MARIAN H. PETTIBONEV£

 Revision of Some

* Species Referred to

Leptonereis, Nicon,

and Laeonereis

(Polychaeta: Nereididae)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 104
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 104

Marian H. Pettibone ReVlSlOIl o f Som e

Species Referred to

Leptonereis, Nicon,

and Laeonereis

(Polychaeta: Nereididae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Pettibone, Marian H. Revision of some species referred to Leptonereis, Nicon, and

Laeonereis (Polychaeta: Nereididae). Smithsonian Contributions to Zoology, number 104, 53 pages, 27 figures. 1971.—Some 47 nereid species, which have been referred directly or indirectly to Leptonereis, Nicon, and Laeonereis, are reviewed and

revised, based on examination of type-specimens, where available, and additional

nereid material. Diagnoses are given for eight genera, two of them emended and two

of them new. Sixteen species (including 16 synonyms) are referred to the eight genera, one of which is considered to be new. Supplementary descriptions and figures are

given for fourteen of them. Four additional species are referred to Platynereis Kinberg, one of which is described and figured. Four species are considered to be

indeterminable.

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

in the Institution's annual report, Smithsonian Year.

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Marian H. Pettibone ReVlSlOn o f Som e

Species Referred to

Leptonereis, Nicon,

and Laeonereis

(Polychaeta: Nereididae)

Introduction

A collection of heteronereids, found swarming in

Tampa Bay, Florida, were sent to me for identification by Mr. John L. Taylor. In the process of attempting to identify them and comparing them with

other nereid species, it appeared to me that a general

review and revision of some of the genera and species

of the family were needed. There has been no general agreement among polychaete systematists regarding the nereid genera Leptonereis, Nicon, and

Laeonereis, as indicated by the following brief review.

Kinberg (1866:178) established three genera in

a new family Niconidea, all characterized by the

absence of paragnaths or soft papillae on the pharynx or proboscis. They were separated by differences

in the parapodia: Nicon (six species), with the

parapodia remaining uniform along the body, Leptonereis (single species), with the parapodia changing gradually, and Nicomedes (single species), with

the parapodia changing abruptly and divided into

three regions. Of the eight species, which were described very briefly by Kinberg, only Leptonereis

laevis was figured in a subsequent publication. These

differences were considered by Claparede (1870:454)

to be of secondary value or associated with the sexual

stages (i.e., heteronereids) and for this reason he

selected Leptonereis (including Nicon and Nicomedes) to be retained. He ranked it as a subgenus

of Nereis, emended it, and included within it a new

Marian H. Pettibone, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560

species, Nereis (Leptonereis) glauca, from the Mediterranean Sea.

The description of Claparede's species was supplemented by Ramsay (1914a: 245), who observed minute papillae on the pharynx. The genus Leptonereis

thus was further emended, resulting in a diagnosis

that applied to L. glauca but not to L. laevis, the

type-species of Leptonereis. Subsequently, Leptonereis

generally was used to include nereids in which the

pharynx lacks paragnaths and with soft papillae

present or absent. This resulted in a heterogeneous

grouping of species, placing together nereids having

little affinity with one another or with the typespecies, as pointed out by Hart man (1945:22).

Some of these species were referred to two new

genera: Laeonereis Hartman (1945:22; 1949:56)

and Australonereis Hartman (1954b: 19).

Based on examination of the type-specimens of

Kinberg, Hartman (1949) proposed that Leptonereis

Kinberg include only the type-species, L. laevis. She

found the type-species of Nicomedes, N. difficilis

Kinberg, to be indeterminable and therefore rejected

the genus. Nicon Kinberg was retained by her and

N. pictus was selected, from the six species included

in the genus by Kinberg, as the type-species. Unfortunately, figures were not added nor was a clear

generic diagnosis included.

After 1949, eight new species were added to Nicon

by Knox (1951), Hartman (1953, 1958, 1965b,

1967) and Berkeley and Berkeley (1964). Two new

species were added to Laeonereis by Day (1957) and

Hartmann-Schroder (1959). Two new species were

added to Leptonereis by Rullier (1963) and Fishelson and Rullier (1969). Nicon has come in turn to
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

include a heterogeneous grouping of species. Some

of the species have been bandied about among the

various genera, resulting in considerable confusion.

As shown in the list of species summarized on

pages 2 to 5, there has been no general agreement

regarding the composition of the genera Leptonereis,

Nicon, and Laeonereis. The emphasis on the presence or absence of papillae on the pharynx has

caused some of the difficulty. The papillae are sometimes difficult to detect, particularly on small specimens in which the pharynx is not extended. Some

of the specimens examined were found to have minute denticles or pectinae on the pharynx, characteristic of Platynereis. Other sources of confusion have

been incomplete or incorrect descriptions, inadequate

figures, and confused terminology.

The purpose of this study, then, has been to review the various species that have been assigned

either originally or subsequently to Leptonereis, Nicon, or Laeonereis, to seek additional characters that

would serve to separate them in more meaningful

ways, to supplement the original descriptions and

figures, to clarify the terminology, and to include

comparable generic diagnoses.

In addition to the abundant collections deposited

in the Smithsonian Institution (USNM), some nereid

material was available for study from the following

Museums: Allan Hancock Foundation, Los Angeles

(AHF), through K. Fauchald; American Museum

of Natural History, New York (AMNH), through

H. S. Feinberg; British Museum (Natural History),

London (BMNH), through J. D. George; Museum

National d'Histoire Naturelle, Paris (MNHNP).

through J. Renaud-Mornant; Naturhistoriska Riksmuseet, Stockholm (NRS), through R. Olerod;

Universitetets Zoologiske Museum, Copenhagen

(UZMC), through J. B. Kirkegaard; Zoologisches

Museum, Berlin (ZMB), through G. Hartwich;

Zoologisches Staats-museum, Hamburg (ZMH),

through G. Hartmann-Schroder. Additional type

material was obtained from the private collections

of Professor F. Rullier at Angers, France, and from

the Department of Zoology, Tel-Aviv University, Israel, through Ch. Lewinsohn. Additional nereids

were sent in exchange by G. A. Knox from the University of Canterbury, New Zealand. Information

concerning type-material deposited in the Museum

of Comparative Zoology, Harvard, was obtained from

H. W. Levi.

I take this opportunity to thank the above-mentioned individuals for their cooperation and help in

arranging for the loans or exchanges of the nereid

material on which this study is based. The manuscript benefited from the suggestions of Meredith L.

Jones and Horton H. Hobbs, Jr., both of the Smithsonian Institution.

Of the 47 nereid species reviewed, 17 were

originally assigned to Leptonereis [or Nereis (Leptonereis)], 15 to Nicon, three each to Nereis and

Tylorrhynchus, two each to Leonnates [or Nereis

(Leonnates)], Ceratocephale, and Laeonereis, and

one each to Nicomedes, Chinonereis, and Stenoninereis.

The species of Nereididae, treated in this report,

are listed below, along with indications of their typelocalities, locations of the type-specimens (when

known), and summaries of their subsequent and

present designations.

SUBSEQUENT AND PRESENT

ORIGINAL DESIGNATION DESIGNATIONS

Nereis Linnaeus, 1758:

1. N. heterocheta Quatre- Holotype examined and refages, 1865:552. Java. ferred to Tylorrhynchus by

Holotype in Museum Grube (1870:312). Referred

National d'Histoire Na- to T. chinensis by Ramsay

turelle, Paris. (1914b:231). Referred to

T. heterochaetus [sic] by

Gravier and Dantan (1932:

671), see page 44.

2. N. culveri Webster, 1880: Referred to Leptonereis by

111. New Jersey and Fauvel (1923b: 124). ReNorth Carolina, eastern ferred to Laeonereis by HartNorth America. Syn- man (1945:21), see page

types in United States 14.

National Museum (US

NM 541, 28178) and

British Museum (Natural History) (BMNHZB

1971.30).

3. N. tridentata Webster, Referred to Ceratonereis by

1880:142. New Jersey, Hartman (1945:21). Specieastern North America. mens from Georgia and

Type-specimens no long- North Carolina examined;

er exist designated herein as typespecies of Websterinereis new

genus, see page 21.

Leonnates Kinberg, 1866 or

Nereis (Leonnates):

1. L. pusillus Langerhans, Questionably referred to Lep1880:279. Madeira. tonereis glauca by Ramsay

(1914a:251); same, without question, by Fauvel

(1914:163). Referred to

Laeonereis glauca by Hart-
NUMBER 10 4

ORIGINAL DESIGNATION

2. N. (L.) ehlersi Augener,

1913:142. Australia.

Leptonereis Kinberg 1866, or

Nereis (Leptonereis):

1. L. laevis Kinberg, 1866:

179. Guayaquil, Ecuador. Syntypes in Naturhistoriska Riksmuseet,

Stockholm (not found

in 1970).

2. N. (L.) glauca Claparede, 1870:454. Gulf of

Naples.

3. N. (L.) cebuensis Grube,

1878:61. Philippine Islands. Syntypes in Zoologisches Museum, Berlin

(ZMB Q3465).

4. N. (L.) vaillanti SaintJoseph, 1888:246.

France.

5. L. vasculosa Giard, 1890:

78. France.

6. N. (L.) inermis Hoagland, 1920:608. Philippine Islands. Holotype

in United States National Museum (USNM

18947).

7. N. (L.) acuta Treadwell, 1923:1237. Brazil.

Paratypes in United

States National Museum

(USNM 19030).

SUBSEQUENT AND PRESENT

DESIGNATIONS

man (1959a: 244). Referred

herein to Websterinereis

glauca, see page 27.

Referred to Leptonereis by

Monro (1938:622). Referred to Laeonereis by Hartman (1949:56). Referred to

Australonereis by Hartinan

(1954b:19).

Type-species of Leptonereis

Kinberg, 1866, by monotypy.

Syntypes examined by Hartman (1945, 1949), see page

6.

Referred to Leptonereis by

Ramsay (1914a: 244). Referred to Laeonereis by Hartman (1945:22). Specimens

from England and France examined (BMNH, MNHNP,

USNM) and referred herein

to Websterinereis, see page

27.

Referred to Laeonereis by

Hartman (1959a: 243, 262).

Syntypes examined and referred herein to Platynereis,

see page 46.

Referred to N. (Leonnates)

pustilus by Augener (1910:

236). Referred to Leptonereis glauca by Ramsay

(1914a:245) and Fauvel

(1914:163).

Referred to Laeonereis glauca by Hartman (1959a: 245).

Specimen from France, identified by Saint-Joseph, examined (MNHNP) and referred herein to Websterinereis

glauca, see page 27.

Nomen nudum.

Possibly Nicon by Hartman

(1949:55). Referred to Laeonereis by Hartman (1959a:

244, 262). Designated as

type-species of Kinberginereis new genus, see page 30.

Referred to Leptonereis culveri by Hartman (1938:14).

Referred to Laeonereis culveri by Hartman (1945:21),

see page 14.

ORIGINAL DESIGNATION

8. L. egregicirrata Treadwell, 1924:13. Antigua,

West Indies. Syntypes

in United States National Museum (USNM

20324).

9. L. foli Fauvel, 1930:520.

New Caledonia. Holotype in Museum National d'Histoire Naturelle,

Paris (MNHNP).

10. N. (L.) distorta Treadwell, 1936:273. China.

Holotype in United

States National Museum

(USNM 20118).

11. L. pandoensis Monro,

1937:242. Uruguay

(freshwater). Syntypes in

British Museum (Natural

History) (BMNH 1937:

4:22: 4-30) and United States National Museum (USNM 43474).

12. L. glauca moniloceras

Hartman, 1940:217.

Southern California and

western Mexico.

13. L. nota Treadwell, 1941:

1. Texas, Gulf of Mexico. Holotype in American Museum Natural

History (AMNH 2896).

14. L. mexicana Treadwell,

1942:1. Lower California, western Mexico. Holotype and paratype in

American Museum Natural History (AMNH

3222).

15. L. punctata WesenbergLund, 1949:289. Persian Gulf. Syntypes in

Universitetets Zoologiske

Museum, Copenhagen

(UZMC).

16. L. zebra Rullier, 1963:

474. Mauritius, Indian

Ocean. Syntypes in collection of F. Rullier

(No. 16-10, W 92).

17. L. davidi Fishelson and

SUBSEQUENT AND PRESENT

DESIGNATIONS

Referred to Nereis by Treadwell (1939:233). Redescribed by Pettibone (1956:284):

syntypes considered to include two species, N. egregicirrata (Treadwell) and N.

allenae Pettibone (1956:

287).

Referred to Laeonereis by

Hartman (1949:55). Holotype examined and referred

herein to Websterinereis, see

page 23.

Considered questionable by

Hartman (1956:255). Referred herein to Tylorrhynchus heterochetus, see page

44.

Referred to Laeonereis by

Hartman (1959a: 245). Syntypes examined and referred

to Laeonereis culveri, see

page 14.

Referred to Nicon moniloceras by Hartman (1958:

265). Referred herein questionably to Platynereis, see

page 48.

Referred to Laeonereis culveri by Hartman (1951:44).

Holotype examined, see page

14.

Types examined and referred to Nicon by Hartman

(1956:253, 274, 279). Perhaps N. moniloceras by Hartman (1959a:245). Types

examined and referred herein to Rullierinereis, see page

35.

Referred to Nicon by Hartman (1958:265). Syntypes

examined and referred herein to Websterinereis, see page

25.

Perhaps Nicon by Hartman

(1965a: 36). Syntypes examined and designated herein

as type-species of Rullierinereis new genus, see page

33.

Holotype examined and re-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ORIGINAL DESIGNATION

Rullier, 1969:62. Red

Sea. Holotype in Department of Zoology,

Tel-Aviv University, Israel (No. E 62/238).

Nicomedes Kinberg 1866:

1. N. difficilis Kinberg,

1866:179. Rio de Janeiro, Brazil. Holotype in

Naturhistoriska Riksmuseet, Stockholm (NRS

572).

Nicon Kinberg, 1866:

1. N. eugeniae Kinberg,

1866:178. Magella n

Strait, South America.

Type-specimens in Naturhistoriska Riksmuseet,

Stockholm.

2. N. loxechini Kinberg,

1866:179. Magella n

Strait, South America.

Holotype in Naturhistoriska Riksmuseet, Stockholm.

3. N. maculata Kinberg,

1866:178. La Plata, Argentina. Holotype in

Naturhistoriska Riksmuseet, Stockholm (NRS

493).

4. N.pictus Kinberg, 1866:

178. Brazil. Syntypes in

Naturhistoriska Riksmuseet, Stockholm (NRS

486).

5. .V. tahitanus Kinberg,

1866:178. Tahiti. Holotype in Naturhistoriska

Riksmuseet, Stockholm

(NRS 494).

6. AT. virgini Kinberg, 1866:

179. Magellan Strait,

South America. Holotype in Naturhistoriska

Riksmuseet, Stockholm

(NRS 576).

7. N. aestuariensis Knox,

1951:225. New Zealand.

Holotype in Canterbury

Museum, New Zealand.

8. N. ehlersi Hartman,

1953:29. Falkland Islands. Syntypes in Naturhistoriska Riksmuseet,

Stockholm (NRS 3863).

SUBSEQUENT AND PRESENT

DESIGNATIONS

ferred herein to Platynereis

species, see page 49.

Type-species of Nicomedes

Kinberg, 1866, by monotypy.

Holotype examined by Hartman (1949:57) and considered to be indeterminable.

Holotype examined, see page

49.

Referred to Nereis by Ehlers

(1897:67) and Hartman

(1949:59).

Holotype examined by Hartman (1949:61) and referred

to Platynereis magalhaensis

Kinberg, 1866.

Holotype examined by Hartman (1949:58). Holotype

examined and selected herein as type-species of Nicon,

see pages 7, 8.

Syntypes examined by Hartman (1949:57) and selected as type-species of Nicon.

Syntypes examined and considered to be indeterminable,

see page 49.

Holotype examined by Hartman (1949:59). Holotype

examined and considered to

be indeterminable, see page

49.

Holotype examined by Hartman (1949:59). Holotype

examined and considered to

be indeterminable, see page

49.

Specimens from type-locality

examined (USNM 43475,

from G. A. Knox), see page

11.

Syntypes examined and referred to N. maculata, see

page 8.

ORIGINAL DESIGNATION

9. N. lackeyi Hartman,

1958:263. Florida. Holotypes in United States

National Museum (US

NM 29627-8).

10. N. peruviana Berkeley

and Berkeley, 1964:128.

Off Peru, pelagic. Holotype and paratype in

United States National

Museum (USNM 32821-

2).

11. A', uncinatus Hartman,

1965b: 86. Off New England.

12. N. abyssalis Hartman,

1967:66. Antarctic.

13. N. benhami Hartman,

1967:66. Antarctic.

14. N. polaris Hartman,

1967:68. Antarctic.

15. AT. maculatus.—Gallardo, 1968:65. South Viet

Nam. In Allan Hancock Foundation (AHF

0252).

Ceratocephale Malmgren,

1867:

1. C. osawai Izuka, 1903:

1. Japan.

2. C. corallicola Reish,

1968:215. Eniwetok,

Marshall Islands. Holotype and paratypes in

United States National

Museum (USNM 38-

392-3).

Tylorrhynchus Grube, 1868:

1. T. chinensis Grube ,

1868:22. China (Shanghai).

2. T. sinensis.—Dawydoff,

1952:88. Indochina.

SUBSEQUENT AND PRESENT

DESIGNATIONS

Referred herein to Stenoninereis martini, see page 39.

Referred herein to Platynereis sp., see page 48.

Referred herein to Rullierinereis uncinata, see page 31.

Generic affinities doubtful.

Specimens from Benham's

record of Nereis loxechini

(BMNH 1928:2:29: 222-

223, 226-228) from McMurdo Sound, the basis of

A', benhami, examined and

referred herein to N. maculata, see page 8.

Generic affinities doubtful.

Referred herein to Rullierinereis gallardoi new species,

see page 39.

Referred to Tylorrhynchus

chinensis by Ramsay (1914b:

231). Referred to T. heterochaetus [sic] by Gravier and

Dantan (1932:671), see

page 44.

Incorrect genus, according

to Pettibone (1970:233).

Referred to Websterinereis

foli, see page 23.

Type-species of Tylorrhynchus Grube, 1868:22, by

monotypy. Referred to T.

heterochaetus [sic] by Gravier and Dantan (1932:671),

see page 44.

Error for T. chinensis.—

Hartman (1959a: 280), see

page 44.
NUMBER 10 4

ORIGINAL DESIGNATION

2. T. bahamensis

mann-Schroder,

Hart1958:

SUBSEQUENT AND PRESENT

DESIGNATIONS

Referred herein to Rullierinereis, see page 31.

Type-species of Chinonereis

Chamberlin, 1924, by original designation and monotypy. Referred herein to Tylorrhynchus heterochetus, see

page 44.

Referred herein to Websterinereis punctata, see page 25.

Type-specimens examined

and referred herein to Laeonereis culvert, see page 14.

238. Bahamas.

Chinonereis Chamberlin, 1924:

1. C. edestus Chamberlin,

1924:81. China (Canton), freshwater. Holotype not present in Museum of Comparative

Zoology, Harvard.

Laeonereis Hartman, 1945:

1. L. ankyloseta Day, 1957:

83. South Africa.

2. L. brunnea HartmannSchroder, 1959:135. El

Salvador, Central America. Holotype and paratypes in Zoologisches

Staats-museum, Hamburg (ZMH 143).

Stenoinereis Wesenberg-Lund,

1958:

1. S. martini WesenbergLund, 1958:9. West Indies (St. Martin). Syntypes in United States

National Museum (US

NM 29726).

Of the 47 species listed above, the following seven

are not discussed further:

Leptonereis egregicirrata Treadwell [Referred to Nereis egregicirrata and N. allenae by Pettibone

(1956)]

Leptonereis vasculosa Giard [nomen nudum]

Nereis (Leonnates) ehlersi Augener [Referred to

Australonereis by Hartman (1945b)]

Nicon eugeniae Kinberg [Referred to Nereis by Ehlers

(1897)]

Type-species of Stenoninereis Wesenberg-Lund, 1958,

by monotypy, see page 39.

Nicon loxechini Kinberg [Referred to Platynereis

magalhaensis by Hartman (1949)]

Nicon abyssalis Hartman [Generic affinities doubtful]

Nicon polaris Hartman [Generic affinities doubtful]

The remaining 40 species are referred to Leptonereis Kinberg (single species), Nicon Kinberg,

emended (two species, two synonyms), Laeonereis

Hartman, emended (single species, four synonyms),

Websterinereis new genus (four species, four synonyms) , Kinberginereis new genus (single species),

Rullierinereis new genus (five species, one of them

new), Stenoninereis Wesenberg-Lund (single species,

one synonym), Tylorrhynchus Grube (single species,

five synonyms), Platynereis Kinberg (four species),

and Indeterminable Nereididae (four species).

All of the genera discussed in this report

{Leptonereis Kinberg, Nicon Kinberg, Laeonereis

Hartman, Websterinereis new genus, Rullierinereis

new genus, Stenoninereis Wesenberg-Lund, Tylorrhynchus Grube, and Kinberginereis new genus) show

a number of characters in common, among which are

the following. The prostomium is subpyriform, with

paired frontal antennae, paired biarticulate palps,

and usually two pairs of eyes (sometimes three pairs;

sometimes lacking?). The tentacular segment is

achaetous and apodous, bearing four pairs of tentacular cirri with distinct cirrophores. The first two

setigerous segments have subbiramous parapodia, the

notopodia lacking notosetae; the remaining segments

have biramous parapodia. The ventral cirri are short,

tapered, and single (not double). The pharynx lacks

typical paragnaths (with or without soft papillae).

The pygidium is provided with a pair of anal cirri.

The eight genera may be separated according to the

following Key.

Key to the Genera

1. Dorsal cirri at base of upper notopodial ligules; neuropodia widi lower ligules (Figures la,

b; 6e,f; I7d,e; \Bc,d) 3

1'. Dorsal cirri with large or elongate cirrophores and distal styles; neuropodia without lower

ligules (Figures 23A,f; 25c,d) 2

2. Notopodia bilobed, with lower acicular lobes continuous with presetal distal lobes and upper

notopodial ligules (latter diminishing posteriorly) ; neuropodia d is tally with diagonal anterior acicular lobes and shorter postsetal lobes (Figure 23h,i,n). Notosetae homogomph

spinigers (Figure 23;). Neurosetae arranged in fan-shaped bundles and composed of homogomph and heterogomph spinigers and heterogomph falcigers (Figure 23/t-m). Pharynx

without papillae Stenoninereis Wesenberg-Lund

2'. Notopodia with low rounded lobes and subacicular ligules; neuropodia thick, bilobed distally with neuroaciculum between (Figure 25c-e). Notosetae hemigomph spinigers (Figure

25/), Neurosetae nearly encircling bilobed neuropodial lobes, arranged in form of "3 " and
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

composed of hemigomph and heterogomph spinigers and heterogomph falcigers (Figure

25c-i). Pharynx with soft papillae on both maxillary and oral rings .... Tylorrhynckus Grube

3. Notosetae homogomph spinigers and homogomph falcigers posteriorly (Figure 18g,;). [Notopodia with two notopodial ligules (Figure I8c,d). Pharynx without papillae].

Rullierinereis new genus

3'. Notosetae only homogomph spinigers (Figures 2ft; 6g) 4

4. Neurosetae all spinigerous—homogomph and hemigomph, without falcigers (Figure 17ft-ifc).

[Neuropodia with bilobed presetal lobes and short rounded postsetal lobes in addition to

lower ligules (Figure 17a» ] Kinberginereis new genus

4'. With some neuropodial falcigers in addition to spinigers (Figures 2»; 6ft) 5

5. Neurosetae only homogomph spinigers and homogomph falcigers with rather long blades

(Figure 6g,h). Dorsal cirri very small—shorter than notopodial ligules (Figure 66-/).

Pharynx with tufts of papillae on maxillary ring (Figure 5c-e,g). [Notopodia with presetal

lobe in addition to two notopodial ligules and neuropodia with presetal and postsetal lobes

in anterior region; ventral cirri very short (Figures 6d-f)] Laeonereis Hartman

5'. Some neurosetae hetevogomph falcigers with relatively long blades (Figures 2i, 5/). Dorsal

cirri longer than notopodial ligules (Figure 2b—e). Pharynx without papillae on maxillary

ring 6

6. Pharynx with papillae on basal or oral ring: pair of larger papillae on area VI, single row

of small papillae on areas VII-VIII (Figures 8a,b; life; 12a; \6a,b).

Websterinereis new genus

6'. Pharynx without papillae 7

7. Upper notopodial ligules in middle and posterior segments large, scale-like, with minute

terminal dorsal cirri. Neuropodia without postsetal lobes Leptonereis Kinberg

T. Upper notopodial ligules not enlarged, scale-like. Neuropodia with digitiform to conical

postsetal lobes (Figures 2d,f,j; 3/) Nicon Kinberg

Abbreviations Used in the Figures

I-VIII, areas of pharynx

acL, acicular lobe

aL, anterior lobe

an, anus

anC, anal cirrus

cph, cirrophore

dC, dorsal cirrus

dLa, dorsal lamella

dLo, dorsal lobe

fAn, frontal antenna

mR, maxillary or distal ring of pharynx

neLi, neuropodal ligule

noLi, notopodial ligule

oR, oral or basal ring of pharynx

pa, palp

pGl, parapodial gland

pL, posterior lobe

poL, postsetal lobe

poLa, postsetal lamella

prL, presetal lobe

tC, tentacular cirrus

tS, tentacular segment

vC, ventral cirrus

vLa, ventral lamella

Leptonereis Kinberg, 1866; emended Hartman, 1945

TYPE-SPECIES.—L. laevis Kinberg, 1866, by monotype. Gender: feminine.

REMARKS.—Leptonereis was proposed to receive a

single species, L. laevis Kinberg, from Guayaquil,

Ecuador. The original description was very sketchy.

Subsequently, figures were added by Kinberg (1910,

pi. 20: fig. 7), including dorsal and ventral views

of the anterior end with the pharynx extended, a

parapodium from setiger 10, and upper and lower

neurosetae. Based on examination of the syntypes,

deposited in the Naturhistoriska Riksmuseet, Stockholm, Hartman (1945, 1949) supplemented the description and included figures of anterior and posterior parapodia, the latter showing the characteristic

greatly enlarged upper notopodial ligule with a minute terminal dorsal cirrus (pi. 3: figs. 5, 6, in Hartman, 1945). Hartman also pointed out that the types

were atokous individuals and not epitokes, as Grube

(1878:62) had supposed, and that L. laevis was the

only known nereid which could be referred to

Leptonereis, as defined in her revision of the genus.

Unfortunately, the syntypes could not be found recently in the Stockholm Museum, according to Mr.

Olerod (in correspondence), and were not available
NUMBER 10 4

for the present study. The following diagnosis is based

on the descriptions and figures given by Kinberg

(1866:179; 1910:53, pi. 20: fig. 7) and Hartman

(1945:21, pi. 3: figs. 5, 6; 1949:56, pi. 8: figs. 5, 6).

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs

of eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Dorsal cirri at bases of upper notopodial ligules in

anterior region. Notopodia with two ligules, upper

one gradually increasing in size, becoming greatly

enlarged, oval, scalelike, with minute dorsal cirri

inserted terminally in middle and posterior parapodia.

Neuropodia with setigerous lobe and lower ligule.

Ventral cirri short, tapered. Notosetae [homogomph?]

spinigers only. Neurosetae homogomph [?] and heterogomph spinigers and heterogomph falcigers with

rather long blades. Pharynx with paired jaws, without paragnaths or papillae.

Following Hartman (1945, 1949), Leptonereis Kinberg is considered to include the single type-species:

L. laevis Kinberg from Ecuador. The other 16 species

(pages 3-4), described originally under Nereis

(Leptonereis) or Leptonereis, are referred to seven

other genera, including Kinberginereis (1) Laeonereis (3) , Nereis (1) , Platynereis (3) , Rullierinereis

(2) , Tylorrhynchus (1) , Websterinereis (4) , and one

nomen nudum.

Nicon Kinberg, 1866; emended

TYPE-SPECIES.—N. maculata Kinberg, 1866, selected herein. Gender: feminine.

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs of

eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Parapodia of first two setigers subbiramous, notopodium represented by dorsal cirrus and single ligule.

Dorsal cirri at bases of upper notopodial ligules.

Notopodia with two ligules and small supraacicular

or presetal lobe between them. Neuropodia with

conical or bilobed acicular lobes, digitiform or conical

postsetal lobes, and lower ligules. Ventral cirri short,

tapered. Notosetae homogomph spinigers only. Neurosetae homogomph spinigers, heterogomph falcigers,

and with or without heterogomph spinigers. Pygidium

with paired anal cirri. Pharynx with paired jaws,

without paragnaths or papillae.

REMARKS.—Nicon Kinberg was established for six

new species: N. eugeniae, N. loxechini, and N. virgini

from the Magellan Strait, N. maculata from Argentina, N. pictus from Brazil, and N. tahitanus from

Tahiti. The species were only briefly described and

figures were lacking. No type-species for Nicon was

designated. The genus was characterized by the

pharynx lacking paragnaths and papillae and by

having parapodia uniform along the body. Claparede

(1870:454) placed Nicon under Leptonereis Kinberg. Nicon eugeniae was referred to Nereis by

Ehlers (1897:67) and has been so recorded by

various polychaete workers. Nicon loxechini was subsequently assigned by Ehlers (1908:73) and Benham

(1921:65) to Nereis, and by Monro (1930:107) to

Leptonereis. As pointed out by Hartman (1949:61),

however, these records are confused and are referred

herein to Nicon maculata. Nicon loxechini Kinberg

was referred to Platynereis magalhaensis Kinberg

by Hartman (1949:61). The other four species were

not recorded in the literature until 1949 when Hartman examined Kinberg*s type-specimens, supplemented their descriptions to some extent and, for

some of them, added figures. For Nicon Kinberg,

Hartman selected N. pictus as the type-species without, however, figuring it or giving a clear diagnosis.

In order to clarify its identity, I examined the

syntypes of N. pictus. As indicated on page 49, I

found the specimens to be in such poor condition that

no adequate diagnosis could be prepared. This was

true also for the holotypes of N. tahitanus and N.

virgini. Thus, all three species are to be considered

indeterminable. The syntypes of Nicon maculata are

better preserved, and, for this reason, I have selected

N. maculata as the type-species of Nicon. The diagnosis of the genus and the redescription of the typespecies are based on the syntypes of N. maculata, as

well as on additional material that was compared

with the syntypes.

In addition to the 15 species originally assigned

to Nicon Kinberg (page 4), three additional species

were referred to it by Hartman (1958:265), including the following:

1. N. moniloceras (Hartman, 1940), as Leptonereis

glauca moniloceras. Southern California and

western Mexico.

2. N. mexicana (Treadwell, 1942), as Leptonereis.

Lower California, western Mexico.

3. N. punctata (Wesenberg-Lund, 1949), as Leptonereis. Persian Gulf.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Of the 18 species, which were originally placed in

Nicon or subsequently referred to it, four species are

retained in the genus as revised, including N. maculata (with synonyms N. ehlersi and N. benhami) and

other genera, including Nereis (1), Platynereis (3),

Rullierinereis (3), Stenoninereis (1) and Websterinereis (1); three species are considered to be indeterminable and two species to have doubtful generic

N. aestuariensis; nine species are referred to five affinities (N. abyssalis, N. polaris).

Key to the Species of Nicon

Neuropodial ligulcs of anterior region large, thick, rounded; neuropodia with conical acicular and digit if orm postsetal lobes (Figure 2b—g). Neurosetae homogomph spinigers and

heterogomph falcigers only (Figure 2h,i) N. maculata

Neuropodial ligules of anterior region conical; neuropodia with bilobed anterior and conical

postsetal lobes (Figure 3d-g). Neurosetae heterogomph spinigers in addition to homogomph

spinigers and heterogomph falcigers (Figure 3t—m) N. aestuariensis

Nicon maculata Kinberg

FIGURES 1, 2

Nicon maculata Kinberg, 1866:178.

Nereis loxechini.—Ehlers, 1908:73, pi. 8: figs. 8-12.—Benham 1921:65, pi. 8: figs. 67-75; 1927:50. [Not Nicon

loxechini Kinberg, 1866]

Not Leptonereis maculata.—Treadwell 1928:469. [ = Rullierinereis mexicana (Treadwell)]

Leptonereis loxechini.—Monto 1930:107; 1936:138. [Not

Nicon loxechini Kinberg 1866]

Nicon maculatus.—Hartman 1949:58.—Not Gallardo 1968:

65. [ = Rullierinereis gallardoi new species]

Nicon ehlersi Hartman, 1953:29.

Nicon benhami Hartman, 1967:66, pi. 18: figs. A-C.

MATERIAL EXAMINED.—Off La Plata, Argentina,

Eugenie Expedition—holotype of Nicon maculata

(NRS 493). ANTARCTIC: Falkland Islands, Burdwood Bank, 53°45'S, 61°10'W, shell fragments and

stones, 137-150 meters, Swedish Antarctic Expedition, station 59, 12 September 1902—3 syntypes of

Nicon ehlersi (NRS 3863).

Off Falkland Islands: 49°S, 62°W, sand, gravel,

stones, 146 meters, Discovery Expedition, station WS

97, 18 April 1927—2 specimens (USNM 43445, from

BMNH); 49°S, 59°14'W, fine dark sand, 251-225

meters, Discovery Expedition, station WS 99, 19

April 1927—1 specimen (USNM 43446, from

BMNH). [as Leptonereis loxechini, by C. Monro]

McMurdo Sound off Glacier tongue, about 8 miles

north of Hut Point, mud and undecomposed animal

remains, 347-457 meters, Terra Nova Expedition,

1910, station 316—1 specimen (BMNH 1928: 2: 29:

226-228). 5 miles north of Inaccessible Island, mud,

406-441 meters, Terra Nova Expedition, 1910, station 314—1 specimen (BMNH 1928: 2: 29:

222-223). [as Nereis loxechini Kinberg by W. B.

Benham]

McMurdo Sound near Cape Hallett, 72°05.8'S,

172°15.2'E, 392 meters, Deep Freeze III, USS

"Atka," station 23, 12 January 1958—16 specimens

(USNM 43395). McMurdo Sound east of Cape Hallett, 72°08'S, 172°10'E, 434 meters, Deep Freeze III,

USS "Burton Island," station 3, 13 January 1958—

2 specimens (USNM 43396). Davis Sea, 66°32.84'S,

93°90'E, 146 meters, Deep Freeze III, USS "Burton

Island," station 5, 29 January 1958—1 specimen

(USNM 43394). Nansen Island, Palmer Peninsula,

66°20'S, 67°47'W, 326 meters, Deep Freeze IV, USS

"Edisto," station 31, 24 March 1959—3 specimens

(USNM 43393).

TYPE-MATERIAL.—The holotype of Nicon maculata

(NRS 493) consists of three fragments, an anterior

one of 8 setigers, a middle one of 19 setigers, and a

posterior one of about 25 setigers with a narrow

posterior growth zone, with a total length of about

15 mm and a width of 1.5 mm, including setae. The

pharynx was extended and the jaws had been removed and are now missing, as noted by Hartman

(1949:58) when she examined the type.

The three syntypes of Nicon ehlersi Hartman (NRS

3863) consist of a larger anterior fragment of 47

setigers, 62 mm in length and 8 mm in width, including setae, and two smaller specimens, one of

which is complete. The pharynx was not extended but

two of the syntypes had been slit open, revealing the

absence of paragnaths and papillae.

DESCRIPTION.—Length up to 120 mm, width up to

10 mm, including setae, segments up to 140. Prostomium (Figure 2a) with more or less conspicuous

middorsal depression on anterior half, with tapered
NUMBER 10 4

frontal antennae, stout biarticulate palps; eyes large,

anterior pair larger than posterior. Tentacular segment nearly twice length of following segment; tentacular cirri of different lengths, longest extending

to setiger 5—9, sometimes indistinctly annulated

distally. Parapodia of first two setigers (Figure 2b,c)

with dorsal cirrus and single notopodial ligule; neuropodia similar to those of following setigers; neurosetae homogomph spinigers and heterogomph falcigers.

Parapodia of anterior region (Figures \a,b) 2d,e)

with cirriform dorsal cirri extending beyond parapodial ligules, two bluntly conical notopodial ligules

and more or less developed small supraacicular lobe

between them; neuropodium with conical acicular

lobe and digitiform postsetal lobe (ovate process referred to and figured by Benham, 1921, on pi. 8: figs.

69, 70, 72), thick rounded to bullet-shaped lower

neuropodial ligule; ventral cirri short, tapered. Notosetae all homogomph spinigers, blades spinous, tapering rather abruptly to slender tips (Figures \d;2h).

Both upper and lower bundles of neurosetae homogomph spinigers similar to notosetae and heterogomph falcigers with rather long blades (Figures \c\

2i\ no heterogomph spinigers). Parapodia of middle

(Figure 2f,g) and posterior (Figure 2)) regions with

rami more widely separated; dorsal cirri becoming

more elongate posteriorly; notopodial ligules more

sharply conical; postsetal neuropodial lobes less conspicuous than in anterior region; lower neuropodial

ligules smaller, subconical. Setae similar to anterior

region except heterogomph falcigerous neurosetae

of upper bundles much stouter, darker, and with

shorter blades (Figure 2k; usually two in number);

blades often missing and stems somewhat worn, appearing as simple stout spines (Figure 2/).

Pygidium prominent, with long anal cirri. Pharynx

lacking paragnaths or papillae; jaws dark ambercolored, each with about 16 oblique teeth along cutting edge (pi. 7: fig. 8, in Ehlers, 1908; pi. 8: fig. 75,

in Benham, 1921).

REMARKS.—The species is characterized by the

absence of paragnaths and papillae on the pharynx,

the relatively large eyes, the conical or digitiform

postsetal neuropodial lobes, the relatively large, thick

lower neuropodial ligules, especially in the anterior

region, and the neurosetae composed of only homogomph spinigers and heterogomph falcigers [no heterogomph spinigers, as stated by Monro (1936:139)

and Hartman (1949:58)]. In the middle and posterior parapodia, a few neuropodial heterogomph

falcigers in the upper bundles are much thicker than

the others; the blades are often broken off, thus

appearing as simple setae.

Nicon loxechini Kinberg was found by Hartman

(1949:61) to have denticles on the pharynx and was

FIGURE 1.—Nicon maculata (holotype of iV. maculata, NRS 493): a, Parapodium from anterior region, posterior view; b, same, anterior view; c, neuropodial heterogomph falciger from

same; d, homogomph spiniger from same. [Figures c, d, not to scale]
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Nicon maculata (USNM 43394): a, Anterior end, dorsal view; b, first parapodium,

posterior view; c, second parapodium, anterior view; d, parapodium from anterior region, posterior view; e, same, anterior view; /, parapodium from middle region, posterior view; g, same,

anterior view; h, notopodial homogomph spiniger from same; i, neuropodial heterogomph falciger from same; ;, parapodium from posterior region, posterior view; k, stout heterogomph

falciger from upper neuropodial bundle of same; /, same, with blade broken off and stem

somewhat worn. [Figures h, i, k, I, not to scale]

fAn
NUMBER 10 4 11

referred to Platynereis magalhaensis Kinberg. The

specimens described and figured by Ehlers (1908)

and Benham (1921), as Nereis loxechini, were not

the same as Kinberg's species. Hartman (1953) gave

the new name Nicon ehlersi for Ehlers' record and

included Benham's record. Later, Hartman (1967)

proposed the new name Nicon benhami for Benham's

record. The presence of thick supraacicular heterogomph falcigers in the neuropodia of N. benhami

was given as one of the distinguishing characters

between N. benhami and N. ehlersi. However, examination of the syntypes of the latter species revealed their presence and both species are herein

considered to be synonyms of N. maculata. The other

distinguishing character mentioned by Hartman was

the presence of prolonged dorsal cirri in N. benhami.

These occur in the posterior parapodia of N. ehlersi

also. Two of the specimens from McMurdo Sound,

identified by Benham (1921) as iV. loxechini, were

examined (BMNH); they were found to agree with

N. maculata, as described herein.

Gravid females with no sign of epitoky have been

found. Monro (1930:108, as Leptonereis loxechini)

described a male heteronereid.

DISTRIBUTION.—Common Antarctic and subantarctic species. In 92 to 1153 meters.

Nicon aestuariensis Knox

FIGURES 3, 4

Nicon aestuariensis Knox, 1951:225, pi. 50: figs. 41-46.—

Estcourt 1966:179, figs. 1-4, pi. 1: figs. 1-3; 1967a: 70;

1967b:390.

MATERIAL EXAMINED.—Heathcote estuary, New

Zealand, I. N. Estcourt, collector, from G. A. Knox.—

16 atokous specimens and 1 partially metamorphosed

male heteronereid (USNM 43475).

DESCRIPTION.—Length up to 250 mm, width up

to 6 mm, segments numerous. Prostomium (Figure

3a,c) with faint middorsal depression on anterior

half, tapered frontal antennae, stout biarticulate

palps; eyes moderately large, anterior pair larger

than posterior (eyes much larger on heteronereids).

Tentacular segment nearly twice length of following

segment; tentacular cirri of different lengths, longest

extending to about setiger 5 (4—6). Parapodia of

first two setigers (Figure 3d,e) with dorsal cirrus and

single notopodial ligule; neuropodium similar to those

of following setigers; neurosetae homogomph and

heterogomph spinigers and heterogomph falcigers.

Parapodia of anterior region (Figure 3f-h) with

subulate dorsal cirri shorter than notopodial ligules,

two conical notopodial ligules, and shorter presetal

lobe between them; neuropodium with bilobed anterior lobe—smaller digitiform one above and larger

rounded one below neuroaciculum, longer conical

postsetal lobe, and conical lower neuropodial ligule;

ventral cirri short, subulate. Notosetae all homogomph spinigers, with blades finely spinous, shorter

(few upper ones, Figure 3i) and longer (Figure 3;).

Neurosetae arranged in S-shaped bundle nearly encircling bilobed anterior lobe (Figure 3h). Supraacicular bundle neurosetae homogomph spinigers with

long blades (posterior and upper part bundle, Figure

3;), heterogomph spinigers with short blades (anterior part bundle, Figure 3/), and heterogomph

falcigers with relatively long blades (anterior part

bundle, Figure 3m). Subacicular bundle of neurosetae heterogomph spinigers with long blades (posterior part bundle, Figure 3k), heterogomph spinigers

with short blades (lower and anterior part bundle,

Figure 3/), and heterogomph falcigers (anterior part

bundle, Figure 3m). Parapodia of posterior region

(Figure Ac,d) somewhat modified: dorsal cirri extending beyond notopodial lobes; presetal notopodial

and neuropodial lobes smaller; notopodial ligules

with conspicuous coiled glands; notosetae few homogomph spinigers; upper bundle neurosetae few,

slightly stouter heterogomph falcigers (Figure Ae);

lower bundle neurosetae few heterogomph spinigers.

Parapodia of middle region of slightly modified

male heteronereid (Figure 4a,b) showing early stages

of extra lamellae: dorsal lamellae medial to dorsal

cirri, large rounded neuropodial postsetal lamella

and ventral lamella on bases of ventral cirri; notosetae homogomph spinigers; upper bundle neurosetae homogomph spinigers with long blades and

heterogomph spinigers "with short blades; lower bundle neuropodia heterogomph spinigers with long and

short blades (usual type setae completely replaced

by swimming setae in fully developed heteronereids,

according to Knox, 1951: 226).

Pygidium with pair long anal cirri. Pharynx lacking

paragnaths or papillae, dark brown jaws, each with

about 10 oblique teeth along cutting edge (Figure

3a-c).

BIOLOGY.—N. aestuariensis is known from estuaries in New Zealand. Additional contributions to

its life history, breeding biology, and habits have
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Nicon aestuariensis (USNM 43475: a, small complete specimen; b, slightly larger

specimen; c-m, larger, partially metamorphosed male heteronereid): a, Anterior end, with

pharynx extended, dorsal view; b, same, ventral view; c, same, dorsal view; d, first parapodium,

posterior view; e, second parapodium, anterior view; /, parapodium from anterior region, posterior view; g, same, anterior view; h, diagrammatic end view of same showing arrangement

of setae; i, homogomph spiniger with relatively short blade; ;, same, with long blade; k, heterogomph spiniger with long blade; I, same, with relatively short blade; m, heterogomph falciger.

[Figures i—m, not to scale]
NUMBER 10 4 13

FIGURE 4.—Nicon aestuariensis (USNM 43475: a, b, partially metamorphosed male heteronereid; c-e, small complete specimen) : a, Parapodium from middle modified region, posterior

view; b, same, anterior view; c, parapodium from posterior region, posterior view; d, same,

anterior view; e, heterogomph falciger from same. [Figure e, not to scale]

been reported by Estcourt (1966:179; 1967a: 70;

1967b: 390). It burrows in the mud, where it is a

bottom-deposit feeder, and is able to tolerate wide

variations in salinity (from 0 to 29%o). It is one of the

most abundant animals in the upper reaches of Heathcote Estuary. N. aestuariensis is one of the few nereid

species that has become well adapted to the estuarine

environment that reproduces as epitokes and not in

the atokous stage. It appears to have a two-year life

cycle. At sexual maturity it forms heteronereids and

undergoes a typical swarming periodicity, spawning

at the surface. The spent adults die after spawning.

The eggs are moderate in size (about 150/x) and

demersal, developing into ciliated larvae. Early development presumably takes place in the surface

layers of the mud, the larvae being absent from the

plankton. Small specimens were found near the surface of the sediment, with large specimens occupying

burrows as deep as 40 centimeters.

DISTRIBUTION.—New Zealand.

Laeonereis Hartman, 1945; emended

TYPE-SPECIES.—Nereis culveri Webster, 1880, by

original designation. Gender: feminine.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs

of eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Parapodia of first two setigers subbiramous, notopodium represented by dorsal cirrus and single ligule.

Dorsal cirri at bases of upper notopodial ligules.

Notopodia with two ligules and additional presetal

lobe in anterior parapodia. Neuropodia with lower

ligules and additional presetal and postsetal lobes in

anterior parapodia. Ventral cirri very short. Notosetae

homogomph spinigers only. Neurosetae homogomph

spinigers and, more posteriorly, some homogomph

falcigers with rather long blades. Pygidium with

paired anal cirri. Pharynx with paired jaws, without

paragnaths; with tufts of papillae on maxillary or

distal ring and pair of large papillae on oral ring

(area VI).

REMARKS.—In addition to L. culvert (Webster)

as type-species, Hartman (1945:21) included Nereis

(Leptonereis) glauca Claparede (1870) in Laeonereis. The latter species is referred herein to Websterinereis. In the brief diagnosis of Laeonereis given by

Hartman, she indicated the presence of heterogomph

falcigerous setae. On the contrary, the only falcigers

present are homogomph, with rather long blades, as

shown in the figures of Webster (1886, pi. 7: fig.

32), Fauvel (1923b, fig. 5e), Monro (1937, fig. If),

and Hartmann-Schroder (1959, fig. 98). Also Hartman indicated that the soft papillae were arranged

in tufts on both the oral and maxillary rings of the

pharynx. As shown in the figure of Webster (1886,

pi. 6: fig. 24), the tufts of papillae are confined to

the maxillary ring; the oral ring has a pair of

triangular dorsal papillae (area VI).

Subsequently two additional species were added to

Laeonereis—L. ankyloseta from South Africa by Day

(1957) (referred herein to Websterinereis punctata)

and L. brunnea from El Salvador by HartmannSchroder (1959) (referred herein to L. culveri).

Nereis (Leptonereis) acuta Treadwell (1923) and

Leptonereis nota Treadwell (1941) were both synonymized with L. culveri by Hartman (1938:14;

1951:44). Four additional species were referred to

Laeonereis by Hartman (1959a, in Catalogue):

Nereis (Leptonereis) cebuensis Grube (1878) from

the Philippine Islands (referred herein to Platynereis), Nereis (Leptonereis) inermis Hoagland (1920)

from the Philippine Islands (referred herein to Kinberginereis), Leptonereis foli Fauvel (1930) from

New Caledonia (referred herein to Websterinereis),

and Leptonereis pandoensis Monro (1937) from

Uruguay (referred herein to Laeonereis culveri).

Laeonereis culveri (Webster)

FIGURES 5-7

Nereis culveri Webster, 1880:111 [plates referred to but not

published]; 1886:140, pi. 6: figs. 23-30, pi. 7: figs. 31,

32 [reprinting of 1880 with addition of figures].—Ferguson and Jones 1949:440.

Leptonereis culveri.—Fauvel 1923b: 124, fig. 5.—Hartman

1938:14.

Nereis (Leptonereis) acuta Treadwell, 1923:1237, figs. 1-5.

Leptonereis pandoensis Monro, 1937:242, fig. la-f.

Laeonereis culveri.—Hartman 1945:21; 1951:44, pi. 13:

figs. 3, 4; 1954a:415; 1956:253; 1959a:243, 245, 262;

1959b:539, 540.—Behre 1950:12.—Hedgpeth 1950:111.

—Carpenter 1956:93, 101.—Rioja 1958:257.—Frankenberg and Burbanck 1963:87.—Wells and Gray 1964:72.

—Oglesby 1965:621.—Mazurkiewicz 1969:1146; 1970:1,

figs. 1-27.

Leptonereis nota Treadwell, 1941:1, figs. 7-10.

Laeonereis brunnea Hartmann-Schroder, 1959:135, figs. 92-

99.

MATERIAL EXAMINED. — CONNECTICUT: Mystic

River estuary, 41°22'10"N, 71°59'42/'W, depth less

than 1 meter, salinity on bottom 31%o, 24 August

1964, M. Mazurkiewicz, collector—60 specimens

(USNM 33288). NEW JERSEY: Beesley's Point, Great

Egg Harbor, coarse sand and gravel, about half tide,

H. E. Webster, collector—10 syntypes of Nereis culveri (USNM 541; BMNH). MARYLAND: Broom's

Island, about 10 miles from mouth of Patuxent River,

30 May 1939, J. P. E. Morrison, collector—3 specimens (USNM 22244). Patuxent River at Long Point,

1 mile south of Benedict and N.E. of St. Leonard,

Calvert County, August 1968, S. L. H. Fuller, collector—3 specimens (USNM 40127-8). Upper Chesapeake Bay in Fox Creek, tributary of Rhode River,

22 August 1969, J. Vogel, collector—5 specimens

(USNM 41604). VIRGINIA: Norfolk peninsula in

Willoughby Bay and off Hampton Bar, August 1940,

F. F. Ferguson, collector—12 specimens (USNM

22245-6). NORTH CAROLINA: Beaufort, summer

1877, Mr. C. L. Culver, collector—11 syntypes of

Nereis culveri (USNM 28178). Cape Hatteras area

in Sandy Bay, 12 July 1959, J. L. Taylor, collector—

1 specimen (USNM 43400). Currituck Sound, 19

October 1960, J. A. Kerwin, collector—5 specimens
NUMBER 10 4 15

(USNM 33292). Beaufort, 1967, W. B. Vernberg,

collector—24 specimens (USNM 38742). SOUTH

CAROLINA: Charleston, March 1960, R. Lunz, collector—4 specimens (USNM 33291). GEORGIA: Sapelo Island on Raccoon Bluff Beach, Marsh at high

tide, and Blackbeard Creek, sand and mud flat, 1961,

M. Gray, collector—10 specimens (USNM 33294-6).

High Point, Sapelo Island, muddy sand and Spartina,

1962, D. Frankenberg, collector — 4 specimens

(USNM 33297). FLORIDA: East coast in St. John's

River drainage system, Brevard County, taken on

surface after using Rotenone, 13 May 1956, W.

McLane, collector—numerous specimens (USNM

28083). West coast at Seahorse Key near Cedar

Key, sand, 1959, 1960, J. L. Taylor, collector—65

specimens (USNM 30045, 33289-90). Gulf coast at

Apalachicola and Bald Point, 1960, L. Oglesby, collector—17 specimens (USNM 39640-1). MISSISSIPPI: Biloxi, sandy-mud flat, October 1943, M. W.

Williams, collector—2 specimens (USNM 22242).

East Ocean Springs at Davis Bay, about 2 feet,

brackish water on sheltered shore of sandy mud, 17

August 1949, R. L. Caylor, collector—5 specimens

(USNM 22093). Ocean Springs east of Henderson

Point, 30°18'N, 89°17'W, 21 March 1968, W. Langley, collector—1 specimen (collection Gulf Coast

Laboratory). LOUISIANA: Grand Isle, sandy beach,

E. H. Behre, collector—3 specimens (USNM 22243).

TEXAS : Offats Bayou, Galveston—holotype of heptoner eis nota (AMNH 2896). Upper part Galveston

Bay, center of Trinity Bay, mud, 18 December 1969,

G. E. Williams, collector—1 specimen (USNM

43401). BRAZIL: Santos, February 1914, H. Luderwaldt, collector—3 paratypes of Nereis (Leptonereis)

acuta (USNM 19030). URUGUAY: Arroyo de Pando,

Canelones, in freshwater, E. H. Cordero, collector—6

syntypes of Leptonereis pandoensis (BMNH 1934:4:

22:4-30; USNM 43474). EL SALVADOR, CENTRAL

AMERICA: Gulf of Fonseca, Isla Perrico, G. Hartmann-Schroder, collector—holotype of Laeonereis

brunnea (ZMH 143). Gulf of Fonseca—Isla Perrico,

Jaltepeque and Jiquilisco estuaries, G. HartmannSchroder, collector—numerous paratypes of Laeonereis brunnea (ZMH 143; USNM 43403).

TYPE MATERIAL.—The syntypes of Nereis culveri

from Great Egg Harbor, New Jersey (USNM 541)

include nine complete specimens and one anterior

fragment. One of them has a length of 55 mm, a

width of 4 mm, including setae, and 110 segments.

The syntypes from Beaufort, North Carolina (USNM

28178) include four complete specimens and seven

anterior fragments. The pharynx was not extended

on any of the syntypes.

The three paratypes of Nereis (Leptonereis) acuta

Treadwell (USNM 19030) are incomplete posteriorly,

from 19 to "25 mm in length, 4 mm in width, including setae, and up to 72 segments. None of them

had the pharynx extended, but dissection showed

the presence of the paired triangular papillae on

the oral ring and some tufts of papillae on the maxillary ring; being colorless, they are difficult to see

(not mentioned by Treadwell). The more posterior

parapodia are modified as in L. culveri; some of the

lower neurosetae are homogomph falcigers with

rather long blades (not described by Treadwell).

The syntypes of Leptonereis pandoensis (BMNH

1934:4:22:4-30; USNM 43474) include three smaller complete specimens, one of which has a length

of 42 mm, a width of 4 mm, including setae, with

about 125 segments, and three larger specimens incomplete posteriorly; one with 51 anterior segments

has a length of 32 mm and a width of 5 mm, including setae. None of the syntypes had the pharynx

extended, as noted by Monro; one of them had been

dissected.

The holotype of Leptonereis nota Treadwell

(AMNH 2896) is incomplete posteriorly, with a

length of 55 mm, a width of 2 mm, including parapodia, and composed of 86 segments. It is rather

dark, showing the transverse banding posteriorly.

The setae are mostly broken. It is a female having

large yolky eggs. The pharynx was partially extended,

showing the characteristic tufts of papillae (Figure

5c,d; not mentioned by Treadwell).

The holotype of Laeonereis brunnea from the Gulf

of Fonseca (ZMH 143) consists of anterior and

middle fragments totaling 41 segments (67, according to Hartmann-Schroder), 14 mm in length (about

20 mm—Hartmann-Schroder), and 2 mm in width,

including setae. The numerous paratypes vary in

size from small specimens of 21 segments with posterior growth zone, 2 mm in length and 0.5 mm in

width, to one of 55 segments with posterior growth

zone, 12 mm in length and 1 mm in width. The

types of L. brunnea all appear to be juveniles.

DESCRIPTION.—Length up to 60 mm, width up to

5 mm, including setae, segments up to 140. Body

widest about segment 8, diminishing rapidly an-
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Laeonereis culvert (a,b, syntype of Nereis culveri from New Jersey, USNM 541 ;

c,d, holotype of Leptonereis nota from Texas, AMNH 2896; e, specimen from North Carolina,

USNM 38742; f,g, specimens from Florida, USNM 28083): a, Anterior end, dorsal view,

prostomium partially withdrawn into tentacular segment; b, same, ventral view; c, anterior

end, dorsal view, with pharynx partially extended; left parapodium missing; d, same, ventral

view; e, ventral view anterior end with pharynx fully extended; /, ventral view anterior end

with pharynx partially extended showing basal or oral ring; g, frontal view of extended pharynx.

teriorly and gradually posteriorly. Variable in coloration : iridescent whitish to yellowish or reddish brown;

posterior region sometimes with two broken bands

per segment; conspicuous dark glandular areas in

dorsal ligules and bases of dorsal and ventral cirri.

Prostomium relatively small compared with broad

body, with deep anterior notch, sometimes appearing

as triangular depression on anterior half; frontal antennae and biarticulate palps short; eyes subequal,

moderately large (Figures ba-c; 6a; la). Tentacular

segment about twice as long as following segment;

tentacular cirri relatively short, longest extending to

about setiger 2 (1—3). First two setigers (Figures

6b,c) with dorsal cirri and single notopodial ligule;

neuropodia similar to those of following segments.

Parapodia of anterior region (Figures 6d-f; 7c,d)

larger, with more numerous lobes. Dorsal cirri much

shorter than ligules. Notopodia with larger triangular

upper ligules, much smaller lower ligules, and still

smaller presetal lobes between them. Neuropodia

with rounded presetal and postsetal lobes and oval

lower ligules. Ventral cirri very small. Notosetae and

neurosetae only homogomph spinigers (Figure 6g).

Parapodia of middle and posterior segments smaller,

with fewer lobes (Figure 6i-l; le-h). Presetal notopodial and both presetal and postsetal neuropodial

lobes inconspicuous; neuropodial ligules much

smaller. Upper bundles of neurosetae homogomph

spinigers; lower bundles with homogomph spinigers

and f alcigers with rather long blades (Figure 6h).

Pygidium with paired anal cirri. Jaws of pharynx

with 12 or more teeth extending to near tip; oral or

basal ring bare except for pair of conical papillae

or triangular areas on middorsal side (area VI);

maxillary or distal ring with tufts of cylindrical

hooked papillae on all areas as follows: middorsal

group (area I) with about 5 papillae (3—6); dorsolateral groups (area II) with about 12 papillae (10—

14); ventrolateral groups (area IV) with about 10

papillae (8-12), 3 midventral tufts (area III) of

smaller papillae (Figures 5c-g; 7b). Juveniles with

fewer papillae (Hartmann-Schroder, 1959, for L.

brunnea).

REMARKS. — L. brunnea Hartmann - Schroder

(1959) was based upon what appear to be juveniles

of L. culveri. They have fewer papillae on the maxillary ring of the pharynx, probably associated with

their smaller size. The fifth parapodium on Figure
NUMBER 10 4 17

FIGURE 6.—Laeonereis culveri (specimen from Florida, USNM 30045) : a, Anterior end, dorsal

view; b, parapodium from setiger 1, posterior view; c, para podium from setiger 2, anterior

view; d, parapodium from setiger 3, anterior view, e, parapodium from anterior region, posterior view; /, same, anterior view; g, neuropodial homogomph spiniger from same; h, neuropodial homogomph falciger from same; i, parapodium from middle region, posterior view; ;,

same, anterior view; k, parapodium from posterior region, posterior view; /, same, anterior view.

[Figures g,h, not to scale]

94 by Hartmann-Schroder incorrectly shows four

neuropodial lobes instead of three.

BIOLOGY.—L. culveri is typically an intertidal,

brackish, warm water species along much of the

eastern and southeastern coasts of the United States

where it abounds in estuaries, brackish streams, coves,

and sandy shoals. It occupies extensive sand flats

arid provides an important source of food for shore

birds (Hartman, 1959b: 539). It was found to be

the most important burrowing form on the salt flats
FIGURE 7.—Laeonereis culvert (specimens from Sapelo Island, Georgia, USNM 33294): a, Anterior end, dorsal view; b, frontal view extended pharynx; c, parapodium from anterior region,

posterior view; d, same, anterior view; e, parapodium from middle region, posterior view;

/, same, anterior view; g, parapodium from posterior region, posterior view; h, same, anterior

view.

in the Aransas Refuge, Texas, being resistant to

periods of low salinity and temporary droughts

(Hedgpeth, 1950:111). In Salt Lake, Brevard

County, Florida, in the St. John's River drainage,

specimens were observed to emerge by the thousands

after treatment with Rotenone on 13 May 1956

(observation of Wm. McLane, et al.). According to

Carpenter (1956:101), L. culvert is one of the most
NUMBER 10 4 19

common annelids in the Alligator Harbor area,

Florida, where it was found from mean high tide

downward to below mean low tide. In the Mystic

River estuary, Connecticut, L. culvert appeared to

be distributed in aggregates and occurred in greatest

abundance intertidally in the upper ten centimeters

of soft, fine sediments in habitats where salinities

fluctuated widely, with a maximum range per tidal

cycle from 0.5 to 30 %o (Mazurkiewicz, 1970). Specimens have been maintained experimentally in freshwater (Webster, 1880:113). The species is euryhaline,

known from freshwater (Uruguay, as L. pandoensis—

Monro, 1937:242), from water of low salinity (1 to

6%0 in Florida—Oglesby, 1965:624), and higher

salinities (Florida—Carpenter, 1956:93, 101).

They form tough tubes of mucoid-lined burrows,

fitting the body very closely, often reddish brown

or rustcolored (Webster, 1880:113; Mazurkiewicz,

1970). They are nonselective deposit feeders, with

characteristic sinuous fecal strings being ejected and

accumulating on the surface of the sediment around

their burrow entrances (Mazurkiewicz, 1970).

The mature worms remain atokous, both sexes being

identical in external morphology. They show no

structural changes at the time of reproduction except

that the eyes become a little larger, the body coloration changes from orange-brown to yellow, then to

green, and the muscles of the body wall undergo histolysis, resulting in somewhat shorter and more fragile

bodies. The reproduction and development of L. culvert in the Mystic River estuary, Connecticut, was

followed by Mazurkiewicz (1970). Their life span

includes a single spawning period and normally does

not exceed one year. Spawnings occur during a period

when the water temperature exceeds 20° C from June

through August and possibly into September. While

swimming, presumably along the bottom, the males

shed their sperm through retractile pygidial papillae.

The females spawn within their mucoid tubes, their

eggs being extruded through rents in the body wall.

The mature eggs are moderate in size (135-162 n),

demersal, and are found in clumps adhering to the

walls of the tube. The aeration currents, produced by

dorsoventral undulations of the females, presumably

draw the sperm into the tubes, where fertilization

takes place. The spent females remain in their tubes,

the eggs and embryos being ventilated by the aeration

currents produced by the females. The spent worms

die ten to sixteen days after spawning. Developmental

stages include nonciliated embryos with three pairs

of setal sacs, later developing into benthic larvae with

ciliary bands and occupying burrows in the upper two

centimeters of the sediment, being absent from the

plankton. The ciliary bands are used to ventilate their

burrows. Larvae of three to five setigers feed predominantly on benthic diatoms. All life history stages

appear to tolerate rigorous environmental conditions.

According to Mrs. Winona Vernberg (in correspondence), L. culveri from North Carolina served as

the second intermediate host for the trematode Zoogonus lasius. Mazurkiewicz (1969:1146; 1970) reported a high incidence of infection in L. culveri from

Connecticut by parasitic sporozoans. The life cycle of

the eugregarine sporozoan, Gonospora pilosa Mazurkiewicz, was found to be synchronized with the reproduction of the host, the parasite reproducing only

after the host had spawned.

DISTRIBUTION.—East coast North America from

Connecticut (Mystic River estuary) to Florida (east

coast), Gulf of Mexico (Florida to Mexico), Central

America (El Salvador), east coast South America

from French Guinea (Fauvel, 1923b: 124), Brazil,

and Uruguay (freshwater).

Websterinereis new genus

TYPE-SPECIES.—Nereis tridentata Webster, 1880.

Gender: feminine.

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs of

eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Parapodia of first two setigers subbiramous, notopodium represented by dorsal cirrus and single ligule.

Dorsal cirri at bases of upper notopodial ligules. Notopodia with two ligules and usually additional presetal lobe in anterior parapodia (inconspicuous in W.

glauca). Neuropodia with postsetal lobes and lower

ligules; with or without presetal lobes distinct from

acicular lobes. Ventral cirri short, tapered. Notosetae

homogomph spinigers only. Neurosetae homogomph

and heterogomph spinigers and heterogomph falcigers. Pygidium with paired anal cirri. Pharynx with

paired jaws, without paragnaths; with papillae on

basal or oral ring: large papillae dorsally (area VI)

and single row of minute papillae ventrally (areas

VII-VIII).
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ETYMOLOGY.—The genus is named for Harrison

Edwin Webster, pioneer worker on the Polychaeta.

Four species (together with four synonyms) are

referred herein to Websterinereis, including the following:

W.glauca (Claparede) [including Leonnates pusillus Langerhans; Nereis (Leptonereis) vaillanti SaintJoseph]

W. tridentata (Webster)

W. foli (Fauvel) [including Ceratocephala corallicola Reish]

W. punctata (Wesenberg-Lund) [including Laeonereis ankyloseta Day]

REMARKS.—A species, such as Nereis tridentata,

does not fit into the more or less accepted scheme of

the nereid genera. Webster (1880, 1886) reported

and figured it as having three small paragnaths on

the ventral surface of the basal or oral ring (area

VII) of the pharynx; they were described as being

circular or elliptical, flat, horny, and brown; in addition, two minute fleshy papillae on either side of the

dorsal midline of the same basal ring (area VI) were

observed on living specimens but not when preserved.

Paragnaths were not observed on the pharynx by

Hartman (1945: 21) when she referred the species

to Ceratonereis (in the latter genus, the paragnaths

are confined to the maxillary or distal ring and absent

from the oral ring). Better choices would have been

Eunereis (with paragnaths confined to the oral ring)

or Leptonereis or Nicon (without paragnaths). The

evanescent character of the so-called paragnaths of

N. tridentata is due to the fact that they are not of

the usual type of horny paragnaths found in the Neredidae. They are low, nodular papillae with thickened cuticle (Figure 86). The three midventral ones,

having a slightly thicker cuticular layer, may be

brownish but fade on preservation. Thus they are

easily overlooked, especially when the pharynx is

withdrawn and observed only by dissection.

For Leptonereis foli, Fauvel (1930) described the

slit pharynx as having an inflated rounded vesicle on

area VI and a row of papillae on areas VII-VIII

(exact number of papillae not stated; examination

of the holotype revealed a row of seven papillae).

For Ceratocephala corallicola, Reish (1968) described each area VI with a pair of conical papillae

and areas VII-VIII with eight papillae in a single

row (examination of the type-specimens revealed a

single large papilla on area VI and a row of seven

papillae on areas VII-VIII).

For Leptonereis punctata, Wesenberg-Lund (1949)

observed two small papillae asymmetrically placed on

the ventral side of the oral ring of the extended pharynx on one of the two syntypes (examination of the

syntypes revealed a row of seven minute papillae).

For Laeonereis ankyloseta, Day (1957) observed a

truncate fleshy lobe on area VI and a single row of

six [perhaps seven?], soft, rather indistinct papillae

on areas VII-VIII, the three medial ones on area

VII sometimes brown. The latter would correspond

to the three "paragnaths" of N. tridentata.

Most of the above-mentioned species were included under Laeonereis by Hartman (1945, 1959a).

L. culveri, the type-species Laeonereis, however, has

tufts of papillae on the maxillary ring of the pharynx,

in addition to parapodial differences.

In the upper neuropodial bundles of the middle

and posterior parapodia, stout heterogomph falcigers

appear. When the blades are broken off, they may

appear as stout simple acicular setae. In W. punctata

(and its synonym ankyloseta), the blades are fused to

the stems, forming ankylosed setae. In the posterior

tail regions of male heteronereids of W. glauca, stout

simple setae occur in the uniramous parapodia.

Key to Species of Websterinereis

1. Anterior parapodia with presetal notopodial lobes small, inconspicuous (Figure 14<f-/).

W. glauca

V. Anterior parapodia with presetal notopodial lobes distinct (Figures 8f,g; \0c,d; I2f,g) .... 2

2. Lower neuropodial bundles without heterogomph spinigers (except for sporadic ones in

posterior neuropodia). Parapodial rami of middle and posterior regions not widely separated (Figures 10c,d; lle,f) W. foli

2'. With heterogomph spinigers in lower bundles of neuropodia (Figures 9e; 12<r,i"). Parapodial

rami of middle and posterior regions widely separated (Figures 9a-c; \3a,b,d,e) 3

3. Posterior parapodia with ankylosed simple setae in upper bundles of neuropodia (Figure

1 3 /) •_ ; • W. punctata

3'. Posterior parapodia without ankylosed simple setae in upper bundles of neuropodia.

W. tridentata
NUMBER 10 4 21

Websterinereis tridentata (Webster), new combination

FIGURES 8, 9

Nereis tridentata Webster, 1880:113; 1886:142, pi. 7: figs.

33-40.

Ceratonereis tridentata.—Hartman 1945:21, pi. 3: figs. 3,

4; 1951:48.—Wells and Wells 1962:149.—Wells 1965:

123.

MATERIAL EXAMINED.—GEORGIA, off Sapelo Island,

M. Gray, collector: off Sea Buoy, 20 meters, 7

March 1961—4 specimens (USNM 33264); off

Sapelo Whistle, 18 meters, 8 November 1961—2

specimens (USNM 33266) ; off Sapelo Whistle, 20

meters, 7 February 1962—1 specimen (USNM

33265). NORTH CAROLINA: off Beaufort, 34°28'N,

76°O7'W, 40 meters, sandy mud, 30 September 1965,

J. H. Day, collector—6 specimens (USNM 43404).

Off Beaufort, on coral in 10 meters, L. R. McCloskey,

collector—1 specimen (USNM 43402).

DESCRIPTION.—Length up to 65 mm, width up to

3.5 mm, including parapodia, segments up to 105.

Prostomium with faint to deep middorsal depression

on anterior half, short tapered frontal antennae, and

stout biarticulate palps; eyes large, anterior pair being larger than posterior pair (Figure 8a). Tentacular segment about twice length of following segment;

tentacular cirri short, longest extending to about setiger 2 (1—4). Parapodia of first two setigers with

dorsal cirrus, notopodial ligule, neuropodial postsetal

lobe, lower ligule, and ventral cirrus (Figure 8c,d);

upper bundle of neurosetae composed of homogomph

spinigers with long blades, heterogomph spinigers with

short blades, and heterogomph falcigers with relatively

long blades (Figure 9d-f); lower bundle neurosetae

all heterogomph spinigers with short blades. Third

parapodium with two notopodial ligules, short presetal lobe, and fan-shaped bundle of homogomph

spinigers with long blades; neuropodium similar to

those of following setigers (Figure Be).

fAn

FIGURE 8.—Websterinereis tridentata (USNM 43403): a, Dorsal view anterior end, with

pharynx partially extended; b, same, ventral view; c, parapodium from setiger 1, posterior view;

d, parapodium from setiger 2, anterior view; e, parapodium from setiger 3, anterior view;

/, parapodium from anterior region, posterior view; g, same, anterior view.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Parapodia of anterior region (Figure Sf,g) with

dorsal cirri emerging from basal parts of upper notopodial ligules and extending slightly beyond them;

notopodium with upper ligules large, subtriangular

and lower ones smaller, subconical; presetal notopodial lobes, between them, somewhat smaller than

lower ligules; presetal and postsetal neuropodial

lobes subequal in size to notopodial presetal lobes;

lower neuropodial ligules subequal to upper notopodial ligules; short subulate ventral cirri. Thus, anterior parapodia provided with six lobes in addition to dorsal and ventral cirri: three notopodial

(two ligules and presetal lobe) and three neuropodial (presetal and postsetal lobes and lower ligule).

Notosetae all homogomph spinigers with long blades

(Figure 9d). Upper bundle of neurosetae composed

of homogomph spinigers with long blades and heterogomph falcigers with relatively long blades (Figure

9/) ; lower bundle of neurosetae composed of heterogomph spinigers with short blades (Figure 9e) and

heterogomph falcigers with relatively short blades

(Figure 9g).

Parapodia of middle region (Figure 9a,b) with

notopodia and neuropodia widely separated; notopodial presetal lobes diminished in size but still distinct; neuropodial presetal lobes indistinct from

acicular lobes. Upper bundles of neurosetae homogomph spinigers with long blades; lower bundles of

neurosetae with heterogomph spinigers with short

blades and heterogomph falcigers with short blades

(Figure 9h).

Parapodia of posterior region (Figure 9c) modified somewhat; both notopodial and neuropodial presetal lobes indistinct, resulting in four lobes: two

notopodial ligules, postsetal neuropodial lobe and

lower neuropodial ligule. Upper bundle of neurosetae

FIGURE 9.—Websterinereis tridentata (USNM 43403): a, Parapodium from middle region,

posterior view; b, same, anterior view; c, parapodium from posterior region, posterior view;

d, homogomph spiniger with long blade; e, heterogomph spiniger with short blade; /, heterogomph falciger with relatively long blade; g, heterogomph falciger from lower neuropodial

bundle of anterior region; h, same, from middle region; i, same, from posterior region,

;, stouter heterogomph falciger from upper neuropodial bundle of posterior region; k, same,

with blade missing. [Figures d—k, not to scale]
NUMBE R 10 4 23

homogomph spinigers with long blades and 1—2 stout

heterogomph falcigers with short blades (Figure 9;') ;

blades sometimes missing, then appearing as stout

simple setae (Figure 9A;) ; lower bundle of neurosetae

heterogomph spinigers with relatively short blades and

heterogomph falcigers with short blades (Figure 9i).

Pygidium with pair long anal cirri. Jaws of

pharynx (Figure 8a,b) with numerous sharp teeth.

Maxillary or distal ring bare, without paragnaths

or papillae; basal or oral ring with subconical papillae on dorsal surface (area VI) and row of seven

papillae on ventral surface (areas VII-VIII); papillae circular or elliptical, flat, horny, middle three

(area VII) with thicker cuticle, sometimes brown

in color but fading on preservation.

BIOLOGY.—W. tridentata forms compact tubes of

sand grains and is associated with hard substrates,

shelly bottoms and corals. In North Carolina and

Florida, it was found by Wells and Wells (1962) and

Wells (1965) to be a pest of the calico scallop

Aequipecten gibbus where it penetrates areas between the shell and mantle, forming U-shaped tubes

of fine sand grains loosely held together by mucus,

and causing large oval or crescent-shaped mud blisters on the shells. The anterior ends of the worm may

project into the mantle cavity. They appear to be

facultative parasites, parasitizing the scallop but also

occurring among the epifauna of the shells. Reproduction, as far as is known, occurs in the atokous

condition.

DISTRIBUTION.—East coast North America from

New Jersey to Florida, Gulf of Mexico (Mississippi,

Texas).

Websterinereis foli (Fauvel) new combination

FIGURES 10, 11

Leptonereis foli Fauvel, 1930:520, fig. 3.

Laeonereis foli.—Hartman 1949:55; 1959a: 245.

Ceratocephala [sic] corallicola Reish, 1968:215, fig. 4:1-5.

MATERIAL EXAMINED.—He des Pins, New Caledonia, 1928, Mm. A. Pruvot-Fol, collector—holotype

of Leptonereis foli (MNHNP).

Bogen Island, Eniwetok Atoll, Marshall Islands, 12

September 1956, D. Reish, collector—holotype and

7 paratypes of Ceratocephala corallicola (USNM

38392-3).

TYPE MATERIAL.—The holotype of L. foli consists

of an anterior fragment of 62 segments, 17 mm long

and 1 mm wide, including setae. The pharynx was

not extended but had been slit.

The holotype of C. corallicola consists of about 90

segments, 12 mm long and 1 mm wide, including

setae. The pharynx was not extended. Of the seven

paratypes, four are complete and three consist of

anterior fragments. The pharynx is partially or completely extended on four of them.

DESCRIPTION.—Length up to 20 mm, width up to

1 mm, including parapodia, segments up to 112.

Prostomium with faint to deep mid-dorsal depression

on anterior half, short tapered frontal antennae, and

stout biarticulate palps; eyes moderately large, anterior pair being larger than posterior pair (Figures

10a, lla). Tentacular segment longer than following segment; tentacular cirri with styles short to

long, longest extending to about setiger 4 (2—7).

Parapodia of first two setigers with dorsal cirrus,

notopodial ligule, neuropodial postsetal lobe, lower

ligule and ventral cirrus (Figure 106) ; upper bundle

of neurosetae composed of homogomph spinigers and

heterogomph falcigers with relatively long blades

(Figure \0e) ; lower bundle of neurosetae heterogomph falcigers (no heterogomph spinigers).

Parapodia of anterior region (Figure \\c,d) with

dorsal cirri emerging from basal parts of upper notopodial ligules and extending beyond them; upper

notopodial ligules rather large, subtriangular; lower

notopodial ligules smaller, subconical; presetal notopodial lobes, between them, smaller than lower

ligules; presetal and postsetal neuropodial lobes subequal in size to notopodial presetal lobe; lower neuropodial ligules slightly smaller than upper notopodial

ligules; short subulate ventral cirri. Notosetae all

homogomph spinigers (Figure \\g). Upper bundle of

neurosetae composed of homogomph spinigers and

heterogomph falcigers with short blades; lower bundle of neurosetae composed of heterogomph falcigers

(Figure \\h).

Parapodia of middle region (Figures \0c,d; lie,/)

with notopodial presetal lobes diminishing in size;

neuropodial presetal lobes indistinct from acicular

lobes; upper bundle of neurosetae homogomph spinigers (Figures 10/; lh") and single stouter heterogomph falciger with short blade (Figures \0g; 11;) ;

lower bundle of neurosetae with heterogomph falcigers bearing short blades (Figures \0h; Ilk). Posterior parapodia sometimes with single heterogomph

spiniger having relatively short blade in lower bundle

of neurosetae.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Websterinereis foli (holotype of Leptonereis foli, MNHNP): a, Anterior end,

dorsal view; b, first parapodium, anterior view; c, parapodium from middle region, posterior

view; d, same, anterior view; e, heterogomph falcigerous neuroseta from first parapodium;

/, homogomph spiniger; g, heterogomph falcigerous neurosetae from upper bundle of middle

region; h, same, from lower bundle. [Figures e-h, not to scale]

9

FICURE 11.—Websterinereis foli (paratypes of C. corallicola, USNM 38393): a, Anterior end,

dorsal view, pharynx partially extended; b, same, ventral view; c, parapodium from anterior

region, posterior view; d, same, anterior view; e, parapodium from middle region, posterior

view; /, same, anterior view; g, homogomph spiniger from anterior region; h, heterogomph

falcigers from anterior region; «, homogomph spiniger from middle region; ;, heterogomph

falciger from upper bundle neuropodium; k, same, from lower bundle neuropodium. [Figures g-k,

not to scale]
NUMBE R 10 4 25

Pygidium with long anal cirri. Jaws of pharynx

(Figure \\b) bearing numerous sharp teeth; maxillary or distal ring bare, without paragnaths or papillae; oral or basal ring with large subconical papilla

on dorsal surface (area VI) and row of seven papillae on ventral surface (areas VII-VIII).

REMAR K S.—Pettibone (1970:233) previously

pointed out that C. corallicola was incorrectly assigned to Ceratocephale. W. foli is associated with

corals, and is probably much more widely distributed

than the two locality records might indicate. Due to

its small size, it easily escapes notice.

DISTRIBUTION.—Central Pacific (New Caledonia,

Marshall Islands). Associated with corals.

Websterinereis punctata (Wesenberg-Lund) new

combination

FIGURES 12, 13

Leptonereis punctata Wesenberg-Lund, 1949:289, figs. 21-

23.

Laeonereis ankyloseta Day, 1957:83, fig. 5a-j; 1960:321.—

Hartman 1959a: 243.

Nicon punctata.—Hartman 1958:265; 1959a:245, 274.

MATERIAL EXAMINED.—Persian Gulf, G. Thorson,

collector: station 25A, 14 March 1937, 28°58'N,

49°30'E, 49 meters, sand with very little clay—

larger syntype (UZMC) ; station 45D, 31 March

1937, 27°42'N, 51°25'E, 58 meters, clay mixed with

sand, shells—smaller syntype (UZMC).

TYPE MATERIAL.—The larger syntype from station

25A is incomplete posteriorly, 40 mm in length, 3

mm in width, including setae, and 70 segments; it

is rather flaccid. The smaller syntype from station

45D is a complete specimen of about 90 segments,

30 mm in length and 2 mm in width, including

setae; it is a female containing relatively large eggs;

the eyes are larger than those in the other syntype.

The pharynx is extended in both syntypes.

DESCRIPTION.—Length up to 50 mm, width up to

FIGURE 12.—Websterinereis punctata (smaller syntype of Leptonereis punctata from station

45D, UZMC): a, Anterior end, lateral view; b, first parapodium, posterior view; c, second

parapodium, anterior view; d, homogomph spiniger from same; e, heterogomph spiniger from

same; /, parapodium from anterior region, posterior view; g, same, anterior view: h, neuropodial heterogomph falciger from same; i, neuropodial heterogomph spiniger from same.

[Figures d,e,h,i, not to scale]
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

9

FIGURE 13.—Websterinereis punctata (smaller syntype of Leptonereis punctata from station

45D, UZMC): a, Parapodium from middle region, posterior view; neurosetae mostly missing;

b, same, anterior view; c, stout heterogomph falciger from upper bundle neurosetae from same;

d, parapodium from posterior region, posterior view; e, same, anterior view; /, stout simple seta

from upper bundle neurosetae from same; g, heterogomph falciger from lower bundle neurosetae from same; h, heterogomph spiniger from lower part of lower bundle neurosetae from

same. [Figures c,f-h, not to scale]

3 mm, including setae, segments up to 90. Prostomium with short tapered frontal antennae and

stout biarticulate palps; moderately large eyes, anterior pair larger than posterior pair (Figure 12a).

Tentacular segment about twice length of following

segment; tentacular cirri with cylindrical cirrophores,

longest extending to about setiger 6. Parapodia of

first two setigers with dorsal cirrus and notopodial

ligule; neuropodium with postsetal lobe, short presetal

lobe, lower ligule, and ventral cirrus (Figure \2b,c) ;

upper bundle of neurosetae composed of homogomph

spinigers (Figure \2d) and heterogomph spinigers

(Figure 12e); lower bundle of neurosetae heterogomph spinigers with blades long (upper ones) to

short (lower ones).

Parapodia of anterior region (Figure 12f,g) with

dorsal cirri emerging from basal parts of upper notopodial ligules and extending slightly beyond them;

upper and lower notopodial ligules subconical and

subequal; presetal notopodial lobe, between ligules,

rounded and somewhat smaller; postsetal neuropodial

lobe elongate-conical; presetal neuropodial lobe

short; lower neuropodial ligules subequal to notopodial ligules; short subulate ventral cirri. Notosetae

all homogomph spinigers with long blades (Figure

I2d). Upper bundle of neurosetae composed of few
NUMBER 10 4 27

homogomph spinigers with long blades and heterogomph falcigers with relatively long blades (Figure

\2h) ; lower bundle of neurosetae composed of upper

heterogomph spinigers with long blades, heterogomph

falcigers with relatively long blades and lower heterogomph spinigers with short blades (Figure 12i).

Parapodia of middle region (Figure 13a,b) with

notopodia and neuropodia more widely separated;

notopodial presetal lobes diminishing in size; neuropodial presetal lobes indistinct from acicular lobes.

Upper bundle of neurosetae homogomph spinigers

with long blades and few (1—4) stout heterogomph

falcigers with bifid tips, blades showing signs of fusion to stems (Figure 13c); lower bundle neurosetae

with heterogomph spinigers and heterogomph falcigers, both bearing short blades.

Parapodia of posterior region (Figure \?>d,e) with

notopodial presetal lobes further diminished in size.

Upper bundle of neurosetae homogomph spinigers

with long blades and few (1—2) stout simple falcigers formed by complete fusion of blades to stems,

their tips bifid, differing in appearance due to wear

(Figure 13/). Lower bundle of neurosetae heterogomph spinigers with long blades, heterogomph falcigers with short blades (Figure 13g), and heterogomph spinigers with relatively short blades (Figure

13/t).

Jaws of pharynx (Figure 12a) with about ten

teeth. Maxillary or distal ring bare, without paragnaths or papillae; basal or oral ring with large subconical papillae on dorsal surface (area VI) and

row of seven small papillae on ventrolateral surface

(areas VII-VIII), middle three (area VII) somewhat larger.

REMARKS.—-Wesenberg-Lund observed only two

small asymmetrically placed papillae on the ventral

surface of the oral ring of the extended pharynx on

the smaller syntype; seven papillae were observed by

me on both syntypes. The stout simple neurosetae

were found in both syntypes of L. punctata, not on

just the larger one, as indicated by Wesenberg-Lund.

Day (1957:84) indicated that L. ankyloseta was

closely allied to L. punctata. The two species were

distinguished by the greater number of papillae on

the pharynx of the former and by the form of the

enlarged tips of the stout simple neurosetae. Examination of the syntypes of L. punctata obliterates these

distinctions and the two are herein considered to be

synonymous.

BIOLOGY.—In the Persian Gulf, W. punctata was

collected in 49 to 59 meters on bottoms of sand, clay

and shells. In South Africa, it was dredged in 2 to

55 meters on bottoms of sand, mud, rock, shells and

coral and was found to be common in sheltered sand

banks. Day (1960:321) collected a heteronereid in

Mossel Bay in 9 meters (not described).

DISTRIBUTION.—Persian Gulf (northern and central part) and South Africa (False Bay to Natal and

Mozambique). In 2 to 58 meters.

Websterinereis glauca (Claparede) new combination

FIGURES 14—16

Nereis (Leptonereis) glauca Claparede, 1870:454, pi. 7:

fig. 3.

Leonnates pusillus Langerhans, 1880:279, pi. 14: fig. 10.

Nereis (Leptonereis) vaillanti Saint-Joseph, 1888:246, pi.

10: figs. 113-123, pi. 11: fig. 124.

Nereis (Leonnates) pusillus.—Augener 1910:236.

Leptonereis glauca.—Ramsay 1914a: 244, pi. 1: figs. 1-10.—

Fauvel 1914:163, pi. 12: figs. 5-23; 1923a:333, fig. 129,

a-d.—Fage and Legendre 1927:90, figs. 9, 10.

Laeonereis glauca.—Hartman 1945:22; 1949:55; 1959a:

243, 262.

MATERIAL EXAMINED.—Plymouth, England, 6 December 1913, C. Crossland, collector—2 specimens

(BMNH 1938:7:1:1-6).

Dinard, France, collection of M. le Baron de SaintJoseph, no. 20, 1911 (as Leptonereis vaillanti) —

atokous female with large eggs (MNHNP). Cherbourg, France, 25 May 1922, collection of P. Fauvel—2 male heteronereids (BMNH 1928:4:26:

827-9).

Cape Pescade, near Algers, 5 December 1924—13

females (MNHNP; USNM 43399). Cape Matifou,

Algeria, 22 April 1924, at night-light—numerous

male heteronereids (MNHNP; USNM 43398).

DESCRIPTION.—Length up to 35 mm, width up to

3 mm, including setae, segments up to 65. Prostomium with short subulate frontal antennae and stout

biarticulate palps; eyes moderately large (Figure

14a). Tentacular segment about twice length of following segment; tentacular cirri short, longest extending to about setiger 4 (3—7). Parapodia of first

two setigers with dorsal cirrus, notopodial ligule,

neuropodial postsetal lobe, ligule and ventral cirrus

(Figure \4b,c) ; upper bundle neurosetae composed

of homogomph spinigers and heterogomph falcigers;

lower bundle with heterogomph spinigers and falcigers.
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Websterinereis glauca (atokous female with large eggs, from Dinard, MNHNP):

a, Anterior end, dorsal view; b, first parapodium, anterior view; c, second parapodium, posterior view; d, third parapodium, anterior view; e, anterior parapodium, posterior view; /, same,

anterior view; g, posterior parapodium, posterior view; h, same, anterior view; i, homogomph

spiniger from anterior region; ;', heterogomph falcigers from anterior region; k, same, from

upper bundle posterior region; /, heterogomph spiniger with short blade from lower bundle

neuropodium of posterior region. [Figures :'—/, not to scale]

Parapodia of anterior region (Figure I4d-f) with

dorsal cirri emerging from basal parts of upper notopodial ligules and extending slightly beyond them;

upper and lower notopodial ligules subequal, subconical, with low presetal notopodial lobe between

them; neuropodia with anterior subconical acicular

lobe and slightly longer postsetal lobe; lower neuropodial ligule smaller than notopodial ligules; ventral

cirri short subulate. Notosetae all homogomph spinigers with long blades; upper bundle of neurosetae

homogomph spinigers with long blades (Figure 14t)

and heterogomph falcigers (Figure 14/) ; lower bundle of neurosetae with or without heterogomph spinigers with short blades; with heterogomph falcigers.

Parapodia of posterior region (Figure I4g,h) somewhat modified; upper notopodial ligules larger than

lower ones; lower neuropodial ligules smaller than

neuropodial postsetal lobes. Upper bundle of neurosetae homogomph spinigers and few stouter heterogomph falcigers (Figure 14k) ; lower bundle of

neurosetae heterogomph spinigers with relatively

short blades (Figure 14/) and heterogomph falcigers.

Pygidium with anal cirri (Figure 15c). Jaws of

pharynx (Figure I6a,b) with numerous teeth (10—

15). Maxillary or distal ring bare, without paragnaths

or papillae; basal or oral ring with pair of large
NUMBE R 10 4 29

FIGURE 15.—Websterinereis glauca (female heteronereid from Cape Pescade, near Algers,

MNHNP) : a, Parapodium from anterior region, anterior view; b, parapodium from middle

region, posterior view; with mixture of long swimming and usual type of setae; c, posterior

end, dorsal view.

anC

papillae on dorsal surface (area VI) and row of

about seven (5—9) minute papillae on ventrolateral

surface (areas VII-VIII).

DESCRIPTION OF FEMALE HETERONEREID.—Body divided into three regions: anterior region of 15—18

setigers with parapodia unmodified, middle region of

about 16 modified setigers, and posterior unmodified

region of about 12 setigers. Parapodia of anterior region (Figure 15a), posterior region, and pygidium

(Figure 15c) similar to those of atokous individuals.

Parapodia of middle region (Figure 15fc) enlarged,

with long swimming setae in addition to usual

shorter notosetae and neurosetae (not replacing

them).

DESCRIPTION OF MALE HETERONEREID.—Body divided into three regions: anterior region of 14—15

setigers, middle region of about 30 setigers (16—45),

and posterior region of 10—12 setigers forming sort

of tail. Prostomium with eyes enlarged (Figure 16a).

Dorsal cirri of anterior seven setigers enlarged, clavate, increasing in size posteriorly (Figure I6c,d);

ventral cirri of anterior seven setigers thicker than

those of following setigers.

Parapodia of middle region enlarged and considerably modified (Figure I6f,g). Dorsal cirri cirriform (not lobulated) ; dorsal lamellae on medial

bases of dorsal cirri; large foliaceous postsetal lamellae

developed from postsetal neuropodial lobes, extending

beyond ligules; winglike ventral lamellae on bases of

long cirriform ventral cirri. Usual type of notosetae

and neurosetae completely replaced by numerous

long swimming setae.

Parapodia of posterior tail region uniramous (Figure 16i) ; notopodia absent except for large subulate

dorsal cirri; neuropodia with lower ligules and ventral

cirri. Upper bundle of neurosetae composed of few

homogomph spinigers and one to two stout dark

simple acicular setae with hooked tips (Figure 16;) ;

lower bundle of neurosetae consisting of one to two

simple acicular setae. Pygidium with paired stout

dorsal lobes and anal cirri (without papillated area

around anus; Figure \6h).

REMARKS.—Claparede (1870) described and figured a female of the atokous form. Langerhans

(1880) described and figured the male heteronereid

(as Leonnates pusillus). The three forms of the

species—atokous nereid, female and male heteronereids—were recognized by Saint-Joseph [1882, as N.

(Leptonereis) vaillanti], L. pusillus was questionably

referred to Leptonereis glauca by Ramsay (1914a)

and by Fauvel (1914) without question. Nereis

(Leptonereis) vaillanti was referred to N. (Leonnates) pusillus by Augener (1910) and to L. glauca

by Ramsay (1914a) and by Fauvel (1914).

BIOLOGY.—W. glauca has been found on pilings,

among compound ascidians, sponges, and other sedentary forms, and on bottoms of mud and stones. Male

heteronereids were found in large numbers in the
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

neL

FIGURE 16.—Websterinereis glauca (male heteronereid from Cape Matifou, Algeria, MNHNP) :

a, Dorsal view of anterior end, with pharynx extended; b, same, ventral view; c, first parapodium, anterior view; d, parapodium from setiger 7, anterior view; e, parapodium from setiger 8,

anterior view; /, modified parapodium from middle region, posterior view; g, same, anterior

view; h, posterior end, lateral view; i, parapodium from posterior region; ;, neuropodial hook

from same. [Figures i,j, not to scale]

plankton but females were taken only rarely at the

surface (Fage and Legendre, 1927). The females are

only partly transformed to heteronereids—there may

be no trace of swimming setae, or swimming setae

may appear in addition to the usual setae but not

replacing them. Large eggs are developed (up to

240 fi in diameter, according to Ramsay, 1914a) and

laid in the tube. Development is presumably direct.

DISTRIBUTION.—Northeast Atlantic from Great

Britain to Madeira, Mediterranean.

Kinberginereis new genus

T Y p E-S p E c i E s.—Nereis (Leptonereis) inermis

Hoagland, 1920. Gender: feminine.

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs of

eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Parapodia of first two setigers subbiramous; notopodium with dorsal cirrus, notopodial ligule and slender

notoaciculum. Dorsal cirri at bases of upper notopodial ligules. Notopodia with two ligules. Neuropodia with bilobed presetal lobe, short rounded postsetal lobe, and subsetal ligule. Ventral cirri short,

tapered. Notosetae homogomph spinigers only. Neurosetae all compound spinigers — homogomph and

slightly heterogomph or hemigomph. Pharynx with

paired jaws and soft papillae on oral ring only.

ETYMOLOGY.—The genus is named for Johan Gustaf Hyalmar Kinberg, pioneer worker on the Polychaeta.

REMARKS.—Kinberginereis resembles Tylonereis

Fauvel, with type-species T. bogoyawlenskyi Fauvel

(1911:376) from the Persian Gulf and Indian Ocean.

In Tylonereis, the neurosetae are homogomph spinigers only; the dorsal cirri are very short, much shorter

than the notopodial ligules; notoacicula are absent on

the first two setigers; the pharynx bears tufts of papillae on both maxillary and oral rings.

Kinberginereis inermis (Hoagland) new combination

FIGURE 17

Nereis (Leptonereis) inermis Hoagland, 1920:608, pi. 47:

figs. 8-12.
NUMBER 10 4 31

TYPE-MATERIAL.—Cliff Island, Philippine Islands,

13 meters, mud bottom, Albatross station D5346, 26

December 1908—holotype (USNM 18947). The

holotype consists of an anterior fragment of 29 setigerous segments, 12 mm in length, and 5 mm in

width, including parapodia. The pharynx is not extended.

DESCRIPTION.—Body flattened dorsoventrally, segments much compressed. Prostomium partially withdrawn into achaetous tentacular segment, with deep

anterior notch between bases of conical frontal antenna; stout palps more or less telescoped, appearing two to three segmented; eyes moderately large,

on posterior half of prostomium (Figure 17a). Tentacular segment about equal in length to following segment but larger and inflated laterally and ventrally;

tentacular cirri with long cylindrical cirrophores;

styles of longest cirri extending to about setiger 13

(posterodorsal pair missing—perhaps even longer).

Parapodia of first two setigers with dorsal cirrus,

single notopodial ligule, and slender notoaciculum

with curved distal tip; neuropodium similar to those

of following segments (Figure \7b,c).

Anterior biramous parapodia (Figure \7d,e) with

cirriform dorsal cirrus and two conical notopodial

ligules subequal in length; small bundle of notosetae

emerging between ligules; notosetae all homogomph

spinigers with long blades, finely serrated along one

side (Figure \7h). Neuropodium with bilobed presetal lobe—subequal conical lobes and shorter

rounded postsetal lobe; subsetal neuropodial ligule

similar in size and shape to notopodial ligules. Neurosetae very numerous; upper bundle all homogomph

spinigers with long blades similar to notosetae (Figure

17 h) ; lower bundle slightly heterogomph or hemigomph spinigers, upper few with blades shorter,

curved, with long spines along basal part (Figure

17i), middle ones with blades longer, straight, and

finely serrated along one side (Figure 17;), and lower

ones with shorter blades (Figure 17/;). Ventral cirri

tapered, shorter than neuropodial ligule.

More posterior parapodia (Figure 17f,g) somewhat modified; cirriform dorsal cirri more elongated,

extending far beyond notopodial ligules; upper notopodial ligules decreasing in size, becoming considerably smaller than lower notopodial ligules.

Pharynx (observed only when cut) without paragnaths or papillae except for fleshy knobs on areas VI

of oral ring. Hooked jaws amber-colored, with serrations or teeth along most of concave border—about

20 teeth.

DISTRIBUTION.—Philippine Islands. In 13 meters.

Rullierinereis new genus

TYPE-SPECIES.—Leptonereis zebra Rullier, 1963.

Gender: feminine.

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs

of eyes. Tentacular segment apodous and achaetous;

four pairs of tentacular cirri with distinct cirrophores. Parapodia of first two setigers subbiramous;

notopodium represented by dorsal cirrus and single

ligule. Dorsal cirri at bases of upper notopodial

ligules. Notopodia with two ligules, upper one sometimes fused with dorsal cirri (in posterior notopodia of

zebra; from setiger 3 on, in bahamensis). Neuropodia

with acicular lobes and lower ligules. Ventral cirri

short, tapered. Notosetae homogomph spinigers and,

more posteriorly, homogomph falcigers. Neurosetae

homogomph and heterogomph spinigers and heterogomph falcigers. Pygidium with paired anal cirri.

Pharynx with paired jaws, without paragnaths or

papillae (latter present on oral ring in bahamensis).

ETYMOLOGY.—The genus is named for Francois

Rullier, eminent worker on the Polychaeta.

REMARKS.—The following nereid species are referred to Rullierinereis:

R. mexicana (Treadwell, 1942), as Leptonereis.

Lower California, western Mexico.

R. bahamensis (Hartmann-Schroder, 1958), as Tylorrhynchus. Bimini, Bahamas.

R. zebra (Rullier, 1963), as Leptonereis. Mauritius,

Indian Ocean.

R. uncinata (Hartman, 1965b), as Nicon uncinatus.

Off New England.

R. gallardoi new species, as Nicon maculatus.—Gallardo, 1968. South Viet Nam.

The five species lack paragnaths on the pharynx

and are provided with one to two homogomph falcigers in the more posterior notopodia (not present

in male heteronereids lacking posterior unmodified

regions). R. mexicana is known only as sexual epitokes. The other four species are small and heteronereids have not been observed.

Hartman (1965a: 36) indicated that L. zebra

should perhaps be referred to Nicon because the

proboscis is smooth. However, the species may be

separated from members of the genus Nicon on other

characters. R. bahamensis was incorrectly assigned
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 17.—Kinberginereis inermis [holotype of N. (L.) inermis, USNM 18947]: a, Anterior

end, dorsal view, prostomium partially withdrawn in tentacular segment, posterior pair eyes

visible through transparent integument dotted in; styles of two upper right and second upper

left tentacular cirri missing; b, first parapodium, anterior view; c, second parapodium, posterior

view; d, parapodium from anterior region, anterior view; e, same, posterior view; /, parapodium

from middle region (about setiger 26), anterior view; g, same, posterior view; h, homogomph

spinigerous notoseta; i, slightly heterogomph or hemigomph spiniger from upper part of lower

neurosetal bundle; ) , same, from middle of bundle; k, same, from lower part of bundle.

[Figures h—k, not to scale]

fAn
NUMBER 10 4 33

to- the genus Tylorrhynchus Grube by HartmannSchroder. In the latter genus, the neuropodia are

provided with well-developed anterior and posterior

lips but without lower neuropodial ligules.

R. bahamensis was reported as having two achaetous tentacular segments (perhaps distorted in connection with the extended pharynx?). Also, beginning

with setiger 3, the upper notopodial ligules are fused

with the dorsal cirri (this is true only in the posterior

notopodia of R. zebra). The maxillary ring of the

pharynx is provided with some soft papillae in R.

bahamensis; papillae were not observed in the other

four species. This character, however, is difficult to

observe on small specimens, especially when the

pharynx is not extended.

R. uncinata was reported as having only heterogomph spinigers and falcigers in the neuropodia (no

homogomph spinigers).

Key to the Species of Rullicrincreis

1. Upper notopodial ligules fused with dorsal cirri from setiger 3 on (pi. 39: figs. 16, 17, in

Hartmann-Schroder, 1958). Pharynx with papillae on oral ring. [Notopodial homogomph

spinigers with blades short, with bifid tips, pi. 39: fig. 18, in Hartmann-Schroder].

*R. bahamensis

1'. Upper notopodial ligules distinct from dorsal cirri (except in posterior parapodia of R.

zebra). Pharynx without papillae 2

2. Upper notopodial ligules diminishing in size posteriorly and fused with dorsal cirri (Figure

18/). Neuropodial heterogomph falcigers with short blades (Figure 18A, i). [Notopodial

homogomph falcigers with blades short, oval, Figure 18;"] R. zebra

2'. Upper notopodial ligules distinct from dorsal cirri in posterior region. Neuropodial heterogomph falcigers with moderately long blades (Figures 19k, 22g,k,n) 3

3. Neuropodial heterogomph falcigers with bifid tips (pi. 12: fig. c, in Hartman, 1965). Dorsal

cirri shorter than notopodial ligules (at least on setiger 18, pi. 12: fig. a, in Hartman).

[Notopodial homogomph falcigers elongate-oval, spinous, pi. 12: fig. b, in Hartman].

*R. uncinata

3'. Neuropodial heterogomph falcigers with entire tips (Figures 19k, 22g,k,n). Dorsal cirri

extending beyond notopodial ligules (Figures I9f,g, 22d,i,l) 4

4. Notopodial homogomph falcigers with blades moderately long, spinous, with blunt curved

tips (Figure 20e) R. mexicana

4'. Notopodial homogomph falcigers with blades short, elongate-oval, smooth (Figure 22;).

R. gallardoi

'Specimens not examined in this study.

Rullierinereis zebra (Rullier) new combination

FIGURE 18

Leptonereis zebra Rullier, 1963:474, fig. 1: 1-12.—Hartman 1965a:36.

MATERIAL EXAMINED.—Mauritius, Indian Ocean,

1962, C. Michel, collector—2 syntypes (no. 16-10

and slide W-92, collection of F. Rullier).

TYPE-MATERIAL.—One of the eight syntypes (no.

16-10) is a female, 9 mm in length, 1 mm in width,

including setae, and 43 segments, containing very

large eggs. A smaller syntype (on slide W-92) is

6 mm long, 1 mm wide, with 35 segments. The

pharynx is withdrawn on both syntypes.

DESCRIPTION.—Length up to 10 mm, width up to

1 mm, including setae, segments up to 48. Anterior

segments banded with brown granular pigmentation.

Anterior region (about 13 setigers) flattened dorsoventrally, segments short, with parapodia more conspicuous than those posterior. Posterior region cylindrical, inflated, with segments well delineated; body

wall thin, transparent, dark parapodial acicula, ventral nerve cord and glands (three per parapodium)

appearing prominently; sometimes filled with very

large eggs—their diameter about half that of body

(Figure 18/). Prostomium (Figure 18a) with short

tapered frontal antennae, stout biarticulate palps, and

small subequal eyes. Achaetous tentacular segment

up to twice length of following segment, tentacular

cirri rather short, longest extending to setiger 1-3.

Parapodia of first two setigers (Figure 186) with

large, thick dorsal cirrus and single notopodial ligule;

neuropodium similar to those of following setigers.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 18.—Rullierinereis zebra (syntype no. 16-10 of L. zebra, collection of F. Rullier) :

a, Anterior end, dorsal view, prostomium partially withdrawn into tentacular segment, posterior

pair eyes dotted in; b, first parapodium, posterior view; c, third parapodium, posterior view;

d, parapodium from anterior region, posterior view; e, same, anterior view; /, parapodium

from posterior region, anterior view, with single large egg; g, neuropodial homogomph spiniger

from setiger 3; h, neuropodial heterogomph falciger from setiger 3; i, same, from anterior

region; j , notopodial homogomph falcigers from posterior region; k, heterogomph spiniger from

lower bundle neuropodium, from posterior region. [Figures g—k, not to scale]

Parapodia biramous from setiger 3 posteriorly

(Figure 18t). Dorsal cirri subulate, extending beyond

notopodial ligules. Upper and lower notopodial ligules

subconical and subequal in size; notosetae homogomph spinigers, few in number (1-2). Neuropodial

acicular lobes subtriangular, without prominent accessory lobes; neuropodial ligules subequal in size

and shape to notopodial ligules. Ventral cirri short,

subulate. Upper bundle of neurosetae (4-5) homogomph spinigers (Figure 18g) and single heterogomph falciger (Figure I8h) ; lower bundle of neurosetae heterogomph falcigers (4). Following parapodia

of anterior region (Figure \8d,e) with notopodial

ligules large, subtriangular; upper bundles of neurosetae with homogomph spinigers (4-5) and single

heterogomph falciger; lower bundles with heterogomph falcigers (4—6) with somewhat shorter blades

(Figure 18i).

Parapodia of middle and posterior regions gradually modified (Figure 18/) ; upper notopodial ligules

and dorsal cirri somewhat fused; lower notopodial

and neuropodial ligules much smaller. Notosetae a

single homogomph falciger with short rounded blade

(beginning about setiger 20) ; in far posterior region,

blades of homogomph falcigers more or less fused to

stems, forming simple setae (Figure 18j; not mentioned by Rullier). Upper bundle of neurosetae

homogomph spinigers (5-6) and single heterogomph

falciger; lower bundle of neurosetae heterogomph

spinigers (2) with short blades (Figure 18&; not

mentioned by Rullier) and single heterogomph

falciger.

Pygidium with paired glands, ventral anal cirri and

dorsal anus. Pharynx without paragnaths or papillae,

basal ring (observed on slit pharynx) appearing indistinctly lobulated; jaws with 7-8 teeth. Repro-
NUMBER 10 4 35

duction in atokous condition, having huge eggs (300/t

in diameter, according to Rullier).

DISTRIBUTION.—Indian Ocean (Mauritius).

Rullierinereis mexicana (Treadwell), new combination

FIGURES 19-21

Leptonereis maculata.—Treadwell 1928:469. [Not Kinberg,

1866].

Leptoneris mexicana Treadwell, 1942:1, figs. 2-9.

Nicon mexicana.—Hartman 1956:253, 279; 1958:265;

1959a: 245, 274.—Berkeley and Berkeley 1958:402 (part;

not smaller male heteronereid).

MATERIAL EXAMINED.—Lower California, outside

Topolobampo Bay, 25°30'N, 109°12'W, 17 November

1935—holotype (male heteronereid) and paratype

(female heteronereid) (AMNH 3222). Marina Madre

Island, western Mexico, at night-light off shore, 20

September 1956, W. L. Klawe, collector—male heteronereid (USNM 35813).

Pacific side Panama, Balboa, Canal Zone, at oil

dock, 4 February 1937, S. F. Hildebrand, collector—

2 male heteronereids (USNM 24481).

Florida, Boca Ciega, Tampa Bay, swarming at 6

to 10 P.M., 3 April 1968, J. L. Taylor, collector—

3 male (USNM 43406) and 28 female heteronereids

(USNM 43405).

Galapagos, Arcturus Oceanographic Expedition:

Seymour Bay, 0°27'N, 90°19'W—female and more

than 60 male heteronereids (AMNH 2621; USNM

43407); 0°19'N, 89°57'W, on surface—1 male heteronereid (AMNH 2614).

TYPE-MATERIAL.—The male heteronereid (holotype) has a length of about 50 mm, a width of 3 mm,

including setae, and has very numerous segments;

the pharynx is extended. The female heteronereid

(paratype) is incomplete posteriorly, having a length

of 26 mm and a width of 3 mm, including setae;

the pharynx is not extended but, when slit, no paragnaths or papillae were observed, although the

ridges may appear as indistinct large papillae; large

eggs are present in the body cavity.

DESCRIPTION OF FEMALE HETERONEREIDS.—Length

up to 70 mm, width up to 4 mm, including setae,

segments up to 200. Body formed of three regions:

anterior unmodified (about 50 segments), middle

modified (about 55 segments) and posterior unmodified (up to 100 segments). Prostomium with short

tapered frontal antennae, stout biarticulate palps,

and large eyes (Figure 19a). Tentacular segment

about same length as following segment, with rather

short tentacular cirri, longest extending to setiger

4-5. Parapodia of first two setigers with dorsal cirrus

and single notopodial ligule; neuropodium similar

to those of following segments (Figure \9d,e). Dorsal

cirri on anterior five setigers slightly modified—thicker

basally, with curved tips; ventral cirri of anterior two

to four setigers slightly modified—thicker than those

of following setigers.

Parapodia of anterior region (Figure 19/,g) with

tapered dorsal cirri extending slightly beyond notopodial ligules, two conical notopodial ligules, conical

anterior neuropodial acicular lobe, shorter truncate

postsetal lobe, and lower conical neuropodial ligule;

ventral cirri slightly shorter than dorsal cirri. Notopodia with relatively few homogomph spinigers having long spinous blades (Figure \9h). Upper bundle

of neurosetae homogomph spinigers, similar to notosetae, and heterogomph falcigers with rather long

blades (Figure 19A:). Lower bundle of neurosetae

heterogomph spinigers with relatively short blades

(Figure 19i,;) and heterogomph falcigers similar to

those of upper bundle.

Parapodia of middle modified region with numerous swimming setae completely replacing normal

setae in both notopodia and neuropodia (Figure

20a-c). Conical dorsal lamella developed medial to

base of dorsal cirrus; notopodial ligules more widely

separated by enlarged, rounded supraacicular notopodial lobe bearing numerous modified swimming

setae. Neuropodial acicular lobe paddle-shaped, bearing fan-shaped bundle of modified swimming setae;

large oval postsetal lamella developed from postsetal

neuropodial lobe and extending to about level of

notopodial ligules; lower neuropodial ligule large,

subconical; base of ventral cirrus enlarged, forming

asymmetrical winglike ventral lamella.

Parapodia of posterior unmodified region similar

to those of anterior region; dorsal cirri longer, more

slender (Figure 20d). Notosetae few in number

(sometimes absent), consisting of single homogomph

spiniger and stouter homogomph falciger with spinous

blade and blunt curved tip (Figure 20e). Upper

bundle of neurosetae homogomph spinigers with long

blades and single stout heterogomph falciger (Figure

20/). Lower bundle of neurosetae mostly heterogomph spinigers with blades long (upper ones) to

short (lower ones); lowest neuroseta sometimes

heterogomph falciger.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

9

FIGURE 19.—Rullierinereis mexicana (female heteronereid from Tampa Bay, Florida, USNM

43405): a, Anterior end, dorsal view, pharynx extended; b, same, ventral view; c, posterior

end, dorsal view; d, first parapodium, posterior view; e, second parapodium, anterior view;

/, parapodium from anterior region, posterior view; g, same, anterior view; h, homogomph

spiniger from same; i, heterogomph spiniger from upper part of lower bundle of neurosetae

from same; ;', same, from middle part of lower bundle; k, heterogomph falciger from lower

part of lower bundle from same. [Figures h-k, not to scale]

Pygidium with papillated area around anal opening and ventral anal cirri (Figure 19c). Pharynx

stout, muscular, without paragnaths or definite papillae; jaws dark amber-colored, each with about 11

teeth (Figure I9a,b).

DESCRIPTION OF MALE HETERONEREIDS.—Length

up to 55 mm, width up to 4 mm, including setae, segments up to 180. Body formed of two regions (posterior unmodified region lacking) : anterior unmodified (about 50 segments) and posterior modified

(about 135 segments). Prostomium and tentacular

segment similar to those of female (Figure 2\a,b).

Dorsal cirri of anterior seven setigers and ventral cirri

of anterior five setigers thick, with terminal papillae

or slender curved tips (Figure 2\d,e); dorsal cirri of

setiger 7 slightly longer than preceding ones. Parapodia of anterior unmodified region similar to those

of female heteronereids (Figure 2\d-g).

Parapodia of posterior modified region with numerous swimming setae completely replacing normal

setae in both notopodia and neuropodia (Figure

21 h,i). Bulbous dorsal lamella developed medial to

base of dorsal cirrus; dorsal cirrus elongated, lobulated—about 6-8 rounded lobes along one side; noto-
NUMBER 10 4 37

FIGURE 20.—Rullierinereis mexicana (female heteronereid from Tampa Bay, Florida, USNM

43405) : a, Modified parapodium, posterior view; b, same, anterior view; c, swimming seta

from same; d, parapodium from posterior region, anterior view; e, stout notopodial homogomph

falciger from same; /, stout heterogomph falciger from upper bundle of neuropodia from

same. [Figures c,e,f, not to scale]

podial ligules enlarged, with rounded supraacicular

notopodial lobe bearing numerous modified swimming setae. Neuropodial acicular lobe paddle-shaped,

bilobed distally in region of neuroaciculum, bearing

fan-shaped bundle of modified swimming setae; very

large postsetal lamella developed from postsetal neuropodial lobe and extending beyond level of notopodial ligules; lower neuropodial ligule large; subconical; base of ventral cirrus greatly enlarged, forming asymmetrical winglike ventral lamella.

Pygidium with large papillated area around anal

opening (more extensive than in female) and anal

cirri (Figure 21<r). Pharnyx similar to that of female.

REMARKS.—The figure of the first parapodium by

Treadwell (1942: fig. 3) was inverted. The first two

setigers have uniramous or subbiramous parapodia,

not just the first setiger, as stated by Treadwell

(1942: 3). The notopodial homogomph falcigers of

the posterior unmodified parapodia in the female

heteronereids were not noted in previous publications.

Leptonereis mexicana was referred to Nicon by

Hartman (1956:279). Later, Hartman (1959a: 245)

indicated that N. mexicana was perhaps referable to

N. moniloceras (Hartman) ; however, the two species

do not have the same type of compound falcigerous

neurosetae.

DISTRIBUTION.—Known only from epitokous stages.

Galapagos, Lower California, western Mexico, Pacific side of Panama, Gulf of Mexico (Tampa Bay,

Florida).
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Rullierinereis mexicana (a, male heteronereid from western Mexico, USNM

35813; b-i, male heteronereid from Tampa Bay, Florida, USNM 43406): a, Anterior end,

lateral view; b, anterior end, dorsal view; c, posterior end, dorsolateral view; d, first parapodium, posterior view; e, second parapodium, anterior view; /, parapodium from anterior region,

posterior view; g, same, anterior view; h, parapodium from modified region, posterior view

(setae not shown); i, same, anterior view.

-via
NUMBE R 10 4 39

Rullierinereis gallardoi new species

FIGURE 22

Nicon maculatus.—Gallardo 1968:65, pi. 15: figs. 3-7. [Not

Kinberg, 1866].

MATERIAL EXAMINED.—Nha Trang, South Viet

Nam, 26 meters, coarse sand with shell and coral

debris, station 164, 25 February 1960, V. A. Gallardo,

collector—holotype (AHF 0252).

DESCRIPTION.—Length of holotype 40 mm, width 2

mm, including setae, segments about 64. Body colorless except for faint brownish wide band on setiger 3

and narrow bands on following anterior segments.

Body cylindrical, tapering gradually posteriorly. Prostomium (Figure 22a) with short tapered frontal antennae and stout biarticulate palps; eyes moderately

large, anterior pair slightly larger than posterior pair.

Tentacular segment nearly twice length of following

segment; tentacular cirri rather short, longest extending to setiger 2. Parapodia of first two setigers with

large subulate dorsal cirri and single notopodial

ligule; neuropodia similar to those of following

setigers (Figure 226).

Parapodia biramous from setiger 3 on (Figure

22c,d). Dorsal cirri subulate, extending beyond notopodial ligules. Upper and lower notopodial ligules

subconical and subequal in size; notosetae few in

number (1-2), homogomph spinigers with rather

short blades (Figure 22e). Neuropodia with conical

anterior acicular lobes and short rounded postsetal

lobes (no conspicuous accessory lobes) ; neuropodial

ligules subequal in size and shape to notopodial

ligules. Ventral cirri short, subulate. Upper bundle

neurosetae homogomph spinigers (3-4) with rather

long blades (Figure 22/) and heterogomph falcigers

(2) with moderately long blades (Figure 22g) ; lower

bundle neurosetae heterogomph spinigers (1—2) with

rather short blades (Figure 22h) and heterogomph

falcigers (4-5).

Parapodia of middle and posterior regions slightly

modified (Figure 22*',/) ; dorsal cirri longer and more

slender, cirriform; lower neuropodial ligules smaller

than notopodial ligules. Notosetae single stout homogomph falciger with short, elongate-oval blade (beginning about setiger 18; blades often missing; (Figure 22;,m). Upper bundle of neurosetae homogomph

spinigers (7-8) and stouter heterogomph falcigers

(1-2) with slightly shorter blades (Figure 22A:,n) ;

lower bundle of neurosetae heterogomph spinigers

(3-5) and heterogomph falcigers (2-3).

Pygidium with ventral anal cirri and dorsal anus.

Pharynx without paragnaths or papillae; horny jaws

each with about 6 teeth (Figure 22a). Body cavity

of holotype with moderately large eggs (Figure 22i).

ETYMOLOGY.—The species is named for Victor A.

Gallardo, the collector and worker on the Polychaeta.

DISTRIBUTION.—South Viet Nam.

Stenoninereis Wesenberg-Lund, 1958

TYPE-SPECIES.—S. martini Wesenberg-Lund, 1958,

by monotypy. Gender: feminine.

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two to three

pairs of eyes. Tentacular segment achaetous and

apodous; four pairs of tentacular cirri with distinct

cirrophores. Parapodia of first two setigers subbiramous; notopodium with dorsal cirrus and delicate

notoaciculum. Dorsal cirri with elongate cirrophores

and short subulate distal styles. Notopodia bilobed,

forming lower acicular lobes continuous with presetal

distal lobes and upper notopodial ligules present in

anterior region but disappearing in posterior region.

Neuropodia with acicular lobes but without ligules.

Ventral cirri short, tapered. Notosetae homogomph

spinigers only. Neurosetae homogomph and heterogomph spinigers and heterogomph falcigers. Pygidium

with paired winglike lobes and anal cirri. Pharynx

with paired jaws, without paragnaths or papillae.

A single species, S. martini, and one synonym,

Nicon lackeyi Hartman, are referred to Stenoninereis.

The genus is unique among the nereid genera in having dorsal cirri with elongate cirrophores and short

distal styles. The parapodia have bilobed notopodia

anteriorly and simple neuropodia, lacking extra neuropodial ligules.

Stenoninereis martini Wesenberg-Lund

FIGURES 23, 24

Stenoninereis martini Wesenberg-Lund, 1958:9, figs. 2—4.

[Published July 1958]

Nicon lackeyi Hartman, 1958:263, figs. 1-5. [Published Aug.

ust 1958]

MATERIAL EXAMINED.—Devil's Hole Swamp near

Simson Bay Bridge, St. Martin Island, West Indies,

4 August 1949, P. W. Hummelinck, collector—2 syntypes of Stenoninereis martini (USNM 29726).

Warm Mineral Springs, Sarasota County, Florida,
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Rullierinereis gallardoi new species (holotype, AHF 0252): a, Anterior end, dorsal

view; pharynx extended; b, first parapodium, posterior view; c, parapodium from anterior region, posterior view; d, same, anterior view; e, notopodial homogomph spiniger from same;

/, neuropodial homogomph spiniger from upper bundle of same; g, neuropodial heterogomph

falciger from same; h, neuropodial heterogomph spiniger from lower bundle of same; i, parapodium from middle region, anterior view; ;, notopodial homogomph falciger from same;

k, neuropodial heterogomph falciger from same; /, parapodium from posterior region, anterior

view; m, stout notopodial homogomph falciger from same, with blade missing; n, stout neuropodial heterogomph falciger from same; o, neuropodial homogomph spiniger from upper bundle

of same; p, neuropodial heterogomph spiniger from lower bundle of same. [Figures e-h, j , k,

m-p, not to scale]

O P
NUMBER 10 4 41

J. Lackey, collector—holotype and 2 paratypes of

Nicon lackeyi (USNM 29627-8).

TYPE-MATERIAL.—One of the syntypes of S. martini (in two pieces) has a total length of 4 mm, a

width of 1 mm, including setae, and 26 segments.

The other syntype has a length of 3.5 mm, 1 mm in

width, and 24 segments.

The holotype of N. lackeyi is a female filled with

eggs, 6 mm in length, 2 mm in width, including setae,

and 32 segments. The two paratypes are 3 and 4 mm

in length, 1 mm in width, including setae, comprising

23 and 30 segments.

DESCRIPTION.—Length up to 6 mm, width up to

2 mm, including setae, segments up to 34. Prostomium (Figures 23a,b; 24a,b) with anterior medial

notch, tapered frontal antennae, stout biarticulate

palps, and moderately large eyes, anterior pair larger

than posterior pair; sometimes additional small pair

near posterior border. Tentacular segment shorter

than following segments; tentacular cirri with long

cylindrical cirrophores and filiform styles, longest extending to about setiger 7 (4-9). Parapodia of first

two setigers with slender notoaciculum within short,

conical, basal cirrophore of dorsal cirrus; style short

(sometimes missing) ; neuropodia similar to those of

following segments (Figure 23a,f,g).

Parapodia of anterior region with rami well separated (Figure 23h,i). Dorsal cirri with elongate, tapered basal cirrophores and short, subulate distal

styles; notopodia elongate, distally bilobed, forming

lower acicular lobe continuous with presetal distal

lobe and upper notopodial ligule, lobes digitiform and

subequal in length; notopodial lobes and lower border

of dorsal cirrophores ciliated. Neuropodia with diagonally truncate anterior acicular lobes and shorter

rounded postsetal lobes. Ventral cirri short, tapered.

Notosetae homogomph spinigers with long, finely

spinous blades (Figure 23;). Neuropodium with fanshaped group of neurosetae; upper few heterogomph

spinigers with short blades (Figure 23k), supraacicular homogomph spinigers with long blades (Figure

23;'), few subacicular heterogomph spinigers with

blades relatively short, curved, with long basal spines

(Figure 23/), and heterogomph falcigers with blades

long (upper ones) to short (lower ones; Figure 23m),

finely spinous, and with hooked tips.

More posteriorly, dorsal cirrophores gradually

shortened, upper notopodial ligules gradually diminishing and finally nearly absent (Figure 23n).

Pygidium with flat winglike expansions lateral and

ventral to anus, with anal cirri attached laterally

(Figures 23d,e; 24<:). Pharynx without paragnaths

or papillae; paired jaws each with about 9 teeth

(6-10; Figure 23c).

REMARKS.—The descriptions of Stenoninereis martini Wesenberg-Lund and Nicon lackeyi Hartman

were both published in 1958, the former in July, the

latter in August. Examination of the type-specimens

of both species indicates that the species are synonymous. The parapodial setae were incompletely described and not figured by Wesenberg-Lund. Hartman emended Nicon Kinberg to include N. lackeyi,

as well as nine other nereid species. The only distinguishing character they have in common is an

absence of paragnaths or papillae on the pharynx. In

the present revision, some of these species are assigned

to other genera.

BIOLOGY.—S. martini, a minute aberrant nereid,

lives under unusual conditions. The type-locality of

S. martini is Devil's Hole Swamp, a sinkhole pond of

limestone area about 150 meters from shore on St.

Martin Island. It is subject to tidal movements, with

a bottom of rocks with mud, detritus, and algae, a

pH of about 8 and salinity of 13.8%e (WesenbergLund, 1958: 6). The type-locality of N. lackeyi is

Mineral Springs, Sarasota County, Florida, a large

first magnitude spring emerging from a deep boil,

having a constant temperature of 86° F, pH about

7.2, and salinity about 17%o containing some hydrogen sulfide but no dissolved oxygen, and characterized by a unique flora and fauna (Lackey, 1957:255;

Hartman, 1958:265). A flourishing population of N.

lackeyi was found in this environment, which is unusual for its high temperature and concentration of

certain minerals. The females have giant-size eggs.

Development apparently is direct, without epitoky.

Adults, kept alive in the laboratory by Dr. James

Lackey, continued to give rise to young stages (Hartman, 1958:265).

DISTRIBUTION.—Florida (Warm Mineral Springs,

Sarasota County), West Indies (Devil's Hole Swamp,

St. Martin).

Tylorrhynchus Grube, 1868

TYPE-SPECIES.—T. chinensis Grube, 1868, by monotypy. Gender: masculine. = T. heterochetus (Quatrefages, 1865).

Chinonereis Chamberiin, 1924. Type-species: C. edestus
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.—Stenoninereis martini (syntypes of S. martini, USNM 29726) : a, Anterior end,

dorsal view, parapodia on right side not shown; tips of dorsal cirri of setigers 3 and 4 broken

off; b, anterior end, ventral view; styles of all but two of tentacular cirri missing; c, tip of jaw;

d, posterior end of female, dorsal view; right anal cirrus broken off and tip of left one broken;

e, posterior end of male, ventral view; anal cirri broken off; /, first parapodium, anterior view;

dorsal cirrus either absent or broken off; g, second parapodium, posterior view; h, parapodium

from anterior region (setiger 5), posterior view; i, same (setiger 6), anterior view; j , homogomph spiniger from same; k, heterogomph spiniger from upper neuropodial bundle from

same; /, heterogomph spiniger from upper part of lower neuropodial bundle from same; m, heterogomph falcigers from lower neuropodial bundle from same; n, parapodium from posterior

region, posterior view. [Figures c,)-m, not to scale]

Chamberlin, 1924, by original designation and monotypy.

Gender: feminine. =T. heterochetus (Quatrefages, 1865).

DIAGNOSIS.—Prostomium subpyriform, with paired

frontal antennae, biarticulate palps, and two pairs of

eyes. Tentacular segment achaetous and apodous;

four pairs of tentacular cirri with distinct cirrophores.

Parapodia of first two setigers subbiramous; notopodium with dorsal cirrus and ligule. Dorsal cirri with
NUMBER 10 4 43

anC

FIGURE 24.—Stenoninereis martini (paratype of Nicon lackeyi, USNM 29628): a, Anterior

end, dorsal view; only bases of tentacular cirri shown; b, same, ventral view; c, posterior end,

ventral view.

large thick cirrophores, enclosing large parapodial

glands, and distal styles. Notopodia with low, rounded

acicular lobes and subacicular ligules. Neuropodia

thick, bilobed distally forming small rounded supraacicular and subacicular lobes; without ligules. Ventral cirri short and subulate. Notosetae hemigomph

spinigers (homogomph, with accessory spine). Neurosetae hemigomph and heterogomph spinigers and

heterogomph falcigers with relatively long blades.

Pygidium with paired anal cirri. Pharynx with paired

jaws, without paragnaths but with soft papillae on

both maxillary and oral rings.

REMARKS.—Tylorrhynchus differs in a number of

ways from the usual pattern in the Nereididae. The

following points may be mentioned: 1. single subacicular notopodial ligules (no neuropodial ligules) ; 2.

dorsal cirri with large cirrophores, enclosing large

parapodial glands with lateral openings, and distal

styles (the cirrophores corresponding to the usual

upper notopodial ligules) ; 3. thick neuropodia, bilobed distally, with circular arrangement of the neurosetae (not the usual fan-shaped upper and lower

bundles) ; 4. all setae with articulations hemigomph or

heterogomph (none strictly homogomph) ; 5. sexual

epitokes formed by transformation of the anterior

part of the body, with the posterior part cast off.

The genus is represented by a single species, T.

heterochetus (Quatrefages), and five synonyms.

Quatrefages (1865) described Nereis heterocheta,

based on a single epitoke from Java; the description

was brief and no figures were provided. Grube (1868)

described and figured Tylorrhynchus chinensis, based

on a single male epitoke from Shanghai, China, and

established the genus Tylorrhynchus. Grube (1870),

after examining Quatrefages' holotype in the Paris

Museum, indicated that Nereis heterochaeta [sic]

belonged to his genus Tylorrhynchus. Izuka (1903)

described Ceratocephale osawai from the west coast

of Japan, including atokous and swarming epitokous

stages of both males and females, the Japanese

Palolo; he tentatively referred his species to Ceratocephale in preference to creating a new genus for it.

Ramsay (1914b) synonymized the three species but

chose to use T. chinensis because of Grube's more

complete description, including figures. Based on a

study of specimens from Indochina (Tonkin and
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Annam), Gravier and Dantan (1932) synonymized

the three species and followed the law of priority by

using Tylorrhynchus heterochaetus [sic] (Quatrefages). T. sinensis (error for T. chinensis) was introduced in the literature by Dawydoff (1952).

Chamberlin (1924) described Chinonereis edestus, based on a gravid female (not epitokous) from

freshwater in Canton, China, collected by Mr. Arthur

S. Campbell who reported that the worms were common in the rice-fields surrounding Canton and in the

low-lands as far as Hongkong and were used as food.

Chamberlin's description was brief, incomplete, and

confusing in some respects; there were no figures.

The type-specimen is not present in the Museum of

Comparative Zoology, Harvard, where it was supposed to have been deposited (H. W. Levi, in correspondence) . Chinonereis is herein referred to Tylorrhynchus and C. edestus to T. heterochetus. The latter species was reported from Canton, Amoy, and

Shanghai, China, by Wu and Chen (1963), where

it is known as a mixohaline and freshwater species;

they suspected that the two species were synonymous.

Treadwell (1936) described Nereis (Leptonereis)

distort a from Amoy, China, based on a single male

epitoke collected by T. Y. Chen. Examination of the

holotype, deposited in the United States National

Museum, showed it to be the same as T. heterochetus

and is described below. It is of interest to note that

Monro (1934) identified a specimen, also from

Amoy by the same collector, as T. heterochaetus

[sic].

T. bahamensis Hartmann-Schroder (1958) from

the Bahamas was incorrectly referred to Tylorrhynchus, as shown by the presence of lower neuropodial

ligules, notopodial homogomph falcigers, and neuropodial heterogomph falcigers with short blades. The

species is herein referred to Rullierinereis (see page

31).

Tylorrhynchus heterochetus (Quatrefages)

FIGURE 25

Nereis heterocheta Quatrefages, 1865:552.

Tylorrhynchus chinensis Grube, 1868:22, pi. 2: fig. 3.—

Ramsay 1914b: 231 (synonymy).

Nereis heterochaeta [sic].—Grube 1870:312.

Ceratocephale osawai Izuka, 1903:1, pis. 1, 2.

Chinonereis edestus Chamberlin, 1924:81.

Tylorrhynchus heterochaetus [sic].—Gravier and Dantan

1932:671, figs. 1-3 (synonymy).—Monro 1934:363.—Wu

and Chen 1963:20, 31, pi. 2: figs. B-G.

Nereis (Leptonereis) distorta Treadwell, 1936:273, fig. 20af.

Tyllorhynchus [sic] sinensis.—Dawydoff 1952:88 (error for

T. chinensis]

Tylorrhynchus heterochetus.—Hartman 1959a: 260, 280.

MATERIAL EXAMINED.—Near Amoy, China, T. Y.

Chen, collector.—holotype of Nereis (Leptonereis)

distorta (USNM 20118).

TYPE-MATERIAL.—The holotype of Nereis (Leptonereis) distorta consists of a male epitoke, 70 mm

in length, 8 mm in width, including parapodia, and

61 setigers, with posterior end missing (characteristic

of epitokes). The pharynx was not extended but

had been slit open, the pharynx removed and is

now missing. Modified parapodia with swimming

setae begin on setiger 25.

DESCRIPTION OF MALE HETERONEREID.—Prostomium with deep middorsal depression on anterior

half, subulate frontal antennae, and stout biarticulate palps; eyes large, anterior pair much larger than

posterior pair (Figure 25a). Tentacular segment

longer than following segment; tentacular cirri with

styles relatively short, longest extending to about setiger 5. Parapodia of first two setigers with dorsal

cirrus and ligule only; neuropodium similar to those

of following segments (Figure 256).

Parapodia of anterior unmodified segments (Figure 25c-e) with rami indistinctly separated, thick,

opaque, only tips of the acicula evident externally.

Dorsal cirri with thick subrectangular cirrophores

and short subulate distal styles; parapodial glands

in basal part, with openings laterally. Notopodia with

low rounded postsetal lobe and shorter presetal lobe

continuous with lower conical ligule. Neuropodia

thick, with subtriangular posterior and bilobed anterior lobes, above and below neuroaciculum. Ventral cirri short, subulate. Notosetae relatively few,

short, arranged in vertical row above notoaciculum.

Notosetae slightly heterogomph or hemigomph spinigers, with blades rather short (Figure 25/). Neurosetae nearly encircling upper and lower neuropodial

lobes with gap in middle anteriorly, setal bundle arranged in form of "3" . Dorsoposterior neurosetae

hemigomph spinigers with long, finely spinous blades

(Figure 25g); ventroposterior neurosetae heterogomph spinigers with long blades, with more prominent spines basally (Figure 25/z) ; upper and lower

anterior neurosetae heterogomph falcigers with relatively long blades (Figure 25i).

Parapodia of modified region somewhat enlarged
NUMBER 10 4 45

g

FIGURE 25.—Tylorrhynchus heterochetus, male heteronereid [holotype of Nereis (Leptonereis)

distorta, USNM 20118]: a, Anterior end, dorsal view; first three setigerous segments compressed unnaturally; b, first parapodium, posterior view; c, parapodium from anterior region

(setiger 16), posterior view; d, same, anterior view; e, diagrammatic cross-section of same,

showing arrangement of setae; /, hemigomph spinigerous notoseta; g, hemigomph spinigerous

neuroseta; h, heterogomph spinigerous neuroseta; i, heterogomph falcigerous neuroseta; j , parapodium from middle modified region, posterior view; k. same, anterior view; I, heterogomph

swimming seta from same. [Figures f—i, /, not to scale]

and flattened but similar to anterior parapodia, without addition of extra lamellae (Figure 25j,k). Cirrophores of dorsal cirri wider, with more prominent

parapodial glands; notopodial ligules more pointed;

usual setae replaced by numerous heterogomph swimming setae (Figure 25/). Pharynx with denticulate

jaws and soft papillae on both maxillary and oral

rings, papillae somewhat variable in number and size

(pi. 2: fig. 3a,b, in Grube, 1868: pi. 2: figs. 10, 11,

in Izuka, 1903; pi. 2: figs. B,C, in Wu and Chen,

1963).

BIOLOGY.—T. heterochetus is a shallow water,
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

brackish and freshwater nereid found burrowing in

the bottom mud of the mouths and tributaries of

rivers, including canals, ditches and marshes. They

burrow in the bottom mud and move about on the

surface of the substrate during flood tides. They are

active and voracious, feeding on various aquatic animals and plants. Sometimes they have become pests

in the rice fields. This atokous stage has been variously called "saltwater millepede," "Itome" by the

Japanese (meaning thread worm), and "Wohchung"

by the Chinese (meaning rice or field worm). The

atokous forms have been reported up to 250 mm in

length, 4 mm in width, and 300 segments.

At sexual maturity, the anterior portions of the

worms, representing about a third of the total body

segments, become filled with eggs or sperm and transformed into heteronereids or epitokes, modifications

including the enlargement of the eyes, expansion of

the parapodia, enlargement of the segments, and development of swimming setae. The posterior twothirds of the body takes no part in the transformation

and undergoes degeneration. These transformed

adults with their shrunken tail regions, when still on

the bottom, are referred to as "Hori-Bachi" (meaning dug-out Bachi).

During reproduction, in a lunar periodicity, the

anterior modified portions break away from the posterior regions, leave the bottom and swarm in large

numbers near the water surface, when they are

called "Bachi" or Japanese Palolo. They vary in

length from 33 to 130 mm, with a width of 3 to 11

mm, and consist of 60—78 segments. The swimming

setae begin about setiger 24. The posterior ends are

missing except sometimes remnants may remain.

DISTRIBUTION.—Indo-Pacific: Java, China, Indochina, Japan. Estuarine and freshwater.

Platynereis Kinberg, 1866

Platynereis cebuensis (Grube) new combination

FIGURES 26, 27

Nereis (Leptonereis) cebuensis Grube, 1878:61.

Laeonereis cebuensis.—Hartman 1959a: 243, 262.

MATERIAL EXAMINED.—Cebu, Philippine Islands,

Semper, collector—2 syntypes (ZMB Q3465).

TYPE-MATERIAL.—The larger syntype is a partially metamorphosed male heteronereid of about 83

setigers, 24 mm in length and 4 mm in width, including setae; the pharynx is extended. The smaller syntype is atokous, with about 40 setigers, a length of

13 mm and a width of 2 mm, including setae.

DESCRIPTION.—Body pigmented with reddish

brown dots on prostomium and anterior segments,

forming transverse bands; more posteriorly, pigmented areas confined to lateral parts of body medial

to parapodia, notopodial and neuropodial ligules.

Prostomium (Figure 26a,b) suboval, with paired subulate frontal antennae, stout biarticulate palps; two

pairs of eyes moderately large (small atokous syntype) to large (partially metamorphosed male syntype). Tentacular segment subequal in length to following segment; four pairs of tentacular cirri, upper

two pairs conspicuously long, extending beyond setiger 11 (tips broken). Parapodia of anterior two setigers subbiramous, notopodium represented by subulate dorsal cirrus and single notopodial ligule (Figure

26e,f) ; neuropodium with anterior conical acicular

lobe and prominent postsetal lobe and neuropodial

ligule; ligules and postsetal lobe elongate-conical.

Upper bundle of neurosetae homogomph spinigers;

lower bundle of neurosetae heterogomph spinigers

with long (upper ones) to short (lower ones) blades.

Ventral cirri subulate.

Dorsal cirri of anterior seven and ventral cirri of

anterior five setigers slightly modified—thicker, cylindrical on basal halves, with curved tips (Figure 26

e-h). Biramous parapodia of more anterior setigers

(Figure 26g,h) with two thick, rounded notopodial

ligules, conical anterior neuropodial acicular lobe,

shorter postsetal lobe, and thick, rounded neuropodial

ligule. Notosetae homogomph spinigers (Figure 26i).

Upper bundle of neurosetae consisting mostly of

homogomph spinigers and few heterogomph falcigers

(Figure 26;) ; lower bundle of neurosetae with few

heterogomph spinigers and mostly heterogomph

falcigers. More posterior parapodia of anterior region (Figure 27a,b) with dorsal cirri more elongate,

cirriform; two notopodial ligules elongate-conical,

widely separated, with low rounded notopodial acicular lobe between them; neuropodium with subequal

anterior acicular and postsetal lobes; neuropodial

ligules subconical; ventral cirri tapered. Setae similar to those of more anterior parapodia.

Middle and posterior parapodia of partially metamorphosed male heteronereid showing beginning

stages of developing extra lamellae (Figure 27c,d) :

dorsal cirri slightly scalloped along one margin, exhibiting early development of lobulated dorsal cirri

characteristic of male heteronereids; dorsal lamella
NUMBER 10 4

FIGURE 26.—Platynereis cebuensis [syntypes of Nereis (Leptonereis) cebuensis, ZMB Q3465:

a, smaller syntype; b-j, larger syntype—partially metamorphosed male heteronereid]: a, Anterior end, dorsal view; part of tentacular cirri missing or regenerating; b, dorsal view of anterior end with pharynx extended; some of tentacular cirri with tips broken; c, same, ventral

view; d, posterior end, dorsolateral view, right anal cirrus missing; e, first parapodium, posterior view; /, second parapodium, anterior view; g, parapodium from setiger 6, posterior view;

h, parapodium from setiger 7, anterior view; i, homogomph spiniger from same; ;, heterogomph

falciger from same. [Figures i,j, not to scale]

medial to base of dorsal cirrus somewhat enlarged;

neuropodial postsetal lamella paddle-shaped, and

winglike ventral lamella on base of ventral cirrus

somewhat enlarged. None of setae replaced by swimming setae. Notosetae consisting of numerous homogomph spinigers and few homogomph falcigers,

blades with hooked tips (Figure 27^). Upper bundle

of neurosetae homogomph spinigers and heterogomph

falcigers (Figure 27/); lower bundle of neurosetae

heterogomph spinigers and falcigers.

Pygidium with conical lateral lobes and long anal

cirri (Figure 26d). Pharynx (Figure 26b,c) with pair

of jaws each with four subterminal teeth; faint denticles, characteristic of Platynereis, present on both

rings; maxillary or distal ring: dorsally (areas I and

II) with transverse ridge, without denticles; midventrally (area III) elongate-oval area of five transverse rows; ventrolaterally (area IV) arched groups
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

neLi

vLa

vC

FIGURE 27.—Platynereis cebuensis (larger syntype—partially metamorphosed male heteronereid,

ZMB Q3465) : a, Parapodium from setiger 11, posterior view; b, parapodium from setiger 12,

anterior view; c, parapodium from middle region, posterior view (partially modified, showing

partial development of enlarged lamellae); d, same, anterior view; e, notopodial homogomph

falciger from same; /, neuropodial heterogomph falciger from same. [Figures e,f, not to scale]

of three rows; basal or oral ring: middorsally (area

V) bare: dorsolaterally (area VI) two faint rows;

ventrally (areas VII-VIII) 7 groups of two rows

each.

REMARKS.—The pharynx is extended on the larger

syntype but the minute denticles of the Platynereis

type were overlooked by Grube. The pharynx of the

smaller syntype is not extended; when dissected, it

also revealed the characteristic denticles. Grube

(1878) described Nereis (Platynereis) fusco-rubida

from the Philippines in the same publication in

which N. (Leptonereis) cebuensis was named. Neither

species was figured. Unfortunately, types of the

former species were not found in the Zoologisches

Museum, Berlin (according to G. Hartwich, in correspondence) and the species must be considered

questionable.

DISTRIBUTION.—Philippine Islands.

[ ?] Platynereis sp.

Leptonereis glauca moniloceras Hartman, 1940:217, pi. 34:

figs. 42-46.

Nicon moniloceras.—Hartman 1958:265; 1959a:245, 274.

REMARKS.—The species, described from a female

heteronereid dredged in 73 meters off Catalina Island, Southern California, and associated with

brachiopod and sponge clusters, is questionably referred to Platynereis. The necessity for this reassignment is suggested by the oval prostomium, conspicuously long tentacular cirri, the parapodial lobes and

setae—all agreeing with Platynereis. Perhaps the

minute pharyngeal denticles were overlooked.

Platynereis sp.

Nicon peruviana Berkeley and Berkeley, 1964:128, figs. 1, 2.
NUMBER 104 49

MATERIAL EXAMINED.—South Pacific off Peru, approximately 5°S, 81 °W, at night-light, 10 April

1959, P.N. Sund, collector—holotype and paratype

(USNM 32821-2).

REMARKS.—The types are male heteronereids. The

pharynx of the holotype had been dissected and is

now missing. The pharynx of the paratype was dissected but the denticles are not clearly visible. The

specimens appear, however, to be a typical Platynereis.

Platynereis sp.

Leptonereis davidi Fishelson and Rullier, 1969:62, fig. 1AD.

MATERIAL EXAMINED.—Bay of David, Entedebir

Island, Red Sea, on Sargassum, 13 March 1962—

holotype (No. E62/238, Department of Zoology, TelAviv University, Israel).

REMARKS.—The holotype consists of a small specimen, 8 mm in length, 1 mm in width, including setae,

and about 55 segments with a posterior growing

zone. The pharynx was not extended but had been

dissected; however, no denticles were observed. Since

the specimen is small and the pharynx was not extended, this character would be difficult to observe.

Nevertheless, the parapodial lobes, setae, and prominent glandular masses are of the typical Platynereis

type.

Indeterminable NEREIDIDAE

1. Nicomedes difficilis Kinberg

Nicomedes difficilis Kinberg, 1866:179.—Hartman 1949:57.

MATERIAL EXAMINED.—South Atlantic off Rio de

Janeiro, Brazil, 55-73 meters, Eugenie Expedition—

holotype (NRS 572).

REMARKS.—The holotype consists of the posterior

end of a heteronereid. As indicated earlier by Hartman (1949:57), both the genus and species must be

considered indeterminable because of the brevity of

the original description, absence of figures, and the

poor condition of the holotype.

2. Nicon pictus Kinberg

Nicon pictus Kinberg, 1866:178.—Hartman 1949:57.

MATERIAL EXAMINED.—South Atlantic off Rio de

Janeiro, Brazil, 20°30'S, 40°55'W, 40-55 meters,

Eugenie Expedition—4 syntypes (NRS 486).

REMARKS.—The syntypes consist of two complete

specimens, 19 and 12 mm in length, 1 mm in

width, and about 50 segments, and two anterior

fragments. They are in poor condition, transparent,

and covered with crystals and fibers. Since the species

has not been, nor can it now be, adequately described or figured, N. pictus should be considered

indeterminable.

3. Nicon tahitanus Kinberg

Nicon tahitanus Kinberg, 1866:178.—Hartman 1949:59, pi.

8: figs. 7, 8.

MATERIAL EXAMINED.—Tahiti near Papeete, in

sand and corals, 16.5 meters—holotype (NRS 494).

REMARKS.—The holotype consists of anterior and

middle fragments totaling 36 segments, 8 mm in

length and 1 mm in width. Accompanying the specimen is a thin tube covered with debris. The pharynx

had been removed and is now missing. Eggs are

present in the body cavity, indicating that it was not

a juvenile, as stated by Hartman (1949:59). The type

material is too poor for an adequate description. For

the present, N. tahitanus should be considered indeterminable.

4. Nicon virgini Kinberg

Nicon virgini Kinberg, 1866:179.—Hartman 1949:59.

MATERIAL EXAMINED.—York Bay, Magellan Strait,

C. A. Virgin collector, Eugenie Expedition—holotype (NRS 576). Port Famine, Magellan Strait, 14.6

meters, Eugenie Expedition—4 specimens (NRS

575). Magellan Sound, South America, Eugenie

Expedition—1 specimen (NRS 1647).

REMARKS.—Of the three lots collected by the

Eugenie Expedition and identified by Kinberg as N.

virgini, only NRS 576 was apparently collected in

the type-locality. The holotype consists of about 70

segments, 20 mm long and 1 mm wide. It is in poor

condition being covered with crystals and with the

blades of the setae mostly broken. NRS 575 contains

four anterior fragments in poor shape. NRS 1647

contains anterior and middle fragments of a larger

specimen; the pharynx is extended and paragnaths

are present. In the absence of figures, an adequate
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

description, and the poor condition of the holotype,

N. virgini should be considered indeterminable.

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* U.S. GOVERNMENT PRINTING OFFICE: 1971- 137—337/4

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