THE VELIGER The Veliger 50(2):149-162 (June 20, 2008) © CMS, Inc., 2007 & Revision of the Protobranch Species Described by Dautzenberg Fischer (1897) with Description of a New Species and Taxonomic Comments on Bathyspinula (Bivalvia, Nuculanoidea) RAFAEL LA PERNA Dipartimento di Geologia e Geofisica, Universita di Bari, Via Orabona 4, 1-70125 Ban, Italy (e-mail: [email protected]) Abstract. In 1897, Philippe Dautzenberg and Henry Fischer described six deep water protobranchs from the North Atlantic {Princesse-Alice expeditions, 1894, 1896): Leda excisa (Philippi) var. subexcisa, Leda bernardi, Leda allaudi, Leda mirmidina, Leda mabillei and Malletia perrieri. Almost all of these taxa are only known from the original description, with no further records in the modern literature. The present revision, based on the original material, led to the following combinations: Bathyspinula subexcisa, Ledella bernardi, Yoldiella allaudi (lectotype designated), Microgloma mirmidina, Nuculana mabillei and Tindariaperrieri. A new species is described as Yoldiella dautzenbergifrom material misidentified as Leda allaudi. Taxonomic comments are given for the genus Bathyspinula Filatova, 1958. Thesubfamily Bathyspinulinae Coan & Scott, 1997 (= SpinulinaeAllen& Sanders, 1982nom. inval), formerly in the family Nuculanidae, is raised to full family rank within the Nuculanoidea. Tindariopsis Verrill & Bush, 1897 is also assigned to the Bathyspinulidae. INTRODUCTION with the original station number in parenthesis. Dautzenberg & Fischer (1897) described six new deep aDnaudtzielnlbusetrrgat(io1n9s27)asalstohroespeorotreidgitnhaellsyampeudbelsicsrhiepdtiobnys water protobranchs from the North Atlantic (Prin- Dautzenberg & Fischer (1897). cesse-Alice expeditions 1894, 1896): Leda excisa (Phi- lippi) var. subexcisa, Leda bernardi, Leda mabillei, Leda The type material is from the Musee Oceanographi- que de Monaco, the Institut Royal des Sciences allaudi, Leda mirmidina and Malletia perrieri. Almost Naturelles de Belgique, Bruxelles and the Monterosato adlelscorifpttihoens,e wtiatxha naroe founrltyherknroewconrdfsroimn tthhee moroidgeinranl collection, Museo Civico di Zoologia, Rome. A list of literature on the North Atlantic molluscs. A single this material is reported in Table 1. species, Spinula subexcisa (Dautzenberg & Fischer, woTrhkefnoulcluolwsanitdheclsascshiefimceatiboyn aOdcokpetlemdainnnth&e pWraesreennt 1897), was included in two taxonomic works (Clarke, 1961; Allen & Sanders, 1982), but without examination (1998),exceptforthepositionofthegenusBathyspinula. of the type material. The present work offers a The following abbreviations are used: exp(s). - systematic revision of these poorly known species, expeditions), sh(s) - complete shell(s), paired valves; v(s) - valve(s); IRScN - Institut Royal des Sciences based on the original material. This paper also gives an MOM Naturelles de Belgique, Bruxelles; - Musee occasion to discuss the taxonomy of the genus Bath- Oceanographique de Monaco, MCZ - Museum of yspinula and its systematic position within the Nucu- Comparative Zoology, Harvard University, Cam- lanoidea. bridge, MZR - Museo Civico di Zoologia, Rome. MATERIAL AND METHODS SYSTEMATICS The Princesse-Alice stations from which Dautzenberg Family Nuculanidae H. & A. Adams, 1858 & Fischer (1897) described the species dealt with in the present work are reported in Figure 1. Dautzenberg Genus Nuculana Link, 1807 (1927) renumbered all the stations from the Princesse- Nuculana mabillei (Dautzenberg & Fischer, 1897) Alice, Hirondelle and Prince de Monaco expeditions (1886-1913) as a single series, with longitudes west of (Figures 2a-e) Greenwich, whereas the original longitudes were west of Paris. In the present work, station numbers and Leda mabillei Dautzenberg & Fischer, 1897:207. pi. 6, longitudes are according to Dautzenberg's (1927) list, figs 9, 10. Page 150 The Veliger, Vol. 50, No. 2 B Corvo Flores :n703 Graciosa 698 SaoJorge 39 Faial ••^—=/15s3-49 I Terceira Pico AZORES SaoMiguel 738O SantaMaria » 37- 25 J L J I I I I I I Figure 1. Monaco expeditions, stations 503, 698, 703, 738, 1114, 1349. Leda mabillei - Dautzenberg, 1927:291, pi. 8, figs 25, minuta, from which it differs mainly by being less 26. elongate and more triangular in shape, more robust and convex. The occurrence ofN. mabilleiin a deep water station Types: Monaco exps., st. 503 {Princesse-Alice 1894, st. 101), 47°10'N, 5°47'45"W, 748-1262 m. 1 v, MOM isshaplulzozlwingw,atearss.speMcioerseoovfetrh,is tgheinsusspteycpiiecsalliys oncoctuarbliyn 21160, holotype. different from some deep water nuculanids with a Distribution: Only known from a single deep water long, slender and bent rostrum which can be assigned station offthe Bay of Biscay. to Thestyleda Iredale, 1929 (Di Geronimo & La Perna, 1997). The valve ofN. mabilleicould have underwent a Remarks: The single, poorly preserved type right valve down-slope transport from outer shelfbottoms, as also is notably robust and convex, triangular-elongate in suggested by its poor preservation status. shape, with a short, truncate, bicarinate rostrum and a wide, slightly concave postero-dorsal area. The sculp- Genus Ledella Verrill & Bush, 1897 ture consists ofcommarginal ridges, somewhat irregu- & Ledella bernardi (Dautzenberg Fischer, 1897) lar in spacing and strength, slightly coarser posteriorly (Figures 2a, d). The hinge is relatively strong, with a (Figures 3a-f) A triangular, oblique ligament pit (Figure 2e). shallow, poorly defined rostral ridge is present internally. The Leda bernardi Dautzenberg & Fischer, 1897:206, pi. 6, rostrum tip is slightly broken (Figure 2d), giving figs 5, 6. appearance ofan oblique truncation, as in the original Leda bernardi - Dautzenberg, 1927:289, pi. 8, figs 21, description {rostrum oblique truncatum). 22. Leda mabillei can be easily assigned to the genus Nuculana bernardi - Clarke, 1962:52. Nuculana. It is different in many respects from the two well known North Atlantic species ofNuculana, i. e. N. Types: Monaco exps., st. 738 (Princesse-Alice 1896, st. pernula (O.F. Miiller. 1776) and N. minuta (O.F. MOM wMieirlelerr,epo17r7t6e)d.bGyoSocdhioitltluest&raWtiaornsenof(1t9h9e2s)e.tDwuoestpoectihees h1o09l)o,ty3p7e°.4S0'aNm.e2s6t°a2ti5o'n15a"sWh.ol1o9t1y9pem,,2v1sv,,IRScN 12213185/85,. short rostrum, Nuculana mabillei is more similar to N. Distribution: Only known from a single deep water station, west of Sao Miguel, Azores. Table 1 Remarks: Leda bernardi was described from a single List ofthe type material. rightvalve(MOM)(Figures 3a-c), buttwoothervalves from the same station as the holotype are present at MOM IRScN MZR IRScN (Figures 3d-f): one of them is fairly well Leda excisa var. sube.xcisa 5 vs 2 vs preserved and younger, the other is badly preserved. Leda bernardi 1 V 2 vs The shell is ovate-elongate, not particularly convex, Leda mabillei 1 V moderately robust, shortly rostrate, with avery shallow Leda allaudi 1 V 1 V subrostral sinuation and an obscureposterior keel. The LMaeldlaetmiiarpmeirdriineari 21vVs 16 vs gurmobwothisstrsitareonagnldy ilolpdiesftihnoegdy,rawtie.delTyhespascuerdfacceombmeaarr-s R. L. Perna, 2007 Page 151 Figure 2. MNuOcuMlanamabillei(Dautzenberg& Fischer, 1897). a-e. Holotype (Dautzenberg& Fischer, 1897:pl. 6, figs 9, 10), length 10.35 mm, 21160. 9F.i4g0urmem3,. scLaeldeelblaarb=ern1armdmi,(DMauOtzMenb2e1r1g58&. dF,isec.hetro,po1t8y9p7e),. la-ecn.gtHho6l.o1t1ypmem,(DaIuRtSzceNnbe1r2g38.&f.Fitsocpheort,yp1e8,97l:epnlg.th6,8f.i2g0sm5,m6,),IlReSngctNh 1238/5. Page 152 The Veliger, Vol. 50, No. 2 ginal ridges, becoming better defined near the ventral zenberg& Fischer, 1897; Dautzenberg, 1927) cannot be margin. The hinge is relatively strong, with a triangu- confirmed. dlaerf,ineodb,linqoute plaritgiacmuelnatrlypidt.eepT.heThepalllairavlalssihneulsl iisswwoerlnl Remarks: The shell of Yoldiella allaudi is delicate, markedly convex, ovate, distinctly inequilateral in in all three valves. shape and with a sculpture of thin, irregularly spaced This species shows remarkable similarities with a group of deep-water North Atlantic species which commarginal ridges. The hinge is thin, with anterior and posterior rows ofteeth ofsimilar length, separated includes Leda oxira Dall, 1927, Leda semen Smith, s1i8m8i5l,isLAeldellelna&parHvaannVaerhr,ill19&89.BuTshhe,y18s9h8areanadnLeodveatlel-a bovyaTtahe,esmm1aa7lt0le,jriatamrliiannogfmulLaaexrdialmiauglamlmaeulndeitngatptiht..MTOhMe lcaornvsalistsshelolfias elongate shell, a short rostrum, a poorly defined single valve (st. 703), illustrated by Dautzenberg & subrostral sinuation and a strongly opisthogyrate Fischer (1897:pl. 6, figs 7, 8). Three valves from the kumnboow.n Offrotmhesteh,e LeWdeesltla Esuirmiolpiseains tBhaesionnl(yAlslpeencie&s aslalmauedis,tawthieonreaarse ptrheesetnwtoaottIhRerScvNa:lvoesneboeflotnhgemtoisaLneedwa Hannah, 1989), whereas Leda oxira, L. semen and species herein described. Ledella parva are from the Western Atlantic (Smith, Additional material labelled as Leda allaudi, from & & 1885; Verrill Bush, 1898; Dall, 1927; Allen two other stations, is present at IRScN: Monaco exps., bHyannAlalhe,n 1&989H).anAnllahof(1t9h8e9m) wwehroe raelfseorraedttteompLteeddelltao sstt.. 11314194,, 3383°°3559''3300""NN,,288°°102'54'54"5,"W8,5112m5,0omf,f CAazsoarbelsaanncda Vseyrnroilnlym&isBeusLhe,de1l8l9a8)p.aTrhviaswpirtohblLeemdaissheamrednto(sreeesoallvseo, (TFhiegumraete1r),iablutfrnoomstpheeciAmzeonrepsroivnecslutdoesbe3Lveadlvaesalloafudain. since the type material of Leda semen is destroyed unidentified species, here kept as Yoldiella sp. A, and (Allen & Hannah, 1989), but the examination of the two poorly preserved, unidentifiable shells. The mate- original illustrations (Smith, 1885:pl. 19, figs 2, 2a; rial from offCasablanca includes Yoldiella semistriata & Verrill Bush, 1898:pl. 81, fig. 1) suggests a distinct (Jeffreys, 1879) (2vs, 1 sh), YoldiellaseguenzaeBonfitto status for both species. Another deep water species, & Sabelli, 1995 (2 vs) and an unidentified species (1 v), Ledella librata Dell, 1952 from the Challenger Plateau, here kept as Yoldiella sp. B. Some of this material is New Zealand, seems notably similar to the group of illustrated in Figure 5. Atlantic species. In order to fix the identity of Leda mabillei, a The systematic position ofthese species is not clear. lectotype was designated (Figures 4a, b): it is the left They actually recall Ledella, but differ bybeing notably valve from st. 703 (MOM), illustrated by Dautzenberg elongate and with a strongly opisthogyrate umbo. The & Fischer (1897). The other valve from the same species ofLedella usually have a rather sharp posterior station at IRScN is a paralectotype (Figures 4c, d). keel, a well defined subrostral sinuation and a pointed None of the many species of Yoldiella known from rostrum (e.g., Waren, 1978; Allen & Hannah, 1989; La theAtlantic (e.g., Waren, 1989; Allenetal., 1995; Salas, Perna et al., 2004). However, no other genus so far 1996) seemsparticularly similarto Y. allaudi, except for described seems to provide a better position for this Y. subaequilatera(Jeffreys, 1879) and the followingnew group of species. species, as discussed below. Genus Yoldiella Verrill & Bush, 1897 Yoldiella dautzenbergi n. sp. & Yoldiella allaudi (Dautzenberg Fischer, 1897) (Figures 4e-h) (Figures 4a-d) Type material: Holotype and one paratype (left valves), Leda allaudi Dautzenberg & Fischer, 1897:207, pi. 6, IRScN, 1238/03. NLuecfdiuaglsaal7nl,aau8.adlil-auDdaiu-tzCelnabreker,g,1916922:75:22.90, pi. 8, figs 23, 24. T1y89p6e, lsto.ca7l4i)t,y:39M°o2n1a'2c0o"Ne,xp3s1.°,06st'.W,701336{0Prmi.ncesse-Alice Etymology: Named after Philippe Dautzenberg, Bel- gian malacologist (1849-1935). Types: Monaco exps., st. 703 {Princesse-Alice 1896, st. MOM 74), 39°21'20"N, 31°06'W, 1360 m, 1 v, 21156, Description: Shell small, thin walled, ovate, poorly lectotype; 1 v, IRScN, 1238/03, paralectotype. elongate, subequilateral, moderately convex. Umbo at mid line, small, slightly opisthogyrate, distinctly >n: Only known from a single, deep water protruding from shell outline. Posterior end well stat it of Flores, Azores. Other records (Daut- rounded, anterior end obscurely rostrate, slightly R. L. Perna, 2007 Page 153 FigureM4O. Ma-d. Yoldiellaallaudi(Dautzenberg&Fischer, 1897). a,bLectotype(Dautzenberg&Fischer, 1897:pl. 6,figs7, 8),length 5.16, 21156. c, d. Paralectotype, length 4.01 mm, IRScN 1238/03. e-h Yoldiella dautzenbergi n. sp. e, f. Holotype, length 4.02 mm, IRScN 1238/03. g, h. Paratype, length 4.08 mm, IRScN 1238/03. narrower than anterior. Ventral margin moderately Remarks: The material of Yoldiella dautzenbergi n. sp. convex, with a faint slope break at postero-ventral is from the lot of Leda allaudi at IRScN (see under transition. Subrostral sinuation almost absent. Sculp- Yoldiella allaudi). ture consisting of growth striae and well incised, Yoldiella dautzenbergi is much less convex, slightly irregularly spaced commarginal lines, giving appear- less elongate and more equilateral than Y. allaudi, with ance of wide, flat ribs. Hinge plate thin, delicate. the posterior end just slightly narrower than the Dentition taxodont, with chevron-shaped teeth in two anterior one. The umbo is less opisthogyrate and series ofsimilar length, with about ten teeth anteriorly slightly smaller, the ventral margin less convex. In both and posteriorly. Anteriorrow slightlyconvex, posterior species there is a faint slope break at the postero- row almost straight. Ligament pit small, triangular, ventral transition, but it is more distinct in Y. sunken. Adductor muscle scars ovate, of similar size. dautzenbergi. The subrostral sinuation is almost absent Pallial sinus narrow, moderately deep. Prodissoconch in Y. dautzenbergi and the sculpture consists of deeply ovate, 280 um in maximum diameter. incised, irregularly spaced commarginal lines, rather Measurements: holotype 4.02 mm in length, 2.97 than ofthin ridges. Thelarval shellis smaller than in Y. mm in height, 1.05 mm in width; paratype 4.08 X 2.90 allaudi. Thelargest valve of Y. allaudiis about 5 mm in X 0.95 mm. length, whereas the two valves of Y. dautzenbergi are about 4 mm, but the material is too scant for assessing Distribution: Only known from a single, deep water a size difference between the two species. A station, east of Flores, Azores. close resemblance also exists with Yoldiella Page 154 The Veliger, Vol. 50, No. 2 F1i2g38u/r0e3.5.c, ad,.bY.olYdoiledlileallsaesgeumeinsztareiaBtoan(fJietftforey&s,Sa1b8e7l9l)i.,M1o99n5a.cMooenxapecdoiteixopnesdi(t1i9o0n1s).(s1t9.011)1,14s,t.of1f11C4a,saobflfaCnacsaa,blleanngctah,3l.e5n6gtmhm2,.5I8RmSmc,N IRScN 1238/03. e, f. Yoldiellasp. A, Monaco expeditions (1902), st. 1349, Azores, length 3.62 mm, IRScN 1238/03. g, h. Yoldiella sp. B. Monaco expeditions (1901), st. 1114, offCasablanca, length 4.86 mm, IRScN 1238/03. subaequilatera (Jeffreys, 1879), a poorly known deep Ledamirmidina-Dautzenberg, 1927:292, pi. 8, figs27- water species from the Northeast Atlantic, dealt with 30. by Waren (1989:p. 235, figs 10a, b). The new species is Nuculana mirmidina - Clarke, 1962:53. slightly less elongate and less equilateral than Y. subaequilatera, and more convex, with a narrower umbonal angle and with a better defined sculpture. Types: Monacoexps., st. 698 (Prtintcewsse-Aliceexp.M1O98M6, ssit.. o6y9)),, 1io8t4o6min,, .3i9y°1ii1'iNn,, 3ju044'40"Wvv,, 2z. vvss,, iviv. Genus Microgloma Sanders & Allen, 1973 21157, syntypes; 16 vs, MIRScN 1239/02, syntypes. Microgloma mirmidina (Dautzenberg & Fischer, Distribution: Only known from a single, deep water 1897) station, south-east ofFlores, Azores. (Figures 6a-g) Remarks: The position of the family Pristiglomidae Sanders & Allen, 1973 in the superfamily Nuculoidea, Leda mirmidina Dautzenberg & Fischer, 1897:208, pi. as proposed by Sanders & Allen (1973), was criticized 6, figs 11-14. by Ockelmann & Waren (1988) who assigned Pristi- R. L. Perna, 2007 Page 155 Figure 6. a-g. Microgloma mirmidina (Dautzenberg & Fischer, 1897). a, b. Syntype (Dautzenberg & Fischer, 1897:pl. 6, figs 12, 14), length 1.84 mm, IRScN 1239/02(ligamentpitenlargedby breakingorcorrosion), c. Syntype, length 1.47 mm, IRScN 1239/02. d, e. Syntype, lenMgtOh M1.58 mm, IRScN 1239/02. f, g. Syntype, length 1.63 mm, scale bar =0.5 mm, IRScN 1239/02. h, i. Syntype, length 1.57 mm, 21157. gloma Dall, 1900 and Microgloma Sanders & Allen, in M. mirmidina and generally well-defined in the other 1973 to the Nuculanidae. They also presented strong species. mm evidence for the progenetic character of Microgloma. The largest syntype is 1.84 in shell length Three further species ofMicrogloma were known, all (Figures 6a, b), the others 1.5-1.6 mm. Microgloma from the Atlantic (Sanders & Allen, 1973; Ockelmann mirmidina therefore is notably larger than M. yongei, &Waren, 1998): M. yongeiSanders&Allen, 1973 (type M. tumidula and M. pusilla which are about 1 mm in species), M. tumidula (Monterosato, 1880) (= M. shell length (Allen & Sanders, 1973; Ockelmann & turnerae Sanders & Allen, 1973) and M. pusilla Waren, 1998). The shell shape changes notably with (Jeffreys, 1879). The last two species occurin European growth, from dorso-posteroventrally oblique to poste- waters. AnotherEuropean species, Phaseolusguilonardi riorly elongate, whereas the other species grow almost Hoeksema, 1983 is provisionally placed in Microgloma, isometrically and equilaterally, as seen in the growth but it clearly belongs to a different group, as discussed series of M. yongei and M. tumidula reported by by Ockelmann & Waren (1998) and La Perna (2003). Sanders & Allen (1973). This is probably due to the Microgloma mirmidina is somewhat similar to M. relatively large size of M. mirmidina, allowing this yongei and M. tumidula in the ovate-subrectangular species to follow a growth pattern more similar to that shape, whereas M. pusilla is distinctly egg-shaped. All ofnormal sized bivalves, whereas the other species are these species have a comparatively robust, notably too small for manifesting marked allometric changes. convex shell, with a sculpture of thin ridges near the At a size larger than 1.3—1.5 mm, the growth of M. ventral margin. The muscle scars are slightly buttressed mirmidina produces a stepped shell edge, giving a box- Page 156 The Veliger, Vol. 50, No. 2 like appearance. As observed by Ockelmann & Waren (1984) included this genus in the subfamily Ledellinae, (1998), such a growth pattern which at a smaller extent family Ledellidae Allen & Sanders, 1982. Ockelmann & occurs in the other species ofMicrogloma, provides an Waren (1998) kept the Nuculanidae as a single, increase in shell volume and counterbalances the effects undivided family; a systematic view markedly different ofminiaturization. from the multi-taxa classification by Allen & Sanders Besides the small size, Ockelmann & Waren (1998) (1986). However, as discussed below, there are good remarked two other synapomorphies for the Micro- reasons for keeping Bathyspinulain a separate position, gloma species: the enlarged innermost teeth of the left at a full family rank. valve and the radially wrinkled surface of the The adults of Bathyspinula posses a long, mainly prodissoconch. The first character is not present in external, amphidetic ligament, with a small internal M. mirmidina (Figure 6g), but admittedly it is not component (Allen & Sanders. 1982; Di Geronimo & La always present or clearly developed in the other species Perna, 1996) (Figures 7a, b; see also the good (e.g., Ockelmann & Waren. 1998:fig. 9f)- However, the illustrations by Knudsen, 1970). The internal ligament hinge of M. mirmidina is similar to that of the tends to a semi-external position and part ofit can be congeners, with slightly chevron-shaped to rather stout seen externally, between the umbones ofclosed valves teeth and a small, elongate ligament pit. The ligament (Figure 7b). This condition is more evident in the pit is slightly oblique, with the anterior end apparently juvenile stages, which possess a proportionally larger, external or semi-external (Figure 6g). Itis similartothe clearly semi-external ligament pit (Figure 7c). The oblique ligament pit ofajuvenile specimen of Yoldiella other nuculanids, such as Nuculana, Ledella and philippiana (Nyst, 1845) illustrated by Ockelmann & Yoldiella have ajuvenile, external amphidetic ligament Waren (1998:fig. 3b), which differs by beingposteriorly becomingfully internal with growth, aswell-document- external. This supportsthe hypothesis byOckelmann & ed by Ockelmann & Waren (1998), or leaving a small Waren (1998:11) for the progenetic origin of Micro- external relict as in Jupiteria (La Perna et al., 2004). gloma from Yoldiella or Ledella. Theligament ofBathyspinulaisthenmuchmoresimilar The larval shell of M. mirmidina is ovate, about to that ofthe families MalletiidaeH. &A. Adams. 1858 180 |J.m in length, notably smaller than that of M. (Sanders & Allen, 1985), Tindariidae Verrill & Bush, yongei (290 urn) and M. tumidula (260-270 um), more 1897 (Sanders & Allen, 1977) and Neilonellidae similarto that ofM.pnsilla(195-218 um), accordingto Shileyko, 1989 (Waren, 1989; Allen & Sanders, 1996; the data by Sanders & Allen (1973) and Ockelmann & La Perna, 2007), all with a well-developed external Waren (1998). Under optical magnification the prodis- ligament and a smaller internal component in the soconch surface shows an unresolved sculpture and it adults, than to that of the other nuculanids. None of was not possible to ascertain if it corresponds to the these families provide a suitable position for Bath- radially wrinkled pattern reported by Ockelmann & yspinula, for the following reasons: 1) malletiids have a Waren (1998). subrectangular, posteriorlytruncate orbluntly rostrate, Family Bathyspinulidae Coan & Scott, 1997 ppoooorrllyy rsocsutlrpattueresdhelslhewlli;th2)noneitlroanceelloifdssuhbarovsetraanl osvualtceu,s Genus Bathyspinula Filatova, 1958 and postero-ventral sinuation; 3) tindariids have a roundish, not rostrate shell and are asiphonate (Bath- Filatova & Shileyko (1984) pointed out the preoc- yspinulahas well developed, united siphons: Filatova& cupied status ofSpinula Dall, 1908 by Spinula Herrich- Shileyko, 1984; Allen & Sanders, 1982). A full family Schaeffer. 1856 (Lepidoptera). They replaced the genus rankis therefore adopted forthe BathyspinulinaeCoan name Spinula with Bathyspinula Filatova, 1958, for- & Scott, 1997 (= Spinulinae Allen & Sanders, 1982). merly subgenus of Spinula, and erected the new The family Bathyspinulidae also provides a suitable subgenus Acutispinula. Accordingly, Bathyspinula in- position for Tindariopsis Verrill & Bush, 1897, instead & cludes the subgenera Bathyspinula (Bathyspinula) and of the Tindariidae (Verrill Bush, 1898), Malletiidae B. (Acutispinula). Species ofthe latter differ by a finer, (Dall, 1898; Vokes, 1980; Laghi, 1986) or even & almost absent sculpture and a longer, sharper rostrum Nuculanidae, subfamily Ledellinae (Allen Sanders, (Allen & Sanders, 1982; Filatova & Shileyko. 1984; 1996). The type species, Malletia (Tindaria) agathida Coan et al., 2000). The type species are Bathyspinula Dall, 1889 has the same ligament type as Bathyspinula, (B.J oceanica (Filatova, 1958) and Bathyspinula with a "well-marked dorsal ligamental furrow and a (Acutispinula) calcar (Dall, 1908), respectively. small notch or «socket» under the beak" (Verrill & Allen & Sanders (1982) erected the monogeneric Bush, 1897, 1898; see also Dall, 1898:582). Tindariopsis & subfamily Spinulinae (invalidly based on an junior agathidahas a shallowpallial sinus (Dall, 1898; Allen homonym, replaced with Bathyspinulinae by Coan & Sanders, 1996) and cannot be assigned to the Tindar- Scott, 1997) in the family Nuculanidae to contain the iidae (which lack a pallial sinus), as suspected byVerrill genus Bathyspinula, whereas Filatova & Shileyko & Bush (1898). On the other hand, the pointed, keeled R. L. Perna, 2007 Page 157 Spinula subexcisa - Clarke, 1962:52 (?). Spinula subexcisa - Allen & Sanders, 1982:21, figs 22, 23, 27, 28. Types: Monaco exps., st. 698 (Princesse-Alice 1896, st. 69), 39°11'N, 30°44'40"W, 1846 m, Azores, 5 vs, IRScN 1238/01, syntypes; Monterosato coll., 2 vs, MZR 14423, syntypes. Other material examined: Challenger exp. (1973), st. 4, 56°52'N, 10°01'W, 1993 m, Rockall Trough, 4 shs, 1 v, MCZ 348787 (Allen & Sanders, 1982). Chain 106 exp. (1972), st. 318, 50°26.8'-50c27.3'N, 13°19.9'- MCZ 13°20.9'W, 2506 m, off West Ireland, 5 shs, 348785. Distribution: Bathyspinula subexcisa is known from the North Atlantic (West Europe and Azores), in 1846- 2506 m. Remarks: The history of Bathyspinula subexcisa is closely linked toNucula excisa Philippi, 1844, described from the Plio-Pleistocene of Southern Italy (Philippi, 1844: p. 46, pi. 15, fig. 4; Di Geronimo & La Perna, 1996). According to Allen & Sanders (1982), the records of Malletia excisa by Jeffreys (1876, 1879) and of Leda excisa by Smith (1885) from the North Atlantic could have been based either on Bathyspinula subexcisa or on Bathyspinula hilleri Allen & Sanders, 1982, both occurring in the North Atlantic, the latter with a much wider Atlantic distribution. Also the Figure 7. a-c. Ligament characters of Bathyspinula. a. record ofSpinula subexcisa from the South Atlantic by CbB.haatlhBlayetsnhpygisenpruilneauxpl.asu(bh1ie9lx7lce3ri)is,a(stA.l(lD4e,anulte&znegtnShbaen5rd.eg0r1s,&mm1,F9i8s2M)c,CheZrs,t.34D188S7928737),., Cspleacrikese, p(o1s9s6i1b)lywBaasthypsrpoibnaubllayhilblaersie.dHeoncoampadirfefderehnits length 4.93 mm, MCZ 348807. c. Bathyspinula excisa (Phi- specimens with material from the Jeffreys coll. and lippi, 1844), Archi, southern Calabria, Early-Middle Pleisto- found them "identical to M. excisa, as Jeffreys cene, length 3.25 mm, author's coll. Scale bars: = 1 mm. understood it." Thisisthefirst timethetypematerial ofBathyspinula subexcisa is revised. The sole illustrations so far rostrum and the well-defined subrostral sulcus make available for this species were the drawings by Allen Tindariopsis similar to Bathyspinula and markedly & Sanders (1982). different from malletiids and neilonellids, whereas the The most obvious differences from Bathyspinula resemblance with Ledella is due to convergence. A excisa (Figures 8n-q) lie in the shallower subrostral series ofgood illustrations, though with some misiden- sinus and in the finer sculpture. Bathyspinula subexcisa tification, was published by Laghi (1986:pl. 8, figs la- also differs bybeinglessconvexandmore delicate, with 6c; Nuculana cfr. pusio Philippi of figs la,b is a a shorter rostrum and a less distinct rostral keel. Tindariopsis species), including the holotype of Tindar- Allen & Sanders (1982) reported Bathyspinula sub- iopsis agathida. excisa from a single station (Challenger exp. 1973, st. Bathyspinula (B.) subexcisa (Dautzenberg & 4a)n.dSiotmacetuoaflltyhimsamtactheersiatlhewatyspeexmaatmeirnieald.(MFiicgruorsecso8pfi-ci,) Fischer, 1897) anastomosing radiating lines, are present along the (Figures 7a, 8a-k, r, s) psruebrsoesnttrailnsuBlactuhsy;stphienyulaareesxicmiislaar(tDoithGeemriocnriomscoul&ptuLrae Ledaexcisavar. subexcisaDautzenbeg& Fischer, 1897: Perna, 1996:pl. 2, figs 1, la). This character is not 205. visible in the type material of B. subexcisa, most Leda (Neilo) excisa var. subexcisa - Dautzenberg, probably due to the poor preservation. Other speci- 1927:295. mens (Chain 106 exp. 1972, st. 318) differ by having a Page 158 The Veliger, Vol. 50, No. 2 Figure 8. a-k. Bathyspinula subexcisa (Dautzenberg & Fischer, 1897). a. b. Syntype, length 6.81 mm, IRScN 1238/01. c, d. Syntype, length 6.52 mm, IRScN 1238/01. e. Syntype, length 5.47 mm, IRScN 1238/01. f. Challenger exp. (1973), st. 4, length 43..4049 mmmm,, MMCCZZ334488778877..j.g,Chh.aiCnha1l0l6enegxep.r,esxtp..31(81,97l3e)n,gtsth.44.,42lemngmt,h M5.C01Zm3m4,87M85C.Zk.3C4h8a7i8n7.10i.6Cehxapl.,lesnt.ge3r18e,xpl.en(g1t9h735).,35stm.m4,,lMenCgtZh 348785. 1, m. Bathyspinulahilleri(Allen&Sanders, 1982). 1. St. DS23, length 3,42 mm, MCZ348807. m. St. DS23, length4.93 mm.