Chaetozone carpenteri McIntosh, 1911 from the Mediterranean Sea and records of other bi-tentaculate Cirratulids

Abstract

Many species of Cirratulidae have been recorded from the Mediterranean Sea since the descriptions and figures of the Naples fauna in one of the first comprehensive studies of polychaetes by delle Chiaje, 1823–30. This original publication included only multi-tentaculate cirratulids and what we would now identify as Cirriformia and Cirratulus. Since delle Chiaje's publication, 25 bi-tentaculate taxa have been recorded from the Mediterranean. During recent sampling programmes Chaetozone carpenteri McIntosh, 1911, a bi-tentaculate cirratulid, has been recorded from several Mediterranean sites and is redescribed. All records of C. setosa will need to be re-examined as they may have been misidentified.

Keywords:

• Cirratulidae
• Mediterranean
• re-description
• Chaetozone carpenteri

Introduction

Since delle Chiaje, 1823–30, one bi-tentaculate genus, Monticellina Laubier, 1961 and several new species, e.g. Caulleriella bioculata (Keferstein, 1862), Aphelochaeta multibranchis (Grube, 1863) and Chaetozone carpenteri McIntosh, 1911, have been described from the Mediterranean.

Various benthic surveys in the Mediterranean have identified bi-tentaculate cirratulids in their samples: Spain (Rioja 1928; Ariño 1987), France (Laubier 1960, 1962b; Bellan 1964), the Adriatic (Katzmann 1973a, b), the Gulf of Naples (Claparède 1868), southern Italy (Gambi et al. 1986), the check list of Italy (Fabio 2003), off the coast of North Africa: Algiers (McIntosh 1911), Morroco (Fauvel 1936; LeChapt, 1994), Tunisian coast (Zaâbi et al. 2009; Ayari et al. 2009; Zaâbi, personal communication), Aegean Sea (Arvanitidis 2000; Simboura & Nicolaidou 2001), Turkey (Çinar et al. 2004; Çinar & Ergen 2007) and Egypt (Abd-Elnaby 2009). Most of these publications are faunal lists and do not contain detailed descriptions or illustrations of the characters required for confident identification. The lists include designations such as Chaetozone sp., which may encompass more than one species. Often, authors have applied names published from areas outside the Mediterranean, e.g. Chaetozone caputesocis (St Joseph, 1894) which was based on samples from Dinard, northern coast of France, Caulleriella alata (Southern, 1914) from the west coast of Ireland and Monticellina dorsobranchialis (Kirkegaard, 1959) which was based on samples from the coast of West Africa (Table I).

Table I. Records of bitentaculate cirratulids from the Mediterranean

Mediterranean historical records with author:	Adriatic sea samples	Tyrrhenian Sea samples
Chaetozone	Chaetozone	Chaetozone
C. carpenteri McIntosh 1911	C. carpenteri	C. carpenteri
Ariño 1987; Fauna Iberica
C. corona Berkeley & Berkeley 1941
Çinar & Ergan 2007
C. setosa Malmgren, 1870
McIntosh 1911; Laubier et al. 1962a, 1962b; Vatova, 1931, 1935, 1949; Fauvel 1934, 1936, 1940; Kiseleva 1964; Gamulin-Brida et al. 1968; Bellan 1964, 1969; Banse 1970; Zavodnik 1971, 1979; Katzmann 1972, 1973a, 1973b, 1983; Amoureaux 1976, 1979,1983a; Zavodnik et al. 1978, 1985, 1987; Požar-Domac 1978, 1983, 1994; Gambi et al. 1983; Aleffi et al. 1987–1988, 2003; Zavodnik & Vidaković 1987; Županović & Jardas 1989; Zahtila 1990; Rebzani-Zahaf 1992; Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003; Solis-Weiss et al. 2004; Ayari et al. 2009
Chaetozone cf setosa
Ayari et al. 2009
C. ‘A’ McIntosh 1911
C. gibber Woodham & Chambers, 1994
Çinar et al. 2004; Zaabi et al. 2009
Chaetozone sp.
Gambi & Giangrande 1986; Simboura & Nicolaidou 2001; Çinar 2005	Chaetozone sp.?	Chaetozone sp.?
Caulleriella	Caulleriella	Caulleriella
C. alata (Southern, 1914)
Gambi et al. 1983; Gambi & Giangrande 1986; Giangrande & Gambi 1986; Arvanitides 2000; Simboura & Nicolaidou 2001 Fabio 2003; Çinar 2005; Ayari et al. 2009; Solis-Weiss et al. 2004
C. bioculata (Keferstein, 1862)
Banse 1959, 1970; Amoureux & Katzmann 1971; Katzmann 1971, 1972, 1973a, 1983; Amoureux 1975; Velimirov & Sint 1975; Požar-Domac 1978, 1994; Gambi et al. 1983; Hartmann-Schröder 1996; Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003; Aleffi et al. 2003; Çinar 2005; Çinar & Ergen 2007; Ayari et al. 2009; Abd-Elnaby 2009; Zaâbi et al. 2009.
C. caputesocis (Saint-Joseph, 1894)
Bellan 1964, 1969, 1976; Zavodnik 1971, 1979; Katzmann 1972; Požar-Domac 1978, 1983, 1986, 1994; Fassari 1983; Gambi & Giangrande 1986; Giangrande & Gambi 1986; Županović & Jardas 1989; Hartmann-Schroder 1996, Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003
C. cf caputesocis
Katzmann 1973a, 1983; Zaâbi et al. 2009.
Caulleriella sp. Çinar 2005
C. cf alata Çinar & Ergen 2007; Zaâbi et al. 2009
C. sp1 Çinar & Ergen 2007
C. sp2 Çinar & Ergen 2007
C. zetlandica (McIntosh, 1911)
Bellan 1964; Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003; Abd-Elnaby 2009	?C. cf zetlandica	?C. cf zetlandica
Monticellina	Monticellina	Monticellina
M. heterochaeta Laubier 1961
Gambi et al. 1983; Gambi & Giangrande 1986; Ambrogi et al. 1987; Zavodnik & Vidaković, 1987; Zahtila 1990; Požar-Domac 1994; Arvanitides 2000; Çinar 2005
M. dorsobranchialis (Kirkegaard, 1959)
Arvanitides 2000; Simboura & Nicolaidou 2001; Aleffi et al. 2003; Fabio 2003.
M. cf. tesselata (Hartman, 1960)
Çinar 2005	? M. heterochaeta	? M. heterochaeta
Tharyx	Tharyx sp?	Tharyx sp?
T. killariensis (Southern, 1914)
Požar-Domac 1978, 1994; Gambi et al. 1983; Simboura & Nicolaidou 2001; Fabio 2003
C. cf .killariensis
Katzmann 1973a, 1983; Zaâbi et al. 2009
Tharyx retierei LeChapt 1994
Aphelochaeta	Aphelochaeta sp?	Aphelochaeta sp?
Aphelochaeta filiformis (Keferstein, 1862)
vatova 1942; Ayari et al. 2009
A. monilaris (Hartman, 1960)
Simboura & Nicolaidou 2001
A. multibranchis (Grube 1863)
Stossich 1883; Carus 1884; Graefe 1905; Fauvel 1936; Vatova 1942; Laubier 1962b; Bellan 1964; Marcuzzi 1972; Katzmann 1973a, 1973b, 1983; Požar-Domac 1978; Gambi et al. 1983; Gambi & Giangrande 1986; Giangrande & Gambi 1986; Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003
A. marioni (Saint Joseph, 1894)
Fauvel 1936; Bellan 1964; Zavodnik 1971, 1979; Katzmann 1972, 1973a, 1973b, 1983; Zavodnik et al. 1978; Amoureux 1979, 1983a, 1983b; Gambi et al. 1983; Gambi & Giangrande 1986; Gillet 1985–1986; Požar-Domac 1986, 1994; Zahtila 1990; Arvanitides 2000; Simboura & Nicolaidou 2001; Fabio 2003; Aleffi et al. 2003; Abd-Elnaby 2009; Ayari et al. 2009; Zaâbi, personal communication
A. cf marioni
Zavodnik & Vidaković 1987

During recent benthic sampling in the central area of the Mediterranean off the coasts of Italy and Croatia, cirratulids were found in most samples. Many taxa were similar to species described from the North Atlantic, e.g. Caulleriella zetlandica McIntosh, 1911 and Chaetozone gibber Woodham and Chambers, 1994, but there were doubts about the application of these names as the Mediterranean environments are significantly different, e.g. temperature and tidal range. Closer examination of several specimens of Chaetozone revealed characters that were not similar to the North Atlantic material, e.g. the position of very large spines from about chaetiger 6. This species was identified as Chaetozone carpenteri and is re-described.

Materials and methods

In the North Adriatic, the Ruđer Bošković Institute, Rovinj, sampling programme employed the research vessel ‘Vila Velebita’ to record the long-term changes of the soft bottom communities. There were three offshore monitoring stations located on the map (Figure 1): (1) SJ 005 (45°18.4′N 13°18.0′E), 31 m, characterised by a muddy detritic bottom, silty sand; (2) SJ 007 (45°17.0′N 13°16.0′E), 31 m, transitional community of detritic bottom enriched with silt, silty sand; (3) SJ 107 (45°02.8′N 13°19.0′E), 37 m, characterised by a coastal detritic bottom, silty sand.

Figure 1. Map of sampling areas.

In the Tyrrhenian Sea, the Italian Institute for Environmental Protection and Research (ISPRA) sampling programme employed the research vessel ‘Astrea’. This was primarily for the environmental monitoring of sub-marine electrical pipelines between Sardinia (Fiume Santo) and the mainland, North West Sardinia, (40°53.87′N–8°32.78′E), 42 m, sandy bottom. Another locality was Penisola Sorrentina near Naples (40°39.03′N 14°23.50′E), 80 m, sandy mud. The samples from Procida Island, Gulf of Naples (40°44.52′N 14°01.30′E), 98 m, muddy bottom, were obtained by a commercial company, Bioservice, as part of a monitoring exercise to examine the impact of Tuna (Thunnus thynnus) harvesting cages. Sediment characteristics are reported for Adriatic and Tyrrhenian stations from which Chaetozone carpenteri material was examined.

Sediment characteristics are reported for Adriatic and Tyrrhenian stations from which Chaetozone carpenteri material was examined (Table II). All samples were collected using a 0.1 m² van Veen grab and samples were sieved with a 1 mm mesh and fixed in 4% formalin and transferred to 74% ethanol. Specimens were examined using a Wild M7, Heerbrugg and Leica optical microscopes. All drawings were prepared with the aid of a camera lucida.

Table II. Adriatic and Tyrrhenian location details

	Adriatic SJ 005	Adriatic SJ 007	Adriatic SJ 107	North west Sardinia	Procida Island, near Naples	Penisola Sorrentina near Naples
Geographic	45°18.4′N	45°17.0′N	45°02.8′N	40°53.87′N	40°44.52′N	40°39.039′N
position	13°18.0′E	13°16.0′E	13°19.0′E	8°32.78′E	14°01.30′E	14°23.50′E
Depth (m)	31	31	37	42	98	80
Sediment type	Silty sand	Silty sand	Silty sand	Silty sand	Mud	Sandy mud

Specimens deposited at the National Museums of Scotland as NMSZ: 2010.66.67 and 68.

Taxonomic account

Chaetozone carpenteri McIntosh, 1911

(Figure 2)

Figure 2. Chaetozone carpenteri. A, Whole animal dorsal view, from Procida, 2007. NMSZ. 2010.068. B, Anterior region, lateral view, NMSZ. 2010.066. C, Anterior region 11 chaetigers, dorsal view, Rovinj, 2003. NMSZ.2010.066. D, Posterior region, last 6 chaetigers. Rovinj, 2003. NMSZ.2010.066. E, Chaetiger 24th–25th middle region in TS, Rovinj, 2003. NMSZ. 2010.066. F, Chaetiger posterior region in TS, Rovinj, 2003. NMSZ 2010.066. G, Chaetae: (a) fine short capillary, (b) posterior spine, (c) anterior spine, (d) capillary long, Rovinj, 2003. NMSZ 2010.068.

Chaetozone carpenteri McIntosh, 1911: 166, Pl. 6, figs 5, 5c–e.

Type material examined

Spain, off Cape Finisterre, Porcupine Expedition 1870, syntype NHM 1921.5.1.3234. It is in two parts, the anterior fragment measures 10 mm for 30 chaetigers and the posterior fragment measures 5 mm for 25 chaetigers. The other syntype from Spain off Cape Guardia, NHM 1921.5.1.3233, is a middle region of about 5 mm and 1 mm for 20 chaetigers which has no prostomium, peristomium or branchiae and is considered unidentifiable.

Material examined

Tyrrhenian Sea, Procida Island, BP6, 2007 Reg: NMSZ.2010.068, 1 specimen of 50 chaetigers, measuring 14 × 2 mm; 1 specimen of 60 chaetigers, measuring 23 × 2 mm; Sardinia, Punta Tramontana, PT7_C3, 2008, 1 specimen of 55 chaetigiers measuring 13 × 2 mm; Croatia, Rovinj, SJ 107 12.08.03 Reg: NMSZ.2010.066, 1 specimen of 55 chaetigers measuring 15 × 2 mm; SJ 007 07.06.04 Reg: NMSZ.2010.067 1 specimen of 55 chaetigers measuring 11 × 2 mm.

Description

Most of the following description is based on the specimens from Rovinj and Procida Island.

Maximum body length 23 mm for 60 chaetigers. Body surface smooth, iridescent and more or less the same width from the anterior to the posterior region. The anterior dorsal surface is slightly rounded and the ventral surface flattened with a longitudinal groove. Posterior segments are concertina-like. The prostomium is conical with a pair of eyes but no nuchal grooves. Peristomium achaetous, with a dorsal raised area posterior to the eyes; ventral mouth and a pair of grooved palps (only palp bases present) originating from the dorsal surface posterior to raised area. Branchiae present from the first chaetiger on the dorsal surface of the outer edge of the chaetae, between the parapodial folds (most of the branchiae are lost during sampling or preservation; consequently, a detailed description of their distribution along the length of the body is not possible). Parapodia all biramous with notopodial and neuropodial lobes, very thin and joined in the mid-lateral line but clearly visible. Chaetae arranged in fan-shaped rows.

Chaetae of four types:

1.	awl-shaped capillaries, 8–10 in anterior notopodia, 4–6 in neuropodia, dorsal chaetae longer in middle body region, up to 6 mm;
2.	very large spines, up to 0.5 mm long, 2–3 in notopodia and neuropodia, from anterior segments, around the 6th–9th segment to the middle body region;
3.	long spines up to 4 mm, from approximately the 20th segment, 3 in each notopodia and neuropodia increasing to 5–6 in posterior segments;
4.	fine, short capillaries in posterior segments between the long spines. Pygidium: small, flat, rounded lobe.

Remarks

In a confusing publication of various cruises, samples and figures of Chaetozone carpenteri McIntosh, 1911 was described from three Porcupine Expedition stations sampled during 1870. They were (1) Bono Bay, on the coast of Algiers, in 25 fathoms (45 m), (2) off Cape Guardia and (3) Cape Finisterre, Atlantic coast of Spain. The specimen from Bono Bay (NHM.1921.5.1.3232), Algeria has been lost, but McIntosh states that the specimens off Spain had the same characteristic features. The Bono Bay specimen is the only sample from the Mediterranean and the other two specimens are in poor condition. On closer examination of the Cape Finisterre specimen, obvious spines from the 7th/8th segment were observed which is in agreement with the samples from the Tyrrhenian Sea and Adriatic coasts.

The spines are clearly visible in McIntosh's figure but his description of the spine (=crotchet) distribution varies when compared to the figures. McIntosh did not mention spines in anterior segments but the type material is in very poor condition with most chaetae damaged. The lack of complete chaetae means an accurate interpretation of which segment has the first spine is impossible. In his figures (5c and e), McIntosh illustrates flat, striated spines which is correct when compared to the type material. The figure of the tenth foot (5b) is illustrated with 4 spines in notopodia and neuropodia, but the description does not mention a specific number; instead, there is a statement that they have attained great length. The exact number of spines depends on how you measure and interpret a spine as many chaetae are damaged. In the samples from Croatia and Italy there are 2–3 spines in anterior notopodia and neuropodia. The McIntosh figures do not include a whole animal and the description does not include any mention of tentacular palps or branchiae.

Chaetozone A McIntosh, 1911 (p. 165) was described from the same Porcupine Expedition of 3 September 1870 from the Bay of Tunis and is registered as NHM 1921.5.1 3239.

It is in two fragments without a prostomium or posterior region. One fragment has no chaetae and after examination is now considered unidentifiable.

Conclusion

In his description of C. carpenteri McIntosh clearly states that crotchets (=spines) are present from the 10th segment. He also wanted to separate this new species from C. setosa and stated that ‘the crochets differ in their great size’. The spines are very obvious in the notopodia and along with the other characters such as number of segments and arrangement of capillaries are useful for identification of Chaetozone species. The other two Chaetozone species with eyes recorded from the Mediterranean are C. corona from Turkey (Çinar & Ergan 2007) and C. gibber from Turkey and Tunisia (Çinar et al. 2004; Zaâbi et al. 2009). The records of C. corona described by Çinar from Izmir Bay are doubtful as the original description was based on samples from California (Berkeley & Berkeley 1941) and subsequent records are from the NE Pacific (Hartman 1969; Blake 1996). In the description of C. corona from Turkey spines are present in neuropodia from the first chaetiger and there are 62 chaetigers for 19.5 mm body length, whereas Chaetozone gibber is separated from C. carpenteri by the raised dorsal hump in anterior segments, spines from the middle body region and up to 200 chaetigers for 20 mm body length. It is probable that many specimens of C. carpenteri from the Mediterranean have been misidentified. All records of this genus will need to be reassessed. The current distribution is off the coasts of Atlantic Spain, Tunisia, Italy and Croatia in shallow water up 98 m and in silty/sand sediment.

Acknowledgements

We would like to thank E. Sherlock (NHM) for the loan of type material and Alex Muir for his bibliographic help and comments.