Contr. Tert. Quatern. Geol. 29(3-4) 73-137 3 tabs, 9 pis. Leiden, February 1993

Danian and Spirorbidae from NE Belgium

and SE Netherlands: K/T boundary extinction,

survival, and origination patterns

ManfredJäger

Rohrbach Zement

Dotternhausen, Germany

SE Netherlands: K/T extinction, survival, and Jager, Manfred. Danian Serpulidae and Spirorbidae from NE Belgium and boundary

— Contr. Tert. 3 tabs, 9 pis. Leiden, February 1993. origination patterns. Quatern. Geol., 29(3-4): 73-137,

the Palaeocene Geulhem Member Formation) Twenty-two serpulid and spirorbid species and subspecies from early (Danian) (Houthem

illustrated, discussed in detail and to the following Cycloserpula, as exposed in the type area of the Maastrichtian Stage are assigned genera:

Rotularia, Metavermilia ?, Placostegus, Neomicrorbis, Bipygmaeus, Pileolaria ? and Neodexiospira ?. Three Serpula?, Cementula, Ditrupa, Pyrgopolon,

Rotularia tobar and R. (R.) tobar vroenhovensis, while additional new new subspecies are erected, viz. Neomicrorbis parietalis jagti, (R.) gracilis

Within the Geulhem Member there but moderate of described in nomenclature as are known. are species are necessity open they poorly

changes in the serpulid/spirorbid faunas, which result mainly from differences between hard and soft substrates. Of twenty-two

recorded from the classic Danian ofDenmark, which corroborates the correlation ofthe Geulhem (sub)species sevenhave previously been

Member with the while two show differences at the subspecies level. Although data are at present insufficient to type Danian, slight middle and determine precisely the position ofthe Geulhem Member within the type Danian succession, faunal relationships suggest a

Danian for this member. A of Danian faunas with Maastrichtian late Danian rather than an early age comparison serpulid/spirorbid

world shows that the former in fact a slightly impoverished ones from the Maastricht area as well as from other parts of the represent

the K/T these either shallow and Thirteen and became extinct at or near boundary: were fauna ofMaastrichtian type. genera subgenera

substrate, or and few Most warm water forms which for fixation needed a large genera subgeneracomprising only species. (sub)genera

and above the crossed the K/T boundary, the majority (seventeen to twenty-four) with different species below boundary, some (eight) the with identical few and the Danian/Palaeocene. species. Comparatively (seven) genera subgenera originated during Generally,

which the are similar those described for other benthic suspension/detritus feeders, on extinction, survival and origination patterns to

K/T boundary event had only a limited influence.

— NE SE Netherlands, new Key words Serpulidae, Spirorbidae, Maastrichtian, Danian, K/T boundary, Belgium, taxa.

Dr M. Jager, Rohrbach Zement, D(W)-7466 Dotternhausen, Germany.

Collins added few others. The two Contents (1988, fig. 1) a

most important localities in the environs of Introduction 73 p. Maastricht the former Curfs Geulhem are quarry at Systematic descriptions p. 74 (now Ankersmit Holding BV) and the temporary Discussion 110 p. Canal Vroenhoven- sections along the Albert near 110 Comparisons p. Riemst between the Vroenhoven and Veldwezelt K/T extinction, survival and boundary origi- The section the Ankersmit bridges. at quarry was nation events p. 112 designated stratotype of the Geulhem Member 115 Acknowledgements p. (Houthem Formation) by Felder (1975); this will References 115 p. presumably remain well exposed through future

quarrying. However, only eight metres of Danian Introduction in strata are exposed there, this contrast to c 12

Localities — Meijer (1959) described four sections metres in the Albert Canal sections. Van der Ham

in the and der Ham al. exposing early Palaeocene (Danian) strata (1988) van et (1987) discussed the

of the Maastrichtian while & of this difference in thickness with ref- type area Stage, Jagt implications 74

the distinctive echinoid faunas result the that erence to highly from fact more samples from the

the recently collected from uppermost part of the latter locality were studied.

Geulhem Member along the Albert Canal. During

1985 and 1986 there were ample opportunities for

Number of Albert Canal Geulhem extensive collecting, but, unfortunately, the various specimens small could be outcrops not combined to a standard > 200 extremely frequent extremely frequent section. if Most, not all, outcrops are no longer 101-200 very frequent extremely frequent accessible. 51-100 frequent very frequent

Most studied herein come from the samples 21-50 common frequent

Albert Canal those of 11-20 sections, e.g. Meijer (1959), not rare common

6-10 now incorporated into the collections of the Geolo- rare not rare 3-5 very rare rare gisch-Palaontologisches Institut und Museum der 2 extremely rare very rare Universitat in addition to and Hamburg, samples 1 extremely rare extremely rare specimens collected by E. Voigt, J.W.M. Jagt and

M. Jager. Only few of the samples studied were

collected from the Geulhem section by M. Meijer SYSTEMATIC DESCRIPTIONS and E. Voigt.

Stratigraphy — Meijer (1959) subdivided the sections Family Serpulidae Rafinesque, 1815 exposed at Geulhem and along the Albert Canal Subfamily Filograninae Rioja, 1923

lower unit Genus 1956 into two units, a comprising more Cycloserpula Parsch, (= coarsely-grained, glauconitic biocalcarenite with Glomerula Regenhardt, 1961 sensu spines of the echinoid Tylocidaris hardouini (Desor, Briinnich Nielsen, 1931; Omasaria

1855) and few specimens of the inarticulate bra- Regenhardt, 1961) chiopod Danocrania geulhemensis (Kruytzer & Meijer,

and unit of less — Goldfuss, 1831. 1958), an upper consisting coarse- Type species Serpula flaccida — sediment with of the Remarks A more detailed grained many specimens cra- study (in comparison with of niid brachiopod and few spines of the echinoid T. Jager, 1983) and early

has shown doubt that bruennichi Ravn, 1928. The reader is referred to Jagt serpulids, beyond the late

& Collins and is (1988) Jagt & Janssen (1988) for a Bajocian (Dogger) species, flaccida Goldfuss, 1831

and more detailed discussion. congeneric, very probably even conspecific

— 1820. Correlation with the type Danian Tylocidaris bruennichi with, gordialis von Schlotheim, Parsch (1956)

of is an important key index fossil for the middle Dan- designated flaccida type species Serpula (Cycloser- ian in above while of Denmark; as mentioned this species pula), gordialis was designated type species

in the unit of the Geulhem A Glomerula Briinnich occurs upper Member. by Regenhardt (1961).

of this member with the Nielsen had the precise correlation type (1931) previously suggested name Glomerula Danian succession is currently impossible (see dis- but had failed to designate a type species, cussion injagt & Collins, 1988). which is why Regenhardt's designation is valid.

— Glomerula is of Synonymies The taxa, Cycloserpula lombricus, Consequently, a junior synonym

Cycloserpula. In of the fact that Glomerula was Pyrgopolon (Pyrgopolon) mosae mosae, P. (P.) m. ciplyana spite well also and defined Briinnich Nielsen, and that the com- and Bipygmaeus pygmaeus occur are com- by with in the late Cretaceous bination the name renders a clear moner underlying strata. specific pic- of the characteristic of the These species and subspecies will be described in ture shape tube, Cycloser-

was used artificial omnium for detail(Jager, in prep.) in the Illustrated Catalogue of late pula as an gatherum all kinds of circular tubes distinct Cretaceous fossils from Limburg (The Netherlands) and lacking longitudi-

To nal adjacent areas (J.W.M. Jagt, editor). avoid repeti- sculpture.

here and — tion, synonymies are kept to a minimum, Diagnosis Tube solitary, or a number of inter-

is made contributions in and tubes In reference to now press twined forming clusters. clusters, tubes nor- in preparation. mally not parallel to each other; only occasionally

— Below is tabulated the of the bundles be less Frequency frequency may more or parallel. various in the sections studied. Differen- Tube in species two very long, very slowly increasing diame-

curved with anterior tube ces between the Geulhem and Albert Canal sections ter, strongly, parts mostly 75

for- the does in central growing thigmotaxically along posterior parts, not, genus not appear Europe

three- before the mid-Late Toarcian. It is absent ming two-dimensional meanders, two- or probably and dimensional charac- from the bifrons Zone of Somerset (England), irregular spirals or, highly from the mid-Late Toarcian teristically, three-dimensional chaotic knots. Very with certainty jurense

in the limestone nodule in SW Ger- often the substrate is a foreign body only layer (sensu stricto)

tube the At another late Toarcian nodule level, posterior part, whereas in the anterior part many. only

older tube used substrate. At the of one foot above the level, sud- parts are as start jurense Cycloserpula and it remains of tube growth and between the strongly curved tube denly becomes fairly common one with there be curved the commonest until Danian time, a parts may straight or only slightly serpulids Bathonian. which not become erect decline the segments, may or may slight during of i.e. tube above the substrate. The dominance straight, non-coiled,

' life in results from the common Breakage of tube during frequently occurs: a parts ;'flaccida probably mollus- tube then from the of the occurrence and comparatively large size of new part grows cavity

first shells in the late of SW so irregularly broken tube end, the new tube at can Bajocian Germany,

lesser that the tubes could across a wide substrate having a slightly (rarely strongly) diameter grow of curved than the old area: is herein considered but a variety and often being more strongly ’flaccida’ tube. gordialis.

the tube which often resemble a disor- There is no regular ornament on exte- Jurassic specimens,

of former weak rolled have a more constant rior, except for lines breakage, derly up rope, slightly

in in few than do Cretaceous and Danian individ- growth lines some specimens, and, a curvature

in which consecutive and Maastrichtian and Danian specimens, peculiar, uals, straight narrowly

curved often occur. the of occasionally perforate, rucksack-like protuberances. parts However, range exceeds this differ- Tube wall consisting of cylinder layer only, parable variation within a single sample

attachedtube the is in layer absent. The base of parts is thus ence; curvature probably dependent part

this weak attachment the environment. not broadened by a border, on deformation The of features increases being somewhat strengthened by basal array special through

the column and reaches dur- of the circular cross-section, so that the tube clings stratigraphic an acme

cemented it. the Maastrichtian and Danian. The closely to the substrate, but is not to ing peculiar

Even in diagenetically relatively strongly altered 'trilobate narrowing of the tube's lumen' mentioned

tube often be loosened (1983, for material from the early sediments, parts may easily by Jager p. 25)

from the substrate without damaging the tube wall. Cenomanian onwards, has lately been observed also

in of middle Green- In contrast to the virtual absence of external specimens Aptian age (Lower

the from the Isle of ornament and generally simple tube structure, sand) Wight (material presented by

Prof. Dr It thus cross-section and the tube interior display an enor- Herwig Wulf). appears appropriate

the most to subdivide the material of mous array of special features, important long-ranged Cycloserpula is the 'trilobate of the tube's into a and a more advanced species. It perhaps being narrowing primitive

unknown in where to fix the lumen'. Most of these features are any not quite clear, however, boundary

for there is few other serpulid species. It should be stressed, how- between these, only relatively mate-

found in limited number of rial of Cretaceous At least, all ever, that they are only a early Early age. Juras- material in and even do not occur at sic to and the Tithonian) specimens any sample (up including the all in some populations or strata. represents primitive species, gordialis (von

— and Comparison In clusters of Filograna Oken, 1815 the Schlotheim, 1820) sensu stricto, specimens

tubes often less and of the from at least the middle onwards are more or parallel, most occurring Aptian

the the advanced for which the special features of Cycloserpula, 'trilobate narrow- represent more species,

lumen' in name lombricus Defrance, 1827b is here ing of the tube's particular, are absent. provi-

— Sinemurian mid-Late Toar- used. Stratigraphic range ?, sionally The social form, de C. is cian- (middle Lutetian), Oligocene ?, post- plexus (J. Sowerby, 1829) considered be but of lombricus, Oligocene ?. now to a subspecies

Remarks — Sinemurian specimens from Sicily (Len- while in gordialis there is no discrimination between

tini, 1974, pi. 12, fig. 2; pi. 16, fig. 10) and a single solitary and social forms.

early Sinemurian specimen from Wellendingen (SW Very probably, Serpulites contorquatus von Schlot-

before to If heim, 1820 belongs to as well, but from Germany) me may belong Cycloserpula. Cycloserpula 76

and illustrated medium-sized the original description (without illustrations no (1972, fig. 2g) a specimen

it be types designated) cannot determined whetherit of ’Glomerula’ sp. from the New Zealand Oligocene,

the the social form. In the and recorded similar tubes from the late represents solitary or (p. 750)

fur Naturkunde of the Miocene and Holocene. As collections of the Museum Eocene, early even long

I unable the tube of Schmidt's and Humboldt-Universitat Berlin, was to trace as structure Fleming's

of the from Aachen and from the material remains of those any ten specimens unknown, interpretation

is difficult. English Chalk mentioned by von Schlotheim nor species

other labelled I did find any specimen contorquatus. a single specimen of the solitary subspecies from the Cycloserpula lombricus (Defrance, 1827b)

English Chalk, without a species name on the PI. 1, Figs 1-20

label. Whether or not von Schlot- accompanying ? 1820 96. Serpulites contorquatus von Schlotheim, p. heim this for the used original description 1827b ? lombricus 571. specimen Serpula Defrance, p.

is unknown. Since there seems to be no material of

lot be determined whether Defrance did the type left, it cannot — (1827b, p. 571) not provide an

of lombricus did he contorquatus is a senior synonym or plexus. illustration, nor designate a type specimen,

and the is brief It cannot be ruled out totally, that, contrary to description very ('Ses tuyaux, qui

' Jager's view (1983), Sarcinella’ sarcinella Regenhardt, n'ont qu'une demi-ligne de diametre, sont lisses et

1961 is referable but in fact entortilles Localities mentioned not to Filograna, repre- sur eux-memes

like of the 'craie Beauvais Meudon'. The Craie sents an atypical Filograna- variety Cycloserpula are a et a

lombricus This be solved de Meudon is of late Hebert plexus. problem can only Campanian age. (1855)

of features of tube stated that lombricus and through an examination special gordialis are conspecific,

in additional material from the which that lombricus is structure type means very probably a genu-

locality of this species. ine species of Cycloserpula.

— For full discussion of varieties and In the Maastricht-Aachen-Liege area, specimens Description a

from the Geulhem Member display nearly exactly special features the reader is referred to Jager (1983,

the of features in material and same array special as seen pp. 24, 25, 29-31) Jager (in prep.).

from the underlying Meerssen Member of the In the Geulhem Member of the Maastricht-

the tube diameter is Maastricht Formation (late Maastrichtian). Thus, Aachen-Liege area on average

at least 0.7-1.3 with maximum of 1.8 A contin- the Danian and Maastrichtian specimens, mm, a mm.

the tube this in contrast from this area, undoubtedly belong not only to uous range of sizes is observed,

but also the the white chalk facies of and few other localities same species, to same (unnamed) vari- to

the in northern and also ety. Cycloserpula crosses K/T boundary mor- Germany (Jager, 1983, p. 25)

phologically unchanged. to the chalky facies of the Gulpen Formation of the

Rovereto the where two size According to (1904a, p. 12) only Maastricht-Aachen-Liege area,

difference between be eximplicata (Rovereto, 1904a) groups may distinguished.

Most from the Geulhem Member (nom. nov. pro gordialis sensu Leymerie, 1846a, p. specimens are

from attached substrate 369, pi. 13, fig. 16) and Cycloserpula specimens to a (calcareous algal crusts,

is the small the Maastrichtian of Riigen (NE Germany) bryozoans etc., rarely algal filaments). Often a

smaller size of the former. Doncieux substrate is as an on (1926, p. 19, negative preserved impression

pi. 2, figs 26-29), who provided relatively good pho- the underside of the tube. Typically in high-energy

tographs of eximplicata, recorded this species from biotopes, the tube first forms meanders (PI. 1, Fig. 1)

the Ypresian to the middle Lutetian. However, then three-dimensional chaotic knots (= 'Rohren-

before from around the verlaufa' of 20a- material me Palaeocene/ Jager, 1983, p. 29) (PI. 1, Figs 2a,

Eocene boundary of the Tremp Basin (NE Spain) c), but these forms could be studied in a few speci-

presented by Drs R.H.B. Fraaye, cannot be distin- mens only, since most tubes were fragmentary.

is curved tube guished from lombricus. Thus, eximplicata herein Straight or slightly parts, which,

considered be with lombricus. because of the absence of substrate on to synonymous impressions

above the sub- During the Eocene and post-Eocene other Cyclo- the tube's circumference, grew erect

much less but do serpula-shaped tubes apparently do not grow to the strate, are common, occur (PL 1,

The Eocene with maximum of 16 In the size ofjurassic and Cretaceous species. Figs 3-5), a length mm.

1955b white chalk and northern ’Serpula’ maeandrica Schmidt, closely resem- chalky facies of Germany bles small meanders of these erect tube are often broken at the very Cycloserpula. Fleming parts top 77

successive tube Oxfordian of Dotternhausen before and, following a sharp curve, parts (SW Germany)

the erect less me. enwrap part more or regularly

b' This varia- Series of at the ('Rohrenverlauf ofjager, 1983, p. 29). oblique step-shaped swellings do in the Geulhem Mem- tube's interior occur tion is rare in the materialfrom the (Jager, 1983, p. 30)

this Geulhem Member 1, 1, lower of ber. In a single specimen envelopment does (PI. Fig. part pho-

is rather but less common than in chalk facies. occur, but irregularly developed (PI. 1, Fig. tograph), are

— inter- the Discussion The is a 6). In another fragmentary specimen, younger following provisional of the features in the tube tube is only slightly curved and is directed in the pretation special seen

third of the In with the broken In a structure a same way as part (PI. 1, Fig. 5). present species: Serpulinae

in knot: the of the tube specimen, tube breakage occurred a well-developed parable layer, thickening from addition of material tube in a direction differentfrom the wall results mainly an new part grew

broken old tube (PI. 1, Fig. 2a, b). onto the tube's exterior. In Cycloserpula, however, in

less which there is tube wall Thetube circumference is sometimes more or no parable layer, thickening

less result from material to the tube's regularly circular, but at times more or can only adding

at the side interior. Thus, the unknown soft of deformed. Especially convex of curved parts Cycloserpula

of the should have been extended to the tube parts, rucksack-shaped protuberances anteriorly give

mentioned wider and had tube's exterior occasionally occur (not prolonged tube parts a diameter, a

either still closed leave for byjager, 1983), which are (PI. 1, gradual taper posteriorly to enough room a

broken of the tube wall. In the modern Figs 7, 8) or open (PI. 1, Figs 9, 10). thickening species,

The thickness of the tube wall is extremely vari- Filograna implexa Berkeley, 1828, however, tube

also added the inner but the able. Sometimes it is rather thin (PI. 1, Figs 4, 11), material is on surface,

soft but specimens in which the lumen is minimised by parts taper only slightly (see Faulkner, 1930, fig.

thick tube wall In an extremely are not very rare. 3a).

order thick-walled specimens the cylinder layer of the tube Moreover, in to produce an occasionally

six thick tube wall in with may consist of up to (and perhaps even more) very a species rapid longi-

which tudinal the area of of tube layers (PI. 1, Fig. 12a) are developed concen- growth, aggregation If material should have been fairly long. one trically or, more frequently, eccentrically, leaving accepts

in the of of the soft circular or crescentic interstices, as seen cross- hypothesis a gradual taper parts,

the of several tube wall section (PI. 1, Figs 12a, 13, 16, 19). common occurrence layers

The cross-section of the innermost tube layers is sheathed into one another concentrically or eccen-

but cross-sec- with crescentic lumina between (as seen in mostly circular or deformed circular, trically

lumen' tions with 'trilobate narrowing of the tube's cross-section) are easily explained by a rapid move

all An forward the of the (PI. 1, Figs 14-16) are not at rare. 8-shaped of soft parts during prolongation

anterior end. At knot fragment shows trilobate narrowing at both tube's a certain position of the

tube's the soft became ends (PI. 1, Fig. 14), but not in between, as seen in length, parts suddenly nar-

Trilobate and thus the diameter of the new inner tube areas where the tube wall is broken. nar- rower,

diminished rowing is brought about by three gradually increas- wall layer also abruptly, leaving a cavity

between the tube wall ing and decreasing longitudinal swellings of up to layers.

about 3 in in Sometimes the innermost tube wall leaves mm length (several cm long an layer little for the soft The abdo- extraordinary early Cenomanian specimen before only very room parts.

whereas lumen should therefore have into thin me), a kidney-shaped (not men- men tapered a very

of there where the of the innermost tioned by Jager, 1983) is the result a single longi- 'tail', aggregation

Such thin tudinal swelling (PI. 1, Figs 12b, 17-19). tube wall layer took place. a tail, however,

the small inner tube does not in F. The corkscrew of Occasionally, layer may occur implexa. shape

within the lumen of the outer the inner tube wall in some grow trochospirally layer specimens requires

in number of but this does in F. tube layer. Distinct corkscrews occur a a corkscrew-shaped tail, not occur either. specimens from the Geulhem Member (PI. 1, Fig. implexa To the 'trilobate of the tube's 20a-c), but the most instructive corkscrews have explain narrowing

been collected from the Meerssen Member (see lumen' the soft body's posterior end ought to have

in The of been trilobate but in all individuals and Jager, prep.). same type corkscrew struc- shaped, not

in but and indeter- Such ture also occurs a unique, worn only during some (repeated) growth stages. a

minate tube (not assignable to Cycloserpula) from the repeated short-time change of abdominal morphol- 78

of sexual Two of the three described below are ogy is usually related to phases activity. species

of sexual to albeit with in view of However, generally swelling organs assigned Serpula, a query,

the 'trilobate of the the fact that show similarities to certain occurs, whereas narrowing they some

of Recent tube's lumen' requires a periodic contraction the (but not all) species of this or of closely

of the third abdomen. In F. implexa, 'posteriorly the body termi- related genera. The generic assignment

- is uncertain. nates in two lobes the anal papillae' (Faulkner, species very

the 'trilobate of As the tubes are of so little value in identification, 1930, p. 113). Perhaps narrowing

well the corkscrew the tube's lumen' as as shape of palaeontology cannot contribute anything to a the inner tube had in the reconstruction of the of the wall a special function phylogeny genus Serpula. high-energy biotopes inhabited by Cycloserpula to Ten Hove (1984) and ten Hove & Jansen-Jacobs ensure a more effective anchorage for the soft parts. (1984) pointed out the relationships between Recent

In F. 'in the tube is notched of and other Recent implexa, some cases species Serpula serpulid genera

features of the and of the dorsally, and when this is so the faeces are even by referring to soft parts

removed' more effectively (Faulkner, 1930, p. 118). opercula.

Such is unknown in a notch Cycloserpula.

The rucksack-shaped protuberances resemble the Serpula ? insiticia (Regenhardt, 1961) brood chambers of Recent Chitinopoma arndti PI. 2, Figs 1-13 Zibrowius, 1983 and C. serrula (Stimpson, 1854). In

these 1961 insiticia 54, Cycloserpula lombricus, however, structures often v Proliserpula (Proliserpula) Regenhardt, p. pi.

3. be 6, fig. are too irregularly developed for them to true — ascendens 9. ? 1965 Spirorbis Rasmussen, p. 35, pi. 8, fig. brood chambers (ten Hove, in litt., February 1990).

— Comparison Cycloserpula gordialis shows a more reg-

-— tube Type Regenhardt described a long fragment ular curvature and fewer of the above-mentioned in detail, but did not illustrate the specimen; it is in there is special features, particular, no 'trilobate here illustrated in PI. 2, Fig. 8a, b. On microfossil narrowing of the tube's lumen.' cell no. which contains three he — 739, specimens, Geographic and stratigraphic distribution Albert Canal wrote 'Holotypus/groBes Stiick/Nr. 739'. In the sections, lower part of the Geulhem Member: legend of the plates, however, he designated the left- extremely frequent, upper part, extremely frequent. hand specimen of his plate 6, fig. 3 (the shortest of Geulhem lower of the Geulhem Mem- section, part the three) holotype. This designation is the valid ber: extremely frequent, upper part, very frequent. here in 9a- one. The holotype (illustrated PI. 2, Fig. tube's There are no striking differences in the shape is curved c) a slightly fragment of an erect tube, 2.5 in of of or size or the frequency occurrence special with 1.3 Tube mm long, two peristomes mm apart. features within the Geulhem Member of these 1.3 diameteris 0.9 mm between the peristomes, mm localities.

at the The lower lobes of are — peristomes. peristomes Occurrence elsewhere Middle Aptian-Eocene (mid- only weakly developed. Geulhem Member, col- dle Lutetian), of Belgium, France, Spain, Great Bri- lected from burrow infill at the K/T boundary, tain, Austria, Germany, Denmark, Sweden, USA, former Curfs at & quarry Geulhem, Voigt Meijer Poland, Czechoslovakia, CIS and India. leg., GPI Hamburg Collections, type catalogue, no.

Subfamily Serpulinae MacLeay, 1840 739.

1758 — The tube has Genus Serpula Linne, Description an initial non-spiral part

(PI. 2, Figs lb, 2a, 3), followed mostly by a spiral (PI.

Type species — Serpula vermicularis Linne, 1767. 2, Figs la-b, 2a, 4, 7) with normally 1-3, rarely 4

— of Discussion It is impossible to give a diagnosis turns. Diameter and height of spiral are generally

is defined charac- In the is attached the tube, because the genus by 3-5 mm. most specimens, spiral by

of the soft base the teristics parts (Hartmann-Schroder, a broad to a more or less planar substrate,

and because the tubes of various then truncated and 1971, p. 527), spiral resembling a cone, being

different from the sub- Recent species of the genus are very less frequently cylindrical. Occasionally each other. strate is small, the spiral then being inversely conical

' ’ in the is In addition, palaeontology name Serpula or forming irregular loops. Spiral coiling predom- has been used omnium but dextral as an gatherum for all kinds inantly sinistral, specimens are not rare. of serpulid tubes: Serpula has often been used almost An umbilicus is always present. Turns of the spiral

cemented the between as a synonym of the family Serpulidae. appear together, sutures 79

Clusters of three biont Rovereto, 1901 are com- them being indistinct. up to speci- Protulophila gestroi found in tubes of the 2, mens are known (PI. 2, Fig. la, b). monly present species (PI.

but do in other and Theanterior tube part (PI. 2, Figs 2, 5-10) is often Figs 1, 2, 4), not occur serpulid from the Geulhem Member, with obliquely, less commonly horizontally or steeply, spirorbid species

curved. to the of a of Placostegus erect, and is straight or slightly Length up exception single specimen Recent of P. The diameter increases to erectus A on 4.0 mm. tube gradually a (see below). paper examples

the and is in in maximum of 1.2 mm between peristomes gestroi preparation (Zibrowius &Jager, prep.).

Discussion — described and 1.8 mm at the peristomes. Regenhardt figured only

but the erect anterior tube A probably Normally there is no longitudinal ornament, part. spiral

before have keel of to the species was illustrated by two specimens me a irregular belonging present

A furrow is the of the Rasmussen Serpula ? insiticia is here assigned height. present at beginning (1965). resembles Peristomes the to the as it in some respects erect tube part. are very rare on genus Serpula,

the erect the Recent 1901) and spiral part, but are very common on part Crucigera zygophora (Johnson, both 4 are the which are representatives (PI. 2, Figs 1, 2, 6-12). Up to peristomes closely genus Cementula,

and the of the the spaced: they are sharp, short broad, upper group comprising genera Serpula, Hydro-

Cementula and the first- halfbeing broader than the lower half. The circum- ides, Crucigera, Spiraserpula,

named 'main of this ference of the peristomes is near-circular, with a big taxon being the genus' group

in the for historical lobe in the half and two small lobes reasons. upper

— The of the and the erect lower half. In weakly developed peristomes these Comparison shape spiral

indistinct. In the middle of the tube of Conorca 1961 is similar, but lobes may be upper Regenhardt, very

the here the tube's cross-section is surface and occasionally also at the two lobes on triangular, quad-

the lines underside the peristomes slightly protrude. On the rangular or pentangular, growth protrude

in the the tube's and base of the tube also more centre of upper sur- spiral part the top may clearly much than in lines are visible and the are longer protrude (PI. 2, Fig. 4). No growth face, peristomes

the the Rotularia is on the tube's exterior. The cross-section of present species. hisingeri (see below)

in the attached to the substrate its attached tube parts is inversely U-shaped, generally by non-spiral

The lumen The tube initial tube and not by the first turn of the erect part circular. is circular. part only,

The is lower, and are wall is moderately thick, and consists of a thicker, spiral itself. spiral peristomes

and thinner absent. The Recent yellowish, smooth outer parable layer a Crucigera zygophora (Johnson,

which the more than three times (maxi- white inner layer, probably represents 1901) grows larger

6 to inner parable layer plus cylinder layer (PI. 2, Fig. 3). mum diameter at peristomes mm according ten

wrinkles Hove & 1984, and In abraded specimens fine transverse may Jansen-Jacobs, p. 167), peri-

in the white well and tube are less common. Cemen- sometimes be seen layer as as on stomes erect parts

No 'internal tula has similar the inner surface of the yellowish layer. a tube structure (smooth outer sur-

in of the absence tube structures' such as those seen Spiraserpula (see face, 'melting together' spiral's turns,

At base of of 'internal tube The of Cementula below) occur. the sides of the structures'). general shape

tube cavities the is also similar, but those of the Cemen- attached parts, sometimes longitudinal peristomes

cells much more and rarely a construction of short [resembling tula/Spiraserpula group protrude clearly

of the tube surface and the those of Neovermilia ampullacea (J. de C. Sowerby, in the centre upper at

visible. lobes the underside. In the of the 1829), but less distinct] are on addition, spirals

in A single specimen (PI. 2, Fig. 13a-b) has two Cementula/Spiraserpula group are general flatter, so that several in with the tabulae following the first spiral turn, plus remains turns are contact substrate,

tabula of the initial of this in the in which of a third just anterior part contrast to present species only

the first is. the spiral. The tabulae are oblique, as they are fixed turn

inner side and distribution — Albert Canal to the tube wall at the spiral's more Geographic stratigraphic

lower of the Geulhem Member: not posteriorly and at the spiral's outer side more ante- sections, part

vaulted side rare. Geulhem section: riorly. The tabulae are lightly (concave rare, upper part, extremely of the Geulhem Member: anterior) and have two horizontal rows of minute lower part very frequent,

the and one on the lower rare. perforations, one on upper upper part, extremely

the Occurrence elsewhere — side. A somewhat larger perforation occurs at Regenhardt (1961, p. 54) similar from the border of the tabula at the inner side of the spiral. recorded one or two specimens

Danian of Fakse The few The characteristic structures of the hydroid sym- (Denmark). fragments 80

Fakse from in my collection do not allow a decision higher level than the others (PI. 2, Fig. 19b), to

their sub- as to being conspecific or not. rounded trapezoid (PI. 2, Figs 14c, 17b) or

squarish (PI. 2, Figs 16b, 18b). The lumen is

circular. Serpula ? aff. trilineata Roemer, 1841 The tube wall (PI. 2, Fig. 19a, b) is white and PI. 2, Figs 14-19 moderately thick, and consists of a thicker outer

aff. 1841 Serpula trilineata Roemer, 102. p. parable layer, a thin, slightly brighter inner parable aff. 1983 Janita trilineata (Roemer, 1841) —Jager, 72, pi. 9, p. wall layer, over which the easily splits, and a mod- fig. 2 (with additional synonymy), thin In the half of the aff. 1987 ? 27-30. erately cylinder layer. upper Janita sp. —Jager, p. 42, fig. 1; pi. 1, figs tube the aff. 1988 ? 74, 2. ratio of thickness of the three is Janita sp. —Jager, p. fig. layers

roughly 2-5 : 1 : 1-2, but the inner parable layer

— failed becomes thicker towards the base and form Type Jager (1983, p. 72) to retrace may

material in the Roemer-Pelizaeus- rows of short cells those in Roemer's type very comparable to seen

Museum the whereabouts of Neovermilia but less that in at Hildesheim, present ampullacea, distinct, so sections the base which is unknown. most appears to be massive. When

— the is the inner Description The tube is attached to large sub- cylinder layer worn, surface of the

shows strates (mainly oyster shells and calcareous algal parable layer a fine transverse striation.

but also its — ? aff. trilineata crusts, planar bryozoan colonies) over Discussion Serpula resembles some

entire with broad border. Recent length a Possibly erect species of the genera Serpula and Hydroides,

anterior tube but S. cf. parts were occasionally present, e.g. kaempferi (Kinberg, 1867) (see Imajima,

these have not been preserved. 1978, fig. 2s-u), H. exaltata (Marenzeller, 1884) (see

The tube forms U's (PI. 2, Figs 15-18) and irregu- Imajima, 1976, fig. 4j), H. externispina Straughan, lar of U/2 loops up to turns (PI. 2, Fig. 14), some- 1967 (see Imajima, 1976, fig. 3k). However, there is

times there in also certain are several successive a a resemblance to the Recent loops single genus specimen. The length of the tube reaches 25-30 Metavermilia (see below). For the time being, the

with tube diameter to a is here referred the with mm; gradually increasing species to genus Serpula a maximum of 2.0 of broad basal bor- mm (exclusive query.

or 3.0 of this There is — ? trilineata stricto is der) mm (inclusive border). no Comparison Serpula sensu very

in but its cross-section is longitudinal ornament juveniles (see juvenile part similar, commonly more in PI. but in medium-sized and adult rounded than that of aff. the keels 2, Fig. 14b), trilineata, may tubes there are always three straight (i.e. non-undu- occasionally be weak or undulate, and peristomes

smooth do and rounded keels 2, not occur at and the base show late), (PI. Figs 14-19). all, may longi-

The two peripheral keels are slightly wider than the tudinal cavities. median keel (PI. 2, Fig. 18a). Jager (1983, 1987, 1988) erroneously assigned

Transverse is in NW and sim- ornament already present juve- trilineata from Germany conspecific or niles (PI. 2, Fig. 14b) and in most of the medium- ilar material from the Maastricht Formation of the sized and adult tubes in PI. 2, 14b, 15; the Recent (distinct Figs Maastricht-Aachen-Liege area to genus less distinct in PI. absent from PI. in this 2, Figs 16-18, 2, Janita: genus the keels are much stronger and

Fig. 19). It apparently becomes weaker and finally undulate.

' ’ disappears in adults, but the material at hand does The Eocene Serpula trilineata J. de C. Sowerby,

allow substantiate this. It consists of wrinkles is not to 1844 a much smaller and, most probably, diffe-

which in the of or pustules slightly protrude centre rent species. Neovermilia ampullacea generally has only the surface and at the base. in keel keel at upper Especially a single or no all, and its peristomes are

of U's and the concave parts loops pustules are large much better developed.

devel- with (PI. 2, Figs 14b, 15). Weakly to moderately Comparisons similar Recent species as

annular rather oped peristomes are rare (PI. 2, Figs described and illustrated by Imajima (1976, 1978)

and in the follows: cf. 15, 16a), they slightly protrude same way are as Serpula kaempferi has three or five

do the wrinkles. keels in the fixed tube and as part a cylindrical erect

The cross-section is with the anterior with less keels trapezoid, periph- part pronounced or no keels eral keels the of the but forming upperedges trapezium, at all, with small flaring peristomes. Hydroides the side of the exaltata the upper trapezium being lightly con- has, according to description, many

that the median keel is situated these vex, so on a slightly growth rings, (however, are not distinct in the 81

and the diameter inner surface in where small illustration), grows to larger sizes, except areas fragments

3 at the aperture being mm. Hydroides externispina of the non-striate cylinder layer are preserved.

closely resembles aff. trilineata, but weak peristomes Discussion — The present species resembles Neover-

other de or transverse ornament apparently occur milia ampullacea (J. C. Sowerby, 1829), a long-

more commonly. ranging and widely distributed species in the Late

and distribution — Albert Canal Geographic stratigraphic Cretaceous, showing an enormous range of varia-

lower of the tion and into the Danian sections, part Geulhem Member: not (see Jager, 1983), ranging

Geulhem lower of Denmark. but rare, upper part, rare. section, part Slightly more triangular, probably

of the Geulhem Member: extremely rare, upper conspecific, forms occur around the Palaeocene/

trilineata stricto of the Basin part, extremely rare. Serpula sensu Eocene boundary Tremp (NE Spain;

in Drs R.H.B. (or a closely related species) occurs the late leg. Fraaye). Regenhardt (1961)

Maastrichtian Gronsveld, ? Schiepersberg, Emael assigned ampullacea to the genus Proliserpula

Meerssen Members of the Maastricht and ? Regenhardt, 1961, but a comparison of ampullacea

Formation. with the descriptions and figures of the Pliocene-

Occurrence elsewhere — Late Santonian (NW Ger- Recent Neovermilia falcigera (Roule, 1898) in

Zibrowius many): Serpula trilineata sensu stricto. & ten Hove (1987) has shown that the

tubes of similar these two species are closely in

? In other Serpula sp. many respects. some features, however,

there PI. 3, Figs 13-15; PI. 7, Fig. 1 are differences, but similarities of ampullacea to

another species of Neovermilia Day, 1961, N. globula Description — The tube diameter increases gradually

(Dew, 1959), make an assignment of ampullacea to to rapidly to a maximum of less than 3 mm. The

this more than tube is attached the substrate for its genus probable. to longest part,

— itself from Comparison Characteristic, though facultative, the anterior part, however, may dislodge

features ofN. are absent from the the substrate Most ampullacea present (PI. 3, Fig. 14). specimens are fine and few form and of species, e.g. large size, transverse ornament, curved, a even loops trochospirals annular In view of the wide of but these peristomes. range up to 2'/ 2 turns (PI. 3, Fig. 13), are only variation of often Often keel ampullacea, single specimens can- fragmentarily preserved. a median or be but the material PI. but not distinguished beyond doubt, crest occurs (PI. 3, Fig. 14; 7, Fig. 1), some from the Geulhem Member of the Maastricht- specimens lack longitudinal ornament. Transverse whole is different. and Aachen-Liege area as a clearly A ornament is only weakly developed, there are

few from this member cannot distinct fragments be distin- no peristomes. guished from Serpula ? aff. trilineata (see above) A single well-preserved tube fragment (PI. 3, Fig.

shows attachment beyond doubt; however, typical examples of this 14) a posterior to a narrow, species three longitudinal keels. unpreserved substrate; anteriorly it dislodges itself possess

Geographic and stratigraphic distribution— Albert Canal from the substrate for the last 1 mm of its length, sections, lower of the Geulhem Member: but without marked erection. The apertural diame- part very Geulhem is 1.3 An undulate rare, upper part, common. section, lower ter mm. strong longitudinal crest

and the lines of the Geulhem Member: extremely protrudes over the aperture, growth part rare,

part, absent. also protrude distinctly towards the crest. upper

Occurrence elsewhere — Unknown. Another well-preserved fragment (PI. 7, Fig. 1), originally attached to a large planar bryozoan col- Genus Cementula Regenhardt, 1961 emend. is almost 3 has ony, relatively large (diameter mm), Pillai (in press) and a strong longitudinal crest transverse growth

towards the the — lines protruding crest and towards Type species Cementula sphaerica Brtinnich Nielsen, base. 1931.

The cross-section is in Remarks — The Cementula first used inversely U-shaped some name was by

in with convex sides Briinnich Nielsen who failed specimens, near-triangular (1931), to designate a others. there is Thus it Generally a longitudinal cavity on type species. was Regenhardt (1961), who each side of the tube's base but introduced the He (PI. 3, Fig. 14c), validly generic name. distin-

cellular construction of the base occasionally a guished two genera, Cementula and Spiraserpula in Neovermilia occurs (PI. 3, Fig. 15b) as ampullacea. Regenhardt, 1961, mainly on the shape of the spi-

There striation the tube's is a fine transverse on which either be flatter. As the ral, may higher or 82

ofRecent shape of the spiral varies, Jager (1983, p. 47; 1991) On the basis of Pillai's studies (in press)

Cementula. similar and fossil Cementula-like united these genera under Very serpulids, two species

in Recent be with 'internal tubes also occur serpulid species usually groups may now distinguished: one

stricto tube structures' i.e. assigned to Serpula sensu (see Lommerzheim, (henceforth: ITS), longitudinal

of the tube wall into the lumen, and one 1979, p. 142;Jager, 1983, pp. 47-49). protrusions

Table 1. Number of specimens of serpulid and spirorbid taxa collected from the Geulhem Member as exposed in the Albert Canal

Collection sections (Vroenhoven-Riemst, Belgium). Lithologic log based on Meijer (1959). Specimens from the J.W.M. Jagt

Collections and are now included in the Natuurhistorisch Museum Maastricht (prefix NHMM, see plates). Meijer’s Voigt’s

collections of the Institut der Universität GPI samples are housed in the Geologisch-Pälaontologisches Hamburg (prefix

Hamburg). 83

oldest lithuus without ITS. In Recent species this difference is graphically species, ’Serpulites’ von

in 1820 correlated with distinct differences the soft parts, Schlotheim, from the Middle Mus- mikesia which made Pillai decide to separate them at the chelkalk of Germany, Spiraserpula Ziegler &

level. From the absence of ITS 1980 from the late Triassic of the generic or presence Michalik, western

similar fossil and unnamed alone in otherwise very tubes, one Carpathians an unique specimen

rather the from the of should expect a separation at subgeneric early Sinemurian Wellendingen (men-

levels Pillai found ITS in the tioned it is uncertain whether or specific only. type above), they belong to

in the all. species of Spiraserpula and noted their absence the Cementula/Spiraserpula group at

of and thus the members of the type species Cementula, two genera Unequivocal Cementula/Spiraser-

known from the Pliensbachian are now distinguished on the basis of an utterly pula group are early

before from different main distinctive character than originally onwards (the oldest specimens me are

intended (Regenhardt, 1961). the ibex Zone of Caen, France, and the lower

of and SW Ger- Diagnosis — The increase in diameter of the tube is davoei Zone Schomberg Bisingen,

attached substrate To the end of the middle these slow; it is to a or to posterior tube many). Jurassic

of its remain rather and it is therefore parts over most length. Following upon an species uniform, of the available initial several that least some non-spiral part are one or regular or possible at specific viz circinnalis irregular, mostly planar but occasionally taller spi- names, complanata (Goldfuss, 1831),

short in in rals, often with or long non-spiral parts (Miinster Goldfuss, 1831), filaria (Goldfuss, 1831) between. On the surface the tube and ramentum 1874) are upper parts (Dumortier, synonymous.

is often from the Toarcian bifrons Zone of cemented which no Specimens appear together, why Somerset lack ITS and therefore suture is seen. Transverse ornament consists of bird- probably are assig-

which and often nable Cementula. wing-like 'alae', are widely spaced to The Oxfordian and do not occur at all. In contrast to the closely similar spirolinites

ITS the in Goldfuss, with its Spiraserpula there are no protruding on inner (Miinster 1831) sharp longitudi-

The anterior tube become nal keel and alae is distinct, whereas tube surface. part may sharp utterly

silicified from the Tithonianof Nattheim raised above the substrate and possess sharp quadri- specimens

rounded 'alae' resemble the and middle partite peristomes consisting of on (SW Germany) again early

surface and on the the upper two pointed processes Jurassic species.

underside. Often there furrow There is record far of the Cemen- is a longitudinal on practically no so in the Cretaceous. In the underside. tula/Spiraserpula group early

— has often the late Cretaceous and Palaeocene this and Comparison Spiraserpula a very similar, group,

shows ITS main in shows wider of more planar tube, (the distinctive Spiraserpula particular, a range

and evolution. the feature), and there are differences in the soft parts. variation a more rapid Following

of the subdivision of the into The differentiation of Cementula from Laqueoserpula discovery group species

1979 — both ITS with or without ITS, Lommerzheim, genera lacking Jager's (1983, p. 48) hypothesis

is not quite clear. Laqueoserpula has a thick-walled, of a polyphyletic derivation of large species (mainly

from small wide tube with only few spiral turns visible from the Spiraserpula) species (mainly Cementula)

bulbous and be outside. The peristomes are more only can no longer supported.

The anterior Pillai's and own studies of the occasionally are birdwing-shaped. According to my

tube's Cementula Brunnich tube part often steeply erects. interior, sphaerica Nielsen,

— Middle Triassic-Sinemurian ?, 1931 Stratigraphic range (Maastrichtian), Spiraserpula spiraserpula

Pliensbachian-Campanian ?, Maastrichtian-Re- Regenhardt, 1961 (Campanian), S. versipellis

1961 and S. cent. Regenhardt, (Maastrichtian) scrupea

— Maastrich- 1961 must be left in Remarks Up to now, only Campanian, Regenhardt, (Maastrichtian)

their whereas S. subditiva tian, Danian and Recent species were assigned to original genus,

1961 is Cementula. Cementula and Spiraserpula as currently defined. Regenhardt, now assigned to Material of C. Brunnich 1931 Other Cretaceous species still need to be assigned to applanata Nielsen,

either of these while for most from the Danian of Fakse lacks ITS. In genera, Jurassic speci- (Denmark) material of unnamed from the is an mens assignment hampered by poor preservation Spiraserpula species

of material which Palaeocene of Kazachstan ITS in of the tube's interior or scarcity are present (Pillai,

forbids sectioning. Thus Cementula and Spiraserpula press). Spiraserpula adunca Regenhardt, 1961

here In strati- is a of are treated together. the two (Maastrichtian) species Placostegus. 84

Many more Cenomanian-Palaeocene species of a shape typical of the Cementula/Spiraserpula

beenrecorded and discussed Lommerzheim the tube's exterior is but have by group. Generally, smooth,

who omitted Cemen- a few (1979),Jager (1983, pp. 47-49, large spirals are granulate. Many specimens

tula ? cauteshabitans Lommerzheim, 1979 and C. ? are broken and show the tube's interior from which

ITS absent. applanata Briinnich Nielsen, 1931) andjager (1991). are — in Santonian S. Discussion Cementula from Fakse These species, as well as material of applanata

material from spiraserpula and C. sphaerica recorded byjager (1983) (topotypical Rejstrup was not avai-

lable for and C. subditiva from and S. parceproma Regenhardt, 1961 from the Cam- study) the Maastricht-

but panian, should still be thoroughly examined for Aachen-Liege area are closely comparable, not

of ITS. identical: it is often presence or absence although impossible to assign

It that the small Maastrichtian C. single to either as a appears species, specimens species, 'populations' whole several differences sphaerica, produced two lineages of larger species in display (see below). The

in fact sub- the Danian: one distinctly umbilicate, more planar two taxa may represent geographic

and which are best the lineage (C. applanata subditiva) a higher lineage species, separated at subspecific with its main species, C. contorta Briinnich Nielsen, level.

1931 which C. ? 1961 (of ventosa Regenhardt, may be a synonym). Cementula applanata subditiva (Regenhardt, Of fossil species it is uncertain post-Palaeocene 1961) whether they belong to the Cementula/Spiraserpula PI. 3, Figs 1-12 all. For Recent the reader is group at species

v 1961 Spiraserpula subditiva Regenhardt, 42, pi. 1, fig. 9. referred to Pillai & ten Hove (in prep.). p.

— The illustrated is Cementula applanata Brünnich Nielsen, 1931 Type specimen by Regenhardt the holotype by original designation; it is here reil- — 1926 Br. N. Serpula (Hydroides) aplanata fnc] 0dum, p. in 3 4 lustrated PL 3, Fig. 2a-c; it represents (or ?) 165.

to considerable The 1931 Cementula Briinnich spirals overlying a degree. large applanata Nielsen, p. 97, pi. 2, fig. anterior of maximum 7. spiral (diameter spiral 7 mm,

diameter of tube 1.4 mm) has c 4 turns, a narrow,

— 2-3 less Type Briinnich Nielsen (1931) illustrated but a deep umbilicus, 4 very prominent (plus

which did It alae its anterior 1 and keel single specimen, he not designate type. distinct) on '/a turns a on is from the Danian Bryozoan Limestone ofRejstrup its anterior 2/3 turn. The aperture is slightly

far (Denmark), and is preserved in the collections of the bulbous with a tooth-like process. As as can be

in broken and in Mineralogical and Geological Museum of seen a posterior spiral damaged

of the anterior ITS absent. Copenhagen University. parts spiral, are Geul-

— The is based hem Albert sections km mark Description following description on Member, Canal at material from the Bryozoan Limestone of Fakse: 23,850, sample no. 125 in the Meijer Collection

Tubes and spirals are medium-sized to moderately (1959), GPI Hamburg Collections, type catalogue

the maximum tube diameter 3 725. large, being mm, no. that 8 — A of the spiral mm. The spirals are planar or, Description single exceptional specimen (PI. 3,

curved the of more often, following curvature cylin- Fig. 1) attached to a slender cylindrical bryozoan

initial tube of drical substrates. Generally several spirals of the colony has a very slightly curved part

other considerable 0.5-0.6 in diameterand of 4 same specimen cover each to a mm a preserved length

due the small size of the sub- The tube then the in degree to relatively mm. enwraps bryozoan as seen

of strate, forming an irregular three-dimensional clus- Pegmaticula wegneri (Jager, 1983) forming a spiral

Umbilicate and non-umbilicate in l'/2 and is raised above the substrate. ter. spirals occur turns, finally

In all other the does about equal numbers. A longitudinal keel occurs specimens spiral not enwrap a

of few only in the anterior part very specimens, foreign body.

often there is indistinctrounded much more an edge Nearly all specimens have become detached from instead. Alae in the their or less and substrate and are common, especially outer (more planar large)

lack the initial tube Most whorls of large spirals, but also, though less dis- non-spiral part. specimens

show 3 4 and tinctly developed, in posterior tube parts. Erect 2, some or perhaps even more spirals,

less anterior tube parts are present, showing peristomes most spirals being more or planar (PI. 3, Figs 85

2-5), but occasionally very irregular and high (PI. 3, comprises but a single spiral, and only rarely does

In Figs 6, 7). Spirals generally cover each other for an umbilicus occur. C. contorta Briinnich Nielsen,

than half 1931 ? more of their surface. Sometimes short the turns become fairly high. C. boryszewiensis

remains of non-spiral, more or less straight tube Pugaczewska, 1967 from the Dano-Montian of Po-

visible between the land is often parts are spirals (PI. 3, Fig. 8). granulate. Spiraserpula spiraserpula

The diameterof the with 4 1961 from the is largest (anterior) spiral, c Regenhardt, Campanian indis-

in ornamented is the but turns, is 10.2 mm an exceptionally large speci- tinctly as present species, the

diameter of is normal. each other lesser men (PI. 3, Fig. 3): a 5-7 mm spirals envelop to a degree at least

In number of the last half takes 'a when the substrate and ITS a specimens turn was large, are present.

In S. from the Meerssen Member ITS short cut' by growing irregularly across the upper scrupea occur,

and the An side of the spiral (PI. 3, Figs 5, 7). Nearly always spirals are more depressed. unnamed

in from the of there is a distinct, narrow deep umbilicus the Spiraserpula species Palaeocene

centre of the spiral (see PI. 3, Figs 2a, b, 3, 4, 6). The Kazachstan is larger and has ITS, and the spirals

increases often become tube diameter slowly, and reaches a maxi- very high.

of 1.0-1.5 and distribution— Albert Canal mum mostly mm, rarely 2 mm. Geographic stratigraphic

of Spirals rarely bear longitudinal and transverse sections, lower part the Geulhem Member: not

in the and in Geulhem ornament (only holotype [PI. 3, Fig. 2] rare, upper part, not rare. section, lower

additional keel be of the Geulhem Member: an specimen [PI. 3, Fig. 5]): a may part very rare, upperpart,

present on the anterior 2/3 turn of the spiral, and absent.

alae the anterior 12/3 Occurrence elsewhere — Unknown. some may occur on turns.

There is no granulation.

The anterior tube above the sub- Genus Rotularia 1827a part may erect Defrance, (= Spi-

but short ofless than 3 in rulaea Tubulostium strate, only fragments mm Bronn, 1827,

length have been preserved (PI. 3, Figs 5a, 9, 10). Stoliczka, 1868, ? Rotulispira Chip-

all of the tube lonkar & Nearly fragments erect part possess a Tapaswi, 1973)

quadripartite peristome (PI. 3, Figs 9, 10) similar to

those of other specimens of the Cementula/Spiraser- Type species — Serpulat spirulaea Lamarck, 1818.

On underside there often Remarks — Lom- pula group. the occurs a Contrary to Regenhardt (1961),

and keel furrow (PI. 3, Figs 9b, 10c), occasionally a merzheim (1979, and in litt., 1984) and Knight-

The cross-section of the tube I still think that Rotularia and similar instead. erect part (PI. Jones (1981),

3, Fig. 9c) is very slightly depressed-circular. genera such as Conorca Regenhardt, 1961, and

The tube wall (PI. 3, Figs 3b, 7c, 11, 12a, b) is Orthoconorca Jager, 1983 should be referred to the

and consists of instead of the moderately thin, a lightly yellowish Serpulidae sensu stricto, to

outer parable layer, as thick as or thicker than the Spirorbidae.

other and the In Rotularia well in the tube two layers together cementing turns, a as as Spirorbidae, has

white inner with striation small parable layer transverse a non-spiral part (not clearly expressed by

its and white In the on interior, a innermost cylinder layer, Jager, 1983). Spirorbidae this tube part is

smooth on its interior and often broken or dissolved small and delicate and in very usually not preserved

(in part preserved in PI. 3, Figs 3b, 11). There are no adult tubes. However, in Rotularia, Conorca and

ITS related it is distinct and — (PI. 3, Figs 7c, 12a, b). genera more longer not

— Discussion Regenhardt (1961) thought that the only in absolute size, but also in relation to the size

of the and with in the of the of the adult tube — and is often shape aperture (bulbous alae) spiral very

holotype was a feature characteristic of the species. preserved. Rotularia is also much larger than spiror-

Flowever, most specimens of the species lack this bids, with only few of the 'normal' spirorbid species

it also the size of Conorca and the attribute, and, moreover, occurs occasionally reaching none reaching

in other of the size species Cementula/Spiraserpula group. of at times very large Rotularia. Moreover, if

— Cementula Comparison applanata applanata grows to Rotularia were to have had a calcareous operculum

while non-umbilicate have been slightly larger sizes, spirals are like 'normal' spirorbids, this should now

found. Alae not restricted to found least in of the and more commonly are at some large common

the of but also in of this its outer parts large spirals, may occur species genus. Therefore, operculum was

tube show calcareous. The of adult juvenile parts. Large spirals may granula- probably not spiral coiling

tion. C. a. sphaerica Briinnich Nielsen, 1931 often tubes is an insufficient criterion for placing Rotularia 86

other Ser- of Rotularia tumida de C. with the Spirorbidae, as non-spirorbid or a gastropod, (J. Sowerby,

be of less is a of but a pulidae are also more or spirally coiled, e.g. 1829) species Nogrobs, may part Rotularia. Cementula and Spiraserpula (Pillai & ten Hove, in phylogenetic lineage leading to

that the tribus Rotulariae Some of the Schmidt to exclude prep.). It is my opinion species preferred

because from Rotularia in fact of this Regenhardt, 1961 is but an artificial taxon, are true species genus, the Conorca and viz. (Mantell, 1822), callosa (Stoliczka, genera Rotularia, Gynaeconitis sowerbyi

1961 related. 1868). de C. and Regenhardt, are probably not closely polygonalis (J. Sowerby, 1829), quin-

— Tube small initial Additional material of Diagnosis with a non-spiral part quecarinata (Roemer, 1841).

followed less ovata and umbonata attached to a substrate, by a more or (J. Sowerby, 1813) (J. Sowerby,

conical is needed substantiate their regularly, tightly coiled planar or inversely 1813) to assignment to

the spiral with coils cemented to each other, but nor- present genus.

The 'Definite but Rotularia first mally not cemented to the substrate. spiral poorly preserved

The of the in the Ururoan 1967, 354, usually has an umbilicus. diameter spiral appear stage' (Stevens, p.

= As Stevens did generally exceeds its height. Coiling of the spiral is Pliensbachian-Toarcian). not pro- decided predominantly sinistral but occasionally dextral. vide figures of these specimens it cannot be

thick whether Rotularia The tube wall consists of two layers, a outer they actually belong to or Nogrobs.

and The same holds for a number of one which may appear to comprise two layers a non-figured speci-

Stevens tube wall be thickened mens from other strata recorded thin inner one. The may by by (1967,

tube the from callous material. The free anterior non-spiral pp. 354, 356, 357). Especially specimens

out be of is sometimes or curved. the Tithonian turn species part very long, straight early may the identification of the — and Comparison In most other spirally coiled genera of Nogrobs, early-middle Callovian material is uncertain. Ball Serpulidae the spiral is attached to the substrate by (1960, p. 21)

Orthoconorca conica referred & Bird 250 = at least the entire first turn. (von compressa Young (1828, p.

is but in this and concinnus de C. 1829, 195) from the Hagenow, 1840) an exception, J. Sowerby, p.

late Toarcian of Yorkshire and tumidus de C. other species of Orthoconorca and Conorca the very J. 1829 from the 'Corallian' slender shape of the spiral differentiates them from Sowerby, (p. 195) to whereas assign- Rotularia. Many species recorded by Rovereto Rotularia, Jager (in press b) prefers

In (1904b) are coiled Serpulidae, not Rotularia. some ment to Nogrobs.

known from literature Rotularia have descended from during Jurassic species exclusively may Nogrobs

Rotularia from Callovian-Oxfordian as at that time some data it is difficult to distinguish dme, spe-

1808 in cies of lost the characteristic cross- Nogrobs de Montfort, (see Jager, press b). Nogrobs square the umbilicus cross-section of the latter is section and had a callus as seen The genus normally filling

and in least but it is modified in in some of Rotularia, the Valanginian square or at quadrangular, species the reached the size of spiral part of some species. Generally, a square large planar Nogrobs spirals non-coiled Rotularia. this is based cross-section occurs in the free, anterior However, assumption on very

of the but there it be circular in few and literature data and needs to part tube, even may specimens only

In the be substantiated. some specimens. general, however, presence

absence of cross-section in the anterior The Rotularia is divided into four or a square genus usually

facilitates from Rotularia. This subdivision 1979; part distinguishing Nogrobs subgenera. (Lommerzheim,

Pliensbachian/Toarcian-Oxfor- is followed it is Stratigraphic range — Macellari, 1984) here, although

have dian ?, early Kimmeridgian-late Eocene. uncertain whether these subgenera really a

Remarks — Schmidt (1955a) presented a detailed monophyletic origin. Some species described by

of the of his whom is Rovereto revision present genus. Although most early authors, amongst (1904b),

followed I be without additional data data and conclusions are here, disagree cannot fully discussed with his assignment to Rotularia of a number of and are therefore omitted here. is The Rotularia with bicarinate illus- species, viz. ampullacea (J. de C. Sowerby, 1829) a sp. a periphery species of Neovermilia, lithuus (von Schlotheim, 1820) trated and described by Fleming (1972, p. 751) from is of middle Triassic the Tithonianof New Zealand to be a a poorly preserved species age, early appears is uncertain Aus- is a of Co- true Rotularia but the (? trochiformis (von Hagenow, 1840) species , subgenus

is not The lituola norca, heliciformis (Goldfuss, 1831) probably trorotularia). species (Leymerie, 1846b)

is from the be referable to Rotularia, subcarinata (Goldfuss, 1833) Neocomian cannot appreciated

internalmould and either be lack of literature data. an only, may a species through a 87

— R. (Austrorotularia) Macellari, 1984 has a 'tricari- stoliczkai (Chiplonkar & Tapaswi, 1973), inflata

keel in in both 1973 and varians 1973 ?. nate present juvenile, adult or stages. Pasternak, Pasternak,

In the R. (Praerotularia) and Tube commonly thickened by an external callosity' northern hemisphere,

and in central It is almost restricted to Rotularia as a whole became (Macellari, 1984, p. 1104). rare, India be absent from the Turonian the southern hemisphere, but occurs also in Europe they appear to

to whereas in the (Stoliczka, 1868) and Japan (Yabe & Nagao, 1928). (or Coniacian) Maastrichtian,

from the southern number of It was originally recorded Aptian- hemisphere a species are fairly

the Cenomanian-Turonian but it that Rotularia indet. common. Maastrichtian, appears sp. Obviously,

event affected the in the north- described and figured by Stevens (1967, p. 353) boundary genus only from the ? early Kimmeridgian of Antarctica and ern hemisphere.

1961 has recorded also from the early and middle Kim- R. (Tectorotularia) Regenhardt, a planar meridgian of New Zealand should also be referred spiral and a quadrangular to hexangular cross-sec-

would make tion. Hauterivian-Maastrichtian. to this subgenus, which Austrorotularia Range: Species

couloni and not Praerotularia is the stratigraphically oldest of included are: quinquecarinata (Roemer, 1841), the subgenera. Defrance, 1827a ?, gregaria (Etheridge, 1907), land- C. Cretaceous species of Austrorotularia include: ereri (Mallada, 1887), polygonalis (J. de Sowerby, rotuloidea andina (Camacho, 1949), callosa (Stoliczka, 1868), 1829), discoidea (Stoliczka, 1868), Chip-

ornata (Wilckens, 1922),,fallax (Wilckens, 1910), tenui- lonkar & Tapaswi, 1973, westfalica Lommerzheim,

laevis Macellari, 1984, zinsmeisteri Macellari, 1984 1979, rotula (Goldfuss, 1831), cf. discoidea (Stoliczka,

rotula ? and Macellari 1868) sensu Wilckens, 1924, (Morton, 1834) sp. (1984).

— 1979 to rotula Goldfuss, R. (Praerotularia) Lommerzheim, appears (non 1831).

R. below. be the 'main line' of Cretaceous Rotularia with the i (Rotularia): see

Rotularia became extinct at the Eocene- most species and perhaps also the most specimens. Obviously,

It comprises fairly large tubes with transversely Oligocene boundary. — and with without Mode It is clear that Rotularia most wrinkled surfaces or simple longi- of life com-

occurred soft substrates silt tudinal ornament. Range: Hauterivian-Eocene monly on (clay, marl,

and/or often associated with other soft-bot- (Eocene according to Lommerzheim, 1979, p. 174; sand),

Cainozoic of such personally I know of no occurrence tom dwelling serpulids as Pentaditrupa or

In of the adult this subgenus). Ditrupa. contrast to the size tube, small substrate for Rotulispira Chiplonkar & Tapaswi, 1973 (p. 209) Rotularia needed only a very set-

is of R. it tlement. the adult tubes often broke probably a synonym (Praerotularia); appears Probably spiral off from the and acted doubtful whether the features mentioned by Chip- juvenile non-spiral parts as

sunken into the seafloor lonkar & Tapaswi (p. 209), having detachmentof an anchor lying on or partly

with the the apertural portion without thickening by cal- aperture pointing obliquely upwards.

without of the Macellari found that in T. careous deposition and constriction (1984, p. 1103) (Aus-

is bored carnivorous tube; also the detached portion of the tube very trorotularia) specimens by gastro-

in Rotularia’ these were close much longer than what is found are pods probably predation traces to

the and he concluded that distinctive features important enough for separation aperture, perhaps

Rotularia burrowed into the soft substrate with the at the generic level. The only specimen of Rotulispira

of R. almost How- known to date is the holotype stoliczkai Chip- aperture pointing vertically upwards.

the anterior of the lonkar & Tapaswi, 1973. ever, position gastropod preda-

of the include: tion traces could also be the of Species present subgenus ovata (J. explained by position the worm's soft in the anterior tube. More- Sowerby, 1813) ?, phillipsii (Roemer, 1841), umbonata parts

it is not for Rotularia to have bur- (J. Sowerby, 1813) ?, sowerbyi (Mantell, 1822), over, necessary into the but the pupoides (Stanton, 1901) ?, australis Cox, 1953, kitchini rowed actively sediment, spiral part

kitchini of adult tubes have been covered with sedi- (Bonarelli & Nagera, 1921) ?, cf. ?, concava (J. might

damesii without to Sowerby, 1813), chathamensis Boreham, 1959, ment passively being disadvantageous

(Noetling, 1884), saxonica Miiller, 1966, spirulaeoides the . I agree with Wrigley (1951, p. 182, on

R. that, 'The various modes of coil- Glaessner, 1958, gregaria (Etheridge, 1907) sensu (R.) bognoriensis) from almost discoid have Regenhardt, 1961, dorsolaevis Ball, 1960 ?, shackletoni ing, to conic, may

the rate of accumulation of London (Wilckens, 1910), libyca Maccagno, 1946, libyca var. depended on the ornata Maccagno, 1946, solarioides (Wanner, 1902), Clay sediment. Where this was rapid, creature 88

rise its in its had to tube a conical form to keep pp. 170, 177), pseudospirulaea (Oppenheim, 1901),

in clear of silt. With slower accu- (Gabb, 1860), nysti (Galeotti, 1837), spi- aperture water, leptostoma rulaea nummularia Schlotheim, mulation, discoid forms might prevail.' (Lamarck, 1818), (von

= of The total absence of Rotularia from the late Cre- 1820) junior synonym spirulaea, mayeri

Defrance, 1827a = taceous chalk facies of central Europe may possibly (Rovereto, 1898), complanata

of be explained by the competition of similar spiral junior synonym spirulaea, clymenioides (Guppy,

such Conorca and var. crassa soft-bottom dwelling species as 1866), angulosa (Chenu, 1842), angulosa

Orthoconorca conica. (Doncieux, 1926), corbarica (Munier-Chalmas in

Hebert, 1882) (fide Rovereto, 1904b), dickhauti

in subcarinata Subgenus Rotularia Defrance, 1827a (White, 1881) (illustrated White, 1882), (Goldfuss, 1833) ?, cristata Defrance, 1827a ?, lituus Type species — Serpula spirulaea Lamarck, 1818. Defrance, 1827a (non lithuus von Schlotheim, Diagnosis — The medium-sized spiral generally is 1820]). planar to low trochospiral, with longitudinal orna- Records from the early Oligocene [nummula (von either and when ment present or absent, present Koenen, 1891) and koeneni Rovereto, 1898 [= tumida there is no quadrangular to hexangular cross- sharp have been (von Koenen, 1891) non Sowerby] not section of the tube. Transverse ornament is but discus- confirmed and are probably erroneous, see there and weak, are no laminated growth lines, no The record of sion in Schmidt (1955a, p. 178). peristomes. Rotularia cf. pseudospirulaea by Jakubowski & Musial

— Discrimination from Austrorotularia is Comparison doubtful. (1977) from the Polish Tortonian is very Three keels be also in not always easy. may present

Rotularia, but in these cases the tube is not thickened

external The tubes of Tectorotularia by an callosity. Rotularia (Rotularia) hisingeri (Lundgren,

have a to hexangular cross- sharp quadrangular 1891) Praerotularia has section. larger and a trans- grows PI. 4, Figs 14-19 versely wrinkled surface. Longitudinal ornament is lithuus von 98. non 1820 Serpulites Schlotheim, p. either absent or weakly developed. 1831 lituus ? 6 vidi, Serpula Hisinger, p. 134, pi. 3, fig. (non — Danian-late Eocene. Stratigraphic range fide Schmidt, 1969). — Remarks The derivation of the — present subgenus, 1837 Serpula Lituus [sic\ Schloth. Hisinger, p. 20, pi. 4,

from 8. either from Austrorotularia or Tectorotularia, is fig.

1891 Serpula Hisingeri Lundgren, 119. clear. Derivation the former is p. not yet from taxon — 1920 24. Serpula Hisingeri Lundgren Rosenkrantz, p. because of the fact that in the strati- favoured — 165. 1926 Serpula (Spirorbis) Hisingeri Lundgr. 0dum, p. ofAustrorotularia the tri- — youngest species 1931 Briinnich graphically Spirorbula Hisingeri Lundgr. sp. Nielsen,

becomes but then this 1-4. carinate keel indistinct, again p. 103, pi. 3, figs

Pal- 1955a Rotularia (Lundgreen) [.sic] — Schmidt, 176. should not be overestimated. Already in the hisingeri p.

1960 Spirorbula hisingeri (Lundgr.) — Kiihn, 164. there is of variation far p. aeocene a wide range as as 1969 Spirorbis (Dexiospira) hisingeri (Lundgren), 1891 — morphology is concerned. Schmidt, p. 36. Palaeocene species include: hisingeri (Lundgren, — of the Pal- speci- 1891). Probably, at the beginning Type Hisinger's (1831, pi. 3, fig. 6) original

is lost there had been unknown men Schmidt, 1969, p. aeogene, a radiation; an probably (see 37).

had shows a with five turns and a 'survivor' species produced a considerable Hisinger's figure spiral

anterior tube its number of different but similar species, all of small protruding straight part, length

therefore consid- the diameter of the size. The following 'species' are exceeding spiral.

have from of tobar is stated to come ered to represent but subspecies (Gardner, Hisinger's specimen

the of but 1935), viz. mcglameryae (Gardner, 1935), sp. (Gar- Klinteberg (Gotland, Sweden),

that this dner, 1935), tobar (Gardner, 1935), gracilis n. subsp. Regnell {in Schmidt, 1969) remarked must

mistake. Most it from the and vroenhovensis n. subsp. be a probably originated

Eocene species include: bognoriensis (Mantell, Danian of Skane (southern Sweden).

— The tube has less 1822), cortezi (Gardner, 1939), horatiana (Gardner, Description a more or straight

= initial 1939), leymeriei Rovereto, 1898 nom. nov. pro non-spiral part (PI. 4, Figs 14c, 15b, 16b, 17,

of which short of 4 quadricarinata Münster sensu Leymerie, 1846a (— 18c, 19c), only a fragment up to

is The attachment substrate is angulosa Chenu, 1842, according to Schmidt, 1955a, mm preserved. to a 89

of the in is not a true Rotularia, the preserved only a single atypical specimen (PL 4, hisingeri shape spiral

this tube then of Fig. 19c, see below). The diameter of part being just a matter convergence. — Rotularia differs from other is up to 1.0 mm. Comparison hisingeri the absence of The diameterof the spiral is rather small, 4.0-5.2 species of the genus by general orna-

the conical, but low ment and by its small size. mm, spiral being inversely very

3'A and distribution— Albert Canal (PI. 4, Figs 14b, 16b, 18b, 19b), with up to turns Geographic stratigraphic Geulhem Member: in specimens from the Maastricht-Aachen-Liege sections, lower part of the com-

Geulhem lower and to 5 turns in Danish and Swedish part, rare. section, area, up mon, upper very and of the Geulhem Member: specimens according to Hisinger (1831, 1837) part absent, upper part,

Briinnich Nielsen (1931). The umbilicus is invaria- absent.

Occurrence elsewhere — Danian of Sweden bly wide. Of eighteen sufficiently well-preserved spi- Denmark,

and Austria. rals from the Maastricht-Aachen-Liege area fifteen

sinistral and three are (PI. 4, Figs 14, 17, 19) are

dextral (PI. 4, Figs 15, 16, 18). The turns of the Rotularia (Rotularia) tobar (Gardner, 1935) spirals are cemented together. 281, 26, 1-5. the 1935 Tubulostium tobar Gardner, p. pi. figs The tube diameter gradually increases up to

Tubulostium tobar Gardner — 19. 1939 Gardner, p. transition from the spiral to the protruding anterior — 177. 1955a Rotularia tobar (Gardner) Schmidt, p. be swollen and part, where the tube may slightly — diameter. In Gardner the reach to 2.0 mm in (1939) designated holotype 1.5 mm, rarely up Type is almost anterior illustrated in 1-3. It an the horizontally protruding straight part specimen figs with tube which be 4.2 in of 6 mm in diameter a (PL 4, Figs 15, 16), may up to mm planar spiral of 1.0 from the Wills length, the tube diameter decreases slightly. diameter (at aperture) mm,

bears Point Pal- The tube normally no ornament. Rarely Formation, Upper Midway Group (late

Station be shallow but indistinct United States Geological Survey there may a very wide, aeocene),

furrow side of the 6 miles south of McFarland sheep pens and longitudinal on the upper spiral 11754,

miles southeast the Windmill and weak at the 27 of Eagle Pass, on a very longitudinal edge periphery In there be low ranch road, Maverick County, Texas. United of the spiral. rare instances, may (Jacal)

and the States National Museum Collections, no. 370940. swellings at the end of the spiral at begin-

anterior tube A —■ 'Shell small tube coiled ning of the straight part. single Description a angular with surface incision in its which almost in one flattened, specimen shows an spiral was a single plane,

and the other The tube, possibly brought about by an injury. slightly depressed centrally.

in The cross-section of the tube is circular (PI. 4, increasing very slowly diameter, rudely quadrate lined ill-defined the tube wall in cross the outer by Figs 14b, 16b, 18b, d, 19b), being section, angles

outer between the flattened. moderately thick to thick. The lumen is circular, spirals; margin spirals

and lower surfaces concave. sometimes slightly eccentric (PI. 4, Fig. 18d). The The upper very feebly

thicker and Outer of the the tube wall comprises a outer layer a margin aperture subquadrate, coherent the coil. thinner inner layer. The tube's exterior is smooth. inner circular, firmly to preceding

Evidence of former lines discern- Only when the inner layer is destroyed by weather- growth probably which the surface bear ible in certain parallel aperture ing does the inner appear to a trans- rugosities wrinkle the shell. characterized and ... verse striation. minutely [...]

discoid outline and the outer A single specimen (PI. 4, Fig. 19) differs from the by the quadrate mar-

of the others in the character of its attachment: not only gin aperture.' (Gardner, 1935, p. 281). — tube attached Discussion The shows an was the non-spiral initial part to an species 'astonishing

but amount of variation' 1935, Gar- unpreserved planar large substrate, excep- (Gardner, p. 281). dner listed from different localities from tionally also the spiral. The base with which the specimens

the substrate in shows the Wills Point and Kincaid tube was attached to part a (Upper Midway)

Formations. Some of the cellular construction with very short closely spaced (Lower Midway) speci- localities third with ? mens from certain 'develop a spiral cells (PI. 4, Fig. 19c). Confusion Serpula insit- it the keel, almost in icia would have been possible in this case, were not midway upon equal prominence

those the of the keel'. is for the fact that in all other features this specimen to at margins Also, 'the apex

the Maverick with normal R. as a rule more County agrees wholly hisingeri. depressed upon

— ruled that forms' Wills Point 'than those Discussion It cannot be wholly out (= Formation) upon 90

of the Colorado River drainage' (= Kincaid relatively small, the spiral being generally very low,

Formation). almost totally planar. The cross-section of the tube

stated when is with the side of Gardner (1939, p. 19) already com- sharply square to trapezoid upper

paring mcglameryae with tobar that the 'range of varia- the trapezium mostly positioned at the periphery of

between tion in these forms is well and the more the not very known, spiral, rarely oblique upper

side and the The of the possibly these two Midway species may be only periphery. two upper edges

varietally distinct.' These two American species and trapezium bear small, sharp granulate keels. In

the two taxa from the Maastricht-Aachen-Liège most specimens a third, small sharp keel in the

below similar in and of the side. The transverse area described are very shape centre trapezium's upper

consists of somewhat are probably closely related to such an extent that ornament minute, undulate

they should not be distinguished at the species level. ridges. The tube wall is moderately thin to just

but but thick. They possibly represent ecological varieties, moderately

— The tube has initial are here considered of tobar Description a non-spiral subspecies , perhaps gen- part,

uine geographic subspecies. In the Maastricht- of which only short fragments of up to 2.5 mm in

neither and 1.0 mm in diameter are Aachen-Liege area, a phylogenetic progres- length up to preserved.

of the sub- The attachment of this initial tube the sion nor a geographic separation two area part to

found. substrate is The diameter of the species can be not preserved. spi-

— Rotularia is smaller and ral is 4.5-7 7 also the Comparison hisingeri easily generally mm; mm being

the near-absence of maximum diameter. The is even lower than distinguished by any ornament, spiral

of with of whereas R. dickhauti (White, 1881) is larger, but in that tobar vroenhovensis, hardly any trace

There 5 the similar otherwise. Whether the latter is to be consid- becoming trochospiral. are up to turns,

another of tobar is and wide. ered to represent yet subspecies umbilicus always present nearly always remains to be determined. In R. leptostoma (Gabb, Of sixty-eight sufficiently well-preserved spirals

the sinistral and 1860) spiral is more distinctly trochospiral at thirty-two are (PI. 5, Figs 1, 3, 4) thirty- least in juveniles, and the two sharp keels at the six dextral (PI. 5, Fig. 2).

Rotularia In the is In spiral's periphery are more closely spaced. rare instances, spiral irregular. a single horatianum and has the of the substrate is (Gardner, 1939) grows larger a specimen impression seen,

distinct The three keels which that several turns of the very transverse ornament. means juvenile spiral of of several of tobar must have been attached to this substrate. many specimens subspecies directly

(gracilis in particular) resemble those of R. (Aus- In another specimen (PI. 5, Fig. 4) the spiral had

but this three reached at least 5 mm in when trorotularia), subgenus grows two to already diameter,

the tube broke the 3/4 whorl dis- times as large, and its umbilicus is narrower. at periphery at

the of tance from the aperture. At point breakage

the animal then formed 9 tube a new mm long Rotularia (Rotularia) tobar gracilis n. subsp. of the old The tube growing out one. new at first PI. 1-6 5, Figs x followed the periphery of the old one for /\ whorl,

— then short anterior In third Derivatio nominis gracilis (L.) meaning graceful, in formed a straight part. a

the tube had also broken. Here the comparison with tobar vroenhovensis the tube being specimen new

somewhat thinner first from the for short slightly smaller and having a part at grew away spiral a the wall. distance, then turned back to grow along

— of of 0.9 in Type The specimen figured in PI. 5, Fig. 1 is here periphery the spiral, leaving a cavity mm

of the designated holotype; it is a sinistral spiral (6.5 mm length and 0.4 mm in width between the turns

In in diameter) with 4'/2 turns, followed by a short, spiral. a fourth specimen (PI. 5, Fig. 3) the last '/2 follow but obliquely broken and somewhat distorted straight turn did not the periphery of the spiral, anterior tube Ornament is well its underside. part. developed. grew across

The third longitudinal keel is relatively weak, but The tube diameter gradually increases up to the

of the transition from the the anterior present. Albert Canal sections, upper part spiral to protruding

Geulhem Museum where the tube is Member, Natuurhistorisch part, often slightly (c 45°) distorted

Maastricht NHMM Collections, no. 1992201-1 around its longitudinal axis (PI. 5, Figs la, c, 4a, 4f),

(leg. M. Jager). but scarcely swollen, and may reach a maximum

— of Diagnosis The diameter of the tube and spiral are diameter 1.5 mm. In the horizontally protruding 91

which be Rotularia tobar vroenhovensis straight anterior part (PI. 5, Fig. 5), may (Rotularia)

the diameter remains n. subsp. up to 8.7 mm long, fairly tube has three PI. 6, 1-6 constant. Generally, the (PI. 5, Figs Figs keels. The 2d, 3b, 4c), more rarely two small sharp

and forms Derivatio nominis — After the of Vroenhoven/ tube's base is somewhat enlarged village

and sides NE the Albert Canal rounded granulate edges on the left right Riemst, Belgium, near

and lower side of the The sections. {i.e. at the upper spiral).

marked at the — The illustrated in PI. 1 is transverse ornament is especially Type specimen 6, Fig.

and where small here The sinistral keels edges, some granules or pus- designated holotype. spiral (diam-

7 3 has three At the tules but occurs also in between. eter height mm) turns. may appear, mm, transition from the the The lumen is circular. In the tube wall (PI. 5, Figs spiral to anterior tube part

the the tube diameteris 2.3 mm. At the broken anterior le, 4b, 5c, 6) two layers can be distinguished, and the tube wall thicker than the inner one. end the tube diameter is 1.8 outer layer being slightly mm,

often is a delicate inter- is thick there, but this is caused Between them there very apparently very by

material. another tube schlot- stice formed by disintegration of tube serpulid (very probably Ditrupa

in which Here the tube wall breaks easily, case a heimi) filling the lumen (cone-in-cone structure).

Member with bruennichi rather indistinct transverse striation becomes Geulhem (top) Tylocidaris

visible. Ravn, 1928 and D. schlotheimi, Albert Canal sections

— would soft Museum Discussion The planar spiral suggest a at Vroenhoven-Riemst, Natuurhistorisch

In Maastricht NHMM 1992202-1 bottom habitat with a low sedimentation rate. Collections, no.

samples yielding tobar gracilis, the soft bottom (leg- J.W.M. Jagt).

in — the tube and dweller Ditrupa schlotheimi, which occurs great Diagnosis The diameter of spiral are the numbers, is practically the only other serpulid moderately small, spiral being inversely conical, but low low. The cross-section of associated. relatively to very

— the tube is rounded with the side of Comparison In the closely similar tobar s. str. only a trapezoid upper between few of the specimens have three longitudinal keels, the trapezium positioned obliquely upper

the side and the The side of the and those from type locality possess only two spiral periphery. upper

resembles flat band keels. In tobar gracilis, however, the three-keeled ver- trapezium a protruding laterally

left two-keeled version. and a rounded on the and sion is commoner than the forming edge right

Transverse is absent. Rotularia lobar vroenhovensis is slightly more margins. ornament normally

tube diameter, a The tube wall is thick. trochospiral, has a distinctly larger relatively

and — The tube has initial far less distinct longitudinal transverse orna- Description a non-spiral part,

of which short 1.5 in wall. Most of the of to mm ment and a thicker tube specimens only fragments up

and 1.0 in diameter two subspecies from the Maastricht-Aachen-Liege length up to mm are preserved.

referred either of these Its attachment a substrate is not The area are easily to on account to preserved. of is 5.5-7 of their morphology; transitional specimens are diameter the spiral generally mm, more

does it reach maximum of 8 mm. Most rare. rarely a

Specimens of Rotularia spirintorta (Rovereto, spirals are fairly regular, only few are slightly irregu-

the is 1904b) from around the Palaeocene/Eocene lar (PI. 6, Fig. 4). Although spiral relatively

trend is dis- boundary of the Tremp Basin (NE Spain, leg. Drs low, a trochospiral recognisable (less

R.H.B. similar but tinct in PI. more so in PI. Fraaye) possess a very ornament, 6, Figs lc, le, 2b, 6, Figs

their 5b, The of the is more variable are easily distinguished by high trochospirals. 4b, 6). height spiral than it is in R. in tobar does In tobar mcglameryae the transverse ornament and, in hisingeri or gracilis. Rarely

keel the of the reach 80 % of its most specimens, the third longitudinal are height spiral diameter,

the diameter decreases absent. and at the top of the spiral

that the becomes Geographic and stratigraphic distribution — Albert Canal again, so spiral barrel-shaped (PI.

lower of the Member: to 4'/2 turns, the umbilicus is sections, part Geulhem 6, Fig. 6). Up always suffi- Geulhem section, present and wide. Of forty-eight absent, upper part, frequent. nearly always sin- lowerpart of the Geulhem Member: extremely rare, ciently well-preserved spirals twenty-seven are

istral 6, 1, 3, 4, and twenty-one dextral upper part, very rare. (PI. Figs 6)

Occurrence elsewhere — Unknown. (PI. 6, Figs 2, 5). 92

The tube diameter increases to the has of gradually up trochospiral, generally (its range variation is

transition from the the anterior and spiral to protruding at present unknown) three longitudinal edges

tube where the tube often is distorted 'shows under the lens rather part, slightly a peculiar granular or

and swollen and where it reach etched (PI. 6, Fig. la) may a an appearance' (White, 1881, p. 161).

of In the — maximum diameter 2.3 mm. horizontally Geographic and stratigraphic distribution Albert Canal

anterior tube which the protruding straight part, may sections, lower part of Geulhem Member:

be to 6 mm 6, the diameter up long (PI. Fig. 3), extremely rare, upper part, frequent. Geulhem sec-

decreases lower of the slightly. tion, part Geulhem Member: absent,

The tube has two In most longitudinal edges. upper part, absent.

specimens these edges are prominent only laterally, Occurrence elsewhere — Unknown.

whereas the upper side of the band is smooth (PI. 6,

Figs lb, c, e, 3, 5b). However, in a few specimens Genus Ditrupa Berkeley, 1835 these also the side of edges are prominent on upper

the band — (PI. 6, Figs 2b, d, 4b). Rarely does a third Type species Dentalium subulatum Deshayes, 1825 =

weak occur in the of the Dentalium arietinum O.F. longitudinal edge centre Miiller, 1776 (see ten Hove

band is band. This with the edges present not only & Smith, 1990).

the but also the tube on spiral, on Hove & protruding part Diagnosis (see ten Smith, 1990, p. 102; data

6, in brackets added the — (PI. Figs 2d, 3). by present author) Tube Transverse ornament is generally absent. Only free, not attached to substratum, elongated curved

do fine wrinkles fusiform rarely or irregular swellings (mostly shaped like an elephant's tusk with

about of brought by regeneration injuries occur. the curvature slightly decreasing towards the ante-

The tube section is rather variable in detail. The rior circular in cross-section. Tube end), open at tube wall reaches its maximum thickness at the both ends, broadening anteriorly though exterior

transition from the to the anterior before the tube mouth. spiral protruding tapers just (In general, no the lumen circular. part (PI. 6, Figs Id, 2c), being longitudinal edges). Tube wall consisting of variably

The tube wall consists of two the outer and often layers, being (thick) hyaline outer opaque (thin, bearing

1'/2 4 times thick the inner. when the to as as Only a transverse striation) inner layer.

inner — layer has been destroyed by weathering, does Range Danian-Recent.

the inner show — surface a transverse striation. Often Remarks Derivation of from the Ditrupa pentangu-

water movement washed tubes of D. schlotheimi lar or genus Pentaditrupa Regenhardt, 1961 (early Sin- other cylindrical or conical objects (cone-in-cone emurian-Danian) via 'mixed populations' of Penta- into the lumen. structure) Moreover, recrystallisa- ditrupa subtorquata (Miinster in Goldfuss, 1831) which has filled the interstices between tion, these (middle Turonian-Danian) comprising specimens and the tube is in the with objects wall, relatively strong either pentangular, subpentangular or circular

interval from which most of the specimens were cross-section, (as suggested by Jager, 1983, p. 56), collected. Both has been post-mortem phenomena provide an now substantiated by ten Hove's examina-

for the lumen explanation fact why the to tion of the tube structure in appears (see Jager, press a).

amount to 20 % of the tube diameter only in a Except for the variety of P. subtorquata with circu- number of specimens. lar cross-section, all of the hitherto examined (some

' Discussion — Rotularia tobar vroenhovensis is found in being in need of further studies) so-called Ditrupa’ sediments and is reladvely coarse-grained thought from the early Jurassic to the late Cretaceous have

to have been better adapted to slightly higher turned out to differ from true Ditrupa. The five conditions than tobar of its energy gracilis on account ‘Ditrupa’ species recorded by Tate {in Tate & Blake, thicker wall and distinct slightly more trochospiral 1876) all differ from true Ditrupa. ’Ditrupa’ varicosa

1960 has nodular shape. Ball, swellings or more rarely

— Comparison R. tobar has an even lower gracilis annular constrictions and a different tube structure. smaller spiral, a distinctly tube diameter, a much Ditrupa antarctica Wilckens, 1907 has an irregular marked and more longitudinal transverse ornament D. dentalina curvature, while (Leymerie, 1851) and a thinner tube wall. R. tobar be mcglameryae resem- appears to closely related to Pyrgopolon (see bles but t. vroenhovensis, has a still lower with Lommerzheim spiral below). (in litt., February 1984) now only scarcely rounded R. dick- considers all his longitudinal edges. Cenomanian ’Ditrupa’ species to be hauti is but less (White, 1881) closely similar, related to P. mosae de Montfort, 1808. The two 93

of & and Museum of species of ’Ditrupa’ Chiplonkar Tapaswi (1973) port, Mineralogical Geological Collections. are dubious. Copenhagen University

— The tube is curved rather The stratigraphically youngest pentangular tubes Description tusk-shaped,

sometimes almost with longitudinal edges have been described from weakly (PI. 4, Figs 1-9), being

' the Danian of Denmark and Israel [see Ditrupa’ straight (PI. 4, Fig. 10), with curvature decreasing

' tube diameter interjuncta Jessen & 0dum, 1923 and Hamulus subtor- towards the anterior end. The gradu-

increases of 0.8-1.5 and a quatus’ of Avnimelech (1941), both being typical ally to an average mm,

maximum of 1.6 Towards the the representatives of P. subtorquata]. Ten Hove (in lilt., mm. aperture

Recent and dimin- September 1989) wrote, 'I have never seen a outer tube diameter slightly gradually

thousands ishes and the Ditrupa (and I have seen literally of speci- again, apertural region usually cannot

tube'. Thus it be discriminated from tube mens) starting off with an angular clearly posterior parts.

that alto- is the swollen club-shaped (PI. appears longitudinal edges disappear Only rarely aperture

— — the is gether even in early ontogenetic stages during 4, Figs 6a, 7a). Exceptionally, apertural region the Cainozoic. bottleneck-shaped (PI. 4, Fig. 8). The maximum

is tube is 40 to Rosenkrantz The oldest unequivocal Ditrupa D. schlotheimi length c mm according from the Danian (see below). During the Cainozoic (1920).

there is the tube shape changed only slightly, and discrimi- Occasionally, an irregular narrow, sharp often this nation of the various species described is rather longitudinal furrow (PI. 4, Figs 10, 11), but

for few millimetres in which difficult. does not extend but a

case it either disappears altogether or reappears

about torsion of Ditrupa schlotheimi Rosenkrantz, 1920 after an interruption, brought by the furrow PI. 4, Figs 1-13 the animal in its tube. Adjacent to two

slightly elevated, rounded longitudinal edges may 1820 Dentalites laevis 93 from pars von Schlotheim, p. (specimens of these occur, and on the other sides two edges, Copenhagen only, those from the Muschelkalk weak furrows. excluded). there may be very

non 1822 Dentalites laevis von fig. 2. Schlotheim, p. 107, pi. 32, The tube surface is usually smooth. In some

1847 Serpula liberaeformis M. u. H. — Anonymous, p. 118. specimens growth lines (especially visible in abraded 1863 Ditrypa liberaeformis Mus. Univ. Hafn. Terkelskov. — tubes), wrinkles, transverse constrictions and bulges Morch, 467. p. in the anterior are seen 1920 Schlotheimi 9. (particularly part) (PL 4, Figs Ditrupa Rosenkrantz, p. 25, pi. 2, figs 8,

Schlotheimi Rosenkr. — 167. The tube and lumen circular 1926 Ditrupa 0dum, p. 2, 4, 5a, 7a, 8, 10). are

1931 Schlotheimi Rkz. — Briinnich Ditrupula sp. Nielsen, p. (PI. 4, Figs 5b, 6b, 7b), the tube wall being mod- 95. erately thick. 1961 Ditrupa (Ditrupa) schlotheimi Rosenkrantz — and 65-80 The outer parable layer is thick, equals 72. Regenhardt, p. % of the entire wall thickness. In sec- 1965 schlotheimi — 26. longitudinal Ditrupa Rasmussen, p. 35, pi. 8, fig.

— tions the laminae 1967 Ditrupa (Ditrupa) schlotheimi Rosenkrantz, 1961 \sic\ (PI. 4, Figs 12, 13) parable are

1-3. Pugaczewska, p. 184,pi. 1, figs 1-4; pi. 2, figs occasionally visible, and between the laminae and 1967 Ditrupa schlotheimi Rosenkrantz 1920 — Hucke & the tube's longitudinal axis is an angle of c 25-30°. 42, 8. Voigt, p. 98, pi. fig. At the ofthe outer and inner the wall — junction layers non 1973 Ditrupa (Ditrupa) cf. schlotheimi Rosenkrantz, 1920

a white a 9-11. separates easily, making layer bearing Pasternak, p. 33, pi. 5, figs

— visible non schlotheimi 1920 transverse striation 4, 1984 Ditrupa Rosenkrantz, Ziegler, p. strong (PI. Fig. 13). 237, pi. 6, fig. 5. Often fragments of small specimens have been schlotheimi & table 1. 1988 Ditrupa —Jagt Janssen, tube. This washed into the lumen of a larger cone- 1988 Ditrupa schlotheimi Rosenkrantz, 1920 —Jagt & Col- be confused with in-cone structure should not a 190. lins, p. tube Some tubes show more complex structure. pre-

— Rosenkrantz dation traces of carnivorous gastropods. Type The specimen illustrated by

Discussion — Within the Geulhem Member, the (1920, pi. 2, figs 8, 9) was designated 'holotype' by pre- the but in fact these are two sent fine-grained Ziegler (1984, p. 238), species obviously preferred in the specimens. Rosenkrantz's figure 8 is here designated lithofacies, although it is not rare coarse-

it tube with facies either. Thus, in contrast to P. subtor- lectotype; is an anterior fragment aper- grained

it confined substrates in low diameter 1.5 11 was not to ture, c mm, length c mm (measure- quata, muddy ments taken from the illustration). Late Danian, energy settings.

— of and lower Crania Limestone, Copenhagen, southern Comparison A variety Campanian 94

fossil Maastrichtian representatives of Pentaditrupa subtor- variation, and assignment of species to a cer-

with but circular tain is The most quata moderately curved tubes and genus problematical. practicable

solution this to be the cross-sections lacking longitudinal edges may to problem appears place-

but of these small three-keeled with closely resemble the present species, generally ment species annu-

darker and the lar in Metavermilia, and with flar- grows larger, has a slightly colour, peristomes species

and tube diameter decrease in the outer tube diameter near the aper- ing peristomes rapidly increasing

in de C. ture is more distinct. Vermiliopsis (e.g. fluctuata J. Sowerby, 1829;

1973 from Briinnich such ’Ditrupa’ cf. schlotheimi sensu Pasternak, dorsolineata Nielsen, 1931). Formerly,

classification the fact that the Campanian-Maastrichtian of European Russia a was substantiated by

is curved be referred Metavermilia lacks funnel- more strongly and cannot to apparently flaring (=

it but the present species; may belong to Pentaditrupa. shaped) peristomes (ten Hove, 1975, p. 57),

— described such Geographic and stratigraphic distribution Albert Canal Imajima has subsequently peri-

lower of the Geulhem for Metavermilia M. sections, part Member: stomes spicata Imajima, 1977,

M. truncata 1978 extremely frequent, upper part extremely frequent. gravitesta Imajima, 1978, Imajima,

Geulhem Mem- M. which record has made Geulhem section, lower part of the and ovata Imajima, 1978,

the classification of fossil tubes difficult. ber: extremely frequent, upper part extremely more

frequent. Another problem is that similar three-keeled spe-

— cies also in the and Occurrence elsewhere Danian of Poland, Denmark, occur genera Serpula Hydroides.

in Lommerzheim described in northern Germany erratic boulders only. (1979, p. 158) a species

Metavermilia with four weak under the name n. sp.

' Genus Metavermilia Bush, 1904 emend. keels from the early to middle Cenomanian. Ser-

Zibrowius, 1971 pula’ cincta Goldfuss, 1831 is either a primitive repre-

of Meta- sentative of Filograna or a specialised species Type species — Vermilia multicristata Philippi, 1844. vermilia. The Vepreculina Regenhardt, 1961 is — genus Diagnosis Most species are small, some medium- be of Metavermilia. For now considered to a subgenus sized to large. In most species the tube is attached to ’Serpula’ trilineata Roemer, 1841, which was erro- its entire but also the substrate over length, some neously referred to Janita or Janita ? byjager (1983, have tube and of M. erect parts, (Vepreculina) above. 1987, 1988), see fimbriata (Regenhardt, 1961) only the non-attached

In and tube parts are known. some species loops Metavermilia ? sp.

irregular spirals are common. The tube diameter Pl. 7, Figs 2, 3 increases slowly to moderately rapidly.

is — The five minute tubes before me are A distinct longitudinal ornament usually pre- Description the underside of sent (except in M. taenia Zibrowius, 1971), consisting preserved as impressions on a tubes of of 9 which be or bryozoan colony (PL 7, Fig. 2), Placostegus up to keels, may straight undulate,

denticulate with lobate alatus 7, P. erectus, ? insiticia and smooth or or spines or pro- (PI. Fig. 3), Serpula

S. ? the tubes of the tuberances, or the keels are replaced by longitudinal sp. Probably present species

and of Transverse is were dissolved. rows granules or spines. ornament aragonitic diagenetically

4 of tube are generally less well developed, and occasionally con- Although only up to mm length

while all are more or less distinctly sists of annular swellings, some species possess preserved, specimens

of them flaring peristomes. The cross-section is circular to curved, some are hook-shaped or S-shaped.

The tube diameter increases to to rounded triangular to quadrangular. gradually rapidly

— Toarcian-Recent. a maximumof mostly 0.3-0.6 mm, in one specimen Stratigraphic range Early

— all five there Remarks In and Cretaceous 0.8 mm. In specimens are five narrow many Jurassic but keels small to medium-sized attached tubes sharp and fairly straight (non-undulate) on sequences of the tube. three the and lateral sides are with a simple sculpture usually consisting of upper They

than keels and more at the periphery longitudinal weakly to moderately pro- slightly closely spaced In about annular ’Ser- on the side. two three nounced peristomes are common (e.g. upper specimens

small but which are not pula’ tricristata Goldfuss, 1831). Obviously, they sharp peristomes occur,

the of Recent such Ver- and not In one of them (PI. 7, belong to group genera as flaring protruding.

these situated close to the miliopsis, Pseudovermilia, Metavermilia, Neovermilia, Jan- Fig. 2) peristomes are

A third 7, has one ita, Filograna and others. However, within these gen- aperture. specimen (PI. Fig. 3)

and wide of an additional era tube morphology shows a range well-preserved, highly fragmen- 95

The Hamulus 1834 and Turbinia which at the genera Morton, tary, peristome, slightly protrudes show Michelin, 1845 also Ornatoporta Gar- median keel. The other two specimens do not (and possibly 1961 and the of all five is dner, 1916, Triditrupa Regenhardt, peristomes. The cross-section specimens 'Octogonae' of Regenhardt, 1961; no material was inversely U-shaped (PI. 7, Fig. 3b). Danian resemble second in — late closely Jager's group Comparison Amongst Cretaceous and examined) calcareous bear those of tube structure as well as in having a species, the tubes some resemblance to here be and so that too are regarded to Vermiliopsis dorsolineata(Briinnich Nielsen, 1931), operculum, they

both of Pyrgopolon, rather than of V. fluctuata (]. de C. Sowerby, 1829), which subgenera separate

five but to sizes, and genera. have keels as well, grow larger medium-sized to tubes. Diagnosis Tube generally very generally have well-preserved with diameter at least in In V. dorsolineata the keels are more closely spaced large, rapidly increasing

lateral sides the attached of most Many species the most of the part species. on upper side, leaving have anterior tube and a long erect part. Longitudinal free. The peristomes are somewhat flaring pro- in keels keels often six or seven number, are trude at the median keel, at the two peripheral or crests,

whereas lack ornament. A V. keels are often some species and at the base. In fluctuata the common, few A relatively complex undulate, and the peristomes are somewhat flaring species possess peristomes. well towards tube structure is often developed. Perforate and distinctly protruding at the lateral sides

in and in contrast to the side. tabulae occur some species, the upper the is — Albert Canal majority of other Serpulidae, operculum Geographic and stratigraphic distribution great with lower of the Geulhem Member: calcareous (probably aragonitic), a sections, part entirely marked differentiation into cucullus and rare. Geulhem sec- clearly extremely rare, upper part, very

of the Member: calcar. tion, lower part Geulhem extremely Hauterivian absent. Stratigraphic range Early ?, early rare, upper part, Cenomanian-Recent. Occurrence elsewhere — Unknown.

1808 Genus Pyrgopolon de Montfort, (— Sclerostyla de Montfort, 1808 Morch, 1863, Hepteris Regenhardt, Subgenus Pyrgopolon

1961)

Diagnosis — Tube medium- to large-sized, with tube

— in the attached mosae de 1808. diameter increasing part, Type species Pyrgopolon Montfort, rapidly in fossils. Remarks — A closer comparison of fossil species of which is fragile and often poorly preserved

and with Recent The base of the attached generally shows a Pyrgopolon, ‘Sclerostyla’ Hamulus part

loan from Dr H.A. cellular construction from a ‘Sclerostyla’ kindly received on ten well-developed (absent Uni- lacking 'Favosites'-struc- Hove (Instituut voor Taxonomische Zoölogie, few specimens), invariably modern In in from Mr Alex Muir Tabulae occur in fossils, versiteit van Amsterdam), and ture. species.

and in the of and Dr G.T. Pillai (Natural History Museum Lon- the attached tube part posterior region

into classi- the erect the tube wall splits an don), has shown thatjager's (1983, pp. 79-82) part, easily and inner is in the differen- an fication of ‘Sclerostyla’ part wrong: outer (= outer parable) (= cylinder) has of is but over a weak median which tiation between two groups species correct, layer layer, probably

of different Thus, the the Recent (and some of the Palaeogene) species a mineralogical composition.

much related and thus less intact inner often be ‘Sclerostyla’ are more closely more or layer may

probably consubgeneric with his second group removed easily from the outer layer. bears than with his first The erect tube of fossil species either ('Zweite Gruppe') rather ('Erste part keels has Gruppe'). seven longitudinal edges or or no longi-

' the of S. tudinal ornament at all, occasionally there are Consequently, type species ‘Sclerostyla ’,

and the of wrinkles. In Recent the ornament ctenactis Morch, 1863, type species transverse species The both differ. There is no 'honeycomb' ornament. Pyrgopolon, P. mosae de Montfort, 1808, belong may striate makes a has a funnel-shaped, radially to Group 2 now, which Sclerostyla junior operculum

slender circular of cucullus and a calcar, to triangular synonym Pyrgopolon. the in cross-section. For the first group, name Septenaria Middle Cenomanian-Recent. Regenhardt, 1961 is available. Stratigraphic range 96

mosae mosae de The of the is 5.5 Pyrgopolon (Pyrgopolon) layers. length fragments up to mm,

1808 the 1.7 and Montfort, diameter up to mm. Much longer some-

PI. ? ? 6 8, Figs 1-4, 5, what thicker specimens are known from the

Maastricht Formation. The cylinder layer is fairly 1808 Mosae de 394. Pyrgopolon Montfort, p. 395, text-fig. on p. thick and thin The posteriorly, anteriorly. cross-

section (PI. 8, Fig. 5b, c) generally is circular, but

— has far been be Type Obviously no type specimen so may occasionally subtriangular posteriorly. Montfort illustrated De a — designated. (1808, p. 394) Discussion Hitherto, P. m. mosae was not known to

unattached anterior tube lon- straight, part lacking occur in the Palaeogene. Specimens found in the

but with transverse gitudinal ornament, growth Geulhem Member could be considered to have been

'dans l'interieur de la de Mae- lines, from montagne reworked from the underlying Maastricht Forma-

stricht'. The subterranean of Maastricht and them galleries tion, perhaps some of really have. Three

have been excavated in the Nekum Member be rule the mostly arguments can put forward to out possi- of the Maastricht Formation (late Maastrichtian). bility of all of them being reworked:

— tube have been col- Description Only fragments - although the material from the Geulhem Member

lected from the Geulhem Member. or Opercula is fragmentary, a few specimens are relatively com-

external moulds such known from the of are only plete and unabraded,

Maastrichtian The - of the (Cupedo, 1980). great majority although many Geulhem Member speci-

of the the tube collected from the fragments represent erect part. mens were basal metre in the two

Remains of the attached also tube are there part compar- sections studied, are also several specimens

atively rare in the Maastrichtian, in comparison Canal and (Albert sections) many specimens (Geul- with the of the The mass occurrence erect hem from 8 parts. section) strata some m (first locality) or

inner has been parable layer normally dissolved, several metres (second locality) above the K/T

and in residues the Geulhem Mem- washing (from boundary.

ber well from as as the Maastricht the - in Formation) the late Maastrichtian Meerssen Member, P. m.

is found from the outer parable layer separated ciplyana is the dominant subspecies, whereas in the

Geulhem Member P. and few cylinder layer. many m. mosae only P.

The rather thin to thick generally moderately m. ciplyana are found.

outer is almost invaria- — parable layer (PI. 8, Figs 1-4) Comparison Pyrgopolon (P.) m. ciplyana (de Ryckholt,

in two is the whole has the tube bly fragmentary. Only specimens 1852) seven longitudinal edges on erect

circumference one 8, preserved: specimen (PI. Fig. part. Close to the aperture these edges disappear

the one from the Geulhem Member show and number distinct 1), only to a of transverse wrinkles or ribs

remains of the attached tube has a the part, complete appear. Dissociated parts of cylinder layer can-

erect tube of 1.8 mm in and 4.0 mm be part only length not distinguished from P. m. mosae. The opercula in diameter the which the at aperture, equals aver- are different.

diameterof from the Geulhem Mem- clava sub- age specimens Pyrgopolon (P.) m. (Lamarck, 1818), a

ber as well as from the Maastricht Formation. The species unknown from the Maastricht-Aachen-

second is of specimen (PI. 8, Fig. 2) a fragment the Liege area, but common at Ciply, St Symphorien

5.7 and diameter 2.3 and Mons erect part, length mm, a mere (southern Belgium) and at Kopinge

mm. (southern Sweden, Campanian), is strongly curved

The outer surface is but in a in its with the tube wall generally smooth, erect part, becoming very few there weak moderate thick towards specimens are to trans- the aperture.

the of verse wrinkles 8, la, near or the outer be (PI. Fig. b) aperture Fragments parable layer may weak or irregular longitudinal furrows or edges, confused with bivalve shells, while bits of the cylin-

at the base of the tube as der be confused especially erect part, layer may with chelae of decapod known from the Maastrichtian. Six with often specimens crustaceans, which, however, are oval in

cross-section. a more distinct longitudinal ornament are assigned

the and — to subspecies P. mosae ciplyana (see below). The Geographic stratigraphic distribution Albert Canal inner surface of shows fine lower of outer parable layer a sections, part the Geulhem Member: fre- transverse striation 8, 3b, Geulhem (PI. Figs 4). quent, upper part, very rare. section,

The is slender cylinder layer (PI. 8, Figs 5, 6) lower part of the Geulhem Member: extremely fre-

and consists of several laminatedconcentric conical, quent, upper part, extremely rare. This subspecies 97

also in the Member Forma- and — Canal occurs Lanaye (Gulpen Geographic stratigraphic distribution Albert

of Geulhem Member: tion), Valkenburg, Schiepersberg, Gronsveld, sections, lower part the very

Members absent. Geulhem lower Emael, Nekum and Meerssen (Maastricht rare, upper part, section,

in the of the Geulhem Member: Formation) Maastricht-Aachen-Liege area, part extremely rare,

also in and is extremely common in some strata. upper part, absent. This subspecies occurs

Occurrence elsewhere — Campanian ?, late Maastrich- the Lanaye to Emael Members exclusively in

Kunrade and in the Meerssen Memberof the tian of northern Germany (Hemmoor, a few atypi- facies, cal fragments overlooked by Jager, 1983), Belgium, Maastricht Formation.

Sweden ? and CIS. Occurrence elsewhere — Late Maastrichtian, Belgium.

Pyrgopolon (Pyrgopolon) mosae ciplyana Subgenus Septenaria Regenhardt, 1961

(de Ryckholt, 1852)

PI. 8, Figs 7-9, ? 5, ? 6 Type species — Sclerostyla (Septenaria) septenaria

1961. — de 1850 de Regenhardt, 1852 Ditrupa ciplyana Ryckholt, Ryckholt, p. 124,

— Tube medium- to with pi. 6, fig. 26. Diagnosis very large-sized,

rapidly to moderately rapidly increasing diameterin

Type — De Ryckholt's original specimen is from the attached part, which is often well preserved.

whereabouts is Ciply (southern Belgium); its present The attached part in most species has one or several unknown. keels. The base lacks cellular longitudinal edges or

— Description The best-preserved specimen was construction, but shows ‘Favosites’ structure, i.e. the found after the plates had been prepared and is interior of the lower half of the tube wall consists of therefore illustrated. It is from the minute 'tubes' with not a fragment many oblique-polygonal many

the Geulhem base of the Geulhem Member at sec- transverse bottoms, closely resembling the Pal-

of the tube aeozoic of smaller tion comprising the outer layer erect coral genus Favosites, but much

8 and 3 with At least in tabulae part, mm long mm wide, seven size). some species, occur rounded longitudinal edges and transverse ribs in occasionally.

five thin-walled The tube wall does weak between. The other are fragments, not comprise a layer, so all smaller than 4 in and of the it is the inner of the mm length width, that impossible to extract part

often tube wall from the outer parable layer. Up to three, more two outer part.

keels In the tube often nine longitudinal edges or are preserved (PI. 8, Figs erect part seven (or five, or

In there is furrow keels The 7a, 8, 9). some specimens a on the more) longitudinal edges or occur. erect

keel of the inner surface underneath the edge or part often has, the attached part less often has 'hon-

surface Transverse ribs outer (PI. 8, Fig. 7b). are eycomb' ornament {i.e. the outer layer of the tube

visible in of the five wall consists of minute hollow only one fragments (PI. 8, Fig. polygonal 'honey-

from the which somewhat and 7a): no fragments aperture are available, combs', are oblique, protrude

becomes At the inner in where this ornament stronger. an apertural direction). surface of but all fine The is unknown in all but most, not specimens a trans- operculum one species;

is Lommerzheim described verse striation seen (PI. 8, Fig. 7b). (1979, pp. 160, 161, fig. 8)

— collected from burrow Discussion The samples opercula belonging to P. (S.) cf. dilatata (d'Archiac, infill at the K/T boundary occasionally contain 1847), but from the description it cannot be deter-

from their specimens which, judging preservational mined whether or not these were found in situ.

from the Meerssen — Hauterivian style, obviously stem underlying Stratigraphic range Early ?, early

Member (burrow walls). These specimens have been Cenomanian-late Maastrichtian, Danian (see dis- omitted from Tables 1-3. However, some specimens cussion), early Eocene-early Oligocene ?.

and in the Geul- very probably are indigenous occur

above the K/T hem Member up to 1 m boundary. Pyrgopolon (Septenaria) sp.

— In P. well PI. 4 Comparison (P.) m. mosae longitudinal as 7, Fig. as transverse ornament is absent or at least weaker

— The sole available from the and more irregular. Dissociated parts of the cylinder Description specimen

be Geulhem Member is broken both and layer cannot distinguished. There are, however, posteriorly

confused differences in opercula. Fragments may be anteriorly, is 12 mm long and has a diameter of

bivalve and 3.5-4.5 It consists of short of with shells, cirripedes decapod mm. a very fragment

the attached with the socle and crustaceans. part a long, slightly 98

curved distorted of — and distinctly fragment the erect Stratigraphic range Oxfordian-Recent.

tube The attached has — part (PI. 7, Fig. 4a, b). part a Remarks Triangular tubes without protruding

median and rounded teeth single strong crest two longi- from the Pliensbachian are attached apically

tudinal in the the becomes their edges, erect part crest only, being free for longest part. They occur

lower into and there six (changing a keel), are commonly in soft bottom settings (claystones) and

rounded Fine wrinkles their mode of life similar that of edges. transverse occur (PI. probably was to

The cross-section is do 7, Fig. 4a, b). drop-shaped at Pentaditrupa or Ditrupa. Presumably, they not

the transition from the attached the to erect part (PI. belong to Placostegus, and their triangular cross-sec-

and near-circular in the 7, Fig. 4c), (subheptangular) tion is a matter of convergence.

' erect part (PI. 7, Fig. 4d). ‘Favosites’ structure does The late Jurassic Serpula’ prolifera Goldfuss, 1831

in the socle of the lower half of the occur not only with unusually strong peristomes might well be the

in tube, but also the median crest on top of the stratigraphically oldest species of Placostegus.

attached part (PI. 7, Fig. 4b, c). There is no 'honey- During the late Cretaceous and especially in Pal-

comb' ornament. time of which aeocene many species occur, some

Discussion — Three of the species subgenus are were assigned by Regenhardt (1961) to Eoplacostegus.

known from the late Maastrichtian Meerssen Mem- An examination of fossil and Recent material has

ber in the the differences Maastricht-Aachen-Liege area, pre- shown that the between Eoplacostegus and

sent the small specimen possibly being only representative Placostegus (Jager, 1983, pp. 96, 98) are too to

in the Geulhem Member. The accompanying label warrant generic separation. Lommerzheim (1979,

that the which the states sample yielded specimen p. 163; in litt., February 1984) shares this view.

was collected from burrow infill directly at the K/T In the late Maastrichtian of the Maastricht-

Geulhem and the of it P. aduncus boundary at notes possibility Aachen-Liege area, (Regenhardt, 1961),

reworked from the late Maastrich- referred is being underlying erroneously to Spiraserpula originally, a

tian. the However, same sample also contains speci- common species.

mens of the exclusively Danian species Serpula ? Paliurus Gabb, 1876, which closely resembles

insiticia and Neomicrorbis in the has club- cingulatus exactly same Placostegus, comparatively sharp edges, a

state of preservation (unworn, same colour and fine like swollen apertural region, and perhaps a cal-

In structure). addition, Pyrgopolon (Septenaria) careous operculum (Lommerzheim, 1979, p. 164).

becomes towards the of the Meerssen It is clear whether these differences rarer top not warrant a

Member. Thus, I assume this specimen to be generic separation.

autochthonous. 1961 stricto Conorca Regenhardt, sensu probably

Geographic and stratigraphic distribution — Albert Canal descended from Placostegus during the late Cre-

lower of the Geulhem Member: sections, part taceous (Jager, 1983, p. 98).

absent. Geulhem absent, upper part, section, lower alatus of the Geulhem Member: Placostegus (Brünnich Nielsen, 1931) part extremely rare, PI. 10-18 absent. 8, Figs upper part,

Occurrence elsewhere — Unknown. 1931 alata Brunnich 19. Serpentula Nielsen, p. 109, pi. 3, figs 18,

Genus 1844 — Placostegus Philippi, (= Eoplacostegus Type Briinnich Nielsen's (1931, pi. 3, fig. 18)

is It is curved Regenhardt, 1961) original designated lectotype herein. a

of the attached tube with and fragment part, a keel a

Type species — Serpula tridentata Fabricius, 1780. single birdwing-like peristome (ala). Middle Danian, — In not Coral Diagnosis many, but all, species the Limestone, Fakse (Denmark), Mineralogical attached tube forms and part often loops. The tube Geological Museum of Copenhagen University cross-section is keel Collections. generally triangular with a on the side. The lines The attached upper growth are curved for- Description posterior part (PI. 8, Figs wards towards the keel. The anterior tube is curved the free anterior part 10-13) or twisted, tube becomes less the cross-section more or steeply erect, part (PI. 8, Figs 11-18) usually steeply erect (PI. 8,

in and circular being triangular most species, in Figs 12b, 13a), straight, curved or rarely twisted and others. In but there In the most, not all, species are three often distorted. attachedpart the tube diame-

teeth the The tube far short protruding at aperture. diameter ter (as as can be seen in the fragments increases to Tubes of modern increases 1.5 slowly gradually. spe- available) gradually up to generally c cies translucent. with are usually mm, a maximum of 2 mm. 99

? I?? (P.) 7 (P.) indet.

(R.) r 1 ?

? ? trilineata ula gracilis sp. Serpula us vroenhovensis (Septenaria) et Cycloserpula aff. Placostegus Neomicrorbis Neomicrorbis cingulatusBipygmaeus pygmaeus Pileolaria Spirorbidae, Neodexiospira Neodexiospira lombricus Serpula? insiticia Serpula Cement Rotularia hisingeri(R.)Rotularia (R.) tobar Rotularia tobar Ditrupa Metavermilia Pyrgopolon Pyrgopolon Pyrgopolon alatus Placostegus erect PileolariaSP.I fcfij. gen, sp. m h!L_ hLI Isp.l Hi' applanata subditiva schlotheimi parietalis jagti mosae mosaemosae ciplyana

Table 2. Number of specimens of serpulid and spirorbid taxa collected from the Geulhem Member as exposed in the Geulhem

(Ankersmit) section (Geulhem, Valkenburg aan de Geul, The Netherlands). Lithologic log based onMeijer (1959). Meijer’s

Institut der Universität and Voigt’s samples are housed in the collections of the Geologisch-Paläontologisches Hamburg

(prefix GPI Hamburg).

anterior tube sections. The attached In the attached part there is a single non-denticu- to or posterior late longitudinal keel (PI. 8, Figs 10-12), which part forms a socle (PI. 8, Figs 12b, 13a, b), the lateral

form the tube. In the erect there of which somewhat to the two appears to rest upon part parts protrude

three lower of the erect are three roundedlongitudinal edges, forming edges part.

The ornament consists of sharp protruding teeth at the aperture (PI. 8, Fig. transverse birdwing-like

in intervals 18). Generally, the two lower edges are more closely peristomes (alae) wide, irregular (PI. 8,

Between the towards the spaced than they are to the upper edge. Figs 10-12, 14, 15, 17); they protrude

there is median furrow keel and towards the base. At the underside of the two lower edges a (PI. 8, Figs

15 erect form a small on either side of 12b, 13b, c, 14a, b, c, d). part they bulge median furrow but tend Most of the fragments (PL 8, Figs 11-13) show the the (PI. 8, Fig. 14a, b), they

where furrow itself. transition from the attached to the erect part, to efface over the

The the tube wall is thicker and less fragile as compared cross-section of the attached part (PI. 8, Fig. 100

of V/i sinistral 11) is strongly rounded subtriangular to inversely loop up to turns anteriorly, (PI. 7,

U-shaped with convex sides, that of the erect part Figs 5, 7b, d, e) and dextral (PI. 7, Fig. 10) loops

both At the base there is mod- (PI. 8, Figs 13c, 14b, 15d, 16) being drop-shaped on occur. a generally

broad border account of the unequal distances between the three erately broad, rarely very (PI. 7, Fig.

thick does the border form but edges. The tube wall is moderately to thick, 6). Rarely a narrow, high,

from the attached the socle in especially at the transition to caterpillar-like (PI. 7, Fig. 7a-c) as seen

is of the late Maastrichtian P. erect part. The lumen circular. many representatives

The tube wall is crystalline yellowish. In most aduncus (Regenhardt, 1961).

and The anterior tube becomes under specimens the wall appears to be homogeneous, part erect an

thicker of c 45-60° but this only occasionally may a outer parable layer angle (PI. 7, Fig. 9), angle may

somewhat thinner inner be also be smaller and a parable layer greater (PI. 7, Fig. 10), or (PI. 7,

of thin Erect tube often curved distinguished. Occasionally remains a very Fig.7d, e). parts are slightly

does the inner and cylinder layer occur. Only rarely distorted (PI. 7, Fig. 10).

distinct The In the attached the tube diameter surface show a transverse striation. part, gradually

in tube wall lacks cells and longitudinal cavities. increases, the erect part it remains constant (PI. 7,

— In the tube diameter is 1.0 Comparison Placostegus erectus (Briinnich Nielsen, Fig. 11). general, c mm,

with of 1.4 of the 1931) is smaller, has no 'alae', a different tube struc- a maximum mm. Fragments

different cross-section the reach 14 in of ture and a of erect part. attached part up to mm length, the

which The alae of Cementula and Spiraserpula, erect part up to 6 mm.

resemble those P. alatus the side In the attached often undu- closely of on upper part a keel, slightly

of construction is distinct but in of the tube, are a more complex at late, generally very (PI. 7, Figs 5-8),

the In the the underside of erect part. these genera some specimens there is only an indistinct edge (PI. tubes form spirals, whereas P. alatus does not. 7, Fig. 10). The keel is usually non-denticulate, less

Geographic and stratigraphic distribution — Albert Canal commonly denticulate (PL 7, Figs 6, 7b, d, e) consis-

of the Geulhem Member: of forward-directed little which sections, lower part com- ting low, teeth, may

not Geulhem lower either occur on of the keel or it. In mon, upper part, rare. section, top replace two

of the Member: it is that the denticulation ends part Geulhem not rare, upper part, specimens seen absent. exactly at the transition from the attached to the

Occurrence — Coral Limestone At the of the lateral walls elsewhere (Middle erect part. lower half two

Danian), Fakse (Denmark). there often is a weak seam-like furrow on either side

(PI. 7, Fig. 7b, d).

In the there three Placostegus erectus (Brünnich Nielsen, 1931) erect part are fairly sharp edges

PI. 5-14 non-denticulate of 7, Figs or keels (PI. 7, Fig. 11), over most

their length, but forming three sharp protruding Goldfuss, 238, 70, 15. v non 1831 Serpula erecta p. pi. fig. teeth at the (PI. 7, Figs 9, 12-14). The 1926 Br. N. — aperture Serpula (Pomatocerus) [sic] erecta 0dum, p. keel and tooth often bear median 167. upper a narrow

Briinnich 1 At the lower of the two 1931 Serpentula erecta Nielsen, p. 99, pi. 2, figs furrow (PI. 7, Fig. la). part

13. 12, lateral walls there is a weak seam-like furrow on

1961 (Eoplacostegus) erectus (Nielsen) — Eoplacostegus in the attached each side (PI. 7, Fig. 1 la, b) as seen 61. Regenhardt, p. At part. the underside of the erect part there often is 1961 Eoplacostegus (Concamerarius) inexpectatus Regenhardt, 1 10. an obtuse median edge (PI. 7, Figs lc, 12b, 13), less p. 62, pi. 9, fig. commonly a seam-like furrow.

— absent Type Brünnich Nielsen's (1931, pi. 2, fig. 13) Transverse ornament is often or very

herein. It is Some show weak original is designated lectotype a frag- weakly developed. erect parts may

ment showing the transition from the attached to peristomes, especially in their anterior regions,

the where the teeth of former erect part, which is not very steep. Bryozoan protruding apertures may

Min- Limestone (Middle Danian), Fakse (Denmark). be seen (PI. 7, Fig. 14a, b).

Museum The of the attached is eralogical and Geological of Copenhagen cross-section part strongly

University Collections. rounded-triangular to inversely U-shaped or subcir-

— with the Description A rather small species, cular, with obtuse edges and convex sides, that of

attached tube part curved irregularly (PI. 7, Fig. 6), the erect part being triangular (PI. 7, Figs 12b, 13,

wide twisted (PI. 7, Fig. 8) or forming a generally 14c) with usually sharp edges and near-straight (nei- 101

lateral The tube in close Within the ther convex nor concave) walls. grow proximity. a single species wall of the attached is that of the direction of is part fairly thick, coiling (sinistral or dextral) fairly

the lumen is circular. with the of the Neomicrorbis erect part moderately thin; constant exception genus

of The tube orna- The tube wall is chalky white, and consists and a few species of other genera.

of the is well The when parable and cylinder layer. The inner part ment developed. operculum, parable layer is well developed, 0.5 to 3 times (at the known, is invariably calcareous.

1 1.5 times thick as the — Rotularia, Conorca and other keels), generally to as cylin- Comparison genera

is also form but their der layer. Between these two layers there a lami- always or occasionally spirals,

white initial and often As nated very thin bright layer (PI. 7, Fig. 11c, parts are larger well-preserved.

which the The inner mentioned above, in the Ser- top), over wall easily loosens. placement family striation. The is for these surface does not show a transverse pulidae preferred genera.

cavities. — ?, - base lacks cells and longitudinal Stratigraphic range

A single specimen (from Geulhem, GPI Hamburg Recent.

— Neomicrorbis 1904a Collections) shows the characteristic structures Remarks The genus Rovereto, that there caused by the hydroid symbiont Protulophila gestroi is here included in spite of the fact are

the soft of this and Rovereto, 1901. differences between parts genus

Discussion — A single specimen (PL 7, Fig. 14a-c) is of'normal' Spirorbidae (see Zibrowius, 1972).

have existed since the Cam- here illustrated, on account of a circular operculum- Spirorbidae perhaps

and with cer- like object with fine, concentric and radial orna- brian (Lommerzheim, 1979, p. 166) the since the Ordovician ment that occupies the lumen near the aperture tainty (Ruedemann, 1934).

of in marine the (PI. 7, Fig. 14c). The operculum of Recent species They are common many strata up to

The tube subse- late Triassic and also in fresh and brackish Placostegus is not calcareous. was water from the quently intentionally broken, and the alleged oper- settings early (Schweitzer, 1983)

the late Triassic authors culum turned out to be a fragment of an echinoid to (Kelber, 1986). Many

attention these non-marine spine. have focused their at

Comparison — Placostegus alatus is larger and has alae, spirorbids, for post-Triassic and Recent representa-

well. marine. In the Palaeozoic most its tube structure being different as Placostegus tives are exclusively

has but there also dextral tridentiger (Regenhardt, 1961) a sharp triangular species are sinistral, are spe-

keels from the onwards. cross-section, and occasionally the are cies Palaeozoic

somewhat undulate and irregular. P. aduncus Many of the Palaeozoic and Triassic spirorbids

ribs of them could (Regenhardt, 1961) is larger, and transverse are are poorly known in detail; some even

its of variation wider. P. indis- be rather than worms & common; range being gastropods (Burchette

has have been described from tinctus (Briinnich Nielsen, 1931) more distinct Riding, 1977). Opercula

view of I growth lines, while P. dentatus (Briinnich Nielsen, the Cretaceous onwards. In this, agree undu- with Lommerzheim in the old 1931) has strongly denticulate edges, and P. (1979, p. 167) using has Microconchus 1839 for Pal- liferus (Briinnich Nielsen, 1931) more strongly genus name Murchison,

ornamented edges. aeozoic spirorbids again, instead of uniting them

and distribution — Albert Canal with the Recent 1800. Trias- Geographic stratigraphic genus Spirorbis Daudin,

the Geulhem Member: fre- sic be in sections, lower part of spirorbids should treated the same way.

Geulhem There is distinct break in quent, upper part, very frequent. section, a spirorbid phylogeny

of the Geulhem Member: from the to the Cretaceous, lower part very frequent, early Jurassic early

when have not upper part, common. Spirorbidae entirely disappeared,

— and Occurrence elsewhere Danian of Denmark and but have become much rarer not so widely

northern Germany (in erratic boulders). distributed [with the exception of the common spe-

cies planorbiformis (Miinster in Goldfuss, 1831)] in

which the Family Spirorbidae Pillai, 1970 comparison with Serpulidae, during

di- Jurassic become distinctly commoner and more

— ousted the Diagnosis [of tube and operculum] Tube with a very verse. Perhaps Serpulidae spirorbids

caused the small, fragile and generally unpreserved non-spiral from their marine habitats. What disap-

of few millimetres of freshwater is still a of initial part, followed by a spiral pearance spirorbids matter

diameter only. Generally the tubes do not inter- conjecture.

often From the Cenomanian onwards, twine to form clusters, but many specimens early Spiror- 102

have become diverse and The oldest known of Neo- bidae common again. stratigraphically species

undescribed from the Lommerzheim Cenoma- microrbis is an as yet species (1979, p. 167) compared Tithonian coral limestone of Nattheim (SW nian spirorbid species with the Recent taxa Spirorbis early

and which resembles the late Cretaceous N. sensu stricto, Pileolaria, Metalaeospira, Eulaeospira Germany),

crenatostriatus crenatostriatus in size, ornament and Janua, and stated that in reality his Cenomanian

did of these. In numbers of sinistral and dextral individuals, species probably not belong to any equal

but unknown. addition, Bipygmaeus and Neomicrorbis are already its operculum is yet

in Cenomanian. definite be stated on a present the Nothing can single speci- Hauterivian of Unfortunately, the succession of the first fossil men from the Heligoland (Stühmer

known of Recent et 21, occurrence (as to date) genera (or al., 1982, pi. fig. 10). the late Cretaceous Neomicrorbis is similar fossil genera) is not in accord with the phy- During fairly and logenetic scheme elaborated by Pillai (1970) and common. Its tube ornament is very variable,

Zibrowius based there transitions between the complemented by (1972), on are many extremes.

remarked that it was research into Recent material (for details see below). Jager (1983, p. 121) already

quite possible that the three late Cretaceous 'spe-

Genus Neomicrorbis Rovereto, 1904a (= Spiror- cies' crenatostriatus (Miinster in Goldfuss, 1831), hage-

bula Howell, 1962) nowii Jager, 1983 and subrugosus (Miinster in Gold-

fuss, 1831), which usually are easily distinguished by

— in Gold- their I now Type species Serpula crenatostriatus Miinster ornament, are not really good species.

them varieties of fuss, 1831. regard as ecological or subspecies — crenatostriatus. Some obser- Diagnosis [of tube and operculum] Spiral relatively a single variable species,

vations Recent this view: large (diameter about 5 mm), within a single species on Spirorbidae support remarked 'Tubes or subspecies sinistral and dextral specimens occur Knight-Jones (1978, p. 234) that, lack in about equal numbers. Tubes are either smooth or that usually bear longitudinal ridges commonly

them in the of ornamented with longitudinal rows of spines or younger stages growth (Crisp, Bailey

granules, longitudinal keels, transverse wrinkles, & Knight-Jones, 1967; Knight-Jones, Knight-Jones

and lack them funnel-shaped peristomes. & Llewellyn, 1974) may throughout

life in reduced The operculum is entirely calcareous, bilaterally areas of water movement (Gee,

This observation conflicts with observations symmetrical, and consists of a more or less massive, 1964).'

and less late Cretaceous of which N. concave or convex cucullus a large, more or on Neomicrorbis, cre-

keeled calcar. natostriatus subrugosus appears to have preferred a

— than the Comparison Nearly all other Spirorbidae are much higher energy setting longitudinally orna-

smaller. In the of the direc- mented N. crenatostriatus. great majority species c.

within In the anterior tube tion of coiling is constant a species. some Neomicrorbis specimens

— Recent Tithonian-Recent. becomes erect like a corkscrew. Stratigraphic range Early part Spiror-

— soft Remarks Neomicrorbis, which has serpulid-like bidae show a similar intraspecific variation (Knight-

thoracical and 'The terminal coils of parts with relatively many segments Jones, 1978, pp. 234, 235):

tubes ascend become when in dense a spirorbid-like tube and operculum (Zibrowius, may or evolute,

filaments could be considered to an inter- or covered algal or 1972) represent aggregations ... by

In mediate, close to the common ancestor of the fami- encrustations (Gee & Knight-Jones, 1962). deep

'It of the tube lies Serpulidae and Spirorbidae: is certainly a silty areas most can become erect link between the two families ...' (Knight-Jones, (Knight-Jones, Knight-Jones & Bregazzi, 1973; Bai-

stated & 1981, p. 197). Although Jager (1983, p. 120) ley-Brock Knight-Jones, 1977).' of N. that Neomicrorbis should therefore occur already in The taxonomie position fawarensis from the of Israel is pre-Cretaceous strata, it has still not proved possible Avnimelech, 1941 Campanian

in uncertain the of variation of the direction to demonstrate their occurrence units predating as range N. the early Tithonian. Thus from the fossil record it of coiling is unknown; it may perhaps be c.

that this is offshoot appears genus a relatively young subrugosus.

be With ornamented of the Serpulidae. The scenario could then as regard to longitudinally speci-

in Palaeocene of New Zealand follows: spirorbids could either have descended mens: the there are

individuals that from N. cre- fromjurassic or Cretaceous species of Neomicrorbis or are indistinguishable c.

Triassic natostriatus In the Dan- from Palaeozoic or species of Microconchus. (see Fleming, 1972, fig. 4d). 103

ian of Denmark and the tube rather this northern Germany (erratic part erects abruptly; near-verti-

similar N. boulders) some specimens are also very to cal tube part is distorted and slightly curved and

but in others in the number 3.5 The is c. crenatostriatus, changes only up to mm long. aperture generally and of elements of strength longitudinal ornament flaring as are the peristomes, reaching a maximum

[e.g. in N. serratus (Briinnich Nielsen, 1931)] or in the tube diameter of 1.4 mm.

take that At the end of the there shape of the spiral (see below) place, so a spiral are 5-7 longitudinal taxonomic is called for. and 13 the which thin separation keels, up to at aperture, are

Within the group of prevalently transversely and low, but fairly distinct and show a tendency to

there is from the become divided ornamented specimens, a change up into rows of minute forward- late the Danian N. teeth. Cretaceous N. c. subrugosus to directed

The is unknown cingulatus (see below). operculum as yet.

Jager (1983, p. 122) stated that an operculum Discussion —,N. parietalis s. str. from Fakse and p. jagti

showed from the Danian of Kagstrup (Denmark) no (see below) from the Maastricht-Aachen-Liege area

differences to from the late Cre- but identical. For significant opercula are very similar, not some speci-

number of northern German taceous from a mens an assignment to either subspecies may be localities. the cucullus but the However, of the operculum difficult, 'populations' as a whole show sev- of the new described below is concave as eral differences. The be subspecies two taxa may geographic

this in the Recent species, N. azoricus, in contrast to subspecies, and are differentiated at that level. the convex cucullus ofN. crenatostriatus. But, because of the specialised shape of spiral in the new species, Neomicrorbis parietalis jagti n.subsp. this is probably not the direct ancestor of the Recent PI. 9, Figs 1-7

but rather offshoot. species, an

As to Eocene and Recent species (see Jager, 1983, Derivatio nominis — In honour ofJohn W.M. Jagt, in

of his the late Cre- p. 122) no new data are available. recognition publications on

Neomicrorbis shows wider of varia- an even range taceous and Palaeogene of the Maastricht area. tion in the Palaeogene than in the late Cretaceous. Type — The complete tube illustrated in PL 9, Fig.

If the Eocene species described by Rovereto (1904a) 1, is designated holotype. The small dextral initial from the locality Kressenberg (Bavaria, Germany) spiral of 2.8 mm in diameter has seven longitudinal

then this like followed 2 are true Neomicrorbis, genus, Rotularia, keels, by a barely mm long attached would have lost its about the intermediate tube and vertical anterior significance at part a part

Eocene/Oligocene boundary, but unlike Rotularia, erecting 4 mm above the base, reaching 1.8 mm in

is still diameter its end and with Neomicrorbis extant. at a flaring aperture

surrounded by sixteen slightly protruding longitudi-

Neomicrorbis parietalis (Brünnich Nielsen, nal keels. Geulhem Member, 0-0.4 m above the

1931) K/T boundary, Albert Canal sections north of

Vroenhoven Natuurhistorisch Museum 1931 Brunnich bridge, Spirorbula parietalis Nielsen, p. 105, pi. 2, figs 30, Maastricht NHMM 1992204-1 31. Collections, (includ-

eleven ing paratypes, leg. J.W.M. Jagt).

— Brunnich illustrated but — The tube small initial Type Nielsen (1931) a Diagnosis has a spiral,

and this not It which is followed single specimen, was designated type. by a short, attached intermediate is from the middle Danian Limestone of Bryozoan part, growing laterally away from the spiral, and

and is of the collections of erected anterior tube Fakse (Denmark), part finally by a long steeply part. the and Museum of There Mineralogical Geological The umbilicus is usually closed by a callosity.

six Copenhagen University. are to seventeen narrow, sharp, weakly denticu-

— is based Description The following description on late longitudinal keels, and weak flaring peristomes

material. The tube forms initial and The tube wall topotypical an a flaring aperture. apparently

of 2 mm in diameter. Sinistral consists of inner planar spiral up to an parable layer only (a cylinder and dextral in about numbers. is The spirals occur equal layer not preserved). operculum has a con-

The umbilicus is rather and closed narrow usually cave cucullus.

the the tube remains — The tube has and is by a callosity. Following spiral, Description a planar base, attached to the substrate, but the contact with the attached by a moderately broad border on a large,

whorl is lost. After this short intermediate which in previous planar substrate, most specimens is pre- 104

did served as an impression only. In rare instances Weak flaring peristomes, which often are not

In than lines an oyster shell serve as a substrate. contrast to stronger growth are common (PI. 9, Figs

late Cretaceous of the many specimens present 4-6).

attachment thin The cross-section is in the genus an on cylindrical algal (?) inversely U-shaped

attached circular in the and the filaments has not been observed. part, erect part at

initial The tube wall The tube has an planar spiral (PI. 9, Figs 1, aperture (PI. 9, Figs le, 2, 4c, 5b, 6b).

with 2, 4-6) of generally 3 (maximum 4) turns and a is moderately thick, and the lumen circular. rather which is closed The tube in view of the fact that the narrow umbilicus, usually by structure:

The final diameter of is not the tube to a callosity (PI. 9, Figs 4b, 5b). cylinder layer preserved, appears the is Of 130 well- consist of inner spiral 2.5-3.7 mm. sufficiently an parable layer only. The inner preserved specimens fifty are sinistral (PI. 9, Fig. 6) surface is frequently somewhat constricted by sharp, and eighty dextral (PI. 9, Figs 1-5). backward-directed rings, in irregular distances,

the tube remains attached which continue the Following the spiral to as flaring peristomes from out- the substrate, but the curvature diminishes dis- side into the tube's interior (not illustrated for the

whorl tinctly, so that the contact with the previous present species, but see PI. 9, Fig. 9, N. cingulatus). of the is lost The The sole available of the spiral (PI. 9, Figs la, 2, 4a, 5a, 6a). operculum present spe-

of this intermediate tube does cies found in situ in length part usually (PI. 9, Fig. 7) was a typical, not exceed the spiral diameter. dextral tube. The total length of the operculum is

The transition to the erect part is rather abrupt, 0.8 mm, its cucullus (diameter a mere 0.5 mm)

socle in at the end of the attached part occurs a (PI. 9, being circular outline (PI. 9, Fig. 7d) and slightly

The lateral of the base to in the Figs 2, 4c). parts protrude concave, a near-perpendicular position to form two edges between the base and the underside axis of the calcar. This (PI. 9, Fig. 7a-c) is slender of the small below with sides and The erect part, constructing a cavity near-parallel slight curvature.

in lateral alae and carina the erect part (similar to the one seen Pyrgopolon). two the interna are well

anterior tube whereas the carina be The part (PI. 9, Figs la, b, d, 2, 3, developed, externa appears to

5a, b, 6a, b) at first becomes erect under an angle absent.

into Discussion — In remains of the usually in excess of 45°, then generally curves a many samples, spiral

above the base. tube vertical position and grows to 5 mm part are remarkably rare, most specimens

It is distorted and curved in the direc- of the slightly same being fragments erect part. They appear to tion of the older have been water movement coiling as spiral (PI. 9, Figs la, 3). transported by away

The is often from habitat. aperture flaring (PI. 9, Figs lb, c, e, 3) the original

the the oscitans better as are peristomes (of proles type, see Neomicrorbis parietalis jagti was obviously

the increase in action than Regenhardt, 1961). At the aperture adapted to current was N. c. cre- the tube's inner diameter exceeds that of its which lower outer natostriatus, preferred energy settings diameter. and which is absent from the tuffaceous chalk facies

The tube diameter increases moderately rapidly of the Maastrichtian type area (however, it does

in in the The of in the attached part, and rapidly the erect part, occur Gulpen Formation). return the where it reach 1.8 Neomicrorbis above the K/T in may up to mm. genus just boundary

The keels this is longitudinal are similarly shaped (PI. 9, area striking.

but the — resembles miniature Figs 1-6), they are situated mainly on upper Comparison N. p. parietalis a

their lower tube part, whereas the sides, especially version of the present subspecies. The general shape

fewer keels: six nine be of the tube of these is but halves, bear or no to may two taxa near-equal, counted at the end of the spiral, twelve to seventeen length and diameter of tube of the former remain

the their number much smaller. N. does at aperture. During ontogeny parietalis s. str. not represent increases by insertion of new keels between previous juvenile individuals of N. p. jagti, as, even when

of addition tubes with anterior ones, and, at the base the erect part, by complete erect parts are com-

keels the lower halfof the tube. the diameter of the and of several new on pared, spiral part the length

is suture-like furrow of the intermediate in There a narrow longitudinal on part are considerably smaller

In the underside of the erect part, with the neighbour- the former. contrast to the present subspecies, the

the number of is than ing keels being more widely separated than dextral spirals only slightly higher

keels the number of sinistral all. other keels. Rarely do the slightly protrude at ones or not at Moreover,

keels in the aperture (PI. 9, Fig. 1). the longitudinal parietalis s. str. are not only 105

— with low fewer in number, but also distinctly thinner and Description The tube starts a planar to lower, and denticulate. However, the absence of inversely conical spiral (PI. 9, Fig. 8), attached with denticulationin be the result of abrasion broad border to sub- p. jagti may an occasionally relatively large during transport in a high-energy environment. strates (here: planar bryozoan colonies and other

is intermediate individual of the In N. c. crenatostriatus, there no same species [PI. 9, Fig. 8b, small

attached tube between the and lower small sub- non-spiral part spiral specimen right-hand side]) or to

thin branch in the sinistral Fakse the erect part, and the cucullus of the operculum is strates (a bryozoan

In Sinistral and dextral The convex and much more massive. N. serratus specimen). spirals occur.

and weaker lines has 3'/2-4 and (Briinnich Nielsen, 1931) stronger spiral turns a narrow, deep umbilicus,

within which is filled The of longitudinal ornament co-occur a single probably not by a callosity.

' diameter of the reaches 5 in tube. Spirorbula’ multilineata Briinnich Nielsen, 1931 spiral at most mm

' wide while Brün- and to 4.5 in the Maastricht- has a umbilicus, Spirorbula’ ampla Denmark; up mm

In nich Nielsen, 1931 has a thinner tube wall. the Aachen-Liege area. The tube diameter increases

the Recent of operculum of species, N. azoricus gradually to rapidly to a maximum 2 mm.

the cucullus is also but in At the end of the the tube often forms Zibrowius, 1972, concave, spiral a

the axis of the small socle and becomes almost an oblique position to longitudinal erect very steeply,

and calcar, the latter being wider, having a triangu- vertically (PI. 9, Figs 8, 9a). This erect part is cork- lar outline. screw-shaped and continues the directionof coiling

— of the In the Geographic and stratigraphic distribution Albert Canal spiral. Maastricht-Aachen-Liege area,

of the Geulhem Member: this corkscrew be to 2.5 mm and raise sections, lower part very may up long

Geulhem the 4.3 mm above the substrate frequent, upper part, common. section, aperture up to (PI. lower of the Geulhem Member: not In Denmark the erect be part rare, upper 9, Fig. 8). part may even

At the Albert Canal sections part, extremely rare. considerably longer. there is maximum 1.50 does Trans- a frequency approximately Longitudinal ornament not occur.

above the K/T in fossiliferous lenses consists of several to m boundary, verse ornament many small,

hardouini but with the spines of the echinoid Tylocidaris narrow sharp flaring peristomes (PI. 9, Fig. 8c,

In there fine (Desor, 1855). d). some specimens are transverse

Occurrence elsewhere — This sub- wrinkles well The cross-section is is a geographic as (PI. 9, Fig. 8).

species of N. parietalis parietalis (Briinnich Nielsen, inversely U-shaped in the spiral, and circular in the

from the Danish 1931) and southern Swedish erect part (PI. 9, Fig. 8b), the lumen being circular.

Danian. Thetube wall is moderately thick, the tube structure

(PI. 9, Fig. 9) resembling that of N. parietalis jagti (see

above). Neomicrorbis cingulatus (Brünnich Nielsen, Discussion — In both specimens figured by Briinnich

1931) Nielsen (1931) the spiral is dextral. In view of the PI. 9, Figs 8, 9 fact that sinistral tubes of the present species have so

far been Lommerzheim not described, (1981, p. 42, 1831 Miinster — non Serpula cingulata Goldfuss, p. 233, pi. 69, fig. 3) referred to fifty-eight small dextral tubes of fig. 4a-c.

1926 Br. N. — 166. similar from the Palaeocene of the Serpula (Spirorbis) cingulata 0dum, p. shape Emperor

Brunnich 1931 Nielsen, 111, pi. 3, figs Seamounts ‘Circeis - Spirorbula cingulata p. as ? sp. (cingulata [Nielsen]

24. 23, Recent of Circeis dextral Wuchsform)'. species are — 1981 Circeis ? - Lom- non sp. (cingulata [Nielsen] Wuchsform). and possess an operculum of characterisdc shape, as merzheim, 42, 3. p. fig. unknown from N. 1983 122. yet cingulatus. Neomicrorbis cingulatus (Nielsen 1931) —Jager, p. in the Fakse However, a sample from type locality

Type — The specimen illustrated by Brunnich supplied by Mr and Mrs Krause (Wennigsen), there

Nielsen (1931, pi. 3, fig. 24) is here designated lec- is a fine and characteristic but sinistral specimen of

It has and N. the five from the totype. a relatively small dextral spiral a cingulatus. Amongst specimens long corkscrew-shaped erect anterior part with Maastricht-Aachen-Liege area, two (although frag-

Middle and also sinistral. many narrow but sharp peristomes. Danian, mentary dubious) specimens are

sinistral and dextral Coral Limestone, Fakse (Denmark). Mineralogical Thus, specimens appear to co-

in about numbers in Denmark well and Geological Museum of Copenhagen University occur equal as as

Collections. in the Maastricht-Aachen-Liege area, and, more- 106

the tubes of Remarks — All known of B. over, cingulatus are comparatively large European specimens

sinistral. Thus it is for 'normal' spirorbids, and distinctly larger than pygmaeus are doubtful whether

Lommerzheim's Seamount dextral from the middle Palaeocene Emperor specimens. ‘Bipygmaeus’ sp.

the of the Therefore, present species is assigned to Neo- Emperor Seamounts described by Lom- merzheim microrbis for the time being; Lommerzheim's speci- (1981, p. 52) really belong here.

another Circeis. As mens belong to genus, possibly to

definite long as opercula are unknown, a assignment Bipygmaeus pygmaeus (von Hagenow, 1840) cannot be made. PI. 6, 9, ?7, ?8. Lommerzheim stated that Figs (1981, p. 42) correctly

the corkscrew-shape of the erect tube part might be 1840 667. Serpula pygmaea von Hagenow, p. in several Recent as a reaction to present spirorbids, 1983 v Bipygmaeuspygmaeus (Hagenow, 1840) —Jager, p. 132,pi

environmental conditions bottom or (muddy 16, figs 13-15. incrustation by algae), which makes the corkscrew-

into low value. The — A been shape a feature of diagnostic Type lectotype has not yet designated.

Denmark — All from the few specimens of the present species from Description specimens Geulhem

and from the show Member Maastricht-Aachen-Liege area a are attached by a broadborder to a planar

wide of variation: the be range erect part may long or curved substrate (mostly a calcareous crust or a

but also other or short or even absent. The same holds for late bryozoan colony, on secondary hard-

Cretaceous species of the genus (Jager, 1983, pi. 15, grounds), but not to thin cylindrical algal filaments.

4 with the other attached the compare fig. figures). However, Occasionally, specimens are to same

keeping in mind that the corkscrew-shape of the substrate in close proximity.

tube is facultative it aid — Almost all less erect part a feature, may specimens are more or planar

in combinationwith size, ornament and direction of sinistral spirals (PI. 6, Figs 7, 8) with approximately

— in the from moderate coiling distinguishing present species 3 turns and with a to fairly wide umbil-

other spirorbids and serpulids. icus. The diameterof the spiral is generally 1.0-1.4

— Neomicrorbis crenatostriatus is the tube diameter at the Comparison subrugosus mm, aperture being 0.4-0.5

similar, but this has generally fewer and weaker mm. A distinct ornament is entirely missing. Many

Circeis ? - their but peristomes. sp. ( cingulatus [Nielsen] specimens start raising apertures, do not

1981 is smaller Wuchsform) sensu Lommerzheim, lose contact with the previous whorl of the planar

and dextral. In Recent of Circeis This be the of exclusively species spiral. appears to beginning the coni-

the transverse ornament is weaker or absent. cal which is spiral, very characteristically developed

Geographic and stratigraphic distribution— Albert Canal in late Cretaceous specimens.

lower of the Geulhem Member: in sections, part very In the Geulhem Member, however, only a

absent. Geulhem lower rare, upper part, section, single specimen (PI. 6, Fig. 9a-c) is the planar spiral

of the Geulhem Member: part very rare, upperpart, (diameter 1.3 mm) followed by a well-developed absent. conical of spiral only 3/4 turn. This spiral grows

Occurrence elsewhere — Danian of Denmark and upwards by diminishing the diameter of the spiral southern Sweden. and by closing the umbilicus. The height of the

conus is at least 0.8 mm (aperture damaged). The

Genus 1961 ofthe Bipygmaeus Regenhardt, sides conus are near-straight, neither concave

and the between nor convex, suture the turns is a

Type species — Serpula pygmaeus von Hagenow, 1840. narrow indistinct line. This is the only specimen

— sinistral. Tube with Small, starts a from the Geulhem Member that be identified Diagnosis may

which rests a conical with planar spiral, upon spiral beyond doubt.

an followed a cork- Discussion — In the late upward decreasing diameter, by Cretaceous, many adult

erected anterior of the well- screw-shaped, nearly vertically specimens present species possess a part. developed, near-vertical corkscrew, but in the Geul-

— is similar Proto- Comparison Bipygmaeus to Recent hem Member none of the specimens available does,

and and has laeospira, Paralaeospira Helicosiphon, which are only one a characteristic conical median shown be in Pillai's tube to relatively primitive (1970) part. The great majority of the Geulhem Mem- scheme. ber has specimens a simple planar spiral lacking

— Cenomanian-Danian. distinct Range Early ornament. These may be juvenile individ- 107

characteristic fea- often form teeth at the The of uals, but, in the absence of any aperture. operculum brood ruled that of them Romanchella lacks a chamber. tures, it cannot be out some

close — Cenomanian middle actually belong to different taxa. There is a Stratigraphic range Early ?,

lack of Recent. resemblance to Pileolaria ? sp. 1, the orna- Santonian-Neogene ?,

distinctive feature. Remarks — Pileolaria to the most progressive ment being the only, but striking, belongs in Pillai's scheme Thus the question arises whether the planar spirals Recent genera (1970, p. 148).

B. Pileolaria Thus caution is called for in Cretaceous are more closely related to pygmaeus or to referring

whether third taxon be involved. to this as long as their opercula ? sp. 1, or a may species genus

them remain unknown. The status of the two early An argument against interpreting as juveniles

that in with raised Cenomanian Pileolaria)’ species of B. pygmaeus is some specimens ‘Spirorbis (aff.

is distinct furrow between the recorded Lommerzheim (1979) is uncertain: aperture, there a very by

in is and its ornament rather weak, whorls, which is never developed typical vastus very large articulatus is dextral and has flaring specimens. strong,

Comparison — Late Cretaceous representatives of B. peristomes.

from attached kronsmoorensis 1983 from the mid- pygmaeus northern Germany not only ‘Spirorbis’ Jager,

filaments. The dle and Maastrichtian to hard substrates, but also to algal Santonian early (a single

the to somewhat size from area, planar spirals may grow a larger specimen Maastricht-Aachen-Liege

2 and of 4 Meerssen be resembles (diameter slightly over mm up to turns; Member, may conspecific)

Geulhem Member. All and a the from the are see Muller, 1966, p. 1055) may possess specimens

in individuals. The sinistral and three keels. A Pil- rounded longitudinal edge some possess longitudinal

in the is much and consists of eolaria-like also occurs conical spiral more common species commonly

Kazachstan 2'/2 instead of Maastrichtian of northern (material up to turns (Muller, 1966, p. 1055) Prof. Dr D. It is uncertain only 3/4. presented by Naidin).

' 1 is far whether macrotuberculatus Pileolaria ? sp. closely comparable as as Spirorbis (Neomicrorbis)’

size, broad border, direction of coiling (exclusively Miiller, 1966 from the early Maastrichtian belongs

because its sinistral, usually planar spirals with only slight erec- to the present genus, spines are unusu-

conical fol- tion of the aperture, very rarely spiral ally strong.

but has three The Cainozoic species belong to lowing planar spiral) are concerned, following may

the but in need of detailed distinct longitudinal keels. present genus, are

' — Albert Canal studies: Briinnich Nielsen, 1931 Geographic and stratigraphic distribution Spirorbula’ expansa

' lower the Member: Danian of sections, part of Geulhem very (middle Fakse, Denmark), Spirobis’

Geulhem section, lower conoideus Lamarck, 1818 (Eocene), ‘S.’ conoideus rare, upper part, frequent. 1904a the Geulhem Member: Lamarck var. Rovereto, (Lutetian), part of extremely rare, gouetensis

absent. ‘S.’ Rovereto, 1904a (Helvetian), ‘S.’ spirin- upper part, dollfusi

Occurrence elsewhere — Middle Santonian, early tortus Rovereto, 1898 (Helvetian), ‘S.’ declivis Reuss,

Maastrichtian-Danian of northern Germany and 1860 (Tortonian) and ‘S.’ catagraphus Rovereto,

Denmark. 1904a (Pliocene).

Pileolaria ? sp. 1 Genus Pileolaria Claparede, 1868 PI. 6, Figs 11, 12, 14

— attached — 1868. All are or were by Type species Pileolaria militaris Claparede, Description specimens curved sub- — with or without a broad border to a planar or slightly Diagnosis Tube sinistral, longi-

The anterior to a calcareous a bryozoan tudinal and/or transverse ornament. strate (mostly crust, but thin tube ascend. The has a brood colony or an oyster shell, etc.), not to part may operculum filaments. chamber, for details see Knight-Jones (1984, p. cylindrical algal less sin- 111). Nearly all specimens are more or planar with 3 — of the Recent istral approximately Comparison Most species genus spirals (PI. 6, Figs 11, 12) wide umbilicus. The 1897 are also sin- turns and with a moderately Romanchella Caullery & Mesnil, As in diameter of the is 1.1-1.6 that of the tube istral, but several species are dextral. Pileolaria, spiral mm, Three in but the 0.4-0.5 mm. distinct, longitudinal keels are developed some species at aperture very and On keels in the smooth, non-granulate non-denticu- not in others. average, are stronger generally

keels keels in the latter and more late, and strong longitudinal equal former than in genus, equally 108

less but in distances from each other. Keels do not form teeth are more or planar at first, most speci-

the the tends without at aperture. mens aperture to ascend, forming

in does the Many specimens raise their apertures, but do not a cone. Only one specimen apertural whorls and lose contact with the previous whorl of the planar region lose contact with the previous

In the 0.7 above the substrate. spiral. a single specimen (PI. 6, Fig. 14) planar erects mm

is followed conical Two three smooth spiral (of 1.2 mm diameter) by a to weak, to faintly granulate

of V2 This keels occur. The tube'ssurface is some- spiral only turn. spiral grows upwards by longitudinal diminishing its diameter and closing the umbilicus. what curved, thus the median keel lies on a slightly

The of the is level than the other The inner keel height cone at least 0.6 mm (aperture higher two. may

of the be absent. Fine wrinkles be damaged). The sides cone are near-straight, transverse may present,

the between the the wrinkles the median keel neither concave nor convex, suture slightly protrude at

indistinct line. and form small the The turns being a narrow may a lip over aperture.

— with but tube wall is thin. Discussion For a comparison very similar,

Discussion — In unornamented, Bipygmaeus pygmaeus, see above. some specimens, discrimination

— In Pileolaria ? 2 the keels between Pileolaria ? 1 and 2 is difficult. Comparison sp. (see below) sp. sp.

weaker and often whereas — Recent are indistinct, transverse Comparison Amongst species, Nidificaria

The tube is but nidica and ovata wrinkles do occur. slightly smaller, (Knight-Jones, 1978) Simplaria (Vine,

somewhat The tube wall is both the becomes higher. appar- 1977), previously assigned to present ently thinner. genus, are similar, although the latter is larger (coil

In 'Spirorbis’ kronsmoorensis Jager, 1983 the keels diameter 2.0 mm).

— often bear granules, whereas they are smooth in the Geographic and stratigraphic distribution Albert Canal

lower of the Geulhem Member: present species. sections, part very

lower Amongst Recent species, P. quasimilitaris (Bailey, rare, upper part, not rare. Geulhem section,

be indistinct and of the Geulhem Member: 1970) (keels may or absent) part absent, upper part,

Nidificaria pocillator (Vine, 1977) (previously referred absent.

teeth Occurrence elsewhere — Unknown. to Pileolaria, and closely related, keels forming

diameter at the aperture, coil 2.0 mm) are similar.

1977 is The Recent armiger Vine, even Metalaeospira indet. Spirorbidae gen. et sp. and its keel is than larger (3.0 mm), median stronger PI. 6, Fig. 10 the peripheral ones.

Several Recent species of Romanchella (perrieri Discussion — Often the tube surface of small spiror- Caullery & Mesnil, 1897; quadricostalis Knight- bids is damaged to such an extent that only the base Jones, 1973; solea Vine, 1977) have three well-devel- of the planar spiral is preserved. Sinistral spirals four but the keels oped keels as well, or even or five, ? probably belong either to Bipygmaeus pygmaeus or to the form teeth at aperture. Romanchella bicava Pileolaria ? 1 2. sp. or sp. Knight-Jones, 1978 is similar, but dextral. Damaged dextral spirals, however, are here Geographic and stratigraphic distribution — Albert Canal referred because well- to Neodexiospira ? sp., amongst lower of the Geulhem Member: sections, part very small dextral this is the preserved spirorbids genus rare, part, not rare. Geulhem section, lower upper in Member. only one to occur the Geulhem part of the Geulhem Member: absent, upper part, Geographic and stratigraphic distribution— Albert Canal extremely rare. sections, lower part of the Geulhem Member: com-

Occurrence elsewhere — Unknown. Geulhem lower mon, upper part, frequent. section,

of the Geulhem Member: part extremely rare, Pileolaria ? sp. 2 absent. upper part, PI. 6, Fig. 13 Occurrence elsewhere — Unknown.

— The tubes attached Description are either by a small calcareous have Genus 1970 border to planar crusts or Neodexiospira Pillai, (= Dexiospira

1897 become detached of their substrates (bryozoans ?, Caullery & Mesnil, non Ehrenberg, algae ?). All specimens are sinistral spirals with 1859)

3 and with wide approximately turns a moderately

1.0-1.4 and — umbilicus, coil diameter being mm, Type species Neodexiospira formosa (Bush, 1904).

Remarks — & 1897 tube diameter at the aperture 0.4-0.5 mm. Spirals Dexiospira Caullery Mesnil, was 109

the recognised as a junior homonym of Dexiospira Maastrichtian) may belong to present genus.

oldest of Ehrenberg, 1859, probably a protozoan (see The stratigraphically genuine species

Pillai known date is undescribed Knight-Jones, 1984, p. 110). (1970, p. 143) Neodexiospira to a yet

in from the Emael to Meerssen designated formosa as type species; Knight-Jones (Jager, prep.) species

mentioned Members of the (1984, p. 110) erroneously pseudocorrugata Maastricht-Aachen-Liège area,

this the of which is reminiscent of the (Bush, 1904) as type species, being type Recent N. foraminosa

Dexiospira Caullery & Mesnil, 1897. (Moore & Bush, 1904), in having perforations in its

— with brood chambers found Diagnosis Tube dextral in most species. Generally tube, and an operculum

in in situ. longitudinal keels, often three number, occur.

Between them transverse ornament is often seen. Lommerzheim (1981) described Janua (Dex-

cf. The tube wall has longitudinal rows of small per- iospira)’ pseudocorrugata (Bush, 1904), ‘J. (D.)’ qua-

The has siacuta 1981 and bilineata forations in some species. operculum a Lommerzheim, ‘J. (D.)’

'distal with from the Palaeocene. In cf. brood chamber, part of brood chamber (Schmidt, 1951) pseu-

also the is calcified walls' (Knight-Jones et al., 1975, p. 93). docorrugata and quasiacuta operculum

Comparison — The tube of Janua de Saint-Joseph, known.

stricto is but lacks Miocene include lovisatoi 1894 sensu very similar, obviously species ‘Spirorbis’

perforations. Rovereto, 1904a (Tortonian), ‘S.’ heliciformis (von

— late and ‘S.’ bilineatus Stratigraphic range Early Maastrichtian ?, Eichwald, 1830) (Schmidt, 1951)

described Maastrichtian-Recent. (both Sarmatian). Belokrys (1984) quite a

'

Remarks — is of the most number of of which Neodexiospira one progres- species ‘Spirorbis (Dexiospira) ’,

scheme in sive Recent genera in Pillai's (1970, p. 148). probably are conspecific part.

is Pleistocene in the late Maastrichtian there- nevianii Rovereto, 1904a a Its occurrence may ‘Spirorbis’

has been fore be surprising at first, but definite proof species.

furnished by opercula found in situ. ? 1 Neodexiospira sp. time there finds Already in Cenomanian are PI. 6, Figs 15, 16 resembling Recent Metalaeospira and Eulaeospira (see

Palaeocene and — Lommerzheim, 1979). Metalaeospira Description Spiral attached without a border to a

the basis of Circeis have been recognised on opercula planar substrate (mostly calcareous crusts, oyster

bored by Lommerzheim (1981). Metalaeospira, Eulaeospira shells, walls of cavities by bivalves into coarse-

in Pillai's and Circeis are also fairly progressive grained limestones), not to algal filaments. Some-

scheme. times several specimens are attached to the same

Other Cenomanian spirorbids resemble the substrate in close proximity.

stricto. All available small dextral Recent genus Spirorbis sensu Knight-Jones specimens are planar

the Eocene S. ele- 2'A and with (1981, p. 197) compared species spirals with approximately turns a

1827c with the is still in gans Defrance, genus Protolaeospira. moderately wide umbilicus. The aperture

It that the different constructions of the and shows appears contact with substrate no tendency to

Recent their different diameter of the is 0.6-1.0 Spirorbidae, e.g. equipment erect above it. The spiral

for brood have been evolved in protection, might a mm, that of the tube at the aperture approximately

radiation the middle rapid during Cretaceous, 0.2-0.3 mm.

consists 4 5 whereas the evolution of other Recent spirorbid Longitudinal ornament of 3, or

mentioned here could have been slower small but rather keels, the middle genera not (mostly 3) sharp than the during the Cainozoic. one(s) at a somewhat higher level periph-

weak annular I doubt that the Cenomanian species (gamigensis eral ones. Only rarely are peristomes

tube wall is thin and and Geinitz, 1872 and bipartita Reuss, 1845) compared seen. The fragile, no

with the Recent visible. by Lommerzheim (1979) genus perforations are

all — carinatus Miiller, 1964 was Janua are at spirorbids. Comparison ‘Spirorbis’

' attached to small filaments The generic status of Spirorbis’ labiatus Jager, algal or seagrass roots,

is and the tube 1983 (late Santonian-late Maastrichtian) uncer- cross-section is more angular. ‘Spi-

is has 4 5 tain, whereas ‘S.’junior Jager, 1983 (late Maastnch- rorbis’junior Jager, 1983 similar, or keels, a

' but to tian) and Scarinatus Muller, 1964 (junior hom- less rounded, more trapezoid square cross-

carinatus and the ascend somewhat onym of carinatus Daudin, 1800, Montagu, section, aperture may

1803 and carinata Defrance, 1827c) (early above the substrate. Amongst Recent spirorbids, 110

of well the fact that fewer several species Neodexiospira as as Janua pag- discrepancies are explained by

enstecheri (de Quatrefages, 1865) are similar. samples (and fewer specimens) were available from

Geographic and stratigraphic distribution — Albert Canal the Geulhem section.

the Geulhem Member: sections, lower part of not At both localities, the lower unit of the Geulhem

Geulhem lower Member is somewhat richer in rare, upper part, rare. section, part serpulid/spirorbid

taxa. Some taxa from of the Geulhem Member: extremely rare, upper uncommon are absent the

absent. unit P. part, upper [Pyrgopolon (Septenaria) sp., (P.) m. cip-

Occurrence elsewhere — Unknown. lyana, Neomicrorbis cingulatus, and Neodexiospira ? sp. 2).

However, three taxa are distinctly commoner in the

than in the lower viz. Neodexiospira ? sp. 2 upper unit, Rotularia (R.) tobar

PI. 6, Fig. 17 vroenhovensis, R. (R.) tobar gracilis and Ditrupa schlot-

heimi, the latter being extremely common in the Description — On the inner surface of an oyster shell lower unit, but reaching an acme (mass occurrence) dextral a single planar spiral with 11/4 regular in the These three all soft- upper part. species are turns is preserved, its coil diameter being 0.6 mm, bottom dwellers attached tube lacking parts the diameter of the tube at the aperture < 0.2 mm, [Ditrupa), or with only very small attachment areas above the the aperture not ascending substrate. [Rotularia), whereas taxa preferring the lower unit all This spiral differs from all other spirorbids from the attached tube of variable possess an part length. Geulhem Member by leaving a distinct space Thus the differences between the faunas of the two between the resembles the turns, so that spiral a units appear to roughly correspond to changes in heteromorphic ammonite. main lithofacies types: preponderate hard bottom The cross-section of the tube is squarish, with two soft bottom settings below, preponderate ones the left and The sharp edges on upper upper right. above. It should be stressed, however, that both side is concave between these upper slightly edges, lithofacies types may co-occur throughout the sec- lateral sides but do form bor- slightly convex, not a tions studied, and that the fauna of the few hard- der at the base. bottom of the unit is samples upper fairly closely Discussion — The relationship of this enigmatic spec- comparable to that of the lower unit. The imen with Recent genera is puzzling. only rea-

it with the is son for comparing genus Neodexiospira COMPARISONS its mode of coiling, but it should be borne in mind

that other Recent 1. Denmark spirorbid genera are dextral as

be well. In spite of its very regular coil it cannot The for is the most important area comparison type is in fact ruled out that this specimen a juvenile area of the Danian Stage. Seven of twenty-two taxa serpulid. found in the Geulhem Member of the Maastricht- Geographic and stratigraphic distribution — Albert Canal Aachen-Liege area have also been recorded from Geulhem Member: sections, lower part of the Denmark, viz. Cycloserpula lombricus, Rotularia (R.) absent, upper part, absent. Geulhem section, lower hisingeri, Ditrupa schlotheimi, Placostegus alatus, P. of the Geulhem Member: part extremely rare, erectus, Neomicrorbis cingulatus and Bipygmaeus pygmaeus. absent. upper part, Two additional similar Danish taxa are very to Occurrence elsewhere — Unknown. and there small differences specimens are only at

the subspecies level, viz. Cementula applanata subditiva DISCUSSION and Neomicrorbis parietalis jagti. 0dum (1926) and

of the Tables 1 to 3 show a comparison geographic Brunnich Nielsen (1931), who studied the serpulid/

and stratigraphic distribution of serpulids and spi- spirorbid faunas from the Danish Danian, divided

rorbids in the Geulhem Member in the this into units: older and as exposed stage two younger Danian,

Maastricht-Aachen-Liege area. with the latter including the Middle Danian of

The serpulid and spirorbid faunas from the two Fakse. Four of the seven plus two species and sub-

studied almost The sole strik- above localities are identical. species mentioned have been recorded from

difference is the of of the older well from the while ing rareness representatives as as younger Danian, the Rotularia the Geulhem section. In addi- the other five D. P. genus at (R. (R.) hisingeri, schlotheimi, alatus, tion, some extremely rare species are absent from N. parietalisand B. pygmaeus) are only known from the

the other of these consid- one or locality. However, some younger. Bipygmaeus pygmaeus should not be 111

Cycloserputa lombricus Serpula insiticia

Serpula sp. aff. trilineata

Cementula applanata subditiva Rotularia(R.) hisingeri Rotularia(R.)

tobar gracilis Rotularia (R.)

tobar vroenhovensis

Ditrupa schlotheimi

? Serpula K Metavermilia to __ Pyrgopolon (P.)

mosae mosae Pyrgopoton (P.) mosae ciplyana Pyrgopolon (Septenaria) &—.—: Placostegus alatus Placostegus erectus

Neomicrorbis parietalisjagti Neomicrorbis cingulatus Bipygmaeus pygmaeus Pileolaria ? sp.1

PileolariaI 7 sp-2 Spirorbidae, inde gen, et sp. et_

NeodexiospiraI 7 SP.I

'?? Neodexiospira"- Jsp.2

Table 3. Frequency of all serpulid and spirorbid recorded from the Geulhem Member as exposed along the Albert Canal and at

details Geulhem. For see text. 112

in and from the ered in this respect, as it appeared already the Rancocas Formation of Maryland of late Cretaceous. Placostegus alatus is restricted to the Danian or late Palaeocene New Jersey.

Middle Danian of Fakse (Coral Limestone), whereas 10. New Zealand D. schlotheimi, N. parietalis and R. hisingeri have been Fleming (1972) recorded comparable species from recorded from several localities in Denmark, and

the Palaeocene: a dubious Cementula from the latest from other countries (see below). the Teurian-Waipawan, a species of Ditrupa from

Teurian (= Danianand early late Palaeocene) and a 2. Northern Germany species of Neomicrorbis from the Waipawan (= latest Some of the taxa found in Denmark are also known Palaeocene to early Eocene). from northern Germany, exclusively from erratic

here. boulders, which is why they are omitted 11. Emperor Seamounts, NW Pacific

Lommerzheim(1981) recorded several species from 3. Southern Sweden the Palaeocene, some of which were compared with Rotularia (R.) hisingeri, N. parietalis, N. cingulatus and and B. there N. cingulatus pygmaeus. However, are other known from Danian in species are strata in that differences size and direction of coiling, so Skane Nielsen, 1931). (Briinnich rela- this fauna does not appear to have any close

tionship with the Geulhem Member assemblages. 4. Belgium (Mons Basin)

Rasmussen (1965) described and illustrated C. K/T BOUNDARY EXTINCTION, SURVIVAL AND ORIGINA- D. schlotheimi and third from the lombricus, a species TION EVENTS Danian of Ciply (Mons Basin).

Of the twenty-two species and subspecies known to

also 5. Poland date from the Geulhem Member, four have

been recorded from the late Pugaczewska (1967) recorded C. lombricus, D. schlot- underlying Cretaceous, heimi third from the Dano-Montianof viz. C. and a species P.lombricus, Pyrgopolon (P.) mosae mosae, (P). m. the Boryszew borehole. ciplyana and B. pygmaeus. In the Maastricht-Aachen-

Liege area these four species are known from most

6. Austria of the members from the Lanaye Member to the

in From the Danian of Bruderndorf, Kiihn (1960) Nekum Member, and all occur the Meerssen recorded R. hisingeri and two additional species. Member. C. lombricus and B. pygmaeus occur in pre-

Member the Lanaye strata as well. In the Geulhem

the 7. Kazachstan extremely variable C. lombricus display same

diverse Palaeocene from of features it does in the Meerssen Amongst a faunaof species array special as various localities in Kazachstan furnished by Prof. Member, and the two subspecies of Pyrgopolon (P.)

also well with from the Dr D. Naidin (Moscow), are C. lombricus and a spe- mosae agree representatives Meerssen whereas B. shows cies of Rotularia, similar to R. (R.) lobar gracilis, but Member, pygmaeus much and differences. larger more trochospiral, both from the (intraspecific ?)

Danian of Kaurtakapy, western Kazachstan. The status of Serpula aff. trilineata is not quite

certain. It be with late Cretaceous may conspecific

8. Crimea trilineata or not.

and In the E. Voigt Collection at Hamburg University The remaining seventeen species subspecies

from the Geulhem Member have been recorded there is a single specimen of Placostegus, resembling not

from Cretaceous neither in the Maastricht- P. erectus, but at least twice as large. strata,

Aachen-Liege area nor anywhere else. A few of

Danian the 9. United States them are known from the or Palaeocene

Gardner (1935, 1939) described and illustrated sim- of other countries (see above). However, almost all ilar Rotularia of these S. aff. trilineata species, which are herein regarded as species (including ) are closely subspecies of R. tobar, viz tobar tobar and tobar related to other congeneric species, or, in the case of of related mcglameryae, from the Palaeocene Midway Group of Ditrupa/Pentaditrupa to species a closely

and that there real in the Texas, Alabama possibly Mississippi. Gardner genus, so are no newcomers

and discussed several faunas in the with the (1916) Regenhardt (1961) spe- serpulid/spirorbid area onset

of of the Geulhem Member. cies resembling our species of Placostegus from the deposition 113

also within the much advanced Not only on a global scale, but more attempt at providing an over-

Maastricht-Aachen-Liege area are the relationships view of serpulid/spirorbid evolution and extinction between late Cretaceous and Danian serpulid/spi- around the K/T boundary.

of material rorbid faunas close: in the Geulhem Member only With the exception from northern

and the the and the three (sub)species of Rotularia unique Germany, Maastricht-Aachen-Liege area

2 have counter- the Seamounts, the literature specimen of Neodexiospira ? sp. no Emperor comprises

the ? but limited well-documented parts in the Maastrichtian of area. Serpula stratigraphically ser- insiticia and S. differ from Cre- faunas. In addition, modern sp. considerably pulid/spirorbid papers if make the The remain- biozonations men- taceous species referred to genus. describing rarely, ever, related tion of faunas. This is the ing sixteen species and subspecies are closely serpulid/spirorbid why scheme used below is of to Maastrichtian taxa occurring in the area. stratigraphic necessity crude for Thus, the serpulid/spirorbid faunas of the Geul- rather crude, too really a proper descrip-

tion hem Member comprise mainly in character of an extinction event of short duration.

It could be follow Danian Maastrichtian faunal elements that experienced a quite revealing to spe-

and Eocene and slightly higher rate of evolution at the K/T bound- cies into the late Palaeocene, to see

similarities between the faunas of the what at the ary. The happens Eocene/Oligocene boundary;

of the Geulhem data available date for detailed Maastricht Formation and those to are too scanty a

Member in view of the of extinction/survival and are not surprising more or description origination

in these intervals. less comparable environmental and lithofacies con- patterns stratigraphic ditions that prevailed during the deposition of these Genera and in the Maastrich- units. However, a slight impoverishment of the ser- 1. subgenera present but absent from the pulid/spirorbid faunas around the K/T boundary tian, Cainozoic, becoming should be noted: twenty-four species and subspecies extinct either during the course of the Maastrichtian

Meerssen or at the K/T are known from the Member, while boundary:

herein from the Geulhem - Protula 1943. In the United twenty-two are recorded i (Longitubus) Howell,

Member. States, this subgenus ranges from the early Campa-

sections nian Merchantville Formation to the In principle, Danish show a comparable early Maastrichtian Navesink it picture, although environmental conditions and Formation; probably

differed from those of became extinct before the K/T lithofacies types considerably long boundary.

Here Dan- known the Maastricht-Aachen-Liege area. too, Jereminella Lugeon, 1919. Poorly genus, a

Maastrichtian been recorded from the ian lithofacies types closely resemble single species having

'The Danian Maastrichtian of restricted ones. reconstituted, typical commu- a area (see Regenhardt,

be less nities in can considered div- 1961, many respects p. 34).

of erse duplicates Maastrichtian communities Serpula (n. subgen.).

(Hakansson and Thomsen, 1979). It is noteworthy, - Pegmaticula Regenhardt, 1961. Originally intro-

however, that some specialized ecological types, duced as a subgenus of Gynaeconitis Regenhardt,

which Maastrichtian chalk but altered in are important in the 1961, diagnosis entirely (Jager, prep.),

in the Danian' contain and tubes with communities, are virtually lacking to spiral non-spiral a gener-

'smooth' surface and an occasional (Birkelund & Hakansson, 1982, p. 381). ally 'secondary

For detailed discussion tube' Nebenrohre of a more of serpulid/spiror- (= Jager, 1983, pp. 11, 38):

' below. 1961 Parsi- bid faunal changes at the K/T boundary see turpificiata Regenhardt, (type species),

lists below and sub- monies’ an undescribed sub- The comprise not only genera wegneri Jager, 1983,

in the of undescribed from the genera occurring Maastricht-Aachen-Liege species wegneri, an species

and Maastrichtianof and area, but all genera subgenera previously Kazachstan, delphinula (Gold-

Oxfordian-Tithonian recorded from the Maastrichtian and/or Pal- fuss, 1831) of age.

of Parsimonia 1961 ?. It is clear aeocene over the world. These lists are a prelimi- Regenhardt, not

include revised whether this into the as nary nature, and in part (yet genus ranges Maastrichtian, unpublished) data, data in need of confirmation, doubts exist on the status offrustulenta (Regenhardt,

in few data from the literature and, a cases, simply 1961). 1961. repeated here without comment. In comparison Mucroserpula Regenhardt,

- 1979. Discrimination with the works of Briinnich Nielsen (1931), LaqueoserpulaI Lommerzheim,

between this and Cementula is Regenhardt (1961) and Howell (1962), this is a genus not entirely clear. 114

- Nogrobs (Tetraditrupa) Regenhardt, 1961. See Jager (Marinoni, 1878) [unspecified Cainozoic], kaufmanni

for details. and (in press b) (Mayer-Eymar, 1887) [Eocene] ouyenensis

This is detailed studies Ornatoporta Gardner, 1916. either a sub- (Chapman, 1913) [early Miocene];

of but needed. genus Pyrgopolon or a closely related, poorly are 1844 known taxon. Maastrichtian (Monmouth Forma- Placostegus Philippi, (= Eoplacostegus

tion), United States. Regenhardt, 1961).

Paliurus 1876. See Lommerzheim Conorca Regenhardt, 1961. Seejager (1983). Gabb, (1979, p.

- Protectoconorca Jager, 1983. 163). Orthoconorca 1983. - Rotularia (Tectorotularia). Jager, Jager's (1983) interpreta-

tion of this of - Rotularia (Austrorotularia). genus was too wide; a group species

reminiscent of the is known type species to range

Genera and in from the late Cretaceous into the Danian. 2a. subgenera present Maastrichtian

but - Pileolaria 1868. See (or Mesozoic) as well as in Cainozoic, with 1 Claparede, Jager (in prep.).

either side the K/T Pillai, 1970. See different species on of -Neodexiospira Jager (in prep.).

boundary. - Spirorbis Daudin, 1800 ?, Metalaeospira Pillai, 1970

? and Pillai, 1970 ?. Lommerzheim - Protula (Protula) Risso, 1826. Unpublished material Eolaeospira

did than of Hauterivian age from France. (1979, pp. 167, 169, 170) no more compare

Cenomanian material with and refer these - Josephella Caullery & Mesnil, 1896. not to Recent in litera- - Serpula Linne, 1758. genera. Spirorbis palaeontological ___

Pillai ture has often been used for all - Spiraserpula Regenhardt, 1961 emend. (in Spirorbidae gen.

press). indet.

- Cementula Regenhardt, 1961 emend. Pillai (in

press). 2b. Genera and subgenera of which at least one

- Rotularia Lommerzheim, 1979 ?. (Praerotularia) the K/T (sub)species crosses boundary mor- Lommerzheim this sub- According to (1979, p. 174) phologically unchanged (given in [] brackets)

into the Eocene, but I know of no genus ranges Filograna Oken, 1815. [F. sodalis (Goldfuss, 1831)]. Cainozoic species. However, one of the three sub- The occurrence in the Danian of Denmark is in K/T of Rotularia must have crossed the genera need of confirmation. this should Otherwise, genus boundary to give rise to the Palaeogene subgenus R. be in category 2a. (Rotularia). Cycloserpula Parsch, 1956 (= Glomerula Regenhardt,

- de Montfort, 1808. Nogrobs (Nogrobs) Seejager (in 1961 Brünnich sensu Nielsen, 1931). [C. implicata b). press (von Hagenow, 1840)]. ?. See Lommerzheim - Galeolaria Lamarck, 1818 Pentaditrupa Regenhardt, 1961. [P. subtorquata (1979, p. 153). (Münster in Goldfuss, 1831).

- Pomatoceros Philippi, 1844. Discrimination from Vermiliopsis de Saint-Joseph, 1894. [V. dorsolineata Dorsoserpula Parsch, 1956 not clear. (Briinnich Nielsen, 1931) and V. fluctuata (J. de C.

- Metavermilia (Metavermilia) Bush, 1904. See Lom- Sowerby, 1829)]. merzheim (1979, p. 158). Neovermilia Day, 1961 (= Proliserpula Regenhardt,

- Metavermilia (Vepreculina) Regenhardt, 1961. 1961). [N. ampullacea (J. de C. Sowerby, 1829)].

- Filogranula Langerhans, 1884. Pyrgopolon (Pyrgopolon) de Montfort, 1808. [P. (P.)

- Pseudovermilia Bush, 1907 ?. See Lommerzheim de 1808 and mosae mosae Montfort, P. (P.) m. ciplyana (1979, p. 159). (de Ryckholt, 1852)].

- de 1894. See Lommerzheim Janita Saint-Joseph, crenatostriatus Neomicrorbis Rovereto, 1904a. [N. c. (1979, 157). p. (Münster in Goldfuss, 1831)].

- Pyrgopolon (Septenaria) Regenhardt, 1961. Upon a Bipygmaeus Regenhardt, 1961. [B. pygmaeus (von closer inspection of Cainozoic and Recent material Hagenow, 1840)]. of this the the only representatives subgenus are sole

specimen from the Geulhem Member described

herein and possibly gallica (Wrigley, 1951) of early Protulophila gestroi Rovereto, 1901, a hydroid sym-

biont in tubes also the K/T bound- Eocene-early Oligocene age. serpulid crosses

- Pyrgopolon (Hamulus) Morton, 1834. Three Cai- ary and ranges from Pliensbachian to Recent (see

nozoic viz. Zibrowius & in species may belong here, hexagonum Jager, prep.). 115

in the attachment tubes. Several forms 3. Genera and subgenera first occurring area, e.g. spiral

which needed attachment Palaeocene a large area, e.g. many

Recorded with shallow-water became extinct. Hydroides Gunnerus, 1768. a query specialised dwellers,

forms often also from the middle Palaeocene by Lommerzheim These shallow water are warm

recorded Howell forms The decrease in (1981, p. 36); also by (1948, p. 5) water (reef inhabitants). tem-

from the late Palaeocene Vincentown Formation of perature at the K/T boundary generally affected

New Jersey. warm water forms more than it did cold water

Rotularia (Rotularia) Defrance, 1827a. The phy- forms.

logenetic relationships between the four subgenera Thus the Danian serpulid and spirorbid faunas

the the K/T do the Danian faunas recognised is unclear and separation at represent, as bryozoan (Voigt,

boundary is slightly arbitrary. 1985), an impoverished Maastrichtian fauna with

1835. but few Ditrupa Berkeley, very new taxa.

1845. Recorded As result of level and -Pyrgopolon (Turbinia) Michelin, by a sea changes sedimentary

Thanetian Pal- Wrigley (1950) from the (late breaks in the great majority of K/T boundary sec-

aeocene) onwards. tions, these show breaks in the palaeontological and

Circeis de Saint-Joseph, 1894. See Lommerzheim evolutionary record (see Birkelund & Hakansson,

The known Palaeocene is which the number (1981). only species C. 1982, fig. 11), explains great of

Briin- in 2a. paleocaenicum Lommerzheim, 1981; cingulatus taxa category

nich Nielsen, 1931 is here referred to the genus ACKNOWLEDGEMENTS Neomicrorbis.

I wish to extend best thanks to Prof. Dr E. - Paralaeospira Caullery & Mesnil, 1897. See Lom- my Voigt, Dr W. Weitschat and H.-W. Lienau for material merzheim (1981). from the Institut der - Cubiculovinea Lommerzheim, 1981. Geologisch-Palaontologisches

Universitat Hamburg received on loan; Drs J.W.M.

Jagt, H. and I. Krause, Drs R.H.B. Fraaye, Prof. Dr The K/T boundary witnessed the mass extinction D. Naidin and Prof. Dr H. Wulffor of dinosaurs, ammonites, belemnites, inoceramid providing com-

Dr H.A. ten Dr A. Lom- bivalves, planktonic foraminifera, phytoplankton parative material; Hove, Dr T.G. Pillai and Dr H. Zibrowius for and other As far level bottom merzheim, groups. as commu- discussions Recent and fossil the K/T is fruitful on nities are concerned, however, boundary serpulids;

fourth-class and W. and H. for listed only as a extinction event by Tilgner Jager kindly preparing

the for the For critical Boucot prints plates. reading, (1990, p. 22, fig. 1). of the and At the K/T boundary the families that disap- improvement English suggesting

is thanked. A first of all and improvements, J.W.M. Jagt Sachmit- peared were consumers, secondly telbeihilfe received from the phytosymbionts, whereas suspension feeders were (project Ja 385/2-1) Deutsche (DFG) is less affected (Roy et al., 1990, fig. 6). Sheehan & Forschungsgemeinschaft grate-

after breakdown of Hansen (1986) stated that, a fully acknowledged.

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1-121, 2 tabs, 8 pis.

PLATE 1

Cycloserpula lombricus (Defrance, 1827b). Houthem Formation, Geulhem Member.

attached tube lower of the tube’s interior. Fig. 1. Meander-shaped part, part figure showing oblique step-shaped swellings on (broken)

Albert Canal sections, north of km mark 24.000, x 11.5. Coll. GPI Hamburg, no. 3300.

2. Tube three-dimensional chaotic knot of with after of tube the does Fig. forming (normal shape tube) an injury: breakage new part

not follow the direction of the previous part, a - overall view, b - detail showing breakage (upper right), U-shaped new part and

remains of broken old lower Geulhem section, burrow infill the K/T and part (upper right to centre). at boundary, x 5.8 (a) x

11.5 (b). Coll. GPI Hamburg, no. 3301.

Fig. 3. Undulate erect tube part. Geulhem section, burrow infill at the K/T boundary, x 7.7. Coll. GPI Hamburg, no. 3302.

4. 5.8. Coll. GPI 3303. Fig. Curved, fairly long erect part, same sample as Fig. 3, x Hamburg, no.

Fig. 5. Erect tube part, showing breakage and straight new part (upper half figure), same sample as Fig. 3, x 11.5. Coll. GPI Hamburg,

no. 3304.

6. After of the the old rather Albert Canal 11.5. Fig. breakage erect part, new part ’enwraps’ part irregularly. sections, as sample 121, x

Coll. GPI Hamburg, no. 3305.

Closed Albert Canal 15.3. Coll. GPI Fig. 7. rucksack-shaped protuberances (left), sections, as sample 121, x Hamburg, no. 3306.

8. Closed in rather tube. Albert Canal 15.3. Coll. GPI Fig. rucksack-shaped protuberances (right) irregular sections, as sample 121, x

Hamburg, no. 3307.

Broken tube. Canal Fig. 9. open rucksack-shaped protuberances in knot-shaped Albert sections, north of km mark 24.000, x 11.5.

Coll. GPI Hamburg, no. 3308.

10. Broken and Albert Canal x 15.3. Coll. GPI Fig. open rucksack-shaped protuberances (centre right). sections, as sample 138,

Hamburg, no. 3309.

11. Cross-section of thin-walled tube Albert Canal 15.3. Coll. GPI 3310. Fig. a erect part. sections, as sample 121, x Hamburg, no.

12. Two cross-sections of tube of thick-walled - small eccentric six oftube wall visible with crescentic Fig. a knot, a very lumen, layers

interstices in between, b - kidney-shaped lumen. Albert Canal sections, north of km mark 24.000, x 15.3. Coll. GPI Hamburg,

no. 3311.

Fig. 13. S-shaped tube with eccentric lumen and two layers oftube wall with crescentic interstices visible in both cross-sections. Albert

Canal sections, as sample 138, x 19.2. Coll. GPI Hamburg, no. 3312.

Fig. 14. 8-shaped fragment of knot with ’trilobate narrowing’ at both ends (one visible, left), but without ’trilobate narrowing’ in

between. Albert Canal north of km mark 19.2. Coll. GPI 3313. sections, 24.000, x Hamburg, no.

Fig. 15. Cross-section of rather thick-walled tube with ’trilobate narrowing’. Albert Canal sections, sample 127 III, x 19.2. Coll. GPI

3314. Hamburg, no.

Cross-section of thick-walled tube with crescentic ’trilobate Fig. 16. very interstices between the layers (lower third) and with

narrowing’. Albert Canal sections, sample 118, x 15.3. Coll. GPI Hamburg, no. 3315.

with lumen visible in both cross-sections. Albert Canal 23. Coll. GPI Fig. 17. S-shaped fragment kidney-shaped sections, sample 118, x

Hamburg, no. 3316.

Fig. 18. Curved fragment with kidney-shaped lumen visible in both cross-sections. Albert Canal sections, sample 118, x 19.2. Coll. GPI

Hamburg, no. 3317.

Fig. 19. Cross-section of rather thick-walled tube with crescentic interstices between the layers (upper right) and with kidney-shaped

lumen. Albert Canal sections, as sample 121, x 15.3. Coll. GPI Hamburg, no. 3318.

20. Three-dimensional knot with of tube wall - different Fig. outer layer broken, showing corkscrew-shaped inner layer, a-c views of

same specimen. Albert Canal sections, as sample 138, x 11.5. Coll. GPI Hamburg, no. 3319. 121\1

PLATE 1 122

Yorkshire Lias. London Stevens, G.R., 1967. UpperJurassic fossils from Ellsworth Land, Tate, R., &J.F. Blake, 1876. The (J. van

viii + 475 + xii 23 West Antarctica, and notes on UpperJurassic biogeography Voorst), pp., pis.

ofthe South Pacific region. — N.Z.J. Geol. Geophys., 10(2): Vine, P.J., 1977. The marine fauna ofNew Zealand: Spirorbinae

— Mem. N.Z. Inst., 68: 345-393, 45 figs. (Polychaeta: Serpulidae). Oceanogr.

Stimpson, W., 1854. Synopsis of the marine Invertebrata of 1-68, 35 figs, 6 tabs.

about the mouth of the of 1985. The ofthe bound- Grand Manan, or the region Bay Voigt, E., Bryozoa Cretaceous-Tertiary

Ordovi- — In: G.P. Larwood & C. Nielsen Fundy, New Brunswick. Smiths. Contr. Knowl., 6: 1-67, ary. (eds). Bryozoa:

2 pis 1-3 (not seen). cian to Recent. Fredensborg (Olsen & Olsen): 329-342,

Stoliczka, F., 1867-1868. The Gastropoda of the Cretaceous tabs.

rocks of Southern India. — Mem. geol. Surv. India, Pal- Wanner, J., 1902. Die Fauna der obersten weissen Kreide der

xiii 497 28 Wiiste. — 91-152, aeont. Indica, 5: + pp., pis. libyschen Palaeontographica, 30(2): pis

Straughan, D., 1967. Some Serpulidae (Annelida: Polychaeta) 13-19.

— Univ. 1881. of invertebrate fossils from from Heron Island, Queensland. Qld Papers, 1(2): White, C.A., Descriptions new Mesozoic and Cenozoic rocks of Arkansas, 27-45, 5 figs. the Wyoming,

Stuhmer, H.H., C. Spaeth & F. Schmid, 1982. Fossilien Helgo- Colorado, and Utah. — Proc. U.S. natl Mus., 3: 157-165.

fossils from Arkansas lands, Teil 1, Trias und Unterkreide. Otterndorf(Niederelbe White, C.A., 1882. On certain Cretaceous

184 9+19 70 and Colorado. — Proc. U.S. natl Mus., 4: 136-139, 1 Verlag), pp., figs, pis. pi.

PLATE 2

Serpula ? insiticia (Regenhardt, 1961). Houthem Formation, Geulhem Member.

- initial tube Geulhem Fig. 1. Cluster ofthree specimens affected by the symbiont Protulophila, b remains ofnon-spiral part (left centre).

3320. section, burrow infill at K/T boundary, x 7. Coll. GPI Hamburg, no.

b - tube affected a - overall remains of initial tube (left), Fig. 2. Low spiral with erect part, by Protulophila, view, showing non-spiral part

GPI 3321. aperture with peristome. Geulhem section, burrow infill at K/T boundary, x 7 (a), x 14 (b). Coll. Hamburg, no.

Fig. 3. Spiral detached from substrate, bottom view, showing straight initial tube part and structure of tube wall. Geulhem section,

burrow infill at K/T boundary, x 10.5. Coll. GPI Hamburg, no. 3322.

and base Albert Canal Fig. 4. Spiral ofjuvenile specimen, affected by Protulophila, lateral view, showing protruding top at aperture.

10.5. Coll. GPI 3323. sections, as sample 138, x Hamburg, no.

3324. Fig. 5. Spiral with long erect tube part. Geulhem section, burrow infill at K/T boundary, x 10.5. Coll. GPI Hamburg, no.

above K/T 14. Coll. GPI Fig. 6. Erect tube part with three peristomes. Geulhem section, some metres boundary, x Hamburg, no.

3325.

burrow infill K/T 10.5. Fig. 7. End of broken spiral with long erect tube part with one peristome. Geulhem section, at boundary, x

Coll. GPI Hamburg, no. 3326.

8. Erect tube described with three - lateral b - frontal view of Geulhem Fig. part by Regenhardt (1961), peristomes, a view, aperture.

section, burrow infill at K/T boundary, x 14. Coll. GPI Hamburg, no. 3327.

- tube with - view, b - lateral view, c frontal view of Geulhem section, Fig. 9. Holotype, erect part two peristomes, a top aperture.

burrow infill at K/T boundary, x 14. Coll. GPI Hamburg, no. 739.

Canal 14. Coll. GPI 3328. Fig. 10. Erect tube part with three peristomes. Albert sections, as sample 121, x Hamburg, no.

Erect tube from behind. Geulhem above K/T 10.5. Coll. GPI Fig. 11. part, peristome seen section, some metres boundary, x

Hamburg, no. 3329.

above K/T 10.5. Coll. GPI Fig. 12. Erect tube part, strongly developed peristome. Geulhem section, some metres boundary, x

3330. Hamburg, no.

- of third Fig. 13. Spiral detached from substrate, oblique bottom view, showing two perforated tabulae (right centre), a remains a

tabula (lower left). Geulhem section, sample 105, x 21. Coll. GPI Hamburg, no. 3331.

Serpula ? aff. trilineata (Roemer, 1841). Houthem Formation, Geulhem Member.

Fig. 14. Near-complete tube, a - overall view, b - detail showing loops, well-developed ornament of adult tube and juvenile part (right

- Canal sections, x 5.3 x 10.5 Coll. GPI centre) lacking longitudinal ornament, c aperture. Albert as sample 121, (a), (b, c).

Hamburg, no. 3332.

Fig. 15. U-shaped fragment with weak longitudinal ornament and in places strong transverse ornament including peristomes. Albert

Canal sections, sample 121, x 8.7. Coll. GPI Hamburg, no. 3333.

16. with and weak weak Fig. U-shaped fragment strong longitudinalornament transverse ornament including peristomes, a -top/lateral

- Coll. GPI 3334. view, b aperture. Albert Canal sections, sample 121a, x 8.7 (a), x 10.5 (b). Hamburg, no.

- Fig. 17. U-shaped fragment with strong longitudinal ornament and weak transverse ornament, a - top view, b cross-sections. Albert

Canal sections, sample 121a, attached to same substrate as Fig. 16, x 10.5. Coll. GPI Hamburg, no. 3334.

the median - b - cross-section. Albert Fig. 18. U-shaped fragment, the two keels at the periphery being broader than keel, a top view,

Canal sections, sample 121a, x 10.5. Coll. GPI Hamburg, no. 3335.

of details oftube Fig. 19. Fragment oflargest tube seen, with stronglongitudinal ornament and lack transverse ornament, showing some

- b - cross-section. Geulhem 8.7. Coll. GPI 3336. wall, a top view, section, topmost strata, x Hamburg, no. 123\2

PLATE 2 124

Wilckens, O., 1907. Die Lamellibranchiaten, Gastropoden etc. Ocean. —J. fish. Res. Board Canada, 28(10): 1373-1383, 4

der oberen Kreide Siidpatagoniens. — Ber. naturf. Ges. figs.

i. 15: 2-9. 1972. Une actuelle du Freiberg Br., 97-166, pis Zibrowius, H., espece genre Neomicrorbis

Wilckens, O., 1910. Die Anneliden, Bivalven und Gastropoden Rovereto (Polychaeta Serpulidae) decouverte dans l'etage

der antarktischen Kreideformation. — Wiss. Ergebn. bathyal aux Azores. — Bull. Mus. natn Hist. nat. Paris,

schwed. Siidpolarexped., 3(12): 1-132, 4 pis (not seen). (3)39(33, Zool.): 423-430, 1 fig.

The Cretaceous of New 1983. arndti Wilckens, O., 1922. Upper gastropods Zibrowius, H., Chitinopoma n. sp., an incubating

Zealand. — N.Z. Dept Mines, Geol. Surv., Paleont. Bull., 9: bathyal serpulid polychaete from Saint-Paul Island, southern

42 2 1 5 Indian Ocean. — 1 v + pp., figs, map, pis. Tethys, 11(1): 21-24, fig.

Wilckens, O., 1924. Lahillia and some other fossils from the Zibrowius, H., & H.A. ten Hove, 1987. Neovermilia falcigera

Senonian of New Zealand. — Trans. N.Z. Inst., 55: Upper (Roule, 1898), a deep- and cold-water serpulid polychaete

539-544. common in the Mediterranean Plio-Pleistocene. — Bull. biol.

Wrigley, A., 1950. Les opercules de serpulides de l'Eocene du Soc. Wash., 7: 259-271, 6 figs. - Bassin de Paris. Bull. Soc. geol. Fr., (5)19: 499-505, 5 figs. Ziegler, V., 1984. Family Serpulidae (Polychaeta, Sedentaria) — Wrigley, A., 1951. Some Eocene serpulids. Proc. Geol. from the Bohemian Cretaceous basin. — Sbornik Narodniho

66 Assoc., 62(3): 177-202, figs. Muzea v Praze, 39B(4): 213-254, 8 pis.

fossils from Hokkaido: Yabe, H., & T. Nagao, 1928. Cretaceous Ziegler, V., & J. Michalik, 1980. Late Triassic serpulids

Annelida, and Lamellibranchiata. — Sci. Gastropoda Repts (Annelida, Polychaetia, Sedentarida) in the western Car-

Tohoku Imp. Univ., Sendai, 9: 77-96, pis 16, 17. pathians. — Geol. Zbornik, Geol. Carp., 31(4): 627-640, 13

Young, G., & J. Bird, 1828. A geological survey of the Yorkshire figs.

367 coast. Whitby [2nd ed.], pp., 19 pis (not seen).

Zibrowius, H., 1971. Revision of Metavermilia Bush (Poly-

with of three chaeta, Serpulidae), descriptions new species Manuscript received 31 May 1992, revised version accepted 25

from off Portugal, Gulf of Guinea, and Western Indian October 1992.

PLATE 3

Cementula applanata subditiva (Regenhardt, 1961). Houthem Formation, Geulhem Member.

1. initial tube the tube the and then Fig. Atypical specimen; following a near-straight part (left), enwraps substrate, a bryozoan,

ascends above this Geulhem without 8.7. Coll. GPI 3337. (right). section, stratigraphic data, x Hamburg, no.

Fig. 2. Holotype, a - top view, b - top/lateral view, c - detail showing anterior tube part with alae, aperture and keel. Albert Canal

sections, sample 125, x 5.3 (a, b), x 10.5 (c). Coll. GPI Hamburg, no. 725.

Fig. 3. Exceptionally large spiral, a - top view, b - bottom view, detail, showing tube interior. Geulhem section, without stratigraphic

data, x 5.3 (a), x 8.7 (b). Coll. GPI Hamburg, no. 3338.

view. Albert Canal Fig. 4. Slightly oblique spiral, top sections, hardground at km mark 23.850, x 5.3. Coll. GPI Hamburg, no. 3339.

5. with alae and keel last half ’short cut’ and - Fig. Spirals anteriorly, turn taking a by growing across spiral finally becoming erect, a

lateral view, b - top view. Geulhem section, without stratigraphic data, x 7. Coll. GPI Hamburg, no. 3340.

6. Tall broken. Lateral view. Albert Canal sections, Coll. GPI Fig. spirals, tube partially sample 125, x 5.3. Hamburg, no. 3341.

Fig. 7. Tall spirals, tube partially broken, last half turn taking a ’short cut’ by growing across spiral, a - top view, b - lateral view, c -

seen detail of other lateral view, showing tube interior, especially absence of ’internal tube structures’ (ITS) such as in

Albert Canal 5.3 8.7 Coll. GPI 3342. Spiraserpula. sections, sample 121a, x (a, b), x (c). Hamburg, no.

8. Bottom view of tube interior and intermediate of Albert Canal Fig. spiral, showing non-spiral part (left centre). sections, sample 120,

x 10.5. Coll. GPI Hamburg, no. 3343.

9. oferect anterior tube with alae, a - view, b - bottom view, - cross-section. Albert Canal sections, Fig. Fragment part top c as sample

121, x 14. Coll. GPI Hamburg, no. 3344.

Fig. 10. Fragment oferect anterior tube part with alae, a - top view, b - lateral view, c - bottom. Albert Canal sections, as sample 121, x

14. Coll. GPI Hamburg, no. 3345.

Fig. 11. Bottom view of spiral, showing tube interior. Albert Canal sections, sample 121, x 8.7. Coll. GPI Hamburg, no. 3346.

Cluster of broken, tube interior, of ’internal tube Fig. 12. spirals, showing especially absence structures’ (ITS) as seen in Spiraserpula, a, b

- lateral views. Geulhem section, sample 105, x 10.5. Coll. GPI Hamburg, no. 3347.

? Houthem Geulhem Member. Serpula sp. Formation,

13. Broken bottom tube interior. Albert Canal Coll. GPI Fig. spiral, view, showing sections, sample 121, x 5.3. Hamburg, no. 3348.

Fig. 14. Tube with projecting crest, a - top view (aperture right), b - lateral view, c - cross-section of broken posterior end, showing

sections oflongitudinal cavities at base, d - aperture. Albert Canal sections, as sample 138, x 14. Coll. GPI Hamburg, no. 3349.

Fig. 15. Tube fragment, a - lateral view, b - bottom view, showing cellular construction oftube base and interior. Albert Canal sections,

Coll. GPI sample 121, x 8.7. Hamburg, no. 3350. 125\3

PLATE 3 126

PLATE 4

Ditrupa schlotheimi (Rosenkrantz, 1920). Houthem Formation, Geulhem Member.

Fig. 1. Fairly complete tube. Albert Canal sections, Geulhem Member, sample B, x 5.8. Coll. Natuurhistorisch Museum Maastricht,

no. NHMM 1992200-1.

Fig. 2. Anterior tube fragment, with wrinkles. Albert Canal sections, Geulhem Member, sample B, x 5.8. Coll. Natuurhistorisch

Museum Maastricht, no. NHMM 1992200-2.

Coll. Natuurhistorisch Fig. 3. Fairly complete tube. Albert Canal sections, Geulhem Member, sample A, x 5.8. Museum Maastricht,

no. NHMM 1992200-3.

Fig. 4. Fairly complete tube, with bulges. Albert Canal sections, Geulhem Member, sample B, x 5.8. Coll. Natuurhistorisch Museum

Maastricht, no. NHMM 1992200-4.

- lateral view, b - cross-section, Albert Canal sections, Geulhem Member, B, x Fig. 5. Anterior tube fragment, with wrinkles, a sample

9.6 11.5 Coll. Natuurhistorisch Museum NHMM 1992200-5. (a), x (b). Maastricht, no.

- - cross-section, Albert Canal sections, Fig. 6. Anterior tube fragment, swollen club-shaped near aperture, a lateral view, b

Vroenhoven, Geulhem Member, sample A, x 9.6 (a), x 15.3 (b). Coll. Natuurhistorisch Museum Maastricht, no. NHMM

1992200-6.

a - lateral view, b - cross-section, Albert Canal Fig. 7. Anterior tube fragment, with wrinkles, swollen club-shaped near aperture,

of Geulhem Member, x 9.6 x 11.5 Coll. Natuurhistorisch Museum Maastricht, no. NHMM sections, upper part (a), (b).

1992200-7 (leg. J.W.M. Jagt, no. 3043).

Fig. 8. Anterior tube fragment, with wrinkles and bottle-neck shape of apertural region, Albert Canal sections, Geulhem Member,

Coll. Natuurhistorisch Museum NHMM 1992200-8. sample B, x 9.6. Maastricht, no.

9.6. Coll. GPI 3351. Fig. 9. Fragment with irregular curvature, Albert Canal sections, sample 180, x Hamburg, no.

Fig. 10. Fragment of straight tube, with longitudinal furrow and transverse bulges, Albert Canal sections, sample 180, x 5.8. Coll. GPI

Hamburg, no. 3352. of Geulhem section, Geulhem Member above Fig. 11. Tube fragment, with longitudinalfurrow, view convex side. hardground, Voigt

3353. 1960, x 11.5. Coll. GPI Hamburg, no.

Fig. 12. Oblique longitudinal section, showing tube structure. Albert Canal sections, Geulhem Member, sample A, x 15.3. Coll.

Natuurhistorisch Museum Maastricht, no. NHMM 1992200-9.

tube and striation interior. Geulhem of Fig. 13. Longitudinal section, showing structure strong transverse on section, topmost part

Geulhem Member, x 15.3. Coll. GPI Hamburg, no. 3354.

Rotularia (Rotularia) hisingeri (Lundgren, 1891)

Immature anterior tube - b - lateral - bottom Fig. 14. individual, spiral lacking straight part, a top view, view, showing aperture, c view,

showing broken straight initial tube part. Albert Canal sections, sample 127(2), x 7.7 (a-c). Coll. GPI Hamburg, no. 3355.

with anterior tube - b - bottom short initial tube Albert Canal Fig. 15. Spiral long straight part, a top view, view, showing straight part.

sections, sample 127(2), x 5.8. Coll. GPI Hamburg, no. 3356.

- Fig. 16. Spiral with long straight anterior tube part, a top view, b - lateral view, showing short straight initial tube part (lower centre)

and cross-section. Albert Canal 127 7.7. Coll. GPI 3357. sections, sample III, x Hamburg, no.

Fig. 17. Spiral with short straight anterior tube part and fairly longstraight initial tube part (centre to lower right), bottom view, Albert

Canal sections, sample 127(1), x 9.6. Coll. GPI Hamburg, no. 3358.

with anterior tube - b - lateral cross-section ofanterior tube - bottom Fig. 18. Spiral short straight part, a top view, view, showing part, c

view, showing short straight initial tube part, d - cross-section of anterior tube part. Albert Canal sections, sample 127(1), x 9.6

Coll. GPI (a, c), x 7.7 (b), x 15.3 (d). Hamburg, no. 3359.

attached substrate broad - b - lateral - bottom Fig. 19. Extraordinary specimen to by base, a top view, view, showing cross-section, c

view, showing near-straight initial tube part and attachment area with cellular construction of base. Albert Canal sections,

III, Coll. GPI no. 3360. sample 127 x 7.7 (a), x 9.6 (b, c). Hamburg, 127\4

PLATE 4 128

PLATE 5

Rotularia (Rotularia) lobar gracilis n. subsp. Houthem Formation, Geulhem Member.

Fig. 1. Holotype, a - top view, b - detail showing ornament, c - detail showing distorsion at the base of the (broken) straight anterior

tube d - lateral view, detail - - part, showing ornament, e lateral view including cross-section of straight anterior tube part, f

bottom detail - bottom overall view. Albert Canal Geulhem view, showing ornament, g view, sections, Member,sample A, x 7

Coll. (a, e, g), x 10.5 (b-d, f). Natuurhistorisch Museum Maastricht, no. NHMM 1992201-1.

2. - b - detail - lateral view with d - detail view Fig. Spiral, a top view, showing ornament, c slightly damaged aperture, of other lateral

showing ornament, e - bottom view. Albert Canal sections, Geulhem Member, sample A, x 7 (a, c, e), x 8.7 (b), x 10.5 (d). Coll.

Natuurhistorisch Museum Maastricht, no. NHMM 1992201-2.

3. the the but its underside, view, - Fig. Spiral, last ½ turn (damaged) not following spiral periphery, growing across a -top b detail of

lateral view showing ornament, c - bottom view, showing irregular last ½ turn (damaged). Albert Canal sections, Geulhem

7 10.5 Coll. Natuurhistorisch Museum NHMM Member, sample A, x (a, c), x (b). Maastricht, no. 1992201-3.

4. the tube from the broken old - distorsion base of Fig. Spiral, following an injury, new growing part, a top view, injury left, at straight

anterior - tube part, b aperture of new tube at end of straight anterior tube part, c - lateral view, showing injury (left half) and

somewhat of old tube d lateral/bottom view, - detail damagedaperture part (right), -oblique e showing injury (top centre) and

new tube growing from old part (top left), f - bottom view (injury right, new tube top). Albert Canal sections, Geulhem Member,

sample A, x 7 (a, c, d, f), x 14 (b, e). Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992201-4.

5. of anterior tube - b - bottom - cross-section. Canal Fig. Long fragment straight part, a top view, view, c Albert sections, Geulhem

Member, sample A, x 10.5 (a, b), x 14 (c). Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992201-5.

6. Broken detail tube Canal Geulhem Fig. spiral, showing structure, Albert sections, Member, sample A, x 14. Coll. Natuurhistorisch

Museum Maastricht, no. NHMM 1992201-6. 129\5

PLATE 5 130

PLATE 6

Rotularia (Rotularia) tobar vroenhovensis n. subsp. Houthem Formation, Geulhem Member.

tube b - detail transition - view, at the base of the anterior showing Fig. 1. Holotype, a top showing swelling (broken) straight part,

- lateral d - detail cross-section of anterior tube with tube spiral/straight anterior tube part (right), c view, showing straight part,

schlotheimi inside - other lateral f - bottom view. Albert Canal of Geulhem of ? Ditrupa lumen, e view, sections, upper part Natuurhistorisch Museum Maastricht, NHMM 1992202-1 Member,x 7 (a, c, e, f), x 10.5. (b), x 14 (d). Coll. no. (leg. J.W.M.

Jagt, no. 2678).

- view, b - lateral view, c - detail cross-section of Fig. 2. Spiral with relatively stronglongitudinal edges, a top showing (broken) straight

- transition aanterior - bottom view. anterior part, with object inside, d lateral view, showing spiral/straight part (right), e

Albert Canal of Geulhem 7 x 14 x 10.5 Coll. Natuurhistorisch Museum sections, upper part Member, x (a, b, e), (c), (d).

Maastricht, no. NHMM 1992202-2 (leg. J.W.M. Jagt, no. 2678).

3. with anterior tube view. Albert Canal sections, of Geulhem Member, x 5.3. Coll. Fig. Spiral very long straight part, top upper part

NHMM 1992202-3 Natuurhistorisch Museum Maastricht, no. (leg. J.W.M. Jagt, no. 2678).

- - Albert Canal sections, Fig. 4. Slightly irregular spiral with relatively strong longitudinaledges, a top view, b oblique lateral view.

of Geulhem 7. Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992202-4 upper part Member, x (leg. J.W.M. Jagt,

no. 2678).

- lateral b - lateral view. Albert Canal of Geulhem x Fig. 5. Relatively high trochospiral, a oblique view, sections, upper part Member,

NHMM 1992202-5 7. Coll. Natuurhistorisch Museum Maastricht, no. (leg. J.W.M. Jagt, no. 2678).

with anterior tube lateral view. Albert Canal ofGeulhem 7. Fig. 6. Barrel-shaped spiral straight part, sections, upper part Member, x

Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992202-6 (leg. J.W.M. Jagt, no. 2678).

Bipygmaeus pygmaeus (von Hagenow, 1840) ?

Albert Canal 21. Coll. GPI 3361. Fig. 7. Planar spiral, aperture slightly erect. sections, sample 125, x Hamburg, no.

Canal north of km mark Coll. GPI 3362. Fig. 8. Planar spiral, aperture slightly erect. Albert sections, 24,000, x 21. Hamburg, no.

Bipygmaeus pygmaeus (von Hagenow, 1840)

Fig. 9. Planar spiral, followed by conical spiral, a - top view, b - oblique lateral view, c - lateral view.

21. Coll. GPI 3363. Albert Canal sections, sample 125, x Hamburg, no.

indet. Spirorbidae gen. et sp.

Coll. GPI 3364. Fig. 10. Base of broken spiral, top view. Albert Canal sections, north of km mark 24.000, x 21. Hamburg, no.

Pileolaria ? 1 sp.

11. Planar Albert Canal 127 21. Coll. GPI 3365. Fig. spiral, aperture slightly erect. sections, sample II, x Hamburg,

- - - lateral view. Albert Canal sections, Fig. 12. Planar spiral, aperture slightly erect, a top view, b oblique lateral view, c sample 127(1),

Coll. GPI 3366. x 21. Hamburg, no.

Pileolaria ? 2 sp.

- - Coll. GPI Fig. 13. Planar spiral, aperture slightly erect, a top view, b oblique lateral view. Albert Canal sections, sample 180, x 21.

Hamburg, no. 3367.

Pileolaria ? 1 sp.

Fig. 14. Planar spiral, followed by conical spiral, lateral view. Albert Canal sections, north of km mark 24.000, x 21. Coll. GPI

3368. Hamburg, no.

Neodexiospira ? sp. 1

Planar - view, b - lateral view. Albert Canal as 121 same substrate as Pl. Fig. 15. spiral, a top oblique sections, sample (attached to 2,

Fig. 14), x 28. Coll. GPI Hamburg, no. 3332.

Fig. 16. Planar spiral. Albert Canal sections, sample 121, x 28. Coll. GPI Hamburg, no. 3369.

? 2 Neodexiospira sp.

17. Planar with free between the Geulhem 28. Coll. GPI 3370. Fig. spiral space turns. section, sample 105, x Hamburg, no. 131\6

PLATE 6 132

PLATE 7

? Serpula sp.

1. of attached tube with - b - lateral view. Albert Canal Geulhem Fig. Large fragment part strong crest, a top view, oblique sections,

Member, sample B, x 6.3. Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992203.

Metavermilia ? sp.

Fig. 2. Hook-shaped curved tube, bottom view of impression on substrate showing tube’s exterior surface. Albert Canal sections,

sample 121a, x 21. Coll. GPI Hamburg, no. 3371.

tube - bottom view of on substrate tube’s exterior surface, b - Fig. 3. S-shaped fragment, a impression showing impression showing

inversely U-shaped cross-section. Albert Canal section, north of km mark 24.000, x 21. Coll. GPI Hamburg, no. 3372.

Pyrgopolon (Septenaria) sp.

- - view, - cross-section transition attached/erect tube Fig. 4. Tube fragment, a lateral view, b opposite lateral c at parts, showing

x x x structure, d - cross-section oferect tube part. Geulhem, burrow infill directly at K/T boundary, 6.3 (a, b), 10.5 (c), 8.4 (d).

Coll. GPI Hamburg, no. 3373.

Placostegus erectus (Brünnich Nielsen, 1931)

Fig. 5. Attached tube part, forming loop, showing undulate keel and transition attached/erect tube part (lower right). Albert Canal

12.6. Coll. GPI 3374. sections, sample 125, x Hamburg, no.

Fig. 6. Attached tube part, curved, showing denticulate keel. Albert Canal sections, sample 125, x 12.6. Coll. GPI Hamburg, no. 3375.

Fig. 7. Two specimens: attached tube part, showing narrow, high, caterpillar-like socle lifting tube relatively highly above substrate

- lateral - cross-section. Albert Canal 16.8 21 Coll. GPI (bryozoan), a view, c sections, sample 127(2), x (a), x (c). Hamburg,no.

3376. b - lateral view, showing narrow, high, caterpillar-like socle lifting tube relatively highly above substrate, d - opposite

lateral view, showing transition attached/erect part (right), e - top view, showing denticulate keel. Albert Canal sections,

12.6. Coll. GPI 3377. sample 125, x Hamburg, no.

Fig. 8. Attached tube part, including transition attached/erect tube part, twisted S-shaped, showing keel. Albert Canal sections, as

sample 121, x 12.6. Coll. GPI Hamburg, no. 3378.

9. Tube transition attached tube teeth lateral Fig. fragment, showing (right, damaged)/erect part (upper left), forming at aperture,

view. Albert Canal sections, as sample 138, x 12.6. Coll. GPI Hamburg, no. 3379.

Attached tube with and distorted Fig. 10. part, forming narrow loop but weak longitudinal edge, steep slightly erect part (right, broken

anteriorly). Albert Canal sections, sample 127 II, x 12.6. Coll. GPI Hamburg, no. 3380.

Fig. II. Long fragment of erect tube part, showing strong longitudinal ornament, a - top view, b - oblique lateral/bottom view, c -

bottom view, of wall Geulhem section, Coll. showing structure damaged tube (top). sample 105, x 12.6. GPI Hamburg, no.

3381.

of - Fig. 12. Fragment erect tube part, with teeth at aperture (tips of some teeth broken off), a oblique lateral view (left)/bottom view

b - Albert Canal km mark 23.850, 16.8 21 (right), showing longitudinal ornament, aperture. sections, hardground at x (a), x (b).

Coll. GPI Hamburg, no. 3382.

13. of tube with teeth Geulhem section, some Fig. Aperture erect part, (some broken oil). metres above K/T boundary, x 16.8. Coll.

GPI Hamburg, no. 3383.

of - - Fig. 14. Fragment erect tube part, with teeth at aperture and strong transverse ornament, a top view, b lateral view (upper side

- section of echinoid broken after left), c aperture, showing spine resembling operculum. Specimen intentionally photography.

Geulhem section, sample 105, x 16.8 (a, b), x 23.5 (c). Coll. GPI Hamburg, no. 3384. 133\7

PLATE 7 134

PLATE 8

Pyrgopolon (Pyrgopolon) mosae mosae de Montfort, 1808

- view, - Fig. 1. Outer parable layer, complete but short erect tube part, a top showing transverse wrinkles near aperture, b lateral

of base of attached tube - Albert Canal 8.4. Coll. view, showing remnants part (lower left), c aperture. sections, as sample 138, x

GPI Hamburg, no. 3385.

of GPI Fig. 2. Outer parable layer, large fragment erect tube part. Albert Canal sections, as sample 138, x 10.5. Coll. Hamburg, no.

3386.

- Fig. 3. Outer parable layer, small fragment oferect tube part, a exterior, b - detail of interior,showing longitudinal section of tube

wall with lamellae and striation. Albert Canal 10.5 16.8 Coll. GPI parable (top) transverse sections, sample 125, x (a), x (b).

Hamburg, no. 3387.

Fig. 4. Outer parable layer, small fragment oferect tube part. Detail of interior, showing transverse striation. Geulhem section, some

above K/T Coll. GPI metres boundary, x 16.8. Hamburg, no. 3388.

Pyrgopolon (Pyrgopolon) mosae subsp. indet.

of - lateral view, - anterior - cross-section. Fig. 5. Fragment cylinder layer, a b cross-section, c posterior Geulhem section, sample 105,

x 10.5 (a), x 16.8 (b). Coll. GPI Hamburg, no. 3389.

Fig. 6. Fragment ofcylinder layer, lateral view. Geulhem section, some metres above K/T boundary, x 12.6. Coll. GPI Hamburg, no.

3390.

Pyrgopolon (Pyrgopolon) mosae ciplyana (de Ryckholt, 1852)

Fig. 7. Outer parable layer, small fragment, a - lateral view of exterior, showing keels and transverse ribs, b - interior, showing

transverse striation and furrow underneath keel, - cross-section, keels. Albert Canal longitudinal c showing sections, as sample

138, x 12.6. Coll. GPI Hamburg, no. 3391.

Fig. 8. Outer parable layer, small fragment, showing longitudinal edges. Albert Canal sections, as sample 138, x 12.6. Coll. GPI

Hamburg, no. 3392.

Fig. 9. Outer parable layer, small fragment, showing longitudinal keels. Albert Canal sections, sample 180, x 16.8. Coll. GPI

Hamburg, no. 3393.

Placostegus alatus (Brünnich Nielsen, 1931)

of curved attached tube view Fig. 10. Fragment part, oblique lateral (direction ofgrowth from right to lower left), showing keel and

peristomes. Albert Canal sections, north of km mark 24.000, x 10.5. Coll. GPI Hamburg, no. 3394.

Fig. 11. Fragment of transition attached/erect tube parts, cross-section of attached part, showing keel and peristomes. Albert Canal

sections, 120, x 16.8. Coll. GPI sample Hamburg, no. 3395.

12. oftransition attached/erect tube - view, weak - lateral Fig. Fragment part, a top showing ornament, b oblique view, showing socle

at anterior end ofbase and median furrow at underside oferect part. Albert Canal sections, as sample 121, x 10.5. Coll. GPI

Hamburg, no. 3396.

13. of transition attached/erect tube - lateral b - bottom Fig. Fragment parts, a view, showing socle, view, showing socle and median

furrow at underside oferect part (right), c - cross-section of erect tube part. Albert Canal sections, sample 125, x 12.6 (a, b), x

Coll. GPI 16.8 (c). Hamburg, no. 3397.

14. of erect tube median furrow and underside and - bottom b -cross-section. Fig. Fragment part, showing bulges at peristome, a view,

Albert Canal sections, sample 125, x 12.6 (a), x 16.8 (b). Coll. GPI Hamburg, no. 3398.

15. of tube - b - lateral - bottom median d - Fig. Fragment erect part, showing peristome, a top view, view, c view, showing furrow,

cross-section. Albert Canal sections, sample 121, x 10.5 (a-c), x 16.8 (d). Coll. GPI Hamburg, no. 3399.

of tube cross-section. Albert Canal sections, Coll. GPI no. 3400. Fig. 16. Fragment erect part, sample 121, x 16.8. Hamburg,

Fig. 17. Fragment of curved erect tube part, top view, showing peristomes. Albert Canal sections, sample 121, x 10.5. Coll. GPI

Hamburg, no. 3401.

18. of curved tube and with - view b - bottom view Fig. Fragment erect part, showing ornament aperture teeth, a top (aperture left),

(aperture right). Albert Canal sections, sample 179, x 12.6. Coll. GPI Hamburg, no. 3402. 135\8

PLATE 8 136

PLATE 9

Neomicrorbis parietalis jagti n. subsp.

- lateral intermediate tube tube and torsion of Fig. 1. Holotype, a view, showing spiral (left), part (bottom centre), erect part (right) this,

b - another lateral view, showing spiral and erect part with aperture, c - oblique top view, d - other lateral view, showing erect

- surrounded keels. Albert Canal sections, above K/T b, d, part, e aperture, by protruding 0-0.4 m boundary, x 12.6 (a, e), x 8.4

(c). Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992204-1 (leg. J.W.M. Jagt, no. 872).

Fig. 2. Very small spiral with intermediate and erect tube part, showing socle and aperture. Albert Canal section, sample 125, x 16.8.

Coll. GPI 3403. Hamburg, no.

3. Erect tube torsion. Albert Canal 12.6. Coll. GPI 3404. Fig. part (broken), showing sections, sample 120, x Hamburg, no.

4. intermediate tube and short - keels and b - Fig. Paratype. Spiral, part very erect part, a top view, showing peristomes, oblique top

in - lateral view socle and Albert Canal 0-0.4 above K/T view, showing callosity umbilicus, c showing aperture. sections, m

boundary, x 12.6 (a, b), x 8.4 (c). Coll. Natuurhistorisch Museum Maastricht, no. NHMM 1992204-2 (leg. J.W.M. Jagt, no.

872).

5. intermediate tube and - lateral view b - Fig. Paratype. Spiral, part erect part, a oblique showing peristomes, top view, showing keels,

aperture and callosity in umbilicus. Albert Canal sections, 0-0.4 m above K/T boundary, x 12.6. Coll. Natuurhistorisch

Museum NHMM 1992204-3 Maastricht, no. (leg. J.W.M. Jagt, no. 872).

6. intermediate tube and - and b - Fig. Paratype. Spiral, part erect part, a oblique top view, showing peristomes slightly irregular keels,

lateral view, showing aperture. Albert Canal sections, 0-0.4 m above K/T boundary, x 10.5. Coll. Natuurhistorisch Museum

Maastricht, no. NHMM 1992204-4 (leg. J.W.M. Jagt, no. 872).

Fig. 7. Operculum, isolated from typical tube, a-c - lateral views, showing cucullus and calcar, d - top view of cucullus. Albert Canal

GPI sections, sample 125, x 33.6. Coll. Hamburg, no. 3405.

Neomicrorbis cingulatus (Brünnich Nielsen, 1931)

Fig. 8. Large tube with strong transverse ornament, a - oblique top view, showing corkscrew-shaped erect tube part, b - top view,

showing aperture and small second specimen (lower right) of same species, c, d - lateral views oferect part, showing ornament.

Geulhem section, burrow infill at K/T boundary, x 8.4 (a, b), x 12.6 (c, d). Coll. GPI Hamburg, no. 3406.

9. Broken interior - lateral view b - broken bottom Fig. tube, showing structure, a (broken spiral bottom, erect part upper right), spiral

view. Albert Canal Coll. GPI sections, sample 121, x 12.6. Hamburg, no. 3407. 137\9

PLATE 9 138

established in 1967 Thus he attempted to reconstruct environmental ities of a working group, princi-

environ- unravel and conditions for fossil biota, using data on pally to Paratethys stratigraphy pal- whole of mental requirements of modern species inhabiting aeogeography, but soon extended to the

and of Asia and Africa. This the marine shelf as a starting point. Europe, parts project

correlations between the basins He collected comparative material himself, through suggested Neogene

Black the of central and southern North diving sessions in the Baltic Sea, the Sea, western, Europe, the shores of Adriatic and the Caribbean. He also organised Africa and the Near East, as far east as

Cuban—Czecho-Slovak expeditions in the Carib- the Indian Ocean. It unified the work of hundreds

for intimate collabora- of and and the result of bean and was responsible the geologists palaeontologists, of the tion with the Institute for Marine Research in this elfort is the synthesis entitled 'Neogene

and in which Rovinj during 1965-1975. Mediterranean Tethys Paratethys',

successions of the basins Apart from his scientific work for the Academy he the Neogene stratigraphic

his to of this extensive described and illustrated. attempted to pass on knowledge younger area are

awards from institutions and universities in generations by lecturing at the Comenius University Many

Czecho-Slovakia and abroad were conferred on Jan (Bratislava), and at the J.E. Purkyne, now Masaryk

he lector in Senes in of his scientific contributions. University (Brno), where was appointed recognition

1967. In the he took his doctor's The medal of Stur, the decoration of same year degree gold Dionyz

meritfor the furtherance of the natural sciences and at Bratislava, on a paper on Neogene palaeogeogra-

in data the medal awarded to him. He phy of the western Carpathians; fact, pre- ofJ.E. Purkyne were

valid for the whole At member ofthe Czecho-Slovak National Com- sented are of central Europe. was a

mittee for and member the this time, he reintroduced and defined the term Geology, an honorary of

marine basins and Austrian Societies. Paratethys for the, in part, isolated of Hungarian Geological Senes nine and western, central and eastern Europe. Jan published monographs more

these than 180 in domestic and Of his many papers published during years, papers foreign journals, the und Neo- and co-authored editions of various one entitled 'Chronostratigraphie geological stratotypen', which he edited and co-authored, was maps.

The of the He was a not in scientific circles, of special importance. regional stages personality, only

Miocene (Badenian, Sarmatian, Pannonian, Pon- but also in everyday life: in spite of his scientific

had the central Euro- he ceased be and this tian) not only a bearing on burdens, never to human, trait made him but were also as far as character so popular and appreci- pean stratigraphy, important

in all of the world ated. He there for advice in word correlations in the Neogene parts was always help or

deed. He knew how the beauties of and the establishment of global chronostratigraphie or to appreciate

and and active he stages were concerned. nature art, as an mountaineer,

the of this diver he His organisational talent did not stop at publishing experienced heights earth, as a

of From 1967 the of the as an amateur on subjects chronostratigraphy. explored depths sea, pilot

member Strati- he ventured the he had onwards, he was a of the National skies, as a philatelist a

of and graphic Committee of the Czecho-Slovak Academy proper sense detail, as an organiser a sense

he the for It is in this that of of Sciences, and in the same year joined synthesis. way most us remem-

order managing committee of the Regional Committeeon ber him; we should follow his example, in to

and Mediterranean Neogene Stratigraphy (RCMNS), pass on his message to younger generations

and in the advancement of which was appointed president at the Lyon congress actively support geology,

1971. The fact that he held this position for two he lived for.

in the framework of the terms, a rare phenomenon

InternationalUnion of Geological Sciences (IUGS), Dr Michal Kovac

in this In he elected Institute is relevant respect. 1979, was Geological

World Slovakian of Sciences president of the Subcommission on Neogene Academy

Stratigraphy (SNS). Dubravska Cesta 9

well his Czecho-Slovakia The above-mentioned talent, as as person- 84226 Bratislava,

the fore of the ality, came to during the execution

extensive correlation project, IGCP-UNESCO no.

which based the 25 during 1973-1983, was on activ-