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\3 Biodiversity 

Proceedings of the Biological Society of Washington. 

Washington, Biological Society of Washington 

v. 102 Mar-June 1989: 
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102(1), 1989, pp. 116-123 





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- 



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- 
sonian Institution. 

Taxonomic Characters 

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, 



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- 

Systematic s 

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, 




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 



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 



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- 
tailed comparison. 

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- 
sent 2 

1 b. Three antennae tucked into nuchal 



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- 
toral fellowship, Smithsonian Institution. 

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