JAMES A. BLA Revision of the

Genus Polydora from

f The East Coast of

North America

Polychaeta: Spionidae)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 75
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 75

James A, Blake Revision of the

Genus Polydora from

The East Coast of

North America

(Polychaeta: Spionidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Blake, James A. Revision of the Genus Polydora from the East Coast of North

American (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, number 75, 32 pages, 1971.—Twelve species of Polydora from the east coast of North

America are described on the basis of recent collections as well as material deposited in various museums. The study includes 11 previously described species,

one new species, 10 synonyms, and one nomen nudum. Diagnosis of the genus

Polydora, with a key to the species covered, are included.

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

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

For ale by the Superintendent of Document*, U.S. Government Printing Office

Wathington, D.C. 20408 • Price 45 cent* (paper cover)
James A. Blake Revision of the

Genus Polydora from

The East Coast of

North America

(Polychaeta: Spionidae)

The family Spionidae is among the largest, in number of species, of the Polychaeta. They are typically

marine bottom-deposit feeders and occur mostly in

coastal waters. The family is rare in the deep sea.

The spionids have a pair of long peristomial

palps which are used in feeding. Each palp has a

ciliated groove along which particles move toward

the mouth; some of these particles are used to

build tubes while others are utib'zed as food. The

sorting mechanism, however, is not as well developed as in some other polychaete families.

The Spionidae has been divided into several

subfamilies (Sdderstrom, 1920; Orrhage, 1964). The

subfamily Spioninae contains the genera Pygospio,

Spio, Paraspio, Polydora, Bpccardia, and Pseudopolydora. The three latter genera comprise the

polydorid complex in which the fifth setigerous

segment is modified and includes specialized setae

or spines. The structure of the spines form useful

taxonomic characters. Major systematic treatments

of the polydorids include works by Carazzi (1893),

Mesnil (1896), Sdderstrom (1920), Fauvel (1927),

Hartman (1941), Woodwick (1964), and Day

(1967).

Our knowledge of spedes of Polydora from the

east coast of North America comes largely from the

studies of Verrill (1880, 1881, 1885a, b), Webster

(1879a, b), Andrews 1891a, b), Moore (1907), and

Dr. James A. Blake is on the staff of the Pacific Marine Station, University of the Pacific, Dillon Beach, Marin County,

California, 94929.

Hartman (1942, 1943, 1944, 1945, 1965). These

five authors described eleven species of Polydora.

Four additional spedes, previously known from

other geographic areas, were reported. Special mention should be made of the paper by Bosc (1802),

in which the description of the type spedes of the

genus, P. cornuta Bosc from Charleston, South

Carolina, was given. Unfortunately, the description

of that spedes was superfidal and spedfic determination is today impossible. Thus, the type-spedes

of Polydora is indeterminable.

Considerable confusion exists with regard to the

four spedes of Polydora erected by Verrill. Only

for P. concharum Verrill (1880) are adequate descriptions and figures given (Verrill, 1885a; Hartman, 1942, 1944) and type-material available.

Polydora gracilis Verrill (1880) and P. tubifcx

Verrill (1885b) were described but never figured,

while P. littorea Verrill (1881) is a nomen nudum.

Hartman (1944) belatedly published a number of

Verrill's unpublished plates, which contained some

figures of Polydora, including P. concharum and

P. littorea.

Webster (1879a, b) described P. hamaia,P. caeca,

and P. ligni. Polydora caeca, however, was dedared

a junior homonym by Hartman (1943) and renamed P. websteri. Polydora hamata was referred

to Boccardia by Blake (1966). Polydora aggregata

was named and the larvae described by Blake

(1969b).

The other spedes described from eastern North
2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

America include Polydora commensalis Andrews

(1891a), P. colonia Moore (1907), and P. tetrabranchia Hartman (1945). Four species from other

regions have also been reported to occur in eastern

North America, including P. ciliata (Johnston)

and P. quadrilobata Jacobi, reported by Procter

(1933); P. socialis (Schmarda), reported by Hartman (1945); and P. caulleryi Mesnil, reported by

Hartman (1965).

Despite these numerous references and others,

only Hartman (1945) provides useful keys and descriptions to a number of species. Unfortunately,

that study is limited to North Carolina and excludes several species found in adjacent areas. The

present study is an attempt to reevaluate older records and to provide new information on species of

Polydora for the entire eastern seaboard. New collections come from Newfoundland to South Carolina. Most were collected by the writer, while others

were collected by various individuals, acknowledged

in the text. The bulk of these collections have been

deposited in the United States National Museum.

Field collections of adult Polydora were made

intertidaUy and subtidally from shells and sediment.

As a result of the author's location in Maine, extensive collections were made in that area. Subtidal samples were obtained with either a biological

dredge or a 0.05 m* Ponar Grab. Sediment was

washed with sea water through fine-mesh screens.

The tubes were removed and placed in jars in running seawater.

In the laboratory, the tubes were carefully dissected and the worms removed. Polydora from shells

were removed after cracking the shells with a hammer or pliers.

Adults were relaxed in 0.15% propylene phenoxytol prior to fixation in 10% formalin or

Bouin's fluid. After several days in the fixative, the

worms were removed and stored in 70% ethyl

alcohol.

Older collections of Polydora, deposited in the

Allan Hancock Foundation, University of Southern

California (AHF), Academy of Natural Sciences

of Philadelphia (ANSP), United States National

Museum (USNM), and the Yale Peabody Museum

of Natural History (YPMNH) were also examined.

The following people assisted by arranging for

the loan of specimens from their respective Museum: Marian H. Pettibone (USNM), Willard

Hartman (YPMNH), and Olga Hartman (AHF).

The following people were of assistance during

visits to their Museums by helping locate specimens

and providing working space: Olga Hartman and

Kristian Fauchald, of the Allan Hancock Foundation, during my visit in September 1968; R.

Tucker Abbott, of the Academy of Natural Sciences

in Philadelphia, during my visit in October 1968.

On numerous occasions, Keith H. Woodwick of

Fresno State College, Fresno, California, provided

working space, offered advice and allowed me to

examine his extensive collections of Polydora from

California.

Additional collections were received from David

Grant and Katherine D. Hobson of the SystematicsEcology Program, Woods Hole, Massachusetts

(SEP); Anders Eliason of the Naturhistoriska

Museet, GSteborg, Sweden (NMGS); Francois

Rullier of the University Catholique d'Angers,

France; Luden Laubier, Laboratoire Arago,

Banyuls-Sur-Mer, France; P. Korringa, Rijksinstituut voor Visserij-Onder, Bergen-Op-Zoom, The

Netherlands; John W. Evans, Memorial University

of Newfoundland, St. John's, Newfoundland; John

D. Davis, Smith College, Northampton, Massachusetts; and Sarah A. Haigler, Virginia Institute of

Marine Sdence, Gloucester Point, Virginia.

Specimens of Polydora concharum collected by

the writer in August 1968 aboard the R/V Hero

off West Greenland are also included. Complete

results of the Hero cruise will be published later.

Assistance rendered by Mrs. Crystal M. Blake,

Barbara Hilton, Thomas L. Richards, Michael A.

Mazurkiewicz, David Dean and George S. Noyes

is gratefully acknowledged.

The contents of this paper represent portions of

a dissertation submitted in partial fulfillment of the

degree of Doctor of Philosophy, University of

Maine, Orono, Maine. This study was supported in

part by a University of Maine Graduate Fellowship,

National Sdence Foundation Summer Fellowship,

N.S.F. Graduate Traineeship, and N.S.F. Grant

GB-4892.

Genus Polydora Bosc, 1802

TYPE-SPECIES.—Pofydora cornuta Bosc, 1802.

DIAGNOSIS.—Prostomium rounded or bifid anteriorly, extending back as a caruncle. Eyes present

or absent. First setigerous segment with or without

notosetae. Setiger 5 greatly enlarged, with spedal-
NUMBER 75

ized spines and companion setae arranged in a single

curved row. Posterior notosetae may include specialized spines or hooks as well as capillaries.

Hooded hooks usually begin on setiger 7. Branchiae

begin on segments posterior to setiger 5 and are not

fused to the notopodial lobes. Pygidium enlarged,

saucer-like or divided into lobes.

The following species of Polydora are covered in

this report (synonyms in parentheses; those marked

with an asterisk are new synonyms):

1. P. ligni Webster, 1879b

(•P. littorea Verrill, 1881. Nomen nudum)

(P. amarincola Hartman, 1936)

2. P. webiteri Hartman, 1943

(P. caeca.—Webster. 1879a.—Not Oersted. 1843)

S. P. aggregata Blake, 1969b

4. P. tetrabranchia Hartman, 1945

5. P. caulleryi Mesnil, 1897

(P. carazzi Mclntosh, 1909)

(P. brachycephala Hartman, 1936)

6. P. quadrilobata Jaoobi, 188S

(•P. tubifex, Verrill, 1885b)

(P. littorea.—Hartman, 1944)

7. P. colonia Moore. 1907

(•P. ancistrata Jones, 1962)

(P. hoplura inhaca Day, 1957)

8. P. commensalis Andrews, 1891a

(P. ciliata brevipalpa Zaii. 1933)

9. P. concharum Verrill, 1880

10. P. socialis (Schmarda, 1861)

(Leucodore socialis Schmarda, 1861)

(P. socialis plena Berkeley and Berkeley, 1936)

(?P. gracilis Verrill, 1880)

11. P. anoculata Moore, 1907

12. P. hartmanae new spedes

(P. anoculata.—Hartman, 1945.—Not Moore, 1907)

Key to the Species of Polydora from the East Coast of North America

(Figure numbers in parentheses)

1. Hooded hooks begin on setiger 7 2

Hooded hooks begin on setigers 10-17. Palps very short (Ha). Modified spines of

setiger 5 with long lateral sheaths (116, c) P. commenmUs (p. 17)

2. Hooded hooks with a constriction on the shaft (2d,e) 3

Hooded hooks without a constriction on the shaft (8/) 6

S. Nuchal tentacle present (la) P. Upd (p. 5)

Nuchal tentacle absent 4

4. Modified spines of setiger 5 bifid, with subterminal collars (lOd-f). Posterior notopodial hooks present (lOh-k) P. coUmim (p. 15)

Modified spines of setiger 5 falcate, with lateral accessory structures. Posterior

notopodial hooks absent. 5

5. Modified spines of setiger 5 with lateral sheaths or flanges (36-g). Bores in calcareous

stniaures. P. websteri (p. 6)

Modified spines of setiger 5 with weakly developed lateral shelves or knobs (46-«).

Forms mat of tubes on rocky shores. P. mggregtlm

6. Posterior notopodial spines present, awl-shaped (8g; 9;)

Posterior notopodial spines absent

7. Modified spines of setiger 5 large, falcate, with pectinate or bushy tops (8c, d)

P. caulleryi

Modified spines of setiger 5 bifid, or with bushy tufts between the teeth (9e, g)

P. qusdrOobatm

8. Modified spines of setiger 5 bifid, or with well developed lateral teeth or sheaths.

Modified spines of setiger 5 simple, falcate or with a weakly-developed accessory

thelt 11

9. Branchiae begin on setiger 7. Modified spines of setiger 5 bifid, with two large teeth

(Jb, c) P. tetrmbrmnchU (p. 10)

Branchiae begin on setigers 9-12. Modified spines on setiger 5 falcate, with lateral

sheath or tooth 10

10. Modified spines of setiger 5 falcate, with lateral sheath or tooth and fine hairs in

concavity (166-A). Branchiae begin on setigers 11-12 P. hartmmnme (p. 25)

Modified spines of setiger 5 falcate, with lateral sheath or tooth not covered with fine

hairs (15fr-i). Branchiae begin on setigers 9-10. P. anocuUtm (p. 25)

11. Modified spines of setiger 5 falcate, with weakly-developed lateral shelf (126-d).

Pygidium with four unequal lobes (12;). Caruncle extending no further than setiger

(p.

(p-

(p8)

7

8

11)

13)

9
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ficuu l.—Polydora Kgni: a, Anterior end, dorsal view; b, setiger 1, anterior view; c, posterior

end, dorsal view.
NUMBER 75

5. Larval pigment retained only in juveniles. Posterior capillary notosetae of three

sizes P. concharum (p. 20)

Modified spines of setiger 5 falcate, with subterminal boss (146). Pygidium with 1-3

lobes (13c, d). Caruncle extending to setigers 4-9. Larval pigment retained on most

adults. Posterior capillary notosetae of one size P. sodattt (p. 20)

A

FIGURE 2.—Polydora ligni: a, Companion seta of setiger 5;

b, c, heavy spines and companion setae of setiger 5; d, e,

hooded hooks.

Polydora ligni Webster

FIGURES 1-2

Polydora ciliatum Agassiz, 1867, pp. 323-330, pis. 9-10.—

Verrill, 1873, pp. 51, 309, 345, 364, 453. 603, pi. 14. fig. 78.

Not Johnston, 1838.

Polydora littorea Verrill, 1881, p. 301 (nomen nudum).—

Hartman, 1944, pp. 336, 340, pi. 18: fig. 10, not pi. 18:

fig. 9.

Polydora ligni Webster, 1879b, p. 119. pi. 5: figs. 45-47.—

Webster and Benedict, 1884, p. 729.—Sdderstrom, 1920, pp.

265-267. figs. 170-174.—Cowlcs, 1930, p. 341.—Berkeley and

Berkeley, 1936. pp. 471-472; 1952. p. 19. figs. 31-33; 1954.

p. 464.—Friedrich, 1937, pp. 345-347, figs. 5-6; 1938. p. 133.

—Rioja, 1943, pp. 232-244; 1947, p. 207.—Mortenson and

Galtsoff. 1944, pp. 164-165.—Graham and Gay, 1945, p. 379.

—Hartman, 1941, pp. 309-310, figs. 47-49; 1945, p. 32; 1951.

p. 82; 1954, p. 10; 1961, p. 29; 1969, pp. 137-138, figs. 1-6.—

Hartman and Reish, 1950, p. 28.—Smidt, 1951, pp. 63-65 —

Hannerz, 1956, pp. 106-111, figs. 37-38.—Jones. 1961, p.

266.—Eliason, 1962, p. 52.—Reish, 1963a, pp. 25, 30; 1963b,

pp. 265-270; 1964a. pp. 202-207; 1964b, pp. 86-92.—Wells

and Gray, 1964, p. 73.—Galtsoff, 1964, p. 421-425.—Cory,

1967, pp. 71-89.—Berkeley, 1968, p. 560.—Blake, 1969b, pp.

4-10, figs. 1-4.

Polydora amarincola Hartman, 1936, p. 49, figs. 6-10.

MATERIAL EXAMINED.—Maine (Cobscook Bay;

Lamoine Beach; Penobscot River; Damariscotta

River; Sheepscot River), Massachusetts (Woods

Hole; West Falmouth; Hadley Harbor), Connecticut (Noank), New Jersey (Raritan Bay, coll. D.

Dean), North Carolina (Beaufort).

DESCRIPTION.—Largest specimens measure 32 mm

in length and have up to 80 segments. The prostomium is bifurcated and flares laterally. There are

four eyes, arranged as a trapezoid. A nuchal tentacle is present on the caruncle at about the level

of setiger 1. The caruncle continues posteriorly as

a narrow ridge through setiger S. A ciliated groove

lies on either side of the caruncle (Figure la).

Setiger 1 has well-formed parapodial lobes but

contains only capillary neurosetae (Figure 16).

Setigers 2-4 and 6 contain fascicles of finely winged

capillary setae. After setiger 6 the number of

notosetae gradually diminishes, until in posterior

segments only a tuft of long capillary setae remains.

Bidentate hooded hooks begin on setiger 7 (Figure

2d, e). There are up to 15 hooks per neuropodium

and no accompanying capillary setae. Hooks of the

same neuropodium may have the main fang either

at a right angle to the shaft or bent to about 70°.

A constriction and curve are prominent on the

shaft. A striated hood covers the teeth.

Setiger 5 is modified to include a row of heavy

spines with closely adhering companion setae (Figure 2a-c). The former have a small accessory tooth

which may be considerably worn. The latter have

a delicate feathery end; when depressed, the tip is

seen to be forked.

Branchiae are long and thin, beginning on setiger

7 and continuing to near the end of the body.

The pygidium is a large, flaring cup with a dorsal

gap (Figure lc).

Color in life is light tan with red blood vessels.

Except for lateral black pigment spots often present

in juveniles, the body is unpigmented.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—Polydora littorea Verrill is herein

considered synonymous with P. Ugni. The species

was named by Verrill (1881) but was never described. Verrill erected the name littorea to replace

the P. ciliatum of Agassiz (1867), which he considered to be distinct from the European P. ciliata.

The figures of the adults and larvae of Agassiz's P.

ciliatum appear to be P. Ugni. Plate 18, Figure 10,

was cited by Verrill but not published until Hartman (1944) issued Verrill's plates. A nuchal tentacle is not evident and Figure 10 is indeterminable

m other respects. Figure 9, however, cited by Hartman as P. littorea, is not the same species. It resembles P. quadrilobata Jacobi both in general

shape and structure of the pygidium.

The species in North America most closely related to P. Ugni is P. nuchalis Woodwick from

California (Woodwick, 1953), both species having

a nuchal tentacle. Polydora nuchalis differs in that

no accessory structure is present on the modified

spines of setiger 5. Further, the companion setae

are of different structure.

ECOLOGY.—Polydora Ugni is a common inhabitant of estuaries in North America. It constructs

fragile tubes of silt on tidal flats or attached to

shells, rocks and wood. The species does not bore

into calcareous structures. Galtsoff (1964) reported

that on several occasions in Delaware Bay the reproduction of P. Ugni was so rapid that nearly every

live oyster of the affected area was killed by a deposit of mud several inches thick. Mud particles suspended in the water are caught on the worm's palps

and carried toward the head by ciliary action. The

silt is then formed into a tube in the manner described for other species by Hempel (1957b) and

Dorsett (1961).

DISTRIBUTION.—East and west coasts of North

America, Gulf of Mexico and northern Europe.

Polydora websteri Hartman

FlGUKE S

Polydora ciliata.—Procter, 19SS, p. 142.—Kavanaugh, 1941,

pp. 32-34.—Lunz, 1940. p. S10; 1941, pp. 273-283.—Abbott,

1946, pp. 15-18, pi. 3.—Medcof, 1946, pp. 498-515.—Plaine,

1952. pp. 121-123. fig. 1.—Turner and Hanks, 1959. pp.

109-111.—Landers, 1967, pp. 63-66, figs. 1-2. Not Johnston, 1838.

Polydora caeca.—Webster, 1879a, pp. 252-253, pL 9: figs.

119-122.—Andrews. 1891a, p. 291. Not Oersted. 1843.

Polydora websteri.—Hartman. 1943, pp. 70-72, fig. 1; 1945,

p. 33; 1951, pp. 81-82; 1954. p. 415; 1961. pp. 99-100, pis.

16-17; 1966b, p. 223; 1969, pp. 151-152, 4 figs.—Loosanoff

and Engle, 1943, pp. 69-78.—Grice, 1951. pp. 1-10.—Mackin

and Cauthron, 1952. pp. 14-24.—Menzel, 1955, p. 112.—

Hopkins, 1958, pp. 268-277, figs. 1-6.—Owen, 1957, pp.

35-46.—Rioja, 1960, p. 304.—Mackenzie and Shearer, 1961,

pp. 105-111—Wells and Gray, 1964, p. 73.—Galtsoff. 1964,

pp. 421-425.—Forbes, 1966, pp. 280-281.—Davis, 1967, pp.

67-72, figs. 2-3.—Evans, 1969, pp. 775-782.—Blake. 1969a,

pp. 814-815. fig. 2; 1969b, pp. 10-16, figs. 5-11.—Haigler,

1969. pp. 821-828, figs. 1-3.

MATERIAL EXAMINED.—Maine (Lamoine Beach;

Damariscotta River; Boothbay Harbor region.

Ocean Point), Newfoundland (St. Mary's Bay, 10-

20 meters, coll. J. W. Evans), Quebec (Gaspe

Peninsula, coll. J. D. Davis), Connecticut (Milford,

Type AHF 1569; Noank), Virginia (Gloucester

Point, coll. J. L. Simon), North Carolina (Beaufort), South Carolina (Charleston, coll. M. Chamberlain), California (Morro Bay, coll. K. H.

Woodwick).

DESCRIPTION.—Polydora websteri is a small slender species, measuring up to 20 mm in length and

having about 100 segments. In life it is light tan

with red branchiae, palps and blood vessels. Body

pigmentation is variable, absent in some specimens

while others have dense anterior and posterior black

pigment.

A prostomial anterior incision may be distinct,

weak or absent (Figure 3a). Specimens from New

England and eastern Canada tend to have a weakly

incised or rounded prostomium, while in specimens

from the middle Atlantic and southern states, it

tends to be more strongly incised. Postlarval forms

from Maine have a rounded prostomium. The

caruncle terminates bluntly on setiger 2. Eyes are

present or absent; if present, there are four, disposed as in Figure 3a.

Setiger 1 has only capillary neurosetae and elevated notopodial lobes. Setigers 2-4 have blunt

notopodial lobes with fascicles of winged capillary

setae. The neuropodial lobes are greatly reduced

and contain winged capillary setae. Setiger 6, and

those immediately following, contain dorsal fascicles

of winged capillary setae. The number of setae per

notopodium gradually diminishes in the middle

segments. In far posterior segments, only three or

four winged capillary setae are found in each notopodium. On setiger 7, bidentate hooded hooks replace the neuropodial winged capillary setae. The

hooks number about six in a series and have the
NUMBER 75

FicintE S.—Poiydora websteri: a, Anterior end, dorsal view; b-g, heavy modified spines of setiger

5 showing different views and degrees of wear; h, pennoned companion seta of setiger 5; i,

superior dorsal winged seta of setiger 5; /, hooded hook; k, posterior end, dorsal view; I,

pygidium, posterior view.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

main fang at a right angle to the shaft. Each hook

has a constriction on the shaft (Figure 3;*).

Setiger 5 is twice as large as either 4 or 6; the

parapodia contain a row of heavy modified spines

(Figure 36-g) alternating with pennoned companion setae (Figure 3/t) . One or two superior dorsal,

winged setae (Figure 3t) and a ventral fascicle of

short winged setae are present. The modified spines

are falcate and have a lateral flange or sheath. Figure Sb-g illustrates different views and degrees of

wear of these spines.

Branchiae begin on setiger 7. They are small at

first, reaching full size on setigers 11 and 12 and are

absent from posterior segments.

The pygidium is cup-shaped with a distinct dorsal notch (FigureS*,/).

REMARKS.—During the course of these studies,

no specimens of P. ciliata have been identified from

eastern North America. Hartman (1945) has already suggested that P. ciliata may not even occur

in eastern North America. On this basis earlier

records are herein referred to P. websteri. Both

Blake (1969a) and Haigler (1969) reported that

the hooded hooks of P. websteri begin on setiger 7

and not 8, as previously reported by Hartman

(1943).

ECOLOGY.—The species is best known as a pest of

commercial oysters, but is abundant in numerous

intertidal and shallow water calcareous substrates,

especially in eastern North America. In Maine the

species has been found boring into shells of the

scallop Placopecten magellanicus and the gastropods Littorina littorea, Thais lapillus, Crepidula

fornicata, and Lunatia heros.

DISTRIBUTION.—East coast of North America from

Quebec and Newfoundland to Florida, Gulf of

Mexico, California, Oregon, and Hawaii.

Polydora aggregate Blake

FICUBES 4-6

Blake, 1969b, pp. 16-21, figs. 12-15. (Description of larvae.)

MATERIAL EXAMINED.—Maine (Lamoine Beach

State Park; Boothbay Harbor region, Ocean Point).

The holotype (USNM 42246) and 10 paratypes

(USNM 42247-8) have been deposited in the

United States National Museum. Additional paratypes have been deposited in the Allan Hancock

Foundation.

DESCRIPTION.—This is a small species measuring

6-8 mm in length and having 35-40 segments. The

body has a characteristic reticulate, dark-brown pigment pattern of varying intensity on the peristomium and first six to eight segments (Figure 4a).

The caruncle has two lateral clear areas which in

life are seen to contain ciliary grooves. Posterior

segments are generally not darkly pigmented, although the pygidium is often rimmed with black.

The prostomium is bifid, although in some preserved specimens the anterior end curves downward

and appears rounded when seen from above. The

caruncle ends abruptly on setiger 2. There are four

eyes, the anterior pair cup-shaped and the posterior

pair oval (Figure 4a).

Setiger 1 has only neurosetae, although a well

developed notopodial lobe is present. Setigers 2-4

have blunt notopodial lobes with long and short,

winged capillary setae. The neuropodial lobes are

small, with those of setigers 2-4 becoming progressively shorter. Beginning on setiger 6, the notopodia

contain fascicles of long and short, winged capillary

setae. The number of setae per fascicle gradually

diminishes in the middle region, and in posterior

segments only a few long, thin, capillary setae are

present. The winged capillary neurosetae are replaced by bidentate hooded hooks beginning on

setiger 7 (Figure 4g). Each hook has a constriction

on the shaft. Five or six hooks are present on most

segments, although fewer are found in posterior segments.

Setiger 5 is about twice the size of preceding segments. The parapodia contain a curved row of

heavy spines (Figure 4b-e) alternating with pennoned companion setae (Figure 4/). Tufts of

winged setae are located above and below the heavy

spines. The heavy spines are falcate, with a small

lateral accessory structure appearing as a small

elevation, cleft on one side and rounded on the

other. This structure can be classed neither as a

tooth nor flange. Deeper unworn setae have a long

tapered mucron on their tips.

Branchiae begin on setiger 7 and are absent from

posterior segments.

The pygidium is cup-shaped with a wide dorsal

gap (Figure 4h).

REMARKS.—The name Polydora aggregata was

prematurely introduced in a larval study (Blake,

1969b). The adult description is given here for the

first time.
NUMBER 75

Ficuu 4.—Pofydora aggregata: a, Anterior end, dorsal view; b-e, heavy modified spines of setiger

5 in different views; /, pennoned companion seta of setiger 5; g, hooded hook; h, posterior end,

dorsal view.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

•

FKU U 5.—Lamoine Beach, Maine: a, Foregrouod ihows a

•olid mat of P. aggregate tubes; b, clustered tubes of P. aggregat* between rocks and broken shells of Mytilus edulis.

The snail is Uttorina tittorem.

Polydora aggregate is closely related to P. limicola

(Annenkova, 1954). The two spedes are similar in

habitat and morphology. They differ, however, in

that P. limicola has a small, triangular tooth on the

side of the modified spines of setiger 5, whereas

P. aggregate has the structure described above.

ECOLOGY.—At Lamoine Beach, P. aggregata occurs in dense mats on and between rocks, exposed

at low tide (Figure 5a). The mats are so thick in

some areas they appear to stabilize the beach. At

the lowest tide levels, the dense mats of tubes give

way to tube dusters or dumps, which are attadied

to rocks, shells, or algae (Figures 5b; 6a, b). Occasionally, the orbiniid Naineris quadricuspida

(Fabricius), phyllododds, chironomid larvae and

nematodes are associates in the mats of tubes. At

Ocean Point the spedes was found among holdfasts

of Chondrus crispus.

Ficuu 6.—Lamoine Beach, Maine: a, Rock with tube duster)

of P. aggregata; b, tube dusters of P. aggregata.

DISTRIBUTION.—Maine, on semiexposed rocky

shores.

Polydora tetrabranchia Hartman

FlCUKE 7

Hartman, 1945, p. 54, pi. 1: figs. 7-10.

MATERIAL EXAMINED.—North Carolina (Beaufort,

Types, AHF N1563).

DESCRIPTION.—This is a small spedes, measuring

about 6 mm in length and having about SO segments. The prostomium has a weak anterior incision. There are no eyes or nuchal tentacle. The

caruncle extends to setiger 1.

In the spedmens examined, notosetae were not

seen on setiger 1, although Hartman (1945) indicated they were present Slender, fine, winged,
NUMBER 75 11

FicmtE T.—Polydora tetrabranchia: a, Hooded hook; b, c,

heavy modified spines of setiger 5.

capillary notosetae occur in setigers 2-4, 6, and succeeding segments. The neuropodia of setigers 1-4

and 6 contain winged capillary setae. Hooded hooks

begin on setiger 7 with no more than three hooks

per segment. They have the main fang bent at

nearly a right angle to the recurved shaft (Figure

7a).

The modified spines of setiger 5 are distally bifid,

with two large teeth (Figure 7b-c). There are

usually two or three spines in each parapodium, accompanied by slender pennoned setae.

The pygidium is a simple, collar-like ring.

Branchiae begin on setiger 7 and usually number

only four pair.

DISTRIBUTION AND ECOLOGY.—Polydora tetrabranchia bores in shell fragments and is known

only from Beaufort, North Carolina.

Polydora cauUeryi Mesnil

8

Polydora cauUeryi Mesnil, 1897. pp. 88-89, pi. 3:figs. 12-14.—

Southern, 1914, p. 104.—Mclntoah, 1915, pp. 210-212, pi.

100:fig. 8. pi. 106:fig. 5.—Fauvel, 1927, pp. 54-45, fig.

19/-/I.—Hartman, 1951, pp. 83-84; 1965, pp. 149-150. pi.

28:figs. b-d.—Pettibone, 1954, pp. 282-284, fig. 32*^.—

Hannerz. 1956, pp. 123-126, figi. 43-44.—Eliason, 1962, p.

53.—Blake. 1969b. p. 52. fig. 38.

Polydora carazxi Mclntoth, 1909, pp. 172-173, pi. 5: figs. 4-6.

Polydora brachycephala Hartman, 1936, pp. 48-49, figs. 3-5;

1941, p. 307; 1944, p. 238; 1954, p. 10; 1961, p. 29; 1969, figs.

1-3.—Filfce. 1958, p. 190.—Reish, 1959. pp. 38, 61, 67. 70,

75; 1964b, pp. 86-92.

MATERIAL EXAMINED.—Maine (Damariscotta

River), Connecticut (Noank), Massachusetts (Cape

Cod Bay, coll. SEP), California (Dillon Beach, coll.

K. H. Woodwick), Sweden (Kattegatt, NMGS).

DESCRIPTION.—Largest specimens measure 50 mm

in length, are 2.2 mm wide, and have over 150 segments. Rust colored pigmentation has been observed

dorsally on the anterior segments of some sped*

mens. The color in life is light tan with prominent

red blood vessels. A female which had recently

spawned was yellow.

The anterior end is prominently wedge-shaped

(Figure 8a). The prostomium is deeply incised

anteriorly with two rounded lobes. The caruncle

extends back through setigers 3 or 4. Eyes are present or absent; if present, a small oval pair lies posterior to a more widely spaced cup-shaped pair.

Palps are slender.

Setigers 1-4 have flattened, bladelike parapodial

lobes, the notopodial lobes being somewhat auriculate (Figure 8b, h, t). The notosetae of setiger 1

are short, slender capillaries (Figure Sh) while

those of setigers 2-4 are long, slender, winged

capillaries (Figure 8t). Setiger 6 and succeeding

segments contain notopodial fascicles of long,

winged capillary setae. In far posterior segments,

the capillary notosetae are replaced by a curved row

of stout, sharply pointed spines (Figure 8g). Neurosetae of setigers 1-4 and 6 are long, slender capillaries. Bidentate hooded hooks begin on setiger 7,

accompanied by a tuft of capillary setae. The hooks

are recurved, with a main fang at nearly right angles

to the shaft and have a smaller secondary tooth

(Figure 8/). There are about 13 hooks on anterior

setigers, the number gradually diminishing until in

far posterior segments only about 5 can be found in

a parapodium, accompanied by one or two capillary

setae.

Setiger 5 is large, with a heavy musculature,

overlapping setiger 6. The characteristic large,

bushy-topped modified spines are arranged in an
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICUBE S.—Pofydora caulleryi: a, Anterior end in dorsal view; b, anterior end, lateral view; c,

heavy modified spine and companion capillary seta of setiger 5; d, heavy spine of setiger 5; e,

dorsal seta of setiger 5; /, hooded hook; g, posterior notopodial spine; h, parapodium of setiger 1,

anterior view; i, parapodium of setiger 3, anterior view; /, pygidium, posterior view.
NUMBER 75 IS

anteriorly elevated semicircle, accompanied by a

few very fine, capillary companion setae (Figure

8c, d). A vertical row of dorsal, winged setae is

located anterior to and above the heavy spines

(Figure 8e). The neurosetae consist of a tuft of

curved capillary setae.

Branchiae begin on setiger 7, being small at first

and reaching full size on setiger 11. They are absent

from the last third of the body.

The pygidium consists of four equal lobes (Figure 8;).

REMARKS.—The synonymy of Polydora caulleryi

and P. brachycephala, proposed by Pettibone

(1954), is herein supported. California specimens

possess posterior spines and agree in other respects

with New England representatives. Specimens from

the Kattegat (Sweden) did not differ from American material. Pelagic larvae obtained in New England waters agree well with those described by

Hannerz (1956) from Sweden (Blake, 1969b).

DISTRIBUTION AND ECOLOGY.—Polydora caulleryi

occurs in Europe, New England, Alaska and California. It ranges from low intertidal (Pettibone,

1954; personal observations) to continental shelf

depths of 200 meters (Hartman, 1965).

Polydora quadrilobata Jacobi

FlCUKE 9

Polydora quadrilobata Jacobi, 1883, pp. 1-37, 2 pis.—Mesnil,

1897, pp. 87-88, pi. 3: figs. 9-11.—Mclntosh, 1909, pp.

170-172. pi. 5:figs. 2-3; 1915. p. 236, pi. 98:figs. 13, 17. pj.

106: fig. 4.—Eliason, 1920, pp. 45-46; 1962, p. 52.—Fauvel,

1927, p. 52, figs. 18 /-n.—Annenkova. 1931, pp. 203-205;

1932, p. 139, figs. 6-9.—Procter, 1933. p. 142—Friedrich,

1938, p. 133.—Berkeley and Berkeley. 1943. p. 130; 1954.

p. 464.—Uschakov, 1955. p. 278. figs. 94a-d; 1965, p. 250*

fig. 9ia-d— Rasmussen, 1956, p. 64.—Hannerz, 1956, pp.

122-123.—Hempel, 1957a, pp. 276-278; 1957b, pp. 100-135,

fig. 5.—Hartman, 1961, p. 29; 1969, pp. 145-146, 4 figs.—

Khlebovitsch, 1961, p. 201.—Jones, 1961, p. 288.—Blake,

1969b, pp. 37-51, figs. 27-37.

Polydora tubifex Verrill, 1885b, pp. 438-439.—Sumner et al.,

1913, p. 624.

Polydora littorea.—Hartman, 1944, pi. 18: fig. 9. Not Verrill.

1881.

MATERIAL EXAMINED.—Maine (Cobscook Bay;

Lamoine Beach; Damariscotta River), Connecticut

(Noank), Sweden and Denmark (NMGS).

DESCRIPTION.—Individuals measure up to 15 mm

in length and have about 70 segments. The anterior

end has brown, reticulated pigmentation dispersed

on the sides of the peristomium and on the anterior

and posterior margins of setigers 1-7. The intensity

of the pigmentation varies between individuals,

but has been found in some degree on all specimens

examined from New England. The color in life is

light tan,with red blood prominent in the branchiae, palps, and blood vessels.

The prostomium is distinctly bifid on its anterior

margin and continues posteriorly as an indistinct

caruncle to about the middle of setiger 3 (Figure

9a). Four to six eyes are present in nearly a straight

transverse line. The two outer pairs may be fused

into a single mass. The palps are long and flexible,

with a prominent ciliated groove.

Setiger 1 has capillary notosetae and neurosetae

(Figure 96). Setigers 2-4 and 6 have neuropodial

fascicles of finely-winged, capillary setae. The

notosetae of setigers 2-4 are winged capillaries

(Figure 9c). Bidentate hooded hooks begin on

setiger 7 (Figure 9i) and are accompanied by two

or three fine capillary setae. There are usually no

more than three or four hooks in any one parapodium. The notosetae of setiger 6 and succeeding

setigers include fascicles of winged capillaries. The

number of capillary notosetae diminishes in the

middle segments, and they are completely replaced

by heavier, sharply pointed spines in far posterior

segments. These spines are disposed in a semicircular arrangement and give the posterior end a

spinous appearance (Figure 9j-k).

Setiger 5 lacks parapodial lobes and has three

types of setae (Figure 9d) . The dorsal setae consist

of a horizontally curved row of heavy, modified

spines (Figure 9e, f) and a vertical curved row of

shorter limbate setae (Figure 9g, h) located at the

anterior end of the horizontal row, resulting in a

J-shaped arrangement when the specimen is viewed

from the sides. The heavy spines have a unique

structure (Figure 9e, f). The distal end curves and

is bifurcated at the tip, forming two nearly equal

teeth between which is located a fine bushy tuft.

When worn, the teeth and tufts may be reduced. A

small fascicle of winged setae is located ventral to

the modified spines.

Branchiae begin on segment 7. They are short

and stubby, not reaching full size until setiger 12 or

13. They are absent from the posterior third of

the body.

The pygidium has four subequal lobes (Figure
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FicwtE 9—Potydora quadrilobata: «, Anterior end, donal view; b, setigrr 1, anterior view; c,

•etiger 3, anterior view; d, anterior end, lateral view; e, f, heavy ipincs of tedger 5; g, h, limbate

•etae of wtiger 5; i, hooded hook; /, posterior notopodial spinet; k, posterior end, donal view.
NUMBER 75 15

9k). When seen on living specimens under reflected

light, they are white.

REMARKS.—Soderstrom (1920) considered P.

quadrilobata to be a juvenile of P. caulleryi. This

synonymy has not been supported by subsequent

workers. There is no question, however, that the

two species are closely related. A comparison between European and American specimens of P.

quadrilobata revealed no morphological differences.

Polydora tubifex Verrill (1885b) is here referred

to P. quadrilobata. The original description was

incomplete and no setal details were given. The

description as given by Verrill, would apply to

several species. The eyes were said to be nearly in

a straight line; the "caudal appendage" was small

and consisted of four short lobes; and the anterior

segments had "greenish-black" marks between segments. Polydora quadrilobata is the only spedes uncovered to date in New England to which this

combination of characteristics agrees. J. Percy

Moore (MS) described and figured P. tubifex. His

description, however, differs markedly from that

given' by Verrill. Moore's spedmen in the Academy

of Natural Sdences of Philadelphia was examined.

Consisting of a single specimen in two parts, it

agreed with Moore's description but dearly belonged to Polydora commensalis.

ECOLOGY.—Polydora quadrilobata is a common

spedes in New England. It is easily collected intertidally at Lamoine Beach and Cobscook Bay State

Parks (Maine) where it occurs in sandy mud. At

Noank, Connecticut, the spedes is abundant off the

pier at the Marine Research Laboratory of the

University of Connecticut, where it occurs in silty

mud. The tubes stand erect in the substrate, are

rusty colored, and are formed of fine silt. The

spedes is usually associated with other spionids,

maldanids, and sabellids. It was dredged in 10-12

meters in Clark Cove on the Damariscotta River,

Maine, where it occurs in muddy substrate and is

associated with Polydora socialis and a sabellid,

Laonome sp.

DISTRIBUTION.—Europe, eastern Canada, New

England, southern California, western Pacific.

Polydora colonia Moore

FIGURE 10

Polydora colonia Moore, 1907. pp. 199-201, pi. 15: figs. 18-23.

—Sumner, et al., 1915, p. 625.—Annenkova, 1938, p. 178.—

Hartman, 1945. pp. 32-33.—Uschakov, 1955. p. 273, fig.

94g-;; 1965. p. 252. fig. 94g-;.

Polydora ancistrata Jones, 1962, pp. 185-187, figs. 55-65.

Polydora hoplura inhaca Day, 1957, p. 99, fig. 6ft-/; 1967. p.

468, fig. 18.2n.

MATERIAL EXAMINED.—Massachusetts (Holotype,

ANSP 2S24, Woods Hole; Hadley Harbor), North

Carolina (Beaufort, AHF 1546), Jamaica (paratypes of P. ancistrata, in the collections of Keith H.

Woodwick).

DESCRIPTION.—Polydora colonia is a small spedes,

measuring up to 5 mm in length and having about

35 segments. The prostomium appears anteriorly

rounded in preserved specimens (Figure 10a). In

life it is seen to have a slight anterior bifurcation,

which is turned under when preserved. The caruncle extends to setiger 2. The palps are short and

reach posteriorly to setigers 8 or 9. Eyes and other

body pigmentation are lacking (Figure 10a).

Setiger 1 contains only capillary neurosetae. The

neurosetae of setigers 2-4 and 6 are short, winged

capillaries. Hooded hooks begin on setiger 7. They

are bidentate with a large fang and small secondary

terminal tooth; the shaft has a constriction (Figure

lOg). The notosetal fasddes of setigers 2-4, 6 and

succeeding segments contain long capillary setae.

The notosetae diminish in number in posterior

segments and, in the last six or seven segments,

are accompanied by one or two specialized hooks

(Figure 10/). The more anterior hooks are slender

(Figure \Qh, t) and not as curved as those which

follow (Figure 10;, k). The latter are recurved and

falcate.

Setiger 5 is larger than the preceding and succeeding setigers. The modified spines are bifid,

with two unequal teeth and a subterminal collar

extending halfway around the spine (Figure lOd-f).

Alternating with the three or four large spines are

pennoned companion setae (Figure 10c). There is

a small bundle of slender, winged, dorsal setae located above the modified spines (Figure 106). A

ventral tuft of three or four winged capillary setae

is also present.

Branchiae begin on setiger 7, continuing through

the middle segments; they are absent from the posterior one-half of the body. They are small and do

not overlap dorsally (Figure 10a).

The pygidium consists of a flattened disk, the

sides of which gradually merge into the last segment, leaving a wide dorsal gap (Figure 10/).
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGUK 10.—Potydora coIonia: a, Anterior end, dorsal view; b, superior dorsal seta of setiger 5;

e, pennoned companion seta of setiger 5; d-f, heavy spines of setiger 5 in different views, g,

hooded hook; h-k, posterior notopodial hooks representing a sequential change in structure on

progressively more posterior segmenu; /, posterior end, dorsal view.
NUMBER 75 17

REMARKS.—Polydora ancistrata Jones is herein

referred to P. colonia Moore. The differences between the two species, as discussed by Jones (1962),

have not been substantiated by recent, wellpreserved material collected from the type locality

of P. colonia (Woods Hole, Massachusetts). Polydora hoplwra inhaca Day (1957) also belongs in the

synonymy, as suggested in a footnote by Day (1967).

The species most closely related to P. colonia is

P. spongicola Berkeley (emend. Woodwick, 1963).

The specimens of the latter from California and

MazatlAn, Mexico (courtesy of Dr. Keith H. Woodwick) were examined by the writer. In P. spongicola

the specialized spines of setiger 5 have a large terminal tooth surrounded by a flange, while in P.

colonia there are two teeth surrounded by a collar.

ECOLOGY.—Polydora colonia inhabits tubes

within sponges, forming compact, soft masses attached to pilings and rocks in areas of water movement.

DISTRIBUTION.—Massachusetts to North Carolina,

Jamaica, South Africa, northwest Pacific.

Polydora commensalis Andrews

FlCUftE 11

Polydora commensalis Andrews, 1891a, pp. 25-35, 2 pis.—

1891b, pp. 291-292, pi. 15: fig. 27.—Cowles, 1930, p. 344.—

Berkeley and Berkeley, 1936, pp. 469-471; 1952, pp. 18-19,

figs. 29-30.—Annenkova, 1938, p. 178, fig. 14—Hartman,

1941, p. 308; 1945, p. 32; 1961, p. 29; 1969, pp. 133-134,

4 figs.—Rioja, 1943, p. 229.—Hartman and Reish, 1950, p.

28.—Uschakov, 1955. p. 272, fig. 93; 1965, p. 250, fig. 93 —

Wells and Gray, 1964, p. 73.—Hatfield, 1965, pp. 356-368,

figs. 1-5.—C. Berkeley, 1968, p. 560.—Blake. 1969a, pp.

815-816, fig. 3; 1969b, pp. 21-24. figs. 16-18.

Polydora sp. Berkeley, 1927, p." 420.

Polydora ciliata brevipalpa Zaks, 1933, p. 129.

MATERIAL EXAMINED.—Maine (Damariscotta

River), Massachusetts (Cape Cod Bay, coll. SEP;

North Falmouth, as P. tubifex coll. J. P. Moore,

ANSP 1516), Connecticut (Noank), North Carolina (Beaufort).

DESCRIPTION.—Specimens measure up to 30 mm

in length, are 2.5 mm in width, and have over 100

segments. The prostomium is rounded or weakly

emarginate on its anterior margin; the entire head

is usually contracted into the first setiger (Figure

lla). There is no caruncle. Four small eyes are

arranged in a nearly straight line. The palps are

unusually short and covered with numerous, fine

papillae, each with a sensory hair. The color in life

is dark tan with a greenish cast on numerous segments. Red blood vessels are prominent.

Setigers 1-4 have well developed noto- and neuropodial lobes with fascicles of long capillary setae

(Figure 1 la) . Se tigers 3 and 4 have some additional

short capillary notosetae. Bidentate hooded hooks

begin on setigers 12-17 (rarely 10-17). The hooks

are accompanied by a few capillary neurosetae. The

hooks have a main fang surmounted by a shorter

secondary tooth (Figure lid). The secondary tooth

is reduced in hooks from far posterior setigers and,

when viewed from certain angles, is difficult to see

(Figure lie). There are 5 or 6 hooks in anterior

setigers, increasing to about 13 in middle setigers,

and reduced to 4 or 5 posteriorly.

The setae of setiger 5 are modified to include a

row of large curved spines and a small tuft of capillary neurosetae. The heavy spines have a characteristic long lateral flange or sheath (Figure lib, c).

Branchiae are long and straplike, with membranous margins. They begin on setiger 6 and continue to the end of the body.

The posterior end of the animal is dorsoventrally

flattened (Figure I If). The anus is surrounded by

a ring of small papillae, the number of which is

variable, depending upon the age of the animal- In

young juveniles 4 papillae may be present, increasing to 14 in the largest adults. An intermediate

stage with 8 papillae is shown in Figure 11/.

REMARKS.—Polydora commensalis deviates widely

from other spedes of the genus and should be removed from Polydora. However, the writer has

knowledge of two undescribed species in California

which appear to be closely related to P. commensalis. Until these new collections are studied in detail, the generic placement of P. commensalis had

best be delayed.

The specimens, identified as P. tubifex Verrill

and described by Moore (MS), were examined and

found to be P. commensalis (ANSP 1516). Polydora

tubifex Verrill (1885b) is here (p. 13) referred

to P. quadrilobata.

ECOLOGY.—Polydora commensalis is known only

from shells containing hermit crabs. In Maine, the

mollusc shells include Littorina littorea and Thais

lapillus, occupied by the hermit crab Pagurus

longicarpus. Shells covered with the hydroid

Hydractinia echinata appear to have a higher infestation of P. commensalis.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ficuu 11.—Poly dora commetualis: a. Anterior end, donal view; b, c, heavy spines of se tiger 5;

d, e, hooded hooks; /, posterior end, donal view.
NUMBER 75 19

Fsscu 12.—Polydora concherum: a, Anterior end, dorsal view; b-d, heavy ipinet of ledger 5 In

different view*; e, pennoned companion seta of ledger 5; /, superior dorsal geniculate sett of

•etiger 5; g, setiger 1, anterior view; h, posterior setiger, dorsal view (not to scale); i, posterior

end, dorsal view; /, pygidium, posterior view; *, /, hooded books; m, n, winged capillary

notosetae.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—East and west coasts of North

America; North Japan Sea.

Polydora conchamm Verrill

Ficinti 12

Polydora concharum Verrill, 1880, pp. 174-176.—Webster and

Benedict. 1884, p. 729; 1887, p. 737.—Sumner, et al., 1913,

p. 624.—Procter, 1933, p. 142—Hartman, 1942, pp. 64-66,

figs. 121-123; 1944, pp. 336-340, pi. 19: fig. 10 (not fig. 8).—

Evans, 1969, pp. 775-782.—Blake. 1969a, p. 816, fig. 4;

1969b, pp. 32-36, figs. 23-26.

Dipolydora concharum.—Verrill, 1881, p. 320; 1885a, p. 688,

pi. 43: fig. 186.

MATERIAL EXAMINED.—West Greenland (Hero

Station 29F) , Newfoundland (St. Mary's Bay, 10-20

meters, coll. J. W. Evans), Maine (Damariscotta

River; Gulf of Maine, Types YPMNH 2687), Massachusetts (Cape Cod Bay, coll. SEP).

DESCRIPTION.—Polydora concharum is a long,

slender spedes reaching lengths up to 140 mm and

having over 300 segments. In life it is light tan with

red branchiae and blood vessels. Juveniles have

been found to retain larval pigment for a time after

metamorphosis. In adults the body is largely colorless, although some specimens have been found

with a line of dark pigment on the anterior borders

of some segments, first appearing in setiger 6.

The prostomium is distinctly bifid on the anterior margin. The caruncle continues posteriorly

as a low ridge to the anterior border of setiger 5

(Figure 12a). Eyes are either present or absent;

when present, there are four in a trapezoidal arrangement. The palps are relatively short in comparison to the length of the animal and extend

posteriorly for only 15 to 20 segments.

Setiger 1 has both noto- and neurosetae (Figure

12g). The former are fine capillaries, while the

latter consists of winged capillaries. Setigers 2-4

have well developed postsetal lobes and large

spreading fascicles of winged capillary noto- and

neurosetae. Notosetae of setiger 2 are arranged in

two successive rows, while those of setigers 3, 4, 6,

and those immediately following have notosetae in

three rows, those of each successive row being longer

(Figure 12m, n). Notosetae from more posterior

setigers lack wings; they are represented by three

sizes of straight capillary setae (Figure 12h). The

winged neurosetae of setigers 2-4 and 6 are arranged in two rows. Bidentate hooded hooks (Figure 12A, /) begin on setiger 7 and are accompanied

by two or three capillary setae.

Setiger 5 is larger than proceeding and succeeding

setigers. The setae are arranged in three bundles:

a group of geniculate setae (Figure 12/) located

anterior and dorsal to a semicircular row of large

heavy spines (Figure \2b-d), alternating with

smaller pennoned companion setae (Figure 12e);

and a small tuft of ventral winged neurosetae. The

heavy spines have a weakly-developed accessory

shelf, horizontal to the main shaft. The structure

was not observed by Verrill (1880) or Hartman

(1942).

Branchiae begin on setigers 7-9. They are small

anteriorly, reaching full size on setiger 11 or 12 and

are absent from the posterior half of the body.

The pygidium is small and has four lobes, the

dorsal pair being smaller than the ventral pair

(Figure 12i,;).

ECOLOGY AND DISTRIBUTION.—Cape Cod, Massachusetts, to Newfoundland, in dredged shells and

decayed wood; West Greenland.

Polydora sociaUs (Schmarda)

FIGURES 13, 14

Leucodore socialis Schmarda, 1861, p. 64, figs, a-c, pi. 26: fig.

209.

Polydora socialis.—Mesnil, 1896, pp. 193-194, pi. 12: figs.

30-32—Ehlers, 1901, p. 165.—Hartman, 1941, pp. 310-311,

pi. 48: figs. 41-42; 1945, pp. 33-35; 1948, p. 37; 1951. p. 83;

1953. pp. 43-44; 1961, p. 29; 1966a, p. 19. pi. 19:figs. 1-2;

1969, pp. 147-148. 2 figs.—Hartman and Reish, 1950, p. 28.

—Rioja, 1943, p. 230.—Hartmann-Schrdder, 1962, pp.

137-138, figs. 167-168; 1965. pp. 209-211, figs. 200-203 —

Wells and Gray, 1964, p. 73.—Long, 1968. p. 348.—Blake,

1969a. pp. 816-817. fig. 5; 1969b. pp. 24-31. figs. 19-22.

Polydora socialis plena Berkeley and Berkeley, 1936, pp.

468-469; 1952, p. 22—Reish, 1968, p. 82.

?Polydora gracilis Verrill, 1880, p. 174.

MATERIAL EXAMINED.—Maine (Damariscotta

River), Massachusetts (Cape Cod Bay, coll. SEP),

Connecticut (Mystic River), North Carolina (Beaufort) , South Carolina (Charleston, coll. M. Chamberlain), California (Mugu Lagoon, AHF N1417;

Cayucos), British Columbia (paratypes of P. socialis

plena, USNM 32704), Chile (Lund University

Chile Expedition 1947-48, deposited AHF).

DESCRIPTION.—Specimens of Polydora socialis

from New England measure up to 55 mm in length

and have over 400 segments. Individuals examined
NUMBER 75

21

Ficuu IS.—Polydora sodalis: a, Anterior end, dorsal view; b, diagram of ledgers 4-7 in lateral

view (not to scale); c, pygidium, posterior view; d, posterior end, dorsal view.

from North and South Carolina were not as large.

Sexually mature females are orange. Larval pigmentation is retained on most adults (Figure 13a),

including both dorsal and ventral bands.

The prostomium is deeply notched on its anterior

margin. The caruncle extends to setigers 4-9. Eyes

are present or absent; when present, they number

four to six: a circular pair high up on the prostomial ridge, and more anterior second and third

pairs, which may be fused laterally on the prostomial ridge (Figure 13a). The palps are long and

prehensile in life; upon preservation, however, they

contract and extend posteriorly only to setigers 12-

15.

Setiger 1 has capillary setae in both noto- and

neuropodia (Figure 14a). Setigers 2-4, 6 and succeeding setigers have two types of posteriorly directed notosetae, an anterior row of short curved

capillaries and a posterior row of long, stout capillaries. In far posterior setigers only a few laterallydirected capillary notosetae are present The

neuropodia of setigers 2-4 and 6 have posteriorly

directed capillary setae. Bidentate hooded hooks

begin on setiger 7. Two or three capillary setae ac-
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Fwtnut 14.—Poly dor a socialis: a, Se tiger 1, anterior view; b, fascicle of modified spines and

pennooed setae from aetiger 5; e, hooded hook from anterior setiger; d, hooded hook from

median setiger; e, f, hooded hooks from posterior sedgers; g, gisard-like structure from an adult

(not to scale).
NUMBER 75 23

company the hooks for several setigers, then disappear, and appear again in far posterior setigers. In

anterior setigers, the hooks have a 45° angle between

the main fang and the secondary tooth (Figure

14c). In posterior setigers, the main fang is longer

and narrower, while the upper tooth is smaller and

closely adhering to the shaft and diminishing the

angle (Figure 14e).

Setiger 5 is large and well developed (Figure

15a, b). The setae include a small bundle of dorsal

geniculate setae lying at the anterior end of a

semicircular row of alternating heavy modified

spines and smaller pennoned companion setae (Figure 14b). The heavy spines are falcate and have a

subterminal protuberance. Ventral and posterior

to the above is a small tuft of winged setae.

Branchiae begin on setiger 8 (rarely 7-9) and

continue to near the posterior end.

The pygidium typically has a large ventral lobe

and two small dorsal ones, with a dorsal gap (Figure 15c, d). In some specimens, however, the three

lobes are continuous.

In P. socialis an unusual grinding apparatus has

been observed between the esophagus and intestine.

In young specimens, this "gizzard-like" structure is

thin and transparent, consisting of four teeth, each

with a central, inwardly directed cusp. In older

specimens, the teeth are fragmented and lie as inclusions in four longitudinal muscles (Figure 14g).

This structure is similar to one figured by Carazzi

(1893) for P. flaw.

REMARKS.—The type material of P. socialis

(Schmarda) from Chile (Vina del Mar) was redescribed by Mesnil (1896). Hartman (1941, 1948,

1961), Rioja (1943, 1947), and Hartman and

Reish (1950) have extended the species range from

Mexico to Oregon and Alaska. Hartman (1953,

1966a) has reported the species from the Falkland

Islands. Records from eastern North America include Hartman (1945) and Wells and Gray (1964)

from the Carolinas and Hartman (1951) from Florida. In each of these reports, the specimens lacked

notosetae on setiger 1.

A new subspecies, P. socialis plena, was described

by Berkeley and Berkeley (1936, 1952) from British Columbia. Reish (1968) reported P. socialis

plena from Baja California. This subspecies differed

in having notosetae on setiger 1. With the record

of Reish, P. socialis plena overlapped the range of

P. socialis (sensu stricto), a fact which seemed unusual to this investigator.

Early in the present study it became apparent

that all specimens of this species from the east coast

of North America, from Maine to South Carolina,

possessed notosetae on setiger 1, thus agreeing with

the subspecies plena. Since the notosetae are small,

it seemed possible that they had been overlooked in

earlier east and west coast reports. During a visit

to California, it was possible to examine specimens

of P. socialis collected by Dr. Keith H. Woodwick

from various California localities. All possessed

notosetae on setiger 1. Collections in the Allan

Hancock Foundation were also examined. The

specimens which Hartman (1941) had reported

from Mugu Lagoon, southern California, were

found to have notosetae on setiger 1, as were specimens identified as P. socialis from the Lund

University Chile Expedition (unpublished), now deposited in the Allan Hancock Foundation. Further,

specimens of P. socialis from localities in Chile described by Hartmann-Schrdder (1962, 1965) also

possessed notosetae on setiger 1. The presence of

notosetae on setiger 1 on all specimens from Chile

to British Columbia, and from South Carolina to

Maine eliminates the need for the subspecies plena.

The "gizzard-like" structure described above was

found on all specimens examined and the structure

of the modified spines of setiger 5 was found to be

the same.

Specimens, described as Polydora gracilis Verrill

by J. P. Moore (MS), were examined in the Philadelphia Academy of Natural Sciences. They agreed

with P. socialis. Blake (1969a) placed P. gracilis in

synonymy with P. socialis but this is at least questionable, since no type material of P. gracilis exists

and the description is not adequate for specific determination.

ECOLOGY.—Polydora socialis constructs tubes in

sediment and may form extensive beds (Hartman,

1941). In the Damariscotta River, Maine, P. socialis

is found subtidally in sediment and to a lesser extent in shell. In the former, it forms tubes of silt

with a firm mucoid lining, and is associated with

P. quadrilobata and Laonome sp. In the latter it

bores into shells of dead Mercenaria and living

Placopecten magellanicus associated with P. concharum and P. websteri.

DISTRIBUTION.—East and west coasts of North

America; Gulf of Mexico; Chile; Falkland Islands.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FKUBE 15.—Potydora anoculata: a. Anterior end, dorsal view; b-i, heavy modified spine* ef tetiger

5 seen in different views and degrees of wear; /, hooded hook.
NUMBER 75 25

Polydora anoculata Moore

FICUBE 15

Polydora anoculata Moore 1907, pp. 197-199, pi. 15:fig». 7-17,

25.—Sumner et al., 191S, p. 625.

MATERIAL EXAMINED.—Massachusetts (Cape Cod

Bay, coll. SEP; Holotype, ANSP 2338).

DESCRIPTION.—Polydora anoculata is a thin, slender species measuring up to 20 mm in length and

having about 100 segments. The prostomium is

anteriorly bifurcated. Eyes are absent. The caruncle

extends posteriorly to setiger 5 (Figure 15a).

Setiger 1 has both noto- and neurosetae. The

former are thin, slender capillaries, while the latter

have narrow wings. Setigers 2-4, 6 and succeeding

setigers contain winged capillary notosetae. The

slender capillary neurosetae of setigers 2-4, and 6

are almost completely replaced by hooded hooks on

setiger 7. The hooks have a recurved shaft and no

constriction (Figure 15/).

Setiger 5 contains three groups of setae: a bundle

of pennoned dorsal setae anterior to a row of heavy

spines alternating with pennoned companion setae,

and a ventral tuft of delicate pennoned setae. The

heavy spines have a lateral flange, which resembles

a tooth when worn (Figure 15b-i). An additional

small spur is occasionally seen on some spines.

Branchiae begin on setigers 9 or 10 (Figure 15a).

They are small at first, attaining full size on setiger

11; they are absent from posterior setigers.

The pygidium is disklike, with a deep dorsal

notch.

REMARKS.—Polydora anoculata resembles -P.

giardi Mesnil and P. trideniiculata Woodwick in

branchial arrangement. It differs from the latter in

the structure of the modified spines of setiger 5

(Woodwick, 1964). The differences between P.

giardi and P. anoculata, however, have not been

entirely resolved in the present study. Specimens

labeled P. giardi were obtained from Europe, but

they proved to be another species. It may be that

the two species are conspecific. Additional material

is needed for comparison.

ECOLOGY.—Moore (1907) reported P. anoculata

associated with colonies of Amaroecium pellucidum, on piles among Cynthia, and in sponge together with P. colonia. In Cape Cod Bay it was

found on a shell-gravel bottom.

DISTRIBUTION.—Known only from the Woods

Hole region, Massachusetts.

Polydora hartmanae, new species

FIGURE 16

Polydora anoculata—Hartman 1945, p. SS. Not Moore 1907.

MATERIAL EXAMINED.—The specimens from

Bogue Sound, near Beaufort, North Carolina, described by Hartman (1945) as P. anoculata, were

reexamined. They proved to be an undescribed

species. The holotype and one paratype are deposited in the Allan Hancock Foundation.

DESCRIPTION.—The types consist of two anterior

ends and two posterior fragments, with the pygidium missing. One anterior end with 78 segments

measured 10 mm in length.

The prostomium is distinctly bifid in its anterior

margin, extending posteriorly as a low caruncle to

setigers 4 or 5. Eyespots and nuchal tentacle are

lacking. The peristomium is greatly enlarged on

the lateral and anterior margins (Figure 16a).

Setiger 1 has fascicles of capillary setae in both

the noto- and neuropodia. The neurosetae are finely

unilimbate. Setigers 2-4 contain spreading fascicles

of winged, capillary noto- and neurosetae. The

notopodia of setiger 6 and succeeding setigers contain double fascicles of long and short, winged capillary setae, the number gradually diminishing

posteriorly. No specialized posterior notosetae were

observed. Bidentate hooded hooks begin on setiger

7, with 5 or 6 per neuropodium; they have a recurved shaft. The teeth are delicate and have a reduced angle between them. (Figure 161).

Setiger 5 contains three groups of setae: a cluster

of 4 or 5 large, geniculate setae (Figure 16A), located anterior and dorsal to a semicircular row of

heavy spines (Figure 16b-h) alternating with pennoned companion setae (Figure 16*,;), and a small

tuft of winged neurosetae below the large spines.

The heavy spines are distally falcate, with a lateral

flange, the latter being broken or worn on all except newly-formed spines. Tufts of fine hairs arise

from the concavity separating the flange from the

main fang and, to a lesser degree, on the exterior

of the flange (Figure 166-g).

Branchiae begin on setigers 11 and 12, being

short and stubby anteriorly and reaching full size

on about segment 16. They are absent from posterior segments.

The pygidial structure is unknown.

REMARKS.—Polydora hartmanae differs from
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Some taxonomic characteristics of

Species of

POLYDORA

aggregate

anoculata

caulleryi

colonia

commensalis

concharum

hartmanae

ligni

quadrilobaU

sodalit

tetrabranchia

websteri

branchiae:

a. begin on

setiger

b. on posterior

segments

a. 7

b. absent on

posterior

half

a. 9-10

b. abtent on

posterior

half

a.7

b. absent on

posterior

third

a.7

b. absent on

posterior

half

a. 6

b. present

a. 7-9

b. absent on

posterior

half

a. 11-12

b. absent

a.7

b. present

a.7

b. absent on

posterior

third

a. 8 (rarely

7-9)

b. absent on

last segments

a.7

b. absent from

setiger 11

a.7

b. absent

Notosetae

on first

setiger

absent

present

present

absent

present

present

present

absent

present

present

absent

absent

Modified

posterior

notopodial

spines or

hooks

absent

absent

awl-shaped

spines

falcate

recurved

hooka

absent

absent

absent

absent

awl-shaped

spines

absent

absent

absent

the species

Modified

spines

of 5th

setiger

falcate.

with

accessory

knob

falcate.

with

accessory

flange

falcate.

with

bushy

top

bifid tip.

with subterminal

collar

falcate.

with

long

accessory

sheath

falcate.

with

lateral

sheath

falcate,

with

lateral

flange and

fine hairs

falcate,

with small

accessory

tooth

bifid tip,

with fine

hairs between teeth

falcate.

with subterminal

boss

bifid tip,

with 2

large teeth

falcate, with

lateral

flange or

sheath

of Polydora from eastern North America

Eyes

4

0

0-4

0

4

0-4

0

4

4-6

0-6

4

4

Hooded hooks:

a. begin on

setiger

b. have constriction on shaft

a.7

b. present

a.7

b. absent

a.7

b. absent

a.7

b. present

a. 10-17

b. absent

a.7

b. absent

a.7

b. absent

a.7

b. present

a.7

b. absent

a.7

b. absent

a.7

b. absent

a.7

b. present

Pygidium

disk-like

disk-like

4 equal

lobes

disk-like

4-14

papillae

4 lobes,

dorsal pair

smaller rii*n

ventral pair

?

disk-like

4subequal

lobes

3 lobes or

disk-like

collar-like

disk-like

Other

features

pigmented

pattern in

anterior

region

-

-

-

commensal in

gastropod shells

occupied by

hermit crabs;

palps short

comparatively long

species; S sizes of

posterior capillary

notosetae

_

nuchal tentacle

present

pigmented pattern

in anterior body

region

gizzard-like structure present;

larval pigment retained on adults

very small

species

_
NUMBER 75 27

Ficuiut 16.—Potydora hartmanae new species: a, Anterior end, donal view; b-h, heavy modified

spines of setiger 5 seen in different views and degrees of wear; i-j, companion setae of setiger 5;

k, donal geniculate seta of setiger 5; I, hooded hook.
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

other species in the structure of the modified spines

of setiger 5.

DISTRIBUTION AND ECOLOGY.—The species is

known only from North Carolina. It was reported

by Hartman (1945, p. 33) as being ". . . dredged

from a bottom of broken shell fragments." It is

not known whether the species actually bored into

the shells or inhabited sediment.

Summary

Twelve species of Polydora have been identified

from collections obtained from various localities

along the east coast of North America. A key to

the species, descriptions and figures are included.

The adults of P. aggregata are described for the

first time. Polydora hartmanae is new to science.

New synonymies include P. littorea Verrill, referred

to P. ligni Webster, P. tubifex Verrill, referred to

P. quadrilobata Jacobi, and P. ancistrata Jones, referred to P. colonia Moore.

Four species to date have been reported only

from eastern North America: Polydora aggregata

from Maine, P. hartmanae and P. tetrabranchia

from North Carolina and P. anoculata from Massachusetts. Polydora concharum ranges from New

England to Newfoundland and West Greenland.

Each of these species may be expected to have a

wider range when additional collections become

available. The other seven species of Polydora have

a wider distribution.

Five species were found boring into shells or other

calcareous structures: P. commensalis, P. con*

charum, P. socialis, P. tetrabranchia and P. websteri.

Five species were found in sediment: P. aggregata,

P. caulleryi, P. ligni, P. quadrilobata and P. socialis.

The latter species is found in both sediment and

shell. Polydora colonia is found in sponges, while

P. anoculata occurs in association with tunicates

and sponges.

Some taxonomic characteristics of the twelve species are summarized in Table 1.

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32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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