Fenestrulina variorugosa Rosso & Di Martino, 2025
|
publication ID |
https://doi.org/10.3897/zookeys.1254.157989 |
|
publication LSID |
lsid:zoobank.org:pub:959351D6-F778-4E6A-9AD5-97B9A190E0A7 |
|
DOI |
https://doi.org/10.5281/zenodo.17237603 |
|
persistent identifier |
https://treatment.plazi.org/id/9BB7F1AA-3432-5707-8BCB-5C43E1A623E0 |
|
treatment provided by |
|
|
scientific name |
Fenestrulina variorugosa Rosso & Di Martino |
| status |
sp. nov. |
Fenestrulina variorugosa Rosso & Di Martino sp. nov.
Figs 1 View Figure 1 , 17 View Figure 17 , 18 View Figure 18 , 19 View Figure 19 , 20 View Figure 20 , 22 View Figure 22 , 23 View Figure 23 , 24 View Figure 24 ; Tables 1 View Table 1 , 4 View Table 4
Fenestrulina malusii (Audouin): Zabala et al. 1993: list of species; Chimenz Gusso et al. 2014: 166 (pars), fig. 83 a, b. View in CoL
Type material.
Italy • Holotype 1 large ovicellate colony without ancestrula on a rhizome of Posidonia oceanica (Linnaeus) Delile. Mediterranean, Sicily Strait, W Sicily, Egadi Islands, Formica Isle, sample EFI 20 ; 37°59'14"N, 12°25'34"E; 8 m depth; Oct. 2007; scuba diving; A. Sinagra leg.; PMC.B 43.23.2.2024.a GoogleMaps . Italy • Paratype 1 juvenile colony including 15 non-ovicellate autozooids around the ancestrula; same details as the holotype; PMC.B 43.23.2.2024.b.1 GoogleMaps .
Other material examined.
Italy • 2 living colonies on the shell of a living specimen of Lithophaga lithophaga (Linnaeus, 1758) . Mediterranean, Italy, Tyrrhenian Sea, NE Sicily, Secca di Levante, Capo Milazzo Peninsula, sample MI_SdL_G ; 38°14'43"N, 15°14'26"E; 33 m depth; Coralligenous biocoenosis; 17 May 2024; scuba diving; G. Donato leg.; PMC Rosso-Collection I.H.B.160.a.1 a GoogleMaps . Italy • 5 living colonies on a soft-bodied alga. Mediterranean, Italy, Tyrrhenian Sea, NE Sicily, Secca di Ponente, Capo Milazzo Peninsula, sample MI_SdPn_G ; 38°16'28"N, 15°13'22"E; 33 m depth; Coralligenous biocoenosis; 6 May 2024; scuba diving; G. Donato leg.; PMC Rosso-Collection I.H.B.160.a.1 b GoogleMaps . France • 1 living colony with only few functional autozooids on a dead coral fragment. Mediterranean, Liguro-Provençal basin, Cassis, Cassidaign Canyon ; no coordinates available; 300 m depth; Bathyal Corals biocoenosis; 21 Jun. 1969; J.-G. Harmelin leg.; PMC J-GH-Collection F.H.B.160.b.1 . France • 5 living ovicellate and non-ovicellate colonies on a bioconcretion of white corals hosting a few Crania anomala (Müller, 1776) specimens and several cryptic arciid and mytilid bivalves, with 1 out of 5 colonies encrusting the outer shell of an arciid. Mediterranean, Liguro-Provençal basin, off Banyuls-sur-Mer, sample 5 ; 42°4.30'N, 3°25'E; 200–300 m depth; Bathyal Corals biocoenosis; Jun. 1984; J.-G. Harmelin leg.; PMC J-GH Collection F.H.B.160.c GoogleMaps . France • 1 living colony encrusting the underside of a stone. Mediterranean, Liguro-Provençal basin, Port Cros Park, Cave of the Bagaud Island ; 43°00.9'N, 6°21.6'E; 7 m depth; June 1984; J.-G. Harmelin leg.; PMC J-GH Collection F.H.B.160.d GoogleMaps . France • 1 living colony encrusting the inner surface of an empty shell of a dead Pinna nobilis (Linnaeus, 1758) . Mediterranean, Liguro-Provençal basin, Veyron Plateau , off Marseille; coordinates not available; 24 m depth; 23 Sep. 1983; J.-G. Harmelin leg.; PMC J-GH Collection F.H.B.160.e . Tunisia • 2 colonies. Mediterranean, Sicily Strait, off Tabarka ; coordinates not available; 86 m depth; J. Jullien Collection; MNHN_IB_2008_2590 .
Diagnosis.
Fenestrulina with lobate pseudopores characterised by three or four irregularly curving spinules, thickening and flattening towards the centre without meeting, arranged in a single row, some becoming semicircular, leaning against rim of frontal shield; globose ovicell rimmed by several small peripheral pores with variable endooecial ornamentation, ranging from gently nodular, to faintly ribbed and scalloped at the periphery, or prominently rough with radial crests.
Description.
Colony encrusting, multiserial, unilaminar (Fig. 17 A View Figure 17 ), irregularly shaped in relation to the substrate morphology, ~ 1 mm in diameter. Interzooidal communications via pore-chambers, two proximolateral, two distolateral (130–150 μm), one distal (~ 120 μm) (Figs 18 A, B, E View Figure 18 , 19 C, F View Figure 19 ), each with 6–10 round pores, 6–9 μm in diameter (Fig. 18 B View Figure 18 ).
Autozooids hexagonal, distinct, contiguous, boundaries marked by narrow, deep grooves widening at triple junctions, exposing upper parts of sub-vertical lateral walls (Figs 17 B – E View Figure 17 , 19 A – E View Figure 19 ). Frontal cryptocystidean area outlined by a thin, slightly raised rim, more pronounced at pseudopore level, lining orifice proximally and laterally, developing long (mean length 148 μm, n = 11) lappets on both sides of orifice (Figs 17 B – E View Figure 17 , 18 A, F, G View Figure 18 , 19 A – C, F, G View Figure 19 ). These lappets occasionally extending distally and almost encircling the orifice but never joining (Fig. 19 A View Figure 19 ). Two elliptical, occasionally one elongate, cryptocystidean areas distal to the orifice, between oral spines, each with one or two pseudopores, rarely more (Figs 17 B – E View Figure 17 , 18 A, C, F View Figure 18 , 19 F, G View Figure 19 ). Frontal surface gently convex, more raised at ascopore level, smooth, perforated by 32–45 pseudopores, 12–20 in periancestrular autozooids (Figs 18 E View Figure 18 , 19 C View Figure 19 ). Pseudopores mostly located in distal half of autozooid, arranged in two rows between orifice and ascopore, one or two rows in lateral lappets (Figs 17 B, E View Figure 17 , 18 A, C, F View Figure 18 ), in a single row along lateral zooidal margins, sparse or absent proximally (Figs 17 C – E View Figure 17 , 19 View Figure 19 ). Pseudopores on a level with frontal shield, irregularly subcircular to slightly lobate, semicircular (Figs 17 D View Figure 17 , 19 View Figure 19 ) or slit-like (Fig. 17 E View Figure 17 ) along lateral rim; each with three or four (rarely more) spinules projecting, thickening and flattening or branching centrally, never merging (Figs 18 C, D, F View Figure 18 , 23 H View Figure 23 ). Circular pseudopores without spinules in regenerated autozooids in damaged colony areas (Fig. 20 C – F View Figure 20 ), sometimes occluded by underlying gymnocystal calcification (Fig. 20 D, E View Figure 20 ). Basal wall largely uncalcified.
Primary orifice transversely D-shaped, hinge-line straight with two tiny denticles near proximal corners; distal rim fairly denticulated (Fig. 18 C View Figure 18 ). Two oral spines in most autozooids (Figs 17 B – E View Figure 17 , 18 C, F, G View Figure 18 ), rarely three (Figs 18 G View Figure 18 , 19 G View Figure 19 ), four observed in the first periancestrular ones (Fig. 18 E View Figure 18 ), ~ 100 μm long (base diameter 15–25 μm), located along distal curvature (Figs 17 B – E View Figure 17 , 18 C, G View Figure 18 ). Intrazooidal regeneration may alter spine count (e. g., Fig. 18 F View Figure 18 ). Only two, barely visible, spines in ovicellate autozooids, lining ovicell margins near proximal rim (Figs 17 C, D View Figure 17 , 19 D, H View Figure 19 ).
Ascopore ~ 90 μm proximal to orifice (Figs 17 View Figure 17 , 18 A, C, D, G View Figure 18 , 19 F, G View Figure 19 ), within a reniform field of smooth gymnocyst marked by a slightly raised rim, in contact with the arched proximal rim of the frontal shield in the presence of an ovicell (Fig. 17 E View Figure 17 ) or fusing with it (Fig. 19 D, H View Figure 19 ); large transversely C-shaped lumen between the distal wide tongue and the arched proximal border; rim irregularly denticulated including tiny spindle-like spinules and larger, platy to branched denticles (Figs 17 B, E View Figure 17 , 18 C, D View Figure 18 , 19 F, G View Figure 19 ).
Ovicell subglobular, prominent, restricted proximally to fit orifice width, slightly obscuring distal part of orifice, not closed by the operculum, produced by the distal autozooid (Figs 17 C, D View Figure 17 , 19 D, H View Figure 19 ). Endooecium well calcified, gently nodular and faintly ribbed and scalloped to prominently rough with blunt spiny processes and radial crests at periphery but smoother proximally; rimmed by a row of ~ 15 quadrangular pores, separated by calcified bridges, often reduced in diameter by secondary calcification (Fig. 17 C View Figure 17 , arrowed); proximal margin with narrow upturned rim just at corners above oral spines. Calcified part of ectooecium consisting of a narrow (~ 30 μm) elevated rim of gymnocyst lining the row of pores.
Ancestrula tatiform (Figs 18 E View Figure 18 , 19 C View Figure 19 , 24 F, G View Figure 24 ), irregularly oval, similar size to first periancestrular autozooids, gymnocyst apparently narrow, largely covered by periancestrular autozooids in examined material, with ten spines: five around orifice (three distal, closely spaced, two more proximally placed, at a greater distance, aligned with proximal margin of operculum), five around proximal half of opesia (widely and regularly spaced). Opesia oval (305 μm long by 220 μm wide) surrounded by a narrow (~ 15 μm), almost smooth cryptocyst. Two longitudinally elongated cryptocystidean areas (2 or 3 pores each) between distal triad and two more proximal oral spines (Fig. 18 E View Figure 18 ). Budding pattern: one distal, two distolateral, two proximolateral and two proximal autozooids (Fig. 18 E View Figure 18 ).
Kenozooids small, triangular to quadrangular, elongate, filling empty spaces between autozooids in areas without evidence of reparation, including few relatively large pseudopores with 5–7 denticles giving a stellate appearance (Figs 18 G View Figure 18 , 19 H View Figure 19 ).
Etymology.
Referring to the variability of the ovicell ornamentation, especially the variable degree of the endooecial rugosity.
Remarks.
Fenestrulina variorugosa sp. nov. resembles F. barrosoi Álvarez, 1993 , an Atlanto-Mediterranean species described from the Alboran Sea at depths of 15–20 and 50–60 m ( Álvarez 1993), and later recorded at 112–120 m depth ( Ramalho et al. 2022), and from the Galician coast on seagrasses at ~ 15 m depth ( Reverter Gil et al. 2019). Both species share the general morphology of ovicells and autozooids, including the shape and location of the ascopore. However, all morphological measurements, except for the autozooidal length of the holotype, tend to be smaller in F. barrosoi than in F. variorugosa sp. nov. In F. barrosoi , pseudopores are always distributed in a single row along lateral and proximal margins, without any doubling or absence in these areas, as observed in F. variorugosa sp. nov. Fenestrulina barrosoi typically has 3–6 oral spines, most commonly four or five based on Álvarez (1993: 833), although his fig. 1 on p. 832 shows zooids with three spines, rather than the 2–4 (mostly two), seen in F. variorugosa sp. nov. Importantly, the proximal pair of spines in F. barrosoi is stout and shifted proximally. Colonies with rugose ovicells resemble F. kalliste sp. nov., but the latter species has heavier ornamentation, with more developed spiny processes and a central area with prominent and often transversal crests. The pseudopore number, distribution and morphology differ significantly, and the proximal oral spines in F. kalliste sp. nov. are bifurcated. Rough ornamentation of the endooecium is also seen in F. cavernicola sp. nov. and F. juani , but in these species, the frontal shield is dimpled. In F. cavernicola sp. nov., the endooecium is spinier and bordered by a large peripheral fissure with only a few pores, while in F. juani , the ornamentation consists of prominent tubercles.
The variability in F. variorugosa sp. nov. primarily pertains to the ornamentation of the ovicell endooecium, which ranges from nearly smooth to highly rough. Autozooid size and shape also vary. Although usually elongate hexagonal and arranged in regular alternating rows, repair in some damaged areas may cause changes in size and shape of some autozooids, including the formation of lateral prominences and relatively enlarged orifices (Figs 18 E View Figure 18 , 19 A – C View Figure 19 ). Orifices also seem to be dimorphic, becoming slightly wider in ovicellate than in non-ovicellate zooids (Figs 17 C, D View Figure 17 , 19 H View Figure 19 ). In a single case, a deformed autozooid had an orifice as long as (133 μm vs 112–137 μm) but decidedly wider (227 μm vs 144–170 μm) than average (Fig. 19 B View Figure 19 ). Frontal pseudopores are very close to each other and can often fuse. The denticles on the proximal side of the orifice are very inconspicuous and barely recognisable in some colonies.
Regeneration is common in this species, particularly in specimens living on soft-bodied algae from the Capo Milazzo area (Figs 19 View Figure 19 , 20 View Figure 20 ). In a colony from sample MI _ SdL _ G, autozooids were mostly damaged at orifice level, with intramurally budded autozooids showing smaller orifices and limited proximal areas of the frontal shield, usually extending distal to the ascopore (Fig. 19 E, H, I View Figure 19 ). Heavier damage, also affecting larger sectors of the autozooidal frontal shield, was observed in a colony from sample MI _ SdPn _ G (Fig. 20 View Figure 20 ), including multiple regeneration events per autozooid (Fig. 20 A, C View Figure 20 ), regeneration with opposite polarity (Fig. 20 B View Figure 20 ), and the production of oral closure plates (Fig. 20 B, C View Figure 20 ). Detachment of the original frontal shield after regeneration left traces in the regenerated one (Fig. 20 E View Figure 20 , arrowed). Regeneration in damaged areas produced elevated irregularly shaped autozooids with circular pseudopores lacking spinules, overarching the underlying colony layer (Fig. 20 F View Figure 20 ). Similar non-spinulose pseudopores occur in F. caseola Hayward, 1988 originating from Mauritius and later reported from other Indo-Pacific localities including Australia ( Hayward 1988; Tilbrook 2006; Bock 2025 https://bryozoa.net/cheilostomata/fenestrulinidae/fenestrulina_caseola.html). However, unlike in F. variorugosa sp. nov., those pseudopores can be occluded by the complete coalescence of spinous processes.
Colonies figured in Chimenz Gusso et al. (2014) as F. malusii resemble F. variorugosa sp. nov. in their general appearance, including the size, morphology and location of pseudopores against the frontal cryptocystidean rim. The two distal spines and the slightly ribbed ovicell are also similar. Colonies from a depth of 86 m off Tabarka ( Tunisia) are highly likely to correspond to this species, based on stereomicroscope images kindly provided by Dr. P. Lozouet.
Habitat distribution.
To date, Fenestrulina variorugosa sp. nov. has been collected from a variety of habitats ranging from shaded and plant-rich areas on the shallow shelf to the upper slope, in association with white corals. Our colonies were found on roots of P. oceanica , collected in a flat, rocky area predominantly covered by algae from the Infralittoral Algae biocoenosis, surrounded by the Posidonia Meadows biocoenosis ( CoNISMa 2009), as well as on soft-bodied algae forming the canopy of the Coralligenous biocoenosis. A few colonies were also observed colonising the shell of a L. lithophaga specimen still inside its bore-hole in the coralligenous concretion, suggesting that this species can thrive in cryptic habitats. A few colonies were found on dead coral fragments between 200 and 300 m depth in canyons off Spain and France (partly published in Zabala et al. 1993). However, the habitat distribution may be incomplete, as the information regarding the habitats of the specimens examined by Chimenz Gusso et al. (2014) and those from Tabarka ( Tunisia) at the MNHN is not available.
Geographical distribution.
In addition to its type locality in the Egadi Archipelago (W Sicily), F. variorugosa sp. nov. has also been found in the Aeolian Archipelago (SE Tyrrhenian Sea) and in the north-western part of the Liguro-Provençal basin. The species distribution is further expanded when considering the colonies from off Tabarka ( Tunisia), extending its occurrence also to the southern part of the Sicily Strait. Most colonies examined by Chimenz Gusso et al. (2014) were collected from several localities in the Tyrrhenian Sea, and subordinately from the Aegean Sea ( Turkey), further expanding the known distribution of this species in the Mediterranean.
| MNHN |
Museum National d'Histoire Naturelle |
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
|
Kingdom |
|
|
Phylum |
|
|
Class |
|
|
Order |
|
|
SuperFamily |
Schizoporelloidea |
|
Family |
|
|
Genus |
Fenestrulina variorugosa Rosso & Di Martino
| Rosso, Antonietta, Di Martino, Emanuela, Donato, Gemma, Figuerola, Blanca, Gerovasileiou, Vasilis, Siddiolo, Chiara, Sinagra, Alessandro, Sanfilippo, Rossana & Sciuto, Francesco 2025 |
Fenestrulina malusii (Audouin): Zabala et al. 1993 : list of species; Chimenz Gusso et al. 2014: 166 (pars), fig. 83 a, b.
| Chimenz Gusso C & Nicoletti L & Bondanese C 2014: 166 |
| Fenestrulina malusii (Audouin): Zabala et al. 1993 : list of species; Chimenz Gusso et al. 2014: 166 (pars), fig. 83 a, b. |