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PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 116-123
REVISION OF THE FAMILY LYSARETIDAE, AND
RECOGNITION OF THE FAMILY
OENONIDAE KINBERG, 1865
G. Kent Colbath
Abstract. — As recognized by previous authors, the family Lysaretidae Kin-
berg, 1865 includes genera with dissimilar jaw morphology and composition.
Lysaretidae is restricted here to include only Lysarete Kinberg, characterized
by jaws mineralized with calcite and relatively short carriers with transverse
musculature. Oenone Savigny, Halla Costa and Tainokia Knox & Green are
removed into the family Oenonidae Kinberg, 1865. These genera are charac-
terized by heavily sclerotized jaws which are not mineralized, and by long
carriers with oblique -longitudinal musculature.
Hartman (1944) published a family-level
revision of the eunicoid polychaetes which
was widely accepted. Drawing on the work
of Kinberg (1865) and Ehlers (1868), she
placed heavy emphasis on the configuration
of the jaws as a taxonomic character.
Hartman recognized three basic maxil-
lary jaw patterns within the superfamily Eu-
nicea: the labidognath pattern ascribed to
the Eunicidae, Onuphidae, and Lumbriner-
idae; the prionognath pattern in the
Arabellidae and Lysaretidae; and a third
pattern (later named ctenognath by Kielan-
Jaworowska 1966) found only in the Dor-
villeidae. Labidognath jaws are character-
ized by having two relatively short, broad
carriers, in contrast to the two long, slender
carriers with a ventral median ligament (or
"third carrier") which characterize priono-
gnath jaw apparatuses. In the ctenognath
condition the maxillae consist of two to four
rows of numerous separate denticles.
During a study of the composition and
fossilization potential of polychaete jaws, I
divided eunicoid jaws into three groups
based on their composition (Colbath 1986).
Labidognath jaws are mineralized with cal-
cium carbonate in the form of aragonite in
the Onuphidae and Eunicidae, and calcite
in the Lumbrineridae. Labidognath jaws are
also weakly sclerotized, and the outer pro-
tein wall is readily leached in 2% KOH
(Hartmann-Schroder 1967, Colbath 1986).
In contrast, prionognath and ctenognath
jaws are amorphous to x-rays, do not react
with HC1, and are relatively resistant to
leaching in KOH (Colbath 1986).
The jaws of species of Lysarete Kinberg,
1865 are mineralized with calcite (Colbath
1986), and exhibit other labidognath mor-
phological characters. Other genera previ-
ously included within the Lysaretidae have
jaws of the prionognath type, prompting a
revision of the family, and recognition of
the family Oenonidae Kinberg, 1865.
History of Study
Early classifications of the eunicoid poly-
chaetes were based exclusively on features
of the external anatomy, and produced ar-
bitrary groupings of genera which are dis-
similar internally (Hartman 1944:1). First
to emphasize the importance of jaw parts
in classification was Kinberg (1865). Kin-
berg recognized ten families which he di-
vided into four major groups (labeled in-
formally as A-D). He defined group C by
the presence of mandibles with separated
right and left halves (an ontogenetically
variable feature), and included only La-
VOLUME 102, NUMBER 1
randa Kinberg, 1865, which was considered
unrecognizable by Hartman ( 1 944: 1 0, 1 948:
98). Kinberg's other groups correspond to
the labidognath (A), prionognath (B), and
ctenognath (D) jaw types presently recog-
nized. Kinberg considered the Lysaretae (sic)
and Oenonidea (sic) distinct, including the
former in group A and the latter in group B.
Ehlers (1868:280-282) also emphasized
the importance of jaw configuration in clas-
sifying these polychaetes, but did not for-
malize the distinctions. Instead, he recog-
nized a single family Eunicea (sic), which
he informally divided into the Eunicea lab-
idognatha and Eunicea prionognatha. Eh-
lers (1868) did not stress the morphology of
maxillary carriers as strongly as did Kin-
berg, but rather subdivided the eunicoids
according to whether the anterior maxillae
are arranged in a semicircle (labidognatha)
or in parallel rows (prionognatha).
In contrast to Kinberg, Ehlers (1868)
placed the genus Lysarete close to Aglau-
rides Ehlers (=Oenone Savigny) in the
prionognath subdivision. He later (Ehlers
1887:107-108) acknowledged the close
similarity between the jaws of Lysarete and
those of Lumbriconereis Grube (=Lumbri-
neris Blainville), but did not emend his ear-
lier taxonomic treatment.
Hartman (1944:2) essentially formalized
the 1 868 scheme of Ehlers using the family-
level nomenclature of Kinberg. She retained
six of the ten families recognized by Kin-
berg, placing the Oenonidae into synonymy
with the Lysaretidae.
Fauchald (1970:118) removed Iphitime
Marenzeller from the Lysaretidae, and
erected the family Iphitimidae. He later
(Fauchald 1977) erected the order Eunicida
to include the seven families within the su-
perfamily Eunicea, plus two additional fam-
ilies (Histriobdellidae, Ichthyotomidae)
which were not assigned to a superfamily.
Materials and Methods
All lysaretids in the collection of the U.S.
Museum of Natural History (USNM) were
examined for the present study, and selected
specimens were dissected for examination
of the jaws. Additional specimens of Ly-
sarete were obtained on loan from the Flor-
ida Department of Natural Resources
(FSBC). The holotype of L. brasiliensis Kin-
berg, 1865 was obtained from the Swedish
National Museum, Stockholm.
One specimen of Lysarete brasiliensis with
jaws in excellent condition (USNM 55884)
was used for the chemical and mineralogical
analyses, and for scanning electron micros-
copy. The maxillary apparatus was re-
moved, dehydrated in an alcohol series, and
prepared for SEM by critical point drying.
The specimen was initially coated with car-
bon only, and chemically analyzed using
x-ray dispersion. The specimen was then
coated with gold palladium for routine scan-
ning electron microscopy. The mandibles
were removed, photographed with a Wild
400 wide-field optical microscope, and ana-
lyzed with the Gandalfi x-ray spectrometer
in the Department of Mineralogy, Smith-
Jaw morphology and composition. —X-ray
dispersion analysis of the maxillae of Ly-
sarete brasiliensis indicates the presence of
abundant calcium carbonate. The mandi-
bles are mineralized with calcite, as are jaws
in the lumbrinerids (Colbath 1986). Calci-
fication is also evident in L. australiensis
Benham, 1915, in that the blades of the
mandibles are white, and the maxillae are
dark brown rather than black. The presence
of calcite is inferred by comparison with L.
In contrast to Lysarete, jaws in species of
Oenone Savigny, Halla Costa and Tainokia
Knox & Green (commonly included in the
Lysaretidae) are not mineralized, but rather
are composed almost exclusively of sclero-
protein. As a consequence, both maxillae
and mandibles are jet black in color. The
distinction between heavily sclerotized and
calcified jaws is apparent upon inspection,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and was described by Benham (1915:233,
236-237). Benham made no comment about
the utility of this distinction above the
species level, however.
Other aspects of jaw morphology are cor-
related with the difference in mineraliza-
tion. In Lysarete the maxillary carriers are
relatively short (approximately equal to the
length of the maxillae) and broad (Fig. lb),
with no ventral ligament (= "third carrier").
A thick band of transverse muscles joins the
carriers dorsally (Fig. 2f ), allowing them to
be pulled closed in the "trap" motion de-
scribed by Wolf (1980) for other labido-
gnath jaws. The MI (maxilla I) elements in
the maxillary apparatus are the same length
(Fig. 2c), which allows the anterior tips to
come together in a pincer-like fashion as
part of the trap motion. The posterior dorsal
surfaces of the Mis exhibit an interlocking
ridge and furrow system (Fig. 2e), which
may help to lock the apparatus in a closed
position (Wolf 1980).
In species of Oenone, Halla and Tainokia
the carriers are narrow and much longer than
the combined length of the maxillae, and
are not equiped with a transverse muscle
band. Instead, a series of long, oblique-lon-
gitudinal muscles are attached along the
length of the carriers. A ventral ligament is
present. The right and left MI elements may
be of different lengths (Crossland 1924;
Hartman 1944; Kielan-Jaworowska 1966),
and no basal ridge and furrow system is
present to lock the jaws in a closed position.
Such jaws are presumably incapable of per-
forming a trap motion, as Wolf (1980) also
concluded for the arabellids.
Soft-part morphology. —The genera tra-
ditionally included within the Lysaretidae
share an elongated post-setal lobe and
prominent dorsal cirrus on posterior para-
podia. The morphology of the head, how-
ever, is distinctly different in each of the
four genera under discussion. Although the
difference between Lysarete and the other
genera may not merit consideration as a
family-level character, the anterior mor-
phology is none-the-less a useful key char-
In Lysarete, three antennae emerge from
the posterior margin of the prostomium, and
may lie flat in a V-shaped notch (Fig. la).
In adult specimens the notch extends pos-
teriorly across the peristomial rings, and in
some cases across the first one or two seti-
gers. Halla has three antennae which may
lie back in a semicircular notch in the two
peristomial rings (Fig. le). Tainokia has a
single antennae, and a slight invagination
which extends posteriorly only partially
across the first of two peristomial rings
(Knox & Green 1972, fig. 1). In Oenone,
three antennae are tucked into a nuchal fold
(Fig. Id), no notch is present, and dorsally
the peristomium has only one distinct ring.
Setae and acicula.— All of the worms
considered here have simple geniculate se-
tae. In addition, bifid hooks have been re-
ported in Oenone and Halla. In Lysarete,
Oenone, and Tainokia the geniculate setae
are smooth (Knox & Green 1972, figs. 6, 7;
pers. observ.). In Halla, however, the dorsal
setae are denticulate, and the ventral setae
are marked by distinctive oblique striae (Fig.
2a, b; Fauvel 1923:427, fig. 169 g, h).
Both described species of Lysarete have
black acicula. In Oenone fidgida (Savigny,
1818) and Halla parthenopeia (Chiaje, 1828)
the acicula are amber, while Knox & Green
(1972:433) described the acicula in Taino-
kia iridescens as colorless. Black acicula are
thus useful for distinguishing Lysarete from
the other three genera, supporting the fam-
ily-level distinction based on jaw morphol-
Order Eunicida Fauchald, 1977
Family Lysaretidae Kinberg, 1865
Emended diagnosis, — Eunicoid poly-
chaetes with jaws mineralized with calcite.
Mandibles flat with semicircular growth
rings anteriorly (Fig. 2d). Maxillary carriers
no longer than combined length of maxillae,
VOLUME 102, NUMBER 1
Fig. 1 . a-c, Lysarete brasiliensis, holotype, Brazil; a, Dorsal view, anterior end, illustrating V-shaped notch
in peristomium; b. Dorsal view, MI elements and carriers, muscle attachment stippled; c, Labial view, MIIL-
MVL. d, Oenonefulgida, USNM 17729, Dry Tortugas, dorsal view, anterior end, note tips of antennae protruding
from nuchal fold, e, Hatta parthenopeia, USNM 5 147, Bay of Naples, dorsal view, anterior end, note semicircular
notch in peristomium. All scale bars = 1 mm.
articulated by prominent transverse mus-
cular bundle, no ventral ligament. Maxillary
jaws symmetrical. MI elements of identical
length, distally falcate with prominent sec-
ondary fang, additional small denticles may
be present at base. Three short antennae
arise from base of prostomium. Elongate
post-setal lobe and flattened dorsal cirrus
(or notopodium) developed in posterior
parapodia. Simple geniculate setae present,
bifid hooks absent.
Comparison.— The Lysaretidae are dis-
tinguished from the Oenonidae based on
jaw morphology and mineralogy. Lysarete,
the only genus presently recognized within
the family, is further distinguished from the
Oenonidae in having black acicula, and a
V-shaped notch in the dorsal surface of the
The Lysaretidae are similar to the Lum-
brineridae in having: 1) jaws mineralized
with calcite; 2) flat mandibles with a semi-
circular blade; 3) symmetrical maxillary
jaws; 4) transverse musculature on the car-
riers. The Lysaretidae are distinguished from
the Lumbrineridae by: 1) well-developed
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. a, b, Halla panhenopeia, USNM 5147, Bay of Naples; a, SEM dentate setae from dorsal bundle,
posterior parapodium, scale bar =10 fim; b, SEM striate setae from ventral bundle, scale bar = 50 (im. c-g,
Lysarete brasiliensis, USNM 55884, Gulf of Mexico; c, SEM maxillary apparatus, dorsal view, scale bar = 1
mm; d, Light photomicrograph mandibles, ventral view, scale bar = 1 mm; e, SEM MI elements, oblique-lateral
view, note ridges and grooves on bases of Mis, scale bar = 0.5 mm; f, SEM carriers, oblique-lateral view, carriers
canted at an angle to Mis, note transverse muscle band, scale bar = 0.2 mm; g, SEM anterior maxillary jaws,
oblique-lateral view, scale bar = 0.2 mm.
dorsal cirri; 2) a prominent secondary fang
on the MI elements of the maxillae; 3) the
absence of hooded hooks. Kuwaita Moham-
mad, 1973 is intermediate between the Ly-
saretidae and Lumbrineridae in having an-
tennae on the prostomium, but clearly
belongs in the Lumbrineridae based on the
3 criteria above (Mohammad 1973:34, 36,
figs. 6, 7).
Distribution.— Lysarete includes one
VOLUME 102, NUMBER 1
species recorded from the east coast of North
and South America in low latitudes, and a
second species from a single locality east of
Australia. Larymna Kinberg, 1865 may also
belong in the Lysaretidae based on jaw mor-
phology (Hartman 1948:99). The type and
only specimen (from Mozambique) is so
poorly preserved that, according to Hart-
man, attribution cannot be made below the
family level, and Larymna should be re-
garded as unrecognizable.
Genus Lysarete Kinberg, 1865
Type species.— L. brasiliensis Kinberg,
Lysarete brasiliensis Kinberg, 1865
Figs, la-c, 2c-g
Lysarete brasiliensis Kinberg 1865:570;
1910:49, pi. 17,fig.30.-Ehlers 1887:107-
108, pi. 33, figs. 1-8.— Rioja 1944:130-
131. figs. 47-50.— Hartman 1951:65, pi.
14, figs. 7, 8.-Orensanz 1975:106-108,
fig. 8. -Gardiner 1976:213, figs. 29b-e.-
Gilbert 1984:43-6-43-7, figs. 43-3, 43-4.
lOenone brevimaxillata. — Treadwell 1931:
1-3, figs. 4-8.
Remarks. — In her re-appraisal of Kin-
berg's type material, Hartman (1948:8) did
not consider the holotype of Lysarete bra-
siliensis, but instead referred to an earlier
revision by Ehlers (1887:107-108, pi. 33).
Ehlers did not figure the Brazilian type of
L. brasiliensis, but rather figured a specimen
collected from Florida. The posthumous
publication of Kinberg's plates (Kinberg
1910) did not include illustrations of the
jaws of the holotype. The maxillae are there-
fore illustrated here (Fig. lb, c).
On the holotype, maxillary jaws III and
IV each have a prominent anterior denticle,
with two progressively smaller denticles
posteriorly. The jaws in a specimen from
Argentina (Rioja 1944, fig. 50) are similar,
except that the Mill elements have four
denticles instead of three.
Most specimens examined from the east
coast of the United States have five to six
denticles on the Mill and four to seven den-
ticles on the MIV elements (Fig. 2g). The
anterior three denticles on the Mill element
are sub-equal in length, unlike those of the
One specimen from Florida (FSBC I
31008) has Mill jaws like the type, how-
ever, and a specimen from Argentina illus-
trated by Orensanz( 1975, fig. 8-3) has Mills
and IVs indistinguishable from most of the
North American specimens. There is thus
some overlap in jaw morphology between
the North and South American populations.
The number and shape of denticles on an-
terior jaws is variable in eunicoid poly-
chaetes, and the two populations are ten-
tatively considered conspecific in spite of
their disjunct geographic ranges.
The type and only specimen of Oenone
brevimaxillata Treadwell, 1931 was never
deposited (H. S. Feinberg, AMNH. pers.
comm. 1985), and is presumed lost. Tread-
well's (1931, fig. 8) illustration of the max-
illae is schematic, and insufficient for de-
Family Oenonidae Kinberg, 1865
Emended diagnosis. —Jaws jet black,
composed primarily of scleroprotein, no
crystalline mineral phases present. Mandi-
bles H-shaped. Maxillary carriers much
longer than anterior jaws, attached to bun-
dles of oblique-longitudinal muscles, ven-
tral ligament present. Maxillae commonly
asymmetrical, Mis differ slightly to mark-
edly in length. One or three antennae arise
from base of prostomium. Elongate post-
setal lobe and prominent dorsal cirri de-
veloped on posterior parapodia. Geniculate
setae present, bifid hooks present or absent.
Key to the genera of Oenonidae
la. One or 3 antennae recline in notch
in peristomium, 2 peristomial rings
dorsally, bifid hooks present or ab-
1 b. Three antennae tucked into nuchal
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
fold, 1 peristomial ring dorsally,
bifid hooks present Oenone
2a. One antenna, peristomial notch
weakly developed, bifid hooks ab-
sent, setae smooth Tainokia
2b. Three antennae, distinct semicir-
cular peristomial notch, bifid hooks
present (small individuals only),
dorsal setae dentate, ventral setae
Genus Halla A. Costa, 1844
Type species. —Halla parthenopeia
(Chiaje) Costa, 1844:63.— Lysidice parthe-
nopeia Chiaj e, 1828:175.
Remarks. — Fauvel (1923:427, fig. 169f)
described Halla parthenopeia as having bi-
fid hooks, in addition to the setae figured
here (Fig. 2a, b). Hooks are absent in para-
podia from the large specimen of H. par-
thenopeia examined here (USNM 5 147), nor
were hooks recorded from the large speci-
men figured by Ehlers (1868:28, figs. 27,
28). The type of Halla okudai Imajima
(1967:437) is also a large worm, and lacks
hooks (Imajima & Hartman 1964:267-268).
Individuals of Halla which attain large size
may lose their bifid hooks. A similar con-
dition prevails in several large species with-
in the Eunicidae (K. Fauchald, pers. comm.
Distribution. — All western hemisphere
specimens deposited in the USNM collec-
tion as Halla proved to be misidentified
members of Oenone or Lysarete. Halla may
be restricted to the Old World, with docu-
mented reports from the Mediterranean
(Fauvel 1923) and Japan (Imajima 1967).
I thank Kristian Fauchald for extensive
discussions about the taxonomy of eunicoid
polychaetes, and for critically reading the
manuscript. Thomas H. Perkins arranged
the loan of material from the Florida De-
partment of Natural Resources. H. S. Fein-
berg, American Museum of Natural His-
tory, provided information concerning the
type of Oenone brevimaxillata Treadwell.
This research was supported by a post-doc-
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