This article was downloaded by: [University of Arizona]On: 27 June 2014, At: 06:37Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH,UK

Annals and Magazine of NaturalHistory: Series 13Publication details, including instructions forauthors and subscription information:http://www.tandfonline.com/loi/tnah19

On some polychaetes from theFirth of Clyde and the West ofScotlandR.B. Clark a & P.G. Dawson aa Department of Zoology , University of BristolPublished online: 10 Aug 2009.

To cite this article: R.B. Clark & P.G. Dawson (1963) On some polychaetes from theFirth of Clyde and the West of Scotland, Annals and Magazine of Natural History:Series 13, 6:71, 641-657, DOI: 10.1080/00222936308651411

To link to this article: http://dx.doi.org/10.1080/00222936308651411

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all theinformation (the “Content”) contained in the publications on our platform.However, Taylor & Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness,or suitability for any purpose of the Content. Any opinions and viewsexpressed in this publication are the opinions and views of the authors, andare not the views of or endorsed by Taylor & Francis. The accuracy of theContent should not be relied upon and should be independently verified withprimary sources of information. Taylor and Francis shall not be liable for anylosses, actions, claims, proceedings, demands, costs, expenses, damages,and other liabilities whatsoever or howsoever caused arising directly orindirectly in connection with, in relation to or arising out of the use of theContent.

This article may be used for research, teaching, and private study purposes.Any substantial or systematic reproduction, redistribution, reselling, loan,

sub-licensing, systematic supply, or distribution in any form to anyone isexpressly forbidden. Terms & Conditions of access and use can be found athttp://www.tandfonline.com/page/terms-and-conditions

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

ON SOME POLYCI-IAETES F g O M THE FIgTI:I OF CLYDE AND THE WEST OF SCOTLAND.

By R. B. CLARK and P. G. DAwson,

Department of Zoology, University of Bristol.

THE list of polychaetes known from the Clyde Sea Area published in 1960 (Clark, 1960) was based on a compilation made at the beginning of 1957. Since then, a great many worms have been collected from the Firth of Clyde, mostly from sub-littoral deposits and mostly by successive classes in marine biology held at the Millport Marine Station. These collections have been sent to the authors for examination and have yielded several worms of general taxonomic interest, and a number of species not previously known from the west of Scotland. The relatively small number of new Clyde records is partly a test imony to the effectiveness of previous surveys in the area and part ly due to the fact that dredging tends to be carried out in the same places year by year. Unfortunately, a number of collec- tions were general ones made during the classes, and details of the locality, depth, substratum, etc., were not always recorded. All that can be said of them is that they were made from well-known dredging grounds near Millport and not in the remoter parts of the Clyde Sea Area.

A few specimens collected before the compilation of the fauna list, but not previously reported upon, and others collected off the west coast of Scotland, outside the Clyde area, have been included in this account.

The nomenclature follows that of Hartman (1959) in the main, although this involves the alteration of some generic names which have become familiar by long usage.

Specimens of all the species discussed are included in the reference collection of polychaetes at the Marine Station, Millport, and, where possible, duplicates have been deposited in the Royal Scottish Museum, Edinburgh.

We are extremely grateful to Dr. R. B. Pike for sending us the greater part of this collection of polychaetes for examination, and also to the Director of the Millport Marine Station and the Keeper of Zoology in the Royal Scottish Museum for financial assistance.

Family APHRODITIDAE Savigny. Harmothog joubini Fauvel.

Fauvel, 1923, p. 70.

Two specimens taken in dredgings. Details of time and place not recorded. A third specimen was found in the tube of a large maldanid from the Fairlie Channel, off Farland Point, at a depth of 20-30 m. on August 30th, 1957.

A.M.N.H. ser. 13, vol. vi. 41

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

642 R . B . Clark & 1 ). G. Da~son : polychaetes from Scotland

The species has not previously been recorded from British waters and it is only known from its original discovery by Fauvel (1914) in deep water in the Atlantic, 60 miles S.S.E. of Cape St. Vincent. The Clyde specimens differ from Fauvel 's only in that the elytra, instead of being uniformly whitish, are marked with a faint brown line of pigment around the posterior margin and across the middle, rather in the manner of the pigmentation of the elytra of H. lunulata.

Pholog minuta (Fabricius).

[ Pholo~ synophthalmica Clapar~de.] Fauvel, 1923, p. 120.

P. minuta was included in the 1960 fauna list on the strength of the record of a single specimen collected many years ago at Millport by David Robertson. Typical specimens of this species have never been found in the Clyde since then, although McIntosh (1900) claimed that it occurs " everywhere on the shores of Britain from Shetland to the Channel Islands "

In recent years, nine specimens answering to the description of P. synophthalmica have been collected at Millport. Of these, four were taken in bot tom samples on August 2nd, 1957, from a muddy-sand substratum in the Fairlie Channel (depth not recorded), and five were taken there on April 6th, 1959, in a silk tow net dragged over the surface of the sand at a depth of 20 m. This species has not previously been recorded from British waters.

Several species of Pholo$" are very similar to P. minuta and it is doubtful if they can all be separated. Marion and Bobretzky (1875) and Saint-Joseph (1887) supported Clapar~de (1868) in his separation of P. synophthalmica from P. minuta, although Saint-Joseph described a variety, P. minuta var. dinardensis, which is intermediate between the two. Both McIntosh (1900) and Fauvel (1923) regarded P. synophthalmica as a variety of P. minuta without formally designating it as such, and Har tman (1959) assigns it to the latter species. Three other species occupy similar positions. Pholo@ inornata Johnston (1839) was regarded as a variety of P. minuta by Fauvel (1923), and Pett ibone (1963) reduces it to synonymy with this species. Pholog eximia Michaelsen (1897) is another variety of P. minuta (Fauvel, 1923) and P. tuberculata Southern (1914) is regarded as yet another variety by Pett ibone (1953) and Southward (1956).

P. synophthalmica is distinguished from P. minuta on the following grounds: the eyes of the former species are fused on each side of the prostomium, the antennee are bulbous at the base, the papillae on the elytra are not moniliform, and there are no fine spines on the shafts of the compound, hooked chaetee. In addition, the elytra of P. minuta are generally marked with fine brown spots, while those of P. synophthalmica are unpigmented. In all these respects the Clyde specimens agree with the description of P. synophthalmica, but the value of these distinctions is questionable. The eyes of P. minuta are contiguous (McIntosh, 1900) and

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 643

in the specimens collected by Southward (1956), they are fused. The swelling at the base of the antennae of P. synophthalmica is very slight and not a convincing difference between the species. The papillm on the elytra of present specimens are not moniliform, but they have a swollen and ridged appearance at their base. The degree of spinulation on the shaft of the hooked ehaetse differs in varieties of P. minuta (Fauvel, 1923) and also appears to be an unreliable character.

The Clyde specimens are relatively small. The four intact specimens measure between 4 and 6 mm. long and include 28-36 segments; they may well have been young worms. From our knowledge of growth processes in other polychaetes, the possibility that the features of typical P. minuta may develop later appears considerable.

Family SYLLIDAE Grube. Odontosyllis gibba Clapar~de.

Fauvel, 1923, p. 275.

Twelve specimens varying between 5 and l0 mm. and 30 to 41 segments long. Of these, six were dredged in Kames Bay, MiIlport, from a sandy substratum at a depth of 8 m. on April 20th, 1960. The remainder were collected from unspecified areas of the Firth of Clyde on April 26th, 1959. The eolour of several of the specimens is olive green, shading to yellowish- white ventrally, with a dense brown patch at the dorsal base of alternate parapodia. Other specimens are uniformly yellowish-white apart from the pigment spots on the parapodia. The species is new to the Clyde fauna.

Eusyllis blomstrandi Malmgren. Fauvel, 1923, p. 293.

A single specimen dredged in shallow water in White Bay, Cumbrae, on December 24th, 1953. I t was found in a clump of the hydroid Tubularia and was very phosphorescent. I t was previously known in British waters only in the neighbourhood of Plymouth (Marine Biological Association, 1957).

Family NEPItTYDIDAE Grube. Nephtys caeca (Fabricius).

Fauvel, 1923, p. 365.

This species, though frequently reported, is, in fact, uncommon on most British coasts (Clark and Haderlie, 1960; Clark, Alder and McIntyre, 1962) and in the recent fauna list (Clark, 1960) it was pointed ou t that many of the records of intertidal N. caeca from the Clyde undoubtedly should refer to N. cirrosa Ehlers, and the only authenticated record of the former species was in the sub-littoral of White Bay, Cumbrae, at depths between 6 and 27 m. (Clark and Milne, 1955). Two further specimens of N. caeca were dredged from the Fairlie Channel off Lion Rock and in mid-channel at a depth of 40 m. on August 2nd, 1957.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

644 R . B . Clark & P. G. Dawson : polychaetes from Scotlana

Family SPHAERODORIDAE Malmgren.

Sphaerodorum claparedii Greeff. Fauvel, 1923, p. 379. Two specimens, 2 mm. and 3 mm. long, were taken in the dredge in

Kames Bay, Millport, at a depth of 40 m. on April 20th, 1960. The species is new to the Clyde fauna.

Family GLYCERIDAE Grube. Glycera capitata Oersted.

Fauvel, 1923, p. 385. Gemmill (1901) and Chumley (1918) claimed that this species was

common in all parts ofthe Clyde Sea area. However, it was not rediscovered during the intensive sublittoral sampling carried out between 1944 and 1950, and G. rouxii Audouin and Milne-Edwards was found to be the dominant glycerid of muddy and sandy deposits. I t was suggested (Clark, 1952, 1960) that there might have been a confusion of species by the earlier investigators and reports of the presence of G. capitata in the Clyde were viewed sceptically. Twelve specimens of this species have now been found in a general collection of worms from the area. Details of place and time were not recorded.

This species has a very wide distribution on both sides of the Atlantic (Southern, 1914; Fauvel, 1923; Stop-Bowitz, 1941; Hartman, 1944a; Pett ibone, 1963), in the Arctic (Stop-Bowitz, 1948; Wesenberg-Lund, 1950 a), Persian Gulf and Indian Ocean (McIntosh, 1924) and Pacifio (Aziz, 1938). I t has been recorded from the Irish Sea (Southward, 1956) and its occurrence in the Clyde is therefore not surprising.

Family EUI~ICIDAE Grube. Marphysa belli (Audouin and Milne-Edwards).

Fauvel, 1923, p. 410. The anterior end of a fairly small individual, 2 mm. broad, was dredged

in Kames Bay, Mfllport, at a depth of 60 m. on September 21st, 1954. This species has previously been reported only from the south-west of the British Isles, at Clare Island (Southern, 1914), Plymouth (Marine Biological Association, 1957), and is new to the Clyde fauna.

Hyalinoecia bilineata Baird. Fauvel, 1923, p. 422. Three specimens in membranous tubes slightly encrusted with sand,

collected in the Fairlie Channel off Lion Rock at a depth of 35 m. on August 2nd, 1957. The branchim begin at the fourth setiger in all the specimens and the dorsal surface of each segment is marked with two brown lines which extend the whole length of the worm. There is a patch of brown pigment at the base of each parapodium. This species has been reported from a variety of sites around the south-western British Isles (McIntosh,

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 645

1910; Southern, 1914; Marine Biological Association, 1957) and also off North Unst in the Shetlands (MeIntosh, 1910). I t appears to be a warm- water species and has not previously been found in the Clyde.

Lumbrinereis gracilis (Ehlers). Fauvel, 1923, p. 432.

A single specimen was taken off Little Cumbrae on August 24th, 1954. I t agrees exactly with Fauvel 's description of the species and is new to the Clyde fauna.

Family ORBINIIDAE Hartman. Orbinia cuvierii (Audouin and Milne-Edwards).

Fauvel, 1927, p. 12, as Aricia cuvieri.

Two specimens were collected in Loch Sween, Argyllshire, on September 29th, 1957, at a depth of 12-15 m. from a substratum of black mud. This species has never been found in the Clyde and has not previously been recorded from the west of Scotland. Elsewhere it occurs on south-western coasts of the British Isles (Southern, 1914; Bassindale and Barrett, 1957 ; Marine Biological Association, 1957) and also in the Irish Sea (McIntosh, 1910; Moore, 1937).

Family SPIONIDAE Grube.

Scolelepis ciliata (Keferstein). Fauvel, 1927, p. 30.

A single specimen was collected from sub-littoral deposits. Details of time and place are not recorded. This species, known from the Norwegian and Swedish coasts (SSderstrSm, 1920), the North Sea and Channel (Fauvel, 1927), has previously been recorded from British waters only in the Irish Sea (Southward, 1956).

Nerinides cantabra Rioja. Fauvel, 1927, p. 31.

Single, incomplete specimen collected in sub-littoral deposits. Details of time and place are not recorded. The only previous British record of this species is from Blacksod Bay, south-west Ireland (Southern, 1914, as ~V. longirostris). I t is known from the French and Spanish Atlantic coasts (Saint-Joseph, 1894; Mesnil, 1896; Rioja, 1919).

Aonides oxycephala (Sars). Fauvet, 1927, pp. 39, 40. Four badly damaged anterior ends up to 25 mm. long, and a number of

posterior fragments, probably of the same species were collected east of Little Cumbrae from coralline deposits at depths of 4 to 6 m. on August 24th, 1954.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

646 R . B . Clark & P. G. Dawson : polychaetes from Scotland

The four specimens of which positive identification can be made are intermediate between A. oxycephala and A. paucibranchiata Southern (1914), a species described from south-west Ireland and which Fauvel (1927) suggests may be a young form of A. oxycephala. The relevant characteristics of each are given in the following table.

A. oxycephala Clyde specimens A. paucibranchiata

Length 50-100 ram.

Prostomium with cephalic tentacle

Eyes smull, arranged in a trapezium.

20-24 pr. branchiae

Hooked chaet~e bidentate.

25 mm. (incomplete)

No cephalic tentacle, posterior margin obtuse.

Eyes small, arranged in a trapezium.

11-15 pr. branchim.

Hooked chaetse bidentate.

18 ram. (incomplete)

No cephalic tentacle, posterior margin obtuse.

Eyes with a lens.

10-11 pr. branchiae.

Hooked chaet~e tr identate.

More northerly specimens of A. oxycephala may differ slightly from those Fauvel is likely to have examined, since McIntosh (1915) figures the anterior end of this species and shows the posterior margin of the prosto- mium to be obtuse and almost to resemble that of A. paucibranchiata. I t is impossible to evaluate the significance of the difference between the eye structure of the two forms, and Fauvel (1927) suggests that the difference in the number of branchim and in the shape of the hooked chaet~e could well be due to differences in age and state of maturity.

Unless better evidence can be found, it seems preferable to include A. paucibranchiata in A. oxycephala; it is impossible to assign the Clyde specimens to either species in their narrow difinition. The genus has not previously been reported in the Clyde area.

Spiophanes kroyeri Grube. Grube, 1860, p. 88.

Fragments of a single specimen taken in the Fairlie Channel off Lion Rock at a depth of 35 m. on August 2nd, 1957. This arctic species was previously known from the Clyde only from an old record of a single specimen collected off Davarr Island by Gemmill (Clark, 1960).

Family CIRRATULIDAE Carus.

Cirriformia tentaculata (Montagu). Fauvel, 1927, p. 91, as Audouinia tentaculata.

Two specimens. Details of time and place of collection are not recorded. This species, common in more southerly British waters, was not previously known from the Clyde.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

It. B. Clark & P. G. Dawson : polychaetes from Scotland 6~7

Family SCALIBREGMIDAE )JIalmgren.

Polyphysia crassa (Oersted). F a n vel, 1927, p. 127, as E u m e n i a crassa.

A single specimen, probably ref~rrable to this species, was collected from a sticky, black mud deposit off Cloch. In its contracted state it is 14 ram. long, and includes 32 segments. The anterior end is dilated and about 5 mm. in diameter. At segments 12-13, the body narrows suddenly to a diameter of 3 mm., and there is a short " tail "region 0.5 ram. in diameter, composed of the terminal four or five segments. The body configuration is somewhat reminiscent of that of Scalibregma. The specimen lacks gills.

The presence of 4-6 pairs of gills on anterior segments of Polyphysia is chief feature that distinguishes this species from Lipobranchius (~[cIntosh, 1869, 1915; Cunningham and Ramage, 1888; Ashworth, 1901; Fauvel, 1927) and Eliason (1920) has suggested that Lipobranchius is only a young, immature, abranchousformofPolyphysia. The branchim of scalibregmids appear to be an extremely variable feature of the worms. Scalibregma inflatum Rathke typically has 4 or 5 pairs of branchise, but McIntosh (1915) remarks that they may be retractile, and specimens from Ireland (Ashworth, 1909; Southern, 1914) and Millport (Clark 1954) apparently lack them altogether. Ditlevsen (1911) and Stop-Bowitz (1945) have found up to 4 pairs of branchise on mature specimens of Lipobranchius jeffreysii (Mclntosh) and regard them as an epitokal modification of the abranchiate, immature form. However, mature worms in the Clyde do not possess branchise (Clark, 1954). Clearly, the presence or absence of gills cannot be regarded as a character of taxonomic significance in these scalibregmids.

Mclntosh (1915) notes that the anus of L. jeffreysii is surrounded by short papillae, but that the anal segment of Polyphysia lacks papillze, although the rectal region may be everted in preserved worms as a crenate, trumpet-shaped projection. This is the condition of the present specimen. The presence or absence of papillae around the anus may be a sufficient means of separating P. crassa from L. jeffreysii, bu t without examining a series of specimens, which we do not have at our disposal, it is impossible to be certain of this.

A more subjective difference between the present specimen and typical Lipobranchius is the manner in which it has contracted. The body forms a swollen anterior end with an abrupt transition to a narrower section, followed by the final four or five segments which are extremely narrow and form a " tail ". None of the several hundred Lipobranchius jeffreysii from the Clyde, that we have examined, have contracted in this way. Usually, both alive and preserved, they have the appearance of a blunt sausage. The quite different aspect of the present specimen from L. jeffreysii suggests more strongly than any preciso taxonomic character, tha t it is not the latter species but P. crassa, the absence of gills notwith- standing.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

648 R . B . Clark & P. G. Dawson : polychaetes from Scotland

P. crassa has previously been recorded from British waters only by Johnston (1865) from the Staples, by Cunningham and Ramage (1888) from the Firth of Forth, and by McIntosh (1915) from the stomach of haddocks at St. Andrews. Elsewhere, it has been found in the north Atlantic and Arctic (McIntosh, 1915; Fauvel, 1927; Pettibone, 1956).

Family OPHELIIDAE Malmgren. Ophelia rathkei McIntosh.

Tebble, 1952, p. 567.

The identity of the Clyde Ophelia has been in doubt for some years. Three species have, at various times, been described from the area. O. limacina (Rathke) was recorded from a number of beaches by Gemmill (1901) and Stephen (1929). In 1935, McGuire described a new species, O. cluthensis, from ](ames Bay, Millport, and in his detailed s tudy of the anatomy of the Kames Bay worms, Brown (1938) referred a number of the earlier records of O. limacina to this species. Tebble (1952) re- examined all the available British specimens of Ophelia and found that specimens from Kames Bay in the British Museum collection were in fact O. rathkei. Brown's description of the animals he studied does not agree with McGuire's of O. cluthensis, but rather with O. rathl~ei. There appear to be no type specimens extant of O. cluthensis and it may be concluded that McGuire's description of this species is actually a faulty account of O. rathlcei.

Since O. limacina appears not to occur in British waters (Tebble, 1952), the only authenticated record of Ophelia from the Clyde so far is of 0. rathlcei, from Karats Bay. The large population of Ophelia known from Stephen's investigation (in Brown, 1938) to exist in the intertidal zone of Fairlie Sands has now been re-examined and proves also to be of O. rath/cei.

Family CAPITELLIDAE Grube.

Capitella capitata (Fabricius). Fauvel, 1927, p. 154.

The only previous record of this species from the Clyde was of a single specimen collected many years ago in the intertidal muddy sand in Balloch Bay, Cumbrae (Clark, 1960). Large numbers have since been collected from the spoil grounds 2 miles north of Cloch Light. The substratum here is composed of sticky, black, polluted mud. C. capitata is widely distributed in British waters and usually occurs in muds containing large amounts of decaying organic matter.

? Mediomastus sp. A single, incomplete specimen collected from sublittoral deposits.

Details of time and place not recorded. I t differs from most genera and all known species, but because of its damaged condition, i t is impossible to be certain of some important characters, and for this reason is not named.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 649

The anterior fragment is 52 mm. long (contracted) and composed of 65 segments. I t is unpigmented in alcohol. The body formula is as follows: first segment achaetous, the next four segments with capillary chaetm in both rami, the following six segments with hooded hooks in both rami. These eleven segments constitute the thorax. The hooks are almost as long as the capillary chaetse, and have a single stout tooth surmounted by five, possibly more, smaller teeth arranged in a transverse series. The hood is somewhat inflated. Abdominal segments have only hooded hooks which have a smaller hood than those in the thoracic segments and they have a single large tooth surmounted by three finer teeth disposed in a triangle. The segments are biannulate. The first few abdominal segments are the same length as the thoracic segments. No branchim are visible and there are no copulatory chaetm.

Genera which have approximately the same body formula are Heteromastus Eisig, Capitomastus Eisig, Cclpitata Hartman and Mediomastus I4artman, but the Clyde specimen differs from all of them in some respects.

Heteromastus has twelve thoracic segments made up of 1 achaetous, 5 with capillary chaet~e, 6 with hooded hooks. In respects other than this, the present specimen resembles Heteromast~ts. The six posterior thoracic segments have hooks. There are no copulatory chaet~, and the worms have biannulate segments. Heteromastus possesses branchiee, but these commence on the 80th segment, and since the Clyde fragment includes only 65 segments, this is not a critical difference.

Male Capitomastus have 4 thoracic segments with capillary chaet~e, as for the Clyde worm, but only 5 succeeding thoracic segments with hooks, making a total of 10 thoracic segments. Both sexes of Capitomastus have copulatory chaetm and, furthermore, the worms consist of no more than 40-50 segments and reach a length of only 15-20 ram.

Capitata, a genus described from the Pacific coast of North America by t ta r tman (1947) has a similar segmental formula to the Clyde specimen, but there is a transition from thoracic to abdominal segments and the abdominal segments have a mixturo of capillary chaet~e and hooks, features which the Clyde specimen does not share.

Mediomastus is another genus described from the Pacific coast of North America by Har tman (1944 b). I t has the same segmental formula as the Clyde specimen and, like it, lacks copulatory chaet~e. The only respects in which the two described species of Mediomastus, M. californiensis Hartman (1944 b) and M. glabrus Hartman (1960), differ from the Clyde specimen are in size (the Clyde worm is larger) and in the length of the thoracic hooks, which are longer in the Clyde specimen. Furthermore, the segments of Mediomastus are uniannulate and not biannulate.

I-Iartman (1947) has shown convincingly that most capitellid genera can be separated on the basis of their segmental formulae. I f this principle is accepted, then the Clyde specimen is undoubtedly a species of Mediomastus, although the only other known species of the genus appear to be confined to the Pacific coast of North America between southern California and Oregon. Alternatively, it must be admitted that an aberrant individual

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

650 R . B . Clark & P. G. Dawson : polychaetes from Scotland

may depart from the normal segmental formula of the species following a disturbance of the normal growth processes. The Clyde specimen differs from Heteromastus only in that it possesses one less thoracic segment with capillary chaetse; and might possibly be an abnormal member of that genus. Because of the damaged state of the specimen, it is impossible to resolve this problem, but the use of the segmental formula has proved so valuable in capitellid taxonomy that it would be improper to assume that the specimen is aberrant in this respect without further evidence. I t is therefore referred tentatively to Mediomastus.

Family MALDANIDAE Malmgren. Clymenella cincta (Saint-Joseph).

Fauvel, 1927, p. 182.

A single incomplete fragment composed of the prostomium and eight anterior chaetigers, measuring 37 mm. long and 4 mm. in diameter, dredged in the Clyde area. Details of time and place are not recorded. Apart from the fact that the posterior rim of the cephalic plate is slightly crenate, the worm answers exactly to Fauvel's (1927) description which is rather more complete than that by Saint-Joseph (1894).

This appears to be the first rediscovery of the species since its original description by Saint-Joseph (1894) of an anterior fragment of 12 chaetigers collected at Dinard.

The genus Clymenella Verrill is a confused one. I t was vaguely defined by Verrill (1873, 1900) and subsequent attempts to give the definition greater precision have not received general acceptance. Fauvel (1927), characterized the genus by the presence of a collar on the anterior end of the fourth segment, which enfolds the end of the segment next anterior to it, although one form of C. rubrocincta lacks such a collar (Arwidsson, 1922; Monro, 1937; but see also Berkeley and Berkeley, 1941). This is not a critical species, however, and is sometimes included in other genera (Arwidsson, 1906; Berkeley and Berkeley, 1941; Hartman, 1959). Following Fauvel's definition of the genus, two groups of species are included, as Monro (1937) pointed out. These are C. cincta (Saint Joseph), C. minor Arwidsson and Clymenella sp. Monro (1937), which have acicular hooks in the neuropodium of the first three chaetigers, and C. torquata Verrill, C. somersi Verrill, C. elongata Moore (and possibly also C. rubrocincta Johnson and C. insecta Ehlers), which have typical uncini in the three anterior chaetigers, comparable to those of succeeding segments, in place of the acicular hooks. Various devices have been adopted to overcome this difficulty.

Arwidsson (1906, 1911) distributed the species between the following genera:

Axiothella (with glandular areas not arranged in bands, no pre-anal raised-ring), including somersi.

Clymenella (glandular areas in bands, pre-anal ring, collar segment), including torquata, elongata, minor (and presumably also cincta).

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 651

Euclymene (glandular areas in bands, pre-anal ring, no collar segment, first three chaetigers with acicular hooks), including insecta.

Unnamed genus (glandular areas in bands, pre-anal ring, no collar segment, first three chaetigers with uncini), including rubrocincta.

Monro (1937) regarded the collar on the fourth chaetiger, the raised pre- anal ring, and the form of the glandular areas as of little generic signifi- cance, and suggested that those species with neuropodial acicular hooks on the first three chaetigers should be included in Clymene (i.e. cincta and minor) and those with uncini (i.e. torquata, somersi, elongata, rubrocincta, insecta) in A'xiothella. This view has not been generally accepted,talhough Berkeley and Berkeley (1941) follow Monro in including rubrocincta in Axiothella, and Har tman (1945, 1959) has adopted something approaching the same solution to the problem.

t Iar tman (1945) retained Clymenella for torquata, but suggested that the other species should be distributed between Axiothella and Praxillella, a subgenus of Clymene in Fauvel 's (1927) and Monro's (1937) classification. Recently, Har tman (1959) has revised this disposition of the species, and Clymenella is now held to include torquata, cincta, tentaculata, minor and ? elongata; somersi and rubrocincta are referred to Axiothella; and insecta is referred to Euclymene. Even in this restricted sense, Clymenella includes species with acicula and uncini in the first three chaetigers, and it might be desirable in any future revision of the Maldanidae to separate cincta and minor (with acicula) from torquata, tentaculata and elongata (with uncini) into separate genera and in other respects revert to a system much closer to that proposed by Arwidsson (1906). Pending a revision of the iVIaldanidm, however, we refer the present species to Clymenella, in the inclusive sense, on the understanding that it is a candidate for re- allocation.

The only species of Clymenella previously known from British waters is C. torquata Leidy, an American species believed to have been introduced to the Whitstable area when Blue Point oysters were laid on the Whitstable Beds in 1936 (Newell, 1949 a, b). This species has so far not been reported from elsewhere on the British coast to our knowledge, though ff it can be transplanted from the American Atlantic coast to the European, there seems no reason why, with the traffic in oysters, it should not suffer a similar accidental transportation in Europe.

C. cincta can be readily distinguished from C. torquata by the following characteristics:

The neuropodia of the first three segments of C. cinta contain only stout acicular spines (1 in the first segment, 2 in the second, 1 in the third, in the Clyde specimen) whereas in C. torquata there are uncini (7-10 in the first segment, 5-7 in the second and third (Newell, 1949 b)) . The acicular spines of C. cincta are very obvious and can be distinguished under low-power magnification. This appears to be the readiest means of distinguishing the species.

The nuchal organs of C. torquata extend almost the whole length of the cephalic plate from near its anterior margin to the levet of the notches in

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

652 R . B . Clark & P. G. Dawson : polychaetes from Scotland

its border. In C. cincta, they are short and extend no more than one-third the length of the cephalic plate from its anterior margin.

The posterior margin of the cephalic lobe of C. torquata is distinctly crenate, but only slightly so in C. cincta.

The notches in the cephalic border are pronounced in C. torquata and are placed near its posterior end; in C. cincta they are slight and occur about half way along the side of the border.

Praxillura longissima Arwidsson. Arwidsson, 1906, p. 27.

Eight anterior fragments of this rather rare species have been collected from sub-littoral deposits around the Isle of Cumbrae. I t has probably been collected once before, for Arwidsson (1922) recorded finding a single fragment of "? Praxillura sp." from the shore of Cumbrae. The present specimens are 79 mm. (19 chaetigers), 42 mm. (22), 39 mm. (23), 18 mm. (13), 16 ram. (13), 12 ram. (11) and 8 mm. (12) in length. The eighth specimen is also 8 mm. long but has an indeterminate number of segments.

The species was described in exemplary detail by Arwidsson (1906) and supplementary accounts of it have been given by Wesenberg-Lund (1948, 1950 a) and Eliason (1962). The prostomium lacks the dorsal rim and mid-dorsal ridge that are features of many other maldanids, and bears a pair of prominent crescent shaped nuchal organs near its anterior margin. The mouth is surrounded by a well marked lip and inside it are two longitudinal ridges which bear short papillm, giving them a crenate appearance. The first segment is achaetous and is almost completely fused with the prostomium. The most anterior chaetigerous segments are about as long as broad, but further back along the body they become elongated. The increase in relative length of the segments becomes noticeable by about the 10th to 15th chaetiger. A number of anterior segments (between 4 and 14, according to Eliason (1962)) bear a single, stout acicular spine in the neuropodium; very occasionally there are two such spines in a neuropodium. All other neuropodia bear a transverse row of a small number of uncini (up to 11) with 4-6 teeth at the tip and a single barbule or hair arising from beneath the lowest and largest tooth. All notopodia bear capillary chaet~e.

Eliason (1962), who has had the opportunity of examining a greater number of specimens of this species than other writers, points out tha t the species is more variable than was previously supposed and this view is supported by the condition of the Clyde specimens, no two of which are identical. The number of anterior chaetigers bearing acicular spines in the neuropodia varies from 6 to 11, and in three of the seven worms, the most posterior of the segments bearing spines is asymmetrical, with a spine on one side and uncini on the other. There is generally only a single spine in a neuropodium, though occasionally a second, smaller spine flanks the first. Except in the more anterior segments, the neuropodial chaetm are born on tori; the segment on which these first become prominent

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 653

varies between the 10th and 14th ch~etigers in the Clyde specimens. The chaet~e and glandular patches occur in the anterior third of the segment in the anterior region of the body, but further back they shift to the posterior part of the segments. According to Arwidsson (1906) this backward shift of the tori occurs between chaetigers 16 and 17, but in one Clyde specimen the shift occurs between chaetigers 18 and 19, and in another, the backward shift is not apparent although the fragment is 22 chaetigers long. The indistinctness of the segmental boundaries does not permit the determination of the level of the change in position of the tori in a third specimen which is more than 17 segments long. There is no evidence of a backward shift of the tori in any of the other, shorter fragments. All anterior segments have one or two dorsal, transverse bands of brownish pigmentation, but the great variability of pigmentation of the worms reported by Arwidsson (1906) and Eliason (1962) suggests that this is not a useful character.

One of the Clyde specimens 8 mm. long, differs from the others in a puzzling manner. The prostomium and first, achaetous segment are normal. They are followed probably by four small biannulate segments, narrower than the first segment and only one-third as long as broad. These appear to be achaetous, although one segment may have a single capillary chaeta. The next three segments are somewhat longer than broad and bear an acicular spine in each neuropodium. The remaining five segments of the fragment have uncini in the neuropodia. Apart from the four small and apparently achaetous segments, this fragment resembles typical P. longissima. Possibly it was in the process of regenerating when collected, bu t if so, it is most unusual that the prostomium and first segment should have been fully differentiated and have reached full size while the succeed- segments are so dimunitive.

The great variability of P. longissima remarked by Eliason (1962) and shown by the Clyde specimens, makes it likely that other species which have been described are identical with it, particularly as they are known only from their original discovery and have been described in inadequate detail. Thus, P. ornata Verrill (1879) from the Massachusetts coast is Mmost identical with P. longissima (Eliason, 1962); ? Nichomachella picta Ehlers (1887) dredged south of Florida and certainly a Praxillura (Arwidsson, 1906; Har tman, 1959), also appears to be indistinguishable apart from its bright yellow eolour. P. maculata Moore (1923) taken off Santa Cruz Island, California, is known from a single incomplete specimen and this, too, appears indistinguishable from P. longissima.

Although comparatively rare, indisputable P. longissima has a wide distribution in the arctic, having been recorded from the east and west coasts of Greenland, north of the Faeroes, off Spitzbergen, Jan Meyen and Novaya Zemlya and from the Kara Sea (Arwidsson, 1906; lVauvel, 1911 ; Wesenberg-Lund, 1948, 1950 a, b, 1953) and penetrates into the north Atlantic as far south as Nova Scotia (Fauve], 1932), off the coasts of Iceland (Wesenberg-Lund, 1951), in the Kat tegat and Skaggerak (Eliason, 1920, 1962), as well as in the Clyde.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

654 R . B . Clark & P. G. Dawson : polychaetes from Scotland

Family TEREBELLIDAE Malmgren.

Axionice maculata (Dalyell). Fauvel, 1927, p. 263 as Pista maculata.

A single, incomplete specimen was collected in Etterick Bay, Bute. This species has been reported from Clare Island (Southern, 1914) and

the Irish Sea (Southward, 1956), but not previously from the Clyde. I t is evidently rare in British waters.

LysiUa loveni Malmgren. Fauvel, 1927, p. 286.

Five specimens have been collected, along With Eupolymnia nesidensis (delle Chiaje) from nests of the lamellibranch Lima hyans on the Skelmorlie Bank. The depth here is about 15 m. and the bot tom is composed very largely of shell gravel. The size range of this species is quoted by Fauvel (1927) as 30-60 mm. long and 5-6 mm. in diameter. The Clyde specimens are all small, 10-15 mm. by 1-1.5 mm.

L. loveni has previously been recorded from St. Magnus Bay, Shetland (McIntosh, 1922) and Port Erin, Isle of Man (Moore, 1937). Elsewhere it appears to have a wide distribution in the north Atlantic and Arctic (l~auvel, 1927 ).

Family SABELLII)AE Malmgren. Chone infundibuliformis Kroyer.

Fauvel, 1927, p. 334.

This species was previously known from a single specimen taken in Etterick Bay, Bute, at a depth of 37 m. (Clark, 1960). Six specimens have now been dredged in the Fairlie Channel.

SUMMARY.

Seventeen species of polychaete worm hitherto unknown from the Fir th of Clyde and the west coast of Scotland, together with a number of others, are reported upon. Three of these have not previously been recorded in British waters: Harmothogjoubini Fauvel, Clymenella cincta (Saint-Joseph) and undetermined capitellid probably belonging to the genus Mediomastus. Both Harmotho~ joubini and Clymenella cincta have been rediscovered for the first time since their original description. Only two species of .Mediomastus are known, both from the Pacific coast of the United States. The Clyde specimen does not appear to belong to either, but its damaged condition makes precise determination impossible, and the possibility tha t it is an abnormal Heteromastus, though unlikely, is not excluded. Pholo~ synophthalmica Clapar~de is another species new to British waters, bu t the Clyde specimens are in some respects intermediate between this species and P. minuta (Fabricius). In view of the known variability of the latter, it is probably preferable to regard P. synophthalmica as a

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R. B. Clark & P. G. Dawson : polychaetes from Scotland 655

variant of P. minuta. Similarly, specimens intermediate between Aonides oxycephala (Sars) and A. paucibranchiata Southern have been found, which cast doubt on the validity of the latter species. Praxillura Iongissima previously known only from a generic record of a single fragment collected in the Clyde, has now been positively identified. I t has not been reported elsewhere in British waters.

R E F F E R E N C E S

ARDWIDSSON, A. 1906. S tud ien fiber die s k a n d i n a v i s c h c n u n d a rk t i s chen Ma ldan iden Zool. Jb. suppl . 9, 1-308•

---. 1911. Die Maldan iden . Wiss. Ergeb. schwed. Sudpolar-Exped., 1901-1903 6 (6), 1-44. 1922. Sys t ema t i c no te s on s o m e ma ldan ids . K. svensk. Vet.-Akad. Handl. 63 (7),

1-46. ASHWORTR, J . H . 1901. The a n a t o m y of Scalibregma inflatum R a t h k e . Quart. J. micr.

Sci. 45, 237-309. 1909. P o l y c h a e t a of t h e coas t s of I re land. I. Arenicol idae a n d Scal ibregmidae .

Fisheries, lreland, Sci. lnvest. {1908) 7, 1-4. AZlZ, N . D . 1938. F a u n a of K a r a c h i . 2. Po lychae tes . ~Iem. Dept. Zool., Univ. Punjab

1, 19-25. BASSINDALE, R . a n d BARRETT, J . H . 1957• The Dale Fo r t ma r ine fauna.. Proc. Bristol

Nat. Sac. 29, 227-328. BERKELEY, E. a n d BERKELEY, C. 1941. O n a collection of Po lychae t a f r o m s o u t h e r n

California. Bull. S. Calif. A cad. Sci. 40, 16-60. BROWN, R . S . 1938. T h e a n a t o m y of the po lychae te Ophelia cluthensis McGuire, 1935.

Proc• Roy. Sac. Edinb. 58, 135-160. CHUMLEY, J . 1918. The fauna of the Clyde Sea Area. {Glasgow: Un iv . Press) . CLAPAR~DE, E . 1868. Les Ann61ides Ch~topodes d u Golfe de Naples . Mere. Sac• Phys.,

Geneva 19, 313-584. CLARK, R . B . 1952. New records of sub- l i t tora l po lychae tes f rom the Clyde Sea Area,

wi th a descr ip t ion of a n e w species. Proe. Roy. Sac. Edinb., B 65, 1-26. 1954. Pelagic s w a r m i n g of Scal ibregmidae (Polychaeta) . Ann. Rapt8 Scot. mar.

biol. Assoc. (for 1952-3), 20-22. 1960. The fauna of the Clyde Sea Area. Polychaeta, with keys to the British genera.

(Millport: Scot. mar . biol. Assoc.) . - - . ALDER, J . R. a n d MCI~TYRE, A . D . 1962. T h e d i s t r ibu t ion o f Nephtys a r o u n d tho

coast of Scot land. J. anita. Ecol. 31, 359-372. • a n d HADERLIE, E . C . 1960. T h e d i s t r ibu t ion of 1Vephtys cirrosa a n d Nephtys hombergi

on t h e s o u t h - w e s t e r n coas t of E n g l a n d a n d Wales . J. anita. Ecol. 29, 117-137. - - . a n d MILLE, A. 1955. The sub- l i t tora l f a u n a of two s a n d y bays on the Isle of Cumbrae ,

F i r th of Clyde. J . mar• biol. Assoc. U.K. 34, 161-180. Cu~rt~OHAM, J . T. a n d RAMAQE, G• A. 1888. Po lychae t a s eden ta r i a of the F i r t h of Fo r th .

Trans. Roy. Sac. Edinb. ~ 635-684. DrrLEVSEN, H . 1911. Anne l ids f rom t h e D a n m a r k - E x p e d i t i o n . Medd• Gronl. 45, 411-432. E H ~ R S , E. 1887. F lor ida-Annel iden . R e p o r t on the a n n e l i d s of the dredging expedi t ion

of the U.S. coas t s u r v e y s t e a m e r " Blake ". ~lem. 2~lus. camp. Zool. Harvard 15, 1-335.

ELIASO~,A. 1920. : B i o l o g i s c h - f a u n i s t i s c h e U n t e r s u c h u n g e n a u s d e m 0 r e s u n d . Po lychae ta . Lund Univ. Aarskr. ii, 16 (6), 1-103.

• 1962. U n d e r s S k n i n g a r 5ver 0 r e s u n d X X X X I . Wei te re U n t e r s u e h u n g e n tiber die P o l y c h a e t e n - f a u n a des 0 r e s u n d s . K.fysiogr. Sallsk. Handl. N.F. , 73 (9), 1-98.

FAUV~L, P. 1911. Ann$1ides Polych~tes . In : Duc d'Orlgans campagne arct~tue de 1907. Pp . 1-45.

• 1914. A n n ~ l i d e s P o l y c h ~ t e s n o n - p 6 1 a g i q u e s p r o v e n a n t d e s c a m p a g n e s d e l ' H i r o n d e U o et de la Pr incesse-Al ice (1885-1910). Bes. Camp. Sei..~/ionaco 46, 1-432. 1923. Po lyeh~tes e r ran tes . Fauna de France 5.

. 1927. Po lych~tes s~dentaires . Faune de France 16. • 1932. A n n ~ l i d e s P o l y c h ~ t e s p r o v e n a n t d e s c a m p a g n e s d e l ' H i r o n d e l l e I I ( 1 9 1 1 - 1 9 1 5 ) .

Res. Camp. sci. Monaco 85, 1-50•

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

656 R . B . Clark & P. G. Dawson : polychaetes f rom Scot land

GEMMILL, J . F . 1901. Mar ine worms . In : El l iot t , G. F . S., Laur ie , M. a n d Murdoeh, J . B. (eds.) Fauna, flora and geology of the Clyde area. (Glasgow: Local c o m m i t t e e for m e e t i n g of Br i t . Assoc.). Pp . 359-363.

GRUBE, A . E . 1860. Beschre ibung neuer oder wenig b e k a n n t e r Annel iden . 4. Be i t r ag : Zahlre iche G a t t u n g e n . Arch. Naturg. 26, 71-118,

HXRTMA~, O. 1944 a. N e w E n g l a n d Annel ida . P a r t 2• Bull. Amer. Mus. nat. Hist . 87, 327-344. 1944 b. P o l y e h a e t o u s anne l ids f rom California, inc luding the descr ip t ions of two

n e w genera a n d n ine n e w species. Allan Hancock Pacif. Exped. 10, 239-307. 1945. Marine anne l ids of N o r t h Carolina. Bull. Duke Univ. mar. Sta. 2, 1-54. 1947. P o l y c h a e t o u s annel ids . P a r t V I I . Capitell idae. Allan Hancock Pacif.

Exped. 10, 391-449. 1959. Cata logue o f t h e po lychae tous anne l ids of the world. Occ. Pap. Allan

Hancock Edn 23, 1-628. 1960. T h e ben th ic f a u n a of the deep bas ins off S o u t h e r n California. P a r t 2. Allan Hancock Pacif. Exped. 22, 69-215.

JOHNSTON, G. 1839. Miscel lanea zoologica. The Br i t i sh Aphrodi tacea . Ann. Mag. nat. Hist. i, 2, 424-441. 1865. A catalogue of British non-parasitieal worms in the collection of the British

Museum. (London: Trus tees , Bri t . Mus.) . MARINE BIOLOGICAL ASSOCIATIObI'. 1957. Plymouth marine fauna (3rd edit.). M~LRIO~, A. F. a n d BOBRETZKY, N. 1875. E t u d e des Ann61ides du golfe de Marseille.

Ann. Sci. nat. Zool. vi, 2, 1-106. McGUIRE, I . P . 1935. Note on a n e w species of po lychae te (Ophelia cluthensis). Scot•

Nat. 212, 45-46. M c I ~ r o s ~ , W . C . 1869. O n t h e s t r u c t u r e of t h e Br i t i sh n e m e r t e a n s , a n d some new Br i t i sh

annel ids . Trans. Roy• Soc. Edinb. 25, 305-433. 1900. British marine annclids• (London : R a y See.) 1 (2).

~ . 1910. British marine annelids. (London: R a y See.) 2 (2). • 1915• British marine annelids. (London: R a y See.) 3 (1). • 1922. British marine annelids. (London: R a y See.) 4 (1).

- - . 1924. Notes f rom t h e G a t t y m a r i n e l abo ra to ry , No. 46. Ann. Mag. nat. Hist. ix, 14, 1-52.

MESNIL, F . 1896. ]~tudes de morphologie e x t e r n e ehez les Ann61ides. Les Spionidiens des cStes de la Manehe. Bull. Sci. Er. Belg. 29, 110-287.

MICHAELSEN, W. 1897. Die P o l y e h a o t e n f a u n a tier d e u t s e h e n Meer, e inschl iessl ich de r b e n a e h b a r t e n u n d v e r b i n d e n e n Gebiete. Wiss. Meere~untcrs. dtsch. Metre, N.F. , 2 (1), 1-216.

MONRO, C. C . A . 1937. Po lychae ta . Sci. Repts John Murray Exped., 1933-34, 4, 243-321. MOORE, H . B . 1937. Mar ine f a u n a of t h e Isle o f M a n . Prec. L'pool biol. See. 50, 1-293. MOORE, J . P . 1923. The p o l y e h a e t o u s anne l ids d r e d g ed b y t he U.S.S. " Alba t ross " off

coas t of s o u t h e r n California in 1904. IV. Sp ion idae to Sabellari idae. Prec. Philad. Acad. nat. Sci. 75, 179-259.

NEWELL, G . E . 1949 a. T h e occurrence of a species o f Clymenella Verrill (Po lyehae ta , Maldanidae) on t h e n o r t h K e n t coast . Nature, Lend. 168, 648. 1949 b. Clymenella torquata (Leidy), a p o l y c h ae t e new to Br i ta in . Ann. Mag.

nat. Hist. xii, 2, 147-155. PETTIBONE, lV[• H . 1953. Some scale bearing polychaetes of Puget Sound and adjacent

waters. (Seatt le: Un iv . W a s h i n g t o n Press) . - - . 1956. Mar ine po lyehae te w o r m s f r o m L a b r a d o r . Prec. U.S. nat. Mus. 105, 531-

584. • 1963. Mar ine po lyehae te w o r m s of t h e N e w E n g l a n d region. P a r t 1. Aphrod i t idae

t h r o u g h Trochae t idae . Bull. U.S. nat. Mus . 227, 1-356. RIoJA, E . 1919. Adiciones a la f a u n a de An61idos del Cantabr ieo . Rev. Acad. Cien.

Madrid 17, 54-79. SAINT-JOSEPh, A. de. 1887. Anndl ides Po lychb tes des e6tes de Dinard . Ann. Sei. nat.

Zool. vii, 1, 127-270. • 1894, Ann61ides Po lyeh~tes des cStes do D i n a r d . Trois i~me Pa t t i e . Ann. Sei. nat.

Zool. vii, 17, 1-395. S~DERSTR6M, A. 1920. Studien i~ber die Polychaetenfamilie Spionidae. (Uppsa la : I n a u g .

diss.). STeP-BowITZ, C. 1941. Les Glycdriens de Norvbge. Nytt Mag. Naturv. 82, 181-250.

• 1945. Les Soalibregmiens de Norvbge. !Vyt$ Mag. Naturv. 85, 63-87. • 1948. Sur les Polych~tes arctiques. Tromse -~lus. Aars. 66 (2), 1-58.

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4

R . B . C l a r k & P . G. D a w s o n : p o l y c h a e t e s f r o m S c o t l a n d 657

SOUTHERLY, R. 1914. Archiannelida and Polychaeta. Clare Island survey. Prec. Roy. l r~h Acad. 81 (47}, 1-160.

SOUTI~WARD, E . C . 1956. On some Polyehaeta of the Isle of Man. Ann. Mag. nat. Hist. xii, 9, 257-279.

STEPHEN, A.C. 1929. Studies on the Scottish marine fauna: the fauna of sandy and muddy areas of the tidal zone. Trans. Roy. Soc. Edinb. 56, 291-306.

TEBBLE, N. 1952. On three ~pecies of the genus Ophelia (Polychaeta) from British and adjacent waters. Ann. Mag. nat. Hist. xii, 5, 553-571.

VERRILL, A° E. 1873. Report on the invertebrate animals of Vineyard Sound and adjacent waters, with an account of the physical conditions of the region. Rep. U.S. Comm. Fish., (1871-2). 295-778. 1879. Notice of recent additions to the marine Invertebrata of the northeastern

coast of America, with descriptions of new genera and species and critical remarks on others. Prec. U.S. nat. Mus. 2~ 165-205. 1900. Turbellaria, Nemertinea and Annelida of the Bermudas. Trans. Corm• Acad.

• Arts Sci. 10, 573-610. WESEZ~BERG-LuIqD, E. 1948• Maldanidae (Polyehaeta) from west Greenland waters. Medd.

Oranl. 184 (9), 1-58. 1950 a. Polychaeta. Danish Ingolf Exped. 4 (14), 1-92. 1950 b. The Polyehaeta of west Greenland with special reference to the fauna of

Norde Stromfjord, Kvane and Bredefjord. Medd. Gronl. 151 (2), 1-171. 1951. Polychaeta. Zoology of Iceland 2 (19), 1-182.

. . . . . 1953. The zoology of east Greenland. Polyehaeta. Medd. Grovel. 122 (3), 1-169.

A . M . N . H . se r . 13, v o l . vi . 42

Dow

nloa

ded

by [

Uni

vers

ity o

f A

rizo

na]

at 0

6:37

27

June

201

4