KRISTIAN FAUCHA A Revision of Six

Species of the

Fla vus-Bidentatus

W Group of Eunice

f (Eunicidae: Polychaeta)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 6

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 6

Fauchaid A Revision of Six Species

of the Flavus-Bidentatus

Group of Eunice

(Eunicidae: Polychaeta)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON
SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTIO N

The emphasis upon publications as a means of diffusing knowledge was expressed

by the first Secretary of the Smithsonian Institution. In his formal plan for the Institution, Joseph Henry articulated a program that included the following statement:

"It is proposed to publish a series of reports, giving an account of the new discoveries

in science, and of the changes made from year to year in all branches of knowledge

not strictly professional." This keynote of basic research has been adhered to over the

years in the issuance of thousands of titles in serial publications under the Smithsonian

imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and

continuing with the following active series:

Smithsonian Annals of Flight

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in History and Technology

In these series, the Institution publishes original articles and monographs dealing with

the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts,

synoptic interpretations of data, or original theory in specialized fields. Each publication is distributed by mailing lists to libraries, laboratories, institutes, and interested

specialists throughout the world. Individual copies may be obtained from the

Smithsonian Institution Press as long as stocks are available.

S. DILLON RIPLEY

Secretary

Smithsonian Institution

Contribution number 324 from the Allan Hancock Foundation

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

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1969

For tale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price 30 cents (paper coyer)
ABSTRACT

The species here revised have yellow, bidentate subacicular hooks and branchiae

limited to a short anterior region. They include E. biannulata Moore (1904), E.

kobiensis Mclntosh (1885, holotype examined), E. segregata (Chamberlin, 1919a,

restricted), E. semisegregata, new species, E. valens (Chamberlin, 1919b, types examined), and E. websteri, new name for E. longicirrata Webster (1884, holotype

examined). The relationship between the six species is discussed.

The species here revised include E. biannulata

Moore (1904), E. kobiensis Mclntosh (1885), E.

segregata (Chamberlin, 1919a, restricted), E. semisegregata, new species, E. valens (Chamberlin, 1919b),

and E. websteri, new name for E. longicirrata Webster

(1884). The six species all belong to the flavus-bidentatus group as defined by Hartman (1944, p. 100) in that

they have yellow, bidentate subacicular hooks.

Branchiae are present from setiger three and limited to

a short anterior region in all species here treated.

Eunice valens has been found intertidally and in

shallow subtidal areas in California and Washington.

E. biannulata is known from shallow water in California and western Mexico. E. websteri is known from

Bermuda and possibly from the West Indies. Authentic

records of E. kobiensis come from shelf-depths off

Japan; records from the eastern Pacific Ocean are

limited to British Columbia and Washington (material

from Berkeley and Berkeley's collections in USNM).

These four species are found in shallow water and do

not appear to occur on the deeper continental slopes.

The two last species, E. segregata and E. semisegregata,

come from slope-depths; E. segregata is known from

Panama to southern California and E. semisegregata

from one locality off southwestern Mexico.

The characters used to separate the six species include the relative length of the occipital tentacles, the

maximal number of branchial filaments, the shape of

the composite hooded hooks, the distribution and shape

of the subacicular hooks, and the relationship between

the length of the prostomium and the peristomial segKristian Fauchald, Allan Hancock Foundation, University of

Southern California, Los Angeles, California 90007.

ments. A compilation of some of these characters and

others is given in Table 1.

The shape of the different parapodial lobes appears

to be constant within a given range of segments, but it

was considered inadvisable to investigate this further

on the sparse material of type-specimens available.

Three regions may be recognized along the body: prebranchial, branchial, and postbranchial. One parapodium from each region was examined in detail and

illustrated; other parapodia from the same region were

compared superficially. Provisionally it appears that

some species (E. valens and E. segregata) have very

characteristic parapodial lobes. The development

of the different parapodial lobes is most distinct in the

branchial segments; lobes from the prebranchial and

postbranchial regions tend to be similar in all species

examined. The character of the parapodial lobes cannot be used taxonomically except in conjunction with

other characters since the lobes remain unknown in

most species of Eunice.

Type-specimens were borrowed from different

museums; I am indebted to the following institutions

and persons for allowing me to examine types placed

in their care: the trustees of the British Museum (Natural History) (BMNH, Dr. J. David George), Museum of Comparative Zoology, Harvard University

(MCZ, Dr. H. Levi), and United States National

Museum (USNM, Dr. Marian H. Pettibone).

The study was supported by grant number B5-1780

from the National Science Foundation to the Allan

Hancock Foundation. The Allan Hancock Foundation

not only permitted me to examine specimens in their

collections but gave me material support as well. Dr.

Olga Hartman and Dr. Marian Pettibone gave

valuable advice and critically read the manuscript.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Species Included in this Revision

1. Maximal number of branchial filaments 8 or less 2

Maximal number of branchial filaments 11 or more 3

2. Occipital tentacles with moniliform articles 1 • E. biannulata Moore

Occipital tentacles with cylindrical articles 2. E. kobiensis Mclntosh

3. Maximal number of branchial filaments 34-38 3. E. semisegregate, new species

Maximal number of branchial filaments less than 20 4-

4. Median occipital tentacle reaches setiger 8 5

Median occipital tentacle barely reaches setiger 2 4. E. valens (Chamberlin)

5. Composite hooks with strongly curved distal teeth; anterior dorsal cirri with seven or eight

articulations 5. E. websteri, new name

Composite hooks with nearly straight distal teeth; anterior dorsal cirri with one or two

articulations 6. E. segregate (Chamberlin)

TABLE 1.—Some taxonomic characters of six species of Eunice

[Based on the specimens here treated]

Species of

Eunice

biamudata

kobiensis

segregate

sanisegregate

valens

websteri

Branchiae

present on

setigers

(.number)

3-53

3-41

3-39

3-69

3-76

3-53

Maximal

number of

branchial

filaments

6-8

8

12-15

34-38

11

15

Structure

of first

bronchia

single

single

single

pectinate

single

single

Occipital

tentacles

moniliform

cylindrical

cylindrical

cylindrical

cylindrical

cylindrical

Subacicular

hooks present

from setiger

(number)

36

30

36

51

43

31

Shape of

distal

tooth of

composite

hook

curved

curved

straight

straight

straight

curved

Structure

of dorsal

cirri of

anterior

segments

articulated

smooth

articulated

smooth

articulated

articulated

1. Eunice biannulata Moore, 1904

FIGURE 1

Eunice biannulata Moore, 1904, pp. 487-490, pi. 37: figs.

10-18, pi. 38: fig. 42.—Berkeley and Berkeley, 1939, p.

335.—[?] Hartman, 1939, p. 13.—Fauchald [in press].

Eunice longicirrata.—Hartman, 1944, pp. 104-107, pi. 6:

figs. 118-122, partim [not E. longicirrata Webster].

MATERIAL.—La Jolla, shale and sandstone reefs just

north of caves, 1940, Allan Hancock Foundation station 1210-40, 1 specimen (USNM 38983).

REMARKS.—The type of E. biannulata was collected

by E. C. Starks at San Diego. The present specimen

comes from La Jolla and fits the original description

well except for some trivial differences mentioned

below. It is a complete specimen with 192 setigers and

is 148 mm long and 4.5 mm wide with setae.

The median occipital tentacle had 21 articles, the

inner lateral ones had 13, and the outer lateral tentacles

had 7 each in the type-specimen (Moore, 1904, p.

488). The median tentacle has 19 articulations (Figure

Id), the inner lateral ones have 11 and the outer

lateral tentacles have 5 each in the present specimen.

The dorsal cirri were described as having three

articles in anterior setigers, the two distal ones together

as long as the proximal one. The dorsal cirri in the

present specimen (Figure la) have three articulations,

but the two distal articles together are shorter than the

proximal one. Maximal number of branchial filaments

(Figure le) is seven in the present specimen; it was

originally described as six to eight.

The setae (Figure \b) are similar to those described

by Moore; the subacicular hooks (Figure \c) are

present from setiger 36 in the La Jolla specimen under
NUMBER 6

FIGURE 1.—Eunice biannulata Moore (USNM 38983): a, second parapodium, X25; b, composite hooded hook, 60th setiger, X570; c, subacicular hook, 60th setiger, X285; d, anterior end,

X10; e, 28th parapodium, X25.

review and were present from setigers 40-50 in the

type-specimens.

DISCUSSION.—E. biannulata is the only species

known in this group (flavxis-bidentatus with a limited

number of branchiae) that has occipital tentacles with

moniliform articles.

Eunice biannulata was separated from E. websteri

by Berkeley and Berkeley (1939, p. 335 as E. longicirrata) on the number of articulations of the anterior

dorsal cirri. The number of articulations has been

shown to be somewhat variable (Fauchald, in press)

and cannot be used as the only character to separate

the two species. All specimens from southern California and western Mexico reported by Berkeley and

Berkeley (1939) have monilif orm tentacles. Other

records of this species and E. websteri ( = E. longicirrata auctores) from the eastern Pacific Ocean have

been discussed in some detail elsewhere (Fauchald, in

press).

DISTRIBUTION.—E. biannulata is known from southern California and western Mexico (Moore, 1904;

Berkeley and Berkeley, 1939; Hartman, 1944 and

Fauchald, in press). It is found intertidally and in very

shallow water. The possible wider distribution is not
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

known since E. biannulata has been confused with several related species.

2. Eunice kobiensis Mclntosh, 1885

FIGURE 2

Eunice kobiensis Mclntosh, 1885, pp. 278-280, pi. 38: figs.

12-13, pi. 20A: figs. 1-3.—Izuka, 1912, pp. 117-118, pi.

13: figs. 11-12.—Berkeley and Berkeley, 1948, p. 90, fig.

135. [Not E. kobiensis Akesson, 1967.]

[?]Eunice gracilis Moore, 1903, pp. 440-441, pi. 25: figs.

46-48.

Eunice longicirrata.—Imajima and Hartman, 1964, p. 256

[not E. longicirrata Webster].

MATERIAL.—Holotype (BMNH 1885:12:1:197) off

Kobe, Japan, Challenger.

DESCRIPTION.—The holotype is an incomplete specimen with 131 setigers; it is 55 mm long and 3.5 mm

wide with setae. It is brown and lacks color pattern.

The prostomium (Figure 2/) is twice as wide as long;

the palpi are visible from the dorsal side and the

anterior incision reaches the base of the median occipital tentacle. All occipital tentacles are slender; the

outer lateral ones reach the first setiger; each has four

articulations. The -inner lateral tentacles reach the

fourth or the fifth setiger; each has five articulations.

The median tentacle reaches the tenth or eleventh

setiger and has seven articulations. All articles are

cylindrical. A pair of eyes is present posterior to the

bases of the outer lateral occipital tentacles.

The cylindrical first peristomial segment is twice as

long as the first setiger; the second peristomial segment

is slightly shorter than the first setiger; the slender

peristomial cirri are smooth and barely reach the

anterior edge of the first peristomial segment.

Prebranchial parapodia (Figure 2b) have short and

truncate presetal and postsetal lobes; the setal lobes

are truncate with a short dorsal extension covering the

dorsalmost aciculum. The slender dorsal cirri are

nearly as long as half the width of the body; each has

one basal articulation. The large ventral cirri are

digitate. Parapodia in the branchial region (Figure 2g) have rounded parapodial lobes; all lobes are

of the same length and there is no prolongation of the

setal lobes associated with the acicula. The dorsal cirri

are shorter and slenderer than in the prebranchial segments; each retains one basal articulation. The ventral

cirri have slight basal swellings, but retain the large

digitate tip. Parapodia in the postbranchial region

(Figure 2a) have short, truncate parapodial lobes; the

long dorsal cirri are smooth; the ventral cirri are

digitate and somewhat slenderer than in the prebranchial region; the basal swellings are absent.

Branchiae (Figure 2g) are present on setigers 3-41;

the three first and the last two pairs are single filaments;

all others are pectinate. The maximal number of branchial filaments is eight, but most branchiae have only

six or seven. The branchial stem is short and thick; the

branchiae are not held erect over the dorsum; the

branchial filaments are long.

Setae are broken off in most setigers; anterior setigers have simple limbate setae and composite hooks.

The composite hooks (Figure 2c) have very short appendages; each has a thick, curved distal tooth and a

small, conical proximal tooth. The short hoods are

blunt; the proximal part of the hoods and the upper

ends of the shafts are serrated. Pectinate setae (Figure

2e) are present in median and posterior setigers; each

is distally straight with one slender whiplike prolongation and eleven to twelve teeth. Subacicular hooks

(Figure 2d) are present from setiger 30 to the end of

the fragment; one or two hooks are present in a parapodium. Each hook has a large proximal tooth and a

small distal tooth; the hoods are short and blunt.

The pharyngeal apparatus has been dissected out

and was found free in the tube with the specimen; it

resembles the figure in Mclntosh (1885, fig. 37)

closely and it is assumed to be from the type-specimen.

Maxilla I is falcate; maxilla II has seven teeth left and

eight right; left maxilla III has seven teeth; the

combined right maxillae III + IV have twelve teeth;

left maxilla IV has ten teeth; both maxillae V and the

mandibles are missing, but are assumed to have been

present.

DISCUSSION.—E. kobiensis resembles E. segregata

and E. websteri. The maximal number of branchial

filaments is 8 in E. kobiensis and 12-15 and 15 in E.

segregata and E. websteri, respectively. The development of the parapodial lobes in the branchial regions

differs markedly in the three species and the proportions of the anterior end are different.

Eunice gracilis has been considered synonymous with

E. kobiensis (Imajima and Hartman, 1964, p. 256, as

E. longicirrata); it apparently differs from E. kobiensis

in the maximal number of branchial filaments which is

eleven in E. gracilis (Moore, 1903, p. 441) and eight

in E. kobiensis. The median occipital tentacle appears

to be somewhat shorter in relation to the inner lateral

ones in E. gracilis than in E. kobiensis. The differences
NUMBER 6

FIGURE 2.—Eunice kobiensis Mclntosh (BMNH 1885:12:1:197): a, 126th parapodium, X50;

b, second parapodium, X50; c, composite hooded hook, 126th setiger, X570; d, subacicular

hook, 126th setiger, X570; e, pectinate seta, 28th setiger, X635; /, anterior end, X10; g, 28th

parapodium, X50.
are slight, and the two species are here doubtfully considered synonymous.

Eunice longicirrata Imajima and Hartman (1964)

fits the description of E. kobiensis rather than that of

E. websteri [=E. longicirrata Webster].

Records from the eastern Pacific Ocean (Berkeley

and Berkeley, 1948, p. 90) are limited to Washington

and British Columbia. E. kobiensis Akesson (1967) is

here referred to E. valens; the illustration of the anterior end of the adults (Akesson, 1967, fig. 21) shows

that the specimens had very short occipital tentacles

and the relationship between the prostomium and the

peristomial segments is similar to that found in E.

valens. The setae found in the larvae (Akesson, 1967,

fig. 8) do not resemble setae in any of the six species

here considered and may be characteristic larval setae.

DISTRIBUTION.—E. kobiensis is known from Japan

in 8 to 50 fathoms, but may be widely distributed in

the western Pacific Ocean.

6. Eunice segregate (Chamberlin, 1919a) restricted

FIGURE 3

Leodiee segregata Chamberlin, 1919a, pp. 237-240, pi. 54:

figs. 1-4, partim.—Treadwell, 1923, p. 7, partim.

Eunice segregata.—Fauchald pn press].

MATERIAL.—Off southern California 33°33' N, 120°

I7'3O" W, 534 fathoms, agassiz trawl, green sand, Globigerina, 26 April 1911, Albatross station D 5695, one

specimen (USNM 19153).

REMARKS.—Leodiee segregata was based on four

specimens from three Albatross stations: stations 3354

and 3358 from off Panama (each with a single specimen) and station 3417 from off southwestern Mexico

(two specimens). Only the two specimens from station

3417 have been recovered, one is in the United States

National Museum (USNM 19398) and the other in

the Museum of Comparative Zoology, Harvard University. These two specimens do not agree with the

original description and were mentioned by Chamberlin (1919a) as different from the two specimens from

Panama; they form the basis for a new species, E.

semisegregata (see below).

A specimen from Albatross station D 5695 off southern California, identified by Treadwell (1923) as

Leodiee segregata (USNM 19153), agrees closely with

the original description of Chamberlin and has been

used as a basis for a redescription of the species.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REDESCRIPTION.—The present specimen is incomplete and has 92 setigers; it is 60 mm long and 4.7 mm

wide with setae and is olive colored without color pattern. The body is cylindrical except for a slight dorsal

flattening in the posterior end.

The prostomium (Figure 3g) is slightly wider than

long; the anterior incision is deep and narrow. Eyes

are absent. The palpi are poorly marked dorsally; the

ventral parts are rounded. The occipital tentacles are

in a crescentic arrangement with the outer lateral pair

in front of the others. All occipital tentacles are articulated, but the articulations are faint and visible only

as septa inside the tentacles. Each of the outer lateral

occipital tentacles reaches setiger 3 and has seven or

eight articulations; each of the inner lateral ones

reaches setiger 6 and has nine to ten articulations. The

median tentacle reaches setiger 8 and has ten

articulations.

The cylindrical first peristomial segment is slightly

longer than the prostoniurn. The second peristomial

segment is as long as the first setiger. The peristomial

cirri reach the middle of the prostonium; each has

eight or nine faint articulations.

The first setiger (Figure 3e) has low, transverse presetal and postsetal lobes; the setal lobe is conical. Both

cirri are well developed; the large, digitiform dorsal

cirrus is nearly as long as hah* the width of the body

and has two articulations. Dorsal cirri in the branchial

region (Figure 3a) have one articulation; those in

the postbranchial region (Figure 3/) are smooth. The

dorsal cirri decrease gradually in size from setiger 3

to the last setiger. The blunt ventral cirrus in the first

setiger is thick; basal swellings are present in branchial

and postbranchial setigers, but become less distinct in

the posterior end; ventral cirri are shorter in the posterior than in the anterior end.

The branchial segments (Figure 3a) have blunt,

well-developed setal lobes; the presetal lobes are low,

transverse folds that continue round the ventral edges

of the setal lobes. The large postsetal lobes are leaf

shaped and envelope the setal lobes and the presetal

lobes from behind. Postbranchial setigers (Figure 3/)

have reduced postsetal lobes and the setal lobes are

shorter than in the branchial setigers.

Branchiae (Figure 3a) are present on setigers 3-39;

the first two and the last pair are single filaments; all

others are pectinate. The maximal number of branchial

filaments is 12-15 and the branchiae cover the dorsum

completely where best developed.
NUMBER 6

FIGURE 3.—Eunice segregate. (Chamberlin) (USNM 19153): a, 21st parapodium, X25; b,

pectinate seta, second setiger, X1350; c, composite hooded hook, second setiger, X570; d, subacicular hook, posterior setiger, X570; e, second parapodium, X25; /, posterior parapodium,

X25; g, anterior end, X10.

829-129 O—69 2
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The dorsal setal fascicles have numerous slender

capillary setae with serrated cutting edges and a number of pectinate setae in all setigers. Each pectinate

seta (Figure 36) is narrow and straight with 6-12

teeth and one margin prolonged; pectinate setae have

fewer (6-8) teeth in anterior, than in posterior

(11—12) setigers. Composite, bidentate hooded hooks

(Figure 3c) with blunt hoods are present in all ventral

fascicles. Each hook has a slightly curved distal tooth

and the proximal tooth is bluntly conical. The hoods

and the upper ends of the shafts are serrated. Acicula

number two in a parapodium except in far posterior

setigers where only one is present; each is conical and

has a slightly bent tip. Subacicular hooks (Figure 3d)

are present from setiger 36 to the end of the fragment.

Each is yellow and bidentate with poorly marked neck;

the proximal tooth is larger than the distal one and

blunt. The hoods are short and rounded.

The pharyngeal apparatus was not dissected.

DISCUSSION.—E. segregata as redescribed here differs in some respects from the original description

(Chamberlin, 1919a). The occipital tentacles are obscurely articulated, not smooth as stated by Chamberlin. Two of the original paratypes of E. segregata (as

E. semisegregata below) have clearly articulated occipital tentacles, so apparently Chamberlin overlooked

this character. Branchiae are present on setigers 3-39

in the present specimen; the holotype had branchiae on

setigers 3-43 according to Chamberlin. The maximal

number of branchial filaments was 21 in the holotype

according to Chamberlin; the present specimen has

12-15.

These differences are here considered minor and

except for the discrepancy in the number of branchial

filaments well within the patterns of variability established for other species in the genus (Fauchald, in

press).

Eunice segregata resembles E. kobiensis, but differs

from the latter in the structure of the parapodial lobes;

the difference is most apparent in the branchial region.

Presetal and postsetal lobes are distinct and differ in

length from the setal lobes in E. segregata and are similar in length and shape in E. kobiensis. The peristomial

cirri in E. segregata reach the middle of the prostomium and barely reach the anterior edge of the first

peristomial segment in E. kobiensis. The prostomium is

shorter than the peristomium in E. segregata and of

the same length as the peristomium in E. kobiensis.

Eyes are absent in E. segregata and present in E.

kobiensis.

Eunice segregata also resembles E. valens and E.

websteri. The composite hooded hooks have strongly

curved distal teeth and slender proximal teeth in E.

websteri and weakly curved distal teeth and blunt,

conical proximal teeth in E. segregata. The anterior

dorsal cirri have several articulations each in E. websteri and only one or two in E. segregata. The occipital

tentacles do not reach the first setiger in E. valens; in

E. segregata even the short outer lateral ones reach the

third setiger.

DISTRIBUTION.—E. segregata is known from slope

depths from southern California to Panama.

3. Eunic semisegregata, new species

FIGURE 4

Leodice segregata Chamberlin, 1919a, pp. 237-240, pi. 54:

fig. 5, partim.

MATERIAL.—Off southwestern Mexico, 16°32'00"N,

99°48'00"W, 493 fathoms, green mud, small beam

trawl, 11 April 1891, Albatross station 3417, two specimens, holotype (Museum of Comparative Zoology,

Harvard University), paratype (USNM 19398).

DESCRIPTION.—The holotype is an incomplete specimen with 91 setigers; it is 65 mm long and 7 mm wide

with setae. It is evenly grey without any color pattern.

The body is cylindrical and only slightly flattened in

the posterior end.

The prostomium (Figure Ac) is shorter than wide

and has a shallow anterior incision, both halves are

rounded. The occipital tentacles are inserted in a

straight line; the outer lateral ones reach the first

setiger; each has three articulations (in the paratype,

broken in the holotype). The inner lateral occipital

tentacles reach setiger 5; each has four articulations.

The median tentacle reaches setiger 7 and has five

articulations. The cylindrical first peristomial segment

is a little longer than the prostomium; the second

peristomial segment is half as long as the first one and

as long as the first setiger. The short peristomial cirri

are smooth; each reaches the middle of the first peristomial segment.

All setigers have low, transverse presetal and postsetal lobes (Figure 4rf). The rounded setal lobes are

longer than the other lobes. The very thick dorsal cirrus is smooth and blunt in the prebranchial region;

the ventral cirrus is shorter than the dorsal one, but
NUMBER 6

FIGURE 4.—Eunice semisegregata, new species (Museum of Comparative Zoology, Harvard

University): a, aciculum, 70th setiger, X570; b, subacicular hook, 70th setiger, X570; c, anterior end, X5; d, 25th parapodium, X10; e, composite hooded hook, 4th setiger, X570.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

similar in shape. The dorsal cirri are slender in the

branchial region; they are similar in shape, but shorter

in the postbranchial region. Ventral cirri are basally

swollen and lack a distinct tip in the branchial region;

a thick, blunt tip reappears in the postbranchial

region, and in the last segments present the ventral

cirri are similar to those in the prebranchial region.

Branchiae (Figure 4d; Chamberlin, 1919a, pi. 54,

fig. 5) are present in setigers 3-69; all branchiae are

pectinate. The first branchia has six branchial filaments; where best developed each branchia has 34-36

(38 in the paratype) branchial filaments. The branchial stem is slender and the long branchial filaments

are thin. The number of branchial filaments is

rapidly reduced posterior to setiger 35; only eight or

nine filaments are present in the last branchia. The

branchiae cover the dorsum completely in most of the

branchial region.

The dorsal setal fascicles have slender, limbate setae

with one cutting edge serrated. Pectinate setae were

not seen, but most setae were broken off; they are

assumed to be present. Ventral fascicles have composite, hooded hooks (Figure \e); each hook has a

straight appendage with a slightly curved distal tooth

and a blunt, conical proximal tooth. The hoods are

blunt, but appear worn. Acicula (Figure 4a) number

two in a parapodium, each is yellow and has a bluntly

conical tip which is slightly bent. Subacicular hooks

(Figure 46) are present from setiger 51; each is

bidentate with both teeth of the same length and

shape.

The pharyngeal apparatus has been dissected; the

following description is rewritten after remarks about

the specimens from this station in Chamberlin (1919a,

p. 239). Maxilla II has eight teeth left and nine right;

left maxilla III has ten or eleven teeth; the combined

right maxillae III + IV have nine teeth; left maxilla TV

has nine teeth. Maxillae V and the mandibles were

not specifically mentioned by Chamberlin for the

specimens from this station.

DISCUSSION.—The specimens here described as E.

semisegregata were specifically mentioned by Chamberlin (1919a, p. 238). He stated that: "In a paratype

from Sta. 3417 the first branchiae, i.e., those of somite

V, have numerous branches, but on posterior somites

the number gradually decreases." As noted above, the

specimens were also mentioned for the differing

pharyngeal apparatus.

Eunice semisegregata differs from E. segregata in

that all branchiae are pectinate in the former; the

latter have the first two and the last branchiae single.

The character of the first branchiae is not generally a

valid specific character; the first simple branchiae are

often lost in large specimens; but in species with

branchiae from setiger 3, the presence and character

of the branchiae are remarkably conservative. This has

been shown for five species of Eunice from western

Mexico (Fauchald, in press).

The peristomial cirri reach the middle of the first

peristomial segment in E. semisegregata and the middle

of the prostomium in E. segregata. The dorsal cirri in

the first setigers are short and thick in E. semisegregata and long and slender in E. segregata. The

maximal number of branchial filaments is 12-15 (possibly 20-21, see comment above) in E. segregata and

34-38 in E. semisegregata. The subacicular hooks are

present from setiger 51 in E. semisegregata and from

setiger 36 in E. segregata.

DISTRIBUTION.—E. semisegregata is known from one

locality off Acapulco, Mexico, in 493 fathoms.

4. Eunice valens (Chamberlin, 1919b)

FIGURE 5

Leodice valens Chamberlin, 1919b, pp. 257-258, pi. 1: figs.

6-8.

Eunice kobiensis.—kktsson, 1967, pp. 141-192, figs. 1-22

[not E. kobiensis Mclntosh, 1885].

lEunice tridentata.—Hartman, 1944, pp. 114-115, pi. 7:

figs. 145-150, partim [not E. tridentata Ehlers, 1905].

MATERIAL.—Mendocino, California, collection of A.

Agassiz, holotype (MCZ 120) and paratype (MCZ

121).

DESCRIPTION.—The holotype is a complete specimen with 179 setigers; it is 140 mm long and 7 mm

wide with setae. It is evenly pink with a slight brown

tinge and lacks color pattern. The anterior end of the

body is cylindrical; the median and posterior end is

slightly flattened dorsally. The anal cirri are as long

as the ten last setigers. The paratype has been dried

out and consists of several unidentifiable fragments.

The prostomium (Figure 56) is wider than long

and has a deep anterior incision; each half is evenly

rounded. A pair of black eyes is present posterior to

the bases of the outer lateral occipital tentacles. Each

of the outer lateral occipital tentacles has five articulations; the inner lateral ones have ten articulations

each and the median tentacle has eleven articulations.
NUMBER 6 11

a

FIGURE 5.—Eunice valens (Chamberlin) (MCZ 120): a, 17th parapodium, X18; b, anterior

end, X5; e, subacicular hook, 150th setiger, X285; d, composite hooded hook, 13th setiger,

X570.

All articles are cylindrical and only faintly marked

externally. The outer lateral pair of tentacles reaches

the second penstomial segment and are somewhat

stouter than the other tentacles. The inner lateral pair

reaches the first setiger and the median tentacle

reaches the second setiger.

The cylindrical first peristomial segment is twice as

long as the first setiger; it projects forward like a collar

and covers the bases of the occipital tentacles. The second peristomial segment is only half as long as the

first setiger. The peristomial cirri are slender; each

has seven or eight articulations and reaches the front

edge of the first peristomial segment

The first setigers have short, transverse presetal

lobes; the setal and postsetal lobes are truncate; the

setal lobes are a little longer than the presetal ones,

but shorter than the postsetal lobes. The parapodia

in the branchial region (Figure 5a) are similar to the

first ones, but the postsetal lobes are reduced to transverse folds in the first branchial segments. The setal

lobes become conical and the presetal and postsetal

lobes are visible only as low folds in the postbranchial

region.

The dorsal cirri are large in the prebranchial region

and in the first branchial setigers; they are surpassed

by the branchiae where these are fully developed even
12

if the cirri themselves are not reduced in size. Dorsal

cirri in the postbranchial region are again very prominent. Each digitiform, thickset cirrus is divided into

two to five irregular articles. The thickset ventral cirri

in the prebranchial and early branchial setigers are

blunt; farther back the bases become very swollen, but

the distinct, blunt tips of the cirri are present in all

setigers.

Branchiae (Figure 5a) are present in setigers 3-76.

The first two and the last eight or nine pairs are

simple filaments; all others are pectinate with long,

stout branchial filaments. The branchiae are best

developed in setigers 15-30; the maximal number of

branchial filaments is eleven.

Composite hooded hooks (Figure 5d) are found in

ventral fascicles in all setigers. They have smooth

shafts which are distally enlarged; each appendage is

triangular with a slightly curved distal tooth; the proximal tooth is blunt and nearly at right angles with the

distal one. The short hoods are blunt and serrated

near the bases of the appendages. Smooth, slightly

curved capillary setae are found in dorsal fascicles in

all parapodia; pectinate setae were not seen. Acicula

number three or four in a parapodium; each is light

yellow and has a strongly bent, conical tip. Subacicular

hooks (Figure 5c) are present from setiger 43 to the

end of the body. Each is tapering evenly and is light

yellow; the proximal tooth is larger than the distal one

and only at a slight angle with the axis of the hook.

The description of the pharyngeal apparatus is

rewritten from Chamberlin (1919b); it was not dissected in the holotype and the condition of the paratype does not permit confirmation of the description.

The mandibles have calcified cutting edges with three

rounded teeth each. Maxilla II has six teeth left and

eight right; left maxilla III has eight or nine teeth;

the combined right maxillae III + IV have eight teeth;

left maxilla IV has six teeth. Chamberlin gave no information on the dentition of either maxillae V; his

illustration of maxillae I (pi. 1: fig. 8) shows that

these are strongly falcate.

DISCUSSION.—E. valens resembles E. biannulata, E.

kobiensis, and E. websteri. It differs from E. biannulata

and E. websteri in that the distal teeth of the composite

hooks are nearly straight in the former and curved in

the two latter. The occipital tentacles are very short in

E. valens, the longest tentacle, which is the median one,

barely reaches the second setiger; the occipital tentacles

reach at least setiger 5 in the three other species.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The specimens reported by Akesson (1967) from

San Juan Archipelago, Washington, as E. kobiensis had

very short occipital tentacles and are here considered

to belong to E. valens.

DISTRIBUTION.—E. valens is known from the typelocality at Mendocino in northern California and from

the San Juan Archipelago, Washington.

5. Eunice websteri, new name

FIGURE 6

Eunice longicirrata Webster, 1884, pp. 318-319, pi. 12:

figs. 74-80.

PLeodice longicirrata.—-Treadwell, 1921, pp. 11-14, figs. 3 -

12, pi. 1: figs. 1-4.

MATERIAL.—Bermuda, collection of G. Brown

Goode, 1876-77, holotype (USNM 4792).

DESCRIPTION.—The holotype is a complete specimen with about 150 setigers; it is about 120 mm long

and 4 mm wide with setae. The parapodia of setigers

1-9 and 21-32 on the left side have been removed and

are mounted on slides; parapodia from the postbranchial region and the whole posterior end also have been

mounted on slides. The measurements given above are

estimates of the length and width of the complete

specimen. The specimen had been dissected for the

pharyngeal apparatus and the illustration of the anterior end is partially reconstructed.

The prostomium (Figure 6c) is wider than long

with a short, narrow anterior incision; both halves are

anteriorly truncate and the lateral margins are nearly

straight. The occipital tentacles are inserted in a crescent; all are about equidistant from each other. A

pair of black eyes is present posterior to the outer

lateral tentacles. The outer lateral occipital tentacles

reach the second peristomial segment; each has five

or six articulations. The inner lateral tentacles reach

setiger 5; each has six or seven articulations. The median tentacle reaches setigers 8 or 9 and has eight

articulations. All articles are cylindrical.

The cylindrical first peristomial segment is as long

as the prostomium. The second peristomial segment

is only one third as long as the first one, but of the

same length as the first setiger. The long peristomial

cirri are slender; they reach the middle of the prostomium ; each has six or seven articulations.

The presetal lobes in the first parapodia (Figure 6e)

are obliquely truncate; the setal lobes are triangular

and the postsetal lobes are low, transverse folds. The

branchial parapodia (Figure 6d) are similar to the
NUMBER 6 13

FIGURE 6.—Eunice websteri, new name (USNM 4792): a, composite hooded hook, first setiger,

X469; b, subacicular hook, median setiger, X192; c, anterior end, X10; d, median parapodium,

X25; *, second parapodiiun, X50.
14

first ones; the setal lobes become more truncate; t)Ut

this may be an artifact since the tissue has recede0-

from the acicula which are left bare over a long dis*

tance. The dorsal cirri are similar in length and shal^

in all parapodia, but are most conspicuous in the pr e "

branchial and postbranchial regions. Each of the dorsa '

cirri in the prebranchial region has seven or eight cylln~

drical articles; those in the branchial region have 0 n e

or two articles each. The postbranchial dorsal ci r n

have four or five articles each. The ventral cirri

 i n

the prebranchial and anterior branchial setigers £r e

thick and digitiform; ventral cirri in the posterior

branchial and postbranchial regions have distinct ba 5^

swellings.

Branchiae (Figure 6d) are present on setigers 3-?3.

The first and the last ten branchiae are single fiia~

ments; all others are pectinate. The branchiae are fu^y

developed on setigers 5-33; the maximal number °f

branchial filaments is fifteen.

Long, slender capillary setae are present in dorsa^

fascicles, and composite, hooded, bidentate hooks

 i n

ventral fascicles in all parapodia. Each hooded ho°k

(Figure 6a) has a strongly curved distal tooth and

 a

sharply pointed proximal tooth at right angles to t n e

axis of the appendage; both teeth are similar in si?e-

The hoods are blunt and serrations are absent. Pec*1"

nate setae were not seen. Acicula number three

 o r

four in a parapodium; each is light yellow and is e'x~

panded distally to form a bluntly conical tip. Sv^~

acicular hooks (Figure 6b) are present from setiger 3^ '•>

they occur singly in most parapodia, but two hoc>ks

are sometimes present. Each is light yellow and b ^

a short distal tooth and a much larger proximal tooth

at right angles to the axis of the hook.

The pharyngeal apparatus is in position in the speP"

men, but the dissection has cut through parts of i*

so the number of teeth cannot be counted; the ja ^

appear to be of the usual kind.

DISCUSSION.—The name E. longicirrata is preoccupied in the combination Eunice (Nicidion) long**

cirrata Kinberg (1865, p. 564) which is a synonym oi

E. (N.) cariboea Grube (see Hartman, 1948, p. 8O0-

The species from Bermuda is renamed here in hono r

of Dr. G. E. Webster who originally described it. 1Tie

records by Treadwell (1921) from the West Indies ar e

doubtful; the occipital tentacles were described as

articulated and shown in the illustration (pi. 1, fig. 2)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

as nearly moniliform and the composite hooks (textfig. 9) were shown to have very short proximal teeth.

Eunice websteri has been confused with at least

three of the species treated here, E. biannulata, E.

valens, and E. kobiensis; the relationship between the

four species has been discussed above and is tabulated

in Table 1.

DISTRIBUTION.—E. websteri is known from Bermuda

and possibly from the West Indies. It may be widely

distributed in this area, but it has been confused with

other species so the total distribution remains unknown.

Literature Cited

Akesson, B.

1967. The Embryology of the Polychaete Eunice kobiensis. Ada Zoologica, 48:141-192, figures 1-22.

Berkeley, E., and C. Berkeley

1939. On a Collection of Polychaeta, Chiefly from the

West Coast of Mexico. The Annals & Magazine

of Natural History, London, series 11, 3:321-346,

figures 1-12.

1948. Annelida, Polychaeta Errantia. Canadian Pacific

Fauna, 9b(l): 1-100, figures 1-160.

Chamberlin, R. V.

1919a. The Annelida Polychaeta. Memoirs of the Museum of Comparative Zoology, Harvard University,

48:1-514, plates 1-80.

1919b. Pacific Coast Polychaeta Collected by Alexander

Agassiz. Bulletin of the Museum of Comparative

Zoology, Harvard University, 63:251-276, plates

1-3.

Ehlers, E.

1905. Anneliden der Sammlung Schauinsland. Ergebnisse

einer Reise nach dem Pacific, Schauinsland 1896—

97. Zoologische Jahrbu'cher Abt. fur Systematik,

Okologie und geographie der tiere, Jena, 22:281—

302, plate 9.

Fauchald, K.

In press. Polychaetous Annelids of the Families Eunicidae,

Lumbrineridae, Iphitimidae, Arabellidae, Lysaretidae and Dorvilleidae from Western Mexico. 350

manuscript pages, 25 plates. Monographs in Marine Biology, Allan Hancock Foundation.

Hartman, O.

1939. The Polychaetous Annelids Collected by the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, 98:1-22, figures 1-3.

1944. Polychaetous Annelids, part 5: Eunicea. Allan

Hancock Pacific Expedition, 10(1): 1-238, plates

1-18.

1948. The Marine Annelids Erected by Kinberg with

Notes on Some Other Types in the Swedish State

Museum. Arkiv for Zoologi, Stockholm, 42A(1) :

1-137, plates 1-18.
NUMBER 6

Imajima, M., and Hartman, O.

1964. The Polychaetous Annelids of Japan. Allan Hancock Foundation Publications, Occasional Paper,

26:1-452, plates 1-38.

Izuka, A.

1912. The Errantiate Polychaeta of Japan. The Journal

of the College of Science, Imperial University of

Tokyo, Japan, 30(2): 1-262, plates 1-24.

Kinbcrg, J. G. H.

1865. Annulata Nova. Ofversigt af Kongl. VetenskapsAkademiens Forhandlingar, Stockholm, 21:559-

574.

Mclntosh, W. C.

1885. Report on the Annelida Polychaeta Collected by

H.M.S. "Challenger" during the Years 1873-76.

In Reports on the Scientific Research of the "Challenger/' Zoology, 12:1-554, plates 1-55 and

1A-39A.

15

Moore, J. P.

1903. Polychaeta from the Coastal Slope of Japan and

from Kamchatka and Bering Sea. Proceedings of

the Academy of Natural Sciences of Philadelphia,

55:401-490, plates 23-27.

1904. New Polychaeta from California. Proceedings of

the Academy of Natural Sciences of Philadelphia,

56:484-503, plates 37-38.

Treadwell, A. L.

1921. Leodicidae of the West Indian Region. Publication, Carnegie Institution of Washington, 293:

1-131, figures 1-467, plates 1-9.

1923. Polychaetous Annelids from Lower California with

Description of New Species. American Museum

Novitates, New York, 74:1-11, figures 1-8.

Webster, H. E.

1884. Annelida from Bermuda. United States National

Museum Bulletin, 25:305-327, plates 7-12.

US. GOVERNMENT PRINTING OFFICE: 1000 O—329-129



Publication in Smithsonian Contributions to Zoology

Manuscripts for serial publications are accepted by the Smithsonian Institution Press, subject

to substantive review, only through departments of the various Smithsonian museums. NonSmithsonian authors should address inquiries to the appropriate department. If submission is

invited, the following format requirements of the Press will govern the preparation of copy. (An

instruction sheet for the preparation of illustrations is available from the Press on request.)

Copy must be typewritten, double-spaced, on one side of standard white bond paper, with

V/z" top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompanied

by the original artwork. Duplicate copies of all material, including illustrations, should be retained

by the author. There may be several paragraphs to a page, but each page should begin with a new

paragraph. Number consecutively all pages, including title page, abstract, text, literature cited,

legends, and tables. The minimum length is 30 pages of typescript and illustration.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic

titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters

and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings

may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date, page) with a full reference at the end

of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent

lines indented about three spaces. Within a taxon, use a period-dash (.—) to separate each

reference. Enclose with square brackets any annotation in or at the end of the taxon. For

references within the text, use the author-date system: "(Jones, 1910)" or "Jones (1910)!" If

the reference is expanded, abbreviate the data: "Jones (1910, p. 122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively;

however, if a group of figures is treated as a single figure, the individual components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 96) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on pages

separate from the text and not attached to the artwork.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and

page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when

used alone).

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.