HELMUT E. Z1BR0W

Some Little Known

Genera of Serpulidae

(Annelida: Polychaeta)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 42
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 42

Helmut W. Zibrowius Revie w o f

Some Little Known

Genera of Serpulidae

(Annelida: Polychaeta)

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CITY OF WASHINGTON

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ABSTRACT

Zibrowius, Helmut W. Review of Some Little Known Genera of Serpulidae

(Annelida: Polychaeta). Smithsonian Contributions to Zoology 42:1-00. 197-.

Serpulids which had been classified in the genera Vermilia, Miroserpula, Chitinopoma, Chitinopomoides, Hyalopomatus, Hyalopomatopsis, Cystopomatus, and

Protis are examined and discussed.

The genus Miroserpula should be included in the genus Chitinopoma, as the

type-species of both genera are identical. C. serrula, single known species of the

genus Chitinopoma, is widely distributed in the Arctic and Boreal Atlantic

(America and Eurasia). It is characterized by incubation in special brood chambers

on the tube of adult specimens. As C. groenlandica, it has often been confused

with a quite different species from the Pacific coast of North America. For this

latter species, first described as Hyalopomatopsis occidentalis, the new genus

Pseudochitinopoma has been created. Chitinopomoides wilsoni, an insufficiently

known species from bathyal depths off Antarctica, may have some affinities with

the genus Chitinopoma. Affinities of Chitinopomoides with the genera Spirobranchus and Serpula are excluded.

The bathyal and abssyal forms from the Atlantic (Arctic Basin, Azores, Madeira)

and from the Mediterranean, described under the generic names Hyalopomatus

and Hyalopomatopsis, and the bathyal Cystopomatus from Antarctica, are closely

related. The latter two genera are to be considered as synomyms of the former.

The validity of the specific distinctions of these forms, based on the known

material, is questionable.

The genus Protis, known from the Atlantic and Pacific, includes forms which

in Arctic regions may be found in lesser depths, but which are at least bathyal

when not abyssal at lower latitudes (Azores, Romanche deep, Gulf of Mexico, off

California). The specific distinctions of these forms are questionable. A serpulid

from the infralittoral at Puerto Rico, described as P. torquata, does not belong to

the genus Protis.

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

in the Institution's annual report, Smithsonian Year.
Helmut W. Zjbrowius OI

Some Little Known

Genera of Serpulidae

(Annelida: Polychaeta)

The present work aims at revising as far as possible

several genera of serpulids which are little known

or monospecific and which are mainly represented

by Atlantic forms living in Arctic and Boreal regions or at great depths in lower latitudes. Forms

from the Pacific coast of North America and Antarctica are also included in these considerations. Some

of these forms, which by reason their very different

morphology should have been classified in separate

genera, were placed together, whereas others, despite their great similarities, were classified in separate genera.

Most of the species discussed are known only from

few specimens in the collections of many museums

throughout America and Europe. In the present

study, I have been able to go beyond mere bibliographic research only by the cooperation of many

people who gave me the opportunity to examine the

material of these collections. For the loan of material, I have the pleasure to express here my thanks

to Mrs. G. Hartmann-Schroder, Zoologisches

Staatsinstitut und Zoologisches Museum, Hamburg

(ZSZM); to Messrs. A. Capart, Institut Royal des

Sciences Naturelles, Bruxelles (IRSN); A. G.

Chabaud, Museum National d'Histoire Naturelle,

Paris (MNHN), A. Eliason, Naturhistoriska Museet,

Goteborg (NMG); J. D. George, British Museum,

(Natural History), London (BMNH); J. B. Kirkegaard, Universitetets Zoologiske Museum, Copenhagen (UZMC); E. Kirsteuer, American Museum of

Natural History, New York (AMNH); H. W. Levi,

Helmut W. Zibrowius, Station Marine d'Endoume, Marseille,

France.

Museum of Comparative Zoology, Cambridge, Massachusetts (MCZ); and especially to Dr. Marian

H. Pettibone, United States National Museum

(USNM), Washington, D. C, who also helped by

her suggestions and by correcting the English text.

This study was aided in part by a grant to Dr.

Pettibone.

The following genera and species are covered in

this study (synonyms in parentheses):

Chitinopoma Levinsen, emended

(=Miroserpula Dons)

(=Microserpula of Hartman)

C. serrula (Stimpson), new combination

(=C. fabricii Levinsen)

(=Miroserpula inflata Dons)

C. groenlandica (Morch), indeterminable

Pseudochitinopoma new genus

P. occidentalis (Bush), new combination

(=Chitinopoma greenlandica of Pixell)

(=Chitinopoma groenlandica of

Berkeley and Berkeley)

Chitinopomoides Benham

C. wilsoni Benham

Hyalopomatus Marenzeller

(=Hyalopomatopsis Saint-Joseph)

(=Cystopomatus Gravier)

H. claparedii Marenzeller

Hyalopomatus sp.

(=Gen. et sp. ? Eliason)

H. marenzelleri Langerhans

Hyalopomatus sp.

(= Protis arctica of Bellan)

H. langerhansi Ehlers

H. macintoshi (Gravier), new combination

( = Cystopomatus macintoshi Gravier)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Protis Ehlers

P. arctica (Hansen)

(=P. simplex Ehlers)

P. pacifica Moore

Pomatostegus stellatus (Abildgaard)

(=Protis torquata Hoagland)

Family SERPULIDAE Savigny

Genus Chitinopoma Levinsen, 1883, emended

Miroserpula Dons, 1931. [Type-species: Miroserpula inflata

Dons, 1931, by monotypy. Referred herein to Chitinopoma

serrula (Stimpson, 1854). Variant spelling: Microserpula.]

Gender: feminine. Type-species: Chitinopoma

fabricii Levinsen, 1883, by monotypy. Referred

herein to Chitinopoma serrula (Stimpson, 1854).

DIAGNOSIS.—Special setae of the first thoracic segment with well-separated distal limbate zones and

proximal denticulate wings. Thoracic sickle-setae

present. Abdominal setae geniculate with a long

triangular blade. Thoracic uncini saw-shaped with

about 10 teeth, the anterior tooth simple, not

bifurcate. Abdominal uncini in all segments raspshaped. Opercular stalk without pinnules or wings.

Thoracic membranes short, not united at the ventral face of the last thoracic segment. Thoracic setigerous segments 7.

REMARKS.—According to this emended definition,

the genus Chitinopoma includes only the widespread North Atlantic Vermilia serrula Stimpson

1854, a tube-incubating species later described by

Dons (1931) as Miroserpula inflata. Referring to

Levinsen's description of C. fabricii, in which no

sort of incubation was mentioned, Dons and followers (Brattstrom, Thorson, Wesenberg-Lund)

considered the breeding form as the type of a different genus, in spite of the nearly identical setae

and uncini described in both forms. As proved by

examination of breeding specimens, there are in

fact no differences between the setae and uncini of

Levinsen's Chitinopoma fabricii and Don's Miroserpula inflata. It has further been found that most

of the material in the collections of the Copenhagen

museum (UZMC), labeled Chitinopoma fabricii (at

least some of these determinations seem to have

been made by Levinsen himself), is identical with

Don's Af. inflata. The existence of a North Atlantic

nonbreeding Chitinopoma species different from

C. serrula (Stimpson, 1954) (=Vermilia serrula,=

Miroserpula inflata) has never been proved.

Chitinopoma serrula (Stimpson, 1854), new

combination

FIGURE 1

Vermilia serrula Stimpson, 1854, pp. 29-30.—Quatrefages, 1865,

p. 561.—Packard, 1863, pp. 403, 418; 1867, p. 292.—Verrill,

1873, p. 439; 1874a, pp. 39, 41, 43; 1874b, p. 412; 1874c, pp.

499, 503, fig. 3, pi. 6: fig. 1; 1874d, pp. 351, 355, 358, 361,

362, pi. 4: fig. 3; 1879, p. 11; 1881, pp. 289, 293, 295, 306,

310, 324.—Hartman, 1944, p. 343, pi. 22 (54): fig. 5; 1948;

p. 50—Treadwell, 1948, p. 69.

Serpula (Vermilia) serrula?—Dawson, 1860, pp. 28-29, fig. 2.

Chitinopoma fabricii Levinsen, 1883, pp. 194-196, 199, 203,

309, pi. 2: fig. 4, 8f, pi. 3: fig. 20; 1887, p. 300.—Fauvel,

1911a, p. 45.—Derjugin, 1915, pp. 357-358, 363.—Augener,

1928, p. 817.—Ditlevsen, 1929, pp. 54, 78, 79.—Dons, 1934a,

pp. 24-27.—Friedrich, 1939, p. 127.—Brattstrom and

Thorson, 1941, p. 21.—Brattstrom, 1945, pp. 1, 20, 21.—

Wesenberg-Lund, 1951, pp. 130, 156, 167; 1953a, pp. 7, 17;

1953b, p. 138.

Chitinopoma greenlandica [sic].—Bush, 1904, pp. 224, 229,

pi. 37: figs. 3, 9; pi. 40: fig. 31.—Nelson-Smith, 1967, p. 37

[in part]. [Not Hydroides norvegica var. groenlandica

Morch, 1863.]

Chitinopoma groenlandica.—Hartman, 1948, pp. 44, 51; 1959,

p. 580. [Not Hydroides norvegica var. groenlandica Morch,

1863.]

Miroserpula inflata Dons, 1931, pp. 3-5; 1934a, pp. 24-27;

1934b, pp. 35-37; 1937, pp. 73-76—Brattstrom and Thorson,

1941, pp. 21-30— Brattstrom, 1945, pp. 1-22.—Thorson,

1946, pp. 135, 136, 141, 145, fig. 77—Wesenberg-Lund, 1950,

p. 61; 1951, pp. 129, 140, 155, 181; 1953a, pp. 6-7; 1953b,

pp. 113, 123, 138, 140, 149, chart 26—Nair, 1962, p. 41.—

Nelson-Smith, 1967, pp. 36, 37, fig. 27.—Boisen Bennike,

1968, pp. 1, 5.

Mikroserpula inflata.—Thorson, 1936, p. 35.

Microserpula inflata.—Hartman, 1948, p. 50.

Microserpula serrula.—Hartman, 1959, p. 582.

MATERIAL EXAMINED.—Bay of Fundy: 1 spec,

Verrill coll., 1872 (USNM 13375). Casco Bay,

Maine; empty tubes, Verrill coll., 1873 (USNM

13291). Gulf of Maine: 1 spec, Packard coll., 1873

(USNM 13322). Eastport, Maine: 1 spec, Packard

coll., 1866 (UZMC). Kara Sea: 2 spec, "Dijmphna"

expedition (UZMC). Greenland: several spec, on

algae, Schmidt coll. (UZMC). Lindenovf jord, Greenland: 1 spec, Bertelsen coll., 23 July 1935 (UZMC).

Angerrok, Upernarvik, Greenland: 1 spec, Salomonsen coll., 29 June 1936 (UZMC). Faroes: 1 spec.

(UZMC). Oersund: empty tubes, Eliason coll.

(NMG). East-Spitzbergen: several spec (ZSZM).
NUMBER 42

FIGURE 1.—Chitinopoma serrula (a, d-j, from specimen from Maine, USNM; b, c, from specimens

from Spitsbergen, ZSZM): a-c, opercula; d, special seta of first thoracic segment; e; thoracic sickleseta; /, abdominal seta; g, h, thoracic uncinus, lateral and edge views; i, j , abdominal undnus,

lateral and edge views.

DESCRIPTION.—Tube white, subtriangular in cross

section, and adherent along its whole length to the

substratum. The lateral faces may be wrinkled

transversely by growth marks. Median keel more

or less denticulate. Tubes of adult specimens may

have one or several pairs of calcareous brood chambers ("ovicells"), one chamber on each side of the

keel. The mouth of the tube may be widened out in

irregular lobes. The length of the animal may attain

20 mm, the operculum diameter 500 /*. About 50

abdominal segments in adult specimens. Collar

large and trilobed. Thoracic membranes short, extending not further than the second setigerous segment. Gill tuft composed of 5 to 7 filaments on each
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

side. No gill membrane. The second filament, on

the right or on the left side, forms the smooth nonpinnate opercular peduncle. Operculum almost

radially symmetrical and covered by a horny distal

plate of brown color. Generally, this horny plate is

more or less concave but variations can sometimes

be observed: plate almost flat, or slightly convex,

or with a more or less large knob or ridge near its

dorsal rim. First thoracic setigerous segment with

two types of setae: capillary setae and special setae

with a distal limbate zone and a proximal wing of a

few well-marked teeth, both zones well separated

by a gap as long as the wing. The other thoracic

segments with limbate setae and capillary ones.

Sickle-shaped setae with a short proximal limbate

zone present in segments 3 to 7. Abdominal setae

geniculate. No long capillary setae in the last segments. Thoracic uncini saw-shaped with about 9

to 12 teeth, the anterior one simple, not bifurcate.

The abdominal notopodial uncini in all segments

rasp-shaped.

Chitinopoma serrula is mainly characterized by

the incubation of the larvae, up to the stage of

metatrochophore, inside paired brood chambers

with calcareous walls which are attached to the

tube on both sides of the median keel.

DISTRIBUTION.—New England, Gulf of Maine, Bay

of Fundy, Gulf of St. Lawrence (Gaspe* and Labrador), Strait of Belle Isle, East and West Greenland,

Iceland, Faroes, Spitsbergen, between Hope Island

and Bear Island, Novaja Zemlya, Isle of Vaigach,

Gulf of Kola, Murman coast, entire coast of Norway, West coast of Sweden, Oersund, North-West of

Jutland. From the tidal zone down to more than

300 m.

The tube of Chitinopoma serrula has been found

attached to various such solid objects as stones,

shells of Mollusca and Brachiopoda, and Bryozoa

with calcareous skeleton, as well as on comparative

soft substratum, such as the mantle of ascidian

species and even on algae. Nair (1962) included C.

serrula (as Miraserpula inflata) in the list of species

collected during a study on fouling organisms made

in the Bergen area. Nevertheless, the species should

not be considered as a main fouling organism.

REMARKS.—The species has a large distribution

throughout the Arctic and Boreal Atlantic Ocean

where it has been described under different names,

chiefly as Vermilia serrula Stimpson on the American coast, as Chitinopoma fabricii Levinsen from

Greenland and Faroe waters, and as Miroserpula

inflata Dons on the coast of Norway. Of the generic

names, Chitinopoma and Miroserpula, created specially for the same species, the former is the older

one. Thus the species should be named Chitinopoma serrula (Stimpson, 1854).

The only publication in which Vermilia serrula

and Miroserpula inflata are considered as identical

(under the name of Microserpula inflata [sic], seems

to be the "Catalogue of the Polychaetous Annelids

of the World" by Hartman (1959), in which, however, no proof for this synonymy is given. Apparently the Scandinavian authors, when treating

of the morphology, breeding, geographical distribution and ecology of Miroserpula inflata (Dons,

Brattstrom, Thorson, Wesenberg-Lund), completely

ignored the former American publications containing some very good descriptions and figures of the

tube of Vermilia serrula well characterized by its

brood-chambers (Dawson, 1860; Verrill, 1874c,

1874d). Hartman (1944) did not refer to these latter

publications when she published the plates prepared by Verrill on the New England Polychaeta

or when she (Hartman, 1948, p. 50) discussed the

affinities of Chitinopoma and Miroserpula: "Chitinopoma may be nearly related to Microserpula [sic]

Dons for which the single known species, M. inflata

Dons, has been recorded from Arctic Seas (Brattstrom, 1945). In the latter, the tube is said to have

ovicels distally."

The original description of Vermilia serrula

Stimpson (1954), based on material from the Bay of

Fundy, is incomplete and only refers to the characters of the tube. No type material is known to exist.

Dawson (1860) then found this species on the coasts

of the Gulf of St. Lawrence (Gaspe" and Labrador).

Giving a detailed description of the tube with its

brood chambers near the orifice, he supposed that

"As they appear only in certain specimens, they may

perhaps be connected with the function of reproduction, and be of the nature of ovi-capsules."

Packard (1863, 1867) noticed the presence of V.

serrula in the Strait of Belle Isle. Verrill (1873, 1874

ad) mentioned V. serrula among the species

dredged off the coast of Maine and observed that

the brood chambers of the tube are only present

when mature but wanting in immature specimens.

All these early references of V. serrula are cited by

Verrill (1881) in a historical sketch on New England
NUMBER 42

Annelida, with annotated lists of the species hitherto

recorded.

Bush (1940), in a publication chiefly dealing with

species from Alaska but giving also a revision of the

genera of Serpulidae and mentioning species from

the Atlantic, Mediterranean, and other areas, did

not mention Vermilia serrula Stimpson. Nevertheless Bush (1940, p. 229, footnote, pi. 37: fig. 3, 9, pi.

40: fig. 31) figured two opercula and a special seta

of this species, referring the specimens to Levinsen's

Chitinopoma fabricii (as Chitinopoma greenlandica

[sic]: "Specimens attached to stones from Greenland and to tubes of Nothria conchylega from 32

fathoms off the New England coast are in the Yale

University Museum, and may prove to be the same

as those on the same host from Greenland identified

by Moore (1902) as Serpula sp.; these could not be

compared. The operculum is covered by a thin

chitinous cup-like plate." As clearly shown by the

figures, Bush's specimens were typical Chitinopoma

serrula. Under the name of Vermilia serrula, apparently no other structure of C. serrula than its

tube with the characteristic brood chambers has

been described. Bush (1904) is probably the only

American author who has figured a seta of this

species. The other published figures of V. serrula by

Dawson, Verrill, and Hartman only show the tube

and the operculum.

As the setae and uncini of Vermilia serrula remained unknown, the species was not mentioned

when a system of the Serpulidae, based on these

structures, was proposed (Saint-Joseph, 1894, and

others). Only Quatrefages (1865), referring to

Dawson (1860) for the brood chambers of the tube,

has cited V. serrula among the serpulid species

incertae sedis.

On the basis of material from Norway, and

independently from the American authors, Dons

(1931, 1934a, 1934b, 1937) gave a more complete

description of the species and created for it a new

genus, Miroserpula. Dons always wrote Miroserpula

and not Microserpula [sic], spelling used by Hartman (1948, 1959). Dons' description was taken up

and completed by Thorson (1946). Two further

papers dealt with the geographic distribution (Brattstrom and Thorson, 1941; Brattstrom, 1945). Records of this species are also found in WesenbergLund's papers (1950, 1951, 1953a, 1953b). It should

be noted that the latter author, ignoring the

identity of Vermilia serrula and Miroserpula inflata,

mentioned Chitinopoma serrula, in a survey of the

distribution of Polychaeta in Arctic and BoreoArctic areas, under the first name for the New

England region, and under the other name for

Greenland and other regions of the North Atlantic.

Levinsen (1883) created the genus Chitinopoma

for C. fabricii n. sp., a serpulid found commonly

on shells and algae from Greenland and the Faroes.

Levinsen's description is rather superficial and it

lacks details about the operculum as well as observation of the brood chambers. It can be summarized

as follows: Tube white, subtriangular in cross section, with transverse growth striations and a median

keel which is slightly denticulate. Total length of

the animal attaining 12 mm, diameter about 0.5 mm.

Thoracic setigerous segments 7. Gill filaments 6 on

each side, opercular peduncle non-pinnate. Operculum with a horny dark-coloured distal plate. First

thoracic setigerous segment with capillary setae and

special setae which are provided with a basal wing

of 4 teeth well-separated from the distal limbate

zone. Posterior segments with capillary and limbate

setae, and, beginning with the third segment, with

sickle-shaped setae. Uncini with 9 teeth.

Brattstrom (1945, pp. 19-22) has shown that the

tube of juvenile Chitinopoma serrula (as Miroserpula inflata), still lacking the characteristic brood

chambers and the lobated orifice, could easily be

mistaken for the tube of C. fabricii as described

and figured by Levinsen. As shown by the examination Of museum material from various museums, the

teeth of the keel, which in general are said to characterize the tube of C. serrula (as Miroserpula inflata),

may be much less developed than indicated by

Stimpson (1854), Dawson (1860), Dons (1931), and

others, and the tube would then resemble still more

closely that of C. fabricii, the keel of which should

be rather smooth.

Examination of the material in the collections of

the Copenhagen Museum (UZMC), labelled "Chitinopoma fabricii Levinsen," has shown that all

specimens of Chitinopoma are referable to C.

serrula, among them specimens from Eastport,

Maine, collected by Packard, 1866, and undoubtedly

seen by Levinsen himself. One specimen from the

"Dijmphna" expedition to the Kara Sea, labelled

"Chitinopoma fabricii Levinsen" (undoubtedly

determined by Levinsen and referred to in 1887),

turned out to be a Protula. The specimens from

East Spitzbergen (ZSZM), referred to by Augener
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(1928) as Chitinopoma fabricii, are typical C. serrula,

some of which have tubes with brood chambers.

The records of the North Atlantic Chitinopoma

serrula, as C. fabricii Levinsen, are rare and

Levinsen's original description was never emended.

Levinsen (1887) has cited C. fabricii among the

records from the "Dijmphna" Expedition in the

Kara Sea. Fauvel (1911a) cited a record from the

"Belgica" Arctic Expedition, in the strait between

Novaja Zemlja and the isle of Vaigach. According

to Derjugin (1915), C. fabricii is most abundant

in the innermost part of the White Sea, the Gulf

of Kola. Friedrich (1939) referred to C. fabricii

specimens from the Barents Sea, Augener (1928)—

specimens from Spitzbergen, Ditlevsen (1929) from

the Faroes, and Bush (1904)—specimens from New

England (see above). According to Wesenberg-Lund

(1951), the presence of C. fabricii in Icelandic waters

is not proved. Wesenberg-Lund (1953a) has then in

a rather questionable way referred empty tubes

from different localities on the Norwegian coast to

C. fabricii. From all these regions the true Chitinopoma serrula, with its characteristic brood chambers, is known.

Citations of Chitinopoma serrula, as C. fabricii

Levinsen, often contain errors caused by Levinsen's

insufficient description, or confusions with other

species. For instance, Levinsen's C. fabricii is cited

by Meyer (1888, p. 495) and by Saint-Joseph (1894,

pp. 246, 260, 264, 265) as a species lacking thoracic

membranes. This is a misinterpretation of the short

membranes which end between segments 2 and 3.

In the key to the serpulid genera, Southward (1963,

p. 587) indicates that the genera Miroserpula and

Chitinopoma are related but distinguished by the

basal denticulate expansion of the special setae.

However, belonging to the same species, M. inflata

Dons and C. fabricii Levinsen possess the same type

of special setae. Apparently, a reference to the

Pacific species Pseudochitinopoma occidentalis (see

below), often cited as Chitinopoma groenlandica (or

C. greenlandica—variant spelling), was used for this

key. Under the name of Chitinopoma greenlandica

[sic], Nelson-Smith thus confused Levinsen's Atlantic species together with the Pacific species.

Chitinopoma groenlandica (Morch, 1863) =

indeterminable serpulid

Serpula triquetra Fabricius, 1780, p. 379, no. 374. Not Linne,

1758.

Hydroides norvegica var. gronlandica Morch, 1863, p. 377.

THydroides gronlandica.—Malmgren, 1867, p. 120.

Not Chitinopoma greenlandica [sic.].—Pixell, 1912, p. 790

[ = Pseudochitinopoma occidentalis (Bush); see below.]

REMARKS.—Levinsen (1883, pp. 199, 203) referred

Serpula triquetra Fabricius (1780) to his new species,

Chitinopoma fabricii. However, Fabricius' Serpula

triquetra is indeterminable and had previously been

referred by Morch (1863, p. 377) to Hydroides

norvegica var. gronlandica. Malmgren (1867, p. 120)

mentioned this form as a questionable species, not

yet described: "7Hydroides gronlandica Morch. . .

Hab. in mari gronlandico. Species incerta, nondum

descripta." Nevertheless, Bush (1904, p. 229) and

Hartman (1948, p. 48) considered Levinsen's Chitinopoma fabricii as identical with Malmgren's

"}Hydroides gronlandica", the latter name would

then have priority. Thus Levinsen's species from

the North Atlantic is listed in Hartman's catalogue

(1959) under the name of Chitinopoma groenlandica. It is quite uncertain whether Fabricius1

Serpula triquetra from Greenland is identical with

the species described by Levinsen and which should

be named Chitinopoma serrula (Stimpson) (see

above).

Another reason for dropping the name of Chitinopoma groenlandica is that Pixell (1912), Berkeley

(1930), and Berkeley and Berkeley (1942,1952, 1961)

used the name of Chitinopoma groenlandica (or C.

greenlandica [sic]) for a serpulid from the Pacific

coast of North America quite different from the

North Atlantic species Chitinopoma serrula (see

below under Pseudochitinopoma occidentalis p. 7).

Pseudochitinopoma new genus

Gender: feminine. Type-species: Hyalopomatopsis

occidentalis Bush, 1904.

DIAGNOSIS.—Special setae of the first thoracic segment with a distal limbate zone and a proximal

wing not well separated. No thoracic sickle-setae.

Abdominal setae with a narrow spatulate end and

a long lateral tip. Thoracic uncini saw-shaped with

about 12 teeth, the anterior tooth bifurcate. Abdominal uncini in all segments rasp-shaped. Opercular stalk without pinnules or wings. Thoracic

membranes short, not united at the ventral face of

the last thoracic segment. Thoracic setigerous segments 7.
NUMBER 42

Pseudochitinopoma occidentalis (Bush, 1904),

new combination

FIGURE 2

Hyalopomatopsis occidentalis Bush, 1904, pp. 229-230, pi. 40:

figs. 3, 22; pi. 44: figs. 2, 4, 8, 9.—Moore, 1908, p. 362; 1923,

p. 254.—Hartman, 1942, pp. 89,90.

Chitinopoma greenlandica [sic].—Pixell, 1912, pp. 790-792,

pi. 88: fig. 6, a-c—Nelson-Smith, 1967, p. 37, fig. 29 (in

part). [Not Chitinopoma groenlandica (Morch, 1863).]

Chitinopoma groenlandica.—Berkeley, 1930, p. 74.—Berkeley

and Berkeley, 1942, p. 207; 1952, pp. 129-130, figs. 265, 266;

1961, p. 663. [Not Chitinopoma groenlandica (Morch,

1863).]

Chitinopoma occidentalis.—Hartman, 1948, pp. 49-51, fig. 12,

a-f; 1959. p. 570.

MATERIAL EXAMINED.—Off Brockway Point, Santa

Rosa Island, California: several spec, "Albatross"

St. 4431, 1904 (USNM 17318). Shelton, Washington:

20 spec, Muller coll., 1934 (USNM 36658). Brown

Island, San Juan Archipelago, Washington: 20 spec,

Pettibone coll., 1 August 1935 (USNM 36659). Wasp

Passage, San Juan Archipelago, Washington: several

spec, Pettibone coll., 27 July 1935 (USNM 38726).

Off Nigerson Point, Puget Sound, Washington: 20

spec, Pettibone coll., 29 June 1936 (USNM 36660).

Canoe Bay, Alaska: 3 spec, Alaska King Crab investigation, sta. 17,1940 (USNM 23621). Canoe Bay,

FIGURE 2.—Pseudochitinopoma occidentalis (from specimens from Wasp Passage, Washington,

USNM): a-c, opercula; d, special seta of first thoracic segment; e, abdominal seta; /, g, thoracic

uncinus, lateral and edge views.

368-077 O- 69 - 2
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Alaska: 2 spec, Alaska King Crab investigation, sta.

21, 1940 (USNM 21282). Canoe Bay, Alaska: numerous spec, Alaska King Crab investigation, sta. 24,

1940 (USNM 21272).

DESCRIPTION.—Tube white, solid, attached to the

substratum over its whole length, subtriangular in

cross section, with a more or less marked nondenticulate keel, somewhat similar to the tube of

Pomatoceros triqueter (L.) but smaller (length may

attain 50 mm, according to Hartman, 1948). Surface

of the tube almost smooth, sometimes with slightlyindicated transverse growth striations; median keel

forming a sharp tooth above the aperture. Longest

specimens examined measuring 13 mm (according

to Hartman, 1948, larger individuals measure 15

mm to 20 mm). Opercular diameter attains 800 /*.

Up to 60 abdominal segments. Collar divided into

a large median lobe and a pair of laterodorsal ones

which are continued into short thoracic membranes

ending at the limit of segments 2 and 3. Branchial

lobes short, supporting up to 10 gill filaments on

each side, inserted in a semicircle. No gill membrane. Operculum always on the left side, first nonpinnate filament transformed into an opercular

peduncle completely covered by a thick transparent

cuticle. Distal part of the operculum often tapering,

having more or less conical form, with a still thicker,

brown-colored cuticle. On preserved material, the

opercular tissues are often contracted inside the

cuticle which maintains the original form, thus

giving the impression of two opercula, one enclosed

in the other.

First thoracic setigerous segment with fine almost

capillary setae and special setae possessing a proximal denticulate expansion which is close to the

distal limbate zone and well separated. Setigerous

segments 2 to 7 with only two types of setae, capillary and simple limbate. No thoracic sickle-setae.

Abdominal setae with a narrow spatulate tip drawn

out on one side into a long point with a denticulate

edge. Setae of the posterior segments still narrower,

but not long capillary. Thoracic uncini saw-shaped

with 10 to 12 teeth; anterior tooth bifurcate; base

below the anterior tooth curved and forming a hook

directed slightly upwards. The number of uncini

in the thoracic rows increases slightly from segments

2 to 7. Abdominal uncini in all segments raspshaped.

DISTRIBUTION.—Pseudochitinopoma occidentalis

is known all along the Pacific coast of North

America, roughly between lat. 61 °N and 34°N:

southern coast of Alaskan Peninsula and Kodiak

Island, Chignik Bay, Prince William Sound, West

and East coast of Vancouver Island, Strait of Georgia and Puget Sound, off Santa Rosa Island and

Carmel Canyon, California.

The bathymetric range of the collection from

Alaska (Hartman, 1948) was from shore to 229 m.

The Berkeleys (1952) indicate for Vancouver Island

from lowest tide mark to 128 m. The southernmost

records, from off Santa Rosa Island (Moore, 1923)

and from Carmel Canyon (Berkeley and Berkeley,

1961), were from depths of 70 to 146 m.

REMARKS.—This species, known only from the

Pacific coast of North America, has often been

mistaken for the Atlantic Chitinopoma fabricii

Levinsen, which is identical with Chitinopoma

serrula (see above). In the original description as

Hyalopomatopsis occidentalis, by Bush (1904), most

of its essential structures are mentioned (tube, gill

tuft with operculm, collar and thoracic membranes,

special setae of the first segment, abdominal setae);

this description is completed by figures of the

operculum, the special setae, and the abdominal

setae. According to Bush (1904), the tube is white

and sinuous and has a well-marked keel, thus

resembling the tube of Chitinopoma fabricii as

figured by Levinsen (1883). It is undoubtedly due to

this remark on the similarities of the tubes that

Pixell (1912), when describing in detail Bush's

species, referred it, under the name of Chitinopoma

greenlandica [sic], to Levinsen's Atlantic species.

Possessing uncini of a quite different type, Bush's

species cannot belong to the genus Hyalopomatopsis

(which is a synonym of Hyalopomatus, see below).

The generic name Hyalopomatopsis has nevertheless been applied to this species by Moore (1908,

1923) and by Hartman (1942). The diagnosis of the

genus Chitinopoma, as given by Pixell (1912), is

based on the description of C. fabricii Levinsen

(identical with C. serrula). The latter species has

thoracic sickle-setae whereas Bush's species lacks this

type of setae. The thoracic sickle-setae were later

even incorrectly considered by Berkeley and Berkeley (1952) as a specific character of Bush's species.

Under the name of Chitinopoma occidentalis, Hartman (1948) gave a detailed description of Bush's
NUMBER 42

species, which was indicated to be different from the

Chitinopoma groenlandica reported by Pixell (1912)

and Berkeley (1930) from the northeastern Pacific.

Because of its important differences with Chitinopoma serrula (Stimpson) (=C. fabricii Levinsen),

Bush's species is placed here in the new genus

Pseudochitinopoma. The tabulation below summarizes the differences:

Setae Pseudochitinopoma occidentalis (Bush)

Thoracic uncini anterior tooth bifurcate.

Special setae of the first seg- basal expansion composed of basal expansion composed of a

Chitinopoma serrula

(Stimpson) (= C.

fabricii Levinsen)

anterior tooth simple.

ment

Thoracic sickle-setae

Abdominal setae

more numerous teeth; distal

limbate zone almost continuous with basal expansion.

absent.

narrow and spatulate tip, with

a long lateral point.

few teeth; distal limbate zone

well separated from basal expansion.

present.

geniculate, with a rather long

triangular blade.

Genus Chitinopomoides Benham 1927

Gender: masculine. Type-species: Chitinopomoides

wilsoni Benham, 1927, by monotypy.

DIAGNOSIS.—Special setae of the first thoracic segment with a large knob below the limbate zone [?].

Thoracic sickle-setae present. Abdominal setae geniculate with a long triangular blade. Thoracic

uncini saw-shaped with about 15 teeth; the anterior

tooth simple, not bifurcate. Abdominal uncini raspshaped in all segments. Opercular stalk without

pinnules and wings. Thoracic membranes short, not

united at the ventral face of the last thoracic segment. Thoracic segments 7.

Chitinopomoides wilsoni Benham, 1927

FIGURE 3

Chitinopomoides wilsoni Benham, 1927, pp. 156-158, pi. 5:

figs. 162-173.—Hartman, 1959, p. 570; 1966, pp. 128-131,

pi. 43: fig. 4-7.—Southward, 1963, p. 586.

MATERIAL EXAMINED.—McMurdo Sound, Ross

Sea: holotype, British Antarctic "Terra Nova" Expedition, 1910 (BMNH 1928:2:29:174).

DESCRIPTION.—Tube white, of almost circular

section, 2.2 mm in diameter, with a median keel

smooth and compressed, and two poorly-defined

smooth ridges along the sides. Distal end of the

tube projecting freely at right angles with its main

course. Total length of the animal about 25 mm,

of which the gill filaments account for 5 mm. About

50 abdominal segments. Collar continued in very

short thoracic membranes. Gill tuft composed of

about 14 filaments on each side. Opercular stalk

without pinnules and wings, its exact position unknown. Operculum partly constricted at two places

by deep grooves which, however, extend only for

a

FIGURE 3.—Chitinopomoides wilsoni (from holotype, McMurdo

Sound, BMNH): a, abdominal seta; b, c, thoracic uncinus,

lateral and edge views.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

about three-fourths of the circumference, and may

be artifacts.

First thoracic segment with almost capillary setae

and stouter special setae characterized by a "large

blunt knob some distance below the point or commencement of the wing" (this important detail

could not be verified on the fragments left of the

holotype as all setae of the first segment were

broken). Segments 2 to 7 with capillary setae and

ordinary limbate setae, together with a third type

of setae: in segments 3 to 7 these are typical thoracic

sickle-setae with a short proximal wing and a distal

blade with a blunt denticulation, whereas in segment 2 the setae have the same curved form as those

of the posterior segments, but have a continuous

wing instead of the blunt denticulation in the

distal part. The typical geniculate abdominal setae

with a long triangular blade give way to long capillary setae on the posterior segments. Thoracic uncini

saw-shaped (not rasp-shaped as indicated by Benham) with about 14 to 16 teeth; anterior tooth

simple (not bifurcate as indicated by Benham).

Abdominal uncini rasp-shaped in all segments, with

more teeth in posterior segments than in anterior

segments. Anterior abdominal uncini with at least

12 teeth when seen in profile.

DISTRIBUTION.—Chitinopomoides wilsoni is known

only from one specimen dredged in the Ross Sea

(McMurdo Sound, depth 366 m) by the British Antarctic "Terra Nova" Expedition (1910).

REMARKS.—In spite of the poor condition of the

holotype (gill tuft and operculum lost, only the

right half of the thorax present, with the setae of

the first segment all broken, and the abdomen

damaged) Benham's description can be improved on

many points, chiefly concerning setae and uncini.

The original description and figures of the setae

and uncini differ considerably from my observations

on these structures. So Benham's description of the

special setae has to be accepted with reservations.

The affinities of Chitinopomoides are hypothetical

and its generic justification remains questionable,

as the detailed structure of the special setae is not

well known. According to Benham, the species

agrees with the genera Chitinopoma and Hyalopomatus by the absence of thoracic membranes. However, as shown above, short thoracic membranes

were often considered as mere parts of the collar.

Uncini, setae, and operculum are not similar to

those of Spirobranchus, as indicated by Benham, but

rather to those of Chitinopoma. There are no structures similar to the genus Serpula, except perhaps

the special setae which, as indicated above, are

poorly known.

Genus Hyalopomatus Marenzeller, 1878

Hyalopomatopsis Saint-Joseph, 1894. [Type-species: Hyalopomatus marenzelleri Langerhans, 1884, by monotypy.]

Cystopomatus Gravier, 1911. [Type-species: C. macintoshi

Gravier, 1911, by monotypy. Referred herein to Hyalopomatus macintoshi (Gravier).]

Gender: masculine. Type-species: H. claparedii

Marenzeller, 1878, by monotypy.

DIAGNOSIS.—Special setae of the first thoracic segment with a uniform distal wing or having a proximal denticulate zone more or less separated. No

thoracic sickle-setae. Abdominal setae almost capillary, with only the tip geniculate. Thoracic uncini,

as well as abdominal uncini, rasp-shaped, with numerous small teeth (about 20 when seen in profile);

anterior tooth bifurcated. Opercular stalk without

pinnules or wings, very thin. Sometimes no operculum. Thoracic membranes short, not united at

the ventral face of the last thoracic segment. Thoracic setigerous segments 6.

REMARKS.—The three species mentioned in Hartman's (1959) catalogue under the name of Hyalopomatus claparedii Marenzeller, 1878, Halopomatus

langerhansi Ehlers, 1887, and Hyalopomatopsis

marenzelleri (Langerhans, 1884) are herein described under the common genus Hyalopomatus

Marenzeller because of their great similarities,

which were well recognized by the original authors.

When introducing the new genus Cystopomatus and

discussing the affinities of Cystopomatus macintoshi,

Gravier (1911) did not mention the species of

Hyalopomatus in spite of the great resemblance

shown by these forms with Cystopomatus macintoshi. The four species have the above generic

characters in common.

Differences between the species of Hyalopomatus

to which various authors have called attention (Langerhans, 1884, Ehlers, 1887, Knox, 1959) are those

of the thoracic membranes, special setae of the first

segment, and the thoracic uncini. However, these

proposed differences are not always of specific

validity since they may refer to an incorrect interpretation of short thoracic membranes considered

as collar lobes, superficial examination of setae and

figures of uncini.
NUMBER 42 11

Marenzeller (1878) and Knox (1959) have indicated that Hyalopomatus claparedii has no thoracic membranes and that the setae of the first

segment are simple limbate ones. Apparently the authors did not distinguish between short thoracic

membranes, which are always present at least to

the limit of segments 2 and 3, and the real collar.

Saint-Joseph (1894) followed Langerhans' (1884)

proposition of a generic separation. Hyalopomatus

marenzelleri Langerhans and H. langerhansi Ehlers

were said to differ from H. claparedii Marenzeller

and were referred to the new genus Hyalopomatopsis, characterized by the presence of short thoracic membranes and special setae with denticulate

basal wings.

The generic distinction between Hyalopomatopsis

and Cystopomatus according to a presumed difference in the length of the thoracic membranes

(Southward, 1963, p. 587) is not justified, as the

membranes end at about the same level and all

other details are the. same. As to the setae of the

first thoracic segment, the differences are less important than claimed. The separation into a distal limbate zone and a proximal wing of stronger teeth is

rather indistinct even in the setae of H. marenzelleri,

H. langerhansi and Cystopomatus macintoshi;

there is no wide gap as is found in the special setae

of Chitinopoma serrula (Stimpson). A more careful

examination may reveal that the distal limbate

zone of the setae of H. claparedii is perhaps less

uniform than shown by the very schematic figures

given by Marenzeller. These slight and even questionable differences seem to be rather artificial in

comparison with all characters common to the

species. For Hyalopomatus and Hyalopomatopsis,

Knox (1959) came to the same conclusion.

The abdominal setae of the species in question

have been referred to as capillary. However, closer

examination of H. marenzelleri and of Cystopomatus macintoshi has shown that the abdominal

setae of these species have their distal tip geniculate,

corresponding to about one-tenth of their length.

These setae differ from the geniculate abdominal

setae of Vermiliopsis, Omphalopoma, Chitinopoma,

etc. This detail was over-looked because of the very

small size of the abdominal setae. We may suppose

that the abdominal setae of the closely related H.

claparedii and H. langerhansi belong to the same

type and are not really capillary.

The known species of Hyalopomatus (including

Cystopomatus) are morphologically very similar and

closely allied. The synonymy of some of these forms

is not excluded. The genus contains species mostly

from bathyal depths in zones covering 150° of

latitude, as shown in the list below:

Hyalopomatus claparedii Marenzeller: Arctic Basin (FranzJoseph Land, Kara Sea, north of Ellesmere Land, north of

Alaska, between Hebrides and Iceland?)

Hyalopomatus sp.: Central Atlantic (West of Azores)

Hyalopomatus marenzelleri Langerhans: temperate East Atlantic (Bay of Biscay, Portugal, Josephine Bank, Madeira,

Azores)

Hyalopomatus sp.: Mediterranean (off Algeria, northern

Tyrrhenian Sea)

Hyalopomatus langerhansi Ehlers: tropical West Atlantic

(northwest of Cuba)

Hyalopomatus macintoshi (Gravier): Antarctic Sea (off South

West of Antarctic Peninsula)

Hyalopomatus claparedii Marenzeller, 1878

Hyalopomatus claparedii Marenzeller, 1878, pp. 393-394, pi.

4: fig. 2.—Ehlers, 1887, p. 307.—Mclntosh, 1879, p. 217.—

Langerhans, 1884, p. 278.—Brattstrom and Thorson, 1941,

p. 21.—Brattstrom, 1945, p. 1.—Uschakov, 1957, pp. 1668-

1669.—Knox, 1959, pp. 110-112, pi. 4: figs. 1-5—Hartman,

1959, p. 577.

Hyalopomatus claparedi.—Levinsen, 1883, pp. 195, 197, 310;

1887, p. 300.—Horn, 1882, p. 3.

Hyalopomatus.—Horn, 1931, p. 5; 1934a, pp. 24-26.

DESCRIPTION (taken from literature only).—Tube

white, round and smooth, sometimes with slightly

indicated growth rings; distal end free. Whole

length of the animal 10-23 mm, about half the

length for the gill tuft. About 30 abdominal segments. Collar a thin, transparent membrane composed of three lobes. Thoracic membranes short,

ending at the limit of segments 2 and 3. About 10

gill filaments on each side. Operculum a soft transparent vesicle, without any horny or calcified distal plate. Setae of the first thoracic segment with a

striated and denticulate limbate zone. Similar setae

in the following segments. Anterior and median

abdominal segments without setae; posterior segments with isolated capillary neuropodial setae.

Thoracic uncini roughly rectangular in shape, raspshaped, with numerous small teeth (about 20 when

seen in profile) and an anterior bifurcate process.

Abdominal uncini similar to those of the thoracic

region.

DISTRIBUTION.—Only few records in the Arctic
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Basin: Off southern Franz-Joseph Land (79° 13'1"N,

63°21'47"E, 230 m), off northern Ellesmere Land

(82°39'N to 82°50'N, 93°05'W to 94°10'W, 441 m

to 740 m), off north coast of Alaska (80°47'9"N,

170°48'W, 3622 m), Kara Sea, 128 m, between the

Hebrides and Iceland ? (61°02'N, 12°04'W, 1190 m).

REMARKS.—The original material of this Arctic

species described by Marenzeller was collected by

the Austrian Arctic Expedition (1873) off South of

Franz-Joseph Land. According to Marenzeller, a

serpulid specimen collected by the British "Lightning" Expedition (1868) between the Hebrides and

Iceland perhaps belong to the same species. Only

the transparent and vesicular operculum of this

specimen was described by Claparede in a report

on the Polychaeta of the "Lightning" Expedition

which, after the author's death, was published by

Ehlers (1875, p. 9, pi. 1: fig. 14). Without giving any

details, Levinsen (1887) referred two specimens from

the Kara Sea to Marenzeller's species. More recently,

H. claparedii was only cited as an Arctic species or

mentioned in discussions on related forms. The only

new records are given by Knox (1959) and Uschakov

(1957): some specimens and numerous tubes were

obtained at three stations by Fletcher's Ice Island

(1955) drifting off the northern Ellesmere Land,

two other specimens off the north coast of Alaska

in an abyssal station.

Hyalopomatus sp.

Gen. et sp. ? Eliason, 1951, p. 142.

MATERIAL EXAMINED.—Central Atlantic, west of

the Azores: 1 spec, Swedish Deep Sea Expedition,

sta. 387, 1948 (NMG).

REMARKS.—An undetermined tiny serpulid from

abyssal depths in the Central Atlantic west of the

Azores (40°33'N, 35°24'W to 40°34'N, 35°52'W,

4540 m to 4600 m) must be referred to the genus

Hyalopomatus. No really specific characters have

been observed on this specimen which lacked an

operculum and was in poor condition. The details

observed are general features of the genus Hyalopomatus: thoracic setigerous segments 6, of which 5

have rasp-shaped uncini with numerous teeth (more

than 15 when seen in profile, anterior tooth slightly

bifurcate); thoracic sickle-setae absent; abdominal

capillary setae observed only in the last segments;

thoracic membranes very short. On the basis of its

setae on the first thoracic segment, which have a

uniform simple limbate zone, the serpulid resembles

most closely H. claparedii as described by Marenzeller (1878) and Knox (1959).

Hyalopomatus marenzelleri Langerhans, 1884

FIGURE 4

Hyalopomatus marenzelleri Langerhans, 1884, p. 278, pi. 17:

fig. 49, a-d.—Ehlers, 1887, p. 307.—Knox, 1959, p. 111.

Hyalopomatopsis marenzelleri.—Saint-Joseph, 1894, pp. 224,

261.—Fauvel, 1909, p. 59; 1911b, p. 39; 1914, p. 341; 1932,

p. 39.—Hartman, 1948, p. 48; 1959, p. 577.—Southward,

1963, pp. 578-580, fig. 4, A-J.—Zibrowius, 1968a, p. 177;

1968b, pp. 385-386, 387, figs. 19-23.

MATERIAL EXAMINED.—Madeira: 4 spec, empty

tubes, French "Jean Charcot" expedition, several

stations, 1966. Southern Portugal: several spec and

tubes on corals, French-Portuguese "Faial" Expedition, sta. P. 59, 1957. Josephine Bank: empty tubes,

Swedish "Skagerak" Expedition, station of 31 May

1946 (NMG).

DESCRIPTION.—Tube white, round and smooth,

without keels, diameter about 500 ft. Part attached

to the substratum sometimes with some transverse

peristomes slightly widened out. Total length of the

animal about 11 mm, more than a third for the gill

tuft. Up to 45 abdominal segments. Collar trilobed,

short thoracic membranes ending at the posterior

limit of segment 2. Gill tuft composed of about 7

filaments on each side, first dorsal filament on the

left side non-pinnate, transformed into a very thin

opercular peduncle. Operculum a vesicle of globular

or slightly oval shape, with a distal, slightly convex,

slightly differentiated horny cap. First thoracic

setigerous segment with capillary setae and special

setae, the latter with a distal limbate zone and a

distinct but not well separated proximal wing of

stronger teeth. Segments 2 to 7 with limbate and

capillary setae. No thoracic sickle-setae. Distal part

of the abdominal setae (about one-tenth of their

length) geniculate. Thoracic uncini rasp-shaped

with the anterior tooth bifurcate; when seen in

profile with about 20 teeth. Abdominal uncini of

smaller size but similar.

DISTRIBUTION.—Hyalopomatus marenzelleri is

known from a large zone in the temperate North

Atlantic comprising the Azores, Madeira, and the

Gulf of Biscay. It is common in bathyal depths,

whereas its records from depths of only 100-200 m
NUMBER 42

FIGURE 4.—Hyalopomatus marenzelleri (from specimens from Madeira, "Jean Charcot" Expedition

1966): a, b, opercula; c, special seta of first thoracic segment; d, tip of abdominal seta; e, thoracic

undnus.

are less frequent: Azores (845-1850 m), Madeira

(100-2800 m), Josephine Bank (208-225 m), southern

Portugal (38°16'8"N, 8°56'43"W, about 300 m),

continental slope off Brittany and Scilly Islands

(366-1630 m).

REMARKS.—The original description of H. marenzelleri by Langerhans (1884) is very brief but illustrated by some figures and was based on specimens

from Madeira. More detailed descriptions were

furnished recently by Southward (1963) and by

Zibrowius (1968a, 1698b).

Hyalopomatus gp.

Protis arctica.—Bellan, 1964, pp. 179-180, 226.—Nelson-Smith,

1967, p. 50 (in part). [Not Protis arctica (Hansen, 1878).]

Hyalopomatopsis sp.—Zibrowius, 1968a, pp. 177-178, pi. 9:

fig. 36.

MATERIAL EXAMINED.—Off Algeria: mounted

specimens. Tyrrhenian Sea: 1 spec, "Calypso",

station SME. 1752, 1961.

REMARKS.—In the Mediterranean a form closely

allied to Hyalopomatus marenzelleri Langerhans

(if not the same species) has been recorded from

bathyal depths off the Algerian coast [on the

Gorgonian species Isidella elongata (Esper), in 570-

670 m] and in the Tyrrhenian Sea (South West of

Gorgona Island in 600-660 m). Bellan referred to

these specimens as Protis arctica (Hansen) (quoted

by Nelson-Smith, 1967) but, as shown by Zibrowius

(1968a), these serpulids with only 6 thoracic setigerous segments have the same type of setae and

uncini as Hyalopomatus marenzelleri. All five specimens known from the Mediterranean (4 from

Algeria, 1 from the Tyrrhenian Sea) had no operculum, but were otherwise identical with Hyalopo-
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

matus marenzelleri from Madeira. Rare specimens

lacking an operculum among typical operculate

specimens of this latter species have already been

found at Madeira by Langerhans (1884) and by

Zibrowius (1968a, 1968b). In some populations

there may be a tendency towards a reduction of the

operculum which even in normal individuals is

fixed to a very thin stalk.

Hyalopomatus langerhansi Elders 1887

Hyalopomatus langerhansi Ehlers, 1887, pp. 304-307, pi. 60:

figs. 14-15.—Hartman, 1938, p. 20; 1959, p. 577.—Knox,

1959, p. 111.

Hyalopomatopsis langerhansi.—Saint-Joseph, 1894, pp. 261,

264.

DESCRIPTION (taken from literature only).—Tube

white, somewhat flattened, with slight lateral keels

in the part attached to the substratum, but quite

smooth and of circular cross section in the erect

distal part. Total length of the animal about 11.5

mm, including 3 mm for the gill tuft. About 40

abdominal segments. Collar divided into 3 lobes;

short thoracic membranes ending at the limit of

segments 2 and 3. Gill tuft composed of about 12

filaments on each side. Operculum borne by the

first dorsal filament on the left side which is nonpinnate and longer than the other filaments. Operculum a transparent vesicle, pear-shaped, without

a differentiated distal plate. Special setae of the

first thoracic segment with a group of teeth at the

base of the limbate zone; these setae less numerous

than the setae of the following segments. Abdominal

setae capillary. Uncini of the thoracic region raspshaped with numerous teeth (about 20 when seen in

profile) and an anterior bifurcate process. Abdominal uncini of smaller size but similar.

DISTRIBUTION.—The two original finds of Hyalopomatus langerhansi were collected by the "Blake"

Expedition (1868), northwest of Cuba (near Havana

in 535 m, and 23°42'N, 83° 19^ , in 1573 m). There

seems to be no other record of this species in the

Gulf of Mexico.

REMARKS.—The above summary is taken from

the description and figures by Ehlers (1887), since

further details are not available, as the type material

(MCZ) is in bad condition (Hartman, 1938).

Hyalopomatus macintoshi (Gravier, 1911), new

combination

FIGURE 5

Cystopomatus macintoshi Gravier, 1911a, pp. 315-316; 1911b,

pp. 149-152, pi. 11: figs. 145-152.—Fauvel, 1936, pp. 39-40.—

Hartman, 1959, p. 572; 1966, pp. 128-129, 131, pi. 43: figs.

8-11.

MATERIAL EXAMINED.—West coast of Antarctic

Peninsula: 1 spec, incomplete, Second French

Antarctic Expedition ("Pourquoi-Pas"), 1908-1910

(MNHN). Bellingshausen Sea: 2 spec, incomplete,

"Belgica" Expedition, 1897-1899 (IRSN).

FIGURE 5.—Hylopomatus macintoshi (specimen from the

"Belgica" Expedition, Antarctica, IRSN): a, special seta of

first thoracic segment; b, abdominal seta; c, d, thoracic uncinus, lateral and edge views.
NUMBER 42 15

DESCRIPTION.—Tube white, smooth and of circular cross section, more or less curved or straight and

erect distally for some length. The whole length of

the tube may attain 50 mm, its diameter 750 /*.

Length of the animal up to 12 mm, including 4 mm

for the gill tuft. About 40 abdominal segments.

Collar large and divided into 3 lobes, the laterodorsal ones continued into short thoracic membranes ending at the limit of segments 2 and 3. Gill

tuft composed of about 12 filaments on each side.

Operculum a transparent almost globular vesicle

fixed to a very thin, smooth, non-pinnate opercular

stalk. The specimens studied are in too poor condition to show the insertion of the opercular stalk

or the existence of a gill membrane.

First thoracic segment with capillary setae and

stronger special setae, with a distal limbate zone

and a proximal more coarsely denticulate wing, the

two zones not well separated. The following 5

segments with only two types of setae: capillary and

limbate; no thoracic sickle-setae. The posterior abdominal setae are long and almost capillary, with

their distal tips (about one-fifteenth of their length)

slightly geniculate. The setae of the more anterior

segments are shorter and more geniculate (geniculate part corresponding to about one-tenth of their

whole length). Thoracic uncini rasp-shaped with 2

or 3 teeth in transverse rows, in lateral view with

about 20 teeth; the most anterior tooth large and

bifurcate. The number of thoracic neuropodial

uncini decreases in the posterior segments: there

may be 110 in the second setigerous segment but

only 80 in one row in segment 6. Abdominal notopodial uncini in rows of 20 to 30, rasp-shaped, with

a more curved profile than the rather rectangular

thoracic uncini and with more teeth in transverse

rows.

DISTRIBUTION.—The type specimens collected by

the Second French Antarctic Expedition ("Pourquoi-Pas," 1908-1910) are from Peterman Island off

the west coast of Antarctic Peninsula (about 67 °S,

69°W, no indication of depth, mixed with Serpula

vermicularis var. narconensis). The specimens from

the "Belgica" Expedition are from a large zone in

the Bellingshausen Sea (between 80"48^ and

87°37'W, 70°00'S and 71 "^S, in about 400-500 m).

No other records are known.

REMARKS.—Gravier's (1911a, 1911b) description,

in which some structures are neglected and the

importance of other structures over-estimated, deals

sufficiently well with the essential characters (setae

and uncini, operculum, tube) which are typical

Hyalopomatus characters. These were confirmed by

examination of original material, as well as the

identity of the material of both expeditions. Of

Gravier's material (MNHN), only one incomplete

syntype in poor condition is left (gill tuft and thoracic region, tube fragments). According to Fauvel

(1936), Cystopomatus macintoshi was collected by

the "Belgica" in 3 stations (28 May 1898, no. 996;

8 October 1898, no. 1010; 18 October 1898, no. 893;

the numbers refer to samples or bottles and not to

stations). There were several specimens and at least

two of them still had their operculum, but the

material of these 3 stations still available (IRSN)

is very poor (no tube fragments left; no. 996, one

specimen without a gill tuft; no. 983, one specimen

with an operculum but abdomen lost, compressed

by mounting (?); no. 1010, label only "Cystopomatus

macintoshi ??").

Genus Protis Ehlers, 1887

Gender: feminine. Type-species: Protis simplex

Ehlers, 1887, indicated by Hartman, 1959, p. 588.

Referred herein to Protis arctica (Hansen, 1878).

DIAGNOSIS.—Special setae of the first thoracic

segment with well separated distal limbate zone

and proximal denticulate wing. Thoracic sicklesetae present. Abdominal setae almost capillary,

with a limbate tip. Thoracic uncini saw-shaped with

about 6 teeth, the anterior simple, not bifurcate.

Abdominal uncini rasp-shaped in all segments. No

operculum. Thoracic membranes long, united at

the ventral face of the last thoracic segment. Thoracic setigerous segments 7.

REMARKS.—The first species of this genus to be

known, the North Atlantic Protis arctica (Hansen),

was described under the name Protula arctica and

was transferred by Ehlers (1887) into the genus

Protis created for a similar form from the Gulf of

Mexico, Protis simplex Ehlers, which seems to be

nothing more than a small specimen of Protis

arctica. Another very similar species, Protis pacifica

Moore, was described from the Pacific coast of

North America. Since an operculum is lacking,

which in most other serpulid genera provides good

specific characters, and all other structures are very

similar, the specific distinction of these forms is

rather questionable. It should be based on com-
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

parison of fresh material and not on the more or

less sketchy descriptions which are not always comparable.

As the serpulids from off Algeria considered by

Bellan (1964, p. 179) as Protis arctica do not belong

to the genus Protis but to Hyalopomatus (see above),

Bellan's considerations on the genus Protis (quoted

by Nelson-Smith, 1967, p. 50) are in error. The

genus Protis is not a transition between the genera

Protula and Filograna (=Salmacina); it seems not

even related to them, as claimed by Hartman (1955,

p. 51). Although without an operculum, it should

not be put together with these other non-operculate

genera. It is in fact a well-defined genus (cf. SaintJoseph, 1894, p. 259, and others) and well distinct

from Protula and Filograna with its thoracic uncini

(saw-shaped, with only a few teeth, resembling the

uncini of Serpula) whereas its special setae belong

to a more common type (e.g., the genera Omphalopoma and Filograna).

Protis arctica (Hansen, 1878)

FIGURE 6

Protula arctica Hansen, 1878, pp. 13-14, pi. 10: figs. 1-10;

1882, p. 43, pi. 7: figs. 28-34.—Levinsen, 1883, pp. 202, 309.

—Ehlers, 1887, p. 328.—Bidenkap, 1895, pp. 41, 45, 141.—

WoUebaek, 1912, pp. 120-122, pi. 48: fig. 1-6, pi. 50: fig. 7.—

Saemundsson, 1918, p. 230.—Wesenberg-Lund, 1950, pp.

41-42, chart 66; 1951, pp. 131, 156, 167 (in part?); 1953b,

pp. 114, 123, 138, 139, 149, 158, 162, 164, chart 26.

Protis simplex Ehlers, 1887, pp. 325-328, pi. 56: figs. 10-14.—

Bush, 1904, pp. 227, 229.—Eliason, 1951, p. 142.—Hartman,

1955, p. 51; 1959, p. 588.

Protis arctica.—Fauvel, 1909, pp. 50-51; 1914, pp. 325-327, pi.

30: figs. 18-25.—Eliason, 1951, p. 142—Hartman, 1959, p.

588.—Zibrowius, 1968a, p. 178. Not Bellan, 1964, pp. 179-

180, 226. Not Nelson-Smith, 1967, p. 50 ( = Hyalopomatus

sp.).

MATERIAL EXAMINED.—Romanche Deep in Central Atlantic: 1 spec, Swedish Deep Sea Expedition,

sta. 342 (NMG).

DESCRIPTION.—Tube white, of circular cross section and more or less distinctly marked with

transverse growth striations, similar to a tube of

Protula. Longest tube known 77 mm, diameter

about 3 mm. Total length of the animal may attain

20 mm to 30 mm. Up to more than 100 abdominal

segments. Collar divided into 3 lobes. Thoracic

membranes long, united at the ventral face of the

last thoracic segment. Gill tuft without an operFIGURE 6.—Protis arctica (from specimen from Romanche

Deep, Swedish Deep Sea Expedition, NMG): a, tip of abdominal seta; b, thoracic uncinus.

culum. Up to 23 gill filaments on each side. No

gill membrane [?]. First thoracic segment with capillary setae and stouter special setae which have a

proximal denticulate wing well separated from the

distal limbate zone. Posterior thoracic segments with

sickle-shaped setae with a proximal limbate zone

and a blunt denticulation on the distal edge. Segments 2 to 7 with capillary setae and simple limbate

setae. Abdominal setae of capillary aspect but in the

distal tip with a short and narrow wing. Thoracic

uncini saw-shaped with about 6 teeth, anterior

tooth bigger and simple, not bifurcate. Abdominal

uncini of similar form, in all segments rasp-shaped.

Uncini of the posterior segments with more teeth

than those of the more anterior segments.

DISTRIBUTION.—The original specimens of Protis

arctica described by Hansen (1878, 1882) were collected by the Norwegian North Atlantic Expedition

between Iceland and Norway (65°53'N, 7°18'W,

2127 m, temperature at the botton — 1.1°C). WoUebaek (1912) gave a more detailed description of the

species, but gave no new records. Many new records

from the Arctic North Atlantic were later given by

Wesenberg-Lund (1950, 1951,1953b), but, as proved

by examination of some of this material (UZMC)

(see below), at least some of the records from the

fjords of East Greenland in depths of less than 100

m are records of Protula tubularia and not of Protis

arctica. Records from the fjords of Iceland perhaps
NUMBER 42 17

are not Protis arctica either, whereas the other

records from bathyal and abyssal depths in the

Arctic Atlantic (between Labrador and Greenland,

between Greenland and Iceland, between Iceland

and Jan Mayen, depths down to 2402 m) may well

refer to Protis arctica.

Fauvel (1909, 1914) recorded Protis arctica from

the Azores between Flores and Faial (39°22'48"N,

31°25'15"W, in 1384 m) and between the Azores

and Maderia (36°54'N, 20°46'15"W, in 440 m).

In the Gulf of Mexico the species is known as Protis

simplex Ehlers, 1887 (28°42'N, 83°13'W, in 1572

m). The southernmost record of Protis in the Atlantic (Eliason, 1951) is from the Romanche Deep in

the Central Atlantic (about 01 °N, 18°40'W, in

5250 m).

Protis arctica had not been found in the Mediterranean, where the bottom water in the great depths

would probably be too warm for this species. It has

not been recorded from the Indian Ocean (erroneous quotation of Eliason, 1951, by Bellan, 1964, p.

181, and copied by Nelson-Smith, 1967, p. 50).

REMARKS.—Protis arctica is known by several

more or less detailed and illustrated descriptions

(chiefly Hansen, 1878, 1882; Wollebaek, 1912;

Fauvel, 1914). Although Ehlers (1887) indicated the

differences between the genera Protula and Protis,

Hansen's species was later attributed to the genus

Protula by some authors, chiefly by Wollebaek

(1912) and Wesenberg-Lund (1950, 1951, 1953b).

The latter (1950, p. 42), however, had emphasized

the difference between the uncini of Hansen's

species and those of Protula tubularia (Montagu).

Despite this exact comparison, Wesenberg-Lund

has sometimes mistaken specimens of Protula tubularia for Protis arctica, for 6 lots of serpulids from

East Greenland (UZMC) turned out to be Protula

tubularia and not Protis arctica (Hansen) as indicated by Wesenberg-Lund (1951, p. 114). Some of

these specimens were still in their tubes and could

not have been studied in detail; they must have

been determined on the aspect of the tube. This

casts doubt on the distributional survey of Protis

arctica indicated by Wesenberg-Lund.

There are no confirmed features which allow a

specific distinction between the forms described as

Protis arctica, mostly from the North Atlantic, and

Protis simplex Ehlers from the Gulf of Mexico. The

latter form, described from a single specimen, was

said to differ from the North Atlantic representatives of Protis by its smaller size and smaller number

of gill filaments. Fauvel (1909, 1914) described

specimens of intermediate size and showed that

these differences are insignificant. The synonymy of

Protis simplex with Protis arctica was also accepted

by Eliason (1951), who recorded a rather large specimen from the Central Atlantic near the equator.

Protis pacifica Moore, 1923

FIGURE 7

Protis pacifica Moore, 1923, pp. 253-254.—Hartman, 1955, pp.

51, 52, pi. 4; 1959, p. 588; 1960, pp. 163, 164; 1963, p. 79 —

Hartman and Barnard, 1958, pp. 2, 10, 11, 27, etc., 65.

MATERIAL EXAMINED.—Off Brockway Point,

Santa Rosa Island, California: holotype and 4 paratypes, "Albatross" sta. 4433, 1904 (USNM 17434,

17435).

DESCRIPTION.—Tube white, straight or more or

less tortuous and bent, approximately cylindrical,

of circular cross section as its distal and free end

but slightly flattened where attached to the substratum. No keels, surface somewhat rugose due to

the presence of irregularly spaced annulated growth

rings. Longest tube described 65 mm, thickest one

4.2 mm at the mouth. The length of the animal may

attain 40 mm. Sometimes more than 60 abdominal

segments. Collar divided into 3 lobes, the laterodorsal ones continued in long thoracic membranes

which are united at the ventral face of the last

thoracic segment. Gill tuft without an operculum.

Gill lobes short, with the filaments arranged in a

semicircle. Up to 20 filaments on each side.

First thoracic segment with capillary setae and

stouter special setae, with a well separated distal

limbate zone and a proximal wing of about 6 teeth.

Segments 2 to 7 with capillary setae and simple

limbate ones. Segments 3 to 7 also with sickle-shaped

setae and with a short proximal limbate zone

preceeding the distal edge with a blunt denticulation. Posterior abdominal segments with long capillary setae. Setae of the anterior segments almost

capillary but with a very small narrow limbate

edge just below the tip. Thoracic uncini saw-shaped,

with 6 to 8 teeth, the anterior one simple, not

bifurcate. Abdominal uncini of similar form but

rasp-shaped in all segments. Uncini of the posterior

segments with more teeth than those of the more

anterior segments.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Protis pacifica (from holotype, Santa Rosa Island, USNM): a, special seta of first

thoracic segment; b, thoracic sickle-seta; c, tip of abdominal seta; d, e, thoracic undnus, lateral

and edge views.

DISTRIBUTION.—In the Pacific the genus Protis is

known by one species from off the California coast.

The original specimens of Protis pacifica Moore

were dredged off Santa Rosa Island (depth about

485 m). Many new records were given by Hartman

(1955, 1960, 1963) and by Hartman and Barnard

(1958) from deep basins and submarine canyons off

California (depths from several hundred meters

down to 787 m). It seems to be an exclusively

bathyal species.

REMARKS.—There appears to be a slight difference in the form of the uncini from the Atlantic

Protis arctica. This, however, should be confirmed

by a detailed study of more specimens from both

oceans. Otherwise Protis pacifica is most similar to

Protis arctica.

Pomatostegus stellatus (Abildgaard, 1779)

Protis torquata Hoagland, 1919, pp. 580-581, pi. 32: figs. 9-13.

—Treadwell, 1939, pp. 304-305.—Hartman, 1955, p. 51;

1959, p. 588.

MATERIAL EXAMINED.—Puerto Rico, San Juan,

Condado Bay: 1 spec, lacking operculum, 2 July

1915, holotype of Protis torquata Hoagland

(AMNH). Puerto Rico, St. Thomas, Jamaica,

Quintana Roo (Mexico): several spec. (USNM).

Fernando da Noronha Island, Brazil: 1 spec,

Laborel coll.
NUMBER 42 19

REMARKS.—Protis torquata was described from a

single nonoperculate serpulid specimen from intertidal rock crevices at Condado Bay, Puerto Rico.

The holotype (AMNH) proved to be a young specimen of Pomatostegus stellatus (Abildgaard) which

had accidentally lost its operculum. The base of the

thick opercular stalk is still visible on the left side

of the gill tuft. The specimen agrees in structures,

such as thoracic membranes, setae and uncini, with

complete specimens of Pomatostegus stellatus from

Brazil and from the West Indies (USNM). (See also

the new definition of the genus Pomatostegus,

Zibrowius, 1968a, p. 153).

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NUMBER 42 21

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