New and non-indigenous species of Bryozoa from Iberian waters

. Iberian material originally identi ﬁ ed as Hincksina ﬂ ustroides is revised and ﬁ ve different species are now identi ﬁ ed: the Atlantic species Hincksina ﬂ ustroides is present to the NW of the Iberian Peninsula, whilst the Mediterranean species Hincksina synchysia is here reported for the ﬁ rst time in Iberian waters. Two new species of Hincksina are described, one from the Strait of Gibraltar area


Introduction
This contribution is part of an effort to collate all the previous knowledge about living marine Iberian Bryozoans. This involves compiling bibliography, reviewing material in collections and collecting new material. This effort has yielded several new contributions including corrections of previous identifi cations, new records for the Iberian Peninsula and descriptions of new species (e.g., Reverter-Gil et al. 2016, 2019a, 2019b, 2021a, 2021b. Nonetheless, our overall knowledge remains 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 characterizations of species, and make it possible to detect misidentifi cations and misinterpretations of species concepts as well. Revision of material stored in collections has thus become an essential part of the work of taxonomists, either to redescribe known species or to describe new ones.

Description
Colony encrusting, multiserial, unilaminar or occasionally multilaminar, forming extensive crusts that cover the substrate on which they grow. Autozooids elongate oval or rectangular, arranged in series, separated by shallow grooves. Distal wall generally ascending towards the frontal surface and angularly bent from side to side or arch-like. Gymnocyst reduced to the proximal region. Two (rarely three) conical, hollow processes, generally open at the end, developed on the proximal gymnocyst, half-way between the central line and the lateral margins. These processes may sometimes be rudimentary or even absent in large parts of the colony. The fi rst zooid in each of both series after bifurcation bears a  (Levinsen, 1909), holotype (NHMD-77254), Koh Samet (Gulf of Thailand). A. Autozooids with well-developed spines and gymnocystal processes. B-C. Autozooids with fewer, poorly developed spines and gymnocystal processes. D. Autozooids without spines and gymnocystal processes.
single median process. Cryptocyst granular, poorly developed, absent at the distal end of the opesia, somewhat more evident at its proximal end, leaving an extensive, oval opesia. Communication via two rather large, multiporous rosette-plates situated in the basal corners of the distal wall. The distal half of each lateral wall has a single multiporous rosette-plate. Oral spines absent. The development of marginal spines shows great differences. Some zooids bear up to 8 pairs of marginal spines, not very thick, somewhat fl attened, recurved on the opesia, reaching the middle of the area or even surpassing it, but in general the spines are smaller and fewer, and many zooids are completely spineless. There are no ovicells or avicularia. Irregular intercalary kenozooids, small, scattered, fi lling gaps between autozooids. Ancestrula unknown.

Remarks
Electra angulata was originally described by Levinsen (1909) from colonies collected on a ligneous core fl oating near Koh Samit, Siam (Ko Samet, Thailand). The description is very clear and complete, but not so the only drawing (Levinsen 1909: pl. 22 fi g. 4a), which provides no information on the variability of the species. Levinsen (1909: 149), however, clearly stated that "The best provided ones  (Levinsen, 1909) (MHNUSC-Bry 643), Menorca (Balearic Islands). A. Autozooids with well-developed spines and gymnocystal processes. B. Multiporous rosette-plates. C. Spineless autozooids. D. Second layer of spineless autozooids covering the fi rst layer of spiny zooids.
[autozooids], which in the colonies examined are in a great minority, have on the margin 12 not very thick spines, which reach the middle of the area or even surpass it. A larger or smaller number of them is however often wanting, and many zooecia are altogether without spines. On the proximal gymnocyst we fi nd in most zooecia 2 (more rarely a single median and still more seldom 3) short, thick, conical spines, generally open at the end, which are situated half-way between the central line and the lateral margins. These spines may sometimes be rudimentary, and in many zooecia (with or without marginal spines) they are absent". On page 156 of the same paper, it is also stated that "Here we may fi nd in the same colony some zooecia, which are entirely without spines, and others provided with a larger number of these structures." It is clear then that according to Levinsen (1909) the zooids of E. angulata may have spines and gymnocystal processes at the same time, or only processes, or only spines, or even none of them, and both processes and spines may be very variably developed. We have been able to verify all this variation both in the type material of E. angulata (NHMD-77254 and Fig. 1), in our own material from the Mediterranean and the Strait of Gibraltar (MHNUSC-Bry 643, 708) (Figs 2-3), and in different museum samples (see Material examined above). Nonetheless, later authors have apparantly incorrectly considered that only the presence of spines (as well as gymnocystal processes) is a typical character of the species.
The species also has a marked tendency to form multilaminar colonies due to the overgrowth of some layers over others (Figs 2F,3D). This does not seem to have been pointed out by other authors, but is clearly visible in our own material, a large colony on plastic debris (MHNUSC-Bry 643) ( Fig. 2A-B).
The upper layers are often spineless to a great extent (Figs 2F, 3D).
Membranipora tenella Hincks, 1880 has also been subject to misinterpretation and even considered a synonym of E. angulata or at least misidentifi ed (e.g., Marcus 1937;Silén 1941;Mawatari 1974;Rosso 1994;Subías-Baratau et al. 2022). However, as already stated about twenty years ago by Tilbrook et al. (2001), they are clearly separate species. These authors (Tilbrook et al. 2001: 40) fi rst pointed out as a difference that M. tenella sensu stricto bears no marginal spines (see Fig. 4), but as we have already demonstrated (see above), zooids of E. angulata usually lack them. Tilbrook et al. (2001) did, however, accurately point out another important difference between the two species: the development of the gymnocystal processes in M. tenella, which are far more robust and knob-like, occupying a greater area of the gymnocyst (Fig. 4A). Importantly, the revision of type material of M. tenella (NHMUK 1899.5.1.684, Fig. 4) shows that the proximal cryptocyst is characteristically much more developed in this species than in E. angulata (see Figs 3A, C, 4A).
The resemblance of E. angulata with Conopeum papillorum Tilbrook, Hayward & Gordon, 2001 was already discussed in the original paper (Tilbrook et al. 2001: 40) and we have nothing more to add. Finally, material of E. angulata was also considered as a new species (named Electra inexpectata) in the unpublished PhD by López de la Cuadra (1991), but this species was not formally published. It was reported later from the Strait of Gibraltar area by López de la Cuadra & García-Gómez (1994) as Electra cf. tenella.
The systematic position of E. angulata has also been subject of discussion ever since its original description. Levinsen (1909) placed the species in Electra with some reservations, closely allied to Electra monostachys (Busk, 1854). This position was also accepted by Harmer (1926). But the original author himself (Levinsen 1909: 160) also related his new species to Aspidelectra melolontha (Landsborough, 1852), the type species and, at that time, the only species of his new genus Aspidelectra Levinsen, 1909. Indeed, as Levinsen (1909 stated, both species share an angularly bent distal wall with a multiporous rosette-plate in each of the two basal corners, and 1-2 gymnocystal processes. These processes somehow replace the oral spines, absent in both species. Note that, for this reason, Aspidelectra defensa (Kirkpatrick, 1888) and Aspidelectra densuense Cook, 1968 cannot remain in this genus because they do have true oral spines and lack gymnocystal proximal processes. This requires further discussion, which is beyond the scope of the present work. Anyway, in our opinion E. angulata cannot be placed in this genus either because Aspidelectra is characterized by a frontal shield of fused, fl attened spines.
In contrast, M. tenella was fi rstly placed in Electra Lamouroux, 1816 by Marcus (1937), and the name Electra tenella was widely used by later authors, even although the species cited by Marcus (1937) and others was actually E. angulata. Nikulina (2007Nikulina ( , 2010 and Nikulina & Schäfer (2008) distributed a number of species previously attributed to Electra to new genera, but these did not include E. angulata and M. tenella. Tilbrook & Gordon (2015) pointed out some similarities of both species with Arbopercula bengalensis (Stoliczka, 1869), the type species of the genus Arbopercula Nikulina, 2010, but they inadvertently erred in attributing both species to Arbocuspis Nikulina, 2010 in their paper. This was clearly a lapsus owing to the similarity of both generic names (D.P. Gordon pers. com.). Electra angulata and M. tenella do not conform to Arbocuspis as the cryptocyst in this genus is inconspicuous and spines are branching, bending across the opesia from its proximal end, forming a sort of shield. We agree with the original intention of the authors since the relationship between the three species is evident, but the inclusion of E. angulata and M. tenella in Arbopercula requires modifying the diagnosis of the genus, which also contains a serious error of understanding and an important omission: Firstly, Arbopercula was originally characterized (and named) based on a pair of bifurcating, chitinous spines on the operculum, which are absent in E. angulata and M. tenella. As Tilbrook & Gordon (2015) pointed out, however, these spines may be very small and easily overlooked. In our opinion, this character can be useful to differentiate the type species, but not to characterize the genus, so it should be eliminated from the generic diagnosis. Otherwise, no other species could be integrated into the genus because this character is exclusive of A. bengalensis.
Secondly, the diagnosis of Arbopercula, but also Arbocuspis, is incorrect. Both diagnoses state that the zooids exhibit a pair of distal spines, stout, conical, non-articulated, but these are really gymnocystal blunt processes (not true spines) and are located at the proximalmost end of the succeeding zooid, such as those present in A. bengalensis, E. angulata and M. tenella (but also in A. melolontha), probably replacing the oral spines, absent in these species.
Thirdly, the absence of marginal spines in M. tenella and in many zooids of E. angulata must be included in the diagnosis.
Finally, unfortunately no information was provided about the interzooidal communication pores of both genera. We have no information about A. bengalensis, but E. angulata and M. tenella have on either side a rather large, multiporous rosette-plate situated in one of the basal corners of the distal wall. In our opionion this must be also incorporated into the diagnosis of the genus Arbopercula. Tilbrook & Gordon (2015) tentatively also added Membranipora devinensis Robertson, 1921 to Arbopercula, but the presence of proximal pores and a presumed ovicell prevents the inclusion of this species, at least until it is correctly described.
Material of A. angulata has been frequently incorrectly reported as E. tenella by many authors around the world, growing on different algae, on drifting or beach-stranded plastic or wood, as small colonies on the hulls of pleasure craft plying tropical and subtropical waters, or even as epibionts attached to the scales of sea snakes, shells of living nautilus or carapaces of sea turtles or horseshoe crabs (e.g., Key et al. 1995Key et al. , 1996Pfaller et al. 2008;Gordon 2009;Tan et al. 2011;Subías-Baratau et al. 2022

Remarks
The The biometries of our own material (Table 3) are signifi cantly similar to those included by Berning et al. (2021) in the redescription of the species, except for the size of the avicularia. Indeed, the measurements of these authors are much lower than ours, but this seems to refl ect a calculation error. In fact, measuring on their photographs (Berning et al. 2021: fi g. 1), the sizes of the avicularia are approximately 0.21 × 0.16 mm, very close to our own data. Size is also similar to the material fi gured in the British Fauna (Hayward & Ryland 1998).
The rest of the Iberian material labelled as H. fl ustroides that we have revised, from the Strait of Gibraltar to the Spanish Levant, does not really correspond to the species. This includes the records of H. fl ustroides published by Zabala (1986), López de la Cuadra & García Gómez (1988Gómez ( , 1994, López de la Cuadra (1991) and Templado et al. (2002Templado et al. ( , 2006, which correspond to other species (see below). Other records from the Mediterranean, such as those of Zabala & Gili (1985), Calvín Calvo (1986), Álvarez (1992) or Madurell et al. (2013), very probably do not correspond to this species either, but to H. synchysia (see below). Other species, however, could be involved, calling for new sampling in these areas.
According to Berning et al. (2021), H. fl ustroides was known with certainty only from western Great Britain, the Channel, and the southern North Sea, whereas its southern limit of distribution remained unclear. Here we considerably extend the southern limit of the species, which reaches at least the Portuguese coast around Lisbon. There does not seem to be at present any realiable record or material of H. fl ustroides from the south of Portugal, the area of the Strait of Gibraltar or from the Spanish Mediterranean (Fig. 12B). Berning, Spencer Jones & Vieira, 2021 Figs 6, 12B; Table 4 Hincksina fl ustroides f. crassispinata Gautier, 1962: 50, fi g. 8.

Hincksina synchysia
Hincksina synchysia Berning et al., 2021: 337, fi g. 2. Comparing with the measures of Berning et al. (2021), based on two samples from Portofi no, we fi nd that the autozooids are smaller in our material, while the avicularia are larger.
In contrast, the record of H. fl ustroides from Alboran Island made by Templado et al. (2006) does in fact correspond to Hincksina chimaera sp. nov. and to Hincksina longispinosa Harmelin & d'Hondt, 1992 (see below). In any case, we cannot be certain whether we have revised all of the original material cited by these authors. At the same time, the record of the same species by Álvarez (1992) in the same area is doubtful because the original paper does not include a description or fi gures, whilst the original sample  Gómez (1988Gómez ( , 1994 and López de la Cuadra (1991) really correspond to Hincksina elephantina sp. nov., at least in part (see below). Therefore, at present, H. synchysia seems to be absent from the Alboran Sea, the Strait of Gibraltar area and the whole Atlantic Iberian coast, so it perhaps must be confi ned to the inner part of the western Mediterranean (Fig. 12B).

Differential diagnosis
Hincksina with 12 thick marginal spines that increase in size distally and a pair of thick oral spines, long and gently curved, resembling elephant tusks. Together, the four distal spines give the autozooid a characteristic appearance. Interzooidal avicularia abundant, with an oval mandible directed distally, usually present in dense clumps. Ovicell endozooidal in distal avicularium or autozooid; proximal margin of ooecium raised centrally, producing a developed central peak.

Etymology
The term ‛elephantina' alludes to the appearance of the two oral spines in this species, which are thick, long and gently curved, resembling elephant tusks.

Description
Colony encrusting, multiserial, unilaminar. Autozooids elongate oval, arranged in irregular series, separated by shallow grooves. Gymnocyst reduced to the proximal region, cryptocyst a narrow band with a coarse nodular surface, opesia extensive. Vertical walls with two or more uniporous pore plates per neighbouring zooid. A pair of long oral spines, thick and gently curved, resembling elephant tusks. Frontal membrane overarched by 12 (occasionally more) cylindrical mural spines, thick, tapering towards the uncalcifi ed tip. Spines more or less folded over the frontal membrane, sometimes overlapping in the midline. Generally, the fi rst pair is longer and more vertical but converging medially, while the height of the spines decreases proximally, the most proximal spines being the shortest. Spines apparently unjointed at their base. Avicularia interzooidal, rectangular or square in outline, distal to many zooids; usually pointing distally next to autozooids and laterally or distolaterally in ovicelled zooids; rostrum semielliptical, at an acute angle to colony surface, mandible hinged on a pair of short triangular condyles delimiting an approximately semicircular proximal area distinctly wider than long. In concavities or convexities of the substrate, avicularia tend to form irregular clusters. Ovicell endozooidal in distal avicularium or less frequently in the distal autozooid; ooecium continuous with the gymnocyst of the avicularia or autozooid, forming a broad hemispherical cap; proximal margin raised centrally, producing a developed central peak. Ancestrula not observed.

Remarks
Material here described as H. elephantina sp. nov. was originaly reported as H. fl ustroides by López de la Cuadra & García Gómez (1988Gómez ( , 1994 and López de la Cuadra (1991). However, H. elephantina clearly differs from this and other Iberian species of the genus by several characters: The two oral spines are thick, long, and gently curved, resembling elephant tusks, while the fi rst pair of the lateral spines is also very developed but shorter, curved and converging medially. Taken together, these spines give a characteristic appearance to the oral end of the autozooid (Fig. 7A-C). Moreover, the remaining spines are also stout, cylindrical and thinner at the tip, covering the opesia but not fusing along the medial line. The average number of marginal spines is 12 (Fig. 7A-C). The quadrangular avicularia are abundant, as in other species such as H. synchysia or H. chimaera sp. nov. (see below), but with a semielliptical rostrum distally directed. Moreover sometimes they appear in clusters, a character not seen in other Hincksina (Fig. 7D-F). Finally, the ovicell of H. elephantina sp. nov. can be immersed in an autozooid or in an avicularium (Fig. 7G), a character that Berning et al. (2021) considered exclusive of H. synchysia, but which is also present in H. chimaera (see below). The ovicell of H. elephantina, however, presents a developed central peak (Fig. 7G), which is characteristic of H. fl ustroides according to Berning et al. (2021).
At present, H. elephantina sp. nov. is known only from La Línea, Andalucía (Strait of Gibraltar area), collected at 30-50 m depth (Fig. 12B). López de la Cuadra & García Gómez (1988) and López de la Cuadra (1991) reported two colonies of H. fl ustroides from this locality, but only one is now preserved, here designated as the holotype of the species (MHNUSC 10127). López de la Cuadra (1991) also reported this species from Tarifa at 50-60 m depth, but as the original colony was not preserved we cannot be certain about its identity.

Differential diagnosis
Hincksina with 8-10 mural spines, frequently fl attened and occasionally bifi d, and a pair of oral spines usually fl attened, somewhat triangular, with the beginning of a bifurcation, parallel or aligned oblique to zooidal midline and converging distally. Avicularia interzooidal, with the rostrum semielliptical  (Fig. 9).

Description
Colony encrusting, multiserial, unilaminar. Autozooids elongate oval, arranged in irregular series, separated by shallow grooves. Vertical walls with two or more uniporous pore plates per neighbouring zooid. Gymnocyst reduced to the proximal region, cryptocyst a narrow band with a coarse nodular surface, opesia extensive. A pair of oral spines cylindrical or more usually fl attened, directed upwards, somewhat triangular, with the beginning of a bifurcation with uncalcifi ed pores in the tips, parallel or aligned oblique to zooidal midline and converging distally. Frontal membrane overarched by 8-10 (occasionally fewer) mural spines, initially cylindrical but frequently fl attened, well developed and reaching the zooidal mid-line; occasionally bifi d. Spines apparently unjointed at their base. Avicularia interzooidal, distal to many zooids, usually pointing distolaterally but often distally or even laterally; oval in outline and distinctly longer than wide, widest proximal to condyles; rostrum semielliptical or rounded-triangular, at an acute angle to colony surface, mandible hinged on a pair of short triangular condyles delimiting an approximately semicircular proximal area with an immersed calcifi ed shelf; interior of mandible with a large central oval lucida; gymnocyst usually reduced to cystid corners. Ovicell endozooidal in distal avicularium or autozooid; ooecium continuous with the gymnocyst of the avicularian or autozooidal cystid, forming a short but broad hemispherical cap; proximal margin raised centrally, often producing a central peak. Ancestrula not preserved.

Remarks
Hincksina chimaera sp. nov. exhibits a curious mix of characters from three other Iberian species of the genus: H. fl ustroides, H. synchysia and H. calpensis Reverter-Gil, Souto & Fernández-Pulpeiro, 2012. At fi rst sight, H. chimaera sp. nov. closely resembles H. fl ustroides, also showing a high number of large marginal spines (8-10), fl attened and sometimes bifi d (especially the fi rst pair) overarching the frontal membrane (Fig. 8). Moreover, the ovicell of Hincksina chimaera sp. nov. presents a developed central peak (Fig. 8E), which is characteristic of H. fl ustroides according to Berning et al. (2021).
In contrast, H chimaera sp. nov. shares with H. synchysia the size, shape, orientation and abundance of avicularia (Figs 6 and 8). In addition, the ovicell can be immersed in an autozooid or in an avicularium ( Fig. 8C-D), a character that Berning et al. (2021) considered exclusive of H. synchysia. The number of marginal spines is, however, lower in H. synchysia (6-9) than in H. chimaera (8-10).
Finally, the oral spines of H. chimaera sp. nov. closely resemble those present in the ovicelled zooids of H. calpensis (see Reverter-Gil et al. 2012: fi g. 2b, d), being erect, fl attened and somewhat triangular, with the beginning of a bifurcation and uncalcifi ed pores in the tips, which are parallel or aligned oblique to the zooidal midline and converging distally (Fig. 8B-D). The development of the spines is variable, as in other species of the genus (Berning et al. 2021), but contrary to what was stated by those authors, their development does not seem to be related to the degree of exposure of the colony to the substrate, but with the ontogeny of the colony, because the marginal and oral spines are often fi ner and more cylindrical near the periancestrular region whereas they are more developed and fl attened in the areas furthest away from the colony. This is visible in sample MNCN 25.03/2488, which contains fi ve colonies in different positions in a concretion. A similar variation also occurs in H. calpensis, although in this species the spines are much more markedly developed, forming a frontal shield.
Hincksina chimaera sp. nov. was previously reported from the Alboran Sea by Templado et al. (2006), as H. fl ustroides (in part) without further information. We have revised the original material, conserved at the MNCN (see material examined and Figs 8, 12B). Almost all the material actually corresponds to H. chimaera, but a single sample (MNCN 25.03/2402, Fig. 9) corresponds to an ovicelled colony of Hincksina longispinosa Harmelin & d'Hondt, 1992. This species was known only from its original description in the Gulf of Cadiz at 135-521 m depth, but it was very recently reported in the Alboran Sea at slightly shallower depths (112-120 m) by Ramalho et al. (2022). The present colony also comes from a similar depth (87-213 m) (Fig. 12B).
The record of H. fl ustroides made by Álvarez (1992)

Differential diagnosis
Caberea with two oral spines in the outer distal angle, the outermost stouter, and one spine in the inner distal angle. Opesia obscured by a thick, oval scutum, its distal edge curved or irregular, hiding the operculum. Frontal avicularium monomorphic, small, with rounded mandible, present in almost all autozooids. Ovicell as long as wide, with an irregularly rounded fenestra of uncalcifi ed ectooecium directly above the aperture.

Etymology
Alluding to the geographic origin of the studied material.

Description
Colonies forming erect tufts up to 4 cm high. Branches straight, cylindrical, dividing dichotomously at regular intervals, without visible joints. Autozooids in two alternating longitudinal series, with frontal planes angled at about 120° to each other, defi ning the frontal surface of the branch. Autozooids elongated rectangular, with arched distal end. Opesia oval, constituting about three-quarters of total frontal length, bordered by broad, coarsely granular cryptocyst. Frontal proximal gymnocyst smooth. Two oral spines in the outer distal angle, the outermost stouter; one spine in the inner distal angle. All of them broken in the studied material. Central autozooid in a bifurcation with 4-5 spines. Ovicelled zooids with one spine in each angle. Almost all the opesia obscured by a thick, oval scutum, attached by a thick stalk one-half of the distance down the inner margin of the opesia. Its distal edge is curved or irregular, hiding the operculum; proximal part forming an ovate lobe. Outer edge of scutum does not cover the cryptocyst. Lateral avicularium very small, directly next to the outermost spine and diffi cult to see, with a rounded triangular mandible directed outwards. Frontal avicularium monomorphic, small, present in almost all autozooids, occupying part of the proximal gymnocyst just between the opesia and the stalk of the scutum of the proximal autozooid of the other series. Mandible small, rounded triangular, directed upwards. Ovicell as long as wide, recumbent on distally succeeding autozooid, occupying all its gymnocyst and hiding also the proximal cryptocyst; frontal avicularium of distal zooid displaced, united to distal inner angle of ooecium. An irregularly rounded fenestra of uncalcifi ed ectooecium directly above the aperture. In ovicelled zooids the scutum tends to be slightly shorter, revealing part of the operculum. Abfrontal surface of colony covered by large, proximally tapered vibracula, with long and straight setal grooves, inclined to the branch axis; seta long, fi nely toothed along one edge. Setal groove of the central vibracula in the bifurcation shorter, located on the axis of the branch. Thin kenozooidal rhizoids arising from vibracular chambers, passing proximally along median abfrontal surface of each branch.

Remarks
The great majority of the species of Caberea are distributed around the Pacifi c Ocean (Bock 2023). Only two species have been reported in European waters: the northern species Caberea ellisii (Fleming, 1814) and the supposedly widespread Caberea boryi (Audouin, 1826). Caberea ellisii differs from C. cantabra sp. nov. most obviously by the lack of a scutum.
Although C. cantabra sp. nov. shows some similarities with C. boryi (it was previously reported under this name), both species clearly differ in several characters: Autozooids are clearly smaller in C. boryi (0.37 × 0.23 mm according to Hayward & Ryland 1998). The scutum is completely different, being wider in C. cantabra sp. nov., projecting distally and covering the operculum (Fig. 10A-C), but lacking the projecting blunt process characteristic of C. boryi (Fig. 11B-C). Moreover, in this latter species the edge of the scutum is perfectly parallel to the edge of the opesia, leaving a fi ne separation (Fig. 11B-C), which does not occur in C. cantabra (Fig. 10C).
The lateral avicularium is much smaller in C. cantabra sp. nov., almost inconspicuous and hidden by the base of a spine (Fig. 10C-D). Moreover, the large frontal avicularium that characterizes C. boryi (Fig. 11C) has not been observed.
The fenestra of the ovicell is large, about as long as wide (Fig. 10D), whilst in C. boryi the fenestra is clearly wider than long and usually has a wide calcifi ed rim on the proximal edge ( Fig. 11B-C). Finally, C. boryi seems to be a shallow-water species, ranging from the intertidal to 100 m depth as much (Hayward & Ryland 1998), whilst the material here identifi ed as C. cantabra sp. nov. comes from 342-690 m depth. D'Hondt (1973) reported C. boryi from deep waters to the NW of the Iberian Peninsula. We have examined the only two samples stemming from this area held at the MNHN: MNHN-IB-2008-6926 (stn Thalassa T503, 490 m depth), and MNHN-IB-2008-6935 (stn Thalassa T512, 510-630 m depth). Both actually correspond to C. cantabra sp. nov. (see material examined and Fig. 10D-E). D'Hondt (1973) also reported C. boryi from a nearby locality (stn Thalassa 478, 513-550 m depth), but no preserved material was found, so we cannot be certain about its identity, although it is very likely that it is also C. cantabra. Jullien (1882) also reported C. boryi from a nearby locality (Travailleur stn 39b, 1037 m depth), but the lack of reference material and additional data prevents us from verifying the identifi cation. We have not been able to locate coordinates for the station 39b, not reported by Jullien (1882), whereas stations 39a and 40, also corresponding to the same date, are correctly referenced in Calvet (1907). For the purposes of representation on the map, we have used the coordinates of station 39a (corrected following Ryland 1969) because, in sharing numbering and a very similar depth (1000 m), we assume that they must be located very close together. D'Hondt (1973) also reported C. boryi from several localities in the NW of the Gulf of Biscay at 332-560 m depth, but again the lack of material, description and fi gures prevents us from verifying this record, which in any case should be considered doubtful due to the depth of the samples. Finally, C. cantabra was also reported by d'Hondt (1974, as C. boryi) from off Cape Peñas (Asturias) at 400-690 m depth (stn Thalassa W405; MNHN-IB-2008-7078). Note that localities for each species were not included in this publication, and the only record of this species was taken by us from d'Hondt's handwritten list. The locality is also included in the label of the sample. We have also revised a single sample from Avilés Canyon (Asturias) collected at 342 m depth (MHNUSC 10128) which is here designated as the holotype of C. cantabra sp. nov. (see Material examined and Fig. 10A-C, F-G).
In summary, at present C. cantabra sp. nov. is known only from the north of the Iberian Peninsula, ranging from 342 to 690 m depth, and perhaps even deeper if the record in Jullien (1882 as C. boryi) at 1037 m depth were of the same species (Fig. 12A).   (Levinsen, 1909) and Caberea cantabra sp. nov. B. Species of Hincksina Norman, 1903.

Discussion
The study of the type specimens and other material of E. angulata and M. tenella has enabled us to confi rm that they are different species. Even though Tilbrook et al. (2001) already demonstrated this 20 years ago, this seems to have escaped notice because most authors continue to confuse both species. Indeed, there is widespread confusion, since what is being cited as E. tenella actually corresponds to E. angulata. This species should be placed in the genus Arbopercula, as previously proposed by Tilbrook & Gordon (2015; erroneously as Arbocuspis, lapsus for Arbopercula), but to do so the diagnosis of the genus must be modifi ed. In addition, we highlight the original description of E. angulata, which describes the great variation in spine number and development, a characteristic that seems to be overlooked by later authors.
The revision of original material is essential today. Moreover, the use of the electron microscope enables a better characterization of species. Hincksina fl ustroides has been reported from numerous Iberian localities, both Atlantic and Mediterranean, but the revision of samples that we conducted shows that, in reality, these records correspond to fi ve different species with smaller distributions, two of them new to science (see Fig. 12B): H. fl ustroides in the Atlantic Ocean NW of the Iberian Peninsula, H. synchysia in the Eastern Iberian Mediterranean, H. elephantina sp. nov. in the Strait of Gibraltar, H. chimaera sp. nov. in the Alboran Sea and H. longispinosa also in the Alboran Sea, where it was already cited very recently. This last species was originally described from deep water in the Gulf of Cadiz. Present data reveal that the species is actually distributed on both sides of the Strait of Gibraltar, although its presence in the Mediterranean is limited to the Alboran Sea, which is considered to have a high affi nity with Atlantic and more specifi cally Lusitanian species (Harmelin & d'Hondt 1993;Maldonado & Uriz 1995;Bianchi & Morri 2000;Riesgo et al. 2019). To all these species we should add Hincksina sp. from southern Portugal (see Souto et al. 2014;Berning et al. 2021), which clearly belongs to the same group as all these species. Hincksina calpensis, also present in the area, differs in the scutum of the adult zooids. The western coast of Iberia seems to be dominated by H. fl ustroides and the eastern one by H. synchysia. Few data are available for the north coast, but in the south there are up to fi ve different species. A similar distribution has recently been reported for other genera such as Watersipora Neviani, 1896, with a single species on the western and northern coasts of Iberia [W. subatra (Ortmann, 1890)], another one on the eastern coast (W. cucullata (Busk, 1854]) and three species in the south (W. subatra, W. soleourum Vieira et al., 2014 andW. arcuata Banta, 1969) (see Reverter-Gil & Souto 2019).
Finally, a SEM revision of material from deep waters demonstrates that, at least in Iberian waters, C. boryi is not a species with a wide bathymetric range, but rather two different species, with the deeper material belonging to a new, different species.
The bryozoological fauna of the Iberian Peninsula is one of the best known in European waters. Our own unpublished compilation, based on dozens of articles published over the last century and a half, and the revision of hundreds of samples -both our own and those in museum collections -has yielded approximately 545 Recent species cited in this region BOE 2020, and unpublished data). In comparison, only 556 species have been registered so far in a larger area as studied as the Mediterranean Sea as a whole, where there is an extensive bibliography on bryozoans dating back more than 200 years (Rosso & Di Martino 2016). Moreover, all European waters combined have yielded 945 indigenous species according to Gordon et al. (2019). The description here of three new species and a new record in a supposedly so well-known area -altogether with the description of nine new species and another three new records in the last fi ve years (Ramalho et al. 2018(Ramalho et al. , 2020a(Ramalho et al. , 2020b(Ramalho et al. , 2022Reverter-Gil et al. 2019b;Reverter-Gil & Souto 2021b) -underlines the continued need for purely taxonomic and faunal works: they are key pillars to develop well-designed and useful biodiversity conservation policies (e.g., Wägele et al. 2011;Higgs 2017;Thomson et al. 2018). It is important that the publications include accurate descriptions and/or good fi gures, as well as reference material, to allow future authors to revise previous records. To publish acritic lists of species is of little help for this. Taxonomy and collections are essential to know which species we are really dealing with. Without this information, all subsequent comments are worth little.
Our present results extend our knowledge on the bryozoan fauna in Iberian waters and in Europe as a whole.