Polychactcs from Inter tidal Areas

in Panama, with a Review of

Previous Shallow-Water Records

KRISTIAN FAUCHALD

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 22 1
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SMITHSONIA N CONTRIBUTION S TO ZOOLOGY • NUMBE R 221

Polychaetes from Intertidal Areas

in Panama, with a Review of

Previous Shallow-Water Records

Kristian Fauchald

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Fauchald, Kristian. Polychaetes from Intertidal Areas in Panama, with a Review of Previous Shallow-Water Records. Smithsonian Contributions to Zoology,

number 221, 81 pages, 13 figures, 2 tables, 1977.—A total of 180 species are

listed from both coasts of Panama. The material includes specimens collected

by the Smithsonian Tropical Research Institute and specimens previously reported by Monro and Hartman in a series of papers. Newly described species

are Aphrodita diplops, Eupanthalis perlae, Eunereis paitillae, Neanthes galetae,

Neanthes pseudonoodti, Nereis panamensis, Marphysa amadae, Isolda bipinnata, and Euthelepus pascua. The eastern Pacific Ocean appears to be more

species-rich than the western Atlantic, and relatively fewer species appear widespread in the Pacific than in the Atlantic Ocean. This fact may be because most

of the sampling has been done on hard substrates in the Atlantic, while more

diverse areas have been sampled in the Pacific. Generally, hard-substrate polychaetes are more widely dispersed than soft-bottom forms.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SEMES COVER DESICN- The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Fauchald, Kristian.

Polychaetes from intertidal areas in Panama, with a review of previous shallow-water records.

(Smithsonian contributions to zoology ; no. 221)

Bibliography: p.

Includes index.

1. Polychaeta—Panama. 2. Polychaeta—Pacific Ocean. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 221

QL1.S54 no. 221 [QL391.A6] 591.5 [595M47'097287] 76-6967
Contents

Page

Introduction 1

Family Aphroditidae 2

Family Polynoidae 4

Family Polyodontidae 7

Family Sigalionidae 9

Family Chrysopetalidae 10

Family Amphinomidae 11

Family Phyllodocidae 13

Family Hesionidae 16

Family Pilargiidae 17

Family Syllidae 17

Family Nereidae 22

Family Nephtyidae 33

Family Glyceridae . 34

Family Goniadidae 35

Family Onuphidae 35

Family Eunicidae 37

Family Lumbrineridae 43

Family Arabellidae 44

Family Lysaretidae 45

Family Dorvilleidae 46

Family Orbiniidae 46

Family Spionidae 47

Family Magelonidae 48

Family Chaetopteridae 48

Family Cirratulidae 49

Family Flabelligeridae 51

Family Opheliidae 52

Family Capitellidae 52

Family Maldanidae 53

Family Oweniidae 53

Family Sabellariidae 53

Family Ampharetidae 55

Family Terebellidae 56

Family Trichobranchidae 60

Family Sabellidae 60

Biogeography 63

Literature Cited 69

Index 76

Polychaetes from Intertidal Areas

in Panama, with a Review of

Previous Shallow-Water Records

Kristian Fauchald

Introduction

Intertidal polychaetes were collected from Panamanian rocky shores in connection with a study

of the effects of oil pollution at Galeta Reef in the

Atlantic Ocean, and at Paitilla Beach on the Pacific side of Panama. A representative sample of

the polychaetes were submitted to me for identification. This sample contains 90 species representing

most of the common shallow-water families.

Polychaetes from Panama have been mainly reported by Hartman in a series of papers from the

Allan Hancock Foundation and by Monro in four

papers (1928a, 1928b, 1933a, 1933b). Very few of

the species reported by Hartman and Monro were

recovered in the recent collections, but other members of the same families or genera were found.

This made a complete review of all available materials necessary.

At present, 180 species of polychaetes are known

from Panama. An additional 18 taxa are identified

as to genus, but are known to be different from the

species already named from the area. Nine species

are newly described.

The overall sampling is skewed since the recent

sampling on rocky shores was quantitative and comprehensive, while samples from sandy and muddy

bottoms taken between 1914 and 1937 consisted

largely of dredge samples and informal shovel samKristian Fauchald, Allan Hancock Foundation, University of

Southern California, Los Angeles, California 90007.

pies. The number of soft-bottom forms is underestimated and, especially, the smaller forms are missing.

The rocky shores sampled on either side of the isthmus differ in physiography and algal coverage; they

were sampled for ecological representation, not for

comparative faunistic studies, so the samples give

a relatively poor picture of the total polychaete

fauna of the area. Nevertheless, a comparison between the fauna of the Atlantic and Pacific coastlines of Panama is instructive and is given below.

SAMPLING OF THE ROCKY SHORES.—The Atlantic

site at Galeta Island reef flats has a dead coral substrate and offers five different zones. The Pacific

site at Paitilla Beach has an andesite rock substrate

and has several zones, of which only the lower and

middle intertidal zones were sampled. Each sample

covered a relatively small area but was quantitative, and sampling was repeated over a two-year

period. An area of i/8 m2 was sampled on each occasion in each zone, and a serious effort was made

to sort out all macro-invertebrates, defined for this

purpose as all animals more than 1. mm in one

dimension.

The Atlantic samples were first subjected to

live sorting, in which algal tufts were shaken over

a tray of sea water. The samples were then refrigerated in water for several hours or overnight. This

refrigeration brought out many of the polychaetes

that were burrowed in the coral substrate. Finally

the coral fragments were broken up into one-inch

fragments, and this rubble was sorted through for

the remainder of the animals.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

In the field, the Pacific samples were preserved

in 10 percent formalin and sorted later in the laboratory under a dissecting microscope. The large

barnacles often contained polychaetes; these were

obtained by scraping the outer and inner surfaces

of the tests so that the parietal canals of the barnacles were opened.

The samples are referred to in the systematic section without specific reference to the collector. The

different zones and the collecting dates for the

rocky intertidal samples are summarized below.

Acanthophora Zone

20 Oct 70

22 Oct 70

3 May 73

8 Aug 73

Coralline Zone

8Sep70

2 Oct 70

9 Oct 70

3 Mar 71

8 Aug 71

13 Jun 72

14 Mar 73

Balanus Zone

29 Jan 71

ATLANTIC OCEAN

Laurencia Zone

1 Sep70

8 Oct 70

23 Oct 70

26 Oct 70

27 Oct 70

1 Feb 71

17 Feb 71

15 Jun 71

18 Jun 71

2 Oct 71

26 Oct 71

8 Feb 72

21 Mar 72

26 Mar 72

27 Mar 72

17 Apr 72

20Jul72

26 Mar 73

PACIFIC OCEAN

Hydroid Zone

28 Oct 70

29 Oct 70

26 Apr 71

26 Aug 71

Mangrove Zone

23 Apr 71

Thalassia Zone

7Sep70

29 Sep 70

3OSep7O

2 Oct 70

7 Apr 71

26 Jul 71

2 Oct 71

7 Jun 72

Zoanthus Zone

28 Oct 70

30 Oct 70

Tetraclita Zone

29 Jan 71

31 Jan 71

4 Jan 72

ACKNOWLEDGMENTS.—I would like to thank Dr.

Amada A. Reimer, Pennsylvania State University,

for placing this interesting material at my disposal,

and Dr. Ira RubinofF, Smithsonian Tropical Research Institute, Balboa, Canal Zone, for financial

and other support during this study. The material

was collected under Contract No. 14-12-874 from

the Environmental Protection Agency to the Smithsonian Tropical Research Institute. Dr. J. David

George at the British Museum (Natural History)

(BMNH) and Dr. J0rgen Kirkegaard of the Zoological Museum, Copenhagen (ZMC), loaned me the

material previously identified by Monro. I am very

grateful to both for their help. Most of the writing

of this paper was done at the Santa Catalina Marine Biological Laboratory, and I thank Dr. Russel

L. Zimmer for making laboratory space and other

facilities available to me. Mrs. Elaine Jahn did the

typing and helped put the whole paper together,

for which I am very grateful.

Type specimens deposited in the National Museum of Natural History, Smithsonian Institution,

carry the catalog number of the former United

States National Museum (USNM). Type specimens

in the Hancock collections carry the designation

"Poly."

Family APHRODITIDAE

Aphrodita diplops, new species

FIGURE Ib-f

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (2, holotype, USNM 53084; paratype, USNM

53085).

DESCRIPTION.—The type is a ripe female, 25 mm

long and 5 mm wide with setae, with 36 setigers.

The body is cylindrical and abruptly tapering at

both ends. The ventrum is densely studded with

small papillae. The thin dorsal felt is dull graycolored.

The prostomium (Figure \e) is more wide than

long with two very prominent ocular peduncles;

the facial tubercle is large. Each ocular peduncle

has a pair of large eyes and a single smaller eye.

The larger eyes are directed dorsally and laterally,

and the small eyespots are dorsal. The ceratophore

of the median antenna is densely studded with short

papillae; the antennal style is about twice as long

as the prostomium.

The first parapodia are directed forwards and are

biacicular with small fascicles of setae. The posterior

faces of both noto- and neuropodia are studded with

small papillae. The dorsal and ventral cirri are of

the same length and project forwards about as far

as the tip of the median antenna. The palpi are

about four times as long as the prostomium; they

are evenly tapering and are covered with four rows

of long, very thin cilia.

In all other parapodia, the notopodia are low,

flattened projections; the neuropodia are conical

and have cylindrical bases. The smooth elytrae are

kidney-shaped and oriented in a long axis along the
NUMBER 221

FIGURE 1.—Unopherus canariensis Langerhans: a, anterior

end, dorsal view, X 25. Aphrodita diplops, new species: b,

neurosetae, third setiger, X 160; c, notoseta, median setiger,

X 160; d, neuroseta, median setiger, X 160; e, anterior end,

dorsal view, X 25; /, notoseta, third setiger, X 385.

body of the specimens. A few dark .pigment spots

are present near the dorsal margin of each elytron;

otherwise they are whitish.

Except for the first three pairs, each notopodium

has 13 to 15 light golden or brassy setae arranged in

a nearly closed crescent, with the opening to the

crescent facing anteriorly; the median setae are

longer than the others and project toward the middorsum. All notosetae penetrate the felt, but do not

project from it. Superior and inferior notosetae are

smooth. Median notosetae (Figure 1c) in each

fascicle have two rows of asperities; there is a 90°

angle between the two rows with the asperities

alternating in the two rows. Each asperity is shaped

like a squared scale.

Each median and posterior neuropodium has

four setae; each seta (Figure Id) is distally falcate

and has a slender subterminal tooth; this tooth

may be worn so that the seta appears spurred

rather than bidentate.

Setae in the first three setigers resemble those

found further back, but differ in proportions; thus

each neuroseta (Figure 1 b) has a long, gently curved

tip and a small spur situated considerably more

basally than in the normal neurosetae. Except in

the first setiger, the notosetae (Figure If) have

double rows of triangular scales arranged at 90°

angles to each other and alternating as in the

normal notosetae. The notosetae of the first setiger

are smooth, gently tapering capillaries.

Aphrodita diplops resembles A. armifera Moore

(1910:371-375, pi. 31: figs. 65-66, pi. 32: figs. 67-75),

A. falcifera Hartman (1939a:23-24, pi. 1: figs. 11-15,

pi. 26: figs. 319-320), and A. roulei Horst (1917:

261-262, 3 unnumbered figs.) in that all four species

have spurred or bidentate neurosetae and asperities

on the notosetae.

The median antenna in A. armifera and A.

falcifera is less than one-half the length of the

prostomium; in A. diplops, it is twice the length of

the prostomium. The eyes are small eyespots in

both A. armifera and A. falcifera; the eyes in A.

diptops are large and cover a large part of the

large ocular peduncles.

Aphrodita roulei resembles A. diplops closely,

but differs in that ocular peduncles and, apparently, eyes are absent in the former and prominently present in the latter.

Aphrodita diplops has been found in sand in one

locality at Galeta Reef, Panama, in the western

Atlantic Ocean.

Aphrodita japonica Marenzeller, 1879

Aphrodita japonica Marenzeller, 1879:111. — Hartman,

1939a:21-22, pi. 1: figs. 1-5; 1968:21-23, 5 figs, [unnumbered].

Aphrodita solitaria.—Monro, 1933a:12 [not Essenberg, 1917 =

A. refulgida Moore, 1910].

MATERIAL EXAMINED.—St. Elmo Bay, Perlas

Islands, Panama, dredging and trawling on the

east side of the bay, 6-9 fm, sand and shell (1),

coll. Mortensen.

REMARKS.—The specimen examined is the one

reported as A. solitaria by Monro. As indicated by
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hartman (1939a:23), this specimen is to be referred

to A. japonic a rather than to A. refulgida, to which

A. solitaria Essenberg has been referred.

Aphrodita japouica is known from widely scattered areas in the north and east parts of the

Pacific Ocean.

Family POLYNOIDAE

Key to the Species in the Present Collection

1. Prostomium with 2 antennae; body strongly flattened, completely covered by the chestnutcolored elytrae Iphione ovata

Prostomium with 3 antennae; body and elytrae otherwise 2

2. Lateral antennae inserted ventrally; prostomial peaks present 3

Lateral antennae inserted distally, prostomial peaks absent 4

3. Elytrae smooth with a black region posteriorly Harmothoe balboensis

Elytrae strongly fringed, separated into polygonal field surmounted by large, forked

spines Harmothoe hirsute

4. With 12 pairs of elytrae 5

With more than 12 pairs of elytrae 10

5. Elytrophores with branchiae Chaetacanthus magnificus

Elytrophores smooth 6

6. Notopodia with smooth lancet-shaped setae in addition to setae with transverse rows of

serrations 7

All notosetae tapering smoothly with transverse rows of serrations 8

7. Antennae and dorsal cirri pilose Thormora johnstoni

Antennae and dorsal cirri smooth Thormora taeniata

8. Notosetae much finer than neurosetae, elytrae with all papillae of the same size 9

Notosetae resemble neurosetae in thickness, anterior elytrae with large spinose papillae in

addition to the smaller papillae found on all elytrae Lepidonotus nesophilus

9. Elytrae densely fringed, with marmorated color pattern Lepidonotus humilis

Elytrae sparsely fringed, with alveolated color pattern Lepidonotus crosslandi

10. Eighteen pairs of elytrae 11

Numerous pairs of elytrae 12

11. Elytrae pale gray; except for the first, all elytrae with small, pointed spines only

Halosydna glabra

Elytrae black, except for one small pale ring; all elytrae with large pustulate tubercles in

addition to the small spines Halosydna leucohyba

12. Neurosetae distally cleft Lepidasthema varius

Neurosetae distally entire, with small subtenninal teeth Lepidasthema gigas

Chaetacanthus magnificus (Grube, 1875)

Iphione magnified Grube, 1875:51.

Chaetacanthus magnificus (Grube).—Monro, 1928b:558.—

Hartman, 1939a:28-29.

MATERIAL EXAMINED.—Coiba Island, dredging

off convict settlement in 5-10 fm, (1); Gorgona

Island, dredging close to shore in 15 fm, shell, dead

coral and gravel (1); both coll. Crossland.

REMARKS.—The species is here accepted as defined by the above-mentioned authors. The present

material is under revision by Dr. Marian H.

Pettibone of the Smithsonian Institution.

Chaetacanthus magnificus is considered widely

distributed in warm waters.

Halosydna glabra Hartman, 1939

Halosydna glabra Hartman, 1939a:35-36, pi. 4: figs. 43-50.

Halosydna reticulate—Monro, 1928b:563-565 [not Johnson,

1897 = H. johnsoni Darboux, 1899:246, as Lepidonotus

johnsoni, new name for Polynoe reticulata Johnson].

MATERIAL EXAMINED.—Taboga Island, from

floats of the stage at the end of the hotel pier (27),

coll. Crossland.

REMARKS.—Hartman (1939a: 34-35) indicated

that she thought these specimens belonged to H.

johnsoni or might represent more than one species.

The present material from Taboga all belongs to

one species, H. glabra, and fits very well with this

species, originally described in part on Panamanian

material.
NUMBER 221

Halosydna glabra is known from warm water

areas in the eastern Pacific Ocean from Panama to

Gulf of California, Mexico.

Halosydna leucohyba (Schmarda, 1861)

Polynoe leucohyba Schmarda, 1861:153-154, pi. 36: fig. 308.

Halosydna leucohyba (Schmarda).—Hartman, 1944a:9.

Halosydna fuscomarmorala.—Monro, 1928b:566-567 [not

Grube, 1875].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (9); Colon, coral flat near Lim6n Bay (4), coll.

Crossland.

REMARKS.—The present specimens fit very well

with material collected in other parts of the western

tropical Atlantic Ocean and with the descriptions

of Polynoe pustulata Mclntosh (1855) and P. granulata Ehlers (1887), both of which were considered

synonymous with H. leucohyba (Schmarda) by

Hartman (1944a).

Halosydna leucohyba appears to be one of the

most common intertidal scaleworms in the tropical

western Atlantic Ocean; it has not been found on

the Pacific side of the Isthmus.

Harmothoe balboensis Monro, 1928

Harmothoe balboensis Monro, 1928b:560-561, figs. 9-11.

MATERIAL EXAMINED.—Balboa, Panama, rocks

and rock-pools (1, holotype, BMNH), coll. Crossland.

REMARKS.—The holotype and only known specimen of this species is as described by Monro.

Harmothoe hirsuta Johnson, 1897

Harmothoe hirsuta Johnson, 1897:182-183, pi. 6: figs. 27-29,

pi. 7: fig. 38, pi. 8: figs. 53A-C.—Monro, 1928b:558-559,

fig. 8.—Hartman, 1968:77-78, 6 figs, [unnumbered].

MATERIAL EXAMINED.—Balboa, rocks and rockpools, low tide (2); Gorgona Island, coral (4);

Taboga Island, from floats at the stage at the end

of the hotel pier (1); all coll. Crossland.

REMARKS.—Monro indicated that his specimens

had simple, pointed spines (Monro, 1928b, fig. 8). A

reexamination of his specimens shows that the

elytral spines are covered with small spinelets; and,

especially on smaller specimens, the spines may be

multifid or at least bifid. On larger specimens the

spinelets are small in relation to the main spine,

all of which may thus appear as a single spine in

low magnification.

Harmothoe hirsuta is known from the eastern

Pacific Ocean between southern California and

Panama.

Harmothoe species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1); Balboa, low tide (1); coll. Crossland.

REMARKS.—The present specimens are both incomplete and lack elytrae. They have been assigned

to the genus Harmothoe because they resemble

members of this genus in setal structures and in the

structure of the prostomium.

Harmothoe lunulata var. pacifica Monro (1928b:

559-560) reported from Taboga is not a polynoid

but belongs to the family Hesionidae (see Gyptis

species indeterminate, p. 16). Specimens reported

by Monro (1933a) from the Galapagos Islands have

not been reexamined.

Iphione ovata Kinberg, 1855

Iphione ovata Kinberg, 1855:383.—Monro, 1928b:557-558.—

Hartman, 1939a:27, pi. 3: figs. 31-32.

MATERIAL EXAMINED.—Taboga, shore (4, 2 in

BMNH, 2 in ZMC); Tortolla, 3-5 fm, shell bottom,

14 Dec 1915 (1); Taboguilla, under rocks at low

tide, 11 Nov 1915 (3), all coll. Mortensen. Coiba

Island, volcanic rocks and boulders (2); Gorgona

Island, coral (3), both coll. Crossland.

REMARKS.—All specimens reported above were

identified by Monro and reported in 1928b. They

fit very well with the specimens reported from other

areas of the eastern Pacific Ocean by Hartman

(1939a).

Iphione ovata is known from the eastern Pacific

Ocean in tropical waters.

Lepidasthenia gigas (Johnson, 1897)

Polynoe gigas Johnson, 1897:172-175, pi. 7: figs. 33, 42. 42a,

pi. 8: figs. 48, 48a, 48b, 49.

Lepidasthenia gigas (Johnson).—Hartman, 1968:113-114, 5 figs.

[unnumbered].

Lepidametria virens.—Monro, 1928b:562-563 [not Blanchard,

1849].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Zone (1). Balboa, rocks and rock-pools at low tide

(31); Coiba Island, dead coral and sand (4); coll.

Crossland.

REMARKS.—Lepidasthenia gigas usually has dark,

often greenish blotches on the elytrae, and the

general body color is light yellowish. The superior

neurosetae are as thick as, or thicker than, the

inferior ones. The inferior neurosetae usually have

distinct subterminal teeth, but these teeth may be

worn and visible only in a few setae or as small

protuberances on the side of the setae.

Setal structures in the specimens reported as

Lepidametria virens by Monro (1928b) agree with

those of Lepidasthenia gigas rather than with those

of L. virens, as the latter are characterized by

Hartman (1939a:46-47, pi. 8: figs. 105-110).

Lepidasthenia gigas has been reported as a commensal of terebellid polychaetes from southern

California and the Galapagos Islands.

Lepidasthenia varius Treadwell, 1917

Lepidasthenia varius Treadwell, 1917a:259-260, pi. I: figs.

11-16.—Hartman, 1956:271.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone(l).

REMARKS.—Lepidasthenia varius is characterized

by the deeply incised neurosetae in all setigers.

Lepidasthenia varius is known from the Galapagos Islands and from various areas in the tropical

western Atlantic Ocean.

Lepidonotus crosslandi Monro, 1928

Lepidonotus crosslandi Monro, 1928b:553—555, figs. 1—4.—

Hartman. 1939a:42-43, pi. 5: figs. 63-69.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1); Tetraclita Zone (4). Balboa, rocks and

rock-pools (1); Taboga, 5 fm, coral fragments (3,

syntypes, BMNH); coll. Crossland.

REMARKS.—There is little to add to the original

description. The neurosetae are distinctly unidentate in median setigers; the tips are often thicker

than illustrated by Monro (1928b), perhaps due to

wear. The nearly smooth elytrae have poorly developed fringes, and the low tubercles are of two

size-classes as indicated by Hartman (1939a:43). The

syntypic material of Monro has been reexamined.

Lepidonotus crosslandi has been found in shallow

water in Panama and off Peru.

Lepidonotus hutnilis Augener, 1922

Lepidonotus humilis Augener, 1922:40; 1933:194-195.—Hartman, 1944a:9-10.

Lepidonotus carinulatus.—Monro, 1928b:553 [not Grube,

1878].

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (2); Galeta Reef, Laurencia Zone (1). Balboa,

scrapings from buoy at Canal entrance (2); Taboga,

5 fm (19); Taboga, from dead and broken coral,

1-2 fm, and from the hotel pier (8, reported as

Thormora johnstoni by Monro, 1928b, reidentified

as L. humilis by Dr. Marian H. Pettibone (pers.

comm., Smithsonian Institution); coll. Crossland.

REMARKS.—Lepidonotus humilis is a small intertidal species. The marmorated elytrae have a

strongly developed fringe. The neurosetae are bifid,

even if the secondary tooth sometimes is lost; it is,

however, usually visible in at least some setae in

each neuropodium. Notosetae are fine and nearly

hair-like.

Lepidonotus humilis has been reported from

intertidal areas on both sides of the Isthmus of

Panama and is more widely distributed in the

Caribbean Sea.

Lepidonotus nesophilus Chamberlin, 1919

Lepidonotus nesophilus Chamberlin, 1919:75-78, pi. 4: figs.

1-7. pi. 5: fig. 13.—Hartman, 1939a:38-39. pi. 7: figs. 83-95.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (5). Taboga, from dead coral, 1-2 fm, or from

the hotel pier (1, reported by Monro, 1928b as

Thormora johnstoni; reidentified by Dr. Marian

H. Pettibone, Smithsonian Institution), coll.

Crossland.

REMARKS.—Lepidonotus nesophilus has large

spinose macrotubercles on anterior elytrae and

small, sharply pointed microtubercles on all elytrae.

The elytral fringe consists of very short papillae.

Lepidonotus nesophilus is known from the eastern Pacific Ocean in warm waters.

Thormora johnstoni (Kinberg, 1855)

Lepidonotus johnstoni Kinberg, 1855:384.

Thormora johnstoni (Kinberg).—Monro, 1928b:556-557, figs.

5-7 [in part].—Hartman, 1939a:5O-51, pi. 7: figs. 96-98;

1968:139-140, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Taboga, 4-5 fm, sand and
NUMBER 221

stones, 8 Feb 1916 (1); Taboguilla, approximately

5 fm, shell bottom, 5 Jan 1916 (1), both coll. Mortensen. Taboga, from dead and broken coral, 1-2

fm or from the hotel pier (13), coll. Crossland.

REMARKS.—Thormora johnstoni is well known

from warm waters in the eastern Pacific Ocean. It

is easily identifiable as one of the two local lepidonotins with two different kinds of notosetae:

smooth lancet-shaped ones and ones with transverse

rows of serrations. It differs from the closely similar

T. taeniata, which has similar kinds of setae, in

that the antennae and tentacular cirri are smooth;

these structures are hirsute in T. taeniata. Crossland's material from Taboga has been reexamined

by Dr. Pettibone.

Thormora taeniata (Ehlers, 1887)

Polynoe taeniata Ehlers, 1887:51-52, pi. 10: figs. 1-8.

Thormora taeniata (Ehlers).—Seidler, 1924:92-94.

MATERIAL EXAMINED.—Taboga, from dead and

broken coral and from the hotel pier (39, identified

as T. johnstoni by Monro, 1928b, reidentified by

Dr. Marian H. Pettibone, Smithsonian Institution);

Taboguilla, 5 fm, shell (2).

REMARKS.—The differences between this and the

closely related T. johnstoni have been mentioned

above. This material has been reexamined by

Marian H. Pettibone and will be the subject of a

future publication by Dr. Pettibone.

Thormora taeniata has previously been reported

from the western Atlantic Ocean.

Family POLYODONTIDAE

Key to the Species from Panama

1. Prostomium with large paired ocular peduncles 2

Prostomium without ocular peduncles (eyes small) Eupanthalis perlae, new spedes

2. Superior neurosetae hastate Polyodontes oculea

Superior neurosetae pendllate S

3. Superior neurosetae with distal tuft Panthalis pacifica

Superior neurosetae with long, slender tip extending beyond the pendllate portion

Panthalis morUnuvi

Eupanthalis perlae, new species

FIGURE 2

Eupanthalis kinbergi.—Monro, 1928b:568 [not Mclntosh,

1876].

MATERIAL EXAMINED.—San Jos6 Island, Perlas

Islands, 25 fm, mud and shells (1 holotype, ZMC),

coll. Mortensen.

DESCRIPTION.—The type and only specimen is an

anterior fragment that is 18 mm long and 6 mm

wide without setae, with 46 setigers. The body is

thick and color patterns are absent.

The prostomium (Figure 2b) consists of two

hemispherical lobes separated by a deep median

fissure. Two pairs of eyes are present; one is

just posterior to the widest part of the head

and the second is near the posterior margin of

the head. A pair of frontal antennae is on the

anterior margin; each is long and slender and

about one-half times longer than the length of the

prostomium. Posterior to the prostomium proper

is a thick, folded nuchal organ, which is anteriorly

produced into a low ceratophore fused to the

dorsal surface of the prostomium. The median antenna is attached to this ceratophore; it starts at

the level of the anterior pair of eyes. Posteriorly the

nuchal organ has a small, medial transverse fold.

The palps are about three times as long as the

length of the prostomium; each is thick and is

distally covered with long, slender papillae in a

scattered arrangement. A pair of smooth, tentacular cirri are ventroposterior to the palps; each is

about the same length as the palps; setae are absent in this tentacular segment.

All parapodia are bluntly truncate and have

long, subulate ventral drri; dorsal cirri are similar

in size and shape.

Setae are of several different kinds. In the anterior end of the body, anterior to setiger 17, a

series of short, slender setae with two or three

whorls of short, slender teeth is found in the anterior fascicles (Figure 2/); each seta is subdistally

inflated. The posterior fascicles in these anterior
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Eupanthalis perlae, new species: a, seta from posterior fascicle, setiger 12, X 385;

b, anterior end, dorsal view, X 25; c, superiormost seta, setiger 40, X 385; d, inferiormost

seta, setiger 40, X 385; e, median seta, setiger 40, X 385; f, seta from anterior fascicle, setiger

12, X 385.

setigers have long, slender, gently curved setae

(Figure 2a) with a fine marginal border of thin

hairs. Setae are of four kinds in posterior setigers;

thus in setiger 40 two slender, pilose setae are

found superiormost (Figure 2c) resembling those

found in anterior setigers; below them is a single

straight, finely pilose seta with a smooth tip; then

there are fifteen to sixteen thick, distally recurved

setae (Figure 2e) with a subdistal double row of

spines and a distal tuft of spines; these setae are in

two rows. These are presumably homologous to

the kind equipped with aristae in other species of

the genus, but aristae are absent in all parapodia in

the present specimen. Inferiormost in each fascicle

is a series of gently curved or straight, pilose setae

(Figure 2d); the innermost hairs on these setae may

be very coarse, approaching spines.

Eupanthalis perlae closely resembles other species in the genus, but differs in the position of the

median antenna, which is attached between the

anterior eyes in the present species and at the

posterior margin of the prostomium in all other

species in the genus. Eupanthalis nigromaculata

Grube (1878) has large eyes covering most of the

prostomial lobes, and the palps are smooth. Eupanthalis kinbergi Mclntosh (1876) lacks the whorled

setae in anterior setigers and the position of the

median antenna is different.

The shape of the aristate setae with their double

groups of spines appears unique to E. perlae.

Eupanthalis perlae is described from one locality

on the Pacific side of the Isthmus of Panama.

Panthalis mortenseni (Monro, 1928)

Polyodontes mortenseni Monro, 1928b:569-572, figs. 19-24.

Panthalis mortenseni (Monro).—Hartman, 1939a:87; 1968:143-

144, 3 figs, [unnumbered].

MATERIAL EXAMINED.—Molones, 3-5 fm, 15 Dec

1915, coll. Mortensen (1). Taboga, muddy sandy

beach at low tide, 5 Feb 1916, coll. Mortensen (2,

holotype, BMNH; other specimen in ZMC).
NUMBER 221

REMARKS.—All three specimens reported by

Monro (19286) have been reviewed. They are as

described by Monro. The pencillate setae with

smooth tips are most easily distinguished in parapodia posterior to setiger 30.

Panthalis mortenseni is known from southern

California to Pacific Panama.

Panthalis pacifica Treadwell, 1914

Panthalis pacifica Treadwell, 1914:184-186, pi. 11: figs. 1-7.—

Hartman, 1968:145-146, 4 figs, [unnumbered].

Panthalis jogasimae.— Monro, 1928b:568-569 [not Izuka, 1912].

MATERIAL EXAMINED.—Gorgona Island, 20-30

fm, dredge, fine sand and shell (2), coll. Crossland.

REMARKS.—The specimens are as described by

Monro (1928b); the setal structures correspond to

those of P. pacifica rather than to P. jogasimae.

Panthalis pacifica is known from southern California to Panama in muddy bottoms.

Polyodontes oculea (Treadwell, 1901)

Panthalis oculea Treadwell, 1901:188-189, figs. 14-18.

Polyodontes oculea (Treadwell).—Monro, 1928b:572-575, figs.

25-30— Hartman, 1939a:83-84, pi. 24: figs. 294-299.

MATERIAL EXAMINED.—Taboga, mud and sand

(2); Balboa rock and rock-pools (6), both coll. Mortensen. Taboga, dredging, 1-2 fm, (3); Taboga,

about 10 fm, mud (2); Colon, trawling in Limon

Bay, 5 fm, thin mud (3); all coll. Crossland. Additionally 27 specimens from Port of Spain Roadstead, Trinidad, 4-5 fm, thin mud, were reexamined.

REMARKS.—The identification of the present

specimens with Treadwell's species agrees with the

reinterpretation made by Monro and Hartman.

Polyodontes oculea is known from the Caribbean Sea and from warm water areas in the eastern

Pacific Ocean as far north as Baja California.

Family S1GALIONIDAE

Key to the Species from Panama

1. Prostomium quadrate with a small median antenna Thalenessa lewisii

Prostomium subglobular with large median ceratophore and antenna 2

2. Ceratophore with ctenidia, parapodia sparsely papillose; elytrae without sand incrustations ....

Sthenelais fusca

Ceratophore without ctenidia; parapodia densely papillose, elytrae sand-incrusted

Psammolyce spinosa

Psammolyce spinosa Hartman, 1939

Psammolyce spinosa Hartman, 1939a:72-74, pi. 19: figs. 232-

243; 1968:159-160, 6 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (3); Zoanthus Zone (3).

REMARKS.—Psammolyce spinosa was described

from warm water areas in the eastern Pacific Ocean

and has been reported from as far north as California. The present specimens represent the first

record from the Atlantic Ocean. The specimens

have been directly compared with the California

material; no relevant differences were noted.

Sthenelais fusca Johnson, 1897

Sthenelais fusca Johnson, 1897:185-186, pi. 9: figs. 60. 61,

61a,b; pi. 10: figs. 64, 64a-g.—Monro, 1933a: 16.—Hartman,

1939a:61-62, pi. 13: figs. 153-162; 1968:163-164. 5 figs,

[unnumbered].

Sthenelais variabilis var. colorata.—Monro. 1933a: 14-16, fig. 7

[not Monro, 1924].—Hartman, 1939a:63-64, pi. 13: figs. 163-

166.

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (2); Coiba Island, in sand at low

water, spring tide, fine sea grass and dead coral

bedded in the sand (2); coll. Crossland.

REMARKS.—All these specimens belong to the

same species. Presently, the rather variable S. fusca

cannot be separated into subgroups. Some of the

morphological variations indicated by Hartman

(1939a), however, probably relate to different species.

Sthenelais fusca is known from the eastern Pacific Ocean from Washington to Panama.

Thalenessa lewisii (Berkeley and Berkeley, 1939)

Sigalion lewisii Berkeley and Berkeley, 1939:326-328, figs. 2, 3.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Eusigalion hancocki Hartman, 1939a:59-60, pi. 12: figs. 141-

145, 148-152.

MATERIAL EXAMINED.—Taboguilla, sandy shore

at low tide (1), coll. Mortensen.

REMARKS.—This specimen was not identified and

reported on by Monro (1928b) in his study of the

polychaetes collected by Mortensen.

Thalenessa lewisii is restricted to the warmer

parts of the eastern Pacific Ocean.

Family CHRYSOPETALIDAE

Key to the Species from Panama

1. Body long and slender, caruncle absent 2

Body short, caruncle present S

2. Lancet-shaped notosetae present ventral to the paleae; paleae narrow Bhawania goodei

Lancet-shaped notosetae absent; paleae distally expanded Bhawania rh/eti

3. Paleae of 2 abruptly difiPerent kinds in each fascicle Paleanotus chrysolepis

All paleae similar in width or grading evenly from one end to the other in each fascicle

Chrysopetahtm occidental*

Bhawania goodei Webster, 1884

Bhawania goodei Webster. 1884:308-309, pi. 7: figs. 1-15.—

Monro, 1933a: 18-19.—Day, 1967:118-119, fig. 21a-f.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1); Paitilla Beach, Hydroid Zone (1). Taboga

Island, low water spring tide (2), coll. Crossland.

REMARKS.—Bhawania goodei can be separated

from B. riveti (Gravier), which has also been found

in the Panamanian material, by the presence of

lancet-shaped notosetae ventral to the paleae. The

paleae are usually rather narrow and have a speckled dirty gold color in B. goodei; they are distally

expanded and a polished gold color in B. riveti.

Bhawania goodei is very widespread in warm

water. Day (1967:119) indicated that the species

may be circumtropical.

Bhawania riveti (Gravier, 1908)

Chrysopetalum riveti Gravier, 1908:40; 1909:638-641, pi. 17:

figs. 31-34.

Bhawania riveti (Gravier).—Monro, 1933a: 17-18.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (2); Tetraclita Zone (4). Taboga Island, shore,

collected in beach sandstone at low water, spring

tide (1), coll. Crossland.

REMARKS.—The differences between this species

and B. goodei have been listed above. The present

specimens fit very well with the description given

by Gravier (1909).

Bhawania riveti was originally described from

Peru and has been reported from the Galapagos

Islands (Treadwell, 1928:466), in addition to the

Panamanian records (Monro, 1933a: 17-18).

Chrysopetalum occidentale Johnson, 1897

Chrysopetalum occidentale Johnson, 1897:161-162, pi. 5: figs.

15, 16, pi. 6: figs. 17-19.—Monro, 1933a: 19.—Hartman,

1968:185-186, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Gorgona Island (1), coll.

Crossland.

REMARKS.—According to Monro, the median antenna should be situated between the two pairs of

eyes in the present specimen; in fact it is between

the anterior eyes as usual in this species. The prostomium is large and globular, and the nuchal organ is nearly as large and of the same general shape.

Chrysopetalum occidentale is known from southern California to Panama in shallow water.

Paleanotus chrysolepis Schmarda, 1861

Paleanotus chrysolepis Schmarda, 1861:163, pi. 37: figs. 326-

329.—Monro, 1933a: 19.

MATERIAL EXAMINED.—Taboga Island from the

float at the end of the stage of the hotel pier, coll.

Crossland (fragments).

REMARKS.—The present fragments are unidentifiable to species at the present stage of preservation.

They do belong to the genus Paleanotus. In view

of the close similarities between P. chrysolepis and

P. bellis (Johnson, 1897), to treat these fragments
NUMBER 221 11

as unidentifiable might be considered preferable.

Monro considered the two species synonymous.

Both species remain poorly known; I decided to

leave the present fragments with the current name

until the taxonomic problem has been satisfactorily

clarified.

Paleanotus chrysolepis is possibly widespread in

warm waters.

Family AMPHINOMIDAE

Key to the Species from Panama

1. Body short and depressed 2

Body long and cylindrical S

2. Dorsura with a single, reddish brown median stripe Chloeia entypa

Dorsum with 3 dark longitudinal stripes Chloeia viridis

3. Caruncle reduced or absent, extending maximally through part of the first segment 4

Caruncle large, often with complex lateral folds 6

4. Seven pairs of branchiae, antennae articulated Linopherus canatiensis

Numerous pairs of branchiae, antennae smooth 5

5. Eyes large, branchiae present on all but the first and last few segments ...linopherus oculata

Eyes indistinct; branchiae limited to the first two-thirds of the body Linopherus ambigua

6. Caruncle long and narrow, median fold covering the 2 lateral folds Eurythoe complanata

Caruncle either very wide, or with very well-developed lateral folds visible from above 7

7. Caruncle diamond-shaped with indistinct lateral folds Hertnodice carunculata

Caruncle with very distinct lateral folds, usually spindle-shaped 8

8. Caruncle with wide lateral folds attached basally to the high median ridge

Notopygot ornata

Caruncle with lateral folds in the form of 7 or 8 leaves attached laterally to the high

median ridge Pherecardia striata

Chloeia entypa Chamberlin, 1919

Chloeia entypa Chamberlin, 1919:30-31, pi. 13: figs. 8, 9, pi.

14: figs. 1, 2.—Hartman, 1968:191-192, 5 figs, [unnumbered].

Chloeia pinnata.—Monro, 1933a:7-8, fig. 3 [not Moore, 1911].

MATERIAL EXAMINED.—Gorgona Island, 30 fm,

muddy sand, showing shell fragments after screening with a fine mesh (3), coll. Crossland.

REMARKS.—Chloeia entypa differs from C. pinnata Moore in that the former has a broad, reddish

brown middorsal stripe, which is absent in the

latter. Chloeia pinnata is usually salmon-colored

without any distinct color patterns. The posterior

notosetae are distally serrated in both species;

these serrations point basally in C. entypa and

distally in C. pinnata.

Chloeia entypa is widely distributed in warm

waters in the eastern Pacific Ocean, but it has frequently been confused with C. pinnata so the exact

distribution is difficult to ascertain at the present

time.

Chloeia viridis Schmarda, 1861

Chloeia viridis Schmarda, 1861:144-146, pi. 25: figs. 295-305.—

Monro, 1933a:9-10, fig. 4.—Hartman, 1940a:205.

MATERIAL EXAMINED.—Taboga Island, between

Taboga and Taboguilla, 6-12 fm, soft mud (1),

coll. Crossland.

REMARKS.—Chloeia viridis has a characteristic

pattern of three longitudinal stripes on the dorsum,

usually most distinct in the anterior end. This pattern is still distinct on the specimen reexamined

above.

Chloeia viridis is known from both sides of the

Isthmus of Panama (Hartman, 1944a: 15).

Eurythoe complanata (Pallas, 1776)

Aphrodita complanata Pallas, 1776:109.

Eurythoe complanata (Pallas).—Chamberlin, 1919:28.—Monro,

1933a:4-5—Hartman, 1968:195-1%, 4 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (6); Zoanthus Zone (1). Balboa, pools at lowest

tide-level (1); Taboga Island, from colony of Pocillopora, all in anterior regeneration (5); Gorgona

Island, low water, spring tide, from tubes of Sabellaria (6); Gorgona Island, from coral (2); Coiba

Island, 5 fm. (1); Colon, from coral flat at SW corner of Limon Bay (3); St. Elmo Bay, Per las Island,

shore at low tide (5); all coll. Crossland.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—Eurythoe complanata as presently accepted has a circumtropical geographical dispersal

pattern in the rocky intertidal and in shallow subtidal areas, frequently associated with pockets of

sand under rocks and in coral reefs. It is easily separable from other Panamanian amphinomids by the

long flexuose caruncle and the long, slender body

shape. Hermodice carunculata is as long-bodied, but

the caruncle of this species is wide and nearly

diamond-shaped.

Hermodice carunculata (Pallas, 1776)

Aphrodita carunculata Pallas, 1776:102.

Hermodice carunculata (Pallas).—Fauvel, 1923:131-132, fig.

47a-i— Hartman, 1951:22-25, pi. 5: fig. 1.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1).

REMARKS.—The present specimen is smaller than

specimens usually reported, but otherwise it is very

similar to the species as illustrated by Hartman

(1951).

Hermodice carunculata is found in warm water

areas of the Atlantic Ocean.

Linopherus ambigua (Monro, 1933),

new combination

Eurythoe ambigua Monro, 1933a:6-7, fig. 2.

Pseudeurythoe ambigua (Monro).—Fauchald, 1972:38-40.

MATERIAL EXAMINED.—Taboga Island, bay between Taboga and Taboguilla, 6-12 fm, soft mud

(1, holotype, BMNH), coll. Crossland.

REMARKS.—The specimen is as described by

Monro (1933a). It is known through the original

record from Panama and from North Carolina

(Day, 1973:16). The generic name Pseudeurythoe

is invalid, as indicated below (L. canariensis); thus

the new combination.

Linopherus canariensis Langerhans, 1881

FIGURE la

Linopherus canariensis Langerhans, 1881:109-110, pi. 4: figs.

14a-g.

Pseudeurythoe canariensis (Langerhans).—Fauchald, 1972:38-

41.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1).

REMARKS.—The present specimen (Figure la) fits

very well with L. canariensis as originally described,

except that the first pair of eyes is slightly larger

and half-moon-shaped rather than round as originally described. Branchiae are present from setiger

3; six pairs are present rather than seven as originally described. A caruncle is completely absent.

The posterior antennae and the dorsal cirri of the

first setiger are biarticulated. None of the notopodial setae are subdistally inflated.

The valid genotype for this genus is canariensis

Langerhans, 1881. The generic name Linopherus

is available in this combination and has priority

over Pseudeurythoe Fauvel (1932), which was proposed for the same group of species.

Linopherus canariensis has been reported only

once, from the Canary Islands; the present record is

from the Atlantic side of Panama.

Linopherus oculata (Treadwell, 1941),

new combination

Eurythoe oculata Treadwell, 1941:18, figs. 1-3.

Pseudeurythoe oculata (Treadwell).—Hartman, 1956:274.—

Fauchald, 1972:38-40.

Eurythoe dubia.— Monro, 1933a:5-6, fig. 1 [not Horst, 1912].

MATERIAL EXAMINED.—Balboa, swimming along

the surface alongside the ship in Balboa Dock (1),

coll. Crossland.

REMARKS.—Eurythoe dubia sensu Horst (1912)

is distinctly a species of Eurythoe in that it has a

large caruncle (cf. Fauchald, 1972:38). The specimen reported by Monro (1933a:5-6) belongs to the

genus Linopherus and agrees in all details with L.

oculata as redescribed by Hartman (1956). The generic name Pseudeurythoe is invalid, as indicated

above (L. canariensis); thus the new combination.

Linopherus oculata has been reported twice from

Balboa, Panama; no further records are known.

The large eyes are possibly related to the sexual

maturation. Treadwell's specimen was sexually mature, and Crossland caught the present specimen

swimming at the surface.

Notopygos ornata Grube, 1856

Notopygos ornata Grube, 1856:53.—Monro, 1933a: 10-11, fig.

5.—Hartman, 1940, p. 207.

MATERIAL EXAMINED.—Balboa, pools at the low-
NUMBER 221

est tide-level (9); Taboga Island, from colony of

Pocillopora (5); Gorgona Island, from coral (4);

St. Elmo Bay, Perlas Islands, from shore at low

tide (1); all coll. Crossland.

REMARKS.—Notopygos ornata has a long, spindleshaped caruncle with widely expanded lateral folds

attached basally to the median ridge. The color

pattern of dark bluish patches on a pink background is very characteristic in life (Monro, 1933a,

fig. 5b) but fades very rapidly in alcohol.

Notopygos ornata is known from the western Atlantic Ocean in warm waters as well as from similar

areas in the eastern Pacific Ocean.

Pherecardia striata (Kinberg, 1857)

Hermodice striata Kinberg, 1857:13.

Pherecardia striata (Kinberg).—Monro, 1933a:7.—Hartman,

1940a:207; 1951:25, pi. 5: fig. 2.

MATERIAL EXAMINED.—Coiba Island, low water,

spring tide (5); Taboga Island, from colony of

Pocillopora (2); Gorgona Island, from coral (6);

all coll. Crossland.

REMARKS.—The shape of the caruncle with the

seven or eight large lateral leaves attached to the

narrow median ridge and the color pattern consisting of numerous thin longitudinal pin-stripes are

both characteristic of this species in the area.

Pherecardia striata is known from nearly circumtropical areas.

Family PHYLLODOCIDAE

Key to the Species from Panama

1. Ventral tentacular cirrus of segment 2 folicaceous; others cirriform 2

All tentacular cirri cirriform 3

2. All tentacular segments dorsally complete; dorsal cirri distally bluntly rounded

Steggoa lobocephalica

First tentacular segment dorsally reduced; dorsal cirri distally pointed Sige orientalis ?

3. Five antennae present, nuchal papilla absent 4

Four antennae present, nuchal papilla present 5

4. Dorsum with transverse dark pigmented bands Eumida bijoliata

Dorsum with three longitudinal dark stripes Eulalia myriacyclum

5. Proboscis with proximal papillae in scattered arrangement; dorsal cirri kidney-shaped

Phyllodoce panamensis

Proboscis with proximal papillae in longitudinal rows; dorsal cirri longer than wide 6

6. Dorsal cirri dark red (even in preserved material) Anaitides erythrophyllus

Dorsal cirri colorless or greenish to tan-colored 7

7. Body colorless or white with colorless dorsal cirri Anaitides madeirensis

Body tan colored with tan-colored or greenish dorsal cirri Anaitides lamellifera

Anaitides erythrophyllus (Schmarda, 1861)

Lepadorhynchus erythrophyllus Schmarda, 1861:88, pi. 39:

fig. 232.

Anaitides erythrophyllus (Schmarda).—Hartman, 1951:33.

Phyllodoce oculata Ehlers, 1887:135-140, pi. 40: figs. 4-6.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1).

REMARKS.—The present specimen agrees well

with A. erythrophyllus as described by Ehlers

(1887). The dorsal cirri are rather distinctly pointed,

and the eyes are very large in the prostomium. The

identification cannot be considered certain, since

the separation of the many species of Anaitides

must be considered doubtful at the present time.

Anaitides erythrophyllus is apparently limited to

the warmer parts of the western Atlantic Ocean.

Anaitides lamellifera (Pallas, 1788)

Nereis lamellifera Pallas, 1788:232.

Phyllodoce (Anaitides) lamellifera (Pallas).—Monro, 1933a:22-

24, fig. 10.

MATERIAL EXAMINED.—Taboga Island, from float

at the stage at the end of the hotel pier (7); ? Coiba

Island, dredging off the convict settlement in 5 fm,

smooth bottom with branched Lithothamnion (3);

all coll. Crossland.

REMARKS.—As indicated by Monro, this species
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

is difficult to separate from A. madeirensis. The

specimens agree with the species as described and

illustrated by Monro. The lower edge of the dorsal

cirri is evenly rounded. The specimens from Coiba

Island were not reported by Monro; they are small,

and in relatively poor condition. The identification

is not considered certain.

Anaitides lamellifera is apparently widespread in

warm waters; it may be circumtropical.

Anaitides madeirensis (Langerhans, 1880)

Phyllodoce (Anaitis) madeirensis Langerhans, 1880:307-308,

fig. 44a,b.—Monro, 1933a:21-22.

Anaitides madeirensis {Langerhans).—Hartman, 1968:231-232,

3 figs, [unnumbered].

MATERIAL EXAMINED.—Gorgona Island, from

coral (15); Taboga Island from floats at the end of

the stage of the hotel pier (3); Taboga Island, from

broken branches of coral (1); Balboa, scraped off

buoy brought in from the canal entrance (1); all

coll. Crossland.

REMARKS.—Anaitides madeirensis is here accepted

as having pointed dorsal cirri and the ventral cirri

projecting beyond the tip of the acicular lobe.

There are no setae in the third tentacular segment.

The proboscis has 12 rows of papillae, and there

should be less than 12 papillae in each row. The

specimens are always pale when preserved and

apparently have little color in life. Monro (1933a:

22) remarked that he could not find any differences,

except for the presence of setae in the third tentacular segment, between this species and A. medipapillata Moore; an additional difference lies in the

fact that the latter species is usually very dark

purplish brown in life and when freshly preserved.

Anaitides madeirensis is considered widely distributed in warm waters.

Anaitides species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (3).

REMARKS.—The present specimens are incomplete posteriorly, and most of the tentacular cirri

and dorsal cirri have been lost. The eyes are relatively small and the dorsal cirri are distally bluntly

rounded.

Eteone species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (3). Coiba Island, dredging off the convict

settlement in 5 fm, smooth bottom with branched

Lithothamnion (1), coll. Crossland.

REMARKS.—One of the specimens has been dried

out; the others are either too incomplete or poorly

preserved to allow further identification. The specimen from Coiba Island was not reported by

Monro (1933a).

Eulalia myriacyclum (Schmarda, 1861)

Notophyllum myriacyclum Schmarda, 1861:87, pi. 29: fig. 233.

Eulalia myriacyclum (Schmarda).—Hartman, 1944a: 16.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (3).

REMARKS.—Eulalia myriacyclum has a yellow or

orange base color with dark brown or black longitudinal stripes covering the whole body. It has yet to

be reported from the eastern Pacific Ocean.

Eulalia species indeterminate

Eulalia species.—Monro, 1933a:21.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (3); Paitilla Beach, Tetraclita Zone (11). Taboga Island, from floats at the stage at the end of

the hotel pier (2), coll. Crossland.

REMARKS.—The present specimens fit rather well

with E. viridis as suggested by Monro, but differ in

color patterns and in that the enlarged terminal

tooth on the setal shafts is considerably smaller than

in that species. The specimens do not seem to belong to any species reported from the tropical western Atlantic nor from the eastern Pacific.

Eumida bifoliata (Moore, 1909)

FIGURE 3a

Eulalia (Sige) bifoliata Moore, 1909b:349-350, pi. 16: figs.

31-34.

Eumida bifoliata (Moore).—Hartman, 1968:271-272, 2 figs.

[unnumbered].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1).

REMARKS.—The present specimen agrees with E.

bifoliata as this species was originally described.
NUMBER 221 15

They share the same color pattern and the presence

of two dark crescents posterior to the paired eyes on

the prostomium.

The structure of the parapodia is different; the

present specimen is posteriorly incomplete with

about 25 setigers. The neuropodial acicular lobes

(Figure 3a) are distally equally bifid rather than

unequal as described, and the ventral cirrus is distally rounded rather than pointed. These differences may be related to the size of the specimen

or to the incompleteness of the fragment.

Eumida bifoliata is known from central and

southern California in shallow water. The present

record is from the eastern Pacific side of the Isthmus of Panama.

Phyllodoce panamensis Treadwell, 1917

Phyllodoce panamensis Treadwell, 1917b:428-430, figs. 1, 2.

Phyllodoce (Anaitides) panamensis Treadwell.—Monro,

1933a:24-25, fig. 11.

Anaitides panamensis (Treadwell).—Hartman, 1956:260.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (14); Paitilla Beach, Tetraclita Zone, partially

in the parietal canals in dead Tetraclita (3). St.

Elmo Bay, Perlas Islands, east side of the bay, 6-9

fm, shell and sand (3), coll. Crossland.

REMARKS.—Phyllodoce panamensis is a true Phyllodoce in that the papillae on the proximal part

of the proboscis are in a scattered arrangement

rather than in rows as in Anaitides. The species

FIGURE 3.—Eumida bifoliata (Moore): a, parapodium 22, posterior view, X 95. Syllis gracilis

Grube: b, median seta, median setiger, X 770. Opisthosyllis brunnea Langerhans: c, median

seta, median setiger, X 950. Typosyllis caeca (Monro): d, simple seta, posterior setiger, X 700.

Typosyllis proliferat (Krohn): e, median seta, median setiger, X 950. Autolytus anoplos Monro:

/, anterior margin of pharynx, X 160; g, two teeth from pharynx, X 385; h, median parapodium, anterior view, X 95; i, median seta, median setiger, X 950; ;', anterior end, dorsal

view, X 50; k, simple seta, posterior setiger, X 950.
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

superfically resembles members of the genus Notophyllum in that the large, kidney-shaped dorsal

cirri are imbricated on the back. Treadwell's,

Monro's and Hartman's descriptions of this species

do not make clear that only one species is involved

in the present concept; however, all material cited

above belongs to one and the same species.

Phyllodoce panamensis is known exclusively from

the Pacific coast of Panama.

Sige orientalis? Imajima and Hartman, 1964

Sige macroceros orientalis Imajima and Hartman, 1964:70, pi.

14c-f.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimen agrees with

Sige in that it has the strongly foliose, asymmetrical

ventral tentacular cirrus. It resembles S. orientalis

in that it lacks setae in the second tentacular segment. Dorsal cirri are only present in far posterior

segments so the identification is considered doubtful since the shape of the dorsal cirri is known to

vary along the body.

Sige orientalis is known only from intertidal

areas in northern Japan.

Steggoa lobocephalica (Kinberg, 1866)

Eulalia lobocephalica Kinberg, 1866b:241.—Monro, 1933a:2O,

fig. 9 [not E. lobocephala Schmarda, 1861].

Steggoa lobocephalica (Kinberg).—Hartman, 1948:48, pi. 7:

figs. 2-3.

MATERIAL EXAMINED.—Taboga Island, from

branches of dead coral (12), coll. Crossland.

REMARKS.—These specimens are as described by

Monro and Hartman. The dorsal cirri are basally

truncate, so that the total shape of the dorsal cirri

is narrowly triangular; they are held erect over the

body in the best preserved specimens.

Steggoa lobocephalica is known from the eastern

Pacific Ocean from Chile and Peru to Panama in

shallow water.

Family HESIONIDAE

Key to the Species from Panama

1. Two antennae; palps absent 2

Three antennae; palps present 3

2. Dorsum light tan with brown longitudinal stripes Hesione intertexta

Dorsum dark brownish with light transverse stripes Hesione picta

3. Both parapodial rami well developed; 8 pairs of tentacular cirri Gyptis sp. indet.

Notopodia reduced; 6 pairs of tentacular cirri 4

4. Digitifonn postsetal neuropodial lobe project beyond the other parapodial lobes in median

setigers Ophiodronuu obscurus

All parapodial lobes of the same length Ophiodronuu pugettensis

Gyptis species indeterminate

Harmothoe lunulata var. pacifica Monro, 1928b:559-560 [in

part].

MATERIAL EXAMINED.—Taboga Island, 4-5 fm,

sand and stone (1), coll. Mortensen.

REMARKS.—This specimen is an unidentifiable

species of Gyptis. The specimen reported by Monro

from the Galapagos Islands can be assumed to belong to a species of Harmothoe.

Hesione intertexta Grube, 1878

Hesione interlexta Grube, 1878:102-103, pi. 6: fig. 5.—Monro,

1933a:26.

MATERIAL EXAMINED.—Balboa, pools at the lowest tide levels (1), coll. Crossland.

REMARKS.— Even in preserved specimens, H. intertexta has distinct brown longitudinal lines on

the pale dorsum. It is known from the Pacific and

Indian Oceans in warm waters.

Hesione picta Muller, 1858

Hesione picta F. Muller, 1858:213-214, pi. 6: fig. 3.—Hartman, 1951:35.—Jones, 1962:180.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (2); Galeta Reef, Coralline Zone (1).

REMARKS.—The complex synonymy of this species
NUMBER 221 17

has been worked out and is cited in the works

indicated above. The species is best characterized

by its color pattern; even in preserved specimens,

the pattern is dark with series of light transverse

stripes scattered irregularly along the length of the

body.

Hesione picta is known from Florida to Brazil in

the western Atlantic Ocean.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1).

REMARKS.—Ophiodromus pugettensis can be separated from the rather similar east coast species of

the genus as indicated above. It is previously known

from Washington to western Mexico in the eastern

Pacific Ocean, but this record of the species is the

first from typical tropical waters.

Ophiodromus obscurus (Verrill, 1873)

Podarke obscura Verrill, 1873:589-590, pi. 12: fig. 61.—

Pettibone, 1963:104-105, fig. 28a,b.

Podarke near guanica.—Hartman, 1951:36, pi. 10: figs. 1-3.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (2).

REMARKS.—Ophiodromus obscurus resembles O.

pugettensis closely, but can be separated from it on

the presence of a digitiform postsetal lobe in median setigers in O. obscurus; this lobe projects

clearly beyond all other parapodial lobes. All parapodial lobes are of the same length in O. pugettensis.

Ophiodromus obscurus is known from the western

Atlantic Ocean from New England to the Caribbean region and Gulf of Mexico.

Ophiodromus pugettensis (Johnson, 1901)

Podarke pugettensis Johnson, 1901:397-398, pi. 3: figs. 23-35.

Ophiodromus pugettensis (Johnson).—Hartman, 1968:369-370,

3 figs, [unnumbered].

Family PILARGIIDAE

Synelmis albini (Langerhans, 1881)

Ancistrosyllis Albini Langerhans, 1881:107.

Synelmis albini (Langerhans).—Pettibone, 1966:191-195, figs.

19-21.

Ancistrosyllis gorgonensis Monro, 1933a:26-18, fig. 12.

MATERIAL EXAMINED.—Gorgona Island, dredged

close to shore, 15 fm, shell, dead coral and gravel

(1, holotype, A. gorgonensis, BMNH); Gorgona

Island, from coral (1); both coll. Crossland, det.

Monro.

REMARKS.—The present specimens agree with

5. albini as this species was defined by Pettibone

(1966) who first suggested the synonymy of A. gorgonensis with S. albini. Considerable variation

appears in the species, and a complete revision of

large materials from worldwide areas should be

undertaken in order to elucidate the variability.

As presently defined, S. albini is circumtropical.

Family SYLLIDAE

Key to the Species from Panama

1. Ventral cirri absent 2

Ventral cirri present 4

2. Body yellow with dark purplish brown markings Autolytus anoplos

Body pale yellow or pinkish with no distinct color markings 3

3. Distal tooth longer than the subdistal tooth in the composite setae Autolytus cf. tnagntu

Distal tooth distinctly shorter than the subdistal one in the composite setae

! Autolytus sp. indet.

4. Dorsal cirri distinctly articulated 5

Dorsal cirri smooth or irregularly wrinkled 16

5. Body dorsoventrally flattened; pharynx with a trepan

Trypanosyllis (Trypanedenta) taeniaformis

Body cylindrical; pharynx with a single tooth 6

6. Pharynx with a middorsal tooth in posterior position Opisthosyllis brunnea

Pharynx with a middorsal tooth in anterior positions 7

7. All setae thick and simple HoplosyUis spongicola

At least some setae composite 8
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

8. Simple and composite setae mixed in most segments Syllis gracilis

Simple setae present singly in posterior parapodia only 9

9. Composite setae include both spinigers and falcigers 10

All composite setae falcigers H

10. Large composite setae unidentate; median antenna posterior to the eyes

Langerhansia mexicana

Large composite setae bidentate; median antenna between the eyes Langerhansia cornuta

11. Dorsal cirri with less than 20 articles 12

Dorsal cirri with more than 20 articles IS

12. Dorsum with transverse broad pigment band on every few segments TyposylUs hyalina

Two narrow dark lines across each segment dorsally TyposylUs actculata

13. Dorsum with solid color, without distinct markings .....14

Dorsum with pattern of transverse dark lines 15

14. Dorsum white, eyes absent TyposylUs caeca

Dorsum brownish purple, eyes present TyposylUs proUfera}

15. Enlarged appendages on some composite setae in median setigers; 2 regular transverse lines

across dorsum of each segment TyposyUis fuscosuturata

All composite setae with appendages of similar size; each segment with a series of irregular

transverse markings on the dorsum TyposylUs variegata

16. Prostomium covered posteriorly by a nuchal flap 17

Prostomium without a nuchal flap Pionosyllis sp. indet.

17. Pharynx with 14 teeth OdontosylUs sp. indet.

Pharynx with 6 teeth OdontosylUs polycera

Autolytus anoplos Monro, 1933

FIGURE 3/-A

Autolytus anoplos Monro, 1933a:38-39, fig. 18.

MATERAL EXAMINED.—Galeta Reef, Coralline

Zone (18); Galeta Reef, Laurencia Zone (2). Lim6n

Bay, from wreck covered with sponges, Balanus, etc.

(7, syntypes, BMNH), coll. Crossland.

REMARKS.—A somewhat expanded description of

the species is given below based on the new material

recorded above. The main differences between this

material and the species as described by Monro

are in details in the color pattern and in the

somewhat different length of the anterior appendages.

A complete specimen with 125 setigers is 25 mm

long and 0.5 mm wide. It is cream-colored with

a single dark brown bar at the posterior margin

of each segment. All dorsal cirri are dark brown

except those in the first three setigers, which are

cream-colored. Two pairs of lensed eyes are present.

The body is ventrally flattened and evenly wide

with abruptly tapering anterior and posterior ends.

The prostomium (Figure 3/) is twice as wide as

long and has two pairs of eyes at the posterolateral

half. The two short lateral antennae are slender;

the median antenna is more than twice as long as

the lateral ones and considerably stouter. All

antennae are smooth. The peristomium is a little

longer than the prostomium and has a pair of

slender, relatively short peristomial cirri. The palps

are as long as the prostomium and are free to the

base. They project ventrally. The first dorsal cirri

are very long and terete; each is about as long as

the median antenna, resembling the latter also in

structure and shape. The dorsal cirri of the second

setiger are considerably shorter and more slender,

but are longer and stouter than those in the more

posterior setigers.

All parapodia posterior to setiger 3 resemble

each other. Each has a bluntly conical acicular

lobe (Figure 3/i) and a poorly developed, low

presetal lobe; postsetal lobes are absent. Ventral

cirri are absent; each dorsal cirrus is short and

fusiform.

Composite setae (Figure Si) with short, deeply

bifid appendages are present in all setigers; the

basal part of the appendages is somewhat slenderer

than the distal part. The shaft is strongly dentate.

Each posterior setiger also contains a single bayonetseta (Figure Sk); each has a thickened shaft, which

is distally strongly dentate, and a mucron, which

is slender and finely tapering.

The proboscis has a trepan (Figure 3/) with 25

teeth. The teeth alternate regularly between two

sizes; both kinds have a median ridge and narrowed

wings. The one kind is about one-fourth smaller

than the other (Figure 3g).

The genus Autolytus was reviewed for the
NUMBER 221 19

Japanese fauna by Imajima (1966a) who subdivided

the genus into two, based on the distribution and

numbers of teeth in the trepan. The subgenus

Regulatus should have teeth in groups of nine or

multiples of nine; the subgenus Autolytus sensu

stricto should have all other numbers of teeth. The

teeth in Regulatus should alternate regularly

between large and small; all teeth should be of

approximately the same size in Autolytus sensu

stricto.

The present species has 25 teeth in the trepan,

thus not a multiple of nine; but the teeth alternate

regularly between small and large. The species is,

thus, intermediate between the two subgenera

proposed by Imajima. The subdivision appears

useful, and I do not suggest here that it should

be rejected. The genus Autolytus sensu lato is

large, and anything that can be used to subdivide

it must be considered a help at the present stage of

taxonomic knowledge. The scheme of subgenera in

this genus may have to be extended to include

other characters in addition to the structure of

the trepan.

Autolytus anoplos resembles A. magnus Berkeley

(see below) in the structure of the parapodia and

the composite setae; it differs from the latter in that

it has only 25 teeth in the trepan; A. magnus has

nearly twice that number. The bayonet setae have

a very short dentate region in A. anoplos and a

somewhat longer spinose region in A. magnus.

Other similar species include A. irregularis and A.

spinoculatus both described by Imajima (1966a)

from Japanese waters. Autolytus anoplos differs

from both of these in the color pattern and in the

number and structure of the teeth on the trepan.

Autolytus anoplos is known from two areas near

Colon, Panama.

Autolytus cf. magnus Berkeley, 1923

Autolytus magnus Berkeley, 1923:210, pi. 1: figs. 5-4.—Imajima, 1966a:40-46, figs. 9a-f. lOa-f. lla-d.

MATERIAL EXAMINED.—Paitilla Beach, Tetraclita

Zone (4).

REMARKS.—The present four specimens fit very

well with A. magnus as redescribed by Imajima

(1966a) in the numbers of the teeth in the trepan

and the shape of the teeth. The structure of the

composite setae is also similar. The present specimens, however, have considerably shorter nuchal

epaulettes, and the tentacular cirri are considerably

shorter than shown by Imajima (1966a, fig. 9a). All

present specimens are atokous.

Autolytus magnus is known from Japan and

from western Canada; it has not been reported from

California.

Autolytus species indeterminate

Autolytus sp.— Monro, 1933a:39-41, fig. 17.

MATERIAL EXAMINED.—Balboa, from buoy

brought in from the canal entrance (1); Coiba

Island, dredging off the convict settlement in 5

fm, smooth bottom with branching Lithothamnion

(1); both coll. Crossland.

REMARKS.—Monro gave a rather detailed description of the specimen from Balboa. Nothing can

presently be added to the description since the

specimen is in a rather bad state of preservation.

The identity cannot be further clarified. The specimen from Coiba Island is incomplete and badly

preserved.

Haplosyllis spongicola (Grube, 1855)

Syllis spongicola Grube, 1855:104-105.

Haplosyllis spongicola (Grube).—Imajima, 1966c:220-221, fig.

38a-h.—Hartman, 1968:433-434. 4 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (2); Paitilla Beach, Tetraclita Zone (2).

REMARKS.—The present specimens have simple,

distally bifid setae with a subdistal boss; the body

is relatively stout and the antennae and tentacular

cirri are strongly beaded.

Haplosyllis spongicola is considered more or less

cosmopolitan in warmer waters; it is usually associated with sponges (Monro, 1933a:34; Day, 1973:29).

Langerhansia cornuta (Rathke, 1843)

Syllis comuta Rathke, 1843:164.

Syllis (Ehlersia) cornuta Rathke.—Fauvel, 1923:267-268, fig.

100g-i.—Monro, 1933a:34.

Langerhansia cornuta (Rathke).—Imajima. 1966d:256-259, fig.

51.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (4). Taboga Island, from under the floating

stage of the hotel pier (1) and from broken branches

of dead coral, 1-2 fm (5); Gorgona Island, 15 fm,
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

shell, gravel and dead coral (26); all coll. Crossland.

REMARKS.—The present specimens fit very well

with L. cornuta as described by Fauvel (1923) and

Imajima (1966d). The species is distinguished from

the similar L. mexicana below. Langerhansia

cornuta is known from world-wide areas.

Langerhansia mexicana (Rioja, 1961),

new combination

Ehlersia mexicana Rioja, 1961:291-295, figs. 4-11.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—Langerhansia mexicana can be separated from the closely similar L. cornuta on the

position of the median antenna, which is posterior

to the eyes in the former and between the posterior

eyes in the latter. Rioja (1961:295) indicated thai

the larger composite setae should be bifid in L.

cornuta and unidentate in L. mexicana; this also

is the case in the present specimen.

Langerhansia mexicana is known from the east

coast of Mexico and possibly from Florida. Rioja

(1961) indicated that he thought the material

reported as L. cornuta by Hartman (1951) from

Florida might belong to this species. The present

record is from the Atlantic side of Panama.

described in the genus. Setae are distally bidentate.

The specimen cannot be further described since it

is rather badly preserved.

Opisthosyllis brunnea Langerhans, 1879

FIGURE 3C

Opisthosyllis brunnea Langerhans, 1879:541-543, pi. 31: fig.

7.—Imajima, 1966c:230-233, fig. 42a-n.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (3); Galeta Reef, Laurencia Zone (73); Galeta

Reef, Thalassia Zone (11); Paitilla Beach, Balanus

Zone (5); Paitilla Beach, Hydroid Zone (21); Paitilla

Beach, Tetraclita Zone (7).

REMARKS.—The present specimen agrees with O.

brunnea as originally described and as reviewed by

Imajima (1966c). The only difference noted is that

the serrations along the cutting edges of the

composite setae (Figure 3c) appear to be somewhat

finer than indicated by Imajima (1966c, fig. 42e-h).

This difference is not considered critical.

Opisthosyllis brunnea was described from

Madeira in intertidal areas and has since been

reported from one locality in Japan. The species

appears to be one of the most common syllids in

all intertidal areas in Panama, both on the Atlantic

and Pacific sides of the Isthmus.

Odontosyllis polycera (Schmarda, 1861)

Syllis polycera Schmarda, 1861:72, pi. 28: fig. 219.

Odontosyllis polycera (Schmarda).—Monro. 1933a:36-37, fig.

15.

MATERIAL EXAMINED.—Balboa, low tide at

Panama (19), coll. Crossland.

REMARKS.—Odontosyllis polycera was redescribed

by Monro (1933a) based on the specimens listed

above. They are as described. The species is known

from warm temperate and tropical waters in the

Atlantic and Pacific Oceans.

Odontosyllis species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1).

REMARKS.—The present specimen agrees with

Odontosyllis in that it has short, smooth dorsal

cirri and series of recurved pharyngeal teeth. The

number of teeth, 14, is higher than in any species

Pionosyllis species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1).

REMARKS.—The present specimen is incomplete.

The proboscis is everted. The setae include composite smooth-edged spinigers and composite falcigers with distally expanded, blade-like appendages.

It resembles P. uraja Imajima (1966b: 114-116, fig.

37a-g) from Japan, but is too poorly preserved to

be completely identified.

Syllis gracilis Grube, 1840

FIGURE 36

Syllis gracilis Grube, 1840:77.—Fauvel, 1923:259, fig. 96f-i.—

Monro, 1933a:30.—Imajima, 1966c:248-250, fig. 49a-k.—

Hartman, 1968:463-464, 4 figs, [unnumbered].

Syllis longissima.—Monro, 1933a:30 [not Gravier, 1900].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (2); Paitilla Beach, Tetraclita Zone (15).
NUMBER 221 21

Taboga Island, from broken branches of dead coral

1-2 fm (36), from under the floating pier of the

hotel pier (3), and obtained by breaking up beach

sandstones (2); Gorgona Island, shell, gravel, and

dead coral, 15 fm (3); Balboa Docks, off piles and

large sponges growing in the shade of the quays (12)

and from pile scrapings (1); Colon (1); all coll.

Crossland.

REMARKS.—Syllis gracilis is very widespread in

warm waters. The present specimens differ in no

manner from the species as described by the authors

cited above. Syllis longissima Gravier has unidentate

compound setae; the material reported as 5.

longissima from Balboa by Monro (1933a:30) have

bidentate setae. Syllis gracilis has short dorsal cirri

with 8-10 articles in median setigers. Syllis longissima has alternating long and slightly shorter dorsal

cirri with 8-10 and 12-15 articles, respectively. The

large setae in median setigers are characteristic

(Figure 36).

All of the above records, except the one from

Colon, are from the Pacific side of the Isthmus of

Panama. This fact is probably more due to the

kinds of environment sampled on the two sides than

to a distributional difference between the two

oceans.

Trypanosyllis (Trypanedenta) taeniaeformis

(Haswell, 1886)

Syllis taeniaeformis Haswell, 1886:741.

Trypanosyllis taeniaeformis (Haswell).—Monro, 1933a:35-36.

Trypanosyllis (Trypanedenta) taeniaeformis (Haswell).—Iraajima, 1966c:239-241, fig. 45.

MATERIAL EXAMINED.—Gorgona Island, dredging

close to shore, 15 fm, gravel, shell, dead coral (27),

coll. Crossland.

REMARKS.—The present specimens agree with this

species as reviewed by Imajima (1966c).

Trypanosyllis taeniaformis is known from warm

water shallow areas in the Pacific and Indian

Oceans.

Typosyllis aciculata Treadwell, 1945

Typosyllis aciculata Treadwell, 1945:1-2, figs. 1-5.—Reish,

1950:1-5, 8 figs, [unnumbered].—Hartman. 1968:475-476, 7

figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (2); Paitilla Beach, Hydroid Zone (1); Paitilla

Beach, Tetraclita Zone (1). Coiba Island, dredging

off the convict settlement, 5 fm, smooth bottom with

branched Lithothamnion (1), coll. Crossland.

REMARKS.—Typosyllis aciculata is known from

southern and central California. The present records

come from both sides of the Isthmus of Panama.

The specimens were compared to material from

southern California and no differences could be

noted. Typosyllis aciculata is differentiated from

similar species in the area as indicated in the key.

Typosyllis caeca (Monro, 1933), new combination

FIGURE 3d

Syllis caeca Monro, 1933a:3O-32, fig. 13.

MATERIAL EXAMINED.—Taboga Island, from floats

at the stage at the end of the hotel pier (5, syntypes,

BMNH), coll. Crossland.

REMARKS.—This species belongs to Typosyllis in

that it has a single, simple bidentate seta (Figure

Sk) in each of a few of the posterior setigers; in most

specimens, they are most easily seen in setigers 7

or 8 from the posterior end. Typosyllis caeca

resembles T. variegata (Grube) in setal structures

as indicated by Monro (1933a:32) but differs clearly

in that T. caeca is white without any color pattern,

while T. variegata has series of transverse stripes.

Typosyllis caeca is known only from its original

record.

Typosyllis fuscosuturata (Augener, 1922)

Syllis (Typosyllis) fuscosuturata Augener, 1922:43-44; 1927:52.

Syllis fuscosuturata Augener.—Monro, 1933a:32-34, fig. 14.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Laurencia Zone (3).

Taboga Island, from broken branches of dead coral,

1-2 fm (1); Gorgona Island, 15 fm, shell, gravel and

dead coral (2) and from coral (1); all coll. Crossland.

REMARKS.—This species is characterized by the

enlarged hooks in median setigers. Other species

of the genus that have similar hooks include T.

excilis Gravier (1900) and T. maculata (Imajima,

1966d: 277-279, fig. 59). Typosyllis fuscosuturata

has two brown transverse lines on each segment.

Typosyllis maculata has no color markings on the

body proper but has dark dots on the dorsal cirri.

Typosyllis excilis lacks color patterns.
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Typosyllis fuscosuturata is known from the

Caribbean Sea and the west Indies in addition to

areas in the eastern Pacific Ocean in warm waters.

Typosyllis hyalina (Grube, 1863)

Syllis hyalina Grube, 1863:45-46, pi. 4: fig. 7.—Monro,

1953a:30.

Typosyllis hyalina (Grube).—Imajima, 1966d:271-273, fig. 57.

MATERIAL EXAMINED.—Gorgona Island, 15 fm,

shell, gravel and dead coral (1); Taboga Island,

from under the floating stage of the hotel pier (1);

both coll. Crossland.

REMARKS.—The present specimens fit very well

with the species as redescribed from Japan by

Imajima (1966d); it is known from worldwide

areas.

Typosyllis prolifera? (Krohn, 1852)

Syllis prolifera Krohn, 1852:66.

Syllis (Typosyllis) prolifera Krohn.—Fauvel, 1923:261-262, fig.

97a-g.

Typosyllis prolifera (Krohn).—Imajima, 1966d:292-294, fig.

651a-n.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimens agree with T.

prolifera? except that the appendage of the composite setae (Figure 3<?) is somewhat smaller than as

illustrated and the inferior tooth is smaller and

does not project at as great an angle from the axis

of the appendage.

Typosyllis prolifera has a reported circumtropical

distribution.

Typosyllis variegata (Grube, 1860)

Syllis variegata Grube, 1860:85-87, pi. 3: fig. 6.—Monro,

1933a:28-29.

Syllis (Typosyllis) variegata Grube.—Fauvel, 1923:262, fig.

97h-n.

Typosyllis variegata (Grube).—Imajima, 1966d:292— Hartman, 1968:495-496, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Coralline Zone (13);

Galeta Reef, Laurencia Zone (52); Paitilla Beach,

Hydroid Zone (1); Paitilla Beach, Tetraclita Zone

(5). Taboga Island, from floating stage at the end

of the hotel pier (2); Balboa Docks, scrapings from

piles under the shade of quays (2); Balboa, rocks

and rock-pools at low tide (2); all coll. Crossland.

REMARKS.—The subdistal tooth of the composite

setae is at a slightly greater angle than usual in this

species; otherwise the specimens fit very well with

the species as reported from other areas. The dorsal

color pattern consists of fine dark brown or purplish

bands on each segment.

Typosyllis variegata is cosmopolitan in algal zones

intertidally.

Syllidae, Unidentifiable Fragments

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (fragments and poorly preserved specimens);

Paitilla Beach, Hydroid Zone (8, all epitokes in

bad condition); Paitilla Beach, Tetraclita Zone

(posterior end).

Family NEREIDAE

Key to the Species from Panama

1. Eversible proboscis with soft papillae only Ceratocephala croulandi

Eversible proboscis without soft papillae, but with paragnaths at least on some areas 2

2. Paragnaths present on both rings of the proboscis 5

Paragnaths present either on the maxillary ring or the oral ring, but never on both 3

3. Paragnaths present on oral ring only Eunereis paitillae, new species

Paragnaths present on maxillary ring only 4

4. Bidentate notopodial falcigers in median and posterior setigers Ceratonereis mirabiUs

No bidentate falcigers in any notopodia Ceratonereis irritabilis

5. All paragnaths of the same shape, pointed and conical 6

At least 2 different kinds of paragnaths present 11

6. Median and posterior notopodia with homogomph falcigers 7

Homogomph falcigers absent 9
NUMBER 221 23

7. Appendages of homogomph falcigers bi- or trifid Nereis panamensis, new species

Appendages of homogomph falcigers unidentate 8

8. Appendage of homo- and heterogomph falcigers recurved Nereis riisei

Appendage of homo- and heterogomph falcigers ankylosed Nereis caUaona

9. Notopodial lobe similar in length to all other parapodial lobes in all setigers

Neanthes gale toe, new species

Notopodial lobes distinctly longer than all other lobes in posterior setigers 10

10. Neuropodial falcigers with ankylosed appendages; superior notopodial lobe strongly foliose

Neanthes sucdnea

Neuropodial falcigers with recurved appendages; superior notopodial lobes narrow in all

setigers Neanthes pseudmnoodti, new species

11. Transverse and conical paragnaths present Permereis anderssoni

Pectinate, transverse, and conical paragnaths present 12

12. Area vi with transverse paragnaths 13

Area vi with 1 or 2 rows of small paragnaths Platynereis dumerilii

IS. Dorsal cirrus in posterior parapodia attached distally on the superior notopodial lobe;

areas vu and vm with a single row of alternating small and large paragnaths

Pseudonereis gallapagensis

Dorsal cirrus in posterior parapodia attached subdistally on the superior notopodial lobes;

areas vu and vra with paragnaths in 2 or 3 rows Pseudonereis variegata

Ceratocephala crosslandi (Monro, 1933)

Chaunorhynchus crosslandi Monro, 1933a:46-49. fig. 20.

Ceratocephala crosslandi (Monro).—Hartman, 1952:15.

MATERIAL EXAMINED.—Gorgona Island, dredging

in 30 fm, muddy sand and shell-fragments (1,

holotype, BMNH), coll. Crossland.

REMARKS.—The type is as described and illustrated by Monro. It closely resembles the subspecies

C. crosslandi americana Hartman (1952) from

southern California; the two may not be separable.

Ceratocephala crosslandi is known from the

eastern Pacific Ocean from Panama to Washington.

Ceratonereis irritabilis (Webster, 1879)

Nereis irritabilis Webster, 1879:231-234, pi. 5: figs. 56-64,

pi. 6: figs. 65-69.

Ceratonereis irritabilis (Webster).—Hartman, 1945:20-21, pi.

3: figs. 7-9; 1951:48.

Ceratonereis versipedata.—Monro, 1933a:45-46, fig. 19 [not

Ehlers, 1887].

MATERIAL EXAMINED.—Col6n, from the outer

piles of the quays at the docks (6), coll. Crossland.

REMARKS.—These specimens agree with C.

irritabilis as this species was redescribed by Hartman

(1945). Day (1973:38-39) has retained the separation

between C. versipedata Ehlers (1887) and C. irritabilis. This separation may not be justifiable, but

under any circumstances the present specimens

belong to C. irritabilis as this species is presently

defined. If the two species should be synonymous,

Webster's name has priority.

Ceratonereis mirabilis Kinberg, 1866

FIGURE Aa-c

Ceratonereis mirabilis Kinberg, 1866a: 170.—Hartman, 1948:

71-72.

Ceratonereis lentaculata Kinberg, 1866a:170.—Hartman. 1940a:

218, pi. 35: fig. 47.—Monro, 1933a:45.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (2); Galeta Reef, Laurencia Zone (3); Galeta

Reef, Thalassia Zone (69). Col6n, from piles of the

quays (1); Gorgona Island, from coral (1); both

coll. Crossland.

REMARKS.—Ceratonereis mirabilis has usually a

distinctly bifid prostomium with very long antennae

and long, slender palps. The parapodia are short in

all setigers, but the dorsal cirri are enormously

prolonged in posterior setigers (Figure 4a).

Notopodia have slightly heterogomph falcigers

(Figure 46) in median and posterior setigers; each

appendage is slightly bifid and has a fringe of

long, slender hairs along the cutting edge. Neuropodial falcigers are similar, but are more distinctly

heterogomph (Figure 4c).

Ceratonereis mirabilis appears to be rather widespread in warm waters and may be circumtropical.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Ceratonereis mirabilis Kinberg: a, far posterior parapodium, anterior view, X 160;

b, notopodial falciger, posterior setiger, X 950; c, neuropodial falciger, posterior setiger, X 950.

Platynereis dumerilii (Audouin and Milne Edwards): d, notopodial homogomph falciger,

posterior setiger, X 950; e, neuropodial heterogomph falciger, posterior setiger, X 950; /, posterior parapodium, anterior view, X 160. Pseudonereis gallapagensis Kinberg: g, neuropodial

heterogomph falciger, posterior setiger, X 950; h, posterior parapodium, anterior view, X 95-
NUMBER 221 25

Eunereis paitillae, new species

FIGURE 5

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (2, holotype, USNM 53086; paratype, USNM

53087).

DESCRIPTION.—The holotype is an incomplete

specimen, 6 mm long and 1.2 mm wide without

setae, with 32 setigers. Brown pigment spots are on

the palpophores and laterally on the peristomium,

as well as on the dorsolateral anterior margins of

each of the first parapodia; otherwise the specimen

is light yellow in color. The paratype has been

dried out at one time.

The short prostomium (Figure 5e) is relatively

wide. The palpophores are short; the long palpostyles are slender. The slender antennae do not

reach beyond the tip of the palps.

The four pairs of tentacular cirri are slender;

the anterior dorsal ones are the shortest and barely

reach the posterior margin of the peristomium. The

anterior ventral ones reach the posterior margin of

the first setiger; the posterior dorsal ones reach the

middle of the second setiger, and the longer posFICURK 5.—Eunereis paitillae, new species: a, proboscis, ventral view, X 50; b, large supra-acicular neuropodial falciger,

median setiger, X 950; c, subacicular neuropodial falciger,

median setiger, X 950; d, posterior parapodium. anterior

view, X 160; e, anterior end, dorsal view, X 50.

terior ventral ones reach the posterior margin of the

fifth setiger.

The proboscis (Figure 5a, e)is fully everted in the

holotype; the jaws are rather slender and have

approximately eight teeth along the cutting edge.

The maxillary ring lacks paragnaths. Five paragnaths are on the oral ring; each of these is a

single, transversely elongated piece. One is in

area vi, and one each in areas vn and vm; other

paragnaths are absent.

All parapodia are similar, except that the

notopodia are strongly reduced and asetigerous in

the two first setigers. The notopodia (Figure 5c)

are otherwise distinctly trilobed with a very large

superior lobe, a short slender acicular lobe and a

digitate inferior lobe. The neuropodia have distinct,

but low, pre- and postsetal lobes, which are otherwise simple and bluntly conical. The slender,

digitate dorsal and ventral cirri are attached

basally.

Notopodial setae are all of one kind; each is a

long, slender homogomph spiniger with the cutting

edge of the appendage finely serrated. Neuropodial

setae are of two kinds: supra- and subacicular

fascicles are similar; both contain a number of

heterogomph spinigers and falcigers. One falciger

in each supra-acicular fascicle (Figure 5b) is

considerably larger than the others. It has a short

thick appendage with a strongly recurved, slender

tip and a group of five to seven hairs near the base

of the cutting edge. The other, slender falcigers

(Figure 5d) are similar, but the appendage is considerably less recurved at the tip and the hairs along

the cutting margin are found along the whole

margin.

The genus Eunereis is known for six species in

addition to E. paitillae described above; these six

include E. caeca Hartman (1960:93-94), E. longipes

Hartman (1936:479-480, fig. 53), E. longissima

(Johnston, 1840:178; see Fauvel, 1923:352, fig.

138a-d), E. marri Monro (1939:117, fig. lOa-f), E.

paradoxa 0rsted, 1843a: 177-178, pl. 4: fig. 50 pl. 5:

figs. 63, 64, 66), E. patagonica (Mclntosh, 1885:228-

229, pl. 35: figs. 13-15: pl. 17a: figs. 1-2) and E.

wailesi Berkeley and Berkeley (1954:456-457, figs.

1, 2). These species are rather varied in other

characters, but appear to have one feature in

common: they all lack paragnaths on the maxillary

ring and have at least a few on the oral ring.

Eunereis paitillae differs from all these in the
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

shape and number of paragnaths and in that it

lacks homogomph falcigers in the notopodia.

Neanthes galetae, new species

FIGURE 6a-c

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Laurencia Zone (6,

holotype, USNM 53088; paratypes, USNM 53089,

AHF Poly 1132); Galeta Reef, Thalassia Zone (1).

DESCRIPTION.—The holotype is a complete specimen 12 mm long and 1.5 mm wide without setae,

with 56 setigers. It is yellow with transverse brown

bars and small lateral brown patches on each of the

first 10 to 15 setigers.

FICURE 6.—Neanthes galetae, new species: a, anterior end, dorsal view, X 50; b, far posterior

parapodium. anterior view, X 160; c, neuropodial heterogomph falciger, posterior setiger,

X 950. Nereis panamensis, new species: d, neuropodial heterogomph falciger, posterior setiger,

X 950; e, notopodial homogomph falciger, posterior setiger, X 950; /, posterior parapodium,

anterior view, X 160; g, anterior end. dorsal view. X 50; h, proboscis, ventral view X 50;

i, proboscis, dorsal view. X 50.
NUMBER 221 27

The prostomium (Figure 6a) is longer than wide

and is rounded anteriorly; the antennae do not

project beyond the palps and are attached closely

together near the tip of the prostomium. The

palpophores are massive; the short palpostyles are

rounded. The four tentacular cirri are digitate;

the posterior ventral ones are the longest and reach

setiger three; the others do not reach beyond the

peristomium.

The proboscis (seen in dissection) has paragnaths

on most areas: area i has 2 in tandem, area n has

20 in 2 straight rows, area m has 18 in a rounded

patch, area iv has 25 in a rectangular patch, area

v lacks paragnaths, area vi has 8 in a round

patch, and areas vii-vm have 5 large paragnaths

in a single row.

Parapodia are similar along the length of the

lx>dy, except that the notopodia are slightly prolonged in posterior setigers. Each notopodium

(Figure 66), except the first two which are reduced,

has a short, blunt acicular lobe and a flattened

supra-acicular lobe; the subacicular lobe is thick

and digitate and of the same length as the supraacicular lobe. Each dorsal cirrus, especially in

posterior setigers, is very long and slender; all

dorsal cirri are attached basally. The neuropodia

are bifid; the superior part with the acicula and

setae has a very well-developed, bluntly rounded

postsetal lobe; the presetal lobe is a low fold. The

inferior neuropodial lobe is as long as the superior

one; it is digitate. The ventral cirrus projects as

far as the tip of the neuropodium in all setigers.

Notosetae are all of one kind; each is a long,

slender, homogomph spiniger; the cutting edge of

the appendage is slightly serrated. Neurosetae are

of three kinds. In anterior setigers, a few homogomph spinigers are in the superior fascicle; these

are replaced by heterogomph spinigers in more

posterior setigers. Otherwise, heterogomph falcigers

are found in both superior and inferior fascicles.

Each falciger (Figure 6c) has a short, slender appendage with five or six fine hairs along the cutting

edge; the distal part of the appendage is strongly

recurved.

The genus Neanthes was separated into groups

by Fauchald (1972:69-70, 408-410). According to

this system, N. galetae belongs to group IIB 1 in

that falcigers are present and in that it has the dorsal lobe of the notopodia barely longer than other

lobes in the parapodium. Other species in this

group with few paragnaths on areas vn and vm

include N. agulhana (Day, 1963:406-407, fig. 6d-j),

N. dawydovi (Fauvel, 1937:297-299, fig. la-k),' N.

kerguelensis (Mclntosh, 1885:225-227, pi. 35: figs.

10-12, pi. 16A: figs. 17-18), and N. kerguelensis oligodonta (Augener, 1913:164-166). Neanthes diversicolor (O. F. Muller, 1776), originally listed for this

group, was moved to the genus Hediste by

Hartmann-Schroder (1971:196, as a subgenus); this

change is followed here.

Of these species, N. agulhana, N. dawydovi, and

N. kerguelensis oligodonta lack paragnaths on both

areas i and v; the other species have paragnaths on

i but lack them on v. Neanthes galetae differs from

N. kerguelensis in that the dorsal cirri are twice as

long as the parapodial lobes in the former and no

longer than these lobes in the latter; otherwise the

two species are rather similar.

Neanthes psendonoodti, new species

FIGURE 7

MATERIAL EXAMINED.—Paitilla Beach, Balanus

Zone (4); Paitilla Beach, Tetraclita Zone (18, holotype, USNM 53090; paratypes, USNM 53091, AHF

Poly 1133).

DESCRIPTION.—The holotype is a complete specimen, 18 mm long and 1 mm wide without setae,

with 62 setigers. It is yellowish and has scattered

brown pigment spots on the anterior end.

The prostomium (Figure 7e) is very broad and

has a pair of short frontal antennae that are wellseparated basally. The large palpophores are ovate

and the short palpostyles button-shaped. The anterior eyes are crescentic; the posterior ones are

ovate. The four pairs of tentacular cirri are slender;

three pairs are of similar length and reach approximately setiger 2. The fourth pair is longer and

reaches the posterior margin of setiger 4.

The proboscis (Figure 7a,e) is everted in the

holotype; the jaws are thick and have five or six

distally directed, pointed teeth. Paragnaths are

present on all areas. Area i has three paragnaths

in a transverse row and one in front of the others.

Area n has 21 or 22 paragnaths in rows in a triangular patch. Area m has 36 paragnaths arranged

in 4 rows in an ovate patch. Area iv has 35 or 36
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Neanthes pseudonoodti, new species: a, proboscis, ventral view, X 50; b, far posterior parapodium, anterior view, X 95; c, neuropodial heterogomph falciger, posterior setiger,

X 950; d, posterior parapodium, anterior view, X 160; e, anterior end, dorsal view, X 50.

paragnaths in 6 rows in a quadrangular patch

(the anteriormost row on the right-hand side is not

visible in Figure 7a). Area v has one very large paragnath, and a pair of small paragnaths on either

side represent area vi. Areas vn and vm form a

girdle consisting of two rows of larger, and several

irregular rows of smaller, paragnaths. This girdle

is continued dorsally in the lateral parts of area vi.

Anterior parapodia have both note*- and neuropodia of the same length; each notopodium has a

short, rounded superior lobe; a short, nearly rudimentary acicular lobe and a long, digitate inferior

lobe. The neuropodia are bifid; each superior lobe

is distally truncate and has low, truncate pre- and

postsetal lobes; each inferior lobe is a little shorter

than the superior lobe and digitate. The short

ventral cirri are slightly clavate in all setigers; the

digitate dorsal cirri are attached distally on the

superior lobe of the notopodium. The superior

notopodial lobes become increasingly elongated in

posterior setigers (Figure Id) and are more than

three times the length of all other parapodial parts

in far posterior setigers. The dorsal cirrus remains

of the same size in all setigers and finally appears

as a short, inconspicuous distal part of the large

superior lobe (Figure 76).

All notopodial setae are similar; each is a

homogomph spiniger with a long, slender appendage. Homogomph spinigers are also present in

superior neuropodial fascicles; spinigers in the

inferior neuropodial fascicles are heterogomph.

Heterogomph falcigers are present in both superior

and inferior neuropodial fascicles. Each superior

fascicle contains a single, very large falciger (Figure

7c) with a short, slightly geniculate appendage with

four or five hairs along the cutting edge near the
NUMBER 221 29

base. Inferior fascicles have more numerous, smaller

falcigers of similar shape.

Neanthes pseudonoodti belongs to group IIB2c

according to the system suggested by Fauchald

(1972). Other species in this group include N.

noodti Hartmann-Schroder (1962:129-130, pi. 11:

figs. 65-66, pi. 12: fig. 68, pi. 20: fig. 67), N. ruficeps

(Ehlers, 1905:24-25, pi. 3: figs. 10-15), and N. seridentata Hartmann-Schroder (1959:138-142, figs.

100-110).

The paragnaths on area vi are arranged in a

nearly straight transverse row in N. seridentata;

the other species have these paragnaths arranged

in a small patch. Neanthes ruficeps lack paragnaths

on areas v and vi; N. noodti and N. pseudonoodti

have paragnaths on all areas.

Neanthes noodti and N. pseudonoodti resemble

each other but differ in the distribution and shape

of the paragnaths, especially on areas vn and vm.

The dorsal cirri are nearly half the total length of

the prolonged superior notopodial lobe in N.

noodti and less than one-fifth of the total length in

N. pseudonoodti.

Neanthes succinea (Frey and Leuckart, 1847)

Nereis succinea Frey and Leuckart, 1874:154.—Monro, 1933a:

42-43, fig. 18.

Neanthes succinea (Frey and Leuckart).—Hartman, 1968:529-

530, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (2); Balboa Dock, scrapings from

piles of quays (1); Col6n, from deck of wreck in

Lim6n Bay (1) and from scrapings from piles of

quays (11); all coll. Crossland.

REMARKS.—Neanthes succinea has large foliose

notopodial superior lobes in posterior setigers with

the dorsal cirri attached subdistally. It has been

reported from worldwide areas.

Neanthes species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (8).

REMARKS.—The specimens differ from all species

mentioned above in the structure of the neuropodial falcigers. They are poorly preserved and

cannot be further identified.

Nereis callaona (Grube, 1857)

FIGURE 8/-/I

Nereilepas callaona Grube, 1857:165-166.

Nereis callaona (Grube).—Hartman-Schroder. 1962b:399-400.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (10); Galeta Reef, Thalassia Zone (1); Galeta

Reef, Zoanthus Zone (1); Paitilla Beach, Hydroid

Zone (3); Paitilla Beach, Tetraclita Zone (25).

REMARKS.—The present specimens fit with the

species as reviewed by Hartmann-Schroder (1962)

except that the homogomph and heterogomph

falcigers have the distal parts of the appendages

ankylosed rather than recurved as indicated by

Hartmann-Schroder (Figure 8/,/i). Specimens from

both sides of the Isthmus are similar.

Nereis callaona is rather widespread in the eastern Pacific Ocean; the present records are the first

from the western Atlantic Ocean.

Nereis panamensis, new species

FIGURE 6d-i

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1, holotype, USNM 53139); Galeta Reef,

Laurencia Zone (27, paratypes, USNM 53140 and

AHF Poly 1136).

DESCRIPTION.—The holotype is a complete specimen, 12 mm long and 1.1 mm wide without setae,

with 52 setigers. Two dark brown patches are on the

lateral sides of the prostomium; the remainder of

the specimen is light yellow with scattered brown

dots. The long anal cirri are slender.

The prostomium (Figure 6g) is distinctly pentagonal with a slightly bifid anterior margin. The

long frontal antennae project beyond the tips of

the palps. The palpophores are ovate, and the

short palpostyles are button-shaped. One pair of

dark eyes is present near the posterior margin of

the prostomium. Three of the four pairs of tentacular cirri reach just beyond the peristomium; the

dorsoposterior pair is longer and reaches setiger

5; all tentacular cirri are slender and digitate.

The proboscis (Figure 6h-i) is nearly globose

when fully everted. The jaws are slender and have

seven or eight teeth along the cutting edge.

Paragnaths are present on all areas except i and v.

Area n has three cones in a transverse row; area

HI has three cones in a transverse row; area iv has
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Perinereis anderssoni Kinberg: a, posterior parapodium, anterior view, X 95;

b, neuropodial heterogomph falciger, posterior setiger, X 950. Nereis riisei Grube: c, posterior

parapodium, anterior view, X 50; d, notopodial homogomph falciger, posterior setiger, X 950;

e, neuropodial heterogomph falciger, posterior setiger, X 950. Nereis callaona (Grube):

/, neuropodial heterogomph falciger, posterior setiger, X 950; g, notopodial homogomph ralciger,

posterior setiger, X 950; h, posterior parapodium, anterior view, X 95.

eight cones in two rows, five in the posterior, and

three in the anterior row. Area vi has five cones in

a transverse row, and areas vu and vm have six

large cones in a single row.

The two first pairs of parapodia have reduced

notopodia; otherwise all parapodia are similar.

Each notopodium (Figure 6/) is bifid and has a

distinct acicular lobe and a bluntly conical, thick

inferior lobe; a distinct superior lobe is absent.

Dorsal cirri are large in all setigers, but become

prolonged in posterior setigers. All neuropodia are

bifid; the superior part, which carries the aciculum,

has low, poorly developed pre- and postsetal lobes;

the thick inferior lobe is digitate and projects

beyond the tip of the superior lobe. Ventral cirri

are long and slender.
NUMBER 221

Notopodial setae include homogomph spinigers

and falcigers; falcigers are present only in median

and posterior setigers. Each homogomph falciger

(Figure 6e) has a thick shaft and a short, bifid or

trifid appendage; the third tooth is very small and

may be worn off or absent in some hooks. Neuropodial fascicles contain heterogomph spinigers and

falcigers in both supra- and subacicular positions.

Each falciger (Figure 6d) has a short, slightly curved

appendage with transverse rows of teeth along the

cutting margin.

Nereis panamensis resembles N. jacksoni Kinberg,

1866 (as reviewed by Augener, 1922) in the distribution of paragnaths and the structure of the

homogomph falcigers. A distinct group of species

appears with bidentate or tridentate homogomph

falcigers and reduced superior notopodial lobes.

The earliest described species in the group is N.

jacksoni; others include N. thompsoni Kott (1951:

103-105, fig. 5), N. denhamensis Augener (1913:156-

159, pi. 2: fig. 51, text fig. 16), and N. funchalensis

Langerhans (1880:287-289, see Fauvel, 1927:409,

fig. 138f-n; Monro, 1933a:43, fig. 18).

Nereis jacksoni reducta was described by

Hartmann-Schroder (1960:110-112, figs. 144-146)

from the Red Sea; later Hartmann-Schroder (1965:

121-123, figs. 50-51) indicated that this subspecies

was identical with the main form and withdrew

the name; she also indicated that she thought N.

heirissonensis and N. denhamensis, both described

by Augener (1913) were synonymous with N. jacksoni. There appears to be general agreement that

N. heirissonensis is in fact synonymous with N.

jacksoni, but Hartman (1954:31) retains N. denhamensis as a distinct form.

Nereis panamensis differs from N. jacksoni (and

all species listed above, whether valid or not) on

the distribution of the paragnaths and on the shape

of the appendage of the heterogomph falcigers.

Nereis riisei Grube, 1857

FIGURE Sc-e

Nereis riisei Grube, 1857:162-163.—Monro, 1933a:43-44.—

Hartraan, 1940a:221-222, pi. 33: fig. 37.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Coralline Zone (1);

Galeta Reef, Thalassia Zone (2). Col6n, scrapings

from piles of quays (4); Gorgona Island, fine sand

and shells, 20 fm (11): both coll. Crossland.

REMARKS.—The present specimens fit very well

with the species as described by Hartman (1940).

The homogomph falcigers (Figure 8e) have recurved

appendages that have series of teeth along the

cutting edges. The heterogomph falcigers (Figure

8d) are strongly recurved distally and have dense

series of long, slender teeth along the cutting edges.

Nereis riisei has been reported from both sides

of the Isthmus and appears to be most common in

the western Atlantic Ocean in warm waters.

Perinereis anderssoni Kinberg, 1866

FICURE Sa,b

Perinereis anderssoni Kinberg, 1866:175.—Hartman. 1948:72-

73; 1951:47, pi. 13. fig. 6.

Perinereis bairdii Webster and Benedict, 1884:312, pi. 8:

figs. 22-28—Monro. 1933a:41.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Coralline Zone (2);

Galeta Reef, Laurencia Zone (84); Galeta Reef,

Thalassia Zone (21). Colon, from washings of weeds,

etc. of the coral reef in Limon Bay (29), and from

wreck in Limon Bay (13), from coral flat at the

southwest corner of Limon Bay (13), and from the

outer piles of the quays of the docks (5); all coll.

Crossland.

REMARKS.—Perinereis anderssoni has the superior

notopodial lobes prolonged in posterior setigers

(Figure 8a); the dorsal cirrus is attached near the

middle of the lobe. Each heterogomph falciger

(Figure 8b) has a short, gently curved appendage

with series of teeth along the cutting edges.

Perinereis anderssoni is common in the warmer

parts of the western Atlantic Ocean; it has not been

reported from the Pacific Ocean.

Perinereis species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1).

REMARKS.—The present specimen is badly preserved and cannot be further identified.

Platynereis dumerilii (Audouin and

Milne Edwards, 1834)

FIGURE 4d-f

Nereis Dumerilii Audouin and Milne Edwards, 1934:196-199,

pi. 4A: figs. 10-12.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Plalynereis dutnerilii (Audouin and Milne Edwards).—Hartman, 1968:561-562, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (25). Col6n, from piles of the quays (178);

Taboga Island, at low tide, obtained by breaking

up standstone (1); coll. Crossland.

REMARKS.—Platynereis dumerilii appears to be

rather variable. The present specimens have the

homogomph falcigers (Figure Ad) distinctly bidentate and the tip very strongly curved. Each appendage of the heterogomph falcigers (Figure 4e) has a

distinct subdistal knob and the tip is strongly

curved; the cutting edge is dentate.

The specimen reported from Taboga was found

in a sample otherwise identified as Lysidice ninetta

by Monro.

Platynereis dumerilii is very widespread in warm

waters.

Platynereis species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (4); Galeta Reef, Thalassia Zone (4).

REMARKS.—These specimens have either been

dried out or are so incomplete that they cannot

be further identified.

Pseudonereis gallapagensis Kinberg, 1866

FIGURE 4g,/i

Pseudonereis gallapagensis Kinberg, 1866:174.—Hartman,

1940a:231.—Hartmann -Schroder, 1962b:432-4S4.

MATERIAL EXAMINED.—Paitilla Beach, Balanus

Zone (2); Paitilla Beach, Tetraclita Zone (38).

REMARKS.—Pseudonereis gallapagensis has the

superior notopodial lobe strongly prolonged and

foliose in posterior setigers (Figure Ah); the short

FIGURE 9.—Diopatra chiliensis Quatrefages: a, third parapodium, anterior view, X 50; b, giant

hooded hook, third setiger, X 385; c, normal hooded hook, third setiger, X 385. Pseudonereis

variegata (Grube): d, neuropodial heterogomph falciger. posterior setiger, X 950; e, posterior

parapodium. posterior view. X 52.
NUMBER 221 33

dorsal cirrus is digitate and attaches distally to the

superior lobe. Each heterogomph falciger (Figure

4g) has a large, gently curved appendage with

approximately ten teeth in double rows near the

base.

Pseudonereis gallapagensis may be widespread in

both the Atlantic and Pacific Oceans, but the best

verified records come from the Pacific Ocean. It has

frequently been confused with other species in the

genus, including P. variegata.

Pseudonereis variegata (Grube, 1857)

FIGURE 9d,e

Nereilepas variegata Crube, 1857:164-165.

Pseudonereis variegata (Grube).—Monro,

mann-Schroder, 1962b: 434-435.

1933a:45—HartMATERIAL EXAMINED.—Colon, from the quays (1,

heteronereis); Col6n, from the coral flat at the

southwest corner of Limdn Bay (3); Coldn, from

deck of wreck in Lim6n Bay (2); all coll. Crossland.

REMARKS.—This species is recognized as indicated

in the key and as remarked by Hartmann-Schroder

(1962b). The dorsal cirrus is attached subdistally to

the superior notopodial lobes (Figure 9e) and the

paragnaths are in several rows on areas vn and vm.

The falcigers (Figure 9d) are recurved and have

hairs in single rows.

Pseudonereis variegata may be widespread in

warm waters.

Nereidae species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef, Laurencia Zone (1); Paitilla

Beach, Tetraclita Zone (posterior ends).

REMARKS.—These specimens have been died out

or are incomplete and cannot be further identified.

Family NEPHTYIDAE

Key to the Species from Panama

1. Interramal cirri involute 2

Interramal cirri recurved 3

2. Postsetal lobes absent in posterior setigers; notopodial cirri longer than the interramal

cirri in posterior setigers Aglaophamus tabogensis

Postsetal lobes distinct in all setigers; interramal cirri short and slender in all setigers

Aglaophamus dicirris

3. Dorsolateral surface of body covered by flattened expansions from the dorsum

Nephtys squamosa

Dorsum not covered by expansions from the dorsum 4

4. Interramal cirri present from setiger 8, erect neuropodial cirri present in median setigers ....

Nephtys iwotwoi

Interramal cirri present from setiger 4, erect neuropodial cirri present in anterior setigers ....

Nephtys singular**

Aglaophamus dicirris Hart man, 1950

Aglaophamus dicirris Hartman, 1950:122-124, pi. 18: figs. 1-8.

Nephthys dibranchis.—Monro, 1933a:56-57. fig. 24 [not

Grube, 1878].

MATERIAL EXAMINED.—Gorgona Island, 20 fm,

fine sand and shell fragments (8); Gorgona Island,

30 fm, muddy sand and shell fragments (1); both

coll. Crossland.

REMARKS.—Aglaophamus dicirris has a very characteristic thin prostomium with a strongly arched

anterior margin; a pair of black eyespots are distinct on the posterior part of the prostomium. The

proboscis is covered with small pointed warts; a

median papilla is absent.

Aglaophamus dicirris is known from tropical

America.

Aglaophamus tabogensis (Monro, 1933)

Nephthys tabogensis Monro, 1933a:53-55, fig. 23 [in part].

Aglaophamus tabogensis (Monro).—Hartman, 1950:125.

MATERIAL EXAMINED.—Taboga Island, 6-12 fm,

mud (15, syntypes, BMNH), coll. Crossland, det.

Monro.

REMARKS.—Aglaophamus tabogensis was restricted by Hartman (1950) to include the material

listed above; the remainder reported by Monro

from Panama was separated into a distinct species:
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Nepthys monroi. Aglaophamus tabogensis can be

separated from the only other species of Aglaophamus reported from Panama as indicated above.

Aglaophamus tabogensis is known only from the

type-locality.

Nephtys monroi Hartman, 1950

Nephtys monroi Hartman, 1950:107-108, pi. 17: fig. 1.

Nephthys tabogensis Monro, 193Sa:53-55, fig. 23h [in part].

MATERIAL EXAMINED.—Taboga Island, 6-12 fin,

mud (1), coll. Grassland, det. Monro and Hartman.

REMARKS.—Nephtys monroi was based on a single

large specimen of the collection made at Taboga.

It differs from all other Nephtys along this coast'

on a combination of characters. It has interramal

cirri from setiger 8, erect neuropodial lobes present

in median segments only, and incised preacicular

lobes in anterior setigers.

Nephtys monroi is known only from the original

record.

Nephtys singuktris Hartman, 1950

Nephtys singulaiis Hartman, 1950:98-100, pi. 15: figs. 1-6.

Nephthys sp — Monro, 1933a:51-53, fig. 22.

MATERIAL EXAMINED.—Coiba Island, in sand at

low tide, fine sea grass and dead coral embedded

in the sand (2), coll. Crossland.

REMARKS.—Nephtys singularis has interramal

cirri present from setiger 4 and erect neuropodial

lobes present in anterior setigers. The presetal lobes

are bilobed in all setigers, and the neuropodial

postsetal lobes are greatly expanded in posterior

setigers.

Nephtys singularis is known from shallow-water

areas from western Mexico to Panama.

Nephtys squamosa Ehlers, 1887

Nephtys squamosa Ehlers, 1887:128-131, pi. 37: figs. 7-10.—

Monro. 1933a:52-53—Hartman, 1950:110-111.

MATERIAL EXAMINED.—Gorgona Island, 20 fm,

fine sand and shell (2), coll. Crossland.

REMARKS.—Nepthys squamosa is characterized by

the expanded, imbricated flattened leaves covering

the dorsum in median and posterior setigers. It is

common in shallow water in tropical America,

both on the Atlantic and Pacific side.

Family GLYCERIDAE

Key to the Species from Panama

1. Parapodia with 1 postsetal lobe Glycera oxycephala

Parapodia with 2 postsetal lobes 2

2. Dendritically branched branchiae present Glycera americana

Branchiae absent 3

3. All proboscideal organs tall and slender with longitudinal ribs Glycera tesselata

AH proboscideal organs smooth, either tall and slender or shorter and wide

Glycera abranchiata

Glycera abranchiata Treadwell, 1901

Glycera abranchiata Treadwell, 1901:200-201, fig. 49.—Jones,

1962:183, figs. 41-48.—Fauchald, 1973:21-22. fig. la-e.

Glycera sp.—Monro, 19S3a:57-58, fig. 25.

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (1); Coiba Island, in sand at low

water, fine sea grass and dead coral embedded in

sand (1); Gorgona Island, low water spring tide (2)

and 20 fm, sand and shell (1), all coll. Crossland.

REMARKS.—Glycera abranchiata has been confused with G. tesselata from which it can be separated by the presence of two kinds of proboscideal

organs in the former and only one kind in the

latter. It has previously been reported from sandy

areas in the Caribbean Sea and Panama; the

present records extend its distribution into the

Pacific Ocean in similar environments.

Glycera americana Leidy, 1855

Glycera americana Leidy, 1855:147-148, pi. 11: figs. 49, 50.—

Monro, 1933a:57.—Hartman, 1950:73-75; 1968:613-614, 1 fig.

MATERIAL EXAMINED.—Coiba Island, in sand at

low water, fine sea grass and dead coral embedded

in the sand (2); Gorgona Island, 30 fm, muddy

sand with shell fragments (1); Perlas Island, St.
NUMBER 221 35

Elmo Bay, dredging and trawling on east side of

bay (1); coll. Crossland.

REMARKS.—Glycera americana is one of the most

common glycerids on both sides of the Americas.

The branchiae are usually distinct; if they are

retracted, the species can be identified by the proboscideal organs. They are of two kinds: one is

large and ovate and lacks ridges; the other is

smaller, usually somewhat more slender than the

other kind and equipped with two oblique or transverse ridges.

Glycera oxycephala Ehlers, 1887

Glycera oxycephala Ehlers, 1887:121-123, pi. 41: figs. 7-11.—

Hart man, 1950:70-71, fig. 3, pi. 10: figs. 3, 4.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1); Galeta Reef, Mangrove Zone (1).

REMARKS.—Glycera oxycephala has a single postsetal lobe in all setigers, and the proboscideal

organs are all of one kind, tall and slender with 9

or 10 transverse ridges. The species is most common in warm water, but has been reported as far

north as Oregon on the west coast; it is found on

both sides of the Americas.

Glycera tesselata Grube, 1863

Glycera tesselata Grube, 1863:41-42, pi. 4: fig. 4.—Hartman,

1950:77-78, pi. 10: fig. 11.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1).

REMARKS.—Glycera tesselata is widely distributed

in warm waters. It is characterized by its lack of

branchiae, the two rounded postsetal lobes, and the

tall slender, longitudinally ribbed proboscideal

organs. It can be separated from the similar G.

abranchiata as indicated in the key.

Family GONI ADI DAE

Goniada acicula Hartman, 1940

Goniada acicula Hartman, 1940a:252-254, pi. 44: figs. 132-

141.—Hartman, 1950:31-32, pi. 4: figs. 2-7; 1968:649-650,

5 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1).

REMARKS.—Goniada acicula has strong, gently

curved setae in the posterior notopodia. The posterior body region is missing in the present specimen, but it fits otherwise very well with the species

as described, both in terms of parapodial characters

and in the structure of the proboscideal armament.

Goniada acicula is known from warm waters on

both sides of the Americas.

Family ONUPHIDAE

Key to the Species from Panama

1. First 6 setigers modified and anteriorly directed Americonuphis reesei

Maximally 1 setiger anteriorly directed 2

2. Branchiae single filaments or pectinate S

Branchiae spiralled in anterior segments 5

3. Branchiae single filaments, first present from setiger 23 Nothria gorgonensis

Branchiae pectinate, first present anterior to setiger 10 4

4. Composite spinigers in some anterior setigers Onuphis nebulosa

Composite spinigers absent Onuphis vermillionensis

5. Anterior parapodia with 2 postsetal lobes Diopatra chiliensis

Postsetal lobes single in all setigers 6

6. Anterior hooded hooks tridentate Diopatra denticulate

Anterior hooded hooks bidentate 7

7. Subdistal tooth of hooded hooks at right angles to the shaft of the setae Diopatra ornata

Subdistal tooth of hooded hooks oblique to the shaft of the setae Diopatra cuprea

Americonuphis reesei Fauchald, 1973 MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (&). Americonuphis reesei Fauchald, 1973:22-23, fig. 3a-e.

 r w

Onuphis magna.—Monro. I933a:76 [not Andrews, 1891]. REMARKS.—T he present specimens are as de-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

scribed by Fauchald (1973); the maximal number

of branchial filaments is six or seven in all specimens rather than the 12 found in Americonuphis

magna.

Americonuphis reesei is known from intertidal

and shallow-water sandy and muddy areas in the

Gulf of Panama.

Diopatra chiliensis Quatrefages, 1865

FIGURE 9a-c

Diopatra chiliensis Quatrefages, 1865:342-344.—Monro, 1933a:

72-73, fig. 30.

MATERIAL EXAMINED.—Balboa, rock and rockpools at low tide (8); Coiba Island, in sand at low

tide, fine seagrass and dead coral embedded in the

sand (2); both coll. Crossland.

REMARKS.—I follow Hartman (1944b: 60-61) and

consider D. chiliensis sensu Ehlers (1901:123-125,

pi. 15: figs. 1-12) as a synonym of D. obliqua

Hartman and consider D. chiliensis sensu Monro

as the valid concept of Quatrefages's species.

The presetal lobe of the first setiger (Figure 9a)

is a transverse fold with a deeply incised, flattened

distal lobe; the two postsetal lobes are digitate. The

hooded hooks of the anterior setigers are bidentate;

a few are about three times larger than the others

(Figure 9b-c).

Diopatra chiliensis is known from the southeastern Pacific Ocean.

Diopatra cuprea (Bosc, 1802)

Nereis cuprea Bosc, 1802:142.

Diopatra cuprea (Bosc).—Monro, 1933a:71-72. fig. 29.—

Hartman, 1944b:54-55, pi. 1: figs. 9-14.

MATERIAL EXAMINED.—Col6n, trawling in Limon

Bay, 5 fm, thin mud (1), coll. Crossland.

REMARKS.—Diopatra cuprea and D. ornata can be

separated only with difficulty. The two species,

however, can be consistently separated as suggested

in the key and in the following characters: the

teeth in the hooded hooks are less curved in D.

cuprea than in D. ornata, the papillation of the

tentacular styles is different and the presetal lobe

in anterior setigers is more oblique in D. cuprea

than in D. ornata. The separation is sufficiently

difficult, so it should not be undertaken except

with the aid of comparative material.

Diopatra cuprea is found in the western Atlantic

Ocean from Massachusetts to Brazil; it has never

been reported from the eastern Pacific Ocean.

Diopatra denticulata Fauchald, 1968

Diopatra denticulata Fauchald, 1968:5-7, pi. la-g.

Diopatra dentata.—Monro, 1933a:73-76, fig. 31 [in part, not

Kinberg, 1865].

MATERIAL EXAMINED.—Gorgona Island, dredging,

20 fm (2), and dredging 30 fm, mud, sand and

shell fragments (5); both collected by Crossland.

REMARKS.—Diopatra dentata sensu Monro consists of two species, D. denticulata and D. ornata.

The present specimens have tridentate anterior

hooded hooks and fit very well with D. denticulata

in the details of the presetal lobes in anterior segments. Diopatra dentata Kinberg (1865) has bidentate anterior hooded hooks (Hartman, 1948:86-87,

pi. 12: figs. 1-7).

Diopatra denticulata was described from western

Mexico; the present records extend its distribution

to the Gulf of Panama.

Diopatra ornata Moore, 1911

Diopatra ornata Moore, 1911:273-277, pi. 18: figs. 77-85.—

Hartman. 1944b:55-56, pi. 1: figs. 15-20.—Fauchald,

1968:10-11, pi. 2: fig. c.

Diopatra dentata.—Monro, 1933a:73-76, fig. 31 [in part, not

Kinberg, 1865].

MATERIAL EXAMINED.—Perlas Islands, St. Elmo

Bay, dredging and trawling on east side of bay,

6-9 fm, sand and shell (10), coll. Crossland.

REMARKS.—The separation between D. cuprea

and D. ornata has been indicated above. Monro

(1933a:73) thought Moore's species was identical

with D. dentata from Australia. Hartman (1948:87)

indicated that the two species differed in the distribution of the subacicular hooks, first present from

setiger 15 in D. dentata and from setiger 30 in

D. ornata and in the maxillary formula. The maxillary formula for D. ornata is left: 1-8-7-5 and

right 1-10-7-1; for D. dentata, the formula is

1-9-12-0 and 1-8-9-6; maxilla iv on the left side

is an oval piece without any distinct point. This

difference is sufficiently large so the two species

are considered distinct here.

Diopatra ornata is known from the eastern

Pacific Ocean from Vancouver to Panama.
NUMBER 221 37

Nothria gorgonensis (Monro, 1933)

Onuphis gorgonensis Monro, 1933a:80-82, fig. 34.

Nothria gorgonensis (Monro).—Hartman, 1944b:83.

MATERIAL EXAMINED.—Gorgona Island, fine sand

and shell, 20 fm (1, holotype, BMNH), coll.

Crossland.

REMARKS.—The species is as described by Monro.

The secondary and tertiary teeth in the hooded

hooks are slightly narrower than as illustrated.

Nothria gorgonensis is known only through the

original record.

Onuphis nebulosa Moore, 1911

Onuphis nebulosa Moore, 1911:269-273, pi. 17: figs. 58-68.—

Monro, 1933a:76-78, fig. 32.—Hartman, 1968:699-700, 7

figs, [unnumbered].—Fauchald, 1968:36.

MATERIAL EXAMINED.—Perlas Islands, St. Elmo

Bay, dredging and trawling on east side of the bay,

6-9 fm, shells and sand (15), coll. Crossland.

REMARKS.—Onuphis nebulosa has large, pseudocomposite or simple hooded hooks in anterior

setigers; each hook has the teeth characteristically

strongly curved. Composite spinigers are present in

a number of setigers, usually most easily seen in

setigers 5-15.

Onuphis nebulosa is found in the eastern Pacific

Ocean from California to Panama.

Onuphis vennillionensis Fauchald, 1968

Onuphis vermillionensis Fauchald, 1968:41-42, pi. 11.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (5).

REMARKS.—Onuphis vermillionensis has branchiae present from setigers 7 or 8; the first 9 or 10

setigers have cirrifonn ventral cirri. All anterior

hooded hooks are tridentate and large; simple tridentate hooks are present in 10 or 11 setigers. Composite spinigers are absent. Subacicular hooks are

first present from setiger 12.

Onuphis vermillionensis was originally described

from Golfo de California; the present records are

from the western Atlantic Ocean in very shallow

water.

Family EUNICIDAE

Key to the Species from Panama

1. Prostomium with a single occipital tentacle Nematonereis unicomis

Prostomium with at least 3 occipital tentacles 2

2. Prostomium with 3 occipital tentacles Lysidice ninetta

3.

Prostomium with 5 occipital tentacles

6.

8.

9.

10.

11

Peristomial drri absent 4

Peristomial cirri present 5

4. All composite setae spinigers Marphysa sanguinea

All composite setae falcigers Marphysa amadae, new species

5. Subacicular hooks absent Palola siciliewis

Subacicular hooks present 6

Subacicular hooks and acicula dark or black 7

Subacicular hooks and acicula light yellow 12

7. Branchiae absent, or if present, as single filaments only Eunice (Nicidion) cariboea

Branchiae present, with at least 2 filaments in part of the body 8

Branchiae first present after setiger 10 9

Branchiae first present before setiger 10 10

Branchiae first present from setigers 18-21; subacicular hooks first present from setigers

35-42 Eunice afra

Branchiae first present from setigers 21-26; subacicular hooks first present from setigers

19-26 Eunice filamentosa

Occipital tentacles distinctly articulated; composite hooded hooks with rudimentary subdistal teeth Eunice reducta

Occipital tentacles smooth or wrinkled; composite hooded hooks with subdistal teeth well

developed 11

Occipital tentacles twice as long as the prostomium; branchiae erect Eunice aphrodiuris

Occipital tentacles barely as long as the prostomium; branchiae flaccid Eunice mutilata
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

12. Subadcular hooks bidentate 13

Subacicular hooks tridentate 14

13. Occipital tentacles with moniliform articles; maximal number of branchial filaments 10 ...

Eunice biannUlata

Occipital tentacles with cylindrical articles; maximal number of branchial filaments 15

Eunice websteri

14. Occipital tentacles with cylindrical articles; branchiae present from setiger 3

Eunice vittatopsis

Occipital tentacles with moniliform articles; branchiae present from setiger 4 to 6 15

15. Branchiae in far posterior setigers single filaments Eunice antennata aedificatrix

Branchiae in far posterior setigers with 3 to 5 filaments Eunice antennata, sensu stricto

Eunice afra Peters, 1854

Eunice afra Peters, 1854:611.—Hartman, 1944b: 110-111. pi. 6:

figs. 135-139.—Fauchald, 1970:16-18, pi. 1: figs. h-i.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (3); Galeta Reef, Laurencia Zone (21); Galeta

Reef, Thalassia Zone (12).

REMARKS.—The present specimens agree with

'specimens reported from western Mexico by

Fauchald (1970). Branchiae are present from setigers

18-21; 30 of 36 specimens had the first branchiae

either on setiger 19 or 20. Subacicular hooks are

present from setigers 29-38 in the present material.

The number of branchial filaments varies between

two and three in the present material.

Eunice afra reported from Panama by Monro

(1933a:66-67) is E. mutilata as indicated by

Hartman (1944b).

Eunice afra is known from circumtropical areas;

it appears to be rather variable, as indicated by

Fauchald (1970).

Eunice antennata (Savigny, 1818)

Leodice antennata Savigny, 1818:322.

Eunice antennata (Savigny).—Monro, 1833a:59-60.—Fauchald,

1970:20-22. pi. la-c.

MATERIAL EXAMINED.—Coldn, from rotten wood

from deck of wreck in Limon Bay (2); Gorgona

Island, from coral (5); Taboga Island, from floats of

the stage at the end of the hotel pier (1); all coll.

Crossland.

REMARKS.—Eunice antennata, sensu stricto, has

branchiae from setigers 4-6 with maximally 10

branchial filaments and the branchiae in the posterior end have from three to five filaments. Yellow,

tridentate subacicular hooks are present from setigers 15-25 in most normal-sized specimens; in

really large specimens they may be first present

from setigers 26-28, but this appearance is rare.

Eunice antennata is very widespread, possibly

circumtropical, but some discrepancies in the descriptions from different areas make it difficult to

assess the total distribution of the species.

Eunice antennata aedificatrix Monro, 1933

Eunice antennata aedificatrix Monro, 1933a:60-61.—Fauchald,

1970:22-23.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (2); Galeta Reef, Laurencia Zone (1); Galeta

Reef, Zoanthus Zone (1). Balboa, rock and rockpools at low tide (3), from scrapings off buoy at

canal entrance (3) and from piles of quays (7);

Colon, scrapings off piles of quays (2); coll.

Crossland (types not designated).

REMARKS.—The material from Balboa represents

the syntype material of this subspecies. The subspecies is characterized by having simple, rather

than pectinate, branchiae in the posterior end.

The present specimens have subacicular hooks first

present from setigers 21-34, well within the range

suggested by Fauchald (1970).

Eunice antennata aedificatrix is known from

intertidal areas at or near Balboa and in western

Mexico; the present records also include areas in

the western Atlantic Ocean near Panama.

Eunice aphroditois (Pallas, 1788)

Nereis aphroditois Pallas, 1788:229.

Eunice aphroditois (Pallas).—Monro, 1933a:58-59.—Hartman,

1944b:109-l 10.—Fauchald, 1970:24-25, pi. 3a.b.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef, Laurencia Zone (1). Taboga

Island, from floats at the stage at the end of the

hotel pier (2), coll. Crossland.

REMARKS.—Eunice aphroditois has strongly
NUMBER 221 39

branched branchiae present from setigers 5-7. The

subacicular hooks are present from setigers 15-54;

the first occurrence is strongly dependent on the

size of the specimen. The species is considered

circumtropical and has previously been reported

from both sides of the Isthmus of Panama.

Eunice biannulata Moore, 1904

Eunice biannulata Moore, 1904:487-490, pi. 37: figs. 10-18, pi.

38, fig. 42.—Fauchald, 1969:2-4. fig. 1; 1970:25-26.

Eunice longicirrata.—Monro, 1933a:61-62 [not Webster, 1884

« Eunice webstert].

MATERIAL EXAMINED.—Gorgona Island, from

coral (4), coll. Crossland.

REMARKS.—The present specimens have the

moniliform articles of the antennae and the low

number of branchial filaments characteristic of this

species. It has been found only in the eastern

Pacific Ocean from southern California to Panama.

Eunice filamentosa Grube, 1856

Eunice filamentosa Grube, 1856:56.—Monro, 1933a:65-66, fig.

27.—Hartman, 1944b:107. pi. 6: figs. 123-126.—Fauchald,

1970:31-33, pi. 3c-g.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (3); Gorgona Island, at low tide (2); coll.

Crossland.

REMARKS.—The present specimens are well

within the established limits of variation in terms

of the origin of the branchiae and the subacicular

hooks. The first branchiae are present on setigers

24-34, and the subacicular hooks from setiger 18

appear in all five specimens. Eunice filamentosa

is further characterized by the strongly hammerheaded acicula in posterior setigers and strongly

beaked, bidentate, dark, subacicular hooks.

Eunice filamentosa was described from tropical

west Atlantic areas and is also common in the

eastern Pacific Ocean.

Eunice mutilata Webster, 1884

Eunice mutilata Webster, 1884:315-316. pi. 9: figs. 36, 36a-d40.—Hartman, 1944b: 113-114, pi. 6: figs. 140-141.—

Fauchald, 1970:37-38, pi. Sj-k.

Eunice afra.—Monro, 1933a:66-67 [not Peters, 1854].

MATERIAL EXAMINED.—Taboga Island, from

floats at the end of the hotel pier (5), coll.

Crossland.

REMARKS.—Hartman (1944b: 113), pointed out

that the specimens reported by Monro (1933a)

from Panama, should belong to this species rather

than to E. afra. I concur after having reexamined

the material. Both species are known from the area

and can be separated as indicated in the key.

Eunice mutilata is common in warm waters on

both sides of the Americas.

Eunice (Nicidion) cariboea Grube, 1856

FIGURE 10

Eunice cariboea Grube, 1856:57.—Monro, 1933a:63.

Eunice (Nicidion) cariboea Grube.—Hartman, 1944b: 123-124,

pi. 7: figs. 157-163; pi. 8: fig. 178.—Fauchald, 1970:38-39.

Eunice cariboea var. kinbergii.—Monro, 1933a:63.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (5); Galeta Reef, Coralline Zone (2);

Galeta Reef, Laurencia Zone (261); Galeta Reef,

Thalassia Zone (1). Coldn (16); Coldn, coral flat at

southwest corner of Lim6n Bay (6); Coiba Island,

dredging off the convict settlement, 5 fm, smooth

bottom with branched Lithothamnion (6); Taboga,

n • 18 88 85 28 31

FIRST SUBACICULAR HOOKS ON SETIGEB NUMBER

FIGURE 10.—Distribution of the first occurrence of subacicular hooks in Eunice (Nicidion) cariboea Grube from

Panama.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

at low tide, obtained by breaking up beach sandstone (1); coll. Crossland.

REMARKS.—Eunice cariboea is here taken to

include E. kinbergi as discussed by Fauchald (1970).

All the more than 265 specimens in the newly

identified material lack branchiae. The distribution of the subacicular hooks is similar to the

distribution in specimens from western Mexico.

Figure 10 shows the distribution in specimens in

which this feature could be determined; some

specimens were too incomplete to be included in

the survey. The shape of the eyes varies with the

first start of the subacicular hooks as indicated by

Fauchald (1970).

Eunice (N.) cariboea is known from both sides

of tropical America.

Eunice reducta Fauchald, 1970

Eunice reducta Fauchald, 1970:39-43, pi. 5a-i.

Eunice tridentata.—Monro, 1933a:63-65, fig. 26 [not Ehlers,

1905].

MATERIAL EXAMINED.—Coiba Island, dredging off

the convict settlement in 9 fm (3), coll. Crossland.

REMARKS.—Eunice reducta has reduced subdistal

teeth in the composite hooded hooks, distinctly

articulated occipital antennae, and branchiae first

present from setiger 4 and limited to approximately

one-half of the body. The bidentate subacicular

hooks are black.

The relationship between E. tridentata Ehlers

from New Zealand and the present species was

discussed by Fauchald (1970). Eunice reducta is

known from the eastern Pacific Ocean between

Ecuador and western Mexico.

Eunice vittatopsis Fauchald, 1970

Eunice vittatopsis Fauchald, 1970:50-52, pi. 7a-d.

Eunice vittata.—Monro, 1933a:61 [not delle Chiaje, 1828].

MATERIAL EXAMINED.—Coiba Island, off a whale's

vertebra fished up from 20 fm (3), coll. Crossland.

REMARKS.—These specimens belong to E. vittatopsis in that they have short, distally rounded

hooks on the composite hooded hooks. Subacicular

hooks are present from approximately setiger 30,

and 35 to 40 pairs of branchiae are present.

Eunice vittatopsis was originally described from

Golfo de California; the present record is from the

Gulf of Panama.

Eunice websteri Fauchald, 1969

Eunice websteri Fauchald, 1969:12-14, fig. 6.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (2).

REMARKS.—These two specimens agree with E.

longicirrata as described by Webster (1884); the

name has had to be replaced since it was preoccupied in the combination E. longicirrata

(Kinberg, 1865) which is considered a synonym of

E. (2V.) cariboea.

The species concepts in this group of the genus

are still somewhat confused. Eunice websteri has

been reported from Brazil by Nonato and Luna

(1970:80-81, figs. 80-81); their description indicates

that the first branchiae should be present on setigers

4 or 5 with a maximal number of 10 branchial

filaments. The type of E. websteri has branchiae

from setiger 3 and a maximum of 15 branchial

filaments.

Subacicular hooks are present from setiger 31 in

the type; Nonato and Luna (1970) reported subacicular hooks present from setiger 20 in their

material. These differences may indicate that still

more species are presently involved in the concept

of E. websteri.

Eunice websteri is known from Bermuda and is

probably rather widely distributed in the western

Atlantic Ocean in tropical waters.

Eunice species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone; Galeta Reef, Laurencia Zone; Galeta Reef,

Thalassia Zone; Paitilla Beach, Hydroid Zone.

Coiba Island, dredging off the convict settlement

in 5 fm, smooth bottom with branched Lithothamnion (1), coll. Crossland.

REMARKS.—These posterior ends, median fragments, and poorly preserved specimens cannot be

further identified.

Lysidice ninetta Audouin and Milne Edwards, 1833

Lysidice ninetta Audouin and Milne Edwards, 1833:235, pi.

12: figs. 1-8.—Monro, 1933a:7O-71.—Hartman, 1944b:125.—

Fauchald, 1970:52-53.

Lysidice collaris Grube, 1870:495-496.—Monro, 1933a:69-70.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid
NUMBER 221 41

Zone (4); Paitilla Beach, Tetraclita Zone (3). Balboa,

rocks and rock-pools at low tide (3); Coiba Island,

dredging off the convict settlement in 5 fm, smooth

bottom with branched Lithothamnion (6) and in

sand at low water (1); Gorgona Island, in coral (2);

Taboga Island, at low tide, obtained by breaking

up beach sandstone (5) and from floats at the end

of the hotel pier (1); all coll. Crossland.

REMARKS.—Subacicular hooks are present from

setigers 15-19 in the present material, well within

the range established for the species from western

Mexico (Fauchald, 1970).

All present specimens are from the Pacific side

of Panama, but the species is well known also from

the western Atlantic; it appears to be circumtropical

in dispersal.

Marphysa amadae, new species

FIGURE 11

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (2, holotype, USNM 53094).

DESCRIPTION.—Both specimens are posteriorly

incomplete. The holotype, 34 mm long and 3 mm

wide, has 139 setigers. Preserved in alcohol, both

specimens are ivory-colored without color patterns.

The prostomium (Figure 1 la) is anteriorly deeply

bifid. The five occipital tentacles are smooth and

barely reach beyond the tip of the prostomium.

The ceratophores are short and wide. The first

peristomial segment is twice as long as the second

one, which is similar in length to the first setiger.

Prebranchial parapodia (Figure 116) are all

similar. Each has a rounded adcular lobe, a low

transverse presetal lobe, and a high, evenly rounded

postsetal lobe. The parapodial lobes are best

developed in anterior setigers. The digitate dorsal

and ventral cirri are similar in length. The dorsal

cirrus becomes shorter in late prebranchial segments, and the ventral cirrus is indistinct from late

prebranchial segments to the ends of the two

fragments.

Branchial parapodia (Figure lie) are similar;

each has barely distinct, elongated transverse preand postsetal lobes. The acicular lobe is gently

pointed, depending in part on the degree of protrusion of the acicula. The short dorsal cirri are

digitate; the ventral cirri are represented by blunt

projections.

Branchiae are present from setiger 54 in the

holotype and from setiger 47 in the paratype; the

first branchiae are all simple, but some branchiae

are bifid from approximately setiger 100. The

maximal number of branchial filaments is two.

Anterior setigers have three black, distally pointed

acicula. The number of acicula is reduced in late

prebranchial setigers, and the branchial setigers

have only one aciculum each. Subacicular hooks

FIGURE 11.—Marphysa amadae, new species: a, anterior end,

dorsal view, X 25; b, anterior parapodium, anterior view,

X 61; c, posterior parapodium, anterior view, X 61; d, bidentate composite hook, anterior setiger, X 950; e-g, appendages of composite hooks of postadcular fascicles, median

setiger, X 950.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

are present from setigers 54 in the holotype and

from setiger 45 in the paratype. Each is black and

distally bifid with the proximal tooth at least twice

as thick as the distal one. Composite falcigerous

hooded hooks are present in dense preacicular

fascicles in prebranchial parapodia. The appendages

are of several different kinds. Most setae have short,

distally bifid appendages (Figure lid), but in postacicular fascicles are found setae with long, slender

appendages; each of these appendages (Figure

1 le-g) is distally, evenly bifid, or the proximal tooth

may be reduced to a varying degree so that the

appendage may appear distally simple. Other setae

in postacicular fascicles include two different kinds

of limbate simple setae; these are present in all

setigers. The one kind is long and slender, and the

other is short and comparatively wide.

Marphysa amadae belongs to the group of

Marphysa that have only composite falcigers and

branchiae present over a long region of the body

(Group C2 according to Fauchald, 1970:210). It

differs from all other species in this group in having

small fascicles of composite postacicular setae in

which the distal end of each appendage is prolonged

and modified. All other species in this group have

normal bidentate composite setae.

Marphysa amadae is named for Dr. Amada A.

Reimers, Pennsylvania State University, who collected most of the new material identified in this

study.

Marphysa sanguinea (Montagu, 1815)

Marphysa sanguinea Montagu, 1815:20-21, pi. S: fig. 1.—

Fauchald, 1970:64-66.

Marphysa sanguinea var. americana Monro, I933a:68-69,

fig. 28.

MATERIAL EXAMINED.—Balboa, shore pools at low

tide (1, holotype of var. americana, BMNH), coll.

Crossland.

REMARKS.—As indicated by Fauchald (1970:65),

it is not possible to separate the variety described

by Monro from the main form. The species is

highly variable, but can be characterized in relation

to other forms in the area as having exclusively

composite spinigers and branchiae present over a

long region of the body. The stem of the branchiae

is very short, and the branchiae appear palmately

arranged rather than pectinate in most specimens.

Subacicular hooks may be absent completely.

Marphysa sanguinea is circumtropical as far as is

known.

Marphysa species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—This specimen cannot be completely

identified since it is incomplete posteriorly. The

composite falcigers have short, bidentate appendages

in all setigers present; branchiae are present from

setiger 10.

Nematonereis unicornis (Grube, 1840)

Lumbriconereis unicornis Grube, 1840:80.

Nematonereis unicornis (Grube).—Fauvel, 1923:412-413, fig.

162h-n.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—Nematonereis unicornis differs

sharply from all other eunicids found in the present

material in that it has only a single median antenna.

The species is small and easily overlooked, or the

specimens are assumed to be juveniles of larger

species of the family.

Nematonereis unicornis is probably circumtropical but is poorly known for reasons indicated

above.

Palola siciliensis (Grube, 1840)

Eunice siciliensis Grube, 1840:83.—Monro, 1933a:62.

Palola siciliensis (Grube).—Hartman, 1944b: 131.—Fauchald,

1970:68-69.

Nicidion edentulum Ehlers, 1901:130-131, pi. 16: fig. 11-16.

Eunice siciliensis var. edentulum.—Monro, 1933a:62-63.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (4), Galeta Reef, Coralline Zone (12);

Galeta Reef, Laurencia Zone (14); Galeta Reef,

Thalassia Zone (2); Paitilla Beach, Hydroid Zone

(3); Paitilla Beach, Tetraclita Zone (1). Coiba

Island, dredging off convict settlement, smooth

bottom with branched Lithothamnion, 5 fm (5),

and in 9 fm same location (1); Col6n, coral flat at

southwest corner of Lim6n Bay (1); Gorgona Island,

from coral (2); Taboga Island, shore (1), and

obtained by breaking up beach sandstone (1); all

coll. Crossland.

REMARKS.—Most of the specimens are small, but
NUMBER 221 43

individuals from all environments are sexually

mature or nearly so. All specimens that could be

examined in detail were studied for the structure

of the right second maxilla. All specimens had two

distinct teeth and the third denticle, characteristic

of P. paloloides (Moore) (Fauchald, 1970:67-68),

was absent.

Palola siciliensis is circumtropical.

Palola species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone, Galeta Reef, Laurencia Zone; Galeta Reef,

Zoanthus Zone; Paitilla Beach, Hydroid Zone.

REMARKS.—These posterior ends and median

fragments cannot be further identified.

Family LUMBRINERIDAE

Key to the Species from Panama

1. Branched branchiae present on some anterior setigers Ninoe chilensis

Branchiae absent 2

2. All hooded hooks simple S

Anterior hooded hooks composite 4

3. Hooded hooks first present from setigers 21-45 Lumbrkierit erecta

Hooded hooks present for the first setigers Lumbrineris tetrama

4. Parapodial lobes no longer in posterior than in anterior setigers Lumbrineris latreiUi

At least postsetal lobes prolonged in posterior setigers 5

5. Both pre- and postsetal lobes prolonged in posterior setigers Lumbrineris cruzensis}

Only postsetal lobes prolonged Liumbrineris inflate

Lumbrineris cruzensis? Hartraan, 1944 MATERIAL EXAMINED.—Balboa, rocks and rock-

. . . . „ lft... .. . , „ , 1O -. pools at low tide (1), coll. Crossland.

Lumbrmerts cruzensis Hartman, 1944b: 165-166, pi. 12: figs. ~ _ \'. , ,

263-269.-Fauchald. 1970:83-84. pi. I2:g-j. REMARKS.—Lumbrineris erecta has long postsetal

Lumbrineris sphaerocephala.—Uomo. 1933a:86 [not Schmar- lobes in all setigers, but they are distinctly proda, 1861, nor Ehlers, 1905:33]. longed in the posterior parapodia. Lumbrineris

Lumbrineris latreiUi.—Monm. 1933a:84-85 [in part, not Au- heteropoda, on the Other hand, has long postsetal

douin and Milne Edwards. 1834]. lobe s ^ anterio r ^ p^ ^ parapodia, but they

MATERIAL EXAMINED.—Gorgona Island, dredging are reduced to short blunt projections in median

30 fm, muddy sand and shell fragments (1); Gorgona setigers.

Island, dredging 20 fm, fine sand and shell (1); coll. Lumbrineris erecta is found from southern

Crossland. California to Panama in very shallow water or in

REMARKS.—Both specimens have the distinct the intertidal; it has not been reported from the

rounded prostomium and prolonged pre- and Atlantic Ocean,

postsetal lobes characteristic of L. cruzensis. The

jaw apparatus is missing in both, so the identification must be considered somewhat questionable. Lumbrineris inflate Moore, 1911

Neither of the two species to which these specimens . . . ..-.. , ,„,, „« „ , , ,. ^ , tna

... . . , , , , . Lumbrtnens inflate Moore, 1911:289-291, pi. 19, 20: figs. 128-

onginally were assigned have prolonged posterior i34.-Hartman, i944b:l60-i6l.-Fauchald, 1970:89-91. pL

parapodial lobes. I4a-d.

Lumbrineris cruzensis is known from the eastern

Pacific Ocean and has been reported more widely MATERIAL EXAMINED.—Galeta Reef, Laurencia

from other parts of the Americas. Zone (1); Paitilla Beach, Hyroid Zone (3).

REMARKS.—The maxillary formula is the same in

Lumbrineris erecta (Moore, 1904) the present specimens as in those reported by

Fauchald (1970) from western Mexico, in that each

LumbriconereU^ecta Moore. 1904:490-492. pi. 37: figs. 19-22. maxiU a „ , ha s tbn t Q r i(jm teet h an d eac h ^ ^

Lumbrineris' <rr«:/« (Moore).-Fauchald. 1970:85-87. pi. 13a.b. | V tW O teeth '

 Th e acicul a ar e usuall y ^ ^ brow n

Lumbrineris heteropoda.—Monro. i93Sa:82-83 [not Maren-

 i n sma1 1

 specimens and darker, often nearly black,

zeller. 1879]. in large specimens.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lumbrineris inflata is known from British Columbia to western Mexico in the eastern Pacific Ocean.

A closely similar, perhaps identical, form, L.

albifrons Crossland (1924), has been reported from

the Galapagos Islands, and a series of other species

have been considered synonyms of L. inflata from

time to time (Fauchald. 1970). It is thus very

difficult to assess the total distribution of the

species, but at a minimum it is present both in the

Atlantic Ocean and the Pacific Ocean as indicated

by the above samples.

shells, red weed (1); Gorgona Island, shore at low

tide (1); coll. Crossland.

REMARKS.—The present specimens lack anterior

composite hooks. Lumbrineris sphaerocephala has

such hooks. The parapodial construction and the

setal structures agree with L. tetraura. The jaw

apparatus is missing in the specimen from Coibo

Island; the identity is thus open to some doubt.

Lumbrineris tetraura appears widespread in

warm waters, both in the Atlantic and Pacific

Oceans.

Lumbrineris latreilli Audouin and

Milne Edwards, 1834

Lumbrineris latreilli Audouin and Milne Edwards, 1834:168-

170, pi. 3b: figs. 13-15.—Monro, 1933a:84-«5 [in part].—

Fauchald, 1970:94-97, pi. 15f-h.

MATERIAL EXAMINED.—Gorgona Island, dredging

30 fm, muddy sand with shell fragments (1) and

dredging close to shore in 15 fm, shells dead coral

and gravel (1), coll. Crossland.

REMARKS.—Lumbrineris latreilli is difficult to

identify because the limits of variability in the

species are poorly defined. The present specimens

fit well with the species as identified by Fauchald

(1970) from western Mexico.

Lumbrineris latreilli appears to be widely distributed, but as remarked by Fauchald (1970), some of

the records may concern related species.

Lumbrineris tetraura (Schmarda, 1861)

Notocirrus tetraurus Schmarda, 1861:117.

Lumbrineris tetraura (Schmarda).—Hartman, 1944b: 147-149,

pi. 8: figs. 175, 190-191. pi. 9: figs. 192-195.—Fauchald,

1970:109-111, pi. 19b-e.

Lumbrineris brevicirra.—Monro, 1933a:83-84 [not Schmarda,

1861].

MATERIAL EXAMINED.—Coiba Island, dredging off

convict settlement in 10-12 fm, fine sand, small

Lumbrineris species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimen has been dried

out, and all setae are broken off.

Ninoe chilensis Kinberg, 1857

Ninoe chilensis Kinberg, 1857:566.—Monro, 1933a:89-91, fig.

36.

MATERIAL EXAMINED.—Gorgona Island, dredging

in 20 fm, fine sand and shell fragments (3), coll.

Crossland.

REMARKS.—The specimens are as described by

Monro. Hooded hooks are present at least from

setigers 10-12, but anterior setae have been broken

in all specimens and they may have been present

even further anteriorly.

Ninoe foliosa and N. longibranchia were described by Fauchald (1972:153-160, pi. 29-32) and

discussed in relation to other known species with

branched branchiae. Ninoe chilensis differs from

these species in that it lacks the free pectinate part

of the branchiae projecting from the parapodia; N.

foliosa and related species have such a structure.

Ninoe chilensis has only been reported from the

eastern Pacific Ocean.

Family ARABELLIDAE

Key to the Species from Panama

1. At least median and posterior parapodia with stout emergent spines 2

Emergent spines absent Arabella mutans

2. Maxilla I proximally dentate 3

Maxilla I proximally smooth Drilonereis nuda

3. Mandibles present Drilonereis falcata

Mandibles absent DrUonereis mexicana
NUMBER 221 45

Arabella mutatis (Chamberlin, 1919)

Cenothrix mutans Chamberlin, 1919:330-332, pi. 61: figs. 1-9,

pi. 62: fig. 1.

Arabella mutatis (Chamberlin).—Monro, 1933a:88-89 [in

part].—Hartman, 1944b:173-174.—Fauchald, 1970:128-130,

pi. 21a-f.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1); Galeta Reef, Thalassia Zone (2); Paitilla

Beach, Hydroid Zone (17). Taboguilla, sandy shore

at low tide, coll. Mortensen, (fragment); Gorgona

Island, coral (1); Taboga Island, washings from very

barren-looking branches of dead coral 1-2 fm (11);

coll. Crossland.

REMARKS.—Arabella mutans has the ventralmost

seta distally bluntly pointed and hooded with a

long, pointed hood in median and posterior setigers.

The maxillary formula for the present specimens

is 9(10)+ 9(10)- 13(14)+ 13(14)- 5 + 5 - 4(5) + 4(5) -

1 + 1 which agrees well with the formula given by

Fauchald (1970) for material from western Mexico.

The relationship between A. mutans and similar

species was discussed by Fauchald (1970). Monro's

concept of a widely distributed, variable species is

not acceptable without more evidence. However, all

material reexamined from his collections belong to

this species.

Arabella mutans is known from the eastern Pacific

Ocean and is now recorded also from the Atlantic

side of the Isthmus of Panama.

Arabella species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (2). Paitilla Beach, Hydroid Zone (1).

REMARKS.—These specimens are poorly preserved.

Drilonereis falcata Moore, 1911

Drilonereis falcata Moore, 1911:298-299, pi. 20: figs. 150-

154.—Fauchald, 1970:135-136, pi. 21g.

Drilonereis filum.—Momo, 1933a:88 [not Claparcde, 1868].

MATERIAL EXAMINED.—Gorgona Island, dredging

close to shore in 15 fm, shell, dead coral gravel (1),

coll. Crossland.

REMARKS.—The present specimen fits with D.

falcata rather than D. filum as these species were

defined by Fauchald (1970). It has dentate bases on

maxillae I, and mandibles are present. It is known

from central California to Panama in the eastern

Pacific Ocean.

Drilonereis mexicana Fauchald, 1970

Drilonereis mexicana Fauchald, 1970:138-140, pi. 23a-c.

Drilonereis nuda.—Monro, 1933a:86-87, fig. 35 [in part, not

Moore, 1909].

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (1), coll. Crossland.

REMARKS.—Drilonereis nuda as identified by

Monro contains material of two species, D. nuda

sensu Moore and D. mexicana. Drilonereis mexicana

has the proximal part of the first maxillae dentate,

and mandibles are absent. Drilonereis nuda lacks

the dentition of the first maxillae.

Drilonereis mexicana is known from western

Mexico and Panama.

Drilonereis nuda Moore, 1909

Drilonereis nuda Moore, 1909a:254-256, pi. 8: figs. 21-23.—

Monro, 1933a:86-87 [in part].—Fauchald, 1970:140-141, pi.

22g.

MATERIAL EXAMINED.—Coiba Island, in sand at

low tide, fine seagrass and dead coral bedded in

sand (2); Gorgona Island, dredging close to shore

15 fm, shells, dead coral and gravel (1); coll.

Crossland.

REMARKS.—The relation between this and similar

species was reviewed by Fauchald (1970); it can be

distinguished from other species in the area as

indicated above. Drilonereis nuda is known from

the eastern Pacific Ocean between California and

Panama.

Family LYSARETIDAE

Oenone fulgida (Savigny, 1818)

Aglaura fulgida Savigny, 1818:326.

Oenone fulgida (Savigny).—Monro, 1933a:91-92.—Fauchald,

1970:143-146, pi. 24a-d.

MATERIAL EXAMINED.—Galeta Reef, Acanthophora Zone (1); Galeta Reef, Coralline Zone (1).

Coiba Island, dredging off the convict settlement,

5 fm, smooth bottom with branched Lithothamnion

(1) and in crevices of Lithothamnia and coral, 5-10

fm (3); coll. Crossland.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—The present specimens have the single

peristomial segment characteristic of this genus, and

the jaw-apparatus and setal structures are similar

to those seen in specimens from western Mexico.

Oenone fulgida is circumtropical in distribution.

Lysaretidae genus and species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (fragment)

REMARKS.—The fragment is of some median

segments and cannot be identified.

2.

Family DORVILLEIDAE

Key to the Species from Panama

Furcate setae present Schistomeringos longicomis

Furcate setae absent 2

Antennae smooth or wrinkled Dorvillea rubrovittatus

Antennae distinctly articulated Dorvillea cerasina

Dorvillea cerasina (Ehlers, 1901)

Staurocephalus cerasinus Ehlers, 1901:263-264.

Dorvillea cerasina (Ehlers).—Hartman, 1944b: 190.—Fauchald,

1970:155-156, pi. 26a-l.

Staurocephalus angolanus.—Monro, 1933a:93 [not Augener,

1918].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1). Colon, scrapings from piles of quays (4),

coll. Crossland.

REMARKS.—The present specimens have the

distinctly articulated antennae of D. cerasina as in

the material from western Mexico (Fauchald, 1970).

The species was originally described from Chile

and has been found commonly in shallow warm

water areas in the eastern Pacific Ocean as well as

from the western Atlantic.

Dorvillea rubrovittatus (Grube, 1855)

Staurocephalus rubrovittatus Grube, 1855:97-98.—Fauvel,

1923:445-446, fig. 177a-l.

Dorvillea rubrovittata (Grube).—Hartman, 1944b: 190.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (3).

REMARKS.—Dorvillea rubrovittatus was retained

as genotype of Dorvillea as restricted by Pettibone

(1961); it lacks furcate setae, and the articulation

of palps and antennae is absent.

Dorvillea rubrovittatus is known from the warmer

parts of the Atlantic Ocean but has never been

found in the Pacific Ocean.

Schistomeringos longicomis (Ehlers, 1901)

Stauronereis longicomis Ehlers, 1901:150-151. pi. 19: figs.

18-21. pi. 20: figs. 4-6.

Schistomeringos longicomis (Ehlers).—Jumars, 1974:107-109,

fig. 2.

Staurocephalus rudolphii.—Monro, 1933a:92-93 [not delle

Chiaje, 1828:176].

Dorvillea rudolphi (delle Chiaje).—Fauchald, 1970:156-159,

pi. 27a-j.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1). Taboga Island, low tide (1), coll. Crossland.

REMARKS.—Schistomeringos longicomis has recently been separated from the similar S. rudolphi

by Jumars (1974:104-109), who also demonstrated

that the generic names used for these species were

invalid and had to be replaced by Schistomeringos.

Schistomeringos rudolphi is known from the Atlantic Ocean, but the material examined here, including the specimens reported by Monro (1933a:92)

belongs to 5. longicomis. Schistomeringos longicomis is known from the eastern Pacific Ocean

from Chile to British Columbia. The record from

Galeta is the first from the Atlantic Ocean.

Family ORBINIIDAE

Haploscoloplos panamensis Monro, 1933

Haploscoloplos panamensis Monro, 1933b: 1045-1046, fig. 1.—

Hartman, 1957:277.

MATERIAL EXAMINED.—Between Taboga and Taboguilla, 6-12 fm, soft mud (3, syntypes, BMNH),

coll. Crossland.

REMARKS.—The species is as described by Monro

and reviewed by Hartman. It is known from

Alaska and from Panama in shallow water.
NUMBER 221 47

Family SPIONIDAE

Key to the Species from Panama

1. With modified setae in setiger 5 2

Without modified setae in setiger 5 4

2. Some modified setae from setiger 5 densely bristle-topped 3

No bristle-topped setae in setiger 5 Boccardia tricuspa

3. Prostomium anteriorly rounded; short notosetae present in first setiger

Boccardia proboscidea

Prostomium anteriorly bifid; notosetae absent from first setiger Boccardia polybranchia

4. Five pairs of branchiae present; hooded hooks in at least some notopodia

—Prionospio heterobranchia texana

At least 30 pairs of branchiae; hooded hooks in neuropodia only Laonice japonica

Boccardia polybranchia (Haswell, 1885)

Polydora (Boccardia) polybranchia Haswell, 1885:275.—Monro,

1933b: 1047.

Boccardia polybranchia (Haswell).—Blake and Woodwick,

1971:39-41.

MATERIAL EXAMINED.—Gorgona, from coral (2)

and dredging close to shore, 15 fm, shells, gravel

and dead coral overgrown with Polyzoa, etc. (2);

Taboga Island, from dead coral in large pieces

just below low water (1); coll. Crossland.

REMARKS.—These specimens agree well with B.

polybranchia as reviewed by Blake and Woodwick

(1971).

The species is widespread, possibly cosmopolitan.

Boccardia proboscidea Hartman, 1940

Boccardia proboscidea Hartman, 1940b:S83-387, fig. la-j.—

Hartman, 1969:95-96, 5 figs, [unnumbered].—Blake and

Woodwick, 1971:40.

MATERIAL EXAMINED.—Paitilla Beach, Balanus

Zone (6).

REMARKS.—The present specimens fit B. proboscidea as reviewed by Blake and Woodwick (1971)

and the description given by Hartman (1969). The

characteristic rounded prostomium and the structure of the modified setae separate it from other

species from this area.

Boccardia proboscidea has been reported from

the eastern Pacific Ocean from Canada to Panama.

Boccardia tricuspa (Hartman, 1939)

Polydora tricuspa Hartman, 1939b: 16-17, fig. 3c-k.

Boccardia tricuspa (Hartman).—Hartman, 1969:99-100, 4

figs, [unnumbered].—Blake and Woodwick, 1971:40.

MATERIAL EXAMINED.—Paitilla Beach, Tetraclita

Zone (2).

REMARKS.—Boccardia tricuspa differs from B.

proboscidea in the structure of the modified setae

in setiger 5. Both species have two kinds of modified setae, including in both cases simple falcate

spines. Boccardia tricuspa has in addition a series

of distally smooth, tripartite modified setae; B.

proboscidea has bifid, distally strongly hirsute setae

in the same position.

Boccardia tricuspa is known from central California to the Galapagos Islands in shallow water

and from the intertidal.

Laonice japonica (Moore, 1907)

Spionides japonicus Moore, 1907:204-206, pi. 16: figs. 31-34.

Laonice japonica (Moore).—Monro, 1933b: 1047.

MATERIAL EXAMINED.—Perlas Islands (1), coll.

Crossland.

REMARKS.—The species is here accepted as reviewed by Monro (1933b). It is very difficult to

separate the species of Laonice; thus the conservative approach of not introducing another name

for the same material would have to apply in this

instance.

Laonice japonica has been reported from Japan

in addition to this record.

Prionospio heterobranchia texana Hartman, 1951

Prionospio heterobranchia texana Hartman, 1951:85.

MATERIAL EXAMINED.—Galeta Reef Thalassia

Zone (1).

REMARKS.—Foster (1971:93) synonymized the
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

two described subspecies of P. heterobranchia

Moore (1907), P. heterobranchia texana Hartman,

and P. heterobranchia newportensis Reish (1959),

with the main form on the grounds that the ranges

of the first occurrences of the hooded hooks overlapped within one sample as much as the difference

between the subspecies. Type material was examined at the time.

I am unwilling to accept the fusion at the present

time since no reference was made to the size of

the specimens examined. Even when an absolute

overlap in characters can be found in other polychaetes, a reference to the size of the specimens

may readily separate the different forms. For conservative reasons, I prefer at this time to retain

the two subspecies as distinct.

Prionospio heterobranchia texana is known from

the western Atlantic Ocean.

Prionospio species indeterminate

Prionospio pinnata (Ehlers).—Monro, 1933b: 1047.

MATERIAL EXAMINED.—Between Taboga and Taboguilla Islands, 6-12 fm, soft mud (2), coll. Crossland.

REMARKS.—The specimens are poorly preserved,

however, the last pair of branchiae is distinctly digitate, not pinnate as it should be in Paraprionospio

pinnata (Ehlers).

Family MAGELONIDAE

Key to the Species from Panama

Hooded hooks of posterior region bidentate, prostomium with frontal horns

Magelona pacifica

Hooded hooks of posterior region tridentate, prostomium without frontal horns

Magelona papillicornis

Magelona pacifica Monro, 1933

Magelona pacifica Monro, 1933b: 1048-1049. fig. 2.—Jones,

1963:22-23.

MATERIAL EXAMINED.—Gorgona Island at low

tide (14, syntypes, BMNH), coll. Crossland.

REMARKS.—The magelonids are presently under

revision by Dr. Meredith L. Jones, Smithsonian Institution. The identification of the two species reported from Panama has been verified using the

key published by Jones (1963), but otherwise the

conservative view has been taken, which is not to

change or introduce any new names into the lists

until better evidence is available.

Magelona pacifica can be separated from M. papillicornis, also reported from this area, as indicated above. It has been reported from southern

California to Panama.

Magelona papillicornis M tiller, 1858

Magelona papillicornis F. Muller, 1858:216.—Monro, 1933b:

1048.—Jones, 1963:22-23.

MATERIAL EXAMINED.—Balboa, rock and rockpools at low tide (1), coll. Crossland.

REMARKS.—Magelona papillicornis can be separated from M. pacifica as indicated above. This

species is possibly widely distributed, but may have

been confused with other, similar species.

Family CHAETOPTERIDAE

Key to the Species from Panama

1. Tube U-shaped, thick-walled and pergamentaceous Chaetopterus variopedatus

Tube straight, thin-walled, usually translucent brown 2

2. Notopodia of median setigers simple Mesochaetopterus alipes

Notopodia of median setigers with at least 2 lobes 3

3. Paired palps with a pair of small tentacles at the base PhyUochaetopterus sp. indet.

Paired palps without tentacles at the base Spiochaetopterus costarum
NUMBER 221 49

Chaetopterus variopedatus (Renier, 1804)

Tricoelia variopedata Renier, 1804:xviii.

Chaetopterus variopedatus (Renier).—Monro, 1933b: 1050.—

Hartman, 1969:209-210, 3 figs, [unnumbered].

MATERIAL EXAMINED.—Gorgona Island, from

coral (1) and dredging close to shore in 15 fm, dead

coral and gravel (tubes); Taboga Island, at low

water spring (tubes), coll. Crossland.

REMARKS.—Chaetopterus variopedatus can be

found both in hard and soft substrates: in the

former, it glues its tube to such objects as kelp

holdfasts, branching corals etc.; in the latter, it

forms the characteristic symmetrically U-shaped

tubes. The tube is open at both ends at all times,

and functionally it does not appear to matter

whether the tube is attached to rocks, etc., or buried

in sediment.

Chaetopterus variopedatus is cosmopolitan.

Mesochaetopterus alipes Monro, 1928

Mesochaetopterus taylori var. alipes Monro, 1928a:92-93, figs.

14-15.

Mesochaetopterus alipes Monro.—Monro, 1933b: 1050-1052,

fig. 3.

MATERIAL EXAMINED.—Taboguilla, sandy shore

at low tide (syntypes, BMNH), coll. Mortensen;

Taboga Island, shore (12); Gorgona Island, at low

water (21); coll. Crossland.

REMARKS.—The species is as described by Monro.

The tubes are thin-walled and are covered externally with sand-grains, resembling tubes of certain onuphids. Monro (1928a: 92) also reported

Mesochaetopterus minutus Potts from Taboguilla

Island. This material has not been reexamined and

the record cannot be confirmed.

Phyllochaetopterus species indeterminate

Phyllochaetopterus sp.—Monro, 1933b: 1050.

MATERIAL EXAMINED.—Taboga Island at low

tide, sand (fragments), coll. Crossland.

REMARKS.—Sufficient parts of the bodies are present so it is possible to identify the present specimens as belonging to either Spiochaetopterus or

Phyllochaetopterus. They are cited as originally

identified by Monro, but have been excluded from

consideration in the faunistic part.

Spiochaetopterus costarum (Claparede, 1870)

Telepsavus costarum Claparede, 1870:80-84, pi. 20: fig. 1.—

Monro, 1933b: 1052-1054, fig. 4.—Hartman, 1969:219-220,

6 figs, [unnumbered].

MATERIAL EXAMINED.—Gorgona Island at low

tide (3), coll. Crossland.

REMARKS.—As pointed out by Gitay (1969:10),

it is impossible to retain the separation between

Telepsavus and Spiochaetopterus. The specimens

are as described by Monro (1933b). The species

is very widespread, possibly cosmopolitan.

Family CIRRATULIDAE

Key to the Species from Panama

l.

2.

A pair of long, grooved palps present 2

Without grooved palps 5

All setae slender capillaries Tharyx sp. indet.

Setae in part acicular spines or hooks 3

3. Acicular spines distally excavate, body dark Dodecaceria concharum

Acicular spines not excavate, body light 4

4. Posterior acicular spines distally entire Chaetoxone sp. indet.

Posterior acicular spines distally bifid Cauleriella sp. indet.

5. Several segments with transverse rows of dorsal tentacles Timarete perbranchiata

Only 1 segment with transverse row of tentacles 6

6. Branchiae present anterior to the segment with the dorsal tentacles 7

Branchiae present from the segment that carries the dorsal tentacles Cirratulus cirratus?

7. Posterior spines dark or black Cirriformia luxuriosa

Posterior spines light yellow 8

8. Spines present from setigers 7-12; dorsum with irregular pattern of dark transverse bars ...

Cirriformia punctata

Spines first present from median segments, dorsum without color patterns

Cirriformia tentaculata
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cauleriella species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimen is incomplete

posteriorly with 27 setigers. It has bifid neuropodial

hooks from setiger 20; notopodial hooks are not

present in any segment. It does not fit in terms

of the distribution of hooks with any described

species in the genus.

Chaetozone species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimen resembles C.

multioculata Hartman, 1961 (Hartman, 1969:239-

240, 2 figs, [unnumbered]) in that it has a series

of eyespots in a row on each side of the prostomium. Hooks are present in the neuropodia from

setiger 7, and notopodial hooks are present from

setiger 8 in the present specimen. Chaetozone

multioculata has neuropodial hooks from setigers

10-13 and notopodial ones from a median setiger

(approximately setiger 20). The present specimen

is about the same size as the ones described by

Hartman, so the difference in distribution does

not appear related to size.

Cirratulus cirratus? (Muller, 1776)

Lumbricus cirratus O. F. Muller, 1776:281, fig. 5.

Cirratulus cirratus}.—Fauchald, 1973:25-26.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (3).

REMARKS.—These specimens, like the ones previously reported from sandy beaches in Panama,

have neuropodial spines first present from setigers

12-14 rather than from setigers 6-11 as in C. cirratus from other areas.

Cirratulus cirratus is considered cosmopolitan in

shallow waters.

Cirriformia luxuriosa (Moore, 1904)

Cirratulus luxuriosus Moore, 1904:493-494, pi. 38: figs. 28-31.

Cirriformia luxuriosa (Moore).—Hartman. 1969:251-252, 2

figs, [unnumbered].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (15); Paitilla Beach, Tetraclita Zone (9).

REMARKS.—Cirriformia luxuriosa has a single,

curved dark spine in each ramus in each of the

median and posterior parapodia in most specimens;

two or three spines are present in each ramus in a

few cases.

The species is known from the eastern Pacific

Ocean from central California to Panama.

Cirriformia punctata (Grube, 1859)

Cirrhatulus punctata Grube, 1859:107.

Cirratulus punctata Crube.—Augener, 1918:465-467.

Cirriformia punctata (Grube).—Hartman, 1956:292.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (34).

REMARKS.—The species is here accepted as revised by Augener (1918) based on material from

Atlantic areas off Africa. Single, gently curved,

yellow hooks are present from setigers 7-10 in the

neuropodia and from setigers 9-12 in the notopodia. Hooks are always present in two more neuropodial than notopodial setigers. The dark transverse bars are characteristic of this species in the

local area.

Cirriformia punctata has been reported from

the tropical Atlantic Ocean.

Cirriformia tentaculata (Montagu, 1808)

Terebella tentaculata Montagu, 1808:110-111, pi. 6: fig. 2.

Audouinia tentaculata (Montagu).—Fauvel, 1927:91-92, fig.

32a-g.

Audouinia polytricha.—Monro, 1933b: 1054 [not Schmarda,

1861].

Audouinia luxuriosa.—Monro, 1933b: 1055 [not Moore, 1904].

MATERIAL EXAMINED.—Paitilla Beach, Tetraclita

Zone (2). Balboa, rock and rock-pools at low tide

(28); Balboa Docks (2); coll. Crossland.

REMARKS.—Cirriformia tentaculata closely resembles C. spirabranchia (Moore, 1904) from California and C. filigera (delle Chiaje, 1828) from

European waters in that all three species have yellow spines starting from a median setiger. Cirriformia spirabranchia has at least five spines in each

ramus in posterior setigers; the other two have

only three. Fauvel (1927:91) separated the two species occurring in European waters on the basis of

the position of the branchiae in relation to die dis-
NUMBER 221 51

tance between the rami in the parapodia. The

branchiae are closer to the notopodia than the

distance between the noto- and neuropodia in C.

tentaculata and at a distance equal to or greater

than the distance between the rami in C. fdigera.

The present specimens agree with C. tentaculata.

Cirriformia tentaculata may be widespread in

warm waters; but due to the close morphological

similarities between the several species of Cirriformia, it is difficult to estimate its total distribution.

Dodecaceria concharum Orsted, 1943

Dodecaceria concharum 0rsted, 1843b:44.—Fauvel, 1927:102-

103, fig. 36a-n.—Hartman, 1969:255-256, 2 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (3).

REMARKS.—The present specimens, all of which

were solitary burrowers in calcareous substrate, fit

very well with the species as described from Europe.

Dodecaceria concharum is cosmopolitan in shallow rocky areas.

Theoryx species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef, Laurencia Zone (1); Paitilla

Beach, Hydroid Zone (1); Paitilla Beach, Tetraclita

Zone (1).

REMARKS.—Theses specimens are incomplete and

cannot be further identified.

Timarete perbranchiata (Chamberlin, 1918)

Ambo perbranchiata Chamberlin, 1918:117-178.

Timarete perbranchiata (Chamberlin).—Hartman, 1969:269

270.

Ambo amerieana Monro, 1933b: 1051-1056.

MATERIAL EXAMINED.—Gorgona Island, 30 fm,

muddy sand (1, syntype of Ambo amerieana,

BMNH), coll. Crossland.

REMARKS.—The present specimen agrees well

with specimens of T. perbranchiata from California.

My unpublished records extend the distribution of

this species from California through western Mexico to Panama and possibly to the Galapagos

Islands.

Cirratulidae species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (fragment); Paitilla Beach, Tetraclita Zone

(fragments).

REMARKS.—These fragments are unidentifiable.

Family FLABELLJGERIDAE

Key to the Species from Panama

1. Neurosetae pseudocomposite; body encased in a mucoid sheath Flabelligera injundibularh

Neurosetae simple; body not encased in a mucoid sheath 2

2. A prolonged, tongue-like branchial membrane present Piromis amerieana

Branchial membrane short and semicircular Pherusa inflate

Flabelligera infundibularis Johnson, 1901

Flabelligera infundibularis Johnson, 1901:417, pi. 12: figs.

124-127.—Hartman, 1969:291-292, 5 figs, [unnumbered].

Flabelligera affinisi—Momo, 1933b: 1056-1057, fig. 5 [not Sars,

1829].

MATERIAL EXAMINED.—Balboa, pools at the lowest tide level (14), coll. Crossland.

REMARKS.—These specimens have the pseudocomposite hooks of F. infundibularis rather than

the fully composite hooks of F. affinis.

Flabelligera infundibularis is known from Alaska

to Panama in shallow water.

Pherusa inflata (Treadwell, 1914)

Trophonia inflata Treadwell, 1914:213-214, pi. 12: fig. S3.

Pherusa inflata (Treadwell).—Hartman, 1969:297-298, 5 figs.

[unnumbered].

Stylarioides papillata.—Monro, 1933b: 1058-1059, fig. 7 [not

Johnson, 1901].

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (27); Galeta Reef, Laurencia Zone (3); Paitilla

Beach, Hydroid Zone (20). Taboga Island, from

dead coral just below low tide level (4), coll. Crossland.

REMARKS.—All specimens are rather small, but
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cannot otherwise be separated from specimens from

California, except that the number of papillae

tends to be slightly lower.

Pherusa inflata is known from Gulf of Mexico

and along the coast from Oregon to Panama.

Piromis americana (Monro, 1928)

Stylarioides capensis var. americana Monro, 1928a:96-97, fig.

16; 1933b: 1057-1058, fig. 6.

Piromis americana (Monro).—Hartman, 1969:305-306, 3 figs.

[unnumbered].—Fauchald, 1973:27.

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (4). Balboa, rock and rock-pools at low tide

(4); Balboa, scrapings from buoy at canal entrance

(1); coll. Crossland.

REMARKS.—The striations of the neurosetae are

completely indistinct in some of the present specimens; otherwise, they are very close to the original

specimens described by Monro. The species is

known from the eastern Pacific Ocean between

Panama and central California.

Family OPHELIIDAE

Ammotrypane species indeterminate

Ammotrypane aulogaster.—Monro, 1933b:1059 [not Rathke,

1843].

MATERIAL EXAMINED.—Gorgona Island, 25 fm

(1), coll. Crossland.

REMARKS.—The present specimen, which is the

one used by Monro (1933b: 1059), cannot belong

to A. aulogaster in that it has prolonged anterior

setae and the anal funnel is complete and equipped

with long anal cirri. It is poorly preserved.

Family CAPITELLIDAE

Key to the Species from Panama

Thorax with 11 setigers with capillary setae Notomastus (Clistomastus) lineatus

Thorax with 13 setigers with capillary setae Dasybranchus lumbricoides

Dasybranchus lumbricoides Grube, 1878

Dasybranchus lumbricoides Grube, 1878:190-191, pi. 10: fig.

4.—Hartman, 1969:373-374, 3 figs, [unnumbered].

Dasybranchus caducus var. lumbricoides Grube.—Monro,

1933b: 1059-1060, fig. 8.

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (3). Gorgona Island, at low water (18), coll.

Crossland.

REMARKS.—Dasybranchus lumbricoides has retractable branchiae which could be identified only

in a few of the present specimens. The hooks resemble those illustrated for D. lumbricoides (Hartman, 1969:373, figs. 2,3) rather than those described for D. glabrus Moore (Hartman: 1969:371,

figs. 1,2).

Dasybranchus lumbricoides was originally described from the Philippine Islands but has since

been reported extensively from the eastern part of

the Pacific Ocean.

Notomastus (Clistomastus) lineatus Claparede, 1870

Notomastus (Clistomastus) lineatus Claparede, 1870:18-20, pi.

17: fig. 4.—Hartman, 1969:395-396, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (2).

REMARKS.—The present specimens have the vesicular superior organs of the abdominal neuropodia somewhat smaller than as illustrated by

Hartman (1969:395, fig. 2); they are otherwise very

similar. The species is known from world-wide

areas and is mostly common in sandy substrates.

Notomastus species indeterminate

MATERIAL EXAMINED.—Taboga Island, washing

from very barren-looking branches of dead coral

(1), coll. Crossland.

REMARKS.—This specimen was found among

specimens of Arabella mutans identified by Monro

(1928a). It is not sufficiently complete to be identifiable as to species.
NUMBER 221 53

Family MALDANIDAE

Key to the Species from Panama

Anus terminal, cephalic plaque with an evenly low rim on all sides Clymenella sp. indet.

Anus dorsal, cephalic plaque with posterior rim much higher than the anterior one

Maldane gorgonensis

Clymenella species indeterminate

Clymene (Euclymene) rubrocincta.—Monro, 1933b: 1062. fig. 10

[not Johnson, 1901].

MATERIAL EXAMINED.—Coiba Island, 19-12 fm,

fine sand with small shells and red weed (1), coll.

Crossland.

REMARKS.—As remarked by Monro, the specimen

is in two pieces. The two pieces may not belong to

the same species; the anterior fragment resembles

Clymenella in that setigers 4 and 5 have deep collars

encircling the segments; such collars are absent

in Axiothella rubrocincta. The posterior fragment

is as described by Monro and could belong to either

a species of Euclymene or Axiothella.

Maldane gorgonensis Monro, 1933

Maldane gorgonensis Monro. 1933b: 1060-1061, fig. 9.

MATERIAL EXAMINED.—Gorgona Island, 30 fm,

muddy sand and shell fragments (1, holotype,

BMNH), coll. Crossland.

REMARKS.—The specimen is as described by

Monro. The anal plaque has vaguely indicated,

subdistal internal grooves that fail to reach the

margin except ventrally where a series of vaguely

indicated notches can be seen.

Maldane gorgonensis is known only through its

original record.

Maldanidae genus and species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (anterior fragment).

REMARKS.—Without a complete specimen it is

impossible to identify maldanids even as to genus

in an area as poorly known as the Gulf of Panama.

Family OWENIIDAE

Owenia collaris Hartman, 1955

Owenia fusiformis collaris Hartman, 1955:46, pi. 2: figs. 6, 7.

Owenia collaris Hartman.—Hartman, 1969:493-494, 4 figs,

[unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (1).

REMARKS.—Owenia collaris differs from O. fusiformis delle Chiaje (1841) in that it has a high

collar covering the base of the tentacular crown,

which is absent in the latter. Owenia collaris is

found from southern California and western Mexico in the eastern Pacific Ocean. The present record is the first from the Atlantic Ocean.

Family SABELLARIIDAE

Key to the Species from Panama

1. Operculum a conspicuous black cone Phragmatopoma attenuata

Operculum of divergent yellow spines 2

2. Opercular spines in 2 series, nuchal hooks present 3

Opercular spines in 3 series, nuchal hooks absent 4

3. Outer opercular spines strongly curved Idanthyrsus petmatus

Outer opercular spines nearly straight Idanthyrsus armatus

4. Middle opercular spines alternating long and short 5

Middle opercular spines all approximately same length 6

5. Inner opercular spines distally serrated SdbeUaria floridensis

Inner opercular spines distally smooth SabeUaria alcocki
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6. Middle opercular spines distally blunty rounded; inner opercular spines distally serrated

Sabellaria moorei

Middle opercular spines distally pointed; inner opercular spines distally smooth

Sabellaria spinulosa

Idanthyrsus armatus Kinberg, 1867

Idanthyrsus armatus Kinberg, 1867:350.—Monro, 1933b: 1066,

fig. 14.

MATERIAL EXAMINED.—Gorgona Island, 20 fm,

fine sand and shell (1), coll. Crossland.

REMARKS.—The specimen is no longer identifiable since all outer opercular spines have been

broken. These spines however, were illustrated by

Monro and are characteristic of this species.

Idanthyrsus armatus is known from shallowwater localities in South and Central America in

the eastern Pacific Ocean.

Idanthyrsus pennatus (Peters, 1854)

Sabellaria (Pallasia) pennata Peters, 1854:613-614.

Idanthyrsus pennatus (Peters).—Monro, 1933b: 1065, fig. 13.—

Hartman, 1944c, pi. 31: fig. 35.

MATERIAL EXAMINED.—Gorgona Island, from

coral (6), and from coral brought in by natives

from about one fathom below low tide (4) and from

low water spring (32); Taboga Island, from floats

at the end of the hotel pier (4); coll. Crossland.

REMARKS.—All the present specimens are identifiable to species. The outer opercular spines are

well illustrated by Hartman (1944c, pi. 31: fig 35).

Idanthyrsus pennatus appears to be circumtropical in distribution.

Phragmatopoma attenuate Hartman, 1944

Phragmatopoma attenuata Hartman, 1944c:352-353, pi. 38:

figs. 90-96, pi. 39: figs. 100, 101.

Sabellaria (Phragmatopoma) virgini.—Monro, 1933b: 1062-

1063, fig. 11 [not Kinberg, 1867].

MATERIAL EXAMINED.—Perlas Islands, St. Elmo

Bay, shore at low tide (numerous), coll. Crossland.

REMARKS.—As indicated by Hartman (1944:353),

this material does not belong to P. virgini, which

lacks the distinct plume present in this species.

The drawing by Monro is slightly misleading since

the outer opercular spines have two distinct teeth

on either side of the distal plumes.

Phragmotopoma attenuata is known from tropical areas in the eastern Pacific Ocean.

Phragmatopoma species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Paitilla Beach, Hydroid Zone (1).

REMARKS.—Both specimens belong to the same

species and resemble P. peruensis Hartman

(1944c:353-354, pi. 37: figs. 84-85, pi. 39: fig. 99, pi.

41: fig. 104) in that the outer opercular spines have

an entire, flattened distal membrane. Both specimens are quite small and have been dried out, so

it was considered best to identify them only to

generic level.

Sabellaria alcocki Gravier, 1906

Sabellaria spinulosa alcocki Gravier, 1906:543.—Hartman,

1944c:339.

Sabellaria alcocki Gravier.—Hartman, 1969:503-504, 5 figs.

[unnumbered].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (28). Balboa (4), coll. Crossland.

REMARKS.—Sabellaria alcocki has frequently been

considered a subspecies of S. spinulosa. The middle

opercular spines are alternating long and short in

the former and are all of the same size in the

latter. The specimens from Balboa were identified

by Monro as S. pectinata var., but were not reported upon in Monro (1933b: 1063).

Sabellaria alcocki is circumtropical.

Sabellaria floridensis Hartman, 1944

Sabellaria floridensis Hartman, 1944c:345-346, pi. 31: figs.

37-41.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Paitilla Beach, Hydroid Zone (5).

REMARKS.—Sabellaria floridensis has the inner

opercular spines distally strongly serrated and the

middle spines alternating long and short. The

present specimens have usually two short for every

long middle spine.

Sabellaria floridensis has been reported from the
NUMBER 221 55

coast of the Gulf of Mexico; the present records

from Panama are from both sides of the Isthmus.

Sabellaria moorei Monro, 1933

Sabellaria pectinala var. moorei Monro, 1933b: 1063-1064,

fig. 12.

Sabellaria moorei Monro.—Hartman, 1944c:339, pi. 30: figs.

24-26.

MATERIAL EXAMINED.—Paitilla Beach, Tetraclita

Zone (1). Balboa, rocks and rock-pools at low tide

(1, holotype, BMNH), coll. Crossland.

REMARKS.—Sabellaria moorei has serrated inner

opercular spines, and the middle ones are short

and distally bluntly rounded. It resembles S. bellis

Hansen (1882) closely, but material from the original locality of Hansen's species will have to be

examined in order to establish the synonymy.

Sabellaria moorei has been reported only from

Panama.

Sabellaria spinulosa Leuckart, 1849

Sabellaria spinulosa Leuckart, 1849:178.—Monro, 1933b: 1064.

—Hartman, 1969:511-512. 5 figs, [unnumbered].

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (many), coll. Crossland.

REMARKS.—The identity of these specimens is

dubious; they are very much smaller than usual

in this species, and the teeth and spines on the

outer opercular spines are very short and thorn-like.

Sabellaria spinulosa is mainly found in the north

Atlantic Ocean but has been reported from San

Francisco Bay.

Sabellariidae species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1).

REMARKS.—The present specimen is in regeneration, and only two spines are present in the operculum. It generally resembles species of Sabellaria.

Family AMPHARETIDAE

I soldo bipinnata, new species

FIGURE \2a-c

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone, 27 March 1972 (paratypes, AHF Poly 1131)

and 20 July 1972 (holotype, USNM 53092).

DESCRIPTION.—The holotype is a complete specimen with 17 thoracic and 55 abdominal setigers; it

is 12 mm long and 1 mm wide without setae. The

anterior part of the body is slightly inflated, and

the abdomen is cylindrical; the pygidium is a thick,

blunt lobe with the anus terminal.

The prostomium (Figure 12a) is trilobed with a

rounded medial lobe and a pair of lateral folds; this

lobation is most distinct when the tentacles are

everted. The peristomium forms a collar ventrally

and is dorsally produced into a pair of low lobes.

The first few segments are completely fused dorsally

and partially fused ventrally. Four segments (Figure 12&) are included in this fused area. The first

has neuropodial needle setae only; the second and

third have notopodial capillary setae also. In addition the large hooks are found dorsally at the level

of the third fused segment. The transverse membrane is low and smooth. Thoracic notopodia are

low and blunt; neuropodia are low ridges. Abdominal neuropodia are truncate, flattened lobes with

the uncini arranged in a row at the apex. Abdominal notopodia are absent.

Eight branchiae are present, fused into two

groups of four with the two groups emerging

from one stem. The branchiae are of two kinds;

the two lateral branchiae in each of the two groups

are long and smooth; the two medial branchiae

are distinctly bipinnate with long, cylindrical

pinnae arranged in pairs on each branchial stem.

The uncini are in single rows. Thoracic and

abdominal uncini (Figure 12c) are similar in size

and shape; each has six teeth arranged in a single

row and increasing evenly in size from the apical

to the basal tooth; the inner margin of each uncinus

has a blunt tip. Tubes were absent.

Isolda contains 5 species, of which 4 have one

pair of bipinnate and 3 pairs of smooth branchiae.

The 5th species, /. whydahaensis Augener (1918:

514), has numerous fine pinnae on the central pair

of branchiae. The 2 groups of branchiae are partially fused in 3 species and widely separated in /.

sibogae Caullery (1944:102-103). /. bipinnata differs

from the other two (/. pulchella Muller (1858:219)

and /. wambroensis Augener (1914:82-87) in distribution of notosetae in the anterior end. Isolda

bipinnata has notosetae first present from the 2nd

setiger; the other two have them first present on the

3rd setiger. Separation of /. pulchella and /. wambroensis appears problematical.
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Family TEREBELLIDAE

Key to the Species from Panama

1. Thoracic uncini in single rows 2

Thoracic uncini in double rows in median and posterior thoracic setigers 5

2. Branchiae absent Polycirrus sp. indet.

Branchiae present 3

3. Two pairs of digitiform branchiae present Euthelepus pascua, new species

Two or 3 pairs of multifid branchiae 4

4. Setae present from first branchial segment; uncini with rectangular bases

Streblosoma crassibranchia

Setae present from second branchial segment; uncini with oval bases Thelepus setosus

5. Uncini pectiniform with all teeth in a single row 6

Uncini crested with a large fang surmounted by a few to numerous smaller teeth 7

6. Thoracic uncini with 5 or 6 teeth Loimia medusa

Thoracic uncini with 4 teeth Loimia annulifilis

7. At least anterior thoracic uncini long-handled 8

All uncini short-handled 13

8. Nephridia basally connected Lantcides taboguillae

Nephridia free from one another 9

9. All thoracic uncini long-handled Pista fasciata

Only anterior thoracic uncini long-shafted 10

10. First pair of branchiae smaller than, or of the same size as, the second pair

Pista brevibranchiata

First pair of branchiae distinctly larger than the second pair 11

11. Expanded region of long-shafted uncini below the middle of the length Pista herpini?

Expanded region of long-shafted uncini just below the head of the setae 12

12. Lappets of segment 3 fused middorsally Pista alata

Lappets of segment 3 not fused Pista elongata

13. Seventeen thoracic setigers present 14

Thorax with numerous setigers TerebeUa gorgonae

14. Eyes absent, a single large tooth above the main fang in each uncinus Eupolymnia regnans

Eyes present, at least 2 teeth above the main fang in each uncinus Eupolymnia nebulosa

Eupolymnia nebulosa (Montagu, 1818)

TerebeUa nebulosa Montagu, 1818:343, pi. 12: fig. 2.

Polymnia nebulosa Montagu.—Fauvel, 1927:257-258, fig.

89a-g.—Monro, 1933b: 1072.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (4); Galeta Reef, Thalassia Zone (1). Coiba

Island, dredging off the convict settlement, 5 fm,

smooth bottom with branched Lithothamnion (13);

Taboga Island, shore (6); coll. Crossland.

REMARKS.—The present specimens agree with

E. nebulosa in that the three pairs of branchiae

decrease in size evenly from the first and in that the

lateral lappets are well developed. Fauvel (1927:258)

indicated that the number of teeth in the crests of

the uncini varies from one to five. This variation is

not the case in the present material where all

specimens have five teeth in the crests. The identification mav thus be somewhat doubtful. The specimens from Coiba Island were identified by Monro

but not published in the report on the Panamanian

fauna.

Eupolymnia nebulosa is widespread in shallow

water.

Eupolymnia regnans Chamberlin, 1919

Eupolymnia regnans Chamberlin, 1919:433-434, pi. 79: figs.

1-3.

Polymnia regnans (Chamberlin).—Monro, 1933b: 1072-1073,

fig. 19.

MATERIAL EXAMINED.—Balboa, pools at the lowest

tide level (2), coll. Crossland.

REMARKS.—This species is identifiable as indicated in the key. The uncini are as illustrated by

Monro (1933b, fig. 19). The numbers of species

included in this genus and the identity of several

of them must remain doubtful until more material

has been studied.
NUMBER 221 57

FIGURE 12.—Melinna bipinnata, new species: a, anterior end, dorsal view, X 50; b, anterior

end, lateral view, X 50; c, thoracic uncinus, lateral view, X 950. Euthelepus pascua, new

species: d, anterior end, ventral view, X 50; e, anterior end, lateral view, X 50; /, late thoracic

uncinus, lateral view, X 950; g, late thoracic uncinus, frontal view, X 950; h, anterior end.

dorsal view, X 50. Terebella gorgonae Monro: i, thoracic uncinus, frontal view, X 700.

Eupolymnia regnans appears limited to the

warmer parts of the eastern Pacific Ocean.

Euthelepus pascua, new species

FIGURE I2d-h

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone, 17 March 1972 (1, holotype, USNM 53093).

DESCRIPTION.—The holotype and only known

specimen is incomplete and consists of 21 thoracic

and 8 abdominal setigers. It is 4.5 mm long and

0.75 mm wide without setae and is evenly tancolored without distinct color patterns.

The prostomium (Figure I2d,e,h) forms a hood

over the mouth; the anterior rim is thickened; eyes

are absent. The peristomium is a complete ring

posterior to the mouth; it carries laterally a single
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pair of short, grooved tentacles; other anterior

appendages are absent. The next two segments are

asetigerous; each carries a pair of digitiform

branchiae. The first pair is about one-third as long

as the second pair and somewhat thicker.

The third postperistomial segment is the first

setiger; it has a tuft of sessile notosetae. Distinct

parapodia are absent in the thorax; the abdominal

neuropodia are slightly raised as ridges. Uncini are

present from the fourth setiger; they are in single

rows. Each uncinus (Figure I2f,g) has a very short,

square base and a large main fang surmounted by a

crest of about 22 teeth in 4 distinct tiers. All uncini

are similar.

The genus Euthelepus is presently known for

three species; these include E. malayensis Caullery

(1944:182, fig. 146), the genotype E. setobalensis

Mclntosh (1885:465-467. pi. 50: fig. 4, pi. 28A: fig.

13), and E. tenuis (Verrill, 1900:663). Euthelepus

malayensis has one pair of branchiae and uncini

from the second setiger. Euthelepus tenuis has

similarly one pair of branchiae, and uncini are

first present from setiger 3. Euthelepus pascua has

two pairs of branchiae, and uncini are first present

from the fourth setiger. Finally, E. setobalensis has

three pairs of branchiae, and uncini appear to be

first present from setiger 3.

There are indications that more than one taxon

may be involved in the present concept of the

genus; thus the genotype has notosetae present

from the first branchial segment, and setae are first

present in postbranchial segments in the present

specimen.

Lanicides taboguillae (Chamberlin, 1919)

Nicolea taboguillae Chamberlin, 1919:425-427, pi. 79: figs.

12, IS.

Lanicides taboguillae (Chamberlin).—Monro, 1983b: 1069-

1070, fig. 17.

MATERIAL EXAMINED.—Balboa, shore on lee side

of Taboga Island, south of the village (5); Col6n,

coral flat at SW corner of Lim6n Bay (1); coll.

Crossland.

REMARKS.—The general appearance and most of

the features of species of Lanicides agree with

Pista, from which it can be separated mainly by

anatomical features.

Lanicides taboguillae is known from both sides

of the Isthmus of Panama in shallow water.

Loimia anntdifilis (Grube, 1871)

Terebella annulifilis Grube, 1871:49.

Loimia medusa var. annulifilis (Grube).—Monro, 1933b: 1069.

MATERIAL EXAMINED.—Taboga Island, shore (1).

coll. Crossland.

REMARKS.—Loimia annulifilis differs from L.

medusa in that it has four teeth in the thoracic

uncini rather than five or six as in the latter.

Loimia annulifilis is known from the Indo-Pacific

region.

Loimia medusa (Savigny, 1818)

Terebella medusa Savigny, 1818:85.

Loimia medusa (Savigny).—Monro, 1933b: 1069.—Hartman,

1969:601-602, 2 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1); Galeta Reef, Thalassia Zone (4); Paitilla

Beach, Hydroid Zone (3).

REMARKS.—Loimia medusa has the characteristic

pectiniform uncini in which all the teeth are in a

single row, usually five or six teeth are present.

The specimens differ in no discernible way from

specimens described from other areas. Loimia

medusa is considered cosmopolitan in shallow water.

It is usually found in sandy substrates, but only

very small amounts of sand are necessary; thus

the species has frequently been reported from hardbottom areas.

Pista alata Moore, 1909

Pista alata Moore, 1909a:273-275, pi. 9: figs. 48-51.—Monro,

1933b: 1066-1068, fig. 15.—Hartman, 1969:611-612, 3 figs,

[unnumbered].

MATERIAL EXAMINED.—Balboa, rocks and rockpools at low tide (7); Coiba Island, in sand at low

water (7) and at low tide among volcanic rocks and

boulders (3); coll. Crossland.

REMARKS.—The characteristic dorsally fused lappets on the third segment make P. alata easily

identifiable on the west coast of the Americas.

According to the labels, the specimens from Balboa

were identified by Monro, but they were not

reported on in his study.

Pista alata is known from southern California

and Panama in shallow water.
NUMBER 221 59

Pista brevibranchiata Moore, 1923

Pista brevibranchiata Moore, 1923:196-197.—Hartman, 1969:

613-614, 4 figs, [unnumbered].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (1).

REMARKS.—The present specimen agrees with

P. brevibranchiata in the structures of the uncini,

both anterior and posterior, but differs in that it

has all branchiae of the same size. Pista brevibranchiata has the second pair very much larger

than the first.

Pista brevibranchiata is known from central and

southern California and western Mexico in intertidal and shelf depths.

Pista elongata Moore, 1909

Pista elongata Moore. 1909a:270-272. pi. 9: figs. 45-47.—

Monro. 1933b: 1068.—Hartman, 1969:619-620, 4 figs, [unnumbered].

MATERIAL EXAMINED.—Balboa, rocks and rock

pools at low tide (9), coll. Crossland.

REMARKS.—The specimens are as described by

Hartman (1969). The long-shafted uncinus has a

narrow, pointed boss rather than the rounded

expanded one, which is characteristic of the other

species of Pista from this area.

This species is known from western Canada to

Panama in shallow water.

Pista fasciata (Grube, 1870)

Terebella (Phyzelia) fasciata Grube. 1870a:513-514.

Pista fasciata (Grube).—Hartman, 1969:621-622, 2 figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef, Laurencia Zone (1); Galeta

Reef, Thalassia Zone (1).

REMARKS.—Pista fasciata has long-shafted uncini

in all thoracic setigers. The branchiae are richly

and loosely branched. It has been reported from

world-wide areas, but appears to be most common

in warm waters.

Pista herpini? Fauvel, 1928

Pista herpini Fauvel, 1928:160-162, fig. 2a-h.—Monro, 1933b:

1068, fig. 16.

MATERIAL EXAMINED.—Gorgona Island, at low

water (1), coll. Crossland.

REMARKS.—The present specimen resembles P.

herpini; but, as remarked by Monro, the shafts of

the long-shafted uncini are stouter than in P.

herpini and are in fact considerably stouter than

as illustrated by Monro. The identity of the specimen remains doubtful.

Polycirrus species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef, Laurencia Zone (1).

REMARKS.—The different species of Polycirrus

are so difficult to identify that it seems foolhardy

to attempt to name any specimens to species without a complete revision of the genus; materials for

this revision are not yet available.

Streblosoma crassibranchia Treadwell, 1914

Strcblosoma crassibranchia Treadwell, 1914:208-209, pi. 12:

figs. 30-31.—Hartman, 1969:641-642, 5 figs, [unnumbered].

Streblosoma verrilli.—Monro, 1933b: 1073-1074, fig. 20 [not

Treadwell. 1911 = Thelepus setosus fide Hartman,

1956:297].

MATERIAL EXAMINED.—Galeta Reef, Thalassia

Zone (2); Paitilla Beach, Hydroid Zone (10). Balboa,

rocks and rock-pools at low tide (numerous); Coiba

Island, in sand at low tide (numerous); coll.

Crossland.

REMARKS.—The present specimens fit very well

with 5. crassibranchia. Setae are present from the

first branchial segment. Each uncinus has a large

main fang and three teeth in a transverse row in

the crest.

The species is known from central California to

Panama; it is present on both sides of the Isthmus.

Terebella gorgonae Monro, 1933

FIGURE 12I

Terebella gorgonae Monro. 1933b: 1070-1071. fig. 18.

MATERIAL EXAMINED.—Gorgona Island, shore (10,

syntypes, BMNH), coll. Crosslands.

REMARKS.—Terebella gorgonae closely resembles

T. californica Moore, 1904, (Hartman, 1969:643-

644, 4 figs.), but differs in the distribution and

structure of the branchiae. The branchiae are on

the first and third setiger in addition to the last
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

presetigerous segment in T. gorgonae; that is, they

are missing on the second setigerous segment. The

branchiae are on consecutive segments in T. californica and include only two pairs. The uncini

(Figure 12*) have a large main fang and a doubletiered crest with three and seven teeth in each tier.

Terebella gorgonae has been reported from the

type-locality and from the Galapagos Islands.

Thelepus setosus (Quatrefages, 1865)

Phenacia setosa Quatrefages, 1865:376-377.

Thelepus setosus (Quatrefages).—Hartman, 1969:649-650, 6

figs, [unnumbered].

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1).

REMARKS.—The present specimen has the attachment points of the uncini in a slightly more subterminal position than is illustrated in Hartman

(1969:649); otherwise the specimen fits very well

within the species as described.

Thelepus setosus is cosmopolitan in shallow

water.

Terebellidae species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (dried-out specimens and posterior ends);

Galeta Reef, Thalassia Zone (fragments and posterior ends); Paitilla Beach, Hydroid Zone (fragment); Paitilla Beach, Tetraclita Zone (posterior

ends); Paitilla Beach, 31 Dec. 1970 (tubes only).

REMARKS.—These fragments cannot be further

identified.

Family TRICHOBRANCHIDAE

Terebellides stroemi? Sais, 1835

Terebellides stroemi Sars, 1835:48—Monro, 193Sb:1075.—

Hartman, 1969:653-654, 7 figs, [unnumbered].

MATERIAL EXAMINED.—Gorgona Island, 30 fm,

muddy sand and shells (1), coll. Crossland.

REMARKS.—The neuropodial spines in setiger 6

are nearly straight and sharply pointed in the

present specimen; they are usually distinctly bent

and blunt in this species.

Terebellides stroemi is cosmopolitan, apparently

in all depths.

Family SABELLIDAE

Key to the Species from Panama

1. Thorax with pennoned neurosetae in addition to the uncini 2

Thorax with only uncini in the neuropodia Chorus sp. indet.

2. Radioles spiralled; with their bases rolled in at the dorsal end 3

Radioles not spiralled; each of their bases semicircular 4

3. Some thoracic notosetae scimitar-shaped Bispira tnonroi

All thoracic notosetae limbate Demonax leucaspis?

4. Some radioles with terminal compound eyes Megalomma vesiculosum

Terminal compound eyes absent 5

5. First setiger with a linear or gently curved series of limbate setae Hypsicomus phaeotaenia

First setiger with a tuft of setae 6

6. All thoracic notosetae slender and limbate; eyes in double rows Sabella melanostigma

Thoracic notosetae of two kinds, slender and broad, eyes in single rows 7

7. Collar 2-lobed and projecting anteriorly ventrally in 2 sharp points Potamilla fonticula

Collar 4-lobed; ventrally recurved or rounded 8

8. Dorsal edge of the radiolar bases deeply deft Pseudopotamitta occelata

Dorsal edge of radiolar bases smooth 9

9. At most 2 eyes per radiole, dorsal lappets distally pointed Pseudopotamilla intermedia

Seven or 8 eyes per radiole; dorsal lappets distinctly triangular ... Pseudopotamilla reniformis

Bispira tnonroi (Hartman, 1961)

FIGURE I3a-b

Distylidia tnonroi Hartman, 1961:129.

Bispira rugosa var. monterea.—Monro, 1933b: 1076-1077, fig.

23 [not Distylia [= Distylidia] rugosa Moore, 1904, nor

Distylia monterea Chamberlin, 1919 = Eudistylia polymorpha (Johnson, 1901)].

MATERIAL EXAMINED.—Balboa, pools at the lowest tide (numerous); Coiba Island, in sand at low
NUMBER 221 61

water, fine seagrass and dead coral bedded in the

sand (2); coll. Crossland.

REMARKS.—The present specimens have the

scimitar-shaped setae characteristic of the genus

Bispira. It differs from Bispira volutacornis

(Montagu, 1804) (Hartman, 1969:659-660, 5 figs,

[unnumbered]) in the shape of the lobes of the

collar. The illustrations give the details of the

anterior end (Figure 13a,b).

Bispira monroi is known from Panama only.

FIGURE IS.—Bispira monroi (Hartman): a, anterior end, ventral view, X 5; b, anterior end,

dorsal view, X 5. Potamilla fonticula (Hoagland): c, bilimbate notoseta, lateral view, X 385;

d, anterior end, ventral view, X 50; e, anterior end, lateral view, X 50; /, thoracic mucronate

seta, X 385; g, abdominal mucronate seta, X 385; h, pennoned seta, X 385; i, bilimbate notopodial seta, frontal view, X 385; /, thoracic uncinus, X 385; k, abdominal uncinus, X 385.

Chone species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1).

REMARKS.—The present specimen can be identified only to genus.

Detnonax leticaspisf Kinberg, 1857

Detnonax leucaspis Kinberg, 1867:354.—Monro, 1933b: 1075-

1076, fig. 22.

MATERIAL EXAMINED.—Galeta Reef, Laurencia

Zone (1). Balboa, rocks and rock-pools at low tide
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(7); Balboa from buoy brought in from the canal

entrance (numerous); coll. Crossland.

REMARKS.—The present specimens fit into

Demonax in that they have spiralled tentacular

crowns, limbate notosetae, two-lobed collars and

smooth abdomens. Monro (1933b) indicated the

presence of D. leucaspis in Panama. His concept of

the species appears very wide; thus, he includes

D. medius (Bush, 1904) into his concept. Based on

the shape of the collar, this species is definitely

distinct, but the whole species concept in this

group appears indeterminate. Until a complete

revision of the genus has been done, the name

previously used for this Panamanian material is

retained.

Hypsicomus phaeotenia (Schmarda, 1861)

Sabella phaeotaenia Schmarda, 1861:35, pi. 22: fig. 188.

Hypisicomus phaeotenia (Schmarda).—Fauvel. 1927:312-314,

fig. 108a-l.—Monro, 1933b:1079.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (11); Galeta Reef, Laurencia Zone (1, a tentacular crown); Paitilla Beach, Hydroid Zone (2).

Balboa, scrapings off piles of quays at the docks

(1); Gorgona Island, from coral (1); Taboga Island,

5 fm off the hotel pier (1) and from low water

spring (8); coll. Crossland.

REMARKS.—Hartman (1944a:24) reported H. circumspiciens Ehlers from Venezuela; in addition to

the present species, H. torquatus (Grube) has been

reported from the area. The synonymy appears at

least in part confused (cf. Day, 1973:125). For this

reason, it seems best to retain the name already

used for specimens from this area. All specimens

examined belong to the same species; they agree

well with the description of this species as given by

Fauvel (1927); but as indicated above, the conclusions drawn here may be incorrect and another

name should possibly be applied to the present

collection.

Hypsicomus phaeotaenia is circumtropical in

distribution.

Megalomma vesiculosum (Montagu 1815)

Amphitrite vesiculosa Montagu, 1815:19-20, pi. 5: fig. 1.

Branchiomma vesiculosum (Montagu).—Fauvel. 1927:315. fig.

109a-q.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (7).

REMARKS.—The present specimens agree in all

essential characters with the species as described

from European waters. The species is widely distributed in warm waters.

Potamilla fonticula (Hoagland, 1919)

FIGURE 13O-*

Parasabella fonticula Hoagland, 1919:579, pi. 31: figs. 3-9.

^Potamilla floridana Augener, 1922:48.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (4); Galeta Reef, Laurencia Zone (36).

REMARKS.—Potamilla fonticula differs from most

other species of Potamilla in that it has a high,

somewhat variable number of thoracic setigers. The

number was originally indicated as 20; in the

present material the number varies from 14 to 22,

but appears to be approximately 20 in all larger

specimens. The segment of change between thorax

and abdomen appears strongly related to the size

of the specimen.

The collar (Figure 13d,e) is deeply cleft ventrally

and the two parts are prolonged anteriorly; palps

were not observed, but are presumably hidden

below the tall collaf folds. Dorsally the collar is

completely reduced, and the base of the tentacular

crown is naked. Laterally the base of the tentacular

crown has a dark brown spot.

Thoracic notopodial setae include spatulate,

mucronate setae, and long, slender bilimbate setae

(Figure \$c,f,i). Thoracic neuropodia have a row of

large avicular uncini (Figure 13;) with a very distinct subdistal thickening and a long, straight shaft.

The pennoned setae (Figure I5h) are subdistally

double-curved and thickened; the distal part is

very thin, flimsy, and often difficult to find. Abdominal setigers have small, very slender uncini (Figure

ISA) without the distinct subdistal thickening and

very strongly spatulate, mucronate setae (Figure

13g). Potamilla floridana Augener agrees with the

present species in so far as it has been described.

Potamilla fonticula was described from Puerto

Rico; the present records are from Panama on the

Atlantic side. It may also be present in Florida.

Pseudopotamilla intermedia Moore, 1905

Pseudopotamilla intermedia Moore, 1905:562-564, pi. 37: figs.

15-22.—Hartman, 1969:727-728, 6 figs, [unnumbered].
NUMBER 221 63

Potamilla brevibranchiata.—Monro, 1933b: 1079 [not Moore,

1905].

MATERIAL EXAMINED.—Balboa, rock and rockpools at low tide (3), coll. Crossland.

REMARKS.—The present specimens differ from

P. brevibranchiata in that the tentacular crown is

too tall, the collar is too well developed, and the

uncini in the last three thoracic setigers are twice

as large as those in anterior setigers. These features

are characteristic of Pseudopotamilla intermedia.

This species is known from California and Alaska.

Pseudopotamilla occelata Moore, 1905

Pseudopotamilla occelata Moore, 1905:559-562, pi. 37: figs.

8-14.—Hartman, 1969:729-730, 5 figs, [unnumbered].

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (2); Paitilla Beach, Tetraclita Zone (1).

REMARKS.—The present specimens have the

deeply cleft dorsal margin of the tentacular crown

characteristic of this species. The species is known

from Alaska and California to western Panama.

Pseudopotamilla reniformis (Muller, 1771)

Amphitrite reniformis O. F. Muller, 1771:194.

Potamilla reniformis (Muller).—Fauvel, 1927:309-310, fig.

107a-l.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1).

REMARKS.—Pseudopotamilla reniformis can be

separated from other species in the area as indicated in the key. The species is common in European waters and has been reported from widely

scattered areas.

Sabella melanostigma Schmarda, 1861

Sabella melanostigma Schmarda, 1861:36, pi. 22: fig. 190.—

Jones, 1962:194.

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (1); Galeta Reef (1), collected on plexiglass,

31 January 1971, coll. C. Birkeland; Paitilla Beach,

Hydroid Zone (2).

REMARKS.—This is the only species of the genus

Sabella identifiable in the present material. The

specimens from Paitilla Beach seem to represent

the first record of this species from the Pacific

Ocean.

Sabella species indeterminate

MATERIAL EXAMINED.—Paitilla Beach, Hydroid

Zone (2).

REMARKS.—These are two small specimens without the distinctive color patterns characteristic of

most species of Sabella.

Sabellidae species indeterminate

MATERIAL EXAMINED.—Galeta Reef, Coralline

Zone (fragment); Galeta Reef, Laurencia Zone (posterior fragment).

REMARKS.—These fragments can be identified as

sabellids only on the distribution of setae.

Biogeography

The fauna of the Atlantic and Pacific sides of

Panama represents the split remnants of a previously unified warm water fauna. Ekman (1953) and

Briggs (1974) summarized the information on the

region, based mainly on the records of crustaceans

and fishes. Generally, the Atlantic side is considered to have the richer fauna in terms of numbers of species and the Pacific side is considered as

depauperate with a fauna derived in the main from

the western Atlantic.

A study of the sandy beaches of central America

(Fauchald, 1973) demonstrated that the populations and species numbers were in fact much higher

on the Pacific side than on the Atlantic in terms of

the polychaetes. The same general picture emerges

from the current study (Tables 1, 2) in that 136

of 179 species are represented on the Pacific side and

only 73 on the Atlantic side.

Tables 1 and 2 survey the occurrence of species

of polychaetes on the Pacific and Atlantic sides of

Panama in relation to the total geographical dispersal of each species. For practical reasons, individual dispersals have been reduced to 14 patterns

as defined below.

la. Cosmopolitan species are shallow-water species

reported from both cold and warm water regions in

all three major oceans in suitable localities. Deepwater dispersal beyond the shelf has not been

considered.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Relation between the local occurrence and geographic dispersal of polychaetes

reported from Panama in terms of numbers of species (numbers in parenthesis = percentages

of total numbers of species reported from each coast line)

Pattern Atlantic Pacific Total species

I a. Cosmopolitan

b. Circumtropical

II Amphi-American

a. Extended

b. Warm water only

III One coast only

a. Atlantic Ocean

1. Extended

2. Warm water only

3. Endemic

b. Pacific Ocean

1. North Pacific

2. Extended East Pacific

north

south

3. Indo-Pacific warm

4. Warm water East Pacific

5. Endemic

10 (13.70)

21 (28.77)

2 (2.74)

16 (21.92)

4 (5.48)

13 (17.81)

7 (9.59)

13 (9.56)

28 (20.59)

1 (0.74)

21 (15.44)

Total number of species 73

4 (2.94)

12 (8.82)

5 (3.68)

7 (5.15)

29 (21.32)

16 (11.76)

136

18 (10.06)

35 (19.55)

3 (1.68)

26 (14.53)

4 (253)

13 (7.26)

7 (3.91)

4 (2.23)

12 (6.70)

5 (2.79)

7 (3.91)

29 (16.20)

16 (8.94)

179

TABLE 2.—Geographic dispersal and local occurrences of polychaetes reported from Panama

(WW = limited to warm water; W-C = found in both warm and cold water; dash = no

other record)

Species Panama General dispersal

Aphrodita diplops

A. japonica

Chaetacanthus magnificus ...

Halosydna glabra

H. leucohyba

Harmothoe balboensis

H. hirsuta

Iphione ovata

Lepidasthenia gigas

L. varius

Lepidonotus crosslandi

L. humilis

L. nesophilus

Thormora Johnstoni

T. taeniata

Eupanthalis perlae

Pant halts mortenseni

P. pacifica

Polyodontes oculea

Psammolyce spinosa

Sthenelais fusca

Thalenessa lewisii

Bhawania goodei

B. riveti

Chrysopetalum ocddentale

Atlantic

Pacific

Pacific

Pacific

Atlantic

Pacific

Pacific

Pacific

Pacific

Atlantic

Pacific

Both

Pacific

Pacific

Pacific

Pacific

Pacific

Pacific

Both

Atlantic

Pacific

Pacific

Both

Pacific

Pacific

North Pacific

Circumtropical

WW Eastern Pacific

WW Western Atlantic

WW Eastern Pacific

WW East-Central Pacific

WW Eastern Pacific

WW Americas

WW Eastern Pacific

WW Western Atlantic

WW Eastern Pacific

WW Eastern Pacific

WW Western Atlantic

WW Eastern Pacific

WW Eastern Pacific

WW Americas

WW Eastern Pacific

W-C Eastern Pacific (north)

WW Eastern Pacific

Circumtropical

WW Eastern Pacific

WW Eastern Pacific
NUMBER 221

65

TABLE 2.—Continued

Species Panam

a

Pacific

Pacific

Pacific

Both

Atlantic

Pacific

Atlantic

Pacific

Pacific

Pacific

Atlantic

Pacific

Pacific

Atlantic

Pacific

Pacific

Atlantic

Pacific

Pacific

Atlantic

Atlantic

Pacific

Pacific

Atlantic

Pacific

Both

Both

Atlantic

Pacific

Both

Both

Pacific

Both

Pacific

Both

Pacific

Atlantic

Both

Pacific

Atlantic

Both

Pacific

Atlantic

Pacific

Both

Both

Atlantic

Both

Atlantic

Both

Pacific

Atlantic

Pacific

Pacific

General dispersal

Circum tropical

WW Eastern Pacific

WW Americas

Circumtropical WW Western Atlanti

c

WW Americas WW Atlantic (both sides

)

Pacific Panama only

WW Americas

Circumtropical WW Western Atlanti

c

Circumtropical

Circumtropical WW Western Atlanti

c

WC Eastern Pacific (north)

Pacific Panama only

Northern Pacific

WC Eastern Pacific (south)

WW Indo-Pacific

WW Western Atlantic

WC Western Atlantic (north)

WC Eastern Pacific (north)

Cosmopolitan

Atlantic Panama only

North Pacific

Circumtropical

Cosmopolitan

WW Western Atlantic

Circumtropical

PCircumtropical

Circumtropical WW Indo-Pacifi

c

WC Eastern Pacific (north)

-WW Americas

Cosmopolitan

Circumtropical

Cosmopolitan

Eastern Pacific

WC Western Atlantic (north)

Circumtropical ---

Cosmopolitan

WC Eastern Pacific (south)

-WW Americas WW Western Atlanti

c

Circumtropical (extended)

WW Indo-Pacific

Circumtropical WW Americas

Paleanotus chrysolepis

Chloeia entypa

C. viridis

Eurythoe complanata

Hermodice carunculata

Linopherus ambigua

L. canariensis

L. oculata

Notopygos ornata

Pherecardia striata

Anaitides erythrophyllvs

A. lamellijera

A, madeirensis

Eulalia myriacyclum

Eumida bifoliata

Phyllodoce panamensis

Sige orientalist

Steggoa lobocephalica

Hesione intertexta

H. picta

Ophiodromus obscurus

O. pugettensis

Synelmis albini

Autolytus anoplos

A. cf. magnus

Haplosyllis spongicola

Langerhansia cornuta

L. mexicana

Odontosyllis polycera

Opisthosyllis brunnea

Syllis gracilis

Trypanosyllis (Trypanedenta)

taeniaformis

Typosyllis aciculata

T. caeca

T. fuscosuturata

T. hyalina

T.prolifera}

T. variegata

Ceratocephala crosslandi

Ceratonereis irritabilis

C. mirabilis

Eunereis paitillae

Neanthes galetae

N. pseudonoodti

N. succinea

Nereis callaona

N. panamensis

N. riisei

Perinereis anderssoni

Platynereis dumerilii

Pseudonereis gallapagensis

P. variegata

Aglaophamus dicirris

A. tabogensis
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Continued

Species

Nephtys monroi

N. singularis

N. squamosa

Glycera abranchiata

G. americana

G. oxycephala

G. tesselata

Goniada acicula

Americonuphis reesei

Diopatra chilicnsis

D. cuprea

D. denticulata

D. ornata

Nothria gorgonensis

Onuphis nebulosa

O. vermillionensis

Eunice afra

E. antennata, sensu stricto

E. antennata aedificatrix

E. aphroditois

E. biannulata

E. filamentosa

E. mutilata

E. (Nicidion) cariboea

E. reducta

E. vittatopsis

E. websteri

Lysidice ninetta

Marphysa amadae

M. sanguined

Nematonereis unicornis

Palola siciliensis

Lumbrineris cruzensist

L. erecta

L. inflata

L. latreilli

L. tetraura

Ninoe chilensis

Arabella mutans

Drilonereis falcata

D. mexicana

D. nuda

Oenonc fulgida

Dorvillea cerasina

D. rubrovittatus

Schistomeringos longicornis

Haploscoloplos panamensis

Boccardia polybranchia

B. proboscidea

B. tricuspa

Laonice japonica

Prionospio heterobranchia texana

Magelona pacifica

M. papillicomis

Chaetopterus variopedatus

Panama General dispersal

Pacific

Pacific

Pacific

Pacific

Pacific

Atlantic

Atlantic

Atlantic

Pacific

Pacific

Atlantic

Pacific

Pacific

Pacific

Atlantic

Atlantic

Both

Both

Both

Both

Pacific

Both

Pacific

Both

Pacific

Pacific

Atlantic

Pacific

Atlantic

Pacific

Atlantic

Both

Pacific

Pacific

Both

Pacific

Pacific

Pacific

Both

Pacific

Pacific

Pacific

Both

Both

Atlantic

Both

Pacific

Pacific

Pacific

Pacific

Pacific

Atlantic

Pacific

Pacific

Pacific

WW Eastern Pacific

WW Americas

WW Western Atlantic

W-C Americas

W-C Americas

Circumtropical

WW Americas

Pacific Panama only

W-C Eastern Pacific (south)

W-C Western Atlantic (north)

WW Eastern Pacific

W-C Eastern Pacific (north)

WW Eastern Pacific

WW Eastern Pacific

Circumtropical

Circumtropical

WW Eastern Pacific

Circumtropical

WW Eastern Pacific

WW Americas

WW Americas

WW Americas

WW Eastern Padfic

WW Eastern Padfic

WW Western Atlantic

Circumtropical

Circumtropical (extended)

Circumtropical

Circumtropical

WW Americas

WW Eastern Padfic

WW Americas

Cosmopolitan

Circumtropical (extended)

W-C Eastern Padfic (south)

WW Eastern-Central Padfic

W-C Eastern Padfic (north)

WW Eastern Padfic

WW Eastern Padfic

Circumtropical

W-C Americas (south)

WW Atlantic (both sides)

Circumtropical

Cosmopolitan

W-C Eastern Padfic (north)

WW Eastern Padfic (extended)

North Pacific

WW Western Atlantic

Circumtropical

Cosmopolitan
NUMBER 221 67

TABI.K 2.—Continued

Species

Mesochaetopterus alipes

Spiochaetopterus costarum

Cirratulus cirratusi

Cirriformia luxuriosa

C. punctata

C. tentaculata

Dodecaceria concharum

Timarete perbranchiata

Flabelligera infundibularis

Pherusa. inftata

Piromis americana

Dasybranchus lumbricoides

Notomastus (Clistomastus) lineatus

Maldane gorgonensis

Owenia collaris

ldanthyrsus armatus

/. petmatus

Phragmatopoma attenuate

Sabellaria alcocki

S. floridensis

S. tnoorei

S. spinulosa

Isolda bipinnata

Eupolymnia nebulosa

E. regnans

Euthetepus pascua

Lanicides taboguillae

Loimia annulifilis

L. medusa

Pista alata

P. brevibranchiata

P. elongata

P. fasciata

P. herpinii

Streblosoma crassibranchia

Terebella gorgonae

Thelepus setosus

Terebellides stroemi?

Bispira monroi

Demonax leucaspis?

Hypsicomus phaeotenia

Megalomma vesiculosum

Potamilla fonticula

Pseudopotamilla intermedia

P. occelata

P. reniformis

Sabella melanostigma

Panama

Pacific

Pacific

Atlantic

Pacific

Atlantic

Pacific

Atlantic

Pacific

Pacific

Atlantic

Pacific

Both

Atlantic

Pacific

Atlantic

Pacific

Pacific

Pacific

Pacific

Both

Pacific

Pacific

Atlantic

Both

Pacific

Atlantic

Both

Pacific

Both

Pacific

Pacific

Pacific

Atlantic

Pacific

Both

Pacific

Atlantic

Pacific

Pacific

Both

Both

Atlantic

Atlantic

Pacific

Pacific

Atlantic

Both

General dispersal

_

Cosmopolitan

Cosmopolitan

W-C Eastern Pacific (north)

WW Western Atlantic

Circumtropical

Cosmopolitan

WW Eastern Pacific

W-C Eastern Pacific (north)

W-C Americas

WW Eastern Pacific

WW Indo-Pacific

Cosmopolitan

-

WW Eastern Pacific

W-C Eastern Pacific (south)

Circumtropical

WW Eastern Pacific

Circumtropical

WW Western Atlantic

Pacific Panama only

W-C Atlantic Ocean (north)

-

Cosmopolitan

WW Eastern Pacific

-

Panama, both sides

WW Indo-Pacific

Cosmopolitan

WW Eastern Pacific

WW Eastern Pacific

W-C Eastern Pacific (north)

Circumtropical (extended)

WW Indo-Pacific

WW Eastern Pacific

WW Eastern Pacific

Cosmopolitan

Cosmopolitan

-

Circumtropical

Circumtropical

Circumtropical

WW Western Atlantic

W-C Eastern Pacific

W-C Eastern Pacific

Cosmopolitan

WW Western Atlantic

Ib. Circumtropical species are found in warm

waters in the three major oceans. They are also

frequently present in the Mediterranean Sea. The

dispersal of this component is centered in the

tropical regions, so that dispersal around the continents in colder waters is assumed to be limited

or precluded.

Ila-b. Amphi-American species include forms

limited to the two American continents, but present

in both the Atlantic and Pacific Oceans. Two sub-
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

patterns can be recognized in that some species have

an extended dispersal into colder waters in at least

parts of their total range (Ha), whereas others are

limited to warm waters (lib).

III. Species limited to only one ocean include a

series of distinct patterns on both sides of the

isthmus. Species with an extended Atlantic distribution (IHal) have been reported from the coasts of

the Mid-Atlantic or New England states of the

United States. The warm water forms (IIIa2) have

been found in the Gulf of Mexico, Caribbean Sea,

and Bermuda in most cases, sometimes also in

Brazil or Venezuela. Endemic species (IHaS) have

never been reported outside Panama; in most cases,

they have been reported only once.

In the Pacific Ocean, North Pacific species (Illbl)

are known from Japan, Canada, and usually from

California, in addition to the present records.

Extended eastern Pacific patterns (IIIb2) can be

separated into two subpatterns; the first including

species known from boreal areas; the second including forms known from antiboreal areas in

addition to the warm-water records. Indo-Pacific

warm-water forms (IHbS) are known from the

central Pacific and usually from the Indian Ocean

as well. Eastern Pacific warm-water species (IIIb4)

are found from southern California to Galapagos

Islands and northern Peru. Finally, a series of

species have never been reported from areas outside

the Gulf of Panama (IIIb5).

A certain number of species have recorded occurrences on both the Atlantic and Pacific coastlines of

Panama. These have been listed in both columns in

Table 1 and, for that reason, the total number of

species showing dispersal patterns Ia.b and lib is

less than the sum of the two columns. The species

found on both sides of the Isthmus include 5 cosmopolitan, 14 circumtropical, and 11 species with

warm water amphi-American dispersal patterns.

An additional 52 species could potentially occur

on both sides. When these are absent from one side,

the absence is either due to incomplete sampling

or to a lack of suitable habitats on either side.

The general picture that emerges demonstrates

three dominant faunal elements on either side. The

widely dispersed, cosmopolitan or circumtropical

forms make up about 42 percent of the total fauna

on the Atlantic side and about 30 percent on the

Pacific side. A distinct amphi-American element,

limited largely to warm waters, makes up 24 percent

of the Atlantic fauna and 16 percent of the Pacific

fauna. Finally, the last faunal element includes

forms limited to one ocean only. This element

makes up about 33 percent of the total fauna on

the Atlantic side and 54 percent on the Pacific side.

Of these large fractions, elements also present in

cold water represent 5 percent of the total fauna in

the Atlantic Ocean; whereas the long-shore patterns

are much better developed in the Pacific Ocean,

where 15 percent of the total fauna are distributed

long-shore. This distribution appears to be related

to the prevalent current-patterns on the two sides.

Geminate species have been recognized in the

Panama region for crabs and fishes (e.g., Briggs,

1974); in general, the state of polychaete taxonomy

is too primitive to allow recognition of sister species even in the best analyzed families.
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70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1969. A Revision of Six Species of the Flavus-Bidentatus

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1972. Benthic Polychaetous Annelids from Deep Water off

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1973. Polychaetes from Central American Sandy Beaches.

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1923. Polychetes errantes. Faune de France, 5:1-488, 181

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1971. Spionidae (Polychaeta) of the Gulf of Mexico and

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1847. Beitrdge tur Kenntnis wirbellosen Thiere mit besonderer Berucksichtigung der Fauna des Norddeutschen Meeres. 170 pages, 2 plates. Braunschweig:

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1967 [1968]. Polychaeta from the Bay of Nha Trang. South

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1860. Beschreibung neuer oder wenig bekannter Anneliden, Beitrag: Zahlreiche Gattungen. Archiv fiir

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1871. Uebersicht der bisher beschrieben Terebellen und

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1875. Bemerkungen uber die Familie . der Aphroditeen.

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1877. Die von der Gazelle mitgebrachten Anneliden, zu

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1878. Annulata Semperiana: Beitrage zur Kenntniss der

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1882. Recherches sur les Annelides recueillies par M. le

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1936. New Species of Polychaetous Annelids of the Family

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1939a. Polychaetous Annelids, part 1: Aphroditidae to

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1939b. The Polychaetous Annelids Collected by the Presidential Cruise of 1938. Smithsonian Miscellaneous

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Polychaetous Annelids. Report on the Allan Hancock Atlantic Expedition, 3:1-33, 2 plates.

Polychaetous Annelids, part 5: Eunicea. Allan

Hancock Pacific Expeditions, 10:1-238,18 plates.

Polychaetous Annelids, part 6: Paraonidae, Mage

lonidae, Longosomidae, Ctenodrilidae and Sabellariidae. Allan Hancock Pacific Expeditions, 10:311-

389. 15 plates.

The Marine Annelids of North Carolina. Bulletin

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plates, 2 charts.

The Marine Annelids Erected by Kinberg with

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137, 18 plates.

Polychaetous Annelids: Goniadidae, Glyceridae,

Nephtyidae. Allan Hancock Pacific Expeditions,

15:1-181, 3 figures. 19 plates.

The Littoral Marine Annelids of the Gulf of Mexico.

Publications of the Institute of Marine Sciences

(University of Texas), 2:7-124.27 plates.

Iphitime and Ceratocephala (Polychaetous Annelids)

from California. Bulletin of the Southern California

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Australian Nereidae Including Description of Three

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Transactions of the Royal Society of South Australia,

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Endemism in the North Pacific Ocean, with Emphasis on the Distribution of Marine Annelids, and

Descriptions of New or Little Known Species.

Pages 39-59 in Essays in the Natural Sciences in

Honor of Captain Allan Hancock on the Occasion

of His Birthday July 26, 1955. 345 pages. Los Angeles: University of Southern California Press.

Polychaetous Annelids Erected by Treadwell, 1891

to 1948, Together with a Brief Chronology. Bulletin

of the American Museum of Natural History (New

York), 109:239-310. 1 plate.

Orbiniidae, Apistobranchidae, Paraonidae and Longosomidae. Allan Hancock Pacific Expeditions,

15:211-393,24 plates.

Catalogue of the Polychaetous Annelids of the

World. Allan Hancock Foundation Publications Occasional Paper, 23:1-629.

Systematic Account of Some Marine Invertebrate

Animals from the Deep Basins off Southern California. Allan Hancock Pacific Expeditions, 22:69-

216, 19 plates.

Polychaetous Annelids from California. Allan Hancock Pacific Expeditions, 25:1-226. 34 plates.

Atlas of Errantiate Polychaetous Annelids from

California. 828 pages. Los Angeles: Allan Hancock
72

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Foundation; University of Southern California.

1969. Atlas of Sedentariate Polychaetous Annelids from

California. 812 pages. Los Angeles: Allan Hancock

Foundation; University of Southern California.

Hartmann-Schroder, G.

1959. Zur Okologie der Polychaeten des Mangrove-EsteroGebietes von El Salvador. Beitrage zur Neotropischen Fauna (Jena), 1:69-183, 188 figures.

1960. Polychaeten aus dem Roten Meer. Kieler Meeresforschungen, 16:69-125, 185 figures.

1962a. Zweiter Beitrag zur Polychatenfauna von Peru.

Kieler Meeresforschungen, 18(l):109-147, 126 figures.

1962b. Zur Kenntnis der Nereiden Chiles (Polychaeta errantia) mit Beschreibung epitoker Stadien einiger

Arten und der Jugend-Entwicklung von Perinereis

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1965. Zur Kenntnis der Eulitoralen Polychaetenfauna von

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Verhandlungen der Naturwissenschaftliche Verein

(Hamburg), new series, 9:81-161, 85 figures.

1971. Annelida, Borstenwurmer, Polychaeta. Volume 58

in Dahl, Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihren Merkmalen und nach

ihrer Lebensweise. 594 pages, 191 figures. Jena.

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1885. On a Destructive Parasite of the Rock-Oyster (Polydora ciliata and P. polybranchia n. sp.). Proceedings

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275.

1886. Observation on Some Australian Polychaeta. Proceedings of the Linnean Society of New South

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Hoagland, R. A.

1919. Polychaetous Annelids from Porto Rico, the Florida

Keys and Bermuda. Bulletin of the American Museum of Natural History (New York), 41:517-591, 4

plates.

Horn, R.

1912. Polychaeta Errantia of the Siboga Expedition, part

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1917. On a New Aphrodite Species (Aphrodite roulei)

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Imajima, M.

1966a. The Syllidae (Polychaetous Annelids) from Japan,

2: Autolytinae. Publications of the Seto Marine

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1966b. The Syllidae (Polychaetous Annelids) from Japan,

III: Eusyllinae. Publications of the Seto Marine

Biological Laboratory, 14:85-116, 10 figures.

1966c. The Syllidae (Polychaetous Annelids) from Japan,

4: Syllinae, 1. Publications of the Seto Marine Biological Laboratory, 14:219-252, 12 figures.

1966d. The Syllidae (Polychaetous Annelids) from Japan,

5: Syllinae, 2. Publications of the Seto Marine Biological Laboratory, 14:253-294, 15 figures.

Imajima, M., and O. Hartman

1964. The Polychaetous Annelids of Japan, parts 1 and 2.

Allan Hancock Foundation Publications Occasional

Paper, 26:1-452, 38 plates.

Izuka, A.

1912. The Errantiate Polychaeta of Japan. Journal of the

College of Science, Imperial University of Tokyo,

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Johnson, H. P.

1897. A Preliminary Account of the Marine Annelids of

the Pacific Coast, with Descriptions of New Species:

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1901. The Polychaeta of the Puget Sound Region. Proceedings of the Boston Society of Natural History,

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Johnston, G.

1840. Mescellanea Zoologica: Contributions towards a History of Irish Annelids. Annals and Magazine of

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Jones, M. L.

1962. On Some Polychaetous Annelids from Jamaica, the

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1963. Four New Species of Magelona (Annelida, Polychaeta) and a Redescription of Magelona longicornis

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Jumars, P. A.

1974. A Generic Revision of the Dorvilleidae (Polychaeta)

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1855. Nya slagten och arter af Annelider. Ofversigt af

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1857. Nya slagten och arter af Annelider. Ofversigt af

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1865. Annulata nova. Ofversigt af Kongl. Vetenskapsakademiens Forhandlingar, 21:559-574.

1866a. Annulata nova. Ofversigt af Svenska Vetenskapsakademiens Forhandlingar, 22:167-179.

1866b. Annulata nova. Ofversigt af Svenska Vetenskapsakademiens Forhandlingar, 22:239-258.

1867. Annulata nova. Ofversigt af Kongl. VetenskapsAkademiens Forhandlingar, 23:337-357.

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1951. Nereidae and Eunicidae of South Western Australia; Also Notes on the Ecology of Western Australia Limestone Reefs. Journal of the Royal Society

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NUMBER 221 73

1852. Uebcr die Erscheinungen bei der Fortpflanzung von

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1879. Die Wurmfauna von Madeira, part 1. Zeitschrift

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1855. Contributions towards a Knowledge of the Marine

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1876. On the Annelida of the Porcupine Expeditions of

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1885. Report on the Annelida Polychaeta Collected by

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1928a. On the Polychaeta Collected by Dr. Th. Mortensen

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1939. Polychaeta. B.AN2.. Antarctic Research Expedition

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1808. New and Rare Animals Found on the South Coast

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1815. Descriptions of Several New or Rare Animals Principally Marine, Found on the South Coast of Devonshire. Transactions of the Linnean Society of

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1818. Descriptions of Five British Species of the Genus

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1904. New Polychaeta from California. Proceedings of the

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1858. Einiges uber die Anneliden Fauna der Insel St.

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1771. Von Wiirmern des sussen und salzigen Wassers. 200

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1843a. Gr0nlands Annulata dorsibianchiata. Kongelige

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1961. New Species of Polychaete Worms from the Atlantic

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Index

(Synonyms and major entry for each species are in italics)

abranchiata, Glycera, 34, 35, 66

acicula, Goniada, 35, 66

aciculata, Typosyllis, 18, 21, 65

aedificatrix, Eunice antennata, 38, 66

affinisf, Flabelligera, 51

afra, Eunice, 37, 38, 39, 66

Aglaophamus dicirris, 33,65

tabogensis, 33, 34, 65

Aglaura fulgida, 45

agulhana, Neanthes, 27

alata, Pista, 56, 58, 67

albifrons, Lumbrineris, 44

Albini, Ancistrosyllis, 17

albini, Synelmis, 77,65

alcocki, Sabellaria, 53, 54, 67

alcocki, Sabellaria spinulosa, 54

alipes, Mesochaetopterus, 48, 49, 67

alipes, Alesochaetopterus taylori var., 49

amadae, Marphysa, 37, 41, 42. 66

ambigua, Eurythoe, 12

Linopherus, 11, 72,65

Pseudeurythoe, 12

Am bo americana, 51

perbranchiata, 51

americana, Ambo, 51

Glycera, 34, 35, 66

Piromis, 51,52, 67

americana, Ceratocephala crosslandi, 23

Marphysa sanguinea var., 42

Slylarioides capeusis, 52

Americonuphis reesei, 35, 36, 66

Amrnotrypane aulogaster, 52

Ammotrypane species indeterminate. 52

Ampharetidae, 55

Amphinomidae, 11

Amphitrite reniformis, 6.*?

vesiculosa, 62

Anaitides erythrophylla. 7?. 65

lamellifera, 13, 14, 65

madeirensis, 13, 14, 65

panamensis, 15

species indeterminate, 14

Ancistrosyllis Albini, 17

gorgonensis, 17

anderssoni, Perinereis. 23, 31, 65

angolanus, Staurocephalns, 46

annulifilis, Loimia, 56, 58, 67

Tcrebella, 58

annulifilis, Loimia medusa var., 58

anoplos, Autolvtus, 17, 18, 19, 65

antennata, Eunice, 38, 66

Leodice, 38

antennata aedificatrix, Eunice, J£, 66

Aphrodita armifera, 3

carnnculata, 12

complanata, 11

diplops, 2, 64

falcifera, 3

japonica, 3

refulgida, 3, 4

roulei, 3

solitaiia. 3, 4

Aphroditidae, 2

aphroditois, Eunice, 37, 38, 39, 66

Nereis, 38

Arabella mutans. 44, 45, 52, 66

species indeterminate, 45

Arabellidae, 44

armatus, Idanthyrsus, 53, 54, 67

armifera, Aphrodita, 3

attenuata, Phragmatopoma, 53, 54, 67

Audouinia luxuriosa, 50

polytricha, 50

tenlaculata, 50

aulogaster, Ammotrypane, 52

Autolytus anoplos, 17,18, 19, 65

irregularis, 19

cf. magnus, 17,79,65

spinoculatus, 19

species indeterminate, 17. 19

bairdii, Perinereis, 31

balboensis, Harmothoe, 4, 5. 65

bellis, Paleanotus, 10

Sabellaria, 55

Bhauania goodei, 10, 64

rived, 10, 64

biannulata, Eunice, 38, 39,66

bifoliata. Eulalia (Sige), 14

Eumida, 13.74, 15,65

bipinnata, Isolda, 55, 67

Bispira monroi, 60, 61, 67

nigosa monterea, 60

volutacornis, 61

Boccardia polybranchia, 47, 66

proboscidea, 47, 66

tricuspa, 47, 66

Branchiomma vesiculosum, 62

brevibranchiata, Pista, 56, 59, 67

Potamilla. 63

brevicirra, Lumbrineris, 44

brunnea, Opisthosyllis, 17, 20, 65

caducus lumbricoides, Dasybranchus, 52

caeca, Eunereis, 25

Sy//w, 21

Typosyllis, 18, 27, 65

callaona, Nereilepas, 29

Nereis, 23, 29, 65

canariensis, Linopherus, 11, 72, 65

Pseudeurythoe, 12

capensis americana, Stylarioides, 52

Capitellidae, 52

cariboea, Eunice, 39

Eunice (Nicidion), 37, i9, 40, 66

cariboea var. kinbergi, Eunice, 39, 40

carinulatus, Lepidonotus, 6

carunculata, Aphrodita, 12

Hermodice, 11,72,65

Cauleriella species indeterminate, 49, 50

Cenothrix mutans, 45

cerasina, Dorvillea, 46, 66

cerasinus, Staurocephalus, 46

Ceratocephala crosslandi, 22, 23, 65

crosslandi americana, 23

Ceratonereis irritabilis, 22, 23, 65

mirabilis, 22, 23, 65

tentaculata, 23

versipedata, 23

Chaetacanthus magnifkus, 4,64

Chaetopteridae, 48

Chaetopterus variopedatus, 48, 49, 66

Chaetozone species indeterminate, 49, 50

Chaunorhynchus crosslandi, 23

chilensis, Ninoe, 43, 44, 66

chiliensis, Diopatra, 35, 36,66

Chloeia entypa, 77, 65

pinnata, 11

viridis, 11, 65

Chone species indeterminate, 60, 61

chrysolepis, Paleanotus, 10, 11, 65

Chrysopetalidae, 10

Chrysopetalum occidentale, 10, 64

circumspiciens, Hypsicomus, 62

Cirratulidae, 49

Cirratulidae indeterminate, 51

Cirratulus cirratus?, 49, 50,67

luxuriosus, 50

punctata, 50

cirratus?, Cirratulus, 49, 50,67

76
NUMBER 221 77

cirratus, Lumbricus, 50

Cirrhatulus punctata, 50

Cirriformia filigera, 50, 51

luxuriosa, 49, 50,67

punctata, 49, 50, 67

spirabranchia, 50

tentaculata, 49, 50, 51, 67

Clymene (Euclymene) rubrocincta, 53

Clymenella species indeterminate, 53

collaris, Lysidice, 40

Owenia, 53, 67

collaris, Owenia fusiformis, 53

complanata, Aphrodita, 11

Eurythoe, 11, 12,65

concharum, Dodecaceria, 49,51, 67

cornuta, Langerhansia, 18,19, 20, 65

Syllis, 19

SyZ/M (Ehlersia), 19

custarum, Spiochaetopterus, 48, 49, 67

Telepsavus, 49

crassibranchia, Streblosoma, 56, 59,67

crosslandi, Ceratocephala, 22, 23, 65

Chaunorhynchus, 23

Lepidonotus, 4, <J, 64

crosslandi americana, Ceratocephala, 23

cruzensis?, Lumbrineris, 43,66

cuprea, Diopatra, 35. 36, 66

Nereis, 36

Dasybranchus caducus lumbricoides, 52

glabrus, 52

lumbricoides, 52,67

dawydovi, Neanthes, 27

Demonax leucaspis?, 60, 61, 62. 67

medius, 62

denhamensis, Nereis, 31

dentata, Diopatra, 36

denticulata, Diopatra, 35, 36, 66

dibranchis, Nephthys, 33

dicirris, Aglaophamus, 33, 65

Diopatra chiliensis, 35, 36, 66

cuprea, 35, 36, 66

dentata, 36

denticulata, 35, 36, 66

obliqua, 36

ornata, 35, 36,66

diplops, Aphrodita, 2, 64

Distylia monterea, 60

rugosa, 60

Distylidia monroi, 60

rugosa, 60

diversicolor, Hediste, 27

Neanthes, 27

Dodecaceria concharum, 49. 5/. 67

Dorvillea cerasina, 4<S, 66

rubrovittatus, 46, 66

rudolphi, 46

Dorvilleidae, 46

Drilonereis falcata, 44, 45, 66

filum, 45

mexicana, 44, 4.5, 66

nuda, 44, 45,66

dubia, Eurythoe, 12

Dumerilii, Nereis, 31

dumerilii, Platynereis, 23, if, 32, 65

edentulum, Eunice sidliensis var., 42

edentulum, Nicidion, 42

Ehlersia mexicana, 20

elongata, Pista, 56, 59, 67

entypa, Chloeia, 11,65

erecta, Lumbriconereis, 43

Lumbrineris, 4.3, 66

erythrophyllus, Anaitides, 13, 65

Lepadorhynchus, 13

Eteone species indeterminate, 14

Eudistylia polymorpha, 60

Eulalia lobocephala, 16

lobocephalica, 16

myriacydum, 13,14, 65

(S/ge) bifoliata, 14

species indeterminate, 14

Eumida bifoliata, 13,14, 15. 65

Eunereis caeca, 25

longipes, 25

longissima. 25

marri, 25

paitillae, 22, 25, 26, 65

paradoxa, 25

patagonica, 25

wailesi, 25

Eunice afra, 37,38, 39, 66

antennata, 38, 66

antennata aedificatrix, 38, 66

aphroditois, 37, 38, 39, 66

biannulata, 38, 39,66

cariboea, 39

cariboea var. kinbergi, 39. 40

filamentosa, 37, ?9, 66

longicirrata, 39, 40

mutilata, 37, 38, 39, 66

(Nicidion) cariboea, 37. 39, 40. 66

reducta, 37, 40, 66

siciliensis, 42

siciliensis var. edentulum, 42

tridentata, 40

vittata, 40

vittatopsis. 38, 40, 66

websteri, 38, 39, 40, 66

species indeterminate, 40

Eunicidae, 37

Eupanthalis kinbergi, 7. 8

nigromaculata, 8

perlae, 7, 8, 64

Eupolymnia nebulosa. 56, 67

regnans, 56, 57, 67

Eurythoe ambigua, 12

complanata, 7/, 12. 65

dubia, 12

oculata, 12

Eusigalion hancocki, 10

Euthelepus malayensis, 58

pascua, 56, 57,58, 67

setobalensis, 58

tenuis, 58

excilis, Typosyllis, 21

falcata, Drilonereis, 44, 45,66

falcifera, Aphrodita, 3

fasciata, Pista, 56, 59, 67

Terebella (Phyzelia), 59

filamentosa, Eunice, 37, 39, 66

filigera, Cirriformia, 50, 51

filum, Drilonereis, 45

Flabelligera affinisi, 51

infundibularis, 51, 67

Flabelligeridae, 51

floridana, Potamilla, 62

floridensis, Sabellaria, 53, 54, 55, 67

foliosa, Ninoe, 44

fonticula, Parasabella, 62

Potamilla, 60, 62,67

fulgida, Aglaura, 45

Oenone, 45, 46, 66

funchalensis, Nereis, 31

fusca, Sthenelais, 9,64

fuscomarmorata, Halosydna, 5

fuscosuturata, Syllis, 21

Syllis (Typosyllis), 21

Typosyllis, 18, 21, 22,65

fusiformis, Owenia, 53

fusiformis collaris, Owenia, 53

galetae, Neanthes, 23, 26, 27, 65

gallapagensis, Pseudonereis, 23, 32, 33, 65

gigas, Lepidasthenia, 4, 5, 6,64

Polynoe, 5

glabra, Halosydna. 4, 5, 64

glabrus, Dasybranchus, 52

Glycera abranchiata, 34, 35. 66

americana, 34, 35, 66

oxycephala, 34, 35, 66

tesselata, 34. 35, 66

Glyceridae, 34

Goniada acicula, 35, 66

Goniadidae, 35

goodei, Bhawania, 10, 64

gorgonae, Terebella, 56, 59, 60, 67

gorgonensis, Ancistrosyllis, 17

Maldane, 53,67

Nothria, 35, 37, 66

Onuphis, 37

gracilis, Syllis, 18, 20, 21, 65

granulata, Polynoe, 5

near guanica, Podarke, 17

Gyptis species indeterminate, 5, 16

Halosydna fuscomarmorata, 5

glabra, 4, 5, 64

johnsoni, 4
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

leucohyba, 4, 5,64

reticulata, 4

hancocki, Ewigalion, 10

Haploscoloplos panamensis, 46, 66

Haplosyllis spongicola, 17,19, 65

Harmothoe balboensis, 4,5,64

hirsuta, 4, 5, 64

lunulata pacifica, 5, 16

species indeterminate, 5

Hediste diversicolor, 27

heirissonensis, Nereis, 31

Hennodice carunculata, 11,12, 65

striata, 13

herpini?, Pista, 56, 59,67

Hesione intertexta, 76, 65

picta, 16, 17,65

Hesionidae, 16

heterobranchia texana, Pfionospio, 47,

48,60

heteropoda, Lumbrineris, 43

hirsuta, Harmothoe, 4, 5,64

humilis, Lepidoriotus, 4, 6, 64

hyalina, Syllis, 22

Typosyllis, 18,22, 65

Hypsicomus circumspiciens, 62

phaeouenia, 60, 62, 67

torquatus, 62

ldanthyrsus aruiatus, 53, 54, 67

pennatus, 53, 54, 67

inflata, Lumbrineris, 43, 44, 66

Pherusa, 51, 52, 67

Trophonia, 51

infundibularis, Flabelligera, 51, 67

intermedia, Pseudopotamilla, 60, 62, 63,

67

intertexta, Hesione, 16,65

Iphione magnified, 4

ovata, 4, 5,64

irregularis, Autolytus, 19

irritabilis, Ceratonereis, 22, 23, 65

Nereis, 23

Isolda bipinnata. 55,67

pulchella, 55

sibogae, 55

warnbroensis, 55

whydahaensis. 55

jacksoni. Nereis, 31

jacksoni reducta, Nereis, 31

japonica, Aphrodita, 3,64

Laonice, 47, 66

japonicus, Spionides, 47

jogasimae, Panthalis, 9

johnsoni, Halosydna, 4

Lepidonotus, 4

johnstoni, Lepidonotus, 6

Thormora, 4, 6, 7, 64

kerguelensis, Neanthes, 27

keiguelensis oligodonta, Neanthes, 27

kinbergii, Eunice cariboea var., 39, 40

kinbergi, Eupanthalis, 7, 8

lamellifera, Anailides, 13,14, 65

Nereis, 13

Phyllodoce (Anaitides), 13

Langerhansia cornuta, 18,19, 20,65

mexicana, 18,20, 65

Lanicides taboguillae, 56, 58,67

Laonice japonica, 47, 66

latreilli, Lumbrineris, 43, 44, 66

Leodice antennata, 38

Lepadorhynchus erythrophyllus, 13

Lepidametria virens, 5,6

Lepidasthenia gigas, 4, 5,6, 64

varius, 4, 6, 64

Lepidonotus carinulatus, 6

crosslandi, 4, 6, 64

humilis, 4, 6,64

johnsoni, 4

johnstoni, 6

nesophilus, 4, 6, 64

leucaspis?, Demonax, 60, 61,62, 67

leucohyba, Halosydna, 4, 5, 64

Polynoe, 5

lewisii, Sigalion, 9

Thalenessa, 5>, 10, 64

lineatus, Notomastus (Clistomastus), 52,

67

Linopherus ambigua, 11,12, 65

canariensis, 11,12,65

oculata, 11,12, 65

lobocephala, Eulalia, 16

lobocephalica, Eulalia, 16

Steggoa, 13. i6, 65

Loimia annulifilis, 56, 58, 67

medusa, 56, 58, 67

medusa var. annulifilis, 58

longibranchia, Ninoe, 44

longicirrata, Eunice, 39, 40

longicornis, Schistomeringos, ¥6, 66

Stauronereis, 46

longipes, Eunereis, 25

longissima, Eunereis, 25

Syllis, 20, 21

lumbricoides, Dasybranchus, 52, 67

Dasybranchus caducus, 52

Lumbriconereis erecta, 43

unicornis, 42

Lumbricus cirratus, 50

Lumbrineridae, 43

Lumbrineris albifrons, 44

brevicirra, 44

cruzensis?, 43. 66

erecta, 43, 66

heteropoda, 43

inflata, 43, 44,66

latreilli, 43, 44,66

sphaerocephala, 43, 44

tetraura, 43, *•/, 66

species indeterminate, 44

lunulata pacifica, Harmothoe, 5, 16

luxuriosa, Audouinia, 50

Cirriformia, 49,50, 67

luxuriosus, Cirratulus, 50

Lysaretidae, 45

Lysaretidae indeterminate, 46

Lysidice collaris, 40

ninetta, 32, 37, 40,41,66

macroceros orientalis, Sige, 16

maculata, Typosyllis, 21

madeirensis, Anaitides, 13,14, 65

Phyllodoce (Anaitis), 14

Magelona pacifica, 48,66

papillicornis, 48,66

Magelonidae, 48

magna, Onuphis, 35

magnifica, Iphione, 4

magnificus, Chaetacanthus, 4, 64

cf. magnus, Autolytus, 17,19,65

malayensis, Euthelepus, 58

Maldane gorgonensis, 53, 67

Maldanidae, 53

Maldanidae indeterminate, 53

Marphysa amadae, 37,41,42, 66

sanguinea, 37, 42, 66

sanguinea americana, 42

species indeterminate, 42

marri, Eunereis, 25

medius, Demonax, 62

medusa var., annulifilis, Loimia, 58

medusa, Loimia, 56, 58,67

Terebella, 58

Megalomma vesiculosum, 60, 62,67

melanostigma, Sabella, 60, 63, 67

Mesochaetopterus alipes, 48, 49, 67

minutus, 49

taylori alipes, 49

mexicana, Drilonereis, 44. 45,66

Ehlersia, 20

Langerhansia, 18, 20,65

minutus, Mesochaetopterus, 49

mirabilis, Ceratonereis, 22,23, 65

monroi, Bispira, 60,61, 67

Distylidia, 60

Nephtys, 33, 34,66

monterea, Bispira rugosa, 60

Distylia, 60

moorei, Sabellaria, 54, 55,67

Sabellaria pectinata, 55

mortenseni, Panthalis, 7, «, 9,64

Polyodontes, 8

mutans. Arabella. 44, 45, 52, 66

Cenothrix, 45

mutilata, Eunice, 37, 38, 39, 66

myriacyclum, Eulalia, 13,14,65

Notophyllum, 14
NUMBER 221 79

Neanthes agulhana, 27

dawydovi, 27

diversicolor, 27

galetae, 23, 26, 27, 65

kerguelensis, 27

kerguelensis oligodonta, 27

noodti, 29

pseudonoodti, 23,27, 28, 29,65

ruficeps, 29

seridentata, 29

succinea, 23, 29, 65

species indeterminate, 29

nebulosa, Eupolymnia, 56, 67

Onuphis, 35, 37, 66

Polymnia, 56

Terebella, 56

Nematonereis unicornis, 37, 42, 66

Nephthys dibranchis, 33

tabogensis, 33, 34

Nephtyidae, 33

Nephlys monroi, 33, 34, 66

singularis, 33, 34, 66

squamosa, 33,34, 66

Nereidae, 22

Nereidae indeterminate, 33

Nereilepas callaona, 29

variegata, 33

Nereis aphroditois, 38

callaona, 23, 29, 65

cuprea, 36

denhamensis, 31

Dinner Mi, 31

funchalensis, 31

heirissonensis, 31

irritabilis, 23

jacksoni, 31

jacksoni reducta, 31

lamellijera, 13

panamensis, 23, 29, 30, 31, 65

riisei, 23, 31, 65

succinea, 29

thompsoni, 31

uesophilus, Lepidonotus, 4, 6,64

Nicidion edentulum, 42

Nicolea taboguillae, 58

nigromaculata, Eupanthalis, 8

ninetta, Lysidice, 32, 37, 40, 41,66

Ninoe chilensis, 43, 44, 66

foliosa, 44

longibranchia, 44

noodti, Neanthes, 29

Nothria gorgonensis, 35, 37,66

Nolocirrus tetraurus, 44

Notomastus (Clistomastus) lineatus, 52,

67

species indeterminable, 52

Notophyllum myriacyclum, 14

Notopygos ornata, 11,12, 13, 65

nuda, Drilonereis. 44. 45,66

obliqua, Diopatra, 36

obscura, Podarke, 17

obscurus, Ophiodromus, 16,17,65

occelata, Pseudopotamilla, 60, 63, 67

occidentale, Chrysopetalum, 10,64

oculata, Eurythoe, 12

Linopherus, 11,72, 65

Phyllodoce, 13

Pseudeurythoe, 12

oculea, Panthalis, 9

Polyodontes, 7, 9,64

Odontosyllis polycera, 18,20,65

species indeterminate, 20

Oenone fulgida, 45, 46,66

oligodonta, Neanthes kerguelensis, 27

Onuphidae, 35

Onuphis gorgonensis, 37

magna, 35

nebulosa, 35,37,66

vermillionensis, 35, 37,66

Opheliidae, 52

Ophiodromus obscurus, 16,17,65

pugettensis, 16,17,65

Opisthosyllis brunnea. 17,20,65

Orbiniidae, 46

orientalis?, Sige, 13,16, 65

ornata, Diopatra, 35, 36, 66

Notopygos, 11,12, 13,65

ovata, Iphione, 4,5,64

Owenia collaris, 53,67

fusifovmis, 53

fusiformis collaris, 53

Oweniidae, 53

oxycephala, Glycera, 34,35,66

pacifka, Harmothoe lunulata, 5, 16

Magelona, 48,66

Panthalis, 7, 9,64

paitillae, Eunereis, 22, 25, 26,65

Paleanotus bellis, 10

chrysolepis, 10, 11, 65

Palola paloloides, 43

siciliensis, 37, 42, 43, 66

paloloides, Palola, 43

panamensis, Anaitides, 15

Haploscoloplos, 46, 66

Nereis, 23, 29, 30, 31, 65

Phyllodoce. 13,15, 16,65

Phyllodoce (Anaitides), 15

Panthalis jogasimae, 9

mortenseni, 7, #, 9,64

oculea, 9

pacifica, 7, 9,64

papillata, Stylarioides, 51

papillicornis, Magelona, 48,66

paradoxa, Eunereis, 25

Paraprionospio pinnata, 48

Parasabella fonticula, 62

pascua, Euthelepus, 56, 57,58,67

patagonica, Eunereis, 25

pectinata moorei, Sabellaria, 55

pectinata var., Sabellaria, 54

pennata, Sabellaria (Pallasia), 54

pennatus, Idanthyrsus, 53,54,67

perbranchiata, Ambo, 51

Timarete, 49, .52, 67

Perinereis anderssoni, 23,31,65

bairdii, 31

species indeterminate, 31

perlae, Eupanthalis, 7,8,64

peruensis, Phragmatopoma, 54

phaeotaenia, Hypsicomus, 60, 62,67

Sabella, 62

Phenacia setosa, 60

Pherecardia striata, 11,13,65

Pherusa inflata, 52,52,67

Phragmatopoma attenuata, 53, 5*. 67

peruensis, 54

species indeterminate, 54

Phyllochaetopterus species indeterminate, 48, 49

Phyllodoce (Anaitides) lamellifera, 13

(Anaitides) panamensis, 15

(Anaitis) madeirensis, 14

oculata, 13

panamensis, 13,15,16,65

Phyllododdae, 13

picta, Hesione, 16, 17, 65

Pilargiidae, 17

pinnata, Chloeia, 48

Paraprionospio, 48

Prionospio, 48

Pionosyllis species indeterminate, 18, 20

Piromis amerkana, 51,52,67

Pista alata, 56,58,67

brevibranchiata, 56, 59,67

elongata, 56, 59,67

fasdata, 56. 59,67

herpini?, 56,59,67

Platynereis dumerili, 23, 31, 32,65

species indeterminate, 32

Podarke near guanica, 17

obscura, 17

pugettensis, 17

polybranchia, Boccardia, ¥7,66

Polydora (Boccardia), 47

polycera, Odontosyllis. 18,20,65

Sy//ii, 20

Polycirrus species indeterminate, 56,59

Polydora (Boccardia) polybranchia, 47

tricuspa, 47

Polymnia nebulosa, 56

regnans, 56

polymorpha, Eudistylia, 60

Polynoe gigas, 5

granulata, 5

leucohyba, 5

pustulata, 5

reticulata, 4

taeniata, 7
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Polynoidac, 4

Polyodontes morlenseni, 8

oculea, 7, 9, 64

Polyodontidae, 7

polytricha, Audouinia, 50

Potamilla brevibranchiata, 63

floridana, 62

fonticula, 60, 62, 67

reniformis, 63

Prionospio heterobranchia texana, 47,

48,66

pinnata, 48

species indeterminate, 48

proboscidea, Boccardia, 47, 66

prolifera, Syllis, 22

Syllis (Typosyllis), 22

prolifera?, Typosyllis, 18, 22, 65

Psammolyce spinosa, 9, 64

Pseudeurythoe ambigua, 12

canariensis, 12

oculata, 12

Pseudonereis gallapageusis, 23, 52, 33, 65

variegata, 23, 33,65

pseudonoodti, Neanthes, 23, 27, 28, 29,

65

Pseudopotamilla intermedia, 60, 62, 63,

67

occelata, 60, 63, 67

reniformis, 60, 63, 67

pugettensis, Ophiodromus, 16, 17, 65

Podarke, 17

pulchella, Isolda, 55

punctata, Cirratulus, 50

Cirrhatulus, 50

Cirriformia, 49, 50,67

pustulate, Polynoe, 5

reducta, Eunice, 37, -#0,66

reducta, Nereis jacksoni, 31

reesei, Americonuphis, 35, 36, 66

refulgida, Aphrodita, 3, 4

regnans, Eupolymnia, 56, bl, 67

Polymnia, 56

reniformis, Amphitrite, 63

Potamilla, 63

Pseudopotamilla, 60, 65, 67

reticulata, Halosydna. 4

Polynoe, 4

riisei, Nereis, 23,57,65

riveti, Bhawania, /0, 64

roulei, Aphrodita, 3

rubrocincta, Clymene (Euclymene), 53

rubrovittatus, Dorvillea, 46, 66

Staurocephalus, 46

rudolphi, Dorvillea, 46

rudolphii, Staurocephalus, 46

ruficeps, Neanthes, 29

rugosa, Distylia, 60

Distylidia, 60

rugosa monterea, Bispira, 60

Sabella melanostigma, 60, 65,67

phaeotaenia, 62

species indeterminate, 63

Sabellaria alcocki, 53, 5*, 67

beilis, 55

floridensis, 53. 54, 55, 67

moorei, 54, 55, 67

pectinata moorei, 55

peclinata var., 54

(Pallasia) pennala, 54

(Phragmatopoma) virgini, 54

spinulosa, 54, 55, 67

spinuloso alcocki, 54

Sabellariidae, 53

Sabellariidae indeterminate, 55

Sabellidae, 60

Sabellidae indeterminate, 63

sanguinea, Marphysa, 37, 42, 66

sanguinea americana, Marphysa, 42

Schistomeringos longicornis, 46, 66

seridentata, Neanthes, 29

setobalensis, Euthelepus, 58

setosa, Phenacia, 60

setosus, Thelepus, 56, 59, 60, 67

sibogae, Isolda, 55

siciliensis, Eunice, 42

Palola, 37, 42, 43, 66

siciliensis edentulum, Eunice, 42

Sigalion lewisii, 9

Sigalionidae, 9

Sige macroceros orientalis, 16

orientalis?, 13,16, 65

singularis, Nephtys, 33,34,66

solitaria, Aphrodita, 3,4

sphaerocephala, Lumbrineris, 43, 44

spinoculatus, Autolytus, 19

spinosa, Psammolyce, 9,64

spinulosa, Sabellaria, 54, 55,67

spinulosa alcocki, Sabellaria, 54

Spiochaetopterus costarum, 48, 49, 67

Spionidae, 47

Spionides japonicus, 47

spirabranchia, Cirriformia, 50

spongicola, Haplosyllis, 17,19,65

Syllis, 19

squamosa, Nephtys, 33, 34, 66

Staurocephalus angolanus, 46

cerasinus, 46

rubrovittatus, 46

rudolphii, 46

Stauronereis longicornis, 46

Steggoa lobocephalica, 13, 76, 65

Sthenelais fusca, 9, 64

Streblosoma crassibranchia, 56, 59,67

verrilli, 59

striata, Hermodice, 13

Pherecardia, 11.75, 65

stroemi?, Terebellides, 60,67

Stylarioides capensis americana, 52

papillata, 51

succinea, Neanthes, 23, 29, 65

Nereis, 29

Syllidae, 17

Syllidae indeterminate, 22

Syllis caeca, 21

cornuta, 19

(Ehlersia) cornuta, 19

fuscosuturata, 21

gracilis, 18,20,21,65

hyalina, 22

longissima, 20, 21

polycera, 20

prolifera, 22

spongicola, 19

taeniaformis, 21

(Typosyllis) fuscosuturata, 21

(Typosyllis) prolifera, 22

(Typosyllis) variegata, 22

variegata, 22

Syne! mis albini, 77, 65

tabogensis, Aglaophamus, 55, 34, 65

Nephthys, 33, 34

taboguillae, Lanicides, 56, 5#, 67

Nicolea, 58

taeniaformis, Syllis, 21

Trypanosyllis, 21

Trypanosyllis (Trypanedenta), 17, 2/,

65

taeniata, Polynoe, 7

Thormora, 4, 7, 64

taylori alipes, Mesochaetopterus, 49

Telepsavus costarum, 49

tentaculata, Audouinia, 50

Ceratonereis, 23

Cirriformia, 49, 50, 51, 67

Terebella, 50

tenuis, Euthelepus, 58

Terebella annulifilis, 58

gorgonae, 56, 59,60, 67

medusa, 58

nebulosa, 56

(Phyzelia) fasciata, 59

tentaculata, 50

Terebellidae, 56

Terebellidae indeterminate, 60

Terebellides stroemi?, 60,67

tesselata, Glycera, 34, 55,66

tetraura, Lumbrineris, 43, *#, 66

tetraurus, Notocirrus, 44

texana, Prionospio heterobranchia, 47

48,66

Thalenessa lewisii, 9, 10, 64

Tharyx species indeterminate, 49, 51

Thelepus setosus, 56, 59, 60, 67

thompsoni, Nereis, 31

Thormora johnstoni, 4, 6, 7, 64

taeniata, 4, 7,64

Timarete perbranchiata, 49, 51,67
NUMBER 221 81

toiquatus, Hypsicomus, €2

Trichobranchidae, 60

Tricoelia variopedata, 49

tricuspa, Boccardia, 47, 66

Polydora, 47

tridentata, Eunice, 40

Trophonia inflata, 51

Trypanosyllis taeniajormis, 21

Trypanosyllis (Trypanedenta) taeniaformis, 17, 21,65

Typosyllis aciculata, 18, 21, 65

caeca, 18, 21, 65

excilis, 21

fuscosuturata, 18, 21,22, 65

hyalina, 18, 22, 65

maculata, 21

prolifera?, 18, 22, 65

variegate, 18, 21, 22,65

unicornis, Lumbriconereis, 42

Nematonereis, 37, 42,66

variegata, Nereilepas, 33

Pseudonereis, 23,}), 65

Syi/ts, 22

Syltis (Typosyllis), 22

Typosyllis, 18,21.22,65

variopedata, Tricoelia, 49

variopedatus, Chaetopterus, 48, 49, 66

varius, Lepidasthenia, 4, 6,64

vermillionensis, Onuphis. 35, 37, 66

verrilli, Streblosoma, 59

versipedata, Ceratonereis, 23

vesiculosa, Amphitrite, 62

vesiculosum, Branchiomma, 62

Megalomma, 60, 62, 67

virens, Lepidametria, 5, 6

virgini, Sabellaria (Phragmatopoma), 54

viridis, Chloeia, ii , 65

vittata, Eunice, 40

vittatopsis, Eunice, 38,40,66

volutacornis, Bispira, 61

wailesi, Eunereis, 25

warnbroensis, Isolda, 55

websteri, Eunice, 38, 39, 40,66

whydahaensis, Isolda, 55



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