European Journal of Taxonomy 157: 1-17

http://dx.doi.org/10.5852/ejt.2015.157

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

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2015 • Reverter-Gil O. & Souto J.

Research article

um:lsid:zoobank.org:pub:7FDDCF57-lF93-4266-B0F7-0486344055DE

Redescription of some species of Bryozoa described by

J. Jullien and L. Calvet in the NE Atlantic

Oscar REVERTER-GIL 1 * & Javier SOUTO 12

1 Departamento de Zooloxia e Antropoloxia Fisica, Facultade de Bioloxia,

Universidade de Santiago de Compostela, 15782 Santiago de Compostela, Spain.

2 Institut fur Palaontologie, Fakultat fur Geowissenschaften, Geographie und Astronomie,

Geozentrum, Universitat Wien, Althanstrasse 14, 1090 Vienna, Austria.

* Corresponding author: [email protected]

2 Email: [email protected]

* urn:lsid:zoobank.org:author:001DFC48-0F01-43AF-90EC-452BE6954DDF

2 urn:lsid:zoobank.org:author:E14FC8A9-40B5-4BF6-8551-3F2AlD991463

Abstract. Five species described by Jullien and/or Calvet from the NE Atlantic are redescribed and

stabilized by typification: 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.

Keywords. Azores, Iberian Peninsula, Strait of Gibraltar, new genus, Trypostegidae.

Reverter-Gil O. & Souto J. 2015. Redescription of some species of Bryozoa described by J. Jullien and L. Calvet

in the NE Atlantic. European Journal of Taxonomy 157: 1-17. http://dx.doi.org/10.5852/ejt.2015.157

Introduction

Towards the end of the nineteenth century several oceanographic expeditions by the research vessels

Travailleur , THirondelle and Talisman collected samples in the Northeast Atlantic. The resulting

bryozoan studies were published by Jullien (1882, 1883), Jullien & Calvet (1903) and Calvet (1906a,

1906b, 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 F. Calvet, a task that has recently been started by various authors (e.g., Reverter-Gil &

Fernandez-Pulpeiro 1999, 2005; Reverter-Gil et al. 2011, 2015; Souto et al. 2011; Beming 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. Fectotypes 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.

European Journal of Taxonomy 157: 1-17 (2015)

Material and Methods

Types and other material stored in the following museums have been studied: Museum national d’Histoire

naturelle, Paris (MNHN), Musee Oceanographique, Monaco (MOM) and the Natural History Museum,

London (NHMIJK).

The longitudes of the sampling stations of the Travailleur , Talisman and / ’Hirondelle cruises, published

by Jullien(1882,1883), Jullien&Calvet( 1903) and Calvet( 1906a, 1906b, 1907), were initially measured

with reference to the Paris meridian. They have here been corrected to the Greenwich meridian (see

Ryland 1969: 238).

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.

Results

Order Cheilostomata Busk, 1852

Infraorder Ascophorina Levinsen, 1909

Superfamily Hippothooidea Busk, 1859

Family Trypostegidae Gordon, Tilbrook & Winston in Winston, 2005

Genus Pulpeirina gen. nov.

um:lsid:zoobank.org:act:50502833-8EFA-4F91-ACE2-6831A613CECC

Diagnosis

Colony encrusting. Frontal wall gymnocystal, perforated by tubular pseudopores prolonged by sub¬

horizontal tubules, arranged radially. Orifice sinuate. Suboral vestigial costae fused in saggital plane,

leaving a narrow slit and a small central pore. Oral spines absent. Communication via small, basal pore-

chambers. Ovicell globular, evenly perforated by small, tubular pores. Orifice dimorphic, closed by

zooidal operculum. Zooeciules and avicularia absent.

Type species

Hippothoa amoena Jullien & Calvet, 1903.

Etymology

This genus is dedicated to Dr. Eugenio Fernandez-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.

Pulpeirina amoena (Jullien & Calvet, 1903) comb. nov.

Figs 1A-E, 2A-E, 3A-E; Table 1

Hippothoa amoena Jullien & Calvet, 1903: 88, pi. 10, fig. 9.

Schizoporella jullieni Calvet in Jullien & Calvet, 1903: 142, pi. 17, figs la-b.

? Smittia guernei Jullien & Calvet, 1903: 103 pars.

? Hippothoa amoena - Calvet 1907: 424. — d’Hondt, 1975: 579.

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Material examined

Lectotype of Hippothoa amoena (here designated)

AZORES: Hirondelle st. 247, Pico Island, 38°24’ N, 28°0E26” W, 30 Aug. 1888, 318 m (MOM INV-

22578).

Paralectotypes of Hippothoa amoena

AZORES: Hirondelle st. 226, Pico-Faial Channel, 38°31’19” N, 28°34 , 31” W, 14 Aug. 1888, 130 m

(MOM INV-22579, MOM INV-22581).

Lectotype of Schizoporella jullieni (here designated)

AZORES: Hirondelle st. 226, Pico-Faial Channel, 38°31’19” N, 28°34 , 31” W, 14 Aug. 1888, 130 m

(MOM INV-22502).

Paralectotypes of Schizoporella jullieni

AZORES: same data as lectotype, several small fragments covered with glue (MOM INV-22523); same

data as lectotype, an encrusting colony, together with Chorizopora sp. and Hippothoa sp. (MOM INV-

22524).

Other material examined

SPAIN: Hirondelle st. 56, 43 o 38 , 30 ,, N, 6 o 08 , 16” W, N Iberian Peninsula, 4 Aug. 1886, 90 m, Jullien

coll., originally labeled as “ Smittia deguernei J. Jullien” (MOM INV-22580).

Fig. 1 . Lectotype of Hippothoa amoena Jullien & Calvet, 1903 (MOM INV-22578). A. Some autozooids.

B. Group of irregular kenozooids. C. Primary orifice. D. Ovicell. E. Distal dietella.

European Journal of Taxonomy 157: 1-17 (2015)

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.

Fig. 2. Pulpeirina amoena (Jullien & Calvet, 1903) comb. nov. A. Autozooids (MOM INV-22502,

lectotype of Schizoporella jullieni Calvet in Jullien & Calvet, 1903). B. Same, ovicellate and non-

ovicellate zooids. C. Autozooids; note the distal dietella (MOM INV-22580). D. Same, primary orifice.

E. Same, ovicells.

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Table 1 . Measurements (in mm) of Pulpeirina amoena (Jullien & Calvet, 1903) comb. nov.

and paralectotypes). SD = standard deviation; N = number of measurements.

(lectotypes

Mean

Minimum

Maximum

Autozooid length

0.626

0.1024

0.463

0.797

Autozooid width

0.399

0.1172

0.244

0.670

Orifice length

0.096

0.0105

0.080

0.117

Orifice width

0.101

0.0099

0.082

0.127

Ovicell orifice length

0.102

0.0065

0.096

0.110

Ovicell orifice width

0.132

0.0100

0.121

0.148

Ooecium length

0.293

0.0191

0.270

0.330

Ooecium width

0.360

0.0334

0.309

0.394

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) and by 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 Aristegui 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

European Journal of Taxonomy 157: 1-17 (2015)

genus. Hippothoa amoena also differs from fossil Trypostegidae provided with suboral vestigial costae

(e.g., Trilophora Lang, 1916, Boreasina Voigt & Hillmer, 1983 and 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. 3 A). 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

Fig. 3. Pulpeirina amoena (Jullien & Calvet, 1903) comb. nov. (MOM INV-22580). A. Simple pores in

a young, marginal autozooid. B. Tubular pores arranged radially. C. An autozooid with radially arranged

tubular pores and a medial horizontal tubule. D. Inner view of an autozooid showing the tubular

pseudopores and horizontal tubules. E. Detail showing tubular pores joining the horizontal tubule.

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

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.

Superfamily Schizoporelloidea Jullien, 1882

Family Escharinidae Til brook, 2006

Genus Escharina Milne Edwards, 1836

Escharina alderi (Busk, 1856)

Figs 4A-E, 5A-E; Table 2

Alysidota alderi Busk, 1856: 311, pi. 9, figs 6-7.

? Lepralia barleei Busk, 1860: 143, pi. 26, figs 1-2.

Schizoporella ovum Jullien, 1882: 16, pi. 15, figs 36-37.

Schizoporella alderi - Hincks 1880: 243, pi. 36, figs 9, 9a, 10.

Schizoporella ovum - Jullien 1883: 512, pi. 15, figs 36-37. — Calvet 1907: 422.

Escharina alderi - Hayward & Ryland 1999: 228, figs 94c-d; 96.

Schizomavella ovum - Reverter-Gil & Fernandez-Pulpeiro 2001: 120.

Material examined

Lectotype of Schizoporella ovum (here designated)

SPAIN: Travailleur st. 40, 44°05 , 00” N, 07 o 14 , 46” W, NW Iberian Peninsula, 15 Aug. 1881, 392 m,

Jullien coll., one colony on a shell, figured specimen (NHMUK 1899.7.1.2348).

Paralectotypes of Schizoporella ovum

SPAIN: same data as lectotype (MNHN 1025, MNHN 2347).

European Journal of Taxonomy 157: 1-17 (2015)

Other material examined

NORWAY: Alysidota alderi , Busk coll., one uniserial colony, with some avicularia but no ovicell, on a

shell (NHMUK 1899.7.1.2143).

UNITED KINGDOM: Alysidota alderi , Shetland, 1859, Busk coll., paratype (NHMUK 1899.7.1.2144);

Alysidota alderi , Shetland, 1859, Busk coll., type (NHMUK 1899.7.1.2145); Escharina alderi var.

barleii , Shetland, Busk coll., marked as ‘Type?’ (NHMUK 1911.10.1.1197); Escharina alderi , Scotland,

22 Oct. 1958, st. 12, 52 m, Ryland coll. (NHMUK 1994.3.14.8).

LOCALITY UNKNOWN: Schizoporella ovum , Talisman 1883, 340 m, no further data, Calvet coll.

(MNHN 3770).

Description

Colony encrusting, composed of linear series of autozooids, or unilaminar, multiserial. Autozooids oval

to hexagonal, separated by sutures. Lrontal shield convex, granular, uniformely perforated by some

25 small rounded pores, often inconspicuous, plus a row of small marginal pores. Distolateral vertical

walls with several dietella. Primary orifice as long as wide. Inner distolateral orifice rim with immersed.

Fig. 4. Escharina alderi (Busk, 1856). A. Group of autozooids (NHMUK 1911.10.1.1197, type of

Alysidota alderi Busk, 1856). B. Same, ovicellate zooid; note the distal marginal pores in the ovicell.

C. Same, detail of a broken ovicell. D. Same, primary orifice. E. Ovicellate zooid and one adventitious

avicularium in a uniserial colony (NHMUK 1899.7.1.2145, paratype of Alysidota alderi Busk, 1856).

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Table 2. Measurements (in mm) of Escherina alderi (Busk, 1856) (lectotype and paralectotypes of

Schizoporella ovum Jullien, 1882). SD = standard deviation; N = number of measurements.

Mean

Minimum

Maximum

Autozooid length

0.640

0.0634

0.546

0.751

Autozooid width

0.442

0.0673

0.331

0.528

Orifice length

0.118

0.0083

0.106

0.130

Orifice width

0.119

0.0086

0.106

0.133

Ooecium length

0.260

0.0138

0.239

0.268

Ooecium width

0.297

0.0209

0.266

0.310

very narrow shelf. Anter slightly horseshoe-shaped, proximal margin straight, with U-shaped sinus, as

deep as wide, occupying one third of proximal border. Primary orifice encircled by smooth, broad band

of gymnocystal calcification. Oral spines absent. A low, conical umbo may be developed proximal to

sinus. Avicularia rare, lateral to orifice, sporadically present in uniserial colonies, apparently absent

in multiserial colonies. Rostrum triangular, raised, directed proximally or proximo-laterally. Ovicell

globular, recumbent on distal autozooid and partially covering orifice of maternal zooid. Surface

granular, with single series of septular pores around basal periphery and sometimes raised into a central

umbo. Ancestrula not observed.

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 1911.10.1.1197

is 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 echantillons 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 & Fernandez-Pulpeiro (2001) following their

revision of the type material at Paris.

European Journal of Taxonomy 157: 1-17 (2015)

The type material of Schizoporella ovum is identical to the sample NHMUK 1911.10.1.1197 JEscharina

alderi var. barleef), 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 Hinclcs (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

Fig. 5. Schizoporella ovum Jullien, 1882. A. Entire colony (NHMUK 1899.7.1.2348, lectotype).

B. Same, primary orifice. C. Same, a broken ovicell showing the marginal pores. D. Ovicellate and non-

ovicellate zooids (MNHN 2347, paralectotype). E. Dietella (MNHN 1025, paralectotype).

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Table 3. Measurements (in:

SD = standard deviation; N

mm) of Escharella guernei (Jullien & Calvet, 1903) comb. nov. (lectotype).

= number of measurements.

Mean

Minimum

Maximum

Autozooid length

0.499

0.0262

0.457

0.549

Autozooid width

0.361

0.0288

0.317

0.427

Orifice length

0.109

0.0046

0.096

0.117

Orifice width

0.098

0.0050

0.088

0.107

Ooecium length

0.242

0.0057

0.238

0.246

Ooecium width

0.275

0.0014

0.274

0.276

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 ma inl y in their mode of growth.

Superfamily Lepralielloidea Vigneaux, 1949

Family Romancheinidae Jullien, 1888

Genus Escharella Gray, 1848

Escharella guernei (Jullien & Calvet, 1903) comb. nov.

Fig. 6A-D; Table 3

Smittia guernei Jullien & Calvet, 1903: 103, pi. 13, fig. 1.

Escharellapseudopunctata Souto, Fernandez-Pulpeiro & Reverter-Gil, 2007: 368, figs 3-4.

? Mucronellapeachi - d’Hondt 1974: 30 (not Johnston, 1847).

Material Examined

Lectotype of Smittia guernei (here designated)

SPAIN: Hirondelle st. 56, 43°38’30” N, 06°08’16” W, NW Iberian Peninsula, 90 m, 4 Aug. 1886,

Jullien coll., originally labeled as “ Smittia deguernei J. Jullien” (MOM INV-22518).

Holotype of Escharella pseudopunctata

FRANCE: English Channel, Pourquoi Pas? st. 101, Jun. 1914 (MNHN 20063).

Paratypes of Escharella pseudopunctata

FRANCE: same data as holotype (MNHN 20064).

Other material examined

SPAIN: Thalassa st. W403, 43°56.2’ N, 5°39.3’ W, Aviles Canyon, N Iberian Peninsula, 280-650 m,

10 Oct. 1970, det. d’Hondt, labeled as “ Mucronella peachi (?)” (MNHN 7069); same data as preceding,

species not marked on original label (MNHN 7070).

European Journal of Taxonomy 157: 1-17 (2015)

Diagnosis

Colony encrusting unilaminar, multiserial. Zooids hexagonal to oval, with smooth, slightly convex frontal.

Pores do not perforate, as tubules run from pores to margins of zooid. Secondary orifice campanulate.

Primary orifice with small, quadrangular lyrula and small, rounded condyles. Six distal spines, retained

in ovicellate zooids. Avicularia not present. Ovicell globular, recumbent on distal autozooid.

Remarks

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. 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).

Fig. 6. Escharella guernei (Jullien & Calvet, 1903) comb. nov. A. Autozooids (MOM INV-22518,

lectotype). B. Same, primary orifice. C. Same, ovicellate zooids. D. Ovicellate and non-ovicellate

zooids; note the ovicells, recumbent or imm ersed (MNHN 7069).

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Smittia guernei does not seem to have been reported under that name since its original description.

Reverter-Gil & Fernandez-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 “ Mucronellapeachi ?” from a Thalassa locality at the Aviles 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

Cha nn el

Superfamily Celleporoidea Johnston, 1838

Family Phidoloporidae Gabb & Horn, 1862

Genus Rhynchozoon Hincks, 1895

Rhynchozoon confusum (Calvet, 1906) comb. nov.

Fig. 7A-E; Table 4

Schizoporella confusa Calvet, 1906b: 217.

Schizoporella confusa - Calvet 1907: 422, pi. 27, figs 12-13.

Material Examined

Lectotype (here designated)

MOROCCO: Talisman 1883 st. 10, 35°26 , 00” N, 06 o 4&46” W, off Cape Spartel, Strait of Gibraltar,

717 m, 10 Jun. 1883, Calvet coll., some fragments detached from a coral (MNHN 4007).

Paralectotypes

MOROCCO: same data as lectotype (MNHN 494, MNHN 1007, MNHN 1008, MNHN 2354).

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 drop¬

shaped, 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

European Journal of Taxonomy 157: 1-17 (2015)

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.

Fig. 7. Rhynchozoon confusum (Calvet, 1906) comb. nov. A. Young autozooids in the colony margin;

note the presence of small adventitious avicularia (MNHN 2354, paralectotype). B. Older zooids with

indistinct margins and well-developed oral knobs (MNHN 494, paralectotype). C. Primary orifice

(MNHN 4007, lectotype). D. Same, ovicell. E. Detail of an adventitious avicularium (MNHN 494,

paralectotype).

REVERTER-GIL O. & SOUTO J., Redescription of some species of Bryozoa

Table 4. Measurements (in mm) of Rhynchozoon confusum (Calvet, 1906) comb. nov. (lectotype and

paralectotypes). SD = standard deviation; N = number of measurements.

Mean

Minimum

Maximum

Autozooid length

0.883

0.1488

0.703

1.000

Autozooid width

0.681

0.0494

0.611

0.785

Orifice length

0.157

0.0130

0.143

0.179

Orifice width

0.162

0.0134

0.138

0.181

Ovicell length

—

0.341

0.344

Ovicell width

—

0.321

0.340

Avicularium length

0.168

0.0214

0.131

0.193

Avicularium width

0.067

0.0086

0.056

0.078

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

Hinclcs, 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.

Other species of Rhynchozoon , as for instance R. haha Hayward, 1988 from the Indo-West Pacific

region or R. sexaspinatum Winston & Woollacott, 2009 from Barbados, may also develop umbos around

the orifice, but the former is distinguished from R. confusum comb. nov. by its round orifice with a

bowl-shaped sinus and knob-like condyles, and the large frontal avicularia; it differs from the latter by

its keyhole-shaped orifice, with a large, rounded sinus and very large condyles, and three distinct types

of avicularia.

Acknowledgements

We are grateful to P. Lozouet (MNHN) and to M. Spencer Jones (NHMUK) for their valuable assistance

during visits and the loan of material, and to M. Bruni (MOM) for the loan of material. We tender special

thanks to L.M. Vieira and J. Winston for communicating unpublished data. Thanks are also due to the

staff of the Servizo de Microscopia Electronica, Universidade de Santiago de Compostela, for the SEM

photographs. Finally, two anonymous reviewers provided suggestions to improve the manuscript.

This work was partially supported by the project “ Fauna Iberica: Briozoos II (Familia Cribrilinidae

- Familia Waters iporidae)” (CGL2010-22267-C07-02), co-financed by the Ministerio de Economia y

Competitividad, Spanish Government, and FEDER. JS thanks the Austrian Science Fund (FWF, Lise

Meitner Program, grant M1444-B25) for financial support.

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Manuscript received: 14 August 2015

Manuscript accepted: 21 October 2015

Published on: 30 November 2015

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, De nm ark