Cainozoic Research, 4(1-2), pp. 97-108, February 2006

Revision of the gastropod family

from the Danian (Early Paleocene) of Fakse, Denmark

Kai+Ingemann Schnetler¹ & Richard+E. Petit²

1 Fuglebakken 14, Stevnstrup, DK-8870 Langd, Denmark; e-mail: [email protected]

2 P.O. Box 30, North Myrtle Beach, SC 29597-0030, U.S.A.; e-mail: [email protected]

Received 11 June 2004; revised version accepted 18 March 2005

The rich mollusc fauna fromthe MiddleDanian ofFakse, Denmark contains ofthe Cancel- very deposits eight species gastropod family lariidae. Ofthese six Unitas anderseni Unitas Plesiotriton Ad- species are previously undescribed, viz: n. sp., aliceae n. sp., stem n. sp., metula Admetula and Tatara danica A for Semitriton is des- rosenkrantzin. sp., faksensisn. sp. n. sp. lectotype biplicatus (Ravn, 1902)

ignated.

Key words:: , ,Cancellariidae, Danian, Early Paleocene, Fakse, Denmark, new species.

Introduction Schilder 1928,Rosenkrantz 1960; SchnetlerefaZ. 2001). A

rare part of this fauna is the gastropod family Cancellarii-

dae, to which Ravn (1933)referred three species only in his

monograph on the Fakse fauna. Since additionalmaterial

has been collected (Spren Bo Andersen and Sten Lennart

Jakobsen in 1972, Alice Rasmussen in the 1990s), a revi-

sion ofthe family is possible. Inthe Rosenkrantzcollection

additional specimens were located. These specimens were

collected by him in 1933 and thus not treated in the revi-

sion by Ravn (1933). Eight species of Cancellariidae are

recognized, of which six are newly described herein, viz:

Unitas anderseni n. sp., Unitas aliceae n. sp., Plesiotriton

Admetula rosenkrantzi Admetula jakobseni n. sp., n. sp.,

faksensis n. sp. and Tatara danica n. sp.

Geological setting and stratigraphy

The limestone in the quarry east ofthe small town ofFakse

in Eastern Zealand has been excavated for several hundred

years. Fakse is situatedin the Danish-Polish Trough, which

is boundedto the north the FennoscandianShield and to structures by Figure 1. Main and facies ofthe Danish area in

the south the A sec- Middle Danian. Compiled by Erik Thomsen, Geologisk In- by Ringkpbing-Fyn High (Figure 1). tion mound is stitut, Aarhus Universitet. (Reproducedfrom Schnetler et through a bryozoan-coral complex exposed

al, 2001). The boundary to the north to the Fennoscandian in the quarry. In the complex a vast numberofinterbedded

Shield is indicated. microfacies results of may be recognized, as early diagene-

sis: Bryzoan limestone, chalky limestone and coral lime-

of Denmark The Danian deposits Fakse, containa very rich stone (Bemecker & Weidlich, 1990;Willumsen, 1995).For

mollusc fauna (Ravn 1902a; 1902b; 1933; Nielsen 1919; a review see Surlyk & Hakansson (1999). Desor (1847) -98 -

introduced the Danian Stage with the typical localities Fakse was discussed by Ravn (1933) and more recently

Fakse Quarry and Stevns Klintand consideredthe stage as Schnetler (2001) noted the fauna in comparison with the

the of the Cretaceous youngestpart System. For manyyears mollusc fauna ofthe younger (Selandian) Lellinge Green- the ofthe Danian has been discussed, sand. The stratigraphic position very rich faunaofthe “nose-chalk” is character- e.g. by Nielsen (1919), Ravn (1925) and Rosenkrantz ised by e. g. numerous Pleurotomariidae, trochoids,

(1938), but is now generally considered as the oldest part Cerithiopsidae, Triphoridae, Ranellidae and. cypraeids. of the Paleocene. Various subdivisions ofthe Danian have Many small species are undescribed. Among the gastro- been and the Fakse has suggested, sequence at been re- pods, supposed sponge eaters were common (Pleurotom- ferred the local bruennichi echinoid to Tylocidaris Zone ariidae, Cerithiopsidae, Triphoridae), while sometrochoids

(0dum 1926, Rosenkrantz 1938) ofMiddle Danian age and were herbivorous and others carnivorous (Fretter & Gra-

NP3 Re- also to nannoplankton zone (Perch-Nielsen 1979). ham, 1978). Emarginula was very common (sponge cently Thomsen (1995) divided the Danish Danian into 9 and detritus eater; Fretter & Graham, 1978).The Ranelli-

calcareous this nannoplankton zones. According to subdivi- dae were carnivorous (Riedel, 1995). Representatives of sion the in Fakse falls into sequence quarry his calcareous the fauna from soft bottom(e.g. Turridae) were very rare.

zones 4 and 5 (Middle Danian A Eulima nannoplankton age). was rather common; this lives parasitic on

echinoids (Muller & Strauch, 1991). Among the bivalves

sessile genera were common, cementing forms like Pycno- Previous work donte and Gryphaeostrea_ or byssate, like representatives of _,t

the Arcidae and Pectinidae (Muller & Strauch, 1991). Li-

Ravn (1902b) recognised no Cancellariidaein the gastro- mopsis. Nuculana and other representatives ofthe infauna

fauna of pod Fakse. He described Tritonium biplicatum, were rare.

it referring to the Cymatiidae ( = Ranellidae) in accordance with at that time. customary placement In his 1933 paper

Ravn revised the faunaand established numerous new spe- Cancellariidaein other Paleocene and Eocene faunas

cies from the so-called “nose-chalk”. This type of lime-

stone is an unconsolidatedcoral limestone in which origi- Cancellariidae are diverse and abundantin the Selandianof

mollusc shells nally aragonitic are preserved due to a trans- Copenhagen (von Koenen, 1885; Ravn, 1939; Schnetler, formationinto calcite (Ravn, 1933; Bernecker & Weidlich, 2001). In the rich mollusc fauna eight cancellariid species

1990). He consideredthree species to belong to the Cancel- have been recorded. From the Danian and Montian of Bel- lariidae:Admete ? biplicata (Ravn, 1902),Admete (Bonel- gium Glibert (1973) mentionedfive species, all established litia) and Uxia These will be discussed in Brian & Cornet (1877) and all referred the sp. sp. species by to genus the systematic part. Unitas Palmer, 1947. Traub (1979) mentioned two rare

Much of the material from the nose-chalk, published by species ofCancellariidaefrom the PaleoceneofHaunsberg,

Ravn (1933) was collected by the late Professor Alfred Austria. The Paleocene of Nuussuaq, West Greenland has

Rosenkrantz, Copenhagen. However, material collectedin yielded a highly diverse cancellariidfauna, consisting of 13 1933 him by was not published by Ravn, because it was species (Kollmann & Peel, 1983). Furthermore they re-

collected later the same year. Rosenkrantz obviously in- ferred one specimen to a new genus, cf. Plesiotriton (p. 67, tended in his later he a publication years as arranged gas- figs 139A-B), indicating placement in Ranellidae. Accord-

from the Paleocene of Fakse, to Beu & tropod species Copenhagen ing Maxwell (1987, p. 23, pi. 29, figs g, h) this and Nuussuaq, West Greenland, and hadartists make draw- specimen should most likely be referred to Turehua Mar-

under his & These 1943. No of ings supervision (Kollman Peel, 1983). wick, representatives any Nuussuaq generaare

hundreds of illustrations housed many prepared are in the present in the Fakse fauna with the possible exception of

Rosenkrantz files in the Museum of the Geological Copenha- juvenile listed herein as Unitas sp. The Eocene of the

gen. Four of the cancellariids treated in this study were Paris Basin contains a rich cancellariidfauna (Cossmann &

drawn those by artists, and Rosenkrantz made a few draw- Pissarro, 1910-13; Pacaud & Le Renard, 1995).The genus

with notes offive established ings accompanying species as Unitas is represented by 25 species and the genus Plesiotri- new in this Unitas anderseni, Plesiotriton steni, ton three list 11 in the paper; by species. They species genus Admetula rosenkrantzi n. Admetula and Bonellitia Jousseaume, 1887 which consider sp., faksensis they a syno- Tatara danica. nym of Admetula, a synonymy not accepted herein. The

Wenz referred Plesiotriton and Semitritonto the from (1938-44) remaining genera the Fakse faunaare not represented.

Beu family Ranelliidae, but & Maxwell (1987) referred The Eocene ofCotentin(North France) contains seven can- both Cancellariidae. genera to Tatara, originally (Fleming, cellariids (Cossmann & Pissarro, 1901), ofwhich six were

included in the also referred referred the 1950) family Ranellidae, was to genus Uxia Jousseaume, 1887and one to the to the Cancellariidaeby Beu & Maxwell (1987). genus Sveltella Cossmann, 1889. Unitas (as Palmer, 1947

is a replacement name for Uxia Jousseaume, 1887) [non

Walker, 1866]. The British Eocene contains seven species

The molluscan fauna referred to the Unitas and 14 Admetula genus (as Uxia) to (as Bonellitia) fide Wrigley 1935.

The mollusc fauna of the bryozoan and coral limestone at -99-

Palaeogeographic conclusions Subfamily PlesiotritoninaeBeu & Maxwell, 1987

Genus PlesiotritonFischer, 1884

Type species Cancellaria volutella Lamarck, 1803 by Of the rare Cancellariidaefrom the Danian of Fakse some original designation. species demonstrate faunal affinities to the New Zealand region. According to Beu & Maxwell (1987) Tatara is known only from the Eocene to Lower Miocene of New Plesiotriton steni n. sp. Zealand and Semitriton from the Eocene ofAustralia and Figures 4, 11 the Eocene of North France (Fresville). Beu & Maxwell

(1987, p. 52) questioned the generic referenceof Semitri-

- Fakse Type locality quarry. ton inopinatus Cossmann & Pissarro, 1905 fromFresville but the characters ofthe Fakse species treatedherein agree Type stratum - Coral Limestone, Middle Danian, Paleo- well with the of Semitriton. Plesiotriton is type a more cene. widespread genus: Cretaceous, Paleocene and Eocene of

USA, Eocene ofthe Paris Basin, Eocene and Oligocene of

Derivationofname - This species is namedafter conserva- Java, Eocene of Algeria, Recent from the western and tor Sten LennartJakobsen, Geological Museumofthe Uni- southwestern Pacific. Unitas has a widespread distribution versity of Copenhagen. in Caenozoic faunas. Beu & Maxwell (1987, p. 14) dis- cussed the distributionofseveral genera of Cancellariidae

Holotype - MGUH 27344 (leg. Sten Lennart Jakobsen, ex and suggested that the known records reflect a very inade- ISL Colin: figure 4). quate knowledge of the fossil record. They found it

that the distribution is caused unlikely pattern by conver-

Paratype - MGUH 27345 (leg. Alice Rasmussen, ex ARF gence of groups, evolving separately in Europe and New Colin: figure 11). Zealand.We agree with theiropinion about our inadequate knowledge of thefossil record and also with theirstatement Additionalmaterial- ARF Colin, 1 juvenile specimen. regarding convergence. The genus Turehua is relatively common in the Eocene and Oligocene of Europe (Hungary, Diagnosis - A slender Plesiotriton with opisthocline axial Germany, Belgium, Denmark) and the Eocene to Pliocene ornament and six primary spirals, of which the medium of New Zealand (Beu & Maxwell 1987). Welle (1997, p. four the are strongest. Anterior canal relatively long. 130) also discussed the relationship between Europe and the Australia- New ZealandRegion and suggested that this

Measurements- The has of holotype a length 15 mm (es- genus was widespread from the Tethys. He presumed the timated) and a width of 5.0 mm. existence ofa large Tethys bioprovince from the European

the region and over Asian to the Australianregion. Records

Description - The shell is rather small, slender and subfusi- of cancellariids from the Cretaceous are few, but the form, height/width ratio almost 3.0. The last whorl equals ofmolluscs from this knowledge period is poor because the half the total shell height, the aperture and canal about 1/3. state of preservation for this class is generally bad due to The protoconch consists of two smooth, convex whorls, the dissolutionof aragonitic shells. which are separated by a deep suture. The small nucleus is

slightly depressed, and the first whorl is planispiral. Imme-

diately before the transition into the teleoconch six spiral Systematic palaeontology cords appear, abapical ones strongest. The transition into

the teleoconchis gradual. The teleoconchof the holotype

consists offive convex whorls separated by a deep suture. Abbreviations- The is defective aperture on the holotype. On the juvenile

the paratype aperture is rounded ovate and rather narrow.

MGUH The collection of the Mu- The labrum is but type Geological broken, seems to have been slightly Copenhagen. seum, thickened. The columellais slightly sinuate with two weak GM material in Registered the Geological Mu- folds. The callus is rather thick and well defined, covering

the seum, Copenhagen. spiral sculpture on the parietal wall. The spiral orna- ISL The collection of K.I. Schnetler, Langa, mentationconsists ofsix primary spiral cords. Of these the

Denmark. four central cords are strongest. Two or three fine secon-

ARF The collection of Alice Rasmussen, Fakse, cords dary spiral soon appear between the primary cords. Denmark. The convex base has numerous weak spiral cords. The ax-

ial sculpture consists of 10-12 slightly opisthocline ribs

which are narrower than their interspaces. Where the cords Class Gastropoda Cuvier, 1797 and ribs intersect, spirally elongate tubercles are formed. Order Wenz, 1938 On the base the axial sculpture gradually weakens. Between CancellarioideaForbes & 1851 Superfamily Hanley, the axial ribs weak collabral growth lines meet the axial Family Cancellariidae Forbes & 1851 Hanley, ribs at an angle ofabout 20°. There are no varices. - 100- - 101 - - 102-

b. Unitas MGUH 27346. 7.5 Figures 2a, anderseni n. sp. Holotype, Height mm.

Unitas MGUH Figures 3a, b. aliceae n. sp. Holotype, 27349. Height 8.8 mm.

4. Figure Plesiotritonsteni n. sp, Holotype, MGUH 27344. Estimated height 15.0 mm.

Figure 5. Semitriton biplicatus (Ravn, 1902). MGUH 27352. Height 9.8 mm.

b. Admetula MGUH Figures 6a, rosenkrantzi n. sp. Holotype, 27351. Height 6.7 mm.

Figures 7a, b. Admetulafaksensis n. sp, Holotype, MGUH 27356. Height 10.3 mm.

8. Figure Semitriton biplicatus (Ravn, 1902). Lectotype, MGUH 108. Height 45 mm.

9. Tatara danica Estimated Silicone rubber Figure n. sp. Paratype, MGUH 27355. height 25 mm. cast.

b. Tatara danica MGUH 27354. 11.0 Figures 10a, n. sp. Holotype, Height mm.

11. Plesiotriton n. MGUH 27345. Figure steni sp. Paratype, Height 4.2 mm.

12. n. Figure Unitas anderseni sp. Paratype, MGUH 27347. Height 2.2 mm.

Unitas MGUH 27350. 3.4 Figure 13. aliceae n. sp. Paratype, Height mm.

Unitas MGUH after Figure 14. aliceae n. sp. Paratype, 3121 ( = Ravn, 1933, pi. 1, figs 12a-b).Reproduced drawingby Betty

Engholm in the Rosenkrantz files in the Geological Museum, Copenhagen, Height 2.4 mm.

Unitas 3192 in the Rosenk- Figures 15a, b. sp. MGUH ( = Ravn, 1933, pi. 7, fig. 7). Reproduced after drawing by Betty Engholm

rantz files in the Geological Museum, Copenhagen. Height 3.5 mm.

16. Unitas 3.5 Figure andersenin. sp. Paratype, MGUH 27348. Height mm.

Figures 17a, b. Semitriton biplicatus (Ravn, 1902),MGUH 27353. Height 2.8 mm.

18. Tatara danica MGUH 27357. 2.3 Figure n. sp, Paratype, Height mm.

b. Tatara danica MGUH 27358. 19a and first Figures 19a, n. sp. Paratype, protoconch teleoconch whorl, height 3.0 mm. 19b apical

view of protoconch, bar 0.5 mm. Silicone rubber cast.

Discussion - Rosenkrantz (unpublished drawing with tor S0ren Bo Andersen, Geologisk Institut, Aarhus Univer-

notes) consideredthe species to be a Plesiotriton, differing sity.

only littlefrom typical representatives ofthe genus. Defini-

tive generic placement of this species is impossible but Holotype - MGUH 27346 (leg. Sprcn Bo Andersen, ex.

Plesiotriton seems most suitable. The columellar plications ISL Colin: figure 2).

of P. steni are not as pronounced as they are in other spe-

cies but the ornamentationand overall make this - MGUH 27347 Alice shape a Paratype (leg. Rasmussen: figure

The MGUH 27348 Bo ISL suitable, although probably temporary, placement. 12): (leg. Spren Andersen, ex.

sculpture is very close to that of P. deshayesianus Beu & Colin: figure 16).

Maxwell, 1987 from the middleEocene of the Paris Basin.

That species also has rather weak columellar folds and also Additionalmaterial- ISL Colin, 2 juvenile specimens (leg.

has the Plesiotriton differs from same type suture. steni Spren Bo Andersen); ARE Colin, 6 juvenile specimens.

other described species of the genus in being elongated as

are most species of Tritonoharpa but the lack of varices Diagnosis - A rather stout Unitas with strong varices at

precludes placement therein. intervals of about one half whorl.

- Measurements The holotype has a length of7.5 mm and a

Subfamily CancellariinaeForbes & Hanley, 1851 widthof 3.9 mm.

Genus UnitasPalmer, 1947 ( = Uxia Jousseaume, 1887, Description - The shell is small, subfusiformand relatively

non Walker, 1866) solid. The length/width ratio is almost 2.0. The body whorl

almost 2/3 of total shell equals the height, the aperture

Type species - Cancellaria costulata Lamarck, 1803, by about 0.4, The four adapical teeth are stronger and more

original designation ofUxia Joussaume, 1887[non Walker, widely spaced than the other. The columella is almost

1866]. straight and has three folds, of which the two adapical are

which is stronger and wider spaced than the abapical one,

placed at the transition to the very short, spout-like canal.

Unitas anderseni n. sp. The aperture is ovate and relatively narrow with a distinct

Figures 2a, b, 12, 16 posterior canal. The thickened labrum has nine teeth inter-

nally and runs gradually into the canal without any demar-

- Fakse cation in a curvation, which under the ca- Type locality quarry. regular projects

nal. The columellar callus is relatively thick and well de-

Type stratum — Coral Limestone, Middle Danian, Paleo- fined. There is no pseudumbilicus. On the parietal wall

cene. there is a rather distinct protuberance midway between the

posterior canal and the adapical columellar fold. The pro-

Derivation ofname - This species is named after conserva- toconch consists of 2 V* convex whorls, which are separated - 103-

by a deeply impressed suture. The small nucleus is slightly

and the first 1 Vi whorls Additional - depressed protoconch are planispi- material Two further juvenile specimens were ral and placed slightly obliquely to the axis, the last whorl mentionedby Ravn (1933: 68).

is quickly increasing in strength. The transition into the

teleoconch is and is indicated the of - A gradual by appearance Diagnosis Unitas with a slightly impressed suture and

theteleoconch sculpture. The holotype has three teleoconch no varices.

whorls, which are mediumconvex and separated by a deep

suture. - The spiral ornament consists offour spiral cords, of Measurements The holotype has a length of 8.6 mm anda

which the abapical one is weakest. The ribs are separated width of 3.5 mm.

by wider interspaces. The axial ribs are slightly prosocline and of almost the same width as their interspaces. On the Description - The shell is small, subfusiform and slender,

body whorl they gradually weaken. At intervals of about almost 2.5 as high as wide.

half x a whorl there are strong varices. The almost invisible The protoconch consists of3 h convex whorls, quickly in-

have the same direction growth-lines as the axial ribs. creasing in diameter, which are separated by a distinct su-

ture. The small nucleus is slightly depressed and the first

Discussion - Rosenkrantz H/z Although (unpublished drawing protoconch whorls are planispiral. From the second

with considered this be it is whorl is notes) species to a Plesiotriton, protoconch a spiral cord present under the adapi-

l properly placed in the Unitas. The Unitas is cal and after further A whorl two genus genus suture, spiral cords ap- from the of represented by numerous species Eocene the pear under it. On the terminal Va whorl a fourth spiral cord

Paris the Eocene ofthe southeasternUnited is Basin, States, present immediately above the abapical suture. The spi-

the Dano - Montian of This has ral cords the Belgium, etc. new species are regularly spaced, adapical one a little

some resemblance to Unitas pearlensis (Meyer & Aldrich stronger than the others. The transition into the teleoconch

in 1887) from the Jackson of Meyer, Eocene the Missis- is gradual and visible as the appearance of small tubercles

Alabama.That has axial sippi Embayment, species stronger on the two adapical spiral cords caused by the intersection ribs and more prominent oblong tubercles on the labrum. ofthe axial ribs. The teleoconchofthe holotype consists of

Cossmann & Pissarro illustrated the Eocene three which (1910-1913) whorls, are mediumconvex and separated by a cancellariidsfrom the Paris Basin. According to Pacaud & slightly impressed suture. Last whorl equals 2/3 ofthe total

Le Renard 25 of Unitas in that shell the about (1995) species are present height, aperture 0.4. The aperture is ovate rich fauna. Of the those, only one resembling the Fakse and relatively narrow with a rather distinct posterior canal.

species is Unitas rhabdota (Bayan, 1873) which has small The labrum is thickened and has nine lirationson the rather

nodes formed by the intersection of the spiral cords and flat and broad internal side. The labrum is almost straight axial ribs and small coronations the shoulder. Gilbert at on the adapical part and meets the canal, which is not de-

revised the of the Dano - Montian of (1973) gastropods marcated from it, in a regular curve. The columella is

None of the of Unitas Belgium. Belgian representatives slightly concave with three folds, ofwhich the two adapical resembles the Danish species fromFakse. are stronger than the abapical fold, which is placed at the

transition into the spout-like canal. The canal is turned to

the left. The parietal callus is well demarcated on the colu-

mella, but relatively thin on the parietal wall. Five spiral Unitas aliceae n. sp. cords, decreasing abapically in strength, can be seen Figures 3a, b, 13, 14 through the parietal wash. The spiral sculpture consists of

four cords which primary are soon accompanied by a fifth

1933 Admete - ? biplicata (RAVN) Ravn, p. 68, pi. 1, figs weaker cord. The spiral ornamentationis relatively weak 12a-b Tritonium Ravn, [partim , non biplicatum

on the teleoconch whorls. The convex base has an addi- 1902b], tional five weak spiral cords, with even weaker secondary

cords inserted. The axial sculpture consists of about 16

Type locality - Fakse quarry. slightly prosocline ribs, slightly wider than their interspaces

and gradually weakening abapically. At the intersections

Type stratum - Coral Limestone, Middle Danian, Paleo- with the two adapical spiral cords weak tubercles are

cene. formed. Between the axial ribs very weak growth lines,

the direction having same as the axial ribs, are visible.

Derivation ofname - This species is named after Mrs. Al-

ice Rasmussen, Fakse. Discussion - This species differs from P. anderseni by

having a larger protoconch with a differentornamentation,

Holotype - MGUH 27349 (leg. Sten Lennart Jakobsen, ex whorls relatively higher without varices, and a deeply im- ISL Colin: figures 3a, b). pressed suture. The outline ofthe labrumdiffers in shape,

P. alicae and having a straighter more flaring aspect. It also

Paratype - MGUH 27350 (leg. Alice Rasmussen, ex ARF has thinnercallus and a stronger sculpture. The body whorl Colin: MGUH illustrated Ravn figure 13); 3121, by (1933, does not slope back regularly to the suture but a small an- pi. 1, figs 12a-b) asAdmete ? biplicata 14. (Ravn): figure shoulder is This gled developed. new species has only a - 104-

few large growth lines, none strong enough to be termed aperture 0.6. Height/width ratio 1.4. The protoconch con- varices. described Ravn (1933) and illustrated a juvenile sists of2Vi smoothand convex whorls, separated by a deep specimen as Admete ? biplicata and mentioned two addi- suture. The nucleus is small, and the firstprotoconch whorl tionaljuvenile specimens in MGUH. We have reproduced is planispiral and oblique to the axis. The last protoconch a drawing of this specimen in the Rosenkrantz files, the whorl is quickly increasing in strength. The terminal V4

Geological Museumof Copenhagen. protoconch whorl has three spiral cords, which increase in

strength. The transition into the teleoconchis gradual. The

holotype has two teleoconchwhorls which are convex with

Unitas flat and sp. a adapical part separated by a deep, almost cana-

Figures 15a, b liculate, suture. The aperture is large and ovate. The labrum

is regularly curved and thickened. Internally the labrumhas

1933 Admete - Ravn, 68, 6, 12a- nine denticles. (Bonellitia) sp. p, pi. figs The columella is slightly concave with two

b. strong oblique folds adapically. A third weak fold is the

ofthe canal. ofthe columella margin Only a part is covered

Discussion - to the and illustration According description by a thin, somewhat indistinctly demarcated callus. Spiral by Ravn the found is not with only specimen nonspecific ornamentationconsists of three primary spiral cords visible the two species described above. In fact it seems to match between the sutures, which continue on the teleoconch

” the figures of what was identified as “cf. Coptostoma by whorls without secondary cords. A fourth spiral cord im- Kollmann & Peel (1983: 94, figs. 212a-b). As specific mediately above the suture is mostly hiddenby the follow- identificationis not and even is possible, generic placement ing whorl. The cords are almost as strong as their inter- problematical, it is here left in Unitas. Unfortunately, locat- On spaces. the convex base there are an additional nine ing the specimen in the Geological Museum, Copenhagen spiral cords, decreasing in strength abapically. Axial sculp- collectionhas not been possible. We have reproduced type ture is absent. On the teleoconch three older varices are the of the in the Rosenkrantz files, the drawings specimen visible as slight thickenings ofthe shell, one occurring im- Museumof Geological Copenhagen. mediately before the aperture and the two others with a

distance ofhalfa whorl. Very weak prosocline growth lines

the are strongest on adapical part of the whorl. Genus Admetula Cossmann, 1889

Discussion - The genus Admetula is well represented in

Type species - Cancellaria evulsa (Solander, 1766) (= Paleogene and Neogene Tethyan faunas and survives into

Buccinum evulsum Solander, 1766), by original designa- the Recent fauna. The type species has prominent varices, a tion. feature not present on all species. Admetula rosenkrantzi

differs from all known congeners in its extreme pyriform

shape as most species are either rounded or elongately Admetularosenkrantzi n. sp. rounded.Its placement in Admetula is questionable due to

Figures 6a, b its overall and also the shape presence ofan almost vertical

columella. As there is no other suitable genus it is de-

- Fakse Type locality quarry. scribed here due to the numberand placement of the colu-

mellar folds. Cancellaria latesulcata Koenen, 1885 (an

stratum - Coral Limestone, Middle Danian, Paleo- Type Admetula) was originally figured illustrating three different cene. shell morphologies, one ofwhich (Koenen, 1885, pi. 1, fig.

2d) is rather low spired and rotund but it is not pyriform. In

Derivation name - This is named after the late of species figuring this species Ravn (1939, pi. 3, figs. 17a-b; as Ad- Professor AlfredRosenkrantz. mete) illustrated only the high-spired form.

Holotype - MGUH 27351 (leg. S.B. Andersen, ex ISL

Colin: figures 6a, b). Admetula faksensis n. sp. Figures 7a, b

Additionalmaterial- GM 1977.1375, one adult specimen.

- Fakse Type locality quarry.

Diagnosis - An Admetula with three spiral cords and no axial sculpture. Type stratum - Coral Limestone, Middle Danian, Paleo-

cene.

Measurements- The holotype has a height of6.7 mm and a width of 4.8 mm. The GM 1977.1375 has a - specimen Derivation ofname faksensis (Latin) = from Fakse. height of7.3 mm and a width of 5.2 mm.

Holotype - MGUH 27356 (GM 1977.1376, leg. A. Ro- Description - The shell is small, ovoid-conical and rather senkrantz 1933: figures 7a, b). solid. The last whorl equals 0.8 of the total shell height, the - 105-

Additional - - material ARF Colin, one adult defective Lectotype (designated herein): MGUH.108 ( = Ravn

specimen. 1902, pi. 2, fig. 11): figure 8.

- An Admetulawith - Diagnosis a spiral ornament consisting Material MGUH 27352 (leg. S.B. Andersen, ex ISL of four low spiral cords, separated by narrow furrows. Colin: figures 5a, b); MGUH 27353 (leg. S.B. Andersen,

Varices occur at an interval ofabout half a whorl. ex ISL Colin: figures 16a, b); ISL Colin, 10 juvenile

specimens (leg. Spren Bo Andersen); APR Colin, 24 juve-

Measurements- The holotype has a height of 10.6 mm and nile specimens.

a widthof 7.3 mm. The additional specimen has a height of

11.1 mm and a width of 7.6 mm. Measurements - The lectotype, illustrated by Ravn (1902,

pi. 2, fig. 11), has a height of45 mm and a widthof 24 mm.

Description - The shell is rather small and ovoid-conical.

The height/width ratio is about 1.5. Last whorl equals 0.8 Description - The adult shell is moderately to medium of the total shell the 0.6. The height, aperture protoconch large and subfusiform. The protoconch consists of 2 Vi

consists ofabout three convex whorls, separated by a deep smooth and convex whorls, separated by a deep suture. The

suture. The nucleus is small, and the whorls are regularly nucleus is small, not depressed, and the first whorl is

increasing in diameter. The transition into the teleoconch is planispiral. On the terminal V* whorl seven or eight spiral

The has and half teleo- gradual. largest specimen two a cords are present. Transitioninto the teleoconch is gradual.

conch which and whorls, are convex separated by a deep Largest specimen has four teleoconch whorls, which are

There is subsutural The is wideand suture. a ramp. aperture convex and separated by a distinct suture. Height/width ovate, going into the canal without constriction. The la- ratio about 2.0. Last whorl equals about0.75 of the total brum has a projecting margin, which is thin on the holo- shell height, aperture and canal about 0.5. Aperture rather

On the other known there three close-set wide and labrum with den- type. specimen are pyriform, thickened, numerous

margins, resulting in a more thickened labrum, which has ticles internally. On the type, illustrated by Ravn there are

four tubercles internally. The columellais rather short and about 30 denticles. Columellaalmost straight, with two almost straight and bears three folds, of which the adapical oblique folds. Canalrather short and turnedto the left. The

is less than the others. The weak fold is columella callus is oblique abapical thin and not very well demarcated. The

the margin of the canal, which is short and wide, slightly convex body whorl is constricted near the base. The spiral

turned the left. The callus is to thin, but well defined, cov- ornamentation consists of seven or eight primary cords

the columella.On the additional there ering only specimen separated by narrower interspaces. The abapical spiral cord

tubercles the wall. The is are two on parietal spiral ornament more or less covered by the following whorl. On the

consists of four low which whorls ribs The base broad, cords, are separated by younger secondary occur. and the

much narrower interspaces. On the convex base there are neck of the canal have numerous ribs. The axial ornament

10further cords, decreasing in strength abapically. There is consists of about 30 poorly definedand slightly prosocline

axial but lines no sculpture, slightly prosocline growth can ribs that gradually weaken on the body whorl. Old aper- be observed. Varices occur at an interval of about half a tures varices, intervals of appear as usually at about one

whorl. half whorl.

Discussion - Rosenkrantz (unpublished notes) considered Discussion - This species differs from Admetula rosen- this be but he krantzi, described above, by having flat, narrowly spaced species to a Semitriton, obviously thought a

new might be needed. The species matches the de- spiral cords and prominent varices. Furthermore, the spe- genus scriptions and illustrationsof Semitritonin Beu & Maxwell cies is more elongated. This species matches Admetulawell (1987) for the multispiral protoconch. The Semitri- by its columella, varices and the rounded body whorl. except illustrated ton protoconchs by Beu & Maxwell (1987) are

paucispiral with a bulbous nucleus. This character thus

be of value. Ravn Genus SemitritonCossmann, 1903 seems to only specific (1902b) desig- nated no holotype, illustrating two internal moulds (1902b,

pi. 2, figures 12, 13) and a wax duplicate of an external Type species - Plesiotriton dennanti Tate, 1898, by origi- impression (plate 2, figure 11). As this last specimen illus- nal designation. trates all characteristic features ofthe species except for the

protoconch, it is here designated as lectotype. Ravn (1933,

68; pi. 1, figs 12a-b) referred three juvenile specimens Semitriton biplicatus (Ravn, 1902b) p. from the nose-chalk to this species. Those specimens, Figures 5, 8, 17a, b have four however, only primary spiral cords and are

juvenile specimens of Unitas aliceae n. described 1902b Tritonium - 228 biplicatum (M.U.H.) n. sp. Ravn, p, sp., herein. This differs from other known of (24); pi.2, figs 11-13. species species

1933 Admete (?) biplicata (Ravn) - Ravn, p. 68; pi. I. figs Semitriton by its multispiral protoconch and its larger size.

12a-b. [partim non 1, figs 12a—b = Unitas It is rather common in the collections of the , pi. Geological

aliceaea n. sp.]. Museum, Copenhagen, but most specimens therein are in- - 106-

temal moulds. Ravn mentioned35 (1933) specimens. strong oblique plaits and up to six (in most cases four)

weaker nodules, decreasing anteriorly in size. The heavy Genus Tatara 1950 Fleming, but narrow callus covers the columella, and on the adult

specimen it has a raised and well defined edge, forming a

- Marshall & Murdoch, Type species Cymatium pahiense collararound the aperture. Spiral ornament consists offive

1921, by original designation. cords that continuationsof spiral are the spirals on the pro-

toconch. The spirals are considerably weaker than their

interspaces. The adapical spiral cord soon disappears, and

Tatara danica nov. sp. the abapical one is partly covered by the suture. The three 9, 10a, b, 18, 19a, b. Figures medium strong spirals cords areregularly distributedon the

whorl and are much weaker than their slightly concave

- Fakse Type locality quarry. interspaces. From the first teleoconch whorl and onwards

fine the secondary spirals appear, seven ones on largest

stratum - Coral Limestone, Middle Danian, Paleo- Type specimen. The axial ornamentation consists of about 20

cene. prosocline ribs on the first teleoconchwhorl, increasing in

number to 30 on the adult specimen. The ribs are only

= Derivation ofname - danicus (Latin) Danish. slightly wider than their interspaces. At the intersectionsof

the spiral and axial sculpture small pointed tubercles are

Holotype - MGUH 27354(leg. Spren Bo Andersen, ex ISL formed. The concave interspaces between the spiral cords Colin: 10a, b). figures are finely cancellatedby the fine secondary cords and the

incremental growth lines. Old apertures are visible as

- MGUH 27355 Spren Bo Andersen, ex Datatypes (leg. varices, with intervals of about half to two thirds of a ISL Colin: figure 9); MGUH 27357 (figure 18); MGUH whorl. The growth lines are parallel to the axial ribs.

27358 (leg. Spren Bo Andersen, ex ISL Colin: figures 19a, b). Discussion - Juvenilespecimens lack the raised columellar

shield and are rather similar to Semitriton biplicatus, but

Additionalmaterial- 17 specimens, juvenile, from mostly differ distinctly by their characteristic sculpture. S. biplica- the “nose-chalk” and two adult specimens from the coral tus has a higher numberoffine, narrowly spaced spirals of limestone (leg. Spren Bo Andersen, ISL Colin); ARF almost equal strength. Rosenkrantz (unpublished notes)

Colin, one adult and two from the juvenile specimens considered the species to be a Sassia. In fact, juvenile

“nose-chalk” and one adult specimen from the coral lime- specimens are rather difficult to separate fromjuvenile rep- stone; MGUH Colin(GM 1977.1373and 1977.1374), two resentatives of Sassia as both genera have strong varices specimens from the “nose-chalk”. and rather similar sculpture. Sassia is represented by sev-

eral species inthe Fakse fauna, of which only Sassia faxen-

- A slender Tatara with two columellar Diagnosis strong sis (Ravn, 1933) has been established. This new species plaits and to six columellar nodules. The of the up edge seems to be related to T. revoluta (Finlay, 1924) as figured

outer lip is moderately wide and bears nine denticles that by Beu & Maxwell (1987, p. 54; fig. 2N, pi. 27 j, k). It are elongate abaxially. differs by having only two columellar plaits and a less wide

edge on the thickened labrum. Also, the tubercles on the

Measurements- The holotype has a height of 11 mm and a parietal wall are more distinct. Tatara flemingi Beu & widthof 5.5 mm. Largest specimen has an estimated height Maxwell, 1987 (p. 53, pi. 27, figs a-e) is shorter and wider, of about 25 mm. lacks varices on most of the spire and has a sinuous aper-

ture and outer lip. Furthermore the spiral ornament is much

Description - The shell is rather small to moderate large and the number of the is finer, ridges on outer lip higher. and ratio about 2. Last whorl subfusiform, height/width Tatara pahiensis (Marshall & Murdoch, 1921), has four equals about 0.8 of the total shell height, about aperture columellar plaits and is larger and wider (see Beu & Max- 0.6. well, 1987, p. 53, pi. 27, figs f, g). It has coarse cancellate 3 Protoconch consisting of 2 A convex and smooth whorls and wide with sculpture a very outer lip 11 coarse trans-

a suture. Nucleus is small and separated by deep slightly verse ridges. Tatara transenna (Suter, 1917) is short and depressed. The first protoconch whorl is planispiral and inflated, also with very coarse cancellate sculpture. obliquely placed to the axis ofthe shell. Terminalhalfpro- toconch whorl with five spiral cords. Transition into the teleoconch is The has three gradual. largest specimen Incerta sedis teleoconch whorls, which are convex and separated by a

deep suture. The is elongate-ovate and relatively aperture 1933 Uxia - Ravn, 68, 7a-b. sp. p. pi. 7, fig. narrow. The labrum is thickened and has a flat internal

Discussion - to the and illustrations side, bearing nine denticles, which are elongate abaxially. According description

Ravn the has no orna- Up to six nodules are visible on the parietal wall. These by specimen (MGUH 3199) spiral

mentation, but close-set axial ribs on the first teleoconch nodules are oblique to the spirals. The columellabears two - 107-

whorl. The drawings of the specimen in the Rosenkrantz Cossmann, M. & Pissarro, G. 1910-13. Iconographie complete

files show about 10 fine des de I'Eocene des spirals on the neck ofthe canaland coquilles fossiles environs de Paris. Pa-

weaker ris. 2 volumes. some more spirals suggested on the base. On the Desor, E. 1847. Sur le terrain danien, nouvel etage de la craie. terminal part of the body whorl there are no axial ribs. The Bulletin de la Societe Geologique de France 2(4), 179-182. columellahas three strong oblique folds, and the short ca- Finlay, H.J. 1924. New shells from New Zealand Tertiary beds. nal is turned to the left. Transactions and Proceedings of the New Zealand Institute The general outline and ornament suggests that the speci- 61,259-270. men might be a juvenile Mitridae. Fischer, P. 1884. Manuel de conchyliologie et de paleontologie

conchyliologique, ou histoire naturelle des mollusques vi-

vants et fossiles. Paris, LibrairieF. xxiv + Savy, 1369pp., 23

Acknowledgements pis [Fasc. 7, pp. 609-688, 1884].

Fleming, C.A. 1950, The molluscan fauna ofthePahl greensands,

We are greatly indebtedto Alice Rasmussen, Fakse, Spren North Auckland. Transactions of the Royal Society ofNew

Bo Andersen, Arhus and Sten Lennart Jakobsen, Copenha- Zealand 78, 236-250, pi. 24.

for donationof Forbes, E. & Hanley, S. 1850-1851. A history of British Mol- gen material. Sten Lennart Jakobsen is fur- lusca and their shells 3, 1-320 (1850), 321-616 (1851). thermorethanked for valuablehelp in locating thematerial London (J. van Voorst). in the Geological Museum, Copenhagen and for photo- Fretter, V. & Graham, A. 1978. The Prosobranch Molluscs of graphic work. Spren Bo Andersen, Geological Institute, Britain and Denmark. Journal ofMolluscan Studies, Supple- University of Arhus helped with photographic work, and ment 1, 1-37. Erik Thomsen from the same Institute made the SEM pho- Gilbert, M. 1973. Revision des Gastropodes du Danien et du tos and drew text-fig. 1. Keld Gregersen, Sorp the gave Montien de la Belgique. Institut Royal des Sciences senior author assistance with Naturelles Memoire 116 photographic equipment. de Belgique. 173, pp. allowed Betty Engholm, Sorp kindly us to reproduce her Jousseaume,F.P. 1887. La famillede Cancellariidae(Mollusques

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