments. Prostomium (Fig. 5 a) rounded to slightly hexagonal, slightly wider than long. Style
of median antenna smooth, very long, gradually tapering; lateral antennae similar, but shorter.
Palps smooth, conical, gradually tapering. Two pairs of small pointed eyes on posterior half of
prostomium, anterior pair on more or less widest part, posterior pair in front of hind margin. Ten-
tacular cirri of similar length, about as long as median antenna. Facial tubercle indistinct. Buccal
cirri moderate in length, not much longer than following ventral cirri. Parapodia of second seg-
ment with or without notochaetae. Prechaetal neuropodial lobes of anteriormost segments elon-
gate, enclosing strongly hooked neurochaetae (Fig. 5 i). Syntype and one specimen from The Gulf
reaching 33 segments show 15 pairs of elytra arranged as indicated in the generic diagnosis. Elytra
(Fig. 5 b) large, completely smooth, more-or-less oval, sometimes with a small notch. Styles of
dorsal cirri smooth, without a subterminal swelling, very long, extending far beyond neurochaetae.
Dorsal tubercles inconspicuous. Parapodia (Fig. 5 c, 5 d) with subconical notopodia of moder-
Fig. 5: Australaugeneria rutilans (USNM 080102). a: anterior end; b: elytra from anterior body; c: right parapodium from midbody
segment, anterior view; d: same, posterior view; e: notochaetae; f: upper neurochaeta; g: lower neurochaetae; h: neurochaetae from
posteriormost segments; i: hooked neurochaeta from anteriormost segments.
58 T. Wehe
ate size. Neuropodia long, slender, bluntly rounded distally, prechaetal lobe slightly longer than
postchaetal one. Ventral cirri smooth, gradually tapering, long, almost reaching tip of neuropodia.
Notochaetae (Fig. 5 e) few, slightly curved, sabre-like, dentate along convex border, with blunt,
unidentate tips (not bidentate, see remarks on generic diagnosis). Neurochaetae falcate, faintly
spinous or smooth, upper ones (Fig. 5 f ) more slender with moderate subdistal swelling and blunt
tips, lower ones (Fig. 5 g) slightly stouter with more prominent subdistal swelling below uni- or bi-
dentate tips. Upper neurochaetae in anterior segments with more distinct spines. Neurochaetae of
posteriormost segments with distinct bidentate tips (Fig. 5 h). Transverse bands of cilia sometimes
present dorsally and ventrally on body surface, patches of cilia also found dorsally and ventrally on
the parapodia.
R e m a r k s : The specimen mounted on a microscope slide from the collections of the IEA
is most probably one of the numerous specimens reported and described as Acholoe astericola by
Amoureux et al. (1978). The description and figures provided in this work suggested that the
specimens were misidentified, which was confirmed by an examination of the specimen.
A. rutilans may be easily distinguished from A. michaelseni by the presence of spines on the
notochaetae, but it is more difficult to separate from A. pottsi. According to Pettibone (1969 d),
the two first-mentioned species differ in the absence of notochaetae on segment two and more
strongly bent hooks in A. rutilans, whereas notochaetae on segment two are present in A. pottsi but
the hooks are not as strongly bent. The specimen (IEA Z5) has notochaetae on segment two and
also possesses strongly bent hooks, and so these two species may only represent variations of one
and the same species.
B i o l o g y & e c o l o g y : Referring to a note in the diary of Prof. Semper, Grube (1878) re-
marked that the species was found on Xenia sp. and was of the same red colour as its host. Accord-
ing to Pettibone (1969 a), the species is associated with alcyonarians.
D e p t h r a n g e : Unknown; reported elsewhere between 11 and 16 m (Pettibone 1969 a).
D i s t r i b u t i o n : Red Sea, The Gulf. The species and genus represent a new record for both
the Red Sea and The Gulf. Elsewhere: Reported from Philippines and South-West Australia. (Pet-
tibone 1969 a).
Gastrolepidia Schmarda, 1861

Gastrolepidia Schmarda, 1861. — Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857.
1. Band: Turbellarien, Rotatorien und Anneliden. Zweite Hälfte: 159.
Gastrolepidia. — Hanley 1989, The Beagle, Records of the Northern Territory Museum of Arts and Sciences 6(1): 16.
Ty p e s p e c i e s : Gastrolepidia clavigera Schmarda, 1861.
D i a g n o s i s : Body flattened dorsoventrally, elongate, up to 60 segments. Prostomium with-
out cephalic peaks. Ceratophores of lateral antennae inserted terminoventrally, converging mid-
ventrally. Median antenna with bulbous, clavate style. All antennae with a distinct terminal filum.
Palps stout, gradually tapering. Two pairs of eyes on posterior half of prostomium. Tentacular
and dorsal cirri shaped as style of median antenna. Buccal segment with nuchal fold. Elytra up to
31 pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, 35, 36, thereafter usually on alternat-
ing segments, sometimes varying and/or asymmetrical. Elytra large, smooth, with folds on anterior
margin where dorsal cirri of antecedent segments arise. Notopodia small, tapering to acicular proc-
ess. Neuropodia with well developed lobes, prechaetal one longer than postchaetal. 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 up-
per ones and slightly curved, unidentate tips in middle and lower ones. Anal cirri similar to other
cirri.
R e m a r k s : Gastrolepidia is monospecific and occurs as a commensal on holothurians

(Britayev & Zamishlyak 1994, Britayev et al. 1999, personal observation on the Great Barrier
Reef, Norman Reef ).
Gastrolepidia clavigera Schmarda, 1861 Fig. 6 a‑h

Gastrolepidia clavigera Schmarda, 1861 — Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis
1857. 1. Band: Turbellarien, Rotatorien und Anneliden. Zweite Hälfte: 159, pl. 36: fig. 316; Seidler 1923, Archiv für
Naturgeschichte 89 (Abt. A/11): 142; Hanley 1989, The Beagle, Records of the Northern Territory Museum of Arts and Sci-
ences 6(1): 17‑23, figs 8‑10; Britayev et al. 1999, Cahiers de Biologie Marine 40(4): 366‑368, figs 19‑23; Barnich et al.
2004, Species Diversity 9: 302‑303.
Ty p e l o c a l i t y : Sri Lanka (“Ceylon”), Indian Ocean.
S p e c i m e n s e x a m i n e d : A r a b i a n S e a , S o c o t r a I s l a n d : 1 spm., off Quara, 12°18.877'N/53°37.493'E,
15‑17 m, coll. M. Apel, 31.03.1999, SMF 13528; 1 spm., Quatananh Bay, west of the Ras, 12°21.280'N/53°32.614'E, 10‑11 m,
under corals, coll. M. Apel, 31.03.1999, SMF 13529.
M e a s u r e m e n t s : SMF 13528: cs (41) 15/6/4. SMF 13529: af (40) 24/8/6 [af (14) 7 + mf (16) 10 + mf (5) 4 + mf (5) 3].
D e s c r i p t i o n : Body flattened dorso-ventrally, tapering anteriorly and posteriorly. Pros-
tomium (Fig. 6 a) wider than long, oval to hexagonal, without cephalic peaks. Ceratophores of
antennae broad, cylindrical, styles broken off in both specimens. Palps long, gradually tapering
(very slender in SMF 13528), smooth. Two pairs of large eyes dorso-laterally on posterior half
of prostomium, close to each other. Tentacular cirri broken off. Facial tubercle distinct. Everted
pharynx on distal borders with two semicircles of 11 soft papillae. Buccal segment with distinct
semilunar nuchal fold, and long buccal cirri with clavate style and short terminal filum. 18 pairs
of elytra on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, 35, 36, 38 (SMF 13528) according
to arrangement of elytrophores. Elytra missing. Dorsal cirri (Fig. 6 b) with conical cirrophore and
long, stout, smooth, clavate style, abruptly terminating into slender terminal filum. Dorsal tuber-
cles inconspicuous. Parapodia (Fig. 6 c, 6 d) with elongate, subtriangular notopodia. Neuropodia
deeply incised with well developed lobes; prechaetal, acicular lobe slightly longer, subtriangular,
and postchaetal one shorter, bluntly rounded. Ventral cirri very small and slender, gradually taper-
ing. No nephridial papillae observed. Prominent ventral lamellae (Fig. 6 e) on bases of parapodia
from segment 3 onwards. Notochaetae (Fig. 6 f ) moderate in number, short to long, curved, with
numerous rows of spines and slightly notched tips. Neurochaetae (Fig. 6 g, 6 h) longer, upper slen-
der, with numerous rows of spines below slightly notched tips, middle and lower ones stouter, with
fewer rows of spines below hooked, unidentate tips.
R e m a r k s : The genus is monospecific. Although the elytra are missing the specimens clearly
belong to Gastrolepidia clavigera.
B i o l o g y & e c o l o g y : The species is found as a commensal on holothurians. Britayev &
Zamishlyak (1994, 1996) provided detailed information on the biology of this species.
D e p t h r a n g e : So far recorded between 10 and 17 m; reported elsewhere from the intertidal
down to 20 m (Hanley 1989, Britayev et al. 1999).
D i s t r i b u t i o n : Arabian Sea: Socotra Archipelago. This is the first record of the species from
the Arabian Sea. Elsewhere: Throughout tropical Indo-West Pacific: i.a. Madagascar, Sri Lanka,
South China Sea, Papua New Guinea, Bismarck Sea, Australia, Western Pacific Islands (Hanley
1989, Britayev et al. 1999, Barnich et al. 2004).
Iphioninae Kinberg, 1856

D i a g n o s i s : Body short, oval, flattened ventrally and arched dorsally. Segments less than
40 (29 in Iphione and Iphionella, up to 39 in Iphionides Hartmann-Schröder, 1977). Prostomium
without cephalic peaks, with two lateral antennae inserted terminally or with short, rudimentary
60 T. Wehe
Fig. 6: Gastrolepidia clavigera (SMF 13529; b from SMF 13528). a: anterior end; b: dorsal cirrus; c: right parapodium from mid-
body segment, anterior view; d: same, posterior view; e: body in ventral view with ventral lamellae; f: notochaeta; g: upper neuro-
chaeta; h: lower neurochaeta.
anterolateral extensions only. Median antenna absent or represented by a small occipital papilla
(rarely with a distal style). Palps stout, conical. With or without two pairs of eyes. Tentaculophores
with chaetae. Second (buccal) segment with or without nuchal fold. Elytra 13 pairs in Iphione and
Iphionella on segments 2, 4, 5, 7, alternating to 23, 27, or up to 20 pairs continuing on alternate
segments behind 23 in Iphionides. Elytra large, covering dorsum, stiff, tightly imbricate, surface
divided into honeycomb-like compartments; margins with or without fringing papillae. Dorsal
cirri on papillate styles. Dorsal tubercles bulbous. Notopodia of moderate size, with acicular proc-
ess. Neuropodia dorsally and ventrally not deeply incised; prechaetal, acicular lobe with acicular
process; postchaetal lobe shorter, rounded. Notochaetae very numerous, forming thick bundles,
slender, bipinnate, with capillary tips, or moderate in number, spinous. Neurochaetae numerous,
with uni- or bidentate tips. Pygidium without anal cirri.
R e m a r k s : The authorship of Iphioninae has usually been assigned to Baird (1865) by e.g.
Pettibone (1986 a), Hanley & Burke (1991 a). However, as for the Polynoidae and Polynoinae,
the first available name for the family group according to §36 of the ICZN is Iphionea in Kinberg
(1856). See remarks on history above.
Iphione Kinberg, 1856

Iphione Kinberg, 1856. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 12 (9&10): 383.
Iphione. — Pettibone 1986 a, Smithsonian Contributions to Zoology 428: 6; Hanley & Burke 1991 a, Mémoires du Muséum
National d’Histoire Naturelle (A) 151: 38‑39.
Ty p e s p e c i e s : Polynoe muricata Savigny in Lamarck, 1818.
D i a g n o s i s : Body stout, arched dorsally and flattened ventrally, chiton-like in appearance,
with 29 segments. Prostomium withdrawn into anterior segments. Two papillate, lateral antennae
inserted terminally, with anterior prolongations of prostomium forming long ceratophores. Palps
finely papillate. Two pairs of eyes. With or without occipital papilla on posterior part of prosto-
mium. Bulbous facial tubercle present. Buccal segment with nuchal fold. Third segment not visible
dorsally, parapodia wedged between those of segments two and four. Elytra 13 pairs on segments
2, 4, 5, 7, alternating to 23, 27, tightly imbricate, covering dorsum completely; surface with hexa-
or polygonal honeycomb-like areas with secondary areolae, with or without spinous tubercles;
margins with or without fringing papillae. Dorsal cirri with prominent cylindrical cirrophores
and papillate styles. Dorsal tubercles prominent, transversely elongate, flattened. Notopodia small,
with acicular process. Neuropodia large, conical with a projecting prechaetal, acicular lobe and a
rounded postchaetal one, both lobes with small papillae distally. Notochaetae very numerous, in
dense bundles, slender, bipinnate, with fine capillary tips. Neurochaetae numerous, stout, with
several rows of spines below slightly hooked, blunt, uni- or bidentate tips. Dorsal anal ridge on
segments 23‑29.
Key to the species of Iphione
1 Elytra with smooth margins I. ovata

– Elytra with fringing papillae....................................................................................... 2
2 Large, erect conical tubercles on posterolateral surface of elytra. Fringing papillae in-
serted submarginally I. muricata
– Small, conical tubercles on posterolateral surface. Fringing papillae inserted margin-
ally or even ventrally I. henshawi
62 T. Wehe
Iphione henshawi Pettibone, 1986 a Fig. 7 a‑h

Iphione henshawi Pettibone, 1986 a. — Smithsonian Contributions to Zoology 428: 21‑23, figs 8, 9.
Ty p e l o c a l i t y : Hilo, Hawaii, Pacific Ocean.
S p e c i m e n s e x a m i n e d : R e d S e a , S u d a n : 1 spm., Sanganeb Atoll, south-west tip, 40 m, from living Seriatopora
hystrix, 2 April 1991, SMF 13609.
M e a s u r e m e n t s : cs (29) 12/7/6.
D i a g n o s i s : Elytra posterolaterally with several low, conical, spinous tubercles with few distal
papillae; margin without smooth areas, with digitiform, spinous fringing papillae inserted margin-
ally or ventrally.
D e s c r i p t i o n : Prostomium (Fig. 7 a) without distinct occipital papilla. Styles of lateral antennae
papillate, slightly longer than ceratophores, gradually tapering. Palps stout with several rows of small
papillae. Two pairs of eyes on posterior half of prostomium, anterior pair on more or less widest part,
posterior one in front of hind margin almost hidden by nuchal fold. Tentacular cirri shaped as antennae.
Nuchal fold with two lateral protrusions, covering posterior margin of prostomium. Buccal cirri longer
and inserted more proximally than following ventral cirri. A pair of small dorsal nodules present behind
nuchal fold. Elytra (Fig. 7 b) yellowish to brown, thick, stiff, tightly imbricate, covering body completely.
First pair small, rounded to oval, second and third pair curved, boomerang-shaped, with a broad notch
anteriorly, posterior ones larger, more or less reniform, last pair small with median notch leaving anus
uncovered. Surface of elytra completely covered with hexa- or polygonal, honeycomb-like compartments
with secondary areolae (Fig. 7 c), no smooth rim. Honeycomb compartments of median and anterior
area more prominent, posterior ones becoming smaller. Posterolaterally with several low, conical tuber-
cles, mostly with few long papillae on their tips (Fig. 7 d). Laterally with several club-shaped papillae, in-
serted marginally or even ventrally, column spinous, distally with long, slender papillae (Fig. 7 e). Elytra
not encrusted. Elytrophores large, flattened, very broad. Dorsal tubercles similar to elytrophores. Dorsal
cirri with large, cylindrical cirrophore and long, papillate style with terminal filum, extending beyond
chaetae. Parapodia (Fig. 7 f) with distinct notopodia, more or less conical. Neuropodia with a more-or-
less subtriangular, prechaetal lobe and slightly shorter, bluntly rounded, postchaetal one. Both lobes with
minute papillae distally. Ventral cirri papillate, short, gradually tapering. Notochaetae (Fig. 7 g) numer-
ous, in dense bundles, whitish, fine, bipinnate (feathery), tapering to capillary tips. Neurochaetae (Fig.
7 h) copper-coloured, stout, numerous, subdistally with several rows of spines and slightly curved, blunt,
unidentate tips. Neurochaetae of few anterior segments more slender, tapering to pointed tips.
R e m a r k s : The specimen examined agrees very well with the description given by Pettibone
(1986 a). The elytra have only been removed from some anterior segments, in order not to damage
the rare specimen more than necessary, and so the dorsal nodules were observed on the segments
behind the nuchal fold only. Nephridial papillae were not observed. There are no large spinous
tubercles on the surface of the elytra, which clearly separates this species from I. muricata. Fur-
thermore, there are no smooth rims on the elytra and the fringing papillae are distinctly inserted
marginally or even ventrally. The neurochaetal spines are also coarser than in I. muricata.
B i o l o g y & e c o l o g y : Unknown, probably associated with corals.
D e p t h r a n g e : So far recorded from 40 m; elsewhere reported from 13‑271 m (Pettibone
1986 a).
D i s t r i b u t i o n : Red Sea. This is the first record since the type specimens. Elsewhere: Hawai-
ian Islands (Pettibone 1986 a).
Iphione muricata (Savigny in Lamarck, 1818) Fig. 8 a‑h

Polynoe muricata Savigny in Lamarck, 1818 . — Histoire naturelle des Animaux sans vertèbres... vol 5: 308; Savigny 1822,
Description de l’Égypte 1(3): 21; Savigny 1826, Description de l’Égypte. 2nd ed., Histoire naturelle. Minéralogie.-Zoologie,
Animaux Invertébrés 21: 347.
Fig. 7: Iphione henshawi (SMF 13609). a: anterior end; b: elytron from left anterior segment; c: compartment with secondary are-
olae on elytra; d: tubercle on elytra surface; e: tubercle on elytra margin; f: left parapodium from segment 5, posterior view; notopo-
dium not visible; g: notochaeta; h: neurochaeta.
Iphione muricata. — Kinberg 1857, Ventenskapliga Iakttagelser på H. Maj: t Konung Oscar den Förstes befallning utgifna af K.
Svenska Vetenskaps-Akademien. Zoologi 1: 8; Quatrefages 1866, Histoire naturelle des Annelés marins et d’eau douce. An-
nélides et Géphyriens: 266‑268; Gravier 1902, Nouvelles Archives du Muséum 4 e série 3: 226‑231, textfigs 232‑239, pl. 9:
figs 129‑135; Fauvel 1919, Archives de Zoologie Expérimentale et Générale 58: 334; Fauvel 1933, Mémoires de l’Institut
64 T. Wehe
d’Égypte 21: 40‑41; Monro 1939 b, Annals and Magazine of Natural History 11(4): 168 [partim, see I. ovata]; Fauvel
1955, Annales de l’Institute Océanographique Paris 30: 102; Fauvel 1957, State of Israel Ministry of Agriculture Division of
Fisheries. The Sea Fisheries Research Station, Haifa 13(7): 3; Fauvel 1958, Israel Ministry of Agriculture Division of Fisher-
ies. The Sea Fisheries Research Station, Haifa 16(7): 1; Storch 1967 a, Kieler Meeresforschungen 23(2): 154‑155, figs 1‑3,
5; Fishelson & Rullier 1969, Israel Journal of Zoology 18: 53; Kiseleva 1971, Naukova Dumka, Kiev: 45; Ben-Eliahu
1972 b, Israel Journal of Zoology 21: 195; Amoureux et al. 1978, Israel Journal of Zoology 27: 68; Pettibone 1986 a,
Smithsonian Contributions to Zoology 428: 9‑16, figs 1‑5; Barnich et al. 2004, Species Diversity 9: 291‑292.
Harmothoe dictyophora. — Fauvel 1927, The Transactions of the Zoological Society of London 22/4(1): 414 [not Grube 1878].
Ty p e l o c a l i t y : Île de France (= Mauritius), Indian Ocean.
S p e c i m e n s e x a m i n e d : S u e z C a n a l : 1 spm. as Harmothoe dictyophora, Egypt, Suez Canal, coll. “Cambridge Ex-
pedition to the Suez Canal”, 12 October 1924, BMNH 1926.11.12.138; 1 microscope slide, Suez, (probably belongs to BMNH
1926.11.12.138), IEA R/76. — R e d S e a : 1 spm., Mer Rouge, Coll. de M. le Baron de Saint-Joseph, 1911, MNHN A274;
1 spm., coll. E. Rüppell, BMNH 2003.95; 1 spm., Coll. Mus. Marseille, ZMUC-POL 1366; 3 spms., coll. E. Rüppell, 1827,
SMF 2641; 8 spms., Egypt, Suez, coll. Jousseaume, 1894, det. M.Ch. Gravier, 1901, MNHN A274; 1 spm., Egypt, Gulf of Suez,
28°14'N/33°23'E, 22 m, muddy sand and madreporarians, coll. R.Ph. Dollfus, 25 December 1928 [collection data supplemented
according to Billard (1933)], MNHN A274; 20 spms., Egypt, Suez, coll. Jousseaume, 1894, det. M.Ch. Gravier, 1901, MNHN
A274; 3 spms., coll. Jousseaume, 1893, MNHN A28; 1 spm., Egypt, Suez Bay, entrance of canal, coll. Gruvel & Moazzo, Jan.
- Feb. 1934, MNHN A894; 1 spm., epi et endofaune de Méandrines, coll. Drach, Cherbonnier & Mercier, 1952, MNHN A364;
1 spm., Egypt, Suez, coll. E. Bannwarth (probably 1912‑13 according to Schäfer, 1963), det. Michaelsen, ZMH V8365; 1 spm.,
Egypt, Gulf of Suez, African coast, Gimsah Bay, coll. Hartmeyer, det. Augener 1920, ZMH V6120; 1 spm., Egypt, Suez, coll. E.
Bannwarth 1914 (probably 1912‑13 according to Schäfer 1963), ZMH P‑E186; 1 spm., coll. E. Bannwarth, Cairo S.V., 10 Janu-
ary 1913, SMF 13633; 2 spms., coll. E. Bannwarth, Cairo S.V., 31 January 1913, SMF 13625; 2 spms., coll. E. Bannwarth, Cairo
S.V., 16 May 1914, SMF 13627; 1 spm., Egypt, Gulf of Suez, coll. McAndrew's Expedition, BMNH 1869.7.8.7; 4 spms., Egypt,
Gulf of Suez, coll. J.K. Lord, BMNH 1870.12.23.34; 1 spm., Sudan, Sanganeb Atoll, north jetty, western side, 1 m, from dead Sty-
lophora sp., stones, 8 April 1991, SMF 13629; 1 spm., Sudan, Sanganeb Atoll, south jetty, western side, 1 m, from dead Stylophora
sp., 6 April 1991, SMF 13622; 1 spm., Sudan, Sanganeb Atoll, inner reef, 12 m, from dead Stylophora pistillata, 15 April 1991,
SMF 13630; 3 spms., Sudan, Sanganeb Atoll, south jetty, eastern side, 1 m, from dead Stylophora pistillata, 4 April 1991, SMF
13626; 1 spm., Sudan, Sanganeb Atoll, south-eastern top of the reef, 1 m, from dead Stylophora pistillata, coll. D. Fiege, 26 Septem-
ber 1992, SMF 13634; 1 spm., Sudan, Sanganeb Atoll, north jetty, 1 m, from dead Stylophora sp., coll. D. Fiege, 1 October 1992,
SMF 13632; 1 spm., Sudan, Suakin, coll. P. Robinson, BMNH 1885.8.3.1; 1 spm., Sudan, “Pearl fishery investigations”, coll. C.
Crossland, October 1904 to May 1905, BMNH 1924.6.18.211‑220; 6 spms., coll. C. Crossland, BMNH 1941.4.4.168‑172;
1 spm., Eritrea, near Assab, bay between Gt. Hanish and Suyul Hanish Islands, 13°39'30''N/42°43'00''E, dredge, 29 m, sand,
shells, corals, coll. “John Murray Expedition”, H.E.M.S. Mabahiss, 17 September 1933, BMNH 1937.9.2.44‑45; 2 spms., as
Iphione sp., Eritrea, Bay of Assab, Sanabor Island, “R/V Akademik Petrovsky”, cruise 9, A.B. Tzetlin, 13 February 1980, ZMUM;
1 spm., Bay of Assab, Sanabor Island, “R/V Akademik Petrovsky”, cruise 9, A.B. Tzetlin, 14 February 1980, ZMUM; 1 spm., Bay
of Assab, Sanabor Island, “R/V Akademik Petrovsky”, cruise 9, A.B. Tzetlin, 14 February 1980, ZMUM; 1 spm., Assab, Eritrea,
25 June 1884, NHMW V310; 22 spms., “Pola Expedition”, 1895‑1898, NHMW V313; 2 spms., Jiddah, Saudi-Arabia, “Pola
Expedition”, December 1895, NHMW V17054. — G u l f o f A d e n : 8 spms., Djibouti, Gulf of Tadjoura, Obock, coll. Jous-
seaume, 1895, MNHN A274; 1 spm., Djibouti, Gulf of Tadjoura, Musha Islands, Maskalla Island, coll. M.Ch. Gravier, 23 January
1904, MNHN A274; 4 spms., Djibouti, Gulf of Tadjoura, coll. Coutière, 1897, MNHN A274; 1 spm., Djibouti, Gulf of Tad-
joura, sand, east of residence, coll. M.Ch. Gravier, 1904, MNHN A274; 2 spms., Djibouti, Gulf of Tadjoura, Musha Islands, from
polyps, coll. M.Ch. Gravier, 23 January 1904, MNHN A274; 1 spm., Djibouti, Gulf of Tadjoura, from madreporarian coral, coll.
M.Ch. Gravier 17 January 1904, MNHN A274; 4 spms., Djibouti, Gulf of Tadjoura, Pingoin and Metéore reefs, dredge, 18‑20 m,
coll. M.Ch. Gravier, 27 January 1904, MNHN A274; 1 microscope slide, Djibouti, M.Ch. Gravier, IEA D‑54. — A r a b i a n
S e a : 1 spm., Socotra Island, near Hawlaf, west of the jetty, 12°40.888'N/54 4.556'E, at Galaxea astreata community, 6‑8 m, coll.
M. Apel, 28 March 2000, SMF 13623; 1 spm., Socotra Island, near Qadub, between Hadibo and airport, 7‑8 m, 13 March 1999,
SMF 13631. — T h e G u l f : 1 spm., Karan Island, near Jubail, Kingdom of Saudi Arabia, under coral rubble, 10‑12 m, coll. M.
Apel, 16 July 1992, SMF 13624; 1 spm., Karan Island, near Jubail, Kingdom of Saudi Arabia, under coral rubble, 5‑9 m, coll. M.
Apel, 18 July 1992, SMF 13628.
M e a s u r e m e n t s : BMNH 1926.11.12.138: cs (28) 7/5/4. MNHN A274: cs (29) 18/13/11. BMNH 2003.95: cs (29)
18/13/10. ZMUC-POL 1366: cs (29) 29/13/10. SMF 2641: cs (29) 20/12/10, cs (29) 20/13/10, cs (29) 25/13/11. MNHN
A274: cs (29) 25/15/11, cs (29) 16/10/7, cs (29) 16/10/7, cs (29) 14/10/7, cs (29) 15/9/12, cs (29) 18/12/8, cs (29) 18/9/6, cs
(29) 15/11/8. MNHN A274: cs (29) 16/12/9. MNHN A274: cs (29) 17/12/8, cs (29) 13/8/6, cs (29) 14/11/8, cs (29) 15/11/8, cs
(29) 18/13/9, cs (29) 12/8/6, cs (29) 20/12/9, cs (29) 20/13/9, cs (29) 20/13/9, cs (29) 13/11/8, cs (29) 12/8/6, cs (29) 12/9/6, cs
(29) 18/12/9, cs (29) 11/8/6, cs (29) 12/11/8, cs (29) 14/11/7, cs (29) 12/9/7, cs (29) 14/13/9, cs (29) 13/10/8, cs (29) 15/11/8.
MNHN A28: cs (29) 18/13/9, cs (29) 20/12/8, cs (29) 23/14/10. MNHN A894: cs (29) 20/14/10. MNHN A364: cs (29) 12/7/5.
ZMH V8365: cs (29) 22/16/11. ZMH V6120: cs (29) 20/13/9. ZMH P‑E186: cs (29) 24/15/12. SMF 13633: cs (29) 19/13/9.
Fig. 8: Iphione muricata (SMF 13628: a, c-h; NHMW V310: b). a: anterior end; b: 7th right elytron with submarginally inserted
fringing papillae directed upwards; c: compartment with secondary areolae on elytra d: spinous tubercle on elytra surface; e: fring-
ing papillae; f: left parapodium from segment 16, posterior view; g: notochaeta; h: neurochaeta.
SMF 13625: cs (29) 26/14/11, cs (29) 19/14/10. SMF 13627: cs (29) 24/13/10, cs (29) 25/13/10. BMNH 1869.7.8.7: cs (29)
15/13/11. BMNH 1870.12.23.34: cs (29) 21/11/9, cs (29) 22/13/10, cs (29) 21/12/9, cs(29) 17/10/7. SMF 13629: cs (29) 7/5/3.
SMF 13622: cs (29) 6/3.5/2.5. SMF 13630: cs (29) 8/4/3. SMF 13626: cs (29) 12/9/6, cs (29) 12/7/5, cs (29) 10/7/5. SMF
13634: cs (29) 12/9/7. SMF 13632: cs (29) 10/8/6. BMNH 1885.8.3.1: cs (29) 13/9/6. BMNH 1924.6.18.211‑220: cs (29)
8/5/4. BMNH 1941.4.4.168‑172: cs (29) 20/14/11, cs (29) 14/11/8, cs (29) 15/13/10, cs (29) 13/10/8, cs (29) 8/6/3, cs (29)
9/7/5. BMNH 1937.9.2.44‑45: cs (29) 14/8/5. ZMUM: cs (29) 14/11/8, cs (29) 8/7/5. ZMUM: cs (29) 11/8/6. ZMUM: cs (29)
66 T. Wehe
8/5/3.5. NHMW V310: cs (29) 17/7/5. NHMW V313: cs (29) 9/4/3, cs (29))9/4.5/3, cs (29) 13/7/5, cs (29) 12/5/3.5, cs (29)
10/5/3.5, cs (29) 11/5/3.5, cs (29) 13/6/4, cs (29) 16/7/4.5, cs (29) 16/7/4, cs (29) 14/6/4, cs (29) 21/9/6, cs (29) 22/10/6, cs (29)
15/7/5, cs (29) 17/8/5, cs (29) 16/7/5, cs (29) 17/7/4.5, cs (29) 15/7/5, cs (29) 18/8/6, cs (29) 25/10/7, cs (29) 18/8/6, cs (29)
16/7/5, cs (29) 17/8/6. NHMW V17054: cs (29) 16/8/6, cs (29) 15/7/5. MNHN A274: cs (29) 14/11/7, cs (29) 22/14/10, cs
(29) 19/12/8, cs (29) 20/13/9, cs (29) 17/9/6, cs (29) 15/13/9, cs (29) 15/11/7, cs (29) 15/10/7. MNHN A274: cs (29) 5/4/2.5.
MNHN A274: cs (29) 19/10/7, cs (29) 24/13/9, cs (29) 15/11/8, cs (29) 11/8/5. MNHN A274: cs (29) 15/11/8. MNHN A274:
cs (29) 9/7/5, cs (29) 15/8/6. MNHN A274: cs (29) 16/9/7. MNHN A274: cs (29) 9/4/3, cs (29) 10/5/3, cs (29) 15/8/6, cs (29)
11/6/4. SMF 13623: cs (29) 25/14/10. SMF 13631: cs (29) 12/8/6. SMF 13624: cs (29) 16/12/8. SMF 13628: cs (29) 20/14/10.
D i a g n o s i s : Elytra posterolaterally with few rows of stout, spinous, slightly bent tubercles;
margin with spinous fringing papillae.
D e s c r i p t i o n : Prostomium (Fig. 8 a) sometimes with occipital papilla, often hidden from
view by nuchal fold or missing. Styles of lateral antennae papillate, slightly longer than ceratophores,
tapering to distinct terminal filum. Palps stout with several rows of small papillae. Two pairs of eyes
on posterior half of prostomium, anterior pair on more or less widest part, posterior one in front of
hind margin of prostomium. Tentacular cirri shaped as antennae. Rounded nuchal fold on buccal seg-
ment mostly covering posterior part of prostomium. Buccal cirri longer and inserted more proximally
than following ventral cirri. Some anterior segments with a pair of dorsal nodules (always on 4 and 5,
variable on other segments); eventually additional pairs on more posterior segments. Elytra (Fig. 8 b)
yellowish to reddish-brown, thick, stiff, tightly imbricate, covering body completely. First pair small,
rounded to oval, second and third pair curved, boomerang-shaped, with a broad notch anteriorly,
following ones larger, more or less reniform, last pair small with a median notch, leaving anus uncov-
ered. Elytra surface almost completely covered with hexa- or polygonal, honeycomb-like compart-
ments with secondary areolae (Fig. 8 c), anteriorly and laterally eventually with small, smooth rim.
Honeycomb compartments of median and anterior part more prominent, posterior ones becoming
smaller. Posteriorly and laterally with one to few, curved rows of large, spinous tubercles (Fig. 8 d),
usually erected diagonally in posterior direction, slightly bent, with larger spines on posterior surface.
Additional, more or less spinous tubercles of different development occurring on posterior part of the
elytra. This part often thickly encrusted, hiding honeycombs and tubercles. Laterally with long fring-
ing papillae (Fig. 8 e) inserted submarginally on surface, column spinous, distally with long, slender
papillae. Elytrophores large, flattened, very broad. Dorsal tubercles similar to elytrophores. Dorsal
cirri with large, cylindrical cirrophore and long, papillate style with terminal filum, extending beyond
chaetae. Parapodia (Fig. 8 f) with distinct notopodia, more or less conical. Neuropodia with subtrian-
gular, prechaetal lobe, and slightly shorter, bluntly rounded, postchaetal one. Both lobes distally with
minute papillae. Ventral cirri papillate, short, gradually tapering. Notochaetae (Fig. 8 g) numerous, in
dense bundles, whitish, bipinnate (feathery), tapering to capillary tips. Neurochaetae (Fig. 8 h) cop-
per-coloured, stout, numerous, subdistally with several rows of fine spines and slightly curved, blunt,
R e m a r k s : I. muricata is easily distinguished from I. ovata by the presence of fringing papil-
lae on the elytra (see also the remarks under I. ovata). Depending on the size of the specimens, this
fringe is more or less prominent and the large dorsal tubercles on the surface may also be more or
less developed.
B i o l o g y & e c o l o g y : The life pattern of I. muricata was described by Storch (1967 a)
and, together with its body shape it is very similar to the Polyplacophora. Individuals rest during
the day, clinging to stones or corals, and only moving a little during the night. Individuals are not
capable of free swimming or wriggling movements. The species typically occurs in the intertidal
and shallow waters, often associated with corals.
D e p t h r a n g e : So far recorded between 1 and 22 m; elsewhere reported down to 60 m
(Pettibone 1986 a).
D i s t r i b u t i o n : Suez Canal, Red Sea, Gulf of Aden, Arabian Sea: Socotra Archipelago, The
Gulf. The species is here recorded for the first time from the Arabian Sea and The Gulf. Elsewhere:
Reported throughout the Indian and Pacific Oceans (Pettibone 1986 a, Barnich et al. 2004).
Iphione ovata Kinberg, 1856 Fig. 9 a‑i, Pl. 1 a

Iphione ovata Kinberg, 1856. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 12(9&10): 383; Kinberg 1857,
Ventenskapliga Iakttagelser på H. Maj: t Konung Oscar den Förstes befallning utgifna af K. Svenska Vetenskaps-Akademien.
Zoologi 1: 8, pl. 3: fig. 8; Pettibone 1986 a, Smithsonian Contributions to Zoology 428: 16‑19, fig. 6; Barnich et al.
2004, Species Diversity 9: 292.
Iphione muricata. — Monro 1939 b, Annals and Magazine of Natural History 11(4): 168 [partim, see I. muricata; not Savigny
in Lamarck, 1818].
Iphione reticulata. — Amoureux et al. 1978, Israel Journal of Zoology 27: 68‑69, fig. 2 [new synonymy].
Ty p e l o c a l i t y : Honolulu, Oahu, Hawaii, Pacific Ocean.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Iphione reticulata, Red Sea, Gulf of Aqaba, artificial lagoon at Elat, coll. L.
Fishelson, MNHN POLY TYPE 1430 [formerly IEA NS. 7039] + microscope slide, IEA Y97’. — R e d S e a : 1 spm., coll. E.
Rüppell, 1827, SMF 13566; 7 spm., “Pola Expedition”, 1895‑1898, NHMW 17060; 3 spm., Gulf of Suez, Ras Kanisa, about 1 km
offshore, 1‑2 m, coral reef, from dead corals, coll. C. Lewinsohn, 17 October 1972, NMNHT VR 25061; 3 spm., Gulf of Aqaba,
2 km north of Dahab, reef flat, 0‑2 m, from dead corals, coll. C. Lewinsohn, 5 November 1981, NMNHT VR 25060; 1 spm.,
Gulf of Aqaba, Elat, coll. J. Dafni, 1 June 1973, NMNHT VR 25062; 1 spm., Gulf of Aqaba, Elat, coll. 10‑12 October 1949,
BMNH 1956.8.8.14; 1 spm., as I. muricata in Monro (1939 b), Sudan, Suakin harbour, on coral, coll. C. Crossland, BMNH
1941.4.4.173; 1 spm., Sudan, Port Sudan, Wingate Reef, 5 m, from living Pocillopora sp., coll. D. Fiege, 20 September 1992, SMF
13572; 1 spm., Sudan, Sanganeb Atoll, inner reef, 12 m, from living Acropora sp., 9 April 1991, SMF 13569; 1 spm., Sudan, San-
ganeb Atoll, north jetty, edge of the lagoon, 1 m, from dead Pocillopora sp., coll. D. Fiege, 29 September 1992, SMF 13570; 1 spm.,
Sudan, Sanganeb Atoll, eastern side, south jetty, 1 m, from living Stylophora pistillata, 4‑15 April 1991, SMF 13565; 1 spm., Sudan,
Sanganeb Atoll, eastern side, north jetty, 1 m, from dead Stylophora sp., 5 April 1991, SMF 13567; 1 spm., Sudan, Sanganeb Atoll,
western side, north jetty, 1 m, from living Stylophora pistillata, 8 April 1991, SMF 13568; 1 spm., Sudan, Wingate Reef, 5 m, from
dead Pocillopora sp., coll. D. Fiege, 21 September 1992, SMF 13562; 1 spm., Sudan, Sanganeb Atoll, north jetty, 1 m, from dead
Stylophora sp., coll. D. Fiege, 1 October 1992, SMF 13563; 1 spm., Sudan, Wingate Reef, middle of reef, 5 m, from dead corals,
coll. D. Fiege, 29 September 1992, SMF 13564; 2 spms., Sudan, Wingate Reef, middle of reef, 5 m, from dead corals, coll. D.
Fiege, 22 September 1992, SMF 13571. — A r a b i a n S e a : 1 spm., Socotra Archipelago, Abd al-Kuri Island, west coast, off fish-
ing village, 12°13.870'N/52°03.774'E, 10 m, under coral rubble, coll. M. Apel, 4 April 2000, NHCY; 1 spm., Socotra Island, Ras
Irisal, Ghales, 12°31.876'N/54°32.444'E, 16‑17 m, under stones, coll. M. Apel, 10 April 2000, SMF 13574; 1 spm., Socotra Is-
land, Qalansiyah Bay, 12°41.026'N/53°28.309'E, 7‑8 m, under rocks, coll. M. Apel, 10 March 1999, SMF 13573; 1 spm., Socotra
Island, off Hawlaff dune, 12°40.519'N/54°4.170'E, 4‑5 m, coll. M. Apel, 21 March 1999, NHCY.
M e a s u r e m e n t s : Holotype MNHN POLY TYPE 1430: cs (28) 7/3/2. SMF 13566: cs (29) 10/5/3.5. NHMW 17060: cs
(29) 9/4/3, cs (29) 17/7/5, cs (29) 8/4/3, cs (29) 10/5/3.5, cs (29) 15/7/5, cs (29) 6/3/2.5, cs (29) 8/4/3. NMNHT VR 25061: cs
(29) 23/12/9, cs (29) 21/11/9, cs (29) 19/10/8. NMNHT VR 25060: cs (29) 10/6/4, cs (29) 10/7/5, cs (29) 13/7/5. NMNHT VR
25062: cs (29) 14/9/7. BMNH 1956.8.8.14: cs (29) 10/5/4. BMNH 1941.4.4.173: cs (29) 13/7/5. SMF 13572: cs (29) 12/7/5.
SMF 13569: cs (29) 6/4/3. SMF 13570: cs (29) 5/4/3. SMF 13565: cs (29) 10/5/4. SMF 13567: cs (29) 10/7/6. SMF 13568: cs
(29) 10/6/5. SMF 13562: cs (29) 13/10/8. SMF 13563: cs (29) 8/6/4. SMF 13564: cs (29) 8/5/4. SMF 13571: cs (29) 5/4/3, cs
(22) 3/2.5/1.8. NHCY: cs (29) 11/5/4. SMF 13574: cs (29) 11/8/6. SMF 13573: cs (29) 20/13/9. NHCY: cs (29) 20/10/7.
D i a g n o s i s : Elytra posterolaterally with numerous, conical, spinous tubercles; margin
smooth, without fringing papillae.
D e s c r i p t i o n : Prostomium (Fig. 9 a) with occipital papilla (sometimes absent), often hid-
den by nuchal fold. Styles of lateral antennae papillate, slightly longer than ceratophores, tapering
to a distinct terminal filum. Palps stout with several rows of small papillae. Two pairs of eyes on
posterior half of prostomium, anterior pair on more or less widest part, posterior one in front of
hind margin of prostomium. Tentacular cirri shaped as antennae. Rounded nuchal fold on buccal
segment covering posterior part of prostomium. Buccal cirri longer and inserted more proximally
than following ventral cirri. Some anterior segments (4‑6) with a pair of dorsal nodules (sometimes
indistinct), some posterior segments (about 17‑22) with a single middorsal nodule. Elytra (Fig.
9 b) yellowish to reddish-brown in ethanol, thick, stiff, tightly imbricate, covering body com-
pletely. First pair small, rounded to oval, second and third pair curved, boomerang-shaped, with a
68 T. Wehe
Fig. 9: Iphione ovata (NHMW V313: a-e; BMNH 1956.8.8.14: f-i). a: anterior end; b: 6th right elytron; c: compartments on elytra;
d: secondary areola; e: tubercle on elytra surface; f: left parapodium from segment 16, posterior view; g: papillae on lateral margin
of parapodia; h: notochaeta; i: neurochaeta.
broad notch anteriorly, following ones larger, more or less reniform, last pair small with median notch,
leaving anus uncovered. Elytra surface almost completely covered with hexa- or polygonal, honeycomb-
like compartments with secondary areolae (Fig. 9 c, 9 d), anteriorly and laterally with small, smooth
rim. Honeycomb compartments of median and anterior area more prominent, posterior ones becoming
smaller. Posterolateral surface with numerous spinous, slender, conical tubercles, distally with prominent
spines, erected diagonally in posterior direction (Fig. 9 e). Additional, more or less spinous tubercles of
different development occur in this part of elytra. Elytra often thickly encrusted, hiding honeycombs
and tubercles. Well developed tubercles as described only visible in larger and very clean specimens.
Elytrophores large, flattened, very broad. Dorsal tubercles similar to elytrophores. Dorsal cirri with large,
cylindrical cirrophore and long, papillate style with terminal filum, extending beyond chaetae. Parapo-
dia (Fig. 9 f) with distinct notopodia, more or less conical. Neuropodia with a subtriangular, prechaetal,
acicular lobe, and a slightly shorter, bluntly rounded, postchaetal one. Both lobes with minute papillae
distally (Fig. 9 g). Ventral cirri papillate, short, gradually tapering. Notochaetae (Fig. 9 h) numerous, in
dense bundles, whitish, slender, bipinnate (feathery), tapering to capillary tips. Neurochaetae numerous
(Fig. 9 i), copper-coloured, stout, subdistally with several rows of fine spines and slightly curved, blunt,
R e m a r k s : I. ovata can be clearly distinguished from all other species of the genus by the
absence of fringing papillae on the elytra. In comparison to I. muricata, the spinous tubercles are
rather slender and not arranged regularly in rows. However, this character is rather variable and not
useful for distinguishing between the two species, especially there are not enough specimens avail-
able for comparison. On average I. ovata appears to be smaller than I. muricata (compare measure-
ments) and tends to be brightly coloured and more yellowish or orange in ethanol, while the latter
is larger and appears darker and more brownish.
Amoureux et al. (1978) described Iphione reticulata. This species was not treated in the review
of the subfamily by Pettibone (1986 a), since the holotype was not available on loan from the IEA
where it was assumed to be deposited. While working in the Museum National d’Histoire Naturelle
in Paris, the holotype of I. reticulata was re-discovered in the collection there. The incomplete original
description of I. reticulata was based on a single, very small juvenile specimen of about 7 mm length
with about 28 segments (the specimen is strongly curled and the last segments are very tiny). Accord-
ing to Amoureux et al. (1978), I. reticulata can be distinguished from I. muricata by the absence of
the fringing papillae on the elytra and from I. ovata by the appearance of the elytral tubercles.
The smallest specimens of I. muricata and I. ovata examined here have been compared with
the type specimen of I. reticulata. The elytra of I. muricata specimens have fringing papillae (less
than in larger specimens, but distinct) and few, but large spinous tubercles, whereas specimens of
I. ovata have no fringing papillae and only rudimentary tubercles, as is the case in I. reticulata. The
elytron depicted by Amoureux et al. (1978) shows none of these tubercles, but large smooth rims
on the elytron surface. The smooth rims are indeed distinct in the type specimen, but not as large
as is shown in the figure in the original description. Such distinct smooth rims were also observed
in small specimens of I. ovata. I. reticulata is here regarded as a junior synonym of I. ovata.
B i o l o g y & e c o l o g y : According to Pettibone (1986 a) and the collection data listed
above, the species typically occurs in the intertidal and in shallow depths, and seems to be closely
associated to coral reefs.
(Pettibone 1986 a).
D i s t r i b u t i o n : Red Sea, Arabian Sea. This is the first record of the species from the Arabian
Sea. Elsewhere: Reported throughout tropical and subtropical regions of the Indian Ocean and the
Eastern, Central and Western Pacific (Pettibone 1986 a, Barnich et al. 2004).
Lepidastheniinae Pettibone, 1989

D i a g n o s i s : Body usually elongate, with numerous segments. Prostomium without cephalic
peaks, with three antennae. Median antenna with a large ceratophore in anterior notch of prosto-
70 T. Wehe
mium, lateral antennae with distinct ceratophores inserted terminally, with ceratophores formed
by anterior prolongations of prostomial lobes. Palps stout, conical. Two pairs of eyes usually on
posterior half of prostomium. Tentaculophores without chaetae. Buccal segment with or without a
nuchal fold. Elytra usually numerous pairs on segments 2, 4, 5, 7, alternating to 23, then variable
in arrangement (exception: Benhamipolynoe, with elytra arrangement variable posterior to segment
15). Elytra surface smooth, sometimes with scattered micropapillae; margin usually without fring-
ing papillae, sometimes with few fringing papillae on anterior elytra. Dorsal cirri with smooth
styles. Dorsal tubercles absent or inconspicuous. Notopodia small or vestigial. Neuropodia dorsally
and ventrally deeply incised, pre- and postchaetal lobes of similar length, bluntly rounded. Noto-
chaetae very few or absent. Neurochaetae few to moderate, with uni- or bidentate tips, shape vari-
able. Pygidium with a pair of anal cirri.
R e m a r k s : Neither Pettibone (1989 b) nor Hanley & Burke (1991 a) referred to cerato-
phores on the lateral antennae. For reasons explained in the remarks section under the Lepidonoti-
nae, the genera belonging to Lepidastheniinae are also considered to possess distinct ceratophores
(see Barnich & Fiege 2003, Barnich & Fiege 2004).
Key to the genera and species of Lepidastheniinae
1 Segments numerous, up to 100 or more. Elytra numerous, on segments 2, 4, 5, 7,

…, 23, 26, 29, then on every third............................................................................. 2
– Segments about 46‑49. Elytra 22 pairs on segments 2, 4, 5, 7, …, 41 and 42
Hyperhalosydna striata
2 Parapodia with large globular vesicles dorsally and ventrally. Elytra margins with
small fringing papillae Lepidasthenia vesiculosa
– Parapodia surface smooth. Elytra margins smooth Lepidasthenia nuda
Hyperhalosydna Augener, 1922

Hyperhalosydna Augener, 1922 a. — Arkiv för Zoologi 14(8): 6.
Hyperhalosydna. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 53.
Ty p e s p e c i e s : Lepidonotus striatus Kinberg, 1856.
D i a g n o s i s (emended): Body flattened, segments about 46 to 49. Styles of antennae with a
subterminal swelling and terminal filum. Palps stout, conical. Two pairs of eyes. Second (buccal)
segment without nuchal fold. Elytra 22 pairs on segments 2, 4, 5, 7, alternating to 41, 42, cover-
ing dorsum completely; surface smooth or with micropapillae, margin without fringing papillae.
Dorsal cirri with large, more or less globular cirrophore and dorsoventrally flattened style, with-
out subterminal swelling. Dorsal tubercles more or less conspicuous. Notopodia small, pin-like.
Notochaetae very few, 0‑3, very small, finely spinous. Neurochaetae stout, falcate, subdistally with
a slight swelling and several rows of spines below bidentate tips (some ventral chaetae in anterior
segments with unidentate tips).
R e m a r k s : The number of pairs of elytra has been the subject of controversy in the literature,
as discussed in Hanley & Burke (1991 a). The latter considered the number of pairs to be constant
with 22. This observation is confirmed here, since all complete specimens examined agree with this
number. The generic diagnosis is emended with respect to the dorsal cirri, which are considered to
possess styles without subterminal swellings.
Hyperhalosydna striata (Kinberg, 1856) Fig. 10 a‑g, Pl. 1 b, c

Lepidonotus striatus Kinberg, 1856. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 12(9&10): 384.
Hyperhalosydna striata. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 54‑57, fig. 17.
Ty p e l o c a l i t y : Port Jackson (Sydney), Australia, Western Pacific.
S p e c i m e n s e x a m i n e d : A r a b i a n S e a : 2 spms., South Arabian Coast, coll. “John Murray Expedition 1933‑34”,
BMNH 1937.9.2.42‑43; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑10 m, under stones and
coral rubble, coll. T. Wehe, 09 April 2000, SMF 13554; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E,
8‑10 m, under stones and coral rubble, coll. T. Wehe, 09 April 2000, SMF 13551; 1 spm., Socotra Island, Ras Qataninh Bay,
12°21.293'N/53°32.659'E, 8‑10 m, under stones and coral rubble, coll. T. Wehe, 09 April 2000, SMF 13552; 1 spm., Socotra
Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑10 m, under stones and coral rubble, coll. T. Wehe, 09 April 2000, SMF
13553; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑10 m, under stones and coral rubble, coll. T. Wehe,
09 April 2000, det. T. Wehe, 2001, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑10 m, under
stones and coral rubble, coll. T. Wehe, 09 April 2000, det. T. Wehe, 2001, NHCY; 1 spm., Socotra Island, south coast of Socotra,
Steroh, Nogid, 12°19.006'N/53°52.845'E, 13‑15 m, rocky ground, coll. M. Apel, 14 March 1999, det. T. Wehe, 2001, NHCY.
M e a s u r e m e n t s : BMNH 1937.9.2.42‑432: af (17) 9/6/4, af (20) 8/5/4. SMF 13554: cs (48) 33/4/3. SMF 13551: cs
(48) 30/3.5/2.5. SMF 13552: cs (47) 26/5/4. SMF 13553: cs (49) 37/6/5. NHCY: cs (48) 32/6/5. NHCY: cs (48) 27/3/2. NHCY:
cs (42) 12/2.7/2.5.
D i a g n o s i s : Elytra surface with five reddish-brown longitudinal stripes, submarginally with
a band of numerous microtubercles, and smooth margin.
D e s c r i p t i o n : Prostomium (Fig. 10 a) slightly wider than long. Antennae smooth, with sub-
terminal swelling and terminal filum. Styles of lateral antennae much shorter than median one. Palps
smooth, conical, long, gradually tapering. Two pairs of eyes, anterior pair dorsolaterally on more or
less widest part of prostomium, posterior pair in front of hind margin. Everted pharynx dorsally and
ventrally with two semicircles of nine, soft border papillae, and two pairs of jaws. Tentacular cirri
similar in length and shape to median antenna, dorsal ones slightly longer than ventral ones. Facial
tubercle distinct. Buccal cirri very long, inserted proximally on parapodia, shaped as tentacular cirri.
Elytra (Fig. 10 b) large, overlapping. First pair rounded to oval, following ones oval. Elytra with five
reddish-brown longitudinal stripes (more or less visible in ethanol preserved specimens) and white
spot above elytrophore; surface mostly smooth except for a band of numerous microtubercles ante-
rolaterally (Fig. 10 b, 10 c); very few additional scattered microtubercles may occur. Dorsal cirri with
large, globular to cylindrical cirrophore; style dorsoventrally flattened, smooth, without subterminal
swelling, abruptly tapering to slender terminal filum. Dorsal cirri usually erected dorsally between
elytra. Dorsal tubercles indistinct. Parapodia (Fig. 10 d, 10 e) with very small notopodia, pin-like,
with minute pre- and postchaetal lobes. Neuropodial lobes diagonally rounded. Ventral cirri short,
gradually tapering. Nephridial papillae distinct from about segments 10‑12 to 43‑45. Single noto-
chaeta usually present, minute, mostly hidden between notopodial lobes or by larger acicula, slightly
curved, with numerous rows of spines below unidentate tip. Neurochaetae (Fig. 10 f, 10 g) much
larger, numerous, slightly thickened subdistally, with several rows of spines below bidentate tips (very
rarely some ventral ones unidentate). Neurochaetae of segments two and three slightly different, with-
out a subterminal swelling and more numerous rows of spines. Anus dorsally on posterior segments.
R e m a r k s : The record of Hyperhalosydna striata from The Gulf by Wesenberg-Lund (1949)
could not be confirmed. The specimen in question is referred to Parahalosydnopsis arabica n. sp.
described herein.
B i o l o g y & e c o l o g y : According to Hanley & Burke (1991 a), the species is free-living or
associated with eunicid polychaetes, in the tubes of which it was found. Specimens from Socotra Island
where found amongst coral rubble, as were the specimens described by Hanley & Burke (1991 a).
(Hanley & Burke 1991 a).
D i s t r i b u t i o n : Arabian Sea. New record. Elsewhere: Reported throughout the Indo-West
Pacific: e.g. from Madagascar, Malay Archipelago, Philippines, Australia, Chesterfield Islands,
72 T. Wehe
Fig. 10: Hyperhalosydna striata (SMF 13553). a: anterior end; b: 9th right elytron; c: microtubercles on elytra; d: left parapodium
from segment 24, anterior view; e: same, posterior view; f: upper neurochaeta; g: lower neurochaeta.
Fairway Reefs, New Zealand, Japan (Day & Hutchings 1979, Fauvel 1953, Hanley & Burke
1991 a, Hoagland 1920, Knox 1951, Seidler 1923).
Lepidasthenia Malmgren, 1867

Lepidasthenia Malmgren, 1867. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 24(4): 139.
Lepidasthenia. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 50‑51; Barnich &
Fiege 2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 559: 88.
Bouchiria Wesenberg-Lund, 1949. — Danish scientific investigations in Iran 4: 255‑257 [new synonymy].
Ty p e s p e c i e s : Polynoe elegans Grube, 1840.
D i a g n o s i s : Body cylindrical in cross-section, vermiform, elongate, usually with numer-
ous segments (up to 100 and more). Styles of antennae smooth, without a distinct subterminal
swelling. Palps stout, conical. Two pairs of eyes. Second (buccal) segment with or without nuchal
fold, sometimes with dorsal papillae. Elytra numerous pairs on segments 2, 4, 5, 7, alternating to
23, 26, 29, continuing on every third segment to end of body; surface mostly smooth, sometimes
with scattered papillae, margin without fringing papillae. Dorsal cirri with prominent cylindri-
cal cirrophore and smooth, tapering style. Notopodia very small or vestigial. Neuropodia with or
without globular papillae dorsally or ventrally. Notochaetae absent (rarely a single one in anterior
segments). Neurochaetae slender, capillary and/or stout, falcate, with rows of spines below uni- or
bidentate tips.
R e m a r k s : The description of the genus Bouchiria was based on a single incomplete speci-
men. As mentioned by Wesenberg-Lund (1949), there are only two small tubercles in the posi-
tion where the palps normally arise, which were assumed to be rudimentary. This was followed by
Fauchald (1977), who placed the genus in the Lepidonotinae. The missing palps and the small
tubercles that are present are here regarded as representing an artificial character state. Palps are
known in all scale worm genera and it appears highly unlikely that they have been reduced in
Bouchiria. Palps missing in preserved specimens is not an unusual situation and has been observed
several times during this study.
Wesenberg-Lund (1949) originally suggested a relationship between Bouchiria and lepi-
dastheniid-like genera, e.g. Lepidasthenia or Hyperhalosydna. Nevertheless, she erected the new
genus mainly based on the missing palps and the presence of globular vesicles on the parapodia,
which were erroneously assumed to be unique amongst the Polynoidae.
If the palps are considered to be broken off, Bouchiria agrees with Lepidasthenia in all other
respects. Slender, spinous neurochaetae are found on the anterior segments of Lepidasthenia spe-
cies (e.g. L. brunnea Day, 1960), as well as globular vesicles on the parapodia (e.g. L. brunnea, L.
mossambica Day, 1962, L. vietnamica Averincev & Uschakov, 1977). Bouchiria is therefore here
synonymised with Lepidasthenia.
Lepidasthenia nuda (Grube, 1870) Fig. 11 a‑j

Polynoe nuda Grube, 1870. — Archiv für Naturgeschichte 36(1): 289‑291.
Lepidasthenia elegans. — Fauvel 1933, Mémoires de l’Institut d’Égypte 21: 41‑42 [not Grube, 1840].
Ty p e l o c a l i t y : Red Sea.
S p e c i m e n s e x a m i n e d : L e c t o t y p e h e r e d e s i g n a t e d : Polynoe nuda Quatrefages, Mer Rouge, “A1(R.) - 1868
- N°32 b”, coll. M. Botta, MNHN POLY TYPE 1434. P a r a l e c t o t y p e : Polynoe nuda Quatrefages, Mer Rouge, “A1(R.) - 1868
- N°32 a”, coll. M. Botta [second label states: syntype Polynoe nuda (Quatrefages, MS) Grube, 1870 = Lepidasthenia elegans (Gru-
be), M. Pettibone, 1975], MNHN POLY TYPE 1441. — R e d S e a : 1 spm., Gulf of Suez, coll. Hartmeyer, January 1902, det.
Michaelsen 1903, ZMH V6128; 1 spm., Gulf of Suez, coll. R.Ph. Dollfus, 25 December 1928, MNHN A360; microscope slide,
Gulf of Suez, coll. R.Ph. Dollfus, IEA R/71; probably belonging to MNHN A360. — A r a b i a n S e a : 1 spm., Socotra Island,
Dimitri, west of Rhiy di-Adho, 12°37.349'N/54°17.589 E, 25‑30 m, under dead corals & soft corals, coll. M. Apel, 3 April 1999,
SMF 13640.
M e a s u r e m e n t s : Lectotype MNHN POLY TYPE 1434: cs (85) 88/8/6 [af (36) 33 + mf (13) 18 + pf (36) 37]. Paralecto-
type MNHN POLY TYPE 1441: af (45) 30/7/5. ZMH V6128: cs (81) 58/9/7. MNHN A360: cs (72) 48/7/5 [af (29) 18 + pf (43)
30]. SMF 13640: cs (84) 47/8/6 [af (42) 25 + mf (12) 8 + mf (5) 4 + pf (25) 10].
D i a g n o s i s : Elytra smooth. No vesicles on parapodia. No giant neurochaetae. Neurochaetae
all bidentate, only the large uppermost one being unidentate. Neurochaetae on anterior segments
very slender, as are most upper and lower ones in following parapodia.
74 T. Wehe
R e d e s c r i p t i o n based on lectotype designated here: Prostomium (Fig. 11 a) about as wide

as long. Antennae of paralectotype (left lateral and median antenna broken off on lectotype)
smooth, with very slight subterminal swelling and distinct terminal filum; styles of lateral antennae
shorter than median one. Palps stout, smooth, conical, long, abruptly tapering to short filiform tip
(palps on paralectotype smaller and shorter). Two pairs of eyes (hardly visible on paralectotype),
anterior pair dorsolaterally on more or less widest part of prostomium, posterior pair in front of
hind margin. Tentacular cirri similar in length and shape to median antenna on paralectotype,
dorsal ones slightly longer than ventral ones (right dorsal one on lectotype comparatively short,
ventral one broken off ). Facial tubercle not visible on lectotype, pharynx slightly everted (facial
tubercle on paralectotype small). Buccal cirri very long, inserted proximally on parapodia, shaped
as tentacular cirri. Elytra (Fig. 11 b) not covering dorsum, large and oval on anterior segments of
paralectotype (missing on lectotype), gradually becoming smaller and more rounded along body.
Elytra smooth, translucent, with leaf-like venation. Dorsal cirri with large, but short cylindri-
cal cirrophore and stout, smooth style tapering abruptly to terminal filum, without subterminal
swelling. Styles gradually decreasing in length and size along body, on anterior segments extending
far beyond neuropodia, posteriorly hardly reaching lateral borders of the same. Dorsal tubercles
absent. Parapodia (Fig. 11 c, 11 d, 11 e) with very small, conical notopodia in anterior segments,
becoming vestigial from about segments 20 to 30 and absent thereafter. Neuropodial lobes bluntly
rounded, of similar length. Ventral cirri smooth, short, gradually tapering. Nephridial papillae
distinctly present from about segment 27 to 30. Notochaetae absent. Neurochaetae falcate, vary-
ing along body; slender on numerous anterior segments, with a slight subterminal swelling only,
numerous spines and bidentate tips (Fig. 11 f ); neurochaetae of following parapodia with the same
arrangement of chaetae, but gradually decreasing in size towards posterior body: uppermost neu-
rochaetae few, very slender, shaped like those of anterior segments (Fig. 11 g); first neurochaetae
below slender ones largest, with subterminal swelling and few spines below blunt, unidentate tip
(Fig. 11 h); lower accompanying large neurochaetae gradually decreasing in size ventrally, with
subterminal swelling and few spines below bidentate tips (Fig. 11 i); ventralmost neurochaetae
again very slender, short, with minute bidentate tips (Fig. 11 j). No single giant chaeta, which ex-
ceeds all other chaetae in size, present.
R e m a r k s : According to Grube (1870), a re-arrangement of the polychaete collection in the
MNHN was made, based on the voluminous work by Quatrefages (1866). However, as pointed
out by Grube (1870), there is no description of Polynoe nuda by Quatrefages (as indicated on the
original labels) in the monograph by Quatrefages (1866). Even though Grube (1870) credited the
authorship to Quatrefages, which was followed by Hartman (1959), it was Grube who described
the species for the first time and the authorship belongs to him. Following Grube (1870), there is
no doubt that the lectotype designated here and the paralectotype represent the specimens exam-
ined by Grube for his description of P. nuda. Unfortunately, the description provided by Grube,
though very long, is not detailed enough to distinguish the species from others within the genus.
Since Grube did not designate a holotype, a lectotype is designated here within the framework of a
redescription. The longer and, in terms of parapodial and chaetael characters, the better preserved
specimen was chosen as lectotype. Species of Lepidasthenia are difficult to distinguish, mainly by
the type and arrangement of the neurochaetae along the body, and the genus is in urgent need of
revision. However, based on a comparison of the characters examined on the specimens and those
described in the literature, it seems that there is no other species that agrees with L. nuda. A syn-
onymy with L. elegans, the type specimens of which were examined, is ruled out since there are no
giant chaetae present in L. nuda.
Fig. 11: Lepidasthenia nuda (lectotype, MNHN A28: c-m; paralectotype, MNHN A28: a, b). a: anterior end; b: right elytron from
anterior segment; c: right parapodium from segment 14, anterior view; d: left parapodium from segment 29, anterior view; e: left
parapodium from segment 69, posterior view; f: neurochaeta from d; g: uppermost slender neurochaeta from e; h: uppermost large
neurochaeta from same; i: middle neurochaeta from same; j: lower neurochaeta from same.
D e p t h r a n g e : So far recorded between 25 and 30 m; unknown elsewhere.

D i s t r i b u t i o n : Red Sea, Arabian Sea: Socotra Archipelago. This is the first record of L.
nuda from the Arabian Sea.
76 T. Wehe
Lepidasthenia vesiculosa (Wesenberg-Lund, 1949) n. comb. Fig. 12 a‑e

Bouchiria vesiculosa Wesenberg-Lund, 1949. — Danish scientific investigations in Iran 4: 255‑257, figs 1, 2.
Ty p e l o c a l i t y : Būshehr (= Bushire), The Gulf.
S p e c i m e n s e x a m i n e d : H o l o t y p e : The Gulf, Iran, Bushire, 41 miles S, ¾ E of B ū shire light-buoy,
28°17'14''N/50°49'E, Petersen-grab, 43 m, very fine light mud with H2S, coll. G. Thorson, 26 March 1937, ZMUC POL-1368.
M e a s u r e m e n t s : af (17) 7/5/3.5.
D i a g n o s i s : Elytra with small fringing papillae. Cirrophores of dorsal cirri, dorsal and ven-
tral surface of parapodia with bulbous vesicles. Neurochaetae of anterior segments slender with
capillary or blunt tips.
D e s c r i p t i o n : Prostomium (Fig. 12 a) slightly wider than long. Style of median antenna bro-
ken off. Styles of lateral antennae, slender, smooth, without a subterminal swelling, with a terminal
filum. An indistinct nuchal fold seems to be present. Palps broken off, two inconspicuous tubercles
instead. Two pairs of rounded eyes on posterior half of prostomium, anterior pair dorsolaterally
slightly in front of widest part of prostomium, posterior pair dorsally in front of hind margin. Ten-
tacular cirri very long, smooth, without subterminal swelling, gradually tapering. Facial tubercle dis-
tinct. Buccal cirri long, inserted proximally on parapodia, shaped as tentacular cirri. Elytra (Fig. 12 a)
small, leaving middorsum uncovered, more-or-less oval, smooth, with very small fringing papillae on
lateral borders. Dorsal cirri with a small, cylindrical cirrophore and smooth style gradually tapering to
a terminal filum; dorsal cirrus on segment three about twice as long as parapodium, length decreasing
on following segments, not reaching lateral border of parapodia on segment 16. Dorsal tubercles ab-
sent. Parapodia (Fig. 12 b) with very small, conical to vestigial notopodia. Neuropodial lobes bluntly
rounded, prechaetal one longer than postchaetal. Parapodia with few large, stalked, globular vesicles
(Fig. 12 c) on dorsal and ventral surface, and along cirrophore of dorsal cirri. Ventral cirri short, grad-
ually tapering. Notochaetae absent. Neurochaetae (Fig. 12 d, 12 e) slender, with few spines below
unidentate tips; upper ones more bent with capillary tips, lower ones straight with blunt tips.
R e m a r k s : The species was described from a very small anterior fragment, which is in bad
condition, and so important neurochaetal characters of the more posterior segments are unknown.
Nevertheless, some of the characters exhibited come very close to other species described in Lepi-
dasthenia. Parapodial vesicles for example are present in L. brunnea Day, 1960, L. mossambica Day,
1962, L. vietnamica Averincev & Uschakov, 1977, and slender capillary chaetae are known from
e.g. L. brunnea (see also Day 1967, Barnich & Fiege 2003). The latter species may also have papil-
lae on the margins of the elytra as figured (but described as smooth) in Barnich & Fiege (2003).
A synonymy with L. brunnea seems possible. However, until there is a revision of the relevant type
specimens, it cannot be decided whether both species are valid. Until then, L. vesiculosa is regarded
as a separate species.
D e p t h r a n g e : So far recorded from 43 m.
D i s t r i b u t i o n : The Gulf.
Lepidonotinae Willey, 1902

D i a g n o s i s : Body short to long, with a fixed number of segments (26) or numerous (up to
70 and more). Prostomium without cephalic peaks, with three antennae, usually two pairs of eyes
on posterior half of prostomium. Median antenna with large ceratophore in anterior notch of pros-
tomium. Lateral antennae inserted terminally or subterminally on prostomium, with ceratophores
formed by anterior prolongations of prostomial lobes. Palps stout, conical. Usually with two pairs
of eyes, arrangement variable. Tentaculophores with or without chaetae. Second (buccal) segment
with or without nuchal fold. Elytra with fixed number of pairs (e.g. 12, 15, 17, 18, 20) or numer-
Fig. 12: Lepidasthenia vesiculosa (holotype, ZMUC POL-1368). a: anterior end; b: right parapodium from posterior part of frag-
ment, posterior view; c: ventral parapodial vesicle; d: upper neurochaetae; e: lower neurochaeta.
ous on segments 2, 4, 5, 7, alternating to 23, then variable in arrangement. Elytra surface smooth
or variously ornamented with tubercles, papillae, ridges, etc., margin with or without fringing
papillae. Dorsal cirri usually with smooth styles. Dorsal tubercles more or less conspicuous. Noto-
podia moderate to small. Neuropodia dorsally and ventrally not deeply incised; pre- and postcha-
etal lobes of about similar length, with or without acicular process. Notochaetae few to numerous,
rarely absent; shape variable. Neurochaetae moderate to numerous, with uni- or bidentate tips,
shape variable. Pygidium with a pair of anal cirri.
R e m a r k s : According to the diagnosis given by Hanley & Burke (1991 a), the antennae are
always smooth in the Lepidonotinae. There are, however, a few exceptions as papillae are present
on the median antenna of Halosydnopsis and Parahalosydnopsis. Both of these genera, together with
Lepidonopsis and Hermenia, are unique amongst the Lepidonotinae for another character, the pos-
session of papillae on the distal edges of the neuropodial lobes.
Hanley & Burke (1991 a) stated that the lateral antennae do not posses distinct ceratophores.
This view is not accepted here. The styles of the lateral antennae do not emerge continuously from
the prostomium, but are clearly separated from structures similar to the median ceratophore, even
though these structures themselves are prolongations of the prostomial lobes. As there is obviously
a separation into a basal carrier structure and a distal style, and since also the cirrophores of the
dorsal cirri are continuous with the parapodia and are unquestionably carrier structures, members
78 T. Wehe
of the Lepidonotinae are considered to possess lateral antennae with distinct ceratophores (see also
Barnich & Fiege 2003).
Some of the genera currently considered as belonging to the Lepidonotinae possess lateral an-
tennae, which are inserted subterminally (see Fig. 2 d), e.g. Alentia or Heteralentia. There is some
confusion between the type of arrangement of the lateral antennae in Heteralentia mentioned by
Hanley & Burke (1991 a) in the description (“terminally”) and in the remarks section (“subter-
minally”). The present author agrees with these authors and Pettibone (pers. comm. in Hanley &
Burke 1991 a) that these two genera together with Paralentia Uschakov, 1982 and Ophthalmonoe
Petersen & Britayev, 1997 may constitute a distinct subfamily of their own. However, the erection
of a new subfamily is postponed until more specimens have been studied.
Key to the genera of Lepidonotinae
1 Body always with 26 segments................................................................................... 2

– Body always with more than 26 segments.................................................................. 4
2 Only first or first to third pairs of elytra large and distinct. Neurochaetae very stout
with only one or two spines subdistally Hermenia
– All elytra distinct. Neurochaetae with several rows of spines subdistally...................... 3
3 All notochaetae of the same type, spinous Lepidonotus
– Notochaetae of two types: outer palisade of stout, spinous ones, and inner bundle of
slender, lancet-shaped ones Thormora
4 Segments up to 75. Antennal styles and parapodial lobes smooth. Elytra 18 pairs or
more on segments 2, 4, 5, 7, …, 23, 26, 29, 32, 35, 38, then variable Heteralentia
– Segments up to 48. Style of median antenna and neuropodial lobes with papillae
distally. Elytra numerous pairs on segments 2, 4, 5, 7, …, 25, 28, 30, …, 42, 43, 45,
47, 48, 50, alternating to end of body Parahalosydnopsis
Hermenia Grube, 1856

Hermenia Grube, 1856. — Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjöbenhavn for Aaret 1856: 44.
Hermenia. — Pettibone 1975, Proceedings of the Biological Society of Washington 88(22): 234; Hanley & Burke 1991 a, Mé-
moires du Muséum National d’Histoire Naturelle (A) 151: 61‑62.
Ty p e s p e c i e s : Hermenia verruculosa Grube, 1856.
D i a g n o s i s : Body with 26 segments, short, dorsoventrally flattened, subrectangular in
cross-section, integument tuberculate and/or papillate. Ceratophores of lateral antennae inserted
terminally. Styles of antennae smooth, with subterminal swelling. Palps stout, conical, gradually ta-
pering. Two pairs of eyes present. Tentaculophores with chaetae. Second (buccal) segment without
nuchal fold. Elytra 12 pairs on segments 2, 4, 5, 7, alternating to 23, firmly attached to indistinct
elytrophores; first to third or first pair only large and distinct, remaining ones very small, merely
covering elytrophore. Surface and margins of elytra with papillae and very large spinous tubercles.
Dorsal cirri very small, with cylindrical cirrophore and smooth style with subterminal swelling.
Notopodia very small, almost inconspicuous. Neuropodia large, truncate, pre- and postchaetal
lobe of similar length with papillae distally. Notochaetae few slender, finely spinous, or absent.
Neurochaetae falcate, very stout, subdistally with 1 or 2 large teeth and slightly bent, blunt tip.
Key to the species of Hermenia
1 First three pairs of elytra large. Neurochaeta with two subterminal spines
H. acantholepis
– Only first pair of elytra large. Neurochaetae with a single prominent subterminal
spine H. verruculosa
Hermenia acantholepis (Grube, 1876) Fig. 13 a‑c

Polynoe acantholepis Grube, 1876. — Jahres-Bericht der Schlesischen Gesellschaft für vaterländische Cultur 53: 61.
Hermenia acantholepis. — Fauvel 1933, Mémoires de l’Institut d’Égypte 21: 41; Pettibone 1975, Proceedings of the Biological
Society of Washington 88(22): 239‑241, figs 3, 4; Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire
Naturelle (A) 151: 62‑64, fig. 19.
Ty p e l o c a l i t y : Upolu (Samoa), Philippines, West Pacific.
S p e c i m e n s e x a m i n e d : S y n t y p e : no type location on label, as Lepidonotus acantholepis, Coll. Grube (single elytron
only, probably from syntype according to Hartwich 1993), ZMB 1078. — R e d S e a : 1 spm., Gulf of Suez, from vermetid bloc,
coll. R.Ph. Dollfus, 30 December 1928, MNHN A894; 1 spm., Gulf of Suez, coll. R.Ph. Dollfus, 1928, MNHN A403; 1 spm.,
Gulf of Suez, coll. R.Ph. Dollfus, 30 December 1928, MNHN A360.
M e a s u r e m e n t s : MNHN A894: cs (26) 18/7/5. MNHN A403: cs (26) 13/5/4. MNHN A360: cs (26) 14/6/4.5.
D i a g n o s i s : First three pairs of elytra large to moderate in size, following ones small, incon-
spicuous. Neurochaetae subdistally with two distinct spines.
D e s c r i p t i o n : Body stout, subrectangular, flattened ventrally, arched dorsally, thickly ver-
rucose. Prostomium rounded, withdrawn into anterior segments. Median antenna with large, cy-
lindrical ceratophore in anterior notch of prostomium, style long, smooth, with subterminal swell-
ing and terminal filum. Lateral antennae similar, but shorter. Palps stout, conical, long, smooth,
gradually tapering. Eyes not visible. Tentaculophores with two chaetae each, tentacular cirri shaped
as lateral antennae, but longer, dorsal ones slightly longer than ventral ones. Facial tubercle dis-
tinct. Second segment with buccal cirri longer than following ventral cirri, shaped as antennae and
tentacular cirri. First three pairs of elytra distinct, large, gradually decreasing in size, only first pair
touching medially; first pair rounded (Fig. 13 a), second one reniform (Fig. 13 b), third one oval.
Following ones much smaller, only covering elytrophores, rounded. Surface and margin covered
with very large, globular, regularly ornamented macrotubercles with scattered papillae in between.
Margin with digitiform fringing papillae. Dorsal cirri with comparatively small, cylindrical cir-
rophore and short, slender styles with subterminal swelling and terminal filum. Dorsal tubercles
inconspicuous. Parapodia with very small to vestigial notopodia. Neuropodia large, truncate
and papillate distally, with subtriangular, prechaetal, acicular lobe and slightly shorter, rounded
postchaetal one. Ventral cirri short, gradually tapering. Nephridial papillae not observed. Notocha-
etae very few, sometimes broken off, very slender, short to long, thread-like, finely spinous. Neu-
rochaetae (Fig. 13 c) numerous, much stouter, subdistally with two distinct spines below bluntly
rounded, unidentate tips. Neurochaetae of segments 2 to 4 more slender, shorter, with several rows
of spines.
B i o l o g y & e c o l o g y : According to Hanley & Burke (1991 a), the species is found
amongst living corals and coral rubble.
D e p t h r a n g e : Unknown; elsewhere reported between 16 and 95 m (Hanley & Burke
1991 a).
D i s t r i b u t i o n : Red Sea: Gulf of Suez. Elsewhere: All over Indo-West Pacific: Madagascar,
Indonesia, Annam, Sri Lanka, Maldives, Philippines, Australia, Chesterfield Islands, Fairway Reefs,
New Caledonia, Samoa (Fauvel 1953, Pettibone 1975, Hanley & Burke 1991 a).
80 T. Wehe
Fig. 13: Hermenia acantholepis (MNHN A894). a: 1st right elytron; b: 2nd right elytron; c: neurochaeta.
Hermenia verruculosa Grube, 1856 Fig. 14 a, b; Pl. 1 d

Hermenia verruculosa Grube, 1856. — Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjöbenhavn for Aaret 1856: 44.
Hermenia verruculosa. — Pettibone 1975, Proceedings of the Biological Society of Washington 88(22): 235‑238, figs 1, 2.
Ty p e l o c a l i t y : St. John, Virgin Islands, West Indies, Caribbean Sea.
S p e c i m e n s e x a m i n e d : A r a b i a n S e a : 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E,
8‑10 m, under stones and coral rubble, coll. T. Wehe, 9 April 2000, SMF 13608.
M e a s u r e m e n t s : cs (26) 18/8/6.
D i a g n o s i s : First pair of elytra large, following elytra small, inconspicuous. Neurochaetae
subdistally with a single prominent spine.
D e s c r i p t i o n : Body stout, subrectangular, flattened ventrally, arched dorsally, thickly ver-
rucose. The specimen exhibits a distinct colouration pattern: dorsal tubercles on elytrigerous seg-
ments brownish, giving the specimen a transversely striated appearance; with two smaller, much
darker transverse stripes on the joint between two segments, also by coloured tubercles; the small
elytra appearing as dark dots; antennae, tentacular- and buccal cirri with three brownish bands,
one basally, one subterminally below swelling and another one distally; dorsal cirri with only two
bands, above and below subterminal swelling. Prostomium withdrawn into anterior segments.
Median antenna with large, cylindrical ceratophore in anterior notch of prostomium, style broken
off. Styles of lateral antennae smooth, with subterminal swelling and terminal filum. Palps stout,
conical, long, gradually tapering, with minute papillae. Eyes hardly visible because of withdrawn
prostomium. Tentaculophores with two chaetae each, tentacular cirri shaped as lateral antennae,
but longer. Facial tubercle distinct. Second segment with buccal cirri longer and inserted more
proximally than following ventral cirri, shaped as antennae and tentacular cirri. First pair of elytra
large (Fig. 14 a), distinct, following elytra very small, inconspicuous, partially embedded in in-
Fig. 14: Hermenia

verruculosa (MNHN
A894). a: 1st left
elytron; b: neurochaeta.
tegument. All elytra rounded to oval. Surface with globular micropapillae and large, ornamented
macrotubercles; margin with short fringing papillae. Dorsal cirri very small in comparison to
parapodium, with cylindrical cirrophore and a short, slender style with subterminal swelling and
terminal filum. No dorsal tubercles. Parapodia with small to vestigial notopodia. Neuropodia large,
truncate and papillate distally, with a subtriangular, prechaetal, acicular lobe and slightly shorter,
rounded postchaetal one. Ventral cirri short, gradually tapering. Nephridial papillae from about
segment 12 onwards. Notochaetae absent or only one slender, short, thread-like, finely spinous
chaeta present. Neurochaetae (Fig. 14 b) much stouter, subdistally with a single prominent spine
below bluntly rounded, unidentate tips. Neurochaetae of segments 2 to 4 slightly more slender,
shorter, with few more spines subdistally.
R e m a r k s : The specimen fits very well with the description given by Pettibone (1975).
Minor differences are the presence of minute papillae on the palps and the absence of such on the
cerato- and cirrophores.
B i o l o g y & e c o l o g y : According to Pettibone (1975), the species was observed living in
crevices of coral rock and was also commensal with ophiuroids. It has a similar coloration pattern
and arrangement of tubercles as the disc of the host ophiuroid, which provides good camouflage.
The specimen examined was sorted from coral rubble.
(Pettibone 1975).
D i s t r i b u t i o n : Arabian Sea: Socotra Archipelago. First record outside the tropical Western
Atlantic. Elsewhere: Gulf of Mexico, Caribbean Sea (Pettibone, 1975).
Heteralentia Hanley & Burke, 1991

Heteralentia Hanley & Burke, 1991 a. — Mémoires du Muséum National d’Histoire Naturelle (A) 151: 58.
Ty p e s p e c i e s : Polynoe ptycholepis Grube, 1878.
D i a g n o s i s (emended): Body dorsoventrally flattened, elongate, up to 48 segments, tapering
posteriorly. Ceratophores of lateral antennae inserted subterminally. Styles of antennae smooth.
Palps stout, conical, gradually tapering. Two pairs of very large eyes present. Tentaculophores
without chaetae. Buccal segment with large nuchal fold covering posterior part of prostomium.
Elytra 18 or more pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, 35, 38, then variable
in arrangement. Elytra surface with scattered microtubercles, margin without fringing papillae.
82 T. Wehe
Dorsal cirri with smooth, long, tapering styles. Notopodia small, with very long acicular process.
Neuropodia with longer, prechaetal, acicular lobe with acicular process, and shorter postchaetal
lobe. Notochaetae about as thick as neurochaetae, with blunt tips. Neurochaetae more numerous,
subdistally enlarged, faintly spinous, with bidentate tips.
R e m a r k s : The generic description provided by Hanley & Burke (1991 a) was emended
as regards the segment number and the arrangement of the lateral antennae. In their diagnosis
Hanley & Burke state that the lateral antennae are inserted “terminally...slightly ventral to median
antenna”. In the remarks section, however, they describe the lateral antennae as inserted “subter-
minally”, contrary to the terminally inserted antennae in the genus Alentia for example. Specimens
examined during this study clearly showed lateral antennae inserted subterminally (see the critical
discussion of the characters above), in Heteralentia as well as in Alentia. The latter genus is not rep-
resented in the area studied, and so additional material from the Atlantic Ocean (France, Roscoff,
SMF 13200) and descriptions and figures in the literature (Chambers & Muir 1997, Barnich
& Fiege 2003) were compared. In the arrangement of the lateral antennae and in the presence
of prominent acicular processes, those genera clearly differ from other lepidonotid genera such
as Lepidonotus, Hermenia or Thormora. The present author agrees with Pettibone (pers comm.
in Hanley & Burke 1991 a) that at least Alentia and Heteralentia should be referred to a separate
subfamily.
Furthermore, in the opinion of the present author, the status of Heteralentia needs to be con-
firmed. In the remarks section of the generic diagnosis, Hanley & Burke (1991 a) discuss only the
similarities and differences between Heteralentia and the genera Alentia and Paralentia Uschakov,
1982. Heteralentia is monospecific, and only represented by H. ptycholepis. Originally described as
Polynoe ptycholepis by Grube (1878), this species was referred to Allmaniella McIntosh, 1885 by
Horst (1917), which was followed e.g. in later works by Fauvel (1932, 1953) and Hartman
(1959). Although referring Allmaniella ptycholepis to Heteralentia and synonymising a further spe-
cies described as Allmaniella nuchalis by Hartman (1974 a), no explanation was given as to why
the genus Allmaniella was not accepted. As already pointed out by Horst (1917), there is no spe-
cial diagnosis given by McIntosh (1885) for Allmaniella. Even though the description may not be
very detailed, the characters described by the latter for Allmaniella agree very well with those for
Heteralentia. According to the diagnosis given by Hanley & Burke with 18 or more pairs of elytra,
the records by Horst (1917) were included in the list of synonyms although he reported only
15 to 17 pairs. In order to clarify these anomolies, a revision of the specimens in question is badly
needed. For the time being, Heteralentia is accepted here since the specimens examined agree with
the more detailed description provided by Hanley & Burke (1991 a).
Heteralentia ptycholepis (Grube, 1878) Fig. 15 a‑h

Polynoe ptycholepis Grube, 1878. — Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8): 39‑40, pl.
2: figs 6, 6 a.
Allmaniella ptycholepis. — Horst 1917, Siboga-Expeditie 24b: 79‑80, pl. 18: figs 6‑9.
? Allmaniella nuchalis. — Hartman 1974 a, Journal of the Marine Biological Association of India 16(1): 201‑203, fig. 1.
Heteralentia ptycholepis. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 58‑61, fig.
18; Barnich et al. 2004, Species Diversity 9: 293‑294.
Ty p e l o c a l i t y : Lapinig (= Lapinin) Island, Philippines, West Pacific.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Polynoe ptycholepis, Lapinig Island, Philippines, coll. Semper, ZMB 1153.
H o l o t y p e : Allmaniella nuchalis, Gulf of Oman, 23°35'N/58°49'E, 121‑124 m, rock dredge, International Indian Ocean Expe-
dition, coll. “Anton Bruun”, 3 December 1963 (station data according to Griffin 1974), LACM-AHF POLY 1193. — A r a b i a n
S e a : 1 spm., Socotra Archipelago, rocks of Kal Farun, 12°26.049'N/52°08.070'E, 15 m, coll. M. Apel, 3 April 2000, SMF 13547;
1 spm., Socotra Archipelago, Abd al-Kuri, Ras Anjara Bay, 12°09.981'N/52°22.282'E, amongst coral rubble, 7‑9 m, coll. T. Wehe,
6 April 2000, NHCY.
M e a s u r e m e n t s : Holotype ZMB 1153 in extremely bad condition and could not be measured. Holotype LACM-AHF
POLY 1193: af (18) 8/4.5/3.5 + mf (4) 3. SMF 13547: cs (48) 20/7/5. NHCY: cs (46) 18/5/3.5 [af (16) + mf (6) + pf (24)].
D i a g n o s i s : Microtubercles on elytra with three blunt tips. Notochaetae faintly spinous with
blunt tips. Neurochaetae with faint spines on convex and concave edge, upper ones notched, lower
ones distinctly bidentate.
D e s c r i p t i o n : Body flattened dorsoventrally, elongate, gradually tapering posteriorly. Prosto-
mium (Fig. 15 a) oval to hexagonal, wider than long. Ceratophore of median antenna small, cylindri-
cal, style long, smooth, slender, with slight subterminal swelling below terminal filum. Ceratophores
of lateral antennae similar to median, styles slightly shorter. Palps very long, gradually tapering (cor-
rugated appearance (Fig. 15 a) in SMF 13547 due to preservation). Two pairs of very large rounded
eyes, anterior pair on widest part of prostomium, posterior pair in front of hind margin. Tentaculo-
phores with short acicular process and tentacular cirri similar to median antenna. Facial tubercle small.
Second (buccal) segment with a large, subtriangular to rounded nuchal fold, covering posterior part of
prostomium. Buccal cirri shaped as tentacular cirri, inserted more proximally than following ventral
cirri. Elytra (Fig. 15 b) 20 or 21 pairs, posterior to segment 38 on 40, 43, 44, (45). Elytra large, oval
to rounded, thin, translucent, margin often folded frill-like; surface on anterior third with numerous
microtubercles with three blunt tips (Fig. 15 c); margins without fringing papillae. Dorsal cirri with
large, cylindrical cirrophore and long, smooth style without distinct subterminal swelling below ter-
minal filum. Dorsal tubercles inconspicuous. Parapodia (Fig. 15 d, 15 e) with very small notopodia,
with a very long and slender, acicular process. Neuropodia with a slightly longer prechaetal, acicular
lobe terminating in a slender, triangular acicular process and a shorter, bluntly rounded, postchaetal
one. Ventral cirri short, conical, gradually tapering. Nephridial papillae distinct from about segment
8 to 11. Notochaetae (Fig. 15 f) moderate in number, slightly curved, faintly spinous, with blunt, un-
identate tips. Neurochaetae (Fig. 15 g, 15 h) long, slightly swollen subdistally with several rows of fine
spines; subdistal swelling more prominent in lower neurochaetae; tips of upper neurochaetae blunt
and slightly notched, middle and lower ones distinctly bidentate, with prominent secondary tooth.
R e m a r k s : The neurochaetae and elytra tubercle present on the specimens from the Socotra
Archipelago differ slightly from those figured by Hanley & Burke (1991 a), in that the secondary
tooth of the middle and lower neurochaetae is more prominent and the tips of the elytra tubercle
are more broad and rectangular, as figured by Grube (1878). The holotype of Allmaniella nuchalis
agrees very well with these specimens and the descriptions provided by Hanley & Burke (1991 a)
and Horst (1917), respectively. The neuropodial acicular process is slightly longer than the figures
by Hartman (1974 a) and Hanley & Burke (1991 a). All these slight differences are considered to
be variations of the same species. However, since the elytra of Allmaniella nuchalis are missing, the
specimen is only tentatively referred to H. ptycholepis.
D e p t h r a n g e : So far recorded between 7 and 15 m; elsewhere possibly down to 124 m
(Hartman 1974 a).
D i s t r i b u t i o n : Arabian Sea: Socotra Archipelago, possibly Gulf of Oman (see remarks on
Allmaniella nuchalis above). This species is reported for the first time from the Arabian Sea. Else-
where: Indo-West Pacific: Mauritius, Hong Kong, South China Sea, Papua New Guinea, Chester-
field Islands, Samoa (Hanley & Burke 1991 a, Barnich et al. 2004).
Lepidonotus Leach, 1816

Lepidonotus Leach, 1816. — Supplement to the 4 th, 5 th, and 6 th editions of the Encyclopaedia Britannica 1(2): 451.
Lepidonotus. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 64‑65; Barnich &
Fiege 2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 559: 82.
84 T. Wehe
Fig. 15: Heteralentia ptycholepis (SMF 13547). a: anterior end; b: right elytron from midbody; c: microtubercles on elytra; d: right
parapodium of segment 21, anterior view; e: same, posterior view; f: notochaeta; g: upper neurochaeta; h: lower neurochaeta.
Ty p e s p e c i e s : Aphrodita clava Montagu, 1808.

D i a g n o s i s : Body short with 26 segments, dorsoventrally flattened, subrectangular in cross-
section. Ceratophores of lateral antennae inserted terminally. Styles of antennae smooth, with or
without subterminal swelling. Palps stout, conical, gradually tapering. Two pairs of eyes usually
present. Tentaculophores usually with chaetae. Second or buccal segment with or without nuchal
fold or nuchal nods. Elytra 12 pairs on segments 2, 4, 5, 7, alternating to 23, usually large, cov-
ering dorsum completely; surface variably ornamented with tubercles, papillae, vesicles, spines,
ridges, mounds, etc., or rarely smooth; margin with or without fringing papillae. Dorsal cirri usu-
ally with prominent bulbous or cylindrical cirrophore and smooth styles, with or without subdistal
swelling. Notopodia small or inconspicuous. Neuropodia with pre- and postchaetal lobes of simi-
lar length. Notochaetae rarely absent, usually with numerous rows of spines and unidentate tips.
Neurochaetae stouter than notochaetae, falcate, with few to many rows of spines subdistally below
uni- or bidentate tips.
R e m a r k s : Seidler (1923) prepared a very extensive review of the Lepidonotinae, providing
descriptions and keys to more than 50 species of Lepidonotus. Unfortunately, however, his work
has some disadvantages. There are almost no illustrations to supplement the descriptions, nor is
it clear from the text which specimens or types were examined. Obviously some descriptions were
prepared by him, while others (indicated by the name of the author of the original description in
brackets) are just translations, most probably indicating that no specimens of these species were
examined by Seidler himself. Furthermore, the systematics proposed by Seidler (1923) need to
be re-examined, since some mistakes have been found and reported here (see L. cristatus and L.
impatiens). Nevertheless, Seidler’s work is invaluable in providing base-line data and access to the
literature on lepidonotid genera.
Key to the species of Lepidonotus
1 Elytra with fringing papillae....................................................................................... 2

– Elytra without fringing papillae.................................................................................. 4
2 Elytra surface with numerous long, slender papillae L. natalensis
– Elytra surface without long, slender papillae............................................................... 3
3 Anterior pair of eyes dorsolaterally on widest part of prostomium, posterior pair in
front of hind margin. Elytra with warty and carinate tubercles. Neurochaetae biden-
tate L. carinulatus
– Both pairs of eyes dorsolaterally on prostomium. Elytra with smooth tubercles. Neu-
rochaetae unidentate L. tenuisetosus
4 Elytra with two keels on posterior half L. glaucus
– Elytra without keels................................................................................................... 5
5 Elytra with sac-like protuberances, conical macrotubercles and numerous microtu-
bercles with four spines L. cristatus
– Elytra without sac-like protuberances, without fourpointed microtubercles................ 6
6 Elytra with nail-like microtubercles............................................................................ 7
– Microtubercles not nail-like....................................................................................... 8
7 Nail-like tubercles numerous and very dense on elytra surface. Uni- to trifid microtu-
bercles in between. Warty macrotubercles absent. L. impatiens
– With large warty macrotubercles and only a few scattered nail-like tubercles. Uni- to
multifid microtubercles in between L. polae
8 Elytra almost smooth, with minute microtubercles. Elytra surface with clusters of
unpigmented cells L. purpureus
– Elytra with ornamented macro- and microtubercles, without clusters of unpigmented
cells L. clava
86 T. Wehe
Lepidonotus carinulatus Grube, 1869 Fig. 16 a‑o

Polynoe (Lepidonotus) carinulata Grube, 1869, Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin
aus dem Jahre 1869: 488; Grube 1878, Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8):
26‑27, pl. 3: figs 2‑2 b.
Lepidonotus carinulatus. — Willey 1905, Ceylon Pearl Oyster Fisheries Supplementary Reports 30: 248‑249, pl. 1: figs 7‑11;
Fauvel 1911, Archives de Zoologie Expérimentale et Générale, Cinquième Série 4(11): 367, fig. 1; Fauvel 1918, Bulletin
du Muséum d’Histoire Naturelle 24(5): 330; Fauvel 1919, Archives de Zoologie Expérimentale et Générale 58: 330; Auge-
ner 1922 b, Archiv für Naturgeschichte, Abteilung A 88(7): 8‑11, figs 3‑3 b; Seidler 1923, Archiv für Naturgeschichte 89
(Abt. A/11): 72‑73; Fauvel 1933, Mémoires de l’Institut d’Égypte 21: 37; Monro 1939 b, Annals and Magazine of Natu-
ral History 11(4): 169; Wesenberg-Lund 1949, Danish scientific investigations in Iran 4: 252‑253; Fauvel 1955, Annales
de l’Institute Océanographique Paris 30: 103; Mohammad 1971, Journal of Zoology 163: 287‑288; Mohammad 1972 b,
Hydrobiologia 40(4): 554; Day 1975 Records of the Western Australian Museum 3(3): 179‑180, figs 2 g‑k; Amoureux et
al. 1978, Israel Journal of Zoology 27: 63; Hanley & Burke 1990, Proceedings of the Third International Marine Biological
Workshop: The Marine Flora and Fauna of Albany, Western Australia 1: 216‑218, fig. 6, Hanley & Burke 1991 a, Mémoi-
res du Muséum National d’Histoire Naturelle (A) 151: 65‑68, fig. 20; Imajima 1997, National Science Museum Monographs
13: 95‑98, figs 45‑46; Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 60‑64, figs 7, 8; Barnich
et al. 2004, Species Diversity 9: 297‑298.
Lepidonotus tenuisetosus. — Mohammad 1971, Journal of Zoology 163: 288 [partim, see L. tenuisetosus; not Gravier, 1902].
S p e c i m e n s e x a m i n e d : S y n t y p e : Lepidonotus carinulatus, Red Sea, Coll. E. Grube, ZMB 1072. 3 Syntypes + micro-
scope slide: Lepidonotus carinulatus, Red Sea, coll. C. Ehrenberg, ZMB 491. — R e d S e a : 1 spm., Gulf of Suez, reef, coll. R.Ph.
Dollfus, 30 December 1928, MNHN A894; 3 spms., Gulf of Suez, 28°14'N/33°23'E, 22 m, muddy sand and madreporarians,
coll. R.Ph. Dollfus, 25 December 1928 [collecting data supplemented according to Billard (1933)], MNHN A360; 1 spm., Gulf
of Suez, 28°14'N/33°23'E, 22 m, muddy sand and madreporarians, coll. R.Ph. Dollfus, 25 December 1928 [collecting data sup-
plemented according to Billard (1933)], MNHN A894; 4 spms., Egypt, Gulf of Suez, Sinai Peninsula, Tor (= At-Tur), coll. Hart-
meyer, det. Augener as Lepidonotus sp., ZMH V.6142; 1 spm., Agig Bay, ca. 8 m, coll. C. Crossland, 1904‑05, BMNH
1941.4.4.189; 17 spms., Sudan, “Pearl fishery investigations”, coll. C. Crossland, October 1904 - May 1905, BMNH 2003.84‑93;
3 spms., Sudan, Sanganeb Atoll, 40 m, southern tip (south jetty), coll. D. Fiege, 2 October 1992, SMF 13581; 1 spm., Sudan,
Sanganeb Atoll, 50 cm, southern tip (south jetty), from dead Stylophora pistillata, coll. D. Fiege, 25 September 1992, SMF 13578;
2 spms., Sudan, Sanganeb Atoll, 50 cm, south west tip, from dead coral, coll. D. Fiege, 1 October 1992, SMF 13580; 1 spm., Su-
dan, Sanganeb Atoll, 12 m, inner reef, from dead Acropora sp., coll. D. Fiege, 9 April 1992, SMF 13579; 5 spms., Ile Abulat (=Abū
Latt Island), Saudi Arabia, 19°58'N/40°7'E, “Calypso”, with Syllis variegata, coll. Drach, Cherbonnier and Mercier, 1952 [collecting
data supplemented according to Fauvel (1955)], MNHN A364. — A r a b i a n S e a : 1 spm., Socotra Island, Qaryih,
12°38.924'N/54°14.387'E, 10‑12 m, coll. T. Wehe, 20 April 2000, NHCY; 1 spm., Socotra Island, Qaryih,
12°38.924'N/54°14.387'E, 10‑12 m, coll. T. Wehe, 20 April 2000, NHCY; 1 spm., Socotra Island, SE of Qualansiyah,
12°40.264'N/53°27.204'E, 6‑7 m, from living Stylophora sp., coll. M. Apel, 8 March 1999, NHCY; 1 spm., Socotra Island, SE of
Qualansiyah, 12°40.264'N/53°27.204'E, 6‑7 m, from dead Pocillopora (?) sp., coll. M. Apel, 8 March 1999, SMF 13587; 1 spm.,
Socotra Island, SE of Qualansiyah, 12°40.264'N/53°27.204'E, 6‑7 m, from dead Pocillopora (?) sp., coll. M. Apel, 8 March 1999,
NHCY; 1 spm., Socotra Island, SE of Qualansiyah, 12°40.264'N/53°27.204'E, 6‑7 m, from dead Pocillopora (?) sp., coll. M. Apel,
8 March 1999, SMF 13593; 8 spms., Socotra Island, Qualansiyah Bay, 12°41.026'N/53°28.309'E, 3‑5 m, patchy rocky substrate
with gravel and sand, coll. M. Apel, 10 March 1999, SMF 13589; 14 spms., Socotra Island, Qualansiyah Bay,
12°41.026'N/53°28.309'E, 3‑5 m, patchy rocky substrate with gravel and sand, coll. M. Apel, 10 March 1999, SMF 13583;
2 spms., Socotra Island, Qualansiyah Bay, 12°41.026'N/53°28.309'E, 3‑5 m, patchy rocky substrate with gravel and sand, from
dead branching coral, coll. M. Apel, 10 March 1999, NHCY; 3 spms., Socotra Island, Steroh, Nogid, S‑coast of Socotra,
12°19.006'N/53°52.845'E, 13‑15 m, bedrock, coll. M. Apel, 14 March 1999, NHCY; 2 spms., Socotra Island, Hawlaf, off the
jetty, 12°41.062'N/54°4.508'E, 12‑14 m, Galaxea-colony on sandy ground, coll. M. Apel, 19 March 1999, NHCY; 1 spm., So-
cotra Island, off Hawlaf dune, 12°40.519'N/54°4.170'E, 3‑5 m, coll. M. Apel, 21 March 1999, SMF 13592; 1 spm., Socotra Is-
land, off Hawlaf dune, 12°40.519'N/54°4.170'E, 3‑5 m, coll. M. Apel, 21 March 1999, NHCY; 4 spms., Socotra Island, off
Hawlaf dune, 12°40.519'N/54°4.170'E, 3‑5 m, from Acropora sp., coll. M. Apel, 21 March 1999, SMF 13582; 8 spms., Socotra
Island, off Hawlaf dune, 12°40.519'N/54°4.170'E, 3‑5 m, from dead Acropora cf. valida, coll. M. Apel, 21 March 1999, SMF
13590; 3 spms., Socotra Island, Shanitan, 600 m off cold store, 12°40.156'N/54°2.850'E, 8 m, from almost completely dead Acro-
pora sp., coll. M. Apel, 12 March 1999, SMF 13586; 1 spm., Socotra Island, Ras Bidou, SW of Qualansiyah,
12°40.282'N/53°26.064'E, 12‑14 m, under stones and coral rubble, coll. M. Apel, 30 March 1999, NHCY; 1 spm., Socotra Is-
land, Ras Qataninh Bay, 12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E, 18‑20 m, dredge, corals and algae on bedrock, coll.
T. Wehe, 9 April 2000, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E,
18‑20 m, dredge, corals and algae on bedrock, coll. T. Wehe, 9 April 2000, SMF 13585; 1 spm., Socotra Island, Ras Qataninh Bay,
12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E, 18‑20 m, dredge, corals and algae on bedrock, coll. T. Wehe, 9 April 2000,
Fig. 16: Lepidonotus carinulatus (BMNH 1969.261). a: anterior end; b: 1st right elytron; c: 2nd right elytron; d: warty macrotubercle
from first elytron; e: same, from second elytron; f: warty, submarginally located microtubercles from 1st elytron; g: same, from 6th
elytron; h: carinate microtubercles and pigmentation; i: digitiform papillae on elytra surface; j: fringing papillae; k: right parapodi-
um from midbody segment, anterior view; l: same, posterior view; m: notochaetae; n: upper neurochaetae; o: lower neurochaetae.
SMF 13588; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E, 18‑20 m, dredge,
corals and algae on bedrock, coll. T. Wehe, 9 April 2000, SMF 13584; 1 spm., Socotra Island, Ras Qataninh Bay,
12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll. T. Wehe, 9 April 2000, NHCY; 1 spm., Socotra Island,
Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll. T. Wehe, 9 April 2000, SMF 13591;
1 spm., Socotra Island, Ras Eriyhan-Quariyah, west of di-Hamri, 12°38.924'N/54°14'387'E, 10‑12 m, coll. T. Wehe, 10 April
88 T. Wehe
2000, NHCY; 1 spm., Socotra Island, near Hawlaf, 1‑2 m, rocky substrate intermixed with sandy patches, coll. T. Schils, 27 April
2000, NHCY; 1 spm., south of Oman, 18°03'30''N/57°02'30''E, lithothamnion, 38 m, triangular dredge, coll. “John Murray Ex-
pedition 1933‑34”, 29 October 1933, BMNH 1937.9.2.46; 2 spms., south of Oman, 19°22'36''N/57°53'00''E, rock, shingle,
shells, lithothamnion, 13.5 m, triangular dredge, coll. “John Murray Expedition 1933‑34”, 2 November 1933, BMNH
1937.9.2.47‑49; 2 spms., south of Oman, 18°03'30''N/57°02'30''E, lithothamnion, 38 m, triangular dredge, coll. “John Murray
Expedition 1933‑34”, 29 October 1933, BMNH 1937.9.2.50‑51; 2 spms., south of Oman, 19°22'36''N/57°53'00''E, rock, shin-
gle, shells, lithothamnion, 13.5 m, triangular dredge, coll. “John Murray Expedition 1933‑34”, 2 November 1933, BMNH
1937.9.2.52‑54; 1 spm., Buleji, Karachi, Pakistan, intertidal rock, 6 September 1989, J. Mustaquim, SMF 13577; 1 spm., Hawks
Bay, Karachi, Pakistan, intertidal rock, 20 December 1990, J. Mustaquim, SMF 13576. — G u l f o f O m a n : 1 spm. + 1 micro-
scope slide, 7 miles WNW of Ras Makki, 25°25'N/59°28'E, Petersen-grab, 14 m, sand with a little claq, coll. Løppenthin, 1 April
1938, ZMUC POL-1370. — T h e G u l f : 9 spms., 15 miles of coast of Oman (now UAE), 24°55' to 25°10'N/54°40' to
55°10'E, dredge, coll. “Mission Bonnier - Perez, Côtes d’Arabie”, 14‑16 March 1901, MNHN A21; 2 microscope slides belonging
to MNHN A21, IEA P.59’; IEA P.59; 1 spm., Qatar, 25°47'N/52°17'E, “Umitaka-Maru” December 1968, det. Mohammad [col-
lecting data supplemented according to Mohammad (1972 b)], BMNH 1969.261; 1 microscope slide, Bahrain, coll. M.N. Bo-
goyawlensky, 12 November 1902, IEA P.55’; 1 microscope slide, Bahrain, Bouchir (=Būshehr), coll. M.N. Bogoyawlensky, IEA
P.56’; 1 microscope slide, Bahrain, coll. M.N. Bogoyawlensky, IEA P.57; 1 spm., north of Abū ‘Alī island, Saudia Arabia, 9 m, whip-
coral habitat, coll. M. Apel, 12 November 1992, SMF 13575; 2 spms., Abū ‘Alī island, Saudia Arabia, 5‑10 m, dredge, sand with
corals, coll. R. Janssen, 1 June 1995, SMF 13642; 1 spm., “Pole Reef ”, from Porites sp., 1‑3 m, coll. M. Apel, 22 February 1993,
SMF 13650; 1 spm., Kuwait, 29°20'N/48°06'E, on rocks near low-water mark, stratified calcerous sand stone and light to dark grey
sand (with very little mud), coll. & det. B.M. Mohammad, November 1967 - December 1968 [collecting data supplemented ac-
cording to Mohammad (1971)], BMNH 1969.187; 1 spm., as Lepidonotus tenuisetosus in Mohammad (1971), Kuwait, coll. & det.
B.M. Mohammad, November 1967 - December 1968 [collecting data supplemented according to Mohammad (1971)], BMNH
1969.188; 1 microscope slide, Coveik, coll. M.N. Bogoyawlensky, 6‑7 April 1902, IEA P.56; 1 spm., as Lepidonotus jacksoni, Khor
Abdullah, subtidal, 2 m, 25 °C, coll. H.E. Gruner, 12 October 1971, ZMH P‑13640; 2 spms. + 1 microscope slide, 12 miles ENE
of Būshehr light-ship, 26°38'N/51°10'E, dredge and trawl, 23 m, shells, gravel, coll. G. Thorson, 27 March 1937, ZMUC POL-
1369; 1 spm. + 1 microscope slide, 5 miles SE of Tunb-light, 26°12'N/55°22'E, Petersen-grab, 38‑60 m, coll. Løppenthin, 7 April
1938, gravel and shells, ZMUC POL-1371; 1 spm. + 1 microscope slide, 2 miles S. by W. of Duwwan, 26°33'N/54°33'E, Petersen-
grab, 12 m, clay with a little sand, coll. Løppenthin, 8 April 1938, ZMUC POL-1372; 2 spms., Bouchir (=Būshehr), Iran, III-
1902, coll. M.N. Bogoyawlensky, MNHN A411; 4 microscope slides, Bouchir (=Būshehr), Iran, coll. M.N. Bogoyawlensky, March
1902, IEA P.53’; P.54; IEA P.54’; P.55.
M e a s u r e m e n t s : Syntype ZMB 1072: cs (26) 10/4/3. Syntypes ZMB 491: cs (26) 11/3.5/2.5, cs (26) 10/3.5/2.5, cs (26)
8/2/1.5. MNHN A894: af (10) 4/2/1.5 + mf (13) 6. MNHN A360: cs (26) 7/2.2/1.5, cs (26) 9/3/2, cs (26) 8/3/2 [af (13) 4 + pf
(13) 4]. MNHN A894: cs (22) 2.2/1/0.7. ZMH V.6142: cs (26) 10/2/1.7, cs (26) 12/3.5/2.8, cs (26) 12/3.5/2.8, cs (26) 8/2/1.7.
BMNH 1941.4.4.189: af (20) 11/3/2.5. BMNH 2003.84‑93: cs (26) 15/4.5/3, cs (26) 17/6/4, cs (26) 19/6/4, cs (26) 13/4/3, cs
(26) 15/5/4, cs (26) 17/5/4, cs (26) 17/6/4, cs (26) 15/5/3, cs (26) 18/5.5/4, cs (26) 13/5/3, cs (26) 15/5/4, af (12) 8/4.5/3.5, af
(20) 13/4/3, af (15) 9/4.5/3.5, af (18) 13/4.5/3.5, af (18) 10/5/4, af (18) 9/4/3. SMF 13581: cs (26) 5/1.5/1, cs (26) 6/2.2/1.8,
cs (26) 3/1.3/1. SMF 13578: cs (26) 7/2.5/1.5 [af (15) 4 + pf (11) 3]. SMF 13580: cs (26) 5/2/1.5, cs (26) 8/3/2. SMF 13579: af
(20) 4/2/1.5. MNHN A364: cs (26) 11/4/3, cs (26) 7/2.5/2, cs (26) 8/2.5/1.8, cs (26) 11/3/2.5, cs (26) 11/4/3 [af (13) 5 + pf (13)
6]. NHCY: cs (26) 6/2.5/1.8. SMF: cs (26) 8/3/2. NHCY: cs (26) 13/4/3. SMF 13587: cs (26) 8/4/3. NHCY: cs (26) 8/2/1.8.
SMF 13593: cs (26) 12/5/4. SMF 13589: cs (26) 9/2/1.7, cs (26) 8/2.4/2, cs (26) 8/2.5/2, cs (26) 9/2.5/2, cs (26) 13/4/3, cs (26)
11/3.5/2.5, cs (26) 9/3.5/2.5, af (13) 4/3/2. SMF 13583: cs (26) 11/4/3, cs (26) 10/4/3, cs (26) 9/3.2/2.8, cs (26) 10/3.5/2.5,
cs (26) 7/3.5/2.5, cs (26) 9/3/2, cs (26) 11/4/3, cs (26) 10/4/3, cs (26) 10/3.5/2.5, cs (26) 11/5/4, cs (26) 10/5/4 [af (15) 6 +
pf (11) 4], cs (26) 12/4/3 [af (17) 7 + pf (9) 5], af (21) 9/4/3, af (19) 7/4/3. NHCY: cs (26) 7/2.5/2, cs (26) 5/1.5/1.2. NHCY:
cs (26) 9/3.2/2.5, cs (26) 11/3.5/2.5 [af (14) 6 + pf (12) 5], af (16) 5/2.5/2. NHCY: cs (26) 11/3.5/2.5, cs (26) 5/1.5/1. SMF
13592: cs (26) 10/3/2. NHCY: cs (26) 13/3/2.5. SMF 13582: cs (26) 12/3/2.5, cs (26) 11/4/3, cs (26) 9/2.5/2, cs (26) 10/3.5/2.5.
SMF 13590: cs (26) 11/3.2/2.7, cs (26) 10/3.5/2.5, cs (26) 13/4/3, cs (26) 8/2.5/2, cs (26) 11/3.5/2.5, cs (26) 8/2.2/1.8, cs (26)
8/2/1.5, cs (26) 8/2.5/2. SMF 13586: cs (26) 9/3/2, cs (26) 11/ 3.5/2.5 [af (13) 5 + pf (13) 6], cs (26) 9/3/2 [af (13) 4 + pf (13) 5].
NHCY: cs (26) 12/4/3. NHCY: af (14) 7/5/4. SMF 13585: cs (26) 13/4.5/3.5. SMF 13588: cs (26) 14/6/4. SMF 13584: cs (26)
13/3.5/2.5. NHCY: cs (26) 10.5/4/3 [af (13) 6 + pf (13) 4.5]. SMF 13591: cs (26) 14/5/4. NHCY: cs (26) 10/3/2. NHCY: cs (26)
12/4/3. BMNH 1937.9.2.46: cs (26) 12/4/3. BMNH 1937.9.2.47‑49: cs (26) 12/4/3, cs (26) 16/6/5. BMNH 1937.9.2.50‑51: cs
(26) 12/4/3, cs (26) 13/5/4. BMNH 1937.9.2.52‑54: cs (26) 13/5/4 [af + pf ], cs (26) 10/3/2. SMF 13577: cs (26) 12/5/3. SMF
13567: cs (26) 13/4.5/3.5. ZMUC POL-1370: cs (26) 9/4/3. MNHN A21: cs (26) 8/3.5/2.5, cs (26) 7/3/2, cs (26) 7/4/3, cs
(26) 6/3.5/2.5, cs (26) 9/4/3, cs (26) 10/4.5/3 [af (15) 5 + pf (11) 5], cs (23) 4/1.5/1, af (22) 7/4/3, af (19) 6/4/3, pf (5), pf (13).
BMNH 1969.261: cs (26) 14/4/3. SMF 13575: cs (26) 9/3/2. SMF 13642: af (18) 4/1.8/1.2, af (13) 3/2/1.5. SMF 13650: cs (26)
10/3.2/2.2. BMNH 1969.187: cs (26) 13/4/3. BMNH 1969.188: cs (26) 8/3/2. ZMH P‑13640: cs (26) 11/4/3. ZMUC POL-
1369: cs (26) 7/2/1.5, cs (26) 9/3/2. ZMUC POL-1371: af (12) 5/5/4. ZMUC POL-1372: cs (26) 11/4/2.5. MNHN A411: cs
(26) 9/3/2, af (24) 8/3.2/2.5. IEA P.53’: cs (26) 8/2.5/1.8.
D i a g n o s i s : Without nuchal fold, with two nuchal nodules. Elytra surface reticulate, with
oval to rounded macro- and microtubercles, anterior ones flattened, smooth or carinate, central
and posterolateral ones warty; margin with fringing papillae. Neurochaetae bidentate.
D e s c r i p t i o n : Body flattened dorsoventrally, subrectangular in cross-section. Prostomium
(Fig. 16 a) rounded to hexagonal, about as wide as long, sometimes withdrawn into anterior seg-
ments. Styles of antennae similar, median one longer than lateral ones, with subterminal swelling
and terminal filum. A band of dark pigmentation may be present subterminally. Palps stout, conical,
often abruptly terminating in a filiform tip. Two pairs of eyes, anterior pair dorsolaterally on more
or less widest part of prostomium, posterior pair in front of hind margin. Tentaculophores with one
to three chaetae. Tentacular cirri shaped as median antenna, of similar length. Facial tubercle usu-
ally distinct. Buccal segment without nuchal fold, but with two nuchal nodules on anterior margin
(rarely inconspicuous). Buccal cirri longer and more proximally inserted than following ventral cirri,
shaped as tentacular cirri. Elytra covering dorsum completely. First pair of elytra rounded to oval (Fig.
16 b), second one slightly reniform (Fig. 16 c), following larger, oval. Surface covered with rounded
to oval, flattened tubercles, giving elytra a reticulated appearance. Conical or globular macrotubercles
in central part of elytra. Macrotubercles (Fig. 16 d, 16 e) and posterolaterally inserted microtubercles
(Fig. 16 f, 16 g) warty, with more or less distinct nodules or pointed projections. Microtubercles in
anterior part flattened, smooth or with carina (Fig. 16 h). Additional digitiform papillae (Fig. 16 i)
scattered over surface. Warty tubercles more prominent on anterior elytra. Margins with dense fringe
of long, slender, digitiform papillae (Fig. 16 j) posterolaterally, often covered by inorganic matter.
Dorsal cirri with large, globular to cylindrical cirrophore and long style with slight subterminal swell-
ing and terminal filum. Dorsal tubercles small, nodular. Parapodia (Fig. 16 k, 16 l) with small, coni-
cal notopodia. Neuropodia stout, with slightly longer, subtriangular, prechaetal, acicular lobe and a
bluntly rounded, postchaetal lobe. Ventral cirri short, conical, gradually tapering. Nephridial papillae
usually from about segment 7 to 8 onwards. Notochaetae (Fig. 16 m) in dense bundles, short to long,
with dense rows of spines and pointed to capillary tips. Neurochaetae (Fig. 16 n, 16 o) stouter than
notochaetae, falcate, subdistally thickened, with several rows of spines, distal spines usually much
larger than basal ones, and bidentate tips with small secondary tooth. Sometimes lower neurochaetae
appear unidentate, which seems to be due to abrasion. Neurochaetae on second segment more slen-
der, with more numerous rows of spines and eventually true unidentate tips.
R e m a r k s : There has been some confusion about the appearance of the tubercles on the
elytra, and descriptions of L. carinulatus vary considerably in this respect (see Grube 1869, Grube
1878, Willey 1905, Horst 1917, Augener 1922 b, Day 1975, Hanley & Burke 1990, Hanley
& Burke 1991 a, Imajima 1997). Unfortunately, Grube (1869, 1878) as well as Augener (1922 b),
who re-examined Grube’s types and the specimens from the Philippines, did not mention the oc-
currence of any warty tubercles. However, re-examination of the syntypes during this study clearly
demonstrated the presence of both smaller warty tubercles near the posterolateral margin and
larger, more weakly warty tubercles in the central part of the elytra. It seems that the appearance of
warty tubercles is somewhat variable amongst specimens as well as along the body. Some specimens
appear to have strongly warty tubercles, whereas in other specimens they are much more smooth.
The latter is true of the syntypes. In general, distinct warty tubercles are found on more anterior
elytra. Comparing two specimens from both ends of the range, one might be tempted to consider
them as two different species. However, the examination of about 130 specimens revealed gradual
transitions among specimens, making a discrete separation of two species impossible.
B i o l o g y & e c o l o g y : L. carinulatus is most commonly found on sandy substrates and
amongst corals (see above), but is also reported from algae, sea grass and pearl oyster banks (Hart-
mann-Schröder in Hartmann-Schröder & Hartmann 1991).
90 T. Wehe
D e p t h r a n g e : So far recorded between the intertidal and 60 m; elsewhere reported down to
200 m (Hartmann-Schröder in Hartmann-Schröder & Hartmann 1991).
D i s t r i b u t i o n : Red Sea, Arabian Sea: Socotra Archipelago, Gulf of Oman, The Gulf. Else-
where: All over Indo-West Pacific: i.a. Madagascar, Sri Lanka, Indonesia, Philippines, Hong Kong,
South China Sea, Australia, New Caledonia, Japan (Willey 1905, Fauvel 1953, Hanley & Burke
1991 a, Hanley 1992, Imajima 1997, Barnich et al. 2004).
Lepidonotus clava (Montagu, 1808) Fig. 17 a‑j

Aphrodita clava Montagu, 1808. — Transactions of the Linnean Society of London 9(2): 108‑109, pl. 7: fig. 3.
Lepidonotus clava. — Fauvel 1927, The Transactions of the Zoological Society of London 22/4(1): 413; Fauvel 1955, Annales de
l’Institute Océanographique Paris 30: 102; Tebble & Chambers 1982, Royal Scottish Museum Studies: 19‑20, figs 24‑25;
Chambers & Muir 1997, Synopsis of the British Fauna (New Series) 54: 114, fig. 31; Barnich & Fiege 2003, Abhandlun-
gen der Senckenbergischen Naturforschenden Gesellschaft 559: 82‑86, fig. 43, pl. 2.1.
Ty p e l o c a l i t y : South Devon, Great Britain.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Southern coast of Devon, Great Britain, Coll. Montagu, Mus. Leach (presump-
tive [according to label in vial]), BMNH ZB.1972.60. — S u e z C a n a l : 1 spm., Port Said, coal basin on eastern bank of harbour,
5 m, sand and mud, coll. “Cambridge Expedition to the Suez Canal”, 28 December 1924 [collecting data supplemented accord-
ing to Fox (1926)], BMNH 1926.11.12.3; 4 spms., Port Said, coal basin on eastern bank of harbour, 5 m, sand and mud, coll.
“Cambridge Expedition to the Suez Canal”, 19 December 1924 [collecting data supplemented according to Fox (1926)], BMNH
1926.11.12.5‑8. — R e d S e a : 1 spm., off Saudi Arabian coast, Île Abulat (=Abū Latt Island), 19°58'N/40°7'E, beach near
camp, 0‑2 m, “Calypso”, coll. Drach, Cherbonnier und Mercier, 1952 [collecting data supplemented according to Fauvel (1955)],
MNHN A364.
M e a s u r e m e n t s : Holotype: cs (26) 30/11/9. BMNH 1926.11.12.3: cs (26) 25/7/5. BMNH 1926.11.12.5‑8: cs (26)
43/11/9, cs (26) 26/6/4.5, cs (26) 24/7/5, cs (26) 26/8/7. MNHN A364: cs (26) 9/2.5/2.
D i a g n o s i s : Without nuchal fold. Elytra leaving parts of middorsum uncovered, with warty
macrotubercles and smooth or ornamented microtubercles, margin without fringing papillae.
Notochaetae very fine. Neurochaetae unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 17 a) rounded to hexagonal,
about as long as wide, sometimes withdrawn into anterior segments. Styles of antennae similar,
median style longer than lateral styles, with subterminal swelling and terminal filum. Palps stout,
conical, long, gradually tapering, more or less abruptly terminating into a short terminal filum,
with rows of minute papillae. Two pairs of eyes (not visible on BMNH 1926.11.12.3), anterior
pair dorsolaterally on more or less widest part of prostomium, posterior pair in front of hind mar-
gin, hidden if prostomium withdrawn. Everted pharynx (Fig. 17 b) distally with two semicircles
of 9 papillae each and two pairs of chitinous jaws. Tentaculophores with one to four chaetae. Ten-
tacular cirri shaped as antennae, as long as or longer than median antenna, dorsal tentacular cirri
longer than ventral cirri. Facial tubercle distinct. Second segment without nuchal fold. Buccal cirri
longer and inserted more proximally than following ventral cirri, shaped as tentacular cirri. Elytra
moderate in size leaving parts of middorsum uncovered. First pair rounded (Fig. 17 c), second
reniform, following elytra broad, oval (Fig. 17 d). Central and posterior surface with prominent,
scaly or warty ornamented macrotubercles (Fig. 17 e), remaining surface with scattered smooth
or ornamented microtubercles, and small, cylindrical papillae. Macrotubercles on anteriormost
pairs of elytra more numerous and raised conical, becoming fewer in number and more flattened
on posterior elytra. Margins without fringe of papillae. Dorsal cirri with large, globular to conical
cirrophore and long, sometimes slightly dorsoventrally flattened style with subterminal swelling
and terminal filum. Dorsal tubercles indistinct. Parapodia (Fig. 17 f, 17 g) with very small, conical
notopodia. Neuropodia large, with subtriangular, prechaetal, acicular lobe and bluntly rounded,
postchaetal lobe of similar length. Ventral cirri short, conical, gradually tapering. Nephridial papil-
lae usually distinct from about segment 8‑9 onwards. Notochaetae (Fig. 17 h) in dense bundles,
Fig. 17: Lepidonotus clava (BMNH 1926.11.12.5‑8). a: anterior end; b: everted pharynx; c: 1st left elytron; d: 7th left elytron e:
warty tubercle; f: right parapodium from segment 12, anterior view; g: same, posterior view; h: notochaetae; i: upper neurochaeta;
j: lower neurochaeta.
92 T. Wehe
short to long, with numerous rows of spines, below blunt or pointed tips. Neurochaetae (Fig.
17 i, 17 j) much stouter, falcate, subdistally with slight swelling and several rows of spines, distal
larger than basal spines, and slightly bent, blunt, unidentate tips. Neurochaetae of second segment
slightly different, more slender with more numerous rows of spines, intermediate to following neu-
rochaetae.
R e m a r k s : The specimen reported by Fishelson & Rullier (1969) (see Wehe & Fiege
2002) could not be located for examination. However, the other specimens examined confirm the
occurrence of the species in the Red Sea and the Suez Canal. According to the literature, L. clava is
widely distributed in boreal and subtropical waters, but the specimens reported here confirm also
its occurrence in tropical regions.
B i o l o g y & e c o l o g y : The species is found in sand and mud, as well as between algae, in
coralligène, in sea grass beds and on hard grounds (Barnich & Fiege 2003).
D e p t h r a n g e : So far recorded between 0 and 5 m; elsewhere reported down to 160 m ac-
cording to Barnich & Fiege (2003).
D i s t r i b u t i o n : Suez Canal: Port Said on Mediterranean entrance, Red Sea. Elsewhere:
North Atlantic, south to West Africa, Mediterranean Sea, Indian Ocean, North Pacific (Campoy
1982, Tebble & Chambers 1982, Hartmann-Schröder 1996, Chambers & Muir 1997, Barnich
& Fiege 2003).
Lepidonotus cristatus (Grube, 1876) Fig. 18 a‑k, Pl. 2 b, c

Polynoe cristata Grube, 1876. — Jahres-Bericht der Schlesischen Gesellschaft für vaterländische Cultur 53: 57, 62.
Polynoe impatiens. — Quatrefages 1866: 221‑223 [partim, see L. impatiens; not Savigny in Lamarck, 1818].
Polynoe (Lepidonotus) impatiens. — Grube 1869, Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu
Berlin aus dem Jahre 1869: 488‑489 [not Savigny in Lamarck, 1818].
Polynoe (Lepidonotus) cristata. — Grube 1878, Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8):
27‑29, pl. 2: fig. 3.
Lepidonotus cristatus. — McIntosh 1885, Report on the scientific results of the voyage of H.M.S. Challenger during the years
1873‑76, R.N. Zoology 12: 67‑69, pl. 11: figs 2, 3, pl. 171: fig. 1, pl. 11A: figs 10, 11; Gravier 1902, Nouvelles Archives
du Muséum 4 e série 3: 210‑214, textfigs 214‑218, pl. 7: figs 104‑110, pl. 9: fig. 136; Potts 1910, The Transactions of the
Linnean Society Series 2, Zoology 8: 332‑333 [not var. ornata & var. echinata]; Fauvel 1919, Archives de Zoologie Expéri-
mentale et Générale 58: 329; Day 1967, A monograph on the Polychaeta of Southern Africa. Part I. Errantia: 82, fig. 1.14.
f‑I; Barnich et al. 2004, Species Diversity 9: 298.
Lepidonotus oculatus Seidler 1923. — Archiv für Naturgeschichte 89 (Abt. A/11): 43‑47, figs 4‑8 [not Baird, 1865].
Ty p e l o c a l i t y : Philippines.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Lepidonotus oculatus, Australian seas?, BMNH 1863.9.23.3. H o l o t y p e :
Lepidonotus cristatus ornata, Salomon Islands, ca. 110‑220 m, “Sealark Expedition”, coll. J.S. Gardiner, M.A. Cains, 3 July 1905,
BMNH 1924.3.1.110. H o l o t y p e : Lepidonotus cristatus echinata, Mauritius, “Sealark Expedition”, coll. J.S. Gardiner BMNH
1924.3.1.141. — R e d S e a : 1 spm., Gulf of Suez, coll. E. Bannwarth, 1912, SMF 13676; 1 spm., Suez, coll. E. Bannwarth,
1914, ZMH PE96; 1 spm., as Polynoe impatiens, “M. Botta, Mer Rouge, A1(R.) - 1868 - No 16 a”, MNHN A28; 1 spm. + mi-
croscope slide, as Lepidonotus impatiens, coll. C. Ehrenberg, ZMB 492; 1 spm., Port Sudan, coll. Werner, 29 April 1914, NHMW
17064; 1 spm., Perim Island, coll. Dr. Jousseaume, 1894, det. C. Gravier, 1901, MNHN A21; 2 spms., Berenice (Egypt), Jiddah,
Mamuret-el-Hamidije (harbour of Al Lith), Saudi Arabia, “Pola Expedition”, 1895‑1898 [collecting data supplemented according
to Schefbeck (1991)], NHMW V185; 1 spm., Dschidda (= Jiddah), Saudi Arabia, “Pola Expedition”, end of December 1895 [col-
lecting data supplemented according to Schefbeck (1991)], NHMW 17061. — G u l f o f A d e n : 1 spm., Djibouti, coll. M.H.
Coutière, 1897, det. C. Gravier, 1901, MNHN A21; 1 spm., Aden, coll. Dr. Jousseaume, 1895, det. C. Gravier, 1901, MNHN
A21; microscope slide, Djibouti, Gulf of Tadjoura, Musha Island, det. C. Gravier, IEA D‑52, probably belonging to MNHN
A21 from Djibouti). — A r a b i a n S e a : 1 spm., Socotra Island, Ras Adho (or Ras Hammadara), 12°38.534'N/54°15.770'E,
10 m, coll. M. Apel, 25 March 1999, SMF 13675); 1 spm., Socotra Island, Hawlaf, 12°40.662'N/54°4.497'E, from Acropora sp.,
4‑5 m, coll. M. Apel, 25 March 2000, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑12 m, coll.
M. Apel, 9 April 2000, SMF 13674.
M e a s u r e m e n t s : SMF 13676: cs (26) 38/14/10. ZMH PE96: cs (26) 50/22/16. MNHN A28: cs (26) 34/15/13. ZMB
492: cs (26) 40/15/12. NHMW 17064: cs (26) 18/9/6. MNHN A21: cs (26) 35/15/11. NHMW V185: cs (26) 26/12/10 [af (13)
12 + pf (13) 14], cs (26) 32/13/10 [af (13) 16 + pf (13) 16]. NHMW 17061: cs (26) 22/14/10. MNHN A21: cs (26) 33/11/8.
Fig. 18: Lepidonotus cristatus (SMF 13675: a; SMF 13674: b-k). a: anterior end; b: 9th right elytron; c: macrotubercle; d: micro-
tubercles; e: left parapodium from segment 2, anterior view; f: left parapodium from segment 14, anterior view; g: same, posterior
view; h: notochaeta; i: upper neurochaeta; j: lower neurochaetae; k: neurochaeta from e.
MNHN A21: cs (26) 21/12/8. SMF 13675: cs (26) 24/15/11. NHCY: cs (26) 37/17/13. SMF 13674: cs (26) 35/17/12 [af (15)
20 + pf (11) 15].
D i a g n o s i s : With nuchal fold and nuchal nodules. Elytra with two large, sac-like protuber-
ances, with prominent conical, pointed macrotubercles and four-pointed microtubercles, without
fringing papillae. Neurochaetae unidentate.
D e s c r i p t i o n : Body very stout, flattened dorsoventrally. Prostomium (Fig. 18 a) rounded to
hexagonal, wider than long, often withdrawn into anterior segments. Antennae with long, smooth
styles, with subterminal swelling and terminal filum. Styles of lateral antennae only slightly shorter
94 T. Wehe
than median style. Palps very stout, conical, gradually tapering, sometimes abruptly terminating
into a short terminal filum, with (larger specimens) or without minute papillae. Two pairs of eyes,
hind margin, sometimes hidden by nuchal fold, if prostomium withdrawn. Everted pharynx on
distal borders with two semicircles of 11 soft papillae each and two pairs of chitinous jaws. Tentac-
ulophores very long and slender with up to four long, fine chaetae (ZMH PE 96), missing on oth-
er specimens. Tentacular cirri shaped as antennae, about as long as median antenna, dorsal longer
than ventral cirri. Facial tubercle distinct. Buccal segment with more or less distinct semilunar
nuchal fold. Dorsum behind nuchal fold often raised rectangularly, with two more or less distinct
nuchal nodules on anterior edge. Buccal cirri longer and inserted more proximally than following
ventral cirri, shaped as tentacular cirri. Elytra (Fig. 18 b, 18 e) very large, more-or-less oval, typi-
cally folded frill-like. Surface with two very large sac-like protuberances posterior to attachment
scar of elytrophore, sometimes connected to each other medially. Protuberances and posterolateral
margin with prominent, conical, smooth, pointed macrotubercles (Fig. 18 c). Remaining surface
completely covered by microtubercles with four tips (Fig. 18 d), near to margin becoming smaller,
with three or two spines only. Margin without fringing papillae. Elytra in ethanol with irregularly
brown patches scattered over surface, with distinct dark spot above attachment of elytrophore.
Dorsal cirri with very large, globular to conical cirrophore and long style with subterminal swelling
and terminal filum. Dorsal tubercles distinct, globular. Parapodia (Fig. 18 e, 18 f, 18 g) with small
notopodia, more or less conical, with short acicular process. Neuropodia large, with triangular,
prechaetal, acicular lobe, tapering to short acicular process and bluntly rounded, shorter, postcha-
etal lobe. Ventral cirri short, conical, gradually tapering. Nephridial papillae distinct from about
segment 7 onwards. Notochaetae (Fig. 18 h) moderate in number, in loose bundles, about as thick
as or only slightly thinner than neurochaetae, with numerous rows of spines. Neurochaetae falcate,
subdistally with slight swelling and several rows of spines, distal much larger than basal spines,
slightly bent, blunt, unidentate tips. Subterminal swelling on upper neurochaetae (Fig. 18 i) long-
er, more slender, with more numerous spines than lower neurochaetae (Fig. 18 j). Neurochaetae of
second segment modified, much more slender, with more numerous rows of spines and gradually
tapering tips (Fig. 18 k). Neurochaetae of following two segments intermediate.
R e m a r k s : L. cristatus is one of the large-bodied species within the genus, with very large and
conspicuous elytra. Specimens examined, especially those from the Socotra Archipelago, agree very
well with the descriptions and illustrations provided by McIntosh (1885) and Gravier (1902).
According to Grube (1876), the neurochaetae are bidentate. However, referring to his work dating
to 1878, it becomes clear, that he regarded the most distal spine of the subterminal row of spines
as a secondary tooth and thus the tips as bidentate. It is not clear from Grube’s work (1876, 1878),
whether his specimens showed a nuchal fold or not. Unfortunately, the type specimens seem to be
lost, since they are neither mentioned in the catalogue of type specimens by Wiktor (1992) nor by
Hartwich (1993). However, in several other descriptions, i.a McIntosh (1885), Gravier (1902)
or Day (1967 a), a more or less distinct nuchal fold or nuchal nodules are mentioned and illus-
trated, as was observed in the larger specimens examined here. In smaller or improperly preserved
individuals, these structures may be indistinct. Day (1975) had already discussed the difficulties
with these and other characters in Lepidonotus, which often exhibit unnatural conditions due to
preservation artefacts.
Seidler (1923) erroneously synonymised L. cristatus with L. oculatus Baird, 1865, even though
his description undoubtly refers to L. cristatus. Re-examination of the holotype of L. oculatus con-
firmed that it is a separate species (see also remarks on L. spinosus in Hanley & Burke (1991 a),
and Hanley (1993) for detailed description of L. oculatus).
Fig. 19: Lepidonotus glaucus (SMF 13550). a: anterior end; b: 1st right elytron; c: 2nd right elytron; d: 10th right elytron; e: micro-
tubercles from anterior surface; f: microtubercles from posterior surface; g: parapodium from segment 12, anterior view; h: same,
posterior view; i: notochaetae; j: upper neurochaeta; k: lower neurochaeta.
96 T. Wehe
For comparison, the types of L. cristatus ornata and L. cristatus echinata, both of them de-
scribed by Potts (1910), were also re-examined. The first-mentioned subspecies is clearly different
from the nominal species and was synonymised and redescribed as L. spinosus by Hanley & Burke
(1991 a), based on freshly collected individuals from the Chesterfield Islands. The second subspe-
cies also differs without doubt from the nominal species. Only a single specimen of this subspecies
has been reported by Horst (1917) in addition to the original description. However, according to
the description by Horst, this record seems to be doubtful, since the characteristic oval patches on
the elytra are missing. Lepidonotus cristatus echinata Potts, 1910 should also be elevated to species
rank in future works involving this species.
The examination of one of the specimens described by Quatrefages (1866) as Polynoe impa-
tiens revealed that it belongs to L. cristatus (see also remarks on L. impatiens).
The specimens reported by Fishelson & Rullier (1969) and Amoureux et al. (1978) could
not be located for examination, with the exception of a microscope slide (IEA Y100’) labelled as
“L. cristatus, Mer Rouge, 4266”, which probably belongs to one of the specimens reported by these
authors. The elytron and parapodium mounted on the slide, however, belong to Lepidonotus polae
n. sp. described in the present paper.
(Horst 1917).
D i s t r i b u t i o n : Red Sea, Gulf of Aden, Arabian Sea: Socotra Archipelago. The species is
reported for the first time from the Arabian Sea. Elsewhere: Throughout the Indo-West Pacific (i.a.
McIntosh 1885, Willey 1905, Potts 1910, Horst 1917, Fauvel 1953, Barnich et al. 2004).
Lepidonotus glaucus (Peters, 1854) Fig. 19 a‑k, Pl. 2 d

Polynoe glauca Peters, 1854. — Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königlich Preussischen Akad-
emie der Wissenschaften zu Berlin aus dem Jahre 1854: 610; Peters 1855, Archiv für Naturgeschichte 21(1): 38.
Polynoe (Lepidonotus) quadricarinata Grube, 1868 a. — Jahres-Bericht der Schlesischen Gesellschaft für vaterländische Cultur 45:
50; Grube 1868 b, Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 18: 630; Grube
1869, Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin aus dem Jahre 1869: 486‑488.
Lepidonotus obscurus Gravier, 1902. — Nouvelles Archives du Muséum 4 e série 3: 218‑222, textfigs 224‑227, pl. 8: figs
118‑122.
Lepidonotus carinatus Potts, 1910. — The Transactions of the Linnean Society Series 2, Zoology 8: 334‑335, pl. 18: fig. 1.
Lepidonotus stellatus Baird, 1865. — Journal of the Linnean Society of London, Zoology 8: 185; Fauvel 1919, Archives de Zoolo-
gie Expérimentale et Générale 58: 329‑330.
Lepidonotus glaucus. — Seidler 1923, Archiv für Naturgeschichte 89 (Abt. A/11): 79‑81; Monro 1939 b, Annals and Maga-
zine of Natural History 11(4): 169; Day 1957, Annals of the Natal Museum 14(1): 60‑61; Day 1965, Bulletin of the Sea
Fisheries Research Station 38(7): 16; Kiseleva 1971, Naukova Dumka, Kiev: 76; Hanley & Burke 1990, Proceedings of the
Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia 1: 221‑223, fig.
8; Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 68‑69, fig. 21.
Lepidonotus platycirrus Day, 1951. — Annals of the Natal Museum 12(1): 11‑14, textfigs 2 a‑f.
Lepidonotus hedleyi Fauvel 1955. — Annales de l’Institute Océanographique Paris 30: 103 [not Benham, 1915]; ? Fishelson &
Rullier 1969, Israel Journal of Zoology 18: 54.
Ty p e l o c a l i t y : Mozambique, Indian Ocean.
S p e c i m e n s e x a m i n e d : H o l o t y p e + 2 microscope slides: Lepidonotus glaucus, Mozambique, ZMB 18. 6 Syntypes:
Lepidonotus quadricarinatus, Tor (= At-Tur), Sinai Peninsula, Gulf of Suez, Red Sea, coll. C. Ehrenberg, ZMB 490. 1 S y n t y p e
(dried out): Lepidonotus quadricarinatus, Red Sea, Coll. E. Grube, ZMB 1079. 2 S y n t y p e s (?): Polynoe quadricarinata, Red Sea,
coll. Frauenfeld, 1855, MPW 546. 1 S y n t y p e : Polynoe (Lepidonotus) quadricarinata, Tor (= At-Tur), Sinai Peninsula, Gulf of
Suez, Red Sea, coll. C. Ehrenberg, USNM 074368 (exchange with ZMB 490). 8 S y n t y p e s : Lepidonotus obscurus, Djibouti,
Gulf of Tadjoura, Gulf of Aden, coll. M.H. Coutière, 1897, det. C. Gravier, 1902, MNHN POLY TYPE 1432. H o l o t y p e :
Lepidonotus carinatus, Seychelles, Praslin Reef, Indian Ocean, coll. “Sealark Expedition 1905”, J.S. Gardiner, BMNH 1924.3.1.62.
H o l o t y p e : Lepidonotus stellatus Baird, 1865, Australian Seas?, coll. Bowerbank, BMNH 1863.9.23.4. H o l o t y p e : Lepidonotus
platycirrus, Port St. Johns, South Africa, coll. J. Day, BMNH 1961.16.5. — R e d S e a : 8 spms., as Lepidonotus platycirrus (second
label saying L. glaucus), Suakin harbour, Sudan, coll. C. Crossland 1905, BMNH 1941.4.4.190/4; 2 spms., Sudan, Sanganeb Atoll,
north jetty, 2 m, from dead Stylophora sp., coll. Hamza, 29 September 1992, SMF 13548; 1 spm., Sudan, Sanganeb Atoll, north
jetty, edge of the lagoon, 1 m, from dead Pocillopora sp., coll. D. Fiege, 29 September 1992, SMF 13550; 7 spms., as Lepidonotus
quadricarinatus, Jiddah, Mamuret-el-Hamidije (harbour of Al Lith), Saudi Arabia, “Pola Expedition”, 1895‑1898 [collecting data
supplemented according to Schefbeck (1991)], NHMW V17058; 3 spms., as Lepidonotus hedleyi, off Saudi Arabian coast, Île Abu-
lat (=Abū Latt Island), 19°58'N/40°7'E, from Acropora sp., “Calypso”, coll. Drach, Cherbonnier und Mercier, 1952 [collecting data
supplemented according to Fauvel (1955)], MNHN A364; 1 spm., as Lepidonotus hedleyi Benham, epi- and endofauna of Porites II,
“Calypso”, coll. Drach, Cherbonnier und Mercier, 1952 [collecting data supplemented according to Fauvel (1955)], MNHN A364;
1 spm., as Lepidonotus hedleyi, epi- and endofauna of Madreporia, “Calypso”, coll. Drach, Cherbonnier und Mercier, 1952 [col-
lecting data supplemented according to Fauvel (1955)], MNHN A364; Elytra only, as Lepidonotus hedleyi, epi- and endofauna of
Madreporia, “Calypso”, coll. Drach, Cherbonnier und Mercier, 1952 [collecting data supplemented according to Fauvel (1955)],
MNHN A364; microscope slide, as Lepidonotus hedleyi, off Saudi Arabian coast, Île Abulat (= Abū Latt Island), 19°58'N/40°7'E,
coll. “Calypso”, Drach, Cherbonnier und Mercier, 1952 [collecting data supplemented according to Fauvel (1955)], IEA A‑64’.
— G u l f o f A d e n : 2 spms., as Lepidonotus stellatus, Djibouti, Gulf of Tadjoura, from Hircinia sp., elytra blue, coll. C. Gravier,
12 January 1904, MNHN A21; 1 spm., as Lepidonotus stellatus (= L. obscurus), Djibouti, Gulf of Tadjoura, Musha Islands, Maskalla
Island, from polyps, coll. C. Gravier, 23 January 1904, MNHN A21; 1 spm., as Lepidonotus stellatus, Djibouti, Gulf of Tadjoura,
“Plateau du Serpent”, on and under stones, coll. C. Gravier, 7 January 1904, MNHN A21; 1 spm., as Lepidonotus stellatus (= L.
obscurus), Djibouti, Gulf of Tadjoura, E of housing, in muddy sand, coll. C. Gravier, 1904, MNHN A21; microscope slide, as
Lepidonotus stellatus (L. obscurus), Djibouti, IEA D.52’, probably belonging to MNHN A21. — A r a b i a n S e a : 1 spm., Socotra
Island, 8 km E of Rhiy di Qatanhin, Nogid, southern coast of Socotra, off Steroh, 12°18.877'N/53°37.413'E, 15‑17 m, on black
holothurian, coll. M. Apel, 31 March 1999, NHCY; 1 spm., Socotra Island, Ras Qatanhin, 12°21.293'N/53°32.659'E, 8‑10 m,
under stones and coral rubble, coll. T. Wehe, 9 April 2000, SMF 13549.
M e a s u r e m e n t s : Holotype ZMB 18: cs (26) 33/10/8. Syntypes ZMB 490: cs (26) 14/6/5, cs (26) 11/4/3, cs (26)
13/5/4, cs (26) 18/6/5, cs (26) 13/5/4; cs (26) 22/6/5. Syntype ZMB 1079: cs (26) 15/5/3.5. Syntypes MPW 546: cs (26) 18/6/4,
cs (26) 9/5/3. Syntype USNM 074368: cs (26) 12/5/4. Syntypes MNHN POLY TYPE 1432: cs (26) 18/7/5, cs (26) 11/5/3, cs
(26) 14/3.5/2.5, cs (26) 17/5.5/4, cs (26) 15/6/4, cs (26) 13/6/4, cs (26) 17/5/3, cs (26) 16/6/4. BMNH 1941.4.4.190/4: cs (26)
16/7/5, cs (26) 13/5/4, cs (26) 10/4/3, cs (26) 13/5/4, cs (26) 9/4/3, cs (26) 6/3/2, cs (26) 10/3.5/2.5, cs (26) 11/5/3. SMF 13548:
cs (26) 9/4/3 [af (14) 5 + pf (12) 4], cs (26) 9/3.5/2.5 [af (12) 4 + pf (14) 5]. SMF 13550: cs (26) 12/5/3.5. NHMW V17058:
cs (26) 15/7/5, cs (26) 13/6/5, cs (26) 11/5.5/4, cs (26) 15/7/5, cs (26) 13/6/4, cs (26) 12/7/5, af (?) 5/4/3. MNHN A364: cs
(26) 10/4/3, cs (26) 12/4/3, cs (26) 12/4/3. MNHN A364: af (13) 8/5/3.5. MNHN A364: cs (26) 12/5/4. MNHN A21: cs (26)
25/10/8, cs (26) 17/6/4. MNHN A21: cs (26) 12/6/4. MNHN A21: cs (26) 15/6.5/5. MNHN A21: cs (26) 12/5/3.5. NHCY: cs
(26) 15/5/3. SMF 13549: cs (26) 14/4/3.5.
D i a g n o s i s : Without distinct nuchal fold, with two nuchal nodules. Elytra with two keels
and scattered, carinate microtubercles, without fringing papillae. Neurochaetae uni- and weakly
bidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 19 a), rounded to hexagonal,
about as wide as long, sometimes slightly withdrawn into following segment. Antennae with long,
smooth styles, without subterminal swelling, gradually tapering to terminal filum. Lateral antennae
slightly shorter than median antenna. Palps stout, conical, long, gradually tapering, often abruptly
terminating in very short pointed tip. Two pairs of eyes, anterior pair dorsolaterally on more or less
widest part of prostomium, posterior pair in front of hind margin. Everted pharynx terminally with
two semicircles of nine soft papillae and two pairs of chitinous jaws. Tentaculophores usually with
one chaeta. Tentacular cirri shaped as antennae, as long as or longer than median antenna, ventral
tentacular cirri slightly shorter than dorsal ones. Facial tubercle distinct. Buccal segment without
distinct nuchal fold, but with a pair of anteriorly inserted small nuchal nodules. Buccal cirri longer
and inserted more proximally than following ventral cirri, shaped as tentacular cirri. Elytra large,
covering dorsum completely. First pair rounded (Fig. 19 b), second reniform (Fig. 19 c), following
elytra oval (Fig. 19 d), outer margin often folded. Surface with more or less carinate microtubercles
(Fig. 19 e, 19 f ), slightly increasing in size from anterior to posterior, slightly more hemispherical
in posterolateral area. Two prominent keels running parallel to longitudinal body axis from at-
tachment scar region of elytrophore to posterior border (only rarely indistinct). Margin without
fringing papillae. Pigmentation, if it has not faded in ethanol, giving elytra a dark appearance, with
98 T. Wehe
the exception of a light spot above attachment of elytrophore. Dorsal cirri with large, globular to
cylindrical cirrophore and sometimes flattened, long, smooth style, gradually tapering to a terminal
filum. Dorsal tubercles more or less distinct, globular. Parapodia (Fig. 19 g, 19 h) with small, more
or less triangular notopodia. Neuropodia large, with rounded to subtriangular, prechaetal, acicular
lobe and bluntly rounded, postchaetal lobe of about similar length. Ventral cirri short, conical,
gradually tapering. Nephridial papillae from about segment 7 to 10 onwards. Notochaetae (Fig.
19 i) moderate in number, in small bundles, slightly curved, with numerous rows of spines and
blunt to pointed tips. Neurochaetae (Fig. 19 j, 19 k) numerous, much stouter, falcate, subdistally
slightly swollen with several rows of spines below slightly hooked, uni- or weakly bidentate tips.
Neurochaetae of second and third segment more slender, with more numerous rows of spines.
R e m a r k s : Even though the species is characterised by the unique feature of two distinct
keels on the elytra, it has been repeatedly described under several new names which were later syn-
onymised again (see Seidler 1923, Day 1957). Re-examination of the relevant types has confirmed
these synonymies. There are slight differences among specimens in some characters, i.a. in the dis-
tinctiveness of the nuchal fold and nuchal nodules. The tips of the neurochaetae appear to be vari-
able. In some specimens, a few bidentate neurochaetae with a small but distinct secondary tooth
are present, e.g. syntypes of L. quadricarinatus. However, numerous neurochaetae have bidentate
tips with a very small tooth only (see Fig. 19 k), or the tips are distinctly unidentate (Fig. 19 j).
Specimens reported from the Red Sea by Day (1965) and Kiseleva (1971) were not examined.
However, according to the description provided by Day and because of the unique elytra characters
exhibited in this species, there is no reason to doubt the identification by these authors.
The specimens reported by Fauvel (1955) and Fishelson & Rullier (1969) as L. hedleyi were
listed under L. purpureus in Wehe & Fiege (2002) according to synonymies given in Hartman
(1959) and Day (1967 a). However, specimens collected during the expedition of the Calypso and
described as L. hedleyi by Fauvel (1955) were all mis-identified and clearly belong to L. glaucus.
The specimen reported by Fishelson & Rullier (1969) could not be located and so the status of
this specimen remains doubtful.
B i o l o g y & e c o l o g y : In addition to the free-living forms, specimens were also found to
be living commensally within the tubes of species of the polychaete genus Eunice Cuvier, 1817
(Hanley & Burke 1990).
D i s t r i b u t i o n : Red Sea, Gulf of Aden, Arabian Sea: Socotra Archipelago. The species repre-
sents a new record for the Arabian Sea. Elsewhere: All over Indo-West Pacific (Seidler 1923, Day
1967, 1975, Hanley & Burke 1990).
Lepidonotus impatiens (Savigny in Lamarck, 1818) Fig. 20 a‑j

Polynoe impatiens Savigny in Lamarck, 1818. — Histoire naturelle des Animaux sans vertèbres…Vol. 5: 309; Savigny 1822,
Description de l’Égypte 1(3): 24; Savigny 1826, Description de l’Égypte. 2nd ed., Histoire naturelle. Minéralogie.-Zoologie,
Animaux Invertébrés 21: 350; Quatrefages 1866, Histoire naturelle des Annelés marins et d’eau douce. Annélides et Géphy-
riens: 221‑223 [partim, see L. cristatus].
Polynoe (Lepidonotus) striata Grube 1877. — Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin.
Aus dem Jahre 1877: 509 [not Kinberg, 1856].
Lepidonotus impatiens var. meridionalis Augener, 1913. — Die Fauna Südwest-Australiens. Ergebnisse der Hamburger südwest-
australischen Forschungsreise 1905 4(5): 103‑106 [new synonymy].
Lepidonotus australiensis Seidler, 1923. — Archiv für Naturgeschichte 89 (Abt. A/11): 49‑51, figs 9‑10 [new synonymy].
Fig. 20: Lepidonotus impatiens (lectotype, MNHN A28). a: anterior end; b: 7th right elytron; c: single nail-like tubercles in dorsola-
teral view; d: nail-like tubercles in dorsal view; e: uni- to trifid microtubercles; f: right parapodium from segment 14, anterior view;
g: same, posterior view; h: notochaeta; i: upper neurochaeta; j: lower neurochaeta.
100 T. Wehe
S p e c i m e n s e x a m i n e d : S y n t y p e : “Savigny, Mer Rouge, A1(R.) — 1868 — No16 b”, MNHN POLY TYPE 1433.
H o l o t y p e : Lepidonotus impatiens meridionalis, Koombana Bay, 6‑7 engl. miles S.W. of Bunburry, Australia, 14,
5‑18 m, coll. 28 July 1905 [collecting data supplemented according to Augener (1913)], ZMH V7887. H o l o t y p e :
Lepidonotus australiensis, Mermaidstreet, NW Australia, “Gazelle”-Expedition, ZMB 897.
M e a s u r e m e n t s : Syntype MNHN POLY TYPE 1433: cs (26) 26/10/8. Holotype ZMH V7887: cs (26) 22/11/8 [af (13)
10 + pf (13) 12]. Holotype ZMB 897: cs (26) 36/12/9.
D i a g n o s i s : With nuchal fold and nuchal nodules. Elytra with nail-like and uni- to trifid
tubercles, without fringing papillae. Neurochaetae unidentate.
R e d e s c r i p t i o n b a s e d o n s y n t y p e : Body compressed, especially parapodia, and
epidermis partially displaced. Body very stout, subrectangular in cross-section. Prostomium (Fig.
20 a) rounded, withdrawn into anterior segment, only anterior half visible. Ceratophores of anten-
nae stout, cylindrical. Style of median antenna broken off, lateral styles smooth, with subterminal
swelling and long terminal filum. Palps broken off. Eyes not visible. Pharynx only slightly everted,
no border papillae or jaws visible. Tentaculophores without chaetae. Tentacular cirri shaped as
lateral antennae, dorsal cirri longer than ventral cirri, both longer than lateral antennae. Facial
tubercle distinct. Buccal segment with prominent rectangular nuchal fold and a pair of anteriorly
directed nuchal nodules. Buccal cirri longer and inserted more proximally than following ventral
cirri, shaped as tentacular cirri. Elytra (Fig. 20 b) large, leaving parts of middorsum uncovered.
Elytra fleshy and strongly folded, first pair rounded to oval, following elytra broad and oval. Sur-
face very densely covered with nail-like tubercles (Fig. 20 c, 20 d), appearing as a velvet surface at
low magnification. In between nail-like tubercles numerous thorn-like, uni-, bi- and trifid tubercles
on posterolateral surface (Fig. 20 e). Margin smooth. Dorsal cirri with large, globular to cylindri-
cal cirrophore and long, smooth style with subterminal swelling and long, slender terminal filum.
Dorsal tubercles indistinct. Parapodia (Fig. 20 f, 20 g) with small, conical notopodia. Neuropodia
stout, short, diagonally truncate, with prechaetal, acicular lobe subtriangular and postchaetal lobe
more rounded, both of similar length. Ventral cirri stout, short, conical, gradually tapering. Ne-
phridial papillae from about segment 11 onwards. Notochaetae (Fig. 20 h) moderate in number,
only slightly curved, with dense rows of faint spines and blunt, unidentate tips. Neurochaetae (Fig.
20 i, 20 j) numerous, stouter, falcate, subdistally slightly swollen with several rows of spines, distal
much more prominent than basal spines, and slightly hooked, blunt, unidentate tips.
R e m a r k s : In the collection of the Muséum National d’Histoire Naturelle, Paris, two speci-
mens were located labelled as Polynoe impatiens and described in Quatrefages (1866). Accord-
ing to Quatrefages (p. 221), one of the specimens was collected by M. Botta, whereas the other
specimen was examined by Savigny: “Celui qui avait servi à Savigny …”, thus implying a possible
type status for the latter specimen. Since Savigny (1822, 1826) mentioned three specimens, this
specimen would represent a syntype. Re-examination of both specimens revealed that they belong
to two different species. The one collected by M. Botta on which Quatrefages (1866) based his
description clearly represents L. cristatus, as does another specimen reported as Polynoe (Lepidono-
tus) impatiens by Grube (1869).
Unfortunately, the description and illustrations of Polynoe impatiens provided by Savigny
(1822, 1826) are no longer sufficient, especially as regards the most important characters of the
elytra, to identify a specimen as belonging to this species. However, according to the statement
by Quatrefages (1866), the second specimen represents a syntype of L. impatiens. The species is
therefore regarded as valid and is here redescribed based on this syntype specimen.
Revision of both the type specimens of Lepidonotus impatiens var. meridionalis Augener,
1913 and Lepidonotus australiensis Seidler, 1923 showed that they are identical with L. impatiens,
and both are consequently regarded as junior synonyms.
D e p t h r a n g e : Unknown; elsewhere recorded between 14, 5‑18 m (Augener 1913).

D i s t r i b u t i o n : Red Sea. Elsewhere: Australia.
Lepidonotus natalensis Day, 1951 Fig. 21 a‑j

Lepidonotus natalensis Day, 1951. — Annals of the Natal Museum 12(1): 9‑11, fig. 1 e‑l [removed from synonymy with L.
tenuisetosus (sensu Day 1967)].
Lepidonotus tenuisetosus Fauvel 1927. — The Transactions of the Zoological Society of London 22/4(1): 414 [not Gravier,
1902].
Ty p e l o c a l i t y : Port Edward, Natal, South Africa, Indian Ocean.
S p e c i m e n s e x a m i n e d : Holotype: Lepidonotus natalensis, South Africa, Port Edward, Natal, coll. Day, BMNH
1961.16.1. — S u e z C a n a l : 1 spm., as L. tenuisetosus, Ballah Signal Station, between Port Said and Lake Timsah, “Cambridge
Expedition to the Suez Canal”, 27 November 1924 (according to a second label, identified as L. natalensis by Pettibone 1974) [col-
lecting data supplemented according to Fox (1926)], BMNH 1926.11.12.4; microscope slide, Ballah Signal Station, between Port
Said and Lake Timsah, coll. M. Fox, 27 November 1924 [collecting data supplemented according to Fox (1926)], IEA R/79’ (most
probably belonging to specimen BMNH 1926.11.12.4). — A r a b i a n S e a : 1 spm., as Lepidonotus jacksoni, Pakistan, Karachi,
Buleji, intertidal rock, coll. J. Mustaquim, 12 June 1997, SMF 13639.
M e a s u r e m e n t s : Holotype BMNH 1961.16.1: cs (26) 19/5/4. BMNH 1926.11.12.4: cs (26) 11/4/3. SMF 13639: cs
(26) 10/3/2.5.
D i a g n o s i s : Without nuchal fold. Elytra margin and surface with long slender, digitiform
papillae, surface with smooth or irregularly pointed microtubercles. Notochaetae partially thread-
like. Neurochaetae unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 21 a) rounded to slightly
hexagonal, about as wide as long. Styles of antennae broken off. Palps stout, conical, gradually ta-
pering to pointed tip. Two pairs of eyes, anterior pair on more or less widest part of prostomium,
posterior pair in front of hind margin. Tentaculophores with few chaetae. Tentacular cirri broken
off. Facial tubercle distinct. Buccal segment without nuchal fold. Buccal cirri longer and inserted
more proximally than following ventral cirri, shaped as tentacular cirri. Elytra (Fig. 21 b) large,
medially doomed. First pair rounded, second reniform, following elytra oval-like. Elytra surface
covered with simple, conical or somewhat irregularly pointed microtubercles (Fig. 21 c), and
numerous very long papillae on posterolateral surface (Fig. 21 d). Margin posterolaterally with
a dense fringe of long, slender papillae (Fig. 21 e), and small clavate sensory papillae (Fig. 21 f ).
Dorsal cirri with large, globular to conical cirrophore, styles without distinct subterminal swelling.
Dorsal tubercles indistinct. Parapodia with small, more-or-less triangular notopodia. Neuropo-
dia (Fig. 21 g, 21 h) with subtriangular, prechaetal, acicular lobe, slightly longer than rounded,
postchaetal one. Ventral cirri short, slender, gradually tapering. Nephridial papillae from segment
7 onwards. Notochaetae (Fig. 21 i) slender, short to long, partially thread-like, with fine, pointed
tips. Neurochaetae (Fig. 21 j) stouter than notochaetae, falcate, subdistally slightly thickened, with
few rows of spines below slightly bent, blunt, unidentate tips.
R e m a r k s : The specimen BMNH 1926.11.12.4, originally described as Lepidonotus tenuise-
tosus by Fauvel (1927), is here identified as L. natalensis (as it was by Pettibone, see above), after
comparison with the holotype. It agrees very well with the latter in the main characters, i.a. ar-
rangement of eyes, long and slender papillae on the elytra surface, elytra tubercles, fine, thread-like
notochaetae, and unidentate neurochaetae. A loose elytron with warty microtubercles is considered
to belong to another species.
Day (1967 a) synonymised L. natalensis which he himself described (Day 1951) with L. tenui-
setosus. This synonymy is difficult to understand, since both species clearly differ in several respects,
most of all in the arrangement of the eyes and the presence or absence, respectively, of long, slender
papillae on the elytra surface. L. natalensis is therefore removed from the synonymy with L. tenui-
setosus.
102 T. Wehe
Fig. 21: Lepidonotus natalensis (BMNH 1926.11.12.4: a, b, g-j; SMF 13639: c-f). a: anterior end; b: left elytron from midbody; c:
microtubercles; d: papilla on elytra surface; e: fringing papillae; f: sensory papillae on elytra margin; g: parapodium from midbody
segment, anterior view; h: same, posterior view; i: notochaetae; j: neurochaetae.
D i s t r i b u t i o n : Suez Canal, Arabian Sea: Karachi. This is the first record of the species from
the Suez Canal and the Arabian Sea. Elsewhere: South Africa (Day 1951).
Lepidonotus polae n. sp. Fig. 22 a‑j

Lepidonotus cristatus. — ? Fishelson & Rullier 1969, Israel Journal of Zoology 18: 53; ? Amoureux et al. 1978, Israel Journal
of Zoology 27: 62 [not Grube, 1876].
S p e c i m e n s e x a m i n e d : Holotype: Tor (= At-Tur), Sinai Peninsula, Gulf of Suez, Red Sea, Egypt, “Pola Expedition”,
1895‑1898, NHMW 17059. — R e d S e a : Microscope slide with single elytron and parapodium, as Lepidonotus cristatus, Mer
Rouge, 4266, IEA Y100’ (most probably belonging to one of the specimens reported as L. cristatus by Fishelson & Rullier (1969)
and Amoureux et al. (1978) according to the type of label).
M e a s u r e m e n t s : cs (26) 14/8/6.
D i a g n o s i s : With nuchal fold and nuchal nodules. Elytra with multifid microtubercles and
ornamented cone-like macrotubercles, without fringing papillae. Neurochaetae unidentate.
D e s c r i p t i o n : Body subrectangular in cross-section. Prostomium (Fig. 22 a) rounded to
hexagonal, withdrawn into following segment, posterior part not visible. Antennae with long,
smooth styles with subterminal swelling and slender terminal filum, lateral styles shorter than
median style. Palps stout, conical, long, gradually tapering, terminating in short pointed tip. Eyes
not visible. Tentaculophores with single chaeta. Tentacular cirri shaped as antennae, about as long
as median antenna, dorsal slightly longer than ventral cirri. Facial tubercle distinct. Buccal seg-
ment with slightly rectangularly raised nuchal fold with a pair of small, anteriorly directed nuchal
nodules. Buccal cirri longer and inserted more proximally than following ventral cirri, shaped as
tentacular cirri. Anteriormost pairs of elytra missing, following elytra completely covering dorsum.
Elytra (Fig. 22 b) large, more-or-less oval, soft, strongly folded. Surface densely covered with small,
uni- to multifid (Fig. 22 c) and nail-like microtubercles (Fig. 22 d); in central area and along pos-
terior margin with few, ornamented cone-like macrotubercles (Fig. 22 b, 22 e). Margin without
fringing papillae. Dorsal cirri with large, globular to cylindrical cirrophore and long, smooth style,
subterminal with slight swelling and long, slender terminal filum. Dorsal tubercles inconspicuous.
Parapodia (Fig. 22 f, 22 g) with small, more or less conical notopodia. Neuropodia with triangular,
prechaetal, acicular lobe and bluntly rounded, postchaetal lobe of similar length. Ventral cirri short,
conical, gradually tapering. Nephridial papillae from segment 7 onwards. Notochaetae (Fig. 22 h)
moderate in number, slightly curved, with numerous rows of fine spines and blunt, unidentate
tips. Neurochaetae (Fig. 22 i, 22 j) stouter, falcate, subdistally only slightly swollen, with several
rows of spines, distal longer than basal spines, and with slightly hooked, blunt, unidentate tips.
R e m a r k s : Seidler (1923), who reviewed the lepidonotid genera, listed more than 50 species
within the genus Lepidonotus. Since then, at least 20 to 30 additional species have been described.
Unfortunately, many of the descriptions are insufficient for distinguishing species, and this is also
true for the descriptions or mere translations of original descriptions provided by Seidler (1923).
No description was found corresponding to the characters exhibited by the specimen examined,
and it is therefore described here as new. Although there are no characters which are not to be
found among other species, the combination of characters is unique: a nuchal fold with nuchal
nodules, the type of spinous microtubercles and ornamented macrotubercles, elytra margins with-
out fringing papillae, and neurochaetae with unidentate tips. The specimen seems to be close to
L. permixturus Hanley & Burke, 1991 a. However, no ornamented, conical macrotubercles were
described from the four specimens examined by these authors.
E t y m o l o g y : The species name was chosen in recognition of the first important deep-sea expedi-
tion to the Red Sea in the years 1895‑1896 and 1897‑1898 by the Austrian Research Vessel “Pola”.
104 T. Wehe
Fig. 22: Lepidonotus polae n. sp. (holotype, NHMW 17059). a: anterior end; b: 11th left elytron; c: microtubercles; d: nail-like
tubercles e: macrotubercle; f: left parapodium from segment 10, anterior view; g: same, posterior view; h: notochaeta; i: upper
neurochaeta; j: lower neurochaeta.
D i s t r i b u t i o n : Red Sea: Gulf of Suez.
Lepidonotus purpureus Potts, 1910 Fig. 23 a‑i, Pl. 2 a

Lepidonotus purpureus Potts, 1910. — The Transactions of the Linnean Society Series 2, Zoology 8: 334, pl. 18: fig. 3; Seidler
1923, Archiv für Naturgeschichte 89 (Abt. A/11): 79; Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences
12(1): 66‑70, fig. 11.
Ty p e l o c a l i t y : East Africa.
S p e c i m e n s e x a m i n e d : 1 Syntype: Lepidonotus purpureus, Wasin, British East Africa (= Kenya) [collected at Zanzibar
according to Potts (1910)], coll. C. Crossland, 1902, BMNH 1924.3.1.104. — R e d S e a : 1 spm., as Lepidonotus carina-
tus, bay between Gt. Hanish and Suyul Hanish Islands, 13°39'30''N/42°43'00''E, 29 m, dredge, coll. “John Murray Expedi-
tion”, 17 September 1933, BMNH 1937.9.2.56; 1 spm., as Lepidonotus carinatus, bay between Gt. Hanish and Suyul Hanish
Islands, 13°39'30''N/42°43'00''E, 26 m, dredge, coll. “John Murray Expedition”, 17 September 1933, BMNH 1937.9.2.57;
1 spm., Eritrea, Assab Bay, Sanabor Island, “R/V Akademik Petrovsky”, cruise 9, 2, A.B. Tzetlin, coll. 14 February 1980,
ZMUM. — A r a b i a n S e a : 1 spm., as Lepidonotus carinatus, south coast of Oman, 19°22'36''N/57°53'00''E, 13.5 m, coll.
“John Murray Expedition”, 2 November 1933, BMNH 1937.9.2.55; 3 spms., Buleji, Karachi, Pakistan, 12 July 1997, coll. J.
Mustaquim, SMF 13546; 1 spm., Socotra Archipelago, Abd al-Kuri Island, W‑coast, “Trident Bay”, in prime channel of the
“trident”, 12°13.457'N/52°4.263'E, 10‑20 m, coll. M. Apel, 4 April 2000, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay,
12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E, 18‑20 m, dredge, corals and algae on bedrock, coll. T. Wehe, 9 April
2000, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21'17''N/53°32'07''E to 12°21'78''N/53°32'39''E, 18‑20 m,
dredge, corals and algae on bedrock, coll. T. Wehe, 9 April 2000, SMF 13543; 1 spm., Socotra Island, Ras Qataninh Bay,
12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll. T. Wehe, 9 April 2000, SMF 13542; 1 spm., Socotra Is-
land, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll. T. Wehe, 9 April 2000, NHCY;
1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll. T. Wehe, 9 April
2000, NHCY; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑11 m, under stones and coral rubble, coll.
T. Wehe, 9 April 2000, SMF 13544; 1 spm., Socotra Island, Ras Qataninh Bay, 12°21.293'N/53°32.659'E, 8‑11 m, under stones
and coral rubble, coll. T. Wehe, 9 April 2000, SMF 13545.
M e a s u r e m e n t s : Syntype: cs (26) 21/8/6. BMNH 1937.9.2.56: cs (26) 20/6/4. BMNH 1936.9.2.57: cs (26) 12/4/3.
ZMUM: cs (26) 20/6.5/5 [af (12) 8 + pf (14) 12. BMNH 1937.9.2.55: cs (15 + small regenerated segments) 7/3/2.5. SMF 13546:
cs (26) 13/5.5/4, cs (26) 16/6/4, cs (26) 12/6/4. NHCY: cs (26) 15/6/4.5. NHCY: cs (26) 20/5.5/4.5 [af (19) 14 + pf (7) 6]. SMF
13543: cs (26) 14/5/4. SMF 13542: cs (26) 14/5/4. NHCY: cs (18, posterior end regenerated) 5/3/2. NHCY: cs (26) 11/3.5/2.5.
SMF 13544: cs (26) 13/4/3. SMF 13545: cs (26) 13/4.5/3.
D i a g n o s i s : With nuchal fold and nuchal nodules. Elytra with clusters of unpigmented cells
and microtubercles, without fringing papillae. Neurochaetae bidentate.
hexagonal, about as wide as long, sometimes withdrawn into anterior segments. Antennae with
long, smooth styles, with slight subterminal swelling and terminal filum. Styles of lateral anten-
nae slightly shorter than median antenna. Palps stout, long, gradually tapering. Two pairs of eyes,
hind margin, sometimes hidden if prostomium withdrawn. Everted pharynx with two semicircles
of nine soft papillae distally and two pairs of chitinous jaws. Tentaculophores with one chaeta each.
Tentacular cirri shaped as antennae, dorsal longer than ventral cirri, both as long as or longer than
median antenna. Facial tubercle distinct. Buccal segment with nuchal fold, often rectangularly
raised, with two distinct nuchal nods on anterior edge. Buccal cirri longer and inserted more proxi-
mally than following ventral cirri, shaped as tentacular cirri. Elytra (Fig. 23 b) large, covering dor-
sum. First pair rounded, second reniform, following elytra oval-like. Surface in anterior part with
globular microtubercles (Fig. 23 c), sometimes appearing flattened, caldera-like, eventually absent
on posteriormost elytra; typically clusters of about 4 to 8 unpigmented cells (Fig. 23 d) spread over
surface, even recognisable under compound microscope if reddish-brown pigmentation of elytra
has faded. Posterior elytra often with two indistinct folds from central part to posterior edge, but
no distinct keels as in L. glaucus. Margin without fringing papillae. Dorsal cirri with large, globular
106 T. Wehe
Fig. 23: Lepidonotus purpureus (SMF 13546). a: anterior end; b: 3rd right elytron; c: microtubercles; d: clusters of unpigmented cells; e:
right parapodium from segment 14, anterior view; f: same, posterior view; g: notochaeta; h: middle neurochaeta; i: lower neurochaeta.
to conical cirrophore and long, smooth style, sometimes flattened dorsoventrally, with slight sub-
terminal swelling below terminal filum. Dorsal tubercles distinct, globular. Parapodia (Fig. 23 e,
23 f ) with small, more or less triangular notopodia. Neuropodia large, with rounded, prechaetal,
acicular lobe and more bluntly rounded, slightly shorter, postchaetal lobe. Ventral cirri short, coni-
cal, gradually tapering. Nephridial papillae usually from about segment 8 onwards. Notochaetae
(Fig. 23 g) moderate in number, slightly thinner than neurochaetae, finely spinous, with pointed
to capillary tips. Neurochaetae (Fig. 23 h, 23 i) stouter, falcate, subdistally with slight swelling and
several rows of spines below slightly hooked, bidentate tips. Secondary tooth often worn away,
leaving a small subterminal scar. Few unidentate neurochaetae. Neurochaetae of second segment
slightly modified, more slender, with more numerous rows of spines.
R e m a r k s : Following Hartman (1959), Wehe & Fiege (2002) listed the records of L. hedleyi
by Fauvel (1955) and Fishelson & Rullier (1969) as synonyms of L. purpureus. A re-examina-
tion, however, revealed the specimens in question belong to L. glaucus (see above).
D e p t h r a n g e : So far recorded between 8 and 29 m; elsewhere reported from the inter-
tidal.
D i s t r i b u t i o n : Red Sea, Arabian Sea: Socotra Archipelago and Karachi. The specimens
listed above represent the first record of the species from the Red Sea. Elsewhere: East Africa,
Madagascar, Australia, New Zealand (Knox 1960, Day 1967).
Lepidonotus tenuisetosus (Gravier, 1902) Fig. 24 a‑l

Euphione tenuisetosa Gravier, 1902. — Nouvelles Archives du Muséum 4 e série 3: 222‑226, textfigs 228‑231, pl. 8: figs 123‑126;
Fauvel 1911, Archives de Zoologie Expérimentale et Générale, Cinquième Série 4(11): 368.
Lepidonotus tenuisetosus. — Seidler 1923, Archiv für Naturgeschichte 89 (Abt. A/11): 25‑27; Mohammad 1971, Journal of
Zoology 163: 288; Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 70‑72, fig. 12; Barnich et
al. 2004, Species Diversity 9: 300‑301.
Ty p e l o c a l i t y : Djibouti, Gulf of Tadjoura, Gulf of Aden.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Euphione tenuisetosa, Djibouti, Gulf of Aden, coll. H. Coutière, 01 January
1897, MNHN POLY TYPE 263. — R e d S e a : 3 spms., Sherm Habban [near to Al Wajh], Saudi Arabia, “Pola Expedition”,
1895‑1898 [collecting data supplemented according to Schefbeck (1991)], NHMW 17062; 1 spm. [no label in vial, but ac-
cording to note on larger collection jar in the NMHN from the Red Sea], MNHN A480. — A r a b i a n S e a : 1 spm., Socotra
Island, 12°41.151'N/53°28.625'E, intertidal, coll. M. Apel, 11 April 1999, SMF 13558; 1 spm., Buleji, Karachi, Pakistan, inter-
tidal rock, 22 December 2001, coll. J. Mustaquim, SMF 13561. — G u l f o f O m a n : 1 spm., Mina al Fahl, north of Muscat,
Oman, coll. Mai 1984, ZMUC POL-1360; 2 spms., Mina al Fahl, north of Muscat, Oman, coll. Mai 1984, ZMUC POL-1361.
— T h e G u l f : 11 spms., Kuwait, 29°20' to 29°22'N/48°02' to 48°06'E, stratified calcareous sand stone and light to dark grey
sand (with very little mud), coll. M.-B.M. Mohammad, 23 November 1967‑28 November 1968 [collecting data supplemented ac-
cording to Mohammad (1971)], BMNH 1969.188; 1 spm. + microscope slide, JMWS project, Jubail, 27°05'12''N/49°35'04''E,
sand, coll. D. Fiege, 29 November 1991, SMF 13559; 1 spm. + microscope slide, JMWS project, Ras Az-Zwar - Marduma Bay
region, 27°22'26''N/49°12'58''E, intertidal, sand-rock, coll. D. Fiege, 28 November 1991, SMF 13556; 1 spm., JMWS project,
Ras Az-Zwar - Marduma Bay region, 27°25'41''N/49°13'53''E, intertidal, rock flat, coll. D. Fiege, 7 December 1991, SMF 13557;
1 spm., JMWS project, Jub. Kas. Arc, sandbank, coll. D. Fiege, 9 December 1991, SMF 13555; 2 spms. + microscope slide, JMWS
project, Ras Az-Zawr - Marduma Bay region, 27°25'12''N/49°11'17''E, intertidal, rock-sand, coll. D. Fiege, 9 December 1991,
SMF 13560; 1 spm., Abu Dhabi, Jazirat Dagalah, 24°12.327'N/52°56.113'E, coral reef with corals mainly dead, 2‑5 m, coll. J.D.
George, 19 November 1999, Coll. George; 1 spm., Abu Dhabi City, Sheraton Hotel pontoon, coll. J.D. George, 14 April 1999,
Coll. George; 1 spm., Abu Dhabi, off Ruwais & Sir Bani Yas, 24°13.520'N/52°43.815'E, from floating mooring rope, coll. J.D.
George, 16 November 1999, Coll. George; 1 spm., Abu Dhabi, Baya Sila harbour, intertidal, under boulder collection, coll. J.D.
George, 14 November 1997, Coll. George; 2 spms., Abu Dhabi, Suwaihal, W of Jebel Dhanna, coll. J.D. George, 14 March 1996,
Coll. George; 4 spms., Abu Dhabi, Dabb’iya, 24°15.762'N/54°06.113'E, ADCO jetty with fouling organisms, 1‑5 m, coll. J.D.
George, 21 November 1999, Coll. George; microscope slide, as Euphione tenuisetosa, Bouchir (=Būshehr), coll. Bogayawlenski,
March 1902, IEA P.50; microscope slide, as Euphione tenuisetosa, Bouchir (=Būshehr), coll. M.N. Bogayawlenski, March 1902, IEA
P.53.
M e a s u r e m e n t s : Holotype MNHN POLY TYPE 263: cs (26) 10/6/4. NHMW 17062: cs (26) 9/4.5/3, cs (26) 9/4/3,
cs (21, posterior end regenerated) 6/4.5/3. MNHN A480: cs (26) 15/4.5/3.5. SMF 13558: cs (26) 15/5.5/4.5 [af (13) 7 + pf
(13) 8]. SMF 13561: cs (26) 20/6/4. BMNH 1969.188: cs (26) 11/4/3, cs (26) 15/5/4, cs (26) 10/3/2, cs (26) 10/3, 5/3, cs (26)
15/5.5/4, cs (26) 10/4/3, cs (26) 9/3.5/3, cs (26) 10/4/3, cs (26) 9/3/2.5, cs (26) 10/4/3, cs (26) 11/5/4. ZMUC POL-1360: cs
(26) 5/1.5/1. ZMUC POL-1361: pf (8) 4/4/2.5, pf (13) 9/3/2. SMF 13559: cs (26) 14/4.5/3.5 [af (17) 9 + pf (9) 5]. SMF 13556:
108 T. Wehe
Fig. 24: Lepidonotus tenuisetosus (BMNH 1969.188). a: anterior end; b: 1st right elytron; c: 2nd right elytron; d: microtubercles; e:
macrotubercles; f: fringing papillae; g: globular papillae on elytra margin; h: left parapodium from segment 14, posterior view; i:
same, anterior view; j: notochaeta; k: neurochaetae; l: neurochaeta from segment 2.
cs (26) 15/5/4. SMF 13557: cs (26) 15/5/4. SMF 13555: cs (26) 10/3.5/2.5. SMF 13560: cs (26) 16/5.5/4.5, cs (26) 13/5/4. Coll.
George: af (12) 5/4/3. Coll. George: af (16) 5/2.2/2. Coll. George: cs (26) 7/2.5/2 [af (15) 4 + pf (11) 3]. Coll. George: cs (26)
18/6/5 [af (14) 9 + pf (12) 9]. Coll. George: cs (26) 17/5/4, cs (26) 11/3/2.7. Coll. George: cs (26) 12/4/3.5, cs (26) 5/2/1.9, af
(13) 5/3/2.5, af (12) 2.5/1.8/1.7.
D i a g n o s i s : Both pairs of eyes dorsolaterally on prostomium. Tentaculophores achaetous.

Without nuchal fold. Elytra with smooth micro- and macrotubercles, with fringing papillae. Neu-
rochaetae unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 24 a) rounded to oval,
about as wide as long, sometimes withdrawn into following segment. Antennae with long, smooth
styles, gradually tapering to terminal filum, without subterminal swelling. Styles of lateral anten-
nae shorter than median one. Palps stout, conical, long, gradually tapering, sometimes abruptly
terminating in a short tip. Two pairs of eyes close to each other dorsolaterally on prostomium.
Tentaculophores achaetous. Tentacular cirri shaped as antennae, dorsal longer than ventral cirri,
both pairs about as long as or longer than median antenna. Facial tubercle distinct. Buccal segment
without nuchal fold. Buccal cirri longer and inserted more proximally than following ventral cirri,
shaped as tentacular cirri. First pair of elytra rounded (Fig. 24 b), second reniform (Fig. 24 c), fol-
lowing elytra oval-like. Surface with conical to globular microtubercles (Fig. 24 d), slightly increas-
ing in size from anterior to central and posterior surface. Some macrotubercles (Fig. 24 e) centrally
and posteriorly with blunt or somewhat warty tips; macrotubercles largest on first pair of elytra.
Margin with fringe of digitiform (Fig. 24 f ) and partially with minute globular papillae (Fig. 24 g).
Elytra irregularly pigmented and with dark spot above attachment of elytrophore if colour has not
faded. Dorsal cirri with large, globular to cylindrical cirrophore and long, gradually tapering style,
without subterminal swelling below terminal filum. Dorsal tubercles more or less distinct, flattened
to globular. Parapodia (Fig. 24 h, 24 i) with small, almost triangular notopodia. Neuropodia with
bluntly rounded to subtriangular, prechaetal, acicular lobe and bluntly rounded, postchaetal lobe
of similar length. Ventral cirri short, conical, gradually tapering. Nephridial papillae from about
segment 8 onwards. Notochaetae (Fig. 24 j) moderate in number, slender, with rows of spines and
fine tips. Neurochaetae (Fig. 24 k) stouter, falcate, subdistally slightly thickened, with few rows
of spines, below blunt, unidentate tips. Neurochaetae of second segment altered (Fig. 24 l), more
slender, with more numerous rows of spines and fine tips.
R e m a r k s : A peculiar character only found in a few species of Lepidonotus is that both pairs
of eyes are close to each other, situated dorsolaterally on the prostomium. The conical macrotu-
bercles are distinct only in larger specimens and might be absent in smaller ones, as in the juvenile
specimens from the Gulf of Oman. The specimen reported by Fauvel (1927) belongs to L. na-
talensis (see above).
D e p t h r a n g e : So far recorded down to 5 m; elsewhere reported down to 27 m (Barnich &
Fiege 2003).
D i s t r i b u t i o n : Red Sea, Gulf of Aden, Arabian Sea: Socotra Archipelago and Karachi, Gulf
of Oman, The Gulf. Elsewhere: India, Vietnam, South China Sea, New Caledonia, Eastern Medi-
terranean Sea (Uschakov 1982, Hanley 1992, Barnich & Fiege 2003, Barnich et al. 2004).
Parahalosydnopsis Pettibone, 1977

Parahalosydnopsis Pettibone, 1977. — Essays on polychaetous annelids in Memory of Dr. Olga Hartman: 44‑45.
Ty p e s p e c i e s : Parahalosydnopsis hartmanae Pettibone, 1977.
D i a g n o s i s (emended): Body elongate with up to 75 segments, flattened dorsoventrally,
subrectangular in cross-section. Ceratophores of lateral antennae inserted terminally. Styles of
antennae with subterminal swelling; style of median antenna much longer, with few papillae sub-
distally. Palps stout, conical, gradually tapering. Two pairs of eyes present. Tentaculophores with
chaetae. Second (buccal) segment without nuchal fold. Elytra numerous pairs on segments 2, 4, 5,
7, alternating to 25, 28, 30, alternating to 42, 43, 45, 47, 48, 50, alternating to end of body. Elytra
110 T. Wehe
Plate 1: a: Iphione ovata, preserved specimen, SMF 13572, Port Sudan (length 12 mm, width 7 mm); b, c: Hyperhalosydna striata,
preserved specimen, SMF 13553, Socotra Island (length 37 mm, width 6 mm); d: Hermenia verruculosa, preserved specimen, SMF
13608, Socotra Island (length 13 mm, width 8 mm); e: Parahalosydnopsis tubicola, preserved specimen, anterior fragment, SMF
13619, The Gulf (length 20 mm, width 4 mm); f: Paralepidonotus ampulliferus, SMF 13599, Karachi (length 30 mm, 12 mm).
covering dorsum; surface with scattered microtubercles, margins with or without fringing papillae.
Dorsal cirri with smooth styles and subterminal swelling. Notopodia very small, inconspicuous.
Neuropodial lobes of similar length, with papillae on distal borders. Notochaetae moderate in
number, very fine, thread-like, faintly spinous. Neurochaetae much stouter, with few pairs of scale-
like spines subterminally and unidentate tips.
Plate 2: a: Lepidonotus purpureus, preserved specimen, SMF 13546, Karachi (length 13 mm, width 5.5 mm); b: Lepidonotus crista-
tus, preserved specimen, SMF 13674, Socotra Island (length 35 mm, width 17 mm); c: elytron from same; d: Lepidonotus glaucus,
preserved specimen, dorsal view of elytra, SMF 13549, Socotra Island (length 14 mm, width 4 mm).
R e m a r k s : The diagnosis given by Pettibone (1977) was emended for the number of possi-
ble segments and a papillate rather than a smooth style of the median antenna. Specimens belong-
ing to this genus with more than 50 segments can easily be recognized by the unique sequence of
112 T. Wehe
the elytra along the body. Only one of about 50 specimens examined show an irregularity in this
arrangement.
Key to the species of Parahalosydnopsis
1 Anterior elytra with fringing papillae. With ampulla-like papillae on elytra surfaces
P. arabica
– Elytra margin smooth. Without ampulla-like papillae P. tubicola
Parahalosydnopsis arabica n. sp. Fig. 25 a‑m

Hyperhalosydna striata Wesenberg-Lund 1949. — Danish scientific investigations in Iran 4: 257 [not Kinberg, 1856].
S p e c i m e n s e x a m i n e d : H o l o t y p e : S u d a n : Red Sea, coll. C. Crossland, October 1904 - May 1905, BMNH
2003.73. 16 P a r a t y p e s : same data as holotype, BMNH 2003.74‑83. 2 P a r a t y p e s : same data as holotype, SMF 12312,
SMF 12313. — R e d S e a : 13 spms., Sudan, coll. C. Crossland, October 1904 - May 1905, BMNH 2003.319‑328. — T h e
G u l f : 1 spm. + microscope slide, off eastern side of Kharg Island, north of Bouchir (=Būshehr), 29°00'N/50°01'E, seine, 12.5 m,
shell, sand, stones, corals, coll. G. Thorson, 6 March 1937, ZMUC POL-1342; 1 spm., Abu Ali Island, near Jubail, Kingdom of
Saudi-Arabia, subtidal grass, coll. 7 March 1999, CEWRD. 1 spm., 52°34'39.2'N/24°42'31.5''E, dive-operated scoop, coll. M.
Stachowitsch & R. Kikinger, SMF 13662; 1 spm., 54°9'16.9'N/24°34'30.6''E, dive-operated scoop, coll. M. Stachowitsch & R.
Kikinger, SMF 13663.
M e a s u r e m e n t s : Holotype BMNH 2003.73: cs (59) 20/3/2.5. Paratypes BMNH 2003.74‑83: cs (59) 19/3/2, cs (61)
21/3/2.5, cs (58) 18/3/2.5, cs (60) 22/4.5/3.5, cs (46) 18/3/2.5, cs (44) 13/3/2.5, cs (61) 19/3.2/2.5, cs (62) 24/3/2.5, cs (50)
14/3/2.5, cs (62) 19/3/2.5, cs (65) 23/3/2.5, cs (61) 32/3.5/2.5, cs (61) 25/4/3, cs (52) 12/3/2.5, cs (47) 14/3/2 cs (53) 16/3/2.5.
Paratype SMF 12312: cs (66) 25/3/2.5. Paratype SMF 12313: cs (62) 23/3.5/2.5. BMNH 2003.319‑328: af (40) 16/4/3, af (45)
19/3.5/3, af (34) 14/4/3, af (44) 16/3/2.5, af (38) 15/3/2.5, af (45) 16/4.5/3.5, af (19) 6/3/2.5, af (19) 6/3/2.5, af (24) 8/3.5/3, af
(19) 7/4/3, af (21) 9/4/3, af (15) 4/2.5/2, af (15) 5/3/2.5. ZMUC POL-1342: af (36) 15/3/2.5. CEWRD: cs (68) 33/3.5/3.2. SMF
13662: af (21) 5/2/1.6. SMF 13663: af (36) 6/1.7/1.5.
D i a g n o s i s : First pair of elytra with distinct fringe of papillae, following anterior ones with
few fringing papillae, median and posterior elytra without. Surface with irregularly ridged or
pointed microtubercles and balloon-like papillae. Neurochaetae with 2 to 7 pairs of broad flat-
tened spines subdistally.
D e s c r i p t i o n : Holotype a complete specimen about 20 mm length, 2, 5 or 3 mm (includ-
ing chaetae) width and 59 segments with minute last segments. Segment number amongst para-
types ranges from 44 to 66. Body oblong, flattened dorsoventrally. Prostomium (Fig. 25 a) round-
ed. Styles of antennae with subterminal swelling and long, slender terminal filum. Style of median
antenna (broken off on holotype) much longer than styles of lateral antennae, with papillae on
subterminal swelling. Palps very long, stout, conical, with numerous minute papillae, gradually ta-
pering, terminating more or less abruptly to fine tips. Two pairs of eyes, anterior pair dorsolaterally
on more or less widest part of prostomium, posterior pair in front of hind margin. Tentaculophores
with one or two chaetae. Tentacular cirri with subterminal swelling without papillae and terminal
filum, dorsal slightly longer than ventral cirri, both as long as or shorter than median antenna.
Facial tubercle distinct. Buccal segment with buccal cirri longer and inserted more proximally than
following ventral cirri, shaped as tentacular cirri. Elytra numerous, depending on segment number
(holotype with 30 pairs), covering dorsum completely. First pair rounded (Fig. 25 b), second pair
slightly reniform to oval (Fig. 25 c), following elytra larger, oval (Fig. 25 d), becoming smaller
again on posteriormost segments. Elytra attached off centre on elytrophores. First pair with dis-
tinct fringe of short, digitiform papillae on anterolateral margin (Fig. 25 b, 25 h), and very short
papillae on medial margin; surface on central and outer parts with numerous irregularly ridged or
pointed microtubercles, central and lateral larger than anterior papillae. Scattered small nodular
Fig. 25: Parahalosydnopsis arabica n. sp. (holotype, BMNH 2003.73). a: anterior end; b: 1st right elytron; c: 2nd left elytron; d: 13th
left elytron e: microtubercles of central elytra surface from c; f: microtubercles of lateral elytra surface from c; g: balloon-shaped
papilla; h: fringing papillae; i: right parapodium from segment 10, anterior view; j: same, posterior view; k: notochaeta; l: upper
neurochaeta m: lower neurochaeta.
114 T. Wehe
or digitiform papillae present between microtubercles. Second pair and following anterior pairs of
elytra with only few, digitiform fringing papillae on lateral margin and small knob-like papillae on
inner margin, ridged or pointed microtubercles more concentrated above region of elytrophore and
submarginal surface (Fig. 25 e, 25 f ). In between slender, balloon-shaped or ampulla-like papillae
(Fig. 25 g). On more posterior elytra, fringing papillae disappearing, number of microtubercles
and papillae decreasing and concentrated on central and submarginal areas, leaving most areas of
surface smooth. Dorsal cirri with large, globular to cylindrical cirrophore and long, smooth style
with subterminal swelling and terminal filum. Dorsal tubercles small, flattened or inconspicuous.
Parapodia (Fig. 25 i, 25 j) with vestigial notopodia. Neuropodia diagonally truncate distally; pre-
chaetal, acicular lobe slightly longer than postchaetal one, both with digitiform to globular papillae
on distal borders. Neuropodia and ventral body surface more or less distinct with scattered minute
papillae. Ventral cirri short, gradually tapering. Nephridial papillae distinct from about segment
8 onwards, small knob-like on first few segments, following papillae very prominent, cylindrical.
Notochaetae (Fig. 25 k) moderate in number, delicate, thread-like, finely spinous, tapering to cap-
illary tips, often thickly covered by inorganic matter. Neurochaetae (Fig. 25 l, 25 m) much stouter,
falcate, subdistally with slight swelling, with 2 to 7 pairs of broad flattened spines below slightly
curved, blunt, unidentate tips. Pygidium with a pair of anal cirri.
R e m a r k s : After preservation the colour faded, and if any remains the elytra are irregularly
brown pigmented and possess a small dark spot above the attachment scar of the elytrophore. On
some specimens, papillae on the median antennae are indistinct. Also the minute papillae on the
ventral body surface and parapodia could not be observed on every specimen. The elytra arrangement
is very constant, and only one specimen shows an irregularity from the general pattern on one side of
the body posterior to segments 42/43, BMNH 2003.74‑83. Another specimen exhibits a dorsal cirrus
on segment 4 instead of an elytron, but this is not a variation in the arrangement of the elytra within
this genus or species but rather a clear anomaly from the general pattern found in Polynoidae.
Two species, P. tubicola (Day, 1973) and P. hartmanae Pettibone, 1977, were previously known.
Pettibone (1977) described or redescribed, respectively, both of them and provided a key. P. arabica
n. sp. is somewhat intermediate between these two species and is therefore described as new. It differs
from P. tubicola by the presence of fringing papillae on the elytra, and differs from P. hartmanae by
the number of spines on the neurochaetae and the much less well developed fringing papillae.
Although incomplete and in bad condition, the specimen described as Hyperhalosydna striata
by Wesenberg-Lund (1949), clearly belongs to this species.
B i o l o g y & e c o l o g y : P. tubicola is known to occur as a commensal in tubes of terebel-
lid polychaetes, and this might also be true for P. arabica n. sp. The specimens BMNH 2003.73,
BMNH 2003.74‑83, BMNH 2003.319‑328, SMF 12312, formerly together in a single vial with
other unidentified specimens and obviously sorted out previously as scale worms from a larger
sample, show remnants of what appears to be terebellid tentacles attached to their bodies. In the
vial of the CEWRD specimen there are parts of a terebellid-like tube together with the specimen.
A commensal life pattern is therefore assumed.
D e p t h r a n g e : 12.5 m.
D i s t r i b u t i o n : Red Sea, The Gulf.
E t y m o l o g y : The species is named for its occurrence on the Arabian Peninsula.
Parahalosydnopsis tubicola (Day, 1973) Fig. 26 a‑k, Pl. 1 e

Lepidasthenia tubicola Day, 1973. — Journal of the Linnean Society of London 52: 341‑343, fig. 2 A‑E.
Parahalosydnopsis tubicola. — Pettibone 1977, Essays on polychaetous annelids in Memory of Dr. Olga Hartman: 47‑50, fig. 4;
Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 74‑76, fig. 14.
Fig. 26: Parahalosydnopsis tubicola (SMF 13619). a: anterior end; b: 2nd left elytron; c: 21st right elytron d: microtubercles of central
surface from b; e: microtubercles of lateral surface from b; f: microtubercles of central part from c; g: right parapodium from seg-
ment 41, anterior view; h: same, posterior view; i: notochaeta; j: upper neurochaeta; k: lower neurochaeta.
116 T. Wehe
Ty p e l o c a l i t y : Mirkae Wada, Ratnagiri, Maharashtra State, Western India, Indian Ocean.

S p e c i m e n s e x a m i n e d : T h e G u l f : 1 spm., Jebel Ali, Kingdom of Saudi Arabia, from sea grass bed, coll. M. Apel,
19 March 1998, SMF 13618; 2 spms., Jebel Ali, Kingdom of Saudi Arabia, from sea grass bed, coll. M. Apel, 19 March 1998, SMF
13619.
M e a s u r e m e n t s : SMF 13618: cs (71) 36/5/3 [af (23) 10 + mf (19) 12 + pf (29) 14]. SMF 13619: af (47) 20/4/3, pf (64)
20/4/3.
D i a g n o s i s : Elytra without fringing papillae. Surface with irregularly ridged or pointed mi-
crotubercles. Neurochaetae subdistally with 2 to 6 pairs of broad spines.
D e s c r i p t i o n : Body slightly flattened dorsoventrally, elongate. Prostomium (Fig. 26 a)
rounded. Style of antennae with subterminal swelling and terminal filum. Median style much
longer with few papillae on swelling (all styles broken off on specimen SMF 13618). Palps long,
stout, conical with minute papillae, gradually tapering. Two pairs of eyes, anterior pair dorsolater-
ally on more or less widest part of prostomium, posterior pair in front of hind margin. Tentacu-
lophores with up to two very fine chaetae. Tentacular cirri shaped as antennae, smooth, not as
long as median antenna. Facial tubercle distinct. Buccal cirri longer and inserted more proximally
than following ventral cirri, shaped as tentacular cirri. First pair of elytra rounded to oval, second
pair more elongate-oval (Fig. 26 b), following elytra broader oval (Fig. 26 c), becoming smaller
again on posteriormost segments. Scar of attachment off centre. Margin without fringing papillae,
surface above elytrophore and on lateral parts with irregularly ridged or pointed microtubercles
(Fig. 26 d, 26 e, 26 f ), central microtubercles slightly larger than lateral ones. On posterior elytra
ridged or pointed microtubercles becoming more concentrated above and laterally to attachment
scar of elytrophore, leaving most parts of surface smooth. Dorsal cirri with conical to cylindrical
cirrophore and long, smooth style with slight subterminal swelling and terminal filum. Dorsal tu-
bercles small to indistinct. Parapodia (Fig. 26 g, 26 h) with very small to inconspicuous notopodia.
Neuropodia rounded to diagonally truncate; prechaetal, acicular lobe only slightly longer than
postchaetal lobe, both with digitiform papillae on distal borders. Neuropodia and ventral body sur-
face more or less distinct with minute papillae. Ventral cirri short, gradually tapering. Nephridial
papillae distinct from about segment 10 onwards. Notochaetae (Fig. 26 i) moderate in number,
delicate, thread-like, faintly spinous, tapering to capillary tips, often thickly covered by inorganic
matter. Neurochaetae (Fig. 26 j, 26 k) much stouter, falcate, subdistally with slight swelling and
with 2 to 6 pairs of broad flattened spines below slightly curved, blunt, unidentate tips.
R e m a r k s : As can be deduced from the sequence of the arrangement of the elytra in Paraha-
losydnopsis, the anterior and posterior fragment of SMF 13619 belong to two different specimens.
The posterior fragment is 64 segments long starting with segment 12. This species or the genus
can therefore reach at least 75 segments. The generic diagnosis was emended accordingly.
B i o l o g y & e c o l o g y : According to Day (1973), commensal in tubes of terebellid polycha-
etes. The specimens examined here had been sorted before examination and it is unknown whether
they were collected from terebellid tubes or not.
D e p t h r a n g e : Unknown; elsewhere reported from the intertidal (Day 1973).
D i s t r i b u t i o n : Arabian Sea: Karachi (Rasheed & Mustaquim 2003), The Gulf. Elsewhere:
South of Bombay, Western India, Indian Ocean (Day 1973).
Thormora Baird, 1865

Thormora Baird, 1865. — Journal of the Linnean Society of London, Zoology 8: 199.
Thormora. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 75.
Ty p e s p e c i e s : Thormora jukesii Baird, 1865.
D i a g n o s i s (emended): Body with 26 segments, short, dorsoventrally flattened, subrec-
tangular in cross-section. Ceratophores of lateral antennae inserted terminally. Styles of antennae
smooth, with subterminal swelling. Palps stout, conical, gradually tapering, usually papillate.
Two pairs of eyes usually present. Tentaculophores usually with chaetae. Second (buccal) segment
without nuchal fold. Elytra, 12 pairs on segments 2, 4, 5, 7, alternating to 23, covering dorsum
completely or leaving parts of middorsum uncovered; surface ornamented with microtubercles
and papillae; margin with or without fringing papillae. Dorsal cirri with large, flattened cirrophore
and smooth styles with or without subdistal swelling. Notopodia small. Neuropodia with pre- and
postchaetal lobes of similar length. Notochaetae of two kinds: outer palisade of stout chaetae with
blunt tips, and an inner bundle of slender chaetae with lancet-shaped tips. Neurochaetae stouter
than notochaetae, falcate, with rows of spines below uni- or bidentate tips.
R e m a r k s : Based on the specimens examined, it is not obligatory for the genus that the mid-
dorsum remains uncovered by the elytra as stated in the diagnosis by Hanley & Burke (1991 a).
The generic diagnosis is emended in this regard.
Key to the species of Thormora
1 Papillae on palps may be Y‑shaped or simple. Elytra with large warty macrotubercles.
Margin with small globular papillae T. jukesii
– Papillae on palps simple. Elytra with smooth margins and smooth macrotubercles
T. salvati
Thormora jukesii Baird, 1865 Fig. 27 a‑o

Thormora jukesii Baird, 1865. — Journal of the Linnean Society of London, Zoology 8: 199‑200; Hanley & Burke 1991 a,
Mémoires du Muséum National d’Histoire Naturelle (A) 151: 75‑77, figs 25, 26; Rasheed & Mustaquim 2003, Pakistan
Journal of Marine Sciences 12(1): 79‑81, fig. 16; Barnich et al. 2004, Species Diversity 9: 301.
Polynoe (Lepidonotus) trissochaetus. — Ehrenberg & Grube in Grube, 1869, Monatsberichte der Königlich Preussischen Akad-
emie der Wissenschaften zu Berlin aus dem Jahre 1869: 485‑486.
Lepidonotus (Thormora) jukesii. — Seidler 1923, Archiv für Naturgeschichte 89 (Abt. A/11): 88‑91.
Lepidonotus (Thermora) [sic.] jukesi. — Fauvel 1955, Annales de l’Institute Océanographique Paris 30: 103.
Lepidonotus (Thormora) jukesi. — Ben-Eliahu 1977 a, Israel Journal of Zoology 25: 158.
Ty p e l o c a l i t y : Australia ?
S p e c i m e n s e x a m i n e d : H o l o t y p e : Thormora jukesii, Australia ?, Coll. Jukes, BMNH 1952.1.12.1. S y n t y p e
+ microscope slide: Lepidonotus trissochaetus, Red Sea, coll. C. Ehrenberg, ZMB 489. — R e d S e a : 1 spm., Jubal (=Gûbàl),
Gulf of Suez, madreporarian reef, coll. R.Ph. Dollfus, 19 December 1928, MNHN A894; 1 spm., as Lepidonotus jukesii, Gulf of
Aqaba, opposite to Wadi Kabila, endofauna in Dendropoma sp., coll. N.M. Ben-Eliahu, 30 April 1971, BMNH 1976.262‑271;
1 spm., Sudan, Sanganeb Atoll, inner reef, 12 m, from dead Acropora sp., coll. 9 April 1991, SMF 13530; 1 spm., Kunfuda,
Saudi Arabia, “Pola Expedition”, 1895‑1898, NHMW 16654; 1 spm., Abulat Island, “Calypso”, coll. M. Drach, M.M. Cherbon-
nier & C. Mercier, 1952, MNHN A364. — A r a b i a n S e a : 1 spm., Socotra Island, Ras Eriyhan-Quariyah, west of di-Hamri,
12°38.924'N/54°14.387'E, 10‑12 m, coll. T. Wehe, 20 April 2000, NHCY; 2 spms., Socotra Archipelago, Darsha Island, north-
eastern side, 1 km east of central plain, 12°7.273'N/53°17.598'E, 6‑8 m, coll. M. Apel, 8 April 2000, SMF 13531.
M e a s u r e m e n t s : Holotype BMNH 1952.1.12.1: cs (26) 37/10/8. Syntype ZMB: cs (26) 14/5/4. MNHN A894: cs
(26) 14/5/3. BMNH 1976.262‑271: af (13) 6/3.5/3 + mf (8) 5. SMF 13530: cs (26) 10/4/3. NHMW 16654: cs (26) 18/7.5/5.5.
MNHN A364: cs (26) 13/5/4. NHCY: cs (26) 11/4/3. SMF 13531: cs (26) 17/6/4, cs (26) 20/8/6.
D i a g n o s i s : Elytra with large, slightly warty macrotubercles, erect on anterior elytra, globu-
lar on following ones, and carinate microtubercles. Margin with scattered small globular papillae.
Neurochaetae unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Colour mostly faded in alcohol, and if rem-
nants of pigmentation are present, antennae and cirri with brown band below subterminal swell-
ing, elytra with dark brown patches, scar of attachment outlined by thin dark line, and transverse
brownish bands on dorsum. Prostomium (Fig. 27 a) rounded to oval, slightly wider than long,
118 T. Wehe
Fig. 27: Thormora jukesi (SMF 13531). a: anterior end; b: y‑shaped papillae on palps; c: 1st right elytron; d: 4th right elytron; e:
macrotubercles from c; f: macrotubercle from d; g: carinate microtubercles; h: globular papillae on elytra surface; i: globular papillae
on elytra margin; j: left parapodium from segment 10, anterior view; k: same, posterior view; l: stout notochaeta from outer pali-
sade; m: lancet-shaped notochaeta from inner bundle; n: upper neurochaeta; o: lower neurochaeta.
sometimes strongly withdrawn into anterior segment. Antennae with a long, smooth style, with
subterminal swelling and terminal filum. Styles of lateral antennae shorter than median style. Palps
stout, long, gradually tapering, abruptly terminating into a short terminal filum, with rows of very
prominent bifurcate, Y‑shaped or simple papillae (Fig. 27 b). Two pairs of eyes, anterior pair dor-
solaterally on more or less widest part of prostomium, posterior pair in front of hind margin. Eyes
may be hidden from view if prostomium withdrawn (see Fig. 27 a). Everted pharynx with two semi-
circles of nine soft papillae distally and two pairs of chitinous jaws. Tentaculophores with one or two
long chaetae each. Tentacular cirri shaped as antennae, dorsal longer than ventral cirri, both shorter or
as long as median antenna. Facial tubercle distinct. Buccal cirri longer and inserted more proximally
than following ventral cirri, shaped as tentacular cirri. Elytra (Fig. 27 c, 27 d) comparatively small,
leaving middorsum mostly uncovered. Elytra with outer margins curled upwards. First pair rounded
to slightly quadrate, second small oval, becoming broader and oval in following elytra. Surface with
two types of tubercles: faintly rugose macrotubercles scattered over surface, very prominent and erect
on first pair of elytra (Fig. 27 e), gradually decreasing in size and becoming more globular on fol-
lowing elytra (Fig. 27 f); globular to flattened, carinate microtubercles (Fig. 27 g) confined to lateral
parts of surface, numerous from second pair of elytra onwards. Small globular papillae scattered over
surface (Fig. 27 h). Similar papillae scattered along margin (Fig. 27 i), but no distinct fringe of papil-
lae. Dorsal cirri with large, globular to conical, slightly flattened cirrophore and long, smooth style,
with slight subterminal swelling and terminal filum. Dorsal tubercles absent. Parapodia (Fig. 27 j,
27 k) with small more or less conical notopodia. Neuropodia large, with weakly triangular, precha-
etal, acicular lobe and bluntly rounded, postchaetal lobe of similar length. Ventral cirri short, conical,
gradually tapering. Nephridial papillae distinct from about segment 9‑10 onwards. Notochaetae very
numerous, in dense tuft-like bundles, of two kinds: outer palisade of short, stout, curved chaetae with
rows of spines below blunt, unidentate tips (Fig. 27 l), and inner bundle of long, fine, smooth chaetae
with lancet-shaped tips (Fig. 27 m). Neurochaetae stouter, falcate, subdistally with slight swelling and
several rows of spines below slightly hooked, blunt, unidentate tips (Fig. 27 n, 27 o).
R e m a r k s : Some characters appear to be somewhat variable. For example, Y‑shaped papil-
lae on the palps were not observed on every specimen. Also, the middorsum is uncovered only in
larger specimens, whereas it is covered in smaller ones. The notochaetae are sometimes arranged in
very dense bundles directed in a lateral direction, or the notochaetae are spread, forming a hemi-
spherical dorsally directed tuft. In the latter case, chaetae are often broken off and are obviously
more vulnerable in this position. It is assumed that the different position of the notochaetae is due
to the contracted state of the musculature and is not a specific difference.
The type specimen of Lepidonotus trissochaetus (ZMB 489) is labelled as lectotype and was des-
ignated as such by Pettibone in 1973, according to Hartwich (1993). As stated by the latter, this
designation was never published and the specimen is here regarded as a syntype.
Specimens reported in Fauvel (1957, 1958) and Amoureux (1981) could not be located for
examination. However, the occurrence of the species in the Red Sea is confirmed by the specimens
examined here.
D e p t h r a n g e : So far recorded between 6 and 12 m; elsewhere reported from intertidal to
95 m (Hanley & Burke 1991 a, Hanley 1993).
D i s t r i b u t i o n : Red Sea, Arabian Sea: Socotra Archipelago and Karachi (Rasheed & Mus-
taquim 2003). Elsewhere: Malay Archipelago, Hong Kong, South China Sea, Australia, New Cal-
edonia, North-Western Pacific (Fauvel 1953, Rullier 1972, Uschakov 1982, Hanley & Burke
1991 a, Hanley 1992, 1993, Barnich et al. 2004).
Thormora salvati Rullier, 1972 Fig. 28 a‑m

Lepidonotus (Thormora) salvati Rullier, 1972. — Expédition Française sur les récifs coralliens de la Nouvelle-Calédonie 6: 31‑33, fig.1.
Ty p e l o c a l i t y : Bay of Saint-Vincent, New Caledonia, Western Pacific.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Lepidonotus (Thormora) salvati, Bay of Saint-Vincent, New Caledonia, coll.
Salvat, 15 November 1961, MNHN POLY TYPE 1335. — R e d S e a : 3 spms., Gulf of Aqaba, 5 km north of Dahab, reef flat,
120 T. Wehe
from dead coral, 0‑2 m, coll. C. Lewinsohn, 5 November 1981, NMNHT VR 25059. — A r a b i a n S e a : 1 spm., Socotra Island,
south coast, Mafrihin, fish landing site, 12°24.679'N/54°13.626'E, 0‑1 m, out of stranded sponge, coll. M. Apel, 22 April 2000,
SMF 13641.
M e a s u r e m e n t s : Holotype MNHN POLY TYPE 1335: cs (26) 8/4/3 [af (12) 3 + pf (14) 5]. NMNHT VR 25059: cs
(26) 23/8/6, cs (26) 20/7/5, cs (26) 15/5.5/4. SMF 13641: cs (26) 24/7/5 [af (15) 13 + pf (11) 11].
D i a g n o s i s : Elytra with small, smooth macrotubercles, erect on anterior elytra, globular on
following elytra, with carinate microtubercles. Margin with or without small globular papillae.
Neurochaetae unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Pigmentation of elytra hardly faded, almost com-
pletely green to black, with a distinct white spot above attachment of elytrophore. Brownish bands
visible subterminally on antennae and cirri. Styles of lateral antennae also brownish basally. Prostomi-
um (Fig. 28 a) strongly withdrawn into all specimens. Antennae with long, smooth style, with slight
subterminal swelling and terminal filum, median style longer than lateral styles. Palps stout, long,
gradually tapering, abruptly terminating into terminal filum, with rows of simple papillae (Fig. 28 b).
First pair of eyes present dorsolaterally on more or less widest part of prostomium, posterior pair in
front of hind margin. Both pairs hidden from view due to withdrawal (see Fig. 28 a). Tentaculophores
with one or two long chaetae each. Tentacular cirri shaped as antennae, both pairs about as long as
median antenna. Facial tubercle distinct. Buccal cirri longer and inserted more proximally than fol-
lowing ventral cirri, shaped as tentacular cirri. Elytra (Fig. 28 c, 28 d) large, covering dorsum com-
pletely. Elytra with outer margins curled upwards. First pair rounded to quadrate, following elytra oval
to rectangular. Surface with two types of tubercles: erect, conical, almost smooth macrotubercles on
first pair of elytra, becoming smaller and globular on following elytra (Fig. 28 e); globular to flattened,
carinate microtubercles (Fig. 28 f) confined to lateral parts of surface. Small globular papillae scattered
over surface (Fig. 28 g). Margin completely smooth. Dorsal cirri with large, conical, slightly flattened
cirrophore and long, smooth style, without distinct subterminal swelling, with slender terminal filum.
Dorsal tubercles absent. Parapodia (Fig. 28 h, 28 i) with small more or less conical notopodia. Neuro-
podia large, prechaetal and postchaetal lobe bluntly rounded and of similar length. Ventral cirri short,
conical, gradually tapering. Nephridial papillae distinct from segment 8 onwards. Notochaetae very
numerous, in dense tuft-like bundles, of two kinds: outer palisade of short, stout, curved chaetae with
rows of spines below blunt, unidentate tips (Fig. 28 j), and inner bundle of long, fine, smooth chaetae
with lancet-shaped tips (Fig. 28 k). Neurochaetae stouter, falcate, subdistally with a slight swelling and
several rows of spines below slightly hooked, blunt, unidentate tips (Fig. 28 l, 28 m).
R e m a r k s : The characters of the specimens examined here agree very well with the holotype,
with the slight difference that no small globular papillae were found on the margins of the elytra of
the non-type specimens.
There are about 10 species described as belonging to the genus Thormora, depending on the
synonymies accepted (see Day 1975, Hanley 1993). According to the literature, all these species
are very similar but there are no published keys to distinguish them. T. salvati is very similar to T.
jukesii, with the exception of Y‑shaped papillae on the palps in some specimens of T. jukesii. There
are also rather small differences in the quality of the macrotubercles on the elytra, which appear
to be larger and more rugose in L. jukesii, whereas they are smaller and somewhat smoother in T.
salvati. A revision of Thormora is needed to clarify whether these differences are mere variations or
are a specific difference.
B i o l o g y & e c o l o g y : Specimens from the Gulf of Aqaba were found amongst corals, and
the one from Socotra Island from sponge cavities.
D e p t h r a n g e : So far recorded between 0 and 2 m; elsewhere unknown.
D i s t r i b u t i o n : Red Sea, Arabian Sea: Socotra Archipelago. This is the first record of the
species from the Red Sea and the Arabian Sea. Elsewhere: New Caledonia (Rullier 1972).
Fig. 28: Thormora salvati (SMF 13641). a: anterior end; b: papillae on palps; c: 2nd right elytron; d: right elytron from midbody; e:
macrotubercles; f: microtubercles; g: globular papillae; h: left parapodium from segment 10, anterior view; i: same, posterior view; j:
stout notochaeta from outer palisade; k: lancet-shaped notochaeta from inner bundle; l: upper neurochaeta; m: lower neurochaeta.
122 T. Wehe
Polynoinae Kinberg, 1856

D i a g n o s i s : Body usually short, rarely with more than 50 segments. Prostomium with or
without cephalic peaks, with three antennae. Median antenna with a large ceratophore in anterior
notch of prostomium, lateral antennae with distinct ceratophores inserted ventrally or terminoven-
trally on prostomium. Palps stout, conical. Usually with two pairs of eyes, arrangement variable.
Tentaculophores with or without chaetae. Second (buccal) segment with or without nuchal fold.
Elytra often 15 pairs on segments 2, 4, 5, 7, alternating to 23, then variable in arrangement, often
continuing on 26, 29, 32; some genera with more or less than 15 pairs. Elytra surface usually or-
namented with various tubercles or papillae, margin with or without fringing papillae. Dorsal cirri
usually with papillate styles. Dorsal tubercles usually distinct. Notopodia large to small, with ac-
icular process. Neuropodia dorsally and ventrally not deeply incised; prechaetal, acicular lobe often
with supraacicular process; postchaetal lobe bluntly rounded, slightly shorter. Notochaetae usually
numerous, shape variable. Neurochaetae usually numerous, uni- or bidentate, shape variable. Py-
gidium with a pair of anal cirri.
Key to the genera of Polynoinae
1 Prostomium without cephalic peaks........................................................................... 2

– Prostomium with cephalic peaks (sometimes indistinct in Paradyte and Hololepi-
della).......................................................................................................................... 4
2 Elytra with poly- or hexagonal compartments, without tubercles or papillae
Gaudichaudius
Elytra smooth or with small tubercles. Without any compartments........................... 3
3 Parapodia with ventral lamellae Paralepidonotus
– Parapodia without ventral lamellae Malmgreniella
4 Elytra numerous pairs on segments 2, 4, 5, 7, …, 23, 26, 29, 31, 34, 36, 38, alternat-
ing to end of body. Segments up to 90 Hololepidella
– 15 or 16 pairs of elytra on segments 2, 4, 5, 7, …, 23, 26, 29, 32 (35)...................... 5
5 16 pairs of elytra on segments 2, 4, 5, 7, …, 23, 26, 29, 32, 35 Australonoe
th
– 15 pairs of elytra on segments 2, 4, 5, 7, …, 23, 26, 29, 32 (rarely 16 pair on 34)... 6
6 At least neurochaetae with spinous pouches............................................................... 7
– Neurochaetae spinous, but spines not forming pouches............................................. 8
7 Notochaetae sabre-like, smooth or with very few spines along convex edge (cephalic
peaks on prostomium sometimes indistinct) Paradyte
– Notochaetae with numerous rows of spinous pouches (usually 15 pairs of elytra,
rarely 16) Subadyte
8 All neurochaetae with unidentate tips........................................................................ 9
– Neurochaetae with uni- and bidentate tips Harmothoe
9 Neuro- and notochaetae distinctly spinous. Notochaetae directed outwards Eunoe
– Neuro- and notochaetae faintly spinous. Notochaetae in hemispherical tuft, directed
dorsally (“hedgehog-like’) Hermadion
Australonoe Hanley, 1993

Australonoe Hanley, 1993. — Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and Fauna of
Rottnest Island, Western Australia 2: 306‑307.
Ty p e s p e c i e s : Australonoe willani Hanley, 1993.

D i a g n o s i s : Body flattened dorsoventrally, with up to 39 segments. Prostomium with
cephalic peaks. Lateral antennae inserted ventrally, with small ceratophores. Styles of antennae
papillate lateral styles shorter than median style. Palps stout, papillate, gradually tapering. Two
pairs of eyes, anterior pair anterodorsally slightly in front of widest part of prostomium, posterior
pair in front of hind margin. Tentaculophores with chaetae. Second (buccal) segment with small
nuchal fold, with buccal cirri longer and inserted more proximally than following ventral cirri.
Elytra 16 pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, 35, completely covering body.
Surface with few microtubercles and few small mounds, margins without fringing papillae. Dorsal
cirri with cylindrical cirrophore and papillate style. Dorsal tubercles prominent, nodular. Notopo-
dia large, with acicular process. Neuropodia with longer, prechaetal, acicular lobe with digitiform,
supraacicular process, and shorter postchaetal lobe. Notochaetae about as stout as neurochaetae,
slightly curved, with rows of spines and blunt, unidentate tips. Neurochaetae falcate, with rows of
spines below uni- and bidentate tips.
Australonoe willani Hanley, 1993 Fig. 29 a‑g

Australonoe willani Hanley, 1993. — Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and
Fauna of Rottnest Island, Western Australia 2: 307‑309, fig. 1 A‑I; Barnich et al. 2004, Species Diversity 9: 301‑302.
Lagisca flaccida. — Fauvel 1933, Mémoires de l’Institut d’Égypte 21: 40 [not Potts, 1910].
Ty p e l o c a l i t y : Little Pioneer Bay, Rottnest Island, Western Australia.
S p e c i m e n s e x a m i n e d : R e d S e a : 1 spm., as Lagisca flaccida, Jubal, “sur Madrèpore”, coll. R.Ph. Dollfus, 29 De-
cember 1928, MNHN A360; microscope slide, Gulf of Suez, Jubal, coll. R.Ph. Dollfus, IEA R/69 (probably belonging to MNHN
A360); 1 spm., det. Augener 1920 as Harmothoe sp., Gulf of Suez, coll. Hartmeyer, ZMH P‑24603.
M e a s u r e m e n t s : MNHN A360: cs (39) 15/5.5/3.5 [af (22) + pf (17)]. ZMH P‑24603: cs (39) 15/5/3.5.
D i a g n o s i s : Second segment with slender nuchal fold. Elytra surface with nodular micro-
tubercles in anterior third and with few conical raised mounds on posterior half, margin without
fringing papillae. Neurochaetae bidentate.
D e s c r i p t i o n : Body flattened dorsoventrally, tapering posteriorly, with 39 segments. Pros-
tomium (Fig. 29 a) with prominent cephalic peaks. Median antenna with large, cylindrical cer-
atophore, style broken off in both specimens. Lateral antennae with papillate styles, tapering to
a filiform tip. Palps stout, conical, long, papillate, gradually tapering. Two pairs of small pointed
eyes, anterior pair dorsolaterally before horizontal midline of prostomium, posterior pair in front
of hind margin. Tentaculophores with small acicular lobe and one chaeta each. Tentacular cirri
papillate, longer than lateral antennae. Second (buccal) segment with small, slender, subtriangular
nuchal fold. Buccal cirri longer and inserted more proximally than following ventral cirri, shaped
as tentacular cirri. Elytra (Fig. 29 b) soft, translucent; first pair small, rounded, following larger,
oval. Anterior third of surface with nodular microtubercles (Fig. 29 c), posterior half with a row
of few conically raised mounds. Margin without fringing papillae. Dorsal cirri with cylindrical
cirrophore and long, papillate style with terminal filum. Dorsal tubercles well developed, nodular.
Parapodia (Fig. 29 d) with comparatively large, conical notopodia, with a short acicular process.
Neuropodia with longer, subtriangular, prechaetal lobe, with supra-acicular process, and shorter,
bluntly rounded postchaetal lobe. Noto- and neuroaciculae penetrate epidermis. Ventral cirri short,
papillate, gradually tapering. Nephridial papillae distinct from about segment 6 to 7 onwards.
Notochaetae (Fig. 29 e) numerous, in dense bundles, short to long, slightly curved, with numerous
rows of serrations below blunt, unidentate tips. Neurochaetae (Fig. 29 f, 29 g) numerous, in verti-
cal fans, about as stout as and longer than notochaetae, falcate, subdistally with slight swelling and
numerous rows of spines below bidentate tips, with secondary tooth set well back from tip. Upper
and middle neurochaetae distinctly bidentate, lower ones with smaller secondary tooth.
124 T. Wehe
Fig. 29: Australonoe willani (ZMH V6141). a: anterior end; b: 5th right elytron; c: microtubercles; d: right parapodium of segment
20, posterior view; e: notochaeta; f: middle neurochaeta; g: lower neurochaetae.
R e m a r k s : The specimens examined differ only slightly from the description given by
Hanley (1993), in that the small mounds on the elytra are arranged more regularly in a row rather
than in a patch. The neurochaetae appear all to be bidentate, while the lower ones, which at first
glance seem to be unidentate, show an abrasion where the secondary tooth would be assumed to
be. This is the first record of the species apart from the original description. The specimens re-
ported as Lagisca flaccida by Amoureux et al. (1978), as listed in Wehe & Fiege (2002), could not
be located for examination.
D e p t h r a n g e : Unknown; elsewhere recorded from 0.5 m (Hanley 1993).
D i s t r i b u t i o n : Red Sea: Gulf of Suez. This is the first record of the species for the Red Sea.
Elsewhere: South China Sea, Western Australia: Rottnest Island (Hanley 1993, Barnich et al.
2004).
Eunoe Malmgren, 1865

Eunoe Malmgren, 1865. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 21: 61.
Eunoe. — Barnich & Fiege 2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 559: 29.
Ty p e s p e c i e s : Eunoe nodosa Sars, 1861: 58 [subsequently designated by Uschakov (1955 b)
according to Pettibone (1963)].
D i a g n o s i s : Body short, about 35 to 45 segments, rounded in cross-section. Prostomium
with cephalic peaks. Lateral antennae inserted ventrally, with small ceratophores. Styles of anten-
nae usually papillate, lateral ones much shorter than median one. Palps stout, gradually tapering.
Two pairs of eyes, anterior pair on more or less widest part of prostomium, posterior pair in front
of hind margin. Tentaculophores with chaetae. Buccal segment without nuchal fold, with buccal
cirri longer and inserted more proximally than following ventral cirri. Elytra 15 pairs on segments
2, 4, 5, 7, alternating to 23, 26, 29, 32, covering dorsum. Elytra surface usually variously orna-
mented with tubercles and papillae, margin with or without fringing papillae. Dorsal cirri with
cylindrical cirrophore and papillate style. Dorsal tubercles small. Notopodia large, with acicular
process. Neuropodia with longer prechaetal, acicular lobe with supra-acicular process, and shorter
postchaetal lobe. Noto- and neuroacicula penetrating epidermis. Notochaetae stout, with numer-
ous rows of spines. Neurochaetae more slender, subdistally with numerous rows of spines below
unidentate tips.
Eunoe yedoensis (McIntosh, 1885) Fig. 30 a‑k

Eunoe yedoensis McIntosh, 1885. — Report on the scientific results of the voyage of H.M.S. Challenger during the years
1873‑76 under the command of Captain George S. Nares, R.N., F.R.S. and the late Captain Frank Tourle Thomson, R.N.
Zoology 12: 75, pl. 15: fig. 4, pl. 19: fig. 9, pl. 10 A: figs 11, 12.
Eunoe depressa. — Rosenfeldt 1989, Senckenbergiana biologica 69(1/3): 216 [not Moore, 1905].
Ty p e l o c a l i t y : Yedo, Japan, Northwest-Pacific.
S p e c i m e n s e x a m i n e d : R e d S e a : 15 spms., det. P. Rosenfeldt 1985, as Eunoe depressa, 21°10'80''N/37°34'00''E,
MeSedA-I, “Sonne” 02, closing trawl, 733‑757 m, coll. H. Thiel & M. Türkay, 16 October 1977 [collecting data supplemented ac-
cording to Thiel (1980)], ZMH P‑18642; 4 spms., det. P. Rosenfeldt 1985, as Eunoe depressa, 20°54'90''N/37°26'10''E, MeSedA-I,
“Sonne” 02, closing trawl, 490‑588 m, coll. H. Thiel & M. Türkay, 17 October 1977 (revised by R. Barnich 2000, as Eunoe yedoen-
sis), SMF 3800; 11 spms., det. P. Rosenfeldt 1985, as Eunoe depressa, 21°12'00''N/37°26'80''E, MeSedA-I, “Sonne” 02, 724‑747 m,
coll. H. Thiel & M. Türkay, 15 October 1977 (revised by R. Barnich, 2000 as Eunoe yedoensis), SMF 3825.
M e a s u r e m e n t s : ZMH P‑18642: cs (40) 10/3/2, af (18) 4/3/2, af (17) 3/3/2, af (16) 4/3/2, af (18) 4/3/2, af (22) 4/3/2,
af (18) 3/3/2, af (25) 5/3, 2/2.5, af (20) 3/2/1.2, af (20) 4/2/1.2, af (22) 5/3/2, af (22) 5/3/2, af (20) 4/3/2, af (20) 5/2.5/1.5, af
(25) 7/3/2. SMF 3800: af (30) 7/3/2, af (25) 5/3/2, af (24) 3/1.5/0.8, af (21) 2/1.9/0.8, pf (7) 1/2/1. SMF 3825: cs (37) 8/3/2,
cs (32) 5/2/1, cs (33) 7/2.5/1.8, cs (35) 7/3/2, af (14) 2/1.5/0.8, af (18) 4/3/2, af (25) 4/3/2, af (16) 3/3/2, af (16) 3/3/2, af (29)
7/3/2, af (22) 6/3/2.
126 T. Wehe
D i a g n o s i s : Elytra margin with small fringing papillae, surface with thorn-like microtuber-
cles. Notochaetae with very faint rows of spines, tips blunt. Neurochaetae with rows of spines and
pointed, unidentate tips.
D e s c r i p t i o n : Body short, rounded to rectangular in cross-section. Prostomium (Fig. 30 a)
with prominent cephalic peaks. Ceratophores of antennae distinctly broader than styles. Median
antenna with style about twice as long as styles of lateral antennae. All styles papillate, gradually
tapering. Palps long, gradually tapering, smooth. Facial tubercle mostly distinct. Tentaculophores
with up to four chaetae each. Tentacular cirri shaped as antennae, about as long as median an-
tenna, dorsal slightly longer than ventral cirri. Buccal cirri much longer than following ventral
cirri, inserted proximally, shaped as antennae and tentacular cirri. First pair of elytra rounded,
following reniform to oval (Fig. 30 b). Elytra surface covered with conical, pointed, thorn-like
microtubercles (Fig. 30 c) and few, scattered, digitiform papillae (Fig. 30 d). Lateral margin with
several short papillae (Fig. 30 e). Dorsal cirri with globular cirrophore and long, slender, papillate
style, gradually tapering to a long terminal filum. Dorsal tubercles indistinct. Parapodia (Fig. 30 f,
30 g) with notopodia small, conical, tapering to short acicular process. Neuropodia with slightly
longer, subtriangular, prechaetal lobe, with supra-acicular process, and shorter rounded postchaetal
lobe. Noto- and neuroacicula penetrate epidermis. Ventral cirri short, papillate, gradually tapering.
Nephridial papillae very small, from about segment 10 onwards if present. Notochaetae (Fig. 30 h)
numerous, arranged in tufts, short to long, sabre-like, with numerous but very faint rows of spines
only visible at high magnification (>40 x), and blunt, unidentate tips. Neurochaetae (Fig. 30 i,
30 j, 30 k) numerous, in vertical fan, more slender than notochaetae, longer, falcate, subdistally
with only a slight swelling and numerous rows of fine spines below pointed, unidentate tips.
R e m a r k s : The occurrence of E. depressa Moore, 1905 in the Red Sea, as listed in Wehe &
Fiege (2002), could not be confirmed.
D e p t h r a n g e : So far recorded between 490 and 757 m; elsewhere reported from about
170 m (Imajima & Hartman 1964).
D i s t r i b u t i o n : Red Sea. This is the first report of E. yedoensis from the Red Sea. Elsewhere:
Japan (Izuka 1912, Imajima & Hartman 1964).
Gaudichaudius Pettibone, 1986

Gaudichaudius Pettibone, 1986 a. — Smithsonian Contributions to Zoology 428: 33‑34.
Ty p e s p e c i e s : Iphione cimex Quatrefages, 1866.
D i a g n o s i s : Body flattened dorsoventrally, with less than 40 segments. Prostomium with-
drawn into anterior segments, without cephalic peaks. Lateral antennae inserted ventrally or termi-
noventrally. Styles of antennae papillate, lateral much shorter than median style. Palps stout, grad-
ually tapering. Two pairs of eyes, anterior pair on more or less widest part of prostomium, posterior
pair in front of hind margin. Tentaculophores with chaetae. Buccal segment with nuchal fold, with
buccal cirri longer and inserted more proximally than following ventral cirri. Elytra 15 pairs on
segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, covering dorsum completely. Elytra surface with
honeycomb-like polygonal compartments without tubercles, margin with fringing papillae. Dorsal
cirri with cylindrical cirrophore and long, papillate style. Dorsal tubercles transversely elongate or
forming digitiform extensions. Notopodia of moderate size, with acicular process. Neuropodia
with longer, prechaetal, acicular lobe with acicular process, and shorter postchaetal lobe. Notocha-
etae more slender than neurochaetae, numerous, short with blunt tips or long, slender with capil-
lary tips. Neurochaetae numerous, falcate, with rows of spines below unidentate tips.
Fig. 30: Eunoe yedoensis (SMF 3825). a: anterior end; b: detached elytra; c: microtubercles; d: papilla on elytra surface; e: fringing
papillae; f: left parapodium from segment 12, anterior view; g: same, posterior view; h: notochaetae; i: upper neurochaeta; j: middle
neurochaeta; k: lower neurochaeta.
128 T. Wehe
Gaudichaudius cimex (Quatrefages, 1866) Fig. 31 a‑k

Iphione cimex Quatrefages, 1866. — Histoire naturelle des Annelés marins et d’eau douce. Annélides et Géphyriens: 270.
Gaudichaudius cimex. — Pettibone 1986 a, Smithsonian Contributions to Zoology 428: 34‑37, figs 16, 17; Rasheed & Mus-
taquim 2003, Pakistan Journal of Marine Sciences 12(1): 51‑54, figs 2, 3; Barnich et al. 2004, Species Diversity 9: 303.
Ty p e l o c a l i t y : Malacca Strait, Malay Archipelago, Indo-West Pacific.
S p e c i m e n s e x a m i n e d : A r a b i a n S e a : 1 spm., Pakistan, Karachi, Clifton, intertidal sand, coll. J. Mustaquim,
12 November 1998, SMF 13643; 1 spm., as ?Lepidonotus sp., International Indian Ocean Expedition, 17°54'N/72°23' to 72°27'E,
46‑55 m, coll. 13 November 1963 [collecting data supplemented according to Hartman (1974 a)], LACM-AHF AB201 A.
M e a s u r e m e n t s : SMF 13643: cs (38) 15/5.5/4. LACM-AHF AB201 A: af (18) 4/3/2.
D i a g n o s i s : Lateral antennae inserted ventrally. Honeycomb-like compartments on elytra
close-fitting. Dorsal tubercles with digitiform branchiae. Notochaetae mostly long, slender with
capillary tips.
D e s c r i p t i o n : Body short, flattened dorsoventrally. Prostomium (Fig. 31 a) almost hexago-
nal, slightly wider than long. Median antenna with very large ceratophore and long, papillate style
with subterminal swelling and terminal filum. Lateral antennae much smaller, inserted ventrally;
ceratophores and papillate styles very slender, without subterminal swelling, nearly completely
hidden underneath ceratophore of median antenna. Palps stout, abruptly tapering to pointed tips,
with rows of very small papillae. Tentaculophores with few, slender chaetae. Tentacular cirri of
similar length, shaped as median antenna. Facial tubercle small. Buccal cirri longer and inserted
more proximally than following ventral cirri, shaped as tentacular cirri. Elytra (Fig. 31 b) yellowish,
tough, tightly imbricate; first pair oval, following elytra reniform. Elytra surface covered with close-
fitting hexa- or polygonal compartments (Fig. 31 c), which are slightly roughened (no secondary
areolae as in Iphione); scattered rounded spots between compartments and smooth surface along
margins give rise to minute papillae (Fig. 31 d); numerous, slender fringing papillae are found pos-
terolaterally (Fig. 31 e). Dorsal cirri with globular cirrophore situated on distal third of parapodia;
long, papillate style with subterminal swelling and terminal filum extending beyond chaetae. Dor-
sal tubercles with long, digitiform branchiae (Fig. 31 f, 31 g). Parapodia (Fig. 31 f, 31 g) with coni-
cal notopodia tapering to acicular process. Neuropodia with subtriangular, prechaetal, acicular lobe
and shorter, rounded postchaetal lobe. Ventral cirri with prominent cirrophore and short, papillate,
gradually tapering style. Notochaetae very numerous, of two types: outer palisade of short, strongly
bent chaetae with blunt tips (Fig. 31 h), and inner tuft of long, slender chaetae with capillary tips
(Fig. 31 i); both with numerous rows of spines. Neurochaetae (Fig. 31 j, 31 k) only slightly stouter,
longer, falcate, subterminally only faintly thickened, with several rows of spines below straight or
slightly curved, unidentate tips.
R e m a r k s : The specimens examined agree very well with the description provided by Petti-
bone (1986 a). Minor differences are the presence of minute papillae on the palps and the presence
of microtubercles on the rounded spots of the elytra, which have been described as “stomata-like”
by Pettibone (1986 a) or Rasheed & Mustaquim (2003). Both characters could be easily over-
looked. The species was reported by Pettibone (1986 a) from the Arabian Sea, but it was not
listed in Wehe & Fiege (2002) since none of the references listed in the synonymy by Pettibone
refers to specimens collected from this region rather than from south and eastern India. However,
the species was recently reported from the coastal waters of Pakistan by Rasheed & Mustaquim
(2003).
B i o l o g y & e c o l o g y : According to Kumaraswamy Achari (1977) as cited in Pettibone
(1986 a), the species lives as a commensal in the shells of hermit crabs. Adaptions to this habitat
are the flattened body and the unusual polynoid feature of digitiform branchiae on the dorsal
tubercles (which are well known in the burrowing species of Sigalionidae), as pointed out by Pet-
tibone (1986 a).
Fig. 31: Gaudichaudius cimex (SMF 13643). a: anterior end; b: detached elytron; c: compartments and papillae on elytra; d: surface
of elytra close to margin; e: fringing papillae; f: right parapodium from segment 16, anterior view; g: same, posterior view; h: noto-
chaeta of outer palisade; i: notochaeta of inner tuft; j: upper neurochaeta; k: lower neurochaeta.
130 T. Wehe
D e p t h r a n g e : So far recorded from the intertidal; elsewhere reported from shallow water to
50 m (Pettibone 1986 a).
D i s t r i b u t i o n : Arabian Sea. Elsewhere: Bay of Bengal, Burma, Malacca Strait, Indochina,
South China Sea, Yellow Sea (Pettibone 1986 a, Barnich et al. 2004).
Harmothoe Kinberg, 1856

Harmothoe Kinberg, 1856. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 12(9&10): 386.
Harmothoe. — Barnich & Fiege 2000, Journal of Natural History 34: 1891‑1893; Barnich & Fiege 2003, Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 559: 31‑33.
Ty p e s p e c i e s : Harmothoe spinosa Kinberg, 1856.
D i a g n o s i s : Body flattened dorsoventrally, with up to 50 segments. Prostomium with ce-
phalic peaks. Lateral antennae inserted ventrally, with small ceratophores. Styles of antennae papil-
late (rarely smooth), lateral ones much shorter than median one. Palps stout, gradually tapering.
Two pairs of eyes, anterior pair in a variable arrangement, either anteroventrally beneath cephalic
peaks, or dorsolaterally on more or less widest part of prostomium, posterior pair always in front
of hind margin of prostomium. Tentaculophores with chaetae. Second (buccal) segment without
nuchal fold, with buccal cirri longer and inserted more proximally than following ventral cirri.
Elytra 15 pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, more or less covering body,
but leaving tail uncovered in larger specimens. Elytra surface usually ornamented with microtuber-
cles, with or without macrotubercles and papillae, margin with or without fringing papillae. Dorsal
cirri with cylindrical cirrophore and papillate style. Dorsal tubercles usually nodular. Notopodia
moderate to large, with acicular process. Neuropodia with longer, prechaetal, acicular lobe with
supraacicular process, and shorter postchaetal lobe. Tips of noto- and neuroaciculae penetrating
epidermis. Notochaetae as stout as or stouter than neurochaetae, with distinct rows of spines and
blunt, unidentate tips. Neurochaetae falcate, with fewer rows of spines subdistally below bi- and
unidentate tips.
R e m a r k s : A worldwide revision of the genus is currently being undertaken by R. Barnich
and D. Fiege (Senckenberg Research Institute, Frankfurt a.M.). The taxonomy in Harmothoe and
related genera is highly confused. In order to avoid misidentifications and additional taxonomic
confusion, the specimens reported and described here are in accordance with the current state of
this project.
Key to the species of Harmothoe
1 First pair of eyes anteroventrally. Elytra surface not divided into compartments. Mac-
rotubercles absent...................................................................................................... 2
– First pair of eyes dorsolaterally. Elytra surface divided into hexa- or polygonal com-
partments. With macrotubercles................................................................................ 4
2 Elytra with microtubercles confined to anterior third of surface H. marerubrum
– Elytra surface completely covered by microtubercles.................................................. 3
3 Microtubercles on posterior half of elytra distinctly conical, pointed, almost thorn-like.
All neurochaetae bidentate, with deep incision and large secondary tooth H. liaoi
– Microtubercles on posterior half of elytra globular to conical. Neurochaetae bi- and
unidentate. Secondary tooth not as prominent H. grisea
4 Macrotubercles conical with one or two (rarely three) pointed tips H. dictyophora
– Macrotubercles bottle-shaped, with several bluntly rounded tips H. hirsuta
Harmothoe dictyophora (Grube, 1878) Fig. 32 a‑h

Polynoe dictyophorus Grube, 1878. — Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8): 44‑45, pl.
15: fig. 9.
Harmothoe dictyophora. — Fauvel 1918, Bulletin du Muséum d’Histoire Naturelle 24(5): 330; Hanley & Burke 1990, Proceed-
ings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia 1:
208‑211, fig. 3; Mustaquim 1991, Pakistan Journal of Zoology 23(4): 362, fig. 1; Rasheed & Mustaquim 2003, Pakistan
Journal of Marine Sciences 12(1): 54‑58, figs 4, 5; Barnich et al. 2004, Species Diversity 9: 306.
Ty p e l o c a l i t y : Philippines.
S p e c i m e n s e x a m i n e d : A r a b i a n S e a : 2 spms., Pakistan, Karachi, Buleji, intertidal rock, coll. Mustaquim, 21 Au-
gust 1997, SMF 13621. — T h e G u l f : 1 spm., Pole Reef, in and on dead and living Porites sp., 1‑3 m, coll. M. Apel, 22 Febru-
ary 1993, SMF 13620; microscope slide, “les bancs perliers, par 10 à 16 brasses, à environ 15 milles de la côte d’Oman”, 24°55'N
to 25°10'N/54°40'E to 55°10'E, dredge, coll. M.J. Bonnier & M. Ch. Pérez, 14‑16 March 1901 [the second label on the slide
states Suez; however, no specimen was caught there according to Fauvel (1918)], IEA P.58’.
M e a s u r e m e n t s : SMF 13621: cs (36) 12/5/4, cs (34) 12/5/4. SMF 13620: cs (36) 14/5/3.
D i a g n o s i s : Anterior pair of eyes dorsolaterally. Elytra surface divided into polygonal com-
partments; macrotubercles conical, spine-like, with uni-, bi- or trifid tips; margin with long, slen-
der fringing papillae. Neurochaetae mostly bidentate with secondary tooth large and set well back
from tip.
D e s c r i p t i o n : Body broad, flattened dorsoventrally. Prostomium (Fig. 32 a) rounded to
hexagonal, as wide as long. Styles of antennae strongly papillate, tapering to slender terminal filum.
Palps finely papillate. Two pairs of eyes, anterior pair dorsolaterally on more or less widest part
of prostomium, posterior pair in front of hind margin. Tentaculophores with up to four chaetae
each. Tentacular cirri about as long as median antenna, shaped as it. Facial tubercle small. Buccal
segment with small knob anteriorly. Buccal cirri longer and inserted more proximally than follow-
ing ventral cirri, shaped as tentacular cirri. Elytra (Fig. 32 b) large, covering dorsum. First pair of
elytra rounded, following elytra reniform to oval. Surface divided into polygonal compartments,
along anterior margin with smooth rim. Polygonal compartments near posterior border with large
conical, spine-like macrotubercles, with single or bifid tip (Fig. 32 c), on anterior elytra also trifid
macrotubercles may occur. Tubercles on each elytron gradually increasing in size from anterior to
posterior, but decreasing in number, up to six small microtubercles may be found in anterior fields
and a single large macrotubercle in posterior fields. Few, slender, filiform papillae scattered over
surface. Margin posterolaterally with a fringe of numerous, long, slender papillae. Dorsal cirri with
large, cylindrical cirrophore and long, strongly papillate style with terminal filum. Dorsal tubercles
distinct, nodular. Parapodia (Fig. 32 d, 32 e) with large, conical notopodia with short acicular
process. Neuropodial lobes of similar length, prechaetal lobe subtriangular, with digitiform supra-
acicular process. Postchaetal lobe bluntly rounded. Noto- and neuroaciculae penetrating epidermis.
Ventral cirri short, papillate, gradually tapering. Nephridial papillae from about segment 6 to
7 onwards. Notochaetae (Fig. 32 f ) numerous, in dense bundles, short to long, slightly curved,
with numerous rows of spines below fine, pointed tips. Neurochaetae (Fig. 32 g, 32 h) numerous,
in vertical fans, about as stout as notochaetae, longer, falcate, subdistally with numerous rows of
spines below bidentate tips on which the secondary tooth is large and set well back from tip; few
lower neurochaetae with unidentate tip. Neurochaetae of segment two much more slender than
following neurochaetae.
R e m a r k s : There are several species in Harmothoe with a pattern of polygonal compartments
on the elytra, but all of them may be distinguished by the shape of the macrotubercles. In H. dic-
tyophora they are uni-, bi- or even trifid, with the tips always sharply pointed. Usually, only uni- or
bifid tubercles are desribed or figured (see e.g. Fauvel 1953, Hanley & Burke 1990, Mustaquim
1991). It seems that the number of tips is variable and that it decreases from the anterior to pos-
terior elytra. Harmothoe hirsuta Johnson, 1897 is very close to H. dictyophora, but differs clearly
132 T. Wehe
Fig. 32: Harmothoe dictyophora (SMF 13620: a; SMF 13621: b-h). a: anterior end; b: right elytron from midbody; c: macrotu-
bercles and fringing papillae; d: left parapodium from segment 16, anterior view; e: same, posterior view; f: notochaeta; g: upper
neurochaeta; h: lower neurochaeta.
with respect to the macrotubercles. Specimens reported by Fauvel (1911, 1933) as H. dictyophora
have proved to belong to H. hirsuta (see description below), but those mentioned in Fauvel (1919,
1927) could not be located. The occurrence of H. dictyophora in the Suez Canal cannot be con-
firmed.
D e p t h r a n g e : So far recorded between 1 and 3 m; elsewhere reported from the intertidal
down to 20 m (Day 1967, Hanley & Burke 1990).
D i s t r i b u t i o n : Arabian Sea: Pakistan, The Gulf. Elsewhere: All over Indo-West Pacific:
South Africa, Madagascar, Vietnam, Sri Lanka, Bay of Bengal, Malay Archipelago, South China
Sea, Philippines, Australia (Fauvel 1953, Barnich et al. 2004).
Harmothoe grisea (Ehrenberg & Grube in Grube, 1869) Fig. 33 a‑l

Polynoe (Harmothoe) grisea Ehrenberg & Grube in Grube, 1869. — Monatsberichte der Königlich Preussischen Akademie der
Wissenschaften zu Berlin aus dem Jahre 1869: 489‑490.
Harmothoe boholensis Fauvel, 1911. — Archives de Zoologie Expérimentale et Générale, Cinquième Série 4(11): 369; Fauvel
1927, The Transactions of the Zoological Society of London 22/4(1): 415‑416; Fauvel 1933, Mémoires de l’Institut d’Égypte
21: 37‑38 [not Grube, 1878].
Harmothoe minuta Fauvel, 1933. — Mémoires de l’Institut d’Égypte 21: 38 [partim, see also Hololepidella nigropunctata; not
Potts, 1910].
Harmothoe goreensis Amoureux et al., 1978. — Israel Journal of Zoology 27: 64 [not Augener, 1918].
Ty p e l o c a l i t y : Tor (= At-Tur), Gulf of Suez, Red Sea, Sinai Peninsula, Egypt.
S p e c i m e n s e x a m i n e d : L e c t o t y p e d e s i g n a t e d h e r e : Harmothoe grisea, Tor (= At-Tur), Sinai Peninsula, Gulf
of Suez, Red Sea, coll. C.G. Ehrenberg, ZMB 493. 2 p a r a l e c t o t y p e s : same data as lectotype, ZMB 11172. 1 p a r a l e c t o -
t y p e (was dried out) + microscope slide: Harmothoe grisea, Red Sea, Coll. Grube, ZMB 1140. — S u e z C a n a l : 2 spm., as
Harmothoe boholensis, Bitter Lake, coll. “Cambridge Expedition to the Suez Canal”, 17 October 1924, BMNH 1926.11.12.126;
1 spm., as Harmothoe boholensis, Balise, coll. “Cambridge Expedition to the Suez Canal”, 1924, BMNH 1926.11.12.127; 2 spms.,
as Harmothoe boholensis, island in the Little Bitter Lake, coll. “Cambridge Expedition to the Suez Canal”, 24 November 1924 [col-
lecting data supplemented according to Fox (1926)], BMNH 1926.11.12.128; 3 spms., as Harmothoe boholensis, Canal Company’s
Signal Station, coll. “Cambridge Expedition to the Suez Canal”, 15 October or November 1924 [collecting data supplemented ac-
cording to Fox (1926)], BMNH 1926.11.12.129; 1 spm., as Harmothoe boholensis, Toussoum Signal Station, coll. “Cambridge Ex-
pedition to the Suez Canal”, 29 November 1924 [collecting data supplemented according to Fox (1926)], BMNH 1926.11.12.130;
2 spms., as Harmothoe boholensis, eastern side of Suez-Port Taufiq, intertidal sand flats, coll. “Cambridge Expedition to the Suez
Canal”, 25 November 1924 [collecting data supplemented according to Fox (1926)], BMNH 1926.11.12.131; 1 spm., as Har-
mothoe boholensis, Kantara, coll. “Cambridge Expedition to the Suez Canal”, 27 November 1924, BMNH 1926.11.12.132;
1 spm., as Harmothoe boholensis, Ferry Post, dredge, coll. “Cambridge Expedition to the Suez Canal”, 4 December 1924, BMNH
1926.11.12.133; 1 spm., as Harmothoe boholensis, western shore of Lake Timsah, coll. “Cambridge Expedition to the Suez Canal”,
10 December 1924 [collecting data supplemented according to Fox (1926)], BMNH 1926.11.12.134/135; 2 spm., Kabret Sig-
nal Station, as Harmothoe boholensis, coll. “Cambridge Expedition to the Suez Canal”, October 1924, BMNH 1926.11.12.136;
1 spm., as Harmothoe boholensis, Balise, coll. “Cambridge Expedition to the Suez Canal”, coll. M. Fox, 21 December 1924, MNHN
A403; microscope slide, as Harmothoe boholensis, Little Bitter Lake, dredge, mud, 2‑3 m, coll. “Cambridge Expedition to the Suez
Canal”, 24 November 1924 [collecting data supplemented according to Fox (1926)], IEA R/76’; microscope slide, as Harmothoe
boholensis, Kantara, coll. “Cambridge Expedition to the Suez Canal”, 27 November 1924 [collecting data supplemented according
to Fox (1926)], IEA R/78; microscope slide, as Harmothoe boholensis, Little Bitter Lake, on buoys, 0‑4 m, coll. “Cambridge Expe-
dition to the Suez Canal”, 17 October 1924 [collecting data supplemented according to Fox (1926)], IEA R/78’. — R e d S e a :
3 spms., det. H. Augener 1920 as Harmothoe sp., Tor (= At-Tur), Sinai Peninsula, Gulf of Suez, coll. Hartmeyer, January 1902,
ZMH V6140; 5 spms., det. H. Augener 1920 as Harmothoe sp., Tor (= At-Tur), Sinai Peninsula, Gulf of Suez, coll. Hartmeyer,
January 1902, ZMH V6141; 1 spm., as Harmothoe boholensis, Gulf of Suez, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m,
fine sand, amongst sponges, coll. R.Ph. Dollfus, 28 November 1928 [collecting data supplemented according to Billard (1933)],
MNHN A360; 1 spm., as Harmothoe minuta (Potts, 1910), Gulf of Suez, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m,
fine sand, coll. R.Ph. Dollfus, 28 November 1928 [collecting data supplemented according to Billard (1933)], MNHN A894;
microscope slide, as Harmothoe boholensis, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m, fine sand, coll. R.Ph. Dollfus,
28 November 1928 [collecting data supplemented according to Billard (1933)], IEA R/69’; microscope slide, as Harmothoe goreen-
sis, Gulf of Aqaba, Marsa-el-At, 25 m, from sponge, coll. 1973 [collecting data supplemented according to Amoureux et al. (1978)],
IEA Y99; 1 spm., Port Sudan, coll. Werner, 29 April 1914, NHMW 17260; 1 spm., as Harmothoe sp., Sudan, Suakin Harbour,
coll. C. Crossland, BMNH 1941.4.4.219; 11 spms., as Harmothoe sp., coll. C. Crossland, BMNH 1941.4.4.220‑4; 2 spms., Mersa
Sheikh, “Pola Expedition”, 1895‑1898, NHMW V17056. — T h e G u l f : Microscope slide, as Harmothoe boholensis, Bouchir (=
Būshehr), coll. M.N. Bogoyawlensky, 1 March 1902, IEA P.52.
M e a s u r e m e n t s : Lectoype ZMB 493: cs (39) 13/3/2 [af (22) 8 + pf (17) 5]. Paralectotypes ZMB 11172: cs (37)
9/2.5/1 [af (22) 5 + pf (15) 4], cs (38) 8/3/2 [af (25) 5 + pf (13) 3]. Paralectotype ZMB 1140: cs (36) 7.5/3/2. BMNH
1926.11.12.126: cs (37) 14/4.5/3.5, af (20) 6/4/3. BMNH 1926.11.12.127: cs (38) 12/4/3 [af (12) 3 + mf (16) 6 + pf (10)
3]. BMNH 1926.11.12.128: cs (39) 13/4/3 [af (18) 6 + pf (21) 7], cs (38) 12/3/2 [af (20) 6 + mf (8) 3 + pf (10) 3]. BMNH
1926.11.12.129: cs (37) 9/3/2.5, cs (35) 9/3.5/2.5, af (25) 7/3.5/2.5. BMNH 1926.11.12.130: af (17) 5/3.5/2.5, mf (16) 7, pf
134 T. Wehe
Fig. 33: Harmothoe grisea (lectotype, ZMB 493). a: anterior end; b: 1st left elytron; c: 2nd right elytron; d: microtubercles; e: digiti-
form papillae on surface; f: fringing papillae; g: parapodium from segment 27, anterior view; h: same, posterior view; i: notochaeta;
j: upper neurochaeta; k: middle neurochaeta; l: lower neurochaeta.
(21) 9. BMNH 1926.11.12.131: cs (34) 11/3/2 [af (18) 5 + pf (16) 6], cs (34) 10/3/2 [af (12) 3 + mf (4) 2 + pf (18) 5]. BMNH
1926.11.12.132: cs (39) 18/3/2. BMNH 1926.11.12.133: af (22) 5/3.5/2.5. BMNH 1926.11.12.134/135: af (22) 9/3.5/2.5.
BMNH 1926.11.12.136: cs (32) 9/3/2 [af (12) 3 + mf (7) 3 + pf (13) 3], af (18) 5/3.5/2.5 + mf (7) 2 + mf (6) 2. MNHN A403: cs
(37) 12/4/3 [af (22) 7 + mf (6) 2 + pf (9) 3]. ZMH V6140: cs (35) 8/3/2, cs (37) 10/2.5/1.8, cs (38) 10/3/2. ZMH V6141: cs (37)
7/2.2/1.8 [af (22) 4 + pf (15) 3], cs (36) 9/2.2/1.8, cs (37) 9/3/2, cs (37) 8/2.5/2, cs (34) 8/2.5/2. MNHN A360: af (25) 4/2.5/1.9.
MNHN A894: cs (35) 8/2/1.5. NHMW 17260: cs (37) 14/6/4 [af (16) 5 + mf (2) 1 + pf (19) 8]. BMNH 1941.4.4.219: cs (40)
17/6/4 [af (28) 12 + pf (12) 5]. BMNH 1941.4.4.220‑4: cs (39) 10/3/2 [af (18) + pf (21)], cs (37) 7/3/2, cs (37) 11/3/2 [af (16) 5
+ mf (9) 3 + pf (12) 3], cs (33) 8/3/2, cs (35) 8/2.5/2, cs (37) 9/2/1.5, cs (37) 8/2/1.5, af (30) 12/4/2.5, af (26) 6/2.5/1, 5, af (17)
3.5/2.5/1.5, af (18) 5/3/2. NHMW V17056: af (19) 4/5/3.5; mf (9) 3/5.5/4 + pf (13) 5.
D i a g n o s i s : Anterior pair of eyes anteroventrally. Elytra surface with globular to conical,

blunt-tipped microtubercles and digitiform papillae; margin with fringing papillae. Neurochaetae
mainly bidentate with secondary tooth set back from tip.
R e d e s c r i p t i o n based on lectotype designated here: Body flattened dorsoventrally. Pros-
tomium (Fig. 33 a) rounded to slightly hexagonal, about as wide as long. Style of median antenna
much longer than lateral styles; all styles strongly papillate, with slender terminal filum. Palps
stout, gradually tapering. Two pairs of eyes, anterior pair anteroventrally, slightly behind cephalic
peaks, posterior pair in front of hind margin. Tentaculophores with single chaetae on lectotype,
sometimes two in other specimens. Tentacular cirri about as long as median antenna, shaped as
it. Facial tubercle small. Buccal segment with small anterior knob. Buccal cirri longer and inserted
more proximally on parapodia than following ventral cirri, shaped as tentacular cirri, but not as
long. Elytra (Fig. 33 b, 33 c) covering dorsum (missing on posterior half of lectotype), first pair
rounded, following elytra reniform to oval. Surface covered with low, globular microtubercles in
anterior part, becoming slightly taller and more conical in posterior part (Fig. 33 d). Along ante-
rior border with small, smooth rim, with exception of first pair of elytra. Numerous digitiform pa-
pillae scattered over posterolateral surface (Fig. 33 e). Elytra margin posterolateral with digitiform
papillae of same type (Fig. 33 f ), gradually decreasing in size on posterior border. Dorsal cirri with
cylindrical cirrophore and strongly papillate style, gradually tapering, without distinct subterminal
swelling. Dorsal tubercles distinct, nodular. Dorsum with two transverse bands of cilia on each seg-
ment. Parapodia (Fig. 33 g, 33 h) with large conical notopodia. Neuropodia with slightly longer,
subtriangular, prechaetal, acicular lobe, with digitiform supraacicular process. Postchaetal lobe
slightly shorter and bluntly rounded. Noto- and neuroaciculae penetrating epidermis. Ventral cirri
short, papillate, gradually tapering. Nephridial papillae small, distinct from about segment 6 to
7 onwards. Notochaetae (Fig. 33 i) numerous, as stout as or slightly stouter than neurochaetae,
short to long, curved, with numerous rows of spines below blunt, unidentate tips. Neurochaetae
(Fig. 33 j, 33 k, 33 l) numerous, in vertical fans, falcate, with slight swelling and several rows of
spines subdistally, mainly with bidentate tips, on which the secondary tooth is set back from the
tip; few lower neurochaetae with unidentate tips.
R e m a r k s : The original description of H. grisea is brief and Grube (1869) gave no figures.
So far as the most important distinguishing characters within the genus Harmothoe are concerned
- the chaetae and elytra - Grube’s description unfortunately does not provide the detail needed
today. The species has not been reported since the original description, and is redescribed here.
Hartwich (1993) reported four syntypes, all of which were examined. A lectotype was chosen for
the purpose of redescription, namely the one specimen that most closely fits the original descrip-
tion, especially with regard to size. The paralectotypes are in bad condition and some of the char-
acters, e.g. the number of chaetae on the tentaculophores, the nephridial papillae, are not visible or
differ from the character states found in the lectotype. This is also the case with several of the non-
type specimens. However, these variations in character states are not regarded as of specific impor-
tance, and all of the specimens agree in the main diagnostic characters such as the arrangement of
eyes, chaetae and elytra structure. Although the species is reported here for the second time only, it
seems to be very common in the Suez Canal and the Northern Red Sea.
Apart from the early accounts by Savigny (1822) and Quatrefages (1866), Grube’s works
(1868 b, 1869) contain the very first available descriptions of polychaetes from the Red Sea. For
reasons unknown, the original description and the existence of H. grisea must have escaped the at-
tention of subsequent workers.
Fauvel (1911) identified a specimen collected from The Gulf as Harmothoe boholensis (Grube,
1878), mainly because of the pigmentation of the elytra, which in his opinion was similar to that
136 T. Wehe
figured for Polynoe boholensis by Grube (1878) and to which he referred as a “Maltese Cross”, a
term not used by Grube (1878). In contrast to Grube (1878), Fauvel placed his specimen in Har-
mothoe, based on the prostomial characters. Several more specimens were subsequently reported
as Harmothoe boholensis by Fauvel (1919, 1927, 1933) and other workers (Fishelson & Rullier
1969, Ben-Eliahu 1972 b, Amoureux et al. 1978), always referring to Fauvel’s early work. How-
ever, none of these authors evaluated properly other important characters described and figured by
Grube (1878), such as the prostomial shape and antennal arrangement, and the presence of ventral
lamellae on the bases of the parapodia. These characters do not fit with those of Harmothoe, as was
already recognised by Horst (1915) who erected the genus Paralepidonotus for this kind of charac-
ter combination. However, this was clearly not recognised by Fauvel (1919), since he again placed
the species in Harmothoe. It was not until Hanley (1991) that Horst’s (1915) opinion was con-
firmed, and Polynoe boholensis Grube, 1878 was again assigned to Paralepidonotus but this time as
a junior synonym of Paralepidonotus indicus (Kinberg, 1856). Hanley (1991) in turn synonymised
all the above-mentioned records of Harmothoe boholensis with Paralepidonotus indicus, without
recognising from the relevant descriptions that they do all indeed belong to the genus Harmothoe,
which is clearly expressed in Fauvel (1919: 333).
Revision of the relevant specimens published as H. boholensis and listed in Wehe & Fiege
(2002) has revealed that they belong to two different species of Harmothoe. Specimens described in
Fauvel (1911, 1927, 1933) belong to H. grisea, as described above, whereas specimens mentioned
in Fauvel (1919, 1955) and Ben-Eliahu (1972 b) belong to Harmothoe liaoi Barnich, Fiege &
Sun, 2004 or are indeterminable. The specimens reported in Fauvel (1957), Fishelson & Rullier
(1969) and Amoureux et al. (1978) could not be located for examination and their taxonomic
status remains unknown.
There is only a single microscope slide with two parapodes and an elytron belonging to the
specimen reported from The Gulf by Fauvel (1911), which represents the sole record from that
area. The slide is in bad condition, but the observed characters agree well with those seen on the
type specimens of H. grisea.
Three specimens from lot ZMH V6141 do not possess elytra. However, they agree in all other
characters with the remaining species of this series and also with the type specimens, and are here
referred to H. grisea.
Harmothoe goreensis as reported by Amoureux et al. (1978) was synonymised with H. bellani
Barnich & Fiege, 2000 by Barnich & Fiege (2000). There is only one remaining microscope
slide with mounted elytra and parapodia from the specimen recorded by Amoureux et al. (1978).
Re-examination revealed that the characters shown by the elytra and parapodia now appear to be
closer to H. grisea rather than to H. bellani, which is restricted to the Mediterranean Sea.
D e p t h r a n g e : So far recorded between 0 and 35 m.
D i s t r i b u t i o n : Suez Canal, Red Sea, The Gulf.
Harmothoe hirsuta Johnson, 1897 Fig. 34 a‑h

Harmothoe hirsuta Johnson, 1897. — Proceedings of the California Academy of Sciences, Third series, Zoology 1(5): 182, pl. 6:
figs 27, 28, pl. 7: fig. 38, pl. 8: figs 53 a‑c; Hartman 1968, Atlas of the errantiate polychaetous annelids from California:
77, figs 1‑6; Ruff, 1995, Taxonomic atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Chan-
nel. The Annelida Part 2. Polychaeta: Phyllodocida (Syllidae and scale-bearing families), Amphinomidae, and Eunicidae 5:
130‑132, fig. 3.10.
Harmothoe dictyophora Fauvel 1911. — Archives de Zoologie Expérimentale et Générale, Cinquième Série 4(11): 370; Fauvel
1933, Mémoires de l’Institut d’Égypte 21: 38 [not Grube, 1878].
Ty p e l o c a l i t y : San Pedro, California, Northeast Pacific.
Fig. 34: Harmothoe hirsuta (NMNH A360). a: anterior end; b: left elytron from midbody; c: macrotubercles and fringing papillae;
d: right parapodium from segment 6, anterior view; e: same, posterior view; f: notochaeta; g: middle neurochaeta; h: lower neuro-
chaeta.
138 T. Wehe
S p e c i m e n s e x a m i n e d : R e d S e a : 1 spm., as Harmothoe dictyophora, Gulf of Suez, 70 m, sand and mud, coll. R.Ph.
Dollfus, 28 December 1928 [collecting data supplemented according to Billard (1933)], MNHN A360; 1 spm., as Harmothoe
dictyophora, Gulf of Suez, coll. R.Ph. Dollfus, 30 December 1928, MNHN A360. — T h e G u l f : Microscope slide, as Harmothoe
dictyophora, Bouchir (= Bushire, Būshehr), coll. M.N. Bogoyawlensky, 1 March 1902, IEA P.52’.
M e a s u r e m e n t s : MNHN A360: af (14) 4/4/3 + mf (6) 2. MNHN A360: cs (36) 10/4/3.
D i a g n o s i s : Anterior pair of eyes dorsolaterally. Elytra surface divided into polygonal com-
partments; macrotubercles bottle-shaped with an irregular crown of several bluntly rounded tips;
margin with long, slender fringing papillae. Neurochaetae mostly bidentate with secondary tooth
large and set well back from tip.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 34 a) rounded to hexago-
nal, as wide as long. Styles of antennae strongly papillate, tapering to a slender terminal filum.
Palps with fine papillae. Two pairs of eyes, anterior pair dorsolaterally on more or less widest part
of prostomium, posterior pair in front of hind margin. Tentaculophores with two chaetae each.
Tentacular cirri about as long as median antenna, shaped as latter. Facial tubercle small. Buccal
segment with small triangular protrusion anteriorly, but no nuchal fold. Buccal cirri longer and
inserted more proximally than following ventral cirri, shaped as median antenna and tentacular
cirri. Elytra (Fig. 34 b) large, covering dorsum. First pair of elytra rounded, following elytra reni-
form to oval. Surface divided into polygonal compartments, but with a smooth rim along anterior
margin. Polygonal compartments on posterior half (Fig. 34 c) with large bottle-shaped macrotu-
bercles, with an irregularly-shaped crown of blunt tips. Tubercles on elytra gradually increasing
in size from anterior to posterior, but decreasing in number, numerous microtubercles in anterior
compartments and single large macrotubercle in posterior compartments. More slender and coni-
cal macrotubercles (like those in H. dictyophora) present immediately submarginal, with one to five
tips. Margin posterolaterally with a fringe of numerous, long, slender papillae; some papillae swol-
len distally, possibly due to preservation. Dorsal cirri with large, cylindrical cirrophore and long,
strongly papillate style with terminal filum. Dorsal tubercles distinct, nodular. Parapodia (Fig.
34 d, 34 e) with large, conical notopodia with short acicular process. Neuropodial lobes of similar
length, prechaetal lobe, subtriangular, terminating to short acicular process, with digitiform supra-
acicular process. Postchaetal lobe bluntly rounded. Noto- and neuroaciculae penetrating epidermis.
Ventral cirri short, papillate, gradually tapering. Nephridial papillae indistinct, from about seg-
ment 7 to 8 onwards. Notochaetae (Fig. 34 f ) numerous, in dense bundles, short to long, slightly
curved, with numerous rows of spines below blunt, unidentate tips. Neurochaetae (Fig. 34 g, 34 h)
numerous, in vertical fans, slightly more slender and longer than notochaetae, falcate, subdistally
with numerous rows of spines below bidentate tips, on which the secondary tooth is large and set
well back from tip; few lower neurochaetae with unidentate tips.
R e m a r k s : The species is very similar to H. dictyophora, but both species clearly differ in the
shape of the tubercles on the elytra. Moreover, the notochaetae are pointed in the latter, whereas
they are blunt in H. hirsuta.
D e p t h r a n g e : So far recorded down to 70 m; elsewhere reported from the intertidal down
to 98 m (Ruff 1995).
D i s t r i b u t i o n : Red Sea: Gulf of Suez, The Gulf. This is the first record of the species from
the Red Sea and The Gulf. Elsewhere: Northeast Pacific: California, Panama (Ruff 1995).
Harmothoe liaoi Barnich, Fiege & Sun, 2004 Fig. 35 a‑k

Harmothoe liaoi Barnich, Fiege & Sun, 2004. — Species Diversity 9: 307‑309, fig. 4.
Harmothoe boholensis Fauvel 1919. — Archives de Zoologie Expérimentale et Générale 58: 332‑334; Fauvel 1955, Annales de
l’Institute Océanographique Paris 30: 102; Ben-Eliahu 1972 b, Israel Journal of Zoology 21: 196 [not Grube, 1878].
Harmothoe gilchristi Amoureux et al. 1978. — Israel Journal of Zoology 27: 65 [not Day, 1960].
Ty p e l o c a l i t y : Yezhu Island, Yalong Bay, Hainan Island, South China Sea.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Yalong Bay, Yezhu Island, 3‑7 m, in dead coral, dive, coll. D. Fiege & R. Sun,
20 March 1992, IOCAS. — S u e z C a n a l : 1 spm., as Harmothoe boholensis, left of MBL, on coral rock, 10 cm, coll. Hebrew
University Expeditions to the Suez Canal, 17 September 1970, HUJ Poly 497; 1 spm. as Harmothoe boholensis, HUJ Poly 485.
— R e d S e a : microscope slide, as Harmothoe gilchristi, “Mer Rouge”, 1973 [most probably from specimen reported in Amoureux
et al. (1978): Gulf of Aqaba, from artificial lagune at Elat, amongst algae], IEA Y98’; 1 spm., Sudan, Sanganeb Atoll, S‑jetty, east
side, 1 m, from dead Stylophora pistillata, coll. 4 April 1991, SMF 13540; 1 spm., Sudan, Port Sudan, Wingate Reef, 5 m, from
living Pocillopora sp., coll. D. Fiege, 20 September 1992, SMF 13539; 1 spm., Sudan, Sanganeb Atoll, S‑jetty, 2 m, from living
(table coral) Acropora sp., coll. D. Fiege, 30 September 1992, SMF 13534; 1 spm. Sudan, Sanganeb Atoll, SE reef flat, from dead
Stylophora pistillata, coll. D. Fiege, 26 September 1992, SMF 13533; 2 spms. + microscope slide, Sudan, Port Sudan, Wingate Reef,
5 m, from dead corals, coll. D. Fiege, 22 September 1992, SMF 13535; 1 spm., Sudan, Sanganeb Reef, S‑jetty, 20 m, from living
Stylophora sp., coll. D. Fiege, 27 September 1992, SMF 13537; 2 spms., Sudan, Sanganeb Atoll, N‑jetty, east side, 1 m, from dead
Stylophora sp., coll. 5 April 1991, SMF 13538; 1 spm., Sudan, Sanganeb Atoll, S‑jetty, west side, 1 m, from dead Stylophora sp., coll.
6 April 1991, SMF 13541; 1 spm., Assab Bay, Sanabor Island, R/V “Akademik Petrovsky”, cruise 9, A. Tzetlin, 14 February 1980,
ZMUM; microscope slide, as Harmothoe boholensis, Abulat Island, 19°58'N/40°7'E, epi- and endo-fauna of Porites II, coll. “Ca-
lypso”, M. Drach, M.M. Cherbonnier & C. Mercier, 1952 [collecting data supplemented according to Fauvel (1955)], IEA A‑65’.
— G u l f o f A d e n : 2 spms., as Harmothoe boholensis, Gulf of Tadjoura, Djibouti, Musha Islands, amongst polyps, coll. Ch.
Gravier, 23 January 1904, MNHN A25; 1 spm., as Harmothoe boholensis, Gulf of Tadjoura, Djibouti, before mouth of Ambouli
river, dredge, 6 m, coll. Ch. Gravier, 1904, MNHN A25; 1 spm., as Harmothoe boholensis, Gulf of Tadjoura, Djibouti, “Météore”
Reef, dredge, 20 m, coll. Ch. Gravier, 29 February 1904, MNHN A25; microscope slide, as Harmothoe boholensis, Gulf of Tadjoura,
Djibouti, from Hircinia sp., coll. Ch. Gravier, 12 January 1904, IEA D.53; probably belonging to MNHN A25; microscope slide,
as Harmothoe boholensis, Gulf of Tadjoura, Djibouti, “Météore” Reef, coll. Ch. Gravier, 1904, IEA D.53’; probably belonging to
MNHN A25. — A r a b i a n S e a : 1 spm., Socotra Island, off Hawlaf dune, 12°40.519'N/54°4.170'E, 4‑5 m, from dead Acropora
cf. valida, coll. M. Apel, 21 March 1999, SMF 13532; 1 spm., Socotra Island, Qualansiyah Bay, 12°41.026'N/53°28.309'E, 3‑5 m,
patchy rocky substrate with gravel and sand, coll. M. Apel, 10 March 1999, SMF 13536.
M e a s u r e m e n t s : Holotype IOCAS: cs (38) 28/6/4 [af (20) 15 + pf (18) 13. HUJ Poly 497: cs (35) 13/5/3 [af (18) 6 +
pf (17) 7]. HUJ Poly 485: cs (36) 12/2.5/1.8. SMF 13540: cs (39) 14/4.5/3 [af (14) 4 + pf (25) 10]. SMF 13539: cs (37) 10/4/3.
SMF 13534: cs (38) 15/4/2.5 [af (14) 5 + pf (24) 10]. SMF 13533: cs (33) 6/2.5/1.5 [af (14) 2.5 + pf (19) 3.5]. SMF 13535: af
(14) 2.5/3/2, af (18) 5/3.5/2.5, mf (5) 1.5, mf (6) 2. SMF 13537: af (22) 4/3/2 + mf (6) 2. SMF 13538: cs (38) 13/5/3.5 [af (16) 5
+ pf (22) 8], cs (38) 14/4/3 [af (16) 5 + pf (22) 9]. SMF 13541: cs (37) 14/4/3. ZMUM: af (16) 3.5/2.5/1.8. MNHN A25: cs (37)
10/4/3, cs (33) 10/5/3. MNHN A25: cs (37) 13/5/3.5. MNHN A25: cs (37) 19/6/4 [af (24) 11 + pf (13) 8]. SMF 13532: cs (38)
12/5/3.5 [af (16) 5 + pf (22) 7]. SMF 13536: cs (41) 20/6/4 [af (20) 10 + mf (8) 5 + pf (13) 5.
D i a g n o s i s : Anterior pair of eyes anteroventrally. Elytra surface with conical, pointed micro-
tubercles and digitiform papillae; margin with fringing papillae. Neurochaetae all bidentate with
very large secondary tooth well set back from tip.
hexagonal, about as wide as long. Styles of antennae strongly papillate, with a slender terminal
filum. Style of median antenna much longer than ventral style. Palps stout, gradually tapering. Two
pairs of eyes, anterior pair anteroventrally, slightly behind cephalic peaks, posterior pair in front of
hind margin. Tentaculophores with one or two chaetae. Tentacular cirri about as long as median
antenna, shaped as latter. Facial tubercle small. Buccal segment sometimes with small knob. Buc-
cal cirri longer and inserted more proximally on parapodia than following ventral cirri, shaped as
tentacular cirri, but not as long. Elytra (Fig. 35 b) covering dorsum, first pair rounded, following
ones reniform to oval. Surface of elytra covered with low, globular, slightly carinate microtubercles
in anterior part, becoming distinctly taller, conical and pointed in posterior part (Fig. 35 c). Along
anterior border with a small, smooth rim, except in first pair of elytra. Numerous digitiform pa-
pillae scattered over posterolateral surface (Fig. 35 d). Elytra margin posterolateral with fringe of
short, digitiform papillae (Fig. 35 e). Dorsal cirri with cylindrical cirrophore and strongly papillate
style, gradually tapering, without distinct subterminal swelling. Dorsal tubercles small, nodular.
Dorsum with two transverse bands of cilia on each segment. Parapodia (Fig. 35 f, 35 g) with large
conical notopodia. Neuropodia with slightly longer, subtriangular, prechaetal, acicular lobe, with
digitiform supraacicular process. Postchaetal lobe slightly shorter and bluntly rounded. Noto- and
140 T. Wehe
Fig. 35: Harmothoe liaoi (SMF 13540). a: anterior end; b: left elytron from midbody; c: microtubercles; d: papillae on elytra
surface; e: fringing papillae; f: right parapodium from segment 11, anterior view; g: same, posterior view; h: notochaeta; i: upper
neurochaeta; j: middle neurochaeta; k: lower neurochaeta.
neuroaciculae penetrating epidermis. Ventral cirri short, papillate, gradually tapering. Nephridial
papillae indistinct, only visible in a few specimens on median segments. Notochaetae (Fig. 35 h)
numerous, as stout as or slightly stouter than neurochaetae, short to long, curved, with numerous
rows of spines below more pointed, unidentate tips. Neurochaetae (Fig. 35 i, 35 j, 35 k) numerous,
in vertical fans, falcate, with slight swelling and several rows of spines subdistally, with bidentate
tips on which the secondary tooth is large and well set back, and appearing deeply incised.
R e m a r k s : The species is very similar to H. grisea, and there is no unique character to sepa-
rate them. It differs from the latter in that the first pair of eyes is located more anteriorly, there is a
greater size gradient in the microtubercles from anterior to posterior on the elytra, the microtuber-
cles are distinctly pointed and almost thorn-like, the tips of the notochaetae are more slender and
more pointed, and the bidentate tips of the neurochaetae possess a much more prominent second-
ary tooth.
Specimens reported as H. boholensis have proved to belong to this species or to H. grisea. A mi-
croscope slide with parapodia and elytra belonging to Harmothoe gilchristi reported in Amoureux
et al. (1978) was examined. The characters agree exactly with those of Harmothoe liaoi, to which
the specimen is here referred.
D e p t h r a n g e : So far recorded between 0 and 20 m; elsewhere reported from 3‑7 m (Bar-
nich et al. 2004).
D i s t r i b u t i o n : Suez Canal, Red Sea, Gulf of Aden, Arabian Sea: Socotra Archipelago. The
species represents a new record for these areas. Elsewhere: South China Sea (Barnich et al. 2004).
Harmothoe marerubrum n. sp. Fig. 36 a‑l

S p e c i m e n s e x a m i n e d : H o l o t y p e : Tor (= At-Tur), Sinai Peninsula, Gulf of Suez, Red Sea, Egypt, det. Augener,
1920 as Harmothoe sp., coll. Hartmeyer, ZMH P‑24601. P a r a t y p e : same data as holotype, ZMH P‑24602.
M e a s u r e m e n t s : Holotype ZMH P‑24601: cs (36) 13/4/2.5. Paratype ZMH P‑24602: cs (34) 11/3.5/2.
D i a g n o s i s : Anterior pair of eyes anteroventrally. Elytra surface with conical microtubercles
confined to a band on anterior third and scattered micropapillae; margin with a fringe of short
papillae. Neurochaetae bi- and unidentate.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 36 a) rounded, slightly
longer than wide. Style of median antenna broken off in holotype, long in paratype, with slight
subterminal swelling and terminal filum. Styles of lateral antennae much shorter, without sub-
terminal swelling. All antennae strongly papillate. Palps stout, conical, long, gradually tapering.
Epidermis partially detached on palps of holotype. Two pairs of eyes, anterior pair anteroventrally
below cephalic peaks, posterior pair in front of hind margin. Tentaculophores with one chaeta
each. Tentacular cirri as long as median antenna, shaped as latter. Facial tubercle small. Buccal
segment without knob or nuchal fold. Buccal cirri longer and inserted more proximally than
following ventral cirri, shaped as tentacular cirri. Elytra large, leaving middorsum uncovered on
holotype and covered on paratype. First pair rounded (Fig. 36 b), following elytra reniform to oval
(Fig. 36 c). Surface of first pair almost smooth, with very few scattered, globular to conical micro-
tubercles (Fig. 36 d) and few digitiform papillae (Fig. 36 e). Following elytra with microtubercles
confined to anterior and few papillae on remaining surface. Margin of elytra with fringe of numer-
ous very short papillae and few longer papillae in between (Fig. 36 f ). Pigmentation in alcohol as a
horizontal band above and posterior to attachment scar of elytrophore, becoming broader on inner
and smaller on outer margin, resulting in rhomboid-shaped figure above pairs of elytra. Dorsal cirri
with large, conical cirrophore and long, strongly papillate, gradually tapering style. Dorsal tuber-
cles nodular. Parapodia (Fig. 36 g, 36 h) with large, conical notopodia. Neuropodia with longer,
prechaetal, subtriangular, acicular lobe, with digitiform supraacicular process, postchaetal lobe
shorter, bluntly rounded. Noto- and neuroaciculae penetrating epidermis. Ventral cirri short, coni-
cal, papillate, gradually tapering. Nephridial papillae from segment 6 onwards. Notochaetae (Fig.
142 T. Wehe
Fig. 36: Harmothoe marerubrum n. sp. (holotype, ZMH V6140). a: anterior end; b: 1st left elytron; c: left elytron from midbody;
d: microtubercles; e: papillae on elytra surface; f: fringing papillae; g: right parapodium from segment 18, anterior view; h: same,
posterior view; i: notochaeta; j: upper neurochaeta; k: middle neurochaeta; l: lower neurochaeta.
36 i) numerous, slightly stouter than neurochaetae, short to long, slightly curved, with numerous
rows of spines below blunt, unidentate tips. Neurochaetae (Fig. 36 j, 36 k, 36 l) numerous, in ver-
tical fans, long, falcate, subdistally with slight swelling and numerous rows of spines below mainly
bidentate tips; some upper and lower neurochaetae with unidentate tips.
R e m a r k s : The species is very similar to H. waahli (Kinberg, 1856) in the appearance and ar-
rangement of the microtubercles on the elytra and the elytral pigmentation. The neurochaetal tips
are also of the same type. Nevertheless, Augener (1922 a) as well as Pettibone (1993 b) examined
the type specimen of the latter species, and both concluded that there is no fringe of papillae on
the elytra, and, furthermore, no digitiform papillae on the surface were mentioned. Since both of
these two characters were clearly observed on the specimens examined, they are here described as a
new species.
D i s t r i b u t i o n : Red Sea: Gulf of Suez.
E t y m o l o g y : The name refers to the sea where the species was found; Latin mare = sea, Latin
ruber = red.
Hermadion Kinberg, 1856

Hermadion Kinberg, 1856. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 12(9&10): 386.
Hermadion. — Pettibone 1953, Some scale-bearing polychaetes of Pudget Sound and adjacent waters: 20.
Ty p e s p e c i e s : Hermadion magalhaensi Kinberg, 1856.
D i a g n o s i s (emended): Body flattened dorsoventrally, with up to 65 segments. Prostomium
with cephalic peaks. Lateral antennae inserted ventrally, with small ceratophores. Styles of anten-
nae papillate, lateral styles much shorter than median. Palps stout, gradually tapering. Two pairs of
moderate to large eyes, anterior pair on more or less widest part of prostomium, posterior pair in
front of hind margin. Tentaculophores with chaetae. Second (buccal) segment without nuchal fold,
with buccal cirri longer and inserted more proximally than following ventral cirri. Elytra 15 pairs
on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, leaving posterior end of body uncovered.
Elytra surface weekly ornamented with microtubercles, margins with fringing papillae. Dorsal cirri
with cylindrical cirrophore and papillate style. Dorsal tubercles nodular. Notopodia large, with
acicular process. Neuropodia with longer, prechaetal, acicular lobe with long digitiform acicular
process, and shorter postchaetal lobe. Tips of notoaciculae penetrating epidermis, those of neu-
roacicula not. Notochaetae arranged in a hemispherical, dorsally directed tuft, as stout as or stouter
than neurochaetae, with faint rows of spines and blunt, unidentate tips. Neurochaetae with faint
rows of spines subdistally below blunt, unidentate tips.
R e m a r k s : The best diagnosis was given by Pettibone (1953). In order to make possible a
better comparison with other genera, especially with Robertianella McIntosh, 1885, the diagnosis
was emended for the following characters. Styles of antennae and cirri are papillate in Hermadion,
whereas they are smooth in Robertianella, whilst the latter genus has bill-shaped neurochaetae in
contrast to pointed, unidentate neurochaetae in Hermadion.
Hermadion africanus Hartman, 1974 Fig. 37 a‑i

Hermadion africanus Hartman, 1974 a. — Journal of the Marine Biological Association of India 16(2): 205‑206, fig. 3.
Ty p e l o c a l i t y : Mozambique, Western Indian Ocean.
S p e c i m e n s e x a m i n e d : S y n t y p e : Portuguese East Africa, Mozambique Channel, International Indian Ocean Ex-
pedition, 22°30'S/36°07 E, 859‑960 m, with hexactinellid sponge spicules, coll. “Anton Bruun”, 1 October 1964, LACM-AHF
POLY 1179. S y n t y p e : Gulf of Oman, International Indian Ocean Expedition, 23°35'N/58°49'E, 121‑124 m, coll. “Anton
Bruun”, 3 December 1963, USNM 58220 [collecting data for both specimens supplemented according to Griffin (1974)].
144 T. Wehe
Fig. 37: Hermadion africanus (SMF 13635). a: anterior end; b: detached elytron; c: thorn-like tubercles in dorsal and lateral view;
d: papillae on surface of elytra; e: fringing papillae; f: right parapodium from segment 24, anterior view; g: same, posterior view; h:
notochaeta; i: neurochaeta.
— R e d S e a : 31 spms., “Meteor” cruise No. 5/2, MINDIK, 22°35.2'N/36°45.9'E to 22°34.8 N/36°46.2'E, 772‑779 m, beam-
trawl, 7 February 1987, SMF 13635; 2 spms., “Meteor” cruise No. 5/2, MINDIK, 19°52.2'N/37°35.3'E to 19°53.4 N/37°35.1'E,
747‑778 m, beamtrawl, 1 March 1987, SMF 13614; 2 spms., “Meteor” cruise No. 5/2, MINDIK, 22°04, 2'N/37°10.0'E
to 22°05.0 N/37°09.3'E, 600 m, beamtrawl, 9 February 1987, SMF 13610; 1 spm., “Meteor” cruise No. 5/2, MINDIK,
22°15.4'N/37°44.7'E to 22°14.4 N/37°45.0'E, 1095‑1116 m, beamtrawl, 8 February 1987, SMF 13615; 2 spms., “Meteor” cruise
No. 5/2, MINDIK, 19°18.2'N/38°15.5'E to 19°18.2 N/38°14.8'E, 537‑681 m, beamtrawl, 28 February 1987, SMF 13611;
1 spm., “Meteor” cruise No. 5/2, MINDIK, 22°08, 4'N/37°28.9'E to 22°09.2 N/37°29.5'E, 827‑863 m, beamtrawl, 2 March
1987, SMF 13613; 2 spms., “Meteor” cruise No. 5/2, MINDIK, 22°52.7'N/37°03.4'E to 22°52.5 N/37°03.5'E, 880‑884 m,
beamtrawl, 7 February 1987, SMF 13612 [collecting data for all specimens supplemented according to Nellen et al. 1996 b].
M e a s u r e m e n t s : Syntype LACM-AHF POLY 1179: af (42) 13/3.5/2. Syntype USNM 58220: cs (47) 20/5/3. SMF
13635: cs (38) 8/4/3, cs (35) 8/3/2, cs (32) 6/3/2, af (25) 7/4/2.3, af (33) 8/3.5/2.5, af (16) 3/3/2, af (20) 6/3.5/2, af (22)
7/3.5/2.8, af (28) 7/3/2, af (16) 4/3/2, af (28) 7/3.5/2.8, af (20) 5/3/2, af (15) 4/3/2, af (20) 5/3/2, af (18) 5/3/2, af (22) 5/3/2, af
(18) 5/3/2, af (31) 10/3/2, af (31) 8/3/2, af (20) 5/3/2, af (18) 5/3/2, af (16) 4/3.5/2.8, af (16) 4/3.5/2.8, af (32) 10/3/2, af (23)
7/3.5/2.5, af (25) 7/3.5/2.8, af (18) 5/3/2, af (16) 4/3.5/2.8, af (14) 2.5/2.2/1.5, af (16) 4/3/2, af (17) 4/3/2, af (15) 4/3/2, mf (7)
2, mf (9) 3, pf (16) 4, pf (17) 5, pf (17) 3.5, pf (7) 18. SMF 13614: af (16) 3/2.5/1.8, af (16) 4/3.5/2.8, mf (4) 1.5. SMF 13610: af
(20) 6/3.5/2.2, af (16) 4/2.5/1.8, mf (11) 4, mf (17) 5. SMF 13615: af (21) 6/3/2.2. SMF 13611: af (15) 4/3/2, af (25) 8/2.5/1.8.
SMF 13613: af (18) 6/3.5/2.5. SMF 13612: af (18) 5/3.2/2.2, af (20) 3.5/2/1.2, mf (10) 3.
D i a g n o s i s : Elytra with thorn-like microtubercles. Noto- and neurochaetae faintly spinous,
with blunt, unidentate tips.
D e s c r i p t i o n : Body tapering posteriorly, dorsoventrally flattened, sometimes rounded
in cross-section. Segments up to 47 (USNM 58220), complete specimens from “Meteor” cruise
5 with 35 and 38 segments. Prostomium (Fig. 37 a) wider than long, oval to hexagonal, with ce-
phalic peaks (inconspicuous on type specimens, but very distinct on all other specimens). Median
antenna with large cylindrical cirrophore and slender, papillate style. Lateral antennae with broad
ceratophore and a shorter papillate style than in median antenna. Palps conical, gradually tapering,
smooth. Two pairs of moderate to large eyes, anterior pair on more or less widest part of prosto-
mium, posterior pair in front of hind margin. Tentaculophores with up to four chaetae each and
long, slender, papillate tentacular cirri, tapering to terminal filum. Facial tubercle distinct. Buccal
segment with buccal cirri longer and inserted more proximally than following ventral cirri, shaped
as tentacular cirri. Elytra (Fig. 37 b) oval; surface with thorn-like microtubercles (Fig. 37 c) gradu-
ally increasing in size from anterior to posterior, leaving anterior part of elytra smooth (with the
exception of anteriormost elytra), and scattered, digitiform, slender papillae (Fig. 37 d); margin
with fringe of short papillae posterolaterally (Fig. 37 e). Dorsal cirri with small, slender cirrophore
and slender, papillate style tapering to terminal filum, extending beyond neurochaetae. Dorsal
tubercles small, nodular. Parapodia (Fig. 37 f, 37 g) with large, conical notopodia with distinct
acicular process. Notoacicula penetrating epidermis. Neuropodia with subtriangular, prechaetal,
acicular lobe with long acicular process (more prominent in type specimens), and shorter, rounded
postchaetal lobe. Neuroacicula not penetrating epidermis. Ventral cirri short, with few papillae,
gradually tapering. Nephridial papillae inconspicuous, visible on a few median segments on type
specimens. Notochaetae (Fig. 37 h) arranged in a hemisphere above parapodia, giving the speci-
mens a hedgehog-like appearance; as stout as or stouter than neurochaetae, with median swelling
and few rows of faint spines medially, tips blunt, unidentate. Neurochaetae (Fig. 37 i) with swell-
ing and few rows of faint spines subdistally, below blunt tips; some neurochaetae appear to have
very faintly notched tips.
R e m a r k s : The description given is based on the non-type specimens collected by the “Me-
teor” cruise 5, unless stated otherwise, since the type specimens have been very accurately described
and figured by Hartman (1974 a). The non-type specimens exhibit the same characters as the syn-
types, with the exception of the more prominent neuropodial acicular processes in the type speci-
mens. In the non-type specimens, some tips of the neurochaetae appear to be very faintly notched.
However, such differences may be due to size and are not considered to be a specific difference. A
few specimens are infested with parasitic copepods.
B i o l o g y & e c o l o g y : The species is a deep-sea form, since it has been collected from sev-
eral different sites at depths from 120 down to over 1100 m.
146 T. Wehe
D e p t h r a n g e : So far recorded between 121 and 1116 m; elsewhere reported from

859‑960 m (Hartman 1974 a).
D i s t r i b u t i o n : Red Sea, Gulf of Oman. H. africanus is reported for the first time from the
Red Sea. Elsewhere: Indian Ocean: Mozambique (Hartman 1974 a).
Hololepidella Willey, 1905

Hololepidella Willey, 1905. — Ceylon Pearl Oyster Fisheries Supplementary Reports 30: 251.
Hololepidella. — Pettibone 1969 c Proceedings of the Biological Society of Washington 82: 49; Hanley & Burke 1991 a, Mé-
moires du Muséum National d’Histoire Naturelle (A) 151: 28.
Ty p e s p e c i e s : Hololepidella commensalis Willey, 1905.
D i a g n o s i s (emended): Body flattened dorsoventrally, tapering posteriorly, with up to
90 segments. Prostomium with subtriangular lobes, with or without distinct cephalic peaks. Lat-
eral antennae inserted ventrally, with broad ceratophores and much more slender styles. Styles usu-
ally smooth, lateral shorter than median style. Palps stout, gradually tapering. Two pairs of eyes on
posterior half of prostomium. Tentaculophores with or without chaetae. Second (buccal) segment
without nuchal fold, with buccal cirri longer and inserted more proximally than following ventral
cirri. Elytra numerous pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 31, 34, 36, 38, alter-
nating to end of body (some irregularity in posterior region might occur). Elytra soft, translucent,
surface smooth or with scattered microtubercles, without fringing papillae. Dorsal cirri with cylin-
drical cirrophore and smooth style. Dorsal tubercles distinct, nodular. Notopodia of moderate size,
with long acicular process. Neuropodia with longer, prechaetal, acicular lobe, tapering to acicular
process, and shorter postchaetal lobe. Notochaetae moderate in number, short to long, as stout or
stouter than neurochaetae, slightly curved, with several widely spaced teeth along convex edge and
blunt, unidentate tips. Neurochaetae longer, more slender, straight, with spinous rows below uni-
dentate or notched, bidentate tips.
R e m a r k s : The generic diagnosis is emended for the presence of chaetae on the tentaculo-
phores and cephalic peaks on the prostomium. The presence of chaetae on the tentaculophores
may vary within the genus. In the diagnosis given by Pettibone (1969 c), the tentaculophores
were described as achaetous. This was followed by Hanley (1989), who accordingly placed the
genus in the Arctonoinae. Hanley & Burke (1991 a) subsequently examined chaetae on the in-
dividuals from the Chesterfield Islands and once again referred the genus to the Harmothoinae (=
Polynoinae). However, the tentaculophores are stated to be “usually without setae” in the diagnosis
provided by the latter authors. As all of the specimens examined during this study possess at least
two or three chaetae on the tentaculophores, their presence has to be considered to be the normal
state rather than an exception. Cephalic peaks have been described as indistinct or absent by Pet-
tibone (1969 c), Hanley & Burke (1991 a) or Britayev et al. (1999). This character state seems
to be variable. In almost every specimen examined here, the prostomium is deeply notched and the
prostomial lobes are not rounded anteriorly but taper to distinct anterolateral cephalic peaks.
Hololepidella nigropunctata (Horst, 1915) Fig. 38 a‑i

Polynoe nigro-punctata Horst, 1915. — Zoologische Mededeelingen 1: 20; Horst 1917, Siboga-Expeditie 24b: 104, pl. 21: figs
15‑17.
Hololepidella nigropunctata. — Pettibone 1969 c, Proceedings of the Biological Society of Washington 82: 50, fig. 2; Amoureux
et al. 1978, Israel Journal of Zoology 27: 67; Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle
(A) 151: 28‑30, fig. 7; Pettibone 1993 a, Smithsonian Contributions to Zoology 538: 4‑7, figs 2‑3; Britayev et al. 1999,
Cahiers de Biologie Marine 40(4): 365‑366, fig. 13; Barnich et al. 2004, Species Diversity 9: 311‑312.
Harmothoe lunulata ssp. synaptae Amoureux et al. 1978. — Israel Journal of Zoology 27: 65 [not Saint-Joseph, 1888].
Harmothoe minuta Fauvel 1933. — Mémoires de l’Institut d’Égypte 21: 38 [partim, see also Harmothoe grisea; not Potts, 1910].
Fig. 38: Hololepidella nigropunctata (SMF 13607: a, d-i; SMF 13604: b, c). a: anterior end; b: detached elytron; c: microtubercles;
d: left parapodium from segment 14, anterior view; e: same, posterior view; f: notochaeta; g: upper neurochaeta; h: middle neuro-
chaeta; i: lower neurochaeta.
Ty p e l o c a l i t y : Ambon, Indonesia, Indo-West Pacific.

S p e c i m e n s e x a m i n e d : R e d S e a : 2 spms., as Harmothoe minuta, Gulf of Suez, 29°11' to 29°7'30''N/32°55'20''
to 32°59'30''E, 35 m, fine sand, coll. R.Ph. Dollfus, 28 November 1928 [collecting data supplemented according to Billard
(1933)], MNHN A360; 2 spms. as Harmothoe minuta, Gulf of Suez, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m, fine
sand, coll. R.Ph. Dollfus, 1928 [collecting data supplemented according to Billard (1933)], MNHN A403; 3 spms., as Harmothoe
minuta, Gulf of Suez, 28°54' to 28°49'N/32°44' to 32°47'E, 25‑31 m, sand and madreporarians, coll. R.Ph. Dollfus, 8 December
148 T. Wehe
1928 [collecting data supplemented according to Billard (1933)], MNHN A894; 7 spms. as Harmothoe minuta, Gulf of Suez,
29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m, fine sand, coll. R.Ph. Dollfus, 28 November 1928 [collecting data sup-
plemented according to Billard (1933)], MNHN A894; 1 spm. as Harmothoe minuta, Gulf of Suez, 28°54' to 28°49'N/32°44'
to 32°47'E, 25- 31 m, sand and madreporarians, coll. R.Ph. Dollfus, 8 December 1928 [collecting data supplemented according
to Billard (1933)], MNHN A894; 3 spms. as Harmothoe minuta, Gulf of Suez, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E,
35 m, fine sand, coll. R.Ph. Dollfus, 28 November 1928, MNHN A894; microscope slide, as Hololepidella nigropunctata, “Mer
Rouge”, IEA Z1’; microscope slide, as Harmothoe minuta, Gulf of Suez, 29°11' to 29°7'30''N/32°55'20'' to 32°59'30''E, 35 m, fine
sand, coll. R.Ph. Dollfus, 28 November 1928 [collecting data supplemented according to Billard (1933)], IEA R/68’; microscope
slide, as Harmothoe minuta, Gulf of Suez or Aqaba?, coll. R.Ph. Dollfus, IEA R/68; 1 spm., as Harmothoe minuta, Gulf of Aqaba,
29°29' to 29°28'45''N/35 ° to 35°1'15''E, 25 m, sand, coll. R.Ph. Dollfus, 4 February 1929 [collecting data supplemented accord-
ing to Billard (1933)], MNHN A775; 1 spm., Sudan, Sanganeb Atoll, south jetty, 12 m, from dead Pocillopora sp., coll. D. Fiege,
29 September 1992, SMF 13604; 1 spm., Sudan, Sanganeb Atoll, 12 m, from dead and living Pocillopora sp., coll. 1 April 1991,
SMF 13602; 1 spm., Sudan, Sanganeb Atoll, south west tip, 22 m, from living Seriatopora sp., coll. D. Fiege, 3 October 1992, SMF
13607; 1 spm., Sudan, Sanganeb Atoll, south jetty, 50 m, from dead Stylophora sp., coll. D. Fiege, 25 September 1992, SMF 13606;
1 spm., Sudan, Sanganeb Atoll, south jetty, east side, 1 m, from living Stylophora pistillata, 4‑15 April 1991, SMF 13603; 1 spm.,
Sudan, Sanganeb Atoll, south jetty, 12 m, from dead Pocillopora sp., coll. D. Fiege, 29 September 1992, SMF 13601. — G u l f o f
A d e n : 1 spm., “Meteor” cruise No. 5/2, MINDIK, 12°30.9'N/44°47.7'E to 12°31.2 N/44°48.4'E, 76 m, beamtrawl, 16 March
1987 [collecting data supplemented according to Nellen et al. 1996 b], SMF 13605.
M e a s u r e m e n t s : MNHN A360: af (29) 4/2.2/1.5, af (37) 8/2.5/1.5. MNHN A403: cs (43) 6/3/2, af (27) 5/3/2.
MNHN A894: af (26) 6/4/3, af (12) 3/4/3, af (27) 5/3/2, mf (6) 2/3.5/3. MNHN A894: af (14) 1.5/1.5/1. af (9) 1/1.5/1. af (19)
3/2/1.2. af (10) 2/2/1.5. af (20) 3/2/1.2. af (10) 1/1.5/1. af (25) 5/2.5/1.8. MNHN A894: cs (35) 5.5/2/1.5 [af (23) 4 + pf (12)
1.5]. MNHN A894: af (32) 5/2/1.5, af (20) 3.5/2.5/2, af (18) 3/2.5/2. MNHN A775: af (20) 3.5/3/2 + mf (11) 2. SMF 13604:
af (37) 8/3/2. SMF 13602: af (29) 3.5/1.8/1. SMF 13607: cs (36)7/3/2 [af (20) 4 + pf (16) 3]. SMF 13606: af (23) 2.5/2/1. SMF
13603: af (21) 4/2/1. SMF 13601: af (19) 3/1.5/0.8. SMF 13605: af (26) 5/1.5/1.
D i a g n o s i s : Elytra mostly smooth, few scattered microtubercles anteriorly, without fring-
ing papillae. Neuropodia with sharply pointed prechaetal, acicular lobe. Notochaetae with 8 to
13 widely spaced spines along convex border. Neurochaetae more slender, with rows of spines and
notched, bidentate tips.
D e s c r i p t i o n : Body flattened dorsoventrally, tapering posteriorly. Prostomium (Fig. 38 a)
slightly wider than long, lobes subtriangular, tapering to anterolateral cephalic peaks. Median antenna
with comparatively slender cylindrical ceratophore, style broken off on all specimens. Lateral anten-
nae with very broad ceratophores and smooth, short, tapering styles. Palps stout, smooth, conical,
gradually tapering. Eyes small, first pair on more or less widest part of prostomium behind horizontal
midline, posterior pair in front of hind margin. Sometimes ciliated tufts visible below ceratophore
of median antenna, situated on the medial bases of the lateral ceratophores. Tentaculophores with
2 to 3 chaetae. Tentacular cirri with long smooth style, gradually tapering. Facial tubercle small. Buc-
cal cirri longer and inserted more proximally than following ventral cirri, shaped as tentacular cirri.
Elytra mostly detached, where still present overlapping medially and posteriorly. Elytra (Fig. 38 b)
large, round to oval, thin, translucent; surface with leaf-like nervature and few scattered microtu-
bercles (Fig. 38 c) near anterior border; margin without fringing papillae. Dorsal cirri with slender,
cylindrical cirrophore and long, smooth, gradually tapering style extending beyond neurochaetae.
Dorsal tubercles distinct, nodular. Parapodia (Fig. 38 d, 38 e) with conical notopodia tapering to
long acicular process. Neuropodia with longer, triangular, prechaetal, acicular lobe tapering to sharply
pointed acicular process, and shorter, bluntly rounded, postchaetal lobe. Ventral cirri short, slender,
gradually tapering. Nephridial papillae small, if present from about segment 10 onwards. Parapodia
with patches of cilia laterally on cirrophores and dorsal tubercles. Notochaetae (Fig. 38 f) stouter than
neurochaetae, moderate in number, slightly curved, with about 8 to 13 widely spaced teeth along
convex border and blunt, unidentate tips. Neurochaetae more slender, longer, falcate; upper chaetae
(Fig. 38 g) slender with numerous rows of spines and minutely notched tips, middle chaetae (Fig.
38 h) more swollen subdistally with several rows of spines below distinct notched, bidentate tips,
lower chaetae (Fig. 38 i) with lesser rows of spines.
R e m a r k s : Tufts of cilia on the ceratophores of the lateral antennae are described for the first
time. Cilia on the parapodia were not mentioned or illustrated by Hanley & Burke (1991 a), but
were figured by Pettibone (1969 c, 1993 a) and described by Britayev et al. (1999). Most specimens
are incomplete, and those that are complete are rather short with not more than 43 segments.
Harmothoe minuta (Potts, 1910) is doubtful (Barnich & Fiege 2000) and was listed as such in
Wehe & Fiege (2002). All the specimens reported by Fauvel (1933) belong to H. nigropunctata or
to Harmothoe grisea (Ehrenberg & Grube in Grube, 1869).
Specimens of Hololepidella nigropunctata and Harmothoe lunulata ssp. synaptae reported by
Amoureux et al. (1978) were not examined. Both records have been synonymised with H. ni-
gropunctata by Pettibone (1993 a). However, two microscope slides were made available from
the Institut d’Écologie Appliquée, Angers. Parapodia mounted on the slide IEA Z1’ belong to
Hololepidella nigropunctata (except one most probably belonging to Harmothoe grisea), whereas
parapodia and elytra found on the slide IEA Y98 and labelled as “Harmothoe lunulata Delle Chiaje,
mer rouge, 1973” are indeterminable.
B i o l o g y & e c o l o g y : Specimens examined from Sanganeb Atoll were mostly found free-
living on corals, but the species is also known to occur as a commensal with ophiuroids (Pettibone
1969 c, Hanley & Burke, 1991 a, Britayev et al. 1999).
D e p t h r a n g e : So far recorded between 1 and 76 m; elsewhere reported from 1‑65 m
(Hanley & Burke 1991 a, Britayev et al. 1999).
D i s t r i b u t i o n : Red Sea. Elsewhere: All over Indo-West Pacific: Mozambique, Malay Archi-
pelago, South China Sea, Timor Sea, Papua New Guinea, Bismarck Sea, northern Australia, Ches-
terfield Islands, Solomon Islands, Marshall Islands, Hawaii, Marquesa Islands, Hong Kong, Japan
(Hanley & Burke 1991 a, Britayev et al. 1999, Barnich et al. 2004).
Malmgreniella Hartman, 1967

Malmgreniella Hartman, 1967. — Alan Hancock Monographs in Marine Biology 2: 37.
Malmgreniella. — Pettibone 1993 a, Smithsonian Contributions to Zoology 538: 9‑10; Barnich & Fiege 2001, Journal of
Natural History 35: 1121; Barnich & Fiege 2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft
559: 56.
Ty p e s p e c i e s : Malmgreniella dicirra Hartman, 1967.
D i a g n o s i s (emended): Body flattened dorsoventrally, with up to 46 segments. Prostomium
usually without cephalic peaks. Lateral antennae inserted terminoventrally, with broad cerato-
phores. Styles of antennae papillate (rarely smooth), lateral shorter than median style. Palps stout,
gradually tapering. Two pairs of eyes, first pair slightly in front of widest part of prostomium, pos-
terior pair in front of hind margin. Tentaculophores usually with chaetae. Second (buccal) segment
without nuchal fold, with buccal cirri longer and inserted more proximally than following ventral
cirri. Elytra 15 pairs on segments 2, 4, 5, 7, alternating to 23, 26, 29, 32, more or less covering
body, leaving tail uncovered in larger specimens. Elytra surface with or without microtubercles,
macrotubercles and papillae, margins with or without fringing papillae. Dorsal cirri either all with
cylindrical cirrophore and long, often papillate, style, or with two types of styles: short, basally bul-
bous and long tapering styles. Dorsal tubercles nodular. Notopodia of moderate size. Neuropodia
with longer, prechaetal, acicular lobe with or without supraacicular process, and shorter, postcha-
etal lobe. Tips of noto- and neuroaciculae penetrating epidermis. Notochaetae as stout as or stouter
than neurochaetae, with distinct rows of spines and blunt or pointed, unidentate tips. Neurocha-
etae falcate, with fewer rows of spines subdistally below bi- and unidentate tips.
R e m a r k s : The diagnoses by Pettibone (1993 a) and Barnich & Fiege (2001, 2003) are
emended for the possible presence of macrotubercles, as found in M. vesicudenta, and the presence
150 T. Wehe
of two different types of dorsal cirri, as originally mentioned by Hartman (1967) and confirmed
in the species description of M. dicirra — but not considered in the generic diagnosis — by Pet-
tibone (1993 a).
Key to the species of Malmgreniella
1 Elytra with few microtubercles confined to anterior surface of elytra. Balloon-shaped

macrotubercles absent M. murrayensis
– Elytra with crested, balloon-shaped macrotubercles along posterior margin
M. vesicudenta
Malmgreniella murrayensis Pettibone, 1993 Fig. 39 a‑i

Harmothoe lunulata Monro, 1937. — Scientific Reports, Zoology 4(8): 258 [not Delle Chiaje, 1822 [1830]].
Malmgreniella murrayensis Pettibone, 1993 a. — Smithsonian Contributions to Zoology 538: 19‑22, fig. 14.
Ty p e l o c a l i t y : South Arabian Coast, Arabian Sea.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Malmgreniella murrayensis, South Arabian Coast, 18°03'30''N/57°02'30''E,
“John Murray Expedition”, 38 m, net: TD4, bottom: Lithothamnion, in cavity of sponge, coll. 29 October 1933, BMNH
1937.9.2.28.
M e a s u r e m e n t s : Holotype BMNH 1937.9.2.28: cs (35) 8/2.5/2 [af (19) 4 + pf (16) 4].
D i a g n o s i s : Elytra with few, globular microtubercles on anterior surface; margin smooth.
Neurochaetae bidentate, with secondary tooth large and set well back from tip.
hexagonal, about as wide as long. Ceratophores of lateral antennae broad, styles short, papil-
late, gradually tapering to a slender terminal filum; median style broken off. Palps stout, conical,
smooth, abruptly terminating to a slender terminal filum. Two pairs of eyes, anterior pair dorso-
laterally slightly in front of widest part of prostomium, posterior pair in front of hind margin.
Tentaculophores with single chaeta each, tentacular cirri broken off. Facial tubercle small. Buccal
cirri papillate, longer and inserted more proximally than following ventral cirri. Elytra (Fig. 39 b)
mostly detached, reniform to oval; surface mostly smooth, with few globular microtubercles next
to anterior border (Fig. 39 c); margin without fringing papillae. Dorsal cirri with distinct, conical
cirrophore and long, gradually tapering, papillate style. Dorsal tubercles distinct, nodular. Parapo-
dia (Fig. 39 d, 39 e) with large, subtriangular notopodia, tapering to acicular process. Neuropodia
with long, almost triangular, prechaetal, acicular lobe and shorter, rounded postchaetal lobe. Noto-
and neuroaciculae pointed, sometimes penetrating epidermis. Ventral cirri papillate, short, gradu-
ally tapering. Notochaetae (Fig. 39 f ) numerous, as stout as or stouter than neurochaetae, short to
long, curved, with numerous rows of spines below blunt tips. Neurochaetae (Fig. 39 g, 39 h, 39 i)
numerous, in vertical fan, falcate, with numerous rows of spines subterminally below bidentate
tips, with large secondary tooth set well back from tip; upper and lower neurochaetae somewhat
more slender than median neurochaetae.
D e p t h r a n g e : Previously recorded from 38 m.
D i s t r i b u t i o n : Arabian Sea.
Malmgreniella vesicudenta (Hanley & Burke, 1991) n. comb. Fig. 40 a‑o

Harmothoe vesicudenta Hanley & Burke, 1991 a. — Mémoires du Muséum National d’Histoire Naturelle (A) 151: 24‑26, fig. 5.
Ty p e l o c a l i t y : Chesterfield Islands, Coral Sea.
Fig. 39: Malmgreniella murrayensis (holotype, BMNH 1937.9.2.28). a: anterior end; b: detached elytron; c: microtubercles; d: left
parapodium from midbody, anterior view; e: same, posterior view; f: notochaeta; g: upper neurochaeta; h: middle neurochaeta; i:
lower neurochaeta.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Harmothoe vesicudenta, Coral Sea, Chesterfield Island, MNHN POLY TYPE

1338. — R e d S e a : 1 spm., Sudan, Sanganeb Atoll, 22 m, from living Seriatopora sp., 3 October 1992, SMF 13617; single
elytron, Sudan, Sanganeb Atoll, reef flat, from dead Stylophora pistillata, 26 September 1992, SMF 13616.
M e a s u r e m e n t s : Holotype MNHN POLY TYPE 1338: cs (37) 13.5/4.5/3.5. SMF 13617: af (24) 6/4/2.5.
152 T. Wehe
D i a g n o s i s : Elytra with globular microtubercles and balloon-shaped macrotubercles with a

crest of blunt teeth; margin with fringing papillae. Neurochaetae bidentate.
as wide as long. Ceratophores of lateral antennae broad, styles short, papillate, gradually tapering
to a slender terminal filum; median style broken off. Palps stout, conical, abruptly terminating in
a short terminal filum, finely papillate. Two pairs of eyes, anterior pair dorsolaterally on more or
less widest part of prostomium, posterior pair in front of hind margin. Tentaculophores without
chaetae. Only ventral tentacular cirri present, much longer than lateral antennae, papillate, gradu-
ally tapering. Facial tubercle small. Pharynx slightly everted, no papillae or jaws visible. Buccal cirri
(Fig. 40 b, 40 c) all detached. Small rounded elytra (1st pair?), others reniform to oval; surface cov-
ered with globular microtubercles, very small and smooth in anterior part (Fig. 40 d), becoming
gradually larger and more rugose or spinous on posterior surface (Fig. 40 e, 40 f ); submarginally
along posterior margin with balloon-shaped macrotubercles with a crest of blunt teeth (Fig. 40 g);
scattered small digitiform papillae on surface (Fig. 40 h); outer margin with sparse fringe of long
slender papillae (Fig. 40 i). Dorsal cirri with small, conical cirrophore and short, gradually taper-
ing, papillate style. Dorsal tubercles small, nodular. Parapodia (Fig. 40 j, 40 k) with large conical
notopodia, gradually tapering. Neuropodia with long, almost triangular, prechaetal, acicular lobe
and shorter, rounded postchaetal lobe. Noto- and neuroaciculae finely pointed, distinctly penetrat-
ing epidermis. Ventral cirri short, gradually tapering. Nephridial papillae from about segment
12 onwards. Notochaetae (Fig. 40 l) numerous, as stout as or stouter than neurochaetae, short to
long, curved, with numerous rows of spines below blunt tips. Neurochaetae numerous, in vertical
fan, falcate, of two types: supraacicular longer, with few, larger spines subdistally and finely biden-
tate tips (Fig. 40 m); subacicular shorter, with more distinct subterminal swelling and numerous,
faint rows of spines below more distinct bidentate tips (Fig. 40 n, 40 o).
R e m a r k s : The specimens agree in detail with the holotype and the description provided by
Hanley & Burke (1991 a). The species is, however, referred to the genus Malmgreniella because
of the possession of terminoventrally inserted lateral antennae and the absence of a supraacicular
process on the neuropodia.
D e p t h r a n g e : Previously recorded from 22 m; elsewhere reported from 16‑64 m (Hanley
& Burke 1991 a).
D i s t r i b u t i o n : Red Sea. First record since the original description. Elsewhere: Coral Sea:
Chesterfield Islands, Fairway Reefs (Hanley & Burke 1991 a).
Paradyte Pettibone, 1969

Paradyte Pettibone, 1969 a. — Proceedings of the Biological Society of Washington 82: 12‑13.
Paradyte. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 13.
Ty p e s p e c i e s : Polynoe crinoidicola Potts, 1910.
D i a g n o s i s (emended): Body flattened dorsoventrally, elongate, tapering posteriorly, segment
number variable, with up to 50 or more. Prostomium with or without distinct cephalic peaks. Lat-
eral antennae inserted ventrally, with small ceratophores. Styles of antennae smooth, lateral shorter
than median style. Palps stout, gradually tapering. Two pairs of eyes of moderate size, anterior
pair on more or less widest part of prostomium, posterior pair in front of hind margin. Tentacu-
lophores without chaetae. Second (buccal) segment without nuchal fold, with buccal cirri longer
and inserted more proximally than following ventral cirri. Elytra 15 pairs on segments 2, 4, 5, 7,
Fig. 40: Malmgreniella vesicudenta (SMF 13617). a: anterior end; b: smallest detached elytron (1st?); c: elytron from midbody; d:
microtubercles on anterior surface; e: same from median surface; f: same from posterior surface; g: macrotubercles; h: papillae on
surface; i: fringing papillae; j: right parapodium from segment 18, anterior view; k: same, posterior view; l: notochaetae; m: supra
acicular neurochaeta; n: upper subacicular neurochaeta; o: lower subacicular neurochaeta.
154 T. Wehe
..., 23, 26, 29 and 32, leaving posterior end uncovered. Elytra thin, translucent, surface smooth,
margin without fringing papillae. Dorsal cirri with cylindrical cirrophore and smooth style. Dorsal
tubercles inconspicuous. Notopodia small to moderate. Neuropodia with longer, prechaetal, acicu-
lar lobe, and shorter postchaetal one. Notochaetae stouter than neurochaetae, sabre-like, smooth or
with few teeth along convex edge, tips entire or notched. Neurochaetae more slender, of two kinds:
supraacicular slender, with single semilunar pouch below numerous subdistal rows of spines and
slightly hooked, bi- or unidentate tips; subacicular stouter, falcate, with single semilunar, spinous
pouch and unidentate tips.
R e m a r k s : According to Pettibone (1969 a) and Hanley & Burke (1991 a), the supraacicu-
lar neurochaetae have bifid tips. Individuals of P. levis (Marenzeller, 1902) examined here possess
neurochaetae with entire or only very faintly notched tips, and for this reason the above diagnosis
has been emended.
Key to the species of Paradyte
1 Notochaetae mostly smooth, 1 to 3 spines may be present on notochaetae of anterior

segments. Supraacicular neurochaetae with unidentate tips P. levis
– Notochaetae with 1 to 6 spines. Supraacicular neurochaetae with notched tips
P. crinoidicola
Paradyte crinoidicola (Potts, 1910) Fig. 41 a‑h

Polynoe crinoidicola Potts, 1910. — The Transactions of the Linnean Society Series 2, Zoology 8: 337, pl. 18: fig. 10, pl. 21: figs
39‑41.
Paradyte crinoidicola. — Pettibone 1969 a Proceedings of the Biological Society of Washington 82: 13‑16, fig.7; Hanley &
Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 13‑15, fig. 1.; Britayev et al. 1999, Cahiers
de Biologie Marine 40(4): 369‑371, figs 24‑29; Barnich et al. 2004, Species Diversity 9: 312.
Adyte crinoidicola. — Hanley 1984, The Beagle. Occasional papers of the Northern Territory Museum of Arts and Sciences 1(10):
88‑90, fig. 1.
Scalisetosus longicirrus Fishelson & Rullier, 1969. — Israel Journal of Zoology 18: 54‑55 [not Schmarda, 1861].
Ty p e l o c a l i t y : Maldive Archipelago, Indian Ocean.
S p e c i m e n s e x a m i n e d : R e d S e a : 2 spms., det. as Scalisetosus sp. var.?, Suez, Egypt, on Comatus sp., coll. E. Ban-
nwarth [1912/13 according to Schäfer (1963)], ZMH PE-166; 1 spm., Gulf of Aqaba, Elat, coll. Diesel, SMF 13638; microscope
slide, as Scalisetosus longicirrus, Gulf of Aqaba, Eilat, coll. L. Fishelson, 21 January 1966, IEA Y.29’.
M e a s u r e m e n t s : ZMH PE-166: cs (51) 19/2.5/2.2 [af (36) 14 + pf (15) 5], pf (27) 12/4/3.5. SMF 13638: af (14)
3.5/2.2/1.8 + mf (31) 12.
D i a g n o s i s : Notochaetae with 1 to 6 teeth along convex border, with notched or blunt tips.
Neurochaetae with semilunar pouches; supraacicular with bidentate tips; subacicular neurochaetae
smooth or faintly spinous.
D e s c r i p t i o n : Body flattened dorsoventrally. Prostomium (Fig. 41 a) rounded, about as
wide as long, lobes bluntly rounded, without cephalic peaks. Style of median antenna broken off,
styles of lateral antennae slender, smooth, gradually tapering. Palps stout, smooth, conical, gradu-
ally tapering. Two pairs of eyes, anterior pair on more or less widest part of prostomium, posterior
pair in front of hind margin. Tentacular cirri shaped as lateral antennae, but longer. Facial tubercle
indistinct. Buccal cirri inserted more proximally, but only slightly longer than following ventral
cirri. Elytra (Fig. 41 b) large, oval, with off-centre attachment scar near lateral border, surface
with leaf-like venation. Dorsal cirri with cylindrical cirrophore, and long smooth style, gradually
tapering and extending beyond neurochaetae. Dorsal tubercles absent. Parapodia (Fig. 41 c, 41 d)
Fig. 41: Paradyte crinoidicola (SMF 13638). a: anterior end; b: detached elytron; c: left parapodium from segment 18, anterior view; d: same,
posterior view; e: dorsally directed notochaeta; f: laterally directed notochaetae; g: supraacicular neurochaeta; h: subacicular neurochaetae.
with notopodia of moderate size, about triangular, with short acicular process. Neuropodia with
enlarged, bluntly rounded to diagonally truncate, prechaetal, acicular lobe, forming a thin fringe
along more stout, inner, triangular part. Neuroacicula not reaching tip of lobe, embedded in stout
156 T. Wehe
part. Postchaetal lobe much shorter, bluntly rounded. Ventral cirri short, slender, gradually taper-
ing. Nephridial papillae from about segment 9 onwards, very small, inconspicuous. Two ciliated
ridges visible dorsally on each segment. Notochaetae (Fig. 41 e, 41 f ) arranged tuft-like, stouter
than neurochaetae, translucent, sabre-like, with 1 to 6 teeth on convex borders (more distinct in
anterior segments than in posterior segments), with blunt (mostly dorsally directed) and notched
(mostly laterally directed) tips. Neurochaetae (Fig. 41 g, 41 h) thinner, of two kinds: upper cha-
etae more slender, with semilunar pouches subdistally and numerous rows of spines below slightly
hooked, bidentate tips; lower chaetae slightly stouter, with semilunar pouches subdistally, without
or only with very faint rows of spines below more strongly hooked, unidentate tips.
R e m a r k s : The specimens reported by Fauvel (1957, 1958) and Amoureux et al. (1978)
could not be located for examination. Since there is no description provided in the works by Fau-
vel, these records have to remain questionable. Referring to the short description in Amoureux et
al. (1978), it seems most likely that the specimens belong to Paradyte crinoidicola. However, the
occurrence of the species in the Red Sea is confirmed by the specimens examined here.
B i o l o g y & e c o l o g y : The species is known to be associated with crinoids (Hanley 1984,
Britayev et al. 1999). Two of the specimens examined were found on Comatus sp. (= most prob-
ably Comanthus) according to label information.
D e p t h r a n g e : Unknown; elsewhere reported between 2.5 and 88 m (Hanley 1984,
Hanley & Burke 1991 a, Britayev et al. 1999).
D i s t r i b u t i o n : Red Sea. Elsewhere: All over Indo-West Pacific: i.a. South Africa, Maldives,
Malay Archipelago, Hong Kong, South China Sea, Papua New Guinea, Bismarck Sea, Australia,
Japan, Caroline -, Marshall -, and Solomon Islands (Hanley 1984, Hanley & Burke 1991 a,
Britayev et al. 1999, Barnich et al. 2004).
Paradyte levis (Marenzeller, 1902) Fig. 42 a‑h

Scalisetosus levis Marenzeller, 1902. — Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie
der Wissenschaften 72: 575‑576, pl. 3: figs 12, 12 A; Fauvel 1933, Mémoires de l’Institut d’Égypte 21: 38‑40, fig.1; Hart-
man 1974 a, Journal of the Marine Biological Association of India 16(1): 207‑209.
Paradyte levis. — Imajima 1997, National Science Museum Monographs 13: 6‑7, figs 3, 4.
Ty p e l o c a l i t y : Kagoshima, Nagasaki (Japan) (and/or ?) Hongkong (China), Northwest Pacific.
S p e c i m e n s e x a m i n e d : R e d S e a : 1 spm., Gulf of Suez, coll. R.Ph. Dollfus, 26 December 1928, MNHN A775;
2 spms., as Paradyte crinoidicola, Gulf of Suez, 28°14 N/33°23'E, 22 m, muddy sand and madreporarians, on sponges, coll. R.Ph.
Dollfus, 24 December 1928 [collecting data supplemented according to Billard (1933)], MNHN A894; 1 spm., Gulf of Suez,
27°52'45'' to 27°48'45''N/33°45'15'' to 33°46'30''E, 70 m, muddy sand, coll. R.Ph. Dollfus, 28 December 1928 [collecting data
supplemented according to Billard (1933)], MNHN A360; microscope slide, as Scalisetosus (crinoidicola) levis, Gulf of Suez, coll.
R.Ph. Dollfus (probably belonging to MNHN A775), IEA R/70’; microscope slide, as Scalisetosus (crinoidicola) levis, Gulf of Suez,
27°52'45'' to 27°48'45''N/33°45'15'' to 33°46'30''E, 70 m, muddy sand, coll. R.Ph. Dollfus (probably belonging to MNHN
A360), IEA R/70’. — G u l f o f O m a n : 1 spm., as Scalisetosus levis by Hartman, 23°35'N/58°49'E, International Indian Ocean
Expedition, “Anton Bruun”, 121‑124 m, removed from tubes of Eunice tubifex, 3 December 1963; 13 specimens, 12 paratypes, 2 in
exchange with USNM) (collecting data supplemented according to Griffin (1974)), LACM-AHF POLY 1333; 2 spms., as Scalise-
tosus levis by Hartman, 23°35'N/58°49'E, International Indian Ocean Expedition, “Anton Bruun”, 121‑124 m, removed from tubes
of Eunice tubifex, 3 December 1963 [collecting data supplemented according to Griffin (1974)], USNM 58216; 1 spm., as Scali-
setosus levis by Hartman, 22°05'N/59°47'E to 22°08'N/59°48'E, International Indian Ocean Expedition, “Anton Bruun”, 55 m,
4 December 1963 (exchange from AHF) [collecting data supplemented according to Griffin (1974)], USNM 58217.
M e a s u r e m e n t s : MNHN A775: af (12) 4/4/2 + mf (8) 3 + mf (2) 1 + pf (15). MNHN A894: af (17) 3/3/2, af (14) 6/6/4
+ mf (4) 3 + pf (22) 8. MNHN A360: cs (35) 11/5/3 [af (19) 7 + pf (16) 4]. LACM-AHF POLY 1333: cs (74 ) 31/4/3 [(af (22) 8
+ pf (52) 23]. USNM 58216: af (34) 12/5/3, af (49) 20/4/3. USNM 58217: af (48) 25/6/4 + mf (20) 9.
D i a g n o s i s : Elytra margin sometimes with few sensory papillae. Notochaetae smooth on
middle and posterior segments, tips unidentate. Neurochaetae with semilunar pouch; supra- and
subacicular ones with unidentate tips.
Fig. 42: Paradyte levis (USNM 58216). a: anterior end; b: detached elytra; c: sensory papillae on elytra margin; d: right parapodium from
midbody segment, anterior view; e: same, posterior view; f: notochaetae; g: supraacicular neurochaeta; h: subacicular neurochaeta.
D e s c r i p t i o n : Body flattened dorsoventrally, elongate. Prostomium (Fig. 42 a) deeply

notched anteriorly, rounded to hexagonal, about as wide as long, lobes subtriangular, pointed, with
more or less distinct cephalic peaks. Style of median antenna long, gradually tapering to filiform
tip (present only on specimen MNHN A775). Styles of lateral antennae similar (present on speci-
158 T. Wehe
men LACM-AHF POLY 1333, USNM 58216), but more slender. Palps stout, smooth, conical,
gradually tapering to filiform tips. Two pairs of eyes, anterior pair dorsolaterally on more or less
widest part of prostomium, posterior pair in front of hind margin. Tentacular cirri shaped as an-
tennae, about as long as median antenna. Facial tubercle more or less distinct. Buccal cirri longer
and inserted more proximally than following ventral cirri, shaped as tentacular cirri. Elytra (Fig.
42 b) large, rounded to oval, with off-centre attachment scar near to lateral border. Surface smooth,
margin without fringe of papillae, eventually with few, very small, scattered sensory papillae (Fig.
42 c). Dorsal cirri with cylindrical cirrophore and long, smooth style gradually tapering and ex-
tending beyond neurochaetae. Dorsal tubercles absent. Parapodia (Fig. 42 d, 42 e) with notopodia
of moderate size, subtriangular, with acicular process. Neuropodia with longer, more triangular,
prechaetal, acicular lobe and shorter, rounded, postchaetal lobe. Neuroacicula reaching tip of lobe.
Aciculae of both rami may penetrate epidermis. Ventral cirri short, slender, gradually tapering. No
distinct nephridial papillae. Two ciliated ridges visible dorsally on each segment. Notochaetae (Fig.
42 f ) arranged tuft-like, stouter than neurochaetae, translucent, sabre-like, with blunt, unidentate
tips; in some anterior segments up to three small spines may be present on convex border of noto-
chaetae, in following segments notochaetae smooth, rarely with a single spine. Neurochaetae thin-
ner, of two kinds: supraacicular chaetae (Fig. 42 g) more slender, subdistally with semilunar pouch
and numerous rows of spines below slightly hooked, unidentate or very faintly notched tips; sub-
acicular chaetae (Fig. 42 h) slightly stouter, with semilunar pouch and fewer rows of spines below
more strongly hooked unidentate tips.
R e m a r k s : The specimens agree very well with the original description of P. levis by Maren-
zeller (1902), with the description given by Hartman (1974 a) - who referred the specimens cor-
rectly to Marenzeller’s species -, and also with the remarks on P. levis by Pettibone (1969 a: 16)
under P. tentaculata (Horst, 1915).
The specimen reported from the Arabian Sea in Hartman (1974 a) could not be located for
examination so the occurrence of the species in this area remains unknown.
B i o l o g y & e c o l o g y : The species seems to be associated with other invertebrates, but not
obligatorily with one particular group. It is found i.a. amongst sponges, tube dwelling polychaetes
(see above) and corals (Imajima 1997).
D e p t h r a n g e : Previously recorded between 22 and 124 m; elsewhere reported from
15‑33 m (Imajima 1997).
D i s t r i b u t i o n : Red Sea: Gulf of Suez, ? Arabian Sea, Gulf of Oman. Elsewhere: Japan (Ima-
jima 1997).
Paralepidonotus Horst, 1915

Paralepidonotus Horst, 1915. — Zoologische Mededeelingen 1: 8.
Paralepidonotus. — Hanley 1991, Invertebrate Taxonomy 4: 1053‑1054; Hanley & Burke 1991 a, Mémoires du Muséum Na-
tional d’Histoire Naturelle (A) 151: 30‑31.
Ty p e s p e c i e s : Polynoe ampullifera Grube, 1878.
D i a g n o s i s : Body flattened dorsoventrally, broad, with about 40 segments. Prostomium
without cephalic peaks. Lateral antennae inserted terminoventrally, with broad ceratophores. Styles
of antennae papillate. Palps long, gradually tapering. Two pairs of eyes on posterior half of prosto-
mium. Tentaculophores achaetous. Second (buccal) segment without nuchal fold, with buccal cirri
longer and inserted more proximally than following ventral cirri. Elytra 15 pairs on segments 2,
4, 5, 7, alternating to 23, 26, 29, 32, completely covering dorsum. Elytra large, surface variously
ornamented with spines, tubercles, papillae and vesicles, margin with fringing papillae. Dorsal
cirri with cylindrical cirrophore and papillate style. Dorsal tubercles prominent. Notopodia large,
with short acicular process. Neuropodia with prechaetal lobe longer than postchaetal lobe, with or
without supraacicular process. Conspicuous semilunar ventral lamellae on bases of parapodia next
to nephridial papillae. Notochaetae numerous, with numerous rows of spines and blunt, uniden-
tate tips. Neurochaetae slightly thinner than notochaetae, with several rows of spines below uni- or
bidentate tips.
Key to the species of Paralepidonotus
1 Larger microtubercles branched, with up to four blunt tips P. erythromaris

– Larger microtubercles chisel-shaped, not branched P. ampulliferus
Paralepidonotus ampulliferus (Grube, 1878) Fig. 43 a‑m, Pl. 1 f

Polynoe ampullifera Grube, 1878. — Mémoires de l’Académie Impériale des Sciences de St.-Pétersbourg, série 7, 25(8): 35‑36, pl.
3: fig. 5.
Harmothoe ampullifera. — Fauvel 1911, Archives de Zoologie Expérimentale et Générale, Cinquième Série 4(11): 368‑369;
Wesenberg-Lund 1949, Danish scientific investigations in Iran 4: 253‑255.
Paralepidonotus ampulliferus. — Hanley & Burke 1990, Proceedings of the Third International Marine Biological Workshop:
The Marine Flora and Fauna of Albany, Western Australia 1: 211‑213, fig. 4; Hanley 1991, Invertebrate Taxonomy 4:
1055‑1062, figs 1‑5; Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 76‑77, fig. 15; Barnich et
al. 2004, Species Diversity 9: 315‑316.
Ty p e l o c a l i t y : Bohol Island, Philippines.
S p e c i m e n s e x a m i n e d : 1 S y n t y p e : Philippines, Bohol, Coll. Grube/Semper, ZMB 1138. — R e d S e a : 1 spm.,
Egypt, Suez, coll. E. Bannwarth [1912/13 according to Schäfer (1963)], det. Augener as Harmothoe sp., ZMH V.8130; 1 spm.,
Sudan, Sanganeb Atoll, 10‑15 m, coll. 19 September 1992, SMF 13595. — A r a b i a n S e a : 2 spms., Buleji, Karachi, Pakistan,
intertidal rock, coll. J. Mustaquim, 22 December 2001, SMF 13598; 1 spm., Buleji, Karachi, Pakistan, intertidal rock, coll. J.
Mustaquim, 12 September 1998, SMF 13596; 1 spm., Karangi, Karachi, Pakistan, intertidal mud, coll. J. Mustaquim, 10 Decem-
ber 1991, SMF 13597. — T h e G u l f : 1 spm. + microscope slide, as Harmothoe ampullifera‚ rev. R. Hanley as Paralepidonotus,
coast off Namakdan on the Island Quishm (Tavila), 26°35'14''N/55°29'E, tidewater zone, coll. G. Thorson, 18 April 1937,
ZMUC POL-1347; 1 spm. + microscope slide, as Harmothoe ampullifera, JMWS project, Ras Az-Zawr - Marduma Bay region,
27°22'38''N/49°15'55''E, intertidal, sand-rock, coll. D. Fiege, 28 November 1991, SMF 13600; 5 spms., as Harmothoe ampullifera‚
JMWS project, Jubail Field Research Center, sandbank, coll. D. Fiege, 9 December 1991, SMF 13594; 1 spm., W Jubail, intertidal,
coll. M. Apel, 17 October 1992, SMF 13599; microscope slide, as Harmothoe ampullifera, Bahrain, coll. M.N. Bogoyawlensky,
5 May 1902, IEA P.51; microscope slide, as Harmothoe ampullifera, Bahrain, coll. M.N. Bogoyawlensky, 7 May 1902, IEA P.51’.
M e a s u r e m e n t s : Syntype ZMB 1138: cs (38) 19/6/4. ZMH V.8130: cs (38) 25/11/7. SMF 13595: cs (36) 8/3/2 [af (21)
5 + pf (15) 3]. SMF 13598: cs (40) 27/10/8, cs (39) 23/9/6. SMF 13596: cs (38) 19/7.5/5. SMF 13597: af (35) 26/13/10. ZMUC
POL-1347: cs (38) 23/8/6. SMF 13600: cs (38) 25/10/7. SMF 13594: cs (37) 16/6/4, cs (38) 13/6/4, cs (38) 15/6.5/4.5, cs (38)
20/7/5, af (32) 13/5/3.5. SMF 13599: cs (38) 30/12/9.
D i a g n o s i s : Larger microtubercles on elytra chisel-shaped, not branched.
wider than long. Styles of antennae papillate, moderate in length, without subterminal swelling,
tapering to terminal filum, median style longer than lateral styles. Palps stout, long, gradually
tapering. Two pairs of eyes, anterior pair dorsolaterally on widest part of prostomium in front of
horizontal midline, posterior pair dorsally in front of hind margin. Tentaculophores achaetous.
Tentacular cirri shaped as median antenna, similar in length. Facial tubercle distinct. Buccal cirri
(Fig. 43 b) large, covering dorsum completely. First pair rounded, following styles slightly reniform
to oval, more posterior broad reniform to rounded. Surface usually near posterior and outer margin
with large, balloon-shaped vesicles or ampullae (Fig. 43 c). Surface covered with microtubercles,
smaller and knob-like in anterior part, becoming gradually larger and chisel-shaped (Fig. 43 d) in
posterior part. Digitiform papillae scattered over surface (Fig. 43 e) and forming lateral fringe (Fig.
160 T. Wehe
Fig. 43: Paralepidonotus ampulliferus (ZMUC POL-1347: a-f, j-m; SMF 13598: g, h). a: anterior end; b: detached elytron; c: am-
pullae on elytra; d: chisel-shaped microtubercles; e: papillae on surface; f: fringing papillae; g: left parapodium from segment 22,
anterior view; h: same, posterior view; i: ventral lamellae; j: notochaeta k: upper neurochaeta; l: middle, bidentate neurochaetae; m:
lower neurochaeta.
43 f ). Elytra surface greyish in specimens preserved in ethanol with ampullae dark, if pigment not
faded. Dorsal cirri with long, cylindrical cirrophore and long, strongly papillate, gradually tapering
style extending beyond neurochaetae. Dorsal tubercles prominent, globular. Parapodia (Fig. 43 g,
43 h) with conical notopodia of moderate size with short acicular process. Neuropodia with longer,
triangular, prechaetal, acicular lobe and shorter, bluntly rounded, postchaetal lobe. Aciculae some-
times penetrate epidermis. Ventral cirri short, slender, gradually tapering to filiform tip. Nephridial
papillae distinct from about segment 6 onwards. Prominent ventral lamellae (Fig. 43 i) on the
bases of each parapodium from segment 4 onwards. Notochaetae (Fig. 43 j) slightly stouter than
neurochaetae, numerous, in dense bundles, short to long, slightly curved, with numerous rows
of spines and blunt, unidentate tips. Neurochaetae (Fig. 43 k, 43 m) longer, straight, subdistally
slightly thickened with numerous rows of spines and mostly unidentate tips; some median neuro-
chaetae also with bidentate tips (Fig. 43 l); rows of spines decreasing in lower neurochaetae.
R e m a r k s : Species identification is difficult without the elytra. According to Hanley (1991),
the absence of a supraacicular process on the neuropodia and the presence of only a few bidentate
neurochaetae would point to P. ampulliferus. However, Hanley’s description of P. erythromaris
was not detailed enough, since bidentate tips are also found in this species on the middle and up-
per neurochaetae (see the description and remarks on P. erythromaris below). For this reason the
occurrence of P. ampulliferus in the Suez Canal (Fauvel 1927) could not be confirmed, since the
specimen deposited in The Natural History Museum, London, (BMNH 1926.11.12.137) has lost
all its elytra, and the microscope slide from the Institut d’Écologie Appliquée, Angers (IEA R/79),
has only parapodia in very bad condition.
The specimen reported as H. ampullifera in Fauvel (1932), collected by the Investigator from
the Arabian Sea, could not be examined and the status of the specimen remains unknown. The
specimen examined from Pakistan waters, however, confirms the occurrence of the species in the
Arabian Sea.
D e p t h r a n g e : Previously recorded between 10 and 15 m; elsewhere from inter- and subti-
dal (Hanley 1991).
D i s t r i b u t i o n : Red Sea, Arabian Sea: Karachi, The Gulf. Elsewhere: All over Indo-West
Pacific: East Africa, India, Indonesia, Malay Archipelago, South China Sea, Philippines, Australia,
New Caledonia, Tonga, Marshall Islands (Hanley 1991, Barnich et al. 2004).
Paralepidonotus erythromaris Hanley, 1991 Fig. 44 a‑k

Paralepidonotus erythromaris Hanley, 1991. — Invertebrate Taxonomy 4: 1062‑1065, fig. 6.
Lepidonotus ampulliferus. — Gravier 1902, Nouvelles Archives du Muséum 4 e série 3: 214‑218, textfig. 219, pl. 7: figs
111‑113, pl. 8: figs 127‑128 [not Grube, 1878].
Ty p e l o c a l i t y : Perim Island, Red Sea.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Perim Island, Red Sea, coll. Dr. Jousseaume, 1894, det. Ch. Gravier, 1901 as
Lepidonotus ampulliferus (= Polynoe ampullifera), MNHN POLY TYPE 1431. — R e d S e a : 1 spm., as Harmothoe sp., R/V
“Akademik Petrovsky”, cruise 9, Assab Bay, Sanabor Island, Montipora sp., coll. A. Tzetlin, 13 February 1980, ZMUM. — G u l f
o f A d e n : 1 spm., Djibouti, Gulf of Tadjoura, coll. Dr. Jousseaume, 1893, det. Ch. Gravier, 1901 as Lepidonotus ampulliferus (=
Polynoe ampullifera), MNHN A21; 2 spms., Djibouti, Gulf of Tadjoura, coll. H. Coutière, 1897, det. Ch. Gravier, 1901 as Lepido-
notus ampulliferus (= Polynoe ampullifera), MNHN A21.
M e a s u r e m e n t s : Holotype MNHN POLY TYPE 1431: cs (38) 16/9/6. ZMUM: cs (38) 13/5/3.5 [af (14) 5 + pf (24) 8].
MNHN A21: cs (38) 22/9/6. MNHN A21: cs (38) 18/6/4, af (19) 11/8/5.5.
D i a g n o s i s : Larger microtubercles on elytra branched, with 1 to 4 blunt tips.
about as wide as long. Styles of antennae papillate, moderate in length, without subterminal swell-
ing, tapering to terminal filum, median style longer than lateral styles. Palps stout, long, gradually
162 T. Wehe
tapering. Two pairs of eyes, anterior pair dorsolaterally on widest part of prostomium in front of
horizontal midline, posterior pair dorsally in front of hind margin. Tentaculophores achaetous.
Tentacular cirri shaped as median antenna, similar in length. Facial tubercle distinct. Buccal cirri
(Fig. 44 b) mostly detached (holotype only with two detached elytra), large, soft, according to size
covering dorsum completely. Surface with globular, balloon-shaped vesicles or ampullae (Fig. 44 c)
on posterior half, mainly near posterior and lateral borders. Surface with microtubercles ranging
from small, blunt knobs to larger, branched tubercles with about one to four tips (Fig. 44 d), the
latter more confined to posterior and lateral surface. Numerous, scattered, small, digitiform to
clavate papillae on surface (Fig. 44 e). Lateral margin with fringe of long, digitiform papillae (Fig.
44 f ). Dorsal cirri with large, cylindrical cirrophore and long, strongly papillate, gradually tapering
style extending beyond neurochaetae. Dorsal tubercles distinct, globular. Parapodia (Fig. 44 g) with
conical notopodia of moderate size, acicular process indistinct. Neuropodia with longer, triangular,
prechaetal, acicular lobe and shorter, bluntly rounded, postchaetal lobe. Sometimes aciculae pen-
etrate epidermis. Ventral cirri short, slender, gradually tapering to filiform tip. Prominent ventral
lamellae on the bases of each parapodium from segment 4 onwards, nephridial papillae from seg-
ments 5 to 7 onwards (indistinct in holotype). Notochaetae (Fig. 44 h) numerous, short to long, as
thick as or slightly thicker than neurochaetae, with numerous rows of spines and blunt, unidentate
tips. Neurochaetae (Fig. 44 i, 44 j, 44 k) longer, with subdistal swelling, mostly unidentate; upper
chaetae with more numerous rows of spines, lower chaetae with fewer rows and fine, unidentate
tips. Few upper or middle neurochaetae bidentate with distinct secondary tooth (often worn off as
in holotype).
R e m a r k s : Four specimens in three different vials and stored in one collection jar (A21) were
located in the MNHN, Paris. According to his remarks, Hanley (1991) knew of 4 specimens but
obviously examined only the single specimen, which he designated as the holotype. None of the
four specimens was labelled as holotype, nor was there any indication that they were examined by
Hanley. However, according to the congruence between the original label and the information
provided by Hanley (e.g. only two remaining elytra with the holotype), there is no doubt which of
the specimens represents the holotype of P. erythromaris.
Hanley emphasised the presence of only unidentate neurochaetae on the holotype, which in
addition to the different elytra characters, distinguishes the species from P. ampulliferus. Indeed,
there are no distinct bidentate neurochaetae on the holotype, but close examination reveals subter-
minal scars where a secondary tooth was most probably inserted. Unfortunately, Hanley did not
examine the remaining specimens on which distinct bidentate neurochaetae are present. P. erythro-
maris and P. ampullaris are therefore very similar and cannot be separated by this character.
D i s t r i b u t i o n : Red Sea, Gulf of Aden.
Subadyte Pettibone, 1969

Subadyte Pettibone, 1969 a. — Proceedings of the Biological Society of Washington 82: 8.
Subadyte. — Hanley & Burke 1991 a, Mémoires du Muséum National d’Histoire Naturelle (A) 151: 15‑16; Barnich & Fiege
2003, Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 559: 76.
Ty p e s p e c i e s : Polynoe pellucida Ehlers, 1864.
D i a g n o s i s : Body flattened dorsoventrally, elongate, tapering posteriorly, segment number
variable, with about 40. Prostomium with or without distinct cephalic peaks. Lateral antennae
inserted ventrally, with small ceratophores. Styles of antennae usually papillate. Two pairs of eyes,
Fig. 44: Paralepidonotus erythromaris (MNHN A21). a: anterior end; b: detached elytron; c: ampullae; d: microtubercles; e: papillae
on elytra surface; f: fringing papillae; g: right parapodium from midbody segment, anterior view; h: notochaeta; i: upper neurocha-
eta; j: middle, bidentate neurochaetae; k: lower neurochaeta.
anterior pair on more or less widest part of prostomium, posterior pair in front of hind margin.
Tentaculophores achaetous. Second (buccal) segment with or without nuchal fold, with buccal
164 T. Wehe
cirri longer and inserted more proximally than following ventral cirri. Elytra 15 (or 16) pairs on
segments 2, 4, 5, 7, ..., 23, 26, 29 and 32 (34), covering body. Elytra with small to large papillae
and without microtubercles, margins with fringing papillae. Dorsal cirri with cylindrical cirro-
phore and papillate style. Dorsal tubercles inconspicuous. Notopodia small, with short acicular
process. Neuropodia incised dorsally and ventrally, with longer acicular lobe with or without cir-
riform process, and shorter, postchaetal lobe. Notochaetae about as stout as neurochaetae, curved,
with rows of semilunar, spinous pouches and blunt or notched tips. Neurochaetae all similar, with
semilunar, spinous pouches below subdistal rows of spines and slightly hooked bidentate tips.
Subadyte albanyensis Hanley & Burke, 1990 Fig. 45 a‑i

Subadyte albanyensis Hanley & Burke, 1990. — Proceedings of the Third International Marine Biological Workshop: The Marine
Flora and Fauna of Albany, Western Australia 1: 204‑208, figs 1, 2.
Scalisetosus pellucidus Hartmann-Schröder 1960. — Kieler Meeresforschungen 16(1): 69 [not Ehlers, 1864].
Scalisetosus fragilis Ben-Eliahu 1972 b. — Israel Journal of Zoology 21: 196 [not Claparède, 1868].
Subadyte pellucida Amoureux 1983 a. — Bulletin du Muséum National d’Histoire Naturelle 4(5): 724 ; Dexter 1987, Israel
Journal of Zoology 34: 132 [not Ehlers, 1864].
Ty p e l o c a l i t y : Quaranup, Albany, Western Australia, Indian Ocean.
S p e c i m e n s e x a m i n e d : S u e z C a n a l : 1 spm., as Scalisetosus fragilis (= pellucida), Egypt, Suez Canal, El Qantara,
coll. Hebrew University Expeditions, 21 June 1967, HUJ Poly 483; 1 spm., as Scalisetosus fragilis (= pellucidus), Egypt, Suez Canal,
Great Bitter Lake E of Deversoir, coll. Hebrew University Expeditions, 13 January 1969, HUJ Poly 484. — R e d S e a : 1 spm., as
Subadyte pellucida, Egypt, Sinai, Gulf of Aqaba, El Hamira, coll. D. Dexter, 11 July 1979, MNHN A896; 4 spms., as Scalisetosus
pellucidus, Egypt, Ghardaqa (=Hurghada), coll. A. Remane & E. Schulz, 29 March 1956, ZMH P‑14001; 6 spms., Sudan, San-
ganeb Atoll, south west tip, 22 m, from living Seriatopora sp., coll. D. Fiege, 3 October 1992, SMF 13637.
M e a s u r e m e n t s : HUJ Poly 483: af (13) 3/3/2 + mf (5) 2 + mf (3) 1. HUJ Poly 484: af (9) 1/1/0, 5. MNHN A896: af
(14) 1.5/1.5/0.8. ZMH P‑14001: cs (18) 1, 8/1.2/0.6, af (12) 2/2/1.2, af (9) 1.5/2/1.2, af (10) 1.9/2/1.2, pf (10) 1.9/2/1.2, pf (13)
2.5. SMF 13637: cs (28) 4/1.5/1, cs (28) 4/1.5/1, af (22) 3/1.2/0.8, af (16) 2/1.5/1, af (18) 2.5/1.5/1, af (13) 2/1.5/1.
D i a g n o s i s : Elytra surface with short to long, digitiform papillae, subdistally constricted,
with terminal knob. Neuropodial acicular lobe without distal process. Notochaetae with up to
seven spinous pouches along convex border.
D e s c r i p t i o n : All individuals in poor condition, tiny and mostly fragmented. Almost all
appendages missing on specimens. Body flattened dorsoventrally. Prostomium (Fig. 45 a) oval,
wider than long, lobes bluntly rounded, without cephalic peaks. Styles of antennae broken off.
Palps smooth, stout, conical, gradually tapering. Two pairs of large eyes (often not visible), anterior
pair on more or less widest part of prostomium, posterior pair in front of hind margin. Everted
pharynx with two semicircles distally of nine soft papillae each and two pairs of chitinous jaws.
Tentacular cirri long, papillate, gradually tapering to filiform tip. Facial tubercle small. Buccal seg-
ment without nuchal fold. Buccal cirri longer and inserted more proximally than following ventral
cirri. Elytra (Fig. 45 b) at least 12 pairs on largest specimen according to elytrophores. Only a few
detached elytra present, small, oval, with off-centre attachment scar. Surface without tubercles,
with several short to long, digitiform, subdistally constricted and terminally knob-like papillae
(Fig. 45 c). Along outer and posterior margin with small clavate or knob-like papillae (Fig. 45 d).
Dorsal cirri with more or less cylindrical cirrophore, and papillate, gradually tapering style. Dorsal
tubercles absent. Parapodia (Fig. 45 e, 45 f ) with notopodia of moderate size, subtriangular, with
short, acicular process. Neuropodia with longer, triangular, prechaetal, acicular lobe and shorter,
bluntly rounded, postchaetal lobe. Ventral cirri short, slender, gradually tapering. Nephridial pa-
pillae distinct from about segment 8 onwards. Notochaetae (Fig. 45 g) as stout as neurochaetae,
translucent, curved, mostly with 4 to 6 spinous pouches on convex border and blunt or weakly
notched tips. Neurochaetae (Fig. 45 h, 45 i) longer, falcate, with single semilunar spinous pouch
below several rows of faint spines and bidentate tips.
Fig. 45: Subadyte albanyensis (ZMH P‑14001). a: anterior end; b: detached elytron; c: papillae on surface; d: papillae on margin; e:
parapodium (segment ?), anterior view; f: same, posterior view; g: notochaetae; h: upper neurochaeta; i: lower neurochaeta.
166 T. Wehe
R e m a r k s : All the specimens are very small anterior fragments or complete juveniles. None
of the specimens is complete or has all the elytra present. They are in bad condition and almost
all antennae, cirri and elytra are missing. Nevertheless, based on the remaining characters, espe-
cially the appearance of the few elytra, parapodial lobes and chaetae, the specimens agree very well
with the characters of S. albanyensis described by Hanley & Burke (1990), especially as regards
the papillae on the elytra and the pouches on the notochaetae. Some of the specimens examined
have previously been reported as Subadyte pellucida (Ehlers, 1864). However, as pointed out by
Hanley & Burke (1990), S. pellucida is an Atlantic species and none of the specimens examined
here shows the characters that are typical for S. pellucida, such as 20 pouches on the notochaetae or
ovoid papillae on the elytra. According to Hanley & Burke (1991 a), who discussed the characters
of the six species so far described in the genus, S. albanyensis is closest to S. chesterfieldensis Hanley
& Burke, 1991 a, but differs from the latter by the presence of more numerous pouches on the
notochaetae and longer papillae on the elytra, which is confirmed here.
Specimens reported from the Red Sea by Kiseleva (1971), Amoureux et al. (1978) and
Amoureux (1983 b), as listed under S. pellucida in Wehe & Fiege (2002), could not be located for
examination, and so the status of these specimens remains unknown.
B i o l o g y & e c o l o g y : Associated with sponges and sea grass (Hanley & Burke 1990).
D e p t h r a n g e : Previously recorded from 22 m; elsewhere reported from intertidal to 3 m
D i s t r i b u t i o n : Suez Canal, Red Sea. First record since the original description. Elsewhere:
Albany, Western Australia (Hanley & Burke 1990).
Uncopolynoinae n. subfam.
D i a g n o s i s : Body very small, fragile, with at least 40 segments (probably several more in
complete specimens). Prostomium without cephalic peaks, with three antennae. Median antenna
with large ceratophore in small anterior notch of prostomium, lateral antennae without cerato-
phores, small papilla-shaped styles inserted terminally on prostomial lobes. Palps stout, conical.
Two pairs of eyes present. Tentaculophores without chaetae. Second (buccal) segment without
nuchal fold. Styles of all appendages smooth. Elytra at least 12 pairs on segments 2, 4, 5, 7, al-
ternating to 23, following insertion unknown. Elytra surface smooth, margin without fringing
papillae. Dorsal cirri with smooth styles. Dorsal tubercles inconspicuous. Notopodia absent. Neu-
ropodia deeply incised in anteriormost parapodia with prominent lobes; lobes inconspicuous on
following segments. Notochaetae absent. Neurochaetae very few, of three types: hooks, clavate and
bidentate neurochaetae. Pygidium with a pair of anal cirri.
R e m a r k s : The numbers of the segments and elytra, and the arrangement of the latter in
complete specimens, remain unknown. Hartmann-Schröder (1960) reported an anterior frag-
ment of 44 segments and the elytra in an alternate arrangement behind segment 23. This fragment
is obviously damaged, since it could not be found. An anterior fragment of 25 segments, how-
ever, had two cirrigerous segments on 24 and 25, contradicting the observation by Hartmann-
Schröder.
Fauchald (1977) placed the species in the Harmothoinae (= Polynoinae). However, based on
the currently accepted concept of subfamilies, the characters exhibited in Uncopolynoe corallicola
do not allow this species to be classified within any of them. The combination of a weekly bilobed
prostomium, the presence of three antennae, the terminally inserted, papilla-like lateral antennae
without ceratophores, the tentaculophores without chaetae, the complete absence of notopodia
and notochaetae, the neuropodia not incised and three different types of neurochaetae including
clavate ones, is unique amongst Polynoidae. Even though no complete specimen is known, it ap-
pears legitimate to erect a new subfamily based on the available characters.
Uncopolynoe Hartmann-Schröder, 1960

Uncopolynoe Hartmann-Schröder, 1960. — Kieler Meeresforschungen 16(1): 69.
Ty p e s p e c i e s : Uncopolynoe corallicola Hartmann-Schröder, 1960.
D i a g n o s i s : As for subfamily.
Uncopolynoe corallicola Hartmann-Schröder, 1960 Fig. 46 a‑d

Uncopolynoe corallicola Hartmann-Schröder, 1960. — Kieler Meeresforschungen 16(1): 70‑71, pl. 1: figs 1, 6, pl. 3: figs 2‑5;
Amoureux et al. 1978, Israel Journal of Zoology 27: 72; Amoureux 1981, Rapports et Procès-Verbaux des Réunions. Com-
mision Internationale pour l’ exploration scientifique de la mer Méditerranée Monaco 27(2): 207; — Amoureux 1983 b,
Cahiers de Biologie Marine 24: 364, 368.
S p e c i m e n s e x a m i n e d : H o l o t y p e : Gubal (=Djubal), Gulf of Suez, Red Sea, Egypt, 1 m, from alcyonarian coral,
coll. S. Gerlach, 29 October 1957, ZMH P‑13967; 8 P a r a t y p e s : Gubal (=Djubal), Gulf of Suez, Red Sea, Egypt, 1 m, from
alcyonarian coral, coll. Gerlach, 29 October 1957, ZMH P‑13968. — R e d S e a : 2 spms., Jubal (=Djubal), Gulf of Suez, Red Sea,
Egypt, from alcyonarian coral, coll. R.Ph. Dollfus, 29 December 1928, MNHN A894.
M e a s u r e m e n t s : Holotype, paratypes and non-type individuals all broken into numerous tiny fragments, therefore total
length and segment number unknown. Largest anterior fragment of one paratype: af (25) 4 /0.5/0.4.
D e s c r i p t i o n : Body very small, fragile, subrectangular in cross-section. Prostomium (Fig. 46 a)
weekly bilobed, broad oval to cordiform, wider than long, sometimes withdrawn into second segment.
Three antennae inserted on about same level; median antenna with large, cylindrical ceratophore in an-
terior notch of prostomium, style broken off on all specimens. Lateral antennae without ceratophore,
styles short, papilla-like, smooth, clearly separated from prostomium. Palps stout, conical, gradually ta-
pering. Two pairs of eyes, anterior pair on anterior half of prostomium on more or less widest part, pos-
terior pair on posterior half, slightly more narrowed. Small sac-like pharynx everted in some specimens,
no papillae or jaws visible. Tentacular cirri mostly missing, long and smooth if present, gradually taper-
ing. No facial tubercle. Buccal cirri not much longer than following ventral cirri, but inserted more
proximally. Most elytra broken off and missing. Elytra oval, soft, smooth, translucent, with off-centre
attachment scar (Fig. 46 a). Dorsal cirri with cylindrical cirrophore; styles mostly broken off, if present
smooth, long, gradually tapering, extending beyond neurochaetae. Dorsal tubercles absent. Parapodia
without notopodia and notochaetae. Neuropodia stout, conical; on segments 2 to 4 with distinct,
rounded pre- and postchaetal lobes, enclosing bases of neurochaetae (Fig. 46 a); following neuropodia
bluntly rounded without neuropodial lobes. Ventral cirri stout, conical, gradually tapering, longer than
neuropodia (except in anteriormost segments). Nephridial papillae large, cylindrical on some paratype
fragments, or absent. Notochaetae absent. Neurochaetae of three kinds: stout, strongly bent hooks
(Fig. 46 b) on segments 2 to 4 (up to four per parapodium); neurochaetae on following segments sub-
distally with a triangular swelling below slightly hooked, bidentate tips (Fig. 46 c), and shorter, curved,
clavate neurochaetae with several rows of spines below bluntly rounded tips (Fig. 46 d); usually a single
bidentate and clavate neurochaeta present in each neuropodium, rarely two of each kind.
R e m a r k s : Although the species was well described and figured by Hartmann-Schröder
(1960), some small additions are necessary. The prostomium and buccal segment are not fused, but
instead the prostomium is withdrawn into the latter. The elytra are smooth. Single noto- as well as
a single neuroaciculae are present. There are no small spines on the subdistal swelling of the falcate
neurochaetae. The segment number may be at least 45, since Amoureux et al. (1978) described a
specimen with 45 segments and Hartmann-Schröder an anterior fragment of 44 segments.
Specimens reported by Amoureux et al. (1978) and Amoureux (1983 b) could not be located
for examination. However, the specimens reported by Amoureux (1981), which were collected by
168 T. Wehe
Fig. 46: Uncopolynoe corallicola (MNHN A894). a: anterior end; b: neurochaetal hook from anterior segment; c: falcate neurochaeta
from midbody segment; d: clavate neurochaeta from midbody segment.
R.Ph. Dollfus in 1928 and have been examined here, clearly belong to this species and there is no
reason to doubt the other identifications.
B i o l o g y & e c o l o g y : Unknown. All specimens were found on alcyonarian corals.
D e p t h r a n g e : So far recorded from 1 m.
D i s t r i b u t i o n : Gubal Island, Northern Red Sea.
Unconfirmed records
Due to the unavailability of specimens, the presence or absence of the following species re-
ported in Wehe & Fiege (2002) could not be confirmed. However, the occurrence of most of
these species in the seas surrounding the Arabian Peninsula seems unlikely, for reasons discussed in
the remarks below.
A recent contribution by Rasheed & Mustaquim (2003) to the polynoid fauna of the coastal
waters of Pakistan, Arabian Sea, became known to the present author during the final preparation
of this manuscript. However, thanks to an earlier donation to the Senckenberg Museum by Prof.
J. Mustaquim, I was able to study several specimens from some of the localities mentioned in the
work by Rasheed & Mustaquim (2003), even though these authors report several more specimens
and species which were not examined during the present study. These records with additional com-
ments are also listed below.
Harmothoe branchiata Hartman, 1974

Harmothoe branchiata Hartman, 1974 a. — Journal of the Marine Biological Association of India 16(1): 203‑204, fig. 2.
R e m a r k s : Hartman (1974 a) reported six specimens of H. branchiata. Only the holotype
from Madagascar could be located, but not the specimens reported from the Arabian Sea. How-
ever, examination of the holotype (LACM-AHF POLY 1337) revealed a misinterpretation of the
name-giving character by Hartman. No parapodium could be observed, as was figured by Hart-
man. Instead, the specimen follows the usual pattern in Harmothoe with distinct, but rather small
dorsal tubercles. It is assumed that the cirrophore of a broken dorsal cirrus or even a broken cirro-
phore and a small dorsal tubercle were confused. The status of this specimen is therefore question-
able, and it may be synonymous with another species.
Harmothoe goreensis (Augener, 1918)

Harmothoe goreensis. — Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 58‑60, fig. 6.
R e m a r k s : This species was previously known only from the North to South-East Atlantic
and the Mediterranean Sea (Barnich & Fiege 2000). The only specimen reported as H. goreensis
from the Red Sea and from the entire Arabian region by Amoureux et al. (1978) was referred to H.
grisea in the present paper. The record by Rasheed & Mustaquim (2003) would extend the distri-
bution of H. goreensis to the Indian Ocean.
Harmothoe imbricata (Linnaeus, 1767)

Harmothoe imbricata. — Hartmann-Schröder 1960, Kieler Meeresforschungen 16(1): 69; Fishelson & Rullier 1969, Israel
Journal of Zoology 18: 52; Amoureux et al. 1978, Israel Journal of Zoology 27: 65‑66.
R e m a r k s : Unfortunately, the descriptions given in the references mentioned above are in-
sufficient and no drawings were provided which might corroborate the occurrence of this species.
Furthermore, of the numerous specimens belonging to the genus Harmothoe studied here, none
were found to belong to H. imbricata (see also Barnich & Fiege 2000). Harmothoe imbricata is
widely reported from the Arctic, the whole northern hemisphere, the Western to the Eastern Medi-
terranean, and the Black Sea (Barnich & Fiege 2003). Records from outside these areas appear
doubtful.
Harmothoe impar (Johnston, 1839)

Harmothoe cf. impar. — Amoureux et al. 1980, Cahiers de Biologie Marine 21(4): 388.
R e m a r k s : Harmothoe impar was only tentatively reported in a brief communication on
polychaetes found among the sponge Fasciospongia cavernosa, without any description being given.
Since the specimen in question was only tentatively identified and the distribution of the species
is similar to that of H. imbricata (Hartmann-Schröder 1996), the occurrence of the species in the
seas surrounding the Arabian Peninsula is unlikely.
Harmothoe spinifera (Ehlers, 1864)

Harmothoe cf. spinifera. — Amoureux et al. 1980, Cahiers de Biologie Marine 21(4): 388.
R e m a r k s : See Harmothoe impar.
Lepidasthenia stylolepis (Willey in Lloyd, 1907)

Lepidasthenia stylolepis Willey in Lloyd, 1907. — Records of the Indian Museum 1: 260.
R e m a r k s : This species was described from the northern part of The Gulf. The type
specimen(s) could not be located. The status and occurrence of the species thus remain uncon-
firmed.
Lepidonotus hedleyi Benham, 1915

Lepidonotus hedleyi. — Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 64, fig. 9.
170 T. Wehe
R e m a r k s : The specimens reported as L. hedleyi by Rasheed & Mustaquim (2003) clearly be-
long to L. purpureus Potts, 1910. The description and figures provided by Rasheed & Mustaquim
(2003) exactly match the syntype and other specimens of L. purpureus studied and described in the
present paper. Hartman (1959) listed L. hedleyi as a synonym of L. purpureus. The occurrence of
L. hedleyi in the Arabian Sea cannot be confirmed here.
Lepidonotus jacksoni Kinberg, 1856

Lepidonotus jacksoni. — Rasheed & Mustaquim 2003, Pakistan Journal of Marine Sciences 12(1): 64‑66, fig. 10.
R e m a r k s : A single specimen from the same location as reported by Rasheed & Mustaquim
(2003), and previously identified and labelled as L. jacksoni by J. Mustaquim, has been identified
here as L. natalensis Day, 1951. The specimens described as L. jacksoni and figured by Rasheed &
Mustaquim (2003) are similar to this specimen as well as to the holotype of L. natalensis examined
in course of our study. It appears that all these specimens might well belong to the same species.
However, the original description and figures of L. jacksoni provided by Kinberg (1856, 1857) are
poor, and subsequent descriptions are also inadaequate and differ from each other, e.g. Augener
(1922), Fauvel (1932, 1953), Day (1967 a). The elytra — the most important character — of the
type specimens have never been figured properly (see Kinberg 1857, Augener 1922). To confirm
whether L. natalensis might be a junior synonym of L. jacksoni, a syntype (SMNH-Type-389) was
examined during this study. Unfortunately, all the elytra are lost on this specimen, making a com-
parison impossible.
Malmgreniella castanea (McIntosh, 1876)

Malmgrenia castanea. — Amoureux et al. 1978, Israel Journal of Zoology 27: 68.
R e m a r k s : The sole record of this species in the Red Sea was reported without providing any
description. Malmgreniella castanea, like the species of Harmothoe discussed above, is only known
to occur outside tropical waters, in the Western Mediterranean and the North East Atlantic. It
might be confused with M. andreapolis, M. darbouxi Pettibone, 1993 a, and M. lunulata (Delle
Chiaje, 1822 [1830]) according to Barnich & Fiege (2003). However, all of these species also oc-
cur only in the Mediterranean Sea or the North East Atlantic. The presence of this species in the
Red Sea is therefore doubtful.
Polyeunoa laevis (McIntosh, 1885)

Polyeunoa laevis. — Amoureux et al. 1978, Israel Journal of Zoology 27: 67.
R e m a r k s : A single specimen was reported from the Red Sea. However, the discription is
inadequate and no drawings were provided. The presence of this species appears highly unlikely,
since it has only been reported from antarctic and subantarctic waters (Pettibone 1969 c).
Polynoe fumigata Grube, 1868

Polynoe fumigata Grube, 1868 b. — Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 18:
630‑631.
R e m a r k s : Polynoe fumigata was described from the Red Sea. The status and occurrence of
this species remains unconfirmed. The type specimen(s) appears to be lost since the species is not
listed by Wiktor (1992) or Hartwich (1993) as present in the collections where Grube worked.
Zoogeography
It is necessary to emphasise that the conclusions presented here on zoogeographic aspects, and
especially on endemism, can only be regarded as preliminary, mainly for the following reasons: (1)
Although based on about 800 specimens collected all around the Arabian Peninsula, the sampling
effort is very sketchy and unevenly spread. This is true for the collecting sites themselves as well as
for the different habitats or depths sampled. The occurrence and distribution of the species report-
ed do not necessarily reflect genuine patterns of distribution but reflect to a certain degree the re-
search activities carried out in the past. Furthermore, aspects concerning distribution patterns dic-
tated by ecological requirements or relative abundances of species are not considered here, simply
because the number of specimens and the associated collecting data available are not sufficient for
any conclusions on these topics to be drawn. (2) Our current knowledge of the Polynoidae in the
Arabian region is fragmentary, but so is our knowledge of the whole Indo-West Pacific. It is there-
fore extremely difficult to draw any conclusions as regards the number and presence of endemic
species. Many species recorded only from the seas surrounding the Arabian Peninsula may have
a much wider distribution and may be found in other locations with increased sampling efforts.
With this problem in mind, endemic species are mentioned only tentatively and are indicated by
quotation marks, e.g. “endemic” species.
Faunal composition
The occurrence of 23 genera and 44 species of Polynoidae is here confirmed, that is to say 14 % of
the genera and 5 % of the species worldwide. The distribution of each species is illustrated in Table
1. Diversity in terms of species richness is highest in the Red Sea (33), followed by the Arabian Sea
(20), The Gulf (11), the Suez Canal (6), the Gulf of Aden (6), and the Gulf of Oman (6) (Fig. 47).
Based on literature data, Wehe & Fiege (2002) listed 49 Polynoidae species occurring in the
seas surrounding the Arabian Peninsula, whilst registering doubts as to the taxonomic status of
nine species. After the revision of the relevant specimens given here, the occurrence of only 26 of
these species can be confirmed. These 26 species together with 14 new records and four species
new to science are described in the systematic part of this work and are listed in Table 1.
Two species are referred to other genera, i.e. Lepidasthenia vesiculosa (originally described as
Bouchiria) and Malmgreniella vesicudenta (originally described as Harmothoe), and Bouchiria is
synonymised with Lepidasthenia. Three species are considered to be junior synonyms: Iphione retic-
ulata is referred to I. ovata, and Lepidonotus impatiens meridionalis and L. australiensis are referred
to L. impatiens.
The presence of eight more species of the 49 listed in Wehe & Fiege (2002), i.e. Harmothoe
branchiata, Harmothoe imbricata, Harmothoe impar, Harmothoe spinifera, Lepidasthenia stylolepis,
Malmgreniella castanea, Polyeunoa laevis, Polynoe fumigata, remains unconfirmed, since the relevant
specimens were not available for study. However, the occurrence of most of these species in the
region appears unlikely, as explained in the “unconfirmed records’ section above. These species are
not listed in Table 1 and are not considered in the zoogeographic analysis.
Furthermore, 15 of the species reported in Wehe & Fiege (2002) do not occur in the study
area, since all of the relevant specimens examined were indeterminable or were referred to other
species (Table 2).
Comparison of the polynoid species listed by Wehe & Fiege (2002) and the species recorded
here shows that only 56 % (according to Second Kulczynski and Ochiai/Otsuka coefficients) of
172 T. Wehe
Table 1: Geographic distribution of Polynoidae species in the seas surrounding the Arabian Peninsula, the Mediterranean Sea, the
Indo-West Pacific, the East Pacific and Atlantic Ocean. According to the literature many of the species listed below appear to have a
much wider distribution (at least as regards the Indo-West Pacific) than is reflected by the references chosen here in order to confirm
their occurrence outside Arabian waters. However, the references chosen are considered to be the most reliable, since all the species
listed are well described and figured, and they adequately demonstrate the occurrence and often wide distribution of the species in
question outside the seas surrounding the Arabian Peninsula. ○: according to Wehe & Fiege (2002). ●: according to the present
work. ■: according to Barnich & Fiege (2003), Mediterranean Sea and East Atlantic. □: according to Hanley & Burke (1991a),
Chesterfield Islands and Fairway reefs, Coral Sea. ◘: according to Hanley & Burke (1990), Albany, Western Australia. ◙: according
to Hanley (1993), Rottnest Island, Western Australia. ♦: according to Barnich et al. (2004), Hainan Island, South China Sea. ◊:
according to Imajima (1997), Sagami Bay & Sea, Japan. ⌂: according to Pettibone (1969a), Philippines and South West Australia.
▲: according to Pettibone (1986), Hawaiian Islands. ▼: according to Pettibone (1975), Gulf of Mexico and Caribbean Sea. ◄:
according to Pettibone (1977), India, Indian Ocean. : according to Rullier (1972), New Caledonia. ∇: according to McIntosh
(1885), Japan. ∗: according to Ruff (1995), California and Panama. nr = new record, ns = new species, ? = tentative identification,
(E) = “endemic” in Arabian region, AO = Atlantic Ocean, AS = Arabian Sea, EM = Eastern Mediterranean, EP = East Pacific, GA =
Gulf of Aden, GO = Gulf of Oman, GU = The Gulf, IWP = Indo-West Pacific, MS = Mediterranean Sea, RS = Red Sea, SC = Suez
Canal.
SC RS GA AS GO GU MS IWP AO EP
Acholoinae
Acholoe astericola (Delle Chiaje, 1841) ○●nr ■ EM ■
Pararctonoella marginopapillata n. sp. (E) ●ns
Arctonoinae
Australaugeneria rutilans (Grube, 1878) ●nr ●nr ⌂
Gastrolepidia clavigera Schmarda, 1861 ●nr ♦
Iphioninae
Iphione henshawi Pettibone, 1986 ●nr ▲
Iphione muricata (Savigny in Lamarck,
○● ○● ○● ●nr ●nr □♦
1818)
Iphione ovata (Kinberg, 1856) ○● ●nr □♦
Lepidastheniinae
Hyperhalosydna striata (Kinberg, 1856) ●nr ○ □
Lepidasthenia nuda (Grube, 1870) (E) ○● ●nr
Lepidasthenia vesiculosa (Wesenberg-
○●
Lund, 1949) (E)
Lepidonotinae
Hermenia acantholepis (Grube, 1876) ○● □◊
Hermenia verruculosa Grube, 1856 ●nr ▼
Heteralentia ptycholepis (Grube, 1878) ●nr ○●? □♦◊
Lepidonotus carinulatus (Grube, 1869) ○● ○● ○● ○● ■ □◘♦◊
Lepidonotus clava (Montagu, 1808) ○● ○● ■ EM ■
Lepidonotus cristatus (Grube, 1876) ○● ○● ●nr ♦
Lepidonotus glaucus (Peters, 1855) ○● ○● ●nr □◘◙
Lepidonotus impatiens (Savigny in
○● ●
Lamarck, 1818)
Lepidonotus natalensis Day, 1951 ●nr ●nr ●
Lepidonotus polae n. sp. (E) ●ns
Lepidonotus purpureus Potts, 1910 ○●nr ●nr ●
Lepidonotus tenuisetosus (Gravier, 1902) ○ ●nr ○● ●nr ●nr ○● ■ EM ♦
Parahalosydnopsis arabica n. sp. (E) ●ns ●ns
Parahalosydnopsis tubicola (Day, 1973) ●nr ◄
Thormora jukesii Baird, 1865 ○● ●nr □◙♦
Thormora salvati Rullier, 1972 ●nr ●nr 
Polynoinae
Australonoe willani Hanley, 1993 ●nr ◙♦
Eunoe yedoensis (McIntosh, 1885) ●nr ∇
SC RS GA AS GO GU MS IWP AO EP
Gaudichaudius cimex (Quatrefages, 1866) ●nr ♦
Harmothoe dictyophora (Grube, 1878) ○ ○ ●nr ○● ◘◙♦
Harmothoe grisea (Ehrenberg & Grube
●nr ○● ●nr
in Grube, 1869) (E)
Harmothoe hirsuta Johnson, 1897 ●nr ●nr ∗
Harmothoe liaoi Barnich, Fiege & Sun,
●nr ●nr ●nr ●nr ♦
2004
Harmothoe marerubrum n. sp. (E) ●ns
Hermadion africanus Hartman, 1974a ●nr ○● ●
Hololepidella nigropunctata (Horst, 1915) ○● □♦
Malmgreniella murrayensis Pettibone,
○●
1993 (E)
Malmgreniella vesicudenta (Hanley &
●nr □
Burke, 1991a)
Paradyte crinoidicola (Potts, 1910) ○● ■ EM □♦
Paradyte levis (Marenzeller, 1902) ○● ○ ○● ◊
Paralepidonotus ampulliferus (Grube, 1878) ○ ●nr ○● ○● ◘♦◊
Paralepidonotus erythromaris Hanley,
○● ●nr
1991 (E)
Subadyte albanyensis Hanley & Burke,
●nr ●nr ◘◙
1990
Uncopolynoinae
Uncopolynoe corallicola Hartmann-
○●
Schröder, 1960 (E)
the species are common to both lists, i.e. the faunal composition of the Polynoidae in the seas
surrounding the Arabian Peninsula has changed for almost half of its species after revision of the
specimens.
Comparison of species richness in the seas surrounding the Arabian Peninsula with that in the
Mediterranean Sea shows a slightly higher richness in the Arabian region as a whole (44 species)
than in the Mediterranean Sea (41 species) (Barnich & Fiege 2003), although species belonging
to two additional subfamily taxa (Macellicephalinae and Polaruschakovinae) were reported in the
latter. However, looking separately at each individual sea surrounding the Arabian Peninsula, as
well as at the different regions of the Mediterranean Sea as defined by Barnich & Fiege (2003),
reveals a very heterogeneous picture with the Red Sea and the Western Mediterranean being the
regions richest in species (see Fig. 48).
No data regarding the total species richness of the Indo-West Pacific region as a whole are
available in the literature. However, perusal of only a few selected references (see below) is suffi-
cient to show that the species richness is much higher in this large area, with 85 species recorded in
just these few references (Fig. 48).
Zoogeographic affinities within the seas surrounding the Arabian Peninsula

Evaluation of the affinities of the faunas occurring in the different seas surrounding the Ara-
bian Peninsula using the Second Kulczynski and the Ochiai/Otsuka coefficients yielded similar
results (Fig. 49). The only noticeable differences between the two coefficients concerned the values
on the affinities between the Red Sea and both the Gulf of Aden and the Arabian Sea, respectively.
There is a tendency for the highest concordance to be found among adjacent seas, whereas the con-
cordance decreases with increasing geographic distance between the seas being compared. However,
174 T. Wehe
Tab. 2: Species formerly reported in the seas surrounding the Arabian Peninsula (Wehe & Fiege 2002), but not occurring in the
region based on the revision of specimens herein.
Collection numbers for

Status according to Wehe &
Original record Current status specimens not listed in
Fiege (2002)
taxonomic part
? Allmaniella sp. — Monro 1937 ? Allmaniella sp. indeterminable BMNH 1937.9.2.33-39
Nectochaeta caroli — Monro 1937 ? Drieschia pelagica juvenile form BMNH 1937.9.2.40
Michaelsen, 1892
Eunoe depressa — Rosenfeldt 1989 Eunoe depressa Moore, 1905 Eunoe yedoensis McIntosh,
1885
Eunoe pallida — Fauvel 1932 Eunoe pallida (Ehlers, 1908) indeterminable IEA N.12
Harmothoe aequiseta — Amoureux Harmothoe aequiseta (Kinberg, indeterminable IEA Y.99'
et al. 1978 1856) — nomen dubium
Harmothoe goreensis — Amoureux et Harmothoe bellani Barnich & Harmothoe grisea (Ehrenberg
al. 1978 Fiege, 2000 & Grube in Grube, 1869)
Harmothoe gilchristi — Amoureux et Harmothoe gilchristi Day, 1960 Harmothoe liaoi Barnich, Fiege
al. 1978 & Sun, 2004
Harmothoe minuta — Fauvel 1933 Harmothoe minuta (Potts, Harmothoe grisea (Ehrenberg
1910) — nomen dubium & Grube in Grube, 1869) or
Hololepidella nigropunctata
(Horst, 1915)
Iphione reticulata — Amoureux et Iphione reticulata Amoureux et Iphione ovata Kinberg, 1856
al. 1978 al., 1978
Lagisca flaccida — Fauvel 1933 Lagisca flaccida Potts, Australonoe willani Hanley,
1910 — nomen dubium 1993
Lagisca flaccida — Amoureux et al. - -
1978
Lepidasthenia elegans — Fauvel 1933 Lepidasthenia elegans (Grube, Lepidasthenia nuda (Grube,
1840) 1870)
Nectochaeta grimaldii — Monro ? Lepidasthenia grimaldii juvenile form BMNH 1937.9.2.41
1937 (Marenzeller, 1892)
Malmgreniella lunulata — Amoureux Malmgreniella lunulata (Delle indeterminable IEA Y.98
et al. 1978 Chiaje, 1830)
Harmothoe boholensis — Fauvel Paralepidonotus indicus Harmothoe grisea (Ehrenberg
1911; — Fauvel 1927; — Fauvel (Kinberg, 1856) & Grube in Grube, 1869)
1933
Harmothoe boholensis — Fauvel - Harmothoe liaoi Barnich, Fiege
1919; — Fauvel 1955; Ben-Eliahu & Sun, 2004
1972b
Scalisetosus pellucidus — Hartmann- Subadyte pellucida (Ehlers, Subadyte albanyensis Hanley &
Schröder 1960 1864) Burke, 1990
Scalisetosus fragilis — Ben-Eliahu - -
1972b
Subadyte pellucida — Amoureux 1983a - -
this pattern appears to overlap with a second pattern that results from selective sampling efforts, as
can be distinctly seen from the comparatively high concordance of all seas with the Red Sea.
Affinities of the seas surrounding the Arabian Peninsula with the Mediterranean Sea
Based on the revision of specimens as well as on literature data, Barnich & Fiege (2003) reported
four Polynoidae species common to the Mediterranean Sea and the Red Sea, i.e. Harmothoe bellani,
Fig. 47: Number and percentage (%) of Polynoidae species occurring in the seas surrounding the Arabian Peninsula based on speci-
mens revised. Total species number: 44.
Harmothoe gilchristi Day, 1960, Paradyte cf. crinoidicola, and Lepidonotus tenuisetosus. The records
of H. bellani and H. gilchristi are not confirmed here. A single slide with mounted parapodia and
elytra belonging to H. goreensis reported in Amoureux et al. (1978) and referred to H. bellani by
Barnich & Fiege (2000) is referred to Harmothoe grisea in the present paper (see Table 2). Another
slide belonging to H. gilchristi reported in Amoureux et al. (1978) and confirmed by Barnich &
Fiege (2000) is referred to Harmothoe liaoi. The occurrence of Lepidonotus tenuisetosus and Paradyte
crinoidicola in the Red Sea is confirmed.
Lepidonotus carinulatus, one of the most common species occurring in the whole of the Ara-
bian region, was reported by Baratech et al. (1986) from Alboran Island, Western Mediterranean
Sea. Barnich & Fiege (2003) could not confirm the presence of this species in the Mediterranean
Sea and suggested that L. carinulatus is a widespread species in the Indo-West Pacific region and
the Red Sea which might have been introduced into the Mediterranean Sea together with oysters.
By contrast, the occurrence of Acholoe astericola and Lepidonotus clava in the Red Sea and the
Suez Canal - two species widely distributed in the Mediterranean Sea and the North East Atlantic
- is confirmed here.
Only five species of Polynoidae occurring in the seas surrounding the Arabian Peninsula are con-
sidered to be in common with the Mediterranean Sea (see Table 1). Consequently, faunal affinities
between the whole Arabian region and the Mediterranean Sea are low, as can be seen in Fig. 50.
176 T. Wehe
Fig. 48: Comparison of the species richness in the seas surrounding the Arabian Peninsula with parts of the Mediterranean Sea
and the Indo-Westpacific. Only records confirmed by Barnich & Fiege (2003) were considered for the Mediterranean Sea. Indo-
West Pacific species were compiled according to references cited in the text [AD = Adriatic Sea, AE = Aegean Sea, AS = Arabian
Sea, CM = Central Mediterranean, EM = Eastern Mediterranean, GA = Gulf of Aden, GO = Gulf of Oman, GU = The Gulf,
IWP = Indo-West Pacific, RS = Red Sea, SC = Suez Canal, WM = Western Mediterranean].
Affinities of the seas surrounding the Arabian Peninsula with the Indo-West Pacific
Evaluating the faunal affinities of the Arabian region with the Indo-West Pacific region is difficult,
since literature on the Polynoidae from the latter region is very scarce. Only a few reliable refer-
ences are available and these have been used for this comparison. Consequently, the 85 species
reported in these references originate from only a very few locations, mainly the Island of Hainan,
South Chinese Sea (Barnich et al. 2004), Sagami Bay, Japan (Imajima 1997), Albany and Rottnest
Island, Western Australia (Hanley 1993, Hanley & Burke 1990), the Chesterfield Islands and
Fairway Reefs, Coral Sea (Hanley & Burke, 1991 a). The records listed in these publications were
supplemented by material studied from the Indo-West Pacific and according to Hanley (1992),
McIntosh (1885), Pettibone (1969 a, 1977, 1986 a), and Rullier (1972). This selection of refer-
ences cannot be regarded as representative for the whole of the Indo-West Pacific and therefore the
following conclusions have to be considered as preliminary. Nevertheless, comparison of the data
compiled here with that provided in the references mentioned above enables a first approximation
of the species affinities between these regions to be made. Accordingly, 30 of the 44 species re-
corded here also occur in the Indo-West Pacific, some of them apparently being widely distributed
as can be seen from Table 1. The high affinity with the Indo-West Pacific is illustrated in Fig. 51.
Fig. 49: Affinities of the Polynoidae faunas in the seas surrounding the Arabian Peninsula based on different indices. Rates of species
concordance are expressed in %.
178 T. Wehe
Fig. 50: Affinities of the Polynoidae in the seas surrounding the Arabian Peninsula with the Mediterranean Sea based on specimens
revised herein and Barnich & Fiege (2003). Rates of species concordance are expressed in % [AS = Arabian Sea, GA = Gulf of
Aden, GO = Gulf of Oman, GU = The Gulf, MS = Mediterranean Sea, RS = Red Sea, SC = Suez Canal].
Fig. 51: Affinities of the Polynoidae in the seas surrounding the Arabian Peninsula with the Indo-West Pacific based on specimens
revised herein and references cited in the text. Rates of species concordance are expressed in % [AS = Arabian Sea, GA = Gulf of
Aden, GO = Gulf of Oman, GU = The Gulf, IWP = Indo-West Pacific, RS = Red Sea, SC = Suez Canal].
Table 3: Number and percentage (%) of “endemic” Polynoidae species occurring in the different seas surrounding the Arabian Pe-
ninsula based on the present revision of specimens.
Endemics sensu stricto Endemics sensu lato

number % number %
Suez Canal - - 1 17
Red Sea 3 9 4 12
Gulf of Aden - - 1 17
Arabian Sea 1 5 1 5
Gulf of Oman 1 17 - -
The Gulf 1 9 2 18
Endemism
Conclusions on the degree of endemism among the Polynoidae in the Arabian region are difficult,
for the reasons explained above. For the time being, the following 10 species, i.e. 23 % of the total,
are considered to be “endemics”, six (14 %) of which are “endemics” sensu stricto (present in only
one of the seas in question, e.g. only in the Red Sea), whereas four (9 %) represent “endemics” sen-
su lato (present in more than one of the seas in question, e.g. the Red Sea and The Gulf, but not
outside the Arabian region) (Table 1): Pararctonoella marginopapillata n. sp., Lepidasthenia nuda,
Lepidasthenia vesiculosa, Lepidonotus polae n. sp., Parahalosydnopsis arabica n. sp., Harmothoe grisea,
Harmothoe marerubrum n. sp., Malmgreniella murrayensis, Paralepidonotus erythromaris, Uncopoly-
noe corallicola. Looking at the various seas, the distribution and percentage of these “endemics” is
shown in detail in Table 3.
Data for a comparison of the endemism in other polychaete families within the Arabian region
are not available. Six endemic species were reported for the Mediterranean Sea, constituting 9 % of
the fauna in question (Barnich & Fiege 2003).
It is assumed that the rate of “endemics” will decrease with further sampling activities around
the Indo-West Pacific. In general, a decrease in the rate of endemism with increased studies appears
naturally and has been reported elsewhere, e.g. Head (1987) and Quignard & Tomasini (2000).
It therefore appears that Fauvel (1932: 6) was most probably right when he stated: “The fauna of
India does not materially differ from that of the Red Sea, the Persian Gulf, the Philippine Islands
and the Malay Archipelago, while many species from the Pacific Ocean, New Zealand, New Cal-
edonia and Australia are also found in this area”.
Lessepsian Migration
The phenomenon of migration of Red Sea biota through the Suez Canal into the Mediterranean
Sea is known as Lessepsian migration and was extensively studied by Por (1978). So far as polycha-
etes are concerned, nine so-called “high probability Lessepsian migrants” and 14 “low probability
Lessepsian migrants” were reported by this author. However, no Polynoidae were among the spe-
cies listed. Since then only the Nereididae and Serpulidae have been the subject of investigations
(e.g. Ben-Eliahu 1989, 1991 a, 1991 b, Ben-Eliahu & Hove 1992, Zibrowius & Bitar 2003).
Of the five Polynoidae species considered here to be present in the Red Sea as well as in the Medi-
terranean Sea (see above), Lepidonotus tenuisetosus and Paradyte crinoidicola were already indicated
as possible Lessepsian migrants by Barnich & Fiege (2003), since they were only recorded from
180 T. Wehe
Fig. 52: Present known depth range of Polynoidae in the seas surrounding the Arabian Peninsula.
the Eastern Mediterranean. Lepidonotus carinulatus, as mentioned above, is only reported from
the Western Mediterranean Sea and furthermore may represent an invasive organism introduced
with oysters. By contrast, Acholoe astericola and Lepidonotus clava are widely distributed in the
Mediterranean Sea and in the North East Atlantic (Seidler 1923, Pettibone 1996 b, Barnich &
Fiege 2003). Tebble & Chambers (1982) and Hartmann-Schröder (1996), in their comments
on distribution, also mentioned that L. clava occurs in the Indian Ocean. However, no records of
this species could be found in any of the major works on the fauna of the Indian Ocean (e.g. Potts
1910, Horst 1917, Fauvel 1932, 1953, Day 1967, Hartman 1974 a, 1974 b) and its occurrence
therefore appears questionable. Since both species are recorded only from the Suez Canal and the
Red Sea, they may represent Anti-Lessepsian migrants, a rather uncommon direction of migration
through the Suez Canal. Por (1978) listed Scalisetosus fragilis Claparède, 1868 as a possible Anti-
Lessepsian migrant. Unfortunately, the occurrence of this species in the Mediterranean Sea was not
confirmed by Barnich & Fiege (2003).
Depth distribution
In many cases, unfortunately, the habitat data relating to specimens examined during this study are
incompletely documented, especially with regard to depth. The depth distribution of only 29 out
of 44 Polynoidae species is therefore given in Fig. 52. However, even this information is limited,
since it is often based on information for only some of the specimens belonging to the species in
question. The depth range reported for the same species from outside the Arabian area may be
considerably different, as can be seen from the additional depth distribution data given for each
species in the systematic account.
In textbooks on marine biology, the benthic region down to the edge of the continental shelf
is usually referred to as the littoral zone, extending to about 200 m depth (e.g. Gage & Tyler
1991, Tardent 1993). All the Polynoidae species listed in Fig. 52 are reported from this zone
and not deeper than 130 m, with the exception of only two species recorded from bathyal depths
(200‑4000 m, according to Tardent 1993), i.e. Eunoe yedoensis (490‑757 m) and Hermadion afri-
canus (121‑1116 m).
For 13 species not listed in Fig. 52, detailed collecting depths are not available. Nevertheless,
by inferring from data for the rest of the sample, e.g. the collector, or from approximate geographi-
cal information, most of the specimens in question were probably collected in shallow water, or at
least in the littoral zone.
Acknowledgements
For the loan of specimens and providing information I would like to thank the following persons
and institutions: Dr. Y. Benayahu and A. Shlagman (Tel Aviv University, Department of Zoology,
Israel), Dr. M.N. Ben-Eliahu (The Hebrew University, Zoological Collections, Jerusalem, Israel),
Dr. A. Brandt and G. Wegener (Zoologisches Institut und Museum der Universität Hamburg,
Germany), A. Cabrinovich, M. Lowe and A. Muir (The Natural History Museum, London,
United Kingdom), Dr. D. Eibye-Jacobsen (Zoologisk Museum, Copenhagen, Denmark), Dr.
Y.H. Fadlallah (Center for Environment and Water Research Institute, King Fahd University of
Petroleum & Minerals, Dhahran, Kingdom of Saudi Arabia), Dr. K. Fauchald and W.G. Keel
(United States National Museum of Natural History, Smithsonian Institution, Washington D.C.,
U.S.A.), Dr. J.D. George (The Natural History Museum, London, United Kingdom), Dr. P. Gilet
(Institut d’Écologie Appliquée, Angers, France), L.H. Harris (Allan Hancock Foundation, Los An-
geles County Museum, Polychaete Collection, Los Angeles, U.S.A.), Dr. L. Levin and L.L. Lovell
(Scripps Institution of Oceanography, La Jolla, California, U.S.A), S. Lundberg (Naturhistoriska
Riksmuseet, Stockholm, Sweden), Dr. B. Neuhaus (Naturhistorisches Forschungsinstitut, Zentra-
linstitut der Humboldt-Universität zu Berlin, Museum für Naturkunde, Germany), Dr. F. Pleijel
and M.J. d’Hondt (Muséum National d’Histoire Naturelle, Paris, France), Dr. H. Sattmann (3.
Zoologische Abteilung, Naturhistorisches Museum, Vienna, Austria), Dr. A. Tzetlin (Department
of Invertebrate Zoology, Biological Faculty, State University of Moscow, Russia), C. Vollelv (Zo-
ologisk Museum, Universitetet i Oslo, Norway) and Dr. A. Wiktor (Museum of Natural History,
Wrocław, Poland).
For providing working facilities during research visits I would like to express my sincere thanks
to the following persons: Dr. Danny Eibye-Jacobsen (Zoologisk Museum, Copenhagen, Den-
mark), Dr. Birger Neuhaus (Naturhistorisches Forschungsinstitut, Zentralinstitut der Humboldt-
Universität zu Berlin, Museum für Naturkunde, Germany), Dr. Gordon Patterson (The Natural
History Museum, London, United Kingdom), Dr. Fredrik Pleijel (Muséum National d’Histoire
Naturelle, Paris, France) and Dr. Helmut Sattmann (3. Zoologische Abteilung, Naturhistorisches
Museum, Vienna, Austria).
I want to thank Dr. Friedhelm Krupp, Forschungsinsitut & Naturmuseum Senckenberg,
Frankfurt a.M., and Dr. Michael Apel, Museum Wiesbaden (formerly at Forschungsinsitut &
Naturmuseum Senckenberg) for the unique opportunity of joining the marine expedition team in
182 T. Wehe
spring 2000 to the Socotra Archipelago, PR Yemen, within the framework of the United Nations
Development Programme (UNDP)/Global Environment Facility (GEF) project “Conservation
and Sustainable Use of Biodiversity of Socotra Archipelago (UNOPS YEM/96/G32)”. Thanks are
also due to the staff at the “Project Implementation Unit (PIU)” on Socotra Island, especially Dr.
Edoardo Zandri, as well as the representatives of the “Environmental Protection Council (EPC)”
of the PR Yemen, and the other members of the marine biological team, but especially to Dr. M.
Apel for his support during the fieldwork.
For the generous donation of specimens to the Forschungsinsitut & Naturmuseum Senck-
enberg, special thanks are due to Dr. Javed Mustaquim (Centre of Excellence in Marine Science,
University of Karachi, Pakistan), Dr. Peter Garwood (Identichaet, Heaton, Newcastle upon
Tyne, United Kingdom) and Dr. Michael Stachowitsch (Institut für Ökologie und Naturschutz,
Abteilung Meeresbiologie, Universität Wien, Austria).
Special thanks are due to Dr. Ruth Barnich, Sektion für Marine Evertebraten II, Forschungsin-
stitut & Naturmuseum Senckenberg, for valuable discussions and critical comments on this work.
Gisela Adam, Institut für Zoologie, Universität Heidelberg, was an invaluable help in prepar-
ing the plates for this work.
This work was in part financed by the following grants: “Landesgraduiertenförderung” schol-
arship of the Ruprecht-Karls-Universität, Heidelberg, and a scholarship from the Forschungsinsitut
& Naturmuseum Senckenberg, Frankfurt a.M. Research stays at the Natural History Museums in
Paris and Vienna were financed through travel grants from the “Landesgraduiertenförderung”. Re-
search visits to The Natural History Museum, London, and the Zoologisk Museum, Copenhagen,
were generously financed by two grants from these two institutions: SYS-RESOURCE (London)
and COBICE (Copenhagen), both within the frame of the IHP (Improving the Human Potential)
program of the EU. Travel costs to the Socotra Archipelago were refunded by the Ungerer Founda-
tion, Frankfurt a.M.
Finally, I would like to thank Prof. Dr. Dr. h.c. Volker Storch, Institut für Zoologie, Univer-
sität Heidelberg, and especially Dr. Dieter Fiege, Forschungsinsitut & Naturmuseum Senckenberg,
Frankfurt a.M, for their constant support during the preparation of my doctoral thesis, of which
this work forms a part.
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Manuscript submitted: 27 October 2004

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Revision of Scale Worms in Arabian Seas

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FAUNA OF ARABIA 22: 23–197

Date of publication: 18.12.2006

Revision of the scale worms (Polychaeta:

History of polychaete research in the Arabian region

Anatomy and physiology

Biology and ecology

Eidonomy and critical discussion of characters

Materials and methods

HUJ The Hebrew University, Zoological Collections, Jerusalem, Israel

Outline of species description

Suez Canal ↔ Red Sea: 5

Computation of faunal affinity values

Light microscopy, preparation of drawings and photography

Polynoidae Kinberg, 1856

1 Prostomium without antennae................................................................................... 2

20 18 segments, 9 pairs of elytra. Palps conical, short, not visible dorsally

Acholoinae Pettibone, 1996

Key to the genera and species of Acholoinae

Acholoe Claparède, 1870

Acholoe astericola (Delle Chiaje, 1841) Fig. 3 a‑g

Pararctonoella Pettibone, 1996

Pararctonoella marginopapillata n. sp. Fig. 4 a‑j

Arctonoinae Hanley, 1989

Key to the genera and species of Arctonoinae

Australaugeneria Pettibone, 1969

Australaugeneria rutilans (Grube, 1878) Fig. 5 a‑i

Gastrolepidia Schmarda, 1861

R e m a r k s : Gastrolepidia is monospecific and occurs as a commensal on holothurians

Gastrolepidia clavigera Schmarda, 1861 Fig. 6 a‑h

Iphioninae Kinberg, 1856

Iphione Kinberg, 1856

Key to the species of Iphione

1 Elytra with smooth margins I. ovata

Iphione henshawi Pettibone, 1986 a Fig. 7 a‑h

Iphione muricata (Savigny in Lamarck, 1818) Fig. 8 a‑h

Iphione ovata Kinberg, 1856 Fig. 9 a‑i, Pl. 1 a

Lepidastheniinae Pettibone, 1989

Key to the genera and species of Lepidastheniinae

1 Segments numerous, up to 100 or more. Elytra numerous, on segments 2, 4, 5, 7,

Hyperhalosydna Augener, 1922

Hyperhalosydna striata (Kinberg, 1856) Fig. 10 a‑g, Pl. 1 b, c

Lepidasthenia Malmgren, 1867

Lepidasthenia nuda (Grube, 1870) Fig. 11 a‑j

R e d e s c r i p t i o n based on lectotype designated here: Prostomium (Fig. 11 a) about as wide

D e p t h r a n g e : So far recorded between 25 and 30 m; unknown elsewhere.

Lepidasthenia vesiculosa (Wesenberg-Lund, 1949) n. comb. Fig. 12 a‑e

Lepidonotinae Willey, 1902

Key to the genera of Lepidonotinae

1 Body always with 26 segments................................................................................... 2

Hermenia Grube, 1856

Key to the species of Hermenia

Hermenia acantholepis (Grube, 1876) Fig. 13 a‑c

Hermenia verruculosa Grube, 1856 Fig. 14 a, b; Pl. 1 d

Fig. 14: Hermenia

Heteralentia Hanley & Burke, 1991

Heteralentia ptycholepis (Grube, 1878) Fig. 15 a‑h

Lepidonotus Leach, 1816

Ty p e s p e c i e s : Aphrodita clava Montagu, 1808.

Key to the species of Lepidonotus

1 Elytra with fringing papillae....................................................................................... 2

Lepidonotus carinulatus Grube, 1869 Fig. 16 a‑o

Lepidonotus clava (Montagu, 1808) Fig. 17 a‑j