Cainozoic Research, 2(1-2) (2002), pp. 163-170, October 2003

Notes on the systematics, morphology and biostratigraphy of

fossil holoplanktonic ,

13. Considerations on a subdivision of Thecosomata, with the

classification of Limacinidae emphasis on genus group

Arie+W. Janssen

Nationaal Natuurhistorisch Museum (Palaeontology Department), P.O. Box 9517, NL-2300 RA Leiden, the Netherlands: current

address: 12, Trig tal’ Hamrija, Xewkija VCT 110, Gozo, Malta; e-mail: [email protected]

Received 11 June 2003; revised version accepted 15 June 2003

The families Limacinidae, and Peraclididae (, Thecosomata) are raised in rank to Limacinoidea,

and and subfamilies Peraclidoidea, respectively, currently recognised ofthe Cavoliniidae are given full familial status, viz. Creseidae,

Cuvierinidae, Clioidaeand Cavoliniidae.A is preliminary genus group classificationof the Limacinidae proposed, mainlyon morphological

shell features; a new genus, Currylimacina n. gen., is introduced.

KEY WORDS: Mollusca, Gastropoda, Thecosomata, Euthecosomata, Limacinidae, pteropods, new genus, .

Introduction Systematics of Thecosomata

Currently, holoplanktonic gastropods, usually referred to as In the present paper, the systematics of Thecosomata, and of

‘’, are subdivided into two orders, Thecosomata the Limacinidae in particular, is considered. The currently

and Withinthe Thecosomata suborders Gymnosomata. two applied taxonomy is almost entirely based on the relatively

are recognised, namely Euthecosomata and Pseudothe- few, extant species. From the fossil record, however, numer-

cosomata. ous taxa are known which cannot always be incorporated in

The Euthecosomata comprises three families, Limacini- the existing systematic framework satisfactorily.

Cavoliniidae and all of soft of Recent dae, Sphaerocinidae, representatives Although part anatomy most representa-

which are shelled in the adult state. Both Limacinidae and tives of the Euthecosomata has been studied extensively,

Cavoliniidae are known from the fossil the earliest identification of taxa is record, species group predominantly based

occurrence being Late Paleocene, and are still extant; on shell morphology. Limacinidae differfrom Cavoliniidae

known fossils. Withinthe in Sphaerocinidae are exclusively as basically possessing an ultra-dextrally coiledshell which,

Pseudothecosomata three families is illustrated are distinguished, namely however, usually as being sinistral. In the

Peraclididae(adult specimens are shell bearing), Cymbulii- Cavoliniidae, the shell is not coiled, but either coni-

dae shell shed, but adult individuals with (larval a cartilagi- cal/cylindrical (Creseinae, Cuvierininae) or bilaterally sym-

so-called and with ventral and dorsal shell nous pseudoconcha) Desmopteridae (adult metrical, parts more or less shell The latter lacking or pseudoconcha). family, distinctly separated (Clioinae, Cavoliniinae). The Sphaero-

unknown from the fossil record to date, is occasionally cinidae differ from both in showing cavoliniid larval shell

considered to belong to the Gymnosomata (van der Spoel, features and secondary dorso-ventralcoiling ofthe shell.

1976, p. 45). Differences between Limacinidae on the one hand and

In the five larval Cavoliniidae+ Gymnosomata, containing families, only Sphaerocinidae on the other, taken together shells These shed are present. are shortly after hatching, the with a large data set for fossil members, have led me to

adult ‘naked’. few of these larval consider these specimens being Only two groups as superfamilies rather than fami- shells have been described which so far, explains why little lies. This point of view is also based on the fact that both

is known about fossil which date back the and fossil representatives, to Recent pteropods apparently comprise far more

Miocene. taxa at the species level than previously assumed. Examples - 164-

of this are given by e.g. van der Spoel et al. (1993) and In accepting a superfamily Cavolinioidea, the next logical

views would consider the subfamilies Janssen (in press); the observation is also supported by step be to existing (Cresei-

Norris The of fossil Clioinae and families. A expressed by (2000, p. 254). study nae, Cuvierininae, Cavoliniinae) as representatives also yields a more reliable insight into the similar argumentation appears valid for the Pseudothecoso-

subdivision of evolutionary development of this group, which may be mata, which leads to the following (Table 1)

from the the Thecosomata. The ofthe is expected to be far more complex than judged authorship new higher taxa relatively few Recent species alone. attributed according to the Principle ofCo-ordination(ICZN,

also advisable rank the In this context, it might be to art. 36.1, p. 45).

Sphaerocinidae as a superfamily. However, for the time To my knowledge, DNA structures of extant Thecoso-

such studies being I prefer to retain it as a family within the Cavolini- mata have not yet been studied. I expect that

would additional criteria which will either confirm oidea, in view ofthe fact that but very few species are known yield or to ofwhich has been described far. refute considerations this outlinedhere. date, only one so my on subject as

Order Thecosomata de Blainville, 1824

Suborder Euthecosomata Meisenheimer, 1905

Superfamily Limacinoidea Gray, 1847

Family LimacinidaeGray, 1847

Superfamily CavolinioideaFischer, 1883

Family Creseidae Rang, 1828

Family Cuvierinidae van der Spoel, 1967

Family Clioidae van der Spoel, 1967

Family CavoliniidaeFischer, 1883

Family Sphaerocinidae Janssen & Maxwell in Janssen, 1995

Suborder Pseudothecosomata Meisenheimer, 1905

Superfamily Peraclidoidea Tesch, 1913

Family Peraclididae Tesch, 1913

Family Cymbuliidae Cantraine, 1841

Subfamily Cymbuliinae Cantraine, 1841

Subfamily Glebinae van der Spoel, 1976

?Family Desmopteridae Chun, 1889

Table 1. Proposed subdivision ofthe Thecosomata.

outlined here did mine. A taxonomy which is comparable to the one as Fortunately, Jeffery permitted me to publish has recently been proposed by Jeffery (2003). Jeffery consid- my results independently. ered the Thecosomataand Gymnosomata to be suborders; the Euthecosomataand Pseudothecosomatain his scheme are

ofThecosomata. The subdi- Genus level of Limacinidae infraorders Gymnosomata are taxonomy vided into the infraorders Gymnosomata and Gymnoptera.

Within the Euthecosomata, however, Jeffery distinguished The systematics ofthe Limacinidae are here enlarged upon

three based shell from but a single superfamily (Limacinoidea), comprising further, mainly on morphology as gleaned families, namely Cavoliniidae, Limacinidae and Sphaero- existing literature and available collections. No attempt has

but been made unravel in detail the cinidae. I have no opinion on the rank of the higher taxa, yet to undoubtedly quite consider the Limacinoidea and Cavolinioidea to be more complex evolutionary history of this family. This may be

lead natural subdivision. than sufficiently differentto represent separate superfamilies. expected to eventually to a more

I have contacted Paul Jeffery on this subject; he informed me The application of subgeneric names in RecentLimacina is

the lack of data. that his relative ranking of the Thecosomata as given in his frequently seen, despite any phylogenetic In

the result of overall in the of of trends in the website was an adjustment anticipation an analysis evolutionary genus

inclined ranking of the superorders Heterobranchiaand Caenogas- , I am now to use only generic names.

in tropoda, thus subunits ofthe Euthyneura were changed rank (generally reduced, actually). The scheme on his Limacinidae (revised diagnosis) — Marine, holoplanktonic Soft dextral, shell webpage arose largely as a matter of systematic conven- euthyneuran gastropods. part anatomy

of Thecosomata, coiled in an ultra-dextral thin- ience and not from any special study the (seemingly sinistral) spiral, - 165 -

conical-almost with walled,ranging in form from discoidal to high or a subperipheral thickened zone projecting as a ros-

Larval shell Surface the Umbilicus from wide in cylindrical. not clearly separated. orna- trum at aperture. ranging very

ment usually absent, but a divaricate or collabral micro- some discoidal forms, to very narrow in high-conical species,

ornament is found in some forms. In the normally smooth or even entirely absent. The columella is simple, sometimes

forms occasionally a weak spiral or collabral striation may with a weak central fold, twisted, or with a three-dimensional

occur as a senile characteristic. Aperture either simple or torsion. Stratigraphical range: Late Paleoceneto Recent.

widened, with an internally or externally reinforced margin,

Name Type species Designation

Altaspiratella Korobkov, 1966 elongatoidea Aldrich, 1887 original

Chaduma Korobkov, 1966 chadumica Korobkov, 1966 original

ChadumellaKorobkov, 1966 planorbella Korobkov, 1966 original

Crino Gistel, 1848 arctica Fabricius, 17801780 monotype

Embolus Jeffreys, 1869 rostralis Souleyet in Eydoux & Souleyet, 1840 original

Heliconoides d’Orbigny, 1836 inflata d’Orbigny, 1836 Herrmannsen, 1846

Heterofusus Fleming, 1823 retroversus Fleming, 1823 Gray, 18471847

Limacina Bose, 1817 Phipps, 1774 monotype

this Munthea van der Spoel, 1967 trochiformis d’Orbigny, 1836 paper

?PakistaniaIPakistaria Lames,Eames, 19521952 antirotataFames,Eames, 1952 monotype

Planorbella Gabb, 1873 nonnon Haldeman, 1843 imitans Gabb, 1873 monotype

Plotophysops Curry, 19811981 bearnensis Curry, 1981 original

Protomedea Costa, 1861 elata Costa, 18611861 monotype

Tembrock, 1989 Pygmella pygmaea (Lamarck, 1804) original Scaea Philippi, 1844 lunaris Gmelin, 1791 Herrmannsen, 1849

Skaptotion Curry, 1965 bartonense Curry, 1965 monotype

Spiratella Blainville, 1817 helicina Phipps, 1774 monotype

Spirialis Eydoux & Souleyet, 1840 trochiformis d’Orbigny, 1836 Herrmannsen, 18491849

Striolimacina Janssen, 1999 imitans(Gabb, 1873) monotype

ThieleaStrebel, 1908 emend. Tesch, 1913 Strebel, 1908 procera Strebel, monotype

Valvatina Bomemann, 1855 umbilicataBomemann, 1855 monotype

Table 2. Existing names of the genus group in Limacinidae (in alphabetical order).

will when evolution- Available limacinidnames satisfactory taxonomy only be possible

trends within this better understood. At ary family are pres-

The above diagnosis indicates that a wide variety of shell ent, the fossil record still shows too many hiatuses which

of which known This tentative forms is represented, most are only as obscure relationships. explains why only a

fossils; only seven, maybe eight, extant species are included classification is possible at thistime.

in the Limacinidae. Naturally, forthese not only the shells are

for the also and known but, majority, anatomy, ecology

geographical distribution.A considerable numberof papers Recent species

is dedicated to these subjects; there can be no doubt that

these species all belong to the same family. Fossil taxa, To a large extent, the taxonomy proposed here is based on

however, outnumberRecent ones by far. Quite a number of discussions of the systematics of the few extant forms. In

(sub)generic names have been introducedover the years for recent years, this subject was reviewed by e.g. van der Spoel

Recent or fossil limacinids (Table 2). In view ofthe fact that (1967), Rampal (1975) and Wells (1978). Concepts outlined

many fossil Limacinidae differ considerably in shell mor- by the last-named author in particular, who applied mainly

phology from Recent taxa, it is no surprise that especially anatomical features together with some ecological and shell

from the fossil record new genera were introduced. On the characters, appear to be well foundedand acceptable. Wells

other hand, limacinid pteropods have occasionally not been opined that all Recent members ofthe Limacinidae should be

recognised as such and have been erroneously assigned to assigned to the genusLimacina, with in subgenus Limacina

Homalaxis, and Planorbis. str. the bulimoides helicina genera such as Aplexa, Physa In s. species (d’Orbigny, 1836),

spite ofthe number of names available in the genus group, (Phipps, 1774), lesueuri (d’Orbigny, 1836), retroversa

is far from settled. is fearedthat and in limacinid systematics It a (Fleming, 1823) trochiformis (d’Orbigny, 1836); - 166-

Thieleathe helicoides and which is distinct. fossils subgenus species (Jeffreys, 1877) base, usually Amongst there are in Embolus the subgenus species inflata (d’Orbigny, 1836). species which lack an umbilicus altogether. Species of

Anatomical features discussed Wells of have by are position Altaspiratella a high conical shell, but are characterised the mantle ofthe cavity, development parapodia, number of especially by a three-dimensionally twisted columellaand a cell of the of the zones pallial gland, position excurrent lack ofan umbilicus. This genus comprises only four species,

siphon, reproductive mechanisms and presence or absence of namely L. elongatoidea (Aldrich, 1887), L. bearnensis tentacular lobe. he included a In addition, some data on (Curry, 1981),L. multispira (Curry, 1981) and L. gracilens habitat (epipelagic or bathypelagic) and on geographical Hodgkinson, in Hodgkinson, Garvie & Be, 1992, all of Early distributionof species. Shell was used in a to Middle morphology very Eocene age. Of special note is the fact that these restricted sense with Wells himself data that there transition only, confining to on species strongly suggest was a gradual to the biomineralogy ofthe shell wall (prismatic, cross-lamellar, Camptoceratops, to whichthe morphology oftheir apertural or both) and to a highly generalised indicationof shell form reinforcementsin particular seems to point, but that taxon has

‘normal’ (spire or ‘flattened’). never been consideredto belong in the Limacinidae.Forthe

From this threefold subdivision ofthe time Limacina group, a being, I interpret Camptoceratops as a (primitive) few additional distinguishing features of a strictly con- creseid, in accordance with previous authors.

nature chological may be extracted. The five species united For a species from the Miocene of the Dominican Re- in Limacina s. str. all have simple apertural margins, lacking public, Limacina imitans (Gabb, 1873) (see Janssen, 1999,

and all have their shells coiled which differs from all other limacinids widening or reinforcements, p. 13), in having a in a or lower helicina higher spatial spiral (in L. antarctica posteriorly divaricating micro-ornamenton the body whorl,

1854 der the the Woodward, [see van Spoel, 1967, figs 5a, 7a], subgenus name Striolimacina (= Planorbella Gabb, shell be almost and all have umbilicated introduced. may discoidal) 1873, non Haldeman, 1843) was

albeit that the umbilicus is shells, very narrow to nearly Another species which merits separation at the generic

‘ closed in L. bulimoides. level is ‘Skaptotion’ cossmanni Curry, 1981, and its probable

A discoidal shell, with planorboid whorls in which an synonym Skaptotion? reklawensis Garvie in Hodgkinson, internal subperipheral thickening produces an anterior ros- Garvie & Be, 1992. This is a near-spherical species, whose trum that reinforces the second half of the whorl, whorl body body to a large extent encloses earlier whorls, with a characterises the sole referred for which but thickened species to Embolus, faintly apertural margin. This species shows a an older name is available. This species also is the only highly regular collabral micro-omamentwhich is unknown

Recent taxon with brood whileits struc- protection, genital in any other pteropod. Garvie (in Hodgkinson, Garvie & Be, ture differs considerably from that ofother limacinids(Wells, 1992, p. 23) considered S.? reklawensis to be dextral, the

‘ ’ 1978). illustrations however are reminiscent of Skaptotion coss-

The sole of Thielea is much thanall other the ofwhich species larger manni, types I have studied. Additionalmaterial

diameter of 15 This is the from the of limacinids, attaining a mm. only type locality S. cossmanni as well as from locali-

limacinid in which extant the columella demonstrates a ties within the North Sea Basin is before me (collections of distinct coaxial torsion. In it is characterised addition, by a Nationaal Natuurhistorisch Museum, Leiden). The type of bathypelagic way oflife and an ovoviviparous reproductive Skaptotion, S. bartonense Curry, 1965, differs markedly in

with system. being completely discoidal, a large, platform-like

’ ‘ extended apertural margin. To contain Skaptotion coss-

manni here I propose the new genus Currylimacina n. gen.,

Fossil Limacinidae in honour ofDennis Curry, who died recently.

‘Limacina’adornata Hodgkinson in Hodgkinson, Garvie

the subdivision Wells for fossil Applying proposed by (1978) & Be (1992, p. 14, pi. 1, figs 6-9) differs from all known limacinids with the fact limacinids in meets difficulties, being hampered by showing a relatively coarse ornament of spirals that their is far than that and radial elements. Should this be morphological diversity greater species turn out to a found amongst the few Recent forms. pteropod, which I doubt, it would merit separation at the The offossil Thielea only representatives are two Mio- generic level as well.

records of shells that be cene (Early Tortonian) appear to All other names in Table2 are based on type species with conspecific with the Recent species, from Monterotondoand umbilicated shells, having either a reinforced or a simple

in northern Stazzano Italy, and a single, comparable find apertural margin, and varying in shape from discoidalto high from the Pliocene in the conical. It is discoidal (Piacenzian) Estepona area (SE tempting to use a shell form as a major

Spain; see Janssen, in prep.: material contained in collections taxonomic feature, but, as demonstratedby Giirs & Janssen of Nationaal Natuurhistorisch Museum, Leiden and (2002), originally conical species may develop over geologi- Frankfurt Senckenberg Museum, am These records cal time into with discoidal shell. This Main). species a may occur be the of life of this within may explained by bathypelagic way a relatively short time span: for instance, L. valvatina which wouldhave species, virtually precluded its occurrence (Reuss, 1867), a limacinidwith an almost ‘ideal’ Limacina in shallow-water shelf from which of or deposits, most the shape (e.g., HAV ratio close to 1, a regular spire with straight fossil record was obtained. tangentsand no apertural reinforcements). In the North Sea

All Limacina have modem species of s. str. a perforated Basin, this species ranges from the Late Oligocene to the - 167-

Late Miocene, showing only a narrow range of variation, The presence or absence ofapertural reinforcements, on

the the well be characteristic in with solely some changes in H/W ratio. However, during contrary, may applied as a major

ofthe Serravallian inNorth subdivisionof Limacina. Such structures younger portion (‘Langenfeldian’ a apertural appear

Sea Basin chronostratigraphy), L. valvatina suddenly starts in a multitude of forms, e.g. as described for Recent Li- to develop rapidly into two directions. On the one hand, there macina inflata above.

is an increase in H/W ratio (exceeding 1.10; Limacina Other species, independent of a helical or discoidal gramensis Rasmussen, 1968), andon the other a reduction of shape, demonstrate different types of apertural reinforce-

the which results in discoidal ments. These internal external same, eventually a shell, as may occur as a simple or

demonstrated in the lineage L. valvatina f. weinbrechti thickening of the apertural margin, an internalpreapertural

Tembrock, 1989 > L. ingridae Janssen, 1989 > L. wilhelmi- thickening or rim, a terminal widening, flange-like structures,

atlanta The last- denticles. is difficult how these different nae Janssen, 1989 > L. (Morch, 1874). or It to see basically

namedmember in this lineage ranges from the Late Miocene morphologies could have developed from each other; a better

(Late Tortonianor Gramian in North Sea Basin stratigraphy) insight into these trends may eventually lead to distinguishing

to far into the Pliocene. more taxa. Too little is known now to reconstruct reliable

In L. ingridae the shell is much wider than high (H/W lineages, which is why I retain all forms with apertural ratio c. 0.55), with a flat to almost flat apical plane, in which reinforcements in a single ( Heliconoides see below). genus ,

the apical worls may or may not protrude above younger

whorls. The same is true for two stratigraphically younger

species, L. wilhelminaeand L. atlanta, which differby then- Proposed classification ofLimacinidae

peculiar way ofcoiling (Janssen, 1989). None of the species

in this shows reinforcements. the main be lineage, however, any apertural Using features, seven groups may distinguished

If helical and discoidal shell forms occur within a single (Table 3). Table 2 contains twentyone genus-level names for

lineage the conclusion must be that this featureshould not be the Limacinidae, and it is from these that valid names may be

used if the selected. these in order to designate supraspecific taxa, even extremes are Grouping names chronological yields

widely separated. the correct names and theirsynonyms (Table 4).

Body whorl ornamented

with radialelements 1 Relatively coarse spiral ornament, some group Shell with micro-ornamentonly

Micro-ornament backwards divaricating group2

Micro-ornamentcollabral group 3

Body whorl without ornament (except growth lines)

Columellain a three-dimensionalspiral, umbilicus absent group 4

Columella with umbilicus almost simple or torsion, present (sometimes closed)

Apertural margin simple

Shell small, columellawithout distinct torsion group 5

Shell 15 distinct torsionof columella large (to mm), group 6

Apertural margin widened or otherwise reinforced group 7

Table 3. Groupingof limacinid species on the basis of main shell features.

Group 1

no name available (pteropod ?)

Group 2

PlanorbellaGabb, 1873 non Haldeman, 1843

Striolimacina Janssen, 1999

Group 3

CurrylimacinaCunylimacina n. gen.

Group 4

Altaspiratella Korobkov, 1966

Plotophysops Curry, 1981 - 168-

Group 5

LimacinaBose, 1817

Spiratella dedeBlainville, 1817

Heterofusus Fleming, 1823

SpirialisSpinalis Eydoux & Souleyet, 1840

Scaea Philippi, 1844

Crino Gistel, 1848

Valvatina Bomemann, 1855

IPakistania?Pakistania Fames, 1952

Chaduma Korobkov, 1966

ChadumellaKorobkov, 1966

Munthea van der Spoel, 1967

Pygmella Tembrock, 1989

Group 6

Thielea Strebel, 1908 emend. Tesch, 1913

Group 7

Heliconoides d’Orbigny, 1836

Protomedea Costa, 1861

Embolus Jeffreys,Jeffreys, 1869

Skaptotion Curry, 1965

Table 4. Distribution of available generic names over the various groups.

it be determined what Naturally, cannot now to extent a taxonomy. classification such as this reflects naturalrelationships, but all The synonymy ofLimacina and Spiratella (group 5) was available data on Recent forms make it at least probable. discussed by Janssen & Zom (2001). In summary, a new

Additionaldata ruled in is here for the Limacinidae on soft-part anatomy are out fossils, taxonomy proposed (Table 5). As but a further scrutiny of limacinid evolution would seem to mentioned above, I expect Heliconoides in particular to be a be a promising tool to confirm and refine or refute this polyphyletic group.

Genus Type species

Altaspiratella elongatoidea {(Aldrich,Aldrich, 1887)

Currylimacina cossmanni (Curry, 1981)

Heliconoides inflata (d’Orbigny, 1836)

Limacina helicina (Phipps, 1774)

Striolimacina imitans(Gabb, 1873)

= Thielea procera Strebel, 1908 helicoides(Jeffreys, 1877)

‘ New genus ? pteropod ? Limacina'‘Limacina’ adornata Hodgkinson, 1992

Table 5. Proposed subdivision of the Limacinidae (in alphabetical order).

Acknowledgements stalt, Vienna, Austria) commented on the views expressed in

this paper, while Dr J.W.M. Jagt (Venlo, the Netherlands)

Dr P. Jeffery (Natural History Museum, London) is thanked improved the English. for to views Thecosomata permitting me publish my on systematics, although similar ideas expressed by himappear on the worldwideweb. F.P. Wesselingh (Nationaal Natuur- References historisch Museum, Palaeontology Department, Leiden, the

Netherlands) and Dr Mag. I. Zom (Geologische Bundesan- Aldrich, T.H. 1887. Notes on Tertiary fossils, with descriptions - 169-

ofnew Journal the Cincinnati Natural naturae, secundum ordines, species. of Society of classes, genera, species cum

78-83. XIII History 10, characteribus, differentis, synonymus, locis. Editio aucta de Blainville, H.M.D. 1817. Clio. Dictionnaire des Sciences reformata 1(6), 3021-3910.Lipsiae.

naturelles 404-407. A 9, Gray, J.E. 1847. list of the genera of recent Mollusca, their

H.M.D, de Dictionnaire des and Blainville, 1824. Mollusques. synonyma types. Proceedings ofthe Zoological Society of

Sciences naturelles 32, 1-392. London 15, 129-219.

Bomemann, A. 1855. Die mikroskopische Fauna des Septarien- Gtirs, K. & Janssen, A.W. 2002. Revised pteropodbiostratigra-

thones von Flermsdorfbei Berlin. Zeitschrift der deutschen phy for the Miocene of the North Sea Basin. In: Gtirs, K.

geologischen Gesellschaft 7, 307-371, pis 12-21. (ed.). Northern European Cenozoic Stratigraphy [Proceed-

Bose, L.A.G. 1817. Clio. Nouveau Dictionnaire d’Histoire ings of the 8th biannual meeting of the RCNNS, RCNPS],

naturelle 7, 194. 117-131,2 pis. Flintbek (Landesamt fUr Natur und Umwelt

Cantraine, F. 1841. Malacologie mediterraneenne et litorale, ou Schleswig-Holstein). des vivent dans la Mediterranee description mollusques qui Haldeman, S.S. 1840-1845. A monograph of the limniades, or le continent de ainsi des ou sur 1’Italie, que coquilles qui se fresh-water univalve shells ofNorth America, continuedas

trouvent dans les terrains tertiaires avec des observa- the italiens, a monograph of freshwater univalve Mollusca of the tions leur leurs leur leur sur anatomie, moeurs, analogic et UnitedStates, including notices of species in other parts of

gisement. Ouvrage servant de faune malacologique italienne North America. Philadelphia.

et de la fossile de A.N. complement a Conchiliologia subapennina Herrmannssen 1846-1849. Idicis generum malacozoorum

Brocchi. Memoires de VAcademic de Bruxelles Nomina royale 13, primordia. subgenerum, generum, familiarum, 1-173, 6 pis. tribuum, ordinum, classium; adjectis auctoribus, temporibus, C. Bericht Chun, 1889. ilber eine nach den Canarischen Inseln im locis systematicis atque literariis, etymis, synonymis. Prae-

Winter 1887/1888ausgefuhrte Reise. Sitzungs-Berichte der termittuntur Cirripedia, Tunicata et Rhizopoda, 1-2, xxvii +

Akademie xlii 717 kdniglichen preussischen der Wissenschaften zu 637 pp. (1846), + pp. (1847-1849). Kassel (T. Berlin 1889, 519-553, pi. 3. Fischer). Costa, O.G. 1861. Microdoride mediterranea e descrizione de Hodgkinson K.A., Garvie, C.L. & Be, A.W.H. 1992. Eocene

ben conosciuti poco od affatto ignoti viventi minuti e micro- euthecosomatous Pteropoda (Gastropoda)of the Gulf and east-

del xviii + 80 1 1-12. Na- North scropici Mediterraneo, pp., pis A, ern coasts of America. BulletinsofAmerican Paleontol-

13 poli (Iride). ogy 341,5-62, pis. D. 1965. The Curry, English Palaeogene pteropods. Proceedings International Commission on Zoological Nomenclature 1999. the of Malacological Society ofLondon 36,357-371, 1 pi. International code ofzoological nomenclature, fourth edi-

D. 1981. eocenes Curry, Pteropodes de la tuilerie de Gan (Pyre- tion, xxix + 306 pp. London (The International Trust for

nees-Atlantiques) et de quelques autres localites du SW de la Zoological Nomenclature). France. Cahiers de Micropaleontologie 4, 35-44, 1 pi. Janssen, A.W. 1989. Some new pteropod species from the North

Eames, F.E. 1952. A contribution to the study of the Eocene in Sea Basin Cainozoic (Mollusca: Gastropoda, Euthecosomata).

western Pakistan and western India, C. The description of the Mededelingen van de Werkgroep voor Tertiaire enKwartaire

Scaphopoda and Gastropoda from standard sections in the Geologic 26, 91-133, 8 pis. Rakhi Nala and Zinda Pir of the areas western Punjab and in Janssen, A.W., 1995 Systematic revision of holoplanktonic

the Kohat district. Philosophical Transactions oftheRoyal Mollusca in the collections of the ‘Dipartimento di Scienze London Society of B236, 1-168, 6 pis. della Terra’ at Torino, Italy. Museo regionale di Scienze

Eydoux, F. & Souleyet, F.L.A. 1840. Description sommaire de naturali, Torino, Monografie 17, 1-233, 14 pis.

ou 1999. plusieurs pteropodes nouveaux imparfaitement connus, Janssen, A.W. Neogene paleontology in the northern

destines etre Le Revue a publics dans voyage de la Bonite. Dominican Republic, 20. Holoplanktonic mollusks (Gastro-

235-239. zoologique,par la Societe Cuvierienne, Paris, poda: Heteropoda and Thecosomata). Bulletins ofAmerican

Fabricius, O. 1780. Fauna groenlandica, systematice sistens Paleontology 358, 5-40, 5 pis.

animalia occidentalis hactenus A.W. in groenlandiae indagata, Janssen, prep. Holoplanktonic molluscan assemblages

quoad nomenspecificum, triviale, vernaculumque; synonyma from the Pliocene of Estepona (Spain, Malaga).

auctorum locum, of plurium, descriptionem, victum, generatio- Janssen, A.W. in press. Evolutionary development Cuvierinidae

nem, mores, usum, capturamque singuli, prout detegendi oc- (Mollusca, Euthecosomata) during the Cainozoic: a non- secundum observatio- casioflit, maximaqueparte proprias cladistic approach with a re-interpretation of Recent taxa. Jour- 452 I Hafniae nes, pp., pi. et Lipsiae (I.G. Rothe). nalofsystematic Palaeontology. 1880-1887. Manuel de de Fischer,?. conchyliologie et paleon- Janssen, A.W. & Zom, I. 2001. Notes on the systematics, mor- histoire tologie conchyliologique ou naturelle des mollusques phology and biostratigraphy of fossil holoplanktonic Mol-

vivants et d’un les fossiles suivi appendice sur brachiopodes lusca, 11. Limacina Bose, 1817: precedence over Spiratella

D P. i-xxiv + 1369 par Oehlert, 1-12, pp., 23 pis. Paris Blainville, 1817 (Mollusca, Gastropoda: Euthecosomata).

(Savy). Basteria 65, 147-149.

J. 1823. On a reversed of Fleming, species Fusus, (Fusus retro- Jeffery, P. 2003. Gastropod classification, http://www.nhm.ac.uk/ versus). Memoirs ofthe Wernerian Natural History Society palaeontology/i&p/gastroclass/gastroclass.htm#HETERO

4, 498-500, pi. 15. BRANCHIA

Gabb, W.M. 1873. On the topography and of Santo J.G. 1869. British geology Jeffreys, conchology, or an account of the

Transactions the American Soci- Domingo. of Philosophical Mollusca which now inhabit the British Isles and the sur-

49-259. ety n.s. 15, rounding seas, 5. Marine shells and naked Mollusca to the Gistel, J.R.F.X. 1848. Naturgeschichte des Thierreichs. Fur end ofthe Gastropoda, the Pteropoda, and Cephalopoda:

hohere Schulen, i-xvi + 216 atlas 32 pp., pis. Stuttgart with a supplement and other matter, concluding the work,

258 London (Hoffmann). pp., 108 pis. (van Voorst).

F.J. 1791. Caroli Linne Gmelin, a systema naturaeper regna tria Jeffreys, J.G. 1877. New and peculiar Mollusca ofthe Eulimidae - 170-

and other families of Gastropoda, as well as of the Pteropoda, the marineyounger Mioceneformations in Denmark, 2. Pale- procured in the ‘Valorous’ The Annals and ontology. Danmarks expedition. geologiske Undersogelse (2)92,265 pp„ MagazineofNatural History (4)19(112),317-339. 27 pis.

Korobkov, LA. 1966. Krylonogie (Mollusca Pteropoda) Paleoge- Reuss, A.E. 1867. Die fossile Fauna der Steinsalzablagerungen

novykh otlozhenij juga S.S.S.R. Voprosy Paleontologii 5, von Wieliczka in Galizien. Sitzungsberichte der mathema-

71-92, pis 131-139. tisch-naturwissenschaftlichen Classe der kaiserlichen

Meisenheimer, J. 1905. Pteropoda. Wissenschaftliche Ergebnisse Akademie der Wissenschaften Wien 55, 17-182, 8 pis.

der deutschen dem ‘Valdi- S. remarkable Tiefsee-Expedition auf Dampfer Spoel, van der 1967. Euthecosomata, a group with

via’ 1898-1899(9),i-vi + 314 pp., atlas 27 pis. developmental stages (Gastropoda, Pteropoda), 375 pp. Go-

Morch, O.A.L. 1874. Forsteningeme i Tertiaerlagene i Danmark. rinchem (J. Noorduijn).

In: S. der 1976. Meddelelserpaa del 11. skandinaviske Naturfors hermode Spoel, van Pseudothecosomata, Gymnosomata and

484 i Kjebenhavn 1873,274-298. Heteropoda (Gastropoda), pp. Utrecht (Bohn, Scheltema Norris, R.D. 2000. Pelagic species diversity, biogeography, and & Holkema).

evolution. In: Erwin, D.H. & Wing, S.L. (eds). Deep time. Pa- Spoel, S. van der, Bleeker, J. & Kobayasi, H. 1993. From Cavolinia

leobiology’s perspective. Paleobiology, 26 (Suppl.), 236-258. longirostris to twenty-four Diacavolinia taxa, with aphyloge-

Orbigny, A. d’ 1836-1846. Voyagedans I Amerique meridionale(le netic discussion (Mollusca, Gastropoda). Bijdragen tot de

Bresil, la republiqueorientate de I'Uruguay, la republique Ar- Dierkunde62, 127-166, 3 pis.

gentine, laPatagonie, la republiquedu Chili, la republique de Strebel, H. 1908. Die Gastropoden (mil Ausnahme der nackten

Bolivia, larepublique duPerou), executependant les annees Opisthobranchier). In: O. Nordenskjbld (ed.). Wissenschaftli-

1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833, 5(3). Mol- che Ergebnisse der schwedischen Siidpolar-Expedition 1901- lusques’, 49-184 (1836), atlas 20pis (1846). Paris (Bertrand). 1903, 6 (Zoologie) 2, 1-111, 6 pis. Stockholm/Berlin/-

Philippi, R.A. 1844. Enumeratio molluscorum Siciliae cum Paris/London (Lithographisches Institut des General-

viventium tertiaria & & turn in tellure fossilium, quae in itinere stabs/Asher Co./Haar Steinert, Eichler/Dulau& Co).

suo observavit 2, i-iv + 303 pp., pis 13-28. Halis Saxonum Tembrock, M.L. 1989. Neue Spiratella-Arten (Gastropoda, (E. Anton). Opisthobranchier, Pteropoda). Zeitschrift fur angewandte

C.J. 1774. A towards the North Pole undertaken 1 Phipps, voyage Geologic 35, 242-244, pi.

by his Majesty’s Command177, 275 pp. London (W. Bowyer Tesch, J.J., 1913. Mollusca, Pteropoda. In: F.E. Schulze (ed.).

& J. Nichols). Das Tierreich. Eine Zusammenstellung undKennzeichnung

J. 1975. Rampal, Les thecosomes (mollusques pelagiques). der rezenten Tierformen 36, i-xvi + 154 pp., Berlin (Fried-

Systematique et evolution. Ecologie etbiogeographie mediter- berger & Sohn).

485 Aix-Marseille thesis Universite de F.E. in raneemes, pp. (unpubl. Wells, 1978. Subgeneric relationships the euthecosomatous

Provence CNRS Limacina 1817. Journal the Mala- AO 11932). pteropod genus Bose, of

Rang, P.C.A.L. 1828. Notice sur quelquesmollusques nouveaux cologicalSociety ofAustralia 4, 1-5. etablissement appartenant au genre cleodore, et et Woodward, S.P., 1854. A manual of the Mollusca; or, a rudi-

du treatise shells: i-xvi 486 monographic sous-genre creseis. Annales des Sciences mentary ofrecent andfossil + pp.,

naturelles 13,302-319,pis 17, 18. 24 pis. London (J. Weak).

Rasmussen, L.B. 1968, Molluscan faunas and biostratigraphy of