European Journal of Taxonomy 44: 1-25

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ISSN 2118-9773

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2Q13 • Beming B.

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Research article

um:lsid:zoobank.org:pub:F7FD3319-AD9D-4DBB-9755-C541759CQD66

New and little-known Cheilostomata (Bryozoa, Gymnolaemata)

from the NE Atlantic

Bjorn BERNING

Geoseienee Collections, Upper Austrian State Museum, Welser Str. 2Q, 4Q6Q Feonding, Austria

Email: [email protected]

um:lsid:zoobank.org:author:7A351E42-FFD7-44A3-B3DE-CF5251B3A3Fl

Abstract. Based on newly designated type material, four poorly known NE Atlantic cheilostome

bryozoan species are redescribed and imaged: Cellaria harmelini d’Hondt from the northern Bay of

Biscay, Hippomenella mucronelliformis (Waters) from Madeira, Myriapora bugei d’Hondt from the

Azores, and Characodoma stmngulatum, occurring from Mauritania to southern Portugal. Moreover,

Notoplites saojorgensis sp. nov. from the Azores, formerly reported as Notoplites marsupiatus (Jullien),

is newly described. The genus Hippomenella Canu & Bassler is transferred from the lepraliomorph

family Escharinidae Tilbrook to the umbonulomorph family Romancheinidae Jullien.

Keywords. Bryozoa, Cheilostomata, Macaronesia, new species, taxonomy.

Berning B. 2Q13. New and little-known Cheilostomata (Bryozoa, Gymnolaemata) from the NE Atlantic. European

Journal of Taxonomy 44: 1-25. http:/dx. doi.org/lQ.5852/eit.2Ql 3.44

Introduction

Compared with the number of publications on the phylum Bryozoa from the Mediterranean Sea, the

subtropical and warm-temperate NE Atlantic faunas have been fairly neglected during the last decades.

There are only a handful of recent papers that deal with relatively few species from the NW African and

Iberian continental shelf and open ocean islands (e.g., Aristegui 1985; Harmelin & d’Hondt 1992; Fopez

de la Cuadra & Garcia-Gomez 1993, 1996; F6pez-Fe 2QQ6; Beming 2Q12). In fact, the most important

scientific cmises and collections were made in the late 19th to early 2Qth century (e.g., Jullien 1883;

Jullien & Calvet 19Q3; Calvet 19Q7, 1931), resulting in the introduction of most species prior to the

use of scanning electron microscopy (SEM), which is cmcial for the detection of subtle yet important

morphological differences between bryozoan species. It is, therefore, of vital importance to clearly

define the historical species in order to assess the diversity, ecological requirements, and geographic

distribution of bryozoan species in the NE Atlantic.

Owing to the research efforts of individual scientists, and particularly during the ongoing project

“Fauna Iberica: Briozoos”, with the aim to inventory the entire Iberian bryozoan fauna, a number of

NE Atlantic species have already been redescribed or newly introduced very recently (e.g., Reverter-

Gil & Femandez-Pulpeiro 1999a, b; Reverter-Gil et al. 2QQ9, 2Q12; Souto et al. 2Q1Q, 2Q11). Other

modem studies revising selected genera have revealed that, due to the presence of undifferentiated

European Journal of Taxonomy AA\ 1-25 ( 2013 )

species complexes alone, the bryozoan diversity in the NE Atlantic may be considerably underestimated

(Harmelin 1978; Harmelin & Aristegui 1988; Reverter-Gil & Femandez-Pulpeiro 1996; Beming &

Kuklinski 2008; Beming et al. 2008).

The present paper deals with another four poorly known and ill-defined species, which are redescribed

based on their type specimens: Cellaria harmelini d’Hondt, 1973 from the northern Bay of Biscay,

Hippomenella mucronelliformis (Waters, 1899) from Madeira, Myriapora bugei d’Hondt, 1975 from the

Azores, and Characodoma strangulatum (Calvet, 1906), apparently occurring from southern Portugal

to Mauritania. In addition, a species from the Azores, which has previously been recorded as Notoplites

marsupiatus (Jullien, 1883), is newly described.

Material and methods

The analysed material is preserved in the collections of the Natural History Museum in London

(NHMUK), Manchester Museum (MM), the Museum National d’Histoire Naturelle in Paris (MNHN),

the Musee Oceanographique de Monaco (MOM), and the Madeiran Museu Municipal (Historia Natural)

in Funchal (MMF). Bryozoan type specimens were digitally photographed at the NHMUK using a

LEO 1455VP SEM, and at the MNHN using a Tescan VEGA SEM. Both machines allowed imaging

the uncoated specimens with back-scattered electrons in the low vacuum mode. Morphometries were

made on these micrographs using the image software ImageJ. Bryozoan systematics follow the working

classification of D.P Gordon (pers. comm. 2011), who is currently developing the classification scheme

for eventual use in the revised Treatise on Invertebrate Paleontology.

Results

Suborder Neocheilostomina d’Hondt, 1985

Superfamily Buguloidea Gray, 1848

Family Candidae d’Orbigny, 1851

Genus Notoplites Harmer, 1923

Notoplites saojorgensis sp. nov.

um: Isid: zoobank, org: act: 19 A50609-B149-46AC-9A17-237D31A95867

Fig. 1, Table 1

Notoplites marsupiatus (Jullien, 1883) - Calvet 1931: 69.

Non Scrupocellaria marsupiata - d’Hondt 1975: 556, figs 14-16 (part or whole).

Differential diagnosis

There are only two other NE Atlantic Notoplites species that have a marginally fimbriated scutum

covering the entire opesia: N. clausus (Busk, 1884) and V. marsupiatus (Jullien, 1883). N. saojorgensis

sp. nov. differs from these in generally lacking distal orificial spines, in having wider zooecia and larger

scuta, and in having ooecia with a triangular proximomedian fenestra. Moreover, in comparison with N.

marsupiatus, zooids in the new species are much larger, the ooecia are distinctly shorter, and the orifice

is wider than long. Additional differences to N. clausus are that the intemodes are more robust, that the

lateral avicularium is clearly visible in frontal view, and that there are more and longer fissures between

marginal branches in the scutum.

Etymology

The species is named after its type location, Sao Jorge Island (Azores archipelago).

BERNING B., NE Atlantic Bryozoa

Type Material

Holotype

MNHN 4163, a large free eolony, Princess Alice Stn 1349, 1250 m, 19 Aug. 1902, 38°35’30” N

-28°05’45” W (Azores, off south-eentral Sao Jorge Island), on sandy mud of voleanie origin.

Fig. 1. Notoplites saojorgensis sp. nov. (MNHN 4163, holotype). A. Optieal image of eolony showing

the poreelain white zooeeia. B. Overview of eolony showing several intemodes and braneh bifureations.

C. Proximal part of the eolony with numerous, elosely joined rhizoids emanating from proximal and

abfrontal zooids, forming supporting stalks. D. Abfrontal side of intemode with two rhizoids. E.

Autozooids at braneh bifureation; note the basal part of the greatly enlarged spine at the base of the

seutum (lower arrow) and the single spine of normal size in the median zooid at the bifureation point

(upper arrow). F. Close-up of autozooid with distolateral and proximal avieularium. G. Ovieellate

zooids; note the proximomedian, aeutely triangular window in the ooeeium. Seale bars: A, B = 1 mm;

C, D = 200 pm; E, G = 100 pm; F = 50 pm.

European Journal of Taxonomy AA\ 1-25 (2013)

Table 1. Measurements (in |Lim) of skeletal eharaeters of Notoplites saojorgensis sp. nov., taken from the

holotype (MNHN 4163). SD = standard deviation, N = number of measurements.

Mean

Range

Zooid length

932

748-1104

Zooid width

331

275-378

Orifiee length

86-103

Orifiee width

126

106-144

Seutum length

277

251-314

Seutum width

290

234-335

Ooeeium length

322

313-337

Ooeeium width

282

248-302

Avieularium length

62-87

Avieularium width

33-45

Paratypes

Same loeality information as holotype: MOM 42 0877, numerous detaehed intemodes; MOM 42 1213,

several intemodes embedded in Canada balsam on a glass slide.

Description

Colony ereet, jointed, diehotomously branehing, forming a delieate tuft of up to 6.5 em in height,

attaehed by numerous rhizoids that form a supporting stalk; zoarium poreelain white in dried state

(Fig. lA-C). Branehes formed by two series of alternating autozooids, mostly with 4-9 autozooids

between bifureations, zooids opening on one side only (Fig. ID). Branehing points eomposed of a single

proximomedian zooid and two distolateral ones, with the nodes developing immediately distal to the

latter zooids by breakage of the narrow proximal parts of the subsequent zooids (Fig. IE). Autozooids

very elongate, narrowest and tubular proximally, widening distally with the distal half of zooids turned

outwards at an angle of e. 40°, usually (mueh) less than half of total autozooid length oeeupied by the

seutum/membranous area and orifiee (Fig. lE-F); skeletal surfaee smooth, eonvex, zooids separated by

a distinet groove, forming a zigzag line on the abfrontal side (Fig. ID). Rhizoids produeed from a small

pore near proximal end on the abfrontal side of some zooids, elosely approximated to form a single stalk

(Fig. IC-D).

Orifiee slightly raised and displaeed towards the outer zooid margin (Fig. IE), very well-defined, wider

than long and widest in distal third, straight proximal margin formed by distal seutum, opereulum

thiekly selerotised (Fig. IF). Seutum eonvex, about as wide as long, distally branehed with the braneh

ends merging with the eireum-opesial gymnoeyst and eovering the membranous area like a well-fitting

lid, branehes separated by some 14-20 narrow fissures of variable length, some reaehing almost half of

seutum length (Fig. IF); seutum originating from a broad ineonspieuous base near the inner distal zooid

margin just proximolateral to orifiee, with the seutum base oeeasionally bearing a single greatly enlarged

spine (Fig. IE); a smaller single spine distal to orifiee only in median zooids at branehing points (Fig.

IE).

Two adventitious avieularia of about the same size and shape per zooid (Fig. IF): one situated lateral

to orifiee and forming the outer zooid margin, with the triangular rostrum direeted outwards and the

frontal area more or less distally; the other one situated on a raised eystid just proximally to seutum, with

BERNING B., NE Atlantic Bryozoa

the distally slightly downcurved rostrum directed proximally; mandibles in both avicularia hinged on

inconspicuous condyles formed by the distal part of an immersed cryptocystal shelf in the proximal area.

Ovicell globular, prominent, the inner half of ooecium resting on the proximal part of the distal zooid,

slightly longer than wide, surface somewhat uneven and with a proximomedian elongate-triangular

fenestra pointing distally, the proximolateral ooecial margins reaching towards mid-distance of lateral

orifice rim, ovicell opening well arched above orifice (Fig. IG).

An ancestrula was not observed.

Remarks

There are two species that are closely related to Notoplites saojorgensis sp. nov., the types of both of

which have recently been figured and described by Souto et al. (2011): N. marsupiatus (Jullien, 1883),

originally recorded from the continental slope ofNW Spain, and A. clausus (Busk, 1884), sampled some

400 km west of the Azores. All three species share the same type of fimbriated scutum.

Notoplites clausus was later considered to be a junior synonym of N. marsupiatus by Jullien (1888),

Calvet (1907) and other workers. However, Souto et al. (2011) showed that they are distinct species and

reinstated A. clausus. In turn, Notoplites saojorgensis sp. nov., which was identified as A. marsupiatus by

Calvet (1931), is clearly different from these two. The specimens imaged as Scrupocellaria marsupiata

by d’Hondt (1975), without any further comments, are very close to A. clausus owing to the presence

of a distal oral spine, small lateral avicularia, and fewer fissures in the scutum than A. saojorgensis

sp. nov. However, d’Hondt (1975) recorded the specimens from eight stations: four within the Azores

archipelago, two from stations west of the western Azorean islands (i.e., closer to the type location of A.

clausus), and two from stations several hundred kilometres north of the Azores. While the exact origin

of the figured specimens cannot be retraced, it is likely that two or more species were combined under

the name A. marsupiata in that study. These samples could not be analysed during the present project,

as most of them are not present at the MNHN.

Notoplites saojorgensis sp. nov., as most other Notoplites spp., occurs at bathyal depths. The colonies

are anchored in soft sediments via a single stalk formed by numerous long rootlets.

Infraorder Flustrina Smitt, 1868

Superfamily Cellarioidea Fleming, 1828

Family Cellariidae Fleming, 1828

Genus Cellaria^Wis & Solander, 1786

Cellaria harmelini d’Hondt, 1973

Fig. 2, Table 2

Cellaria harmelini d’Hondt, 1973: 374, pi. 1, figs 2-4.

Material examined

Lectotype (here designated)

MNHN 6888, Thalassa Stn 436, 4 Aug. 1967, northern Bay of Biscay, 47°56’ N - 07°52.7’ W, 360 m.

Paralectotype (here designated)

MNHN 6902, Thalassa Stn 451, 4 Aug. 1967, northern Bay of Biscay, 47°57.5’N- 07°50.7’W, 358 m,

on hy droids.

European Journal of Taxonomy AA\ 1-25 (2013)

When introducing this species, d’Hondt (1973) did not indicate type material. The subsequent listing

of the “holotype” and “paratype” of Cellaria harmelini without further information in a catalogue of

the potential type specimens kept at the MNHN (Tricart & d’Hondt 2009) violates Article 73.1.3. of the

ICZN Code (ICZN 1999). Therefore, the lectotype and paralectotype are formally designated here for

the “holotype” and “paratype”, respectively, of Tricart & d’Hondt (2009).

Description

Colony erect, flexible, dichotomously branching, about 3 cm in height and composed of 6-7 intemodes;

intemodes slender (Fig. 2A), cylindrical, composed of 6 alternating longitudinal series of zooecia (4 in

one horizontal row), basal intemodes some 2-4 mm long, distal ones up to 8-9 mm in length, with 1 or

2 fertile zones per intemode producing distinctly thicker {ca. 0.7-0.9 mm) segments, zooecia separated

by a thin ridge flanked by a shallow groove on each side. Autozooecia in sterile segments elongated

subhexagonal (Fig. 2B), successive zooecia in a series not in direct contact and being well spaced,

contact with lateral neighbours in the next series after the alternating one along a straight boundary,

proximal zooecial margin V-shaped. Fertile zooecia elongated hexagonal, about as long and wide as

sterile zooecia, in direct contact with the successive zooecia in the same series and with those in the

alternating neighbour-series, proximal zooidal margin usually straight (Fig. 2D). Central cryptocyst

depressed, framed by a pair of longitudinal ridges reaching from the distolateral zooecium margin lateral

to opesia toward proximal part of zooecium, where they level with the depressed cryptocyst (Fig. 2B);

cryptocyst smooth in autozooecia, in fertile zooecia granular along zooecial margins, particularly so

proximally and distally. Opesia semi-elliptical, broader than long, proximal edge convex with a pointed

Fig. 2. Cellaria harmelini d’Hondt, 1973 (MNHN 6902, paralectotype). A. Overview of an infertile

intemode with a distal avicularium. B. Close-up of an autozooecium. C. An avicularium situated distal

to an autozooecium. D. Ovicellate zooecia, one with a distal avicularium at left. Scale bars: A= 200 pm;

B, C = 50 pm; D = 100 pm.

BERNING B., NE Atlantic Bryozoa

Table 2. Measurements (in jam) of skeletal eharaeters of Cellaria harmelini d’Hondt, 1973, taken from

the paraleetotype (MNHN 6902). SD = standard deviation, N = number of measurements.

Mean

Range

Fength of sterile zooeeia

611

551-659

Width of sterile zooeeia

300

270-325

Opesia length

86-107

Opesia width

129

115-144

Ooeeium length

40-42

Ooeeium width

109

104-115

Avieularium length

123

120-125

Avieularium width

158

154-162

Diameter of sterile braneh part

442

376-505

Diameter of fertile braneh part

690

dentiele near eaeh eomer that is bent at a 90° angle and direeted terminally, distal opesial rim finely

beaded, proximal rim oeeasionally very faintly beaded (Fig. 2B-D).

Opening of fully formed ooeeia roughly trapezoidal (Fig. 2D), very short and broad, situated at the

maternal zooid’s extreme distal end, with the proximal eryptoeyst of the sueeeeding zooeeium or

avieularium forming the distal margin of the ooeeium, slightly less than proximal half oeeluded by a

square plate with small lateral dentieles extending from its lateral edges; “ooeeial” opening in zooeeia

situated in proximal parts of the thiekened fertile intemodal zones, subrounded, plaeed between opesia

and distal zooeeium margin and therefore less distal than in fully formed ooeeia.

Avieularium of fistulosa-ty^Q (Fig. 2C), situated direetly distal to a zooeeium, proximal border straight

and the distolateral outline forming two-thirds of a full eirele in sterile intemodal zones and with a rather

square outline in fertile zones, both forms slightly broader than long. Proximal eryptoeyst in autozooeial

zones short and smooth, extremely redueed distolaterally, thin margin granular all around, proximal

eryptoeyst in fertile zones slightly more extensive and granular; rostmm semieireular, arehed and very

slightly raised with straight proximolateral edges extending into the subeireular opesia, direeted distally

or distolaterally.

Perforations for rhizoidal kenozooids were not observed.

Remarks

Almost all of the speeies’ eharaeters were already given by d’Hondt (1973) in the original deseription

in Freneh, whieh are largely transeribed here. However, whereas some SEM photos of general zoarial

features were given in that publieation, images of eleaned zooeeia were not provided.

The new measurements given above oeeasionally differ from those in the original aeeount. Whereas

some of these differenees may be due either to the few intemodes available in the present study and/or

to the use of different measuring teehniques (optieal with a mieroseope vs. digital from SEM images),

other values in the original paper seem umealistie. For instanee, signifieant morphometrie differenees in

zooeeium and opesia dimensions between fertile and sterile zooeeia, as stated by d’Hondt (1973), eould

not be affirmed in the paraleetotype studied.

European Journal of Taxonomy AA\ 1-25 (2013)

Cellaria harmelini was subsequently reported by Hayward (1978) and Hayward & Ryland (1978) from

several other stations along the northern shelf margin of the Bay of Biseay, whereas d’Hondt (1973)

reeorded the speeies also from a loeation in the southern Bay of Biseay (44°01.6’ N - 07°01.9’ W, 510-

630 m). Most samples were taken from gravelly sediment surfaees, and the speeies’ depth of oeeurrenee

ranges from 180 to 700 m.

The status of Cellaria harmelini subsp. tenuis d’Hondt, 1974, reeorded from the northern and north¬

western Spanish shelf edge, eannot be eommented on at present.

Infraorder Aseophorina Levinsen, 1909

“Grade” Umbonulomorpha Gordon, 1989

Superfamily Lepralielloidea Vigneaux, 1949

Family Romaneheinidae Jullien, 1888

Genus Hippomenella Canu & Bassler, 1917

Hippomenella Canu & Bassler, 1917: 41.

Hippomenella - Brown 1949: 517. —Harmer 1957: 1095. — Gordon 1984: 77. —Zabala & Maluquer

1988: 117.

Type species

Lepralia mucronelliformis Waters, 1899, by original designation.

Diagnosis (amended)

Colonies enerusting, budding intrazooidal. Frontal shield partly umbonuloid, with two or more rows

of lateral areolar pores eneireling an imperforate suboral area; basal pore ehambers with multiporous

septula. Orifiee with eondyles, oral spines present. Ovieell hyperstomial, eetooeeium unealeified,

endooeeial surfaee finely pitted in parts but devoid of any other struetures, not elosed by the opereulum.

Adventitious avieularia present, oeeasionally dimorphie.

Remarks

Sinee its introduetion, the genus Hippomenella has been assigned to a number of different families. The

eonfusion stems from the faet that the original generie diagnosis by Canu & Bassler (1917) ineluded

eharaeters of several superfieially similar, yet strueturally distinet, Reeent and fossil speeies, resulting

in a poorly defined genus. Although Brown (1949) notieed this problem and deseribed the leetotype of

Hippomenella mucronelliformis in great detail, the type speeies still remained relatively poorly known,

as SEM images have never been published to date.

The genus has previously been plaeed in the sehizoporelloid families Hippoporinidae Osbum, 1952

(Brown 1958: 62), Hippopodinidae Levinsen, 1909 (Gordon 1984: 77), Sehizoporellidae Jullien, 1883

(Gordon 1989:43; Gordon e/a/. 1994; Gordon & d’Hondt 1997:25), andveryreeently intheEseharinidae

Tilbrook, 2006 (D.R Gordon, pers. eomm. 2011). Tilbrook (2006) and Hayward & Winston (2011)

regarded Hippomenella as incertae sedis. After SEM analysis of the leetotype of H. mucronelliformis the

generie diagnosis is here revised, and the genus transferred to the lepralielloid family Romaneheinidae

Jullien, 1888 for reasons speeified below. Thus, Hippomenella is now being united with the genus

Waters (1899) initially eonsidered elosely related: the speeies epithet mucronelliformis refers to its

similarity with Mucronella cocczfrea Abildgaard, 1806, whieh today is plaeed in the romaneheinid genus

Escharoides Milne Edwards, 1836.

BERNING B., NE Atlantic Bryozoa

The presence of a ring scar (Fig. 3E), which defines the slightly reduced central umbonuloid part of

the frontal shield in the type species, is one reason for removing the genus from the lepraliomorph

Escharinidae to the umbonulomorph Romancheinidae. The previous assignment of Hippomenella to

lepraliomorph families was partly due to its original placing in the Hippoporininae by Canu & Bassler

(1917) [see the discussion on this subfamily and the family Hippoporinidae in Gordon (1984); both

taxa are now considered as synonyms of Phidoloporidae Gabb & Horn, 1862], Moreover, Brown (1949,

1952) and Gordon (1984) considered the New Zealand species Lepralia vellicata Hutton, 1873 to belong

to Hippomenella. Despite the apparently similar structure of the frontal shield at the zooidal surface,

Gordon (1984: 77) observed that the imperforate umbonuloid part of the shield of E. vellicata is extremely

reduced, which was considered by him to support a placing of the genus in the Eepraliomorpha. The

numerous pores along the marginal frontal shield of the above-mentioned species are all considered here

to be areolar pores, as more than one may contribute to avicularium formation. The presence of over

20 avicularia that are scattered on the surface of a single zooid of L. vellicata {cf. Brown 1949: 519)

supports this interpretation.

The other major source of confusion was a misconception of the ooecium structure of Hippomenella.

The ooecium of H. mucronelliformis was not known when Canu & Bassler (1917) introduced the

genus. Instead they described the ooecia of species that may be placed in the morphologically similar

romancheinid genus Hippopleurifera Canu & Bassler, 1925 (see discussion in Hastings 1966), the

genotype of which (Eschara biauriculata Reuss, 1847) is characterised by a pair of lateral fenestrae in the

ooecium. Moreover, Brown (1949) considered the endooecium to be entirely perforated by pseudopores,

and, based on observations ofE. vellicata, Gordon (pers. comm. 2012) later regarded the ooecium type

of Hippomenella as schizoporelloid. Thus, a number of species with a range of ooecial structures and

morphologies were lumped together in Hippomenella. In contrast, in H. mucronelliformis the calcified

endooecium is not perforated by pseudopores (as in the schizoporelloid type) but merely superficially

pitted in the distolateral part (Fig. 3C, F), and it is lacking any other ooecial structures.

This ovicell type is, in turn, rather similar to the primary ooecium in the genus Escharoides, and

therefore yet another argument for placing Hippomenella in the Romancheinidae. Prior to being covered

by secondary calcification of the distal zooid, the endooecium in Escharoides shows a similarly pitted

surface as in H. mucronelliformis. This can best be seen in fossil specimens of Escharoides in which the

(presumably aragonitic) secondary calcification has vanished (see Beming 2006: figs 71, 73-79).

Comparative analyses of the frontal shield (e.g., the extent of the umbonuloid part) and the ooecium will

be vital in order to umavel the relationship between Hippomenella and Hippopleurifera. For instance, in

the genotype of Hippopleurifera almost the entire frontal shield is perforated by areolar pores, suggesting

that the umbonuloid part is extremely reduced. The ooecial fenestrae in Hippopleurifera supposedly result

from a thick but incomplete cover of secondary calcification produced by the distal zooid, whereas the

exposed endooecium is densely perforated by tiny pseudopores or superficial pits. In contrast, in several

other (fossil) species that have been assigned to Hippopleurifera the extent of the umbonuloid frontal

shield seems to be similarly extensive as in H. mucronelliformis, and there is no secondary calcification

from the distal zooid covering the imperforate endooecium. The occasionally occurring prominent ribs in

some of these species (e.g., in the Pliocene Eschara sedgwicki Milne Edwards, 1836) are probably entirely

of endooecial origin. Additionally, a third group comprises species with apparently schizoporelloid-like

ovicells, i.e., with pseudopores that penetrate the endooecium (e.g., L. vellicata, Hippomenella amaralae

Vieira et al., 2010, Hippopleurifera belizae Winston, 1984, Hippomenella ramula Hayward & Winston,

2011). A thorough revision of these taxa is, therefore, urgently needed but beyond the scope of the present

paper. For this reason, the generic diagnosis given above is rather conservative and may need to be

expanded again if species with other ooecial structures are included in Hippomenella.

European Journal of Taxonomy AA\ 1-25 (2013)

Information on the ancestmla has also been omitted from the generie diagnosis beeause 1 doubt that the

zooid identified and deseribed as sueh by Brown (1949), whieh looks similar to a mature autozooid but

with more proximally positioned avieularia (Fig. 3 A), is indeed the aneestrula. In the type speeimen (and

also observed in another eolony) the region immediately proximal to and around this zooid is oeeupied

by other autozooids, whieh probably eover and disguise the true aneestrula. In faet, remnants of spines

of the true aneestrula ean still be seen that are situated elose to the proximal margin of the first generation

autozooid and whieh are not entirely eovered by the overgrowing zooid. As all other romaneheinid taxa

have a tatiform aneestrula, 1 presume the same applies to Hippomenella.

Hippomenella mucronelliformis (Waters, 1899)

Fig. 3, Table 3

Lepralia mucronelliformis Waters, 1899: 11, pi. 3, figs 15, 21.

Lepralia mucronelliformis -Elorman 1909: 306.

Hippomenella mucronelliformis 1949: 513, figs 1, 2a,b,d,e.

Material examined

Lectotype

MM 3780 (here designated), Madeira (no further information provided as to the exaet loeation or depth),

on bivalve fragment, mounted on slide. Waters eolleetion.

Paralectotype

MMF 42297 (here designated), Madeira (no further information provided as to the exaet loeation or

depth), on bivalve fragment, mounted on slide, J.Y. Johnson eolleetion.

Other material

MM 3781, Madeira (no further information provided as to the exaet loeation or depth), opereula, in

Canada balsam on slide. Waters eolleetion; NHMUK 1947.8.12.1, Madeira (no further information

provided as to the exaet loeation or depth), two eolonies on bivalve shell fragments and several isolated

zooids mounted on slide, from the Canon J. de G. Barreto eolleetion.

Although Brown (1949) stated that the leetotype was ehosen by Norman (1909: 306), this author simply

mentioned that he had “seen the type”. This may not violate Artiele 74.5 of the ICZN Code (ICZN

1999), and may represent a valid designation of the leetotype. However, it is unelear exaetly what

speeimen Norman had seen: the eolony now kept at Manehester Museum or that from the Funehal

Museum. Both speeimens earry identieal labels and the handwriting of Waters, and are elearly syntypes.

Moreover, the slide from the MMF, whieh was eonsidered by Brown to eontain the type speeimen,

originally eomprised two eolonies, judging from the remains of glue in the slide eavity, whereas neither

Waters (1899) nor Norman (1909) explieitly mentioned how many speeimens they were referring to. A

small green label on MM 3780 indieates that this speeimen was figured by Waters (a single autozooid;

pi. 3, fig. 21), whereas a green and red label on MM 3781 denotes this speeimen (supposedly eontaining

opereula) as the figured type speeimen (PI. 3, Fig. 15). However, it is unelear from whieh eolony the

opereula were taken; more importantly, there is not a single opereulum present on the slide today.

They have either deeayed or been lost, as one edge of the eover slide is shattered. This speeimen must,

therefore, not be regarded as a type.

To eonelude, 1 disregard Brown’s (1949) statement that “the type” had been ehosen by Norman (1909),

and designate as leetotype of H. mucronelliformis the figured speeimen MM 3780. The only remaining

syntype speeimen on the slide of sample MMF 42297 beeomes the paraleetotype.

BERNING B., NE Atlantic Bryozoa

Fig. 3. Hippomenella mucronelliformis (Waters, 1899). A. Overview of autozooids in the leetotype (MM

3780); note the extremely long and slender mandibles of the small avieularia. B. Early astogenetie part

of the paraleetotype (MMF 42297), ineluding the zooid interpreted by Brown (1949) to be the aneestmla

(at lower left) but whieh is here eonsidered as the first autozooid; note that the avieularia in early

astogenetie zooids are proximally positioned and direeted. C. Autozooids and an ovieellate zooid at the

eolony margin; note that the forming endooeeium is not perforated by pseudopores and that the suboral

muero is absent in early ontogenetie zooids, forming only during later ontogeny (NHMUK 1947.8.12.1;

photo: M.E. Speneer Jones). D. Close up of orifiee (leetotype, MM 3780). E. Interior frontal shield with

the umbonuloid ring-sear framed by areolar pores (NHMUK 1947.8.12.1; photo: M.E. Speneer Jones).

F. Ovieellate zooid; note the superfieial pits on the distolateral endooeeium (NHMUK 1947.8.12.1;

photo: M.E. Speneer Jones). G. Eateral view of a zooid (distal is to the right), showing five multiporous

pore plates in the vertieal wall (NHMUK 1947.8.12.1; photo: K. J. Tilbrook). Seale bars: A = 400 pm; B,

C = 200 pm; D = 50 pm; E, F, G = 100 pm.

European Journal of Taxonomy AA\ 1-25 (2013)

Table 3. Measurements (in |Lim) of skeletal eharaeters of Hippomenella mucronelliformis (Waters, 1899),

taken from speeimens MM 3780 (leetotype), MMF 42297 (paraleetotype) and NHMUK 1947.8.12.1.

SD = standard deviation, N = number of measurements.

Mean

Range

Zooid length

755

618-922

Zooid width

640

100

441-820

Orifiee length

168

134-190

Orifiee width

133

113-168

Ooeeium length

303

275-350

Ooeeium width

342

310-370

Length of small avieularium

128

83-176

Width of small avieularium

41-89

Length of large avieularium

321

206-402

Width of large avieularium

133

97-157

Description

Colony enerusting, unilaminar, multiserial, forming small patehes. Zooeeia relatively large,

subhexagonal, widest at about mid-distanee, oeeasionally wider than long, separated by deep furrows

(Fig. 3A-C); eommunieation between zooids via up to 5 multiporous pore plates per neighbouring

zooid (Fig. 3G). Frontal shield slightly eonvex proximally, somewhat raising distally towards orifiee,

seeondary ealeifieation forming a blunt prominent suboral muero during ontogeny (Fig. 3C, D), surfaee

rugose to nodular, eentral drop-shaped area imperforate, demareated by a row of densely spaeed areolar

pores with radially aligned intervening ridges indieating the extent of the eentral umbonuloid part of

the frontal shield (Fig. 3C, E), 1-3 additional rows of widely-spaeed pores in the external area towards

zooid margin, partieularly abundant proximolateral of orifiee with 1-2 rows extending between orifiee

and distal zooeeial margin (Fig. 3C). Orifiee elongate oval, longer than wide and usually widest in distal

third, proximolateral edges rounded, the slightly eoneave proximal margin eovered by the overarehing

and distally pointing muero, blunt eondyles formed by a eontinuation and slight inbending of lateral

orifiee margins, direeted proximomedially and delimiting the proximal fourth or fifth of total orifiee

length (Fig. 3D); opereulum strongly selerotised with a pair of lateral ridges where museles attaeh;

usually 6 (range 4-7, in early astogenetie zooids up to 9) oral spines along distolateral orifiee margin,

the proximal pair slightly larger, two in ovieellate zooids (Fig. 3A, B, F).

Ooeeium hyperstomial, depressed globular, slightly wider than long, endooeeial surfaee fairly smooth,

imperforate but distolaterally with numerous small round pits (Fig. 3F), proximal ooeeial margin

eoneave, shallowly arehed, reaehing towards lateral orifiee rim, not elosed by the opereulum.

Avieularia adventitious, dimorphie, usually paired (Fig. 3F), oeeasionally single, rarely absent, situated

lateral or proximolateral to orifiee at zooid margin, direeted laterally or proximolaterally exeept in early

astogenetie zooids where the rostrum points proximally and the avieularia are more proximally positioned

(Fig. 3B); small avieularium with a relatively short, proximally ineurved rostrum, distally parallel-sided,

downeurved and ehute-like with open end, mandible thin and elongate, up to four times the length

of rostrum (Fig. 3A); often one avieularium or sometimes both greatly enlarged, their width usually

exeeeding length of smaller avieularia (Fig. 3C, F), positioned on a slightly enlarged perforated eystid,

rostrum proximally ineurved, distally thin and parallel-sided with an aeute downeurved tip, reaehing

BERNING B., NE Atlantic Bryozoa

over the lateral zooid’s frontal shield, mandible eonfined to rostrum; erossbar in both avieularium types

eomplete without eolumella, proximal opesia semieireular.

Aneestrula not observed.

Remarks

Apart from its type loeation at Madeira, H. mucronelliformis has been reported from NW Moroeeo

by Canu & Bassler (1925: 30) as well as from the Mediterranean Sea by Harmelin (1969: 1208, figs

6-9), Hayward (1974: 371), Zabala (1986: 407, text-fig. 134, pi. 6, figs B, C), and Zabala & Maluquer

(1988: 117, text-fig. 244, pi. 8, fig. H), from depths down to 200 m. However, most of these works

laek a thorough deseription and illustration, and beeause some of these reeords differ in a few aspeets

from the type, I have only ineluded the reeords from Madeira in the synonymy list. For instanee, in

at least some of the speeimens reeorded by Zabala (1986) and Zabala & Maluquer (1988) from the

Western Mediterranean, the suboral muero is not developed or only very redueed, as ean be seen in the

provided SEM images. In turn, the orifiee of the speeimen from the Eastern Mediterranean Sea imaged

by Harmelin (1969) has medially pointing eondyles. Therefore, these reeords need to be eheeked for

eonspeeifity based on SEM observations.

There are also several fossil speeimens that have been reeorded as H. mucronelliformis. Those from

the Plioeene of Sieily (Pouyet & Moissette 1992: 62, pi. 9, fig. 9) very mueh resemble the Reeent

type, whereas the Middle Mioeene speeimens from the Paratethys are similar but probably represent

a distinet speeies, as they have a slightly different orifiee shape and the proximal ooeeial margin is

rimmed (Zagorsek 2010: 168, pi. 143, figs 1-4). These reeords show that the genus Hippomenella has a

eonsiderable fossil history in the Paratethys, and also that the Atlantie-Maearonesian region may have

played an important role in aeting as a refuge for Paratethyan-Mediterranean bryozoan taxa during

the Messinian salinity erisis and/or the Pleistoeene temperature minima (e.g., Beming 2006). S im ilar

temporal and geographie distribution patterns were reeently reported in speeies of the eheilostome

gQX\QX?i Saevitella Bobies, 1956 and Ca/Zopor/fia Neviani, 1895 (see Beming 2012).

“Grade” Eepraliomorpha Gordon, 1989

Superfamily Sehizoporelloidea Jullien, 1883

Family Myriaporidae Gray, 1841

Genus Myriapora de Blainville, 1830

Myriapora bugei d’Hondt, 1975

Fig. 4, Table 4

Myriapora bugei d’Hondt, 1975: 585, figs 23, 25-28.

non Myriapora bugei-El Hajjaji 1992: 250, pi. 15, figs 14-15.

Material examined

Lectotype (here designated)

MNHN IB-2013-3, one dried eolony fragment (former part of MNHN 7481), Jean Charcot, Bia 9 ores

Stn 109, 20 Get. 1971, NW of Flores (Azores), 39°33’ N- 31°17’ W, 190-230 m.

Paralectotypes (here designated)

MNHN 7481, Jean Charcot, Bia 9 ores Stn 109, 20 Get. 1971, NW of Flores (Azores), 39°33’ N- 31°17’

W, 190-230 m; MNHN IB-2013-2, one dried eolony fragment (former part of MNHN 7492), Jean

Charcot, Bia 9 ores Stn 110, 20 Get. 1971, NW of Flores (Azores), 39°33’ N - 31°17.5’ W, 300-350 m;

MNHN 7482, 13 eolony fragments in ethanol, Jean Charcot, Bia 9 ores Stn 109, 20 Get. 1971, NW of

European Journal of Taxonomy AA\ 1-25 (2013)

Flores (Azores), 39°33’ N - 31°17’ W, 190-230 m; MNHN 7488, one eolony fragment in ethanol, Jean

Charcot, Bia 9 ores Stn 110, 20 Oet. 1971, NW of Flores (Azores), 39°33’ N - 31°17.5’ W, 300-350 m;

MNHN 7492, four eolony fragments in ethanol, Jean Charcot, Bia 9 ores Stn 110, 20 Oet. 1971, NW of

Flores (Azores), 39°33’N-31°17.5’W, 300-350 m.

Fig. 4. Myriapora bugei d’Hondt, 1975. A. Colony fragment with elosely spaeed branehes bifiireating at

a 90° angle from the main braneh (MNHN lB-2013-3, leetotype). B. Braneh segment with one whorl of

fertile zooeeia at top, identified by the larger dimorphie orifiee and the radial arrangement of pseudopores

(MNHN lB-2013-2, paraleetotype). C. Distal braneh with early ontogenetie autozooeeia (MNHN 7481,

paraleetotype). D. Close-up of an autozooeeial orifiee (MNHN lB-2013-2, paraleetotype). E. The

dimorphie orifiee of a maternal zooeeium (MNHN lB-2013-2, paraleetotype). Seale bars: A = 2 mm; B

= 200 pm; C = 300 pm; D = 50 pm; E = 100 pm.

BERNING B., NE Atlantic Bryozoa

Table 4. Measurements (in jam) of skeletal eharaeters of Myriapora bugei d’Hondt, 1975, taken

from speeimens MNHN IB-2013-3 (leetotype), MNHN 7481 (paraleetotype), and MNHN IB-2013-2

(paraleetotype). Braneh width was measured on relatively mature braneh regions; younger ones are

distinetly narrower. As skeletal boundaries are invisible on the eolony surfaee, ooeeium dimensions

eould not be measured. SD = standard deviation, N = number of measurements.

Mean

Range

Fongitudinal distanee between eentroids of orifiees

1337

113

1197-1541

Fateral distanee between eentroids of orifiees

741

643-908

Orifiee length in sterile zooeeia

227

216-240

Orifiee width in sterile zooeeia

199

183-211

Orifiee length in maternal zooeeia

227

Orifiee width in maternal zooeeia

266

Braneh diameter

1367

118

1176-1531

The original material available to d’Hondt (1975: 560) was fromBia 9 ores stations 109 and 110. However,

he did not indieate type material when introdueing Myriapora bugei, and the figured speeimen (figs 23,

25-28) was subsequently destroyed (J.-E. d’Hondt pers. eomm. 2012). For the same reasons as stated in

the Material seetion for Cellaria harmelini (see above), the type speeimens of M. bugei were invalidly

eonsidered as “holotype” and “paratypes” by Trieart & d’Hondt (2009), and are formally designated

here as leetotype and paraleetotypes, respeetively.

Description

Colony ereet, rigid, delieate branehing, produeing lateral offsets at an angle of about 90° at irregular

intervals (Fig. 4A). Branehes long (3-4 em), slender and eylindrieal, with the zooids opening all around.

Zooids large, radially arranged in distinet whorls of 4 (rarely 5), with all orifiees at same level and the

distal whorl ideally alternating (Fig. 4B-C); zooeeia elongate reetangular but boundaries invisible on

eolony surfaee. Frontal shield of mature zooeeia gently raising towards orifiee, giving distal branehes

an undulating morphology that is slightly redueed due to frontal ealeifieation during ontogeny (Fig. 4A-

C); frontal shield regularly perforated by numerous elongate pores in spindle-shaped depressions that

are longitudinally aligned (Fig. 4B), surfaee finely granular (Fig. 4D-E). Primary orifiee dimorphie; in

autozooids slightly longer than wide (Fig. 4D), anter horseshoe-shaped, widest slightly distal to mid-

distanee, proximolateral eomers rounded with the short shoulders slightly sloping towards a deep and

broadly U-shaped sinus oeeupying about two-thirds of total proximal width, eondyles eonspieuous,

broad and with rounded edges, not extending beyond shoulders of the proximolateral margin; orifiee

in ovieellate zooids D-shaped (Fig. 4E), of similar length as in autozooids but espeeially the proximal

margin distinetly broader, sinus a very shallow straight edge eomprising four-fifths of total proximal

width, proximolateral shoulders thus relatively short and fairly straight, eondyles similar to those in

autozooids; orifiee beeoming immersed by frontal ealeifieation and oeeasionally elosed during ontogeny

in proximal braneh regions, in whieh ease a small short peristome with a eentral pore remains. No oral

spines.

Ovieells relatively rare, present in all zooids of eertain whorls, indieated by a slightly thieker swelling

of this braneh region and by frontal pseudopores that are radially arranged from the proximal ooeeium

eentre (Fig. 4B, E), ooeeium mostly ineorporated into frontal shields of the distal zooids, surfaee as

frontal shield of zooeeia, boundaries invisible at eolony surfaee, ovieell aperture at an aeute angle to

frontal plane, elosure of the eleithral or subeleithral type.

European Journal of Taxonomy AA\ 1-25 (2013)

No avicularia.

An ancestmla was not present in the available material.

Remarks

Although the genus Myriapora eomprises only a few speeies globally, owing to their mostly ereet and

eonspieuous eolonies, and beeause of the ubiquitous Mediterranean speeies M. truncata (Pallas, 1766),

this bryozoan genus has reeeived a fair amount of attention (e.g., Viskova 1986; Beming 2007; Ferretti

et al. 2007; Rodolfo-Metalpa et al. 2010). Besides M. truncata and the polar M subgracilis (d’Orbigny,

1852), M. bugei is the only other known speeies from the northern Atlantie realm. The most obvious

differenee between M. bugei and M. truncata is that in the former the branehes are eomposed of distinet

whorls whereas zooeeia in the latter are ideally alternating, i.e., apparently arranged in spirals around the

braneh axis. The ooeeia and dimorphie orifiees of M. bugei are here deseribed and figured for the first time.

Originally deseribed from the Azores, the speeies has also been reeorded from the Great Meteor Bank

(Piepenburg & Muller 2004: 61), whieh is loeated some 800 km south of the arehipelago. However,

although the eharaeters are variable and overlapping to some extent, the populations from the Great

Meteor Bank, as well as from the nearby Hyeres and Irving seamounts, slightly differ from the type of M.

bugei in having branehes that predominantly eonsist of five zooids per whorl (oeeasionally four or six),

smaller autozooids, and a narrower autozooeeial sinus (pers. observation). Thus, espeeially eonsidering

the distanee between the Azores and the seamounts, it is likely that these populations represent distinet

speeies, although genetie studies may be needed to elarify this issue.

In the Azores, M. bugei was reported from depths between 190 and 1235 m (d’Hondt 1975). It oeeurs on

roeky, gravelly, sandy and muddy substrata growing on roeks, dead eorals and bivalve shells.

The speeimens El Hajjaji (1992) deseribed and figured as M. bugei from the Late Mioeene of NE

Moroeeo (Mediterranean Sea) are similar to the type but eertainly belong to a different speeies due to a

distinetly narrower sinus and larger eondyles. The status of other eoeval speeimens from NW Moroeeo

(Atlantie) eannot be assessed, as a detailed deseription or figures were not provided by Sefian et al.

(1999: 242).

Superfamily Mamilloporoidea Canu & Bassler, 1927

Family Cleidoehasmatidae Cheetham & Sandberg, 1964

Genus Characodoma Maplestone, 1900

Characodoma strangulatum (Calvet, 1906)

Fig. 5, Table 5

Myriozoum strangulatum Calvet, 1906: 158.

Myriozoum strangulatum - Calvet 1907: 427, pi. 26, figs 11-12.

Cleidochasma strangulatum - Harmelin 1977: 1067, text-fig. 25, pi. 2, figs 1, 4, 7. — Harmelin &

d‘Hondt 1992: 29.

Characodoma strangulatum 1999: 429.

Material examined

Syntype series

MNHN 492, a single eolony fragment on slide, Travailleur Dr. 49, 7 Aug. 1882, Canary Islands, 29° N -

16°08’ W, 3700 m; MNHN 493, a single eolony fragment on slide, Travailleur Dr. 38, 30 Jul. 1882, NW

Moroeeo, 34° 13’30” N - 07°43’ W, 636 m; MNHN 947, a single eolony fragment on slide, Travailleur

BERNING B., NE Atlantic Bryozoa

Dr. 38, 30 Jul. 1882, NW Morocco, 34°13’30” N - 07°43’ W, 636 m; MNHN 2471, a single colony

fragment on slide. Talisman Dr. 96, 15 Jul. 1883, off Cap d’Arguin (Mauritania), 19°18’ N - 18°0E W,

2330 m.

Other material

MNHN 15487, several eolony fragments, Balgim DW07,29 May 1984, SWPortugal, 36°46.1 ’ N-9°27’

W, 1139-1144 m; MNHN 19804, several eolony fragments, Balgim CP92, 8 Jun. 1984, NW Moroeeo,

34°24.3’N-7°30.3’W, 1182 m.

While elearly stating that the original material eomprised speeimens from the Travailleur (stations Dr.

38, 49) and Talisman (station Dr. 96) eruises, Calvet (1906, 1907) did not indieate type speeimens when

introdueing Myriozoum strangulatum. The listing of MNHN 2471 as “leetotype” by Trieart & d’Hondt

(2009) was obviously for mere euratorial purposes and violates Artiele 74.7.3 (see also Deelaration 44)

of the ICZN Code. This designation is thus eonsidered invalid. For reasons given below, I refrain here

from ehoosing as leetotype a speeimen from the syntype series.

Sampling positions of the Talisman and Travailleur eruises, the longitudes of whieh were initially

measured with referenee to the Paris meridian, have here been eorreeted to the Greenwieh meridian by

subtraeting 2°20’14” from the longitude originally given {cf. Ryland 1969: 238).

Description

Colony presumably eellarinelliform (sensu Rosso 1992). Rods up to about 1 em in length, eylindrieal to

oval in eross-seetion with eonstrietions at irregular intervals (Fig. 5A-E), whieh result from breakage and

regeneration of the eolony, or from a reduetion in the number of distally budded zooids; ramifieations

or rhizoids were not observed; in fully developed eolony parts, 3 zooids are simultaneously budded

distally, and the rods are eomposed of an abfr ontal side devoid of orifiees on about one-fourth of the total

perimeter (Fig. 5H), with the zooids opening at the latero-frontal sides (Fig. 5E) and being arranged in a

plaited manner along the rod axis, i.e., zooids aligned in 3 alternating longitudinal series on eaeh side of

a median frontal (zigzag) line; all zooids inelined at about 45° to rod axis, pointing towards the median

frontal line (Fig. 5E, G, J). The zooids in these 3 series are polymorphie and deerease in size towards

the median frontal line: the abfrontal side is exelusively eomposed of the extremely elongated proximal

parts of the latero-abfrontally positioned series of zooeeia of eaeh side (Fig. 5H), while the distal parts

of these zooeeia are bent around the rod axis and open along the lateral sides; zooeeium shape very

elongated hexagonal; the intermediate lateral series eonsist of distinetly shorter subrhomboidal zooeeia

(Fig. 5E, G, J), abutting distally against a zooeeium of the median series from the other side of the

median frontal line; zooeeia of the median series even shorter and subhexagonal (Fig. 5E, G), opening

at an angle of almost 90° with respeet to the latero-abfrontal zooeeia. In between zooids at the median

line, an orbieular opening is oeeasionally present that presumably marks the heterozooid from whieh a

rhizoid is produeed for eolony support (Fig. 5J).

All zooeeia separated by shallow grooves and indistinet meandering sutures owing to seeondary

ealeifieation during later ontogeny; frontal shield slightly eonvex, distally forming a swollen, salient

rim around distolateral orifiee, surfaee distinetly nodular (Fig. 5G); few areolar pores of variable shape,

usually situated in zooeeial eomers. Primary orifiee somewhat immersed, situated at quite a distanee to

the distal zooeeial margin (Fig. 5G), eleithridiate, slightly longer than wide, with a large suborbieular

poster eomprising about three quarters of a full eirele and a very broadly U-shaped anter of about one-

third of total orifiee length and over half of orifiee width, delimited by very short proximomedially

direeted eondyles (Fig. 5F); orifiee dimensions smaller in zooids of the median series than in those of

the lateral and latero-abfrontal series.

European Journal of Taxonomy AA\ 1-25 (2013)

Ovicells were not observed.

Avieularia adventitious, small, transversely elliptieal, usually two per zooeeium (Fig. 5E, G, 1-J):

one in the distolateral eomer that is eloser to the median frontal line, and another one usually at the

proximolateral zooeeial margin; the latero-abfrontal zooids may have another avieularium at the very

proximal zooeeial margin; eystid slightly raised during early ontogeny, later levelled by seeondary

ealeifieation. Rostrum mueh wider than long, semi-elliptieal, with a slightly raised smooth distal rim,

direeted distolaterally (distal avieularia) or proximolaterally (proximal avieularia); erossbar eomplete,

without eolumella; proximal unealeified area transversely oval.

BERNING B., NE Atlantic Bryozoa

Table 5. Measurements (in jam) of skeletal eharaeters of Characodoma strangulatum (Calvet, 1906),

taken from the syntype speeimens MNHN 492, 493 and 2471 (see text for morphometrie variability

between speeimens). Due to the eonvex eolony surfaee, zooeeial measurements of the extremely elongated

latero-abfrontal zooids eould not be taken. SD = standard deviation, N = number of measurements.

Mean

Range

Zooid length (median zooids)

447

421-468

Zooid width (median zooids)

375

356-396

Orifiee length (median zooids)

112

113-107

Orifiee width (median zooids)

103

96-107

Zooid length (lateral zooids)

587

523-648

Zooid width (lateral zooids)

368

304-422

Orifiee length (lateral zooids)

133

120-142

Orifiee width (lateral zooids)

120

116-131

Avieularium length

36-54

Avieularium width

56-78

Braneh diameter

776

678-860

Fig- 5 (page 18). Characodoma strangulatum (Calvet, 1906). A. Eatero-abfrontal view of MNHN 492

(Canary Islands) from the syntype series, whieh is affeeted by Bynesian deeay. B. Eateral view of

MNHN 493 (NW Moroeeo) from the syntype series, a mature eolony in whieh the zooeeia are frontally

thiekened by seeondary ealeifieation; the abfrontal part (to the left) and the autozooeeia from the opposite

side eannot be seen in this view. C. Eateral view of MNHN 2471 (Mauritania) from the syntype series;

the eonstrietion in this eolony was eaused by zooid regeneration after damage and does not mark a

eonventional eonstrietion in growth as seen in other speeimens (photo: P. Kuklinski). D. A long rod

with reversed polarity growth after damage at the eolony eentre, the lower half representing the primary

eolony from whieh the upper part was budded; in both parts growth eonstrietions oeeur that did not

result from eolony breakage (MNHN 19804, NW Moroeeo; photo: J. Souto). E. Close-up of the same

speeimen in lateral view showing ontogenetieally young zooids that laek seeondary ealeifieation; the

three differently positioned, polymorphie autozooids are indieated as “la” (elongated latero-abfrontal

zooids), “1” (lateral zooids of intermediate size), and “m” (small median zooids) (MNHN 19804, NW

Moroeeo; photo: J. Souto). F. Orifiee (MNHN 492, Canary Islands); the ealeium aeetate erystals growing

on the skeletal surfaee are the result of Bynesian deeay owing to the speeimen having been kept enelosed

in a wooden slide in relatively humid eonditions. G. Eate ontogenetie polymorphie autoozooids (MNHN

493, NW Moroeeo); see E for abbreviations. H. The eolony growth margin of another eolony of sample

MNHN 19804 (NW Moroeeo) with two zooids having been budded after a normal eonstrietion in eolony

diameter; view of the abfrontal side of the rod with the extremely elongated latero-abfrontal zooid; the

orifiee is to the left whereas its proximal margin is at the very right (arrow) (photo: J. Souto). I. Close

up of two avieularia (MNHN 19804, NW Moroeeo; photo: J. Souto). J. Yet another eolony of sample

MNHN 19804 (NW Moroeeo) showing the approximate median line at whieh zooids from both eolony

sides meet; note the orbieular opening of a kenozooid at the median line from whieh a rhizoid for eolony

attaehment presumably originates (arrow) (photo: J. Souto). Seale bars: A-C, E, H, J = 200 pm; D = 400

pm; F = 30 pm; G = 100 pm; I = 50 pm.

European Journal of Taxonomy AA\ 1-25 (2013)

Early astogenetic stages absent in the examined material.

Remarks

In a revision of the Mediterranean speeies of the genus Characodoma Maplestone, 1900, Rosso

(1999: 429) first assigned Myriozoum strangulatum to this genus when deseribing the elosely related

Pleistoeene speeies Characodoma reclinatum Rosso, 1999. Although the eharaeteristie ovieell eould not

be observed on any of the available material, the general zooidal and zoarial features of C. strangulatum

eorrespond to those of Characodoma.

However, there are serious problems eoneeming the speeifie taxonomie identity of C. strangulatum that

arise from the speeimens of the syntype series. 1) The four syntypes are small eolony fragments that

are very poorly preserved. Speeimen MNHN 947 is strongly affeeted by Bynesian deeay and eould not

be studied in detail. MNHN 492 and 493 are also affeeted to varying degrees but some eharaeters are

still visible, while in MNHN 2471 all openings are filled with sediment. 2) Important speeies-speeifie

eharaeters are either entirely laeking in these speeimens (e.g., ovieells) or are mostly obseured, sueh as

the primary orifiees, whieh are eoneealed by opereula in MNHN 493, or filled with sediment in MNHN

2471. Cleaning or bleaehing of the speeimens is presumably impossible without damaging or destroying

the fragile eolonies. Moreover, as they are glued onto slides, only a restrieted number of measurements

eould be taken (ef Table 5, and see below). 3) The speeimens are from widely distant loeations, with a

distanee of over 800 km between eaeh of the three sampling regions (Mauritania, Canary Islands, Gulf

of Cadiz). 4) Of the four speeimens of the syntype series, two (MNHN 492 and 2471) resemble the

original figure (Calvet 1907: pi. 26, fig. 11) owing to the presenee of eonstrietions typieal for the speeies.

Neither of these two eolonies ean be unequivoeally identified as the figured speeimen, maybe beeause

the eolony surfaee depleted by Calvet eannot be seen in the mounted speeimens, i.e., they were glued

onto the slides with the wrong side faeing upwards.

Thus, beyond the absenee of ovieells and early astogenetie stages from all material assigned to C.

strangulatum, the few small and poorly preserved speeimens of the syntype series do not allow this

speeies to be preeisely defined. Moreover, the great geographie range of distribution and distanee

between sampling regions raises the question of whether all syntype speeimens do indeed belong to

the same speeies. For instanee, the lateral zooids in speeimen MNHN 493 have slightly smaller orifiees

(mean OL 124, OW 113, N = 3) than the remaining syntype speeimens (mean OL 136, OW 122, N =

11), although the number of measurements is elearly too small to be statistieally reliable. In eontrast, the

speeimens reported from NW Moroeeo by Harmelin & d’Hondt (1992; MNHN 19804), i.e., from the

same region as MNHN 493 and MNHN 497, differ from the original suite of C. strangulatum in having

distinetly larger orifiees (mean OL 168 pm, OW 146 pm, N = 10), both measured in lateral and latero-

abfrontal zooids (MNHN 19804, Fig. 5D, E, H, J).

Therefore, 1 eonsider the speeimens of the syntype series as insulfieiently preserved and laeking

important diagnostie eharaeters for the speeies to be preeisely defined, and it eannot be ruled out that

the syntype series eomprises more than one distinet speeies. Henee, 1 will not designate a leetotype

from the syntype series but suggest instead that a neotype should be ehosen as soon as newly sampled

material from either one of the three original sampling sites beeomes available. In the present study, C.

strangulatum is eonsidered sensu lato, as is its geographie range and depth distribution.

Another ineongruity eoneems the avieularium: in the original figure of C. strangulatum (Calvet, 1907:

pl. 26, fig. 12) the avieularia are depleted as having elongated and rather triangular rostra, instead of the

extremely short and broad, semi-elliptieal rostra reported in other speeimens assigned to this speeies.

Harmelin (1977) aseribed this differenee to a possible intraspeeifie variability between populations

inhabiting different environments. However, in none of the speeimens of the “syntype series” are

BERNING B., NE Atlantic Bryozoa

crossbars visible, either beeause they are broken and lost, or beeause the mandibles are still in plaee,

eoneealing the skeletal stmetures underneath. Another possibility is that they are truly absent. In any

ease, it seems likely that, in the absenee of any visible erossbars, Calvet deeided that they should be,

as in most other bryozoans, perpendieular to the longest avieularian dimension, and haphazardly added

erossbars to the figure. That the avieularium is, indeed, broader than long in C. strangulatum is indieated

by the slightly raised distal margin of the rostrum, while simple erossbars spanning the longest dimension

are present in well-preserved speeimens from eentral and northwestern Moroeeo that were assigned to

C. strangulatum by Harmelin (1977) and Harmelin & d’Hondt (1992), respeetively (Fig. 51).

Characodoma strangulatum has been reeorded from depths between 600 and 3700 m and its geographie

range is provisionally eonsidered to streteh from southern Portugal along the Moroeean shelf, and via

the Canary Islands to the Mauritanian slope, i.e., some 2000 km in N-S direetion. The speeies oeeurs on

muddy substrata and is, as other eongenerie speeies {cf. Rosso 1999), likely to be anehored in the fine

sediment by rhizoids. The rhizoids themselves were never observed, not even in well-preserved speeimens

from the Gulf of Cadiz reeovered during the Balgim eruise (Harmelin & d’Hondt 1992). This may not

be surprising, as Hirose (2011) reeently showed that, in eertain speeies, rhizoids may be translueent and

extremely delieate stmetures, and may thus be among the first tissue to deeay after sampling. However,

the orbieular kenozooidal openings situated along the frontal midline of the C. strangulatum eolonies

do suggest this mode of attaehment (Fig. 5J). This means that the tentaeles of the median zooids faee

towards the substratum when everted, whieh has already been inferred for other speeies of this genus

(Rosso 1999). The relative proximity of the median zooids to the substratum may also explain the smaller

orifiee dimensions (i.e., smaller tentaeles, if these traits are interrelated) in eomparison with zooids from

the lateral and latero-abfrontal series {cf. Table 5). On the other hand, the small orifiees may simply be

eorrelated with the smaller zooid size of the median series owing to eonstmetional eonstraints of the

eolony.

Undamaged zooeeia with one or two intramural buds, reeognisable by the presenee of multiple orifiee rims

within the primary orifiee {cf. Beming 2008), were present in all of the studied eolonies. Reparative growth

within primary zooids may either be triggered by partial predation (Beming 2008) or, in (seasonally)

oligotrophie eonditions, by partial starvation (J.-G. Harmelin pers. eomm. 2008). The typieal eonstrietions

within eolonies of C. strangulatum (Fig. 5D), from whieh the speeifie epithet derives, may also be a

response to seasonal reduetion in food supply during whieh eolony growth slows down or eomes to a

halt. The paueity of ovieells even in long rods, in eoneert with frequent signs of breakage and reparative

growth at eolony eonstrietions, may refieet another adaptation to unfavourable eonditions. Instead of

investing energy in embryos, propagation may more often proeeed by fragmentation of the eolonies at

these eonstrietions (Fig. 5C, D). However, ovieells are eommonly present in a very elosely related, hitherto

undeseribed speeies from the Great Meteor Bank at similar depths (pers. observation). This seamount is

situated in the eentral North Atlantie in eertainly more oligotrophie eonditions than the populations of C.

strangulatum from the eontinental shelf and slope.

Acknowledgements

Mary Speneer Jones (NHMUK) is partieularly thanked for her hospitality and help with the eolleetion,

literature, and for taking some of the images of H. mucronelliformis (as did Kevin Tilbrook, Museum

of Tropieal Queensland, Townsville). Jean-Eoup d’Hondt and Pierre Eozouet (MNHN), Miehele Bmni

(MOM), as well as Henry MeGhie and Kate Sherbum (MM), kindly provided the type speeimens of, and

information on, their respeetive eolleetions. To Piotr Kuklinski (NHMUK) and Javier Souto (Universidade

de Santiago de Compostela) I am grateful for taking some of the photos of C. strangulatum. Furthermore,

I thank J. Souto, Osear Reverter-Gil (Santiago), Andrey Ostrovsky (Universities of St. Petersburg

and Vienna), Dennis Gordon (NIWA Wellington), Jean-Georges Harmelin (Centre d’Oeeanologie de

Marseille), Steven Traeey (NHMUK), and an anoynmous reviewer for fmitful diseussions and exehange

European Journal of Taxonomy AA\ 1-25 (2013)

of vital information. This research received support from the SYNTHESYS Project t http://www.

svnthesvs.info/) . which is financed by the European Community Research Infrastructure Action under

the FP7 “Capacities” Program, and which allowed me to study some of the type material at the MNHN

(FR-TAF-1902).

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Manuscript received: 9 February 2013

Manuscript accepted: 10 April 2013

Published on: 21 May 2013

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

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

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Museum, Eondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Bmssels, Belgium;

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