INTRODUCTION
We are currently undertaking the first stage of a project entitled ‘Iberian Fauna: Bryozoa’, a long-term research project in which we aim to combine and update all previous citations for Bryozoa in the Iberian Peninsula and Balearics, and to provide new data from the collection of new material. For this, in addition to the obligatory literature search, we are revising our own Iberian material and that deposited in diverse collections, whether published or not.
At present, and although a complete catalogue does not yet exist, we consider that the bryozoological fauna of the Iberian Peninsula is one of the best known in European waters, with approximately 450 recent species cited in this region. Despite this, our overall knowledge is still fragmentary and undoubtedly includes many taxonomic errors, which are gradually being corrected in various studies. The greater ease of access to original reference material and, above all, the use of the electron microscope enable better characterization of species, and makes possible to detect misidentifications and misinterpretations of the species concept as well. Revision of material stored in collections has thus become an essential part of the work of taxonomists.
In the present study, we present 3 new species of cheilostomate Bryozoa. All of the material originates from the Iberian Peninsula, and therefore extends our knowledge of the bryozoan fauna in these waters and in Europe.
MATERIALS AND METHODS
We examined Iberian material originally stored in our own collection—both published and unpublished—as well as unpublished material deposited in collections in the Museo Nacional de Ciencias Naturales de Madrid (MNCN). Other reference material was also examined at the Natural History Museum, London (NHM), and Muséum National d'Histoire Naturelle, Paris (MNHN).
The observations were made with a LEO 435 VP scanning electron microscope on uncoated material. The measurements were made in a camera lucida attached to a Leica MZ16 binocular microscope.
Fig. 1. Chaperiopsis hirsuta sp. nov. (MNCN: paratype 2). (A) View of the colony; (B) detail showing ovicellate and non-ovicellate zooids, spines, and distal avicularia; (C) an autozooid; (D) occlusor laminae.
Fig. 2. Chaperiopsis hirsuta sp. nov. (MNCN: paratype 2). (A) Broken spines with regenerations; left above the peduncle of a proximal avicularium; (B) dietellae; (C) a proximal pedunculate avicularium; (D) ancestrula (centre), periancestrular zooids and a proximal avicularium.
Table 1. Measurements (in mm) of Chaperiopsis hirsuta sp. nov. (MNCN: holotype).
Min, minimum; max, maximum.
TYPE MATERIAL
Holotype: MNCN-25.03/3741: north of Columbrete Grande, Islas Columbretes, 20 July 1996, FAUNA IV, Station 295 B1, 43 m (39º54 02′N–39º53 98′N; 000º41 08′E–000º41 11′E). Colony preserved in alcohol 70º.
Paratypes:
Paratype 1: MNCN-25.03/3742: north-east of Cabo de Pera, Mallorca (Balearics), 27 June 1994. FAUNA III, Station 194A, 58–59 m (39º46 09′N–39º44 17′N; 3º22 04′E–3º35 45′E). Colony preserved in alcohol 70º.
Paratype 2: MNCN-25.03/3743: north-east of Cabo de Pera, Mallorca (Balearics), 27 June 1994. FAUNA III, Station 194A, 58–59 m (39º46 09′N–39º44 17′N; 3º22 04′E–3º35 45′E). One uncoated colony in a SEM slide.
Paratypes 3–8: MNCN-25.03/3744-9: north of Columbrete Grande, Islas Columbretes, 20 July 1996, FAUNA IV, Station 295 B1, 43 m (39º54 02′N–39º53 98′N; 000º41 08′E–000º41 11′E). Colonies preserved in alcohol 70º.
Paratypes 9–11: MNCN-25.03/3750-2: north of Columbrete Grande, Islas Columbretes, 20 July 1996, FAUNA IV, Station 295 B1, 43 m (39º54 02′N–39º53 98′N; 000º41 08′E–000º41 11′E). Three uncoated colonies in a SEM slide.
ETYMOLOGY
hirsuta: alluding to the hirsute appearance of the colony.
DESCRIPTION
Colony encrusting, unilaminar, forming copper-coloured crusts, hirsute appearance owing to presence of zooidal spines. Autozooids oval, elongate, in alternating radiating series. Membranous area oval, occupying almost all of the frontal wall. Cryptocyst smooth, more developed proximally, narrowing gradually towards the distal extreme; its internal border finished off by a flared ring. Occlusor laminae distal, V-shaped, arising in the medial portion of the opesia. Gymnocyst smooth, developed mainly proximally. A vertical peripheral lamina arises from the proximal third, surrounding all of the distal part of the zooid; more conspicuous laterally, with appearance of two well-developed laminar wings. Two pairs of basally articulated, thick, straight spines present on the distal rim, even in the ovicellate zooids, their bases placed outside the peripheral lamina. The 4 spines are similar in both thickness and length, which may greatly exceed the zooidal length. The spines, which are never bifurcate, may present different curvatures and regenerations, and may even modify their growth to avoid the adjacent spines (Figure 2A). Autozooids with a medio-distal dietella and two pairs of lateral dietellae. Distal sessile avicularium present in all zooids, attached between the distal-most pair of spines, and originating from the medio-distal dietella; rounded triangular in outline, narrowed at the apertural bar; mandible ogival, directed distally. Occasionally, an avicularium with short peduncle may appear on the proximal gymnocyst of some zooids, this is difficult to see owing to the development of zooidal spines; rostrum and mandible similar to the sessile avicularium. Ovicell hyperstomial, hemispheric and prominent, with a narrow transversal fenestra parallel to the proximal border. Distally shows an avicularium with distally directed rostrum similar to the sessile avicularium but slightly larger; its curved peduncle covers the apex of the ovicell vertically, and is fused with the ectooecium. Ancestrula, surrounded by periancestrular zooids in the material observed and difficult to see; circular, diameter 0.26 mm, with 9 spines: two disto-lateral pairs and 5 proximal.
REMARKS
Chaperiopsis hirsuta sp. nov. characterized by the oval autozooids with a gymnocyst surrounded internally by a flared ring, and by the presence of two pairs of thick spines which may greatly exceed the length of the autozooid. Furthermore, a distal sessile triangular avicularium is attached between the distal-most pair of spines, while a proximal pedunculate avicularium is infrequent.
Some 50 recent species have been described as belonging to the genus Chaperiopsis, most in the southern hemisphere (Bock, Reference Bock2002) (e.g. Busk, Reference Busk1884; Osburn, Reference Osburn1950; Brown, Reference Brown1952; Uttley & Bullivant, Reference Uttley and Bullivant1972; Gordon, Reference Gordon1984, Reference Gordon1986; Hayward & Thorpe, Reference Hayward and Thorpe1988; Reverter Gil & Fernández Pulpeiro, Reference Reverter Gil and Fernández-Pulpeiro1995; Fernández Pulpeiro & Reverter Gil, Reference Fernández Pulpeiro and Reverter Gil1998). However, only one species of the genus is known in European waters, Chaperiopsis annulus (Manzoni, 1870). This species differs greatly from Ch. hirsuta sp. nov. because it has rounder zooids, with a pair of poorly developed distal spines, and a second bifurcate or even trifucate pair; occlusor laminae horseshoe-shaped; proximal avicularia more abundant, with a longer and narrower peduncle; and ovicell with a wider fenestra and often 1 or 2 distal pedunculate avicularia.
Of the remaining species of the genus, many have more or less branched or swollen spines, sometimes more than 4, or the frontal surface is obscured by calcification (subgenus Clipeochaperia Uttley & Bullivant, Reference Uttley and Bullivant1972), which enables them to be clearly differentiated from C. hirsuta sp. nov. Among those with 4 erect distal spines, some lack a sessile distal avicularium, such as Chaperiopsis galeata (Busk, 1854), Chaperiopsis indefensa Hayward & Thorpe, Reference Hayward and Thorpe1988, Chaperiopsis rotundata Hayward & Thorpe, Reference Hayward and Thorpe1988 and Chaperiopsis patulosa (Waters, 1904), although in the latter species it may appear occasionally. In addition, in all of these species the spines are poorly developed. Chaperiopsis cervicornis (Busk, 1854) possesses a distal avicularium, but small and inconspicuous, and additional spines orientated towards the frontal membrane. In Chaperiopsis lanceola Hayward & Thorpe, Reference Hayward and Thorpe1988 and Chaperiopsis protecta (Waters, 1904) the avicularium is acutely triangular, not rounded; in addition, in both species the pair of proximal spines is more developed, and there may be one or two conspicuous proximal columnar avicularia. Chaperiopsis quadrispinosa (Kluge, 1914) and Chaperiopsis rubida (Hincks, 1881) present 4 erect, cylindrical and rather thick spines, all persisting in ovicellate zooids, as in C. hirsuta sp. nov.; however, in the former, the spines are fusiform, with the proximal pair curving medially, and it presents abundant proximal avicularia, whereas the latter has one or two very conspicuous proximal avicularia, with pointed basal processes. In Chaperiopsis spiculata Uttley, 1949, the proximal pair of spines is much more developed than the distal pair, and the proximal avicularia present distal spiny projections. Finally, in Chaperiopsis splendida Gordon, Reference Gordon1986, the proximal pair of spines is much thicker and flatter than the distal pair, and curve above the opesia overlapping and joining the peduncle of the proximal avicularium.
Fig. 3. (A–D) Metroperiella gay sp. nov. (MNCN: holotype). (A) View of the colony with ovicellate zooids and periancestrular area; (B) an ovicellate zooid; (C) primary orifice; (D) ancestrula (partially covered) and periancestrular zooids, (E–F) Codonella atlantica Canu & Bassler, 1928 (MNHN: type); (E) primary orifice; (F) some ovicellate zooids.
Table 2. Measurements (in mm) of Metroperiella gay sp. nov. (MNCN: holotype).
Min, minimum; max, maximum.
TYPE MATERIAL
Holotype: MNCN-25.03/3738: V-16, Ría de Vigo, 11 June 1992. 42º11′30″N 008º5′30″W, 40 m. Coated.
Paratypes:
Paratype 1: Personal collection: V-16, Ría de Vigo, 11 June 1992. 42º11′30″N 008º51′30″W, 40 m. Dry.
Paratype 2: MNCN-25.03/3739: V-16, Ría de Vigo, 11 June 1992. 42º11′30″N 008º51′30″W, 40 m. Dry.
Paratype 3: MNCN-25.03/3740: V-26, Ría de Vigo, 2 August 1985. 45º13′5″N 8º49′58″W, 34 m. Dry.
COMPARATIVE MATERIAL EXAMINED
Codonella atlantica Canu & Bassler, 1928: type. MNHN, Paléontologie, Canu Coll., without registration number. Morocco. Codonellina lacunata Hayward & Hansen, Reference Hayward and Hansen1999: holotype: NHM-1998.7.29.1. English Channel, 7 November 1961. 49º46′N 0º48′W. Coll. N.A. Holme. Paratypes: NHM-1998.7.29.3-6. English Channel, 7 November 1961. 49º46′N 0º48′W. Coll. N.A. Holme.
ETYMOLOGY
gay: the material ascribed to this new species was kept in a closet for years, and now it has come out of the closet.
DESCRIPTION
Colony encrusting, unilaminar, forming small circular patches. Autozooids rounded hexagonal, convex, separated by distinct grooves, placed in alternating series. Frontal shield smooth, uniformly perforated by some 40 circular pores, except in a small area proximal to the orifice and the proximal end of the autozooid. Primary orifice subcircular; proximal border shallowly concave with two small rounded condyles. Laminar peristome well developed in the proximal half of the orifice. Oral spines absent, except in periancestrular zooids, which present 2–3 thin oral spines. Small suboral avicularium, placed on a small umbo just below the orifice, connected to the peristome; oval, with apertural bar complete. Ovicell globular, wider than long, united with the peristome. Ectooecium almost entirely membranous, forming a basal ring of calcification; entooecium marked with radial ridges and perforated by rounded pores except in the central area. Ancestrula tatiform, partially covered in the material examined. Astogenesis begins with two disto-lateral zooids, similar to successive autozooids but smaller, perforated by some 20 pores, and with 2–3 spines.
REMARKS
Four colonies encrusting dead shells were examined; all were collected in the Ría de Vigo in 1985 and 1992, from gravelly seabed. This species is characterized by hexagonal autozooids, with a smooth surface perforated by some 40 circular pores, a vertical laminar peristome developed in the proximal half of the circular primary orifice, and a small oval suboral avicularium.
Metroperiella gay sp. nov. is very similar to Codonellina lacunata Hayward & Hansen, Reference Hayward and Hansen1999, a species described in the English Channel, whose type material we have had the opportunity to study. However, there are clear differences between the two species, the most noteworthy being the presence in M. gay sp. nov. of a suboral avicularium, while in the same position C. lacunata has an oval chamber with a membranous surface. Furthermore, in M. gay sp. nov. the orifice is more rounded and has rounded condyles, the peristome begins towards the lateral half of the edge of the orifice (Figure 3C) and is basically vertical, whereas in C. lacunata the condyles are triangular, the peristome reaches the distal ends of the primary orifice and its edge is clearly flared. Moreover, a small proximal or proximo-lateral area without pores is usually observed in the zooids of M. gay sp. nov. but not in C. lacunata. Finally, in the colonies of C. lacunata the zooids are arranged in very clear radial series.
The present species also presents some similarities to Codonella atlantica Canu & Bassler, 1928, a species from Morocco, poorly described in the original paper; however, we have studied the type material held at the MNHN (Figure 3 E, F). Codonella atlantica presents a larger orifice, with the proximal border more concave and two small triangular condyles; the peristome reaches the distal ends of the primary orifice, like in C. lacunata, but it is somewhat quadrangular in outline, with straight lateral walls and two marked proximo-lateral corners. Zooids are larger, also perforated by rounded pores, but the imperforate central area is more extensive, while there is no imperforate proximo-lateral area. Finally, the ovicell is comparatively smaller.
Hayward & Hansen (Reference Hayward and Hansen1999) placed their species in the genus Codonellina; despite the differences between M. gay sp. nov., C. atlantica and C. lacunata, it is clear that the three species are closely related, and therefore we consider that they should be included in the same genus. However, several authors (e.g. Harmer, Reference Harmer1957; Hayward, Reference Hayward1974; Gordon, Reference Gordon1984; Zabala, Reference Zabala1986) have discussed the possible synonymy between the genera Metroperiella Canu & Bassler, Reference Canu and Bassler1917 and Codonellina Bassler, Reference Bassler1934.
The genus Metroperiella was introduced by Canu & Bassler (Reference Canu and Bassler1917) for Schizoporella lepralioides Calvet in Jullien & Calvet, 1903, a species described in the Azores and widely cited in the Mediterranean. On the other hand, the same authors (Canu & Bassler, Reference Canu and Bassler1927) described the genus Codonella for Lepralia galeata Busk, 1854; as this generic denomination was pre-occupied, Bassler (Reference Bassler1934) substituted it with Codonellina.
Lepralia galeata is a perfectly defined species, which has been re-described by López Gappa (Reference López Gappa1981), and whose type material is deposited in the Natural History Museum, London (registration no: 1854.11.15.265; López Gappa, personal communication, September, 2008).
In contrast, there is no type material of S. lepralioides and moreover, its identity is not clear as numerous authors have suggested the possibility that this species is in fact a junior synonym of Flustra montferrandi Audouin, 1826, a species described in the Red Sea, for which there is also no type material. It would therefore be necessary to design neotypes for both species to be able to resolve this question definitively. Only once this problem has been resolved, can the most likely synonym between the genera Codonellina and Metroperiella be decided on.
At present, and as in recent studies this synonym has been informally accepted (see Tilbrook, Reference Tilbrook2006; Bock, Reference Bock2007; Gordon, Reference Gordon2007), we have placed our species in the genus Metroperiella.
Fig. 4. Schizoporella artabra sp. nov. (A) (MNCN: holotype), view of the colony with some extra avicularia; (B) (MNCN: paratype 4), an ovicellate zooid (centre) and extra avicularia; (C) (MNCN: holotype), primary orifice and three avicularia; (D) (MNCN: paratype 3), primary orifice and one avicularium; (E) (MNCN: holotype), autozooid; (F) (MNCN: paratype 4), ovicellate zooid and extra avicularia.
Table 3. Measurements (in mm) of Schizoporella artabra sp. nov. (MNCN: holotype).
Min, minimum; max, maximum.
Schizoporella dunkeri (Reuss): Lanza Suárez & Fernández Pulpeiro, Reference Lanza Suárez and Fernández Pulpeiro1984: 280; Reverter Gil, Reference Reverter Gil1995: 175.
Schizoporella hesperia Hayward & Ryland: Reverter-Gil & Fernández-Pulpeiro, Reference Reverter Gil and Fernández-Pulpeiro1999: 43, figure 4 A, B.
TYPE MATERIAL
Holotype: MNCN-25.03/3753: Ría de Ferrol, D–12a, 43º27′36″N 008º17′30″W, 13 June 1989, 20 m. Uncoated, on a SEM slide.
Paratypes:
Paratype 1: MNCN-25.03/3754: Ría de Ferrol, D-12a, 42º27′36″N 008º17′30″W, 13 June 1989, 20 m. Dry.
Paratype 2: MNCN-25.03/3755: Ría de Ferrol D-8b, 43º28′03″N 008º18′36″W, 22 October 1990, 16 m. Dry.
Paratype 3: MNCN-25.03/3756: Ría de Ferrol D-18b, 43º28′15″N 008º15′30″W, 22 October 1990, 15 m. Dry.
Paratype 4: MNCN-25.03/3757: Ría de Ferrol, D-12a, 43º27′36″N 008º17′30″W, 13 June 1989, 20 m. Coated.
Paratype 5: personal collection: Ría de Ferrol, D-12a, 42º27′36″N 008º17′30″W, 13 June 1989, 20 m. Dry.
ETYMOLOGY
artabra: the Ría de Ferrol, where the material here ascribed to this new species was collected, is placed in the ‘Artabrian Gulf’.
DESCRIPTION
Colony encrusting, unilaminar, forming whitish crusts. Autozooids oval to hexagonal, irregularly arranged and separated by distinct sutures. Frontal shield convex, smooth to slightly rough, uniformly perforate by small round pores. Primary orifice orbicular with a V-shaped rounded sinus occupying a half of the proximal border; long more than 1/4 of the orifice. Condyles conspicuous, elliptical, with acute corners. No oral spines. Avicularia single or paired, sometimes absent; proximal portion level with the proximal border of the orifice, distal end exceeding the distal end of the orifice; rostrum triangular, disto-laterally directed; crossbar slender, complete. In some material autozooids may present 1–6 extra avicularia, with structure similar to oral avicularia, placed in the margins of the zooid and directed outwards. Ovicell hyperstomial, rounded, scarcely prominent. Frontal surface with a rough imperforate central area; periphery of the ovicell perforated by pores similar to those of the frontal surface of the zooid, and continuous with it. Kenozooids present in some material. Size and shape similar to autozooids, but without orifice; sometimes with a distal adventitious avicularium (Paratype 5).
REMARKS
The genus Schizoporella Hincks, 1877 is well defined, and numerous species have been described throughout the world. Hayward & Ryland (Reference Hayward and Ryland1995) have re-described the species present in the British Isles, but there are undoubtedly many other species in European waters.
Schizoporella artabra sp. nov. characterized by a primary orbicular orifice, with a V-shaped rounded sinus occupying half of the proximal border, flanked by condyles with acute corners; the proximal portion of the avicularia is level with the proximal border of the orifice, while its distal end exceeds the distal border of the orifice. Finally, the ovicell is scarcely prominent. No other species from Europe or from elsewhere, display a similar combination of characters.
The material here ascribed to S. artabra sp. nov. was previously cited by us as Schizoporella dunkeri (Reuss, 1848) (Lanza Suárez & Fernández Pulpeiro, Reference Lanza Suárez and Fernández Pulpeiro1984; Reverter Gil, Reference Reverter Gil1995), and later as Schizoporella hesperia Hayward & Ryland, Reference Hayward and Ryland1995 (Reverter Gil & Fernández-Pulpeiro, Reference Reverter Gil and Fernández-Pulpeiro1999). However, we now consider that the differences between our material and S. hesperia are sufficiently important not to ascribe it to this species. The most important differences include: the absence of umbo in our material, a more orbicular orifice, a more open sinus, condyles with acute corners, and also the position and orientation of the avicularia, situated more distally than in S. hesperia, and forming a smaller angle with the longitudinal axis of the autozooid. Furthermore, in our material we did not observe the presence of the dimorphic zooids described in S. hesperia, but we did observe 1–6 extra avicularia at the edges of the autozooid.
Extra avicularia have been reported in Schizoporella ansata as defined by Canu & Bassler (Reference Canu and Bassler1930) and Gautier (Reference Gautier1962). Ryland (Reference Ryland1968) considered that these records actually belong to Schizoporella magnifica Hincks, 1886; this author also cited extra avicularia in this species. However, S. magnifica (as described by Hayward & Ryland, Reference Hayward and Ryland1999) clearly differs from S. artabra sp. nov. having an orifice that is longer than wide, with a deep, narrow, U-shaped sinus; the avicularia, usually paired and with a voluminous cystid, which occupies an even more distal position than in our material; and by the egg-shaped ovicell.
Other species of the genera Schizoporella: S. dunkeri (Reuss, 1848), S. unicornis (Johnston in Wood, 1844), S. errata (Waters, 1878) and S. cornualis Hayward & Ryland Reference Hayward and Ryland1995, were found at the same site where the material of S. artabra sp. nov. was collected. However, all of these show clear differences, especially in the form of the orifice and the sinus, and in the position and orientation of the avicularia, which make them perfectly distinguishable.
ACKNOWLEDGEMENTS
We are grateful to J.-L. d'Hondt (MNHN–DMPA), J.P. Saint Martin and H. Senut (MNHN–Paléontologie), and M. Spencer Jones (NHM) for valuable assistance during our visits and the loan of specimens and to F. Yagüe (MNCN) for the loan of specimens. We also thank Juan López Gappa and Dennis Gordon for their helpful comments. This work was supported by the project ‘Fauna Ibérica: Briozoos I (Ctenostomados y Queilostomados Anascos)’ (CGL2006-04167), co-financed by the Ministerio de Ciencia e Innovación, Spanish Government and FEDER. O.R.G. and J.S. gratefully acknowledge two SYNTHESYS grants (GB-TAF-4454 and FR-TAF-3988 respectively), which enabled visits to the NHM and the MNHN. The SYNTHESYS Project (http://www.synthesys.info/) is financed by European Community Research Infrastructure Action under the FP6 ‘Structuring the European Research Area’ programme.
