Type species

Cribrilina latimarginata Busk, Reference Busk1884, by monotypy.

Diagnosis

Colony encrusting, multiserial. Autozooids separated by kenozooids. Frontal shield formed by costae, with intercostal pores but without lumen pores. Primary orifice D-shaped; condyle-like processes may be present. Oral spines absent. Secondary orifice may be present. Interzooidal kenozooids with a smooth gymnocyst surrounding a variable developed cryptocyst. Interzooidal vibracular heterozooids present, with elevated cystid and conspicous pivots; vibracular setae long. Adventitious heterozooids absent. Ovicelled zooids dimorphic; ovicell immersed to subimmersed, cleithral, with smooth, imperforate ooecium, sometimes covered by interzooidal kenozooids. Ancestrula similar to autozooids.

Remarks

Jolietina was erected by Jullien, Reference Jullien1886 for Cribrilina latimarginata Busk, Reference Busk1884, from deep water off Montevideo, Uruguay. This little-known genus seems to be restricted to South America, from the Magellanic region (Chile) to Bahia (northeast Brazil); it is characterized by having vibracular heterozooids and elongate interzooidal kenozooids that lie along interzooidal boundaries.

Jolietina latimarginata (Busk, Reference Busk1884)
Figure 2.1, 2.2; Table 1

Reference Busk1884

Cribrilina latimarginata Busk, p. 131, pl. 22, fig. 10.

Reference Waters1888

Cribrilina latimarginata; Waters, p. 22, pl .1, figs. 11, 12.

Reference Moyano G.1984

Jolietina latimarginata; Moyano, p. 56, figs. 16, 17, 33, 34.

Reference López Gappa and Lichtschein1990

Jolietina latimarginata; López Gappa and Lichtschein, p. 20, pl. 9, figs. 3, 4, pl. 15.

Reference Cáceres and Moyano1994

Jolietina latimarginata; Cáceres and Moyano, p. 131, pl. 2, figs. E, F.

Reference Gordon, Voigt, Gordon, Smith and Grant-Mackie1996

Jolietina latimarginata; Gordon and Voigt, p. 91, fig. 2A–D.

Figure 2.

(1, 2) Jolietina latimarginata, MACN-In 43341: (1) general aspect of a colony fragment after treatment with NaOCl; (2) general aspect of an uncleaned specimen, showing its cuticular structures in situ. (3–6) Jolietina pulchra, holotype USNM 8549: (3) general aspect of a colony; (4) autozooids and vibracularia; (5) autozooid and ovicelled zooid; (6) vibracular chamber. (1–5) Scale bars = 200 μm; (6) scale bar = 100 μm.

Table 1.

Measurements (in μm) of Jolietina latimarginata (Busk, Reference Busk1884) (MACN-In 43341), Jolietina pulchra Canu and Bassler, Reference Canu and Bassler1928a (USNM 8549), and Jolietina victoria n. sp. (MLP 36402). N = number of measurements; SD = standard deviation.

Holotype

NHMUK 1887.12.9.524, Challenger station 320.

Occurrence

Jolietina latimarginata has a typical Magellanic distribution, ranging from the cold-temperate waters of southern Chile to the continental slope off Argentina and Uruguay (Busk, Reference Busk1884; Waters, Reference Waters1888; Moyano, Reference Moyano G.1984; López Gappa and Lichtschein, Reference López Gappa and Lichtschein1990; Cáceres and Moyano, Reference Cáceres and Moyano1994; López Gappa, Reference López Gappa2000).

Description

Colony encrusting, unilaminar, multiserial. Autozooids sub-ovate, longer than wide, arranged more or less in transverse rows (Fig. 2.1), separated by kenozooids. Primary orifice transversely D-shaped, distinctly wider than long, proximo-lateral corners with two short lateral condyle-like processes; oral spines absent. Secondary orifice developed as a distal raised rim, slightly cucullate, becoming more so with increasing calcification, obscuring the primary orifice. Spinocyst flat, formed by five or six pairs of costae merging in the middle of the frontal shield, the first pair wider and more robust with a distinct median suture; pairs of costae mostly separated by rows of three pores, with about 30–35 rounded intercostal pores overall; costal bases separated by thin slits (Fig. 2.1, 2.2). Kenozooids elongate, occupying interzooidal margins, their opesiae typically narrow in the middle, wider and triangular at each end, with a smooth gymnocyst surrounding a granular cryptocyst. Interzooidal vibracular heterozooids directed laterally, generally located proximal to the autozooids and touching the kenozooidal gymnocyst, with a large, robust, and elevated cystid with gymnocystal calcification, about half autozooidal length; combined opening reniform, with asymmetrical pivots, one stout, one tiny (Fig. 2.1); vibracular setae long and curved (Fig. 2.2). Ovicells and ancestrula not seen in the present material.

Remarks

Jolietina latimarginata is characterized by having a primary orifice distinctly wider than long, with the proximo-lateral corners having condyle-like processes. The vibracular heterozooids have an elevated cystid with asymmetrical pivots.

The ovicell has been figured by Waters (Reference Waters1888) and illustrated with scanning electron microscopy by López Gappa and Lichtschein (Reference López Gappa and Lichtschein1990). It is semi-immersed, longer than wide, mitriform, imperforate, with a central triangular uncalcified area of ectooecium. The ancestrula and early astogeny of J. latimarginata were described by Cáceres and Moyano (Reference Cáceres and Moyano1994).

This species is frequently found on the stems of the erect cyclostome Hornera americana (d'Orbigny, Reference d'Orbigny1847). Gordon and Voigt (Reference Gordon, Voigt, Gordon, Smith and Grant-Mackie1996) provided a figure of the holotype specimen from H.M.S. Challenger Expedition (NHMUK 1887.12.9.524) found on dead coral at station 320 and of a syntype specimen (NHMUK 1887.12.9.525; erroneously stated to be from the Holocene of Argentina) from the same station, found on the bryozoan Foveolaria elliptica Busk, Reference Busk1884 (Gordon, personal observation, 2020).

Holotype

USNM 8549, Jolietina pulchra, F. Canu and R. Bassler det., Rio de Janeiro, Brazil, 21°48′S, 40°03′W, 128 m, collected 1877 by Steamer Norseman.

Occurrence

Recent, Brazil (Bahia, Rio de Janeiro), 49–128 m (Canu and Bassler, Reference Canu and Bassler1928a; present study).

Description

Colony encrusting, unilaminar, multiserial. Autozooids sub-ovate, longer than wide, arranged more or less in transverse rows, separated by kenozooids (Fig. 2.3). Primary orifice transversely D-shaped to oval, wider than long, proximal margin formed by the first pair of smooth and wider costae, these separated by a thin median suture; oral spines absent (Fig. 2.4, 2.5). Secondary orifice developed as a short collar of calcification around the primary orifice, not obscuring it. Spinocyst arched, formed by six to eight pairs of costae merging in the middle of the frontal shield and separated by rows of about five to eight pores, with about 60–85 rounded intercostal pores overall; costal bases separated by thin slits (Fig. 2.5). Elongate kenozooids lying along interzooidal margins, their opesiae constricted in the middle, drop-shaped to triangular at each end, with smooth gymnocyst surrounding a smooth narrow cryptocyst (Fig. 2.4, 2.5). Vibracular heterozooids generally located proximal to the autozooids and touching the kenozooidal gymnocyst, laterally directed, less than half autozooidal length, the cystid smooth, convex; opesia reniform, with a single median pivot (Fig. 2.4, 2.6). Ovicelled zooid larger than non-ovicelled autozooid; orifice dimorphic, transversely oval, secondary orifice formed by a well-developed collar of calcification obscuring the primary orifice distally, distinctly wider than long (Fig. 2.3, 2.5). Ovicell largely immersed, ooecium smooth, imperforate, resembling a small cap (Fig. 2.5). Ancestrula not seen.

Remarks

This is the first redescription of the species since its original description based on a specimen from Rio de Janeiro, southeast Brazil (Canu and Bassler, Reference Canu and Bassler1928a). Here we examined the holotype but also found other fragments of J. pulchra (USNM 348) studied by Canu and Bassler (Reference Canu and Bassler1928a) that were not included in their paper. This revealed that J. pulchra also occurs in relatively shallower waters (49 m) in Bahia State, northeast Brazil.

Canu and Bassler (Reference Canu and Bassler1928a) stated that J. pulchra was smaller than J. latimarginata, but we found the opposite. Jolietina pulchra is distinguished from J. latimarginata by having larger zooids (385–485 × 324–431 μm in J. pulchra, 324–341 × 230–286 μm in J. latimarginata), more spinocystal pores (60–85 in J. pulchra, 30–35 in J. latimarginata), narrower vibracular chambers (69–78 μm wide in J. pulchra, 129–166 μm wide in J. latimarginata), and shorter vibracular pivots (larger in J. latimarginata). Canu and Bassler's (Reference Canu and Bassler1928a) original material was found on coral fragments.

Type material

Holotype: MLP 36402, one fragment. Paratypes: MLP 36403, five fragments.

Figure 3.

Jolietina victoria n. sp. (1) Holotype, MLP 36402, general aspect; (2) paratype, MLP 36403, detail of three autozooids, vibracularia, and kenozooids; (3) holotype, MLP 36402, autozooids, vibracularia, and kenozooids; (4) paratype, MLP 36403, ovicell. (1–3) Scale bars = 200 μm; (4) scale bar = 100 μm.

Diagnosis

Jolietina with spinocyst formed by eight or nine pairs of costae merging in the middle, separated by rows of about four to six pores, with about 64–96 intercostal pores overall. Vibracular heterozooids with reniform opesia and a robust median pivot.

Occurrence

So far, the species has been found only at the locality “Cabeza de León” (Monte León National Park, Santa Cruz Province, Argentina), in the Punta Entrada Member of the Monte León Formation (early Miocene).

Description

Colony encrusting, unilaminar, multiserial. Autozooids sub-ovate, longer than wide, arranged more or less in transverse or oblique rows, separated by kenozooids (Fig. 3.1–3.3). Primary orifice transversely D-shaped, wider than long, proximal margin formed by the first pair of wider costae, these separated by a thin median suture, proximo-lateral corners of the orifice curved; oral spines absent (Fig. 3.2, 3.3). Secondary orifice developed as a short collar of calcification around the primary orifice, occasionally forming a slightly elevated distal rim but not obscuring the primary orifice. Spinocyst flat, formed by eight or nine pairs of costae merging in the middle of the frontal shield and separated by rows of about four to seven pores, with about 64–96 rounded intercostal pores; costal bases separated by slits (Fig. 3.2, 3.3). Narrow, elongate kenozooids lying along interzooidal margins, their opesiae constricted in the middle, mostly drop-shaped at each end, with smooth gymnocyst surrounding a smooth narrow cryptocyst (Fig. 3.2, 3.3). Vibracular heterozooids generally located proximal to the autozooids and touching the kenozooidal gymnocyst (Fig. 3.2), the cystid smooth, inflated, laterally directed, about one-third of autozooidal length; opesia reniform, with a robust median pivot. Ovicelled zooid with a secondary orifice much wider and shorter than non-ovicelled zooids, formed by a well-developed collar of calcification obscuring the distal part of the primary orifice (Fig. 3.4). Ovicell semi-immersed, with smooth ooecium, imperforate, surmounting the frontal shield of the distal zooid, bordered disto-laterally and partially covered by the interzooidal kenozooids (Fig. 3.4). Ancestrula not seen.

Etymology

Named for the Nao Victoria, the ship that completed the first circumnavigation around the planet, belonging to the Fernão de Magalhães fleet, Portuguese captain of the first European expedition that explored the study area in the sixteenth century. Used as a noun in apposition.

Remarks

Jolietina victoria n. sp. is similar to J. pulchra in having a more-perforated frontal shield (64–96 and 60–85 intercostal pores in J. victoria and J. pulchra, respectively) when compared with J. latimarginata (with 30–35 rounded intercostal pores overall), a similar orifice with rounded proximo-lateral corners but lacking condyle-like processes in the orifice (D-shaped and having proximo-lateral condyles in J. latimarginata). Jolietina victoria differs from both J. latimarginata and J. pulchra mainly in having a spinocyst formed by more costae (five or six pairs in J. latimarginata, six to eight pairs in J. pulchra, but eight or nine pairs in J. victoria).

Jolietina victoria has semi-immersed ovicells partially covered by gymnocystal calcification, which are distinct from immersed ovicells without gymnocyst in J. pulchra.

Type species

Lepralia figularis Johnston, Reference Johnston1847. By original designation. Recent.

Occurrence

Eocene to Recent.

Remarks

Figularia was erected by Jullien (Reference Jullien1886) for Lepralia figularis Johnston, Reference Johnston1847 from the British Isles, while Rosso et al. (Reference Rosso, di Martino and Ostrovsky2020) very recently provided a redescription of the type species as well as a revised diagnosis of the genus. Gordon (Reference Gordon1984) mentioned the range of morphological variation attributed to the genus, including the presence or absence of lumen pores, variable length of costal field, and the characters of the ectooecium. Figularia and Jullienula Bassler, Reference Bassler and Moore1953 were inexplicably downgraded by Soule et al. (Reference Soule, Soule and Chaney1995) to subgenus rank within Reginella Jullien, Reference Jullien1886. Yang et al. (Reference Yang, Seo, Min, Grischenko and Gordon2018) defended the status of Figularia, Jullienula, and Reginella as distinct genera. According to them, Figularia should include taxa with a well-developed lateral gymnocyst and an ooecium that is typically bifenestrate or with ectooecial pseudopores but suggested that the latter character needs reexamination; species with pseudoporous ooecia might not be congeneric. As redefined by Rosso et al. (Reference Rosso, di Martino and Ostrovsky2020), species of Figularia have only a single lumen pore (pelma) in each costa, the ooecium is bifenestrate (or tetrafenestrate if each fenestra divides in two while forming), and there are vicarious avicularia. Accordingly, Figularia currently comprises 18 species: Figularia arnouldi Buge, Reference Buge1956; Figularia carinata (Waters, Reference Waters1923); Figularia clithridiata (Waters, Reference Waters1887); Figularia dimorpha Figuerola, Gordon, and Cristobo, Reference Figuerola, Gordon and Cristobo2018; Figularia figularis (Johnston, Reference Johnston1847); Figularia fissa (Hincks, Reference Hincks1880); Figularia fissurata Canu and Bassler, Reference Canu and Bassler1929; Figularia haueri (Reuss, Reference Reuss1848); Figularia hilli (Osburn, Reference Osburn1950); Figularia japonica Silén, Reference Silén1941; Figularia mernae Uttley and Bullivant, Reference Uttley and Bullivant1972; Figularia pelmatifera Gordon, Reference Gordon1984; Figularia philomela (Busk, Reference Busk1884); Figularia rhodanica Li, Reference Li1990; Figularia speciosa (Hincks, Reference Hincks1881); Figularia spectabilis Rosso, Di Martino, and Ostrovsky, Reference Rosso, di Martino and Ostrovsky2020; Figularia tenuicosta (MacGillivray, Reference MacGillivray1895); and Figularia triangula Powell, Reference Powell1967. Rosso et al. (Reference Rosso, di Martino and Ostrovsky2020), however, reported smooth ooecia in Figularia planicostulata Canu and Lecointre, Reference Canu and Lecointre1928 (Miocene, France), but this species is characterized by having bifenestrate ooecia (MNHN.F.R53650); thus, it is considered to belong to the genus Figularia.

Di Martino and Taylor (Reference Di Martino and Taylor2018) assigned Emballotheca capitifera Canu and Bassler, Reference Canu and Bassler1929 from the Philippines to Figularia, but this species is morphologically distinguished from other Figularia by having a reduced costal shield surrounded by a pseudoporous gymnocyst (non-pseudoporous in Figularia) and ovicells with multiple ectooecial pseudopores (bifenestrate in Figularia). Two other Indo-Pacific species with pseudopores in the gymnocyst and ectooecium were also previously included in Figularia: Figularia lepida Hayward, Reference Hayward1988 from Mauritius and Figularia gemina Tilbrook, Reference Tilbrook2006 from the Solomon Islands. These now belong to Vitrimurella Winston, Vieira, and Woollacott, Reference Winston, Vieira and Woollacott2014, in its own family Vitrimurellidae Winston, Vieira, and Woollacott, Reference Winston, Vieira and Woollacott2014. Vitrimurella was erected for Gemellipora lata Smitt, Reference Smitt1873 and includes species with a dimorphic orifice, small costal field, pseudoporous gymnocyst, and pseudoporous ectooecium that were previously assigned to both Figularia and Tremoschizodina Duvergier, Reference Duvergier1921. Initially, the genus included five tropical species: Vitrimurella anatina (Canu and Bassler, Reference Canu and Bassler1928b); Vitrimurella fulgens (Marcus, Reference Marcus1955); Vitrimurella gemina Tilbrook, Reference Tilbrook2006; Vitrimurella lata (Smitt, Reference Smitt1873); and Vitrimurella lepida (Hayward, Reference Hayward1988). The type specimen of E. capitifera illustrated by Di Martino and Taylor (Reference Di Martino and Taylor2018, figs 106–109), however, allowed Rosso et al. (Reference Rosso, di Martino and Ostrovsky2020) to also include it in this genus. The Pacific species (i.e., V. capitifera, V. gemina, and V. lepida) are distinguished from the Atlantic species by having hyperstomial ovicells (immersed in Atlantic species) and ooecia with a proximal fissure (absent in Atlantic species). Winston (Reference Winston2016) included Vitrimurellidae in the superfamily Hippothooidea because of the pseudoporous gymnocystal frontal shield, vestigial costae, and pseudoporous ooecia, similar to those in some Trypostegidae. This family may have a pliophloean ancestor (Gordon, Reference Gordon, Herrera-Cubilla and Jackson2000), but the lack of a strictly pliophloean orifice and the absence of an avicularian crossbar in trypostegids suggest that Vitrimurellidae is unrelated to Hippothooidea. However, the morphology of the vicarious avicularia (spatulate with complete crossbar), extent of the gymnocyst, and small suboral costal field in Vitrimurella suggest a relationship with Figularia and with the cribrilinoidean genus Euthyroides Harmer, Reference Harmer1902 (Euthyroididae Levinsen, Reference Levinsen1909). An extant species from the Caribbean, Figularia? [sic] ampla Canu and Bassler, Reference Canu and Bassler1928b (USNM RB 7494), also has pseudoporous ooecia as well as traces of a well-developed costal field like some Figularia species, with pseudopores extending onto costae, hence resembling lumen pores; it is here reassigned as Vitrimurella ampla (Canu and Bassler, Reference Canu and Bassler1928b) n. comb., suggesting this family is better placed in Cribrilinoidea rather than Hippothooidea.

Among other species with multiple ectooecial pseudopores previously attributed to Figularia (considered doubtful by Rosso et al., Reference Rosso, di Martino and Ostrovsky2020), two fossil species—Membraniporella rugosa Maplestone, Reference Maplestone1901 from the Miocene of Australia and Figularia ryukyuensis Kataoka, Reference Kataoka1961 from the Pleistocene of Japan—are better assigned to Reginella or Hayamiellina Grischenko and Gordon in Grischenko et al., Reference Grischenko, Gordon, Nojo, Kawamura, Kaneko and Mawatari2004 rather than Figularia. A final conclusion concerning their placement, however, requires revision of the respective type material. Despite similarities among Figularia and four other cribrilinid taxa—Hayamiellina, Juxtacribrilina Yang et al., Reference Yang, Seo, Min, Grischenko and Gordon2018 (formerly regarded as a subgenus while here elevated to genus rank), Jullienula, and Reginella—these latter genera seem to be mutually more closely related by having a well-developed costal area with two or more costal lumina (pelmatidia), compared with core Euthyroididae and Vitrimurellidae.

At least three species with remarkable costate ooecia, Figularia pulcherrima Tilbrook, Hayward, and Gordon, Reference Tilbrook, Hayward and Gordon2001, Figularia jucunda Canu and Bassler, Reference Canu and Bassler1929, and Figulina tahitiensis Waters, Reference Waters1923, are here assigned to Costafigularia n. gen. (see the following). Figularia contraria Lagaaij, Reference Lagaaij1963, is quite distinct from Figularia (and from Costafigularia and Vitrimurella) in having smaller female zooids; this species requires reexamination and should be reassigned to a distinct genus (currently under review, J.E. Winston, personal communication, 2020).

Five species still require further investigation: Figularia? crassicostulata Canu and Bassler, Reference Canu and Bassler1920 (Priabonian, Florida, USA) and Figularia peltata (Reuss, Reference Reuss1874) (Miocene, Austria) apparently lack pelmata on the costae; Figularia duvergieri Bassler, Reference Bassler1936 (Miocene, France) has an orifice with denticulate anter; Figularia echinoides Brown, Reference Brown1952 (early Oligocene, New Zealand) has mostly two tubular pelmata on each costa and the ovicell was not seen, so it remains unknown if it was bifenestrate; Figularia kenleyi Brown, Reference Brown1958 (Miocene, Australia) has pelmata only on suboral costae.

Figularia elcanoi new species
Figure 4; Table 2

Figure 4.

Figularia elcanoi n. sp. (1) Holotype, MEF 6799, group of ovicelled zooids; the arrows show three pore plates on a vertical wall; (2) paratype, MEF 6800, ovicelled zooids; the arrows show the gymnocyst and the pair of processes on the lateral margins of the orifice; (3) holotype, MEF 6799, detail of spinocyst; (4) holotype, MEF 6799, detail of ovicells and frontal shields; the arrows show the pair of orificial processes. (1, 2) Scale bars = 200 μm; (3) scale bar = 50 μm; (4) scale bar = 100 μm.

Table 2.

Measurements (in μm) of Figularia elcanoi n. sp. (MEF 6800). N = number of measurements; SD = standard deviation.

Type material

Holotype: MEF 6799, a bilaminar fragment. Paratype: MEF 6800, a bilaminar fragment.

Diagnosis

Colony bilaminar. Zooids rectangular, more than three times longer than wide. Gymnocyst extremely reduced, barely visible proximally. Frontal shield formed by more than 6–12 pairs of alternating costae with a circular lumen pore near the midline. Orifice with a pair of processes on the lateral margins. Ectooecial fenestrae large, separated by a median suture.

Occurrence

This species was found only at its type locality, “Punta del Marqués”, Chubut Province, in the lower layers of Chenque Formation (early Miocene).

Description

Colony erect, bilaminar (Fig. 4.1). Ovicellate zooids rectangular, more than three times longer than wide, ordered quincuncially, separated by deep grooves, lateral margins almost straight (Fig. 4.1, 4.2). Orifice in ovicellate zooids wider than long, proximal margin straight or slightly concave, oral spines not seen, but a pair of blunt processes is constantly present on the lateral margins (Fig. 4.2, 4.4). Gymnocyst smooth, extremely reduced, barely visible proximally in ovicellate zooids (Fig. 4.2). The frontal surface is covered by a shield formed by more than 6–12 pairs of alternating costae separated by slits and fused along the midline; first pair of costae wider and more robust than the others, forming the proximal margin of the orifice; each costa with a small circular lumen pore near the midline (Fig. 4.3). Ovicell hyperstomial, ooecium globose, longer than wide, protruding over the proximal gymnocyst and the most proximal pairs of costae of the distal zooid, with two very large ectooecial fenestrae occupying most of its frontal surface, separated by a median suture (Fig. 4.2, 4.4). Communication between zooids by multiporous septula in the lateral walls (Fig. 4.1). Vicarious avicularia and ancestrula not seen.

Etymology

Honorific for the explorer Juan Sebastián Elcano, who commanded the flagship Victoria from the Philippines to Spain, completing the first circumnavigation of the Earth in 1522.

Remarks

Figularia elcanoi n. sp. is characterized by its bilamellar colonies, the extreme reduction of the gymnocyst, and the pair of large, deep fenestrae in the ovicell. In the two fragments examined, all the zooids seen on both sides of the colony were ovicellate. A gymnocyst not covered by the preceding ovicell can be seen in a zooid at the bottom of Fig. 4.2. The number of costae of each zooid could not be counted precisely because the proximal part of the spinocyst is covered by the ovicell of the preceding zooid.

Bilamellar colonies are not common in the genus Figularia. As far as we know, they occur in F. kenleyi Brown, Reference Brown1958 from the Miocene of southeastern Australia (Brown, Reference Brown1958) and in the recently described species F. spectabilis, from the early Pleistocene deepwater sediments of northeast Sicily, Italy (Rosso et al., Reference Rosso, di Martino and Ostrovsky2020).

Romero et al. (Reference Romero, Brezina, Bremec and Casadío2018) figured an unnamed encrusting Figularia from the Puerto Madryn Formation (early–late Miocene, Patagonia) growing on oyster shells. It is clearly related to F. elcanoi but differs in its narrower costae and shorter zooids and ectooecial fenestrae.

Type species

Figularia pulcherrima Tilbrook, Hayward, and Gordon, Reference Tilbrook, Hayward and Gordon2001. Recent.

Diagnosis

Colony unilaminar, encrusting. Frontal shield spinocystal, with costal field variable in length, surrounded by marginal gymnocyst; each costa with a single lumen pore; intercostal pores present. Vicarious avicularia often present. Ovicell hyperstomial, with costate ooecium.

Occurrence

Recent, Pacific.

Etymology

Costafigularia n. gen. is derived by combining the Latin costa, rib, with Figularia, alluding to the remarkable costate ooecia of this genus.

Remarks

Figularia pulcherrima Tilbrook, Hayward and Gordon, Reference Tilbrook, Hayward and Gordon2001 from the western Pacific has a well-developed smooth gymnocyst with a costal field like Figularia figularis, but it also has remarkable costate ooecia. Costafigularia n. gen. is here erected for three Indo-Pacific species: Costafigularia pulcherrima (Tilbrook, Hayward, and Gordon, Reference Tilbrook, Hayward and Gordon2001) n. comb. (type species; NHMUK 1998.8.4.71, Recent, Iririki Island, Efate, Vanuatu), Costafigularia jucunda (Canu and Bassler, Reference Canu and Bassler1929) n. comb. (USNM RB 7994, Recent, Phillipines), and Costafigularia tahitiensis (Waters, Reference Waters1923) n. comb. (Recent, Tahiti).

Costafigularia jucunda n. comb. (Canu and Bassler, Reference Canu and Bassler1929, p. 241, pl. 22, fig. 3) and C. tahitiensis n. comb. (Waters, Reference Waters1923, p. 571, pl. 18, fig. 1) are known only from their original publications; thus, a proper review and descriptions of the type material are still required.

Waters (Reference Waters1923, p. 571) characterized Costafigularia tahitiensis n. comb. as having a distinct ovicell “which has at the distal end an area with a number of large prominent papillae at the end of the rays.” His figure suggests a similar ooecial structure as in C. pulcherrima n. comb., with no clear differences in autozooid morphology between these two species. No type specimen of C. tahitiensis n. comb. has been located, but it is supposed that the type material is housed at the Manchester Museum, with other specimens studied by Arthur Waters. Costate ooecia described in both Costafigularia pulcherrima n. comb. and Costafigularia tahitiensis n. comb. are slightly different from ooecia with marginal pseudopores described in Costafigularia jucunda n. comb.

Jullienula is distinguished from Costafigularia n. gen. by having no lateral gymnocyst (gymnocyst well developed in Costafigularia), and its ooecium is reduced in size, possibly derived from one or two costae (ooecium hyperstomial with three or more costae in Costafigularia).

Type species

Membraniporella magellanica Calvet, Reference Calvet1904a. Smyth Channel, southern Chile. By monotypy.

Remarks

Parafigularia was erected by Moyano (Reference Moyano G.1984) for Membraniporella magellanica Calvet, Reference Calvet1904a (=Cribrilina patagonica Waters, Reference Waters1905) from the Magellanic region. The genus is characterized by fertile zooids with an ovicell connected with the avicularian chamber and modules composed of an autozooid, a distal avicularium, and a distalmost kenozooid. Two other species are included (Moyano, Reference Moyano G.2011), the subantarctic Parafigularia darwini Moyano, Reference Moyano G.2011 and the Antarctic Parafigularia discors (Hayward and Taylor, Reference Hayward and Taylor1984). Kuklinski and Barnes (Reference Kuklinski and Barnes2009) erected the genus Inferusia, with Inferusia taylori Kuklinski and Barnes, Reference Kuklinski and Barnes2009 from the Scotia Arc as type species. It is characterized, inter alia, by its triangular avicularia associated with ovicells. They compared Inferusia with three genera: Figularia, Filaguria Moyano, Reference Moyano G.1991, and Klugerella Moyano, Reference Moyano G.1991, but neglected to mention Parafigularia while also including Figularia discors in Inferusia (cf. Moyano, Reference Moyano G.2011). The type species of Parafigularia and Inferusia have in common the same kind of ovicell and modules composed of autozooids, avicularia, and kenozooids. They differ, however, in the development of the spinocyst (reduced in P. magellanica, greatly developed in I. taylori) and kenozooids (interspersed between autozooids in I. taylori, distal in P. magellanica). We rather believe that these are differences at the specific level and that Inferusia should be synonymized with Parafigularia due to similarities in their ovicells and modular structure.

Unfortunately, Moyano (Reference Moyano G.2011) inadvertently introduced Parafigularia darwini while being unaware of the existence of Inferusia taylori, described two years earlier (Kuklinski and Barnes, Reference Kuklinski and Barnes2009). The comparison of SEM images and the geographic proximity of the type localities of both nominal species leave no doubt regarding their synonymy, resulting in Parafigularia taylori (Kuklinski and Barnes, Reference Kuklinski and Barnes2009) n. comb.

Species of Parafigularia seem to be distinguishable from other species with a spinocystal frontal shield by having a broad poster with proximal corners with shallow indentations in the gymnocystal calcification. Similar indentations are also known in two genera, Eurystomella Levinsen, Reference Levinsen1909 and Integripelta Gordon, Mawatari, and Kajihara, Reference Gordon, Mawatari and Kajihara2002, currently placed in the catenicelloid family Eurystomellidae Levinsen, Reference Levinsen1909. Despite the morphological similarities between Parafigularia and Eurystomellidae, the relationship between these taxa remains uncertain and requires further studies.

Holotype

MLP 36404, fragment of a colony.

Figure 5.

Parafigularia pigafettai n. sp., holotype, MLP 36404. (1) General aspect of the colony; the left and middle arrows show two kenozooids; the right arrow shows the distalmost kenozooid; (2) detail of two zooids; (3) autozooids, ovicell, and avicularia; (4) detail of spinocyst. (1) Scale bar = 500 μm; (2, 3) scale bars = 100 μm; (4) scale bar = 50 μm.

Table 3.

Measurements (in μm) of Parafigularia pigafettai n. sp. (MLP 36404). N = number of measurements; SD = standard deviation.

Diagnosis

Parafigularia with spinocyst covering two-thirds of frontal shield, with 7–10 costae fused in the midline. Interzooidal avicularia distal to each infertile zooid, rostrum triangular, directed disto-laterally, transverse bar incomplete; with an associated distalmost kenozooid.

Occurrence

This species has so far been found only at the locality “Cabeza de León,” Monte León National Park, Santa Cruz Province. The specimens were found in the Punta Entrada Member of the Monte León Formation (early Miocene).

Description

Colony encrusting, unilaminar, multiserial. Autozooids subhexagonal, 370–590 × 200–370 μm, ordered quincuncially, separated by deep grooves (Fig. 5.1). Spinocyst convex, comprising about two-thirds of the frontal shield, formed by 7–10 flattened costae (frequently four or five pairs) separated by narrow intercostal spaces and fused in the midline; first pair of costae more robust and wider than the others (Fig. 5.2). Gymnocyst smooth, restricted to the lateral margins and the proximal third of the frontal shield. Orifice wider than long, bell-shaped, anter separated from the poster by proximolateral indentations, forming a pair of robust lateral condyle-like projections; proximal margin shallow W-shaped (Fig. 5.2, 5.4). Interzooidal avicularia distal to each infertile zooid, wider than long, opesia rounded, cryptocyst well developed, rostrum triangular, directed disto-laterally, transverse bar incomplete; sometimes with an associated distalmost kenozooid (Fig. 5.1). Fertile zooids with ovicell formed by the proximal cystid of the distal avicularium similar to those of infertile zooids; ovicell closed by the operculum, surface of ooecium smooth (Fig. 5.3). Zooids communicating via pore chambers in the disto-lateral walls (Fig. 5.1). Kenozooids consisting of gymnocyst, cryptocyst, and an elliptical opesia may occur between autozooids (Fig. 5.1).

Etymology

Honorific for the Italian scholar and explorer Antonio Alberto Pigafetta, who published the book Relazione del primo viaggio intorno al mondo after completing the first circumnavigation of the world together with Juan Sebastián Elcano.

Remarks

Parafigularia pigafettai n. sp. differs from the remaining species of the genus mainly in the number of costae and the area occupied by the spinocyst (4–7 costae covering a small area in P. magellanica, 9–10 pairs in P. discors, 9–11 pairs covering the whole frontal shield in P. taylori n. comb., 4–5 pairs occupying two-thirds of the frontal shield in P. pigafettai). The only colony examined encrusted the inner surface of a bivalve.