Redescription of some species of Bryozoa described by J . Jullien and L . Calvet in the NE Atlantic

Five species described by Jullien and/or Calvet from the NE Atlantic are redescribed and stabilized by typifi cation: Hippothoa amoena, Schizoporella confusa, S. jullieni, S. ovum and Smittia guernei. Three new synonymies are established: Schizoporella jullieni with Hippothoa amoena, S. ovum with Escharina alderi, and Escharella pseudopunctata with Smittia guernei. A new trypostegid genus, Pulpeirina gen. nov., is erected for H. amoena. Three new combinations are introduced.


Introduction
Towards the end of the nineteenth century several oceanographic expeditions by the research vessels Travailleur, l'Hirondelle and Talisman collected samples in the Northeast Atlantic.The resulting bryozoan studies were published by Jullien (1882Jullien ( , 1883)), Jullien & Calvet (1903) and Calvet (1906aCalvet ( , 1906bCalvet ( , 1907)).In total, about 135 new species were described in these papers, but most of them have not subsequently been redescribed according to current standards.Therefore, it is necessary to give accurate new descriptions, based on the original material and SEM examinations, of the species described by J. Jullien and/or L. Calvet, a task that has recently been started by various authors (e.g., Reverter-Gil & Fernández-Pulpeiro 1999, 2005;Reverter-Gil et al. 2011, 2015;Souto et al. 2011;Berning 2013).
In the present paper we redescribe five further species, which are stabilized by typification.The goal is to provide a better understanding of these species so as to ensure easier future identification and to give a clearer appreciation of their taxonomic affinities.Lectotypes have been selected from the original syntypes, three new synonymies are established and three new combinations are introduced.A new trypostegid genus is erected for one of the redescribed species.

Material and Methods
Types and other material stored in the following museums have been studied: Muséum national d'Histoire naturelle, Paris (MNHN), Musée Océanographique, Monaco (MOM) and the Natural History Museum, London (NHMUK).
The samples were examined with a stereo microscope and uncoated material was photographed with Zeiss EVO LS15 and Inspect S50 scanning electron microscopes, with a back-scattered electron detector in low variable vacuum mode.Measurements were taken with the software ImageJ ® on SEM photographs.

Etymology
This genus is dedicated to Dr. Eugenio Fernández-Pulpeiro, who died in September 2014.He was one of the people responsible for restarting bryozoology in Spain, but also our director, colleague and friend, and above all our best example as a researcher as well as a person.Gender feminine.(Jullien & Calvet, 1903)

Description
Colony encrusting, unilaminar.Autozooids oval to hexagonal, only slightly convex, separated by shallow grooves.Frontal wall smooth, translucent in young zooids, marked with slight transversal undulations; perforated by numerous, evenly spaced, small pseudopores rimmed in surface, prolongated in oblique tubules.One to several rows of pores distal to zooidal orifice.Pores joined by short tubules, visible in transparent walls, forming somewhat radial series running to margins of zooid.Orifice cleithridiate, not terminal; anter transversely oval, surrounded by thin rim; separated from shallow, concave poster by short, pointed condyles and pair of small notches.Edge of poster formed by pair of presumed vestigial costae fused in saggital plane, leaving small central pore and narrow slit.Spines absent.Communication via small, basal pore chambers.Ovicells terminal, prominent, cleithral.Large ooecium formed by distal kenozooid with no visible frontal part (Ostrovsky et al. 2009;Ostrovsky 2013).Ectooecium smooth, evenly perforated by small pseudopores prolongated in tubules, with small central umbo and sometimes growth lines.Ovicell aperture dimorphic, larger than in infertile autozooids, formed by proximal border of ooecium and poster of zooidal orifice; closed by zooidal operculum.Small, irregularly shaped kenozooids (without any orifice) may be present filling small spaces between autozooids.Zooeciules and avicularia absent.Ancestrula not observed.

Remarks
Hippothoa amoena was described by Jullien & Calvet (1903) for three samples collected on shells at two stations near Pico Island, Azores, at depths between 130 and 318 m.The original material is now kept at MOM and a lectotype is designated here.The name of the species appeared in the original paper with two different spellings: 'amoena' and 'amaena'.Both include a ligature and are therefore incorrect according to Art. 32.5.2 of the ICZN Code (ICZN 1999).We select here the first spelling and correct it to amoena (Arts 19.3,24.2.3 and 32).Hippothoa amoena was later reported from its original area between 80 and 115 m by Calvet (1907) andby d'Hondt (1975) between 80 and 1260 m, but this material was not found in MNHN or MOM, so perhaps it has been lost.
Schizoporella jullieni was described by Calvet (Jullien & Calvet 1903), also from material collected at the Azores (strait between Pico and Faial Islands, at a depth of 130 m).The original material is also kept at MOM and a lectotype is designated here.Schizoporella jullieni does not seem to have been reported since its original description.However, Gautier (1962: 121) stated that S. jullieni must be a synonym of Trypostega claviculata (Hincks, 1884), a species that he reported from the Strait of Sicily.This synonymy was subsequently assumed by Arístegui Ruiz (1984) when citing T. claviculata from the Canary Islands.These records differ from S. jullieni in the presence of interzooidal avicularia and the absence of the vestigial suboral costae.A revision of Gautier's original material proves that it belongs to a new species of Trypostega, which will be described in a future paper.
A revision of the type material of H. amoena and S. jullieni demonstrated that they correspond to the same species.Taking into account that H. amoena was described first in the original paper, and reported twice since its original description, we decide here to give precedence to this name, according to Art. 24 of the ICZN Code.
The porous gymnocystal frontal wall, the suboral pair of vestigial costae, and the absence of oral spines in H. amoena are all characters reminiscent of those in several taxa, such as the recently described genus Vitrimurella Winston et al., 2014, some fossil Trypostegidae, or even some Catenicellidae Busk, 1852.However, catenicellids differ most obviously by their erect, flexible and jointed colonies.Vitrimurella differs, among other features, by the ovicell.In the type species of the genus, V. lata (Smitt, 1873), and two other species, the ovicell is small, cap-like, immersed in the distal part of the maternal zooid, and therefore quite different from the large, prominent and uniformly perforated ovicell of H. amoena.In two other species of the same genus, V. gemina (Tilbrook, 2006) and V. lepida (Hayward, 1988), the ooecia are also prominent but possess a median suture, perhaps suggesting that they belong in a different Table 1.Measurements (in mm) of Pulpeirina amoena (Jullien & Calvet, 1903) (e.g., Trilophora Lang, 1916, Boreasina Voigt & Hillmer, 1983and Grammothoa Voigt & Hillmer, 1983) mainly by the absence of zooeciules or avicularia and the dimorphic orifices.Finally, Boreas Morris, 1980 also lacks avicularia, as H. amoena, but it lacks suboral vestigial costae and dimorphic orifices.
Moreover, H. amoena differs from all these taxa by the pseudopores of the frontal wall, which are not simple perforations, as can be seen in young, translucent zooids.The perforations in the surface of the zooids are formed during the calcification of the frontal wall (Fig. 3A).As the calcified layer thickens, the pseudopores extend to form short, oblique tubules.Groups of these oblique tubules are joined together by basal, horizontal tubules arranged forming a somewhat radial series, presumably reaching the zooidal margins (Fig. 3B, E).In some cases there is also a median, longitudinal horizontal tubule (Fig. 3C).The ovicells, however, do not possess horizontal tubules.The horizontal tubules do not seem to actually be excavated in the calcified layer, but rather to be an open space between the calcified layer and the underlying membranous ectocyst (Fig. 3D).If this is the case, this space represents a vestigial hypostegal coelom.This possibility, together with the radial arrangement of the horizontal tubules and the pair of suboral vestigial costae, is perhaps reminiscent of an ancient spinocystal frontal shield.Nevertheless, study of more fresh material is necessary to clearly establish the structure of the frontal wall in H. amoena.Although some related species may have tubular pseudopores (e.g., Trypostega sp.nov.or Vitrimurella spp., L.M. Viera pers.comm.Sep.2014), they are not radially arranged.On the other hand, it will be difficult to prove whether fossil Trypostegidae had radial tubular pseudopores.
Therefore, we erect a new genus, Pulpeirina, for H. amoena, and we tentatively place it in the family Trypostegidae.The structure of the frontal wall in Pulpeirina amoena comb.nov., perhaps reminiscent of an ancient spinocystal frontal wall, may support the suggestion that the Trypostegidae evolved from a pliophloeine ancestor (Gordon 2000).
Pulpeirina amoena comb.nov. is only known with certainty from around Pico Island, Azores, between 80 and 318 m depth (up to 1260 m according to d'Hondt 1975).However, we have found another sample, labeled as Smittia deguernei and presumably collected from the NW Iberian Peninsula, which corresponds to the same species.Smittia guernei was described by Jullien & Calvet (1903) for two samples collected off the NW coast of the Iberian Peninsula at a depth of 90 m.One of the samples (MOM INV-22518) matches the original description and is here designated as the lectotype of the species (see below).The other sample (MOM INV-22580) corresponds to P. amoena comb.nov.(see Fig. 3).Taking into account the evident differences between both species, it seems impossible that Jullien, who originally described both of them in the same paper (or even any other researcher), could have made such a mistake of identification.It seems more probable that the samples were mixed up during their preservation or during their subsequent revision by Calvet, when he undertook the responsibility of finishing the study of the samples collected by the Hirondelle.Therefore, we consider that the geographical origin of this sample of P. amoena comb.nov. is also doubtful.

Remarks
Busk (1856) described Alysidota alderi, a species with zooids in linear series, imperforate ovicells and rare adventitious avicularia.Some years later, he described Lepralia barleei for a single multiserial colony encrusting a shell collected in Shetland, with ovicells with peripheral pores and lacking avicularia (Busk 1860).
Norman (1869) was the first author who considered these two species to be synonymous while reporting A. alderi from Shetland at depths between 50 and 170 fathoms (90-310 m).He also stated that the type specimens of both species were in his collection.Hincks (1880) assumed this synonymy to be correct and transferred the species to the genus Schizoporella.Finally, Marcus (1940) transferred Schizoporella alderi to the genus Escharina.Type and other material labeled as A. alderi is conserved at the NHMUK, but no original material of L. barleei seems to exist.Although the sample NHMUK 1911.10.1.1197is labeled as the type of 'Escharina alderi var.barleei', it contains two colonies on stones, so they do not match the original description of the species.As no original material of L. barleei is preserved and the original description is very vague, it is not possible at present to confirm or reject the proposed synonymy.
Schizoporella ovum was described by Jullien (1882) for an undeterminate number of dead multiserial specimens, with ovicells with peripheral pores and lacking avicularia, collected off the NW Iberian Peninsula at a depth of 392 m.Three type specimens are now kept at NHMUK and MNHN; the one at NHMUK is here designated as the lectotype.Calvet (1907) reviewed the original record by Jullien, stating that the original material consisted of "trois échantillons sur coquilles".This author also reported two further colonies, also on shells, collected by the Talisman 1883 cruise from an unrecorded locality at a depth of 340 m; this material (MNHN 3770), although belonging to the same species, is not part of the type series.Schizoporella ovum does not seem to have been rediscovered since then, but it was transferred to the genus Schizomavella by Reverter-Gil & Fernández-Pulpeiro (2001) following their revision of the type material at Paris.The type material of Schizoporella ovum is identical to the sample NHMUK 1911.10.1.1197('Escharina alderi var.barleei'), so they belong to the same species.Furthermore, S. ovum may be somewhat similar to Lepralia barleei as originally described.On the other hand, the autozooids of S. ovum are identical to the autozooids of E. alderi (see Figs 4-5), as was stated by Hincks (1880: 244) for uniserial and multiserial colonies of S. alderi.Differences between these two species do not appear to be significant.Schizoporella ovum (and L. barleei) develop multiserial colonies, whereas Escharina alderi forms uniserial chains of zooids.According to Norman (1869), both forms of growth may even share the same substrate.In S. ovum (as in L. barleei) avicularia have not been reported, and in E. alderi they are rare.Therefore, their presence is perhaps related to the uniserial mode of growth, but in any case, their presence or absence should not be used to differentiate species.In E. alderi the development of a sporadic umbo in the ovicell and in the autozooids has been reported, while it is absent in the material of S. ovum.Its presence may be related to a different degree of calcification, with no phylogenetic significance.Finally, the ovicell was described as imperforate in E. alderi, while in S. ovum (and L. barleei) it has a series of septular pores around the basal periphery.However, SEM images of the type material of E. alderi demonstrate that the ovicell really has several basal pores (Fig. 4E), similar to those present in the types of S. ovum (Fig. 5D).It should be stated that in the genus Escharina, as in all members of the family Escharinidae, the ovicell has been described as imperforate (see Hayward & Ryland 1999;Tilbrook 2006), but the pores in the ovicell of E. alderi, as well as in other species of the family (see Bock 2014), indicate that the diagnosis of the family should be amended.
In conclusion, we propose here that Schizoporella ovum (and probably also Lepralia barleei) are junior synonyms of Escharina alderi, differing mainly in their mode of growth.

Remarks
Smittia guernei was described by Jullien & Calvet (1903)   Smittia guernei does not seem to have been reported under that name since its original description.Reverter-Gil & Fernández-Pulpeiro (2001) reviewed the original record, without studying the original material, and stated that the species probably belongs in Escharella.The species was reported by d'Hondt (1974) as "Mucronella peachi ?" from a Thalassa locality at the Avilés Canyon, N Iberian Peninsula (see Material examined section).
Recently, Souto et al. (2007) described a new species, Escharella pseudopunctata, from material collected in the English Channel at an unrecorded depth.Examination of the lectotype of S. guernei reveals that E. pseudopunctata is actually a junior synonym.For a complete description of the species and further discussion of its generic assignment see Souto et al. (2007: 368-371).
There are some minor differences between the types of S. guernei and E. pseudopunctata.In the former, the ovicell is globular, recumbent on the distal autozooid (Fig. 6C), but in the latter the ovicell is almost entirely immersed in the distal autozooid (see Souto et al. 2007: fig.4a, c).However, in the Thalassa material the ovicell may be recumbent or immersed in the same colony; this is rather frequent in several other genera of cheilostomatous Bryozoa (A.Ostrovsky, pers. comm. May 2015).Moreover, the secondary orifice seems more markedly bell-shaped in E. pseudopunctata (Fig. 6D; see also Souto et al. 2007: fig. 3d-e).Finally, zooids of E. pseudopunctata are slightly larger than those of S. guernei.
Escharella guernei comb.nov.was collected off the northern Iberian Peninsula at a depth of 90 m and at an unrecorded depth between 280 and 650 m, as well as from an unrecorded locality in the English Channel.

Description
Colony encrusting, multilaminar, forming irregular crusts.Autozooids oval to irregularly polygonal, separated by very fine sutures on shallow grooves in young autozooids; boundaries becoming indistinct in later ontogeny.Frontal shield thick, finely granular, slightly convex, bordered by single series of marginal, round pores; rapidly covered by secondary calcification.Primary orifice orbicular to dropshaped, about as wide as long.Anter distinctly denticulate, with 16 to 25 rounded denticles.Proximal border almost occupied by shallow sinus flanked by two shoulders.Condyles small, inconspicuous, sloping towards sinus.No oral spines.Zooids rapidly developing a tuberculate peristome formed by 2 to 7 stout knobs, granular at apex but frequently broken in examined material; two lateral lobes most frequent and most developed; distal knobs may be absent or smaller.Some zooids may also have a single suboral umbo, smaller that the rest.One (occasionally two) avicularia present in most zooids, placed laterally, at midlength on zooidal margin and projecting beyond zooidal boundaries or lateral to orifice.Rostrum triangular, directed laterally or distolaterally; crossbar complete, without columella.Ovicells subimmersed, acleithral, with ectooecium formed by distal autozooid.Ectooecium covered by secondary calcification except smooth central area.Ovicell opening above zooidal orifice.

Remarks
Schizoporella confusa was described by Calvet (1906b;see also Calvet 1907) for several colonies growing on Lophohelia (=Lophelia) collected at a depth of 717 m near Cape Spartel (Strait of Gibraltar).The original material is now preserved at MNHN and we here designate a lectotype from that material.The species does not seem to have been recorded since its original description.The encrusting colonies, the zooids having only marginal pores, the distally denticulate orifice, with sinus and condyles, the well-developed peristome, the adventitious avicularia, and the ovicell with an exposed area of endooecium are all characters allowing the inclusion of the species in the genus Rhynchozoon Hincks, 1895.Rhynchozoon confusum comb.nov. is characterized by the tuberculate peristome formed by 2 to 7 stout knobs, the lack of a suboral avicularium, and the lateral, triangular avicularium.Among the Atlantic-Mediterranean species of Rhynchozoon, only R. pseudodigitatum Zabala & Maluquer, 1988 may possess well-developed oral nodules, but this species is distinguished, among other features, by the suboral avicularium having an uncinate process defining a pseudospiramen.
for two samples collected off the NW coast of the Iberian Peninsula at a depth of 90 m.In the collections of MOM there are two slides, labeled as types of S. deguernei [sic!], collected at the same station, but only one of them (MOM INV-22518) matches the original description and figure of the species, and it is therefore designated here as the lectotype.The other specimen (MOM INV-22580) corresponds to P. amoena comb.nov.(see above).