MARIAN H. PETTIBONE ReVtSWU of the

Aphroditoid Polychaetes

the Family

Eulepthidae Chamberlin

(=Eulepidinae Darboux,

=Pareulepidae Hartman)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 41
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Smithsonian Institution
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 41

Marian H. Pettibone Revision o f the

Aphroditoid Polychaetes

of the Family

Eulepethidae Chamberlin

(=Eulepidinae Darboux;

=Pareulepidae Hartman)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1969
ABSTRACT

Pettibone, Marian H. Revision of the Aphroditoid Polychaetes of the Family

Eulepethidae Chamberlin (=Eulepidinae Darboux; = Pareulepidae Hartman).

Smithsonian Contributions to Zoology, 41: 1-44. 1969.—The family Eulepethidae is reviewed and revised, based on a reexamination of most of the previous

records in the literature. The family characteristics are summarized and its systematic position among the other families of the superfamily Aphroditoidea is

discussed.

The family is represented by four genera: Eulepethus Chamberlin (=Eulepis

Grube), with a single species; Pareulepis Darboux, including two species and one

synonym; Mexieulepis Rioja, with a single species and one synonym; and Grubeulepis, new genus, including four previously described species and three new species.

In addition, Eulepis challengeriae Mclntosh is considered questionable.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.
Marian H. Pettibone Revision of the

Aphroditoid Polychaetes

of the Family

Eulepethidae Chamberlin

(=Eulepidinae Darboux;

=Pareulepidae Hartman)

Among the scaled polychaetes of the superfamily

Aphroditoidea, the family Eulepethidae consists of

relatively few, widely distributed species. The

earlier descriptions of the species by Grube (1875),

Mclntosh (1885), Treadwell (1901), Horst (1913,

1922), Fauvel (1918), and Berkeley and Berkeley

(1939) were deficient and misleading in many respects. A few additions and corrections for some of

the species were made subsequently by different

authors. Recent descriptions of new species by

Rioja (1961) and Jones (1962) are more complete,

but comparisons with previously described species

are limited by deficiencies in the literature. A reexamination of these species was necessary to fill in

the gaps and clarify the terminology.

The genera and species of Eulepethidae have had

a checkered history, as indicated by the following

brief summary. The first species of this family was

described by Grube (1875, 1878), as Eulepis hamijera, who placed it in the Aphroditeen: Sigalionina.

Mclntosh (1885) added two species of Eulepis and

gave a misleading description of the arrangement of

the elytra for one of them, E. wyvillei, and indicated

that Eulepis required the institution of a new

family, although he placed it in the Polynoidae.

Darboux (1900) established the tribe Eulepidinae in

Marian H. Pettibone, Curator, Division of Worms, Museum

of Natural History, Smithsonian Institution, Washington, D.C.

20560.

the Aphroditiens for Eulepis Grube and the new

genus Par eulepis for P. wyvillei (Mclntosh). Later,

two species of Eulepis were added by Treadwell

(1901). Horst (1913, 1917, 1922) added two species

and Fauvel (1918, 1919) described an additional

species of Eulepis. Augener (1918) raised Darboux's

tribe to family status—Eulepidae [sic]. Chamberlin

(1919) replaced Eulepis Grube, preoccupied, with

the new name Eulepethus and substituted Eulepethinae Chamberlin for Eulepidinae Darboux, including Eulepethus and Pareulepis; at the same

time, Chamberlin transferred the subfamily to the

Sigalionidae.

The Berkeleys (1939) described an additional

species of Eulepethus. Hartman (1939) referred

Eulepethus Chamberlin to Pareulepis Darboux and

proposed a separate family Pareulepidae, based on

the single genus Pareulepis. Fauvel (1940: 1) continued to use Eulepis and Eulepidinae, preferring

not to follow the Law of Priority, with the argument that there was little chance of confusing an

aphroditid with an insect or a reptile. More recently, Rioja (1961) added a new genus and species,

Mexieulepis elongatus, and Jones (1962) added a

new species of Pareulepis.

As indicated above, there has been no general

agreement regarding the family or subfamily among

the scaled polychaetes or Aphroditoidea to which

the various species have been referred. Thus, they

have been placed in the following:

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Aphroditeen: Sigalionina, by Grube (1875, 1878);

Polynoidae, by Mclntosh (1885), Treadwell (1901), and

Augener (1906);

Aphroditiens or Aphroditidae: Eulepidinae, by Darboux

(1900), Horn (1917, 1922), and Fauvel (1919);

Aphroditidae (sensu lato), by Horst (1913), Treadwell (1920),

Fauvel (1939, 1940), Tebble (1955), and Rullier (1965);

Eulepidae [sic], by Augener (1918, 1927);

Sigalionidae: Eulepethinae, by Chamberlin (1919);

Aphroditidae: Polynoinae, by Pruvot (1930), and Day (1951,

1962, 1967);

Sigalionidae, by Treadwell (1939), and Berkeley and Berkeley (1939);

Pareulepidae, by Hartman (1939, 1942, 1944, 1959, 1961),

Rioja (1961), Jones (1962), and Reish (1968).

In the present study, three genera are retained:

Eulepethus Chamberlin (=Eulepis Grube), Pareulepis Darboux, and Mexieulepis Rioja. In addition,

a new genus Grubeulepis is proposed. There is good

basis for separating the species in a family distinct

from the other Aphroditoidea. Eulepethidae Chamberlin is retained, since the nominal genus is Eulepethus Chamberlin, a replacement for Eulepis Grube,

preoccupied. It also predates Pareulepidae Hartman, based on Pareulepis Darboux, the oldest available generic name in the family but with a different

type-species.

Eleven species of eulepethids have been described

previously. Type-material was available for eight

of them. For two of the species for which no typematerial was available, specimens from near the

type-locality, and which seemed to agree with the

original descriptions, were studied. Of the eleven

species, two are considered herein to be synonyms

of previously described species and one is considered

questionable. To the remaining eight species, three

new species are added, based on the reexamination

of some specimens identified as Eulepis or Pareulepis geayi and P. fimbriata. Those two species do

not appear to be as widely distributed as their

records in the literature indicate.

In addition to the specimens of Eulepethidae

deposited in the United States National Museum

(USNM), material was obtained from the following

Museums: Allan Hancock Foundation, Los Angeles

(AHF), through O. Hartman; American Museum of

Natural History, New York (AMNH), through E.

Kirsteuer; British Museum (Natural History), London (BMNH), through J. D. George; Museum of

Comparative Zoology, Harvard (MCZ), through H.

W. Levi; Museum National d'Historie Naturelle,

Paris (MNHNP), through J. Renaud-Mornant;

Rijksmuseum van Natuurlijke Historie, Leiden

(RNHL), through J. van der Land; Zoologische

Museum Universiteit van Amsterdam (ZMA),

through S. van der Spoel; Zoologisches Staats-museum, Hamburg (ZMH), through G. HartmannSchrdder. Additional specimens were received from

D. J. Reish, California State College, Long Beach,

and from F. Rullier, Universite Catholique, Angers,

France. Information concerning types was received

from G. Hartwich of the Zoologisches Museum,

Berlin and from M. E. Caso Munoz of the Instituto

de Biologia Universidad Nacional Aut6mona de

Mexico.

I am indebted to the above-mentioned people for

their cooperation and help and for allowing me to

examine the eulepethid material upon which this

study is based. The manuscript benefited from the

suggestions of F. A. Chace, Jr., and M. L. Jones,

both of the Smithsonian Institution. This study was

aided in part by a grant from the National Science

Foundation (GB-1269).

The following genera and species are recognized

in this study (synonyms in parentheses):

Genus Eulepethus Chamberlin, 1919

(=Eulepis Grube, 1875, preoccupied)

E. hamifer (Grube)

(=Eulepis hamifera Grube, 1875)

Genus Pareulepis Darboux, 1900

P. wyvillei (Mclntosh)

(=Eulepis wyvillei Mclntosh, 1885)

(=Eulepis splendida Treadwell, 1901)

P. malayana (Horst)

(=Eulepis malayana Horst, 1913)

Genus Mexieulepis Rioja, 1961

M. weberi (Horst), new combination

(=Eulepis weberi Horst, 1922)

(=Mexieulepis elongatus Rioja, 1961)

Grubeulepis, new genus

G. fimbriata (Treadwell), new combination

(=Eulepis fimbriata Treadwell, 1901)

G. geayi (Fauvel), new combination

(=Eulepis geayi Fauvel, 1918)

G. ecuadorensis, new species

(=Pareulepis fimbriata.—Hartman, 1939)

G. mexicana (Berkeley and Berkeley), new combination

(=Eulepethus mexicanus Berkeley and Berkeley, 1939)

G. sulcatisetis (Jones), new combination

(=Pareulepis sulcatisetis Jones, 1962)

G. tebblei, new species

(z=Pareulepis geayi.—Tebble, 1955)

G. augeneri, new species

(=Eulepis fimbriata.—Augener, 1918)

(=Eulepis geayi.—Fauvel, 1940)

(=Pareulepis fimbriata.—Rullier, 1965)

Questionable eulepethid

Eulepis challengeriae Mclntosh, 1885
NUMBER 41

Family EULEPETHIDAE Chamberlin, 1919

(=EULEPIDINAE Darboux, 1900; =Pareulepidae Hartman,

1939)

TYPE-GENUS.—Eulepis Grube, 1875, preoccupied;

= Eulepethus Chamberlin, 1919.

Characteristics of the Family Eulepethidae

The body is depressed, subrectangular in shape; it

may be relatively short, consisting of 32 to 40 segments, or more elongate, with up to 70 segments.

The prostomium and tentacular segment (I) are

distinct; both are withdrawn and wedged between

the anterior segments (II-IV) and are partly hidden

from view; the posterior middorsal part of the prostomium may be attached to segment II (Figures la;

6a,b; lie; 156; 24a). Segment III is not visible middorsally, its para podia being wedged between the

parapodia of segments II and IV. The prostomium

(Figures 16; 4a,6; lie; \bc,h) is globular, with three

short antennae lacking distinct ceratophores; the

median antenna is globular to subulate, attached

antercdorsally, the lateral antennae are conical,

attached anteroventrally on the prostomium. A pair

of elongate-conical palps are attached ventrally on

the prostomium; eyes, when present, are very small,

variable in number, hidden by segment II and

easily overlooked, especially when the prostomium

is pigmented or darkened through preservation. The

tentacular segment (I) forms a pair of small lobes

lateral to the prostomium; they are more slender

basally and inflated distally, each provided with two

acicula, a pair of rather short subulate dorsal and

ventral tentacular cirri and two bundles of capillary

setae (Figures lc,d; 6c,d; \\e,g; 15d,e). Between the

prostomium and tentacular lobes is a pair of clubshaped nuchal organs. Segments II and III contribute to the lateral and posterior lips of the ventral

mouth (Figure 66). The buccal ventral cirri of segment II are longer and thicker than those following

(Figure 66,e). When extended, the strong muscular

pharynx is encircled with soft papillae (about 13

pairs) and two pairs of platelike jaws (somewhat

similar to those in the Aphroditidae; Figure lie,/).

Dorsally the segments may be provided with

either elytra, dorsal cirri, branchiae, or posterior

lamellae. The dorsal cirri occur on segments 3 and

6; they are short, subulate, without distinct rirrophores, and attached posteriorly on the notopodia

(Figures la,/; 7a,c; 12a; 156,g). The anterior 12 pairs

of elytra are located on segments 2, 4, 5, 7, 9, II, 13,

15, 17,19,21, 24 [not on segments 2, 3,4, as reported

by Mclntosh (1885) for Eulepis wyvillei and followed by Darboux and Chamberlin for Pareulepis;

the three anterior pairs of elytra appear at first

glance to be attached on consecutive segments, since

segment III is indistinct middorsally]. The elytra

increase in length progressively, the twelfth pair

being extremely elongate and subrectangular (Figures 3a-c; 4g-i; lla-c; 17d-/). The elytral surfaces are

usually smooth, lacking tubercles, but sometimes

with scattered microtubercles; their external borders

are usually notched or fimbriated (flattened, leaflike

processes). The elytriphores are large, cylindrical,

their bases converging middorsally.

Branchiae occur on segments 8, 10, 12, 14, 16, 18,

20, 22,23,25 (10 pairs), 26 (11 pairs), 27 (12 pairs), or

28 (13 pairs). The bases of the branchiae converge

middorsally and are joined medially by a small

crescent-shaped fold; more distally they are inflated

and provided with a dorsal glandular crest and

ventral ciliated bands which are continuous with

ciliated bands on the dorsal surfaces of the parapodia; bulbous to subconical lobes or branchial

cirri are attached distally (Figures 26, 86, 136, 16a).

The branchiae have sometimes erroneously been referred to as dorsal cirri; they correspond rather to

modified dorsal tubercles on the nonelytra-bearing

segments.

In the posterior regions of the more elongate

species (Eulepethus, Mexieulepis), additional elytra

occur on all segments from 27 or 28 on, differing

from the more anterior elytra by their smaller size

and by their attachment to the elytriphores along

their anterior borders; the bases of these elytriphores

also converge medially (Figures 3d, lid). In the

posterior regions of the shorter species (Pareulepis,

Grubeulepis), beginning on segment 26-29, the

elongate inflated dorsal tubercles are provided distally with flattened, oval to lanceolate lamellae;

basally the dorsal tubercles converge medially but

the medial crescent-shaped fold found in the more

anterior branchiae is lacking; the ventral ciliated

bands are also lacking (Figures Ac; 10a,6; 17a). The

elytriphores with their elytra, the branchiae with

their distal lobes or branchial cirri, and the modified dorsal tubercles with their posterior lamellae,

appear to be homologous structures, for they have

similar positions in relation to the body and are

formed from dorsal coelomic outpocketings. The
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pygidium bears a dorsal anus and is provided with a

single, extremely long anal cirrus, which is minutely

papillated along one side; the anal cirrus is usually

on the right side; the left side bears a small bulbous

rudiment (Figure 4c).

The parapodia are biramous, with the setae all

simple, not compound (Figure la,b). The notopodia

are short, cylindrical, with posterior rounded lobes;

the embedded notoacicula have characteristic

hooked tips. The notosetae are slender, capillary,

smooth, and spinous; beginning on segment III,

additional stout, geniculate notopodial hooks occur,

whose distal tips may be fine or blunt, spatulate

(Figure 2c). The neuropodia are thick, rounded,

and paddle-shaped, and supported by stout neuroacicula which are provided distally, on their anterior sides, with characteristic hammer-shaped chitinous plates. The neurosetae are of several kinds: 1-2

upper pectinate setae (Figure 2d); bilimbate or unilimbate capillaries (Figure 2e); short nonlimbate

capillaries and, in addition, there may be long

slender capillaries and stout acicular neurosetae; the

the upper neurosetae of the posterior region may be

modified in various ways. The anterior ventral cirri

are short and tapered (Figure lf,j); those following

are globular, with filamentous tips (Figure 2b).

Geographic Distribution of Eulepethidae

The family is known from relatively few species

from widely scattered areas in the western Indian

Ocean (Madagascar, Mozambique), Red Sea, western Pacific (Indochina, Philippine Islands, Malay

Archipelago, New Caledonia), eastern Pacific (Southern California, Gulf of California to Ecuador), Gulf

of Mexico, western Atlantic (Bermuda, Georgia,

Florida, Panama, West Indies), eastern Atlantic

(West Africa from Gold Coast, French Congo), and

Adriatic. In depth they extend from low intertidal

to 823 meters. The eulepethids are burrowing forms

and are found in sand, silt, soft mud, with muck,

shells, and coral. One species has been reported as

commensal with a tube-dwelling polyodontid polychaete.

Relations of the Eulepethidae to Other Families of

the Aphroditoidea

The Eulepethidae are unique among the Aphroditoidea in their type of branchiae and branchial

cirri, found only on nonelytra-bearing segments;

the parapodia have characteristic hooked notoacicula and hammer-shaped neuroacicular distal plates,

and stout bent notosetae; the pygidium has a single

long papillate anal cirrus; the prostomium and

tentacular parapodia (I) are withdrawn within the

anterior segments (II-IV) and segment III is fused

middorsally with the adjacent segments.

Among the families of the Aphroditoidea, the

Eulepethidae appears to show closest relationship

to the Aphroditidae (sensu s trie to). Both families

have only simple setae, including capillary and stout

notosetae; platelike jaws in the pharynx; globular

prostomium; setigerous tentacular parapodia (segment I) distinct and not fused with the prostomium.

They differ by having three antennae in the Eulepethidae, one in the Aphroditidae; dorsal cirri

are present only on segments III and VI in the

former but present on all nonelytra-bearing segments in the latter.

The Eulepethidae agrees with the Polynoidae in

having only simple setae and three antennae. They

differ in the shape of the prostomium: globular in

the former, bilobed in the latter; the three antennae

lack distinct ceratophores in the former, but are

present in the latter; dorsal cirri are present only on

segments III and VI in the former but occur on all

nonelytra-bearing segments in the latter; branchiae

are present in the former but absent in the latter.

The Eulepethidae agrees with the Sigalionidae in

having branchiae but they are of different types: in

the former they are modified dorsal tubercles on

nonelytra-bearing segments; in the latter they are

cirriform and found on the dorsal tubercles, as well

as on the elytriphores; dorsal cirri are few in the

former (on segments III and VI) and few or lacking

in the latter (sometimes on segment III). They differ

in that the neurosetae are all simple in the former

but some neurosetae are compound in the latter; the

prostomium and tentacular parapodia (I) are distinct in the former, but often fused in the latter.

Abbreviations Used in the Figures

I-IV, segments

aC, anal cirrus

br, branchia

brC, branchial cirrus

brF, branchial fold

buC, buccal cirrus
NUMBER 41

dC, dorsal cirrus

dTc, dorsal or upper tentacular cirrus

elph, elytriphore

lAn, lateral antenna

mAn, median antenna

nuO, nuchal organ

pa, palp

pLa, posterior lamella

vC, ventral cirrus

vTc, ventral or lower tentacular cirrus

Key to the Genera of Eulepethidae

1. Elytra 12 pairs, increasing in length progressively, followed by smaller foliaceous lamellae

posteriorly 2

1'. Elytra more than 12 pairs, anterior 12 pairs increasing in length progressively, followed by

smaller pairs of elytra continuing to posterior end 3

2. Elytra with lateral borders notched (Figure 4g-i) Pareulepis Darboux

2'. Elytra with lateral borders fimbriated (flattened, leaflike processes, (Figure Yld-f)

Grubeulepis, new genus

3. Elytra with lateral borders notched Eulepetkus Chamberlin

(—Eulepis Grube, preoccupied)

3'. Elytra with lateral borders fimbriated (flattened leaflike processes) . . Mexieulepis Rioja

Genus Eulepethus Chamberlin, 1919

Eulepis Grube, 1875. [Type-species: E. hamifera Grube, 1875,

by monotypy. Gender: feminine. Preoccupied by (Dalman

manuscript) Billberg, 1820 (Lepidoptera).]

Eulepethus Chamberlin, 1919. [New name for Eulepis Grube,

preoccupied. Gender: masculine.]

DIAGNOSIS.—Body elongate, segments about 60

(60-70). Elytra numerous pairs; first 12 pairs larger,

with lateral notch, on segments 2, 4, 5, 7 . . . 21, 24;

smaller elytra with margins entire, on all segments

beginning with segment 28; elytriphores fused

medially. Dorsal cirri on segments 3 and 6. Branchiae 12 pairs, on segments 8, 10 .. . 22, 23, 25, 26,

27.

Eulepethus hamifer (Grube)

FIGURES 1-3

Eulepis hamifera Grube, 1875, p. 71; 1878, p. 52, pi. 3: fig. 8.

—Fauvel, 1939, p. 261.

Eulepethus hamifera.—Treadwell, 1920, p. 592.

Pareulepis hamifera.—Hartman, 1959, p. 123.

MATERIAL EXAMINED.—Albatross station D5235,

east coast of Mindanao, Philippine Islands, 80

meters, soft mud bottom, 9, May 1908—1 specimen

(USNM 17483).

DESCRIPTION.—Length 29 mm (37-40 mm—Grube,

Fauvel), width 6 mm, including setae, segments

about 60 (up to 70—Fauvel). Anterior 12 pairs elytra

becoming more elongate progressively, each with

lateral notch (first pair lacking on specimen

examined; Figure $a-c); elytriphores large, inflated,

converging middorsally (Figure la). Smaller elytra

on all posterior segments, beginning on segment 28;

elytra with entire borders, attached by anterior

borders to elytriphores, which are fused medially

(Figure 3d). Branchiae elongate, inflated, ciliated

on under side, with distal appendage or branchial

cirrus narrowed to digitiform tip (Figure 2b). Dorsal

cirri on segments III and VI subulate (Figure la,/).

Prostomium partially hidden by elytriphores of

segment II (not attached); median antenna short,

subconical, inserted on anterior fourth of prostomium; lateral antennae slightly longer, conical,

inserted terminally; ventral palps elongate-tapered,

extending slightly beyond parapodial lobes; 2 pairs

small eyes, anterior pair formed of scattered pigment spots; nuchal organs oval, lateral to prostomium (Figure \b, c). Tentacular para podia (I)

with short subulate tentacular cirri, ventral pair

slightly longer than dorsal pair; 2 acicula with

hooked tips; 2 bundles of long, spreading capillary

setae (Figure la, c, d). Ventral buccal cirri on segment II much stouter than those following (Figure

\e).

Biramous parapodia supported by reddish ambercolored acicula and neuropodial hammer-shaped

distal plates (Figure 2a, b). Notopodial acicula with

hooked tips. Notopodial capillary setae smooth and

finely spinous, forming long, spreading bundles on

posterior part of notopodia; stout golden notopodial

hooks, beginning on segment III, either smooth or

finely spinous along curved edge, with tips finely

tapered or flattened, spoon-shaped (Figures \g, 2c).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

m

FIGURE 1.—Eulepethus hamifer (USNM 17483): a, Anterior end, dorsal view, elytra missing;

b, prostomium, dorsal view, with dorsal side segment II pulled back; c, prostomium and segment I, lateral view from left side; d, segment I, medial view; e, parapodium from segment II,

anterior view; /, parapodium from segment III, anterior view (base of dorsal cirrus dotted);

g, notopodial hooks from same; h, upper pectinate neuroseta from same; i, bilimbate neuroseta

from same; /, parapodium from segment IV, posterior view.
NUMBER 41

FIGURE 2.—Eulepethus hamifer (USNM 17483): a, Middle elytrigerous parapodium, anterior

view; b, middle branchial parapodium, posterior view; c, stout notopodial hooks from same;

d, upper pectinate neuroseta from same; e, limbate capillary neuroseta from same.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Neuropodial acicula with conspicuous hammershaped distal plates; thin-walled inflated areas on

anterodorsal parts of neuropodial bases (Figure 2a).

Neurosetae of several kinds: 1-2 upper pectinate

setae (Figures \h, 2d); bilimbate capillary setae of

anterior parapodia with short fine tips (Figure It);

unilimbate setae with longer fine tips (Figure 2e);

lower posterior bundles of long capillary setae on

anterior parapodia (Figure 1;); lower posterior

groups short capillary setae (Figure 26); neurosetae

of posterior region similar (without some stouter

upper neurosetae, as in some species of eulepethids).

Ventral cirri of anterior few segments slender,

tapered (Figure 1/,;); remainder globular, with short

slender tips (Figure 2b). Pygidium with single long

anal cirrus on right side, papillated along one side.

Pharynx not extended.

DISTRIBUTION.—Philippine Islands, Indochina

(Annam). In 80 meters.

REMARKS.—The type-specimen of Eulepis hamifera Grube from the Philippine Islands is no longer

available, at least it is not present in the Zoological

Museum Berlin (G. Hartwich, in litteris) nor in the

Zoological Museum Hamburg (G. Hartmann-Schroder, in litteris). The Albatross specimen from the

Philippine Islands, recorded by Treadwell (1920),

seems to agree for the most part with the descriptions and figures of Grube (1875, 1878).

Genus Par eulepis Darboux, 1900

TYPE-SPECIES.—Eulepis wyvillei Mclntosh, 1885,

by original designation. Gender: feminine.

DIAGNOSIS.—Body short, segments about 37 (34-

38). Elytra 12 pairs, on segments 2, 4, 5, 7 .. . 21,

24; elytra with lateral borders notched. Dorsal cirri

on segments 3 and 6. Branchiae 12 pairs, on segments 8, 10 . . .22, 23, 25, 26, 27. Posterior lamellae

begin on segment 28.

REMARKS.—Par eulepis Darboux was supposedly

separated from Eulepis Grube by the elytra inserting on segments 2, 3, 4, 6, 8 .. . 18, 20, 23 in the

former and on segments 2, 4, 5, 7, 9 . . . 19, 21, 24

in the latter. This, however, was based on an erroneous observation by Mclntosh (1885) when the parapodia of segment 3 were overlooked. There are

other characters, however, by which E. wyvillei, the

FIGURE 3—Eulepethus hamifer (USNM 17483): a, Right elytron from anterior region; b, right

elytron from middle region; c, right twelfth elytron; d, pair of elytra and elytriphores from

posterior region.
NUMBER 41

type-species of Pareulepis, may be separated from

E. hamifera, the type-species of Eulepis. The latter

genus, being preoccupied, was replaced by Eulepethus Chamberlin.

Pareulepis wyvillei (Mclntosh)

FIGURES 4, 5

Eulepis wyvillei Mclntosh, 1885, p. 131, pi. 19: fig. 11, pi. 20:

figs. 2, 3, pi. 24: figs. 2, 3, pi. 25: fig. 11, pi. 14A: figs. 4-6,

pi. 32A: fig. 7.

Eulepis splendida Treadwell, 1901, p. 189, figs. 19-22— Augener, 1906, p. 128.

Eulepethus sptendidus.—Treadwell, 1939, p. 195, fig. 24.

Pareulepis wyvillei.—Hartman, 1942, p. 95; 1959, p. 123.

MATERIAL EXAMINED.—Challenger station 33, off

Bermuda, 32° 21' N, 64° 35' W, 796 meters, coral

mud, 4 April 1873—holotype Eulepis wyvillei

(BMNH 1885: 12: 1: 105a).

Mayaguez Harbor, Puerto Rico: Fish Hawk

station 6062, 46-55 meters, sand, mud, shells, 20

January 1899—holotype E. splendida (USNM

15916); Fish Hawk station 6065, 7-11 meters, 20

January 1899—para type E. splendida (USNM

15917).

Grenada, Caribbean Sea, Blake station 247, 311

meters—1 specimen (MCZ 2235).

DESCRIPTION.—Length 18-37 mm, width 4-5 mm,

including setae, segments 35-38. Elytra 12 pairs,

becoming more elongate posteriorly, with lateral

notch (Figure 4g-t); first pair elytra (missing on

types of E. wyvillei and E. splendida; present on

MCZ 2235) with margin entire except for a notch

on inner anterior border; without papillae. Branchiae elongate, inflated, ciliated on underside; distal

branchial cirrus oval, tapering distally (Figure 5b).

Dorsal cirri, on segments 3 and 6, subulate (Figure

4e). Posterior lamellae, beginning on segment 28,

subcordiform to lanceolate (Figure 4c).

Posterior half of prostomium covered by segment

II, attached along midline; median antenna short,

conical, inserted anterodorsally on prostomium;

lateral antennae slightly longer, inserted more

ventrally; ventral palps elongate-tapered, extending

beyond tentacular cirri; eyes not visible; nuchal

organs oval, lateral to prostomium (Figure 46).

Tentacular parapodia (I) with short subulate tentacular cirri, subequal in length or with ventral pair

slightly longer than dorsal pair; 2 acicula; 2 tufts

capillary setae (Figure 4a, b). Ventral buccal cirri

segment II thicker than following ventral cirri

(Figure 4d).

Biramous parapodia supported by dark reddish

amber-colored acicula and neuropodial hammershaped distal plate (Figures 4d-f; 5a). Notopodial

acicula with hooked tips. Notopodial capillary setae

smooth and spinous, forming long, spreading bundles on posterior parts of notopodia; stout reddish

amber-colored notopodial hooks, beginning on segment III, smooth, with tips finely tapered or

flattened spoon-shaped (Figure 5c). Neuropodial

acicula with conspicuous hammer-shaped plates;

thin-walled inflated areas on dorso-anterior parts of

neuropodial bases (Figures 5a). Neurosetae of several

kinds: 1-2 upper pectinate setae (Figure 5d); limbate

capillaries with tips elongate, curled (Figure 5e);

lower non-limbate capillaries (Figures 5/); neurosetae of posterior region similar to anterior region

(without some stouter upper neurosetae, as in some

species of eulepethids). Ventral cirri of anterior few

segments slender, tapered (Figure 4e, f); rest globular, with short slender tips (Figure 5a, b). Pygidium

with single long anal cirrus on right side; cirrus

finely papillate along one side (Figure 4c). Pharynx

not extended.

DISTRIBUTION.—North Atlantic off Bermuda,

Caribbean region. In 7 to 796 meters.

REMARKS.—The single holotype of Eulepis

wyvillei, deposited in the British Museum, is now

in two pieces; most of the setae are broken off; all

the elytra are missing except for part of the ninth

right elytron; the worm consists of 35 segments, the

last one being small. The original description by

Mclntosh was erroneous and misleading in several

respects. There are 12 pairs of elytriphores on segments 2, 4, 5 . . . 21, 24 (not on segments 2, 3, 4, 6,

8, etc., since the parapodia of segment 3 were overlooked); Mclntosh indicated 15 pairs of elytra,

which included some of the larger posterior

lamellae. The one remaining elytron has a simple

lateral notch (not a double notch, as shown on

plate 25: figure 11 by Mclntosh). The dorsal cirri on

segment 6 were referred to as branchiae by Mclntosh.

Augener (1906: 129) indicated that Eulepis splendida possibly agreed with E. wyvillei. Hartman

(1942: 95) synonomized the two species.
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Pareulepis wyvillei (holotype Eulepis splendida, USNM 15916): a, Prostomium and

left tentacular parapodium (I), dorsal view (right I had been removed; dorsum of segment II

had been cut back); dotted area indicates place of attachment to underside of segment II; b,

same, lateral view of left side; c, posterior end, dorsal view, right side not completely drawn;

anal cirrus broken; d, parapodium from segment II, anterior view; e, parapodium from segment

III, posterior view; /, parapodium from segment IV, anterior view; g, left anterior elytron; h,

left middle elytron; i, left twelfth elytron.
NUMBER 41 11

FIGURE 5.—Pareulepis wyvillei (holotype Eulepis splendida, USNM 15916): a, Middle elytrigerous

parapodium, anterior view; b, middle branchial parapodium, posterior view; c, notopodial

hooks; d, upper pectinate neuroseta; e, limbate capillary neurosetae; /, lower nonlimbate

neuroseta.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pareulepis malayana (Horst)

FIGURES 6-10

Eulepis malayana Horst, 1913, p. 164, fig. 2; 1917, p. 128,

pi. 15: figs. 5-7.

Pareulepis malayana.—Harttnan, 1959, p. 123.

MATERIAL EXAMINED.—Siboga station 47, Bay of

Bima, 55 meters, mud with patches of fine coral

sand, 8/12 April 1899—1 specimen (USNM 39166;

mixed with Leanira vulturis Horst). Siboga station

204, Buton Strait, 4° 20' S, 122° 58' E, 75-94 meters,

sand with dead shells, 20 September 1899—syntype

(RNHL 1052). Siboga station 260, west of Great Kei

Island, 5° 36.5' S, 132° 55.2' E, 90 meters, sand, coral

and shells, 16/18 December 1899—syntype (ZMA

219).

DESCRIPTION.—Length 15-21 mm, width, including

setae, 4-6 mm, segments 34-37. Elytra 12 pairs,

becoming more elongate posteriorly, with lateral

notch; first pair elytra (absent on syntypes; present

on USNM 39166) with anterior margin papillate

(Figures Se, f; 9c, d). Branchiae elongate, inflated,

ciliated on underside, with distal branchial cirrus

tapered to digitiform tip (Figures 86, 9a). Dorsal

cirri subulate, posterior on notopodia of segments 3

and 6 (Figure la, c). Posterior lamellae, beginning

on segment 28, conical to lanceolate (Figure 10a, b).

Prostomium covered on posterior half by segment

II, attached along midline; median antenna short,

conical, inserted anterodorsally on prostomium;

lateral antennae slightly longer, inserted more

ventrally; ventral palps elongate-tapered, extending beyond tentacular cirri; 2 pairs small eyes on

posterior part prostomium; nuchal organs oval,

lateral to prostomium (Figure 6a-c). Tentacular

parapodia (I) each with 2 short subulate tentacular

cirri, subequal in length, 2 acicula, and 2 tufts

capillary setae (Figure 6a-d). Ventral buccal cirri on

segment II longer and stouter than those following

(Figure 6b, e).

Biramous parapodia supported by reddish ambercolored acicula and neuropodial hammer-shaped

distal plates (Figures 7a-c; 8a,b). Notopodial acicula

with hooked tips. Notopodial capillary setae smooth

and spinous, forming long spreading bundles on

posterior parts of notopodia; stout reddish ambercolored notopodial hooks, beginning on segment

III, smooth, with tips finely tapered or flattened

spoon-shaped (Figures Id, 8c, 10c). Neuropodial acicula with conspicuous hammer-shaped plates; thinwalled inflated areas on dorso-anterior parts of

neuropodial bases (Figure 86). Neurosetae of several

kinds: 1-2 upper pectinate setae (Figure 6/); limbate

capillaries with tips elongate, curved (Figures 6g, le,

8d, 96); smooth capillary setae on lower and posterior parts of neuropodia; in posterior neuropodia,

upper neurosetae slightly stouter and darker than

lower neurosetae, with tips curved downward and

faintly spinous along curved edge (Figure 10a,b,d,e).

Ventral cirri of anterior few segments slender, tapered (Figure 7a-c); rest globular, with short slender

tips (Figure 8a). Pygidium with single long anal

cirrus on right side; cirrus finely papillate along one

side. Pharynx not extended.

DISTRIBUTION.—Malay Archipelago. In 55 to 94

meters.

REMARKS.—P. malayana, from the Malay Archipelago, is close to P. wyvillei, from the Caribbean.

They differ in the following characters:

P. wyvillei (Mclntosh) P. malayana (Horst)

First elytra: With anterior notch Entire, with fringe of

papillae

Eyes: None Two pair small eyes on

posterior part prostomium

Neurosetae Upper and lower Upper neurosetae slightof posterior neurosetae similar ly stouter and darker

parapodia: than lower neurosetae,

with tips bent downward and finely spinous along curved

edge

Genus Mexieulepis Rioja, 1961

TYPE-SPECIES.—M. elongatus Rioja, 1961, by original designation, referred herein to M. weberi

(Horst, 1922). Gender: feminine.

DIAGNOSIS.—Body elongate, segments about 50

(37-62). Elytra numerous pairs; first 12 pairs larger,

on segments 2, 4, 5, 7 .. . 21, 24; smaller elytra on

all segments beginning on segment 27 or 28, with

elytriphores fused medially; elytra with lateral

margins fimbriated. Dorsal cirri on segments 3 and

6. Branchiae 11-12 pairs, on segments 8, 10 .. . 22,

23, 25, 26 (and sometimes 27).
NUMBER 41

e

FIGURE 6.—Pareulepis malayana (USNM 39166): a, Anterior end, dorsal view, elytra removed; b,

anterior end, ventral view; c, prostomium and tentacular parapodium (I), lateral view; d, tentacular parapodium (I), medial view; e, parapodium from segment II, anterior view; /, upper

pectinate neuroseta from same; g, limbate capillary neuroseta from same.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Pareulepis malayana (USNM 39166): a, Parapodium from segment 3, posterior view;

b, parapodium from segment 4, anterior view; c, parapodium from segment 6, posterior view; d,

notopodial hook from same; e, limbate capillary neuroseta from same.
NUMBER 41 15

FIGURE 8.—Pareulepis malayana (USNM 39166): a, Middle elytrigerous parapodium, posterior

view; b, middle branchial parapodium, anterior view; c, notopodial hook from same; d, limbate

capillary neuroseta from same; e, right first elytron; /, right third elytron.
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 9.—Pareulepis malayana (syntype, ZMA 219): a, Middle branchial parapodium, anterior

view, b, limbate capillary neurosetae from same; c, right middle elytron; d, right twelfth elytron.
NUMBER 41 17

FIGURE 10.—Pareulepis malayana (syntype, ZMA 219): a, Posterior lamelligerous parapodium,

anterior view; b, same, posterior view; c, notopodial hooks from same; d, upper neurosetae

from same; e, lower neuroseta from same.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Mexieulepis weberi (Horst), new combination

FIGURES 11-14

Eulepis weberi Horst, 1922, p. 199, fig. 2—Augener, 1927,

p. 43.

Pareulepis weberi.—Hartman, 1959, p. 123.

Mexieulepis elongatus Rioja, 1961, p. 238, figs. 1-29.—Hartman, 1965, p. 15.

MATERIAL EXAMINED.—Spaansche Haven, Curasao, West Indies, van der Horst, collector—holotype (ZMA 220). Spaansch Water, Curasao, van der

Horst, collector, 4 May 1920—1 specimen (ZMH

10324).

Sapelo Beach, Sapelo Island, SE Georgia, low

intertidal, muddy sand, J. M. Teal, collector, 25

June 1957—1 specimen (USNM 32359).

DESCRIPTION.—Length of holotype 52 mm, width,

including setae, 12 mm, segments 52. Length of

other specimen from Curasao (ZMH 10324) 13 mm,

width, including setae, 3 mm, segments 37. Length

of specimen from Georgia (USNM 32359) 75 mm,

width, including setae, 11 mm, segments 62. Anterior 12 pairs elytra larger, fimbriated along external border (flattened, leaflike processes); first pair

elytra fimbriated on anterior border; elytral surface

smooth except for scattered microtubercles on anterior part; elytriphores large, inflated, converging

middorsally (Figure 11 a-c, e). Smaller elytra on all

posterior segments, beginning on segment 27 [specimens from Curasao and Veracruz (Rioja, 1961)] or

segment 28 (large specimen from Sapelo Island);

elytra with few lateral processes or simply notched,

attached by anterior borders to elytriphores, which

are fused medially (Figure lid). Branchiae 11-12

pairs, ciliated on underside, with distal branchial

cirri (Figure 136). Dorsal cirri, on segments 3 and 6,

subulate (Figures lie, 12a).

Prostomium partially hidden by and attached to

underside of segment II; median antenna short,

oval, inserted anterodorsally on prostomium; lateral

antennae longer, conical, inserted anteroventrally;

ventral palps elongate-tapered, extending slightly

beyond parapodial lobes; no eyes visible; nuchal

organs clavate, lateral to prostomium (Figure lie).

Tentacular parapodia (I) with short subulate tentacular cirri, ventral pair slightly longer than dorsal

pair; 2 acicula with hooked tips; 2 bundles long,

spreading capillary setae (Figure \\e,g). Ventral

buccal cirri of segment II stouter and longer than

those following (Figure llh).

Biramous parapodia supported by reddish ambercolored acicula and neuropodial hammer-shaped

distal plates (Figures 12a,b; 13a,6). Notopodial

acicula with hooked tips. Notopodial capillary setae

smooth and finely spinous, forming long spreading

bundles on posterior parts of notopodia; some of

capillary setae on segment II stouter, with spinous

rows (Figure Hi); stout reddish amber-colored

notopodial hooks, beginning on segment III,

smooth, with tips finely tapered or flattened spoonshaped (Figures 11/, 12c, 146). Neuropodial acicula

with conspicuous hammer-shaped distal plates.

Neurosetae of several kinds: 1-2 upper pectinate

setae (Figures 12d, 13c); upper and lower limbate

and nonlimbate capillaries (Figures 12e,g; 13d,/);

lower posterior bundles of long capillary setae

(Figures llh, 12a,6); middle stout smooth acicular

setae, beginning on segment II (Figures 11A, 12/,

13e). In posterior region, upper neurosetae stout,

bent downward, with tips slender or blunt; lower

neurosetae slender, slightly limbate, with slender

tips curved upward (Figure 14a,c,d). Ventral cirri of

anterior few segments slender, tapered (Figure

12fl,6); rest globular, with short distal tips (Figure

13a,6). Pygidium with single long anal cirrus on

right side; cirrus papillate along one side; left cirrus

short bulbous. Pharynx extended on specimen from

Georgia, with 13 pairs larger papillae and small

lateral pair; 2 pairs platelike chitinous ridges (not

toothlike, as in Polynoidae or Sigalionidae; (Figure

He,/).

DISTRIBUTION.—West Indies (Curasao), Gulf of

Mexico (Veracruz), Georgia (Sapelo Island). Low

intertidal.

REMARKS.—The types of Mexieulepis elongatus

Rioja no longer exist (M. E. Caso Munoz of the

Instituto de Biologia Universidad Nacional Automona de Mexico, in litteris). Based on the original

figures and description, it appears to agree with

M. weberi.

M. weberi is the largest of the known species of

Eulepethidae, the specimen from Sapelo Island

reaching a length of 75 mm and a width of 11

mm for 62 segments. Horst's type-specimen from

Curasao is 52 mm long. Rioja indicated his specimens had a length of 50-56 mm. Augener's specimen

from Curacao is only 13 mm long.
NUMBER 41 19

FIGURE 11.—Mexieulepis weberi (USNM 32359): a, Right third elytron; b, right eighth elytron; c,

right twelfth elytron; d, four of right elytra and elytriphores from posterior region (anterior

to posterior); e, dorsal view, anterior end, first two right elytra removed; pharynx extended;

posterior part prostomiutn covered by segment II (place of attachment to midline stippled); /,

distal tip of extended pharynx, anterior view; g, par apod him from segment I, lateral view; h,

parapodium from segment II, anterior view; i, stouter capillary notoseta from same; ;, notopodial

hooks from same; k, stout acicular neuroseta from same; /, limbate neuroseta from same.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12.—Mexieulepis weberi (USNM S2359): a, Parapodium from segment III, posterior view;

b, parapodium from segment V, posterior view; c, notopodial hook from same; d, pectinate

neuroseta from same; e, upper limbate neuroseta from same; /, middle stout acicular neuroseta

from same; g, lower limbate and nonlimbate neurosetae from same.
NUMBER 41 21

FIGURE 13.—Mexieulepis weberi (USNM 32359): a, Elytrigerous parapodium from middle region,

posterior view; b, branchial parapodium from middle region, anterior view; c, upper pectinate

neuroseta from same; d, upper limbate capillary neuroseta from same; e, middle acicular neuroseta from same; /, lower limbate capillary neuroseta from same.
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Mexieulepis weberi (USNM 32359): a, Elytrigerous parapodium from posterior

region, anterior view; b, notopodial hook from same; c, upper neurosetae from same; d, lower

neuroseta from same.

Grubeulepis, new genus

TYPE-SPECIES.—Eulepis fimbriata Treadwell,

1901. Gender: feminine.

DIAGNOSIS.—Body short, segments about 38 (32-

40). Elytra 12 pairs, on segments 2, 4, 5, 7 .. . 21,

24; elytra with lateral borders fimbriated. Dorsal

cirri on segments 3 and 6. Branchiae 10-13 pairs, on

segments 8, 10 .. . 22, 23, 25, (26, 27, 28). Posterior

lamellae beginning on segments 26-29.

Key to the Species of Grubetdepis, New Genus

1. Lateral processes of elytra digitiform, not biarticulate (Figures ITd-f, ZOd-f, 23/-i). [First pair

elytra entire or with few minute papillae (Figures 15a, 23e). Last pair elytra (12th) with 13-21

lateral processes (Figures 17/, 20/, 23»)] 2

1'. Some lateral processes of some or all elytra biarticulate (Figures 27/, 29e-h, 30/-/», $lh-k) . 4

2. Upper neurosetae of posterior region not stouter than lower ones, not bent downward (not

spinous; Figure 17a, c). Without acicular neurosetae in some anterior segments. [Notopodial

hooks smooth (Figure 16c). Branchiae 12 pairs] G. fimbriata (Treadwell)

2' Upper neurosetae of posterior region stouter than lower ones and bent downward. With acicular neurosetae in some anterior segments (Figures 186, d; 2\e,g) 3

3. Upper neurosetae of posterior region finely spinous along bend, tapering to fine tips (Figure

23a-c). Notopodial hooks smooth (Figure 22c). Branchiae 12 pairs

G. ecuadorensis, new species

3' Upper neurosetae of posterior region tapering abruptly to fine tips (not spinous; Figure 20a, b).

Notopodial hooks spinous along bend (Figure 19c). Branchiae 13 pairs . G. geayi (Fauvel)
NUMBER 41 23

4. With acicular neurosetae in some anterior segments (Figure 25e, i). Last elytra (12th) each

with more than 20 (22-25) entire processes (Figure 27/i). Some anterior elytra with biarticulate processes (Figure 27/). Notopodial hooks spinous on upper distal part (Figure 26c).

[Upper stouter neurosetae of posterior region serrulate along the bend (Figures 24e, 27c).

Branchiae 11 pairs, 12th may be transitional.] . • • G. mexicana (Berkeley and Berkeley)

4'. Without acicular neurosetae in any anterior segments. Last elytra (12th) each with less than

15 lateral processes. Most elytra with biarticulate processes. Notopodial hooks smooth . . 5

5. Last elytra (12th) each with 5-6 subrectangular lateral processes (Figure iOh, ilk). First elytra

each with 12-13 papillae on anterior border (Figures 30c, 31/). [Upper stouter neurosetae of

posterior segments finely spinous along bend (Figure Sid). Branchiae 12 pairs.] ... .

G. augeneri, new species

5' Last elytra (12th) each with 10-12 digitiform lateral processes (Figure 29/t). First elytra each

with 7-8 papillae on anterior border (Figure 8, in Jones, 1962) 6

6. Neurosetae of anterior segments bilimbate, with long tapering tips. Upper stouter neurosetae

of posterior segments spinous along bend, tips limbate or scoop-shaped (Figure 25, in Jones,

1962). Branchiae 10 pairs * G. sulcatisetis (Jones)

6'. Neurosetae of anterior segments bilimbate, with short tips (Figure 28d). Upper stouter neurosetae of posterior segments smooth, tips limbate, spatulate (Figure 29c). Branchiae 11 pairs.

G. tebblei, new species

Grubeulepis fimbriata (Treadwell),

new combination

FIGURES 15-17

Eulepis fimbriata Treadwell, 1901, p. 190, figs. 23, 24.

Eulepethus fimbriatus.—Treadwell, 1939, p. 196, fig. 25.

Pareulepis fimbriata.—Hartman, 1944, p. 14.

Not Eulepis fimbriata.—Augener, 1918, p. 153 (=Grubeulepis

augeneri, new species).

Not Pareulepis fimbriata.—Hartman, 1939, p. 79 (=G. ecuadorensis, new species, and G. mexicana).—Rullier, 1965,

p. 16 (=Grubeulepis augeneri, new species.)—Reish, 1968,

p. 73 (=Grubeulepis mexicana).

MATERIAL EXAMINED.—Mayaguez Harbor, Puerto

Rico, Fish Hawk station 6061, 33 meters, sand, mud,

20 January 1899—4 syntypes (USNM 15640). Mayaguez, Puerto Rico, 11-15 meters, 10 December 1954,

M. J. Allen, collector—3 specimens (AHF).

Caledonia Bay, Panama (Atlantic), shore, April

1939, station A 52-39—1 specimen (AHF).

DESCRIPTION.—Length 14-24 mm, width, including setae, 4-6 mm, segments 37-38. Elytra 12 pairs,

becoming more elongate posteriorly; first pair elytra

smooth except occasionally with 2 clavate papillae

on anterior border; rest of elytra with variable

number lateral leaflike processes arranged between

anterior and posterior rounded lobes; twelfth pair

each with 15-17 processes (Figures 15a, lld-f).

Branchiae 12 pairs, inflated, with distal branchial

cirri (Figure 16a). Dorsal cirri, on segments 3 and

6, subulate (Figure 15g). Posterior lamellae, beginning on segment 28, oval (Figure 17a).

Prostomium covered by segment II and attached

along midline on posterior half; median antenna

short, globular, inserted anterodorsally on prostomium; lateral antennae slightly longer, conical,

inserted more ventrally; ventral palps elongatetapered, extending slightly beyond tentacular cirri;

2 pairs small eyes on anterolateral border (easily

overlooked); nuchal organs oval, lateral to prostomium (Figure 156-d, h). Tentacular parapodia

(I) with short subulate tentacular cirri, ventral pair

slightly longer than dorsal pair; 2 acicula; 2 tufts

capillary setae (Figure \bd,e,h). Ventral buccal cirri

on segment II thicker and longer than those following (Figure 15/).

Biramous parapodia supported by light ambercolored acicula and neuropodial hammer-shaped

distal plates (Figures 16a, 6; 17a). Notopodial acicula with hooked tips. Notopodial capillary setae

smooth and spinous, forming long spreading bundles on posterior parts of notopodia; stout light

amber-colored notopodial hooks, beginning on segment III, smooth, with tips finely tapered or flattened spoon-shaped (Figures 16c, 176). Neuropodial

acicula with hammer-shaped distal plates; thinwalled inflated areas on anterodorsal parts of neuropodial bases (Figure 166). Neurosetae of several

kinds: 1-2 upper pectinate setae (Figure 16d); limbate and nonlimbate capillaries (Figures 16e, 17c);

neurosetae of posterior region similar to anterior

region (without any stouter upper neurosetae; without acicular neurosetae in anterior segments). Ventral cirri of anterior few segments slender, tapered

(Figure 15g); those of following segments globular,
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

e

FIGURE 15.—Grubeulepis fimbriata (a-g, syntypes, USNM 15640; h, Puerto Rico, AHF): a,

Anterior end, dorsal view; b, anterior end, dorsal view, first two pairs elytra removed; c, prostomium, dorsal view; shaded area indicates place of attachment to underside of segment II; d,

prostomium and tentacular segment (I), dorsal view; e, tentacular parapodium (I), lateral view; /,

parapodium from segment II, anterior view; g, parapodium from segment III, posterior view; h,

prostomium and tentacular segment, dorsolateral view (palp not shown).
NUMBER 41 25

FIGURE 16.—Grubeulepis fimbriata (syntypes, USNM 15640): a, Middle branchial parapodium,

posterior view; b, middle elytrigerous parapodium, anterior view; c, notopodial hooks from

same; d, upper pectinate neuroseta from same; e, neurosetae from same.
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

e

FIGURE 17.—Grubeulepis fimbriata (syntypes, USNM 15640): a, Posterior lamelligerous parapodium, posterior view; b, notopodial hooks from same; c, neurosetae from same; d, left third

elytron; e, left eighth elytron; /, left twelfth elytron.

with short slender tips (Figure I6a,b). Pygidium

with single long anal cirrus on right side; cirrus

finely papillate along one side. Pharynx not

extended.

DISTRIBUTION.—West Indies, Atlantic side

Panama. Intertidal to 33 meters.

REMARKS.—The original description of Eulepis

fimbriata by Treadwell (1901), based on four specimens from Puerto Rico, was sketchy and poorly

illustrated. Reexamination of additional specimens,

identified as this species, indicates that most of them

should be referred to other species. This includes

records by Augener (1918) and Rullier (1965) from

West Africa, by Hartman (1939) from southern

California and Ecuador, and by Reish (1968) from

the Gulf of California. Both Hartman (1939) and

Rullier (1965) included the Indo-Pacific Eulepis

geayi under Pareulepis fimbriata. Thus, G. fimbriata is not as widely distributed as its records in

the literature indicate.
NUMBER 41 27

Grubeulepis geayi (Fauvel), new combination

FIGURES 18-20

Eulepis geayi Fauvel, 1918, p. 503, fig. 1, ah; 1919, p. 335,

pi. 15: figs. 17-21, pi. 17: figs. 76-79.—Pruvot, 1930, p. 17,

pi. 2: figs. 51-61.—Day, 1951, p. 14.

Pareulepis geayi.—Hartman, 1959, p. 123.—Day, 1962, p. 632;

1967, p. 45, fig. 1.3, q-u.

Not Eulepis geayi.—Fauvel, 1940, p. 9 (=Grubeulepis augeneri, new species).—Tebble, 1955, p. 79 (=Grubeulepis

tebblei, new species).

MATERIAL EXAMINED.—Delagoa Bay, Inhaca Island, southeast Africa, intertidal—1 specimen

(BMNH 1952: 1: 2: 1).

DESCRIPTION.—Length 35 mm (21-40 mm), width,

including setae, 8 mm (5-8 mm), segments 39 (36-39).

Elytra 12 pairs, becoming more elongate posteriorly; first pair (missing on specimen examined)

with 3-4 papillae on anterior margin (Day, 1967);

remaining elytra with variable number lateral leaflike processes between anterior and posterior

rounded lobes [for example, 3 on second, 8 on

eighth, and 17 (left) and 21 (right) on twelfth)];

processes digitiform, not articulate (Figure 20d-f).

Branchiae 13 pairs, inflated, with distal branchial

cirrus (Figure 196). Dorsal cirri on segments 3 and

6, subulate (Figure 186). Posterior lamellae, beginning on segment 28, subreniform to subconical

(Figure 20a).

Prostomium covered by segment II, attached middorsally on posterior half; median antenna short,

conical, inserted anterodorsally on prostomium; lateral antennae conical, inserted more ventrally;

palps elongate-tapered, extending beyond tentacular cirri; no eyes visible; nuchal organs clavate, lateral to prostomium (Figure 18a). Tentacular parapodia (I) narrower basally, enlarged distally, each

with dorsal and ventral tentacular cirri, subequal in

length, 2 acicula and 2 bundles of smooth and

spinous capillary setae. Ventral buccal cirri on segment II thicker and longer than those following.

Biramous parapodia supported by reddish ambercolored acicula and neuropodial hammer-shaped

distal plates (Figures I9a,b; 20a). Notopodial acicula with hooked tips. Notopodial capillary setae

smooth and spinous, forming long spreading bundles emerging from posterior parts of notopodia;

stout reddish amber-colored notopodial hooked setae, spinous along bend and distal part, tapering

FIGURE 18.—Grubeulepis geayi (BMNH 1952: 1: 2: 1): a, Prostomium, dorsal view, turned slightly

to right; b, parapodium from segment III, anterior view; c, upper neuroseta from same; d, middle

acicular seta from same; e, lower neuroseta from same.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Grubeulepis geayi (BMNH 1952: 1: 2: 1): a, Middle elytrigerous parapodium, posterior view; b, middle branchial parapodium, anterior view; c, hooked notosetae from same; d,

upper pectinate neuroseta from same; e, upper neurosetae from same; /, lower neurosetae from

same.
NUMBER 41 29

distally to fine tips or flattened spatulate (Figure

19c). Neuropodial acicula with hammer-shaped distal plates. Neurosetae of several kinds: 1-2 upper

pectinate setae (Figure 19d); limbate capillaries

(Figure \8c,e); slender setae, tapering gradually to

long capillary tips (Figure 19/); slightly stouter

setae tapering rather abruptly to capillary tips

(Figure 19e); few stout acicular neurosetae in some

anterior segments (Figure 18d). In posterior region,

some of upper neurosetae much stouter than lower

neurosetae, bent downward, tapering abruptly to

fine tips (Figure 20a-c). Pygidium with single long

anal cirrus on right side; left cirrus short, bulbous.

Pharynx not extended.

DISTRIBUTION.—Indo-west-Padfic (Madagascar,

Mozambique, Natal), Red Sea, South-west-Pacific

(New Caledonia). Intertidal. May be commensal

with Polyodontes melanotus, according to Day

(1962).

REMARKS.—The type-specimens of Eulepis geayi

from Madagascar apparently are no longer available, at least they were not found in the Museum

FIGURE 20.—Grubeulepis geayi (BMNH 1952: 1: 2: 1): a, Posterior Iamelligerous parapodium,

anterior view; b, upper neurosetae from same; c, lower neuroseta from same; d, left second elytron;

e, left eighth elytron; /, left twelfth elytron.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

National d'Historie Naturelle, Paris (J. RenaudMornant, in litteris) or in the collections of Polychaeta at the University Catholique, Angers (F. Rullier, in litteris). The specimen from Inhaca Island,

deposited in the British Museum (Natural History)

and identified by J. H. Day, seems to agree with

Fauvel's original description. Reexamination of additional specimens, identified as belonging to this

species, indicates that they should be referred to

other species. This includes the record of Fauvel

(1940) from the Adriatic and that of Tebble (1955)

from West Africa. Thus, G. geayi is not as widely

distributed as its records in the literature indicate.

Eulepis geayi was referred to Pareulepis fimbriata

(Treadwell) by Hartman (1939). In her Catalogue,

Hartman (1959) listed the two species as distinct.

Grubeulepis ecuadorensis, new species

FICURES 21-23

Pareulepis fimbriata.—Hartman, 1939, p. 79, pi. 23: figs. 280,

282-285, 287, 288 (part; Velero station 216-34). [Not Eulepis

fimbriata Treadwell, 1901.]

MATERIAL EXAMINED.—Cape San Francisco, Ecuador, 36.5 meters, muck, Velero station 216-34,

11 February 1934—holotype (AHF).

DESCRIPTION.—Length of holotype 35 mm, width,

including setae, 8 mm, segments 40, last small.

Elytra 12 pairs, becoming more elongate posteriorly;

first pair oval, with entire margins (no papillae);

rest of elytra with variable number lateral leaflike

processes (3-13) arranged between anterior and posterior rounded lobes; processes digitiform, not articulate (Figure 23e-i; plate 23: figs. 285, 287, in

Hartman, 1939). Branchiae 12 pairs, inflated, with

distal branchial cirrus (Figure 22a). Dorsal cirri, on

segments 3 and 6, subulate (Figure 21e; plate 23:

fig. 280, in Hartman, 1939). Posterior lamellae, beginning on segment 28, subcordiform to subconical

(Figure 23a,6).

Prostomium covered by segment II and attached

middorsally by more than half its length; median

antenna short, oval, inserted anterodorsally on prostomium; lateral antennae conical, inserted more

ventrally; palps elongate-tapered, extending beyond

tentacular cirri; 3 pairs small eyes anterolaterally in

close groups; nuchal organs clavate, lateral to prostomium (Figure 21a). Tentacular parapodia (I) narrower basally, enlarged distally, each with dorsal

and ventral tentacular cirri, subequal in length,

2 acicula and 2 bundles of smooth and spinous

capillary setae (Figure 21a,&). Ventral buccal cirri

on segment II thicker and longer than those following (Figure 21c).

Biramous parapodia supported by light ambercolored acicula and neuropodial hammer-shaped

distal plates (Figures 22a,b; 23a,b). Notopodial acicula with hooked tips. Notopodial capillary setae

smooth and spinous, forming long, spreading bundles emerging from posterior parts of notopodia;

stout light amber-colored notopodial hooks, beginning on segment HI, smooth, tapering distally to

fine tips or flattened spatulate (Figure 22c). Neuropodial acicula with hammer-shaped distal plates.

Neurosetae of several kinds: 1-2 upper pectinate

setae (Figure 22d); widely bilimbate neurosetae with

short tips in anterior parapodia (Figure 2ldJ,h);

limbate and non-limbate neurosetae with tapering

tips (Figure 22e,f); single dark acicular neuroseta in

segments 3-8 (Figure 21g). Upper neurosetae of posterior region much stouter than lower ones, curved

downward, tapering to fine tips, finely spinous

along bend (Figure 23a-d). Ventral cirri of anterior

few segments subulate, tapered (Figure 21e); following ventral cirri globular, with short slender tips

(Figure 22a,6). Pygidium with single long anal cirrus on right side; cirrus finely papillate along one

side; left cirrus short, bulbous. Pharynx not

extended.

DISTRIBUTION.—Central Pacific (off Ecuador). In

36.5 meters.

Grubeulepis mexicana (Berkeley and

Berkeley), new combination

FICURES 24-27

Eulepethus mexicanus Berkeley and Berkeley, 1939, p. 328,

figs. 4-7.

Pareulepis fimbriata.—Hartman, 1939, p. 79, pi. 23: figs. 281,

286 (part; Velero station 770-38; Mission Bay, California);

1961, p. 54.—Reish, 1968, p. 73. [Not Eulepis fimbriata

Treadwell, 1901.]

MATERIAL EXAMINED.—Isle Grande Bay, Mexico,

M/S Stranger, 11 meters, fine sand, 8 April 1937,

W. Williams, collector—holotype of Eulepethus

mexicanus (USNM 32893).

Bahia de los Angeles, Gulf of California, Mexico,

30 meters, brown silty very fine sand, station 45,

27 April 1962, J. L. Barnard, J. R. Grady, and
NUMBER 41 31

e

FIGURE 21.—Grubeulepis ecuadorensis, new species (holotype, AHF): a, Prostomium and tentacular

segment, left lateral view; b, tentacular parapodium, lateral view; c, parapodium from segment

II, anterior view; d, bilimbate neurosetae from same; e, parapodium from segment III, posterior

view; /, upper bilimbate neuroseta from same; g, middle acicular seta from same; h, lower bilimbate neurosetae from same.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

e

FIGURE 22.—Grubeulepis ecuadorensis, new species (holotype, AHF): a, Middle branchial parapodium, anterior view; b, middle elytrigerous parapodium, posterior view; c, notopodial hooks

from same; d, upper pectinate neuroseta from same; e, middle neuroseta from same; /, lower

neuroseta from same.

D. J. Reish, collectors—3 specimens (USNM 39283).

Off San Juan Light, Guatemala, 13-20 meters,

sand, shells, mud, Velero station 770-38, 11 January

1938—2 specimens (AHF).

Southern California: Mission Bay, intertidal—

4 specimens (AHF). Newport Bay, intertidal—

1 specimen (AHF). Velero station 4720-56, 33° 37'

39" N, 117° 58' 16" W, 15 meters, dark green silt,

broken shells, 21 November 1956—3 small specimens (AHF). Velero station 6731-59, Newport

shelf, 34° 17' 53" N, 119° 21' 05" W, 7 meters, very

fine silty sand, 6 December 1959—4 small specimens

(AHF).

DESCRIPTION.—Length up to 33 mm, width, including setae, up to 9 mm, segments up to 37.

Holotype of E. mexicanus rather small, with length

of 15 mm for 38 segments. Elytra 12 pairs, becoming

more elongate posteriorly; first pair of elytra with

anterior fringe of minute papillae (about 7); rest of

elytra with variable number digitiform lateral processes (3-25) arranged between anterior and posterior

rounded lobes; some of processes on more anterior

elytra biarticulate (Figures 24a; 2.1 e-h; pi. 23: figs.

281, 286, in Hartman, 1939). Branchiae 11 pairs,

inflated, with distal branchial cirrus (Figure 26&).

Dorsal cirri, on segments 3 and 6, subulate (Figures

24a, 25e). Posterior lamellae, beginning on segment

27, oval to lanceolate (Figures 24c, 27a).
NUMBER 41 33

FIGURE 23.—Grubeulepis ecuadorensis, new species (holotype, AHF): a, Posterior lamelligerous

parapodium, anterior view; b, same, posterior view; c, upper neurosetae from same; d, lower

neuroseta from same; e, left first elytron; /, left second elytron; g, left third elytron; h, left

seventh elytron; i, right twelfth elytron.

Prostomium nearly covered by segment II and

attached along midline; median antenna short,

globular, inserted anterodorsally on prostomium;

lateral antennae longer, conical, inserted more ventrally; ventral palps elongate-tapered, extending

slightly beyond tentacular cirri; 3 to 5 pairs small

eyes visible (easily overlooked); nuchal organs oval,

lateral to prostomium (Figures 24a,b; 25a). Tentacular parapodia (I) with short subulate tentacular

cirri, lower pair slightly longer than upper pair;

2 acicula; 2 tufts capillary setae (Figures 24a, 25a, b).

Ventral buccal cirri on segment II thicker and

longer than those following (Figure 25 a, c).

Biramous parapodia supported by light amber-
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 24.—Grubeulepis mexicana (holotype, USNM 32893): a, Anterior end, dorsal view, left

two elytra removed; left parapodium II had been previously removed; b, prostomium, left

lateral view (not to scale); c, posterior lamelligerous parapodium, posterior view; d, hooked

notoseta from same; e, upper neuroseta from same; /, lower neuroseta from same.

colored acicula and neuropodial hammer-shaped

distal plates (Figure 26a,b). Notopodial acicula with

hooked tips. Notopodial capillary setae smooth and

spinous, forming long, spreading bundles on posterior parts of notopodia; stout light amber-colored

notopodial hooks, beginning on segment III, with

tips finely tapered or flattened spoon-shaped, finely

spinous on upper distal part (Figures 24d, 25/, 26c,

276). Neuropodial acicula with hammer-shaped distal plates. Neurosetae of several kinds: 1-2 upper

pectinate setae (Figure 25g); stout acicular neurosetae in some anterior segments (Figure 25i); limbate

and non-limbate capillary neurosetae (Figures 25d,

h,j; 26d). Upper neurosetae of posterior region

stouter and darker than lower neurosetae, bent

downward, corrugated or serrulate along bend (Figures 24c,e,f; 27a,c,d). Ventral cirri of few anterior

segments slender, tapered (Figure 25e); rest globular, with short slender tips (Figure 26a,b). Pygidium

with single long anal cirrus on right side; cirrus

finely papillate along one side. Pharynx not

extended.

DISTRIBUTION.—Southern California, Gulf of California, Mexico (Isle Grand Bay), Guatemala.

Intertidal to 30 meters.

REMARKS.—The small oval specimens from southern California (Velero stations 4721-56 and 6731-59),

reported by Hartman (1961) as P. fimbriata, have

only 16-18 segments; the elytral papillar processes

are 2 jointed; the upper neurosetae are serrated and

stouter than the lower neurosetae; they appear to be

the young of G. mexicana.

Fauvel (1940: 9) referred Eulepethus mexicanus

to Eulepis geayi. Hartman (1959: 123) referred the

same species to Pareulepis fimbriata. The three

species are herein considered to be distinct.
NUMBER 41 35

FIGURE 25.—Grubeulepis mexicana (Mission Bay, southern California, AHF): a, Left lateral view

anterior end, segment II cut back; b, tentacular segment (I), lateral view; c, parapodium from

segment II, anterior view; d, upper, middle and lower neurosetae from same; e, parapodium

from segment III, anterior view; /, notopodial hooks from same; g, upper pectinate neuroseta

from same; h, upper limbate capillary neuroseta from same; t, middle acicular neuroseta from

same; ;', lower limbate and nonlimbate capillary neurosetae from same.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Grubeulepis mexicana (Mission Bay, southern California, AHF): a, Middle elytrigerous parapodium, anterior view; b, middle branchial parapodium, posterior view; c, notopodial

hooks from same; d, upper, middle and lower neurosetae from same.

Grubeulepis sulcatisetis (Jones), new

combination

Pareulepis sulcatisetis J<

1965, p. 15.

[ones, 1962, p. 174, figs. 1-27—Hartman,

MATERIAL EXAMINED.—Green Bay, Jamaica, West

Indies, 11 February 1960, I. M. Goodbody, collector

—holotype and paratype (AMNH 3605, 3606).

DESCRIPTION.—Length 9.5-10 mm, width, including setae, 2.4 mm, segments 32. Elytra 12 pairs, becoming more elongate posteriorly; first pair with

anterior fringe of minute papillae (about 7); rest of

elytra with variable number digitiform processes

(4-10) arranged between anterior and posterior

rounded lobes; some of processes on more anterior

elytra biarticulate, some on posterior elytra tripartite (Figures 8-11, in Jones, 1962). Branchiae 10

pairs, inflated, with distal branchial cirrus (Figure 7,

B-E, in Jones, 1962). Dorsal cirri, on segments 3 and

6, subulate (Figures 3, 6, 7,A, in Jones, 1962). Pos-
NUMBER 41 37

FIGURE 27.—Grubeulepis mexicana (Mission Bay, southern California, AHF): a, Posterior lamelligerous parapodium, anterior view; b, notopodial hooks from same; c, upper neuroseta from

same; d, lower neurosetae from same; e, right first elytron; /, right second elytron; g, right sixth

elytron; h, right twelfth elytron.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terior lamellae pyriform, beginning on segment 26

(Figure 7, F-H, in Jones, 1962).

Prostomium covered by segment II and attached

middorsally by more than half its length; median

and lateral antennae subequal, short, conical; 3

pairs small eyes (Figures 1,2, in Jones, 1962). Tentacular parapodia (I) with dorsal and ventral tentacular cirri subequal in length (Figure 4, in Jones,

1962).

Neurosetae of several kinds: 1-2 upper pectinate

setae; bilimbate and unilimbate capillaries; slender

capillaries on lower posterior face of neuropodia;

without acicular setae in some anterior segments.

Upper neurosetae of posterior segments stouter than

lower neurosetae, bent downward, spinous along

bend, with tips scoop-shaped (Figures 24, 25, in

Jones, 1962). Pharynx with about 14 pairs distal

papillae (15-1-18—Jones) and 2 pairs chitinous

plates (no definite teeth).

DISTRIBUTION.—West Indies (Jamaica).

REMARKS.—Jones did not distinguish between

dorsal cirri, branchiae, and posterior lamellae (Figure 7, AH, in Jones, 1962).

Grubeulepig tebblei, new species

FIGURES 28-29

Pareulepis geayi.—Tebble, 1955, p. 79, fig. 2. [Not Eulepis

geayi Fauvel, 1918.]

MATERIAL EXAMINED.—Gold Coast, West Africa,

off Accra and off Christiansborg Castle, Buchanan

survey, 11-13 meters, station 123—holotype (BMNH

1953: 1: 623); station 118—paratype (BMNH 1953:

1: 622).

DESCRIPTION.—Length 13-17 mm, width, including setae, 4 mm, segments 33. Elytra 12 pairs, becoming more elongate posteriorly; first pair with

fringe of about 8 papillae on anterior border; rest

of elytra with variable number of digitiform lateral

processes—about 12 on twelfth elytra, some biarticulate (Figure 29e-h). Branchiae 11 pairs, inflated, with distal branchial cirrus (Figure 28e).

Dorsal cirri, on segments 3 and 6, subulate (Figure

28c). Posterior lamellae, beginning on segment 27,

subcordiform to lanceolate (Figure 29a).

Prostomium nearly covered by segment II and

attached along midline on posterior half; median

antenna short, conical, inserted anterodorsally on

prostomium; lateral antenna short, conical, inserted

more ventrally; ventral palps elongate-tapered, extending slightly beyond tentacular cirri; 2 pairs

small eyes on posterior part; nuchal organs oval,

lateral to prostomium (Figure 28a). Tentacular

parapodia (I) with short subulate tentacular cirri,

subequal in length or ventral pair slightly longer

than dorsal pair; 2 acicula; 2 tufts capillary setae

(Figure 28&). Ventral buccal cirri on segment II

thicker and longer than those following.

Biramous parapodia supported by reddish ambercolored acicula and neuropodial hammer-shaped

distal plates (Figure 28e). Notopodial acicula with

hooked tips. Notopodial setae smooth and spinous,

forming long, spreading bundles on posterior parts

of notopodia; stout reddish amber-colored notopodial hooks smooth, with tips finely tapered or spatulate (Figures 28/, 29&). Neuropodial acicula with

hammer-shaped distal plates. Neurosetae of several

kinds: upper pectinate setae; bilimbate setae with

short tips in some anterior parapodia (Figure 28d);

lower group slender limbate and non-limbate setae

tapering gradually to slender tips (Figure 28g).

Upper neurosetae of posterior parapodia stouter

than lower neurosetae, bent downward, with tips

limbate, spatulate (Figure 29a,c,d). Ventral cirri of

anterior few segments slender, tapered (Figure 28c);

rest of ventral cirri globular, with short slender tips

(Figure 28e). Pygidium with single long anal cirrus

on right side; short rudiment on left side. Pharynx

not extended.

DISTRIBUTION.—West Africa (Gold Coast). In 11-

13 meters.

REMARKS.—The prostomium was incorrectly

shown by Tebble (1955)—it is not bifurcate anteriorly and the median antenna is located anterodorsally and not posteriorly on the prostomium.

Grubeulepis augeneri, new species

FIGURES 30-31

Eulepis fimbriata.—Augener, 1918, p. 153, pi. 3: figs. 39-41,

text-fig. 10. [Not Treadwell, 1901.]

Eulepis geayi.—Fauvel, 1940, p. 9, fig. la. [Not Fauvel, 1918.]

Pareulepis fimbriata.—Rullier, 1965, p. 16. [Not Eulepis

fimbriata Treadwell, 1901.]

MATERIAL EXAMINED.—French Congo, Sett£

Cama, A. Hupfer, collector—holotype (ZMH 592).

Adriatic, station 434—paratype (MNHNP). Togo,

West Africa, 6° 05' N, 1° 32' E, 34 meters, fine
NUMBER 41 39

FIGURE 28.—Grubeulepis tebblei, new species (holotype, BMNH 1953: 1: 623): a, Prostomium,

dorsal view, turned slightly to right; stippled area indicates point of attachment to II; b, tentacular parapodium (I), lateral view; c, parapodium from segment III, anterior view; d, bilimbate

neuroseta from same; e, middle branchial parapodium, anterior view; /, notopodial hooks from

same; g, neurosetae from same.

muddy sand—paratype (F. Rullier, private collection).

DESCRIPTION.—Length of holotype 11.5 mm,

width, including setae, 4 mm, segments 33, last one

small. Length of paratype from Adriatic 17 mm,

width, including setae, 3.5 mm, segments 35. Length

of paratype from Togo 11 mm, width, including

setae, 4 mm, segments 33, last one very small. Elytra

12 pairs, becoming more elongate posteriorly (Figures $0e-h; 31/-A); first pair subtriangular, each with
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 29.—Grubeulepis tebblei, new species (hototype, BMNH 1953: 1: 623): a, Lamelligerous

parapodium from posterior region, anterior view; b, notopodial hooks from same; c, upper

neurosetae from same; d, lower neuroseta from same; e, left third elytron; /, left fifth elytron;

g, left eighth elytron; h, right twelfth elytron.
NUMBER 41 41

FIGURE 30.—Grubeulepis augeneri, new species (paratype MNHNP): a, Prostomium, left lateral

view; stippled area indicates point of attachment to II; b, tentacular parapodium (I), medial

view; c, parapodium from segment II, anterior view; d, neurosetae from segment IV; e, left first

elytron; /, left third elytron; g, left seventh elytron; h, left twelfth elytron.

12-13 papillae on anterior and lateral borders; rest

of elytra subreniform to subrectangular, lateral

borders with single notch or up to 8 digitiform or

wide processes between anterior and posterior

rounded lobes, some biarticulate. Branchiae 12

(holotype and paratype from Togo) or 13 (paratype

from Adriatic) pairs, inflated, with distal branchial

cirrus (Figure 31a). Dorsal cirri, on segments 3 and

6, subulate. Posterior lamellae, beginning on segment 28 or 29, subcordiform to subconical (Figure

31&).

Prostomium covered by segment II and attached

middorsally on posterior half; median antenna

short, conical, inserted anterodorsally on prostomium; lateral antennae conical, inserted more ventrally; palps elongate-tapered, extending beyond

tentacular cirri; 2 pairs small eyes on lateral border;

nuchal organs oval, lateral to prostomium (Figure

30a). Tentacular para podia (I) narrower basally,

enlarged distally, each with dorsal and ventral

tentacular cirri, subequal in length or ventral

slightly longer, 2 acicula and 2 bundles of smooth
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 31.—Grubeulepis augeneri, new species (holotype, ZMH 592): a, Middle branchial parapodium, anterior view; b, posterior lamelligerous parapodium, posterior view; c, upper pectinate

neuroseta from same; d, upper neurosetae from same; e, lower neuroseta from same; /, right

first elytron; g, right second elytron; h, right third elytron; i, right fifth elytron; /', right eighth

elytron; k, right twelfth elytron.
NUMBER 41 43

and spinous capillary setae (Figure 306). Ventral

buccal cirri on segment II thicker and longer than

those following (Figure 30c).

Biramous parapodia supported by light ambercolored acicula and neuropodial hammer-shaped

distal plates (Figure 31a). Notopodial acicula with

hooked tips. Notopodial capillary setae smooth and

spinous, forming long, spreading bundles on posterior parts of notopodia; stout light amber-colored

notopodial hooks, beginning on segment III,

smooth, with tips tapered to fine tips or flattened

spatulate. Neuropodial acicula with hammer-shaped

distal plates. Neurosetae of several kinds: 1-2 upper

pectinate setae (Figure 31c); limbate capillary neurosetae with long tapering tips (Figure 30d); lower

more slender nonlimbate neurosetae; without acicular neurosetae in anterior segments. Upper neurosetae of posterior region stouter than lower ones,

curved downward, tapering to fine tips, faintly

spinous along bend (Figure $\b,d,e). Ventral cirri of

anterior few segments subulate, tapered; following

ventral cirri globular, with short clavate tips (Figure $la,b). Pygidium with single long anal cirrus on

right side; cirrus finely papillate along one side; left

cirrus short, rudimentary. Pharynx not extended.

DISTRIBUTION.—West Africa, Central-east Atlantic

(French Congo, Togo), Adriatic. In 34 meters.

Questionable eulepethid: genus and species

questionable

Eulepis challengeriae

Mclntosh, 1885, p. 134, pi. 20: fig. 1, pi. 23: fig. 1, pi. 24:

fig. 1, pi. 14A: figs. 7, 8.

Pareulepis challengeriae.—Hartman, 1959, p. 123.

MATERIAL EXAMINED.—Challenger station, off

Sombrero and St. Thomas, West Indies, 713 to 823

meters—holotype (BMNH 1885: 12: 1: 104).

REMARKS.—The holotype consists of an anterior

fragment of 18 segments. Only three elytra remain,

the left one on segment 13 and a pair on segment 15.

The elytral margins are entire, with only a slight

indication of a lateral notch. The absence of a

distinct notch or fimbriated external margin on the

elytra sets E. chalengeriae apart from the other

species of eulepethids. Most of the notopodial hooks

are broken off; one of the remaining ones shows a

spinous margin along the bend, as indicated by

Mclntosh (1885, pi. 14a: fig. 7). Because of the lack

of the posterior end and the poor condition of the

single specimen, the generic status and additional

specific characters of E. challengeriae must remain

doubtful.

Literature Cited

Augener, H.

1906. Westindische Polychaeten. Bulletin of the Museum

of Comparative Zoology at Harvard College in Cambridge, 43: 91-196, plates 1-8.

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1927. Polychaeten von Curasao. Bijdragen tot de Dierkunde, Amsterdam, 25: 39-82,9 figures.

Berkeley, E., and Berkeley, C.

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intertidal and estuarine Polychaeta of Natal and

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1962. Polychaeta from Several Localities in the Western

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1967. A Monograph on the Polychaeta of Southern Africa.

Part 1. Errantia. The British Museum (Natural History), Publication No. 656: 1-458, 108 figures.

Fauvel, P.

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Bulletin du Muse"um National d'Histoire Naturelle,

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1939. Annelides polychetes de l'lndochine recueillies par

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1878. Annulata Semperiana. Memoires de VAcademie Imperiale des Sciences de St.-Petersbourg, ser. 7, 25 (8):

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44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hartman, O.

1939. Polychaetous Annelids. Part I. Aphroditidae to Pisionidae. Allan Hancock Pacific Expeditions, 7 (1):

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1942. Report on the Scientific Results of the Atlantis Expeditions to the West Indies under the Joint Auspices

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1959. Catalogue of the Polychaetous Annelids of the World.

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figures.


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