ION NOliniliSNI NVINOSHillNS S3IHVyail LIBRARIES SMITHSONIAN INSTITUTION NOlinili

_ >

in - -z to *^2

11 LIBRARIES SMITHSONIAN INSTITUTION NOlifUliSNI NVINOSHillNS SIIBVaail LIBRAR

— in ^ in — in ~

in ^ w

Q ''i2L£S-^ — Q ~-^iiisgx Q xO\Tbc^

ION ^ NOliniliSNI ““NVlNOSHimS S3iaVHan~'LIBRAR I ES^SMITHSONIAN"" INSTITUTION ^NOlinilJ

73

m ^ m

(/) — CO _ to

BRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHIIWS S3iyVHan LIBRAR

in z in z

i s i

> ' 5 X^sviiS!' > ■ 5

in 2

NOliniliSNI NVINOSHIIWS S3IBVyan libraries SMITHSONIAN INSTITUTION NOIinili

an libraries smithsonian institution NoiiniiiSNi nvinoshii/js S3iavaan librar

■|0N NOliniliSNI NviNOSHiiws S3iavaan libraries smithsonian institution Noiinii.

CO
2

X
in
O

> 5 ^ 2

2 CO 2 c/3

n libraries smithsonian institution NOliniliSNI NviNOSHims S3iavaa

~ in ^ ^ in 5

4 . I lj I ^-<'T7w7!*Vw 1 1 1 >^USV/l4

C/>

1_ LI B RAR
z

in

ION ** N0iiniiiSNi'"NviN0SHiiws^s3 I a va a n libraries smithsonian institution Noiinii.

z r- z r- 2 <2 ^ ^ 2: ^ ^

O x;STrr(7;>\ _ vv o ~ O >5^. m xToolJoX ^

i^y CO ^ z m 'in * " ■' m ^

an~LIBRARI ES^SMITHS0NIAN~INSTITUTI0N^ NOliniliSNI “nvinoshiiims S3iavaail_l-IBRAR

osHiii^s S3iyvyan libraries Smithsonian institution^ NoiiniiisNi^NviNOSHims^sa i ^

i %

HSONIAN^ INSTlTUTION‘^NOIinillSNl^NVINOSHilWS^S3 IBVaan^LIBRARI ES^SMITHSONIAN _ INST

OSHillNS^S3iavaan"'uBRARIES^SMITHSONIAN“;iNSTITUTION^NOIiniliSNi;^NVlNOSHillNS^S3

o O O ~ °

HSONIAN INSTITUTION NOIifliliSNI NVINOSHimS S 3 I B V B 3 11

BRARIES SMITHSONIAN INSl

CO ^ ^ z

O ^

c^' 5 ^

A- ^

osHims S3 lava a

m
UJ

q:

w

B RAR I Es'^SMITHSONIAN INSTITUTION NOlifUliSNI _ NVINOSHil S 3

CO ~ CO — ^

03

‘HSONIAN~'lNSTlTUTION NOIlfUliSNI NVINOSHimS S 3 I a V a 3 11^ L I B R A R I E S^ SMITHSONIAN^ INS

osHims'^sa i avaa n u brar i es'^smithsonian iNSTiTUTiON,„NoiiniiiSNi NviNosHims^sa

. (C\ - C/7 ^ --

HSONIAN^ INSTlTUTION^NOlifUliSNI NVINOSHillMs'^S 3 I ava 3 I1_LI BRAR I ES SMITHSONIAN INS

rr\ — (r\

Q _ XiOiiiSJiZ o " — o ^

IOSHimS^S3 I ava a n^LI B RAR l ES^SMITHS0NIAN^INSTITUTI0N^N0lini!lSNI^NVIN0SHimS^S3

TO O’ ^ °

za
>

faj 33
rn

— CO

rucr\MiAM IMCTITIITinM MnfinillQM

MWIM/^CliMIAIC' 0-1 \ \JW\J

'i inoAPlF^; 5^MITH^nNlAN

.'f" ■ '

j*te. ■

a*** r::_ *

i

|5\.« ^

I'^V;. 'I'. . ■

' ■wm

Wf ■'■ ■

'.tC,..

f

' :' ff''

> A

■li

-.v „■

/<■

J ^ •*%,,, , V

V' • .,'•' ■ ,

.' "''i’

;; 'a '<

‘ft

;'T<f ’

-k;.

«•!

.:. 'V'S

A.

-

'. '■ ‘ ivrsi'j.

■r-

1^

,- m'S'

'WJ'.

.. .. ”. ,r<* • .;

■ 'V- jO/;"' V

^■' 'i-i; .„>4r -■

■ ■ m

‘■K.. !;

w .v-*

' s ^.■'

«, . ■? JntV'v

., Kfii'V i jji;;/-,

fc'.'Ai'fi’S

VOLUME 6

Palaeontology

1963

PUBLISHED BY THE
PALAEONTOLOGICAL ASSOCIATION
LONDON

Dates of publication of parts in Volume 6

Part 1, pp. 1-218, pis. 1-26
Part 2, pp. 219-396, pis. 27-56
Part 3, pp. 397-596, pis. 57-80
Part 4, pp. 597-770, pis. 81-108

26 April 1963

27 May 1963

7 October 1963
10 December 1963

THIS VOLUME EDITED BY N. F. HUGHES, W. S. McKERROW, GWYN THOMAS, AND ISLES STRACHAN

PRINTED IN GREAT BRITAIN

CONTENTS

Part Page

Amos, A., and Boucot, A. J. A revision of the brachiopod family Leptocoeliidae 3 440

Arkell, W. J., and Callomon, J. H. Lower Kimeridgian ammonites from the drift of
Lincolnshire 2 219

Balme, B. E. Plant inicrofossils from the Lower Triassic of Western Australia 1 12

Barnard, T. The morphology and development of species of Marssonella and Pseiido-

textulariella from the Chalk of England 1 41

Boucot, A. J. See Amos, A.

Breimer, a. See Joysey, K. A.

Bulman, O. M. B. On Glyptograptiis dentatiis (Brongniart) and some allied species 4 665

Callomon, J. H. See Arkell, W. J.

Chaphekar, N. Some calamitean plants from the Lower Carboniferous of Scotland 3 408

Chuplac, Ivo. Phyllocarid crustaceans from the Silurian and Devonian of Czecho-
slovakia 1 97

Clemens, W. A. Late Jurassic mammalian fossils in the Sedgwick Museum, Cambridge 2 373

— Wealden mammalian fossils 1 55

CooGAN, A. H. See Wilson, E. C.

CoRDEY, W. G. The genera Brotzenia and Voorthiiysenia (Foraminifera) and Hofker’s
classification of the Epistomariidae 4 653

Cox, E. R. The Rhaetic-Hettangian bivalve genus Pteroniya Moore 3 582

Craig, G. Y., and Hallam, A. Size-frequency and growth-ring analyses of Mytiliis edidis
and Cardium edide, and their palaeoecological significance 4 731

Currie, Ethel D., and George, T. Neville. Catalogue of described and figured speci-
mens in the Begg Collection in the Hunterian Museum of the University of Glasgow 2 378

Delcourt, a. F., Dettmann, M. E., and Hughes, N. F. Revision of some Eower
Cretaceous microspores from Belgium 2 282

Dettmann, M. E. See Delcourt, A. F.

Downie, C. ‘ Hystrichospheres ’ (acritarchs) and spores of the Wenlock Shales (Silurian)
of Wenlock, England 4 625

— and Sarjeant, W. A. S. On the interpretation and status of some hystrichosphere

genera 1 83

Doyle, J. C. A new heteromorph ammonite from the Lower Cretaceous of Yorkshire 3 575

Elliott, G. F. A Palaeocene teredinid (Mollusca) from Iraq 2 315

— Problematical microfossils from the Cretaceous and Palaeocene of the Middle East 2 293

George, T. Neville. See Currie, Ethel D.

Hallam, A. Observations on the palaeoecology and ammonite sequence of the Fro-
dingham Ironstone (Lower Jurassic) 3 554

— See also Craig, G. Y.

Hendry, R. D., Rowell, A. J., and Stanley, J. W. A rapid parallel grinding machine
for serial sectioning of fossils 1 145

House, M. R., and Pedder, A. E. H. Devonian goniatites and stratigraphical correlations
in Western Canada 3 491

Hudson, J. D. The ecology and stratigraphical distribution of the invertebrate fauna of
the Great Estuarine Series 2 327

— The recognition of salinity-controlled mollusc assemblages in the Great Estuarine

Series (Middle Jurassic) of the Inner Hebrides 2 318

Hughes, N. F. See Delcourt, A. F.

Jackson, D. E., and Lenz, A. C. A new species of Mouograptits from the Road River
Formation, Yukon 4 751

Jeletzky, j. a. Malayomaorica gen. nov. (Family Aviculopectinidae) from the Indo-
Pacific Upper Jurassic; with comments on related forms 1 148

Joysey, K. A., and Breimer, A. The anatomical structure and systematic position of
Pc/ifflWfliPis (Blastoidea) from the Carboniferous of Spain 3 471

CONTENTS

Part Page

Kaye, P. The ostracod genus Neocythere in the Speeton Clay 2 274

— The ostracod species Orthonotacythere inversa (Cornuel) and its allies in the Speeton

Clay of Yorkshire 3 430

Lagaaij, R. Cupiiladria canariensis (Busk) — portrait of a bryozoan 1 172

Lenz, a. C. See Jackson, D. E.

Logan, A. The new brachiopod genus Howseia from the Permian Magnesian Limestone
of Durham 4 754

Love, L. G. The composition of Pyritosphaera barbaria Love 1957 1 119

Miller, T. G. The bryozoan genus Polypora M‘Coy 1 166

Moore, L. R. Microbiological colonization and attack on some Carboniferous miospores 2 349

Pant, D. D., and Verma, B. K. On the structure of leaves of Rhabdotaeiiia Pant from the
Raniganj coalfield, India 2 301

Pedder, a. E. H. Alaiophyllum mackenziense sp. nov., a Devonian tetracoral from Canada 1 132

— See also House, M. R.

Philip, G. M. Two Australian Tertiary neolampids, and the classification of cassiduloid
echinoids 4 718

Reyment, R. a., and Sandberg, P. Biometric study of Barremites siibdifficilis (Karakasch) 4 727

Ross, J. R. P. Lower Permian Bryozoa from Western Australia 1 70

— Trepostome Bryozoa from the Caradoc Series, Shropshire 1 1

Rowell, A. J. See Hendry, R. D.

Samanta, B. K. Two new species of Discocyclina (Loraminifera) from the Upper Eocene
of Assam, India 4 658

Sandberg, P. See Reyment, R. A.

Sarjeant, W. a. S. See Downie, C.

Skwarko, S. W. a new Upper Cretaceous Ophiuroid from Australia 3 579

Smith, J. D. D., and White, D. E. Cambrian trilobites from the Purley Shales of
Warwickshire 3 397

Spjeldn.es, N. Some silicified Ordovician fossils from South Wales 2 254

— Some Upper Tremadocian graptolites from Norway 1 121

Stanley, J. W. See Hendry, R. D.

Stephenson, D. G. The spines and diffuse fascioles of the Cretaceous echinoid Echino-
corys scutata Leske 3 458

Stephenson, N. G. Growth gradients among fossil monotremes and marsupials 4 j615

Stevens, G. R. The systematic status of Oppel’s specimens of Belemnites gerardi 4 690

Verma, B. K. See Pant, D. D.

Waines, R. H. See Wilson, E. C.

Waterhouse, J. B. New Zealand species of the Permian bivalve Atomodesma Beyrich 4 699

Watson, D. M. S. On growth stages in branchiosaurs 3 540

White, D. E. See Smith, J. D. D.

Wills, L. J. Cyprilepas holmi Wills 1962, a pedunculate cirripede from the Upper Silurian

of Oesel, Esthonia 1 161

Wilson, E. C., Waines, R. H., and Coogan, A. H. A new species of Komia Korde and the
systematic position of the genus 2 246

Wilson, R. B. The lamellibranch genus Prothyris in the Upper Devonian and Carboni-
ferous of Great Britain 1 136

Wood, A. The British Carboniferous species of Girvanella (Calcareous Algae) 2 264

Wright, C. W. Cretaceous ammonites from Bathurst Island, Northern Australia 4 597

VOLUME 6 • PART 1

Palaeontology

APRIL 1963

PUBLISHED BY THE
PALAEONTOLOGICAL ASSOCIATION
LONDON

Price £3

THE PALAEONTOLOGICAL ASSOCIATION

The Association was foimded in 1957 to further the study of palaeontology. It holds
meetings and demonstrations, and publishes the quarterly journal Palaeontology.
Membership is open to individuals, institutions, libraries, &c., on payment of the
appropriate annual subscription:

Institutional membership . £5. 5s. (U.S. $15.50)

Ordinary membership ... . £3. 3^. (U.S. $9.50)

Student membership . . . . £2. 2s. (U.S. $6.50)

There is no admission fee. Student members will be regarded as persons receiving
full-time instruction at educational institutions recognized by the Council. Subscrip-
tions are due each January, and should be sent to the Treasurer, Professor P. C.
Sylvester-Bradley, Department of Geology, The University, Leicester, England.

Palaeontology is devoted to the publication of papers (preferably illustrated) on all
aspects of palaeontology and stratigraphical palaeontology. Four parts are pubhshed
each year and are sent free to all members of the Association. Members who join for
1963 will receive Volume 6, Parts 1 to 4.

All back numbers are stiU in print and may be ordered from B. H. Blackwell,
Broad Street, Oxford, England, at the prices shown below (post free) :

Vol. 1 (for 1957-8) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 2 (for 1959) in 2 parts at £2 or U.S. $6.00 per part.

Vol. 3 (for 1960) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 4 (for 1961) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 5 (for 1962) in 4 parts at £3 or U.S. $9.00 per part.

A complete set. Volumes 1-5, consists of 18 parts and costs £40 or U.S. $120.

Manuscripts on all aspects of palaeontology and stratigraphical palaeontology are
invited. They should conform in style to those already published in this journal, and
should be sent to Mr. N. F. Hughes, Sedgwick Museum, Cambridge, England.
A sheet of detailed instructions for authors will be supplied on request.

TREPOSTOME BRYOZOA FROM THE
CARADOC SERIES, SHROPSHIRE

by JUNE R. P. PHILLIPS ROSS

Abstract. Mesotrypa bitlmani sp. nov., Amplexopora thomasi sp. nov., Homotrypa sp. A, and Homotrypa sp. B
occur in the Hoar Edge Limestone in the lower part of the Caradoc Series, near Evenwood and Hamage. These
trepostome species show similarities with species from Cincinnatian strata in Ohio, Indiana, and Anticosti
Island.

The four trepostome species described in this paper were collected from the lower part
of the Caradoc Series in the Evenwood and Harnage area in southern Shropshire (text-
fig. 1). Mesotrypa buhnani sp. nov., Amplexopora thomasi sp. nov., and Homotrypa sp.
A occur in the Harknessella siibquadrata horizon in the section at Evenwood Quarry
(text -fig. 2). Mesotrypa buhnani is also found in two quarries in the Eloar Edge Lime-
stone south-west of Harnage Grange, and M. buhnani sp. nov., A. thomasi sp. nov.,
and Homotrypa sp. B occur in this formation at the Wilderness, Wilstone Farm House.
These species are from fossil collections made during the mapping of the Shrewsbury
District and locality data and identifications made by Dr. C. J. Stubblefield and Dr. G.
Elies are listed in Pocock et al. (1938). The cryptostome Phaenopora stubblefieldi Ross
(1962) from the Hoar Edge Limestone also came from these collections and was located
in the upper part of the 15-foot section in Evenwood Quarry in the Harknessella
subquadrata horizon which overlies lower units containing graptolites of the Nema-
graptus gracilis Zone. The succeeding sandstone and shale beds above the Hoar Edge
Limestone contain graptolites regarded as characterizing the Diplograptus multidens
Zone.

TEXT-FIG. I. Index map of Evenwood-Harnage area, Shropshire.

Few bryozoan species have been described from the Caradoc Series and the occurrence
of species of Amplexopora, Homotrypa, and Mesotrypa which have been recorded from

[Palaeontology, Vol. 6, Part 1, 1963, pp. 1-11, pi. 1-3.]

C 1015

B

2

PALAEONTOLOGY, VOLUME 6

Ordovician strata in North America, Greenland, England, Estonia, Russia, and
Australia, adds to the knowledge of the distribution of these three genera. Mesotrypa
bulmani sp. nov. is similar to M. orbiculata from the Cincinnatian Arnheim Formation
of south-eastern Indiana, M. patella from the Whitewater Formation of south-western
Ohio (text-fig. 3), and M. infida from the Rhinidictya beds in Minnesota. It shows little
similarity to M. lens (M‘Coy) (Spjeldnass 1957) from the Caradocian Horderly Sand-
stone of eastern Shropshire.

Amplexopora thomasi sp. nov. resembles A. pustulosa from the Cincinnatian Waynes-
ville Shale of south-western Ohio, A. ampla from the Cincinnatian Leipers Limestone of
central Tennessee and the Cincinnatian Fairmount Member of south-western Ohio, and
A. billingsi from the Cincinnatian English Head and Vaureal Formations of Anticosti
Island.

Homotrypa sp. A is similar to a rare species, H. diimosa, from the Cincinnatian Fair-
mount Member of south-western Ohio, and Homotrypa sp. B has many morphologic
features that resemble H. obliqua from the Cincinnatian Fairmount and Corryville
Members of south-western Ohio, and H. wortheni from the Cincinnatian Whitewater
Formation of south-western Ohio and south-eastern Indiana.

Collection numbers with the prefixes WM and RR, and specimen numbers with the prefix GSM
refer to material in the Geological Survey Museum, London.

SYSTEMATIC DESCRIPTIONS

Family amplexoporidae Miller 1889
Genus amplexopora Ulrich

1882 Amplexopora Ulrich, p. 154.

1960 Amplexopora Ulrich; Boardman, pp. 16-20.

1920 Acaiithotrypella Vinassa de Regny, p. 221.

Tvpe species. Amplexopora cingulata Ulrich (1882, pp. 126, 254-6, pi. 11, figs. 5, 5a-c); designated by
Ulrich (1882, p. 255).

Diagnosis. Colonies are ramose, encrusting, massive, or rarely bilaminate. Monticules
are commonly present. Zooecial openings are polygonal and are enclosed by generally
integrate zooecial walls. Mesopores are generally not abundant. Acanthopores are
commonly abundant and occur at the junctions of zooecial walls. In axial regions the
zooecial tubes have slender walls. In the peripheral regions the thickened zooecial walls
have a laminate microstructure in which the outer parts of adjacent zooecial walls have
steeply inclined laminae that intersect at acute angles. Diaphragms are present in the
peripheral and subperipheral regions and may be flat, curved, or cystoidal. Some species
display cystiphragms.

Amplexopora thomasi sp. nov.

Plate 1, figs. 1-10

Material. Collections WM 1265 (including paratype GSM 102009), WM 1267, WM 1273 (including
holotype GSM 102004, paratypes GSM 102005, GSM 102006), WM 1274 (including paratype GSM
102007), from the top part of Evenwood Quarry; and collection WM 1279, from the middle part of
Evenwood Quarry; Hoar Edge Limestone, Hoar Edge Group; Caradoc Series; Ordovician.

Description. In tangential sections the slender, integrate zooecial walls enclose polygonal

J. R. P. PHILLIPS ROSS: CARADOC TREPOSTOME BRYOZOA 3

or subpolygonal zooecial openings which vary considerably in diameter (PI. 1, figs. 1, 7,
10). One to two mesopores are generally located at the junctions of the zooecial walls
(PI. 1, figs. 1, 10). Acanthopores are rare and, if present, are small, dense spots in the
zooecial walls.

Transverse sections of ramose colonies display axial regions with thin-walled poly-
gonal zooecial tubes and peripheral regions with thickened, integrate zooecial walls and

UPPER CAMBRIAN

TEXT-FIG. 2. Measured section of the Hoar Edge Group in Evenwood Quarry (after Pocock et cil. 1938).

diaphragms crossing the zooecial tubes. Transverse sections of bilaminate colonies dis-
play a dark medial line lacking microstructure and with zooecia on both sides of it.
The zooecial tubes have thin walls in the medial region and thickened, integrate walls
and diaphragms in the peripheral region (PI. 1, figs. 2, 4).

In longitudinal sections of ramose colonies the axial regions display very broadly
crenulate zooecial walls that are very slender. For some distance the walls diverge only
slightly from the axis of growth of the branch and then bend abruptly into the peripheral
region. In this region they thicken and are more closely crenulate and very slightly
beaded. The zooecial openings are slightly oblique at the periphery. The zooecial and
mesospore walls have an integrate, laminate microstructure as noted in the Diagnosis of
Amplexopora (PI. 1, fig. 9). The very slender, widely spaced diaphragms crossing the
zooecial tubes pass into the steeply inclined wall laminae (PI. 1, fig. 9). In the mesopores,
which develop at the base of the peripheral region, the diaphragms are closely spaced.

4

PALAEONTOLOGY, VOLUME 6

In longitudinal sections of bilaminate colonies the zooecial tubes curve gently to the
periphery and are crossed by widely spaced diaphragms. The zooecial openings are
oblique to the periphery (PI. 1, fig. 8).

TABLE 1

Measurements of Amplexopora thomasi sp. nov. (in millimetres)

Catalogue no.

GSM 102007

GSM 102004
holotype

Diameter of zoarial stem ....

6-7

6-7

Diameter of zooecial opening

. min.

014x016

0-15X0-15

No. of zooecia per 2 mm. ....

max.

0-31 X 0-22
6-8

0-23 X 0-21
6-8

Combined thickness of adjacent zooecial walls
region .......

in peripheral

004-006

004-006

Diameter of mesopore ....

. min.

003x003

0-05

Diameter of acanthopore ....

max.

OlOxO-12

0-01-0-04

010

001-002

No. of acanthopores per zooecium

0-6

0-5

No. of diaphragms per 1 mm. in zooecium in peripheral region

4-6

3-5

No. of diaphragms per 1 mm. in mesopores.

14-20

11-15

Width of peripheral region

1-2

2-0

Ratio: width of peripheral region of zooecium/total width of
zooecium. .........

0-4

0-6

Remarks. The species is characterized by slender zoarial stems with wide peripheral
regions in which the thickened, crenulate, and slightly beaded zooecial and mesopore
walls are distinctly integrate and in which mesopores have closely spaced diaphragms
and the zooecial tubes have widely spaced diaphragms. Acanthopores, if present, are not
prominent.

Amplexopora thomasi is similar to A. billingsi (Bassler 1928) from the Ordovician
(Cincinnatian) English Head and Vaureal Formations, Anticosti Island. A. billingsi has

EXPLANATION OF PLATE 1

Figs. 1-10. Amplexopora thomasi sp. nov. 1, Tangential section showing integrate walls between zoo-
ecial tubes and mesopores, collection WM 1273, holotype GSM 102004, X 20. 2, Transverse section
of a bilaminate colony showing narrow peripheral region of thickened walls, collection WM 1273,
GSM 102005, X 20. 3, Part of longitudinal section in peripheral region showing thick, laminate
zooecial walls and diaphragms in zooecial tubes and mesopores, collection WM 1274, GSM 102007,
X 50. 4, Part of transverse section of bilaminate colony showing integrate, laminate zooecial walls
in peripheral region, collection WM 1273, GSM 102005, X 50. 5, Part of a longitudinal sectioninthe
peripheral part of a ramose colony showing the finely crenulate zooecial walls, collection WM 1273,
GSM 102004, X 20. 6, Part of a longitudinal section of a ramose colony showing very slender walls
in the axial region and thickened walls in the peripheral region, collection WM 1274, GSM 102007,
X 20. 7, Tangential section showing general arrangement of zooecial openings, collection WM 1274,
GSM 102007, X 20. 8, Longitudinal section of a bilamrnate colony showing diaphragms across zoo-
ecial tubes, collection WM 1265, GSM 102009, X 50. 9, Part of a longitudinal section in peripheral
part of a colony showing integrate, laminate zooecial walls, and thin diaphragms, collection WM
1274, GSM 102007, X 100. 10, Tangential section showing distinct boundaries between adjacent
zooecial walls, collection WM 1273, GSM 102004, x50.

Palaeontology, Vol. 6

PLATE 1

PHILLIPS ROSS, Ordovician Bryozoa

J. R. P. PHILLIPS ROSS: CARADOC TREPOSTOME BRYOZOA

5

fewer zooecial openings per 2 mm., fewer mesopores, and more numerous diaphragms
in the zooecial tubes. Both species have inconspicuous acanthopores, closely spaced
diaphragms in the mesopores, and crenulate zooecial walls. A. thomasi, A. pustidosa
Ulrich (1890) from the Ordovician (Cincinnatian) Waynesville Shale, Hanover, Ohio,
and A. ampla Bassler (1904) from the Ordovician (Cincinnatian) Leipers Limestone,
Nashville, Tennessee, are similar in the diameter of their zoarial stems and have about
the same number of zooecia per 2 mm. A. pustiilosa and A. ampla differ from A. thomasi
in having diaphragms in the axial region, and A. pustidosa commonly has acanthopores
at the junctions of the zooecial walls and has few mesopores in the peripheral region.

SYSTEM

<

O

>

o

Q

cr

0

1

SERIES

<

I-

o

o

_s_

GROUP

FORMATION

MEMBER

RICHMOND

elkhorn

WHITEWATER

LIBERTY

WAYNESVILLE

ARNH EIM

MAYS-

VILLE

McMillan

CORRYVILLE

FAIRVIEW

FAIRMOUNT

EDEN

\j-Mesotrypa patella
\^Homolrypa wortheni

Amptexopora

pustutosa

-M. orbiculata

■H. obtiqua

H, dumosa
H. obHqua
■A, ampla

TEXT-FIG. 3. General geologic column of formations and members in the Cincinnatian Series in south-
western Ohio and south-eastern Indiana from which Bassler (1915) reported the bryozoan species
that are here compared with the Caradocian species.

Amplexopora thomasi appears to have some similarity to A. murchisoni (SpjeldntES
1957), the type of which is from ‘Horderley’, Shropshire, but its exact locality and strati-
graphic position is not known. Unfortunately the type material of A. murchisoni is
partly recrystallized so that recognition of the species is difficult. However, Spjeldnaes
(1957, text-figs. \a, b) illustrates the typical wall microstructure of species of Amplexopora
and his text-fig. \b appears to show small mesopores and also acanthopores at the junc-
tions of the zooecial walls.

The species is named in honour of Dr. H. Dighton Thomas, British Museum (Natural
History).

Family monticuliporidae Nicholson
Genus mesotrypa Ulrich 1893

Type species. Mesotrypa infida (Ulrich 1886, p. 88).

Diagnosis. Colonies are discoidal, hemispherical, conical, or encrusting expanses. Lower
surfaces of discoidal colonies are flat or concave. Polygonal or cylindrical zooecial tubes
grow directly upwards from the basal lamina and are crossed by flat and curved dia-
phragms. Mesopores are present in both the proximal and distal parts of a colony and
have closely spaced diaphragms. Zooecial openings are polygonal in some colonies
and circular in others. Mesopore openings are rectangular or polygonal and may be very

6

PALAEONTOLOGY, VOLUME 6

small. Acanthopores are prominent where the zooecial openings are angular and meso-
pores are small, and they are small and indistinct in colonies where zooecial openings are
round and mesopores are large. The zooecial and mesopore walls display a granular
microstructure at low magnifications.

Mesotrypa bulmani sp. nov.

Plate 2, figs. 1-9; Plate 3, figs. 2, 4

Material. Collections WM 1262 (including paratype GSM 102003) and WM 1274 (including paratype
GSM 102008), from the top part of Evenwood Quarry. Collection WM 1279 (including paratypes
GSM 102011-102015), from the middle part of Evenwood Quarry. Collections RR 2635 (including
paratype GSM 102016), RR 2636 (including paratype GSM 102019), RR 2637, and RR 2638, from
the old quarry in the wood near Coundmoor Brook. Collections RR 2669 and RR 2672 (including
holotype GSM 102017), Wylde’s Quarry, near Harnage Grange. All collections from Hoar Edge
Limestone, Hoar Edge Group; Caradoc Series; Qrdovician.

Description. Many of the discoidal colonies, 10 to 13 mm. diameter at their bases, and
1 to 2 mm. high, have flat or concave bases. The encrusting colonies vary from small,
cylindrical forms, 1-5 to 2 mm. in diameter, to broad, explanate forms that extend across
fragmented shells and other material.

In tangential sections the zooecia are variable in size and shape. In a well-developed
discoidal colony, GSM 102017, almost round zooecial openings and polygonal or
rectangular mesopores fill the spaces between clusters of large, polygonal zooecial
openings and a few mesopores. In these clusters where the mesopores are very few,
acanthopores, 0-02 to 0-03 mm. in diameter, are present at the junctions of the zooecial
walls (PI. 2, fig. 7) ; where the mesopores are distinctly visible between the zooecia, the
acanthopores are generally lacking (PI. 3, fig. 4). In an encrusting form, GSM 102003,
and a discoidal form, GSM 102015, the polygonal zooecial openings are separated by
a few mesopores, and prominent acanthopores, 0-03 to 0-06 mm. in diameter, lie at the
junctions of the zooecial walls (PI. 2, fig. 8). Deeper tangential sections of colonies
display more slender walls enclosing polygonal tubes (PI. 2, fig. 5). Walls between
zooecia and mesopores are slender and amalgamate.

In longitudinal sections the zooecial tubes may be slightly reclined for a short distance
along the basal lamina (PI. 2, fig. 4). Acanthopores and mesopores are commonly
present a short distance above the basal lamina and extend to the distal parts of colonies.

EXPLANATION OF PLATE 2

Figs. 1-9. Mesotrypa bulmani sp. nov. 1, Part of longitudinal section of subperipheral and peripheral
parts of colony showing diaphragms in the zooecial tubes and mesopores, collection WM 1279,
GSM 102011, x50. 2, General aspect of transverse section, collection WM 1279, GSM 102012, x20.
3, Part of a longitudinal section of a multilaminate colony showing thin, granular walls in the peripheral
region, collection RR 2635, GSM 102016, X 50. 4, Longitudinal section of multilaminate colony
showing slender zooecial walls and diaphragms in the zooecial tubes and mesopores, collection WM
1279, GSM 102013, X 20. 5, Deep tangential section showing distinct acanthopores at the junctions
of very slender zooecial walls, collection WM 1274, GSM 102008, X 50. 6, Tangential section show-
ing polygonal zooecial openings, numerous mesopores, and very fine acanthopores, collection RR
2636, GSM 102019, X 50. 7, 8, Tangential sections showing distinct acanthopores at junctions of
zooecial walls; collection RR 2672, holotype GSM 102017, X 50; and collection WM 1262, GSM
102003, x50, respectively. 9, Qblique longitudinal section showing diaphragms in the mesopores
and in the long zooecial tubes, collection WM 1279, GSM 102011, X20.

Palaeontology, Vol. 6

PLATE 2

PHILLIPS ROSS, Ordovician Bryo/.oa

J. R. P. PHILLIPS ROSS: CARADOC TREPOSTOME BRYOZOA

7

As the walls, mesopores, zooecial tubes, and acanthopores have some variation in their
direction of growth, a longitudinal section may pass across and then back into a meso-
pore, acanthopore, or even a zooecial tube. The zooecial and mesopore walls thicken
slightly as they extend to the periphery (PI. 2, figs. 4, 9). The very slender zooecial walls
have a very fine, laminate microstructure that commonly appears granular at low
magnifications. The clear inner parts of the zooecial walls lining the zooecial tubes
consist of fine laminae lying at 45 degrees to the inner boundary. The outer parts of
adjacent zooecial walls appear as dark bands of intertonguing convexly curved, granular

TABLE 2

Measurements of Mesotrypa bulmani sp. nov. (in millimetres)

Catalogue no.

GSM 102017

No. of zooecia per 2 mm. .......

8-10

Diameter of zooecial opening ......

. max.

0-22 X 0-22

min.

016x016

Diameter of zooecial opening in monticule ....

. max.

0-29 X 0-29

min.

0-29 X 0-26

Combined thickness of adjacent zooecial walls in peripheral region

002

Diameter of mesopore ........

. max.

010x015

min.

006x003

Diameter of acanthopore .......

001-003

No. of acanthopores per zooecium .....

2-4

in monticules

No. of diaphragms per 1 mm. in zooecium ....

6-8

No. of diaphragms per 1 mm. in mesopore ....

about 25

laminae. The thin diaphragms crossing the zooecial tubes and mesopores pass into the
wall laminae after bending abruptly at the zooecial wall boundary. Acanthopores have
slender, dark walls enclosing clear axial spaces. In encrusting forms the zooecia extend
upright to the periphery and in discoidal forms the zooecial tubes curve broadly near
the periphery (PI. 2, figs. 4, 9). Diaphragms are present throughout the length of the
zooecial tubes and are commonly flat, or convexly or concavely curved, and in the peri-
pheral region may be more closely spaced and include additional kinds such as compound
and cystoidal diaphragms (PI. 2, fig. 4). The mesopores have closely spaced diaphragms
throughout their length (PI. 2, figs. 3, 4, 9).

Remarks. The species is characterized by small, commonly discoidal colonies with round
or subpolygonal zooecial openings, sparsely distributed mesopores, abundant dia-
phragms throughout the length of the mesopores, widely spaced diaphragms in the
zooecial tubes, and small acanthopores which are most prominent in the monticules.

Mesotrypa bulmani is similar to M. infida (Ulrich 1886, 1893) from ‘the middle third
of the Trenton Shales at Minneapolis’ ( = the Rhinidictya and Ctenodonta beds of Ulrich
1893, p. /) in the form of the zoarium and the shape of the zooecial openings. M. bul-
mani has smaller zooecial openings both in and between the monticules, has fewer
acanthopores per zooecium, and lacks funnel-shaped diaphragms. M. bulmani resembles
M. orbieulata Cumings and Galloway (1913) from the Ordovician (Cincinnatian)
Arnheim Formation near Harmon’s Station, Indiana, in the size and form of the colonies.

PALAEONTOLOGY, VOLUME 6

and in the arrangement of the diaphragms in the zooecial tubes and mesopores.
M. bidmani has a greater number of zooecia per 2 mm., fewer and smaller mesopores
between the zooecia, and generally smaller acanthopores. M. bidmani M. patella
(Ulrich 1890) from the Ordovician (Cincinnatian) Whitewater Formation, Oxford, Ohio,
in having about the same number of zooecia per 2 mm., small acanthopores of about
0-02 mm. diameter, and in the arrangement of diaphragms in the mesopores and zooecial
tubes. M. bidmani has fewer mesopores between the zooecial openings in smaller
colonies.

The species is named in honour of Professor O. M. B. Bulman, Sedgwick Museum,
Cambridge.

Genus homotrypa Ulrich 1882

Type species (by original designation). Homotrypa curvata Ulrich (1882, p. 242, pi. 10, figs. 7, la-d).

Diagnosis. Colonies are ramose and encrusting. Monticules may be present. In tangential
sections, wide integrate zooecial walls enclose polygonal or subpolygonal zooecial
openings. Acanthopores with dense, concentrically laminate walls commonly occur at
the junctions of zooecial walls. Mesopores are sparse and may be present only in clusters.
In longitudinal sections zooecial walls are thin in the axial region and thicken in the
peripheral region where they display a laminate microstructure. The thick laminae of the
diaphragms and cystiphragms pass into the wall laminae and form part of the zooecial
walls. These wall laminae are steeply inclined to the boundary of the zooecial tube and
laminae of adjacent walls abut in an irregular dark band or line. Cystiphragms and
diaphragms are abundant in the peripheral region.

Homotrypa sp. A

Plate 3, figs. 1, 3, 5, 6

Material. Single fragment of a colony in collection WM 1265, from the top of Evenwood Quarry;
Hoar Edge Limestone, Hoar Edge Group ; Caradoc Series ; Ordovician. Eigured specimen GSM 1 020 1 0.

Description. The specimen is part of a ramose colony. In tangential sections polygonal
mesopores are distributed around the polygonal and subpolygonal zooecial openings

EXPLANATION OF PLATE 3

Figs. 1, 3, 5, 6. Homotrypa sp. A. Collection WM 1265, GSM 102010. 1, Tangential section showing
general arrangement of zooecial openings and clusters of large zooecia, X 20. 3, Tangential section
showing subpolygonal zooecial openings, mesopores, and crescentic lines of cystiphragms curving
across zooecia, x 50. 5, Part of an oblique transverse section in the peripheral region showing curved
laminate microstructure of zooecial walls, X 50. 6, Part of a longitudinal section in the subperipheral
and peripheral regions showing cystiphragms lining the zooecial tubes and diaphragms crossing
the zooecial tubes, X 50.

Figs. 2, 4. Mesotrypa bulmani sp. nov. 2, Transverse section of a small colony showing the slender
zooecial walls and mesopores with closely spaced diaphragms, collection WM 1279, GSM 102014,
X 20. 4, Tangential section showing general arrangement of zooecial openings, collection RR 2672,
GSM 102017, x20.

Figs. 7-9. Homotrypa sp. B. Collection RR 1481, GSM 102018. 7, Part of an oblique longitudinal
section in the peripheral region showing cystiphragms and diaphragms in the zooecial tubes, X 50.
8, General aspect of longitudinal section in peripheral region showing closely spaced diaphragms and
cystiphragms in zooecial tubes, x 20. 9, Tangential section showing distinct acanthopores at the
junctions of zooecial walls, x 50.

Palaeontology, Vol. 6

PLATE 3

PHILLIPS ROSS, Ordovician Bryozoa

J. R. P. PHILLIPS ROSS: CARADOC TREPOSTOME BRYOZOA

9

(PI. 3, figs. 1, 3). Dense acanthopores are occasionally observed at the junctions of the
zooecial walls and they may indent the inner parts of the zooecial walls into the zooecial
tubes. Curved cystiphragms cross the zooecial openings (PI. 3, fig. 3). The zooecial walls
are generally integrate but may also appear amalgamate.

In longitudinal sections the zooecial tubes curve in a broad arc and are oblique to the
periphery (PI. 3, fig. 6). In the peripheral region the zooecial walls thicken and display
a laminate microstructure. The inner parts of the zooecial and mesopore walls consist of
steeply inclined laminae and the laminae of adjacent zooecial walls intertongue and
abut in a steeply pitched arc to form a dark, outer band (PI. 3, fig. 5). Cystiphragms
commonly located only on the distal wall in the peripheral region are widely spaced so
that adjacent cystiphragms are not always in contact (PI. 3, fig. 6). Widely spaced, flat
diaphragms cross the zooecial tubes in the peripheral region. The mesopores lack dia-
phragms and acanthopores were not readily observed in longitudinal sections.

TABLE 3

Measurements of Homotrypa sp. A (in millimetres)

Catalogue no.

GSM 102010

Diameter of zoarial stem .........

3-5

No. of zooecia per 2 mm. .........

9-10

Diameter of zooecial opening ....... max.

018x016

min.

013x013

Combined thickness of adjacent zooecial walls in peripheral region

002-004

Diameter of mesopore ......... max.

008x006

min.

002x003

Diameter of acanthopore .........

001

No. of acanthopores per zooecium .......

0-4

No. of diaphragms per 1 mm. in zooecium in peripheral region

0-2

Width of peripheral region .........

0-77

Ratio: Width of peripheral region of zooecium/total width of zooecium

0-41

Remarks. The species is characterized by slender, ramose stems in which the peripheral
region contains very slender zooecial walls and very few diaphragms and few cysti-
phragms, the latter generally located on the distal zooecial walls. Acanthopores are
sparse, and mesopores are numerous.

Homotrypa sp. A, which is represented by a single specimen in the collections studied,
is similar to a rare species, H. dumosa Bassler (1903), from the Ordovician (Cincinnatian)
Fairmount Member of the Fairview Formation at Covington, Kentucky, and Cincin-
nati, Ohio. The two species are similar in having slender zooecial walls in the peripheral
region and in having few acanthopores, few diaphragms, and few cystiphragms in the
peripheral region. Insufficient material does not permit adequate definition of this species
from the Hoar Edge Limestone.

Homotrypa sp. B
Plate 3, figs. 7-9

Material. A single fragment of a colony in collection RR 1481, The Wilderness, 18 chains north-west
of Wilstone Farm House; Hoar Edge Limestone, Hoar Edge Group; Caradoc Series; Ordovician.
Figured specimen GSM 102018.

10

PALAEONTOLOGY, VOLUME 6

Description. The specimen is part of a ramose colony with overgrowths. In tangential
sections the subpolygonal zooecial openings are crossed by cystiphragms (PL 3, fig. 9).
Prominent acanthopores with dense, laminate walls and a clear axial region lie at the
junctions of the zooecial walls and may project into the zooecial tubes. An occasional
acanthopore penetrates the sides of the zooecial walls. Mesopores are rare and, if
present, occur as small polygonal structures between the zooecia. The zooecial walls are
generally integrate.

In longitudinal sections, which display mainly the peripheral region and only the
crushed, delicate walls of the axial region, the zooecial tubes are filled with flat and
cystoidal diaphragms and overlapping cystiphragms (PI. 3, figs. 7, 8). The inner parts of
the zooecial walls consist of steeply inclined laminae and laminae of adjacent zooecial
walls intertongue in a distally convex arc which extends distally as a dark, rolled band.
The thick laminae forming the diaphragms and the cystiphragms pass into the wall
laminae. Acanthopores appear as long, clear tubes cutting through the zooecial walls.

TABLE 4

Measurements of Homotrypa sp. B (in millimetres)

Catalogue no.

GSM 102018

Diameter of zoarial stem .......

6-9

No. of zooecia per 2 mm. .......

8-9

Diameter of zooecial opening ......

. max.

0-23 X 0-22

min.

010x010

Combined thickness of adjacent zooecial walls in peripheral region

002-006

Diameter of mesopore ........

. max.

006 X 0-02

min.

003x002

Diameter of acanthopore .......

0-03-005

No. of acanthopores per zooecium ......

0-2

No. of diaphragms per 1 mm. in zooecium in peripheral region.

10-11

Remarks. This species is characterized by a peripheral region which has thick zooecial
walls, few but distinct acanthopores, few mesopores, and numerous cystiphragms and
diaphragms.

Homotrypa sp. B and H. obliqua Ulrich (1882) from the Ordovician (Cincinnatian)
Fairmount and Corryville Members at Cincinnati, Ohio, are similar in having about the
same number of zooecia per 2 mm., few mesopores, and a similar arrangement of dia-
phragms and cystiphragms. Homotrypa sp. B has larger, more prominent acanthopores.
Homotrypa sp. B and H. wortheni (James 1882; Bassler 1903) from the Whitewater
Formation at Oxford, Ohio, and Richmond, Indiana, are similar in having about the
same number of zooecia per 2 mm., few mesospores, and thick zooecial walls in the
peripheral region. Homotrypa sp. B has fewer and more prominent acanthopores.

Insufficient material does not permit adequate definition of this species from the Hoar
Edge Limestone.

Acknowledgements. I express my sincere thanks to Dr. C. J. Stubblefield, Director of the Geological
Survey of Great Britain, for the loan of the material and for assistance in examining the bryozoan
specimens while in London; and to Dr. F. W. Anderson and Mr. J. D. D. Smith, also of the Survey,
for generous assistance while examining the collections. I gratefully acknowledge financial support for
this study from the National Science Foundation, Washington, D.C., U.S.A.

J. R. P. PHILLIPS ROSS: CARADOC TREPOSTOME BRYOZOA

11

COLLECTION LOCALITIES
HOAR EDGE LIMESTONE

WM 1262, WM 1265, WM 1267, WM 1273, and WM 1274, from the upper part of Evenwood Quarry
at the east end of Black Dick’s coppice, 300 yards south of Evenwood, Shropshire: WM 1262 has
Phaenopora stubblefieldi and Mesotrypa bulmani', WM 1274 has Amplexopora thomasi, M. bulmani,
and P. stubblefieldi', WM 1267 has A. thomasi and P. stubblefieldi', WM 1273 has M. bulmani,

A. thomasi, and P. stubblefieldi', WM 1265 has A. thomasi, Homotrypa sp. A, P. stubblefieldi, and
Mesotrypa sp.

WM 1 279, from the middle part of Evenwood Quarry at the east end of Black Dick’s coppice, 300 yards
south of Evenwood, Shropshire: Amplexopora thomasi and Mesotrypa bulmani.

RR 1481, The Wilderness, 18 chains north-west of Wilstone Farm House, Shropshire; Homotrypa sp.

B, Amplexopora! sp.

RR 2635, RR 2636, RR 2637, and RR 2638, old quarry in the wood near Coundmoor Brook, over
I mile south-west of Harnage Grange, Shropshire: Mesotrypa bulmani. Locality as indicated by
Pocock et al. (1938, fig. 28).

RR 2669 and RR 2672, Wylde’s Quarry, 30 chains south-west of Harnage Grange, Shropshire: Meso-
trypa bulmani. Locality as indicated by Pocock et al. (1938, fig. 28).

REFERENCES

BASSLER, R. s. 1903. The structural features of the bryozoan genus Homotrypa, with descriptions of
species from the Cincinnatian Group. Proc. U.S. nat. Mus. 26, 565-91, pi. 20-25.

1915. Bibliographic index of American Ordovician and Silurian fossils. Bull. U.S. nat. Mus. 92,

1521 pp.

■ 1928. Bryozoa. /// twenhofel, w. h.. Geology of Anticosti Island. Mem. geol. Surv. Can. 154,

143-68, pi. 5-14.

BOARDMAN, R. s. 1960. A revision of the Ordovician bryozoan genera Batostoma, Anaphragma, and
Amplexopora. Smithson, misc. Coll. 140 (5), 1-28, 7 pi.

COMINGS, E. R. and GALLOWAY, J. J. 1913. Stratigraphy and paleontology of the Tanner’s Creek section
of the Cincinnati Series of Indiana. Rep. Ind. Dep. Geol. 37, 35-478, 20 pi.

JAMES, u. P. 1882. Descriptions of Cincinnatian and other Paleozoic fossils. The Paleontologist (6), 50.
(Not seen.)

1883. Descriptions of Cincinnatian and other Paleozoic fossils. Ibid. (7), pi. 1, fig. 2. (Not seen.)

POCOCK, R. w., WHITEHEAD, T. H., WEDD, c. B. and ROBERTSON, T. 1938. Shrewsbury District including
the Hanwood Coalfield. Mem. geol. Surv. U.K., 297 pp.

ROSS, J. R. p. p. 1962. Early species of: he bryozoan genus Phaenopora from the Caradoc Series, Shrop-
shire. Palaeontology, 5, 52-58, pi. 9.

SPJELDN/LS, N. 1957. A redescription of some type specimens of British Ordovician Bryozoa. Geol.
Mag. 94, 364-76, pi. 12, 13.

ULRICH, E. o. 1882. American Paleozoic Bryozoa. Cincinnati Soc. nat. Hist. 5, 121-75, 232-57, pi.

10, 11.

■ 1886. Report on the lower Silurian Bryozoa, with preliminary description of the new species.

Minnesota geol. and nat. Hist. Surv., 14th Ann. Rept., 57-103.

1890. Paleozoic Bryozoa. Illinois geol. Surv., Geology and Paleontology, 8, 283-688, pi. 29-78.

1893. On Lower Silurian Bryozoa of Minnesota. Minnesota geol. and nat. Hist. Surv., Geology

of Minnesota, 3 (1), Paleontology, 96-332, 28 pi. 1895. (Author’s separate, 1893.)

and BASSLER, R. s. 1904. A revision of the Paleozoic Bryozoa. Pt. 2, Trepostomata. Smithson.

misc. Coll. 47, 15-55, pi. 6-14.

viNASSA DE REGNY, p. 1921. Sulla classificazione dei treptostomidi. Atti Soc. ital. Sci. nat. 59 (for
1920), 212-31.

JUNE R. P. PHILLIPS ROSS

Illinois State Geological Survey,
Urbana, Illinois, U.S. A.

Manuscript received 22 February 1962

PLANT MICROFOSSILS FROM THE
LOWER TRIASSIC OF WESTERN AUSTRALIA

by B. E. BALME

Abstract. Seventeen species of plant microfossils, made up of nine spores, six pollen grains, and two bodies of
uncertain function, are described from the Kockatea Shale, a marine formation of early Triassic age, occurring
in the Perth Basin, Western Australia. Nine new species are proposed and a new form genus Liindbladispora is
instituted to include certain trilete spores of probable lycopodiaceous affinities. Species belonging to the genus
Taeniaesporites (Leschik) are described for the first time from the southern hemisphere.

Microfloras from the Kockatea Shale differ completely from any known to occur in Australian Permian
sediments and are considered to represent a specialized plant community which succeeded the Glossopteris-¥\ora
at the end of Permian time. They resemble in some respects Upper Permian and Triassic microfloras from Europe,
and Scythian assemblages from the Peace River area in Canada. The palaeofloristic and stratigraphical implica-
tions of the palynological data are discussed.

Permian and older rocks in the northern part of the Perth Basin, Western Australia,
are, in some places, overlain by the Kockatea Shale, a marine deposit of considerable
stratigraphic importance. At two localities it contains ceratitic ammonoids and other
marine invertebrate fossils of Lower Triassic (Scythian) age. It is, therefore, the only
Triassic sequence in Australia which can so far be dated unequivocally in terms of the
Austrian marine standard. The Kockatea Shale consists predominantly of greenish,
grey, and red shale with occasional siltstone and sandstone units. It has been known for
many years in bores near Geraldton, but its Triassic age was unsuspected until 1957,
when Dr. P. E. Playford discovered ammonoids in a core from 1,470 feet in the Gerald-
ton Racecourse Bore. One of these has been identified as an ophiceratid of upperm.ost
Permian or basal Triassic age (Glenister and Furnish 1961). A rich and well-preserved
assemblage of spores, pollen grains, and microplankton was recovered from the same
core and on the basis of this, a correlation was proposed between the Kockatea Shale
and the Blina Shale in the Canning Basin (McWhae, Playford, Lindner, Glenister, and
Balme 1957). Refinement of the earlier age determinations became possible in 1959
following the drilling, by the Commonwealth Bureau of Mineral Resources, of a strati-
graphic test bore at Beagle Ridge, about 80 miles south of Geraldton. This bore (BMR
10) penetrated over 1,000 feet of Kockatea Shale, and cores from the unit yielded marine
faunas at several horizons. Palaeontologists of the Bureau of Mineral Resources, Can-
berra, are still studying this material, but identifications of some of the key genera have
recently been published by Dickins (in Dickins, McTavish, and Balme 1961). They include
the ophiceratid Subinyoites and the pelecypods Claraia and cf. Bakevillia, and demon-
strate the Scythian age of the Kockatea Shale.

Proved exposures of the Kockatea Shale are known only from a small area in the upper
reaches of the Greenough River (text-fig. 1) although it may correlate with part of the
Chapman Group of Arkell and Playford (1954). It has been recognized in bores at
Geraldton, Beagle Ridge, and in the vicinity of Indarra, a small township on the
Mullewa road about 45 miles east of Geraldton. In coastal bores the Kockatea Shale
has a thickness of up to 1,000 feet, but it thins rapidly to the east of Geraldton and is less

[Palaeontology, Vol. 6, Part 1, 1963, pp. 12-40, pi. 4-6.]

B. E. BALME: TRFASSIC PLANT MICROEOSSILS

13

than 300 feet thick in bores close to the Urella Fault. Erosion in the Triassic and Lower
Jurassic may partly explain this easterly attenuation but it seems that the Scythian
shoreline lay to the west of the present line of the Darling Fault. Spores and pollen
grains are most abundant and best preserved in sediments from the Eradu-Indarra
district and it is unlikely that these were deposited far from the Lower Triassic coastline.

TEXT-FIG. 1. Generalized geological map of part of the Perth Basin showing localities of the samples

examined.

Nevertheless, all samples so far examined from the Kockatea Shale contain hystricho-
sphaerids, usually in enormous numbers, and there can be little doubt that the unit is
marine, even in its easterly occurrences.

Poor exposures and a scarcity of reliable sub-surface data obscure the stratigraphical
relationships of the Kockatea Shale. It is clearly transgressive on the Permian, for it
overlies Artinskian marine strata in Beagle Ridge Bore and continental Upper Permian
sediments at Indarra. Below Geraldton it rests directly on Archaean metasediments.

Triassic beds of probable continental origin overlie the Kockatea Shale in Beagle
Ridge Bore; elsewhere it is succeeded by marine or continental Jurassic.

Microfloras from the Kockatea Shale are intrinsically interesting, but their excellent
preservation and the precision with which they can be dated, give them an especial

14

PALAEONTOLOGY, VOLUME 6

stratigraphical and phytogeographical importance. They throw light on the fate of the
GJossopteris-¥\ox2L in Western Australia and can be fairly closely compared with
assemblages occurring in Lower Triassic sediments in other parts of the world. The
emerging palynological evidence hints, therefore, that plant microfossils may provide
a means of placing an upper limit on the Permian System in Australia.

Source and storage of samples. Seventeen samples from the Kockatea Shale provided the material
for the present account. They were obtained from five boreholes and a shallow water-well in the nor-
thern part of the Perth Basin and their general localities are shown in text-fig. 1 . Further details of the
samples and their origin are as follows :

Well at Point 217 (28° 34' 30" S., 115° 7' 10" E.)

Sample 44070. Pale greenish-grey claystone. Depth 30 ft.

471 Mile Peg Bore (28° 39' 55" S., 115° 15' 20" E.)

Sample 43813. Greenish-grey shale. Depth 741-749 ft.

Sample 43815. Pale grey claystone. Depth 808-945 ft.

Sample 43816. Pale greenish-grey shale. Depth 948-968 ft.

Sample 43819. Brownish-grey sandy shale. Depth uncertain.

Sample 43820. Brownish-grey sandy shale. Depth uncertain.

Kockatea Creek No. 19 Bore (28° 33' 40" S., 115° 13' 30" E.)

Sample 43305. Pale greenish-grey claystone. Depth 139-190 ft.

Geraldton Racecourse Bore (28° 47' 36" S., 114° 39' 16" E.)

Sample 44342. Pale grey claystone. Depth 1,273 ft.

Sample 44497. Greenish-grey, red-stained, claystone. Depth 1,465 ft.

Geraldton Municipal Bore (28° 46' 42" S., 114° 35' 50" E.)

Sample 47563. Pale greenish-grey sandy shale. Depth 1,012-1,096 ft.

Sample 47564. Yellowish sandy shale. Depth 1,177-1344 ft.

Sample 47565. Pale greenish-grey calcareous shale. Depth 1,346-1,386 ft.

BMR. 10 (Beagle Ridge) Bore (29° 49' 38" S., 114° 58' 30" E.)

Sample 43938. Pale grey sandy siltstone. Depth 2,131-2,141 ft.

Sample 43939. Grey micaceous shale. Depth 2,223-2,233 ft.

Sample 43940. Grey micaceous shale with ammonoids. Depth 2,405-2,415 ft.

Sample 43941. Grey sandy shale. Depth 2,802-2,812 ft.

Sample 43942. Grey calcareous shale. Depth 3,203-3,213 ft.

Sample numbers in the above list, and throughout the text, refer to the general collection of the Depart-
ment of Geology, University of Western Australia. Small reference samples of each of the sediments
treated are lodged in this repository as are the slides of the type specimens and spore assemblages. The
original samples, which provided the source of reference material, are held either by the Commonwealth
Bureau of Mineral Resources, Canberra, or by the Geological Survey of Western Australia. Type
specimens are mounted singly in glycerine jelly and sealed with bees-wax.

Techniques. Acid insoluble microfossils were extracted by boiling the crushed, carbonate-
free sediment in 50 per cent, hydrofluoric acid and treating the residue by the modified
Schulze method described by Balme and Hassell (1962). The residues were lightly
stained with Safranin ‘O’ and examined by means of a Leitz ‘Ortholux’ microscope
using a PI Apo Oel 100/T32 objective for detailed study of morphography. Photo-
micrographs were taken on Ilford Pan F film using the same oil immersion objective.
They were developed in Kodak D76 and printed on Ilford ‘Plastika’ paper.

Acknowledgements. The author is indebted to the Director of the Commonwealth Bureau of Mineral
Resources, Canberra (Mr. J. M. Rayner), for supplying material from the Beagle Ridge Bore and for
permitting publication of the data obtained. The former Government Geologist of Western Australia
(Mr. H. A. Ellis) also generously permitted the author access to the collections of the Geological
Survey. Thanks are due to the Management of West Australian Petroleum Proprietary Limited, who

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

15

virtually initiated the investigation and at all times were co-operative and helpful. The author is par-
ticularly grateful for the factual information and stimulating discussion provided by two geologists of
that Company, Dr. P. E. Playford and Mr. S. P. Willmott. Dr. J. Jansonius, Calgary, Dr. Wilhelm
Klaus, Vienna, and Mr. N. F. Hughes, Cambridge, have been patient and helpful correspondents, and
the photomicrographs are the work of Mr. K. Bauer.

SYSTEMATIC PALYNOLOGY

With the exception of a short paper by Hennelly (1958) and the recent contribution on
megaspores by Dettmann (1961), no formal papers have been published on Australian
Triassic palynology. Preliminary descriptions of microfloras from the Leigh Creek and
Ipswich coals have been presented by Taylor (1953) and de Jersey (1949) respectively,
but neither of these authors attempted to apply a binomial nomenclature to their plant
microfossils. In the northern hemisphere, however, palynologists have been increasingly
occupied with Upper Permian and Triassic sediments during the past five or six years,
and papers from Russia, western Europe, and North America have proved surprisingly
relevant to the present account. Malyavkina (1953) was the first to describe a varied
assemblage of Triassic spores. These came from the Keuper of the Russian Pre-Urals, but
Malyavkina used no photographs and adopted a rather informal taxonomy. Her account
is, therefore, less useful as a systematic reference than the subsequent papers of Potonie
and Klaus (1954), Leschik (1955, 1956), Klaus (1960), and Jansonius (1962).

Leschik’s papers are well illustrated and provide a basis for the classification and
comparison of isolated spores and pollen grains of Triassic age. Nevertheless, they
contain certain nomenclatural inconsistencies, and some of the new genera and species
proposed were poorly defined or based on misinterpretation of morphographic features.
Critical revision of Leschik’s systematics has been undertaken by Potonie (1958),
Klaus (1960), and Jansonius (1962), but no stable taxonomy has yet evolved. This applies
particularly to the classification of disaccate pollen grains assignable to the Infraturma
Striatiti of Pant. Further discussion of this and other taxonomic problems will be found
in subsequent sections of the present paper.

Taxonomic criteria are based on the publication of Potonie and Kremp (1954), and
subsequent papers by Potonie. Certain of the suprageneric form taxa proposed by
Potonie and Kremp and later authors have also been adopted. However, no attempt has
been made to apply the all-embracing scheme of higher form categories favoured by
many recent workers. This system has certain advantages in bringing apparent order to
the bewildering variety of trilete spores in Upper Devonian and Carboniferous micro-
floras. Nevertheless, it seems unwise, at this stage of our knowledge of the morphology
and affinities of fossil spores, to obscure possible natural relationships by over-rigid
codification.

Terminology. With few exceptions morphographic terms have been taken from the
glossaries provided by Potonie and Kremp (1955) and Couper (1958). Amb was defined
by Erdtman as the outline of a spore or pollen grain viewed from the direction of the
polar axis, and the term cavate is used in the sense of Dettmann (1961), to describe a
spore in which the exoexine and intexine have become detached.

Measurements, unless otherwise stated, are taken on specimens preserved in full
polar view, and the dimensional nomenclature is based on that of Couper (1958, text-
fig. 7, p. 101).

16

PALAEONTOLOGY, VOLUME 6
Anteturma sporites H. Potonie
Turma triletes Reinsch

Genus punctatisporites (Ibrahim) Potonie and Kremp 1954
Type species. Punctatisporites pimctatus Ibrahim, Upper Carboniferous, West Germany.

Punctatisporites fimgosus sp. nov.

Plate 4, figs. 10, 11

Holotype. Slide 47544. Paratype. Slide 47545.

Diagnosis. Amb circular, periphery smooth, off-polar compressions common and exine
frequently ruptured. Trilete, scar distinct, laesurae straight and often of unequal length
but seldom extending more than about half-way to the equatorial margin. Groove of
commissure visible in some specimens. Exine very thick, with fine irregularly distributed
pits visible under oil immersion. Narrow anastomosing pits and channels sometimes
developed, particularly in the area of the proximal pole (PI. 4, fig. 11). These channels
are probably due to partial destruction of the exine, either during fossilization or as a
result of the maceration process.

Dimensions. Diameter 83-119 p; 15 specimens.

Locus typicus. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio nominis. Latin fungosus = ‘spongy’, from the texture of the exine.

Description. Holotype diameter 114 ^ in slightly off-polar view. Exine 6-7 p thick with
irregularly disposed shallow pits less than 1 p in diameter. The pitting is slightly more
pronounced in the vicinity of the proximal pole.

Remarks and comparisons. Punctatisporites fungosus may be distinguished from the
Lower Permian species P. gretensis Balme and Hennelly by its larger size and relatively
thicker, finely pitted exine. It occurred, usually in small numbers, in all the samples
examined and has also been found in the Blina Shale (Canning Basin, Western Australia).

Known stratigraphical range in Australia. Lower Triassic.

Genus osmundacidites Couper 1953

Type species. Osmundacidites wellmanii Couper, Upper Jurassic, New Zealand.

EXPLANATION OF PLATE 4

Magnifications X 600

Figs. 1-2. Osmundacidites senectus. 1, Holotype. 2, Paratype 47542.

Fig. 3. Tetraporina sp. cf. Azonotetraporinal horologia Staplin.

Figs. 4-5. cf. Schizosporis sp.

Fig. 6. Lycopodiacidites pelagius, holotype.

Fig. 7. Lycopodiacidites sp.

Figs. 8-9. Lundbladispora brevicida. 8, Paratype 47547. 9, Holotype.

Figs. 10-11. Punctatisporites fungosus. 10, Holotype. 11, Paratype, showing pitted and channelled exine.

Palaeontology, Vol. 6

PLATE 4

Bly ^

7M^%

iS^^SSi

BALME, Triassic plant microfossils

'ifi!,- a-

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

17

Osmundacidites senectiis sp. nov.

Plate 4, figs. 1, 2

Holotype. Slide 47541. Paratypes. Slides 47542 and 47543.

Diagnosis. Amb circular, periphery finely notched. Trilete, scar distinct, laesurae straight
and extending about three-quarters of the distance to the distal margin. Exine 1-2 p
thick, sculptural elements small and varied, including cones, grana, and sub-baculate
processes. Basal diameter of individual processes 1-3 p, height 1-2 p. Off-polar com-
pressions common, suggesting that the spore was originally spherical, and the flattened
exine usually has one or more broad, crescentic, folds.

Dimensions. Diameter 58-79 p; 25 specimens.

Locus typicus. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio itominis. Latin seuectus = ‘aged’.

Description. Holotype preserved in full polar view. Diameter 72 p, exine about 2 p thick
with irregularly distributed but fairly closely spaced sculptural elements. Among these,
short cones predominate but grana and sub-baculae also occur; eighty-nine processes
were counted around the periphery. In the holotype sculptural elements were less pro-
minent in the region of the proximal pole, but this reduction was not obvious in the
paratypes.

Remarks and comparisons. This species is assigned to Osmundacidites because of the
variability in form of its sculptural elements and its general conformity with Couper’s
diagnosis. Some hesitation is felt in extending so considerably the stratigraphical range
of a ‘half-natural’ genus, but Klaus (1960) has assigned Carnian spores to Osmundaci-
dites and the known antiquity of the family Osmundaceae may be invoked in justification
of the use of Couper’s genus.

Osmundacidites senectus has finer and more regular sculptural elements than O. well-
manii Couper or O. comaumensis (Cookson) and is larger than O. alpina Klaus. It is
fairly common in most samples from the Kockatea Shale and occurs also in the Blina
Shale and in the Erskine Sandstone, a continental Triassic formation which overlies the
Blina Shale. Occasional specimens have also been found in sediments from the Narra-
been Group in New South Wales.

Known stratigraphic range in Australia. Lower to (?) Middle Triassic.

Genus lycopodiacidites (Couper) R. Potonie 1956
Type species. Lycopodiacidites buUerensis Couper, Upper Jurassic, New Zealand.

Lycopodiacidites pelagius sp. nov.

Plate 4, fig. 6

Holotype. Slide 47551. Paratype. Slide 47550.

Diagnosis. Amb subtriangular to circular, periphery undulate. Trilete, scar clearly
defined, laesurae straight and extending about three-quarters of the distance to the

c

0 1015

18

PALAEONTOLOGY, VOLUME 6

equatorial margin. Exine 4-7 jx thick on the distal side, slightly thinner on the proximal.
Proximal face smooth or slightly rugulose. Distal face sculptured into low, rounded,
irregular and rather indistinct ridges. Ridges 2-7 ix wide and 1-3 jx high, occasionally
anastomosing to form a crude reticulum but usually separate from one another, giving
the distal surface an irregular rugose pattern.

Dimensions. Diameter 86-100 /x; 10 specimens.

Locus typicus. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio nominis. Latin pelagius = ‘pertaining to the sea’, from the wavy appearance of the distal
surface.

Description. Holotype diameter 97 p, exine 6-8 p thick, measured in optical section
along the equatorial margin. Laesurae about 35 p long and closed along their entire
length. Distal ridges 1-3 p high and 3-6 p wide, occasionally anastomosing and enclos-
ing rounded lumina.

Remarks and comparisons. Lycopodiacidites kuepperi Klaus, from the Upper Triassic of
Austria, is slightly smaller and has more clearly defined distal rugae. L. pelagius was not
common in the Kockatea Shale, but occasional specimens were recognized in most
samples. It has not been found in any other deposit.

Known stratigraphic range in Australia. Lower Triassic.

Figured specimen. Slide 47557.

Lycopodiacidites sp.
Plate 4, fig 7

Description. Amb strongly rounded triangular, periphery undulate. Trilete, scar strongly
defined, laesurae extending about half-way to the equatorial margin. Edges of laesurae
turned outwards forming a triangular opening at the proximal pole. Exine about 6 p
thick, with an indistinctly rugose distal surface; proximal face perforate. Perforations
less than 1 /a in diameter, irregularly distributed, and particularly concentrated around
the proximal polar area.

Dimensions. Diameter 79-91 p', 5 specimens.

Locality of figured speeimen. Well at Point 217, Upper Greenough River area (sample 44070), Western
Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. Lycopodiacidites sp. may represent L. pelagius in a corroded
condition, but it was too rare adequately to assess this possibility. It has not therefore
been named formally, but specimens maintaining the characters described above occurred
sporadically in several of the assemblages.

Known stratigraphic range in Australia. Lower Triassic.

Genus kraeuselisporites (Leschik) Jansonius 1962
Type species. Kraeuselisporites dentatus Leschik, Keuper, Switzerland.

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

19

Generic name. The inclusion of Kraeuselisporites in the Unterabteilung Zonaletes by
Leschik (1955) has led to difficulties in the application of his generic name. It was
rejected by Klaus (1960, p. 142), in favour of Styxisporites Cookson and Dettmann, on
the ground that Leschik’s description and interpretation were incorrect. Certainly the
original diagnosis of Kraeuselisporites was inadequate but the illustrations were fairly
clear, and, as Jansonius has noted, a weak trilete scar is visible on some of Leschik’s
figured specimens. Styxisporites and Kraeuselisporites have a similar structure, but the

TEXT-FIG. 2. a, Limdbladispora playfordi gen. et sp. nov. lateral view, b, L. playfordi diagrammatic
section parallel to the polar axis, c, Kraeuselisporites ciispidus sp. nov., lateral view, d, K. ciispidus
diagrammatic section parallel to polar axis. All x800 approx, a.p. apical papillae; exo. exoexine;

int. intexine.

form of the flange and lips seems to provide a means of distinguishing between the two
genera. In Styxisporites the flange is translucent to almost transparent and clearly de-
limited from the central body, and the prominent laesurae are bordered by fairly strong
lips.

Kraeuselisporites cuspidus sp. nov.

Plate 5, figs. 9, 10, 11 ; text -figs. 2c, d
Holotype. Slide 47552. Paratypes. Slides 47553 and 47554.

Diagnosis. Amb rounded triangular, periphery dentate. Exine cavate consisting of u
zonate exoexine enclosing a thin intexine. Trilete, scar weak but usually clearly visible,
laesurae extending to the inner margin of the zona. Distal side hemispherical, proximal
flattened pyramidal. Exoexine 2-3 p thick on distal side and 1 p or less on the proximal.

20

PALAEONTOLOGY, VOLUME 6

Surface of exoexine (including zona) scabrate; distal surface and zona bearing heavy
irregularly disposed spines 5-10 /x long, 5-6 /x in basal diameter, and 3-15 /x apart.
Intexine thin, smooth, partially or completely detached from the exoexine, and usually
folded.

Dimensions. Total diameter 71-112 /x, width of zona 9-25 ju.; 25 specimens.

Locus typicus. Kockatea Creek No. 19 Bore, 1 39-190 ft. (sample 43305), Upper Greenough River area.
Western Australia. Kockatea Shale, Lower Triassic.

Derivatio iiominis. Latin ciispis = ‘a point’.

Description. Holotype diameter 91 p, zona 12-16 p wide. Distal surface and zona bearing
about forty heavy spines. Spines about 8 p long with a basal diameter ranging from
3 to 5 ;ix. In the holotype the intexine is detached from the exoexine but unfolded. Para-
type 47553 has a zona of irregular width and a shrunken, rather crumpled intexine.

Remarks and comparisons. Kraeuselisporites cuspidiis is much larger than any of the
forms of Kraeuselisporites described by Leschik. Styxisporites cooksonae Klaus is of
comparable size, but its spines are confined to the distal surface and do not occur on the
zona. K. apicidatiis Jansonius is smaller and has more numerous spines than K. cuspidus.

Kraeuselisporites cuspidus was recorded from all the samples examined but was gener-
ally most common in material from the Upper Greenough River-Indarra area. It occurs
also in the Blina Shale but undoubted specimens have not been recognized in other
Triassic deposits. Other species of Kraeuselisporites occur in the Australian Permian.

Known stratigraphic range in Australia. Lower Triassic.

Kraeuselisporites saeptatus sp. nov.

Plate 6, figs. 8, 9, 10

Holotype. Slide 47549. Paratypes. Slides 47555, 47556, and 47561.

Diagnosis. Amb rounded triangular, periphery slightly ragged. Exine cavate consisting
of a zonate exoexine enclosing a thin intexine. Trilete, scar usually distinct, laesurae
straight with weak lip development visible on some specimens and extending to the
inner margin of the zona. Distal hemisphere strongly inflated, proximal face forming a
low pyramid (see PI. 6, fig. 8). Exoexine 2-3 p thick on the distal side, less than 1 p thick
on the proximal. Surface of exoexine (including zona) scabrate with a ‘scaly’ texture.

EXPLANATION OF PLATE 5

Magnifications X 600, except fig. 7.

Figs. 1-3. Lundbladispora willmotti. 1, Holotype, distal surface showing sculpture. 2, Holotype, proxi-
mal surface. 3, Paratype 47533, showing ruptured proximal face and spinose sculpture.

Figs. 4-8. L. playfordi. 4, Holotype. 5, Paratype 47838, with indistinct scar and faint apical papillae.
6, Paratype 47536, small specimen with slight lip development. 7, Specimen dissected intexine show-
ing apical papillae, x 700. 8, Adherent tetrad showing free intexine and equatorial thickening, Para-
type 47539.

Figs. 9-11. Kraeuselisporites cuspidus. 9, Holotype. 10, Paratype 47553, with irregular zona. 11,
Paratype 47554.

Palaeontology, Vol. 6

PLATE 5

BALME, Triassic plant microfossils

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

21

irregularly disposed grana sometimes present on the distal hemisphere. Zona gradually
becoming thinner towards its outer periphery. Intexine thin, smooth, with occasional
fine marginal folds but rarely crumpled.

Dimensions. Diameter 51-76 /x, width of zona 6-18 /x; 25 specimens.

Locus typiciis. Geraldton Racecourse Bore, 1,465 ft. (sample 44497), Western Australia. Kockatea
Shale, Lower Triassic.

Derivatio nominis. Latin saeptum = ‘an enclosure’.

Description. Holotype diameter 72 p, zona 10-13 p wide, diameter of intexine 42 p.
Intexine weakly folded around its margin and probably still adhering to the exoexine on
the proximal side. Paratype 47555 is preserved in lateral view and has a polar diameter
of 53 p.

Remarks and comparisons. Kraeuselisporites saeptatus dilfers from any species previously
assigned to Kraeuselisporites in its lack of distinct sculptural elements. However,
Jansonius (1962) has included in the genus forms with weakly developed distal grana
(cf. K. piinctatus Jansonius) and the present assignation does not seem an undue exten-
sion of his interpretation of Kraeuselisporites. Eventually it may become expedient
to create a new form genus for unsculptured forms of the K. saeptatus-iypQ, but this has
been deferred here in order to avoid excessive monotypy.

In lateral compressions K. saeptatus is easily distinguished from Densosporites by its
strongly inflated distal hemisphere and flattened proximal surface. Simplicisporites
laciniatus Leschik appears, from the original illustration (Leschik 1955, pi. 5, fig. 5),
to have a similar structure to K. saeptatus, and so do the two forms referred to Aequitri-
radites by Nilsson (1958, p. 47). Rogalska (1956, pi. 31) also has illustrated comparable
types from the Liassic of Poland.

Kraeuselisporites saeptatus is one of the most common species in the Kockatea Shale,
especially in the sediments from BMR 10 (Beagle Ridge) Bore. It occurs also in the
Blina Shale.

Known stratigraphical range in Australia. Lower Triassic.

Genus lundbladispora gen. nov.

Type species. Lundbladispora willmotti sp. nov.. Lower Triassic, Western Australia.

Diagnosis. Trilete spores with a maximum diameter of less than about 150 p. Exine
cavate with a finely structured exoexine enclosing a thin-walled intexine. Exoexine
thinner on the proximal than on the distal side and with a narrow equatorial thickening.
Surface of exoexine scabrate with a spongeous appearance. Distal side devoid of sculp-
ture or bearing small cones, spines, or grana. Sculptural elements absent or considerably
reduced on the proximal surface. Intexine thin-walled, smooth, with three apical papillae
(cf. Bharadwaj 1958) in the area of the proximal pole. Intexine often eccentrically placed
with respect to the equatorial margin of the exoexine.

Remarks and comparisons. The distinctive characters of Lundbladispora are as follows:
(a) The relatively thick finely textured exoexine with an equatorial thickening.

22

PALAEONTOLOGY, VOLUME 6

{b) The presence of a thin, frequently eccentrically placed, papillate intexine.

(c) Virtual restriction of sculptural elements, when they occur, to the distal surface
and equatorial region.

Dispersed megaspores of comparable structure to Limdbladispora have been described
from Palaeozoic and Mesozoic sediments and assigned to a number of form genera.
These include Duosporites Hoeg, Bose, and Manum, Banksisporites Dettmann, and
Bacutriletes (van der Hammen). Few previously described microspores closely resemble
Limdbladispora, but a possible exception is Aculeisporites variabilitis Jansonius, which
possesses a papillate intexine and a hnely granulose and spinose exoexine. Microspores
illustrated by Lundblad (1948) from the Lower Triassic of Greenland appear indis-
tinguishable from Limdbladispora playfordi; they were associated with the lycopo-
diaceous strobilis Selaginellites polaris and may be its microspores.

Cavate exines occur frequently in the spores of modern species of Selagi/iella (Knox
1950, Harris 1955) and they are known also in fossil spores obtained from lycopodiaceous
strobili (Lundblad 1948, 1950, Bharadwaj 1958). Another lycopodiaceous character of
Limdbladispora is the restriction of sculptural elements to the distal face (see Couper
1958, p. 105). In so far as one is justified in inferring affinities from spore morphography
the evidence favours lycopodiaceous, and possibly selaginellid, affinities for Limd-
bladispora.

The genus is named after Dr. B. Lundblad, Stockholm University, in recognition of
her contributions to our knowledge of Triassic floras.

Limdbladispora willmotti sp. nov.

Plate 5, figs. 1, 2, 3

Holotype. Slide 47532. Paratypes. Slides 47533, 47534.

Diagnosis. Amb circular or strongly rounded triangular, periphery spinose, exine cavate.
Trilete, scar indistinct, laesurae poorly defined, extending about two-thirds of the dis-
tance to the equatorial margin. Exoexine 4-6 g thick on the distal side, about 1 p thick,
and sometimes ruptured, on the proximal. Slightly depressed contact areas sometimes
visible on the proximal face. Exoexine scabrate, distal surface and narrow equatorial
thickening ornamented with cones or spines. Sculptured elements 1-2 p in basal dia-
meter, 1-3 p long and 1-10 p apart, absent or much reduced in size on the proximal face.
Intexine smooth and thin-walled, bearing weak tetrad markings with three apical papil-
lae, one in each inter-radial area.

Dimensions. Total diameter 71-86 p, diameter of intexine 49-65 p; 25 specimens.

Locus typicus. Kockatea Creek No. 19 Bore, 139-190 ft.. Upper Greenough River area (sample 43305),
Western Australia. Kockatea Shale, Lower Triassic.

Derivatio nominis. After Mr. S. P. Willmott, geologist of West Australian Petroleum Pty. Ltd.

Description. Holotype diameter 78 p, diameter of intexine 53 p. Proximal face extremely
thin and translucent, bearing a faint, slightly sinuous tetrad scar. Sculptural elements 2 p
in basal diameter, 1-3 p high and irregularly disposed. In paratype 47533 the proximal
face is ruptured and the spines are fewer and more prominent, ranging up to 6 ft long.

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

23

Paratype 47534 is preserved in semi-lateral view and has a total diameter of 86 ft and
a polar diameter of about 81 ft.

Remarks and comparisons. Limdbladispora willmotti is distinguished from L. playfordi
by its spinose sculpture, larger size, and the coarser texture of its exoexine. It was present
in all samples from the Kockatea Shale and occurs also in the Blina Shale.

Known stratigraphical range in Australia. Lower Triassic.

Lundbladispora playfordi sp. nov.

Plate 5, figs. 4-8; text-fig. la, b

Holotype. Slide 41537. Paratypes. Slides 47538, 47536, and 47839 (tetrad).

Diagnosis. Amb circular or strongly rounded triangular, periphery smooth, exine cavate.
Trilete, scar indistinct or clearly defined, lips sometimes present, laesurae straight or
slightly curved, length variable but usually extending almost to the equatorial margin.
Exoexine scabrate, 2-4 p thick on the distal face, slightly thinner on the proximal,
thickened equatorially in a zone about 3-5 p wide. Intexine thin, smooth, frequently
folded, and bearing three apical papillae in the region of the proximal pole (PI. 5, fig. 7).
In polar compressions the intexine is usually eccentrically placed with respect to the
equatorial margin of the exoexine. Exoexine and intexine detached (PI. 5, fig. 8).

Dimensions. Total diameter 40-71 p, diameter of intexine 24-48 p; 25 specimens.

Locus typicus. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio nomiiiis. After Dr. P. E. Playford, now of the Geological Survey of Western Australia.

Description. Holotype diameter 66 p, diameter of intexine 42 p, equatorial thickening
2-3 ft wide. Eaesurae poorly defined, 20-22 p long, and curved. Paratype 47839 consists
of four spores in an adherent tetrad; these show the species to be biconvex with a polar
diameter of about 60 p. In Paratype 47838 the tetrad scar is scarcely visible but the
proximal face is ruptured, exposing the papillate intexine.

Remarks and comparisons. Lundbladispora playfordi was abundant in all the samples
examined from the Kockatea Shale and has also been found in the Blina Shale and rarely
in the Erskine Sandstone. Its resemblance to spores from the Lower Triassic of Green-
land has already been noted, but no other obviously similar forms have been previously
described. Endosporites papillatus Jansonius has a detached, papillate intexine, but the
Canadian species is said to have an extremely thin exoexine and is devoid of equatorial
thickening.

Known stratigraphical range in Australia. Lower to (?) Middle Triassic.

Lundbladispora brevicula sp. nov.

Plate 4, figs. 8, 9

Holotype. Slide 47546. Paratypes. Slides 47547 and 47548.

24

PALAEONTOLOGY, VOLUME 6

Diagnosis. Amb rounded triangular, periphery spinose, exine cavate. Trilete, scar dis-
tinct, laesurae curved or sinuous, with narrow elevated lips and extending almost to the
equatorial margin. Exoexine 2-3 /x thick on distal side, I g or less on the proximal with
an equatorial zone of thickening 2-4 jU. wide. Surface of the exoexine finely spongeous ;
ornamented on the distal side and along the equatorial margin, with sparsely dis-
tributed cones and spines. Sculptural elements about 2 ixin basal diameter, 1-4 ju. high.
Intexine thin, smooth and with three faint apical papillae sometimes visible. The intexine
is never notably eccentric and is apparently attached proximally to the exoexine.

Dimensions. Total diameter 41-56 ju,, diameter of intexine 29-39 /x; 20 specimens.

Locus typiciis. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio nominis. Latin breviculiis = ‘rather small’.

Description. Holotype diameter 51 ;u-, diameter of intexine 36 p. Laesurae sinuous with
raised lips about 2 p wide on either side of the commissure. Sculpture consists of about
sixty irregularly disposed cones and spines, 1-3 p in basal diameter and 2-4 p high.
Except for slight size differences the paratypes show little variation from the holotype.

Remarks and comparisons. Lundbladispora brevicida is distinguished from L. willmotti
by its smaller size, prominent tetrad scar, and relatively larger spines. It was present in
all the samples examined and most common in material from the Upper Greenough
River area. The only other Australian sediment in which it is known to occur is the
Blina Shale.

Known stratigraphical range in Australia. Lower Triassic.

Anteturma pollenites R. Potonie
Turma saccites Erdtman
Subturma disaccites Cookson

Genus vitreisporites Leschik 1955 (= Caytonipollenites Couper 1958)

Type species. Vitreisporites signatiis Leschik, Keuper, Switzerland.

Vitreisporites pallidas (Reissinger) Nilsson 1958
Plate 6, fig. 7

1938 Pityopollenites pallidus Reissinger, Palaeontographica, 84b, 84.

1950 Pityosporites pallidus (Reiss.) Reissinger, Palaeontographica, 90b, 109; pi. 15, figs. 1-5.
1958 Caytonipollenites pallidus (Reiss.) Couper, p. 150; pi. 26, figs. 7-8.

1958 Vitreisporites pallidus (Reiss.) Nilsson, p. 78; pi. 7, figs. 12-14.

Figured specimen. Slide 47529.

Description. Disaccate pollen grain. Central body elongate oval, proximal face infra-
granulate, distal germinal area smooth and very thin. Sacci microreticulate, slightly
inclined distally, and a little longer than the central body.

Dimensions. Length of body 16-25 p, breadth of body 12-21 p, total breadth of grain
26^0 p; 4 specimens.

B. E. BALME: TRIASSIC PLANT MICROEOSSILS

25

Locality of figured specimen. Well at Point 217, Upper Greenough River area (sample 44070), Western
Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. In form and size these Triassic specimens fall within the range
of Cooper’s diagnosis of CaytonipoIIenites pallidiis. Quantitatively the species is un-
important in the Kockatea Shale, but has been recorded because of its possible palaeo-
botanical interest. Vitreisporites was described by Jansonius from the Scythian of
Canada, but in Europe the oldest clear record is in Keuper strata (Leschik 1955).
Pollens similar to Vitreisporites pallidas occur rarely in Upper Permian strata in the
Canning Basin, but are not common before the Lower Jurassic.

Known stratigraphical range in Australia. Upper Permian to Lower Cretaceous.

Genus platysaccus (Naumova) R. Potonie and Klaus 1954
Type species. Platysaccus papilionis R. Potonie and Klaus, Zechstein, Austria.

Platysaccus sp. cf. P. papilionis R. Potonie and Klaus 1954

Plate 6, fig. 12

1954 Platysaccus papilionis R. Potonie and Klaus, p. 539; pi. 10, figs. 11, 12.

1955 Lueckisporites fusus [pars] Balme and Hennelly, p. 92; pi. 1, figs. 6, 9.

Figured specimen. Slide 47527.

Description. Disaccate, strongly diploxinoid pollen grain. Central body subcircular,
thick-walled, proximal surface smooth or slightly rugulose. Sacci much larger than the
central body, microreticulate, attached distally with their inner bases separated by a
narrow germinal area.

Dimensions. Diameter of central body 47-53 p, length of sacci 71-87 p, total breadth of
grain 104-127 p; 8 specimens.

Locality of figured specimen. Kockatea Creek No. 19 Bore, 139-190 ft.. Upper Greenough River area
(sample 43305), Western Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. Platysaccus cf. papilionis falls within the broad limits pro-
posed for the original species by Potonie and Klaus. It occurred sporadically in the
Kockatea Shale and provides one of the few links between Permian and Triassic micro-
floras in Western Australia.

Platysaccus papilionis has been recorded by Jansonius from the Scythian of Canada
and by several authors from the European Zechstein. Pollen grains of apparently similar
structure occur also in the Polish Liassic (Rogalska 1956, pi. 21, fig. 1).

Known stratigraphical range in Australia. Lower Permian to (?) Middle Triassic.

Infraturma striatiti Pant
Genus striatites (Pant) Pant 1955

Type species (designated by Pant). Pityosporites sewardi Virkki, Upper Permian, New South Wales,
Australia.

Generic name. Striatites is one of the genera assigned by Potonie (1958) to the Infra-
turma Striatiti which includes all fossil, disaccate pollen grains bearing transverse bands

26

PALAEONTOLOGY, VOLUME 6

of exoexinal thickening on their proximal faces. Such pollen grains are particularly charac-
teristic of Permian microfloras from the ‘Gondwanaland’ countries and are common also
in Permian and Triassic sediments from the northern hemisphere. Because of their great
variety and abundance it is difficult to establish a satisfactory systematic scheme for the
Striatiti and many ill-conceived and inapplicable taxa have appeared in palynological
literature. It is obvious that the taxonomy of the whole group needs revision, but this
cannot be attempted without considering the great diversity of Permian forms. In the
present account the generic name Striatites is used to accommodate members of the
Striatiti with well-developed sacci and more than about six bands of proximal thicken-
ing. It therefore includes forms which Potonie placed in Striatites, Limatisporites, and
Striatopodocarpidites.

Striatites sp. cf. Taeniaesporites antiquus Leschik 1956

Plate 6, fig. 13

1956 Taeniaesporites antiquus Leschik, p. 134; pi. 22, fig. 4.

Figured specimen. Slide 47526.

Description. Disaccate, sacci joined equatorially in some specimens, haploxinoid
pollen grains. Central body oval in polar view. Proximal face of central body bearing
about fifteen transverse bands of exoexinal thickening separated by narrow clefts in
which the intexine is exposed. Proximal bands scabrate and 5-9 p wide. Sacci
fairly large, attached and inclined distally, separated on the distal side by a thin ger-
minal area consisting of exposed intexine. Sacci microreticulate with clearly defined,
slightly radially elongate lumina.

Dimensions. Length of body 76-1 14 ;u,, breadth of body 58-88 p, length of sacci 81-116 p;
breadth of sacci 49-61 p; 5 specimens.

Locality of figured specimen. Well at Point 217, Upper Greenough River area (sample 44070), Western
Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. In its large size and occasional pseudo-monosaccate appear-
ance Striatites sp. resembles Lueckisporites richteri Klaus from the German Zechstein.
However, Klaus’s species has heavier and better-defined proximal thickenings. The
specimens of Taeniaesporites antiquus illustrated by Leschik look similar to some ex-
amples of Striatites sp., but Leschik’s diagnosis is imprecise and detailed comparisons
are not possible.

EXPLANATION OF PLATE 6

Magnifications X600, except fig. 13.

Figs. 1-3. Taeniaesporites obex. 1, ITolotype, proximal side. 2, Holotype, distal side. 3, Paratype 47558,
showing transverse, proximal scar.

Figs. 4-6. Taeniaesporites sp. cf. T. noviaulensis Leschik.

Fig. 7. Vitreisporites pallidus (Reissinger).

Figs. 8-10. Kraeuselisporites saeptatus. 8, Holotype. 9, Paratype 47555, lateral view. 10, Specimen
showing outline of the intexine.

Fig. 11. Crustaesporites sp.

Fig. 12. Platysaccus sp. cf. P. papilionis Potonie and Klaus.

Fig. 13. Striatites sp. cf. Taeniaesporites antiquus Leschik. X440.

Palaeontology, Vol. 6

PLATE 6

BALME, Triassic plant microfossils

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

27

Striatites sp. is assignable to Limalisporites as that genus was interpreted by Potonie
(1958), but recent doubts have been expressed as to the validity of Liiiiatisporites (Klaus
1960, Dr. G. F. Hart, personal communication) and use of the name should be dis-
continued pending clarification of its status. No form clearly identical with Striatites sp.
is known from the Australian Permian, but similar types occur rarely in the Blina Shale.

Known stratigraphical range in Australia. Lower Triassic.

Genus taeniaesporites (Leschik) Jansonius 1962
Type species. Taeniaesporites kraeuseli Leschik, Keuper, Switzerland.

Generic name. The genus Taeniaesporites was proposed by Leschik (1955, p. 38) on the
basis of the following brief diagnosis: ‘ Mikrosporen mit zwei Luftsacken. Der sporen-
korper ist durch 6 und mehr Streifen zerlegt.’ These characters scarcely justified the
establishment of a new genus, for they do not distinguish Taeniaesporites from several
pre-existing form genera. Leschik’s subsequent attempt (Leschik 1956) to change the
type species of Taeniaesporites did nothing to clarify the application of his generic name.

Potonie did not accept Taeniaesporites on the ground that Leschik had misconstrued
the structure of T. kraeuseli, which, Potonie argued, had only three proximal striae and
belonged in Lueckisporites R. Potonie and Klaus. Leschik’s photographs bear out
Potonie’s interpretation and any attempt to prevent the proliferation of generic names
among the Striatiti commands sympathy. Nevertheless, T. kraeuseli was adequately
illustrated and is a well-characterized and distinctive species. It could certainly have been
included in Lueckisporites as that genus was originally conceived, but having accepted
the restriction of Lueckisporites to pollens of the L. virkkiae type, it was not altogether
consistent for Potonie to reject Taeniaesporites.

Leschik’s views on the scope of his genus have never been clearly expressed, but he
has vigorously defended the separation of Taeniaesporites and Lueckisporites on mor-
phographic and stratigraphic grounds. His arguments are strongly supported by the
evidence from Western Australia. Here forms similar to Taeniaesporites kraeuseli and
T. noviaulensis are common in Lower Triassic sediments, but pollens resembling Luec-
kisporites virkkiae have never been observed.

Jansonius (1962) has recently proposed an emendation of Taeniaesporites which
restricts and clarifies the genus. In this the generic name Taeniaesporites is restricted to
striatitid forms with less than six, and most commonly four, bands of transverse proximal
thickening. These exoexinal bands are commonly inflated and separated by transverse
clefts in which the intexine is exposed. The clefts are wider and more clearly defined than
in other genera of the Striatiti. Under the diagnosis proposed by Jansonius both hap-
loxinoid and diploxinoid forms may be assigned to Taeniaesporites, for he minimizes
the relative size of body and bladder as a criterion in the classification of the Striatiti.
One may agree with Jansonius that many Russian authors have over-emphasized this
character, without subscribing unreservedly to his view. In the future it may prove con-
venient to reserve Taeniaesporites for haploxinoid forms and create a new genus for
thick-walled diploxinoid species such as Taeniaesporites transversundatus Jansonius and
T. obex sp. nov.

28

PALAEONTOLOGY, VOLUME 6
Taeniaesporites sp. cf. T. noviaiilensis Leschik 1956
Plate 6, figs. 4-6

1956 Taeniaesporites noviaiilensis Leschik, p. 134; pi. 22, figs. 1, 2.

71962 Taeniaesporites novimiindi Jansonius, p. 63.

Figured specimens. Slides 47522, and 47524.

Description. Disaccate pollen grain. Central body subcircular or oval in polar view,
intexine thin and translucent, exoexine confined to the proximal side, infrareticulate and
slightly thicker than the intexine. Exoexine divided by three, or sometimes four, trans-
verse clefts in which the intexine is exposed; the proximal side, therefore, has four or
five bands of transverse exoexinal thickening. A short, transverse, linear, scar is usually
visible within the cleft which passes through the proximal pole and from this it is inferred
that the species was formed in planar tetrads. Sacci attached, and slightly inclined
distally. Structure of sacci microreticulate ; mesh lumina fine near the bases of the sacci,
becoming coarser towards their peripheries. Distal germinal area fairly broad, con-
sisting of thin, smooth intexine.

Dimensions. Maximum diameter of central body 42-62 /x, breadth of sacci 26-36 /x, length
of sacci 43-70 p, total breadth 72-90 /x; 30 specimens.

Locality of figured specimens. Well at Point 217, Upper Greenough River area (sample 44070), Western
Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. Specific discrimination within the genus Taeniaesporites is not
easy, partly because of the rigidity of the emended diagnosis and partly because the
forms encountered have a fairly wide range of apparently continuous variation, both
in size and in the arrangement of the proximal thickenings. Jansonius recognized nine
species from the Canadian Lower Triassic, but some of these are rather arbitrarily
characterized and may prove difficult to maintain. A comparatively wide range of
variation has been allowed in the specimens assigned here to Taeniaesporites cf. noviau-
lensis and this latitude may be gauged from the three specimens illustrated in Plate 6,
figs. 4-6.

The Western Australian form is similar in size to Leschik’s species which it also
resembles in arrangement and structure of its proximal thickenings and the presence of a
short, transverse, tetrad scar. A possible distinction lies in the internal structure of the
sacci. In Taeniaesporites noviaiilensis the lumina are radially elongate. The mesh is
variable in the Australian species and appears coarser in poorly preserved specimens.
There is some uncertainty, therefore, concerning its reliability as a specific character.

Taeniaesporites cf. noviaulensis is a key species in the microflora of the Kockatea
Shale. It is easily recognized and occurred in all samples, usually making up between 3
and 10 per cent, of the total spore-pollen assemblage. It occurs commonly in the Blina
Shale, less frequently in the Erskine Sandstone and in continental Triassic sediments
overlying the Kockatea Shale in BMR 10 (Beagle Ridge) Bore. Rare specimens have
been found in Triassic coals from the Springfield Basin, South Australia, but no repre-
sentatives of Taeniaesporites have been recorded from the Leigh Creek Coal Measures.
In the Sydney Basin, New South Wales, a few examples have been recognized in as-
semblages from the Collaroy Claystone (Middle Narrabeen Group), and similar forms

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

29

occur red fairly commonly in a sample from the base of the Narrabeen Group, collected
in the Nattai River area.

The distribution of Taeniaesporites is discussed further in a subsequent section of
this paper.

Known stratigraphical range in Australia. Lower to (?) Upper Triassic (mainly Lower
Triassic).

Taeniaesporites obex sp. nov.

Plate 6, figs. 1-3

Holotype. Slide 47531. Paratype. Slide 47558.

Diagnosis. Disaccate, more or less diploxinoid pollen grain. Central body circular in
polar view, intexine very thin, translucent and faintly punctate. Proximal face bearing
four prominent transverse bands of heavy exoexinal thickening, separated by clefts in
which the intexine is exposed. Subsidiary clefts sometimes present in the thickenings
giving rise to isolated patches of discontinuous exoexine. Thickenings infragranulate and
about 2 p thick. A short transverse, linear scar passes through the proximal pole. Sacci
crescentic in polar compressions, longer than the central body, attached and inclined
distally. Sacci microreticulate, mesh clearly defined, lumina radially elongated near the
base of the sacci, becoming more equidimensional towards their outer margins.

Dimensions. Diameter of central body 36-43 p, breadth of sacci 25-37 p, length of sacci
43-57 p, total breadth 69-81 p.

Locus typicus. Well at Point 217, Upper Greenough River area (sample 44070), Western Australia.
Kockatea Shale, Lower Triassic.

Derivatio nominis. Latin obex = ‘a barred gate’, from the appearance of the proximal surface.

Description. Holotype dimensions — diameter of central body 38 p, sacci 30 x45 p and
and 29 x46 p, total breadth 72 p. The holotype is only slightly compressed and bears
four bands of thickened exoexine. An elongated patch of discontinuous thickening occurs
between the two major bands and is visible at the right-hand side of the central body in
Plate 6, fig. 1 . Thickenings 6-8 p wide, with notched margins, and infragranulate sculp-
ture. The paratype has a wider polar cleft than the holotype and displays a clear trans-
verse scar.

Remarks and comparisons. Taeniaesporites obex is fairly rare in the Kockatea Shale and
did not occur m all the samples. It is known from the Blina Shale and similar forms occur
in the lower part of the Narrabeen Group in New South Wales. The circular central body
and heavier proximal thickenings distinguish it from T. noviaulensis.

No species obviously resembling Taeniaesporites obex has been described from Euro-
pean sediments, but Jansonius (1962) has recorded forms with heavy discontinuous
proximal thickenings from the Scythian of Canada. Of these, T. transversimdatus is
closest to the Australian species but it is smaller than T. obex and its proximal thickenings
are markedly constricted away from the pole.

Known stratigraphical range in Australia. Lower Triassic.

30 PALAEONTOLOGY, VOLUME 6

Genus crustaesporites Leschik 1956

Type species. Crustaesporites globosus Leschik, Zechstein, Germany.

Figured specimen. Slide 47525.

Crustaesporites sp.
Plate 6, fig. 1 1

Description. Trisaccate or irregularly disaeeate pollen grain. Central body oval, intexine
thin, finely punctate and translucent; proximal side bearing faint, roughly parallel bands
of exoexine. Bands infrareticulate, 2-5 p- wide and about 1 p apart. Sacci attached
marginally slightly on the distal side of the equator and inclined distally. Sacci narrow
and elongate, occasionally lobed; microreticulate with a clearly defined mesh.

Dimensions. Central body 91 x78 p, sacci 73 x31 p, 76 x27 p, 90 x27 p, figured speci-
men only.

Locality of figured specimen. Kockatea Creek No. 19 Bore, 139-190 ft.. Upper Greenough River area
(sample 43305), Western Australia. Kockatea Shale, Lower Triassic.

Remarks and comparisons. Crustaesporites sp. is rare in the Kockatea Shale. Representa-
tives of the genus have been found in Australian Upper Permian deposits (cf. Balme
and Hennelly 1955, pi. 4, fig. 44) and are known also from the German Zechstein (Le-
schik 1955, Klaus 1955, Grebe 1957) and the Lower Triassic of Canada (Jansonius 1962).

Jansonius interpreted Crustaesporites as a monosaccate form in which the sacci were
irregularly constricted to give a multisaccate appearance. In the few specimens seen in
microfloras from the Kockatea Shale the sacci were clearly separated, but the species
was too rare to judge its range of variation. However, it is certainly possible to envisage
a graduation from monosaccate to multisaccate specimens. Crustaesporites has clear
affinities with the striatitid pollens and not, as Leschik suggested, with the Podosporites-
Microcachryidites group.

INCERTAE SEDIS

Genus schizosporis Cookson and Dettmann 1958
Type species. Schizosporis reticulatus Cookson and Dettmann, Cretaceous, South Australia.

cf. Schizosporis sp.

Plate 4, figs. 4, 5

Figured specimens. Slide 47530.

Description. Spheroidal body of uncertain function. Body wall 2-4 p thick, two-
layered, inner layer very thin and smooth. Outer layer unsculptured, sometimes with
occasional scattered pores. Most specimens are split roughly into two halves, along a
line of weakness in the body wall. Some, however, are unruptured and in others the
incipient split appears as a faint line. The form resists compression and most specimens
are strongly three-dimensional.

Dimensions. Diameter 18-41 p; 60 specimens.

Locality of figured specimens. Geraldton Racecourse Bore, 1,465 ft. (sample 44497), Western Australia.
Kockatea Shale, Lower Triassic.

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

31

Remarks and comparisons. The generic assignation is tentative as the Triassic form is
considerably smaller than any included in Schizosporis by Cookson and Dettmann
(1958). It is similar in general structure to Schizosporis spriggi Cookson and Dettmann,
but is smaller and has a relatively thicker body wall. Jansonius (1962) has assigned similar
bodies to the genus SpheripoUenites (Couper) Jansonius, but judging from Couper’s
original diagnosis and illustrations (Couper 1958, p. 158, pi. 31, fig. 12) the propriety of
Jansonius’s emendation is dubious. SpheripoUenites scabratus Couper, the type species,
was said by Couper to resemble the pollen grain of PagiophyUum connivens, except in
its lack of a thickened equatorial zone. From this comment, and from the original
specific diagnosis, S. scabratus appears close to the Western Australian Jurassic species
Exesipollenites tumulus Balme, and is certainly distinct from Laevigatosporites scissus
Balme and Hennelly, which Jansonius includes in SpheripoUenites.

From its morphography and its distribution in the Kockatea Shale cf. Schizosporis sp.
is unlikely to be the spore or pollen grain of a vascular plant. It is usually extremely
abundant in samples with a high microplankton content, and is rare or absent else-
where.

SpheripoUenites balmei Jansonius may be identical with Schizosporis sp., but the
Canadian species is said to have a faint intrapunctate or interpunctate pattern within
the body wall. This feature was not convincingly exhibited by any specimens of cf.
Schizosporis sp. which have been closely studied.

Known stratigraphical range in Australia. Lower Triassic.

Genus tetraporina Naumova 1950

Type species. Tetraporina antiqua Naumova 1950, Lower Carboniferous, U.S.S.R. (designated Potonie
1960, p. 130).

Tetraporina sp. cf. Azonotetraporina ? horologia Staplin 1960

Plate 4, fig. 3

1960 Azonotetraporina ? horologia Staplin, p. 6; pi. 1, figs. 4, 6.

Figured specimen. Slide 47528.

Description. Test quadrilateral, two opposite sides concave, others straight or slightly
convex. Surface smooth or faintly punctate, thickness of test about 1 p. Arcuate folds
about 10 p long at each corner of the test.

Dimensions. Maximum length 36-46 p; 8 specimens.

Locality of figured specimen. Geraldton Racecourse Bore, 1,465 ft. (sample 44497), Western Australia.
Kockatea Shale, Lower Triassic.

Remarks and comparisons. Tetraporina sp. belongs to a group of morphographically
similar bodies, of obscure origins, which is widely distributed in Carboniferous sedi-
ments (Naumova 1950, Teteriuk 1958, Staplin 1960). It is rare in the Kockatea Shale,
but its occurrence extends the stratigraphical and geographical range of Tetraporina.
Churchill (1960) has reported the presence of a form closely resembling Tetraporina sp.

32 PALAEONTOLOGY, VOLUME 6

in modern lake sediments and considers that it is the aplanospore of a member of the
Cyanophyceae.

Known stratigraphical range in Australia. Permian to (?) Recent.

MICROPLANKTON

Except in one core (2,131-2,141 ft.) from BMR 10 (Beagle Ridge) Bore, all samples
from the Kockatea Shale were characterized by extraordinarily large numbers of micro-
plankton. Almost invariably they were many times more plentiful than spores or pollen
grains, and a single slide from Gerldton Raceourse Bore (sample 44497) was estimated
to contain over 25,000 specimens of hystrichosphaerids and leiospheres. Considered
individually the various forms present are not particularly distinctive, for they are simple,
spinose hystrichosphaerids, not differing obviously from species known to range from
the Silurian onwards. On the other hand, both de Jekhowsky (1961) and Jansonius (1962)
have noted that microplankton are particularly common in Lower Triassic marine
sediments from widely separated parts of the world. Jansonius has already remarked on
similarities between the microplankton suites of the Kockatea Shale and the Canadian
Toad-Grayling Formation. All the common Western Australian forms are closely
similar to species described by Jansonius, although the Canadian suites are more diverse,
and in addition to hystrichosphaerids contain dinoflagellate tests. The following list
includes all the common microplankton types in the Kockatea Shale, and is presented
without taxonomic comment.

Wilsonastrum colonicum Jansonius abundant

Wilsonastnim spp. abundant

Miahystridiiim setasessitante Jansonius common
M. breve Jansonius common

M. sp. cf. M. uiconspieiaim (Deflandre) common

Colonial unicellular thallophytes similar to Botryococcus were common in some
samples, particularly those from the Upper Greenough River area.

COMPOSITION OF THE ASSEMBLAGES

Text-fig. 3 shows the quantitative distribution of spore and pollen species in the
seventeen samples studied. These figures are based on counts of about 200 specimens,
except for three samples in which spores were rare and poorly preserved. In addition, the
relative proportions of spores (including pollen grains) to microplankton have been
estimated for each assemblage and these are expressed as a fractional ratio in the final
column of text-fig. 3. [

Except in the uppermost sample from BMR 10 Bore microplankton always out- 1
number spores, although their relative dominance varies within wide limits. The j
assemblages from BMR 10 Bore show gradually decreasing microplankton propor-
tions towards the upper part of the section and a similar trend is present in the Geraldton
Racecourse and Municipal Bores and in the Al\ Mile Peg Bore; although these were not
sampled in sufficient detail to enable confident interpretations. Frequency of micro-
plankton in a sediment may be influenced by a variety of factors the relative importance

B. E. BALME: TRIASSIC PLANT MICROFOSSILS

33

LOCALITY

Point 217 Bore

Kockatea Creek No. 19 Bore

6*

0
CO

o>

V

CL

1

Geraldton Racecourse Bore

Geraldton Municipal Bore

O

<n

o «

— o>

XJ

5

2

(ci ctf
*o»
o

(D

IN FEET

SPECIES

o

cn

139 — 190

uncertain

uncertain

741 — 749

800 — 945

940 — 968

1273

146 5

1012— 1096

1177 — 1344

1346 — 1366

2131 — 2141

2223 — 2233

2405—2415

2802—2012 1

3203 -3213

SPORES

Punctatisporites fungosus

5

3

X

2

5

1

5

6

12

4

9

7

4

34

5

3

2

Osmundacidites sensctus

26

6

3

6

2

X

3

1 I

lO

4

6

4

4

24

4

3

3

Lycopodiaciditss pelagiuii

2

2

2

1

X

1

1

1

2

2

I

2

2

I

1

I

Lycopodiocites gp.

1

1

X

1

1

1

I

1

2

1

X

X

I

Krasuselisporitss cujipidus

1

lO

16

13

lO

X

8

3

16

4

4

8

4

lO

5

5

I

K. sQc^tatu6

3

36

20

24

36

9

24

21

58

76

34

39

76

106

104

31

5

Lundbladispora playfordi

42

98

lO!

85

129

S2

las

127

43

71

9§

09

71

91

63

58

15

L . willmotti

24

13

7

9

7

2

13

24

12

3!

13

21

31

11

7

9

5

L . brevicula

39

!6

26

38

6

1

!8

S

4

18

2

6

18

8

2

0

2

POLLEN GRAINS

Vitreisporites pallidus

1

!

X

1

2

X

I

1

Platysaccus cf. papilionis

1

X

2

I

1

X

I

X

1

Striatitcs sp.

4

I

4

!

X

1

4

3

4

2

S

4

4

X

X

I

Tatniaesporitcs cf.
noviaulcnsis

60

22

23

34

7

3

II

16

19

16

20

17

16

11

14

5

7

T. ob«x

4

4

6

9

1

1

2

3

1

3

6

5

3

5

4

I

2

CruitoEsporites sp.

4

1

1

X

1

X

X

X

X

X

total count

2U

2!5

210

225

207

69

211

221

18!

235

199

205

235

307

209

124

4S

SPORE,,..-'^

.-^^mIcroplankton ratio

24

V

IS

1^

7

1^

2

o\

if

37

6

^5

60

I

i/

15

is

TEXT-FIG. 3. Distribution of spore and pollen species in samples from the Kockatea Shale.

C 1015 D

34

PALAEONTOLOGY, VOLUME 6

of which is difficult to assess, particularly when dealing with a group of such divergent
origins as the Hystrichosphaeridae. Reeent studies (Muller 1959, Staplin 1961) suggest
that the most favourable environment for the preservation of large numbers of hystri-
chosphaerids is a fairly deep, offshore basin in which turbidity is low. The most direct
environmental interpretation of the palynologieal data from BMR 10 Bore is that the
depositional basin gradually shallowed during the formation of the Kockatea Shale.
Lithological evidence presented by McTavish (1960) favours the same concept, for
coarser-grained sediments become more common in the upper part of the seetion, and
the uppermost 100 feet or so are characterized by lingulid brachiopods, worm tubes,
wood fragments, and other concomitants of the deltaic environment.

A singular feature of both the microplankton suites and the accompanying spore-
pollen assemblages is their remarkable lack of diversity. Almost invariably Wilsonastriim
was the dominant microplankton genus and in most samples the only other common
forms were one or two species of Micrhystridium. The microplankton suite, consisting
of few speeies and enormous numbers of individuals, suggests, therefore, a restricting
but highly productive environment. High salinity may have been the main cause of this
restriction, and could also be invoked to explain the general paucity of invertebrate
fossils in the Kockatea Shale. Nevertheless, it would be unwise to imply that salinity
was the only possible controlling factor.

Lack of variety in the spore-pollen assemblages is even more marked than a casual
glance at text-fig. 3 would suggest, for of the fifteen species described only nine were
present in every sample. Limdbladispora playfordi was almost always the most abundant
species, and spores, especially forms with cavate exines, always outnumber pollen
grains. In both paucity of species and relative abundance of pteridophyte spores, micro-
floras from the Kockatea Shale contrast notably with those from Australian Permian
sediments. Assemblages from the Upper Permian in particular, are characterized by
their heterogeneity and by a high content of striatitid, disaccate pollen grains (Balme
1962).

It may be argued that impoverishment of the Lower Triassic microfloras is a function
of the conditions of deposition of the Koekatea Shale and does not necessarily imply
specialization of their parent floras. Almost certainly the plant microfossils have been
carried long distances from their sources and selective forces undoubtedly operated dur-
ing transportation. The abundance of Kraeuselisporites and Limdbladispora, genera of
probable lycopsid origin, suggests, for example, that elements of coastal swamp floras
were heavily represented in the plant microfossil assemblages.

Even allowing for the possibility of selective transportation and preservation, however,
it seems unlikely that these factors alone can explain the impoverishment in species of
these Lower Triassic microfloras. Such impoverishment characterizes assemblages from
the Kockatea Shale wherever it has been sampled, even in areas which must have lain
close to the shoreline. Qualitatively similar microfloras have, in addition, been recovered
from the Blina Shale, the closest outcrop of which lies about 1,000 miles north of Gerald-
ton. Inadequate as the palynologieal data may be from the phytogeographic point of
view, they can most reasonably be interpreted as recording the presence of highly
specialized floras in western marginal areas of the Australian continent during early
Triassic times.

Phytogeographic changes of the magnitude postulated can only have resulted from

B. E. BALME: TRIASSIC PLANT MICROEOSSILS

35

major climatic changes at the end of the Permian. It would be rash to attempt a dog-
matic inference on the direetion of these changes from the palynological evidence alone,
although the sudden appearance, in the Western Australian Scythian, of Taeniaesporites
and other European Zechstein and Triassic forms suggests increasing aridity. The
lithology of the Kockatea Shale is compatible with a desertic climate in the adjoining
mainland, as it is predominantly fine-grained and contains red layers in Geraldton
Racecourse Bore. Clearly this ferric iron cannot have resulted from conditions within
the basin of deposition, for even the red shales contain abundant plant microfossils
which could not have survived prolonged oxidation. It may be concluded that most of
the iron was transported in the ferric state, and partially or completely reduced after
deposition in a marine environment.

Nowadays it would be reactionary to invoke a desertic climate to explain the origin of
red coloration in sediments. On the other hand, many authorities (Dunham 1953, Dun-
bar and Rodgers 1958, p. 217) are agreed that arid conditions favour, and may even be
essential to, the preservation of ferric oxides during prolonged transportation.

PHYTOGEOGRAPHIC AND STRATIGRAPHIC SIGNIFICANCE OF THE

MICROFLORAS

Because of the precision with which the Kockatea Shale can be dated its microfloras
have an intrinsic importance to both stratigraphers and palaeobotanists, and when the
unusual composition of the assemblages is also taken into account, their significance is
further enhanced. In Western Australia at least, it is probable that an upper limit to
the Permian System can be established on a palynological basis, and that floral changes
of great magnitude were initiated at the beginning of the Triassic Period. These changes
involved the extinction of the major elements of the Glossopteris-¥\ox2i and their replace-
ment by other plant groups, some of which possibly originated in the northern hemisphere
during Upper Permian time.

To expand these conclusions on the fate of the GIossopteris-¥\oxsi a brief consideration
of the main characteristics of Australian Permian microfloras is a necessary preliminary.
Certain aspects of these microfloras have been discussed by Klaus (1958), Balme (1962)
and more cursorily by other authors. Wherever sediments containing the typical Glos-
sopteris-2LSSociation of plant macrofossils have been examined palynologically, they
have yielded well-defined and basically similar assemblages of spores and pollen grains.
In post-glacial Permian sediments these microfloras are characterized generally by con-
siderable diversity, and particularly by their high content of disaccate pollen grains with
striate proximal faces. In Western Australia microfloras of this type make their first
appearance in glacial deposits, which form the basal units of the Permian successions
in each of the major sedimentary basins. With modifications they persist throughout the
Artinskian and attain their maximum variety in Upper Permian times. It is uneertain
whether the Upper Permian is completely represented in any Australian state and there
is evidence of a break in sedimentation between Permian and basal Triassic sediments
in both the Canning and Perth Basins in Western Australia. Nevertheless, the hiatus
cannot be a long one, for the Hardman Member of the Liveringa Formation in the
Canning Basin contains marine fossils which enable it to be correlated with the Upper
Productus Limestone of the Salt Range (Dickins and Thomas 1954), in turn equivalent

36

PALAEONTOLOGY, VOLUME 6

to the Dzhulfian or uppermost Permian of Russia. If a non-sequence exists in the Upper
Permian of the Canning Basin it can, therefore, represent only part of the Dzhulfian
Stage.

Typically Permian microfloras occur in the Liveringa Formation wherever it has been
examined, and these compare closely with assemblages from the Indarra Beds in the
Perth Basin, the Newcastle and Illawarra Coal Measures in New South Wales, the
Upper Bowen Group and its equivalents in Queensland and the Cygnet Coal Measures in
Tasmania. In other words the palynological evidence strongly implies the existence of
a pan-Australian flora in Upper Permian times. This, in classic terms, was the Glossop-
tcra-Flora.

Basal Triassic microfloras from Western Australia, represented by assemblages from
the Kockatea Shale and Blina Shale, are quite unlike any known from Australian Per-
mian strata. A few Palaeozoic types fe.g. Platysaccus cf. papiliouis) occur rarely in the
Mesozoic, but the most prominent Scythian forms, such as Limdbladispora and Taeni-
aesporites, have never been observed in Australian Permian sediments. Only one con-
clusion seems possible from the microfloral evidence, namely that, in Western Australia,
Lower Triassic floras were quite distinct from those of the Upper Permian. From this it
follows that the Glossopteris-Flora. did not survive into the Mesozoic, at least in the
western part of the present Australian continent.

Virtually nothing is known of the megascopic organs of these Lower Triassic plants
which superseded the Glossopteris-Flora. No identifiable plant megafossils have so far
been recorded from the Kockatea Shale, but plants occur rarely in the upper part of the
Blina Shale at Erskine Range in the Canning Basin. These have not been studied in
detail, but include equisetalean, and probably lycopodiaceous, stem fragments. Dicroi-
dium, the frond genus which particularly characterizes Triassic continental sediments
in eastern Australia, has been reported from the Erskine Sandstone, a continental
Triassic unit conformably overlying the Blina Shale (Antevs 1913, Brunnschweiler 1954)
but is not known to occur in the Perth Basin. On palynological grounds it is unlikely
that the pteridospermous group which bore Dicroidium-foliage was an important com-
ponent of the Scythian floras in Western Australia. Neither the Kockatea Shale nor the
Blina Shale contains the pollen genus PteruchipoUenites (in the sense of Couper 1958),
which is always abundant in association with Dicroidiiim in the Triassic of New South
Wales, Queensland, and South Australia. PteruchipoUenites is present in the Erskine
Sandstone, and in the Perth Basin its first known occurrence is in continental Triassic
sediments which overlie the Kockatea Shale in BMR 10 (Beagle Ridge) Bore. If, as is
implied here, PteruchipoUenites can be used as an index for the Dicroidium-FXora, it
follows that this did not become established in Western Australia until after early
Scythian time. Its first appearance cannot be dated unequivocably, but the sediments in
which PteruchipoUenites occurs in Beagle Ridge Bore conformably overlie the Kockatea
Shale, with an apparently transitional contact, and are likely to be late Scythian or early
Middle Triassic. From the preceding discussion it is concluded that plant microfossils
from the Kockatea Shale record the existence of a specialized and fairly short-lived flora
which became established in the late Permian or early Triassic and was replaced by the
Dicroidium-Flora, probably before the end of the Scythian Age.

Whether this flora existed in eastern Australia is conjectural, in view of the inadequacy
of palynological data and the difficulties of correlating Australian Triassic sediments. On

B. E. BALME; TRIASSIC PLANT MICROFOSSILS

37

a priori grounds microfloras similar to those from the Kockatea Shale would be expeeted
to occur in the lower part of the Narrabeen Group in New South Wales. However, from
the scanty published evidence, and from the few samples examined by the present author,
this is not so. Taeniaesporites was present in a shale collected from the base of the Nar-
rabeen Group in the Nattai River district of New South Wales, but in other ways the
microflora from this sample resembled those from the Erskine Sandstone rather than
the Kockatea Shale. Hennelly (1958) described a rather poorly preserved microflora
obtained from the bottom 75 feet of the Narrabeen Group in the Illawarra District of
New South Wales. In this, disaccate pollen grains of the PteruchipoI/enites-typQ (Pityo-
sporites reticulatus Hennelly) were apparently dominant and in general composition the
assemblages look younger than those from the Kockatea Shale. Two hypotheses may be
advanced to explain the differences between microfloras from the base of the Triassic
System in New South Wales and Western Australia. It is possible that a distinct floral
province existed in the western part of the Australian continent during Scythian times
and that elements of the Dicroiciiwn-Flora. migrated from eastern Australia late in the
Lower Triassic. Alternatively, part of the Lower Triassic may be missing in the Sydney
Basin and the conformable contact of the Narrabeen Group with the underlying Illa-
warra Coal Measures may obscure a sedimentary hiatus representing at least part of
the Scythian Stage. Further speculation is unjustified on the published data available,
but a convincing answer should follow the assessment of information on the Triassic
of eastern Australia at present being accumulated by palynologists of the Bureau of
Mineral Resources, Canberra.

The most obviously interesting record from the Kockatea Shale is that of the disac-
cate pollen genus Taeniaesporites, which had not previously been reported from the
southern hemisphere. Taeniaesporites belongs to the diverse complex of disaccate pollen
grains with transverse thickenings on their proximal faces, which have been assigned
by Potonie to the Infraturma Striatiti. However, its structural characters set it apart
from other genera of the Striatiti, and are so distinctive that it can hardly be doubted
that at least the haploxinoid species of Taeniaesporites were derived from closely related
plants. Such species have been reported from widely separated regions in the northern
hemisphere, occurring in Upper Permian and Triassic strata.

In Australia Taeniaesporites has never been found in Permian sediments and appears
to be confined to the Triassic. Rare specimens have been found in the upper part of the
Narrabeen Group and in coals from the Springfield Basin, South Australia. These South
Australian coals almost certainly correlate with the Leigh Creek Coal Measures and may
be of Upper Triassic or Rhaetic age (Dettmann 1961). Although Taeniaesporites prob-
ably ranges into the Australian Upper Triassic, it appears to be particularly charac-
teristic of Scythian strata, for it is only known in abundance from the Kockatea Shale
and Blina Shale. Records from the northern hemisphere indicate that Taeniaesporites
has a different stratigraphic distribution in western Europe. One of its oldest documented
occurrences is in the Hilton Plant Beds in Westmorland (Jansonius 1962) and it is
known also from the German Zechstein (Potonie and Klaus 1954, Leschik 1956). Little
published information is available on the palynology of the Buntsandstein or Muschel-
kalk equivalents, but Taeniaesporites (Lueckisporites) kraeuseli was listed by Klaus
from the Carnian of Austria, and similar species are known from the Keuper (Leschik
1955, Pautsch 1958). Post-Keuper records are dubious, but the species Protosacculina

38

PALAEONTOLOGY, VOLUME 6

glabrescens Malyavkina from the Rhaetic of Kazakhstan may belong to Taeniaesporites,
and a single poorly preserved specimen of the genus from the Swedish Liassic was illus-
trated by Nilsson (1958).

On the available evidence Taeniaesporites appeared earlier, and persisted longer, as
a prominent microfloral element in Europe, than it did in Australia. Perhaps the appar-
ent differences in the time of first appearance should not be stressed, in view of the pos-
sibility that strata homotaxial with the Upper Zechstein are not represented in Australia.
The abundance of Taeniaesporites throughout the European Triassic, on the other hand,
contrasts with its virtual restriction to the Lower Triassic in Australia and these dis-
crepancies in distribution call for some explanation. One interpretation of the evidence
is that the parent plants of Taeniaesporites were adapted to the more or less desertic
environments which characterized the Triassic Period in western Europe. The occurrence
of Pleuromeia, and other Triassic genera, in the Erskine Sandstone (Brunnschweiler
1954) suggests that desertic plant communities of the European type were stabilized in
Western Australia during the early Triassic. These did not survive the extensive climatic
changes which were initiated probably in the late Scythian, leading to the establishment
of the Dicroidium-¥\om and the formation of thick coal measures in many parts of the
Australian continent.

The only detailed published palynological study of Lower Triassic strata from the
northern hemisphere was carried out by Jansonius on sediments from the Scythian sec-
tion of the Toad-Grayling Formation in Alberta, Canada. Assemblages from the Toad-
Grayling Formation are more diverse than those from the Kockatea Shale and contain
higher proportions of disaccate pollens, but clear similarities exist between the Western
Australian and Canadian microfloras. Taeniaesporites is common in both and the Alber-
tan assemblages also contain KraeuseJisporites, Limdbladispora (cf. Aculeisporites vari-
abilis Jansonius), Vitreisporites, Striatites, Crustaesporites, and Platysaccus. Close
resemblances between the microplankton suites have already been noted by Jansonius,
and it seems reasonable to infer that the Australian and Canadian microfloras originated
from basically similar communities of parent plants.

Disaccate pollen grains of the Taeniaesporites-type are known from the Lower Triassic
of Madagascar (de Jekhowsky, personal communication), and the wide distribution of
these forms in basal Mesozoic sediments suggests that the postulated late Permian
phytogeographic changes were not confined to Western Australia. There is an urgent
need for further information on early Mesozoic microfloras, particularly from India and
the countries of the southern hemisphere, but the evidence at present available draws
attention to the possibility of establishing a palynological basis for the inter-continental
correlation of Lower Triassic strata.

REFERENCES

ANTEVS, E. 1913. The results of Dr. Mojberg’s Swedish scientific expedition to Australia. V. Some

Mesozoic plants. K. svenska Veteiisk Akad. Hand. 52, 1-6.

ARKELL, w. J. and PLAYFORD, p. E., 1954. The Bajocian ammonites of Western Australia. Phd. Trans.

Roy. Soc. Loud., Ser. B, 237, 547-605.

BALME, B. E. 1962. Palynological evidence bearing on the development of the Glossopteris-^ora.

Evolution of Living Organisms (Melbourne University Press), 269-80.

• and HASSELL, c. w. 1962. Upper Devonian spores from the Canning Basin, Western Australia.

Micropaleontology, 8, 1-28.

B. E. BALME; TRIASSIC PLANT MICROFOSSILS 39

BALME, B. E. and HENNELLY, J. p. F. 1955. Bisaccate sporomorphs from Australian Permian coals.
Aust. J. Bot. 3, 89-98.

BHARADWAJ, D. c. 1 958. On Porostrobiis zeilleri Nathorst and its spores with remarks on the systematic
position of P. beruholdi Bode and the phylogeny of Densosporites Berry. Palaeobotaiiist, 7, 67-75.
BRUNNSCHWEiLER, R. o. 1954. Mcsozoic Stratigraphy and history of the Canning Desert and Fitzroy
Valley, Western Australia. J. Geol. Soc. Aust. 1, 35-54.

CHURCHILL, D. M. 1960. Living and fossil unicellular algae and aplanospores. Nature, Loud. 186,
493-4.

COOKSON, I. c. and dettmann, m. e. 1959. On Schizosporis, a new form genus from Australian Cre-
taceous deposits. Micropaleontology, 5, 213-16.

couper, r. a. 1958. British Mesozoic microspores and pollen grains. Palaeontographica, 103b, 75-179.
DETTMANN, M. E. 1961. Lowcr Mcsozoic megasporcs from Tasmania and South Australia. Micro-
paleontology, 7, 71-86.

DiCKiNS, J. M., M^TAViSH, R. A. and BALME, B. E. 1961. The Beagle Ridge Bore. Aust. Oil and Gas J.
7, 20-21.

DUNBAR, c. o. and RODGERS, J. 1958. Principles of Stratigraphy. New York. John Wiley.

DUNHAM, K. c. 1953. Red coloration in desert formations of Permian and Triassic age in Britain.

Congr. Geol. Intern., Comptes rendus, 19^ session, Alger 1952. Sect. VII, fasc. VII, 25-29.
GLENiSTER, B. F. and FURNISH, w. M. 1961. The Permian ammonoids of Australia. J. Paleont. 35,
673-736.

GREBE, H. 1957. Zur Mikroflora des niederrheinischen Zechsteins. Geol. Jb. 73, 51-74.

HARRIS, w. F. 1955. A manual of the spores of New Zealand Pteridophyta. N.Z. Dept. sci. and ind.
Res. Bull. 116, 1-186.

HENNELLY, J. p. F. 1958. Sporcs and pollens from a Permian-Triassic transition, N.S.W. Proc. Linn.
Soc. N.S. W., 83, 363-9.

JANSONius, J. 1962. Palynology of Permian and Triassic sediments. Peace River area, western Canada.
Palaeontographica, 110b, 35-98.

DE JEKHOWSKY, B. 1961 . Sur quclqucs hystrichospheres permo-triasiques d’Europe et d’Afrique. Rev.
micropaleontologie, 3, 207-12.

DE JERSEY, N. J. 1949. Principal microspores in the Ipswich coals. Univ. QlcL, Dept. Geol. Papers, 3
(9), 1-12.

KLAUS, w. 1955. liber die Sporendiagnose des deutschen Zechsteinsalzes und des alpinen Salzge-
birges. Z. deutsch. geol. Ges. 105, 776-88.

1958. Some lower mesophytic microspores of Europe with remarks on their relation to the

Gondwana-microflora. J. Palaeont. Soc. India, 3, 151-5.

■ 1960. Sporen der karnischen Stufe der ostalpinen Trias. Jb. Geol. B.A. 5, 107-83.

KNOX, E. M. 1950. The spores of Lycopodium, Phylloglossum, Selaginella and Isoetes and their value
in the study of microfossils of Palaeozoic age. Trans, bot. Soc. Edinb. 35, 207-57.

LESCHiK, G. 1955. Die Keuperflora von Neuewelt bei Basel. II. Die Iso- und Mikrosporen. Schweiz,
palaont. Abh. 72, 1-70.

1956. Sporen aus dem Salzton des Zechsteins von Neuhof bei Fulda. Palaeontographica, 100b,

122-42.

LUNDBLAD, B. 1948. A sclaginelloid strobilus from east Greenland (Triassic). Medd. dansk. geol.
Foren. 11, 351-63.

1950. On a fossil Selaginella from the Rhaetic of Hy Hinge, Scania. Svensk. bot. Tidskr. 44, All-

86.

mYavish, r. a. 1961. Completion report on Bore BMR 10, Beagle Ridge, Western Australia. Bur.
Min. Resourc. Aust. Rec. 1960/31.

M^WHAE, J. R. H., PLAYFORD, P. E., LINDNER, A. W., GLENISTER, B. F. and BALME, B. E. 1958. The Strati-
graphy of Western Australia. J. Geol. Soc. Aust. 4, 1-161.

MALYAVKiNA, V. s. 1953. Upper Triassic, Lower Jurassic and Middle Jurassic spore-pollen assemblages
from the eastern and western Pre-Urals. Leningrad and Moscow V.N.I.G.R.I. Trud., n.s., 75, 93-147.
[In Russian.]

MULLER, J. 1959. Palynology of Recent Orinoco delta and shelf sediments: Reports of the Orinoco
Shelf Expedition; Volume 5. Micropaleontology, 5, 1-32.

40 PALAEONTOLOGY, VOLUME 6

NAUMOVA, s. N. 1950. PolIcn of angiosperm type from Lower Carboniferous deposits. Izv. Akad.

naiik. SSSR, ser. geol. 3 (1950), 103-13. [In Russian.]

NILSSON, T. 1958. Uber das Vorkommen eines mesozoischen Sapropelgesteins in Schonen. Lunds
Univ. Arssk. F.N. Ard. 2. 54 (10), 1-111.

PAUTSCH, M. 1958. Keuper sporomorphs from Swierczyna, Poland. Micropaleontology, 4, 321-5.
POTONiE, R. 1958. Synopsis der Gattungen der Sporae dispersae. II. Teil: Sporites (Nachtrage),
Saccites, Aletes, Praecolpates, Polyplicates, Monocolpates. Beili. Geol. Jb. 31, 1-114.

1960. Synopsis der Gattungen der Sporae dispersae. III. Teil. Ibid., 39, 1-189.

and KLAUS, w. 1954. Einige Sporengattungen des alpinen Salzgebirges. Geo/. 7/5. 68, 517-44.

■ and KREMP, G. 1954. Die Gattungen der palaozoischen Sporae dispersae und ihre Stratigraphie.

Ibid. 69, 111-94.

1955. Die Sporae dispersae des Ruhrkarbons. I. Teil. Palaeontographica,9^^, \-\36.

ROGALSKA, M. 1956. Spore and pollen analysis of the Liassic deposits of the Mroczkow-Rozwady
area in the Opoczno District. Warsaw Inst. Geol. Bulk 104, 1-89.

STAPLiN, F. L. 1960. Upper Mississippian plant spores from the Golata Formation, Alberta, Canada.
Palaeontographica, 107b, 1-40.

1961. Reef-controlled distribution of Devonian microplankton in Alberta. Palaeontology, 4,

392-424.

TAYLOR, G. H. 1953. The spore Content of the Leigh Creek coal. S. Aiist. Dept, of Mines Mining Rev.
99, 155-70.

TETERiUK, V. K. 1958. On the discovery of open-pored pollen grains of Palaeozoic angiosperms. DokL
Akad. nauk. SSSR 118, 1034-5. [In Russian.]

THOMAS, G. A. and DiCKiNS, J. M. 1954. Correlation and age of the marine Permian formations of
Western Australia. Aust. J. Sci. 16, 219-23.

B. E. BALME

Department of Geology,
University of Western Australia,

Manuscript received 20 March 1962 Nedlands

THE MORPHOLOGY AND DEVELOPMENT OF
SPECIES OF MARSSONELLA AND
PSEUDOTEXTULARIELLA FROM THE
CHALK OF ENGLAND

by TOM BARNARD

Abstract. The morphology and stratigraphical importance of species of Marssoiiella and Pseudote.xtiilariella
from the Chalk of England are discussed. The type specimens of Pseudotextidariella preserved in the Cushman
collection have been re-examined. This genus from the Cenomanian has been re-studied particularly in relation
to the internal structures, which are re-described. The relationships of several conical arenaceous Foraminifera
are discussed.

In 1953 the author {in Barnard and Banner 1953) deseribed species of Marssonella and
Pseudotextulariella from parts of the Chalk succession in England. These species were
not dealt with in any detail, but formed part of a general paper on faunas from the Chalk,
no emphasis being laid on their relative stratigraphical importance. Abundant new
material has become available to the author and a more complete account of the mor-
phology, development, and occurrence of these two genera may now be given. Much
new evidence has been found concerning the detailed structure of Pseudotextulariella
cretosa (Cushman), and its relationships and derivations may be discussed.

The present work was carried out during the tenure of a Leverhulme Fellowship,
which enabled the author to visit the U.S.A. and study Cushman’s specimens in the
Smithsonian Institution, Washington. The author is grateful to Dr. C. A. Cooper for
access to these specimens. The author wishes to acknowledge a grant from the Central
Research Fund, University of Fondon, which enabled him to collect additional material
from a number of localities in southern and eastern England.

Specimen numbers with the prefix P indicate specimens in the British Museum (Natural History)
collections.

SYSTEMATIC DESCRIPTIONS
Genus marssonella Cushman 1933

Genotype. Textukiria trochus d’Orbigny.

Gaudryina oxycona Reuss 1860 (designated by Cushman 1933u, p. 36; 1933/t, p. 121)
is a synonym of Textularia trochus d’Orbigny 1840, p. 45, pi. 4, figs. 25, 26, which pre-
dates Gaudryina oxycona Reuss 1860 and was redesignated genotype by Barnard and
Banner 1953, p. 204.

Marssonella ozawai Cushman
Text-figs. \a-c

1936 Marssonella ozawai Cushman, p. 43, pi. 4, figs. I0<7, b.

1937 Marssonella ozawai Cushman, p. 59, pi. 6, fig. 18.

1953 Marssonella ozawai Cushman; Barnard (in Barnard and Banner), p. 205, pi. 19, figs, la, b.
[Palaeontology, Vol. 6, Part 1, 1963, pp. 41-54, pi. 7.]

42

PALAEONTOLOGY, VOLUME 6

Description. Conical test, almost circular in cross-section. The test is coarsely arena-
ceous, composed of a mosaic of almost equal-sized quartz grains, with little calcareous
cement. These grains protrude through the cement so that the surface of the test is
rugose. A globular proloculum is followed by one to three whorls of three to five small
crescentic or reniform chambers. The test then becomes biserial, the height of the cham-
bers increases slowly, but the diameter increases rapidly, so that initially the edges of
the test diverge up to an angle of 60°. After this stage, which sometimes continues up
to about half the height of the test, the diameter of the test may remain almost constant
giving a cylindrical late portion. Rarely the diameter increases rapidly giving a pseudo-
flange to the later chambers, or it may increase slowly producing a large conical test.

TEXT-FIG. 1. Marssoiiella ozawai Cushman. All specimens from
the Cenomanian, a, b. From the Portsdown borehole, 1055-1060
feet, P45013. c, From the marls 4 feet above ‘Chloride Mari’,

Arlsey, Bedfordshire, P45014. All specimens X90.

The septa are almost flat in some varieties, but often become slightly constricted and
indented, sometimes near the apertural end or occasionally the whole of the biserial
portion. The apertural faces vary from almost flat, in the commonest variety, to either
slightly convex or, in rare specimens, concave. Generally the apertural faces are reni-
form in shape. The basal aperture varies from slit-like to reniform in shape. The test
is usually straight, but occasionally may be slightly arcuate or even arranged in several
stages. At certain levels, long slender almost cylindrical variants occur.

Localities. Numerous.

Horizon. Schloenbachia varians zone.

Dimensions and hypotypes. P45013 (text-figs. \a, b). Height 0-99 mm., diameter 0-49 mm. P45014 (text-
fig. Ic). Height 1-33 mm., diameter 0-57 mm.

Marssonella turris (d’Orbigny)

Text-figs. 2a-h

1840 Textidaria turris d’Orbigny, p. 46, pi. 4, figs. 27, 28.

1890 Textidaria turris d’Orbigny; Burrows, Sherborn, and Bailey, p. 553, pi. 8, fig. 15.

1928 Textidaria turris d’Orbigny; Franke, p. 131, pi. 12, fig. 3.

1937 Textidaria turris d’Orbigny; Cushman, p. 58, pi. 5, fig. 28; pi. 6, figs. 1, 2, (5?).

1957 Marssonella turris (d’Orbigny); Hofker, p. 83, text-figs. 84a-c, 85a-e.

TOM BARNARD; MARSSONELLA AND P SEU DOTEXTU LARIELLA

43

Description. The test is generally a long narrow cone, which can vary to almost cylin-
drical, or to a cone with an apical angle of about 60°. The size of the test varies consider-
ably, and also the rugosity of the wall. Usually the test is composed of a mosaic of small
equi-dimensional quartz grains set in a moderate amount of calcite cement. Occasionally
at certain horizons large quartz grains are cemented with little calcite. The quartz grains
are usually almost flush with the surface of the test, but when large grains are present
they are not deeply set in the cement, but protrude so that the surface is rough. Often in
forms with the coarser test, grains of minerals other than quartz are included, such as
glauconite, ilmenite, &c.

TEXT-FIG. 2. Marssoiiella tunis (d’Orbigny). a-e, Showing variation in shape of the test, from the
Cenomanian, Schloeubachia varians zone, from Sundon, near Dunstable, Bedfordshire, P45016, x90.

/-/;, Showing the variation in the shape of the apertural face and the position of the aperture.

Variations occur in the shape of the test. Occasionally there are breaks in the normal
growth-rate which give the appearance of rejuvenation. Later chambers are often ill-
formed or damaged, growth is irregular, and the tests appear to be flanged. Sometimes
the tests are slightly arcuate. A globular or hemispherical proloculum is followed by
two or three whorls of three to five crescentic chambers which show little or no increase
in height. This stage in the growth of the test is difficult to interpret. A biserial stage is
soon reached and continues throughout the test. The sutures are straight and scarcely
visible except in decorticated specimens, or in the end-ehambers of the tests, where they
may be constricted, ribbed, or flush with the surface of the test. The apertural face varies
from crescentic to reniform in shape, and from highly convex to flat. Occasionally
furrows run down the apertural face towards the aperture. The aperture is either slit-
like, crescentic, or reniform.

Localities. Numerous.

Horizon. Cenomanian-Lower Senonian.

Dimensions and hypotypes. P45016 (text-figs. 2a-e). a. Height 0 46 mm., diameter 0-38 mm. b, Height
0-49 mm., diameter 0-38 mm. c. Height 0-87 mm., diameter 0-46 mm. d. Height 0-68 mm., diameter
0-42 mm. e. Height 0-46 mm., diameter 0-42 mm.

44

PALAEONTOLOGY, VOLUME 6
Marssonella trochus (d’Orbigny)

Text-figs. l>a-q

1840 Textularia trochus d'Orbigny, p. 45, pi. 4, figs. 25, 26.

1854 Textularia comilus Reuss, p. 72, pi. 26, figs, la, b.

1860 Gaudryina oxycoua Reuss, p. 229, pi. 12, fig. 3.

1891 Textilaria cf. conulus Reuss; Beissel, p. 68, pi. 13, figs. 23-29.

1899 Gaudryina oxycoua Reuss; Egger, p. 38, pi. 4, figs. 1-3.

1899 Haplophragmium petiolus Egger, p. 143, pi. 2, figs. 37-39.

1925 Gaudryina oxycoua Reuss; Franke, p. 15, pi. 1, figs. 20a, b.

1928 Gaudryina oxycoua Reuss; Franke, p. 143, pi. 13, figs. 8u, b.

1928 Textularia trochus d’Orbigny f. siibconica Franke, p. 131, pi. 12, fig. 1.

1928 Textularia trochus d’Orbigny f. typica Franke, p. 131, pi. 12, fig. 3.

1937 Marssonella oxycoua (Reuss); Cushman, p. 56, pi. 5, figs. 27-29; pi. 6, figs. 1-17.

1946 Marssonella cf. oxycoua (Reuss); Schijfsma, p. 38, pi. 1, figs. 12a, b.

1953 Marssonella trochus (d’Orbigny); Barnard (in Barnard and Banner), p. 204, text-figs. 5o-s.
1957 Marssonella oxycoua (Reuss); Hofker, p. 85, text-figs. 86-90.

Description. The conical test varies slightly in shape. During the initial growth-stages the
early part of the test is conical with walls diverging at a wide angle as the diameter
increases. This is often followed by a period when the diameter does not increase, and
a cylindrical growth-stage is produced. Sometimes there is an abrupt increase in the
diameter of the test, so that rapid expansion of the end-chambers produces a ‘flanged’
test. Often there is an abrupt halt in growth, which is then resumed, following a similar
pattern as before; this gives the appearance of rejuvenation.

The globular proloculum is followed by whorls of five, four, and three chambers,
before the biserial stage is reached. In conical forms the biserial crescentic chambers are
overlapped by succeeding chambers.

The septa are usually flat, but can be concave, or convex, often with a slight depression
towards the aperture. The face of the end-chamber varies in shape from semicircular,
crescentic, to reniform. The aperture varies from its usual position, basal and flush with
the surface of the test, to a vertical apertural face. Occasionally this is excavated and
the aperture is almost hidden. The aperture is usually slit-like, but can be hemi-elliptical
or reniform. The test is smooth with slight constrictions at the septa. The wall is com-
posed of fine quartz grains set in a calcareous cement. Sometimes isolated specimens
have up to five radial partitions developed in the last chambers only.

Locality of figured specimens. Whitlingham, near Norwich, Norfolk.

Horizon. Upper Chalk. Belemnitella mucronata zone.

Dimensions and hypotypes. P45015 (text-fig. 2a). Height 0-95 mm., diameter 0-84 mm. P45015 (text-fig.
2b). Height 0-95 mm., diameter 0-72 mm.

Marssonella conoidea (Marie)

Text-figs. Aa-c

1941 Textularia conoidea Marie, p. 63, pi. 2, fig. 20.

Description. The test is a regular blunt cone, which does not show a great deal of variation
in shape. The initial stages are often a curved-sided cone; later the sides of the test
become parallel so that it appears bullet-shaped. The cross-section is almost circular.
A globular proloculum is followed by whorls of five to three chambers, but rapidly the

TOM BARNARD: MARSSONELLA AND PSEUDOTEXTULARIELLA

45

TEXT-FIG. 3. Marssonella trochus (d’Orbigny). a, b, X 90, from the Upper Chalk, Belemnitella miicro-
tiata zone, from Whitlingham, near Norwich, Norfolk, P45015. c, d. Showing the triserial and biserial
initial stages of the test, e, A specimen showing the simple interior to the chambers with no radial
partitions./. Showing remnants of radial partitions in the interior of the shell, g, Showing chambers with
normal simple interiors, h-k. Outline lucida drawings showing variation in the shape of the cones.
I-n, Showing the variation in the shape of the apertural face, o-cj. Showing the position of the aperture

and the slope of the apertural face.

test becomes biserial. This initial part of the test is extremely difficult to interpret due to
the rough surface. The wall of the test is rugose. In the initial part, coarse grains of quartz
and sponge spicules are set in a thick calcite cement. However, in most of the test the
coarse grains protrude above the surface of the cement. Characteristic of this species is
that coarse grains appear to be arranged in a rough rib-like pattern, one rib above each
suture. The sutures are in deep constrictions, and have flat, concave, or convex surfaces.
Often the slit-like aperture is deeply set, almost covered by a lip-like extension of the
chamber. In these forms there is a tendency for the chambers of one side of the test to
develop, whereas those on the other decrease in size so that the test becomes almost

46

PALAEONTOLOGY, VOLUME 6

uniserial, especially in forms where the aperture becomes central and deep, appearing to
serve both sets of chambers.

Locality. H. Attock’s Pit, New Catton, Norwich, Norfolk.

Horizon. Belemnitella miicronata zone.

Dimensions. Height 1T4 mm., diameter 0-76 mm.

Hypotype. P45017.

The development and stratigraphical oeeurrenee o/Marssonella. In 1953 the author drew
attention {in Barnard and Banner 1953, p. 204) to the extreme variation which occurs

in the size and shape of the test, as well as the
shape of the final chamber and aperture. The
species of Marssonella from the Chalk were
thought (ibid.) to be one group; however, since
then numerous specimens have been obtained
from widely scattered localities as well as con-
tinuous sections, and it has become increasingly
apparent that the group consists of a few species
which prove to be of stratigraphical value.

At some horizons abnormally large forms are
produced, at others the general character of the
wall changes, so that an abundance of coarsely
rugose forms is found. Text-fig. 5 shows in dia-
grammatic form the general history of the species
of Marssonella and Pseudotextidariella (another
conical form, although probably unrelated to
Marssonella, is often confused with it) through-
out the Chalk. Almost confined to the Ceno-
manian is M. ozawai Cushman, a large bullet-
shaped species with a rugose wall. This species is
easily separated from its contemporaneous, but longer-ranged form M. turn's d’Orbigny.
The latter is the root-stock of the species of Marssonella from the Chalk.

Marssonella turris d’Orbigny is usually an acute-angled cone, with a smooth wall.
However, at certain horizons there is a tendency for the test to become rugose. Forms
with coarser grains in the test generally occur in the Cenomanian, where the coarser silt-
grade quartz is found in greater profusion than in higher levels in the Chalk. Generally
there is a steady increase in the amount of calcareous cement as the species ranges into
higher zones in the Chalk.

However, even here more rugose forms occur at certain levels, which can often be
correlated with the size of the quartz grains in the Chalk. Although the test varies con-
siderably in shape, two main trends occur. At some levels in the higher zones of its
range there is a tendency for long, narrow, almost cylindrical variants to occur abun-
dantly at sporadic horizons. Also at about the same time (Micraster cor-testudinarium
zone), broader, blunter cones develop, which may be the forerunners of Marssonella
trochus d’Orbigny and its synonymous form M. oxycona Reuss. In M. troehus d’Orbi-
gny there is considerable variation in the shape of the test, as shown in text-figs. 3h-k
and 3l-q. There is, however, a general tendency for the test to increase in size until the

TEXT-FIG. 4. Marssonella conoidea (Marie).
a, X 90, from the Upper Chalk, Belemnitella
miicronata zone, H. Attock’s Pit, New Cat-
ton, Norwich, Norfolk, P45017. b, c. Top
view showing the sunken aperture; b, lipped
and basal; c, circular and central.

TOM BARNARD; MARSSONELLA AND PS UEDOTEXTULARIELLA

47

TEXT-FIG. 5. Diagram showing the ranges and development of some conical foraminifera
from the Chalk (Upper Cretaceous).

Be/emnitella mucronala zone, where large specimens occur, several times the volume of
earlier ones. Often in these forms the end-chambers increase abruptly in diameter so that
the test appears to be flanged.

At some horizons isolated specimens have irregular radial partitions developed in the
end-chambers only. This is rare, but gives a superficial resemblance to Pseudotextiilariella

48

PALAEONTOLOGY, VOLUME 6

cretosa Cushman, and its Turonian derivatives. However the absence of chamberlets in
the later chambers and radial partitions in the early part of the test, separates the two
forms.

Marssonella conoidea ( Marie) occurs in the Actinocamax quadratus and Belemnitella
mucronata zones, and although it resembles M. ozawai Cushman in shape, it is the only
species of Marssonella from the Chalk which has a rib-like ornament composed of coarse
quartz grains arranged in rough lines above the sutures. Also it has a much more regular
test than other species.

Summarizing the history of the species of Marssonella from the Chalk, it may be stated
that one main stock (M. Unris d’Orbigny) occurs, giving rise, at later horizons, to blunt
conical forms, which, when sorted by natural selection, produce such species as M.
trochus d’Orbigny, and its large, often flanged variety, M. oxycona Reuss.

Genus pseudotextulariella Barnard 1953

Genotype. Textulariella cretosa Cushman.

In Barnard and Banner 1953, p. 198, the author proposed the genus Pseudotextidari-
ella with the holotype Textulariella cretosa Cushman, specimens of which had been
obtained by Cushman from the Cenomanian, at Barrington near Cambridge. Unfor-
tunately, although the manuscript was sent in during 1952, it was not published until
1953. Since then Thalmann (1955, p. 53) has established that under Art. 25 of the Inter-
national Rules of Zoological Nomenclature, Pseudotextulariella is legitimate.

The author has now had the opportunity of studying Cushman’s specimens in the
National Museum, Washington, and resulting from this work, certain points can be
added to enable a more accurate definition of the genus. More information has also been
obtained based on the study of the internal structures.

Pseudotextulariella cretosa (Cushman)

Plate 7, figs. 1-6, 8; text-figs. 6a-d, la-f, 8o-c

1932 Textulariella cretosa Cushman, p. 97, pi. 11, figs. 17-19.

1937 Textulariella cretosa Cushman, p. 61, pi. 6, figs., 26-28.

1948 Textulariella cretosa Cushman; Williams-Mitchell, p. 97, pi. 8, fig. 1.

1953 Pseudotextulariella cretosa (Cushman); Barnard (in Barnard and Banner), p. 198, text-
figs. 6b-i.

Description. Test conical, broad or narrow, usually straight, but occasionally slightly
arcuate, particularly at the initial end. Initial whorl triserial, following a globular or sub-
globular proloculum. A biserial arrangement of the chambers is soon established and

EXPLANATION OF PLATE 7

Figs. 1-6, 8. Pseudotextulariella cretosa (Cushman). All specimens from the Cenomanian, Barrington,
near Cambridge. Fragments of tests to show the distribution and development of the radial and
horizontal partitions. 1, x40. 2-6, x45. 8, X 60.

Fig. 7. Textulariella pacifica Cushman. X 60. Recent, from the H.M.S. Challenger Expedition, St. 33,
435 fathoms, Bermuda, to show the development of the radial partitions and the central cavity. The
partitions do not reach the centre of the test as in Pseudotextulariella cretosa (Cushman). BM No.
1962.3.16 1-2.

Palaeontology, Vol. 6

PLATE 7

X 40

X 45

X 45

X 45

X 45

X 60

X 45

X 60

BARNARD, Chalk Foraminifera

TOM BARNARD; MARSSONELLA AND PSEUDOTEXTULARIELLA 49

remains for the rest of the test. The initial chambers are subglobular, but those in the
biserial arrangement become almost semicircular to reniform in transverse section. The
height of these chambers varies considerably so that the chambers may have almost
parallel faces or sometimes they converge rapidly to become acutely pointed at the
medial suture, where the chambers alternate.

The height of the chambers varies from an
eighth to a quarter of the diameter, and the
sides of the test diverge at a constant angle.

The walls separating the chambers are often
marked at the surface of the test by strong
raised ribs; however, on other specimens
the sutures are either flush with the surface
of the test or even slightly constricted.

In all the numerous specimens examined,
the chambers, after the initial stage of the
test had been passed through, are divided
by an almost regular series of chamberlets.

They occur arranged through the biserial
portion of the test, and often become multi-
plied so that two to five tiers of chamber-
lets are developed. The chamberlets are
developed progressively, and usually regu-
larly, appearing at the surface of the test in
weathered or decorticated specimens. With
the application of water or clove oil to the
surface of the test, the pattern of the chamberlets is often seen, and sectioning is not
always necessary. The aperture is generally slit-like to reniform in shape, and may be
flush with the surface of the apertural face or raised on a surface almost at right angles
to the face of the chamber. In some forms irregular depressions occur in the apertural
face from the periphery to the aperture.

Specimens. U.S. National Museum, Washington. 17624 (text-fig. 6a), holotype. 25295 (text-fig. 6b)
holotype. 17625 (fext-figs. 6c, d), parafypes.

Locality. Charing, Kent, England.

Horizon. Lower Chalk, Schloenbachia varians zone.

TEXT-FIG. 6. Pseiidotextidariella cretosa (Cush-
man). All specimens in the U.S. National
Museum, Washington, from the Lower Chalk,
Schloenbachia varians zone, from Charing, Kent.
a, 17624, holotype. b, 25295, holotype. c, d, 17625
paratypes.

Development of the radial septa and ehamberlets. Although somewhat irregular and vari-
able, a progressive development of both the radial septa and tiers of chamberlets takes
place in both ontogeny and phylogeny. The phylogeny, based on variation in successive
populations occurring within the Cenomanian, is beyond the scope of the present paper.
However, the general ontogenetic development is described below, and is shown in
text-figs. 7 and 8.

Two main trends in ontogenetic development are shown in text-figs, la-f in forms
which only develop radial septa. Text-figs, la, b, c, e represent transverse sections of the
biserial part of the test. Near the initial end (a) only a few vertical radial septa are formed,
stretching from the periphery to join with a circular wall surrounding the apertural

C 1015

E

50

PALAEONTOLOGY, VOLUME 6

areas. This stage is rapidly followed by one in which the septa alternate in length,
apparently forming two generations, one group reaching the circular central wall,
whereas the other is shorter and confined to a small peripheral area.

This growth stage is shared by both main variants, but divergence now takes place

TEXT-FIG. la-f. Pseiidotextulariella cretosa (Cushman). Diagrams
to show the development of the radial partitions.

in later growth stages. In text-fig. 7c the septa again are in two main groups, a longer set
reaching the centre, and the peripheral group as in earlier growth stages. However, the
latter may do one of two things. Usually they elongate until coalescence with the central
wall results, or they become bifid or irregular, and sometimes bend, joining the longer
sutures about half-way to the centre of the chambers. Text-fig. Id shows the regular
development of a few radial vertical septa, when the smaller peripheral septa have
reached the centre. In both text-figs. Id and / the central circular wall has been omitted
for clarity. In the other trend (text-figs. 7c and /) the primary vertical septa reach the
central wall, but secondary and even shorter tertiary septa are formed and confined to
the peripheral part of the test. Often the smaller tertiary septa are arranged irregularly
and do not usually occur in the gap between every primary and secondary septa. Also
these secondary and tertiary septa are not the same height for their whole length, but
taper rapidly towards the lower chamber wall. Text-figs. 8a-c are block diagrams con-

TOM BARNARD: MARSSONELLA AND PSEUDOTEXTULARIELLA

51

structed to show development of tiers of chamberlets by the growth of not only vertical
radial septa, but several generations of horizontal septa parallel to the chamber sutures.
Text-fig. 8(3 shows a few primary radial partitions developed, with secondary alternating
partitions coalescing. Near the periphery one horizontal secondary septum is formed, con-
fined to the peripheral area and producing two tiers of almost rectangular chamberlets.

i

TEXT-FIG. Sa-c. Pseudotextiilariella cretosa (Cushman). Block
diagrams showing the development of the radial and horizontal
partitions.

A further development is shown in text-fig. 86, where further horizontal septa are
added, one above and one below the earlier secondary septum. The latter has now grown
to extend almost to the centre of the test, whereas the two later septa again are short and
confined to the peripheral zone of the test. Text-fig. 8c shows an even further develop-
ment with four secondary septa of varying length, producing five tiers of chamberlets.

THE AFFINITIES OF THE CONICAL FORAMINIFERA

The relationships and development of the conical foraminifera from the Cretaceous
to Recent are of considerable interest and importance. In spite of the numerous publica-
tions in which mention of these forms is made, little is known concerning their affinities.

52

PALAEONTOLOGY, VOLUME 6

As well as the genera dealt with in detail in earlier parts of this paper, suggestions are
made below as to the possible derivation of some other Cretaceous forms.

There are two root-stocks from which most of the genera of conical foraminifera may
have been derived, namely Marssonella and Arenobulimina. MarssoneUa consists largely
of a biserial arrangement of the chambers which rapidly follows an initial part made up
of one or two whorls, composed of three, four, or five chambers. As has already been
pointed out, at certain horizons in the Upper Senonian there is a tendency for isolated I
specimens to produce simple radial partitions dividing the chambers. It is considered I
that this tendency is repeated to some degree at least three times in the history of the
group. i

(a) In the Cenomanian Pseiidotextiilariella occurs, a genus in which complex internal |

structures are developed. Radial partitions stretch towards the centre of the test and then ]
are united in a wall surrounding a central circular hole. Later horizontal partitions and )
further generations of chamberlets are added, resulting in a genus with extremely com-
plex internal structures. h

(b) In the Upper Senonian, forms develop with a few simple radial partitions only i

occurring in the end-chambers. |

(c) A genus found in the late Tertiary to Recent is Textulariella, in which a thick ■
zone of radial partitions develops in the wall-zone, and does not reach far towards the j
centre. This character serves to distinguish this genus from Pseudotextulariella.

It is considered that these groups are somewhat similar repetitive forms which con- ’■
verge towards a common plan of development, but which do not produce identical
forms. Text-fig. 9 shows the suggested affinities of the various conical genera. ^

The initial coil in Pseudotextulariella suggests that it may have arisen from Arena- •
buliniina (Barnard, in Barnard and Banner 1953). It is more probable, however, that ,
Marssonella, Pseudotextulariella, and Arenobulimina have arisen at slightly different
stratigraphical levels from either a common ancestor or closely similar ancestral stocks. >
Also the complex internal structures and the progression of these throughout the Ceno- t
manian in Arenobulimina are closely parallel to those of Pseudotextulariella, and may f
show the close affinities between the two. It is probable that Marssonella is not the root-
stock giving rise to Pseudotextulariella. Arenobulimina occurs first in the Albian in
England and the evolution of this genus is the subject of a paper (Barnard and Banner) *
in preparation. In North America species of Arenobulimina appear to be few, and their ’
history apparently terminates in Albian times, whereas in north-west Europe the genus ■
continues and forms an important part of the Chalk faunas. t

In the Albian of England there are a few species of Arenobulimina and these are |
associated with the argillaceous environments of the Gault clay and do not develop good i
internal structures. However, as soon as the Upper Greensand and Cenomanian facies, f
with their additional limestones, are encountered the internal structures develop in the f
species.

A form superficially similar to Pseudotextulariella is found abundantly in the Walnut ■ |
Formation (Lower Albian) of Texas. This species, Dictyoconus walnutensis (Carsey), j \
possibly related to a later Upper Cretaceous genus, Colomia, has recently been studied j
by Maync (1955). i

Somewhat similar developments of chamberlets are shown in various groups of ;T
Cretaceous species belonging to the genera Dictyoconus and Coskinolina and those of ; |

( I

|- .

I

I

I

TOM BARNARD: MARSSONELLA AND PSEUDOTEXTULARIELLA

53

Pseudotextulariella. Although the distribution of these genera has not been studied in
detail, it would appear that somewhat similar internal structures develop and evolve in
almost the same pattern in different faunal provinces. Dictyoconus and Coskinolina
appear in the Texas-Mexico basins, through Florida, and in the Tethyan areas of Europe,
Spain, southern France, sometimes migrating farther north to occur in intermediate
areas such as Aquitaine. Arenobulimina and Pseudotextulariella occur in the north-west

RECENT

QC a

U 4^

Tixtulorietlo

CL —

£

3 ul

1-

L. TERTIARY

z

<

z

o

o

z

Ld

o

tn

N

TURONIAN

s

Z

<

z

C

<

b

2

b

O

z

u

u

C

■§

e

b

40

0

a:

N

c:

Cv

LU

Z

<

b

1''

N

/

/

Ar

3 _

to

7

b

common

CC

tc

o

ancestor

o

N /

s

\ /

o

y

TEXT-FIG. 9. Diagram showing the affinities of some
genera of conical foraminifera.

European basins. The ranges of the two faunas do not coincide, although there is some
overlap. Dictyoconus and Coskinolina reach their maximum development in the Albian,
Pseudotextulariella in the Cenomanian, whereas Arenobulimina is longer ranged from
Albian to Palaeocene, with several periods when development of certain characters
reached a maximum.

In parts of the Upper Cretaceous basin of California, the genus Colomia has been
recorded sporadically. This genus appears closely similar in many aspects to Dictyoconus
walnutensis (Carsey), except that the development of internal structures may be sporadic,
as with later forms of Marssonella oxycona Reuss in Europe. Certain authors (Hofker
1958, Cushman and Bermudez 1948) have recorded internal structures, whereas others
(notably Bandy 1951) have not. It is possible that Colomia may not be closely related to
Dictyoconus, as it is a calcareous genus.

54

PALAEONTOLOGY, VOLUME 6

REFERENCES

BANDY, o. L. 1951. Upper Cretaceous Foraminifera from the Carlsbad area, San Diego County,
California. J. Paleont. 25, 488-513.

BARNARD, T. and BANNER, F. T. 1953. Arenaceous Foraminifera from the Upper Cretaceous of Eng-
land. Quart. J. geol. Soc. Loud. 109, 173-216, pi. 7-9.

BEissEL, I. 1891. Die Foraminiferen der Aachener Kreide. Abh. preuss. geol. Landesanst., n.f., 3,
1-78, pi. 1-16.

BURROWS, H. w., SHERBORN, c. D. and BAILEY, G. 1890. The Foraminifera of the Red Chalk of York-
shire, Norfolk and Lincolnshire. J. R. micr. Soc. 7, 549-66, pi. 7-11.

CUSHMAN, J. A. 1932. Textidaria and related forms from the Cretaceous. Coutr. Cushman Lab. 8 (4),
86-97, pi. 11, figs. 17-19.

■ 1933u. Some new foraminiferal genera. Ibid. 9 (2), 32-38, pi. 3-4.

1933fi. Foraminifera, their classification and economic use. Spec. Publ. Cushman Lab. 4, 1-349,

pi. 1-31.

1936. New genera and species of the families Verneuilinidae and Valvulinidae and of the sub-
family Virgulininae. Ibid. 6, 1-71, pi. 1-8.

1937. A monograph of the foraminiferal family Valvulinidae. Ibid. 8, 1-210, pi. 1-24.

and BERMUDEZ, p. J. 1948. Co/um/u, a new genus from the Upper Cretaceous of Cuba. Contr.

Cushman Lab. 24, 12-13, pi. 2.

EGGER, J. G. 1899. Foraminiferen und Ostrakoden aus den Kreidemergeln der Oberbayerischen Alpen.
Abh. barer. Akad. Wiss. 21, 1-230, pi. 1-27.

FRANKE, A. 1925. Die Foraminiferen der pommerschen Kreide. Abh. geol. -palaeont. Inst. Greifswald,
6, 1-96, pi. 1-8.

1928. Die Foraminiferen der Oberen Kreide Nord- und Mitteldeutschlands. Abh. preuss. geol.

Landesanst., n.f., 3, 1-207, pi. 1-18.

HOFKER, J. 1957. Foraminiferen der Oberkreide von Nordwestdeutschland und Holland. Beih. Geol.
Jb. 27, 1-464.

1958. The taxonomic position of the genus Colomia Cushman & Bermudez, 1948. Contr. Cush-
man Fdn. 9 (2), 34—35.

MARIE, p. 1941 . Les foraminiferes de la Craie a Belemnitella mueronata du Bassin de Paris. Mem. Mus.
Hist, nat., Paris, N.S., 12, 1-296, pi. 1-37.

MAYNC, w. 1955. Coskinolina sunnilandensis n.sp., a Lower Cretaceous (Urgo-Albian) species. Contr.
Cushman Fdn. 6 (3), 105-11, pi. 16-17.

ORBiGNY, A. d'. 1840. Mcmoire sur les Foraminiferes de la Craie Blanche du Bassin de Paris. Mem.
Soe. geol. Fr. 4, 1-51, pi. 1-4.

REUss, A. E. 1854. Beitrage zur Charakteristik der Kreideschichten in den Ostalpen, besonders im
Gosanthale und am Wolfgangsee. S.B. Akad. Wiss. Wien, 7, 1-156, pi. 1-31.

1860. Die Foraminiferen der westphalischen Kreideformation. Ibid. 40, 147-238, pi. 1-13.

SCHIJFSMA, E. 1946. Foraminifera from the Hervian (Campanian) of Southern Limburg. Meded.geol.
Sticht., N.S., C (5), 7, 1-174, pi. 1-10.

THALMANN, H. E. 1955. New names for foraminiferal homonyms; II. Contr. Cushman Fdn. 6 (1), 53.
wiLLiAMS-MiTCHELL, E. 1948. The zonal value of Foraminifera in the Chalk of England. Proe. Geol.
Ass. 59 (2), 91-112, pi. 8-10.

TOM BARNARD

Department of Geology,
University College,
Gower Street, London, W.C. 1

Manuscript received 26 March 1962

WEALDEN MAMMALIAN FOSSILS

by WILLIAM A. CLEMENS

Abstract. Only two of the previous identifications of specimens considered to be teeth of Wealden mammals
can be accepted without reservation. A special collecting technique including both chemical and mechanical
processes facilitated the discovery of eight more Wealden mammalian fossils. Five were found in the Cliff End
Bone Bed, a part of the Ashdown Beds, and the remainder in the Paddockhurst Bone Bed, a part of the Grin-
stead Clay. These fossils give additional information about the morphology of the multituberculate Loxaidax
valdensis and demonstrate the presence of a symmetrodont and eupantothere in England in the early (pre- Aptian)
Cretaceous.

The fossil record of Mesozoic mammals is limited. Large collections of mammalian
fossils of late Jurassic age have been made in the Purbeck Beds of England and the
Morrison Formation of the United States. The Albian mammalian fauna is known from
fossils found in the Trinity Sand of Texas. The only record of mammals that certainly
lived in the interim between the Purbeckian and Albian consists of the rare specimens
discovered in the Wealden of England. Two mammals found in the Husin Series of
Manchuria also may be of post-Purbeckian and pre- Albian age, but the correlation of the
strata in which they were found is still open to question (see Patterson 1956, pp. 30-31).
The first specimen considered to be a mammalian fossil discovered in the Wealden of
England was described in 1891 ; in the two following decades four more specimens were
found. After this initial activity almost fifty years elapsed before another Wealden
mammalian fossil was discovered. Tn the autumn of 1960, working under the auspices
of a National Science Foundation Postdoctoral Fellowship, I began a survey of British
Mesozoic mammals and the localities at which they were found. At the suggestion of
Dr. K. A. Kermack, one of the first localities visited was Cliff End, where three of the
Wealden mammalian fossils had been found in a bone bed cropping out on the shore.
Blocks of the bone bed found scattered along the beach proved to contain mammalian
fossils. As word of my interest in Wealden mammals spread, I was told of another
promising bone bed in Paddockhurst Park that also proved to contain isolated mammalian
teeth. Some of these fossils were exhibited at a meeting of the Geological Society
(Clemens 1960). This paper is an interim report on a research project that is being con-
tinued under the direction of Dr. K. A. Kermack of the Department of Zoology, Uni-
versity College, London.

All catalogue numbers cited are from the British Museum (Natural History) Catalogue
of Fossil Mammalia.

PREVIOUS INVESTIGATIONS

In 1891 A. Smith Woodward announced the discovery of a tooth of a Wealden
mammal. Two years later Richard Lydekker described another specimen and in 1911
Woodward reported on three more. Study of these fossils and the circumstances of their
discovery has brought to light facts confirming the unpublished suggestions of several
palaeontologists that the identification of some of the fossils as teeth of Wealden

[Palaeontology, Vol. 6, Part 1, 1963, pp. 55-69.]

56

PALAEONTOLOGY, VOLUME 6

mammals is incorrect. The first fossil thought to be a Wealden mammalian tooth
(M 13 134) was found by Charles Dawson in a bone bed within the Wadhurst Clay
exposed in the ‘Old Roar Quarry’ near Hastings. It was an isolated tooth having both
the heavily worn crown and root preserved. Later it was damaged and all that remains
is a fragment of the root. Woodward (1891, p. 586) described the crown of the tooth as
being ‘ . . . supported by two roots, which are robust, of nearly equal size and depth, in-
completely separated in the upper half and terminating obtusely’. Loss of parts of the
tooth has revealed that, although deeply grooved on the surface, the root contains a
single pulp cavity rather than two as would be expected if this structure were two
incompletely separated roots.

After comparing this Wealden tooth (M13134) with haramiyid teeth and the Mg of
Ctenacodon (P/agiaiilax) minor, Woodward tentatively identified it as the molar of a
multituberculate and allocated it to the genus Plagiaidax under the provisional name
P. dawsoni. The similarities on which this identification was based are neither numerous,
detailed, nor fully convincing. Simpson (1928, p. 52) commented that the tooth ‘. . . is
probably Plagiaulacid, although its generic and specific affinities are quite undeter-
minable’. The presence of a single pulp cavity in the root reduces the probability that
the tooth is a plagiaulacid molar, which has two roots, and strongly suggests that it is
not the tooth of a mammal.

In 1893 Lydekker identified another fossil (M5691) as an incisor of a Wealden multi-
tuberculate. In the discussion following the presentation of Lydekker’s paper at the
Geological Society, Sir John Evans recounted the history of the discovery of the tooth:
‘ He [Evans] found it at Hastings, in a block of Tilgate Grit which formed part of a heap
by the side of the sea-shore, and almost immediately afterwards gave it to Prof. Prestwich,
in whose collection it was mislaid for a period of over thirty years. On again coming
across it. Prof. Prestwich placed it at the disposal of the speaker . . .’ (Lydekker 1893,
p. 283). In the course of the same discussion, Dawson stated: ‘ ... it was unfortunate that
the specimen had been taken from a loose block, because at Hastings stones foreign to
the district and miscellaneous drifted stones from the shore were frequently broken up
for road-metal. From the limited view permitted him of the specimen that evening he
was unable to identify the matrix as from the Hastings district; and he did not recognize
the fragmentary specimen before them as a portion of a mammalian tooth.’

Dawson appears to have been alone in questioning the identification of this fossil as
a mammalian tooth. Study of the specimen substantiates Lydekker’s observation that
there is no way of distinguishing it from a rodent incisor. Because it was and still is
thought to be unlikely that the Rodentia had evolved as a discrete group in the Early
Cretaceous and, because multituberculates possessed somewhat rodent-like incisors,
Lydekker and others concluded that this tooth was probably a multituberculate incisor.
Excluding the specimen in question, the available fossil record of multituberculates
demonstrates that incisors closely resembling those of rodents were evolved only in one
Tertiary subfamily, the Eucosmodontinae. The convergence was far from complete and
isolated incisors of these and other multituberculates can be distinguished from the
ineisors of rodents (Jepsen 1937). On the basis of the size, curvature, and cross-section
of the crown and the distribution of enamel, the tooth described by Lydekker can be
identified as an incisor of a rodent. The only attribute of the fossil that precludes this
identification is its supposed stratigraphic provenance. The fossil was preserved in a

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

57

block of stone picked up on the shore. Its stratigraphic provenance was determined
through identification of the enclosing sediment as ‘Tilgate Grit’, a name once applied
to calcareous sandstones now known to be present at various levels within the Hastings
Beds. This identification was challenged by Dawson and later Simpson (1928, p. 192)
commented: ‘The specimen, moreover, is not in such a sandstone [Tilgate Grit] but in
a very crumbly clay, so that its exact level must be considered very dubious.’

To the best of my knowledge no teeth closely resembling the fossil described by
Lydekker have been found in situ in English Wealden strata. The tooth from Hastings
is similar in morphology and, superficially at least, in mode of preservation to upper
incisors of Theridomys sp. found in argillaceous. Tertiary strata cropping out on the Isle
of Wight and adjacent parts of the mainland. The nearest outcrops of these strata are
over 70 miles away from Hastings. Similar incisors of rodents also have been found in
Tertiary strata of the London basin that extend to within approximately 35 miles of
Hastings. Dawson was first to suggest that the fossiliferous block might not have come
from the Wealden cropping out near Hastings but might have been introduced from
some other area. Although well removed from the Hastings district, the Tertiary strata
of the Hampshire and London basins are not so distant as to preclude all possibility of
the specimen being accidentally introduced from either area.

In summary, if an identification were based on dental morphology alone the tooth in
question would be identified as a rodent incisor. The only controverting evidence is the
supposed stratigraphic provenance of the fossil. The identification of the enclosing sedi-
ment as Tilgate Grit is incorrect, and there is no unequivocal evidence to support the
former assignment to the Early Cretaceous. The block containing the fossil possibly was
a fragment of a Tertiary stratum drifted on to the beach at Hastings or accidentally
introduced during road construction. Although final judgement should be reserved until
our knowledge of the morphology of Wealden multituberculates is more complete, the
available evidence suggests that the fossil (M5691) probably is the incisor of a Tertiary
rodent rather than a Wealden multituberculate.

In 1911 Woodward described three more specimens that had been discovered at Cliff
End by P. Teilhard de Chardin and F. Pelletier, who were assisting Dawson. Two of
these fossils are multituberculate teeth. One of them (M10480), the type of Loxaulax
valdensis Simpson, had been described in detail by Woodward (1911, pp. 278-9) and
Simpson (1928, pp. 49-50). The second (M10481) is probably a fragment of an of
a member of the same species. Both teeth are still imbedded in sediment that is litho-
logically similar to the blocks of Cliff End Bone Bed found by our field parties. The
third tooth was described by Woodward (1911, p. 278) as follows: ‘. . . most of the crown
has decayed, but the two divergent roots are well displayed, the one somewhat stouter
than the other’. This fossil was not recorded in the British Museum (Natural History)
Catalogue of Fossil Mammalia in 1912 when the other two teeth from Cliff End were
assigned specimen numbers. Recently a specimen was found in the collection with the
following label: ‘Mammal tooth, Fairliglit, Ashdown Sand, N. Hastings, C. Dawson
15/1/1911.’ This fossil, now catalogued as M20241, agrees with Woodward’s brief
description and probably is one of the fossils discovered by Teilhard de Chardin and
Pelletier. The crown is almost completely destroyed; only a small band of enamel or
enamel-like substance remains. The coronal dimensions are: length = 3T mm., width
> 1 -2 mm. The root consists of a large, cylindrical, grooved body approximately 1 -8 mm.

58

PALAEONTOLOGY, VOLUME 6

long that terminates in two divergent branches, the largest of which is approximately
1-1 mm. long. Even though the root is bifid apically it does not resemble the roots of
Late Jurassic or Early Cretaceous mammalian teeth. Also the teeth of these mammals
are, for the most part, smaller than M20241. The mammalian affinities, if any, of this
tooth will have to be regarded as indeterminable.

Thus of the fossils collected in and prior to 1911 only Ml 0480 and Ml 0481 remain as
unquestionable Wealden mammalian teeth. The mammalian affinities of two of the other
fossils, M13134 and M20241, are doubtful, and the tooth described by Lydekker,
M5691, probably was not part of the dentition of a Wealden multituberculate. After
1911 no more Wealden mammalian fossils were discovered until 1960; but interest in
the stratigraphy and the fauna of the Wealden did not decline. Of the studies carried on
after 1911, the work of P. Allen, especially his reports on the stratigraphy (Allen 1959)
and a study of Wealden bone beds (Allen 1949), and Simpson’s (1928) review of Wealden
mammals, are especially pertinent.

DESCRIPTIONS OF THE LOCALITIES AND THE MAMMALIAN EOSSILS

Grid coordinates given in the locality descriptions are those of the National Grid and
lie in the 100-kilometre square 51 (TQ).

(1) CLIFF END BONE BED

Location and collecting technique. The exact locality where Teilhard de Chardin and
Pelletier collected was not described in detail by Woodward (1911). Many years later
Allen, using a description of the site obtained from Woodward in 1938, relocated the
bone bed and published a locality description (Allen 1960o, p. 11) in which he stated:
‘. . . rapid erosion of the cliff's has re-exposed the celebrated bone-bed . . .’. On our
collecting trips in the autumn of 1960 and the following spring an outcrop of the bone
bed could not be found; apparently the forces of erosion had destroyed or covered what
they once revealed. Loose blocks of bone bed containing the new mammalian fossils
were, however, collected from the wave-cut platform and the shingle; most were found
within 100 yards of the foot of the cliff. These blocks ranged in size from cobbles to
a fragment of the stratum nearly 4 feet in diameter and were distributed over an area
(text-fig. 1) extending northward from a point approximately 2,000 feet north-east of
Haddock’s Cottages (88751280) to a point near the northern end of the cliff (88851305).
Because no bone beds could be found cropping out in the area, it is assumed that all the
loose blocks were derived from the intermittently exposed Cliff End Bone Bed described
by Allen.

The Cliff End Bone Bed is a thin stratum composed of the comminuted bones and
teeth of fish, amphibians, reptiles, and, very rarely, mammals concentrated in a hard,
calcareous, coarse sandstone. The stratum, part of the Ashdown Beds, is never more than
4 or 5 inches thick, and its upper surface is ripple-marked. In order to find the minute
mammalian fossils a large quantity of rock, 2 or 3 hundredweights, had to be processed.
After some experiment the following technique was found to be most efficient: The
calcareous cement was dissolved with dilute formic acid (a 10 to 20 per cent, solution).
The material that remained was washed and dried and then passed through a set of
graded sieves. The coarser fraction, material caught in 12-mesh and coarser sieves, was
sorted without further treatment. The finer fraction was placed in a mixture of bromo-

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

59

form or tetrabromethane and dibromethane to give the density required to separate the
quartz from the residue of fossils and heavy minerals. This residue was then washed and
sorted. Separating the quartz was of particular importance because it greatly reduced
the volume of fine fraction that had to be sorted under a microscope.

TEXT-FIG. 1 . Cliff End Bone Bed, locality map. Area depicted in map on left shaded in the

index map on right.

Mammalian fauna. The only fossils in earlier collections that remain as unquestionable
teeth of Wealden mammals are the two multituberculate teeth found at Cliff End. One
fossil, M10480, is the type specimen of Loxaiilax valdensis; the other, M10481, is also
a fragment of a multituberculate molar identified as ‘ ? Loxaulax sp.’ by Simpson (1928).
To date five more mammalian fossils have been discovered in blocks of the Cliff End
Bone Bed.

Subclass ALLOTHERiA Marsh 1880
Order multituberculata Cope 1884
Family plagiaulacidae Gill 1872
Genus loxaulax Simpson 1928

Loxaulax valdensis (Woodward 1911)

Specimen M21098 (text-fig. 2). The presence of a small expansion of the crown lateral
to the two rows of cusps, the morphology of the cusps,
and the morphology and proportions of the crown (length
= 2-4 mm., width = 1-9 mm.) indicate that M21098 is
probably a right M^. The first (anterior) and second lingual
cusps are demarcated by a shallow groove, a deeper in-
dentation separates the second and third, and the third
and fourth are separated by a deep cleft. The first, second,
and third lingual and two labial cusps are of approximately
equal height; the fourth lingual cusp is lower. Both the
posterior lingual and posterior labial cusps are slightly
selenodont with the flatter face anterior. The anterior edge
of the crown in i;oncave to allow to abut against the posterolingual corner of M^.

TEXT-FIG. 2. Loxaulax valdensis
Simpson, right M21098,
X 15. Occlusal view.

60

PALAEONTOLOGY, VOLUME 6

On the labial side of the anterior labial cusp there are several irregular, enamel-lined
pits that do not appear to be the result of abrasion during mastication or post-mortem
damage. The occlusal surface of the labial expansion of the crown slopes gradually from
the anterior labial cusp to the edge of the crown; it is smooth and there is no evidence
of a cusp or ridge.

Although the molars of plagiaulacids have only two main rows of cusps, in some
species, for example, Bolodon osborni, the upper molars also have small expansions of
the crown or cuspules in the areas in which a third cusp row is present on ptilodontid or
taeniolabidid molars. Another character of the plagiaulacids is that the lengths of the
molars of an individual are approximately equal while in members of the other families
the first molars tend to be longer than the second. Simpson (1928, p. 49) tentatively
identified the type specimen of Loxaulax valdensis as a left Mj. M2 1098 is only 0-4 mm.
longer than the type specimen. Although not so pronounced as in the type, its posterior
cusps are vaguely selenodont. The labial expansion of the crown appears to be rela-
tively larger than that found on the M^’s of Purbeck multituberculates. In contrast the
posterior cusps of the M‘^ of B. osborni, which is smaller, have a more distinctly seleno-
dont shape than those of M21098. Because M21098 is approximately the same length as
the type of L. valdensis, because some of the cusps of both teeth are slightly selenodont,
and because there is at present no evidence demonstrating the presence of more than one
species of multituberculate in the Cliff End fauna, M2 1098 is provisionally allocated to
L. valdensis. Future collections will no doubt yield evidence of other multituberculates.
However, for the purposes of this preliminary report the best procedure is to refrain
from the introduction of species on the probability that they will be validated by subse-
quent discoveries.

Acomparison of M21098 with M10481, a fragment of a molar identified as Loxaulax

sp.’ by Simpson (1928, p. 50), suggests that the orientation of the fragment proposed by
Simpson is not correct. If Ml 0481 be regarded as a fragment of the posterior end of an
M‘\ then a number of resemblances to M21098 are apparent. The cusps of M 10481 that
Simpson thought might correspond to the ‘antero-external’ cusps of the type specimen
are comparable to the posterior lingual cusps of M2 1098. The labial side of the base of
the anterior labial cusp of Ml 0481, using the orientation proposed here, is slightly
enlarged, suggesting the presence of a labial expansion similar to that of M2 1098. The
preserved cusps of M 1 048 1 differ from their counterparts on M2 1 098 in that the posterior
lingual cusp is as high as, but not so well separated from, the penultimate lingual cusp.
Differences of this kind have been found in large samples of the molars of ptilodont
multituberculates and cannot be accorded great significance.

Specimen M2 1099 (text-fig. 3). This tooth is a plagiaulacid, anterior upper premolar,
that is, P^, P^, or P®. The bulbous crown is surmounted by three main cusps and a small
cuspule that is little more than a ridge on the surface of the enamel. Coronal dimensions
are: length = T4 mm., width =1-2 mm. There are two distinct but not divergent roots.
M21099 closely resembles the P^ of Bolodon osborni, which has three main cusps and an
anterolabial cuspule.

Specimen M21 106 (text-fig. 4). There are four main cusps on the crown of this anterior
upper premolar. Two of them, probably the lingual cusps, are higher than their lateral

W. A. CLEMENS: WEALDEN MAMMALIAN EOSSILS

61

counterparts. The labial cusps are of different size; judging from the morphology of other
multituberculate premolars the smaller cusp can be identified tentatively as the antero-
labial cusp. There is a small cuspule anterior to the anterolingual cusp and another
cuspule posterior to the posterolabial cusp. The crown is 0-9 mm. long and 0-8 mm. wide
and was supported by two roots.

a

b

c

TEXT-FIG. 3. Loxaiilax valdensis Simpson, anterior upper premolar, probably
a left premolar, M21099, X 15. a. Labial view; b, occlusal view; c, lingual

view.

TEXT-FIG. 4. Loxaiilax valdensis Simpson, anterior upper premolar, probably
a right premolar, M2 1 106, X 30. a. Labial view; b, occlusal view; c, lingual view.

Most of the anterior upper premolars of Jurassic plagiaulacids have three main cusps.
The of Bolodon osborni that has a small fourth cusp or cuspule and the of Psalodon

polens could be considered exceptions. The anterior upper premolars of ptilodontids
have, in many species, a more complex cuspation. M21106, which has four main cusps,
is more closely comparable to anterior upper premolars of certain Late Cretaceous
ptilodontids. Probably M2 1 106 is not a P^ or P^, because in Jurassic plagiaulacids these
teeth are proportionately longer relative to width and have more than four cusps. It
can be suggested that M2 1099, which is the larger and has a simpler crown, is a P^ or P'^,
and M21 106 is a P^ or P^. Both teeth are of the size that would be expected for premolars
of Loxaidax valdensis and are provisionally allocated to this species. If these identifica-
tions are correct the premolar dentition of L. valdensis is slightly more ptilodont-like
than those of the Jurassic plagiaulacids.

Specimen M2 1100 (text-fig. 5). One side of the crown of this single-rooted incisiform
tooth is smooth and convex; the other is divided into three concave facets. There are

62

PALAEONTOLOGY, VOLUME 6

three small cuspules at the base of the medial facet. None of the incisors preserved in

English Jurassic mammalian fossils closely
resembles this tooth. The lower incisors
of Trioracodon ferox and Spalacotheriiim
thcuspidens are grossly similar but the
concave sides of their crowns are not
subdivided and basal cuspules are not
present. Closer resemblances can be found
to the P of the American Jurassic plagi-
aulacid Psalodon fort is, and the upper in-
cisors of some Late Cretaceous and early
Tertiary ptilodontids. Therefore M21 100 is
provisionally identified as an upper incisor
of a multituberculate and allocated to Loxaidax valdensis.

Subclass THERIA Parker and Haswell 1897
Infraclass pantotheria Simpson 1929
Order eupantotheria Kermack and Mussett 1958
Family dryolestidae Marsh 1879
Genus cf. melanodon Simpson 1927

cf. Melanodon goodrichi Simpson 1929

Specimen M21101 (text-fig. 6). The fossil consists of the lingual part of the crown and
part of the lingual root of an upper molar of a eupantothere. The lingual cusp is high
and from its slope three prominent, heavily worn ridges extend labiad. Even though the

TEXT-FIG. 5. Loxaulax valdensis Simpson, probably
an upper incisor, M21 100, X 7-5. a and b. Lateral
views.

molar, M21 101, Xl5. a. Mesial view; Z), occlusal view; r, distal view.

specimen suffered post-mortem abrasion, the facets on the ridges are so distinct that they
must have been well developed prior to the death of the individual. One of the lateral
facets is slightly expanded suggesting the presence of a small cusp.

The Cliff End fossil differs from the molars of the species of Amblotheriinn in the
presence of a large, distinct, median transverse ridge; a structure not present on the
molars of A. pusillum and small on A. nanimi molars. The median transverse ridges on
molars of Kurtodon pusiUus are larger and more distinct than those on the molars of
A. nanum, but not so prominent as their counterpart on M2 1101. Also neither of the

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

63

lateral transverse ridges on the molars of K. pusillus have an expansion similar to that
found on one of the lateral transverse ridges of M21101. As far as can be determined
from the illustrations, M21101 is closely comparable to molars of Melanodou goodrichi
from the American Morrison Formation (Simpson 1929, pi. xv, figs. 1, 2). The molars
of the type specimen of M. goodrichi have three distinct transverse ridges, there is an
expansion of the posterior transverse ridge immediately labial to the lingual cusp, and
the transverse ridges appear to have wear facets along their crests. Although not to be
construed as a positive or even provisional identification, on the basis of the few avail-
able characters the comparison with M. goodrichi appears to be close.

TEXT-FIG. 7. Paddockhurst Bone Bed, locality map. Area depicted in map on left shaded
on the index map on right. Geologic details taken from a hitherto unpublished map by
Professor J. H. Taylor. Abbreviations: UTWS, Upper Tunbridge Wells Sand; GC,
Grinstead Clay; LTWS, Lower Tunbridge Wells Sand.

(2) PADDOCKHURST BONE BED

Location and collecting technique. This bone bed was discovered by Professor J. H. Taylor
in the course of mapping the geology of Paddockhurst Park. The fossiliferous stratum
cropped out in a now abandoned clay pit (53281334) near a park house south-west of
Turners Hill (text-fig. 7). The bone bed, which is part of the Grinstead Clay, is a grey,
shell-rich limestone. The vertebrate fossils concentrated in it are comminuted but are
not so fragmentary or heavily abraded as those found in the Cliff End Bone Bed. During
the autumn of 1960 Kermack and 1 were fortunate to receive several samples of the bone
bed. In January 1961 with the help of Professor Taylor, Dr. Cox, and Miss Lees we col-
leeted all the fragments of the bone bed that were still visible. Further collecting will
involve a large amount of preliminary work for in addition to slumping of the pit walls
and the presence of luxuriant undergrowth, a number of trees have blown over and
toppled into and across the pit.

The process used to free the vertebrate fossils from the rock is similar to that applied to
the Cliff End Bone Bed. After the rock had been treated with dilute formic acid, usually
all that remained with the fossils was a small residue of clay and ferruginous material.

64

PALAEONTOLOGY, VOLUME 6

Some of the residual clay could be broken down by thoroughly washing and drying the
material left after the acid treatment and then placing it in a hydrogen peroxide solution
(30 vol.). On completion of the chemical treatment the fossils and other residual materials
were passed through a graded set of sieves. The finer fractions were sorted with the aid
of a binocular microscope.

Mammalian fauna. To the best of my knowledge the specimens described below are the
only mammalian fossils discovered in the Paddockhurst Bone Bed.

? Subclass ALLOTHERiA Marsh 1880

Specimen M21102 (text-fig. 8). The crushed root is long and appears to have been of
approximately the same diameter throughout its length, but it may have been con-
stricted apically. One side of the crown is a
shallow, concave basin that is subdivided by a
low, longitudinal ridge; the other side (shown
in text-fig. 8) is convex and smooth. The tip of
the crown has been blunted by wear and the
eminence behind the apex may be the base of
a cusp. Among the Mesozoic mammals only the
multituberculates are known to have had large, long, somewhat gliriform incisors; there-
fore M21102 is allocated with a query to the Subclass Allotheria.

Subclass THERIA Parker and Haswell 1897
Infraclass pantotheria Simpson 1929
Order symmetrodonta Simpson 1925
Family spalacotheriidae Marsh 1887
Genus cf. spalacotherium Owen 1854

cf. Spalacotherium tricuspidens Owen 1854

Specimen M2 1 103 (text-fig. 9). The paraconid and metaconid of this left lower molar are
close to the base of the protoconid, separated from it by shallow grooves. The basal
cingulum encircles the crown, and a small anterior and a slightly larger, posterior cingular
cusp are present. On the posterolabial side of the crown there are two indentations in the
cingulum ; probably the lingual indentation received the anterior end of the following
molar. The convex lingual side of the crown is higher than the labial; the dimensions of
the crown are: length = 0-80 mm., width = 0-55 mm.

The acute angle formed by the trigonid cusps and the presence of a continuous lingual
cingulum indicate that M21 103 is a molar of a spalacotheriid symmetrodont. Five genera
have been allocated to this family: the Late Jurassic Spalacotherium, Tinodon, Ewy-
lambda, and Peralestes (based on an upper dentition and probably a synonym of
Spalacotherium) and the Early Cretaceous Spalacotheroides. The molars of Tinodon
differ from M2 1 103 in their wide-angled trigonid and the absence of a labial cingulum.
Among the presently known genera of spalacotheriids, Eurylambda, known only from
an upper molar, probably is most closely related to Tinodon. Allocation of these two
genera to the Spalacotheriidae, the family to which M21 103 certainly pertains, has been

TEXT-FIG. 8. ?Allotherian, incisor, M21102,
X 9. Lateral view.

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

65

questioned recently by Patterson (1956, p. 87). M21103 is more closely comparable to
the molar of Spalacotheroides, but it differs in several important characters. The angle
formed by the trigonid cusps of the Spalacotheroides molar is more acute, the crown is
shorter relative to its width, and the basal cingulum is not continuous around the
protoconid. M21 103 is smaller than any of the molars that have been referred to Spalaco-
therium tricuspidens, but otherwise it resembles any of the anterior molars of this species.
The posterior molars of S. tricuspidens, particularly M7, differ from the more anterior
molars in the smaller size of the protoconid and greater separation of the cusps of the

TEXT-FIG. 9. Cf. Spalacotlieriiim tricuspidens Owen, left lower molar, M21103, X 30.
a. Labial view; b, occlusal view; c, lingual view.

trigonid. The length of an anterior molar, to M5, of S. tricuspidens ranges from
1-6 mm. to 1-8 mm. If M21103 is correctly identified as an anterior molar its smaller
size (length = 0-80 mm.) indicates that it was part of the dentition of a spalacotheriid
distinctly smaller than S. tricuspidens. In themselves this difference in size and the
temporal separation of the symmetrodonts found in the Paddockhurst Bone Bed and the
Purbeck Beds do not warrant the establishment of a new species. Because there is an
excellent chance that more material of this Wealden symmetrodont will be discovered,
M21103 has not been assigned to a genus or species; only a comparison with Spalaco-
therium tricuspidens is suggested.

? Subclass THERiA Parker and Haswell 1897

Specimen M2 1 104 (text-fig. 10). The crown of this small, trenchant premolariform tooth
(length 0-85 mm., width 0-25 mm.) consists of a large, central cusp and smaller
anterior and posterior accessory cusps. On one side of the crown there is a short ridge
on the slope of the anterior accessory cusp and a longer but less distinct, cingulum-like
expansion of the base of the crown on the posterior half of the central cusp. The two
well-separated and divergent roots, which appear to be complete, have narrowly oval
cross-sections ; their length and disposition suggest that M2 1 104 is a deciduous premolar.

Identification of M2 1104 poses a number of problems. Considering the roster of
mammals that could be expected in an Early Cretaceous fauna only the representatives
of one order, Multituberculata, lacked premolars grossly similar to M21 104. In spite of

C 1015

F

66

PALAEONTOLOGY, VOLUME 6

the lack of great morphologic differentiation, the premolars of other Late Jurassic and,
presumably, Early Cretaceous mammals differ in a few minor characters that might be
of taxonomic significance. The premolars of triconodonts have at least a lingual cingulum
or a definite expansion of the lingual base of the crown. Anterior and posterior accessory
and cingular cusps are common. The roots of the permanent premolars are close
together, nearly parallel, and almost twice as long as the crown is high. The DP4 of
Tricofiodon mordax, the only triconodont deciduous premolar discovered so far, is
molariform. It appears unlikely that M21104 is a triconodont premolar. Allocation to

TEXT-FIG. 10. ? Therian, premolar, M21104, x30. a. Lateral view; b, occlusal view.

the Order Docodonta also can be regarded as improbable for the premolars of Docodou,
a Late Jurassic member of this order, have a distinct lingual cingulum and exhibit other
differences in morphology. The premolars of many pantotherian species have a lingual
cingulum, but in some species of eupantotheres — for example, note the upper premolars
of Amblotherium pusillum — basal cingula are indistinct or absent. ThePaddockhurst Bone
Bed is high in the sequence of Wealden strata and mammals of eutherian-metatherian
grade might have been present at the time it was deposited. M21104 is a type of pre-
molar that might be expected in such a mammal. In summary, M2 1104 is certainly not
a multituberculate premolar and probably not a triconodont or docodont premolar.
Its allocation to the subclass Theria is based on both a process of elimination and the
observation that the premolars of some Mesozoic therians lack a basal cingulum.

Although no attempt will be made to describe the stratigraphy of the Cliff End and
Paddockhurst Bone Beds in detail, a few comments are necessary in order to interpret
the significance of the mammalian fossils.

Derived Upper Kimeridgian and basal Portlandian ammonites and other inverte-
brates were discovered by Allen {\960b) in the Top Lower Tunbridge Wells Pebble Bed.
This would suggest that the Wealden mammalian fossils, especially those found in the
coarse sands of the Cliff End Bone Bed, might have been derived from older deposits.
The Ashdown Beds in which the Cliff End Bone Bed occurs have been interpreted by
Allen (1959) as delta-face and shore-face deposits formed during a period of deltaic
advance. As would be expected in a unit of this origin, the vertebrate fossils from the
Cliff End Bone Bed are comminuted, rolled, and abraded. However, most of the speci-
mens of derived Jurassic ammonites found in the Top Lower Tunbridge Wells Pebble
Bed are internal casts of parts of a camera or one or two camerae, the largest being

a

b

STRATIGRAPHY

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

67

approximately half a whorl (Arkell 1960); and all the derived invertebrates found in this
pebble bed give evidence of transport and extended or intensive weathering. It is doubt-
ful if the small mammalian fossils found in the Cliff End Bone Bed could have survived
similar weathering and transport in addition to the fragmentation and abrasion involved
in their concentration and deposition on the delta face. In contrast to the Ashdown
Beds, the Grinstead Clay has been interpreted as a unit deposited during the lacustrine
transgression of the Lower Tunbridge Wells delta (Allen 1959). The vertebrate fossils
found in the Paddockhurst Bone Bed appear to be less fragmented and abraded than
those found in the Cliff End Bone Bed, indicating that they too probably were not
derived from older strata.

Additional evidence that the Cliff End mammals were not derived from Jurassic beds
comes from a study of the animals themselves. The little that is known of the dentition
of Loxaulax valdensis suggests that this species had a slightly more ptilodont-like
dentition than the Purbeckian plagiaulacids. Assuming that the ptilodont multi-
tuberculates evolved from a plagiaulacid ancestor, the stage of evolution of L. valdensis
suggests that the species is of post-Purbeckian age. The differences between the other
Wealden mammals and their Purbeckian counterparts can also be interpreted as being
indicative of a post-Purbeckian age. However, knowledge of the phylogeny of Mesozoic
mammals is far from complete and any age-determination based on an assessment of the
stages of evolution of these mammals can only be regarded as a suggestion. Finally,
Allen (1960Z?) has determined that the derived Jurassic fossils found in the Top Lower
Tunbridge Wells Pebble Bed were transported into the basin by a river flowing from the
north. In contrast he found that the pebbles in the Cliff End Bone Bed were brought in
from the north-east. Although the latter sediments contain glauconitic sand apparently
derived from Jurassic strata. Upper Kimeridgian and basal Portlandian pebbles from
the north-east are absent or rare. None of these lines of evidence conclusively demon-
strates that the mammalian fossils found in the Wealden bone beds were not derived
from older strata, but in concert they make this improbable.

Neither a precise determination of the stratigraphic position of the Cliff End Bone
Bed within the Ashdown Beds nor an exact correlation of its time of deposition with the
standard chronology based on the European marine sequence has been accomplished as
yet. Currently the bulk of the Wealden of England is regarded as being of post-Tithonian
age (Allen 1955, 1959, and Hughes 1958). Certainly the fossil localities are of pre-
Aptian age. The differences between members of the Cliff End and Purbeck mammalian
faunas suggest, but do not in themselves confirm, that the Cliff End Bone Bed is younger
than the mammal-bearing strata of the Purbeck Beds. Hughes (1958), on paleobotanical
evidence, allocates the Ashdown Beds to the Valanginian. Allen (1955, 1959) has
correlated the period of deposition of the Ashdown Beds with the Berriasian regression.
Thus a post-Purbeckian age is indicated for the Cliff End Bone Bed. The Paddockhurst
Bone Bed is part of the Grinstead Clay, a unit stratigraphically higher than the Ashdown
Beds. Using paleobotanical data, Hughes (1958) allocated the Tunbridge Wells Sand-
stone, which includes the Grinstead Clay, to the later Hauterivian. Allen (1959) sug-
gested that the Grinstead Clay was deposited during a lacustrine transgression associated
with later Valanginian or early Hauterivian changes in the level of the Neocomian sea in
the Paris basin.

In summary, it appears that the Wealden mammalian fossils were not derived from

68

PALAEONTOLOGY, VOLUME 6

older strata. The mammalian fauna of the Cliff End Bone Bed is probably of early
Neocomian age, whereas that of the Paddoekhurst Bone Bed is distinctly younger. Both
faunas are post-Purbeckian and pre-Aptian in age.

CONCLUSION

The discovery of these few Wealden fossils has increased knowledge of the morpho-
logy of the plagiaulacid multituberculate Loxaulax valdensis; extended the demonstrated
range of the Eupantotheria, previously limited to the Middle and Late Jurassic; and
established the presence of the Symmetrodonta in England in the Early Cretaceous. The
mammalian fauna of the Cliff End Bone Bed, which is part of the Ashdown Beds, con-
sists of L. valdensis and a dryolestid eupantothere; the fauna of the younger Paddock-
hurst Bone Bed includes a spalacotheriid symmetrodont and possibly another therian
and a plagiaulacid. These discoveries indicate that with proper collecting techniques
more mammalian fossils probably can be obtained from these and other Wealden bone
beds.

Acknowledgements. My sincere thanks are extended to Dr. K. A. Kermack for his continuing and
extensive help and encouragement. I am indebted also to Professor J. H. Taylor for showing us the
Paddoekhurst Bone Bed and giving me the opportunity to quote from his unpublished work on the
geology of the area, and to Professor P. Allen for specimens and helpful comments. I also gratefully
acknowledge the assistance received from Drs. E. I. White and A. Sutcliff, who gave me the privilege
of studying material under their care, and Dr. Barry Cox, Mr. H. A. Toombs, and Mr. B. Newman,
who allowed me to process their samples of the Paddoekhurst Bone Bed. Miss Patricia Lees and Mrs.
Francis Mussett have provided valuable assistance in both field and laboratory work. Thanks are due
also to Professor P. B. Medawar for making available the necessary laboratory facilities. The field and
laboratory work was carried out during the tenure of a National Science Foundation Postdoctoral
Fellowship at University College, London, and this report completed at the Museum of Natural History,
The University of Kansas.

REFERENCES

ALLEN, p. 1949. Notes on Wealden bone-beds. Proc. Geol. Assoc. 60, 275-83.

1955. Age of the Wealden in north-western Europe. Geol. Mag. 92, 265-81.

1959. The Wealden environment: Anglo-Paris basin. Phil. Trans. Roy. Soc. B, 242, 283-346.

1960(7. Geology of the Central Weald: a study of the Hastings Beds. Geol. Assoc. Centenary

Guide, No. 24, 1-28.

\96Qb. Strand-line pebbles in the mid-Hastings Beds and the geology of the London uplands.

General features. Jurassic Pebbles. Proc. Geol. Assoc. 71, 156-68.

ARKELL, w. j. 1960. Report on derived Jurassic ammonites from the top lower Tunbridge Wells
Pebble Bed. Appendix to allen, 19601?.

CLEMENS, w. A. 1960. Explanation of an exhibit of specimens of new Wealden mammals. Proc. Geol.
Soc. London, 1588, 90.

HUGHES, N. F. 1958. Palaeontological evidence for the age of the English Wealden. Geol. Mag. 95,
41-49.

JEPSEN, G. L. 1937. A Paleocene rodent, Paramys ataviis. Proc. Amer. Philos. Soc. 78, 291-301, pi. 1.
LYDEKKER, R. 1893. On a mammalian incisor from the Wealden of Hastings. Quart. J. Geol. Soc.
London, 49, 281-3.

PATTERSON, B. 1956. Early Cretaceous mammals and the evolution of mammalian molar teeth.
Fieldiana: Geol. 13, 1-105.

SIMPSON, G. G. 1928. A catalogue of the Mesozoic mammalia in the Geological Department of the
British Museum. London, British Museum (Nat. Hist.), x+215, 12 pis.

W. A. CLEMENS: WEALDEN MAMMALIAN FOSSILS

69

SIMPSON, G. G. 1929. American Mesozoic mammalia. Mem. Peabody Mas. 3, xv+171, pi. 1-32.
WOODWARD, A. s. 1891. Oil a mammalian tooth from the Wealden Formation of Hastings. Proc.
Zool. Soc. Loudon, 585-86.

• 1911. On some mammalian teeth of the Wealden of Hastings. Quart. J. Geol. Soc. London,

67, 278-81.

WILLIAM A. CLEMENS

Museum of Natural History
and Department of Zoology,
The University of Kansas,

Manuscript received 20 February 1962 Lawrence, Kansas, U.S.A.

LOWER PERMIAN BRYOZOA FROM WESTERN

AUSTRALIA

by JUNE R. P. PHILLIPS ROSS

Abstract. Scattered bryozoan localities in the Lyons Group contain three species of Stenopora, S. dickinsi
sp. nov., 5. fisheri sp. nov., and S. lyndoni sp. nov., that show little similarity to previously described species.

The fenestellid Polypora lyndoni occurring at all the bryozoan localities studied has close affinities with two
species from the Lower Permian of eastern Australia and one species from the Asselian of Timor. These species
appear to be primitive members of a group of species described from the upper part of the Sakmarian Series of
the Urals.

Fossils are generally rare and sporadic in the Lyons Group, which crops out in a belt
extending from the Lyndon River to the Wooramel River in the Carnarvon (North-
West) Basin, Western Australia (text-fig. 1). The group is about 4,600 feet thick in the
Moogooree area and thins southward to 2,640 feet in the Wooramel River area (Condon
1954, pp. 40, 43).

The Lyons Group begins with the Permian? lepidodendroid plant-bearing Harris
Sandstone, which rests conformably on the Pre-Cambrian basement, and consists of
tillite, greywacke, siltstone, sandstone, and conglomerate. It is conformable with the
overlying Permian (Upper Sakmarian or Lower Artinskian) Callytharra Formation
(Table 1).

Previous studies of fossils from the Lyons Group include description of brachiopods
by Prendergast (1943), Coleman (1957), and Thomas (1958), description of molluscs by
Dickins (1956, 1957), description of two species of Foraminiferida by Crespin (1958),
and a record of brachiopods, molluscs, Bryozoa, and crinoids by Dickins and Thomas
(1959). The Bryozoa occur in thin, local calcareous lenses within the tillite, sandstone,
and conglomerate beds.

Age of the fauna of the Lyons Group. From the faunal data in strata overlying the Lyons
Group, Teichert (1941, 1947) considered the Lyons Group to be Sakmarian in age.
Later studies of the faunas from the Lyons Group, summarized by Dickins and Thomas
(1959), and from the overlying strata have supported Teichert’s findings.

Relations of bryozoan species in the Lyons Group. The scattered bryozoan collections that
were examined from the Lyons Group indicate that the bryozoan species range laterally
and vertically throughout the group. At locality ML 6, considered to be in the lower part
of the Lyons Group, the bryozoans Stenopora dickinsi sp. nov., and Polypora lyndoni
sp. nov. occur with the following brachiopods and molluscs listed by Dickins and Thomas
(1959): Trigonotretal sp. indet., Neospirifer sp. indet., Linoproductus (Cancrinella)
lyoni Prendergast (1943), Rhynchonellacea gen. indet., Rostrospiracea ? gen. indet.,
Deltopecten lyonsensis Dickins (1957). Higher in the succession in the upper part of the
Lyons Group in the northern part of the Carnarvon Basin at locality ML 90, the
bryozoans Stenopora dickinsi, S. lyndoni, and Polypora lyndoni, are associated with

[Palaeontology, Vol. 6, Part 1, 1963, pp. 70-82, pi. 8-11.]

J. R. P. PHILLIPS ROSS: LOWER PERMIAN BRYOZOA

71

Stutchburia variabilis Dickins (1957) and Deltopecten lyomensis Dickins (1957); and at
localities ML 106, ML 107, and ML 109 in the upper part of the Lyons Group the
bryozoans Stenopora dickinsi and Polypora lyndoni occur with the following fossils listed
by Dickins and Thomas (1959); Trigonotreta sp. nov., Linoproductus (Cancrinel/a) lyoni
Prendergast (1943), Neospihfer sp. nov., Pseudosyrinx sp. nov., Stutchburia variabilis
Dickins (1957), Astartila condoni Dickins (1957), Cleobis sp., Eurydesma sp. indet.,
Schizodus crespinae Dickins (1957), Deltopecten lyonsensis Dickins (1957), Aviculopecten
sp. indet., Peruvispira umariensis (Reed) (1928), Mourlonia? lyndonensis Dickins (1957),

TABLE 1

Suggested correlation of Permian units referred to in text

Carnarvon

Basin,

Ural Mis., Western

U.S.S.R. Australia

Yarrol

Basin,

Queensland,

Australia

Hunter River
Valley,
N.S. W.,
Australia

South
Coast,
N.S. W.,
Australia

Eastern

Tasmania,

Australia

Timor

Glass Mts.,
Texas,
U.S.A.

Kungurian

Artinskian

Westley

Park

Tuffs

?.

Sakmarian

Callytharra
Formation
? ?-

9.

Asselian

Lyons

Group

Lake Creek Allandale

Beds Formation

(Eurydesma
cor datum)

Lochinvar

Formation

Bitauni

Beds

Lenox Hills

c

ai

0

Formation

d

S

e>3

Berriedale

Neal Ranch

O

Limestone

Formation

Keeneia carnarvonensis Dickins (1957), Calceolispongidae gen. et sp. nov., Conularia sp.,
and Calcitornella stepliensi (Howchin) (1894).

The fenestellid Polypora lyndoni is closely comparable with two species of Polypora
occurring in the lower Permian of eastern Australia; one of the species, Polypora
keppelensis Crockford 1962, is from the Lake Creek Beds, Queensland, and the other,
P. pertinax, is from the Eurydesma cordatum horizon. New South Wales. It also closely
resembles P. tripliseriata Bassler from the Asselian Bitauni Beds of Timor. All these
species appear to be primitive members of a group of species including P. tuberculifera,
P. punctata, P. subvariicellata, and P. nadinae described from the Sterlitamak Beds
(upper part of the Sakmarian Series) of the Urals.

The stenoporids from the Lyons Group show little similarity with previously described
species from Australia. However, few species of Stenopora have been described from the
lower part of the Permian succession in eastern Australia. The four bryozoan species of

72

PALAEONTOLOGY, VOLUME 6

the Lyons Group show no similarities with those in the overlying Callytharra Formation.
The genera occurring in the Callytharra Formation described by Crockford (1951),
including Hexagonella, Evactinopora, Protoretepora, Penniretepora, Ramipora, Streblo-
trypa, Rhombocladia, and StrebJocladia, were not found in the Lyons Group.

TEXT-FIG. 1. Index map to bryozoan collection localities in the Lyons Group, Carnarvon Basin,
Western Australia. Hatching indicates outcrop area of Lyons Group. Map after M. A. Condon,

J. M. Dickins, and G. A. Thomas.

Acknowledgements. I express my sincere thanks to Drs. J. M. Rayner, Director; N. H. Fisher, Chief
Geologist ; and J. M. Dickins, Palaeontologist ; of the Bureau of Mineral Resources, Australia, for the
loan of the specimens examined from the Lyons Group. Dr. J. M. Dickins kindly helped locate the
material and compiled information about the collection localities and their stratigraphic position.

Repository. CPC — Commonwealth Palaeontological Collections, Bureau of Mineral Resources,
Canberra, A.C.T., Australia.

SYSTEMATIC DESCRIPTIONS
Order trepostomata

Family stenoporidae Waagen and Wentzel

Genus stenopora Lonsdale

1844 Stenopora Lonsdale, p. 178.

1845fl Stenopora Lonsdale, p. 262.

1943 Stenopora Lonsdale; Crockford, p. 261.

1962 Stenopora Lonsdale; Ross and Ross, pp. 41, 42.

1929 JJlrichotrypa Bassler, p. 55.

J. R. P. PHILLIPS ROSS: LOWER PERMIAN BRYOZOA

73

1844 Tubuliclidia Murchison and Verneuil [nomen nudum], pp. 497-8.

18456 Tubuliclidia Murchison, Verneuil, and Keyserling [nomen nudum], p. 221.

1949 Tubuliclidia Duncan [nomen nudum], p. 131.

Type species. Stenopora tasmaniensis Lonsdale, 1844, p. 161, from the Permian, Tasmania, Australia.

Diagnosis (after Crockford 1943). Colonies are ramose, encrusting, laminar, massive, or
frondescent. The zooecial tubes have thin walls in the axial region and irregularly
thickened, moniliform walls in the peripheral region. Diaphragms are extremely rare or
absent. Acanthopores are well developed, are generally large and very numerous, and
commonly occur in two series. Mesopores lack diaphragms and are generally fewer in
number than the zooecia. Monticules, and less commonly maculae, are generally present
except for some ramose species which lack them.

Stenopora dickinsi sp. nov.

Plate 8, figs. 1-3, 5-8; Plate 9, figs. 1-5, 7-9

Type material. Holotype CPC ML 109-2, and paratypes CPC ML 109-1, ML 109-3; from the upper
part of the Lyons Group, locality ML 109, 410 feet west of ML 106 in the Lyndon River area. Paratypes
CPC ML 106-1 to ML 106-5; from the upper part of the Lyons Group, locality ML 106, approxi-
mately 3 miles north-north-east of Round Hill Well, Lyndon River area. Paratypes CPC ML 90-2,
ML 90-4; 45 feet below the top of the Lyons Group, approximately 2 miles north-north-east of Round
Hill Well, Lyndon River area.

Description. Colonies are slender, branching cylindrical stems, the fragments studied
being 1 to 2 cm. in length (PI. 9, figs. 2, 4). Maculae and monticules were not observed.
Some colonies display broad, flat, proximal bases about 8 mm. in diameter (PI. 9,
fig- 8).

In tangential sections the oval zooecial openings are regularly aligned longitudinally
(PI. 8, figs. 6, 7, 8). The zooecial walls are narrower or about the same width as the
zooecial openings and are penetrated by prominent acanthopores, generally 0-05 to
0-09 mm. in diameter, but smaller acanthopores with diameters as small as 0-02 mm. are
occasionally present. Mesopores are very rare and, if present, are about the same size as
the acanthopores (PI. 8, fig. 6).

In longitudinal sections the zooecial walls are longitudinally laminate in the axial
region and may show two or three narrow bands of monilae (PI. 9, fig. 5). The zooecial
walls thicken considerably in the peripheral region and display low, broad monilae that
have a laminate microstructure in which steeply inclined laminae line the inner parts of
the zooecial walls and pass into the outer part of the zooecial walls as broad, distally
convex laminae. The peripheral region is as wide as the axial region (PI. 8, fig. 2).

In transverse sections the laminate microstructure of the zooecial walls in the peri-
pheral region is well displayed. The laminae curve steeply from the inner part near the
zooecial tube and make a broad distally convex arc in the outer part of the zooecial wall.
Laminae of adjacent zooecia intertongue inconspicuously in the outer region (PI. 8,
fig. 3). In the peripheral region the steeply inclined laminae of the acanthopores cut
through the zooecial walls (PI. 9, figs. 1, 3). In the axial region where the zooecial walls
are slightly moniliform, the polygonal zooecial tubes may have acanthopores at their
wall junctions (PI. 8, fig. 5; PI. 9, fig. 1).

Remarks. The species is characterized by colonies of small diameter with five to ten

74 PALAEONTOLOGY, VOLUME 6

distinct acanthopores in slender, moniliform walls. Mesopores and diaphragms are very
rare.

Stenopora dickinsi is not closely similar to species described either from the Permian
of the Fitzroy Basin (Crockford 1957) or from eastern Australia. It shows certain

TABLE 2

Measurements of Stenopora dickinsi sp. nov. (in mm.)

CPC numbers

ML 109-2

ML 90-2

ML 90M

ML 106-1

ML 106-5

Diameter of zoarial stem

No. of zooecia per 2 mm. longitu-

3-5-40

5-6

4-5

4

2-3-2-5

dinally .....

3-5-4-5

3-4-5

3-4

3

4

Diameter of zooecial opening max.

0-35 x 0-21

0-40x0-24

0-31 xO-21

0-39x0-17

0-33x0-21

min.

0-24 X 01 6

0-24x0-15

0-26x0-12

0-26x0-15

0-21X0-12

Diameter of mesopore .

—

—

—

—

(0-05-0-08) X
0-05

Diameter of acanthopore

003-009

0-03-0-07,

generally

0-05-0-07

0-05-0-09

0-02-0-09

0-04-0-10

Zooecial wall thickness long.

01 1-0-27

007-0-26

0-10-0-39

0-15-0-23

0-10-019

lat.

01 3-0-29

0-07-0-21

0-07-0-23

0-12-0-28

0-10-0-16

No. of diaphragms in zooecial tube

0-1

—

—

—

—

Width of peripheral region .

Ratio: width of peripheral part of
zooecium/total width of zoo-

0-88-0-90

1-26

1-4-1-6

1-54

0-55

ecium .....
No. of large acanthopores per zoo-

0-5

0-5

0-64

0-63

0-66

ecial opening ....
No. of small acanthopores per zoo-

3-8

5-8

6-10

5-8

5-8

ecial opening ....
No. of mesopores per zooecial

0-1

—

—

0-1

0-1

opening .....

—

—

—

—

0-2

similarities to S. gracilis (Dana) (Crockford 1943) from the Westley Park Tuffs, Permian,
New South Wales, Australia, in its broad basal attachment, size of its zoarial stem,
number of zooecial openings per 2 mm., remote monilae across the axial region, broad
monilae in the peripheral region, and sparse distribution of mesopores. However,

EXPLANATION OF PLATE 8

Figs. 1-3, 5-8. Stenopora dickinsi sp. nov. 1, Tangential section showing oval zooecial openings and
zooecial walls pierced by large acanthopores, locality ML 109, CPC ML 109-2, X 50. 2, Longi-
tudinal section showing slender zooecial walls in axial region and greatly thickened, laminate walls
in abraded peripheral region, locality ML 106, CPC ML 106-5, x20. 3, Part of transverse section
showing laminate microstructure of zooecial walls in the peripheral region, locality ML 90, CPC
ML 90-2, X 50. 5, Transverse section showing thin-walled, polygonal zooecia in axial region and
thickenedzooecial wallsintheperipheralregion,localityML 109, CPC ML 109-3, x20. 6,Tangential
section showing large acanthopores and occasional mesopore around zooecial openings, locality
ML 106, CPC ML 106-2, X 50. 7, Tangential section showing longitudinal alignment of zooecial
openings, locality ML 109, CPC ML 109-2, X 20. 8, Tangential section showing laminate micro-
structure in zooecial walls and acanthopore walls, locality ML 109, CPC ML 109-2, X 100.

Fig. 4. Stenopora fisheri sp. nov. Part of longitudinal section showing moniliform zooecial walls in
peripheral and subperipheral regions, locality ML 107, CPC ML 107-4, x20.

Palaeontology, Vol. 6

PLATE 8

PHILLIPS ROSS, Lower Permian Bryozoa

J. R. P. PHILLIPS ROSS: LOWER PERMIAN BRYOZOA

75

S. dickinsi has slightly larger zooecial openings, narrower zooecial walls, fewer acantho-
pores, and a considerably wider peripheral region. parallela Crockford (1945) from
the Permian, Mt. Wellington, Tasmania, is similar to S. dickinsi in the size of its zoarial
stem, diameter of the zooecial openings, broad monilae in the peripheral region, and
remote monilae in the axial region. It differs from S. dickinsi in having fewer acantho-
pores per zooecium, fewer zooecia per 2 mm., and a narrower peripheral region.

The species is named in honour of Dr. J. M. Dickins, Bureau of Mineral Resources,
who has studied extensively the Permian faunas of Western Australia.

Stenopora fisheri sp. nov.

Plate 8, fig. 4; Plate 10, figs. 1-9

Type material. Holotype CPC ML 107-4, paratypes CPC ML 107-2, -5, -8; uppermost part of the
Lyons Group, locality ML 107, Lyndon River area. Paratype CPC ML 109-4, upper part of the Lyons
Group, locality ML 109, Lyndon River area.

Description. Colonies are slender, cylindrical branching stems (PI. 10, figs. 2, 3). Monti-
cules and maculae were not observed.

TABLE 3

Measurements of Stenopora fisheri sp. nov. (in mm.)

CPC numbers

ML 107-4

ML 107-5

ML 109-4

Diameter of zoarial stem .

5-8

5-7

2-5-3-0

No. of zooecia per 2 mm. longitudinally

3-4

3-5-4

4-4-5

Diameter of zooecial opening

max.

0-48 X 0-36

0-43x0-28

0-46x0-32

min.

0-26X019

0-28x0-17

0-32x0-21

Diameter of mesopore

max.

0-38X0-12

0-16x0-13

0-05x0-05

min.

005x005

0-08x0-05

0-26x0-19

Diameter of acanthopore .

002-016

0-02-0-08

0-02-0-13

Zooecial wall thickness

long.

005-008

0-10-0-21

0-05-0-16

lat.

005-0 17

0-10-0-21

0-04-0-16

No. of diaphragms in zooecial tube

0-1 or 2

0

0

Width of peripheral region

1-76; 2-0

2-2; 2-5

0-88; 0-66

Ratio: width of peripheral part of zooecium/total

width of zooecium

0-68; 0-57

0-66-0-70;

0-66; 0-60

0-55

No. of large acanthopores per zooecial opening .

5-8

5-6

4-5

No. of small acanthopores per zooecial opening .

0-2

1-2

0

No. of mesopores per zooecial opening

1-5

0-3

1-3

In tangential sections the zooecial openings are irregularly spaced and enclosed by
slender walls considerably narrower than the zooecial openings (PI. 10, figs. 1, 4, 6-8).
The narrow walls are penetrated by large, dense acanthopores that may protrude into
the zooecial openings (PI. 10, figs. 4, 7). Numerous shallow mesopores also lie in the
zooecial walls between adjacent zooecia (PI. 10, figs. 7, 8).

In longitudinal sections the zooecial walls are slender and longitudinally laminate in
the axial region (PI. 8, fig. 4). The zooecial tubes curve in a broad arch into the peripheral
region and are slightly oblique to the zoarial surface. The zooecial walls thicken in the

76

PALAEONTOLOGY, VOLUME 6

peripheral region and display narrow, elongate monilae (PI. 10, fig. 5) and have a
laminate microstructure in which steeply inclined laminae line the inner parts of the
zooecial walls and pass into the outer part of the zooecial walls as broad distally convex
laminae (PI. 10, fig. 9). Laminae of adjacent walls merge inconspicuously in this outer
part or are penetrated by the steeply inclined laminae of the acanthopore walls. Dia-
phragms are generally lacking.

Remarks. Stenopora fisheri is characterized by slender, branching, colonies having large
zooecia, mesopores, and numerous large acanthopores enclosed in slender zooecial and
mesopore walls. It shows little similarity with previously described species, although it
resembles S. dickinsi in its slender, cylindrical stems and numerous, distinctive acantho-
pores. It differs from this species in having larger zooecial openings, larger acanthopores,
and numerous large mesopores.

The species is named in honour of Dr. N. H. Fisher, Bureau of Mineral Resources,
Australia.

Stenopora lyndoni sp. nov.

Plate 11, figs. 1-3, 6-10

Type material. Holotype CPC ML 90-1, paratype CPC ML 90-3; 45 feet below the top of the Lyons
Group; locality ML 90, Lyndon River area, approximately 2 miles north-north-east of Round Hill
Well; lat. 23° 21' S., long. 114°41'E.

Description. Colonies are slender, cylindrical, branching stems (PI. 11, figs. 6, 8, 9).
Maculae and monticules were not observed.

In tangential sections the oval zooecial openings are irregularly spaced in longitudinal
ranges (PI. 11, figs. 1, 3, 7). The elongate zooecia are separated by walls as thick as the
width of the zooecial openings. Numerous large (0-07 to 0-09 mm. diameter), dense
acanthopores penetrate the zooecial walls. Small acanthopores, 0-02 mm. in diameter,
having less dense walls, are sparse. Mesopores are small and sparsely distributed.

EXPLANATION OF PLATE 9

Figs. 1-5, 7-9. Stenopora dickinsi sp. nov. 1, Part of transverse section showing acanthopores at
junction of zooecial walls in axial region and thickened zooecial walls in peripheral region, locality
ML 106, CPC ML 106-5, x 50. 2, External aspect of fragment of colony, locality ML 106, CPC
ML 106-3, X 5. 3, Part of transverse section in peripheral region showing steeply inclined laminae of
acanthopore walls cutting through the more gently curved laminae of the zooecial walls, locality
ML 106, CPC ML 106-5, X 100. 4, External aspect of fragment of colony, locality ML 106, CPC
ML 106-2, X 5. 5, Longitudinal section showing slender walls, partly crushed, in axial region and
thickened, slightly moniliform zooecial walls in peripheral region, locality ML 109, CPC ML 109-2,
X 20. 7, Part of longitudinal section showing narrow peripheral region of thickened walls, locality
ML 106, CPC ML 106-5, X 50. 8, External aspect of zoarial fragment showing broad base of attach-
ment, locality ML 106, CPC ML 106-1, X 5. 9, Part of transverse section showing thin-walled poly-
gonal zooecia in the axial region, and thickened zooecial walls in the peripheral region, locality
ML 90, CPC ML 90-2, X 20.

Figs. 6, 10-12. Polypora lyndoni sp. nov. 6, External aspect of fenestrate colony showing meshwork
pattern and zooecial openings, locality ML 107, CPC ML 107-20, X 5. 10, External aspect of basal
part of funnel-shaped zoarium, locality ML 109, CPC ML 109-10, x 2. 11, General aspect of inner
surface of funnel-shaped zoarium near its base showing zooecial openings, locality ML 109, CPC
ML 109-10, X 2. 12, Inner surface of basal part of colony showing arrangement of zooecia and rudi-
mentary fenestrules, locality ML 107, CPC ML 107-22, x2.

Palaeontology, Vol. 6

PLATE 9

PHILLIPS ROSS, Lower Permian Bryozoa

J. R. P. PHILLIPS ROSS; LOWER PERMIAN BRYOZOA

77

In transverse sections the zooecial tubes are polygonal in the axial region. The zooecial
walls are greatly thickened in the wide peripheral region but are only slightly moniliform.
The zooecia and shallow mesopores have laminate walls in which steeply inclined laminae
line the inner parts of zooecial walls and pass into the outer parts of the zooecial walls as
broad, distally convex laminae. Laminae of adjacent walls merge inconspicuously or are
penetrated by the steeply inclined laminae of the acanthopore walls (PI. 11, figs. 2, 10).

TABLE 4

Measurements of Steuopora lyndoni sp. nov. (in mm.)

CPC numbers

ML 90-1

ML 90-3

Diameter of zoarial stem ....

5

4-5

No. of zooecia per 2 mm. longitudinally .

6

4-5

Diameter of zooecial opening . . . max.

0-26X0-19

0-23X0-13

min.

019X0-13

009x007

Diameter of mesopore .....

(005-006)x002

(0-04 to 0 06) x (0 04 to 0 05)

Diameter of acanthopore ....

002-0-09

002-0-07

Zooecial wall thickness .....

008-0-21

0-1 3-0-26

Width of peripheral region ....
Ratio: width of peripheral part of zooecium/total

1-8

not det.

width of zooecium .....

0-68

not det.

No. of large acanthopores per zooecial opening

3-4

3-6

No. of smalt acanthopores per zooecial opening

1

1-3

No. of mesopores per zooecial opening .

0-3

0-3

Remarks. Stenopora lyndoni is characterized by slender zoarial stems and small zooecial
openings enclosed by wide zooecial walls that are penetrated by prominent acanthopores
and small mesopores. It dilfers from S. dickinsi in its smaller zooecial openings and in
having small mesopores, 0-3 per zooecial opening.

Order cryptostomata
Family fenestellidae
Genus polypora M‘Coy
1844 Polypora M‘Coy, p. 206.

Type species. Polypora dendroides M‘Coy, 1844, p. 206, pi. 29, fig. 9.

Diagnosis. Infundibuliform or flabellate Fenestellidae having zooecia arranged in three
or more rows on the branches, except just after bifurcation, where only two rows may be
present.

Polypora lyndoni sp. nov.

Plate 9, figs. 6, 10-12; Plate 11, figs. 4, 5

Type material. Holotype CPC ML 107-20, paratypes CPC ML 107-21 to -28, uppermost part of the
Lyons Group, locality ML 107, Lyndon River area. Paratypes CPC ML 109-10 to -16; upper part of
the Lyons Group, locality ML 109, Lyndon River area. Paratypes CPC ML 6-5 to -10, locality ML 6
(34 miles west of north of ‘ Moogooree ’ Homestead), considered to be located in the lower part of the
Lyons Group and in its lowest known marine beds. Paratypes CPC ML 90-10 to -15, 45 feet below the
top of the Lyons Group; locality ML 90, Lyndon River area.

78

PALAEONTOLOGY, VOLUME 6

Description. Colonies are funnel-shaped and grow from circular or oval bases 3 to
15 mm. or more in diameter (PI. 9, figs. 10, 1 1). The zoarial fragments examined are up
to 40 mm. high and above the first 20 mm. the tall, narrow funnels display pronounced
flanging of the fenestrate branches. The zoarial bases of solid calcareous material rise
3 to 8 mm. or more in height, lack zooecia, and may have flanges and rootlets.

The zooecial openings lie on the inner surfaces of the fenestrate funnels and the
delicate meshwork is poorly defined in the first 2 to 5 mm. (PI. 9, figs. 11, 12).
Above this the slender, smooth, and flat zoarial branches each have three rows of
zooecia which extend continuously along the branches (PI. 9, fig. 6). Four zooecia are
generally present just before bifurcation. The obverse surfaces of the branches display
fine striae that form irregular polygons and circles around the zooecial openings. The
reverse surfaces of the colonies display longitudinally striate branches. The meshwork
formula is 8^-13 | 7^10 |1 15-17 | 2-3. The branches are very straight and the mesh-
work remains regularly developed. The zooecial openings, which are without peristomal
rims, do not project into the fenestrules. The lateral rows of zooecia commonly encroach
onto the narrow dissepiments. The fenestrules are oval or circular on the obverse
surface but commonly appear square, polygonal, or circular on the reverse surface. On
the obverse surface the short, narrow dissepiments are commonly depressed below the
level of the branches.

In shallow tangential sections the zooecial walls have an amalgamate, laminate
microstructure, and the round zooecial openings do not show hemisepta. In very shallow
tangential section bands of small capillaries, 0-01 mm. in diameter, fill the zooecial walls,
weave around the zooecial openings, and extend across the dissepiments (PI. 1 1 , fig. 5).
Tubules, 0-05 to 0-08 mm. in diameter, occur at the junctions of zooecial walls (PI. 11,
fig. 5). Deep tangential sections show hexagonal, basal zooecial sections (PI. 11, fig. 4).

Remarks. Polypora lyndoni sp. nov. is characterized by a delicate, fenestrate meshwork of
slender branches, three rows of zooecia across a branch, small fenestrules, fine capillaries
of 0-01 mm. diameter, and occasional tubules at the junctions of zooecial walls. It
closely resembles P. pertinax Laseron (Crockford 1941, pp. 412-13) from the Eurydesma
cordatum horizon, Allandale, N.S.W. The two species are similar in the number of
branches and fenestrules per 10 mm., in having about the same number of zooecia per
2 mm., and the occasional small tubules at the junctions of zooecial walls. P. lyndoni has

EXPLANATION OF PLATE 10

Figs. 1-9. Stenopora fisheri sp. nov. 1, Tangential section showing arrangement of zooecial openings,
slender zooecial walls, mesopores, and acanthopores, locality ML 109, CPC ML 109-4, x20.
2, 3, External aspect of branching zoarial fragments, locality ML 107, CPC ML 107-7, ML 107-8,
respectively, x 5. 4, Part of tangential section showing zooecial opening with slender zooecial walls
that are penetrated by larger acanthopores and mesopores, locality ML 109, CPC ML 109-4, X 100.
5, Part of longitudinal section in peripheral region showing moniliform walls, locality ML 107,
CPC ML 107-4, X 50. 6, Tangential section showing numerous mesopores and acanthopores around
zooecial openings, locality ML 107, CPC ML 107-4, X 20. 7, Tangential section showing large
acanthopores and mesopores between large zooecial openings, locality ML 107, CPC ML 107^,
X 50. 8, Tangential section showing zooecial openings, prominent acanthopores, and mesopores,
locality ML 109, CPC ML 109-4, X 50. 9, Part of longitudinal section in peripheral region showing
laminate, moniliform zooecial walls, locality ML 107, CPC ML 107-4, x50.

Palaeontology, Vol. 6

PLATE 10

-v**4

r ;■ r

;:Tt.Hl»^,

PHILLIPS ROSS, Lower Permian Bryozoa

J. R. P. PHILLIPS ROSS: LOWER PERMIAN BRYOZOA

79

ra

hJ

m

C

H

80 PALAEONTOLOGY, VOLUME 6

a greater number of zooecial openings per fenestrule, and its dissepiments are wider than
those in P. pertinax.

Polypora lyndoni is similar to P. keppelensis Crockford (1962) (new name for homonym
P. mimita Crockford 1946) from the Lake Creek Beds, Rockhampton, Queensland,
in its fine meshwork, in having three rows of zooecia across each branch, and in its
obverse surface ornamentation. P. lyndoni has wider branches, shorter fenestrules,
generally two or three zooecia per fenestrule, wider dissepiments, and lacks high
peristomes around the zooecial openings.

Polypora lyndoni is similar to P. tripliseriata from the Bitauni Beds of the Permian of
Timor, which are probably equivalent in age to the Lenox Hills Formation in the upper
part of the standard Wolfcampian Series, Glass Mountains, Texas, and upper part of the
Asselian Series, Ural Mountains, Russia (personal communication, C. A. Ross 1962).
The two species are similar in the dimensions of their fenestrules, number of branches
and fenestrules per 10 mm., number of rows of zooecia across a branch, and presence of
capillaries and tubules in the zooecial walls. P. lyndoni has wider branches and fewer
zooecia along a fenestrule.

In general features Polypora lyndoni is similar to a group of species including
P. tubercidifera, P. punctata, P. subvariicellata, and P. nadinae described by Shulga-
Nesterenko (1952) from the Sterlitamak (upper part of the Sakmarian) of the Urals.
However, these species display more highly specialized structures in the zooecial walls
and around the zooecial openings. P. sargaensis and P. kulivoki, described by Trizna
(1948) from the Permian of the Sulva River, apparently also belong to the same group
of species but display more specialized features.

COLLECTION LOCALITIES

ML 90. Approximately 2 miles north-north-east of Round Hill Well, Winning Station, Lyndon River
area. Lat. 23° 21' S., long. 1 14° 41' E. 45 feet below the top of the Lyons Group. Stenopora dickinsi;
S. lyndoni; Polypora lyndoni.

ML 106. Approximately 3 miles north-north-east of Round Hill Well, 1 mile west of Kialawibri Creek
road-crossing. Winning Station, Lyndon River area. Lat. 23°21'S., long. 114°42'E. Upper part of
Lyons Group. Stenopora dickinsi; Polypora lyndoni.

EXPLANATION OF PLATE 1 1

Ligs. 1-3, 6-10. Stenopora lyndoni sp. nov. 1, Tangential section showing zooecial openings separated
by thick zooecial walls that bear numerous large acanthopores and small mesopores, locality ML 90,
CPC ML 90-3, X 50. 2, Part of transverse section showing outer edge of axial region and thick,
laminate zooecial walls in peripheral region, locality ML 90, CPC ML 90-1, X20. 3, Tangential
section showing regular arrangement of large zooecial openings, distinct acanthopores, and small
mesopores, locality ML 90, CPC ML 90-1, X 50. 6, 8, 9, External aspect of branching zoarial frag-
ments, locality ML 90, CPC ML 90-1, ML 90-1, ML 90-3, respectively, X 5. 7, Tangential section
showing laminate microstructure of zooecial walls penetrated by acanthopores and mesopores with
laminate walls, locality ML 90, CPC ML 90-3, X 100. 10, Part of transverse section showing gently
curved laminae of zooecial walls and more steeply inclined laminae of acanthopore walls, locality
ML 90, CPC ML 90-1, X 50.

Ligs. 4, 5. Polypora lyndoni sp. nov. 4, Deep tangential section showing hexagonal form of basal
zooecial sections, locality ML 109, CPC ML 109-11, X20. 5, Tangential section showing round
zooecial openings, fine capillaries and an occasional tubule in the zooecial walls, and small fene-
strules, locality ML 107, CPC ML 107-20, x20.

Palaeontology, Vol. 6

PLATE 11

PHILLIPS ROSS, Lower Permian Bryozoa

J. R. P. PHILLIPS ROSS: LOWER PERMIAN BRYOZOA

81

ML 107. 100 feet west of ML106, Lyndon River area. Lat. 23° 21' S., long. 1 14° 42' E. Uppermost part
of the Lyons Group. Stenopora fislieri; Polypora lyndoni.

ML 109. 410 feet west of ML 106, Lyndon River area. Lat. 23° 21' S., long. 1 14° 42' E. Upper part of
the Lyons Group. Stenopora dickinsi; S. fislieri', Polypora lyndoni.

ML 6. 34 miles west of north of ‘ Moogooree’ Homestead. Lat. 24° S., long. 115° 17' E. Considered to
be located in the lower part of the Lyons Group and in its lowest known marine beds. Stenopora
dickinsi; Polypora lyndoni.

REFERENCES

BASSLER, R. s. 1929. The Permian Bryozoa of Timor. Paldontologie von Timor, 16, pt. 28, 37-90,
23 pi.

COLEMAN, p. J. 1957. Permian Productacea from Western Australia. Bull. Bur. Min. Resour. Aiist.
40, 1-147.

CONDON, M. A. 1954. Progress report on the stratigraphy and structure of the Carnarvon Basin,
Western Australia. Rep. Bur. Min. Resour. Aust. 15, 1-163.

CRESPiN, I. 1958. Permian Foraminifera of Australia. Bull. Bur. Min. Resour. Aust. 48, 1-207.
CROCKEORD, J. 1941. Permian Bryozoa of eastern Australia. Pt. 1 . A description of some previously-
named species of Fenestrellinidae (Fenestellidae). J. roy. Soc. N.S.W. 74, 397-418, pi. 18, 19.

1943. Permian Bryozoa of eastern Australia. Pt. 3. Batostomellidae and Fenestrellinidae from

Queensland, New South Wales, and Tasmania. Ibid. 76, 258-67, pi. 15.

1945. Stenoporids from the Permian of New South Wales and Tasmania. Proc. Linn. Soc.

N.S. W. 70, 9-24, pi. 2, 3.

1946. A bryozoan fauna from the Lake’s Creek Quarry, Rockhampton, Queensland. Ibid. 70,

125-34.

1951. The development of bryozoan faunas in the Upper Palaeozoic of Australia. Ibid. 76,

105-22.

1957. Permian Bryozoa from the Fitzroy Basin, Western Australia. Bull. Bur. Min. Resour.

Aust. 34, 1-134, 21 pi.

1962. Correction of a bryozoan name. J. Paleont. 36, 840.

DiCKiNS, J. M. 1956. Permian pelecypods from the Carnarvon Basin, Western Australia. Bull. Bur.
Min. Resour. Aust. 29, 1-42, 6 pi.

1957. Lower Permian pelecypods and gastropods from the Carnarvon Basin, Western Australia.

Ibid. 41, 1-74, 10 pi.

and THOMAS, g. a. 1959. The marine fauna of the Lyons Group and the Carrandibby Formation

of the Carnarvon Basin, Western Australia. Rep. Bur. Min. Resour. Aust. 38, 65-96.

DUNCAN, H. 1949. Genotypes of some Paleozoic Bryozoa. J. Wash. Acad. Sci. 39, 122-36.

LONSDALE, w. 1844. Description of six species of corals from the Palaeozoic Formation of Van
Diemen's Land. In darwin, Charles, Geologic Observations on the volcanic islands, visited during
the voyage of the H.M.S. Beagle, together with some brief notices on the geology of Australia and the
Cape of Good Hope, Appendix, 161-9. London.

1845fl. Polyparia. In strzelecki, p. e. de. Physical Description of New South Wales and Van

Diemen's Land, 262-8, pi. 8, 9. London.

18456. Description of some characteristic Palaeozoic corals of Russia. In murchison, r. l,

VERNEUiL, E., and KEYSERLING, A. The Geology of Russia in Europe and the Ural Mountains, 591-634,
pi. A. London.

M'COY, F. 1844. A synopsis of the characters of the Carboniferous Limestone Fossils of Ireland. 207 pp.,
29 pi. Dublin.

MURCHISON, R. I. and VERNEUIL, E. 1844. Note sur les equivalents du systeme permien en Europe,
suivie d’un coup d’oeil general sur I’ensemble de ses fossiles et d’un tableau des especes. Bull. Soc.
geol.fr., ser. 1, 1, 475-517.

prendergast, K. L. 1943. Permian Productinae and Strophalosiinae of Western Australia. J. roy.
Soc. W. Aust. 28, 1-61.

ROSS, J. R. p. and ROSS, c. a. 1962. Faunas and correlations of the late Paleozoic rocks of northeast
Greenland. Pt. 4, Bryozoa. Medd. Gronland, 167 (7), 1-65, 18 pi.

C 1015 G

82 PALAEONTOLOGY, VOLUME 6

SHULGA-NESTERENKO, M. I. 1952. LowcE Permian Bryozoa from the area of the Urals. Trud. Inst.

paleont. Akad. nauk SSSR, 37, 1-84, 16 pi. (In Russian.)

TEiCHERT, c. 1941. Upper Paleozoic of Western Australia: correlation and paleogeography. Bull.

Amer. Ass. Petrol. Geol. 25, 371-415.

^1947. Stratigraphy of Western Australia. Ibid. 31, 1-70.

THOMAS, G. A. 1958. Permian Orthotetacia of Western Australia. Bull. Bur. Min. Resour. Aiist. 39,
1-115.

TRizNA, V. B. 1948. Permian Bryozoa from the Sulva River. Trud. vsesoyuz nauch.-issled. Inst. Geol. 31,
137-88, 20 pi. (In Russian.)

JUNE R. P. PHILLIPS ROSS

Illinois State Geological Survey,

Manuscript received 21 March 1962 Urbana, Illinois, U.S.A.

ON THE INTERPRETATION AND STATUS OF
SOME HYSTRICHOSPHERE GENERA

by c. DOWNiE and w. A. s. sarjeant

Abstract. The taxonomic status of the genera Micrhystridium, Baltisphaeridium, Veryhachiimi, Midtiplici-
sphaeridiiim, Leiospliaeridia, Leiosphaeridium, Frotoleiosphaeridium, and Archaeobystrichosphaeridium is ex-
amined. The generic names Midtiplidsphaeridium, Leiosphaeridium, and Protoleiospbaeridium are rejected : the
genera Micrhystridium, Baltisphaeridium, Leiospliaeridia, and Veryhachium are redefined : and lists are given of
the species now considered to belong to these genera, the generic attribution of 140 species being changed.

In his recent paper describing assemblages of Devonian microplankton from Alberta,
Canada, F. L. Staplin (1961) has made a number of proposals relating to the taxonomy
of the hystrichospheres which we consider not acceptable. These proposals have been
briefly criticized by Eisenack (1962); they are here discussed fully and the reasons for
their rejection are outlined. The status of the genera concerned and of some related
genera is examined and the attribution of species to these genera is clarified.

1 . Micrhystridium — Baltisphaeridium

Deflandre (1937) erected the genus Micrhystridium to include all globular hystricho-
spheres having a shell diameter generally less than 20 p, regardless of the character of
their appendages : he designated as type species Micrhystridium inconspicuum (Deflandre
1935), a form having thorn-like processes with closed tips. The distinction between this
genus and the genus Hystrichosphaeridium, erected in the same paper, lay only in
dimensions, Hystrichosphaeridium then comprising all forms whose shells were not
differentiated into fields by sutures, again regardless of appendage character. In 1958
this latter genus was split by Eisenack, who, retaining the nscme. Hystrichosphaeridium for
forms which, like the type H. tubiferum (Ehrenberg 1838) Deflandre 1937, possess tubular
spines open distally, erected a new genus, Baltisphaeridium, for forms having spines
closed distally, the type species being B. longispinosum (Eisenack 1931). No alteration
was made to the diagnosis of Micrhystridium.

Staplin proposes the abolition of the upper size limit for Micrhystridium and a re-
definition of the genus on the basis of the spine character of its type species, causing it to
correspond exactly with Baltisphaeridium. On this basis he proposes abandonment of the
latter genus and the reattribution of its species to Micrhystridium.

The hystrichospheres having spines closed distally exhibit an overall range in shell
diameters from 5 ju, to 240 p. Consideration of the size ranges of individual species
reveals two major groupings, two foci of morphology: this has been graphically repre-
sented by Downie (1958, p. 334) in an analysis of a Tremadocian hystrichosphere
assemblage. The boundary between these two major groupings has been accepted by most
workers as being sensibly drawn at 20 p. While species exist which overlap this boun-
dary from either side, the number of species having a size mode between 20 and 25 p is
very small: of 204 Palaeozoic species and infraspecific groups attributable to the genera

[Palaeontology, Vol. 6, Part 1, 1963, pp. 83-96.]

84

PALAEONTOLOGY, VOLUME 6

Micrhystridium and Baltisphaeridium (186 species and 18 infraspecies), only 10 overlap
the boundary and only 1, B. ramusculosum (Deflandre 1942), has a size mode actually on
the boundary. Of 164 Mesozoic and Tertiary species and infraspecific groups attributable
to the genera Micrhystridium and Baltisphaeridium (117 species and 46 infraspecies and
forms related to named species), only 8 overlap the boundary and only one, M. piUferum
(Deflandre 1937), apparently has a size mode actually on the boundary; this species is
based on a single specimen. (Size ranges are quoted in detail in the accompanying
systematic section.) The few forms transgressing the size boundary separating the
genera may be regarded as extremes in series surrounding morphological foci: trans-
gressions of the boundary are usually slight and by extremes in the size range of a parti-
cular species whose mode is clearly below or above the boundary.

The forms having a shell diameter smaller than 20 p, the micrhystridia, appear at
many horizons in the Upper Palaeozoic and Mesozoic (Carboniferous, Permian, Trias-
sic, and Jurassic) to correspond in their concentration to differences in environment,
tending to be either very abundant or extremely few in number in particular assemblages.

Although Staplin’s emendation of the genus Micrhystridium may be legally valid in
terms of the existing Codes of Nomenclature, his proposal to remove from the diagnosis
the one distinguishing characteristic specified by its author renders the genus meaning-
less : to do so would be to make it synonymous with Hystrichosphaeridium as defined at
the time of publication of the two generic diagnoses. We consider that the genera
Micrhystridium and Baltisphaeridium express two natural size modes, and that the size
boundary separating them should be retained at least until more compelling evidence for
emendation of the genera is available.

We feel, however, that the genus Micrhystridium, as at present defined, is rather
heterogeneous, since it comprises not only forms having processes with simple tests, not
divided into fields by crests, and with simple spines closed distally, but also forms whose
shells are divided into fields and others that have processes open distally. We consider
with Staplin that it should be restricted to forms having spines with closed tips: that
forms having shells whose surfaces are divided into fields by crests should be transferred
to the genus Cymatiosphaera and forms having tubular processes to the genus Hystri-
chosphaeridium. If in the future it transpires that within the two latter genera there exists
a similar division into natural size modes, new taxa may then be erected to express this
difference.

2. Veryhachium

The name Veryhachium was introduced in 1954, being applied to a species, V. trisulcum,
which was not at that time validly published. This was subsequently rectified in 1958 and
the genus should date from then. The genus was emended by implication by one of us
(Downie 1959) when forms with an ellipsoidal test and a single spine were removed to
a new genus, Deunffia. Veryhachium then included hystrichospheres with polygonal tests
and a small number of spines. In recent publications (Eisenack 1959, Staplin 1961),
markedly contrasted interpretations of the genus have been evident and we consider
a clarification of the situation necessary.

It is evident from recent publications (Eisenack 1959, Jekhowsky 1961) that mor-
phological transitions exist between Veryhachium and both Micrhystridium and Balti-
phaeridium. Eisenack (p. 207) recognizes these difficulties and prefers to retain in

C. DOWNIE, W. A. S. SARJEANT: SOME H YSTRICHOSPHERE GENERA 85

Baltisphaeridium any polygonal form which, by other morphological criteria, is linked
to typically round-bodied forms, e.g. B. oligospinosum and B. visbyense, which accord
in all respects with the diagnosis of Veryhachium. He rightly states that the generic
boundary would otherwise separate forms that are closely related.

The principle that Eisenack is enunciating, that taxonomic units should express natural
relationships, is one with which the authors are in general agreement. However, rigid
application of this principle to the hystrichospheres would result in the shrivelling or
disappearance of most existing genera, with a resultant vast increase of species within the
few remaining (Micrhystridium emend. Staplin would, for example, contain about 800
species). It would, moreover, produce a false picture of the real situation where, although
overlap and gradation may occur at the fringes, there do exist morphological groupings
with intermediates considerably rarer than typical forms. Even Eisenack does not apply
his principle rigorously, for he has several times emphasized the existence of continuous
gradation between leiospheres and hystrichospheres, and yet separates them into
distinct genera.

We consider that the generic name Veryhachium should be applied to forms having
a low number (generally 3-8) of hollow spines arising from a polygonal or subpolygonal
test. The combination of shell shape and spine number is considered to define the genus,
and the views of Staplin, that all species having a polygonal test should be placed within
this genus, regardless of spine number, are rejected as contrary to the original intention
of Deunlf.

3. Diornatosphaera

The status of this genus and of related genera erected by Timofiev (1959) has been
recently reviewed by Deflandre and Deflandre-Rigaud (1961). The decisions of these
authors are accepted in their entirety, and their paper is welcomed for its clarification and
regularization of a potentially difficult taxonomic situation.

4. MultipUcisphaeridium

This proposed new genus (Staplin 1961) is distinguished from Baltisphaeridium and
Micrhystridium in the character of the spine tips: forms having unmodified spine tips are
referred to the two latter genera, while forms having multifurcate, expanded, or dis-
sected spine tips are referred to MultipUcisphaeridium. The number of species that would
be referable to this latter genus in the Palaeozoic is not large (20 species, at present
allocated to the genera Micrhystridium, Baltisphaeridium, and Archaeohystrichosphaeri-
dium), but in the Mesozoic and Tertiary it is considerable.

MultipUcisphaeridium is considered morphologically ill-founded for the following
reasons :

(а) While many species do have entirely simple or entirely complex spine tips, there
are a number of species whose representatives consistently have both simple and furcate,
or complex, spine tips.

(б) In some individual species, the normal range of variation includes specimens with
only simple spines, specimens with only furcate, &c., spines, and specimens with spines
of mixed types.

(c) The species having in common the character of furcate, &c., spine tips show no

86

PALAEONTOLOGY, VOLUME 6

distinctive combinations of other characters to suggest that they constitute a morpho-
logically separate entity. Examination of assemblages at the same or earlier horizons
frequently discloses a higher degree of morphological similarity to species with un-
branched spines than to other species of Multiplicisphaeridium. There is no evidence for
the view that Multiplicisphaeridium constitutes an evolutionary chain of species ; on the
contrary, there is much evidence that the species with furcate or complex spine tips
repeatedly arose, by a trend of increasing spine complexity, from forms with simple
spines.

{d) When combinations of other characters are brought into account, there are seen
to be a number of groupings within Baltisphaeridium as here defined; species within
these groupings may have simple, furcate, or complex spines. Separation of forms having
spines of the two latter types into a distinct genus would cut across these groupings and
would obscure natural relationships.

In the Palaeozoic, a series of increasing spine complexity is clearly shown in two cases :
Baltisphaeridium polygonale (Eisenack 1931)-^5. digitatum (Eisenack 1938); Ordovician.
B. oligofurcatum (Eisenack 1954)— >5. meson (Eisenack 1955)— >5. brevifurcatum (Eise-
nack 1954); Silurian. Other Palaeozoic species having intermediate characters are listed
below.

(a) Species having both simple spines and knobbed, bi- or trifurcate spines.

ORDOVICIAN. Baltisphaeridium longispinosum (Eisenack 1931) Eisenack 1958 ; B. nuda-
tum (Timofiev 1959) comb, nov; B. ordovicum (Timofiev 1959) comb, nov; B. rjabinini
Timofiev 1959) comb. nov.

SILURIAN. B. ramusculosum (Deflandre 1942), Downie 1958.

DEVONIAN. B. cf. digitatum (Deunff 1954) comb. nov.

(h) Species having only briefly furcate spines.

ORDOVICIAN. B. circumscissum (Timofiev 1959) comb, nov; B. setigerfurcatum
(Timofiev 1959) comb. nov.

DEVONIAN. B. lewisi (Deuntf 1954) comb, nov.; B. microfurcatum Deunff 1957.

A large number of Mesozoic and Tertiary species show intermediate characteristics:

{a) Species having both simple spines and knobbed, bi- or trifurcate spines.

JURASSIC. Baltisphaeridium polytrichum sensu Downie 1957 non Valensi 1947;
B. varispinosum Sarjeant 1959; B. cf. multifurcatum (Deflandre 1947), Klement 1960;
B. lumectum Sarjeant 1960«; B. cf. fimbriatum (White 1842), Sarjeant 1960a.

CRETACEOUS. B. multifurcatum (Deflandre 1937) Klement 1960; B. pseudhystri-
chodinium (Deflandre 1937) comb, nov.; B. striolatum (Deflandre 1937) comb, nov.;
B. cf. pseudhystrichodinium (Deflandre 1937), Lejeune-Carpentier 1941 ; B. longifiircatum
(Firtion 1952) comb, nov.; B. cf. striolatum (Deflandre 1937), W. Wetzel 1952; B. oli-
gacanthurn (W. Wetzel 1952) comb. nov. and its subspecies.

EOCENE. Micrhystridium cf. stellatum Deflandre 1942, Deflandre and Cookson 1955;
Baltisphaeridium geometricum (Pastiels 1948) comb. nov.

OLIGOCENE. B. cf. pseudhystrichodinium (Deflandre 1937), Gocht 1952.

C. DOWNIE, W. A. S. SARJEANT: SOME HYSTRICHOSPHERE GENERA 87

(b) Species having simple spines together with others of complex form.

JURASSIC. Baltisphaeridium vestitum (Deflandre 1938) Sarjeant 19606.

CRETACEOUS. B. flosculus 1937) comb. nov. ; B. horridimi (Deflandre 1937)

comb, nov.; B. heteracanthum (Deflandre and Cookson 1955) comb, nov.; B. tridacty-
lites (Valensi 1955) comb. nov.

(c) Species having capitate (c.), knobbed (k.), or only briefly furcate (f.) spines.

JURASSIC. Baltisphaeridium cf. intermedium (O. Wetzel 1933), Deflandre 1938 (f.);

Micrhystridium bigoti Deflandre 1947 (f.); M. variabile Valensi 1953 (c. or f.); B. downiei
Sarjeant 1960« (c., k., or f.); B. alf. fimbriatum (White 1842), Sarjeant 19606 (f.); B. tri-
buliferum Sarjeant 19626 (f.); M. rhopalicum Sarjeant 19626 (k. or f.).

CRETACEOUS. Baltisphaeridium fimbriatum (White 1842), Sarjeant 19606(h); 5. inter-
medium (O. Wetzel 1933) comb. nov. (f.); B. alf. multifurcatum (Deflandre 1937),
Firtion 1952; B. rhabdophorum (Valensi 1955) comb. nov. (f.); B. huguonioti (Valensi
1955) comb. nov. (f.).

EOCENE. B. colligerum (Deflandre and Cookson 1955) comb. nov. (c. or f.).

OLIGOCENE. B. longofilum (Maier 1959) comb. nov. (f).; B. echinoides (Maier 1959)
comb. nov. (c.).

There are thus a considerable number of species whose spine characters indicate a
morphogenetical position between forms having wholly simple spines and forms having
multifurcate, expanded, or dissected spines all of one type. The use of the structure of
spine tips cannot be considered a satisfactory single criterion for determining generic
assignment ; the genus Multiplicisphaeridium would, if accepted, be an agglomeration of
end members of numerous evolutionary trends rather than the focus of a morphological
grouping. All, or nearly all, existing genera have been established in the view that they
have genetic meaning and express natural groupings : the discovery of overlap with other
genera at the extreme limits of their morphological spread does not necessarily invalidate
this view. In the case of Multiplicisphaeridium, such a view cannot be held even at the
inception. We propose therefore that this genus be abandoned and that the present
distribution into genera of the species comprising it be retained.

5. The leio spheres

The generic name Leiosphaeridium Timofiev apparently first appeared in a caption to
an illustration (1956) and was not therefore validly published until 1959, when a descrip-
tion was given. (Figures unaccompanied by diagnoses do not constitute valid publication
either under the Botanical or Zoological Rules at this date.) It was intended as an altera-
tion of Leiosphaera Eisenack (1938) and as such it is invalid. The type species of the
genus Leiosphaera, L. solida, was recognized by Eisenack (1958) as conspecific with
Tasmanites huronensis (Dawson 1871); he thus abandoned the genus, attributing to
Tasmanites such species as accord with its diagnosis and setting up a new genus,
Leiosphaeridia, to accommodate forms not attributable to Tasmanites. The name Leio-
sphaeridium was proposed as an amendment of Leiosphaera and presumably Timofiev
regarded L. solida as its type species; Leiosphaeridium is thus a junior synonym of
Leiosphaera and hence of Tasmanites. If the single species attributed to Leiosphaeridium
by Timofiev were considered as its type, its characters are such as to render it attributable

88

PALAEONTOLOGY, VOLUME 6

to Leiosphaeridia, of which the genus then beeomes a junior synonym. Whichever view-
point is taken, the name Leiosphaeridium must be abandoned. It is clear that at the time
of writing, Timofiev was not aware of the existence of the genus Leiosphaeridia.

The genus Protoleiosphaeridiwn Timofiev was likewise not validly published until 1959
and its type species, P. conglutinatum, was not so designated until 1960. The size ranges
quoted for this genus and for Leiosphaeridium are 20-30 p and 200-300 p : these size
ranges are confusing, since the six species attributed by Timofiev to the former genus
have a range 20-50 p and the single species attributed to the latter genus has a size range
of 100-125 /X.

While the name Leiosphaeridium cannot be retained, whether in the original meaning
of Timofiev or under the amended diagnosis of Staplin (rejected names cannot be re-used
under either the Zoological or the Botanieal Rules), Protoleiosphaeridium might be
retained as a subgenus of Leiosphaeridia and restrieted to thin-walled forms having a
diameter less than 50 p. The type of Eisenack’s genus has a size range 80-140 p: how-
ever, the total range size of all species attributed to this genus is 23-1 50 /.i and there is
no indication of any division into size modes. We feel that Protoleiosphaeridium should
be treated as a synonym of Leiosphaeridia and rejected, and that its species should be
reattributed to the latter genus.

The statement in Eisenack’s diagnosis that a pylome exists in Leiosphaeridia is to be
interpreted as indicating that this is a structure present in some species but not in all. He
does not even consider possession of a pylome to be a factor of specific value (Eisenack,
in lift.).

Timofiev (1959) also established seven other genera in the family Leiosphaeridiaceae.
Of these, Vavososphaeridium (reticulate ornament), Orycmatosphaeridium {pitted surface),
Lophosphaeridium (tubercular ornament), and Trachysphaeridium (shagreen ornament)
might be considered subgenera of Leiosphaeridia based on dilferent shell ornament. Some
of the speeies listed under Leiosphaeridia could probably be attributed to one or other of
these subgenera: however, without examination of type material, the authors do not feel
competent to make such reallocation. Timofiev’s other genera Zonosphaeridium (thick
walls), Trematosphaeridium (perforate test), and Symplassosphaeridium (clusters of vesi-
cles) seem either to be the product of accidents of preservation or to be quite different
kinds of structure from the typical leiospheres: judgement on them is, for the present,
reserved, although the considerable resemblance between Zonosphaeridium (thick walls:
? = Tasmanites) and the genus Tasmanites may be noted.

6. Archaeohystrichosphaeridium

Staplin does not consider this genus of Timofiev (1959) in his taxonomic review. The
genus was defined as follows: ‘Vesiele with diameter 10-35 /x, thin, ornamented with
simple, blunt, or pointed spines, sometimes forking on the periphery. Colour pale
yellow.’

The range of diameters quoted by Timofiev is in faet 18-70 /x: only ten of seventy-nine
species so far attributed to it fail to exceed 35 p. No type species has yet been designated,
so that the genus has not yet been validly published. As at present defined, the genus is
indistinguishable from Baltisphaeridium Eisenack and may be by intent a synonym of
that genus. The thinness of the shell walls might afford a basis for differentiation at
subgeneric level. The authors have not reattributed the species of Archaeohystricho-

C. DOWNIE, W. A. S. SARJEANT: SOME H YSTRICHOSPHERE GENERA

89

sphaeridium here, since they hope that a validation and redefinition of the genus may be
forthcoming.

SYSTEMATIC SECTION

Evitt (1961) has recently drawn together evidence concerning the relationships of
the hystrichospheres and has demonstrated the presence in many genera and species,
though by no means in all, of structures indicating relationship to the dinoflagellates.
In particular he has shown that the genus Hystrichosphaera, on which the Order
Hystrichosphaeridea is based, is attributable to the Dinophyceae. A reclassification of
the hystrichospheres, following this work, is at present in preparation ; for the present,
classification is made to generic level only. Known range of shell diameters is quoted
for the species of Micrhystridiwn and BaJti sphaeridium: where species are reallocated
to these genera, they were previously in Hystrichosphaeridium unless otherwise stated.

The authors would like to emphasize that species have been attributed to the various
genera on the basis of published descriptions and allocations, and that the inclusion of
a species name does not necessarily imply endorsement of its status. Some of the species
included differ morphologically only slightly, if at all, from others previously described ;
others, e.g. those of Sannemann (1955) here attributed to Baltisphaeridium, probably
constitute distinct genera whose definition is not possible on present knowledge.

Genus baltisphaeridium Eisenack 1958, emend.

Emended diagnosis. Hystrichospheres with spherical to oval shells not divided into fields
or plates, bearing i numerous processes, simple, branching or ramifying, hollow to
solid, always with closed tips. The processes are not connected together distally and no
outer shell, complete or incomplete, is present : the processes are most often of a single
basic type, but processes of two or more types may be present. Mean and modal dia-
meter of shell greater than 20 jtt.

Discussion. While the spines of species attributed to Baltisphaeridium and Micrhystri-
dium are most often hollow, some or all spines of particular species may be partly or
wholly solid. The hollowness or otherwise of the spines is extremely difficult to deter-
mine in many instances, particularly in species of Micrhystridiwn. On a single specimen,
spines may be in part hollow, in part solid; Evitt {in litt.) notes that this is the case in B.
ferox and in B. floscidus. Species occur whose spines are hollow throughout their length
and closed only at the tip; others with spines solid for most of their length, but with a
cavity at the base. Sannemann (1955) has described a number of species in which there is
a solid central ‘spire’ within the spine cavity. The importance of distinguishing between
these alternatives is doubtful; for the present, no such distinction is made.

Type species. Baltisphaeridium loiigispinosum (Eisenack 1931) Eisenack 1958 (40-15 p); Cambrian-
Ordovician.

Other species (Palaeozoic). Baltisphaeridium aculeatum (Timofiev 1959) comb. nov. (70-90 /x) ; B. alloiteaui
(Deunff 1955) comb. nov. (formerly Micrhystridium) (25-30 /x); B. amiidatum (Timofiev 1959) comb,
nov. (93-96 p) \ B. apiculatimi (Timofiev 1959) comb. nov. (50-60 p); B. arrectum (Timofiev 1959) comb,
nov. (70-100 /x).

B. bimargiuatum (Timofiev 1959) comb. nov. (60-80 p); B. bohemicum (Eisenack 1934) comb. nov.
(120 /x); B. brevifurcatum (Eis. 1954) comb nov. (60 p); B. brevispiuosum (Eis. 1931) Eisenack 1958

90

PALAEONTOLOGY, VOLUME 6

(50-70 /x); B. brevispinosiim (Eis. 1931) subsp. callosum Sannemann 1955 (200 yx); B. brevispinosum
(Eis. 1931) subsp. castaneioides Sannemann 1955 (200 p)', B. brevispinosum (Eis. 1931) var. granulifera
Downie 1959 (16-32 ;u.); B. brevispinosum (Eis. 1931) var. nanum Deflandre 1942 (25-26 /x); B. brevi-
spinosimi (Eis. 1931) var. wenlockensis Downie 1959 (16-32 /x); B. breviciliatum (Staplin 1961) comb,
nov. (formerly Micrhystridium) (32-37 p).

B. castaneum (Eisenack 1934) comb. nov. (150-160 p)\ B. circumscissurn (Timofiev 1959) comb. nov.
(70-90 jLx); B. cognition (Timofiev 1959) comb. nov. (50-100 jix); B. coniferum (Sannemann 1955) comb,
nov. (340/x); B. conspicuum (Timofiev 1959) comb. nov. (106-109 |tx); B. corallinum Eisenack 1959
(70-95 /x); B. corollatum (Timofiev 1959) comb. nov. (91-99 p)\ B. crassiechinatum (Staplin 1961) comb,
nov. (formerly Micrhystridium) (24-33 p) \ B. cristatiim (Downie 1958) Eisenack 1958 (23-30 /x).

B. differtum (Sannemann 1955) comb. nov. (250 p); B. digitatum (Eisenack 1938) Eisenack 1959 (65-
75 p) \ B. dignum (Sannemann 1955) comb. nov. (150 /x); B. diploporum (Eis. 1951) comb. nov. (formerly
Micrhystridium) 35^0 p).

B. cc/n'«oj//»z(Staplin 1961)comb. nov. (formerly Micrhystridium) (17-22 /x); B. eisenackium (Deunff
1958) comb. nov. (60 p)\ B. eisenackium (Deunff 1958) var. crozonensis Deunff 1958 (25 p)\ B. eisenacki
(Sannemann 1955) comb. nov. (125 /x); B. eoplanktonicum (Eisenack 1955) Downie 1959 (20 p);
B. erraticum (Eis. 1954) comb. nov. (35-50 jix); B. franconium (Sannemann 1955) comb. nov. (200 p);
B. gotlandicum (Eis. 1954) comb. nov. (46 p).

B. hippocrepicum (Timofiev 1959) comb. nov. (69-78 yx) ; B. hirsutoides (EiSQndiCk. 1939 emend. Eisenack
1951) Eisenack 1958 (20-50 p)\ B. hirsutoides (Eis. 1939 emend. Eis. 1951) var. hamatum Downie 1958
(18-40 yx); B. hymenoferum (Eis. 1938) Eisenack 1958 (65 yx); B. hystrichoreticulatum (Eis. 1938) comb,
nov. (40-45 /x); B. inconspicuum (Timofiev 1959) comb. nov. (45-65 yx); B. integrum (Sannemann 1955)
comb. nov. (300 yx).

B. lewisi (Deunff 1954) comb. nov. (45 yx); B. longispinosoides (Sannemann 1955) comb. nov. (120-
200 yx); B. longispinosum (Eisenack 1931) forma filifera Eisenack 1959 (40-75 yx); B. longispinosum (Eis.
1931) forma latiradiata Eisenack 1959 (40-75 yx); B. longispinosum (Eis. 1931) forma robusta Eisenack
1959 (40-75 yx); B. lophophorum Eisenack 1959 (30 yx); B. lucidum (Deunff 1958) comb. nov. (80-89 yx).

B. inacropylum'EismdLck 1959 (60-68 yx); B. meson (Eis. 1955) comb. nov. (54-62 yx); B. micracan-
thum Eisenack 1959 (68-88 yx); B. microfurcatum (Deunff 1957) comb. nov. (22 p); B. microspinosum
(Eis. 1954) Downie 1959 (58 yx); B. miiltipilosum (Eis. 1931) Eisenack 1958 (45-60 yx); B. midtipilosum
(Eis. 1931) subsp. validum Sannemann 1955 (130-180 yx); B. mutabile (Sannemann 1955) comb. nov.
(150 yx); B. nudatum (Timofiev 1959) comb. nov. (66-69 p).

B. octospinum (Staplin 1961) comb. nov. (formerly Micrhystridium) (24— 33yx); B. oligofurcatum
(Eisenack 1954) comb. nov. (56 p); B. ordovicum (Timofiev 1959) comb. nov. (80-84 yx); B. pateiim
(Timofiev 1959) comb. nov. (60-100 yx); B. paucispinum (Deunff 1954) comb. nov. (formerly Very-
hachium) (24-28 yx); B. piriferum (Eis. 1954) comb. nov. (62 yx); B. polygonale (Eis. 1931) Eisenack 1959
(75-230 yx); B. quadriradiatum (Timofiev 1959) comb. nov. (93-106 yx).

B. ramispinosum (Staplin 1961) comb. nov. (formerly Multiplicisphaeridium) (19-29 yx); B. ramuscu-
losum (Deflandre 1942) Downie 1959 (18-23 yx); B. ramusculosum (Defl. 1942) var. macrocladum
Deunff 1955 (30 yx); B. rigens (Timofiev 1959) comb. nov. (90-100 yx); B. rjabinini (Timofiev 1959)
comb. nov. (58-70 yx); B. robustispinosum Downie 1959 (30 yx); B. robiistum (Sannemann 1955) comb,
nov. (240 yx); B. robustum (Sannemann 1955) s\xhs>y>. fission Sannemann 1955 (15 yx).

B. sanneoianni ( Deunff 1958) comb. nov. (formerly Micrhystridium (24 p); B. sedecimspinosum
(Staplin 1961) comb. nov. (formerly Veryhachium) (19-25 yx); B. sericum (Deunff 1954) comb. nov.
(formerly Micrhystridium) (23-26 yx); B. setigerfurcatum (Timofiev 1959) comb. nov. (81-84 yx); B.
sexradiatum (Timofiev 1959) comb. nov. (55-60 yx); B. spicatum (Staplin 1959) comb. nov. (formerly
Multiplicisphaeridium) (63-72yx); B. spiciferum (Deunff 1955) comb. nov. (23-31 yx); B.spiniscens (Timo-
fiev 1959) comb. nov. (80-1 10 yx); B. spinoglobosum (^tayAm 1961) comb. nov. (formerly Micrhystri-
dium) (17-24 yx); B. sprucegrovensis (Staplin 1961) comb. nov. (formerly ? Multiplicisphaeridium)
(35-44 yx); B. stellaeforoie (Timofiev 1959) comb. nov. (88-99 yx); B. sueciciim Eisenack 1959 (50-67 yx).

B. tiniofeevi Deunff 1961 (25-30 yx); B. trifurcation (Eisenack 1931) Eisenack 1958 (50-70 yx); B.
trifurcation (Em. 1931)forma breviradiataExsQnac\c\959 (70 p); B. trifurcatum (Eis. 1931) forma
rar//x7?a Eisenack 1959(45yx); B. trifurcation (Eis. 1931) forma Eisenack 1959(52 yx); B. trifurcation
(Eis. 1931) forma paucifurcatum Eisenack 1959 (37-52 p); B. trifurcatum (Eis. 1931) subsp. consoniim
Sannemann 1955 (150yx); B. trifurcatum subsp. procerum Sannemann 1955 (120 yx); B. triplicativum

C. DOWNIE, W. A. S. SARJEANT: SOME H YSTRICHOSPHERE GENERA 91

(Timofiev 1959) comb. nov. (62-73 /x); B. tnincatum (Staplin 1961) comb. nov. (formerly Multipli-
dsphaeridium) (28-38 /x); B. tuberatum (Downie 1958) comb. nov. (15-30 p,)', B. tuberosum (Sannemann
1955) comb. nov. (125 /x).

B. venustum (Sannemann 1955) comb. nov. (150^i); B. vigivitispimim (Staplin 1961) comb. nov.
(formerly Micrhystridium) (24 /x); B. zouale (Timofiev 1959) comb. nov. (93-106 /x).

Remarks. The seventy-nine species attributed XoArchaeohystrichosphaeridiumhyTimo^QV
(1959) have not been reattributed to Baltisphaeridiiim here (although all are capable of
such reattribution) in view of the possibility of a redefinition of the genus. Sannemann
(1955) refers to a species, Hysirichosphaeridium saalfieldensis Eisenack, of which no
description could be located: this species is therefore omitted from consideration. The
species Baltisphaeridium lewisi, B. paucispinosum, and B. sericwn were published as
names accompanying figures (Deunff 1956): since no text descriptions have been yet
published, their validity is doubtful.

Other species {Post-Palaeozoic). Baltisphaeridium armatum (Dellandre 1937) comb. nov. (25 X 18-20 p,)\
B. asteroideum (Maslov 1956) comb. nov. (20-28 /x); B. biformoides (Eisenack 1954) comb. nov. (50-
67 /x); B. brevispi/iosiim Pastiels 1948 non Eisenack 1931 (40-55 fxx 22-25 /x).

B. centrocarpum (Dellandre and Cookson 1955) Gerlach 1961 (54-80 /x); B. claviculorum (Deflandre

1938) comb. nov. (30 /x); B. claviferum (Wilkinson 1849) comb. nov. (c. 100 /x); B. clavigerum (Deflandre
1937) comb. nov. (c. 45 /x); B. colligerum (Deflandre and Cookson 1955) comb. nov. (33 /x).

B. danicum (W. Wetzel 1952) comb. nov. (36-12 p,); B. deusicomatum (Maier 1959) Gerlach 1961
(53-95 X 86-88 /lx); B. dictyophorum (Cookson and Eisenack 1958) comb. nov. (lip); B. diffbrme
(Pritchard 1845) comb. nov. (c. 30 p); B. divergeus (Eisenack 1954) comb. nov. (52-83 /ix); B. dowuiei
Sarjeant 1960t? (32-40 X 38-46 /x); B. echiuoides (Maier 1959) comb. nov. ( 70-80 /x); B. ehrenbergi
(Deflandre 1947) Sarjeant 1961 (42-46 /x).

B. ferox (Deflandre 1937) comb. nov. (46-120/16); B. fimbriatum (White 1842) Sarjeant 1959 (31-
77 /x); B. c(. fimbriatum (White 1842), Valensi 1955 (34-35 p) \ B. cf. fimbriatum (White 1842), Sarjeant
1960(7 (dimensions not obtained); B. flosculus (Deflandre 1937) comb. nov. (38-45 /x); B. geometricum
(Pastiels 1 948) comb. nov. (44-60 p) ; B. gilsouii (Kutferath 1 950) comb. nov. (formerly Micrhystridium)
(30 /x); B. gramdosum (Deflandre 1937) Sarjeant 1962a (33-55 X 18-26 /lx).

B. heteracanthum (Deflandre and Cookson 1955) comb. nov. (56-107 p); B. cf. hirsutoides (Eisenack

1939) Gocht 1952 (c. 80 /u.); B. hirsutum (Ehrenberg 1838) comb. nov. (35-46 /ix); B. hirsutiim (Ehr.
1838) forma minor O. Wetzel 1933 (16-28 /lx); B. hirsutum (Ehr. 1838) forma various O. Wetzel 1933
{35-16x25-60 p)\ B. hirsutum subsp. arnplum W. Wetzel 1955 {96 p): B. cf. hirsutum (Ehr. 1838)
Pastiels 1948 (35-40 p) ,B. c{. hirsutum (Ehr. 1838) Cookson, 1953 (26-44 /x); B. cf. hirsutum (Ehr. 1838),
Cookson and Eisenack 1958 (dimensions not stated); B. horridum (Deflandre 1937) comb. nov. (55-
57 /lx); B. huguouioti (Valensi 1955) comb. nov. {29-33x25-29 p).

B. iaculigerum Klement 1960 {29-35 p)\ B. intermedium (O. Wetzel 1933) comb. nov. (dimensions
not quoted); B. cf. intermedium (O. Wetzel 1933) Deflandre 1938 (30-35 /lx); B. inusitatum Klement 1960
{52 p); B. lobospinosum (Gocht in Weiler 1956) comb. nov. (90-95 /x); B. longifurcatum (Firtion 1952)
comb. nov. (40-46 /x); B. longofilum (Maier 1959) comb. nov. {59 p)\ B. lumectum Sarjeant 1960a
(45-65 p).

B. machaerophorum (Deflandre and Cookson 1955) Gerlach 1961 (41-54 /tx); B. maHeoferum (White
1842) comb. nov. (63-85 /u.); B. mariannae (Philippot 1949) comb. nov. {c. 40 p); B. mixtispincsiim
Klement 1960 (46-60 X 37-52 /lx); B. nndtifurcatum (Deflandre 1937) Klement 1960 {45-65 p); B. cf.
midtifurcatum (Defl. 1937), Firtion 1952 (40-60 /lx); B. cf. nndtifurcatum (Defl. 1937), Klement 1960,
(58-63 p); B. neptuni Eisenack 1958 (40-60 /lx).

B. oligacanthum (W. Wetzel 1952) comb. nov. (mean 12 p); B. oligacanthum (W. Wet.) var., W.
Wetzel 1952 (90 p); B. oligacanthum (W. Wet. 1952) subsp. complanatum W. Wetzel 1952 (66-12 p);
B. oligacanthum (W. Wet.) subsp. grandulatum W. Wetzel 1952 (60 p); B. oligacanthum (W. Wet. 1952)
subsp. Stella W. Wetzel 1952 (36-42 /lx); B. oligacanthum (W. Wet. 1952) subsp. velatum W. Wetzel
1952 {60 p).

B. palmatum (Deflandre and Courteville 1939) comb. nov. (c. 54 p); B. cL palmatum (Defl. and Court.

92

PALAEONTOLOGY, VOLUME 6

1939), Valensi 1955 (45 /x); B. panniforme Gerlach 1961 (46-68 X 53-105 jU.); B. parvispimim [Deflandre
1937] (Cookson and Eisenack 1958) Klement 1960 (40-84 X 20-46 /u.); B. pattei (Valensi 1948) Sarjeant
1960n (34-37 /x); B. pectiniforme Gerlach 1961 (32-42 /x); B. pilosian (Ehrenberg 1843) Sarjeant 1961
(33-60 /x); B. pilosum (Ehr. 1843) var. longispinosum Sarjeant 1961 (51 x28/x); B. placacanthum
(Deflandre and Cookson 1955) comb. nov. (49-54 /x); B. plicatiim (Maier 1959) comb. nov. (45jlx);
B. polytricluim (Valensi 1947) Sarjeant 1960a (33-60 p); B. polytrichum Downie 1957 non Valensi 1947
(40 /x); B. pseudhystrichodinium (Dell. 1937) comb. nov. ( 38-45 X 49-54 /x); B. pseudhystrichodinium
(Dell. 1937) subsp. magnum W. V^etzel 1955 (98 /x + ); 5. cf. pseudhystrichodinium 1937), Lejeune-
Carpentier 1941 (dimensions similar to that species); B. cf. pseudliYstrichodinium 1937), Gocht
1952 (55x72ju); B. cf. M.pumile (O. Wetzel 1932), Valensi 1948 (26-30 /x).

B. ramidiferum (Deflandre 1937) comb. nov. (35-45 p)\ B. cf. ramidiferum (Defl. 1937), Gocht 1952
(63 x50 p)\ B. reginaldi (Mantell 1846) comb. nov. (?); B. rehdense (Maier 1959) comb. nov. (59-66 /x);
B. satiirnium (Maier 1959) comb. nov. (70 /x); B. seminiidum (W. Wetzel 1952) comb. nov. (30-72 ;x);
B. spiculatum (White 1842) comb. nov. (c. 160 ^u); B. spinosum (White 1842) comb. nov. (40-102 /x);
B. striolatum (Defl. 1937) comb. nov. (40-58 /x); B. cf. striolatum (Defl. 1937), W. Wetzel 1952 (72 p);
B. striatoconus (Deflandre and Cookson 1955) comb. nov. (52-57/x); B. stimuliferum (Defl. 1938)
Sarjeant 19606 (20-26 /x).

B. tiara (Klumpp 1953) comb. nov. (44-68 /x); B. cf. tiara (Klumpp 1953), Deflandre and Cookson
1955 (44x58;x); B. triangulatum Gerlach 1961 (36-60 X 33-48 ;u); B. tribuliferum Sarjeant 19626 (53-
62x48-58 /x); B. tridactylites (Valensi 1955) comb. nov. (40 X 33 p)\ B. trifurcatum Pastiels 1948 non
Eisenack 1931 (30-50 p) \ B. cf. trifurcatum (Eis. 1931), Gocht 1952 (55 p).

B. varispinosum Sarjeant 1959 {c. 48 /x); B. vestitum (Deflandre 1938) Sarjeant 19606 (38-70 /x); B.
cf. vestitum (Defl. 1938), Deflandre 1941 (65 /x); B. whitei (Deflandre and Courteville 1939) Sarjeant
1959 (56-60 jtx); B. xanthiopyxides (O. Wetzel 1933) Klement 1960 (28-30X 10-12 p).

Remarks. Baltisphaeridium spiral isetum de Wit 1943, of which descriptions have been
given only in Dutch, is not treated. Hystricliospliaeridium mensuhim (O. Wetzel 1933) is,
from its description, referred to Veryhachium. H. pterophorum Deflandre and Courteville
1939 is transferred to Cymatiospliaera. H. pumile (O. Wetzel 1933) and H. cf. pilosum
forma nana (O. Wetzel 1933) are transferred to Micrhystridium. The latter form, which
cannot be considered comparable to B. pilosum, is considered probably conspecific with
Micrhystridium incortspicuum and is therefore renamed M. cf. inconspicuum.

Genus micrhystridium Deflandre 1937, emend.

Emended diagnosis. Hystrichospheres with spherical or oval shells not divided into fields
or plates, bearing processes with closed tips, most often simple, rarely branching or
ramifying, without distal connexions of any kind. The processes are generally of one
type only. Mean and modal diameter of shell less than 20 p.

Type species. Micrhystridium inconspicuum (Deflandre 1935) Deflandre 1937 (7-11 /x); Jurassic-
Cretaceous.

Other species (Palaeozoic). Micrhystridium albertensis Staplin 1961 (18 /x); M. angustum Staplin 1961
(14-16/x); M. bacilliferum Deflandre 1946 (8-lOjx); M. bistchoensis Staplin 1961 (c. 17/x); M. diio-
deciaster (Staplin 1961) comb. nov. (formerly Vervhachium) (18/x); M. eatonensis Downie 1959 (12-

22 p).

M. imitatum Deflandre 1942 (8-10 p)\ M. incertum Deunflf 1958 (6-7 p)\ M. mendax Deflandre 1945
(11 p)\ M. micropolygonale Stockmans and Williere 1961 (18 /x); M. nannacanthum Deflandre 1942
(10-14;x); M. parinconspicuum'DQf^^Lr\6.ve 1942 (9-13 /x); M. parvispinum Deflandre 1946 (9-12 /x); M.
parvispinum Defl. 1946 forma major Deflandre 1946 (15 /x).

M. robustum Downie 1958 (10-12 /x); M. shinetonensis Downie 1958 (4—15 /x); M. stellatum Deflandre
1942 (11-16/x); M. stellatum Deflandre 1942 var. inflatum Downie 1959; M. tenuissimum Deflandre

C. DOWNIE, W. A. S. SARJEANT; SOME H YSTRICHOSPHERE GENERA 93

1945 (5 ju,); M. temiissiinwn Deflandre 1945 forma major Deflandre 1946 (10 /u.); M. tomaeeiise Stock-
mans and Williere (lOju,).

Otlwr species (Post-Palaeozoic). Micrhystridiam ambiguum Deflandre 1937 (14— 22ju.): M. cf. amhiguum
Deflandre 1937, Cookson 1953 (13-20/x); M. arachnoides Valensi 1953 (5-8 /x); M. cf. arachnoides
Valensi 1953 (7-8 /x); M. bigoti Deflandre 1947 (13-25 x 12-22 /x); M. biornatiau Deflandre 1937
(15-17 /x); M. castaniinan Valensi 1953 (10/x); M. cometes Valensi 1948 (1 1-19 /x).

M. deflandrei Valensi 1948 (11-19/x); M. densispinum Valensi 1953 (11-13/x); M. echinoides Valensi
1948 (10-17 jtx); M. echinoides Valensi 1948 forma minor Valensi 1953 (7-10 /x); M. fdigernm Valensi
1953 (10-13 /x); M. fragile Deflandre 1947 (7-20 /x); M. ei. fragile Deflandre 1947, Valensi 1953 (12-
16 /x); M. heteracantlmm Deflandre 1937 (8 p).

M. inconspicuum (Defl. 1935) forma bullosa Valensi 1953 (7-10 /x); M. inconspiciium (Defl. 1935)
forma Valensi 1953 (9-13 p) \ M. cf. inconspicuum Deflandre 1935, O. Wetzel 1933 (= Hystricho-
spliaera cL pilosa {oxma nana) (6-20 /tx); A/, cf. inconspicuum (Defl. 1935) Deflandre and Cookson 1955
(dimensions not quoted); M. cf. inconspicuum (Defl. 1935), Sarjeant 1960Z> (6~9 p).

M. lagynophorum Valensi 1953 (13-18 /x); M. leptothrix Valensi 1953 (17-22 ju); M. lucasi Valensi
1953 (14-15 jtx); M. mastigophorum Valensi 1948 (14x 16/u,); M. cf. mendax Deflandre 1945, Sarjeant
1961 (11 ^i); M. multispinosum Pastiels 1948 (c. 15 p); M. nannacanthum Valensi 1953 non Deflandre
1942 (10;u); M. operosum Deflandre 1937 (8-5-11 /x); M. pachydermum Deflandre and Cookson 1955;
M. piliferum Deflandre 1937 (20-22 /x); M. piveteaui Valensi 1953 (19 X 18 ;x); M. polyedricum Valensi
1948 (6x8 p); M. pumile (O. Wetzel 1933) comb. nov. (12-20 p).

M. rarispinum Sarjeant \960b (8-15 ;ix); M. recurvatum Valensi 1953 (10-21 /x); M. recurvaturn
Valensi 1953 forma brevispinosa Valensi 1953 (12-14ju,); M. recurvatum Valensi 1953 forma longi-
spinosa Valensi 1953 (13-14 p) \ M. recurvatum Valensi 1953 forma multispinosa Valensi 1953 ( 14-15 /x);
M. recurvatum Valensi 1953 forma reducta Valensi 1953 (1 1 /x); Af. cf. recurvatum Valensi 1953, Valensi
1953 (8 p); M. cf. recurvatum forma multispinosa Valensi 1953, Sarjeant 1960A (22xl9/x); M.
rhopalicum Sarjeant 1962A (10-15 p)\ M. roquesi Valensi 1948 (16 p).

M. scutospinum Valensi 1953 (14jlx); M. singulare Firtion 1953 ( 14-5 x 18-5 /u,); M. cf. stellatum
Deflandre 1942, Valensi 1953 (up to 18 /x); M. cf. stellatum Deflandre 1942, Deflandre and Cookson
1955 (14-19 ju); ?AF. stellatum Deflandre 1942, Sarjeant 1959 (c. 25 p)\ M. sydus Valensi 1953 (5-7 p);
M. tenuissimum Valensi 1953 non Deflandre 1945 (5 p)\ M. variabile Valensi 1953 (15-16 jtx).

Remarks. The species of Micrhystridium having tubular spines, M. fucosum Valensi
1955 and M. paulinae Valensi 1953, are transferred to the genus Hystrichosphaeridiiim
Deflandre 1937 emend. Eisenack 1958. The species having a shell surface divided by
ridges into fields, M. dictyophoriim Valensi 1953, M. placophoriim Valensi 1948, M.
reticidatum Deflandre 1937, and M. cf. reticidatum Deflandre 1937, Cookson 1953, are
transferred to the genus Cymatiosphaera O. Wetzel 1933 emend. Deflandre 1954. M.
polyedricum forma reducta Valensi 1953 is transferred to the genus Veryhachium and
given specific status under the name Veryhachium valensii (Valensi 1953) comb. nov.

Genus veryhachium Deunflf (1954) 1958, emend.

Emended diagnosis. A genus of hystrichospheres having polygonal or subpolygonal
tests bearing a small number (in general 3-8) of hollow pointed spines with closed tips.
Size of tests 10 /u. to 40 /x, rarely smaller or greater.

Type species. Veryhachium trisulcum Deunff 1958; Ordovician.

Discussion. Distinction from Michystridium and Baltisphaeridium is based on the shape
of the test and the number of spines. Polygonal and subpolygonal forms with nine or
more spines are excluded unless it can be demonstrated that the additional spines are
‘accidental’ and that the individuals are members of a typical Veryhachium group (such
forms are known in the Permian). V. sedecimspinosum Staplin 1961, with fifteen to

94

PALAEONTOLOGY, VOLUME 6

sixteen spines, and V. duodeciaster Staplin, with twelve spines, are therefore transferred
respectively to Baltisphaeridium and to Micrbystridiurn. V. ambiguurn DeunflF is stated to
have nine processes : unless it can be shown that the extra spines are to be considered
‘accidental’, this species also should be transferred to Baltisphaeridium. Certain species
hitherto placed in Micrhystridiwn are transferred to Veryhachium: species are also
transferred to this genus from Baltisphaeridium, Hystrichosphaeridium, and Palaeote-
tradinium.

Distinction between Veryhachium and Estiastra Eisenack depends on the relationship
of the processes to the test. In the latter genus the bases of the processes meet at an acute
angle and the test cannot be distinguished as a separate structure. In Veryhachium, on the
other hand, the test is visibly distinct from the processes, although where the latter have
wide, flaring bases they curve into the test walls and the point of junction cannot be
accurately indicated. Eisenack has placed only one species in Estiastra', few others fit the
diagnosis accurately. V. centrigerum Deunff 1957 accords well with Estiastra', the species
V.florigerum Deunff 1957, V. libratum Deunff 1957, and V. visbyense (Eisenack 1959) all
show transitional features. It is considered for the present best that all four species should
be retained in Veryhachium, the boundary between the two genera appearing quite
arbitrary.

Other species {Palaeozoic). Veryhachium ambiguurn Deunff 1955; V. asymmetricum Deunff 1954; V.
bacifer (Eisenack 1935) Deunff 1954; V. balticum (Eis. 1951) Deunff 1954; V. brevitispinosum Staplin
1961; V. bulbiferum (Deflandre 1944) Deunff 1954; V. centrigerum Deunff 1957; V. ? clava Deunff
1958; V. criicistellatum Deunff 1955.

V. eisenacki Deunff 1954; V. europaeum Stockmans and Williere 1961 ; V. exasperatum Deunff 1955;
V. florigerum Deunff 1957; V. formosum Stockmans and WiUiere 1961; V. furcillatum Deunff 1955;
V. geometrician (Deflandre 1944) Deunff 1954; V. hebetatum Deunff 1957 ; V. heterogonum Deunff 1955.

V. irregulare Jekhowsky 1961; V. irregular e Jekhowsky 1961 forma pyrula Jekhowsky 1961; V.
irregulare Jekhowsky 1961 forma subhexahedron Jekhowsky 1961 ; V. irregulare Jekhowsky 1961 forma
subtetraedron Jekhowsky 1961; V. lairdi (Deflandre 1946) Deunff 1954; V. ledanoisi Deunff 1957;
V. libratum Deunff 1957.

V. ? macroceras Deunff 1958; V. mamillatum Deunff 1954; V. minor Staplin 1961; V. minutum
Downie 1958; V. octoaster Staplin 1961; V. oligospinosum (Eisenack 1934) Deunff 1954; V. polyaster
Staplin 1961 ; V. reductum (Deunff 1958) Jekhowsky 1961 ; V. remotum Deunff 1955; V. rhomboidium
Downie 1959; V. riburgense Brosius and Bitterli 1961 (pars).

V. staurasteroides (Deflandre 1942) Deunff 1954; V. stelligerum Deunff 1957; V. stelligerum Deunff
var. robustum Deunff; V. tetraedron Deunff 1954c; V. tetraedron var. wenlockium Downie 1959; V.
trispinosum (Eisenack 1938) Deunff 1954; V. trisulcum Deunff var. Venetian Deunff 1958; V. visbyense
(Eisenack 1959).

Remarks. The species Veryhachium asymmetricum, V. eisenacki, V. mamillatum, and
V. tetraedron were published as names accompanying figures (Deunff 1954); since no
text descriptions have yet been published, their validity is doubtful.

Other species. {Post-Palaeozoic). Veryhachium hyalodermum (Cookson 1956) comb. nov. (formerly
Palaeotetradinium)', V. mensulum (O. Wetzel 1933) comb, nov.; V. tetraxis (Sarjeant 19606) comb. nov.
(formerly Micrhystridium)', V. valensii (Valensi 1953) comb. nov. ( = Micrhystridiian polyedricum
forma rediicta Valensi 1953).

Genus leiosphaeridia Eisenack 1958, emend.

Emended diagnosis. Spherical to ellipsoidal bodies without processes, often collapsed or

95

C. DOWNIE, W. A. S. SARJEANT: SOME HYSTRICHOSPHERE GENERA

folded, with or without pylomes. Walls granular, punctate or unornamented; thin.
Without divisions into fields and without transverse or longitudinal furrows or girdles.

Type species. Leiospbaeridia baltica Eisenack 1958; Ordovician.

Discussion. Eisenack’s diagnosis is emended to exclude reference to colour, since this is
considered to reflect degree of staining by humic substances rather than any intrinsic
difference. Investigations have shown that single species of fossil microplankton can
exhibit great variation in colour with provenance, and that colour modification can result
from laboratory treatment with oxidizing solutions. The species described under the
invalid names Frotoleiosphaeridium and Leiosphaeridium are transferred to this genus.

Other species {Palaeozoic). Leiospbaeridia aurata (Deflandre 1945) Eisenack 1958; L. cambriense
(Timofiev 1959) comb, nov.; L. conglutinata (Timofiev 1959) comb, nov.; L. cryptogramdosa (Staplin
1961) comb, nov.; L. diaphania (Staplin 1961) comb, nov.; L. eisenackia (Timofiev 1959) comb, nov.;
L. fastigatirugosa (Staplin 1961) comb, nov.; L. faveolata (Timofiev 1959) comb, nov.; L. fragile
Downie 1958.

L. granulata (Eisenack 1938) Eisenack 1958; L. gramdifera (Staplin 1961) comb, nov.; L. granulosa
(Staplin 1961) comb, nov.; L. indefinita (Timofiev 1959) comb, nov.; L. major (Staplin 1961) comb,
nov. ; L. megacystis (Eisenack 1937) Eisenack 1958 ; L. mierocystis (Eis. 1937) Eisenack 1958 ; L. micro-
gramdifera (Staplin 1961) comb, nov.; L. inicrosaetosa (Staplin 1961) comb, nov.; L. miniita (Staplin
1961) comb, nov.; L. nervata (Timofiev 1959) comb, nov.; L. orbiculata (Staplin 1961) comb. nov.

L. papillata {Staplin 1961)comb. nov.; L. (Eisenack 1938) Eisenack 195S: L.parvigranulosa

(Staplin 1961) comb, nov.; L. retigera (Deflandre 1945) Eisenack 1958; L. sorediforme (Timofiev 1959)
comb, nov.; L. temdssima Eisenack 1958; L. voigti Eisenack 1958; L. wenlockia Downie 1959; L.
wimani (Timofiev 1959) comb. nov.

Other species {Post-Palaeozoic). Leiospbaeridia aptiana Eisenack 1958; L. chytroeides Sarjeant 19626;
L. cf. granulata Eisenack 1938, Gocht 1952; L. byalina (Deflandre 1941) comb, nov.; L. cf. micro-
cystis (Eis. 1937), Valensi 1953; L. cf. reticulata Eisenack 1938, Gocht 1952; L. similis Cookson and
Eisenack 1960; L. scrobiculata (Deflandre and Cookson 1955) Eisenack 1958.

Acknowledgements. The authors acknowledge the valuable assistance and comments received from
Professor P. Allen and Mr. J. R. L. Allen, University of Reading; Dr. W. R. Evitt (Jersey Production
Research Co., U.S.A.); Mr. N. F. Hughes (Sedgwick Museum, Cambridge); Mr. G. Norris (New
Zealand Geological Survey); and Mr. D. A. E. Spalding (The Museum, Kingston-upon-Hull, Yorks.)*

REFERENCES

DEFLANDRE, G. 1937. Microfossiles des silex cretaccs. 11. Ann. Paleont. 26,51-103.

— — and DEFLANDRE-RiGAUD, M. 1961. Nomenclature et systematique des hystrichospheres {sen. lat.).

Observations et rectifications. Paris; Lab. de Micropaleontologie. 1-15.

DEUNFF, j. 1954. Veryhachium, genre nouveau d’hystrichospheres du Primaire. C. R. Somm. Soc.
geol. fr. 13, 305-6.

1957. Micro-organismes nouveaux (Hystrichospheres) du Devonien de I’Amerique du Nord.

Bull. Soc. geol. et min. de Bretagne, 2, 5-14.

DOWNIE, c. 1957. Microplankton from the Kimeridge Clay. Quart. J. Geol. Soc. Land. 112,413-34.

1960. Deunffia and Domasia, new genera of hystrichospheres. Micropaleontology, 6, 197-202.

■ , WILLIAMS, G. L. and SARJEANT, w. A. s. 1961. Classification of fossil microplankton. Nature, 192,

471.

EISENACK, A. 1958o. Tasmanites Newton 1875 and Leiospbaeridia n.g. als Gattungen der Hystricho-
sphaeridia. Palaeontographica, 110a, 1-19.

19586. Mikroplankton aus dem norddeutschen Apt nebst elnigen Bemerkungen fiber fossile

Dinoflagellaten. Neues Jb. Geol. Palaeont. Abb. 106, 383-422.

96 PALAEONTOLOGY, VOLUME 6

EISENACH, A. 1959. Ncotypen baltischer Silur — Hystrichospharen und neue Arten. Palaeontographica,
112 a, 193-211.

1962. Einige Bemerkungen zu neueren Arbeiten iiber Hystrichospharen. Neiies Jb. Geo!. Pal.

Mb. 2, 92-101.

EViTT, w. R. 1961. Observations on the morphology of fossil dinoflagellates. Micropaleontologv, 7,
385-420.

JEKHOWSKY, B. DE, 1961. Sur quelques hystrichospheres permo-triasiques d’Europe et d’Afrique. Rev.
Micropal. 3, 207-12.

SANNEMANN, D. 1955. Hystrichosphaerideen aus dem Gothlandium und Mittel-Devon des Franken-
waldes und ihr feinbau. Senck. leth. 36, 321-46.

SARJEANT, w. A. s. 19616. The hystrichospheres: a review and discussion. Grana Palynologica, 2,
101-11.

STAPLiN, F. L. 1961. Reef-controlled distribution of Devonian microplankton in Alberta. Palaeonto-
logy, 4, 392-424.

TiMOFiEV, B. V. 1956. Hystrichosphaeridae from the Cambrian. Dokl. Akad. nauk SSSR 106, 130-2.
(In Russian.)

1959. The ancient flora of the Pre-Baltic and its stratigraphic significance. Mem. VNIGRl, 129,

1-350. (In Russian.)

VALENSi, L. 1953. Microfossiles des silex du Jurassique moyen. Remarques petrographiques. Mem.
Soc.geol.fr. 68, 1-100.

CHARLES DOWNIE

Department of Geology,
University of Sheffield

WILLIAM A. S. SARJEANT
Department of Geology,
University of Nottingham

Manuscript received 28 February 1962

PHYLLOCARID CRUSTACEANS FROM THE
SILURIAN AND DEVONIAN OF CZECHOSLOVAKIA

by I VO CHLUPAC

Abstract. Recent discoveries of phyllocarids in the Silurian and Devonian of the Barrandian area in Bohemia
and the Upper Devonian of the Moravian Karst area are described. Nearly complete specimens of Aristozoe
parabolica Perner and Fygocaris schuberti Perner have been found, permitting a more detailed diagnosis of the
family Aristozoidae, to which the genus Heroldina Broili has also been referred. Ceratiocaris cornwalliseiisis
damesi subsp. nov. from the Czech Silurian and Ceratiocaris (?) coherbaria sp. nov. from the Middle Devonian
are described.

Concavicaris desiderata (Barrande) from the Czech Middle Devonian and C. incola sp. nov. from the Moravian
Upper Devonian are described and assigned to the genus Concavicaris Rolfe (formerly Colpocaris Meek),
previously known only from the American Carboniferous. The systematic position of the genus Montecaris Jux
is discussed and the description of M. brunnensis Chlupac is amplified.

Phyllocarids from the Silurian and Devonian of Czechoslovakia are known partly
from the classical Barrandian area in central Bohemia and partly from the Moravian
Karst area in central Moravia. In the Barrandian area phyllocarids have been discovered
in beds of Silurian, Lower and Middle Devonian age; in the Moravian older Palaeozoic
rocks they have so far been found only in the Upper Devonian.

The fundamental description of the Bohemian Silurian and Devonian phyllocarids was
published in J. Barrande’s appendix to the first volume of his Systeme silurien du centre
de la Boheme (1872). Important facts about Bohemian phyllocarids were added in papers
by Novak (1885^z, b\ 1886a, b), but Novak died before his large prepared work was
published. In this century Perner (1916, 1919) has studied the Czech phyllocarids and
described new forms from the uppermost Silurian, and Gurich (1929) has discussed
questions of systematics. Otherwise, with the exception of the author’s brief communica-
tion (19606), the interesting phyllocarid fauna of the Bohemian Silurian and Devonian
has not been studied by modern methods. In the recent literature only sporadic mention
can be found of the occurrence of single species, e.g. Boucek (1938), Prantl and Pfibyl
(1948), and Chlupac (1953, 1955).

In Moravia the first remains of phyllocarids were only recently discovered in the Upper
Devonian of the southern part of the Moravian Karst (Chlupac 19606).

During new systematic researches in the Silurian and Devonian of central Bohemia as
well as in the Devonian of Moravia we have succeeded in finding numerous further
specimens. These finds of more completely preserved individuals of formerly known
species or of new forms improve our knowledge of the Czechoslovakian phyllocarid fauna.
The present paper deals with some finds of systematic and stratigraphical importance.

Acknowledgements. For this study the Bohemian Silurian material from the collections of Diplom-
geologist R. Horny, F. Kalfus, Dr. M. Snajdr, and J. Vanek was used besides the material collected by
the author. The author wishes to thank Miss O. Hofmanova for allowing him to study her specimens
from the Devonian of the Moravian Karst. Professor Dr. R. Kettner and Diplomgeologist J. Obrhel
made it possible for the author to study the older material from the collections of the Geological-
Palaeontological Institute of the Charles University, and R. Horny gave access to the originals in the

[Palaeontology, Vol. 6, Part 1, 1963, pp. 97-118, pi. 12-16.]

C 1015 H

98

PALAEONTOLOGY, VOLUME 6

Barrandeum of the National Museum of Czechoslovakia. The author wishes to express his thanks to
all the above-mentioned scientists for their willingness and understanding as well as his sincere appre-
ciation to Dr. W. D. I. Rolfe, Hunterian Museum, University of Glasgow, for his kind comments and
help in the comparative study of phyllocaris.

Repositories. The described material is deposited mainly in the author’s collection in the Central
Geological Institute in Prague (in the text abbreviated to ICh). Other specimens used in the paper are in
the collections of the National Museum in Prague (abbreviated to NM) and Geological-Palaeontological
Institute of the Natural Science Faculty of the Charles University (abbreviated to KU). The photo-
graphs were made by R. Horny (National Museum, Prague) and H. Vanova (Central Geological
Institute, Prague).

SYSTEMATIC DESCRIPTIONS

Order phyllocarida Packard 1879
Suborder ceratiocarina Clarke 1900

Family aristozoidae Giirich 1929

The representatives of this family have been known only on the basis of isolated parts
of the carapace. New finds of almost complete specimens of the genus Aristozoe Barrande
and Pygocaris Perner, in which the carapace, abdomen, and telson are preserved con-
nected in their original position, make it possible to improve the diagnosis of the family.

Revised diagnosis. Carapace bivalved, with prominent nodes in the anterior part and
distinct marginal rim. Small number (3, in Heroldina 5) of abdominal segments preserved
outside the carapace, the last of which is strikingly elongated. The caudal part consists of
one long spine only — a telson, which is usually dorso-ventrally curved. Furca not found.

Remarks. The bivalved carapace without a dorsal plate and free rostrum of genera of the
family Aristozoidae proves that they belong to the suborder Ceratiocarina. The most
outstanding feature of the family is that the caudal portion is formed by a single spine,
the telson, usually curved, while traces of the furca have not been ascertained even in
complete, well-preserved individuals (e.g. even of the genus Heroldina). Novak (1886Z?),
who first called attention to the absence of furca, had, however, only isolated carapaces
at his disposal. In this the representatives of the family Aristozoidae differ not only from
other families of the suborder Ceratiocarina, but also from other phyllocarids and
archaeostracans in general, to which according to other features they undoubtedly
belong. In any case the family Aristozoidae Giirich represents an independent group,
markedly differentiated from other families but showing closest relation to the suborder
Ceratiocarina Clarke, especially to the family Echinocarididae Clarke. The configuration
of the caudal part in the family Aristozoidae is, however, an exceptional phenomenon
among the Archaeostraca. The new discoveries indicate that we cannot regard the
assignment of the genus Aristozoe and other related genera to ostracods given by
Barrande (1872) and in the Treatise on Invertebrate Paleontology, Part Q (Moore 1961),
as correct. The comments of W. D. I. Rolfe (ibid., p. Q429) are, therefore, more than
justified.

Genera. The genera Pygocaris Perner and Orozoe Barrande belong to the family Aristo-
zoidae together with the type genus Aristozoe Barrande. The genus Heroldina Broili
without any doubt belongs here. It agrees with Aristozoe and Pygocaris in having a

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS 99

greatly elongated last abdominal segment and especially in having the characteristic
caudal part represented by the single curved telson (Broili 1929). To this family the
author refers the genus Callizoe Barrande only with reserve, the nodes of which are
situated more antero-ventrally than in other genera and the valves show weak lateral
carinae.

Genus Aristozoe Barrande 1872
Aristozoe parabolica Perner 1919

Plate 13, figs. 1-3; text-fig. 1

1916 Aristozoe parabolica Perner, pp. 5, 6, pi. 1, figs. 9-11 (Czech description).

1919 Aristozoe parabolica Perner, p. 228, pi. 1, figs. 9-11 (German description).

1929 Aristozoe parabolica Perner; Gurich, p. 62, text-figs. 7-14.

1934 Aristozoe parabolica Perner; Straelen and Schmitz, p. 100.

Lectotype. Perner 1919, pi. 1, figs. 9-11, NM Br042.

Type locality. Kosof, near Praha, Czechoslovakia.

Horizon. Lochkov formation, Radotin-Limestones facies, Lochkovian, Zone of Monograptiis lier-
cynicus, Silurian.

Emended description. The description of the carapace was given by Perner (1916, 1919).
The main features of the carapace are the characteristic and almost parabolic outline of
the ventral border of the valves, the closeness of the length to the width of the valves
(5:4 to 8:7), an obliquely truncate postero-lateral region, a narrow marginal rim, two
nodes in the antero-dorsal region, the posterior one larger and more convex than the
anterior one, semi-elliptical, and dorso-ventrally elongated. It is necessary, however, to
add to Perner’s description and illustrations that the postero-dorsal border is not so
broadly rounded as Perner states. The original depicted in Perner’s pi. 1, fig. 9 is badly
damaged and incompletely preserved in the postero-dorsal part (in the positive entirely
broken off), and in this respect the illustration is wrongly reconstructed. From better-
preserved specimens (see PI. 13, figs. 2, 3) it is evident that the marginal rim extends as
far as the dorsal border, where the postero-dorsal angle is obtuse.

The recent material (coll. ICh 105) presents a strongly compressed specimen of A.
parabolica, in which besides the carapace even the abdominal segments and telson in
natural position are preserved (see text-fig. 1 ). The carapace of this individual is ill pre-
served owing to flattening. The assignment to A. parabolica can be proved by the outline
preserved in greater part, a narrow marginal rim and smooth surface (the nodes rendered
indistinct by deformation). The right and the left valves of the carapace of this individual
overlap each other only partially: the left valve was posthumously displaced ventrally
relative to the right valve so that a considerable part of the ventral sector of the left valve
remains uncovered by the right valve, causing the apparently unusually large dorso-
ventral width of the carapace. In the antero-dorsal part two free portions of the carapace
are observed which could be interpreted as a broken rostrum. Owing to the strong
compression of the valves the possibility that these are secondarily separated fragments
of the antero-dorsal part of the carapace cannot be excluded.

Outside the carapace, in addition to the telson, fragments of three abdominal segments
have been preserved. The first has been preserved only as three isolated fragments of the
posterior part of the segment. The second segment has been preserved more completely,
its width considerably exceeding its length (approx. 1 : 3) while the outline shows that in

100

PALAEONTOLOGY, VOLUME 6

the dorsal part the segment was longer than in the ventral. The postero-dorsal part is
pointed and projects postero-dorsally. The last abdominal segment is considerably
elongated, about four times longer than the penultimate one. The proportion between
sagittal length and width after lateral compression is 2 ; 1. At the proximal border of the
segment in the antero-dorsal part there is a swelling, which in spite of the compression
is clearly perceptible. At a distance of 9 mm. from the proximal end of the last segment
runs a dorso- ventral transverse furrow which could possibly be considered as a dividing-

TEXT-FiG. I. Aristozoe parabolica Perner. ICh 105, laterally compressed specimen
with abdomen in natural position, left valve of the carapace displaced ventrally.

line between two abdominal segments. The fact that the dividing-line is interrupted in
the middle and that the distinct swelling characteristic of the anterior part of the last
segment cannot be found near this line is incompatible with this interpretation. The
analogy with the closely related genus Pygocaris Perner also tends against this explana-
tion. Slight traces of dorso-ventrally running irregular transverse striae are preserved
on the last segment.

The telson, which is articulated with the last abdominal segment and lacks the distal
part, shows a slight dorso-ventral curvature. It bears two distinct small longitudinal
ridges; at the dorsal border traces of insertions of bristles are partly preserved. Remains
of the furca have not been found.

Dimensions. ICh 105 (PI. 13, fig. 1 ; text-fig. 1), max. length of the carapace 55 mm, dorso-ventral width
of the carapace 42 mm., length of the abdominal part without telson 35 mm. Length of the last
abdominal segment 24 mm., width of the last abdominal segment 1 1 mm. ICh 102 (PI. 13, figs. 2, 3),
max. length of the carapace 47 mm., width 38 mm.

Remarks. A. parabolica Perner has hitherto been known only on the basis of isolated
valves of the carapace. The described specimen with preserved abdominal part, in spite
of its less favourable and somewhat incomplete preservation, is of considerable signifi-
cance as it constitutes the first specimen of Aristozoe in which carapace and abdominal
part of the body are preserved in natural position. This confirms to a great extent the
correctness of Novak’s reconstruction (1885) which was made on the basis of isolated

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS

101

carapaces and segments of A. regina Barrande from the Lower Devonian. It is significant
that only a small number (3) of abdominal segments was observed, which is analogous
with Pygocaris Perner and obviously characteristic of the family Aristozoidae.

From the Lochkov Formation of the Kosor locality Novak (1886^, pp. 15, 16)
described a minute isolated telson with longitudinal ridges which he designated as
Aristozoe solitaria Novak. His specimen agrees with the telson of A. parabolico described
above in having distinct small longitudinal ridges, dorso-lateral insertions of bristles,
and a slight curvature. The incomplete preservation of the telson of A. paraboUca
the precludes identification of the species with A. solitaria, especially since in the same
strata several related phyllocarids occur together.

Occurrence. A. parabolica has so far been found only at the classical locality in the
Silurian-Devonian boundary beds near Kosor (quarries at Cerna rokle). All known finds
come from the blackish-grey thin-bedded fine-grained limestones and dark calcareous
shales which alternate with the limestones. The beds belong to the Lochkovian (i.e. to
the Lochkov Limestones in the Radotin facies), Monograptus hercyniciis Zone, upper-
most Silurian. The beds are probably younger than the British Upper Ludlovian as
mentioned by Boucot (1960). They are assigned to the uppermost Silurian in accordance
with the conclusion of the Prague symposium (Svoboda 1958). Specimens are not
abundant; about twenty discoveries have so far been made, and except for one whole
described specimen consist of valves of the carapace only.

Genus Pygocaris Perner 1919
Pygocaris schuberti Perner 1919

Plate 12, fig. 8; Plate 13, figs. 4, 5; text-fig. 2

1916 Pygocaris schuberti Perner, pp. 2-5, pi. 1, figs. 1-5 (Czech description).

1919 Pygocaris schuberti Perner, pp. 226, 111, pi. I, figs. 1-5 (German description).

1929 Pygocaris schuberti Perner; Giirich, pp. 54, 62; text-pl. 5, fig. 16.

1934 Pygocaris schuberti Perner; Straelen and Schmitz, p. 193.

1960 Pygocaris schuberti Perner; Krestovnikov, p. 428, fig. 1254.

Lectotype. Perner 1916, 1919, pi. 1, figs. 1-4, NM Cel593.

Type locality. Kosor, near Praha, Czechoslovakia.

Horizon. Lochkov Formation, Radotin-Limestones facies, Lochkovian, Zone of Monograptus her-
cynicus, Silurian.

Emended description. Perner (1916, 1919) described the carapace as possessing the
following distinctive features : a prominent tapered-off projection at the anterior border
of the valves and a tapering postero-dorsal part of the valves; an outstanding and fairly
broad marginal rim separated off by a deep border groove; a greater number (4?) of
nodes in the antero-dorsal region ; and a characteristic ornamentation formed by promi-
nent longitudinal anastomosing and often bent small ridges of unequal length.

Besides a number of carapaces which confirm the description given by Perner, a young
specimen has been found among the new material in which the carapace, abdominal
segments, and telson are preserved in the natural position (ICh 100, PI. 12, fig. 8; PI. 13,
figs. 4, 5), and which allows Perner’s specific description to be amplified. The specimen is
preserved in a lateral position and is strongly compressed laterally so that the carapace

102

PALAEONTOLOGY, VOLUME 6

does not show the original convexity. Only the posterior half of the carapace is preserved,
and by its characteristic outline, postero-dorsally tapering, broad marginal rim, and
traces of striation preserved at the dorsal border of the carapace, clearly belongs to P.
schuberti.

Outside the carapace traces of three abdominal segments are preserved in addition to
the telson. The first, having been evidently partly covered by the carapace, is incom-
pletely preserved; only the posterior, thickened border of the segment, running sub-
parallel to the posterior border of the carapace, is more distinct. Being laterally compressed
the second segment has an almost sub-trigonal outline and the dorsal border was
evidently longer than the ventral border. The length at the dorsal border approximately
equals the dorso-ventral length. As in ^4. parabolica the segment terminates posteriorly
in a postero-dorsal projection. The last abdominal segment, unlike the previous ones,
is well preserved despite the lateral compression. The length of the segment is conspicu-
ous; it is two and a half times longer than the dorso-ventral width. At the anterior
border the segment was evidently more massive, as shown by its convex surface. At the
antero-ventral border a laterally situated node is preserved; the swelling of the segment
and the elevation of this part are possibly due to muscular insertions. The postero-dorsal
border of the segment may have run out in a short caudal projection which partly over-
lapped the head of the telson.

The head of the telson was articulated with the posterior border of the last segment,
which was embayed. On the surface of the last segment an ornament is perceptible
formed by dorso-ventrally running, slightly un-
dulating, small ridges. This type of ornament
corresponds well with the sculpture of the sur-
face of the carapace in F. schuberti. Only the
proximal half of the telson is preserved. It is
distinctly bent in the dorso-ventral direction and
carries two longitudinal ridges. The head is
slightly enlarged and the spine tapers gradually
backwards. Traces of bristle insertions are not
preserved. Furcal rami have not been found,
although the articulation between the last ab-
dominal segment and telson is well preserved.

EXPLANATION OF PLATE 12

Figs. 1, 2. Concavicaris iiicola sp. nov. Hady, near Brno. 1, Holotype, ICh 176, carapace (left valve),
x2-6. 2, Paratype, ICh 178fl, carapace (right valve), x2-3.

Figs. 3-5. Concavicaris desiderata (Barrande). Pekarek mill, near Chotec. 3, Paratype D, ICh 193, left
valve of the carapace showing the rostral plate, X 2-5. 4, Paratype A, ICh 194, carapace (left valve),
x2-6. 5, Paratype B, ICh 192, carapace (left valve), x3.

Figs. 6, 7. Ceratiocaris (?) coherbaria sp. nov. Karlstejn. 6, Restored telson and furca, after holotype,
ICh 126n, x2. 7, Detail showing the sculpture of the telson, after ICh 123, x4-6.

Fig. 8. Pygocaris schuberti Perner. Kosor, near Praha. Paratype A, ICh 100. Incompletely preserved
carapace, abdominal segments and telson in natural position, x 1.

Figs. 9, 10. Ceratiocaris cornwallisensis damesi subsp. nov. 9, Paratype, ICh 137, detail of the distal
portion of telson (lateral view) showing pits marking points of insertion of bristles situated ventro-
laterally; Samor near Liten; x4-7. 10, Paratype, ICh 250, detail of compressed telson showing the
stripe of pits marking points of insertion of bristles; Kosof; x2.

TEXT-FIG. 2. Pygocaris schuberti Perner.
ICh 110, laterally compressed abdomen.

Palaeontology, Vol. 6

PLATE 12

CHLUPAC, Phyllocarid crustaceans

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS

103

The shape of the abdominal part of the carapace is also visible in another specimen
(ICh 110), which, by analogy with that described above, the author also refers to
P. scJmberti. It is an isolated flattened abdomen with telson, preserved in lateral posi-
tion (text-fig. 2). In addition to the telson three abdominal segments are clearly visible;
the first two are much shorter than the last. The last segment is more than three times
longer than it is wide; it is, however, impossible to exclude deformation caused by
compression. The telson, characteristically slightly but distinctly dorso-ventrally curved,
preserves for part of its length a number of insertions of bristles located dorso-laterally.
Traces of furca rami are lacking.

Dimensions (in mm.)

Holotvpe
(NM Cel593)

Paratype A
(ICh 100)

Paratype B
(ICh 110)

Length of carapace ....

c. 103

c. 35

—

Dorso-ventral width of carapace

58

c. 25

—

Length of last abdominal segment

—

16

20

Dorso-ventral width of last abdominal segment

—

6

7

Length of telson ....

■

c. 35-40
(20 mm. pre-
served)

38

Remarks. New finds of the abdomen of P. schuberti show a distinct resemblance to
Aristozoe parabolica. Their common features are especially the small number of abdomi-
nal segments, a considerable elongation of the last segment, and a distinctly curved
telson. It is important to note that in neither of these species could traces of furcae be
established.

The telson of P. schuberti differs by its stronger curvature and shorter head from the
isolated telson described by Novak (1886Z^) from the same formation as Aristozoe
solitaria. On the other hand the possibility that the specimen described by Fritsch (1907)
as Onchus siluricus Fritsch is conspecific with P. schuberti cannot be excluded. Fritsch’s
original from the Kotys-locality, near Kon“prusy (coll. NM Ce 1456), probably from
the lower part of the Lochkov formation in the Radotin facies, is doubtless a fragment
of an aristozoid phyllocarid. The curved spine bearing longitudinal ridges represents
the telson and the fragments interpreted by Fritsch as intraclavicula and scapula are the
remains of abdominal segments. The resemblance between Fritsch’s original and the
telson of P. schuberti is striking. As these occur in the older horizon of the Lochkov
limestones (Monograptus imiformis Zone) where the occurrence of P. schuberti has not
yet been proved by the discovery of carapaces, and as the telsons of different represen-
tatives of the family Aristozoidae resemble each other so closely that it is not yet possible
to exclude assignment to another species, the author does not at present consider
‘'Onchus' siluricus synonymous with P. schuberti.

Occurrence. P. schuberti is so far only known with certainty from the classical locality at
Kosof, south-west of Prague, where it occurs in the same beds as A parabolica, with
a rich marine fauna (Chlupac 1953). It is found in intercalations of dark fine-grained

104

PALAEONTOLOGY, VOLUME 6

limestones and in highly calcareous shales, and occurs about as abundantly as A. para-
bolica; twenty-five specimens have so far been found.

Family Ceratiocarididae Salter 1863
Genus Ceratiocaris M'Coy 1849

Ceratiocaris coruwallisensis Copeland 1960

1960 Ceratiocaris coruwallisensis Copeland, pp. 49, 50; pi. 8, figs. 1, 2; pi. 9, fig. 5.

Ceratiocaris coruwallisensis damesi subsp. nov.

Plate 12, figs. 9, 10; Plate 14, figs. 3-5; Plate 15, figs. 1-4; text-figs. 3-5

18866 Ceratiocaris damesi Novak, p. 676 (nomen nudum).

Derivation of name. After Mr. W. Dames; the original manuscript name suggested by Novak has been
used.

Holotype. Two last abdominal segments and telson, NM Br283 (PI. 15, fig. 1 ; text-fig. 3).

Type locality. ‘Cerna rokle’, Kosor, near Praha, Czechoslovakia.

Horizon. Lochkov Formation, Radotin-Limestones facies, Lochkovian, Zone of Monograptus her-
cynicus, Silurian.

Material. Two incomplete carapaces; a great number of caudal appendages partly with abdominal
segments; and about thirty mandibles.

Description. Carapace incompletely known, lacking anterior region (text-fig. 5). Outline
suboval, the dorsal border much less convex than the ventral one, posterior border
truncate, a narrow marginal rim, the border line distinctly marked. Surface sculpture
not observed.

The abdominal part has not yet been found with a carapace in natural position, so that
the number of abdominal segments is unknown. In several specimens disarticulated

EXPLANATION OF PLATE 13

The specimens in Figs. 1-3 and 5 were coated with ammonium chloride before being photographed.

Figs. 1-3. Aristozoe parabolica Perner. Kosor, near Praha. 1, Paratype A, ICh 105, laterally compressed
specimen showing abdominal portion and telson in natural position (left valve of the carapace dis-
placed ventrally relative to the right valve), X 1 T . 2, 3, Paratype B, ICh 102, left valve of the carapace
only slightly compressed showing the vaulting and nodes differently illuminated, X IT.

Figs. 4, 5. Pygocaris schuberti Perner. Kosor, near Praha. Paratype A, ICh 100. 4, Incompletely
preserved carapace, abdominal segments and telson in natural position, X 1. 5, Detail of the same
specimen showing the articulation of the last abdominal segment and telson, x 2.

EXPLANATION OF PLATE 14

The specimens in Figs. 1-3 and 5 were photographed under alcohol, and the specimen in Fig. 4 was
coated with ammonium chloride before being photographed.

Figs. 1, 2. Montecaris brunnensis Chlupac. Bedfichovice, near Brno. 1, Paratype C, ICh 200u.
2, Counterpart of the same specimen, ICh 2006, X 1 .

Figs. 3-5. Ceratiocaris coruwallisensis damesi subsp. nov. Kosor, near Praha. 3, Paratype NM 1114,
laterally compressed telson showing the backwardly curved posterior projection in the proximal
portion, X 1-8. 4, ICh 109, isolated non-compressed mandible of a big specimen, x 1. 5, ICh 115i7.
Compressed mandibles and isolated fragments of the carapace, x 0-9

Palaeontology, Vol. 6

PLATE 13

CHLUPAC, Phyllocarid crustaceans

Palaeontology, Vol. 6

PLATE 14

CHLUPAC, Phyllocarid crustaceans

IVO CHLUPAC; SILURIAN AND DEVONIAN PHYLLOCARIDS

105

abdominal segments and caudal appendages are, however, pre-
served near the remains of carapaces, so that there is no doubt
that they belong to the same subspecies.

It is impossible to decide which abdominal segments the
isolated specimens represent, but two last abdominal segments
are preserved in natural position. The penultimate segment, e.g.
in the holotype (text-fig. 3; PI. 15, fig. 1), shows a subquadrate
outline and is much shorter than the last segment (the ratio of
its length to the length of the last segment is 1:2-5). Several
pieces show sculpture in the anterior part formed by fine anas-
tomosing small ridges and striae of unequal thickness. In front
they run subparallel to the anterior border, but they soon turn
obliquely postero-ventrally.

The last abdominal segment is considerably elongated and is
about two and a half times longer than the penultimate segment.
It is more or less flattened on all specimens so that the original
contours are not preserved. It has a subquadrate outline, the
length exceeding at least twice the dorso-ventral width. At the
anterior border the segment, even in the flattened specimens, is
arched and suggests the contours of the original segment. In this
position the sculpture is very often preserved and is seen to
consist of unequally thick small ridges and striae as on the
penultimate segment. Anteriorly the striae are subparallel to
the anterior border, but they soon turn away postero-ventrally
from the dorsum. On the holotype it can be clearly observed
that these small ridges and striae on the ventral side form acute
angles with vertices pointing backward. The posterior part of
the last segment is unornamented in all the specimens so far
known. Near the posterior border the segment was obviously
thickened as in the anterior region, although to a lesser
extent.

The most frequently found specimens are large caudal ap-
pendages, represented by a long telson and shorter lateral spines.
The telson is long and dagger-shaped, with a conspicuously
broadened strongly vaulted head, and preserves its convexity
at least in part, even in the compressed specimens. The telson
is almost straight and gradually tapers backwards. In large
specimens only was it possible to establish that the distal part
of the telson is very moderately dorsally curved. The head of
the telson is provided anteriorly with a narrow articulation plate.
The head carries on both sides of the ventral surface two strong
distinct tooth-like projections, the anterior of which is simply
pointed, while the posterior one is produced into a short curved
spine (PI. 14, fig. 3). The head bears traces of sculpture formed as
in the abdominal segments by irregular small ridges and striae
running postero-ventrally from the anterior border.

TEXT-FIG. 3. Ceratiocaris
cornwallisensis damesi
subsp. nov. Holotype
NM Br283, laterally
compressed two last ab-
dominal segments and
caudal appendages.

106

PALAEONTOLOGY, VOLUME 6

A cross-section of the telson cannot be obtained from the large, more or less com-
pressed specimens. When the specimens are laterally compressed only two longitudinal
carinae are noticeable, one at the dorsal border, and the other at the ventral. Another
lateral longitudinal line, indistinctly marked in relief, runs postero-ventrally from the

head. It was possible to study in detail the cross-section
and nature of the telson on undeformed young speci-
mens from the locality Samor, near Liteh. The telson
shows five longitudinal ridges. A prominent dorsal
ridge running along the median line of the telson has a
sharp crest, and two dorso-lateral ridges have rounded
crests as do the two ventro-lateral ridges. The ventral
concavity of the telson in the proximal part is con-
spicuous where the section of the spine is star-shaped.
This ventral concavity dies out posteriorly and the sec-
tion of the spine becomes subpentagonal. The ventral
concavity disappears completely in the distal part of
the telson, and the two ventro-lateral ridges fuse into
one so that the section of the telson becomes sub-
quadrate (text-fig. 4).

The pits marking the insertions of bristles are situated
laterally on the telson between the dorso-lateral and
ventro-lateral ridges. In the paratype from Kosof (ICh
250) it is possible to establish that the pits do not form
a single row but several longitudinal rows of rounded
pits of different sizes (PI. 12, fig. 10). The largest pits
form a row near the dorso-lateral ridge, under which
are three to four rows of tiny insertions not quite
regularly arranged, the size of the latter decreasing ventrally. Some similarity exists with
the specimens described by Barrande (1872) as Ceratiocaris tardus from the Devonian
of Bohemia.

In the adult specimens the lateral spines of the furca are approximately two-thirds of
the length of the telson (in the young specimens the ratio of the length of telson to
lateral spines ranges around 5 : 3). The spines taper posteriorly more rapidly than the
telson, and older specimens (text-fig. 3; PI. 15, fig. 3) show a slight dorsal curvature.
Only two longitudinal ridges are visible on the laterally compressed specimens, namely
at the dorsal and ventral border. The unflattened specimens from Samor show a subovate
cross-section of the spines, sharp dorsally. The exterior surface of the lateral spines

TEXT-FIG. 4. Ceratiocaris cornwal-
lisensis damesi subsp. nov. ICh 139,
telson with furcal rami of a young
specimen showing the section of
the central spine.

EXPLANATION OF PLATE 15

The specimens in Figs. 3 and 4 were coated with ammonium chloride before being photographed.

Figs. 1-4. Ceratiocaris cormvallisensis damesi subsp. nov. Kosof, near Praha. 1, Holotype, NM Br283,
two last abdominal segments and caudal appendages laterally compressed, 0 3. 2, Paratype A,
ICh 143, two last abdominal segments and caudal appendages in lateral position, x0 5. 3, ICh 113o,
incompletely preserved and laterally compressed caudal appendages showing a slight bending of the
spines, X0 7. 4, ICh 142, isolated abdominal segment showing the sculpture, X 1-8.

Palaeontology, Vol. 6

PLATE 15

CHLUPAC, Phyllocarid crustaceans

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS 107

shows no traces of sculpture. Only in the casts from Samor were irregular, fine, postero-
dorsally oriented ridges observed which indicate the sculptured interior surface of
the integument. Traces of circular insertions of bristles form a longitudinal row on the
lateral spines just under the dorsal ridge, but they are rarely preserved.

TEXT-FIG. 5. Ceratiocan's coniwallisensis dawesi subsp. nov. KU 1208. Displaced
carapace, mandibles, and abdominal segments.

In addition to the carapace, abdominal segments, and caudal appendages, mandibles
are known, but they are usually isolated from the other parts of the shield. In several
cases, however, they were found either directly within the carapace (but not in natural
position) or close to the carapace so that the assignment to C. cormvaUisensis damesi
subsp. nov. is certain. The mandibles may be up to 50 mm. long. The corpus mandibulae
is produced into a comparatively long sickle-shaped process formed of a much thinner
carapace than that over the oral part. The arcuate oral part proper is very massive and
carries four to six paired, partly alternating pointed teeth, which have radially running
furrows on the surface.

Dimensions. Ceratiocaris coniwallisensis damesi subsp. nov. is probably the largest Bohemian phyllocarid,
and of all Bohemian species only C. bohemica Barrande attains similar dimensions. The specimens
found indicate that the length of the whole body can be estimated as being up to 70-80 cm.

Dimensions (in mm.)

Holotvpe
(NM Br283)

Paratype A
UCh 143)

NM 35615

Length of penultimate abdominal segment

37

—

—

Length of last abdominal segment .

90

70

—

Length of telson ......

210

c. 150

c. 300

Length of lateral spine .....

117

85

210

The carapace (paratype C, KU 1208) is about 200 mm. long, the maximal dorso-ventral width being
110 mm. In ten young specimens from Kosof and Samor the length of the telson varies between 17 and
50 mm., the length of the lateral spines being 1 1-31 mm.

108

PALAEONTOLOGY, VOLUME 6

Remarks. The described specimens from the uppermost Silurian of Bohemia were
formerly mentioned by a number of authors as Ceratiocaris damesi Novak (Novak’s
manuscript name) but without description and illustration. Copeland (1960) has
recently described C. cornwallisensis from the Upper Silurian of Cornwallis Island in the
Arctic area of Canada, and it is very closely related to Ceratiocaris damesi Novak MS.
Both forms have the same general structure and agree in the main features of sculpture
of caudal appendages and abdominal segments. A more detailed comparison of the
specimens figured by Copeland (1960) and the material from Bohemia reveals, however,
some differences of lesser importance. The difference between the length of the telson and
the lateral spines is in the Bohemian specimens somewhat greater than in the specimens
figured by Copeland. In the Canadian form no curvature of the lateral spines has been
established. From Copeland’s description there seem to be differences in the cross-section
of the telson but, considering the compression of the Canadian material, they are prob-
ably only secondary. According to Copeland’s description the length of the last
abdominal segment is much shorter than in the Bohemian specimens; however, accord-
ing to W. D. I. Rolfe {in lift.), the last segment in the holotype of C. cornwallisensis is
longer (61 mm.); the indication of a shorter length was due to the mistaking of casual
cracks for the boundary between segments. The comparison shows that despite close
relationship it is not possible to regard the Canadian and the Bohemian forms as com-
pletely synonymous. Since a number of features in the Bohemian material could not be
established in the described Canadian material, the author considers it appropriate to
designate the Bohemian form as a separate subspecies which may be a geographical
mutation of the Canadian species.

Occurrence. The occurrence of C. cornwallisensis damesi subsp. nov. is restricted in the
Barrandian to the uppermost Silurian Lochkov Formation (Lochkov stage) in which it is
abundant in some places. Most of the material comes from the Radotin and Kosor
facies of this formation from the Cerna rokle quarries near Kosof (Zone of Monograptus
hercynicus), where it is accompanied by a rich marine fauna. This subspecies has been
found in the same beds at Pfidoli near Velka Chuchle, in the former Podoli Cement
Works in Praha 14, and in the Svarcava valley near Solopysky, &c. (for more detailed
records see Chlupac 1953). The subspecies was also found in the organo-detritic facies of
the same age in the Kotys Limestones, at Samor, near Liten, where it occurs frequently
in beds of light grey limestones with a rich trilobite fauna.

Cera1iocaris{l) coherbaria sp. nov.

Plate 12, figs. 6, 7; Plate 16, figs. 4-9

1960fl Ceratiocaris sp. nov. Chlupac, pp. 152, 157, &c.

Derivation of name. From the Latin co- and herbaria = herbaceous, indicating the common occurrence
with plant remains.

Holotype. Telson figured on Plate 16, fig. 7; ICh 126.

Type locality. ‘Volfova rokle’, Karlstejn, near Beroun, Czechoslovakia.

Horizon. Srbsko Formation, lower part (Kacak Beds), Upper Middle Devonian (Lower Givetian).
Material. Twenty-five telsons, mostly with furca.

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS 109

Diagnosis. A species referred only with reserve to the genus Ceratiocaris M‘Coy,
characterized by a diagnostic ornament on the telson and furca. The ornament is formed
on both sides of the median ridge of the telson by dense fine broken lines, the angles of
which point anteriorly, and on the lateral spines by distinct oblique lines. The telson
bears a pair of small lateral telson-spines.

Description. The only part of the shield known so far is the telson and paired lateral
spines (furca). The telson is produced as a straight central spine of medium length and is
slightly longer than the lateral spines. It tapers rapidly posteriorly at an angle of 7-10°,
and the telson head is only slightly inflated. A distinct median dorsal keel runs the length
of the telson. In well-preserved specimens a pair of lateral spines may be seen just over
half-way along the telson; the spines are slightly curved and point obliquely backwards.
The characteristic ornament of the telson consists of fine broken lines, which form
sharply pointed arches, with vertices pointing anteriorly. When dorsally coujpressed
these arches are arranged symmetrically in relation to the longitudinal keel. Only some
specimens show a row of closely spaced circular pits marking points of insertion of
lateral bristles. The lateral spines (furcal rami) are almost as long as the central spine.
They taper rapidly posteriorly at an angle of about 5-8°, while the dorsally compressed
specimens have their distal ends slightly bent inwards towards the central spine. A
longitudinal keel runs along the interior margin of the lateral spines, and some speci-
mens show a row of tiny circular pits marking the insertions of setae. The ornament on
the lateral spines consists of striae running obliquely from the interior to the exterior
margin; it is more distinct than on the telson. The striae become finer towards the
exterior margin. The angle formed by the exterior sides of both spines of the furca is
fairly constant and varies from 60° to 80°.

Dimensions (in mm.)

Holotvpe
ilCh 126)

Paratvpe A
ilCh 124)

Paratvpe B
ilCh 133)

Length of telson along midline

37

21

20

Width of proximal part of telson

6

3-3

3-3

Length of lateral spines along exterior margin

26

15-2

17

Angle between exterior margins of lateral spines

c. 65°

60°

70°

Remarks. Since only caudal appendages have so far been found their generic assignment
must remain questionable. The author only tentatively refers the specimens described
above to Ceratiocaris because the characteristic ornament of the telson differs from all
other representatives of the genus. Similar ornament may be seen in Bohemian Silurian
representatives of the group of Ceratiocaris stygia Salter, e.g. C. scharyi Barrande, which
have an ornament of pointed arches on the last abdominal segment, although this
ornament does not usually continue on to the telson. A certain analogy in the ornament
can be seen in the specimens described by Novak (1886u) as Phasganocaris piigio,
which according to Giirich (1929) represents the telson and possibly the abdominal seg-
ment of Aristozoe memoranda Barrande. The shape and spacing of the lines are, how-
ever, quite different. On the other hand the ornament of oblique lines on the lateral
spines is similar to that of a number of phyllocarids, e.g. of the genus Mesothyra Hall

110

PALAEONTOLOGY, VOLUME 6

from the family Rhinocarididae (especially of Mesothyra neptimi Hall) and of the genus
Montecaris Jux (e.g. M. antecedens Chlupac), &c. The fact that neither the carapace nor
other parts of the body except caudal appendages have so far been found suggests
that as in many other ceratiocarids only the caudal appendages were suitable for
fossilization. It is interesting to note that most of the specimens of caudal appendages
are preserved in dorsal or ventral position and only exceptionally in lateral position. The
fairly constant angle formed by the lateral spines suggests a close connexion between
furca and telson. The weight of the central spine being greater the specimens were
necessarily deposited on the sea bottom on the dorsal or ventral side, while the lateral
spines remained symmetrically extended on both sides of the telson.

C.(?) coherbaria sp. nov. is the youngest known phyllocarid from the Central Bohe-
mian earlier Palaeozoic and comes from beds where no phyllocarids have previously
been found.

Distribution. Ceratiocaris{l) coherbaria sp. nov. appears rarely in dark calcareous shales
of the Kacak division of the Srbsko Beds which form the base of the Givetian in the
Barrandian area. Both an abundant marine fauna (tentaculitids, lamellibranchs, brachio-
pods, &c.) and especially terrestrial plants occur together with this species (see Chlupac
1960; Obrhel 1961). During recent study C.(?) coherbaria sp. nov. was found in several
localities, e.g. in Karlstejn-Volfova rokle and at the locality ‘U dubu’ near the forestry,
Srbsko, Koda, and the hill Korensky vrch. For further details of localities and
accompanying fauna see the author’s stratigraphical paper (1960a).

Genus concavicaris Rolfe 1961

Remarks. The commonly used name Colpocaris Meek 1872 has been replaced by
Coneaviearis Rolfe, since the former was a junior homonym of Colpocaris von Meyer
1862 (see Rolfe 1961).

Orientation of the earapace. The orientation of the valves of the carapace is disputed in
this genus. While Meek (1875) considered the part with a prominent sinus-like incision
to be posterior, most of the later authors, e.g. Clarke (1900), Giirich (1929), Cooper
(1932), Shimer and Shrock (1944), regarded the sinus as being anterior. By analogy in
many other phyllocarids the posterior thorny projections and incisions are more promi-
nent than the anterior ones, and the conspicuous broadening of the carapace towards
the sinus in Concavicaris suggests that Meek’s original orientation is correct. This is

EXPLANATION OF PLATE 16

The specimens in Figs. 2-5 and 9 were photographed under alcohol, and those in Figs. 6-8 were coated
with ammonium chloride before being photographed.

Figs. 1, 2. Concavicaris incola sp. nov. Hady, near Brno. ICh I76a. 1, Carapace (left valve), X 1-7.

2, The same specimen photographed under alcohol, X L7.

Fig. 3. Concavicaris desiderata (Barrande). Pekarek mill, near Chotec. Paratype A, ICh 194, carapace,
x2-5.

Figs. 4-9. Ceratiocaris (?) coherbaria sp. nov. Karlstejn, near Beroun. 4, Paratype B, ICh 1236, caudal
appendages, x2-4. 5, Paratype A, ICh 124, caudal appendages, x2-4. 6, Paratype C, ICh 122, the
right lateral spine showing the sculpture, X L9. 7, Holotype, ICh \26a, caudal appendages, x2T.
8, ICh 1266, counterpart of the same specimen, x2T. 9, The same specimen, detail of the telson
showing the small lateral telson-spines, x 5-8.

Palaeontology, Vol. 6

PLATE 16

CHLUPAC, Phyllocarid crustaceans

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS

111

confirmed by the presence in a new American species of a rostral plate at the opposite
end to that with the sinus (personal communication, W. D. I. Rolfe).

Distribution. Concavicaris has so far been recorded from the North American Lower and
Upper Carboniferous only. To this genus the author here refers the species Concavicaris
desiderata (Barrande) from the Bohemian early Middle Devonian, previously regarded
as an ostracod, and C. incola sp. nov. from the Upper Devonian of Moravia. Thus the
stratigraphic range of the genus is extended to the Middle Devonian and the geo-
graphical distribution to Central Europe.

Concavicaris desiderata (Barrande 1872)

Plate 12, figs. 3-5; Plate 16, fig. 3; text-fig. 6

1872 Leperditia desiderata Barrande, p. 530, pi. 34, figs. 27, 28.

1938 Ceratiocaris sp. nov. Boucek, p. 168.

1959 ColpocarisO desiderata (Barrande); Chlupac, pp. 460, 491.

Holotype (by monotypy). NM CF1271. Figured by Barrande 1872, pi. 34, figs. 27, 28; refigured here as
text-fig. 6.

Type locality. Vavra mill at Chotec, Czechoslovakia.

Horizon. Daleje Shales, Gyroceratites gracilis Zone, Lower Eifelian, Devonian.

Redescription. Carapace broadly suboval with a distinct sinus, the length being approxi-
mately twice the width. Antero-dorsal part of the carapace pointed, with a tapering
projection in some specimens which can be interpreted as a rostral plate in natural
position. The dorsal border of the carapace is less convex than the ventral and runs in
a weak arch to the upper end of the sinus. The ventral border is strongly convex and runs
arcuately from the pointed anterior border to the postero-ventral end of the sinus. The
ventral marginal rim is indistinct and separated only by a shallow marginal line.
The rim reaches its maximum width at the ventral border, and narrows arcuately
anteriorly and posteriorly. A conspicuous postero-ventral arcuate sinus incises the
posterior of the carapace. The dorsal border forms an acute angle with the dorsal limb
of the sinus and forms a tapering postero-dorsal process which exceeds the length of the
process enclosed between the ventral border and the lower part of the sinus. The surface
of the valves is smooth, but two or three furrows are commonly present in the ventral
region, parallel to the border. These are, however, not genuine ornament but the border
lines of the opposite valve of the carapace which have been impressed through the sur-
face of the carapace during the natural shifting of the valves under pressure. Numerous
irregular longitudinal ridges are the result of compression and of secondary deforma-
tions, as their course differs in individual specimens and they are absent in better-
preserved specimens.

Dimensions (in mm.)

Holotype
(NM CF1271)

Paratvpe A
UCh 194)

Paratype B
(ICh 192)

Paratvpe C
UCh 211)

Length of valve .

19

20

21

25

Maximum width of

valve

9

10

10

13

112

PALAEONTOLOGY, VOLUME 6

Remarks. Barrande (1872) described and figured this species as an ostracod, Leper ditia
desiderata Barrande, from the locality ‘Wawrowitz’ (Vavra mill), near Chotec. Later
Boucek (1938) mentioned specimens of a phyllocarid which he considered a new species
of the genus Ceratiocaris M‘Coy from the same stratigraphic horizon (Daleje shales)
from the Pekarek mill locality, near Solopysky. The author’s collections from Pekarek
mill proved the identity of the specimens collected by Boucek with Barrande’s species
Leperditia desiderata. Barrande’s figure of this species is considerably and inaccurately

idealized. Study of the original specimen
(NM CF1271) shows a conspicuous posterior
sinus which is filled, however, with broken
fragments of the carapace. The shape of the
anterior and posterior border is also different
from that figured by Barrande, but it corre-
sponds well with the new discovery from
Pekarek mill. Otherwise Barrande’s original
is much damaged by pressure so that it does
not show detail of the surface of the valves
(text-fig. 6).

C. desiderata differs from the American
Carboniferous representatives of the genus especially in its shorter and broader carapace,
and smaller, less incised sinus, placed more postero-ventrally. From the Upper Devonian
species C. incola sp. nov. it differs in more elongated valves, less curved ventral border,
and less incised sinus. Concavicaris desiderata is the oldest representative of the genus
yet known.

Occurrence. Barrande’s original comes from a hard calcareous intercalation in the Daleje
shales from Vavra mill, near Chotec, in Bohemia. This species has recently been found
in the lowest part of the Daleje shales at the locality near Pekarek mill, near Solopysky
(Chlupac 1959, p. 460). Here it abounds in a thin calcareous intercalation with very
abundant tentaculitids, Nowakia cancellata (Richter), Styliolina clavulus (Barrande), and
other fossils. Stratigraphically this horizon is in the lower part of the Gyroceratites
gracilis Zone, Lower Eifelian, lowest Middle Devonian.

Concavicaris incola sp. nov. i

Plate 12, figs. 1,2; Plate 16, figs. 1,2
Derivation of name. From Latin incola, resident.

Holotype. Carapace figured on Plate 12, fig. 1, and Plate 16, figs. 1, 2. ICh 176a. !

Type locality. Hady, near Brno, Ruzena-quarry, Moravian Karst, Czechoslovakia. '

Horizon. Dark platy Hady-Limestones, Famennian, Upper Devonian. ■

Material. Six carapaces and damaged fragments of body and abdominal spines.

Description. Carapace suboval, deep, with a prominent semicircular sinus in the posterior
part, the length being about one and a half times the width. The maximum width is
reached in the posterior half. The antero-dorsal part of the carapace slightly pointed. I

6 5mm

TEXT-FIG. 6. Concavicaris desiderata {Bawdinde).
Holotype NM CF1271, laterally compressed
carapace (posterior part damaged).

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS 113

Dorsal border slightly convex, ventral border strongly convex. Ventral marginal rim
narrow, in the anterior part indistinct; ventral border line very fine and narrow. The
postero-dorsal projection between the dorsal border and the upper part of the sinus is
produced into a ventrally curved, beak-shaped process. It is longer and sharper than the
postero-ventral process between the ventral border and the lower part of the sinus. The
ventral border line continues to the sinus, rims its margin, and dies out just before the
dorsal border. In some specimens an indistinct line parallel to the dorsal border may
have been produced by shifting and bending of the dorsal border of the opposite valve.
The surface of the carapace is smooth.

Holotvpe

Paratvpe A

Paratvpe B

Dimensions (in trim.)

(ICh 176)

(ICh 178)

(ICh 177)

Length of valve .....

17-5

16

25

Maximum width of valve

10-5

10

15

Remarks. All the available material is flattened so that the original rotundity is not
perceptible. Fragments of abdominal spines found in the same layer are very incom-
pletely preserved, but some show longitudinal ridges.

Concavicaris incola sp. nov. differs from the American Carboniferous species C. ely-
troides (Meek), C. woodfordi (Cooper), and C. bradleyi (Meek) mainly in the greater
dorso-ventral width and lack of sculpture. Its form is more reminiscent of that of the
Upper Carboniferous C. siniiata (Meek and Worthen), the anterior region of which is,
however, even more distinctly pointed and the dorsal border less convex. C. desiderata
(Barrande) has a different length-to-width ratio, a more gently curved ventral border, and
a less incised sinus.

Occurrence. C. incola sp. nov. was found in dark Hady Limestones with calcareous shale
intercalations in the north-east wall of the big quarry ‘Ruzena’ at Hady, near Brno, in
Moravia. It occurs very abundantly in one thin layer. The accompanying ostracod fauna
confirms the Upper Devonian age, namely the lower part of the Clymenia Zone (Famen-
nian Va), i.e. the lower part of the Dasberg substage according to the German division.
Further specimens doubtfully referred to this species have been found in the northern
part of the Hady plateau in the same formation of thin bedded dark limestones of
Famennian age.

Family echinocarididae Clarke 1900
Genus montecaris Jux 1959

1959 Montecaris Jux, pp. 167, 168.

1960Z) Montecaris Chlupac, p. 639.

1960 Montecaris Jux, p. W JA.

1961 Baituganocaris Krestovnikov, p. 28.

Systematic position. The genus Montecaris was initially assigned by Jux (1959) and
Chlupac (1960Z?) to the family Ceratiocaridae of the suborder Ceratiocarina as the an-
terior region of the carapace was poorly known. After finding more completely pre-
served specimens Jux (1960) reassigned this genus to the family Rhinocarididae of the

C 1015

I

114

PALAEONTOLOGY, VOLUME 6

suborder Rhinocarina. Krestovnikov (1961) refers the specimens designated by him as
Baituganocaris to the same family, and these are doubtless congeneric with Montecaris.
The critical character for classification is the presence or absence of the dorsal median
plate which in rhinocaridids separates both valves of the carapace, which come together
at one place only. The median plate has not been established in Montecaris. In the newly
found well-preserved specimen of M. brunnensis Chlupac described below both valves of
the carapace touch each other in a simple dorsal line without any trace of median plate.
The valves are slightly separated from each other only in the anterior quarter where they
leave space for an elongated triangular rostral plate.

The lack of dorsal plate suggests that the assignment to the order Ceratiocarina is
correct. The sculpture of the carapace and the shape of caudal appendages would
correspond both to rhinocaridids and echinocaridids. The abdominal segments with
their distinct postero-lateral spines suggest a relationship with echinocaridids. In spite of
this Montecaris shows some features reminiscent of rhinocaridids, for example the whole
shape of the carapace with the conspicuous mid-posterior spines, especially in the
German specimens, suggests a relationship with, for example, the genera Dithyrocaris
Scouler and Mesothyra Hall. The shape of the Moravian specimens, however, is closer
to that of Elyrnocaris Beecher. A similar shape of carapace is not, however, unknown in
Ceratiocarina, as Galenocaris Wells proves.

It would appear that the relationship to the suborder Ceratiocarina is much closer
than to the Rhinocarina. From the present study the author thinks it better to refer
Montecaris to the family Echinocarididae of the suborder Ceratiocarina.

Distribution. The genus Montecaris was described originally from the higher Middle or
lower Upper Devonian of the Rhineland where it is represented by M. strimensis Jux
1959 and M. lehmanni Jux 1960. At the same time its representative M. antecedens Chi.
was found in the uppermost Silurian (Lochkovian) of Bohemia and M. brunnensis
Chlupac in the Moravian Upper Devonian (Chlupac 1960Z>). Caudal appendages un-
doubtedly belonging to Montecaris Jux were described and figured by Copeland (1960)
as Spathiocaris? sp. from the Upper Devonian of British Columbia, Canada. The genus
Baituganocaris Krestovnikov from the Upper Devonian of the Transvolga region in the
U.S.S.R. is evidently synonymous with Montecaris. Of five species described by Krestov-
nikov (1961) M. tatarica (Krestovnikov) seems to be the only species adequately
described and thus objectively justified. From the material known to date we may con-
clude that Montecaris had a wide geographical distribution and ranges from the upper-
most Silurian to the Upper Devonian.

Palaeoecology. All previously described records of Montecaris come from purely marine
sediments of similar character. German and Moravian species occur in a facies of thin-
bedded platy limestones with many intercalations of dark calcareous shales. The Cana-
dian specimen occurs in dark shales. The specimens from the Chugurovo Beds of the
Transvolga region come from beds of alternating shales, limestones, and siltstones. It is
obvious that Montecaris, like Ceratiocaris, found suitable life conditions in the facies of
alternating limestones and shales, i.e. in quiet-water conditions with a rich planktonic
and nektonic marine fauna (cf. Chlupac \960b). The shallow-water environment of a
reef or organo-detritic facies was evidently inimical to Montecaris.

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS

115

Montecaris bnmnensis Chlupac 1960
Plate 14, figs. 1,2; text-fig. 7

1960 Montecaris hnamensis sp. nov. Chlupac, pp. 640-2, text-figs. 3, 4; pi. 2, figs. 1,2;
pi. 3, figs. 1, 2.

Holotype. Chlupac 1960, pi. 2, figs. 1, 2. Coll. ICh. 18.

Type locality. Bedfichovice, near Brno, Moravian Karst, Czechoslovakia.

Horizon. Dark platy Hady Limestones, Famennian, Upper Devonian.

Description of new material. The newly found almost complete specimen of M. briin-
nensis from the type-locality makes it possible to augment the author’s original descrip-
tion. The new specimen (ICh 200a, b) is preserved in dorsal view and shows the carapace
and abdomen in natural position. The antero-dorsal carapace incision caused by the
arcuate truncation of the antero-dorsal part
of the valves is very noticeable. The straight
median dorsal line in the new specimen is very
clear. In the anterior quarter the valves are
slightly separated from each other. In this space
between the valves are preserved fragments of
the test that can be interpreted as a rostral plate,
but they do not distinctly project into the antero-
dorsal carapace incision. A dorsal median plate
has not been found and both valves of the cara-
pace simply meet at the midline.

Distinct nodes are not detectable on the an-
tero-dorsal part of the valves but as in the
holotype there are more massive parts of the
carapace and irregularities which are probably
due to the impression of mandibles through the
carapace. The ventral margin of the valves is
moderately arcuate but is truncate posteriorly
and forms the slightly concave posterior margin.

On the surface of the left valve an oblique stripe
is preserved in the antero-ventral region as in
the holotype, and which runs from the anterior
border in a postero-ventral direction. In addition
there is another longitudinal stripe not clearly
marked in the ventral sector of the left valve
produced probably by the dorsal compression
of the carapace. Three dorso-ventrally running,
equally spaced transverse furrows are dis-
cernible in the posterior half of the carapace at
the dorsal line on both valves, and probably represent the course of segments covered
by the carapace (text-fig. 7).

In front of the anterior border of the valves of the carapace two disconnected stripes
are indistinctly visible formed of the same material as the valves, but finely dispersed.

TEXT-FIG. 7. Montecaris brunnensis Chlupac.
Paratype B, ICh 200a, specimen dorsally
compressed.

116

PALAEONTOLOGY, VOLUME 6

The stripe in front of the right valve is striking, especially when under alcohol. It runs
subparallel to the frontal border of the carapace. By analogy with other rare examples of
phyllocarids with appendages preserved the author considers these structures to be the
remains of antennae. Articulation or other particulars cannot, however, be established
on the specimen.

Four abdominal segments are preserved outside the carapace and increase in length
backwards. The ratio of dorsal length to width in the first is 1 :4-5, in the second 1 : 3 to
1 : 3-5, in the third 1 : 2-3, and in the last the length equals the width. The outline of all the
segments is subquadrate. Short postero-lateral spines, which were well preserved in the
previously described holotype and paratype, are not well seen in the new specimen.

The caudal appendages are only partly preserved ; only the proximal part of the telson
with its broad head and the left lateral spine of the furca are preserved. The latter agrees
in shape with the caudal appendages of M. brimuensis. The characteristic telson bristles
are not preserved.

Dimensions. Paratype B, ICh 200«, b. Approximate sagittal length of whole exoskeleton 140-150 mm.
Maximum length of carapace 60 mm. Maximum width of carapace 58 mm. Length of abdominal part
of shield without caudal appendages 41 mm. Length and width respectively of abdominal segments:
first segment, 6 mm., 29 mm.; second segment, 8 mm., 26 mm.; third segment, 10 mm., 22 mm.; last
segment, 17 mm., 17-18 mm.

Remarks. The new specimen of M. brimnensis is important especially because of its good
preservation of the anterior region of the carapace. The prominent antero-dorsal incision
of the carapace is different from the anterior region of the carapace of the related species
M. lehmaimi Jux, as reconstructed by Jux (1960, text-fig. 3). On the other hand the shape
of the frontal part of the carapace corresponds well with the photographs of M. lehmanni
given by Jux (1960, pis. 142, 143) which in the antero-dorsal part also shows a similar
incision, interpreted by Jux as purely secondary. The new specimen of M. brimnensis
clearly proves the natural existence of the frontal incision of the carapace as in many
other phyllocarids.

M. brimnensis agrees with M. lehmanni in some features, e.g. in the general shape of
the carapace, broad posterior incision, longitudinal stripes in the ventral region of the
valves, and the shape of the caudal appendages. M. brimnensis differs, however, in
having a rounded postero-lateral region without spiny projections ; in the greater number
(4) of abdominal segments (M. lehmanni shows 3) ; in the much shorter postero-lateral
spines on the abdominal segments; and in having fewer bristles on the telson and
narrower lateral spines.

On the basis of new knowledge presented by Jux (1960) the comparison with M. stru-
nensis Jux may be completed by the fact that in M. brunnensis postero-lateral spiny
projections on the carapace are not developed while in M. strimensis they are distinctly
marked. A comparison of the location of nodes in the antero-dorsal part cannot be made
owing to the flattening of the Moravian material. The species M. tatarica (Krestovnikov)
agrees with M. brimnensis in the simple midline and probably also in the outline of the
posterior part of the carapace, but it has a different telson-length ; furcal-length ratio.

The traces interpreted by the author as probable remains of antennae are noteworthy.
Similar traces have so far been found only in very rare cases. They are well known and
excellently preserved, for example, in Nahecaris stuertzi Jaekel from the Rhenish Lower

IVO CHLUPAC: SILURIAN AND DEVONIAN PHYLLOCARIDS

117

Devonian (cf. Broili 1928, 1929). The preservation of the specimen of M. bnmnensis
unfortunately does not allow more detailed studies of these organs.

The three transverse lines visible on both sides of the midline in the posterior half of
the carapace probably mark the boundaries between the three anterior abdominal seg-
ments covered by the carapace valves. As these transverse furrows are shorter than the
abdominal segments and die out 7 mm. away from the dorsal line, the possibility that
they represent the remains of hinge organs cannot be excluded. The author considers the
first explanation, however, to be more probable.

Occurrence. M. bninnensis is so far known only from the Upper Devonian thin-bedded
Hady limestones at the type-locality, Bedfichovice, near Brno, in the Moravian Karst.

REFERENCES

BARRANDE, J. 1872. Systemc sihirieu dll Centre dc la Bohemc. Supplement ail vol. 1 . Text, Atlas. Prague-
Paris.

BOUCEK, B. 1938. O poloze s Ceratiocaris n.sp. v dalejskych bfidlicich u Roblina. Cas. ndr. Mas. 112,
(2), 168.

BOUCOT, A. J. 1960. Lower Gedinnian brachiopods of Belgium. Mem. Inst. geol. Univ. Louvain, 21,
281-324, pi. 9-18.

BROILI, F. 1928. Beobachtungen an TVo/iea/ra. Abli. bayer. Akad. iViss. I-IS, p\. \.

1929. Beobachtungen an neuen Arthropodenfunden aus den Hunsruckschiefern. Ibid. 253-80,

pi. 1-5.

CHLUPAC, I. 1953. Stratigraphical investigation of the border strata of the Silurian and the Devonian
in Central Bohemia. Sboniik Ustr. list. geol. 20, odd. geol. 277-380, pi. 1-5.

1955. Stratigraphical study of the oldest Devonian beds of the Barrandian. Ibid. 21 (1954), odd.

geol. (2), 91-244, pi. 1-9.

1959. Facial development and biostratigraphy of the Daleje Shales and Hlubocepy Limestones

(Eifelian) in the Devonian of Central Bohemia. Ibid. 25 (1958), odd. geol. 445-51 1, pi. 1-5.

1960rt. Stratigraphical investigation of the Srbsko Beds (Givetian) in the Devonian of Central

Bohemia. Ibid. 26 (1959), odd. geol. (1), 143-85, pi. 1-3.

19606. Die Gattung Montecaris Jux (Crustacea Phyllocarida) im alter Palaozoikum der

Tschechoslowakei. Geologie, 9, H. 6, 638-49, pi. 1-3.

CLARKE, J. M. 1900. Phyllocarida in eastman, c. r., zittel, k. a. von. Textbook of Palaeontology, 1,
653-6. London.

1913. Ibid. 1, 748-54.

COOPER, c. L. 1932. A crustacean fauna from the Woodford Formation of Oklahoma. J. Paleont.
6, 346-52, pi. 52.

COPELAND, M. J. 1960. Ncw occurrcnccs of Ceratiocaris and Ptychocaris (Phyllocarida) from the
Canadian Arctic. Bull. geol. Surv. Can. 60, 49-54, pi. 8-10.

FRiTSCH, A. 1907. Miscellanea Palaeontologica. 1, Palaeozoica. Prague.

gOrich, g. 1929. Silesicaris von Leipe und die Phyllokariden iiberhaupt. Mitt, miii.-geol. {St) Inst.
Hanib. 2, 21-90, pi. 1.

JUX, u. 1959. Phyllocariden-Reste aus dem oberen Mitteldevon der Bergisch-Gladbach-Paffrather
Mulde. Palaont. Z. 33, 166-71, pi. 18.

1960. Montecaris lehmaiini, a new crustacean from the Rhenish Devonian and the problem of its

systematic position. J. Paleont. 34, 1129-52, pi. 142, 143.

KRESTOVNIKOV, V. N. 1960. Pliyllocarida in Osnovy paleontologii, tsclilenistonogie, trilobitoobraznye
i rakoobraznye, 425-9. Moscow.

1961. Novye rakoobraznye fillokaridy paleozoja ruskoj platformy, Urala, Timana i Donbasa.

Trudy geol. hist. Akad. nauk SSSR. 52, 1-67.

MEEK, E. B. 1875. Report on some of the invertebrate fossils of the Waverly Group and Coal Measures
of Ohio. Rep. geol. Surv. Ohio, 2 (2), 317-21, pi. 18.

118 PALAEONTOLOGY, VOLUME 6

MOORE, R. c. (ed.) 1961. Treatise on Invertebrate Paleontology, Part Q, Arthropoda 3: Crustacea,
Ostracoda. New York.

NOVAK, o. 1885u. Remarques sur le genre Barrande. S.B. bbhn. Ges. Wiss. 239-43, pi. 1.

■ — ^ — • 18856. Nouveau Crustace phyllocaride de I’etage F-fa en Boheme. Ibid. 343-7, pi. 1.

1886fl. Note sur P/mgn/wcnra, nouveau Phyllocaride de Tetage F-fa, en Boheme. Ibid. 498-501,

pi. 1.

1 8866. Zur Kenntniss der Fauna der Etage F-fi in der palaeozoischen Schichtengruppe Bohmens.

Ibid. 660-85, pi. 1, 2.

OBRHEL, J. 1961. Die Flora der Srbsko-Schichten (Givet) des mittelbohmischen Devons. Sbornik
Ustr. list. geol. 26 (1959), odd. paleont. 7-46, pi. 1-14.

PERNER, J. 1916. O novych phyllocaridech z pasma F-fj. Rozpr. H. ceske Akad. 25, No. 40, 1-8, pi. 1.

1919. Ueber neue Phyllocariden aus der Bande F-E. Bull. int. Acad. Prague, 21 (1917), 225-30,

Pl- 1-

PRANTL, F. and PRiBYL, A. 1948. Revision of the Bohemian Silurian Eurypterida. Rozpr. geol. Ust.
10, 1-116, pl. 1-8.

ROLFE, w. D. I. 1961. Concavicaris and Quasicaris, substitute names for Colpocaris Meek, 1872, and
Pterocaris Barrande, 1872. /. Paleont. 35, 1243.

SFUMER, H. w. and SHROCK, R. R. 1944. Index fossils of North America. New York.

STRAELEN, V. VAN and SCHMITZ, G. 1934. Crustacea Phyllocarida ( = Archaeostraca). Fossilium
Catalogus. I: Animal ia, 64, 1-246.

SVOBODA, J. (ed.) 1958. Prager Arbeitstagung iiber die Stratigraphie des Silurs und des Devons.
Beschluss. Ceskoslovenskd Akademia ved, Praha.

IVO CHLUPAC

Ustfedni ustav geologicky CSSR,
Malostranske nam. 19,
Praha I, Czechoslovakia

Manuscript received 5 April 1962

THE COMPOSITION OF PYRITOSPHAERA
BARBARIA LOVE 1957

by LEONARD G. LOVE

Abstract. A bulk sample of Pyritosphaera barbaria Love 1957 is shown by microchemical analysis, following
other indications, to be dominantly organic in composition.

Pyritosphaera barbaria was isolated (Love 1957) by removal of the pyrite by nitric acid
from pyrite microspheres of the Pumpherston Shell Bed and adjacent strata of the
Scottish Lower Carboniferous Lower Oil Shale Group. The residual bodies, which
average about S pin size and are of much the same diameter as the spheres from which
they come, are spherical and microgranular in appearance. They were regarded as being
organic on the basis of their pale brown colour, translucency, isotropic character, and
chemical inertness except to further oxidation or combustion. Further they were regarded
as the remains of micro-organisms on the basis of the regularity of their form, coupled
with their great abundance in an environment in which such material might be expected
to flourish. The suggestion that they might have had an active significance in the forma-
tion of the sulphide is not of immediate concern here. Similar forms were obtained by
Love (1962fl, b) from pyrite spheres of Upper Palaeozoic and Mesozoic rocks and much
detail of the manner of occurrence was noted. Somewhat similar forms were recorded by
Love and Murray from pyrite spheres from Recent sediments.

Direct analysis of the material of P. barbaria of Carboniferous age has been made
difficult both by problems in adequately concentrating the pyrite spheres from the rock
and, of greater importance, in obtaining samples in which the pyrite spheres, and thus
the P. barbaria bodies, were free from other organic material which often encloses them,
A sample recently examined, however, conformed with the latter requirement as far as
detailed microscopic examination of the pyrite and of the residues could show, and
yielded a good separation of pyrite spheres. It came from Namurian marine shale of
Ri age, from River Noe, Derbyshire, England.

The pyrite was dissolved with brominated concentrated nitric acid acting for one hour
at room temperature and the residue consisted dominantly of P. barbaria. Preliminary
examination of a portion by emission spectrograph between copper electrodes indicated
at least 35 per cent, of carbon by the method developed by Dennen (1957), whose
observation that a substantial upward correction may be necessary is amply borne out.
Electron probe micro-analysis of the spherical bodies in another part of the sample
indieated (to a 1 per cent, limit) the absence of iron (which might be expected as insoluble
ferric oxide), and of sulphur, silica, calcium, and other metallic elements. Carbon cannot
be detected by this method. On a third portion normal microchemical analysis indicated
56 per cent, carbon. Based on bulk analysis of Carboniferous ‘kerogen’ (Down 1939) this
might represent as much as 80 per cent, organic matter but perhaps as low as 70 per cent,
if the oxidizing acid had removed any organic material, as indeed is indicated by the
C:H ratio being 15-7:1 for the oxidized sample compared with that of 11-3:1 for the

[Palaeontology, Vol. 6, Part 1, 1963, pp. 119-20.]

120

PALAEONTOLOGY, VOLUME 6

unoxidized material quoted below. The balancing weight of the sample, proportionately
less by volume, was observed to consist of heavy minerals while the P. barbaria material
was observed to be only a little heavier than water. Infra-red spectroscopy showed the
presence of undoubted organic compounds, since C-H bonds, carbonyl groups, and
possibly N-H groupings were recognized.

Part of a parallel original unoxidized sample of pyrite spheres was shown by micro-
chemical analysis to consist of approximately 85 per cent, pyrite (40-9 per cent. Fe, 45T
per cent. S) and 3-4 per cent, carbon by weight. The latter perhaps represents 4-25 per
cent, of organic material by weight. Heavy minerals and some insoluble residue from the
use of hydrofluoric acid were seen to be present (giving 1 1 -2 per cent, ash) but no visible
organic matter. A ratio of organic matter to pyrite in the spheres of 1 : 20 by weight is
therefore given.

This demonstration of the organic composition of P. barbaria puts the argument that
these are remains of micro-organisms occurring in situ in the pyrite on a surer basis but
is not in itself proof. The data will be of value for comparison with other sediments but
this work is not complete and will be discussed elsewhere (Love, in the press).

Acknowledgements. This work was carried out while the author was on a U.S. A.E.C. grant at Yale
University, on leave from the University of Sheffield. The rock material was kindly made available by
the Peabody Museum, Yale University. Emission spectrography was carried out by Professor W. H.
Dennen in the Cabot Laboratory, M.I.T. ; the electron probe micro-analysis was carried out by Dr. H.
Adler at the U.S. Geological Survey, Washington ; and the infra-red spectroscopy by Professor W.
Lwowski of Yale University Chemistry Department. To all these the author’s thanks are due. Micro-
chemical analysis was carried out by the Schwarzkopf Microanalytical Laboratory, New York.

REFERENCES

DENNEN, w. H. 1957. Spectrographic determination of carbon in sedimentary rocks by using direct-
current arc excitation. Spectrochim. Acta, 9, 89-97.

DOWN, A. L. 1939. The analysis of the Kerogen of Oil Shales. J. Inst. Petrol. 25, 230-7.

LOVE, L. G. 1957. Micro-organisms and the presence of syngenetic pyrite. Quart. J. geol. Soc. Land.
113, 429-40.

1962u. Biogenic primary sulfide of the Kupferschiefer and Marl Slate. Econ. Geol. 57, 350-66.

\961b. Eurther studies on micro-organisms and the presence of syngenetic pyrite. Palaeontology,

5, 449-59, pi. 63, 64.

Pyrite spheres in sediments. The biogeochemical cycle of sulfur isotopes: Symposium, ed. M. L.

Jensen, Yale University. In the press.

and MURRAY, J. w. 1963. Biogenic pyrite in Recents ediments of Christchurch Harbour, England.

Amer. J. Sci. In the press.

LEONARD G. LOVE

Department of Geology,
University of Sheffield,

St. George’s Square, Sheffield 1

Manuscript received 21 May 1962

SOME UPPER TREMADOCIAN GRAPTOLITES

FROM NORWAY

by NILS SPJELDN^S

Abstract. New material of Upper Tremadocian graptolites from the Oslo Region, preserved in full or half
relief in pyrite, is described. Bryograptus ramosiis Brogger 1 882 is shown to have distinct bithecae. ‘‘Didymograptiis'
kiaeri Monsen 1925 is also shown to have bithecae and is made the type of the new genus Kiaerograptus.
Adelograptiis bidmani sp. nov. is a peculiar species, which might have affinities both with the anisograptids and
the leptograptids and might be intermediate between them. The presence of bithecae in some graptoloid-like
forms in the Upper Tremadoc suggests that the presence of graptoloids in beds older than the Arenig might be
questionable. The material present might indicate that a number of graptolite lineages are more complex than
previously assumed, and that several lineages in the dichograptids and other graptoloid groups originated
directly and independently from the dendroids.

Because of the bad state of preservation of the material, the details of the structure of
most Upper Tremadocian graptolites are virtually unknown. The specimens are generally
preserved as mineralized films, and only the number of branches, angle of bifurcation,
and the gross outline of the thecae can be observed.

The author recently succeeded in finding some specimens of Upper Tremadocian
graptolites from the Oslo Region which are preserved in full or half relief in pyrite. The
specimens come from the Ceratopyge Shale (Zone 3 a^) and are found in the upper part
of this formation, about 0-4-1 -8 m. below the Ceratopyge Limestone (Zone 3 ay), in
two road sections at Slemmestad, about 20 km. SW. of Oslo.

The general stratigraphy, and an outline of the geology of the area is given by Stormer
(in Holtedahl and Dons 1960, pp. 11-24, 45-47). One locality is about 300 m. ENE. of
Stormer’s locality 4 (loc. cit., text-fig. 25), and the second is along the same road, about
1| km. to the SW. The horizon is rich in graptolites, but only a few of them are preserved
in full relief.

Parts of the specimens preserved in full relief are sometimes distorted and swollen
in a rather peculiar way which is interpreted as a result of the pyritization. The specimens
were probably originally filled with, or replaced by, an iron-sulphide gel (possibly hydro-
troilite or melnichovite); later the sulphide gel crystallized into pyrite, and the specimens
became compressed by the diagenesis of the sediment. In the cases where the quantity of
the gel was small, or where it was very rich in water, the specimens were compressed to
mineral films. This is the usual preservation of the Tremadocian graptolites from the
Oslo Region, and several other areas. In some few specimens, the sulphide content of the
gel was sufficient to give an exact replica in massive pyrite of the specimens, or at least
one in half relief. In some cases again the gel seems to have swollen, probably due to high
osmotic pressure, and the fossils became swollen and distorted. This should be borne in
mind when working with pyritized materials, as the process of fossilization may have
caused quite considerable changes in the morphology of the specimens, and may make
statistical, biometric work on such graptolites rather difficult.

In many specimens the pyrite has been oxidized, and the specimens are preserved as

[Palaeontology, Vol. 6, Part 1, 1963, pp. 121-31, pi. 17, 18.]

122

PALAEONTOLOGY, VOLUME 6

casts partly filled with iron oxides. These specimens are studied on, and photographed
from latex moulds. The moulds were made from latex emulsion stained with Indian ink
and were whitened with ammonium chloride for photography.

The few specimens preserved in full relief reveal a number of details, not previously
observed, especially the presence of bithecae in some specimens. In Bryograptus ramosus
this was expected, even if it had not previously been possible to demonstrate their
presence. It was more surprising to find bithecae in " Didymograptus' kiaeri Monsen,
which superficially looks like an extensiform didymograptid. Based on the early strati-
graphic occurrence of this species, and some structural features (especially the long nema
and the oblique sicula), the author suspected it was not an ordinary didymograptid.
Bulman (1941, 1950) had also suggested that this species might be related to the Aniso-
graptids, and this view has gained strong support from the material presented here.

A new genus, Kiaerograptiis, is made to accommodate the two-branched, extensiform
Anisograptids with distinct bithecae.

Acknowledgements. The author is deeply indebted to Professor O. M. B. Bulman, Cambridge, for advice
and encouragement in preparation of this paper, and to Dr. Henningsmoen for having read the manu-
script critically. The photographs were made by Faculty photographer B. Mauritz and the author.

Bryograptus ramosus Brogger 1882

Plate 17, figs. 6-9; text-fig. 1

1882 Bryograptus ramosus Brogger, p. 37, pi. 12, fig. 21.

1925 Bryograptus ramosus Brogger; Monsen, pp. 160-2, pi. 1, fig. 9, text-figs. 3a-c.

1954 Bryograptus cf. ramosus Brogger; Bulman, p. 34, pi. 4, fig. 9.

Description. Some specimens presumably belonging to this species are preserved in full
relief, and show distinct bithecae, situated alternately on both sides of the branches.
Because of this pattern of budding, the apertures of the autothecae also occur alternately
in two rows, instead of in one single row, which is usually the case in Graptoloid
rhabdosomes.

The sicula is long, and needle-shaped, and the budding of the proximal part is easily
seen in the best specimens (PI. 17, fig. 8). The rhabdosome is initially rather elongate,
and in at least one specimen (PL 17, fig. 7) there are definitely three primary branches.

EXPLANATION OF PLATE 17

All specimens belong to Paleontologisk Museum, Oslo, Norway. The specimens shown in figs. 3, 4, 7,
8, 9 are photographed from latex casts coloured with Indian ink and coated with ammonium chloride.
The specimens shown in figs. 1 and 2 are coated with ammonium chloride, and the ones shown in
figs. 5 and 6 are photographed by the maximum reflection method.

Figs. 1-5 Kiaerograptiis kiaeri (Monsen). 1, Distal part of a branch showing bithecae, PMO 72833a;
X 13. 2, 3, Two proximal parts, PMO 72833b (2) and PMO 72834a; X 10. 4, Specimen showing
proximal part in somewhat oblique view, PMO 72834b; X 10. 5, Lectotype, the specimen figured by
Monsen (1925, text-fig. 5, pi. 2, fig. 16), PMO 60212a; x4. Specimens 1-4 from Slemmestad, 20 km.
SW. of Oslo; 5 is from Stensberggaten in Oslo; all from Zone 3 a^.

Figs. 6-9. Bryograptus ramosus Brogger. 6, Lectotype, a large rhabdosome from Zone 3 a)S at Vest-
fossen. Biker (70 km. W. of Oslo). Probably the specimen figured by Brogger (1882), pi. xii, figs. 21,
21a; PMO 72829; X 2. 7, 8, Two proximal parts of young rhabdosomes. Zone 3 a/3, Slemmestad.
7, PMO 72831 ; X 9. 8, PMO 72832; X 7. 9, Fragments of two branches showing bithecae; Zone 3 a/S
Slemmestad; PMO 72830; x20.

Palaeontology, Vol. 6

PLATE 17

SPJELDNiElS, Upper Tremadocian graptolites

NILS SPJELDN/ES; UPPER TREM ADOCI AN GRAPTOLITES

)23

There are probably several different Bryograptids in the Upper Tremadocian of the
Oslo Region. In the present material there are a number of different types, which vary
as to size of the branches, and shape of rhabdosome. Monsen
(1925, pp. 162-5, pi. 1, figs. 10-1 1, text-fig. 4a-Z)) described
a new species from this horizon in Oslo, but my studies do
not quite agree with the description given. The specimens
discussed here as B. ramosus agree with the lectotype (PL 17,
fig. 6) both in the thecal measurements, so far as they can
be identified, and in the elongate rhabdosome with infrequent
branching in the proximal part. Other specimens, including
some of those referred to B. ramosus by Monsen (1925) have
more frequent branching, and more rapidly expanding rhab-
dosomes.

It was anticipated by Bulman (1941, p. 106, 1954, p. 34)
that this species had three primary branches, and therefore
was a true Bryograplus. This is quite evident from the present
material.

The real number of branches is known only in a few
species of Bryograptus, and as far as the author knows
bithecae have been recorded only in B. ramosus. It is pos-
sible that a number of the species now referred to Bryo-
graplus do not have bithecae, and some might have only two
primary branches.

KIAEROGRAPTUS gen. IIOV.

Diagnosis. Dendroid graptolite genus, probably referable to
the family Anisograptidae. Sicula of graptoloid type, rhab-
dosome consisting of two branches of equal width, in one
plane (extensiform type of branching). Distinct bithecae
placed alternately on both sides of branches. Autothecae
resembling the dichograptid type. Initial budding of den-
droid type, except that the bithecae related to the early auto-
thecae of each branch appear to be missing.

Type species. Didymograptiis kiaeri Monsen 1925, from the Upper
Tremadoc (Zone 3 a^S) of the Oslo Region.

Remarks. The genus is at present monotypic, and its distri-
bution is therefore identical with that of the type species.
For discussion of affinities, see remarks on K. kiaeri.

TEXT-FIG. 1 . Bryograptus
ramosus Brogger, from the
Ceratopyge Shale, Zone 3 a^,
at Slemmestad, Oslo Region,
Norway. A young rhabdo-
some in full relief, showing
three primary branches,
bithecae, and rather late
secondary branching. The
specimen is slightly distorted
from excessive pyritization.

PMO 72831; X 13-5.

Kiaerograptus kiaeri (Monsen 1925) comb. nov.

Plate 17, figs. 1-5; text-fig. 2

1925 Didymograptiis kidri Monsen, pp. 172-5, pi. 2, figs. 9-10, 12-14, 16, pi. 4, figs. 6-8,
text-fig. 5a-c.

1925 Didymograptiis kidri var. regidaris Monsen, pp. 175-6, pi. 2, figs. 11, 15.

124

PALAEONTOLOGY, VOLUME 6

Type data. The lectotype, selected now by Monsen, in miiseo, is PMO 60212a, from Zone 3aj3, subzone
b, at Stensberggaten in Oslo. The lectotype of D. kiaeri regidaris is PMO 60219, from the same horizon
and locality.

Material. In the present material there are more than 100 specimens most of which are compressed.
About twenty-five specimens and fragments are preserved in relief, and seven of them show the
proximal part in relief. The original types of Monsen, and about fifteen topotypes have also been used.

Description. The rhabdosome consists of two horizontal branches, up to more than 2-5
cm. long each. Because most of the specimens are fragmentary or too densely crowded,
it is difficult to give exact measurements for the length of the branches. The branches are

TEXT-FIG. 2. Kiaerograptns kiaeri (Monsen) from the Ceratopyge Shale, Zone 3 a^S,
at Slemmestad, Oslo Region, Norway. Somewhat diagrammatic drawings. A, Speci-
men in obverse view; about X 14. b. Proximal part of a distorted specimen showing
how both thU and th2^ (or rather the stolotheca from which th2^ buds) originates
high up on the sicula; based on specimen PMO 72834b, Plate 17, fig. 4; about X 14.

1-5 mm. wide. The autothecae are short, straight, and oblique with rather short overlap
in the distal parts of the branches, and curved downwards with more overlap in the
proximal parts. There are about 11 to 12 autothecae in 10 mm., and they are about
0-65 mm. in longer diameter. The bithecae are placed alternately on each side of the
branches. They are somewhat irregularly cylindrical (about 0-3 mm. in diameter) and their
apertures are higher up than those of the autothecae.

The sicula is long, and inclined, often considerably so. The inclination seems always
to be towards the second (thF) branch, but since it is difficult to tell which side is the
obverse in most of the compressed specimens, and there are few uncompressed ones, this
cannot be stated without some reservation. The nema is long (more than 8 mm. in most
specimens).

The budding of the initial part of this species differs considerably as could be expected,
from that found in Didymograptus. Only specimens showing the obverse views of the

NILS SPJELDN/ES: UPPER TREM ADOCI AN GRAPTOLITES

125

proximal part are well preserved, and all attempts to prepare the reverse side from them
have been in vain. The reverse side is therefore known only from a distorted specimen
(PI. 17, fig. 4), and a fragmentary specimen (PI. 17, fig. 3), where the left part of the
proximal portion is broken away. Owing to this and the sources of error introduced by
the preservation, the interpretation given must be regarded as tentative (cf. text-figs.
2a, b).

Discussion. The type of budding is more related to that found in the Dendroids, as
described by Bulman (1936, text-figs. 24o, b) but it differs in that the first theca gives rise
to two other primary thecae very high up on the sicula. One of them (X in text-fig. 2) is
interpreted as the first bitheca, and the other as the first theca of the second branch. (In
order to conform with Bulman’s figure; the term X should be bi 1; first theca (P), should
be P+P+thP, &c. Since the present material does not allow such a delicate anatomic
discrimination, the terminology has been somewhat simplified.)

Except for the first (X), bithecae are not found at the first thecae. Normally they do
occur at the third or fourth autothecae on each branch. There are no specimens in which
the first bithecae can be seen in both branches simultaneously because of the fragmentary
nature of the specimens, but in the branches where this feature can be clearly observed,
five specimens show the first bitheca after the third autotheca (such as in text-fig. 2a),
and three specimens after the fourth theca. The number of observations of this kind is
higher than the total of well-preserved proximal parts owing to the fact that the proximal
part is compressed in a number of specimens where the branches are preserved in relief.

The sicula is long and conical, and where the branches originate it bends slightly
towards the second branch. In this feature the distal part of the sicula resembles the
distal part of the first thecae in Didymograptiis, and the usual appearance of the
specimen (PI. 17, fig. 5) is at first glance puzzling, because the sicula resembles the first
thecae, and the X resembles the sicula in externally similar Dichograptids. Other speci-
mens (PI. 17, fig. 4) reveal the three thecae originating from the proximal part of the
sicula.

A certain amount of variation is found in this species, but it is difficult to discriminate
between real variation, and changes due to different preservation. In some specimens the
theca X is almost as long as the sicula, and especially in the small specimens there are
distinct slits between the distal parts of the thecae (cf. Monsen 1925, pi. 2, figs. 15-16,
text-fig. 5a). This is less well developed in old (large) specimens, and in the distal part of
the branches.

Remarks. The only species referred to Kiaerograptus at present is the type species, but it
possibly includes several species now referred to Didymograptiis. Among the Trema-
docian and Lower Arenigian didymograptids which might be suspected to have bithecae,
many belong to the geometricus-iy^t of branching, with two straight or slightly curved
branches meeting at about 120-150° {D. pritchardi T. S. Hall, D. taylori T. S. Hall,
D. klotschichini Obut 1961, and Didymograptiis sp. of Bulman 1954, pi. 5, figs. 7-9).
D. primigenius Bulman shows the increase in width of branches found in the D. extensus
group, and might be a real graptoloid. More similar to K. kiaeri in rhabdosome shape
are D. abnormis Hsii, D. novas Berry 1960 and D. latiis T. S. Hall, the latter of which is
found in Zone 3 ba (the basal Arenig) in the Oslo District.

126

PALAEONTOLOGY, VOLUME 6

D. norvegiciis Monsen (1925, pp. 176-7, pi. 2, figs. 6-7; pi. 4. figs. 4-5; text-fig. 6) is
another Tremadocian species, from the same horizon as K. kiaeri. Like Tetragraptus
kolderupi Monsen it is found only at the type locality in Oslo (Stensberggaten), which
is also the type locality for Triograptus osloeusis Monsen. All these three species have the
same size and shape of the sicula, the same width of the branches, and the same type and
size of thecae. The latter are easily identified, being widely conical, and apparently loosely
connected. In the type material there are, besides several hundred branch fragments,
about ninety-five good specimens of Triograptus osloeusis, five to six of D. norvegicus,
and two to three of T. kolderupi. It is impossible to separate these three species on branch
fragments only, but very easy to discriminate between them and all other graptolites in
the same horizon.

This might indicate either that they all belong to one species with a somewhat erratic
mode of branching, or that they belonged to a very rapidly developing lineage. The latter
was the view held by Monsen (1925, p. 171). The presence of specimens with abnormal
budding in Triograptus os/oeusis, such as the specimen figured by Monsen (1925, pi. 3,
fig. 7), where the third branch might not be primary, but seems to bud from the second
theca in one of the other branches, might suggest the former hypothesis. It is also possible
that the peculiar thecal structure made the branches brittle, and D. norvegicus may only
be specimens of T. osloeusis from which one branch has been broken off. This problem
will, however, have to be studied in more detail on better material, and at present it may
only be concluded that the three species in question evidently form a closely related, and
possibly isolated, group.

Monsen (1925, pp. 175-6, pi. 2, figs. 11, 15) distinguished a separate variety of K. kiaeri
(regidaris). After studying the type material, and the new material described here, the
author is inclined to regard the specimens referred to this variety as falling within the
normal limits of variation of K. kiaeri.

In spite of having only two primary branches, this genus seems to fit conveniently into
the family Anisograptidae. This family is somewhat heterogeneous, embracing all few-
branched dendroids, even if their number of primary branches is unknown, or less than
three. The asymmetrical sicular part of K. kiaeri suggests that it might have developed

EXPLANATION OF PLATE 18

All specimens belong to Paleontologisk Museum, Oslo, Norway. The specimens shown in figs. 2, 8,
and 9 are photographed from latex casts coated with ammonium chloride. The other specimens are
photographed directly, without coating.

Figs. 1-10. Adelograptus? bidmani sp. nov. 1, Proximal parts of two specimens, the upper one a left-
handed specimen in obverse view, and the lower one a right-handed specimen in reverse view
(text-fig. 3a is based on the lower specimen); PM072835b-c; X 13. 2, Distal branch showing ‘Lepto-
graptoid’ thecae; PMO 72836c; X 6, 5. 3, Proximal part, ‘right’ specimen, reverse view; PM072835g;
X 13. 4, Proximal part, ‘left’ specimen, reverse view; PMO 72835h; X 13. 5, Proximal part, ‘right’
specimen, obverse view; PMO 72835i; X 13. 6, Proximal part, ‘right’ specimen, obverse view, having
only the first bitheca and thU; PMO 72835e; X 13. 7, Proximal part, ‘right’ specimen, obverse view
showing the small thU; PMO 72835d; x 25. 8, Proximal part, ‘right’ specimen, reverse view, with
a constriction of the proximal part of the sicula (prosicula?); PMO 72836a; x 21. 9, Proximal part,
‘right’ specimen, obverse view, showing sicula, first bitheca and thU. The peculiar fringes on the
aperture of the sicula might be due to pyritization; PMO 72836b; x21. 10, Proximal part of two
adult specimens; the upper one a ‘left’, and the lower, the holotype, a ‘right’; PMO 72835f and
PMO 72835a (holotype); x25.

Palaeontology, Vol. 6

PLATE 18

SPJELDNiElS, Upper Tremadocian graptolites

NILS SPJELDN/ES: UPPER TREM ADOCIAN GRAPTOLITES 127

from originally three-branched forms, and the genus is therefore, at least provisionally,
referred to the Anisograptidae.

Adelograptus? bulmani sp. nov.

Plate 18, figs. 1-8; text -figs. 3, 4

Type data. The holotype is PMO 72835a, a rhabdosome with five thecae preserved in half relief in
pyrite. It is from the upper part of the Ceratopyge Shale (Zone 3 a^) in a road section in Bodalen, about
1 -5 km. WSW. of Slemmestad, in the Oslo District, Norway. On the same slab as the holotype there are
several other specimens of this species in various stages of development, some of which are figured in
this paper. They are preserved in pyrite, in half or full relief.

Material. About fifty specimens, mostly proximal parts, preserved on two small shale slabs from the
type locality.

Diagnosis. Species probably referable to Adelograptus, having the system of initial bud-
ding and general shape of rhabdosome found in that genus. The sicula is very long, and
the long, overlapping thecae have almost leptograptoid apertures. Only two branches
have been observed; if further branching occurs, it is irregular and in the distal parts of
the rhabdosome. No bitheca except the first one has definitely been ascertained. Left-
and right-handed forms occur.

Description. The rhabdosome is small, consisting of two horizontal to slightly reclined
branches. In most of the specimens in which the sicula is preserved, the branches are
short, three or four thecae on each as a maximum. There are a few specimens with longer
branches, up to six thecae.

The thecae are long, gradually tapering and with a considerable overlap. Three thecae
are observed in a cross-section of a branch. The thecae occur alternately on both sides
of the branch, and cross in their initial part, just after budding. The apertures of the first
thecae on each branch are curved slightly down, but subsequently are directed distally.
In the specimens preserved in half relief, most of the thecae seem to be of the dichograptid
type, or intermediate between this and the leptograptoid. Among the specimens pre-
served in full relief, there are some branch fragments, presumably belonging to this
species, which appear to be more like the leptograptoid type of thecae (PI. 18, fig. 2).

The proximal part is preserved in an astonishing number of the specimens, in fact
most of the material seems to consist of immature specimens, showing the sicula, and
only some few thecae (text-figs. 3-4). In some specimens (PI. 18, fig. 8) a well-defined
prosicula-like structure is seen. This feature is not consistently found, and might be due
to the capricious effects of the pyritization. The sicula is long and narrowly conical,
almost cylindrical. The first theca (P) buds from the proximal part of the metasicula,
follows it until about half-way down, and turns at right angles to the sicula. The
proximal part of the first theca is irregularly cylindrical. It is also remarkable in budding
alternately from the right (normal) and left side of the sicula. The ‘left’ specimens are
fewer in number than the normal ones, but because of the difficulty in determining this
feature with absolute certainty, no exact ratio between the two forms can be given. It
appears to be about 1:3 or 1:4.

Subsequent budding seems to be somewhat different in the ‘left’ and normal specimens,
but because of the lack of ‘left’ material preserved in reverse view (only two specimens)
this could not be definitely stated with the present material.

128

PALAEONTOLOGY, VOLUME 6

In addition to these thecae there is a special one, which buds from before the origin
of and is interpreted as the first bitheca.

In the branches, bithecae have not been observed. This does not exclude their presence,
because a number of the specimens are broken at the critical points (PI. 18, fig. 7), and in

i \

TEXT-FIG. 3. Adelograptiisl biilmani sp. nov. from the Ceratopyge Shale, Zone 3 at
Slemmestad, Oslo Region, Norway, a. Proximal part of a ‘right’ specimen (drawing
based on specimen PMO 72835b, Plate 18, fig. 1); about X 30. b. An obverse view
(drawing based on specimen PMO 72835d, Plate 18, fig. 7); about x30. S, sicula; Te,

first bitheca.

others diffuse swellings of the branches are observed both where bithecae are to be expec-
ted, and in other places. These swellings are in most cases probably due to preservation
(pyritization), but might in some cases be due to bithecae. Such suspect structures are
generally found in the more distal parts of the branches, but have occasionally also been
observed in connexion with the early thecae (cf. the extreme left part of the lower
specimen in PI. 18, fig. 7). The first bitheca is always large and distinct, and if other
bithecae are present, they must be considerably smaller and less developed.

The branching of the rhabdosome is also difficult to ascertain. In A. Inmnebergensis
the second-order branching generally takes place after the second or third thecae. A great

NILS SPJELDN^S; UPPER TREM ADOCI AN GRAPTOLITES 129

number of specimens of ? buhnani with short branches do not show any second order
branching. This indicates that if second-order branching was present, it took place
after the fifth theca, or by lateral budding in branches longer than five theca. None of
the longer branch fragments ascribed to this species show branching, but in the same
beds there is what is believed to be another species of Adelograptus which differs from
bidmani in having somewhat thinner branches, and shorter, more conical sicula. This
species, which is less well preserved than the material of bidmani, shows second-order
branching rather regularly at the fourth or fifth thecae. Unless the preservation is very
good, it might be difficult to discriminate between distal branches of these two species.

TEXT-FIG. 4. Adelograptus"! biilmaiii sp. nov. from the Ceratopyge Shale, Zone 3 a/3,
at Slemmestad, Oslo Region, Norway, a. The proximal part of a ‘left’ specimen in
reverse view (drawing based on specimen PMO 72836a, Plate 18, fig. 8); about x 30.

B, Proximal part of ‘right’ specimen in obverse view (drawing based on specimen
PMO 72836b, Plate 18, fig. 9); about X 30. PS, prosicula; S, sicula; Te, first bitheca.

Remarks. The affinities of this species are somewhat obscure. It superficially resembles
a primitive Leptograptid in the thecal budding in the proximal part, in the direction
of the thecae (except the first ones), and in the general shape of the rhabdosome. It differs
from the later Leptograptids in the almost cylindrical metasicula, which is regarded
as a dendroid feature (Kozlowski 1960), in having the primary branches diverging
near the middle of the metasicula instead of near the aperture, in the presence of the first
bitheca, and in the existence of ‘left’ and ‘right’ forms.

There is a suggestion (text-fig. 4) that th2^ arises from thU and, if this should prove
to be correct the type of development, differs fundamentally from any of the types
described by Bulman (1955, p. V56).

A closer examination of the initial and distal budding suggests that it is closely related
to that in Adelograptus (cf. Stubblefield 1929, text-figs. 2, 4, 6, and especially Bulman
1941, text-fig. 1) except that the bithecae (other than the first one), are apparently
missing.

0 1015

K

130

PALAEONTOLOGY, VOLUME 6

It is possible to imagine a development of the leptograptoid type of budding directly
from that found in this species, but until intermediate forms are found and much more
detailed studies are made, this hypothesis could not be seriously considered. If, however,
the distal bithecae really are missing in A.? buJmani, it might represent a transitional
form between the Dendroids and Graptoloids such as Kiaerograptus. As mentioned
above, it is difficult to show this and other important anatomical features beyond doubt,
and the species is therefore, with some doubt, left in Adelogmptus for the time being.
There can be little doubt that it is a descendant of one of the typical representatives of
that genus.

GENERAL REMARKS

The presence of bithecae in virtually all upper Tremadocian species which are sufficiently
well preserved to show this feature clearly, might indicate that there were few if any
graptoloid graptolites before the Arenig. Most Tremadocian graptolites could either
be referred to the dendroids, or they are preserved only as mineralized films, and their
alleged graptoloid affinities are based only on their astogenic resemblance to various
Dichograptids. The discovery of a two-branched, extensiform dendroid indicate that
such criteria might not be valid in all cases, and in order to prove that species belong to
the graptoloids, more evidence than number of branches and angle of branching is
necessary. It is possible that the widening of the stipes characteristic of the didymo-
graptids of the exfensus-group, which is less prominent in Kiaerograptus kiaeri, might
be useful in specimens where the presence or absence of bithecae could not be ascertained.
It is still possible that there were some graptoloid graptolites in the Tremadoc, but they
were certainly fewer than hitherto supposed, and the remaining records must be care-
fully checked before they can be definitely accepted as graptoloids.

It is also possible that a number of species in the lower Arenig now parading as dicho-
graptids really are few-branched dendroids, only that the bithecae are overlooked because
of the state of preservation of the material. Such species might be suspected in the
Bryograptus-Tetragraptus fructicosus-Didymograptus protobifidus lineage, and in didy-
mograptids of the geometricus type, because their constant angle of branching (120-
150°) might indicate that they are descendants of three-branched forms. Didyrnograptus
latus T. S. Hall, which was shown by Thomas (1961, p. 9) to be as highly developed as
the later D. hirundo, might also belong to an entirely different group, perhaps that of
Kiaerograptus kiaeri, or might be one of its descendants. Both have, in contrast to the
c.\-/c/?5U5-group, long nemas and branches of uniform width.

The presence of bithecae in K. kiaeri has also some phylogenetic significance, show-
ing that the tendency to branch-reduction went as far as two-branched forms in the
dendroid graptolites. It also gives support to Bulman’s (1954n, p. 208; 1958, p. 161)
view that the transition between the dendroids and graptoloids was a gradual one,
taking part in several lineages independently. This transition was not necessarily con-
temporaneous in all lineages, but seems to have taken place frequently about the
Tremadoc-Arenig boundary.

K. kiaeri is an intermediate species between the Dendroidea and the Graptoloidea.
The autothecae, the general shape of the sicula, and the absence of bithecae at the first
autothecae are graptoloid features, whereas the initial budding and the bithecae at the
distant parts of the branches are dendroid ones.

NILS SPJELDN/ES: UPPER TREMADOCIAN GRAPTOLITES 131

Urbanek (1960) in a recent paper discussed the evolutionary changes in graptolite
colonies. He drew attention to the well-known fact that in Monograptids new thecal
types are often introduced in the proximal part of the colonies. This might also be the
case with K. kiaeri, where the new type of thecae (without bithecae) is found in the
proximal part of the branches. If more complete material had been present, it might
have been possible to follow the development from a purely dendroid two-branched
form through types like K. kiaeri to extensiform graptolites without bithecae, such as in
the evolutionary lineages referred to by Urbanek, but in this case from one order to
another. This might indicate that studies on thecal variation might be of considerable
phylogenetic importance in the Dendroids and Dichograptids also.

REFERENCES

BROGGER, w. c. 1882. Die Siho ischen Etageii 2 und 3 im Kristimuagebiet unci auf Eker. viii+376 pp.,
12 pi. Christiania (Oslo).

BULMAN, o. M. B. 1936. On the Graptolites prepared by Holm. Part. VII. The Graptolite fauna of the
Lower Orthoceras Limestone of Halludden, Oland, and its bearing on the evolution of the Lower
Ordovician Graptolites. Arkiv f. Zool. 28A, 17, 107 pp.

■ 1941. Some Dichograptids of the Tremadocian and Lower Ordovician. Ann. Mag. Nat. Hist.,

sen 11,7, 100-21, pi. 2.

1950. Graptolites from the Dictyonema shales of Quebec. Quart. Journ. Geol. Sac. London, 106,

63-99, pi. 4-8.

1954. The Graptolite fauna of the Dictyonema shales of the Oslo Region. Norsk Geol. Tidsskr.

33, 1-40, pi. 1-8.

1954. Status of invertebrate paleontology; Graptolithina. Bull. Mus. Comp. Zool. 112, 201-15.

1955. Graptolithina. In moore, r.c. ed.: Treatise on Invertebrate Paleontology, Part V, 101 pp.

— — 1958. The sequence of Graptolite faunas. Palaeontology, 1, 159-73.

HOLTEDAHL, o. and DONS, J. A. 1957. Geological guide to Oslo District. Skrifter Vidensk. Akad. Oslo, L
Mat.-Nat. Kl. 1957, 3, 86 pp.

KOZLOWSKi, R. 1960. Calyxdendrurn graptoloides n. gen. n. sp., a Graptolite intermediate between the
Dendroidea and the Graptoloidea. Acta Paleont. Polonica, 5, 107-25.

MONSEN, A. 1925. Uber eine neue Ordovicische Graptolitenfauna. Norsk Geol. Tidsskr. 8, 147-87. 4 pi.

STUBBLEEiELD, c. J. 1929. Notes On some early British Graptolites. Geol. Mag. 66, 268-85.

THOMAS, D. E. 1960. The zonal distribution of Australian Graptolites. Journ. Proc. Rov. Soc. New
South Wales, 94, 1-58, 15 pi.

URBANEK, A. 1960. An attempt at biological interpretation of evolutionary changes in Graptolite
colonies. Acta Paleont. Polonica, 5, 127-234, 3 pi.

NILS SPJELDN.CS

Institutt for Geologi,
Blindern, Oslo,

Manuscript received 30 March 1962 Norway

ALAIOFHYLLUM MACKENZIENSE SP. NOV.,
A DEVONIAN TETRACORAL FROM CANADA

by A. E. H. PEDDER

Abstract. The fauna of the Kee Scarp Formation at the type locality near Norman Wells, Northwest Terri-
tories, is reviewed, and its only common tetracoral is described as Alaiophylliim mackenziense sp. nov. It is con-
cluded that the fauna is either Givetian or early Frasnian in age.

The age of the Kee Scarp Formation is not clear from a study of published reports. In
1959 the writer visited the type exposure near Norman Wells, Northwest Territories,
and collected, among others, the fossils described below. He is indebted to Triad Oil
Co. Ltd., for whom the work was carried out, for permission to publish this paper and
to present the type specimens to the Geological Survey of Canada (abbreviated to GSC
below).

Fauna and age of the type exposure of the Kee Scarp Formation

When first studied and described (Stelck 1944, pp. 15, 16; in Hume 1945, p. 35) the
Kee Scarp Formation was considered to be Upper Devonian. Although this opinion has
prevailed with some geologists (Warren and Stelck 1956, p. 11; Storey 1961, p. 499), it
has been challenged by others (Crickmay 1957, p. 1 1 ; Bassett 1961, pp. 492-4), who regard
the Kee Scarp Formation as upper Middle Devonian. Galloway (1960, p. 621) considered
it to be lower Upper Devonian, but later (in Collinson 1960, p. 1218) abandoned his
earlier opinion and regarded it as upper Middle Devonian. Lenz (1961, table 1 ) considered
the lower part as either uppermost Middle, or lowermost Upper Devonian, and the
upper part as very low Upper Devonian.

Bassett (1961) extended the use of the term Kee Scarp to include the Ramparts Lime-
stone of the Fort Good Hope area, because Ramparts as a stratigraphical name is pre-
occupied. Unfortunately this obscures the relationship between the two limestones,
which are not physically one and the same, since the Kee Scarp is a discrete reef. Deter-
mination of its age must be based on the fauna at the type locality and not on those from
other localities in limestones subjectively correlated with it.

Warren and Stelck (1956, pi. 15, figs. 25-27) figured a specimen from the type locality
as Indospirifer sp. The presence of AdoJfia [= Indospirifer] would strongly suggest a
middle Frasnian age, but examination of the Kee Scarp specimen’s micro-ornament
excludes it from that genus. At the present time the specimen does not help resolve the
problem of the formation’s age.

Lenz (1961, p. 504, pi. 3, figs. 6, 7) compared a coral from the formation with Mocgeen
gallica Lang and Smith, which is a middle Frasnian species. He stated that the Kee
Scarp coral has horseshoe dissepiments, but these cannot be seen in the figure of the
longitudinal section and in fact the coral resembles that group of Givetian corals which
includes Aulophyllunf richardsoni Meek and other undescribed species.

Other forms reported in the literature (Warren and Stelck 1956; Crickmay 1957;

[Palaeontology, Vol. 6, Part 1, 1963, pp. 132-5, pi. 19.]

A. E. H. PEDDER: A NEW DEVONIAN TETRACORAL

133

Galloway 1960; Lenz 1961) from the type locality are: Stwmatoporella damnoniensis
Nicholson, Stromatopora sp. cf. S. planulata Hall and Whitfield sensii Nicholson,
Clathrocoilona aboena Yavorsky, Cladoporal sp., Alveolites sp., Aiilopora sp. cf. A. con-
ferta Winchell, Trachypora sp., Schizophoria sp., productoid indet., Atrypa or Spinatrypa
sp., ambocoeliinid, and thick-shelled pelecypods. The following were collected at the
type locality by the writer (Triad Oil Co. Ltd. collections D107, 108, 135): calcispheres,
Alaiophyllum mackenziense sp. nov.. Alveolites sp., Scoliopora sp., Thamnopora sp.,
Idiostronia sp., Amphipora ramosa (Phillips), Stachyodes sp., Hennatostroma sp.,
Spinatrypa sp., Straparollus (Euomphahis) sp. and thick-shelled pelecypod fragments.

None of these forms provides a sure index of their age. Alaiophyllum, however,
although a rare genus, is significant. It was first described from three widely scattered
Givetian deposits in Russia and has subsequently been recognized by the writer in the
Maligne equivalent and Swan Hills Member of the Beaverhill Lake Formation of
Alberta (these Canadian specimens are being described by the writer for publication by
the Geological Survey of Canada). The Maligne Formation contains Timanites, indicat-
ing that it is early Frasnian, and the Swan Hills Member, as shown by its fauna, is also
about the same age. The known range of Alaiophyllum, therefore, is Givetian to early
Frasnian and it may be concluded that the age of the Kee Scarp reef also falls within
these limits.

SYSTEMATIC DESCRIPTION
Family stauriidae Edwards and Haime 1850, sensu lato
Genus alaiophyllum Goryanov 1961

Type species (original designation). Alaiopliylliini janishevskyi Goryanov 1961 , pp. 71-73, pi. 8, figs. \a-
3<r/. Givetian stage, Boord Ridge, southern Ferghana, U.S.S.R.

Remarks. Goryanov referred the genus doubtfully to the Phillipsastraeidae. However,
the poorly and variably developed dissepimentarium and thick stereozone appear to
relate Alaiophyllum more closely to the Stauriidae.

Alaiophyllum mackenziense sp. nov.

Plate 19, figs. 1-6

Name derivation. Mackenzie River, Northwest Territories, Canada.

Types. Holotype and paratypes 1 and 2 are GSC 16850-2 respectively.

Type stratum. Kee Scarp Formation — the lower 140 feet which are exposed.

Type locality. Kee Scarp, 6 miles ENE. of Norman Wells, Northwest Territories, Canada.

Description. Compound tetracoral with loosely fasciculate corallum and gently sinuous
subcylindrical corallites. Calice and exterior surface not exposed in the types. All the
specimens collected at the type locality were tightly embedded in matrix and corallites
were not obviously parts of particular colonies; thus the size of the entire corallum is
unknown.

Transverse sections of individual corallites circular, or nearly so, with diameters of
up to 14-0 mm., generally about 12-5 mm. ; adjacent corallites touching or up to 30-0 mm.
apart. Epitheca smooth, commonly enveloped by stromatoporoids, reinforced by a

134

PALAEONTOLOGY, VOLUME 6

prominent peripheral stereozone, normally about 1-0 mm. thick but as much as 3-0 mm.
thick in parts of some corallites. Stereozone composed of dilated septal ends and
lamellar tissue. In some sections inner less-prominent sterozones are formed by scleren-
chyme deposited on the dissepiments. Septa roughly radially arranged, differentiated in
two orders and of such variable length that some minor septa are as long as major ones.
Major septa number 20 to 25 in corallites with diameters of 10-0 to 14-0 mm., but not
necessarily more numerous in larger corallites. Minor septa very variably developed,
some suppressed and discernible only in the microstructure of the stereozone. Dissepi-
ments normal or in ‘herringbone’ arrangement where the minor septa are suppressed.

Longitudinal sections with subparallel epitheca and up to 50-0 mm. in length. Stereo-
zone broad, surface rough and enveloped by stromatoporoids in places. The dissepi-
mentarium is generally between one-quarter and one-third the total width of the lumen;
dissepiments moderately long and gently curved, peripheral ones commonly coated with,
or even embedded in, sclerenchyme. Tabulae broad, but mostly incomplete, sinuous,
spaced at 1 to 6, typically 3 or 4, per mm. Trabeculae not well displayed in type material,
directed upwards and inwards at an angle of about 26° to the walls of the corallite and
very slightly divergent distally.

Remarks. The species is similar to another species of Alaiophyllum occurring rarely in
the Maligne Formation of the central Alberta Rocky Mountains. It is distinguished by
its broader dissepimentarium and narrower tabularium with consistently closer-spaced
and less complete tabulae. Further, perhaps less diagnostic, distinctions are that the
Alberta specimens have sclerenchyme coating on some of the tabulae as well as the dis-
sepiments and have individual corallites of more variable diameters. Both species almost
invariably grew with ectoparasitic stromatoporoids. A. jarushevskyi Goryanov has
shorter septa and a narrower dissepimentarium than A. mackenziense. It also has clearer
traces of the septa within the stereozones, but this may be due to differences in preserva-
tion.

REFERENCES

BASSETT, H. G. 1961. Devonian stratigraphy, central Mackenzie River Region, Northwest Territories,
Canada. In Geology of the Arctic, ed. G. O. Raasch, 1, 481-98.

COLLINSON, c. 1960. Correction. J. Paleont. 34, 1218.

CRiCKMAY, c. H. 1957. Elucidation of some western Canada Devonian formations. Published by the
author in Calgary.

GALLOWAY, J. J. 1960. Devonian stromatoporoids from the lower Mackenzie Valley of Canada.
J. Paleont. 34, 620-36.

GORYANOV, V. B. 1961. A new rugose genus from the Middle Devonian of Southern Ferghana
Paleont. Zluirnal, 1961, 1, 70-73. (In Russian.)

HUME, G. s. 1945. The Lower Mackenzie River area. Northwest Territories and Yukon. Mem. Geol.
Surv. Canada, 273.

EXPLANATION OF PLATE 19

All figures are X 4.

Alaiophyllum mackenziense sp. nov., lower 140 feet of the Kee Scarp Formation at Kee Scarp, 6 miles
WNW. of Norman Wells, Northwest Territories, Canada.

Figs. 1, 2, 6. Holotype, GSC 16850. 1, 6, Longitudinal sections; 2, transverse section.

Figs. 3, 5. Paratype 1, GSC 16851, longitudinal sections.

Fig. 4. Paratype 2, GSC 16852, ti'ansverse section.

Palaeontology, Vol. 6

PLATE 19

PEDDER, Alaiophyllum mackenziense

9

*

'J;. :’.'

r

-y-*

'^„v.

A. E. H. PEDDER: A NEW DEVONIAN TETRACORAL 135

LENZ, A. c. 1961. Devonian rugose corals of the lower Mackenzie Valley, Northwest Territories. In
Geology of the Arctic, ed. G. O. Raasch, 1, 500-14.

STELCK, c. R. 1944. Schooner Creek. Canol Report 34, on open file. Government of Canada, Depart-
ment of Northern Affairs.

STOREY, T. p. 1961. Devonian stratigraphy — Norman Wells region (abstract). In Geology of the Arctic,
ed. G. O. Raasch, 1, 499.

WARREN, p. s. and stelck, c. r. 1956. Reference fossils of Canada, pt. 1, Devonian faunas of Western
Canada. Spec. Pap. Geol. Assoc. Canada, 1.

Manuscript received 13 August 1962

A. E. H. redder
Triad Oil Co. Ltd.,
535-7th Avenue S.W.,
Calgary, Alberta,
Canada

THE LAMELLIBRANCH GENUS PROTHYRIS IN
THE UPPER DEVONIAN AND CARBONIFEROUS

OF GREAT BRITAIN

by R. B. WILSON

Abstract. The morphology and distribution of the species of the lamellibranch genus Prothyris Meek, occurring
in the Upper Devonian and Carboniferous of Great Britain, are reviewed. Of the three Upper Devonian species
described by Whidbome (1896), P. scalprata is regarded as conspecific with P. contorta, and it is concluded that
P. recta should be removed from the genus. A new Upper Devonian species, P. stubblefieldi, is described. Four
species are recognized in the Carboniferous; P. breviformis Wilson, P. obhtiga Wilson, P. scotica sp. nov., and
P. carbomria sp. nov. Some general comments are made on the mode of life, environment, and evolution of the
genus.

The lamellibranch genus Prothyris Meek 1869, type species Prothyris elegans Meek,
was first recorded from the Upper Carboniferous of Illinois and Nebraska (Meek and
Worthen 1869, p. 172). American authors have since described several species, showing
that the genus ranged from the Upper Devonian to the Upper Carboniferous in North
America. Whidborne (1896, pp. 86-89) first recognized the genus in Europe when he
described three species from the Upper Devonian of Devon. Hind (1906, p. 148) identi-
fied a shell as Prothyris elegans from the Scottish Namurian, the first record in the
British Carboniferous. Since then, various occurrences of Prothyris have been cited from
several horizons and the present paper attempts to examine the morphology and distri-
bution of the known British species.

For permission to examine specimens in their care, the writer is grateful to the Director of the
Geological Survey and Museum, Mr. A. E. Blackwell of the North Devon Athenaeum, Barnstaple,
and Mr. A. G. Brighton of the Sedgwick Museum, Cambridge. This paper is published with the per-
mission of the Director of the Geological Survey. Specimens with registration numbers prefixed by
GSM are housed in the Geological Survey and Museum, London, and those by GSE are in the
Collection of the Geological Survey, Edinburgh.

Genus prothyris Meek 1869

Meek (in Meek and Worthen 1869, p. 172) proposed the name Prothyris for a lamel-
libraneh genus found in the Upper Carboniferous of Nebraska and Illinois, accom-
panying the announcement with a general description of the genus, but he did not assign
any species to it. Later Meek (1871, pp. 8-9) gave a much fuller account, and selected
Prothyris elegans Meek as the type species. His generie diagnosis is as follows: ‘Equi-
valve, very inequilateral, longitudinally oblong, valves compressed or moderately con-
vex, nearly closed or a little gaping behind and more or less widely gaping in front,
where the hiatus is increased in size by a nearly rectangular notch in the margin mainly
below the middle; beaks depressed and very near the anterior end with a small ridge
usually extending from the anterior side of each to the corner of the anterior marginal
notch ; dorsal margin without escutcheon or lunule, being erect and sharp behind the
beaks. Surface merely marked with striae of growth. Hinge and interior unknown.’

[Palaeontology, Vol. 6, Part 1, 1963, pp. 136-44, pi. 20.]

R. B. WILSON: PROTHYRIS IN THE DEVONIAN AND CARBONIFEROUS 137

Only minor additions need be appended to this diagnosis. Immediately posterior and
parallel to the ridge running from the umbo to the anterior marginal notch, here called
the anterior notch, a shallow groove is present on well-preserved specimens of most spe-
cies examined. Although the hinge has not been observed, several specimens of Carboni-
ferous species show a structure in the umbonal region which may indicate some degree
of dentition. It consists of two subparallel short lines, about 1 mm. long in a valve of
12 mm. length, running postero-ventrally from the umbo, and preserved as faint im-
pressions or ridges. These may indicate poorly developed cardinal teeth; this would
support the suggestion made by Meek (1871, p. 8) that the genus has affinities with the
Solenidae.

Hind (1908, p. 352) suggested that the ear-shaped process, anterior to the ridge
running from the anterior notch to the umbo, here called the anterior lobe, contained
the anterior adductor muscle. He also thought that the anterior notch marked a byssal
opening. Wilson (1961, p. 102) questioned these suggestions as P. oblonga Wilson
appears to have the anterior adductor scar posterior to the ridge bounding the anterior
lobe, and it seems unlikely that a bivalve gaping at both ends, and probably of burrowing
habit, would possess a byssus. Elias (1957, pp. 741-3) recorded radial markings on the
valves of P. soleniformis and described, but did not figure, a simple hinge apparatus. He,
also, regarded Prothyris as a subgenus of Solen Linnaeus, but the present writer con-
siders that the internal and hinge characters are still not sufficiently well known and that
the anterior notch and lobe are of generic significance.

DEVONIAN SPECIES

Whidborne (1896, pp. 86-89, pi. 9, figs. 12-18), in his study of the Devonian faunas of
the south of England, described three species from the Upper Devonian of north
Devon, which he assigned to Prothyris. All the surviving syntypes of these species have
been re-examined.

^Prothyris' recta Whidborne
Plate 20, fig. 3

Discussion. The remaining type material of Prothyris recta (Whidborne 1896, pp. 86-87,
pi. 9, figs. 12-14) consists of two of the three original figured specimens and several in-
determinate cited specimens, all in the Geological Survey Collection, London. The
specimen illustrated as fig. 13 cannot be found. Of the figured specimens, GSM 6085
(op. cit., fig. 12) is incomplete anteriorly and posteriorly, and is preserved in a dark-
grey limestone. It is too poorly preserved to be used for determining the specific charac-
ters. GSM 87343 (op. cit., fig. 14), now refigured (PI. 20, fig. 3), is an internal mould of
a left valve, with the ventral margin missing; it is also preserved in a dark-grey limestone.
It is subquadrate in outline, about 19 mm. long, and the height is estimated to be about
14 mm. An angular anterior umbonal ridge runs half-way towards the ventral margin,
and a rounded posterior umbonal ridge extends to a point half-way down the posterior
border. Between these ridges, the valve is moderately convex and there are several in-
distinct short ridges radiating from the umbo. The anterior margin has irregularities
caused by the granular fracture of the matrix. The largest of these irregularities is about
one-third of the way down ; Whidborne interpreted it as the anterior notch of Prothyris.

138

PALAEONTOLOGY, VOLUME 6

There is no ridge extending from this feature to the umbo, and the present writer inter-
prets it as fortuitous and of no morphological significance. Whidborne (1896, p. 87)
stated that the species was of variable shape, but the specimen GSM 87343, just de-
scribed, is the only one of the remaining syntypes which is reasonably well preserved; it
is here selected as lectotype of the species ‘P.’ recta. As it does not possess any of the
diagnostic characters of Prothyris, the writer excludes it from that genus.

Prothyris con tort a Whidborne

Plate 20, figs. 1 , 2

Prothyris contorta Whidborne 1896, pp. 87-88, pi. 9, figs. 15-16(7.

Prothyris scalprata Whidborne 1896, pp. 88-89, pi. 9, figs. 17-18«.

Discussion. An examination of the type specimens of P. contorta and P. scalprata has
shown that only one species is involved. As the only factor relevant to determining which
specific name is valid is that of page precedence, it is suggested that the name scalprata
be regarded as a junior synonym of contorta.

Lectotype. The specimen figured by Whidborne (1896, pi. 9, fig. 15), marked as such, and housed in the
North Devon Athenaeum, Barnstaple, is here selected as lectotype. It is an internal mould of a left
valve, preserved in dark-grey limestone from Kingdon’s Shirwell, north Devon.

EXPLANATION OF PLATE 20

All the lamellibranch photographs (figs. 1-9) are twice natural size. The accompanying outline draw-
ings are natural size; they were prepared by drawing in waterproof ink the outlines of the shells on

X 2 photographs, bleaching the photographs, and then reducing the remaining outlines photographi-
cally. Figs. 1-9 were prepared by Mr. W. D. Fisher and the author.

Figs. 1, 2. Prothyris contorta Whidborne. 1, GSM 7044, both valves. Upper Devonian, Plaistow Mill
Quarry (Sloly Beds), north Devon. Figd. by Whidborne as P. scalprata (1896, pi. 9, fig. 17). 2, Speci-
men in the North Devon Athenaeum, Barnstaple, lectotype, left valve. Upper Devonian, from King-
don’s Shirwell, north Devon. Figd. by Whidborne (1896, pi. 9, fig. 15) as F. contorta.

Fig. 3. ‘’Prothyris' recta Whidborne. GSM 87343, lectotype, left valve, Upper Devonian, South Cave,
Baggy, north Devon. Figd. by Whidborne (1896, pi. 9, fig. 14).

Fig. 4. Prothyris stnbblefieldi sp. nov. GSM 87387, holotype, left valve. Upper Devonian, at 1,458| feet
in Willesden Borehole, north London.

Fig. 5. Prothyris oblonga Wilson. GSE 11755, holotype, left valve. Lower Carboniferous, at 3,175i2 feet
in Archerbeck Borehole, Canonbie, Dumfriesshire.

Fig. 6. Prothyris breviformis Wilson. GSE 11758, holotype, right valve. Lower Carboniferous, at 3,393
feet in Archerbeck Borehole.

Figs. 7, 8. Prothyris scotica sp. nov. 7, GSE 11719, holotype, right valve, from mudstone under Calmy
Limestone, Scottish Upper Limestone Group, Namurian (Ej), shore 135 yards south-east of
Hospital, Kirkcaldy. 8, GSE 1 1827, paratype, both valves, roof of Index Limestone, Namurian (Ej),
at 2843 feet in Powis Mains No. 1 Bore, Stirlingshire.

Fig. 9. Prothyris carbonaria sp. nov. GSM 87388, holotype, left valve, Mansfield Marine Band, Coal
Measures, at 1,724^ feet in Lindholme Borehole, Yorkshire.

Fig. 10. View of serial sectioning machine from side away from the operator showing general layout
and position of the coolant drip-feed system. Height of machine to top of coolant tank 20 inches.

Fig. 11. Detail of compound slide rest viewed from operating side. Longitudinal feed handle on the
right, cross-feed handle on the left. Magnetic positioner mounted on the tool post block in front
of the cupped diamond wheel.

Fig. 12. Detail of magnetic positioner showing specimen attached to steel mounting block held by the
positioner.

Palaeontology, Vol. 6

PLATE 20

WILSON, Prothyris

HENDRY, ROWELL, and ST AN LEY, parallel grinding machine

R. B. WILSON: PROTHYRIS IN THE DEVONIAN AND CARBONIFEROUS 139

Description. Small, elongate, subrhomboidal, moderately convex, the maximum con-
vexity occurs along the poorly defined umbonal ridge which fades out half-way to the
posterior end of the valve. The umbo is situated at the anterior fifth of the length of the
shell; it is prosogyrous and inconspicuous. The anterior end is symmetrically rounded
but interrupted by a re-entrant notch from which a narrow, ill-defined ridge runs to the
umbo. The dorsal margin is short and straight, passing into the long, gently convex,
postero-dorsal border which ends in the acutely rounded postero-ventral extremity of
the valve. The ventral margin is very gently convex. Two subparallel linear ridges run
from the umbo half-way to the ventral margin. The postero-dorsal region is marked by
two broad radiating ribs on the internal mould and by three radiating striae on the
exterior of the shell. The musculature and hinge characters are not seen.

Dimensions.

Length Height
mm. mm.

Lectotype ...... 16 6

GSM 7044 (Whidborne, p). 9, fig. 1 7) . 17 6-5

Remarks. Whidborne (1896, p. 87), in his description of P. contorta, mentioned that the
ventral margin is slightly concave, as shown in his illustration of the specimen now chosen
as lectotype (op. cit., pi. 9, fig. 15). This outline was false, caused by some matrix, since
removed, which covered part of the margin. The other specimen he figured as P. contorta
(pi. 9, fig. 16) is very poorly preserved and indeterminate. Of the four specimens he
illustrated as P. scalprata, GSM 7044 (pi. 9, fig. 17) is now refigured (PI. 20, fig. 1) and
referred to P. contorta', Sedgwick Museum specimen H 363 (pi. 9, fig. 18) is poorly
preserved, but is probably P. contorta as now described, and the two specimens shown
on pi. 10, figs. 16, 17, are indeterminate lamellibranch fragments.

Prothyris stubbleficldi sp. nov.

Plate 20, fig. 4

Discussion. Stubblefield (1947, p. 20) reported an occurrence of Prothyris from a bore-
hole sunk by the D’Arcy Exploration Company at Willesden, north London. It occurred
at a depth of 1,458| feet in a lilac-stained mudstone of Upper Devonian age. Although
only one specimen is known, it is reasonably well preserved and merits description.

Holotype. GSM 87387, a left valve, in the Geological Survey Collection, London.

Description. Small, elongate, dorsal and ventral margins subparallel, diverging very
slightly towards the posterior end. Umbo inconspicuous, placed at anterior end of
dorsal margin. A rounded, posterior umbonal ridge extends almost to the postero-
ventral angle, giving the valve a moderate convexity. The anterior end is incomplete
ventrally but a distinct anterior lobe is present, separated from the remainder of the
valve by a narrow, rounded ridge, posterior to which a shallow groove occurs. The
dorsal and ventral margins are almost straight, the posterior border is obliquely trun-
cate making an angle of about 45 degrees with the ventral margin. The postero-ventral
angle is sharply rounded. Dimensions of the holotype are: length 18-5 mm., height 6 mm.

Remarks. P. stubbleficldi differs from the other British Devonian species, P. contorta,

140

PALAEONTOLOGY, VOLUME 6

by having a relatively longer dorsal margin, by the absence of radial striae on the postero-
dorsal region, and by the subparallel alignment of its dorsal and ventral margins. It
differs from P. bergica Drevermann, the German Devonian (Etroeungt) species (Drever-
mann 1902, pp. 498-9, pi. 14, fig. 15), in that the latter species has convex dorsal and
ventral margins. Several species were described in the last century from the Devonian of
North America, but the descriptions are meagre. Some of them appear to lack an anterior
notch and lobe, suggesting that they may belong to some other genus. Until these species
are revised, any comparisons with them must be unsatisfactory but P. stubblefieldi
shows no marked resemblance to any of the American species so far described.

CARBONIFEROUS SPECIES

The earliest occurring species recorded from the British Carboniferous are P. ob/onga
Wilson and P. breviformis, which were found in beds of Lower Visean age in the Archer-
beck Borehole, Canonbie, Dumfriesshire (Lumsden and Wilson 1961, p. 11). They have
been described already (Wilson 1961, pp. 101-3), but, for the sake of comparison, their
salient features are given below.

Prolhyris oblonga Wilson
Plate 20, fig. 5

Prolhyris oblonga Wilson 1961, pp. 101-2, pi. 3, figs. 5-7.

Description. Relatively small, elongate, the average length/height ratio is 2-75. Dorsal
margin slightly arched, the ventral one convex anteriorly and straight posteriorly, while
the posterior end of the shell is bluntly rounded. The anterior notch is well developed,
almost right-angled, situated about half-way up the anterior end. At this notch the valves
gape appreciably. A poorly developed posterior umbonal ridge is present, antero-ventrally
to which the shell is marked with fine, regular, concentric striae and obscure radiating
striae. On the postero-dorsal area, only faint concentric striae are seen. Dimensions of
holotype: length 9-8 mm., height 3-4 mm. The largest specimen seen is about 15 mm.
long.

Prolhyris brevifonnis Wilson
Plate 20, fig. 6

Prothyris brevifonnis Wilson 1961, pp. 102-3, pi. 3, figs. 8-10.

Description. Relatively small, elongate-quadrate, the length/height ratio is approxi-
mately 2, which feature distinguishes it from all other known species. The dorsal margin
is gently convex, the ventral one convex at the extremities and straight in the middle,
while the posterior end of the shell is bluntly rounded and subtruncate. The anterior
notch is small and inconspicuous, and the anterior lobe is narrow in the antero-posterior
direction. There is no umbonal ridge, the maximum convexity being in the sub-umbonal
region. The only markings seen on the shell are fine, concentric striae and obscure radial
striae, both confined to the antero-ventral part of the shell. Dimensions of holotype:
length 8-9 mm., height 4-2 mm. The largest specimen seen is 10-5 mm. long.

R. B. WILSON: PROTHYRIS IN THE DEVONIAN AND CARBONIEEROUS 141

Prothyris scotica sp. nov.

Plate 20, figs. 7, 8

Prothyris elegans. Hind non Meek 1908, p. 353, pi. 2, figs. 48-50.

Prothyris sp. nov. Wilson 1962, p. 514, pi. 66, fig. 26.

Holotype. GSE 11719, right valve, internal mould, in silty mudstone under the Calmy Limestone,
Scottish Upper Limestone Group (Eg), on shore 185 yards south-east of Hospital, Kirkcaldy, Fife.

Paratype. GSE 11827, both valves, open, attached, in mudstone above the Index Limestone, Scottish
Upper Limestone Group, at 2841- feet, in Powis Mains No. 1 Bore 1959 (N.C.B.), Stirlingshire.

Description. Elongate, average length/height ratio (from twenty-three specimens) is
2-9, dorsal and ventral margins parallel. The umbo is situated near the anterior end of
the dorsal margin, but not raised above it. The dorsal margin is straight, except for its
posterior third which slopes ventrally in a very gentle convex curve to meet the almost
vertically truncate posterior border. The ventral margin is almost straight for most of its
length, but anteriorly it curves relatively sharply into the anterior notch which is almost
rectangular. The anterior lobe is prominent, its height being about twice its length. It is
separated from the main part of the valve by a narrow ridge running from the umbo and
inclined anteriorly. Posterior to this ridge a shallow groove occurs. A distinct umbonal
ridge runs to the postero-ventral angle, angular at its proximal end but becoming less
well-defined distally. The anterior lobe and that part of the shell anterior and ventral to
the umbonal ridge are marked by fine, regular, concentric striae, but over the umbonal
ridge and on the postero-dorsal region the striae tend to coalesce into irregular plications.
The valves gape at the anterior notch and at the posterior end. The shell is partly pre-
served on the paratype and is seen to be very thin. Hinge and interior not seen.

Dimensions.

Length
mm.

Holotype 20

Paratype 21-5

The largest specimen seen is 30 mm. long.

Remarks. The species just described was first recognized as a Prothyris by Hind (1908),
who identified it as the American species P. elegans Meek. This latter species differs from
P. scotica in being relatively longer, length/height ratio 3-5 (Meek, in Hayden 1872,
p. 223), its dorsal margin is arched, and there is no distinct umbonal ridge. P. scotica
appears to be similar to P. soleniformis Elias, a late Mississippian form from Oklahoma,
in its general dimensions and by possessing an umbonal ridge, but the illustration of the
holotype of the latter species (Elias 1957, pi. 90, fig. 3) shows a shell markedly different
in outline from that just described.

P. scotica is found in mudstones, silty mudstones, and, more rarely, in ironstones in the
Scottish Midland Valley, ranging from the Hosie limestones up to the lower part of the
Passage Group. In terms of the goniatite zonal scheme, this range is from the uppermost
part of P2 to the top of Eg as it is represented in Scotland.

Wilson (1961, p. 93) named a specimen as P. aff. soleniformis Elias from the Archer-
beck Borehole, Dumfriesshire. The horizon is in the roof of the Linns Limestone, prob-
ably near the top of the Visean. A similar form has now been found at the same horizon

Height

mm.

7

c. 8

142

PALAEONTOLOGY, VOLUME 6

in Penton Linns, Canonbie. Neither specimen is well preserved but they show a close
resemblance to P. scotica. This occurrence of Prothyris in the Canonbie area, Dum-
friesshire, is about the horizon of the Tyne Bottom and Jew limestones of north-east
England. From a band between these limestones in the Roddymoor Boring in Co.
Durham (Woolacott 1923; Lee 1924), a specimen of Prothyris has now been recognized.
It is numbered T 3897*^ in the collections of the Geological Survey, Leeds. Both valves
are present, but the shell appears to be a juvenile form, incomplete posteriorly, and it is
specifically indeterminate.

Prothyris has recently been recorded from Northern Ireland (Fowler and Robbie
1961, p. 234). It occurred in the Millstone Grit (E) at approximately 150 feet in Palm
Lodge No. 1 Borehole. The specimen, NIR 774, at present housed in the Geological
Survey Collections, London, is part of a right valve, but both anterior and posterior ends
are missing. Sufficient is preserved however, to suggest that it is a specimen of P. scotica.

From the Millstone Grit of the north of England, two specimens of Prothyris are in
the Geological Survey Collections, Leeds. They are HS 4591 and CS 844a and come
from Chalk Beck, 2 miles north of Caldbeck, Cumberland, and the Cayton Gill Beds
(Dunham and Stubblefield 1945, pp. 241-3) in Watergate Beck at Bay Croft Hill Farm,
3 miles west of Pateley Bridge, Yorkshire, respectively. They are both incomplete, but
sufficient is preserved to suggest that they have affinity with P. scotica.

Prothyris carbonaria sp. nov.

Plate 20, fig. 9

Holotype. GSM 87388, left valve, internal mould, in mudstone from the Mansfield Marine Band,
Coal Measures, at 1,724^ feet in Lindholme Borehole, about 10 miles east-north-east of Doncaster,
Yorkshire. It is slightly incomplete on the ventral and posterior margins.

Description. Relatively small, elongate-oval, average length/height ratio (from seven
specimens) is about three, dorsal and ventral margins almost parallel. The umbo is
inconspicuous, situated almost at the anterior end of the cardinal border. The dorsal
margin is very gently convex, with a shallow groove parallel and ventral to it. The
anterior lobe is narrow, length about one-third of height, and its height is about one-
third of the height of the valve. The lobe is separated from the main part of the valve by
a faint linear ridge at right angles to the dorsal margin, with a shallow groove posterior
to it. The anterior notch forms an oblique angle passing with a gentle curve into the ventral
margin, which is straight for most of its length. The posterior end of the valve is almost
vertically truncate, gently convex, its dorsal and ventral extremities symmetrically
curved. There is no umbonal ridge, and the valves are only gently convex with slight
gapes at the anterior notch and the posterior end. The anterior lobe and antero-ventral
part of the valve are marked by fine, regular, concentric striae, whereas the postero-dorsal
area has irregular plications apparently formed by the coalescence of groups of striae.
Hinge and interior unknown. The approximate dimensions of the holotype are; length
15-5 mm., height 5-5 mm. The largest specimen seen is approximately 24 mm. long.

Remarks. Although P. carbonaria is similar in its dimensions to P. scotica, it differs from
the latter in having an arched dorsal margin and no umbonal ridge; its anterior lobe is
relatively smaller, and the ridge dividing the lobe from the valve is normal to the dorsal

R. B. WILSON: PROTHYRIS IN THE DEVONIAN AND CARBONIEEROUS 143

margin, while in the latter species it is inclined anteriorly when traced from the umbo.
P. carbonaria resembles P. elegans in having a groove parallel to the cardinal edge and in
the general shape of the posterior end, but it is distinguished by being relatively shorter
than the American species, by having a relatively smaller anterior lobe, and by the ridge
posterior to the anterior lobe lying at right angles to the hinge line.

P. carbonaria appears to be a rare species in Britain. Few specimens were available
for study, and they all came from the Mansfield Marine Band of the Yorkshire and
Nottinghamshire Coalfields and the equivalent horizons in the South Wales and Bristol
Coalfields.

GENERAL COMMENTS

Protbyris has been shown to range from the Upper Devonian to the Upper Carboni-
ferous in Britain, as it does in North America. With the exception of P. contorta, found
in limestone, it normally occurs in mudstones and silty mudstones. The shell matter is
very thin, and the normal faunal associates are thin-shelled molluscs, suggesting that the
environment was not rich in carbonates. The fact that the valves gape anteriorly and
posteriorly, is evidence that the animal was a burrower. The valves are commonly
found associated, either open wide or closed, but with one slipped slightly relative to the
other. This suggests little disturbance of the bottom deposits, but it is also probably
a reflection of the ability of the ligament to withstand decay until the valves were en-
tombed.

From the evidence of the five British species known at present, several changes took
place in the valve shape during the evolution of the genus. The Devonian species have
the dorsal and ventral margins diverging posteriorly, whereas the Carboniferous forms
have these borders more or less parallel. The postero-ventral angle in the Devonian
species is acute, but this angle increases to almost a right angle in the later species. The
anterior lobe tends to be better defined in the Carboniferous, with the maximum develop-
ment of this feature being displayed by P. scolica. These general trends in the evolution
of the valve shape appear to have been paralleled in the North American species.

REFERENCES

DREVERMANN, F. 1902. Ucbcr cine Vertretung der Etroeungt-Stufe auf der rechten Rheinseite. Zeit.
Deiitsch. geol. Gesell. 54, 480-524.

DUNHAM, K. c. and STUBBLEFIELD, c. J. 1945. The stratigraphy, structure and mineralization of the
Greenhow mining area, Yorkshire. Quart. J. Geol. Soc. 100, 209-68.

ELIAS, M. K. 1957. Late Mississippian fauna from the Redoak Hollow Formation of southern Oklahoma.
Part 3. Pelecypoda. J. Paleont. 32, 1-57.

FOWLER, A. and robbie, j. a. 1961. Geology of the country around Dungannon. Mem. Geol. Surv.
N. Ireland.

HAYDEN, F. V. 1 872. United States Geological Survey of Nebraska and portions of the Adjacent Territories.
Washington.

HIND, w. 1906. In Snmni. Prog. Geol. Surv. for 1905.

1908. On the lamellibranch and gastropod fauna found in the Millstone Grit of Scotland. Trans.

Roy. Soc. Edin. 46, 331-59.

LEE, G. w. 1924. On the faunal sequence of the Carboniferous rocks met in the Roddymoor Boring,
Co. Durham. Sninm. Prog. Geol. Surv. for 1923, 146-9.

LUMSDEN, G. I. and WILSON, R. B. 1961. The stratigraphy of the Archerbeck Borehole, Canonbie, Dum-
friesshire. Bull. Geol. Surv. G.B. 18, 1-89.

144

PALAEONTOLOGY, VOLUME 6

MEEK, F. B. and WORTHEN, A. H. 1869. Description of New Carboniferous fossils from the western
States. Proc. Acad. Nat. Sci. Philadelphia, for 1869, 137-72.

1871. Descriptions of some new types of Palaeozoic shells. Arner. J. Conch. 7, 4-10.

STUBBLEFIELD, c. J. 1947. Abst. Proc. Geol. Soc., No. 1437.

WHiDBORNE, G. F. 1896-1907. A monograph of the Devonian fauna of the south of England. 3. Palaeont.
Soc.

WILSON, R. B. 1961. Palaeontology of the Archerbeck Borehole, Canonbie, Dumfriesshire. Bad. Geol.
Sla v. G.B. 18, 90-106.

1962. A review of the evidence for a Nebraskan fauna in the Scottish Carboniferous. Palaeontology,

4, 507-19.

WOOLACOTT, D. 1923. On a boring at Roddymoor Colliery, near Crook, Co. Durham. Geol. Mag.
60, 50-62.

R. B. WILSON

Geological Survey Office,
19 Grange Terrace,
Edinburgh, 9

Manuscript received 22 March 1962

A RAPID PARALLEL GRINDING MACHINE FOR
SERIAL SECTIONING OF FOSSILS

by R. D. HENDRY, A. J. ROWELL, and J. W. STANLEY

For many years the internal structures of fossils have been investigated by means of
orientated serial sections. There are two essential requirements of any machine used for
this purpose : the successive surfaces must be parallel and the interval between them must
be known and preferably capable of being controlled. Croft (1950) gave a brief historical
account of the development of equipment designed for this purpose. In the same paper
he gave details of an instrument that he had designed capable of producing ground
surfaces at an interval of 10 ft, a considerable improvement in precision over previous
machines. Perhaps the most notable advance in equipment for serial sectioning since 1950
is that described by Jefferies, Adams, and Miller (1962), who have successfully designed
and produced a machine capable of automatically grinding and producing a photograph
of the surface of the specimen every 100 ft.

Problems currently being investigated in the Department of Geology, University of
Nottingham, include studies of the variation of internal structures in species of brachio-
pods and ontogenetic studies of these forms. With the material used, both these projects
require a serial sectioning instrument with an accuracy comparable to that of the Croft
grinder. A second requisite is that, as in the Croft grinder, the specimen should be
readily removable from the instrument to enable an acetate peel to be made of the
ground surface. For our purpose the acetate peels have an advantage over other methods
of recording the ground surface of the fossil, since in material that has not recrystallized
they record the micro-structure of the shell, which itself may provide some information
on the ontogenetic history of the various structures, particularly those associated with
the cardinalia. Either ‘wet’ or ‘dry’ peel techniques can be used. We employ the former as
we find it gives better resolution and freedom from bubbles and with the materials used
has a speed comparable with that of the ‘dry’ method.

The projects were initiated using a hand-operated Croft grinder. Manual methods are,
however, extremely time-consuming and laborious if much sectioning has to be done.
The machine described below was devised in an endeavour to improve the efficiency and
speed with which successive peels could be produced and has proved very successful.

Basically the machine consists of a small lathe unit with several attachments. Grinding
of the surface of the specimen is accomplished by passing it across the face of a diamond-
impregnated cup wheel. The movement and cutting action are similar to that of the larger
machine used by Olsen and Whitmore (1944) in their studies of fossil mammals. Their
machine, however, was designed for handling relatively large specimens and its standards
of accuracy would be unacceptable when dealing with invertebrates, particularly if they
were small.

Among the existing relatively low-priced lathes that are available, the small models
initially designed for watch-making are well suited for adaption to a serial sectioning

(Palaeontology, Vol. 6, Part 1, 1963, pp. 145-7 pi. 20.]

C 1015

L

146

PALAEONTOLOGY, VOLUME 6

machine for use with small- and medium-sized invertebrate fossils. They are compact
and designed to work to close limits. The machine used was constructed from basic
units of the Pultra lathe, obtainable from Smart and Brown (Machine Tools) Ltd.,
Biggleswade, Beds. The units employed were a lathe bed, chip tray base plate, high-
speed head-stock, compound slide rest, and an arbor for holding the cup wheel. The
compound slide rest is screw operated, with precision ground threads of 1 mm. pitch.
Micrometer collars, adjacent to the handles, are graduated in of a revolution,

each graduation representing a movement of 10 ;u,.

The 3-inch diameter electrometallie diamond cup wheel, with 240-grade diamond, is
mounted on the arbor held in the headstock spindle of the lathe. The machine is Vee-
belt driven by a | h.p. electric motor mounted beneath the bench, producing a wheel
speed of approximately 5,200 r.p.m. Coolant in the form of a soluble cutting fluid
(Sternopal) is supplied by drip feed from a can mounted at the rear of the machine, the
waste suds running through the lathe bed, down a plastic tube to a large bottle also
beneath the working bench. A ‘Perspex’ guard was formed and mounted round the wheel
to reduce spray from the coolant.

The specimen holder consists of a permanent magnet chuck, in this case an ‘Eclipse’
magnetic positioner from which the ‘V’ shape on the face away from the wheel was
removed. The positioner is screwed on to the tool post block of the lathe. It has a work-
ing face of 63-5 X 51 mm.

The specimens are embedded in polyester resin. ‘Ceemar’ cold-setting resin has been
used and found very satisfactory, but extensive tests on other brands have not been
carried out and many others may be equally suitable. Small polythene moulds, sold for
making ice cubes, make good containers in which to carry out the embedding process,
because the resin is adhesive to most common materials except polythene and expands
slightly during setting. The embedded specimens are attached by tough wax (obtainable
from Cutrock Engineering Co. Ltd.) to mild steel mounting blocks. A number of mount-
ing blocks were made of 6-25 mm. mild steel, both sides being ground to obtain accuracy
and maximum holding power. The blocks are all 50 mm. long and of various widths, the
actual width of mount used depending on the size of the specimen. The mounting block
with specimen fixed can then be readily attached to the magnetic chuck and this switched
on. No trouble has been experienced with movement of the specimen on the mounting
block or with movement of the block on the magnetic chuck. Care has to be used to keep
the working face of the chuck and the back of the mounting block clean if the high
precision of the machine is to be utilized.

With the present arrangement of the machine the maximum area it can grind is
50 X 40 mm. and the specimen can be of any length up to 25 mm. It is capable of grinding
successive surfaces at a minimum spacing of 10 ;U.; the grinding time for removal of this
increment varies slightly with the size of the specimen and the nature of its matrix, but is
in the order of 10-15 seconds. If, as often is the case, larger spacings are required between
successive sections, these may be removed in increments of up to 50 by successive
passes across the diamond wheel.

The cycle for a specimen is as follows. The embedded specimen, attached to its steel
block, is fitted to the magnetic chuck while clear of the diamond wheel. Longitudinal and
cross feeds are adjusted to bring the specimen block just into contact with the grinding
surface of the wheel. The cross feed is wound back and the longitudinal feed advanced

R. D. HENDRY ET AL.\ A RAPID PARALLEL GRINDING MACHINE 147

not more than 50 fi. Advancing the cross feed now causes the removal of 50 fi from the
block surface, this process is repeated until the specimen is exposed. At this stage the
block is removed from the chuck and details recorded in the desired manner, the longi-
tudinal feed reading having been noted in readiness for the next grinding operation.
After recording the details of the surface of the specimen, it is returned to the chuck, the
longitudinal feed then advanced the desired increment between 10 ft and 50 ft over the
previous setting and the surface ground as before.

With the ‘wet’ acetate peel technique used, the longest stage in the preparation is the
drying of the acetate solution on the block surface. A solution of 50 per cent, by volume
of liquid ‘Cerric’ DL4013 cellulose acetate and acetone produces a quick-drying film,
which with care is free from bubbles. It is necessary to dilute the solution occasionally
during use to compensate for the loss of acetone by evaporation. A thin syrupy consis-
tency should be maintained to enable a smooth coat to be brushed on quickly. The
drying time can be reduced by using only a very thin layer of acetate solution. The result-
ing film has little strength, but after its margins have been freed, it may be readily peeled
from the block using cellulose adhesive tape (‘Scotch’ tape or ‘Sellotape’). It has been
found convenient to process a number of blocks in such a way that the acetate on the
earlier ones dries while the following blocks are in turn ground, etched, and coated with
acetate solution. Using ten blocks, the acetate on the first may be peeled after the etching
and painting of the last is finished, thus a continuous cycle may be maintained by a single
operator without having to wait for material to dry. Using the above solution and ten
blocks it is possible to produce at least twenty-five peels per hour, which is a very great
improvement over any of the hand-operated methods that have been tried. A further
advantage is the high degree of cleanliness of the method ; it is not necessary after grinding
to use more than a paper tissue to dry the block prior to etching.

The total cost of the machine as illustrated (PI. 20) and described is in the order of
£135.

We are indebted to Professor W. D. Evans for his interest and encouragement in the work, to F. Ban-
croft for modifying the existing lathe, and to J. Eyett for taking the photographs.

REFERENCES

CROFT, w. N. 1950. A parallel grinding instrument for the investigation of fossils by serial sectioning.
J. Paleont. 24, 693-8.

JEFFERIES, R. p. s., ADAMS, J. B. and MILLER, R. c. 1962. Automatic serial sectioning machine for fossils.
Nature, Loud. 193, 1166-7.

OLSEN, F. R. and WHITMORE, F. c., Jr. 1944. Machine for serial sectioning of fossils. J. Paleont. 18,210-15.

R. D. HENDRY, A. J. ROWELL, J. W. STANLEY
Department of Geology,
University of Nottingham,

Manuscript received 30 July 1962 Nottingham

MALAYOMAORICA GEN. NOV. (FAMILY
AVICULOPECTINIDAE) FROM THE
INDO-PACIFIC UPPER JURASSIC;
WITH COMMENTS ON RELATED
FORMS

by J. A. JELETZKY

Abstract. Critical reinvestigation of the aviculopectinid form ‘Aucella' malayomaorica Krumbeck from the
late Jurassic of the Indo-Pacific region indicates that it does not belong either to the genus Buchia (= Aitcella)
or to any other known aviculopectinid genus. A new genus Malayomaorica is erected to receive ‘Aiicella' malayo-
maorica. This genus is characterized by; (1) presence of a submedian furrow-like excavation in the ligamental
pit of the left valve; (2) presence of an irregularly shaped tooth-like bulge in front of the ligamental pit at the
anterior margin of the left ligamental plate; (3) presence of a strong ligamental ridge rising above the true
dorsal shell margin in the right ligamental plate; and (4) a wide to very wide byssus notch.

Malayomaorica gen. nov. is much more closely allied to Meleagrinella, Aucellina, and Arctotis than to Buchia.
This genus is, accordingly, placed in subfamily Oxytominae and not in Buchiinae (= Aucellinae). The genus
Aiicelliua is also placed in the Oxytominae, as it appears to be an independent offshoot of Meleagrinella un-
related to Buchia.

The boreal genus Arctotis is the closest known ally of Malayomaorica. The latter is accordingly interpreted as
an offshoot of the Arctotis lineage, which immigrated, together with the true Buchia species, into the Indo-
Pacific region during the early part of the Upper Jurassic.

The purpose of this paper is to establish a new genus for a peculiar aviculopectinid
pelecypod from the Upper Jurassic (Kimeridgian) of Indonesia and New Zealand, as
well as to investigate the morphological distinctions and probable phylogenetic ties of
this biochronologically important genus.

^AitceUa' malayomaorica Krumbeck 1923 has been the source of a considerable taxo-
nomical and biochronological confusion ever since its original description by Boehm
(1911) under the name of Aitcella plicata Zittel. This species has been assumed to be an
‘archaic’ member of the genus Buchia (better known as Aitcella) closely allied to the
so-called Buchia racliata-impressae species group from the Oxfordian stage of Europe
(Krumbeck 1923, pp. 70-74). It was, consequently, widely assumed to be of the early
Upper Oxfordian age in terms of the international standard stages by the European
workers who studied Upper Jurassic rocks and faunas of Indonesia and New Zealand
(e.g. Krumbeck 1923, 1934; Wandel 1936; Stolley 1929). At about the same time, how-
ever, Crickmay (1932, p. 5) has made an astute but undeservedly neglected observation
that: ^Aitcella from the more southerly localities, e.g. India, and New Zealand, are very
different, and hardly congeneric.’ This conclusion appears to be quite correct, at least so
far as ‘‘Buchia' malayomaorica (and possibly ‘‘Buchia'’ misolica) is concerned.

The Oxfordian dating of ^Aitcella' malayomaorica beds has been recently refuted by
Eleming (1958, p. 379; 1960, p. 267, table 1), who was able to demonstrate their early to
mid-Kimeridgian age. Eleming (1958, p. 380) has, furthermore, concluded that: ‘two
parallel lineages [of Buchia', writer’s remark], one represented by malayomaorica and

[Palaeontology, Vol. 6, Part 1, 1963 pp. 148-160, pi. 21.]

J. A. JELETZKY: MALAYO MAO RIC A GEN. NOV. FROM THE JURASSIC 149

misolica, the other by the plicata-subpallasi group, lived in the south-east Tethyan region
during the Kimeridgian-Tithonian.’

Several Upper Jurassic Buchia forms of the Canadian Western Cordillera appear to
be closely allied to and are possibly conspecific with Buchia spitiensis (Holdhaus),
B. legwninosa (Stoliczka), and B. blanfordiana (Stoliczka) of the Indo-Pacific region
(Jeletzky 1950, pp. 28-29; 196-). The present paper explains the reasons for the state-
ment (Jeletzky 196-) that ‘ ^"Buchia’' malayomaorica is rather different from and prob-
ably not congeneric with either the boreal Buchia (better known as Aucella) or the
Buchia plicata-subpallasi species group of the Indo-Pacific region.’

The writing of this paper has been facilitated by valuable comments from Dr. C. A. Fleming, Senior
Palaeontologist of the New Zealand Geological Survey, who also has kindly provided the writer with
several well-preserved specimens of" Aucella' malayomaorica Krumbeck 1923 from the Kawhia section.
It is published by permission of the Director, Geological Survey of Canada.

SYSTEMATIC DESCRIPTION

Family aviculopectinidae Etheridge, Jr., 1906 emend. Newell 1938
Subfamily oxytominae Ichikawa 1958

Genus malayomaorica gen. nov.

Plate 21, figs. 1-2

Derivation of name. Malayo for the Malay Archipelago; maorica for Maori, the name of the New
Zealand native people. Gender feminine.

Type species. Aucella malayomaorica Krumbeck (1923, pp. 65-76, pi. clxxiii, figs. 2-12,
17; pi. clxxvii, figs. \2>a-b). Early to mid-Kimeridgian beds of Indonesia and New
Zealand.

Diagnosis. Ligamental plate of the left valve separated into two parts by a submedian
transverse furrow. This transverse furrow is a modification of the spoon-like excavation
of the ligamental pit of some other members of the family. This furrow extends all across
the ligamental plate from the very tip (underside) of the beak to its lower margin. The
lower margin of the ligamental plate is indented at the lower end of the furrow.

The anterior part of the ligamental plate is markedly thickened ; it carries an irregularly
shaped bulge and an articulation furrow (= ‘Gelenkgrube’) near the anterior margin.

The byssus notch of the right valve is wide to very wide and broadly triangular in
shape. The middle part of the ligamental plate of the right valve has an irregularly
swollen, strong ligamental ridge, which rises above the true dorsal shell margin and is
directed obliquely upward and forward. This ligamental ridge fits into the transverse
furrow of the left ligamental plate.

Type specimen and synonymy. No previous selection of the type specimen of Malayo-
maorica malayomaorica (Krumbeck 1923) is known to the writer. The left valve repre-
sented in pi. clxxiii (2), fig. 4 of Krumbeck’s (1923) work is, therefore, selected herewith
as the lectotype of the type species of the genus Malayomaorica. It is from the so-called
Aucella horizon of Portuguese Timor, and was collected between Bele and Tooi near
Kampong Kamlachi, in calcareous marl with Inoceramus. According to Dr. Fleming, it
is preserved in Geol. Pal. Inst., Bonn.

150

PALAEONTOLOGY, VOLUME 6

Fleming (1958, p. 378), Glaessner (1945), and Marwick (1953) have placed Aucel/a
boehmi Marwick 1926 into the synonymy of malayomaorica Krumbeck 1923. Most
other workers concerned (beginning with Krumbeck 1934) have, of course, been aware
of the fact that " Aucella' plicata Boehm non Zittel from Kawhia Point is conspecific
with ' Aucella' malayomaorica Krumbeck; they have, however, overlooked the fact that
this New Zealand form was formally named by Marwick.

There is little doubt that 'Aucella' boehmi Marwick 1926 is both congeneric and con-
specific with Malayomaorica malayomaorica (Krumbeck 1923). The characteristic furrow-

EXPLANATION OF PLATE 21

(Unless otherwise indicated, all specimens are reproduced natural size. To facilitate the comparison of
their shapes, the valves of the pelecypods figured are, as a rule, oriented with their hinge margins
uppermost.)

Figs. 1a-g. Malayomaorica malayomaorica (Krumbeck 1923). New Zealand, North Island, north-east
coast of Heteri Promontory, Kawhia Harbour. Heterian stage (early to mid-Kimeridgian). A topo-
type of Biichia boehmi Marwick 1926. GSC Cat. No. 17006.

Left valve of a variant with a strongly reduced anterior ear. The semi-smooth appearance of most
of the anterior part of the valve (fig. 1e) appears to be due to its abrasion. 1a: Dorsal view of the
beak part. 1b: Oblique posterior view of the ligamental plate. Note the furrow-like excavation of the
middle part of the ligamental pit. The ligamental plate occupies all of the underside of the beak.
Ic: Posterior view. Note the predominance of the concentric, lamellar growth-lines on this part of
the valve. Id: Anterior view. Note the rounded, tooth-like bulge at the anterior rim of the ligamental
plate and the pronounced embayment of the valve’s margin underneath it. 1e: View from the left.
Note the strongly reduced anterior ear and well-developed posterior ear. If: The beak part and the
ligamental plate, x 3. Oriented exactly as in fig. 1b. 1g: Inside view.

Figs. 2a-f. Malayomaorica malayomaorica (Krumbeck 1923). Same locality and age as for the specimen
shown in figs. 1a-g. GSC Cat. No. 17007. Complete shell with its valves almost in normal position
relative to each other. A variant with typically developed anterior and posterior ears of the left valve
and typically developed and well-preserved sculpture of both valves. The flattened, folded appearance
of the toothlike bulge at the anterior margin of the ligamental plate of the left valve (figs. 2a and 2f)
appears to be an individual feature peculiar to this specimen.

2a: Right valve viewed from the right. The transversally grooved ligamental plate of the left
valve and the underside of the left beak are clearly visible. Note that the ligamental plate occupies
all of the underside of the left beak up to its very tip. The wide gap between the byssus ear and the
anterior ear of the right valve is characteristic of Malayomaorica. 2b: Left valve viewed from the
left. Note the predominance of radial ribbing over the concentric growth-lines and the nodose to
spinose appearance of radial ribs at their intersections with the concentric growth-lines. This
ornament contrasts strongly with that of Biichia erringtoni (Gabb) shown in figs. 8a, b, f. 2c:
Anterior view of both valves. 2d : Posterior view of both valves. 2e : Dorsal view of the beak parts of
both valves. 2f: Beak part of the right valve, X 3 viewed from the right. Oriented as in fig. 2a. Note
the sculpture of the surface of the byssus ear and the marked bend of the latter toward the left valve.
Fig. 3. Biichia mosqueiisis (Buch non Keyserling non Lahusen). Canada. N.W.T., Mackenzie District.
Richardson Mountains. East slope of Aklavik Range, southern side of nameless creek falling into
Husky Channel, about f mile south of the top of Mount Gifford. Collected on the float of the basal
part of lower shale-siltstone division. Middle to late Kimeridgian. GSC Cat. No. 17008. An early
form of the species somewhat transitional to Biichia ex gr. coiiceiitrica {Sov^erhy]-erriiigtoiii (Gabb).
An internal cast of right valve viewed from the right. The central part of the valve, including its beak
part, is strongly abraded.

Fig. 4. Biichia mosqueiisis ( Buch non Keyserling non Lahusen). Same locality and age as for the speci-
men shown in fig. 3. GSC Cat. No. 17009. An early form of the species somewhat transitional to
Biichia ex gr. coiiceiitrica {So'^erhy)-erriiigtoiii (Gabb). Inside view of the right valve with the well-
preserved shell layer. The concentric ornament is just as well developed as on the surface of the shell.

Palaeontology, Vol. 6

PLATE 21

JELETZKY, Malayomaorica

J. A. JELETZKY; M ALAYO M AO RIC A GEN. NOV. FROM THE JURASSIC 151

like excavation of the ligamental pit is neither visible in Krumbeck’s (1923) drawings
nor clearly described in the text. This feature was, however, excellently figured and
described by Wandel (1936, p. 457, pi. xvii, figs. 2-3), who has been able to study
Krumbeck’s (1923) original material of the species.

Age and correlation. Krumbeck (1923, expl. of pi. clxxii (2), fig. 2) has tentatively dated
the Malayomaorica maJayomaorica beds of Indonesia as of early Upper Oxfordian age.
Fleming (1958, p. 379; 1960) has recognized more recently, however, that in New Zealand

while the radial ornament is subdued compared with its development on the shell’s surface. Note the
simply grooved, Meleagrinella-Wke appearance and orientation of the inside surface of the byssus
ear, which differs from that of the younger Bitchia species (e.g. B. piochii var. russieiisis shown in
figs. 5d, e).

Figs. 5a-e. Buchia piochii (Gabb) var. russieiisis (Pavlow). Canada, N.W.T., Franklin District. Prince
Patrick Island, Mould Bay, lat. 76° 16' 30" N., long. 1 19° 27' 9" W. Collected on the float of Mould
Bay Formation (restricted). Late Portlandian (s. str.) GSC Cat. No. 17010. A well-preserved right
valve.

5a: View from the right. Note the pronounced bend of the byssus ear toward the left valve. The
byssus ear of Buchia mosqueusis (Buch) shown in figs. 3-4 is, in contrast, oriented more or less in the
plane of contact of the valves. 5b: Anterior view. Note that the anterior surface of the byssus ear is
oriented almost perpendicular to the plane of contact of the valves. 5c: Posterior view. 5d: Inside
view, X 3. Note the pronouncedly spoonlike appearance of the inside surface of the byssus ear con-
trasting with the simply grooved appearance of the same in Buchia mosqueusis (Buch) shown in
fig. 4. 5e: Dorsal view of the beak part, X 3, to show structural detail of the right ligamental plate and
byssus ear. Note that the byssus ear is oriented almost perpendicularly to the plane of contact of the
valves (plane of symmetry of the shell).

Figs. 6a-c. Buchia keyserliugi (Lahusen 1888) s. lato. East Greenland, Trail 0. Locality 92 of Donovan
(Medd. om Gronl. Bd. Ill, No. 4, 1953, pp. 51-52). Middle or late Lower Valanginian. Property of
Min. og Geol. Mus., Copenhagen.

A Buchia crassicoUis-Wke variant of the species, possibly identical with B. crassicollis var. gracilis
Sokolov, 1908 non Lahusen 1888. A well-preserved internal cast of the left valve with considerable
patches of well-preserved shell layer. Note the pronouncedly lamellar appearance of concentric
growth-lines and the absence of any radial ornament.

6a: Anterior view. 6b: Viewed from the left. 6c: Inside view of the beak and hinge areas, x3.

Figs. 7a-b. Buchia piochii (Gabb) var. russieiisis (Pavlow). Same locality and age as for the specimen
shown in figs. 5a-e. GSC Cat. No. 1701 1. A somewhat distorted and cracked left valve with excellently
preserved shell layer.

7a: Inside view of the complete valve. 7b: Inside view of the beak part, X 3, to show structural
detail of the beak and ligamental plate. The ligamental plate is broken in the middle of the ligamental
pit. The structure of the ligamental plate and the beak is closely similar to that of Buchia keyserliugi
s. lato shown in figs. 6a-c.

Figs. 8a-f. Buchia coiiceiitrica (Sowerby) var. erriiigtoiii (Gabb). Canada, Western Cordillera of British
Columbia. Tyaughton Lake area, north side of a pass If miles north-west of Peak of Sheba. Lower
part of Eldorado Group. Early Kimeridgian? GSC Cat. No. 17012.

A complete internal cast with valves in approximately normal position relative to each other.
Small patches of the inner shell layer are preserved around the right beak and near the posterior
margin of the right valve. The byssus ear is largely preserved as an internal cast. 8a: Left valve,
viewed from the left. 8b: Right valve viewed from the right. The ligamental plate and beak of the
left valve are clearly visible above the beak of right valve. 8c: Anterior view; 8d: Posterior view.
8e: Dorsal view of the beak part. 8f: Beak part of the right valve and the inside view of the left
valve, X 3. Note the far-reaching similarity of the structure of the byssus ear to that of Buchia
mosqueusis (Buch) shown in fig. 3 and its dissimilarity with that of B. piochii var. russieiisis shown in
figs. 5a-e. The left beak is clearly separated from the ligamental plate and protrudes above the latter
as is the case in younger Buchia species.

152

PALAEONTOLOGY, VOLUME 6

this species occurs in early to mid-Kimeridgian rocks. The M. malayomaorica beds of
Indonesia are, therefore, believed by the writer to be of early to mid-Kimeridgian age.

Morphological and taxonomic remarks. The morphology and infra-specific variability
of M. malayomaorica have been described in detail by Krumbeck (1923), Marwick
(1926), and Wandel (1936). The present discussion of its morphology can, therefore, be
limited to description, illustration, and taxonomic analysis of those morphological
features that are diagnostic or otherwise significant at the generic and family level.

The generally accepted reference of M. malayomaorica to the genus Buchia {=Aucella)
is based exclusively on its morphological similarity to such European forms as "Aucella''
impressae Quenstedt and 'A.' radiata Trautschold. This is stressed in the following
statement made by Krumbeck (1923, p. 73) in the original description of M. malayo-
maorica'. Tm ganzen kann es nach meiner Ansicht nicht zweifelhaft sein, dass Aucella
malayomaorica den alteren Aucellen und zwar vor allem der Impressae-Radiata-GxwppQ
am nachsten steht und in dieser mit A. impressae die meisten und engsten Beriihrungs-
punkte aufweist. Von den ubrigen mir bekannten Aucellen ^ — vielleicht ausgenommen A.
plicata Zitt, sp. — weicht sie . . . ab.’ This conclusion has been accepted by all subsequent
workers in its entirety. As we shall see, however, only Krumbeck’s claim of the far-
reaching similarity of M. malayomaorica to "A.' impressa and 'A.’ radiata is apt and valid.
His assumption that these two European forms are true representatives of Buchia is, on
the contrary, erroneous and based on their misinterpretation by Pompeckj (1901).

Malayomaorica malayomaorica typically possesses a fairly well-developed anterior
ear in the left valve (Krumbeck 1923, p. 66; Wandel 1936, p. 457; this paper, PI. 21,
figs. 2a, b, f), although this ear can become strongly reduced and barely noticeable in
some specimens (Wandel 1936, pi. xvii, figs. 9, 11 ; this paper, PI. 21, figs. 1a, e, g).

The taxonomic significance of the presence or absence of the anterior ear in the left
valve of Buchia has been discussed by Pompeckj (1901) in connexion with his reference
of such aberrant forms as ^A.' radiata Trautschold and 'A.' impressae to this genus. The
interpretation of these two Oxfordian forms as ‘archaic’ representatives of Buchia con-
necting this genus with Pseudomonotis (Meleagrinella of recent usage) led Pompeckj
(1901, pp. 327, 339, pi. xv, figs. 12, 16) to the revision of the generally accepted diagnosis
of the former genus. Pompeckj concluded that the oldest representatives of Buchia
possess a distinct, although short, anterior ear in the left valve, which becomes lost in the
younger representatives of the genus. This conclusion has been strongly criticized by
Sokolov (1908, 1912, and elsewhere).

Sokolov (1908, p. 3) insisted that the possession of an anterior ear in both valves and
the peculiar shape of their beaks excludes "Aucella’ radiata and "A.’ impressae from the
genus Buchia. Only the right valve figured by Pompeckj (1901, pi. xv, figs. 2, 5) under the
name of Aucella impressae is a representative of Buchia according to Sokolov (1908,
p. 3, footnote). A new name, A. pompeckii, was proposed for this form by Sokolov (1908),
only to be passed later (Sokolov 1912, pp. 115-16) into the synonymy of Buchia lata
(Trautschold).

To strengthen his argument Sokolov (1912, pp. 108-13; pi. 2, figs. \\a, b, \2a-\7>b)
has redescribed and refigured the original specimens oi" Aucella’’ Trautschold and

was able to show that this species is in all respects a typical representative of the group
of species formerly included in Pseudomonotis s. lato but now separated under the name

J. A. JELETZKY: MALAYOMAORICA GEN. NOV. EROM THE JURASSIC 153

of Meleagrinella. Sokolov (1912, pp. 115-16) points out in the same paper that the
oldest-known true Biichia species, such as B. lata (Trautschold), occur already in middle
to late Callovian rocks of Russia. They antedate, therefore, the Oxfordian 'A.' impressae
and radiata, which have been assumed to be the oldest representatives of Buchia by
Pompeckj (1901, pp. 327, 337). B. lata (Trautschold) is, moreover, already a typical
representative of the genus in the complete absence of the anterior ear of the left valve
and in the sculpture (see below). There is little doubt that Sokolov’s (1908, 1912) criti-
cisms of Pompeckj’s (1901) treatment of 'A.’ radiata and impressae are fully valid.

Although published in German, in part at least (Sokolov 1908, pp. 26-27), Sokolov’s
conclusions have been completely ignored by those European workers who have made
the most thorough study of Malayomaorica maJayomaorica and unreservedly accepted
Pompeckj’s (1901) interpretation of Buchia. Krumbeck (1923, p. 71) actually stated that
he did not use Sokolov’s works, except for his plates.

As already seen, the presence of the anterior ear in the left valve suffices to exclude
M. malayomaorica from the genus Buchia, as all its true representatives are characterized
by the complete absence of the same. Its placement in this genus would probably never
have been seriously contemplated except for the authority of Pompeckj’s (1901) work
and the unfortunate neglect of Sokolov’s (1908, 1912) research.

Only Krumbeck (1923, p. 67) describes the ornamentation of M. malayomaorica in
any detail. He stresses the strong variability of the ornamentation and the common
occurrence of the specimens with more or less obsolete sculpture in all population
samples available. Specimens characterized by the presence of the concentric ornament
alone on parts or all of the shell are also common in his material.

Judging by the New Zealand specimens in the writer’s possession (PI. 21, figs. 1-2),
the marked variation in strength and character of ornamentation from one specimen to
another (or from one part of the shell’s surface to another within a single specimen) is at
least partly related to the degree of abrasion of the shells concerned prior to or after
their burial. This abrasion (e.g. in specimen shown in PI. 21, figs. 1a-g) was apparently
facilitated by the restriction of ornamentation to the outer layers of the shell. The writer
does not deny the strong variability of the ornamentation and the presence of specimens
with obsolete sculpture. The sculpture of the specimen of M. malayomaorica shown in
Plate 21, figs. 2a, b, f is, nevertheless, assumed to be typical of the species. The speci-
mens of M. malayomaorica figured by Boehm (1911, pi. 2, figs. 2-A, non 1) and the
specimen figured by Marwick (1926, pi. 71, fig. 10) are similarly strongly sculptured.
The same is, finally, true of the herewith selected lectotype of the species (see above).

The ornament of the well-preserved specimens studied (PI. 21, figs. 2a, b, f) consists
of closely spaced, fine, sharp, and straight radial ribs intersecting more widely spaced
concentric growth-lines (or growth-ridges). In our material the radial ribs are generally
speaking stronger developed on the left valve, where they tend to be about as strongly
developed as the concentric growth-lines (PI. 21, fig. 2b). In the right valve the concentric
growth-lines or growth-ridges are generally speaking more strongly developed than the
radial ribs (PI. 21, figs. 2a, f). The concentric growth-lines tend to be imbricated and
lamellar in appearance in the best-preserved specimens; they also tend to become
markedly frilled and to form small, ill-defined tubercles and spines whenever intersected
by the radial ribs (PI. 21, figs. 2a, b, f). The intersection of the radial and concentric
elements of sculpture, which are about equally strongly developed, results in the markedly

154

PALAEONTOLOGY, VOLUME 6

reticulate appearance of some parts of the shell’s surface. In the specimens available,
I was not able to observe clearly the intercalation of the second-order radial ribs with
the first-order radial ribs observed by Krumbeck (1923, p. 67).

The sculpture of Malayomaorica malayomaorica described above is quite unlike that
of Buchia. As it can be seen in Plate 21, figs. 3, 6a, b, 8a, b, f, the ornament of Buchia
is usually visible on the surface of internal casts as well as on the surface of the shell.
Furthermore, as already noted by Sokolov (1912, p. 110), the concentric growth-lines
(or growth-ridges) of Buchia are always much more pronounced than the radial ribs.
The latter are either of a very low relief or almost unnoticeable, even in Buchia ex gr.
conceutrica (Sowerby) (= ? B. bronni Rouillier) (PI. 21, figs. 8a, b, f), and are mostly
completely absent in the majority of younger Buchia species (PI. 21, figs. 6a, b). The
imbricated appearance of the concentric growth-lines of Malayomaorica malayomaorica
and the presence of small tubercles and spines on them is also quite foreign to Buchia
and characteristic of Meleagriuella. The same is true of the intercalation of the second-
order radial ribs observed by Krumbeck (1923, p. 67). On the whole the ornamentation
of Malayomaorica malayomaorica is distinctly more similar to that of Meleagriuella than
to that of Buehia. It differs, however, from that of Meleagriuella in much greater strength
of the concentric growth-lines (or growth-ridges). In Meleagriuella the radial ribs are,
indeed, always much stronger developed than the concentric growth-lines, which can be
sometimes quite weak (as, for example, in Meleagriuella radiata Trautschold, see Sokolov
1912, pi. 2, figs. 11(7, 12(7, c). The intercalation of the second-order radial ribs with
those of the first order is, finally, much better developed in Meleagriuella than in
Malayomaoriea.

The sculpture of Arctotis (Bodylevsky 1960, p. 44) differs from that of Malayomaorica
in the same way as that of Meleagriuella. It is, furthermore, coarser in the best-known
representatives of the genus (Bodylevsky 1960, pi. 7, figs. 1(7, b, 2a; Borissiak 1915, pi. 11,
figs. 3, 6, 12).

The sculpture of Aucelliua aptieusis (d’Orbigny) Pompeckj 1901 and other Barremian-
Aptian representatives of the genus is essentially similar to that of Meleagriuella and
Malayomaorica. The radial ribs strongly predominate over the concentric growth-lines.
The latter are, furthermore, often imbricated, tuberculate and/or spinose whenever they
cross the radial ribs (Pompeckj 1901, pp. 353-4, pi. xvi, figs. \a, 5a, and the writer’s own
observations on Canadian specimens). The sculpture of Aucelliua gryphaeoidesi^oV'iQxby),
on the contrary, is more like that of Buchia than that of older Aucelliua species. This is,
however, probably a secondary phenomenon, reflecting the convergent development of
Aueelliua and Buchia.

Pompeckj’s (1901, pp. 328-32, 336, expl. of plates) nomenclature of morphological
elements of the ligamental plate of Aviculopectinidae is used in the following discussion
with some additions (e.g. excavations of the ligamental pit).

The edentulous ligamental plate of the left valve of Malayomaorica malayomaorica is
wider than that of any other aviculopectinid genus (PI. 21, figs. 1b, If). The ligamental pit
of this plate is subdivided into two parts by a deep transverse groove, which is broadly
V-like in cross-section (PI. 21, figs. 1b, f). This groove begins underneath the very tip
of the short, blunt, centrally located beak, and extends right across the ligamental plate.
Both parts of the ligamental plate slope markedly toward this groove in its proximity.
The lower margin of the ligamental plate is more or less clearly indented at the lower

J. A. JELETZKY: MALAYOMAORICA GEN. NOV. FROM THE JURASSIC 155

end of the transverse groove discussed above (PL 21, figs. 1b, f, g). In the specimens
available to the writer, as well as in those figured by other workers, the posterior part
of the ligamental pit is one and a half to three times longer than its anterior part
(PI. 21, figs. If, 2e). It is covered by numerous, more or less sinuous, longitudinal striae
(PI. 21, figs. If, 2f). An oblique, more or less sharp rim slanted posteriorly separates the
posterior part of the ligamental pit from the still longitudinally striated inner surface of
the posterior ear (PI. 21, figs. If, 2f). A curved, low, longitudinal ridge may occur in the
middle of the posterior part of the ligamental pit (PL 21, figs. If, 1g).

The anterior part of the ligamental pit is covered by sinuous, longitudinal striae,
similar to those of its posterior part. The shell is markedly strengthened underneath this
part of the pit. A more or less pronounced bulge occurs in front of the anterior part of
the ligamental pit (PL 21, figs. 1b, d, f, g, 2a, e, f). This bulge is often more or less
regularly rounded (PL 21, figs. Id, If), but it may also be quite irregularly shaped, flat-
topped, and folded over (PL 21, figs. 2a, c, f). This bulge occupies the inner surface of
the anterior ear of M. malayomaorica. The inner surface of the bulge described above is
also covered by irregularly curved, fine or coarse striae. An irregularly curved groove or
depression, corresponding to the articulation pit (== ‘Gelenkgrube’ of Pompeckj 1901,
p. 336) of Biichia and other aviculopectinid genera, separates this bulge from the liga-
mental plate proper. This articulation groove or depression persists right across the
ligamental plate, is finely or coarsely striated, and may ramify in some of the specimens
studied. The presence of striations on the bulge and articulation groove of Malayo-
maorica described above suggests that these features have also been covered by the
ligament.

The anterior margin of the left valve is usually more or less markedly concave just
below the above-described bulge (PL 21, figs. 1a, b, d, 2c). In some of the specimens
studied the anterior margin of this valve forms a broadly rounded, deep embayment at
that place. This embayment sharply separates the anterior ear from the rest of the shell
(PL 21, figs. 1a, b, d).

The above-described structure of the left ligamental plate of Malayomaorica differs
sharply from that of Buchia. In Buchia the bottom of the ligamental pit is essentially
flat and appears to lack any excavations or furrows similar to the transverse furrow of
M. malayomaorica (PL 21, figs. 6c, 7a, b; Pompeckj 1901, pi. xv, figs. 17, 18, 21 ; pi. xvi,
figs. 4, 6d). The left beak of Buchia is, at that, situated at the extreme anterior end of the
ligamental plate, well forward of the ligamental pit and immediately above the articula-
tion pit (= ‘Gelenkgrube’). The articulation pit of Buchia differs, furthermore, from that
of Malayomaorica (and from that of all other Jurassic and Cretaceous aviculopectinid
genera) in its regularly hoof-like shape, greater depth, and well-outlined, near vertical
rims (PL 21, figs. 6c, 7a, b; Pompeckj 1901, pi. xv, figs. 17-18). Finally, no structure
homologous to the ligamental bulge of Malayomaorica discussed above is present in
Buchia.

The structure of the left ligamental plate of Aucellina is only imperfectly understood.
Judging by the description and figures of Pompeckj (1901), and some Canadian speci-
mens available to the writer, Aucellina is more similar, in this and other respects, to
Meleagrinella and Arctotis than to Buchia. It can, accordingly, hardly be a member of
the subfamily Buchiinae Cox (see below).

The structure of the left ligamental plates of Meleagrinella and Arctotis is much more

156

PALAEONTOLOGY, VOLUME 6

similar to that of Malayomaorica than to that of Buchia. To begin with, all better-known
representatives of MeleagrineUa possess an irregularly rounded, more or less transverse
excavation of the middle part of the ligamental pit equivalent to the transverse furrow of
Malayomaorica. The anterior rim of this excavation may, furthermore, become steep and
furrow-like in appearance (Pompeckj 1901, pi. xv, fig. 15; Borissiak 1909, pi. ii, fig. \Aa\
Bodylevsky 1960, p. 44). The central excavation of MeleagrineUa and Arctotis differs,
nevertheless, from the homologous transverse furrow of Malayomaorica in being broad,
irregularly rounded in outline, and broadly concave in cross-section. Malayomaorica
differs, furthermore, from MeleagrineUa (but apparently not from Arctotis) in lacking,
completely, the tooth-like bulge separating the ligamental pit from the articulation pit.
This tooth-like bulge appears to be present in all better-known species of MeleagrineUa
(Pompeckj 1901, pi. xv, figs. 15-16; Borissiak 1909, pi. i, fig. 12c; pi. ii, figs. 14c, 15c).
The ligamental plate of Malayomaorica has, instead, an irregular, tooth-like bulge at its
anterior rim forward from the furrow-like articulation pit (= ‘Gelenkgrube’) (PI. 21,
figs. 1a, b, d; 2a, f). This bulge appears to be absent in MeleagrineUa and Arctotis.

The articulation pit of MeleagrineUa, Arctotis, and probably Aucellina, differs strongly
from that of Malayomaorica (and Buchia, see above) in being more like a slight sinus
than a regular pit or furrow. The articulation pit of MeleagrineUa, Arctotis, and probably
Aucellina, is an only slightly depressed and ill-defined area of pronouncedly bent and
lamellated concentric growth-lines situated on the underside of the anterior ear of the
left valve well forward from its beak (Pompeckj 1901, pi. xv, figs. 15-16, 19; pi. xvi,
figs, 2>a, 6b, \0a; Borissiak 1909, pi. i, fig. 12c; pi. ii, figs. 14c, 15c; 1915, pi. 11, figs. 7-9;
Bodylevsky 1960, pi. 7, fig. \a).

The ligamental plate of the left valve of M. malayomaorica occupies all of the under-
side of its short and blunt beak (PI. 21, figs. 1b, f, 2f). The left beak is, thus, not separated
at all from the ligamental plate. In typical representatives of Buchia and Aucellina, in
contrast, left beaks are more or less clearly separated from and protrude above the left
ligamental plate (PI. 21, figs. 6a, c, 7a-b; Pompeckj 1901, pi. xv, fig. 14; pi. xvi, figs.
6c, lOa). This separation of left beaks from the ligamental plates apparently exists also
in the true ‘archaic’ Buchia species (PI. 21, figs. 8b, c, f), although it is not so well
expressed in these forms. The beak structure of Barremian-Aptian Aucellina species is
unknown.

The structure of the left beak of MeleagrineUa is much more similar to that of Malayo-
maorica than to that of Buchia and Aucellina. As pointed out by Borissiak (1909, p. 10,
pi. i, fig. 12c), the left beak of M. echinata var. donetzensis protrudes only slightly beyond
the upper margin of the ligamental plate. Pompeckj’s (1901, p. 325, pi. xv, figs. 15, 16)
descriptions and drawings indicate that similar relationships prevail also in MeleagrineUa
echinata and M. impressae.

The best-known representatives of Arctotis are indistinguishable from Malayomaorica
in the structure of their left beaks. In Arctotis lenaensis (Lahusen), the type species of
this genus, the ligamental plate occupies all of the underside of the short, blunt, and
centrally located left beak (Borissiak 1915, pi. 11, figs. 7-9), just as in Malayomaorica
malayomaorica. The same appears to be true also of A. intermedia (Bodylevsky 1960,
pi. 7, fig. la).

The ligamental plate of the right valve of Malayomaorica malayomaorica possesses
a strong ligamental ridge in its middle part. This ridge has been well described and figured

J. A. JELETZKY: MALAYOMAORICA GEN. NOV. EROM THE JURASSIC 157

by Wandel (1936, p. 458, pi. xvii, figs. \a-c, 4b-c, 11); it rises above the true dorsal
shell margin and is directed obliquely forward.

The above-described ligamental ridge of Malayomaorica is completely absent in the
right valve of Buchia to the best of the writer’s knowledge (PI. 21, figs. 3, 4, 5a, d, e,
8b, f; Pompeckj 1901, pi. xv, figs. 2, 5, 6, and 9). A similar ligamental ridge, is however,
present at least in some (probably in all) representatives of AuceUina, Meleagrinella,
and Arctotis (Pompeckj 1901, pi. xv, fig. 7; pi. xvi, figs. \a, b; Bodylevsky 1960, pi. 7,
figs. 2a, b). The ligamental ridge of Arctotis intermedia Bodylevsky 1960 is, moreover,
particularly similar to the ligamental ridge of Malayomaorica malayomaorica in its
shape, position on the ligamental plate, &c.

Other characteristic features of the right valve of Malayomaorica malayomaorica are :
the stoutness and exceptionally large size of the byssus ear (PI. 21, figs. 2a, f), the large
size, the great to very great width and the broadly triangular shape of byssus notch, the
marked bend of the byssus ear toward the left valve, and strong ornamentation of the
external surface of the byssus ear (PI. 21, fig. 2f). In some specimens (PI. 21, figs. 2a, f)
the anterior end of the byssus ear is covered by a fold-like flap of the previously de-
scribed tooth-like bulge of the left valve. If considered in relation to the total shell’s
dimensions, the byssus ear of M. malayomaorica is probably larger and stouter than
that of any other Jurassic and Cretaceous aviculopectinid genus, with the possible
exception of some small-sized representatives of Arctotis (e.g. A. intermedia Bodylevsky
1960).

The byssus ear and notch of Malayomaorica malayomaorica differ strongly from the
homologous features of the true ‘archaic’ Buchia species of the northern hemisphere
{Buchia of the B. lata-concentrica-moscjuensis species group); they are at the same time
relatively more similar to the byssus ear and notch of the younger Buchia species.

The mid-Callovian to Kimeridgian ‘archaic’ Buchia of the B. lata-concentrica-mos-
quensis species group are characterized by the relatively small byssus ear extending for-
ward and upward more or less in the plane of contact of the valves (PI. 21, figs. 3-4, 8b,
f; Pompeckj 1901, pi. xv,fig. 2). The groove occurring on the inner surface of the byssus
ear in these Buchia species is, unlike that of younger Buchia species, open at its anterior
end (PI. 21, figs. 4, 5d). The byssus ear of the presently discussed ‘archaic’ Buchia
species is thus rather similar to that of Meleagrinella (Pompeckj 1901, pi. xv, figs. 1, 7).
This explains the previously discussed erroneous assignment of such Meleagrinella species
as M. radiata and M. impressae to Buchia (Pompeckj 1901).

In the latest Jurassic and early Lower Cretaceous Buchia species, beginning with
B. piochii (Gabb, s. lato) (= B. russiensis Pavlow and B. mniovnikensis Pavlow) (PI. 21,
figs. 5a-e), the byssus ear gradually becomes relatively larger and stouter and bends more
and more toward the left valve. In the Valanginian Buchia forms, such as B. keyserlingi
s. lato (Pompeckj 1901, pi. xv, figs. 6, 8-10, 21) the external surface of the byssus ear is
directed almost perpendicularly to the plane of contact of the valves (plane of symmetry
of the shell). At the same time, the anterior margin of the byssus ear begins to extend,
ridge-like, across the anterior end of the canal that runs on the inner side of the byssus
ear. A distinctly spoon-like shape of byssus ear (when seen from inside; PI. 21, figs.
5d, e) results; this biochronologically important feature also reaches its maximum
development in the Valanginian Buchia keyserlingi-crassicollis species group. These
morphologically progressive changes of the byssus ear in the course of evolution of

158

PALAEONTOLOGY, VOLUME 6

Buchia have already been clearly recognized by Pompeckj (1901, p. 326). They have
however, been, completely neglected by later workers.

In spite of its greater similarity to the byssus ear of the younger Buchia species, the
byssus ear of Malayomaorica differs from it in its large and widely gaping byssus notch.
As noted by Pompeckj (1901, p. 326, pi. xv, figs. 6, 8-10, 20-21), the byssus notch of the
younger Buchia species is all but closed. Only a short and extremely narrow fissure
remains at the anterior end of the ear in these species. The byssus notch of all Jurassic
Buchia species, including that of its true ‘archaic’ representatives (PI. 21, figs. 3, 4, 8b,
f), is also much narrower than that of Malayomaorica.

The byssus ear and byssus notch of MeleagrineUa differ from those of Malayomaorica
in about the same way as do the byssus ear and notch of the true ‘archaic’ Buchia species
(see above). The byssus ear and notch of MeleagrineUa are, furthermore, proportionally
smaller than those of any known Buchia species (see Pompeckj 1901, pi. xv, figs. 1, 4, 7).

The byssus ear and notch of Arctotis are more similar to those of Malayomaorica
than are those of any other Jurassic and Cretaceous aviculopectinid genus. As noted in
the description, and clearly visible in the figures, of Bodylevsky (1960, p. 45, pi. 7, figs.
\a, 2a-h), Arctotis intermedia possesses a short, stout, and large byssus ear accompanied
by a widely gaping byssus notch. The byssus ear of A. lenaensis (Borissiak 1915, pi. 11,
fig. 10, text-fig. 1) is also similarly large and stout; it is, however, much longer than the
byssus ear of Malayomaorica and A. intermedia. The byssus ears of the Arctotis species
mentioned above differ from the byssus ear of Malayomaorica in the apparently com-
plete absence of the inward bend described above.

Because of the morphological peculiarities of its hinge described above, the articu-
lation of valves of Malayomaorica differs markedly from that of Buchia. In the latter
genus, the byssus ear of the right valve enters the articulation pit of the left valve (Pom-
peckj 1901, pi. XV, figs. 20-21). In Malayomaorica, however (Wandel, 1936, p. 458):
‘dient die Gelenkgrube nicht, wie Krumbeck annimmt, zur Aufnahme des Byssusohres,
sondern zur Aufnahme des wulstig verdickten und schrag nach oben gerichteten Vor-
sprung “v” der Bandplatte der rechten Klappe (Taf. XVII, Fig. lb, c u. 11), der wie ein
Zahn in sie eingreift (vgl. Taf. XVII, Fig. 4b). Das Byssusohr schmiegt sich dem pra-
umbonalen Teil der Bandplatte der linken Klappe an (Taf. XVII, Fig. 4a).’

The above description of Wandel (1936) is quite apt, except that his ‘Gelenkgrube’
corresponds to the furrow-like excavation of the ligamental pit of the writer rather than
to the true articulation pit (= ‘Gelenkgrube’) of Pompeckj (1901).

In MeleagrineUa, Arctotis, and Aucellina, the articulation of valves is essentially
similar to that of Malayomaorica. For Aucellina and MeleagrineUa this is clearly indi-
cated by drawings of Pompeckj (1901, pi. xv, fig. 19; pi. xvi, figs. 4, 6b). For Arctotis this
is clearly indicated by Bodylevsky’s (1960, pi. 7, fig. la) figures of A. intermedia. The
writer was, furthermore, able to check on the reliability of the above data when studying
the still undescribed Canadian representatives of Aucellina and Arctotis.

Malayomaorica malayomaorica (Krumbeck 1923) itself and M. aff. malayomaorica
(Fleming 1958) are the only species that can be placed with assurance in the genus
Malayomaorica at the present time.

The generic nature of'Aucella sp.? aus der Gruppe der A. malayomaorica' (Wandel
1936, pp. 461-2, text-figs, la-c) from the Demu limestone and Lilinta marly limestone
of Misol is uncertain, although the apparent absence of a ligamental ridge in the

J. A. JELETZKY: MALAYOMAORICA GEN. NOV. EROM THE JURASSIC 159

drawings of its right valve does not favour its assignment to Malayomaorica. A detailed
study of well-preserved ligamental plates of the left valves of this form is needed for its
definite generic assignment.

The generic position of ‘Aucella misolica' of Krumbeck (1934, pp. 454-5, pi. xv, fig.
\0a-b) is also uncertain, as the type material of this species consists of a unique, poorly
preserved right valve. It probably is a representative of Malayomaorica, however, as
Biichia aflf. misolica (Krumbeck) from the Lower Tithonian of New Zealand has been
recognized by Fleming (1958, p. 379; 1960, p. 267, table 1) as a younger representative
of Malayomaorica malayomaorica lineage. It is to be hoped that Buchia aflF. misolica
will be described and figured in the near future.

All other Buchia species hitherto described from the Indo-Pacific region seem to be
true representatives of the genus, in the present state of our knowledge at least (compare
Jeletzky 196-).

CONCLUSIONS

'Aucella malayomaorica Krumbeck, 1923’ neither belongs to the genus Buchia
{ = Aucella), as hitherto assumed, nor is closely allied to this genus. It differs from Buchia
in all diagnostic morphological features and so cannot even be placed in the subfamily
Buchiinae Cox, 1953 (= Aucellinae Fischer, 1887).

''Aucella malayomaorica’ is much more closely allied to such aviculopectinid genera
as Meleagrinella, Aucellina, and Arctotis. It is particularly similar morphologically to
the boreal genus Arctotis. From this genus it differs, however, in the furrow-like appear-
ance of the excavation of its left ligamental pit, the occurrence of a tooth-like bulge in
front of the articulation pit (= ‘Gelenkgrube’) of the left valve, and in the marked bend
of the byssus ear toward the left valve. These morphological distinctions fully warrant
the erection of a new genus Malayomaorica for 'Aucella malayomaorica’ , particularly
as it differs even more strongly from Meleagrinella and Aucellina. Malayomaorica is
sufficiently close to Arctotis and Meleagrinella to be included in the subfamily Oxyto-
minae Ichikawa, 1958. Aucellina is also a member of the Oxytominae in the writer’s
opinion. This genus differs strongly from Buchia in most of its diagnostic morphological
features, while being rather similar to Meleagrinella in these same features. It is inferred,
therefore, that Aucellina is only homoeomorphically similar to Buchia, and that it is an
independent offshoot of Meleagrinella.

Malayomaorica is considered provisionally to be an offshoot of Arctotis lineage,
which had penetrated the Indo-Pacific region during the early part of the Upper Jurassic
together with true Buchia species belonging to the Buchia lata-concentrica-mosquensis
species group. The morphological similarities of Malayomaorica to Meleagrinella and
Aucellina are not sufficiently far-reaching to assume its being directly related phylo-
genetically to either of these two genera.

REFERENCES

BODYLEVSKY, V. I. 1960. Novyi pozdneyurskii predstaviteP avikulopektinid Taimyra (A new late
Jurassic representative of aviculopectinids from Taimyr) VNIGI (VSEGEI), Novye vidy drevuikh
rastenii i bespozvoiiocimykh SSSR, Pt. 2, Gosgeoltekhizdat, Moskva, 44-45, pi. 7. (In Russian.)
BOEHM, G. 1911. Grenzschichten zwischen Jura und Kreide von Kawhia (Nordinsel Neuzeelands).
Neues Jb. Min. Geol. Paldont. Bd. I, 1-24, 2 pi.

160

PALAEONTOLOGY, VOLUME 6

BORissiAK, A. A. 1909. Die Pelecypoden der Jura-Ablagerungen im Europaischen Russland. IV. Avicu-
lidae. Mem. Com. GeoL, n.s., 44, 1-26, 2 pi. (In Russian with German summary.)

1915. On Fseiidomonotis (Eiimorphotis) lenaensis Lah. sp. (= Himiites lenaensis Lah.). Tiavaiix

Musee GeoL Pierre le Grand, Acad. Imper. des Sciences, Petrograd, t. VIII, 6, 1914, 141-52, pi. 11
(In Russian.)

CRiCKMAY, c. H. 1932. A new Jurassic ammonite from the coast range of California. American Midland
Naturalist, 13, (1), 1-7, pi. 1.

FLEMING, c. A. 1958. Upper Jurassic fossils and hydrocarbon traces from the Cheviot Hills, North
Canterbury. New Zealand Journ., Geol. and Geophys. 1 (2), 375-94.

1960. The Upper Jurassic sequence at Kawhia, New Zealand, with reference to the ages of some

Tethyan guide fossils. Intern. Geol. Congress, Kept. XXI Sess., Norden, Pt. XXI, Other Subjects,
Copenhagen, 264-9.

GLAESSNER, M. F. 1945. Mesozoic fossils fiom the Central Highlands of New Guinea. Proc. Roy. Soc.,
Victoria, 56, 2, 154.

ICHIKAWA, K. 1958. Zur Taxionomie und Phylogenie der triadischen ‘Pteriidae’ (Lamellibranch.).
Mit besonderer Beriicksichtigung der Gattungen Claraia, Eumorphotis, Oxytoma und Monotis.
Palaeontographica, Abt. A, Bd. HI (5-6), 131-212, pi. 21-24.

JELETZKY, J. A. 1950. Stratigraphy of the west coast of Vancouver Island between Kyuquot Sound and
Esperanza Inlet, British Columbia. Paper Geol. Survey, Canada, 50-37, 1-52.

• 196-. Late Upper Jurassic and Early Lower Cretaceous fossil zones of the Canadian western

Cordillera, British Columbia. Bulletin Geol. Surv., Canada (in press).

KRUMBECK, L. 1923. Zur Kenntnis des Juras der Insel Timor, sowie des Aucellen-Horizontes von Seran
und Burn. Paldontologie von Timor, Stuttgart, Lief. XII, 1-120, 6 pi.

1934. Die Aucellen des Malms von Misol. Neues Jb. Min. Geol. Paldont. Beil. Bd. 71b, 422-69,

pi. 14-16.

MARWICK, J. 1926. Myalinidae from the Jurassic of New Zealand, a new genus and new species. Trans.
& Proceed., New Zealand Institute, 56 (new issue), 304-6, pi. 71.

1953. Divisions and faunas of the Hokonui System (Triassic and Jurassic). Bull. Geol. Survey,

New Zealand, Pal. 21, 96.

POMPECKJ, J. p. 1901. Ueber Aucellen und Aucellen-ahnliche Eormen. Neues Jb. Min. Geol. Paldont.
Beil. Bd. XIV, 319-68, pi. 15-17.

SOKOLOV, D. 1908. Aucellen vom Timan und von Spitzbergen. Mem. Com. Geol., n.s., 36, 1-29 3 pi.
(In Russian with German summary).

1912. Originaly i paratipy K. E. Roulie i G. A. Trautshol’da v kollekstii Earenkolya iz Gal'iovoi

(Types and paratypes of K. E. Rouillier and G. A. Trautshold in Eahrenkohl’s collection from
Gal'iova). Travaux Musee Geol. Pierre le Grand, Acad. Imper. Sciences, St. Petersburg, t. VI (4),
97-1 19, 2 pi. (In Russian.)

STOLLEY, E. 1929. Uber Ostindische Jura-Belemniten. Paldontologie von Timor, Lief. 16, 91-213, 9 pi.,
1 pi. in text.

WANDEL, G. 1936. Beitrage zur Kenntnis der jurassischen Molluskenfauna von Misol, Ost-Celebes,
Buton, Seran und Jamdena. Neues Jb. Min. Geol. Paldont. Beil. Bd. 75b, 447-526, pi. 15-20.

J. A. JELETZKY

Geological Survey of Canada,
601 Booth Street,
Ottawa, Ontario,

Manuscript received 24 April 1962 Canada

CYPRILEPAS HOLMI WILLS 1962,

A PEDUNCULATE CIRRIPEDE FROM THE
UPPER SILURIAN OF OESEL, ESTHONIA

by LEONARD J. WILLS

ABSTRACT. An account is given of about fifty examples of a pedunculate cirripede, Cyprilepas holini Wills 1962,
some still attached to the chitinous skin of Euryptenis fischeri Eichw. The bivalve chitinous shell and attachment
peduncle of the cirripede are described. Comparison is made with the Cypris-\3.rva. and early post-larval stages of
the present-day Lepas.

Biologists and ecologists, as well as palaeontologists, may be interested in the dis-
covery of what appears to be a pedunculate cirripede in rocks as ancient as the Silurian ;
and to find that this sessile animal had a bivalve chitinous shell not unlike that of the
Cypm-larva invariably present in all cirripedes. Because of this resemblance to
the Cypns-lsLTva, the fossil has been named Cyprilepas holnii (Wills 1962, p. 567), the
specific name being in honour of Gerhard Holm, who in his classic paper (1898) on
Eurypterus fischeri Eichw. figured one on his pi. 4, fig. 22 as ‘ Kiemenblatter ?, zwei
zusammenhaftende’ of Eurypterus, although on p. 39 he rejected this ascription.

Discovery and mode of occurrence. While engaged in etching out the chitinous skins of
E. fischeri from the silty Upper Silurian limestone of Oesel, Esthonia, I cut one specimen
(ElO) longitudinally into two parts (R. and L.) before treatment with acid. As etching
proceeded I noticed about ten of the small objects which form the subject of this paper
waving about in the acid. All were clustered round the first bit of Eurypterus to appear
(PI. 22, figs. 1, 2). The majority lay round its edges, but a few were on its flattened surface
(PI. 22, fig. 3). Most of them floated free as the etching proceeded, but in one or two
cases I had to prise them off with a fine needle. Ultimately I extracted more than fifty
from the two halves of the specimen, and a number can still be observed attached to
various appendages now in the final mount, BU 739, and on two leg-joints mounted as
BU 748. A few were seen when etching another specimen. As noted above. Holm also
isolated one example from the same locality.

As the etching of ElO proceeded it became clear that this specimen of Eurypterus
was a moulted skin that had lain ventral side up on the sea-bed, and had then been
partially covered by calcareous silt. Gentle currents had carried away the metastoma,
and displaced and broken up the greater part of the last two appendages, bits of which
were found well away from the body. Since all the shells were attached to upward-
stretching parts of the appendages or to the drifted bits, it seems certain that these parts
were exposed above the layer of silt and were solid objects to which the floating larvae
of this sessile organism could anchor themselves.

Morphology. The fossils consist of a chitinous bivalve shell, not unlike a small Isaura
[Esther ia] or a Cypris, and a cylindrieal chitinous attachment-stalk, both now flattened.

(Palaeontology, Vol. 6, Part 1, 1963, pp. 161-5, pi. 22.]

C 1015

M

162

PALAEONTOLOGY, VOLUME 6

By analogy with modern pedunculate cirripedes these two parts may be regarded as the
capitulum and peduncle respectively, joined at the anterior end of the shell. The com-
bined length of shell and stalk varies from about 2-2 mm. to 4-8 mm. (PI. 22, figs. 4-9).

(a) The shell. The two valves of the shell are always squashed flat with some dis-
tortion and folding. Between the two valves there is usually a small amount of silt, but
in no case have I observed any evidence of a cavity such as would be expected if the
fossil, prior to acid etching, had had calcareous plates comparable to those that reinforce
the very tenuous chitin of the bivalve outer-skin of a post-larval present-day Lepas.
Usually the valves lie fairly symmetrically one above the other, and often it appears as
if one is larger than the other, as in ostracods (PI. 22, figs. 5, 6). In this condition most
are roughly oval, approximately two-thirds as wide as long; but the smaller ones have
almost circular outlines. The length of the shell, apart from the stalk, varies from 1-4 to
3-0 mm. In a few cases the compression has opened the two valves at the posterior end
(PI. 22, figs. 7, 8). The hinge appears to occupy the full length of the carinal side and to
extend part-way across the rather blunt anterior end to the points where the free edges
of the two valves merge on either side into the skin of the stalk. The edges of the shells
are slightly thickened. The valves have been crushed irregularly, but show no growth-
lines at all.

The chitin of a single valve appears structureless at low magnifications, but under
a high power it displays a faint meandering mesh-pattern (PI. 22, fig. 11). On three
specimens, however, there is a definite hexagonal patterning visible on the chitin (PI. 22,

EXPLANATION OF PLATE 22

Cyprilepas and Lepas.

Figs. 1-12. Cyprilepas holmi Wills, attached to or associated with Eurypterus fischeri Eichw., Upper

Silurian, Oesel, Esthonia.

I, 2. Two views of a single specimen attached to an edge of a segment of an appendage of E. fischeri
(black), moving about in the etching acid which was disturbed between successive exposures, thus
altering the posture of the shell.

3. Two segments of an appendage of E. fischeri with four specimens of Cyprilepas (outlined in ink)
held virtually in their original positions by ‘Durofix’ which is itself blurred by adherent silt-
particles. BU 748.

4-6. Young examples, two possibly showing the attachment-disc {di). BU 744, 750, 749 (para-
types).

7-9. Larger examples showing the two valves of the shell and variations in the stalk as it is pre-
served. BU 745, 746 (paratypes), and 743 (holotype).

10. Hexagonal patterning on part of the shell of the holotype (BU 743). Cf. fig. 14.

II. Linear markings on a single valve of the shell. BU 747.

12. ? Papillae seen through the double skin of the stalk (BU 745). Cf. fig. 15.

Figs. 13-15. Lepas australis Darwin. Australian and New Zealand waters. Recent.

13. Late CyprisAdiiva showing rudiments of the calcareous plates (ic, scutum; te, tergum; ca,
Carina) together with the hair-clad stalk (ha) of the future post-larval animal, all within the
chitinous bivalve shell (sh) of the Cypm-larva with its attachment-disc (di). Cirri (ci).

14. Part of the chitinous shell of a Cypris-larva at the stage seen in fig. 13, after decalcification,
showing the impress of the hexagonal patterning on one of the calcareous plates. Cf. fig. 10.

15. A detached post-larval individual after the first post-attachment moult and the loss of the
Cypris-Uke shell; pa, papillate stalk (cf. fig. 12); other lettering as in fig. 13.

Palaeontology, Vo I, 6

PLATE 22

Scale
Figs. 4-9

14

I lO'OSmm

1mm

1mm

I 0-05 mm 10

1 0-05 mm 11

,0-05 mm 12

WILLS, Upper Silurian pedunculate cirripede

L. J. WILLS; A SILURIAN PEDUNCULATE CIRRIPEDE

163

fig. 10). The significance of this as evidence for the original existence of calcareous plates
within the shell is discussed below.

{b) The peduncle, now compressed, was a cylindrical structure opening out from the
body of the capitulum at the anterior end of the hinge. In some cases it can be seen that
its sides are continuations of the two valves. At the distal end where it was attached to
the Eiiryptems, there is either a sucker-like disc (PI. 22, fig. 6) or a sharp point (PI. 22,
figs. 5, 7). The latter is probably due to fracture, but the disc has only been seen in two
or three cases. The stalk may be a simple tapered tube (PI. 22, figs. 5, 8), but more
usually is divided into from two to four parts by transverse bands that appear to be
either thickenings or, more probably, folds of the skin. The bands are usually obscured
by silt. Their number bears no obvious relationship to the length of the stalk. I incline
to the view that they functioned to give flexibility and perhaps muscle attachments in
what must have been an organ capable of retraction and of bending.

The texture of the chitin of the stalk is sometimes distinctive. It then appears to be
ornamented by transverse rows of drop-like markings which are probably minute
papillae on the two superimposed layers of skin (PI. 22, fig. 12).

The ratio of stalk-length to shell-length lies between 2 and 3, but it varies irregularly
— in some cases, no doubt, because the stalk is incomplete.

Affinities. Modern bivalve shells that are sessile belong either to the Mollusca, Brachio-
poda, or Crustacea. Only the last (in the pedunculate Cirripedia, such as the Lepadidae)
presents any points of similarity to Cyprilepas.

For purposes of comparison, I obtained through the kindness of Dr. B. Patel of the
Menai Straits Marine Biology Station and of Dr. T. M. Skerman of the New Zealand
Oceanographic Institute, both Cv/^rw-larvae and young adults of Lepas. In these one
can recognize the growth stages described by Caiman (1909).

Every Cirripede begins as a Nauplius-ldLXva. and then passes through a Cy/?m-larva
stage. In the latter the animal is enclosed within a bivalve chitinous shell. At first the
larva is free : later it attaches itself to some object by a disc secreted by its two anten-
nules. In the later development of the Cypris-loxya. (PI. 22, fig. 13) there appears inside
the chitinous Cypris-Wkt shell, a single hair-clad stalk (the pair of antennules is no
longer visible) connecting the disc (external to the shell) to the capitulum wliich dis-
plays rudiments of five calcareous plates {ca, sc, te). The plates are milky-white struc-
tures exhibiting a conspicuous hexagonal surface-patterning.

After the first moult the Cypris-like, shell disappears and a miniature adult (PI. 22,
fig. 1 5) now has its soft parts contained within two valves of tenuous chitin lined by five
calcareous plates arranged in two pairs with the fifth or carinal plate along the hinge.
The outside of the plates has the same hexagonal patterning as in the larva. The cover-
ing of very tenuous chitin thickens somewhat in the sutural strips between the plates.
This whole structure is the capitulum. Its chitin is continued into a strong papillate
peduncle.

The shells of the Cypm-larva of Lepas are all about the same size, c. 1-6 mm. in
length. The youngest post-larval individual that 1 have seen has a total length of 1 -8 mm.,
of which T3 mm. is shell. The smallest known Cyprilepas has a total length of 2-2 mm.,
of which 1-3 mm. is shell. These figures suggest that all the known specimens of Cypri-
lepas were early post-larval individuals.

164

PALAEONTOLOGY, VOLUME 6

In order to get a true comparison of the young Lepas with the fossils which had been
extracted by an acid etch, and consequently must have lost any calcareous plates that
may have been present originally, one Cypm-larva at the stage represented by Plate 22,
fig. 13, and several very small post-larval individuals like the one shown on Plate 22,
fig. 15, were treated with acid to remove the calcareous plates; and the soft parts,
including as much of the lining of the valves as possible, were dissected away. A very
tenuous film of chitin remained, which shows :

{a) in the larva, a very distinct patterning of hexagons (PI. 22, fig. 14) representing the
impress of the ends of the prismatic columns that formed the rudimentary calcareous
plates, but no growth-lines to define the limits of the plates;

{h) in the post-larval individuals, a similar hexagonal patterning is to be seen, but only
on certain small areas of each plate. The outlines of the plates, however, are clearly \
indicated by narrow growth-lines conspicuous round their edges where the plate i
adjoined a sutural strip. Earlier growth-lines, where seen, are extremely narrow. j

There is a remarkable resemblance between the hexagonal patterning on the inside j
of the decalcified chitin of Lepas and the admittedly less-distinctly hexagonal markings (
seen on three specimens of Cyprilepas, noted above in the description of the shell. It \
remains a matter of speculation whether the latter can be accepted as valid evidence of
the original presence of a calcareous plate or plates within its shell; but the complete
absence from the shell of any growth-lines or thickened sutural strips, makes it certain
that there were not five plates as in Lepas. If there were plates at all, there were two,
one lining the inside of each chitinous valve, and perhaps leaving their impress only on
certain parts of it, as was noted in the post-larval Lepas.

The peduncle seen within the CyprA-larva shell of Lepas is clad with hairs (PI. 22,
fig. 13/7a), but after the first moult it is covered with papillae (PI. 22, fig. \5pa). The
markings seen on some of the stalks of Cyprilepas (PI. 22, fig. 12) probably represent
papillae, but in no case has it been possible to find only a single layer of chitin on which
to check this point. All that can be claimed is that the peduncle skin structure in the
fossil resembles that of the post-larval rather than that of the larval state in Lepas.

The attachment-disc of Lepas is conspicuous in the advanced CyT^ra-larva. As growth
proceeds it becomes obscured by the stalk, and fully adherent to the supporting object.

In line with this is the observation that in the fossils it is on the smaller individuals that
the disc can be recognized with some certainty (PI. 22, figs. 4, 6).

CONCLUSIONS

j

1. The capitulum of Cyprilepas consisted of a bivalve chitinous shell resembling !
that of a Cypris-\2iXV2L of a cirripede, but equally like that of an adult entomostracan,
such as Isaura [Estheria] or Cypris. This chitinous shell may have been reinforced
by a lining of calcareous matter possessing a prismatic texture which produced a
hexagonal patterning on the chitin that has occasionally been preserved. There is no
evidence that such calcareous matter, if it existed, formed five independent plates, like
those in Recent pedunculate lepadid cirripedes.

2. The peduncle of Cyprilepas resembles that found in early post-larval stages of
Lepas in having a thick papillate chitinous sheath. The constrictions usually seen in it
in the fossils are not matched in any of the small adults of Lepas that I have examined.

L. J. WILLS: A SILURIAN PEDUNCULATE CIRRIPEDE

165

3. The size of the whole organism and the possession of a papillate peduncle show
that Cyprilepas was an early post-larval stage of a lepadomorph pedunculate cirripede,
differing from young post-larval Lepadidae in the absence of the five calcareous plates
forming the capitulum of the latter, and from Scalpellidae in the absence of numerous
plates from the capitulum and of calcareous scales from the peduncle.

4. If Cyprilepas be accepted as a lepadomorph cirripede, the range of that sub-order
is extended from the Middle Carboniferous {Praelepas jaworskii Tschernischew) to the
Upper Silurian (the Ordovician and Silurian machaeridians are now excluded from the
cirripedes). The Lepadidae are not known before the Eocene.

Specimens with the prefix BU are in the collection of types and figured specimens in
the Department of Geology, University of Birmingham. Of the figured specimens,
BU 743 on Plate 22, fig. 9, is the holotype; the others, BU 744-50, are paratypes.

REFERENCES

CALMAN, w. T. 1909. Crustacca. In A Treatise on Zoology: Part VII, Appendiculata, ed. E. Ray
Lankester. London.

HOLM, G. 1898. Ueber die Organisation des Eurvpterus fischeri Eichw. Mem. Acad. Sci. St. Peters-
boiirg, Ser. VIII, 8 (2), 1-57.

WILLS, L. J. 1962. A pedunculate cirripede from the Upper Silurian of Oesel, Esthonia. Nature,
Land. 194, 567.

LEONARD J. WILLS
Farley Cottage,
Romsley,

Manuscript received 5 July 1962 Near Birmingham

THE BRYOZOAN GENUS POLYPORA M‘COY

by T. G. MILLER

ABSTRACT. The gcnus Polypora M‘Coy 1844 is redefined by reference to the type material. M‘Coy’s four original
species of Polypora are redescribed, one being only tentatively retained within the genus, and one being assigned
to Fenestella Lonsdale.

During the last 118 years more than 250 species of fenestrate bryozoa have been assigned
to the genus Polypora. It is probable that imperfect knowledge of the characters of the
genus has led to inclusion of species with quite different affinities. Shulga-Nesterenko,
Morozova, and Nekhoroshev have attempted to subdivide the genus into species-
groups based on the number of rows of zooecial chambers in the branches. In the light
of the present re-examination it seems likely that a more valid subdivision of what is
certainly an unwieldy genus will be based on the presence or absence of nodes on the
obverse of the branches, and the arrangement of these with respect to a central carina,
together with consideration of the organization and location of the zooecial chambers in
relation to the ‘basal plate’.

SYSTEMATIC DESCRIPTIONS
Specimens with the prefix NMI are in the National Museum of Ireland, Dublin.

Order cryptostomata Shrubsole and Vine 1882
Family fenestellidae King 1850
Genus polypora M‘Coy 1 844

Type species. Polypora dendroides M‘Coy 1844, p. 206; pi. 29, fig. 9; Carboniferous, Tournaisian,
Ireland.

lyPCofs diagnosis. ‘Expanding, interstices round, branching, having on one side from
three to five rows of pores, the margin of which is never raised, interstices connected by
thin, transverse, nonporiferous dissepiments.’

Emended diagnosis. Unifoliate fenestrate expansion of straight or slightly sinuous,
regularly bifurcating, zooecia-bearing, non-carinate branches joined at regular intervals
by transverse non-poriferous dissepiments. Branches circular or elliptical in cross-section,
carrying regularly spaced elevated nodes on the central line of the obverse. Zooecia
regularly arranged in three or more rows in the branches, the number increased for
a short length below a bifurcation and similarly diminished above. In cross-section the
zooecial chambers lie side by side on a flat basal plate in the plane of the expansion. In
deep transverse section the zooecial base-shape is elongate-hexagonal to irregular poly-
gonal. In shallow transverse section the calcareous investment of the branches is seen to
be perforated by small ‘tubules’ which may be locally clustered, particularly round the
margin of the zooecial apertures, and more compactly to form the cores of the branch

[Palaeontology, Vol. 6, Part 1, 1963, pp. 166-71, pi. 23-24.]

T. G. MILLER: THE BRYOZOAN GENUS POLYPORA M'COY 167

nodes. Obverse of zoarium smooth, granular, or striated. Reverse similar to obverse but
may also carry more or less regularly spaced node-like projections.

Polypora dendroides M‘Coy
Plate 23, figs. 1-3

Polypora dendroides M‘Coy 1844, p. 206; pi. 29, fig. 9.

NPCofs description. ‘Flat, fan-shaped, interstices thick, regularly branched, divari-
cating; dissepiments very thin, frequently oblique, and placed at nearly equal distances;
fenestrules large, rhomboidal, poriferous face, with five rows of small impressed pores
arranged in quincunx, reverse longitudinally striated.

‘This beautiful species is chiefly remarkable for the degree of divergence with which the
interstices branch, and the small number of rows of pores. Length one inch, width one
and a half inches; the interstices are about one line apart and half a line thick.’

Material. (1) Lectotype NMI XXIX.9a, Carboniferous ‘Slate’, Tournaisian, Hook Head, Fethard, Co.
Wexford, Eire. Fragment of zoarium, roughly rectangular (30 X 25 mm.), showing obverse, on matrix
of hard dark grey calcareous mudstone full of crinoidal and bryozoan fragments. (2) Homeotype
NMI G.39:1962, Upper Tournaisian, Hook Head, Co. Wexford, Eire. Fragment of zoarium
(25 X 20 mm.), partly obscured, showing obverse, in matrix of hard grey shaly bioclastic limestone.

Micrometric formulae. (1) Lectotype NMI XXIX.9a. (2) Homeotype NMI G.39:1962. (3) Polypora
bukhtarmensiformis Nekhoroshev 1956, p. 210, Lower Carboniferous Ulbinskaya suite, Rudnyi Altai,
and Fominskaya zone, Kuznetsk Basin.

(For explanation of column headings see Miller 1962, p. 120; in addition, R/B = number of rows of
zooecia in branch, ND = base-diameter of branch nodes.)

R/B

B/10

D/10

Z/5

N/5

B„ (mm.)

ZD (mm.)

ND (mm.)

1.

i 4 6

6-8

4-5-1

13-16

5-7

0-5-0-9

0-15X0-2

0-13

2.

4

7-11

5

13-U15-^

5-7

0-6-0-8

019

0-13

3.

4-6

6-8

4k-5k

14-16

7

0-8-0-85

0-12X0-18

7

Description. Zoarium probably flat, fan-shaped. Branches thick, straight, sub-elliptical
in cross-section. Dissepiments short, half the diameter of the branches. Fenestrules
long, with rounded ends, occasionally oval, width about equal to the diameter of the
branches. Zooecia normally in four rows, but increased to six below a bifurcation, and
diminished to three immediately above. Zooecial apertures oval, margins minutely sub-
asteroid due to the presence of fifteen or sixteen short thin internal projections resem-
bling the septa of corals. Branch nodes on obverse circular at base, high conical in
profile, with grooved sides and rounded perforate tops. Zooecial bases elongate-
hexagonal (eH) to irregular polygonal.

Discussion. The lectotype is considered to be the one used by M‘Coy for his lower figure.
In the Griffith Collection of the National Museum of Ireland it is labelled ‘? Type for
structure’. Under the same catalogue number (XXIX. 9) is a second specimen, hitherto
supposed to be a syntype, labelled ‘? Type for form’. This is a fragment of a zoarium,
roughly semicircular, 40x 25 mm., showing the reverse, in a matrix of pale grey crystal-
line limestone, from the Carboniferous ‘Upper’ Limestone of Blacklion, Co. Cavan,
Eire. The rock at this locality is a reef-facies limestone of Upper Visean (B.2) age. It is

168

PALAEONTOLOGY, VOLUME 6

clear that this second specimen (now labelled XXIX.9Z)), which, in spite of showing only
the reverse of a zoarium, must have been used for M‘Coy’s upper figure, is not conspecific
with its companion, and must therefore be discarded from the type material of Polypora
dendroides. It is probably a specimen of P. (?) verrucosa (see p. 169), but as the obverse
is not visible a precise determination cannot be made.

In shallow transverse section the finely granular (or minutely porous) calcareous
tissue of the main zoarial skeleton is seen to be penetrated by small ‘tubules’. These
appear in the section as clear spots surrounded by a darker zone without a sharp
margin. The tubules are 10-1 2, u in diameter and are distributed fairly evenly in the
skeletal tissue 40-80/.1 apart. This wide separation of the tubules is replaced by a closer
packing in two situations : (a) round the margin of the zooecial apertures, where fifteen or
sixteen are placed against the margin, about their own diameter apart, or a little more
(12-1 5 /a); and (b) at the site of the branch nodes, where usually eight tubules appear to
bend together and coalesce into an asteroid space surrounding a solid central structure.
Similar structures were figured (but not discussed ) by Ulrich (1 890) in Polypora cesteriensis
Ulrich and P. halliana Prout.

This formation of branch nodes by a growing together or bunching of tubules suggests
that the carinal nodes in Fenesteda may have a similar structure. The tentative interpre-
tation by Bassler (1953, p. 120) of these carinal nodes as possibly homologous with the
acanthopores of the Trepostomata — an interpretation followed by me (Miller 1961,
p. 223) — may have to be revised. It is interesting to note in this connexion that in general
appearance and arrangement the tubules in Polypora recall the micracanthopores of
the Ordovician bifoliate cryptostome genus Pachydictya recently described by Phillips
(1960, pi. 5, fig. 4).

Polypora marginata M‘Coy
Plate 24, fig. 3

Polypora marginata M'Coy 1844, p. 206; pi. 29, fig. 5.

M' Coy's description. ‘Interstices thick, irregularly bifurcate ; sides margined ; dissepiments
thin; fenestrules small, elongate, oval, or approaching to a square form; reverse with
direct, deep, longitudinal striae; poriferous surface, with five alternating rows of pores,
and interjacent, waved striae.

‘The broad, flat, margin of the branches distinguishes this rare species at a glance
from any of the other corals likely to be confounded with it.’

Material. Holotype NMI XXIX. 5, Carboniferous ‘Upper’ Limestone, Upper Visean (horizon in
Upper Dibunophyllum zone (Dj = Pp, fide Fowler and Robbie 1961, p. 80), Killymeal, Dungannon,
Co. Tyrone, Northern Ireland. Fragment of zoarium, 1 1 X 25 mm., showing reverse, on matrix of dark
grey compact crystalline limestone.

EXPLANATION OF PLATE 23

Figs. l-3i. Polypora dendroidesyVCoy. 1, Obverse of zoarial fragment, X 4-5. Holotype NMI XXI.X.9a.
2, Obverse, showing arrangement of zooecial apertures on branch surface, x 20. Holotype NMI
XXIX.9(7. 3, Shallow transverse section showing arrangement of tubules round apertural margins,
and at site of a branch node (irregularly shaped light patch with dark diffuse margin at top right),
X 100. Homeotype NMI G.39:1962.

Palaeontology, Vol. 6

PLATE 23

MILLER, Polypora

T. G. MILLER: THE BRYOZOAN GENUS POLYPORA M‘COY

169

Micronietric formula

R/B

B/10

D/10

Z/5

N/5

Bvv (mm.)

ZD (mm.)

ZB

NMI XXIX.5

4 5 8

5-9

4-5

15J-18

7

0-6-M

0-1 2-0- 15

eH

Description. Branches thick, rather sinuous, compressed-elliptical in cross-section.
Dissepiments thin. Fenestrules rectangular. Zooecial chambers in five rows, increasing
to eight below, and diminishing to four above, a bifurcation. Zooecial apertures small,
circular, slightly less than their own diameter apart; the outer rows commonly projecting
slightly into the fenestrule so as to produce a sinuous margin. Zooecial bases elongate-
hexagonal. Details of branch nodes unknown. Reverse minutely striated parallel to long
axes of branches and dissepiments, without auxiliary projections.

PoJypora (?) verrucosa M^Coy
Plate 24, fig. 2

Polypora verrucosa M‘Coy 1844, p. 206; pi. 29, fig. 6.

M'Coy s descripton. ‘Interstices rarely bifurcating, regular, equal, rounded; dissepiments
thin, distant; fenestrules rectangular, five times as long as wide, about one-third wider
than the interstices, equal; obverse with four rows of prominent, wart-like pores, about
ten in each row to the length of a fenestrule; between the pores are waving longitudinal
striae; reverse nearly smooth.’

Material. Holotype NMI XXIX.6, Carboniferous ‘Upper' Limestone, Upper Visean (horizon in reef-
facies, upper Beyrichoceras zone (Bj = T>p,fide Hodson 1956), Blacklion, Co. Cavan (near Enniskillen,
Co. Fermanagh), Eire. Fragment of zoarium, 25 X 10 mm., showing patches of obverse, on matrix of
grey crystalline limestone full of fenestellid bryozoan fronds.

Micrometric formula

R/B

B/10

D/10

Z/5

N/5

B^v (mm.)

ZD (mm.)

ZB

NMI XXIX.6

3 4 5

5i-74

2-3

11-12

7

0-48-0-58

01 0-0- 13

7

Description. Branches of moderate thickness, straight, subparallel, circular in cross-
section, dissepiments thin. Fenestrules long, narrow, rectangular. Zooecial chambers in
four rows, increasing to five below, and diminishing to three above, a bifurcation.
Zooecial apertures relatively small, circular, with prominent peristomal collars, placed
nearly twice their diameter apart along the branches; apertures on the outer rows
project slightly into the fenestrules so as to produce a beaded effect. Obverse surface
deeply sculptured in ridges and grooves streamlined past and between the zooecial
apertures. No nodes on the branches. Reverse with fine longitudinal striations. Shape of
zooeeial base unknown.

Discussion. The striking feature of this species is the absence of nodal structures from
the centre line of the branches on the obverse, combined with the ridged and grooved
sculpturing of the calcareous investing tissue. Both these characters recall the condition
in the fenestellid genus Levifenestella Miller 1961. The question whether M‘Coy’s P. ver-
rucosa should be detached as a member of a genus distinct from Polypora must await
the recognition of similarly characterized species in other bryozoan faunal assemblages.
For the present it is tentatively assigned as P. (?) verrucosa.

170

PALAEONTOLOGY, VOLUME 6

Genus fenestella Lonsdale 1839
Fenestalla papillata (M‘Coy)

Plate 24, fig. 1

Polypora papillata M‘Coy 1844, p. 206, pi. 29, fig. 10.

M' Coy' s description. ‘Interstices rarely bifurcating, very narrow, rounded ; fenestrules cir-
cular or oval; poriferous surface smooth, with three alternating rows of pores; reverse
smooth, with a small papillated pore at the origin of most of the dissepiments.

‘This species is remarkable for its large and round fenestrules, which are of such a size
that the interstices and dissepiments appear disproportionably thin. The prominent
pores on the outer side have caused me to doubt the propriety of placing this coral in
the same genus with the preceding species [P. marginata]. It is very rare.’

Material. Holotype NMI XXIX. 10, Carboniferous ‘Upper’ Limestone, Upper Visean (horizon in
reef-facies, upper Beyriclioceras zone (Bj = Di), fide Hodson 1956), Blacklion, Co. Cavan (near
Enniskillen, Co. Fermanagh), Eire. Fragment of zoarium, 20x20 mm., showing reverse, on matrix of
massive grey calcilutite.

Micrometric formulae

B/10

D/10

Z/5

N/5

B^ (mm.)

ZD (mm.)

ZB

1. NMI XXIX.IO

16

8

15-17

27-30

0-30-0-40

0-10

hH

2. F. polynodosa Miller
1961 ...

12-17

7-9

15-18

32-36

0-30-0-40

0-15

sT

Description. Normal reticulate fenestellid meshwork of carinate zooecia-bearing branches
with inter-branch dissepiments. Branches relatively stout, straight, or slightly sinuous.
Dissepiments thinner than the branches. Carinae narrow, with a row of small, closely
set nodes with elliptical bases. At a few places the line of carinal nodes becomes sinuous,
the nodes lying alternately on either side of the central line, thus approaching the con-
dition of the carinal nodes in Minilya Crockford 1944. Zooecial apertures large, circular,
slightly more than their own diameter apart, with prominent peristomal collars, indenting
the fenestrules, four or five to a fenestrule. Zooecial bases of variable shape, from tri-
angular through subtriangular (sT) to hemi-hexagonal (hH). Reverse longitudinally
striated, with scattered non-perforate steep-sided elevations, some of which lie opposite
the ends of dissepiments.

Discussion. It is difficult to understand M‘Coy’s attribution of this species to his genus
Polypora. The holotype, which is undoubtedly the one figured by M‘Coy, shows the
reverse of a zoarial fragment. By grinding down part of this fragment it has been possible
to show the characteristic fenestellid features of the obverse. There is nowhere any trace

EXPLANATION OF PLATE 24

Fig. 1. Fenestella papillata (M‘Coy). Holotype NMI XXIX. 10, polished surface slightly oblique to
plane of zoarial expansion showing at top left the arrangement of carinal nodes, x 17.

Fig. 2. Polypora (?) verrucosa M‘Coy. Holotype NMI XXIX.6, part of obverse, x 16.

Fig. 3. Polypora marginata M‘Coy. Holotype NMI XXIX.5. Reverse of zoarium with partial trans-
verse section (polished surface) at left, showing arrangement of zooecial apertures, x 14.

Palaeontology, Vol. 6

PLATE 24

MILLER, Polypora

T. G. MILLER: THE BRYOZOAN GENUS POLYPORA M‘COY

171

of the ‘three alternating rows of pores’ of M‘Coy’s description. On the contrary, the
typical fenestellid rows of zooecial apertures separated by a noded carina can be clearly
distinguished.

F. papillata is close to F. polynodosa Miller from the Irish Tournaisian. It differs in
having a slightly smaller number of zooecial apertures to a fenestrule, in the smaller
zooecial apertures, and in the slightly less closely packed carinal nodes. Also, the zooecial
base in F. papillata shows a strong tendency to become stabilized as hemi-hexagonal,
and is distinguishable from the predominantly subtriangular base in F. polynodosa.

Acknowledgements. I am indebted to the Director of the National Museum of Ireland, for allowing me to
examine further material from the Griffith Collection; and to Dr. J. S. Jackson, Keeper of the Natural
History Division of the same Museum, for information about the stratigraphic position of the various
type-localities.

REFERENCES

BASSLER, R. s. 1953. Treatise on Invertebrate Palaeontology (R. C. Moore, Ed.), Part G: Bryozoa,
xiii+253 pp. Univ. Kansas Press and Geol. Soc. Amer.

FOWLER, A. and robbie, j. a., 1961. Geology of the country round Dungannon. Mem. Geol. Siirv. U.K.
HODSON, F. 1956. Carboniferous goniatites from Blacklion, Co. Leitrim {sic), Ireland. Geol. Mag.
93, 41,42.

M‘COY, F. 1844. A Synopsis of the Characters of the Carboniferous Limestone Fossils of Ireland. 207 pp.,
29 pi. Dublin.

MILLER, T. G. 1961. New Irish Tournaisian fenestellids. Geol. Mag. 98, 493-500.

1962. North American species of FenesteUa from the Carboniferous of Great Britain and Ireland.

J. Paleont. 36, 120-5.

NEKHOROSHEV, V. p. 1956. Lower Carboniferous Bryozoa of the Altai and Siberia. Trudy vses. nauch.
issled. geol. Inst., n.s., 13, 419 pp., 57 pi.

PHILLIPS, J. R. p. 1960. Restudy of types of seven Ordovician bifoliate Bryozoa. Palaeontology, 3, 1-25.
ULRICH, E. o. 1890. Palaeozoic Bryozoa. In Palaeontology of Illinois, Geol. Surv. HI. 8, 285-688.

T. G. MILLER

Department of Geology,
University of Keele,
Keele, Staffs.

Manuscript received 28 July 1962

CUPULADRIA CANARIENSIS (BUSK)— PORTRAIT

OF A BRYOZOAN

by R. LAGAAIJ

ABSTRACT. The lunulitiform bryozoan Cupiiladria canariensis (Busk) is a benthonic marine organism, whose
calcareous colonies can easily be recognized with a hand-lens or under the microscope.

This species is eurybenthic (2-> 300 fathoms), eurythermal (12-31° C.), reasonably euryhaline (28-37%„),
and requires a stable quartz and/or carbonate sand bottom. It is at present widely distributed over the continental
shelves of the Atlantic and East Pacific between the 14° C. surface isocrymes and had an equally wide distribution
during the Late Tertiary and Quaternary.

The occurrence of C. canariensis in Miocene and Pliocene marine sediments of the southern part of the North
Sea basin calls for sea-water surface temperatures at least 9° C. higher than those obtaining at present in this
area. Its occurrence in the Miocene, Pliocene, and Early Pleistocene marine sediments of Spain, Italy, and Rhodes
suggests that the Mediterranean was somewhat less saline in the geological past than it is at present.

Although lunulitiform Bryozoa range from the Upper Cretaceous to Recent, C. canariensis first appears in
the Lower Miocene (Aquitanian). Its presence or abseirce among lunulitiform bryozoan assemblages may serve
as a criterion for establishing the Oligocene-Miocene boundary in sequences of ancient tropical and subtropical
shelf sediments on both sides of the Atlantic. On this criterion a large part of the ‘Caribbean Oligocene' is to be
considered as Lower Miocene.

Cupuladria canariensis was named in 1859 by the English bryozoologist George Busk,
who discovered it in material collected from the sea bed in the neighbourhood of
Madeira and the Canary Islands. Later in the same year he reported its occurrence in
the Pliocene Coralline Crag in East Anglia. In the last hundred years the number of
records has increased enormously, and the data are widely dispersed in the biological
and palaeontological literature. It seemed worth while to try to assemble these widely
scattered data so as to give a comprehensive picture of the species in space and time.

Such a synthesis is of considerable geological interest. It will be shown that a study of
Cupuladria canariensis cdLn not only give an insight into certain ecological, climatological,
and hydrological conditions in the geological past, but can also help resolve the problem
of determining the Oligocene-Miocene boundary in sequences of ancient tropical and
subtropical shelf deposits.

Its interest for the palaeo-ecologist is implied in a variant of Grimsdale’s golden rule
for systematic palaeontologists: . . one detected synonym is worth from ten to one

hundred new species’ (1951, p. 467). Perhaps this account will help to establish that one
ecologically well-known species is worth more than a hundred tabulae rasae.

ZOOGEOGRAPHY AND ECOLOGY

1. Distribution

Cupuladria canariensis (Bryozoa: Cheilostomata, Anasca) belongs to the so-called
lunulitiform Bryozoa (PI. 25, fig. 1), all of which possess the same zoarial form as the
genus Lunulites Lamarck, 1816. The fully grown colony, or zoarium, has the shape of
a dome or flattened cone and consists of a single layer of cells, or zooecia, opening on
the convex side of the dome. In addition to the normal cells, or autozooecia, in which

(Palaeontology, Vol. 6, Part 1, 1963, pp. 172-217, pi. 25-26.]

R. LAGAAIJ; THE BRYOZOAN CUPULADRIA CANARIENSIS 173

the polypides reside, there are other cells, the vibracula, each bearing a long, whip-like
vibracular seta. In the genus Cupiiladria there is, without exception, a vibraculum situ-
ated distally to each autozooecium ; the tip of each seta is capable of describing a 180°
arc in the median plane of its autozooecium (text-fig. 1).

Practically nothing is known with certainty concerning the function of these setae.
It has been suggested, notably by Busk (1854, pp. 100, 104, 106; 1859, p. 79), that in
certain lunulitiform species the setae might be ‘subservient to locomotion’. Alternative

TEXT-FIG. 1. a, Cupuladria canariensis', b, Discoporella umbel lata.

Vibracula with vibracular setae (after Norman 1909).

views are that they function as defensive or cleansing organs (Busk 1859, p. 79). In my
opinion, it is clear that they serve to stir up the water in the colony’s immediate vicinity,
not so much in order to fan food particles towards the polypides’ mouths as to prevent
clay particles settling on the colony.

If this view is correct, then the possession of these setae must be a valuable asset to
the lunulitiform Bryozoa, since it would make them some of the least sensitive to clay
sedimentation. It is certainly no coincidence that of all the possible zoarial growth-
forms it is precisely two lunulitiform genera, Cupuladria and Discoporella, that venture
most closely, on both sides, to the mouths of the Mississippi distributaries (text-fig. 2).
On this map the distribution pattern of the Bryozoa can clearly be divided into three
zones :

a. An inner zone between the shore and the 5 fathom line, in which swell and breakers
begin to disturb the sea bottom. Here, where the water is turbulent, no lunulitiform
Bryozoa occur (for reasons given on p. 187) though other types do, such as those that
attach themselves to plants or shells.

b. An outer zone between the 50 and 100 fathom lines, having a steep slope and an
irregular topography and situated on the outer edge of the continental shelf. Deposition
of clay along the outer margin of many continental shelves is notoriously slight to non-
existent (Kuenen 1939; Shepard 1948, p. 160), and the Gulf of Mexico is no exception
(Phleger 1959, p. 650; 1960, p. 288). The low rate of deposition and the local presence
of hard substrata explain why Bryozoa with other growth-forms have been able to
settle in this zone.

174

PALAEONTOLOGY, VOLUME 6

c. A middle zone, which coincides with the broad plateau lying between the 5 and 50
fathom lines. A large part of the clay brought down to the sea by the Mississippi is
deposited on this plateau, and it is in this area that only lunulitiform species of Bryozoa,
being equipped with vibracular setae, are able to survive.

number of Bryozoa
per 30 gramme sample

Bryozoa present, but in
unknown quantity

1-3

4-10

11-30

31-100

>100

□

barren sample

lunulitiform Bryozoa

as percentage of total ^

assemblage

lunulitiform Bryozoa
present, but percentage "
unknown

TEXT-FIG. 2. Predominance of lunulitiform Bryozoa in the bottom sediments of the Mississippi Delta
area. Sources: (a) collections from the Exploration and Production Research Laboratory, Shell
Development Company, Houston, Texas, now at ksepl, Rijswijk; (b) Parker 1956.

There are five characteristics which in combination are responsible for the ability of
this type of bryozoan to occupy an exceptionally wide range of environments :

(i) The ability to tolerate a certain amount of clay sedimentation owing to the posses-
sion of vibracular setae.

(ii) The ability to exist on almost any kind of bottom as long as the latter consists of
small particles.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

175

(iii) The ability to withstand a wide range of temperatures (eurythermal).

(iv) The ability to withstand moderate salinity variations (euryhaline).

(v) An insensitivity to hydrostatic pressure, light penetration, and other factors
directly concerned with depth.

TEXT-FIG. 3. Recent distribution of Ciipuladria canariensis, showing confinement between 14° C. surface
isocrymes (isocrymes after Sverdrup, Johnson and Fleming 1960 and Wust 1960).

This combination of characteristics is the reason for the wide area of distribution of
Cupuladria canariensis, which includes the tropical and subtropical Atlantic, the eastern
Pacific, and the Mediterranean (text-fig. 3), and which has been generally the same through-
out the Later Tertiary and Quaternary (text-fig. 4). Yet its distribution is not merely
wide on a global scale. Within fairly restricted areas such as the Gulf of Mexico (text-
figs. 5, 6) and the Nigerian shelf (text-fig. 7), or in the marine Pliocene of the Low Coun-
tries (text-fig. 8) and in the Miocene basin of eastern Venezuela (text-fig. 9), it is also
widely distributed on a provincial scale.

2. Larval stage

One may wonder whether there is not perhaps a sixth characteristic contributing to
this organism’s wide distribution : the duration of its larval stage. Like all Bryozoa, the
lunulitiforms are sessile, colonial organisms but they possess a free-swimming larval

176

PALAEONTOLOGY, VOLUME 6

Stage. The larvae are able to swim by means of their cilia, and the duration of this free-
swimming stage is conceivably one of the factors contributing to the geographical
distribution of a benthonic species (Cloud 1959, p. 951).

Unfortunately, the larval stage of C. canariensis is still unknown. Within the order
Cheilostomata, however, two completely different types of larvae occur. One of these,
the so-called Cyphonautes larva, which is characterized by the possession of a functional

alimentary canal, may spend a period of up to two months in this condition. The large
majority of the Cheilostomata, however, have the second type of larva, in which this
structure is rudimentary or entirely lacking. According to current views, such larvae
are therefore drastically limited in the duration of the free-swimming stage (to no more
than 12-24 hours, depending on the supply of yolk), so that they do not become truly
pelagic. Nevertheless, there are species of Cheilostomata that have this second type of
larva and which, in spite of this, have a very wide, or almost cosmopolitan distribution
(e.g. Microporella ciliata).

This is a baffling paradox with which every student of Bryozoa is sooner or later
confronted and for which various solutions have been proposed, e.g. continental drift,
dispersal via ancient archipelagos or shelf bridges, or trans-oceanic rafting on floating
objects by surface currents. Recently Cheetham (1960), in a stimulating paper, discussed

R. LAGAAIJ; THE BRYOZOAN CUPULADRIA CANARIENSIS

177

TEXT-FIG. 5. Quantitative distribution of Ciipiiladria canariensis in bottom sediments of the north-eastern Gulf of Mexico.

178

PALAEONTOLOGY, VOLUME 6

the merits of each of these three hypotheses in the light of Early Tertiary cheilostome
distribution. He clearly favours the third alternative, but did not fully consider the possi-
bility of long-distance dispersal in the larval stage. Yet it is precisely this fourth alterna-
tive that most strongly suggests itself in the case of those zoarial form-groups that are
most unlikely to become attached to seaweed or other ‘rafts’. If the distribution of such
species, including C. canariensis, is amphi- Atlantic, and if the Wegenerian hypothesis that
the surface and bottom configuration of the Miocene Atlantic Ocean were drastically

TEXT-FIG. 6. Quantitative distribution of Cupuladria canariensis in bottom sediments of the north-
western Gulf of Mexico.

different is rejected, the conclusion is inevitable that their free larval stage must under
certain conditions be, and have been, able to last a long time.

Harmer (1910, p. 520) suggested ‘that it does not follow that because we know that
a larva may, under favourable conditions, fix itself a few minutes after it becomes free,
we should be justified in assuming that that larva would not retain for a long period the
power of undergoing a normal metamorphosis should it be drifted away from suitable
fixing-grounds’. Silen’s (1944, pp. 30, 31) hypothetical concept of external food absorp-
tion in larvae which are devoid of an alimentary canal is also interesting in this connexion.

3. Substratum

After a brief or protracted period of wandering, the larva settles on a hard substratum.
It would seem to make a very careful choice in this matter, as if it knew in advance that
the substratum on which it settled would have to be raised above the sea floor and even-
tually become lodged in the apex of the conical structure which is the adult zoarium.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 179

TEXT-FIG. 7. Quantitative distribution of Cupitladria canariensis in bottom sediments of the Nigeria shelf (12° C. bottom isotherm from data filed

at KSEPL, Rijswijk).

180

PALAEONTOLOGY, VOLUME 6

ll^OUa+APPELSCHA y

#H0DeN

0OOST£f!HOOT zteLlWO*^ i

6ftEDA

♦seppE*

^ '’|#W(LI«(ARSD0NCK

># >Aan,twerp

PLIOCENE IN INNER TO MIDDLE NERITIC FACIES
PLIOCENE IN MIDDLE TO OUTER NERITIC FACIES

st.janS^

STEEN

TEXT-FIG. 8. Distribution of Cupuladria canariensis in the eastern part of the Pliocene North Sea basin
(distribution of marine facies after van Voorthuysen 1956).

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

181

Above all, Iherefore, this substratum must not be too big, and it is for this reason that
w'e usually find coarse quartz grains, glauconite pellets, small shell fragments, broken
echinoid spines, even moderately large Foraminiferida such as Amphistegina gibbosa and
Globorotalia menardii, or fragments of other lunulitiform Bryozoa functioning as nuclei
for Cupuladria. The ideal biotope for the lunulitiform Bryozoa is therefore a small-
particle (quartz and/or carbonate sand) bottom. They never occur on a bottom consisting

TEXT-FIG. 9. Distribution of Cupuladria canariensis in the Miocene basin of eastern Venezuela

(shaded area).

entirely of clay, and are never found encrusting rocks, large pebbles, piles, larger shells,
or other such large objects. It is not without reason that lunulitiform colonies are
commonly called ‘free’ (though this is incorrect) as opposed to ‘encrusting’ forms.
So unusual, in fact, is the appearance of an ‘encrusting’ C. canariensis that Silen (1942,
p. 14) refers to a juvenile colony on a small stone from Anguilla in the West Indies as
only ‘possibly belong[ing] to this species’.

The question arises whether the larva’s power of discrimination is real or apparent.
On the one hand one might suppose that the larva settles indiscriminately on all kinds
of substrata and that it develops into a colony only on those that are suitable. In that
case the selection is the work of external circumstances and not of inherent ‘intelligence’
in the larvae. On the other hand, selectivity on the part of pelagic larvae has definitely
been observed in other groups of marine invertebrates. According to Thorson (1955, p.
390), Wilson (1952 and earlier papers) has shown ‘that the larvae of several polychaetes.

182

PALAEONTOLOGY, VOLUME 6

when ready to metamorphose, will critically examine the bottom substratum to which
they are exposed. If they find it attractive, they settle. If they find it less attracth e or
directly repellent, they will continue their pelagic life for days or even weeks. During
such a prolonged larval life these larvae test the substratum at intervals as they are
transported by the current directly over the bottom.’

As far as Cupuladria is concerned, the following example is pertinent. Plate 25,
figs. 2a, b, show two specimens of Cupuladria (not C. canariensis, but an allied species,
having affinity with C. pyrifornds Busk) from the subsurface Oficina formation (Mio-
cene) of eastern Venezuela. The larvae of both specimens, and of several others from the
same shale samples, originally singled out, and settled upon, specimens of Planorbidinella
trinitatensis (Nuttall) and persistently neglected several other species of smaller
Foraminiferida. This choice seems to have been prompted by the absence of quartz
sand; for in a sandy facies with both quartz grains and Planorbidinella available in
good quantities, such as is found along the southern boundary of the eastern Venezuela
basin, settling invariably occurred on the quartz grains only.

4. Astogeny

Having attached itself to a suitable substratum, the larva then rapidly undergoes its
metamorphosis into the first individual, the ‘ancestrula’, of the future colony. Further
development takes place by a process of budding. It is interesting to follow the juvenile
colony through the early stages of this development, especially since Harmer in 1931
(p. 162) was still able to say: ‘there is no conclusive evidence with regard to the earliest
stages in the discoidal or conical colonies, and a mere count of the number of surround-
ing zooecia is not enough to settle the matter’, and no pertinent observations have been
made since then.

By the time the number of zooecia has increased to twenty-six, the zoarium has passed
through several separate stages of growth (text-fig. 10):

(a) The single ancestrula. Despite prolonged searching I have never observed a single
ancestrula. Thus it would seem that the process of budding sets in very soon after the
metamorphosis is complete.

(b) The three-cell zoarium. This, the earliest zoarial growth stage observed, invariably
consists of three zooecia forming the pattern shown in text-fig. lOZ?. I have recorded
several dozens of these three-cell colonies. They display distinct bilateral symmetry.
From their mutual relationships it may be inferred that the ancestrula has given rise to
two proximo-lateral first-generation zooecia.

(c) The four-cell zoarium. A fourth zooecium is added proximally in the plane of sym-
metry (text-fig. 10c). The circle is closed and the colony is ready to start its radial growth.

{d) The six-cell zoarium. By the addition of two zooecia in the angles between the
ancestrula and the two first- generation zooecia, the colony assumes the shape of a six-
pointed star (text-fig. lOrZ). The plane of symmetry through the ancestrula is still clearly
apparent.

(e) The eight-cell zoarium. Two new proximo-lateral zooecia appear (text-fig. 10c).

(/) The ten-cell zoarium. Two more zooecia are added distally, one on either side of the
ancestrula (text-fig. 10/; PI. 25, fig. 3).

(g) The twelve-cell zoarium. Two new lateral zooecia appear (text-fig. lOg). At this
stage the colony is still markedly stelliform; the bilateral symmetry can be seen without

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

183

TEXT-FIG. 10. Early stages of zoarial growth in Cupidadria canariensis (based on a series of Recent

specimens from Nigeria).

difficulty. No zooecia are yet in proximal contact with the vibracula of preceding cells,
but this situation changes in the next stage. This earliest, central, part of the colony
is therefore relatively less densely covered by vibracular setae than later, peripheral,
additions. It is therefore probably no coincidence that preferably (although not ex-
clusively) these central zooecia sometimes regenerate into large vibracula of the type first
described by Hastings (1930, p. 714; see also Silen 1942, text-fig. 8), whereas in other
lunulitiform species they tend to develop a calcareous closure.

(h) The eighteen-cell zoarium. Six new zooecia have appeared at the periphery (text-
fig. lO/i), probably via two intermediate stages. The first shght departure from the
original plan of symmetry is now apparent, since the proximal and the distal indentations

184

PALAEONTOLOGY, VOLUME 6

at the periphery in the median line are too large to be filled by a single zooecium and yet
too small to accommodate a pair of zooecia. The filling of the gap therefore takes place
asymmetrically.

(z) The twenty-six-cell zoarium. A further departure from bilateral symmetry takes place
(text-fig. lOr). By now the colony has attained a diameter of T9 mm. and is well on its
way to adulthood.

Two points emerge from these observations:

i. Waters’s (1926, p. 426 and text-fig.) concept of a double ancestrula in Cupuladria,
the two being turned in opposite directions, and each giving off" three distal
zooecia, is untenable. Although Waters specifically mentioned C. canariensis in
connexion with his observations, it is clear from the occurrence of partially closed
zooecia and from the provenance of his material (Princess Charlotte Bay, Queens-
land, Australia), that he was actually dealing with C. guineensis (Busk).

ii. Silen’s (1942) theory of spiral growth is no longer valid in the case of the genus
Cupuladria.

I should like to emphasize that the astogeny outlined above only applies to ideal
cases, and that deviations from this scheme are common and may appear at an early
stage (PI. 25, fig. 4). Usually such deviations are closely bound up with irregular con-
figurations of the small-particle substratum, or with an excentric position of the ances-
trula on the substratum. The eight-cell zoarium figured in Plate 25, fig. 4 deviates in that
it has developed a zooecium (on the left) in a position that would normally not be
occupied until the twelve-cell stage, while the usual place for the eighth zooecium (on
the right-hand side) remains vacant. Occupation of the latter position, which projects
beyond the edge of the particle, would have involved building a stronger dorsal wall
than the extremely thin one required in the position now preferred, where it is supported
by the substratum. Obviously less building energy is required for growth on the sub-
stratum than for expansion beyond its edges.

5. Mode of life of lunulitiform colonies

The mode of life of the adult colony and, closely connected with that, its orientation
with respect to the sea bottom, are controversial matters, and widely differing suggestions

EXPLANATION OF PLATE 25

Fig. 1. Cupuladria canariensis (Busk). Adult colony, a. View of the convex surface; b. View of the con-
cave surface. G.S. 173, Pliocene of Proefboring 41, Reek, Netherlands, 15-00-20 00 m. x 10. (After
Lagaaij 1952.)

Fig. 2. Cupuladria sp. Concave side of juvenile colonies, showing larval predilection for the foraminiferid
Planorbulinella trinitatensis (Nuttall). Lower Miocene (Oficina formation) of Texas Petroleum Com-
pany well Mata-1, Estado Anzoategui, Venezuela; a, 8,160-8,170 ft.; b, 8,300-8,320 ft. X20.

Fig. 3. Cupuladria canariensis (Busk). Juvenile colony (ten-cell stage) in normal symmetrical develop-
ment. Recent, Mees Cremer 1959 Sta. 98, Nigeria, 14 fms. x20.

Fig. 4. Cupuladria canariensis (Busk). Juvenile colony (eight-cell stage), showing early departure from
bilateral symmetry. Recent, Mees Cremer 1959 Sta. 376, Nigeria, 11 fms. x20.

Fig. 5. Cupuladria canariensis (Busk). Regenerated colony, showing sector of original large flattened
conical colony with peripheral outgrowth in radial direction. Recent, Ras-el-Amouch, Mediterranean,
45 fms. X 10. (After Dartevelle 1935.)

Palaeontology, VoL 6

PLATE 25

LAGAAIJ, Cupidadria canariensis (Busk)

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 185

have been made, none of which, according to Harmer (1931), have been supported by
pertinent observations. Four tenable hypotheses have been put forward (text-fig. 11).

As Harmer (1931, pp. 150-1) has shown, the conventional view that the colony rests
on its flat ‘base’ (text-fig. 1 \a) was first disputed by Maplestone (1910, p. 3), who expressed
the opinion that ‘the conical forms in their living state have the base uppermost’.
Although at this time Maplestone was only referring to the conical colonies of the genus

RESTING ON BOTTOM WITH
a GROWING EDGE

ATTACHED BY ROOTLETS
SPRINGING FROM CIRCUM-
ANCESTRULAR "RADICULAR"
200ECIA

ALGAL FROND

FIXED UNDER THE FRONDS J ROTATION AND ASCENSION
C OF algae fl TO ESCAPE BURIAL

TEXT-FIG. 1 1 . Various hypotheses regarding the mode of
life of lunulitiform colonies.

Conescharellina, later authors have extended his contention to include other genera
whose colonies are conical. Moreover, Maplestone was not certain whether the conical
colonies (of Conescharellina) hang point downwards from some foreign object, or whether
they retain this position on the bottom by means of anchoring filaments.

Canu (1915, p. 21) adopted and elaborated on both hypotheses. His early reasoning
is not generally known (cf. Harmer 1931, p. 151), but later re-statements of his views
have profoundly influenced later workers. It is therefore necessary to quote Canu in full
on this point: ‘Les especes flottantes comme les Limiilites ont le zoarium conique, la
pointe en bas. Tantot il est maintenu sous les Algues [text-fig. 11c], tantot il est attache
a de petits objects par des radicelles [text-fig. 11/?]. Celles-ci proviennent de Zoecies
radiculaires (= Zoecies avortees de d’Orbigny) disposees autour de I’ancestrule. . . .
Les zoecies radiculaires sont d’abord des zoecies hydrostatiques. La larve, en effet, se
fixe sur un grain de sable ; I’ancestrule qui se developpe emet immediatement des zoecies
radiculaires qui I’enveloppent et permettent a I’animal de commencer son ascension
sous une Algue loin du sable dangereux a son developpement. . . . Soit attaches par des
radicelles, soit retenus sous les Algues, les Limiilites, par leur forme turbinee, sont de
position tres instable: ils chavirent au moindre filet d’eau. L’animal maintient sa position

186

PALAEONTOLOGY, VOLUME 6

normale a I’aide de longs filaments articules appeles vibraculaires. Ce sont done des
appareils de stabilisation iin pen analogues au balancier des danseurs de corde.’

The same views, essentially unchanged, are repeated by Faura and Canu (1916),
Canu and Bassler (1920, pp. 238 ff.), and Canu and Lecointre (1927, p. 35). Waters
(1921) was quick to oppose these views, which have in fact largely been confined to the
French school. I refer in particular to his statement (1921, p. 401): ‘. . . though sometimes
the growth is on a much larger stone, as in some specimens of Cupuladria canariensis
from Petit Tahou, Liberia. It would seem impossible for a colony so heavily weighted
to float, nor can we think it could float in a reversed position.’ In 1926, however, Waters
(1926, p. 425) wrote more cautiously: ‘What we have called the upper surface is, in the
ancestrular and early stages, at the top, even though there may be a subsequent reversal ’

Finally, Canu and Bassler, realizing how difficult it was to see how a conical colony
with its apex downward could ‘maintain its equilibrium even in the water, in a position
absolutely contrary to the ordinary laws of statics’, invoked rotation as a means of
conserving its position (text-fig. \\d), first for Conescbarellina (1929, p. 482) and later
for Limidites (1931, p. 9): ‘Ils vivent done I’apex en bas au voisinage du fond sableux.
Ils s’en degagent par rotation et ascension pour eviter I’enlissement’, and (1931, p. 19)
‘Les Limidites sont de petites coupes en perpetuelle rotation pour se degager du sable
et changer de place’.

Dartevelle also attributed a planktonic mode of life to Limidites (1933, p. 69) and to
Cupuladria (1943, p. 108): ‘Leur mode de vie est semblable a celui, bien connu, des
Limidites, e’est-a-dire que la colonie flotte entre deux eaux, les zoecies tournees vers le
dessous, la face concave vers le haut. . . .’ The designation ‘face superieure’ for the con-
cave side and ‘face inferieure’ for the convex, celluliferous side of lunulitiform colonies
still persists in recent French literature (Vigneaux 1949; Buge 1957).

Dartevelle (1933, p. 57), moreover, provided a novel explanation of the role played
by the small foreign particle at the downward directed apex of the cone : ‘La presence de
ce substratum constitue par un morceau de coquille, une nummulite, un grain de sable,
contribue a maintenir I’equilibre de la colonie et a I’empecher d’etre chaviree par les
vagues’ ; it served, in other words, as ballast, and as such would profoundly influence the
shape of the colony. According to Dartevelle (loc. cit., p. 70) the lighter the substratum,
the flatter the zoarial cone, and, conversely, the heavier the particle, the higher and more
dome-shaped the colony would have to become in order to keep the ballast as low as
possible: ‘. . . ce qui gouverne done la colonie, e’est la souci d’eviter le renversement et
de maintenir le meilleur equilibre possible au sein du fluide.’

Harmer’s presidential address to the Linnean Society of London in 1931 made it
abundantly clear that Canu and Bassler’s reasoning was based entirely on inference and
not on direct observation. He added (1931, p. 151): ‘I have failed to find any evidence that
is really conclusive with regard to the question at issue. Except for Whitelegge’s very
brief account, I am acquainted with no observations made on living specimens, and in
my judgment the matter should for the present be regarded as undecided.’ Harmer’s
statement is still as valid today as it was thirty years ago. Here is clearly a case where
laboratory experiment could be of value. It should not be too difficult to collect some
living specimens of Cupuladria and keep them under observation in a sea-water
aquarium.

Another example of reasoning by inference, the emphasis in which is laid on the

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 187

orientation of the ancestrula and of later zooecia with respect to the small-particle
substratum, occurs in Silen (1947, pp. 5-6, 8, 15 and text-fig. 8). He concluded (correctly,
in my opinion) that the colonies of Cupidadria rest freely on the sea bottom with the
apex pointing upwards. Silen thereby reverted to the conventional view (see, however,
Silen 1942, p. 13).

Two further arguments, both arising from the ecology of C. caiiariensis, support this
conventional view, or rather conflict with the hypothesis of a planktonic mode of life
postulated by Canu and his school. In the first place, if C. canariemis were indeed a
planktonic organism, one would expect to find its skeletons distributed over different
kinds of bottom and a very wide range of depths. However, its remains are found only
on the small-particle bottoms to which it is confined in life by the special requirements of
the larvae. Secondly, there is a direct relationship between the maximum depths at
which C. canariemis has been observed in the various marine areas, and the temperature
of the bottom water in those areas (see p. 189). Both these observations suggest that
C. canariemis is a truly benthonic organism.

6. Minimum depth

Cupidadria canariensis occurs mainly on small-particle bottoms and is hence a full
member of that rather select level-bottom community described by Petersen (cf. Thorson
1955). A type of small-particle bottom from which it is excluded, however, is one where
the sand grains have a tendency to shift under the influence of water movements. The
minimum depth at which the larvae of C. canariensis and other lunulitiform species,
such as DiscoporeUa umbellata (Defrance), can settle and develop into adult colonies
therefore depends on the intensity of the water turbulence over the bottom, i.e. on the
depth of the wave base and on the strength of the bottom currents. Since both these
factors are determined by the degree of exposure of the shelf and by oceanographical
factors, the minimum depth mentioned above varies from place to place (text-fig. 12).

Cupidadria colonies are therefore unlikely to be washed up on the shore, and there are
no records that this has ever occurred, although ‘one dead and worn’ colony of D.
umbellata has been found on the shore at Balboa, Panama Canal Zone (Hastings 1930,
p. 718). As Stach (1936, p. 63) has put it, ‘their free mode of life prohibits their existence
in the littoral zone where wave action is strongly felt’. The lunulitiform Bryozoa thus
seem to be confined to the stable small- particle bottoms below wave base. This conclusion
is quite contrary to that of Dartevelle (1933, 1935), who inferred an agitated, current-
infested biotope from the common occurrence of broken and regenerated lunulitiform
colonies in the Eocene of Belgium. Yet there is no reason why fragmentation should be
due solely to mechanical breakage in a highly turbulent environment; the destructive
activity of other marine organisms in deeper, quieter water might just as easily be respon-
sible (Ginsburg 1957, p. 83). It is known that holothurians (sea cucumbers) include the
lunulitiform Bryozoa in their diet. Silen (1942, p. 13) records eight colonies of C. cana-
riensis taken from the stomach of the sea-urchin Meoma ventricosa, and I have observed
the occurrence of fragments and of several entire colonies of C. canariensis among the
coarser debris in the stomach of a holothurian in the north-western Gulf of Mexico
(Cavalier 1956 Station 227, at a depth of 37-5 fathoms). Dartevelle (1935) gives Recent
examples of regenerated zoaria of C. canariensis (PI. 25, fig. 5) taken from the Medi-
terranean locality Ras-el-Amouch at a depth of 45 fathoms, which is well below that at

188

PALAEONTOLOGY, VOLUME 6

NEW

ORLEANS

TAMPA

PORT ARTHUR

PORT

HARCOURT

GULF OF MEXICO

GULF OF GUINEA

GALVESTON

GULF OF MEX CO

TEXT-HG 12. Minimum depths at which Cupiiladria canariensis has been found in various areas.
Notice shallowest occurrences in sheltered, deepest occurrences in exposed biotopes.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

189

which sand transport could occur, let alone transport and breakage of far larger objects
such as bryozoan colonies.

7. Maximum depth {minimum temperature)

The maximum depth of occurrence of Cupid adria canariensis is determined by the
temperature of the bottom water. The maximum depths at which C. canariensis has been
found in three marine provinces is as follows:

NW Gulf of Mexico

Fathoms

138

NE Gulf of Mexico

117

Straits of Florida

122

Jamaica

150

E of Jacksonville, Florida

440

N of St. Thomas, Virgin Islands

300^70

Senegal

118

Nigeria

120

Gulf of Mexico

I W and S confines of Sargasso Sea
I Equatorial West Africa

Since the recent geographical distribution of C. canariensis is roughly limited both in the
Atlantic and in the eastern Pacific by the 14° C. surface isocrymes (text-fig. 3) (and it
must be remembered that the bottom water at shelf depths will be a few degrees colder)
it may reasonably be assumed that this approaches the minimum temperature at which
C. canariensis can survive. The maximum depths of occurrence and the corresponding
bottom-water temperatures of approximately 12° C. observed in various marine areas
seem to confirm this assumption (text-fig. 13).

The deep occurrences along the western and southern confines of the Sargasso Sea
are of particular interest. In all three cases the data refer to living specimens. The
hydrography of this region (text-fig. 14) is almost unique in featuring a lenticular body
of water of uniform temperature (18° C.) and salinity (approx. 36-5%o) down to a depth
of 300-400 metres (Worthington 1959). Below this depth a gradual decrease of tem-
perature takes place down to the main thermocline. Clearly it is only the peculiar
temperature conditions prevailing in this area that permit the occurrence of C. canariensis
at such unusually great depths.

8. Maximum temperature

It is at the shallow end of its depth range that one has to seek the maximum temperature
which the species will tolerate. This will be in shallow coastal waters, where the effects
of atmospheric heat exchange are most strongly felt.

It will be readily understood, however, that the determination of this parameter will
be far less precise than that of the minimum temperature, since the absolute maxima
vary from year to year. Thus, C. canariensis may conceivably form part of the bottom
fauna of a shallow bay during the normal summer of a particular year, but the following
year an abnormally hot summer may bring its occupation of that bay to a sudden
termination. The best approximation will therefore be found by taking the average
maximum temperature in the warmest month, recorded over a number of years.

Areas where such shallow occurrences of C. canariensis coincide with the necessary
amount of regularly recorded temperature data are Tampa Bay, Florida, and Breton

190 PALAEONTOLOGY, VOLUME 6

Sound, Louisiana, where C. canariensis occurs at minimum depths of 4 and 2 fathoms
respectively (text-fig. 12).

According to the records published by the U.S. Coast and Geodetic Survey (1955),
the average maximum temperature in the warmest month (July) for the years 1947-54
at St. Petersburg (Tampa Bay) was 31° C. Since this figure is based on surface measure-
ments, the corresponding value for the shallow bottom water is bound to be somewhat
less extreme.

N.W. GULF OF MEXICO N.E. GULF OF MEXICO SARGASSO SEA

GULF OF GUINEA

TEXT-FIG. 13. Relation between deepest occurrences of Cupuladria canariensis and bottom-water

temperature.

In Breton Sound, where C. canariensis is known to occur in 2-3 fathoms depth in the
lee of Breton Island (Parker 1956), surface-water temperatures during the summer reach
an average absolute maximum of about 31-32° C. (Scruton 1956, p. 2937). According to
Scruton (loc. cit.), ‘vertical stratification is stronger in the summer than at other times,
so that bottom temperatures in Breton Sound proper probably seldom reach 32°’.

9. Maximum salinity

The clue to the maximum salinity that C. canariensis will tolerate lies in the Medi-
terranean. From its occurrences elsewhere, on the Florida shelf and in the West Indies,
one may infer that it still thrives in salinities of approximately 36-5%o. In the Medi-
terranean, on the other hand, salinities at the surface are everywhere higher than 37%o
(and increase with depth), except where the surface current carries water of Atlantic
origin and an original salinity of about 36-25%o through the Strait of Gibraltar along the

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

191

temperature in ® C

10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

deepest known occurrences of CUPULADRIA CANARIENSIS

TEXT-RG. 14. Deepest known occurrences of Cupuladria canariensis are found where deeply descending
warm water mass of Sargasso Sea intersects with sea bottom. Shading in ocean area indicates the occur-
rence of water of 18° C. at 300 m. depth (map and graphs after Worthington 1959).

north coast of Africa as far east as Tunisia (Sverdrup, Johnson and Fleming 1960,
pp. 643, 646).

The extent of this Atlantic water in the Mediterranean coincides exactly with the
Recent Mediterranean distribution of C. canariensis (text-fig. \5d). The easternmost
Mediterranean record of the species is from Cape Rosa, Algeria, 40 km. east of Bone.
C. canariensis, like all other lunulitiform species, is conspicuously absent among the
bryozoan fauna of the eastern Mediterranean, for example in Tunisia (Canu and
Bassler 1930), Egypt (O’Donoghue and De Watteville 1939), and Syria (Gautier 1957).
Since this entire area lies south of the 14° C. surfaee isocryme (text-fig. 15d) and since

192

PALAEONTOLOGY, VOLUME 6

bottom temperatures in the Mediterranean, even at great depths, nowhere drop below
about 13° C. (Nielsen 1912; Furnestin 1960), it is clear that here the limiting factor is
not the water temperature but the >37%o salinity.

This conclusion has the interesting implication that the Miocene, Pliocene, and early
Pleistocene Mediterranean (text-fig. 15n-c) was somewhat less saline than it is at present.
The difference was most marked in the Sicilian, when C. canadensis (and by inference
salinities <37%o) extended into the Levantine basin (Rhodes), where present-day
salinities constantly remain about 39%o (Wiist 1960, figs. 2, 5; pi. 7).

10. Minimum salinity

Data on the minimum salinity which C. canadensis will tolerate must be inferred from
its occurrence in the shallow coastal waters, where precipitation and run-off are most
effective in lowering the salinity.

Reduced salinities have been observed in the following shallow areas where C.
canadensis is known to occur;

a. Breton Sound, leeward of Breton Island, Louisiana; depth 2-3 fathoms (Parker
1956); bottom salinity 28-5%o, measured in the autumn of 1951 during flood tide; the
ebb tide produces still lower salinities (Scruton 1956, p. 2927).

b. South of Calcasieu Pass, Louisiana; depth 5-5 fathoms (text-fig. 12). Salinities
measured at neighbouring stations (Bandy 1954, fig. 8, Sta. 106, Sta. 108) are of the order
of 28-28- 5 %Q. To all appearances these figures are based on surface measurements, but
in this shallow turbulent part of the Gulf of Mexico vertical stratification is bound to
be slight and the corresponding bottom salinities will therefore probably be not very
different.

C. canariensis is absent in the Gulf of Mexico off Grand Isle, Louisiana, presumably
because here salinities at 10 feet below the surface may drop periodically to as low as
21 %o in June and July, and 14%o in February (Geyer 1950, p. 103). The lowest monthly
average in this area is 22-6%o (March 1949).

11. Recognition

Another, more subjective, factor which affects the boundaries of the known distri-
bution of C. canadensis is the ease with which the species can be recognized. Large
undamaged colonies may easily attain a diameter of 1 -5 cm., and their aesthetically satis-
fying shape makes them conspicuous among the other members of the macro-fauna.
Unfortunately, C. canadensis is rather fragile and during rough handling in nature o** in

EXPLANATION OF PLATE 26

Fragments of fossil Cupuladria canariensis (Busk) from various localities.

Fig. 1. Detail of the broken surface, showing the characteristic fine, parallel, vertical striation. Lower
Miocene (Carapita formation) of Mene Grande Oil Company well SB-133, Estado Monagas,
Venezuela, 3,825-3,840 ft. X40.

Fig. 2. Middle Miocene (Reinbek/Dingdener Stufe) of Twistringen, SSW of Bremen, Germany, x 20.
Fig. 3. Lower Miocene of Shell-BP Petroleum Development Company well Ituk-1, Nigeria, 3,320-
3,780 ft. x20.

Fig. 4. Lower Miocene (Chickasawhay formation) of Limestone Creek, Wayne County, Mississippi,
U.S.A. X20.

Fig. 5. Upper Miocene (Cubagua formation) of Socony Mobil Oil Company well Cubagua-1 , Cubagua,
Venezuela, 218 ft. x20.

Palaeontology, Vol. 6

PLATE 26

LAGAAIJ, Cupuladria canariensis (Busk)

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

193

Fresh-water lokes and marshes

\////\ Brackish-wafor areas

tzz]

Marine

TEXT-FIG, 15. Fossil and recent distribution of Ciipiiladria canariensis in Europe and North Africa
(Middle Miocene and Pliocene base maps after Wills 1951).

C 1015

o

194

PALAEONTOLOGY, VOLUME 6

the laboratory is apt to break, almost always along the radial lines on the concave
surface. The smallest recognizable fragments take the form of tiny rectangular prisms
(PI. 26, fig. 1), whose dorsal ends correspond to the rectangular compartments visible on
the concave surface (text-fig. 16). These compartments are perforated, and so, con-
sequently, are the dorsal ends of the individual prisms. The vertical faces are flat and show
fine, parallel, vertical striation (PI. 26, fig. 1). Thus even very small fragments can be
recognized (PI. 26, figs. 2-5), and for this reason C. canariensis may be regarded as a
component of both the macro- and the microfauna.

APPLICATIONS
1. Shifts in sea-water temperature since the Miocene

Palaeobotany holds the key to the interpretation of Tertiary climates in north-west
Europe. It is known that the north-west European climate gradually cooled from tropical
(Eocene), through subtropical (Miocene and early Pliocene) to warm temperate (late
Pliocene) conditions, after which even more marked and rapid cooling to subarctic
conditions introduced the Pleistocene.

At present both summer and winter air temperatures in north-west Europe are strongly
influenced by the temperature of the water in the eastern Atlantic and the North Sea.
It is not unreasonable to assume, therefore, that the subtropical and oceanic climates
prevailing in north-west Europe during the Miocene and Pliocene periods bore a direct
relationship to considerably higher sea-water temperatures in these latitudes.

Such an hypothesis is strongly supported by a comparison of the present-day occurrences
of C. canariensis with those during Miocene and Pliocene times (text-fig. 17). As has
been already mentioned (p. 189), the northern boundary of its present area of distri-
bution in the northern hemisphere coincides
with the 14° C. surface isocryme (text-fig. 3).
Consequently, Miocene and Pliocene winter
surface-water temperatures in the North Sea
basin must have been at least 14° C. February
surface-water temperatures in this area now
average between 5° and 6° C. {fide Sverdrup,
Johnson, and Fleming 1960, chart II). In the
Miocene and even in the Pliocene the water
in the southern part of the North Sea must
therefore have been at least 8° or 9° C. warmer
than it is at present. This conclusion accords
well with the figures derived by the Polish
palaeobotanist Szafer for the amount by
which Pliocene air temperatures in Europe
north of the Alps exceeded those now pre-
vailing: January +11°, July +9°, yearly average 4-9° C. {fide Godwin 1956, p. 296).

This conclusion is particularly interesting, however, because both in the Middle
Miocene (text-fig. 1 5a) and in the Pliocene (text-fig. 1 5b) the land area at present occupied
by the British Isles was connected to the continent of Europe. C. canariensis (and the
entire warm-water fauna associated with it) must therefore have migrated to the North
Sea area of those times by a path lying to the north of Scotland. In view of the present-

TEXT-FiG. 16. Ciipitladria canariensis. Detail of
the concave surface (after Busk 1859a).

R. LAGAAIJ; THE BRYOZOAN CUPULADRIA CANARIENSIS

195

□ Land (in Pliocene)

□ Fresh and brackish water areas
(in Pliocene)

RECENT

B14°C surface isocryme
(after Wust, I960)

RECENT occurrences of
" Cupuladria conariensis

PLIOCENE occurrences of
^ Cupuladria canariensis

INFERRED PLIOCENE
14® and I5°C surface isocrymes,
patterned on the position
of the present 5° and 6® isocrymes
(after Sverdrup, Johnson ft
Fleming, I960)

TEXT-FIG. 17. Shift of sea water temperatures in the Eastern Atlantic and Mediterranean since the
Pliocene (Pliocene base map after Wills 1951).

196

PALAEONTOLOGY, VOLUME 6

day confinement of the species by the 14° C. surface isocryme, this isotherm must have
lain to the north of Scotland in both the Miocene and Pliocene periods and probably
resembled in shape the present-day 5° C. isocryme, which under the influence of the
Gulf Stream curves around Scotland and penetrates into the North Sea (text-fig. 17). The
stranding of a loggerhead turtle, Caretta caretta (Linn.), on the Norwegian coast in
December 1951 (Willgohs 1953) and the capture of a flying fish, Cypsilunis hetenims, in
Oslo Fiord in 1848 and in 1937 (Bruun 1938) are indications that even today the Gulf
Stream occasionally carries tropical and subtropical marine organisms along this ancient
migration route.

During the Pleistocene C. canariemis did not have a chance to migrate once more into
the North Sea region (text-fig. 15c). This sea- water temperature did not rise sufficiently
to allow this, even in the interglacial periods, when the climate was somewhat warmer
than it is now. From the composition of the molluscan fauna of the Dutch Eemian (the
last, late-Pleistocene, interglacial), van Straaten (1956, pp. 224, 225) concluded that the
temperature of the sea-water at that time could only have been about 4° C. higher than
that of the present North Sea, and might have been no more than 2° or 3° C. higher. It is
interesting to note that Spaink (1958, p. 31) concluded that the southern element of this
Eemian fauna must also have reached the Dutch coastal area by the route north of
Scotland.

2. The Oligocem-Miocene boundary in the Gulf Coast and Caribbean areas

During the past decade, several attempts have been made to correlate the Tertiary
formations on either side of the Atlantic. The problem of defining the Oligocene-Miocene
boundary in the Caribbean area has been closely connected with these attempts and has
still not definitely been solved.

A detailed discussion of this problem lies outside the scope of this paper. Those who
are interested are referred to the paper by Fames (1953), through whose activity the
problem became critical, to the ensuing discussion between Stainforth (1954) and Fames
(1954), to the later restatement of the problem by Stainforth (1960u; 1960Z?, with
extensive bibliography) and to the renewed discussion between Fames et al. (1960u;
19606) and Stainforth (1960c). The arguments put forward by Fames et al. have
now been stated in full detail in their book on Mid-Tertiary stratigraphical correlation
(1962).

Briefly, the Oligocene-Miocene boundary in the Caribbean area has undergone a pro-
gressive lowering during the past decade. All concerned agree that this move was justi-
fied; there is no agreement, however, about the level in the Caribbean sequence of
planktonic foraminiferal zones where this boundary should finally be drawn. Stainforth
( 1 9606, p. 226) is undecided ‘whether the whole Globigerina ciperoensis zone and part of
the Globigerina dissiniilis zone or only part of the Globigerina ciperoensis zone represents
the whole Oligocene’. Fames (1955, p. 86), on the other hand, implied complete absence
of marine Oligocene sediments in Trinidad by his statement that ‘probably all the Cipero
formation is of Lower Miocene age’. Fames et al. (1960u, 19606, 1962) have since cor-
robated this view and have even extended it so that not only the Cipero but also the
underlying San Fernando formation is included. If their opinion is correct, a major
hiatus occurs in the Trinidad sequence between the Eocene (Navet and equivalents) and
the Miocene (San Fernando, Cipero and equivalents) marine deposits.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 197

It is important to discover whether, and to what extent, the lunulitiform Bryozoa, and
in particular Cupu/adria canariensis, throw light on the matter.

Lunulitiform Bryozoa, which range from the Upper Cretaceous to Recent, have long
been of interest to palaeontologists. Lumilites, the eponymous genus, was introduced
by Lamarck in 1816 and since then a considerable number of fossil species have been
recorded from Tertiary and Quaternary strata both in Europe and in North America.
The accompanying range charts (text-fig. 18) show the time-stratigraphic distribution of
these species in the two hemispheres.

Comparison of these two charts reveals several interesting facts :

a. There are no Eocene and Oligocene species of Limulites common to both hemi-
spheres. It is possible that a thorough systematic revision of the group might alter this
situation. But whether it would or not, it is significant that there is a similar lack of
common species among the contemporaneous larger Eoraminiferida of both hemi-
spheres.

b. The disappearance of the genus Lumdites clearly did not take place contem-
poraneously in both hemispheres. In North America it probably no longer occurs in
situ in the post-Vicksburg formations, whereas in Europe it persists into the Pliocene. It
should be borne in mind, however, that several living species of Lumdites have been
recorded from Australian seas.

c. Several names common to both charts, of which one is C. canariensis, first appear
in the Miocene of Europe and in the post-Vicksburg formations of the southern U.S.A.
This new appearance of modern lunulitiform genera and species, which, as has been seen
in the case of C. canariensis, possess exceptional environmental tolerance leading to
wide and rapid dispersal, may well serve as a criterion for establishing the Oligocene-
Miocene boundary in sequences of ancient tropical and subtropical shelf sediments on
both sides of the Atlantic.

By this criterion the Vicksburg group correlates with some part of the European Oligo-
cene (absolute hegemony of Lumdites), whereas such post-Vicksburg formations as the
Chickasawhay of Mississippi and Alabama and the subsurface Upper Frio of Texas
cannot be considered older than Aquitanian (since both contain C. canariensis).
Although most North American stratigraphers, e.g. Cooke et al. (1943), MacNeil
(1944), and oil companies do not share this view, it is clear that it was held as long ago
as 1934 by others, such as Howe (1934) and McGuirt (1941).

Some of the evidence underlying text-fig. I8b is brought out in greater detail on the
correlation chart of the Gulf Coast Oligocene and Miocene formations (text-fig. 19),
which shows the known occurrences of the genus Lumdites and C. canariensis according
to the published records and my own observations. Conspicuous on this chart is the
wide distribution of these lunulitiform Bryozoa throughout the marine Tertiary sedi-
ments, even though information on several southern states is still incomplete.

It is interesting to study the Caribbean area in the light of what is known about Europe
and the Gulf Coast, where modern lunulitiform species and genera first appear in the
Aquitanian and post-Vicksburg formations respectively.

It has already been shown (text-fig. 9) that lunulitiform Bryozoa, in this case C.
canariensis, are widely distributed throughout the ‘Oligocene-Miocene’ basin of eastern
Venezuela. The vertical distribution of C. canariensis in various parts of the basin is

198

a) in Europe

HOLOCENE a RECENT

(£. ATLANTIC a hlEDITERRANEAN)

I

1

1 — n*

PLEISTOCENE

1

1

1 1

PLIOCENE

[TfT~T

1

1

y 1

MIOCENE

PONTIAN

SARMATIAN

TORTONIAN

I

1

HELVETIAN

1

si

1

1

■

BURDIGALIAN

t

II

1

1

1

AQUITANIAN

tml

ill

1

1

OLIGOCENE

CHATTIAN

TTTl

RUPELIAN

LATTORFIAN

liiniurr.

EOCENE

BARTONIAN

TT

LEDIAN

LUTETIAN

YPRESIAN

PALAEONTOLOGY, VOLUME 6

! £C CD

at < 3 Z <

a ^ ^ <

< S 2 U 2

o o a: < ^

< a. <> in

q: => CL o u

</) </) '

LU uj

K »- !

< s

Q: >-

3 ui

< _i

<C ®

o 6

o 2
; h; o d m

cc -.3 CE

< <

tt =>

H o

' ^
' « 3 g

^ 1 1 s

3 tD Z

CO ^

m q: O

s ^ >

<0-5

2

° £

m 9- cc

iiis

3 3 3 _,

OuUJliJUJUJUJUJliJliJ

ujttttttttt

COZZZZ2ZZZZ

___________

' 3 Z' O 3 3 O

3 3 3
Z 2 Z
3 3 3

X 3

^ 5 z

^ 5 < < z

2 < UJ O <

:0 ^ 2 U

rr 2 o V)

> m <
CO > -^ z a:

j £

0- < < S 3

g 3 = 5 £

3 -I 0. (D _|

CO CO CO CO CO

UJ UJ UI UJ UJ

t t t t t

-I _l _l _J -I

D 3 3 3 3

2 z 2 Z Z

3 3 3 3 3

OX _ I

I f 1 ^ i

< - S =

Xtoo :ii5tEo<-Q:5

^UjUIrvCOiO'^CuSoS

3 < ^

« > -
CD 3 ^
tr o <

S ^ o ^ *

: o = UJ
J Z CE 03

? s S i

1 < <

■ I O <

[ < < j
: £E (E
) Q o X
c < < p
_|_I_J_|_J0-
0 0 3 3 3

- Z 2

9 P ^ I s:

o ^ CO = S2

Q ^ S O 3

<<<<<<
(E CE CE (E (E CE

Q O O O O Q

< < < < < <1

-I _l -I -I -I _l

3 3 O 3 3 z>

0. 0. Q. Q. O. Q.

-I -J _4 O O

b) in North America

m o

« CE-
^ u <

CD >

CO 2
• O ffl

9 a, CD
< z H
§2<
g o g

<0 t/> Q.

UJ 5

2 ^ 2 li. UJ

3 c 3 (K m

I- CL H

CP CD
CD UJ ^

<0 9

CO

9 :? W

< t ^
- f « s

UI ^ UI ^

8i5^

CD _J U. ^

^ - P < ^

I- UJ Z -J

-1 O <

ffl — 2 to

=> <
(D QC
2 UJ
< U.

I- H K -J -I

to CO < CO

UJ UJ — I UJ

O t t ri t:

UJ o
_i 2
UJ D

ujuj — ^ujS2

1-33033

3 3 H 3 3

[25 “

o S B

— CD
CO <

' CO o 3

0 ZT CD

: o > <
3^2

£E 2 P

'Sis
' > [2 •-
I uj uj

t § t

1 2 -

3

Ij <« <

E UJ
CE CD
^ 2

2 2

■

► u z ®

J *-* o
: < CD to
< § 2
^sg

o “ t

^ 3

s ? ?

^ UI CD UI O ^

<

I '<0 ’(O < ^ ^

I UJ UJ rr DC a:

L- L- S: Q

< O

- 3

=> 3 O

-JtO_IOO_l-J-J_l-lJoJj_IO-J«jJ-l

HOLOCENE a RECENT

IT

PLEISTOCENE

1

PLIOCENE

1

MIOCENE

■ all

OLIGOCENE

VICKSBURG

i;

DD™

1 n

EOCENE

JACKSON

„„„„„nnnnlJLJ

CLAIBORNE

WILCOX

u

TEXT-FIG. 18. Time-Stratigraphic distribution of Tertiary and Quaternary lunulitiform species.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 199

shown in text-fig. 20. As far as the problem of the Oligoeene-Miocene boundary is
concerned, the oldest oeeurrences are the most relevant.

In northern Guarico the earliest oeeurrenee is to be found in the upper part of the
‘Oligocene’ Roblecito formation of well GXB-5. In the Rio Areo of northern Monagas
C. canariensis oeeurs at the base of the Areo Shale, direetly above its eontaet with the
Los Jabillos formation (text-fig. 21). The Areo Shale is generally eonsidered to be the
lateral equivalent of the lower part of the Naricual formation of the Bareelona area.

S. TEXAS

LOUISIANA

W. MISSISSIPPI

E. MISSISSIPPI

SW. ALABAMA

NW. FLORIDA

PLIO-

CENE?

GOLIAD

"CITRONELLE FM."

"CITRONELLE FM."

"CITRONELLE FM,"

CITRONELLE FM, (Type)

"CITRONELLE FM."

MIOCENE

LAGARTO

FLEMING CLAY

PASCAGOULA CLAY

PASCAGOULA CLAY

PASCAGOULA CLAY

CHOCTAW- \
1 HATCHES ]
[ STAGE 1

CANCELLARIA
^ FACIES

ARCA^

YOLDIA )^£CPH0RA

OAKVILLE

HATTIESBURG CLAY

HATTIESBURG CLAY

HATTIESBURG CLAY

3

S ? -
3 « c
< c

R0VE^\ S ^

CATAHOULA FM.
A ANAHUAC
A UPPER FRIO
LOWER FRIO

^ CATAHOULA FM.

m

CATAHOULA FM.

CATAHOULA FM.

■ PAYNES HAMMOCK FM.

TAMPA LIMESTONE

A CHICKASAWHAY FM

A CHICKASAWHAY FM.

SUWANNEE LIMESTONE

? ?

BUCATUNNA CLAY

BUCATUNNA CLAY

OLIGOCENE

VICKSBURG

GROUP

VICKSBURG

0YRAM MARL LENTIL

VICKSBURG GROUP

MINT SPRING MARL LENTIL

CS BYRAM MARL

BYRAM MARL

BYRAM MARL

GLENDON LIMESTONE

GLENDON LIMESTONE

CIlS GLENDON LST,

MARIANNA EQUIVALENT
MINT SPRING MEMBER
*— -* OF MARIANNA

O MARIANNA LST.
MINT SPRING MEMBER
.Of MARIANNA

MARIANNA LST.

MARIANNA LST.

FOREST HILL SAND

FOREST HILL — -yX

_ REOBLUFF CLAY

RED BLUFF CLAY

RED BLUFF CLAY EQUIVALENT

LUNULITES SPP. A CUPULADRIA CANARIENSIS

TEXT-FIG. 19. Occurrences of LwuiUtes spp. and of Ciipiiladria canariensis throughout the Oligocene

and Miocene formations of the Gulf Coast.

whieh itself was considered to be of Lower Oligocene, and possibly even Upper Eoeene,
age (Renz et al. 1958, p. 576). This age assessment can no longer be maintained. Since
they contain C. canariensis, both the upper part of the Roblecito and the Areo Shale (and
for that matter the Narieual) cannot be of more than Aquitanian (or post-Vicksburg)
age. The correctness of this interpretation is reinforeed by the faet that C. canariensis is
aeeompanied in both eases by Discoporella umbellata, another ‘modern’ lunulitiform
species.

It is interesting to compare these oeeurrences with the Caribbean planktonie fora-
miniferal zonation loeally established by the Creole Petroleum Corporation.

According to Creole, the Areo Shale represents the upper two-thirds of the Globorotalia
opima opima zone and the whole of the Globigerina ciperoensis ciperoensis zone. The
oldest known oceurrenees of C. canariensis at the base of the Areo Shale thus fall in the
lower half of the G. opima opima zone.

The underlying Los Jabillos formation has hitherto generally been considered as
Upper Eoeene (fide Feo-Codeeido 1956, p. 331; Renz et al. 1958, p. 576). This age
assessment, too, needs revision in the light of newly aequired evidenee. According to
Creole, the Los Jabillos formation must be post-Eocene, since its stratigraphical position
falls slightly above the base of the G. opima opima zone.

200

PALAEONTOLOGY, VOLUME 6

TEXT-FIG. 20. Occurrences of Cupuladria canariensis throughout the Tertiary formations in the eastern
Venezuela basin (map and correlation chart after Renz et al. 1958, modified).

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

201

Again according to Creole, the Los Jabillos formation is in turn underlain by shales
with Globigerim anipliapertura (not present in the Rio Areo outcrop section shown in
text-fig. 21). It is this G. ampliapertura zone that
on planktonic foraminiferal evidence correlates
with the Vicksburg (Bolli 1957, p. 107). Thus one
might expect to find a different suite of lunuliti-
form Bryozoa at this level. Unfortunately, there
are no earlier records of this group of Bryozoa
from Eastern Venezuela than those mentioned
above. All one can say is that the Vicksburg, as
it is defined in the southern U.S.A., is charac-
terized, from the point of view of the lunulitiform
bryozoan sequences, by the absolute hegemony
of the genus LwmJites (text-fig. 18Z?) and there-
fore correlates with some part of the European
Oligocene (text-fig. 18a). This would imply that
the G. ampliapertura zone is also Oligocene.

In summary it appears that Eames et al. were
substantially correct in lowering the Oligocene-
Miocene boundary in the Caribbean area farther
than any of their critics were prepared to go.

From the presence of modern lunulitiform
Bryozoa it must be concluded that at least the
upper two-thirds of the Globorotalia opima opima
zone is Miocene (Aquitanian or younger). On
the other hand, Eames et al. (1960n, p. 448; 1962,
pp. 48, 49, fig. 5) would definitely seem to be
going too far in wishing to submit the Vicksburg
to the same rejuvenation course. The Vicksburg
lunulitiform bryozoan assemblages have a dis-
tinct Oligocene character, and if the planktonic foraminiferal correlation is correct, this
would imply that the Globigerina ampliapertura zone of the Caribbean sequence of
planktonic foraminiferal zones is also Oligocene. The Oligocene-Miocene boundary in
the Caribbean, subject of so much discussion in the past decade, would thus be pin-
pointed in eastern Venezuela somewhere between the top of the Globigerina ampli-
apertura zone and the top of the lower one-third of the Globorotalia opima opima zone.

The earliest occurrences of C. canariensis in Nigeria of which I am aware have so far
been found in the wells ltuk-1 (range 3,320-3,780 feet) (PI. 26, fig. 3) and Ituk-2 (range
2,770-2,950 feet) in the Calabar flank area, and in Ihuo-1 (range 2,062-2,085 feet).

Acknowledgements. The author wishes to thank the staffs of the museums, the numerous oil companies,
and the many friends and colleagues who have provided the material which forms the basis of this
study, or who have offered suggestions and help in many other ways and whose contribution is not
specifically mentioned in the text.

He is particularly indebted to the staff of the Creole Petroleum Corporation, who, in their comments
on the manuscript, made available important information on the planktonic foraminiferal zonation in
eastern Venezuela and suggested several corrections to text-figs. 20 and 21.

CUPULADRIA CANARIENSIS

TEXT-FIG. 21. Earliest occurrence of Cnpii-
ladria canariensis in the Rio Areo out-
crop section, northern Monagas, eastern
Venezuela (from unpublished Company
reports); 1:5,000.

202

PALAEONTOLOGY, VOLUME 6

The Dutch manuscript was translated by Mr. R. W. Burke; Mr. W. B. Mulder executed the
text figures with great care; Messrs. J. Fournier and J. H. H. van Gigch took some of the photo-
graphs.

REFERENCES

BANDY, o. L. 1954. Distribution of some shallow-water Foraminifera in the Gulf of Mexico. Prof.
Pap. U.S. geol. Sla v. 254-F, 125-40.

BOLLi, H. M. 1957. Planktonic Foraminifera from the Oligocene-Miocene Cipero and Lengua Forma-
tions of Trinidad, B.W.I. Bull. U.S. nat. Mas. 215, 97-123, pi. 22-29.

BRUUN, A. F. 1938. A new occurrence of flying-fish {Cvpsiliiriis heteriirus) in Oslo Fiord. Nytt. Mag.
Naturv. 78, 295-9.

BUGE, E. 1957. Les Bryozoaires du Neogene de I’ouest de la France et leur signification stratigraphique
et paleobiologique. Mem. Mas. Hist, nat., Paris (n.s.), 6, 1-435, pi. 1-12.

BUSK, G. 1854. Catalogue of Marine Polyzoa in the Collection of the British Museum, 2, 55-120,
pi. 69-124. British Museum, London.

1859. A Monograph of the Fossil Polyzoa of the Crag. Palaeont. Soc. Monogr., i-xiii, 1-136,

pi. 1-22.

CANU, F. 1907. Bryozoaires des terrains tertiaires des environs de Paris. Ann. Paleont. 1, 57-88,
pi. 9-12.

1915. Le Systeme hydrostatique zoarial des Bryozoaires chilostomes. C. R. Soc. geol. Fr. 3-4,

21-22.

and BASSLER, r. s. 1920. North American Early Tertiary Bryozoa. Bull. U.S. nat. Mus. 106,

1-879, pi. 1-162.

1929. Bryozoaires eocenes de la Belgique conserves au Musee Royal d’Histoire Naturelle

de Belgique. Mem. Mus. Hist. nat. Belg. 39, 1-69, pi. 1-5.

1930. Bryozoaires marins de Tunisie. Ann. Sta. oceanogr. Salammbd, 5, 1-91, pi. 1-13.

1931. Bryozoaires oligocenes de la Belgique conserves au Musee Royal d’Histoire Naturelle

de Belgique. Mhn. Mus. Hist. nat. Belg. 50, 1-24, pi. 1-4.

and LECOiNTRE, G. 1927. Les Bryozoaires cheilostomes des Faluns de Touraine et d’Anjou.

Mhn. Soc. geol. Fr. (n.s.), 4, 19-50, pi. 6-11.

CHEETiTAM, A. H. 1960. Time, migration, and continental drift. Bull. Amer. /455. Petrol. Geol. 44,
244-51.

CLOUD, p. E. 1959. Paleoecology — retrospect and prospect. J. Paleont. 33, 926-62.

COOKE, c. w., GARDNER, J. and WOODRING, w. p. 1943. Correlation of the Cenozoic Formations of the
Atlantic and Gulf Coastal Plain and the Caribbean Region. Bull. Geol. Soc. Amer. 54, 1713-23.

DARTEVELLE, E. 1933. Contribution a I’etude des Bryozoaires fossiles de I’Eocene de la Belgique.
Ann. Soc. zool. Belg. 63 (1932), 55-116, pi. 2-4.

1935. Zoarial regeneration of free Polyzoa. Ann. Mag. nat. Hist. (10), 15, 559-61, pi. 19.

1943. In RUTSCH, r. Die Mollusken der Springvale Schichten (Obermiocaen) von Trinidad

(British-West-Indien). Verb. 7taturf. Ges. Basel, 54, 107-8.

1952. Bryozoaires fossiles de I’Oligocene de I’Allemagne. Paldont. Z. 26, 181-204.

EAMES, F. E. 1953. The Miocene/Oligocene boundary and the use of the term Aquitanian. Geol. Mag.
90, 388-92.

1954. The Caribbean ‘Oligocene’. Geol. Mag. 91, 326-7.

1955. The Miocene/Oligocene boundary in the Caribbean region. Ibid. 92, 86.

EAMES, F. E., BANNER, F. T., BLOW, w. H. and CLARKE, w. J. 1960fl. Mid-Tertiary stratigraphical
palaeontology. Nature, Fond. 185, no. 4711, 447-8.

19606. [Comments on staineorth 1960c.] Nature, Land. 187, no. 4738, 679-80.

1962. Fundamentals of Mid-Tertiary stratigraphical correlation, i-viii, 1-163,

pi. 1-17. Cambridge.

faura, m. and canu, f. 1916. Sur les Bryozoaires des terrains tertiaires de la Catalogue. Treb. Inst,
catal. Hist. nat. 1-137, pi. 1-9.

FEO-CODECIDO, G. 1956. ‘Los Jabillos formation’. In Stratigraphical Lexicon of Venezuela (English
edition). Bol. Geol., Spec. Publ. 1, 331-2.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 203

FURNESTiN, J. 1960. Hydrologic de la Mediterranee occidentale (Golfe du Lion, Mer catalane, Mer
d’Alboran, Corse orientale), 14 juin-20 juillet 1957. Rev. Trav. Off. Peches marit. 24, 5-119.
GAUTIER, Y. V. 1957. Premiere faunule des Bryozoaires des cotes syriennes. Vie et Milieu, 7,
554-61.

GEYER, R. A. 1950. The occurrence of pronounced salinity variations in Louisiana coastal waters.
J. Mar. Res. 9, 100-10.

GiNSBURG, R. N. 1957. Early diagenesis and lithifaction of shallow-water carbonate sediments in
south Florida. Spec. Piibl. Soc. Ecoii. Paleont. Mineral. 5, 80-99.

GODWIN, H. 1956. The history of the British flora, \ \-2S4, p\. 1-26. Cambridge.

GRiMSDALE, T. F. 1951. Correlation, age determination, and the Tertiary pelagic Foraminifera. Proc.
Third World Petrol. Congr. 1, 463-74.

HARMER, s. F. 1910. ‘Polyzoa’. h\ The Cambridge Natural History, 2, ASS-iM.

1931. Recent work on Polyzoa. Proc. Linn. Soc. Loud., Session 143, 1930/1, 113-68.

HASTINGS, A. B. 1930. Chcilostomatous Polyzoa from the vicinity of the Panama Canal collected by
Dr. C. Crossland on the cruise of the S.Y. ‘St. George’. Proc. zool. Soc. Lond. (1929), 697-740,
pi. 1-17.

HOWE, H. V. 1934. Preliminary paleontologic analysis of the Upper and Lower Chickasawhay mem-
bers of the Catahoula formation. Shreveport Geol. Soc. Guidebook, 11th Ann. Field Trip, 22-28.
KUENEN, PH. H. 1939. The cause of coarse deposits at the outer edge of the shelf. Geol. en Mijnb.
(N.S.), 1, 36-39.

MCGUiRT, J. H. 1941. Louisiana Tertiary Bryozoa. Bull. geol. Surv. La. 21, i-xiii, 1-177, pi. 1-31.
MacNEiL, F. s. 1944. Oligocene stratigraphy of southeastern United States. Bull. Amer. Ass. Petrol.
Geol. 28, 1313-54.

MAPLESTONE, c. M. 1910. On the growth and habits of the Biporae. Proc. roy. Soc. Viet. 23, 1-7.
MENCHER, E. et al. 1953. Geology of Venezuela and its oil fields. Bull. Amer. Ass. Petrol. Geol. 31,
690-777.

NIELSEN, J. N. 1912. Hydrography of the Mediterranean and adjacent waters. Rep. Danish oceanogr.
Exped. Medit. 1908-1910, 1, 77-191, pi. 2-11.

o’donoghue, c. H. and DE WATTEViLLE, D. 1939. The fishery grounds near Alexandria. XX. Bryozoa.

Notes and Mem. Fouad I Inst. Hydrobiol. Fish. 34, 1-58.

PARKER, R. H. 1956. Macro-invei tebiate assemblages as indicators of sedimentary environments in
East Mississippi Delta region. Bull. Amer. Ass. Petrol. Geol. 40, 295-376, pi. 1-8.

PHLEGER, F. B. 1959. Sedimentary patterns of Foraminifera, northern Gulf of Mexico. Int. Oceanogr.
Congr. Prepr. 649-50.

1960. Sedimentary patterns of microfaunas in northern Gulf of Mexico, in Recent sediments,

northwest Gulf of Mexico. Amer. Ass. Petrol. Geol. 267-301, pi. 1-6.

PURI, H. s. 1953. Contribution to the study of the Miocene of the Florida Panhandle. Bull. Geol.
Surv. Florida, 36, 1-345, pi. 1-17.

RENZ, H. H. et cd. 1958. The eastern Venezuela Basin. In Habitat of Oil. Amer. Ass. Petrol. Geol.
551-600.

SCRUTON, p. c. 1956. Oceanography of Mississippi Delta sedimentary environments. Bull. Amer.
Ass. Petrol. Geol. 40, 2864-952.

SHEPARD, F. p. 1932. Sediments of the continental shelves. Bull. geol. Soc. Amer. 43, 1017-40.

1948. Submarine Geology, i-xvi, 1-348. New York.

siLEN, L. 1942. On spiral growth of the zoaria of certain Bryozoa. Ark. Zool. 34A, 1-22, pi. 1-5.

1944. The anatomy of Labiostomella gisleni Silen (Bryozoa Protocheilostomata). Kungl. Svenska

Vetensk-Akad. Handl. (3), 21, 1-111, pi. 1-5.

1947. Conescharellinidae (Bryozoa Gymnolaemata) collected by Prof. Dr. Sixten Bock’s expedi-
tion to Japan and the Bonin Islands 1914. Ark. Zool. 39, 1-61, pi. 1-5.

SPAiNK, G. 1958. De Nederlandse Eemlagen. I. Algemeen overzicht. Wet. Meded. Kon. Ned. Natuur-
hist. Veren. 29, 1-44.

STACH, L. w. 1936. Correlation of zoarial form with habitat. J. Geol. 44, 60-65.

STAiNFORTH, R. M. 1954. Comments on the Caribbean Oligocene. Geol. Mag. 91, 175-6.

■ 1960n. Estado actual de las correlaciones transatlanticas del Oligo-Mioceno por medio de

foraminiferos planctonicos. Bol. Geol., Publ. espez. 3, Terc. Congr. Geol. Venezol. 1, 382-406.

204

PALAEONTOLOGY, VOLUME 6

STAiNFORTH, R. M. 19606. Current status of transatlantic Oligo-miocene correlation by means of
planktonic Foraminifera. Rev. Micropal. 2, 219-30.

1960c. The American Oligocene. Nature, Loud. 187, no. 4738, 678-9.

STRAATEN, L. M. J. u. van. 1956. Composition of shell beds formed in tidal flat environment in the
Netherlands and in the Bay of Arcachon (France). Geol. en Mijnb. (n.s.), 18, 209-26.

SVERDRUP, H. u., JOHNSON, M. w., and FLEMING, R. H. 1960. The Oceans, i-x, 1-1087. Englewood
Cliffs, N.J.

THORSON, G. 1955. Modern aspects of marine level-bottom animal communities. J. Mar. Res. 14,
387-97.

ULRICH, E. o. and bassler, r. i. 1904. Bryozoa. In Miocene, Maryland geol. Siirv., pp. 404-29, pi. 109-
18.

u.s. COAST and geodetic survey. 1955. Surface water temperatures at tide stations Atlantic coast
North and South America. Spec. Pitbl. U.S. Dept. Commerce, 278, 1-69.

viGNEAUX, M. 1949. Revision des Bryozoaires Neogenes du Bassin d’ Aquitaine et essai de classifica-
tion. Mem. Soc. geol. Fr. (n.s.), 28, 1-155, pi. 1-11.

vooRTHUYSEN, J. H. Van. 1956. Plioceen. Mariene afzettingen. In Geologische geschiedenis van Neder-
land, 1-154, pi. 1-16. ’s-Gravenhage.

WATERS, A. w. 1921. Observations upon the relationships of the (Bryozoa) Selenariadae, Cone-
scharellinidae, &c., fossil and Recent. J. Linn. Soc. (Zool.), 34, 399-427, pi. 29, 30.

1926. Ancestrula of Cheilostomatous Bryozoa. V. Cupularia, &c. Ann. Mag. nat. Hist. (9), 18,

424-33, pi. 18.

wiLLGOHS, J. F. 1953. Common Loggerhead Caretta caretta (Linne) stranded in western Norway.
Univ. Bergen Arb. 1952, Naturv. R. 17, 1-8.

WILLS, L. J. 1951. A palaeogeographical atlas of the British Isles and adjacent parts of Europe, 1-64,
pi. 1-22. London.

WILSON, D. p. 1952. The influence of the nature of the substratum on the metamorphosis of the larvae
of marine animals, especially the larvae of Ophelia bicornis Savigny. Ann. Inst. Oceanogr. 27,
49-156.

WORTHINGTON, L. V. 1959. The 18° water in the Sargasso Sea. Deep-sea Res. 5, 297-305.

wOsT, G. 1960. Die Tiefenzirkulation des Mittellandischen Meeres in den Kernschichten des Zwischen-
und des Tiefenwassers. Dtsch. hydrogr. Z. 13, 105-31.

APPENDIX

1. Synonymy

1959« Cupularia canariensis busk, g. On some Madeiran Polyzoa. Collected by J. Yales Johnston, Esq.

Zoophytology. Quart. J. micr. Sci. 7, 66, pi. 23, figs. 6-9.

18596 Cupularia canariensis Busk: busk, g. A Monograph of the Fossil Polyzoa of the Crag. Palaeont.
Soc. Monogr., p. 87, pi. 13, fig. 2.

1868 Cupularia canariensis Busk: manzoni, a. Saggio di Conchiologia foss. Subappennina, p. 71.

1869 Cupularia canariensis Busk: manzoni, a. Bryozoi Pliocenici Italian!. S.B. Akad. Wiss. Wien,

59, p. 26, pi. 2, fig. 17.

1873 Membranipora canariensis (Busk): smitt, f. a. Floridan Bryozoa collected by Count L. F. De
Pourtales. II. Kungl. Svenska Vetensk.-Akad. Handl. 11, p. 10, pi. 2, figs. 69-71.

1877fl Cupularia canariensis Busk: manzoni, a. I briozoi fossili del Miocene d’Austria ed Ungheria.

II. Denkschr. Akad. Wiss. Wien, 37, p. 72, pi. 17, fig. 56.

18776 Cupularia canariensis Busk: manzoni, a. Bryozoaires du Pliocene superieur de file de Rhodes.
Mem. Soc. geol. Fr. (3), 1, p. 67.

1878 Cupularia canariensis Busk: broeck, e. van den. Esquisse geologique & paleontologique des

depots plioctoes des environs d’Anvers. Ann. Soc. Malac. Belg. 9 (1874), p. 274.

1879 Cupularia canariensis Busk: seguenza, g. Le formazioni terziarie nella provincia di Reggio
(Calabria). Mem. Accad. Lined (3), 6, pp. 131, 371.

Cupularia canariensis Busk: coppi, f. Paleontologia modenese o guida al paleontologo. Modena,
p. 123.

1881

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS 205

1885 Limulites conica Defrance var. depressa: lorie, j. Contributions a la geologic des Pays-Bas.

Resultats geologiques et paleontologiques des forages de puits a Utrecht, Goes et Gorkum.
Arch. Mus. Teyler (2), 2, p. 133, pi. 3, fip. 15a, b.

1887 Cupidaria canariensis Busk: pergens, e. Pliocane Bryozoen von Rhodos. Ann. natiirh. {Mus.)
Hofnuis. 2, p. 31.

1 889 Cupidaria canariensis Busk : waters, a. w. Supplementary Report on the Polyzoa collected by

H.M.S. Challenger during the years 1873-1876. Rep. Sci. Res. Voy. Challenger, Zoology,
31, pt. 79, p. 36.

1 890 Cupidaria canariensis Busk : namias, i. Contributo ai briozoi pliocenici delle provincie di Modena

e Piacenza. Boll. Soc. geol. ital. 9, p. 506.

1891 Cupidaria canariensis Busk: neviani, a. Contribuzione alia conoscenza dei briozoi fossili italiani.

Briozoi postpliocenici del sottosuolo di Livorno. Ibid. 10, p. 130.

1894 Cupidaria canariensis Busk: broeck, e. van den. Materiaux pour la connaissance des depots

pliocenes superieurs rencontres dans les derniers travaux de creusement des Bassins Maritimes
d’Anvers, Bassin Africa (ou Lefebre) et Bassin America. Bull. Soc. beige Geol. 6 (1892), p.
139.

1895a Cupidaria canariensis Busk: neviani, a. Briozoi fossili della Farnesina e Monte Mario presso
Roma. Palaeontogr. ital. 1, p. 101.

\895b Cupidaria canariensis Busk: neviani, a. Briozoi neozoici di alcune localita dTtalia, I. Boll. Soc.
Roniana Stiidi Zool. 4, pp. 238, 243.

1895 Cupidaria canariensis Busk: de angelis, j. Descripcion de los Briozoos fosiles pliocenicos de

Cataluha. Barcelona, p. 9, pi. B, figs. 6-9.

1896 Cupidaria canariensis Busk: neviani, a. Briozoi neozoici di alcune localita dTtalia, III. Boll.

Soc. Roniana Studi Zool. 5, p. 121.

1 898 Cupidaria canariensis Busk : neviani, a. Briozoi neozoici di alcune localita dTtalia, V. Ibid. p. 38.

1900 Cupidaria canariensis Busk: neviani, a. Briozoi terziari e posterziari della Toscana. Boll. Soc.

geol. ital. 19, p. 362.

1901 Cupidaria canariensis Busk: neviani, a. Briozoi neogenici delle Calabrie. Palaeontogr. ital.

6 (1900), p. 168.

1905 Cupidaria canariensis Busk: neviani, a. Briozoi fossili di Carrubare (Calabria). Boll. Soc. geol.
ital. 23, p. 522.

1907 Cupidaria canariensis Busk: calvet, l. Bryozoaires. Exped. sci. ‘'Travailleur' et ^Talisman

1880-1883, 8, p. 393.

1908 Cupidaria canariensis Busk: canu, f. Iconographie des Bryozoaires fossiles de 1’ Argentine.

An. Mus. nac. B. Aires, 17, p. 275, pi. 5, figs. 8-10.
non 1908 Cupidaria canariensis Busk: robertson, a. The incrusting chilostomatous Bryozoa of the
west coast of North America. Univ. Calif. Publ. Zool. 4, p. 314, pi. 24, figs. 90, 9\=Dis-
coporella umbel lata (Defrance).

1909 Cupidaria guineensis Busk: norman, a. m. The Polyzoa of Madeira and neighbouring islands.

J. Linn. Soc. (Zool.), 30, p. 289, pi. 37, figs. 2-6 (non Cupidaria guineensis : Busk, G. Catalogue
of Marine Polyzoa in the Collection of the British Museum, 2 (1854), p. 98, pi. 1 14, figs. 1-5).
1913a Cupidaria canariensis Busk: canu, f. Contributions a I’etude des Bryozoaires fossiles. IV.

Pliocene d’Alger. Bull. Soc. geol. Fr. (4), 13, p. 124.

\9\3b Cupidaria canariensis Busk: canu, f. Contributions a I’etude des Bryozoaires fossiles. X. Astien
de Valle Andona. Ibid., p. 128.

1914 Cupidaria guineensis Busk: osburn, r. c. The Bryozoa of the Tortugas Islands, Florida. Publ.

Carnegie Instn. 182, p. 194 (non Cupidaria guineensis: Busk, G. Catalogue of Marine Polyzoa
in the Collection of the British Museum, 2 (1854), p. 98, pi. 114, figs. 1-5).

1914 Cupidaria canariensis Busk: faura, m. Sobre la presencia de un briozoo vivente, la Cupidaria
canariensis descubierto en los terrenos miocenicos de Cataluna. Bol. Soc. esp. Hist. nat. 14,
p. 397.

1916 Cupidaria canariensis Busk: faura, m. and canu, f. Sur les Bryozoaires des terrains tertiaires

de la Catalogne. Treb. Inst, catal. Hist. nat. p. 77, text-figs. 4a, b, pi. 3, figs. 7, 8.

1917 Cupidaria canariensis Busk: canu, f. Les Bryozoaires fossiles des Terrains du Sud-Ouest de la

France. X. Burdigalien. Bull. Soc. geol. Fr. (4), 16 (1916), p. 137, pi. 3, figs. 4-6.

PALAEONTOLOGY, VOLUME 6

206

1919a Cupularia canariemis Busk; canu, f. and bassler, r. s. Bryozoa of the Canal Zone and related
areas. Bull. U.S. nat. Mits. 103 (1918), p. 119, pi. 53, figs. 5-7.

\9\9b Cupiilaclrici canarie?isis (Busk): canu, f. and bassler, r. s. Fossil Bryozoa from the West Indies.
Publ. Carnegie Instn. 291, p. 78, pi. 1, figs. 8-10.

1920a Cupuladria canariensis (Busk): canu, f. Contributions a I’etude des Bryozoaires fossiles. XV.

Redonnien du Pigeon-Blanc (Loire-Inferieure). Bull. Soc. geol. Fr. (4), 19 (1919), p. 213.
\910b Cupuladria canariensis (Busk); canu, f. Contributions a I’etude des Bryozoaires fossiles. XVI.
Scaldisien d’Anvers. Ibid., p. 213.

1 920 Cupuladria canariensis ( Busk) : duvergier, j. Note sur les Bryozoaires du Neogene de I’Aquitaine.

Act. Soc. linn. Bordeaux, 72, p. 149.

1921 Cupuladria canariensis (Busk): cipolla, f. I briozoi pliocenici di Altavilla presso Palermo. G. Sci.

nat. econ. Palermo, 32 (1918-20), p. 31, pi. 2, figs. 22-24.

1921 Cupularia canariensis Busk : waters, a. w. Observations upon the relationships of the (Bryozoa)
Selenariadae, Conescharellinidae, &c., fossil and Recent. /. Linn. Soc. (Zool.), 34, p. 410
(partim), pi. 29, figs. 1-5; pi. 30, figs. 11, 12, 21, 22, 25.

1923 Cupuladria canariensis (Busk): canu, f. and bassler, r. s. North American Later Tertiary
and Quaternary Bryozoa. Bull. U.S. nat. Mas. 125, p. 28, pi. 1, figs. 7-9.

1923 Cupuladria biporosa canu, f. and bassler, r. s. North American Later Tertiary and Quaternary

Bryozoa. Ibid., p. 29, pi. 47, figs. 1, 2.

1924 Cupuladria canariensis (Busk): cipolla, f. I briozoi del Quaternario inferiore dei dintorni di

Palermo. Nota preliminare. Boll. Soc. Sci. nat. econ. Palermo, 6, p. 7.

1925a Cupuladria canariensis (Busk): canu, f. and bassler, r. s. Contribution a I’etude des Bryo-
zoaires d’Autriche et de Hongrie. Bull. Soc. geol. Fr. (4), 24 (1924), p. 673.

19256 Cupuladria canariensis (Busk): canu, f. and bassler, r. s. Les Bryozoaires du Maroc et de
Mauritanie. Memoire. Mem. Soc. Sci. nat. Maroc, 10, p. 13.

1925 Cupuladria canariensis (Busk): canu, e. and lecointre, g. Les Bryozoaires cheilostomes des

Faluns de Touraine et d’Anjou. Mem. Soc. geol. Fr. (n.s.), 4, p. 17, pi. 3, figs. 10, 11.

1925 Cupuladria canariensis (Busk): Mansfield, w. c. Miocene Gastropods and Scaphopods from
Trinidad, British West Indies. Proc. U.S. nat. Mus. 66, pp. 5, 8.
non 1926 Cupularia canariensis Busk: waters, a. w. Ancestrulae of Cheilostomatous Bryozoa. Part
V. Cupularia, &c. Ann. Mag. nat. Hist. (9), 18, p. 426, text-fig. on p. 425, pi. 18, figs. 1, 10
= Cupuladria guineensis (Busk).

1928a Cupuladria canariensis (Busk): canu f. and bassler, r. s. Fossil and Recent Bryozoa of the
Gulf of Mexico Region. Proc. U.S. nat. Mus. 72 (14), p. 15, pi. 1, figs. 7-9.

19286 Cupuladria canariensis (Busk): canu, f. and bassler, r. s. Les Bryozoaires du Maroc et de
Mauritanie. 2™*^ Memoire. Mem. Soc. Sci. nat. Maroc, 18, p. 16.

1929 Cupuladria canariensis (Busk); canu, f. and bassler, r. s. Bryozoa of the Philippine Region.

Bull. U.S. nat. Mus. 100, p. 73, pi. 3, figs. 1, 2.

1 930 Cupuladria canariensis (Busk) : Hastings, a. b. Cheilostomatous Polyzoa from the vicinity of the

Panama Canal collected by Dr. C. Crossland on the Cruise of the S.Y. ‘St. George’. Proc.
zool. Soc. Lond. 47, p. 714, pi. 8, figs. 38, 40.

1934 Cupulardia [51c] canariensis (Bu?,k.): howe, h. v., hanna, m. a., and gravell, d. w. Fossil plates

with explanations. Fossils present in subsurface formations of Clarke and Wayne Counties,
Mississippi, and in wells in South Mississippi. Shreveport Geol. Soc., Guide Book 11th Ann.
Field Trip, pi. 1 , figs. 27, 28.

1935 Cupuladria canariensis (Busk): dartevelle, e. Zoarial regeneration of free Polyzoa. Ann. Mag.

nat. Hist. (10), 15, p. 559, pi. 19, figs. 1, 2.

1936 Cupuladria canariensis (Busk): scotti, p. Briozoi fossili del Miocene della Collina di Torino

(Collezione Rovasenda). Atti Accad. Torino, 71, p. 410.

1 938 Cupularia canariensis Busk : gillard, p. a. Sur les Bryozoaires helvetiens des Faluns de la Vienne.
C. R. Soc. geol. Fr. 9, p. 155.

1940 Cupuladria canariensis (Busk): osburn, r. c. Bryozoa of Porto Rico with a resume of the West

Indian bryozoan fauna. Sci. Surv. P. R. 16, p. 354.

1941 Cupuladria caaar/c/w/.y (Busk): mcguirt, j. h. Louisiana Tertiary Bryozoa. Bull. Geol. Surv. La.

21, p. 46, pi. 1, figs. 1-3, 5-6, 8.

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

207

1942 Cupuladria canariensis (Busk): silen, l. On spiral growth of the zoaria of certain Bryozoa.

Ark. Zool. 34A, p. 13, pi. 4, figs. 15, 16.

1943 Cupuladria canariensis (Busk): dartevelle, e. In rutsch, r. Die Mollusken der Springvale

Schichten (Obermiocaen) von Trinidad (British-West-Indien). Verh. naturf. Ges. Basel, 54,
p. 107.

1945 Cupuladria canariensis \ huge, e. Empreintes et monies internes de Bryozoaires de I’Hel-

vetien et du Redonien de Bretagne. Bull. Soc. geol. Fr. (5), 15, p. 590.

1946 Cupuladria canariensis (Busk): roger, j. and huge, e. Les Bryozoaires du Redonien. Ibid.

16, pp. 218, 226.

1 947 Cupuladria canariensis (Busk) : osburn, r. c. Bryozoa of the Allan Hancock Atlantic Expedition,

1939. Rep. Allan Hancock Atlant. Exped. 5, p. 10.

1949 Cupuladria canariensis (Busk): vigneaux, m. Revision des Bryozoaires Neogenes du Bassin

d’Aquitaine et essai de classification. Mem. Soc. geol. Fr. (n.s.), 28, p. 38, pi. 2, figs. 8, 9.

1950 Cupuladria canariensis (Busk): osburn, r. c. Bryozoa of the Pacific Coast of America. I.

Cheilostomata-Anasca. Rep. Allan Hancock Pacif. Exped. 14, p. 33, pi. 3, figs. 2, 3.

1951 Cupuladria canariensis (Busk): malecki, j. Contribution a la connaissance des Bryozoaires du

Miocene de Benczyn. Ann. Soc. geol. Pologne, 19 (1949), p. 488.

1952 Cupuladria canariensis (Busk): lagaaij, r. The Pliocene Bryozoa of the Low Countries, &c.

Meded. geol. Sticht., serie C-V-5, p. 33, pi. 2, figs. \a, b.

1953 Cupuladria canariensis (Busk): lagaaij, r. The vertical distribution of the lunulitiform Bryozoa

in the Tertiary of the Netherlands. Ibid, (n.s.), 7, p. 15, pi. I, figs. \a, b.

1955 Cupuladria canariensis (Busk): gautier, y. Bryozoaires de Castiglione. Bull. Statist. Aquicidt.

Peche Castiglione (n.s.), 7, p. 231.

1956 Cupuladria canariensis (Busk): barker, r. h. Macro-invertebrate assemblages as indicators of

sedimentary environments in East Mississippi Delta region. Bull. Amer. ^4.^5. Petrol. Geol.
40, pp. 311, 335, pi. 4, fig. 25.

1957 Cupuladria (Busk): buge, e. Les Bryozoaires du Neogene de I'ouest de la France, &c.

Mem. Mus. Hist, nat., Paris (n.s.), 6, p. 139, pi. 9, fig. 5; pi. 10, fig. 3.

1957 Cupuladria canariensis (Busk): gautier, y. Sur quelques Bryozoaires des Cotes d’Algerie.

Result. Camp. "Professeur Lacaze Duthiers', 2, suppl. 6 ( Vie et Milieu), p. 102.

1 959 Cupuladria canariensis (Busk) : soule, j. d. Results of the Puritan- American Museum of Natural
History expedition to western Mexico. 6. Anascan Cheilostomata (Bryozoa) of the Gulf of
California. Amer. Mus. Novit., No. 1969, p. 8.

1961 Cupuladria canariensis (Busk): reguant, s. Los briozoos del Neogeno espanol. Notas Inst,
geol. Esp. 62, 111.

1961 Cupuladria canariensis (Busk): galopim de carvalho, a. m. Note sur des Bryozoaires du Plio-
cene de Pombal. Bol. Soc. geol. Portug. 14, 97, pi. 1, figs. 1-3.

1961 Cupuladria canariensis (Busk): GHiURCA, v. Contributii la cunoa§terea faunei de Bryozoare din

Transilvania (II). Revizuirea taxonomica a Bryozoarelor de la Lapugiu si Biuturi publicate
de A. Koch. Stadia Univ. Babes-Bolyai (2), 1, table.

1962 Cupuladria canariensis (Busk): gautier, y. v. Recherches ecologiques sur les Bryozoaires

chilostomes en Mediterranee occidentale. Thesis, Aix-en-Provence, 53.

All cited papers are listed in either the References or the Synonymy.
bmnh: British Museum (Natural History).
usnm: United States National Museum.

ksepl: Koninklijke/Shell Exploratie en Produktie Laboratorium, Volmerlaan 6, Rijswijk Z.H.,
Netherlands.

text-fig. 3. The selected localities shown are:

2. Documentation of maps and figures

Reference or
repository

Reference or

repository

1 . Cape Rosa, Algeria

2. Ras-el-Amouch, Algeria

Dartevelle 1935

BMNH

3. Oran, Algeria

4. Tangier Bay

Waters 1921
BMNH

208

PALAEONTOLOGY, VOLUME 6

Reference or

Reference or

repository

repository

5.

Vanneau Sta. XXIX, Morocco

Canu and Bass-

28.

Lat. 28° 4' N, long. 93° 44' W,

KSEPL

ler 19256

40 fms.

6.

Cape Sagres, Portugal

BMNH

29.

Lat. 25° 47' N, long. 96° 27' W,

KSEPL (ex Mr.

7.

Off Punta Delgada, Azores

Silen 1942

50 fms.

R. W. Barker)

8.

Madeira

Busk 1859a;

30.

Albatross Sta. 2361, NE of

USNM

Norman 1909;

Cabo Catoche

Silen 1942

31.

Port Antonio, Jamaica

USNM

9.

‘entree de la Bocayna’, Canary

Calvet 1907

32.

Caroline Sta. 55, Samana Bay,

USNM

Islands

Santo Domingo

10.

Gran Canaria, Canary Islands

Silen 1942

33.

Off Guanica Harbor, Puerto

Osburn 1940

11.

Lat. 27° 16' N, long. 23° 21' W

Silen 1942

Rico

12.

Cap Blanc, Mauritania

Calvet 1907

34.

St. Thomas, West Indies

Silen 1942

13.

Cape Verde, Senegal

BMNH

35.

St. Martin, West Indies

Silen 1942

14.

Conakry, Guinee

BMNH

36.

Albatross Sta. 2145, NNW of

Silen 1942

15.

Petit Tahou, Liberia

Waters 1889,

Aspinwall

1921

37.

Cape la Vela, Colombia

Osburn 1947

16.

Accra, Ghana

BMNH

38.

Aruba, Netherlands Antilles

Osburn 1947

17.

SE of Lagos, Nigeria, 13 fms.

KSEPL

39.

Tortuga Island, Venezuela

Osburn 1947

18.

Off Fishtown River, Nigeria,

KSEPL

40.

Margarita Island, Venezuela

Osburn 1947

14 fms.

41.

Barbados, West Indies

BMNH

19.

N of Fernando Poo, 42 fms.

KSEPL

42.

Lat. 9° 28' N, long. 60° 8' W, 39

KSEPL (ex Dr. D.

20.

Calypso Sta. 45, W of Libre-

BMNH

fms.

J. Nota)

ville, Gabon

43.

Berbice, British Guiana

BMNH

21.

Albatross Sta. 2597, off Cape

USNM

44.

Off Bahia, Brazil

Waters 1889

Hatteras

45.

Cedros Island, Lower Cali-

Osburn 1950

22.

Albatross Sta. 2415, E of Jack-

USNM

fornia

sonville, Fla.

46.

Albatross Sta. 3012, Gulf of

USNM

23.

Albatross Sta. 2639, Straits of

Canu and Bass-

California

Florida

ler 1928a

47.

Albatross Sta. 2826, Gulf of

Canu and Bass-

24.

Tortugas, Florida

Smitt 1873;

California

ler, 1929

Osbum 1914;

48.

Clarion Island, Mexico

Osburn 1950

Silen 1942

49.

Socorro Island, Mexico

Osbum 1950

25.

Albatross Sta. 2405, off Cedar

Canu and Bass-

50.

West coast of Mexico

Osburn 1950

Keys, Fla.

ler 1928a

51.

West coast of Costa Rica

Osburn 1950

26.

S of Mobile, Alabama

Parker 1956

52.

West coast of Panama

Osburn 1950

27.

Lat. 28° 40' N, long. 90° 14' W,

KSEPL

53.

Gorgona, Colombia

Hastings 1930

19 fms.

54.

Galapagos Islands

Osburn 1950

55.

Ecuador

Osbum 1950

TEXT-FIG. 4. The selected localities shown are:

Miocene

1 . Twistringen, SSW of Bremen,

Germany

2. Beeringen, Netherlands

3. Wyneghem, near Antwerp,

Belgium

4. Vienna Basin, Austria

Aa. Benczyn, near Wadowice,
Poland

5. Turin, Italy

6. Reggio, Calabria, Italy

7. Ferriere I’Arcon, Indre-et-

Loire, France

8. Dept. Gironde, France

9. Barcelona province, Spain

Reinbek/Dingdener Stufe, Middle
Miocene (PI. 26, fig. 2)
Diepboring 15, 154-1 59 m., Middle
Miocene

Lower Diestian (Deumian), Upper
Miocene

Tortonian, Middle Miocene
Tortonian, Middle Miocene
Miocene

Tortonian, Middle Miocene
Pontilevian, Middle Miocene

Aquitanian and Burdigalian, Lower
Miocene

Helvetian, Middle Miocene

ex Dr. C. W. Drooger
Lagaaij 1953
Lagaaij 1952

Manzoni 1877a; Canu and Bassler
1925n

Malecki 1951

Scotti 1936
Seguenza 1879

Canu and Lecointre 1925; Buge
1957

Canu 1917; Duvergier 1920; Vig-
neaux 1949

Faura 1914; Faura and Canu 1916

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

209

10. Niger Delta, Nigeria

1 1 . Calabar area, Nigeria

12. Missellele River, 10 km. N of

Tiko, E of Mount Cam-
eroon, Nigeria

13. French Cameroons

14. Monroe County, Florida

15. Jackson’s Bluff, Ochlockonee

River, Leon County, Florida

16. Shell Bluff, Shoal River, N

of Mossyhead, Walton
County, Fla.

17. Limestone Creek, Wayne

County, Mississippi

18. East Baton Rouge Parish,

Louisiana

19. Laguna Atorcosa, Cameron

County, Texas

20. Goliad County, Texas

21. Gulf of Mexico, off Nueces

County, Texas

22. Bowden, Jamaica

23. Cercado de Mao, Santo Dom-

ingo

24. Port Limon, Costa Rica

25. Rio Coro, at Caujarao, Distr.

Miranda, Estado Falcon,
Venezuela

26. Cabo Blanco, near Maiquetia

Airport, Distr. Federal,
Venezuela

27. Estado Guarico, Venezuela

28. Estado Anzoategui, Vene-

zuela

29. Estado Monagas, Venezuela

30. Estado Monagas, Venezuela

3 1 . Cubagua Island, Venezuela

32. Concord quarry. Point h
Pierre, Trinidad

Exploration well Oloibiri-1 , 9,560-
9,570', Miocene

Exploration well Ituk-1, 3,320-
3,780', Miocene (PI. 26, fig. 3)
Lower Miocene

Middle Miocene

Caloosahatchee formation, Mio-
cene

Choctawhatchee, Miocene

Shoal River (type locality), Mio-
cene

Chickasawhay, Miocene (PI. 26,
fig. 4)

Superior Oil Prod. Company’s
Duplantier Community No. 1
well, 7,671-7,687', Heterostegina
zone; 8,091-8,101', Marginulina
zone, Catahoula formation, Mio-
cene

Shell-Continental Fee 1 , 1 1 ,8 10-

11,850', Marginulina zone, Mio-
cene

Housh, Thompson & Crown Cen-
tral C. G. Wood 1, 3,163-
3,193', updip limit of Anahuac
shale wedge, Miocene

Gulf Oil Corp. and others. Block
889, 3-A, 10,300', Upper Frio,
Miocene

Bowden Beds, upper Middle Mio-
cene

Middle Miocene

Middle Miocene

Caujarao formation, upper Middle
Miocene

Playa Grande formation. Upper
Miocene

Creole Petroleum Corp., GXB-5,
Roblecito formation, Miocene

Mene Grande Oil Company, OG-1 ,
2,125'; 3,250', Freites formation;
3,750-4,625', Oficina formation,
Miocene

Creole Petroleum Corp., ORC-2,
3,900-4,200', Carapita forma-
tion, Miocene

Compania Shell de Venezuela,
Cerro Negro-3, Freites forma-
tion ; Oficina formation, Miocene

Socony Mobil Oil Company, Cuba-
gua-1, 218', Cubagua formation.
Upper Miocene (PI. 26, fig. 5)

Concord marl, Miocene

KSEPL

KSEPL

KSEPL

KSEPL

Canu and Bassler 19196; 1923
Canu and Bassler 19196; 1923

KSEPL

KSEPL (ex Mr. R. W. Barker)
McGuirt 1941

KSEPL (ex Dr. D. D. Bannink)

Shell Oil Company, Corpus Christ!
Division

KSEPL (ex Mr. E. M. Wilkins)

Canu and Bassler 1919u; 19196;
1923

Canu and Bassler 19196; 1923
Canu and Bassler 1923

KSEPL

KSEPL

Compania

Caracas

Shell

de

Venezuela,

Compania

Caracas

Shell

de

Venezuela,

Compania

Caracas

Shell

de

Venezuela,

Compania

Caracas

Shell

de

Venezuela,

KSEPL

KSEPL (ex Dr. E. Th. N. Spiker)

C 1015

P

210

PALAEONTOLOGY, VOLUME 6

Pliocene

I. Sutton, Suffolk, England
II. Antwerp, Belgium

III. Heumensoord, Netherlands

IV. Roden, Netherlands

V. Contigne, Maine-et-Loire,
France

VI. Pigeon-Blanc (Landreau),
near Nantes, France
VIn. Pombal, Portugal
VII. El Ampurdan, near Bar-
celona, Spain

VIII. Valle Andona, Asti, Pied-
mont, Italy

IX. Province of Modena, Italy
X. Altavilla, near Palermo,
Sicily, Italy

XI. Nador, near Algiers, Algeria
XII. Terrebonne Parish, Louis-
iana

XIII. Minnitimmi Creek, Bocas

Island, Almirante Bay,
Panama

XIV. Camarones, 10 km. E of

Esmaralda, Ecuador

Pleistocene

A. Livorno, Italy

B. Monte Mario and Farnesina,

near Rome, Italy

C. Carrubare, Calabria, Italy

D. Palermo, Sicily, Italy

E. Rhodes

F. Gulf of Mexico, Mississippi

River Delta area, Louisiana

G. Gulf of Mexico, High Island

area

H. Gulf of Mexico, off Mata-

gorda County, Texas

J. Cabo Blanco, near Maiquetla

Airport, Distr. Federal,
Venezuela

K. Estado Monagas, Venezuela

L. Puerto Militar, Bahia Blanca,
Argentine

Gedgravian, Pliocene
Scaldisian (prob. Luchtbal hori-
zon), Pliocene

Boring 1928, 102-40-10900 m.,
‘Scaldisian’, Pliocene
Boring 114/23, 184-50-215-00 m.,
Pliocene

Redonian, Pliocene

Redonian, Pliocene

Pliocene

Pliocene

Astian, Pliocene

Plaisancian, Pliocene
Pliocene

Plaisancian, Pliocene
Terrebonne Gas Company Fee
4^ 1, 2,300', Pliocene
Pliocene

Pliocene

Sicilian, Pleistocene
Calabrian

Sicilian?

Sicilian?

Sicilian

South Pass, Block 6 Area, State
Lease 2590 1, 500-1,100', ‘Upper
Marine’, Pleistocene
Federal Block A-104, Shell-Con-
tinental ‘Neptune-1’, 775-790';
835-1,000', Pleistocene
Ohio Oil Company, State of Texas
403 44: 1, 537-599'; 662-754';
1,935-1,997', Pleistocene
Mare formation. Pleistocene?

Compania Shell de Venezuela,
Guanipa-1, 640-670', Paria for-
mation, Pleistocene
Pampean, Pleistocene

Busk 18596
Canu 19206

Lagaaij 1952

KSEPL (ex Dr. J. H. van Voort-
huysen)

Roger and Buge 1946; Buge 1957

Canu 1920a; Roger and Buge
1946; Buge 1957
Galopim de Carvalho 1961
De Angelis 1895

Manzoni 1869; Canu 19136

Namias 1890
Cipolla 1921

Canu 1913a
McGuirt 1941

Canu and Bassler 1928a

KSEPL (ex Mr. J. Brouwer)

Neviani 1891

Manzoni 1869; Neviani 1895

Neviani 1905
Cipolla 1924

Manzoni 18776; Pergens 1887;
Silen 1942

KSEPL (ex Mr. J. J. Gouty)

KSEPL

KSEPL

KSEPL

Compania Shell de Venezuela,
Caracas

Canu 1908

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

211

TEXT-FIG. 5. Sources: {a) Collections from the Exploration and Production Research Laboratory,
Shell Development Company, Houston, Texas, now at ksepl, Rijswijk, Netherlands; {b) Parker 1956;
and (c) various other sources (see list) :

Depth

in

Depth

in

Lat. N

Long. IV fms.

Lat. N

Long. W

fms.

1.

Cavalier 1956 Sta. 128

29=02'54"

89°28'26"

5-5

47.

Atlantis 1951 Sta. 168

28°51'30"

83=3930"

12

2.

Parker 1956 Sta. 410

29°27'

89°15'

2

48.

169

28°54'

83°34'30"

11-5

3.

321

29°30'30"

89°15'

2

49.

170

28°55'

83°28'30"

11

4.

322

29°30'30"

89T0'

2

50.

171

28°56'30"

83°22'

8

5.

G. 2074

29°30'

88°52'

7-5

51.

172

28°58'

83°16'30"

7?6?

6.

Parker 1956 Sta. 346

29°24'

88°54'

10

52.

173

28»59'

83°12'30"

5

7.

349

29°22'

88°51'

15

53.

Off Egmont Key, Florida

27°36'27"

83°08'57"

11-5

8.

316

29°12'

88°43'

39

54.

27°36'27"

83°04'48"

10-5

9.

G. 2021

29°09'

88°43'

43

55.

27°36'27"

83°02'42"

8-5

10.

G. 2028, Sandpile Bank

20°04'40"

88°43'

49

56.

27°36'27"

83°00'30"

8-5

11.

Parker 1956 Sta. 311

29=22'

88°32'

32

57.

27°36'27"

82°53'48"

7

12.

309

29=23'

88°22'

30

58.

ITl&ir

82°51'42"

5-5

13.

308

29°26'

88°20'

30

59.

Tampa Bay, Florida

27»38'o9"

82°37'27"

4-5

14.

244

29°26'

88°08'

29

60.

27°39'54"

82°36'06"

5-5

15.

U.S.S. Hydrographer

61.

27°4i'i4"

82°34'43"

4-5

1941/2, Sta. 50

29°46'05"

86°58'

103-8

62.

27°46'

82°31'36"

4

16.

U.S.S. Hydrographer

63.

Vema-3 1954 BT# 18

2TQ?,'

83°30'

23

1941/2, Sta. 23

29°3U

86°26'

77

64.

Albatross 1885 Sta. 2409

17.

U.S.S. Hydrographer

(Silen 1942)

2TW

83'>21T5"

26

1941/2, Sta. 45

29°33'

86°08'

21

65.

Vema-3 1 954 Dredge =|:{: 2

26°50'

83°40'

29

18.

U.S.S. Hydrographer

66.

SSW of John's Pass,

1941/2, Sta. 40

30°15'

86°ir

15

Florida

26°25'

83°32'

34

19.

U.S.S. Hydrographer

67.

Vema-3 1954 BT4:ii2

26°10'

84°20'

96

1941/2, Sta. 39

30°13'

86°08'

12-5

68.

Vema-3 1954 Dredge 4i: 1

25°50'

84°30'

117

20.

Off Panama City, Florida

30°05'58"

85°45'39"

10

69.

Tortugas, Florida (Os-

21.

30°0T24"

85°51'54"

14-5

burn 1914)

10

22.

30°00'45"

85°52'45"

15

70.

SW of Tortugas, Florida

23.

Albatross 1885 Sta. 2405

(Silen 1942)

40 m.

(Canu and Bassler

71.

Rebecca Shoal, Florida

1928a)

28°45'

85°02'

30

(Silen 1942)

15 m.

24.

Fish Hawk Sta. 7157

72.

Albatross 1885 Sta. 2315

(usnm)

29=23'

83°41'45"

9

(Silen 1942)

24°26'

8r48'15"

37

25.

Off Cedar Keys, Florida

28°58'48"

83°16'42"

5-5

73.

NW of Sombrero Key,

26.

29°00'06"

83°15'

5

Florida

24°39'36"

81°04'12"

6-5

27.

29°01'24"

83°13'20"

4-5

74.

S of Sombrero Key, Flo-

28.

29°02'36"

83°ir38"

4

rida

24°35'30"

8U06'55''

25

29.

29°03'55"

83°09'50"

3-5

75.

SE of Molasses Reef

30.

29°05'16"

83°08'

3-5

Light, Fla.

24°54'

80°15'30"

122

31.

Atlantis 1951 Sta. 152

28°09'

85°07'

100

76.

Albatross 1886 Sta. 2639

32.

153

28°11'30"

85°02'

80

(Canu and Bassler

33.

154

28°14'

84°57'

64

1928a)

25°04'50"

80°15T0''

56

34.

155

28°16'30"

84°52'

43

77.

Due E of Carysfort Light,

35.

156

28°19'

84°46'30"

32

Fla.

25°13T5"

80"07'

78

36.

157

28°22'

84°14'

34

78.

Off Turtle Harbor, Flo-

37.

158

28°24'30"

84°36'

33

rida (usnm)

20

38.

159

28=27'

84°3U

28

79.

Due E of Triumph Reef,

39.

160

28°30'

84°25'30"

25

Fla.

25°28'30"

80°05'45"

45

40.

161

28°32'

84°20'

22

80.

Due E of Triumph Reef,

41.

162

28°35'

84°14'

20

Fla.

25°28'30"

80°05'

58

42.

163

28°38'

84°08'

20

81.

Off Fowey Light, Florida

25°30'48"

80°03'36"

80

43.

164

28°41'

84°02'

17

82.

Fowey Light, Florida

44.

165

28°43'30"

83°56'

17

(Canu and Bassler

45.

166

28°46'

83°50'

16

1928a)

40

46.

167

28°49'

83°44'

14

212

PALAEONTOLOGY, VOLUME 6

TEXT-FIG. 6. Sources: (a) Collections from the Exploration and Production Research Laboratory, Shell
Development Company, Houston, Texas, now at ksepl, Rijswijk, Netherlands; (b) Parker 1956:

Depth

in

Lat. N

Long. ]V

fins.

1.

Cavalier 1956 Sta. 12

28°43'18"

95°19'09"

9

2.

20

28°47'30"

95°15'24"

10

3.

21

28°45'16"

95°15'24"

10-5

4.

23

28°42'47"

95°14'30"

12

5.

28

28°37'03"

95°09'54"

15

6.

30

28°34'50"

95°08'05"

16

7.

35

28°44'48"

95°08'05"

11-5

8.

Univ. of Houston,

Nos. 1^

28°40'

94°33'

15

9.

Vema-3 1954 Core

i|t 65 (Top 12")

28°10'48"

94°15'

30

10.

Heald Bank, N of

Heald Bank Light

8-5

11.

Heald Bank

29°04'

94°14'

10-5

12.

Cavalier 1956 Sta. 331

29°03T5"

94°18'45"

6-5

13.

330

29°01'06"

94°18'45"

8

14.

329

28°58'45"

94°18'45"

6

15.

328

28°55'42"

94°18'06"

11-5

16.

327

28°49'36"

94°16'42"

13-5

17.

326

28°43'44"

94°14'54"

15

18.

High Island area. Block

A- 104

28°42'12"

94°01'36"

15-5

19.

Cavalier 1956 Sta. 325

28°37'06"

94°13'14"

18

20.

324

28°33'

94°12'15"

20

21.

323

28°26'06"

94°10'30"

23

22.

321

28°17'30"

94°08'30"

27-5

23.

320

28° 15 '24"

94°03'54"

31-5

24.

319

28°09'06"

94°00'00"

38

25.

318

28°02'54"

93=5130"

40

26.

317

27°58'45"

93°49'45"

50

27.

316

27°54'15"

93°47'45"

50

28.

315

27°48'45"

93=4530"

92

29.

314

27°44'12"

93°43'48"

138

30.

312

27°48'

93°47'36"

102

31.

311

27°51'30"

93°47'

70

32.

310

27°53'45"

93°46'30"

60

33.

309

27°56'15"

93°46'

57

34.

308

28°04'

93°44'20"

40

Depth

in

Lat. N

Long. \V fins.

35.

Cavalier 1956 Sta. 307

28°09'30"

93°43'15"

35-5

36.

. 306

28°16'15"

93°42'15"

35

37.

. 305

28=22T2"

93°40'30"

30

38.

. 304

28°29'06"

93°38'30"

24

39.

. 303

28°37'24"

93°36'24"

18-5

40.

. 302

28°43'

93°34'45"

15

41.

. 301

28°49'

93°33'15"

12-5

42.

. 300

28°52'30"

93032'

11

43.

. 299

28°59'15"

93°30'15"

11

44.

. 298

29°06'

93°28'12"

11

45.

. 297

29°11'30"

93=26"

9

46.

. 296

29=1 5'09"

93°24'08"

8-5

47.

Sabine Bank

area.

Sta. 2

29=i9'20"

93°36'30"

7

48.

Cavalier 1956 Sta. 294

29=2 r04"

93°21'18"

8-5

49.

. 293

29°23'48"

93°20'04"

6-5

50.

. 292

29°24'42"

93°20'

7

51.

. 291

29°26'24"

93=19'55"

4

52.

. 290

29°27'28"

93»19'50"

6-5

53.

. 289

29°30'21"

93°19'38"

6-5

54.

. 288

29°33'30"

93°19'26"

6-5

55.

. 286

29°40'35"

93°19'40"

5-5

56.

. 273

28°31'30"

9 1=49' 15"

24

57.

. 278

28=05' 15"

91°58'

38

58.

. 236

28°42'24"

90°59'38"

7

59.

. 235

28°39'50"

90°59'38"

8-5

60.

. 217

28°33'12"

90°59'50"

14

61.

. 226

28°08'30"

9i°or

56

62.

. 227

28°06'30"

91°0L

37-5

63.

. 229

28°04'

91°0L

57

64.

. 166

28°37'

90° 16' 12"

21-5

65.

. 175

28°10'

90°13'30"

66

66.

. 180

28°39'55"

90=1 3 '45"

19

67.

. 128

29°02'54"

89°28'26"

5-5

68.

Parker 1956, Sta

410

29=27'

89°15'

2

69.

. 321

29°30'30"

89°15'

2

70.

. 322

29°30'30"

89°10'

2

71.

. 346

29=24'

88°54'

10

TEXT-FIG. 7. Source: samples kept at ksepl, Rijswijk, Netherlands; the co-ordinates and depths are
listed below:

Mees C renter
1959 Stations

Lat. N

Long. E

Depth

in

fins.

1

158

6° 11 '20"

3°21'26"

44

2

320

6°20'04"

3=29'i4"

13

3

169

6°18'24"

3034,22"

19

4

168

6°20'31"

3°46'15"

14

5

170

6° 11 '50"

3°32'04"

42

6

162

6°07'25"

3°30'01"

63

7

163

6°08'45"

3°33'22"

52

8

164

6°10'24"

3°37'26"

44

9

171

6°11'25"

3°37'12"

43

10

165

6°12'00"

3°42'32"

41

11

318

6°11'01"

3°46'21"

47

12

215

6° 13 '59"

4°11'38"

12

13

147

6°05'40"

4°25'41"

21

14

212

5°42'43"

4°40'30"

37

15

203

5°32'00"

4°49'38"

27

Depth

Mess Crenier

in

1959 Stations

Lat. A'

Long. E

fms.

16

206

5°24'01"

4=39'51"

120

17

133

5°19'58"

5°01'29"

16

18

137

5°12'46"

4=46'59"

109

19

310

4°54'09"

5°08'26"

25

20

311

4=48'55"

5°02'55"

45

21

312

4=47'34"

5°01'38"

55

22

107

4°37'17"

5°24'22"

12

23

108

4=34'47"

5°21'18"

18

24

109

4=32' 19"

5°18'04"

29

25

377

4°24'10"

5°35'49"

10

26

376

4°23'33"

5°33'53"

11

27

375

4=23 '01"

5°32'04"

13

28

374

4°22'34"

5°29'53"

14

29

373

4°22'07"

5°27'58"

17

30

372

4=21 '41"

5°25'58"

21

R. LAGAAIJ: THE BRYOZOAN

Depth

Mees Cremer

in

J959 Stations

Lat. N

Long. E

fms.

31

371

4°21'14"

5°23'58"

27

32

299

4=25 '07"

5°34'55"

9

33

300

4°2T56"

5°31'46"

13

34

301

4° 18 '26"

5 027 '29"

21

35

96

4° 15 '39"

5°42'55"

9

36

97

4°12'28"

5°40'38"

12

37

98

4°09'20"

5°38'25"

14

38

99

4°05'44"

5°35'37"

17

39

100

4°02'10"

5°33'06"

26

40

101

3059,55"

5031,23"

34

41

220

4°06'56"

5°37'31"

16

42

221

4°04'50"

5°40'50"

17

43

222

4°00'55"

5°43'11"

18

44

223

3°57'06"

5°45'07"

23

45

224

3°52'51"

5°47'47"

31

46

395

4°04'34"

5°50'10"

14

47

396

4°04'09"

5°52'23"

14

48

397

4°02'47"

505434"

14

49

398

4'=01'25"

5°56'21"

15

50

399

4°00'37"

5°58'38"

15

51

400

3°59'54"

6°00'59"

16

52

401

3°58'57"

6°03'18"

18

53

402

3°58'17"

6°05'42"

21

54

403

3057,27"

6°07'50"

23

55

388

4°06'28"

6°02'25"

10

56

389

4°04'40"

6°0T34"

12

57

390

4°02'41"

6°00'52"

13

58

391

4°00'49"

6°00'08"

14

59

392

3°59'01"

5°59'32"

16

60

393

3°57'10"

5°58'58"

22

61

394

3°55'37"

5°58'24"

24

62

83

4°10'39"

6°10'25"

7

63

387

4=08 '44"

6°06'38"

8

64

84

4°05'41"

6°07'53"

11

65

85

4°02'49"

6°07'04"

14

66

86

4°00'14"

6°06'26"

17

67

87

3°56'42"

6°05'02"

23

68

81

4°09'13"

6°18'27"

9

69

344

4°iril"

6°26'59"

8

70

345

4°04'33"

6°26'11"

15

71

346

3°57'37"

6°24'40"

31

72

74

40 1231"

6°31'22"

8

73

287

4° 10' 19"

6°43'14"

11

74

288

4°06'I6"

6°43'20"

18

75

289

4W11"

6°43'24"

32

CUPULADRIA CANARIENSIS 213

Depth

Mess Cremer

in

1959 Stations

Lat. N.

Long. E

fms.

76

291

3°50'04"

6°43'19"

61

77

77

4°13'41"

6°47'27"

9

78

78

4°14'05"

6°51'59"

9

79

79

4°14'41"

6“56'58"

8

80

11

4°08'46"

7°01'31"

16

81

6

4°06'46"

7°00'54"

20

82

10

4°02'31"

7°02'05"

29

83

7

4°00'31"

7°01'12"

36

84

404

3°56'30"

6°58'42"

48

85

334

4°11'18"

7°18'47"

15

86

335

4°05'35"

7°18'12"

22

87

338

3°46'57"

7°18'17"

91

88

243

4°14'41"

7°35'01"

10

89

244

4° 10' 18"

7°34'56"

17

90

245

4°05'55"

7°34'59"

25

91

246

4°01'37"

7°34'54"

32

92

247

3°57'16"

7°34'56"

42

93

260

3°50'30"

7°30'10"

76

94

248

3°51'58"

7°35'00"

71

95

259

3°48'34"

7“39'49"

93

96

274

4°16'25"

7°42'38"

9

97

23

4°15'58"

8°01'50"

12

98

24

4° 11 '50"

8°02'33"

14

99

31

3057,38"

8°04'39"

36

100

28

3°55'27"

8°05'30"

42

101

29

3°51'43"

8°05'49"

68

102

32

3'’56'07"

8°08'35"

40

103

34

3°52'40"

8°18'02"

49

104

36

3049' 19"

8°17'22"

100

105

255

3°47'59"

8=20' 19"

62

106

37

3°47'02"

8°20'17"

91

107

73

4°05'38"

8°26'10"

22

108

72

4°05'21"

8°30'16"

26

109

254

3°52'38"

8°28'02"

40

no

38

3°44'50"

8°23'50"

75

111

39

3°43'08"

8°27'22"

56

112

46

4°06'34"

8°34'35"

34

113

45

4°03'36"

8°36'10"

31

114

44

3°58'36"

8°35'43"

36

115

43

3°54'05"

8°35'01"

41

116

42

3°48'53"

8°33'59"

44

117

41

3°44'39"

8°33'01"

44

118

40

3°40'36"

8°32'11"

44

119

70

3°48'10"

8°43'41"

42

TEXT-FIG. 8. Distribution of Pliocene marine facies after van Voorthuysen 1956, fig. 26. Eighteen
borings and outcrop localities listed in Lagaaij 1 952, p. 33, are shown, to which the following occurrences
north of the Rhine have here been added :

Wanneperveen-2

Oud-Appelscha

Roden

Kernboring NAM
114/23

171-50-224-50 m.
1 87-25-201 -00 m.
191 -50-21 3 00 m.

214

PALAEONTOLOGY, VOLUME 6

TEXT-FIG. 9. Southern boundary of eastern Venezuela Basin after Mencher et al. 1953, fig. 7; northern
boundary coincides with ‘mountain front’.

The following Miocene occurrences are shown:

Cabo Blanco, near Maiquetia Airport, Distr. Federal .....

Cubagua Island, Socony Mobil Oil Company, Cubagua-1, 218' ....

Estado Guarico, Creole Petroleum Corp., GXB-5 ......

. . Socorro-3 ......

Estado Guarico, Venezuelan Atlantic Refining Company, Tucupido-2
Estado Guarico, Sinclair Oil & Refining Company, Piedra Azul-1
Estado Guarico, Venezuelan Atlantic Refining Company, Guama-1 .

Estado Guarico, Companla Shell de Venezuela, Shothole 6043 ....

.... . . Shothole 1 146 ....

Estado Guarico, Sinclair Oil & Refining Company, Camoruco-1, 2,350-2,790'

Estado Anzoategui, Companla Shell de Venezuela, VC-1

Cagigal-1, 750-775'

.. 4,450-5,025' .

Cagigal-2, 2,200-2,225' .

.. 6,775-6,800' .

Estado Anzoategui, Mene Grande Oil Company, Casca-1, 750-1,225'
. . 3,375-3,425'

Estado Anzoategui, Phillips Oil Company, B-1, 625-1,175' . . . .

Estado Anzoategui, Creole Petroleum Corp., El Roble-1 (RPN-1), 925-950', 1,300-
1,325'

Estado Anzoategui, Socony Mobil Oil Company, Guario-1, 1,675-1,700',
2,200-2,225'

Estado Anzoategui, Mene Grande Oil Company, OG-1, 2,125-2,150', 3,250-3,275'

3,750-1,650' .

Estado Anzoategui, Texas Petroleum Company, Mata~l .....
Estado Monagas, Creole Petroleum Corp., El Salto-1 .....

Estado Monagas, Texas Petroleum Company, Sotil-1 .....
Estado Monagas, Creole Petroleum Corp., Hato-1 3 . . . . . .

Estado Bolivar, Companla Shell de Venezuela, Cerro Negro- 1 ....

.... . . Cerro Negro-2 ....

Estado Monagas, Companla Shell de Venezuela, Cerro Negro-3

Estado Anzoategui, Companla Shell de Venezuela, outcrop sample St. 2992

.... . . outcrop sample St. 3000

.... . . outcrop sample Ol. 445 .

Estado Monagas, Companla Shell de Venezuela, outcrop sample Ol. 375, Rio Areo

.... .. outcrop sample Ol. 381, Rio Areo

Estado Monagas, Mene Grande Oil Company, SB- 13 3, 925-6,250'

Estado Monagas, Creole Petroleum Corp., JGE-28 ......

Monaq-1 . . . . .

. . J-274, 5,325-6,700' . . . .

.. ORC-1, 2,800-3,800'

. . ORC-2, 3,900-4,200'

. . ORC-9

Estado Monagas, Companla Shell de Venezuela, ORS-3 .....
Estado Monagas, Creole Petroleum Corp., QGE-26, 2,900-4,300'

Playa Grande formation
Cubagua formation
Roblecito formation
Chaguaramas formation
Chaguaramas formation
Chaguaramas formation
Chaguaramas formation
Upper Santa Ines
Upper Santa Ines
Chaguaramas formation
Lower Santa Ines
Naranja equivalent
Verde equivalent, Oficina
formation
Moreno member
Verde member, Oficina
formation
Blanco member
Moreno member, Oficina
formation

Blanco member, Oficina
formation

Blanco member, Oficina
formation

Azul member, Oficina
formation
Freites formation
Oficina formation
Oficina formation
Freites formation, Oficina
formation
Oficina formation
Freites formation ; Oficina
formation
Oficina formation
Freites formation ; Oficina
formation

Freites formation ; Oficina
formation
Capaya formation
Capaya formation
Capaya formation
Areo shale
Areo shale

Carapita and Capaya
formations

Buena Vista shale ( = Areo
shale)

Carapita formation
Carapita formation
Carapita formation
Carapita formation
Carapita formation
Carapita formation
Carapita formation

R. LAGAAIJ: THE BRYOZOAN CUPULADRIA CANARIENSIS

215

Estado Monagas, Creole Petroleum Corp., QGE-29. ..... Carapita formation

Trinidad, Point-a-Pierre, Shell Trinidad Ltd., outcrop sample Ho. 207, Concord
Quarry ............. Concord marl

Trinidad, Caroni County, Couva Ward, Springvale, near Couva, about J to 1 mile
S of Milton, outcrop sample (Mansfield 1925; Dartevelle 1943) . . . . Springvale formation

TEXT-FIG. 12. The four stations shown are:

Parker 1956, Sta. 321, lat. 29° 30' 30" N, long. 89° 15' W
Tampa Bay, Florida, lat. 27° 46' N, long. 82° 31' 36" W
Cavalier 1956 Sta. 286, lat. 29° 40' 35" N, long. 93° 19' 40" W
Mees Cremer 1959 Sta. 83, lat. 4° 10' 39" N, long. 6° 10' 25" E

TEXT-FIG. 13. The eight stations shown are:

NW Gulf of Mexico: bathythermogram — Mabel Taylor 1932 Sta. 1106

C. canancniw— Cavalier 1956 Sta. 314, 27° 44' 12" N, 93° 43' 48" W
NE Gulf of Mexico: bathythermogram — Mabel Taylor 1932 Sta. 903

C. canariensis — Atlantis 1951 Sta. 152, 28° 09' N, 85° 07' W
Sargasso Sea: bathythermogram — Atlantis 1933 Sta. 1483, 21° 46' N, 62° 48' W
C. canariensis — Caroline 1933 Sta. 94, 18° 39' N, 65° 03' 30" W
Gulf of Guinea: bathythermogram — Meteor 1926 Sta. 235, 3° 33' 36" N, 5° 6' 42" E

C. canariensis — Mees Cremer 1959 Sta. 206, 5° 24' 01" N, 4° 39' 51" E

TEXT-FIG. 14. The three occurrences shown are:

Albatross Sta. 2415, lat. 30° 44' N, long. 79° 26' W, 440 fms.

Port Antonio, Jamaica, 150 fms.

Caroline 1933 Sta. 94, lat. 18° 39' N, long. 63° 03' 30" W, 300-470 fms.

TEXT-FIG. 15. The Middle Miocene occurrences shown are those listed under text-fig. 4 as Nos. 1, 2, 4,
4n, 5, 6, 7, 8, and 9; the Pliocene occurrences shown are those listed under text-fig. 4 as Nos. I, III,
IV, VI, VIo, VII, VIII, IX, X, and XI ; the Early Pleistocene occurrences shown are those listed under
text-fig. 4 as Nos. A, B, C, D, and E; the Recent occurrences shown are those listed under text-fig. 3
as Nos. 1, 2, 3, 4, 5, and 6.

TEXT-FIG. 17. The Pliocene occurrences shown are those listed under text-fig. 4 as Nos. I, II, III, IV, V,
VI, VIu, VII, VIII, IX, X, and XI; the Recent occurrences shown are those listed under text-fig. 3 as
Nos. 1, 2, 3, 4, 5, and 6.

TEXT-FIG. 18fl. Compiled from Canu 1907; Canu and Bassler 1923, 1929, 1931; Dartevelle 1933, 1952;
Manzoni 1869; Vigneaux 1948, and others. Not shown on this chart, on the assumption of being
junior synonyms, are:

Liinulites rhomboidalis Von Munster [ = Discoporella umbellata (Defrance); fide Canu and Bassler 1929, p. 11]
Lumdites angusticostata Canu and Bassler [= Liinulites siibplena Keuss] fide Dartevelle 1933, p. 57; 1952, p. 183]

TEXT-FIG. \%b. Compiled from Canu and Bassler 1920, 1923, 1928fl; McGuirt 1941 ; Ulrich and Bassler
1904 and others; and incorporating own observations, e.g. the occurrence of Lumdites ligidata (Canu
and Bassler) in the Byram marl at Byram, Pearl River, Mississippi, and the occurrences of Ciipitladria
canariensis, C. doina (d’Orbigny) and Discoporella umbellata (Defrance) in the Pleistocene of the
Gulf Coast at the localities listed under text-fig. 4 as Nos. F and G.

TEXT-FIG. 19. Correlation chart largely based on one made by Mr. R. W. Barker, with the south Texas
column (1) here added and the NW Florida column (6) adapted to include the latest views of Puri
(1953). The following occurrences of Lumdites spp. are shown:

Column 2 (Louisiana)

Rosefield, Catahoula Parish ...... Vicksburg (McGuirt 1941)

Column 3 (W Mississippi)

Mint Springs Bayou, near Vicksburg .... Mint Springs marl, ksepl (ex Mr. J. J. Gouty)
Byram .......... Byram marl (Canu and Bassler 1920)

216

PALAEONTOLOGY, VOLUME 6

Column 4 (E Mississippi)

Red bluff, Wayne County .......

Horton’s Mill Creek, near Highway 45, Wayne County

Locality CX, W bank of Chickasawhay River, Wayne County

Column 5 (SW Alabama)

One mile N of Monroeville ......

St. Stephens Bluff, Tombigbee River .....
Column 6 (NW Florida)

Road cut, U.S. Highway 90, at Chipola River

Red Bluff clay (Canu and Bassler 1920)
Basal Marianna limestone, ksepl (ex Mr.
R. W. Barker)

Paynes Hammock sand (1 fragment), ksepl
(ex Mr. R. W. Barker)

‘Chimney Rock’ member of Marianna lime-
stone (Canu and Bassler 1920)

Glendon, ksepl (ex Mr. E. H. Rainwater)

Marianna limestone, ksepl

The following occurrences of Ciipiiladria canariensis are shown:

Column 1 (S Texas)

Gulf of Mexico, off Nueces County, Gulf Oil Corp. and others.

Block 889, 3-A, 10,300'

Goliad County, Housh, Thompson & Crown Central C. G.

Wood#l, 3,163-3,193'

Column 2 (Louisiana)

East Baton Rouge Parish, Superior Oil Producing Company’s
Duplantier Community No. 1 well, 7,671-7,687' .

East Baton Rouge Parish, Superior Oil Producing Company’s
Duplantier Community No. 1 well, 8,091-8,101' .

Column 4 (E Mississippi)

Limestone Creek, Wayne County .....
Column 5 (SW Alabama)

Choctaw Bluff, Alabama River, Clarke County .

Column 6 (NW Florida)

Shell Bluff, Shoal River, N of Mossyhead, Walton County .
One mile below Baileys Ferry, Chipola River
Jackson Bluff, Ochlockonee River, i mile above Florida
Highway 20 bridge .......

Ibid

W. D. McDaniel’s farm, near Red Bay, Walton County.

Upper Frio, ksepl (ex Mr. E. M. Wilkins)
Updip limit of Anahuac shale wedge, ksepl

Heterostegina zone, Catahoula formation
(McGuirt 1941)

Marginiilina zone, Catahoula formation
(McGuirt 1941)

Chickasawhay, ksepl (ex Mr. R. W. Barker)

Chickasawhay, ksepl (Mr. W. McGlamery
Coll. ; ex Mr. R. W. Barker)

Shoal River (type locality), ksepl

Chipola marl (Canu and Bassler 1923)

Choctawhatchee stage, Ecplwra facies, Pec-
ten Bed, KSEPL

Choctawhatchee stage, Cancellaria facies,
KSEPL

Choctawhatchee stage. Area facies, marl
with Area rubisiana, ksepl

TEXT-FIG. 20. The following occurrences of Ciipuladrio canariensis are shown:

Column 1 (Northern Guarico)

Creole Petroleum Corp., GXB-5 ......

Sinclair Oil & Refining Company, Piedra Azul-1
Venezuelan Atlantic Refining Company, Tucupido-2 .

Column 2 (Greater Anaco area)

Compania Shell de Venezuela, Cagigal-1, 750-775'

. . 4,450-5,025

Mene Grande Oil Company, Casca-1, 750-1,225'

. . 3,375-3,425' .

Socony Mobil Oil Company, Guario-1, 1,675-1,700', 2,200-2,225'
Phillips Oil Company, B-1, 625-1,175' .....

Column 3 (Greater Oficina area)

Texas Petroleum Company, Mata-1 ......

Mene Grande Oil Company, OG-1, 2,125-2,150', 3,250-3,275' .

. . 3,750-4,650'

Column 4 (Temblador area)

Compania Shell de Venezuela, Cerro Negro-1 ....
Compania Shell de Venezuela, Cerro Negro-3 ....

Roblecito formation
Chaguaramas formation
Chaguaramas formation

Naranja equivalent

Verde equivalent, Oficina formation

Blanco member

Moreno member, Oficina formation
Azul member, Oficina formation
Blanco member, Oficina formation

Oficina formation
Freites formation
Oficina formation

Oficina formation
Freites formation

R. LAGAAIJ; THE BRYOZOAN CUPULADRIA CANARIENSIS

217

Column 5 (NW Anzoategui: ‘mountain front’)

Compania Shell de Venezuela, VC-1 . . . . .

Column 6 (N Anzoategui)

Compania Shell de Venezuela, outcrop samples St. 2992, St. 3000
Column 7 (Northern Monagas)

Creole Petroleum Corp., JGE-28 ......

Mene Grande Oil Company, SB-133, 925-6,250'

Creole Petroleum Corp., J-274, 5,325-6,700' . . . .

Lower Santa Ines

Capaya formation

Buena Vista shale (=Areo shale)
Carapita and Capaya formations
Carapita formation

R. LAGAAIJ

Koninklijke/Shell Exploratie
en Produktie Laboratorium,
Volmeiiaan 6,
Rijswijk Z.H.,

Manuscript received 12 May 1962 Netherlands

PRINTED IN GREAT BRITAIN
AT THE UNIVERSITY PRESS, OXFORD
BY VIVIAN RIDLER
PRINTER TO THE UNIVERSITY

J

i

W

<-rJh&

lb.

THE PALAEONTOLOGICAL ASSOCIATION

COUNCIL 1962-3

President

Professor T. Neville George, The University, Glasgow, W. 2
Vice-Presidents

Professor O. M. B. Bulman, Sedgwick Museum, Cambridge
Professor W. F. Whittard, The University, Bristol
Dr. W. H. C. Ramsbottom, Geological Survey Office, Leeds

Treasurer

Professor P. C. Sylvester-Bradley, The University, Leicester
Assistant Treasurer

Dr. T. D. Ford, The University, Leicester
Secretary

Dr. C. H. Holland, Department of Geology, Bedford College, London, N.W. 1

Editors

Mr. N. F. Hughes, Sedgwick Museiun, Cambridge
Dr. W. S. McKerrow, University Museum, Oxford
Dr. Gwyn Thomas, Department of Geology, Imperial College of Science, London, S.W. 7

Other members of Council

Dr. F. T. Banner, British Petroleum Company, Sunbury on Thames
Dr. F. M. Broadhurst, The University, Manchester
Dr. A. J. Charig, British Museum (Natural History), London
Dr. L. R. Cox, British Museum (Natural History), London
Dr. W. T. Dean, British Museum (Natural History), London
Dr. C. Downie, The University, Sheffield
Dr. R. Goldring, The University, Reading
Dr. J. C. Harper, The University, Liverpool
Dr. M. R. House, The University, Durham
Dr. J. W. Neale, The University, Hull
Dr. R. J. G, Savage, The University, Bristol
Mr. J. J. D. Smith, Geological Survey and Museum, London
Dr. C. D. Waterston, Royal Scottish Museum, Edinburgh
Mr. C. W. Wright, London

Overseas Representatives

Australia: Professor Dorothy Hill, Department of Geology, University of Queensland, Brisbane
Canada: Dr. D. J. McLaren, Geological Survey of Canada, Department of Mines and Technical
Surveys, Ottawa

India: Professor M. R. Sahni, Department of Geology, Panjab University, Chandigarh
New Zealand: Dr. C. A. Fleming, New 2fealand Geological Survey, P.O. Box 368, Lower Hutt
West Indies and Central America: Dr. L. J. Chubb, Geological Survey Department, Kingston, Jamaica
Eastern U.S.A.: Professor H. B. Whutington, Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge 38, Mass.

Western U.S.A.: Dr. J, Wyatt Durham, Department of Paleontology, University of California,
Berkeley 4, Calif.

PALAEONTOLOGY

VOLUME 6 • PART 1

CONTENTS

Trepostome Bryozoa from the Caradoc Series, Shropshire. By june r. p. pmLLiPS

ROSS 1

Plant microfossils from the Lower Triassic of Western Australia. By B. E.

BALME 12

The morphology and development of species of Marssonella and Pseudo-
textulariella from the Chalk of England. By tom Barnard 41

Wealden mammalian fossils. By william a. clemens 55

Lower Permian Bryozoa from Western Australia. By June r. p. Phillips ross 70

On the interpretation and status of some hystrichosphere genera. By c. downie
and w. A. s. sarjeant 83

Phyllocarid crustaceans from the Silurian and Devonian of Czechoslovakia.

By rvo chlupaC 97

The composition of Pyritosphaera barbaria Love 1957. By Leonard g. love 119
Some Upper Tremadocian graptolites from Norway. By nils spjeldn.es 121

Alaiophyllum mackenziense sp. nov., a Devonian tetracoral from Canada.

By A. E. H. PEDDER 132

The lamellibranch genus Prothyris in the Upper Devonian and Carboniferous of
Great Britain. By R. B. wilson 136

A rapid parallel grinding machine for serial sectioning of fossils. By r. d.
HENDRY, A. J. ROWELL, and J. W. STANLEY 145

Malayomaorica gen. nov. (Family Aviculopectinidae) from the Indo-Pacific
Upper Jurassic; with comments on related forms. By j. a. jeletzky 148

Cyprilepas holmi Wills 1962, a pedunculate cirripede from the Upper Silurian
of Oesel, Esthonia. By Leonard j. wills 161

The bryozoan genus Polypora M‘Coy. By t. g. miller 166

Cupuladria canariensis (Busk) — ^portrait of a bryozoan. By r. laagaaij 172

PRINTED IN GREAT BRITAIN AT THE UNIVERSITY PRESS, OXFORD
BY VIVIAN RIDLER, PRINTER TO THE UNIVERSITY

VOLUME 6 * PART 2

Palaeontology

MAY 1963

PUBLISHED BY THE
PALAEONTOLOGICAL ASSOCIATION
LONDON

Price £3

THE PALAEONTOLOGICAL ASSOCIATION

The Association was founded in 1957 to further the study of palaeontology. It holds
meetings and demonstrations, and publishes the quarterly journal Palaeontology.
Membership is open to individuals, institutions, libraries, &c., on payment of the
appropriate annual subscription :

Institute membership . . . . £5. 5j. (U.S. $15.50)

Ordinary membership . . . . £3. 3>s. (U.S. $9.50)

Student membership . . . £2. 2s. (U.S. $6.50)

There is no admission fee. Student members will be regarded as persons receiving
full-time instruction at educational institutions recognized by the Council; on first
applying for membership, they should obtain an application form from the Secretary
or the Treasurer. Subscriptions are due each January, and should be sent to the
Treasurer, Dr. T. D. Ford, Department of Geology, The University, Leicester,
England.

Palaeontology is devoted to the publication of papers (preferably illustrated) on all
aspects of palaeontology and stratigraphical palaeontology. Four parts are published
each year and are sent free to all members of the Association. Members who join for
1963 will receive Volume 6, Parts 1 to 4.

All back numbers are stiU in print and may be ordered from B. H. Blackwell,
Broad Street, Oxford, England, at the prices shown below (post free) :

Vol. 1 (for 1957-8) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 2 (for 1959) in 2 parts at £2 or U.S. $6.00 per part.

Vol. 3 (for 1960) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 4 (for 1961) in 4 parts at £2 or U.S. $6.00 per part.

Vol. 5 (for 1962) in 4 parts at £3 or U.S. $9.00 per part.

A complete set. Volumes 1-5, consists of 18 parts and costs £40 or U.S. $120.

Manuscripts on aU aspects of palaeontology and stratigraphical palaeontology are
invited. They should conform in style to those already published in this journal, and
should be sent to Mr. N. F. Hughes, Sedgwick Museum, Cambridge, England.
A sheet of detailed instructions for authors wMl be supphed on request.

LOWER KIMERIDGIAN AMMONITES FROM
THE DRIFT OF LINCOLNSHIRE

by the late w. J. arkell and J. h. callomon^

Abstract. A collection of ammonites is described from the drift of Lincolnshire ; they are of Lower Kimeridgian
age, Mutabilis Zone. Almost all of the specimens belong to the genera Rasenia and Aiilacostephamis, and include
several new species which are grouped into two new subgenera of Aiilacostephamis-. Xenostephaims and Xeiio-
stephanoides. These new forms have only recently been found in situ in England, and are known from the
Kimeridge Clay of both east and west Scotland.

This paper describes a collection of ammonites made mostly by Dr. R. G. Thurrell
while mapping in the Lower Cretaceous of north Lincolnshire, and now deposited in
the Sedgwick Museum, Cambridge. The material is all from the drift, and hence does
not provide primary stratigraphic evidence ; its publication seems of interest, however,
for a number of reasons. The various forms were found in such intimate admixture,
often in the same block of stone, that there can be little doubt that they give a fairly
representative picture of a closely contemporaneous population. Their age is Lower
Kimeridgian, Mutabilis Zone. The material is in rock-facies and only slightly crushed,
whereas the known exposures in the outcrop of the shales of the Mutabilis Zone
in this country yield, mostly, only wholly flattened shells, which have not attracted
collectors in the past. Our knowledge of the fauna of the Mutabilis Zone is therefore
scanty.

The fauna consists overwhelmingly of forms of the perisphinctid subfamily Aulaco-
stephaninae at the stage where Rasenia changes to Aiilacostephamis, and includes several
new and previously little-known forms of great interest. The classification of the genus
Rasenia is at present still in a state of confusion, but a revision of Aiilacostephamis has
been published by Ziegler (1962). New names will therefore be mostly confined here to
the forms being described.

In referring to specimens the following abbreviations will be used:

R, MS, SW : Thurrell collection, material from Ranby, Market Stainton, and South
Willingham (Lincolnshire) respectively.

BM : British Museum (Natural History).

SM : Sedgwick Museum, Cambridge.

We wish to thank Dr. R. G. Thurrell for making his material available to us for description;
Dr. B. Ziegler for discussions and criticisms, and for very kindly allowing us to see the manuscript
of his monograph on Aulacostephanus-, and Mr. A. G. Brighton (Sedgwick Museum), Dr. M. K.
Howarth (British Museum), and Mr. R. B. Wilson (Geological Survey, Edinburgh) for help with
material in collections in their custody.

^ This paper is based in part on a provisional manuscript prepared by Dr. Arkell shortly before his
illness in 1956. 1 am grateful to Mr. A. G. Brighton for passing the manuscript on to me for completion
after Dr. Arkell’s death in 1958. J. H. C.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 219-45, pis. 27-33.]
C 1213 Q

220

PALAEONTOLOGY, VOLUME 6

SYSTEMATIC DESCRIPTIONS

Family perisphinctidae Steinmann 1890
Subfamily aulacostephaninae Spath 1924 (= raseniinae Schindewolf 1925)

This subfamily encompasses a well-defined offshoot of the main perisphinetid tree.
It leads back to such typically perisphinetid forms as Pictonia in the basal Lower
Kimeridgian, with evolute coiling, round whorl-section, and smooth, regular ribbing
which passes over the venter without interruption. It terminates in end-forms of Aula-
costeplianus which are of quite different appearance, with compressed, angular whorl-
section, smooth tabulate or grooved venter, and strong straight ribbing reminiscent in style
of the Cretaceous genus Hoplites. Between these extremes, further wide variations in
coiling and novel forms of ornamentation occurred, so that the whole subfamily,
although in itself closely delimited, covers an unusually wide range of forms. Attempts
at classification have usually started from a broad division of the subfamily into two
parts: an earlier part grouped around the genus Rasenia, leading to the later forms
grouped around Aulacostephanus. Besides these, there has, however, sprung into exis-
tence a large number of other generic names created often on the basis of insufficient
material and stratigraphy, inadequate type specimens, and the requirements of phylo-
genetic theory. The resulting confusion cannot be fully analysed here, but, in order to
justify the classification which will be adopted in this paper, a number of points have
to be discussed.

{a) Generic classification. It seems sufficient to divide the subfamily into only two
genera: Rasenia and Aulacostephanus. Further grouping according to sculpture can be
conveniently incorporated at subgeneric level.

{b) Dimorphism. In common with apparently most other perisphinctids, the Aulaco-
stephaninae seem to have been dimorphic, and it is possible to classify most of the shells
as macroconchs or microconchs (for full discussion, see Callomon 1957, 1963). As else-
where, it seems helpful to incorporate this distinction at subgeneric level.

(c) The Rasenia-Aulacostephanus boundary. The transition between these genera is
so gradual that, in drawing a dividing-line, many difficulties arise. Development occurred
in more than one character of the sculpture, and the breaks in the development of
different characters occurred at different times. If the development of a single character
were made the sole criterion, an unnatural classification would result. Thus, for example,
the presence or absence of a ventral smooth band alone might seem a tempting dis-
tinction to use as criterion; but, as is in fact seen in the material being here described,
among a collection of shells which in most other respects are clearly conspecific, some
have a ventral smooth band, others not. Moreover, many have it at one stage of their
ontogeny, but not throughout. The best approach seems to be to group specimens of
a contemporaneous population according to species, taking all their characters into
account. A particular arrangement of species into genera is then of lesser importance.
The arrangement of borderline species used here has been chosen to accord fairly
closely with the one used by Dr. Ziegler in his monograph of the genus Aulaco-
stephanus, but no claims are made for its uniqueness. It serves also to bring out resem-
blances between the Lincolnshire material and the fauna of the Mutabilis Zone farther
south.

W. J. ARKELL AND J. H. CALLOMON; KIMERIDGIAN AMMONITES

221

Genus rasenia Salfeld 1913

Type species. Rasenia involuta (Salfeld MS) Spath 1935, designated by ICZN Opinion 426.

Microconchs. The following subgenera may be recognized, arranged in order of de-
creasing coarseness of ribbing.

1. PRORASENiA Schindewolf 1925. Type species: P. quenstedti Schindewolf, figured 1926,
pi. xix, fig. 1. (Synonym Desmosphinctes Schindewolf 1925, type species: Perisphinctes
mniovniketisis Nikitin 1884, p. 30, pi. ii, figs. 11, 12.) Small, strongly ribbed species with
lappets. Its special character is the presence of strong triplicate ribbing on the inner
whorls, reverting to biplicate perisphinctoid on the final body-chamber. Spath (1935,
pi. 14, 15) has figured and named three species from England, and Waterston (1951,
pi. 2) records them from Scotland.

2. RASENIA sensu stricto. Salfeld attached the manuscript name involuta to a number of
specimens in the British Museum, of which Spath selected and figured only one. As, how-
ever, Spath is formally the author of the species, this specimen is holotype of involuta
Spath rather than lectotype of involuta Salfeld, and both species and genus have to be in-
terpreted by Spath’s figured specimen. This is broken and slightly incomplete, and, in the
absence of the last few sutures and peristome, it is impossible to tell conclusively whether
the specimen is adult or not. All of the preserved last three-quarters whorl is, how-
ever, body-chamber, and it gives the impression near the end of slight contraction
indicating maturity. There exist in the collections other specimens (e.g. Geol. Survey Coll.
25550-1) which agree with the holotype closely in all respects including size, and which,
although also without peristomes, carry similar amounts of body-chamber. A com-
plete adult from Scotland has been figured by Waterston (1951, pi. ii, fig. 8a, b). Its final
diameter is 40 mm., and the body-chamber is terminated by a final constriction. The
peristome shows the stump of a lappet. It seems certain, therefore, that Rasenia s.s. was
fully grown at c. 40 mm. diameter with strong unmodified ribbing maintained to the end,
and hence microconch; and it seems probable that it carried at least short lappets.
Its special character is the differentiation of strong, sinuous primary and fine, fasciculate
secondary ribbing. Inner whorls are always well covered, involute and inflated.

In a recent revision of the genus Geyer (1961, p. 86), rather than re-examine the type
specimen of involuta Spath himself, quotes Spath’s statement describing it as ‘small,
inner whorls . . .’, which, whatever significance one may attach to size in taxonomy, it is
not; and interprets Rasenia sensu stricto as covering large, typically macroconch forms.
Rasenia s.s. is now capable of precise interpretation, and it is hoped that the confusion
of the past has found an end.

Geyer does point out that, if Rasenia sensu lato is taken to include Involutieeras
Salfeld as a subgenus (see below), involuta Spath 1935 is junior homonym of involuta
(Quenstedt 1849), and proposes Rasenia anglica nom. nov. = involuta Spath non
Quenstedt.

3. RASENioiDES Schindewolf 1925, p. 335. Type species by original designation: Nautilus
striolaris Reinecke 1818, which Schindewolf states should be interpreted by Am. striolaris
Quenstedt pars 1888, pi. 124, fig. 8, non Reinecke. Reinecke’s specimen seems to be lost,
but his figure represents recognizable characters which seem to differ considerably from

222

PALAEONTOLOGY, VOLUME 6

those shown by Quenstedt’s figure, notably in whorl-section and coiling. There is
nothing in Reinecke’s description to indicate a smooth venter, which, in this case, puts
Quenstedt’s specimen in Aulacostephamis', and, as small ‘ striolarian ’ Rasenids of the
thermarum group are common in the Mutabilis Zone (see below), Rasenioides fills a
useful role, interpreted here as microconch subgenus characterized by very fine, dense
fasciculate ribbing. Lappets proper are not known, although some specimens carry a
slightly raised sinuous collar on the mouth-border.

(pRORASENioiDES Schindewolf 1925, p. 338; type species by original designation:

P. transitorius Schindewolf, which is nom. nov. for Am. striolaris Quenstedt pars 1888,
pi. 107, figs. 12, 13, non Reinecke, with additional specimens figured by Schindewolf
1926, p. 507, pi. xix, figs. 3-5, resembles R. striolaris Reinecke more than Am. striolaris
Quenstedt 1888, pi. 124, fig. 8. However, as the latter is not strictly the type species of
Rasenioides whereas, according to the Rules, the former is, Prorasenioides and Raseni-
oides are here regarded as synonymous.)

Macroconchs

4. ZONOVIA Sasonov 1960. Type species by original designation : Amm. nralensis d’Orbigny
1845. Large, evolute, coarse-ribbed forms with smooth apertures.

5. EURASENIA Geycr 1961, p. 87, 90. Type species: Amm. Rolandi Oppel 1863. Large,
coarsely ribbed forms intermediate between Zonovia and Invohiticeras.

6. INVOLUTICERAS Salfeld 1913. Type species by monotypy (Salfeld 1914, p. 175) or
subsequent designation (Spath 1931): Amm. involutns Quenstedt 1849. Large, involute,
compressed forms, finely ribbed on inner whorls, becoming smooth.

7. ?SEMiRASENiA Geyer 1961, p. 87, 92. Type species: Amm. Mdsehi Oppel 1863. The f

holotype of mdsehi is 45 mm. in diameter, and if Oppel’s statement in the legend to his |'
figure, that the suture-line shown is the last, is to be believed, it carries three-quarters j
whorl body-chamber and is hence nearly complete. Geyer states the final size to be |

‘probably about 70 mm.’ without giving reasons based either on Oppel’s specimen, |

which he refigures but does not redescribe, or additional material, of which he cites |
only one additional, even smaller specimen. The peristome remains unknown, and I
maximum size and ontogeny of mature examples in doubt. |

The only other species included by Geyer in Semirasenia is Amm. thermarum Oppel. §
The holotype of this species seems to us to fit equally well into Rasenioides, however. ^

Subgenus rasenia s.s. Salfeld •'

1 . Rasenia (Rasenia) sp. ind. |

cf. Rasenia evoluta (Salfeld MS), Spath 1935, pp. 48-49, pi. 14, fig. 6a, b. /

Material. One (MS 128). h

Description. Diameter 17 mm., sutures invisible, but at least half the outer whorl appears ;
to be body-chamber. Coiling moderately evolute; ribbing straight, coarse and strong,
modifying on the last quarter-whorl to typically rasenid fasciculate style, passing un- '
interruptedly over the venter.

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 223

Spath assumed that R. evoliita was a large species (1935, p. 49, footnote). Like R.
involuta, however, he figured only a single specimen (BM C.8046 from Market Rasen),
which is therefore holotype. It is a wholly septate nucleus, maximum diameter only
24 mm., and on comparing the actual specimen with the type of R. involuta (which is
broken and shows the inner whorls well) at similar diameters, it is hard to detect even the
minutest differences between them. Whatever else Spath may have had in mind, R.
involuta and R. evoluta as defined by the monotypes are the same, and the latter name
may be dropped with little regret.

Subgenus rasenioides Schindewolf (— prorasenioides Schindewolf)

1 . Rasenia {Rasenioides) cf. transitoria (Schindewolf)

Plate 33, fig. 2

Ammonites striolaris Quenstedt 1888 pars, pi. 107, figs. 12, 13.

Prorasenioides transitoriiis Schindewolf 1925, p. 338.

Material. One (MS 80a).

Description. The specimen is crushed, but agrees well with Quenstedt’s figures. The
ribbing passes uninterruptedly over the venter where visible. Sutures and peristome are
missing, but at the maximum diameter of 32 mm. at least some of the outer whorl appears
to be body-chamber. The umbilical seam uncoils slightly near the end, indicating maturity.

2. Rasenia {Rasenioides) cf. and aff. lepidula (Oppel)

Plate 32, figs. 19-21

cf. Ammonites lepidulus Oppel 1863, p. 242, pi. 67, fig. Aa, b.

Material. Five, more or less fragmentary (MS 1, 33-35, 53).

Description and comparisons. Oppel’s figured type specimen was wholly inadequate to
form the basis of a new species, only the final quarter-whorl being recognizable. For-
tunately, this shows the specimen to have been a complete adult, and the name is used
here for forms of the R. thermarum group which are rather larger and more evolute
than R. thermarum, and in which the secondary ribs cross the round venter with perhaps
some weakening but without actual interruption.

Agreement is best between Oppel’s figure and the terminal fragment shown in Plate 32,
fig. 21. This shows the peristome: the aperture is sinuous, preceded by a broad, shallow
constriction and provided with a raised collar, as in R. thermarum, with no lappets;
whereas Oppel’s specimen was armed with lappets.

Oppel’s species is very close to Rasenia eulepida Schneid (see below). This, however,
has a prominent smooth band on the venter {teste Dr. Ziegler), but whether this serves
to distinguish it from R. lepidula it is impossible to say in the absence of more iiffor-
mation on the inner whorls of the latter. Schneid also stated that the distinguishing
feature of R. lepidula was the persistence of the characteristic sheaves of multiple secon-
daries arising from each primary rib right to the end, whereas in R. eulepida near the
aperture the ribbing reverts to simple bi- or triplication.

224

PALAEONTOLOGY, VOLUME 6

3. Rasenia {Rasenioides) thermarum (Oppel)

Plate 32, figs. 13-18

Ammonites thermarum Oppel 1863, p. 243, pi. 65, fig. Sa, b.

Rasenia (Semirasenia) thermarum Geyer 1961, p. 106, pi. 8, fig. 9, (? 10).

Materia!. Eight, and fragments (MS 9, 14, 31-32, 36, 64, 78, 118; R 20).

Description. All of the specimens seem to carry some body-chamber, and in one (PL 32,
fig. 14) the peristome is preserved on one side. The impression that they are fully grown
and nearly complete is supported by lack of remains of similar shells of larger am-
monites. The average size of the eight specimens is 19-5 mm. : Oppel’s holotype is 20 mm.
in diameter, with nearly complete body-chamber, and he says: ‘All the specimens
found so far possess a nearly complete body-chamber. The partly visible peristome
appears to be straight, without showing the formation of lappets.’ The peristome ob-
served here is also without lappet, but is sinuous and bears a slightly raised collar; it is
preceded by a broad shallow constriction.

The Lincolnshire material therefore agrees well with Oppel’s specimens, and differs
only in being slightly less inflated than the holotype. On some specimens the secondaries
weaken on the venter, which is round, but they are not fully interrupted. Geyer considers
R. thermarum to be a somewhat larger species, up to 40 mm. in diameter, effectively
reinterpreting it on the basis of a specimen previously figured by de Loriol (1878, pi. xiii,
fig. 5), which is variocostate like Aulacostephanus mdschi. The total material available
to him was only three specimens, however, including the holotype; and as there is no
reason to believe that the holotype is in any way imperfect, incomplete, or untypical,
but in fact representative of a readily recognizable group as shown by the Lincolnshire
material, we prefer to continue to interpret R. thermarum as closely defined by Oppel’s
type, and retain it in Rasenioides.

4. Rasenia {Rasenioides) cf. striolaris (Reinecke)

Plate 31, figs. 6a, b

Nautilus striolaris Reinecke 1818, p. 77, figs. 52, 53.

cf. Rasenia (Rasenioides) striolaris Geyer 1961, p. 10.

Material. Two (MS 29, figured; MS 141).

Description. The figured specimen is only 16-5 mm. in diameter but carries half a whorl
body-chamber. The umbilicus is only some 20 per cent, of the diameter, and inner
whorls are almost wholly concealed. The second specimen is crushed, but similar.

Reinecke did not state the magnification of his figure, and his specimen seems to be
lost. The present material agrees well with it, if we assume that his figure was about
natural size. Geyer has designated and figured a neotype (1961, pi. 21, fig. 3) without,
however, an adequate description. Thus, there is no mention of whether it, or supporting
material, carries any body-chamber; there is no ventral view; and no indication of the
form of the peristome. Moreover, the specimen differs considerably from Reinecke’s
figure in being much more evolute with very little inclusion of inner whorls, quite unlike
both Reinecke’s side-view and section.

W. J. ARKELL AND J. H. CALLOMON; KIMERIDGI AN AMMONITES

225

Subgenus zonovia Sasonov 1960
1. Rasenia (Zonovia) sp. ind.

Plate 33, fig. ha, b

Material. One fragment (R 2).

Description. The specimen consists only of a quarter of two contiguous wholly septate
whorls, corresponding to a diameter of about 45 mm. The whorl-section is round,
inflated, depressed ; the ribbing blunt, subdued, coarse. Short, straight bullate primaries
trifurcate into strong secondaries, which cross the venter with sight weakening but no
interruption.

Genus aulacostephanus Sutner and Pompeckj in Tornquist 1896

Type species. Ammonites miitabilis d’Orbigny non Sowerby = Ammonites pseudomutabilis de Loriol
1874, designated by ICZN Opinion 302. A lectotype from the d’Orbigny collection has been figured
by Ziegler (1962, pi. 15, fig. 8).

The boundary between Rasenia and Aulacostephanus is here being drawn such that
most forms assigned to the latter have a smooth band on the venter in at least one stage
of their ontogeny. Simultaneously, the whorl-section becomes more compressed and
quadrate, the venter tending to flatten ; and inner whorls are markedly more evolute and
serpenticone than in Rasenia so that the secondary ribs on them are often partly exposed.
The umbilici are in consequence also usually shallow, with often characteristically
smooth, gently sloping or even barely perceptible walls. These features together allow
a fairly sharp line to be drawn between the two genera.

The material from Lincolnshire falls into four subgenera: two pairs each consisting of
a microconch and a macroconch. One of these pairs of subgenera, Aulacostephanites and
Aulacostephanoides, differs but little from Rasenia and includes the forms well known
from the Mutabilis Zone elsewhere. The other pair of subgenera, Xenostephanoides and
Xenostephanus, is new. A few specimens have been known previously, but the presence
of a considerable number in the material under consideration gives it a peculiar stamp of
great interest.

Microconchs

1. aulacostephanites Ziegler 1962. Type species: Rasenia eulepida Schneid 1939, p. 146,
pi. V, figs. 13, 13n. Small, evolute forms with lappets; ornament similar to that of
Rasenioides. Smooth band on venter, at least on inner whorls. In crushed material from
the typical Mutabilis Zone shales in the south of England Rasenioides and Aulaco-
stephanites may be indistinguishable.

2. XENOSTEPHANOIDES subgen. nov. Type species: Aulacostephanus (Xenostephanoides)
thurrelli sp. nov. (see below). Small species (c. 40-50 mm.); coiling evolute, particularly
on inner whorls; whorl-section rounded quadrate. Ribbing strong; short bullate pri-
maries with sheaves of strong, straight secondaries often separated from the primaries
by a smooth band on the whorl-side. The style of the ribbing is already much more
like the ‘hoplitid’ of the later aulacostephanoids than the sinuous one of the rasenids.
On the final body-chamber it may revert to simple biplication, as in Prorasenia. The

226

PALAEONTOLOGY, VOLUME 6

venter carries a smooth band, sometimes accentuated by a groove. Peristomes with
oblique final constrictions and short lateral projections not really long enough to call
lappets.

Macroconchs

3. AULACOSTEPHANOiDES Schindewolf 1925. Type species, by original designation: Amm.
desmonotus Oppel. The type specimen of Oppel’s species (1863, p. 241, pi. 67, fig. la, b)
seems to have been lost during the war. His figure establishes the style of ribbing, coiling,
and umbilicus well, and shows clearly a smooth band on the venter. Oppel stated,
however, that no sutures were visible anywhere, and that the peristome was unknown ;
and it is therefore impossible to say whether the specimen (the only one known to Oppel)
was fully grown and complete, or merely the nucleus of a much larger form. This makes
the species a highly unsuitable one on which to found a genus, and interpretations have
been correspondingly varied. Dr. Ziegler has redescribed the species on the basis of
additional material. Among this is one specimen (Ziegler 1962, pi. 2, fig. 14) which
resembles the lost holotype extraordinarily closely, with unfortunately the same short-
comings. Others lead from this to more or less complete adults, which are large and
nearly smooth with simple peristomes. We follow this interpretation of A. desmonotus,
which is thus a macroconch.

The subgeneric characters of Aulacostephanoides are therefore: macroconch; rasenid
ornamentation on the inner whorls which fades, becoming smooth on the final body-
chamber; ventral smooth band on at least the inner whorls; simple peristomes. From
general experience involuteness of coiling seems of lesser importance, and Dr. Ziegler
shows that Aulacostephanoides can usefully accommodate a gradation of forms ranging
from the discoidal A. linealis (Quenstedt) to the true evolute A. rnutabUis (Sowerby).
Several species of the subgenus differ but little from forms still included in Rasenia, the
dividing-line from which is arbitrary, as discussed above.

4. XENOSTEPHANUS subgen. nov. Type species: Aidacostephanus (Xenostephanus) ranby-
ensis sp. nov. (see below). Large species (c. 120 mm.); coiling stout, depressed, and
evolute — particularly on the inner whorls. Primary and secondary ribbing well differen-
tiated on inner and middle whorls, the former tending to extreme bullae or tubercles on
the umbilical margin, the latter characteristically strong, triplicate, with intercalatories,
fusing into the primaries or sometimes separated from them by a narrow smooth band
on the whorl sides. Venter flat on inner whorls, with smooth band, sometimes accen-
tuated by a groove. On the body-chamber the ribbing becomes subdued and degenerates
to more perisphinctid form: primary ribs biplicate smoothly or become merely single
widely spaced ridges. The whorl-section becomes round, and ribbing passes over the
venter without interruption. Peristomes simple.

Inner and intermediate whorls in some species are almost fully homeomorphic with
the quite unrelated earlier genus Reineckeia; outer whorls are as in Rasenia (Zonovia),
especially R. uraJensis (d’Orbigny).

Xenostephanus and Xenostephanoides form 2LlypicdX macroconch and microconch pair,
and it would be difficult to find a clearer demonstration of the dimorphism among the
perisphinctids. The inner whorls are so similar that it would be hard to sort them out
in immature, incomplete, or fragmentary material.

W. J. ARKELL AND J. H. CALLOMON; KIMERIDGIAN AMMONITES

227

Subgenus aulacostephanites Ziegler 1962
1. Aulacostephanus {Aulacoslephanites) cf. eidepidus (Schneid)

Plate 32, figs. 1-8

Rasenia eulepida Schneid 1939, p. 146, pi. v, figs. 13, 13o.

Aulacostephanus {Aulacostephanites) eidepidus Ziegler 1962, p. 44, pi. 341, figs. 2-7, 8 partini,
? 11, 14.

Material. Twenty-five (24 from MS; and R 5), and fragments.

Description and remarks. The Lincolnshire material seems to be consistently somewhat
smaller than the type (33 mm.), for the average diameter of seventeen complete adults is
25 mm. (standard deviation 3-9 mm. = 15-5 per cent.). The style of ribbing, however,
is the same, including the degeneration of the ribbing near the end, which Schneid cited
as diagnostic compared with Am. lepididus Oppel. Peristomes are preserved in several
specimens. There is usually a slight terminal constriction and a raised mouth-border.
Lateral extensions may be short (PI. 32, figs. 3, 5), or long enough to qualify as lappets
(figs. 2, 10). Smooth bands on the venters are generally present, but narrow and incon-
spicuous, and they die out on the last half-whorl of body-chamber.

The question of the generic position of the specimens here described raises in its most
acute form the difficulty attending attempts to divide Aulacostephanus from Rasenia.
The ventral smooth band is undoubtedly much less prominent here than in the type of
A. eidepidus and other south German material, and the Lincolnshire forms might, on this
account, rest more comfortably in Rasenia (Rasenioides), with possibly gradations to
R. thermarum (Oppel). On the other hand, the style of coiling and ribbing resembles A.
eidepidus much more closely than R. thermarum or any other Rasenioides so far described
(with the possible exception of R. lepidida (Oppel), which we exclude from consideration
because of the imperfection of the type); and there is gradation to the forms described as
A. ebrayoides below, which have quite prominent smooth bands on the venter.

It seems therefore that we are here sampling the Rasenia-Aulacostephanus boundary
at a slightly dilferent stage from that observed in central Europe. There the distinction
between Rasenioides thermarum and Aulacostephanus eidepidus is clear, and the latter
grades into more coarsely ribbed forms which nevertheless retain the rasenid (fasciculate)
style of secondary ribbing, and which Ziegler designates A. peregrinus nov. In contrast,
these forms appear to be absent in the Lincolnshire material, which, instead, shows more
or less gradation from A. ebrayoides (unknown in central Europe), via A. eidepidus, to
R. thermarum. Whether these differences reflect slight differences in ages, or ecological
factors (geographic subspeciation), it is at present hard to say.

A. eulepidus is also found in the Mutabilis Zone of Mull (cf. PI. 33, figs. 4-6).

2. Aulacostephanus {Aulacostephanites) ebrayoides sp. nov.

Plate 30, figs. 14-18

Material. Nine (eight from MS, and R 16), and fragments.

Description. The holotype, MS 5 (PI. 30, fig. 14) is 26 mm. in diameter, with half-whorl
body-chamber preserved. The remaining material is of similar size. Coiling is evolute,
the whorl-section quadrate. Ribbing is dense, but strong and sharp, straight, triplicate

228

PALAEONTOLOGY, VOLUME 6

or biplicate with intercalatories ; the furcation point is very low on the whorl-side, so that
the primaries (about twenty-two on the outer whorl) are reduced to short ridges on the
umbilical margins. There is a distinct smooth band on the venter on inner whorls, fading
somewhat on the body-chamber.

Comparisons. The species differs from A. eulepidus through the reduced contrast between
primary and secondary ribbing, and the straightness and coarseness of the latter. Its
name reflects the close homeomorphism with the Bathonian genus Ebrayiceras Buckman
(Morphoceratidae), e.g. E. pseudo-anceps (Ebray) (see Arkell 1955, pi. xvii, figs. 7-11).

There are occasional specimens with more flexuous, but still coarse, secondary ribbing
(PI. 32, figs. 9, 10, with short lappets), which may link A. ebrayoides with A. eulepidus.
These coarse forms of Aulacostephanites occur also in Mull (PI. 33, fig. 7).

Quenstedt’s Amm. cf. striolaris pars, 1888, pi. 107, fig. 18 seems to resemble A.
ebrayoides closely in its ribbing, but differs in having a much more prominent ventral
smooth band, and a deep terminal constriction (which may, however, have only varietal
significance.)

3. Aidacostephanus {Aulacostephanites) sp. aff. desmonotus (Oppel)

Plate 32, figs. 11,12

cf. Ammonites desmonotus Oppel 1863, p. 241, pi. 67, figs. \a, b.

Material. Three (MS 76, 1 15; R 17).

Description and comparisons. All three specimens attain between 20 and 23 mm. and
appear to carry some body-chamber. They resemble the other specimens described here
under Rasenia thermarum in ornament, but are distinctly more compressed, have promi-
nent ventral smooth bands and the characteristic shallow umbilicus of Oppel’s Amm.
desmonotus. The latter, however, probably attained a large size (see discussion above).

4. Aidacostephanus {Aulacostephanites) sp. aff. mdschi (Oppel)

Plate 32, figs. 22-24

cf. Ammonites mdschi Oppel 1863, p. 240, pi. 65, fig. 2a, b.

Ammonites cf. striolaris Quenstedt pars 1888, p. 969, pi. 107, fig. 17.

Material. Eleven (MS, ten; R 1), and fragments (MS).

Description and comparisons. These specimens are characterized by the absence of primary
ribbing on the inner whorls, which carry only very dense, fine secondary ribbing. The
whorl-section is compressed, umbilicus shallow, and there is a ventral smooth band.
Agreement with Oppel’s figure is fairly close, except that A. mdschi shows no trace of
a ventral smooth band, is bigger, and has the primary ribs, when they appear, lower on
the whorl-side. The best agreement is with Quenstedt’s specimen.

EXPLANATION OF PLATE 27

Fig. \a-c, Aidacostephanus (Xenostephanus) ranbyensis sp. nov. var. elshamensis var. nov. Adult, with
half a whorl body-chamber. BM C.47908, T. B. Parks coll., from Elsham, Lines. All natural size.

Palaeontology, Vol. 6

PLATE 27

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES

229

Subgenus aulacostephanoides Schindewolf, emend. Ziegler
1. AuJacostephanus {Aulacostephanoides) mutabilis (J. de C. Sowerby)

Plate 31, fig. 3

Ammonites mutabilis J. de C. Sowerby 1823, 4, p. 145, pi. 405, fig. 1.

Rasenia mutabilis Salfeld 1913, p. 129.

Pararaseuia mutabilis Arkell 1933, p. 612, pi. xxxix, figs. 5, 5a (holotype refigured).

Rasenia mutabilis Arkell 1951, p. 188.

Aulacostephauus {Aulacostephanoides) mutabilis Ziegler 1962, p. 62, figures and plates.
Material. One (MS 80).

Description. The specimen, half an ammonite, is somewhat crushed; maximum diameter
175 mm., wholly septate, the last few septa approximated and degenerate. The complete
shell must have attained some 210 mm. Inner whorls are evolute; the whorl-section is
at all stages compressed with barely perceptible umbilical walls. Well-differentiated, fine,
dense, secondary ribbing is visible to at least 60 mm. with occasional feeble constrictions
and flared ribs; primary ribs persist to 90 mm. Thereafter the shell is wholly smooth.
Venter on inner whorls not visible.

Remarks. There has been confusion in the past in interpreting this species, largely as a
result of Sowerby’s poor illustration. The holotype (BM 43934) has a maximum diameter
of 71 mm. and is wholly septate, with none of the degeneration and approximation of the
last sutures rather typical of adults of this species. Despite the sudden end of primary
ribbing at a shallow constriction at 52 mm., it seems certain that the holotype is the inner
part of what was a much larger shell. The secondary ribbing on the venter is interrupted
by a distinct smooth band. The whorl-section is compressed and angular. All these points
are exemplified in another specimen figured here (PI. 31, fig. 3, from Cambridgeshire)
which resembles the holotype very closely, and is also wholly septate to 105 mm. It also
shows something of the inner whorls, invisible in the holotype, which are evolute,
exposed, and densely ribbed.

As for the generic position of the species, the style of ribbing on middle whorls is that
of Rasenia; but the compressed whorl-section, evolute inner whorls, shallow umbilical
walls, and ventral smooth band place it firmly in Aulaeostephanus. [This view is contrary
to that expressed previously by Dr. Arkell (1951, p. 188); but on re-examining both the
holotype and supporting material, and in the light of Dr. Ziegler’s study of the whole
genus Aulaeostephanus it seems the only possible one. J. H. C.]

2. Aulaeostephanus {Aulaeostephanoides) sp. nov.?, aff. mutabilis (Sowerby)

Plate 31, figs. 1, 2

Material. Two, and fragments (MS 37, 102; R 5).

Description. This species resembles A. mutabilis in all respects, but seems to be much
smaller. Thus the specimen figured in Plate 31, fig. 1 is apparently adult, and complete
with peristome at the same size (110 mm.) at which A. mutabilis itself (PI. 31, fig. 3) is
still septate. The same specimen is rather more evolute than A. mutabilis, and all ribbing
has disappeared at 60 mm. The specimen shown in Plate 31, fig. 2 has inner whorls
which are evolute and densely ribbed as in ^1. mutabilis; likewise the imprint of the flat

230

PALAEONTOLOGY, VOLUME 6

venter of the penultimate whorls in the outer (body-chamber) whorls shows the secon-
dary ribbing to be dense, with occasional feeble constrictions and slightly flared ribs,
and interrupted by a smooth band.

Similarly, small forms (for macroconchs) are known from the Kimeridge Clay of
Cromarty (Scotland). They are to be described by Dr. Ziegler, who proposes the name
R. askepta for them. They differ from those described here, however, in having oval
whorl-section, involute inner whorls, and secondary ribbing uninterrupted on the venter,
i.e. belong still to Raseuia. Their precise age is not known.

3. Aiilacostephanus (Aulacostephanoides) circumplicatus (Quenstedt)

Plate 31, figs. 4, 5; Plate 33, figs. 9-11

Ammonites circumplicatus Quenstedt 1888, p. 978, pi. 107, fig. 19.

Auuuouites cf. trifurcatus Quenstedt 1888, p. 971, pi. 107, fig. 21.

Tobolia pseudotrifurcata Sasonov 1960, p. 156, pi. v, fig. 4 (= nom. nov. for, and reproduction
of original figure of Amm. cf. trifurcatus Quenstedt, cited above).

Material. Qne, and two fragments (MS 51, 120; R 19).

Description. The specimen shown in Plate 31, fig. 4 is septate to only 55 mm., and if adult,
could not have exceeded 80 mm. complete. The whorl-section is slightly compressed, and
in both figured specimens the secondary ribbing is weakened over the venter, but not
actually interrupted.

Remarks and comparisons. The present material agrees with A. circumplicatus in size and
style of ribbing; but the holotype has an exceptionally prominent and broad smooth
ventral band. This is, however, in part due to a deformity, and, according to Dr. Ziegler,
the ventral smooth band is unusually variable in the species, some specimens being
practically without.

There is a considerable resemblance to Rasenia gemmata Schneid (1939, p. 147, pi. v
(ix), figs. 9, 9a), which, however, differs in being inflated, with depressed whorl-section,
and is more involute.

The species is common in the Kimeridgian exposure on Mull (PI. 33, figs. 9-11), and
some of the specimens have a clearly defined ventral smooth band.

Subgenus xenostephanoides nov.

1. Aidacostephanus {Xenostephanoides) thurrelli sp. nov.

Plate 30, figs. 1-3

Material. Eight, and fragments (R, five; MS, two; SW, eleven).

Description. The holotype (R 1 1 ; PI. 30, fig. 1) is 44 mm. in diameter; septate to 40 mm..

EXPLANATION OF PLATE 28

Fig. \a-e, Aidacostephanus (Xeuostephamis) ranbyensis sp. nov. Holotype; adult, with five-eighth whorl
body-chamber preserved (lu). Suture-line at ^ shown in fig. le, X 2. Thurrell coll. R 6, from Ranby,
Lines.

Fig. 2, Aidacostephanus {Xenostephauus) cf. ranbyensis sp. nov. Body-chamber fragment, showing the
peristome. Thurrell coll. SW 23, from South Willingham, Lines.

All natural size, except fig. \e.

Palaeontology, Vol. 6

PLATE 28

S

PFPp!

WK

le

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 231

with a quarter-whorl body-chamber preserved. The last three septa are approximated,
with some degeneration. The coiling is extremely evolute, and the dimensions (between
and through ribs or tubercles) at 45 mm. are: 24-5-26-5, 25-5-31, 49. Primary ribs are in
the form of tubercles on the umbilical margin (20 per whorl at 45 mm. ; 21 at 30 mm.);
secondaries are trifid or bifid with frequent intercalatories, separated from the primaries
by a nearly smooth lateral spiral band. On the last half-whorl, the secondaries revert to
widely spaced, very coarse bifid without intercalatories indicating, together with septal
approximation, that the specimen was adult. The secondary ribbing is extremely strong
and coarse, resembling that of A. imdorae (Pavlow) of the Eudoxus Zone, but is quite
different on the venter. In A. ihurrelli it is interrupted by a deep narrow smooth band,
accentuated by a groove, with no tendency to form ventro-lateral tubercles.

The paratype R 25 (PI. 30, fig. 2) resembles the holotype closely. It is septate to only
34 mm., but shows the complete body-chamber of five-eighth whorl with peristome at
50 mm. The ribbing reverts to strictly biplicate on the last quarter whorl; an oblique
constriction follows an isolated triplicate rib. The lip of the aperture is flared into a collar
on the venter, and has a slight lateral projection too short to be called a lappet proper.

Presumed inner whorls are shown in Plate 30, fig. 3, although no septa are visible.

The species or a close ally seems to be present in the Kimeridgian of Mull (cf. PI. 33,
fig. 8) to judge by inner whorls; but the imprint of the outer whorls is too badly dis-
torted to be certain. There is also a specimen from the Cromarty coast (Royal Scottish
Museum, Edinburgh, Hugh Miller coll.).

2. Aulacostephanus {Xenostephanoides) lindensis sp. nov.

Plate 30, figs. 4-13

Derivation. Lindum is the Roman name for Lincoln.

Material. Twenty-seven (SW, twenty-one; R, three; MS, four), and fragments.

Description. The material is plentiful but poorly preserved, and no single specimen is
complete. It is with some hesitation, therefore, that we select the specimen MS 13 (PI. 30,
fig. 10) only formally as holotype and designate the other nine figured specimens (PI. 30,
figs. 4-9, 11-13) paratypes, although for purposes of description it would be best to
regard all ten as syntypes.

Sutures are rarely visible, but from the difference in preservation between outer and
inner whorls (which are usually flattened) it seems probable that most of the specimens
carry some body-chamber. The holotype itself definitely does, and was presumably
complete at about 30 mm. diameter. Another fragment (SW 59) shows the final peris-
tome with oblique constriction, flared collar, and short incipient lappet (just as in A.
thurrelli) at a final diameter of c. 40 mm. The species is thus small.

The coiling is moderately involute, the whorl-section stouter than in A. thurrelli.
Ribbing is dense and moderately strong, with three, sometimes four, secondaries per
primary, separated, as in A. thurrelli, by a lateral smooth band. The venter is flat, broad,
with a narrow smooth band but no groove. Inner whorls are shown in Plate 30, fig. 13.

Comparison. The species most closely resembles Aulacostephanus thurrelli although the
stouter, more involute coiling and finer ribbing appear to be consistently distinct. It
stands, like A. thurrelli, apart from all other contemporaneous aulacostephanids ; but,

232 PALAEONTOLOGY, VOLUME 6

in its resemblance to some later forms of the A. eudoxus group, it heralds the future
development of the genus.

A. lindensis has also been found in the drift of Norfolk (SM J48657, one specimen,
Baden-Powell coll.).

For completeness it is convenient to mention here another form which belongs to this
group.

3. Aidacostephauus {Xenoslephanoides) scoticus nom. nov. = Aidacostephanus cf. autissi-
odoremis (Cotteau) Spath 1935, p. 49, pi. 13, figs. 8a, b (BM C13216)

Remarks. The specimen figured by Spath from Culgower, eastern Scotland, seems
unique. It carries some body-chamber on which the otherwise trifid ribbing reverts to
biplication, indicating maturity, as in /I. thurrelli; A. scoticus is therefore a small species
and not a young specimen of giant forms like Rasenia borealis Spath, as Spath sup-
posed. Its evoluteness and style of ribbing, particularly on the inner whorls, are typical
of Xenoslephanoides. It differs from A. {X.) thurrelli principally in having rather more
inffated inner whorls, about nine more ribs per whorl, and a less-pronounced ventral
smooth band, the ventral ribbing on the inner whorls being weakened without actual
interruption. The exact horizon is unknown, although there are other Mutabilis Zone
faunas from the same area.

Subgenus xenostephanus nov.

1 . Aidacostephanus {Xenostephanus) ranbyensis sp. nov.

Plate 28, figs. \a~e, 2; Plate 29, fig. 4
Material. Two (R 6; BM C.47909), and fragments (SW).

Description. The holotype (R 6) is a splendidly preserved internal cast of 120 mm.
maximum diameter, septate to 81 mm., with the last three sutures approximated and
simplified. The body-chamber is broken, but, to judge from the highly degenerate last
few visible ribs, represented very nearly up to the peristome and hence occupying five-
eighths of a whorl.

Dimensions: at 115 mm.: 23, c. 26, 48.

80 mm. : 25-27, 30-33, 54.

55 mm.: 25-27, 29-31, 55.

Ribs: at 40-50 mm.: 19 per whorl; 70-90 mm.: 22; 115 mm.: c. 25.

EXPLANATION OF PLATE 29

Fig. \a, b, Aulacostephamis {Xenostephanus) anceps sp. nov. Holotype; adult, with one-quarter whorl
body-chamber preserved, and umbilical suture indicating former extent to five-eighths of a whorl.
BM 50761, old collection, no history.

Fig. la, b, Aidacostephanus {Xenostephanus) staintonensis sp. nov. Syntype I; the cross marks what
appears to be the last septum. Thurrell coll. MS 142, from Market Stainton, Lines.

Fig. 3, Aidacostephanus (Xenostephanus) staintonensis sp. nov. Syntype II; wholly septate, last few
septa approximated. SW 2/35, from South Willingham, Lines.

Fig. 4, Aidacostephanus {Xenostephanus) cf. ranbyensis sp. nov. Fragment at commencement of the
body-chamber. SW 34.

All natural size.

Palaeontology, VoL 6

PLATE 29

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 233

In the umbilicus nothing is visible before a diameter of 20 mm. Thereafter the primary
ribbing consists of conspicuous dorsolateral tubercles, only slightly bullate, and separ-
ated from the strong rectiradiate ribs that spring from them by a lateral smooth band.
The ribs are bifid with regular intercalatories, which, on the last septate whorl, tend to
join up to form trifid bundles, while fresh intercalatories come in. On the body-chamber
the ribbing weakens, the primaries degenerate to bullae and triplicate, finally biplicate,
ribs. The last few ribs are single, coarse and irregular, indicating the end of growth. The
peristome is not preserved. The venter, where it emerges, has a groove, but no tubercles.
The groove fades to a mere smooth band before septation ceases, this coming about
as the ribbing loses relief, and persists to the end even when the ventral ribbing is
barely perceptible. The whorl-section on inner whorls is quadrate, angular, and
becomes rounded towards the end. The suture-line is a typically perisphinctid one, with
slender widely spaced lobes on a straight base-line.

The peristome is preserved on another fragment (PI. 28, fig. 2). It is straight and simple,
with slight forward sweep at the umbilical seam.

There is another specimen in the British Museum (C47909, T. B. Parks coll., from
Elsham, Lines., only 72 mm. in diameter, wholly septate, somewhat crushed, which
agrees with the holotype in all details, including matrix. The species has also been found
in the drift of Norfolk (SM J48657, Baden-Powell coll., one specimen).

Remarks. Inner whorls are practically indistinguishable from A. (X.) thwrelli, and, as
remarked previously, it would be hard to find a clearer demonstration of ammonoid
dimorphism. Also remarkable is the close homeomorphism between Xenostephanus
and Reineckeia s.s. of the Callovian, yet another instance showing the tendency of
ammonites to evolve repeatedly through similar forms.

la. A. (A.) ranbyensis var. elshamensis var. nov.

Plate 27, fig. \a-c

Material. One (BM C.47908, T. B. Parks coll., from Elsham, N. Lines.).

Description. Size c. 130 mm. (slight distortion), septate to 103 mm. with approximation
and degeneration, half a whorl of body-chamber preserved with indications of former
extent to three-quarters of a whorl, so that the maximum size of the complete shell must
have been 150 mm.

Dimensions: at 100 mm.: 25, 29-34, 57.

Ribs: at 130 mm.: 25 per whorl; 120: 23; 100: 22; 80: 20; 60: 19; 40: 19; 30: 20.

The primary ribs are bullate, triplicate with low furcation point up to 80 mm. ; they then
revert to biplicate, perisphinctid ridges to 120 mm., and single simple ribs thereafter.
The secondaries are strong, in regular triplicate sheaves, to about 80 mm., then rapidly
weaken and become biplicate at high furcation points, and finally fade altogether. They
show some weakening in the middle of the venter where first visible, although there is no
fully smooth band, and after about 90 mm. pass over the venter with no interruption.

Comparison. Agreement with the holotype of A. ranbyensis is close in proportions and
rib-count. The variety differs in becoming bigger, with a more inffated body-chamber.

234

PALAEONTOLOGY, VOLUME 6

and having less differentiation between primary and secondary ribbing so that a lateral
smooth band is present only on the innermost whorls.

2. Aulacostephanus {Xenostephanus) staintonensis sp. nov.

Plate 29, figs. 2, 3

Material. Two (MS 142, SW 2), and fragments (SW).

Description. Neither specimen is really sufficiently complete to serve as sole basis of the
species, and the two figured specimens are syntypes.

Syntype I (PI. 29, fig. 2): maximum diameter 57 mm., apparently septate to only
35 mm. with live-eighths whorl body-chamber. Neither ribbing nor coiling, where last
visible, show any signs of modification whatever, and the specimen seems to be genuinely
immature.

Dimensions: at 55 mm.: 32, c. 33, 45.

Ribs: at 60 mm.: 22 per whorl; 50: 21 ; 40-20: 20.

Inner whorls evolute, becoming more inflated. Primary ribs moderately bullate, trifid
on inner whorls, changing to regular triplicate. Strong secondaries with ventral smooth
band, fading at the end.

Syntype II (PI. 29, fig. 3): maximum diameter 115 mm., wholly septate (details of
sutures invisible), flattened through crushing.

Ribs: at 115 mm.: c. 30 per whorl; 70: c. 27 (counts based on half-whorls).

Ribs on inner whorls triplicate or biplicate with intercalatories ; sharp bullate pri-
maries, no lateral smooth band; ventral smooth band. Ribbing has started to modify a
90 mm. to perisphinctid style; and at 115 mm. only straight primaries of low relief
remain. The ventral smooth band persists until the entire venter is smooth.

Comparisons. The species differs from A. ranbyensis in being markedly more densely
ribbed and involute. The primary and secondary ribbing are less well differentiated,
so that there is no lateral smooth band even on inner whorls, which are less openly
exposed. In this sense the species stands in the same relation to A. ranbyensis as A. lindensis

EXPLANATION OF PLATE 30

Figs, f-3, Aulacostephanus {Xenostephanoides) thurrelli sp. nov. \a, b, Holotype; adult, with one-
quarter whorl body-chamber preserved. Thurrell coll. R 1 1, from Ranby, Lines. 2a, b. Fragmentary
but complete adult, with peristome showing slight lappet. R 19/25. 3a, b. Inner whorls, no sutures
visible. SW 39.

Figs. 4-13, Aulacostephanus {Xenostephanoides) lindensis sp. nov. 4, SW 31, no sutures visible but
probably one-half whorl body-chamber. 5, SW 29, last third whorl body-chamber. 6a, b, SW 33,
no sutures visible, but probably at least one-half whorl body-chamber. 7, SW 26, ditto. 8a, b, SW 60,
ditto. 9a, b, SW 9, ditto. 10a, b, Holotype, MS 13, seven-tenths whorl body-chamber. 11, SW 53,
remarks as 6. 12a, MS 38, ditto. 13a, 6, MS 18, wholly septate. All Thurrell coll.

Figs. 14-18, Aulacostephanus (Aulacostephanites) ebrayoides sp. nov. 14, Holotype; last half whorl
body-chamber; MS 5. 15, MS 56, half whorl body-chamber. 16a-c, MS 57, ditto. 17a-c, MS 45,
eight-tenths whorl body-chamber. 18a-c, MS 71, six-tenths whorl body-chamber; perhaps transi-
tional to A. (A.) eulepidus (Schneid). All Thurrell coll.

All natural size.

Palaeontology, Vol. 6

PLATE 30

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 235

stands to A. thiirrelli. An example of this species has also been found in the drift of
Norfolk (SM J48658, a double mould, Baden-Powell coll.)-

3. Aulaco Stephanas {Xeno Stephanas) aneeps sp. nov.

Plate 29, fig. 1

Material. One (the holotype, BM no. 50761).

Description. Maximum size 95 mm., septate to 85 mm. with severe approximation and
degeneration of the last (and only visible) sutures, and a quarter whorl body-chamber.
The umbilical suture indicates a former extent of the final body-chamber to five-eighths
whorl, so that the complete shell attained a maximum diameter of c. 110 mm.

Dimensions; at 90 mm.; 27, 29-33, 51 (cast).

at 75 mm. ; 27, 36-43, 52 (test).

Ribs; at 100-80 mm.; 21 per whorl; 60-50; 20; 40; 19; 30-20; 18.

This is the most inflated of the species described here, with a whorl-section at 75 mm.
depressed almost to the point of being coronate. In contrast, the body-chamber contracts
and reverts to normal perisphinctid dimensions. Similarly, the ribbing, which is charac-
terized by very high, sharp bullate primaries on the umbilico-lateral margin giving rise to
regular sheaves of very coarse triplicate secondaries, modifies to rather widely spaced
and in part biplicate ribs of low relief on the body-chamber. The secondaries show only
the faintest weakening on the round venter where first visible (at 60 mm.); they smooth
out altogether on the body-chamber.

Affinities and remarks. A. aneeps bears undeniably close resemblance to various species
of Rasenia from the Cymodoce Zone of Market Rasen, including R. aralensis (d’Orb.),
and, in the absence of a ventral smooth band, to place it in any but this genus might
seem unnatural. Its inner whorls, however, are quite different: evolute, with flat whorl
sides and the differentiation, with lateral smooth band, of primary and secondary ribbing
characteristic of Xeno Stephanas. The aulacostephanid characters of angular whorl-section
and ventral smooth band seem here to make their first appearance in Xenostephanas
proterogenetically.

The specimen has the test preserved on all but the last half whorl, and it is therefore
conceivable that an internal cast might show a ventral smooth band not visible on the
outside; and that the difference in this respect between A. {X) aneeps and the other
forms of Xenostephanas described here, mostly casts, might only be apparent and attribu-
table to differences of preservation. This seems unlikely, however, for a specimen of
A. (X.) ranbyensis from Norfolk (SM J48657) shows the same ventral smooth band on
both internal cast and external mould of the same part of the body-chamber at 60 mm.
diameter.

The holotype was presented to the British Museum by Prof. J. Morris in 1867 ; there is
no other recorded history. Its preservation and matrix are, however, so precisely similar
to the other material here described that there can be little doubt that it came from a
similar source. It seems to have been one of the specimens which Salfeld included in his
MS species evolata; but its inner whorls are quite different from those of the specimen
Spath selected as type for his species (see above). Besides oysters, the matrix carries an
imprint of Rasenia cf. t her mar am (Oppel).

C 1213

R

236

PALAEONTOLOGY, VOLUME 6

Xenostephanus seems to occur in the Kimeridgian of Skye. A single specimen of A.
(X.) cf. or aff. anceps (PI. 33, fig. 1) was collected by Mr. W. G. Cordey at Kildorais,
Trotternish, in the spring of 1961, and we thank him for presenting it to us. It is crushed,
in clay, and encrusted with an oyster on the inner whorls. Neither septa nor venter are
discernible, but the ornament is typically that of Xenostephanus, and enough is visible
of the inner whorls to show that they are evolute just as in A. anceps. The Skye specimen
differs in having rather fewer ribs on the middle and outer whorls; at 85 mm.: 17 per
whorl; 70: 16; 50: 15; 30: c. 18. The specimen was unfortunately found without any
supporting fauna.

For completeness, we include here:

4. Aulacostephanus (? Xenostephanus) sp. indet. = Ammonites cf. trifurcatus Quenstedt

1888, p. 971, pi. 107, fig. 24

Remarks'. Quenstedt’s figure shows an extremely evolute shell, maximum size c. 85 mm.,
with ornament typical of Xenostephanus'. well-spaced bullate primary ribs leading to
triplicate, strong secondaries with intercalatories. Unfortunately neither venter nor
sutures are anywhere visible. At 85 mm. there are 18 ribs per whorl; at 60: 17; at 40: 15.
The species differs from A. ranbyensis therefore in being less densely ribbed; also, in
having the primaries half-way up the whorl-side rather than on the umbilico-lateral
margin.

Nautilus trifurcatus Reinecke (1818, p. 75, pi. v, fig. 49) is quite different: inflated,
involute, with barely bullate primary ribbing. Spath (1935, p. 49, footnote) thought
Quenstedt’s specimen belonged to Rasenia evoluta Salfeld MS, and in the sense that the
specimen described above as A. {X.) anceps appears to have been a syntype of Salfeld’s
unpublished manuscript species he was probably correct. It is unfortunate that he
chose to publish the name on the basis of only a small nucleus (see above).

If the above identification is correct, Quenstedt’s specimen seems to be the only
representative of these early aulacostephanitids so far known from more southerly
regions. It was found in a volcanic erratic at Floriansberg, and ascribed to White Jura §.

There are a number of other boreal forms intermediate between Rasenia and Aulaco-
stephanus proper which, when more is known about them, may turn out to be closely
related to Xenostephanus.

Aulacostephanus groenlandicus Ravn (1912, p. 492, pi. xxxvii, fig. 3a-c, from Koldewey
Island, 76° N.; Frebold 1930, p. ix, fig. 4, from Spitsbergen) has the evolute coiling and
indications of a ventral smooth band, but is densely ribbed and lacks the tuberculate

EXPLANATION OF PLATE 31

Figs. 1 , 2, Aulacostephanus (Aulacostephanoides) sp. nov. ? aff. mutabilis (J. de C. Sowerby). 1 , Complete
adult, with seven-tenths whorl body-chamber, and peristome transposed unto the figured side (white
dotted line) as it is developed on the reverse side. Thurrell coll. MS 37. 2, MS 102; the outer whorl
is body-chamber.

Fig. 3, Aulacostephanus (Aulacostephanoides) mutabilis (J. de C. Sowerby). Wholly septate example,
from a nodule-bed in Kimeridge Clay near Haddenham Station, Cambridgeshire. SM J.29157.

Figs. 4, 5, Aulacostephanus (Aulacostephanoides) circumplicatus (Quenstedt). 4, MS 120, with some
body-chamber. 5a, b, MS 51b, outer whorl body-chamber.

Fig. 6a, b, Rasenia (Rasenioides) cf. striolaris (Reinecke). MS 29, with half whorl body-chamber.

All natural size.

Palaeontology, Vol. 6

PLATE 31

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 237

primaries. Precise age unknown, but Kimeridgian of mutabilis age is known to be present
in east Greenland (J. H. C. 1957).

Aidacoslephanus Pischmae Khudyaev 1932 (p. 646, text-fig. 2, pi. 1 fig. 1, from
Petchoraland) seems to be very close to Xenostephanus, if not actually to belong to it.
Khudyaev’s text-fig. 2 shows the whorl-section of inner whorls to be depressed with
tuberculate or bullate ribbing of high relief, as in A. (X.) anceps. Unfortunately the plate
shows only the outer whorl — body-chamber — which has perisphinctid bi- and triplicate
ribbing with subtriangular, compressed whorl-section, and, in the absence of better
material, affinity yNdh Xenostephanus must remain in doubt. A. pischmae seems equally
close to some of the large, coarsely, but bluntly, ribbed forms like A. quenstedti Durand,
which Dr. Ziegler places in Pararasenia Spath.

A. pischmae is type species by monotypy of the genus Sarygidia Khudyaev, proposed
in the legend of text-fig. 2 (p. 647) only. The type species comes from the Timan, Petch-
ora, north Russia, whereas Sary-goul (Saragula) is in the southern Urals, where, to judge
by Pavlow’s plates (1886), the group does not occur. Possibly Khudyaev withdrew the
name on account of this but forgot to remove it also from the underline of the text-figure.
The original name is invalid, having been proposed after 1931 without diagnosis or
citation of a type species. It has been revived by Sasonov (1960, p. 157), who has formally
designated A. pischmae as type species. He figures no additional material, however, and
so Sarygidia Sasonov 1960, though valid, will continue to be regarded here as synony-
mous with Pararasenia Spath, emend. Ziegler.

Pomerania ilovaiskyi Sasonov 1960 (p. 162, pi. 3, figs. 1, la, pi. iv, figs. 2, 2a, b). This
species resembles Xenostephanus in having evolute inner whorls; a clear ventral smooth
band without tubercles; in its size; and in the extreme degeneration of the ribbing on the
body-chamber. It dilTers in being somewhat more compressed than any of the species
here described, and much more densely ribbed: at 50 mm. it has twenty-six ribs per
whorl. The primary ribs in consequence never attain the highly differentiated tubercu-
late character of those in, for example, ranbyensis, or even A. (Xenostephanoides)
scoticus, which is similarly densely ribbed. There are also prominent constrictions which
Xenostephanoides does not have. Sasonov’s material came from the Sosva basin, on the
eastern side of the northern Urals.

Family cardioceratidae Siemiradzki 1891
Subfamily cardioceratinae Siemiradzki 1891
Genus amoeboceras Hyatt 1900

Type species. Ammonites cdternans von Buch 1832.

Subgenus amoebites Buckman 1925

Type species. Amoebites akanthophorus Buckman 1925.

1. Amoeboceras {Amoebites) cf. kitchini (Salfeld)

Plate 32, fig. 26a, b

Cardioceras Kitchini Salfeld 1913, p. 423.

Cardioceras Kitchini Salfeld 1914, p. 129.

Cardioceras Kitchini Salfeld 1915, p. 189, pi. xix, figs. 8-17; pi. xx, figs. 15, 16.

Amoeboceras {Amoebites) kitchini Spath 1935, pp. 30-31 (neotype designated); pi. 1, fig. 9a, b.

238

PALAEONTOLOGY, VOLUME 6

Anioeboceras (Amoebites) kitchini Waterston 1951, p. 42, pi. ii, fig. Aa, b.

Amoeboceras {Amoebites) aflf. rasenense Waterston 1951, pi. ii, fig. 1.

Material. One (MS 48).

Description. The specimen is 29 mm. in diameter; septate to 20 mm., the last sutures
show none of the degeneration or approximation otherwise almost invariably found in
adults of the Cardioceratidae, so that the specimen seems to have been immature. It
agrees well, as far as it goes, with the neotype (Salfeld 1915, pi. xx, fig. 16), although it
does not, and could not at that size, show any of the looped ribbing or ventro-lateral
clavi of the adult A. kitchini.

Remarks: Salfeld, in his English paper (Oct. 1913), based the species on a drawing in
Woodward, 1895, but in his German paper (1914) he compared Woodward’s figure with
another in de Loriol (1876), which he queried in his monograph (1915). It is possible to
interpret the species only as from the 1915 monograph, when photographs were pub-
lished, including that of the Scottish neotype. Unfortunately, even the latter does not fix
the species unambiguously, being itself incomplete and recognizable only up to 27 mm. ;
and both the neotype of A. kitchini and the specimen described here might be hard to
distinguish from inner whorls of larger specimens differing in their outer whorls and
described under various names, e.g. A. akanthophorus Buckman, A. salfeldi Spath (nom.
nov. for Card, pingiiis Salfeld pars, 1915 pi. xx, fig. 14), or A. rasenense Spath (1935, pi. 1,
fig. 6a, b).

The amoeboceratids of the Lower Kimeridgian are as yet scarcely known. The tuber-
culate forms of Amoebites are of particular interest. Most of those hitherto published
are of the A. kitchini group, from the Baylei-Cymodoce Zones, and so we take the

EXPLANATION OF PLATE 32

Figs. 1-8, Aulacostephamis {Aiilacostephamtes) cf. eulepidus (Schneid). \a, b, MS 112, eight-tenths
whorl body-chamber with lappet on reverse side. 2a, b, MS 69, with lappet. 2a, b, MS 27, with
short lappet. Aa, b, MS 77, nine-tenths whorl body-chamber. 5a, b, MS 58, no sutures visible, but
peristome preserved with lappet and collar. 6a, b, MS 66, no sutures visible, la-c, R 5, with onset
of lappet. 8u, 6, MS 72, outer whorl body-chamber.

Figs. 9, 10, Aulacostephamis (Aiilacostephanites) ebrayoides sp. nov., transitional to A. (A.) cf. eulepidus
(Schneid). 9a-c, MS 8, with lappet. lOu, b, MS 10 with lappet.

Figs. 11, 12, Aulacostephamis {Aiilacostephanites) aflf. desmonotiis (Oppel). \\a, b, R 17, no sutures
visible, but probably part of the outer whorl is body-chamber. \2a, b, MS 115, ditto.

Figs. 13-18, Raseuia (Rasenioides) thernuiriim (Oppel). \2a, b, MS 14, with probably some body-
chamber. \Aa, b, MS 118, with eight-tenths whorl body-chamber, peristome, and slight lappet.
1 5a-c, MS 9, nine-tenths whorl body-chamber. 16a-c, MS 32, remarks as 1 1. I7a, b, MS 36, ditto.
18u, b, R 20, ditto.

Figs. 19-21, Raseuia (Rasenioides) cf. lepidiila (Oppel). \9a, b, MS 34, eight-tenths of last whorl body-
chamber. 20a, b, MS 53, a fragment with outer whorl body-chamber. 2la, b, MS 35, body-chamber
with peristome.

Figs. 22-24, Aulacostephamis {Aiilacostephanites) sp. aff. nioschi (Oppel). 22, MS 136, body-chamber
fragment. 22a, b, MS 24, with eight-tenths whorl body-chamber. 2Aa, b, MS 19, no sutures visible.

Fig. 25a~c, Anioeboceras {Amoebites) cricki (Salfeld). MS 111, outer whorl body-chamber with peri-
stome.

Fig. 26a, b, Amoeboceras {Amoebites) kitchini (Salfeld). MS 48, with half whorl body-chamber.

All natural size.

Palaeontology, Vol. 6

PLATE 32

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 239

opportunity to illustrate two examples of A. (Amoebites) salfeldi Spath from the Muta-
bilis Zone of Mull (PI. 33, figs. 12, 13). Development via the ventro-lateral clavi of A.
kitchini to the fully looped ribs with tubercles of A. salfeldi seems systematic and may be
of help in dating beds in regions, e.g. east Greenland, in which rasenids are absent.

2. Amoeboceras {Amoebites) cf. cricki (Salfeld)

Plate 32, fig. 25a-c

Cardioceras Cricki Salfeld 1914, p. 129.

Cardioceras Cricki Salfeld 1915, p. 191, pi. xix, figs. 2-6.

Cardioceras Kapffi Salfeld pars (non Oppel) 1915, pi. xix, fig. la-c.

Material. One (MS 111), and two fragments (MS).

Description. The figured specimen is about 22 mm. in diameter. The last half-whorl is
body-chamber and the peristome is preserved, with ventral rostrum. The whorl-section
is quadrate, compressed, with sharp shoulders and flat venter.

Remarks. The history of the name is similar to that of kitchini and interpretation is
possible only as from the 1915 monograph. Salfeld’s best figured specimen (pi. xix, fig. 2)
attains 25 mm., and is slightly more densely ribbed than the one described here. The
species seems to stay small and not to develop the modified ribbing characteristic of the
A. kitchini group.

ORIGIN AND AGE OF THE FAUNA

The Lincolnshire material has been collected at four localities (from north to south):
Elsham, 9 miles east of Scunthorpe and 15 miles north-north-west of Market Rasen;
South Willingham, 6| miles south-east of Market Rasen; Market Stainton, 9h miles
south-east of Market Rasen; and Ranby, | mile south of Market Stainton.

Elsham. The material in the British Museum (T. B. Parks coll.) came from ‘huge
boulders in sand’ found in a small pit on the Chalk Wolds J mile from Elsham chalk-pit.
The ammonites were briefly mentioned by Spath (1954), who described them as ‘probably
new and undescribed species of Rasenia'.

South Willingham. The following are extracts from notes kindly supplied by Dr. Thurrell.

‘Specimens labelled SW were collected from the village at a point 250 yards east of the
church where a water-main trench intercepted the boulders within the Chalky Boulder
Clay.’

Market Stainton. ‘Specimens labelled MS were collected from two massive boulders
(each about 3 feet across) in the corner of a large field 500 yards south-east of the church.’

Ranby. ‘Specimens labelled R were collected on the roadside 600 yards east of Ranby
church.

‘The MS and R specimens are from blocks which are only 725 yards apart, but it is
quite obvious that the blocks have been dumped. Enquiries failed to locate their exact
origin, except that the MS blocks came from the centre of the field in the corner of which
they lie.

240

PALAEONTOLOGY, VOLUME 6

‘It seems certain that all the blocks come from — or are likely to occur in — the lowest
levels in the Boulder Clay spread and one might presume also that they are of fairly local
origin. Mapping of the boulder-clay suggests ice-movement from north to south over the
existing Jurassic clay vale, the stream being confined by the Chalk Wolds escarpment
not a mile to the east of Market Stainton. Evidence of possible derivation from the
North Sea basin is lacking.’

The matrix and preservation of all the material is very similar : fine grey siltstone, some
pieces crowded with fossils including, besides the ammonites, small gastropoda, pectens,
oysters, and Astarte. There is no known outcrop of Lower Kimeridgian in such facies
in England, nor has any been found in borings. The origin of the material must remain
conjectural. Some of it has found its way to Norfolk, although for boulders of the size
of those found in Lincolnshire transport over such distances seems improbable. The
Kimeridgian is known to thin rapidly towards the Market Weighton axis, only some
30 miles north of Market Rasen, which influenced sedimentation in Jurassic times; and
a more sandy development in its immediate vicinity, perhaps under chalk cover or even
some way out to sea, would not be unexpected.

The question arises : do all these boulders contain identical faunas and hence are they,
geologically speaking, all of the same age? The distribution of the material among the
various localities is shown in Table 1, counting only identifiable specimens (some un-
numbered fragments are indicated ‘-f ’). The species of Aulacostephaninae can be divided
into two groups: those known elsewhere from the Mutabilis Zone — group (A); and
species of the subgenera Xeuostephanus and Xenoslephanoides peculiar to the present
collection (B). Whereas species of the second group (B) were found at all the localities,
those of the first (A) occurred only at Ranby and Market Stainton, and not at South
Willingham. There exist boulders, therefore, which contain exclusively the Xeuostephanus-
Xenostephanoides fauna. This leads to two possibilities for the age of the latter.

Firstly, if segregation is not general, and there also exist boulders containing both
groups of species, then these have the same age which is Mutabilis Zone sensu stricto.
The segregation of species in some parts of a ‘contemporaneous’ deposit would require
explanation, but cases of this have been observed in other deposits of dogger facies.

EXPLANATION OF PLATE 33

Fig. 1, Aulacostephanus (Xeuostephanus) aff. anceps sp. nov. Kildorais, Trotternish, Skye. W. G.
Cordey coll.

Fig. 2, Rasenia (Rasenioides) cf. transitoria (Schindewolf). MS 80a, Thurrell coll., with probably some
body-chamber.

Fig. 3o, b, Rasenia (Zonovid) sp. ind. R 2, wholly septate.

Figs. 4-6, Aulacostephanus (Aulacostephanites) eulepidus (Schneid). Latex casts of crushed impressions
from Kimeridgian shales of Mull (Geol. Surv. Scotland, nos. M4512i, T4734c, T1981d).

Fig. 7, Aulacostephanus (Aulacostephanites) spp. aff. eulepidus (Schneid) or ebrayoides sp. nov. As
above, from Mull (T1970d).

Fig. 8, Aulacostephanus (Xenostephanoides) aff. thurrelli sp. nov. or scoticus sp. nov. Mull (M4525i).

Figs. 9-11, Aulacostephanus (Aulacostephanoides) circumplicatus (Quenstedt). 9, Imprint of smooth
band on the venter shown at ‘v’. 10, Imprint of venter of Amoeboceras in the umbilicus. Mull
(M4516i, T4732C, T4733c).

Figs. 12-13, Amoeboceras (Amoebites) salfeldi Spath. Mull (T1991d, T4721c).

All natural size. Figs. 4-13 are photographs under oblique lighting of latex moulds coated with

magnesium oxide.

Palaeontology, Vol. 6

PLATE 33

CALLOMON and ARKELL, Kimeridgian ammonites

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES 241

The fossils occur notably in isolated pockets, the chance remains of accumulations
through wave-action at different moments over considerable periods of time in shifting,
unconsolidated sandbanks whose relics we must consider to be geologically of uniform
age. Segregation in such pockets might then be a reflection of ecology, particularly if the
remains are those of organisms that were gregareous.

TABLE 1. Summary of distribution of species in the Lincolnshire Drift

R

MS

SW

E

A. Rasenia {Rasenia) sp.

1

(Rasenioides) transit oria

1

cf. & aff. lepidida

5

thennarnm

1

8

cf. striolaris

2

iZonovia) sp.

1

Aidacostephamis

( Aidacostephanites) eidepidus

1

24

ebravoides

1

8

aff. desmonotus

1

2

aff. mdschi

1

10

(Aidacostephanoides) mutabilis

1

aff. mutabilis

1

2

circumplicatus

1

2

B. {Xenostephanoides) thurrelli

5

2

11

lindensis

3

4

21

(Xenostephanus) ranbyensis

1

1^

1

— var. elshamensis

1

staintonensis

1

1 +

anceps

Amoeboceras

{Amoebites) cf. kitchini

1

cf. cricki

3

Totals

17

77

34+

2

E = Elsham.

Alternatively, if the segregation of species is rigorous, we must deduce differences of
age. This could not have been large, for on zoological grounds Xenostephanus and
Xenostephanoides would fit into the faunal succession best as descendants of at least the
earliest of the well-known rasenids of the Cymodoce Zone, and certainly as precursors
of the aulacostephanids of the Eudoxus Zone (the lower part of the former Pseudo-
mutabilis Zone sensu lato — for revision, see Ziegler 1961). On geological grounds it
would also seem unlikely to have a selection of faunas of two zones, 1 and 3, with no trace
of the intermediate zone 2. At most, therefore, it seems possible that Xenostephanus and
Xenostephanoides mark a separate subdivision or subzone of the Mutabilis Zone.

The two alternatives above can only be resolved by further collecting, and careful
attention should be paid to analysis of new material boulder by boulder.

Note added in proof {1 February 1963). The pit at Elsham which yielded Xenostephanus
has been recently re-examined by P. E. Kent and R. Casey {Proc. Geol. Soc., London,

242

PALAEONTOLOGY, VOLUME 6

1605, 1963). They report that the sands and doggers exposed there are in fact in situ,
showing the presence of a local sandy development of Lower Kimeridgian immediately
under the Chalk. This had previously been mapped as Spilsby Sandstone (Neocomian).

The Xenostephamis fauna has also been found in drift at Miningsby, 4 miles west of
Spilsby, at the southernmost end of the Wolds. Two specimens are figured by Ziegler
(1962, pi. 22, figs. 5-6, 13-14; University of Nottingham Collection).

COMPARISON WITH OTHER AREAS

East Scotland. Lower Kimeridgian is exposed at two places: Eathie, on the Moray Firth,
south of Cromarty; and the Brora-Helmsdale strip in Sutherlandshire.

The Eathie exposure seems to lie mainly in the Baylei-Cymodoce Zones (Waterston
1951), although nodules washed up on the beach indicate some higher beds offshore.
The Lincolnshire fauna does not appear to be represented among them.

A more complete section is to be found in Sutherlandshire. The collections contain
material from all the zones of the Lower Kimeridgian, but there seems to be no system-
atically collected succession (for summary, see Bailey and Weir 1932). The specimens
that have found their way into the museums have tended to be the well-preserved ones
out of mudstone nodules, often found loose on the shore; among them is A. {Xeno-
stephanoides) scoticus sp. nov. (see above). Of the crushed ammonites from the rest of the
great thickness of shales making up most of the succession there is little record.

Some of the lower of these shales — the Loth River Shales — are highly fossiliferous.
The following section was recorded during a brief visit (J. H. C. 1956), in the river gorge
south of Loth station, just east of the railway bridge on the north bank of the river.
From above:

Loth River Shales

6.

5.

4.

3.

2.

Black paper shales with sandy intercalations and seams; to below rail-level at ft.

the north end of the bridge. seen c. 15

Soft yellow sandstone or sand, cross-bedded, with large doggers; variable c. 6

Black varved paper shales with sandy white layers; many crushed ammonites,
especially 2 ft. from the top:

Amoeboceras {Amoehites) beaiigrandi (? Sauvage) Spath (1935, pi. 5, fig. 4).

A. (A.) sp. aff. kitchini (Salfeld).

Rasenia? cf. mdschi (Oppel).

Aidacostephanus {Xenostephanoides ?) cf. lindensis sp. nov. also profuse plant

remains, and Aucella sp. 6

Sandstone, doggery, cross-bedded, barren 2 to 8

Black paper shales, as above, with sandy layers 10 to 15

in.

0

0

0

0

0

Allt na Cade Sandstone

1. Sandstone, white, massive, weathering black

seen 1 5 0

This section was not recorded in connexion with the present work, and only a few speci-
mens from bed 4 were brought back. There seems little doubt about the Aidacostephanus,
however, and more careful collecting from this section might settle the precise age of the
Lincolnshire material.

West Seotland. There are indications of the Lincolnshire fauna from two localities:
Kildorais, northern Skye, and Duart Bay, eastern Mull (see Arkell 1933, p. 114, fig. 19).

W. J. ARKELL AND J. H. CALLOMON; KIMERIDGIAN AMMONITES 243

The only description of the Skye section so far published is by MacGregor (1934,
p. 400), although it seems to be one of the most complete and fossiliferous successions
from Lower Oxfordian to Lower Kimeridgian in Britain (J. H. C. 1961). The Pseudo-
cordata Zone (uppermost Oxfordian), with profuse Ringsteadia, and Baylei Zone (Lower
Kimeridgian) with typical Pictonia are thick and easily recognizable. Higher beds are
present, but much less fossiliferous and badly covered by beach material: from one of
these Mr. W. G. Gordey obtained the Xenostephanns shown in Plate 33, fig. 1. Mac-
Gregor submitted his material to Spath for identification, and in a more detailed account
in a thesis (1931) mentions 'Pararasenia pseudomutabilis’ in text. Illustrations show
nothing identifiable of posX-baylei age, and search has failed to find the material. Thus
there is so far no unconfirmed material which could throw light on the age of the
single Xenostephamis mentioned above.

The occurrence in Mull is in a tiny faulted patch of baked blue shale on the right bank
of the stream flowing into Duart Bay (Lee and Bailey 1925). The identification of the
ammonites has had a varied history. They were first reported by Buckman to be Callo-
vian Kosmoceras and Reineckeia. This was corrected by Spath (1932, p. 149) and Arkell
(1933, p. 371), who identified them as Amoeboceras and Rasenia; but their precise age
was not mentioned and none of them has so far been described. They are in fact from
the Mutabilis Zone and the following species can be recognized in the collections of the
Scottish Geological Survey:

Aulacostephamis (Aiilacostephanites) eulepidus (Schneid) (PI. 33, figs. 4-6).

— — cf. ebrayoides sp. nov.

— — aff. nibscld (Oppel).

— (Aiilacostephanoides) mutabilis (Sow.).

— — circumplicatus (Quen.) (PI. 33, figs. 9-11).

— — linealis (Quenstedt).

— (Xenostephanoides) cf. thinrelli sp. nov. (PI. 33, fig. 7).

— {Xenostephamis) aflf. ranbyensis sp. nov.

Amoeboceras (Amoebites) beaiigraiidi (Sauvage).

— — beaiigramli Spath (non Sauvage).

— — cricki (Salfeld).

— kitchini (Salfeld).

— — salfeldi Spath (PI. 33, figs. 12, 13).

— {Eiiprionoceras) cf. sokolovi Sokolov and Bodylevsky.

Thus there is considerable similarity with the Lincolnshire fauna. The Xenostephamis
resembles the one from Skye, but is too poorly preserved to be worth figuring. Amoebo-
ceras predominates (this seems to apply to most Scottish Lower Kimeridgian), and the
small fine-ribbed rasenids seem to be rare or absent. This further suggests that the
Mutabilis Zone may be capable of subdivision.

England. Nowhere between Yorkshire and Dorset does the Mutabilis Zone appear to
exceed a few feet in thickness, and very little is known about it in detail. Locally it is
condensed or absent. Once again the museum material is largely that from nodules,
found in many of the small brick-pits which have now all vanished. Such material from
Lincolnshire is fairly abundant, but contains no Xenostephamis. This, too, may indicate
that these forms occur at a somewhat different level from the rest of the Mutabilis fauna.
Otherwise, if the Lincolnshire drift boulders are of relatively local origin, there would
have to be a very sharp ecological boundary in the neighbourhood of the Humber.

244

PALAEONTOLOGY, VOLUME 6

South Germany. Due largely to the work of recent years the faunal successions are now
known in considerable detail. The fauna of the Mutabilis Zone seems to be confined to
the White Jura S 1-2 (Ziegler 1962) (Tenuilobatum Zone partim, Badener Schichten of
Switzerland), but constitutes only a minor part of the whole fauna. It seems unlikely
therefore to become the type area for subdivisions of the Mutabilis Zone. Only one
specimen of ? Xenostephanus has so far been found (see above) and that not in place.
This confirms the impression that this group is primarily characteristic of the boreal
faunal realm.

REFERENCES

ARKELL, w. J. 1933. The Jurassic System in Great Britain. Oxford.

— — 1951. Proposed use of the plenary powers to designate the type species of the genus ‘Aulaco-
stephanus’ Tornquist, 1896 (Class Cephalopoda, Order Ammonoidea) (Jurassic). Bull. Zool.
Nomencl. 2, 188-90.

1955. Monograph of the English Bathonian Ammonites, Pt. v. Palaeont. Soc. Monogr.

BAILEY, E. B. and WEIR, J. 1932. Submarine Faulting in Kimmeridgian Times in East Sutherland.
Trans, roy. Soc. Edinb. 57, 429-67, pi. 1-3.

CALLOMON, J. H. 1957. Field Meeting in the Oxford Clay of Calvert and Woodham Brick Pits,
Buckinghamshire. Proc. Geol. T55. 68, 61-64.

1963. Sexual dimorphism in Jurassic ammonites. Trans. Leicester lit. phil. Soc., 57 (In press).

FREBOLD, H. 1930. Vcrbreitung und Ausbildung des Mesozoikums in Spitzbergen. Skrifter om
Svalbard og Ishavet, no. 31.

GEYER, o. F. 1961. Monographic der Perisphinctidae des unteren Unterkimeridgium (Weisser Jura y,
Badenerschichten) im siiddeutschen Jura. Palaeontographica, A 117, 1-157, pi. 1-22.

KHUDYAEV, I. 1932. The fauna of the Upper Kimmeridgian deposits of Timan. Bull. Geol. Serv.
U.S.S.R. 51, 645-53, pi. 1.

LEE, G. w. and BAILEY, E. B. 1925. The pre -Tertiary geology of Mull, Loch Aline and Oban. Mem.
Geol. Surv. Scotland.

LORiOL, p. DE. 1874. In LORiOL, p. DE and PELLAT, E. Monographic paleontologique et geologique
des etages superieures de la formation jurassique des environs de Boulogne-sur-Mer. Mem. Soc.
phys. Geneve, 23, 253-407, pi. 1-10.

1876. Monographic paleontologique des couches de la zone a Ammonites tenuilobatus (Badener

Schichten) de Baden (Argovie). Mem. Soc. pal. Suisse, 3, (pt. I), 1-32, pi. 1-4.

1878. Ibid. (pt. Ill), 77-200, pi. 13-22.

MACGREGOR, M. 1931. An account of the Sedimentary rocks of north Trottcmish, Islc of Skye. Thesis,
University of London (unpublished).

— ^ — 1934. The sedimentary rocks of North Trotternish, Isle of Skye. Proc. Geol. 45, 389-413.
NIKITIN, s. N. 1885. Die Cephalopodenfauna der Jurabildungen des Gouvernements Kostroma.
Verb, russ.-kais. Min. Ges. St. Petersburg, 20, 13-86, pi. 1-8. (Also published at the same time in
Russian, Mem. Comm. geol. St. Petersbourg, 2, no. 1.)

OPPEL, A. 1863. Uber jurassische Cephalopoden. Pal. Mitth. Mus. bayer. Staates, 3, 163-266, pi. 51-74.
ORBiGNY, A. d’. 1845. In MURCHISON, R. I., VERNEUiL, E. DE, and KEYSERLiNG, A. DE. Gcologie de !a
Russie d' Europe et des Montagues de TOural. Vol. II: Paleontologie. Systeme Jurassique: Mol-
losques. London and Paris.

PAVLOW, A. 1 886. Les ammonites de la zone a Aspidoceras acanthicum de 1 ’est de la Russie. Mhn.
Comm. geol. St. Petersbourg, 2, 1-91, pi. 1-10.

QUENSTEDT, F. A. 1887-8. Die Ammoniten des Schwdbischen Jura. Band 3: Der Weisse Jura. 818-944,
pi. 91-102 (1887); 945-1140, pi. 103-26 (1888). Stuttgart.

RAVN, J. p. J. 1912. On Jurassic and Cretaceous fossils from N.E. Greenland. Medd. om Gronl. 45, 10,
433-500, pi. 32-38.

REiNECKE, J. c. M. 1818. Maris protogaei Nautilos et Argonautas vulgo Cornua Ammonis in Agro
Coburgi et vicino reperiundos. . . . Coburg.

W. J. ARKELL AND J. H. CALLOMON: KIMERIDGIAN AMMONITES

245

SALFELD, H. 1913. Certain Upper Jurassic Strata of England. Quart. J. geol. Soc. Loud. 69, 423-30.

1914. Die Gliederungdesoberen Jura inNordwesteuropa. Miner. Bd.37, 125-246.

1915. Monographie der Gattung Cardioceras Neumayr und Uhlig, Teil I. Die Cardioceratiden

des oberen Oxford und Kimmeridge. Z. deutsch. Geol. Ges. 67, 149-204, pi. 16-20.

SASONOV, N. T. 1960. Novye dannye ob Oksfordskikh i kimeridjskikh ammonitakh. Paleontolo-
gicheskvi Sbornik, Leningrad, 3, Trudyi, Vyipusk xvi, 133-61, pi. 1-6. (English translation: Nat.
Lend. Lib. RTS 2019, D.S.I.R., London.)

SCHINDEWOLF, o. H. 1925. Entwurf einer Systematik der Perisphincten. Neues Jb. Miner. ,Bq\\. BA.
B 52, 309-43.

1926. Zur Systematik der Perisphincten. Neties Jb. Miner., Beil. Bd. B 55, 495-517, pi. 19.

SCHNEID, T. 1939. Uber Raseniiden, Ringsteadiiden und Pictoniiden des nordlichen Lrankenjura.
Palaeontographica, A 89, 117-86, pi. 1-14.

sowERBY, J. and sowERBY, J. DE c. 1812^6. The Mineral Conchology of Great Britain. London. 7 vols.
SPATH, L. F. 1931. Revision of the Jurassic Cephalopod Launa of Kachh (Cutch). Pal. Indica, n.s.
9, mem. 2, part iv, 279-550, pi. 48-102.

1932. The invertebrate faunas of the Bathonian-Callovian deposits of Jameson Land (East

Greenland). Medd. om Granl. 87, no. 7.

1935. The Upper Jurassic invertebrate faunas of Cape Leslie, Milne Land. I. Oxfordian and

Lower Kimmeridgian. Ibid. 99, no. 2.

1954. In DENNISON, V. D. and parks, t. b. Geology. Trans. Lines. Nats. Union, 13, 191-3.

TORNQUiST, A. 1896. Die dcgenerierten Perisphinctiden des Kimmeridge von Lc Havre. Abb. scinveiz.
paldont. Ges. 23, 1-43, pi. 1-8.

WATERSTON, c. D. 1951. The stratigraphy and palaeontology of the Jurassic rocks of Eathie (Cromarty).
Trans. Roy. Soc. Edinb. 53, 33-51, pi. 1, 2.

WOODWARD, H. B. 1895. The Jurassic Rocks of Britain. V. The Middle and Upper Oolites of England
(Yorkshire excepted). Mem. Geol. Siirv. Gt. Brit.

ZIEGLER, B. 1961. Stratigraphische und zoogeographische Beobachtungen an Aulacostephanus
(Ammonoidea — Oberjura). Paldont. Z. 35, 79-89.

1962. Die Ammonitengattung Aulacostephanus im Oberjura (Taxionomie, Stratigraphie, Bio-

logie). Palaeontographica, A 119, 1-172, pi. 1-22.

J. H. CALLOMON
University College, London,
Gower St., London, W.C. 1

Manuscript received 21 November 1961

A NEW SPECIES OF KOMIA KORDE AND THE
SYSTEMATIC POSITION OF THE GENUS

by E. C. WILSON, R. H. WAINES, and A. H. COOGAN

Abstract. Study of Komia eganemis sp. nov. from the Middle Pennsylvanian (Atokan) rocks of eastern Nevada
suggests that the genus be assigned to the Stromatoporoidea (Kingdom Animalia) rather than the Rhodophyceae
(Kingdom Plantae). Komia has been reported from rocks of Middle Carboniferous, Early Pennsylvanian, and
Middle Pennsylvanian (Atokan and Desmoinesian) ages. K. eganensis and Fusulinella acuminata define a
restricted biostratigraphic zone within the Middle Pennsylvanian (Atokan) rocks of eastern Nevada.

The Palaeozoic fossil Komia Korde (1951, p. 181) has been reported from eastern
Europe, central Japan, and western North America (Johnson 1960, p. 51 ; 1961, p. 86).
The following historical resume presents previous investigations concerning Komia and
is intended to serve as background for consideration of the genus.

This study is one in the series on the stratigraphy and palaeontology of the Ely No. 3
Quadrangle, White Pine County, Nevada. A list of previous papers in this series may be
found in Wilson and Langenheim (1962, p. 495).

Historical resume. Korde (1951, p. 181) described the genus Komia with Komia abimdans
as the type species (loc. cit., text-figs. 4, 5; pi. 2, figs. 3, 4) and considered it to be a
rhodophytean alga.

Maslov(1956, p. 21), in questioning the validity, in part, of Korde’s (1951, pp. 175-82)
study, suggested that Komia might be referable to the Echinodermata.

Johnson (1957, pp. 13, 80) cited Komia as a fossil alga. Later (1960, p. 45; 1961, p. 85)
he placed Komia among several genera of calcareous red algae of uncertain affinity. He
remarked (1960, p. 51 ; 1961, p. 86) that Komia has long been called ^Desmoinesia' [nomen
nudum] by certain North American geologists and that it occurs in the ‘Middle Car-
boniferous of Russia, Lower Pennsylvanian of central Japan, Des Moines group (Penn-
sylvanian) of west Texas and New Mexico’. He also figured Komia sp. ? from the
Pennsylvanian of southern New Mexico (1960, pi. 19; 1961, pi. 25).

Kordeophyton Rezak, apparently listed as a junior synonym of Komia Korde in Johnson
(1961, p. 286), appears to be a nomen nudum. We have been unable to verify that it has
ever been published elsewhere.

Illustrations (reconstructions?) labelled Komia abundans Korde appeared in Drush-
chits and Yakubovskaya (1961, pi. 2, fig. 10). They considered it (ibid., p. 44) to be a
Middle Carboniferous rhodophytean alga.

Komia sp. was recorded, without description or figures, from the Pennsylvanian of
Utah by Mollazal (1961, p. 26) and Wright (1961, p. 154). We have found no other
reports of the genus.

SYSTEMATIC DESCRIPTION

The morphological terminology used is largely that of Galloway (1957, pp. 350-60).
Type specimen and locality numbers refer to the collections of the Museum of Paleon-
tology, University of California, Berkeley (abbreviated to UCMP).

[Palaeontology, Vol. 6, Part 2, 1963, pp. 246-53, pi. 34-35.]

E. C. WILSON, R. H. WAINES, A. H. COOGAN: KOMIA KORDE

247

Phylum COELENTERATA Frey and Leuckart 1847
Class HYDROZOA Owen 1843
Order stromatoporoidea Nicholson and Murie 1878
Family Incertae sedis
Genus komia Korde 1951

Type species. Komia abiiiidans Korde (1951, p. 181, text-figs. 4, 5; pi. 2, figs. 3, 4).

Diagnosis. Coenosteum small, cylindrical, ramose, composed of a broad outer region
and a narrow axial cylinder; outer region formed of superposed, truncated cones of
perforate laminae and interlaminar to continuous pillars with minutely trabeculate, en
jet d'eau (Steiner 1932, pp. 24-28, text-figs. 3, 5) microstructure; axial cylinder composed
of elongate grooves curving upwards and outwards from axis and partially enclosed
abaxially by incomplete, perforate, distally thickening walls.

Remarks. In this study we have chosen to consider the Order Stromatoporoidea in the
broad concept presented by Lecompte (1956, p. F127). The combined features of Komia
appear so singular that it has been impossible to compare the genus closely with other
stromatoporoid genera except on the basis of individual characters. For this reason we
have not placed Komia in a family. Furthermore, we do not wish to erect a new family
on the basis of one genus and two species.

In its cylindrical and ramose form, in its possession of a differentiated axial region,
and in its possession of regular laminae and pillars, Komia most closely resembles two
Middle Devonian stromatoporoids, Idiostroma and Dendrostroma (see Galloway 1957,
pp. 443-4, pi. 34, figs. 8, 9). Komia differs significantly from these two genera in the
nature of the pillar microstructure, the relatively small size of the coenosteum, the
comparatively close spacing of the skeletal elements, the structure of the axial region,
and the coarsely porous nature of the laminae. Although differing in most characters,
Komia resembles the Permo-Triassic stromatoporoid Disjectopora (see Lecompte 1956,
p. F138, text-figs. Ill, 112) in the presence of a rectilinear perforate network of laminae
and pillars. The microstructure of the pillars of Komia most closely approaches that of
Mesozoic stromatoporoids such as Steinere/la (see Lecompte 1956, p. FI 38, text-fig. 2c)
and DehorneUa and Astroporina (see Hudson 1960, pi. 26).

In summary, Komia closely resembles the Middle Palaeozoic stromatoporoids men-
tioned above in its ramose and cylindrical form and in the relative regularity of its pillars
and laminae. It more closely resembles the Mesozoic and Late Palaeozoic stromato-
poroids in the more open and porous nature of the laminae, in the microstructure
of the pillars, and, to a lesser degree, in the relatively minute dimensions of the skeletal
framework.

Korde (1951, p. 181) interpreted Komia as an alga with a decidedly branched cylin-
drical thallus formed of a hypothallus of a small bundle of elongate, cellular filaments
and a thick perithallus of dichotomously branching, cellular filaments. Our study demon-
strates that Komia is a stromatoporoid with a non-cellular coenosteum formed of (1)
an outer region (= perithallus of Korde) composed of trabeculate pillars and perforate,
conical laminae, and (2) an axial cylinder (== hypothallus of Korde) composed of elon-
gate, abaxially walled grooves. To our knowledge, this combination of structures is not
known to occur in undoubted algae (Paul C. Silva, oral communication to Wilson and

248

PALAEONTOLOGY, VOLUME 6

Waines, July 1962, agrees). Calcareous algal skeletons are composed of cells or fine
tubules, or are gross as in some of the Cyanophyta, or are uniquely specialized as in the
Dasycladaceae and Charophyta. Komia is like none of these. On the other hand, the
presence of coarsely perforate laminae and trabeculate pillars strongly suggests affinities
with some of the Late Palaeozoic and Mesozoic stromatoporoids.

Our differences with Korde (1951, text-fig. 5) in the interpretation of the axial section
of Komia are illustrated, very diagrammatically, in text-fig. l.The conceptacle (?) cham-
bers in Korde’s reconstruction (see text-fig. la) were neither mentioned nor otherwise
illustrated by her (Korde 1951) and were not observed by us in our material. Since Korde

TEXT-FIG. 1. Diagrammatic representations of axial sections of Komia Korde showing differing
structural interpretations, a, after Korde; b, present paper.

(in Hit., May 1962), after examination of a thin section of K. eganensis, found our fossils
closely similar to her K. abimdans, we discount the possibility that our widely differing
interpretations are both correct and founded on non-related fossils.

Maslov’s (1956, p. 21) passing suggestion of similarities between Komia and some
echinoderms seems invalid since the discrete skeletal units do not behave as single calcite
crystals in polarized light, but are composed of microcrystalline fibres.

The two undescribed illustrations labelled Komia abimdans by Drushchits and Yaku-
bovskaya (1961, pi. 2, fig. 10) appear, possibly in agreement with our observations, to be
less cellular than the reconstruction of Korde (1951, p. 181, text-fig. 5). However, the
inclination of the laminae is totally different from that described in Korde (loc. cit.) and
in this study. Furthermore, the transverse section lacks the characteristic concentric

EXPLANATION OF PLATE 34

Photographs not retouched; figs. 1 and 5 photographed by transmitted light, figs. 2, 3, and 4 by
reflected light.

Figs. 1-5. Komia eganensis sp. nov. 1, 2, Axial section of holotype UCMP 30781 and transverse
sections of two other specimens, x 28. 3, 4, Transverse section of paratype UCMP 30782; 3, X 88 ;
4, x28. 5, Transverse section of paratype UCMP 30783, X 30. Twin central zones in figs. 4 and 5
indicate sections were cut just below a dichotomous branch.

Palaeontology, Vol. 6

PLATE 34

WILSON, WAINES and COOGAN, Komia

E. C. WILSON, R. H. WAINES, A. H. COOGAN: KOMIA KORDE 249

nature of the laminae of Komia and the structure of the inner zone is very obscure.
Because these important inconsistencies were not justified by Drushchits and Yaku-
bovskaya (1961) we feel that their figures either may represent fossils which are not
referable to Komia or else may be erroneously executed reconstructions. We therefore
have elected to disregard them as accurate representations of Komia.

Komia eganensis sp. nov.

[by Wilson and Waines]

Plate 34, figs. 1-5; Plate 35, figs. 1-4

External features (PI. 35, fig. 3). Coenosteum ramose, uniformly cylindrical in mature
portions, tapering distally to blunt conical tips, exceeding 5-2 mm. in length, up to 1-7
mm. in diameter; surface very minutely granulate; basal region of growth not observed.

Transverse section (PI. 34, figs. 1, 2 in part, 3-5; PI. 35, fig. 1). Coenosteum circular,
varying from 0-2 to 1-7 mm. in diameter, formed of an outer zone of concentrically
arranged laminae and radially arranged pillars and a central zone of irregular con-
struction.

Outer zone. Laminae concentric, usually complete but in places incomplete and merg-
ing with subjacent laminae, regularly spaced, numbering 6 to 7 in 0-33 mm., varying re-
spectively from 1 to 1 3 with coenosteal diameters of 0-2 to 1 -7 mm., irregular in thickness,
varying from 0-015 mm. between pillars to 0-035 mm. in vicinity of pillars, in many
places uniperforate between pillars with pores varying from 0-005 to 0-03 mm. or more
in width (best observed in tangential section); pillars generally spool-shaped between
laminae, less frequently expanding abaxially and tapering axially, occasionally irregular
or incomplete, confined to one interlamina or continuous or superposed through 2 to 5
or more interlaminae, generally from 0-01 to 0-035 mm. in width, numbering from 5 to 7
in 0-33 mm.; galleries generally circular, less frequently semicircular or horizontally
rectangular with vertical constrictions, generally 0-01 to 0-035 mm. in height, varying
from 0-01 to 0-165 mm. in width though averaging from 0-015 to 0-035 mm.; filaments
(possibly representing incompletely developed pillars) occasional, irregular, generally
vertically disposed, confined to interlaminae, generally less than 0-005 mm. in width;
microstructure of pillars fibrous with fibres curving upwards and outwards from central
portions of pillars (PI. 35, fig. 2).

Central zone. Generally circular, averaging 0-15 to 0-25 mm. in diameter, bounded by
innermost lamina of outer zone, generally composed of three intergrading portions:
innermost portion composed of an amoebiform complex of frequently interconnected
galleries; galleries measuring up to 0-05 mm. in width, frequently intersected by pillar-
like and filament-like structures with dimensions similar to those of comparable struc-
tures in outer zone; middle portion interfingering with gallery complex within and
grading to more regularly constructed outer portion, composed of an irregular perforate
network of galleries and pillar-like, lamina-like, and filament-like structures with dimen-
sions similar to those of comparable structures in outer zone; outer portion generally
interlamina-like with normal pillars and galleries as in outer zone.

Axial section (PI. 34, figs. 1, 2). Coenosteum elongate, ramose, of uniform width in
mature portions, tapering distally to blunt tips, greatly exceeding 5-0 mm. in length.

250

PALAEONTOLOGY, VOLUME 6

Up to 1-1 mm. in width, formed of an outer zone of regular laminae and pillars and
a central zone of less regular construction.

Outer zone. Laminae generally inclined downwards and outwards about 1 5 degrees
from the axis in inner portion of outer zone, becoming subparallel to parallel to surface
of coenosteum in outer portion, successively emerging at surface in a recessively over-
lapping manner, terminating axially by merging indistinctly with elements of central zone
and terminating unobtrusively on surface; complementary laminae on either side of
central zone generally terminating in apposition axially, rarely terminating in apposi-
tion surficially; dimensions and mutual disposition of laminae, laminar pores, pillars,
filaments, and galleries similar to comparable elements in outer zone of transverse
section ; microstructure of pillars similar to that of pillars in transverse section.

Central zone. Slender, elongate, continuous, averaging 0-2 mm. in width, generally
composed of two intergrading portions: inner portion composed of a complex of
irregular, perforate, indistinct filament-like structures gently curving upwards and out-
wards from axis and irregular, elongate, generally indistinct galleries frequently inter-
sected by irregular filament-like structures and very rarely intersected by thin transverse
structures; heights of galleries and widths of filament-like structures similar to those of
comparable elements in transverse section; outer portion merging indistinctly with inner
margin of outer zone, composed of a complex of thickening distal extremities of the
curved filament-like structures of the inner portion, pillar-like and filament-like struc-
tures and round irregular galleries; dimensions of galleries and structural elements
similar to those of comparable elements in outer zone.

Tangential section (PI. 35, fig. 4). Pillars round to irregular, generally separate, occasion-
ally coalescent; usually 0-015 to 0-030 mm. in diameter, numbering about 20 to 25 in
0-28 of a square mm., galleries coalescent about pillars; laminae appearing as irregular
perforate areas; laminar pores generally round, usually 0-015 to 0-035 mm. in diameter,
numbering about 15 in 0-28 of a square mm.

Documentation. Holotype 3078 1 and paratypes 30782 to 30790 are from locality B.4854. Approximately
fifteen polished sections and eighteen thin sections, exhibiting a total of several hundred specimens of
K. eganensis, were prepared and studied. In addition, serial photographs of successive transverse
sections of one coenosteum served to reveal more fully the nature of the skeletal framework.

Name. The species is named after the Egan Range in eastern Nevada, from which many of the speci-
mens of K. eganensis used in this study were obtained.

Occurrence. K. eganensis was collected from six localities in eastern Nevada, which are listed below.

Discussion. K. eganensis is similar to K. abundans in the overall form and size of the
coenosteum and in the general spacing of the laminae. In the former species, less closely
spaced pillars (5 to 7 in 0-33 mm. v^. 8 or more in the latter) and a larger diameter (0-2
vs. 0 08 mm.) of the axial cylinder are considered characteristic.

EXPLANATION OF PLATE 35

Photographs not retouched; figs. 1, 3, and 4 photographed by reflected light; fig. 2 by polarized light.

Figs. 1-4. Komia eganensis sp. now. 1, Transverse section near distal extremity of coenosteum, X 105.
2, Microstructure of pillars, X250. 3, Fragments of silicified coenostea, x5-4. 4, Tangential section
showing perforate lamina, paratype UCMP 30784, X75.

Palaeontology, Vol. 6

PLATE 35

WILSON, WAINES and COOGAN, Komia

251

E. C. WILSON, R. H. WAINES, A. H. COOGAN: KOMIA KORDE

Komia sp. ?, figured by Johnson (1960, pi. 19; 1961, pi. 25), is not described and is
insufficiently well illustrated for specific comparison.

The material upon which the present study is based occurs as abundant, fragmentary,
partially silicified coenostea in a medium grey-brown, fine-grained, biofragmental lime-
stone. Due to some as yet unidentified character inherent in the structure of the unaltered
skeletal elements, much more morphological detail was revealed by the study of polished

TEXT-FIG. 2. Map of part of eastern Nevada
showing localities from which Komia eganensis
sp. nov. was collected.

TEXT-FIG. 3. Generalized columnar section
of part of the Ely Limestone near Sunny-
side, Nevada, showing stratigraphic distri-
bution of Komia eganensis in relation to
some other fossils.

and slightly acid-etched surfaces under oblique reflected light (e.g. PI. 34, figs. 2-4; PI. 35,
figs. 1, 4) than was revealed by thin sections under transmitted light (e.g. PI. 34, figs. 1, 5).
The nature of the microstructure of the pillars, however, was best observed by means of
the latter method. We recommend use of both methods in studying Komia. It is felt that
observations limited only to transmitted light (e.g. Korde 1951, pi. 2, figs. 3, 4, text-fig. 4;
and Johnson 1960, pi. 19; 1961, pi. 25) may have restricted some investigators’ interpre-
tations of the structure of Komia.

STRATIGRAPHIC DISTRIBUTION

Komia is widespread in the Pennsylvanian rocks of the western United States. Johnson
(1960, p. 51) reported its presence in west Texas and New Mexico in rocks of Desmoi-
nesian age. Mollazal(1961,p.26)and Wright (1961, p. 154) reported it in rocks of Derryan

C 1213

s

252

PALAEONTOLOGY, VOLUME 6

(= Atokan) age in the Ely Limestone of Millard County, Utah, and the Oquirrh For-
mation of Tooele County, Utah, respectively. Mollazal (1961, p. 26) further commented
that he had observed Komia elsewhere ‘in eastern Great Basin in Derryan to Desmoi-
nesian limestones’. In the Ely Limestone of east-central Nevada and in the Bird Springs
Formation of southern Nevada, K. eganensis occurs with Fusulinella spp. of Atokan age.
Fragmentary remains with associated fusulinids and other bioclastic debris form co-
quinoid limestones. Korde (1951, p. 181) reported similarly that Komia together with
foraminifera substantially make up many limestone beds along the Un’ya River in the
northern Ural Mountains of Russia. Johnson (1960, p. 51) reported Komia in the Lower
Pennsylvanian of central Japan, but did not further discuss its occurrence there.

In a characteristic section near Sunnyside, Nye County, Nevada (text-fig. 3), Komia
eganensis is associated with Fusulinella acuminata Thompson in the Ely Limestone from
about 1,500 to 1,820 feet above the base. Its lowest occurrence is 150 feet above the
Chaetetes-Profusulinella faunizone. Komia is not known to occur with Profusulinella,
Fusulinella devexa Thompson, or younger faunas in this area.

In the Ely Limestone at Moorman Ranch, White Pine County, Nevada, K. eganensis
occurs 375 feet above Caninia torquia (Owen) and 150 feet above the Chaetetes-Profusuli-
nella faunizone. In the Arrow Canyon Mountains, Clark County, Nevada, K. eganensis
and Fusulinella acuminata are present 1,375 feet above the base of the Bird Spring
Formation and 40 feet above the Chaetetes-Profusulinella faunizone. The association of
K. eganensis and F. acuminata Thompson apparently characterizes a restricted biostrati-
graphic zone in eastern Nevada.

LOCALITIES

The abbreviations T. 00 N., R. 00 E. appearing below represent Township 00 North, Range 00 East,

Mt. Diablo Base and Meridian.

B.4851. Bird Spring Formation, Arrow Canyon Quadrangle, Clark County, Nevada. Two- to three-
foot resistant limestone ledge 1,285 feet above base of formation near red painted number 26 and
yellow painted ‘C’. The outcrop is in Arrow Canyon, accessible from U.S. Elighway 93 by turning
west past a ranch house 12 miles north of Glendale Junction on to a dirt road which passes through
a dump and into the canyon. Collected by Coogan.

B.4853. Ely Limestone, lllipah Quadrangle, White Pine County, Nevada. ‘Moorman Ranch’ locality.
Komia occurs with fusulinids 1,360 feet above base of Ely Eimestone exposed on a spur overlooking
U.S. Highway 50 (north side). The line of traverse is marked with many wooden stakes. Komia can
be collected at the stake with a brass ring numbered 485, approximately 150 feet above a well-
defined bed with silicified Chaetetes. Collected by Coogan.

B.4852. Ely Limestone, T. 8 N., R. 62 E., Lincoln County, Nevada, near Sunnyside. Locality is
1,490 feet above base of Ely Limestone in section due east of Silver Spring, which lies a short
distance north of Shingle Pass road at the base of the west side of the Egan Range. Collected by
Coogan.

B.7873. Same section as B.4852, 20 feet higher.

B.7874. Same section as B.4852, about 310 feet higher and 10 feet below a red number 7 painted on
the rocks.

B.4854. Ely Limestone, Ely No. 3 Quadrangle, White Pine County, Nevada. Locality lies in T. 14 N.,
R. 62 E. just south of crest of ridge bordering north side of south fork of Willow Creek near 9040-
foot contour line as shown on USGS Advance Sheet for the SE. quarter of the Ely No. 3 Quadrangle,
1958 Edition. Grey limestone unit with K. eganensis, Chaetetes favosiis, Multithecopora hypatiae,
and Atokan fusulinids. Collected by R. L. Langenheim, Jr.

E. C. WILSON, R. H. WAINES, A. H. COOGAN: KOMIA KORDE

253

Acknowledgements. J. Harlan Johnson (of the Colorado School of Mines), Wayne L. Fry, and Paul C.
Silva (paleobotanist and botanist, respectively, of the University of California, Berkeley), kindly
examined some of our specimens and discussed possible algal affinities. Drs. Fry and Silva also read
the manuscript and suggested apposite changes. K. B. Korde, of the Paleontological Institute, U.S.S.R.
Academy of Sciences (Moscow), graciously examined one of our thin sections of Komia eganensis and
agreed that it was properly placed in her genus. Thanks are due to J. Wyatt Durham of the University
of California (Berkeley) for criticisms and suggestions made during this study and for reading the
manuscript. R. L. Langenheim, Jr., of the University of Illinois, supplied the fossils which instigated
this study and contributed locality and stratigraphic data for his material.

Coogan contributed specimens from and information regarding localities B.4851-3 and B. 7873-4,
text-figs. 1 and 3, and the section entitled Stratigraphic Distribution. In addition, he assisted with
translations. Wilson and Waines are responsible for the remainder of the paper, including the descrip-
tion of K. eganensis sp. nov.

REFERENCES

DRUSHCHiTS, V. V. and YAKUBOVSKAYA, T. A. 1961. PaleobotanickeskH atlas. Univ. Mosk., Moscow,
179 pp., 64 pi.

GALLOWAY, J. J. 1957. Structure and classification of the Stromatoporoidea. Bull. Amer. Paleont. 37
(164), 333-480, pi. 31-37.

HUDSON, R. G. s. 1960. The Tethyan Jurassic Stromatoporoids Stroinatoporina, Dehornella, and
Astroporina. Palaeontology, 2, 180-99, pi. 24-28.

JOHNSON, J. H. 1957. Bibliography of fossil algae: 1942-1955. Colo. Sell. Min. Quart. 52, 1-92.

1960. Paleozoic Solenoporaceae and related red algae. Ibid. 55, 1-77, pi. 1-23.

• 1961. Limestone-building Algae and Algal Limestones. 297 pp., 14 tables, 139 pi. Boulder, Nev.

KORDE, K. B. 1951. Novye rody i vidy izvestkovykh vodoroslei iz kamennougoTnykh otlozhenii
Severnogo Urala. Trud. Mosk. obscli. ispy. prir., otd. geol. 1, 175-82, pi. 1-3.

LECOMPTE, M. 1956. Stromatoporoidca. In bayer, f. m. et al. Treatise on Invertebrate Paleontology,
ed. R. c. MOORE, Part F, Coelenterata, F107-F144, text-figs. 86-114. Lawrence, Kansas.

MASLOV, v. p. 1956. Iskopaemye izvestkovye vodorosli SSSR. Akad. nauk SSSR, Trud. Inst. geol.
nauk, 160, 1-301, pi. 1-86.

MOLLAZAL, Y. 1961. Petrology and petrography of Ely Limestone in part of eastern Great Basin.
Brigham Young Univ. Geol. Stud. 8, 3-35.

STEINER, A. 1932. Contribution a I’etude des Stromatopores secondaires. Bull. Labs. Univ. Lausanne,
50, 1-117, pi. 1-14.

WILSON, E. c. and langenheim, r. l., jr. 1962. Rugose and tabulate corals from Permian rocks in
the Ely Quadrangle, White Pine County, Nevada. J. Paleont. 36, 495-520, pi. 86-89.

WRIGHT, R. E. 1961. Stratigraphic and tectonic interpretation of Oquirrh Formation, Stansbury
Mountains, Utah. Brigham Young Univ. Geol. Stud. 8, 147-66,

E. C. WILSON

Paleontology Department,
University of California,
Berkeley 4, California

R. H. WAINES

Paleontology Department,
University of California,
Berkeley 4, California

A. H. COOGAN

Humble Oil and Refining Company,

Manuscript received 4 August 1962 Houston, Texas

SOME SILICIFIED ORDOVICIAN FOSSILS FROM

SOUTH WALES

by NILS SPJELDN^S

Abstract. A preliminary description is given of a silicified fauna from the Upper Llanvirn or Lower Llandeilo
of South Wales. The material consists mostly of trilobites, ostracods, and bryozoans. One new genus and two
new species of ostracods are named, and the ontogeny of Tallinnella compUcata (Salter) is described. Because
of the silicified and fragmentary nature of the material, most of the numerous bryozoan species have not been
named formally.

During the author’s studies on some Ordovician brachiopods (Spjeldnaes 1959), it was
discovered that a slab of limestone, stated to come from the Llandeilo of Llan Mill, near
Narberth, Carmarthenshire, was rich in silicified fossils. The specimen, which belongs
to the Geological Survey and Museum, London (no. 85415), is part of an old collection,
and further details concerning the horizon and locality are not known. A description of
the localities near Llan Mill is given by Strahan et al. (1914, pp. 31-33). The age of the
fauna is discussed below.

The unsilicified fossils found on the slab are: Macrocoelia (?) Uandeiloensis (Dav.),
Sowerbyella antiqua Jones, Horderleyella cf. subcarinata MacGregor, DaJmanella cf.
parva Williams, and other brachiopods (for the brachiopod fauna, see MacGregor 1961),
indeterminate gastropods and cystid fragments.

The brachiopods, molluscs, echinoderms, and some of the trilobites are not silicified,
and are preserved as moulds in the shale partings only. In spite of the small size of the
slab (about 20x8x2 cm.), it yielded, after solution in hydrochloric acid, several hun-
dred silicified fossils, mainly ostracods, bryozoans, and trilobites. The silicified fossils
are described below.

Except for the fragmentation, which might be a local feature only, the silicified speci-
mens are excellently preserved, almost equal to the famous material from the Edinburgh
Eormation in Virginia, U.S.A. (cf. Whittington and Evitt 1956).

The author is deeply indebted to Dr. Stubblefield, Geological Survey and Museum,
London, for the loan of the specimens. Dr. Gunnar Henningsmoen and Dr. A. L. Guber
for inspiring discussions about the ostracods, and to Miss B. Mauritz and Mr. O. Bryn-
hildsrud for photographing the specimens.

SYSTEMATIC DESCRIPTIONS
Class TRILOBITA

Marrolithus inflatus incipiens Williams 1948
Plate 36, fig. \a~b

1948 Marrolithus inflatus incipiens — Williams, Geoi. Mag. 85, p. 77, text-fig. 6, pi. vi, fig. 2.

Materiai. Very plentiful (about 120 fragments), but fragmentary. In most specimens, only one-half of
the fringe is preserved, and this prevents a statistical study of the material.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 254-63, pi. 36-37.]

NILS SPJELDN^S: SILICIFIED ORDOVICIAN FOSSILS

255

The wide range in size of the specimens allows a study of the variation of the fringe.
In the largest specimens, the raised pits are more numerous, and they approach M.
inflatus inflatus. In the smaller specimens the number of raised pits gradually decreases
with size, and in the smallest specimens they are absent.

The preservation of the material shows that the pits are continuous, and hyperbolic in
cross-section. The raised pits are steeper (more cylindrical) than the ordinary ones.

In Tretaspis, from studies made by Stormer (1930), it is known that the pits are hour-
glass-shaped, parted by a suture in the middle. In M. inflatus incipiens no suture is
observed, and the curvature of the walls of the pits is the reverse of that found in Tretaspis.

Other trilobites. The other silicified trilobites are Flexicalymene cambrensis, indeter-
minable asaphid fragments, and a small cephalon of an Ampyx sp., which was broken
during preparation.

Class CRUSTACEA
Subclass OSTRACODA
Tallinnella complicata (Salter)

Plate 36, figs. 9-13, text-fig. 1

1947 Tetradella complicata (Salter) — Harper, GeoL Mag. 84, pp. 345-53 (for further references,
see this paper).

Material. Four hundred valves which were sufficiently complete for measurement, and a large number
of fragmentary valves. As mentioned above, it is possible that this is topotype material of Salter’s
species.

Description. The markedly preplete valves are rather flat in profile in adult specimens,
whereas in the larval instars they are more regularly swollen. In all stages the surface is
smooth, except for the four distinct lobes, two of which (LI and L3) protrude over the
hinge-line.

The smaller larval valves also have a marginal row of short, thin spines which are not
found in the adults.

The hinge itself is generally smooth in both valves; some teeth-like structures were
observed, but, as usual in silicified specimens, it is very difficult to determine if such
delicate structures are original or due to somewhat capricious silicification. It is, anyhow,
not a constant feature in the present material.

Some slight variation was observed in the material present, especially in the size of the
swollen ends of LI and L3, and in the outline of the domicilium.

Text-fig. 1 records measurements of 400 valves. Several growth stages are discernible
(I-VI). Some of them have the size theoretically expected (growth factor about 1-26, cf.
Spjeldnaes 1951), but between the third and fourth larval stage the difference in size is
much smaller than expected, especially the width, which is almost the same in both.

If the growth series demonstrated by the stages VI-V-IV is extrapolated according to
Brooke’s law, the theoretical stage III should be about 1 mm. wide and L5-L6 mm.
long. Some few specimens of this size are found, in addition to the normal stage III. If,
similarly, the growth series of the larger stages I-II-III are extrapolated, the theoretical
stage IV will fit with some few scattered specimens, and not with the majority observed
in stage IV. This peculiar disconformity in the growth series might be due either to a
mixing of two populations, or an unknown feature in the ontogenetic development of
T. complicata.

256

PALAEONTOLOGY, VOLUME 6

Opik (1937) suggested that Tallinnella showed sexual dimorphism, and this is con-
firmed by the present material. About 30 per cent of the adult specimens show a promi-
nent velar flange in the anterior part (PI. 36, fig. 12), and about 40 per cent are without
this flange (PI. 36, fig. 13). In the rest, this feature could not be observed clearly. Varia-
tion in size of the flange was observable but small, and all specimens in which the inside
was well preserved could be referred to one of the two dimorphs without doubt.

TEXT-FIG. ]. TaUitmella complicata (Salter) from Llan Mill, near Narberth, Geol. Suiv. Mus.
No. 85415. Measurements of 400 valves, showing instars.

Remarks. This species resembles in most features T. (?) bohemica (Barr.) as described by
Jaanusson (1957, pp. 342-3, pi. x, fig. 3). The lobes are stronger, and the extralobal area
is not visible in lateral view in T. complicata. The limitation of the genus TaUitmella is not
considered here, the term is used in the wide sense of Henningsmoen (1953, pp. 213-14),
and not in the more restricted sense of Jaanusson (1957). T. complicata differs consider-
ably from the type species, T. dimorpha, especially in the development of the extralobal
area.

Genus gunnaropsis gen. nov.

Diagnosis. Four-lobed ostracods with LI, L2, and L3 each developed as a sharp crest, and
L4 as a gentle swelling. A histial (?) flange starts at the upper anterior end of the domi-
cilium, turns almost horizontal below Ll-3, and stops abruptly in the lower part of L4.
A velar (?) flange starts just in front of the middle of the valve, and runs parallel to the
histial flange almost to the upper posterior corner, where it stops abruptly. Dimorphism
not observed.

NILS SPJELDN^ES: SILICIFIED ORDOVICIAN FOSSILS

257

Type species. Gimnaropsis cristata sp. nov.

Remarks. This genus resembles some from the Lower Ordovician, such as Rigidella
Opik, ProtaUirmeUa Jaanusson, and TaUinneUma Jaanusson. It differs from them, and,
as far as the author knows, from most other Palaeozoic ostracods in that the velar and
histial flanges are not parallel to the margin of the valves. Gimnaropsis also differs from
the three mentioned genera in that its four lobes are not distinctly united, and that L4
is developed as a rounded ridge, contrasting with the sharp edges of the other lobes.
In the other three genera, the differences between the lobes in this respect are much less
strong.

The family relationship of Gimnaropsis is at present obscure, especially since the
presence or absence of dimorphism, and the possible type of dimorphism, cannot be
ascertained because of the small amount of material present. The three other genera
mentioned, as well as Tallinnella, have been referred to the Quardijugatorinae by Jaan-
usson (1957, pp. 338-40), but it is rather doubtful if Gimnaropsis can be accommodated
to this subfamily.

Gimnaropsis cristata sp. nov.

Plate 36, fig. 6

Diagnosis. Same as diagnosis for genus.

Description. Outline of valves slightly preplete, with long, straight hinge-line. LI, L2, and
L3 have sharp crests, and LI and L2 are connected ventrally, forming a sharp V. L4 is
well developed, but lacks a crest in its dorsal part. L3 protrudes over the hinge-line, and
this is generally the case also with LI and L4, but to a lesser extent.

There are two flanges; one, which is interpreted as the velar, starts below Ll-2, at the
margin of the valve, and ends abruptly beside L4, just before reaching the hinge-line.
The distance from the margin of the valves increases gradually. Below Ll-2, it is found
at the margin of the valve; at the end, it is in close contact with L4, some distance from
the margin.

The other flange, which is interpreted as a histial one, starts at the upper anterior end
of the valve, near the margin, and the distance from it gradually increases. Below Ll-2,
it joins the connecting lobe and turns horizontally. It ends abruptly in the ventral part
of L4.

Fifteen specimens of this species were found, the larger ones being about 1-5 mm.
long and 0-9 mm. wide. Some smaller specimens, about 1-0 mm. long and 0-55 mm.
wide, probably represent a larval stage. No dimorphism is observed, and it is possible,
also, that the larger specimens are not adults.

Ceratopsis britannica sp. nov.

Plate 36, fig. 7

Diagnosis. A long and low Ceratopsis species with a prominent carina-like bulge con-
necting LI and L4.

Description. The holotype is 1-3 mm. long and 0-7 mm. wide. L4 is faintly developed,
L3 is prominent and rounded, L2 is shorter, but still prominent, and L4 continues into
the rounded triangular ‘horn’ with strong horizontal striation, which is characteristic for

258

PALAEONTOLOGY, VOLUME 6

this genus. A histial (?) bulge unites L4 and possibly LI. It is blunt-edged, and directed
outwards. A velar flange is developed, and is widest in the anterior part of the valve.
No sexual demorphism was observed among the about 20 specimens studied.

Some of them are rather small (LO-Ll mm. long and 0-55 mm. wide), representing
larval stages, and as usual the lobes are bulbous and diffuse. The horn on L4 is also
proportionally much longer in the smaller specimens. This species differs from the
other members of the genus in the proportions, and in the strong development, of the
carinal bulge.

Lomalobolbina sp.

Plate 36, fig. 8

Seven specimens of this species are present, both tecnomorphs and heteromorphs.
An average specimen is 1-2 mm. long. Because of the development of a histial dolon,
the sigmoidal sulcus S2, and the strongly developed posteroventral lobe, it probably
belongs to Lomatobolbina Jaanusson (1957, pp. 395-9). It resembles L. mammillata
(Thorslund) cf. Jaanusson (1957, pi. 12, figs. 6-8) in the absence of a distinct node on the
posteroventral lobe, but differs from it in outline, and in the more anterior position of S2.

Other ostracods. In addition to the species described above, there are also some smooth
specimens in the material present. One species represented probably belongs to Concho-
primitia, but the number of undamaged valves is too small (4-5 specimens) for a detailed
description. The other smooth valves are very small, and of different shapes. Some of
them might be young larval valves.

Phylum BRYOZOA

Bryozoans are very abundant in the material, but they are generally fragmentary. In
order to give a modern description of Ordovician bryozoans, thin sections are absolutely
necessary, especially for the Trepostome bryozoans, and also for Cryptostomes. As it is

EXPLANATION OF PLATE 36

All specimens belong to the Geological Survey and Museum, London. The specimens have not been

whitened and the photographs are not retouched. All the specimens come from the Upper Llanvirn

or Lower Llandeilo at Llan Mill, near Narberth, Carmarthenshire.

Fig. 1. Marrolithus inflatm incipiens Williams. Upper (fig. \d) and lower (fig. \b) sides of a fragment
of the fringe, x about 6.

Figs. 2, 4. Mesotrypa aff. le?ts (M‘Coy). 2, Lower surface of a zoarium, showing numerous mesospores
surrounding the zooecia. 4, Upper surface of another zoarium, showing only very few and small
mesospores, X about 8.

Fig. 3. Orbigiiyella favulosa (7) (Phillips). Upper surface of a fragmentary zoarium, X about 8.

Fig. 5. Indeterminate trepostome bryozoan D, showing central tube in a fragmentary zoarium, X 18.

Fig. 6. Gmmaropsis cristata gen. and sp. nov. Holotype. Large, complete valve, X 27.

Fig. 7. Ceratopsis britanmca sp. nov. Holotype, x 27.

Fig. 8. Lomatobolbina sp., X 27.

Figs. 9-1 3. Tallinnella complicata (Salter). 9, 10, Young instars of the two last larval stages, X 27. 11,

Exterior of an adult valve with histial dolon (female?), x 27. 12, Interior of an adult valve with histial
dolon (female?), x 27. 13, Interior of an adult valve without histial dolon, X 27.

Palaeontology, Vol. 6

PLATE 36

m

‘w

'*»%

3

mmB

SPJELDNyES, Silicified Ordovician fossils

•ifK:

NILS SPJELDN^S: SILICIFIED ORDOVICIAN FOSSILS

259

difficult to get good thin sections from silicified material, description of the species from
the present material must remain incomplete at present. Therefore no new species have
been named even though the whole fauna is new. The external features, as usual in
silicified material, are very well preserved, and this, combined with the few features of
the interior which were observed, gives some indication of the type of fauna found.

Order trepostomata
Orbignyella favulosa ? ( Phillips)

Plate 36, fig. 3; Plate 37, figs, \2a-b

(?) 1957 Orbignyella favulosa (Phillips 1848) — SpjeldnEes, pp. 367-8, pi. 12, figs. 3, 7. (For further
references, see this paper.)

Thin encrusting zoaria probably referable to this species are common. They differ from
the type in being thinner, and smaller in diameter. The typical curved diaphragms are
observed only in the larger specimens, but the maculae can be seen clearly, and acantho-
pores and mesopores are absent, both at the base and surface of the zoaria.

The type of this species (GSM 56404, cf. Spjeldnses 1957, p. 367) comes from the
Llandeilo Limestone of Llan Mill, and the present material might therefore be topotypes.
The detailed locality and horizon is, however, not known either for the type or for the
present material.

Mesotrypa aflf. lens (M‘Coy)

Plate 36, figs. 2, 4

Externally this species resembles O. favulosa, but it has smaller zooecia, less-developed
maculae, and shows the characteristic development of the mesopores. They are clearly
visible at the base of the specimens (PI. 36, fig. 2), but very few are seen at the upper
surface (PI. 36, fig. 4).

The present material differs from the types (cf. Spjeldnccs 1957, pp. 368-70, pi. 13, figs.
5, 7, text-fig. 1c-e) in its smaller size and less-developed maculae. There are also fewer
tabulae in the zooecia.

Species of this kind, usually referred to as "Nebulipora lens' in the older literature, are
common in the British Middle Ordovician, and seem to replace the Diplotrypa petro-
politana, which is the most common trepostome bryozoan in the Scandinavian-Baltic
Province, and rather rare in Britain.

Indet. Trepostome A

Plate 37, fig. 7

Ramose zoaria, with branches about 2-5 mm. thick. Bifurcation unknown. The
elongate-rounded zooecia have thick walls, and are almost completely surrounded by
the numerous, rounded polygonal mesopores. There are numerous small acanthopores.
No tabulae have been observed in the zooecia (only the outer part of the exozone is
preserved), but the mesopores appear to be tabulated.

This species is the most common ramose trepostome bryozoan in the material.

260

PALAEONTOLOGY, VOLUME 6

Indet. Trepostome B

Plate 37, fig. 6

Thin ramose zoaria (branches approximately 0-5 mm. in diameter) without branching
in the fragments at hand. Zooecia rather large (approximately 0-06 mm. in diameter),
polygonal, and with few mesopores. Walls thin, and no acanthopores observed. Only
few diaphragms have been observed in the zooecia and in the mesopores. A considerable
variation is found as to number and arrangement of the mesopores, but maculae are not
developed.

This species recalls thin-branched species of Hallopora, but generic identification
cannot be confirmed without thin sections.

Indet. Trepostome C

Plate 37, fig. ia~b

A small zoarium forming a hollow cone about 2 mm. in diameter, and 1 mm. high.
Large, polygonal zooecia with thick walls. No mesopores and acanthopores observed.
No diaphragms.

The conical shape might either be due to incrustation of a conical object, or it might be
the normal shape of the zoarium. The material is insufficient for a final decision.

Indet. Trepostome D

Plate 36, fig. 5

Externally the zoaria appear to be normal ramose ones, with 1-1 -2 mm. diameter, but
cross-sections show that the colony was formed around a hollow tube with 0-3 mm.

EXPLANATION OF PLATE 37

All specimens belong to the Geological Survey and Museum, London. They come from the Upper

Llanvirn or Lower Llandeilo of Llan Mill, near Narberth, Carmarthenshire. The specimens have been

whitened with ammonium chloride, but the photographs have not been retouched.

Figs. 1-5. Arthrostylid cryptostome bryozoans. Six fragments of zoaria, three of which (figs. 1, 2,
and 5) have the articulated base preserved. They illustrate the wide range of variation found in this
material, X 30.

Fig. 6. Indeterminate trepostome bryozoan B. Fragment of a zoarium with unusually many meso-
pores, X 18.

Fig. 7. Indeterminate trepostome bryozoan A. Fragment of the surface of a zoarium, X 18.

Fig. 8. Indeterminate trepostome bryozoan C. The only complete zoarium seen from above (fig. 8o)
and below (fig. %b), X 18.

Figs. 9, 10. Indeterminate bifoliate cryptostome bryozoan A. 9, A fragment of an old zoarium,
showing rounded zooecial apertures, and zooecia covered with surface sculpture, X 18. 10, A frag-
ment of a young zoarium, X 18.

Fig. 11. Indeterminate bifoliate cryptostome bryozoan B. Fragmentary zoarium, Xl8.

Fig. 12. Orbigjiyella /aw//o5a (Phillips). Half of a zoarium seen from above (fig. I2b) and below
(fig. \la), xl8.

Palaeontology, Vol. 6

PLATE 37

10 “'

■rf^C-

JKfffJf

• -

SPJELDN^S, Silicified Ordovician fossils

NILS SPJELDN^S: SILICIFIED ORDOVICIAN FOSSILS

261

diameter. The zooeeia are similar to species B in shape, they are generally slightly larger,
and have more mesopores.

The hollow tube might either be a part of the zoarium, formed by the bryozoan, or it
might be the tube filled by an organism encrusted by the bryozoan. Both cases are
well known in Ordovician bryozoans.

Order cryptostomata
Indet. Bifoliate Cryptostome A
Plate 37, figs. 9-10

Nine fragments are known of this species. Young parts of the zoaria are sharp-edged,
the older ones blunt-edged, about 1 -5 mm. wide. The thickness also increased with age.
There are five to seven rows of zooecial apertures, which are arranged diagonally. They
are elongate, drop-shaped in the young, and more regularly circular in the old ones.
Along the margin of the branches and between the zooeeia there are longitudinal ridges.
In some older specimens (PI. 37, fig. 9) this sculpture covers some of the zooeeia.

The considerable variation found in this species can be attributed to the different age
of the fragments, such variation is known within a single zoarium both of fossil and
recent bryozoans. The marked sculpture recalls Artbropora (= Graptodichtya), but this
cannot be confirmed without thin sections of better-preserved material.

Indet. Bifoliate Cryptostome B

Plate 37, fig. 11

This species, which is found only in two small fragments, differs considerably from
species A. The zooecial apertures are larger and more regularly arranged in diagonal
rows, there are no peristomes, and no sculptured edge of the branches, and no sculpture
between the zooecial apertures. The species recalls certain species of Escharopora, but
also in this case it is not wise to refer it to a genus without thin sections.

Arthrostylid cryptostomes
Plate 37, figs. 1-5

The arthrostylid bryozoans are second in number only to the flat, discoid trepostomes.
They occur in a variety of types having 3, 4, 5, or even 6 rows of zooeeia, and a number
of different sculptures, and sizes of zooecial apertures.

A number of new genera and species have been founded on such fragmentary silicified
material (cf. Bassler 1953, pp. G128-G130), but the author has no wish to follow this
procedure. Studies, on both fossils and recent species with fine-branched jointed zoaria,
show that considerable variation is found in numbers of zooecial rows and in sculpture
in different parts of the same zoarium (cf. Marcus 1940, text-figs. 97, 113). Some of these
species also have primary, secondary, and tertiary branches which are rather different,
especially in number of zooecial rows, and in length of segments.

Nevertheless, the present material shows such a variation in sculpture and size of
zooeeia that it must be assumed that more than one species is present. For the reasons
stated above, these species remain unnamed because of the scanty and incomplete
material at hand.

262

PALAEONTOLOGY, VOLUME 6

Other bryozoans. Besides those mentioned here, there are a number of others in the
material, chiefly very fragmentary trepostomes. One of them has very oblique zooecial
apertures, and resembles Bythopora.

THE AGE OF THE FAUNA

Judging from the presence of Marrolithus inflatus incipiens, and the trilobite and
brachiopod fauna which accompany this subspecies, the age of the fauna would be
either Upper Llanvirn {D. murchisoni Zone) or Lower Llandeilo. The stratigraphy of
these forms are well known from the type Llandeilo district through the papers of
Williams (1948, 1950) and earlier authors.

The ostracod and bryozoan fauna described here resemble faunas generally found in
Caradoc beds. Lomatobolbina is not recorded below the Crassicauda Limestone (upper-
most Llandeilo), and has its main distribution in the Caradoc. TaUinnella has been
reported from Lower Ordovician onwards, but the Bohemian section of the genus, to
which T. complicata belongs, is not elsewhere reported from beds older than the Caradoc.
Conchoprimitia, on the other hand, is mainly a Lower Ordovician genus. The ostracod
faunas known from the Scandinavian Llanvirn and Lower Llandeilo are strikingly
different from that described here (Henningsmoen 1953Z7, Jaanusson 1957).

The bryozoan fauna, although it cannot be identified as to species and genus, resembles
in general pattern those found in the Caradoc beds of Scandinavia, Estonia, and eastern
U.S.A. (especially the Edinburgh Formation of Virginia). The resemblance with the
latter might partly be due to facies and preservation since the arthrostylids, which are so
prominent in many silicified bryozoan faunas of this age, are easily overlooked in other
preservations. The few known bryozoan faunas of this age (Oil Creek Formation in
Oklahoma, and Kanosh Shale in Utah) have a quite different set of species, which are
considerably more primitive-looking and resemble the Lower Ordovician fauna.

The apparent difference from other early Ordovician faunas is perhaps because very
few bryozoan and ostracod faunas of Llanvirn-Llandeilo age are well known, and it is
quite possible that the picture of faunal distribution will have to be changed when
material from other facies and areas are studied.

A check of the exact correlation between the type Llandeilo and the standard graptolite
succession might, however, be advisable in light of the new material presented here.

REFERENCES

BASSLER, R. s. 1953. Bryozoo, In moore, r. c. : Treatise on Invertebrate Paleontology. Part G. 1-253.
Lawrence, Kansas.

harper, j. c. 1947. (Salter) and some Caradoc species of the genus. Geol. Mag.

84, 345-53.

HENNINGSMOEN, G. 1953fl. Classification of Paleozoic straight-hinged Ostracods. Norsk Geol. Tidsskr.
31, 185-288.

1953^. The Middle Ordovician of the Oslo Region, Norway. 4, Ostracoda. Ibid. 32, 35-56.

JAANUSSON, V. 1957. Middle Ordovician Ostracodes of Central and Southern Sweden. Bull. Geol.
Inst. Univ. Uppsala, 37, 173-442.

MACGREGOR, A. R. 1961. Upper Llandeilo brachiopods from the Berwyn Hills, North Wales. Palaeonto-
logy, 4, 177-209.

MARCUS, E. 1940. Mosciyr. In Danmarks Fauna, 46, \-40l. Kobenhavn.

NILS SPJELDN/ES: SILICIFIED ORDOVICIAN FOSSILS 263

OPiK, A. 1937. Ostracoda from the Ordovician Uhaku and Kukruse formations of Esthonia. Publ.
Geol. Inst. Univ. Tartu, 50, 1-74.

SPJELDN/ES, N. 1951. Ontogeny of Beyrichia jonesi Boll. Jouni. Paleont. 25, 745-55.

1957. A redescription of some type specimens of British Ordovician bryozoa. Geol. Mag. 94,

364-76.

STORMER, L. 1930. Scandinavian Trinucleidae. Skr. Vid. Akad. Oslo, I Mat.-Naturv. Kl. 4, 1-111.

1959. The Double Cardinal Process in the oldest Strophomenids. Norsk Geol. Tidsskr. 39, 13-23.

STRAHAN, A., CANTRILL, T. C., DIXON, E. E. L., THOMAS, H. H. and JONES, O. T. 1914. The geology of
the South Wales Coalfield, part XL The country around Haverfordwest. 1-262. Mem. Geol.
Survey.

WHITTINGTON, H. B. and EViTT, w. R. 1953. Silicificd Middle Ordovician trilobites. Mem. Geol. Soc.
America, 59, 1-137.

WILLIAMS, A. 1948. The Lower Ordovician cryptolithids of the Llandeilo District. Geol. Mag. 85,
65-88.

1953. The geology of the Llandeilo District, Carmarthenshire. Quart. Journ. Geol. Soc. London,

108, 177-208.

NILS SPJELDN/ES
Institutt for Geologi,
Blindern,

Oslo,

Manuscript received 28 May 1962 Norway

THE BRITISH CARBONIFEROUS SPECIES OF
GIRVANELLA (CALCAREOUS ALGAE)

by ALAN WOOD

Abstract. The type specimens, or topotypes, of the known species of Girvanella from the Lower Carboniferous
rocks of Britain are redescribed. The two species found in the Girvanella Band in northern England are identical
with those described by Wethered from the Avon Gorge, whieh occur at a very similar horizon. It is shown that
the type of preservation and the amount of decay before burial affects appearances of the specimen in thin
section, blurring specific characters. Various growth-forms of a species may be recognized, but such forms do
not seem to be of value for stratigraphic work.

The best-known occurrence of Girvanella is in the Carboniferous Limestone of northern
England, where Garwood (1912, 1924) found algal nodules to be useful as horizon
markers at a level which he took as the base of the Dg zone. Though this Girvanella
Band has been recorded over hundreds of square miles of north-western England, and
bands have been found at a similar horizon in Derbyshire and in Northumberland, no
specific names have been given by British authors to the forms occurring there, nor have
their ranges in time or space been ascertained. This is due in part to the inherent diffi-
culties of identification of such a featureless fossil, but even more to uncertainty as to
the exact characters of the already named species. Garwood himself (1931) when
describing a new species, G. staminea, from Ci beds at Bewcastle, could only refer to the
types found in the typical Girvanella Band as ‘showing two sizes of threads’, with a
reference to his two figures o^ " Girvanella sp.’’ (1924). The position will be made easier
in the future by the discovery of the type specimens of the species described by Wethered
(1890) from the Avon Gorge, which were contained in a large collection of thin sections
given by the late Judge Wethered to the University of Bristol. These were kindly put at
my disposal by Professor W. E. Whittard. Though the original drawings did not show
all the features which might be useful for specific identification, they were very accurate
representations, and it has proved possible to pick out all the figured specimens of both
species.

Perusal of Wethered’s paper will show that he was chiefly concerned with the origin of
oolitic granules. Already (1889) he had demonstrated that the pisoliths of the Pea Grit
in the Inferior Oolite of Gloucestershire were of organic origin, and in 1890 he was con-
cerned with both Jurassic and Carboniferous rocks, Girvanella being considered to be the
main agent in the formation of ooliths. It seemed possible, he thought, to trace a series
from clear examples of Girvanella, through those ooliths in which the Girvanella structure
was in process of being destroyed, to ooliths with only a trace of organic structure, the
end-member being perhaps those with a regular radial crystalline arrangement. This pre-
occupation with the origin of ooliths has introduced peculiar difficulties in the choice of
a type specimen for G. diicii.

In these delicate species of Girvanella accurate comparisons can only be made on
enlarged photographs, repeatedly checked under the microscope. A uniform enlarge-
ment of 450 diameters has been used. When whole-plate photographs are held side by

[Palaeontology, Vol. 6, Part 2, 1963, pp. 264-73, pi. 38-40.]

ALAN WOOD: BRITISH CARBONIFEROUS GIRVANELLA

265

side, comparison is rendered easy. It becomes clear that there are a number of different
growth habits among species whose internal and external diameters appear to be the
same. When the eye can hardly observe these in thin section and the memory cannot
retain the details while one slide is changed for another on the microscope stage, the
practical value of these subtle differences almost vanishes. If more species were erected,
based on those features which have become familiar to the writer through a long period
of observation they would be unidentifiable by other workers, except when a full range
of species was available for study. Consequently the present observations are limited to
named species or figured specimens.

SYSTEMATIC DESCRIPTIONS

Girvanella wetheredii Chapman 1908 (= G. incnistans Wethered non Bornemann)

Plate 38, fig. 1 ; Plate 39, fig. I

Diagnosis. Flexuous, winding, interlaced tubes, not observed to taper, wrapping around
a central foreign body, perhaps also free. External diameter, measured on circular cross-
sections, 0-013 mm. to 0-015 mm., internal diameter ranging between 0-006 and 0-009 mm.
Closeness of packing variable; wall dark, fine-grained, thickness generally about
0-003 mm.; septa not observed. Branching occurring at a rather wide angle; sometimes
the branches come off close together from the same side of the parent thread. In one
case the branch bends round rapidly to grow subparallel with the others, in a manner
reminiscent of Ganvoodia.

Observations. Bornemann (1886) described the genus Siphonema, type species (here
chosen) S. incrustans, from a pebble of Silurian limestone found in Pleistocene glacial
drift and presumed to be from the Baltic area. This genus, as pointed out by Hinde
(1887), was a synonym of Girvanella, so that G. incrustans of Wethered (1890) was
invalid. Chapman (1908) noticed this and proposed the name G. wetheredii to replace it.

The original figures of this species were of the same specimen at different magnifica-
tions, one (la) being reversed in reproduction. The slide in which the type specimen is
contained has the locality ‘B below New Rd.’ engraved on the glass, and the labels read
‘Carboniferous M. L. Clifton’ and ‘No. 1. B. Below New Rd. Dolomite’. The rock is
a fragmental foraminiferal limestone, the matrix being of granular interlocking crystals
of calcite whose grain size ranges between 0-01 and 0-05 mm. Scattered in the granular
matrix are organic fragments ranging in size from 0-02 mm. to foraminifera 0-6 mm.
across, and even larger fragments of brachiopods and gastropods. The algal growths
occur around circular or slightly elliptical bodies about a millimetre in diameter, which
are probably sections of Productid spines. These are hollow, with transparent fibrous
walls which show a black cross under crossed nicols, and their centres are filled with
matrix. A considerable amount of finely disseminated pyrite occurs in the algal growth,
particularly near the nucleus, and this has invaded the material of the spine itself. Owing
to the thickness of the section, the tubuli of G. wetheredii are difficult to make out, but
they seem to have been continuous all round the nucleus, though dense structureless
calcite intervenes between the nucleus and the visible threads. Similarly, tubes are diffi-
cult to discern in the outer layers. The whole gives an impression of a nodule formed by

266

PALAEONTOLOGY, VOLUME 6

growth of various creatures around a common centre, some of which left no trace of
their presence beyond a dense precipitate. Around the nodule a narrow discontinuous
zone of clear calcite occurs, in continuity with a vein penetrating it, as if the nodule had
shrunk slightly away from the surrounding matrix and the gap had been filled with
material from solution.

One small portion of this algal nodule was figured at a higher magnification by
Wethered, and is the portion in which the threads can be most clearly seen. It is figured
in Plate 38, fig. 1, and is here chosen as the type specimen. The tubuli are more closely
packed than in the other examples described later, and are confusedly interwoven.
Probably they are cut parallel to the surface on which they grew, and they are seen to
branch as they spread over it. Direct measurement on circular cross-sections of threads
gives internal diameters varying from 0-006 mm. to 0-009 mm., with the average nearer
the latter figure. The outer limits of the walls are extraordinarily difficult to determine
owing to the slice not having been ground thin enough, but the two largest cross-sections
approach 0-015 mm. in diameter while some are smaller, down to perhaps 0-012 mm.
The photograph, which has not been touched up, but whose contrast has been increased
by every possible means, gives an impression of clear definition not borne out when the
specimen is observed by eye under the microscope.

Two other slides, both engraved with the words ‘New Rd. Oolite’, are almost certainly
from the same bed, since they match the type slide in grain size, foraminiferal content,
and especially in the algal growths being around identical nuclei. Since the slides are
thinner the tubuli are more clearly seen, and an enlarged figure (PI. 39, fig. 1) is given
here. All the measurements discussed below have been taken from these two slides. In
one case G. wetheredii occurs in contact with the nucleus, and in another it reaches the
outer surface of the nodule. In all cases it seems to have had a general encrusting habit
and to have taken a real part in the growth of the nodule. Layers of dense calcite devoid
of tubuli are constantly present, as in the type specimen. The nucleus of one nodule is
a piece of fragmental limestone with a clear crystalline matrix different from that of the
rest of the section.

When enlarged photographs were studied it was found that cross-sections showing
truly circular interiors were more rare than had been anticipated, most sections showing
irregularities due probably to partial collapse of the external mould after death. Circular
cross-sections showing the thickness of the wall were even rarer, usually one side or
another being in contact with and blending into the dense matrix. It was interesting to
note that measurements of internal tube diameter on longitudinal sections were on the
average slightly over 0-001 mm. less than those measured on circular cross-sections,
while measurements of the external diameter measured in the same way were in error
by nearly 0-002 mm. Presumably Wethered made his measurements on longitudinal

EXPLANATION OF PLATE 38

Fig. 1. Girvanella wetheredii Chapman. Type specimen (here chosen). Branching can be seen in three
places. Upper Di, Avon section, Bristol. Geological Survey Collection, GSM. PF1923.

Fig. 2. G. staminea Garwood. Type specimen. Tubuli indistinct for reasons discussed in text. Ci,
White Beck, Bewcastle, Cumberland. British Museum (N.H.) Collection, V43735.

Fig. 3. G. ottonosia Pia. Topotype. Branching visible near the centre of the photograph. Km, Avon
section. Geological Survey Collection, GSM. PF2047.

Palaeontology, Vol. 6

PLATE 38

WOOD, Girvanel/a

ALAN WOOD: BRITISH CARBONIFEROUS GIRVANELLA

267

sections, since he stated that the diameter was 0-01 mm. The thickness of the wall varied
from rather over 0-0015 mm. to 0-003 mm., the latter being nearer the average figure.

The two slides from the Wethered collection show slightly different growth habits or
types of preservation. In one the tubes remain for relatively long distances in the plane
of the section. In the other, cross-sections of the walls are more obvious, and the impres-
sion of free and rapid growth parallel to the surface of the nodule is less strong. The
tubes also appear to be more contorted. There is no doubt that both specimens are from
the same level, perhaps from the same hand specimen, for the reasons given above.
A similar range of variation in appearance can be seen in specimens from the Girvanella
Band at Hull Pot, Yorkshire, in the Garwood collection in the British Museum, Natural
History (Slides V43730 and 157B). The difference in appearance probably depends in
part on a small variation in thickness of the thin section, but also is likely to depend on
an original slight difference in the position of the threads with respect to the angle at
which light fell on the algal nodule and also, perhaps, to the influence of competing
organisms. In other cases, the threads are well separated and even more contorted. This
is well seen in the specimen figured by Garwood in 1924 from Hull Pot in which the
tubes are rendered conspicuous by a dark coating, and their contortion and the degree
of separation are noteworthy. It seems likely that this specimen is conspecific with
Wethered ’s specimens, since similar variations in degree of contortion have been
described in Girvanella problematica itself (Wood 1957).

As stated in the introduction, G. wetheredii has not been positively identified since
Wethered’s time, though in 1932 Pia, who had made collections from the Carboniferous
Limestone of England, stated that in his opinion G. staminea Garwood was identical
with ‘G. incrustans\

Type specimen. Figured on Plate 38, fig. 1 , deposited in the Geological Survey Museum, South
Kensington, GSM. PF1923.

Locality. A bed of dark nodular limestone, with argillaceous material, 9 inches thick, immediately
below an oolitic bed 10 feet thick, a short distance below the junction of the road from Clifton with
that along the banks of the River Avon. Upper D1 (Reynolds 1921, pp. 233-4). National Grid
Reference 31/563734.

Girvanella ottonosia Pia 1937
Plate 38, fig. 3

Diagnosis. Highly contorted flexuous tubes, circular in section, forming a pelhcle around
a foreign body with a digitate Oltonosia-\ik& outer surface. External diameter of tubes
usually about 0-007 mm., internal diameter 0-004-0-005 mm., wall thickness 0-001-
0-002 mm. Occasionally one tube may be as much as 0-007 mm. in internal diameter,
but in general the threads are remarkably uniform in size. Branching apparently dichoto-
mous, but contorted, usually at an angle of less than 40°.

Observations. The specimens are preserved in a fragmental limestone, consisting of
crinoid ossicles and plates, broken gasteropods, polyzoa and lamellibranch shells,
enclosed in a granular calcite matrix. Brachiopods are apparently absent. The matrix is
of two types, one yellowish in transmitted light, probably nearly in its original state
except as modified by grain growth, average grain size 0-02-0-03 mm., and a clearer
more translucent portion irregularly dispersed around and among the organic fragments.

C 1213

T

268

PALAEONTOLOGY, VOLUME 6

This is slightly coarser and probably represents a cavity infilling. The range of size of the
organic fragments is from 0-2 mm. upwards to 7 mm., the size of the largest algal nodule.
The rock was probably rapidly deposited as a mass of transported fragments in a cloud
of fine silty calcite which partially filled the interstices between the fragments.

The algae are preserved as thin coatings on polyzoa and around recrystallized lamelli-
branch (?) shells. The coating ranges in general from 0-3 to 0-5 mm. in thickness
(maximum T4 mm.), and displays the digitate outer margin described by Pia. Sections
parallel to the surface of the alga show that the projections are irregularly peg-hke,
rather than parallel-sided wrinkles. The spaces between the outgrowths were certainly
present before burial; they are filled generally by the yellowish original matrix described
above and occasionally by the later clear drusy grains. The algal growth must have been
quite stiff and crisp to withstand transport and to retain this surface appearance. There
is not sufficient evidence that the algal threads originally grew parallel to the present
outer margins of the projections, though occasional portions of a section suggest this.
On the other hand the interstices do not have the form of later burrows which had
destroyed part of a once continuous coating. The absence of growth parallel to the
margins may be due to the former presence of a living, uncalcified layer, which decayed
before fossilization; certainly the whole series of outgrowths gives the impression of
being an original feature.

Locality. Lower Limestone Shales, Km, Avon Gorge (Clifton Side), Bristol.

Material. Four slides from the Wethered Collection, labelled ‘Lower Limestone Shales, Clifton,
Bristol’, or some variant of this. Figured specimen in the Geological Survey Museum, GSM. PF2047.

Girvanella staminea Garwood 1931
Plate 38, fig. 2; Plate 39, fig. 2

Diagnosis. Flexuous winding interlaced tubes, not observed to taper, wrapping around
a central foreign body. External diameter, measured on circular cross-sections, 0-011-
0-013 mm. internal diameter 0-006-0-008 mm. Closeness of packing variable, wall dark,
fine-grained, thickness about 0-003 mm., septa not observed. Branching occurring at
a wide angle.

Observations. The type specimen forms part of a single composite algal nodule, without
matrix, showing a well-preserved specimen of Garwoodia gregaria (Nich.) enveloped
by a thin layer of Girvanella threads. These extend in varying states of preservation
nearly around the central spherical alga, being separated at intervals by thin layers and
irregular blebs of clear calcite, often showing traces of organic structure. This Girvanella
layer is in turn surrounded by another organically deposited portion which forms the
outer part of the nodule. In this there are a variety of encrusting organisms, worm tubes.

EXPLANATION OF PLATE 39

Fig. 1. Girvanella wetheredii Chapman. Topotype. To show growth habit. Compare tube-diameter
and growth habit with G. staminea (below). Upper Di, Avon section, Bristol. Geological Survey
Collection, GSM. PF1925.

Fig. 2. G. staminea Garwood. To show tube size, branching, and general growth-habit. Compare
with G. wetheredii (above). ? C,, Cambeck, Spadeadam, Cumberland. British Museum (N.H.)
Collection, V43734.

Palaeontology, Vol. 6

PLATE 39

WOOD, Girvanella

ALAN WOOD: BRITISH CARBONIFEROUS GIRVANELLA 269

‘bean-shaped organisms’, and a few small, nearly stifled layers of Garwoodia. The nodule
was thus formed in a period to be measured in months rather than weeks. Growth of
the Girvanella layer, itself only an insignificant portion of the whole, was interrupted by
colonization of other organisms or by boring creatures or both, which gave rise to the
clear patches described above.

Though the slide is labelled ‘too thick’ in Garwood’s handwriting, the polarization
colours of the clear calcite infilling of the Girvanella tubes are bright, and the indefinite
appearance and lack of character of the algal tubes must be ascribed to partial decay
before lithifaction. The nodule must have lain on the sea floor for some time, and
colonization by other creatures would affect the preservation of this specimen if only
by the penetration of their basal cells. Much, if not all, of the indefinably different
impression this specimen gives is to be explained in this manner.

The original diagnosis is: ‘This species is distinguished from other Carboniferous
species previously described by the minute size of its tubes, which measure 0-006 mm.
in diameter.’ It is certain that Garwood made his measurements on the bright circles
which represent infilled cross-sections of the tubes, and thus give the internal diameter
of the threads. All previous measurements of species of Girvanella had been made on
the threads as seen in longitudinal sections, including the thickness of the wall. Because
of the close packing of the threads in this specimen, and because of subsequent changes,
longitudinal sections are always indefinite. A few cross-sections of the whole thread can
be distinguished; and the measurements cited above fall into the range of variation of
G. wetheredii, as do the measurements of internal diameter.

Nevertheless, other better-preserved material from north Cumberland contained in
the Garwood collection seems to show that a species with slightly slimmer threads than
G. wetheredii is characteristic of this horizon in Cumberland. The specimen figured on
Plate 39, fig. 2 is from Cambeck, Spadeadam, north Cumberland, and is said by Gar-
wood to come from a horizon in C2. Reference to Garwood’s map (1931) will show
that this locality is only just above the Main Algal Series, from which the type specimen
of G. staminea was obtained. In the Cambeck specimen well-preserved Girvanella threads
closely envelop a Garwoodia nodule, and are enclosed in a fine-grained bioclastic lime-
stone containing occasional tiny rounded quartz grains. They closely resemble the type
specimen in appearance, growth, habit, and even in the ecological niche they have chosen
to inhabit. The preservation is good, the threads are clear, and branching can be seen.

Comparison with the figures of G. wetheredii shows that the average diameter of the
threads is slightly less, though the growth-habit is similar. Such a difference may be due
to a differing environment, but it seems more probable that there was a real, though by
now almost inappreciable, distinction between the C^ and Cj species and that charac-
teristic of the upper part of the and base of the D2 zones.

Type specimen. Figured on Plate 38, fig. 2, deposited in the British Museum (N.H.), V43735.

Locality and horizon. ‘Main Reef’, Main Algal Series; White Beck, Bewcastle, Cumberland; Ci.

Girvanella ducii Wethered 1890
Plate 40, figs. 1-2

Diagnosis. Gently flexuous tubes, generally loosely interwoven or not in contact, some-

270

PALAEONTOLOGY, VOLUME 6

times closely wrapped around a foreign body. External diameter, measured on circular
cross-sections, ranging from 0-026 to 0-033 mm., average 0-029 mm., internal diameter
ranging from 0-015 to 0-020 mm., average 0-018 mm. Wall dark, fine-grained, thickness
generally about 0-006 mm., branching irregularly dichotomous, at a varying angle. No
cell-walls observed.

Observations. The original figures of G. ducii were from three different thin sections and
two localities. I here choose the original of Wethered’s figure 2a as the type; this
specimen is figured on Plate 40, fig. 1. It occurs in a fragmental limestone, in which the
majority of organic fragments have a pellicle of algal or inorganic radial fibrous coating
up to 0-1 mm. thick. The fragments are often rounded and range in size from 0-05 mm.
to nearly 1 mm. in diameter, the latter being rare. The average grain size is about 0-4 mm.
Some foraminifera, in particular, have no precipitated layer around them. The fragments
are closely packed, but rarely in contact, and the matrix is transparent and granular,
with an average grain size of about 0-01 mm. The minute nodule containing G. ducii is
the largest fragment in the slide, measuring 2-8 mm. in longest diameter. The margins
are dark and composed of fine-grained ‘ algal dust ’ in which threads of G. ducii occur,
while the centre is occupied by clear calcite similar to the matrix and containing similar
murky ooliths and organic fragments. In the clear calcite near the centre, tubuli of G.
ducii are common, and are not so closely packed as in the less transparent margins.
There is no suggestion of a concentric arrangement of the threads, but this may be due
to the nodule being irregular in shape and excentrically cut. This suggestion is supported
by the presence of ooliths in the apparent centre of the nodule. Branching is not seen
in this specimen.

Wethered stated that the diameter of the tubuli was 0-02 mm. Measurement on his figure
2a gives a diameter of 0-03 mm. for the external diameter of the larger cross-sections,
a figure which is substantiated by the present measurements (see above). It is, however,
particularly noticeable that the threads seem to be of greater external diameter when
they are not touching, and are surrounded by clear calcite. This fact has not affected
the measurements given in the diagnosis above, since it happens that only one cross-
section occurs in this region, but it is very noticeable when measurements are made on
longitudinal sections of threads. These measurements in the dense closely packed regions
average 0-027 mm. for external diameters, while in the clear area with loosely packed
threads, the average is slightly above 0-030 mm., with a maximum of 0-035 mm. In each
case the threads were measured at the point of greatest diameter and greatest interior
brightness, this being presumed to indicate nearness to a median section. The difference
is probably due in the main to the outermost boundaries of the walls being easily visible
in the areas of clear calcite, while they are taken to be part of the matrix in the more
crowded regions.

The second specimen figured by Wethered (fig. 2b) shows rather indistinct tubuli

EXPLANATION OF PLATE 40

Figs. 1-2. Girvanella ducii. Wethered. 1, Type specimen (here chosen). Tubes winding loosely in
a fine-grained limestone with cavity infillings. Upper Dj, Avon section, Bristol. Geological Survey
Collection, GSM. PF1924. 2, Specimen from Girvanella Band, Yorkshire, to show growth-habit and
tube diameter identical with the Bristol specimen. Base of D2, Hull Pot, Pen-y-Ghent, Yorkshire.
British Museum (N.H.) Collection, V44935.

Palaeontology, Vol. 6

PLATE 40

WOOD, Girvanella

ALAN WOOD; BRITISH CARBONIFEROUS GIRVANELLA 271

partly enveloping a compound nucleus. The internal diameter of the tubuli is between

0 01 7 mm. and 0-020 mm., and they are surrounded by a dense structureless ‘algal dust’.
The largest diameter of the whole granule is 0-7 mm. The chief interest of this specimen
is that it shows that G. diicii could grow concentrically around a nucleus and envelop it,
as do other species of Girvanella. The tubes are markedly less flexuous than in G. wetheredii,
and they lie at one side of the nodule only. This may well be due to attrition before
fossilization.

The original of Wethered ’s figure 2c came from Tortworth Park, Gloucestershire, and
is an algal aggregate enclosed in radially disposed calcite crystals. It occurs in an oolitic
limestone in which every organic fragment is covered with a thin layer of radiating
crystals; the matrix is entirely recrystallized. It is clear that the fragment of Girvanella
is detrital and had been rounded before being incorporated into the sediment. Unfortu-
nately no circular cross-section of the threads is visible. However, there is no doubt that
the tubuli are markedly thinner than in the type specimen, measuring only 0-016 mm.
in external diameter on longitudinal sections, and they must belong to another species.
The internal diameter measured in the same way varies from 0-008 to 0-010 mm.

The choice of a type specimen is not entirely straightforward. On p. 272 of Wethered’s
paper a measured section of the New Road Oolite is given, six beds being listed. Con-
cerning the topmost bed, he stated ‘a form of Girvanella was here discovered to which

1 propose to give the name of Girvanella Ducii, for reasons to be presently explained.
It consists of an aggregation of tubules similar to those represented in PL XI, figs. 2a
and b; but in this instance the outlines are to a large extent obliterated by mineral
changes. . . .’ On page 273 he wrote, when describing bed 3, ‘In this bed we again meet
with the organism I propose to call Girvanella Ducii, occurring in loose aggregations
as represented in fig. 2a.’ At first sight it seems difficult to come to any other conclusion
than that the type specimen must be that first mentioned by Wethered, unfigured and
undescribed. However, when one considers his interest in the origin of ooliths, and the
fact that the measured section was described in logical order from above downwards,
it becomes apparent that Girvanella ducii was here referred to as an agent in oolith
formation, without any intention of making a type specimen from among the tubules
of Bed 1.

The specimen figured by Garwood (1924, pi. xix, fig. 2) from the Girvanella Band at
Hull Pot, Penyghent, is undoubtedly G. ducii. A similar specimen from the same locality
(B.M.N.H. slide V44935) is identical with the type specimen in dimensions, growth habit,
and branching. It is made up of a lax aggregation of tubes forming a small part of a typical
‘algal’ nodule, 15 mm. long, which shows a series of irregular concentric layers around
a fragment of a brachiopod shell. The whole is contained within a fragmental foramini-
feral and crinoidal limestone.

The fact that Wethered gave an incorrect diameter for the tubuli of G. ducii might
well have led to confusion, but fortunately no other species has been named from
Carboniferous rocks with a tube diameter of 29 ft. Indeed most authors, including
Maslov (1956), quote diameters of around 20 ft for the specimens they ascribe to G. ducii.
The growth-habit of the specimens from the Urals, and those figured by Dangeard (1948)
from Belgium is closely similar to that of typical G. ducii, and the difference in external
diameters requires explanation. Part of the difference may be explained by other workers
measuring tube diameter on longitudinal sections, rather than on cross-sections and

272

PALAEONTOLOGY, VOLUME 6

a part may be due to grain growth in the matrix of the limestone, whereby the outer part
of the calcareous envelope has been taken over and incorporated in the crystals around.

G. liebusi Paul is stated to have a tube diameter of 20 jji. It is interesting to note, in
view of the remarks given above concerning the preservation of G. ducii, that Paul (1938)
stated that his species with elongated weakly curved threads did not build nodules, and
thus is separated from other forms of Girvanella, though the threads were individually
identical with those of Girvanella so that a generic distinction was not possible. Later
in 1938 he added that this alga was found in a coarsely crystalline limestone with
foraminifera, a habitat that algae usually avoid. In this respect too it is similar to G. ducii
and the limestone figured does not look any coarser than the beds in which G. ducii
usually occurs. In view of these facts, and the incorrect magnification given by Paul for
the type figure (the true magnification is around 25 diameters) it is possible that G. liebusi
is synonomous with G. ducii.

Both Dangeard (1948) and Maslov (1935) mention the occurrence of cell walls
transverse to the tubes, in specimens they consider to belong to this species and Dangeard
gives a fine photograph of these. Similar features are occasionally seen in specimens from
Britain, but observation is very difficult at the magnifications that have to be employed,
and the present author has not been able to feel certain that they are not simply the
boundaries of the relatively more clear crystals filling the interior of the tubes.

Though this form has frequently been referred to by other authors, for example Pia
(1932, 1937) and Johnson (1945), the original incorrect statement concerning the tube
diameter necessitates a re-examination of the evidence in each case.

Tvpe specimen. Figured on Plate 40, fig. 1, deposited in the Geological Survey Museum, GSM.
PF1924.

Locality. A bed of blue limestone with some oolitic spherules, 2-5 feet thick, lying immediately below
the thin dark limestone with G. wetheredii. Upper Dj.

Girvanella nicholsoni (Wethered) 1886

This appears to be a rare species, and there is nothing new to add to the description
given by Wood (1941).

NOTE ON THE ‘GENUS’ GIRVANELLA

Johnson (1946) considered the present author’s use of the generic name Girvanella for
so coarse a species as G. n/c/to/wn/ ‘unfortunate’. There can be no doubt that the various
species included above in one genus are not congeneric. G. ottonosia Pia, for instance,
with strongly contorted minute tubuli and peculiar growth habit is very different from
G. ducii Wethered. The latter does, however, have many characters in common with
G. problematica Nich. and Eth., the type species. At some time in the future it will be
necessary to split off the forms with highly contorted minute threads as a separate genus,
and at that time the form with very wide tubuli now called G. nicholsoni could con-
veniently and logically resume the name Mitcheldeania, at present suppressed as a
synonym.

REFERENCES

BORNEMANN, j. G. 1886. Die Verstcinerungen des Cambrischen Schichtensystems der Insel Sardinien.
Nova Acta Kgl. Leap. Carol. Deiitsch. Akad. Naturf. 41, 1-83.

ALAN WOOD: BRITISH CARBONIFEROUS GIRVANELLA 273

CHAPMAN, F. 1908. On the relationship of the genus Girvanella, and its occurrence in the Silurian
Limestones of Victoria. Rep. Australasian Assoc. Adv. Sci. 1907, 377-86.

DANGEARD, L. 1948. Contribution a I’etude des genres Girvanella et Spbaerocodiiim. Bull. Mus. Roy.
Hist. Nat. Belg. 24, no. 2, pp. 1-3.

GARWOOD, E. j. 1912. The Lower Carboniferous succession in the north-west of England. Quart. J.
Geol. Soc. London, 68, 449-582.

1931. The Tuedian Beds of northern England and Roxburghshire, east of the Liddel Water.

Ibid. 87, 97-157.

and GOODYEAR, E. 1924. The Lower Carboniferous succession in the Settle District and along

the line of the Craven Eaults. Ibid. 80, 184-271.

HiNDE, G. J. 1887. Review of Bornemann (1886). Geol. Mag. lA, 226-9.

JOHNSON, J. HARLAN 1946. Lime-secreting algae from the Pennsylvanian and Permian of Kansas.
Bull. Geol. Soc. Amer. 57, 1087-1119.

MASLOV, V. p. 1935. Some Palaeozoic calcareous algae of the Southern Urals. Trans. All-Uiuon Inst.
Min. Prod., 72. (Not seen, quoted from Maslov 1956.)

1956. Eossil calcareous algae of the U.S.S.R. Proc. Inst. Geol. Sci., Acad. Sci. U.S.S.R., 160.

(In Russian.)

PAUL, H. 1938. Unterkarbonische Kalkalgen und Calcisphaeren Deutschlands. Jb. Preuss. Geol.
Landes f. 1937, 58, 276.

19386. Girvanella liebusi Paul in der unteren Dibunophyllum-Zone des Velberter Sattels.

Palaeont. Z. 20, 317-18.

PiA, J. 1932. Die Girvanellen des englischen Kohlenkalkes. Anzeiger Akad. Wiss. Wien, M. N. Klasse,
68, 94-99.

— — 1937. Die wichtigsten Kalkalgen des Jungpalaeozoikums und ihre geologische Bedeutung.

Deiixieme Congres Etudes Strat. Carbonifere, Heerlen, 2, 765.

REYNOLDS, s. H. 1921. The lithological succession of the Carboniferous Limestone (Avonian) of the
Avon section at Clifton. Quart. J. Geol. Soc. London, 77, 213-45.

WETHERED, E. 1886. On the structure and organisms of the Lower Limestone Shales, Carboniferous
Limestone and Upper Limestones of the Lorest of Dean. Geol. Mag. 23, 529-34.

1889. On the microscopic structure of the Jurassic pisolite. Ibid. 26, 196-200.

1 890. On the occurrence of the genus Girvanella in oolitic rocks, and remarks on oolitic structure.

Quart. J. Geol. Soc. London, 46, 270-81.

WOOD, A. 1941. The Lower Carboniferous calcareous algae Mitcheldeania Wethered and Garwoodia
gen. nov. Proc. Geol. Assoc. 52, 216-26.

1957. The type-species of the genus Girvanella (Calcareous algae) Palaeontology, 1, 22-28.

ALAN WOOD

Department of Geology,
University College of Wales,

Manuscript received 8 August 1962 Aberystwyth

THE OSTRACOD GENUS NEOCYTHERE IN THE

SPEETON CLAY

by P. KAYE

Abstract. Six species of the genus Neocythere Mertens are described from the Barremian and Albian beds in
east Yorkshire. Three species are new, one of which is the earliest known occurrence of the genus. The genus
Centrocythere is regarded as a subgenus of Neocythere s.l.

The material which forms the basis for this paper is part of a large fauna from the upper
part of the Speeton Clay studied as the topic for a thesis at the University of Hull.

The bulk of the material is of Albian age and in some cases has been obtained from
inland exposures. The upper Red Chalk at Speeton is extremely hard, but the equivalent
strata inland is found to be softer and easier to break down. The species have, however,
all been recorded from Speeton, but are generally poorly preserved. Away from the coast
exposures are very rare and samples have only been obtained from South Cave near
Market Weighton (Grid Ref. 925326) and West Heslerton near Malton (Grid Ref.
913769).

The Albian strata at Speeton falls into two divisions : a thin series of grey-green marly
clays comparable to the Gault Clay of the south of England and a thick sequence of
Red Chalk. There is a gradual transition in colour and hme content between the two
divisions, and an exact boundary is hard to draw. At the base of the Gault Clay there
is a 6-inch seam of glauconitic clay and phosphatic nodules known as ‘The Greensand
Streak’. This seam is underlain by dark clays of Aptian age, and it has yielded fragments
of ammonites indicative of the reguJoris Subzone. Apart from belemnites, the Gault and
Red Chalk are almost barren of macrofossils, and consequently detailed zoning has been
impossible. The author, in his observations over the last five years, has found the litho-
logical section, published by C. W. Wright in Swinnerton’s Monograph of Lower
Cretaceous Belemnites, to be extremely sound.

All of the specimens are catalogued and deposited in the collection of the Department
of Geology at Hull (HU).

SYSTEMATIC DESCRIPTIONS

Family progonocytheridae Sylvester-Bradley 1948
Genus neocythere Mertens 1956

Type species. Neocythere vanveeni Mertens 1956.

This genus was erected by Mertens (1956) for forms similar to Cy there concentrica
Reuss, but having an amphidont hinge and an accommodation groove in the left valve.
There is a great deal of uncertainty about the interpretation of the forms related to
this group, and, until quite recently, all Cretaceous ostracoda having concentric orna-
ment and slight ventral tuminity were referred to Cythere concentrica Reuss. Mertens

[Palaeontology, Vol. 6, Part 2, 1963, pp. 274-81, pi. 41.]

P. KAYE: THE OSTRACOD NEOCYTHERE

275

has done extensive work on these forms and has redescribed Reuss’s types. Unfortu-
nately, the material was not well enough preserved, particularly the hinge structure, to
allow correct generic assignment, and the species, together with related forms named
for the first time, were left tentatively in the genus 'Cylhere\ Mertens did, however,
distinguish two new genera, Neocythere and Centrocy there, on the basis of the hinge
structure.

On examination of the hinge of species referred to the genus ‘Cythere\ it is found

BARREMIAN

APTIAN

ALBIAN

L.

M.

U.

L.

M.

u.

L -

M

U

Neocythere (N.)protovanveeni sp.nov.*

• •

N.(N.) vanveeni MERTENS 1956 -

N. (Physocythere) semilaeva sp. nov.^

N.(P) hleroglyphica sp.nov.-’^

N.(P) lingenensis MERTENS 1956 +

N.tCentrocythere) denticulata MERTENS

Neocythere sculpta (CORNUEU1846 °

...

Neocythere mertensi OERTLI 1958 °

"Cythere" favosa MERTENS 1956°

Cythere" semiconcentrica MERTENS °

Cythere" steghausi MERTENS 1956 °

Species of Progonocytheridae from the Speeton Clay and equivalent beds in N.W. Europe.
f= Speeton & N.W, Europe, “’= N.W. Europe only; Speeton only.

TEXT-FIG. 1

that they differ from Neocythere in being symmetrical merodont and in having no
accommodation groove. Though these differences do occur, it is not, in the author’s
opinion, sufficient to warrant generic separation. It has been shown on many occasions
that the hinge in ostracods is unreliable as a basis for higher taxonomic division, and
therefore to split this group of very similar ostracoda on that basis is considered unwise.
It is proposed, therefore, to include all these forms under the genus Neocythere s.l. and
to create new subgenera for the different hinge varieties. Neocythere s.s. is used for
forms having an amphidont hinge, whilst the new subgenus Physocythere is erected for
the forms grouped by Mertens under 'Cythere' and having a merodont hinge. On the
same basis, the genus Centrocythere, which differs from Neocythere s.s. only in the
nature of the anterior tooth in the right valve, is therefore included as a subgenus of
Neocythere s.l.

The ranges of the species of Neocythere found at Speeton together with related forms
are given in text-fig. 1 .

276

PALAEONTOLOGY, VOLUME 6

Subgenus neocythere Mertens 1956
Type species. Neocythere vanveeni Mertens 1956.

Diagnosis. A subgenus of Neocythere s.l. including forms similar to the genotype having
an inflated, ventrally tumid carapace. The hinge in the right valve consists of two
terminal crenulate teeth, between which is a locellate furrow deepened at its anterior end
to form a smooth circular socket. The left valve is complementary, and possesses a
distinct accommodation groove above the median element.

Neocythere {Neocythere) vanveeni Mertens 1956
Plate 41, figs. 23, 25

Cythere concentrica Dupper 1952, p. 106, pi. 4, fig. 14.

Neocythere vanveeni Mertens 1956, p. 205, pi. 12, figs. 12-1% \ pi. 14, figs. 100-2.

Material. Mounted specimens HU 17.C. 10. 1-10, from 1 foot above the Greensand streak. Middle
Albian at Speeton.

Measurements Length Height Total width

Male carapace 0-65 mm. 0-35 mm. 0-42 mm.

Female carapace 0-65 mm. 0-38 mm. 0-42 mm.

Description. Carapace rounded, inflated, ventrally tumid. Greatest height at one-third
length, greatest width at mid-length. Dorsal margin strongly arched in the left valve,
but straight with marked cardinal angles in the right valve. Ventral margin straight or
slightly convex, being obscured in lateral view by the prominent ventral tumidity.
Anterior margin rounded, slightly asymmetrical ventrally ; posterior margin forming a
blunt point at mid-height.

The surface is ornamented with concentric ribs, which parallel the anterior, ventral,
and posterior margins. These ribs are rounded and prominent ventrally, but elsewhere
are rather indistinct. There are three major concentric ribs, but many other subsidiary
ribs are enclosed within them. The ventral surface is crossed by several strong longitu-
dinal ribs. A low, rounded muscle node occurs just in front of the centre of the lateral
surface.

A narrow duplicature occurs with a steep drop into the inflated interior of the valve.
Inner margin and line of concrescence coincide. Radial pore canals short, straight, and
few in number. Normal pore canals numerous, concentrated along the crests of the
concentric ribs.

The hinge of the right valve consists of two bar-like terminal teeth divided into four
or five distinct lobes. These teeth are separated by a long, straight furrow, which is
coarsely crenulate and deepened at its anterior end, to form a smooth oval socket. In the
left valve there are two long, shallow, crenulate sockets, partially open to the interior
of the valve, and separated by a long, high, prominent median bar. This bar is denticulate
and often enlarged at its anterior end into a boss-like tooth. Above the median bar lies
a broad, shallow accommodation groove.

Sexual dimorphism is well marked, with the females being higher than the males.

Juveniles are common, and differ from the adult principally in the fact that the hinge
is symmetrical, merodont in all the instars, and also in the fact that the ornament is
much less strongly developed. Certain of the earliest moults appear smooth. A constant

P. KAYE; THE OSTRACOD NEOCYTHERE

111

feature of the hinge in all the instars is the presence of an accommodation groove in the
left valve. The muscle node is also more prominent in many of the moult stages.

Remarks. Neocythere vanveeni is easily recognized by the smoothly arched dorsal margin
in the left valve, and by its hinge. The amphidont nature of the hinge is not always
apparent in the left valve, but can always be seen in the right. The presence of the
accommodation groove is diagnostic.

The specimens found here are seen to be rather smaller than those found by Mertens
in Germany. It is therefore possible that no adult specimens have been found. The
largest forms are rare, and the bulk of the material found belongs to the early instars.
This fact is borne out by the poor sexual dimorphism shown, and the less strongly
developed hinge than the German forms.

Neocythere {Neocythere) protovanveeni sp. nov.

Plate 41, figs. 1-3, 5, 6

Holotype. A female right valve from 14 feet above the base of ‘Lower B’ (Lower Barremian) at
Speeton, no. HU 17.C.1L1.

Other watericil. Three paratypes mounted as HU 17.C.12.1-3.

Diagnosis. A species of Neocythere s.s. basically similar to N. vanveeni but having the
dorsal margin of the left valve less arched, and the posterior margin more bluntly
pointed.

Measurements Length Height Total width

Holotype female right valve, HU 17.C.1 1.1. 0-65 mm. 0-37 mm. 0-20 mm.

Paratype male left valve, HU 17.C. 12. 1. 0-67 mm. 0-41 mm. 0-20 mm.

Description. Carapace rounded, inflated, ventrally tumid. Greatest height at one-third
length, greatest width at mid-length. Dorsal margin of the left valve slightly arched, but
with a flattened central portion. Dorsal margin of the right valve straight, with marked
cardinal angles. Ventral margin convex, but obscured in its central portion by the
ventral tumidity. Anterior margin rounded ventrally asymmetrical; posterior margin
forming a blunt point at mid-height. Muscle node not well defined. Surface ornamented
with concentric ribs, which are most prominent ventrally and arranged as in N. vanveeni.
The dorsal third of the valve is, however, noticeably smooth. Internal features are
identical to N. vanveeni. Radial pore canals are short, straight, and number ten anteriorly
and four posteriorly. Normal pore canals are numerous and arranged along the crests
of the concentric ribs. Hinge identical to N. vanveeni, having the median element notiee-
ably subdivided, but with a deeper and narrower accommodation groove in the left valve.

Remarks. This species is very similar to N. vanveeni, to which it is almost certainly related.
It differs in the nature of the dorsal margin, in the width and depth of the aecommodation
groove in the left valve, and also in the shape of the posterior margin.

Subgenus physocythere subgen. nov.

Type species. "Cythere' lingenensis Mertens 1956.

Diagnosis. A subgenus of Neocythere s.l., showing the characteristic inflated ventrally
tumid shape of the genus, but differing from Neocythere s.s. in the structure of the hinge.

278

PALAEONTOLOGY, VOLUME 6

The hinge is symmetrical, consisting in the right valve of two strong, terminal, crenulate
cusps separated by a long, straight locellate groove. In the left valve there is no accom-
modation groove, but a wide shelf occurs above the median element sloping down to the
dorsal margin. The hinge elements as a whole are much stronger than in Neocythere s.s.

Neocy there (Physocythere) Hngenensis Mertens 1956

Plate 41, figs. 15-22, 24

'Cythere" costala Mertens 1956, p. 190, pi. 9, figs. 27-30; pi. 10, figs. 31-32.
non Cythere costata M‘Coy 1844, p. 165, pi. 23, fig. 11.

'Cythere' Hngenensis Mertens 1958, p. 359.

Material. Mounted specimens HU 17.C.13.1-7 from the Middle Albian at Speeton.

Measurements Length Height Total width

Female carapace 0-69 mm. 0-39 mm. 0-41 mm.

Male carapace 0-73 mm. 0-39 mm. 0-41 mm.

Description. Carapace egg-shaped, inflated, ventrally tumid. Greatest height at one-
quarter length, greatest width at mid-length. Dorsal margin straight or slightly concave ;
ventral margin convex. The tumidity is not strong enough to obscure the ventral margin.
Anterior margin broadly rounded, slightly asymmetrical ventrally; posterior margin
triangular, forming a blunt point at mid-height. A muscle node is not developed.

Ornament consists of a reticulate honeycomb-like network of ridges covering the
lateral surfaces. This network is bordered by three major concentric ribs paralleling the
anterior, ventral, and posterior margins. These are most strongly developed ventrally,
and often have the uppermost rib accentuated into a keel-like prolongation which forms
an ear-like projection when the valve is viewed dorsally. A series of longitudinal ribs

EXPLANATION OF PLATE 41

All figures X 50

Figs. 1-3, 5, 6. Neocythere {Neocythere) protovanveeni sp. nov., Barremian. All figured specimens from
14 feet above the base of Lower ‘B’ at Speeton. 1, Right valve female (holotype) lateral view
HU. 17.C.11.1. 2, Right valve female (holotype) internal view HU. 17.C.11.1. 3, Left valve male
(paratype) lateral view HU. 17.C.12.1. 5, Carapace (paratype) dorsal view HU. 17.C.12.2. 6, Cara-
pace (paratype) ventral view HU. 17.C.12.2.

Figs. 4, 7. Neocythere {Physocythere) semilaeva sp. nov.. Upper Albian, South Cave. 4, Left valve
(holotype) lateral view HU. 17.C.16.1. 7, Carapace (paratype) from right HU. 17.C.17.1.

Figs. 8-12, 14. Neocythere {Physocythere) hieroglyphica sp. nov. 8, Left valve (holotype) dorsal view
HU. 17.C.14.1. 9, Right valve lateral view HU. 17.C.15.1. 10. Left valve (holotype) lateral view
HU. 17.C.14.1. 11. Right valve dorsal view HU. 17.C.15.1. 12. Right valve internal view HU.
17.C.15.1. 14. Left valve (holotype) internal view HU. 17.C.14.1.

Fig. 13. Neocvthere ? {Centrocythere) denticulata Mertens, Upper Albian. Carapace from left HU.
17.C.18.1.

Figs. 15-22, 24. Neocythere {Physocythere) Hngenensis Mertens, Middle Albian. 15, Left valve female
dorsal view HU. 17.C.13.1. 16, Right valve male lateral view HU. 17.C.13.3. 17, Left valve female
lateral view HU. 17.C.13.1. 18, Right valve male dorsal view HU. 17.C.13.3. 19, Right valve female
internal view HU. 17.C.13.2. 20, Female earapaee ventral view HU. 17.C.13.4. 21, Female carapace
dorsal view HU. 17.C.13.4. 22, Right valve female lateral view HU. 17.C.13.2. 24, Left valve female
internal view HU. 17.C.13.1.

Figs. 23, 25. Neocythere {Neocythere) vanveeni Mertens, Middle Albian. 23, Right valve female lateral
view. HU. 17.C.10.1. 25, Left valve female lateral view HU. 17.C.10.2.

Palaeontology, Vol. 6

PLATE 41

KAYE, Neocythere

P. KAYE: THE OSTRACOD NEOCYTHERE

279

crosses the ventral surface. Duplicature narrow, dropping steeply into the inflated
interior of the valve. Inner margin and line of concrescence coincide.

The hinge of the right valve consists of two bar-like teeth divided into four or five
lobes. These teeth are separated by a long, straight locellate furrow. In the left valve
there are two elongated divided sockets separated by a long straight denticulate bar.
Above the bar, there is a broad flat shelf sloping to the dorsal margin but no accom-
modation groove.

Remarks. This species can be differentiated from other members of the group by the
shape of the dorsal margin and by the ornament.

Neocythere (Physocythere) hieroglyphica sp. nov.

Plate 41, figs. 8-12, 14

Holotype. A left valve from the Red Chalk, bed N4 at West Heslerton, no. HU 17.C.14.1.

Other material. Specimens no. HU 17.C.15.1-6, from the uppermost Red Chalk at South Cave.

Diagnosis. A species of the subgenus Physocythere having a straight to concave dorsal
margin, and an ornament of ridges arranged in a hieroglyphic pattern.

Description. Carapace strongly inflated, ventrally tumid. Greatest height at one-quarter
length, greatest width at mid-length. Dorsal margin straight or slightly concave with
rounded cardinal angles. Ventral margin convex, and obscured in its central portion by
the ventral tumidity. Anterior margin rounded, ventrally asymmetrical; posterior margin
triangular, forming an acute upturned point at mid-height. There is usually a flattened
rim along the dorsal margin, particularly in the left valves.

Lateral surfaces strongly ornamented with a pattern of concentric ribs enclosing an
irregular, hieroglyphic-like pattern of ridges. These concentric ribs radiate from the centre
of the dorsal margin, and parallel the other three margins, being most prominent
ventrally. Several sinuous ridges run from the centre of the dorsal margin, across the
lateral surface, branching and forming the hieroglyphic-like pattern with the concentric
ribs. The marginal areas are smooth. Inner margin and line of concrescence coincide.
Radial pore canals short and straight, numbering eight anteriorly and three posteriorly.
Normal pore canals large, but rather rare; situated along the crests of the concentric ribs.

Hinge very strongly built. In the right valve there are two large, elongated terminal
cusps, each divided into five lobes. They are separated by a long, straight locellate furrow
open ventrally, but bordered dorsally by a smooth bar. In the left valve there are two
large, strongly divided sockets separated by a long, straight denticulate bar. Above the
median element there is a broad, flat shelf sloping down to the dorsal margin.

Remarks. N. (P.) hieroglyphica appears to be restricted to the Red Chalk. It differs from
the other members of the group in the ornament, the greater inflation, and the strong
hinge.

Holotype left valve, HU 17.C.14.1
Right valve, HU 17.C.15.1.

Measurements

Length Height Total width

0-69 mm. 0-46 mm. 0-26 mm.

0-73 mm. 0-39 mm. 0-24 mm.

280

PALAEONTOLOGY, VOLUME 6

Neocythere (Physocythere) semilaeva sp. nov.

Plate 41, figs. 4, 7

Holotype. A left valve from the uppermost Red Chalk at South Cave; no. HU 17.C.16.1.

Other material. Five paratypes mounted as HU 17.C.17.1-5.

Measurements Length Height Total width

Left valve holotype, HU 17.C.1 6.1. 0-60 mm. 0-39 mm. 0-26 mm.

Description. Only left valves and one complete carapace of this species are known, and
therefore internal details of the right valves are lacking.

Carapace rounded, strongly tumid ventrally. Greatest height and width at mid-length.
Dorsal margin of the left valve strongly arched, forming a broad obtuse angle at about
the centre of the valve. Ventral margin convex, but obscured in lateral view by the
tumidity. Anterior margin broadly rounded, posterior margin subtriangular, forming a
rounded point at mid-height. The lateral surfaces are largely smooth, but show traces
of concentric ribbing. A large inflated rib follows the ventral margin and can be vaguely
traced around the edge of the lateral surface. This ridge is swollen ventro-laterally almost
forming alaeform expansions. The ventral surface is smooth.

The hinge of the left valve consists of two strong divided terminal sockets separated
by a long, straight denticulate bar. Above the median element there is a broad, flat shelf.

Occurrence. This species is only known from the Red Chalk (Upper Albian).

Remarks. N. {P.) semilaeva differs from the other species of the genus in the shape oi
the dorsal margin, the prominent ventral rib, and the smooth dorso-lateral surface.

Subgenus centrocythere Mertens 1956
Subgenotype. Centrocythere denticulata Mertens 1956.

Diagnosis. A subgenus of Neocythere s.l. showing the usual inflated shape and ventral
tumidity, but having a characteristic hinge arrangement. In the right valve the anterior
tooth is step-like, the back portion being twice as high as the front. The posterior tooth
is crenulate, and the median element consists of a locellate furrow, deepened at its
anterior end to form a smooth socket. There is an accommodation groove above the
median element in the left valve.

Neocythere ? {Centrocythere) denticulata Mertens 1956
Plate 41, fig. 13

? Cythere punctatula Jones 1849, p. 11, pi. 1, fig. 2a-n.

Centrocythere denticulata Mertens 1956, p. 204, pi. 11, figs. 66-71; pi. 14, figs. 97-99.
Brachycythere concentrica (Reuss); Deroo 1956, p. 1512, pi. 3, figs. 35, 36.

Material. Five carapaces, from the Red Chalk at Speeton.

Diagnosis. As only complete carapaces have been found, details of the hinge are unknown
and the material is therefore only tentatively referred to this subgenus. It is placed here
on a basis of the shape of the dorsal margin and posterior end, and also on the orna-
mentation.

Measurements Length Height Total width

Carapace, HU 17.C. 18.1. 0-60 mm. 0-39 mm. 0-35 mm.

P. KAYE; THE OSTRACOD NEOCYTHERE

281

CONCLUSIONS

The genus Neocythere has been subdivided into three subgenera on a basis of hinge
structure. The subgenera are all represented in the Speeton Clay, and are of similar shape
and ornamentation. Five of the six species are of Albian age. Of these, three are restricted
to the Upper Albian, but the other two are also found in the Middle Albian. Neocythere
{Neocythere) Protovanveeni sp. nov. occurs in the Lower Barremian and is the lowest
form of the genus in the Cretaceous yet described, though the author has in his posses-
sion specimens of this, or a closely related form, from the Hauterivian Tealby Clay of
Lincolnshire. It is possibly ancestral to the other forms, but, as the genus is absent or
very rare from the Middle and Upper Barremian, Aptian, and Lower Albian, this cannot
be proved with certainty. Of the six species, three are new, two belong to known German
species, and one is tentatively compared to a known species.

REFERENCES

ENNIS, w. c. 1937. The Upper Beds of the Speeton Clay. Trans. Hull Geol. Soc. 7, 130-8.

JONES, T. R. 1849. A monograph of the Entomostraca of the Cretaceous formation of England.
Palaeontogr. Soc. 1-40, 7 pi.

• and HiNDE, G. j. 1890. A supplementary monograph of the Cretaceous Entomostraca of England

and Ireland. Ibid. 1-70, pi. 1-4.

MERTENS, E. 1956. Zur Grenzziehung Alb. Cenoman in Nordwestdeutschland mit Hilfe von Ostra-
coden. Geol. Jahrb. 72, 173-230, 7 pi.

■ 1958. ' Cythere' lingenensis n.n. pro Cythere costata Mertens 1956. Ibid. 75, 359.

REUSS, A. E. 1846. Die Versteinenmgen der Bdhmisclien Kreideformation. Stuttgart, E. Schwelzerbart.
II. Bd., 1-148, pi. 14-51.

WRIGHT, c. w. 1955. In: swinnerton: A monograph of British Eower Cretaceous Belemnites,
Palaeontogr. Soc. 1-86, pi. 1-18.

P. KAYE

Department of Geology,
The University,

Manuscript received 3 September 1962 Reading

REVISION OF SOME LOWER CRETACEOUS
MICROSPORES FROM BELGIUM

by A. F. DELCOURT, M. E. DETTMANN, and N. F. HUGHES

Abstract. Most of the new Wealden spore species described in 1955 by Delcourt and Sprumont are refigured;
some have been placed in different genera. Two infra-specific taxa in the genus Concavissimisporites have been
raised to specific rank. The diagnoses of the genera Biretispohtes, Concavissimisporites, and Aequitriradites have
been emended.

Many of the spore species described by Delcourt and Sprumont (1955) in their pioneer
work on the Belgian Wealden have subsequently been widely reported from Lower
Cretaceous strata. Some of them are type species of important Mesozoic form-genera of
wide distribution, and a clearer understanding of them is obviously of great importance.
We have therefore re-examined the type specimens of those species instituted in 1955 and
in 1959 (French Wealden) which need clarification. As a result of our investigation
several species are transferred to different genera and some genera are emended; two
infra-specific taxa are raised to specific rank, as we consider that subdivisions of form
species of virtually unknown affinities are inappropriate. All the spores are now discussed
in the order in which the various groups are treated by Potonie (1956, 1958, 1960).

We have faced considerable nomenclatural difficulty over the current Russian policy
(Bolkhovitina 1961, Samoilovitch et al. 1961, and others) of placing large numbers of
fossil dispersed spore species in Recent (extant) genera. The tangle of names in micro-
fossil nomenclature is unfortunately unavoidable in a subject developing so quickly in
several different continents, but this particular (Russian) solution seems to be totally
undesirable. Knowledge of Mesozoic plants can only be obtained by studying all the
fossils patiently; no advantage accrues from implying on very slender evidence that such
genera as Lygodium Swartz and Anemia Swartz were present in the Lower Cretaceous.
Search is being made continuously for the ancestors of these genera, but the circumscrip-
tion of Lygodium Swartz cannot be blindly and indefinitely extended into the past on the
basis of dispersed spores alone, without making the Recent taxon meaningless. We there-
fore intend to continue using and improving the generally recognized fossil spore taxa.
We also notice that there has recently been conflict within the U.S.S.R. on this subject,
in the two contrasted papers in the same volume by Zauer et al. (1960) and Zaklinskaya
et al. (1960).

Some of the new photographs are oriented in the same way as the originals, but this is
not always possible because some of the originals are ‘mirror images’ due to reversal in
photographic printing. Some slight technical faults in a few of the new photographs are
noted in the plate legends. A few of the type slides have deteriorated badly and this is
noted in the worst cases. All the preparations remain in the collection of M. Delcourt
at Ath, Belgium.

Acknowledgements. We are grateful to Dr. W. Mullenders, of Louvain, for providing facilities for one of
us (M. E. D.) to study some of the type material mentioned, while Delcourt was away abroad.

Palaeontology, Vol. 6, Part 2, 1963, pp. 282-92, pis. 42-45.]

DELCOURT, DETTMANN, HUGHES: CRETACEOUS MICROSPORES

283

SYSTEMATIC SECTION

Anteturma sporites H. Potonie 1893
Turma triletes Reinsch 1881, emend. Potonie and Kremp 1954
Subturma azonotriletes Luber 1935
Infraturma laevigati Bennie and Kidston, emend. Potonie 1956
Genus cyathidites Couper 1953

Type species. Cyathidites australis Couper 1953.

Discussion. The slight extension of the genus by inclusion of further species makes it
a more purely morphographic unit, and less dependent on the affinities implied by Couper
(1953, 1958). Following the emendation of Delcourt and Sprumont (1955) Concavi-
sporites Pflug is retained for smooth spores having a concave amb and proximal ‘kyr-
tomes’. In uncompressed spores the kyrtomes constitute an abruptly convex exine
area surrounding the laesurae; in compressed spores it is reduced to arcuate folds in the
proximal inter-radial regions (Potonie and Kremp 1955, p. 13; Klaus 1960, p. 122).
Few Lower Cretaceous spores attributed to Concavisporhes exhibit kyrtomes around
the laesurae.

Comparison. Deltoidospora has straight sides (Potonie 1956). From the Palaeozoic, Leio-
iriletes (convexly triangular amb) was validated in 1954 (Potonie and Kremp).

Cyathidites punctatiis (Delcourt and Sprumont) comb. nov.

Plate 42, figs. 1-4

1955 Concavisporites punctatus Del. and Spr., p. 25, pi. 1, fig. 8; pi. 2, fig. 2.

1955 Concavisporites baldurnensis Del. and Spr., p. 24, pi. I, figs. 7 a, b.

1955 Cingulatisporites cavus Del. and Spr. p. 38, fig. 8.

1961 Lygodium asper (Bolk.) Bolkhovitina (pars), p. 86.

Description. The equatorially tapering ‘laesurate lips’ are narrow oblique exposures of
the slightly thickened exine adjoining the laesurae; undehisced (or uncompressed) spores
do not show this ‘feature’, which is elucidated by Van Campo (1961, figs. 3-8). As men-
tioned by Couper (1958), the wall ‘has a rather undulose appearance and is not punctate’;
the species (holotype — PI. 42, fig. 3) can therefore be included in Cyathidites, which is
described as psilate.

Discussion. The species baldurnensis included uncompressed (or inflated) specimens (PI.
42, figs. 1-2) of punctatus, the latter being the name selected by us for retention (Art. 57
of Code of Botanical Nomenclature, 1961 — Montreal). The holotype of the species cavus
is a specimen of punctatus with much adherent mineral matter (PI. 42, fig. 4). The
Canadian spore figured by Pocock (1962) as C. punctatus may be distinct in possessing
a sculpture of ‘distinct granules’.

Genus biretisporites Delcourt and Sprumont 1955, emend.

1955 Biretisporites Del. and Spr., p. 40.

1955 Punctatisporites Ibrahim, emend. Del. and Spr. (pars), p. 29.

1957 Hymenophyllumsporites Rouse, p. 363.

C 1213

U

284

PALAEONTOLOGY, VOLUME 6

Emended diagnosis. Microspores trilete with a triangular to subtriangular amb. Laesurae
enclosed within elevated lips which are upturned extensions of the proximal exine. Exine
smooth or almost smooth.

Type species. B. potoniaei Del. and Spr. 1955.

Comparison. The type species was originally considered to have equatorial thickenings
(Del. and Spr. 1955, Potonie 1956). The holotype, however, has no more than a smooth,
uniformly thick exine and a trilete aperture enclosed within elevated lips. Biretisporites
is therefore synonymous with Hymenophyllumsporites, which was instituted and clearly
defined by Rouse (1957). It is distinct from Dictyophyllidites Couper, the exine of which
is thickened about the laesurate margins (‘margo’ of Couper 1958). The type species of
Psilatriletes van der Hammen ex Potonie resembles Biretisporites, but its diagnosis does
not mention elevated laesurate lips.

Biretisporites potoniaei Delcourt and Sprumont 1955

Plate 42, figs. 12-14; Plate 44, fig. 11

1955 Biretisporites potoniaei Del. and Spr., p. 40, fig. 10.

1955 Pimctatisporites pileolus Del. and Spr., p. 30, pi. 2, fig. 5.

1955 Pimctatisporites nidosiis Del. and Spr., p. 31, fig. 4 (cf. Kremp, W. 1958, Catalogue of fossil

spores and pollen, 4, 17).

1956 Psilatriletes pileolus (Del. and Spr.) Potonie, p. 15.

Holotype. Delcourt and Sprumont 1955, p. 40, fig. 10; here refigured on Plate 42, figs. 12, 13. Distal
oblique aspect.

Description. Microspore trilete, biconvex; amb convexly subtriangular, diameter 4Sp.
Laesurae 23 p long; lips narrow (l-2p wide), elevated (2-3 p high) extensions of the
promimal exine. Exine smooth, 2-5p thick. The specimen has a median fold in the distal
exine, as does another specimen shown on Plate 44, fig. 1 1 .

Discussion. The further specimens of Psilatriletes (al. Pimctatisporites) pileolus (Del. and
Spr. 1959, pi. 5, fig. 15) are here refigured on Plate 42, fig. 14; they conform to the above
description of potoniaei as does the designated holotype of pileolus on which the apparent
granules are mineral grains.

Infraturma apiculati Bennie and Kidston, emend. Potonie 1956
Genus concavissimisporites Delcourt and Sprumont, emend.

1955 Concavissimisporites Del. and Spr., p. 25.

1961 Swartz; Bolkhovitina (pars), p. 100.

1961 Lygodium Swartz; Ivanova (pars) in Samoilovitch et al., p. 90.

1962 Concavisporites Pflug; Pocock (pars), p. 46.

Emended diagnosis. Microspores trilete ; amb triangular with concave to almost straight
sides. Exine of uniform thickness (inclusive of sculpture), verrucate; verrucae more or
less uniformly developed and evenly distributed over entire spore surface.

Type species. C. verrucosus Del. and Spr., emend.

Discussion. The genus was originally instituted to include trilete, azonate microspores

DELCOURT, DETTMANN, HUGHES: CRETACEOUS MICROSPORES 285

characterized by a ‘concavissime’ amb. Potonie (1956) suggested, but did not formally
propose, that the genus should be restricted to contain only such spores with verrucate
sculpture. The genus is here formally restricted to include verrucate spores which have
concave, but not necessarily strongly concave, ambs; it is distinct from Trilobosporites
Pant ex Potonie, which has a differentially thickened exine and/or larger sculptural
elements about the radial regions at the equator.

Concavissimisporites verrucosus Delcourt and Sprumont 1955, emend.

Plate 42, figs. 5-7

1955 C. verrucosus forma verrucosus Del. and Spr., p. 25, pi. 2, fig. \a.

1961 Lygodium verrucosus (Del. and Spr.) Bolkhovitina, p. 100, pi. 37, figs. 2a, b.
non 1955 C. verrucosus f. crassatus Del. and Spr., p. 26.
non 1962 Concavisporites verrucosus (Del. and Spr.) Pocock, p. 46.
non 1962 Concavisporites verrucosus var. minor Pocock, p. 47.

Holotype. Delcourt and Sprumont 1955, p. 25, pi. 2, fig. \a, here refigured on Plate 42, figs. 5-7. Proximal
aspect.

Description. Amb 90 p. in diameter (60 p in maximum radial length), triangular with
concave sides and broadly rounded angles. Laesurae 43 p long (2/3 spore radius), straight,
and with linearly arranged verrucae along margins. Exine 3 -5p thick (inclusive of sculp-
ture); verrucae closely spaced, l-2p high, semicircular in optical section, and with
circular to polygonal bases 2-4 p in diameter.

Comparison. This specimen is considered specifically distinct from the holotypes of the
two validly designated infra-specific taxa, f. crassatus and var. minor, which are (below)
elevated to specific rank.

Discussion. C. verrucosus is distinct from spores assigned to Concavisporites variverru-
catus by Couper (1958, p. 142, pi. 2, figs. 4-5); it is considerably larger, more strongly
concave, and has more regular verrucae.

Concavissimisporites crassatus (Del. and Spr.) sp. nov.

Plate 42, figs. 9-1 1

1955 C. verrucosus f. crassatus Del. and Spr., p. 26, pi. 2, fig. lb.
non 1962 Concavisporites verrucosus (Del. and Spr.) Pocock, p. 46.

Holotype. Delcourt and Sprumont 1955, pi. 2, fig. lb, here refigured on Plate 42, figs. 9-11. Proximal
aspect.

Description. Amb triangular with rounded angles and concave sides, l%p in diameter
(50 p in maximum radial length). Laesurae 43 /a long (4/5 spore radius), straight; margins
obliquely exposed and with linearly arranged verrucae. Exine 3 p thick (including sculp-
ture); with low, dome-shaped verrucae which are 1-3 p in basal diameter and spaced
1-3 p apart.

Comparison. The species differs from C. verrucosus in having smaller and more widely
spaced verrucae and longer laesurae.

286

PALAEONTOLOGY, VOLUME 6

Concavissimisporites minor (Pocock) sp. nov., comb. nov.

1962 Concavisporites verrucosus var. minor Pocock, p. 47, pi. 5, fig. 75.

Discussion. The imperfectly preserved holotype is distinct from C. verrucosus and C.
crassatus in having smaller verrucae which are more widely spaced.

Genus pilosisporites Delcourt and Sprumont 1955
Pilosisporites verus Delcourt and Sprumont 1955

Text-fig. 1

Comparison. Text-fig. 1 shows the broad-based form of the sculptural elements of the
holotype. The other difference from P. trichopapillosus is in the close spacing of sculptural
elements in the equatorial radial regions.

TEXT-FIG. 1, Detail of sculpture of Pilosisporites verus,
holotype (Del. and Spr. 1955, pi. 4, fig. 1). X 1000.

Infraturma murornati Potonie and Kremp 1954
Genus lycopodiumsporites Thiergart ex Delcourt and Sprumont 1955

Discussion. The genus was only validated in 1955 by selection of a ‘lectogenotype’, L.
agathoecus (Potonie). Nilsson (1958, p. 45) used this genus but Krutzsch (1959, pp. 159,
164, 165) suggested that the genus should be abandoned; although we sympathize with
his reasons we do not find his argument convincing. The two species mentioned below
could be placed in Klukisporites Couper 1958, if this genus should fall.

EXPLANATION OF PLATE 42

All figures X 500, unless otherwise stated.

Figs. 1-4. Cyathidites punctatus (Del. and Spr.) comb. nov. 1-2, Distal aspect, specimen of Concavi-
sporites baldurnensis D. and S. 1955, prep, e; 1, Low focus; 2, Mid focus. 3, Holotype, Cyathidites
punctatus, optical section ; aspect distal. 4, Proximal aspect of holotype of Cingulatisporites cavus D.
and S. 1955, p. 38; optical section, air bubble subcentral.

Figs. 5-7. Concavissimisporites verrucosus D. and S. emend., Holotype. 5, Mid focus. 6, Detail, X 1000,
mid focus. 7, X 1000, high focus.

Figs. 8-11. Concavissimisporites crassatus (D. and S.) sp. nov., holotype, proximal aspect (D. and S.
1955, pi. 2, fig. la). 8, High focus. 9, Mid focus. 10, Detail, X 1000, mid focus. 1 1 , X 1000, high focus.

Figs. 12-14. Biretisporites potoniaei D. and S. 1955. 12, Holotype, oblique distal aspect; mid focus. 13,
Low focus. 14, Spores illustrated as Psilatriletes pileolus (D. and S.) D. and S. 1959, pi. 5, fig. 15.

Note. Two extraneous hairs are present in figs. 6, 7, 10, 11.

Palaeontology, Vol. 6

PLATE 42

DELCOURT, DETTMANN and HUGHES, Lower Cretaceous Spores

DELCOURT, DETTMANN, HUGHES: CRETACEOUS MICROSPORES 287

Lycopodiumsporites triarcuatus Delcourt and Sprumont 1955
Plate 43, figs. 1, 2, 5

Description. The damaged holotype is refigured (PI. 43, fig. 1) and supported by two
other specimens from the same material (PL 43, figs. 2, 5). The proximal face is smooth,
unsculptured; other features as previously described. The two formae mentioned (1955)
are not validly published.

Lycopodiumsporites elongatus Delcourt and Sprumont 1955
Plate 43, figs. 3, 4

Description. The holotype is refigured. Again the proximal face is smooth, unsculptured.

Genus cicatricosisporites Potonie and Gelletich 1933

Type species. C. dorogensis Pot. and Gell. ; Eocene, Hungary.

Discussion. Kedves (1961 , p. 124) selected neotypes {sic) for C. dorogensis, and rediagnosed
the species to exclude, among other things, the many Mesozoic references (Couper 1958,
and others) ; he did not entirely succeed in this, but the purpose is clear and we agree
that none of the illustrated Lower Cretaceous specimens conforms with the species. We
therefore follow and amplify the policy of Delcourt and Sprumont (1955).

Cicatricosisporites hallei Delcourt and Sprumont 1955
Plate 43, figs. 6, 7

Description. The holotype is refigured. The muri are regular with sharply defined edges,
and are flat-topped; they are twice as wide as the intervening canals (lumina). The
spacing of four canals and muri in ll^i, is as previously illustrated (1955, text-fig. 1).

Comparison. C. hallei differs from C. dorogensis in possessing flat-topped muri with
sharp edges.

Cicatricosisporites sewardi Delcourt and Sprumont 1955
Plate 43, fig. 8

Description. The holotype is refigured as a photograph; diameter 55/x, not as previously
stated. The spacing (four canals and muri) is about lip only and thus differs little from
C. hallei’, the canals are, however, relatively wider, and the muri more round-topped
with less sharp edges.

Cicatricosisporites mohr hides Delcourt and Sprumont 1955

Discussion. It was not possible to improve on the original figure of this species because
the holotype is now orientated in a lateral aspect, and does not appear to be well preserved.

288

PALAEONTOLOGY, VOLUME 6

Turma zonales Bennie and Kidston, emend. Potonie 1956
Subturma auritotriletes Potonie and Kremp 1954
Infraturma auriculati Schopf, emend. Potonie and Kremp 1954
Genus trilobosporites Pant ex Potonie 1956

1956 Trilobosporites Pant ex Potonie, p. 55.

1961 Lygodium Swartz; Ivanova (pars) in Samoilovitch et al., p. 90.

Type species. T. hamionicus (Del. and Spr.) Potonie 1956.

Restated diagnosis. Microspores trilete; amb triangular. Exine differentially thickened in
equatorial radial regions where sculptured valvae are developed. Sculptural elevations
(grana, verrucae) larger and sometimes coalescent on valvae.

Discussion. Trilobosporites is distinct from Concavissimisporites on distribution of sculp-
tural elements and presence of valvae. Matonisporites Couper has a smooth exine and
Cibotiumidites Maljavkina is defined by Potonie (1960, p. 62) as cingulate.

Trilosbosporites hannonicus (Del. and Spr.) Potonie 1956

Plate 43, figs. 9, 10

Holotype. Delcourt and Sprumont 1955, p. 24, pi. 2, fig. 3. Refigured here on Plate 43, figs. 9, 10;
distal aspect.

Description. Amb concavely triangular with truncated angles, diameter 58 /x. Laesurae 25/i
long; margins weakly thickened and obliquely exposed. Exine 2-5/x thick, 4^ in equa-
torial radial regions where verrucate valvae are developed ; verrucae low (c. 1 pt high),
closely spaced, and with circular to polygonal bases 3-4 /x in diameter. Remainder of
exine with smaller, verrucae and granules, which are polygonal to circular in outline and
1-3 /X in basal diameter.

Remarks. Potonie (1956, p. 55) described the exine wrongly as microreticulate. It is now
shown to be verrucate.

Trilobosporites bernissartensis (Del. and Spr.) Potonie 1956

Plate 43, figs. 1 1-14

Holotype. Here figured as a photograph, Plate 43, figs. 11, 12; distal aspect.

EXPLANATION OF PLATE 43
All figures X 500 unless otherwise stated.

Figs. 1, 2, 5. Lycopodiumsporites triarcuatus D. and S. 1955. 1, Holotype, proximal aspect; mid focus.

2, Prep. B301, proximal aspect, high focus. 5, Prep. B103, distal aspect, high focus.

Figs. 3, 4. L. elongatus D. and S. 1955. Holotype, distal aspect. 3, Mid focus. 4, High focus.

Figs. 6, 7. Cicatricosisporites hallei D. and S. 1955. Holotype, oblique distal aspect. 6, X 1000, high
focus. 7, Mid focus.

Fig. 8. C. sewardi D. and S. 1955. Holotype, proximal aspect, high focus.

Figs. 9, 10. Trilobosporites hannonicus (D. and S. 1955). Holotype, distal aspect. 9, Low focus. 10,
X 1000, mid focus.

Figs. 1 1-14. Trilobosporites bernissartensis (D. and S. 1955). 11, Holotype, distal aspect, optical section.
12, High focus, proximal surface. 13, Single mount X3, Loc. Baudour Bed 1, proximal aspect, optical
section. 14, Single mount XI 4, Loc. Baudour Bed 1, proximal aspect, mid focus.

Palaeontology, VoL 6

PLATE 43

DELCOURT, DETTMANN and HUGHES, Lower Cretaceous spores

DELCOURT, DETTMANN, HUGHES: CRETACEOUS MICROSTORES 289

Description. The holotype is not well preserved and the isotype could not be located;
two other specimens from the same material are shown in Plate 43, figs. 13-14. As stated
by Couper (1958, p. 141), the proximal face is sculptured.

Genus appendicisporites Weyland and Krieger 1953
Appendicisporites ethmos Delcourt and Sprumont 1959
Plate 44, figs. 12, 13

Description. The holotype is here refigured. Distal aspect, diameter 60 /x, not as previously
stated. Sculpture as in Cicatricosisporites, with four canals and muri in about 12 p; muri
pitted throughout, pits up to 1/x diameter; the pitting is very constant.

Remarks. This species may have to be placed in Plicatella Maljavkina ex Potonie 1960;
this should be determined, however, by the origin of the ‘appendices’ (Dettmann 1963,
in press), rather than by their length (Potonie 1960, p. 50). The holotype of A. ethmos
does not illustrate this clearly.

Subturma zonotriletes Waltz 1935
Infraturma cingulati Potonie and Klaus 1954
Genus densoisporites Weyland and Krieger 1953
Densoisporites triradiatus Delcourt and Sprumont 1955

Plate 45, figs. 7-9

Holotype. Delcourt and Sprumont 1955, text-fig. 1 1 ; here refigured on Plate 45, fig. 7. Proximal aspect,
corroded; diameter 125ju., not as previously stated.

Description. Laesurae straight, extending to the periphery, and with elevated (3 p high)
lips. Tapering cingulum, 25 p wide. The second specimen (PI. 45, figs. 8, 9) is smaller
(96 /x) and shows the surface detail more clearly.

Genus baldurnisporites Delcourt and Sprumont 1955

Discussion. Potonie (1956, p. 60) regarded the genus as insufficiently defined, but placed it
in the Cingulati; he removed the megaspore species Minerisporites marginatus (Dijkstra)
from Baldurnisporites, with which we agree. The specimens of the two remaining species
are refigured below, but further specimens seem to be rare in the preparations ; there is
a slight suspicion that these spores may be perinate specimens of Cyathidites (al. Con-
cavisporites) punctatus (Del. and Spr.) comb. nov.

Type species. B. cennius.

Baldurnisporites cernuus Delcourt and Sprumont 1955
Plate 44, figs. 7, 8

Type specimens. The holotype (Del. and Spr. 1955; pi. 3, fig. 5) has been accidentally
destroyed. The isotype (Del. and Spr. 1955, p. 44) is now figured on Plate 44, figs. 7, 8.

Baldurnisporites papillosus Delcourt and Sprumont 1955
Plate 44, figs. 9, 10

Description. The holotype, distal aspect, diameter 100/x, is refigured. The laesurae do not

290

PALAEONTOLOGY, VOLUME 6

appear to reach the flange as originally described; the ‘baton’ supporting the flange is
clearly seen near one radius.

Infraturma zonati Potonie and Kremp 1954
Genus aequitriradites Delcourt and Sprumont, emend. Cookson and Dettmann 1961

1955 Aequitriradites Delcourt and Sprumont, p. 44.

1961 Selaginellidites Krasnova in Samoilovitch et al., p. 38.

1961 Aequitriradites D. and S. emend. Cookson and Dettmann, p. 426.

Restated diagnosis. Microspores tetrahedral, with a membraneous zona. Exine inaper-
turate or hilate; with structural and/or sculptural modifications at and about the distal
pole, where a ‘hilum’ may be formed as the result of a natural exinous breakdown.
Proximal tetrad mark distinct, or only faintly represented, especially towards the pole.

Type species. A. dubius.

Aequitriradites dubius Delcourt and Sprumont 1955

Plate 45, figs. 1-4

1955 A. inconspicuus Delcourt and Sprumont, p. 45, pi. 2, fig. 6.

Holotype. Delcourt and Sprumont 1955, pi. 3, fig. la only (not fig. lb — see below); here
refigured on Plate 45, figs. 1-3.

Description. Proximal aspect, diameter 85ju; exine l-Sp- thick, coarsely granulate except
in a circular area (25 p in diameter) about the distal pole, where it is composed of discrete,
low, polygonal-based (2-3 ju diameter) irregular verrucae. Zona 20 p wide, scabrate, with
four short (5-6 p long), radially oriented ridges which emanate outwards from its inter-
radial inner margin. Proximal tetrad mark comprises two, long (30 p), sinuous ridges
(2p wide) which extend from pole to zone.

EXPLANATION OF PLATE 44

All figures x 500 unless otherwise stated.

Figs. 1-6. Rouseisporites grauospeciosus (D. and S.) comb. nov. 1, Holotype, distal aspect, high focus.
2, Mid focus. 3, Single mount X7, Baudour Bed 1, distal aspect, mid focus. 4, High focus. 5, Single
mount X21, Baudour Bed 1, distal aspect, high focus. 6, Mid focus.

Figs. 7, 8. Baldurnisporites cernuus D. and S. 1955, Isotype, proximal aspect. 7, High focus. 8, Mid focus.

Figs. 9, 10. B. papillosus D. and S. 1955. 9, Holotype, distal aspect, mid focus. 10, Detail, X 1000.

Fig. 11. Biretisporites potoniaei D. and S. 1955; prep. 9, proximal aspect.

Figs. 12, 13. Appendicisporites etiimos D. and S. 1959. 12, Holotype, distal aspect, high focus. 13,
Mid focus.

EXPLANATION OF PLATE 45

All figures X 500.

Figs. 1-4. Aequitriradites dubius D. and S. 1955. 1-3, Holotype, proximal aspect. 1, Low focus. 2, Mid
focus. 3, High focus. 4, Distal aspect, holotype of A. inconspicuus D. and S. 1955, pi. 2, fig. 6; high
focus.

Figs. 5, 6. Schizaeoisporites phaseolus D. and S. 1955; holotype, lateral aspect. 5, High focus. 6, Low
focus.

Figs. 7-9. Densoisporites triradiatus D. and S. 1955. 7, Holotype, proximal aspect, mid focus. 8, Prep. 9,
proximal aspect, high focus. 9, Mid focus.

Palaeontology, Vol. 6

PLATE 44

DELCOURT, DETTMANN and HUGHES, Lower Cretaceous spores

Palaeontology, Vol. 6

PLATE 45

DELCOURT, DETTMANN and EFUGHES, Lower Cretaceous spores

DELCOURT, DETTMANN, HUGHES: CRETACEOUS MICROSPORES 291

Discussion. Delcourt and Sprumont’s holotype of A. inconspicuus (pi. 2, fig. 6), which is
here refigured on Plate 45, fig. 4, has a coarsely granulate exine and is nonspecific with
A. dubius. Delcourt and Sprumont’s specimen in pi. 3, fig. lb has a rugulo-reticulate
sculpture, and belongs to a separate undescribed species.

Genus rouseisporites Pocock 1962
Type species. R. reticiilatiis Pocock 1962.

Discussion. This genus appears to be distinct from Zlivisporis Pacltova 1961, and Seducti-
sporites Chlonova 1961, on the presence of zonal invaginations in each radial, equatorial
region.

Rouseisporites granospeciosus (Del. and Spr.) comb. nov.

1955 Cingulatisporites granospeciosus Del. and Spr., p. 39, pi. 4, fig. 2.

Holotype. Delcourt and Sprumont 1955, p. 39, pi. 4, fig. 2. Here refigured on Plate 44, figs. 1-2. Distal
aspect. Diameter 65p.

Deseription. Microspore zonate, biconvex; amb subcircular. Sclerine two-layered con-
sisting of a granulate inner layer {2p thick) and an enveloping membraneous, zonate
outer layer. Zona smooth (8p, wide) with a flask-shaped invagination (Sp diam.) in each
region at the equator. Proximal surface smooth except for a faint tetrad mark. Distal
surface with radially orientated, bifurcating, muroid ridges (2-3 ju, high).

Diseussion. Two other specimens are figured on Plate 44, figs. 3-6. The species has been
removed from Cingulatisporites on the basis of its clearly special characters of the zona.
C. euskirehensoides Delcourt and Sprumont 1955, p. 38, might have required similar
treatment, but we were unable to illustrate the holotype further.

Turma monoletes Ibrahim 1933
Subturma azonomonoletes Tuber 1935
Infraturma sculptatomonoleti Dybova and Jachowicz 1957
Genus schizaeoisporites Potonie 1951
Sehizaeoisporites pbaseolus Delcourt and Sprumont 1955

Plate 45, figs. 5, 6

Deseription. The holotype is refigured as a photograph; lateral view, length 68 p.. Speci-
men dense, as originally implied.

REFERENCES

BOLKHOviTiNA, N. A. 1961. Fossil and Recent spores of the family Schizaeaceae. Trud. Geo/. Inst. Akad.
Nauk SSSR, 40, 1-176, pis. 41. [In Russian.]

CHLONOVA, A. F. 1961. Spores and pollen of the upper part of the Upper Cretaceous from the Eastern
part of the Western Siberian lowland. Trud. Inst. Geol. Geophys. Akad. Nauk SSSR Novosibirsk, 7,
1-138, 17 pi. [In Russian.]

COOKSON, I. c. and dettmann, m. e. 1961. Reappraisal of the Mesozoic microspore genus Aequitrira-
dites. Palaeontology, 4, 425-7.

couPER, R. A. 1953. Upper Mesozoic and Cainozoic spores and pollen grains from New Zealand.
N.Z. Geol. Surv. Pal. Bull. 22, 77 pp.

— — 1958. British Mesozoic microspores and pollen grains. Palaeontographica, 103B, 75-179, pi. 15-31.

292

PALAEONTOLOGY, VOLUME 6

DELCOURT, A. F. and SPRUMONT, G. 1955. Les spores et grains de pollen du Wealdien du Hainaut. Mem.
Soc. Belg. GeoL, n.s., 4, 1-73, 4 pi.

1959. Spores, grains de pollen, Hystrichospheres et Peridiniens dans le Wealdien de Feron-

Glageon. Ami. Soc. Geol. Nord, 79, 29-64, pi. 3-7.

DETTMANN, M. E. 1963. (In press.)

KEDVES, M. 1961. Etudes palynologiques dansle bassinde Dorog — II. Pollen et Spores, 3, 1, 101-53, 9 pi.
KLAUS, w. 1960. Sporen der Karnischen Stufe der ostalpinen Trias. Jb. geol. Bimdesanst. Oesterr.,
Sonderb. 5, 107-82, 11 pi.

KREMP, G., AMES, H. T. and KOVAR, A. J. 1958. Catalog of fossil spores and pollen, 4.

KRUTZSCH, w. 1959. Mikropalaontologische untersuchunge in der Braunkohle des Geiseltales. Geologie,
Beiheft 21-22, 1-425, 49 pi.

PACLTOVA, B. 1961. On some plant microfossils from the Upper Cretaceous in the South-Bohemian
basin I. Shorn. Ustred. Ust. Geol., Odd. palaeont., Praha, 26, 47-102, 24 pi.

PococK, s. A. j. 1962. Microfloral analysis and age determination of strata at the Jurassic-Cretaceous
boundary in the Western Canada plains. Palaeontographica, lllB, 1-95.

POTONiE, R. 1956-60. Synopsis der Gattungen der Sporae Dispersae, l-III. Belli. Geol. Jb. 23, 31, 39.

— — and KREMP, G. 1955. Die sporae Dispersae des Ruhrkarbons^ — -I. Palaeontographica, 98B, 1-136.
SAMOiLOViTCH, s. R. et al. 1961. Pollen and spores of Western Siberia; Jurassic to Palaeocene. Trad.

VNIGRI, Leningrad, 177, pp. 657, 84+65 pi. [In Russian.]

VAN CAMPO, M. 1961. Mecanique aperturale. Grana Palyn. 2, 3, 93-97.

ZAKLiNSKAYA, E. D. et al. 1960. Taxonomy and nomenclature of fossil spores and pollen. In Pre-
Quaternary Micropalaeontology. Report Soviet Geol., XXI Int. Geol. Congr., Problem 6, 167-71,
[In Russian; English summary.]

ZAUER, V. V. et al. 1960. Taxonomy, Nomenclature and Description Order of fossil spores and pollen.
Ibid. 159-66.

A. F. DELCOURT

College St. Julien,

Ath, Belgium

M. E. DETTMANN
N. F. HUGHES

Department of Geology,
Sedgwick Museum,

Manuscript received 21 August 1962 Cambridge

PROBLEMATICAL MICROFOSSILS FROM THE
CRETACEOUS AND PALAEOCENE OF THE

MIDDLE EAST

by GRAHAM F. ELLIOTT

Abstract. During studies of the non-foraminiferal microfossils, especially the algae, of the Middle East over
the last decade a considerable number of problematical organisms have been noted. These include some whose
allocation is doubtful, at varying taxonomic levels, or whose position has only recently been established, as well
as new or unusual examples of known families and genera. A selection of these, Cretaceous and Palaeocene in
age, is now placed on record as a further contribution to Tethyan micropalaeontology.

1. Calcareous Algae

Pseudolithothamnium was described by Pfender (1937) as an organism apparently similar
in growth-form and appearance in reflected light to the melobesioid algae (of which
Lithothamnium and Lithophyllum are common examples), but differing in the transparent
appearance of the tissues in thin section, in which condition the melobesioids appear
more or less uniformly dark. This distinction was ascribed to differences in the calcifi-
cation during life. Pfender saw no close resemblance in the cell-structures of Pseudolitho-
thanmhim to those of the melobesioids, and although she described apparent sporangial
structures she did not regard the fossil as close to this algal family, and only doubtfully
referred it to the Rhodophyta (red algae). She recorded it from rocks ranging from
Lower Cretaceous to Oligocene in age, over a very wide area of central and southern
Europe, the Mediterranean Basin, the Middle East, and Madagascar.

Pseudolithothamnium was figured from the Upper Cretaceous of the Carpathians by
Andrusov (1939), and recorded from both Upper Cretaceous and Palaeocene of northern
Iraq by Elliott (1955, 1960). These records, following Pfender, were all referred to the
single described species P. album.

More recently a convincing case has been put forward by Massieux and Denizot
(1962) for the identity of the fossil Pseudolithothamnium with the Recent Ethelia. This is
an encrusting lamellar red alga (family Squamariaceae), with a central mesothallus
margined by upper and lower perithalli; the reproductive structures are temporary
deciduous outgrowths from the upper surface. The internal structures are not normally
easy to prepare in orientated section owing to the contorted and twisted nature of the
thallus. Fossil records of Pseudolithothamnium are all based on random cuts in thin-
section or fractured surfaces, and it is not surprising that the detailed structure of Ethelia
has not been seen in them before. The somewhat doubtful alleged reproductive structures
in the fossils are discounted and ascribed to mineralization and fracture.

Ethelia is a calcified warm-water genus; the calcium carbonate is in the form of
aragonite, unlike the melobesioids, which are calcitic. This initial difference is considered
the probable cause of the different appearance of the fossils in thin-section. It may be
added that similar exceptions are known in largely aragonitic groups, e.g. Pseudovermi-
porella (calcitic) in the Dasycladaceae (largely aragonitic).

[Palaeontology, Vol. 6, Part 2, 1963, pp. 293-300, pi. 46-47.]

294

PALAEONTOLOGY, VOLUME 6

Massieux and Denizot show similar structure in Recent Atlantic Ethelia fosliei and
French Eocene PseudolitJwihamnium album Pfender, but wisely do not attempt to equate
the long-range fossil Pseudolithothamnium album with any of the modern botanical
species of Ethelia, and therefore propose the combination Ethelia alba (Pfender 1937).

The Iraqi material now figured from the Palaeocene of Kurdistan, a richly algal level,
comes from the Kolosh Formation at Rowanduz and the Sinjar Limestone at Sarmord.
The Rowanduz slide (PI. 47, fig. 1) shows a tangle of thalles of Ethelia intergrown
with those of the melobesioid Archaeolithothanmium (and also with other algae, not
shown). Ethelia is transparent, Archaeolithothanmium dark, except for the chains of
sporangial cavities. Although the section shows a considerable area of Ethelia, only at
one spot (PI. 47, fig. 2) was it possible to discern a vertical cut of the thin thallus
showing the central mesothallus and upper and lower perithalli. The thickness of this
thallus-section (0-26 mm.) is comparable with that of the Recent Ethelia (0-23 mm.)
given by Massieux and Denizot (1962, fig. 2). Much of the section shows as highly
oblique cuts through successive laminae about 0-10-OT2 mm. thick; this is even better
shown on the Sarmord specimen (PI. 47, fig. 3), where the varying laminae range up to
0-24 mm. in thickness as seen, traversed by a curved-radial cell arrangement, and the
section is presumably very highly oblique through a gently curved flat growth. This may
not be the same species, but Pfender (1937, fig. 2) shows a somewhat similar section of
Turkish Palaeocene material, with similar transparent tissues, and she refers to the
curved-radial ‘jet d’eau’ cell-arrangement. In the Recent Ethelia the species biradiata and
fosliei show considerable differences in section (Weber-Van Bosse 1921).

Ethelia alba is easily distinguished from the associated solenoporoid Parachaetetes
asvapatii, a much more massive species in which the curved-radial structure is much
coarser, and which is normally yellowish or buff in thin-section appearance. It is also
different from those melobesioids in which diagenesis has altered the dark calcite to
a translucent appearance with partial obliteration of cell-structure, a condition seen not
uncommonly in various Tertiary limestones.

2. Bacinella

Bacinella was diagnosed by Radoicic (1959), from the Lower Cretaceous of Jugoslavia,
as nodular and encrusting algae whose interior is constructed of irregular cells of different
size and form, filled with crystalline calcite, whereas the intercellular mass is crypto-
crystalline; the dermal structure is differentiated. In her description she states that, when
exceptionally preserved, B. irregularis may show a subdermal structure like that of
Lithocodium aggregatum Elliott.

Elliott (1956) described L. aggregatum, also Lower Cretaceous in age, as irregular
nodular growths largely formed of thin superimposed thalles : within the outer zone of
definite Lithocodium structure there occurs in some, but not all, cases a very coarse
irregular inner mesh or spongy mass, described originally as ‘a larger mass of irregular
tubules, never radial and presumably of Lithocodium itself’. This inner structure is the
same as in Bacinella.

There is thus often a close association between these two doubtfully algal growths;
in spite of frequent recrystallization, they have been seen to occur both separately and
together in favourable thin-sections of Middle Eastern material. Moreover, Lithocodium
is itself often intimately intergrown with the lamellar stromatoporoid Burgundia.

G. F. ELLIOTT: PROBLEMATICAL MICROFOSSILS

295

It seems likely that Lithocodiiim and BacineUa may be constructed by the same organ-
ism, but, pending a full investigation on favourable material, it seems best to use both
names, for the appropriate structures. Both are typically Lower Cretaceous, but Radoicic
(1959) states that she has seen structures like BacineUa in both the Upper Jurassic and
Upper Cretaceous. Upper Jurassic species of Lithocodiiim have been described by Endo
(1961) from Japan. The example now figured (PL 48, fig. 3) is from the Qamchuqa
Limestone, Albian level, of Zewiya, Pir-i-Mugrun, Sulaimania, north-east Iraq.

3. Microcalamoides

As described by Bonet (1956) from the Lower Cretaceous (Barremian to Lower Albian)
of Mexico, this is a problematic microfossil in the form of a thin-walled cylinder, fluted
or grooved with about thirty longitudinal sulci, and ranging from about 0-2 to nearly
TO mm. in diameter; the fossils are all fragmentary or incomplete. Bonet discusses the
pelagic associations, filamental and perhaps ramifying structure, and possible algal
nature of this organism, which he describes as M. diversus, with varieties A, B, and C.

It is now recorded from the Lower Cretaceous Yamama Formation (Valanginian-
Hauterivian) of Iraq (PI. 47, fig. 4). The example shown is larger (diameter 1-69 mm.)
and approximates most closely to Bonet’s type C ; fragments of type B have been seen
also. It is associated with debris of Permocalcidus, dasyclads, and BacineUa', all are
shallow-water algae, but fragmented and washed out to a deeper zone, and it could be
a pelagic addition to this debris from the littoral zone.

As well as the Mexican and Iraqi records, Microcalamoides has been seen by the writer
from the Lower Cretaceous of Borneo, and so it has a very wide distribution.

4. Calymenospongia

Studies of calcareous algae from the Middle East (bibliography in Elliott 1960; 1961)
were largely carried out on thin-sections, owing to the nature of the material collected,
though weathered or loose specimens were examined whenever available. A sample from
the Palaeocene of Bekhme, Erbil Liwa, northern Iraq, consisted of a loose detrital
serpentine sand rich in dissociated calcareous algae. Some of these, such as Cymopolia
and Halimeda, were instantly recognizable; some, e.g. Pagodaporella (Elliott 1956),
proved on dissection to be already well known in thin-section, and the solid fossil per-
mitted description of the genus. The earlier reconstruction from sections of the external
appearance of Dissoeladella by Pia (Rama Rao and Pia 1936) was proved correct from
specimens in this sample. Among these little algae was a fossil which, similar in size-
range and general appearance to the others, was revealed on sectioning to be, not an
alga, but a small calcisponge.

Genus calymenospongia gen. nov.

Small calcisponge of uniserial consecutive near-spherical hollow growths, fused at
junctions, and communicating internally by large, single apertures; a few scattered,
irregularly placed pores penetrating outer surface of growths. Palaeocene, Iraq.

Type species. C. kiirdistanensis sp. nov.

296

PALAEONTOLOGY, VOLUME 6

Calymenospongia kurdistanensis sp. nov.

Plate 46, figs. 1, 2, 3

Description. This sponge is about 2-3 mm. long (incomplete), with a maximum diameter
of 0-72 mm. It consists of a simple, straight or gently curved, single series of consecutive
swelhngs delimited by constrictions, like a row of fused spheres. These are circular in
transverse section : in the measured example quoted above there are six (one incomplete),
the others having a height of 0-42 mm. A few scattered coarse pores are to be seen on
each swollen surface. Internally, a longitudinal section reveals that each swelling or unit
is hollow and thin-walled, with a wall-thickness of 0-03 mm. measured at the maximum
diameter of the unit. Top and bottom, inside from the external constrictions, the walls
continue inwards to fuse with those of the next unit in a swollen thickening; this is
perforated in the centre by a large intercommunicatory pore. The vertical thickness of
the swollen fused internal walls, and the horizontal diameter of the intercommunicatory
pores which they encircle, are about the same, 0T5 mm. The scattered pores of the
external surfaces are filled with calcite, but are believed to have originally perforated the
wall: the diameter of one seen in seetion is 0-026 mm. The wall-structure throughout
shows as mottled grey calcite interpreted as fused spicules; thin-sections of material
from the famous Lower Cretaceous calcisponge locality of Faringdon, England, show
a similar appearance.

Holotype. BM S8335 (PI. 46, fig. 1 ), Kolosh Formation (Palaeocene), Bekhme, Erbil Liwa, northern Iraq.
Paratypes. BM S8336, S8337 (PI. 46, figs. 2, 3), same locality and horizon.

Remarks. This sponge is closest to Sollasia (Steinmann 1882) from the Carboniferous of
Spain, resembling it in size and general simplicity of structure, but differing in having
a wall showing one layer only, unlike the two of Sollasia, and in showing a more complete
fusion internally at the junctions of the walls of the consecutive units. This latter feature
is probably only a consequence of the different wall-structure. The group of sponges to
which Sollasia belongs has recently been carefully revised in great detail by Seilacher
(1962), on the occasion of the description of new and exceptionally well-preserved
material. He places Sollasia in the family Celyphiidae de Laubenfels, superfamily Aporata
Seilacher, suborder Sphinctozoa Steinmann, order Pharetronida Zittel. Like Sollasia,
Calymenospongia is catenulate and cryptosiphonate. Seilacher interprets the wall-structure

EXPLANATION OF PLATE 46

Fig. 1. Calymenospongia kurdistanensis gen. et sp. no\. Holotype, BM S8335, x50. Kolosh Forma-
tion, Palaeocene; Bekhme, Erbil Liwa, northern Iraq.

Figs. 2, 3. C. kurdistanensis, paratypes, BM S8336, S8337, vertical and transverse thin-sections, X 50;
same locality and horizon.

Fig. 4. Coptocampylodon lineolatus gen. et sp. nov. Holotype, BM Z988, X 56. Sarmord Formation,
Hauterivian, Lower Cretaceous; Surdash, Sulaimania Liwa, north-east Iraq.

Figs. 5, 6. C. lineolatus, lateral and terminal views of two examples, same locality and horizon, x 56.

Fig. 7. Favreina prusensis Parejas. BM Z996, thin-section, X 56. Chia Gara Formation, bottom
Cretaceous (Berriasian); Ser Amadia, Mosul Liwa, northern Iraq.

Fig. 8. Coptocampylodon lineolatus. Paratype, BM Z993, thin-section, X 56; Garagu Formation, sub-
surface Lower Cretaceous, Fallujah Well, Dulaim Liwa, Iraq.

Palaeontology, Vol. 6

PLATE 46

ELLIOTT, Problematical microfossils

G. F. ELLIOTT: PROBLEMATICAL MICROFOSSILS

297

of the Palaeozoic genus as spicules embedded in an original calcareous sclerosome,
giving a rigid wall in life and hence occasioning intermittent segment growth. The hollow
segments seen in this fossil are considered to have contained soft sponge tissue originally.
These interpretations are applicable to Calymenospongia also.

Calymenospongia is the first member of the Sphinctozoa to be described from post-
Cretaceous strata. It is easily distinguished from the somewhat similar Sphaerocoelia
(Upper Jurassic-Cretaceous), which has a markedly perforate wall.

Simple sponges of this kind, of microfossil dimensions, may be of much more common
occurrence than has been realized hitherto,

5. Coptocampylodon

Grooved spicules of Lower Cretaceous age have been familiar to micropalaeontologists
handling Middle Eastern material for many years, and have usually been referred to as
Aciciilaria sp. Acicularia s.s. (Tertiary and Recent) is a genus of dasyclad algae with a
terminal calcified sporangial disc which eventually falls apart into radial petal-hke
calcareous spicules each containing numerous, tiny, spherical, sporangial cavities. More
or less similar algal structures of varying form occur from the Jurassic onwards, usually
as small dissociated bodies, and have been described as species of Acicularia s.l. At least
three are known from the Lower Cretaceous: a small, spherical, sporangial body "Ter-
quemelld’ sp., the ovoid A. antiqua Pia, and the long, irregular^, elongata Carozzi. Trans-
verse cuts of the Middle East spicule referred to above are not unlike similar sections of
worn examples of the acicularians, but numerous longitudinal and obhque cuts reveal
that it is different. It is now described and its possible biological nature discussed.

Genus coptocampylodon gen. nov.

Small solid cylindrical calcareous bodies, longer axis gently curved or irregular, cir-
cular in cross-section but deeply incised by longitudinal grooves, ends irregularly rounded.
Lower Cretaceous; Middle East and Borneo.

Type species. C. lineolatiis sp. nov.

Coptocampylodon lineolatus sp. nov.

Plate 46, figs. 4, 5, 6, 8; Plate 48, fig. 2

Description. Solid short cylindrical calcareous bodies (units), up to 3-0 mm. in length
(incomplete) and varying from about 0-25 to 1-0 mm. in diameter. The longer axis of
the unit is gently curved or slightly irregular: the ends, when not obviously broken, are
irregularly rounded. The outer surface, itself usually smooth, but in some examples
showing a feeble transverse lamination, is deeply incised by parallel equidistant longi-
tudinal grooves, from five to eight in number. In transverse section these grooves are
seen to widen inwards and often to be near-circular in cross-section, with an internal
diameter of 0-04-0-05 mm. The circular transverse section of the whole unit is notched
by the grooves and has the appearance of a coarse sparsely spoked cog-wheel or of a
stellate structure with truncated rays. Longitudinal sections show the parallel, matrix-
filled grooves, either curved or passing out of the plane of the section owing to their
irregularity.

298

PALAEONTOLOGY, VOLUME 6

Holotype. BM Z988 (PI. 46, fig. 4); Sarmord Formation (Hauterivian level). Lower Cretaceous;
Surdash, Sulaimania Liwa, north-east Iraq. Paratypes. BM Z993) (PI. 46, tig. 8), subsurface Garagu
Formation, Lower Cretaceous; Fallujah Well, Dulaim Liwa, Iraq.

Other material. Numerous examples, solid and thin-section, from the Lower Cretaceous of Iraq. Seen
in thin-section from the Lower Cretaceous of Borneo.

Remarks. As stated above, transverse sections of Coptocampylodon have previously been
interpreted as acicularian, but the longitudinal sections disprove this, since all known
acicularian spicules, long or short, have near-spherical sporangial cavities giving circular
cross-sections at all angles of cut.

Nothing similar to the units of Coptocampylodon is known amongst the algae or the
spicular elements of calcareous sponges. The mineral structure is not that of echinoderm
skeletal elements, and the form is quite different from that of ‘alcyonarian spicules’.
These latter are the minute embedded calcareous elements of the horny corals or sub-
class Octocorallia. In the octocoral family Isididae Lamouroux there occur genera
showing an alternation of horny nodes and calcareous internodes ; the latter are known
fossil, notably in the genus Moltkia (Upper Cretaceous). Typically these isolated cal-
careous bodies are larger than those of Coptocampylodon, and show conspicuous calicular
pits ; they are also associated with the larger, dichotomously branching structures charac-
teristic of the genus. However, comparison of the varied series of skeletal remains of
Moltkia figured by Voigt (1958) shows that some of the smaller dissociated elements are
very similar to the Iraqi fossil (cf. Voigt 1958, pi. 3, figs. 9, 10; Moltkia isis Steenstrup).
A comparison may also be made with the elements of the decorticated axis of Isis (Recent).
It is therefore suggested that Coptocampylodon comprises the skeletal remains of a small
octocoral in which horny and calcareous joints alternated. Coptocampylodon is earlier
(Lower Cretaceous) than these genera; most pre-Upper Cretaceous records of fossils
referred to the octocorals seem doubtful (Bayer 1956; Voigt 1958; Hantzschel 1958),
but this allocation seems the most likely.

EXPLANATION OF PLATE 47

Fig. 1 . Transparent Ethelia (Pseudolithothamnium), contrasted with dark ArchaeoUthothammiim showing
chains of conceptacles. BM V44845, thin-section, x40. Kolosh Formation, Palaeocene; Rowanduz,
Erbil Liwa, northern Iraq.

Fig. 2. Ethelia alba (Piendev). BM V44845, thin-section, xlOO. Vertical cut of single growth showing
central mesothallus with upper and lower perithalli. Kolosh Formation, Palaeocene; Rowanduz,
Erbil Liwa, northern Iraq.

Fig. 3. Ethelia albatPiendcr). BM V44844, thin-section, X 30. Sinjar Limestone, Palaeocene; Sarmord,
Sulaimania Liwa, north-east Iraq.

Fig. 4. Microcalamoides diversusPone\.,ci. iyy>cC. BM Z992, thin-section, x28. Subsurface Yamama
Formation, Valanginian-Hauterivian; Makhul no. 1 well, Mosul Liwa, northern Iraq.

EXPLANATION OF PLATE 48

Fig. 1. Favreina montana sp. nov. Syntypes, BM Z997, thin-section, X56. Qamchuqa Formation,
Cretaceous ( Albian) ; Gund-i-Shikavt, Erbil Liwa, northern Iraq.

Fig. 2. Coptocampylodon lineolatiis gen. et sp. nov. BM Z994, random cuts in thin-section, X 56.
Yamama Formation, subsurface Lower Cretaceous (Valanginian-Hauterivian); Makhul no. 1 well,
Mosul Liwa, northern Iraq.

Fig. 3. Bacinella irregularis R-didoicic. BM Z995, thin-section, x28. Qamchuqa Formation, Creta-
ceous (Albian) ; Zewiya, Pir-i-Mugrun, Sulaimania Liwa, north-east Iraq.

Palaeontology, Vol. 6

PLATE 47

ELLIOTT, Problematical microfossils

Palaeontology, Vol. 6

PLATE 48

I

2

ELLIOTT, Problematical microfossils

"V, I .

*

e

>

C±M

G. F. ELLIOTT: PROBLEMATICAL MICROFOSSILS

299

6. Favreina

The peculiar microfossils named Favreina by Bronnimann (1955) are short, solid,
cylindrical pellets perforated by varying numbers of fine longitudinal canals in arrange-
ments showing as characteristic patterns in cross-section. They were first figured from
the Swiss Jurassic (Joukowsky and Favre 1913), but we owe to Parejas (1935, 1948) the
recognition of these as the faecal pellets of certain crustaceans, the Anomura, in which
the fleshy structure of the stomach initiates perforations in the gut-content of largely
inorganic ingested material. Since different genera and species of living Anomura show
distinctively different pellet-patterns (Moore 1932), numerous species of Favreina have
been described on the morphology of the fossils; two recent review studies are those of
Bronnimann and Norton (1961) and Elliott (1962). Two new records can now be added
to the Middle East species listed and described in the latter study.

Favreina prusensis Parejas
Plate 46, fig. 7

Remarks. This distinctive species (Parejas 1948) from the Upper Jurassic (Portlandian)
of Turkey, shows from 66 to 136 canals, set in a complicated pattern of zigzag lines as
seen in transverse section. No doubt this pattern increased in complexity with the growth
of the individual originating the pellets. Listed for comparison by Elliott (1962) on the
strength of Parejas’s described material, it is now recorded from the Upper Chia Gara
Formation, Berriasian, of Ser Amadia, Mosul Liwa, northern Iraq. This example, 0-57
mm. in diameter, shows over 100 canals: the arrangement may be interpreted as similar
to that in the type material.

Favreina montana sp. nov.

Plate 48, fig. 1

Description. Pellet of up to T3 mm. in length and 0-6 mm. diameter, cylindrical, with
circular or near-circular cross-section. Canals very fine, circular to comma-shaped in
section, about sixty in number, seen in some examples to be paired by very fine lines.
Pattern of canals in transverse-section not definite, but obscurely spiral.

Syntypes. BM Z997 (PI. 48, fig. 1); Qamcluiqa Formation (Albian level), Cretaceous; Gund-i-Shikavt,
Erbil Liwa, northern Iraq.

Other material. Numerous random cuts in thin-section, same locality and horizon.

Remarks. F. montana resembles the Lower Cretaceous F. cuviUieri Bronnimann in size
and arrangement of paired canals, but possesses double the number of canals. It is easily
distinguished from the third Cretaceous species occurring in the Middle East, F. kitrdi-
stanensis Elliott, which shows eight large canals thinly crescentic in section.

Specimens in the Department of Palaeontology, British Museum (Natural History), which are
referred to in the text, have the prefix BM.

REFERENCES

ANDRUSOV, D. 1 939. Role des Thallophy tes dans la constitution des roches sedimentaires des Carpathes
tchecoslovaques. Man. Soc. Sci. Boheme, ann. 1938, no. II.

C 1213

X

300

PALAEONTOLOGY, VOLUME 6

BAYER, F. M. 1956. OctocoralHa. Treatise on Invertebrate Paleontology (ed. R. C. Moore), Part F,
166-231. Lawrence, Kansas.

BONET, F. 1956. Zonificacion Microfaunistica de las Calizas Cretacicas dell Este de Mexico. Int. geol.
Congr. 20th sess., Mexico. Contr. Asoc. Mex. Geol. Petrol.

BRONNiMANN, p. 1955. Microfossils incertae sedis from the Upper Jurassic and Lower Cretaceous of
Cuba. Micropaleontology, 1, 28-51.

— ^ — and NORTON, p. 1961. On the classification of fossil fecal pellets and description of new forms
from Cuba, Guatemala and Libya. Eel. geol. Helv. 53, 832-42.

ELLIOTT, G. F. 1955. Fossil calcareous algae from the Middle East. Micropaleontology, 1, 125-31.

1956. Further records of fossil calcareous algae from the Middle East. Ibid. 2, 327-34.

1960. Fossil calcareous algal floras of the Middle East with a note on a Cretaceous problematicum

Hensonella cylindrica gen. et sp. nov. Quart. J. geol. Soc. Load. 115, 217-32.

1961. The sexual organization of Cretaceous Permocalcidiis (Calcareous Algae). Palaeontology,

4, 82-84.

1962. More microproblematica from the Middle East. Micropaleontology, 8, 29-44.

ENDO, R. 1961. Calcareous Algae from the Jurassic Torinosu Limestone of Japan. Sci. Rep. Saitama
Univ. (B), Comm. Vol. R. Endo, 55-75.

HANTZSCHEL, w. 1958. Oktokoralle Oder Lebcnsspur? Mitt, min.-geol. (St.) Inst. Hainb. 21, ll-Sl.

JOUKOWSKY, E. and favre, j. 1913. Monographie geologique et paleontologique du Saleve (Haute-
Savoie, France). Mem. Soc. Phys. Geneve, 37, 296-519.

LAUBENFELS, M. w. DE. 1955. Porifera. Treatise on Invertebrate Paleontology (ed. r. c. moore). Part E,
21-112. Lawrence, Kansas.

MASSiEUX, M. and denizot, m. 1962. Sur la valeur du genre Pseudolithothamniinn J. Pfender (Cretace-
Eocene) et son rapprochement avec le genre Ethelia Weber van Bosse (Algue Floridee Squamariacee,
actuel). C.R. Acad. Sci., Paris, 254, 2626-8.

MOORE, H. B. 1932. The faecal pellets of the Anomura. Proc. roy. Soc. Edinb. 52, 296-30S.

PAREJAS, E. 1935. L'organisme B de E. Joukowsky et J. Favre. Arch. Sci. phys. nat. (5), 17, 221-4.

1948. Sur quelques coprolithes de crustaces. Arch. Sci. Geneve, 1, 512-20.

PFENDER, J. 1937. Sur un organisme constructeur des calcaires Cretaces et Nummulitiques: Pseiido-
lithothaniniian album nov. gen. nov. sp. Bull. Soc. geol. Fr. (5), 6, 303-8.

RADOicic, R. 1959. Some problematic microfossils from the Dinarian Cretaceous. Bull. Serv. geol.
geophys. Serbie, 17, 87-92. (In Serbian).

RAMA RAO, L. and PiA, J. 1936. Fossil Algae from the uppermost Cretaceous beds (the Niniyur Group)
of the Trichinopoly District, S. India. Palaeont. indica, n.s., 21 (4), 1-49.

SEiLACHER, A. 1962. Die Sphinctozoa, cine Gruppe fossilcr Kalkschwammc. Abh. math.-nat. Kl. Akad.
(Viss. Mainz. Jhrg. 1961, 10, 721-90.

steinmann, G. 1882. Pharetronen-Studien. Neues Jb. Min. Geol. Paldont. 2, Vi9-9\ .

VOIGT, E. 1958. Untersuchungen an Oktokorallen aus der oberen Kreide. Mitt. min. -geol. (St.) Inst.
Hamb. 27, 5-49.

WEBER- VAN BossE, A. 1921. Liste des Algues du Siboga. II, Rhodophyceae. U pt. Protofloridae,
Nemalionales, Cryptonemiales. Rept. Siboga-Exped., 89.

GRAHAM F. ELLIOTT

Geological Department,
Iraq Petroleum Co. Ltd.,
214 Oxford Street,
London, W. 1

Manuscript received 1 October 1962

ON THE STRUCTURE OF LEAVES OF
RHABDOTAENIA PANT FROM THE
RANIGANJ COALFIELD, INDIA

by D. D. PANT and b. k. verma

Abstract. A new species of Rbabdotaeuia Pant (R. fibrosa) and additional details of epidermal structure of
Rhabdotaenia danaeoides (Royle) Pant, based on a number of leaf fragments, collected from the Raniganj coal-
field, India, are described. It is pointed out that these two species and R. harkinii Pant, are all structurally
distinct.

Tongue-shaped leaves found in the Glossopteris flora include, besides Glossopteris and
Gangamopteris, the three other forms, Noeggerathiopsis, Palaeovittaria, and Taeniopteris.
Pant (1958) has assigned some of these Taeniopteris leaves to a new genus Rhabdotaenia,
in which he distinguishes two species, R. danaeoides, based on three incomplete leaves
from the Raniganj coalfield, India, and R. harkinii, based on a single leaf fragment from
East Africa. A third species, R. fibrosa, which shows interstitial fibres, is described here
from the Raniganj coalfield. Besides these, additional characters of R. danaeoides are
also described from numerous compressions of leaves of the species from the same
locality.

Genus rhabdotaenia Pant

Rhabdotaenia fibrosa sp. nov.

Plate 49, figs. 1-7; text-figs. 1, 2b-e

Diagnosis. Leaf large, elongated oval, length unknown, 3-5-9-S cm. wide, lamina widest
in middle region, tapering towards apex. Apex acute, margins entire, midrib up to 3 mm.
wide in middle region, narrower towards apex, showing many longitudinal strands
towards base, fewer above. Lateral veins parallel, arising at an acute angle from midrib
and almost immediately arching outwards and running at about 90° to midrib. Vein
ends near margin sometimes slightly curved towards apex, but marginal veins or loops
not seen. About one-fifth of total number of lateral veins forked once, forking either at
point of origin from midrib or elsewhere in lamina. Lorked veins less frequent towards
apex. Concentration of veins 15-27 per cm. near midrib and 19-31 per cm. towards
margin. Veins normally up to 167 p. thick. Areas between lateral veins often showing
interstitial fibres, fibres up to 45 p thick.

Upper epidermis of lamina nonstomatiferous. Cells between veins sinuous-walled,
amplitude of sinuosities of cell walls averaging at 5 p, average lateral distance between
two antinodes 9 p. Cells rectangular, arranged in longitudinal rows parallel to veins,
typically averaging at 93 p long x 36 p wide. Average thickness of walls 5 p in middle
region and 8 p near midrib. Cells over veins narrower and longer with less sinuous sides.
Surface of cells vaguely mottled, papillae not seen.

Lower epidermis of lamina stomatiferous. Cells between veins usually averaging at
79 p long X 60 p wide, cell walls slightly more sinuous than in upper, amplitude of

[Palaeontology, Vol. 6, Part 2, 1963, pp. 301-14, pi. 49-50.]

302

PALAEONTOLOGY, VOLUME 6

sinuosities of cell walls 7 jx, lateral distance between two antinodes 15 Cells less regu-
larly arranged than those of upper epidermis and elongated in various directions. Cells
over veins narrower, elongated in direction of veins, sides of cells over veins less sinuous.
Surface of cells clearly mottled, occasionally showing median hollow papillae. Trichomes
absent. Stomata haplocheilic, unevenly distributed in areas between veins and normally
absent in vein courses. Guard cells irregularly orientated, exposed, generally 51 jx longx
\9 IX wide, stomatal opening 6 fi wide. Subsidiary cells 3-5, irregularly shaped like
ordinary epidermal cells; polar subsidiary cells unspecialized and like laterals.

Upper and lower epidermis of midrib showing rectangular or polygonal cells tending
to be arranged in longitudinal rows; cell rows forming bands of narrow almost straight-
walled cells (above longitudinal strands of midrib), alternating with bands of somewhat
wider slightly sinuous-walled cells (in areas between midrib strands). Surface of epi-
dermal cells above and near midrib often showing longitudinal surface striations.

Upper and lower cuticles of lamina like respective epidermal layers, surface mottled,
cell outlines usually well defined over veins but often obscure between them.

Holotype. Specimen No. 229 of Divya Darshan Pant Collection at present located in Botany Depart-
ment, University of Allahabad, Allahabad, India. ,

Locality ami horizon. Raniganj coalfield, India. Lower Gondwana.

Description. The species is based on nine fragments of leaves, including one apex and
eight pieces representing middle portions. None of our specimens shows the basal part
of the leaves. The single apex is acute but its concentration of veins and epidermal
structure are like those found in other fragments assigned to this species although
interstitial fibres are seen only at a few places in the apical fragment. The width of
various fragments from middle region of leaves varies from 3-5 to 9-5 cm. Lateral veins,
which run almost straight up to the margin, are 62-167 p thick. Displaced lateral veins,
like those of R. danaeoides and R. harkinii, sometimes appear sinuous (PI. 49, figs. 1, 7;
text-fig. 1h, i). In an incident-light examination, small compressed creases and folds in
the lamina may sometimes appear like anastomoses between lateral veins, but such
apparent anastomoses can be clearly made out in transmitted-light examination of
celloidin pulls, as the cells above vein courses run straight without anastomoses. Actual
connexions between veins were never seen.

Interstitial fibres, which are 22-45 p thick, come out of the lateral veins, usually in the
middle part of the lamina. For some distance the fibres run between two veins and

TEXT-FIG. I. Rhabdotaenia fibrosa sp. nov. A, Lower cuticle of lamina showing obscure epidermal cell
outlines. 23 Id; x267. b. Lower epidermis of lamina showing an enlarged stoma with lignified lateral
and polaiTamellae (the lamella at the lower pole is broken (see also PI. 49, fig. 3). 232a; x267. c. Lower
epidermis of midrib showing elongated cells above vein and longitudinal rows of slightly sinuous-
walled rectangular cells between them. 232a; x83. d. Lower epidermis of lamina with irregularly
arranged sinuous-walled cells and stomata. 232a; x83. e. Side veins of lamina (thicker lines) showing
interstitial fibres (thinner lines) between them (see also PI. 49, fig. 4). 232; x 10. f. Upper epidermis
of lamina showing longitudinal rows of almost straight-walled narrow cells above a vein and wider
cells with more sinuous sides on its two flanks. 232b; X 83. g, Broader upper epidermal cells in area
between two veins. 229; x 83. h. Apical portion of lamina showing venation (see also PI. 49, fig. 1).
233; X 1. I, Middle portion of lamina showing furcate or simple parallel lateral veins diverging at a
wide angle from longitudinally striped midrib (see also PI. 49, fig. 7). 229; X 1.

TEXT-FIG. 1

304 PALAEONTOLOGY, VOLUME 6

TEXT-FIG. 2. A, Rhabdotaenia danaeoides, b-e, Rhabdotaenia fibrosa sp. nov. A, Middle portion of
lamina showing venation (see also PI. 50, fig. 1). 251 ; x 3/2. b, Xylem element from midrib showing
scalariform thickenings. 231a; x800. c, Xylem element from midrib showing alternate biseriate pits
(see also PI. 49, fig. 5). 231a; x800. d, Xylem element from midrib showing spiral thickenings (see
also PI. 49, fig. 6). 231a; x 800. e, Xylem element from a vein showing annular thickenings (see also

PI. 49, fig. 2). 234; X800.

D. D. PANT AND B. K. VERMA; LEAVES OF RHABDOTAENIA 305

they may then take an oblique course to re-enter the same or another vein and subse-
quently they may emerge once again on the same or the opposite side. Sometimes
they meet or cross other fibres running between two veins (PL 49, fig. 4; text-fig. 1e).
Marginal fibres could not be seen clearly at any point.

Amplitude of sinuosities of cell walls in upper epidermal cells varies from 3 to 14 yn
and the lateral distance between two antinodes from 5 to 12 /x. Cells between veins are
rectangular and 62-128 /x long and 26-48 [j- wide. The thickness of their lateral walls
varies, these being generally thicker over and near midrib. The cell walls are 4-7 /x thick
in areas between veins and 6-11 /x near and over midrib.

Cells between two veins in the lower epidermis are 48-1 12 /x long and 40-80 /x wide.
Surface of cells is clearly mottled and shows well-marked lighter oval or rounded areas,
like those of R. haykinii. Surface striations are generally present in the cells over and
near the midrib. These run parallel to the longitudinal axis of cells but never radiate
from their centre. The guard cells are 37-63 [x long and 1 5-26 /x wide and the stomatal
opening is 4-7 /x. The stomata are generally exposed and not protected by subsidiary
cells as in R. danaeoides, or their papillae as in R. harkinii. The guard cells show clear
lignified lateral and polar lamellae of gymnosperm type, which are dissolved after
maceration with concentrated nitric acid and potassium chlorate, followed by ammonia.

The side veins and midrib in various leaf fragments show tracheids in abundance.
Some of them showing clear spiral (PI. 49, fig. 6; text-fig. 2d) and annular (PI. 49, fig. 2;
text-fig. 2e) thickenings, but most of them being scalariform (PI. 49, fig. 5; text-fig. 2b).
Some tracheids of the midrib also show alternate biseriate pits (PI. 49, fig. 5 ; text-fig. 2c).
The fibrovascular elements of midrib and veins sometimes show rounded or oval pinhole-
like pits as in R. danaeoides.

Comparison and discussion. Rhabdotaenia fibrosa agrees with Macrotaeniopteris feddeni
Feistmantel in shape, size, midrib characters, and above all in having about the same
concentration of veins per centimetre. Therefore, perhaps it would have simplified
matters if the present leaves were also included under the old species M. feddeni, but
epidermal structure of that species is unknown. Six specimens of M. feddeni in the
Museum of the Geological Survey of India, Calcutta (Nos. 5200, 5203, 5204, 5205,
5206, and 5498, figured by Feistmantel 1881, pi. xxi a, fig. 3; pi. xxii a, figs. 1-4; and
1886, pi. 1 a, fig. 1 respectively), were examined to see if they could yield any epidermal
and cuticular preparations, but none of them shows any carbon. All are fragments of
the middle regions of leaves impressed on ferruginous stone and even in specimens
where veins are otherwise clear, no thinner lines representing impressions of interstitial
fibres are seen (cf. Glossopteris fibrosa Pant, 1958). Accordingly, for the present we
presume that our specimens of R. fibrosa are distinct from M. feddeni. In fact we consider
it possible that two specimens (Nos. 5200 and 5498) of taeniopterid leaves, which
Feistmantel assigned to M. feddeni, may not belong to it because they are bigger than
others and show clear impressions of side veins running at an acute angle to the midrib
(cf. Palaeovittaria), instead of running at right angles to it as is typical of Rhabdotaenia.

Among comparable Lower Gondwana forms whose epidermal structure is known,
Rhabdotaenia fibrosa differs from other species of the genus in having characteristic
interstitial fibres. In addition, R. fibrosa differs from R. danaeoides in having an acute
apex (the apex in R. danaeoides is obtusely pointed). It differs from both R. danaeoides

306

PALAEONTOLOGY, VOLUME 6

and R. harkinii in having a higher concentration of veins per centimetre {R. fibrosa has
15-27 veins per cm. near midrib and 19-31 veins towards margin; R. harkinii has 15-18
veins per cm. near midrib and 24-28 veins near margin and R. danaeoides has 7-16 veins
per cm. near midrib and 8-20 veins towards margin). In 7?. harkinii about half of the total
number of veins are forked, but in R. fibrosa only one-fifth are forked as in R. danaeoides.
The epidermal structure of the three species is also quite distinct. The upper epidermal
cells in R. fibrosa are rectangular and arranged in rows parallel to the lateral veins. Their
anticlinal walls are sinuous and generally unevenly thickened, their thickness is generally
4-7 /X in areas between veins over the lamina and 6-1 1 fi in the region of the midrib.
The upper epidermal cells in R. danaeoides and R. harkinii are polygonal and irregularly
arranged. Their sides are straight. The surface of upper epidermal cells in R. danaeoides
sometimes shows obscure marks of hollow papillae, but these have not been seen in
R. fibrosa (or even in R. harkinii). The sides of lower epidermal cells in all the three
species are sinuous, but the walls in R. fibrosa and R. danaeoides show many more
smaller undulations, while the waves in walls of lower epidermal cells in R. harkinii are
fewer and larger. The figures depicting the amplitude of sinuosities and the lateral dis-
tance between two antinodes in the lower epidermis of the three species are:

Name of species

Amplitude of sinuosities

Lateral distance
between two antinodes

Range

Average

Range

Average

R. fibrosa

3-13 p

Ip

11-21 p

1 5 p

R. danaeoides

5-\6p

1 1 p

8-24 p

\2p

R. harkinii

13-28 p

\9p

20-43 p

3 1 p

Surface striations are present almost in every cell of the lower epidermis in R. harkinii.
In R. fibrosa and R. danaeoides these are present usually in the lower epidermal cells of
the midrib region. In R. harkinii the surface papillae are very prominent and generally
present in most cells of the lower epidermis, but in R. fibrosa and R. danaeoides they occur
only occasionally in the cells of the lower epidermis of the lamina. The guard cells of
stomata in R. fibrosa are always exposed and never protected by papillae of surrounding
cells as in some stomata of R. harkinii. The tracheids in the side veins of R. fibrosa show
annular and spiral thickenings besides scalariform elements like those described by Pant
(1958) in R. danaeoides and R. harkinii. Some tracheids of the midrib in R. fibrosa also
show alternate biseriate pits.

R. fibrosa may also be compared with Glossopteris fibrosa Pant on account of the
occurrence of interstitial fibres. However, it clearly differs from G. fibrosa in having side

EXPLANATION OF PLATE 49

Figs. 1-7. Rhabdotaenia fibrosa sp. nov. 1 , Apex of a leaf. 233; xl. 2, Xylem element from a lateral
vein showing annular thickenings. 234; x552. 3, Lower epidermis showing an enlarged stoma with
polar and lateral lignin lamellae and epidermal cells with sinuous anticlinal walls. 232a; X447.
4, A portion of lamina showing lateral veins running parallel to each other (thicker lines) and a few
interstitial fibres (thinner lines) between them. 232; x 15. 5, Xylem elements from midrib showing
alternate biseriate pits on the lower, left side and also scalariform thickenings elsewhere. 231a;
X 552. 6, Xylem elements from midrib showing spiral thickenings. 231a; X 1220. 7, Fragment of
middle region of a leaf. 229, xl.

Palaeontology, Vol. 6

PLATE 49

2

PANT and VERMA, Leaves of Rhabdotaenia

D. D. PANT AND B. K. VERMA: LEAVES OF RHABDOTAENIA

307

TEXT-FIG. 3. Rhabdotaenia danaeoides. A, Apical portion of leaf showing arrangement of veins (see
also PI. 50, fig. 1). 240; X 4/3. b, An epidermal cell of midrib showing surface striations. 241e; X 533.
c, Middle and basal portions of a leaf showing venation. 245; x4/3. d, Margin of leaf with veins
slightly curved towards the apex. 248d; xc. 5. e, A few cells between the veins in text-fig. 5e, more

magnified. 238p; x 267.

308

PALAEONTOLOGY, VOLUME 6

veins running at right angles to the midrib and in the absence of anastomoses between
them. Moreover, the epidermal structure of the two forms is quite distinct.

Rhabdotaenia danaeoides (Royle) Pant
Plate 50, figs. 1-2; text-figs. 2a, 3-6

Synonymy. See Pant (1958).

Emended diagnosis. Leaves oval, oblong, petiolate, 3-9 • 4 cm. wide and up to 15-9 cm.
long, widest in middle region, apex obtusely pointed, base more or less abruptly con-
tracted, margins entire, sometimes slightly undulate. Midrib up to 6 • 5 mm. wide below,
narrower towards apex, showing numerous longitudinal strands in basal part, fewer
above. Lateral veins parallel, arising at an acute angle from midrib and almost immedi-
ately arching outwards and running at about 90° to midrib, vein ends near margin often
slightly curved towards apex. At an average about one-fifth of lateral veins forked once,
forking either at points of origin from midrib or elsewhere in lamina, but mostly in
middle region; twice forked veins very rare; other veins remain unforked. Cross con-
nexions between veins present but extremely rare. Concentration of veins 7-16 per cm.
near midrib (near points of origin) and 8-20 per cm. towards margin.

Upper epidermis of lamina without stomata. Cells between veins straight-walled,
polygonal, sometimes showing obscure marks of papillae. Marginal epidermal cells
longitudinally elongated, almost straight-walled, leaf substance at margin showing elon-
gated thick- walled cells.

Lower epidermis of lamina stomatiferous. Cells between veins irregularly shaped,
64-0-1 32-8 [JL (93-3 /x) long and 35-2-92-8 /x (60-8 ix) wide, walls sinuous sometimes
showing rounded marks of median hollow papillae, becoming gradually straight near
midrib. Cells over veins narrower, elongated in direction of veins, walls almost straight.
Stomata usually confined to areas between veins, rarely over veins, somewhat unevenly
distributed, frequency about 66 stomata per sq. mm., orientation irregular. Stomata
haplocheilic. Guard cells 16-28 ^ix wide and 32-64 /x long, stomatal opening 1-6-8 p.
wide. Stomata exposed or more or less covered by subsidiary cells but never overarched
by subsidiary cell papillae. Subsidiary cells 3-5, irregular, not forming a ring, polar
subsidiary cells like laterals, size and shape of subsidiary cells similar to ordinary epi-
dermal cells.

Upper and lower epidermis of midrib showing straight-walled cells above veins and
slightly sinuous-walled cells between them, cells tending to be arranged in longitudinal
rows. Cells of lower epidermis showing fine longitudinal surface striations.

Upper cuticle of lamina showing cells with somewhat obscure outlines, cell walls thin.
Lower cuticle of lamina stomatiferous like lower epidermis. Cells between veins with

EXPLANATION OF PLATE 50

Figs. 1, 2. Rhabdotaenia danaeoides. 1, Apical and middle portion of a leaf showing venation. The
lighter patches are caused by the removal of carbon in pulls. 240; X 1. 2, Lower epidermis show-
ing an exposed stoma. 245b; x471.

Figs. 3, 4. Rhabdotaenia harkinii; V34454. 3, Lower epidermis showing sinuous-walled cells and
irregularly placed protected stomata, X 193. 4, Lower epidermis with papillate epidermal cells and
an exposed stoma, x 400.

Palaeontology, Vol. 6

PLATE 50

PANT and VERMA, Leaves of Rhahdotaenia

• ‘’i!-

t

\

c

D. D. PANT AND B. K. VERMA: LEAVES OF RHABDOTAENIA

309

TEXT-FIG. 4. Rhabdotaenia danaeoides. A, Lower epidermis with two stomata, one (left) showing
lignified polar lamellae on the two poles and the other (right) showing a hollow mark of papilla in
one of its subsidiary cells. 238a; X400. b, Lower cuticle showing a stoma and obscure sinuous-walled
cells (under phase-contrast microscope). 308n; x301. c, Lower epidermis showing sinuous-walled
cells. The stomata are irregularly placed in between the veins. 238a; xl25.

310

PALAEONTOLOGY, VOLUME 6

obscure cell walls usually appearing broken, but sometimes showing sinuous waves ; cell
walls below veins less obscure, straight.

Neotvpe. Brit. Mus. (N.H.) Palaeont. Dept. No. V4191. Figured Royle (1833, pi. 2, fig. 9) and Arber
(1905, pi. 5, fig. 1).

Isotypes. Specimen Nos. 237-330 deposited in the Divya Darshan Pant Collection, at present located
in Botany Department, University of Allahabad.

Locality and horizon, Raniganj coalfield, India, Lower Gondwana.

Description. The emended diagnosis is based on ninety-five leaf fragments from the
Raniganj coalfield, India (including five apices, one base, and eighty-nine pieces from the
middle region of leaf laminae). Some of the leaves appear to have been quite narrow,
about 3 cm., but others were up to 9-4 cm. wide. The midrib was up to 6-5 mm. wide in
the basal part. Lateral veins, which run almost straight up to the margin, are sometimes
slightly curved upwards towards the apical region (text-fig. 3d). In places the veins
appear sinuous (PI. 50, fig. 1 ; text-figs. 2a, 3a), but, as already pointed out by Pant (1958),
such sinuosities are obviously due to a displacement of the veins inside the epidermal
and cuticular coat of the leaf, when its more delicate mesophyll tissue has partially or
wholly dissolved.

The margins of simple spathulate leaves are seldom torn. In the absence of regular
anastomoses between the side veins, like those found in Glossopteris and Gangamopteris,
it was difficult to understand how such large laminae remained intact, but a specimen
with a well-preserved margin reveals that this was possibly due to the presence of longi-
tudinal strands of thick-walled cells running parallel to the margin (text-fig. 6b, c). The
thick-walled marginal cells appear to be fibrous, but at one point (text-fig. 6c) their walls
show pits like those of xylem, suggesting that there could even be some marginal
tracheids present among them.

The number of bifurcated veins is usually higher in the middle region. The concen-
tration of veins per cm. is higher in the apical and basal portions (at the point of their
origin from the midrib). In different leaf fragments the ratio of once-forked lateral veins
and those remaining unforked ranges from one-third to one-fifteenth.

The typical sinuous-walled cells of the lower epidermis are clearly seen in most of the
specimens, but fragments of the upper epidermis are seen only in one specimen. The cells
of upper epidermis seem to be relatively less sinuous-walled or with almost straight walls
like the cells over the meshes of the veins in the midrib (text-fig. 3e).

The lateral veins and midrib show abundant tracheids. Some of them have spiral
(text-fig. 6f) and reticulate thickenings (text-fig. 6e), but most of them are scalariform
(text-fig. 6a). Along with the typical xylem cells of the veins and the midrib are seen some
thick-walled elongated elements with oval or rounded pinhole-like pits. They may repre-
sent fibres or tracheids (text-fig. 6d, and also Pant 1958).

Comparison and discussion. Pant (1958) has already discussed the differences between
Rhabdotaenia and other leaves of Taeniopteris type which possess haplocheilic stomata,
such as Doratophyllum (Harris 1932) and Bjuvea (Florin 1933). The Jurassic leaves of
NipaniophyUum raoi (Sahni 1948) also have haplocheilic stomata. Like Rhabdotaenia,
their midribs contain a number of parallel veins. However, they belong to a different age
and their size, form, and epidermal structure are clearly different. Moreover, they occur

TEXT-FIG. 5. Rhabdotaenia danaeoides. a, Lower epidermis with a single enlarged stoma showing two
lateral lignin lamellae. 241a; x400. b, Upper cuticle showing obscure outlines of cells (under phase-
contrast microscope). 308j; X 144. c, Epidermis of midrib showing straight-walled cells arranged in
longitudinal rows. 238e; X 125. d, Upper epidermis showing straight-walled cells some of which show
median papillae. 238a; X 125. e, Epidermis of midrib showing straight-walled cells, above its parallel
veins and slightly sinuous-walled cells between them. All cells tend to be arranged in longitudinal
rows. A few cells of the fibrovascular midrib bundles are also seen. Note the simple pits and scalari-
form thickenings in some of these. 238p; Xl25.

PALAEONTOLOGY, VOLUME 6

312

TEXT-FIG. 6. Rhabdotaenia danaeoides. A, Xylem of midrib showing scalariform elements. 238p; x800.
B, Portion of a lamina showing thick-walled marginal cells arranged parallel to the margin of leaf.
261a; x 125. c, A portion of b, more magnified. 261a; x400. d, Fibrovascular elements from midrib
showing pinhole-like pits. 275; X 800. e, Xylem elements from a vein showing reticulate and scalari-
form elements. 264; x800. f, Xylem elements of a vein showing spiral elements. 275; x800.

D. D. PANT AND B. K. VERMA: LEAVES OF RHABDOTAENIA

313

associated with their characteristic fructifications, the like of which have never been
reported associated with Rhabdotaenia. The mode of preservation of the two leaves is not
comparable and we therefore assume that they are clearly different. Pant (1958) has also
compared Rhabdotaenia with other Lower Gondwana leaves, but their cuticular structure
is unknown.

The general form and epidermal structure of Rhabdotaenia are nevertheless comparable
with those of Glossopteris, Gangamopteris, and Palaeovittaria, and it is quite likely that
Rhabdotaenia too will ultimately prove to belong to a plant of the same general alliance.

Pant (1958) distinguished the two species Rhabdotaenia danaeoides and R. harkinii, on
the basis of concentration of veins per centimetre, the relative frequency of forking veins
and certain epidermal characters. The epidermal structure of R. danaeoides wsis, however,
described by Pant from very small bits of epidermis, and the range in the concentration
of veins and other characters was also unknown in both the species. Our study of ample
material of R. danaeoides has shown that the concentration of veins in this species is
never so high as in R. harkinii. R. danaeoides has on an average about one-fifth of the
lateral veins which fork once and only a single vein in one specimen is twice-forked, but
in R. harkinii about half of the total number of veins are once-forked and a number of
them twice. Among other differences between the two species is the usual occurrence of
a median prominent papilla and surface striations in the cells of R. harkinii. Striations
and surface papillae are present in R. danaeoides as well, but in this species papillae occur
relatively rarely in the lower epidermal cells and surface striations are usually restricted
to the lower epidermal cells of the midrib. The anticlinal walls of the epidermal cells
show numerous small waves in R. danaeoides. The waves of the walls in R. harkinii are
fewer and larger. The amplitude of sinuousities in the cell walls of lower epidermal cells
and the distance between two antinodes of sinuous-walled cells are also different in both
the species (see above). The stomata of R. harkinii are usually protected by papillae of
the surrounding subsidiary cells (PL 50, fig. 3) which may be drawn apart and expose the
underlying guard cells (PI. 50, fig. 4) or they may partially or wholly cover the stoma
(Pant 1958, pi. 20, fig. 5). The stomata in R. danaeoides are never so well protected. The
surface of epidermal cells in R. harkinii shows well-marked lighter oval or rounded areas
which may be surface pits like those present in the Cycadales (Pant and Nautiyal 1963).
In R. danaeoides such areas are seemingly present but less marked.

Acknowledgements. We are grateful to Sri A. P. Sharma and Sri B. K. Palit for their generous help
in collecting fossils in their respective collieries in the Raniganj coalfield, India. The material was
collected jointly by a party consisting of the authors and Prof. H. N. Andrews, Dr. D. D. Nautiyal,
Dr. G. K. Srivastava, and Miss Bharati Mehra. The authors are thankful to their other colleagues of
the collection party for help in picking up specimens of Rhabdotaenia. We are thankful to Prof.
P. Maheshwari, Chairman of the Biological Committee of the C.S.I.R., for his keen interest in the
progress of the scheme under which the present work has been carried out. Our thanks are also due
to Sri M. V. A. Sastry, Palaeontologist-in-charge, Geological Survey of India, Calcutta, for permission
and help in examining the type and syntype material of Macrotaeniopteris feddeni and Taeniopteris
danaeoides, in the Survey collections. Lastly we thank the Council of Scientific and Industrial Research
for financial assistance and the award of a Junior Research Assistantship to one of us (B. K. V.).

REFERENCES

ARBER, E. A. N. 1 905. Catalogue of the Fossil Plants of the Glossopteris Flora in the Department of Geology,
British Museum {Natural History). Ixxiv -1-255 pp.

314

PALAEONTOLOGY, VOLUME 6

FEiSTMANTEL, o. 1880-1. The fossil flora of the Gondwana System. The flora of the Damuda and
Panchet Divisions. Palaeont. indica, 3 (12), 1-149.

1886. The fossil flora of the Gondwana System. The fossil flora of some of the coalfields in

Western Bengal. Ibid. 4 (12), 1-66.

FLORIN, R. 1933. Studien iiber die Cycadales des Mesozoikums. Nebst Erorterungen fiber die
Spaltoffniingsapparate der Bennettitales. K. Sveiiska Vetensk. Akad. Hand!. 12 (5), 1-134.

HARRIS, T. M. 1932. The fossil flora of Scoresby Sound, East Greenland. 2. Description of seed plants
Incertcie sedis, together with a discussion of certain Cycadophyte cuticles. Medd. Grenland. 85 (3),
1-112.

PANT, D. D. 1958. The structure of some leaves and fructifications of the Glossopteris flora of Tanga-
nyika. Bull. Brit. Mils. (N.H.) Geol. 3 (4), 127-75.

and NAUTIYAL, D. D. 1963. Cuticular and epidermal studies of some modern cycadean leaves,

sporangia and seeds. Seiickenbergiana (in press).

ROYLE, J. F. 1 833. Illustrations of the Botany and Other Branches of the Natural History of the Himalayan
Mountains and of the Flora of Cashmere. 2 vols., lxxxii + 472 pp.

SAHNi, B. 1948. The Pentoxyleae; a new group of Jurassic gymnosperms from the Rajmahal Hills
of India. Bot. Gaz. 110 (1), 47-80.

D. D. PANT
B. K. VERMA

Botany Department,
University of Allahabad,

Manuscript received 4 August 1962 India

A PALAEOGENE TEREDINID (MOLLUSCA)

FROM IRAQ

by GRAHAM F. ELLIOTT

Abstract. The teredinid mollusc Bankia (Bankiella) kurdistanensis sp. nov. is described from an association
of tubes, valves, and pallets in the Palaeocene of Iraq.

Wood bored by Teredo and allied genera is a common Tertiary fossil at various levels
and localities, as in the London Clay of England. Current zoological classification of
these highly modified molluscs is based neither on the valves nor on the conspicuous
tubular shell-lined borings, but on the morphology of the little calcareous structures
known as pallets, which in life are associated with the fleshy siphons. Such pallets are not
commonly recorded as fossils. Although the Iraqi material described below was in solid
preservation not permitting the isolation of pallets, numerous thin-sections show these
and other structures excellently preserved.

The specimens, of petrified dicotyledonous wood showing well-preserved structure
and riddled with calcite-filled teredinid burrows, were collected from the Kolosh Forma-
tion (van Bellen 1959). This is composed of sands and sandstones of detrital serpentine,
chert, and radiolarite, with subordinate shales and limestones, and outcrops extensively
in northern Iraq. It has yielded foraminiferal faunas and an algal microflora of Palaeo-
cene-Lower Eocene age, as well as some molluscs and other macrofossils. The present
material comes from three separate localities, but all the pallets seen in sections are of
one distinctive type, suggesting that one species only is present.

In a revision and classification of the Teredinidae, Bartsch (1922) subdivided the genus
Bankia Gray 1 842, which comprises those forms whose pallets show cone-in-cone struc-
ture. Such pallets consist of a proximal rod-like portion or stem, from which a series of
laterally flattened cones, opening distally, arise one within the other, together forming
a blade-like structure. In the subgenus Nausitora the cones are fused on one flattened
side and backed by a calcareous deposit (‘thick periostracum’ of Bartsch). In the three
subgenera Bankia s.s., Bankiella, and Neobankia the cones are free and covered by a thin
investing membrane; distally this membrane is fimbriated in Bankia s.s., entire in Ban-
kiella, and denticulated in Neobankia.

Nausitora-ysiWQis showing calcareous fusion might be expected to occur as fossils and
have in fact been recorded from the Lower Eocene London Clay (Wrigley 1930). The
Iraqi fossils now discussed show pallets with free cone-in-cone structure and entire
margins. It would at first sight seem doubtful that the membrane permitting subgeneric
differentiation would be preserved, and hence that Bankia s.l. would be the best allocation
that could be made. However, Stinton (1957) described denticulated-margin pallets
recognizable as Neobankia from the Upper Eocene Barton Clay. This preservation is
probably inferior in fine detail to that of the material now examined. It seems likely,
therefore, that the entire margins seen are original, and for this reason the fossil is
referred to Bankiella.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 315-17, pi. 51-52.]

C 1213 Y

316

PALAEONTOLOGY, VOLUME 6

Genus bankia Gray 1842
Subgenus BankieUa Bartsch 1921
Bankia {BankieUa) kurdistanensis sp. nov.

Plates 51, 52

Diagnosis. Bankia with pallets of 4-3 mm. length or more, maximum diameter about
2-5 mm., of flattened cone-in-cone type, about fifteen cones with distal margins entire,
free stem presumed relatively short. Valves incompletely known; tubes typically variable
in length and diameter, incipiently septate in large examples.

Holotype. BM LL30332 (PI. 51, fig. 1), Kolosh Formation (Palaeocene), Dohuk, Mosul Liwa,
northern Iraq. Paratypes. BM LL30333-5 inch (PI. 51, figs. 2, 3; and PI. 52, figs. 1, 2), Kolosh
Formation (Palaeocene), Dohuk, Mosul Liwa, and Koi Sanjak, Erbil Liwa, northern Iraq.

Other material. Specimens from the Kolosh Formation (Palaeocene), of Shaqlawah and Koi Sanjak,
Erbil Liwa, and Dohuk, Mosul Liwa, northern Iraq.

Description, {a) Tubes. The calcite- or matrix-filled shelly tubes show great variety both
in size and degree of crowding. In diameter they vary from 1-5 to 12-0 mm. The larger
examples may have reached a length of 20 cm. or more, though this cannot be proved
from the pieces of bored wood collected. These specimens show all the usual varieties of
wood penetrated by straight parallel tubes, small, crowded, and twisted tubes, &c. Some
larger tubes show a regular, narrowly annular internal septation (PI. 52, fig. 2); in an
example of 5 mm. tube-diameter the septa are 2 mm. apart. This feature is developed in
individuals of some Recent teredinid species. In one fossil tube the section shows two
smaller, conjoined but complete tubes, associated with matrix. This is presumably the
broken apertural end, out of position: such a constricted double apertural siphonal tube
has been described from the London Clay by Sowerby (1815), Davis (1936), and Wrigley
(1940); the last, by analogy with Recent species, did not regard it as of specific value.
Only two examples have been seen in the Iraqi material (BM LL30331).

{b) Valves. The valves are known only from cross-sections of two paired examples
(PI. 51, fig. 2; PI. 52, fig. 1). Presumably they were of the specialized pattern common to the
family (Bartsch 1922, Stinton 1957), functioning as excavating tools. The distinctive

EXPLANATION OF PLATE 51

Bankia (BankieUa) kurdistanensis sp. nov., thin-sections, Xl5. Kolosh Eormation, Palaeocene;

Dohuk, Mosul Liwa (fig. 1); and Koi Sanjak, Erbil Liwa, northern Iraq (figs. 2, 3).

Eig. 1. Vertical section of pallet (in two pieces) at right angles to maximum width of the pallet blade.
In calcite-filled burrow, in dicotyledonous wood. Holotype, BM LL30332.

Eig. 2. Oblique section through blade of pallet, showing stem, cones, and concave pallet-face; also
the two valves. Post-mortem association in burrow. Paratype, BM LL30334.

Pig. 3. Oblique and transverse sections of two adjacent pallets in burrow. Not in position held during
life. Paratype, BM LL30334.

EXPLANATION OF PLATE 52

Bankia (BankieUa) kurdistanensis sp. nov., thin-sections, Xl5. Kolosh Eormation, Palaeocene;

Dohuk, Mosul (fig. 1); and Koi Sanjak, Erbil Liwa, northern Iraq (fig. 2).

Pig. 1 . Approximately vertical section of two associated valves, one showing well the external ridges
used for mechanical excavation. In burrow, in wood. Paratype, BM LL30333.

Fig. 2. Portion of longitudinal section of large burrow, showing spaced annular septa on inside of
shelly lining to burrow (conspicuous calcite parting on one side). Paratype, BM LL30335.

Palaeontology , Vol. 6

PLATE 51

ELLIOTT, Palaeocene teredinid

Palaeontology, Vol. 6

PLATE 52

ELLIOTT, Palaeocene teredinid

G. F. ELLIOTT: A PALAEOGENE TEREDINID

317

fine concentric ridges (‘dental ridges’ of part of the outer valve surface) are seen in Plate
52, fig. 1 .

(c) Pallets. Thin-sections reveal numerous pallets, single or paired, occurring usually
at the inner, closed end of the burrows. This is not the position in life, when the pallets
are adjacent to the siphons at the mouth of the burrow, but presumably most such fossil
pallets did in fact belong to the individual inhabiting the burrow in which they now
occur. In each of four random cuts the count of component cones is about fifteen; these
structures are seen to be closely set, laterally flattened, with one face convex and the
other slightly concave with median convexity over the stem. The only possible trace of
accessory detail on the oblique cuts through the margins, which were entire, is seen at the
lateral margins, which may have been produced into spines as in Recent species. The
length of the free stem, proximal to the cones or blade of the pallet, is not known, but
as it is infrequently seen in random cuts of different burrows, as compared with the
blades, it is suggested that it was relatively short. There is great variation in the lengths
of free stem to blade in the pallets of different Recent teredinid species.

A Recent Bankiella species with pallets of about the same size is Bankia {BankieUa)
edmondsi (Balakrishnan Nair 1956), from the Madras coast, India. This agrees in the
close-set cones (eleven in the type specimen) with entire margins except for lateral spines,
but possesses a proportionally longer stem and a narrower blade.

Pallets of Naiisitora, Neobankia, and Bankia s.l. are all known from the English
Eocene, as well as those of Teredo (Psdoteredo) and remains of the pallet-less Teredina
(Wrigley 1930, Stinton 1957; see also Davis 1936, Moll 1942), so the present occurrence
of BankieUa in the Iraqi Palaeocene is in keeping with an early Tertiary differentiation
of the group.

Specimens in the collections of the Department of Palaeontology, British Museum (Natural History),
which are referred to in the text have the prefix BM.

REFERENCES

BALAKRISHNAN NAIR, N. 1956. Shipwomis from India. 1. Report on ten species of shipworms from
the Madras coast. Rec. Indian Mus. 52, 387-414.

BARTSCH, p. 1922. A monograph of the American shipworms. Bull. U.S. nat. Mus. 122, 1-51.

DAVIS, A. G. 1936. The London Clay of Sheppey and the location of its fossils. Proc. Geol. Ass., Land.
47, 328-45.

MOLL, F. 1942. Die fossilen Terediniden und ihre Beziehung zu den rezenten Arten. Palaeontographica,
94A, 134-53.

sowERBY, J. 1815. The Mineral Conchology of Great Britain, 1 (1812-15), 234, pi. 102. London.
STINTON, F. c. 1957. On the occurrence of Teredinidae in the Upper Eocene of Barton, Hampshire.
Proc. tnalac. Soc. Lond. 32, 167-73.

VAN BELLEN, R. c. 1959. Tertiary. In Lexique stratigraphique internat. 3 (Asie), fasc. lOn (Iraq).
WRIGLEY, A. 1930. Notes on English boring Mollusca, with descriptions of new species. Proc. GeoL
Ass., Lond. 40, 376-83.

1940. The faunal succession in the London Clay, illustrated in some new exposures near London.

Ibid. 51, 230-45.

GRAHAM F. ELLIOTT

Geological Department,
Iraq Petroleum Co. Ltd.,
214 Oxford Street,
London, W.l

Manuscript received 1 October 1962

THE RECOGNITION OF SALINITY-CONTROLLED
MOLLUSC ASSEMBLAGES IN THE GREAT
ESTUARINE SERIES (MIDDLE JURASSIC)

OF THE INNER HEBRIDES

by J. D. HUDSON

Abstract. The Great Estuarine Series was deposited in large, shallow lagoons. Its abundant molluscan faunas
are restricted to very few genera and species, several of which are similar to modem fresh- and brackish-water
forms. Consideration of the nature and history of the brackish-water fauna, together with direct comparison
with the modem brackish-water environments of the Texas coast, results in an interpretation of the faunas as
being controlled very largely by salinity variations in time. The salinities ranged from freshwater to fully marine,
but were usually intermediate.

The purpose of this paper is to show that the molluscan faunas of the Great Estuarine
Series lived in brackish waters in a lagoonal environment, and that it is possible to
recognize a series of overlapping faunal assemblages whose composition and distribution
were controlled largely by salinity variations in time. The detailed evidence on which this
thesis is based, the stratigraphical distribution of the faunas, and the ecology of the fossils
concerned, is described in a second paper (this journal, p. 327).

THE GREAT ESTUARINE SERIES

The stratigraphy of the Great Estuarine Series (Upper Bajocian-Bathonian) has been
described elsewhere (Hudson 1962); the Staffin Bay Beds (Upper Bathonian?-Lower
Callovian) of Trotternish are not now included in the Great Estuarine Series. The ‘ Series ’
has long been recognized as of unusual facies, ‘estuarine’ in the wide sense of nineteenth-
century geologists (Judd 1873, 1878), since it bears unmistakable signs of deposition in
very shallow water, and has a fauna which is impoverished in numbers of species but
very abundant in numbers of individuals.

Depositional environment. The conditions of deposition of the Great Estuarine Series
naturally varied from one formation to another, but some generalizations can be made.
The characteristic feature of all the shale and limestone formations is that lithologies and
faunas may vary very rapidly from bed to bed up the succession, but are laterally per-
sistent; some beds can be traced for several miles. There is abundant evidence of very
shallow-water conditions throughout, with frequent mudcracked surfaces, but the area
never became a land surface. These features, together with the brackish-water nature of
the fauna, are best explained if deposition was in extensive shallow lagoons, probably
partially separated by a bar from the open sea. The sandy formations represent the
subaqueous portions of deltas built into these lagoons, whieh persisted throughout the
deposition of the ‘Series’. Such lagoons are very favourable to the establishment of more
or less stable brackish-water conditions; the best modern examples — the bays of the
Texas coast — are discussed below.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 318-26.]

J. D. HUDSON: SALINITY-CONTROLLED MOLLUSC ASSEMBLAGES 319

Nature of the fossil assemblages. Most of the fossils of the Great Estuarine Series are
not in position of life. They occur as accumulations of dead shells, probably winnowed by
wave action, with the valves of lamellibranchs disarticulated. However, the shell beds are
enclosed in fine-grained shales, and there is seldom any evidence of strong currents or of
long distance derivation. So that, although the shell beds are death assemblages, they
are indigenous death assemblages in the sense of Hallam (1960), and represent the general
environment in which they are found, though small-scale patchiness of distribution may
have become blurred. The uniformity of composition of many of the shell beds, especially
some of the more obviously drifted ones, supports this interpretation; and adjacent shell
beds, separated by only a few inches of shale, may have different compositions. There is
much less need to invoke faunal mixing to explain assemblages than might be expected.

SOME ASPECTS OF BRACKISH WATER FAUNAS

The chemical and biological classification of brackish waters has been reviewed by
Hiltermann (1949); and Table 1 is slightly modified from his paper. The biological

TABLE 1. Classification of brackish water. Mainly after Hiltermann {1949)

Biological classification
(mainly after Reniane)

Descriptive term

Total salinity
(parts per thousand)

Freshwater

Freshwater

0-0-5

Freshwater province with reduced number of
species

Oligohaline brackish water

0-5-3

Brackish freshwater

Miohaline brackish water

3-5

Typical brackish water

Mesohaline brackish water

5-9

Brackish-marine

Pliohaline brackish water

9-16-5

Marine province with reduced number of species

Brachyhaline sea water

16-5-30

Marine

Sea water

30^0

(Not considered by Remane)

Hypersaline water

greater than 40

classification is based mainly on the work of Remane at Kiel (see below); it assumes,
of course, reasonable constancy of other environmental conditions. When used of con-
ditions (e.g. brackish lagoon), the word ‘brackish’ usually implies a fairly stable inter-
mediate salinity, contrasted with the tidally fluctuating salinity of an estuary, but never
as stable as that of the sea. The following discussion on modern brackish-water faunas
is based mainly on review articles on the fauna in general (Gunter 1947, Hiltermann
1949, Schmidt 1951, Pearse and Gunter 1957, Hedgpeth 1957) and on special areas
(Segerstrale 1957, Sorgenfrei 1958 on the Baltic, Ladd et al. 1957, Parker 1959, Shepard
et al. 1960 on Texas coast bays).

The most obvious and well-known characteristic of brackish-water faunas, whether
stable-brackish or estuarine, is their paucity in numbers of species, contrasted frequently
with great abundance of individuals. Temperature effects, oxygen deficiency, and
extreme exposure to waves can cause a similar restriction, but can frequently be elimi-
nated as an important cause by regional studies (Sorgenfrei 1958). The explanation of this
effect is that, while brackish waters are often rich feeding grounds, few animals are able
to use them because of physiological difficulties, mainly of osmoregulation, so that those
few which have overcome the difficulties multiply greatly.

320

PALAEONTOLOGY, VOLUME 6

Two Other points from the literature are relevant:

1. Even at quite low salinities the brackish-water fauna is predominantly reduced-
marine; hence the minimum number of species is found, not midway between marine
salinity and freshwater, but at a salinity of around 5-9%o. This was first established by
Remane at Kiel, and the general result seems to be accepted (Hedgpeth 1957, pp. 702-3).

2. The brackish-water fauna is recruited from a small part only of the marine fauna.
Whole phyla or other major taxa are stenohaline (corals, echinoderms, cephalopods,
articulate brachiopods). Even among the molluscs and the crustaceans, which are the
most important groups in brackish waters today, only a few relatively small groups
produce brackish-water forms in any numbers.

Rapid fluctuations in salinity are more deadly to marine animals than slow ones;
hence, for a given gradient of average salinities, estuaries will show a sharper faunal
reduction than stable brackish waters. Stable brackish waters are also more liable to
invasion, at low salinities, by freshwater forms (e.g. Unio at salinities of up to 3-4%o in
the Baltic area). Many animals of brackish lagoons also live in near-by hypersaline ones.
‘This suggests that once an organism is capable of adjusting itself to salinity changes its
range of tolerance may exceed the usual changes in its immediate environment, so that
salinity is not the governing factor and the organism is able to meet unusual conditions,
and colonise extreme environments’ (Pearse and Gunter 1957, p. 147).

This may also help to explain the second observation, that the brackish-water fauna
comes from only a few invertebrate taxa. Few organisms seem actually to require the
fluctuating conditions always associated with low salinity, but once an organism can
withstand them it has a permanent advantage in an evolutionary sense (Hedgpeth 1957,
p. 695). Since brackish-water animals can by definition withstand large short-period
oscillations, usually of temperature and depth as well as of salinity, they are better
equipped to withstand small secular changes than animals adapted to a narrower range
of conditions. These less-adaptable animals must either evolve distinct features to
encounter the changed environment, or become extinct. Therefore estuarine (and
brackish- water) species are often conservative forms with a long history (Hedgpeth 1957,
p. 696). Hedgpeth quotes the oysters as a typical case, and concludes that ‘euryhalinity
runs in families’.

Hutchinson (1960) makes the further point that freshwater faunas, too, are very long-
ranging. He estimates that only 20-30 separate invasions from the sea are required to
account for all the flourishing molluscan faunas of modern lakes and rivers. Their
inability to re-enter any but the most dilute of brackish waters makes these freshwater
forms extremely valuable environmental indicators.

The recognition of fossil brackish-water faunas. An admirable review of this problem has
been given by Schmidt (1951). This has the encouraging result of reaffirming most of the
traditional criteria for brackish water; Schmidt concludes that the following four criteria
should be diagnostic of fossil brackish-water faunas, and hence deposits.

1 . Paucity of species, often forming monotypic shell beds.

2. Absence of stenohaline groups.

3. Presence of euryhaline animals, including special brackish- water forms, notably
lamellibranchs (and also some gastropods) among the macrofauna, and, among the
hard-shelled microfauna, particularly ostracods.

J. D. HUDSON: SALINITY-CONTROLLED MOLLUSC ASSEMBLAGES 321

4. Association of marine and freshwater (not hypersaline) deposits. This means con-
sidering the palaeogeography, e.g. hypersaline deposits should be associated with
evaporites, true brackish deposits with deltaic sands and drifted plant remains.

Am.ong general conditions mentioned by Schmidt as favourable for the production of
brackish-water areas are a wet climate in the source area, leading to high run-off ; and
in the depositional area, leading to small evaporation. The absence of strong sea currents
near the river mouths is favourable; hence lagoons and restricted gulfs are often brackish.
If brackish water is deduced from other considerations, those conditions are corre-
spondingly indicated, a fact which is obviously useful in palaeogeography. Schmidt
gives a review of brackish-water faunas and sediments in the stratigraphical column,
and remarks that their recognition back, as he claims, to the Lower Palaeozoic suggests
that the salt sea is very ancient. As a result of their review, Pearse and Gunter (1957,
pp. 131-3) also conclude that the salinity of the open ocean has been fairly constant.
For the following discussion, which is mainly concerned with relative salinity, it will
therefore be assumed that the Jurassic ocean, like the modern one, had a salinity of 35%q.

THE GREAT ESTUARINE SERIES AS A BRACKISH- WATER DEPOSIT

The Great Estuarine Series, taken as a whole, admirably fits Schmidt’s criteria for a
brackish-water deposit.

1 . The total number of macro-invertebrate species recorded is less than 50, compared
with over 200 species each of gastropods and lamellibranchs as well as numerous
brachiopods, echinoderms, &c., from the English Great Oolite (Cox and Arkell 1948-
50). This result is too extreme to be explained entirely by deficient collecting. Monotypic
shell beds are extremely characteristic.

2. A few feet of strata in the Lower Ostrea Beds are responsible for nearly all records
of stenohaline forms from the Series — one echinoid plate and spine, one polyzoan, one
species of articulate brachiopod, a few records of foraminifera. Corals and cephalopods
are completely unknown.

3. Lamellibranchs, less often gastropods, ostracods, and Euestheria {Estheria auct.),
or some combination of these groups, dominate the fauna at all horizons. They include
several well-known brackish- water forms.

4. Marine beds occur above and below the Series; one or two freshwater beds occur
within it. Drifted plant remains are common in the sandstones. Evaporites are absent
throughout the British Middle Jurassic, and plant remains and deltaic deposits are
common.

The conclusion that the Great Estuarine Series is a brackish- water deposit is not new;
Hugh Miller (1858, especially p. 55) has a very well-argued passage on the subject. It has
been deduced without subdividing the Series, and without considering the fauna except
in the most general terms of phyla and classes.

The appropriate taxonomic level at which to discuss the fauna in more detail is that
of rather broad genera. It is immediately striking that, of the small total number of
genera in the Great Estuarine Series, several are still living in brackish or fresh waters;
they are plausibly regarded as members of those conservative euryhaline or freshwater
groups discussed above. Liostrea {Ostrea s.L), Mytiliis, Utiio, Viviparus, and Euestheria
are the best known. Not only do all these occur in the same group of rocks, which in

322

PALAEONTOLOGY, VOLUME 6

itself strengthens the ‘conservative’ hypothesis, but when separate formations are con-
sidered, and still more when the rocks are examined bed by bed, it is found that they
do not occur indiscriminately mixed. Liostrea hebridica and Mylilus strathairdensis,
which are members of marine-euryhaline genera, both occur most typically in mono-
typic shell beds, recalling modern oyster and mussel beds. Liostrea also occurs in the
marine horizon mentioned above, with genera usually regarded as fully marine {Rhyn-
chofiella, Myopholas, Anisocardia) in assemblages very similar to those of the fully
marine Great Oolite. Unio and Viviparus, on the other hand, which are freshwater at
the present day, do not occur with the marine genera, nor with Liostrea or Mytilus, but
they do occur together. Their other main associate is the extinct lamellibranch Neomio-
don, which forms with Unio and Viviparus the well-known Hastings Beds assemblage
of the Wealden, which has always been regarded as freshwater. Neomiodon, however,
also occurs with the more marine genera mentioned above. It seems to have been
euryhaline, though as usual with such forms it is most abundant numerically in low-
salinity assemblages, often forming monotypic shell beds. A frequent associate of Neo-
miodon and Viviparus is Euestheria, which inhabits fresh or rarely brackish water today.
Ostracods are also most abundant in the beds with Euestheria and Neomiodon.

It is therefore possible to divide the Great Estuarine Series assemblages at least into
more and less saline, as briefly indicated by Anderson (1948) and others, and perhaps
to arrange them in a salinity series. On the whole the Estheria Shales, the Concretionary
Sandstone Series, and the Ostracod Limestones have a low-salinity fauna, and the
Lower Ostrea Beds and the Staffin Bay Beds (= Upper Ostrea Beds and Belemnite
Sands, of Anderson and Cox 1948) a high-salinity one. The Mytilus Shales, and parts
of the Estheria Shales and Lower Ostrea Beds, show rapid alternations within a few
feet (or sometimes a few inches) from one type of fauna to the other, within a fairly
constant gross lithology. These reversed and repeated changes rule out evolution or
long-range immigration as explanations of the faunal changes, and render large changes
of depth most improbable. Salinity changes seem much the most reasonable explanation.
Salinity-controlled assemblages of similar type are known in the Baltic and in the Texas
Bays, where conditions more closely approach those envisaged for the Great Estuarine
Series.

THE TEXAS COAST BAYS AND THEIR MOLLUSCAN FAUNA

These bays have been intensively studied in the last few years, and the results have
now been collected and summarized in Shepard et al. (1960). The earlier reports most
relevant to the present topic are by Ladd et al. (1957) and Parker (1959). The situation
is intermediate between that in large masses of stable brackish water, such as the Baltic,
and that in a typical estuary.

The bays form a complex series of estuaries and lagoons behind a string of sandy
barrier islands, through which are narrow connexions to the Gulf of Mexico. The
climate in the area varies from humid in the north-east, near the Mississippi delta, to
semi-arid in the south-west. The whole system is extremely shallow, less than 9 ft., but
the bays rarely dry up. Tides are negligible. The total salinity variation is large, from
virtually fresh to hypersaline, but in any one part of the system may be constant for
several years. Atypical pattern is a slow build-up to high salinities in a spell of dry years,
leading to an invasion of marine forms, followed by sudden freshwater floods. These

J. D. HUDSON: SALINITY-CONTROLLED MOLLUSC ASSEMBLAGES 323

have been recorded as reducing salinities from 40%,, to 2-4%o in a few weeks, causing
mass-mortalities of the marine forms. The sediment is mainly mud, stratified in the bay
heads where the fauna is sparse, structureless in the lower bays. The sediment of the
barrier islands and passes is more sandy and contains echinoid remains.

The distribution of both micro- and macrofaunas is controlled very largely by salinity,
or at least parallels the salinity contours. Minor variations due to the nature of the
bottom, &c., are superimposed on this salinity-controlled pattern. The fauna of the
waters of very low salinity is very poor in numbers of species, and the change from this
to the ‘mid-estuarine’ fauna inhabiting waters of about half the salinity of sea water is
much sharper than the change from the mid-estuarine to the marine fauna. This accords
with Remane’s observations in the very different environment of the Baltic.

The macro-invertebrate assemblages (especially of the molluscs), as described by Parker
(1959), are of great interest. The bay-head facies, with strong inffuence from rivers and sali-
nity at normal times less than 6%o, has a very sparse fauna with only five macro-inverte-
brate species. In this environment ostracods dominate foraminifera (Ladd etal. 1957).

In enclosed bays of low to variable salinity (12-25%o) the characteristic feature is the
occurrence of ‘reefs’ of Crassostrea virginica, the American estuarine oyster. The only
other common lamellibranch on the reefs is the Mytilid Brachidontes. This association
strongly recalls the Liostrea hebhdica-Modiohis association which occurs both in the
Hebrides and in the Upper Estuarine Series of the Midlands. L. hebridica is thought to
be related to the modern Crassostrea. On the muddy bottoms between the reefs there
are seven macro-invertebrates, including especially the small triangular lamellibranch
Mulinia lateralis. This occurs in all environments, including some hypersaline ones and
the open Gulf coast, but is most common in low salinities. Its behaviour is considered
a good analogy for that of Neomiodon, which it somewhat resembles in morphology.
In times of stable high salinities (> 25%o) these bays are invaded by the fauna normally
characteristic of the more open bays nearer the inlets. This has far more species (34
according to Parker 1959), most of which also inhabit the open Gulf. Lamellibranchs
and gastropods are still dominant. The assemblage at the algal bed horizon of the Lower
Ostrea Beds is thought to be of this kind — a somewhat restricted selection of shallow
water, open-sea forms typical of the Great Oolite limestones.

In his conclusion Parker reaffirms the classical account of the response of animals to
adverse salinities, high or low. In unusual salinities there are few species, but many
individuals. ‘As salinity decreases or increases to normal values (along with relative
stability), the number of species increases and the number of individuals per species
decreases’ (Parker 1959, p. 2158).

SALINITY-CONTROLLED ASSEMBLAGES IN THE
GREAT ESTUARINE SERIES

Rather a striking analogy can be drawn between the well-documented macro-faunal
assemblages of the Texas Bays and those of the Great Estuarine Series. If such a system
of bays and estuaries had existed in the Jurassic — and there is evidence independent of the
fauna to show that it did in the Great Estuarine Series — it is reasonable to suppose that it
would have been inhabited by assemblages of animals adapted to different salinity condi-
tions, as in Texas today. I have argued that the general response of the fauna to salinity

324

PALAEONTOLOGY, VOLUME 6

changes (relative numbers of species and individuals) should be similar, and that the
estuarine genera concerned are likely to have been conservative ones. These are the general
predictions, and direct comparison as outlined above shows that they are fulfilled.

Assuming actual mean salinity values for the habitats of the Jurassic assemblages
having the best analogies with Recent ones, and using ‘overlapping’ of faunas for the
others, I have constructed a ‘salinity series’ or spectrum for the Great Estuarine Series
and Staffin Bay Beds (text-fig. 1). The principal values assumed are: Unio-Viviparus-
Neoniiodon = fresh water or less than 3%o ; Neomiodon as only mollusc = the species
minimum, 5-9%o; Liostrea dominant = Crassostrea reefs of Texas, 12-20%o; marine
assemblage in Lower Ostrea Beds = open bays of Texas, more than 25%o; belemnites
= marine, 35%q. The order which emerges does not involve reversing the salinity
preferences of any of the surviving genera.

Several points must be made in qualification. (1) The order of the assemblages is
obviously more certain than the actual salinity values assigned, which depend upon
several assumptions (e.g. the salinity of the Jurassic ocean). (2) The values assigned are
supposed to represent mean salinities; the salinity doubtless varied widely from time to
time, as happens in Texas, but short-period oscillations have not usually been recorded
in the fossil fauna. (3) The Jurassic assemblages are seen in vertical succession, and it
is not possible as yet to prove lateral passage from one into another. However, except
for the apparent extinction of Mytihis strathairdensis, there is little evidence of evolu-
tionary change in the succession.

The salinity series presented is based entirely on the macrofauna, and especially on
molluscs. Among the microfauna, ostracods are very abundant in what I regard as the
low salinity {Neomiodon-Vivipams-Eiiestheria) assemblages of the Estheria Shales and
Ostracod Limestones. I have not studied these, but Dr. F. W. Anderson informs me
that they are mostly Metacyprids, which would not conflict with the interpretation given.
Foraminifera are apparently absent from most of the Series, but have been seen in thin-
sections and, as chitinous, decalcified shells, in microplankton separations, from the
marine horizon of the Lower Ostrea Beds. A group of samples was examined for spores
and microplankton by Mr. N. F. Hughes and Mr. R. N. Shrivastava. Most of the
samples yielded spores and pollen, but only those from the Garantiana Clay (the bed
immediately below the Great Estuarine Series), the Basal Oil Shale (a transition bed at
the base of the Series), the two algal bed horizons, and the Staffin Bay Beds, yielded
microplankton (dinoffagellates and hystrichospheres). Such microplankton is usually
regarded as marine, and each of the samples yielding it had a marine or brackish-marine
mollusc fauna.

Very few such salinity series have been constructed for fossil assemblages, and very
few indeed from rocks as old as Jurassic. Schmidt (1951) quotes one, with some similari-
ties to mine, prepared for Senonian rocks by Mertin (1939, unpublished thesis). Hilter-
mann (1949) has assigned salinity values to microfossil assemblages; some of these were
criticized by Schmidt (1951). Less precise statements have often been made — marine-
brackish, quasi-marine, &c. — not usually referred to any definition of the terms. Con-
siderable scattered information of this sort exists on the Middle and Upper Purbeck
Beds, which are in many respects similar to the Great Estuarine Series, but no synthesis
has been published. By far the best-known non-marine rocks and faunas are those
of the Coal Measures (Eagar 1960). Large salinity changes can be recognized both

TYPE OF
FAUNA

ASSEMBLAGES IN GREAT ESTUARINE SERIES
AND STAFFIN BAY BEDS.

35-

30

Q

z

<

3

o

X

I—

CL.

<

lb5-

0)

c

-o

<u

u

"Cl

cC

0)

Q- .£

5

3H

05

(Seawater)

Marine,
reduced
number
of species

Brackish -
marine

Typical

Brackish

Water

Brackish -
fresh water

F.W.with
reduced na
of species
Freshwater

(ms. algal bed)

CUSPl DARIA -

PLACUNOPSIS-

MODIOLUS

2

BELEMNITE SANDS (MARINE LAMELLIBRANCHS + BELEMNITES )

(in STAFFIN BAY BEDS)

UPPER OSTREA BEDS (MARINE LAMELUBRANCHS ^UOSTREA &

NEOM/ODON )

NO DIRECT CONNECTION — ■

(l.O-B. ALGAL bed)

UOSTREA-RHYNCHONELLA
MODIOLUS. ETC

Q

MYTILUS

UOSTREA

HEBRIDICA

T A

I

> ^

8
0

UNIO- VIVIPARUS- NEOMIODON- EUESTHERM

TEXT-FIG. 1. Salinity-controlled assemblages in Middle Jurassic faunas. For explanation see text.
Continuous lines show ‘overlapping’ of faunas in the formations indicated.

326

PALAEONTOLOGY, VOLUME 6

palaeontologically and geochemically, with consistent results (Ernst, Michelau, and Tasch
1960), but the effects of minor fluctuations in salinity are difflcult to differentiate from
the large effects of organic content, water turbulence, &c. (Eagar 1960, 1962). It would
be surprising if further work did not extend, and in some respects complicate, the picture
from the Great Estuarine Series, from which very little collecting has been done com-
pared with the Coal Measures. But I am confident that the main thesis — that the faunas
are brackish-water ones controlled mainly by salinity variations — is correct.

Acknowledgements. I wish to thank Dr. G. Larwood for reading the manuscript and suggesting improve-
ments, and Mr. N. F. Hughes and Dr. F. W. Anderson for discussions and information.

REFERENCES

ANDERSON, F. w. 1948. Algal beds in the Great Estuarine Series of Skye. Proc. R. phvs. Soc. Edinb.
23, 123-42.

and cox, L. r. 1948. The ‘Loch Stafhn Beds’ of Skye, with notes on the molluscan fauna of the

Great Estuarine Series. Proc. R. phys. Soc. Edinb. 23, 103-22.
cox, L. R. and ARKELL, w. j. 1948-50. A survey of the Mollusca of the British Great Oolite Scries.
Paloeontograpbical Society, London.

EAGAR, R. M. c. 1960. A Summary of the results of recent work on the palaeoecology of Carboniferous
non-marine lamellibranchs. C.R. 4 Congr. Strat. Carbon., Heerlen, 1, 137-49.

1962. Boron content in relation to organic carbon in certain sediments of the British Coal

Measures. Nature, Land. 196, 428-31.

ERNST, w., MICHELAU, p. and TASCH, K. H. 1960. Vcrglcich des Niveaus von Zollverein I mit dem von
Wyshagen mit Hilfe der Bor-Methode. C.R. 4 Congr. Strat. Carbon., Heerlen, 1, 163-8.

GUNTER, G. 1947. Palaeoecological import of certain relationships of marine animals to salinity.
J. Paleont. 21, 77-79.

HALLAM, A. 1960. A Sedimentary and faunal study of the Blue Lias of Dorset and Glamorgan. Phil.
Trans. Roy. Soc. B. 243, 1^4.

HEDGPETH, J. w. 1957. Estuarics and Lagoons, II, Biological Aspects, in hedgpeth, ed.. Treatise on
Marine Ecology and Paleoecology, Vol. 1 Ecology. Mem. geol. Soc. Amer. 67 (1).

HiLTERMANN, H. 1949. Klassifikation der natui Uchen Brackwasscr. Erdol u. Koble, 1, 4-%.
HUTCHINSON, G. E. 1960. On evolutionary euryhalinity. Amer. J. Sci. 258 A, 98-103.

JUDD, J. w. 1873. The secondary rocks of Scotland. First paper. Quart. J. geol. Soc. Loud. 29, 91-194.

1 878. The secondary rocks of Scotland. Third paper. The strata of the Western Coast and Islands.

Ibid. 34, 660-743.

LADD, H. s., HEDGPETH, J. w. and POST, R. 1957. Environment and facies of existing bays on the
Central Texas Coast, in ladd, h. s. Mem. geol. Soc. Amer. 67 (2).

MILLER, H. 1858. The Cruise of the Betsey . . . Edinburgh.

PARKER, R. H. 1959. Macro-invertebratc assemblages of Central Texas coastal bays and Laguna
Madre. Bull. Amer. Ass. Petrol. Geol. 43, 2100-66.

PEARSE, A. s. and gunter, g. 1957. Salinity, in hedgpeth, j. w. ed.. Treatise on Marine Ecology and
Paleoecology Vol. 1. Mem. geol. Soc. Amer. 61 (1).

SCHMIDT, H. 1951. Erkennbarkeit fossiler Brackwasserabsatze. Z. dtsch. geol. Ges. 9-\6.
segerstrale, s. G. 1957. Baltic Sea, ;« hedgpeth, J.w.ed., Treatise on Marine Ecology and Paleoeco-
logy, Vol. 1. Mem. geol. Soc. Amer. 67 (1).

SHEPARD, F. p. et al. 1960. Recent Sediments, North-west Gulf of Mexico. Tulsa.
sorgenfrei, t. 1958. Molluscan assemblages from the marine Middle Miocene of South Jutland, and
their environments. Damn. geol. Unders. II. Raekke, 79, 1-503.

J. D. HUDSON
Department of Geology,
Sedgwick Museum,

Manuscript received 29 September 1962 Cambridge

THE ECOLOGY AND STRATIGRAPHICAL
DISTRIBUTION OF THE INVERTEBRATE FAUNA
OF THE GREAT ESTUARINE SERIES

by J. D. HUDSON

Abstract. The invertebrate macrofauna of the Great Estuarine Series is reviewed, with special reference to the
ecological inferences which can be drawn from it. Evidence from modem relatives, and from association with
other fossils, is used to deduce the probable ecological preferences, especially of salinity, of each of the common
species. The stratigraphical distribution of the fauna among the formations of the Series is comprehensively
described for the first time. Unio andersoni sp. nov. is described and figured.

This paper records the stratigraphical distribution of the more important faunas of
the Great Estuarine Series (Middle Jurassic) of north-west Scotland, and reviews the
evidence for the ecology of the fossils concerned. The stratigraphy of the Series has been
described elsewhere (Hudson 1962), and is summarized in text-fig. 1.

The fauna is dominated by the Mollusca. As systematic descriptions of most species
have recently been published or are readily available, I have only included morphological
descriptions and taxonomic discussions where my own work has added materially to
existing knowledge. Ecological information, on the other hand, is very scattered in the
literature; part of it comes from modern representatives of fossil groups, and part from
association with other fossils. This information is brought together here, attention being
concentrated on the commonest species or those giving a clear ecological indication,
especially of palaeo-salinity (Hudson, this journal, p. 318).

In the stratigraphical section, all the important invertebrate macro-fossils are listed
(Table 1); this is the first such survey for the whole Great Estuarine Series.

THE INVERTEBRATE FAUNA

The restricted nature of the invertebrate fauna of the Great Estuarine Series as com-
pared with that of the equivalent marine beds of the English Great Oolite Series is
immediately apparent even when allowance is made for deficient collecting.

The common mollusc species of the Great Estuarine Series were mostly described by
Forbes (1851) and Tate (1873). They also identified some fossils with already described
English species. Anderson and Cox (1948) redescribed Forbes’s and Tate’s species, and
assigned them to their correct stratigraphical horizons. They also described some new
species, and reviewed the fauna of the Great Estuarine Series as then defined. In fact
most of this discussion refers to the Staflhn Bay Beds of my classification (Hudson 1962),
the faunas of which are more marine than those of the Great Estuarine Series sensii
stricto, and which are not dealt with in detail here. Some of the lamellibranchs were
further revised by Casey (1952, 1955), and British Great Oolite mollusca generally have
been revised by Cox and Arkell (1948-50). Yen (1948) redescribed some gastropods, but
his conclusions have been disputed. Forsyth (1960) records a few fossils new to the
‘Series’.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 327-48, pi. 53.]

328

PALAEONTOLOGY, VOLUME 6

TABLE 1. List of invertebrate fossils from the Great Estuarine Series

Formation abbreviations from text-fig. 7. * = new record for Great Estuarine Series;

t = single occurrence only.

Formation

Foraminifera: Miliolids, and family indet.*

Echinoderma ; Regular echinoid, plate and spines*
Isocrinuslf
Annelida; " Serpula'

Brachiopoda: Discinisca sp.*'\

' Rliynclionella' cf. conciima (J. Sow.)

Polyzoa: Cyclostome indet. *f

Lamellibranchia:

Modiolus cf. imbricatus J. Sow.

Mytilus (Praemytiliis) strathairdensis Anderson and Cox

Meleagrinella

Pteroperna sp.*

Placimopsis social is Morris and Lycett*

Placunopsis aestiirina (Tate)

Liostrea hebridica (Forbes)

Lopha ip.t

Unio andersoni sp. nov.

Neomiodon brycei (Tate) and N. spp.

Anisocardia {Antiqiiicyprina) cucullata (Tate)

Quenstedtia cf. bathonica (Morris and Lycett)
Quenstedtia? stafiinensis (Forbes)

' Quenstedtia' forbesi Anderson and Cox*

Pleuromya robusta (Tate)

Corbida hebridica Tate

Myopholas acuticostata (J. de C. Sowerby)

Cuspidaria ibbetsoni (Morris)*

Gastropoda:

Neridomus arata (Tate)

Hydrobia praecursor Sandbergerf
Zebina caledonica (Tate)

Tornus praecursor (Tate)

Assiminea skyensis Anderson and Coxf
Procerithium cf. pisoUticum (Hudleston)f
Procerithium {Rhabdocolpus) cf. vetustiim (Phillips)t
Ghbidaria formosal (Morris and Lycett)*t
Globularia hebridica Anderson and Cox
Viviparus scoticus (Tate)

Viviparus bithnyoides (Yen)f
CylindrobuUina inermis (Tate)

Arthropoda:

Phyllopoda: Euestheria murchisoniae (Jones)

Ostracoda: Ostracods including Metacyprids
Bairdia 5p.t

Trace fossils: Pelecypodichnus amygdaloides Seilacher*

L.O.B.

L.O.B.

B.O.S.

L.O.B.

L.O.B.

L.O.B.

L.O.B.

M.S., E.S., L.O.B.
M.S.

B.O.S.

M.S.

M.S., E.S., L.O.B.
M.S., E.S.

E.S., L.O.B.

M.S.

M.S., L.O.B.

M.S., L.O.B. inch
L.O.B.

B.O.S.

O.L.

L.O.B.

L.O.B.

L.O.B.

L. O.B.

M. S., E.S., L.O.B.

L.O.B.

O.L.

L. O.B.

M. S., E.S.

O.L.

O.L.

O.L.

L. O.B.

M. S., E.S.
M.S.-O.L. incl.
O.L.

M.S., E.S., C.S.S.

B.O.S.-O.L. incl.
B.O.S.-O.L. incl.

B. O.S.

C. S.S.

329

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES

My own collection (in the Sedgwick Museum, Cambridge) contains specimens of all
the important species and some additions to the fauna. I have also examined the
collections in the Geological Survey Museum, Edinburgh (through the kindness of
Mr. R. B. Wilson), and have redetermined some of the fossils. I have also been able to
correct the stratigraphical assignment of a few groups of specimens.

<

LU

CC

<

TROTTERNISH

(N.SKYE)

STAFFIN BAY BEDS
(i’U.BATH.-LRCAUL)

RAASAY

LR.CORNBBASH
(top BATHONIAN)

STRATHAIRD

(SSKYE)

CARN MOR SST.
(lr. CALLOVIAN)

EIGG

(u. cretaceous)

MUGK

( tertiary)

yj

cC

UJ

OO

LU

Z
ct
<
3
I —
OO

al

O

MOTTLED

CLAYS

45

OSTRACOD

LIMESTONES

90

LOWER OSTREA
BEDS

65

(Not

exposed )

LOB

MOTTLED

CLAYS

60

OSTRACOD

LST.

80

( Non-

Sequence )

O.L.

20

LR. OSTREA
BEDS

25

LR. OSTREA
BEDS

( Non-

Sequence)

O.L.

30

LR. OSTREA
BEDS

lO

20

CONCRETIONARY
SST. SERIES

250

CONC. SST
SERIES

CONC. SST.
SERIES

CONC. SST
SERIES

120 +

80

190

ESTHERIA

SHALES

120

ESTHERIA SH.
AND

MYTILUS SH.

lOO

ESTHERIA SH
AND

MYTILUS SH.

150

ESTHERIA SH,
AND

MYTILUS SH.

C.S.S.

(" Bose
not

WHITE

SANDSTONE

lOO

WHITE

SST.

50

WHITE

SST.

50

(lR. MYTILUS SK?)

BASAL OIL
SHALE

lO

BASAL OIL
SH.

lO

BASAL OIL
SH

15

lor

50

GARANTIANA ZONE (uR BAJOCIAn}

SST. (7 BAJ.)

TEXT-FIG. 1 . Stratigraphy of Great Estuarine Series. Thicknesses in feet.

THE ECOLOGY OF THE INVERTEBRATE FAUNA

The fauna listed in Table 1 may be readily divided into ecological groups.

(a) The typical fossils of the Great Estuarine Series. Certain species may be con-
sidered with more abundant species of the same genus, so that the ecological inter-
pretation of the ‘Series’ turns very largely on the interpretation of only twelve species
or poups; Modiolus cf. imbricatus, My t Hus (Praemylilus) strathairdensis, Placimopsis
socialis, Liostrea hebridica, Unio andersoni sp. nov., Neomiodon brycei, Cuspidaria

330

PALAEONTOLOGY, VOLUME 6

ibbetsoni; Viviparus scoticus; Tornus praecursor, GlobuJaria hebridica, and Cylindro-
bidlina inermis', Euestheria rnurchisoniae.

These species are discussed in detail below. Taxonomic discussion is included where
necessary.

{b) Six records of marine stenohaline major taxa and eight species of marine mollusca,
confined to the marine horizon of the Lower Ostrea Beds. These are listed and discussed
under Lower Ostrea Beds on p. 340. They were normal shallow-water marine animals.

(c) Three species of marine genera confined to the Basal Oil Shale, a transition bed
at the base of the Series.

{d) The remaining seven species are single records or of very restricted occurrence
(Table 1), so that extended discussion is not possible.

LAMELLIBRANCHS
Modiolus cf. imbricatiis J. Sowerby

Occurrence. A Modiolus comparable to this species, but generally smaller, has been
recorded from several localities and horizons (e.g. Lee 1920, p. 54). I have collected it
from three horizons; from just below the Mytilus Shales algal bed with Placunopsis and
Cuspidaria, from the algal bed horizon of the Lower Ostrea Beds with Rhynchonella
and marine mollusca, and from the Estheria Shales in Raasay alone.

Ecology. Modiolus belongs to a very ancient group, which has always been shallow-
water marine, or even littoral (Newell 1942). Data on modern Modiolus are not so
abundant as on Mytilus, but at least in Britain it apparently lives in somewhat deeper
water, below the littoral zone proper (Yonge 1949), and in the Baltic it does not extend
into water of such low salinity as Mytilus edulis (Sorgenfrei 1958). The genus is thus
marine, shallow-water but not littoral, and able to tolerate limited reduction in salinity.
Its occurrence in the Great Estuarine Series is consistent with this, since it occurs
principally in the more marine beds.

Mytilus (Praemytilus) strathairdensis Anderson and Cox

Preservation. This species (PL 53, fig. 6) was described by Anderson and Cox (1948)
from near the base of the Estheria Shales of Strathaird. Their material consisted of internal
and external moulds in hard black shale. No further occurrences were known, and none
has since been described. The species is, however, abundant on the same horizon in
Eigg, where it characterizes the Mytilus Shales (Hudson 1962). In Eigg the shell is
preserved ; except for a very thin outer layer, which may be calcitic, it is composed of
nacreous aragonite (checked by X-ray powder photograph by Dr. P. Gay). The Myo-
conclia n.sp. of Kitcliin (in Barrow 1908, pp. 21-23) and Lee (1920, p. 54) belongs to
this species (Geological Survey Museum, Edinburgh, T 2963a-T 2968a, from Eigg, and
T 215e, from Raasay).

Occurrence. Except for one doubtful record from the Upper Estheria Shales of north
Eigg, this species is confined to the Mytilus Shales of Eigg, Skye, and Raasay. In these
it is extremely abundant, and except for a few thin beds it dominates the fauna. It
usually occurs in monotypic shell beds, never with Unio or Viviparus, and doubtfully
with Neomiodon.

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 331

Ecology. M. strathairdensis is one of the earliest representatives of its genus, although
subgenerically distinet from Mytilus s.s. At the present day Mytilus is one of the best
examples of a euryhaline marine molluse. In the Baltie area M. edidis inhabits the Gulf
of Finland, one of only four lamellibranehs, out of ninety-two inhabiting the Kattegat,
to do so. The mean salinity at the limit of its range is 6-10%o; there is no doubt that
salinity is the limiting factor (Segestrale 1957, Sorgenfrei 1958). M. edidis is also a com-
mon littoral species in waters of normal salinity. It is extremely gregarious (Newell
1942). There is every reason to suppose that M. strathairdensis, like Midiolus, shared the
general preferences of the Mytilacea, and it almost certainly resembled Mytilus s.s. in
tolerating extreme conditions of salinity and in being gregarious. Its exclusion from the
more certainly marine beds with Placunopsis, Cuspidaria, and Modiolus supports this.
On the other hand, Mytilus has always been a marine-euryhaline, rather than a fresh
or true brackish-water genus, and this is supported by the absence of M. strathairdensis
from beds with Unio and Viviparus. The species does not overlap in stratigraphical range
with Liostrea hebridica, which may have had similar ecological requirements.

Placunopsis socialis Morris and Lycett
Placunopsis aesturina Tate

Previous records. Placunopsis socialis (PL 53, figs. 3, 4) is a well-known species in the
Great Oolite, especially in the Upper Estuarine Series of the Midlands. It has not been
recorded from the Hebrides before. P. aesturina was described by Tate (1873, p. 349,
pi. 12, fig. 12) from the Estheria Shales, and has been recorded from various horizons
and localities in the Great Estuarine Series.

Tate’s description of P. aesturina mentions central umbones, whereas his figure shows
an inequilateral shell. The holotype is in fact inequilateral, like the figure (Geological
Survey Museum, London, 8621). Most of the records from the Great Estuarine Series
are of specimens with central umbones, better referred to P. socialis, or are not specifi-
cally determinable. P. socialis is a very variable species. An inflated form, very common
in the Mytilus Shales, is rather distinct, but also occurs in the Upper Estuarine Series
of the Midlands.

Occurrence. Placunopsis occurs at various horizons in the Mytilus and Estheria Shales,
and in the Lower Ostrea Beds. In the limestones just below the Mytilus Shales algal
bed P. socialis is almost a rock-former, especially the inflated form. It is associated with
Cuspidaria ibbetsoni. Modiolus, Lopha, and rare Neomiodon.

Ecology. Placunopsis is undoubtedly a marine genus. It is gregarious and common in
restricted assemblages, and may have been tolerant of very shallow water and variable
salinity.

Liostrea hebridica (Forbes)

Discussion. This is one of the most important fossils (text-fig. 2) of the Great Estuarine
Series, where it dominates the fauna of the Lower Ostrea Beds. A discussion of the
species, with synonymy and excellent figures, is given by Arkell (1934). In the Great
Oolite Series it was long known as Ostrea sowerbyi. The variety rugosa is not known
in the Hebrides. Most specimens are moderately elongated, but the extreme variety

C 1213

z

332 PALAEONTOLOGY, VOLUME 6

eJongata, which is the commonest form in the Fuller’s Earth at Langton Herring in
Dorset, is uncommon.

The genus Liostrea Douville may conveniently be used for smooth and not strongly
curved Mesozoic oysters, but it is probably only a form genus, as are other ‘genera’ of
oysters based on shell form and sculpture. As is becoming well known, living oysters
can be distributed among three genera — Pycnodonte, deep-water and non-gregarious ;
Osirea s.s., with type species O. edulis; and Crassosfrea, type species the American
oyster, C. virginica (Yonge 1960). The biological differences are considerable, and are
correlated with ecological station : Ostrea is smaller, rounded, larviparous, and marine,
while Crassosfrea is larger, elongated, oviparous, and estuarine. It possesses a structure,

TEXT-FIG. 2. Interiors of right valves of Liostrea hebridica (A, b) and Crassostrea virginica (c), to show
similarity of shell form and position of muscle-scar, a, Langton Herring (tracing of Arkell 1934, pi. iv,
iig.6;L.hebridica\ar.elongata,x 1. B,Ketton, Upper Estuarine Series, original, X 1. c. After Orton, X

the promyal chamber, for removing suspended mud from its system. Correlated with
this is the position of the muscle-scar well back from the hinge-line, as opposed to its
central position in Ostrea. Ranson (1939, 1942) showed that the larval shells, or prodis-
soconchs, afforded a clear distinction between the oyster genera, and demonstrated by
this criterion that some fossil genera could be assigned to either Osirea or Crassostrea
(Gryphaea of Ranson).

In general shape and position of muscle-scar L. hebridica closely resembles Crassostrea
virginica, though it is only about half as large (text-fig. 2). The muscle-scar is regarded
as significant in fossils by Gunter (1950) and Yonge (1960). Hallam (1959) relates
L. irregularis to Ostrea by this criterion. Preliminary work has been done on separating
prodissoconchs from the matrix of L. hebridica shell-beds, especially that of Langton
Herring in the Fuller’s Earth of Dorset. A few have been found which are strongly
inequilateral, like those of Crassostrea and unlike Ostrea (Yonge 1960). It is hoped to
continue this work, but at present both lines of evidence suggest that L. hebridica was
more closely related to Crassostrea than to Ostrea.

Occurrence. The chief occurrence of L. hebridica is in monotypic shell-beds in the Lower
Ostrea Beds. Less often, the shell beds also contain Neomiodon, and this lamellibranch
is perhaps more common in the associated shales. At the marine Algal Bed horizon
L. hebridica occurs with a much greater variety of lamellibranchs (listed in next section)

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 333

and with Rhynchonella. Very similar assemblages occur in the Upper Estuarine Series
of the Midlands. In the Staffin Bay Beds L. hebridica is common, but less completely
dominant.

Ecology. Summaries of the vast literature on oyster ecology are given by Korringa
(1952) and Yonge (1960). Crassosirea, to which L. hebridica is thought to have been
related, has a narrower temperature range than Ostrea, but is much more tolerant of
low and fluctuating salinities; Ostrea extends to salinities of 23%o, Crassosirea to 12%o
(Yonge 1960). Crassosirea in fact does not normally form reefs except in reduced
salinities. In the enclosed bays of Texas the pure C. virginica reefs which flourish in
salinities of 12-20%o are invaded at higher salinities by marine forms, including Ostrea.

The morphological and ecological analogies between L. hebridica and C. virginica,
further discussed in the accompanying paper (p. 323), are so close that it seems reason-
able to infer that L. hebridica, in the form of pure oyster reefs, inhabited shallow,
partially enclosed bays with salinity about half that of sea water.

Unio andersoni sp. nov.

This species (PI. 53, figs. 1, 2) is described at the end of the paper.

Previous records. There has been some doubt about the occurrence of Unio in the Great
Estuarine Series; as Anderson and Cox (1948) show, most of the early records were
erroneous. Unio was, however, correctly identified from Eigg by Kitchin (in Barrow
1908), but was not described. Jackson (1911a, p. 120) records Unio, compared to
U. disiortus Bean, from Eigg. I have examined the specimen (Manchester Museum,
L. 10504), and do not accept the pseudocardinal tooth mentioned by Jackson. There is
no proof that the shell is a Unio.

Occurrence. Reptile Bed and Unio Bed, Mytilus Shales, Eigg; Lower Ostrea Beds(?),
Trotternish.

Ecology. Unio is perhaps the best known of all freshwater lamellibranchs, and the
Unionidae have been exclusively freshwater at least as far back as the Jurassic. The
typical occurrence of present-day Unionidae is in freshwater lakes and rivers. However,
U. andersoni certainly did not live in inland waters, since both the Reptile Bed and the
Unio Bed are followed without a break by beds with marine-brackish fossils. It is
therefore interesting to note that Unio occurs at the present day in the shallow coastal
waters of the Gulf of Einland, with salinities up to a maximum of 3-4%o (Segerstrale
1957). Therefore, if modern Unio is taken as a guide, U. andersoni could have inhabited
coastal lagoons like those envisaged for the deposition of the Great Estuarine Series,
but these must have become temporarily almost freshwater.

The Reptile Bed occurrence, with gastropods usually regarded as marine (see under
Tornus praecursor, below), may be an ecologically mixed one in a condensed deposit,
and in any case I prefer to regard the gastropods, rather than Unio, as euryhaline.

Neoniiodon brycei (Tate) and Neomiodon spp.

Discussion. The genus Neomiodon, the Cyrena of earlier authors, may be considered as a
whole. It is the commonest mollusc of the Series; it dominates the faunas of the Estheria
Shales and Concretionary Sandstone Series, where it forms monotypic shell-beds and is

334

PALAEONTOLOGY, VOLUME 6

a rock-former in the limestones. It also occurs in the Mytilus Shales and the Lower
Ostrea Beds. In the Staffin Bay Beds it forms part of a more varied and more marine
fauna.

The taxonomic history of the Great Estuarine Series Neomiodotis is complicated.
Forbes and Tate described several species of ‘ Cyrena \ mostly from the Staffin Bay Beds.
These were redescribed by Anderson and Cox (1948), who referred some to marine
genera, and erected a new genus Protomiodon, for Cyrena brycei Tate and species con-
sidered congeneric with it. Casey (1955), in a comprehensive paper on the Neomio-
dontidae, showed that Protomiodon was a synonym of Neomiodon. Casey (1952) has also
demonstrated extensive external homeomorphy among such smooth, trigonal-ovate
lamellibranchs as Neomiodon and Staffinella [Protomiodon] staffinensis. Fortunately this
problem concerns mainly the Staffin Bay Beds; the forms which dominate the fauna of
the middle part of the Great Estuarine Series are undoubtedly Neomiodon. Most can
be referred to N. brycei (Tate) and its variety quadrata on shell outline (Anderson and
Cox 1948), but other species, including N. cwminghamii (Forbes), are probably present.

Occurrence and ecology. Since the Neomiodontidae are extinct, it is not possible to draw
ecological conclusions from close modern relatives, and arguments must be based on
associations with other fossils. Casey (1955) considers the family to be the ecological
forerunners, but not the direct ancestors, of the fresh- and brackish-water Corbiculidae,
of the Tertiary and the present day, which they so closely resemble. He sums up the
ecology of Neomiodon itself as ‘fresh or brackish water, gregarious, sometimes associated
with members of marine genera’. He cites evidence from the Purbeck and Wealden of
frequent association with Unio and Viviparus, and much less frequent occurrence with
Modiolus, Corbula, and Pecten. Despite association with marine genera in the Staffin
Bay Beds, Casey considers Middle Jurassic Neomiodon to have had the same ecological
preferences. It is now possible to cite evidence, similar to Casey’s from the Purbeck,
from the Great Estuarine Series itself. Neomiodon occurs with Viviparus, Euestheria, and
metacyprid ostracods at many horizons, and with Unio at two, as well as with Liostrea.
As in the Purbeck, however, it is not usually abundant in precisely the same bed as
marine-brackish genera. The analogy with the Middle Purbeck is the more striking as
the facies is so similar. I therefore support Casey’s conclusions on the ecology of
Neomiodon. The extensive beds with Neomiodon alone may have been deposited in water
just saline enough to exclude Unio and Viviparus, but still too dilute to admit marine
genera; this is the ‘species minimum’ of the Great Estuarine Series.

The trace fossil Pelecypodicimus amygdaloides Seilacher is extremely common in the
thin-bedded sandstones and shales of the Concretionary Sandstone Series. It is formed
as the ‘resting trace’ of a lamellibranch inhabiting shallow, temporary burrows in a thin
layer of sand over mud (Seilacher 1953). In this case Neomiodon was undoubtedly the
lamellibranch responsible, and the occurrence confirms the mode of life of Neomiodon
which could also be predicted from its typically ‘active lamellibranch’ morphology (cf.
Yonge 1949, pp. 228-9).

Cuspidaria ibbetsoni (Morris)

This species (PI. 53, fig. 5) has not previously been positively identified from the
Series, though Anderson and Cox (1948, p. 120) record a probable fragment.

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 335

Occurrence. C. ibbetsoni is abundant at several horizons in the Mytilus and Estheria
Shales, especially just below the algal bed, and also occurs in the Lower Ostrea Beds.
It is associated especially with PJacimopsis socialis, an association also characteristic of
the Upper Estuarine Series of the Midlands,

Ecology. Modern Cuspidaria is deep-water marine, and Cox (1960) has questioned
whether the Jurassic species, which are all shallow-water forms, were related to it, or
were septibranchs at all. C. ibbetsoni occurs widely in the Great Oolite Series in England,
and is especially abundant in the Upper Estuarine Series of the Midlands. The forms
in the clays there have long rostra, like my specimens and unlike those from the Great
Oolite limestones (Cox and Arkell 1948) — a fact which may be ecologically significant.
The occurrence and associations of C. ibbetsoni suggest a shallow marine habitat, and
tolerance of muddy and perhaps brackish conditions.

GASTROPODS

Viviparus scoticus (Tate)

Previous records. This is by far the most important gastropod in the Great Estuarine
Series, occurring in all formations from Mytilus Shales to Ostracod Limestones, and it
has given rise to much controversy. It was described as a Paludina (= Viviparus) by
Tate (1873), and considered as freshwater until Yen (1948) proposed a new genus
BathoneUa for it, and claimed that it was marine. The ecological and taxonomic argu-
ments are linked, since many palaeontologists are unwilling to admit that a marine shell
was a Viviparus, or vice versa. The purely morphological differences between BathoneUa
and Viviparus have been very differently assessed by different authors. Yen (1948) placed
it in a different family from Viviparus', this was accepted by Anderson and Cox (1948).
Arkell (1948) doubted the generic separation, and Watson (1950) firmly rejected it. Cox
(1950) admitted that the distinction could be specific only, but held that since BathoneUa
was marine the generic distinction was desirable. Since I do not consider that the ecology
of BathoneUa differed materially from that of other Viviparus, I propose to reject the
name, regarding the slight morphological differences listed by Yen as of specific rank
only (Watson 1950). BathoneUa bithnyoides Yen 1948 is assumed to be congeneric with
V. scoticus.

Occurrence. Yen (1948), who was originally responsible for the view that V. scoticus was
marine, did not in fact record marine fossils from the same bed, but only from an
adjacent one (see Stratigraphical section). In view of the repeated oscillations of condi-
tions known in the Great Estuarine Series this cannot count as a true association. Cox
(1950) drew up a list of occurrences of V. scoticus in the Great Estuarine Series, mainly
from Anderson’s unpublished records, but including two of Tate’s (1873) lists, which
alone contain undoubtedly marine genera. Cox admits that these may have been com-
piled from more than one bed, and from field experience of the localities concerned I
should regard this as certain. This reduces Cox’s list of associates to Neomiodon,
Euestheria and ostracods, all at many localities, and Hydrobia praecursor and Quen-
stedtia? staffinensis at one each. The only marine genus here is Questedtia, and the
generic attribution of the species is quite uncertain (Anderson and Cox 1948). To this
list I can add, from my own collecting, Unio in two localities, both with Neomiodon also.

I can also add several more occurrences of V. scoticus alone or with Neomiodon,

336

PALAEONTOLOGY, VOLUME 6

Euesiheria, and ostracods, and a great deal of negative evidence, on the non-occurrence
of V. scotkus with marine or marine-brackish genera.

The other well-known occurrences of V. scotkus (or species regarded by Anderson
and Cox (1948) as synonymous with it) are in Oxfordshire (Watson 1950) and Indre,
France (Fischer 1961). Neither yields acceptable evidence of marine associates. Fischer
shows clearly that Viviparus occurs immediately above a lignite bed which rests on an
erosion surface, that it is associated with "Valvatk and 'Planorbis', and that none of
these gastropods is found in the marine beds above and below.

Ecology. Present-day Viviparus is one of the best known of all freshwater gastropods.
There is therefore a presumption that beds containing shells which appear to be fossil
Viviparus are also freshwater, especially if they contain other freshwater fossils. This
conclusion has always been accepted, for instance, in the Wealden Beds. On the basis
of the above review, there seems no reason to depart from it in the case of the Great
Estuarine Series. There only remains, as with Unio, the possibility that Viviparus,
since its habitat in this case was certainly coastal, could tolerate a slight degree of
brackishness in the water. This is quite possible; but the total exclusion of marine genera
shows that salinity stayed on the freshwater side of the ‘species minimum’.

Tornus praecursor (Tate)

Globularia hebridica Anderson and Cox
CylindrobuUina inermis (Tate)

These three small gastropods may be considered together. The type localities are all in
the Estheria Shales of the Portree district, Skye, where the species occur together and
with Modiolus (Tate 1873, Anderson and Cox 1948). Tate referred the species to
Valvata, Neritina (G. hebridica = N. staffinensis Tate non Forbes), and Melania respec-
tively. In referring them to marine genera, Anderson and Cox comment on the difficulty
of generic assignment of imperfectly preserved small gastropods, and in the case of
T. praecursor they were influenced by association with other marine fossils.

Occurrence. Most small gastropods in my collection do not show apertural characters,
and are therefore difficult to determine. However, all three species mentioned above
probably occur in the Reptile Bed of the Mytilus Shales. G. hebridica is the abundant
small naticoid shell of Kitchin (in Barrow 1908, p. 22). Here they are associated with
Unio, as well as with numerous flsh scales and teeth and disaggregated reptilian bones,
so the assemblage is an ecologically mixed one if the gastropods were truly marine.
I have also found CylindrobuUina sp. with young Mytilus strathairdensis and with
Neomiodon brycei. Small, low-spired gastropods, possibly referable to T. praecursor,
occur associated with Neomiodon and Euestheria at several localities in the Estheria
Shales of Skye.

It is possible that these small gastropods were euryhaline, as many small gastropods
are today, but until more taxonomic work has been done on better-preserved material
they are unsafe environmental indicators.

PHYLLOPODA

Euestheria murchisoniae (Jones)

Previous records. In 1862 Jones described this species of ''Estheria^ from material

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 337

collected by Murchison in 1826. His type material appears to be lost. Subsequently
the species, or the genus ' Estheria', has been recorded several times from the Great
Estuarine Series, but no further descriptions or figures have been published. The classi-
fication of fossil ^ Estheria' (itself a preoccupied name) is in a confused state; the genus
accepted here for E. murchisoniae is Euestheria Deperet and Mazeran, of which the type
species is Eu. minuta of the Triassic (Kobayashi 1954).

Occurrence. Euestheria is most abundant in the Estheria Shales, where bedding planes
may be covered by its carapaces. It is associated with Neomiodon, ostracods, fish
fragments, and oogonia of Chara. In the Ostracod Limestones it is again very common,
but less universally present, accompanied by ostracods and fish fragments. It occurs
more locally in the Basal Oil Shale, the Mytilus Shales, and the Concretionary Sandstone
Series.

I have collected twelve samples with abundant Eucv/Z/er/a from the Estheria Shales and
the Ostracod Limestones, and studied the variation within and between them (210
specimens were examined). Carapace outline, position of umbo, number of growth-rings,
and interspace ornament are among rather few features of fossil" Estheria ’which can be
used in classification, but there has been no agreement as to which of them is most
important (compare Raymond 1946, Bock 1953, Kobayashi 1954, Tasch 1956). My
study showed that each sample was reasonably homogeneous within itself, but that
there was considerable variation between samples, especially in interspace ornament
and also in size. This suggests that several taxa are present — these range from varieties
or subspecies to genera or even families, depending on the classification employed. In
general the specimens were smaller and less elongated than those figured by Jones, but
their general outline and number of growth-rings were similar. At present I propose to
retain all the specimens in Eu. murchisoniae. The variations did not have any obvious
stratigraphical significance, but the largest specimens were from the Ostracod Lime-
stones.

Ecology. Modern "Estheria" species are most common in very shallow ponds, including
temporary ‘playas’, in inland basins in warm temperate areas. They mostly inhabit
fresh waters, occasionally brackish lagoons, but never live in the sea (Kobayashi 1954,
Jones 1862, pp. 5-7). The vast majority of fossil " Estherias' come from continental
formations. It is certainly significant that they are completely absent from the marine
Jurassic and Cretaceous of north-west Europe, but occur in the Triassic (including the
Rhaetie), the Great Estuarine Series, the Purbeck, and the Wealden. In the Great
Estuarine Series the associates listed above are all freshwater or freshwater-brackish.

THE STRATIGRAPHICAL DISTRIBUTION OF THE FAUNA

The faunas of the formations of the Great Estuarine Series are here described in
stratigraphical order. All macro-invertebrate species are listed (apart from a few early
records which have not been confirmed) ; vertebrates and microfossils are only mentioned
in passing. Measured sections are given of a few important localities; much fuller
stratigraphical information, including many more measured sections, is in my thesis
(University of Cambridge, Ph.D. 1962). Figures in brackets after locality names refer
to the National Grid.

338

PALAEONTOLOGY, VOLUME 6

a. Basal Oil Shale

The Garantiana Clay, the last marine formation below the Great Estuarine Series,
passes gradually up into the Basal Oil Shale. The lithology changes from blue, blocky
clay to black fissile shale, and the fauna changes from one of ammonites to a peculiar
and restricted assemblage in which fish scales are the commonest fossils.

The gradual passage is best seen in the cliff's above Prince Charles’s Cave, north of
Portree, Skye (513477 to 515490). The Oil Shale here contains ? Quenstedlia cf. bathonica
(Morris and Lycett), with valves attached (cf. Forsyth 1960), Meleagrinella sp. indet.,
indeterminate tiny gastropods and ostracods, in addition to numerous cycloid and
ctenoid fish scales and one almost complete leptolepid fish (identified by Mr. H. A.
Toombs; Sedgwick Museum J.29055). In Raasay the Quenstedtia Shale of Forsyth
(1960), the equivalent of the Basal Oil Shale, contains Isocrinus in its lower part and
Quenstedtia alone in its upper part; just below the White Sandstone Eiiestheria occurs.
At Port na Cullaidh, Elgol, Strathaird, tlie Basal Oil Shale is intermittently exposed;
it is very similar to the Trotternish development and yields abundant fish scales,
Eiiestheria, ostracods, gastropods, and ? Mytilus. The abundance of fish material con-
trasted with the paucity of mollusca, together with the high carbon content, suggest
that the bottom water may have become stagnant at this time. There was probably also
a gradual decline in salinity.

b. The White Sandstone is unfossiliferous.

c. Mytilus Shales

These form the lower part of the Estheria Shales of Eigg, Strathaird, and Raasay
(Hudson 1962). The type section in Eigg, north of Kildonnan (495870), gives the best
record of rapid vertical variations in lithology and fauna in the Great Estuarine Series.
A detailed account will be published elsewhere; the following is a summary of the
faunas, from top to bottom.

9. Algal Bed, 1 ft. 4 in. An ‘algal-dome’ bed partly composed of faecal pellets (Hudson
1962). Placunopsis socialis near the base.

8. Placunopsis Limestones, 6 ft. Hard blue limestones separated by shales. P. socialis,
P. aesturina?, Cuspidaria ibbetsoni. Modiolus sp., Lopha sp., Neomiodon sp. (rare).
Following the interpretations of Section III, this is a high-salinity fauna.

7. Unio Bed, 8 in. Sandy argillaceous limestone. Unio andersoni, common; Neomio-
don, Viviparus. A freshwater fauna.

6. Limestones and shales, 5 ft. Cuspidaria ibbetsoni near base, Neomiodon brycei near
top. First occurrence of these two species. Last occurrence of Mytilus strathairdensis.

5. Silty shale with thin limestones, 30 ft. The main mass of the Mytilus shales; fauna
dominated by M. strathairdensis, forming ‘mussel beds’.

4. Complex Bed, 3 ft. Coarse sandstone with common fish remains. First occurrence
of fully grown M. strathairdensis.

3. Shale, 8 ft. Very abundant M. strathairdensis, but nearly all young individuals.
Near the base a 1-in. bed with Viviparus but no Mytilus.

2. Reptile Bed, 4 in. Hard sideritic limestone. Abundant small gastropods
{Globularia hebridica, Torniis praecursor, Cuspidaria ibbetsoni), common Unio andersoni.

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES

339

Abundant fish teeth, scales and fin-spines, reptile vertebrae, and limb bones. Possibly a
condensed deposit.

1. Shale, 8 ft. Sparsely fossiliferous. Near base, Euestheria murchisoniae (therefore
low salinity) and fish scales like those from the Basal Oil Shale. Base not seen.

These rapid changes of fauna within a fairly constant gross lithology are regarded as
characteristic of lagoonal conditions, and due to salinity changes (see accompanying
paper). The above succession records a gradual (and fluctuating) increase in salinity from
bed 1 to bed 5, to perhaps half the salinity of sea water; a decline in beds 6-7 to almost
fresh water, and a more sudden increase in beds 8-9 to definite marine-brackish salinities,
as judged by the ecology of the fossils deduced in the previous section of this paper.

In Strathaird M. strathairdemis is common, and the Placunopsis limestone below the
algal bed is well developed. It is exposed north of Elgol (Anderson 1948, Hudson 1962),
where it yields Placunopsis socialis, Cuspidaria ibbetsoni, Pteroperna sp.

In Raasay the Mytilus Shales are not well known, but M. strathairdensis occurs and loose
blocks of the Algal Bed and of the Placunopsis limestones have been found (Lee 1920).

d. Estheria Shales

The typical Estheria Shales have an abundant and highly characteristic fauna of
Euestheria, Neomiodon, ostracods, and scattered fish fragments; in certain beds these
are accompanied by Viviparus and small gastropods.

At the Leah River in Trotternish, the type locality (Anderson 1948, pp. 125-6, with
map), the typical lithology is well seen. The black shale immediately overlying Anderson’s
algal limestone, on the south bank of the river just above the road-bridge, yields
Euestheria murchisoniae, a form with radial-linear interspace ornament (in great abun-
dance and well preserved), Neomiodon, ostracods, and fish scales. Similar shale a few feet
higher in the succession, on the north bank, yields Eu. murchisoniae, form with punctate
ornament, Neomiodon, Viviparus, and very abundant ostracods.

The burn north of Rigg Burn (crosses road at 518571) gives a good section, which
affords an excellent example of the characteristic rapid variation of fauna. The following
section is about 100 yards west of the road, and 30 ft. above the base of the Estheria
Shales:

ft. in.

3. Shale with Cuspidaria ibbetsoni, Placunopsis socialis, cf. Quenstedtia,

Procerithium spp. 1 0

2. Shell limestone, Cuspidaria, with shale partings 1 0

1. Shale wiih Euestheria murchisoniae, ostracods, fish fragments, ? Mytilus 2 0

A similar shale to bed 1, about 10 ft. lower in the succession, yielded Eu. murchisoniae,
ostracods, Neomiodon sp., Viviparus scoticus, ? Tornus praecursor. This is a complete
low-salinity assemblage; it is notable that bed 1 above contains Euestheria and Mytilid
shells, but not Viviparus, and may thus represent somewhat higher salinity. Bed 3 is
definitely brackish-marine, with no freshwater-brackish indicators.

In Raasay the shales are identical to those of Trotternish, with abundant Euestheria.
Oolitic limestones are well represented ; one of these yielded Modiolus cf. imbricatus.

In Strathaird the beds are baked by intrusions, and less amenable to search for fossils.
The most notable feature is the occurrence of Liostrea hebridica just above the algal

340

PALAEONTOLOGY, VOLUME 6

bed ( Anderson 1948, p. 127). Higher up, some Viviparus beds occur. In Eigg the shales
yield Euestheria, Neomiodon, ostracods, and Viviparus, with no additions to the Trot-
ternish fauna. Cuspidaria ibbetsoni and Placiuwpsis sp. occur a short way above the algal
bed on the north coast (474907), and I have a queried record of Mytilus from here.
Immediately above is a Viviparus bed.

€. Concretionary Sandstone Series

This is the thickest formation in the Great Estuarine Series, and it is very variable.
Its deposition is an involved study in sedimentary petrology and sedimentation. Its
fauna, however, is monotonous. Neomiodon is the only macrofossil in many beds;
Viviparus is the only other common mollusc. A few beds contain ostracods; Euestheria
is rare. There are isolated records of Placunopsis (from Strathaird, T 4311, T 4318;
Wedd 1910, p. 124), and CylindrobuUina? (from Eigg, T 719a, T 720a, two forms ; Kitchin
in Barrow 1908, p. 24); these are in the Survey Museum, Edinburgh.

My collection includes well-preserved Neomiodon brycei and Neomiodon sp. from the
Lealt River, Trotternish (with brown on white colour banding), from the Valtos shore
section, Trotternish, from Eist, Duirinish (associated with small CylindrobuUina sp.),
and from Raasay. The best-preserved Viviparus scoticus are from Eist and from Strath-
aird, where they occur near the top of the formation on the shore at Cladach a’ Ghlinne
(Wedd 1910, p. 124). The trace fossil Pelecypodicimus amygdaloides is abundant,
especially in Trotternish.

f. Lower Ostrea Beds

These beds, with their extraordinary abundance of oysters, have attracted attention
and comment from the earliest investigators onwards. Everywhere limestones composed
of Liostrea hebridica, alternating with shales, are the dominant lithology. The beds are
exposed over a wider area than any other formation of the Series, and are fairly constant,
though thicker and sandier in Trotternish than elsewhere (Hudson 1962). Exposures
yielding L. hebridica are far too numerous to list here. This account concentrates on
the more extensive fauna of the marine horizon which is constantly associated with
Anderson’s (1948) Algal Bed. The important exposures are described below. Since
knowledge of the distribution of this fauna at present depends more on the vagaries of
collecting than on true geographical variation, it is convenient first to review the fauna
as a whole.

The greatest thickness of the marine beds is not more than 10 ft., in Trotternish and
Waternish, and these beds are always underlain and overlain by ordinary Liostrea beds.
The Algal Bed usually occurs towards the top of the marine beds.

The following members of marine stenohaline groups are confined to this horizon:
foraminifera (several records, mostly of miliolids), echinoids (regular, plates and spines,
2 records), (1 record), Discinisca{\ record), RhynchoneUa cf. concinna (common

in north Skye), polyzoan (cyclostome, 1 record).

Of the mollusca, Liostrea hebridica, Modiolus imbricatus, Cuspidaria ibbetsoni, and
Placunopsis socialis were dealt with above, and the following are confined to this
horizon: Anisocardia (Antiquicyprina) ciicullata (Tate), ' Quenstedtia' forbesi Anderson
and Cox, Pleuromya robusta (Tate), Corbula hebridica Tate, Myopholas acuticostata
(J. de C. Sowerby); Neridomus arata (Tate), Zebina caledonica (Tate), Globularia

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 341

fonnosa? (Morris and Lycett). These are discussed below. In addition there are frag-
ments of Mytilus (Falcimytilus) sp. in my collection (see also Tate 1873, p. 346), and
indeterminate lamellibranchs and gastropods. It is doubtful if Neomiodon occurs. The
smooth, trigonal ovate lamellibranchs {"Cyrenas' of the old authors) in these beds are
usually larger than typical Neomiodon, and in the absence of preserved dentition are
very difficult to determine. Most are probably members of marine genera, like Aniso-
cardia cucullata, which was itself one of Tate’s 'Cyrenas' (cf. Cox 1947, Anderson and
Cox 1948, Casey 1952, 1955).

Anisocardia (Antiquicyprima) cucullata (Tate). The specimens recorded here do not show
dentition, but may be referred to this species on shell outline. Some are not distinguish-
able from small specimens of A. (A.) loweana (Morris and Lycett) from the Great Oolite.

'Quenstedtia' forbesi Anderson and Cox. One specimen from Duntulm agrees with
Anderson and Cox’s figure (1948, pi. 2, fig. 9). The dentition is unknown and the shell
may not be a Quenstedtia (Casey, personal communication).

Pleuromya robusta (Tate). Several specimens from Lovaig Bay, of which the best is
figured (PI. 53, fig. 8), appear to confirm Anderson and Cox’s tentative conclusion that
this species is distinct from P. uniformis(i. Sow.), being shorter and relatively higher.

Corbula hebridica Tate. This small species is quite common at this horizon.

Myopbolas acuticostata (J. de C. Sowerby). This distinctive lamellibranch (PI. 53,
fig. 7) was recorded by Tate (1873, p. 346), but not mentioned by Anderson and Cox.
It is quite common at Duntulm and at Loch Bay. It is widespread in the Great Oolite
Series, including the Upper Estuarine Series of Northants.

Neridomus arata (Tate) and Zebina caledonica (Tate). Two of Tate’s species, revised by
Anderson and Cox (1948). I have no further records.

Globidaria fonnosa? (Morris and Lycett). Two specimens from Loch Bay are doubtfully
referable to this common Great Oolite species.

These are all normal, shallow-water, marine fossils, either identical with or closely
related to species of the English Great Oolite Series. The marine incursion was short-
lived, however, and it seems likely that the lagoons reached almost marine salinity for
a while, rather than that truly open-sea conditions were established. A closely analogous
case is the invasion of the Texas Crassostrea reefs by a more saline fauna, of species
which also inhabit the open Gulf, when salinity rises above 25%o (see Hudson, this
journal, p. 318).

Trotternish succession

The best exposure of the marine horizon is the shore section at Cairidh Ghlumaig,
Duntulm (410737), where Macgregor first discovered Rhynchonella in the Great Estu-
arine Series (Macgregor 1934, Anderson 1948). The section shows, in summary:

ft. in.

3. Shale and limestone, alternating, with Liostrca 10 0
2. Shale, sandstone, and limestone with Liostrca

and Rhynchonella, algal bed 3 in. below top 7 7

1 . Sandstone with plant remains and Liostrca 11 0

342

PALAEONTOLOGY, VOLUME 6

The marine beds, as defined by the range of RhynchoneUa {R. ef. concmna, Macgregor
1934), are no. 2 of the above suceession. A bed of sandy shale, 2 ft. above the base of
bed 2, yielded, in addition to L. hebridica, Myopholas acuticostala, Anisocardia cucidlata,
Quenstedtia forbesi, and indeterminate heterodonts. From bed 2 generally eome
Discinisca sp., Serpida sp. and, according to Macgregor (1934), Modiolus sp.

A stream section | mile SSW. of the above locality (406728), above the road-bridge,
is in beds probably entirely above those just described. It shows 7 ft. of green silty shales,
with Neomiodon, overlain by 5 ft. of Liostrea beds, which also contain PJacunopsis. This
section is important as showing Neomiodon shales within the Lower Ostrea Beds. Loose
blocks of sideritic mudstone, apparently nodules, yielded a well-preserved fauna of
Neomiodon bryeei and N. sp., Unio andersoni, and Viviparus scoticus. All three genera
occur in one hand specimen. This is one of the best freshwater faunas from the Great
Estuarine Series. The loose blocks almost certainly came from the green Neomiodon
shales within the Lower Ostrea Beds, and thus imply a very marked oscillation in
salinity. It is notable that L. hebridica does not occur with the freshwater fossils. It is
just possible, but unlikely, that the blocks came from the Ostracod Limestones higher
up the stream.

Waternish succession

The small inlier at Loch Bay gives excellent exposures of the marine horizon. There
are three sections showing algal beds in the Bay River and its tributaries. They differ in
details but agree in showing two algal beds, separated by 1 to 2\ ft. of shale and oyster
limestone, and probably are all on the same horizon. Anderson (1948, pp. 133-4) gives
a map of the exposures, but only records one algal bed.

The following section is seen in the tributary to the Bay River at grid reference 271537 ;
beds 1-3 in the right bank at a bend in the stream, beds 4-7 in the stream itself a little

farther downstream: .

ft. in

1. Shale and limestone with Liostrea 3 9

6. Algal bed, brecciated 0 9

5. Shale with Liostrea 1 0

4. Algal bed, laminated, with RhynchoneUa and foraminifera 0 9

3. Shale and limestone with well-preserved Liostrea 6 3

2. Sandstone, grey, fine-grained, uncemented 3 6

1. Clay, grey, sandy, base not seen 3 0

The top of bed 2 is somewhat hardened and yields abundant but fragile Placunopsis
socialis, Cuspidaria ibbetsoni, and numerous ostracods.

Near the mouth of the Bay River a richly fossiliferous white limestone outcrops,
overlain by the algal beds and Liostrea beds. It probably yielded many of the fossils
listed by Tate (1873, pp. 346-7) from Loch Bay. My collection includes L. hebridica,
Myopholas acuticostata. Modiolus c^. imbricatus, Anisocardia cucullata, Corbula hebridica,
heterodont lamellibranchs, Globularia formosa? The top of this bed yielded the only
polyzoan from the Great Estuarine Series, a cyclostomatous form revealed by a thin
section.

At Lovaig Bay, about 2 miles WNW. of Loch Bay, there is a small outcrop of Lower
Ostrea Beds below basalt, and richly fossiliferous blocks of limestone are strewn on the

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 343

beach. One of these yielded L. hebridica, Pleuromya robusla (abundant), Corbula
hebridica, Anisocardia cucuUata, My tikis (Falcimytihis) sp. Blocks of the algal bed also
occur.

Duirinish succession

Only loose blocks of the Lower Ostrea Beds occur at List, in the stream draining Loch
Mor and on the shore. They include the algal bed. One block of limestone yielded
Corbula hebridica, Pleuromya robusta, and echinoid spines (in thin-section).

Raasay, Strathaird, and Eigg make no additions to the faunas from north Skye.
A. cucuUata is recorded from Raasay (Lee 1920). A thin-section from Eigg showed a
regular echinoid plate and spine. This occurrence of regular echinoids in the oyster beds
recalls the well-known association of Hemicidaris with the Cinder Bed in the Purbeck.

Muck succession

The typical oyster beds are better exposed in Camas Mor, Isle of Muck, than any-
where else (Hudson 1962). The transition from Concretionary Sandstone Series shales,
with Neomiodon, is also well seen. Harker (1908) included these shales in the Ostrea
Beds, and said that the oysters increased in size up the succession. According to my
observations, L. hebridica comes in sharply, and is its normal size at the base. The algal
bed occurs near the top of the exposed succession. A limestone about 5 ft. below this
reveals numerous foraminifera in thin-section, mostly Miliolids.

Some of the fossils collected by the Geological Survey are wrongly listed in the
Memoir (Harker 1908, p. 33). Lithologies and catalogue entries show that the following
groups of specimens are truly from the Lower Ostrea Beds. I have revised the identifica-
tions. All are in the Survey Museum, Edinburgh.

1 . T 3238a-T 3247a. Base of Ostrea Beds.

Modiolus sp., Placunopsis socialis, fish scales.

2. T 3227a-T 3237a. Probably about 5 ft. below the algal bed.

Liostrea hebridica, Placunopsis sp. or spp., small lamellibranchs and gastropods;

Pholidophorus sp. (ident. by Kitchin).

3. T 3221A-T 3226a. Ostrea Beds isolated among dolerite intrusions, i.e. near top of

formation.

Liostrea hebridica, including some very large specimens.
g. Ostracod Limestones

The fauna of this formation is extremely restricted. The typical lithology is alternating
shale and argillaceous limestone, but the limestones are more muddy and the shales
usually more calcareous than in the Estheria Shales. Mudcracks are very common.
Many of the limestones are crowded with ostracods, and Euestheria is common; Vivi-
parus scoticus is common in certain beds but other molluscs (notably Neomiodon) are
rare. Scattered fish remains are frequent.

The most important exposure in Trotternish is at Kilmaluag Bay (437751, map in
Anderson 1948, p. 129). Near the base of the formation, grey sandy shales with plant
remains (5 ft.) are overlain by white calcareous shales with pyrite nodules and mudcracks

344

PALAEONTOLOGY, VOLUME 6

(2 ft.). Both these beds yield ostracods, Euestheria murchisoniae (unusually large in the
calcareous shale) and lamellibranchs comparable to Quenstedtia? stqffinensis, but
generically uncertain. Higher up a limestone with Viviparus is interbedded with ripple-
marked sandstone; the exact succession is difficult to determine because of discon-
tinuous exposures. Yen (1948) collected fossils from here, but I have been unable to
locate the beds in my succession. They were said to be 9 ft. above the Lower Ostrea Beds.
Yen’s succession, written out from his description (1948, p. 167), is;

ft.

in.

4. Sandy limestone

1

0

3. Black shale

0

H

2. Fine-grained cementstone

0

4

1. Sandy limestone

1

0

Bed 2 yielded two species of Procerithium, ‘the sandy limestone’ (it is not stated which)
yielded Viviparus scoticus, V. bithnyoides {Bathonella of Yen), and Neomiodou sp.
(Cyreua jamesoui Forbes). These Procerithiums are the only marine fossils from the
Ostracod Limestones. I have not found a rock like that containing them elsewhere in
the Great Estuarine Series, but similar small gastropods occur in the Estheria Shales
with Cuspidaria and Placunopsis. They do not occur in the same bed as Viviparus.
Anderson and Cox (1948) record Assiminea skyensis from 5 ft. above the base of the
Ostracod Limestones here, and Cox (1950) records Hydrobia praecursor from the
Ostracod Limestones of Skye, locality unspecified but probably from Trotternish. Both
these genera of small gastropods inhabit brackish water today (Anderson and Cox 1948).

In Strathaird, Eigg, and Muck the Ostracod Limestones yield only ostracods, Eues-
theria, Viviparus scotieus, and fish fragments V. scoticus (‘Paludina’) gives its name to the
Paludina Limestones of Wedd (1910). The Geological Survey’s specimens from Muck
are wrongly listed in the Memoir (Marker 1908, p. 33) as from the Ostrea Beds. Their
true source is easily established from lithology. The fossils (T 3199A-T 3220a) comprise
ostracods, fish fragments, Euestheria murchisoniae, and Viviparus scoticus.

Thus, except for one thin bed at Kilmaluag, the Ostracod Limestones contain a low-
salinity assemblage throughout. The water must have been almost continuously muddy,
and mudcracks show that it very frequently dried up. These facts may account for the

EXPLANATION OF PLATE 53

Figures natural size except 3, 4; figs. 1, 4, 5, 6 coated with ammonium chloride. Sedgwick Museum
registered numbers.

Figs. 1, 2. Uuio audersoui sp. nov. In, b, Holotype, Lower Ostrea Beds?, near Duntulm, Skye; J. 48684.
Reversed stereo-pair, separation 6-5 cm. The specimen is an internal mould. 2, Mytilus Shales, near
Kildonan, Eigg; J.48685. Left valve with nacreous shell preserved.

Figs. 3, 4. Placunopsis socialis Morris and Lycett. Mytilus Shales, near Kildonan, Eigg. 3, J. 48686,
left valve. 4, J.48687, left valve of large specimen of inflated form. Both x 2.

Fig. 5. Cuspidaria ibbetsoni (Morris). Mytilus Shales, near Kildonan, Eigg; J.48688. Left valve.

Fig. 6. Mytilus (Praeniytilus) strathairdensis Anderson and Cox. Mytilus Shales, near Kildonan,
Eigg; J.48689. Left valve with nacreous shell preserved.

Fig. 7. Myopholas acuticostata (J. de C. Sowerby). Lower Ostrea Beds, Duntulm, Skye; J. 48690.
Right valve.

Fig. 8. Pleuromya robusta (Tate). Lower Ostrea Beds, Lovaig Bay, Waternish, Skye; J.48691. n. Right
valve, b. Anterior view, c. Dorsal view.

Palaeontology , Vol. 6

PLATE 53

HUDSON, Middle Jurassic Mollusca

J. D. HUDSON: FAUNA OF THE GREAT ESTUARINE SERIES 345

absence of most of the usual mollusca, especially Neomiodon. It is also possible that
the lagoons at this time were almost isolated from the open sea, so that even if salinity
did increase (for instance during evaporation) marine forms could not enter the area.

h. Mottled Clays

These red beds are almost completely unfossiliferous, and were probably deposited
largely above average water-level.

/. The Great Estuarine Series as now defined (Hudson 1962) ends with the Mottled
Clays. In Trotternish it is overlain by the Upper Ostrea Beds, with a somewhat reduced
marine fauna, and the Belemnite Sands, with a fully marine fauna. These faunas have
been listed by Anderson and Cox (1948). They both include Liostrea hebridica and
Neomiodon hryeei.

SYSTEMATIC DESCRIPTION

Genus unio Retzius, 1788
Unio andersoni sp. nov.

Plate 53, figs. 1,2; text-fig. 3

Diagnosis. Species of Unio with the following generic characters: transversely elongate
lamellibranch, with nacreous shell composed of aragonite. In the left valve, two large

TEXT-FIG. 3. Unio andersoni sp. nov. Interior of left valve, X 2. Somewhat idealized reconstruction
to show Unionid musculature and dentition, l.t., lateral teeth; c.t., cardinal teeth; a. a., anterior
adductor muscle impression; a.p.r., anterior pedal retractor; p.p., pedal protractor; p.a., posterior

adductor; p.l., pallial line.

cardinal teeth anterior to the umbo, with a socket between them; two long lateral teeth,
posterior to the umbo. In the right valve, one large cardinal tooth, sometimes (at least)
divided; lateral teeth not seen, but probably one only. Anterior adductor muscle im-
pression deep, anterior to and below the cardinal teeth. Anterior pedal retractor between
anterior adductor and cardinal teeth ; pedal protractor immediately posterior to anterior
adductor. Posterior adductor large, high, shallow. Pallial line simple. External liga-
ment long, posterior to umbo.

346

PALAEONTOLOGY, VOLUME 6

Specific characters; shell small for a Unio, ovoid, length nearly twice the height,
compressed. Umbones depressed, scarcely projecting above hinge-line, situated at the
anterior third of the length. Outline smoothly ovate, the ventral margin very gently
convex. Inflation smooth, at maximum below the umbo ; posterior ridge feebly developed
or absent. Shell rather thin for a Unio, unornamented except for faint growth-rugae.

Holotype. Sedgwick Museum no. J48684; Lower Ostrea Beds?, Great Estuarine Series (Bathonian),
stream 1 mile south of Duntulm Castle, Isle of Skye, Scotland.

Dimensions. Holotype; length 4-5 cm., height 2-5 cm., thickness through both valves (approximate)

1 -5 cm. These dimensions are approximately constant for well-preserved, apparently adult, individuals,
in the many specimens examined. The maximum length seen was 5-5 cm.

Name after Dr. F. W. Anderson in recognition of his researches on the Great Estuarine Series.

Material. All from the Great Estuarine Series, Middle Jurassic, of the Inner Hebrides.

Locality 1. As for holotype; loose blocks, probably from the Lower Ostrea Beds. The holotype and
four other specimens. The holotype is the most complete specimen known. It originally had some shell
attached, but was developed as an internal mould to show dentition and musculature. Reversed stereo
photographs, used at the suggestion of Dr. C. L. Forbes, enable the mould to be seen as a three-
dimensional shell, and greatly help in interpreting the structures.

Locality 2. Reptile Bed, Mytilus Shales, Eigg. My collection, several specimens mostly fragmentary.
Geological Survey, Edinburgh: T 2843a, T 2862a, T 2878a (cf. Jackson 191 1«, pi. 10, fig. 2, neanic
form of U. distortns), T 2902a, T 2928a-T 2930a.

Locality 3. Unio Bed, Mytilus Shales, Eigg. My collection, fifteen specimens, some with nacreous
shell. Geological Survey, Edinburgh; T 2944a-T 2947a. Almost certainly from this bed, from the
matrix. T 2944a has part of the ligament preserved.

In the Geological Survey, Edinburgh, are some small Unios from the Brora Estuarine Series which
may be conspecific with U. andersoni (R and S 3564-7). None of them shows musculature or dentition.
A small Unio from Brora was referred to U. distortns by Jackson (1911o, p. 120).

Discussion. The shell characters are undoubtedly those of a Unionid (text-fig. 3) and
compare closely with those of modern Unio (e.g. Cox 1961, p. 329). The species appears
to be more closely related to Unio than to Margaritifera, from the list of shell characters
given by Mongin (1961, p. 341).

The only described Middle Jurassic Unios from Britain are from the Yorkshire
‘Estuarine’ Series. In internal characters my species agrees with U. kendalli Jackson
(191 IZ)) from the Lower Estuarine Series (Bajocian), but in shape of shell and lack of
ornament it more closely resembles U. distortus Bean (Jackson 191 la) from the Upper
Estuarine Series (Bathonian?). The internal characters of U. distortus are unfortunately
not accurately known, and it is also about twice as large as my species. Jackson (1911a)
referred it tentatively to Margaritana (= Margaritifera).

Various genera, or subgenera, of Unio, have been proposed for Mesozoic Unionids
from North America and Japan; Dr. L. R. Cox kindly sent me a list of these. The only
one which at all resembles my specimens is Rhabdotophorus Russell 1935, from the
Cretaceous of Alberta. The internal structures, size, and outline are similar, but it has
costae on the post-umbonal slope which my species lacks. The geological horizon and
locality are also very different. Until a comprehensive review of Mesozoic Unionacea
is undertaken it seems best to leave the species in Uitio sensu lato.

Acknowledgements. I wish to thank Dr. F. W. Anderson, Dr. L. R. Cox, and Dr. R. Casey for dis-
cussions on the mollusca, and Dr. Anderson, Mr. R. B. Wilson, and Dr. R. M. C. Eagar for access

J. D. HUDSON; FAUNA OF THE GREAT ESTUARINE SERIES 347

to and loan of specimens in their charge. Dr. G. Larwood read and criticized the first draft of the
manuscript.

REFERENCES

ANDERSON, F. w. 1948. Algal beds in the Great Estuarine Series of Skye. Proc. R. phvs. Soc. Ed'mb.
23, 123^2.

and cox, l. r. 1948. The ‘Loch Staffin Beds’ of Skye, with notes on the molluscan fauna of the

Great Estuarine Series. Ibid. 23, 103-22.

ARKELL, w. j. 1934. The oysters of the Fuller's Earth . . . Proc. Cotteswold Nat. Fid. Cl. 25, 21-68.
— ■ — • 1948. Batlionella and Vivipariis. Geol. Mag. 85, 227.

BARROW, G. 1908. In The Geology of the Small Isles of Inverness-shire. Mem. Geol. Surv. U.K.
H.M.S.O., Glasgow.

BOCK, w. 1953. American Triassic Estherids. J. Paleont. 11, 62-16.

CASEY, R. 1952. Some genera and subgenera, mainly new, of Mesozoic heterodont lamellibranchs.
Proc. malac. Soc. Fond. 29, 121-80.

1955. The Neomiodontidae, a new family of the Arcticacea (Pelecypoda). Ibid. 31, 208-22.

cox, L. R. 1947. The lamellibranch family Cyprinidae in the Lower Oolites of England. Ibid. 27,
141-84.

1948. Batlionella and Vivipariis. Geol. Mag. 85, 313.

1950. Bathonian Lm^uraMike gastropods. Ibid. 87, 228-30.

— — 1960. Thoughts on the classification of the Bivalvia. Proc. malac. Soc. Fond. 34, 60-88.

1961. Observations on the family Cardiniidae (Class Bivalvia). Ibid. 34, 325-39.

and ARKELL, w. J. 1948-50. A survey of the Mollusca of the British Great Oolite Series. Palaeon-

tograpliical Society, London.

FISCHER, j.-c. 1961. Sur I’origine du niveau a coquilles paludiniformes du Bathonien de I’lndre.
C.R. Soc. geol. Fr., Annee 1961, 81-83.

FORBES, E. 1851. On the Estuary Beds and the Oxford Clay at Loch Staffin, in Skye. Quart. J. geol.
Soc. Fond. 7, 104-13.

FORSYTH, I. H. 1960. A marine shell-bed near the base of the Estuarine Series in Raasay. Trans.
Fdinb. geol. Soc. 17, 273-5.

GUNTER, G. 1950. The generic status of living oysters and the scientific name of the common American
species. Amer. Midi Nat. 43, 438.

HALLAM, A. 1959. On the supposed evolution of Gryphaea in the Lias. Geol. Mag. 96, 99-108.
HARKER, A. 1908. In The Geology of the Small Isles of Inverness-shire. Mem. geol. Surv. U.K.
H.M.S.O., Glasgow.

HUDSON, J. D. 1962. The stratigraphy of the Great Estuarine Series (Middle Jurassic) of the Inner
Hebrides. Trans. Fdinb. geol. Soc. 19, 139-65.

JACKSON, J. w. 191 lu. On Uiiio distortus Bean and Alasniodon vetustus Brown, from the Upper
Estuarine Beds of Gristhorpe, Yorks. Naturalist, Fond, for 1911, 104-7, 119-22.

19116. A new species of Unio from the Yorkshire Estuarine Series; with notes on other forms.

Ibid. 211-14.

JONES, T. R. 1862. A monograph of the fossil Estheriae. Palaeontogr. Soc. [Monogr.].

KOBAYASHi, T. 1954. Fossil Estherians and allied fossils. J. Fac. Sci. Tokyo Univ. Sect. 2., 9, 1-192
KORRiNGA, p. 1952. Recent advances in oyster biology. Quart. Rev. Biol. 21, 266-308, 339-58.

LEE, G. w. 1920. The Mesozoic rocks of Applecross, Raasay and north-east Skye. Mem. geol. Surv.
U.K. H.M.S.O.

MACGREGOR, M. 1934. The sedimentary rocks of North Trotternish, Isle of Skye. Proc. Geol. Assoc.,
Fond. 45, 389^06.

MONGIN, D. 1961. 'Unio' valdeiisis Mantell, from the Wealden Beds of England . . . Proc. malac.
Soc. Fond. 34, 340-5.

NEWELL, N. D. 1942. Late Palaeozoic pelecypods: Mytilacea. Univ. Kansas Publ. State geol. Surv.
Kansas, 10, part 2, 1-80.

R ANSON, G. 1939. Le provinculum et la prodissocoque de quelques ostreides. Bull. Mus. Hist. nat.
Paris, 2® ser., 11, 318-31.

A a

C 1213

348

PALAEONTOLOGY, VOLUME 6

RANSON, G. 1942. Note sur la classification des ostreides. Bull. Soc. geol. Fr., 5® ser., 12, 161-4.
RAYMOND, p. E. 1946. The genera of fossil conchostraca — an order of bivalved Crustacea. Bull. Mus.
comp. Zool. Harv. 96, 218-307.

RUSSELL, L. s. 1935. Fauna of the Upper Milk River Beds, southern Alberta. Trans, rov. Soc. Can.,
Series 3, 29, 115-28.

segerstrAle, s. G. 1957. Baltic Sea. 7/;hedgpeth, j. w., ed.. Treatise on Marine Ecology and Paleo-
ecology, vol. 1. Mem. geol. Soc. Amer. 67 (1).

SEiLACHER, A. 1953. Studicn zur Palichnologie II. Die fossilen Ruhespuren (Cubichnia). Neues Jb.
Geol. 98, 87-124.

TASCH, p. 1956. Three general principles for a system of classification of fossil conchostracans.
J. Paleont. 30, 1248-57.

TATE, R. 1873. In BRYCE, J. On the Jurassic rocks of Skye and Raasay. Quart. J.geol. Soc. Lond. 19,
317-51.

WATSON, H. 1950. A note on Bathonian gastropods assigned to freshwater genera. Geol. Mag. 87,
17-25.

WEDD, c. B. 1910. In The geology of Glenelg, Lochalsh and south-east Skye. Mem. geol. Surv. U.K.
H.M.S.O.

YEN, T. c. 1948. On some Bathonian mollusca from Skye. Geol. Mag. 85, 167-71.

YONGE, c. M. 1949. The Sea Shore. London.

— — 1960. Oysters. London.

J. D. HUDSON

Department of Geology,.
Sedgwick Museum,

Manuscript received 16 October 1962 Cambridge

MICROBIOLOGICAL COLONIZATION AND
ATTACK ON SOME CARBONIFEROUS
MIOSPORES

by L. R. MOORE

Abstract. The paper describes Palynomorphites diversifonnis gen. et sp. nov., a saprophytic organism thought
to be related to the Fungi, and which is responsible for some part of the microbiological attack on Carboniferous
miospores. The different organization of this attack on several groups of miospores of widely varied structural
characteristics is described. The miospores were preserved in a chert band within the Upper Oil Shale Group of
the Calciferous Sandstone Series of Scotland at a horizon considered on the evidence of goniatites to be of
Upper Visean age.

During studies of the micropalaeontology and constitution of the Scottish Carboni-
ferous Oil Shales, problems of the recognition and possible interrelationships of the
organic contents of the Oil Shale were encountered. The presence of a relatively opaque
organic groundmass in close association with inorganic material led to difficulties in the
resolution of the finer organic material as seen in normally prepared thin-sections. This
was also true of larger organic entities such as wood cells, spores, or algae embedded in
the groundmass. To some extent these difiiculties were overcome by the production of
microtomed sections and by the application of peel techniques to polished and etched
surfaces. On the other hand, the improved techniques indicated the presence of a con-
siderable content of fine cellular material within the groundmass and upon the surface
of many larger remains such as fragments of wood and spores. The application of
maceration techniques resulted in the isolation of some ingredients e.g. spores and algae,
and provided substantiation of the presence of microbiological remains upon their
surface or within their body. These studies also indicated the possibility of selective,
partial, or complete destruction of the various organic constituents during the varied
maceration processes. However, in favourable preparations considerable quantities of
fine thread-like filaments, or small rounded cells often in groups or chains, were present
within the groundmass and upon subjects embedded within it. This material, present to
some extent in all the oil shales studied, appeared to play a significant part in the con-
stitution of certain types of oil shales.

In order to investigate what appeared to be evidence of microbiological attack, pos-
sibly of a saprophytic character, upon the organic material, other rocks within the Oil
Shale succession were studied. These included calcareous shales, limestones, ironstone
nodules, and chert nodules and bands. This account describes the evidence provided by
one chert band, and from the sum total of the microbiological attack on a wide variety
of organic remains presents that evidence concerned with the attack on miospores.
Subsequent papers will be concerned with the attack on other plant remains by a variety
of microbiological agents.

Brief summary of previous work. There is considerable literature concerned with the
presence of microbiological remains of a type resembling in morphological details those

fPalaeontology, Vol. 6, Part 2, 1963, pp. 349-72, pi. 54-56.]

350

PALAEONTOLOGY, VOLUME 6

herein described. The occurrences relate to most geological systems from the Precam-
brian (Tyler and Barghoorn 1954) to the present day, with particular abundance within
the Tertiary (Meschinelli 1902). The remains have been recorded from a wide diversity
of rock types in association with fossil plant and animal remains, or as isolated entities.
The ubiquitous distribution is striking and references to either Fungi, Algae, or Bacteria
are equally varied and controversial. In many cases the association of microbiological
remains with recognizable fossil plants or animals led to a direct comparison with existing
organisms and their known methods of attack, and resulted in identification of genera
and species which imply genetic affinity with members of these three main classes.

Bacteria have long been regarded as the main instruments of organic decay and an
early record of their fossil occurrence was due to Van Tieghem (1879), who recorded
Bacillus amylobacter from silicified Coal Measures rocks of St. Etienne. This work was
followed by Renault (1896, 1900), who described and named a great variety of bacteria,
both micrococci and bacilli. On the basis of their morphology and association with
particular organic material, a number of genera and species were erected principally for
Carboniferous and Permian forms.

A further group of organisms consisting of fine filaments from 0-7 to 2-0/u, in dia-
meter have received consistent reference in literature. Thus Wedl (1858) described
remains from animal shells of various geological ages as due to the ‘alga’ Saprolegnia
ferox (Kiitzing), while other authors, e.g. Kolliker (1859), referred similar organisms to
the Fungi. Renault (1900) referred to fine aseptate filaments of the saprophytic fungus
Phellomyces in the bark cells of Carboniferous plants ; similar remains from carbonaceous
shales, oil shales, and cannel coals were referred to the saprophyte Anthracomyces can-
nellensis.

Kidston and Lang (1921) described forms of somewhat greater geological age con-
sisting of a felt of aseptate unbranched tubular filaments 2/x in diameter from the Rhynie
chert. The organisms, designated Schizophyta No. 2 and No. 3, were compared with
Archaeothrix contexta and A. oscillatoriformis, and, while they were regarded as possible
fungal hyphae, they were believed to be suggestive of the Trichobacteria, Beggiatoa or
Oscillatoria.

In recent years organisms with affinities between Fungi and Bacteria have received
considerable attention as the Actinomycetes. Although some indecision is evident re-
garding the systematic position of these organisms from the works of Drechsler (1919),
Henrici (1947), Waksman (1950), and Bergey (1948), there is a clear understanding of
their morphological characteristics. Thus according to Waksman, the Actinomycetes are
organisms forming elongated, usually filamentous, cells ; the filaments are usually Ip or
less and do not exceed 1 -5ft in diameter. They multiply by special spores (oidiospores)
or by conidia; the oidiospores are formed by segmentation or simple division of the
filaments, whilst conidia are formed singly at the ends of simple conidiophores. The
genus Actinoplanes, a fungus allied to the Actinomycetes, was responsible for the attack
on certain modern pollen described by Goldstein (1960).

There have been few references to the attack of micro-organisms on fossil spores ; this
is particularly true for those of Carboniferous age. Reinsch (1884) figured and described
what he believed to be fungi on spore coats, and Renault (1900, p. 204) described the
presence of the saprophytic fungus Anthracomyces cannellensis on the surfaces of micro-
and megaspores, and from within the latter.

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 351

The attack of modern aquatic and soil-inhabiting fungi on pollen has long been known
to botanists. Goldstein (1960) gave a summary account of some of this work and de-
scribed the attack by Phycomycetes, both filamentous forms and Chytrids. He draws
attention to the importance and interest of the degradation and destruction of pollen
grains in pollen analysis studies, and amongst other conclusions refers to the different
modes of attack of the various fungi on pollen, and to the greater susceptibility to attack
of some types of pollen grains.

Sangster and Dale (1961) carried out controlled experiments on three genera of pollen,
Populiis, Finns, and Typha, placed in four different habitats: pond, lake, swamp, and bog.
Their work indicated considerable variability in preservation and fossilization of these
genera under identical conditions. They concluded also that the decomposition of a
given pollen is dependent on the environment which affects not only the rate of decom-
position, but the nature of the processes involved. The processes were in part micro-
biological, and, particularly in the cases of Populus and Typha, were due to bacteria,
although these organisms were not described nor the nature of the decomposition dis-
cussed. Other experimental evidence indicated oxidation to be an important factor in
disintegration for certain environments and suggested that the resistance of the exine to
an oxidizing agent favours fossilization. The Finns pollen was very resistant to disin-
tegration and was well preserved in all four sites; Typha pollen was preserved only in the
bog; Popnlns disintegrated at all four sites. The authors concluded that the exines possess
in varying proportions the material ‘sporopollenin’, a substance of extraordinary chemi-
cal resistance; they drew attention to differences in the percentage frequency of species
in the pollen rain spectrum and the fossil spectrum. These differences follow from the
differential preservation of the exine, and also from the fact that bog environments or
stagnant water will not necessarily ‘fossilize’ pollen grains.

Studies on the degradation of modern and near-recent plant tissues have focused
attention upon the importance of the microbiological action of fungi and bacteria as the
most effective agents of degradation. Many authors, including Barghoorn (1952), Siu and
Reese (1953), Savory (1954), and Reese (1959), have described these processes and the
factors which affect susceptibility to microbial attack. Of considerable relevance to the
present study is the readiness with which cellulose is attacked and the effect of the amount
and nature of the non-cellulosic components. Where the latter are resistant, e.g. lignin
and waxes, the more resistant is the structure to microbiological decomposition. Barg-
hoorn (1952) drew attention to the pronounced differences in the rates of degradation of
various cellulosic layers of a single cell wall. This differential degradation of cellulosic
lamellae is stated to be a consistent feature of anaerobic decomposition. It is thought to be
due to chemical differences in the more resistant lamellae of the cellulosic framework and to
the presence of substances, notably lignin, which retard the hydrolysis of cellulose. Lignin
is stated to be a primary factor in retarding the action of many fungi and most bacteria.

The mechanism of microbiological attack upon cellulose described by Siu and Reese
(1953) and Reese (1955, 1959) is dependent upon physico-chemical action. The active
hyphal tip of the fungus produces hydrolytic enzymes which diffuse into the substrate
and digest it, the hypha growing into the digested region and maintaining contact with
the substrate. The growing fungus appears to follow the cellulose distribution and in
certain cases seems to have followed the orientation of the cellulose fibrils (Bailey and
Vestal 1937, Savory 1954, Reese 1959).

352

PALAEONTOLOGY, VOLUME 6

The possibility of an interrelationship between the presence and activity of the attack-
ing organism and the distribution of materials of varied constitution over the spore body
cannot be overlooked. This may be brought about by distribution of material resulting
from major differences in the structure of the spore, or may be due to differential changes
in composition in a given layer or in different layers of the exine. For this reason the
microbiological attack on miospores is presented in a systematic manner in the following
pages.

THE CHERT BAND

Locality and horizon. The chert band occurs within the Midhope Burn of Midlothian,
where it outcrops on the south bank of the stream at the locality described by Kennedy
(1943, p. 12) and within the following section:

ft.

Dark shales with ironstones bands and nodules 30
Hard, bedded ashy limestone 4

Dark shale with plant remains (Lepidostrobus)

Dark shale with entomostraca and fish remains
Grey micaceous shale 1

Entomostracan limestone
Shale with plant detritus

Black coaly shale, irregular distribution in small
pockets

Chert brown or grey in nodules, mammillated upper
surface — plant remains

Calcareous and ferruginous siltstone, containing chert
nodules and banded black chert
Thin coaly shale
Impure ferruginous sandstone
Black, sulphurous carbonaceous shale with entomos-
straca

Oil shale of inferior quality

This section occurs within the Upper Oil Shale Group of the Calciferous Sandstone
Series, and towards the top of that Group. Kennedy (1943, p. 12) observed the chert
band to contain petrified plant remains and considered that it replaced the normal
position of the Two Foot Coal. The band occurs below the Raeburn Shell Bed which is
exposed within the Midhope Burn and nearer Dovecotes Farm. Stratigraphically the
horizon appears to lie within the Bollandian Stage of the Lower Carboniferous and
possibly within the lower part of that stage, namely Pj. Such a reading of horizon follows
Currie (1954, p. 531) and depends upon the correlation of the goniatite-rich Fordell
Marine Band of Fife, with a horizon near to that of the Raeburn Shell Bed. Currie (1954,
p. 533) also pointed out that corroboration of such correlation follows from the
occurrence and range of Posidonia becheri, which reaches a maximum in the higher
Basket Shell Bed. The latter horizon by analogy with the local maximum in the north
of England would indicate Zone P^b. The chert band probably occurs within P^, possibly
near the base of that zone, and is thus of Upper Visean age.

in.

0

0

6

6

6

2

6

0-2

0-3

0-1

1

2-6

Horizon
of Two Foot
Coal

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 353

Mode of occurrence of the chert. The upper layer of the chert has a nodular or mamillated
surface, the hollows of which are filled by a black laminated coaly shale of very irregular
distribution. The higher or brown chert is remarkably pure and contains well-preserved
macroscopic plant remains ; it is principally from this part of the chert that the organic
remains have been obtained. The lower black and banded chert is organic rich but im-
pure, and evidence of replacement of the original sediment by chert is seen in thin-section
and will be described elsewhere. The combined evidence supports Kennedy’s opinion
(p. 12) that the horizon marks the position of the Two Foot Coal Seam. It points to
penecontemporaneous inundation of a peat or soil by siliceous waters prior to diagenesis
of the underlying sediment. Whether the upper band of the chert contains precisely
autochthonous plant material, or whether that material was introduced by the invading
solutions cannot be ascertained. However, much of the plant material could not have
been transported far in view of the delicate structures preserved. The material represents
the decayed remnants of a terrestrial flora in which the processes of decay were arrested
by saturation in the siliceous water and subsequently preserved in the chert. The methods
of disintegration, the various stages reached, and the selective action of parasitic or
saprophytic attack upon a variety of organic ingredients provides important evidence
of the microbiological agents concerned with the eventual disintegration of organic
material. At the same time, some appreciation of the intermediate and final products of
decay is of inestimable value in a consideration of the organic detritus which would be
swept from a soil or peat into the depositional areas of the oil shale accumulation.

The mode of occurrence of the chert and state of preservation of the organic material
is reminiscent of the remarkable Rhynie chert of Aberdeenshire described by Kidston
and Lang (1921); there is, however, no detailed evidence of peat succession and inun-
dation such as occurred at Rhynie. The wide range of microfossils, including spores and
pollen, wood cells, algae, fungi, and presumed bacteria, together with fragments of insect
remains and possible protozoa, suggests the presence of a very shallow water cover such
as a pool or pond at the time of inundation.

Preservation of plant material. The organic material is preserved within the chert with no
sign of appreciable chemical alteration of the spore coat substances. Such a conclusion
is suggested by examination of thin-sections, and demonstrated by the ease with which
the organic matter can be chemically released by hydrofluoric acid. There is little or no
subsequent alteration of the material by bituminization or coalification processes, such
as would lead to darkening of colour or disruption of structures. Consequently it has
been unnecessary to subject the material to maceration techniques other than that in-
volving hydrofluoric acid digestion; therefore the difficulties of partial or selective de-
struction of some material in attempts to clear the remainder have been avoided.

Chert provides the best possible medium for the preservation and study of micro-
biological remains. It is homogeneous and its hardness prevents compression. The relative
insolubility, and the unjointed and non-porous character of the rock, preclude the intro-
duction of a microbiological population by groundwater or surface solutions. Like-
wise, it is resistant to weathering and erosion and prevents the subsequent introduction
of those microbiological remains associated with rock weathering and disintegration.

Techniques of study. The upper brown or grey chert was cut into a rectangular block and
thus freed from the possibility of surface contamination. Smaller slices from this block

354

PALAEONTOLOGY, VOLUME 6

were placed in polythene containers to which 40 per cent Analar hydrofluoric acid was
added. Alight-brown sludge settled on to the base of the container from which the hydro-
fluoric acid was removed by repeated dilution with distilled water. This was carried out
manually at high acid concentration, and later by means of a suction pump using an
inverted micro-porcelain filter until acid reaction was negligible. The filter was attached to
a suitable cover and set at a constant level within the container, thereby causing a mini-
mum amount of mechanical movement in the organic material and preserving detailed
structures. The organic material, taken direct from the polythene container, was mounted
in glycerine jelly while still moist. Since the difference of refractive indices between the
material and mounting medium was sufficient to provide good resolution of detail at high
magnifications, the majority of the slides were thus prepared. Some of the organic
material was stained with safranin or Cotton Blue with the original objective of im-
proving resolution of detail. However, the material proved to be selective in its absorp-
tion of stain and particularly so with safranin, and this selectivity provided an initial
basis for the recognition of related but scattered organic components; some remains in
the slides were avid in their absorption of Cotton Blue. Concurrently with examination
of these mounts, the study of thin-sections served to demonstrate the presence of similar
material within the chert, but the mounted isolated organic material possessed consider-
able advantages for study of detail. The mounts have been prepared for a number of
years and in some instances a series of photographs taken of the same object. In no
single case have there been any changes in the nature or distribution of the microbio-
logical remains associated with the spores and pollen described in this paper. Under
these circumstances the author is confident that an indigenous microflora was respon-
sible for the saprophytic attack, and that such studies can be conducted with advantage,
particularly when chert is the parent rock.

THE MICROBIOLOGICAL ATTACK

The various forms described are all characterized by a prominent, dark, and relatively
thick cell wall which encloses colourless or pale-yellow contents of high refractivity.
The commonest forms are spherical cells of the coccoid type which range in size from
0-75jLA to but a frequent modification provides larger ovoid cells which reach l-2jx
on the longer axis.

The cells may occur singly or in pairs of either rounded or ovoid forms, and this
arrangement is usually in the ‘end-on’ position when the appearance resembles the
diplococcoid organization. Occasionally a T-shaped grouping of three cells is observed,
but more commonly an irregular grouping of the cells results. Increase in the number of
spherical individuals in the ‘end-on’ position forms a chain which resembles the strepto-
coccal organization. The presence of predominantly ovoid or elongate cells in a linear
arrangement forms a beaded filament. The latter may be short (2-3 /x) or much longer
(10-20 /x) and occasionally show lateral bifurcation; the termination of such a filament is
marked by a highly refractive and larger spherical cell, but there is no clear evidence of
septation. Rod-shaped forms (2-3 /x) resembling bacilli in appearance are present; these
do not show a beaded structure and may possess either bluntly rounded ends or swollen
terminations. While straight forms are more common, simply curved examples have
been observed.

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 355

Filaments with well-defined parallel walls and resembling hyphae in appearance vary
from 0-75 ft to l-5p. in thickness and may be straight, flexuous, orwith polygonal, curved,
or concentric arrangement, and vary considerably in length. Branching of the filaments
is a common feature and the angle of bifurcation most variable from acute to right-
angled and obtuse. At the point of bifurcation there is usually some swelling and the
point is generally marked by the appearance of a spherical cell. Occasionally single
swollen vesicles may occur along the length of a filament, and many filaments end in
a beaded structure or are represented by a beaded filament at certain points. No evidence
of septation has been observed in any of the filaments examined.

Organization of attack. The small spherical cells are commonly randomly distributed,
but they may show some orientation which follows the structure of the host material.
When this occurs their form becomes ovoid and the long axis of the cell lies in the
orientated direction. Under these circumstances pairs of cells in the ‘end-on’ position
result, and the eventual linear arrangement of elongate cells gives rise to a beaded fila-
ment which possesses a prominent orientation. The filaments commonly show a similar
orientated relationship, particularly when they are in part beaded or are in close asso-
ciation with beaded filaments. However, when the filaments are particularly well developed
they may indicate an independence of structural orientation.

The considerable diversity of elements may suggest more than one kind of organism
to be concerned with the attack. However, the similarity in size and appearance, and the
continued association of all these elements, point to a single organism exhibiting various
stages of vegetative development and which may be observed on the several regions of
the same spore. Modification of the host material following upon changes in the degree
of attack may lead to changes in the vegetative form of the attacking organism at a given
point. In this manner the incidence of attack by coccoid or ovoid cells may lead to the
presence of beaded filaments and finally give rise to filaments. The latter in turn may
produce beaded filaments or fragment into cocci when absorption is completed. Some
supporting evidence occurs in cases where well-defined filaments cross areas of totally
destroyed exine, they become beaded in part and possess large terminal spherical cells.

The biological affinity of the attacking organism is in doubt; morphologically it might
be similar to certain bacteria, actinomycetales, or fungi. The filaments respond only
faintly to staining and are merely tinted with Cotton Blue. From the evidence presented
in this paper it is impossible to decide on the parasitic or saprophytic nature of the
attack. Nevertheless, the presence of this or a similar organism on a wide variety of
organic material, such as could only be brought together as a result of continued
processes of decomposition, suggests the action to have been saprophytic.

This saprophytic attack upon various groups of miospores is described below. In some
instances it has been possible to follow the attack from its incidence to its completion by
selecting spores (of the same genus) which clearly show a progression of events. In this
manner some degree of selectivity of the saprophyte upon the host is demonstrated, and
the structural features of the various spores are shown to exert considerable control over
the distribution and manner of attack; the organism follows these features to a great
extent. In spores where there is no marked structural differentiation the ornament of the
exine, particularly in thick- walled forms such as Convulutispora (p.364), plays an important
part in controlling the distribution of the attack and the manner in which it is carried out.

356

PALAEONTOLOGY, VOLUME 6

SYSTEMATIC DESCRIPTIONS
Fungi incertae sedis

The organism in some of its characteristics bears resemblances to the small sapro-
phytic fungus referred to by Renault (1900, p. 334) as Authracomyces cannellensis and
recorded from a variety of Carboniferous and Permian rocks. Renault considered this
fungus to be closely related to the groundmass of the carbonaceous rocks and to be
generated within this medium, spreading on to the other recognizable organic fragments
contained within it. His insistence upon the role of this fungus as a means of producing
organic detritus under certain conditions is reflected in the speciflc name A. cannellensis.
The fungus was described as forming a small plant 5-6^ in height, with straight sinuous
or branching filaments 2-2-9 jj. long and 0-85 wide. Short branches were noted to end in
a spherical conidium. The propagation of individual plants was believed to form a felt
which spread over the organism attacked.

Renault did not provide a systematic diagnosis of generic characters or give a specific
definition, and no reference was made to the method or organization of the attack by the
fungus. Considerable difficulties are involved in interpreting the very general and much-
restricted account of the organism as given by Renault, especially since the present work
has shown a very much wider range of variation on both sides of what might be considered
the mean characteristics referred to by Renault. As a result it is proposed, at a later stage
in these studies, to emend the genus Anthracomyces and to restrict its use for the reference
of similar saprophytic fungi present in the disorganized organic matter of carbonaceous
rocks. The present organism, in view of its considerable morphological variation and the
close relationship of its organization with the structural features of the spores, is referred
to a new genus Falynornorphites. There may well be overlap between the new genus
Palynomorphites and the forms to be included in the emended genus Anthracomyces, but
at the present stage of the investigations it is necessary to refer to form genera.

Genus palynomorphytes gen. nov.

Type species. P. diversiformis sp. nov.

Diagnosis. Single coccoid, ovoid, or bacilli-like cells, or aggregates of such cells ; beaded
or non-beaded filaments, aseptate. 0-75 ft to 1-5 ft in cross-section and occurring on or
within fossil spores.

Palynomorphites diversiformis sp. nov.

Holotype. Plate 54, fig. 13; fungal elements investing a specimen of Densosporites.

Type locality. Slide Mb/Ch/3/4/907260. Chert Band, Midhope Burn, Upper Oil Shale Group of
Calciferous Sandstone Series (Scotland), Upper Visean, Pi subzone.

Diagnosis. Coccoid or ovoid cells 0-75ft to l-5ft arranged in a linear ‘end-on’ position,
and associated aseptate filaments 0-75 ft to l-5ft in width.

Description. Cell walls dark and well defined; cell contents highly refractive, colourless
or pale yellow. The filaments may be straight, flexuous, or curved, or exhibit simple
bifurcation to produce a rounded or reticulate pattern. At the point of bifurcation
a rounded cell is present and filaments may terminate in a swollen rounded cell.

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES

357

Coccoid cells may occur alone as on Pimctatisporites, Plate 55, fig. 10, or be associated
with beaded filaments only as on Pimctatisporites, Plate 56, fig. 2c. Filaments both beaded
and non-beaded may occur in close association and of very variable length. They may
occur together or singly without the normal association of coccoid cells. These features
are exemplified by Lycospora, Plate 54, fig. 1 1 .

Comparison. P. diversiformis sp. nov. in certain characteristics bears resemblance to A.
cannellensis Renault (1900, pi. 22, figs. 4—10; text-fig. 24) but A. cannellensis Renault
(1900, p. 334) was defined as ‘a plant 5-6 p high, with straight branching or sinuous
filaments 2-2-9 p long and 0-85/x wide’.

DESCRIPTION OF MICROBIOLOGICAL ATTACK
ON VARIOUS MIOSPORES

Schulzospora (PI. 54, figs. 1, 2, 3).

The grain of Schulzospora consists of a circular central body, with an enveloping
saccus which may be attached to the proximal polar region of the central area.

The three grains illustrated represent the varying degrees of saprophytic investment
observed for this genus. Thus Plate 54, fig. 1 indicates incipient attack by isolated
irregularly distributed coccoid cells, while in Plate 54, fig. 2, with a greater intensity of
action, some pattern of distribution becomes apparent. With the heavier concentrations
near the equatorial margins of the saccus, and in a region near the junction of the saccus
with the central area, there is a tendency to the formation of chains of cells and beaded
filaments. On the saccus alone there is little definite arrangement but many filaments
cross on to the region underlain by the central area where a distinct radial arrangement
is followed. Many of the filaments branch or coalesce to enclose irregularly shaped
polygonal or rectangular areas of pitted or thin exine. In some cases the proximal exine
has been removed from the polar region of the central area as at Plate 54, fig. 2. A high
concentration of saprophytic material around the junction of the saccus and central area
and over the surface of the latter is demonstrated by Plate 54, fig. 3. The concentric
organization of the disturbed wall substance appears to have resulted from attack in
depth which has totally disorganized the saccus covering and the exine of the central
area. Some part of the disorganized infected exine was partly moved from the body of
the spore during mounting and is seen (PI. 54, fig. 3) to consist of coccoid cells and
beaded filaments.

Other examples of this type of spore were noted in which the central area was either
partly destroyed or even absent. These may indicate the ultimate effects of the form of
attack upon the saccus and central area, or alternatively the partial destruction of the
enveloping saccus may lead to release of the central body. There appears to be some
relationship between the form of the spore and the resulting organization of attack.
Thus the polygonal or rectangular habit of the filaments enclosing the pits on the flatter
polar region of the central area gives place to a radial orientation of elongate cells and
beaded filaments on the sloping sides of that area. The fringes of the saccus show a less
definite arrangement over their flattened surface, but the concentric orientation of elon-
gate cells followed in Plate 54, fig. 3 appears to be directly related to the presence and
influence of the central body. The dark region on the central area of Plate 54, fig. 1

358

PALAEONTOLOGY, VOLUME 6

consists of a dense mass of very thick-walled dark-brown cells of a type diftering from
those under present description.

Lycospora (PI. 54, figs. 4-12)

The illustrated spores represent common forms in the preparations and they have
been selected to show the varying degree of saprophytic attack and its manifestations
on the various structural units of the spore. There is a close correlation between areas of
microbiological action, the nature and orientation of the attacking cells, and the struc-
ture of the spore. Demonstration of this relationship follows from a study of proximal
and distal surfaces, and as a result of separate consideration of the central area and
cingulum. The latter is further subdivided into the thickened inner region (crassitudo)
and the outer equatorial flange.

Plate 54, fig. 4 depicts in proximal view a typical Lycospora with granular ornament
and carrying incipient attack over its surface by isolated or small groups of coccoid cells.
A concentration of elongate cells and beaded filaments with a linear orientation occurs
along the dehiscence slits of the central area. The crassitudo carries a similar concen-
tration of cells orientated in a concentric manner.

The central area. On the proximal surface along the margins of the triradiate marks, the
concentration of linearly orientated cells, and the subsequent development of beaded
filaments are characteristic but variable features. Plate 54, fig. 4 depicts the earlier stages
of this development, which is continued in Plate 54, figs. 5 and 6. The spores of Plate 54,
fig. 10, and particularly Plate 54, fig. 11, illustrate the eventual production of relatively
thick-walled filaments. Each proximal intertectal segment carries independently orien-
tated elongate cells or beaded filaments traversing the exine between the triradiate marks
in broken or continuous lines, as illustrated by Plate 54, figs. 5, 6, 8. Some of the filaments
either branch or coalesce to enclose rounded, rectangular, or polygonal areas of altered
exine, as shown by the circular pale areas of Plate 54, fig. 5a.

The absence of haptotypic features on the distal surface of the spore leads to a difTerent
organization of attack. In the initial stages this consists of rounded coccoid cells which

EXPLANATION OF PLATE 54

All figures are of specimens from the Chert Band, Midhope Burn, Upper Oil Shale Group of the

Calciferous Sandstone Series (Scotland). Upper Visean, Pi subzone. Magnification x750 unless

otherwise stated. Slides all labelled M.B.Ch.

Figs. 1-3. Schiilzospora sp. 1, 3/5/726.334, proximal surface. 2, 3/7/793.200, proximal surface.
3, 3/3/776.341, distal surface.

Figs. 4—12. Lycospora sp. 4, 3/7/712.260, proximal surface. 5, 3/6/779.403, proximal surface; a, rounded
and polygonal areas of altered exine on central body. 6, 3/6/803.360, proximal surface. 7, 3/6/810.247,
distal surface with reticulum of filaments. 8, 3/6/776.295, distal surface; a, single large lumen; b, con-
centrically orientated elongate cells and radial filaments on the equatorial flange. 9, 3/6/730.256, distal
surface, x 1500. 10, 3/6/766.290, proximal surface. 1 1, 3/6/811.230, proximal view of ‘Palynomor-
phites pseudomorph’, X 1500; a, fragments of the equatorial flange with filaments spreading from the
annular zone. 12, 3/7/690.322, distal view, Xl500.

Figs. 13-17. Densosporites sp. 13, 3/4/907.260, distal surface. Type of PalynoJiwrphites diversifonnis
gen. et sp. nov. investing distal surface of Densosporites sp., X 1500. 14, 3/8/805.188, distal surface,
X 1000. 15, 1/4/798.140, distal surface. 16, 3/5/750.197, presumed distal view, concentric arrange-
ments of filaments within annular region, X 1500. 17, 3/5/782.166, presumed proximal view.

Palaeontology, Vol. 6

PLATE 54

MOORE, Microbiological attack on Carboniferous miospores

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 359

are either randomly distributed or show a very general concentric organization, e.g.
Plate 54, fig. 8. The development of elongate cells, beaded filaments, and short true
filaments (PI. 54, fig. 9) results in the formation of irregular areas from which the exine
has been destroyed; a single lumen is shown in Plate 54, figs. 8a and 9. The eventual
pattern of attack for the distal surface is represented by Plate 54, fig. 7, in which the
surface of the central area is represented by a reticulum of filaments.

The Cingulum, thickened inner zone (crassitudo). One of the most remarkable areas of
investment is the thickened annular ring of exine concentric to and bordering the central
area of the spore. Incipient action on this region (PI. 54, fig. 4) is represented by rounded
or elongate coccoid cells orientated longitudinally in a concentric manner. With more
severe attack prominent filaments lie within the structure, and their strong concentric
orientation is particularly well illustrated by Plate 54, figs. 5, 7, 11. Prominent vesicle-like
swellings give rise to filaments which branch on to or into the central area of the spore,
particularly with respect to the distal surface (PI. 54, figs. 7, 12), but also at some points
on the proximal surface (PI. 54, fig. 1 1). Other filaments radiate from this region into
the flange area towards the equatorial margin of the spore, and are seen in Plate 54,
figs. 1 la, dib.

The Cingulum, equatorial flange. This region extends from the outer area of the thickened
zone to the equatorial margin of the spore and consists of a double layer of thin colour-
less exine. The radiating filaments referred to above cross the flange and either end in
a spherical vesicle at the equator of the spore or are connected with beaded filaments
developed there. The equatorial margin is often markedly indented and carries concen-
trically orientated elongate cells and filaments, e.g. Plate 54, figs. 4, 86 and the lower
equatorial margins of Plate 54, figs. 5, 6, 7. The flange is often much reduced in width and
may be absent.

Plate 54, fig. 4 illustrates incipient attack on the equatorial flange by isolated coccoid
cells with some radial and concentric filaments. The area of the flange is often reduced
by increased microbiological activity upon its equatorial margin, e.g. Plate 54, figs. 5, 7,
8, but equally severe action also takes place from the inner margin of the flange at
a stage when the attack on the thickened inner zone (crassitudo) is well defined. Thus
Plate 54, figs. 5-8 show a colourless and irregularly patterned area from which the exine
of the inner margin of the flange has been removed, due either to the concentration of
radial filaments or, more likely, to the absorptive effect of the concentric filaments in the
neighbouring annular zone. This destruction of the inner flange exine is due to micro-
biologieal action, and not to mechanical effects since the remaining flange exine is
entirely isolated from the central body of the spore, except at those points where the
radial filaments which cross this area act as connecting links. In Plate 54, figs. 5, 6, 8 the
filaments alone hold the remainder of the flange to the body of the spore. Such a fragile
connexion serves to emphasize the importance of microbiological destruction of the
external border of the flange, for it is most unlikely that the apparently eroded con-
figuration of the external margin would result from mechanical disintegration without
disrupting the entire flange structure. The absence of a flange from parts of Plate 54,
fig. 6, its almost total absence from Plate 54, figs. 10-12, the presence of delicate filaments
at the present flange margins of Plate 54, figs. 6, 10, 11, indicate disintegration of the
flange to be principally due to microbiological action, and not to mechanical deformation.

360

PALAEONTOLOGY, VOLUME 6

The remarkable specimen illustrated by Plate 54, fig. 1 1 indicates the essential unity of
the microbiological attack in its later organization, to produce a "Palynomorphites
pseudomorph’ of Lycospora, represented by the filaments developed in the thickened
inner cingulum region and along the triradiate marks. The exine from the central area
has been almost completely removed, while mere fragments of the equatorial flange
remain (PI. 54, fig. lla). The severity of this attack is remarkable, and the preservation
of such a delicate structure indicates the action to have taken place after settlement and
with little or no subsequent mechanical transport. The most frequently occurring forms
exhibiting advanced stages of attack are shown (PI. 54, figs. 10, 12) ; where the equatorial
flange has been removed and a well-defined triradiate attack has developed, the remain-
der of the spore breaks up into three triangular segments. These isolated segments,
bounded on one side by concentric filaments and with linear filaments along two other
tapering margins, are common ingredients of the preparations ; they may contain rem-
nants of partially altered original exine.

The attack on Lycospora is very clearly controlled by the structural features of
the spore. The order of this attack is less clearly defined; it is somewhat variable in the
relative time of onset and in the degree of severity on or within the various areas of
the spore. The effect of the microbiological action is to reduce partially or completely the
equatorial flange, to destroy the exine of the central body, and to cause break-up of
the spore into segments. Where the development of filaments has been particularly
strong a ‘pseudomorph’ of the spore results.

Densosporites (PI. 54, figs. 13-17)

The illustrated spores represent common forms of this genus in the preparations and
are selected to indicate both the methods of microbiological attack and its organization
and relationship to the structure of the spore. The method of attack is very similar to
that described for Lycospora, and since the structure of the two spores is essentially
similar this is to be expected. The weaker development of haptotypic features in Denso-
sporites as compared with Lycospora does not lead to such well-dilferentiated features
of attack on the proximal surface of the former. In many cases the action was so severe
that recognition of the two surfaces of the spore was a matter of considerable difficulty.
For descriptive purposes the following regions of the spore are considered: central area
and cingulum, {a) inner thickened zone, {b) outer equatorial flange.

Central area. Plate 54, fig. 13 indicates the elements which constitute the basis of the
investment. They consist of coccoid cells and elongate bacilli-like fragments in asso-
ciation with beaded filaments and short thick-walled branching filaments often carrying
spherical terminal vesicles. The filaments may be straight or curved and towards the margin
of the central area are somewhat radially directed from the inner thickened zone of the
cingulum ; they appear to originate from dense filaments within this zone. The result is
a loose reticulum of filaments which produces an overall concentric pattern of pentagonal
to polygonal lumena. Within each lumen the exine has been either partially or totally
absorbed or destroyed. Plate 54, fig. 14 indicates a completely reticulate distal surface
with partially absorbed exine, while in Plate 54, fig. 15 fragments of exine are only
present around the margins. Some filaments still retain the elements of reticulate pattern-
ing and preserve filamentous connexions across the destroyed middle portion of the

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 361

central area. The eventual result of microbiological action on the eentral area is to
destroy it completely, as shown by Plate 54, fig. 16.

Cingulum, inner thickened zone. The dense brown region which represents an inner
thickened and annular zone of the cingulum is the most heavily invested part of Denso-
sporites. Upon and within this strueture the elements of microbiological attack form an
anastomosing and interweaving bundle of concentric cellular ‘tissue’ which completely
fills this annulus. This organization is particularly well seen in the spores (PI. 54, figs.
13, 14, 16). At certain points, beaded filaments proliferate into thieker masses from
whence arise other filaments which either pass into and over the central area of the spore
or extend into the equatorial flange as far as the present equatorial margin.

Many spores which show an advanced stage of attack upon this feature (PI. 54, figs.
15, 17) exhibit some degree of disorganization within the concentric structure at certain
points. There is a tendency for the filaments to be replaced by swollen masses of coccoid
or elongate cells, which occur in dense groups without any particular orientation. The
break in outline shown by Plate 54, fig. 17 coincides with such an area and is likely to
coincide with the position of a trilete ray affecting the structure of the inner zone. The
eventual result of the attack is to break the thiekened annular zone into fragments which,
by the concentric organization of filaments, can still be reeognized in the preparation as
fragments of Densosporites.

Cingulum, outer equatorial flange. Where the flange is reasonably complete there is little
significant orientation of the attacking cells. Thus Plate 54, fig. 13 carries coccoid cells
and occasional beaded filaments occur at various points upon the flange. With a greater
degree of investment filaments spread from the annulus in a general radial direction
towards the equator of the flange; these are well seen in Plate 54, figs. 13, 15-17.
The fretted nature of the equatorial margins (PI. 54, 15-17) is governed in part by the
position of filaments emanating from the inner thickened zone. At the same time the
existing margin of the spore is marked by elongate cells and beaded filaments which are
not continuous, but which show a concentric orientation (PI. 54, fig. 13). Thus in part,
the attack on the equatorial flange also takes place from its outer margin; its absence at
some points and partial representation at other points in Plate 54, figs. 15-17 is princi-
pally the result of biological absorption rather than mechanical disintegration.

While the attack on Densosporites is mainly controlled by the structural features of
the spore body, the order of this attack on the various features is less clearly known.
The inner thickened zone of the cingulum may be preferentially selected and rapidly
invested with either concomitant or subsequent spreading of filaments into neighbouring
regions of the spore. Alternatively, the attack may be more widespread initially, but, due
to the rapidity of growth within the annulus, the organization there outstrips that at other
places on the spore. The central area is severely affected at an early stage and its entire
removal may be brought about before major changes are effected in the equatorial flange.
The latter may be partly or entirely removed, and when this coincides with the removal
of the central area a " Palynomorphites pseudomorph’ of Densosporites remains, which
consists of a brown mass of concentric filaments representing the inner thickened zone
or annulus of the cingulum as shown in Plate 54, fig. 16. A comparison of the ‘pseudo-
morphs’ of Densosporites and Lyeospora emphasizes the difference in structure between

362

PALAEONTOLOGY, VOLUME 6

the two spores, and illustrates the part played in structural control by the trilete rays.
The eventual break up of the ‘pseudomorph’ after Densosporites mio fragments completes
the biological disintegration of this type of spore.

Leiotriletes (PI. 55, fig. 1)

Plate 55, fig. 1 illustrates the proximal surface of a thick-walled yellow resinous spore
with broadly rounded apices and slightly concave sides. While the exine on the proximal
surface is present and reasonably complete, that of the distal surface has been partially
destroyed. The microbiological attack on this spore appears to be located in two main
regions, (a) near the equatorial margin, (b) along the triradiate structure.

Near the equatorial margin. The body cavity near the equatorial margin carries beaded
filaments which anastomose but in general follow the spore outline; their beaded charac-
ter is visible at several points near the right-hand margin of Plate 55, fig. 1. That this
feature is not to be confused with an optical effect is demonstrated at Plate 55, fig. 2,
where, after following the equatorial outline, a prominent filament curves into the centre
of the spore. There, in conjunction with a filament which has followed a triradiate ray,
it encloses an elliptical area of pale-yellow partially absorbed exine.

Along the triradiate strueture. The broken wall of exine following this structure is accom-
panied along part of its course by isolated coccoid and elongate cells, and beaded and
non-beaded filaments. Occasionally, beaded filaments with swollen terminal vesicles
branch on to the triradiate segments and enclose the elliptical areas of pale partially
absorbed exine referred to above. The characteristic form of the beaded filament and
terminal vesicle is well seen at Plate 55, fig. \b.

Leiotriletes (PI. 55, figs. 2-5)

The illustrations (PI. 55, figs. 2-4) represent an advanced stage of investment upon
a thick-walled resinous triangular spore possessing blunt or flattened apices and concave

EXPLANATION OF PLATE 55

All figures are of specimens from the Chert Band, Midhope Burn, Upper Oil Shale Group of the

Calciferous Sandstone Series (Scotland). Upper Visean, Pi subzone. Magnification x750 unless

otherwise stated. Slides all labelled M.B.Ch.

Figs. 1-5. Leiotriletes sp. I, 3/6/731.245, proximal surface; a, curved filament partly enclosing elliptical
area of partially absorbed exine; b, beaded filament with swollen terminal vesicle. 2, 3/7/665.289,
distal surface taken with ground-glass filter. 3, Same spore, proximal surface taken with ground-
glass filter. 4, Same spore, distal surface in ordinary light illustrating details of attack. 5, 3/6/
728.245, distal surface, X 1500.

Fig. 6. Granulatisporites sp., 3/7/677.312, distal surface with view of underlying triradiate mark, X 1500;
a, beaded filaments surrounding an area from which the exine has been partially removed; b, curved
filaments enclosing areas of thin or totally absorbed exine.

Figs. 7-9. Convolutispora sp. 7, 3/6/765.248, proximal surface, X 1500. 8, 3/5/826.250, presumed distal
surface, focal plane slightly below equator of spore to show surface detail, x 1500. 9, 3/10/731.170,
surface unknown, advanced stage of attack with well-defined and branching filaments.

Figs. 10-12. Pimctatisporites sp. 10,3/11/790.179, proximalsurface, Xl500. 1 1, 3/7/784.169, proximal
surface, partial investment of the surface of the spore by coccoid cells and marginal filaments, x 1 500.
12, 3/10/768.192, distal surface, coccoid, paired cells, chains of cells, beaded filaments on the surface.

Palaeontology, Vol. 6

PLATE 55

MOORE, Microbiological attack on Carboniferous miospores

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 363

inter-radial margins. The distal and proximal surfaces, taken with a ground-glass filter
in order to illustrate major features, are represented by Plate 55, figs. 2, 3 respectively;
Plate 55, fig. 4 represents the distal surface of the same spore in ordinary light.

Distal surface. The surface is marked by coccoid and orientated elongate cells, beaded
and non-beaded filaments, all arranged in a general concentric manner around areas
where the exine is absent. These areas are illustrated by Plate 55, figs. 2, 4 and although
the distal exine has been entirely removed from most of them, thin fragments remain
nearer the margins of the spores. The attack appears to result from the establishment of
isolated centres and to spread outwards, eventually producing a reticulate pattern. The
margins of the lumena in Plate 55, figs. 2, 4 are formed partly by filaments, but also in
part by ridges of incompletely destroyed or unattacked exine. The remaining exine may
represent a thicker part of the spore wall such as would be the case if the original spore
exine was differentially ornamented ; strong features at the equatorial margin of the spore
suggest this possibility. It is evident that the microbiological action has destroyed well-
defined areas of exine so that either an original structure has been emphasized by partial
destruction of the exine, or a reticulate pattern has been created by the attack and is now
represented on the distal surface.

Proximal surface. The method of degradation on this surface is essentially similar but
for the presence of haptotypic features. Between the thickened exine associated with
a triradiate mark and lip and the equatorial margin along the left-hand side of the spore
the exine has been removed from one continuous vacuity; a similar feature also marks
the right-hand side of the spore. However, in the upper segment the exine between the
triradiate structure and the equatorial margin is more complete and contains smaller
well-defined vacuities which produce a reticulate pattern. The indented marginal form
of the larger vacuoles indicates a more advanced stage of destruction, whereby the inter-
vening exine which formed the margins of the smaller lumena has been destroyed,
leading to coalescence into one larger vacuity.

The evidence provided by this thick-walled resinous spore is of particular interest
since microbiological action on both surfaces of the same specimen is represented. It is
principally superficial with independent action on the two surfaces and which initially
leads to destruction of exine from circular or polygonal areas, with the eventual formation
of large, elongate vacuoles. The thickened exine along the triradiate mark is effective in
restricting the organization of the attack on the proximal surface, but that on the distal
surface is not so restricted. It may to some extent be controlled by other factors such as
an original thickness differentiation or material differentiation of the exine. A further
and similar spore (PI. 55, fig. 5) emphasizes the features of the attack described, and
depicts clearly the nature of the organism responsible. Very little of the original exine
remains on the body of this spore. The spores described represent to a great extent
‘skeletons’ of microbiological decay, and differ in important respects from the ‘pseudo-
morphs’ of Lycospora and Densosporites. Principally they differ in the facts that no major
structural elements of the spore are missing, and that some part of the skeleton consists
of original exine which preserves the spore form. The ‘skeleton’ may break down further
into fragmentary remains ; the thick rounded radial extremities of these spores are pre-
served in the preparations.

B b

C 1213

364

PALAEONTOLOGY, VOLUME 6

Gramdatisporites (PL 55, fig. 6)

The distal surface is represented in the focal plane of the illustration, and the triradiate
mark of the proximal surface is visible beneath it. The spore is much compressed, has
broadly rounded extremities with deeply concave sides, and a thin yellow exine orna-
mented with granules. There is little structural differentiation of the spore body, and
consequently a less well-defined distribution and organization of microbiological attack.
The latter is carried out by elongate orientated cells and beaded filaments, but thick-
walled filaments are present and these in particular occur within the spore body. The
equatorial outline is followed, along most of its length, by prominent filaments which
are particularly well developed and proliferate in the neighbourhood of the radial ex-
tremities. The upper extremity is most severely affected; there, beaded filaments arising
from a marginal filament enclose a region from which the exine has been partially
removed. This lighter-coloured area and its surrounding filaments are shown in Plate 55,
fig. 6a. At the concave lateral margin, beaded and non-beaded filaments follow a modi-
fied course. They swing in towards the centre of the spore by means of prominent curved
but irregular loops and thereby enclose areas of thin or totally absorbed exine (PI. 55,
fig. 6b) so inaugurating a coarse reticulate pattern.

Convohitispora (PI. 55, figs. 7-9)

The spores illustrated are characterized by a very thick resinous exine, and rugulate
ornament. The rugulae may either consist of irregular ridges with a rounded cross-section
and protruberances (PI. 55, fig. 8) or of flattened ridges (PI. 55, fig. 7), and are
separated by lower-lying areas. The three spores illustrated indicate the method of micro-
biological attack. In Plate 55, figs. 7, 8 this organization is well seen upon spores which
retain sufficient original structure to warrant recognition of the genus. The spore in
Plate 55, fig. 9 represents a much more advanced stage of destruction, and although the
thicker irregularly distributed exine is still partially visible, reference to the spore genus
is more doubtful.

The proximal surface of a spore with incipient attack is represented by Plate 55, fig. 7,
where elongate cells and beaded filaments predominate but occasional thick-walled
filaments occur. The distribution is very localized and follows a linear organization in
the neighbourhood of two rays of the triradiate mark and a concentric direction at
several points near the equator of the spore. In places the exine has been destroyed and
circular or irregular perforations are surrounded by elongate cells flanking the thicker
exine of the marginal wall.

Plate 55, fig. 8 represents the distal surface of a spore upon which the attack is very
diversified and locally intense. The lower part of the spore is reasonably unaffected; the
prominent rugulae are devoid of attached cells, but the inter-spaces between the rugulae
carry coccoid cells and short beaded filaments. The right-hand margin of the spore
illustrates a progressively greater intensity of investment in which beaded filaments, occur-
ring in the hollows of the surface, produce a concentric pattern. At the upper margin
of the spore, coalescence of the beaded filaments surrounding the rugulae has resulted in
a linear organization of beaded filaments. The region of greatest investment is the dark
area along the left-hand side of the spore, where the entire surface is covered by a fine
cellular mat of elongate cells and beaded filaments which fan out in a ‘ray-like’ manner
from darker centres. The most severe attack is located on the upper central portion of

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 365

the spore, where a red highly refractive spherical body is seen as a sphere on the plate.
This appears to be a reproductive body caused by fusion of the filaments, or it is the focal
point from which filaments radiate in several directions. Around this body the exine has
been totally destroyed, for the light-coloured areas hereabouts are circular or polygonal
vacuities, many of which have coalesced. The method of destruction of the exine is further
emphasized by Plate 55, fig. 9, in which severe action has culminated in the complete
removal of exine from areas in the central regions of the spore and from deep embay-
ments into its equatorial margin. These areas are surrounded by the thicker remaining
exine which carries filaments, some of which extend into and across the vacuities. With
this more advanced stage significant changes are notable in the nature of the attacking
organism. Thus beaded filaments give way to well-defined thick-walled and larger fila-
ments which are straight or curved and commonly branched. Several branches may arise
from or fuse in a common centre which then carries one or more small spherical vesicles
or radiating groups of beaded filaments. The action is also in depth, for the bundles of
filaments cross one another at lower focal planes. The overall organization produces
a reticulate pattern, and at this more advanced stage of destruction the investing organ-
ism may be less seriously controlled by the structure of the remaining exine. Thus
a broadly concentric organization develops in which filaments follow the spore contour
and are interconnected by other radial filaments.

In thick-walled ornamented spores of the Convohitispora type there are no outstanding
differential structural units of the spore body, and the haptotypic features are of minor
importance in controlling the attack. However, the strong ornamentation produces
hollows of relative protection from which centres of microbiological action could be
generated; the same regions contain thinner and possibly more favourable exine for
organic assimilation. The exine is therefore more rapidly absorbed with the formation
of vacuities, and the rugulae isolated by this means. The resulting configuration passes
from one in which the exine has been removed from smaller circular polygonal or
irregular vacuities, to one of larger irregular but rounded vacuities. A ‘skeleton’ spore
results in which both the remaining exine and the attacking organism play a part. The
breakdown of the spore into a number of isolated, irregularly shaped, and thickened
fragments is envisaged as the final stages in its destruction.

Punctatisporites (PI. 55, figs. 10-12; PI. 56, figs. 1-3)

The spores included in this genus are characterized by a circular outline, with little
structural diflferentiation of the body except in the presence of a frequently well-marked
triradiate feature. They are characterized by an infra-punctate or infra-granular exine.
There is consequently little selectivity of this uniform surface for microbiological activity
and a degree of uniformity pertains in distribution and organization of attack. Thus
Plate 55, fig. 10 illustrates a spore generally free of saprophytic cells and which therefore
exhibits the typical infra-granular exine structure over most of its surface. At one point
a colony of saprophytic cells is clearly visible, which at a higher focal plane consists of
orientated beaded filaments and elongate cells with a radial symmetry. The only other
attack on this spore is associated with the triradiate mark, where the dark areas consist
of linear chains of cells and beaded filaments. A greater concentration of saprophytic
material is illustrated for the proximal surface of the spore (PI. 55, fig. 11), where half of
the surface is covered by coccoid cells often paired in the streptococcal manner. They

366

PALAEONTOLOGY, VOLUME 6

possess little definite or regular orientation, for while some oceur as radial clusters,
others towards the margin show a general orientation following the spore contour. The
dark marginal area on the upper border represents a region of concentrically orientated
filaments.

The complete investment of a distal surface is illustrated in Plate 55, fig. 12, where the
principal elements are coccoid cells. Groups of cells with radial arrangements are found
in association with cells containing two or more individuals in streptococcal arrangement,
or with short beaded hlaments 2-3 /x in length. Over the greater part of the spore the
distribution presents an impression of minute reticulation. The dense marginal distribu-
tion produces a concentric pattern which eventually results in a curvilinear distribution
of beaded hlaments. The spore exine is severely attacked and the isolated mass of
cells lying off the margin of the spore (PI. 55, hg. 13) illustrates the elements responsible
for the investment. A possible further stage of this attack is represented by Plate 56, hg. 1,
in which a change in the form of the saprophytic elements is notable. While the strepto-
coccal-like cells are still common there is a preponderance of beaded hlaments and of
short branching hlaments 2-5 /x long. This development has resulted in an irregular but
distinct reticulate appearance particularly evident in the central regions of the spore.
This hgure (PI. 56, hg. 1) is taken at a slightly higher than median focus to accentuate
the affected and partially destroyed exine, which therefore appears as white lines or areas.
An overall concentric reticulate pattern for most of the spore suggests a contour control,
but this is lost towards the margins. There a distinct radial distribution may indicate
a change in the underlying spore exine or merely a region of originally modihed spore
contour.

The four spores described illustrate the increased areal degradation upon a surface
which shows little differentiation. It is questionable whether the infra-granular structure
of the exine exercises much control upon the organization of attack, and yet it is signifi-
cant that the earliest stages of investment are by minute coccoid cells and beaded fila-
ments only. The spores also indicate the degrees of intensity or stages of the attack, for,
as the microbiological action becomes well established and the exine is made viable,
there are significant changes. These result in the development of branching filaments
with the establishment of a reticulate pattern and the final breakdown of the spore wall.

The spore in Plate 56, hg. 2 is referred to the genus Pimctatisporites with some doubt,
in view of the general lack of original exine visible. The investment is predominantly by
elongate cells, beaded and non-beaded hlaments. The latter are 0-75-T5/(x wide and may
be short branched forms of 2-3 p, or long unbranched forms 10-1 5 /x long. There is a very
general concentric orientation of the saprophytic remains, particularly near the margin
of the spore, where the darker regions consist of both types of hlaments. Several of the
hlaments curve inwards towards the centre and are seen in connexion with other con-
centrations of hlaments near the centre of the spore. There is a secondary concentric
arrangement illustrated by the pale patches of Plate 56, hg. 2, and which exhibits some
diversity and may represent progressive stages in the attack. Thus Plate 56, hg. 2a
represents some areas of thin and altered exine upon which occasional coccoid cells and
hlaments remain; they are surrounded by a diffuse boundary along which occurs a con-
centration of orientated cells and hlaments which form an incomplete concentric boun-
dary. At Plate 56, hg. 2b the structure is larger and the exine from the upper surface of
the spore has been removed. The boundary is formed by concentrically-arranged beaded

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 367

filaments and by filaments which are in connexion with well-defined thick-walled fila-
ments traversing other areas of the spore. The structure at Plate 56, fig. 2c is surrounded
for the most part by a concentric bundle of beaded filaments, the left-hand margin has
suffered mechanically, and the exine is folded upon itself. The exine from the upper
surface of the spore has been removed, but a part of that of the lower surface is still
represented.

This organization appears to be significantly different from that described for other
members of this group and appears to commence with an original cover of saprophytic
material distributed in a broadly concentric pattern. Within this general pattern, local-
ized areas have been affected more seriously than others, and the spread of the attack
outwards results in a concentration of filaments around the margins. Similar features
are shown around the marginal region of the spore (PI. 56, fig. 3).

Microreticulatisporites (PI. 56, fig. 4)

The surface of this spore carries a great number of coccoid- and streptococcal-type
cells of two or three units, dispersed over the surface in irregular groups. There are,
however, two regions of the spore where some organization is noticeable. Firstly, the
equatorial margin is densely covered and the cells are orientated in a concentric manner
and may pass into beaded filaments as near the lower border; the broken left-hand
margin indicates the cellular nature hereabouts and gives little evidence of the presence
of exine. Secondly, the most remarkable organization is that based on the irregularly
shaped reddish-brown and resinous mass in the centre of the spore. Leading to it or from
it, according to its function, is a complicated irregular system of anastomosing lines of
saprophytic cells which are organized, and orientated in the direction of the main struc-
ture. The resulting pattern is the encirclement either completely or partially of areas of
the surface; nearer the centre of the structure the enclosed areas are irregular but they
tend to become hexagonal or polygonal in shape at some distance away. It is significant
that many of the pale areas from which the exine of the upper surface has been removed
are bounded by cells and filaments marking the furthermost extensions of the structure.

The ‘focus’ of the system appears to be a vesicle or vegetative reproductive centre
from which the attack commenced. The organization of the ensuing attack possibly
bears some resemblance to that already described (PI. 56, fig. 2), but comparison is not
obvious. The meandering nature and quasi-reticulate pattern of this particular organ-
ization is symptomatic of microbiological action upon a uniform structure lacking surface
ornamentation.

Knoxisporites (PI. 56, figs. 5, 6)

The spores included in this genus and represented by the above figures have a very
different appearance. The spore in Plate 56, fig. 5 is red-brown with a thick resinous
exine, while Plate 56, fig. 6 represents a pale-yellow central region with a thin colourless
outer zone, the exine being considerably thinner for the spore as a whole. Despite these
differences, there is a similarity of structure, in that both spores consist of a central body
bearing haptotypic features and surrounded by a cingulum. The latter structure is sub-
divided into an inner thickened annulus concentric with the central body and a thinner
outer flange. This differentiation is most marked in Plate 56, fig. 6.

The attack on the heavier spore (PI. 56, fig. 5) is restricted and simple, consisting of

368

PALAEONTOLOGY, VOLUME 6

coccoid cells and beaded filaments irregularly scattered over the outer flange, but with
a tendency to overall concentric distribution. Locally, the coccoid cells occur in small
concentric groups enclosing small circular areas from which the exine has been partially
removed to provide a pitted surface, illustrated at Plate 56, fig. 5a. Filaments follow
a discontinuous but concentric path nearer the margin of the flange and are best seen
where the exine has been partially destroyed, as at Plate 56, fig. 5b. A much greater degree
of attack is shown by the thinner- walled spore (PI. 56, fig. 6, distal surface), and the
organization follows closely upon the structural elements of the spore in a manner
previously described for spores of a similar structural type, e.g. Lycospora and Denso-
sporites.

Central body. A dense infestation of coccoid cells and filaments characterize this region;
the filaments are curved or branched and, while they produce no distinctive pattern,
they tend to surround circular regions from which the exine is either partially destroyed
or absent as in the light areas (PI. 56, fig. 6).

Cingulum, inner thickened zone. The darker-brown thickened zone forming an annulus
around the central body is heavily attacked and both types of filaments are present as
bundles with an entirely concentric orientation which follows this structural feature of
the spore. At various points branching filaments spread both on to the central body and
outwards over the outer flange. At two points on the lower part of the structure (shown
as dark areas on PI. 56, fig. 6) a dense mass of filaments containing swollen vesicles
results in unusual expansion of the concentric structure, and a similar but smaller aggre-
gate occurs in the opposed segment of the annulus. These masses represent a concen-
tration of saprophytic activity at points along the prolongation of the triradiate rays
and where these cross the annulus.

Cingulum, outer equatorial flange. Very little of the original exine remains and even where
the equatorial margin is complete the exine is much altered. The presumed outer margin
is marked by highly refractive beaded filaments (PI. 56, fig. 6) which are often present

EXPLANATION OF PLATE 56

All figures are of specimens from the Chert Band, Midhope Burn, Upper Oil Shale Group of the

Calciferous Sandstone Series (Scotland). Upper Visean Pi subzone. Magnification X 750 unless other-
wise stated. Slides all labelled M.B.Ch.

Figs. 1-3. Punctatisporites sp. 1, 3/11/820.315, presumed distal surface, figure at slightly higher than
median focus to illustrate reticula. 2, 3/11/782.177, presumed distal surface; a, areas of thin and
altered exine with coccoid cells and filaments; b, area bounded by concentric filaments, exine
removed from the upper surface of the spore; c, area bounded by concentric filaments, exine removed
from upper surface over whole area and from lower surface over part of the area (white), X 1500.
3, 3/5/796.186, proximal surface.

Fig. 4. Microreticidatisporites sp., 3/6/710.213, presumed proximal surface.

Figs. 5-6. Knoxisporites sp. 5, 3/13/802.334, proximal surface; a, small circular pits in the exine;
b, filaments with concentric organization. 6, 1/1/784.190, distal surface.

Fig. 7. Auroraspora sp., 1/4/680.161, proximal surface; a, alignment of attacking cells on the saccus
and near the position of the triradiate mark ; b, dark line marking present line of saccus margin.

Fig. 8. Diatomozonotriletes sp., 4/6/737.142, proximal surface; a, circular area surrounded by a fila-
ment, exine destroyed.

Figs. 9-10. Incertae sedis, presumed '' Palynomorphites pseudomorph’, 9, 3/4/850.183, X1500. 10,
3/4/850.184, X 1500.

Palaeontology, Vol. 6

PLATE 56

mMM

MOORE, Microbiological attack on Carboniferous miospores

L. R. MOORE; MICROBIOLOGICAL ATTACK ON MIOSPORES 369

even where the remainder of the flange has been destroyed. Other filaments arise from
the region of the annulus, cross the equatorial flange radially, and are connected or fused
with the equatorial filaments. A ‘ghost’ framework is thereby produced within which the
exine may either be totally destroyed are partly preserved between the filaments. The
equatorial margin has suffered from some degree of mechanical effects, but this is slight
since the margins of the prominent embayments are lined with coccoid cells and beaded
filaments. Along the lower border the equatorial flange and part of the annulus have
been destroyed, yet the line of the equatorial margin is preserved by a beaded filament in
continuity with a well-preserved portion of the flange.

There is a remarkable similarity in the distribution and organization of this attack
with that described for certain stages of attack on Lycospora and Densosporites, based
as it is upon a similarity of spore structure. No other specimens indicated a more ad-
vanced stage of investment on this type of spore, and it was impossible to complete the
comparison for all stages.

Auroraspora (PI. 56, fig. 7)

The illustration represents the proximal surface of a large spore of this genus in which
the focal plane is coincident with the equatorial margin of the spore. The characteristic
features include a pale-yellow resinous central body carrying prominent trilete sutures.
This body is enclosed by a thin-walled colourless saccus which extends beyond its peri-
phery as a double layer of tissue uniting in the equatorial margin of the spore. The
saprophytic action is directed upon the saccus and also upon the central body.

Saccus. This thin colourless membrane shows evidence of original ornamentation in the
form of closely spaced granules, and the attack upon it is best seen in the outer zone
consisting of the proximal and distal layers of the saccus alone. Coccoid cells, beaded
filaments, and branching filaments are common, the latter two form an irregular network
of straight or curved, branching remains. The complexity suggested by Plate 56, fig. 7 is
in part due to the superimposition of elements occurring on both surfaces of the saccus.
There is a general radial direction in the filament distribution, and many are continuous
from the marginal saccus on to its continuation over the surface of the central body.
This is generally evident but particularly so where the influence of the haptotypic struc-
tures has affected the saccus (PI. 56, fig. la). Over portions of the proximal surface the
saccus has been removed from the central body by microbiological activity. Thus the
greater part of the right-hand segment is free (paler coloured); the present margin of the
saccus is represented by the dark line crossing the lower portion of this segment, which
consists of a concentration of linearly arranged coccoid cells and beaded filaments
marking its edge (PI. 56, fig. lb).

The central body. The most prominent regions of attack are along the two margins of the
triradiate ridges. The investment is principally concerned with the saccus, which is de-
stroyed along such lines, and it extends into the exterior wing as seen along the north-east
triradiate mark. The fact that this trilete mark on the central body is not accompanied by
a double row of filaments follows from the destruction of the saccus along one side of
this line and the retreat of its margin to the further dark line in the right-hand segment
(PI. 56, fig. lb). The central body is only exposed in the right-hand segment; here its
surface carries isolated cells and filaments, but no obvious pattern of distribution has

370

PALAEONTOLOGY, VOLUME 6

been established. While the margin of the central body carries a somewhat denser con-
centric distribution of elements, these are predominantly in the saccus and no unusual
development is present.

The attack on this saccate spore appears to be relatively simple, and at the stage
represented consists largely of the destruction or modihcation of the saccus. The promi-
nent radial organization on this part of the spore may follow an original structure
of the integument, either of ornamentation or radial folding. When compared with
spores such as Tensosporites, which possess an inner thickened cingulum which is followed
by annular concentric filaments, the contrast in the organization of attack on this saccate
spore is well marked.

Diatomozouolriletes (PI. 56, fig. 8)

The figure illustrates the proximal view of a typical spore of this genus characterized
by a thick-walled trilete form with sharply rounded extremities and straight sides, the
latter adorned by strong, pointed, spine-like processes. Attack on the proximal surface is
very restricted and generally limited to the region of the trilete mark. The rays are
attended by thick-walled filaments, with prominent swollen cells, which diverge from
the margins of the ray on to the surrounding exine. The left-hand ray is followed by
a filament which branches and recurves to enclose a small circular area at the radial
extremity of the spore, from which the exine has been removed (PI. 56, fig. 8n). Isolated
clusters of cells are scattered over the surface.

Incertae sedis (PI. 56, figs. 9, 10)

The remains represented by the above figures are common ingredients of the pre-
parations and further illustrate the nature and association of the elements here described
as Palyuomorphites. They consist of that organism preserved in a skeletal network, and
may be other examples of ‘'Palynomorphites pseudomorphs’ but with no indisputable
indication of an original organism which may have been replaced; Plate 56, fig. 10 may
possibly bear some resemblance to the ‘pseudomorph after Lycospora'. These remains
serve to illustrate the advanced degree to which microbiological action may progress
under certain conditions without destruction of the evidence. The figures are of value in
providing a comparative basis upon which the selection of material used in the other
illustrations, indicating stages of attack, may be judged.

CONCLUSIONS

Spores of very divergent characters are invested by the organism Palynomorphites
diversiformis gen. et sp. nov. under suitable environmental conditions. There is an
organization in the nature and distribution of the elements which form the basis of the
colonization or attack; this organization is not uniform but dependent either upon the
major structural features or upon the ornamentation of the spore: the organism is there-
fore often selective in its distribution upon a given spore. The major structural features are
important in controlling or modifying the organization of the attack, but in certain
cases serious modification of a structural feature may result, culminating in its partial
destruction or complete removal. The loss of a central body or flange, or the removal of

L. R. MOORE: MICROBIOLOGICAL ATTACK ON MIOSPORES 371

both structural features, exemplifies this modification as in Densosporites and Lycospora.
One effect of the attack on the structural features results in the production of a 'Palyno-
morphites pseudomorph’ of the spore in which the form of certain structural features
is represented by the organism which has destroyed the original spore.

In the case of spores with few structural features and strong ornamentation, particular-
ly that of rugulae on thick-walled spores such as Convohitispora, the attack is controlled
by the surface ornament. The destruction of irregular or reticulate areas of the exine may
follow and eventually result in the formation of a "Palynomorphites skeleton’ of the spore
as in Convohitispora and Leiotriletes.

Where the spore has no marked structural features, and little or no well-defined surface
ornamentation, as in the case of some forms of Pimctatisporites described in this paper,
there is little selectivity and a greater degree of uniformity in distribution results. The
organization is more diverse, but tends to follow a reticulate or concentric pattern. The
eventual degradation of the spore is followed by its disintegration into segments often
controlled, but not necessarily so, by the haptotypic features (e.g. Lycospora).

A spore may be modified by the addition of Palynomorphites cells or filaments, and
confusion with normal ornament may arise with the varied elements and organization of
attack which Palynomorphites assumes. Thus the formation of radial, reticulate, or con-
centric patterns by filaments, and the absorption and pitting of the exine may severely
affect the appearance of the spore.

While the attack on miospores by the agent, Palynomorphites diversiformis, has been
established, it is not possible to provide reliable evidence as to whether this attack was
preferential on certain groups of spores. All spores present show some degree of attack,
and even within the same group there is wide diversity in the intensity of attack. The
most commonly attacked spores are Lycospora, Densosporites, and Schnlzospora in this
order, the first two groups being most regularly and most seriously affected. Some sub-
stantiation for this statement follows from the larger number of recognizable fragments
of these spores in the preparations. It is of interest to record that microbiological attack
has largely destroyed wood cells; only algae of the Botryococcus type are less affected
than the miospores.

The severity of attack on the miospores described may be regarded as unusual and
possibly due to special circumstances of the environment. As described, the environ-
ment was abnormal, but similar attack by the same organism on spores isolated from
other oil shales and organically rich sediments, has been noted by the writer. This has
been particularly well seen where isolation by hydrofluoric acid alone has been used, or
where calcareous sediments have been digested with hydrochloric acid and the organic
material separated.

In conclusion, Palynomorphites diversiformis is not the only organism which attacks
fossil spores; reference to Plate 54, fig. 1 indicates a further organism investing Sclmlzo-
spora, and other presumed saprophytic organisms of a type similar to Polymorphyces
(Moore 1963).

Acknowledgements. The author is indebted to the British Petroleum Company for financial assistance
in support of the work on the Scottish Oil Shales and related studies. To my colleagues, Dr. Neves
and Dr. Sullivan, I am grateful for their generous assistance in the preparation of this paper and for
relevant discussion. Mr. Bryant, Chief Technician in the Department of Geology, has provided the
photographic illustrations, which the author acknowledges with gratitude.

372

PALAEONTOLOGY, VOLUME 6

REFERENCES

BAILEY, I. w. and VESTAL, M. R. 1937. The significance of certain wood-destroying fungi in the study
of the enzymatic hydrolysis of cellulose. Journ. Arnold Arboretum, 18, 196-205.

BARGHOORN, E. s. 1952. Degradation of plant tissues in organic sediments. J. Sed. Pet. 22, 34-41.
CURRIE, E. G. 1954. Scottish Carboniferous goniatites. Trans. Roy. Soc. Edinb. 62, 527-602.
DRECHSLER, c. 1919. Morphology of the genus Actinomyces. Bot. Gaz. 67, 147-64.

GOLDSTEIN, s. 1960. Degradation of pollen by Phycomycetes. Ecology, 41, 543-5.

HENRici, A. 1947. Molds, Yeasts and Actinomycetes. Wiley, New York, 2nd ed.

KENNEDY, w. Q. 1943. The Oil Shales of the Lothians — structure. Area IV : Philipstoun. Geol. Surv.
Scot. Wartime Pamphlet. 27.

KiDSTON, R. and lang, w. h. 1921. On Old Red Sandstone plants showing structure, from the Rhynie
Chert bed, Aberdeenshire. Pt. V. The Thallophyta. Trans. Roy. Soc. Edinb. 52, 831-55.
KOLLiKER, A. w. 1859. On the frequent occurrence of vegetable parasites in the hard structures of
animals. Proc. Roy. Soc. 10, 95-100.

MESCHiNELLi, A. 1902. Eimgorum Eossilimn Omnium, Iconographia. Typis Aloysii Fabris Co. Vice-
tiae, 1-144.

MOORE, L. R. 1 963. On some micro-organisms associated with the scorpion Gigantoscorpio willsi Stormer.

Vid. Akad. Oslo, M.N.Kl. Skr., n.s., 9, 1-14.

REESE, E. T. 1955. Enzymatic hydrolysis of cellulose. Appl. Microbiol. 4, 37^5.

1959. Cellulose Decomposition: Fungi. Friday Harbour Symposia, Marine Boring and Fouling

Organisms. Univ. Washington Press, Seattle, 265-90.
reinsch, p. f. 1884. Micro-Palaeo-Phytologia, Formationis Carboniferae, vols. i-ii. Theo. Kirsche,
Erlangae, Germania.

RENAULT, B. 1896. Recherches SUE les Bacteriacees Fossiles. Ann. Sci. Nat. 8"^ ser. Bot. 215.

1900. Sur quelques microorganisms des combustibles fossiles. Bull. Soc. Ind. Min. 14, 1-460.

SANGSTER, A. G. and DALE, H. M. 1961. A preliminary study of diflferential pollen grain preservation.
Canad. J. Bot. 39, 35-43.

SAVORY, J. G. 1954. Breakdown of timber by Ascomycetes and Fungi Imperfect!. Annals Appl. Biol.
41, 336-47.

siu, R. G. H. and reese, e. t. 1953. Decomposition of cellulose by micro-organisms. Bot. Rev. 19,
377-416.

SOCIETY OF AMERICAN BACTERIOLOGISTS, 1948. Bergey's Manual of Determinative Bacteriology, 6th ed.
Williams & Wilkins, Baltimore.

TYLER, s. A. and BARGHOORN, E. s. 1954. Pre-Cambrian fossils. Science, 119, 606-8.

VAN TiEGHEM, p. 1 887. Sur les Bacillus amylobacter et son role dans le putrefaction des tissues vegetaux.
Bull. Soc. Bot. 24, 128.

1 879. Sur le ferment butrique {Bacillus amylobacter) a I'epoque de la houille. Compt. Rend. Acad.

Sci., Paris, 89, 1102.

WAKSMAN, s. A. 1950. The Actinomycetes, their Nature, Occurrence, Activities and Importance.
Waltham, Massachusetts.

WEDL, c. 1858. tiber die Bedeutung der in den Schalen von manchen Acephalen und Gasteropoden
vorkommenden Canale. Sitzimgsber. d. Kais. Akad. d. Wiss. in Wien, 33, 451.

L. R. MOORE

Department of Geology,
University of Sheffield

Manuscript received 21 May 1962

LATE JURASSIC MAMMALIAN FOSSILS IN THE
SEDGWICK MUSEUM, CAMBRIDGE

by WILLIAM A. CLEMENS

Abstract. The collection of the Sedgwick Museum contains three mammalian fossils found in the Late Jurassic
Purbeck Beds: dentaries of Trioracodon ferox and Spalacotherium triciispidens and a fragment of a skull of
Peralestes longirostris. The dentary of Trioracodon was discovered in 1933. The other two fossils, part of the
Brodie collection, demonstrate that the postcanine dentitions of Spalacotherium and Peralestes consist of three
premolars and seven molars. Phascolestes dubius is shown to be a junior synonym of Peralestes longirostris.

The collection of the Sedgwick Museum contains three mammalian fossils from the
Purbeck Beds of Durlston Bay, Dorset, that have escaped notice in recent studies of
Mesozoic mammals. The whereabouts of two of these fossils, which were discovered in
the Mammal bed by W. R. Brodie and first described by Richard Owen (1854), was
unknown to Simpson when he prepared his monograph of British Mesozoic mammals
and they were listed as lost (Simpson 1928, pp. 100 and 143). Dr. F. R. Farrington began
a report on these fossils, but the pressure of other commitments has prevented him from
completing it. In the course of my study of British Mesozoic mammals, I had the oppor-
tunity to see these fossils, and Dr. Farrington graciously suggested that I should publish
this note communicating their whereabouts and describing certain aspects of their mor-
phology pertinent to other research problems.

Fossils identified by catalogue numbers prefixed with the letter ‘J’ are part of the collec-
tion of the Sedgwick Museum, Cambridge. Catalogue numbers lacking this prefix
pertain to fossils in the collection of the Palaeontology Department, British Museum
(Natural History).

Order triconodonta Osborn 1888
Family triconodontidae Marsh 1887
Subfamily triconodontinae Hay 1902
Genus trioracodon Simpson 1928
Trioracodon ferox (Owen 1871)

Description. Sedgwick Museum No. J13141 is a fragment of a right dentary containing
the canine and all the postcanine dentition. Most of the dentary anterior to P^ has been
broken away and the bulk of the large, curved canine root is exposed. The root extends
beneath P^, and there is a shallow, longitudinal groove along its lingual side. Unfortu-
nately the premolars and molars were broken and the fragments displaced during fossili-
zation. In the initial preparation of the fossil the crown of P3 came free. It was replaced
in a reversed position, thus exposing the labial surface of the crown, which has a narrow
basal cingulum. Specimens of Trioracodon in the collection of the British Museum
(Natural History), in which the labial side of the lower dentition is exposed, demonstrate
that small but distinct labial cingula are present on at least the P3 and P4 ; these cingula
have been overlooked by some students of the triconodonts. The teeth preserved in
J13141 resemble those found in other mandibles of Trioracodon ferox and do not warrant

[Palaeontology, Vol. 6, Part 2, 1963, pp. 373-7.]

374

PALAEONTOLOGY, VOLUME 6

additional comment. Ventral to Mg and the anterior end of the coronoid process there is
a short, internal groove. Although of approximately the same length, the groove is more
dorsal and posterior in position than the internal groove on the dentary of the type
specimen of Trioracodon ferox, No. 47775.

Comments. This fossil was found in 1933 by D. A. Curry. The following locality data
were obtained from him by Dr. C. L. Forbes. The fossil was discovered in Pur beck beds
cropping out in Durlston Bay approximately half-way between Peveril Point and Durl-
ston Head. It came from a band of blue-grey marl between 3 and 6 inches thick, and
softer than the adjacent strata.

Order symmetrodonta Simpson 1925
Family spalacotheriidae Marsh 1887
Genus spalacotherium Owen 1854
Spalacotheriiim tricuspidens Owen 1854

Description. Sedgwick Museum No. Ill 378 is a fragment of a right dentary containing
an incisor, the alveoli of the ultimate incisor and the canine, and the postcanine dentition.

TEXT-FIG. 1. Spalacotherium tricuspidens Owen, reconstruction of part of the lower dentition based on
a fragment of a right dentary containing the penultimate incisor, alveolus of the ultimate incisor,
alveoli and part of the anterior root of the canine, and the postcanine dentition; J11378; x6-2.

Lingual view, broken surfaces blackened.

Since 1854, when this fossil was described and illustrated by Owen (1854, fig. 10), it has
suffered considerable damage especially to its molars. The preserved incisor is a small,
single-rooted tooth. The posterolingual side of its crown is concave. The penultimate
incisor, P^, and Pg, as well as the alveoli for the ultimate incisor and the two-rooted
canine, are preserved in a fragment of the dentary that does not appear to be distorted.
Posterior to Pg the dentary has been broken and the adjacent part displaced. The crown
of Pg was also broken and rotated out of position. In the reconstruction (text-fig. 1),
Pg has been returned to its proper orientation, but gaps have been left between Pg, Pg,
and Ml to indicate that the fossil is broken and distorted in this region. Other fossils
show that in their natural position these teeth were more closely approximated. The
dental arcade of J11378 also may have been distorted by a fracture of the dentary
beneath Mg. The morphology of the postcanine dentition of Spalacotherium tricuspidens
has been fully described by Simpson (1928, pp. 100-1).

Dimensions. Measurements in millimetres of the teeth preserved in No. J11378.

Pi

Ps

Ml

Mg

Mg

Ml

Ms

Me

M,

Length

1-5

1-7

1-7*

1-6

1-7

1-8

1-8

1-7

1-6*

1-4

Width

0-6

0-8

0-9*

1-0

1-3*

1-4*

1-5*

1-5*

1-5

1-2

* Estimated minimal value

W. A. CLEMENS: LATE JURASSIC MAMMALIAN FOSSILS 375

Comments. This fossil is the only specimen of Spalacotherium tricuspidens containing
a complete incisor and the entire, albeit fragmentary, postcanine dentition. With the
exception of the number of incisors, it gives final confirmation to the dental formula
adopted by Simpson (1928, p. 100), l3+-Ci-P3-M7.

Peralestes longirostris Owen

1871 Peralestes longirostris'. Owen, p. 33.

1871 Phascoiestes cliibins', Owen, legend plate I, fig. 40.

Description. Sedgwick Museum No. J11379 is a heavily damaged fragment of a skull
containing an incisor, the canine, and the postcanine dentition. Most of the alveolar

TEXT-FIG. 2. Peralestes longirostris Owen, fragment of the left side of a skull containing an incisor,
the canine and the postcanine dentition; J11379. a. Outline of the skull fragment, x3-3. b. Occlusal
view of X 20. c. Labial view of the incisor, canine, and P^-P®, reconstruction of alveolar

margin shown by a dashed line, X 10. In figs, lb and 2c the broken surfaces of the teeth are stippled.

region of the maxilla and probably part of the premaxilla are preserved. The bone (text-
fig. 2a) has been severely crushed and distorted and the suture delimiting the premaxilla
and maxilla obliterated. The preserved incisor, now lacking the apex of its crown, has
small anterior and posterior accessory cusps. Its root is long and has a longitudinal

376

PALAEONTOLOGY, VOLUME 6

groove near the middle of the labial surface. Because of damage to the specimen it cannot
be determined if a diastema separated the canine and the ultimate incisor. Most of the
canine has been lost, but an indistinct impression in the matrix and Owen’s illustrations
show that the canine was long and lanceolate. (text-fig. 2c), which is directly behind the
canine, is separated from by a diastema approximately 1 millimetre long. The alveolar
edge of the maxilla between these premolars does not appear to be distorted. A smaller
gap now separates P“ and P^, but the intervening bone is broken and these premolars
have been displaced. Probably they were more closely approximated prior to the frag-
mentation of the specimen. P’ has a simple, trenchant crown and lacks accessory cusps
and a labial cingulum. The larger P^ has small anterior and posterior accessory cusps
and a short labial, basal cingulum on the posterior end of the crown. P^ has a large
posterior accessory cusp and a distinct, crenulated labial cingulum. The lengths of the
premolars are as follows: P^ = TO mm., P“= T4 mm., and P^ = T6 mm.

The molars are heavily damaged. Not only were they broken and distorted during
fossilization, but many fragments were lost, apparently in the first attempts to prepare
the fossil. The crowns of the molars are dominated by the large paracone (nomenclature
following Patterson 1956). Ridges along the margins of the crown connect the paracone
with the stylar region. Cusps were present on these ridges, but have been damaged or
destroyed on most of the teeth. Enough remains to indicate that at least one cusp was
present on the anterior, and another on the posterior, side of the crown of most molars.
These cusps appear to have been as distinct as, but possibly smaller than, those found on
the type of Peralestes longirostris. The ridges along the anterior and labial edges of the
crown terminate at a large stylar cusp. Although of different sizes and somewhat vari-
able in position the other stylar cusps are smaller than the anterior cusp. A small but
prominent posterior stylar cusp is present on the labial ridge of most molars. As far as
can be determined, the molars preserved in J11379 are of approximately the same size
and closely resemble the molars preserved in the type specimen of P. longirostris, no.
All At). Recently the type specimen was more fully prepared and cleaned, revealing the
alveoli of M'^ containing fragments of the tooth. Thus the only emendation to Simpson’s
(1928, pp. 105-6) description of the molars of Peralestes required by 111 379 is the
addition of a description of the seventh molar. M'^ (text-fig. lb), like the preceding molars, is
heavily damaged. Its crown appears to have resembled that of M®, differing in its smaller
size, the shallower medial notch on the labial edge of the crown, and, possibly, in the
relatively smaller size or absence of cusps along the anterior and posterior edges of the
crown.

Comments. Simpson (1928, pp. 102-4) has reviewed the evidence for the allocation of
Peralestes to the Spalacotheriidae. He seconded the suggestion that the genus Peralestes
might be based on the upper dentition of Spalacotheriwn tricuspidens. But Simpson
commented (ibid., pp. 103-4): Tt is not to be definitely considered as a synonym of
Spalacotherium, however, until their association can be considered proven beyond any
question.’ Since publication of Simpson’s monograph the upper and lower molars of
an Albian symmetrodont, Spalacotheroides bridwelli, have been described by Patterson
(1955, 1956). Information obtained from these fossils supports the allocation of Pera-
lestes to the Spalacotheriidae, but does not bear directly on the problem of the synonymy
of the Purbeck genera. Now it can be demonstrated that the postcanine dental formulae

W. A. CLEMENS: LATE JURASSIC MAMMALIAN FOSSILS

377

of Peralestes and Spalacotherium are equivalent and that the posterior molars of both,
My, are distinctly smaller than the penultimate molars. This adds support to the sugges-
tion that Peralestes longirostris is a junior synonym of Spalaeotherium tricuspidens, but
it does not prove their association beyond any question.

Recognition of the whereabouts of J 11379 requires one minor change in the currently
accepted classification. This fossil appears to be the specimen originally identified by
Owen (1854, p. 430) as a fragment of the right dentary of Spalacotherium tricuspidens.
The illustration of the specimen (ibid., fig. 12) is not detailed, but the identification can
be based on the gross similarities between J 11379 and the illustration. Also, of the fossils
listed by Owen (ibid., p. 431) as associated with the mammalian fossil, only the ‘. . . jaw
with a few teeth . . .’ and ‘. . . the impression of the dentary bone . . .’ of Macellodus are
missing from the block in which J11379 is preserved. In the text of his monograph on
Mesozoic mammals Owen (1871, pp. 36-37) identified the mammalian fossil as "Incertae
sedis, Peralestes sp. ?’, but in the legend of Plate I he allocated it to a new species of
Phascolestes, P. dubius. Because Owen’s description and illustrations were inadequate,
the whereabouts of the type specimen was unknown, and the referred specimen was
indeterminate, Simpson (1928, p. 143) deemed the species Phascolestes dubius to be a
nomen nudum. Now that the type specimen has been found, P. dubius can be demonstrated
to be a junior synonym of Peralestes longirostris.

Acknowledgements. I am indebted to Mr. A. G. Brighton and Dr. F. R. Farrington, who made it
possible for me to study these Mesozoic mammalian fossils. Thanks are due to Drs. E. I. White and
A. Sutcliffe, who gave me access to the collection of Mesozoic mammals in the British Museum (Natural
History) and extensive assistance with many phases of my research. I am indebted also to Drs. C. L.
Forbes, K. A. Joysey, and K. A. Kermack, and Mrs. Frances Mussett for the help they gave me. It
is a pleasure to cite the craftsmanship of Mr. A. Rixon, who skilfully prepared the type specimen of
Peralestes longirostris. This research was carried out while I was a National Science Foundation Post-
doctoral Fellow at University College, London, and the report was completed at The University of
Kansas.

REFERENCES

OWEN, RICHARD. 1854. On some fossil reptilian and mammalian remains from the Purbecks. Quart.
J. geol. Soc. London, 10, 420-33.

1871. Monograph of the fossil mammalia of the Mesozoic formations. Paleontogr. Soc. 24,

1-115, pi. i-iv.

PATTERSON, B. 1955. A symmctrodont from the Early Cretaceous of northern Tcxas. Fieldiana: Zool.
37, 689-93.

■ 1956. Early Cretaceous mammals and the evolution of mammalian molar teeth. Fieldiana: Geol.

13, 1-105.

SIMPSON, G. G. 1928. A catalogue of the Mesozoic Mammalia in the Geological Department of the
British Museum. London, British Museum (Nat. Hist.), x+215, pi. i-xii.

WILLIAM A. CLEMENS

Museum of Natural History
and Department of Zoology,
The University of Kansas,
Lawrence, Kansas

Manuscript received 27 September 1962

CATALOGUE OF DESCRIBED AND FIGURED
SPECIMENS IN THE BEGG COLLECTION IN
THE HUNTERIAN MUSEUM OF THE
UNIVERSITY OF GLASGOW

by ETHEL D. CURRIE and T. NEVILLE GEORGE

John L. Begg (1874—1958) of Glasgow was a distinguished amateur palaeontologist
who for many years collected fossils notably from the Ordovician and Silurian rocks of
southern Scotland. His specimens, many of them exceptionally well preserved, have been
used by Begg himself and by others in palaeontological and stratigraphical studies that
have enriched Lower Palaeozoic geology. Most of them were presented to, or bought
by, the Hunterian Museum of the University of Glasgow, where they form the Begg
Collection.

The collection is especially rich in fossils of the Bala and Valentian series of the Girvan
district, represented by several hundred trilobites and as many protrematous brachio-
pods, about 100 asteroids, 70 gastropods, 60 lamellibranchs, 60 cystids, and smaller
numbers of inarticulate brachiopods, nautiloids, and crinoids. The local fossils are
supplemented by about 200 other specimens, most of which Begg got by exchange, from
comparable horizons in Scandinavia, Canada, and the U.S.A.; and by about 60 extra-
British Cambrian trilobites. A group of Carboniferous fossils includes about 100 lamelli-
branchs (mainly Scottish), about 80 crinoids, about 70 gastropods (many from the Isle
of Man), and 30 nautiloids; but Carboniferous corals, brachiopods, goniatites, and
polyzoans are very poorly represented. Mesozoic and Tertiary fossils are mostly lacking,
but there is a heterogeneous group of about 50 ammonites, and a few gastropods from
the Pleistocene clays of the Clyde estuary.

This catalogue lists all the described and figured specimens in the Begg Collection.
References are given to the original figure or description of each specimen, and to any
subsequent renaming or redescription. Where possible, references are also given to the
use of several specimens or of supplementary specimens in description, and comment is
made on the manner of preservation of the specimens.

Specimens ‘lost’ or unidentifiable are omitted from reference; a number of type-
specimens that should have come to the Museum, and that may be recorded elsewhere
as being in the Museum collections, were never received from Mr. Begg, and, although
it is likely that he gave or lent them to other workers or deposited them in other insti-
tutions, no record of them is now to be found in Glasgow.

Specimens are listed under Hunterian Museum numbers, but Mr. Begg’s private
numbers (prefaced by BG) are also included in round brackets where the specimens
carry them. An explanation of Mr. Begg’s Starfish Bed horizons may be found in Begg
1947, p. 29.

[Palaeontology, Vol. 6, Part 2, 1963, pp. 378-96.]

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION

379

TRILOBITA

Ordovician

Acidaspis (Ceratocephala) discreta Reed

Internal mould, A1030« (BG2102), and external mould, A 1030/? (BG2103), of an almost complete
specimen.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935r/, p. 38, pi. 1, figs. 22, 22o, h (A1030n), pi. 4, fig. 10 (A1030/?). Whittington 1956, p. 515,
pi. 59, fig. 13; pi. 60, fig. 10(A10306). Whittington placed the species in synonymy with Eroccra/o-
cephala tern'bilis (Reed), to which he referred Begg’s specimen.

Acidaspis {Miraspis) solitaria Reed

Holotype, a small cranidium, A939 (BG3142).

Caradoc, Stinchar Limestone Group, Didymograptus superstes Shales; Aldons, near Girvan.

Reed 1935fl, p. 37. pi. 3, fig. 21.

Ainpyx (Ampyxina) aldone/isis Reed

Syntypes, ‘a few head-shields’, only two of which are specified by number, A919 (BG2124) and
A920 (BG3101). There is also a pygidium, A1126 (BG2141), which according to Reed may be
associated with them.

Caradoc, Stinchar Limestone Group, Didymograptus superstes Shales; Aldons, near Girvan.

Reed 1935«, p. 7, pi. l,fig. 23 (A919). Whittington 1959, p. 487, pi. 28, fig. 10(A1126), figs. 11, 13
(A919).

Ampyx carrickensis Begg

Holotype, A3690, a small cephalon.

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Begg 1947, p. 42, pi. 3, figs. 7, 8.

Asaphus (Parabasilicus) cf. powisi Murchison

Part of a thorax comprising axial lobe and pleura of seven segments, A934 (BG2261).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935i7, p. 12, pi. 2, fig. 13.

Astroproetus reedi Begg

Holotype, A 1082 ( BG6451), a complete specimen.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 1939, p. 375, pi. 6, fig. 2.

Astroproetus whittardi Begg

Holotype, A1083 (BG7707), a complete specimen.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1939, p. 376, pi. 6, fig. 3.

Calymene (Colpocoryphe?) aldouensis Reed
Syntypes, two imperfect cranidia, A1037 {BG2I1 1) and A1038 (BG2110).

Caradoc, Stinchar Limestone Group, Didymograptus superstes Shales; Aldons, near Girvan.

Reed 1935n, p. 48, pi. 1, fig. 14 (A1037), fig. 15 (A1038). Shirley 1936, p. 418.

Calymene (Diacalymeiie) bigener Reed

The description is based on one nearly perfect cephalon with attached thorax, A 1098 (BG929).
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935^, p. 47, pi. 1, fig. 3. Shirley 1936, p. 418.

Clypoproetus asteroideus Begg
Holotype, A1080 (BG7713), a complete specimen.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1939, p. 374, pi. 6, fig. 1.

c c

c n;i3

380

PALAEONTOLOGY, VOLUME 6

Cryptolithiis nicholsoni Reed

The specimen, A3623 (BG142), shows the lower lamella of the fringe.

Ashgill, Pusgill Beck, Westmorland.

Begg 1944, pi. 5, fig. 1.

Cyhele (Cybeloides) loveni Linnarsson var. girvaneiisis Reed
A specimen split to separate the glabella with hypostome attached beneath, A737, A131a, and
A737Z)(BG8217).

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1943n, p. 61, pi. 2, figs. 2, 2a (A737).

Cyhele (Cybeloides) loveni Linnarsson var. radiata Reed

Holotype of the variety, a specimen consisting of a pair of free cheeks displaced from their correct
relative position, together with some broken fragments of the thorax, A 1054 ( BG259).

Ashgill, Upper Drummuck Group, Ladyburn mudstone. Zone of Dicellograptus anceps\ South
Threave, near Girvan.

Reed 1930, p. 195, pi. 9, figs. 2, la-b.

Cybele (Cybelina) monoceros Reed

The description of this species is based on a cranidium, A1067 (BG252). Two small pygidia, A1068
( BG254) and A1069 ( BG253), are said to be probably referable to the same species.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1931n, p. 101, pi. 5, fig. 1 (A1067), fig. 2 (A1068), fig. 3 (A1069). Reed 19316, p. 26.

Cybele (Cybele) nicholsoni Reed

This species is apparently based on a cranidium, A1042 (BG2260).

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Reed 1935a, p. 52, pi. 4, fig. 4.

An isolated right free cheek, A 1043 (BG2273), from the same locality, probably belongs to the same
species (Reed 1935a, p. 53, pi. 2, fig. 14).

Cybele nicholsoin Reed

A hypostome, counterpart moulds, A4127a (BG 1 1903) and A41276(BG11904), is referred tentatively
to this species.

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Begg 1951, p. 367, pi. 1, fig. 14 (A4127a).

Cybele (Cybele?) perversa Reed

Holotype, a small incomplete cranidium, AllOO (BG1052).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1935a, p. 53, pi. 4, fig. 5.

A specimen referred doubtfully to this species (ibid., p. 54) cannot be found in the Begg Collection.

Encrinurus midtisegmentatus (Portlock) var. girvanensis Reed
This variety ‘is represented by one perfect individual [A1040 (BG963)] and by several incomplete
specimens, thoraces, pygidia and head-shields one of which [A1 1 36 (BG960)] shows the hypostome
in position’. Specimens A1040 and A1 136 are labelled, but the others, for which numbers are not
given, cannot be positively identified.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935a, p. 50, pi. 4, fig. 9 (A1040).

A hypostome, A733 (BG7573).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1; Ladyburn, near Girvan.

Begg 1943a, p. 58, pi. 2, fig. 4.

A hypostome, A4128 (BG 11905).

Same horizon and locality.

Begg 19506, p. 367, pi. 1, fig. 15.

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION

381

Encriniirus sublanceolatus Reed

Holotype, a pygidium, A1093 (BG2265).

Caradoc, Stinchar Limestone Group; Glenlochrie, near Girvan.

Reed 1935n, p. 49, pi. 4, fig. 8.

Encriminis aff. trispinosiis Reed

A large cranidium, A4099 (BG3368).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; east brow of Quarrel Hill, New Dailly,
near Girvan.

Tripp 1957, p. 68, pi. 11, fig. 14.

Flexicalymene cf. meeki (Foerste)

A cranidium, A1096 ( BGlOl 1).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; east brow of Quarrel Hill, New Dailly,
near Girvan.

Shirley 1936, p. 404, pi. 29, fig. 8.

Ulaemis balclatchie/isis Reed var. cristata Reed

Holotype of the variety, a cranidium, A935 (BG25).

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Reed 1935n, p. 24, pi. 3, fig. 2.

Ulaemis dnnnmiickensis Reed

Holotype, A3687 (BG6454), a complete pygidium; paratype, A3688 (BG7740), the posterior half
of another.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed in Begg and Reed 1945, p. 260, pi. 1, fig. 3 (A3687), fig. 4 (A3688).

Ulaemis longicapitatiis Reed

A cephalon with the epistome in place, split to expose the hypostome in situ, counterparts, A734
(BG7669«) and A734n (BG7669).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 1943n, p. 59, pi. 2, figs. 1, In (A734).

Ulaemis oblongatiis (Angelin) var. exstans Reed

Holotype of the variety, a small pygidium, A1108 (BG9478).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1941, p. 270, pi. 5, fig. 7.

Ulaemis roemeri Volborth var. transversalis Reed

Holotype of the variety, a pygidium, A1053 (BG290).

Ashgill, Upper Drummuck Group, Ladyburn mudstones, Zone of Dicellograptiis ancepsi South
Threave, near Girvan.

Reed 1930, p. 194, pi. 9, figs. 1, \a, b.

Ulaemis siiperstes Reed

Syntypes, a cephalon and pygidium of the same individual with the thorax crushed between them,
A3630 (BG9685); a cephalon lacking the right free cheek, A3631 (BG9686); a pygidium with
thorax attached, A3632 ( BG9653) ; a pygidium, A3633 (BG9687) ; and a cephalon, A3646 ( BG9680),
which is not figured (p. 60).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1944n, p. 59, pi. 2, figs. 3, 3a (A3630), figs. 4, 4a (A3631), fig. 5 (A3632), fig. 6 (A3633).

Lichas (Metopolichas?) congruens Reed

Holotype, a well-preserved cranidium, A937 (BG2084).

Ashgill, Upper Drummuck Group, Ladyburn mudstones. Zone of Dicellograptiis anceps; South
Threave, near Girvan.

Reed 1935o, p. 34, pi. 3, fig. 10.

382

PALAEONTOLOGY, VOLUME 6

Lidias (Platylichas) crescenticus Reed

The fragments (BG2080 and BG2082) on which Reed based his description are counterparts ; the
holotype thus consists of part of a cranidium with hypostome in place, A915 (BG2082), an
external mould of the complete cephalon, A915c (BG2082), and an external mould of the ventral
surface and hypostome, A915Z) (BG2080).

Ash gill. Upper Drummuck Group, Ladyburn mudstones. Zone of Dicellograptus anceps; South
Threave, near Girvan.

Reed 1935n, p. 29, pi. 3, figs. 13 and 15 (A915), fig. 14 (A915^>), fig. 16 (A915u). Phleger 1937, p. 254
(BG2080<7 misquoted for BG2082). Begg 1943Z), p. 160.

Lidias (Hemiarges) maccullodii Reed

A hypostome in situ, exposed in dorsal view beneath the cephalon which has been partly removed,
A730 (BG5404).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 1943n, p. 56, pi. 2, fig. 6.

Reference is made (p. 56) to four specimens in which ornament is well shown, A743 (BG3305),
A744n (BG3307fl), A745 (BG3219), A746 (BG7561).

Lidias (Acrolichas) tenuilobatiis Reed

A small cranidium, A3647 (BG9783).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1944n, p. 61, pi. 2, fig. 8. Tripp 1958, p. 580, pi. 85, fig. 12.

Lidias (Platylichas) thmivensis Reed

Holotype, a pygidium, A916 (BG2087), and its external mould, A916o (BG2081).

Ashgill, Upper Drummuck Group, Ladyburn mudstones. Zone of Dicellograptus anceps South
Threave, near Girvan.

Reed 1935a, p. 32, pi. 3, fig. 17 (A916), fig. 18 (A916a).

A hypostome, A917 (BG2090), is referred to as possibly belonging to this species (Reed 1935a, p. 33,
pi. 3, fig. 19).

Lidias (Platylichas) vicinus Reed

Holotype, a crushed pygidium with parts of a few thoracic segments attached, A918 (BG2083).

Ashgill, Upper Drummuck Group, Ladyburn mudstones, Zone of Dicellograptus anceps; South
Threave, near Girvan.

Reed 1935a, p. 33, pi. 3, fig. 12.

Otarion elongatus Begg

Holotype, A1089 (BG9325), a nearly complete specimen; paratype, A1090 (BG6453), an external
mould of a small, almost complete specimen.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1939, p. 380, pi. 6, fig. 8 (A1089), fig. 7 (A1090).

Otarion (= Cyphaspis) cf. minima (Cooper)

A small, incomplete cranidium, A1034 (BG1017).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1935a, p. 44, pi. 1, fig. 17.

Paraharpes hornei (Reed)

An almost complete specimen, E4472 (BG759) and E4473 (BG760), counterparts; and part of a
cephalon with fringe, A4475 (BG2362).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.

Whittington 1950, p. 39, pi. 4, fig. 4 (A4473), fig. 6 (A4475). The specimen (BG803) of figs. 1 and
3 cannot be found in the Begg Collection.

Phacops (Monorakos) asteroideus Reed

A complete specimen with pygidium displaced, A1058 (BG594), and a cephalon with part of the
thorax attached, A1059 (BG595).

E. D. CURRIE AND T. N. GEORGE; THE BEGG COLLECTION 383

Ashgill, Upper Drummuck Group, Ladyburn mudstones, Zone of Dicellograptus anceps; South
Threave, near Girvan.

Reed 1930, p. 198, pi. 10, figs. 1, \a, b (A1058), figs. 2, 2a (A1059). Reed 19316, p. 24.

Phacops (Calliops) jiikesi Salter var. viciiia Reed

Holotype, A5370 (BG11768), a nearly complete cephalon.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1945, p. 314, pi. 1, fig. 6.

Phacops {Pterygometopus?) quairelensis Reed
Syntypes, two pygidia, A1056 (BG603), A1057 (BG602).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1930, p. 197, pi. 10, figs. 3, 3a (A1056). Reed 19316, p. 24.

Phacops {Pterygometopus) retardatus Reed

An almost complete specimen, A736 (BG8208a), with part of the cephalon missing so that it presents
an external mould of the hypostome.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1943a, p. 60, pi. 2, fig. 10.

Phillipsinella parabola Barrande

A hypostome in place behind the glabella, in ventral aspect, counterparts, A735 (BG8211) and
A735a (BG8211a).

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1943a, p. 60, pi. 2, fig. 8 (A735).

Platylichas congruens (Reed)

A large individual in which the cephalon and pygidium are crushed and incomplete, (A3967).
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Tripp 1958, p. 579, pi. 85, figs. 7, 8.

Pliomera (Pliomerops) girvauensis Reed
Holotype, part of a cephalon, A1079 (BG445).

Caradoc, Stinchar Limestone Group; Aldons Quarry, near Girvan.

Reed 1930, p. 196, pi. 9, figs. 3, 3a.

Pliomerella serotina Reed

Holotype, part of a cranidium, A1 107 (BG9480), and a fragment of its impression, A1 107a (BG948 1).
Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Reed 1941, p. 268, pi. 5, fig. 1 (A1107), fig. 2 (A1107a).

Proceratocephala terribilis (Reed)

Internal and external moulds, A1030a (BG2102) and A10306 (BG2103), of an almost complete
specimen. The specimen is the holotype of Acidaspis {Ceratocephala) discreta (Reed) (q.v.).
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Whittington 1956, p. 515, pi. 59, fig. 13, pi. 60, fig. 10 (A10306).

Proetiis ardmiUaneusis Begg

Holotype, A3692, part of a small cranidium.

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Begg 1947, p. 42, pi. 3, fig. 3.

Proetus balclatchiensis Begg

Holotype, A4122 (BG11898), an incomplete cephalon and part of the enrolled thorax.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Begg 1951, p. 362, pi. 1, fig. 1.

Proetus fardenensis Begg

Holotype, A3821 (BG11894), a cephalon with part of the thorax.

384

PALAEONTOLOGY, VOLUME 6

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1950, p. 285, pi. 14, hgs. 1, 2.

Proetiis girvanensis Nicholson and Etheridge

Part of a cephalon with hypostome in place beneath the glabella, A731 (BG8341).

Ashgill, Upper Drummuck Group, Starhsh Bed No. 2; Ladyburn, near Girvan.

Begg 1943u, p. 57, pi. 2, hgs. 7, la.

Proetiis mactaggarti Begg

Holotype, A3691 (BG 11871), a complete specimen.

Ashgill, Upper Drummuck Group, Starhsh Bed a few feet above No. 3 ; Ladyburn, near Girvan.
Begg 1947, p. 40, pi. 3, hgs. 1, 2.

Proetiis scobiei Begg

Holotype, A3689 ( BG 11017), a cephalon.

Ashgill, Upper Drummuck Group, Starhsh Bed No. 2; Ladyburn, near Girvan.

Begg, in Begg and Reed 1945, p. 261, pi. 1, hgs. 1, 2.

Proetiis trefoileiini Begg

Holotype, A4123 (BG11899), external mould of an almost complete cephalon.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Begg 1951, p. 364, pi. 1, hg. 2.

Proetiis sp.

A small, incomplete cranidium, counterparts, A4124<7 {BG11900) and A41246 (BG11900u).
Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Begg 1951, p. 365, pi. 1, hg. 3 (A4124u).

Pseudosphaerexochiis graniilatiis (Angelin)

Reed referred to two cranidia (BG2075 and BG2335), but only one of them, A1050 (BG2075), is in
the Hunterian Museum. The other was sent by Begg to the Senckenberg Museum, Frankfurt-
on-Main, there numbered SMF X 929a.

Ashgill, Upper Drummuck Group, Starhsh Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935o, p. 56, pi. 4, hg. 2 (A1050).

Pseudosphaerexochiis siibquadratiis Reed
Part of a cranidium, A1048 (BG748).

Ashgill, Upper Drummuck Group, Starhsh Bed No. 1; Ladyburn, near Girvan.

Reed 1935u, p. 56, pi. 4, hg. 3.

Remopleiirides asteroideiis Reed

Three glabellae, A933 (BG2066), A1127 (BG785), A1128 (BG2341).

Ashgill, Upper Drummuck Group, Starhsh Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935«, p. 11, pi. 4, hg. 6 (A933), hg. 7 (All 27).

Remopleiirides hedstrdmi Reed

Holotype, a cranidium, A3629 (BG9502).

Ashgill, Upper Drummuck Group, Starhsh Bed No. 1 ; Threave Glen (Ladyburn), near Girvan.
Reed 1944u, p. 58, pi. 2, hgs. 2, 2a.

Seleiioharpes youiigi (Reed)

Cephala with fringe, A4468 (BG206) and A4469 (BG205).

Caradoc, Balclatchie Beds; Balclatchie, near Girvan.

Whittington 1950, p. 30, pi. 3, hg. 4 (A4469), hg. 5 (A4468).

Sphaerocoryphe hastata Begg

Holotype, A1095 (BG2151) and A1095a (BG2512), counterparts, part of a cephalon.

Caradoc, Stinchar Limestone Group, Didymograptus superstes Shales ; Aldons Quarry, near Girvan.
Begg 1940fl, p. 302, pi. 4, hg. 8 (A1095).'

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION

385

Sphaerocoryphe thomsoni Reed

Of six specimens referred to, BG7729-31, BG955, BG7727, and BG7728, the first three cannot
be found. The specimens in the Hunterian Museum are a cranidium, All 10 (BG955), and two
specimens on a single slab, A838 (BG7727 and BG7728), in each of which the glabella is removed
and the hypostome exposed.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1940fl, p. 300, pi. 4, fig. 3 (All 10), fig. 6 (A838).

Stygina latifrons (Portlock)

A specimen in which the glabella is removed, exposing an external mould of the hypostome. A732
(BG3308).

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1943fl, p. 58, pi. 2, fig. 5.

Telephiis girvcmensis Reed

Reed implied that the Begg Collection includes more than one example of this species, but only one
specimen, a cranidium, A921 (BG3140), is quoted by number.

Caradoc, Stinchar Limestone Group, Didymogrciptus superstes Shales; Aldons, near Girvan.

Reed 1935n, p. 9, pi. 1, figs. 2, la (A921).

Teratorhynchus sp.

A cephalon and a few thoracic segments, A3823 (BG229).

Caradoc, Balclatchie Group; Dow Hill, near Girvan.

Begg 1950, p. 290, pi. 14, fig. 3.

Tretaspis cerioides (Angelin) var. sortita Reed

Three almost complete specimens, A3625 (BG2251), A3626 (BG24), A3628 (BG23), and two
cephala, A3624 (BG3049), A3627 (BG9515).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 1944, p. 113, pi. 5, fig. 2 (A3624), figs. 3 and 6 (A3625), fig. 4 (A3626), fig. 5 (A3627), fig. 7
(A3628).

Trinucleiis (Cryptolithus) gibbifrons M‘Coy var. praeterita Reed

Reed referred the following specimens to this variety: A930 (BGIOOO). A924 (BG1002), A925
(BG2263), A926 (BG1003), A927 (BG2264), A928 (BG1013), A929 (BG1014), and BG1004. Of
these, BG1004 is missing. The specimens are cephala or parts of cephala, except A928 and
A929, which are pygidia.

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1935n, p. 1, pi. 4, fig. 11 (A924), fig. 12 (A925), fig. 13 (A926), fig. 14 (A927), fig. 15(A928).

Trimideus {Tretaspis) persidcatus Reed

The species appears to be based on one specimen in the John Smith Collection and on several
specimens in the Begg Collection, of which three (BG951, BG964, and BG967) are missing. The
specimens remaining are A1116 (BG968), A1117 (BG974), A1118 (BG975), A1119 (BG980),
A1120 (BG981), and A932 (BG982).

Ashgill, Upper Drummuck, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1935o, p. 6, pi. 4, fig. 16 (A932).

Warburgella rotiindata Begg

Holotype, AI084 (BG9323), and paratype, A1085 (BG6458), both almost complete specimens.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1939, p. 378, pi. 6, figs. 5 and 6 (A1084), fig. 4 (A1085).

Silurian

Acidaspis (Globidaspis) prominens Reed

Holotype, an incomplete cranidium, A1066 (BG735).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 193 lo, p. 100, pi. 5, figs. 5, 5a, b.

386

PALAEONTOLOGY, VOLUME 6

Acidaspis {Miraspis) ultima Reed

Holotype, a small incomplete cranidium, A1065 (BG738).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 193 In, p. 99, pi. 5, figs. 6, 6a.

Cheininis conjimctus Reed

Holotype, a well-preserved cranidium, A1045 (BGl 198), and its external mould, A1045n (BGl 199).
Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1935n, p. 55, pi. 4, fig. 1 (A1045).

Cheimrus elougatiis Reed

The species is based on four syntypes — two incomplete cranidia, A1072 (BG421) and A1073
(BG420), an external mould of a pygidium, A1074 (BG422), and a hypostome, A1075 (BG419).
Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1931n, p. 103, pi. 4, figs. 5, 5a (A1072), fig. 6 (A1073), fig. 7 (A1074); pi. 5, fig. 4 (A1075).
Begg 1943n, p. 62 (A 1075).

Cyphopwetus newlandeusis Begg
Holotype, A3824 (BGl 1895), a small cranidium.

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Begg 1950, p. 287, pi. 14, fig. 7.

lUaemis aemidus Salter
A pygidium, A936 (BG1028).

Middle Valentian, Saugh Hill Group, Upper Pentamerus Bed; Penwhapple Glen, near Girvan.
Reed 1935o, p. 25, pi. 2, fig. 9.

A head-shield (BGl 036), referred to as probably belonging to this species, cannot be found in the
Begg Collection.

Phacops elliptifrous Sars and Boeck

An incomplete hypostome, A742 (BG8715).

? Saugh Hill Group; north of Mulloch Hill, near Girvan.

Begg 1943o, p. 61, pi. 2, fig. 9.

Specimens that show some detail of the ventral ‘interlocking’ furrow of the cephalon, A4141
(BG586), A4142-5.

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Begg 1950, p. 367, pi. 1, figs. 6 and 7 (A4141), fig. 8 (A4142), fig. 9 (A4143), fig. 10 (A4144), fig. 11
(A4145).

Proetiis sp.

External moulds of two pygidia, A4125 (BGl 1901), and A4126 (BGl 1902).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Begg 1950, p. 366, pi. 1, fig. 4 (A4125), fig. 5 (A4126).

Proetus {Cyphopwetus) externus Reed
Holotype, a cranidium, A1032 (BG911).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1935a, p. 42, pi. 2, fig. 15.

Proetus iiiteijectus Reed

Holotype, a small cranidium, A1031 (BG2057).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1935a, p. 40, pi. 3, fig. 23.

Proetus scoticus Reed

Holotype, a cranidium, A1109 (BG9483).

Lower Llandovery, Mulloch Hill Group; Craigens, near Girvan.

Reed 1941, p. 271, pi. 5, fig. 3.

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION 387

Proetus subtriangularis Begg

Holotype, A3822 ( BG11896), a cranidium.

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Begg 1950, p. 287, pi. 14, fig. 6.

Sciitellion (Bronteus) cunctatiim Reed

According to Reed there are three nearly perfect cranidia and two pygidia in the Begg Collection,
which are presumably the syntypes of the species. One of these specimens cannot be found, but
the four figured specimens are available. These are two cranidia, A1061 (BG501) and A1062
(BG503), and two pygidia, A1063 (BG499) and A1064 (BG495).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1931n, p. 97, pi. 4, fig. 1 (A1061), fig. 2 (A1062), fig. 3 (A1063), fig. 4 (A1064).

Carboniferous

Cummingella balladoolensis Reed

The species is represented by three syntypes — two cephala, A3704 (BG8601) and A3705 (BG8601a),
and a glabella, the latter on the same rock-fragment as a pygidium, A3706 (BG8601x).
Carboniferous Limestone, Upper Limestone; Balladoole, Isle of Man.

Reed 1942, p. 668, pi. 10, fig. 1 (A3704), fig. 2 (A3705), figs. 3, 3a (A3706).

Cyphiuioides ashfellensis Reed

Holotype, a complete individual enrolled so that the cephalon is pressed against the pygidium,
A3700 (BG7853).

Lower Carboniferous, Ashfell Limestone; near Kirkby Stephen, Yorkshire.

Reed 1942, p. 654, pi. 8, figs. 1, la, 2.

Eocyphimiim clitheroense Reed
Holotype, a cephalon, A3701 (BG8602).

Lower Carboniferous; Clitheroe, Lancashire.

Reed 1942, p. 656, pi. 9, figs. 1, la.

Eocyphinium? bivium Reed

Holotype, a pygidium in interior view, A3702a (BG9082), and its internal mould, A3702 (BG9089).
Upper Limestone Group, Calmy Limestone; Linn Spout, Dairy, Ayrshire.

Reed 1942, p. 659, pi. 9, figs. 2, 2a, b (A3702).

Griffithides ? ambiguus Reed
Holotype, a pygidium, A3714 (BG7579).

Carboniferous Limestone, Upper Limestone; Balladoole, Isle of Man.

Reed 1942, p. 667, pi. 11, figs. 6, 6a, b.

Weberides dalrieiisis Reed
Holotype, a pygidium, A3707 fBG9080I

Upper Limestone Group, Calmy Limestone; Linn Spout, Dairy, Ayrshire.

Reed 1942, p. 666, pi. 10, figs. 4, 4a,^>.

Weberides mucronatus M‘Coy var. lata Reed
Holotype of the variety, a pygidium, A3713 (BG7851).

Lower Limestone Group; Leslie, Fife.

Reed 1942, p. 663, pi. 11, figs. 5, 5a.

Weberides mucronatus M‘Coy var. traquairi Reed

Syntypes of the variety appear to be a cephalon, A3709 (BM104), and two pygidia, A3710 (BM103)
and A3711 (BM102).

Carboniferous Limestone Series; Garple Burn, Muirkirk, Ayrshire.

Reed 1942, p. 660, pi. 11, fig. 1 (A3709), figs. 2, 2a (A3710) and fig. 3 (A3711).

A pygidium, A3712 (BG7852).

388

PALAEONTOLOGY, VOLUME 6

Lower Limestone Group; Leslie, Fife.

Reed 1942, pi. 11, fig. 4.

Weberides parilis Reed
Holotype, a pygidium, A3708 (BG9094).

Upper Limestone Group, Calmy Limestone; Linn Spout, Dairy.
Reed 1942, p. 664, pi. 10, figs. 5, 5a, b.

Ordovician

BRACHIOPODA

Aegiria? baiclatchiensis (Reed)

An internal mould of a ventral valve, L2795 (BG9477).
Caradoc, Balclatchie Group; Balclatchie, near Girvan.
Reed 1941, p. 273, pi. 5, fig. 6.

Bimuria pernigata Reed

Holotype, L3191 (BG11754), an external mould of a dorsal valve with the hinge-area of the ventral
valve; paratypes, two external moulds of dorsal valves, L3192 (BG11874) and L3193 (BG 11729).
Caradoc, Balclatchie Group; Balclatchie Bridge, near Girvan.

Reed 1945, pp. 313-14, pi. 1, fig. 1 (L3191), fig. 2 (L3192), fig. 3 (L3193).

Chonetes (Eodevonaria) celtica Reed

Holotype, L3315 (BG9748), a dorsal valve with the hinge-areas of both valves.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed, 1944c, p. 217, pi. 3, figs. 2, 2a.

Coelospira? insoleta Reed

Holotype, an internal mould of a complete shell, L2805 (BGS75).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Threave Glen (Ladyburn), near Girvan.
Reed 19356, p. 8, pi. 1, figs. 2, 2a-d.

Lingula girvanensis Begg

Holotype, L2992, a ventral valve.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1947, p. 32, pi. 2, fig. 2.

Lingula ? isosceles Begg
Holotype, L2990.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Begg 1947, p. 36, pi. 2, figs. 7, la.

In a letter (27 June 1949), Begg stated: ‘I have now become convinced that this is a peculiarly pre-
served example of the Hyolithes present in the Balclatchie Beds.’

Lingula pustulosa Begg

Holotype, L2994, a ventral valve.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1947, p. 31, pi. 2, fig. 1.

Lingulasma ? concatenata Begg

Holotype, L2991, a ventral valve; paratype, L2993, an internal mould of a ventral valve with part
of the shell in place.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1947, p. 35, pi. 2, figs. 5, 5a (L2991), fig. 6 (L2993).

Lingulasma ? fardenensis Begg

Holotype, L2996, a ventral valve; paratype, L2997, an internal mould of a ventral valve with part
of the shell attached.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladyburn, near Girvan.

Begg 1947, p. 33, pi. 2, fig. 4 (L2996).

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION

389

Lifigulasma cf. pe?niltinnmi Reed
A ventral valve, L2998.

Ashgill, Upper Drummuck Group, Starfish Bed No. 2; Ladybum, near Girvan.

Begg 1947, p. 33, pi. 2, figs. 3, 3o.

Orthis (Glyptort/iis) bionmta Reed

Holotype, L3194 (BG 11716), external mould of a brachial valve.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1945, p. 309, pi. 1, fig. 5.

Orthis (Barbcirorthis) ci. foraminifera Opik
External mould of a dorsal valve, L2799 (BGS37).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Threave Glen (Ladyburn), near Girvan.
Reed \91>5b, p. 4, pi. 1, fig. 10.

Orthis (Platystrophia) praeciirsor Foerste var. molaris Reed
Holotype of the variety, a ventral valve, L2800 (BGllO).

Whitehouse Group; Shalloch Mill, near Girvan.

Reed 19356, p. 4, pi. 1, fig. 4.

Orthis (Barbarorthis) quarrelensis Reed

Holotype, an internal mould of a ventral valve, L2798 (BG126).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 19356, p. 3, pi. I, figs. 6, 6a.

Plectambonites praeteritus Reed

Holotype, L3195 (BG11726), external mould of a dorsal valve.

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1945, p. 312, pi. 1, fig. 4.

Plectodoiita thraiveusis (Reed) var. divergens Reed

Holotype of the variety, an internal mould of a ventral valve, L3314 (BG9558).

Caradoc, Craighead Mudstones (not Craighead Limestone, as stated); Craighead Quarry, near
Girvan.

Reed 1944c, p. 216, pi. 3, figs. 3, 3«.

Ptychoglyptiis shallocheiisis Reed

Holotype, half of a dorsal valve showing the hinge-area, L2803 (BG144).

Whitehouse Group; Shalloch Mill, near Girvan.

Reed 19356, p. 7, pi. 1, fig. 5.

Ptychoglyptiis virgiiiieiisis Willard var. irregularis Reed

Holotype of the variety is based on one valve showing ornament, L2794 (BG9482).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1941, p. 274, pi. 5, fig. 8.

Rafiiiesqiiina cf. hadelaiidica Holtedahl
A small ventral valve, L2796 (BG9479).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1941, p. 274, pi. 5, fig. 9.

Siphonotreta laggaiiensis Begg

Holotype, L3371, a small ventral valve.

Caradoc, Balclatchie Group; Laggan Gill, near Girvan.

Begg 1947, p. 44, pi. 3, fig. 9.

Sowerbyella cf. iiiidosa Opik

Internal mould of a small ventral valve, L2804 (BG1124).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Threave Glen (Ladyburn), near Girvan.
Reed 19356, p. 7, pi. 1, fig. 8.

390

PALAEONTOLOGY, VOLUME 6

Strophomena plamimbona (Hall) var. girvanensis Reed

Holotype of the variety, a ventral valve in interior view, L3316 (BG9741).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 1944c, p. 218, pi. 3, fig. 5.

Triplecia asteroidea Reed

Syntypes, three internal moulds, one of a complete shell, L2801 (BGS49), which is figured. Of the
other two, only one, L2802 (BG5228), is in the Hunterian Museum.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Threave Glen (Ladyburn), near Girvan.
Reed 1935(>, p. 5, pi. 1, figs. 1, \a-c (U2801).

Silurian

Meristina mullochensis Reed

Syntypes, an internal mould of a ventral valve, L2807 (BG2015), and an internal mould of a complete
shell, L2808 (BG1154).

Lower or Middle Valentian, Mulloch Hill Group; Rough Neuk (L2807) and Mulloch HiU (L2808),
near Girvan.

Reed 19356, p. 10, pi. 1, figs. 9, 9a (L2807).

Orthis (Ortliostrophia) camreganensis Reed

Holotype, an internal mould of a ventral valve, L2797 (BG108).

Upper Llandovery, Camregan Group; Penwhapple, near Girvan.

Reed 19356, p. 2, pi. 1, fig. 7.

Rhynchospirim dispar Reed

Holotype, a small ventral valve, L2806 (BG2030).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 19356, p. 9, pi. 1, fig. 3.

Carboniferous

Fluctaria undata (Defrance)

A ventral valve, L1536.

Visean; Poolvash, Isle of Man.

Muir-Wood and Cooper 1960, pi. 115, figs. 15-19.

^ j . . LAMELLIBRANCHIA

Ordovician

Ambonychinia balclatchiensis Reed

Holotype, a nearly complete shell, S7639 (BG9722).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 19446, p. 213, pi. 2, fig. 2.

Byssonychia quarrelensis Reed

Holotype, an internal mould of a right valve, S7644 (BG 11885).

Ashgill, Lower Drummuck Group, Quarrel Hdl mudstones; Quarrel Hill, near Girvan.

Reed 1946, p. 203, pi. 14, figs. 3, 4.

Ctenodonta collina Reed

Holotype, internal mould of a left valve, S7641 (BG11880).

Ashgill, Lower Drummuck Group, Quarrel HiU mudstones ; Quarrel Hill, near Girvan.

Reed 1946, p. 202, pi. 14, fig. 1.

Cyrtodonta reposita Reed

Holotype, an internal mould of a left valve, S7637 (BG9765).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 19446, p. 210, pi. 2, fig. 5.

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION 391

Goniophorina sp.

A left valve, S7648 (BG11889).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1946, p. 204, pi. 14, fig. 6.

Orthodesma cf. approximation Foerste
An internal mould of a right valve, S7647 (BG11888).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1946, p. 204, pi. 14, fig. 5.

Pletliocardia propinqiia Reed

Holotype, a nearly complete shell, S7638 (BG9762).

Caradoc, Balclatchie Group; Balclatchie, near Girvan.

Reed 19446, p. 211, pi. 2, figs. 3, 3u-c.

Vanuxemia ? girvanensis Reed

Holotype, a small shell, S7564 (BG9484), both valves, side by side but not in contact.

Ashgill, Upper Drummuck Group, Ladyburn mudstones. Zone of Dicellograptiis anceps; South
Threave, near Girvan.

Reed 1941, p. 276, pi. 5, fig. 4.

Vanuxemia aff. sardesoni Ulrich
Internal mould of a left valve, S7645 (BG11886).

Ashgill, Lower Drummuck Group, Quarrel Hill mudstones; Quarrel Hill, near Girvan.

Reed 1946, p. 203, pi. 14, fig. 2.

Silurian

Niicidites seductiis Reed

Holotype, an internal mould of a right valve, S7563 (BG9485).

Middle Valentian, Saugh Hill Group; Newlands, near Girvan.

Reed 1941, p. 276, pi. 5, fig. 10.

Ordovician

GASTROPODA

Archinacella unica Reed
Holotype, S9659 (BG6742).

Ashgill, Upper Drummuck Group, Ladyburn mudstones. Zone of Dicellograptiis anceps \ South
Threave, near Girvan.

Reed 1941, p. 275, pi. 5, figs. 5, 5a.

Carboniferous

Bucaniopsis decussatus (Fleming)

An almost complete specimen, S9679 (BG6988).

Carboniferous, Lower Limestone Group, ‘Chiton Bed’ ; Woodmill, Dunfermline.

Weir 1931, p. 821, pi. 8, fig. 12.

Eoptychia sulcata (de Koninck)

Two almost complete specimens, S9493 and S9494. A young specimen also figured (fig. 3) cannot be
found in the Begg Collection.

Carboniferous, Visean; Balladoole, Isle of Man.

Longstaff 1933, p. 113, pi. 10, fig. 1 (S9493), fig. 2 (S9494).

Euphemus urei (Fleming) var.

A specimen with incomplete apertural margin, S9674 (BG7013).

Carboniferous, Lower Limestone Group, ‘Chiton Bed’; Woodmill, Dunfermline.

Weir 1931, p. 845, pi. 9, fig. 8.

392

PALAEONTOLOGY, VOLUME 6

Glabrocinguhim atornarium (Phillips)

A complete specimen, S9070 (BG9326).

Lower Limestone Group, Charlestown Main Limestone; Woodmill, Dunfermline.

Thomas 1940, p. 40, pi. 2, figs. 2a, b.

Moitr Ionia monensis Weir

This species is founded on two specimens in neither of which is the aperture preserved, S9076
(BG7121), S9077 (BG7122).

Carboniferous, Poolvash Limestone; Balladoole, Isle of Man.

Weir 1925, p. 434, pi. 19, figs. 1, 2 (S9076).

Polytremaria beggi Weir

Syntypes, three specimens, none with complete aperture, S9078 (BG7123), S9079 (BG7125), S9080.
Carboniferous, Poolvash Limestone; Balladoole, Isle of Man.

Weir 1925, p. 435, pi. 19, figs. 3, 4.

Port lock ia cf. blanda (de Koninck)

Two well-preserved specimens, S9082, S9083. In S9082 the outer lip is slightly fractured, and in
S9083 it is absent.

Carboniferous, Poolvash Limestone; Balladoole, Isle of Man.

Weir 1925, p. 436, pi. 19, figs. 5, 6 (S9082). Knight 1933, p. 38.

Scalites ambigiius Weir

Holotype, a well-preserved specimen, S9081 (BG7110).

Carboniferous, Poolvash Limestone; Balladoole, Isle of Man.

Weir 1925, p. 437, pi. 19, figs. 8, 9.

Tychonia near Tychonia omaliana (de Koninck)

One specimen, S9075 (BG7126).

Carboniferous, Poolvash Limestone; Balladoole, Isle of Man.

Weir 1925, p. 433, pi. 19, fig. 7.

Ordovician

ECHINOIDEA

Aulecliimis grayae Bather and Spencer

An external mould of part of a test, E1225 (JLB19), one of the specimens used in description of
the species.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

MacBride and Spencer 1938, p. 98, pi. 12, fig. 2.

Ectinechiniis lamonti MacBride and Spencer

An external mould of part of a test, E1226 (JLB21), and internal moulds of parts of tests, El 127
(JLB22) and El 232 (JLB26), are part of the material used in the description of this species.
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

MacBride and Spencer 1938, p. 99, pi. 12, fig. 4 (E1226), pi. 12, fig. 6 (El 127), pi. II, fig. 4 (E1232).

Eothuria beggi MacBride and Spencer

Holotype, E1224 (JLB31), internal mould of part of a test, and a portion of its external mould,
E1224fl (JLB30).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

MacBride and Spencer, 1938, pp. 95, 126, pi. 16, figs. 1, 2 (E1224).

Ordovician

CRINOIDEA

Anulocrinus driimmuckensis Ramsbottom

Holotype, E3508fl and E35086 (BG9300Z)), counterparts, external moulds of cup and arms.
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 9, pi. 2, fig. 9 (E35086), fig. 10 (E3508u).

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION

393

Amilocriniis thiaivensis Ramsbottom

Paratypes, E3524 (BG3386) and E3496 (BG3389), both external moulds of cup and arms. The
holotype and other paratypes are in the British Museum, and there is also a paratype in the
Lamont Collection (Hunterian Museum).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 8, pi. 2, fig. 8 (E3524).

Archaeoa iniis elevatus Ramsbottom

Holotype, E3488o; paratypes, E3487n, E3488c, E3513, E3529, E36I5; all cups or parts of cups.
Caradoc, Stinchar Limestone Group, Orthis confinis Beds; Minuntion, near Girvan.

Ramsbottom 1961, p. 23, pi. 8, fig. 4 (E3488«), fig. 2 (E3487a), fig. 1 (E3488e), fig. 3 (E3615), text-fig.
11, p. 24 (based on E3487u, E3488u, E3488c, E35I3, E3529).

Cupidocriinis heterobrachialis Ramsbottom

Holotype, E3614 and E3614« (counterparts); paratypes, E3484 and E3484u (BG3377 and BG3377a)
(counterparts), and E3485 (BG3378u); all external moulds of stem, cup, and arms.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 12, pi. 5, figs. 1, 2 (E3614u), fig. 3 (E3484), fig. 5 (E3485).

Dendocriniis grauditiibus Ramsbottom

Paratypes, E3509 (BG7948), E3516 (BG3396), E3518 (BG7950), E3603, E3605, all external moulds
of the anal tube and parts of the stem, cup, and arms. The holotype is in the British Museum
(Natural History).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 15, pi. 4, fig. 1 (E3603), fig. 4 (E3509), fig. 5 (E3518), fig. 6 (E3605), fig. 7
(E3516). (Ramsbottom’s fig. 1 and fig. 2 should be interchanged: fig. 1 is the Hunterian paratype,
not, as stated by Ramsbottom, the holotype, which is fig. 2.)

Diabolocrbms sp.

Internal and external moulds of part of a large cup, E3504u (BG7841a) and E35046 (BG78416).
Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 29, pi. 4, fig. 10 (E35046).

Macrostyloainus cin ifer Ramsbottom

An external mould of stem with cirri, E3492 (BG945I). The holotype and paratypes are in the
Lamont Collection (Hunterian Museum).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 20, pi. 6, fig. 11 (E3492).

Frota.xocriiius girvanensis Ramsbottom

An external mould of stem, cup, and arms, E3606 (BG9475). The holotype is in the British Museum
(Natural History).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 19, pi. 6, fig. 15 (E3606).

Undetermined heterocrinid
An external mould of stem, cup, and arms (E3613).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Thraive Glen (Ladyburn), near Girvan.
Ramsbottom 1961, p. 10, pi. 6, fig. 5.

Carboniferous

Dialiitocriniis aculeatus (T. and T. Austin)

A young specimen in side view, E3459.

Tournaisian; Hook Head, Co. Wexford, Eire.

Wright 1955, p. 247, pi. 59, fig. 4.

Dialiitocrinus cataphractus (T. and T. Austin)

A young specimen in side view, E3472o (BG7349); calyx and arms somewhat crushed, E3473
(BG9467); calyx and arms in side view, E3450 (BG7330).

394

PALAEONTOLOGY, VOLUME 6
Tournaisian; Hook Head, Co. Wexford, Eire.

Wright 1955, p. 248, pi. 59, fig. 1 (E3473), pi. 59, fig. 5 (E3472a), pi. 64, fig. 3 (E3450).

Holcocrinus smythi (Wright)

Calyx and arms in side view, E3464 (BG9446).

Tournaisian, Supra-Dolomite Beds; Hook Head, Co. Wexford, Eire.

Wright 1952, p. 105, pi. 36, fig. 14.

Onychocrimis hibernicus Wright

Paratypes, E3466 (BG9447), calyx and arms in dorsal view, and E3478 (BG7359), part of calyx and
arms in side view.

Tournaisian, Supra-Dolomite Beds; Hook Head, Co. Wexford, Eire.

Wright 1934, p. 261, pi. 15, fig. 1, text-fig. 29 (E3466); 1954, p. 174, pi. 45, fig. 16, text-fig. 93 (E3466).

Ophiurocrinus dactyloides (T. and T. Austin)

Part of calyx and arms in side view, E3467.

Tournaisian; Hook Head, Co. Wexford, Eire.

Wright 1950, p. 21, pi. 6, fig. 3.

Scytalocrinus beggi Wright

Holotype, E3354 (BG8556), calyx and arms in side view.

Visean, Upper Caninia zone; Cass-ny-hawin, Isle of Man.

Wright 1938a, p. 285, pi. 11, fig. 12; 1938Z>, p. 343, pi. 13, fig. 2; 1950, p. 22, pi. 6, fig. 6.

Ulocrimis miciformis (M‘Coy)

A cup with two columnals, E3436 (BG7544).

Lower Limestone Group; Corrieburn, Campsie.

Bather 1920, p. 207, text-figs. 1 and 2. Wright 1939, p. 28.

Ureocrimis bockschii (Geinitz)

A calyx and one arm in side view, E3435.

Lower Carboniferous; Penton Linns, Liddesdale.

Wright and Strimple 1945, p. 225, pi. 9, fig. 10.

Ordovician

INCERTAE SEDIS — CYCLOC YSTOIDES

Cyclocystoides decussation Begg

Syntypes, an almost complete lower plate, E5071 (BG2004), a fragment of its counterpart upper
plate, E5071a (BG2005), together with two specimens in the Lamont Collection.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 1934, p. 220, pi. 11, fig. 2 (E5071a), fig. 3 (E5071).

Part of a lower plate and marginal ring, E5074 (BG9038).

Same horizon and locality.

Begg 19406, p. 24, pi. 1, fig. 3.

Cyclocystoides wrighti Begg

Holotype, E5073 (BG8213) and E5073a (BG8213a), counterparts, complete upper and lower plates
with marginal ring.

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Begg 19406, p. 23, pi. 1, fig. 1 (E5073), fig. 2 (E5073a).

Ordovician

CONULARIIDS

Conularia asteroidea Reed
Holotype, a crushed fragment, S9715 (BG2002).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.
Reed 1933, p. 357, pi. 19, figs. 4, 4a.

E. D. CURRIE AND T. N. GEORGE: THE BEGG COLLECTION 395

Comdaria cimctata Reed

Holotype, a nearly complete specimen, S9714 (BG2310).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1933, p. 357, pi. 19, figs. 3, 2>a-b.

Comdaria mirifica Reed
Holofype, S9711 (BG238).

Ashgill, Upper Drummuck Group, Sfarfish Bed No. 1 ; Ladyburn, near^Girvan.

Reed 1933, p. 356, pi. 19, figs. 1, \a-b.

Comlaria slateri Reed

Holotype, a crushed specimen, S9710 (BG237).

Ashgill, Upper Drummuck Group, Starfish Bed No. 1 ; Ladyburn, near Girvan.

Reed 1933, p. 355, pi. 19, figs. 2, 2a-c.

Comdaria truemani Begg
Holotype, C5980, an almost complete specimen.

Ashgill, Upper Drummuck Group, Starfish Bed a few feet above No. 3 ; Ladyburn, near Girvan.
Begg 1947, p. 37, pi. 3, figs. 4, 5.

Pseudoconularia currieae Begg

Holotype, C5981 and its counterpart C5981a, a large specimen.

Ashgill, Upper Drummuck Group, Starfish Bed a few feet above No. 3; Ladyburn, near Girvan.
Begg 1947, p. 39, pi. 3, fig. 6 (C5981).

REFERENCES

BATHER, F. A. 1920. Some British specimens of Ulocrinus. Trans, geol. Soc. Glasgow, 16, 207-19.

BEGG, J. L. 1934. On the genus Cyclocystoides. Geol. Mag. 71, 220^.

1939. Some new species of Proetidae and Otarionidae from the Ashgillian of Girvan. Ibid. 76,

372-82.

— — 1940fl. A note on the genera Staiirocephalus and Sphaerocoryphe, with the description of a new
species of Sphaerocoryphe. Ibid. 77, 295-304.

1 940Z>. On the genus Cyclocystoides, with the description of a new species from the Ashgillian of

Girvan. Trans, geol. Soc. Glasgow, 20, 21-29.

1943^7. Hypostomes of some Girvan trilobites and their relationship to the cephala. Geol. Mag.

80, 56-65.

1943Z?. Hypostomes of Girvan trilobites. Ibid. 80, 160.

1944. On the fringe of Tretaspis. Ibid. 81, 113-17.

1945. The illaenid pygidium. Ibid. 82, 107-13.

1947. Some new fossils from the Girvan district. Trans, geol. Soc. Glasgow, 21, 29-47.

1950. New trilobites from Girvan. Geol. Mag. 87, 285-91.

— — ■ 1951. Some new Girvan trilobites. Trans, geol. Soc. Glasgow, 21, 362-70.

and REED, F. R. c. 1945. Two new trilobites from Girvan. Ibid. 20, 260-3.

KNIGHT, j. B. 1933. The gastropods of the St. Louis, Missouri, Pennsylvanian outlier: V. The Trocho-
Turbinidae. J. Pcdeont. 7, 30-58.

LONGSTAFF, J. 1933. A revision of the British Carboniferous Loxonematidac. Quart. Journ. geol. Soc.
89, 87-124.

MACBRiDE, E. w. and SPENCER, w. K. 1938. Two new Echinoidea, Aidechinus and Ectinechinus, and an
adult plated holothurian, Eothuria, from the Upper Ordovician of Girvan. Phil. Trans. Rov. Soc.,
B, 229, 31-136.

MUIR- WOOD, H. M. and COOPER, G. A. 1960. The Productacea. Morphology, classification, and life
habits of the Productoidea (Brachiopoda). Mem. Geol. Soc. America, 81.

PHLEGER, F. B. 1937. Further notes on the Lichadacea. J. Paleont. 11, 253-6.

RAMSBOTTOM, w. H. c. 1961. British Ordovician Crinoidea. Mon. Palaeontogr. Soc. 1-36.

396 PALAEONTOLOGY, VOLUME 6

REED, F. R. c. 1930. Notes on some new Ordovician trilobites from Girvan. Ann. Mag. Nat. Hist.
Ser. 10, 6, 193-201.

1931(7. Additional new trilobites from Girvan. Ibid., Ser. 10,7, 97-105.

\9'i\b. Lower Palaeozoic trilobites of Girvan. Mon. Palaeontogr. Soc. Suppl., 2, 1-30.

1933. Some new Ordovician species of Conularia from Girvan. Geol. Mag. 70, 354-8.

• 1935(7. Lower Palaeozoic trilobites of Girvan. Mon. Palaeontogr. Soc. Suppl., 3, 1-64.

1935^. Some new brachiopods from Girvan. Ann. Mag. Nat. Hist., Ser. 10, 16, 1-12.

1941. A new genus of trilobites and other fossils from Girvan. Geol. Mag. 78, 268-78.

1942. Some new Carboniferous trilobites. Ann. Mag. Nat. Hist., Ser. 11, 9, 649-72.

1944u. Five new Ordovician trilobites. Geol. Mag. 81, 58-64.

1944(5. Some new Ordovician lamellibranchs from Girvan. Ann. Mag. Nat. Hist., Ser. 11, 11,

209-15.

1944c. Notes on some new Ordovician brachiopods from Girvan. Ibid., Ser. 11, 11, 215-22.

— — 1945. Notes on some Ordovician brachiopods and a trilobite from the Balclatchie Beds of Girvan.
Ibid., Ser. 11, 12, 309-16.

— ^ — 1946. Notes on some lamellibranchs from Quarrel Hill, Girvan. (With a note on the Quarrel Hill
section by J. l. begg.) Geol. Mag. 83, 201-5.

SHIRLEY, J. 1936. Some British trilobites of the family Calymenidae. Quart. Journ. geol. Soc. Lond.
92, 384-422.

THOMAS, E. G. 1940. Revision of the Scottish Carboniferous Pleurotomariidae. Trans, geol. Soc.
Glasgow, 20, 30-72.

TRIPP, R. P. 1954. Caradocian trilobites from mudstones at Craighead Quarry, near Girvan, Ayrshire.
Trans, roy. Soc. Edinb. 62, 655-93.

1957. The trilobite Encrinurus multisegmentatus (Portlock) and allied Middle and Upper Ordovi-
cian Species. Palaeontology, 1, 60-72.

1958. Stratigraphical and geographical distribution of the named species of the trilobite super-
family Lichadacea. J. Paleont. 32, 574-82.

WEIR, J. 1925. On new Carboniferous Gastropoda from the Isle of Man. Geol. Mag. 61,

1931. The British and Belgian Carboniferous Bellerophontidae. Trans, roy. Soc. Edinb. 56,

767-861.

WHITTINGTON, H. B. 1950. British trilobites of the family Harpidae. Mon. Palaeontogr. Soc. 1-55.

1956. Type and other species of Odontopleuridae (Trilobita). J. Paleont. 30, 504-20.

— — 1959. Silicified Middle Ordovician trilobites. Bull. Mus. Comp. Zool. 121, 371-496.

WRIGHT, J. 1934. New Scottish and Irish fossil crinoids. Geol. Mag. 71, 241-67.

1938(7. Some British Platycrinidae and descriptions of new species. Ibid. 75, 266-87.

— ■ — 19386. Anemetocrinus, n.gen. A five-armed poteriocrinid from the Lower Carboniferous lime-
stones of Scotland. Ibid. 75, 337-46.

1939. The Scottish Carboniferous Crinoidea. Trans, roy. Soc. Edinb. 60, 1-78.

— — 1950-5. The British Carboniferous Crinoidea. Mon. Palaeontogr. Soc.

and STRiMPLE, H. L. 1945. Mooreocrinus and Ureocrinus gen.nov., with notes on the family

Cromyocrinidae. Geol. Mag. 82, 221-9.

ETHEL D. CURRIE
T. NEVILLE GEORGE
Hunterian Museum,
University of Glasgow

Manuscript received 1 October 1962

• ■

‘ .

If

f- ■

\

I

t-

THE PALAEONTOLOGICAL ASSOCIATION

COUNCIL 1963

President

Professor T. Neville George, The University, Glasgow, W. 2
Vice-Presidents

Professor O. M. B. Bulman, Sedgwick Museum, Cambridge
Dr. L. R. Cox, British Museum (Natural History), London
Dr. W. H. C. Ramsbottom, Geological Survey OjBfice, Leeds

Treasurer

Dr. T. D. Ford, Department of Geology, The University, Leicester

Secretary

Dr. C. H. Holland, Department of Geology, Bedford College, London, N.W. 1

Editors

Mr. N. F. Hughes, Sedgwick Museum, Cambridge
Dr. W. S. McKerrow, University Museum, Oxford
Dr. Gwyn Thomas, Department of Geology, Imperial College of Science, London, S.W. 7
Dr. I. Strachan, Department of Geology, The University, Birmingham, 15

Other members of Council

Dr. C. G. Adams, British Museum (Natural History), London
Dr. F. M. Broadhurst, The University, Manchester
Dr. W. J. Clarke, British Petroleum Company, Sunbury on Thames
Dr. W. T. Dean, British Museum (Natural History), London
Dr. C. Downie, The University, Sheffield
Dr. R. Goldring, The University, Reading
Dr. M. R. House, The University, Durham
Mr. M. Mitchell, Geological Survey and Museum, London
Dr. J. W. Neale, The University, Hull
Professor F. H. T. Rhodes, University College, Swansea
Dr. A. J. Rowell, The University, Nottingham
Dr. R. J. G. Savage, The University, Bristol
Dr. C. D. Waterston, Royal Scottish Museum, Edinburgh
Mr. C. W. Wright, London

Overseas Representatives

Australia: Professor Dorothy Hill, Department of Geology, University of Queensland, Brisbane
Canada: Dr. D. J. McLaren, Geological Survey of Canada, Department of Mines and Technical
Surveys, Ottawa

India : Professor M. R. Sahni, Department of Geology, Panjab University, Chandigarh
New Zealand: Dr. C. A. Fleming, New Zealand Geological Survey, P.O. Box 368, Lower Hutt
West Indies and Central America: Dr. L. J. Chubb, Geological Survey Department, Kingston, Jamaica
Eastern U.S.A. : Professor H. B. Whittington, Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge 38, Mass.

Western U.S.A.: Dr. J. Wyatt Durham, Department of Paleontology, University of California,
Berkeley 4, Cahf.

PALAEONTOLOGY

VOLUME 6 • PART 2

CONTENTS

Lower Kimeridgian ammonites from the drift of Lincolnshire. By the late
W. J. ARKELL and J. H. CALLOMON 219

A new species of Komia Korde and the systematic position of the genus. By

E. C. WILSON, R. H. WAINES, and A. H. COOGAN 246

Some sihcified Ordovician fossils from South Wales. By nils spjELDNiES 254

The British Carboniferous species of Girvanella (Calcareous Algae). By
ALAN WOOD 264

The ostracod genus Neocythere in the Speeton Clay. By p. kaye 274

Revision of some Lower Cretaceous microspores from Belgium. By A. F.
DELCOURT, M. E. DETTMANN, and N. F. HUGHES 282

Problematical microfossils from the Cretaceous and Palaeocene of the Middle
East. By G. F. elliott 293

On the structure of leaves of Rhabdotaenia Pant from the Raniganj coalfield,

India. By D. D. pant and B. K. verma 301

A Palaeocene teredinid (MoUusca) from Iraq. By G. F. elliott 315

The recognition of sahnity-controUed mollusc assemblages in the Great Estua-
rine Series (Middle Jurassic) of the Inner Hebrides. By J. D. Hudson 318

The ecology and stratigraphical distribution of the invertebrate fauna of the
Great Estuarine Series. By J. D. Hudson 327

Microbiological colonization and attack on some Carboniferous miospores.

By L. R. MOORE 349

Late Jurassic mammalian fossils in the Sedgwick Museum, Cambridge. By
W. A. CLEMENS 373

Catalogue of described and figured specimens in the Begg Collection in the
Hunterian Museum of the University of Glasgow. By ethel d. currie and
T. NEVILLE GEORGE 378

PRINTED IN GREAT BRITAIN AT THE UNIVERSITY PRESS, OXFORD
BY VIVIAN RIDLER, PRINTER TO THE UNIVERSITY

SNl NIVINOSHillAlS SBI^VyaiT LIBRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHill

m I #%

'CT ' VVV ^ N

ES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHil^^JS SBIBVaaiT LIBRARIES SMITHSONL

SNl NVINOSHilWS S3ldVyan libraries SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHill

Z r- Z r- z

ES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHillAS SaiaVHan LIBRARIES SMITHSONI

CO Z ...■ CO

< S , <

5) CO

V §

CO Z (/)

BRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHil

_ CO Z CO —

liJ ,r^ ^

To

ES SMITHSONIAN INSTITUTION NOliniliSNI NVlNOSHillAlS SaiHVBaiT LIBRARIES SMITHSONI

Z r- z ^ , Z I- Z

O „ XoaTloX O XiVnToJX _ ~ XSFSJrs, O

DO
X)

>

:o
-

_ CO CO ■ _ CO

.SNl NviNosHilws saiavaan libraries Smithsonian institution NoiiniiiSNi nvinoshti
^ ^ ^ 5 ^ ^ ^

I I ^

CO

I
CO

o
z
>

CO z CO ' 2 CO Z CO

lES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHillNS S3iavaail LIBRARIES SMITHSONI

LSNI NVINOSHilWS SaiBVaail LIBRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHil

dvaan libraries Smithsonian institution NoiiniiiSNi nvinoshiiins sainvaan libr

C/) Zv (/) z

X
CO

3> ' 5 >

in 1/ ^ ' Z C/5 2

■ITUTION NOliniliSNI NVINOSHiltMS S3iyvaai1 libraries SMITHSONIAN INSTITUTION NOlil

- V 05 — cn — to ^

_ Z \ ... Z . . .yy,_ Z i . I

Too */qX oj /

<'5X /-i

N

avaan libraries Smithsonian institution NoiifuiiSNi nvinoshiiins saiavaan libf

r- z z r- z

riTUTiON NOliniliSNI NViNosHilws saiavaaii libraries Smithsonian institution noit

avaan libraries Smithsonian institution NoiiniiiSNi nvinoshiuns saiavaan libi

to 2 ^ ^ Z ^ Z

CO ^ .05 ^ Jo 05

o

riTUTiON NOliniliSNI NViNOSHiiiMS S3iavaan libraries Smithsonian institution noii

Z , Z r- Z r- z

(o ~ — CO to X

BRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHimS SBiaVaail LIBf

CO Z V CO Z CO z ,

z =i z: d S N

riTUTION NOliniliSNI NVINOSHilWS S3iaVMBn libraries SMITHSONIAN INSTITUTION NOli

CO

o

z

avaan libraries Smithsonian institution NoiiniiiSNi nvinoshiiiais saiavaan lib

r- Z r- Z r- . z

NOliniliSNI nvinoshiuns SBiavaan libraries smithsonian institution noij

I I r> r» A »-5 I

I i P