Cribrilinid bryozoans from Pleistocene Mediterranean deep-waters, with the description of new species

Abstract Cribrilinid bryozoans originating from Pleistocene deep-water sediments from two localities near Messina (Sicily, Italy)—Capo Milazzo (Gelasian) and Scoppo (Calabrian)—were examined. Five cribrilinid species were found, three in each locality and time interval, with only one species shared. Three species, Cribrilaria profunda n. sp., Glabrilaria transversocarinata n. sp., and Figularia spectabilis n. sp., are new to science. Of the two remaining species, Figularia figularis was already known from local fossil associations, whereas Glabrilaria pedunculata, a present-day Mediterranean species, is recorded for the first time as a fossil. New combinations are suggested for two species previously assigned to Puellina, Cribrilaria saldanhai (Harmelin, 2001) n. comb. and Cribrilaria mikelae (Harmelin, 2006) n. comb. The diagnosis of the genus Figularia was amended to include an erect growth morphology in addition to the encrusting form, and the occurrence of ooecia formed by the distal kenozooid. Following a literature revision of all species currently assigned to Figularia, the new combinations Vitrimurella capitifera (Canu and Bassler, 1929) n. comb. and Hayamiellina quaylei (Powell, 1967a) n. comb. are suggested, and problematic species are listed and briefly discussed. UUID: http://zoobank.org/b7b36152-bf7b-4e00-b6ec-2614b2a58f1b


Introduction
is an extremely large family of cheilostome bryozoans including 127 genera and more than 700 living and fossil species to date, accounting for ∼3% of total bryozoan diversity (Bock, 2020). First appearing ca. 100 Ma, in the Cenomanian, Cribrilinidae underwent a peak of diversification during the Santonian, greatly contributing to the radiation of cheilostomes in the Late Cretaceous (Cheetham, 1971;Jablonski et al., 1997 and references therein). This family is one of the most species-rich in the present-day Mediterranean (Rosso and Di Martino, 2016), as well as in other regions of the world (e.g., Gordon et al., 2019). Cribrilinids exhibit a typical and distinctive costate frontal shield, but also high morphological variability, including different types of heteromorphs (avicularia, kenozooids, articulated and non-articulated spines, etc.) and ovicell structures. A future subdivision of Cribrilinidae into several families or subfamilies is very likely. A more accurate definition of certain genera will, however, require a thorough re-examination of the original material, particularly of the numerous Cretaceous representatives (e.g., Taylor and McKinney, 2006;Rosso et al., 2018), as well as phylogenetic analyses. Genus and species identification are often based on subtle morphological characters, such as those associated with the zooidal orifice and the suboral bar (e.g., Harmelin, 1970Harmelin, , 1978Harmelin, , 2001Harmelin, , 2006Bishop and Househam, 1987), which require scanning electron microscopy (SEM), still lacking in the descriptions of numerous taxa. In fossil material, identification of taxa is also jeopardized by taphonomic filters, with abrasion, corrosion, partial dissolution and recrystallization obliterating fine diagnostic characters. This is particularly true for species introduced in old publications, normally including only brief descriptions and often lacking proper illustrations. Descriptions and revisions of fossil cribrilinids based on detailed illustrations are scarce in the modern literature, especially for specific stratigraphic intervals (Berning, 2006;Taylor and McKinney, 2006;Di Martino and Rosso, 2015). In this context, this paper aims to: (1) document cribrilinid associations from Pleistocene deep-water habitats of southern Italy; (2) illustrate fossil representatives of some established species; (3) describe three new species; (4) amend the diagnosis of the genus Figularia Jullien, 1886, and provide a comparative morphological analysis of species currently assigned to this genus; and (5) propose new *Corresponding author Vertino, 2003). At Scoppo, these sediments unconformably lie on Messinian brecciated evaporitic limestone. They consist of basal rudstones rich in fragments of cold-water corals (i.e., M. oculata, D. pertusum, and D. dianthus) that are overlain by poorly cemented white marls with sparse corals and plates of the cirriped Scillaelepas Seguenza, 1876. These macrofossils, and ostracodes, point to deposition in bathyal environments (Vertino et al., 2013;Sciuto, 2016) in the MNN19b-19c biozones (A. Baldanza, personal communication, 2015), corresponding to the early Calabrian (=Santernian).

Materials and methods
Studied material originates from deep-water sediments cropping out in two different localities near Messina in north-eastern Sicily: Capo Milazzo Peninsula (two outcrops: Cala Sant'Antonino and Punta Mazza) and Scoppo ( Fig. 1; see Geological setting for details). Additional material used for comparison derives from a present-day submarine sample collected at the Apollo Bank off Ustica Island in the Tyrrhenian Sea ( Fig. 1).
At Capo Milazzo, cribrilinid bryozoans were found in "sample 1 (1999)" collected near the top of the layers exposed at Cala Sant'Antonino West; "sample 17 (2000)" and "sample 2015" collected in the central part of Cala Sant'Antonino outcrop; and "sample 4" and "sample 5" collected in biogenic layers near the base of Punta Mazza section, corresponding to "sample 12" and "sample 11" of Sciuto (2014b), respectively. Further information on these samples can be found in Sciuto (2014b) and Rosso and Sciuto (2019). At Scoppo, cribrilinids were found in a test sample associated with a Scillaelepas-rich layer, and in the sample "Scoppo 24 top" coming from uncemented marly sediment.  Rosso et al.-Pleistocene deep-water cribrilinids At the Apollo Bank, coarse sediments associated with the kelp Laminaria rodriguezii Bornet, 1888 were collected at about 60 m depth. Living and dead bryozoan associations were characterized by high species richness, but delivered only one colony (now fragmented) of Figularia figularis (Johnston, 1847) (Di Geronimo et al., 1990).
Sediment was routinely treated (washed, sieved, and dried) at the Paleoecological Laboratory of the University of Catania. All bryozoans were picked from residues larger than 0.5 mm. After preliminary identification under a stereomicroscope, selected uncoated specimens were mounted for scanning electron microscopy (SEM) using a TESCAN VEGA 2 LMU in backscattered-electron/low-vacuum mode at the Microscopical Laboratory of the University of Catania. For the attribution of the specimens to the genera Cribrilaria Canu andBassler, 1929 andGlabrilaria Bishop andHouseham, 1987, we followed the diagnoses in Rosso et al. (2018) summarized herein: Cribrilaria has totally calcified non-pseudoporous ooecia produced by the distal autozooid or kenozooid, interzooidal avicularia of variable size and shape, usually five (4-8) oral spines, and relatively large uncalcified windows of pore-chambers; Glabrilaria has non-pseudoporous ooecia that are exclusively produced by the distal kenozooid, erect or semi-erect avicularia, 6-7 (rarely five) oral spines, small to moderately sized uncalcified windows of pore-chambers. Measurements were obtained from SEM images using the image processing program ImageJ (Schneider et al., 2012). Measurements were tabulated and provided in micrometers. The complete range is given first, followed by the mean value plus/minus standard deviation and the number of measurements taken. In specimens of Glabrilaria, zooidal boundaries were obliterated by recrystallisation with bands of crystals filling the interzooidal grooves. To estimate zooidal size, length was measured from the distal end of the orifice to the mid-point of the crystal band located proximally, while width was measured from mid-point to mid-point of the crystal bands located laterally.
Repositories and institutional abbreviations.-All specimens described and illustrated in this work are part of the Rosso Collection deposited at the Museum of Paleontology of the University of Catania (PMC) under the catalogue numbers reported in the "Systematic paleontology" section. Other abbreviations: MNHN, Muséum national d'Histoire naturelle, Paris; NHMUK, Natural History Museum, London; NMNH, National Museum of Natural History, Smithsonian Institution, Washington DC.
Etymology.-From the Latin profundus, alluding to its deep-water distribution.
Cribrilaria scripta and C. radiata, although similar in appearance to C. profunda n. sp., have smaller zooidal dimensions and larger interzooidal avicularia, and four oral spines occur in most zooids in the latter species (Harmelin, 1970;Bishop and Househam, 1987). Recent specimens of C. scripta sensu Harmelin and Aristegui (1988) from deep waters of the Ibero-Moroccan Bay and Gibraltar Strait, are here attributed to C. profunda n. sp. based on the measurements, the presence of generally five oral spines, and presence of a robust and smooth pair of suboral costae forming a median prominence.
In addition, specimens from the early Messinian of Carboneras (SE Spain) identified by J.-G. Harmelin as Puellina (Cribrilaria) scripta and mentioned in Barrier et al. (1992), without description or illustrations, might belong to C. profunda n. sp.
The Recent Cribrilaria pseudoradiata from the upper bathyal Atlanto-Mediterranean region is also similar to C. profunda n. sp., but has smaller dimensions and lacks interzooidal avicularia.
Cribrilaria profunda n. sp. could possibly correspond to Lepralia planicosta Seguenza, 1880, a cribrimorph species reported from Plio-Pleistocene sediments of the Messina Strait area. Seguenza (1880) distinguished his species from C. scripta, adducing that autozooids were irregularly shaped, with a flat costate shield consisting of several costae, as in C. profunda n. sp. Unfortunately, Lepralia planicosta, supposedly corresponding to Lepralia scripta sensu Manzoni (1875) from the early Pliocene of Castrocaro, was not figured and the type material was lost in 1908 during the Messina earthquake. We refrain from selecting our material as the neotype of L. planicosta because the original description of this species seems insufficient to ensure their conspecificity, and the type localities, although geographically close, are not exactly the same, and neither are the geologic horizons. Seguenza (1880) abstained from illustrating his new species and referred to drawings of L. scripta sensu Manzoni (1875, figs. 25, 25a). Manzoni's specimens, held in the collection of the Museo di Storia Naturale, Geologia e Paleontologia of Florence, should be located and examined before selecting a neotype for this species.
Remarks.-The available specimens are worn and recrystallized, preventing recognition of some diagnostic characters. However, the morphology and morphometrics of autozooids, ooecia, and kenozooids are closely reminiscent of Glabrilaria pedunculata Gautier, 1956, although with a few small differences. The present-day Mediterranean species invariably shows six oral spines and two median pores in the triangular shelf distal to the suboral costae (Bishop and Househam, 1987, fig. 97;Harmelin, 1988, fig. 17a, c;Rosso et al., 2019a, fig. 5e, f). However, both the variability in the number of oral spines and the presence/absence of median pores are considered to be in the range of intraspecific variability in cribrilinids (e.g., C. pseudoradiata Aristegui, 1988 andG. orientalis Harmelin, 1988). The longstalked (=pedunculate) avicularia, originating from basal pore chambers in both autozooids and kenozooids, which are typical of G. pedunculata, were not observed in our fossil specimens. This is likely a taphonomic bias, because such avicularia can be easily detached even in living colonies, as observed in Glabrilaria hirsuta Rosso in Rosso et al., 2018 from the Bahama Bank. In our fossil specimens, zooidal boundaries are mostly covered by neomorphic calcite crystals that prevent the detection of the basal pore chambers from which the pedunculate avicularia are budded. However, in Figure 4.4 and 4.5 (see arrows) the pores potentially producing the avicularia lateral to the ovicell are visible.
Seven oral spines were described in Glabrilaria corbula Bishop and Househam, 1987 and Glabrilaria orientalis lusitanica Harmelin, 1988, two closely related extant species reported from the Atlanto-Mediterranean region and the Gibraltar Strait area, respectively. However, the former species shows an ooecium that is formed by a distal kenozooid which is not distinguishable in frontal view, has 4-6 costae-like ridges arranged in a radial pattern, a flatter autozooidal shield with somewhat carinate costae that are sometimes with a pelma, and two large pores in the suboral shelf (Bishop and Househam, 1987;Harmelin, 1988), while the latter species lacks midline pores in the suboral shelf (Harmelin, 1988). Glabrilaria orientalis lusitanica also has semi-erect interzooidal avicularia (Harmelin, 1988) backed against the ooecium. Six to seven oral spines also occur in Glabrilaria africana (Hayward and Cook, 1983), but this species has numerous variably sized pores in the suboral shelf in addition to semi-erect avicularia associated with the ooecium and squeezed between autozooids.
Etymology.-From the Latin transversus, meaning transversely placed, and carina alluding to the typical median crest of the ooecium.
Remarks.-The co-occurrence of a prominent transverse ridge on the ooecium and a bifid suboral mucro is distinctive of this species. Ooecia with a transverse ridge are known in a few species only. One is the extant Glabrilaria hirsuta Rosso in Rosso et al., 2018 from the Bahama Bank, in which the ridge is, however, very arched to subtriangular and equipped with prominent spine-like processes (Rosso et al., 2018). Furthermore, in G. hirsuta, the number of oral spines (six, four persisting in ovicellate zooids) occasionally increases to seven, the costae have more obvious spine-like processes at the periphery of the frontal shield, the suboral costae form a transverse spiny crest proximal to the orifice, and kenozooids arranged in rows or clusters are very common (Rosso et al., 2018). In the extant Glabrilaria cristata (Harmelin, 1978) from the Hyères and Meteor banks south of the Azores, the ooecial ridge is extremely protruding and situated more proximally towards the orifice, contributing to form a sort of spiny collar around the orifice together with the second pair of suboral costae. These costae bear cockscomb-like spines that are still present but smaller than those on the other pairs (Harmelin, 1978). Oral spines are invariably seven in this species.
Measurements of Glabrilaria transversocarinata n. sp. generally overlap with those of G. cf. G. pedunculata from Capo Milazzo (Table 2), but tend towards the higher values, sometimes exceeding the upper limit. The only exception is the size of the kenozooid, which seems to be smaller, although only based on a single measurement. However, morphological differences, including the number of oral spines, shape of costae, suboral lacuna and ooecia, and the rarity of kenozooids, distinguish the two species.
The two colony fragments available are detached from the substratum, a common feature for bryozoan specimens found in the Capo Milazzo "yellow marl." This may indicate either that the substratum was organic or that selective aragonitic dissolution took place during/before fossilization.
Amended diagnosis.-Colony commonly encrusting, but erect, fan-shaped, or developing erect lobes in some species. Autozooids with variably developed gymnocyst, usually wider proximally; costate shield formed by few to numerous (up to 30) costae, each bearing a pelma (circular to drop-shaped or transversely elongated) varying in size and position. Orifice with well-developed poster and condyles, dimorphic and typically larger in ovicellate zooids. Oral spines absent. Avicularia, when present, vicarious, elongate, and often spatulate, with complete crossbar. Ovicells hyperstomial or subimmersed, cleithral. Ooecium formed by the distal autozooid or kenozooid (sometimes in the same colony), bilobate, consisting of two very large, modified costae, arched and meeting in the midline to form a suture and/or carina; each costa with a wide fenestra. Interzooidal communication via mural pore chambers in the transverse walls and multiporous septula in the lateral walls. Ancestrula only observed in the type species, wider than autozooids, subcircular, with narrow gymnocyst encircling an extensive opesia with differentiated orifice; no spines.
Remarks.-The finding of a new species having morphological skeletal features fitting into the genus Figularia Jullien, 1886, but characterized by erect colony form and a very distinctive and large ooecium formed by a distal kenozooid, led to the examination of species currently placed in this genus (Tables 3, 4).  Figularia was introduced by Jullien (1886, p. 608) who designated Lepralia figularis Johnston, 1847, an Atlanto-Mediterranean extant species, as the type species of the genus, and included an additional fossil species Lepralia elegantissima based on the unique drawing available (Seguenza, 1880, p. 83, pl. 8, fig. 11). This latter species, depicted with oral spine bases, is more likely to be a species of Cribrilaria (see also Remarks on Cribrilaria profunda n. sp.). Oral spines are absent in the type species F. figularis (see Soule et al., 1995, fig. 45C), as well as in all living and fossil specimens found to date (e.g., Figs. 6, 7). The absence of oral spines has also been reported almost consistently in the diagnosis of the genus, with only a few exceptions (e.g., Gordon, 1984). Further diagnostic characters include a complete crossbar in the vicarious avicularia, and the presence of large, symmetrical ectooecial fenestrae and a median carina in the ooecium (see Soule et al., 1995;Hayward and Ryland, 1998;Kukliński and Barnes, 2009;Yang et al., 2018).

Rosso et al.-Pleistocene deep-water cribrilinids
The erect colony-form has never been mentioned in the generic diagnosis before. However, Busk (1884, p. 132) described Figularia philomela as "free; erect or decumbent (hemescharan)." Subsequently, Hayward and Cook (1979, p. 76) found a bilaminar fragment of F. philomela interpreted as part of an erect foliaceous colony possibly arising from an encrusting phase (var. adnata of Busk, 1884). Gordon (1989, p. 15, 16) recorded the occasional occurrence of an erect bilamellar lobe, arising from the adjacent encrusting zooids, in a colony of Figularia mernae Uttley and Bullivant, 1972 from Puysegur Bank, off the South Island of New Zealand. The fanshaped colonies of the newly discovered Figularia species from Capo Milazzo, although often fragmentary (Fig. 8), show a configuration comparable to that observed in F. mernae, with basal zooids elongated and arranged in back-to-back adjacent pairs ( Fig. 8.1, 8.2, 8.6). The lack of a costate frontal shield, with no obvious evidence of breakage, in several proximal/basal zooids, suggests that simplified polymorphs, reminiscent of those in Corbulipora MacGillivray, 1895 (see Bock and Cook, 2001) may occur. However, the raising of the erect fan-shaped portions from an encrusting phase is doubtful until encrusting colonies, or at least isolated encrusting zooids, are found.
The ooecium in Figularia is generally described as bivalved/bifenestrate (Ostrovsky, 2013). In F. figularis, the prominent bilobate ooecium is formed by the distal autozooid, with two costae meeting in the midline leaving a suture and/or forming a slightly raised carina; each costa bearing a large, irregularly shaped and transversely elongate fenestra (membranous area in non-cleaned specimens). The colony fragment of F. figularis from the Apollo Bank (Tyrrhenian Sea, Mediterranean) shows that ooecia formed by the distal kenozooid can co-occur in the same colony in this species (Fig. 6). Though uncommonly reported, and here recorded in F. figularis for the first time, the co-occurrence of ooecia produced by the distal autozooid and kenozooid is known in other cribrilinids, such as Cribrilina punctata (Hassall, 1841), "Puellina" harmeri Ristedt, 1985 (see also discussion in Rosso et al., 2018), Cribrilaria innominata (Couch, 1844) (see Chimenz Gusso et al., 2014),
The kenozooid producing the ooecium in F. figularis shows a crescent-shaped shield of short radial costae, each with a single pelma as in the autozooids, but also with a single intercostal pore (Fig. 6). The same structure is also evident in the fossil species from Capo Milazzo (Fig. 9). Ovicells with ooecia formed by the distal kenozooid also occur in other species currently assigned to this genus, based on examination of available SEM images and, to a lesser extent, drawings (see Table 3).
Ostrovsky (2013, fig. 1.28A) illustrated sectioned decalcified ovicells of F. figularis in which most of the brood cavity is situated in the proximal part of the distal zooid predominantly below the colony surface, thus corresponding to endozooidal type. Whether this position of the brood cavity was an effect of decalcification of the skeleton (and, thus, sagging of the originally raised ooecium) during preparation for sectioning is currently not clear, but this contradicts most descriptions showing hyperstomial ovicells in this species (see references above). Still, a degree of the brood cavity immersion may vary, and, for example, both hyperstomial and subimmersed ovicells are known within the genus Figularia, and hyperstomial, subimmersed, and endozooidal ovicells are described in the different species of Puellina (Ostrovsky, 2013). Subimmersed ovicells were present in Recent colonies of F. figularis from the Mediterranean (A. Ostrovsky, personal observations). Ostrovsky and Taylor (2005) noted the occurrence of species of Figularia-F. clithridiata (Waters, 1887), F. tahitiensis Waters, 1923, andF. pulcherrima Tilbrook, Hayward andGordon, 2001-having costate ooecia (see also Ostrovsky, 2002). Winston et al. (2014) remarked that the occurrence of costate ooecia in F. pulcherrima possibly suggests a better allocation of this species in a distinct genus. Inclusion of costae in the Journal of Paleontology 95(2):268-290 construction of the ooecium has also been observed in Figularia hilli (Osburn, 1950), with two small costae similar to those of the frontal shield added proximally to the larger ooecial halves (see Table 3). Yang et al. (2018), while including pseudoporous ooecia in the diagnosis of Figularia, also suggested the examination of species with multiple ectooecial pseudopores in order to determine if they are genuinely congeneric. These species are here re-assigned to different genera (see also below and Table 4).
A certain variability occurs in the presence/absence of pelmata in the frontal shield, and in their position along the costal length. Sometimes this variability was noted (e.g., Gordon, 1984). Nevertheless, all Figularia species lacking pelmata (i.e., not included in formal descriptions and/or undetectable in available images) are fossil, except "F. philomela var. adnata" (Busk, 1884), suggesting that their absence may be a preservation artefact.
The ancestrula is generally not mentioned in species descriptions to our knowledge. In the amended diagnosis, we include characters of the ancestrula for the first time, based on the ancestrula found in a colony of F. figularis from the Mediterranean illustrated in Rosso et al. (2019b, fig . 5C). The large size of both autozooids and ancestrula (0.65 × 0.67 mm) and the absence of spines are rare and remarkable among cribrilinids, which usually have small, tatiform ancestrulae, and this may have implications on the systematics/phylogeny of this genus within the family Cribrilinidae. However, observation of ancestrulae in additional species is needed to confirm whether this morphology is constant among congeners, which has been proven not to be the case in other cheilostome genera, such as e.g., Escharina Milne Edwards, 1836 (see Berning et al., 2008).
Several species previously assigned to Figularia were recently displaced in different genera of the families Cribrilinidae and Calloporidae (e.g., Vitrimurella, Reginella Jullien, 1886, Inferusia Kukliński and Barnes, 2009, Valdemunitella Canu, 1900; see Bock and Gordon, 2020), and Jullienula Bassler, 1953(Yang et al., 2018. Here, we suggest further displacements: both Figularia? ampla Canu and Bassler, 1928, only tentatively included in Figularia when first described, and Emballotheca? capitifera, Canu and Bassler, 1929, subsequently referred to his new genus Calyptotheca by Harmer (1957) and to Figularia by Di Martino and Taylor (2018), fit better in Vitrimurella, owing to the pseudoporous zooidal gymnocyst and ooecia, and the extremely reduced costate shield. Figularia ryukyuensis Kataoka, 1961 andF. jucunda Canu andBassler, 1929 also need to be revised, pending examination of the type material. These species have pseudoporous ooecia formed by the distal kenozooid without a visible frontal part. Figularia duvergieri Bassler, 1936 has an unusual denticulate proximal orifice margin, and lacks costal pelmata and fenestrae in the ooecium. A detailed revision based on SEM images is needed to confirm generic allocation for these problematic species (Table 4). This issue has been partially addressed by López Gappa et al. (in press).
Occurrence.-Figularia figularis is widely distributed in the Atlanto-Mediterranean area since the middle Miocene (Moissette et al., 1993;Berning, 2006). This species has been commonly reported from shelf habitats, mostly from the deep shelf, often associated with deep coralligenous facies (Di Geronimo et al., 1990;Ballesteros, 2006), and at the shelf break in both the Mediterranean (110-145 m; see Harmelin and d'Hondt, 1992) and the eastern Atlantic as far north as the British Isles (Hayward and Ryland, 1998 Remarks.-Two fossil fragments were found, each consisting of a few zooids (Fig. 7). Zooidal morphological characters allow a reliable identification, even in the absence of ovicells and avicularia. Morphometrics fall within the ranges reported for this species. Inferred teratology in an autozooid resulted in a double-bifurcated frontal shield ( Fig. 7.1, 7.2). This unusual feature also occurs in the type specimen of F. tenuicosta MacGillivray, 1895 from the middle Miocene of Victoria, Australia (Bock, 2020). Although F. figularis exhibits a certain range of morphological variability, some historical records, mostly beyond its confirmed geographical range, proved to be different species (e.g., Brown, 1952). The conspecificity of the colony found on a rock at Armaçao de Pêra in Portugal (Souto et al., 2014) needs to be verified. This colony has an unusual triangular ooecial fenestra with narrow horizontal part and could represent a different species. Figures 8-11, Table 4 Holotype. Diagnosis.-Colony erect, bilaminar with fan-shaped fronds, the tapering proximal terminations possibly consisting of heteromorphs, likely rising from an encrusting phase. Zooidal frontal shield consisting of flat costae, each with a large, elongate drop-shaped pelma placed on its peripheral half; gymnocyst wider laterally and proximally, narrower distally, with faint striations. Vicarious avicularia elongate, spatulate, with extensive rostral palate and complete crossbar. Ovicell subimmersed, presumably cleithral. Ooecium formed by the distal kenozooid with frontally visible costate part, and consisting of two very large, wing-shaped costae merging in the midline producing a longitudinal suture, with two large fenestrae exposing wide areas of endooecium; the costae of the ooecium-producing kenozooid smaller, forming a distal, crescent-shaped crown, each costa with a small pelma.
Description.-Available colony fragments bilaminar, fan-shaped (the largest ∼2 mm long by 3 mm wide); fragments diverging distally at variable angles from a subcylindrical proximal  . 10.5). Costate shield extensive (∼75% of the frontal surface), gently convex, formed by 7-14 flat and smooth costae (maximum basal width 72-111 μm), varying from short and subtriangular proximally to long and parallel sided distally; the suboral pair often the largest (Fig. 10.1-10.5). Costae defined by grooves, connected by an uncertain number of intercostal bridges, presumably 3-4 ( Fig. 10.5), with small oval to subcircular intercostal pores in between. A longitudinal suture marking the costal fusion along zooidal midline (Fig. 11.1). Each costa bearing a single, elongate, drop-shaped pelma with the rounded base placed in correspondence with the base of the costa, while the acute vertex extends up to half to two thirds of costal length. Orifice oval to round, slightly longer than wide, concave proximally, gently arched distally, outlined by a rim of calcification (Fig. 10). Oral spines absent. Avicularia vicarious, infrequent, elongate and slightly asymmetrical, varying in size; rostrum long, spatulate, directed distally and slightly inclined, facing frontally (Figs. 9.1, 11); post-mandibular area short, palate wide, crossbar complete (Fig. 11.3). Ovicell subimmersed, presumably cleithral. A single observed ooecium formed by the distal kenozooid with frontally visible costate shield of 10 costae, longer than wide, wider and slightly more prominent than the ovicellate zooid (Fig. 9). Very large ooecium consisting of two flat, wing-shaped costae converging along the midline, the fusion marked by a longitudinal suture, distally with two small tubercle-like prominences. Large rhomboidal fenestra exposing finely granular endooecium. Orifice of the ovicellate zooid slightly larger than those of autozooids, rounded rectangular. Closure plates or calcified opercula sometimes occluding orifices ( Fig. 10.7).
Etymology.-From the Latin spectabilis, meaning remarkable, exceptional, alluding to the distinctive architecture of the colony and ooecium.
Remarks.-The morphology of the colony, zooids and ooecium distinguish Figularia spectabilis n. sp. from congeners. The flabellate to short, ribbon-like morphology of the colony, with putative heteromorphs placed basally, may suggest the occurrence of basal rhizoids for fixation to the substratum. Alternatively, the connection to an encrusting portion may develop through "sites of articulation" as in Bryobaculum carinatum Rosso, 2002a, occurring in the same sediment.

Discussion
Five species of cribrilinid bryozoans, three of which are new to science, namely Cribrilaria profunda n. sp., Glabrilaria cf. G. pedunculata, G. transversocarinata n. sp., Figularia figularis, and F. spectabilis n. sp., were found in Pleistocene deepwater sediments from north-eastern Sicily. Figularia figularis was already recorded from the area by Seguenza (1880) and Neviani (1900), while C. profunda n. sp. was possibly recorded as Lepralia planicosta (see Remarks above), while the remaining three species, including G. cf. G. pedunculata, represent new records.
Focusing only on deep-water assemblages, cribrilinids are present with three species in both the Gelasian associations from Capo Milazzo and the Calabrian (MNN19b-19c biozones) of Scoppo. These figures are comparable to those found in present-day deep-water associations from the Mediterranean and Atlantic (Bahama Bank), in which cribrilinids usually occur with 2-3 species (Rosso et al., 2018). However, the Gelasian of Capo Milazzo includes at least 46 cheilostome species, and the cribrilinid relative percentage is ∼6%, which is lower than the 10-18% found in present-day assemblages (Rosso and Sciuto, 2019). No comparison can be made for Journal of Paleontology 95(2):268-290 the Calabrian of Scoppo whose bryozoans are still under investigation.
Discovery of a new species of Figularia, F. spectabilis n. sp., led to the emendation of the genus diagnosis and the re-examination of the 32 species and one variety currently assigned to the genus, based on drawings and photographic material available from the literature. This preliminary survey allows us to confidently reassign two species based on published scanning electron micrographs of the type material. The newly proposed combinations are Vitrimurella capitifera (Canu and Bassler, 1929) n. comb. and Hayamiellina quaylei (Powell, 1967a) n. comb., as also suggested by Kukliński et al. (2015). Thirteen species remain doubtful and their assignment to more suitable genera requires examination of the type material (Table 4).
At present, 18 species, including Figularia spectabilis n. sp., match the diagnosis of the genus. This figure will likely change further after a more detailed revision of some fossil species and species left in open nomenclature (see Berning, 2006 for F. haueri and F. figularis; Di Martino et al., 2017 andCook et al., 2018 for two different Figularia spp.) as well as cryptic species/species complexes (e.g., F. clithridiata and F. fissa). Based on our literature review, the diversity of Figularia is reduced by about one-half, from 33 (including F. spectabilis n. sp.) to 18 species, with a revision in the stratigraphic range, but only little variation in the geographic distribution of the genus. The genus possibly appeared in the Cenozoic of Europe and Australia, and commonly occurred in sediments in the European-Mediterranean area during the Miocene. Of the 12 species of Figularia living today, 10 species are found in the Pacific and Australasian region. Only two species, F. figularis and F. dimorpha, fall outside this area, being recorded in the Atlanto-Mediterranean and southwestern Atlantic regions, respectively.
A twofold future investigation is sought. This includes an examination of the type material of all the species in the genus to confirm their status, prioritizing those that appear to remain problematic (see Table 4; issue partially addressed by López Gappa et al., in press), and an accurate re-examination of all species records to refine both the temporal and spatial distribution of the genus and reconstruct its diversification history, as well as disentangle species complexes.