Leufroyia Monterosato, 1884

Subgenus Leufroyia Monterosato, 1884

Type species. Pleurotoma leufroyi Michaud, 1828 View in CoL , S.D.

Crosse (1885) (fide Pusateri et al. 2012). Recent, Mediterranean

coast of France.

Diagnosis. Raphitomids with broad, wavy axial ribs; dense and numerous, low spiral cords (three to four on adapical teleoconch whorl). Microsculpture of dense, rather conspicuous growth lines, or rugae, no granules or pustules. Protoconch with three or four whorls, the apical one wider (at c. 220 to 250 µm diameter) and lower than in the ‘multispiral’ species in the Raphitoma group, and with a weak, incipient rounded keel for a quarter of a whorl at the transition to the teleoconch. The characteristic cancellated sculpture covers 1½ of these whorls, while the apical 1½ whorls are covered by eight to nine punctuated spiral striae.

Remarks. Leufroyia was introduced by Monterosato (1884) as one of two genera, the other being Cordieria ( Monterosato, 1884) , encompassing Raphitoma sensu Bouchet & Gofas 2015 .The diagnosis was brief (“Gruppo ben distinto ad anfratti rigonfi, costati, spiralmente striati; bocca ingrossata internamente, levigata, senza denti nè solchi”) [“Distinct group with convex whorls, with costae, spirally striated; aperture inflated, internally smooth, without teeth or grooves”]. Most of these descriptive terms could be applied to many other species of Raphitoma s.l. However the group was well distinguished by the three species included, R. leufroyi , R. concinna ( Scacchi, 1836) and R. erronea ( Monterosato, 1884) . Later authors have used Leufroyia both as a genus and a subgenus, but have apparently had difficulties in specifying the morphological characters distinguishing the taxon. Defined by van Aartsen et al. (1984:91) as: “...species with noncarinate, but still diagonally cancellate protoconch whorls,...”; by Campani (1999): “Protoconch multispiral of four whorls, with diagonally cancellated sculpture on at least the two lower whorls, not carinate but regularly rounded.” [translated from Italian]; and by Cachia et al. (2001:63): “Protoconch consisting of three rounded whorls, first blunt, last two cancellated, rather oblique ribs on body whorl”. In my opinion (based on the Norwegian material), the main diagnostic morphological characters are the details of the (micro)sculpture as specified above. The protoconch is certainly different from other Raphitoma species, but the number of whorls and the presence or absence of a terminal keel might be of specific rather than generic value, as evidenced by the SEM-photo in Campani (1999) of a protoconch belonging to R. leufroyi which is very similar to my R. concinna , but with four rather than three whorls, and no visible keel.

A single Norwegian species is referable to the subgenus.

Raphitoma (Leufroyia) concinna ( Scacchi, 1836) View in CoL Figures 2F View Figure 2 , 3D and 21 - 23

Pleurotoma concinna Scacchi, 1836:12 View in CoL , Figure 18 View Figure 18 (fide Cretella et al. 2005)

Raphitoma concinna ( Scacchi, 1836) View in CoL - Rolán 1983; Sabelli et al. 1990; Öztürk et al. 2004; Høisaeter 2009; CLEMAM 2014; Gofas 2015b Defrancia Leufroyi Michaud - Jeffreys 1867 View in CoL ; Friele 1874; Norman 1879 non Pleurotoma leufroyi Michaud, 1828:121 View in CoL

Clathurella Leufroyi, Mich. - G.O. Sars 1878

Philbertia leufroyi (Michaud) - Hubendick & Warén 1976; Høisaeter 1986

Raphitoma leufroyi ( Michaud, 1828) View in CoL - Fretter & Graham 1985; Graham 1988; Cachia et al. 2001

Fusus Boothi Brown in J. Smith, 1839:98

Leufroyia Boothii, Brown - Monterosato 1884

Raphitoma boothii View in CoL (Brown in Smith, 1839) - Olsen 1994 Raphitoma (Leufroyia) boothii View in CoL (Brown in Smith, 1839) - van Aartsen et al. 1984; Smith & Heppell 1991; Heppell et al. 1997

Type material. Presumed lost ( Cretella et al. 2005)

Type locality. “ In sinu Neapolitano et Tarentino parum frequens ” ( Cretella et al. 2005).

Material examined. Sixty-two specimens from 31 stations from c. 58°N to 61°N on the coast of Norway. A single juvenile from Hjartøysundet, Bodø , Nordland county, 67°10.7’N, 14°20.3’E, 35 m, coarse shell gravel GoogleMaps .

Description. Based mainly on a specimen from Hillersholmen ( Figure 21 View Figure 21 ), 10.5 x 5.2 mm) and one from O-sundet ( Figure 23C View Figure 23 ), 11.2 x 4.6 mm. Maximum size of those measured, 12.1 mm (specimen from O-sundet, Figure 23A View Figure 23 ). Shell thick and opaque; height 2.04 to 2.45 times the diameter (for shells longer than 9.5 mm), thus extremely variable (compare Figure 21 View Figure 21 with Figure 23C View Figure 23 ). Body whorl 64-72 % (usually between 70 and 72 %) of total shell height. Shell colour variable, from dark brown to bright yellow, usually with dark spiral cords on a light-coloured background. Teleoconch whorls five (for 11-12 mm long specimens), convex with deep and distinct suture. Adapical teleoconch whorl 775-873 µm. Sculpture of numerous wide and dense axial ribs crossed by wide and low spiral cords, 12 on penultimate whorl. Distance between cords almost twice the width of the cord. Tubercles where cords cross axial ribs, low transverse swellings on ribs. Axial ribs fading out towards base, not discernible on siphonal canal. In some large shells hardly any ribs on body whorl. Siphonal canal of varying length usually short (but compare Figure 23F View Figure 23 with 26G). Shallow and wide anal sinus in outer lip near suture. Outer lip thickened but not denticulated. Microsculpture ( Figures 2F View Figure 2 and 21 View Figure 21 ) of growth lines and irregular rugae, never isolated pustules or granules. Protoconch (Figures 3D and 21) of three whorls, nine spiral rows of isolated ‘points’ on the apical 1½ whorl, the rest with diagonal diamonds as in other ‘multispiral’ species of Raphitoma . Diameter of apical whorl 225–250 µm. Apical angle 48°–54.5° (usually more than 50°). Protoconch W/L: 1.1. Protoconch colour same as teleoconch colour, usually with white apical tip ( Figure 21 View Figure 21 ). Radula illustrated in G.O. Sars (1878:Tab. VIII, Figure 3).

DISCUSSION length of the siphonal canal, are useful, but variable characters for distinguishing between species. However, all of these

Of the six species of Raphitoma described in the review, only are hard to describe in a way that is helpful for species three are relatively common in Norwegian inshore waters, identification. The microsculpture ( Figures 1 View Figure 1 and 2 View Figure 2 ) may be and only one is common in waters north of c. 64°N. This the most useful of these, but sometimes even that one is hard review does not, however, pretend to exhaust the diversity of to interprete.

Norwegian members of the genus. Several more or less well The transition zone between protoconch and teleoconch preserved shells from the Skagerrak coast (58°- 59° N) indicate is sometimes a useful morphological detail to confirm the that the list of Norwegian species is longer than the six species determination of a doubtful specimen. This is especially useful named above. Most of the shells not included belong in the R. for distinguishing between R. aequalis and R. linearis . The linearis / R. aequalis / R. obesa group, but also a shell with clear latter species has two whorls of a dark purplish-brown colour affinity to the R. purpurea group but in too bad condition to between whorls of a much lighter hue. This colour difference be properly described, points towards a larger species pool is visible even in specimens stored for a long time in ethanol

(see e.g. Figure 16F View Figure 16 ). The protoconch and transition zone of he described as new, he surprisingly did not compare with R. aequalis on the other hand has an almost uniform milk- members of the R. linearis / R. aequalis complex. However, as chocolate colour. is apparent from his detailed descriptions of shell characters as

Ideally types, or at least topotypes, should be studied well as his holotype photographs (see Figures 10 and 17 above), in order to verify the names of the species treated. No type C. aethus is most likely a synonym of R. aequalis , and C. material or other extralimital museum material has been coccyginus of R. linearis . It is highly unlikely that two species studied for this review, but the fact that the species has been not previously described should be found sympatrically with the described from British material and are easily distinguished common look-alikes in 1967. This synonymy has been accepted from sympatric congeners, is in my opinion sufficient reason in several recent check-lists (e.g. CLEMAM 2015, WoRMS for adopting the well-known and already accepted species 2015). R. linearis (or R. aequalis ) was already in the 1870-ties names: R. linearis , R. aequalis and R. purpurea for the shown to lack a radula (G.O. Sars 1878:348). G.O. Sars regarded Norwegian species. This group is sorely in need of DNA-based the lack of radula to be of generic value, and suggested that R. phylogenies, and when DNA-based analyses are produced, it aequalis (as R. linearis ) should be generically separated from R. might very well turn out that some Scandinavian species are concinna (as R. leufroyi ). More recently it has been extensively different from their British counterparts. Available evidence documented that species in the R. linearis complex lack radula indicates that most Mediterranean species of the genus are (e.g. Sheridan et al. 1973 and Fedosov 2008). Neither G.O. confined to the Mediterranean, but species of the R. linearis / Sars (1878) nor Fedosov (2008) suggested an alternative genus aequalis group as well as the R. leufroyi -groups are exceptions name for this subgroup of Raphitoma . Monterosato (1884) being found along the East Atlantic coast both south of and suggested Cirillia as name for this group, but did not refer to north of the Mediterranean. The acceptance of R. concinna as the lack of radula as a diagnostic character. However, Cirillia the name for Norwegian member of the R. leufroyi -group is is preoccupied, and if the R. linearis complex is accepted partly based on this hypothesis. as a genus-group taxon, it needs another name. Lineotoma

The material studied for this review is practically all the Nordsieck (1977) has been suggested as a replacement name, material ever collected from inshore Norwegian waters. A but Cenodagreutes is older and might thus be the valid name for limited amount of material from deeper waters on the shelf this group if DNA-based phylogenetic analyses should confirm or outer coast are listed in the MOD database (Environmental that they constitute a clade of generic rank. Monitoring Database [MOD https://projects.dnvgl.com/MOD/ Default.aspx?TOOL=ArtUt]). Most of this material is sorted from grab-hauls from 100 to 165 m depth, in the North Sea or ACKNOWLEDGEMENTS on the western slope of the Norwegian Trench and thus outside the geographic area here considered part of the Norwegian I am grateful to Per Johannessen of SAM/UNIFOB and David faunal realm. Osca of Museo Nacional de Ciencias Naturales, Madrid, but

As hinted to above, strict enforcement of the priority rules especially Per Bie Wikander from Grimstad, for providing concerning the family name Raphitomidae , is not optimal. important material. Louise Lindblom, Kenneth Meland and The continued use of Daphnellidae would have led to fewer Solveig Thorkildsen of the Biodiversity Group of the Department taxonomic problems, as Daphnella is based on a well-known of Biology, University of Bergen, for trying to tease out some Recent species. DNA secrets of part of the material for an earlier version of the

Ideally, genera should be based on a phylogenetic analysis. ms. Jon Anders Kongsrud at the University Museum of Bergen The genera in common use today are mostly from a pre- Natural History Collections is thanked for loan of museum phylogenetic era, and need confirmation from DNA-based material. Jean-Paul Kreps from Royal Belgian Institute for analyses. As it is rather impractical to discard all names not Natural Sciences donated specimens of R. purpurea from yet based on a sound phylogenetic analysis, I find it acceptable Bretagne. The late Christoffer Schander kindly read an early to use Leufroyia as the name of a subgenus, as this is well version and offered helpful comments. Last but not least thanks supported by shell morphology, and has been adopted by to Anders Warén for helping with literature and good advice. most Mediterranean authors of check-lists. Most likely the Riccardo Giannuzzi-Savelli and an anonymous referee provided heterogeneous subgenus Raphitoma will be split into several much needed resistance, and forced me to thoroughly reanalyze genus group taxa in the future, one for the R. linearis -group, my conclusions. Giannuzzi-Savelli has given much helpful one for the R. purpurea -group and one for the R. echinata - advice in addition to his review of the ms. group. Both the micro- and the macro-sculpture are sufficiently distinct in each of these groups to support such a splitting. E.H. Smith (1967a) argues that the lack of radula (and other REFERENCES peculiarities in the foregut anatomy) in at least two species from the Northeast Atlantic, justifies the erection of a separate Aartsen JJ van, Menkhorst HP, Gittenberger E. 1984. The marine genus, Cenodagreutes . The two species, from western Scotland, Mollusca of the Bay of Algeciras Spain with general notes on

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Editorial responsibility: Torkild Bakken.

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