Type species.—Eschara radiata Moll, 1803, by original designation.

1988 Puellina (Cribrilaria) scripta; Harmelin and Aristegui, p. 526, figs. 18–19, 24.

1993 Puellina scripta; Harmelin and d'Hondt, fig. 5.

Holotype.—PMC. B27.10.10.2019a. Capo Milazzo Peninsula: Cala Sant'Antonino center, sample 2015: one small fragment including ovicellate zooids and interzooidal avicularia.

Paratypes.—PMC. B27.10.10.2019b. Additional specimens from Capo Milazzo Peninsula: Cala Sant'Antonino West, sample 1 (1999: surface): one specimen; Cala Sant'Antonino center, sample 17 (2000): three specimens; sample 2015: 12 specimens in addition to the holotype. PMC. B27.10.10.2019c. Scoppo: sample 24 top: two specimens.

Diagnosis.—Colonies encrusting, multiserial. Autozooids nearly flat, oval to irregularly polygonal. Basal pore-chambers present. Gymnocyst visible along the zooidal margins. Frontal shield consisting of 14–25 costae with 4–11 intercostal pores/ lacunae. Suboral bar formed by the first pair of widest costae with blunt median prominence and proximal pore. Orifice transversely D-shaped with five (occasionally 6–7) oral spines, four in ovicellate zooids. Interzooidal avicularia with elongate, triangular or parallel-sided, raised rostrum, crossbar lacking. Ovicell hyperstomial, presumably cleithral. Ooecium formed by distal autozooid, with a longitudinal median carina. Kenozooids rare.

Occurrence.—Cribrilaria profunda n. sp. is presently known from the early Pleistocene deep-water deposits of southern Italy (Gelasian of Capo Milazzo Peninsula and early Calabrian of Scoppo, Messina), in the Recent Ibero-Moroccan Gulf (223–990 m depth), the Gibraltar Strait (580 m depth) (Harmelin and Aristegui, 1988), and in the Alboran Sea (205 m) (Harmelin and d'Hondt, 1992, 1993).

Description.—Colonies encrusting, multiserial, unilaminar, the largest observed fragment including a dozen zooids. Zooids large and nearly flat, slightly longer than wide (L/W = 1.15: Scoppo; 1.29: Milazzo), oval to rhomboidal or rarely irregularly polygonal in shape, wider in their proximal half; zooidal boundaries marked by shallow grooves (Figs. 2.1, 2.4, 2.7, 3.1, 3.6). Gymnocyst exposed all along the zooidal margins, usually wider laterally to the orifice and at triple zooid junctions (Figs. 2.1, 3.1, 3.2). Interzooidal communication through basal pore-chambers with windows (∼70 × 20 µm), visible only in some zooids at colony periphery (Fig. 3.5). Frontal shield flat (Figs. 2.1, 2.7–2.9, 3.1, 3.2), consisting of 14–25 wedge-shaped costae (including suboral), narrowing and tapering towards the center of the zooid (maximum basal width 32–65 µm), converging toward a median point or along a median longitudinal, transverse, or trifurcate midline. Costae connected by several intercostal bridges leaving 4–11, regularly spaced, subrectangular lacunae, 8–16 µm long; peripheral pores the largest. Intercostal pores reduced to 4–5 proximally to the first suboral pair of costae (Fig. 2.2). These are shorter and larger than the other pairs, and merge along the zooidal midline leaving a suture with a median pore, and often forming a more or less elevated prominence distally, adjacent to the pore (Figs. 2.2, 3.3, 3.4). Orifice transversely D-shaped, outlined by a raised rim. Orifice bearing five (occasionally up to 7) equally spaced, articulated oral spines (Figs. 2.3, 3.3, 3.4), four persisting in ovicellate zooids (Fig. 2.2). Interzooidal avicularia common, directed laterally or rarely distolaterally, with a variably shaped (often triangular) cystid and an elongate triangular to almost parallel-sided rostrum, raised above or positioned between the costate shield of adjacent autozooids, no crossbar (Figs. 2.1, 2.4, 2.7, 2.9, 3.5, 3.6). Ovicell hyperstomial, presumably cleithral. Ooecium formed by the distal autozooid. Ectooecium smooth, with a longitudinal median elevated carina (Figs. 2.1, 2.2, 2.6, 3.6). A single kenozooid with costate frontal shield numbering 13 costae was observed (Fig. 2.4). Ancestrula not seen.

Etymology.—From the Latin profundus, alluding to its deep-water distribution.

Remarks.—Specimens from Capo Milazzo and Scoppo are very similar in general appearance, including the occurrence of some irregularly polygonal autozooids with a somewhat trifurcate suture in the costate shield. Measurements also largely overlap, although Capo Milazzo material shows more variability. Yet, some specimens from Scoppo show a slightly convex costate shield with fewer costae, a more raised suboral prominence, and more (occasionally 6–7) oral spine bases. Variability in the number of oral spines within the same species is known in other cribrilinids, such as Cribrilaria pseudoradiata Harmelin and Aristegui, 1988. Specimens reported as Cribrilaria scripta (Reuss, 1848) by Harmelin and Aristegui (1988) and Harmelin and d'Hondt (1993) share their characters with the Capo Milazzo material and are here considered conspecific (see below).

Cribrilaria profunda n. sp. is very similar to the Recent C. saginata Winston, 2005 from off Bahia Honda (Cuba) (Winston, 2005) and the Bahama Bank (Rosso et al., 2018). However, C. saginata differs in having a distinctly more extensive proximal gymnocyst, a shorter and squatter orifice (orifice length/orifice width 0.42–0.55 in C. saginata vs. 0.64–0.69 in C. profunda n. sp.), five constant oral spines, and carinated suboral costae. Hincks (1884), and later Neviani (1900), also suggested conspecificity between C. saginata, as C. radiata (Moll, 1803) from Florida, and the middle Miocene (Langhian) Lepralia elegantissima Seguenza, 1880 from southern Calabria (Italy), which is, however, extremely unlikely owing to the great geographic and temporal distance between the two populations. In addition, the only illustration available for L. elegantissima (Seguenza, 1880, pl. 8, fig. 11) is a drawing showing a very distinctive morphology for this species, with ovoidal zooids having a wide and prominent frontal median keel, and seemingly 3–5 suboral tubercles alternating with lacunae.

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.

Type species.—Puellina pedunculata Gautier, 1956, by original designation.

cf. 1956 Puellina pedunculata Gautier, p. 203, fig. 20.

cf. 1966 Colletosia pedunculata; Prenant and Bobin, p. 596, fig. 207 III.

cf. 1970 Cribrilaria pedunculata; Harmelin, p. 93, fig. lg, h, pl. 2, fig. 6.

cf. 1987 Puellina (Glabrilaria) pedunculata; Bishop and Househam, figs. 95–97, tab. 13.

cf. 1988 Puellina (Glabrilaria) pedunculata; Harmelin, p. 31, figs. 9–11.

cf. 2013a Puellina (Glabrilaria) pedunculata; Rosso et al., tab. 17.1.

cf. 2015 Puellina (Glabrilaria) pedunculata; Sanfilippo et al., tab. 2, fig. 5f.

cf. 2019a Glabrilaria pedunculata; Rosso et al., fig. 5e, f.

Holotype.—MNHN-IB-2008-10384, Grand Conclu de Riou (Golfe de Marseille), Mediterranean, Recent.

Occurrence.—Glabrilaria pedunculata is an endemic Mediterranean species, widespread throughout the basin, from the Gulf of Lion to the Aegean Sea. Its presence in the Atlantic is restricted to areas swept by Mediterranean outflow water (Harmelin and d'Hondt, 1992). It has been reported from: (1) shallow-water submarine caves in the Provençal area (Harmelin, 1969, 1970, 1988, 2003), in the Ionian sea (Rosso et al., 2013a, b; Sanfilippo et al., 2015) and Aegean sea (Crete: Harmelin, 1988; Lesvos: Rosso et al., 2019a); (2) cryptic microhabitats from shallow waters (Harmelin, 2003), mid-shelf coralligenous cliffs, and outer shelf “Coralligène de Plateau,” at 100–140 m depth off Lybia and near Santorini (Harmelin, 1988); and (3) at bathyal depths, ∼700 m in the Sicily Strait (Harmelin, 1979, 1988), ∼280 m in the southern Adriatic Sea (D'Onghia et al., 2015), and ∼500 m at Leuca, northeastern Ionian Sea (Mastrototaro et al., 2010), usually associated with cold-water coral habitats. Specimens from the Gelasian of Sicily represent the first fossil record for this species, suggesting its persistence, at least in deep-water settings, in the Mediterranean since the early Pleistocene.

Description.—Colony encrusting, multiserial, unilaminar (Fig. 4.1, 4.6), the largest specimen including at least 50 zooids. Zooids oval, longer than wide (L/W = 1.28), convex, outlined by furrows filled by incipient re-crystallization (Fig. 4). Interzooidal communication through basal pore-chambers, more than 10 visible only in some marginal zooids, with longitudinally elongate windows ∼10 × 20 µm (Fig. 4.4). Gymnocyst narrow, steeply sloping. Costate frontal shield oval and extensive, formed by 13–17 (including suboral) wedge-shaped, prominent costae, 27–45 µm wide at the base, converging towards the midline and forming a slightly raised carina (Fig. 4.4, 4.5). Costae joined by regularly spaced intercostal bridges leaving 6–7 slit-like intercostal pores, ∼7–8 µm long (Fig. 4.5). Only four intercostal spaces occur proximally to the suboral pair of costae, which are flat and merge at the midline forming a triangular shelf, possibly leaving a single round pore (Fig. 4.3, 4.6). Orifice transversely D-shaped (Fig. 4.1, 4.3, 4.6), marked by a raised rim, provided with 6–7 closely spaced, articulated oral spines (Fig. 4.1, 4.3), four persisting in ovicellate zooids (Fig. 4.2, 4.5). Ovicells hyperstomial, presumably cleithral. Ooecium formed by distal kenozooid, with frontally visible small costate shield consisting of three costae (Fig. 4.4); ectooecium smooth, with elevated longitudinal carina (Fig. 4.2, 4.4, 4.5). Avicularia not observed. Abundant small kenozooids recorded in larger colonies, interspersed between autozooids, seemingly polygonal, with boundaries obliterated by recrystallisation, with extensive gymnocyst and costate frontal shield of 6–8 costae (Fig. 4.6). The only ancestrula found seemingly regenerated as a miniature autozooid (Fig. 4.1).

Materials.—Rosso-Collection, collective code: PMC I. Pl. B.81a: Capo Milazzo Peninsula: Cala Sant'Antonino center: sample 2015: three specimens; sample 17 (2000): one specimen; Punta Mazza: sample 4: two specimens; sample 5: one specimen.

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 Harmelin and Aristegui, 1988 and G. orientalis Harmelin, 1988). The long-stalked (=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.

Holotype.—PMC. B28.10.10.2019a: colony consisting of ∼20 autozooids, some ovicellate. Scoppo, sample 24 top, early Calabrian, MNN19b-19c biozones.

Paratype.—PMC. B28.10.10.2019b: small colony fragment including seven autozooids, two ovicellate. Scoppo: sample 24 top, early Calabrian, MNN19b-19c biozones.

Diagnosis.—Colony encrusting, multiserial. Autozooids convex. Gymnocyst narrow. Frontal shield consisting of 12–14 prominent and tuberculate costae with 3–7 intercostal spaces. Suboral pair of costae forming a bifid mucro. Orifice transversely D-shaped with six oral spines, four persisting in ovicellate zooids. Ovicells subimmersed. Ooecium formed by distal kenozooid, surface smooth, with transverse rib. Avicularia not observed. Kenozooids rare.

Occurrence.—Only known from the early Calabrian of Scoppo, Messina.

Description.—Colony encrusting, multiserial, unilaminar, but including superimposed lobes (Fig. 5.1), the largest observed fragment consisting of ∼20 zooids. Zooids oval, longer than wide (L/W = 1.44), convex, the outline hidden by incipient recrystallization (Fig. 5.4). Interzooidal communication through basal pore-chambers visible in some peripheral zooids, with slightly longitudinally elongate windows ∼21 × 18 µm (Fig. 5.2). Gymnocyst very narrow, except for proximal and, occasionally, lateral extensions wedged between neighboring zooids (Fig. 5.2, 5.3). Frontal shield oval and extensive, formed by 14–16 (including suboral) wedge-shaped, prominent, tuberculate costae, 26–47 µm wide at the base, converging towards the midline (Fig. 5.2–5.4). Costae joined by intercostal bridges apparently leaving 6–7 intercostal pores (Fig. 5.2), seemingly reduced to 3–4 proximally to the suboral pair of costae. These are shorter and more robust than the other costae and raised at the midline, forming a bifid mucro (Fig. 5.2, 5.3). Orifice transversely D-shaped, provided with six closely spaced, articulated oral spines (Fig. 5.2), four persisting in ovicellate zooids (Fig. 5.3, 5.4). Ovicells subimmersed. Ooecium formed by the distal kenozooid, with frontally visible costate (4–5 costae) shield and distal band of gymnocyst (Fig. 5.3, 5.4); ooecium with prominent, transverse, straight to slightly arched rib possibly with protruding spikes (lost) (Fig. 5.3, 5.4); an additional thinner and lower longitudinal carina was observed in a single ooecium (Fig. 5.3). Avicularia not observed. Only one kenozooid was observed. It was small, polygonal, with a relatively narrow gymnocyst and costate frontal shield of five radial costae (Fig. 5.3). Ancestrula not observed.

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.

Occasionally, transverse ornamentation has been reported in the ooecia of other cribrilinid genera. A succession of ribs adds to a longitudinal carina in Puellina cassidainsis Harmelin, 1984 from the 3PP submarine cave in the Mediterranean French coast (see Harmelin, 1984, fig. 7b). A cruciform pattern can develop in the ooecia of Cribrilaria macaronensis (Harmelin, 2006), and transverse ridges or wrinkles in Cribrilaria atlantis (Harmelin, 2006), both species previously assigned to Puellina (see Harmelin, 2006, fig. 1).

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.

Type species.—Lepralia figularis Johnston, 1847, by original designation.

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).

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 fan-shaped 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), Puellina saldanhai Harmelin, 2001, and Puellina mikelae Harmelin, 2006. Following Rosso et al. (2018), the latter two species are here allocated to the genus Cribrilaria: Cribrilaria saldanhai (Harmelin, 2001) n. comb. and Cribrilaria mikelae (Harmelin, 2006) n. comb. Both ovicell variants sometimes may appear within the same colony (e.g., in C. punctata and “P.” harmeri) indicating a developmental plasticity of this character (reviewed in Ostrovsky, 2013). A similar plasticity in ovicell formation is only known in some Calloporidae (Ostrovsky and Schäfer, 2003; Ostrovsky et al., 2009; Ostrovsky, 2013) that are presumed ancestors of cribrilinids.

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, and F. pulcherrima Tilbrook, Hayward and Gordon, 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 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).

Suboral costae often differ from the other pairs. In the type species of Figularia, suboral costae merge, forming a smooth, wide shelf facing the orifice, most evident in ovicellate zooids (Fig. 6). Wide suboral costae associated with ovicellate zooids were also observed in F. rhodanica Li, 1990. In F. pelmatifera Gordon, 1984 the suboral pair of costae develops into two lateral, divergent, spinose processes (see Gordon, 1984, pl. 19, fig. E).

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 and F. jucunda Canu and Bassler, 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).

1847 Lepralia figularis Johnston, p. 314.

1966 Figularia figularis; Prenant and Bobin, p. 604, fig. 2010 I–IV, VI.

1998 Figularia figularis; Hayward and Ryland, p. 338, fig. 120, cum syn.

2002 Figularia figularis; Hayward and McKinney, p. 38, fig. 16 D–E.

2006 Figularia figularis; Berning, 2006, p. 49, pl. 3, figs. 7, 10, cum syn.

2014 Figularia figularis; Chimenz Gusso et al., p. 167, fig. 84a–c.

Holotype.—NHMUK 1847.9.16.39, English Channel, Recent.

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).

Materials.—Rosso collection PMC. I. H. B.71.b, Apollo Bank sample: two specimens, Recent; Rosso-Collection PMC I. Pl. B.71.c, Scoppo: sample 24 top: two specimens, early Calabrian, MNN19b-19c biozones.

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.

Holotype.—PMC. B22. 5.4.2015.a: bilaminar fragment including some autozooids and the only observed ovicell. Cala Sant'Antonino, sample Cala Sant'Antonino center, 2015, Gelasian.

Paratypes.—PMC. B22. 5.4.2015.b: additional 39 fragments from the same sample as the holotype, including several fan-shaped colony portions. One fragment from sample 17 (2000), Cala Sant'Antonino center.

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.

Occurrence.—Exclusively known from early Pleistocene (Gelasian) deep-water sediments of Capo Milazzo (NE Sicily, Italy).

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 portion, consisting of four zooids arranged in back-to-back pairs (Fig. 8.1, 8.2, 8.4–8.8). Other fragments of similar size include only the edges of presumably ribbon-like colonies (Figs. 8.3, 10.1). Putative proximal heteromorphs, possibly arising from an encrusting phase and forming the basal stalk, lacking calcified frontal shield. Zooidal boundaries marked by grooves. Zooids large, about twice as long as wide (L/W = 1.72), gently arched distally, wedged proximally. Gymnocyst more extensively exposed proximally and laterally (Figs. 8.2, 8.3, 8.5, 10.1–10.4), locally obliterated by recrystallization (Fig. 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.