Eusyllinae (Polychaeta: Syllidae) from Australia with the description of a new genus and fifteen new species

Abstract

Large collections of Syllidae (Polychaeta) from around Australia, housed at the Australian Museum (Sydney), have been examined and identified. Australian material from the Hamburgische Zoologische Museum der Universität, Hamburg, Germany was also examined. All known Australian species of the subfamily Eusyllinae (Syllidae) are described and figured. Some were examined using the Scanning Electron Microscope to illustrate characters and methods of reproduction in this subfamily. Keys to genera and species are given. A total of 53 species are reported from Australia belonging to 14 genera: Amblyosyllis Grube, 1857 (3 species); Anoplosyllis Claparède, 1868 (1 species); Astreptosyllis Kudenov & Dorsey, 1982 (2 species); Eusyllis Malmgren, 1867 (3 species); Nudisyllis Knox & Cameron, 1970 (1 species); Odontosyllis Claparède, 1863 (9 species); Opisthodonta Langerhans, 1879 (3 species); Paraehlersia San Martín, 2003 (2 species); Paraopisthosyllis Hartmann-Schröder, 1991 (4 species); Pionosyllis Malmgren, 1867 (15 species); Psammosyllis Westheide, 1990 (1 species); Streptodonta n.gen. (1 species); Streptosyllis Webster & Benedict, 1884 (3 species); and Syllides Örsted, 1845 (5 species). A total of 15 new species are described: Amblyosyllis enigmatica, A. multidenticulata, Odontosyllis marombibooral; Opisthodonta hanneloreae, Paraopisthosyllis alternocirra, P. ornaticirra, Syllides tam, Pionosyllis rousei, P. kalimna, P yolandae, P. mariae, P. mayteae, P. ancori, P. koolalya, and P. heterochaetosa. A new name, Pionosyllis hartmannschroederae, is proposed for Typosyllis (Langerhansia) longisetosa Hartmann-Schröder, 1990 as this is junior homonym of Pionosyllis longisetosa (Hartmann-Schröder, 1965). Additionally, five species are new records for Australia: Odontosyllis gravelyi Fauvel, 1930; Opisthodonta morena Langerhans, 1879; Pionosyllis corallicola Ding & Westheide, 1997; Streptodonta pterochaeta (Southern, 1914) and Syllides japonicus Imajima, 1966. A discussion of the reproduction and systematics of the subfamily is given.

Full text

© Copyright Australian Museum, 2006
* author for correspondence
Eusyllinae (Polychaeta: Syllidae) from Australia with the
Description of a New Genus and Fifteen New Species
GUILLERMO SAN MARTÍN
1
* AND PAT HUTCHINGS
2
1
Departamento de Biología (Zoología), Laboratorio de Biología Marina e Invertebrados,
Facultad de Ciencias, Universidad Autónoma de Madrid, Canto Blanco, 28049 Madrid, Spain
guillermo.sanmartin@uam.es
2
Aquatic Zoology, Australian Museum, 6 College Street, Sydney NSW 2010, Australia
path@austmus.gov.au
ABSTRACT. Large collections of Syllidae (Polychaeta) from around Australia, housed at the Australian
Museum (Sydney), have been examined and identified. Australian material from the Hamburgische
Zoologische Museum der Universität, Hamburg, Germany was also examined. All known Australian
species of the subfamily Eusyllinae (Syllidae) are described and figured. Some were examined using
the Scanning Electron Microscope to illustrate characters and methods of reproduction in this subfamily.
Keys to genera and species are given. A total of 53 species are reported from Australia belonging to 14
genera: Amblyosyllis Grube, 1857 (3 species); Anoplosyllis Claparède, 1868 (1 species); Astreptosyllis
Kudenov & Dorsey, 1982 (2 species); Eusyllis Malmgren, 1867 (3 species); Nudisyllis Knox & Cameron,
1970 (1 species); Odontosyllis Claparède, 1863 (9 species); Opisthodonta Langerhans, 1879 (3 species);
Paraehlersia San Martín, 2003 (2 species); Paraopisthosyllis Hartmann-Schröder, 1991 (4 species); Pionosyllis
Malmgren, 1867 (15 species); Psammosyllis Westheide, 1990 (1 species); Streptodonta n.gen. (1 species);
Streptosyllis Webster & Benedict, 1884 (3 species); and Syllides Örsted, 1845 (5 species). A total of 15
new species are described: Amblyosyllis enigmatica, A. multidenticulata, Odontosyllis marombibooral;
Opisthodonta hanneloreae, Paraopisthosyllis alternocirra, P. ornaticirra, Syllides tam, Pionosyllis rousei,
P. kalimna, P. yolandae, P. mariae, P. mayteae, P. ancori, P. koolalya, and P. heterochaetosa. A new
name, Pionosyllis hartmannschroederae, is proposed for Typosyllis (Langerhansia) longisetosa Hartmann-
Schröder, 1990 as this is junior homonym of Pionosyllis longisetosa (Hartmann-Schröder, 1965).
Additionally, five species are new records for Australia: Odontosyllis gravelyi Fauvel, 1930;
Opisthodonta morena Langerhans, 1879; Pionosyllis corallicola Ding & Westheide, 1997; Streptodonta
pterochaeta (Southern, 1914) and Syllides japonicus Imajima, 1966. A discussion of the reproduction
and systematics of the subfamily is given.
SAN MARTÍN, GUILLERMO, & PAT HUTCHINGS, 2006. Eusyllinae (Polychaeta: Syllidae) from Australia with the
description of a new genus and fifteen new species. Records of the Australian Museum 58(3): 257–370.
Records of the Australian Museum (2006) Vol. 58: 257–370. ISSN 0067-1975
www.amonline.net.au/pdf/publications/1466_complete.pdf
This is the second contribution to the Australian Syllidae,
based on the large collections housed in the Australian
Museum from all around Australia, and revision of material
collected and described by Hartmann-Schröder (1979–
1991). The material examined was collected predominantly
from southern Australia, and no material was examined from
north of 14°S on either the west or east coast of Australia.
This paper also summarises material already published by
San Martín (2002, 2005) and San Martín & López (2003).
A general introduction to the family Syllidae is given by

258 Records of the Australian Museum (2006) Vol. 58
San Martín (2005) in his revision of the subfamily
Exogoninae in Australian waters. In this paper, all species
belonging to the subfamily Eusyllinae are described and
figured, and keys for identification provided. Comments
are given on the other genera of Eusyllinae not currently
known from Australia. Subsequent papers will deal with
the subfamilies Syllinae and Autolytinae.
The subfamily Eusyllinae was erected by Malaquin
(1893), for the genera Syllides Örsted, 1845; Opisthodonta
Langerhans, 1879; Pionosyllis Malmgren, 1867; Eusyllis
Malmgren, 1867; Odontosyllis Claparède, 1863; and
Amblyosyllis Grube, 1857. Subsequently, Fauvel (1923)
included the genera Fauvelia Gravier, 1900 (now considered
a doubtful genus), Streptosyllis Webster & Benedict, 1884;
and Parapionosyllis Fauvel, 1923 (now considered as
belonging to the Exogoninae). Later, San Martín (2003)
re-erected Anoplosyllis Claparède, 1868, and erected
Neopetitia and Paraehlersia and placed them in the
subfamily. Other genera considered to belong to the
subfamily are Palposyllis Hartmann-Schröder, 1977;
Miscellania Martín, Alós & Sardá, 1990; Dioplosyllis
Gidholm, 1962; Paraopisthosyllis Hartmann-Schröder, 1991
(although with some doubts, see below); Astreptosyllis
Kudenov & Dorsey, 1982; and Streptospinigera Kudenov,
1983. The genus Psammosyllis Westheide, 1990, which was
not assigned to a subfamily when erected, is herein
considered to belong to the Eusyllinae. In this paper, we
erect a new genus, Streptodonta. Other genera previously
assigned to this subfamily include: Irmula Ehlers, 1913 (now
transferred to Hesionidae); Clavisyllis Knox, 1957;
Lamellisyllis Day, 1960 (both questionable members of this
subfamily); and Rhopalosyllis Augener, 1913 (now assigned
to the Syllinae). Nudisyllis Knox & Cameron, 1970 is redefined.
Currently 14 of these genera are known to occur in Australia.
This subfamily is heterogeneous and probably should
be divided into several monophyletic groups (Glasby, 1994;
San Martín, 2003). Until a detailed analysis (currently in
preparation by senior author) is completed, however, we
prefer to use the traditional classification of Eusyllinae.
A consistent diagnosis of this subfamily is difficult to
provide because of the variability of many of the characters.
Like most syllids, they are dorsally arched, convex, ventrally
flat or concave. Size range of the subfamily Eusyllinae varies
from minute to more than 30 mm long, occupying a similar
range of habitats to members of Exogoninae (see San
Martín, 2005). They often are fragile and Museum
collections rarely contain complete, well-preserved
specimens necessary for detailed studies. Eusyllinae
typically lack colour markings. The palps are fused, partially
fused or free from each other, depending upon the genera.
Even within a genus, the degree of fusion of palps varies.
Nudisyllis was originally described as lacking antennae and
dorsal cirri, but this was based on data obtained from
examination of damaged specimens, intact specimens have
3 antennae and dorsal cirri, so all Eusyllinae have three
antennae that are typically long, extending beyond palps.
Four lensed eyes and sometimes two additional eyespots
are present, although these may fade with time on preserved
material. The peristomium has two pairs of tentacular cirri,
usually long and slender, and may be modified dorsally to
form an occipital flap. Nuchal organs consist of two
dorsolateral, densely ciliated grooves between the
prostomium and peristomium. The genus Amblyosyllis (as
well as Clavisyllis and Lamellisyllis) is characterized by
having a nuchal organ consisting of two nuchal lappets,
which strongly resemble those found in the subfamily
Autolytinae. The pharynx is straight, except in Amblyosyllis,
typically with a conical tooth that is located on the anterior
margin opening or behind the anterior rim, or in the middle
or posterior part of the pharynx. Other genera (Syllides,
Anoplosyllis, Streptosyllis, Astreptosyllis, Streptospinigera)
lack a pharyngeal tooth. These latter genera resemble each
other and perhaps should be removed from the subfamily.
Amblyosyllis has a long, slender, coiled pharynx, with a
trepan, lacks a tooth, and margins of the pharynx are
ornamented with soft papillae. This genus together with
Clavisyllis and Lamellisyllis may also need to be removed
from the Eusyllinae, since they have nuchal lappets instead
of ciliated grooves as nuchal organs.
Parapodia are uniramous, with dorsal and ventral cirri,
although dorsal cirri may be absent on chaetiger 2. Dorsal
cirri typically are long, filiform, smooth, not articulated,
although in Paraopisthosyllis and some species of
Pionosyllis all or some are club-shaped and other species
of Pionosyllis has short, exogonid-like dorsal cirri, although
anteriorly they are long and smooth. Members of the
Eusyllinae reproduce by epigamy (Garwood, 1991; Franke,
1999; San Martín, 2003). The eyes enlarge (see Figs 38A,
22A), colour patterns if present become far more marked
and long, capillary notochaetae develop on middle and
posterior segments (Figs 36D, 45E, 71B), which presumably
are used in swimming. Members of the subfamily
Exogoninae also reproduce by epigamy, but they attach their
eggs either dorsally by means of capillary notochaetae of
females, or ventrally by attaching them to the nephridial
pore (San Martín, 2005). Some members of the genus
Nudisyllis have been shown to brood dorsally, and some
members of Syllides and Pionosyllis breed ventrally, but
other information is lacking. San Martín (2005) has shown
the usefulness of reproductive characters in the Exogoninae
that may also prove useful in the Eusyllinae when more
genera have been examined for these characters.
Some species of syllids are restricted to small geographical
areas, and many examples can be found within the Australian
fauna, but others have apparently wide distributions, and are
regarded as being “cosmopolitan” species. Although
increasingly such so-called “cosmopolitan” species are being
found to represent suites of sibling species (see Westheide,
1971; Westheide & Hass-Cordes, 2001). In some of these
examples, species exhibit no morphological differences and
molecular techniques must be used to separate the species.
Westheide & Hass-Cordes (2001) have used such techniques
on syllids and found it useful. In some cases upon careful
examination of these so-called “cosmopolitan” species has
revealed small but consistent morphological differences and
we have therefore described them as separate species. In other
cases, we have been unable to find any morphological
differences and so tentatively accept that the Australian
Eusyllinae fauna includes these widely distributed species.
Material and methods
The material examined was mainly from the collections in
The Australian Museum (AM), and was collected by many
people including: N. Coleman, G. Wilson, J.K. Lowry, R.T.
Springthorpe, H.E. Stoddart, P.A. Hutchings, A. Murray,
T.J. Ward, P.C. Young, A. Jones, and others. Australian

San Martín & Hutchings: Eusyllinae of Australia 259
material from the Zoologisches Museum of Hamburg
(HZM), collected and identified by Hartmann-Schröder, has
been re-examined. The specimens are preserved in 70%
ethanol after having been fixed in formalin. Examinations
were made using a compound microscope with interference
contrast optics (Nomarsky). Drawings were made using a
camera lucida drawing tube. Scanning Electron Microscope
observations and photographs were made in the SIDI
(Servicio Interdepartamental de Investigación) of the
Universidad Autónoma de Madrid, Spain and at the SEM
unit at the Australian Museum.
Information about aboriginal words for the names of
several new taxa was obtained from Endacott (1973).
Some structures, e.g., nuchal organs or eyespots, are
difficult to see. They are described only when they were
visible on the specimens, although their presence cannot
be excluded and certainly some species lack eyespots.
Categories of sizes of specimens given in the text are:
small (<5 mm in length), medium (5–10 mm in length) and
large (>10 mm in length). In syllids, the length of chaetal
blades within fascicle typically decreases from dorsal to
ventral (dorsoventral gradation); and also the shape and
length of the chaetae may vary along the body, and therefore
all descriptions include this information.
A useful compilation of polychaete terms are provided
by Glasby et al. (2000) and by San Martín (2005).
The Material examined section lists material in a
clockwise direction around Australia.
The following abbreviations are used in the Material examined:
AM The Australian Museum, Sydney, NSW,
Australia
BMNH The Natural History Museum, London, UK
HMZ Zoologisches Museum of Hamburg, Germany
LACM Los Angeles County Museum, Los Angeles, USA
MNHN Museum national d’Histoire naturelle, Paris, France
MNCNM Museo Nacional de Ciencias Naturales de
Madrid, Spain
NFMN Naturhistorisches Forschungsinstitut Museum
für Naturkunde, Berlin, Germany
SMNH Naturhistoriska Riksmuseet, Stockholm, Sweden
SMF Senckenberg Museum, Frankfurt, Germany
USNM National Museum of Natural History, Washing-
ton, D.C, USA
MPUW Museum Przydnicze Uniwersytetu Wroclaws-
kiego, Wroclaw, Poland
Key to genera of Eusyllinae recorded from Australia
1 Body, rough, densely papillated; with short, inflated antennae and
dorsal cirri (Fig. 50A) .......................................................................................... Paraopisthosyllis
—— Body elongated, smooth, without epidermal papillae; antennae
and cirri not inflated (except in some species of Pionosyllis, with
some cirri inflated) ......................................................................................................................... 2
2 Nuchal organs as lappets (Fig. 2C). Pharynx long, highly
convoluted with trepan, without mid-dorsal pharyngeal tooth (Fig.
3A) ............................................................................................................................... Amblyosyllis
—— Nuchal organs as two ciliated grooves between prostomium and
peristomium. Pharynx straight, not convoluted............................................................................. 3
3 Pharynx unarmed ........................................................................................................................... 4
—— Pharynx armed with single tooth, and or pharyngeal denticles .................................................... 7
4 Aciculae of some anterior parapodia with inflated tips (Fig. 82I) ............................... Streptosyllis
—— Aciculae without inflated tips (Fig. 6E) ........................................................................................ 5
5 Dorsal cirri all smooth, more or less club-shaped (Fig. 6A) ....................................... Anoplosyllis
—— Dorsal cirri from chaetiger 3 distinctly articulated (Fig. 90A) ..................................................... 6
6 Dorsal simple chaetae with distinct, longitudinally striated, distal
hood (Fig. 9C). Compound chaetae unidentate. Ventral cirri of
posterior parapodia distinctly elongated.................................................................... Astreptosyllis
—— Dorsal simple chaetae without hood, or if present, small and not
striated. Compound chaetae bidentate. Ventral cirri not elongated ..................................... Syllides
7 Pharynx with incomplete trepan formed by few teeth, backwardly
directed, without mid-dorsal tooth (Fig. 20A) ........................................................... Odontosyllis
—— Pharynx with mid-dorsal tooth ...................................................................................................... 8
8 Pharyngeal tooth surrounded by incomplete circle of small
denticles, forwardly directed, forming incomplete trepan (Fig. 10C)..................................... Eusyllis
—— Single pharyngeal tooth present, trepan absent ............................................................................. 9
9 Blades of compound chaetae with tendon connecting proximal
tooth with margin (Fig. 74D). Pharyngeal tooth located near
anterior rim, on middle of pharynx or posteriorly in mid line (Fig.
74A) ................................................................................................................. Pionosyllis (in part)
—— Blades of compound chaetae lacking such tendon ...................................................................... 10

260 Records of the Australian Museum (2006) Vol. 58
10 Pharyngeal tooth located far from anterior rim ........................................................................... 11
—— Pharyngeal tooth located on, or close to anterior rim ................................................................. 14
11 Tentacular and some dorsal cirri inflated, club-shaped (Fig. 60A) ................... Pionosyllis (in part)
—— Without inflated dorsal cirri ......................................................................................................... 12
12 Several anterior parapodia with distally inflated aciculae (Fig.
81A,C) .............................................................................................................. Streptodonta n.gen.
—— Without distally inflated aciculae ................................................................................................ 13
13 Palps free, not fused at base. Dorsal cirri long, slender, filiform,
twice width of body. Ventral cirri of most anterior parapodia
partially fused to parapodial lobes (Fig. 38E). Pharyngeal tooth
located medially ......................................................................................................... Opisthodonta
—— Palps fused at bases and for most of their length. Dorsal cirri
relatively short, about half width of body, ventral cirri not fused
to parapodial lobes. Pharyngeal tooth located laterally (Fig. 80A) ............................ Psammosyllis
14 Ventral cirri inserted medially or distally on parapodial lobes (Fig.
59B). Dorsal cirri of two lengths, long filiform and extremely
short exogonid-like dorsal cirri that alternate along body (Fig.
59A) ................................................................................................................. Pionosyllis (in part)
—— Ventral cirri inserted at bases of parapodia. Dorsal cirri similar
throughout or else differences between short and long cirri not as
pronounced ................................................................................................................................... 15
15 Antennae, tentacular and anterior dorsal cirri articulated (adults).............................................. 16
—— Antennae and cirri smooth, sometimes rugose, pseudoarticulated ............................................... 17
16 Segments posterior to proventricle fused in units of 2–3 segments
(Fig. 78A). Subcirral papilla absent. Palps completely free .......................... Pionosyllis (in part)
—— Segments not fused. Small subcirral papilla present on parapodia,
shorter than width of dorsal cirrus (Fig. 44B). Palps fused at base ............................... Paraehlersia
17 Dorsal cirri (except some anteriormost) exogonid-like, short,
slightly longer than parapodial lobes (Fig. 56A)............................................ Pionosyllis (in part)
—— Dorsal cirri long ........................................................................................................................... 18
18 Small to minute size (<5 mm in length). Pharynx short, shorter
than proventricle, with a long tooth (Fig. 17A). Compound chaetae
unidentate or provided with small, spine-like proximal tooth ........................................ Nudisyllis
—— Medium to large size (>5 mm in length). Pharynx long. Compound
chaetae bidentate .......................................................................................................................... 19
19 Distinct prechaetal lobe present (Fig. 66B). Acicula straight,
extending beyond parapodial lobes. Blades of compound chaetae
without long, fine spines; without spiniger-like chaetae. Large size,
distinctly macrofaunal (>10 mm in length) .................................................... Pionosyllis (in part)
—— Prechaetal lobe absent. Acicula distally knobbed, with 2 unequal
lobes (Fig. 68J). Blades of compound chaetae with long, fine,
distally ornamented spines (Fig. 63G); sometimes with spiniger-
like chaetae (Fig. 63F). Medium size (>5 mm in length)............................... Pionosyllis (in part)
Genus
Amblyosyllis Grube, 1857
Amblyosyllis Grube, 1857: 186.
Gattiola Johnston, 1865: 195.
Nicotia Costa, 1864: 160.
Pterosyllis Claparède, 1863: 46.
Thylaciphorus Quatrefages, 1865: 55.
Pseudosyllides Czerniavsky, 1882: 173.
Type species. Amblyosyllis rhombeata Grube, 1857 by
monotypy.
Diagnosis. Body less than 5 mm in length, dorsoventrally
flattened, extremely fragile, with few segments; peri-
stomium and last segment without parapodia and chaetae,
each with 2 pairs of cirri. Intersegmental constrictions
strongly marked, midbody segments typically trapezoidal
in shape. Prostomium with 3 antennae, 4 eyes, and usually
2 anterior eyespots, sometimes ventrally located. Palps short,
less than length of prostomium, basally fused, divergent,
usually ventrally folded, and difficult to see dorsally.
Peristomium shorter than following segments, with 2 pairs

San Martín & Hutchings: Eusyllinae of Australia 261
of tentacular cirri, and 2 nuchal organs, forming nuchal
lappets, usually ciliated. Antennae, tentacular and dorsal
cirri long, greater than body width, usually strongly coiled,
sometimes forming skeins, smooth to indistinctly
articulated, fragile. Pigmentary glands on dorsal and ventral
cirri, sometimes forming distinct vesicles. Pharynx long,
slender, highly convoluted, with trepan formed by several
teeth, lacking median tooth. Proventricle proportionally
small to body width. Ventral cirri large, located latero-
posteriorly to parapodial lobes, similar in length. Compound
chaetae, heterogomph, bidentate falcigers, numerous,
present on all chaetigers, other types of chaetae rarely
present; chaetae similar in all species. Pygidium with 2 long
anal cirri, third length of dorsal cirri.
Remarks. Species belonging to this genus appear to be
uncommon and are typically known from damaged
incomplete specimens. Distinguishing species is therefore
difficult. The structure of the nuchal organs and trepan are
useful characters. The method of reproduction not well
known, although a mature male has been observed with
natatory chaetae and one species is known to brood eggs in
a gelatinous mass, suggesting that the species of this genus
are epigamic (Pernet, 1998).
Key to Australian species of Amblyosyllis
1 Nuchal lappets elongated, reaching middle of chaetiger 1 .............................. A. enigmatica n.sp.
—— Nuchal lappets rounded, small, not extending beyond peristomium................................................. 2
2 Dorsal cirri with few, large, distinct vesicles (Fig. 5A). Compound
chaetae with blades all similar, with small dorsoventral gradation
in length (Fig. 5C). Trepan with monocuspid teeth................................................... A. vesiculosa
—— Dorsal cirri with small, numerous granules (Fig. 3A). Compound
chaetae with blades elongated (50–102 µm in length) (Fig. 3D),
with distinct dorsoventral gradation in length. Trepan with 6 penta
cuspid teeth, 2 of them separated by single monocuspid tooth............... A. multidenticulata n.sp.
Amblyosyllis enigmatica n.sp.
Figs 1A–C, 2A–E
?Amblyosyllis granosa.—Augener, 1913: 243. Not Ehlers, 1897: 58.
Material examined. HOLOTYPE (AM W28943) AUSTRALIA:
NEW SOUTH WALES: NW corner of Bowen Is., Jervis Bay,
35°06.81'S 150°46.11'E, dense bryozoan community under
rock ledge, 13 m, coll. P. Serov & G.D.F. Wilson, 8 Dec 1993.
Additional material examined. Elizabeth Reef, Tasman Sea,
29°55.8'S 159°01.3'E, algae, reef flat, near wreck “Yoshin Maru
Iwaki”, 0.5 m, Elizabeth & Middleton Reefs Expedition, 1987,
14 Dec 1987, 6 + 3 on SEM stub (AM W28923).
Description. All specimens incomplete; holotype (Fig. 1A)
mature, epigamic specimen, with regenerating posterior end,
3.5 mm long, 0.6 mm wide, with 10 chaetigers plus
regenerating segments. Body large, segments trapezoidal
(Fig. 2A), especially those of midbody, fragile; some
specimens colourless, others strongly pigmented with 2–4
dorsal transverse dark bands on each segment, more marked
on midbody. Prostomium oval, posteriorly notched, with 4
large red eyes in trapezoidal arrangement, covering lateral
margins (Fig. 1A). Antennae long, inserted on anterior
margin of prostomium, with distinct ceratophores (Fig. 2B).
Palps free, fused basally, ventrally directed. Peristomium
shorter than subsequent segments, with 2 pairs of tentacular
cirri, dorsal ones twice as long as ventral; 2 ciliated,
elongated nuchal lappets (Figs 1A, 2A–C), extending to
middle of chaetiger 1, with dorsal, longitudinal row of
pigment (Fig. 1A). Dorsal cirri long, as wide as body,
indistinctly articulated, with numerous dark, granular
inclusions, irregularly arranged in 3 longitudinal rows (Fig.
1A). Parapodial lobes long, conical, with long, digitiform
prechaetal papilla (Fig. 2D). Ventral cirri conical, large,
broad, arising ventrolaterally (Fig. 2D), upwardly directed,
with granular, dark inclusions (Fig. 1A). Parapodia with
16–18 compound, heterogomph chaetae (Fig. 2E), blades
distinctly bidentate; within fascicle blades exhibiting
dorsoventral gradation in length (Fig. 1C), about 100 µm in
length dorsally, 43 µm in length ventrally, with short spines
on margin; about 6–8 aciculae per parapodium, straight,
distally pointed. Pharynx slender, with several coils (Fig.
1A); trepan composed of 6 teeth each with 5 denticles (Fig.
1B). Proventricle extending through 1.5 segments, with 11–
13 muscle cell rows. Holotype with notoacicula and capillary
notochaetae, curved, digitiform, fleshy, dark structure arising
from cirrophores from chaetiger 6 onwards (Fig. 1A, arrows);
also, some ciliary bands present on lateral areas of first 4
chaetigers (Fig. 1A). Curved structure present on holotype
missing on other material examined; function unknown.
Remarks. This species is characterized by having elongated
nuchal lappets and a trepan with 6 teeth, each with 5
denticles; no species of the genus has been described with
such a combination of characters. The general body form
is almost identical to Amblyosyllis granosa Ehlers, 1897,
from Magellan Strait and Galápagos Islands. Ehlers (1897)
originally described this species with the anterior margin
of the pharynx lacking teeth; subsequently, Westheide
(1974) reported the same species from Galápagos,
describing a trepan with 6 teeth, each formed by a long,
median cusp and 2 lateral, much smaller ones. Examination
of 1 specimen of the type series of Amblyosyllis granosa
from Punta Arenas (Chile) (NFMN 5318) did not reveal
any teeth on anterior margin of pharynx, as originally
described, so, we are describing the Australian material as
a new species A. enigmatica. A previous record of A. granosa
from Australia by Augener (1913) is also referred with
caution to A. enigmatica n.sp. Augener’s material is not
available for examination in the HZM and is presumed lost.
Amblyosyllis granosa, however, could be present in
Australian waters.

262 Records of the Australian Museum (2006) Vol. 58
Fig. 1. Amblyosyllis enigmatica n.sp. (A) anterior end, dorsal view; (B)
trepan; (C) compound chaetae, midbody. AM W28943. Scales: A 0.38
mm, B 48 µm, C 20 µm.
Amblyosyllis formosa (Claparède, 1863) from European
coasts is also similar to A. enigmatica n.sp., with regards to
body form and the presence of long nuchal lappets; this
species, however, has a trepan with 6 unidentate teeth (San
Martín, 2003).
Little is known about reproduction within Amblyosyllis.
The digitiform structures arising from cirrophores of
chaetiger 6 onwards have never been described before, and
their significance is unknown, but they may be used in
reproduction, since they are only present on the single epigamic
specimen examined. Knowing whether such structures occur
in other species of the genus would be of interest.
Habitat. Occurring in algae and colonial bryozoans, in
shallow depths.
Distribution. Australia (Western Australia, New South
Wales).
Etymology. The name of the new species is referred to the
enigmatic structure found on the cirrophores of the holotype.
Amblyosyllis multidenticulata n.sp.
Figs 3A–D, 4A–F
Amblyosyllis spectabilis.—Haswell, 1920 (in part): 108, pl. 13,
figs 4–10. Not Johnston, 1865: 195.
Material examined. HOLOTYPE (AM W502) AUSTRALIA:
NEW SOUTH WALES: Port Jackson, 33°51'S 151°16'E, Feb
1920, coll. W.A. Haswell, mounted on microscope slide.
PARATYPES NW corner of Bowen Is., Jervis Bay, 35°06.81'S
150°46.11'E, dense bryozoan community under rock ledge,
13 m, coll. P. Serov & G.D.F. Wilson, 8 Dec 1993, 3 on
SEM stub (AM W26323); Halfway Reef, 200 m S of
Sullivan Reef, Ulladulla, 35°21.42'S 150°29.31'E, red algae
on rock ledges at base of wall, 13 m, coll. A. Murray, 3
May 1997, 2 (AM W28229); Port Jackson, 33°50'S
151°16'E, 1 (AM W25245), mounted on slide, id. W.A.
Haswell as A. spectabilis.

San Martín & Hutchings: Eusyllinae of Australia 263
Fig. 2. SEM of Amblyosyllis enigmatica n.sp. (A) anterior end,
dorsal view; (B) detail of prostomium and anterior segments;
(C) detail of the nuchal lappets; (D) section of a midbody segment;
(E) compound chaetae, midbody. AM W28923.
Description. Body compact (Fig. 4A), longest complete
specimen examined 5.7 mm long, 0.6 mm wide, with 13
chaetigers and 1 posterior segment without parapodia or
chaetae. Colour pattern variable; holotype collected in 1920
colourless. More recently collected paratypes with several
dark, violet transverse dorsal bands of pigment on each
segment; intensity of bands differs between individuals.
Antennae, tentacular, dorsal and ventral cirri provided with
numerous, dark, small globular glands (Fig. 3A).
Prostomium wider than long, margins rounded, with 2 pairs
of eyes in open trapezoidal arrangement (Fig. 3A), and 2
anterior, ventrally located, small eyespots. Antennae long,
3 times width of prostomium, distinctly articulated, with
numerous articles, arising on anterior margin of prostomium
from distinct ceratophores (Figs 3A, 4B). Palps ventrally
directed (Fig. 4C), forming double lip over mouth, nearly
invisible dorsally (Fig. 3A). Peristomium about half of
length of subsequent segments, with 2 pairs of tentacular
cirri, similar in shape to antennae, dorsal tentacular cirri
longer than antennae and ventral tentacular cirri; 2 nuchal
lappets, small, rounded not extending beyond peristomium
(Figs 3A, 4B). Dorsal cirri similar to antennae and tentacular
cirri, usually coiled. Parapodia broad, long, conical, with
distinct prechaetal, digitiform papilla (Figs 3C, 4D). Ventral
cirri broad, long, located lateroposteriorly on parapodial
lobe. Compound chaetae numerous, up to 25–26 on
midbody parapodia, heterogomph, with long, distinctly
bidentate blades (Fig. 4E,F), with short spines on margin,
and dorsoventral gradation in length of blades (Fig. 3D),
length in midbody segments about 102 µm, 50 µm in length
ventrally on midbody. Parapodia with 5–6 aciculae, straight,
distally pointed. Pharynx long extending to chaetiger 5,
narrow width, with several coils visible inside pharynx (Fig.
3A); trepan formed by 6 large teeth with 5 denticles
separated by 1 unidentate, much smaller tooth (Fig. 3B).
Proventricle small, located on chaetiger 5 (Fig. 3A), with
17–18 muscle cell rows. Pygidium small, trapezoidal, with
2 long anal cirri, half length of dorsal cirri.
Remarks. Haswell (1920) referred his Australian specimens
to the European species Amblyosyllis spectabilis (Johnston,
1865), a poorly known species usually considered as a
synonym of A. formosa, which may represent a suite of
sibling species. Johnston’s description of Gattiola
spectabilis states that the pharynx is short and barrel shaped
without teeth and long nuchal lappets are illustrated. In
contrast, Haswell (1920) reported that teeth were present

264 Records of the Australian Museum (2006) Vol. 58
in the pharynx. Re-examination of this material has shown
that it belongs to a new species, Amblyosyllis multidentic-
ulata. This new species is characterized by having small
nuchal lappets and a trepan comprising 6 teeth each with 5
denticles, which are each separated by 1 small tooth (Fig.
3B); no other described species of this genus has been
described with this arrangement of teeth on the trepan. The
most similar species is Amblyosyllis madeirensis Langer-
hans, 1879, which occurs along European coasts; both
species have small nuchal lappets, similar colour pattern,
and body shape. The European species, however, has only
6 pentacuspid teeth on the trepan, and the segment that bears
the proventricle is distinctly larger than other segments (San
Martín, 2003). We are thus describing the Australian
material as a new species. Haswell (1920) also reported
some specimens 10 mm long, with 14–30 segments, these
were not available for examination and because of their size
we suggest that they probably represent another species.
Habitat. Occurring among bryozoans and others colonial
animals, in shallow depths.
Distribution. Australia (New South Wales).
Etymology. The specific name refers to the highly
denticulated trepan.
Fig. 3. Amblyosyllis multidenticulata n.sp. (A) anterior end, dorsal view; (B) trepan; (C) parapodial lobe and
ventral cirrus; (D) compound chaetae, midbody. A: AM W502 (holotype); B,C: AM W28229 (paratypes); D: AM
W26323 (paratypes). Scales: A 0.38 mm, B 48 µm, C 0.18 mm, D 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 265
Fig. 4. SEM of Amblyosyllis multidenticulata n.sp. (A) complete specimen; (B) nuchal lappets; (C) anterior end, ventral view; (D)
midbody parapodia, dorsal view; (E) superior compound chaeta; (F) inferior compound chaeta. AM W26323 (paratypes).
Amblyosyllis vesiculosa Hartmann-Schröder, 1989
Fig. 5A–C
Amblyosyllis vesiculosa Hartmann-Schröder, 1989: 25, figs 28–33.
Material examined. AUSTRALIA: NEW SOUTH WALES: Lake
Macquarie, 33°03'S 151°38'E algae & epibionts, intertidal, coll.
G. Hartmann-Schröder, holotype (HZM P-19661).
Description. Specimen complete, but broken in two pieces,
in good condition. Body small, 2 mm long, 0.2 mm wide,
with 13 chaetigers. Prostomium rounded; 4 eyes in
trapezoidal arrangement. Antennae inserted on anterior
margin of prostomium, median one with about 11 articles,
lateral ones with about 7 articles (Fig. 5A). Palps ventrally
folded, not visible dorsally. Nuchal lappets rounded,
extending to anterior margin of chaetiger 1 (Fig. 5A).
Peristomium visible dorsally; tentacular cirri similar to
antennae; dorsal tentacular cirri with about 12 articles,
ventral ones with 2–3 articles. Dorsal cirri about twice width
of body, distinctly articulated, with about 12–14 articles;
large, hyaline inclusions common throughout cirri and
antennae (Fig. 5A). Ventral cirri, rounded, similar length to
parapodial lobes, sometimes with large inclusions.
Parapodial lobes length similar to body width, blunt, distally
bilobed (Fig. 5B). Parapodia with compound chaetae, about
13–15 anteriorly, 9 posteriorly, heterogomph, with short,
bidentate blades, with small dorsoventral gradation in
length, with fine spines on margin (Fig. 5C). Pharynx,
strongly coiled, extending to chaetiger 5 (Fig. 5A); trepan

266 Records of the Australian Museum (2006) Vol. 58
Fig. 5. Amblyosyllis vesiculosa Hartmann-
Schröder, 1989 (A) anterior end, dorsal view;
(B) parapodial lobe and ventral cirrus; (C)
compound chaetae. ZMH P-19661. Scales: A
0.5 mm, B 0.2 mm, C 20 µm.
small, formed by 6 unidentate teeth (according to original
description by Hartmann-Schröder, 1989), not verified.
Proventricle on chaetiger 5, about length of segment, with
about 14 muscle cell rows. Pygidium compact, with 2 coiled
anal cirri.
Habitat. Occurring on algae, intertidally.
Distribution. Australia (New South Wales).
Genus Anoplosyllis Claparède, 1868
Anoplosyllis Claparède, 1868: 214.—San Martín, 2003: 134.
Type species. Anoplosyllis edentula Claparède, 1868, by
monotypy.
Diagnosis. Body small, (<5 mm in length), with up to 30
chaetigers. Prostomium rectangular, similar width to anterior
segments, with 2 pairs of eyes and 2 anterior eyespots. Three
antennae. Palps small, fused basally, without median furrow.
Nuchal organs as 2 ciliated grooves between prostomium
and peristomium. Two pairs of tentacular cirri. Antennae,
tentacular and dorsal cirri smooth, club-shaped, tapered
basally, longer than parapodial lobes. Compound chaetae
heterogomph, blades slender, elongate, unidentate or
indistinctly bidentate. Dorsal and ventral simple chaetae present
on some parapodia. Pharynx shorter than proventricle,
unarmed. Proventricle large, almost as wide as body. Some
species brood eggs in gelatinous masses.
Remarks. Fauvel (1923) synonymized Anoplosyllis with
Syllides. Species of Syllides, however, have long, articulated
cirri from chaetiger 3, and bidentate compound chaetae.
Although these genera are closely related, San Martín (2003)
proposed the resurrection of Anoplosyllis, and transferred
species of Syllides with smooth dorsal cirri into the genus.

San Martín & Hutchings: Eusyllinae of Australia 267
Anoplosyllis sexoculata (Hartmann-Schröder, 1962)
n.comb.
Fig. 6A–F
Syllides sexoculata Hartmann-Schröder, 1962: 97, figs 78–80;
1974: 128; 1989: 27.
Material examined. AUSTRALIA: NEW SOUTH WALES:
Between Liverpool Reach and Upper Half Moon Reach,
Hawkesbury R., 33°25'S 150°55'E, coarse sand, 20 m, coll.
A. Jones & party, 26 Aug 1980, 2 (AM W24699); Lower
Half Moon Reach, Hawkesbury R., 33°25'S 150°57'E, coll.
A. Jones & party, 26 Aug 1980, 1 (AM W24700); Tuross
R., at Princes Hwy, Bodalla, 36°04'S 150°04'E, coll. D.F.
Boesch, 23 May 1972, 1 (AM W7465). VICTORIA: Lakes
Entrance, 33°05'S 151°40'E, 1 (HZM P-20037). CHILE:
Niebla bei Valdivia, 39°46'S 73°150'W, holotype (HZM P-
14673), 3 paratypes (HZM P-14674).
Description. Body up to 2 mm long with 20 chaetigers;
complete specimen 1.4 mm long, 0.2 mm wide, 20
chaetigers. Prostomium oval, more than twice as wide as
long; 4 eyes arranged in open trapezoidal pattern and 2
distinct anterior eyespots, similar in size to eyes (Fig. 6A).
Antennae short, similar to width of prostomium, club-
shaped; median antenna inserted between posterior eyes,
lateral antennae near anterior margin of prostomium (Fig.
6A). Palps small, shorter than prostomium. Tentacular cirri
similar to antennae, dorsal ones longer than ventral ones.
Dorsal cirri similar to antennae and tentacular cirri (Fig.
6A), smooth, club-shaped, slightly longer than parapodial
lobes, some with distinct dark inclusion (Fig. 6B), which
opens via terminal pore. Parapodial lobes elongate; ventral
cirri digitiform, shorter than parapodial lobes (Fig. 6B).
Compound chaetae heterogomph, smooth, slender, blades
elongate, thin, unidentate, smooth (Fig. 6C), numbering 12–
18 per parapodium, with dorsoventral gradation in length
of blades, 48 µm in length dorsally, 29 µm in length
ventrally. Dorsal simple chaetae from chaetiger 1, slender,
unidentate, with minute spines on margin (Fig. 6D). Ventral
simple chaetae on posterior chaetigers, similar to dorsal
simple chaetae, but thinner and smooth (Fig. 6F). Aciculae
solitary, slender, distally rounded (Fig. 6E). Pharynx
partially everted, short, probably through 1–2 chaetigers.
Proventricle, massive, through 4 segments (Fig. 6A);
number of muscle cell rows not possible to assess. Pygidium
small, triangular, with 2 filiform anal cirri and compact
median papilla.
Fig. 6. Anoplosyllis sexoculata (Hartmann-Schröder, 1962) (A) anterior
end, dorsal view; (B) midbody parapodium; (C) compound chaetae,
midbody; (D) dorsal simple chaeta; (E) acicula; (F) ventral simple
chaeta. A,B: AM W24700; C–F: AM W7465. Scale: A 0.18 mm, B 48
µm, C–F 20 µm.

268 Records of the Australian Museum (2006) Vol. 58
Remarks. The Australian material is represented by a few
juvenile individuals and while they closely resembles the
type material of Anoplosyllis sexoculata from southern
Chile, they possess a shorter proventricle. At this stage we
are referring them to this species, but mature individuals
are really needed to confirm this.
Habitat. Occurring interstitially in fine to coarse sand, on
algae, on colonies of sabellariids; intertidally to about 20
m.
Distribution. Southern Chile, Namibia, Australia (Victoria,
New South Wales).
Genus Astreptosyllis Kudenov & Dorsey, 1982
Astreptosyllis Kudenov & Dorsey, 1982: 575.
Type species. Astreptosyllis acrassiseta Kudenov & Dorsey,
1982, by original designation.
Diagnosis. Body of meiofaunal size. Prostomium with 2
pairs of eyes and 3 antennae. Palps fused basally, small but
visible dorsally without median furrow. Nuchal organs as 2
ciliated grooves between prostomium and peristomium. Two
pairs of tentacular cirri. Antennae, tentacular cirri and dorsal
cirri of 2 anterior chaetigers unarticulated, club-shaped,
slender basally and distally slightly inflated, longer than
body width; from chaetiger 3 onwards, dorsal cirri
articulated, articles elongated to pyriform, with dark,
glandular inclusions. Parapodial lobes subrectangular on
anterior parapodia, conical and elongated from midbody;
ventral cirri short and broad on anterior parapodia,
posteriorly becoming more elongated, digitiform.
Compound chaetae heterogomph or hemigomph falcigers.
Dorsal simple chaetae thick, provided with distal,
longitudinally striated hood. Ventral simple chaetae
sometimes present on far posterior parapodia, but usually
lacking. Pharynx unarmed, with distal crown of soft papillae.
Pygidium with 2 anal cirri.
Remarks. The genus is only known from Australia. Details
on reproduction unknown, probably it reproduces as other
similar genera by epigamy.
Key to Australian species of Astreptosyllis
1 Compound chaetae similar throughout........................................................................ A. similiseta
—— Compound chaetae of chaetigers 1–6 enlarged, hemigomph, with
short, thick blades (Fig. 7E), different to compound chaetae of
other parapodia (Fig. 7G).......................................................................................... A. acrassiseta
Astreptosyllis acrassiseta Kudenov & Dorsey, 1982
Figs 7A–I, 8A–F
Astreptosyllis acrassiseta Kudenov & Dorsey, 1982: 576, fig. 1.
Material examined. AUSTRALIA: NEW SOUTH WALES: Off
Bass Point, 34°36'S 150°54'E, 50 m, 1 Feb 1990, 1 (AM
W22995). VICTORIA: Port Phillip Bay, 38°16.3'S 144°41.5'E,
medium sand, 13 m, Feb 1971, 1 paratype, (AM W18587).
WESTERN AUSTRALIA: Ned’s Camp, Cape Range National
Park, 21°59'S 113°55'E, limestone reef near shore, fine
sediment & sand from patches in reef, 1 m, coll. H.E. Stoddart,
2 Jan 1984, 1 (AM W26780).
Description. Body up to 3.1 mm long, 0.3 mm wide, with 42
chaetigers. Prostomium rectangular, 2 pairs of eyes in
trapezoidal arrangement; antennae slightly club-shaped,
unarticulated, usually missing; median antenna arising from
posterior margin of prostomium, between posterior eyes, lateral
antennae originating between anterior eyes. Palps large, similar
in length to prostomium, basally totally fused (Figs 7A, 8A),
directed anteriorly, provided with distal constriction. Tufts of
cilia on lateral areas of prostomium and peristomium (Fig.
7A). Tentacular cirri and dorsal cirri of subsequent 2 segments
similar to antennae; dorsal cirri from chaetiger 3 articulated
(Figs 7A, 8A), slender, slightly longer than body width, with
about 6–10 elongated articles, some with 1–2 granular, dark
inclusions (Fig. 7A,C). Parapodia of anterior segments
relatively broad in contrast, to posterior ones, distally truncated,
with some distal incisions (Fig. 7B); becoming elongated and
conical from proventricular segments onwards, with dorsal
band of long cilia (Fig. 7C). Ventral cirri of anterior parapodia
broad, shorter than parapodial length, becoming greatly
elongated posteriorly, longer than parapodial length, arising
from about middle of ventral side of parapodial lobes (Fig.
7C). Anterior 1–6 chaetigers each with 8–10, sometimes 12,
compound chaetae with enlarged shafts, hemigomph
articulation, with short, stout, broad blades, apparently
unidentate or bifid, with distal incision (Figs 7E, 8B), about
8–15 µm long. From chaetiger 7 onwards, parapodia with 5–8
compound chaetae with shafts much slender than those of
anterior chaetae, heterogomph, with fine spines on margin (Figs
7G, 8C,D), and blades slender, unidentate, some with distal
hood, with minute spines on margin, and dorsoventral gradation
in length of blades within fascicle (Fig. 8C), 27 µm in length
dorsally and 15 µm in length ventrally; under SEM, margin
with several rows of spines (Fig. 8E) and distal hood
composed of minute spines (Fig. 8E). Acicula solitary,
distally knobbed (Fig. 7B,H). Dorsal simple chaetae from
chaetiger 1, thick, tip tapered, blunt, covered with
longitudinally striated, rounded hood (Fig. 7D,F); under
SEM, dorsal simple chaetae with “Banksia-like” or
“artichoke-like” appearance. Ventral simple chaetae usually
absent, but one specimen with single thin, smooth, filiform,
ventral simple chaeta (Fig. 7 I ) on each fascicle of last
parapodium. Pharynx through about 7–8 segments, with crown
of 10 soft papillae (Fig. 7A). Proventricle through about 7–8
segments, with 30–32 muscle cell rows. No specimens with
anal cirri present, probably present on undamaged specimens.
Habitat. Occurring in median to coarse sand; from intertidal
to 50 m.
Distribution. Australia (Victoria, New South Wales,
Western Australia).

San Martín & Hutchings: Eusyllinae of Australia 269
Fig. 7. Astreptosyllis acrassiseta Kudenov & Dorsey, 1982 (A) anterior end, dorsal view (antennae and some
tentacular and dorsal cirri missing); (B) anterior parapodial lobe and acicula, dorsal view; (C) posterior parapodium;
(D) dorsal simple chaeta, anterior parapodium; (E) compound chaetae, anterior parapodium; (F) dorsal simple
chaeta, midbody; (G) compound chaetae, midbody; (H) acicula, midbody; (I) ventral simple chaeta. AM W26780.
Scales: A 0.18 mm; C 92 µm; B, D–I 20 µm.
Astreptosyllis similiseta Hartmann-Schröder, 1986
Fig. 9A–H
Astreptosyllis similiseta Hartmann-Schröder, 1986: 40, figs 16–21.
Material examined. AUSTRALIA: SOUTH A USTRALIA: Port
Augusta, 32°30'S 137°13'E, intertidal sediments, 6 Dec
1975, holotype (HZM P-18624) and 3 paratypes (HZM P-
18625), coll. G. Hartmann-Schröder.
Description. Body, delicate, fragile, entire, without colour
markings, 1.8–2.0 mm long, 0.1 mm wide, with 23–28
chaetigers. Prostomium semi-circular, with 4 eyes in open
trapezoidal arrangement; antennae fusiform to club-shaped,
smooth; median antenna longer than combined length of
prostomium and palps, inserted on middle of pharynx,
between anterior eyes; lateral antennae similar in shape to
median antenna, but shorter, inserted near anterior margin
of prostomium (Fig. 9A). Palps small, shorter than

270 Records of the Australian Museum (2006) Vol. 58
Fig. 8. SEM of Astreptosyllis acrassiseta Kudenov & Dorsey, 1982 (A) anterior end, dorsal view (antennae and some tentacular and
dorsal cirri missing); (B) compound chaetae, anterior parapodium; (C) compound chaetae, midbody; (D) dorsalmost compound
chaeta, posterior parapodium; (E) detail of tip of the same; (F) detail of distal end of dorsal simple chaeta, midbody. AM W26780.
prostomium, fused basally (Fig. 9A). Peristomium similar
in size to subsequent segments; tentacular cirri smooth,
slightly club-shaped (Fig. 9A); dorsal tentacular cirri similar
in length to lateral antenna, ventral ones shorter than dorsal
ones. Dorsal cirri of anterior 2 chaetigers similar in shape
to antennae and tentacular cirri, and slightly longer; dorsal
cirri from chaetiger 3 onwards articulated, with some dark
granular inclusions on some articles (Fig. 9A,B), 5–7 articles
on midbody chaetigers. Parapodia slender, elongated;
ventral cirri digitiform, longer on posterior parapodia (Fig.
9B) becoming of similar length to parapodial lobes.
Compound chaetae heterogomph falcigers, with conspicuous
spines subdistally on shafts, and unidentate blades with short,
apparently smooth spines on margin. Anterior parapodia
(1–5 chaetigers) each with several compound chaetae, with
slight dorsoventral gradation in length of blades within
fascicle, 40 µm in length dorsally, 20 µm in length ventrally
(Fig. 9D); on midbody blades of compound chaetae longer,
more marked gradation (65 µm dorsally and 23 µm
ventrally) (Fig. 9F); posterior parapodia with less marked
dorsoventral gradation in length of blades within fascicle
(40 µm dorsally and 23 µm ventrally) (Fig. 9G). Dorsal
simple chaetae thick, distally unidentate, with distinct distal
hood longitudinally striated, similar throughout (Fig.
9C,E,H), present from anterior parapodia. Ventral simple
chaetae absent. Pharynx through about 5 segments;
proventricle through 4.5 segments (Fig. 9A), with 38–40
muscle cell rows. Pygidium rounded, with single digitiform

San Martín & Hutchings: Eusyllinae of Australia 271
Fig. 9. Astreptosyllis similiseta Hartmann-Schröder, 1986
(A) anterior end, dorsal view; (B) posterior end, dorsal view;
(C) dorsal simple chaeta, anterior parapodium; (D) dorsal
and ventralmost compound chaetae, anterior parapodium;
(E) dorsal simple chaeta, midbody; (F) compound chaetae,
midbody; (G) compound chaetae, posterior parapodium; (H)
dorsal simple chaeta, posterior parapodium. ZMH P-18624.
Scales: A,B 0.1 mm; C–H, 20 µm.
anal cirri, similar in shape and length to posterior ventral
cirri (Fig. 9B).
Habitat. Occurring on mud flats with mangroves.
Distribution. Australia (South Australia).
Genus Eusyllis Malmgren, 1867
Eusyllis Malmgren, 1867: 40.
Eudontosyllis Knox, 1960: 105.
Type species. Eusyllis blomstrandi Malmgren, 1867
Diagnosis. Body of medium to small size (10 to <5 mm in
length), cylindrical. Prostomium with 4 eyes and sometimes
2 anterior eyespots. Three antennae. Palps either entirely
free or fused just basally. Nuchal organs as 2 ciliated
grooves, ciliation extending sometimes to prostomium and
peristomium, as well as other segments. Two pairs of
tentacular cirri. Antennae, tentacular and dorsal cirri non-
articulated, sometimes rugose, with pseudoarticulate
appearance. Compound chaetae; dorsal and ventral simple
chaetae on some parapodia. Pharynx with mid-dorsal tooth,
usually large and conspicuous, and incomplete crown of
small denticles, anteriorly pointing. Pygidium with 2 anal
cirri. Reproduction by epigamy (Garwood, 1991).
Remarks. Eusyllis is characterized by having an incomplete
trepan on the pharynx, with all teeth anteriorly directed,
and a mid-dorsal, usually large, tooth. Species of this genus
are typically large, even so the denticles of the trepan are
small and sometimes difficult to observe, which can lead to
misidentifications. Eusyllis brevicirrata Knox & Cameron,

272 Records of the Australian Museum (2006) Vol. 58
1971 from New Zealand, reported in Australia by Hartmann-
Schröder (1985), reproduces by schizogamy (San Martín,
pers. observ., on Australian material) so we suggest that
this species represents a member of the subfamily Syllinae
probably an undescribed genus (San Martín et al., in prep.).
Australian material agrees with the original description based
on New Zealand material. Eudontosyllis has been recently
proposed as a synonymy of Eusyllis by Glasby et al., (in press).
Key to Australian species of Eusyllis
1 Ventral cirri of chaetiger 1 enlarged, flattened leaf-like (Fig. 15B) ........................... E. lamelligera
—— Ventral cirri of chaetiger 1 not enlarged, flattened leaf-like ......................................................... 2
2 Aciculae from midbody greatly enlarged, distally curved,
extending beyond parapodial lobes (Fig. 10G). Compound chaetae
distinctly bidentate (Fig. 10F)....................................................................................... E. assimilis
—— Aciculae not modified (Fig. 13E). Compound chaetae unidentate
(Fig. 13C) ........................................................................................................................ E. kupfferi
Fig. 10. Eusyllis assimilis Marenzeller, 1875 (A) anterior end, dorsal view; (B) midbody parapodium; (C) pharyngeal opening; (D)
aciculae, anterior parapodium; (E) dorsalmost compound chaetae, anterior parapodium; (F) remaining compound chaetae, anterior
parapodium; (G) acicula, posterior parapodium; (H) dorsal simple chaeta; (I) dorsalmost compound chaetae, posterior parapodium; (J)
remaining compound chaetae, posterior parapodium. AM W28878. Scales: A 0.4 mm, B 0.1 mm, C 0.1 mm, D–J 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 273
Eusyllis assimilis Marenzeller, 1875
Figs 10A–J, 11A–F, 12A–C
Eusyllis assimilis Marenzeller, 1875: 158, pl. 3, Fig. 2.—Fauvel,
1923: 294, fig. 112a–g.—San Martín, 2003: 114, figs 52, 53.
?Eudontosyllis aciculata Knox, 1960: 105, figs 113–117.
Material examined. AUSTRALIA: NEW SOUTH W ALES: SW
side of South Solitary Is. 30°12'S 153°16'E, coral rubble,
18 m, coll. R.T. Springthorpe, 24 Jun 1992, 1 (AM
W28942); Taupo Seamount, Tasman Sea, 33°14.21'S
156°10.68'E, rough marl bottom, 133 m, coll. J.K. Lowry
& party on RV “Franklin”, 2 May 1989, 2 on SEM stub
Fig. 11. SEM of Eusyllis assimilis Marenzeller, 1878 (A) anterior end, dorsal view; (B) midbody, dorsal view; (C) detail of
prostomium, dorsal view; (D) everted pharynx; (E) pharyngeal opening; (F) detail of the same (arrows show denticles). AM
W28878, W28879.
(AM W28878); Taupo Seamount, Tasman Sea, 33°16.85'S
156°09.15'E, limestone & sand, 244 m, coll. J.K. Lowry &
party on RV “Franklin”, 2 May 1989, 4 on SEM stub (AM
W28879); Taupo Seamount, Tasman Sea, 33°16.85'S
156°09.15'E, limestone & sand bottom, 244 m, coll. J.K.
Lowry & party on RV Franklin, 2 May 1989, 17 (AM
W28925). SOUTH AUSTRALIA: Lowly Point, 33°00'S
137°47'E, subtidal, 1 (AM W26357); 4 km NW of 5th Creek,
Port Pirie, Spencer Gulf, 33°12'S 137°55'E, subtidal,
Posidonia sp. and Amphibolus spp, 4.6 m, T.J. Ward & party,
Mar 1980, 1 (AM W28234). WESTERN AUSTRALIA: Red
Bluff, Kalbarri, 27°42'S 114°09'E, mixed coralline algae
on rocky shore, 3.5 m, coll. J.K. Lowry, 10 Jan 1984, several

274 Records of the Australian Museum (2006) Vol. 58
(AM W28941); Bundegi Reef near Point Murat, Exmouth
Gulf, 21°49'S 113°11'E, orange finger sponge, 9 m, coll.
J.K. Lowry, 4 Jan 1984, WA-398, 1 (AM W26738).
Additional material examined. Spain: Off Cabo Cros,
Soller, Mallorca, Balearic Is., 39°38.50'N 02°25.13'E,
dredged 59–61 m, 6 (MNCNM 8637); Off Ribadeo, Lugo,
Galicia, 43°40.59'–43°40.25'N 7°02.77'–7°04.35'E, dredged
114–116 m, 8 (MNCNM 8643).
Description. Material examined small to medium size (10
to <5 mm length), described specimen 5 mm long, 0.54
mm wide, with 41 chaetigers (all incomplete specimens),
Fig. 12. SEM of Eusyllis assimilis Marenzeller, 1878 (A) dorsal compound chaeta, midbody; (B) compound chaetae, midbody; (C)
compound chaeta, posterior parapodium. SEM of Eusyllis kupfferi Langerhans, 1879 (D) anterior end, dorsal view; (E) detail of
prostomium and anteriormost segments, dorsal view; (F) anterior end, ventral view. A–C: AM W28878, W28879, D–F: AM W28952.
fragile. Elsewhere; specimens exceeding 40 mm in length
reported. Prostomium oval, deeply incised posteriorly,
forming 2 distinct lobes, densely ciliated on margin (Figs
10A, 11A,C); 4 eyes in open trapezoidal arrangement;
antennae long, slender, much longer than combined length
of prostomium and palps, irregularly pseudoarticulated;
lateral antennae inserted near anterior margin of pro-
stomium, median antenna arising slightly posteriorly to
lateral ones (Figs 10A, 11A,C). Palps broad, similar in length
to prostomium or slightly longer. Peristomium shorter than
subsequent segments; dorsal tentacular cirri long, similar
in shape but longer than median antenna, ventral tentacular
cirri about quarter length of dorsal ones. Occipital flap

San Martín & Hutchings: Eusyllinae of Australia 275
present (Fig. 11A,C). Dorsal cirri of chaetiger 1 elongated,
longer than dorsal tentacular cirri, also indistinctly
articulated; following dorsal cirri irregularly alternating long
cirri, slightly longer than half of body width, and others
distinctly shorter, all smooth, elongated, tapered (Figs 10A,
11B). Parapodia with prechaetal lobes. Parapodia dorsally
with cilia; other groups of cilia dorsally along body close
to dorsal cirri (Figs 10B, 11B). Ventral cirri triangular,
shorter than parapodial lobes (Fig. 11B). Compound chaetae
heterogomph falcigers, with shafts distally covered in
numerous, thin spines. Two different types of compound
chaetae present, one with slender, bidentate blades, both
teeth similar, and short spines on margin (Figs 10E,I, 12A),
located dorsally, others with shorter and larger blades,
strongly bidentate, both teeth similar on anterior parapodia
(Figs 10F, 12B), on posterior parapodia proximal tooth
becoming longer and stouter, located more ventrally within
bundles (Figs 10J, 12C). Anterior parapodia with 3
compound chaetae of slender type, blades about 20–22 µm
in length, and 19–20 compound chaetae of broad type,
blades about 15 µm in length. Posteriorly along body,
number of compound chaetae decreases, posterior parapodia
with 1–2 compound chaetae of slender type, blades 15–22
µm in length, and 6 of broad type, with prominent proximal
tooth, and minute spines or smooth on margin, about 16
µm in length. Dorsal simple chaetae slender, capillaries,
unidentate, with minute subdistal spines on margin (Fig.
10H), present only on posterior parapodia. Ventral simple
chaetae not seen. Anterior parapodia with 2 slender aciculae,
with slightly bent tip (Fig. 10D); posteriorly single aciculum
present, thicker than anterior ones, distally bent and
extending beyond parapodial lobes (Fig. 10G). Pharynx
through about 7–8 segments; opening surrounded by a
crown of 17–20 soft papillae and dense layer of cilia (Fig.
11D–F); plus crown of 10 smaller papillae basally (Fig.
11D); pharyngeal tooth large, located on anterior margin
Fig. 13. Eusyllis kupfferi Langerhans, 1879. (A) anterior end, dorsal view; (B) dorsal compound chaetae,
midbody; (C) remaining compound chaetae, midbody; (D) dorsal simple chaeta; (E) acicula; (F) anterior
end of pharynx; (G) parapodium, midbody. A,F: AM W28404; B–G: AM W28408. Scales: A 0.2 mm, B–
E 20 µm, F 0.1 mm, G 0.18 mm.

276 Records of the Australian Museum (2006) Vol. 58
(Fig. 10C); trepan incomplete, with variable number of teeth,
ranging from few to up to 35 separate teeth present (Fig.
11F, arrows). Proventricle elongate, through about 10
segments, tapering posteriorly, with about 50 muscle cell rows.
Remarks. This species has been reported from a wide range
of habitats and depths and material should be re-examined
and a molecular study would be useful to confirm if this is
a widely distributed species or a suite of sibling species.
We believe that Eudontosyllis aciculata Knox, 1960, is
identical with E. assimilis based on the description. We were
unable, however, to borrow this material.
Habitat. Occurring in all kind of substrates, from intertidal
to more than 500 m depth.
Distribution. North Atlantic, Mediterranean Sea, North
Pacific, New Zealand, Australia (Western Australia, South
Australia, New South Wales).
Eusyllis kupfferi Langerhans, 1879
Figs 12D–F, 13A–G, 14A–E
Eusyllis kupfferi Langerhans, 1879: 552, Fig. 14.—San Martín,
1990: 607, figs 12, 13.
Eusyllis autolytoides Hartmann-Schröder, 1991: 33, figs 47–53.
Odontosyllis multidentata Hartmann-Schröder, 1982: 64, figs 41–
46; 1990: 51, fig. 17.
Fig. 14. SEM of Eusyllis kupfferi Langerhans, 1879 (A) right
section of a midbody segment; (B) dorsal simple chaeta; (C)
dorsalmost compound chaeta, anterior parapodium; (D,E) short
compound chaetae, anterior parapodium. AM W28952.
Material examined. AUSTRALIA: QUEENSLAND: SW Reef, Heron
Is. 23°27'S 151°55'E, reef margin, algae, coral sand & coral debris, 3
Feb 1976, 2 paratypes of Eusyllis autolytoides Hartmann-Schröder, 1991,
(AM W20387). NEW SOUTH WALES: Off old wharf, Richmond R., near
Ballina, 28°52.5'S 153°33.6'E, drift algae, 6 m, coll. S.J. Keable, 5 Mar
1992, 4 (AM W28218); S ledge, Cook Is. 28°11.65'S 153°34.63'E,
surface of massive sponges, 14 m, coll. R.T. Springthorpe, 9 Jun 1993,
1 (AM W28402); S ledge, Cook Is. 28°11.65'S 153°34.63'E, Halimeda
sp., 13 m, coll. E.L.A. Ho, 9 Jun 1993, 1 (AM W28404); SW side of
South Solitary Is. 30°12'S 153°16'E, coral rubble, 18 m, coll. R.T.
Springthorpe, 24 Jun 1992, few (AM W28215); 1 km S of E end of
Spectacle Is. Hawkesbury R., 33°32'S 151°07.5'E, muddy sand, 12 m,
coll. A.R. Jones & A. Murray, 8 May 1984, 1 (AM W22149); Green Pt.
Croppy Pt. Hawkesbury R., 33°33.5'S 151°14.5'E, mud, 6 m, coll. A.
Jones & party, 22 Feb 1980, 1 (AM W196604); E end of Brooklyn Boat
Channel, Hawkesbury R., 33°33'S 151°14'E, coll. A. Jones & party, 18
Dec 1979, 1 (AM W196606); E end of Brooklyn Boat Channel,
Hawkesbury R., 33°33'S 151°14'E, coll. A. Jones & party, 16 May 1980,
1 (AM W196607); E end of Brooklyn Boat Channel, Hawkesbury R.,

San Martín & Hutchings: Eusyllinae of Australia 277
33°33'S 151°14'E, A. Jones & party, 16 May 1980, 1 (AM W196609); E
end of Brooklyn Boat Channel, Hawkesbury R., 33°33'S 151°14'E, coll.
A. Jones et al., 1 Aug 1979, 2 (AM W196611); E end of Brooklyn Boat
Channel, Hawkesbury R., 33°33'S 151°14'E, coll. A. Jones et al., 18
Dec 1979, 1 (AM W196612); E end of Brooklyn Boat Channel,
Hawkesbury R., 33°33'S 151°14'E, coll. A. Jones & party, 18 Dec 1979,
1 (AM W196613); Barrenjoey Head, Broken Bay, 33°35'S 151°20'E,
algae on rocky substrate, 4 m, coll. J.K. Lowry & party 22 Apr 1983, 1
(AM W28408). WESTERN AUSTRALIA: N end of beach, Bundegi Reef,
Exmouth Gulf, 21°49'S 114°11'E, rocky rubble, sticky sediment & brown
algae with epiphytes, 2 m, coll. H.E. Stoddart, 4 Jan 1984, many + 2 on
SEM stub (AM W28952); N end of beach, Bundegi Reef, Exmouth Gulf,
21°49'S 114°11'E, rocky rubble, coralline algae with green epiphyte,
1.5 m, coll. H.E. Stoddart, 4 Jan 1984, several (AM W28953).
Description. Body up to 4.5 mm long, 0.3 mm wide, with
40 chaetigers, slightly dorsoventrally flattened, dorsally
convex along median line of body, with well marked
segments. Distinct colour pattern, with 1 transverse band
of red pigment, sometimes 2 bands, and additional narrow
bands often laterally located on posterior half of segments
(Fig. 13A). Material examined exhibiting great variation in
intensity of pigmentation, from highly pigmented to
colourless individuals. Prostomium twice as large as 2nd
segment, pentagonal to quadrangular, with nuchal notch,
ciliated on both sides (Fig. 13A), double semicircle of cilia
on anterior part of prostomium (Fig. 12D,E); 4 eyes in open
trapezoidal arrangement, occasionally 2 anterior eyespots.
Antennae, tentacular and dorsal cirri fusiform, smooth.
Lateral antennae arising in front of anterior eyes, length
similar to combined length of prostomium and palps, median
Fig. 15. Eusyllis lamelligera Marion & Bobretzky, 1875 (A) anterior end, dorsal view; (B) prostomium
and anteriormost segments, ventral view; (C) pharyngeal armature; (D) compound chaetae, anterior
parapodia; (E) aciculae, anterior parapodia; (F) compound chaetae, midbody; (G) compound chaetae,
posterior parapodia; (H) acicula, posterior parapodia; (I) dorsal simple chaeta; (J) ventral simple chaeta.
AM W28416. Scales: A 0.4 mm, B 0.18 mm, C 0.1 mm, D–J 20 µm.

278 Records of the Australian Museum (2006) Vol. 58
antenna typically much longer than lateral antennae, arising
slightly posteriorly to lateral antennae (Figs 12D,E, 13A).
Palps broad (Fig. 12F), shorter than prostomium.
Peristomium shorter than following segments, sometimes
covered partially by prostomium and fold of chaetiger 1,
provided with some cilia (Figs 12D,E, 13A); dorsal
tentacular cirri shorter than median antenna but longer than
lateral antennae; ventral tentacular cirri similar to lateral
antennae. Dorsal transverse row of cilia on each segment
(Figs 12D, 14A); plus some cilia on dorsal surface of
parapodial lobes (Fig. 14A). Dorsal cirri of chaetiger 1 long,
similar in length to median antenna, longer than body width;
remaining dorsal cirri irregularly alternating between cirri
similar in length to half of body width, and cirri much shorter
(Figs 12D, 13A, 14A). Parapodia short, about third of body
width, thick, conical. Ventral cirri broad, shorter than
parapodial lobes or similar in length (Figs 13G, 14A).
Compound chaetae heterogomph falcigers, shafts with
several thin subdistal spines; blades proportionally short,
unidentate, with rounded tips, and short spines on margin
(Fig. 13B,C). Anterior parapodia with 2 kinds of compound
chaetae, one with relatively thin blades, 3 in anterior
segments, declining to 1 from midbody onwards (Figs 13B,
14C), blades 20–25 µm in length, located dorsally, other
type with thicker and shorter blades (Figs 13C, 14D,E),
about 17 µm in length, all similar, 12–14 per parapodium.
Dorsal simple chaetae bayonet-shaped (Figs 13D, 14B),
from proventricle segments onwards. Ventral simple chaetae
absent. Acicula solitary, distally knobbed, apparently hollow
at tip (Fig. 13E). Pharynx through 4–5 segments; opening rim
smooth on one half and other half with small denticles (Figs
13A,F); material examined exhibiting considerably range in
numbers of denticles present, from few to 20–24. Pharyngeal
tooth large, conical to rhomboidal; pharyngeal opening
provided with crown of about 14 soft papillae (Fig. 13F).
Proventricle longer than pharynx, through 7–8 segments, with
about 50 muscle cell rows. Pygidium with 2 slender, anal cirri.
Remark. Material from all localities listed in the distribution
has been examined by the senior author during other studies,
and no morphological differences between them can be
found. Molecular studies may be useful to confirm if this is
a widely distributed species or a suite of sibling species.
Specimens described as Eusyllis autolytoides by Hartmann-
Schröder (1991) and Odontosyllis multidentata (Hartmann-
Schröder, 1982) agree with the description of Eusyllis
kupfferi, so we consider them as synonymous.
Distribution. Portugal (Madeira), Canary Is., Cuba, Australia
(Queensland, Western Australia, New South Wales).
Habitat. Occurring on hydroids, Rhizophora mangle roots,
on algae, algae with sand, debris, dead corals, sand and
mud, from intertidal to about 20 m.
Eusyllis lamelligera Marion & Bobretzky, 1875
Figs 15A–J, 16A–F
Eusyllis lamelligera Marion & Bobretzky, 1875: 33, pl. 3, figs.
9A–C.—Fauvel, 1923: 294, Fig. 113.—San Martín, 2003: 117,
figs 54, 55.
Eusyllis dentata Hartmann-Schröder, 1990: 50, figs 13–16.
Eusyllis habei Imajima, 1966: 97, text-fig. 31a–k.
Material examined. AUSTRALIA: NEW SOUTH WALES: N side of
Bannister Head, N of Ulladulla, 35°19.15'S 150°29.12'E, grey sponge
from top of boulder, 18 m, coll. K. Attwood, 6 May 1997, 1 (AM
W28960); SW of Bowen Is., Jervis Bay, 35°07.49'S 150°45.77'E, small
white sponge with pink lobes, from seagrass bed, 7 m, coll. P. Serov &
G.D.F. Wilson, 8 Dec 1993, several (AM W28416); SW of Bowen Is.,
Jervis Bay, 35°07.49'S 150°45.77'E, rock on sandy bottom covered in
bryozoa & polychaetes, 7 m, coll. P. Serov & G.D.F. Wilson, 8 Dec
1993, few (AM W28415); Montagu Roadstead, Jervis Bay, 35°02.2'S
150°46.0'E, unvegetated sediment, 12 m, coll. P.A. Hutchings & party,
6 Jun 1990, 1 (AM W28231); NW corner of Bowen Is., Jervis Bay,
35°06.81'S 150°46.11'E, dense bryozoans under rock ledge, 13 m, coll.
P. Serov & G.D.F. Wilson, 8 Dec 1993, several (AM W28959); Darling
Road, near anchorages, Jervis Bay, 35°7.3'S 150°44.1'E, 18 m, coll. P.A.
Hutchings, 23 Jan 1973, 2 (AM W28437); Taupo Seamount, Tasman
Sea, 33°16.85'S 156°09.15'E, limestone & sand bottom, 244 m, coll.
J.K. Lowry & party, on RV “Franklin”, 2 May 1989, 5 on SEM stub
(AM W28877); Taupo Seamount, Tasman Sea, 33°16.85'S 156°09.15'E,
limestone & sand bottom, 244 m, coll. J.K. Lowry & party on RV
“Franklin”, 2 May 1989, many (AM W28927); Reef flat, near wreck
“Yoshin Maru Iwaki”, Elizabeth Reef, Tasman Sea, 29°55.8'S 159°01.3'E,
algae, 0.5 m, coll. Elizabeth & Middleton Reefs Expedition, 1987, 14
Dec 1987, several (AM W28928). TASMANIA: Woodchip Jetty, Spring
Bay, Triabunna, 42°30'S 147°55'E, muddy bottom, 4 m, coll. D. Cropp,
Nov 1982, 1 (AM W199178). SOUTH AUSTRALIA: Billy Lights Point,
Port Lincoln, 34°45'S 135°53'E, stone washings from sheltered intertidal
rocks, coll. I. Loch, 15 Feb 1985, 1 (AM W28929). WESTERN A USTRALIA:
Goss Passage, Beacon Is. 28°25.5'S 113°47'E, dead plates of Acropora
coral covered in coralline algae, 24 m, P.A. Hutchings, 21 May 1994, 1
(AM W28374); N end of Long Is. 28°27.9'S 113°46.3'E, dead coral
substrate covered in coralline & brown algae, 5.5 m, coll. C. Bryce, 22
May 1994, several (AM W28961).
Additional material examined Eusyllis dentata AUSTRALIA: NEW
SOUTH WALES: Angourie Point, south of Yamba, 29°29'S 153°22'E algae,
intertidal, holotype (HZM P-19963), 8 paratypes (HZM P-19964), 1
(HZM P-203313). Eusyllis habei JAPAN: Bos Peninsula, Hubara Port,
1, coll. E. Nishi. Eusyllis lamelligera SPAIN: off Punta Jovo, W. W
Menorca, Balearic Is. 10 m, algae 43°40.27'–43°40.06'N 5°13.36'–
5°14.35'E, 6 (MNCNM 8675); NW Cabo de Lastres, Asturias, Cantabrian
Sea, NE Atlantic, 39°49.66'–39°47.64'N 2°40.78'–2°38.71'E, 146 m, 8
(MNCNM 8688).
Description. Complete specimen 6.3 mm long, 0.5 mm
wide, with 50 chaetigers. Prostomium semi-circular; 4 eyes
in open trapezoidal arrangement and 2 anterior eyespots.
Median antenna more than twice combined length of
prostomium and palps, inserted on middle of prostomium,
lateral antennae shorter than median antenna, inserted in
front of anterior eyes, near anterior margin (Fig. 15A). Palps
broad, similar in length to prostomium (Fig. 15A). Nuchal
organs with distinct ciliation, extending to posterior margin
of prostomium (Figs 15A, 16A). Peristomium similar to
subsequent segments; dorsal tentacular cirri long and
slender, more than twice as long as body width, ventral
tentacular cirri about third length of dorsal cirri. Antennae,
tentacular and dorsal cirri smooth, long, slender, distally
tapered (Fig. 15A), sometimes rugose to weakly pseudo-
articulate (Fig. 16B). Dorsal cirri of chaetiger 1 longer than
dorsal tentacular cirri, remaining dorsal cirri irregularly
alternating long, and short cirri, long cirri much longer than
body width, and short ones, slightly shorter than body width
(Fig. 15A). Ventral cirri of chaetiger 1 enlarged, flattened
leaf-like; following ones conical (Figs 15B, 16B); ventral
cirri with minute pores (Fig. 16D). Lateral tufts of cilia on
each segment (Fig. 16C). Compound chaetae heterogomph,
with subdistal spines on shafts and bidentate blades, both
teeth similar in size, well separated and rounded margin
between both teeth, and fine spines on margin (Figs.
15D,F,G, 16E,F). Anterior parapodia with about 20–25
compound chaetae, with strong dorsoventral gradation in

San Martín & Hutchings: Eusyllinae of Australia 279
size of blades within fascicle, about 40 µm in length dorsally,
21 µm in length ventrally. Progressively along body, number
of compound chaetae per parapodium decreases, to 13–15
on midbody, 10 on posterior parapodia, with thicker shafts
and blades, with less marked dorsoventral gradation in
length of blades; on midbody, blades about 35 µm in length
dorsally, 22 µm in length ventrally, and 30 µm in length
dorsally, and 15 µm in length ventrally on posterior
parapodia (Fig. 15D,F,G). Dorsal simple chaetae on
posterior parapodia, slender, bidentate, both teeth similar,
with short spines on margin (Fig. 15I). Ventral simple
chaetae on far posterior parapodia, similar to dorsal simple
chaetae, but larger and teeth well separated (Fig. 15J).
Anterior parapodia with 2 aciculae, one distally knobbed
with acute tip, other with tricuspid tip, with lateral ones
poorly developed (Fig. 15E); medium and posterior
parapodia each with single acicula, distinctly tricuspid (Fig.
15H). Pharynx through about 7 segments; pharyngeal tooth,
conical, on anterior margin; trepan variable, ranging from
few small teeth present to distinct incomplete crown of up
to 24 teeth (Figs 15C, 16B).
Remarks. This species is characterized by having enlarged,
flattened leaf-like ventral cirri on chaetiger 1, tricuspid
aciculae, and compound chaetae with bidentate blades, with
both teeth of similar size and well separated by rounded
Fig. 16. SEM of Eusyllis lamelligera Marion & Bobretzky, 1875 (A) prostomium; (B) everted pharynx and anteriormost segments,
ventral view; (C) midbody segments, dorsal view; (D) detail of first and second ventral cirri; (E,F), compound chaetae. AM W28877

280 Records of the Australian Museum (2006) Vol. 58
margin. The Australian specimens were described as Eusyllis
dentata by Hartmann-Schröder (1990); and these were
examined and they are indistinguishable from Mediterranean
specimens of Eusyllis lamelligera. This species has been
reported from a wide range of habitats and depths and
material should be re-examined and a molecular study
would be useful to confirm if this is a widely distributed
species or a suite of sibling species.
Eusyllis habei Imajima, 1966, from Japan, is also similar
according to the description and may represent the same
species according to Imajima (1966). The material examined
from Bos Peninsula, Japan appears to be similar to
specimens of E. lamelligera from the Mediterranean and
also to material from the Atlantic and Australia.
Habitat. Occurring in a wide variety of substrates and
depths, intertidally to depths greater than 500 m.
Distribution. North Atlantic, Mediterranean
Sea, Australia (New South Wales,
Tasmania, Western Australia).
Genus Nudisyllis Knox & Cameron, 1970
Nudisyllis Knox & Cameron, 1970: 77, emended.
Type species. Nudisyllis tinihekea Knox & Cameron, 1970.
Diagnosis. Body small, fragile, <5 mm in length.
Prostomium large, about width of 2 segments, with 4 eyes
and sometimes pair of anterior eyespots. Three antennae.
Palps separated, free from each other, sometimes adjacent
at base. Median antenna inserted on middle of prostomium
or anteriorly, on line with lateral antennae. Two pairs of
tentacular cirri. Nuchal organs as 2 ciliated grooves between
prostomium and peristomium. Dorsal cirri on all chaetigers,
long, cylindrical, smooth or slightly rugose. Ventral cirri
not modified. Compound chaetae with both unidentate and
bidentate blades on same parapodium, or all unidentate;
bidentate blades having proximal tooth small, spine-like.
Tips of aciculae tricuspidate or lancet-shaped. Pharynx
small, shorter than proventricle; pharyngeal tooth located
near anterior rim, usually long. Pygidium with 2 anal cirri.
Reproduction by epigamy, probably with dorsal
incubation of eggs (only reported on two species)
(Pierantoni, 1905; Augener, 1913).
Remarks. Nudisyllis was described as being
characterized by a lack of antennae, tentacular
and dorsal cirri. Loss of appendages during
fixation and manipulation is usual in some
species, suggesting that these structures are
absent when in fact they are present. Several
of the specimens examined lack all or some
appendages; those that lack them are
identical to the description and drawings of
Nudisyllis tinihekea, so we assume that are
all the same species; they type specimens
represent damaged specimens, having lost
appendages during fixation. Some other
species described as belonging to Pionosyllis
have the same characters, but differ from
Nudisyllis in being of minute size (<5 mm in
length), having free palps, short pharynx and
a long tooth, and compound chaetae with
unidentate to minutely bidentate blades. So,
we consider Nudisyllis a valid genus, and have
Fig. 17. Nudisyllis tinihekea Knox & Cameron, 1970 (A) anterior
end, dorsal view; (B) posterior end, dorsal view; (C) compound
chaetae, midbody; (D) dorsal simple chaeta; (E) acicula; (F) ventral
simple chaeta. Scales: A,B 68 µm; C–F 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 281
emended the diagnosis and the genus now includes some
species previously considered belonging to Pionosyllis, among
them P. pulligera (Krohn, 1852); P. magnidens Day, 1953;
and P. divaricata (Keferstein, 1862).
Nudisyllis tinihekea Knox & Cameron, 1970
Fig. 17A–F
Nudisyllis tinihekea Knox & Cameron, 1970: 77, figs 6–9.
Pionosyllis pulligera.—Augener, 1913: 221, pl. II, fig. 8, text-
fig. 29. Not Krohn, 1852: 251.
Pionosyllis samsonensis Hartmann-Schröder, 1980: 52, figs 39–
43; 1981: 32; 1982: 65. 1983: 130; 1984: 20; 1985: 69; 1986:
41; 1987: 38; 1991: 35.
Material examined. AUSTRALIA: NEW SOUTH WALES: Barrenjoey
Head, Broken Bay, 33°35'S 151°20'E, algae on rocky substrate, 4 m,
coll. J.K. Lowry & party 22 Apr 1983, 1 (AM W28966). WESTERN
AUSTRALIA: Red Bluff, Kalbarri, 27°42'S 114°09'E, seagrass in shallow
sand on rocky shore, 3.5 m, coll. R.T. Springthorpe, 10 Jan 1984, 2
(AM W28968); Red Bluff, Kalbarri, 27°42'S 114°09'E, mixed coralline
algae on rocky shore, 3.5 m, coll. J.K. Lowry, 10 Jan 1984, several (AM
W28970); Inshore limestone reef, Ned’s Camp, Cape Range National
Park, 21°59'S 113°55'E, frilly Caulerpa sp., 1 m, coll. J.K. Lowry, 2 Jan
1984, several (AM W28971); Inside outer Ningaloo Reef, Cape Range
National Park, 21°59.5'S 113°54.5'E, mixed algae, 2 m, coll. J.K. Lowry,
1 Jan 1984, 2 (AM W28364); Inshore limestone reef, Ned’s Camp, Cape
Range National Park, 21°59'S 113°55'E, sponge covered with epiphytes,
sediment & muddy worm tubes, 1.5 m, coll. R.T. Springthorpe, 2 Jan
1984, 2 (AM W28969); N end of beach, Bundegi Reef, Exmouth Gulf,
21°49'S 114°11'E, rocky rubble, coralline algae with green epiphyte,
1.5 m, coll. H.E. Stoddart, 4 Jan 1984, 1 (AM W28967).
Description. Body up to 3.1 mm long with 34 chaetigers,
according to literature, longest specimen examined 1.2 mm
long, 0.2 mm wide, with 16 chaetigers, extremely fragile,
several specimens lack most appendages, others having lost
all antennae, tentacular and dorsal cirri. Prostomium squared
to pentagonal, densely covered by long cilia on posterior margin
and laterals (Fig. 17A), 4 eyes in trapezoidal arrangement,
located laterally on prostomium; median antenna inserted on
middle of prostomium, fusiform, distally tapered, slightly
longer than combined length of prostomium and palps; lateral
antennae inserted on anterior margin, slightly shorter than
median antenna (Fig. 17A). Palps short, broad, oval.
Peristomium dorsally reduced, covered by chaetiger 1 (Fig.
17A); dorsal tentacular cirri long, about twice as long as median
antenna; ventral tentacular cirri about one third length of dorsal
ones. Dorsal cirri smooth, thick, distally tapered, long on
anterior segments, from proventricle posteriorly, cirri consist
of long and short cirri that alternate; long cirri longer than
body width, and short cirri, shorter than body width (Fig. 17A).
Parapodia conical, ending with rounded papilla, provided with
some tufts of cilia. Ventral cirri digitiform, shorter than
parapodial lobes. Compound chaetae similar throughout, 15
anteriorly, 7 posteriorly, blades elongated, distally slightly
hooked, unidentate, longer ones with subdistal spine, smooth
or provided with fine spines on margin (Fig. 17C); dorsoventral
gradation in length within fascicle, about 42 µm in length
dorsally, 15 µm in length ventrally. Dorsal and ventral simple
chaetae on far posterior parapodia, slender, smooth, unidentate
(Fig. 17D,F). Acicula solitary, with tricuspidate tip (Fig. 17E).
Pharynx short, through 2 segments; single pharyngeal tooth
on anterior margin (Fig. 17A). Proventricle barrel-shaped,
through 3 segments, with about 20 muscle cell rows. Pygidium
semi-circular, with 2 anal cirri, similar in shape and length to
long dorsal cirri (Fig. 17B).
Habitat. Occurring on coralline algae, mixed algae,
Caulerpa, seagrasses on sand, sand with debris, sponges
with epibionts in shallow depths.
Distribution. New Zealand, Australia (New South Wales,
Western Australia, South Australia, Victoria, Queensland).
Genus Odontosyllis Claparède, 1863
Odontosyllis Claparède, 1863: 47.
?Eurymedusa Kinberg, 1865: 61.
Parautolytus Ehlers, 1900: 213.
?Alluaudella Gravier, 1905: 372.
?Atelesyllis Pruvot, 1930: 39.
Pharyngeovalvata Day, 1951: 26.
Odontoautolytus Hartmann-Schröder, 1979: 112.
?Synpalposyllis Hartmann-Schröder, 1983: 132.
Umbellisyllis Sars, 1869: 254.
Type species. Syllis fulgurans Audouin & Milne Edwards,
1833, designated by Hartman, 1959.
Diagnosis. Body of variable size, from <5 mm to >10 mm
in length, with numerous segments, cylindrical, dorsally
highly convex, flattened ventrally. Prostomium with 4 eyes
and, sometimes, pair of anterior eyespots. Three antennae.
Palps broad, free for almost all their length, fused basally.
Peristomium usually reduced dorsally; 2 pairs of tentacular
cirri. Occipital flap present, usually well developed,
covering peristomium dorsally and prostomium partially.
Nuchal organs as 2 ciliated grooves between prostomium
and peristomium, extending sometimes to lateral areas of
prostomium. Dorsal cirri elongated, smooth, distally
tapered, but sometimes short or indistinctly articulated.
Parapodia usually with pre- and postchaetal lobes. Ventral
cirri digitiform to pillow-shaped. Compound chaetae
heterogomph, usually with shafts distally spinose. Dorsal
and ventral simple chaetae present on some parapodia.
Pharynx short, distinctly shorter than proventricle, provided
with several teeth, usually few, pointing backwards,
pharyngeal mid-dorsal tooth absent; pharynx when not
everted situated posteriorly to chaetiger 1, inside tube that
leads to mouth opening on peristomium. Proventricle
usually long and wide, massive. Pygidium with 2 anal cirri.
Reproduction by epigamy; epigamic specimens sometimes
strongly modified and phosphorescent (Fischer & Fischer,
1995; Gaston & Hall, 2000).
Remarks. The structure of the anterior part of the gut in
this genus is distinct and unusual, with the pharynx being
short and the opening located posteriorly to the mouth; a
tube is present that surrounds the pharynx and that leads to
the mouth. This must be removed to see the trepan and the
two lateral plates of the pharynx. Occasionally specimens
lack any teeth on the trepan, and we suggest that individuals
can regenerate these teeth. The genera that we have
synonymized with Odontosyllis have been inadequately
examined previously with regard to the structure of the
anterior part of the gut. For example, GSM examined one
paratype of Pharyngeovalvata natalensis (BMNH
1961.16.16-17, Natal Shore, South Africa Stn. 47.8, coll:
J.H. Day) and it is identical to specimens of Odontosyllis
ctenostoma Claparède, 1868. Both species possess a
pharynx with a trepan; even though the description of the

282 Records of the Australian Museum (2006) Vol. 58
pharynx of Pharyngeovalvata by Day (1951) states that the
pharynx lacks a trepan, the paratype possesses one. We
therefore have synonymized these two species that are the
type species of their respective genera and Pharyngeo-
valvata is synonymized with Odontosyllis. We suspect that
Atelesyllis Pruvot, 1930, is also identical to Odontosyllis
with the pharynx contracted, but the types appear to be lost.
We have also examined the holotype of Synpalposyllis
australiensis Hartmann-Schröder, 1983 (HZM P-17400), a
species known only from the holotype, which is less than 5
mm in length, with the prostomium and the anterior part of
the pharynx damaged so it is not possible to verify the
presence or absence of the palps, but the chaetae and
aciculae are similar to those of O. australiensis. In our
opinion, this specimen may be a juvenile of O. australiensis.
Types of Alluaudella madagascariensis Gravier, 1905 were
not available for examination but this species appears to be
related to some species of Odontosyllis with short dorsal
cirri and indistinct ventral cirri, distally located, and partially
fused, appearing as absent, although this should be checked
on material from the type locality. We have examined type
material of the other species of the genus, Alluaudella
longicirrata Mohammad, 1973, and it is clearly a species
of Odontosyllis. We suggest that it is likely that the genus
Alluaudella should be synonymized with Odontosyllis, and
it certainly does not belong in the subfamily Autolytinae
where it has previously been placed (Gravier, 1905).
The key below includes nine species, an additional two
species have been reported from Australia, but they are not
included in the key. Odontosyllis hyalina Grube, 1878, was
reported by Hartmann-Schröder (1990); and we have
examined the specimen (HZM P-20185) and it is a small,
probably a juvenile specimen, belonging to O. australiensis;
the original description (Grube, 1878) is based on a single,
epigamic specimen (MPUW, 325), in poor condition, but
the compound chaetae are similar to those of O. australiensis
but the shafts are proportionally larger, the blades are shorter,
some of them almost unidentate. Haswell (1920) reported
two specimens of Odontosyllis suteri Benham, 1915, from
Port Jackson, Sydney, as the type description from New
Zealand specimens is incomplete and no details of the
chaetae are given it is therefore difficult to verify Haswell’s
identification, and we suggest that they may represent
individuals of Odontosyllis polycera. So neither of these
records of O. hyalina and O. suteri from Australia are likely
to be valid.
Key to Australian species of Odontosyllis
1 Occipital flap present (Fig. 32A) ................................................................................................... 2
—— Occipital flap absent or indistinct (Fig. 18A) ................................................................................ 6
2 Occipital flap totally covering prostomium and palps ............................. O. marombibooral n.sp.
—— Occipital flap covering only posterior part of prostomium........................................................... 3
3 Antennae, tentacular and dorsal cirri ovoid, globular, or sphaerical
(Fig. 26A) ............................................................................................................ O. globulocirrata
—— Antennae and cirri elongate, not as above ..................................................................................... 4
4 Dorsal cirri short, slightly longer than parapodial lobes. Blades of
compound chaetae elongated, unidentate. Parapodial lobes
elongated and distinctly bifid (Fig. 24C) .............................................................. O. freycinetensis
—— Dorsal cirri long, filiform, distinctly longer than parapodial lobes
Fig. 35A). Compound chaetae either bidentate or bidentate plus
unidentate. Parapodial lobes not elongated ................................................................................... 5
5 Large size (up to 25 mm long). Occipital flap covering almost all
prostomium, reaching to level of base of antennae (Fig. 35A).
Ventral cirri pillow-shaped ............................................................................................ O. polycera
—— Medium size (up to 6.6 mm long). Occipital flap covering only
posterior part of prostomium (Fig. 20A). Ventral cirri ovoid................................ O. australiensis
6 Dorsal cirri ovoid, shorter than parapodial lobes or similar in
length. Ventral cirri distally inserted ........................................................................... O. annulatus
—— Dorsal cirri elongated, distinctly longer than parapodial lobes.
Ventral cirri inserted basally on parapodial lobes ......................................................................... 7
7 Compound chaetae claw-shaped, unidentate or indistinctly
bidentate (Fig. 22C) ........................................................................................................ O. detecta
—— Compound chaetae with some elongated blades, some distinctly
bidentate ......................................................................................................................................... 8
8 Blades of compound chaetae mostly unidentate, and some
bidentate. Dorsum with dark transverse bands or without colour
markings .................................................................................................... O. langerhansiaesetosa
—— All blades bidentate. Dorsum with 3 longitudinal dark rows....................................... O. gravelyi

San Martín & Hutchings: Eusyllinae of Australia 283
Odontosyllis annulatus (Hartmann-Schröder, 1979)
Fig. 18A–G
Odontoautolytus annulatus Hartmann-Schröder, 1979: 112, figs
178–182.
Odontosyllis brevicirra Hartmann-Schröder, 1991: 34, figs 54–56.
Material examined. AUSTRALIA: WESTERN AUSTRALIA:
Port Hedland, 19°38'S 119°31'E, coarse sediments,
intertidal, 28 Sept. 1975, holotype of Odontoautolytus
annulatus (ZMH P-15482) and 2 paratypes (ZMH P-15483).
QUEENSLAND: Gladstone, 23°49'S 151°25'E, coarse sand,
28 Jan 1976, holotype of Odontosyllis brevicirra (HZM P-
20543) and 1 paratype (HZM P-20544). All material
collected Hartmann-Schröder.
Description. Body long, slender, (Fig. 18A), about 4.8 mm
long, 0.3 mm wide, with 47 chaetigers. Prostomium oval,
laterally convex, with 2 lateral and 2 frontal tufts of cilia
(Fig. 18A); 4 eyes in rectangular arrangement, those of
anterior pair slightly larger than posterior ones; antennae
short, digitiform, shorter than prostomium; median antenna
inserted between anterior eyes; lateral antennae inserted near
anterior margin, close to each other, near midline of margin
(Fig. 18A). Palps shorter than prostomium, blunt.
Peristomium shorter than subsequent segments; tentacular
cirri similar to antennae; dorsal ones longer than ventral
ones. Occipital flap absent, although some with peristomium
Fig. 18. Odontosyllis annulatus (Hartmann-Schröder, 1979) (A) anterior end, dorsal
view; (B) posterior end, dorsal view; (C) pharynx opening and trepan; (D) midbody
parapodium; (E) compound chaetae, midbody; (F) acicula; (G) ventral simple chaeta.
HZM P-15483. Scales: A,B 0.2 mm; C without scale, after Hartmann-Schröder (1979);
D 50 µm; E–G 10 µm.
slightly folded resembling small occipital flap. Following
segments biannulated, becoming tri- or tetra-annulated, with
transverse row of cilia usually on second annuli (Fig. 18A).
Dorsal cirri short, oval, shorter than parapodial lobes (Fig.
18A,D); dorsal cirri of chaetiger 1 longer than following
ones, similar in shape and size to dorsal tentacular cirri.
Ventral cirri fused with parapodial lobes, inserted distally
(Fig. 18A,D). Parapodia each with 4 compound chaetae (Fig.
18D), strongly heterogomph, shafts with subdistal large
tooth, and short, smooth, triangular, bidentate blades,
subdistal tooth small, all similar in shape and length (Fig.
18E), 7–8 µm in length. Solitary ventral simple chaeta on
far posterior parapodia of one specimen, slender, smooth,
unidentate (Fig. 18G). Dorsal simple chaetae not observed.
Solitary acicula on each parapodium, distally acuminate
(Fig. 18F). Pygidium large, slightly bilobed, with 2 long,
slender, filiform anal cirri (Fig. 18B). Pharynx through about
2 segments, according to original description, trepan with
6 small teeth (Fig. 18C); dark gland near pharynx opening
(Fig. 18C). Proventricle long and slender, through about
7–8 segments (Fig. 18A), with about 50 muscle cell rows.
Remarks. This species has small ventral cirri, inserted
distally. Hartmann-Schröder (1979) made an incorrect
interpretation of the parapodia, overlooked the ventral cirri
and described it as a new genus, which she placed in the
subfamily Autolytinae. Nygren (2004) synonymized
Odontoautolytus with Odontosyllis. After examination of
the type series of both species, Odontoautolytus annulatus
and Odontosyllis brevicirra, we suggest that they represent
the same species Odontosyllis annulatus.
Habitat. Occurring in coarse sediments, intertidally.
Distribution. Australia (Western Australia, Queensland).

284 Records of the Australian Museum (2006) Vol. 58
Odontosyllis australiensis Hartmann-Schröder, 1979
Figs 19C–F, 20A–H, 21A–F
Odontosyllis australiensis Hartmann-Schröder, 1979: 95, figs 97–
104; 1981: 31; 1984: 20.
?Odontosyllis hyalina.—Hartmann-Schröder, 1990: 50. Not
Grube, 1878: 129.
Odontosyllis fulgurans.—Haswell, 1920: 107. Not Audouin &
Milne Edwards, 1834: 229.
Synpalposyllis australiensis Hartmann-Schröder, 1983: 132, figs
18–20.
Fig. 19. SEM of Odontosyllis polycera Schmarda, 1863 (A) dorsal simple chaeta; (B) ventral simple chaeta. SEM of Odontosyllis
australiensis Hartmann-Schröder, 1979 (C) complete specimen, ventral view; (D) anterior end, dorsal view; (E) prostomium and
anteriormost segments; (F) midbody segments, lateral view. A,B: AM W4344; C–F: AM W28380.
Material examined. AUSTRALIA: NEW SOUTH WALES: 100 m NW
of Julian Rocks, Byron Bay, 28°36.8'S 15°37.8'E, shell and gravel, 15
m, coll. Ho & party, 3 Mar 1992, 1 (AM W28217). WESTERN A USTRALIA:
N end of Long Is., 28°27.9'S 113°46.3'E, dead coral substrate covered
in coralline & brown algae, 5.5 m, coll. C. Bryce, 22 May 1994, 2 on
SEM stub, (AM W28380); Lafontaine Is., Kimberley region, 14°10'S
125°47'E, 15m, coll P.A. Hutchings, 19 July 1988, 1 (AM W28933);
Reef south of Lucas Is., Brunswick Bay, Kimberley region, 15°16'S
124°29'E, 2 m, coll P.A. Hutchings, 24 July 1988, 1 (AM W28932).
Additional material. AUSTRALIA: WESTERN A USTRALIA. Denmark,
Ocean Beach, algae, 18 Nov. 1975, coll. & id. Hartmann-Schröder,
holotype of Synpalposyllis australiensis, (HZM P-17400). Singapore,
syntype of Odontosyllis hyalina Grube, 1878: 129, (MPUW 325).

San Martín & Hutchings: Eusyllinae of Australia 285
Description. Body about 4.4 mm long, 0.3 mm wide, with
44 chaetigers (Fig. 19C); maximum size reported 6.6 mm
long for 55 chaetigers (Hartmann-Schröder, 1981), without
colour pattern, yellowish in alcohol. Prostomium almost
round; 4 eyes in open trapezoidal arrangement, large on
epigamic specimens (Fig. 20A). Antennae originating near
anterior margin, close to each other, median antenna inserted
slightly posterior to lateral ones (Figs 19D,E, 20A),
distinctly longer than combined length of prostomium and
palps, lateral antennae shorter than median one. Palps
shorter than prostomium, ventrally folded (Fig. 21A).
Peristomium reduced dorsally, covered by occipital flap
(Fig. 19D); dorsal tentacular cirri elongated, nearly twice
as long as median antenna, ventral tentacular cirri shorter
than dorsal ones. Occipital flap distinct, small, covering
posterior margin of prostomium (Figs 19D,E, 20A, Fig. 20A
Fig. 20. Odontosyllis australiensis Hartmann-Schröder, 1979 (A)
anterior end, dorsal view; (B) midbody parapodium; (C) dorsalmost
compound chaetae, midbody parapodium; (D) remaining compound
chaetae of the same parapodium; (E) dorsal simple chaeta; (F)
ventral simple chaeta; (G) aciculae, midbody; (H) acicula, posterior
parapodium. A: AM W28396, B: AM W28380, C–F: AM W28395.
Scales: A 0.4 mm, B 48 µm, C–F 20 µm.
drawn slightly vertical). Nuchal organs ciliated, extending
laterally to prostomium (Fig. 19E). Dorsal ciliary transverse
band on each segment (Fig. 19D–F). Antennae, tentacular
cirri and dorsal cirri smooth, more or less elongated, tapered
distally, those of chaetiger 1 long, distinctly longer than
body width, those of chaetiger 2 short, on chaetigers 3 and
4, long, similar in length to body width, from chaetiger 5
onwards, alternating long and short cirri, always shorter
than body width (Figs 19D, 20A). Parapodial lobes conical,
with 2 distal, small lobes, with some cilia dorsally (Fig.
21B). Ventral cirri oval, shorter than parapodial lobes (Figs
20B, 21A). Compound chaetae heterogomph, strongly
spinose distally, also with thin spines on tendon between
shafts and blades, similar throughout, blades short, strongly
bidentate, both teeth similar in size and well separated (Figs
20C,D, 21C,D,F); blades of chaetae within fascicle
exhibiting inverse dorsoventral gradation in length (Figs
20C,D, 21B), blades about 6 µm in length dorsally, 12 µm
in length ventrally on midbody. Anterior parapodia with
about 16 compound chaetae, 12 on midbody, 4–5 on
posterior parapodia. Dorsal simple chaetae on posterior
parapodia, thin, unidentate (Fig. 20E) (minutely bifid,
according to Hartmann-Schröder, 1979), with short spines
on margin (Fig. 21E). Ventral simple chaetae on far posterior
chaetigers, bidentate, with short spines on margin (Figs 20F,
21F). Aciculae distally knobbed, ending with short terminal
process; 3–4 aciculae on anterior parapodia, and 1 on posterior
parapodia (Fig. 20G,H). Pharynx short, through 1–2 segments,
with 5 teeth and 2 lateral plates. Proventricle long and slender
(Fig. 20A), through about 8 segments, with 50 muscle cell
rows. Pygidium with 2 anal cirri similar to dorsal ones.
Remarks. This species seems to be similar to Odontosyllis
hyalina Grube, 1878 from Singapore; the description of that
species was made on the basis of a single, epigamic
specimen, in which some characters can be modified in

286 Records of the Australian Museum (2006) Vol. 58
Fig. 21. SEM of Odontosyllis australiensis Hartmann-Schröder, 1979 (A) prostomium and anteriormost segments, ventral view;
(B) posterior parapodium; (C,D): compound chaetae, midbody; (E) dorsal simple chaeta; (F) ventral simple and compound chaetae.
AM W28380.
comparison to non-epigamic specimens, and it is difficult
to assess if both species are valid or synonymous. The
syntype (MPUW, 325) is in poor condition, covered by
crystals of formalin, but the chaetae appear different (see
above); so at this stage we are accepting both species as
valid.
Habitat. Occurring on algae, dead coral and coarse
sediments, from intertidal to shallow depths.
Distribution. Australia (Western Australia, New South
Wales, Queensland).
Odontosyllis detecta Augener, 1913
Fig. 22A–C
Odontosyllis detecta Augener, 1913: 236, pl. III, Fig. 33, text-
fig. 34; 1927: 153.—Haswell, 1920: 105.—Imajima, 1966:
103, figs 33a–m.—Hartmann-Schröder, 1985: 69.—San
Martín, 1990: 613, fig. 16.
Material examined. AUSTRALIA: NEW SOUTH WALES: Port
Jackson, 33°51'S 151°16'E, donated by W.A. Haswell, Feb
1920, 1 (slide) (AM W501).

San Martín & Hutchings: Eusyllinae of Australia 287
Description. Only examined specimen, mature, epigamic
male, 5 mm long, 0.1 mm wide, with 37 chaetigers, on
permanent, stained slide. Adult, non-epigamic specimens
from Japan, 6–9 mm long, 1 mm wide, with 40–52
chaetigers (Imajima, 1966). Specimen from Port Jackson
with 2 large, black eyes, small palps and coiled cirri (Fig.
22A). Occipital flap absent. 4–5 compound chaetae per
parapodium, with distally spinose shafts and short, curved,
unidentate blades, with short spines on margin (Fig. 22C).
Dorsal simple chaetae from midbody, thin, unidentate,
smooth. Ventral simple chaetae absent. Acicula solitary,
distally knobbed, with short tip. Pharynx through 2
segments, with 5 teeth and 2 lateral plates. Proventricle
through 2 segments (Fig. 22A). Pygidium semi-circular,
with 2 long anal cirri, extending for 6 chaetigers (Fig. 22B).
Japanese non epigamic specimens (fide Imajima, 1966),
with antennae, dorsum and dorsal cirri dark red; with ciliary
bands across dorsum, nuchal ridges present; lacking
occipital flap. Dorsal cirri alternating in length, shorter than
body width, and ventral simple chaetae on posterior
parapodia, unidentate, slightly hooked, with short, fine
spines on margin.
Remarks. As the only Australian material available for
examination was a mature epigamic male, it is difficult to
relate this to the non epigamic individuals recorded from
Japan by Imajima (1966), but we believe them to represent
the same species.
Habitat. Occurring in sand, seagrass, algae; intertidally.
Distribution. Australia (Western Australia, South Australia,
New South Wales), Japan, Cuba.
Fig. 22. Odontosyllis detecta Augener, 1913 (A) anterior end, dorsal view
(epigamic specimen); (B) posterior end, dorsal view; (C) compound chaetae,
midbody. AM W501. Scales: A,B 0.4 mm, C 20 µm.
Odontosyllis freycinetensis Augener, 1913
Figs 23C–F, 24A–G, 25A–D
Odontosyllis freycinetensis Augener, 1913: 234, pl. II, fig. 7, text-
fig. 33 a, b.—Haswell, 1920: 107.
Material examined. AUSTRALIA: NEW SOUTH WALES: Manta Reef,
North West Solitary Is. 30°1.5'S 153°16.5'E, lace bryozoan, 19 m, coll.
R.T. Springthorpe, 25 Jun 1992, 1 (AM W28216); S side of Shelly Beach,
Fairy Bower, 33°48.18'S 151°17.4'E, in between large boulders & under
small ones, 8 m, coll. P.A Hutchings, 24 Oct 1971, 1 on SEM stub (AM
W24679); Grotto Point, Port Jackson, 33°49'S 151°15'E, algae, 4 m,
coll. P. Colman, 18 July 1983, 1 (AM W28915); Summer Cloud Bay,
Wreck Bay, 35°10.5'S 150°41'E, 15 m, coll. P.A. Hutchings, 29 Nov
1971, 1 (AM W26325). WESTERN AUSTRALIA: Off S end of Long Is.
Beacon Is. 28°28.8'S 113°46.3'E, dead coral substrate covered in
coralline algae, 4.5 m, coll. P.A. Hutchings, 25 May 1994, 2 (AM
W28390).
Description. Body broad anteriorly, tapered posteriorly
(Fig. 23C), largest complete specimen examined 10 mm
long, 0.5 mm wide, with 59 chaetigers, sometimes without
colour pattern, but usually with 2–3 dark, dorsal spots on
some anterior chaetigers (Fig. 24A), others dark.
Prostomium oval, large, with long cilia laterally (Fig. 24A);
4 eyes in open trapezoidal arrangement; antennae short,
fusiform, shorter than prostomium (Figs 23D, 24A), median
antenna originating between anterior eyes, lateral antennae
near anterior margin of prostomium, close to median
antennae. Palps small, shorter than prostomium, ventrally
folded (Fig. 24A). Nuchal organs distinct, with long cilia
between prostomium and peristomium (Figs 23D, 24A).
Peristomium distinct, similar in length to subsequent
segments. Occipital flap covering posterior part of
prostomium (Figs 23D, 24A). Tentacular cirri similar in
shape to antennae, but longer. Dorsal cirri fusiform, some
more elongated than antennae, those of chaetiger 1 slightly
longer than others, shorter than half of body width, slightly
longer than parapodial lobes (Figs 23E, 24C). Parapodial
lobes elongated, almost rectangular, distally bilobed (Figs
23E, 24C); anterior parapodia with distal, digitiform papilla;
progressively, posteriorly parapodia becoming more
elongated, and distinctly bilobed distally, and distal papilla
becoming fused. Ventral cirri digitiform, elongated, reaching
or extending beyond parapodial lobes (Fig. 24C).
Compound chaetae slender, with elongated, unidentate
blades, with slightly hooked tips (Figs 23F, 24E,F, 25A),

288 Records of the Australian Museum (2006) Vol. 58
Fig. 23. SEM of Odontosyllis marombibooral n.sp. (A) dorsal compound chaeta, posterior parapodium; (B) ventral compound
chaeta, posterior parapodium. SEM of Odontosyllis freycinetensis Augener, 1913 (C) complete specimen, dorsal view; (D) prostomium
and anteriormost segments; (E) midbody parapodia, dorsal view; (F) anterior compound chaetae. A,B: AM W5496, C–F: AM
W24679.
with margins either smooth or with short spines. Anterior
parapodia with about 16–18 compound chaetae, with
dorsoventral gradation in length of blades within fascicle, 40
µm in length dorsally, 20 µm in length ventrally; number of
compound chaetae per parapodium decreasing posteriorly, up
to about 9 on posterior parapodia, blades of similar lengths
to anterior ones; blades with short, minute subdistal spine
(Fig. 25B–D, arrows). Dorsal and ventral simple chaetae
on posterior parapodia, thin, smooth, dorsal ones bifid (Fig.
24H) and ventral unidentate (Fig. 24I). Anterior parapodia
with 2–3 aciculae, becoming solitary after proventricular
segments, distally acuminate (Fig. 24D,G). Pharynx wide,
through 2–3 segments, with 5 teeth and 2 lateral plates.
Proventricle wide, short, slightly longer than pharynx (Fig.
24A), through about 4 segments, with 28–30 muscle cell
rows. Pygidium rectangular, small, with 2 lateral, short anal
cirri, and 1 median, digitiform papilla (Fig. 24B).
Habitat. Occurring on algae, bryozoans, dead corals; inter-
tidal and shallow depths.
Distribution. Australia (Western Australia, New South
Wales).

San Martín & Hutchings: Eusyllinae of Australia 289
Fig. 24. Odontosyllis freycinetensis Augener, 1913 (A) anterior end, dorsal view;
(B) posterior end, dorsal view; (C) posterior parapodium; (D) acicula, anterior
parapodium; (E) compound chaetae, anterior parapodium; (F) compound chaetae,
posterior parapodium; (G) acicula, posterior parapodium; (H) dorsal simple
chaeta; (I) ventral simple chaeta. AM W28216. Scales: A,B 0.2 mm, C 48 µm,
D–I 20 µm.
Odontosyllis globulocirrata
Hartmann-Schröder, 1981
Fig. 26A–E
Odontosyllis globulocirrata Hartmann-Schröder, 1981: 31, figs
49–51; 1983: 129; 1989: 25.
Material examined. AUSTRALIA: WESTERN A USTRALIA: Cervantes,
30°30'S 115°03'E, fine sand with Posidonia, coll. G. Hartmann-Schröder,
24 Oct 1972, 2 paratypes (AM W17725). NEW SOUTH WALES: Halfway
Reef, 200 m S of Sullivan Reef, Ulladulla, 35°21'25"S 150°29'19"E,
airlift over wall of sponges, Bryozoa, Hydrozoa, 15m, coll. P.B. Berents,
K.B. Attwood & A. Murray, 3 May 1997, 1 (AM W29375).
Description. Paratype examined 8.1 mm long, 0.8 mm wide,
with 43 chaetigers; holotype 10 mm long for 48 chaetigers
according to Hartmann-Schröder. Body broad anteriorly,
tapering posteriorly, lightly coloured on dorsum of some
segments, with 2–4 dark spots (Fig. 26A). Prostomium oval,
about 3 times wider than long; 4 eyes arranged in open
trapezoidal pattern, posterior ones covered by occipital flap
(Fig. 26A); antennae short, globular, median antenna slightly
longer than lateral ones (Fig. 26A,B), originating between
anterior eyes, lateral antennae inserted on anterior margin
of prostomium. Palps small, short. Peristomium dorsally
reduced (Fig. 26A,B). Tentacular cirri globular, similar in
size to antennae. Occipital flap oval, covering only posterior
margin of prostomium (Fig. 26A,B). Dorsal cirri globular,
sphaerical, shorter than parapodial lobes, those inserted
dorsolaterally on chaetigers 1, 4, and 6-8-10-…, slightly
larger and more elongate than those on chaetigers 2, 3, 5-7-

290 Records of the Australian Museum (2006) Vol. 58
Fig. 25. SEM of Odontosyllis freycinetensis Augener, 1913 (A,B), compound chaetae, midbody; (C) long-bladed compound chaeta,
midbody; (D) short bladed compound chaeta, midbody. SEM of Odontosyllis polycera Schmarda, 1863 (E) anterior end and
midbody, dorsal view; (F) anterior end, dorsal view. A–D: AM W24679, E: AM W195387, F: AM W194258.
9-…, inserted adjacent to parapodial lobes (Fig. 26B,C).
Parapodial lobes elongated, distinctly bilobed distally (Fig.
26A–C). Ventral cirri short, conical, inserted near distal part
of parapodial lobes. Compound chaetae numerous, about
20 on midbody parapodia, with smooth shafts, or provided
with minute subdistal spines, and elongate, unidentate
blades, distally slightly hooked, provided with short, fine
spines on margin (Fig. 26E), blades of chaetae within
fascicle exhibiting dorsoventral gradation in length, about
33 µm in length dorsally, 25 µm in length ventrally. Dorsal
and ventral simple chaetae not seen nor previously
described. Aciculae solitary, slender, straight. Pharynx wide,
short, with 7 teeth (fide Hartmann-Schröder; not seen in
material examined) and 2 lateral plates. Proventricle large,
2.5× longer than pharynx (Fig. 26D), with about 40 muscle
cell rows. Pygidium with 2 globular anal cirri, similar to
dorsal ones.
Habitat. Occurring on algae and sediment, intertidal.
Distribution. Australia (Western Australia, New South
Wales).

San Martín & Hutchings: Eusyllinae of Australia 291
Fig. 26. Odontosyllis globulocirrata Hartmann-Schröder, 1981 (A) anterior end, dorsal view; (B) anterior end,
dorsolateral view; (C) midbody parapodia; (D) pharynx and proventricle; (E) compound chaetae, midbody. AM
W17725. Scales: A,B 0.4 mm, C 0.1 mm, D 0.4 mm, E 20 µm.

292 Records of the Australian Museum (2006) Vol. 58
Odontosyllis gravelyi Fauvel, 1930
Figs 27C–F, 28A–H, 29A–F
Odontosyllis gravelyi Fauvel, 1930: 16, figs. 3–4; 1953: 160, figs.
81–82.
Material examined. AUSTRALIA: NEW SOUTH WALES: Careel Bay,
Pittwater, 33°37'S 151°19'E, Zostera, coll. P.A. Hutchings, 4 Nov 1973
(AM W11110). QUEENSLAND: Calliope R., 23°51'S 151°10'E, coll. P.
Saenger, 1974, 7 (AM W199367). Triangular Islets, Shoalwater Bay,
22°23'S 150°31'E, coll. J.A. Lewis & J.R. Forsyth, 1981, 10 on SEM
stub (AM W202645).
Fig. 27. SEM of Odontosyllis langerhansiaesetosa Hartmann-Schröder, 1979 (A) long falciger, midbody; (B) short falciger, midbody.
SEM of Odontosyllis gravelyi n.sp. (C) anterior end, dorsal view; (D) detail of prostomium (arrows showing the mounds with
minute pores); (E) detail of mounds with pores on prostomium; (F) midbody parapodia. A,B: AM W198069, W202639; C–F: AM
W202645.
Description. Body 14 mm long, 1.2 mm wide, with 50
chaetigers; distinct, median longitudinal black band, and
large lateral spots, resembling 3 longitudinal stripes (Fig.
28A). Prostomium small, hidden under most anterior
segments (Figs 27C, 28A–C) (probably contracted), with 2
black lateral spots and posterior transverse band; antennae
short and slender, inserted in front of anterior eyes, median
antenna longer than lateral antennae, inserted slightly
posteriorly (Figs 27C, 28B,C), similar to combined length
of prostomium and palps. Prostomium with 2 slightly raised
mounds with minute pores (see arrows Fig. 27D,E). Two
densely ciliated, semi-circular nuchal organs (Fig. 27C,D).
Palps small, fused basally, ventrally folded. Peristomium

San Martín & Hutchings: Eusyllinae of Australia 293
Fig. 28. Odontosyllis gravelyi Fauvel, 1930 (A) anterior end, dorsal view; (B)
dorsal detail of prostomium; (C) frontal view of prostomium; (D) midbody
parapodium of an epigamic female; (E) pharynx and proventricle; (F) trepan;
(G) long-bladed compound chaetae, midbody; (H) falcigers, midbody. Holotype
AM W11110, except B, AM W202645. Scales: A–C,E: 0.8 mm, D: 0.2 mm, F:
0.18 mm, G,H: 20 µm.
small, dorsally nearly covered by small lobe of chaetiger 1
(Figs 27C, 28B); tentacular cirri similar to antennae, but
slightly longer. Anterior dorsal cirri elongated, smooth,
distally tapered, rugose to pseudoarticulate (Figs 27C, 28A)
becoming progressively shorter along body (Figs 27F, 28D),
slightly longer than parapodial lobes. Parapodia with 2–3
parallel lobes (Figs 27F, 28D, 29A). Ventral cirri short,
pillow-shaped (Figs 27F, 28D). Compound chaetae
numerous, shafts distally spinose, and blades of 2 kinds,
most dorsal ones with elongate, bidentate blades (Fig. 28G),
short spines on margin, and remaining chaetae with short,
triangular, distinctly bidentate blades (Figs 28H, 29A–E);
on midbody, parapodia with about 6–7 compound chaetae
of slender type, 37–45 µm in length, and 28–30 chaetae
with short blades, all similar, about 16–18 µm in length;
compound chaetae slightly longer on anterior parapodia
(Fig. 29A,B). Aciculae slender, distally blunt, numerous in
anterior parapodia, reduced to 2–3 on posterior parapodia.
Dorsal simple chaetae slender, thin, unidentate (Fig. 29C).
Ventral simple chaetae on posterior parapodia, bidentate,
with short spines on margin (Fig. 29C,F). Pharynx through
3–4 segments, with 5 teeth and 2 lateral plates (Fig. 28F).
Proventricle long, slender (Fig. 28E), through 9 segments,
with about 80 muscle cell rows.

294 Records of the Australian Museum (2006) Vol. 58
Fig. 29. SEM of Odontosyllis gravelyi Fauvel, 1930 (A) chaetal fascicle, anterior parapodium; (B) chaetal fascicles, midbody
parapodia; (C) chaetal fascicle, posterior parapodium; (D) compound chaetae, posterior parapodium; (E) falciger, midbody; (F)
ventral simple chaeta. AM W11110.
Remarks. The small lobe on the peristomium is not regarded
as an occipital flap. This species is characterized by its
colour pattern, a lack of an occipital flap, and two kinds of
compound chaetae being present. Odontosyllis trilineata
Imajima, 2003 has a similar colour pattern and chaetae, but
they differ in that O. gravelyi also has long-bladed chaetae,
and O. trilineata has a distinct occipital flap and 9 teeth on
trepan (Imajima, 2003).
Habitat. Occurring in coarse sand and gravel and mud,
shallow depths often in estuarine conditions.
Distribution. Australia (New South Wales, Queensland) and
southeast India.
Odontosyllis langerhansiaesetosa
Hartmann-Schröder, 1979
Figs 27A,B, 30A–K, 31A–F
Odontosyllis langerhansiaesetosa Hartmann-Schröder, 1979: 94,
figs 91–96; 1980: 50; 1990: 51.
Material examined. AUSTRALIA: QUEENSLAND: Triangular Islets,
Shoalwater Bay, 22°23'S 150°31'E, coll. J.A. Lewis & J.R. Forsyth, 1981,
5 + 1 on SEM stub (AM W202639); Calliope R., 23°51'S 151°10'E,
coll. P. Saenger, 1974, few (AM W28947); Auckland Creek, Gladstone,
23°51'S 151°14'E, coarse gravel, 1.3 m, coll. P. Saenger, Apr 1977, 1
(AM W198068); Auckland Creek, Gladstone, 23°51'S 151°14'E, coarse
sand, 2.8 m, coll. P. Saenger, July 1979, 1 on SEM stub (AM W198069);
Branch of Calliope R., Gladstone, 23°51'S 151°10'E, coarse gravel, 2.3

San Martín & Hutchings: Eusyllinae of Australia 295
Fig. 30. Odontosyllis langerhansiaesetosa Hartmann-Schröder, 1979 (A) anterior end, dorsal view; (B) midbody parapodium; (C)
pharynx and proventricle; (D) trepan; (E) long-bladed compound chaetae, anterior parapodium; (F) falcigers, anterior parapodium; (G)
long-bladed compound chaetae, posterior parapodium; (H) falcigers, posterior parapodium; (I) dorsal simple chaeta; (J) acicula; (K)
ventral simple chaeta. A,B,E–K: AM W198073; C: AM W198068. Scales: A 0.4 mm, B 0.18 mm, C 0.1 mm, D 0.2 mm, E–K 20 µm.
m, P. Saenger, July 1979, 1 (AM W198070); Calliope R., Gladstone,
23°51'S 151°10'E, silty sand, 3.4 m, P. Saenger, July 1979, 1 (AM
W198071); Auckland Creek, Gladstone, 23°51'S 151°14'E, coarse sand,
2.3 m, coll. P. Saenger, July 1979, 1 (AM W198072); Black Harry Creek,
Gladstone, 23°51'S 151°10'E, soft mud, 0.7 m, coll. P. Saenger, Sept.
1981, 1 (AM W198073). NEW SOUTH WALES: Green Point, Hawkesbury
R., 33°34'S 151°14'E, mud, 12 m, coll. A. Jones & party, 13 Nov 1979,
1 (AM W28946).
Description. Body broad anteriorly, tapered posteriorly, up
to 9.6 mm long, 0.9 mm wide, 72 chaetigers; colour pattern
variable, some apparently lacking pigmentation to others,
dark, with distinct pattern (Fig. 30A) of dark band on dorsum
of each anterior segment, split into 3 or more spots, and
sometimes on midbody segments. Prostomium oval, about
twice as wide as long; 4 eyes in trapezoidal arrangement.
Median antenna longer than combined length of prostomium
and palps, originating on middle of prostomium; lateral
antennae about half of length of median one, inserted on
anterior margin of prostomium (Figs 30A, 31A). Palps short,
ventrally folded (Fig. 31C). Peristomium distinct dorsally,
slightly shorter than subsequent segments (Figs 30A, 31A),
with dorsal, small lobe covering posterior margin of
prostomium (Fig. 30A). Occipital flap absent. Tentacular
cirri and most anterior dorsal cirri elongated, longer than
body width (Figs 30A, 31A), smooth; remaining dorsal cirri
shorter (Figs 30A, 31B), slightly longer than parapodial
lobes, distally tapered (Figs 30B, 31D). Parapodial lobes

296 Records of the Australian Museum (2006) Vol. 58
Fig. 31. SEM of Odontosyllis langerhansiaesetosa Hartmann-Schröder, 1979 (A) anterior end, dorsal view; (B) midbody parapodia
of an epigamic specimen, showing the natatory chaetae; (C) anterior end, ventral view; (D) cross section of a midbody segment;
(E,F) compound chaetae, anterior parapodium. AM W198069, W202639.
conical; ventral cirri triangular to digitiform, shorter than
parapodial lobes. Compound chaetae with spinose shafts
and 2 kinds of blades; most superior chaetae with elongated,
slender, distally bidentate blades (Figs 30E,G, 31E,F),
margin smooth, and others similar but blades shorter,
unidentate or weakly bidentate (Figs 30F,H, 27A,B); tendon
partially fused to blades. Anterior parapodia with about 6
long-bladed chaetae, 38–60 µm long, and 10 short-bladed
chaetae, with dorsoventral gradation in length of blades,
35 µm in length dorsally, 18 µm in length ventrally.
Progressively along body, number of compound chaetae
per parapodium decreasing to 4 long-bladed, 37–40 µm in
length, and 4–5 short bladed, 25 µm in length dorsally, 15
µm in length ventrally. Dorsal simple chaetae on posterior
parapodia, slender, unidentate, distally blunt, with some
short, subdistal spines on margin (Fig. 30I). Ventral simple
chaetae on most posterior parapodia, bidentate, with fine
spines on margin (Fig. 30K). Several aciculae on anterior
parapodia, decreasing to 1 on posterior parapodia, slender,
distally knobbed (Fig. 30J). Pharynx about half of length
of proventricle (Fig. 30C), through about 6 segments, with
5 teeth and 2 lateral plates (Fig. 30D). Proventricle through
about 9–10 segments, with 50 muscle cell rows.
Habitat. Occurring in coarse sand, silt, gravel, mud, from
intertidal to shallow depths.
Distribution. Australia (Western Australia, Queensland,
New South Wales).

San Martín & Hutchings: Eusyllinae of Australia 297
Odontosyllis marombibooral n.sp.
Figs 23A,B, 32A–C, 33A–F, 34A–E
Material examined. HOLOTYPE (AM W28387) AUSTRALIA:
WESTERN AUSTRALIA: Off jetty adjacent to Fisheries Hut, Beacon Is.
28°25.5'S 113°47'E, dead coral substrate, plate-like Acropora and
Montipora spp., 12 m, coll. P.A. Hutchings, 23 May 1994. PARATYPES:
1 on SEM stub (AM W5496) Blow Holes, Quobba, 24°29'S 113°25'E,
in sponge, 2 m, coll. N. Coleman, 20 Jun 1972; E side of Mangrove Is.
20°56'S 116°09'E, in dead coral, 1 m, coll. Aquinas College, 9 Jan 1968,
1 (AM W194813).
Description. Body broad, robust anteriorly, tapered
posteriorly, 25 mm long, 2 mm wide, with 93 chaetigers.
Distinctly coloured (Fig. 32A) with transverse black rows
and other yellowish areas on preserved specimens; 2 kidney-
shaped black spots on prostomium (Fig. 32B); chaetigers
1, 4, 6, 9 darkly pigmented, subsequent segments, consisting
of one darkly pigmented segment followed by unpigmented
segment, this striped pattern continues to mid body,
following segments with reduced pigmentation; each
segment slightly biannulate with dorsum rugose. Pro-
stomium almost circular, totally covered by occipital flap
(Figs 32A, 34A,B), with 4 eyes in rectangular pattern,
antennae short and thick, shorter than prostomium,
indistinctly wrinkled (Fig. 32B,C), originating close to each
other, all similar size, median antenna originating slightly
posteriorly to lateral antennae, all antennae covered by
occipital flap. Palps triangular, ventrally folded (Fig. 32C).
Peristomium dorsally reduced, covered by chaetiger 1 and
occipital flap (Fig. 32A), forming 2 lobes ventrally (Fig.
32C). Occipital flap large (Figs 32A–C, 34A,B) colourless.
Tentacular and dorsal cirri similar, short, thick, slightly
Fig. 32. Odontosyllis marombibooral n.sp. (A)
anterior end, dorsal view; (B) detail of prostomium,
with the occipital flap turned backwards; (C)
prostomium and anteriormost segments, ventral
view. AM W28387. Scales: 0.4 mm
longer than parapodial lobes, indistinctly articulated (Figs
32A–C, 34A–D). Parapodia elongate, slender, with 2 distal
lobes (Figs 33A, 34A–C). Ventral cirri short, broad,
subdistally inserted (Figs 32A, 33A, 34C,D). Compound
chaetae smooth or with short spines on shafts; within each
parapodium, few compound chaetae, most dorsally located,
with elongate, slender blades, distally bidentate and short
spines on margin; remaining chaetae with shorter and wider
blades that progressively along body, become strongly
bidentate, with short spines on margin (Figs 33B,D, 34E,
23A,B). Anterior compound chaetae shorter and more
slender than posterior ones, about 20 per parapodium, with
dorsoventral gradation in length of blades within fascicle,
38 µm in length dorsally, 16 µm in length ventrally, those
of posterior parapodia, numbering about 15 per parapodium,
46 µm in length dorsally, 26 µm in length ventrally. Dorsal
and ventral simple chaetae not seen. Aciculae slender,
distally broad, numerous on anterior parapodium, numbers
decreasing posteriorly to 2–3 on posterior parapodia (Fig.
33C). Pharynx about half of length of proventricle (Fig.
33E), with 5 teeth and 2 lateral plates (Fig. 33F). Proventricle
long and wide, with about 56 muscle cell rows (Fig. 33E).
Remarks. Odontosyllis marombibooral n.sp. differs from
all other species of the genus in having a large occipital
flap, which covers totally the prostomium and antennae, a
distinctive colour pattern, bifid parapodia with distal ventral
cirri, and short, pseudoarticulated dorsal cirri. No other
species has this combination of characters. The most similar
species is Odontosyllis picta (Kinberg, 1865 described as
Eurymedusa picta), from New Zealand (Ehlers, 1904) (also
questionably reported for Australia), which also has a large
occipital flap, but not as large as the one present in
Odontosyllis marombibooral n.sp. Odontosyllis picta also
has black transverse stripes, but they are arranged in a
different pattern to those of Odontosyllis marombibooral,
and the compound chaetae have all short blades, whereas
Odontosyllis marombibooral, has two types of chaetae
present. Odontosyllis rubrofasciata (Pruvot, 1930), from

298 Records of the Australian Museum (2006) Vol. 58
Fig. 33. Odontosyllis marombibooral n.sp. (A) midbody
parapodial lobe; (B) compound chaetae, anterior segment;
(C) aciculae, posterior parapodium; (D) compound chaetae,
posterior parapodium; (E) pharynx and proventricle; (F)
pharyngeal armature. AM W28387. Scales: A 0.18 mm, B–
D 20 µm, E 0.8 mm, F 0.4 mm.
New Caledonia, (described as Atelesyllis rubrofasciata), and
O. rubrofasciata Grube, 1878 (a possible homonym), have
strikingly similar colour patterns, short dorsal cirri, and
compound chaetae that resemble those present in Odontosyllis
marombibooral, but the occipital flap is much smaller and this
character easily separates these three species. Fauvel, in the
notes given after the description of Atelesyllis rubrofasciata
considered that is a different species to O. rubrofasciata Grube,
1879. Odontosyllis rubrofasciata (Pruvot, 1930) has transverse
stripes, but only from the midbody onwards, the prostomium
has 4 lobes. The chaetae, although similar to those of O.
marombibooral are slightly different, with long blades not as
elongated and short blades with distal tooth not as curved as
those present in O. marombibooral. In the description of
Atelesyllis rubrofasciata (Pruvot, 1930), pharyngeal teeth are
not mentioned as being present, although examining a range
of species of Odontosyllis some individuals appear to lack these
teeth. In all other characters, however, they are identical to
specimens with teeth, suggesting they may lose these teeth
and be able to regenerate them.
Habitat. Occurring in dead corals and sponges, from depths
of 1–12 m.
Distribution. Australia (Western Australia).
Etymology. The name of this species comes from two
aboriginal words, marombi, meaning shield, and booral,
meaning big, or large, in reference to the large occipital
flap present.
Odontosyllis polycera (Schmarda, 1861)
Figs 19A,B, 25E,F, 35A–F, 36A–F
Syllis polycera Schmarda, 1861: 72, pl. 28, fig. 219
Odontosyllis polycera Augener, 1927: 152.—Day, 1967: 260, fig.
12.—Hutchings & Murray, 1984: 32.—Hartmann-Schröder,
1984: 20; 1985: 68, figs 14–17; 1986: 41; 1989: 25; 1990: 51.
?Odontosyllis suteri Non Benham, 1915.—Haswell, 1920: 107.
Material examined. AUSTRALIA: QUEENSLAND: Lagoon, Low Islets,
16°23'S 145°34'E, British Great Barrier Reef Expedition 1928–1929, 3
Oct 1928, id. as O. hyalina, 1 epigamic specimen, (AM W2952). NEW
SOUTH WALES: NE of Mary’s Rock, Cook Is., 28°11.42'S 153°34.79'E,
orange frilly bryozoan, 19 m, coll. R.T. Springthorpe, 8 Jun 1993, 1
(AM W28400); NW of Split Solitary Is., 30°14'S 153°10.8'E, orange
sponge, 14 m, coll. R.T. Springthorpe, 7 Mar 1992, 1 (AM W28219);
Green Point, Hawkesbury R., 33°34'S 151°14'E, mud, 12 m, coll. A.
Jones & party, 13 Nov 1979, 1 (AM W196605); Hawkesbury R., E end
of Brooklyn Boat Channel, 33°33'S 151°14'E. A. Jones & party, 18 Dec
1979, 1 (AM W196420); Grotto Point, Port Jackson, 33°49'S 151°15'E,
algae, 4 m, coll. P. Colman, 18 July 1983, 2 (AM W28407); W of La
Perouse, Botany Bay, 33°59.4'S 151°12.8'E, St. 99, mud, 13 m, coll.
SPCC, 10 Mar 1977, P.A. Hutchings (id.), 1 epigamic female, (+ 2
midbody pieces on SEM stub) (AM W14203); S of Banksmeadow,
Botany Bay, 33°58'S 151°12'E, mud, 19 m, 8 Dec 1976, coll. SPPC, 1
(AM W14197); Botany Bay, 33°59.3'S 151°13.1'E, coll. NSW Fisheries,
31 Jan 1975, 1 epigamic specimen on SEM stub (AM W195387); Botany
Bay, 34°0.5'S 151°11'E, coll. NSW Fisheries, 1 (AM W195520); N of
Kurnell, Botany Bay, 34°00'S 151°12'E, mud, 13 m, 10 March 1977,
P.A. Hutchings (id.), 1 (AM W14204); Port Botany, Botany Bay,
33°58.75'S 151°11.093'E, 7 m, 7 Apr 1992, A. Murray (id.), 1 (AM
W21628); W of La Perouse, Botany Bay, muddy sand, 19 m, 4 Feb
1977, coll. SPCC, 2 (AM W14201); off Bass Point, 34°36'S 150°54'E,
50 m, coll. The Ecology Lab, 1 Feb 1990, several (AM W22990); 100
m, Jervis Bay, 35°06'S 150°44'E, coll. P.A. Hutchings & party, Feb 1989,
1 (AM W20828); Jervis Bay, off Murrays Beach, 35°7.5'S 150°45.5'E,
coll. NSW Fisheries, 25 Apr 1972, 1 on SEM stub (AM W194258);
Jervis Bay, Murrays Basin sandbank, coll. NSW Fisheries, 17 Oct 1972,
1 (AM W194540); Jervis Bay, Murrays Basin 35°7.5'S 150°45.5E, sand,
coll. NSW Fisheries, 17 Oct 1972, 1 (AM W194290); Jervis Bay, off
Murrays Beach, 35°7.5'S 150°45.5'E, NSW Fisheries, Apr 1972, 1 (AM
W17559); Plantation Point, Jervis Bay, 35°4.35'S 150°41.80'E, intertidal
rock platform, coll. A. Murray, 24 Oct 1998, 1 (AM W24937). TASMANIA:
Fancy Point, Bruny Is. 43°16'S 147°19'E, algae, 3–6 m, coll. G. Edgar,
10 Nov 1980, 1 (AM W18189). SOUTH AUSTRALIA: 2 km off First Creek,
Spencer Gulf, Port Pirie, 33°12'S 138°00'E, subtidal, Posidonia and
Amphibolus spp seagrass, 4.1 m, T.J. Ward, Mar 1980, 1 (AM W28233);
Sleaford Bay, Port Lincoln, 34°54'S 135°47'E, algal washings, coll. P.A.
Hutchings, 10 Mar 1979, 4 (AM W26356). WESTERN AUSTRALIA:
Bramble Point, Princess Royal Harbour, 35°02'S 117°55'E, Posidonia
sinuosa, 1–1.5m, coll. P.A. Hutchings & party, Jan 1988, 1 (AM
W20305); Cottlesloe Beach, 6 miles W of Perth, calcareous algae &
Idanthyrsus tubes, 6 m, coll. H. Paxton, 14 Feb 1970, G. Hartmann-
Schröder (id.), 6 (+ 2 fragments on SEM stub), (AM W4344); Red Bluff,
Kalbarri, 27°42'S 114°09'E, mixed coralline algae on rocky shore, 3.5
m, coll. J.K. Lowry, 10 Jan., 1984, 2 (AM W28365).

San Martín & Hutchings: Eusyllinae of Australia 299
Fig. 34. SEM of Odontosyllis marombibooral n.sp. (A) anterior
end, lateral view; (B) detail of prostomium and anteriormost
segments, lateral view; (C) midbody parapodium; (D) transverse
section, posterior end; (E) compound chaetae, anterior
parapodium. AM W5496.
Description. Body robust (Fig. 35E), broad anteriorly, with
numerous segments, one specimen (AM W22990) nearly
complete, 25 mm long, 2 mm wide, with 116 chaetigers,
but specimens twice as long previously reported, pale yellow
in alcohol, without colour pattern. Prostomium oval, with
2 pairs of large eyes in open trapezoidal arrangement;
median antenna longer than combined length of prostomium
and palps, lateral antennae inserted close to median antenna,
on anterior margin of prostomium, nearly half length of
median one (Figs 25F, 35A). Palps divergent, ventrally
folded, free for almost their length, fused basally. Dorsal
tentacular cirri slightly longer than antennae, ventral
tentacular cirri shorter. Peristomium reduced dorsally,
covered by large, long occipital flap, that also covers most
of prostomium (Figs 25E,F, 35A); anterior edge of occipital
flap with band of cilia (Fig. 36A, arrow); midbody segments
divided in two sections by one furrow, each section with
row of cilia (Fig. 36B, arrows). Antennae, tentacular, and
dorsal cirri elongated, smooth, distally tapered, with short
cirrophore (Figs 25E, 35A), dorsal cirri becoming shorter
posteriorly (Fig. 25E); dorsal cirri of chaetigers 1, 3, 4 and
6 long, alternating dorsal cirri long and short on remaining
segments. Parapodia conical, ending in 2 distal lobes (Fig.
35B). Ventral cirri rounded, stout, pillow-shaped, shorter
than parapodial lobes (Figs 35B, 36C,D). Compound
chaetae heterogomph falcigers, with spinose ending shafts
and short, bidentate blades, slightly hooked, proximal tooth
well separated from distal ones (Figs 35E,F, 36F),
sometimes on middle of margin; few, short spines, on
margin, also some thin spines on tendons between shafts
and blades, becoming more marked on posterior chaetae
(Fig. 35E,F). Length of blades of chaetae within fascicle
increasing ventrally (Figs 35E,F, 36E). Anterior parapodia
with about 50 compound chaetae, blades 15–17 µm in length
dorsally, 23 µm in length ventrally; number of compound
chaetae per parapodium diminishing progressively along
body to 25–27 on posterior segments, blades increasing in
length within fascicle from 18 µm dorsally to 25 µm
ventrally. Anterior parapodia with 3 slender aciculae with
trilobed tips, numbers decreasing posteriorly with only 1
on posterior parapodia, similar to anterior ones (Fig. 35D),
but larger. Dorsal simple chaetae on posterior parapodia,
thin, unidentate, with short spines on margin (Fig. 19A).

300 Records of the Australian Museum (2006) Vol. 58
Fig. 35. Odontosyllis polycera Schmarda, 1863 (A) anterior end, dorsal view;
(B) midbody parapodium; (C) pharynx and proventricle; (D) acicula, posterior
parapodium; (E) compound chaetae, anterior parapodium; (F) compound chaetae,
posterior parapodium. AM W22990. Scales: A 0.8 mm, B 0.4 mm, C 0.5 mm,
D–F 20 µm.
Ventral simple chaetae on far posterior segments of few
specimens, with minute subdistal spines on margin, distally
hooked, with short proximal tooth (Fig. 19B). Pharynx short,
through about 4 segments, with 5–8 teeth, and 2 lateral
plates. Proventricle long, relatively slender, more than 3
times length of pharynx (Fig. 35C), through 9–10 segments,
with about 130 muscle cell rows. Pygidium, with 2 long
anal cirri. Several specimens epitokous, with long natatory
chaetae on midbody parapodia (Fig. 36D).
Remarks. This species, which was described from Table
Bay, South Africa, has been reported widely, and a detailed
study should be undertaken to confirm such a wide
distribution and depth range.
Habitat. Occurring in sand, mud, algae, calcareous
substrata, bryozoans, sponges, from intertidal to 90 m
(Hartmann-Schröder, 1984).
Distribution. Angola, Namibia, South Africa, USA
(Southern California), Panama, Indo-Pacific, New Zealand,
Australia (Queensland, New South Wales, Tasmania, South
Australia, Western Australia).
Genus Opisthodonta Langerhans, 1879
Opisthodonta Langerhans, 1879: 547.
Type species. Opisthodonta morena Langerhans, 1879.
Diagnosis. Body long, with numerous segments, stout,
dorsally convex, of macrofaunal size (>10 mm in length).
Prostomium provided with 2 pairs of eyes, and sometimes
2 anterior eyespots. Three antennae. Median antenna

San Martín & Hutchings: Eusyllinae of Australia 301
Fig. 36. SEM of Odontosyllis polycera Schmarda, 1863 (A) occipital flap; (B) detail of dorsum of midbody segments; (C) transverse
section, midbody of an atokous specimen; (D) transverse section, midbody of an epigamic specimen; (E) fascicle of chaetae,
midbody; (F) compound chaetae, midbody. A,B: AM W195387; C: AM W4344; D: AM W14203; E,F: AM W4344.
inserted on middle of prostomium or slightly in front of
anterior eyes. Palps apparently free from each other, basally
fused. Peristomium small, partially covered by prostomium
and first chaetiger, with 2 pairs of tentacular cirri. Nuchal
organs as 2 ciliated grooves between prostomium and
peristomium. Dorsal cirri on all chaetigers, cylindrical, long
to extremely long, smooth or slightly rugose. Parapodia
elongate; ventral cirri of anterior parapodia, ovate,
foliaceous almost completely fused with parapodial lobe,
provided with hyaline inclusions, perhaps glands.
Subsequent ventral cirri conical to digitiform, neither
foliaceous nor fused to parapodial lobes, inserted at base of
parapodia. Chaetal bundles formed of numerous compound
chaetae, including falcigers provided with long, thick
proximal tooth and short distal tooth and sometimes a few
chaetae with long, slender, spiniger-like blades. Dorsal
simple chaetae apparently lacking. Aciculae with button-
shaped tips with crown of spines, or tricuspidate. Pharynx
and proventricle similar in length. Pharynx with crown of
soft papillae on anterior rim and single mid-dorsal tooth,
inserted on anterior third or middle of pharynx. Repro-
duction by epigamy (Garwood, 1991).

302 Records of the Australian Museum (2006) Vol. 58
Key to Australian species of Opisthodonta
1 Pharyngeal tooth inserted mid to far posterior of pharynx. Spiniger-
like chaetae absent ......................................................................................................... O. morena
—— Pharyngeal tooth located in front of middle of pharynx. Spiniger-
like chaetae present .........................................................................................................................2
2 Spiniger-like chaetae about 84 µm in midbody, bidentate. Dark,
transverse rows of pigment on anterior segments ............................................. O. melaenonephra
—— Spiniger-like chaetae about 175 µm in midbody, weakly bidentate
to unidentate. Lacking any colour pattern ..................................................... O. hanneloreae n.sp.
Fig. 37. Opisthodonta hanneloreae n.sp. (A) anterior end, dorsal view; (B) spiniger-like compound chaetae, anterior parapodium; (C)
falcigers, anterior parapodium; (D) spiniger-like compound chaeta, midbody; (E) falcigers, midbody; (F) aciculae, anterior parapodium;
(G) aciculae, midbody. AM W28393. Scales: A 0.18 mm, B–G 20 µm.
Opisthodonta hanneloreae n.sp.
Fig. 37A–G
Material examined. HOLOTYPE (AM W28393). AUSTRALIA:
WESTERN AUSTRALIA: Wallabi Group of Is., 28°23.99'S 113°46.73'E,
shell debris from scallop beds, 39 m, coll. P.A. Hutchings on WA FRV
Flinders, 30 May 1994. PARATYPES Wallabi Group of Is., 28°24'S
113°46.26'E, scallop beds, shell debris, 35 m, coll. P.A. Hutchings on
WA F RV Flinders, 30 May 1994, few (AM W28372); Off S end of Long
Is. Beacon Is. 28°28.8'S 113°46.3'E, dead coral substrate covered in
coralline algae, 4.5 m, coll. P.A. Hutchings, 25 May 1994, 1 (AM
W28950); Goss Passage, Beacon Is. 28°25.5'S 113°47'E, dead coral
substrate, in fine sediment at foot of reef slope, 33 m, coll. P.A. Hutchings,
23 May 1994, 1 (AM W28951).

San Martín & Hutchings: Eusyllinae of Australia 303
Opisthodonta melaenonephra
(Haswell, 1920) n.comb.
Figs 38A–G, 39A–O
Pionosyllis melaenonephra Haswell, 1920: 103, pl. 12, figs 11–
16, pl. 13, fig. 1.—Hutchings & Murray, 1984: 32.
Material examined. AUSTRALIA: NEW SOUTH WALES: Offshore from
Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy mud, 4 m,
coll. A.R. Jones & A. Murray, 9 Nov 1982, 2 (AM W22122); Offshore
from Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy mud, 4
m, coll. A.R. Jones & A. Murray, 5 Aug 1983, 1 (AM W22123); Offshore
from Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy mud, 4
m, coll. A.R. Jones & A. Murray, 11 Nov 1983, 1 (AM W22124); Offshore
from Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy mud, 4
m, coll. A.R. Jones & A. Murray, 17 May 1982, 1 (AM W22126);
Offshore from Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy
mud, 4 m, A.R. Jones & A. Murray, 26 May 1981, 1 (AM W22127);
Offshore from Hungry Beach, Hawkesbury R., 33°35'S 151°17'E, sandy
mud, 4 m, coll. A.R. Jones & A. Murray, 9 Nov 1982, 1 (AM W22128);
mid-stream between Juno Head and Hungry Beach, Hawkesbury R.,
33°34'S 151°16'E, muddy sand, 10 m, coll. A.R. Jones & A. Murray,
9 Nov 1984, 1 (AM W22129); 50 m NE of Green Point, Hawkesbury R.,
33°34'S 151°13.5'E, muddy sand, 4 m, coll. A.R. Jones & A. Murray, 9 Nov
1982, 1 (AM W22131); 50 m NE of Green Point, Hawkesbury R., 33°34'S
151°13.5'E, muddy sand, 4 m, coll. A.R. Jones & A. Murray, 5 Aug 1983, 2
(AM W22132); 300 m NE of Green Point, Hawkesbury R., 33°34'S
151°13.5'E, sandy mud, 5 m, coll. A.R. Jones & A. Murray, 26 May 1981,
1 (AM W22133); 300 m NE of Green Point, Hawkesbury R., 33°34'S
151°13.5'E, sandy mud, 5 m, coll. A.R. Jones & A. Murray, 7 Aug 1981,
1 (AM W22134); 300 m NE of Green Point, Hawkesbury R., 33°34'S
151°13.5'E, sandy mud, 5 m, coll. A.R. Jones & A. Murray, 26 May
1981, 1 (AM W22135); 300 m NE of Green Point, Hawkesbury R., New
South Wales, Australia, 33°34'S 151°13.5'E, sandy mud, 5 m, coll. A.R.
Jones & A. Murray, 11 Nov 1983, 1 (AM W22136); 1 km S of E end of
Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy sand, 12 m,
coll. A.R. Jones & A. Murray, 5 Aug 1983, 1 (AM W22138); 1 km S of
E end of Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy sand,
12 m, coll. A.R. Jones & A. Murray, 21 Aug 1984, 1 (AM W22139); 300 m
NE of Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m,
coll. A.R. Jones & A. Murray, 4 Feb 1983, 1 (AM W22140); 300 m NE of
Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll.
A.R. Jones & A. Murray, 11 Feb 1981, 1 (AM W22141); 300 m NE of
Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll.
A.R. Jones & A. Murray, 27 May 1983, 1 (AM W22142); 300 m NE of
Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll.
A.R. Jones & A. Murray, 11 Feb 1981, 3 (AM W22143); 300 m NE of
Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll.
A.R. Jones & A. Murray, 7 Aug 1981, 1 (AM W22144); 300 m NE of
Green Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll.
A.R. Jones & A. Murray, 7 Aug 1981, 1 (AM W22145); 1 km S of E end
of Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy sand, 12
m, coll. A.R. Jones & A. Murray, 9 Feb 1984, 1 (AM W22147); 1 km S
of E end of Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy
sand, 12 m, coll. A.R. Jones & A. Murray, 21 Aug 1984, 1 (AM W22148);
E end of Brooklyn Boat Channel, Hawkesbury R., 33°33'S 151°14'E,
12 m, coll. A. Jones et al., 18 Dec 1979, 1 (AM W196608). E end of
Brooklyn Boat Channel, Hawkesbury R., 33°33'S 151°14'E, A. Jones et
al., 1 Aug 1979, 5 (AM W196610); 200 m S of E end of Spectacle Is.,
Hawkesbury R., 33°32.5'S 151°13.5'E, sandy mud, 5 m, coll. A.R. Jones
& A. Murray, 11 Nov 1983, 1 (AM W22150). Pittwater, 33°35.87'S
151°18.71'E, mud, 15.9 m, coll. Australian Museum Party, 15 Dec 1994,
1 (AM W23929); Port Jackson, 33°51'S 151°16'E, Feb 1920. Syntypes:
Port Jackson, 33°51'S 151°16'E, Feb 1920, 2 (AM W513), 8 (AM
W6175), 1 on microscope slide (AM W494). Rock platform, Murray’s
Beach, Jervis Bay, 35°07.5'S 150°46'E, dead barnacles encrusted with
sponges, in intertidal pools, 0.5 m, coll. H.E. Stoddart, 28 Jun 1981, 1
(AM W28412); Summercloud Bay, Jervis Bay, 35°10.4'S 150°41.2'E,
15.8 m, coll. P.A. Hutchings, 29 Nov 1971, 1 (AM W28435); Murrays
Basin, Jervis Bay, 35°07.5'S 150°45.5'E, sand, Zostera sp., coll. NSW
State Fisheries, 17 Oct 1972, 1 (AM W194168).
Description. Body long (>10 mm in length) and broad,
fragile, strongly convex dorsally, dark, 1–3 transverse dark
bands on dorsum of anterior segments, as well as two black
Description. Body fragile, all fragmented specimens,
strongly convex dorsally, colourless, longest anterior
fragment 2.8 mm long, 0.6 mm wide, with 18 chaetigers.
Prostomium oval, with 4 eyes in open trapezoidal
arrangement and sometimes 2 anterior eyespots; median
antenna inserted in front of line between anterior eyes, about
twice as long as combined length of prostomium and palps;
lateral antennae inserted near anterior margin, about half
length of median antenna (Fig. 37A). Palps trapezoidal,
sometimes ventrally folded, slightly longer than pro-
stomium. Peristomium similar in length to following
segments; dorsal tentacular cirri long, filiform, longer than
median antenna, ventral tentacular cirri about one third
length of dorsal cirri. Dorsal cirri, as well as antennae and
tentacular cirri, smooth, filiform, alternating irregularly, long
cirri, longer than body width, and short cirri, shorter than
body width (Fig. 37A). Parapodial lobes rectangular, ending
with pre-chaetal lobe. Ventral cirri large, triangular, partially
fused to parapodial lobes on anterior parapodia, with some
granular inclusions, becoming digitiform, not fused to
parapodial lobes, on subsequent segments. Compound
chaetae compound heterogomph, shafts distally spinose, and
two kinds of blades, long, spiniger-like chaetae, anteriorly
bifid (Fig. 37B), apparently unidentate on posterior
parapodia (Fig. 37D), and bidentate falcigers, elongate, with
short spines on margin and margins of blades weakly convex
on anterior parapodia, bidentate, with unequal teeth,
proximal tooth longer and broader than distal tooth (Fig.
37C), becoming more marked on posterior chaetae (Fig.
37E). Anterior parapodia with about 3 spiniger-like chaetae,
blades 90–175 µm in length; and about 50 falcigers with
blades 25–27 µm in length; progressively posteriorly
number of compound chaetae decreasing to 1–2 spiniger-
like, about 105 µm in length, smooth and unidentate, and
25 falcigers, 25–22 µm in length, on mid-posterior
parapodia. Simple dorsal and ventral chaetae not seen.
Aciculae distally slightly enlarged, ending in button, 4 on
most anterior parapodia (Fig. 37F), reducing to 2 on mid-
posterior parapodia, one large and other slender (Fig. 37G).
Pharynx long and slender, through about 11 segments;
pharyngeal tooth long and large, oval, located slightly
anteriorly to middle of pharynx (Fig. 37A). Proventricle
large, through about 6 segments, with 23 muscle cell rows.
Details of posterior end unknown.
Remarks. Opisthodontha hanneloreae n.sp. is similar to
O. melaenonephra, but lacks any colour pattern, the
spiniger-like chaetae are distinctly longer and more slender,
unidentate from midbody onwards. Opisthodonta morena
Langerhans, 1879 (see above), and O. mitchelli Kudenov
& Harris (1995) have much shorter spiniger-like chaetae
than O. hanneloreae.
Habitat. Occurring in dead corals and in shell debris, in
shallow water.
Distribution. Australia (Western Australia).
Etymology. The species is named after Dr Hannelore Paxton
an Australian polychaetologist.

304 Records of the Australian Museum (2006) Vol. 58
Fig. 38. Opisthodonta melaenonephra (Haswell, 1920) (A) anterior
end, dorsal view, epigamic specimen; (B) detail of prostomium
and everted pharynx, dorsal view; (C) anterior end, ventral view;
(D) parapodial lobe, dorsal view; (E) anterior parapodium; (F)
midbody parapodium; (G) pharynx and proventricle. AM W28412.
Scales: A–C,G: 0.18 mm; D 48 µm; E,F: 92 µm.
spots on each palp, with dark pigment on prostomium; one
epigamic specimen (AM W28412) with this pattern well
developed (Fig. 38A), but others less pigmented. (Haswell
recorded an individual 14 mm long, 1 mm wide for 65–75
segments). Prostomium ovate with 4 eyes arranged in open
trapezoidal pattern and, sometimes 2 small anterior eyespots.
Median antenna long, inserted on middle of prostomium,
between anterior eyes, slightly longer than twice combined
length of prostomium and palps; lateral antennae shorter,
about half of median antenna, arising in front of anterior
pair of eyes (Fig. 38A). Palps apparently basally free, ovate
(Fig. 38B), sometimes ventrally folded. Dorsal tentacular
cirri similar in length to median antenna, ventral tentacular
cirri similar to lateral antennae. Antennae, tentacular cirri
and dorsal cirri long, slender, cylindrical, smooth on
midbody and posterior segments, to rugose, not articulated,
supported by short cirrophores or indistinctly articulated
on antennae, dorsal tentacular cirri and most anterior dorsal
cirri (Fig. 38A,E). Dorsal cirri alternating, twice as long as
body width on most anterior segments, and short, similar

San Martín & Hutchings: Eusyllinae of Australia 305
Fig. 39. Opisthodonta melaenonephra (Haswell, 1920) (A) distal end of spiniger-like chaeta, anterior parapodium; (B) blade of spiniger-
like chaeta, anterior parapodium; (C,D) blades of falcigers, anterior parapodium; (E) spiniger-like chaeta, midbody; (F, G ) blades of
falcigers, midbody; (H) spiniger-like blade, posterior parapodium; (I–L) blades and falcigers, posterior parapodium; (M) acicula,
posterior parapodium; (N) ventral simple chaeta; (O) aciculae, anterior parapodium. AM W494. Scales: 20 µm.
in length to body width. Parapodia slender, elongate, sub-
rectangular, with 2 rounded lobes on anterodorsal and
posterodorsal margins of parapodia (Fig. 38D). Ventral cirri
of anterior parapodia foliaceous, thick, partially fused to
base of parapodial lobe, with hyaline inclusions near ventral
surface (Fig. 38E), becoming progressively shorter, not
fused to parapodial lobe, digitiform (Fig. 38F). Anterior
parapodia, except those of 2–3 most anterior segments, with
2–3 compound chaetae with long, slender, spiniger-like
blades, about 125 µm long, indistinctly bifid distally (Fig.
39A,B), with proximal tooth slightly larger than distal tooth
and short, indistinct spines on margin, and about 15
compound falcigerous chaetae, with elongate, bidentate
blades provided with short and slightly hooked distal tooth
(Fig. 39C,D), larger and triangular proximal tooth, and short,
straight spines on margin; dorsoventral gradation in length
of blades within fascicle, 65 µm in length dorsally, 25 µm
in length ventrally, ventral blades larger with marked
difference in size of teeth (Fig. 39E–G); blades with margins
weakly convex, becoming more marked ventrally within
fascicle. Posteriorly, blades of spiniger-like chaetae
becoming progressively shorter, and falcigers with wider
blades; posterior parapodia with single compound chaetae
with elongate, spiniger-like blade (Fig. 39H), about 60 µm
in length, and 3–4 chaetae with falcigerous blades with
dorsoventral gradation in length within fascicle, 50 µm in
length dorsally, 25 µm in length ventrally; spiniger-like
blade with margins weakly convex and distally bidentate,
with both teeth almost equal (Fig. 39I–L). Dorsal simple
chaetae not seen, probably absent. Ventral simple chaetae
only seen on far posterior parapodia of holotype, sigmoid,
stout, strongly bidentate, bearing proximal tooth longer than
distal tooth (Fig. 39N). Anterior parapodia with 3
tricuspidate tipped aciculae (Fig. 39 O) remaining
parapodia, except posteriormost ones, with 2 similar
aciculae. Posteriormost parapodia, each with single acicula,
tip distally rounded, with button (Fig. 39M). Pharynx short
and wide, extending through about 7–8 segments, with
crown of about 16 soft papillae; pharyngeal tooth conical,
large, located just in front of middle of pharynx (Fig.

306 Records of the Australian Museum (2006) Vol. 58
38A,G). Proventricle through 4–7 chaetigers, with about
23 muscle cell rows. Pygidium with 2 anal cirri.
Habitat. Occurring in soft sediments, intertidally to depths
of 50 m; also recorded from dead barnacles covered with
encrusting sponges.
Distribution. Australia (New South Wales).
Opisthodonta morena Langerhans, 1879
Figs 40A–F, 41A–F, 42A,B
Opisthodonta morena Langerhans, 1879: 547, fig. 12.—Pérès,
1954: 107, figs 3–6.—Laubier, 1966: 249.—Storch, 1966: 173,
pl.1, fig. 2.—Hartmann-Schröder, 1971: 100, Figs 1–3.—
Campoy, 1982: 307.—San Martín, 2003: 54, figs 15, 16.
Material examined. TASMAN SEA: Reef flat near Yoshin Maru Iwaki
wreck, Elizabeth Reef, 29°55.8'S 159°01.3'E, reef flat at low tide, 14 Dec
1987, 1 on SEM stub (AM W28874); Taupo Seamount, 33°16.85'S
156°09.15'E, limestone & sand bottom, 244 m, coll. J.K. Lowry & party, 2
May 1989, 2 on SEM stub (AM W28875); Taupo Seamount, 33°16.85'S
156°09.15'E, limestone & sand bottom, 244 m, coll. J.K. Lowry & party, 2
May 1989, 2 (AM W28930). WESTERN AUSTRALIA: Goss Passage, Beacon
Is., 28°25.5'S 113°47'E, dead Acropora plates covered in algae, sponges &
ascidians, 23 m, coll. P.A. Hutchings, 19 May 1994, 1 (AM W28384); N
end of Long Is., 28°27.9'S 113°46.3'E, dead coral substrate covered in
coralline & brown algae, 5.5 m, coll. C. Bryce, 22 May 1994, 1 (AM
W28954). N end of Long Is., 28°28.3'S 113°46.3'E, dead coral substrate,
coralline algae, boring bivalves, 8 m, coll. C. Bryce, 22 May 1994, 1 (AM
W28955); Goss Passage, Beacon Is., 28°25.5'S 113°47'E, dead plates of
Acropora coral covered in coralline algae, 8 m, coll. P.A. Hutchings, 22
May 1994, 1 (AM W28956); Goss Passage, Beacon Is., 28°25.5'S 113°47'E,
dead plates of Acropora coral, 8 m, coll. P.A. Hutchings, 19 May 1994, 1
(AM W28957); Off jetty adjacent to Fisheries Hut, Beacon Is., 28°25.5'S
113°47'E, dead coral substrate, plate-like Acropora & Montipora spp., 12
m, coll. P.A. Hutchings, 23 May 1994, 1 (AM W28958); Reef S of Lucas
Is., Brunswick Bay, Kimberley region, 15°16'S 124°29'E, 2 m, coll. P.A.
Hutchings, 24 July 1988, several (AM W28931).
Fig. 40. Opisthodonta morena Langerhans, 1879
(A) anterior end, dorsal view; (B) ventral view of
anteriormost end; (C) aciculae, anterior para-
podium; (D) acicula, posterior parapodium; (E)
long-bladed compound chaeta, midbody; (F)
compound chaetae, midbody. AM W28930.
Scales: A 0.4 mm, B 0.18 mm, C–F 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 307
Fig. 41. SEM of Opisthodonta morena Langerhans, 1879 (A) anterior end, dorsal view; (B) detail of prostomium (median antenna
missing); (C) anterior end, ventral view; (D) detail of anteriormost end, ventral view; (E) pharyngeal opening; (F) compound
chaetae, anterior parapodium. AM W28874, AM W28875.
Description. Body fragile from proventricular segments,
only anterior fragments examined, longest fragment 3.7 mm
long, 0.4 mm wide, with 26 chaetigers. Prostomium oval to
pentagonal (Figs 40A, 41B), with 2 pairs of eyes in
trapezoidal arrangement and 2 anterior eyespots; median
antenna inserted on middle of prostomium, more than three
times combined length of prostomium and palps; lateral
antennae about half length of median antenna, inserted near
anterior margin of prostomium. Palps broad, large,
trapezoidal, thin, usually ventrally folded (Figs 40B, 41D);
apparently free from each other (Fig. 40A), but fused basally
(Fig. 41B). Peristomium dorsally reduced, covered by
chaetiger 1; dorsal tentacular cirri similar to median antenna,
but shorter, ventral tentacular cirri about two thirds length
of dorsal ones. Posterior margin of prostomium and nuchal
organs densely ciliated (Figs 40A, 41B) dorsal cirri similar
to antennae, smooth, filiform, long and slender, long on
anterior segments, (Figs 40B, 41A), alternating long cirri,
twice body width, and short cirri, shorter than body width,
from proventricular segments onwards. Parapodial lobes
conical, distally bifid (Fig. 40B). Ventral cirri large, ovate,
partially fused to parapodial lobes on anterior segments
(Figs 40C, 41D), conical, shorter and not fused from
proventricular segments onwards (Fig. 41C). Compound
chaetae numerous, 17–23 anteriorly, decreasing to about 7
on midbody, heterogomph, shafts subdistally spinose,
similar throughout; on midbody, 2–3 compound chaetae
with slender, elongate, bidentate blades (Figs 40E, 41F,

308 Records of the Australian Museum (2006) Vol. 58
Fig. 42. SEM of Opisthodonta morena Langerhans, 1879 (A) long-bladed compound chaeta; (B) compound chaetae. SEM of
Pionosyllis yolandae n.sp. (C) anterior end, dorsal view; (D,E) detail of prostomium and anteriormost segments; (F) pharyngeal
opening. A,B: AM W28874, AM W28875; C–F: AM W28876.
42A), about 25–26 µm in length, short spines on margin, 2
compound chaetae with similar blades but shorter, 15–16
µm in length, marginal spines, distally directed, and
remaining compound chaetae with short, bidentate blades,
proximal tooth longer than distal tooth, and few spines on
margin (Figs 40F, 41F, 42B), moderate in length, distally
directed, dorsoventral gradation in length within fascicle,
13 µm in length dorsally, 8 µm in length ventrally. Dorsal
and ventral simple chaetae absent. Anterior parapodia with
3 aciculae, straight, distally broad (Fig. 40C), diminishing
to single acicula posteriorly (Fig. 40D). Pharynx long and
wide, through 9–11 segments, pharyngeal tooth inserted
posteriorly (Fig. 40A), on about chaetigers 7–8; pharyngeal
opening with some soft papillae and dense layer of cilia
(Fig. 42E). Proventricle through 6 segments, with about 24
muscle cell rows.
Remarks. This species has been reported widely from
tropical and subtropical regions; we have examined material
from the Mediterranean, and the European Atlantic coast
and they appear morphologically similar. A molecular study
would be useful to confirm whether this is a widely
distributed species or a suite of sibling species.
Habitat. Occurring interstitially in coarse and coral sand,
seagrasses, mud, from intertidal to depths greater than 240 m.
Distribution. Western Atlantic, Mediterranean, Red Sea,
Australia (New South Wales, Western Australia).

San Martín & Hutchings: Eusyllinae of Australia 309
Genus Paraehlersia San Martín, 2003
Paraehlersia San Martín, 2003: 61.
Type species. Ehlersia ferrugina Langerhans, 1881,
designated by San Martín, 2003.
Diagnosis. Body long, stout, with numerous segments, adults
5 mm or greater in length, dorsally convex. Prostomium with
4 eyes and pair of anterior eyespots. Three antennae. Median
antenna inserted on middle of prostomium. Palps basally fused,
with dorsal furrow. Two pairs of tentacular cirri. Nuchal organs
as 2 ciliated grooves between prostomium and peristomium.
Antennae, tentacular cirri and anterior dorsal cirri of adults
articulated to irregularly articulated depending upon size,
remaining dorsal cirri smooth. Dorsal ciliary bands on
segments. Parapodia without prechaetal lobes; digitiform,
retractile papilla between parapodial lobes and dorsal cirri of
some parapodia. Compound chaetae heterogomph, including
one or more chaetae with spiniger-like blades and several
bidentate falcigerous blades, anteriorly with both teeth similar,
posteriorly with proximal tooth longer and more robust than
distal tooth. Aciculae acuminate. Pharynx and proventricle of
similar size. Pharyngeal tooth anteriorly located. Reproduction
by epigamy (San Martín, 2003).
Key to Australian species of Paraehlersia
1 Falcigers and ventral simple chaetae provided with distal long
spines on margin, reaching or extending beyond level of proximal
tooth.................................................................................................................. P. weissmannioides
—— Chaetae with short spines on margin (Fig. 44D) ................................................ P. ehlersiaeformis
Paraehlersia ehlersiaeformis (Augener, 1913)
n.comb.
Figs 43D–F, 44A–K, 45A–F, 46A–C
Pionosyllis ehlersiaeformis Augener, 1913: 225, figs 31, 32.
Syllis (Ehlersia) ferrugina.—Haswell, 1920: 101, pl. 12, figs 3–
10. Not Langerhans, 1881: 104.
Typosyllis (Langerhansia) ferrugina.—Hartmann-Schröder, 1981:
30; 1987: 37; 1989: 23; 1991: 33. Not Langerhans, 1881: 104.
Material examined. AUSTRALIA: NEW SOUTH WALES: Taupo
Seamount, Tasman Sea, 33°16.85'S 156°09.15'E, limestone & sand
bottom, 244 m, coll. J.K. Lowry & party on RV Franklin, 2 May 1989,
1 on SEM stub (AM W28899); Taupo Seamount, Tasman Sea, 33°16.85'S
156°09.15'E, limestone & sand bottom, 244 m, coll. J.K. Lowry & party
on RV Franklin, 2 May 1989, 22 (AM W28939); Pittwater, 33°35.78'S
151°18.33'E, sand, 13.8 m, coll. Australian Museum Party, 22 Apr 1994,
2 (AM W23925); Pittwater, 33°35.77'S 151°18.34'E, muddy sand, 14.9
m, coll. Australian Museum Party, 9 Oct 1995, 1 on SEM stub (AM
W23926); Bass Point, 34°36'S 150°54'E, 50 m, The Ecology Lab for
RMI/Pioneer Project, 1 Feb 1990, 1 (AM W23063); Halfway Reef, 200
m S of Sullivan Reef, Ulladulla, 35°21.42'S 150°29.31'E, airlift over
sponges, Bryozoa & Hydrozoa, 15 m, coll. K. Attwood & party, 3 May
1997, 1 on SEM stub (AM W28221); N side of Bannister Head, north of
Ulladulla, 35°19.15'S 150°29.12'E, grey sponge from top of boulder, 18
m, coll. K. Attwood, 6 May 1997, few (AM W28223); Murray’s Beach,
Jervis Bay, 35°07.5'S 150°46'E, 10 m, coll. P.A. Hutchings, 23 Jan 1973,
2 (AM W28413); 350 m S of southern entrance to Jervis Bay, 35°7.7'S
150°46.05'E, 23 m, coll. P.A. Hutchings, 22 July 1972, 1 (AM W28436);
S of Worang Point, Calle Calle Bay, Twofold Bay, 37°3.6'S 149°56.5'E,
6.1 m, coll. S. Keable, P. Albertson, 21 Feb 1985, 1 (AM W28440).
SOUTH AUSTRALIA: 7 km NW of Port Davis Creek, Port Pirie, Spencer
Gulf, 33°16'S 137°51'E, subtidal, unvegetated, 9.3 m, coll. T.J. Ward &
party, Mar 1980, 1 (AM W21772); Boston Bay, Port Lincoln 34°51'S
135°51'E, washings from sheltered weedy rock, 2 m, coll. I. Loch, 12
Feb 1985, 1 (AM W28937); Billy Lights Point, Port Lincoln, 34°45'S
135°53'E, stone washings from sheltered intertidal rocks, coll. I. Loch,
15 Feb 1985, 1 (AM W28938). WESTERN AUSTRALIA: Goss Passage,
Beacon Is., 28°25.5'S 113°47'E, dead plates of Acropora coral, 8 m,
coll. P.A. Hutchings, 19 May 1994, several (1 epigamic on SEM stub)
(AM W28373); Goss Passage, Beacon Is., 28°25.5'S 113°47'E, dead
plates of Acropora coral covered in coralline algae, 8 m, coll. P.A.
Hutchings, 22 May 1994, several (AM W28375); NE entrance to Goss
Passage, Beacon Is., 28°27.9'S 113°46.7'E, dead branching Acropora,
coralline & brown algae, 24 m, coll. P.A. Hutchings, 25 May 1994, 4
(AM W28377); Off S end of Long Is., Beacon Is., 28°28.8'S 113°46.3'E,
dead coral substrate covered in coralline algae, 4.5 m, coll. P.A.
Hutchings, 25 May 1994, 1 (AM W28379); Goss Passage, Beacon Is.,
28°25.5'S 113°47'E, dead branching coral covered in coralline algae,
10 m, coll. P.A. Hutchings, 18 May 1994, 1 (AM W28382); Goss Passage,
Beacon Is., 28°25.5'S 113°47'E, dead Acropora plates covered in algae,
sponges & ascidians, 32 m, coll. P.A. Hutchings, 19 May 1994, 1 (AM
W28383); Goss Passage, Beacon Is., 28°25.5'S 113°47'E, dead Acropora
plates covered in coralline algae, 20 m, coll. P.A. Hutchings, 20 May
1994, 1 (AM W28385); E side of West Wallabi Is., 28°27.9'S 113°40.9'E,
in Posidonia root mat, plus epifauna, 1.5 m, coll. P.A. Hutchings, 26
May 1994, several (AM W28388); NE entrance to Goss Passage, Beacon
Is., 28°27.9'S 113°46.7'E, under boulders in coral sand at foot of reef
slope, 33 m, coll. P.A. Hutchings, 25 May 1994, 2 (AM W28389); Wallabi
Group of Is., 28°34.65'S 113°46.46'E, diverse sponges & rubble, 49 m,
coll. P.A. Hutchings on WA FRV Flinders, 28 Jun 1994, 1 (AM W28391);
Wallabi Group of Is., 28°23.61'S 113°45.09'E, sponge & shell debris
from scallop beds, 35 m, coll. P.A. Hutchings on WA FRV Flinders, 30
May 1994, 1 (AM W28394); Off S end of Long Is., Beacon Is., 28°28.8'S
113°46.3'E, dead coral substrate covered in coralline algae, 4.5 m, coll.
P.A. Hutchings, 25 May 1994, 2 (AM W28934); N end of Long Is.,
28°28.3'S 113°46.3'E, dead coral substrate, coralline algae, boring
bivalves, 8 m, coll. C. Bryce, 22 May 1994, few (AM W28972); N end
of Long Is., 28°27.9'S 113°46.3'E, dead coral substrate covered in
coralline & brown algae, 5.5 m, coll. C. Bryce, 22 May 1994, several
(AM W28973); Wallabi Group of Is., 28°27.05'S 113°45.10'E, medium
to fine sand & shell debris from scallop beds, 36.5 m, coll. P.A. Hutchings
on WA FRV Flinders, 30 May 1994, 1 (AM W28935); 5 km offshore,
Bush Bay, 30 km S of Carnarvon, 25°10'S 113°39'E, shallow strap-leaved
seagrass beds, 2 m, coll. J.K. Lowry & R.T. Springthorpe, 6 Jan 1984, 1
(AM W28371); N end of beach, Bundegi Reef, Exmouth Gulf, 21°49'S
114°11'E, rocky rubble, coralline algae with green epiphyte, 1.5 m, coll.
H.E. Stoddart, 4 Jan 1984, 1 (AM W28936); Reef S of Lucas Is.,
Brunswick Bay, Kimberley region, 15°16'S 124°29'E, 2 m, coll. P.A.
Hutchings, 24 July 1988, 1 (AM W28940); north end of beach, Bundegi
Reef, Exmouth Gulf, 21°49'S 114°11'E, rocky rubble, coralline algae
with green epiphyte, 1.5 m, coll. H.E. Stoddart, 4 Jan 1984, several
(AM W28974). Syntype: ZMH (P-8786), Sharks Bay, Western Australia.
Description. Body without colour markings, some
specimens broad anteriorly (Fig. 43D), others slender,
filiform, about 9 mm long, 0.4 mm wide, with 75 chaetigers.
Prostomium oval to circular, 4 eyes arranged in open
trapezoidal pattern. Median antenna long, slender, with
numerous articulations, usually 2–3 times longer than
combined length of prostomium and palps, inserted on
middle of prostomium; lateral antennae similar to median
one, but shorter, inserted in front of anterior eyes. Palps
slightly longer than prostomium, triangular (Figs 43E, 44A).
Peristomium distinct, similar in length to subsequent
segments; tentacular cirri similar to antennae, dorsal
tentacular cirri slightly longer than lateral antennae, ventral
ones about two thirds length of dorsal ones. Anterior dorsal
cirri irregularly articulated (Figs 43E, 44A), becoming

310 Records of the Australian Museum (2006) Vol. 58
Fig. 43. SEM of Paraehlersia weissmannioides (Augener, 1913) (A) compound chaetae, posterior parapodium; (B) dorsal simple
chaeta; (C) ventral simple chaeta. SEM of Paraehlersia ehlersiaeformis (Augener, 1913) (D) complete specimen, dorsal view; (E)
anterior end, dorsal view; (F) lateral view, midbody (arrows showing subcirral papillae). A–C: AM W28224; D–F: AM W23926,
W28221, W28899.
smooth, filiform (Figs 43F, 44A), from chaetiger 5–6,
shorter than anterior ones, alternating irregularly long cirri,
slightly longer than body width, and shorter ones about
equal to body width. Dorsum of each segment with 2 rows
of cilia (Figs 43F, 45B), except anteriormost, with only 1
row (Fig. 43E). Subcirral papilla small (Figs 43F, 44B, 45B
arrows, 45C), difficult to see, present on anterior and
midbody segments. Parapodial lobes sub-rectangular (Fig.
44B). Ventral cirri digitiform, similar in length to parapodial
lobes. Anterior parapodia with up to 12 compound chaetae,
2–4 with elongated, slender, blades about 33 µm in length,
bidentate with both teeth similar (Fig. 44C), and short spines
on margin; remaining chaetae with shorter blades, bidentate,
both teeth similar, and dorsoventral gradation in length
within fascicle (Fig. 44D), 18 µm in length dorsally and 11
µm in length ventrally. Progressively along body, 1–2 blades
of dorsalmost compound chaetae becoming more elongated,
spiniger-like, indistinctly bidentate, with short spines on
margin (Fig. 44F,I), 47 µm in length on midbody, 50 µm in
length on posterior parapodia. Blades of falcigers with large
proximal tooth and shorter distal tooth, short spines on
margin (Figs 44G,J, 45D,F, 46A), 10–15 µm in length on
midbody, 10–11 µm in length on posterior parapodia; about
6–7 falcigers on midbody parapodia and 4–5 posteriorly.
Dorsal simple chaetae from mid-posterior segments, distally
bifid, with short subdistal spines (Figs 44H, 46B). Ventral

San Martín & Hutchings: Eusyllinae of Australia 311
Fig. 44. Paraehlersia ehlersiaeformis (Augener, 1913) (A) anterior end, dorsal view; (B) anterior
parapodium (arrow showing subcirral papilla); (C) spiniger-like chaeta, anterior parapodium; (D)
falcigers, anterior parapodium; (E) aciculae, anterior parapodium; (F) spiniger-like compound
chaeta, midbody; (G) falcigers, midbody; (H) dorsal simple chaeta; (I) spiniger-like compound
chaeta, posterior parapodium; (J) falcigers, posterior parapodium; (K) ventral simple chaeta; (L)
acicula, posterior parapodium. AM W28377. Scales: A 0.18 mm, B 48 µm, C–L 20 µm.
simple chaetae on posterior segments, smooth, strongly
bidentate, proximal tooth long, robust, and distal tooth small
(Figs 44K, 46C). Anterior parapodia with 3 aciculae (Fig.
63E), reducing to 1 from midbody, distally acuminate or
lancet-shaped (Fig. 44L). Pharynx through 7 segments;
pharyngeal tooth on anterior margin (Fig. 44A), surrounded
by crown of 10 soft papillae and layer of cilia (Fig. 45A).
Proventricle through 5 segments, with about 21 muscle cell
rows. Pygidium with 2 long, filiform anal cirri and median
papilla. Some specimens in epigamic reproductive phase,
with long capillary chaetae on some parapodia (Fig. 45E).
Remarks. The specimens from Australia are similar to
specimens of P. ferrugina Langerhans, 1879, which has been
widely reported from Europe and northern Atlantic.
Examination of one syntype of P. ehlersiaeformis (HZM P-
8786) reveals that some blades of chaetae from posterior-
most segments are slightly wider than those described from
P. ferrugina. At this stage, we prefer to regard P.
ehlersiaeformis and P. ferrugina as separate species.
Augener (1913) reported Syllis (Ehlersia) ferrugina from
Western Australia. This record, however, seems to be another
species, probably belonging to the subfamily Syllinae.

312 Records of the Australian Museum (2006) Vol. 58
Fig. 45. SEM of Paraehlersia ehlersiaeformis (Augener, 1913) (A) everted pharynx; (B) subcirral papillae, showed by arrows; (C)
detail of subcirral papilla; (D) compound chaetae, mid-anterior parapodium; (E) midbody parapodium of epigamic specimen,
showing capillaries and notochaetae; (F) falciger, midbody. AM W23926, W28221, W28899.
Habitat. Reported from wide variety of substrates,
especially abundant in algae and coralline concretions, from
shallow waters to depths greater than 100 m.
Distribution. Australia (all States).
Paraehlersia weissmannioides (Augener, 1913)
n.comb.
Figs 43A–C, 47A–I, 48A–F, 49D–F
Pionosyllis weissmannioides Augener, 1913: 223, fig. 30.
?Ehlersia ferrugina non Langerhans, Böggemann & Westheide,
2004: 418, fig. 6.
Material examined. AUSTRALIA: NEW SOUTH WALES: S ledge, Cook
Is., 28°11.65'S 153°34.63'E, sand & shelly grit, 15 m, coll. K. Attwood, 9
Jun 1993,1 on SEM stub (AM W28224); E of North Head, Port Jackson,
33°48.77'S 151°20.98'E, sandy substratum, 60 m, coll. Australian Museum
party, 12 Apr 1989, 1 (AM W20442). WESTERN A USTRALIA: holotype (ZMH
V-7949), Shark Bay, Useless Inlet, 26°08'S 113°21'E, 7 m depth.

San Martín & Hutchings: Eusyllinae of Australia 313
Fig. 46. SEM of Paraehlersia ehlersiaeformis (Augener, 1913) (A) falciger, posterior parapodium; (B) dorsal simple chaeta; (C)
ventral simple chaeta. SEM of Pionosyllis mariae n.sp. (D) complete specimen; (E) anterior end, dorsal view; (F) detail of the same
(arrows showing tufts of cilia). A–C: AM W23926, W28221, W28899; D–F: AM W28873.
Description. Body broad anteriorly (Fig. 49D), tapering
posteriorly, yellowish, 7.2 mm long, 0.3 mm wide, with 61
chaetigers. Prostomium oval, nearly twice as wide as long;
4 eyes in open trapezoidal arrangement, and 2 anterior
eyespots; median antenna longer than combined length of
prostomium and palps, normally more than twice length,
inserted near line between posterior eyes; lateral antennae
about half of length of median antenna, inserted in front of
anterior eyes (Fig. 49E), behind eyespots (Fig. 47A). Palps
broad, slightly longer than prostomium, triangular.
Peristomium distinct, similar in length to following
segments; dorsal tentacular cirri longer than lateral antennae,
shorter than median antenna; ventral tentacular cirri about
two third length of dorsal tentacular cirri. Antennae,
tentacular, and anterior dorsal cirri, elongated, articulated
except basally (Figs 47A, 49E), with up to 20 articles; dorsal
cirri becoming shorter and smoother; posterior to
proventricle segments onwards, shorter than body width,
totally smooth. Nuchal organs as 2 ciliated grooves between
prostomium and peristomium (Fig. 49E,F). Dorsum of each
segment provided with 2 ciliary bands and minute pores
between bands (Fig. 47A,B). Subcirral papilla small,
inconspicuous, but present on anterior and midbody
segments (Figs 47B, 48B arrows, C). Parapodia conical,
slightly elongate, distally bilobed (Fig. 49B). Ventral cirri
digitiform, shorter than parapodial lobes. Most anterior

314 Records of the Australian Museum (2006) Vol. 58
Fig. 47. Paraehlersia weissmannioides (Augener, 1913) (A) anterior end, dorsal view; (B) anterior parapodium, lateral view (arrow
showing subcirral papilla); (C) compound chaetae, anterior parapodium; (D) aciculae, anterior parapodium; (E) spiniger-like compound
chaeta, midbody; (F) falcigers, midbody; (G) dorsal simple chaeta, midbody; (H) spiniger-like compound chaeta, posterior parapodium;
(I) falcigers, posterior parapodium; (J) dorsal simple chaeta, posterior parapodium; (K) ventral simple chaeta; (L) acicula, posterior
parapodium. AM W20442. Scales: A 0.18 mm, B 97.5 µm, C–L 20 µm.
parapodia with about 15 compound chaetae, with strongly
bidentate blades, both teeth well separated from each other,
distal tooth slender, proximal robust, and long, fine spines
on margin, 2–3 most distal ones longer than others, longer
than proximal teeth (Figs 48C, 43D,E). Progressively
posteriorly, most dorsal compound chaetae becoming longer
and slender, spiniger-like, blades 95 µm in length on
midbody, about 100 µm on posterior parapodia with fine,
moderate spines on margin, spiniger like chaetae absent
from last parapodia (Fig. 47E,H), indistinctly bidentate.
Remaining compound chaetae similar to those of anterior
segments (Figs 47F, 48F), progressively along body
becoming larger, with stronger proximal tooth, slightly
hooked (Figs 43A, 47I), reducing to 5. Dorsal simple chaetae

San Martín & Hutchings: Eusyllinae of Australia 315
Fig. 48. SEM of Paraehlersia weissmannioides (Augener, 1913) (A) midbody segments, dorsal view; (B) same (arrow showing
subcirral papilla); (C) subcirral papilla; (D) dorsal compound chaeta, anterior parapodium; (E) ventral compound chaeta, anterior
parapodium; (F) compound chaetae, midbody. AM W28224.
from midbody, slender, distally slightly truncate and bifid,
with short spines on margin (Figs 47G,J, 43B). Ventral
simple chaetae on posterior parapodia, thick, strongly
bidentate, with proximal tooth large, slightly hooked, and
distal tooth much shorter than proximal one, with about 4
long spines on margin, reaching level of proximal tooth (Figs
47K, 43C). Anterior parapodia with 2–3 slender aciculae, one
straight, others distally rounded (Fig. 47D); from proventricular
segments onwards, acicula solitary, with oblique, short tip (Fig.
47L). Pygidium with 2 long anal cirri and median papilla.
Pharynx everted on examined specimens, through about 7
segments; pharyngeal tooth anteriorly located, surrounded by
crown of 10 soft papillae (Fig. 47A). Proventricle rectangular,
through 4–5 segments, with about 22 muscle cell rows.
Remarks. The description of Ehlersia ferrugina by
Böggemann & Westheide (2004) from specimens collected
in coralline sand in Mahé (Seychelles) is similar to
Australian specimens of Paraehlersia weissmannioides and
they may represent the same species; but, the anterior
compound chaetae seem to have both teeth less well separated,
and the dorsal simple chaetae appear to be distally entire, not
truncate and bifid, as occurs in the Australian material.
Habitat. Occurring in sand and shelly grit, in shallow
depths, up to 15m.
Distribution. Australia (Western Australia, New South
Wales); ?Seychelles.

316 Records of the Australian Museum (2006) Vol. 58
Fig. 49. SEM of Pionosyllis yolandae n.sp. (A) compound chaetae, anterior parapodium; (B) detail of the same; (C) parapodial lobe
and acicula, midbody. SEM of Paraehlersia weissmannioides (Augener, 1913) (D) anterior part and midbody, dorsal view; (E)
prostomium and anteriormost segments; (F) detail of nuchal organs. A–C: AM W28876, D–F: AM W28224.
Genus Paraopisthosyllis Hartmann-Schröder, 1991
Paraopisthosyllis Hartmann-Schröder, 1991: 27.
Type species. Opisthosyllis brevicirra Hartmann-Schröder,
1979, designated by Hartmann-Schröder, 1991.
Diagnosis. Body robust, cylindrical, broad anteriorly,
tapered posteriorly, with many segments. Dorsal and ventral
surfaces covered with numerous, small papillae. Pro-
stomium with 4 lensed eyes and 3 antennae. Palps broad,
fused at base, ventrally folded. Peristomium shorter than
subsequent segments, sometimes covered dorsally by
chaetiger 1; 2 pairs of tentacular cirri. Nuchal organs as 2
ciliated grooves between prostomium and peristomium.
Dorsal and ventral cirri on all chaetigerous segments.
Antennae, tentacular, anal and dorsal cirri smooth, enlarged,
club-shaped to foliaceous. Dorsal cirri usually provided with
distinct cirrophores; on some species, dorsal cirri of anterior
segments alternating in size between large and small ones,
larger cirri arising more dorsally. Parapodia with compound,
heterogomph chaetae, and dorsal and ventral simple chaetae
on posterior parapodia. Pharynx wide, with pharyngeal tooth
inserted far from anterior rim. Proventricle wide,
voluminous. Pygidium small, with 2 anal cirri.
Remarks. According to Hartmann-Schröder (1991), this
genus belongs to the Eusyllinae. It has some unusual
morphological characters, however, such as the shape and

San Martín & Hutchings: Eusyllinae of Australia 317
arrangement of dorsal cirri, epidermal papillae, shape of
pharynx and proventricle; its method of reproduction is
unknown. It appears to be closely related to Rhopalosyllis,
and therefore should be considered as a member of the
subfamily Syllinae. Until its method of reproduction is
determined, we are retaining it within the Eusyllinae
according to its original designation. The genus is known
only from Australia.
Key to Australian species of Paraopisthosyllis
1 Dorsal cirri inflated, club-shaped (Fig. 50A) ................................................................................ 2
—— Dorsal cirri otherwise ..................................................................................................................... 3
2 Blades of some dorsal compound chaetae bidentate. Dorsal cirri
alternating in size, larger ones located more dorsally than others ................. P. alternocirra n.sp.
—— All blades unidentate. Dorsal cirri all similar in size except those
of chaetiger 1, inserted close to parapodial lobes (Fig. 53A) ..................................... P. brevicirra
3 Some dorsal cirri foliaceous (Fig. 55A). Pharyngeal tooth located
on anterior half of pharynx ........................................................................................ P. phyllocirra
—— Anterior dorsal cirri provided with distal, digitiform button (Fig.
54A). Pharyngeal tooth located on posterior half of pharynx.......................... P. ornaticirra n.sp.
Paraopisthosyllis alternocirra n.sp.
Figs 50A–E, 51A–H, 52A–I
Opisthosyllis brevicirra.—Hartmann-Schröder, 1982: 57 (in part).
Material examined. HOLOTYPE (AM W26734) AUSTRALIA:
WESTERN AUSTRALIA: Red Bluff, Kalbarri, 27°42'S 114°09'E, mixed
coralline algae, 4 m, coll. J.K. Lowry, 10 Jan 1984. PARATYPES Red
Bluff, Kalbarri, 27°42'S 114°09'E, round-leaved seagrass in shallow sand
on rocky shore, 3.5 m, coll. R.T. Springthorpe, 10 Jan 1984, 3 (AM
W28366); Rocky shore, Red Bluff, Kalbarri, 27°42'S 114°09'E,
dictyotalean alga from cave, 4 m, coll. J.K. Lowry, 10 Jan 1984, 4 (AM
W26784); Rocky shore, Red Bluff, Kalbarri, 27°42'S 114°09'E, brown
algae from surf zone, 0.5 m, coll. H.E. Stoddart, 9 Jan 1984, 1 (AM
W26783); Red Bluff, Kalbarri, 27°42'S 114°09'E, mixed coralline algae,
4 m, coll. J.K. Lowry, 10 Jan 1984, 7 + 1 on SEM stub (AM W28984);
Inshore limestone reef, Ned’s Camp, Cape Range National Park, 21°59'S
113°55'E, sponge covered with epiphytes, sediment & muddy worm
tubes, 1.5 m, coll. R.T. Springthorpe, 2 Jan 1984, 1 (AM W28368).
Additional material examined. WESTERN AUSTRALIA:
Rockingham, Point Peron, 32°17'S 115°44'E 115, algae, intertidal, coll.
G. Hartmann-Schröder, 1 (HMZ P-17049); identified by Hartmann-
Schröder as Opisthosyllis brevicirra.
Description. Complete specimen. Body anteriorly broad,
tapered posteriorly (Fig. 52A), 6.7 mm long, 0.5 mm wide,
with 59 chaetigers. Dark area of pigment dorsally on each
segment, sometimes divided into 2 dorsal areas, anterior
and posterior lateral areas; dark areas forming incomplete
transverse row on posterior segments (Fig. 50A–C). Small,
scattered papillae on lateral and ventral surfaces, more
numerous on dorsum (Figs 50A,C, 52C,D). Prostomium
oval, 4 small eyes arranged in open trapezoidal pattern,
almost in straight line; lateral antennae inserted near anterior
margin, oval, slightly rugose, shorter than combined length
of prostomium and palps, median antenna similar to lateral
but thicker, slightly longer, provided with dark inclusions,
inserted slightly posteriorly to lateral ones, in front of eyes
(Fig. 50A). Palps broad, longer than prostomium, ventrally
folded (Figs 50D,E, 52C). Peristomium shorter than
subsequent segments; tentacular cirri similar in shape to
lateral antennae, but larger. Dorsal cirri of chaetiger 1 distally
inflated, club-shaped, provided with dark inclusions, and
located laterodorsally, anteriorly directed, partially covering
prostomium (Fig. 50A,C,D); dorsal cirri of chaetigers 4 and
6 similar, but less inflated and more laterally inserted,
subsequent dorsal cirri club-shaped, not as inflated as those
of chaetigers 1, 4 and 6, without dark inclusions; alternating
dorsal cirri laterodorsally located and others slightly smaller,
more laterally located, cirri becoming smaller posteriorly
(Fig. 50C,D). Dorsal cirri with cirrophores. Anterior
parapodia with 7 compound, heterogomph chaetae, shafts
with some spines on distal margin, and curved blades; 2
dorsalmost compound chaetae with relatively short,
bidentate blades, with both teeth close to each other, and
short spines on margin (Fig. 51C), 16 µm long; remaining
compound chaetae with smooth, unidentate blades (Figs
51C, 52G), within fascicle slight dorsoventral gradation in
length of blade, 22 µm in length dorsally, 17 µm in length
ventrally. Progressively along body, chaetal blades
becoming shorter and less bidentate; posterior parapodia
with 6 compound chaetae, decreasing to 3 on posteriormost
chaetigers, blades slightly hooked, smooth on margin,
unidentate (Figs 51G, 52H), blades 15 µm in length dorsally,
10 µm in length ventrally. Dorsal simple chaetae slender,
unidentate, smooth (Fig. 51F), present on posterior
chaetigers (Fig. 52I). Ventral simple chaetae smooth, distally
bent, unidentate, present on most posterior parapodia (Figs
51H, 52I). Anterior parapodia with 3–4 aciculae, distally
rounded, single acicula on posterior parapodia (Fig. 51E).
Pharynx wide, through 3–4 segments; pharyngeal tooth
located about one third from anterior margin of pharynx
(Fig. 51A). Margin of pharynx with 10 papillae (Figs 51A,
52E), each papilla with short cilia (Fig. 52F). Proventricle,
longer than pharynx, extending through 3 segments, with
25–30 muscle cell rows. Pygidium small, with 2 anal cirri
with dark inclusions, inflated and oval (Fig. 50B).
Remarks. This species is similar to P. brevicirra Hartmann-
Schröder, 1979 (see below) but P. alternocirra has short
and long dorsal cirri alternating along the anterior part of
the body, distinct dark inclusions on the median antenna
and large dorsal cirri, a colour pattern, and some dorsal
compound chaetae on anterior and midbody parapodia with

318 Records of the Australian Museum (2006) Vol. 58
Fig. 50. Paraopisthosyllis alternocirra n.sp. (A) anterior end, dorsal view; (B) posterior end,
dorsal view; (C) midbody, lateral view; (D) prostomium and anteriormost segments, lateral view; (E)
same, ventral view. A,B,E: AM W26734; C,D: AM W26783. Scales: 0.18 mm
Fig. 51. Paraopisthosyllis alternocirra n.sp. (A) pharynx and proventricle; (B) aciculae, anterior
parapodium; (C) dorsalmost compound chaetae, anterior parapodium; (D) compound chaetae,
anterior parapodium; (E) acicula, posterior parapodium; (F) dorsal simple chaeta; (G) compound
chaetae, posterior parapodium; (H) ventral simple chaeta. AM W26734. Scales: A 0.18 mm, B–
H 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 319
Fig. 52. SEM of Paraopisthosyllis alternocirra n.sp. (A) complete specimen, lateral view; (B) anterior end, lateral
view; (C) prostomium and anterior segments, lateral view; (D) midbody, lateral view; (E) everted pharynx; (F) detail
of pharyngeal papillae; (G) fascicle of chaetae, anterior parapodium; (H) midbody fascicle of chaetae; (I) posterior
fascicle of chaetae. AM W28984.
bidentate blades. In contrast, P. brevicirra has only
unidentate chaetae, dorsal cirri not as inflated and lacks dark
inclusions on the antennae and cirri, and is colourless. The
specimen reported as Opisthosyllis brevicirra by Hartmann-
Schröder (1982) belongs to this new species and differs from
the holotype of the taxon.
Habitat. Occurring in algae, seagrasses, and sponges; from
intertidal to shallow depths.
Distribution. Australia (Western Australia).
Etymology. The specific name refers to the alternating sizes
of the dorsal cirri along the body.
Paraopisthosyllis brevicirra
(Hartmann-Schröder, 1979)
Fig. 53A–C
Opisthosyllis brevicirra Hartmann-Schröder, 1979: 57, figs 43–
45. Not Opisthosyllis brevicirra Hartmann-Schröder, 1982: 57.
Material examined. AUSTRALIA: WESTERN AUSTRALIA: holotype
(HZM P-15499), Port Hedland, close to Highway Motel, 19°38'S
119°31'E, filamentous algae with sand & debris, intertidal, 27 Sept. 1975,
coll. G. Hartmann-Schröder.
Description. Incomplete specimen, 4.5 mm long, 0.5 mm
wide, with 28 chaetigers, without pigment pattern (Fig.
53A). Numerous, scattered, rounded papillae covering

320 Records of the Australian Museum (2006) Vol. 58
Fig. 53. Paraopisthosyllis brevicirra (Hartmann-Schröder, 1979)
(A) anterior end, dorsal view; (B) midbody parapodium; (C)
compound chaetae. HZM P-15499. Not scaled. Modified from
Hartmann-Schröder, 1979.
dorsal and ventral surfaces, extending onto parapodia.
Prostomium oval, 4 eyes arranged in open trapezoidal
pattern; lateral antennae inserted near anterior margin, oval,
shorter than combined length of prostomium and palps,
median antenna similar to lateral but distinctly thicker,
slightly longer, inserted posteriorly to lateral ones (Fig.
53A). Palps broad, longer than prostomium, ventrally folded
(Fig. 53A). Peristomium shorter than subsequent segments;
tentacular cirri similar in shape to lateral antennae, but
longer. Dorsal cirri of chaetiger 1 distally inflated, club-
shaped, laterodorsally located (Fig. 53A). Remaining dorsal
cirri smaller, club-shaped, slightly elongate, not as inflated
as those of chaetiger 1, inserted more laterally (Fig. 53A,B).
Dorsal cirri with distinct cirrophores (Fig. 53B). Parapodia
conical, with 2 distal papillae (Fig. 53B). Anterior parapodia
with 8–9 compound, heterogomph chaetae, shafts with some
spines on distal margin, and curved, unidentate blades,
slightly shorter dorsally than ventrally, smooth on margin
(Fig. 53C). Dorsal and ventral simple chaetae not seen.
Anterior parapodia each with 3–4 aciculae, distally rounded,
reducing to single acicula on posterior parapodia. Pharynx
wide, extending through 3 segments; pharyngeal tooth
located about one third from anterior margin of pharynx
(Fig. 53A). Proventricle large, longer than pharynx, through
3 segments, with 25–26 muscle cell rows.
Remarks. One specimen 1 (HMZ P-17049) identified by
Hartmann-Schröder as Opisthosyllis brevicirra belongs to
P. alternocirra n.sp.
Habitat. Occurring in sand and debris, intertidally.
Distribution. Australia (Western Australia).
Paraopisthosyllis ornaticirra n.sp.
Fig. 54A–K
Material examined. HOLOTYPE (AM W28949) AUSTRALIA:
WESTERN AUSTRALIA: Inshore limestone reef, Ned’s Camp, Cape Range
National Park, 21°59'S 113°55'E, Caulerpa sp., 1 m, coll. J.K. Lowry, 2
Jan 1984.
Description. Complete specimen. Body broad and robust
anteriorly, tapered posteriorly, 7.8 mm long, 0.6 mm wide,
with 63 chaetigers. Segmented pigmentation absent, but
median antenna and anterior dorsal cirri each with 2 dark
transverse areas, large dorsal cirri with more extensive
pigmented area, anal cirri with single transverse band of
pigment (Fig. 54A,C,D). Dorsal and ventral surfaces densely
covered with small, rounded papillae (Fig. 54A).
Prostomium rectangular, with 2 pairs of eyes arranged
almost in line. Lateral antennae inserted in front of eyes,
relatively short and slender; median antenna longer and
thicker than lateral antennae, arising between eyes (Fig.
54A). Palps broad, as wide as prostomium, ventrally folded
(Fig. 54A,B). Peristomium dorsally reduced, covered by
chaetiger 1; dorsal tentacular cirri similar in shape to median
antenna slightly longer and thicker, ventral tentacular cirri
similar but smaller than dorsal ones. Dorsal cirri of
chaetigers 1, 4, and 6 distinctly longer and larger than
remaining dorsal cirri, inflated, except for chaetiger 6, with
digitiform, non-pigmented terminal lobe, arising dorso-
laterally (Fig. 54A). Remaining dorsal cirri with cirrophores;
cirrostyles tapered, fusiform, shorter than half of body
width, with 1–2 bands of dark pigment, midbody dorsal
cirri lacking such pigment (Fig. 54D); posterior dorsal cirri
oval, shorter than those of anterior chaetigers. Parapodia
with anterior and posterior lobes (Fig. 54D). Compound
chaetae with thick shafts, distally provided with strong
serration, and short, strongly bidentate blades, with short,
few spines on margin; anterior parapodia with about 15
compound chaetae, blades slender (Fig. 54F), about 13 µm
in length. Compound chaetae becoming progressively
thicker posteriorly, with blades bidentate; about 10–14
compound chaetae on midbody parapodia, blades 10–12
µm in length, terminal teeth increasing in size ventrally
within fascicle (Fig. 54G). Number of compound chaetae
per parapodium decreasing posteriorly, with only 3 on most
posterior parapodia, similar to those of midbody (Fig. 54I).
Dorsal simple chaetae on posterior parapodia, distally
serrated and bifid (Fig. 54J). Ventral simple chaetae on most
posterior parapodia, thick, strongly bidentate, smooth on
margin (Fig. 54K). Anterior parapodia with 3 aciculae,

San Martín & Hutchings: Eusyllinae of Australia 321
Fig. 54. Paraopisthosyllis ornaticirra n.sp. (A) anterior end, dorsal view; (B) prostomium and palps, ventral view; (C) posterior end,
dorsal view; (D) midbody parapodium; (E) anterior aciculae; (F) compound chaetae, anterior parapodium; (G) compound chaetae,
midbody; (H) acicula, posterior parapodium; (I) compound chaetae, posterior parapodium; (J) dorsal simple chaeta; (K) ventral simple
chaetae. AM W28949. Scales: A–C: 0.18 mm, D: 92 µm, E–K: 20 µm.
distally rounded (Fig. 54E), posteriorly single acicula
present, slightly bent at tip (Fig. 54H). Pharynx wide,
through 3–4 segments; pharyngeal tooth located posteriorly.
Proventricle similar in size to pharynx, with 30–35 muscle
cell rows. Pygidium trapezoidal; anal cirri larger than
posterior dorsal cirri, oval to egg-shaped.
Remarks. This species is easily distinguished from the other
members of this genus by the shape of the enlarged dorsal
cirri of chaetigers 1, 4, and 6, the pigment pattern present
on the dorsal cirri and median antenna, and the presence of
compound chaetae with strongly bidentate blades.
Habitat. Occurring in Caulerpa, in shallow depths.
Distribution. Australia (Western Australia).
Etymology. The species is named after the characteristic
pigmentation pattern of the dorsal cirri.

322 Records of the Australian Museum (2006) Vol. 58
Paraopisthosyllis phyllocirra
Hartmann-Schröder, 1991
Fig. 55A–F
Paraopisthosyllis phyllocirra Hartmann-Schröder, 1991: 27, figs
26–29.
Material examined. HOLOTYPE (HZM P-20533), AUSTRALIA:
QUEENSLAND: Heron Is., Great Barrier Reef, 23°27'S 151°55'E,
coralline sand, intertidal, coll. G. Hartmann-Schröder.
Description. Incomplete specimen, 3.5 mm long, with 32
chaetigers, with some indistinct red-brownish spots dorsally
and laterally. Numerous, scattered, rounded papillae
covering dorsal (Fig. 55A) and ventral surfaces. Prostomium
oval, 4 eyes arranged in open trapezoidal pattern; lateral
antennae inserted near anterior margin, oval, shorter than
combined length of prostomium and palps together, median
antenna similar to lateral, originating slightly posterior to
lateral antennae (Fig. 55A). Palps broad, longer than
prostomium, ventrally folded (Fig. 55A). Peristomium
slightly shorter than subsequent segments; tentacular cirri
similar in shape to lateral antennae, but larger. Dorsal cirri
of chaetiger 1 inflated, leaf-shaped, laterodorsally located
(Fig. 55A), right one missing. Following dorsal cirri smaller,
oval, not as inflated as those of chaetiger 1, arising more
laterally (Fig. 55A,B); some dorsal cirri of posterior
segments flattened, truncated, with some dark pigment (Fig.
55E), alternating with non modified dorsal cirri (Fig. 55F).
Parapodia sub-rectangular, with 4–5 distal papillae (Fig.
55B). Anterior parapodia with 7–9 compound, heterogomph
chaetae, shafts with some spines on distal margin, and
curved, bidentate blades, within fascicle dorsoventral
gradation in length of blades (Fig. 55C), with short spines
on margin. Posterior compound chaetae similar to anterior
ones, but with shorter blades and without any dorsoventral
gradation in length within fascicle (Fig. 55D). Dorsal and
ventral simple chaetae absent. Anterior parapodia with 3
aciculae, distally rounded, single acicula on posterior parapodia.
Fig. 55. Paraopisthosyllis phyllocirra Hartmann-Schröder, 1991 (A) anterior end, dorsolateral view; (B) anterior parapodium; (C) long
and short compound chaetae, anterior parapodium; (D) same, posterior parapodium; (E) modified dorsal cirrus, posterior parapodia;
(F) non-modified dorsal cirrus, posterior parapodium. HZM P-20533. Not scaled. Modified from Hartmann-Schröder, 1991.
Pharynx wide, through 3–4 segments; pharyngeal tooth occurs
about third from anterior margin of pharynx (Fig. 55A).
Proventricle large, longer than pharynx, through 3 segments,
with 25 muscle cell rows.
Remarks. Shape of posterior dorsal cirri were not described
in the original description.
Habitat. Occurring in coralline sand, intertidally.
Distribution. Australia (Queensland).
Genus Pionosyllis Malmgren, 1867
Pionosyllis Malmgren, 1867: 40.
Type species. Pionosyllis compacta Malmgren, 1867 by
monotypy.
Diagnosis. Body ranges from meiofaunal to macrofaunal
size (<5 to >10 mm in length), dorsally convex. Prostomium
with 4 eyes, sometimes also pair of eyespots, sometimes
without eyes. Three antennae. Prostomium sometimes with
cheeks or lobes on large specimens. Median antenna inserted
on middle of prostomium or anteriorly. Palps fused at bases,
with dorsal furrow, or free from each other. Two pairs of
tentacular cirri. Nuchal organs as 2 ciliated grooves between
prostomium and peristomium. Antennae, tentacular cirri and
dorsal cirri smooth, sometimes rugose, long, filiform, all
similar; weakly articulated on some species. Ventral cirri
not fused to parapodial lobes, those of anterior parapodia
somewhat inflated on some species, inserted basally or
distally on parapodial lobes. Heterogomph compound
chaetae with bidentate falcigerous blades, teeth similar in
size; some dorsal anterior and midbody falcigers slightly
longer than others, decreasing in size in dorsoventral
gradation within fascicle; some species with hemigomph
or homogomph articulations and blades of compound
chaetae different to those described above. Dorsal and
ventral simple chaetae on some posterior parapodia,
bidentate, with both teeth similar, sometimes absent. Parapodia

San Martín & Hutchings: Eusyllinae of Australia 323
with or without prechaetal lobes. Pharynx and proventricle
similar in size. Pharyngeal tooth anteriorly located, some
species with tooth on middle of pharynx. Reproduction by
epigamy. Pygidium small, with 2 smooth anal cirri.
Remarks. Pionosyllis as above defined is a heterogeneous,
probably a polyphyletic group in need of revision. Currently,
GSM and other authors are preparing a major revision of
the group.
Key to Australian species of Pionosyllis
1 Blades of compound chaetae with tendon connecting proximal
tooth with margin (Fig. 74D). Pharyngeal tooth located near
anterior rim, on middle of pharynx or posteriorly in mid line ...................................................... 2
—— Blades of compound chaetae without such tendon ....................................................................... 3
2 Pharyngeal tooth located just in front of middle of pharynx.
Compound chaetae including short, strongly bidentate falcigers
(Fig. 74C,D) .............................................................................................................. P. rousei n.sp.
—— Pharyngeal tooth located close to anterior margin of pharynx.
Compound chaetae all elongated, with proximal tooth distinctly
larger than distal tooth ............................................................................................. Pionosyllis sp.
3 Pharyngeal tooth located far from anterior rim (Fig. 65A). Some
dorsal cirri inflated ......................................................................................................................... 4
—— Pharyngeal tooth located on anterior rim or close to (Fig. 56A) .................................................. 5
4 Body dark. Inflated dorsal cirri with distal button (Fig. 60D).
Dorsoventral gradation in length of blades of compound chaetae ................................... P. fusigera
—— Body with small red spots. Inflated dorsal cirri without distal button
(Fig. 65B). Inverse dorsoventral gradation in length of blades of
compound chaetae........................................................................................................... P. kalimna
5 Ventral cirri inserted medially or distally on parapodial lobes.
Dorsal cirri of 2 lengths, long and filiform and extremely short,
exogonid-like dorsal cirri that alternate along body (Fig. 59A) ............................... P. corallicola
—— Ventral cirri inserted at bases of parapodia. Dorsal cirri similar
throughout or else differences between short and long cirri not so
pronounced (Fig. 78A) ................................................................................................................... 6
6 Segments posterior to proventricle fused in units of 2–3 segments.
Palps completely free ........................................................................................... P. yolandae n.sp.
—— Segments not fused. Palps fused at bases ...................................................................................... 7
7 Dorsal cirri (except some anteriormost) exogonid-like, short,
slightly longer than parapodial lobes ............................................................................................. 8
—— Dorsal cirri long ........................................................................................................................... 10
8 Dorsal cirri more or less truncate, with some internal glands (Fig.
70C) .......................................................................................................................... P. mariae n.sp.
—— Dorsal cirri otherwise, without internal glands (Fig. 56D) ........................................................... 9
9 Compound chaetae including spiniger-like, with elongated blades.
Dorsal simple chaetae truncate (Fig. 56I)................................................................ P. ancori n.sp.
—— Compound chaetae only falcigers. Dorsal simple chaetae pointed,
unidentate (Fig. 73G) ............................................................................................ P. mayteae n.sp.
10 Distinct prechaetal lobe present. Acicula straight, extending
beyond parapodial lobes (Fig. 67L). Blades of compound chaetae
without long, fine spines; without spiniger-like chaetae. Large size,
(>10 mm in length) .................................................................................................. P. kerguelensis
—— Prechaetal lobe absent. Acicula distally knobbed, with 2 unequal
lobes. Blades of compound chaetae with long, fine, distally
ornamented spines; sometimes with spiniger-like chaetae. 5–10
mm in length ................................................................................................................................ 11

324 Records of the Australian Museum (2006) Vol. 58
11 Without long, spiniger-like compound chaetae ........................................................................... 12
—— With long, thin, spiniger-like compound chaetae (Fig. 63D)...................................................... 13
12 Anteriormost parapodia distinctly enlarged, with compound
chaetae provided with large, thick shafts, and short blades (Fig.
75B) .................................................................................................................................. P. serrata
—— Parapodia and compound chaetae similar throughout ......................................... P. koolalya n.sp.
13 Anterior parapodia without elongated, spiniger like compound
chaetae ........................................................................................................ P. heterochaetosa n.sp.
—— Spiniger-like chaetae from anterior chaetigers ............................................................................ 14
14 Compound chaetae of anterior segments with coarse serration and
distal tooth more or less broad, rounded. Falcigers without double
curvature ........................................................................................................................ P. augeneri
—— Compound chaetae similar throughout. Some falcigers provided
with double curvature (Fig. 61C)............................................... P. hartmannschroederae n.comb.
Pionosyllis ancori n.sp.
Fig. 56A–M
Material examined
. HOLOTYPE (AM W29244). AUSTRALIA:
QUEENSLAND: Outer Yonge Reef, Great Barrier Reef, 14°36'S 145°38'E,
rock covered with pink coralline algae, encrusting sponges, 9 m, coll.
P.A. Hutchings, 21 Jan 1977. PARATYPES 4 (AM W28455), Outer Yonge
Reef, Great Barrier Reef, 14°36'S 145°38'E, rock with Lithothamnion
& Halimeda, 30 m, coll. P.A. Hutchings, 24 Jan 1977.
Additional material examined. QUEENSLAND: Outer Yonge Reef,
Great Barrier Reef: 14°36'S 145°38'E, rock with Lithothamnion &
Halimeda, 30 m, coll. P.A Hutchings, 24 Jan 1977, 4 (AM W28962).
NEW SOUTH WALES. Elisabeth & Middleton Reef, 33°16.85'S 19°9.15'E,
244 m, Franklin, May 1998; Elisabeth & Middleton Reef, 33°16.85'S
159°9.15'E, 30 m, Lithothamnion & Halimeda, 24 Jan. 77, P.A. Hutchings
coll. 8 (AM W28838); Taupo Seamount, 33°16'51"S 156°09'09"E,
limestone & sand bottom, 244 m, coll. J.K. Lowry, 2 May 1989, 2 (AM
W28839). WESTERN AUSTRALIA Kimberley, Lafontaine Is. 68, 14°10'S
125°47'E, 9–15 m, 19 July 1988, coll. P.A. Hutchings, 1 (AM W28841).
Description. Body slender, strongly filiform, 8 mm long,
0.2 mm wide, with 50 chaetigers. Prostomium sub-
pentagonal, without eyes, only 2 anterior eyespots (Fig.
56A); median antenna inserted near posterior margin of
prostomium, longer than combined length of prostomium
and palps; lateral antennae nearly one third length of median
antenna length, similar in length to palps, inserted
posteriorly to eyespots; 2 transverse furrows behind lateral
antennae (Fig. 56A). Palps long, triangular, free from each
other, fused basally, longer than prostomium. Peristomium
distinct, about half of length of subsequent segments; dorsal
tentacular cirri longer than lateral antennae but shorter than
median one, similar in length to body width; ventral
tentacular cirri about as long as dorsal ones (Fig. 56A).
Dorsal cirri, oval, papilliform, slightly longer than
parapodial lobes, without internal glands, distally tapered,
absent on chaetiger 2 (Fig. 56A). Parapodial glands with
granular material present on segments posterior to
proventricle (Fig. 56A). Parapodial lobes elongate, conical,
with distal papilla (Fig. 56D). Ventral cirri digitiform, shorter
than parapodial lobes. Compound chaetae hemigomph,
shafts smooth, and blades of 2 kinds; most dorsal compound
chaetae with elongate, bidentate blade, short spines on
margin, apparently joined by membrane on anterior and
midbody parapodia (Fig. 56E,G), about 33 µm in length on
anterior parapodia, 27 µm in length on midbody parapodia,
20 µm in length on posterior parapodia (Fig. 56J); remaining
chaetae with shorter blades, bidentate, with short spines on
margin, 15–8 µm on anterior parapodia, 10–8 µm on
midbody parapodia, about 10 µm on posterior parapodia, 3
on anterior and midbody parapodia (Fig. 56F,H), 2 on
posterior parapodia (Fig. 56K). Dorsal simple chaetae
truncated, with few spines on margin (Fig. 56I), present
from mid to posterior parapodia. Ventral simple chaetae on
posterior parapodia, acicular, bidentate with both teeth
hooked, proximal tooth larger than distal one, provided with
small translucent hood (Fig. 56L). Solitary acicula, with
oblique tip (Fig. 56M). Pharynx through 5 segments;
anterior end on chaetiger 1, with small pharyngeal tooth
located on anterior rim (Fig. 56A). Proventricle through
two and half segments, with 30 muscle cell rows. Pygidium
small, semi-circular, with 2 long, filiform anal cirri,
extending for 3 segments (Fig. 56C).
Remarks. Specimens from Western Australia and Elisabeth
and Middleton Reef have dorsal cirri of chaetiger 1 long
and 4 small eyes are visible (Fig. 56B) and have parapodial
glands larger than those present on the holotype. At this
stage we prefer to regard these as variations within
Pionosyllis ancori, as the eyes on some specimens have
become faded, and cirri can easily become detached or
damaged. Pionosyllis ancori n.sp., differs from the other
Australian species of the genus in having some compound
chaetae as long-bladed falcigers, long and simple dorsal
chaetae truncated, and having a thread-like body. The closest
species is Pionosyllis weismanni, which is a much more
robust species, with dorsal cirri on chaetiger 2, and glands
inside the dorsal cirri; the compound chaetae are similar in
both species, but the ventral simple chaetae are smaller than in
P. ancori, with the distal tooth longer and markedly curved.
Habitat. Occurring in encrusting algae and sponges, in
depths of 9 to 244 m.
Distribution. Australia (Queensland, Western Australia,
northern New South Wales).
Etymology. The species is named in honour to Ancor Núñez
Brito, son of our friend who recently died.

San Martín & Hutchings: Eusyllinae of Australia 325
Fig. 56. Pionosyllis ancori n.sp. (A) anterior end, dorsal view; (B) prostomium and anteriormost segments of one specimen with long
cirri on chaetiger 1; (C) posterior end, dorsal view; (D) midbody parapodium; (E) elongated, spiniger-like compound chaeta, anterior
parapodium; (F) falcigers, anterior parapodium; (G) elongated, spiniger-like compound chaeta, midbody; (H) falcigers, midbody; (I)
dorsal simple chaeta. (J) elongated compound chaeta, posterior parapodium; (K) falcigers, posterior parapodium; (L) ventral simple
chaeta; (M) acicula. Scales: A,C 68 µm, B 0.18 mm, D 37 µm, E–M 20 µm. A,C–M: AM W29244; B: AM W28838.

326 Records of the Australian Museum (2006) Vol. 58
Pionosyllis augeneri Hartmann-Schröder, 1979
Figs 57A–J, 58A–F
Pionosyllis augeneri Hartmann-Schröder, 1979: 98, figs 119–125;
1980: 52; 1981: 32, fig. 52. Non Hartmann-Schröder 1991: 35.
Material examined. HOLOTYPE (HMZ P-15474), PARATYPES (HMZ
P-15475, P-21018, P-16796) AUSTRALIA: WESTERN A USTRALIA: Broome,
17°58'S 122°14'E, mangrove, sand & detritus, intertidal. NEW SOUTH
WALES: Palm Beach, Pittwater, 33°35'S 151°19'E, Halophila &
Posidonia seagrass beds, sand, 3 m, coll. J.K. Lowry & R.T. Springthorpe,
28 Apr 1983, 3 (AM W28406) + 1 on SEM stub (AM W28898); Mid-
stream between Juno Head & Hungry Beach, Hawkesbury R., 33°34'S
151°16'E, muddy sand, 10 m, coll. A.R. Jones & C. Watson-Russell, 12
Jan 1977, 1 (AM W22130). SOUTH AUSTRALIA: 1 km NW of 5th Creek,
Fig. 57. Pionosyllis augeneri Hartmann-Schröder, 1979. (A) anterior end, dorsal
view (median and left antennae missing); (B) compound chaetae, anteriormost
parapodium; (C) spiniger-like compound chaeta, mid-anterior parapodium; (D)
compound chaetae, mid-anterior parapodium; (E) spiniger-like compound chaeta,
posterior parapodium; (F) compound chaetae, posterior parapodium; (G) dorsal
simple chaeta, midbody; (H) dorsal simple chaeta, posterior parapodium; (I) ventral
simple chaeta; (J) acicula, posterior parapodium. HZM P-15474. Scales: A 0.1 mm,
B–J 20 µm.
Port Pirie, Spencer Gulf, 33°12'S 137°55'E, 0.8 m, Zostera, T.J. Ward et
al. coll, Mar 1980, 1 (AM W22106), 1 (AM W22105); Boston Bay, Port
Lincoln, 34°51'S 135°51'E, washings from sheltered weedy rocks, 2 m,
coll. I. Loch, 12 Feb 1985, 1 on SEM stub (AM W28880). QUEENSLAND:
Halifax Bay, 19°10'S 146°44'E, 5 m, Queensland Nickel, Feb 1985,
Judell, Platt, Thomas & Assoc., 2 (AM W28206). Halifax Bay, 19°10'S
146°44'E, 5 m, Queensland Nickel, Feb 1985, Judell, Platt, Thomas &
Assoc., 2 (AM W28459). Triangular Islets, Shoalwater Bay, 22°23'
150°31'E, J.A. Lewis & J.R. Forsyth, 1981, 17 (AM W202630); Heron
Island, 23°27'S 151°55'E, sand, intertidal, (HMZ P-21018, P-16796).
Description. Body fragile, holotype 3.8 mm long with 36
chaetigers, longest specimen 4.9 mm long, with 52
chaetigers; normally colourless, but some with black spots.
Anterior part of body enlarged, after proventricle, body

San Martín & Hutchings: Eusyllinae of Australia 327
Fig. 58. SEM of Pionosyllis augeneri Hartmann-Schröder, 1979. (A) incomplete specimen, dorsal view; (B) anterior compound chaetae; (C)
long, spiniger-like compound chaetae, anterior parapodium; (D) falcigers, midbody; (E) compound chaetae, midbody; (F) falcigers with long
basal spines, posterior parapodium. AM W28898.
becoming thinner. Prostomium pentagonal, sometimes eyes
absent in fixed material, but normally 4 eyes in open
trapezoidal arrangement (Fig. 57A). Antennae cylindrical,
smooth, long; median antenna inserted between posterior
eyes, longer than combined length of prostomium and palps;
lateral antennae inserted close to anterior eyes, shorter than
median antenna. Palps longer than prostomium, triangular,
basally fused (Figs 57A, 58A). Peristomium similar in
length to subsequent segments; tentacular cirri similar to
antennae; usually dorsal tentacular cirri longer than ventral
tentacular cirri. Some dorsal cirri of anterior segments long,
slender, filiform, others slightly shorter than body width,
rest shorter than half of body width (Figs 57A, 58A); on
midbody dorsal cirri alternating long and short. Parapodia
of anterior segments large, subrectangular, becoming
slender from proventricle segments (Fig. 57A). Ventral cirri
digitiform, shorter than parapodial lobes. Anterior segments
with transverse row of long cilia; from proventricular
segments, 2 rows of dorsal cilia per segment (Fig. 57A).
Anterior parapodia with numerous compound chaetae,
about 15, with elongated blades, distally stout, with short
subdistal tooth, and coarse spines on margin (Figs 57B,
58B,C), spines slightly longer on ventral chaetae (Fig. 58D);
from proventricle posteriorly, some compound chaetae
becoming more elongated, spiniger-like, with filiform
blades, apparently unidentate (Figs 57C, 58E); other chaetae

328 Records of the Australian Museum (2006) Vol. 58
with blades falcigerous, distally rounded, with subdistal
tooth, and long spines distally directed (Fig. 57D).
Posteriorly, falcigers with shorter blades, with longer and
thinner spines on margin, especially basally (Fig. 57F), and
longer spiniger-like chaetae (Fig. 57E). Midbody and
posterior parapodia with 1 spiniger-like chaetae and 6
falcigers; posteriorly 1 spiniger-like and 4 falcigers (Figs
57F, 58F), some with long basal spines on margin. Dorsal
simple chaetae from midbody, distally provided with spines
of varying length (Fig. 57G), longer spines on posterior
parapodia (Fig. 57H). Ventral simple chaetae on posterior
parapodia, distally bidentate, both teeth similar and well
separated, with some long subdistal spines (Fig. 57I).
Fig. 59. Pionosyllis corallicola Ding &
Westheide, 1997. (A) anterior end, dorsal
view; (B) parapodial lobe; (C) dorsal simple
chaeta; (D) compound chaetae, midbody; (E)
acicula. AM W28944. Scales: A 0.68 µm, B
48 µm, C–E 20 µm.
Anterior parapodia with up to 4 aciculae, some distally
enlarged, numbers declining posteriorly to 1 slender, with
distal, rounded button (Fig. 57J). Pharynx through 6–7
segments; pharyngeal tooth anteriorly located, surrounded
by crown of 10 soft papillae (Fig. 57A). Proventricle shorter
than pharynx, through about 5 segments, with 20–23 muscle
cell rows.
Remarks. Material from Heron Island (HZM P-21018; P-
21017) includes 10 specimens of P. koolalya n.sp described
above and 2 specimens of P. hartmannschroederae.
Distribution. Australia (Western Australia, New South
Wales, Queensland), and possibly Caribbean.
Habitat. Occurring in coarse coralline sand, muddy sand, and
sediment in seagrasses beds, from intertidal to shallow depths.
Pionosyllis corallicola Ding & Westheide, 1997
Fig. 59A–E
Pionosyllis corallicola Ding & Westheide, 1997: 285, fig. 6.
Material examined. AUSTRALIA: WESTERN AUSTRALIA: 5
km offshore, Bush Bay, 30 km S of Carnarvon, 25°10'S
113°39'E, strap-leaved seagrass beds, 2 m, coll. J.K. Lowry
& R.T. Springthorpe, 6 Jan 1984, 1 (AM W28944).

San Martín & Hutchings: Eusyllinae of Australia 329
Description. Body 3.6 mm long, 0.2 mm wide, with 38
chaetigers. Prostomium semi-circular to pentagonal, with
2 tufts of cilia on each side, and 2 anterior eyespots (Fig.
59A). Antennae filiform, long and slender; median antenna
inserted near posterior margin of prostomium, more than 3
times combined length of prostomium and palps; lateral
antennae about half of length of median antenna, inserted
on anterior margin of prostomium, close to eyespots. Palps
triangular to oval, similar in length to prostomium (Fig.
59A). Peristomium similar in length to following segments;
nuchal organs distinct, as 2 ciliated grooves between
prostomium and peristomium; tentacular cirri similar to
antennae, dorsal ones longer than lateral antennae, shorter
than median one; ventral tentacular cirri about two thirds
length of dorsal ones. Tuft of cilia present on each side of
each segment (Fig. 59A). Dorsal cirri all smooth, absent on
chaetiger 2. Two kinds of dorsal cirri; long, filiform, similar
to antennae, distinctly longer than body width, with short
cirrophores, on chaetigers 1, 4, 6, 8 …, those of chaetiger 1
much longer than others, and others short, exogonid-like,
shorter than parapodial lobes, on chaetigers 3, 5, 7 … (Fig.
59A); although those on chaetiger 3 slightly longer than
other short ones. Parapodia with 2 lobes, prechaetal one
slightly bilobed (Fig. 59B). Ventral cirri distally inserted,
digitiform, extending beyond parapodial lobes (Fig. 59B).
Compound chaetae homogomph, with short, poorly
developed bidentate blades provided with thin spines on
margin (Fig. 59D), about 3 per parapodia, similar
throughout. Shafts thick with 3-dimensional articulation
(Fig. 59D). Dorsal simple chaetae unidentate, thick, with
few thin spines on margin (Fig. 59C). Ventral simple chaetae
absent. Acicula solitary, distally expanded laterally (Fig.
59E). Pharynx through 4 segments, pharyngeal tooth on
anterior rim, surrounded by crown of 10 soft papillae (Fig.
59A). Proventricle through 3 segments, with about 14
muscle cell rows. Pygidium semi-circular, with 2 filiform
anal cirri.
Habitat. Occurring interstitially in coralline sand.
Distribution. China (Hainan Is.), Australia (Western Australia).
Fig. 60. Pionosyllis fusigera Augener, 1913.
(A) anterior end, dorsal view; (B) anterior end,
ventral view; (C) anterior parapodium with
non-modified dorsal cirrus; (D) anterior
parapodium with inflated dorsal cirrus; (E)
compound chaetae, anterior parapodium;
(F, G ) aciculae on two different views; (H)
compound chaetae, midbody; (I) compound
chaetae, posterior parapodium. ZMH P-7951.
Scales: A,B 0.18 mm, C,D 48 µm, E–I 20 µm.

330 Records of the Australian Museum (2006) Vol. 58
Pionosyllis fusigera Augener, 1913
Fig. 60A–I
Pionosyllis fusigera Augener, 1913: 227, pl. III Fig. 34, text-fig.
32a–c.
Material examined. Syntype 1 (HZM V-7951) AUSTRALIA:
WESTERN AUSTRALIA: Sharks Bay, Surf Point, Outer Bar,
25°16'S 113°28'E, 1–3.5 m, 16 Jun 1905.
Description. Specimen in poor condition, covered with
crystals of formalin. Body fragile, without colour pattern,
but dark, cylindrical, flattened ventrally, convex dorsally, 4
mm long, 0.5 mm wide for 38 chaetigers, incomplete.
According to Augener (1913), dorsal cirri, prostomium and
palps pigmented brown and anterior segments with brown
dorsal bands. Prostomium sub-quadrangular with rounded
corners; 4 eyes almost equal in size, arranged in open
trapezoidal pattern; 2 nuchal lobes, kidney shaped, located
behind posterior pair of eyes (Fig. 60A). All antennae
located on anterior margin of prostomium, short, similar in
shape and size, slightly enlarged distally, as long as
prostomium; lateral antennae thinner than median one (Fig.
60A). Palps broad, large, fused basally, widely separated
distally, folded ventrally (Fig. 60B). Peristomial ring shorter
than following segments, dorsally covered between first
chaetiger and prostomium and visible only laterally and
ventrally, with two pairs of tentacular cirri. Dorsal tentacular
cirri and some dorsal cirri similar in shape, smooth, thick,
club-shaped, basally slender and gradually becoming wider
distally, terminating in globular papillae; most anterior cirri
slightly shorter than middle and posterior cirri (Fig. 60B);
ventral tentacular cirri similar in shape and size to lateral
antennae. Long, club-shaped dorsal cirri alternating with
shorter, thinner, digitiform cirri, as long as half of body
width (Fig. 60A). Ventral cirri conical to digitiform, distally
pointed, almost as long as parapodia. Parapodia with
prechaetal lobe larger than postchaetal one (Fig. 60C,D).
Compound chaetae heterogomph falcigers; shafts with distal
short spines; blade tips bidentate, distal tooth hooked,
proximal tooth similar in size to distal one (Fig. 60E,H,I).
About 12 compound chaetae on anterior parapodium,
number progressively decreasing posteriorly; dorsalmost
compound chaetae on each parapodium with elongate,
slender, minutely bidentate blade, with numerous fine, short
spines on edge, 60 µm in length on anterior parapodia,
shorter, about 36 µm in length, posteriorly. Blades of
remaining chaetae more strongly bidentate, all similar in
shape with slight dorsoventral and anteroposterior gradation
in size, blades progressively becoming shorter and wider
posteriorly, about 54 µm in length dorsally and 28 µm in
length ventrally on anterior parapodia; blades on posterior
parapodia about 31 µm in length dorsally and 20 µm in
length ventrally (Fig. 60H,I). Anterior parapodia with 2
straight, thick, acuminate, distally pointed aciculae;
posterior parapodia with single acicula (Fig. 60F,G).
Pharynx, removed from specimen, reaching segment 4,
pharyngeal tooth on segment 2 (fide Augener, 1913).
Proventricle barrel shaped, extending through 3 chaetigers
(from 4 to 7) (Fig. 60A) with about 18 muscle cell rows.
Habitat. Occurring in depths of 1 to 3.5m.
Distribution. Australia (Western Australia).
Pionosyllis hartmannschroederae n.nom.
Fig. 61A–F
Typosyllis (Langerhansia) longisetosa Hartmann-Schröder, 1990:
49, figs 9–12.
Pionosyllis augeneri.—Hartmann-Schröder, 1991: 35. Not
Hartmann-Schröder, 1979: 98.
Material examined. AUSTRALIA: NEW SOUTH WALES: Angourie
Point, 29°29'S 153°22'E, algal beds, intertidal, holotype of Typosyllis
(Langerhansia) longisetosa, (HZM P-19661), paratype (HZM P-19962).
WESTERN AUSTRALIA: Goss Passage, Beacon Is., 28°25.5'S 113°47'E,
dead coral substrate, in fine sediment at foot of reef slope, P.A Hutchings,
23 May 1994, 4 (AM W28386); NE entrance to Goss Passage, Beacon
Is, 28°27.9'S 113°46.7'E, terebellids on rocks in coral sand at foot of
slope, 33 m, P.A. Hutchings, 25 May 1994. 1 (AM W26455); inshore
limestone reef, Ned’s Camp, Cape Range National Park, 21°59'S, fine
sediment & sand from patches between reefs, H.E. Stoddart, 2 Jan 1984,
4 (AM W26785); 5 km offshore Bush Bay, 30 km S of Carnarvon 25°10'S
113°55'E, shallow strap-leaved seagrass beds, 2 m, coll. J.K. Lowry &
R.T. Springthorpe, 6 Jan 1984, 1 (AM W28926); Descartes Is.,
Kimberleys, 14°11'S 125°40'E, sandflats & mangroves, coll. P.A.
Hutchings, 20 July 1988, 1 (AM W28397).
Description. Body 1.4 mm long, 0.2 mm wide, with 53
chaetigers. Prostomium oval to circular, with 4 eyes in open
trapezoidal arrangement (Fig. 61A), absent in some
specimens (probably lost after fixation); median antenna
about two and half times longer than combined length of
prostomium and palps, inserted between posterior eyes;
lateral antennae slightly shorter than median antenna,
inserted near anterior margin. Palps broad, longer than
prostomium (Fig. 61A). Peristomium shorter than following
segments; dorsal tentacular cirri elongated, filiform, similar
to median antenna, ventral tentacular cirri smaller. Dorsal
cirri elongate, slender, smooth, filiform, varying in length
in different specimens and in each specimen, some cirri
long, several times longer than body width, and some cirri
shorter than body width, alternating irregularly (Fig. 61A).
Anterior parapodia broad, becoming conical and slender
from proventricle segments onwards. Ventral cirri
digitiform, shorter than parapodial lobes anteriorly,
elongated on posterior parapodia, longer than parapodial
lobes. Compound chaetae similar throughout, including 1–
2 long, slender, filiform spiniger-like chaetae, 180 µm in
length, 1–2 falcigers with bidentate blades, provided with
long spines on margin, especially basally, and distal spines
extending beyond level of distal tooth, straight margin of
blade with convex curvature proximally then concave
subdistally (Fig. 61C), 45–47 µm in length, and about 5
chaetae with bidentae blades, with long distal spines, and
short basal spines (Fig. 61D) and dorsoventral gradation in
length within fascicle, 40 µm in length dorsally, 25 µm in
length ventrally. Dorsal simple chaetae from proventricle
segments, with distinct, short spines on margin (Fig. 61E).
Ventral simple chaetae on posterior segments, distinctly
bidentate, proximal tooth prominent, slightly longer than
distal tooth, and some thin spines on margin, distal one
longer than others, extending to level of distal tooth.
Aciculae distally knobbed, with 2 lateral, unequal lobes (Fig.
61F); 2 aciculae on anterior parapodia. Pharynx slender,
through about 6–7 segments (Fig. 61A). Proventricle
through 4–6 segments, with about 24 muscle cell rows.
Pygidium small, with 2 filiform anal cirri.
Remarks. Hartmann-Schröder described two different
species named longisetosa in two different genera, on the

San Martín & Hutchings: Eusyllinae of Australia 331
Fig. 61. Pionosyllis hartmannschroederae n.nom. (A) anterior end, dorsal view;
(B) spiniger-like compound chaeta, midbody; (C) compound chaetae with
double curvature and long basal spines, midbody; (D) compound chaetae
without double curvature and moderate basal spines, midbody; (E) dorsal
simple chaeta; (F) acicula. AM 28926. Scales: A 0.18 mm, B–F 40 µm.
basis of our study of these two species we regard them as
congeneric and therefore transfer Typosyllis (Langerhansia)
longisetosa into the genus Pionosyllis. Unfortunately this
renders Typosyllis (Langerhansia) longisetosa Hartmann-
Schröder, 1990 as a junior homonym of Pionosyllis
longisetosa (Hartmann-Schröder, 1965). We therefore propose
a new name for Typosyllis (Langerhansia) longisetosa
Hartmann-Schröder 1990 of P. hartmannschroederae.
Habitat. Occurring on sandflats in mangroves, dead coral
substrate, in fine sediment at foot of reef slope, in strap-
leaved seagrass beds, sediments trapped in algae, intertidally
and in shallow waters.
Distribution. Australia (Queensland, Western Australia).
Pionosyllis heterochaetosa n.sp.
Figs 62D–F, 63A–O, 64A–F
Material examined. HOLOTYPE (AM W21629) AUSTRALIA: NEW
SOUTH WALES: S of airport runway extension, Botany Bay, 33°58.13'S
151°11.16'E, muddy sand, 5 m, coll. Australian Museum party, 6 Apr
1992. PARATYPES S of airport runway extension, Botany Bay, 33°58.13'S
151°11.16'E, muddy sand, 5 m, Australian Museum party, 27 July 1992,
3 on SEM stub (AM W22117); S of airport runway extension, Botany
Bay, 33°58.10' 151°11.16'E, muddy sand, 5 m, coll. Australian Museum
party, 27 July 1992, 2 (AM W22110); S of airport runway extension,
Botany Bay, 33°58.13'S 151°11.16'E, muddy sand, 5 m, Australian
Museum party, 6 Apr 1992, 2 (AM W22112); S of airport runway
extension, Botany Bay, 33°58.13'S 151°11.16'E, muddy sand, 5 m, coll.
Australian Museum party, 27 July 1992, 1 on SEM stub (AM W22116). S
of airport runway extension, Botany Bay, 33°58.13'S 151°11.16'E, muddy
sand, 5 m, coll. Australian Museum party, 27 July 1992, 2 (AM W22115).
Description. Body 4.6 mm long, 0.2 mm wide, with 54
chaetigers. Prostomium circular, with 4 eyes in open
trapezoidal arrangement (Fig. 63A); median antenna long,
about 3 times longer than combined length of prostomium
and palps, inserted between posterior eyes; lateral antennae

332 Records of the Australian Museum (2006) Vol. 58
Fig. 62. SEM of Pionosyllis koolalya n.sp. (A) compound chaeta, midbody; (B) dorsal simple chaeta; (C) ventral simple chaeta and falciger,
posterior parapodium. SEM of Pionosyllis heterochaetosa n.sp. (D) anterior end, dorsal view; (E) compound chaetae, anterior parapodium; (F)
fascicles of chaetae, midbody. A–C: paratype, AM W28409; D–F: paratype, AM W22117.
about half or third length of median antenna, inserted near
anterior margin. Palps broad, longer than prostomium (Figs
62D, 63A). Peristomium shorter than subsequent segments;
dorsal tentacular cirri elongated, filiform, similar to median
antenna, ventral tentacular cirri smaller. Dorsum of anterior
segments each with transverse band of cilia (Figs 62D, 63A);
double row from proventricle segments onwards. Dorsal
cirri elongated, slender, smooth, filiform, varying in length
in different specimens and in each specimen, some cirri
long, several times longer than body width, and some cirri
shorter than body width, alternating irregularly (Figs 62D,
63A). Anterior parapodia broad, becoming conical and
slender from proventricle segments posteriorly. Ventral cirri
digitiform, shorter than parapodial lobes anteriorly,
elongated on posterior parapodia, longer than parapodial
lobes. Most anterior parapodia with numerous compound
chaetae, about 10–15, slightly enlarged shafts, and short,
bidentate blades, provided with long spines on margin (Figs
62D,E, 63B), especially on more dorsal chaetae, extending
beyond distal tooth, and dorsoventral gradation in length
within fascicle, about 16 µm in length dorsally, 5–6 µm in
length ventrally. Progressively along body, number of
compound chaetae per parapodium decreasing, and blades
of more dorsal compound chaetae becoming elongate and
more strongly bidentate (Fig. 63C); on chaetiger 5, blades
about 22 µm in length dorsally, 12 µm in length ventrally.
From proventricular segments, parapodia with 2–3,
sometimes only 1, compound chaetae with elongate,

San Martín & Hutchings: Eusyllinae of Australia 333
Fig. 63. Pionosyllis heterochaetosa n.sp. (A) anterior end, dorsal view; (B) compound chaetae, anteriormost parapodium; (C) compound chaetae,
anterior parapodium; (D) spiniger-like compound chaeta, mid-anterior parapodium; (E) compound chaetae, mid-anterior parapodium; (F) spiniger-
like compound chaeta, midbody; (G) compound chaetae, midbody; (H) spiniger-like compound chaeta, posterior parapodium; (I) compound chaetae,
posterior parapodium; (J) dorsal simple chaeta, anterior parapodium; (K) same, midbody; (L) same, posterior parapodium; (M) ventral simple
chaeta; (N) aciculae, anterior parapodium; (O) acicula, posterior parapodium. AM W21629 (holotype). Scales: A 0.18 mm, B–O 20 µm.
spiniger-like blades (Fig. 62F), about 112 µm in length,
weakly bidentate distally, with moderate spines on margin
(Fig. 63D,F,H), and 5–6 compound chaetae with falcigerous,
bidentate blades, slender, margin with long, distinct, distally
pointed basal spines and thin, fine, spines distally (Figs 62F,
63E,G,I, 64B–D), reaching level of distal tooth, and
dorsoventral gradation in length within fascicle, 25 µm
dorsally, 15 µm ventrally. Dorsal simple chaetae from
proventricle segments, distally broad, provided with short
spines on margin (Figs 63J–L, 64E). Ventral simple chaetae
on posterior segments, bidentate, proximal tooth prominent,
slightly longer than distal tooth, and some thin spines on
margin, 2–3 distal ones longer than others, reaching to level
of distal tooth (Figs 63M, 64F). Aciculae distally knobbed,
with 2 lateral, unequal lobes (Fig. 63 O); 2 aciculae on
anterior parapodia (Fig. 63N). Pharynx slender, through

334 Records of the Australian Museum (2006) Vol. 58
Fig. 64. SEM of Pionosyllis heterochaetosa n.sp. (A) chaetae, midbody; (B–D) compound chaetae, midbody; (E) dorsal simple
chaeta; (F) ventral simple chaeta. AM W22116, AM W22117.
about 6–7 segments (Fig. 63A). Proventricle through 6
segments, with about 24 muscle cell rows. Pygidium small,
with 2 filiform anal cirri.
Remarks. Pionosyllis heterochaetosa n.sp. is characterized
by having anterior parapodia without spiniger-like
compound chaetae, which appear in post- proventricle
segments. Pionosyllis serrata and P. koolalya, described
above, also lack spiniger-like chaetae anteriorly, and
posteriorly. All other species of the genus (P. hartmann-
schroederae and P. augeneri described below, as well as P.
spinisetosa San Martín, 1990, from Cuba, and P. anoph-
thalma Capaccioni & San Martín, 1989, from the western
Mediterranean Sea), have spiniger-like compound chaetae
from chaetiger 1 (see San Martín, 1990, 2003), as well as P.
longisetosa (Hartmann-Schröder, 1965) from Chile (see
Hartmann-Schröder, 1965).
Distribution. Australia (New South Wales).
Habitat. Occurring in muddy sand at about 5 m depth.
Etymology. The specific name refers to the presence of
different chaetae on anterior segments compared to others.

San Martín & Hutchings: Eusyllinae of Australia 335
Pionosyllis kalimna n.sp.
Fig. 65A–K
Material examined. HOLOTYPE (AM W28948) AUSTRALIA:
WESTERN AUSTRALIA: N end of beach, Bundegi Reef, Exmouth Gulf,
21°49'S 114°11'E, rocky rubble, sediment & brown algae with epiphytes,
2 m, coll. H.E. Stoddart, 4 Jan 1984. PARATYPES 2 (AM W26779) N end
of beach, Bundegi Reef, Exmouth Gulf, 21°49'S 114°11'E, rocky rubble,
coralline algae with green epiphyte, 2 m, coll. H.E. Stoddart, 4 Jan 1984.
Description. Body 6.4 mm long, 0.3 mm wide, slightly
broad anteriorly, tapering posteriorly, with 39 chaetigers
plus 5–6 segments without chaetae (Fig. 65A,G), darkly
pigmented prostomium and peristomium, anterior segments
covered in numerous red pigment spots (Fig. 65A).
Prostomium oval to rectangular, with 4 eyes in open
trapezoidal arrangement and 2 anterior eyespots; median
antenna inserted between anterior eyes, on middle of
prostomium, fusiform, slightly longer than combined length
of prostomium and palps; lateral antennae similar to median
antenna but less fusiform, inserted in front of anterior margin
of prostomium, ventral in relation to median antenna,
slightly longer than palps. Palps triangular, divergent, free
for most of length, fused basally, slightly shorter than
prostomium (Fig. 65A). Peristomium similar in length to
subsequent segments, with 2 nuchal organs forming ciliated
dorsal pits; dorsal tentacular cirri similar in shape and length
to median antenna, ventral tentacular cirri similar to lateral
ones. Median antenna, dorsal tentacular cirri, and dorsal
cirri of chaetigers 1, 4, and 6, enlarged, fusiform, distinctly
longer than remaining appendages (Fig. 65A); remaining
dorsal cirri not as fusiform, and shorter, about two thirds
length of longer ones. Parapodia with bilobed prechaetal
lobe and trilobed postchaetal lobe, medium lobe enlarged
at base with rounded tip (Fig. 65B). Ventral cirri digitiform,
shorter than parapodial lobes. Compound chaetae
heterogomph, with smooth or nearly smooth shafts
Fig. 65. Pionosyllis kalimna n.sp. (A) anterior end, dorsal view; (B) midbody parapodium; (C)
compound chaetae, anterior parapodium; (D) aciculae, anterior parapodium; (E) compound
chaetae, midbody; (F) acicula, midbody; (G) posterior end, dorsal view; (H) dorsal simple
chaeta; (I) compound chaetae, posterior parapodium; (J) ventral simple chaeta; (K) acicula,
posterior parapodium. AM W28948. Scales: A,G 0.18 mm; B 48 µm; C–F, H–K 20 µm.

336 Records of the Australian Museum (2006) Vol. 58
anteriorly becoming distally serrated on
midbody and posterior segments and
bidentate, short blades, becoming more
bidentate ventrally within fascicle; and
progressively along body (Fig. 65C,E,I).
Compound chaetae of anterior parapodia (Fig.
65C) all similar, about 7 per parapodium, blades
relatively slender with short proximal tooth and
smooth margin or provided with minute spines,
about 10–11 µm in length. Number of
compound chaetae increasing along
body, up to 9 present on posterior
parapodia; within fascicle, blades
with inverse dorsoventral gradation
in length, terminal teeth becoming
larger and with thicker shafts (Fig.
65E,I), on midbody, blades 6 µm
in length dorsally and 9 µm in
length ventrally within fascicle; on
posterior parapodia blades bidentate
and 6 µm in length dorsally and 11
µm in length ventrally within fascicle.
Anterior parapodia with 2 acuminate
aciculae (Fig. 65D), 1 from midbody
(Fig. 65F,K). Dorsal simple chaetae from
mid-posterior parapodia, slender, distally
bifid, with short spines on margin (Fig.
65H). Ventral simple chaetae on posterior
parapodia, thick, smooth, bidentate, both teeth
well separated (Fig. 65J). Pharynx through 4–5
segments; pharyngeal tooth inserted posteriorly
to mid pharynx (Fig. 65A), on chaetiger 3.
Proventricle through 2 segments, large, with about 30
muscle cell rows, difficult to count. Pygidium semi-circular,
with 2 oval, slightly enlarged, anal cirri, slightly longer than
pygidium (Fig. 65G).
Remarks. Pionosyllis kalimna n.sp. is easily distinguished
from P. fusigera Augener, 1913, by the compound chaetae
that are short with two equally well-developed terminal
teeth, enlarged dorsal cirri without a distal button; it also
differs in the structure of the parapodia, and the colour
pattern, black stripes in P. fusigera, and small red spots in
P. kalimna.
Habitat. Occurring in rocky rubble, sediment and brown
algae with epiphytes, in 2 m depth.
Distribution. Australia (Western Australia).
Etymology. The specific name comes from an aboriginal
word, kalimna meaning beautiful.
Fig. 66. Pionosyllis kerguelensis (McIntosh, 1885). (A) prostomium with
everted pharynx, dorsal view; (B) anterior end, dorsal view; (C) anterior
parapodium, posterior view; (D) anterior parapodium, posterior view; (E)
posterior parapodium, posterior view (dorsal cirrus not drawn); (F) chaetigers
1–8, ventral view, right side. A,D: BMNH 1885.12.1.142; B,C,E,F: MNHN
A-921. Scales: A,B 0.56 mm; C,D 0.1 mm; E 0.07 mm; F 0.8

San Martín & Hutchings: Eusyllinae of Australia 337
Pionosyllis kerguelensis (McIntosh, 1885)
Figs 66A–F, 67A–L
Eusyllis kerguelensis McIntosh, 1885: pl. 29, fig. 4, pl. 33 fig. 3,
Pl. 15a fig. 13. Ehlers, 1897: 42; 1913: 473.—Augener, 1924:
376; 1927: 152.—Monro, 1930: 94, fig. 30a-c; 1936: 130.—
Knox, 1951: 73; 1960: 105.—Knox & Cameron, 1998: 49,
figs 99–100.—Hartman, 1953: 20; 1964: 81, pl. 25, figs 2–
3.—Wesenberg-Lund, 1961: 59, fig. 19.—Hartmann-Schröder,
1965: 115, figs 74; 1986: 77.—Averincev, 1982: 19, pl. II,
Figs 1–2.—Hartmann-Schröder & Rosenfeldt, 1988: 39, fig.
16; 1990: 97; 1992: 98.
Pionosyllis comosa Gravier, 1906: 288.—Ehlers, 1913: 473, pl.32,
Figs 1–4.—Benham, 1927: 60.—Monro, 1930: 92; 1936: 128,
fig. 20.—Hartman, 1953: 23; 1964: 85, pl. 26 figs 7–8; 1967: 58.
Pionosyllis cosma [sic] Knox, 1960: 106.—Knox & Cameron
1998: 51.
Pionosyllis kerguelensis San Martín & Parapar, 1997: 291; San
Martín, 2004: 16.
Material examined. KERGUELEN IS. off Christmas harbour, 49°30'S
69°30'W, 200 m, coll. H.M.S. Challenger, presented McIntosh, syntype
of Eusyllis kerguelensis, BMNH 1885.12.1.142 (1 ant. end + 2 median
parts). FALKLAND ISLANDS: Port Albemarle, 52°11'S 60°26'W, 40 m,
sand & algae, coll. Swedish Antarctic Exp. 1901–1903, 8 Sept. 1902,
Pionosyllis comosa, 2 (SMNH 3332, 3728); Burwood bank, 53°45'S
61°10'W, 137–150 m, shell fragments & stones, coll. Swedish Antarctic
Exp. 1901–1903, 12 Sept. 1902, P. comosa 1 (SMNH 3660). ARGENTINA:
coast of northern Argentina: 37°15'S 56°8'W, 100 m, sand & gravel,
coll. Swedish Antarctic Exped. 1901–1903, 23 Dec 1901, P. comosa 1
(SMNH 3826). SOUTH GEORGIA: south fjord in front of Nordenskjöld
glacier, 54°24'S 36°22'W, 210 m, blue grey mud with few small stones,
coll. Swedish Antarctic Exp. 1901–1903, 29 May 1902, Pionosyllis
epipharynx 1 (SMNH 3033); French Antarctic Exp. Charcot: 1910, P.
comosa 1 (MNHN A-76), 1906 (P. comosa). CROZET ISLANDS: 46°27'S
52°00'E, St. 12, DC 12, P. comosa, (MNHN A-921). SOUTH SHETLAND
ISLANDS: Spanish cruise Bentart 94, coll. Hespérides, St. 52, mixed
sediment, 62°43'S 60°27'W, 84 m, 1 (MNCNM 1521); St. 71, rocks,
62°43'S 60°26'W, 50 m, 1 (MNCNM 1523); St. 77, mud & organic rests,
62°40'S 60°40'W, 130 m, 1 (MNCNM 1522). TASMAN SEA: 42°17.99'S
170°00.00'E, 958 m depth, 1 (SMF 13225).
Description. Body dorsally arched, ventrally flattened, with
well defined intersegmental furrows, segments short (Fig.
66B); all specimens fragmented or lacking posterior end;
largest specimen examined (SMNH 3033) 29 mm long, 2.5
Fig. 67. Pionosyllis kerguelensis (McIntosh, 1885) (A–C) dorsal,
median and ventral compound chaetae, anteriormost parapodium;
(D) dorsalmost compound chaeta, anterior parapodium; (E,F)
dorsal and ventral compound chaetae of remaining chaetae, mid-
anterior parapodium; (G) dorsalmost compound chaeta, mid-
posterior parapodium; (H,I) dorsal and ventral compound chaetae
of remaining chaetae, mid-posterior parapodium; (J,K) dorsal and
ventral compound chaetae, posterior parapodium; (L) aciculae.
A–F: BMNH 1885.12.1.142; G–L: MNHN A-76. Scales: 14 µm.

338 Records of the Australian Museum (2006) Vol. 58
mm wide for 34 chaetigers; without colour markings
(preserved specimens). Prostomium with irregular surface,
with longitudinal and transverse grooves forming two
posterior cheeks (Fig. 66A,B); 4 eyes arranged in trapezoidal
pattern, anterior pair subequal or smaller than posterior one,
located on cheeks and often covered by peristomium and
chaetiger 1. Palps broad, long, divergent, distally rounded,
slightly longer than prostomium, basally fused, with dorsal
furrow (Fig. 66A,B). Antennae detached or broken in most
specimens; much longer than prostomium (according to
Hartman, 1964); median antenna more than twice as long
as lateral ones (according to Gravier, 1906), originating just
in front of median cleft, lateral antennae inserted on anterior
margin of prostomium. Peristomium short, dorsally visible,
with 2 pairs of tentacular cirri; dorsal tentacular cirri much
longer than ventral ones, similar in shape to antennae and
dorsal cirri. Long dorsal cirri smooth basally, weakly
crenulated distally, about 4 times as long as body width;
short dorsal cirri slightly longer than body width, alternating
irregularly with long cirri (Fig. 66B–D). Ventral cirri broad,
thick, inflated, almost as large as parapodial lobes on
anterior chaetigers (Fig. 66C,D), except those of chaetigers
1–2 (Fig. 66F), shorter and thinner posteriorly (Fig. 66E).
Parapodial lobes with long digitiform, dorsal prechaetal
papilla (Figs 66C–E). Anterior parapodia with more than
40 compound chaetae, decreasing to about 10 on posterior
parapodia. Compound chaetae heterogomph falcigers; shafts
increasing in width posteriorly along body and dorso-
ventrally within fascicle, with spinose endings. Blades
bidentate, with short, distally directed spines on margin;
with marked dorsoventral and anteroposterior gradation in
both shape and size; dorsal blades on anterior and midbody
parapodia long, finely spinulated on margin, distal tooth
slightly hooked, proximal tooth slightly smaller, ventral
blades basally wider, distal tooth hooked and slightly larger
than proximal one (Fig. 67A–C); posterior segments with
1–2 dorsal chaetae with elongate, slender blades (Fig. 67D)
and remaining chaetae with dorsoventral gradation (Fig.
67E,F); in midbody, this arrangement more marked (Fig.
67G–I); on posterior fascicles, most dorsal chaetae with short
blades (Fig. 67J), blades of remaining falcigers similar in length
to dorsal ones, but wider, proximal tooth long, triangular (Fig.
67K). Blades of anterior compound chaetae about 100 µm in
length dorsally, 40 µm in length ventrally; posterior blades 55
µm in length dorsally, 35 µm in length ventrally. Anterior
parapodia with 4–5 straight, thick, aciculae, pointed tip
protruding beyond parapodial lobes; posterior parapodia with
1–2 aciculae (Fig. 67L). Dorsal and ventral simple chaetae
not seen. Pharynx wide, everted in some specimens,
extending through 8–9 chaetigers, with 2 crowns of papillae
surrounding opening; pharyngeal tooth conical, located on
anterior dorsal rim (Fig. 67A,B). Proventricle barrel shaped,
extending to chaetiger 17–18, slightly longer than pharynx.
Habitat. Occurring in fine grey sandy mud, volcanic mud,
hard bottom, sand and shells, intertidally to 2916 m (Ehlers,
1913).
Distribution. Coast of North Argentina, Antarctic and
Subantarctic seas: Kerguelen Is. South Georgia, Falkland
Is., Magellan area, Davis Sea, Palmer Archipelago, Drake
Passage, South Orkney Is. Antarctic Peninsula, South
Shetland, Australia (Tasmania, New South Wales), New
Zealand, Crozet Is.
Pionosyllis koolalya n.sp.
Figs 62A–C, 68A–P, 69A–F
Pionosyllis augeneri.—Hartmann-Schröder, 1991: 359–125. Not
Hartmann-Schröder, 1979: 98.
Material examined. HOLOTYPE (AM W28945) AUSTRALIA: SOUTH
AUSTRALIA: Speeds Point, Streaky Bay, 32°48'S 134°13'E, coll. P.A.
Hutchings, 14 Mar 1979. PARATYPE 1 on SEM stub (AM W28409). NEW
SOUTH WALES: Bottle & Glass Rocks, Port Jackson, 33°51'S 151°16'E,
substrate unknown, 12m, coll. G. Clark, 11 Dec 1989.
Additional material examined. QUEENSLAND: Heron Is., 23°27'S
151°55'E, sand, intertidal, coll. G. Hartmann-Schröder, id. as Pionosyllis
augeneri (HMZ P-21018, P-16796).
Description. Holotype 12.8 mm long, 0.3 mm wide, with 71
chaetigers; paratype much smaller. Prostomium oval, with 4
eyes arranged in open trapezoidal pattern (Fig. 68A). Median
antenna long, coiled on holotype, several times longer than
combined length of prostomium and palps, inserted between
posterior eyes; lateral antennae distinctly shorter than median
antenna, inserted near anterior margin of prostomium (Figs
68A, 69A–C). Palps broad, similar in length to prostomium or
slightly longer. Tentacular and dorsal cirri long, smooth, coiled
on holotype, long on anterior segments, becoming shorter on
segments beyond proventricle, alternating long cirri (Figs 68C,
69A,B), 2–3 times as long as body width, and short cirri (Fig.
68B), slightly shorter than body width; dorsal cirri with
cirrophores. Ciliary bands on dorsum of each segment and
areas of dorsal cirri (Fig. 68B–D), distinct on holotype (Fig.
68A); anterior segments with single ciliary band (Fig. 69B,C),
double from proventricle segments posteriorly (Fig. 69D).
Parapodial lobes conical, ending with 2 lobes (Fig. 68B,D).
Ventral cirri digitiform, shorter than parapodial lobes anteriorly,
becoming longer posteriorly (Fig. 68B,D). Anterior parapodia
with about 10–14 compound chaetae, slightly enlarged, distally
truncated shafts, provided with short subdistal spines, and
bidentate blades, both teeth similar, with long spines on margin,
pointing distally, those of basal part coarse, distal ones thin,
reaching or extending beyond proximal tooth, 1–3 slightly
longer (Fig. 68E), rest of chaetae within fascicle with slight
dorsoventral gradation in length (Fig. 68F), 26 µm in length
dorsally, 18 µm in length ventrally. Posteriorly, number of
compound chaetae per parapodium decreasing, and blades of
1–3 most dorsal compound chaetae becoming proportionally
longer (Figs 68H, 69F) and rest with dorsoventral gradation in
length (Figs 68I, 69E); midbody parapodia with 1–2 compound
chaetae with relatively long blades, about 31 µm in length,
and 4 compound chaetae 15–25 µm in length; proximal tooth
proportionally longer than those of anterior chaetae (Fig. 62A).
Posterior parapodia with 1 compound chaeta with long blades
(Fig. 68K), 33 µm long, and 3 compound chaetae, similar
to those of midbody, but with distinctly larger proximal tooth
(Figs 68L, 62C), well separated from distal tooth, 18–22
µm in length. Dorsal simple chaetae from midbody, distally
bifid to truncate (Figs 68G, 62B), thicker posteriorly, with
short, coarse spines on margin (Fig. 68 O ). Ventral simple
chaetae on posterior parapodia, sigmoid, thick, strongly
bidentate, both teeth well separated, almost at 90°, proximal
tooth long, broad, slightly hooked, with few, long subdistal
spines, surpassing level of proximal tooth (Figs 68P, 62C).
Two aciculae on anterior segments (Fig. 68M), 1 from
proventricle segments onwards, distally bilobed (Fig. 68J),
thicker on posterior parapodia (Fig. 68N). Pharynx through

San Martín & Hutchings: Eusyllinae of Australia 339
Fig. 68. Pionosyllis koolalya n.sp. (A) anterior end, dorsal view; (B) midbody parapodium with short dorsal cirri; (C) long dorsal cirri,
midbody; (D) posterior parapodium; (E) long-bladed compound chaeta, anterior parapodium; (F) compound chaetae, anterior parapodium;
(G) dorsal simple chaeta, midbody; (H) long-bladed compound chaeta, midbody; (I) compound chaetae, midbody; (J) acicula, midbody;
(K) long-bladed compound chaeta, posterior parapodium; (L) dorsal and ventral compound chaetae, posterior parapodium; (M) aciculae,
anterior parapodium; (N) acicula, posterior parapodium; (O) dorsal simple chaeta, posterior parapodium; (P) ventral simple chaeta.
AM W28409. Scales: A 0.18 mm, B–D 97 µm, E–P 20 µm.
about 6 segments, pharyngeal tooth located on anterior rim.
Proventricle through 4 segments, with about 26 muscle cell
rows. Pygidium small, with 2 anal cirri.
Remarks. Pionosyllis koolalya is similar to P. serrata
described above but differs mainly in lacking enlarged
anterior parapodia with thick compound chaetae with
markedly short blades; remaining compound chaetae are
rather similar, but the dorsal simple chaeta of P. koolalya is
truncated, and slightly bifid in P. serrata. All other species
of the genus are provided with long, spiniger-like chaetae.
These two species also have fewer compound chaetae with

340 Records of the Australian Museum (2006) Vol. 58
Fig. 69. SEM of Pionosyllis koolalya n.sp. (A) incomplete specimen, dorsal view; (B) anterior end, dorsal view; (C) detail of same;
(D) detail of dorsum, midbody; (E) compound chaetae, midbody; (F) long-bladed compound chaeta, and one short-bladed, compound
chaeta. Paratype, AM W28409.
distinctly longer blades than other chaetae in the fascicle,
but they are relatively short and less than half the length of
the longest bladed chaetae. Some material of P. augeneri
(Hartmann-Schröder, 1991) from Heron Island, Queensland,
belong to this new species.
Habitat. Unknown, occurring from intertidal to 12 m.
Distribution. Australia (South Australia, New South Wales,
Queensland).
Etymology. The specific name comes from an aboriginal
word, koolalya, meaning feather epaulettes, in reference to
the ciliated dorsum.
Pionosyllis mariae n.sp.
Figs 46D–F, 70A–J, 71A–F, 72A–C
Material examined. HOLOTYPE (AM W28454) AUSTRALIA:
QUEENSLAND: Outer Yonge Reef, Great Barrier Reef, 14°36'S 145°38'E,
rock with Lithothamnion & Halimeda, 30 m, coll. P.A. Hutchings, 24
Jan 1977. PARATYPE (AM W28921) Outer Yonge Reef, Great Barrier
Reef, 14°36'S 145°38'E, rock covered with pink coralline algae,
encrusting sponges, 9 m, coll. P.A. Hutchings, 21 Jan 1977.
Other material examined. NEW SOUTH W ALES: Taupo Seamount,
Tasman Sea, 33°16.85'S 156°09.15'E, limestone & sand bottom, 244 m,
coll. J.K. Lowry & party on RV Franklin, 2 May 1989, 1 on SEM stub
(AM W28873).

San Martín & Hutchings: Eusyllinae of Australia 341
Fig. 70. Pionosyllis mariae n.sp. (A) anterior end, dorsal view; (B) posterior end,
dorsal view; (C) midbody parapodium; (D) elongate, short spiniger-like, compound
chaeta, anterior parapodium; (E) falcigers, anterior parapodium; (F) anterior
aciculae; (G) dorsal simple chaeta; (H) falcigers, posterior parapodium; (I) elongate,
short spiniger-like compound chaeta, posterior parapodium; (J) ventral simple
chaeta. AM W28454. Scales: A,B 0.18 mm; C 92 µm; D–J 20 µm.
Description. Body slender, filiform (Fig. 46D), 7 mm long,
0.3 mm wide, 54 chaetigers. Prostomium pentagonal to
triangular, with 4 eyes in open trapezoidal arrangement, and
sometimes 2 anterior eyespots. After fixation, however, eyes
may be absent (Fig. 70A). Lateral antennae inserted between
anterior eyes and eyespots, near anterior margin of
prostomium, smooth, similar in length to palps; median
antenna arising between posterior eyes, long, often curled,
distally pseudoarticulated, broken on holotype (Fig. 70A),
more than twice length of prostomium. Palps longer than
prostomium, triangular, directed behind, fused basally. Two
transverse, small, ciliated furrows on prostomium (Figs 46F
arrow, 70A). Peristomium distinct, slightly shorter than
subsequent segments; tentacular cirri elongated, dorsal
tentacular cirri longer than lateral antennae, shorter than
median one, ventral tentacular cirri shorter than dorsal ones
(Figs 46E,F, 70A). Dorsal cirri of chaetiger 1 long, slender,
similar to median antenna, sometimes pseudoarticulated
distally. Dorsal cirri short, slightly longer than parapodial
lobes, fusiform (Fig. 71A,B), distally more or less truncated
(Fig. 70A,C), usually with some fibrillar inclusions, with
terminal pore (Fig. 71C arrow). Dorsal cirri absent on
chaetiger 2, tuft of cilia present (Fig. 46E,F arrow).
Parapodia conical, with distal papilla (Figs 70C, 71A).

342 Records of the Australian Museum (2006) Vol. 58
Fig. 71. SEM of Pionosyllis mariae n.sp. (A) midbody, lateral view; (B) the same of an epigamic specimen, showing the capillary
notochaetae; (C) dorsal pores and distal pore of dorsal cirrus (arrows); (D) detail of dorsal pores (arrow); (E) chaetal fascicle, mid-
posterior parapodium; (F) short, spiniger-like chaeta. AM W28873.
Ventral cirri digitiform, slender, slightly longer than
parapodial lobes on posterior parapodia. Compound chaetae
with hemigomph articulation, smooth distally on shafts;
blades of 2 kinds, most dorsal chaetae with elongated, short
spiniger-like, bidentate blades, spines on margin apparently
jointed by membrane (Figs 70D,I, 71F), remaining chaetae
with short, bidentate blades, with short spines on margin
(Figs 70E, 71E), more strongly bidentate on posterior
parapodia (Figs 70H, 72A). Anterior parapodia each with 1
compound chaetae with elongated blade, about 27 µm in
length, and 4 chaetae with shorter blades, 17–12 µm in
length; posterior parapodia with 1 compound chaetae with
elongate blade and 2–3 short-bladed, all similar in size to
anterior ones. Dorsal simple chaetae from midbody, smooth,
distally truncate (Fig. 72B), with small, indistinct hood (Fig.
70G). Ventral simple chaetae from mid-posterior parapodia,
large, acicular, prominent, with subdistal translucent hood,
bidentate, proximal tooth long, hooked, distal tooth smaller,
also hooked (Figs 70J, 71E, 72C). Anterior parapodia each
with 2 aciculae, one straight, other bent at tip (Fig. 70F);
solitary acicula on remaining parapodia, bent tip, thicker
than anterior ones. Dark glands on each parapodia from
midbody (Fig. 70B) with dorsal pores (Fig. 71C,D arrows).
Pharynx through about 4 segments; pharyngeal tooth
anteriorly located (Fig. 70A). Proventricle rectangular,
through 3–4 segments, with 30–36 muscle cell rows.

San Martín & Hutchings: Eusyllinae of Australia 343
Fig. 72. SEM of Pionosyllis mariae n.sp. (A) falciger, posterior parapodium; (B) dorsal simple chaeta; (C) ventral simple chaeta.
SEM of Pionosyllis serrata (Hartmann-Schröder, 1984) (D) complete specimen, dorsal view; (E) prostomium and palps, dorsal
view; (F) anterior end, dorsal view. A–C: AM W28873, D–F: AM W28913.
Pygidium semi-circular, with 2 long, slender anal cirri (Fig.
70B), similar in length to median antenna. Mature specimen
with notoacicula and natatory chaetae from chaetiger 17.
One specimen epigamic, with natatory chaetae on midbody
and posterior segments (Fig. 71B).
Remarks. Pionosyllis mariae n.sp. is similar to P. weismanni
Langerhans, 1879, a worldwide reported species, that
probably represents a complex of morphologically similar
species (San Martín, 2003), since small differences among
specimens from different parts of the world have been
reported (see Ben-Eliahu, 1977). The Australian specimens
are distinctly smaller than those of the Mediterranean Sea
(7 mm versus 18 mm) and lack dorsal cirri on chaetiger 2,
which are always present on Mediterranean specimens (see
San Martín, 2003).
Habitat. Occurring as cryptic species in encrusting
communities and in sand, in depths of 9 to 244 m.
Distribution. Australia (Queensland, New South Wales).
Etymology. The new species is named after María Capa,
collaborator and friend.

344 Records of the Australian Museum (2006) Vol. 58
Pionosyllis mayteae n.sp.
Fig. 73A–K
Material examined. HOLOTYPE (AM W29245) AUSTRALIA:
QUEENSLAND: Outer Yonge Reef, Great Barrier Reef, 14°36'S 145°38'E,
rock with Lithothamnion & Halimeda, 30 m, coll. P.A. Hutchings, 24
Jan 1977. PARATYPES 5 (AM W28963) Outer Yonge Reef, Great Barrier
Reef, 14°36'S 145°38'E, rock with Lithothamnion & Halimeda, 30 m,
coll. P.A. Hutchings, 24 Jan 1977.
Description. Body slender, less filiform than other species
of the genus above described, 5.4 mm long, 0.2 mm wide,
with 44 chaetigers. Prostomium semi-circular to pentagonal,
apparently without eyes. Median antenna elongate, about
twice combined length of prostomium and palps, rugose to
indistinctly articulated, inserted near posterior margin of
prostomium; lateral antennae smooth, about half length of
median antenna, arising laterally in front (Fig. 73A). Palps
Fig. 73. Pionosyllis mayteae n.sp. (A) anterior end, dorsal view; (B) parapodium,
midbody; (C) dorsal and ventral compound chaetae, anterior parapodium; (D) compound
chaetae, midbody; (E) acicula, midbody; (F) posterior end, dorsal view; (G) dorsal simple
chaeta; (H) compound chaetae, posterior parapodium; (I) ventral simple chaeta; (J)
acicula, posterior parapodium; (K) acicula, anterior parapodium. AM W28963. Scales:
A,F 68 µm; B 48 µm; C–E, G–K 20 µm.
short, broad, stout, shorter than prostomium. Peristomium
similar in length to subsequent segments; nuchal organs as
2 distinct ciliated grooves between prostomium and
peristomium; dorsal tentacular cirri slightly shorter than
lateral antennae, ventral tentacular cirri about two thirds
length of dorsal ones. Dorsal cirri of chaetiger 1 elongate,
smooth, longer than body width, slightly shorter than
median antenna; dorsal cirri of subsequent segments
fusiform, slightly enlarged basally, slightly longer than
parapodial lobes, absent on chaetiger 2 (Fig. 73A).
Parapodial lobes conical, relatively short, stout. Ventral cirri
digitiform, similar in length to parapodial lobes or slightly
longer (Fig. 73B). Large parapodial glands from midbody,
with granular, dark material, additional small glands with
hyaline material present posteriorly on some specimens (Fig.
73B,F). Compound chaetae falcigers, with relatively short,
bidentate blades (Fig. 73C,D,H), hemigomph anteriorly
becoming heterogomph posteriorly, with smooth shafts,
with distinct subdistal spine; blades more strongly bidentate
posteriorly, with short spines on margin or smooth. Anterior
and midbody parapodia with 5 compound chaetae, blades
16 µm in length dorsally, 8 µm in length ventrally; 4
compound chaetae on each posterior parapodia, blades 7–
10 µm in length. Dorsal simple chaetae from mid-posterior
segments, long, straight, acute, distally pointed, with minute
subdistal spines on margin (Fig. 73G). Ventral simple
chaetae on posterior parapodia, thick, strongly bidentate,
teeth at 90°, proximal tooth slightly hooked, with small
translucent hood (Fig. 73I). Acicula solitary throughout,
slender, slightly larger posteriorly, distally expanded
laterally with convex surface (Fig. 73E,J). Pygidium semi-
circular, with 2 long anal cirri, smooth, similar in length to
dorsal cirri of chaetiger 1 (Fig. 73F). Pharynx through 5
segments; pharyngeal tooth on anterior rim (Fig. 73A).
Proventricle through 5 segments, with 20 muscle cell rows.
Remarks. Pionosyllis mayteae n.sp. differs from similar
other species of the genus in lacking compound chaetae

San Martín & Hutchings: Eusyllinae of Australia 345
Fig. 74. Pionosyllis rousei n.sp. (A) anterior end, dorsal view; (B) spiniger-like compound chaeta, anterior parapodium; (C) spiniger-
like compound chaetae, midbody; (D) falcigers, midbody; (E) aciculae, anterior parapodium; (F) aciculae, midbody. AM W28840
(holotype). Scales: A 0.18 mm; B–F 20 µm.
with elongated blades and having distally pointed dorsal simple
chaetae; and ventral simple chaetae bidentate with teeth at 90°.
Habitat. Occurring on encrusting Lithothamnion and
Halimeda at depths of 30 m.
Distribution. Australia (Queensland).
Etymology. The species is named after María Teresa
Aguado (Mayte), collaborator and friend.
Pionosyllis rousei n.sp.
Fig. 74A–F
Material examined. HOLOTYPE (AM W29230), TASMAN SEA: reef
flat near Yoshin Maru Iwaki wreck, Elisabeth Reef, 29°55.48'S
159°01.18'E, Elisabeth & Middleton Reefs Expedition, St. 43, 14 Dec
1987. PARATYPES 4 (AM W28840), reef flat near Yoshin Maru Iwaki
wreck, Elisabeth Reef, TASMAN SEA, 29°55.48'S 159°01.18'E, Elisabeth
& Middleton Reefs Expedition, St. 43, 14 Dec 1987.

346 Records of the Australian Museum (2006) Vol. 58
Additional material examined. TASMAN SEA: Taupo Seamount,
33°16.51'S 156°09.09'E, limestone & sand bottom, 244 m, coll. J.K.
Lowry, on RV Franklin, 2 May 1989, 3 (AM W28836).
Description. Holotype best preserved individual, incom-
plete specimen, 2.2 mm long, 0.6 mm wide, with 22
chaetigers. Body fragile, dorsally arched, cylindrical,
without colour pattern, yellowish in alcohol. Prostomium
oval, with 4 eyes in open trapezoidal arrangement. Antennae
damaged; lateral antennae inserted near anterior margin of
prostomium, median antenna inserted on middle of
prostomium. Palps large, slightly longer than prostomium,
free from each other (Fig. 74A). Nuchal organs ciliated,
extending to lateral margins of prostomium (Fig. 74A).
Peristomium dorsally reduced, covered by chaetiger 1.
Anteriormost segments short, becoming longer from
proventricle onwards. Left dorsal tentacular cirrus missing,
right one appears broken; ventral tentacular cirri shorter
than dorsal cirri. Dorsal cirri smooth, with distinct
cirrophores, elongated, alternating irregularly long and short
(Fig. 74A), long cirri similar in length to body width, shorter
cirri less than half length of longer ones. Parapodial lobes
conical. Ventral cirri elongated, similar in length to
parapodial lobes. Compound chaetae heterogomph, blades
of two kinds, spiniger-like and falcigers. Midbody parapodia
with 2 spiniger-like chaetae, with short spines on margin,
distinctly bidentate, distal tooth hooked, and proximal tooth
curved, with tendon contacting with margin, about 40–35
µm long (Fig. 74C), and 8–11 falcigers, strongly bidentate,
long and hooked distal tooth and long, curved proximal
tooth, provided with distinct tendon contacting with edge
of blade, slight dorsoventral gradation in length of blades
within fascicle, 25 µm dorsally and 15 µm ventrally (Fig.
74D), with thin spines on margin. Anteriormost parapodia
with 1–2 distinctly long, slender, spiniger-like chaetae
present, blades about 86 µm in length, bidentate, with both
blades similar in size, proximal tooth slightly recurved, but
without tendon (Fig. 74B). Dorsal and ventral simple
chaetae not seen. Aciculae tricuspid, 3 on anterior parapodia
(Fig. 74E), 2 on midbody parapodia (Fig. 74F). Pharynx
wide, though about 7–8 segments; pharyngeal tooth conical,
located in front of middle of pharynx (Fig. 74A).
Proventricle through 7 segments, similar in size to pharynx,
with about 28 muscle cell rows.
Remarks. Pionosyllis rousei n.sp. is characterized by having
compound chaetae distinctly bidentate, with both teeth
similar and well separated from each other. Pionosyllis
serratisetosa (López, San Martín & Jiménez, 1997), from
southwestern Mediterranean has similar chaetae, but the
teeth on the blades are not separated and are smaller, the
blades have a different shape; long spiniger-like compound
chaetae are absent, and the pharyngeal tooth is located near
anterior margin of pharynx. Pionosyllis longocirrata (Saint-
Joseph, 1887), from the eastern Atlantic and Mediterranean
Sea, also has the pharyngeal tooth located more anteriorly
than P. rousei, and the blades of compound chaetae either
have a minute distal tooth or none (see San Martín, 2003,
for more details). Pionosyllis templadoi (San Martín, 1991)
from Cuba has a similar body and the position of pharyngeal
tooth is also similar. The blades of compound chaetae,
however, have a small distal tooth and an elongated proximal
tooth (San Martín, 1991). Pionosyllis luquei (San Martín,
1990), also from Cuba, has similar compound chaetae,
although not as strongly hooked, with well-developed teeth,
but the dorsal cirri are shorter, and the pharyngeal tooth is
located more posteriorly (San Martín, 1990).
Habitat. Substrate unknown, recorded from 244 m depth.
Distribution. Tasman Sea (Elisabeth and Middleton Reef).
Etymology. This species is named after Dr Greg Rouse, an
Australian Polychaetologist.
Pionosyllis serrata (Hartmann-Schröder, 1984)
Figs 72D–F, 75A–N, 76A–F, 77A–F
Eusyllis serrata Hartmann-Schröder, 1984: 18, figs 23–26.
Material examined. AUSTRALIA: NEW SOUTH WALES: East of Bondi,
33°53.1'S 151°17.4'E, sand, 30 m, Fisheries Research Institute, 24 July
1989, 2 (AM W24373); Bass Point, 34°36'S 150°54'E, 50 m, coll. The
Ecology Lab, Ready Mixed Industries, 1 Feb 1990, 3 (AM W28922);
Bass Point, 34°36'S 150°54'E, 50 m, coll. The Ecology Lab, Ready Mixed
Industries, 1 Feb 1990, 2 on SEM stub (AM W28913).
Description. Body fragile (Fig. 72D), 8.7 mm long, 0.4
mm wide, with 77 chaetigers, pale in alcohol. Prostomium
oval, slightly wider than long, with 4 eyes in open
trapezoidal arrangement (Fig. 75A); median antenna more
than twice as long as prostomium and palps together,
inserted between posterior eyes, near posterior margin of
prostomium; lateral antennae shorter than median antenna,
similar or slightly longer than combined length of
prostomium and palps, inserted near anterior margin; two
tufts of cilia posterior to insertion of lateral antennae (Fig.
72E). Palps long and basally broad, triangular, longer than
prostomium (Figs 72E, 75A), free for almost entire length,
basally fused. Peristomium dorsally reduced. Anterior
segments with single transverse row of cilia (Figs 72F, 75A);
midbody and posterior segments each with double row of
cilia (Fig. 76A,B). Antennae, tentacular and dorsal cirri long,
slender, filiform, smooth, sometimes coiled, of 2 sizes, long
and short, long ones distinctly longer than body width, short
ones similar in length to half of body width (Figs 72D,F,
75A, 76A,B). Dorsal tentacular cirri long, ventral ones short.
Parapodia of anterior 8–10 chaetigers enlarged, wide,
distally truncated (Fig. 75A) with several small, rounded
lobes distally (Fig. 75M), and numerous, about 12–14 short,
broad compound chaetae, with enlarged shafts and short,
bidentate blades (Fig. 76D), about 22–16 µm in length, with
few, long, thick spines on margin, distal ones extending
beyond proximal tooth (Figs 75B, 76E). Posterior to
proventricle, parapodial lobes becoming conical, slender,
with terminal papilla; compound chaetae from proventricle
parapodia without enlarged shafts, apparently smooth and
distally truncated, and elongated blades with long, thin
spines on margin, distally ornamented, distal ones slightly
longer than others, extending beyond proximal tooth and
sometimes reaching level of distal tooth; most dorsal
compound chaetae with more elongate, bidentate blades
(Figs 75C,F,I, 76F, 77A), about 40 µm in length on post-
proventricle segments, 35 µm on midbody parapodia, 42
µm on posterior parapodia, and several compound chaetae
with shorter blades, and dorsoventral gradation in length
within fascicle (Figs 75D,G,J,K, 76F, 77B,C), 13–26 µm
on proventricle segments, 16–25 µm on midbody parapodia,
16–37 µm on posterior parapodia; compound chaetae 8–10

San Martín & Hutchings: Eusyllinae of Australia 347
Fig. 75. Pionosyllis serrata (Hartmann-Schröder, 1984); (A) anterior end, dorsal view; (B) compound chaetae, anteriormost parapodium;
(C) dorsalmost compound chaeta, anterior parapodium; (D) compound chaetae, anterior parapodium; (E) dorsal simple chaeta, midbody;
(F) dorsalmost compound chaeta, midbody; (G) compound chaetae, midbody; (H) dorsal simple chaeta, posterior parapodium; (I) dorsalmost
compound chaeta, posterior parapodium; (J,K) compound chaetae, posterior parapodium; (L) ventral simple chaeta; (M) aciculae and edge of
anteriormost parapodium; (N) acicula and edge of posterior parapodium. AM W28922. Scales: A 0.18 mm, B–N 20 µm.
on proventricle segments, about 7 on midbody, 4 on
posterior parapodia. Blades of posterior chaetae markedly
bidentate, both teeth well separated, almost at 90°, and
slightly convex (Figs 75J,K, 77D). Dorsal simple chaetae
from midbody, distally weakly bifid (Fig. 75E), with some
spines on margin, becoming thicker posteriorly, with shorter
spines on margin (Figs 75H, 77E). Ventral simple chaetae
on posterior parapodia, thick, strongly bidentate, both teeth
almost at 90°, proximal tooth slightly hooked, and few, long
spines on margin, extending beyond proximal tooth (Figs

348 Records of the Australian Museum (2006) Vol. 58
Fig. 76. SEM of Pionosyllis serrata (Hartmann-Schröder, 1984) (A) midbody, dorsal view; (B) detail of the same; (C) everted
pharynx; (D) anteriormost compound chaetae; (E) compound chaeta, anteriormost parapodium; (F) compound chaetae, midbody.
AM W28913.
75L, 77F). Aciculae distally knobbed, 3 on anterior
parapodia (Fig. 75M), diminishing progressively along body
to 1 (Fig. 75N). Pharynx through about 6–7 segments;
pharyngeal tooth located anteriorly, surrounded by crown
of 10 soft papillae and layer of cilia (Fig. 76C). Proventricle
through 7 segments, with 30 muscle cell rows. Two long,
filiform anal cirri (Fig. 72D).
Habitat. Occurring in sandy substrates, from intertidal to
about 30 m.
Distribution. Australia (Western Australia, New South
Wales).
Pionosyllis yolandae n.sp.
Figs 42C–F, 49A–C, 78A–J
Material examined. HOLOTYPE (AM W28222) AUSTRALIA: NEW
SOUTH WALES: Burrill Rocks, 35°23.39'S 150°28.24'E, on gorgonacean,
24 m, coll. R.T. Springthorpe, 7 May 1997. PARATYPES 1 (AM W28401)
S ledge, Cook Is., 28°11.65'S 153°34.63'E, colonial ascidian, 14 m, coll.
G.D.F. Wilson, 9 Jun 1993; 1 (AM W28964) N side of Bannister Head,
N of Ulladulla, 35°19.15'S 150°29.12'E, grey sponge from top of boulder,
18 m, K. Attwood, 6 May 1997.
Other material examined. TASMAN SEA Taupo Seamount,
33°16.85'S 156°09.15'E, limestone & sand bottom, 244 m, coll. J.K.
Lowry & party, on RV Franklin, 2 May 1989, 3 on SEM stub (AM
W28876); Taupo Seamount, 33°16.85'S 156°09.15'E, limestone & sand

San Martín & Hutchings: Eusyllinae of Australia 349
Fig. 77. SEM of Pionosyllis serrata (Hartmann-Schröder, 1984) (A) dorsalmost compound chaeta, midbody; (B,C) dorsal and
ventral compound chaetae, midbody; (D) ventral compound chaeta, posterior parapodium; (E) dorsal simple chaeta; (F) ventral
simple chaeta. AM W28913.
bottom, 244 m, coll. J.K. Lowry & party, on RV Franklin, 2 May 1989, few
(AM W28924). WESTERN AUSTRALIA: Reef W of groyne, 2 km S of Cape
Peron, 32°16'S 115°41'E, orange sponge in deep channel in limestone reef,
4.5 m, coll. R.T. Springthorpe, 26 Dec 1984, 1 (AM W28965).
Description. Body fragile, slender; only anterior fragments
examined; longest anterior fragment 2.4 mm long, 0.3 mm
wide, with 21 chaetigers; usually colourless, but some
specimens with few dark transverse bands on dorsum of
anterior segments, varying in intensity. Posterior to
proventricle, segments fused, forming suprasegmental
structures of 2–3 segments (Fig. 78A). Prostomium oval,
with 4 eyes in open trapezoidal arrangement; antennae
irregularly articulated, originating on anterior margin of
prostomium, median antenna inserted slightly posteriorly
behind lateral antennae (Figs 42D,E, 78A). Median antenna
with about 15 articles, about twice combined length of
prostomium and palps; lateral antennae about half length
of median antenna, with about 6–8 articles. Palps blunt,
similar in length to prostomium, free from each other,
ventrally folded in several specimens (Figs 42E, 78B).
Peristomium shorter than subsequent segments, forming
small, indistinct occipital flap (Fig. 42C–E); dorsal
tentacular cirri irregularly articulated, similar in length to
median antenna; ventral tentacular cirri about half length
of dorsal ones, almost smooth (Fig. 78A,B). Nuchal organs
as 2 dorsal densely ciliated grooves between prostomium
and peristomium (Figs 42D,E, 78A). Dorsal cirri of
chaetiger 1 elongated, articulated, with about 13 irregular

350 Records of the Australian Museum (2006) Vol. 58
Fig. 78. Pionosyllis yolandae n.sp. (A) anterior end, dorsal view; (B) detail of
prostomium and anteriormost segments, ventral view; (C) spiniger-like
compound chaetae, anterior parapodium; (D) falcigers, anterior parapodium;
(E) spiniger-like compound chaeta, midbody; (F) falcigers, midbody; (G–J)
gradation on number and shape of aciculae from anterior parapodia to midbody.
AM W28876. Scales: A 0.18 mm, B 97.5 µm, C–J 20 µm.
articulations; dorsal cirri of remaining anterior parapodia
slightly elongated, rugose, shorter than body width,
becoming progressively shorter and smooth along body;
midbody dorsal cirri alternating in length, longer ones about
half of length of body width, shorter ones about two thirds
length of longer ones (Fig. 78A). Parapodial lobes conical;
ventral cirri digitiform, slightly shorter than parapodial
lobes; ventral cirri of chaetiger 1 leaf-like, enlarged, located
mid ventrally (Fig. 78B). Anterior parapodia with about 2–
3 compound chaetae with slender, spiniger-like blades
bidentate, with short spines on margin (Figs 78C, 49A),
blades about 36 µm in length; 8–10 compound falcigerous
chaetae with distinctly shorter blades, similar in shape but
slightly larger (Figs 78D, 49B), with dorsoventral gradation
within fascicle, 22 µm in length dorsally, 15 µm in length
ventrally. Progressively along body, number of compound
chaetae with elongated blades decreases to 1–2 in midbody,
32 µm in length, remaining compound chaetae, about 6,
with shorter and wider blades, about 10 µm in length, with
prominent proximal tooth, similar in size to distal tooth or
longer (Fig. 78E,F). Dorsal and ventral simple chaetae not
seen. Anterior parapodia with 2 tricuspid aciculae (Fig.
78G), proventricular segments with single acicula,
becoming progressively larger (Fig. 78H,I), extending
beyond parapodial lobes, with large tooth and 2 smaller
ones (Fig. 78J), similar in shape to subacicular hooks in
eunicids (Fig. 49C). Pharynx proportionally long and
slender (Fig. 78A), through 8–9 segments, anterior rim with
crown of 10 soft papillae and layer of cilia (Fig. 42F);
pharyngeal tooth small, located slightly behind anterior rim.
Proventricle about half of pharynx length, with about 20
muscle cell rows. Details of posterior segments unknown.

San Martín & Hutchings: Eusyllinae of Australia 351
Remarks. Pionosyllis yolandae n.sp. is similar to
Pionosyllis aciculata San Martín, 1990, from Cuba; but,
this latter species has much shorter compound chaetae, with
typical dorsoventral gradation in length of blades within a
fascicle, and lacks elongated, spiniger-like compound
chaetae (San Martín, 1990). Pionosyllis lamelligera (Saint-
Joseph, 1887) has spiniger-like chaetae, but it lacks the
characteristic enlarged, tridentate aciculae, and the posterior
falcigers have the proximal tooth longer than distal tooth
present, whereas in Pionosyllis yolandae the teeth are of
similar size (San Martín, 2003, for a description of
Pionosyllis lamelligera).
Habitat. Occurring on gorgonaceans, ascidians, sponges,
and in sand, from 14 to 244 m depths.
Distribution. Australia (New South Wales, Western
Australia).
Etymology. The species is named after Miss Yolanda Lucas,
who helped us in many ways, especially collaborating with
us on the illustrations of this paper.
Pionosyllis sp.
Fig. 79A–G
?Pionosyllis divaricata Haswell, 1920: 104, pl. 13, figs 2, 3. Not
Keferstein, 1862: 11.
?Eusyllis sp. Hartmann-Schröder, 1984: 19, fig. 27.
Material examined. AUSTRALIA: NEW SOUTH WALES: NE corner
of Clark Is., Port Jackson, 33°51.85'S 151°14.7'E, within encrustation
on outside of bottle, 4.5 m, coll. P.A. Hutchings, 17 Apr 1996, 1 anterior
& 1 posterior fragment (AM W28410).
Description. Both fragments in poor condition; anterior
fragment of 19 segments and mid-posterior fragment of 8
segments. Similar to Pionosyllis rousei, but compound
chaetae more elongate, with longer spiniger-like chaetae
and elongated falcigers; anterior parapodia with numerous
compound chaetae, blades of spiniger-like chaetae about
69 µm long, bidentate, both teeth small and equal in size
(Fig. 79A); falcigers with blades 45–49 µm in length,
proximal tooth distinctly curved but lacking tendon (Fig.
79B). Midbody parapodia with some spiniger-like chaetae,
blades about 89 µm in length, teeth distinctly hooked, but
lacking tendon (Fig. 79C); falcigers with blades 40–51 µm
in length, most provided with tendon (Fig. 79D). Posterior
parapodia with single spiniger-like chaetae, long and
slender, blades 90 µm in length, with proximal tooth
distinctly curved, indistinct tendon (Fig. 79E); falcigers with
both teeth marked, curved, proximal tooth longer than distal
tooth, with distinct tendon between proximal tooth and blade
edge (Fig. 79F) 42–58 µm in length. Dorsal and ventral
simple chaetae not seen. Aciculae distally blunt, with short
pointed tip; 2 acicula on anterior parapodia (Fig. 79G), and
1 on posterior parapodia (Fig. 79H). Pharynx everted, with
crown of 10 long, slender soft papillae, pharyngeal tooth located
near anterior margin of pharynx. Proventricle short and wide,
through 10 segments, with about 27 muscle cell rows.
Remarks. This species does not agree with any other
described species within the genus Pionosyllis. Additional
complete material, however, is necessary before it can be
described as new species.
Fig. 79. Pionosyllis sp. (A) spiniger-like compound chaeta, anterior
parapodium; (B) falcigers, anterior parapodium; (C) spiniger-like
compound chaeta, midbody; (D) falcigers, midbody; (E) spiniger-
like compound chaeta, posterior parapodium; (F) falcigers,
posterior parapodium; (G) aciculae, anterior parapodium; (H)
acicula, posterior parapodium. AM W28410. Scale: 20 µm.

352 Records of the Australian Museum (2006) Vol. 58
Material described by Haswell as Pionosyllis
divaricata could not be located for examination,
and therefore possible identity of this species is
based only on the description and this is reflected in
the synonymies.
Habitat. Cryptic species on encrusting organisms found in
shallow water.
Distribution. Australia (New South Wales and possibly
South Australia).
Genus Psammosyllis Westheide, 1990
Psammosyllis Westheide, 1990: 165.
Type species. Psammosyllis aliceae Westheide, 1990.
Diagnosis. Body small, of meiofaunal size, with few
segments. Prostomium with 4 eyes, and 3 short antennae.
Palps fused entire length or just leaving distal notch. Nuchal
organs as 2 ciliated grooves between prostomium and
peristomium. Two pairs of tentacular cirri. Dorsal cirri on
all segments, smooth, cylindrical, longer than parapodial
lobes, shorter than body width. Ventral cirri digitiform, short.
Parapodial lobes bilobed. Compound chaetae heterogomph,
with short blades. Dorsal and ventral simple chaetae on some
parapodia. Pharynx wide, without papillae on opening,
pharyngeal tooth stout, small, located posteriorly on lateral
position of pharynx. Proventricle massive, barrel-shaped,
long and large. Pygidium with 2 long, filiform anal cirri.
Method of reproduction unknown.
Psammosyllis curticirris
(Hartmann-Schröder, 1983) n.comb.
Fig. 80A–F
Opisthodonta curticirris Hartmann-Schröder, 1983: 130, figs 11–14.
Material examined. AUSTRALIA: NEW SOUTH WALES: Hole-in-the-
Wall, Jervis Bay, 35°07.6'S 150°44.8'E, sandy mud, unvegetated
sediments, 12 m, coll. P.A. Hutchings & party, 18 Aug 1989, 1 (AM
W24706). Paratype WESTERN AUSTRALIA: Dunsborough, 33°36'S
115°06'E, fine sand, intertidal, coll. G. Hartmann-Schröder, 9 Nov 1975,
(AM W196215).
Fig. 80. Psammosyllis curticirris (Hartmann-Schröder,
1983) n.comb. (A) anterior end, dorsal view; (B)
anterior end, ventral view; (C) compound chaetae,
midbody; (D) dorsal simple chaeta; (E) ventral simple
chaeta; (F) aciculae, midbody. AM W24704. Scales:
A 0.1 mm, B 0.18 mm, C–F 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 353
Description. Body proportionally broad in relation to other
species of genus, 3.1 mm long, 0.4 mm wide, with 29
segments. Prostomium oval, width more than twice length;
2 pairs of eyes in open trapezoidal pattern, close to each
other, laterally inserted; antennae shorter than combined
length of prostomium and palps (Fig. 80A), slightly rugose,
median antenna originating between anterior eyes, slightly
longer than lateral antennae; lateral antennae shorter than
prostomium, inserted near anterior margin of prostomium.
Palps broad, fused for almost entire length, except for
distinct terminal notch, ventrally folded on one specimen
(Fig. 80B). Peristomium shorter than following segments;
dorsal tentacular cirri longer than median antenna, similar
in shape, ventral tentacular cirri shorter than dorsal ones.
Dorsal cirri smooth, similar to antennae and tentacular cirri,
rough, slightly enlarged on middle, tapered basally and
distally, longer than parapodial lobes (Fig. 80A), shorter
than half of body width. Parapodial lobes broad, bilobed
dorsally (Fig. 80A). Ventral cirri short, oblong, shorter than
parapodial lobes. Compound chaetae similar throughout,
heterogomph falcigers, blades short, about 12–14 µm in
length, short spines on margin, some apparently unidentate,
others with prominent proximal tooth located on different
plane (Fig. 80C), about 20–25 per parapodium. Dorsal
simple chaetae from anterior segments, unidentate, distally
knobbed and provided with small transparent hood and short
spines on margin (Fig. 80D). Ventral simple chaetae on
posterior chaetigers, smooth, bidentate, both teeth similar
(Fig. 80E). Aciculae distally knobbed (Fig. 80F), 3 present
anteriorly, decreasing to 1 on most posterior chaetigers.
Pharynx wide, through 5–6 segments, pharyngeal tooth
located laterally, near proventricle (Fig. 80A). Proventricle
barrel-shaped, massive, through 4–5 segments, with about
50 muscle cell rows.
Remarks. This species differs in several characters from
the recently emended description of Opisthodonta, in having
fused palps, short cirri, a massive proventricle and large
pharynx without soft papillae as well as a pharyngeal tooth
located laterally, not mid-dorsally; furthermore, Opistho-
donta as redefined above is characterized by having ventral
cirri of anteriormost segments expanded leaf-like, partially
fused with parapodial lobes, whereas in this species the
ventral cirri are short and ovoid. We are therefore
transferring Opisthodonta curticirris Hartmann-Schröder
to the genus Psammosyllis as the above characters clearly
place it in this genus rather than the emended genus
Opisthodonta. The genus Psammosyllis contains only two
other previously described species P. aliceae Westheide,
1990, described from India, and P. wui Ding & Westheide,
1997, described from China. Psammosyllis curticirris differs
from these two species by having a broader body, dorsal
simple chaetae with a distal hood, and compound chaetae
with blades unidentate or bidentate with both teeth on
different planes (see Westheide, 1990; Ding & Westheide,
1997). This represents the first record of the genus in
Australia.
Habitat. Occurring in sandy substrates, from intertidal to
shallow depths.
Distribution. Australia (Western Australia, New South
Wales).
Genus Streptodonta n.gen.
Type species. Opisthodonta pterochaeta Southern, 1914,
herein designated.
Diagnosis. Body long, slender, tapered anteriorly and
posteriorly, with numerous segments. Prostomium
pentagonal to triangular, with 4 eyes and 2 anterior eyespots.
Three antennae. Palps short, fused basally, triangular in
shape. Nuchal organs as 2 ciliated grooves between
peristomium and prostomium. Two pairs of tentacular cirri.
Antennae, tentacular and dorsal cirri elongated, smooth,
distally tapered. Ventral cirri triangular. Compound chaetae
with translucent hood on margin, ornamented with several
rows of minute spines. Dorsal simple chaetae with
translucent hood. Ventral simple chaetae probably absent.
Aciculae of several anterior parapodia distinctly enlarged.
Pharynx and proventricle long, pharyngeal tooth located
laterally and distinctly posteriorly, close to proventricle.
Reproduction by epigamy.
Remarks. The species Opisthodonta pterochaeta differs in
several characters from the emended diagnosis of
Opisthodonta, such as having palps fused basally,
pharyngeal tooth located laterally, extremely long pharynx
and proventricle, and, also having enlarged aciculae on
several anterior segments and with different types of chaetae.
As suggested by Kudenov & Harris (1995), a new genus
was needed for this species.
Streptodonta n.gen., is similar to Streptosyllis in having
enlarged aciculae on some anterior parapodia, but differs
in having a pharyngeal tooth, much longer pharynx and
proventricle, and in the shape of the compound chaetae.
It is probably more closely related to Psammosyllis
Westheide, 1990, than to Opisthodonta, sharing with the
former some unusual characters, such as the position of the
pharyngeal tooth (see Westheide, 1990).
Etymology. The name of the new genus is a combination
of the prefix streptos, from the Greek, meaning twisted, (in
reference to Streptosyllis), and the suffix donta, (in reference
to Opisthodonta), because it shares characters of both
genera, thick, enlarged aciculae, like those present in
Streptosyllis and a posterior pharyngeal tooth, like those
present in Opisthodonta.
Streptodonta pterochaeta (Southern, 1914), n.comb.
Fig. 81E
Opisthodonta pterochaeta Southern, 1914: 30, pl. IV, figs 6A–
G.—Fauvel, 1923: 274, fig. 102d–i.—Hartmann-Schröder,
1971: 104, figs 4–6; 1996: 160.—Campoy, 1982: 304, pl. 24.—
Bachelet, 1990: 174, fig. 1.—Parapar et al., 1993: 370, fig.
4.—San Martín, 2003: 51, figs 13, 14.
Material examined. AUSTRALIA: NEW SOUTH WALES: Bass Point,
34°36'S 150°54'E, 50 m, 1 Feb 1990, 1 (AM W22989).
Description. Body slender, with numerous segments.
Incomplete specimen, 4.2 mm long, 0.4 mm wide, with 31
chaetigers; an entire individual, may reach 10 mm or more.
Prostomium pentagonal to triangular, with 2 pairs of eyes
arranged in open trapezoidal pattern, and 2 anterior eyespots
(Fig. 81A). Median antenna inserted between anterior eyes,
slightly longer than combined length of prostomium and

354 Records of the Australian Museum (2006) Vol. 58
Fig. 81. Streptodonta pterochaeta (Southern, 1914) (A) anterior end, dorsal view; (B) non-
enlarged acicula, chaetiger 1; (C) enlarged aciculae; (D) compound chaetae, posterior
parapodium; (E) dorsal simple chaeta. AM W22989. Scales: A 0.4 mm, B–E 20 µm.
palps, lateral antenna in front of anterior eyes. Palps small,
triangular, basally fused, shorter than prostomium (Fig.
81A). Peristomium slightly shorter than following segments;
dorsal tentacular cirri longer than lateral antennae, shorter
than median, ventral tentacular cirri shorter than dorsal ones.
Dorsal cirri similar in shape to antennae and tentacular cirri,
smooth, elongated, relatively short, shorter than body width
(Fig. 81A), detached on most parapodia. Ventral cirri
triangular, similar in length to parapodial lobes. Compound
chaetae heterogomph falcigers, blades with indistinct distal
tooth and longer, prominent proximal tooth, provided with
translucent hood on margin (Fig. 81D). Numerous
compound chaetae on anterior parapodia, decreasing to 10
on midbody parapodia, blades all similar, but longer ones,
about 20 µm in length, with more prominent distal tooth,
shorter ones about 15 µm in length, with distal tooth poorly
developed. Dorsal simple chaetae on midbody parapodia,
unidentate, with distal, translucent hood (Fig. 81E). Aciculae
of chaetigers 1, and chaetigers 26 onwards slender, knobbed
distally (Fig. 81B); aciculae of chaetigers 2–26, larger,
distally strongly knobbed, with terminal button (Fig. 81C).
Pharynx long, through 20–21 segments; pharyngeal tooth
small, located laterally, close to proventricle, far from
anterior rim of pharynx (Fig. 81A). Proventricle through 7
segments, with about 50 muscle cell rows.
Remarks. The Australian specimen differs from material
described from Europe, in being proportionally smaller,
having smaller palps, shorter antennae and dorsal cirri,
longer pharynx and shorter proventricle, and varying in the
length of chaetal blades. As we have only a single incomplete
specimen, slightly damaged, we prefer to assign it to this
described species. It may represent a new species, but additional
material is required. San Martín (2003, Fig. 14C,D), found
that the chaetal hoods of specimens from the Mediterranean
are composed of several rows of minute spines, the
Australian specimen was not examined under SEM.
Habitat. Occurring in coarse sand, in depths of 6 to 50 m.
Distribution. Eastern Atlantic Ocean, from North Sea to
Straits of Gibraltar, Australia (New South Wales).
Genus Streptosyllis Webster & Benedict, 1884
Streptosyllis Webster & Benedict, 1884: 711.
Type species. Streptosyllis arenae Webster & Benedict,
1884, by monotypy.
Diagnosis. Body small (<5 mm in length), with up to about
40 chaetigers, usually 20–30. Prostomium with 4 eyes and
2 anterior eyespots. Three antennae. Palps fused at bases,
without median furrow, sometimes terminating with 2
papillae, occasionally reduced and only papillae visible.
Nuchal organs as 2 ciliated grooves between prostomium

San Martín & Hutchings: Eusyllinae of Australia 355
and peristomium. Dorsal cirri usually smooth or indistinctly
articulated, club-shaped to elongated, sometimes with
glandular inclusions. Ventral cirri digitiform, sometimes
distinctly longer than parapodial lobes and pseudo-
articulated, arising from middle of parapodial lobes.
Compound chaetae, homogomph to hemigomph, blades
falcigerous or, exceptionally, some spiniger-like chaetae.
Dorsal simple chaetae present usually from chaetiger 1.
Ventral simple chaetae absent. Aciculae from some anterior
parapodia distinctly enlarged. Pharynx unarmed, provided with
crown of soft papillae. Proventricle with poorly defined muscle
cell rows. Reproduction by epigamy (Garwood, 1991).
Key to Australian species of Streptosyllis
1 Palps reduced to 2 small papillae (Fig. 82A). Compound chaetae
similar throughout ........................................................................................................ S. aequiseta
—— Palps not reduced to papillae. Compound chaetae of some anterior
parapodia with distinctly different chaetae to those of remaining
parapodia, with short blades .......................................................................................................... 2
2 Enlarged aciculae and chaetae with short blades on chaetigers 2–
4 (Fig. 84B). Blades of compound chaetae all unidentate (Fig.
84E). Palps with distal articulation .......................................................................... S. biarticulata
—— Enlarged aciculae on chaetigers 2–6 and chaetae with short blades
on chaetigers 2–6. Compound chaetae with both unidentate and
bidentate blades. Palps without distal articulation ................................................. S. magnapalpa
Streptosyllis aequiseta Hartmann-Schröder, 1981
Figs 82A–I, 83A–F
Streptosyllis aequiseta Hartmann-Schröder, 1981: 32, figs 53–
58; 1983: 131, fig. 15; 1984: 21; 1985: 70; 1989: 26.—
Böggemann et al., 2003: 21, figs 3, 4.
Material examined. AUSTRALIA: NEW SOUTH WALES: Weeney Bay,
Botany Bay, 34°01.3'S 151°09.7'E, mud, 1 m, coll. A. Roach & A. Jones,
30 Mar 1995, 1 (AM W23562); Weeney Bay, Botany Bay, 34°01.3'S
151°09.7'E, mud, 1 m, coll. A. Roach & A. Jones, 30 Mar 1995, 1 (AM
W23567). WESTERN AUSTRALIA: Bush Bay, 30 km S of Carnarvon,
25°10'S 113°39'E, lumps of algae on shallow sandflats, intertidal, coll.
H.E. Stoddart, 6 Jan 1984, 2 (AM W27654); inshore limestone reef,
Ned’s Camp, Cape Range National Park, 21°59'S 113°55'E, fine sediment
& sand from patches in reef, 1 m, coll. H.E. Stoddart, 2 Jan 1984, 2 (AM
W26782); N end of beach, Bundegi Reef, Exmouth Gulf, 21°49'S
114°11'E, rocky rubble, coralline algae with green epiphyte, 1.5 m, coll.
H.E. Stoddart, 4 Jan 1984, 1 on SEM stub (AM W28370).
Description. Up to 4.9 mm long for 36 chaetigers (mature
specimen), but usually smaller; examined specimens up to
1.4 mm long, 0.2 mm wide, with 23 chaetigers; fragile, often
broken and damaged. Prostomium oval, with 4 eyes
arranged in an open trapezoidal pattern and 2 anterior
eyespots (Fig. 82A). Antennae smooth, club-shaped, similar
in length to prostomium or longer, median antenna inserted
between anterior eyes, lateral antennae inserted near
eyespots. Palps reduced to 2 small, sometimes indistinct
papillae (Figs 82A, 83B). Peristomium shorter than
subsequent segments; tentacular cirri similar to antennae,
dorsal ones slightly longer than ventral tentacular cirri.
Dorsal cirri similar in shape and size to antennae (Fig. 82A)
with distinct cirrophores. Parapodial lobes elongated,
subrectangular, ending as rounded lobe (Fig. 82B), those
of chaetigers 2–6 enlarged and truncated distally. Ventral
cirri digitiform, elongated, longer than parapodial lobes,
arising from about middle of ventral side of parapodial lobes
(Figs 82B, 83A). Compound chaetae with homogomph
articulations on anterior parapodia (Figs 82C,D, 83D), and
hemigomph on mid to posterior parapodia (Figs 82F, 83C),
provided with distinct subdistal spine on shafts, and
bidentate blades, with short spines on margin. Anterior
parapodia with 2 compound chaetae with elongate blades,
about 21 µm long, distinctly bidentate, both teeth well
separated (Figs 82C, 83D), and 6 compound chaetae with
shorter blades (Fig. 82D), within fascicle blades 10 µm in
length dorsally, 6 µm in length ventrally; difference between
2 types of chaetae becoming progressively less marked
along body; posterior parapodia with 6 compound chaetae
(Fig. 82F), bidentate with short spines on margin,
dorsoventral gradation in length of blades within fascicle,
25 µm long dorsally, 9 µm long ventrally. Dorsal simple
chaetae from chaetiger 1, unidentate, with minute serration
on margin and distinct concave, translucent hood, covering
tip of chaetae, small on simple chaetae of anterior parapodia
(Figs 82E, 83E) but becoming more developed posteriorly
(Figs 82G,H, 83F). Aciculae knobbed at tips, enlarged on
chaetigers 2–6 (Fig. 82I). Pharynx extending through 3–4
segments, with crown of 10 soft papillae on margin (Fig.
83A). Proventricle large, pyriform, extending through about
6 segments and 40–48 indistinct muscle cell rows. Pygidium
small, with 2 anal cirri and median papilla.
Remarks. All Australian specimens have all dorsal cirri
smooth; those from the Seychelles (Böggemann et al., 2003)
have longer dorsal cirri, articulated, with two glands on each
article, except anteriormost, which are smooth. Until more
material is available, it is unknown as to the levels of
variation of articulation of dorsal cirri that may occur within
a species. So at this stage we are tentatively accepting the
records of this species from the Seychelles.
Habitat. Occurring interstitially in coralline sand, fine sand,
mud, on algae, from intertidal to shallow depths.
Distribution. Australia (Western Australia, South Australia,
Tasmania, New South Wales), possibly Seychelles Islands.

356 Records of the Australian Museum (2006) Vol. 58
Fig. 82. Streptosyllis aequiseta Hartmann-Schröder, 1981 (A)
anterior end, dorsal view; (B) posterior parapodium; (C)
dorsalmost compound chaetae, anterior parapodium; (D)
remaining compound chaetae, anterior parapodium; (E)
dorsal simple chaeta, lateral view, anterior parapodium; (F)
compound chaetae, posterior parapodium; (G) two different
views of dorsal simple chaetae, posterior parapodium; (H)
detail of tip, inferior view, dorsal simple chaeta, posterior
parapodium; (I) aciculae from chaetigers 1–8. AM W28370.
Scales: A 0.18 mm, B 92 µm, C–I 20 µm.
Streptosyllis biarticulata Hartmann-Schröder, 1991
Fig. 84A–F
Streptosyllis biarticulata Hartmann-Schröder, 1991: 36, figs 57–61.
Material examined. AUSTRALIA: QUEENSLAND: Heron Is., Great
Barrier Reef, 23°27'S 151°55'E, coarse sand, intertidal, 4 Feb 1976,
coll. G. Hartmann-Schröder, holotype (HZM P-20545).
Description. Body 2.5 mm long, 0.2 mm wide, with 35
chaetigers, incomplete specimen. Prostomium oval, with 4
large eyes in trapezoidal arrangement. Antennae broken (and
not in vial); point of insertion not visible. Palps shorter than
prostomium, fused basally. Peristomium well defined,
shorter than following segments; dorsal tentacular cirri with
about 6 articles, ventral tentacular cirri smooth, shorter than
dorsal ones. Most dorsal cirri detached; dorsal cirri with 4–
7 articles. Enlarged aciculae (Fig. 84C) from chaetigers 2–

San Martín & Hutchings: Eusyllinae of Australia 357
Fig. 83. SEM of Streptosyllis aequiseta Hartmann-Schröder, 1981. (A) anterior end, ventral view; (B) detail of prostomium,
showing the papillae of palps; (C) dorsal compound chaeta, posterior parapodium; (D) same, anterior parapodium; (E) dorsal
simple chaeta, anterior parapodium, lateral view; (F) dorsal simple chaeta, inferior view, posterior parapodium. AM W28370.
4, large basally, distally blunt. Subsequent aciculae and
aciculae of chaetiger 1, similar but much thinner (Fig. 84F).
Compound chaetae with thick shafts, some of them provided
with thin hood distally, and blades distally blunt, unidentate,
with short spines on margin, and slight dorsoventral
gradation in length of blades within fascicle (Fig. 84E), 13
µm in length dorsally, 7 µm in length ventrally. Compound
chaetae of chaetigers 3–4 with distinctly shorter blades,
blunt, short, with thick shafts, some covered by translucent
hood (Fig. 84B). Dorsal simple chaetae from chaetiger 1,
thick, distally blunt, with hood, longitudinally striated (Fig.
84A), becoming thicker posteriorly (Fig. 84D). Ventral
simple chaetae absent. Pharynx extends for about 4
segments. Proventricle extending for 3.5 segments, with
about 35 muscle cell rows.
Habitat. Occurring in coral sand, intertidally.
Distribution. Australia (Queensland).

358 Records of the Australian Museum (2006) Vol. 58
Fig. 84. Streptosyllis biarticulata Hartmann-Schröder, 1991 (A)
dorsal simple chaeta, anterior parapodium; (B) compound chaetae
from parapodia 2–4; (C) enlarged acicula (chaetigers 2–4); (D)
dorsal simple chaeta, midbody; (E) compound chaetae, midbody;
(F) acicula, midbody. ZMH P-20545. Scale: 20 µm.
Streptosyllis magnapalpa Hartmann-Schröder, 1981
Fig. 85A–H
Streptosyllis magnapalpa Hartmann-Schröder, 1981: 33, figs 59–
67; 1982: 65, fig. 47; 1983: 13; 1987: 38.
Material examined. AUSTRALIA: WESTERN AUSTRALIA: Horrocks,
algae, seagrass and sand, 28°23'S 114°26'E, intertidal, 17 Oct 1975,
coll. G. Hartmann-Schröder, paratype (HZM P-16488); Cervantes,
30°30'S 115°03'E, fine sand and Posidonia, intertidal, coll. G. Hartmann-
Schröder, 24 Oct 1975, 1 (HZM P-17019).
Description. Body fragile, without colour markings; largest
complete specimen 2.8 mm long, 0.2 mm wide, with 37
chaetigers. Prostomium oval, with 4 eyes in open trapezoidal
arrangement; tuft of cilia present in front of anterior eyes.
Antennae short, smooth, slightly club-shaped; median
antenna missing (broken and lost), arising on middle of
prostomium; lateral antennae inserted near anterior margin
of prostomium (Fig. 85A). Palps shorter than prostomium,
fused basally. Peristomium slightly shorter than following
segment, provided with some hyaline inclusions (Fig. 85A);
tentacular cirri similar to lateral antennae in shape; dorsal
tentacular cirri slightly longer than lateral antennae, ventral
tentacular cirri shorter than dorsal. Transverse row of cilia
on dorsum of each segment (Fig. 85A). Dorsal cirri of
anterior 2–3 segments similar to antennae and tentacular
cirri, although slightly longer. Subsequent dorsal cirri
articulated, with few articles, 4–6, some of them with
granular, dark inclusions. Aciculae slender, distally blunt
(Fig. 85D); enlarged aciculae on chaetigers 2–6. Compound
chaetae of chaetigers 2–6 with thick shafts, short, unidentate
blades, some covered by translucent hood, with slight
dorsoventral gradation in length of blades within fascicle,
18 µm in length dorsally, and 11 µm in length ventrally
(Fig. 85C). Compound chaetae of subsequent chaetigers
with elongate, unidentate, distally blunt blades, with short
spines on margin, shafts with thick subdistal spine, and
dorsoventral gradation in length within fascicle, 47 µm
dorsally, 22 µm ventrally (Fig. 85F); progressively on
following segments, blades becoming slender, and tips
indistinctly bidentate (Fig. 85G). Anterior parapodia with
about 6 compound chaetae, reducing to 4 on posterior
parapodia. Dorsal simple chaetae from chaetiger 1, thick,
distally blunt, with distal, longitudinally striated hood,
similar throughout body (Fig. 85B,E,H). Pharynx long,
through about 5–6 segments, with distal crown of 10 papillae
and subdistal crown of much smaller papillae (Fig. 85A).
Proventricle through about 5 segments, with 35 muscle cell
rows. Pygidium small, with two long anal cirri and median
short papilla.
Habitat. Occurring in algae, seagrass, sand, from intertidal
to shallow depths.
Distribution. Australia (Western Australia, Victoria).
Genus Syllides Örsted, 1845
Syllides Örsted, 1845: 408.
Type species. Syllides longocirrata Örsted, 1845,
designated by Hartman, 1959.
Diagnosis. Body small, short, with relatively few segments.
Prostomium with 4 eyes, typically pair of anterior eyespots.
Three antennae. Palps fused basally, without median furrow,
sometimes ending with small papilla. Two pairs of tentacular
cirri. Nuchal organs as 2 ciliated grooves between
prostomium and peristomium. Antennae, tentacular cirri,
and dorsal cirri of chaetigers 1 and 2, smooth, non-
articulated, club-shaped to fusiform; dorsal cirri from
chaetiger 3 onwards, distinctly articulated, with glandular
inclusions on some articles. Ventral cirri digitiform.
Compound chaetae heterogomph with blades slender,
usually bidentate. Dorsal simple chaetae from anterior
segments, usually from chaetiger 1. Ventral simple chaetae
present in some species. Pharynx unarmed, with crown of
soft papillae. Reproduction by epigamy, some species brood
eggs ventrally (Heacox & Schroeder, 1978).

San Martín & Hutchings: Eusyllinae of Australia 359
Fig. 85. Streptosyllis magnapalpa Hartmann-Schröder, 1981 (A) anterior
end, dorsal view; (B) dorsal simple chaeta, anterior parapodium; (C)
compound chaetae of chaetigers 2–4; (D) acicula, midbody; (E) dorsal
simple chaeta, midbody; (F) compound chaetae, midbody; (G) compound
chaetae, posterior parapodium; (H) dorsal simple chaeta, posterior
parapodium. ZMH P-17019. Scales: A 0.1 mm, B–H 20 µm.
Key to Australian species of Syllides
1 Blades of some compound chaetae provided with long, basal spurs
on margin (Fig. 89B)...................................................................................................................... 2
—— Without basal spurs on blades of compound chaetae (Fig. 88D).................................................. 4
2 Longest blades of compound chaetae lacking basal spurs, only
present on medium-length blades .................................................................................. S. spinosus
—— Longest blades of compound chaetae with basal spurs................................................................. 3
3 Dorsal simple chaetae distally rounded, with distal hood (several
rows of minute spines, under SEM) ............................................................................. S. tam n.sp.
—— Dorsal simple chaetae pointed, without hood ............................................................. S. japonicus
4 Dorsal simple chaetae distally rounded, with distal hood (several
rows of minute spines, under SEM). Proventricle long, through 4
segments .......................................................................................................................... S. pumilus
—— Dorsal simple chaetae distally bifid, without hood. Proventricle
short, through 2 segments ............................................................................................. Syllides sp.

360 Records of the Australian Museum (2006) Vol. 58
Syllides japonicus Imajima, 1966
Figs 86C–F, 87A–E
Syllides japonicus Imajima, 1966: 112, text-fig. 36.—San Martín,
2003: 142, fig. 69.
Material examined. AUSTRALIA: NEW SOUTH WALES: 1 km SE of
Dangar Is., Hawkesbury R., 33°33'S 151°14'E, shelly mud, 9 m, coll.
A.R. Jones & party, 18 Dec 1979, 1 (AM W28467); Pittwater, 33°35.80'S
151°18.31'E, muddy sand, 14.4 m, coll. Australian Museum Party, 31
July 1995, 2 (AM W23927); 1 km SE of Dangar Is., Hawkesbury R.,
33°33'S 151°14'E, shelly mud, 9 m, coll. A.R. Jones & party, 1 Aug
1979, 1 (AM W28975); 1.5 km SE of Dangar Is., Hawkesbury R., 33°33'S
151°14'E, sandy mud, 7 m, coll. A.R. Jones & party 18 Dec 1979, 1
(AM W28976). TASMANIA: Fancy Point, Bruny Is. 43°16'S 147°19'E,
Fig. 86. SEM of Syllides tam n.sp. (A) compound chaetae of first and second pair (arrows showing basal spurs); (B) dorsal simple
chaeta. SEM of Syllides japonicus Imajima, 1966 (C) anterior end, dorsal view (antennae and cirri missing; arrows showing papillae
of palpi); (D) detail of papilla of palp; (E) dorsal simple chaeta and some compound chaetae; (F) detail of basal spur of compound
chaeta. A,B: AM W196520; C–F: W23927.
algae, 4 m, coll. G. Edgar, 10 Nov 1980, 1 (AM W18204). WESTERN
AUSTRALIA: Inshore limestone reef, Ned’s Camp, Cape Range National
Park, 21°59'S 113°55'E, Caulerpa sp. algae, 1 m, coll. J.K. Lowry, 2
Jan 1984, 1 (AM W26739); Inshore limestone reef, Ned’s Camp, Cape
Range National Park, 21°59'S 113°55'E, small purple sponge with
Caulerpa sp. algae & sticky sediment, 1.5 m, coll. R.T. Springthorpe, 2
Jan 1984, 2 (AM W26781).
Description. Body less than 5 mm in length, densely
covered with golden inclusions on dorsum of each segment
and prostomium (Fig. 87A). Prostomium oval, about twice
as wide as long; 4 small eyes in open trapezoidal
arrangement; antennae missing on most specimens, short,
smooth. Palps broad, fused at bases, shorter than
prostomium, each ending with distal, small papilla (Figs

San Martín & Hutchings: Eusyllinae of Australia 361
Fig. 87. Syllides japonicus Imajima, 1966 (A) anterior end, dorsal
view; (B) dorsal simple chaeta; (C) acicula; (D) dorsalmost
compound chaetae; (E) remaining compound chaetae, midbody
parapodium. AM W28467. Scales: A 0.18 mm, B–E 20 µm.
86C,D, 87A). Tentacular and dorsal cirri mostly missing.
Parapodia elongated, ventral cirri digitiform. Compound
chaetae with elongated, distinctly bidentate blades, arranged
in 5 pairs of chaetae within fascicle, each pair of similar
shape and length; blades of dorsalmost pair with long blades,
about 55 µm long, with basal long spur (Figs 86F, 87D)
and short spines on margin; blades of subsequent pair similar
(Fig. 87E) but shorter, about 44 µm long; remaining
compound chaetae similar in shape, lacking basal spur (Fig.
87E), with dorsoventral gradation in length of blades, 35
µm in length dorsally and 22 µm in length ventrally. Dorsal
simple chaetae from chaetiger 1, thin, pointed (Figs 86E,
87B), with short spines on margin (Fig. 87B). Ventral simple
chaetae absent. Aciculae slender, distally knobbed (Fig.
87C) pharynx everted on most specimens, through about 4
segments when relaxed. Proventricle long, through 7
segments (Fig. 87A), with about 47 muscle cell rows.
Remarks. The original description of the species from Japan
and descriptions of Mediterranean specimens do not
describe the terminal papillae on the palps; these structures
are small and could easily be overlooked. This species is
characterized by having basal spurs on the bases of blades
of the dorsalmost and second pair of compound chaetae;
palps each with a small papilla on the Australian material,
and dorsal simple chaetae, slender. Syllides caribica Licher,
1996, from Aruba Island in the Caribbean is similar, but
differs in the following characters: the spurs on the
compound chaetae are shorter, and have a smooth area
between the spur and the edge of blade, the proventricle is
shorter (through 2 segments), and the dorsal simple chaetae
are slightly thicker (see Licher, 1996). Syllides floridanus
Perkins, 1981 also has papillae on the palps, a long
proventricle and slender, distally pointed dorsal simple
chaetae; but the blades of the dorsalmost pair of compound
chaetae lack basal spurs; they are present only on the second
pair (Perkins, 1981).
All the material examined from Australia is damaged and
while it closely resembles material examined from the
Mediterranean, it may represent an undescribed species, but
complete specimens are needed in order to confirm this.
Habitat. Occurring in sand, sandy mud, rhizomes of
Posidonia, on algae and sponges; in shallow depths.
Distribution. Japan, Western Mediterranean, USA
(Massachusetts), Australia (New South Wales, Tasmania,
Western Australia).
Syllides pumilus Hartmann-Schröder, 1983
Fig. 88A–F
Syllides articulosus pumilus Hartmann-Schröder, 1983: 131, figs
16, 17; 1984: 21; 1986: 42; 1987: 38; 1989: 26.
Material examined. AUSTRALIA: NEW SOUTH WALES: S of airport
runway extension, Botany Bay, 33°58.13'S 151°11.16'E, 5 m, coll.
Australian Museum party, 6 Apr 1992, 1 (AM W23118); Currambene
Creek, Jervis Bay, 35°02'S 150°40'E, intertidal unvegetated sediment,
coll. L. Howitt, Dec 1988, 1 (AM W22608). WESTERN AUSTRALIA:
Dunsborough, 33°36'S 115°06'E, Posidonia & Halophila seagrasses,
intertidal, coll. G. Hartmann-Schröder, 9 Nov 1975, 2 paratypes (AM
W196214); reef W of Groyne, 2 km S of Cape Peron, 32°16'S 115°41'E,
orange sponge in deep channel in limestone reef, 4.5 m, coll. R.T.
Springthorpe, 26 Dec 1984, 1 (AM W28367); Limestone reef, off Ned’s
camp, Cape Range National Park, 21°59'S 113°55'E, sponge with
epiphytic algae, & muddy worm tubes, 1.5 m, coll. R.T. Springthorpe, 2
Jan 1984, 1 (AM W27652).
Description. Body 2.4 mm long, 0.2 mm wide, with 30
chaetigers, dorsum with bright, minute inclusions, density
of inclusions varies between individuals, fragile, most
appendages missing on all specimens. Prostomium oval,
with 4 eyes arranged in open trapezoidal pattern, and 2

362 Records of the Australian Museum (2006) Vol. 58
Fig. 88. Syllides pumilus Hartmann-Schröder, 1983 (A) anterior end, dorsal view; (B) dorsal simple chaeta; (C)
acicula; (D) dorsalmost pair of compound chaetae; (E) second pair of compound chaetae; (F) compound chaetae of
remaining pairs, midbody parapodium. AM W26322. Scales: A 0.18 mm, B–F 20 µm.
anterior eyespots; 3 antennae all similar. Palps basally fused,
shorter than prostomium (Fig. 88A). Peristomium with
hyaline inclusions, larger than inclusions on dorsum;
tentacular cirri elongated, slightly club-shaped, longer than
lateral antennae (Fig. 88A), ventral tentacular cirri about
two thirds length of dorsal cirri. Dorsal cirri of chaetigers
1, 2, elongated, longer than tentacular cirri, similar in length
to body width; from chaetiger 3 onwards, dorsal cirri
articulated (Fig. 88A), with about 10 articles (up to 16, fide
Hartmann-Schröder), cirri mostly broken, articles usually
with 1–2 dark, granular inclusions. Parapodia trapezoidal
dorsally, slightly larger on anterior parapodia, becoming
conical and slender from mid parapodia onwards. Ventral
cirri digitiform, similar in length or shorter than parapodial
lobes. About 15 compound chaetae on anterior parapodia,
numbers diminishing progressively along body to about 10;
shafts with thin subdistal spines; blades elongated, bidentate,
with short spines on margin, without basal spurs (Fig. 88D–
F), with dorsoventral gradation in length of blades within
fascicle, about 50 µm in length dorsally, 20 µm in length
ventrally, on midbody. Dorsal simple chaetae from chaetiger
1, distally blunt, with small distal hood and short spines on
margin (Fig. 88B). Ventral simple chaetae absent. All
parapodia with single acicula, distally knobbed, with
filiform, short tip (Fig. 88C). Pharynx through 4 segments.
Proventricle through 4 segments (Fig. 88A), with 24–30
muscle cell rows.
Remarks. Syllides articulosus Ehlers, 1897, from Magellan
Strait, in South America, appears to be a different species
to the Australian specimens that were described by
Hartmann-Schröder (1983) as a subspecies Syllides
articulosus pumilus. Syllides articulosus has a proventricle
distinctly longer than the pharynx (see Ehlers, 1897, pl. II,
Fig. 52), and the dorsal simple chaetae are distally pointed,
and lack a distal hood (Banse, 1972). We suggest that such
differences are sufficient to warrant elevating the subspecies
of Hartmann-Schröder to a full species Syllides pumilus.
Syllides bansei Perkins, 1981, from Florida and the
Mediterranean Sea, has similar dorsal simple chaetae, but
the blades of the second pair of compound chaetae within
each fascicle are provided with a long basal spur (see
Perkins, 1981; San Martín, 2003).
Habitat. Occurring on sponges, algae, Posidonia, and in
sediments, from intertidal to shallow depths.
Distribution. Australia (Western Australia, South Australia,
Victoria, New South Wales).
Syllides spinosus Hartmann-Schröder, 1979
Fig. 89A–C
Syllides articulosus spinosus Hartmann-Schröder, 1979: 99, figs
126–128; 1981: 34; 1991: 37.
Material examined. AUSTRALIA: NEW SOUTH WALES: NE corner
of Clark Is., 33°51.85'S 151°14.47'E, encrustation on outside of bottle,
5 m, coll. P.A. Hutchings, 17 Apr 1996, 2 (AM W26322). WESTERN
AUSTRALIA: Exmouth, Tantabiddy Creek, 21.56°S 113°58'E, medium
sand, intertidal, coll. G. Hartmann-Schröder, 1 (HZM P-16907).
Description. Body 2.2 mm long, 0.3 wide mm with 30
chaetigers. Dorsal cirri long, with hyaline vacuoles on most
articles (Fig. 89A). Tufts of cilia in front of each anterior
eyespot. Ciliary bands on dorsum of each segment (Fig.
89A). Parapodia with about 5 compound chaetae, relatively
long blades, bidentate, both teeth distinctly separated, with
short spines on margin; blades of 2 medium length

San Martín & Hutchings: Eusyllinae of Australia 363
Fig. 89. Syllides spinosus Hartmann-Schröder, 1979 (A) anterior end,
dorsal view; (B) compound chaetae, midbody; (C) dorsal simple chaeta.
HZM P-16907. Scales: A 0.2 mm, B,C 20 µm.
compound chaetae with long, basal spur (Fig. 89B). Dorsal
simple chaetae from chaetiger 1, distally indistinctly bifid,
with short spines on margin (Fig. 89C). Pharynx through
about 4–5 segments. Proventricle through about 4 segments,
with 40 muscle cell rows (Fig. 89A).
Remarks. The long, basal spurs on medium-length blades
of compound chaetae have not previously been described.
This character is considered to be a useful character to
distinguish between species in this genus. Syllides bansei
Perkins, 1981, has similar compound chaetae, but the dorsal
simple chaetae differ in having a distal hood (Perkins, 1981).
Syllides edentatus Westheide, 1974, also has a similar
arrangement of chaetae, but the pair of the medium size
chaetae have several long spines instead of a single basal
spur as occurs in Syllides spinosus (Westheide, 1974; San
Martín, 2003). Syllides benedicti Banse, 1972, also has a
single long basal spur on the pair of medium-length bladed
chaetae, but the dorsal simple chaetae are distally rounded,
with some spines present in addition to the spines on margin
(Banse, 1972). For a discussion of the differences between
the stem species S. articulosus and S. spinosus see
Hartmann-Schröder, 1979; furthermore, S. articulosus lacks
long basal spurs on any blade (Somaschini & San Martín,
1997).
Habitat. Occurring in coarse and medium sand, on
encrustations, from intertidal to shallow depths.
Distribution. Australia (Western Australia, Victoria, South
Australia, New South Wales).
Syllides tam n.sp.
Figs 86A,B, 90A–G
Syllides longocirrata.—Augener, 1913: 229.—Haswell, 1920:
102.—Hutchings & Murray, 1984: 33. Not Örsted, 1845: 11.
Material examined. HOLOTYPE (AM W28472) AUSTRALIA: NEW
SOUTH WALES: 1.5 km SE of Dangar Is., Hawkesbury R., 33°33'S 151°14'E,
sandy mud, 7 m, coll. Jones & party, 21 Aug 1980. PARATYPES. 50 m NE of
Green Point, Hawkesbury R., 33°33.5'S 151°14.5'E, sandy mud, 4 m, coll.
A.R. Jones & party, 17 May 1982, 1 (AM W24704); 300 m NE of Green
Point, Hawkesbury R., 33°34'S 151°13.5'E, sandy mud, 5 m, coll. A.R.
Jones & A. Murray, 18 Nov 1980, 1 (AM W28463); 1 km S of E end,
Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy sand, 12 m, coll.
A.R. Jones & A. Murray, 5 Aug 1983, 1 (AM W28464); 1 km S of E end,
Spectacle Is., Hawkesbury R., 33°32'S 151°07.5'E, muddy sand, 12 m, coll.
A.R. Jones & A. Murray, 11 Nov 1983, 1 (AM W28465); 1 km SE of Dangar
Is., Hawkesbury R., 33°33'S 151°14'E, shelly mud, 9 m, coll. A.R. Jones &
party, 1 Aug 1979, 1 (AM W28466); 1 km SE of Dangar Is., Hawkesbury
R., 33°33'S 151°14'E, shelly mud, 9 m, coll. A.R. Jones & party, 21 Aug
1980, 1 (USNM 1082992); 30 m SE of Brooklyn boat-channel, Hawkesbury
R., 33°33'S 151°14'E, shelly mud, 7 m, coll. A.R. Jones & party, 18 Dec
1979, 1 (LACM ex AM W28469); 30 m SE of Brooklyn boat-channel,
Hawkesbury R., 33°33'S 151°14'E, shelly mud, 7 m, coll. A.R. Jones &
party, 1 Aug 1979, 1 (AM W28470); 1.5 km SE of Dangar Is., Hawkesbury
R., 33°33'S 151°14'E, sandy mud, 7 m, coll. A.R. Jones & party, 1 Aug

364 Records of the Australian Museum (2006) Vol. 58
1979, 3 (AM W28471); 1.5 km SE of Dangar Is., Hawkesbury R., 33°33'S
151°14'E, sandy mud, 7 m, coll. A.R. Jones & party, 18 Dec 1979, 1 (AM
W28473); Brooklyn boat-channel, Hawkesbury R., 33°33'S 151°14'E, fine
mud, 3 m, coll. A.R. Jones, 16 May 1980, 1 (AM W28474); Midway between
Green Point and Croppy Point, Hawkesbury R., 33°33.5'S 151°14.5'E, mud,
6 m, coll. A. Jones & party, 22 Nov 1980, 1 (AM W196517); E end Brooklyn
boat channel, 1 (AM W196518); Liverpool Reach, Hawkesbury R., 33°25'S
150°56'E, fine sand, 4 m, coll. A. Jones & party, 21 Feb 1980, 1 (AM
W196657); Liverpool Reach, Hawkesbury R., 33°25'S 150°56'E, coarse
sand, 20 m, coll. A. Jones & party, 21 Feb 1980, 3 (AM W196658); Liverpool
Reach, Hawkesbury R., 33°25'S 150°56'E, coarse sand, 20 m, coll. A. Jones
& party, 21 Feb 1980, 21 (AM W196659).
Fig. 90. Syllides tam n.sp. (A) anterior end, dorsal view; (B) posterior end, dorsal view; (C) parapodial lobe; (D) compound chaetae of
longest and second pair, midbody parapodium; (E) compound chaeta of third pair; (F) compound chaeta of fourth pair; (G) compound
chaeta of fifth pair; (H) dorsal simple chaeta; (I) acicula. AM W28472. Scales: A,B 0.1 mm; C 37 µm; D–I 20 µm.
Additional material examined. QUEENSLAND: At mouth of
Althaus Creek, Halifax Bay, 19°10'S 146°36'E, 2 m, coll. Queensland
Nickel, Jan 1977, 2 (AM W28207). NEW SOUTH WALES: 0.5 km E. of
Dangar Is., Hawkesbury R., 33°33'S 150°14'E, coll. A. Jones & party,
21 Aug 1980, 4 on SEM stub (AM W196520); S of airport runway
extension, Botany Bay, 33°58.13'S 151°11.16'E, 5 m, coll. Australian
Museum party, 7 Apr 1992, 1 (AM W21631); SW of airport runway
extension, Botany Bay, 33°58.33'S 151°10.22'E, 7 m, coll. Australian
Museum party, 28 July 1992, 1 (AM W21632); E of Marley, 34°08.08'S
151°09.65'E, sand, 60 m, coll. Fisheries Research Institute (NSW), 2
May 1991, 1 (AM W24377); Bass Point, 34°36'S 150°54'E, 50 m, coll.
The Ecology Lab for RMI/Pioneer Project, 1 Feb 1990, few (AM

San Martín & Hutchings: Eusyllinae of Australia 365
W22994); Montagu Roadstead, Jervis Bay, 35°02.2'S 150°46'E, 12 m,
coll. P.A. Hutchings & party, 6 Jun 1991, 1 (AM W27574). VICTORIA:
Port Phillip Bay, 38°09.3'S 144°42.7'E, sand, 3 m, coll. Marine Pollution
Studies Group, 11 Jun 1971, 2 (AM W16236).
Description. Body about 2.4 mm long, 0.2 mm wide, with
30 chaetigers. Prostomium oval, slightly wider than long; 4
eyes arranged in open trapezoidal pattern and 2 anterior
eyespots; median antenna inserted between posterior eyes,
about one and half times combined length of prostomium
and palps, lateral antennae about two thirds of length of
median antenna, inserted near anterior eyes (Fig. 90A). Palps
similar in length to prostomium, without distal papillae.
Peristomium slightly shorter than subsequent segments, with
hyaline, rounded inclusions; dorsal tentacular cirri similar
to median antenna, slightly shorter; ventral tentacular cirri
about two thirds length of dorsal ones (Fig. 90A). Antennae,
tentacular, and dorsal cirri of chaetigers 1 and 2, elongate,
slender; dorsal cirri of chaetiger 1 longer than median
antenna, those of chaetiger 2 shorter than those of chaetiger
1, similar in length to median antenna. Dorsal cirri of
subsequent segments alternating irregularly with long cirri,
distinctly longer than body width, with about 10–12 articles;
short cirri, similar in length to body width, with about 6–7
articles; articles pyriform, usually with 1 dark inclusion (Fig.
90A). Parapodial lobes elongated, conical, distally bilobed
(Fig. 90C). Ventral cirri digitiform, similar in length to
parapodial lobes. Compound chaetae similar throughout,
shafts with several, long, fine, subdistal spines, and
elongated, bidentate blades, within fascicle 5 pairs of
chaetae, each pair similar in shape and length (Fig. 90D–
G). Blades of most dorsal pair of compound chaetae with
single long, distinct basal spur, and moderate, thin spines
on margin, about 60–62 µm long; blades of second pair
similar to those of most dorsal pair, but shorter, about 52
µm long, with shorter spines on margin (Figs 90D, 86A).
Blades of subsequent pairs without basal spur and having
short spines on margin (Fig. 90E–G), 40, 27 and 15 µm in
length respectively. Dorsal simple chaetae from chaetiger
1, unidentate, distally blunt, provided with distal hood and
short spines on margin (under light microscope) (Fig. 90H);
under SEM, hood consists of several rows of minute spines
(Fig. 86B). Ventral simple chaetae absent. Acicula solitary,
distally knobbed (Fig. 90I). Pharynx through 5–6 segments
(Fig. 83A). Proventricle barrel-shaped, through about 5–6
segments, with 35 muscle cell rows. Pygidium small, semi-
circular, with 2 short, smooth lateral anal cirri and 1 median,
longer cirrus (Fig. 90B).
Remarks. This species has been reported in Australia as
Syllides longocirrata (Örsted, 1845), which only occurs in
the Northern Atlantic and Northern Pacific areas, and differs
from the Australian specimens, which are described as a
new species. Syllides longocirrata has dorsal simple chaetae
that are pointed and without a hood, and the compound
chaetae are unidentate (see Banse, 1972), whereas the
compound chaetae of Syllides tam are distally blunt and
hooded. Syllides japonicus Imajima, 1966 (see below) and
S. floridanus Perkins, 1981, also have dorsal simple chaetae
that are pointed, and without a hood (Perkins, 1981), which
differ from those present in Syllides tam.
Habitat. Occurring in coarse, medium to muddy sand; from
intertidal to 60 m.
Distribution. Australia (Queensland, New South Wales,
Victoria).
Etymology. The species is named after The Australian
Museum.
Syllides sp.
Fig. 91A–D
Material examined. AUSTRALIA: WESTERN AUSTRALIA: N end of
beach, Bundegi Reef, Exmouth Gulf, 21°49'S 114°11'E, rocky rubble &
sticky sediment with brown & epiphytic algae, 2 m, coll. H.E. Stoddart,
4 Jan 1984 (AM W28920).
Description. Body 2.0 mm long, 0.2 mm wide, with 25
chaetigers. Dorsum of each segment with 2–4 transverse
rows, more or less well defined, of small, golden inclusions
(Fig. 91A). Prostomium trapezoidal, with 2 distinct tufts of
long cilia on anterior corners, and 4 eyes in open trapezoidal
arrangement (Fig. 91A); antennae detached. Palps slightly
shorter than prostomium. Peristomium similar in length to
following segments, tentacular cirri missing. Dorsal cirri
of chaetigers 1 and 2 smooth, elongated, longer than body
width; remaining dorsal cirri articulated, long, up to 11
articles, with 1 dark inclusion of granular material in most
articles (Fig. 91A). Compound chaetae with translucent
hooded shafts serrated on edge, and relatively short,
bidentate blades, with short spines on margin, and
distributed in 5 pairs within fascicle, each pair of similar
length (Fig. 91B); dorsal most pair 26 µm in length and
ventral most pair 15 µm in length. Dorsal simple chaetae
from chaetiger 1, slender, slightly bifid, with short spines
on margin, lacking distal hood (Fig. 91C). Acicula solitary,
distally knobbed, with distal, filiform tip (Fig. 91D).
Pharynx long and slender, through 5 segments. Proventricle
short, slender, through 2 segments (Fig. 91A), with about
17 muscle cell rows.
Remarks. This specimen is characterized by having a hood
on the edge of shafts of compound chaetae, and blades
lacking spurs. The most similar species is Syllides gomezi
San Martín, 1990, from Cuba (San Martín, 1990). The
Cuban species, however has a longer proventriculus through
4 segments, instead of through 2 as occurs in Syllides sp.,
and the dorsal simple chaetae are entire, distally pointed,
whereas they are bifid in the Australian species. Other
species of Syllides also lack basal spurs on the blades of
compound chaetae, but none have a hood on the edge of
shafts. Syllides sanyaensis Ding & Westheide (1997)
described from China, is similar, but the dorsal simple
chaetae are hooded whereas those of this specimen lack
such hoods. Since we have a single, somewhat damaged
specimen, we prefer not to describe this species as a new
species.
Habitat. Rocky rubble and sediment with brown and
epiphytic algae, in depths of 2 m.
Distribution. Australia (Western Australia).

366 Records of the Australian Museum (2006) Vol. 58
Fig. 91. Syllides sp. (A) anterior end, dorsal view; (B) compound
chaetae (one of each pair), midbody parapodium; (C) dorsal simple
chaeta; (D) aciculum. AM W28361. Scales: A 0.18 mm, B–D 20 µm.

San Martín & Hutchings: Eusyllinae of Australia 367
ACKNOWLEDGMENTS. This paper was possible by a Visiting
Fellowship from The Australian Museum to the senior author;
we want to express our gratitude to the Museum authorities for
the grant, and all colleagues that gave their help and assistance
during, before and after the stay in Sydney of the senior author.
Also, funds of the Universidad Autónoma de Madrid, Spain,
helped in the travel and stay. A grant from the Spanish Ministerio
de Educación, Cultura y Deporte (Programa de Movilidad
Salvador de Madariaga, ref. PR2002-207) and the Project of the
Spanish Ministerio de Ciencia y Tecnología, BOS2003-01322,
provided financial support. Dr Penny Berents facilitated the study
of the collection. Kate Attwood and Anna Murray did the rough
sorting of the material, previously only identified to family level,
and extracted the specimens of the subfamily Eusyllinae, and,
together with Keyne Monro, managed the collection and checked
the material examined sections for us. Previous work extracting
syllids was mostly done by volunteers. Richard Johnson helped
us with the literature. The comments and suggestions of two
anonymous referees, as well as the efforts of the editor, greatly
improved the quality of the paper. Ms Miranda Lowe, British
Museum Natural History, London (UK), Dr Angelika Brandt and
Gisella Wegener, Hamburgische Zoologische Museum, Hamburg
(Germany) kindly assisted the senior author during his stay in the
Hamburgische Museum to examine specimens for comparison.
We want to express also our gratitude to Miss Yolanda Lucas,
who collaborated on the figures 5, 7, 40, 57, 74, and 80. Dr
Esperanza Salvador (SIDI of the UAM), and Sue Lindsey (AM)
who assisted in the SEM study and photographs.
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