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Ammonicera in Florida: Notes on the Smallest Living Gastropod in tlie United States and Comments on Other Species of ia)

Rüdiger B iele r Paula M . M ikke Is en Center for Evolutionary, and ÍV pa rt in ent ut Invertebrates Fm ironniental Biolt>g\ American Museum of Natural I list on Field Museum oí Nahn ul Histnrv Central Park West at 79th Street Roosevelt Hoad at Lake Shore Drive Neu Vork, Neu York 10024. V S.A. Chicago, Illinois R0f>05r U S A iinkkelUamnh .or«; hieleri^'trnnh i Uii

\BSTRACT ix surprising 11 rat no true omalogvrids were reported in some Caribbean studies that otherwise dealt with min­ Tlie first record ol a species ot m Florida with ute species (eg , De Jong ¿í Coomans, 1938; Rios. an additional record from Yucatan. Mexico i> presented and the gross morpholngx ot the living is described lor 1994). the first time Tins smallest living snail in the United States In addition lo their small size, omalogvrids have is identified as Ai/ifíiuíi/Uí U/ minortalis Rolan, 199:2. origi­ anatomical Features thai sei them apart from eaeno- nally described from Cuba, ("oni pari sous are made with gaslropods with which the\ were usualh grouped closeli similar species, especialK A japonica Habe 1972, a This led to earh spec ulations about their systematic possi b K i/o 11 specific form with known wide distribution iii position. Because of their unusual radii lar characters, the Pacific ( Lean Various taxonomic problems iii the genera G. <) bars ( 1ST-S ■ placed the at the time monntypic Amnic’fiíOTú and are addressed, and current composition of the famib is discussed Lists family as the onix member ol his new higher taxon ot curreutb recognized urna logy rid species in the Atlantic “Prionoglossa.” giving it eipial rank with other groups (Lean [including (lie Mediterranean Sea! are presented such as Tac il i ogl ossa and Ptenoglossa Jeffreys (1859a: km/ awf/.s Florida Levs , lower Hetenibramliia, thought these the onh surviving members of Ammoniet ra < hnalogyra, < »malogvrnidea, At lau iii < Uran, Svs- the otherwise extinct genus Euomphalus Sow vrhv. tematics LS14. F letter (194V showed in a detailed anatomical studx that Omalop/ra atomus (Philippi, IS II differs greatlv from tlie prosobrunchs with which it wax Ira 1NTRODIK T i ON ditiunalh placed. Omalogvrids have regained interest in recent vea is because of their presumed basal po­ The tarni)y Omalogyridae is a pourh known group of sition within the heterobnmchs ( llaszprunar, 1938 extremely small manne snails. Placed in their own su- Ponder, 1999, 1991). Their exact relationships remain perfainih < hualogvruidea thcv are < ■urn ■ 1111 \ c lassilied as uncertain; recent suggestions {Pandei. 199b; Pacaud members of the unresolved lower hete robra neb" gas­

\BBKEVI YTIONS USED ture. Prntoconch Figs 3—1 oí L3 whorls1 ; identical iii SEM but described as ' 3/4 whorl," bv Rolan. 1992), di­ A M N IL American Museum of Natural History. New ameter 126-135 pan, distinctively sculptured with nue York, U S A. major spiral cord at mid-whorl, reticulate sculpture pe­ BMNH Hie Natural I liston Museum, London Unit­ ripheral lo major cord and 3—1 smaller spin'll ridges cen­ ed K ingdom tral to major cord No distinction of a separate larval Field Museum ol Natural I listen, Chicago FMNH shell ( prntoconch II ). indicating the absence of a free USA. swimming lanai stage. Coiling near-planispiral. with M L P Museo de La Plata, Argentina si i glit initial hvperstrophv [compare Figs 3 and 4' Te- M N U N Museo Nacional de Ciencias Naturales. Ma­ leocoiich of about I 3 rounded whorls ( 1 1.5 whorls fide drid, Spain Rolan 1992 '. sculptured with prominent elongated axial MNHN Museo Nacional de Historia Natural Santiago, tuben les regularly spaced, equally sized, beginning im ­ C h ile mediately after prntoconch, nurnhrring 15 19 I I,‘3— 17 ZMB Zoologisches Museum, Humboldt Universität, fide Rolan 1992 ) on body whorl, fading to no axial sculp­ Berlin ( ¿crm me ture at the periphery Tubercles and spaces between M so Scanning Electron Micrograph with fine growth lines. Pcriphcn (Fig 6) uniformly rounded smooth except lor fine growth lines No dis­ RESULTS tinct spiral sculpture (but occasiuiudh with extremely iine lirus sec specimen in Fig. 1) Outer lip (Figs 1-2 F a m ih Omalogyridae (-,O Sars IS7S: 215 [as llo tn - thin, sharp, ending in a single plane perpendicular to the alogyridae] plant of coiling; aperture circular: columella without i ölten erroneoush credited to Fischer, 1385; c,g , Ab- folds or grooves. Head-loot (Figs, 7-8' translucent to hott, 1974) ru arb transparent Animal gliding rapid.lv on short loot, Ammonicera Vavssière, 1-393 with blunt ven active propodium Shell held nearly v er­ Ammonicera minori alis Rolan, 1992 tical h as the snail crawls Transparent operculum on (Figures 1-8) hmdfoot semng as a support for the coil of the shell species.— Aokes and Yokes 1934 IOS figs 7. 7a Head with two finger-shaped tentacles, each held m an (SEM ). erect arch cuning toward the midhue; eves black near ¿Ammonicera ntinortali.s Bolán, 1992; 40 42 figs 10 11 Itcleo- basent tentacles Radula and internal anatium not stud concli 13, 15 i protoconch ) mil SI A li ied. H o lo typ e : (V1NCN 15.95/6794); shell diameter 9 35 Habitat: Animals not observed in situ, but collected mín; ripe locality; north ol Cuba. Baracoa; hnlotvpe from shallow subtidal rocks covered on one or several i nun 4 m depth. surfaces with vannus polychaete worm tubes, marine nl Tvpe material studied; 3 paratypus, AM N II 226459 gae sponges, tunicatus, and numerous other attached or (rom tvpe locality; free-living mollusks Diet unknown (but sec below )

Florida material studied: 2 F lori tia Keys specimens Remarks: Although the diet ot Aw/nmueerfl Minor­ collected and observed alive, one each from station 1 k- talis has not been confirmed, it is hkelv to feed on the 045 [Indian Ka \ Fill Mile Marker 79 Monroe Countv, variety of algal species growing in its subtidal rock hab­ 24°53'25"N, S9°4(V28"\V, G ulf side, rocks in 0,5-1 m itat. based on literature records on the habitat/diet ol am ong Tludassitif Halodule seagrasses, 29 September other Omalogyridae on Codium and Zostera (Omah>- 1996] and station FK-062 [Missouri Key, Mile Marker gyro aftnntts | as Euomphalus nitidissimus |— Jeffreys, 39.5. Monroe County, 24°19'29r'N. SÚ14'21 "W. Gulf 1859a, I359rir on Ulva lO, atomus — F re t 6 j, 1948); on side ol Missouri-Ohm Kr\ bridge subtidal rocks J4 linus (A rota — Nordsiech, 1972); ou I lia and Enter- April 1997 ! Also ernpri shells I roni sta Fk-949 [ Mis­ omorftlui (O atomus Fretter Graham 1978 Gra­ souri ke\ site as above, 12 March 1996], FM N H 279919 ham 1.988): ou Funis, Laminaria, Clad<>plu>rat Coralli­ il shell) FK-057 [Missouri site as above 26 Sep­ na, Cha i Ammonicera rota —Fretter Öc Graham» 1978); tember 1996], FM NH 279911 (19 shells inch SEM ma­ on Padina i Haw aiian A, japonica— Kav 1979); piercing terial), AM NH 288137 (5 shells). All localities wore lulls algal cells and sucking out the contents and depositing marine and w ere sampled b\ the authors b\ rock wash­ egg strands ou bases nl Cladophora ÍA rota /A fischer­ ing brushing and rinsing ol rock surfaces ) iana —Flam 1943 Graham 1983 un Halimeda {A ja- fUOtica and others— S leurs 1985a, 1985c t; on Zostera Distribution: Now known I rom north and south Cha. ('ifstoscira. and epiphvtic diatoms (O atomus. \ coasts ol ( uka .■Bohni 1992 the Florida keys.[ this pa- fischeriana — Gaglini 1993). Bullock et al 1990 provid pci and the Yucatan Peninsula i\nkes N Yokes 1984 1 cd the un ist detailed account, reporting Azorean (1 ato­ Description: Shell Figs 1-2) extremely minute ill­ mus and \ fischeriana from a varieri of algae, including ana 1er 9 34-9 lii ...... 9 2 9 ! nun fiflc Rolan 1992), piai uspi rai tight h coiled, glossy, uniformlv dark brown Asceil.iuict! using the method nf l';ivlin as summarize«. j h> resembling a miuiahiie ammonite m shape and sculp­ |ahlonsk] ¿X Lutz I'KSU 430, fig 4 R Bieler and \} M Mikkolsen 1993 Page 5

Figures l-4>. Ammemom minoitfilis shells b\ scanning electron microscopy 'four different sin-Its F \1 \H 27901 1 :■ 1. Apical view: 2, I miniical view 3. Frotoconch detail ul fig 1 -4. Protncuneli 'detail oí fig 2) 5* Apertural new 6* Dorsal view from abovt- IS seen ill crawling animal'1 Scale bars Fig 1 = 1Ü0 jam Figs 2 lí at same scale ; Fig 3 = 20 pm (Fig 1 at same scale . PageC? 4 Tl 1E NAUTILUS. Vol 111, No. 1

\dditional specimens were described and illustrated as Ammonicera japonica from Haw aii bv Kav ( 1979 92, figs 32A-C ¡SEM as Omalogyra: earlier reported bv Kav ¿X Switzer, 1974:278* table b from Fanning Island I kav mentioned sculpture from 16 to IS axial ribs on the last whorl, the ribs becoming obsolete at the penpherv Spi­ ral striae were not described but faint spiral sculpture is visible in one illustrated shell (Kav, 1979 fig. 32B). Sleurs' {1985a:4-5 pi. 1 figs. 1, 6. 9 [SEM]; as Omal- ogifra) description of A japonica from Papua New Guin­ ea was verv similar. He described the protocol!eh in de­ tail with reticulated sculpture at the ah apical suit- his fig. 9 1 : spiral sculpture of the 0.3 to 0.45 mm large te- leoconch was not discussed, but shows verv faini Iv in one 7 SEM illustration (his íig 6» Fukudas illustration ni this species from the < Jgasawara (Bonin) Islands ( 1994 pi 35, fig. 697a-c; as Qmalogyta" ¡apt m ira ).^ shows no spiral sculpture. The large specimen illustrated (0.4 mm) has about 19 axial ribs The prntoconch and teleocunch sculpture of the sheiks of At rimolii cera minortalis and A japonica are extremelv similar according tu the SEMs provided bv Rolan 1992 ), kav (1979i Fuknda (1994) and Sleurs (1985a4 respec­ tively and suggest svnonvrnv ot the Caribbean and bulu- Pacific species. Faint spiral tt leoconch sculpture appears 8 to occur m some individuals of both nominal species. However, the disjunct distributional pattern makes fur­ Figures 7-S. Ammonicera minortalis, sketch ut living animal. ther studv necessary. No similar form has been de- Maximum shell diameter = 0 42 mm 7, Righi Literal view 8. Morsa] view scribed or recorded trom the eastern Pacific (Shaskv.

As also notus I bv Holán ( 19924 Ammcmiccra minortalis Enteromorpha, (i/sfoseira, Vita. Pterocladia, Pci/stm- is similar to the European A. rota (Forbes & Han lev. m lia, Halopteris, Asparagopsis, as w e ll as Codium Om­ 1859 ' in its teleocunch characters : blit the latter has a alogyra atomus, which the) also found ou Gelidium and gi catei number ul whorls and axial tubercles con (inning Sargassumi was the dominant species on Chondria and over the penpherv I. Ammonicera rota has however a the only molhisk found on Fucus iii that studv ■d" S ven different protoconch without reticulated sculpture (see, eg Rodriguez Babin Thiriot-Qiiiévreux, 1971 DISCUSSION pi 2, F - II; as A fischeriana) Also similar is A plicata Sleurs. 19S5, I mm the Maldives { 1985b:20 11.* figs 2, 7. Species-level identification: Holán (1992) described 10, 13. 14 4 w hich has a larger teleoeonch (0.45 to 0.65 tins species based on empty shells collected horn north mm ) with weaker axil û ni >s and a protocouch lacking the and south coasts ol Cuba (3-29 in). A comparison with reticulate sculpture present in A. mi notialis and A ja­ flu excellent original illustrations ami with paratypie ma­ ponica. terial at ÁM NH proved the identilv of I he Florida Kevs specimens. No other known Atlantic species combines Genus-level identification: Omalogyridae□ y euro nth - such axial teleocunch sculpture with reticulated sculp- comprises three recognized extant genera Ammonia*m ture of its protocol k L, Rolan apparently wás unaware of Vayssitu r 1S93. Omalogyra jell revs, 1859, and Retro* an earlier record ol this form as “Omalogyra speciesA halina Chaster, 1896. The las! walli tvpc species bv bv Voke.s and Yokes i 19841 who collected it in Arrecife monoivpv: AÏ fuscata Chaster 1896) has a sinistra! te- \lacian. about 140 km north of Progreso Yucatan, in leoconch that distinguishes i! from Ammonicera and the (n iIf of Mexico. i )malogi/ni Vccordirig to Rolan i 1992: 42' oiilv Ammonitrra ja­ Handel ( 1988:9) also placed Orbitestella 11 edule, 1917. ponica Habe, 1972. described as japans smallest gas­ m this lamilv but Ponder (1990) showed that this genus tropod trom Honshu is 'superficially similar but it has belongs ui the Valvatoidea, not OtmdogVTuidea Handel ver\ constant spiral striae I labes species (1972:11.5- (198S), who viewed omalogvridsCv as si nail-bodied mem- 116 fiel'. I-I was described as 0 42-9.68 nun in di­ bers of Architectonicidae or \rehitectouienidea (pp 10 ameter dark brow n in color with ’about 16 annulations I7v attempted to introduce a new fossii genus W am- m (ht bí whort* lí.ihe 1972 116). Habe did not phitornaria but did not designate a tvpe species Han mention or illustrate the spiral sculpture noted h\ Rolan lIcI (in Pocken 1995 92' subsrqiientlv provided such a R H iel er and P M, M ikke] sen 1998 Page 5

designation (the Upper Cretaceous Pseudomalaxis stan­ lished biological information about this/these species in­ toni Sohl I960) and thus validated Xeamphito/naria of cludes description of gross anatomv (f ranc, 1948 m i- thai date I not as of 1988 as is frequently cited see [< ZN vous svstein (Itu b e r. 199.8' egg capsules (Franc, 1943 Art. 13(b)]. Amphi tornaria Koken, 1897, and Ncamphi- I summarized by Knudsen 1995]: <3 rah am, 1988), and tomaria Bandek 1993, were then plant'd in a new family, feeding (Graham, 1988), A tupi i: toi nan id ae. bv Baude 1 i 1996), thus removing the Ammonicera should not be confused with Ammoni­ genus again from the Omalogyridae. cerina — see synonymV cif Omahtgyra (below ) The placement of the present species in the genus Ammonicera, rather than Omalogyra, is here accepted REMARKS ON OMALOGYRA AND ITS TYPE because ot: ( I ) its prntoconch sculpture with strong spi­ SPECIES ral ribs and grooves (in contrast to small tubercles in Omalogyra; e.g., Bolán, 1992); (2) tlie presence of dis­ Omalogyra was introduced bv |effreys (1359b in the tinct cephalic tentacles (absent in Omalogyra); ami (3) midst ot an en gageii discussion (with (lark. 1359; ulti­ its strong teleoconch sculpture (absent or weak in Om­ mately involving the identities of Helix nitidissima A d ­ alogyra). The known radii lae <>f Omalogyra and Am ­ ams, 1890 "Skenea "nitidissima senstt Forbes and Han­ monicera (not vet studied h>r \ mino Halis) are sn chi- ley 1850, and Tnuieatella aft anus P h ilip p i, 1841 The nF terent between members of the two nominal genera that current understanding <4 Omalogyra is based on Fret­ Sleurs (1985c: 181i suggested that they might belong to ters 19 48) excellent an atom teal studv on British annuals different families However, published radula! data dil­ identified as O atomus Other published information oil le r even w ithin the1 two genera: Omalogyra rad ulae have this species includes gross anatomv and radula (G. O been described as either uniserial (Jeffreys, 13.59, 1867; Sars 1373) nervous system ilbiber, 19938 spermatozoa Thiele* 1929; Sleurs, 1985c) or with a formula of 1-1-1 ( I leah, 1993). and egg capsules (Graham, 1988; Knud- i.G. O* Sars, 1S78: Thiele, 1929; Egorova, 1991). Those sen, 1995 [However, it should be noted that the accom­ of Ammonicera hav e been described with a tonnula oí panying SEM shell photographs, Knudsen s fig, 5. seem 1-1-1 ( V avssiére, 1893 ) or 1 - 1 -9 - J - 1 (Sieurs* 1985b t c )* to be of a skeneopsid, not of O atomus]). Tlie “eggs" of and so definitiv e conclusions must await a detailed com­ O atomus as described bv Jeffreys (1867) and Lcbour parative studv. (1937) were subsequently recognized as misidentified Unfortunately, the taxonomic history of Ammonicera, glandular s t metu res (Fretter, 1948) Omalogyra, and their included species is exceedingly No t\pe material for anv of these nominal taxa could complex and confused (see discussion below.) lie located; our following discussion thus has to concen­ trate on literature review: Helix nitidissima J, Adams, 1300, was introduced with a short description and three REMARKS ON AUUOXICFJU AND ITS TYPE illustrations (here reproduced in Fig 9b The species was SPECIES accepted and cited, in various generic combinations. b\ Ammonicera was introduced bv Vavssiere (1893:16 ffr) subsequent authors (e.g.. Weinkauffi 1868:266, as “Spira lo r Homalogyra fischeriana Monterosatii, 1869. Ile pro­ nitidissima Adams“ ). Mainj authors have considered H vided a lull anatomical description based on histologv nitidissima ) Adams, 1800. as svnonvmous with Trun­ and studies of the* radula. Franc i 1948:142 ff) and Sleurs catella atomus Philippi. 1341 (e.g., Fischer, 1857; Wein­ (1985a 9) questioned the identity ot Vavssiere s material, kauffi 1363 Fretter ¿X Graham* 1973 Rolan. 1983 Gra­ assuming that his work was based on misidentified 0?n- ham, 1933; Poppe ¿y Goto. 1991; Rosenberg/Malacolog, alogyra rota ” Forbes and Hanley, 1850 ("lo m pared to 1997). The original description bv Adams (1800: 4. ph Gaglini’s descriptions and illustrations (1993:933-04, 1* figs, 22-24) was based on the shell alone: H jelix] 934-03-04 ), Vav ssiere s line drawings of the shell ( 1893; testa duohus anfractibus, subtilissime transverse striata. figs 3-9) seem to represent tvpical A fischeriana in c o l­ Ohs Corneous pellucid, umlulicated; easily distin­ or pattern and relatively fine ereinilations of the periph­ guished by the uncommon hillii ancv of its glossiness ery alli tough the sketched pronounced axial ribbing is Original figure 23, said to be of natural size" i 1300 6 more representative of tlie nominal species A rota. measures nearlv 3 miii, The shell, much too large to be Monterosati) himself considered the two nominal species a European omalogvrid species was subsequently rec­ as varieties of one i< g , Monterosato 1872:38; 1875:29), ognized as “evidently the frv of Zonites rat I infula s [J. and mam recent authors (c g , Fretter ¿S: Graham, 1978; Alder, 1330]/ a land snail, bv Jeffreys 1367 71). The Backeljau et al , 1984: Knudsen, 1995) have deemed liolotvpe ol H nitidissima was not located (K Wav them svnonvmous (see also Höisaeter, 1968; van Aart sen RM N II T per s comm., 1997). The interpretation as a et a i, 1984’. Gaglini (.1993), on the other hand, argued young stage of a British land snail is here accepted; // convincingly for the presence of two svmpatric species. nitidissima Adams is not a senior svnonvmJ y of T atomus W h e th e r nr not tliev will prov e to be synonymous, tliev Much of the interpretation <>l "nitidissima bv sub­ arc without doubt so closelv related and morphologically sequent authors was based on Skenea" nitidissima sensu similar that it will not impact interpretation ot the nom­ Forbes and Hanley 1859, who used this name tor a dif­ inal genus Ai nun mirera (in contrast to Sleurs, 1985a) ferent species Several authors erroneously credited In addition to Vavssiere s extensive description, pub­ Forbes and Hanlev with the description of a new species Paee 6 THE NAUTILUS. Vul U l. No I

/% * 2 4 riate rai Inia). He reported its range as from the Shet­ lands to Sicilv, and probably far beyond these limits (p. I l l 1 based m part ou the synouvmv ol Tntncatclla ato­ ntas ot Philippi, and expressed lus astonishment over Philippi s m istake’’ of describing the amnia] so différ­ © ends iéc., with tapering tentacles). 9 Much of the ensuing confusion was based on (1) the treatment of Philippi’s Italian TnutcaUlla atomus " spec­ imens as members ot the British Omalogyranitidissi­ ma" sensu Forbes and Harilev, and (2) the discrepancy between gross anatomical descriptions of these two spe­ cies i.e. with or without tapering head tentacles re- s pi ctive h

/v it C la rk 1 1859:410—113 text-figure a after reexamining <4, * British annuals reconstituted from dried specimens, dis­ fry agreed with j eft revs and corroborated the correctness of Philippi's figure ol an animal with triangular tentacles, the large eyes embeddedat the center of their bases. “It appears quite char ! hat Mr. Jeffreys has delineated his 10 animal with rounded lobes, or. m other words, with the tentacles retracted (p. 4 LÍ) Jeff re vs ( lS59h:498) re­ Figure 9, Re product i ui i "I original illnslr.il ions ol Ilrln a iti- butted: “What Mr. Clark supposed to be tentacula must dtsskna j- Adams, 1800 (from Adams, 1800 pi 1. figs 22-24' have been the shrivelled lobes of the veil. Fischer Figure IO. Reproduction ot original illustrations ol Tntncatclla (1859:364-367' joined Clark in criticizing ieffrevs atomus Philippi. 1S41 from Philippi, 1841 pi 5, figs 4a-d i 1859c), assuming that the latter had described a larva1 stage with vela instead of tentacles Jeffreys then I860 198-111' m rebuttal of Fischer, affirmed that Ins ob­ 'Skenea nitidissima (c.g Jeffreys, I860; Nordxieck. servations were based on adult specimens without ten­ 1972 Nordsieck 6c Garcia-Talavera, 1979; Gagliui, tacles Finally, m British ( om.hologx Jei'frevs (1867 07- 1993) However. Forbes and Hanley themselves ( 1850 71. pi. F fig. 5. 1869:209, pi. 70, fig. 2) again described 153') cited the species as 'S [kenea] nitulissima, Adam s tile shell and annual in detail, reaffirming his opinion ol with full page and figure reference to Adams original Philippi’s error, but recognized the priority of Hintuil- work it is this misidentified "nitidissima scn.su Forbes ogym atomus (Philippi) over “Skenea nitidissima' of 1 Lm lr\ that enters into the* various longthv published Forbes and H unlex This ‘anatomically corrected Horn- discussions comparing 'nitidissima' and Tninaitclta aft*- tf/ogr/ffl atomus, w ith Skenea" nitidissima sensn Forbes mus and Ílanles m svrmrivmv, is the Omalttaura fittmms de- ■ ' ■ ir lí " "m.’’ . . = Philippi (1841 54 pi 5 figs 4a-d) described and il­ scribt i< 1 in detail b\ f ret I er 1 1948' and that currently lu s tra te d Tvun< ati lia atomus, collected iii S o rre n to forms our concept ol the genus (Campania, southern ItaKHe emphasized that lie was Unfortunately the original fig u re s of Tmneattlla ato­ able to studv tlie animal hi detail at a magnification of mus Philippi I 84 I are in eon 1 liet with the descriptions fid tunes and placed it in Tm neat ella because ol the an­ ot Fretter. Philippik illustrated gross moqthologiea] de­ imais similarity to members of that genus lu 1844, he tails iic . tapering tentacles) are indicative of Amm oni­ re-described the species (p. 134 pi 21 fig. 5; again as et i a Meanwhile, the sketched smooth shell appears m n sp 1 and reproduced his 1-341 illustrations ol T ato­ line with the current concept oi Onutlopfiu In the ab­ mus Philippi's 1341) illustrations, here reproduced in sence of ripe material, it is impossible to explain I his Fig IO. show a living specimen with planispira! shell iii sen pai k \ ll is possible that Philippi’s material con­ i with logarithmic growth ), tapering tentai les an oper­ tained members ol both genera and his illustration is a culum. and a representation of actual size of about 0.5 composite based on more than one species, mm The t\pe material 1 um not been located in Berlin In I lu* interest of nomenclatura! stability, we base or Santiago (von Rmielen ZMB pers comm. 1997 our interpretation ol Philippi's Truncatella atomus M NUN, pers, obs., 1997). on bis ilhislrutioiiAlcscription ol the shell alone (ex­ Forbes and Hanle\ (1850:153-160, pi 73 figs 7. 8: cluding the anatomv in original fig* 4c), thus pre­ described and illustrated a British shell under the name serving this name for“ Skenea' nitidissima sensu S kenrti nititlissima .Adams’ thev did not mention tile Forbes and Hanlev,1850 (non Adams, 1 8 0 0 ), and ar firing animal fliey placed Philippi’s Tntncatclla atomus Omalogyra atomussensu Jeffreys, 13 5 9 , as w ell as with question mark m svnnnvniv Jetlrevs 1 1859a 109- F re tte r, 1948. I I I pi ) figs 15a. b, 16a-c) discussed the species, as The taxonomic contusion has been compounded b\ I notnpluiius nitidissimus, with a sketch ol the animal uncertainty about tin* tvpe species designation and the

showing ciliated head lubes, ilo tentacles, and a umse- date of introduction of (hnalagtfra Some authors teg R, Biejer and P. M, M ilclcelsen, 1998 Page

Warén, 19'SO 12 cited it as having been introduced b\ Waren (1991 7-1 \ the Ivpe spec it1.s was misidentified Jeffreys i I860), with tvpe species Tnt tu'Utella atmuus Wtdi Palaz/l "1867 '.eg. Abbott 1974; Castellanos 1989a ediae sed is m the Archaeogastropoda Vaught. 1939: Rios 1994)* ( hnalogijra aloman .Philippi. 1341 Ule generic name for the reception nl these anom­ Truucuti Ila atomus P h ilip p i, 134 154 pi 5 tig 4a-d ¡excluding alous molli isles' was in tact proposed h\ ¡eflrevs : IST) b tin- sketched animal rn fig 4i | 193' In that paper he referred In name to limmt- S kitted nitidissima (Adams) sema Forbes .Liui llanlcv, 1S50, et phalus nitidissimus" (with reference lo his earlier 1859a, duct. \non Helix nitidissima ] Adams 13091 article), to E R(>ta an usage thus cannot he construed as 'deliberately subsequent citations of the name are not available tor 17? m ustal m tlx meaning ol a previous misuse 1CZN 1985 nom cnclat 1 irai purposes because thev represent manina Art I Lii)]. mula (Mouterosato. 1875:29: 1878:83 Bucquov et ul Truncatella atomus Philippi 1841 [in synonymy | 1384 324) The first available introduction appears to be Skt iira rota Forlies and ITanles 18 >6 Now considered unein that ot Daut/enbcrg arid Duroiichoux fl9l \) her ol AmrUim/rem Skt f¡ tv it 'it a ri nat ti Wehster 18,56 Described as a pnteiilia! new \mmonieerina atomus Aar pallida Mouterosato 1834 species suhscf jiieiith beginning with Jeffreys in Wi hstcr as cited bv Gaglini (1993:928—62) is likewise not avail­ 18,57' considered a varie tv/wu on vm ol S rot d a b le as of that date Monterosatoi usage (1834.22) of Jeff revs t 1867 69 tl ) sviionvmizcd Skenea nitidissima var pallula ' is a nomen malum, as is Ins Homalogyra sensit Forbes and llanlcv under Truncatella atomus, af­ atomus var zonata Mouterosato (1373 33 1. subsequenth ter recognizing the true Helix nitidissima Adams as a cited as var cv colore 2. zonata Monts. bv Bucquoy i land snail Ile also svnonvnuzed Skenea tricarinata W eb­ ul 1 1884:324; likewise a nomen nudum)* ster under Skenea rota No tvpe species was designated llotnalogym atomus var nautiliformis De Gregorio. h ftrcvs I herein changed the generic name to iinnmlo- 1889. was recognized bv Mouterosato (1890:141) as a gf ¡iii. an unjustified emendation The fitst ¿uithors to se­ juvenile of Capulus ungaricus (Linnaeus, 1753) Nev­ lect a tvpe species appear to have been Bucquoy et al ertheless, the name nautiliformis De ( hvg o rio , 1889, (IS84 78) who stated Tvpe: Htnnalogijna ahnnus P h ilip ­ was retained bv some authors to describe an ( Omalogyra pi sp. ( Truncatella ) " morph with a much widened bodv whoii (e.g.. Nord­ We lher'efore offer the following synonymies: sieck. 1972 148: Gaghni tk Gunni G illetti 1978 216. fig Onttl(oei/ra [i tin u s . PS59b 193 tvpe species hv subsequent 2c'1 Guglini (1997923 62-A) irilmduced a new irii m designation of B u cq uo v et a! I 88-1 78 . Vm neat ella ato­ subspecific name for this morph Om aleyi a atomus var mus Philippi, 1341 hiflaiao

Awuinmccrm# O C t usta, 1861 71, tvpe species bv'sub­ Homalogyra atomus var polyzona ’Brusina mss (fide séquent desigual ion ot Dali 19271» 144 .V trtmon- Mouterosato'" in Bucquov et ul 1884:624. pi 37, fig Oferina" \mmon terrina simplex <> C- Gusta 1861 32. Earlier references to a variety polyzona Bursina" bv Preoccupied hv AríimoHiccnna D 0 Costa 1856 Monterosalrj ( 1372 1875 1873- are unavailable because [Protista Phis toxon is usualh placed in sviiouvtuv thev wore stated in svnniiymv oras nomina mula Gag- ol Ammonicera (i g Palazzi & (-aplini. 1979' how­ lini ( 1995:961- 6 1 931-02-3) showed that 41 kis is a po ever its tvpe species hv subsequent designation be­ tenda! synonym of (.). simplex not O atomus longs to Ontfihmtjra. Homalogyra [eilrevs. 1867 67 on unjustified em endation

Note: In the de seri prion of their new genus Transo- ( I K BENT COMPOSITK >N ( >F OMAL( >GYRIDAE malogyia Palazzi and Gaglim I 1979 531 made Ammmi- Recognized western Atlantic Species;| regions nl iccritui simplex G Costa 1361, the tvpe species b\ tvpe localities in brackets | original designation This would make Tramonta!?e a un objective Svnonvin of Aumf< um eri mJ and a subjective Anwionic&r

Vimunmrcra rurm /aarra 17»Lui. 1992 45, figs 24, 26, 23 taia Watson \ 1386 6 7 7 ) added a Homalogyra rhtisicos- fata (?)" record from north of th e island oi i ulcina Ammo/uVcra familiaris R oi m 1992 42, 44, tigs ]6, Is. 20 between Puerto Bico and tile Virgin Islands < halleng- [Cuba] e r" si a! mu 24 715 in e M oore 11971 1 1 5 -1 16 ) doubted Ammonicera lineofuscata R ollii, 1992 14—15, figs. 22, 24—25 both the synommv o f the Ghallenger material and that 27 [Cuba] ol the shallow-water malcrial reported by Duutzenberg Kmnumicera minortalis Rolan. I'492: 10. 42. figs 10-11, 13, 15 [Cuba] (1 3 8 9 16) for the Azores, thus restricting densicostata Ktnmouiccra sculpturata K olii i 1992 40. tigs 9 12. 14 [Guha| again to I he eastern Atlant ic. The spi vies was c< »Tisii len d (hnalogtfia atoma* (Philippi 1S41 54. pi. 5. Kgs. 4a-ri [M ed­ a member uj the eastern Atlantic’ um ilogyrid tanna until iterranean] n re a d y (eg, Sabelli et atL, 1990; Gaglini 1494 . Hom- ( )intilogyrci burdwoodiana (Strehtd, 1908:52, pi e- % alogtpa densicostata was svuonymized under At ¡euom­ [Bnrriwood Bank, south of Falkland Islands] phalus ammoniformis Scguenza. 1 376 and placed as an ( hnalogyra fuscopardalis Bolín. 1992 36 38 figs 1 -7 5, 7 arc hacogasl roperiL ot uncertain affiliations tentatively [Cuba] assigned lo (Warén, 1991 74 If ). Omalogyra taludana ( 'astrllanos, 1939a S8-SVJ figs. I 2 plus sketch of apertura! aspect iii 1989b:pl I. tig. 10 [utf San Recognized eastern Atlantic/Mediterranean Spe­ [urge Culf. Argentina] cies; [regions o l tvpe localities in brackets] Omalogyra zebrina Rulan, 1992:33. Mgs 2. 4, 6 3 [Cul>a¡ Ammonicera burnayi Rol áu. 1941 112. figs. 13-14 [Cape Verde For the western Atlantic, eleven omalogvrid species Archipelago] are currently recognized. C >1 these eight are to date nnl\ Ammonicera fmheriana '.Mouterosato, 1369274—2771 pi 13 known trom (àiba Fill described b\ Holán, 1992). Two Iß I ) I Mediterranean | others Omalogyra hurdiroodiana Í Si rebel 1968) and O \umtonirrni lignea P a la //i 1688 165. (igs. 8, 13'' | M iriei ra [ \uunomcera multistriata Rolan 1991 112, 111 figs 17-16 taludana t astellano.s. 1939 art' known iroin subantarctic L? |< ape \ 'erde Archipelago] w aters of I South Mnerica. Two omalngyrid species are Ammonicera nolai Koláii 1991 114. figs 8-9 [(..’ape AVrrir Ar- now recognized from the east coast ot the United States iC* elnpelago| Ammonicera minortalis and O atomus Vmtnonirera oteroi Rolan, 1991 114. 112, figs. 14-12 |< ’ajie Several other nominal omalogvrid species have been \ erde Archipelago] reported tor tlie western Atlantic Ocean but need to he Ammonicera robusta Rolan. 1991.114—115, figs. 17-18 [Cape excluded from that fauna Verde Archipelago I \itmuuiwrra rota (ForRes N Manlrv, 1854 164, pi 73 fig It) 'Omalogyra planorbis" A nom inal species in the w est pi, 88, figs. L 2) [ïreland[ cm Atlantic frequently cited as an omalogyriri is Lipp- Ammomrcui rotundata (Palazzi 1988 145, tigs. In 21 27 ist es? planorbis Dali 1927a: 131. originally described " cu trom oti Fernandina/ Florida This deep-water species Ammonicera i ordensis Rolan, 1991 149, figs 6-7 |( ape Verde was re-described in detail by Moore 11971 114 116 fig Arch ¡pelagi)] I 1 as Omalogyra planorbis, and subsequently cu led Om­ Omalogyra atomus Philippi, 1341:54 pi 5, figs 4a-tb ¡Med­ alogyra (Ammonicera) plantains íe g Abbott, 1974:81; iterranean! Rios 1994:60). This toxon was placed iii Palazzia W aren, Otnaltigtjra disculus Palazzi 1933:164. figs 1. 26 [Madeira] 1991 as an ’ archaeogastropori group of uncertain a lfil Omalogyra simplex ((.>(7 Costa 1861:72. pi It figs 3 a. h) I Mediterranean | iations, tentatively assigned to Skeneidae (Warén. 1991 (hnalogfjra nnth.m Palazzi 1988 104 figs. 5 15 ¡Madeira! 74, 76). liefnrftu-tnut fuscula (diaster. 1896:2 [Slrail ol Gibraltar] Ammonicera fischeriana"-. N original i les* option Abbott ( 197 I 31 ) icported this spe- atomus, us members ul a moiphologivallv extrerneh sim­ • tes as ( hnalogyra 1 Xm m oniitni ) tlcîïsicostafa from deep ilar species í J antarctica Egorova 1991 wafer oil Portugal I he Azores. and labrador Moure S e v e ra l other nommai omalogvrid species have been (1971 111 showed tliaf the Labrador ( Bulldog ) ma- describi d lor the eastern Atlantic < H these, < )tnalogyra teria I in laet belongs lo I ipphtcs planmhis D ali 1927 aprila Svkes, 1925:192, I mm oi l Portugal was recog­ thus removing tlie Labrador rerun! loi O ' densiios- nized as a nieinbei ol tin* aivhaeogast ropori“ genus Eu- R Bieler an.l V M Mikkelsen. 1998 Page 9

f/üroum ( lotton, 1915 bv \\ arén i \ 991.84' Homalogyra staled on volume cover; dale confirmed in General index granulosa 3 vies 1925, also from oil Portugal was to the lYauxactions ot the Lini lean Sm-iHv of London, placet! in [lie arci iac* igastropnd" gcuus Retigyra W armt, 1876 J. 1989 (see Waren 1992.168', llotaalitgyra paradtoa Ardnini B Lactelli F < triando and G Repetto. 1495 ( .a Mouterosato (? MS ot Svkrs (1925 192) is a nomea taloer» IlhistraLo dr-lle < nnchiglir M an m del Meditcrra- neo Associazione Aimei del Museo Frederici» Eusebio nudum fwu other nominal species introduced hv Sykes Alba 173 pp [9- 23 pp. index]. (// .sitnitisa Svktvs, 1925, and II ('J| marshalli Svkes Backeljaui. T M de Mes er t, Janssens K Prnesmaiis and \\ 1925) are in need of further study (see Palazzi. 1992). Wadei 1434 Attutiomecrti rota iii Norway Gas­ An additional Me< liter ranean species. O ausonia Palazzi. tro p o d a < Omalogyridae' Fauna Norvegica .A ¡5 6 3 1938 was recently made the tvpe ol Palazzia W arén, Rande] k 1433 Repräsentieren die Fj loin ph. Jnirieu eine na­ 1991, .uii! transferred to the urchaeogustroporis," v illi türliche Einheit der CuslropodeiG Mitteilungen ms dem tentative placement iii the Skeneidae (Waren, 1991 (Geologisch Palm »ntnlngischeri Institut del l mversität Nominal species Htnimlogjfra ornata Dautzenhorg 1889 Hamburg, 67: 1-33. p 46 pi. 4, fig. 9a-d), described from the Azores, is still Bandet, k 1996 Sona Iicti rostrophie gastropods trom Iriass!» in need t »1 reinvest igai ii mi. 3t. t ätSsiiiTi Formal ron widi a disi iiSMon on the » 1 tssifit It siicMild be noted that Palazzi 1938 used ausonia lion ol the Allogastinpodj PaLnntologiscla Zeitschrift Itak and 4list ulus little disk as nouns in apposition 70(3/4): 325—365. Bielei R 1942 Gastropod phv ln ge m and systematic* Annual iii lile original descriptions recent usage as Palazzia austenae or "Omalogyra discula" (e g Sal melli et al Review e d Ecology and SvUematks 25 511—353 Bucquov, E , P. Dautzenbergi and C Dollfm 1332—1886 Les 1999; Oiannuzzi-Saveíli et al , 1994: Ardí mu et al 1995) mollusques inarms du Roussillon Tome F Gastropodes are incorrect subsequent spellings J-B Bailliete et Fils Paris 11 1 40. pis 1-5 Feb.

F3S2 , 12• H -34 pis. 6-10 (Aug 18S2> (3) 85-145. pis

acknowledgments 11—15 ( Feb 1335). (4): 13(5-196. pis 16-20 (Aug. 1883). W) 197-222, pis. 21-25 ¡ jan 1834'; (6': 223-25s. pis 2 0 - This project, as part of a broader studv of lower heter 34 Feb 1334 (7 259-243 pis 31-35 (Aug 1334 . s. o branch gastropods was supported under National Sei 299-342, pis 36—40 : Sept 1334' ■ ÙrtUCdSfi, pis. 41 -45 ence Foundation grant DEB4J31S23I to KB Field and ■;Feb 1335 . i 10 .437 418 pis 46—54 Aug. 1885); '1! la bo raton work in Florida was made possible through 119-454, pis 51-55 J a 11 13 3fi i 12' 455-436, j i f s 5 6 - Visiting Scientists Awards hv the Smithsonian Marine 60 ( Apr 1886); (lo i 437-570, pis 61 66 i O t i 18561 Station at Link Pori iSMSLP) to KB: Dr Man F Rice Bullock, R G 1969 (ïtiinlogi/ra atomus (Philippi from Mame and flie station stalt are gratehilh acknowledged lor The Nautilus, 83(2) 70 7 ! Bullock. B C 1995 Tlie distribution ol tlie niollu.scaii tanna their support Field collecting in tlie blonda krvs was associated with the intertidal coralline algal turf of a par supported through supplementär) funding Irorn the R ally submerged volcanic crater, the llheu de A ila Franca. Bertha Le Bus Charitable Trust and Field Museum's Sao Miguel, Ayon s Azoreana Suppl 1995 4-55 Marshall Field Fund Dr Kenneth J. Boss (Museum oí Castellanos. Z. | A de, 1439a Novedades sobre mieromohis- Comparative Zoology Harvard \ niversitV and Richard COS Mibaiitarticos ■: Mollusca, (Gastropoda1 Neotropica E. Petit i North Mvrtle Beach. South (Carolina) kind I v 36(92'; 89-42 [dale given as 1 de denen ihre it*' 1933' provided lilia at me, and Roberto Cipriani (FMNH as­ on first page ot article, hut published m September ot sisted v illi translations YYe thank Dr Seigio Letcher 1989 according to back msidewover of number] IMN1 IN) for hospibditv extended during RBs visit to Castellanos Z | A de 1989b Catalogo Descriptiv o de la Wa- Santiago k at lui1 Wav (BMNH) and Thomas von Bui­ [acotauna Manna Magallanica 1 Mesogastropoda Skene­ telen (ZMB) for in fon nation on tvpe holdings ol their opsidae Omalogyridae Littorinidae. Barleeidae. Eaton respective collections D r Cristian F I tu arte (MLP) for [cliidae. Skcuellidac v Rissoidae Comisión de Invesliga- a specimen loan and Richard E Petit (North Mvrtle ciones Científicas, Provincia de Buenos Aires. La Plata. 44 Beachi as veli as two anonymous reviewers tor their pp. (diaster. G W 1396 Sume new marme Mollusca from Tangu i comments on tlie manuscript The excellent facilities The Journal of Malacology, $( 1) 1—1 pi 1 and helpful stafl of the AMNH and FM NH libraries are (dark, W 1S59 On Ali Jeffreys s CG leanings iii British Con also acknowledged This is SMSLP Contribution no chohigy publislicd in the Annals ot Natural liisiorv Iot 434. I an lí arx- and August 13.53 ;tnd fm | an nan and Febmarv 1359 The Annals and Magazine ol Natural I liston TA LITERATURE CITED 4 0 6 -4 3 4 t.Gosta O G ]87>6 Atti dell'Accadernia poutailiana. Rendicon Abbott. K T 1974 American Seushclls— The M anne Molínsea to Napoli. 7 I not seen] of the Atlantic and Pacific ( oasts of North America, 2 Gusta, t ) G 1361 Microdond» niedih r run cu < m lest ri/io n e de ed Nan Nostraud Reinhold New York [viii] + 663 pp. poco ben conoscmti oil aifutti» ignoti viventi minuti e mi- 24 pis. t roscopiei del mediterráneo Naples xuii + 89 pp 13

A cia tils, j 1890 Descriptions ol some minute British shells pis. Transactions of die I imteari Society. 5 1-6 pi 1 | Article Dali W |] 1427a Smalli shells from dredgings off the south­ was read February 6, 1743 and published m 13(H), as east coast of the United States hv the 1 uited 3 |-.lNjs Irish Ikige IB THE NAUTILUS, Yol I 11. No 1

e lies Strainer Albatross iii 133.3 and ]S.S6 Proceedings 931—91 to - 9 5 . pi 981 93.3-91 to - 9 4 pi 9.38. 934-4)1 of [ f ir \ nited States National Museum 70(266f I 134 to —94, pi 934. 935—91 to —08 pi 9.85

H aii \Y IE 19271 1 Not* oni lie genera ol Costa* Mien Monde (Gaglmi A. and M Cu rin t i Gal let h, 1973 Ah une constdei a/iom The Nautilus, 40:43 1 34 sulla lam Omalogyridae Cuuchiglie, lili 12 2 0 7 -2 1 4 Dautzeiibe 1S89 Révision des im »Ñusques marins ties Giamuizzi-Savelfi R, V PusaterL A Palmeri and ( Ebreo Avon-1 S Contribution a la Lunie muhicologique lies lies 1994 Atlante del le Concii iglie Marine del Mediterrá­ Vyoivs Résultats dos Campagnes Seiri it i fiques Accom­ neo— Atlas oi the Mediterranean Sea Shells.\ol 1 Ar­ plies sur son Yachf par Albert ! Bn nee Souverain de chaeogastropoda). La Con chi glia. Rome 125 pp Monaco Fascicule I 1112 4 pis. Graham A 1938 Molluscs Prosohranch and Pvntmidellid Dautzenbcrg, P. ami P Duroucboux 1913-14 Les mollusques Gastropods Keys and Notes tui the Identification of the de la haie de Saint-Malo Feuille di s jeunes Naturalistes Specu s 2" i d Series eds Kennack, D M and B S 43 '514-suppl : 1191.4' 1-24 14 ¡517-xuppl • i 1914] 25— k Bamesi Synopsrx ot the British Fauna New Scries 2 64 pis, 1—4 [dates test* vau Aartseni-/ /;/. 1934 95| The Unm an Socictv *4 Loudon and The Estuarina and De Gregorio A 1389 Same di tainui molluschi vivent] e ter- Brachis!i-waiter Sciences Association, Bnll/Buckhuvs, Lei­ /u iri del Baeiuo Mediterráneo. Naturalista Siciliano, 8. 26 den New York, Copenhagen, t lologne vii + 662 pages, pp.. 2 pis. llabc 4 1972 The Japans smallest gastropod Ammonit era Dr long, k M and II E. Cornualis 193.3 Manne Gastropods ¡aptm tat. sp uo\ Aenus, The Japanese (oumal of Mala- Iroin Curaçao, Aruba and Bonaire. E | Bnll, Leiden cologv, 31(3)115-116. New York, ( lopoiihagen Cologne 261 pp. Ilaszpnm.il (. 1958 Ori tKë origin and evolution óf major Doekt tv; D T. lii 1993 The sh epioneman gastropods, ex- gastropod groups with special reference to the Strepto­ ( lusne ot the Stenoglossa, ol the C oitor Sand • t lampan- nema Journal ol Molluscan Studies 54- 4 . Gi67—141 i an ot northeastern Mississippi Mississippi Department Healv I \E 1993. Gnmpurativt- sperm ultrustmctim and spor- ot Environmental (t>ualit\, ( )ffii v o f i Geology. Bulletin 129 mtogi-iiesis in basal hetembranch gastropods Valvatoidea 191 pp. Architectonicoidea Rissoelloidea Omalogvrnidi a Pyr­ Egorov a. E 1941 Sulio status delia specie bipolare— Í >n the amidelloidea Mollusca1 Zoologica Scripta 226V1: 268- status ol bipolar species (hiiuloctjni atomus Philippi. 1341 11 hnalogyridae. Heterostropha) La ( 'michigha 22 H öisaetei V 1968 Skt nea nitens, Ammonicera rota r lio s to ­ mia hikisi and Eulimella nitidissima small marine gastro­ bischer, P 1857 Études sur un group* de coquilles de la tam pods new to the Norwegian fauna Sarsia 31 25-84 ¡Ut* des Trochidae (suite), lourual de Coi ici i v lu »logic, 6 Huber, C 1998 On the cerebral nervous system ol marine IBS-176 i Gastropoda i Journal of Mollnscun Stud­ Fiscb«'i'. P 1859. Notes sur le Mollusque désigne sous le nom ies, 59(4): 381 -4 2 0 de Skenea nitidissima Journal de Conehvliogie, 7- (364— IGZN lutemational Commission lui Zoologica] NomencJa-

367 r tnrei 193.5 International Code ol Zoological Noniencla Fischer, P 1835 Manuel de Conchyliologie et de Paléontologie ture 8 tai International Trust for Zoological Nomeucla- ( amcliv liologique ou Histoire Naturelle des Mollusques lure and British Museum Natural Histon 1 Uuulnu \\ \ ivauts et Fossiles, Fasen ule V III |pp 689-784]. F. Saw. + 338 pp Pans. 1 redale T 1917 More molluscan name-cliauges generic and Korbes E and S. Hanlev 1348-1353 A History (»i British specific Proceedings ot the M alaco logical Soi i eh of Lon- Mollusca and their Shells Yol 1 1 477 '1348,2 1—13(1 (1849 481-557 (1850b 3; l-é zo fXSSO), 321-R Ifí (1851 h [ablonski. D and R A Lutz 1989 Molluscan lam ii shell mor- 4 1 3(H) Í18521: Introduction: f-lxxx 1853' John van phologv Ecological and paleontological applications Bp Voorst. London ¡dates teste van Aurtsi n ei ul , I9 8 4 ] »28-877 in Rhoads D C and R A Lutz ¡eds Skeletal Kram \ 1443 Note sm (leu\ l|onialog\ ridés // Fischeriana growth i»t aquatic organisms Topics in geobiolugy Ani I. et II Atomus .Gastéropodes Prosobraiiches) et sur leur New York and London, Plenum Press développement Bulletin de la Société d Histoire Natu­ ¡eftrevs, I G 1359a Ftirtlit*r gleanings in British cnuchoiogv relle de l'Afrique du Nord 39 I 42-145. 11 h ■ Annals and Magazine ot Natura! Hisloix (3).3 106- Fretter \ 1948 The structure and life historv ol sonie minute 120, pis. 2-3. prosobrancTis ol rock pools Skeneitpsis / liant trhis F a b ri­ [ethvvx. I (G LS59li Notes on British Mollusca, in answer to cius ( htuilogtfra atomus P h ilip p i Rissoi Hit diaphana Mr William t lark s remarks mi ( Gleanings in British Con- Aider'1 m*ï opalina Jeffjew Journal ot Ehe M.l- eholiigv The Annals and Magazine of Natural Histoix nue Biological Association oi the 1 mied kingdom 27 (3)3 496-499!* 597-632, pi. 4. [|e llri\s I G I IS59e t )bsen,itinus Lutes par M J -G Jeffreys P o tte r \ and \ (Graham. 1978 Hm prosobruuch molluscs of mii I am mai ihi Skenea nitidissima hmmal de ( oiichvh- Britain and Denmark Part 4— Mamie Rissoacea The ogic 7 361-864 [partial translation of Jrfirevs ]859}i| Journal ot Mollusc,ui Studies. Snppl 6 153-241 Jeffreis. 1.3598 Additional gleanings in British com hologv 1’fie Fukudi 11 1994 M arine Gastropoda Mollusca of the ( >ga Annals and Magazine ol Natural Historv 3 1 139-299

sawara Bornii IsLnuIs Part 2 Neogastropoda. IL torn- 11 ■ 111. ■ \ s. I (. 1869 Sur le Mollusque désigné pai AtM Forbes hiantina inri lussii species with humai accounts Ogasu- el Hani e\ soils h nom Skenea niinlissinui |i nimai tie wutu Research, 29 1-126, niei 42 pis ( Gt niei IV lio lo g ir 3 11)3- 1 1 1 G agit m \ 199.1 p antilla i )malo(g\r(dae Sais (GO, 1378 \ r (el'IVevs I ( : 1367 Bntish ( rinchulogv or an account ui tlie gonautu Stippi I 1. Ernniicratio Molluscorum Mans Nos- Mollusca winch now inhabit tin British Isles .mil the sur- tri i [t +1923-91 to —04 pi 923, 929-91 fo 94 pi 929 n n m d in g seas. V olum e IA M ar im- sin lis m co n tin u a it*m IV Bielerand P M Mikkrlsrn 199S Page I 1

1 4 tin- Gastropoda as lar us the Bulla fumilv John vau Palazzi, 3 1433 Note s ngi i < Imalogvndae mediterranei e mad- Voorst. Lando]i 437 pp., frontispiece + S pis erensi Bultettmo Malacologie;», 24'5—S 101-111 ]effrovs I G. ]364 Bntisli Con* hologv m an ici omii o] the Palazzi S 1492 Note s ngi i Omalogyridae mediterranei e mad­ Mollusca w lin. lí now inhabit dic British Isles and (lie sur­ erense Aggimile Bollettmo Malaculogieo. 28' 5—12 139- rounding seas \ VítiMiif A. Manne shells and naked Mol 144 lusca lo die end ol tfie Gastropoda tile Pteropoda and Palazzi S and A Gagtim 1479 : ivononih notes on the Ris­ Cephalopoda with a supplement and other matter, con­ soidae and related families. The genus Ammonicerina <> cluding die work |nhu van\oorst London 259 pp. fron­ (4 Costa 136] Notizianu L I S M A., 1 29-37 tis piece + 8 + M12 pis Philippi. R A 1341 Z ix »logisch* Bemerkungen Fortsetzung Jeffreys | G 1334 ( in tin- Mollusca procured during the Archiv fur Naturgeschichte 7 1 42 59, pi 5 Lightning'auri ‘Porcupine Fxpedrtioux L86S-L370 Part Philippi, lí A. 1344 Enumerado Molluseonuu Siciliae cum A I! Proceedings of the Zoological Society of London. A iventnun him iu Tellure Tertiaria F o lliu m quae m Li­ 1884; 111-149, p k 9, UL nere sun Observavit, Volumen Secundum 1 Anton H il­ Kav E A 1979 Hawaiian Marine Shells Bishop M use mii le, IV + 303 pp,, pis. 13-28, Press Honolulu wu + 652 pp Prunier AA F 1490 Tlie anatomv and relationships of tin- < )r- Kav E \ and M V Switzer 1974 Molluscan distnhulion bitestelliílae Gastropoda 1 b-ten »branchia Jounial > »I Molluscan SI m lies. 56.4 515 552 patterns m Fanning Island Lagoon and a coin pan sou ol the mollusks ol ihi' lagoon and the seaward reels Pacific Ponder, AA F 1991 M a n n e valv atoidean gastropods -im pi i Science, M 3 h 275-295. cations !or eail\ heterohrauch phvlogem |oumal of \b»L luscan Studies, 57 1 21-82 Knudsen | 1995 * observations ou icproductivc strategy and Poppi-, G T .md A (Goto 1991 European Seushells Volume zoogeography ot some marine pmsobraiich gastropods I Polyplacophora Candotoveata .Snlenogastra, (Gastrop Mollusca from tlie Azores Azoreana, Suppl 1995 135- oda C h ris ta H e m m en W iesbaden 3.52 pp 15S. Bios E (M Haimnvici, [ A A Peres and R A dos Santos Le hour. M A' 1937 Tl i e eggs and lanae ot the Bn tisii pro­ collaborators 1994 Seashells of Brazil 2- ed Fundayiio sol iratichs with special reference to those living m the Gidade and Fundaçâo 1 mversidadc do Rio (Grande M u­ plankton Journal ot die Marine Biological Association ot sen Gee ano gráfico 'Prof Elic/t-r de Carvalho Ríos Rio the Fnited Kingdom, 22 105-166 Grande 863 pp.. 113 pis Le Renard J *7 al leurrent] I uitas Malacologie a GLE- Rodriguez Rabio. G and C Thiriot-(t)uiévrei.L\ 1974 Gastér­ MAM- -Check fast ol European Marine Mollusca. Inter­ opodes de la région de Roscott Etude particulière de la net ¡current TRI < hitp /Avww ninini tr/base/niala- prütuconque Cahiers de Biologie Marine, 15 531-549 co htm l>, last searched Mav 1997] pis. 1—6 M on te im a tu T A 1369, Description d’espèces nouvelles de Rolan Mosquera E 193.5 Moluscos dela Bia de Vi go I Gas la Méditerranée | on mal de GnuchvHulugie 17(8) 274— teropodos Thalassa*. 11 suppl 1 1-333 277. pL 13 Holán Y 1991 La familia Omalogyridae G <) Sars, 1373 Mohtemsato T A 1372 Nutizie mtonio alle coin.Iiighe med- Nh »IIu m u i. Gastropoda '-i i el Archipiélago de ( 'abo Verde l terrai ice Michele Amenta. Palermo 61 pp Graellsii!, 47: 105-116. Monterosati). T A 1375 Nuovu revista de Ile concii iglie med­ Roían E 1992 The familv ( Jmulogyridae G (i Sars, b373 iterraner Atti dell Act adei nia iii Scieii/r, Lettere ed Arti, í Mollusca Gastropoda m Cuba with description of eight Palermo, 5 1 5U new species Ape x. 7 2 3-5—16 Monterosati» T A 1377 Notizie sulle conchighe delia iada di Rosenberg G [current] M ALACOLOG 2 0— a Database ol Civitavecchia At mali Museo ( anco di (Genova 9 407- West*■ m Atlantic (Gastropods. Internet [current FRI 42S . last searched June 1997] Monterosati». T A 1373 Emmiera/ioue e sinonimia delle eon- Sabelth B , R. Gianmrzzi-Savelli an*I D Reduffi 1440-1992 eluglie mediterraner. Giomale Scietize Natu raii o l Econ- Catalogo Ami«»tato dei Molluschi Manni del Mediterrá­ oniiche, Palermo 13: 61-115 neo— Annotated (heck-last *4 Medit erran can Marine Mouterosato T A 1334. Conchighe httorali mediterranei Molli i sU Socictá Italiana *h Malacologia. Librería Natur [contm ] Naturalista Siciliano Paler 4 1-2 21-25 ■dística B*»logúese, Bologna 1 i-xiv, 1-348 1 1.990 2 844- t JdoberiNoy ember | 493 Í1992): 3: 501-731 ¡1992b Monterosato j A 1899 < Mnchiglie delia piofomlitá del mare Sars G O 1378. Bidrag lii Kuudskaben *ini Norges arktiske iii Palermo Naturalista Siciliano Palermo, 9 6' 140—151 lanna. 1 Mollusca région i s Arcticuc Norvegiae Oversigt Moore, D R 1471 A deep water OnWogr/rr/ in the western over de i Norges arktiske Region fnrekommendc BI oddw Atlantic T h e Nautilus, 84(4); 113-117. A AA Broggi * Christiania vvi + 466 pp.. 1 map pis 1 Nordsieck F 1972 Die europäischen M erresschneckeu Opis* 34. I-XA 111 t [lobiam'Ilia m it P\ ra undellidae Rissoacea'—Yoni Fis- Skias lev D R 1439 Th* fainilv í >iiialogVTÍd;ie in die Impical im vr his Kapverden Mittelmeer und Schw ir/es M* * r eastern Pacific The Festivus 2 b 2 17 1 fig [also pub­ Gustav Fischer Stuttgart xiii + 827 pp niei 4 pis lished as abstract with same title The Western Society o! Nordsieck F and F Gan -iu-Talavera 1979 Moluscos marinos Malacologixts Annual Report 21: 21 1989] de i 'ananas v Madera .Gastropoda Aula de ( ul tura de 3leurs. AA' 1935a The marini' inierugastropi »ds from the north­ Tenerife 208 pp., 46 pis e rn coast ol Papua New Guinea i Mollusca: Gastropoda1 Pat aud I -M and [ Le Renard 1996 Révision des mollusques 1 Fanulv < >iualogy ridac with descriptam of tun new spe­ paléogènes du basin de Pans IAr— List» systématique ae cies Bulletin di 1 Institut roval des Sciences naturelles de tuahsée Cossu i an ubui ¿i. 8 4 1995 ■ 151 -137 [published Belgique, 55 2) 31-MI 148,V ' 1-11 pi 1 ¡published 8 1 1 January 1996] March 1935] Page 12 T H E N A U T IL U S , Vol IJ L No. 1

Sleurs, W 1 MS-51) Marine rniciogastropo*Is Iront the Republic water marine Mollusca, Yucatan Peninsula, Mexico Me- *it Maldives 1 Genus Ammonicera Vavssiere, 1395 with soamericun Ecology Institute Monograph 1 Middle description of lour new species Prosol nandua Omalo­ American Research Institute Tu lane Lmversitv New Or­ gyridae Basteri;» 44' 1-3): 19—27 leans.' I'nbhcation 54 \iii + 135 pp jncl oti pix , 1933" Sleurs, W 1985c. Amtnonicern angulata sp nov i roni Laiug published A[>ril 1984 teste R E Petit after corres pun Island, Papua New (Guillea with comments ou the genus deuce with publisher ! \innioniccra Vavssiere 1393 Mollusca: (Gastropoda' An* Waren A 1939. Marine Mollusca described b\ |uhn Gwyn Hales de la Société R ovale Zoolugique de Belgique. fed revs with the location *4 the tvpe material Concho 115(2); 177-181. !« .L ii col Society oí Great Britain and Ireland Special Pub­ Stri liei H 1903 Die Castropoden unit Ausnahme der nack­ lication. I 94 pp 3 pis ten Opi s tl tob rai ich i er Wissenschaftliche Ergebnisse der Warén, A 1991 New and little known Mollusca from Iceland Schwedischen Sndpolar-Expeditiou 1991-1908 6(1 lii ani I Scandinavia. Sarsia, 76: 53-124. pp., 6 pis. Warén, A 1992 New and little known Askeneimnrpfi gastro­ Sykes E R 1925 Ou the Mollusca procured during the ‘Por­ poda trom the Mediterranean Sea and the adjacent Adaro cupine’ Expeditions 1869 70 Supplemental notes. Part Öé Ocean Bollehno M alacologico, 27i 19-12': 149-247 V Proceedings ol the M id act ) logical Society *4 London W atson H B 1386 R e p o rt oti the Scaphopoda and Gastero­ 16(4); 181-193, pi 9. poda collected bv the ff VI 8 Challenger during the vears 1373-76 Report ou the Scientific Results *4 the Vovage I hiele [ 1929 Handbuch der systematischen Wvichtierkunde o f II M S C h a lle n g e r during the years 1873-76 . , .. Zo­ Teil 1 (Loricata, Gastropoda; Prosobranchia). G Fischer. o lo g y lo-: 42) ¡ vi j + 756 p p . pis I —50. C aecidae pis 1-3. |ena. 376 pp. Webster W 1356. Ou a new British species of ,3 ktnra Annals lliorsou. G 1944 The Zoology ot East Greenland. Marine and Magazine *4 Natural History 2)13 156-157 p] 3 (Gastropoda Prosobrunehiata Meddelelser oni Grönland Webster, W 1857 < >u the supposed new British species ot 121(13): 1-181. Skenea Annals and Magazine '>1 Natural History (2'19: van Aartsen | 1 II P M G, Menkhurst and E Cittenberger 269 1934 The marine Mollusca ol the Ba\ * d Algeciras. Spain Wéinkautt, H ( 1363 Die ( onclnlien d» s Mittelmeeres, ihre vvotli genero] notes u ii Mitrella. Marginellidae and Turri­ ■■ geographische mul geologische Verbreitung Baud Í1 dae. Basteria, Stippi 2: 1—135. M ollu sca cephala T Fischer, Kassel vi + 512 pp. Yuugllt. K. C 1989 A ( classification ot the Living Mollusca R Wruz W 1983-1944 Gastropoda, Teil Í Allgemeiner Feil und T Abbott and K [ Boss eds American Mai acui* »gists, Priisobranchia In iSclmidewolf <) II led Handbuch

Melbourne Flonda. \ii + 195 pp der Palauzoologuc Berlin, Borni rage r (1 m i i. 1—249 Vkyssifere, A. 1393 Observations zoologiqties ¿i anatomiques figs 1-47] (1938 ¡2): 241—480 figs 472 1235 ¡ 19.53' >8’ sur 1 . \ntifuument. nouveau genn d*- gitstéropnde proso- 431-729 figs. 1236-2935 i 1939) A 721-969 figs 2084- branche Annales de la Faculté des 3* fences de Marseille 2737 (1940 ': ;5,) 961 1200, figs 2783 5416 19411 dv 3: 15-28, 1 pi. 1201-1396 figs.3 41 7—1211 (1943) (7) 1597-1639. i vu Yokes II E and E H Yokes. 1934 Distribution ot shallow- (1944) The following text is generated from uncorrected OCR.

[Begin Page: Page 1]

THE NAUTILUS 111(1):1-12, 1998

Page 1

Ammonicera in Florida: Notes on the Smallest Living Gastropod

in the United States and Comments on Other Species of

Omalogyridae ( Heterobranchia)

Riidiger Bieler

Center for Evolutionary and

Environmental Biology

Field Museum of Natural History

Roose\elt Road at Lake Shore DriYe

Chicago, Illinois 60605, U.S.A.

[email protected]

Paula M. Mikkelsen

Department of Invertebrates

American Museum of Natural History

Central Park West at 79th Street

New York, New York 10024, U.S.A.

[email protected] ABSTRACT

The first record of a species of Ammonicera in Florida, with an additional record from Yucatan, Mexico, is presented and the gross morphologN' of the living animal is described for the first time. This smallest living snail in the United States

is identified as Ammonicera minoiinli.s Rolan, 1992, origi­

nally described from Cuba. Comparisons are made with closely similar species, especially A. japonica Habe 1972, a

possibly conspecific form with known wide distribution in the Pacific Ocean. Various taxonomic problems in the genera

Ammonicera and Omalogyra are addressed, and current composition of the family Omalogyridae is discussed. Lists of currently recognized omalogvrid species in the Atlantic

Ocean (including the Mediterranean Sea) are presented.

Key words: Florida Keys, Gastropoda, lower Heterobranchia,

Ammonicera. Omalogyra. , Atlantic Ocean. Sys- tematics.

INTRODUCTION

The family Omalogyridae is a poorly known group of extremely small marine snails. Placed in their own su­

perfamily Omalogyroidea, they are currently classified as

members of the unresolved "lower heterobranch" gas­ tropods (e.g., Haszpnmar, 1988; Bieler, 1992; Healy,

1993). Even the most basic taxonomic and distributional information is sketchy for this group, with most faunistic studies missing or dehberatelv omitting the usuallv less- than-one-miilimeter size range of the adult shells. The few studies that have concentrated on this group have

brought many new species to our attention, such as the recent series of works with excellent scanning electron

micrographs by Sleurs of omalogyrids in Papua New

Guinea (198.3) and in the Republic' of Malcbves (1985b),

by Palazzi (1988) in the Mediterranean and Madeira, and those by Rolan in the Cape Verdes (1991) and Cuba

(1992). Whether these areas are exceptional in their high species diversity- of omalogvrids is doubtful, although it

is surprising that no true omalogyrids were reported in some Caribbean studies that otherwise dealt with min­ ute species (e.g., De Jong & Coomans, 1988; Rios,

1994).

In addition to their small size, omalogyrids have anatomical features that set them apart from caeno- gastropods with which they were usually grouped.

This led to early speculations about their systematic

position. Because of their unusual radular characters,

G. O. Sars (1878) placed the at the time monotvpic family as the only member of his new higher taxon

"Prionoglossa," giving it equal rank with other groups such as Taenioglossa and Ptenoglossa. Jeffreys (1859a) thought these animals the only surviving members of the otherwise extinct genus Euomphalus Sowerby, 1814. Fretter (194S) showed in a detailed anatomical study that Omalogt/ra atomii.s (Philippi, 1841) differs greatly from the "prosobranchs" with which it was tra­ ditionally placed. Omalogyrids have regained interest

in recent years because of their presumed basal po­ sition within the heterobranchs (Haszprunar, 1988;

Ponder, 1990, 1991). Their exact relationships remain uncertain; recent suggestions (Bandel, 1996; Pacaud

& Le Renard, 1996) to combine the Omalogyridae with several other families in a superfamily Architec­ tonicoidea are not supported by anatomical data (Hea-

ly, 1993; Huber, 1993).

At the nomenclatural level, certain confusion exists

in the literature about the usage of genus-group

names such as Ammonicera versus Ammonicerina, and about the identity- and authorship of Omalogi/ra's type species.

The discovery of an Ammonicera species in the

Florida Keys, representing the smallest gastropod

known in the United States, is here used to summarize existing data on Omalogyridae in the Atlantic Ocean and to address additional taxonomic problems.

[Begin Page: Page 2] Page

THE NAUTILUS, Vol. Ill, No, 1

ABBREVIATIONS USED

AMNH American Mu.seum of Natural History, New

York, U.S.A.

BMNH The Natural Hi.ston Museum, Ujiidon, Unit­ ed Kingdom

FMNH Field Museum of Natural Historv, Chicago,

U.S.A.

MLP Museo de La Plata, Argentina

MNCN Museo Nacional de Ciencias Naturales, Ma­ drid, Spain

MNHN Museo Nacional de Historia Natural, Santiago,

Chile

ZMB Zoologisches Museum, Humboldt Universität,

Berhn, Germany

SEM Scanning Electron Micrograph RESULTS

FamiK Omalogyridae G. O. Sars, 1S7S: 215 [as Flom- alog>Tidae]

(often erroneously credited to Fischer, 1885; e.g., Ab­

bott, 1974)

Genus Ammonicera Vayssiere, 1893

Ammonicera ininoiioUs Rolan, 1992

(Figures 1-8)

Omalogyra species. — V'okes and Vokes, 1984: 168, ligs. 7, 7a

(SEM).

Ammonicera minortalis Rolan, 1992: 40, 42, figs. 10, 11 (teleo- conch), 1.3, 15 (protoconch) (all SEM).

Elolotype: (MNCN 15.05/6794): shell diameter 0.35

nun: t\pe locahty: north of Cuba, Baracoa; holotype from 4 m depth.

Type material studied: 3 paratypes, AMNEI 226450, from Kpe localit\.

Florida material studied: 2 Florida Keys specimens collected and observed ahye, one each from station FK-

045 [Indian Key Fill, Mile Marker 79, Monroe County,

24°53'25"N. 80°40'28"W, Gulf side, rocks in 0.5-1 m among Thalassia/Flalodulc seagrasses, 20 September 1996[ and station FK-062 [Missouri Key, Mile Marker

39.5, Monroe County, 24°40'29"N, 81°14'21"W, Gulf side of Missouri-Ohio Key bridge, snbtidal rocks, 14

April 1997]. Also emptV shells from sta. FK-040 [Mis­ souri Key site as above, 12 March 1996], FMNFI 279010

(1 shell); FK-057 [Missouri Key site as above, 26 Sep­ tember 1996], FMNFI 279011 (10 .shells incl. SEM ma­ terial), AMNFI 288137 (5 shells). All localities were fully

marine and were sampled by the authors by "rock wash­

ing" (bnishing and rinsing of rock surfaces).

Distribution: Now known from north and south coasts of Cuba (Rolan, 1992), the Florida Keys (this pa­

per), and the Yucatan Peninsula (Vokes & Vokes, 1984).

Description: Shell (Figs. 1-2) extremely minute, di­ ameter 0..34-0.46 mm (0.2-0.4 mm, fide Rolan. 1992),

planispiral, tightly coiled, glossy, uiiii'orinly dark brown, resembling a miniature annnonite in shape and sculp­

ture. Protoconch (Figs. 3A) of 1.3 whorls' (identical in

SEM but described as ".3/4 whorl," by Rolan, 1992), di­ ameter 120-135 [jLm, distinctively sculptured vYith one

major spiral cord at mid-whorl, reticulate sculpture pe­ ripheral to major cord, and .3—4 smaller spiral ridges cen­ tral to major cord. No distinction of a separate larval shell ("protoconch 11 "), indicating the absence of a free swimming larval stage. Coihng near-planispiral, with slight initial hyper.strophy (compare Figs. 3 and 4). Te- leoconch of about 1.3 rounded whorls (1-1.5 whorls fide

Rolan, 1992), sculptured with prominent elongated a.xial tubercles, regularly spaced, equally sized, beginning im­

mediately after protoconch, numbering 15-19 (1.3-17 fide Rolan, 1992) on body whorl, fading to no axial sculp­ ture at the periphei'V. Tubercles and spaces between al.so with fine growth Unes. Periphery (Fig. 6) uniformly rounded, smooth except for fine growth lines. No dis­ tinct spiral sculpture (but occasionally with extremely fine hues, see specimen in Fig. 1). Outer lip (Figs. 1-2) thin, sharp, ending in a single plane perpendicular to the

plane of coiUng; aperture circular; columella without folds or grooves. Flead-foot (Figs. 7-8) translucent to

nearly transparent. Animal ghding rapidh- on short foot, with blunt, very active propochum. Shell held nearly ver­ tically as the snail crawls. Transparent operculum on

hindfoot serving as a support for the coil of the shell.

Plead with two finger-shaped tentacles, each held in an erect arch curxang toward the midline; eyes black, near

base of tentacles. Radula and internal anatomy not stud­

ied.

Flabitat: Anim;ds not observed in situ, but collected from shallow snbtidal rocks covered on one or several surfaces with various polychaete wonn tubes, marine ;d- gae, sponges, tunicates, and numerous other attached or free-living mollusks. Diet unknowm (but see below). Remarks: Although the diet ot Ammonicera minor­ talis has not been confirmed, it is hkely to feed on the variety of algal species growing in its subddal rock hab­

itat, based on literature records on the habitat/tbet of other Omalogyridae: on Codiuin and Zxstera (Omalo­ gyra atomiis [as Eiiomphahis nitidissimiis] — Jeffreys,

1859a, 1859d); on Ulva (O. atomus— Fretter, 1948); on

Fiictis (A. rota — Nordsieck. 1972); on Ulva and Enter- omoiyJia (O. atomtis — Fretter & Graham, 1978; Gra­

ham, 1988); on Fucits, Faminaria, Cladophora. Coralli­

na. Ulva [Ammonicera rota — Fretter & Gr;iliam, 1978); on Padina ( Flawaiian A. japonica — Kay, 1979); piercing algal cells and sucking out the contents, and depositing egg stranils on bases of Cladophora (A. rota I A. fischer­

iana — Franc, 1948; Graham, 1988); on Flalimeda [A. ja ­

ponica and others — Sleurs, 1985a, 1985c); on Zostera,

Ulva, Cifstoseira, and epiphytic diatoms (O. atomiis, A. fisdwriana—C,:vj}m\. 1993). Bullock tf «/., 1990, pnnid- ed the most detailed ;iccount, reporting Azorean O. ato-

iints and ,\ fischeriana from a varieh' of algae, incluihng

' Ascertained using the method of Taylor as summarized by

[alilonski & Fiitz (1980: ;5;}0, fig. 4)

[Begin Page: Page 3] R. Bieler and P. M. Mikkelseii, 199.S

Page 3

Figures 1-6. Amnionicera minortalis, shells by scanning electron nncroscopy (tonr different shells, F'MNFI 279011). 1. Apical view.

2. Umbilical \ie\v. 3. Protoconch (detail of fig. 1). 4. Protoconch (detail of fig. 2). 5. Apertural \iew. 6. Dorsal view (from "above," as seen in crawling animal). Scale bars: Fig. 1 = 100 (j.m (Figs. 2, 5, 6 at same scale); Fig. 3 = 20 (j.m (Fig. 4 at same scale).

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THE NAUTILUS, Vol. Ill, No. 1

8

Figures 7-8. Ammonicera 77iinortalis. sketch of liNing animal.

Maximum shell diameter = 0.42 mm, 7. Right lateral view. 8.

Dorsal view.

Enteromorpha, Cystoseira, Viva, Pterocladia, Petjson-

nelia, Halopteris, Asparagppsis, as well as Codium. Om­ alogyra atomus, which they also found on Gelidium and

Sarga.ssum, was the dominant species on Chondria and the only moUusk found on Funis in that study. DISCUSSION

Species-level identification: Roliin (1992) described this species based on empty shells collected from north and south coasts of Cuba (3-20 m). A comparison with the excellent original illustrations and with paratypic ma­ terial at AMNH proved the identity of the Florida Keys specimens. No other known Atlantic species combines such axial teleoconch sculpture with reticulated sculp­ ture of its protoconch. Rolan apparently was unaware of an earlier record of this iorm, as "Ottialogyra species,"

by Yokes and Yokes (1984) who collected it in Arrecife

Alacran, about 140 km north of Progreso, Yucatan, in the Gulf of Mexico.

According to Rolan (1992: 42), only Ammonicera ja-

ponka Flabe, 1972, described as "Japan's smallest gas­ tropod" from Flonshu, is "superficially similar but it has very constant spiral striae." Flabe's species (1972:11 .'>-

116, figs. 1A) was described as 0.42-0.68 mm in di­ ameter, dark brown in color, with 'about Ifi annulations

in the body whorl" (Flabe, 1972:116). Flabe did not

mention or illustrate the spiral sculpture noted by Rolan.

Additional specimens were described and illustrated as

Ammonicera japonica from Flawaii by Kay (1979:92, figs

32A-C [SEM], as Omalogyra; earlier reported bv Kay &

Switzer, 1974:278, table 1, from Fanning Island). Kay mentioned sculpture "from 16 to 18 axial ribs on the last whorl, the ribs becoming obsolete at the periphen,. " Spi­ ral striae were not described but faint spiral sculpture is visible in one illustrated shell (Kay, 1979:fig. 32B).

Sleurs' (1985a:4-5, pi. 1, figs. 1, 6, 9 [SEM]; as Omal­ ogyra) description of A. japonica from Papua New Guin­ ea was very similar. Ele described the protoconch in de­ tail "with reticulated sculpture at the abapical side" (his fig. 9); spiral sculpture of the 0.3 to 0.45 mm large te-

leoconch was not discussed, but shows very faintly in one

SEM illustration (his fig. 6). Fuk-uda's illustration of this species from the Ogasawara (Bonin) Islands (1994:pl. 35, fig. 697a-c; as "Omalogijla" japonica), shows no spiral sculpture. The large specimen illustrated (0.4 mm) has about 19 axial ribs.

The protoconch and teleoconch sculpture of the shells oi Ammonicera minortaJis and A. japonica are extremely similar according to the SEMs provided bv Rolan (1992),

Kay (1979), Fukuda (1994) and Sleurs (i985a), respec­ tively, and suggest synonymy of the Caribbean and Indo-

Pacific species. Faint spiral teleoconch sculpture appears to occur in some individuals of both nominal species.

Elowever, the di.sjunct distributional pattern makes fur­ ther study necessary. No siniOar fomi has been de­ scribed or recorded from the eastern Pacific (Shasky,

1989). As also noted by Rolan (1992), Ammonicera iniuortalis

is similar to the European A. rota (Forbes & Elanley,

1850) in its teleoconch characters (but the latter has a greater number of whorls and axial tubercles continuing over the periphery). Ammonicera rota has, however, a very different protoconch without reticulated sculpture

(see, e.g., Rodriguez Babio & Thiriot-Quie\Teux, 1974:

pi. 2, F-H; as A. fischeriana). Also similar is A. plicata

Sleurs, 1985, from the Maldives (19S5b:20 f f , figs. 2, 7,

10, 13, 14), which has a larger teleoconch (0.45A to 0.65

mm) with weaker axial ribs and a protoconch lacking the reticulate sculpture present in A. minortalis and A. ja ­

ponica.

Genus-level identification: Omalogyridae currently comprises three recognized extant genera: Ammonicera

Yayssiere, 1893, Omalogyra Jeffreys, 1859, and Retro­ tortina Chaster, 1896. The last (with type species by

monot\pv: R. ftiscata Chaster, 1896) has a sinistral te­

leoconch that distinguishes it from Ammonicera and

Omalogtpa.

Bandel (1988:9) also placed Orbitestella Iredale. 1917,

in this family, but Ponder (1990) showed that this genus

belongs in the Valvatoidea, not OmiJogv roidea. Bandel

(1988), who viewed om;ilog\Tids as small-bodied mem­

bers of Architectonicidae or Architectonicoidea (pp. 10,

17), attempted to introduce a new fossil genus "Neam-

phitomaria." but did not designate a type species. Ban- del (in Dockeiy, 1993:92) subsequently provided such a

[Begin Page: Page 5]

R. Bieler and P. M. Mikkelsen, 1998

PasAe 5

designation (the Upper Cretaceous Psctidomalaxis stan­ toni Sohl, 1960) and thus \ahdated Ncainphitomaria of that date [not as of 1988 as is frequently cited; see ICZN

Art. 13(b)]. Amphitomaria Koken, 1897, and Neamphi- tomaria Bandel, 199.3, were then placed in a new family,

Amphitoniariidae, by Bandel (1996), thus removing the genus agiiin from the Omalogyridae.

The placement of the present species in the genus

Ammonicera, rather than Omalogyra, is here accepted

because of (1) its protoconch sculpture with strong spi­ ral ribs and grooves (in contrast to small tubercles in

Omalogyra; e.g., Rolan, 1992); (2) the presence of dis­ tinct cephiilic tentacles (absent in Omalogyra); and (3)

its strong teleoconch sculpture (absent or weak in Om­ alogyra). The kniovvn radulae of Omalogyra and Am­

monicera (not yet studied for A. mhwrtalis) are so dif­ ferent between members of the two nominal genera that

Sleurs (1985c: 181) suggested that they might belong to different families. However, published radular data dif­ fer even within the two genera: Omalogyra radulae have

been described as either uniserial (Jeffreys, 1859, 1867;

Thiele, 1929; Sleurs, 1985c) or with a formula of 1 -1 -1

(G. O. Sars, 1878; Thiele, 1929; Egorova, 1991). Those of Ammonicera have been described with a fomiula of

1-1-1 (Vayssiere, 1893) or 1-1-0-1-1 (Sleurs, 1985b, c), and so definitive conclusions must await a detailed com­

parative study.

Unfortunatelv, the taxonomic history o{ Ammonicera.

Omalogyra, and their included species is exceedingly complex and confused (see discussion below).

REMARKS ON AMMOTSIICERA AND ITS TYPE

SPECIES

Ammonicera was introduced by Vayssiere (1893:16 ff.) for Homalogi/ra fischeriana Monterosato, 1869. He pro­ vided a full anatomical description based on histology and studies of the radula. Franc (1948:142 ff ) and Sleurs

(1985a:9) questioned the identity of Vavssiere's material, assuming that his work was based on misidentified "Om­ alogyra rota" Forbes and Hanlev, 1850. Compared to

Gaglini's descriptions and illustrations (1993:933-04,

934-03-04), Vayssiere s line drawings of the shell (1893: figs. 8-9) seem to represent typical A. fischeriana in col­ or pattern and relatively fine crenulations of the periph­ ery, although the sketched pronounced axial ribbing is

more representative of the nominal species A. rota.

Monterosato himself considered the two nominal species as varieties of one (e.g., Monterosato 1872:38; 1875:29), and manv recent authors (e.g., Fretter & Graham, 1978;

Backeÿau ct ah, 1984; Knudsen, 1995) have deemed them svnonymous (see also Floisaeter, 1968; van Aartsen et ai. ¡984). Gaglini (1993), on the other hand, argued convdncinglv for the presence of two sympatric species.

Whether or not thev will prove to be sviionvanous, they are without doubt so closely related and mor{)hologic;illy similar that it will not impact interpretation of the nom­

inal genus Ammonicera (in contrast to Sleurs, 1985a).

In addition to Vayssiere's extensive description, pub­

lished biological information about this/these species in­ cludes description of gross anatomy (Franc, 1948), ner­ vous system (Hulier, 1993), egg capsules (Franc, 1948

[summarized by Knudsen, 1995]; Graham, 1988), and feeding (Gniham, 1988).

Ammonicera should not be confused with Ammoni­ cerina — see svnonvmy of Omalogyra (below).

REMARKS ON OMALOGYRA AND ITS TYPE

SPECIES

Omalogyra was introduced by Jeffreys (1859b) in the

midst of an engaged discussion (with Clark, 1859) ulti- mately involving the identities of Helix nitidissima Ad­ ams, 1800, "Skenea" nitidissima .sensii Forbes and Han­

ley, 1850, and "Tntncatella" atomus Philippi, 1841. The current understanding of Omalogyra is based on Fret- ter's (1948) excellent anatomic;il study on British animals

identified as O. atonuts. Other pubhshed information on this species includes gross anatomy and radula (G. O.

Sars, 1878), nervous system (Huber, 1993), spermatozoa

(Healy, 1993), and egg capsules (Graham, 1988; Knud­ sen, 1995 [However, it should be noted that the accom­

panying SEM shell photographs, Knudsens fig. 5, seem to be of a skeneopsid, not of O. atomit.i]). The "eggs" of

0. atomtis as described by Jeffreys (1867) and Lebour

(1937) were subsequently recognized as misidentified glandular stiiictures (Fretter, 1948).

No type material for any of these nominal taxa could

be located; our following discussion thus has to concen­ trate on hterature review: Helix nitidissima ]. Adams,

1800, was introduced with a short description and three

illustrations (here reproduced in Fig. 9). The species was accepted and cited, in various generic combinations, by subsequent authors (e.g., Weinkauff, 1868:266, as "Spira

nitidissima Adams"). Many authors have considered H.

nitidissima J. Adams, 1800, as svnonymous with Tntn­ catella atoinus Philippi, 1841 (e.g., Fischer, 1857; Wein­

kauff, 1868; Fretter & Graham, 1978; Rolan, 1983; Gra­ ham, 1988; Poppe & Goto, 1991; Rosenberg/Malacolog,

1997). The original description by Adams (1800: 4, pi.

1, figs. 22-24) was based on the shell alone: H.[elLx] testa duobus anfractibus, subtihssime transverse striata.

Obs. Corneous, pellucid, umbilicated; easily distin­ guished by the uncommon brilliancy of its glossiness."

Original figure 23, said to be of "natural size" (1800:6)

measures nearlv 3 mm. The shell, much too large to be a European omalogyrid species, was subsequently rec­ ognized as "evidently the fry of Zonites radiatuliis [J.

Alder, 1830]," a land snail, by Jeffreys (1867:71). The

holotype of H. nitidissima was not located (K. Way,

BMNH, pers. comm., 1997). The interpretation as a young stage of a British land snail is here accepted; H.

nitidi.ssima Adams is not a senior svnonvm of T atomiis.

Much of the interpretation of "nitidissima " bv sub­ sequent authors was based on "Skenea" nitidissima sensu

Forbes and Hanley, 1850, who used diis name for a dif­ ferent species. Several authors erroneously credited

Forbes and Hanley with the description of a new species

[Begin Page: Page 6]

Page 6 THE NAUTILUS, Vol. Ill, No. 1

/C-A /CU

tB

JugA

10

Figure 9. Reproduction of original illustrations of Helix niti­ dissima ]. Adams, 1800 (from Adams, 1800: pi. 1, figs. 22-24).

Figure 10. Reproduction of original illustrations of Tnincatella atomiis Philippi, 1841 (from Philippi, 1841: pi. 5, figs. 4a-d).

"Skenea nitidissima " (e.g., Jeffreys, 1860; Nordsieck,

1972; Nordsieck & Garcia-Talavera, 1979; Gaglini,

1993). However, Forbes and Hanley themselves (1850:

158) cited the species as "S.lkcnca] nitidissima, Adams" with full page and figure reference to Adams' original work. It is this misidentified "nitidissima sen.sti Forbes

& Hanley " that enters into the various lengthy published chscussions comparing "nitidissima" and TntncatcHa ato­

mus.

Phihppi (1841:54, pi. 5, figs. 4a-d) described and il­

lustrated Tnincatella atomtis, collected in Sorrento

(Campania, southern lt;ilv). He emphasized that he was able to studv the animal in detail at a magnification of

60 times and placed it in Tnincatella because of the an­

imal's similarity to members ofthat genus. In 1844, he re-described the species (p. 134, pi. 24, fig. 5; again as

"n. sp.") and reproduced his 1841 illustrations of T ato-

miis. Philippi's (1841) illustrations, here reproduced in

Fig. 10, show a living specimen with planispiral shell

(with logarithmic growth), tapering tentacles, an oper­ culum, and a representation of actual size of about 0.5

mm. The t\pe material has not been located in Berlin or Santiago (von Rintelen, ZMB, pers. comm., 1997;

MNPIN, pers. obs., 1997).

Forbes and Flanley (18.50:158-160, pi. 73, figs. 7, 8) described and illustrated a British shell under the name

"S.[hettea] nitidissima, Adams "; tlic\ did not mention the

living animal. They placed Phifippis Tnincatella atomtis. with questicm mark, in .synonymy. Jeffreys ( 1859a: 109-

111, pi. 3, figs. 15a, b, 16a-c) discussed the species, as

Euomphalus nitidi.ssimtis. with a sketch of the animal

(showing ciliated head lobes, no tentacles, and a unise-

riate radula). Fie reported its range as 'from the Shet­

lands to Sicily, and probably far bevond these limits" (p.

Ill) based in part on the s)aionymy of Tnincatella ato­

mus of Pfiilippi, and expressed his astonishment over

Philippi's "mistake" of describing the animal so differ­ ently (i.e., with tapering tentacles). Much of the ensuing confusion was based on (1) the treatment of Philippi's Italian "Tnincatella atomtis" spec­

imens as members of the British "Omalogt/ra nitidissi­

ma" .sensu Forbes and Flanlev, and (2) the tiiscrepancy

between gross anatomical descriptions of these two spe­ cies, i.e., with or without tapering head tentacles, re­ spectively.

Clark (1859:410A13, text-figure), after reexamining

British animals reconstituted from dried specimens, dis­ agreed with Jeffreys and corroborated the correctness of

Philippi's figure of an animal with triangular tentacles, the large eves embedded at the center of their bases. "It appears quite clear that Mr. Jeffreys has delineated his animal with rounded lobes, or, in other words, v\ith the tentacles retracted. . ." (p. 411). Jeffreys (1859b:498) re­

butted: "What Mr. Clark supposed to be tentacula must

have been the shrivelled lobes of the veil. . . " Fischer

(18.59:364-367) joined Clark in criticizing Jeffreys

(1859c), assuming that the latter had described a larVal stage with vela instead of tentacles. Jeffreys then (1860:

108-111), in rebuttal of Fischer, affirmed that his ob­ servations were based on adult specimens without ten­ tacles. Finally, in British Concholog)', Jeffreys (1867:67-

71, pi. 1, fig.' 5; 1869:209, pi. 70, fig. 2) again described the shell and animal in detail, reaffirming his opinion of

Philippis error, but recognized the priority of "Flomal- ogt/ra atomu.s" (Philippi) over "Shenea nitidissima" of Forbes and Hanley. This "anatomically corrected " Hom­ alogyra atomtis, with "Skenea" nitidissima .sensu Forbes and Hanley in sMionvinv, is the Omalogt/ra atomtis de­ scribed in det;iil by Fretter (1948) and that currently forms our concept of the genus.

Unfortunately the original figures of Tnincatella ato­

mtis Philippi, 1841, are in conflict with the descriptions of Fretter Philippi's illustrated gross moqihological de­ tails (i.e., tapering tentacles) are indicative of Ammoni­ cera. Meanwhile, the sketched smooth sheU appears in

line with the current concept of Oinalogt/ra. In the ab­ sence of t\pe material, it is impossible to explain this discrepancy. It is possible that Philippi's material con- tiiined members of both genera and his illustration is a composite based on more than one species.

In the interest of nomenclatural stability, we base our interpretation of Philippis Tnincatella atomtis on hi.s illustration/description of the shell alone (ex­ cluding the anatomy in original fig. 4c), thus pre­ serving this name for A'Skenea" nitidissima sensu

Forbes and Hanley, 1850 (non Adams, 1800), and

Omalogyra atomus sensu Jeffreys, 1859, as well as

Fretter. 1948.

The taxonomic confusion has been compounded by uncertainty about the tvpe species designation and the date of introduction of Omalogyra. Some authors [e.g.

[Begin Page: Page 7]

R. Bieler and P. M. Mikkelsen, 1998

Page 7

Waren, 1980:12) cited it as having been introduced by

Jeffreys (1860), with t\pe species Tnincatella atoinus

Philippi, 1841, by monot>py. Others (e.g., Wenz, 1939:

647-648) gave "O. niticlissiina (Forbes & Hanley)" as tvpe species. The date of description is often erroneously cited as »1867" (e.g., Abbott, 1974; Castellanos, 1989a;

Vaught, 1989; Rios, 1994).

The generic name "for the reception ol these anom­ alous niollusks" was in fact proposed by Jeffreys (1859b:

498). In that paper, he referred bv name to "Eitoin-

phalus niti(li.ssiiini.s" (with reference to his earlier, 1859a, article), to "E. Rota" and its "varietv tricariiuita of Web­ ster." In the referenced article, he additionally stated a synonym for nitidissimus: "I have no doubt that it

is the Tnincatella atoiuti.s of Philippi" (1859a:lll). Jef­ freys did not indicate a t\pe species. Following ICZN

(1985: Art. 69(i)), there are four "originally included

nominal species": Helix nitkli.'i.'iimti J. Adams, ISOO. Now considered a land snail

[Jeffreys recognized the misidentification only in 1867; his

(1859a, b) usage thus cannot be construed as "deliberately used in the meaning of a previous misuse" (ICZN, 1985;

Art. Il(i)].

Tnincatellti atomus Philippi, 1S41 [in sviioimiiy],

Skenea rota Forbes and Hanlev, 1850. Now considered a mem­

ber of Amnumicern.

Skenea tricarinata Webster, 1856. Described as a potential new species; subsequently (beginning with Jeffreys in Webster,

1857) considered a yariety/svnonvni of S. rota-

Jeffreys (1867:69 ff ) synonvmized "Skenea nitidissima"

.icnsii Forbes and Hanley under TnincatcUa atomus, af­ ter recognizing the tnie Helix nitidissima Adams as a

land snail. He iilso svnonvmized Skenea tricarinata Web­ ster under Skenea rota. No tvpe species was designated.

Jeffreys therein changed the generic name to Homalo- giji'a, an unjustified emendation. The first authors to se­

lect a tvpe species appear to have been Buciiuov et al.

(1884:78) who stated "TyjDe: Homalogiira atomus Philip­

pi sp. (TnincatcUa)."

We therefore offer the following sviionymies:

Otnalogiira Jeffreys, 1859b:498; tvpe species by subsequent designation of Bucquoy et al. (1884:78), TnincatcUa nto-

;ni(.s' Philippi. 1841. Ammonicerina O. G. Costa, 1861: 71; tspe species by .sub­ sequent designation of Dali (1927b: 1.34, as "Ammon- ocerina"). Ammonicerina simplex O. G. Costa, 1861.

Preoccupied by Ammonicerina O. G. Costa, 1856

[Protista], This ta.xon is usually placed in synonymy of Ammonicera (e.g., Palazzi & Gaglini, 1979); how­ ever, its tvpe species by subsequent designation be­

longs to Omalogyra.

Homalogyra Jeffreys, 1867:67 dm unjustified emendation).

Note: In the description of their new genus Transo-

malogi/ra, Palazzi and Gaghni (1979:3.3) made Ammon­

icerina simplex O. G. Costa, 1861. the tvpe species by original designation. This woidd make Tran.somalogijra an objective synonym oi Ammonicerina and a subjective svnonvin of Omalogt/ra. However, as pointed out by

Waren (1991:74), the tvpe species was misidentified, with Palazzi & Ciaghni's illustration actually showing a shell of Adeuomphalus ammoniformis Seguenza, 1876.

Waren (1991) thus placed Transomalogijra in the syn­ onymy oi Adeuomphalus Seguenza, 1876, as a genus in- ceriae scdis in the "Archaeogastropoda."

Omalogt/ia atomus (Philippi, 1841)

Tnincatella atonuis Philippi, 1841:.54, pi. 5, fig. 4a-d [excluding the sketched animal in fig. 4c].

Skenea nitidissima (Adams) sensii Forbes and Flanlev. 18.50, et

anct. [non Flelix nitidi.ssimn J. Adams, 1800].

Flomalogijra atomus van vitrea Jeffreys, 1867:69.

Flomalogijra atomus \a.r. fasciata Monterosato, 1877:418.

Notes on other named "varieties":

Flomalogipa atonnis var maculata Dautzenberg and Du- rouchoiFX, 1914:27. The authorship of this name is usu­ ally crethted to Monterosato, 1875 (e.g., Gaghni, 1993:

928-02). Flowever, Monterosato's applications and some subsequent citations of the name are not available for

nomenclatural purposes because they represent nomina

nuda (Monterosati 1875:29; 1878:88; Buctjuoy et al.,

1884:324). The first available introduction appears to be that of Dautzenberg and Durouchoux (1914).

Ammonicerina atomus "var. pallida Monterosato 1884" as cited bv Gaghni (1993:928-02) is hkewise not avail­ able as ofthat date. Monterosato's usage ( 1884:22) of

"var. pallida" is a nomen nudum, as is his Flomalogi/ra atomus var. zonata Monterosato (1878:88), subsequently cited as "var, ex colore 2, zonata Monts." by Bucquoy et al. (1884:324; hkewise a nomen nudum).

Flonudogt/ra atonuis var nautilijonnis De Gregorio, 1889, was recognized bv Monterosato (1890:141) as a juvenile of Capuhis ungaricus (Linnaeus, 1758). Nev­ ertheless, the name nautilifonnis De Gregorio, 1889, was retained bv some authors to describe an Omalogi/ra

morph with a much widened body whorl (e.g., Nord­ sieck, 1972:148; Gaghni & Curini Galletti, 1978:210, fig.

2c), GagUni (1993:928-02-3) introduced a new infra­ subspecific name for this moqih, Omalogyra atomus var.

"inflata."

Homalogi/ra atomus var poh/zona "Brusina mss. (fide

Monterosato)" in Bucquoy et al., 1884:324, pi. 37, fig.

32. Earher references to a varietv' "poh/zona Brusina" by

Monterosato (1872; 1875; 1878) are unavailable because they were stated in sviiommv or as nomina nuda. Gag­

hni' (1993:931-01, 9.31-02-3) showed that this is a po­ tential sviionvm of O. simplex, not O atomus.

CURRENT COMPOSITION OF OMALOGYRIDAE

Recognized western Atlantic Species: [regions of type locahties in brackets]

Ammonicera albospeciosa Rolan, 1992:44, figs. 17, 19, 21

[Cuba] [Begin Page: Page 8]

Page 8

THE NAUTILUS, Vol. Ill, No. 1

Amnwnicera rirrnmrirm Rolan, 1992:45, figs. 23. 26, 28

[Cuba]

Amriumicera familiaris Roldn, 1992:42, 44. figs. 16. 18, 20

[Cuba]

Aintnonicera lineofusciita Rolan. 1992:44-45, figs. 22, 24— 25,

27 [Cuba]

Ammonicera minorfalis Rolan. 1992:40. 42. figs. 10-11. 1.3, 15

[Cuba]

Amnu>nicera sculpturata Rolan, 1992:40, figs. 9, 12, 14 [Cuba]

Omalogyra atomiis (Philippi. 1841:54. pi 5, figs. 4a-d) [Med­

iterranean]

Omalogyra burdwoocliann (Strebel, 190S:.52. pi. 6. fig. 85a-c

[Burdvvood Bank, south ot Falkland Islands]

Omaloayra fiiscopardahs Rolan, 1992:.36. .38, figs, 1, 3, 5, 7

[Cuba]

Onmlogtjra taludana Castellanos, 1989a:88-89, figs. 1, 2 (plus sketch of apertural aspect in 19S9b:pl. 1, fig. 10) [off San

Jorge Gulf Argentina]

Omaloaijra zebrina B-olin. 1992:38. figs. 2. 4. 6. 8 [Cuba]

For the western Atlantic, eleven omalogyrid species are currently recognized. Of these, eight are to date only

known from Cuba (all described by Rolan, 1992). Two others, Omaloaijra bunlwoodiana (Streliel, 1908) and O. taludana Castellanos, 19S9, are faiown from subantarctic waters off South America. Two omalogyrid species are

now recognized from the east coast of the United States;

Ammonicera minorialis and O. afomiis.

Several other nominal omalogyrid species have been reported for the western Atlantic Ocean, but need to be excluded from that fauna:

"Omalogyra piano rbis . A nominal species in the west­ ern Atlantic frequently cited as an om;dogyricl is Lipp­

istes? planorbis Dali, 1927a: 131, originally described from "off Fernandina," Florida. This deep-water species was re-described in detail by Moore (1971: 114-116, fig.

1) as Omalogifra planorbis, and subsequently called Om­ alogyra (Ammonicera) planorbis (e.g., Abbott, 1974:81;

Rios, 1994:60). This taxon was placed in Palazzia Waren,

1991, as an "archaeogastropod" group of uncertain affil­

iations, tentatively assigned to Skeneidae (Waren, 1991 : 74, 76).

"Ammonicera jischctiana": Nortlsieck (1972:149) re­ ferred to "Ammonicera fischeiiana (Monterosato. 1869)

= densecostata [sic] (Jeffreys, 1884)" in "Westindien," without further explanation. This synonymy is erroneous.

The West Indian record for this Mediterranean species

is based on Watsons (1886) "Challenger" materiiil of

"densicostata" as explained in the following.

"Omalogifra (Ammonicera) densicostata": Homalogt/ra densicostata Jeffreys, 1884:129, pi. 10, fig. 1. was de­ scribed from "Porcupine" stations (1098-2002 m) off the coast of Portugal. Additional material from a "Bulldog" cruise off Labrador (2967 m) was also included in the original description. Abbott (1974:81) reported this spe­ cies as Omalogyra (Ammonicera) densicostata Iroui deep water off Portugal, the Azores, and Labrador Moore

(1971:114) showed that the Labrador ("Bulldog") ma­ terial in fact belongs to "Lipj)i.ste.s" planorbis Dali, 1927, thus removing the Labrador record for "O." densicos­

tata. Watson (1886:677) added a "Homalogyra densicos­ tata (?)" record from north of the island of Culebra,

between Puerto Rico and the Virgin Islands ("Challeng­ er" staHon 24, 71.5 m). Moore (1971:11.5-116) doubted

both the syiionviiiv of the Challenger material and that of the shallow-water material reported by Dautzenlierg

(1889:46) for the Azores, thus restricting densicostata again to the eastern Atlantic. The species was considered a member of the eastern Atlantic omalog\Tid fauna until recently (e.g., Sabelli et al., 1990; Gaglini. 199.3). Hom- alogifra densicostata was syiionymized imtler Adeiiom-

phaliis ammonifonnis Seguenza, 1876, and placed as an

"archaeogastropod" of uncertain affiliations, tentatively assigned to Skeneidae (Waren, 1991:74 ff.).

Recognized eastern AtlanticAlediterranean Spe­ cies: [regions of type localities in brackets]

Ammonicera biimayi Rolan. 1991:112, figs. 1.3-14 [Cape Verde

Archipelago]

Ammonicera fischeriana (Monterosato, 1869:274— 275. pi. 13, fig. 1) [Mediterranean]

Ammonicern tignrn (Palazzi. 1988:105, figs. 8. 18) [Madeira]

Ammonicera midlistriata Rolan, 1991:112, 114, figs. 1.5-16

[Cape Verde Archipelago]

Ammonicera nolai Rolan. 1991:110. figs. 8-9 [Cape Verde Ar­ chipelago]

Ammonicera oteroi Rolan, 1991:110. 112, figs. 10-12 [Cape

Verde Archipelago] Ammonicern robusta Rolan, 1991:114-115. figs. 17-18 [Cape

Verde Archipelago]

Ammonicera rota (Forbes & Hanley, 18.50:160, pi. 73, fig. 10;

pi. 88, figs. 1, 2) [Ireland]

Ammonicera rotundata (Palazzi, 1988:105, figs. 10, 21, 27)

[Madeira]

Annnonicera verdensis Rolan, 1991:109, figs. 6-7 [Cape Verde

Archipelago]

Omalogt/ra afomus (Philippi, 1S41:.54, pi. 5. figs. 4a-d) [Med­

iterranean]

Onuilogi/ra di.scutus Palazzi, 1988:104, figs. 1, 20 [Madeira]

Omalogtpa simplex (O.G. Costa. 1861:72, pi. 11 figs. 3 a, b)

[Mediterranean]

Omaloayra undosa Palazzi, 1988:104, figs. 5, 15 [Madeira]

Retrototiina fuscata Chaster. 1896:2 [Strait of Gibraltar]

In the eastern Atlantic, fifteen omalogyrid species are currently recognized, comprising ten species of Ammon­

icera, four of Omalogifra. as well as Retrototiina fuscata

(for Mediterranean records see also Sabelli et al., 1990;

Le Renard et a/./CLEMAM, 1997). Omalogifra atcunus is the only species known from both sides of the Atlantic;

it is widely distributed, ranging from the Mediterranean,

Madeira, and the Azores to Norway, Iceland. Greenland, and in New England (Abbott, 1974; Bullock, 1969, 1995;

Fretter & Gniliam, 1978; Thorson, 1944) from Maine to

Rhode Island. Egorova (1991) recognized material from

Antarctic waters, previously identified and cited as O. atomits, as members of a moiphologicallv extremely sim­

ilar species, O. antarctica Egorova, 1991.

Several other nominal omalogyrid species ha\e been described for the eastern Atlantic. Ofthe.se, Omalogyra apeiia Sykes, 1925:192, Ironi off Portugal, was recog­

nized as a member of the "archaeogastropod" genus Eii-

[Begin Page: Page 9]

R. Bieler and P. M. Mikkelsen. 1998

Paee 9

daronia Cotton, 1945, Ny Waren (1991:80). Flomalogijra granulosa Svke.s, 1925, also from off Portugal, was

placed in the "archaeogastropod" genus Retigyra Waren,

1989 (see Waren, 1992:168). Flomalogijra paiadoxa

"Monterosato (? MS.)" of Svkes (1925:192) is a nomcii

niuliim. Two other noiinn;il species introduced bv Sykes {H. sititiosa Sykes, 1925, and H. (?) marshalli Sykes,

1925) are in need of further study (see Palazzi, 1992).

An additional Mediterranean species, O. ausonia P;ilazzi,

1988, was recently made the t\pe of Palazzia Waren,

1991, and transferred to the "archaeogastropods," with tentative placement in the Skeneidae (Waren, 1991).

Nominal species Homalogi/ra oniata Dautzenberg, 1889

(p. 46, pi. 4, fig. 9a-d), described from the Azores, is still

in need of reinVestigation.

It should be noted that Palazzi (1988) used "ausonia"

(Italy) and "disculus" (little chsk) as nouns in apposition

in the original descriptions; recent usage as "Palazzia ausoniac" or "Oiiwlogi/ra disciila" (e.g.. Sabelli et al.,

1990; Giannuzzi-Saveili ct al. 1994; Arduini ct al.. 1995) are incorrect subsequent spellings.

ACKNOWLEDGMENTS

This project, as part of a broader study of lower heter­ obranch gastropods, was supported under National Sci­ ence Foimdation grant DEB-9318231 to RB. Field and

laboratory' work in Florida was made possible through

Visiting Scientists Awards by the Smithsonian Marine

Station at Unk Port (SMSLP) to RB; Dr Mary E. Rice and the station staff are gratefulK acknowledged for their support. Field collecting in the Florida Keys was supported through supplementary funding from the Bertha LeBus Charitable Trust and Field Museum's

Marshall Field Fund. Dr. Kenneth J. Boss (Museum of

Comparative Zoologw Flarvard Uni\'ersit\) and Richard

E. Petit (North Myrtle Beach, South Carolina) kindly

provided hterature, and Roberto Cipriani (FMNH) as­ sisted with translations. We thank Dr. Sergio Letelier

(MNHN) for hospitality- extended during RB's \isitto

Santiago, Kathie Way (BMNH) and Thomas von Rin- telen (ZMB) for information on tvpe holdings of their respective collections. Dr. Cristian F. Ituarte (MLP) for a specimen loan, and Richard E. Petit (North Myrtle

Beach) as well as tw o anonymous reviewers for their comments on the manuscript. The excellent facilities and helpful staff of the AMNH and FMNH libraries are also ackniowledged. This is SMSLP Contribution no.

434.

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