CAENOGASTROPODA Cox, 1960
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publication ID |
https://doi.org/ 10.1080/00222933.2017.1407833 |
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persistent identifier |
https://treatment.plazi.org/id/AB5AD758-4951-FF85-FE4A-C57AFD9AFD88 |
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treatment provided by |
Felipe |
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scientific name |
CAENOGASTROPODA Cox, 1960 |
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Subclass CAENOGASTROPODA Cox, 1960 View in CoL
Family CERITHIOPSIDAE Adams and Adams, 1853 View in CoL Type species Murex tubercularis Montagu, 1803 by original designation Cerithiopsis horrida di Monterosato, 1874
( Figures 1 View Figure 1 , 2 View Figure 2 (a–m), 3(a))
Confirmed records
Cerithiopsis horrida (Jeffreys ms.) – di Monterosato 1874a: 273.
Cerithiopsis horrida – di Monterosato 1874b: 360.
Cerithiopsis horrida Jeffreys – di Monterosato 1878: 156; di Monterosato 1890: 163.
Cerithiopsis horrida ( Jeffreys, 1885) – Cachia et al. 2001: 186 (plate 26, figure 6a, b); Mifsud 2003: 41 (figure 6a, b); Bartolini 2006: 20 (figure 1a, b); Cossignani and Ardovini 2011: 16, 165 (figures).
Cerithiopsis horrida ‘Jeffreys’ Monterosato, 1874 – *D’ Angelo and Gargiullo 1978: 115 (figure 1).
Cerithiopsis horrida Jeffreys, 1885 – *Warén 1980: 25; Giannuzzi-Savelli et al. 1999: 14, 40–41 (figures 64, 65); Perna 2013: 121 (figure).
Cerithiopsis sp. – Cecalupo and Giusti 1989: 98, 101 (figures 5, 5a).
Cerithiopsis atalaya Watson, 1874 sensu Giannuzzi-Savelli et al. 1999 View in CoL (partim): 34–35 (figure 29).
Cerithiopsis horrida Monterosato, 1874 – Peñas et al. 2006: 68 (figure 75), 72 (figures 95, 96), 74; Peñas et al. 2008: 18; Segers et al. 2009: 35, 92, 412–413 (figure 4); Rolán 2011: 93, 454–455 (figure 27m, o); Scaperrotta et al. 2016: 13, 48 (figure).
Doubtful records
Cerithiopsis horrida Jeffreys ms. – Jeffreys 1874: 114.
Cerithiopsis horrida Jeffreys – Jeffreys 1885: 60 (plate 6, figures 9, 9a); Locard 1898: 381;
Marshall 1911: 194; Sykes 1925: 182–182 (Stations 50 – Mediterranean; Adventure
Bank); Bellini 1929: 42. Cerithiopsis horrida ( Jeffreys, 1885) – Nordsieck 1976: 6–7 (figure 11). Cerithiopsis tubercularis horrida Jef. – Hernández Otero 1975: 118. Metaxia horrida – Silesu and Sosso 1987: 66. Cerihiopsis [sic] horrida Monterosato – De Smit and Bába 2001: 98. Cerirhiopsis [sic] horrida (Monterosato, 1874) – Manousis and Galinou-Mitsoudi 2014: 5.
Incorrect records
Cerithiopsis tubercularis horrida (Jeffreys) – Tenekides 1989: 32, 96 (figure 8) (= Cerithiopsis atalaya Watson, 1885 ).
Cerithiopsis horrida ‘Jeffreys’ Monterosato, 1874 – MiliŠiĆ 1991: 201 (figure); MiliŠiĆ 2007: 202 (figure) (= Metaxia sp. ).
Type locality/localities
di Monterosato (1874a) described Cerithiopsis horrida di Monterosato, 1874 based on one fragment from San Vito Lo Capo and ‘beautiful’ shells from Palermo (north-western Sicily, southern Italy) and ‘an additional locality’. According to the International Code of the Zoological Nomenclature ( ICZN 2012: art. 73, 76), the abovementioned sites should be listed as type localities. However, based on our research, material from San Vito Lo Capo and the ‘additional locality’ is lacking in MCZR, while specimens from Palermo are divided into three different vials (seven sh + two sh + one sh) (see below in Material examined). This material does not properly match any statement by Monterosato, who, soon after the description, further listed additional findings from San Vito Lo Capo ( di Monterosato 1874b), one fragment from Palermo ( di Monterosato 1878), and presumably additional specimens from Palermo ( di Monterosato 1890). Therefore, no certain type material could be located in MCZR.
Material examined
Cerithiopsis horrida di Monterosato, 1874 . Spain (Mediterranean Sea): Punta de la Mona – La Herradura, September 2006, 2 sh ( SBC), 40 m, in biogenic substrate, SCUBA diving on coralline bottom (reported in Bartolini 2006; Scaperrotta et al. 2016) ( Figure 2 View Figure 2 (h, i)); Seco de los Olivos, off Almería, 10 April 2004, 22 sh and frag ( CBC, CSC, and FSC), 98.2–108.8 m, biogenic substrate dredged on rocky bottom (partially reported in Scaperrotta et al. 2016). Italy (Mediterranean Sea): Palermo (Sicily), 10 sh ( MCZR – Monterosato collection – lot. 21173), 50–200 m; off Capraia Island (Livorno), 2010, 2 sh ( CSC), 400 m, biogenic substrate dredged on muddy bottom. Malta (Mediterranean Sea): off Ras il-Wahx, July 1992, 10 sh ( CMC), 120–160 m, dredged in muddy sand and dead coralline algae (partially reported in Cachia et al. 2001) ( Figure 3 View Figure 3 (a)); off Ras il-Wahx, August 2002, 1 lv (dissolved to obtain the radula; radula and operculum deposited in ERC) ( Figure 2 View Figure 2 (a–c, j)), (reported in Mifsud 2003) and 1 sh ( CMC) ( Figure 2 View Figure 2 (d–g)), 120 m, live attached to old line with sponges and other fauna; RV/URANIA – Marcos Cruise, MS 44-Heavy Dredge, start 35°30.506 ʹ N, 14°06.230 ʹ E, 632 m, end 35°31.228 ʹ N, 14°05.698 ʹ E, 467 m, 12 April 2007, 1 sh ( CMC); off north Gozo (36°12.370 ʹ N, 14°23.559 ʹ E), 15 September 2011, 1 lv ( CMC), 350 m, attached to sponges and coralline algae on old sunken fishing lines, in fisherman’ s by-catch; off Ras il-Qaws, 20 May 2011, 1 sh ( NMNHM – 2274), 80–100 m, on coralline algae; off Ras il-Wahx, June 2014, 1 sh ( NMNHM – 3705), 120–140 m, dredged in muddy sand and gravel.
Cerithiopsis horrida Jeffreys, 1885 . Unknown Mediterranean locality: 1 sh ( NHMUK 1885.11.5.3910) ( Figure 2 View Figure 2 (k)). Algeria (Mediterranean Sea): Râs el Amoûch (Tipaza), 2 frag ( NHMUK 1885.11.5.3932–3933) ( Figure 2 View Figure 2 (l, m)).
Distribution
As with many north-eastern Atlantic/Mediterranean molluscan taxa, analysis of the literature on Cerithiopsis horrida di Monterosato, 1874 revealed a chaotic situation, with recent authors often reporting highly inconsistent data or merely copying older literature. Regarding its Atlantic distribution, some researchers considered it as limited to the Lusitanian Sea and Madeira (Nordsieck 1968); including these previous areas and extending it north as far as England (Nordsieck 1976, 1982; Cachia et al. 2001); ranging from Angola to the Lusitanian seamounts, including Azores, Madeira Archipelago, and Canary Islands (Segers et al. 2009); limited to Morocco and Canary Islands (Rolán 2011); or ranging from Gibraltar to Morocco, including Lusitanian seamounts ( Gofas et al. 2012). With regards to its Mediterranean distribution, some authors considered it as limited to some areas [Lusitanian Sea, Sicily ( Italy), and Brindisi (Apulia, Italy): Nordsieck 1976, 1982; southern parts of the Basin: Giannuzzi-Savelli et al. 1999; western parts of the Basin: Cachia et al. 2001; western parts of the southern Mediterranean: Gofas et al. 2012] or the entire basin (D’ Angelo and Gargiullo 1978). Unfortunately, the majority of the statements reported above were not supported by documented material. Acknowledging the risk of discarding some true distributional data, and considering as confirmed only records based on specimens figured and/or examined by us, C. horrida shells were found with certainty in Madeira ( Portugal) (Segers et al. 2009) and Canary Islands ( Spain) (Rolán 2011) in the Atlantic Ocean, and in Spain (e.g. Giannuzzi-Savelli et al. 1999; Bartolini 2006; Peñas et al. 2006), Italy (e.g. di Monterosato 1874a, 1874b; Cecalupo and Giusti 1989), and the Maltese Archipelago ( Cachia et al. 2001; Mifsud 2003) in the Mediterranean Sea, although living specimens were only found in the latter country. Additional potential records come from the UK ( Marshall 1911) and Angola (Segers et al. 2009) in the Atlantic, and from Algeria ( Jeffreys 1874, 1885; Sykes 1925), between Italy and Tunisia (Sykes 1925), Greece ( Manousis and Galinou-Mitsoudi 2014), and Turkey ( Jeffreys 1874) in the Mediterranean, but they were not confirmed here by the analysis of well-documented specimens. In support of our statements, C. horrida was never subsequently reported among cerithiopsids occurring in the UK despite careful research ( Fretter and Graham 1982; Graham 1988), ‘specimens in poor condition’ from Angola may more likely be assigned to Cerithiopsis leopardus Rolán and Gori, 2013 recently described from a nearby geographic area (see Rolán and Gori, 2013 and discussion below), and no records are known between these two remote sites and the more plausible distributional range. In addition, the Algerian records often reported in the literature are uncertain (see below in Remarks), the De Smit and Bába (2001) and Manousis and Galinou-Mitsoudi (2014) records from Greece were impossible to check as no material/photos were available to us [authors’ data: see also Crocetta et al. (2017) for comments on the latter articles], and this taxon was not even doubtfully included in a recent review of molluscan taxa living along coast of Turkey (Öztürk et al. 2014). However, the same doubts also hold true for the following studies from Madeira ( Portugal: Jeffreys 1885; Locard 1898) and Canary Islands ( Spain: Hernández Otero 1975) in the Atlantic, and from off Pantelleria Island (Sykes 1925), the Gulf of Naples ( Bellini 1929), Oristano area (Silesu and Sosso 1987), and Brindisi (Nordsieck 1976) in Italy (Mediterranean), despite the confirmed presence of C. horrida in all these countries. These zoogeographic data are depicted in Figure 1 View Figure 1 .
Ecology and bathymetric range
Little is known about the general ecology and feeding behaviour of this species. Confirmed findings of empty shells come from samplings ranging from 30 to 632 m depths ( di Monterosato 1875; Material examined), although we cannot exclude some of them may be based on subfossil material. Living specimens were found in rocky
substrata with coralline algae and encrusting sponges at 120–350 m depth ( Mifsud 2003; Material examined).
Description
Animal body colour entirely white. Cephalic tentacles long and cylindrical, with black eyes at bases. Foot long and narrow, with a somewhat squared anterior end extending well beyond the head ( Figure 2 View Figure 2 (a)). Radula formula 2 + 1 + 1 + 1 + 2 ( Figure 2 View Figure 2 (b)). Central tooth short, broad, cusps conical, with formula 4 + 1 + 4, the median cusp being the largest ( Figure 2 View Figure 2 (b)). Lateral teeth short, very broad, with nine conical cusps, the second cusp being the largest ( Figure 2 View Figure 2 (b, c)). Marginal teeth comb-like, cusps conical. Inner marginal teeth with formula 2 + 5–6, the first two very slender and filamentous. Morphology of outer marginal teeth not well visible, but presumably with the same formula of inner marginal teeth ( Figure 2 View Figure 2 (c)). Outer teeth narrower than inner teeth ( Figure 2 View Figure 2 (b, c)).
Shell solid, up to 12 mm in height, turriculate, with a regularly conical outline ( Figure 2 View Figure 2 (a, k–m)). Protoconch conical-cylindrical, of 3–4 highly convex whorls, light brown–yellowish in colour ( Figure 2 View Figure 2 (d)). Embryonic shell (protoconch I) of 1.5 whorls, rounded, sculptured by microgranules somewhat spirally arranged, united by very thin threads ( Figure 2 View Figure 2 (e)). Transition area between embryonic shell and larval shell characterized by granules merging into several irregular and beaded spiral threads, subsequently becoming unbeaded and remaining in number of three only ( Figure 2 View Figure 2 (f)). Larval shell (protoconch II) sculptured by axial ribs crossed by three weaker spiral threads. Axial ribs flexuous, opisthocline, much narrower than interspaces, extending from suture to suture (13–16 on last protoconch whorl) ( Figure 2 View Figure 2 (g)). Three widely spaced spiral threads running in the median part of the whorls ( Figure 2 View Figure 2 (f, g)). Transition area between protoconch and teleoconch seems to be marked by a gradual ornamentation change, although only a shell with signs of damage was available to us ( Figure 2 View Figure 2 (g)). Teleoconch of up to 15 flat-sided to slightly convex whorls, sculptured with axial ribs (18–21 on last whorl), and four spiral cords on the last whorl, forming pointed nodules at the intersections, slightly shouldered due to a pronounced ridge on their adapical edge ( Figure 2 View Figure 2 (a, h, k–m)); see also Peñas et al. 2006). Teleoconch cords 2 and 3 starting soon after protoconch–teleoconch transition on the same line of 1st and 3rd protoconch spiral threads, cord 1 starting on 3rd whorl as a subsutural thread, progressively increasing in thickness, becoming equivalent to cords 2 and 3 on 6th–7th whorl (see also Peñas et al. 2006). Cord 4 nodulose, appearing on last whorl as a continuation of the marked suture ( Figure 2 View Figure 2 (h)). Base concave and smooth, except for sinuous growth lines and a spiral ridge encircling columella ( Figure 2 View Figure 2 (i)). Subquadrangular and small aperture, extending into a short, inclined siphonal canal, ending with a sinuous edge ( Figure 2 View Figure 2 (h, i)). Columella straight or barely concave, with an evident callus ( Figure 2 View Figure 2 (h, i)). Outer lip barely thickened inside, slightly crenulated by external sculpture ( Figure 2 View Figure 2 (h)). Teleoconch whitish cream to tan in colour, with darker brown flecks irregularly distributed, base dark brown. Periostracum very thin, transparent.
Operculum horny, thin, almost transparent, ovoidal, and paucispiral, with marginal nucleus ( Figure 2 View Figure 2 (j)).
Remarks
Cerithiopsis horrida di Monterosato, 1874 has a troublesome nomenclatural and taxonomic history. The binomen was first mentioned, without description, by Jeffreys (1874) from Râs el Amoûch ( Algeria) and Izmir (as Smyrne, Turkey, based on MacAndrew’ s material), and one year later made available by Monterosato, who applied it to shells from San Vito Lo Capo and Palermo (Sicily, Italy), using a ‘manuscript name’ by Jeffreys ( di Monterosato 1874a). Finally, 10 years later, Jeffreys (1885) described Cerithiopsis horrida , presumably based on the same taxon described by Monterosato (but see below). Jeffreys’ (1874) usage is a nomen nudum ( ICZN 2012: glossary) and not available, while di Monterosato’ s (1874a) description, despite being devoid of illustrations, contained peculiar distinctive traits, allowing easy identification even by past authors (e.g. the distinctive axial ribs on the protoconch). This leads to the conclusion that Cerithiopsis horrida Jeffreys, 1885 is a primary homonym of Cerithiopsis horrida di Monterosato, 1874 , and has to be considered as permanently invalid ( ICZN 2012, art. 57.2). In spite of that, several authors incorrectly listed Jeffreys as the proper authority (e.g. Nordsieck 1976, 1982; Warén 1980; Giannuzzi-Savelli et al. 1999; Cachia et al. 2001; Mifsud 2003; Bartolini 2006; Cossignani and Ardovini 2011). This chaotic situation was presumably amplified by Warén (1980), who considered C. horrida Monterosato a nomen nudum and listed this taxon as C. horrida Jeffreys , and by Giannuzzi-Savelli (1985), who first reported Warén’ s (1980) statements as incorrect and highlighted the validity of di Monterosato’ s (1874a) description, but subsequently discussed and figured Cerithiopsis horrida Jeffreys as valid, and Cerithiopsis horrida Monterosato as a nomen nudum ( Giannuzzi-Savelli et al. 1999).
Apart from these nomenclatural issues, C. horrida has been often misidentified, placed in inappropriate genera, or even neglected as a valid species in the past. Attention to its taxonomic identity was presumably first drawn by D’ Angelo and Gargiullo (1978), who figured a specimen from the Monterosato collection – MCZR (M. Appolloni, pers. comm.). Despite that, similarities with Cerithiopsis atalaya Watson, 1885 led to some published misidentifications [e.g. Tenekides (1989) and Giannuzzi-Savelli et al. (1999): cf. also Peñas et al. (2006) for discussions on the latter work]. However, shells of C. atalaya morphologically differ from those of C. horrida in protoconch ( C. atalaya : embryonic shell rugose, larval shell sculptured with two median parallel chords crossed by close-set axial riblets) and teleoconch ( C. atalaya : spiral nodules rounded, fourth cord on the last whorl not beaded, more uniform colour) features ( Gofas et al. 2012). In addition, the general rarity of specimens, almost always lacking intact protoconchs (shells with entire protoconchs are, however, present among Monterosato’ s material: specimens to be figured in a paper dealing with the Monterosato collection – M. Appolloni, pers. comm.), as well as the absence of anatomical studies, led to discussions of its generic attribution in the past. Originally described as belonging to Cerithiopsis Forbes and Hanley, 1850 (see di Monterosato 1874a), Warén (1980) suggested it belonged to Metaxia Monterosato, 1884 after the examination of the material in the NHMUK, a taxonomic rearrangement that was blindly followed or doubtfully accepted by several authors (e.g. Piani 1980; Bruschi et al. 1985; Orlando and Palazzi 1985; Silesu and Sosso 1987; Delamotte and Vardala-Theodorou 1994; Giannuzzi-Savelli et al. 1999). Indeed C. horrida is similar to species of the genus Metaxia at a first glance, as also suggested by further misidentifications by MiliŠiĆ (1991, 2007) who reported and figured Metaxia specimens from Croatia identifying them as Cerithiopsis horrida ‘Jeffreys’ Monterosato, 1874, and by Nordsieck (1968) (see below). However, Jeffreys’ material was re-examined by us and is figured for the first time here, and consists of a fairly well-preserved shell from the Mediterranean (NHMUK 1885.11.5.3910) ( Figure 2 View Figure 2 (k)) that clearly belongs to C. horrida , and two other worn shells from Râs el Amoûch (NHMUK 1885.11.5.3932–3933) ( Figure 2 View Figure 2 (l, m)), which may be conspecific. That material does not match Metaxia species at all, having different protoconchs and teleoconch sculptures (see Bouchet 1985; Fernandes and Pimenta 2011; Table 1; Figure 3 View Figure 3 (c)), and therefore Warén’ s (1980) statements are incorrect. On the other hand, we doubt that the NHMUK material is really Jeffreys’ syntypes. In fact, Jeffreys (1885) described the species as from ‘Med. St. Rasel Amoush’. If we consider the previous sentence as correct (which should therefore only involve material from Râs el Amoûch), the specimen originally figured by Jeffreys (1885) does not match the material hosted in NHMUK ( Figure 2 View Figure 2 (l, m)). If we consider the previous sentence as incorrect (it seems to lack the number of the stations between ‘Med. St.’ and ‘Rasel Amoush’, and may also lack a further sampling site: see Sykes 1925, who reported C. horrida from ‘Stations 50, 56, Rasel Amoush, Adventure Bank’), then at least the presence of one specimen from the ‘Mediterranean’ and two specimens from Râs el Amoûch makes more sense. Unfortunately, we are in doubt on where to geographically place the NHMUK 1885.11.5.3910 specimen, which, according to us, is the only one that can be ascribed with certainty to C. horrida . It may have been found in station 50 or in station 56, the first in Algeria and the second between Pantelleria ( Italy) and Tunisia (see Carpenter and Jeffreys 1870), as it does not match the peculiar Adventure Bank form described by Sykes (1925). Notwithstanding the above-mentioned geographic problems, we also point out that Jeffreys’ (1885) original figure shows rounded nodules and the fourth cord on the last whorl unbeaded, as in typical C. atalaya , and that Sykes’ (1925) notes on Râs el Amoûch and Adventure Bank specimens add further doubts as to what Jeffreys (1874, 1885) and Sykes (1925) really analysed. On the other hand, it is noteworthy to point out that the overall number of whorls (14) between the NHMUK 1885.11.5.3910 specimen and that figured by Jeffreys (1885) seems to match. Taking into account previous statements, we are uncertain about the categorization of the three NHMUK specimens as syntypes of C. horrida Jeffreys, 1885 , and are also inclined to exclude Algeria from the confirmed sites for C. horrida (see above). Finally, concerning its validity as a proper species, C. horrida was sometimes considered as a mere subspecies of Cerithiopsis tubercularis (Montagu, 1803) , namely Cerithiopsis tubercularis horrida Jeffreys (e.g. Nordsieck 1968; Parenzan 1970; Tenekides 1989) (see discussions above and below for the wide differences among specimens of the C. tubercularis complex; Table 1; Figure 3 View Figure 3 (b)), or simply neglected in several catalogues and checklists of European or Mediterranean Mollusca (e.g. Sabelli et al. 1990–1992; Arduino et al. 1995; Repetto et al. 2005), and even in recent ones covering molluscan fauna of Italy ( Bodon et al. 1995; Oliverio 2010), from which waters the species was originally described.
Despite what is reported above, and apart from a few inconsistencies, in recent years European malacologists seem more or less to agree in considering C. horrida as a valid species belonging to the genus Cerithiopsis Forbes and Hanley, 1850 ( Gofas 2013) . However, the analysis of living specimens carried out here, as well as the first description of its radula and an entire protoconch, highlight several diagnostic features that are extremely different from those of the type species of Cerithiopsis ( Table 1). A comprehensive bibliographic search for the type species of ~40 genera ascribed to Cerithiopsidae H. Adams and A. Adams, 1853 revealed that C. horrida only shows striking similarities with Retilaskeya Marshall, 1978 ( Table 1; Figure 3 View Figure 3 (d); see also below), a genus based on Retilaskeya zelandica Marshall 1978 from New Zealand. Furthermore, also Marshall (1978) suggested that Cerithiopsis (Metaxia) rugulosa ([sensu] Sowerby) = Cerithiopsis (Metaxia) metaxae [sic for metaxa ] (Chiaje) as illustrated by Nordsieck (1968) could belong to Retilaskeya . Although the original drawing by Nordsieck (1968) is very poor, it shows a teleoconch with three rows of pointed nodules, while the protoconch detail, though quite inaccurate, shows granulated embryonic whorls followed by reticulated larval whorls. This pattern is quite consistent with C. horrida and rules out M. metaxa or Metaxia rugulosa (C. B. Adams, 1850) , suggesting Nordsieck (1968) misidentified the taxon.
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CAENOGASTROPODA Cox, 1960
| Romani, Luigi, Rolán, Emilio, Mifsud, Constantine & Crocetta, Fabio 2017 |
Cerithiopsis horrida ( Jeffreys, 1885 )
| Cossignani T & Ardovini R 2011: 16 |
| Bartolini S 2006: 20 |
| Mifsud C 2003: 41 |
| Cachia C & Mifsud C & Sammut P 2001: 186 |
Cerithiopsis horrida
| Giannuzzi-Savelli R & Pusateri F & Palmeri A & Ebreo C 1999: 14 |
Cerithiopsis sp.
| Cecalupo A & Giusti F 1989: 98 |
Cerithiopsis horrida
| Angelo G & Gargiullo S 1978: 115 |
Cerithiopsis horrida
| Locard A 1898: 381 |
| Jeffreys JG 1885: 60 |
Cerithiopsis horrida
| di Monterosato TA 1890: 163 |
| di Monterosato TA 1878: 156 |
Cerithiopsis horrida
| di Monterosato TA 1874: 273 |
Cerithiopsis horrida
| di Monterosato TA 1874: 360 |
Cerithiopsis horrida
| Jeffreys JG 1874: 114 |