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VOLUME 12 • PART 3

Palaeontology

SEPTEMBER 1969

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
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ship Treasurer, Dr. A. J. Lloyd, Department of Geology, University College,
Gower Street, London, W.C. 1, England.

COUNCIL 1969-70

President : Professor Alwyn Williams, The Queen’s University, Belfast

Vice-Presidents : Dr. W. S. McKerrow, Department of Geology, Oxford
Dr. C. Downie, The University, Sheffield

Treasurer-. Dr. J. M. Hancock, Department of Geology, King’s College, London, W.C. 2

Membership Treasurer : Dr. A. J. Lloyd, Department of Geology, University College,
Gower Street, London, W.C. 1

Secretary: Dr. W. D. I. Rolfe, Hunterian Museum, The University, Glasgow, W. 2

Editors

Mr. N. F. Hughes, Sedgwick Museum, Cambridge
Dr. Gwyn Thomas, Department of Geology, Imperial College, London, S.W.7
Dr. Isles Strachan, Department of Geology, The University, Birmingham, 15
Professor M. R. House, The University, Kingston upon Hull, Yorkshire
Dr. R. Goldring, Department of Geology, The University, Reading, Berks.

Other members

Dr. F. M. Broadhurst, Manchester
Dr. L. R. M. Cocks, London
Dr. C. B. Cox, London
Mr. D. Curry, Northwood
Dr. A. Hallam, Oxford
Dr. Julia Hubbard, London
Dr. J. D. Hudson, Leicester

of Council

Dr. W. J. Kennedy, Oxford

Dr. J. D. Lawson, Glasgow

Dr. E. P. F. Rose, London

Dr. C. T. Scrutton, London

Dr. V. G. Walmsley, Swansea

Professor H. B. Whittington, Cambridge

Overseas Representatives

Australia: Professor Dorothy Hill, Department of Geology, University of Queensland, Brisbane
Canada: Dr. D. J. McLaren, Institute of Sedimentary and Petroleum Geology, 3303-33rd Street
NW., Calgary, Alberta

India: Professor M. R. Sahni, 98 The Mall, Lucknow (U.P.), India

New Zealand: Dr. C. A. Fleming, New Zealand Geological Survey, P.O. Box 368, Lower Hutt
West Indies and Central America: Mr. John B. Saunders, Geological Laboratory, Texaco Trinidad,
Inc., Pointe-a-Pierre, Trinidad, West Indies

Western U.S. A. : Professor J. Wyatt Durham, Department of Paleontology, University of California,
Berkeley 4, Calif.

Eastern U.S. A. : Professor J. W. Wells, Department of Geology, Cornell University, Ithaca, New York

© The Palaeontological Association , 1969

MEDUSAE FROM UPPERMOST PRECAMBRIAN
OR CAMBRIAN SANDSTONES, CENTRAL

AUSTRALIA

by MARY WADE

Abstract. Positive and negative moulds of circular bodies occur largely on depositional undersides of flaggy
sandstones of the Central Mt. Stuart Beds, attributed to the Upper Proterozoic. Respectively they represent
Hallidaya brueri gen. et sp. nov. and Skinnera brooksi gen. et sp. nov., two medusae with strikingly different
gastrovascular systems, each of a scyphozoan grade of complexity but lacking pronounced fourfold symmetry.
They do not closely resemble any form known from Ediacara, South Australia, but add to the slowly accumula-
ting evidence that medusae in the past have been more varied than at present. Hallidaya brueri is also found in
the lower part of the Arumbera Sandstone, WSW. of Alice Springs ; this formation may bridge the Precambrian-
Cambrian boundary.

In the course of geological exploration near Mt. Skinner, Northern Territory, Australia,
a sequence of maroon shales, sandstones, and quartzites was examined by Mr. A. L.
Halliday, geologist, Kennecott Explorations (Australia) Pty. Ltd. He discovered a
fossiliferous slab bearing six casts of an unknown fossil (PI. 68, fig. 1) approximately
4\ miles ENE. of Mt. Skinner, but was unable to trace it to its origin. During later work,
Mr. M. M. Bruer, field assistant with the same company, discovered a sandstone and
shale sequence containing a large number of these fossils approximately 1,100 ft.
higher in the section, 3 miles NNW. of Mt. Skinner. There is a general dip of 15° SW.
and the exposed structure is uncomplicated. No fossils have been found elsewhere in the
sequence, save for a few doubtful ‘worm casts’ still higher. On the bases of regional
reconaissance mapping and lithologic similarity this sequence is considered part of the
Central Mt. Stuart Beds (Halliday, personal communication) of Upper Precambrian
age. This formation is not known to contain fossils elsewhere, but at its northern ex-
tremity is overlain by the Grant Bluff Formation, to which a Cambrian age is assigned,
since it contains a Lower Cambrian Helcionella (N. H. Fisher, pers. comm.). The Central
Mt. Stuart Beds unconformably overlie a very rugged Archaean basement, as the beds at
Mt. Skinner also seem to do (Halliday, pers. comm.) but the contact is here obscured by
alluvium.

With aid from the Australian Research Grant to Professor M. F. Glaessner, and from Kennecott
Explorations, a short visit was made to Mt. Skinner (text-fig. 1) to collect fossils and study their occur-
rence. Mr. Halliday demonstrated the local geology and Mr. Bruer provided guidance in the field and
assistance in collecting; with the permission of Kennecott Explorations he has also prepared text-figs.
1 and 2 from maps and air photos in their possession. I am indebted to Mr. C. C. Brooks of Kennecott
Explorations for making arrangements for the trip and for giving permission to refer to the un-
published Kennecott final report, Mt. Skinner Examination, Northern Territory. Another field trip
to the vicinity of Alice Springs was recently supported by the Australian Research Grant to Professor
M. F. Glaessner. He and Mr. I. M. Thomas have constructively criticized the manuscript.

The material studied is deposited in the collections of the Geology Department, University of
Adelaide, South Australia.

C 6685 A a

[Palaeontology, Vol. 12, Part 3, 1969, pp. 351-65, pis. 68, 69. J

352

PALAEONTOLOGY, VOLUME 12

text-fig. 1. Locality plan showing roads connecting the Mt. Skinner area to the main
north-south (Darwin to Alice Springs) road. This portion of the Northern Territory is
indicated on the inset map of Australia.

The monotonous succession of flaggy, maroon red-beds is occasionally interrupted by
thin, widespread, pale greenish, flaggy beds, and by localized, coarse sediments such as
the strongly cross-bedded, arkosic sandstones and grits that cap two prominent hill-
tops overlooking the main fossil occurrence (text-fig. 2). In both the red and green flag-
stones the coarser sediments are arkosic and sometimes include clay pellets. Small-scale

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

353

text-fig. 2. The Mt. Skinner area (after mapping by A. L. Halliday), showing the known fossil
localities. The enlargement shows the main fossil localities on either side of the valley containing Mt.
Skinner No. 2 drill-hole.

354

PALAEONTOLOGY, VOLUME 12

cross-bedding is very frequent, and scoured surfaces are often quite extensive. Rarely,
masses of detrital mica are found.

A petrological report on samples of maroon and greenish, coarse and fine sedi-
ments was supplied to Kennecott Explorations by Australian Mineral Development
Laboratories (Appendix to Kennecott final report, Mt. Skinner Examination, Northern
Territory). It indicated that the sediments generally may be classified as subgreywackes
and impure siltstones to claystones. The coarser sediments are more strongly cemented
by authigenic quartz, and the red coloration is induced by iron oxides coating the
detrital grains, whether of quartz, rock-fragments, or feldspar. The green bands contain
copper minerals and lack iron oxide coatings on detrital grains. Where there is clay
cement present (as, particularly, in the finer beds) there is less authigenic quartz. Laths
of authigenic mica are present, and there is more evidence of stress and metamorphic
change reported from the petrological study than would be expected from field evidence.
The lithological setting has numerous parallels with the Precambrian fossiliferous
deposits at Ediacara (Wade 1968), though these lack rock fragments and have much less
detrital mica (Goldring and Curnow 1967).

Occurrence and fossilization. The main fossil occurrence lies almost on a direct line
between Kennecott Explorations’ drill-holes Mt. Skinner No. 2 and No. 3 (text-fig. 2),
on the sides of a small valley running ENE.-WSW. On the south side of the valley,
the fossils have been found through about 20 ft. of maroon sandstones and minor shales
above a green band which measured 16 ft. thick (where it was intercepted by the drill-
hole Mt. Skinner No. 2) and in the green band. On the north side they were encountered
more sparsely below and in the green band which here forms a dip slope and is strongly
weathered, outcropping poorly. Except on the steeper slopes of the valley, scree and
rainwash material covers much of the outcrop. One piece of float containing a fossil
was found in the creek between Mt. Skinner No. 2 and the main fossil outcrop; it is
most probably from a further occurrence of the fossils along the strike of the beds.

The fossils occur in two major forms:

A. In the elevated relief on depositional lower surfaces of beds. These are discs,
truncated at the edges, usually depressed in the centre. Their elevation ranges down from
about one-eigth of their diameter to totally flat.

B. Impressions in depositional lower surfaces of beds. Latex casts of these are almost
hemispherical in small specimens, to gently domed with elevation only about one-tenth
of their diameter in large specimens.

Group A. Most of the fossils belong to group A and are sandstone discs, elevated to
flat, and normally depressed in the centre, where the impressions of a number of con-
centrically ringed bodies (here called nuclei) are grouped. Quite a number of specimens
appear as though the rock has broken away to expose the nuclei, and removal of the
surface of one fossil with some partly exposed nuclei revealed their continuation inside
it. In numerous specimens the nuclei are scattered from a grouping around the centre;
in some the bodies seem to have been buried in a partly decomposed condition since
they leave scarcely a mark on the sediment (PI. 69, fig. 5), though the nuclei make firm
impressions and occur also beyond the outline of the body. Clearly, the nuclei have been
much more resistant to decay and to compression by sediment load than the bodies that
enclosed them; nevertheless they can be flattened, giving rise to extra annular rings in

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

355

their outer parts. Their thickest portion, a central boss, is often more resistant to com-
pression but some are compressed excentrically. The common preservation, as a cast of
the body enclosing moulds of the more resistant nuclei, is best considered a positive
composite mould in the sense of McAlester (1962). Although the usual occurrence of
these fossils is on the depositional lower surfaces of flagstones as positive composite
moulds, they have been collected in situ with counterparts in complete reverse on the
upper surface of the underlying bed. In these the mould of the fossil body may be little
altered from the shape it had when the positive composite mould (projecting down from
the bed above) was formed. The nuclei, however, are cast by sediment which infills
their impressions in the overlying bed. It would be tempting to regard them as primary
infillings of hollows in the animals by mud, like the casting of medusa-gonads (Nathorst
1881 ; Walcott 1898) if it were not for the fact that nuclei can be scattered away from the
body and still make impressions in the sediment which covered them (PI. 69, fig. 5).
These casts of nuclei are therefore secondary structures, counterpart casts (Glaessner
and Wade 1966; Wade 1968) set in the mould of the body. It is probably best to employ
the term negative composite mould ( McAlester 1962) for the whole complex preservation.

The partial rings of sediment enclosing the identifiable specimen (PI. 69, fig. 2a)
and its unidentifiable neighbour show that the specimens adhered tightly to the surface,
which is strongly scoured, while the sediment transport was in progress. Twenty or
thirty of these rings were observed on dismembered slabs of the one bedding plane, and
scattered individuals elsewhere, but only a few of the contained specimens were identi-
fiable. Perhaps a certain degree of decomposition caused dead bodies to adhere to the
substrate, for the preservation is exceptionally poor and only one depression that might
have sheltered such a creature in life is known. It is a gently curved hollow with a definite
circular outline, penetrating at least one lamina of sediment, and of the approximate
size to accommodate one of the smaller organisms. Alternatively, this hollow may
represent a specimen which chanced to be buried in the top of a bed; it lies in the
depression between small sedimentary ripples.

The nuclei may be scattered from the centre though still within the disc; many, few,
or none may be seen (PI. 69, figs. 3, 4). Discs without nuclei are not, strictly speaking,
identifiable, as it cannot be proved that a cast with no nuclei, or a positive composite
mould with one or two displaced nuclei, is the same form as a specimen with nuclei
in place. In the field as well as in the collection, sharp-edged casts, flat to gently curved
very low domes, are a rare accompaniment of the positive composite moulds and may
well represent casts of the aboral side, unmodified except by flattening against the
substrate.

Group B fossils are only half as common as group A. All save one are impressions on
lower surfaces. A cast of the internal spaces (PI. 69, fig. 12; text-fig. 5) occurs on an
upper surface. The amount of detail shown on latex casts of the moulds is very variable.
They tend towards a smooth, low, featureless dome but none of the larger specimens
have withstood sediment-load to the extent of being quite smooth, and they show
characteristic depressions. One of the least-flattened is illustrated (PI. 69, fig. 8). Small
specimens, being much more steeply curved and having much smaller radii, withstood
sediment load better, and featureless moulds in this size-range are known. It thus appears
that the usual preservation of group B fossils is as negative composite moulds.

356

PALAEONTOLOGY, VOLUME 12

The only distinction between poorly preserved fossils of group A and group B is that
they are in opposite relief on the same (upper or lower) surfaces; a few of the most
flattened that are possibly group B cannot be distinguished from group A fossils.

Evidence of numerous minute, unidentifiable organisms was seen on slabs like that
shown in Plate 69, fig. 7. Some small casts of medusiform shape (PI. 69, fig. 6) are also
known on the bases of slabs but they are not numerous. On the other hand, slabs with
their lower surfaces literally covered with structures half way between this medusiform
shape and the small load-casts seen on Plate 68, fig. 1 have been found. These minute
load-casts are almost ubiquitous. They could have been initiated by hollows where small
organisms decayed, but where there is evidence of variable thickness of the clay layers,
the load-casts have formed only above the thicker clay. This evidence is supplied by the
shape and particularly the texture (Wade 1968) of the undersurfaces of the sandstone
slabs, which often indicate that clay was almost confined to ripple-mark troughs while
neighbouring ridges were only thinly covered. Jungst (1934) described a cratering of
clay surfaces as clay dehydrates under water but the rather thin layers of clay involved
here are not likely to have been able to form deeply indented surfaces. Dehydration
could, however, have formed a surface dimpled enough to initiate the formation of
these minute load-casts, which have not penetrated into any sedimentary layers below the
clay in which they were formed. This supposition fits the confinement of the load-casts
to the areas of thicker clay better than either the view that the initial surface-irregularities
(and thus the load-casts) are all due to the decay of small organisms, or to scour marks.
Their frequent lack of consistent orientation also seems to rule out formation of the
initial irregularities by flow of water.

Fossil form A
Genus hallidaya gen. nov.

Type species. Hallidaya brueri sp. nov.

Diagnosis. As for type species.

Hallidaya brueri sp. nov.
Plate 68, figs. 1, 3-6; Plate 69, figs. 1-5

EXPLANATION OF PLATE 68

Figs. 1, 3-6. Hallidaya brueri gen. et sp. nov. Positive composite moulds on the undersides of sandstone
slabs. la-/, Paratypes F16461 a-f, X 1 ; example a is the only one appearing to possess a marginal
flange but this is probably due to the superposition of two individuals. Nuclei can be clearly seen
in a-e and less clearly in /. Branched, radial canals appear at the left of c. The five-rayed depression
at the centre of / may represent its mouth; its left and lower rays bifurcate, the lower of the two
right-hand rays traverses one of the four visible nuclei. 3, F16463, x2; specimen most clearly
showing dichotomous branching of radial furrows, nuclei faint. 4, Holotype, F16464a, x2;
specimen with numerous, rather small nuclei; radial furrows show faintly and connect to a circum-
central zigzag furrow which is only partly preserved. 5, Paratype F16465, approx. X 1-5; the other
specimen which shows a zigzag circumcentral furrow with radial furrows connected to its outer points,
and nuclei. 6 a-c, F16471a-c, x2; a, specimen with 13 nuclei; b, c, neighbouring well-preserved

specimens with numerous nuclei.

Fig. 2. Lower surface of slab F16470, x2-6; either the cast of a minute medusiform fossil similar to
that shown on Plate 69, fig. 6, or the counterpart cast of an isolated nucleus of H. brueri.

Palaeontology , Vol. 12

PLATE 68

WADE, Medusae from central Australia

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

357

Diagnosis. Body disc-shaped, with truncated margin. Surface of elevated discs either
flat, depressed in centre above discrete, ringed, inflated structures (here called nuclei),
even more widely depressed, or (rarely) gently curved in a very low dome showing, at
most, several, scattered nuclei. Branched, radial structure sometimes observed in those
with central nuclei; it is clearly preserved only in one specimen where it consists of
dichotomously branching, narrow furrows. Concentric corrugations very rare. Nuclei
relatively resistant to compression and decay, round to oval in plan view; an inflated,
equatorial Tyre’ forms a belt around a central, domed structure. Additional equatorial
rings and ovate furrows on the central dome show on flattened specimens. Normal
attitude of nuclei flat in one plane around centre of body but if crowded may be tilted
and overlapping, 3-13 are known in different specimens.

Holotype. FI 6464a (PI. 68, fig. 4).

Material. 49 well-preserved specimens in which both the over-all dimensions of the disc and the detail
of at least some of the nuclei can be seen and a number of additional, moderately good to poor speci-
mens; all these are positive composite moulds. Two identifiable negative composite moulds. A few
smooth, featureless, casts may also belong here.

Subsequent to the completion of this paper, several specimens of H. brueri were collected about
23 km. WSW. of Alice Springs (133° 4F, 24° 37'), in a clayey siltstone approximately a quarter of the
distance from base to top of the Arumbera Sandstone. This formation is about 600 m. thick here. It is
mapped and recorded in numerous publications, mainly of the Bureau of Mineral Resources, Geology
and Geophysics, Canberra, as uppermost Precambrian to Lower Cambrian, or more rarely as wholly
Lower Cambrian.

Dimensions. The common size-range is between 10 and 30 mm. in a known range of 5-50 mm. On
text-fig. 3 the over-all average diameter of each well-preserved specimen is plotted against the average
diameter of the largest well-preserved nucleus present in it. This choice was made to minimize bias
in the measurements; while flattening would tend to increase all diameters, the vagaries of tilting or a
shallow impression could only act to minimize size, and it would never be possible to establish whether
a small impression was the natural result of a small specimen or induced by preservation. The maximum
size ratio of smallest to largest well-preserved nuclei in one individual is about 2 : 3 in this material.
It has not been possible to recognize nuclei on specimens smaller than one of 5-5 mm., and specimens
without nuclei cannot be identified with certainty. Even with the rather small number of points
available to construct text-fig. 3, it suggests a direct relationship between the size of the over-all dia-
meter and the size of the nuclei.

Description. The fossils present several aspects. A few are gently convex, very low domes
that show, at most, a few nuclei scattered and tilted at any angle (PI. 69, fig. 3, F16467).
These could be aboral sides as they show no trace of a mouth, or tactile or locomotory
organs, nor is there any structure which could obscure their presence. The most com-
mon aspect, whether the specimens are flattened or not, has a group of nuclei extending
from the centre to occupy one-third to one-half of the diameter of the disc. They would
tend to obscure a central mouth or any other insubstantial structure placed in this posi-
tion. In unflattened specimens the surface is often depressed near the group of nuclei
and always depressed where the nuclei are; it also seems decidedly flatter than the
probable aboral side (PI. 68, figs. 1 c-f; PI. 69, fig. 1). From these factors of shape and the
association of scattered, tilted nuclei with the more convex side (PI. 69, fig. 3) it appears
that a planoconvex body with the nuclei lying nearer the flat side is the most natural
original shape to assume. As the nuclei tend to lie horizontally in a group around the
centre, they do not exclude the existence of a centrally situated mouth. Indeed, the

358

PALAEONTOLOGY, VOLUME 12

centre-top specimen (PI. 68, fig. If) possesses a small, five-rayed depression in this
position; it is not a compressed specimen and only faintly shows four nuclei situated
between the rays and over one of them; two of the rays branch dichotomously (text-
fig. 4b). A similar arrangement of three nuclei and a three-rayed impression is known
from one other specimen. A number of the specimens with groups of nuclei also show
a faint, branching, radial structure which is clearly seen only in F16463 (PI. 68, fig. 3),

mm

50

40 •

30

20 •

10 •

7

mm

text-fig. 3. Hallidaya brueri gen. et sp. nov. Scatter diagram of the average diameters of the discs
in millimetres plotted against the diameters of the largest nucleus in each disc. Holotype circled.

where it is seen to consist of dichotomous furrows. There is not enough structure pre-
served to tell whether the pattern of branching is regular in detail, but it is apparently
always dichotomous. No trace of it has yet been observed on any wholly convex side
(i.e. on any of the probable aboral sides), though it shows on some of the most convex
of the specimens with nuclei grouped at their centres (PI. 68, fig. lc). In the holotype
and in F16465 (PI. 68, figs. 4, 5) some radial furrows can be seen to branch from the
outer points of a zigzag furrow which partly encloses the centre and is partly obliterated
by nuclei.

Some of the nuclei appear to have escaped the confines of ruptured or decayed bodies
that may be faintly indicated on the same bedding plane (PI. 69, fig. 5, FI 6468) but
others are found completely free. Even when free of the discs, they normally held up the
sediment long enough to form external moulds on the bottoms of overlying beds. Some
casts of this general size have the proportions of nuclei (PI. 68, fig. 2, F16470) and are

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

359

also found on the bottoms of beds; either they represent counterpart casts of nuclei
which were partly buried in the top of a sediment layer, or the medusiform creatures
responsible for such casts as that shown in Plate 69, fig. 6 can overlap the nuclei in
proportions as well as size. Some casts which appear to represent uncrushed and partly
crushed nuclei are present on a few bedding planes crowded with innumerable small,
wrinkled, circular fossils which have the size range 3-6 mm. and could be either totally
flattened nuclei or the minute medusiform creatures. They have the appearance of
having been strongly inflated but are now flattened to an unidentifiable film (PI. 69,
fig. 7).

text-fig. 4. Hallidaya brueri gen. et sp. nov., approx. X 2. Restored as a translucent medusa showing
the structures illustrated in Plate 68, figs. 1, 3-6; Plate 69, figs. 3, 5. a, oblique exumbrellar view;
b, oblique subumbrellar view; m, mouth (PI. 68, fig. 1/); n, nuclei (many figs.); s, central stomach
(partially visible in PI. 68, figs. 4, 5); r, radial canals (PI. 68, figs, lc, 3-5). The marginal flange shown
here by dashed lines is seen only in Plate 68, fig. 1 a and is probably due to superposition of a larger
specimen on a smaller (several examples of partial superposition are known, e.g. PI. 69, fig. la).

Interpretation. As these fossils are of soft-bodied animals of which no original material
remains, we are left with a record only of the general shape and relative durability of
some of the organs. Association of certain structures in the fossils allows us to assume
their closeness, and dissociation their separation, in the undistorted animal. Besides
circular outline, it is notable that the mass preservation of the discs is convex downward
on lower sediment faces. Another feature is that the nuclei are present as moulds. From
these factors we see that the animals were buried in, or adhered to, sediment surfaces,
and were still present, either dead or alive, when the sand covered them but collapsed
quickly allowing it to cast them. The nuclei, however, did not collapse but were tough
enough to support the sediment until it set into moulds. The nuclei were discrete struc-
tures which could be displaced from their original position but probably would then
also suffer relative displacement such as we observe. In elevated specimens, radial,
branching furrows may show on the side of the disc that has the nuclei in orderly group-
ing, but not on sides with smooth, elevated contours in which only impressions of
displaced nuclei can be seen. (Radial structure can also show, as do nuclei, in flattened
specimens — in these the characters of the side with centrally grouped nuclei always
prevail to some extent over the featurelessness of the side which shows displaced nuclei,
or none.)

Assembling all these characteristics we have a circular body curved in a low dome on
one side, the curve of the dome covering structureless material that disintegrated quickly

360

PALAEONTOLOGY, VOLUME 12

upon burial. The opposite surface is flatter. It is either naturally concave or poorly
supported in the central region. It is most frequently smooth but in some specimens
shows branched, radial furrows across its outer part; these may be related to a sub-
surface system of canals which occasionally forms impressions on the casts, as their
usual absence would be very difficult to explain if they were to be regarded as a system
of surface markings. Their pattern is consistent with medusan — particularly scyphozoan
— radial canals, and they branch from the outer points of a rather zigzag furrow around
the centre, which could represent a central stomach. At about the same level as the
canals we have to place the nuclei. Normally assembled around the centre and lying
in one plane, they must also be beneath the epidermal layer, as they sometimes scarcely
show (PI. 68, fig. 1 b,f ), but the actual depth is probably rather variable: in specimens
in which they are very numerous they overlap somewhat. There is also the possibility
that a central mouth is represented by the star-shaped slits, which are rarely observed
in the centres (PI. 68, fig. 1/). The structure of marginal and submarginal furrows seen
in Plate 68, fig. la may represent two superimposed specimens or, perhaps, a nearly
vertical marginal flange like a velarium.

In summary, the proposed reconstruction indicates a medusiform body almost certain
to be a coelenterate medusa (text-fig. 4), whether Plate 68, fig. la is taken to represent a
marginal flange or not. The nuclei are placed in a position which would allow them to be
connected with the gastral epithelium, which is a common position for reproductive
structures in modern medusae; these also grow throughout life. The behaviour of the
nuclei as firm, discrete bodies cannot be equated with present day medusoid gonad struc-
tures except, perhaps, those in which the embryos are incubated during early development.
They are more probably the early stages of medusoid buds, but their later development
is not known. The possibility of their being food material has to be considered also but

EXPLANATION OF PLATE 69

Figs. 2 and 12 are of the depositional tops of sandstone slabs; all others are from the bases of slabs.

Figs. 1-3, 5. Hallidayci brueri, gen. et sp. nov., paratypes. la-c, F16462 a-c, X 1 ; a, superimposed
specimens; b, the largest individual, showing the strongest development of concentric ridging, which
is also seen on c. 2a, b, F16466 b, Xl; negative composite moulds. 3, F16467, xl; positive
composite mould of probable aboral side showing 3 tilted and 1 horizontally-placed nuclei. 5,
FI 6468, X 2, scattered nuclei, probably of 2 individuals, one of which is centred on the nucleus near
the centre of the picture.

Fig. 4. cf. Hallidaya brueri, FI 6469, X 1; probable aboral side.

Fig. 6. Cast of minute medusiform fossil, F164714, x2.

Fig. 7. Minute fossils which appear to have been circular and inflated but very flattened during
fossilization. FI 6472, x 1. Reverse side of slab shown in fig. 12.

Figs. 8-12. Skinnera brooksi gen. et sp. nov. 8, 9, X 1 ; 10-12, X 2; 8-11, negative composite moulds;

12, internal mould. 8, Paratype F16475k, relatively smooth, large specimen, centre undulating
and partly surrounded by 8 secondary depressions spaced around half the perimeter. 9a, b,
Paratypes, F16475«, b’, a, irregularly flattened, large specimen; b, earliest identifiable growth stage
with 3 large inner depressions and a fourth, smaller depression. 10, Paratype F16476; a small
specimen with 3 inner depressions ringed by 13 visible secondary depressions and a small obscured
area. 11, F16474u, b; a, holotype with 3 large, pouch-shaped inner depressions and 15 secondary
depressions, b, Paratype with 3 inner depressions and 15 secondary depressions. 12, Paratype
internal mould FI 6473, showing 3 inner pouches with broken canals at their inner ends, 15 secondary
pouches attached by double canals to the inner pouches or the central area, and more complex
structures placed between, and toward the margin from, the secondary pouches.

Palaeontology, Vol. 12

PLATE 69

WADE, Medusae from central Australia

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

361

their rather small size range at any size of the body seems against this (text-fig. 3), as is
the lack of oral sides without nuclei, and the lack of specimens that might be partly
digested.

An early view that these bodies might represent some sort of embryo capsule with
developing embryos, rather than an entire animal, seems negated by the constant
growth-rates of the nuclei and the bodies, and the overall medusiform character of the
bodies.

The system of radial canals in H. brueri is of a complexity very rarely reached in the
Hydrozoa but frequently found in the Scyphozoa. The apparent depth of the nuclei
within the body suggests also that they were more likely to have been related to the
gastral epithelium than to have been invaginations in the sub-umbrella. The structure
of the probable mouth resembles the bifurcating mouth-furrows of Rhizostomites
admirandus Haeckel but is not as regular. Although it apparently had only blunt,
finger-like lips like Rhizostomites and similarly lacked a manubrium, it shows no other
similarities to Rhizostomites and stands apart as a primitive scyphozoan-like medusa
without known, close relatives.

Fossil form B
Genus skinnera gen. nov.
Type species. Skinnera brooksi sp. nov.

Diagnosis. As for type species.

Skinnera brooksi sp. nov.

Plate 69, figs. 8-12

Diagnosis. Circular, low-domed, soft-bodied animals, relatively higher in proportion
to width in smaller specimens. Probable original shape plano-convex. Body resilient
but differentially compressible, revealing the pattern of major, internal, pouch-shaped
spaces by its surface depressions, which form a group of 3 large inner depressions near
the centre with a ring of 15 secondary depressions (wherever these can be counted)
outside them. The 3 largest, inner depressions tend to be pouch-shaped with a concave
curve toward the axial region like the inner pouches of a complete internal mould. In
this, a network of paired canals connects the 3 inner pouches to the centre on one side
and to 15 secondary pouches in a ring outside them but minor spaces and connections
near the margin may be reticular. Only the ‘inner’ depressions are seen on the smallest
individuals.

Holotype. F16467# (PI. 69, fig. 10o).

Material. 26 negative composite moulds and 1 internal mould. Several small, smooth, external moulds
and a number of very distorted specimens may belong here. They are described from latex casts of the
natural moulds.

Dimensions. The ratio of height to width decreases markedly with increasing size. This is no doubt
exaggerated by the larger specimens being more prone to sag against the substrate. Most of these are
badly distorted (PI. 69, fig. 9a). The range of average diameter is from 3-9 mm. to 32 mm. with a
decided maximum frequency in the vicinity of 10 mm. Maximum height known is about 2 mm. regard-
less of diameter.

362

PALAEONTOLOGY, VOLUME 12

Description. The low-domed bodies were probably smooth prior to burial but tended to
sag under load into a pattern of ridges and hollows which is standard for various growth
stages, though more obscure in some specimens than others. The smallest specimen
shows, in the cast, a pattern of three large, oval depressions and a fourth smaller one
to one side, with prominent, smooth ridges between them (PI. 69, fig. 9b shows the

text-fig. 5. Skinnera brooksi gen. et sp. nov., F16473, paratype, x3T. Clear areas represent gently
rounded prominences on this internal mould and stippled areas the depressions that bound the
prominences. Depressed areas with obscure structures are lightly stippled. Coincidence of the dominant
shear direction (x, y) in the rock and the radial canals partly obliterates the natural structure near the
margin, as does chipping of the margin above. Inner pouches, pl-p3; secondary pouches, 1-15; of
these, 1, 6, and 11, situated between the inner pouches, are larger than the others. They probably
attached directly to the central area (?stomach) as their paired canals run radially and are much longer
than those between inner and secondary pouches. There is no trace of structure derived from a ring
canal in the plexus of small structures near the margin.

original). This characteristic ridging between the depressions is common to all identifi-
able small specimens. Numerous secondary depressions are lacking in the very smallest
individuals; this may be an artifact of resistance to compression and not due to the
structures they indicate forming at a later growth stage. The inner depressions are the
largest and they are frequently pouch-shaped, concave on the axial side. No specimen
has more than these 3 largest depressions possessing concave axial sides. In the holo-
type the 3 secondary depressions placed between the inner depressions are larger than
the remaining 12 which occupy relatively more cramped positions. The accompanying

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA

363

paratype (PL 69, fig. 11 a, b) has the same number of depressions with similar proportions,
and the structures are again evident in the internal mould (PI. 69, fig. 12) where they are
expressed as sand mounds, apparently casting a system of natural spaces. In this internal
mould a third order of peripheral spaces is also cast by sand but the structures are small,
and obscure in detail. Only the holotype, among the negative composite moulds, shows
any evidence of these peripheral spaces, and that only by a few peripheral depressions
exterior to the secondary depressions. In one or more sectors of several other specimens
the number of secondary structures is 5 to each inner pouch (or depression). One speci-
men has only 3 secondary to one inner depression in the only sector in which they can
be counted. In medium-large specimens (PI. 69, figs. 11 a, 12) the secondary structures
may be radially elongate, to scarcely visible in relatively uncompressed specimens (PI.
69, fig. 8). In a number of the larger specimens the centre is obscure but the pattern of
depressions and ridges in the outer parts is radial. The most enlightening single specimen
(PI. 69, fig. 12; text-fig. 5) shows elevations of sandy sediment occurring in the relative
positions of every depression known from fossils of the external surface. It seems this
is an internal mould of a system of natural spaces in the body. The low, convex disc
on which the internal mould is displayed may approximate the shape of the underside
but it is more likely that the lower surface disintegrated, allowing the entry of sand to
even small canals and spaces. The sand fill is identical with the substrate and the margin
is a very sharply depressed furrow. Text-fig. 5 was drawn from the specimen onto a
5x enlarged photo to achieve accurate proportions and overcome difficulties in illu-
minating the radial structure. Its lightly stippled centre indicates where the internal
cast was either too shallow to form definite markings, or weathered or chipped away.
3 purse-shaped, inner pouches taper to canals at both axial ends. These canals appear
broken off, as though once attached to a central stomach which is not delineated here
because the numerous faint markings in the centre offer too many possible shapes for
unprejudiced restoration. A system of paired canals attaches the 3 inner pouches (or,
between them, the axial ‘stomach’ region) to 15 secondary pouches symmetrically
placed in a ring nearer the margin. These are separated by, and give rise to, smaller
connections and spaces too complex and small for clear preservation. These appear to be
connected in a reticular manner. Canals do not multiply by branching.

Interpretation. The shape of the internal system of spaces and canals closely parallels
the gastrovascular system of a medusa of the scyphozoan grade of complexity. The
central area may be equated to the stomach, with double connections to the 3 large inner
pouches and through these (or by-passing them in the inter-spaces) with double con-
nections to the 15 secondary pouches and further connections to the still smaller sub-
divisions. The present evidence is of dominant 3-fold symmetry. It is tempting to regard
the inner pouches as the direct homologue of the scyphozoan gastric pouches. They may
be, but they differ from any modern scyphozoan system (see, e.g., Mayer 1910, Hyman
1940, Thiel in Rees 1966) in each pouch being connected to the central stomach by 2
widely separated canals; again, the canals to the secondary pouches are also double;
even in the marginal plexus, minor spaces and narrower connections are still recogniz-
able and the whole is distinctly different from the more uniform plexus development of a
number of Semaeostomeae and Rhizostomeae. The similarities are probably due to
convergence and it is not likely that S. brooksi is ancestral to any modern medusa.

364

PALAEONTOLOGY, VOLUME 12

Despite the initial impression of similarity produced by the similar size and curvature
of H. brueri and S. brook si, they have very little in common. H. brueri has narrow,
dichotomously branched, radial canals. S. brooksi has its central stomach leading by
paired canals into 3 inner pouches, and thence to numerous, more marginally placed
pouches from which canals again branch. No example of a branching canal has been
seen, though there may be reticulate connections near the margin. While in H. brueri
reproduction by medusoid buds or incubation in the gonads appears to be firmly estab-
lished, there is no evidence of this in S. brooksi. H. brueri has a normal texture to its
mesogloea and collapses quickly when buried, forming casts. S. brooksi is much more
resistant and under the same conditions forms moulds.

There is little evidence of 4-fold symmetry in these two species or among the medusoid
forms from Ediacara (Upper Precambrian), nor is it a striking feature of the Cambrian
medusoids (Nathorst 1881; Walcott 1898; Moore 1956), though in Mesozoic times it
appears to have been well established. The taxonomic importance of 4-fold symmetry
in medusae has been emphasized by the loss of the less regular forms and the diversifica-
tion of the more regular forms to produce the modern aspect.

The few known fossils do not allow us to reconstruct scyphozoan phylogeny from
fossil material, and the field has been left largely to workers on Recent forms who are
in general agreement on the degrees of relationship within the modern scyphozoans but
in notable disagreement about which gave rise to which (Thiel in Rees 1966). Thiel’s
careful assessment of characters in common leads back to an ever more strongly tetra-
merous ancestral form for the Scyphozoa. Chapman’s work (in Rees 1966) verifies earlier
placement (Kiderlen 1937, vide Moore 1956) of the Conulariida as ancestral to scypho-
polyps and relatively close to Stauromedusae. The long range of the Conulariida, now
extending to Upper Precambrian (Glaessner and Wade 1966) would provide ample time
for the course of differentiation proposed by Thiel. In any discussion on the ancestral
form of the Cnidaria the position of the early medusae, which are not Hydrozoa nor
placeable among modern Scyphozoa, needs consideration. There seems no place in
Thiel’s scheme for the non-tetramerous fossil forms, even for those like Hallidaya and
Skinnera, as complex (in the characters we know) as the younger scyphozoans. Extend-
ing the concept of Scyphozoa beyond its limitation to modern forms, however, it is
reasonable to classify these new forms as parts of an early scyphozoan radiation.

REFERENCES

chapman, d. m. 1966. Evolution of the scyphistoma. 51-75, in rees, w. j., ed. 1966 (q.v.).
glaessner, m. f. and wade, m. 1966. The late Precambrian fossils from Ediacara, South Australia.
Palaeontology, 9, 599-628, pi. 97-103.

goldring, R. and curnow, c. n. 1967. The stratigraphy and facies of the late Precambrian at Ediacara,
South Australia. /. Geol. Soc. Australia , 14 (2), 195-214, pi. 10.
hyman, L. H. 1940. The Invertebrates : Protozoa through Ctenophora. New York and London.
jungst, h. 1934. Zur geologischen Bedeutung der Synarese. Geol. Runds. 25, 312-25.
kiderlen, h. 1937. Die Conularien, fiber Bau und Leben der ersten Scyphozoa. Neues Jahrb. f.

Mineralogie, Beil-Band 77, Abt. B, pp. 113-69, figs. 1-47.
mcalester, a. l. 1962. Mode of preservation in early Paleozoic pelecypods and its morphologic and
ecologic significance. J. Paleont. 36, 67-93.
mayer, a. G. 1910. The medusae of the world. Pubis. Carnegie Instn. 109 (1-3), 1-735.
moore, r. c., ed. 1956. Treatise on Invertebrate Paleontology, Part F, Coelenterata. Geol. Soc. Am. and
Univ. Kansas Press.

M. WADE: MEDUSAE FROM CENTRAL AUSTRALIA 365

nathorst, a. G. 1881. Om aftryck af medusor i Sveriges Kambriska lager. Kongl. Svenska vetenskaps-
akad. Hand. 19 (1), 3-34, pi. 1-6.

rees, w. j., ed. 1966. The Cnidaria and their Evolution. Zool. Soc. London Symp. 16.
thiel, Hj. 1966. The evolution of Scyphozoa. A review. 77-116 in rees, w. j., ed. 1966 (q.v.).
wade, m. 1968. Preservation of soft-bodied animals in Precambrian sandstones at Ediacara Range,
South Australia. Lethaia , 1 (3), 238-67.

Walcott, c. d. 1898. Fossil medusae. Monogr. U.S. Geo/. Surv. 30, 1-201, pi. 1-47.

MARY WADE
Department of Geology
The University of Adelaide
Adelaide

Typescript received 20 July 1968 South Australia

SPECIFIC FREQUENCY AND ENVIRONMENTAL
INDICATORS IN TWO HORIZONS OF THE
CALCA1RE DE FERQUES (UPPER DEVONIAN),
NORTHERN FRANCE

by PEIGI WALLACE

Abstract. Statistical examination of the fauna from two horizons in the Calcaire de Ferques (Frasnian) of
northern France, combined with thin section studies and field observations, lead to the conclusion that while the
fauna of the lowest bed of the formation suffered slight transportation, this did not disrupt the ecological
associations of the fauna and a true life assemblage is represented. The fauna of the upper beds accumulated in
situ in an environment with a gradually increasing argillaceous content, and shows features which may be asso-
ciated with these deteriorating conditions and with the level of background radioactivity.

The Calcaire de Ferques is a highly fossiliferous bioclastic limestone, world-famous for
its rich and varied fauna, which includes topotypes of many well-known species such as
Cyrtospirifer verneuili. It outcrops only in the Palaeozoic inlier of the Boulonnais,
northern France (text-fig. 1), the geology and palaeoenvironments of which have been
redescribed by Ager and Wallace (1967).

The Calcaire de Ferques has been intensively studied in the last hundred years,
notably by Robinson (1920) and by Pruvost and Pringle (1924). It was given its name by
de Verneuil (1838), who compared its fauna with that of the Wenlock Limestone, which
it closely resembles both in faunal groupings and in mode of preservation. However,
Murchison (1840) referred it to the newly erected Devonian System. It was correlated
with the Calcaire de Givet, the type of the Givetian stage, by Godwin Austen (1853),
but Gosselet (1860) compared it to the Calcaire de Rhisnes, which was included by
d’Halloy (1862) in his newly erected Frasnian stage.

No further modifications of its stratigraphical age have been published, but it must
be considered to be of Middle Frasnian age since both the underlying Schistes de Beaulieu
and the overlying Schistes de Fiennes lie within this substage.

Until the end of the last war, the limestone was extensively worked in a series of small
quarries strung out along the outcrop. The degree of exposure is excellent at the time of
writing, but is deteriorating rapidly because many of the smaller quarries are being
infilled with overburden from the more extensively worked Carboniferous Limestone
nearby. The majority of the quarries terminate along the strike at one of the numerous
small dextral wrench faults which dissect the area (Wallace 1968) and which are well
exhibited in the largest quarry, La Parisienne at Beaulieu (text-fig. 1).

The palaeoenvironments of the main part of the Calcaire de Ferques and the
palaeoecological relationships of its fauna are discussed in detail by the author else-
where (Wallace, in press). They may be summarized as follows: a succession of en-
vironments of rapidly increasing and then gradually decreasing depth, passing from the
Zones Subturbulente and Turbulent e of Lecompte (1961) to deeper, quieter, open shelf
conditions with the formation of brachiopod and coral-bearing calcilutites in the middle

(Palaeontology, Vol. 12, Part 3, 1969, pp. 366-81, pi. 70.]

P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 367

part of the formation. The upper part of the formation shows a gradually shallowing
sequence with increasing terrigenous material. No previous account, however, has
attempted to discuss the environment as it changed from the hypersaline lagoons of
the Schistes de Beaulieu into the biostromes of the Calcaire de Ferques, or from the
latter into the subtidal mudflats of the overlying Schistes de Fiennes.

A neglected truism about quarry geology is that the most interesting features in any
quarry are at the top and bottom of the working faces. This is logical, since the excava-
tion normally ceases where the character of the rock changes. It is especially true in

text-fig. 1. Locality map of quarries in the Calcaire de Ferques, showing the outcrop of the limestone
and the position of the dextral wrench faults which displace the formation.

the numerous quarries of the Calcaire de Ferques, where only the main part of the
limestone is massive. The formation dips at about 30°, and huge bedding planes of the
Thin Basal Limestone are exposed in every quarry, providing the large amounts of data
which are considered statistically here. Similarly, the argillaceous nature of the Upper
Beds of the limestone makes them unsuitable for quarrying. They are well exposed along
the southern faces of many of the quarries, where extensive collections have been made.

Although the main part of the limestone is well exposed in one or two quarries,
notably Carriere du Bois, so that a general impression of the succession of environ-
ments may be obtained (Wallace, in press, text-fig. 13), only dip sections are available.
That, and the more massive nature of the rock, make statistically reliable collections
and observations unobtainable.

THIN BASAL LIMESTONE

The basal bed and earliest development of limestone within the Calcaire de Ferques
is the Thin Basal Limestone, 2 cm. thick, resting on the dolomitic sand facies of the

b b

C 6685

368

PALAEONTOLOGY, VOLUME 12

Dolomie de Fiennes and overlain by a moderately fossiliferous sandy dolomite (Ager
and Wallace 1967, bottom of fig. 2). The limestone itself has a red or grey calcilutite
matrix, rich in bioclastic fragments, and contains an extremely abundant and diverse
fauna. Dolomitization affects both the upper and lower surfaces of the bed, but rarely
alters it completely. The fauna is completely unaffected. Iron enrichment commonly
accompanies the dolomitization ; many of the dolomite rhombs are zoned, and the upper
and lower bedding surfaces are commonly dark red.

The population structure of the Thin Basal Limestone is shown in text-fig. 2. In both
this histogram and the similar one for the Upper Beds (text-fig. 4) the population is
expressed as percentage of sample rather than in absolute figures. Although it is realized
that ideally for comparison the population should be expressed in terms of standard
quadrats, this was not possible because of difficulties in the field, especially with the
Upper Beds. Instead a faunal census was taken for each locality. To eliminate observa-
tion and collection bias, the accuracy of this census was checked in two ways. First,
the assistance of two other people, one a geologist and the other a non-geologist in the
field for the first time, was enlisted. Although the size of the collections varied, the
proportions between the various species collected by the author and her two assistants
remained remarkably constant. Secondly, a bulk sample from the exposed surface of the
Thin Basal Limestone was analysed. Again the proportions between the species cor-
responded closely to those in the collections made by hand. Therefore it is considered
that the faunal censuses represent a valid sample of the population.

Numerically, by far the most dominant species in the Thin Basal Limestone is Cyrto-
spirifer verneuili, which consistently forms about 40% of the sample. All growth stages
are present. There is considerable size and shape variation within the species and it might
be possible to erect up to a dozen nominal species for this horizon alone. All these
‘species’ intergrade, however, and both Gosselet (1894) and Vandercammen (1959),
in statistical studies of the species from the Boulonnais and Belgium respectively,
concluded that only one species was present.

Athyris concentrica is rather more common in the west of the outcrop than in the
east, where its place seems to be taken by Productella sabaculeata. This latter species,
with its very delicate spines, might suggest that water conditions were rather quieter
in the east than in the west, although flume experiments (Ferguson, pers. comm.) suggest
that spiny productids may have been adapted to regimes with moderately strong uni-
directional currents. Corals are relatively rare within the Thin Basal Limestone itself,
although they occur abundantly within thedolomitic sand immediately below. Bryozoans
may be more common than suggested by the histogram (text-fig. 2), but their delicate
nature makes fragmentation almost inevitable, and hence counting difficult. For that
reason, only their presence has been indicated and the percentages shown are not a true
representation of abundance.

Even cursory examination of the histogram (text-fig. 2) shows some basic associa-
tions between species. For example, (a) where spiriferids and athyrids are abundant,
corals are virtually absent; ( b ) there is an inverse relationship between Cyrtospirifer
and Athyris ; (c) where corals are abundant, athyrids and rhynchonellids are low, (d)
where productellids are low there is a low specific diversity. The y2 test was used to test
these and all other relationships in the bed. The test was applied 2i-1 n t'mes where
n = number of recognized species or species groups. Thus not only was each individually

0

EAST

text-fig. 2. Histogram of the population structure in the Thin Basal Limestone, Calcaire de Ferques.
Values are expressjd as percentage of the assemblage, rather than as absolute figures, for ease of
comparison. Epifaunas and minor elements omitted. Localities are in order along the strike, not in a

stratigraphic succession.

370

PALAEONTOLOGY, VOLUME 12

listed species considered, but also certain basic groups such as spiriferids, rhyn-
chonellids, cerioid corals, solitary corals and stick bryozoans. This brought n to 37, and
hence the number of tests to nearly 600. Of these relationships, 47 were significant to a
1 0% level, 25 to a 5% level (the level most usually applied by modern ecologists according
to Kershaw (1966) and accepted as valid by Johnson (1962)) and 3 were significant
to less than 0-01%, a very high level of significance indeed. It was considered useful to
include levels higher than 5% (i.e.p = 0-05) in both tables and figures, since this demon-
strates associations and linkages which can be subjectively observed in the field both in
this region and in other Devonian areas, but it should be noted that their statistical
significance is small.

The results of the y2 test are summarized in Table 1. They show a close correlation
between all types of coral, especially Hexagonaria and the solitary corals. Corals are
also fairly strongly associated with other species; for example Spinatrvpa aspera and
(less strongly) Ptyehomaletoechia boloniensis with the cerioid forms; and gastropods,
Tenticospirifer tenticulum and S. aspera with the solitary forms. Cyrtospirifer shows a
negative correlation with both cerioid and solitary forms when it is present in proportions
greater that 30%, which was observed from casual inspection of the histogram, but the
converse association, that of high (>30%) Athyris concentrica with corals is not justified
statistically, although the inverse relationship between Athyris and Cyrtospirifer is
significant to p = 0-07.

Some of the associations, such as Chonetes with Douvillina and Nervostrophia, might
have been expected since the forms concerned were probably adapted to rather similar
environments. Close associations also appear to exist between other brachiopods, such
as the rhynchonellids (both Ptyehomaletoechia and Cupular ostr uni), Athyris, delthyrids,
Schizophoria, and Spinatrypa, and these are also associated with gastropods and Pseuda-
viculopecten. Since Cyrtospirifer occurs in every collection, it is difficult to prove a
statistically valid association between it and the other brachiopods, but logically this
association (which may be only a tolerance of similar conditions) can be seen to
exist.

Negative associations, from which one might possibly deduce some antipathy or
intolerance of similar conditions, occur between Cyrtospirifer (when present in very
high proportions) and gastropods, and between strophomenids and gastropods. A
general negative correlation also occurs between the spiriferid group and the coral
group; it is exhibited by Cyrtospirifer I Hexagonaria, Cyrtospirifer/ massive corals,
Cyrtospirifer/ solitary corals, Tenticospirifer/ Alveolites and delthyrids / D isp hy I him.

Text-fig. 3 is a constellation diagram of the positive associations in the Thin Basal
Limestone. The strongest associations are between the two major coral groups, the
solitary and the massive corals, with a weak association with phaceloid forms. This
plexus is linked more loosely with the Cyrtospirifer/ Athyris association of brachiopods
and with gastropods. Very loose links occur with the remainder of the brachiopods,
the pectinids and the stick bryozoans, which form a separate plexus centred round a
strong association of Ptyehomaletoechia with Schizophoria. A strong link exists between
Chonetes and the strophomenids, but the strophomenids are negatively associated with
several other members of both plexuses, and this association, although positive in
itself, is probably not related to the main constellation.

The only major species which shows no statistically significant association with any

0-001 +ve Hexagonarialsolitary corals 0 08 — ve Del t h y r i d D isphyllwn

0-001 +ve Massive corals/solitary corals 0-08 +ve Cupularostrum\Ptychomaletoechia

0 001 +ve Hexagonarial small solitary corals 0 09 +ve Hexagonarial Disphyllum

0-002 +ve Ptychomaletoechia/Schizophoria 009 — ve Schizophoriaj Disphyllum

0-005 +ve Hexagonarial large solitary corals 010 +ve Tenticospirifer\Cupularostrum

P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 371

<L> <U

C C

o o

JZ ZZ -
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JZ ZZ V-

C* c/3

I + I + + + I + I I I + I + + + I I + + + + I + + + +

666666666666666666666666666

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to VO h

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6666666666666

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0 07 + ve Atrypa squamifera/ gastropod 0-20 +ve Solitary corals/gastropod

0-07 + ve Cyrtospirifer < 30%/ Athyris > 30% 0-20 +ve Atrypids/rhynchonellids

0-08 +ve Cyrtospirifer < 30%/massive corals

Sch

372

PALAEONTOLOGY, VOLUME \2

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P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 373

other is Productella subaculeata. Its strongest association is with S. aspera (positive with
p — 0-16). The general trend of its associations shows that it would form part of the
coral plexus, linked more loosely to the massive forms than to the solitary forms.

Epifaunal encrustation is moderately common at this horizon, the brachiopods
particularly being affected. About 10% of individuals of Cyrtospirifer verneuili and a
rather smaller proportion of A. concentrica are affected. The most common organism,
and often apparently the first to arrive, was Spirorbis. Aulopora frequently encrusts the
fold of the brachial valve of the spiriferids, though it also occurs on the wings. Other
epifaunal elements, such as the encrusting bryozoans Hederella and Paleschara , are
rarer. These encrustations are similar to those observed by Ager (1961) on Spinocyrtia
from the Devonian of Iowa.

Thin-section studies permit close examination of matrix relationships, especially those
of the broken shells and bioclastic fragments (PI. 70, fig. 2; text-fig. 5). Separated
brachiopod valves fell concave downwards in the biocalcarenitic matrix, often leaving
an ‘umbrella structure’ of space beneath them. These spaces were colonized by Girvanella-
like algae, which also acted as a cement for the matrix. The overturned shells and shelly
sand provided a protected environment for the algal colonies and would also have
shielded them from excessive light. Senes (1967) has found that most Recent algae in the
Mediterranean (although not directly comparable to Girvanella , whose affinities are
uncertain) cannot tolerate more than 70% luminosity and many prefer much less, often
living beneath overhanging ledges or amongst rocks and sand-grade sediment. After
the growth of the algae beneath the shells, burial and lithification, drusy growth of
calcite filled the remainder of the cavity. Subsequently dolomitising fluids attacked both
the upper and lower surfaces of the bed.

The highly broken nature of much of the shell debris suggests forceful comminution,
either by waves or by scavenging organisms, but the algae show that once deposited,
the bed remained undisturbed until lithification. It is thus possible that the bed was
formed in rather deeper water than suggested by the comminution of the shell debris,
perhaps by the action of a single storm, dumping shell debris and whole shells in an
alien environment,

The size frequency histograms of the main species of brachiopods (text-fig. 6) mainly
show a normal bell-shaped distribution, which according to Boucot (1953) suggests
post-mortem transport and sorting of the assemblages. Other features which suggest
transported assemblages are the absence of juvenile rhynchonellids, and the remarkable
correlation of endpoints of three of the species, and of the main size peak in all four
species at 1-1 or 1-2 cm. length. It is most unlikely that all four species would achieve
mature shell sizes to within 1 mm. of each other, but size sorting of this kind might be
expected in a transported assemblage. The frequency of all species tails off at 1-5-1 -7 cm.,
again a remarkable correlation, though it should be noted that the spiriferids show a
small peak of larger individuals.

UPPER BEDS

The Upper Beds of the Calcaire de Ferques, varying in thickness from c. 8-10 m.
are noticeably richer in argillaceous material than the underlying bulk of the limestone,
and pass gradually into the shales of the overlying Schistes de Fiennes. The fossils,
although of species similar to or identical with those in the main part of the limestone,

374 PALAEONTOLOGY, VOLUME 12

are markedly bigger, especially those species of brachiopod presumably attached by a
pedicle during life.

The first large individuals to occur are preserved in a matrix of grey bioclastic cal-
cilutite very similar to that of the main part of the formation, but the argillaceous content
of this calcilutite increases rapidly upwards. The fauna within the calcilutite is very well
preserved; for example, atrypids still have their spires and very delicate fringe of lamel-
lose growth lines undamaged (PI. 70, fig. 1 ; text-fig. 4).

Some incipient dolomitisation is present; the proportion increases towards the top of
the formation. Limonitic zoning within the rhombs is frequent, but there is little iron
staining within the rock as a whole, and no iron sulphide is present. The uppermost
limestones are greenish-grey (5 GY 6/1) in colour and frequently have a sandy appear-
ance because of the numerous rhombs of dolomite on their surfaces. They are inter-
bedded with greenish-grey shales.

Although the fossils are the largest within the formation, the fauna is the least diverse,
and epifaunal encrustation similar in species and position to that in the Thin Basal
Limestone is common.

Pathologically deformed specimens of both spiriferids and atrypids form a surprisingly
large proportion of the assemblages, often up to 10%. Deformation usually took place
at a growth-line, and consists either of invagination of both valves or, in the case of
the spiriferids, of a marked branching of the ribs on the wings.

The population structure of the Upper Beds is shown in text-fig. 7. As with the Thin
Basal Limestone, visual inspection indicates an inverse relationship between Cyrto-
spirifer and Athyris. Where C. verneuili forms a dominant (i.e. >30%) part of the sample,
and especially where this is supported by the presence of other spiriferids, the proportion
of Athyris concentrica is markedly lower. It can be seen from Table 2 that the number of
significant associations in the Upper Beds is considerably lower than in the Thin Basal
Limestone, largely as a result of the greatly diminished diversity of the former. Many
of the most important members of the assemblage still remain, and this diminished
diversity has been achieved at the expense of elements such as bryozoa and some of the
less-abundant brachiopods. The constellation diagram (text-fig. 8) shows rather different
groupings from those of the Thin Basal Limestone, with a plexus based on a strong

text-fig. 4. Diagram of the main features of the photomicrograph shown in Plate 70, fig. 1. An atrypid
brachiopod is in presumed position of life. The shell structure can be clearly seen, and the position
of the frills and small pieces broken from them show that the shell has been undisturbed since death,
and was probably killed by burial. Geopetal sedimentation within the shell indicates the horizontal:
it can be seen that more sediment entered through the anterior commissure and piled up against the
first spire. The spires have moved from their position during life, possibly owing to internal decay of
the soft parts. They are surrounded by an oval shadow which may represent the final position of some
of the soft parts. Drusy crystalline calcite fills the upper part of the shell.

text-fig. 5. Diagram of the main features of the photomicrograph shown in Plate 70, fig. 2. Separated
valves of brachiopods, convex up, in a bioclastic, calcarenitic matrix, have left umbrella-shaped
sheltered spaces beneath. The large shell covers a gastropod with an open umbilicus. Girvanella-
type algae grew in the enclosed spaces, carpeting the sheltered bottom and filling the body-chamber of
the gastropod. The remaining space beneath the shell was subsequently filled with drusy crystalline
calcite. Dolomitisation attacked the upper and lower surfaces of the bed: the iron-enriched rhombs
form dark patches on the photomicrograph. Large-zoned rhombs grew in the inner chambers of the
gastropod. The shell structure of the brachiopod shells (probably atrypids) is clearly visible.

TEXT-FIG. 5

376

PALAEONTOLOGY, VOLUME 12

inverse link between Cyrtospirifer and Athyris very loosely linked to a coral/rhynchonel-
UdlSchizophorialstrophomenid plexus. These links are considerably less strong than
those of the Thin Basal Limestone, but the relationships between species are similarly
oriented, i.e. a species does not change the nature of its relationship with another species
from bed to bed, but only the strength of the relationship.

Other associations and occurrences may be noted from the histogram, such as the
presence of fan bryozoans at the eastern end of the outcrop only, and of stick bryozoans
only in the west. It may be noted that gastropods are more important than they were
in the Thin Basal Limestone.

ECOLOGICAL DEDUCTIONS

The environments in which the earliest and latest beds of the Calcaire de Ferques
were laid down were clearly very different, although both fall within the broad classi-
fication of ‘shelf biostrome’.

The Thin Basal Limestone consists of an assemblage of very well-preserved whole
fossils in a coarsely bioclastic matrix. It may well be that much of the comminution of
the shell debris was achieved by scavengers turning over the sediment in their search
for food, rather than by current or wave action. All the fauna (with the possible exception
of the nuculoid lamellibranch found at one locality only) is epibiontic, most are sus-
pension feeders and many are attached forms, which may suggest that this was a firm
bottom fauna. The author considers it more probable, however, that the original sea
floor was covered by a layer of bioclastic debris, since in all species where one may see
an attachment area (such as both solitary and cerioid corals, and exceptionally
Schuchertella), the attachment areas show moulds of shell ornament, either spiriferid,
strophomenid, or schizophorid, or show evidence of attachment to other bioclastic
debris, such as crinoid ossicles. No evidence is found of forms attached to a substrate
such as a rocky sea floor.

Although the bell-shaped curves of the size frequency diagrams suggest a transported
fauna, the fact that y2 tests show significant associations between species argues against
extensive transportation, as does the unabraded nature of all members of the assemblage
The presence of in situ algae beneath overturned shells suggests very little reworking of
the bed. There are three possible solutions of this apparent incompatability :

(a) The curves may not be truly bell-shaped;

(b) The curves are bell-shaped, but true life associations are still represented;

(c) The associations proved by the y2 test may be mechanical rather than ecological.

Thus if the faunal associations are significant ecological associations, it must be
concluded that bell-shaped curves may occur in biocoenoses as well as in thanato-
coenoses, as suggested by Hallam (1967). Moreover, it may be deduced from the work

EXPLANATION OF PLATE 70

Fig. 1. Photomicrograph of a vertical section through the Upper Beds, Calcaire de Ferques; X 10,
(see text-fig. 4).

Fig. 2. Photomicrograph of a vertical section through the Thin Basal Limestone, Calcaire de Ferques;
x 7-5, (see text-fig. 5).

Palaeontology, Vol. 12

PLATE 70

WALLACE, Calcaire de Ferques

P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 377

40-

35-

30-

25-

20-

15-

10-

5‘

25-

20-

15-

10-

5-

TEXT-

Cyrtospirifer

verneuili

X-

1

65 -

60

55-

50

45

40

35-

30-

10 15 20 25 cm 25

Length

20-

15-

Ptychomaletoech/a

boloniensis

io-

£

5 10 15 cm

Length

Athyris

concentr/ca

I

■5 10 15 cm Length

Spmatrypa
aspera

5 10

1-5 cm Length

fig. 6. Size-frequency histograms of the main brachiopod species from a single locality in the
Thin Basal Limestone at Carriere de La Parisienne.

378

PALAEONTOLOGY, VOLUME 12

Locality

Mansel

!

j

I

*f

Bouton

Bouton

La Fdrisienne
W end

specimen s
in sample

58

text-fig. 7. Histogram of the population structure of the Upper Beds of the
Calcaire de Ferques. Values expressed as in text-fig. 2.

of Middlemiss (1962) that extensive transportation would cause almost complete
destruction of brachiopod shells. This is confirmed by many workers on Recent faunas,
especially on molluscs (Schafer 1962, Holme 1961, and Johnson 1965). Thus it may be
concluded that if the assemblage has been transported, it was only over a very short
distance and had little disruptive effect on associations. Hence the fauna is still representa-
tive of a life assemblage.

It is suggested that the fauna lived and accumulated in water of shallow to moderate
depth (i.e. below wave base but not greater than 10 fathoms). A catastrophic event such

P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 379

as a storm subsequently transported both fauna and matrix to its present position where
it was not subject to currents and was quickly cemented, first by algae and then by
crystalline calcite. This interpretation is supported by the field relationships of the bed.
It is thin and developed only relatively locally, being confined to the eastern end of the
outcrop. It immediately overlies a formation whose environment of deposition has been
interpreted as lagoonal (Wallace, in press), the uppermost beds of which are a black
and yellow sandy dolomite containing branching stromatoporoids, a facies interpreted
by Lecompte (personal communication) as ‘sub-lagoonaf. This facies in fact occurs

TABLE 2.

Significant associations in the Upper Beds, as proved by the test.

P

0001

+ ve

Cyrtospirifer > 30 /JAthyris < 30%

001

+ ve

Gastropod /Athyris < 30%

001

+ ve

Schizophoria/ fan bryozoan

0015

+ ve

Rhynchonellids/solitary corals

004

+ ve

Solitary corals \Productella subaculeata

005

+ ve

Schuchertella /gastropod

006

+ve

Gastropod/ Cyrtospirifer > 30%

006

+ ve

Ptychomaletoechia/ Productella subaculeata

010

— ve

Schuchertella/Schizophoria

0-20

+ ve

Tenticospirifer/strophomenids

0-20

+ ve

Productella subaculeata/ strophomenids

0-20

+ ve

Sch iiopho ria / s t ro p h o m e n i d s

0-20

+ ve

Schuchertella/massive corals

0-20

+ ve

Schizophoria/ massive corals

0-20

— ve

Productids, 'gastropods

both below and immediately above the bed, leaving the anomalous situation of a thin
limestone with a good marine fauna sandwiched between lagoonal, possibly strongly
saline, dolomites. It would seem likely that the Thin Basal Limestone thus represents a
marine association washed, perhaps by a sudden storm, into lagoons, which in a short
time were themselves swamped by a sudden marine transgression which laid down the
limestones of the main part of the Calcaire de Ferques.

The higher argillaceous content and lower bioclastic content of the Upper Beds of
the limestone at once suggest a quieter, possibly deeper, environment of deposition than
that of the main part of the limestone, but this initial impression may be erroneous
since they pass up to shales and sandstones interpreted as the products of an intertidal
(and just subtidal) environment (Wallace, in press). The excellent preservation of much
of the fauna and especially that of the atrypid frills, however, indicates that little or no
reworking of the sediment, either by currents or by burrowing organisms, took place,
and that the environment was extremely quiet.

The increase in size of many of the species is most marked in those brachiopods which
presumably were attached by a pedicle during life, and may be a response to the in-
creased argillaceous content of the sediment. It is noticeable, however, that the size
increase does not coincide exactly with the increase in clay content, but is first observed
some metres below; thus the two may not be connected. These large brachiopods do
not show the crowding of growth-lines near the anterior commissure usually considered
as indicative of adulthood, a feature which is seen in smaller specimens from other parts

380

PALAEONTOLOGY, VOLUME 12

of this formation and from other formations in the area. Ager (1963, p. 142) has sug-
gested that in exceptional circumstances ‘gigantism’ in fossils may be a result of delayed
sexual maturity. This effect has also been observed in several living vertebrates, largely
by inhibition of the action of the thyroid (Comfort 1965, pp. 85-6). It is interesting,

p = 001

01 < P < 05

05 < p < 10

10 < p < 25

text-fig. 8. Constellation diagram of positive associations in the Upper Beds of the Calcaire de
Ferques. Symbols as in text-fig. 3. Ath, A thyris ; Cyr, Cyrtospirifer, D/N, Douvi/lina/Nervostrophia ;
Ga, Gastropod; Pro, Prodactellci subaculeata; Pty, Ptychomaletoechia; Schi, Schizophoricr, Schu,

Schuchertel/a

therefore, that this gigantism occurs most markedly in those species which have a
high proportion of pathologically deformed individuals. The two phenomena may be
connected, possibly through an external influence such as increased background radio-
activity affecting the genetic balance of certain species.

The authorship of species mentioned in this paper is as follows: Cyrtospirifer verneuili (Murchison);
Tenticospirifer tenticulum (de Verneuil); Cyrtina heteroclita (Defiance) ; Spinatrypa «.s/;<?ra (Schlotheim) ;
S. explanata (Schlotheim); S. longispina (Bouchard); Atrypa squamifera (Schnur); Cupularostrum
recticostcitum Sartenaer; Ptychomaletoechia boloniensis (Oehlert); Athyris concentrica von Buch;
Terebratula hastata Phillips; Productella subacideata (Murchison); P. dutertrei Rigaux; Chonetes
laevis Davidson; C. douvillei Rigaux; Sieberella brevirostris (Phillips); Schizophoria striatula (Schlot-
heim); Pseudaviculopecten striatus (Hall).

Acknowledgements. The author would like to thank Mr. J. D. MacDougall and her sister Alexandra
for assistance in testing the validity of the sampling methods.

P. WALLACE: INDICATORS IN TWO HORIZONS OF CALCAIRE DE FERQUES 381

REFERENCES

ager, d. v. 1961. The epifauna of a Devonian spiriferid. Q. Jl geol. Soc. Lond. 117, 1-10.

1963. Principles of Paleoecology. 371 pp. McGraw-Hill, New York.

— and Wallace, p. 1967. The environmental history of the Boulonnais, France. Proc. Geol. Ass.
11 (for 1966), 385-417.

boucot, a. j. 1953. Life and death assemblages among fossils. Am. J. Sci. 251, 25-40.
comfort, a. 1965. The Process of Ageing. 152 pp. Weidenfeld and Nicolson, London.
godwin-austen, r. a. 1853. On the series of Upper Palaeozoic groups in the Boulonnais. Q. J! geol.
Soc. Lond. 9, 231-45.

gosselet, j. a. a. 1860. Memoire sur les terrains primaires de la Belgique des environs d'Avesnes et du
Boulonnais. 164 pp. Martinet, Paris.

1894. Etude sur les variations du Spirifer Verneuili. Mem. Soc. geol. N. 4(1), 61 pp., 7 pi.

hallam, a. 1967. The interpretation of size-frequency distributions in molluscan death assemblages.
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halloy, d’o, d’. 1862. Abrege de geologie. 7th edn.

holme, N. a. 1961. The bottom fauna of the English Channel. J. mar. biol. Ass. 41, 397-461.

Johnson, r. g. 1962. Interspecific associations in Pennsylvanian fossil assemblages. J. Geol. 70, 32-55.

1965. Pelecypod death assemblages in Tomales Bay, California. J. Paleont. 39, 32-55.

kershaw, k. a. 1966. Quantitative and Dynamic Ecology. 2nd edn., 183 pp. Edward Arnold, London.
lecompte, m. 1961. Facies marins et stratigraphie dans le Devonien de la Belgique. Annls Soc. geol.
Belg. 85, B 17-57.

middlemiss, f. a. 1962. Brachiopod ecology and Lower Greensand palaeogeography. Palaeontology, 5,
253-67.

murchison, r. i. 1840. Sur les roches devoniennes du Boulonnais et des pays limitrophes. Bull. Soc.
geol. Fr., lre ser., 1 1, 229-57.

pruvost, p. and pringle, j. 1924. A synopsis of the geology of the Boulonnais including a correlation
of the Mesozoic rocks with those of England, with report of excursion. Proc. Geol. Ass. 35, 29-67 .
robinson, j. w. d. 1920. The Devonian of Ferques (Lower Boulonnais). Q. Jl geol. Soc. Lond. 76,
228-36.

Schafer, w. 1962. Actuo-Palaontologie nach Studien in der Nordsee. 666 pp. W. Kramer, Frankfurt.
senes, j. 1967. Repartition bathymetrique des algues fossilisables en mediterranee. Geol. Sb., BratisL,
18, 141-50.

vandercammen, a. 1959. Essai d’etude statistique des Cyrtospirifer du Frasnien de la Belgique.
Mem. Inst. r. Sci. nat. Belg. 145, 1-175.

verneuil, e. de. 1838. Note sur les terrains anciens du Bas-Boulonnais. Bull. Soc. geol. Fr., lre ser., 9,
388-96.

Wallace, p. 1968. The sub-Mesozoic palaeogeology and palaeogeography of northeast France and
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(in press). The sedimentology and palaeoecology of the Devonian of the Ferques inlier, northern

France. Q. J! geol. Soc. Lond.

PEIGI WALLACE

Department of Geology
Imperial College
Prince Consort Road
London, S.W.7

Typescript received 9 July 1968

ON THE STRUCTURE AND RELATIONSHIPS OF
A NEW PENNSYLVANIAN SPECIES OF THE
SEED PACHYTESTA

by THOMAS N. TAYLOR and DONALD A. EGGERT

Abstract. Pachytesta berryvillensis sp. nov., a small trigonocarpalean seed most closely resembling P. pitsilla ,
is described and figured from petrifaction material of Upper Pennsylvanian age collected from southeastern
Illinois.

Two coal balls collected from the Mattoon Formation (of late Pennsylvanian age) near
Berryville, Illinois have revealed 52 structurally preserved seeds of a new type. The seeds
are ovoid, radially symmetrical, and measure 6T-6-8 mm. long and 5-0-5-2 mm. in
maximum diameter. Zonation of the integument is delimited into 3 principal layers;
an outer parenchymatous sarcotesta containing sclerenchyma cells and numerous secre-
tory canals, a middle 2-layered sclerotesta distinguished by the size and disposition
of the individual fibres, and cuticular remains believed to represent remnants of an inner-
most endotesta. Nucellar attachment to the integument is confined to a small pad in the
seed chalaza. Vascular tissue is present in the nucellus, but not observed in the integu-
ment.

SYSTEMATIC DESCRIPTION

Genus pachytesta Brongniart 1874
Pachytesta berryvillensis sp. nov.

Plate 71, figs. 1-6; Plate 72, figs. 1-5; text-fig. 1

Diagnosis. Ovoid seeds 6T-6-8 mm. long and 5-0-5-2 mm. in diameter with rounded
base and short micropylar tube. Integument with three discernible layers; outer
sarcotesta of loosely arranged, thin-walled cells and longitudinally directed secretory
canals, bounded by isodiametric cells with thickened walls and dark contents. Middle
sclerotesta consisting of an outer zone of fibre-like cells one cell deep and oriented at
right-angles to seed axis, and inner zone of identical fibres showing longitudinal orienta-
tion, occasional fibres of inner zone radially disposed and projecting into seed cavity.
Endotesta represented by cuticle. Integument differentiated into three primary ribs,
well-developed in apex, recognizable as thin areas in the sclerotesta at mid-level and
chalaza. Nucellus attached to integument only at base by small disc TO mm. high and
1-2 mm. in diameter, apically differentiated into pollen chamber and nucellar beak.
Nucellar vascular system of 28-35 flattened bundles that terminate at pollen chamber
base; vascular configuration of integument unknown. Megaspore membranes ovoid
and granulose.

Holotype (Plate 71, fig. 1). Slides and peels of Coal Ball 3687 A(5), Paleobotanical Collections, Depart-
ment of Biological Sciences, University of Illinois at Chicago Circle.

[Palaeontology, Vol. 12, Part 3, 1969, pp. 382-7, pis. 71-72.]

T. N. TAYLOR AND D. A. EGGERT: THE SEED PACHYTESTA 383

Paratypes. Slides and peels of Coal Ball 3687 B-D, and Coal Ball 735 in above collection.

Type locality. Berryville (Sec. 7, T 2 N, R 1 3 W), Sumner Quadrangle, Lawrence County, Illinois.
Stratigraphic position. Calhoun Coal, Mattoon Formation, McLeansboro Group.

Age. Late Pennsylvanian.

text-fig. 1 . Median longitudinal section of immature pollen chamber showing fleshy nature of chamber
walls and floor. (The illustration shows only the pollen chamber of the nucellus, no megaspore mem-
brane is illustrated.)

Description.

Integument. The integument consists of three tissue systems. The outer layer, or sarco-
testa, measures up to 1-6 mm. in thickness in the chalazal region (PI. 71, fig. 1 ; PI. 72,
fig. 1), thins to several hundred microns near the mid-level and apex (PI. 71, figs. 4, 5).
It is constructed of thin-walled cells that show variability in both size and shape (PI. 71,
fig. 4; PI. 72, fig. 1). Cells disposed toward the outer limits of the integument are larger
and have more irregular shapes, while those cells more deeply positioned in the layer
are smaller and have more uniform diameters. In the best-preserved specimens the
sarcotesta is bounded by a variable number of layers of isodiametric cells with thickened
walls and dark lumen contents (PI. 71, fig. 4).

Numerous secretory canals up to several millimetres long further delimit the sarco-
testa (PI. 71, fig. 1 ; PI. 72, fig. 5). These canals that lack septa parallel the long axis of
the seed and are most abundant near the periphery of the tissue. The arrow in PI. 72,
fig. 6 shows the position of a 1-2 cell thick layer that is often noticeable between the
inner limits of the sarcotesta and the outer limits of the sclerotesta.

c c

C 6685

384

PALAEONTOLOGY, VOLUME 12

Throughout the length of the seed a uniformly thickened sclerotesta constitutes the
middle zone of the integument. Distally this layer is attenuated into a short micropylar
tube 0-5 mm. long (PI. 71, fig. 1); in the chalaza it extends 0-3 mm. into the seed cavity
to form the nucellar disc (PI. 71, fig. 1 ; PI. 72, fig. 1).

The outer portion of the sclerotesta, approximately one-third of the total thickness
of this layer, consists of thick- walled fibres averaging 76 x 17 /x whose longitudinal axes
are oriented perpendicular to the long axis of the seed ( PI. 71, fig. 2; PI. 72, figs. 3, 6).
The inner segment of the sclerotesta is formed of similar fibres, however, these cells
show an orientation pattern that parallels the seed axis (PI. 72, figs. 3, 6). Fibres of the
inner limits of this layer have lobes that form a series of inwardly facing projections
that sculpture the inner surface of the sclerotesta (PI. 71, fig. 2; PL 72, figs. 3, 6). These
projections from the fibres of the inner surface form a reticulum surrounding large
lacunae which appear as conspicuous spaces in tangential section (PI. 71, fig. 6).

A supposed inner flesh or endotesta, that constitutes the third component of the seed
coat, is not preserved except for cuticular remains that apparently bounded this layer.
This cuticle is seen in Plate 72, fig. 3 and fig. 6 (upper arrow) extending between the tips
of the inwardly facing sclerotestal fibres. Near the base of the seed the cuticle of the
endotesta becomes continuous with the outer cuticle of the nucellus. Orientation of the
cuticular remains suggests that at some stage in the ontogeny of the seed a delicate
endotesta occupied the lacunae formed in the inner surface of the sclerotesta. This
tissue was either not preserved or became disorganized at some stage of the development
of the megasporangium prior to that present in the specimens.

Three equidistantly spaced primary ribs extend radially for the length of the seed. They
are prominent in the apex (PI. 71, fig. 4), but are recognizable in mid-level and chalazal
sections only as interruptions of the inner zone of the sclerotesta (PI. 71, figs. 2, 5).
Secondary ribs were not observed in any of the specimens.

Nucellus. The nucellus is continuous with the integument only at the chalaza where it is
attached to a small disc-like extension of the sclerotesta that forms the nucellar disc (PI.
71, fig. 1 ; PI. 72, fig. 1). Early in development the apical portion of the nucellus forms a
conical pollen chamber and nucellar beak that presumably engaged the lower orifice of the
micropylar tube to ensure that pollen entered the chamber (text-fig. 1). In most of the
specimens the nucellus is crushed and separated from the inner limits of the integument
(PI. 71, fig. l).In some specimens a cutinized epidermis is all that remains of the nucellus.

In several seeds, however, the tissue of the nucellus is thick and shows little distortion.
A median longitudinal section of the pollen chamber of one of these seeds is illustrated
in text-fig. 1. It has a maximum diameter of 2-2 mm. and measures 1-2 mm. high. The

EXPLANATION OF PLATE 71

Figs. 1-6. Pachytesta berryvillensis sp. nov. (n = nucellus, pc = pollen chamber, r = rib, sar = sar-
cotesta, scl = sclerotesta). 1, Near median longitudinal section of seed; note presence of pollen
grains in pollen chamber; C.B. 3687 A(5)bot No. 13, x 12. 2, Transverse section below mid-level;

C.B. 3687 C(2)bot No. 4, Xl6. 3, Transverse section of chalaza showing position of vascular
bundle (arrow) and disposition of sclerotesta fibres; C.B. 3687 C(2)bot No. 8, X 22. 4, Transverse

section of seed apex showing nature of ribs and integument zonation; C.B. 735 E(2)top No. 7,
X20. 5, Mid-level transverse section showing position of ribs (arrows); C.B. 735 C(2)bot No. 25,
X 10. 6, Tangential section of integument showing lacunae between sclerotesta fibres; C.B. 735

F(2)top No. 5, X 20.

Palaeontology, Vol. 12

PLATE 71

TAYLOR and EGGERT, Pennsylvanian seed Pachytesta

T. N. TAYLOR AND D. A. EGGERT: THE SEED PACHYTESTA 385

pollen chamber cavity is 0-7 mm. in diameter and 0-6 mm. high. The walls and pollen
chamber floor are thick and constructed of exceptionally thin-walled cells. Epidermal
cells of the distal end that will ultimately form the nucellar beak have only slightly
thicker radial walls. In these specimens the disorganization and breakdown of cells
at the distal end of the nucellus to form the pollen chamber had apparently only pro-
ceeded a short time prior to fossilization.

The megaspore membrane is thin and characterized by a granulose surface pattern.
Pollen. Approximately 40 pollen grains were observed in the pollen chambers of several
seeds (PI. 72, fig. 2). The largest number observed in a single chamber was 17. All of the
grains are similar and consist of a psilate central body surrounded by a slightly elliptical
bladder with reticulations on the inner surface (PI. 72, fig. 2, 4). Diameter (greatest
length of the bladder) varies from 72 to 81 p. If compared with dispersed small spores
the pollen grains present in the chamber of Pachytesta berryvillensis would be included
in the genus Florinites as interpreted by Schopf, Wilson, and Bentall (1944). Pollen or
pre-pollen grains have now been reported in the following species of Pachytesta : P.
incrassata (Brongniart 1881), P. gigantea (Renault 1896, Taylor 1965), P. vera (Hoskins
and Cross 1946, Taylor 1965), P. hexangu/ata (Stewart 1951), P. illinoensis (Stewart — -
personal communication), P. pusilla (Brongniart 1881), and P. berryvillensis reported
here. Of these seeds P. gigantea, P. hexangulata, P. illinoensis , and probably P. incrassata
all have large bilaterally symmetrical pre-pollen grains of the Monoletes type. Such
grains are known to have been produced by some members of the Pteridospermales.
Pachytesta berryvillensis and P. vera on the other hand contain pollen of the Florinites
type that is known from a large number of pollen organs assignable to the Cordaitales.
Whether the presence of these Florinites grains in seeds demonstrated as belonging to
the Medullosan seed ferns is merely the result of contamination at the time of pollina-
tion, or reflects the occurrence of another group of Palaeozoic plants that have seed fern-
like ovules and Cordaitalean-like pollen grains must await the discovery and report of
additional seeds containing pollen grains and the determination of the biological affinities
of Florinites.

Vascularization. A single vascular bundle penetrates the integument (PI. 71, fig. 3;
PI. 72, fig. 1) and passes into the nucellus where it vascularizes that tissue. At a level
of approximately TO mm. above the point of attachment between nucellus and integu-
ment the nucellar vascular bundles form an almost continuous ring similar to the tracheal
sheath that delimits seeds of the Stephanospernmm type. At higher levels the vasculature
of P. berryvillensis consists of a variable number (28-35) of closely spaced, but clearly
distinct flattened bundles that continue to the region of the pollen chamber. Tracheids
are variable in size and possess scalariform-helical secondary thickenings.

Although several scattered tracheids were observed in association with the sarcotesta,
the general poor preservation of this tissue makes it impossible at this time to determine
whether the integument of P. berryvillensis was vascularized, and if so, the number and
disposition of the bundles.

DISCUSSION

The genus Pachytesta as currently interpreted includes radially symmetrical seeds
having a rounded base and attenuated apex, a multiple-layered integument consisting
of three principal zones, three primary ribs with secondary and tertiary ribs present or

386

PALAEONTOLOGY, VOLUME 12

absent, a nucellus free from the integument except at the base and apically forming a
large pollen chamber, and vascular tissue in the form of distinct bundles in both nucellus
and integument.

Of the 13 recognized species (Taylor 1965) size varies from ovoid seeds 7-0 mm. long
and 4-0 mm. in diameter (P. pusilla) to obovoid forms 11-0 cm. long and up to 6-0 cm.
in diameter (P. incrassata). Variability of the three principal tissues of the integument
(sarcotesta, sclerotesta, endotesta) ranges from those examples where little differentiation
is present in any of the zones (e.g. P. stewartii ) to those forms where the histology is
extremely complex (e.g. six recognizable zones of the sarcotesta in P. saharasperma).
The arrangement of the individual fibre-like cells that make up the sclerotesta are in
three general patterns that have been described by the terms tangential, radial, and
intermediate. These three types of sclerotesta are exemplified by P. illinoensis, P.
hexangulata , and P. hoskinsii respectively.

Radial extensions of the sclerotesta that form the primary ribs may be prominent
with parenchymatous trabeculae as in P. vera , or may be absent and marked only by
indentations or thin areas in the corresponding position within the sclerotesta (e.g.
P. gigantea). Secondary and tertiary ribs may number as many as 42 in P. illinoensis
or be totally absent as demonstrated by P. stewartii. Vascularization of the integument
may vary from 6 bundles in P. shorensis to 54 in P. incrassata. In two previously described
species (P. hoskinsii and P. pusilla) the pattern and number of integumentary vascular
bundles is unknown. The integumentary vascular pattern varies from 8 bundles in
P. olivaeforniis to more than 40 in specimens of P. gigantea. Nothing is known about the
nucellar vascular system of P. noei, and P. incrassata, while in P. pusilla only the presence
of nucellar tracheids has been reported.

When compared with other species of Paehytesta, P. berryvillensis appears most
similar to P. pusilla known only from the Late Pennsylvanian equivalent of France.
Size range for the few specimens reported (Brongniart 1874, Oliver 1904) includes
seeds 6-5— 8-5 mm. long and 4-0-5-0 mm. in maximum diameter with the distal end
attenuated into a short micropylar tube. Thin-walled, isodiametric parenchyma cells
characterize the very incompletely preserved portion of the sarcotesta. The sclerotesta
is similar to that of P. berryvillensis, in that an outer palisade-like layer, and an inner
layer of longitudinally oriented fibres are present. In P. pusilla radially disposed fibres
are absent from the inner surface of the sclerotesta. The presence of an endotesta is
known only from cuticular remains in both species. Vascularization of the nucellus is

EXPLANATION OF PLATE 72

Figs. 1-6. Paehytesta berryvillensis sp. nov. (nd nucellar disc, sar = sarcotesta). 1, Median
longitudinal section of seed chalaza showing position of vascular bundle (arrow) and extent of
sarcotesta; note secretory canals at right of photograph; C.B. 3687 A(5)bot No. 17, x21. 2,

Several Florinites pollen grains within chamber; C.B. 735 C(2)bot No. 70, X250. 3, Longitudinal

section of sclerotesta showing orientation of cells and inward projecting fibres of inner layer; C.B.
3687 A bot No. 12, Xl20. 4, Florinites pollen grain observed on peel preparation; C.B. 735

E(2) top No. 3, x 375. 5, Tangential section of sarcotesta showing longitudinally directed secretory

canals; C.B. 3687 B(l) top No. 27, X 16. 6, Transverse section of sclerotesta showing disposition

of cells and inwardly projecting fibres; upper arrow indicates position of endotesta cuticle and
lower arrow position of thin-walled layer between sarcotesta and sclerotesta; C.B. 735 FI(2)bot
No. 16, X 120.

Palaeontology, Vol. 12

PLATE 72

TAYLOR and EGGERT, Pennsylvanian seed Pachytesta

T. N. TAYLOR AND D. A. EGGERT: THE SEED PACHYTESTA

387

reported in P.pusilla, however, no bundle number is given and nothing is known regard-
ing a vascular system in the integument.

The unique occurrence of radially disposed fibres making up the inner layer of the
sclerotesta of P. berryvillensis extends an already broad range of anatomical variability
of the integument of seeds assignable to the genus Pachytesta. To our knowledge the
only other seed that shows integument ornamentation of the inner surface is Albertlongia
incostata (Taylor 1967) from the Middle Pennsylvanian of Illinois. Specimens of this
taxon are radially symmetrical and measure 2-0 cm. long and 1-4 cm. in diameter.
The apex is rounded; the base extended into a pedicel 2-2 mm. long. Features in com-
mon with the genus Pachytesta include a heterogeneous integument of three principal
tissues, a double vascular system, and a nucellus that is attached to the integument
only at the base. In Albertlongia the inner surface of the sclerotesta appears as an uneven
series of broad ridges and furrows that extend from apex to chalaza, whereas the in-
dividual ornamentations of P. berryvillensis are the result of relatively few inwardly
projecting fibres. The consistent absence of primary ribs of the integument either in the
form of prominent radial extensions, or in the presence of indentations or thin areas
in the sclerotesta of Albertlongia , currently serves to distinguish the genera Pachytesta
and Albertlongia.

Our ability to interpret the significance of internal ornamentation patterns in the
integument of Palaeozoic seeds is extremely limited in that at present we can not properly
distinguish between those features which are consistent throughout ontogeny of a given
species versus those that changed during the course of development. The ability to
distinguish phylogenetic features from ontogenetic stages must await the discovery of
additional seeds that will adequately document the course of Palaeozoic ovule develop-
ment.

Acknowledgement . This research was supported by National Science Foundation Grants GB-6834 and
GB-4126.

REFERENCES

brongniart, a. 1874. Etude sur les graines fossiles trouvees a Fetat silicifie dans le terrain houiller
de Saint-Etienne. Deuxieme partie. C. r. Acad. Sci. (Paris), 79, 427-35.

1881. Recherches sur les graines fossiles si/icifiees. Paris.

hoskins, J. H. and cross, a. t. 1946. Studies in the Trigonocarpales. Pt. 1. Pachytesta vera, a new
species from the Desmoinesian Series of Iowa. Amer. Midi. Nat. 36, 207-50.

Oliver, f. w. 1904. Notes on Trigonocarpus Brong. and Polylophospermum Brong., two genera of
Palaeozoic seeds. New. Phytol. 3, 96-104.

Renault, b. 1896. Bassin houiller et permien d’Autun et d'Epinac. Fasc. 4. Flore fossile. Deuxieme
partie (Text) 1-578, Etudes gites miner. France.

schopf, j. m., wilson, l. r., and bentall, r. 1944. An annotated synopsis of Paleozoic fossil spores
and the definition of generic groups. Rep. Inst. III. St. Geol. Surv. 81, 1-73.
stewart, w. n. 1951. A new Pachytesta from the Berryville locality of southeastern Illinois. Amer.
Midi. Nat. 46, 717-42.

taylor, t. N. 1965. Paleozoic seed studies: a monograph of the American species of Pachytesta.
Palaeontographica, B1 17, 1-46.

1967. Paleozoic seed studies: on the structure of Conostoma leptospermum n. sp., and Albert-
longia incostata n. gen. and sp. Ibid. B121, 23-9.

THOMAS N. TAYLOR
DONALD A. EGGERT
Department of Biological Sciences
University of Illinois at Chicago Circle
Chicago, Illinois

Typescript received 17 September 1968

THE ONTOGENY OF THE THECIDEACEAN
BRACHIOPOD MOORELLINA GRANULOSA
(MOORE) FROM THE MIDDLE JURASSIC OF

ENGLAND

by P. G. BAKER

Abstract. Investigation of fifty-five brachial valves and several sectioned shells from a new locality has led
to the conclusion that in Moorellina granulosa the brachial apparatus shows progressive elaboration from simple
tubercles to very delicate convoluted lobes which are in the form of a ptycholophe. Also that Thecidium forbesi
Moore (1855) should be regarded as an early pre-brachial lobe stage of M. granulosa.

Five ontogenetic stages are described. The argument for an ontogenetic series is strengthened by the changes
occurring in the cardinal process, coincident with the development of the brachial apparatus. Reference is
made to the improbability of finding detached brachial valves showing adult characteristics. It is considered
that the generic distinction between Moorellina Elliott (1953) and Rioultina Pajaud (1966) should be based on
the form of the brachial lobes and not merely on their presence or absence. Specific determination based on the
morphology of the brachial valve is considered unsatisfactory and a technique has been developed for the study
of the internal characters of complete specimens.

The paper records the occurrence of thecidellinids in the Oolite Marl of the Mid-
Cotswolds. The deposit is rather variable and outcrops at a number of localities. It is
well developed on Cleeve Cloud near Cheltenham where it attains a thickness of over
4 m. The Marl is typically a pale, cream-coloured marl, relatively harder and more
oolitic in the upper layers, softer and with the ooliths more scattered towards the base.
It thins in a north-easterly direction towards Chipping Campden and changes litho-
logically in a south-westerly direction, becoming indurated south from Painswick
until it is indistinguishable from the overlying limestone.

Stratigraphically the Oolite Marl was placed by Arkell in the Lower Inferior Oolite
(Upper Aalenian, murchisonae zone). It rests on a clearly recognizable bored surface
of Lower Freestone but cannot be structurally separated from the overlying Upper
Freestone, into which it passes laterally in the vicinity of Stroud.

The material described in this paper was obtained from the northernmost outcrop
of the Oolite Marl two miles to the south of Chipping Campden at Westington Hill
quarry, grid. ref. SP 142368. The Marl occurs at the top of the quarry which exposes
almost 8 m. of Lower Freestone. The collection was made from a bed of soft, pale marl
30 cm. thick, which occurs in the north face of the quarry between two harder bands
and about 60 cm. above the top of the Lower Freestone.

The thecidellinids from this locality occur together with a variety of organo-detrital
remains which are recorded in tabular form in the text. This fauna is essentially similar
to the surf-zone fauna recorded by Nekvasilova (1967) (Nekvasilova in Ager 1965,
p. 146). The occasional large brachiopods which occur are forms which are anatomically
characteristic (Ager 1965) of peri-reefal deposits.

Acknowledgements. The author is indebted to Dr. J. D. Hudson, Geology Department, University of
Leicester and Dr. H. Torrens, Geology Department, University of Keele, for helpful advice during the

[Palaeontology, Vol. 12, Part 3, 1969, pp. 388-399, pis. 73-74.)

P. G. BAKER: BRACHIOPOD MOORELLINA GRANULOSA (MOORE) 389

preparation of this paper. Thanks are due to Mr. M. Talbot, Geology Department, University of
Bristol and Dr. L. R. M. Cocks, Department of Palaeontology, British Museum (Natural History)
for the loan of Corallian and Inferior Oolite material for comparison purposes; to Mr. G. McTurk
for preparing the stereoscan negatives; to Dr. G. F. Elliott for comments concerning certain mor-
phological features; to Dr. A. W. Medd, Instititue of Geological Sciences for identifying the polyzoa
fragments and finally to Professor P. C. Sylvester-Bradley for use of the research facilities of the
University of Leicester.

Registration of Material. The specimens figured and referred to in this paper are to be deposited in the
museum collection of the Department of Geology, University of Leicester. The specimen numbers
quoted refer to their catalogue numbers.

TABLE 1

Residue retained on sieve mesh

Residue type

2mm. mesh

422p

mesh

152p. mesh

A v. ten

Av. °/0

Av. ten

Av. %

Av. ten

>

<

=-°

5gm.samps.

sample

5gm. samps

sample

Igm.samps.

sample

Oolite fragments

3-8

4-7

40-8

0-78

-

-

Ooliths

—

—

1198-8

22-9

148-8

0-8

Brachiopod

69-9

84-3

3555 0

68-0

shell fragments

1 7064-0

98-4

Bivalve

shell fragments

3-8

4-7

45-0

0-84

Echinoderm debris

1-8

2-2

21 4-8

4-1

64-5

0-3

Gastropods

-

-

0-3

0 006

-

-

Thecidellinids

0-2

0-24

14-4

0-27

2-6

001

Other brachiopods

1 -4

1 -6

13-8

0-26

5-7

0-03

Ostracods

-

-

8-4

0-16

38-9

0-2

Polyzoa fragments

1-7

20

112-2

2-1

2-0

001

Annelid worms

0-4

0-48

19-1

0-36

3-5

001

Total

830

5222-6

17330-0

PREPARATION OF MATERIAL

Weathered marl samples were dried and crumbled through a 6-mm. sieve to remove
large fragments. The sieved material was immersed in water and cleaned for ten minutes
in a Dison electrosonic cleaner, marketed by Headland Engineering Developments Ltd.
The samples were then washed until a clean residue was obtained. This was dried and
passed through 2-mm., 422-ju,, and 152-ju, sieves, each residue size being analysed for
faunal content (Table 1). Experience has shown that only the material retained on the
422-^ mesh is likely to yield thecidellinids. The cleaned residue was hand-picked under
a binocular microscope and the brachiopods transferred to glass tubes for further
cleaning. The individual tubes were replaced in the cleaner for periods of 10-20 seconds
until the matrix had been removed. The shells were examined at each phase of cleaning

390

PALAEONTOLOGY, VOLUME 12

to determine whether the treatment should be continued. It should be emphasised that
this is a very slow method of collecting as the thecidellinid content of the residue is
approximately 0-3%. The collection of 172 specimens is comprised as follows:

Complete specimens selected for sectioning were cemented to glass slides, using a
mixture of Lakeside cement (obtainable from Cutrock Engineering Co.) and chloroform.
This remains plastic for sufficient length of time to allow correct orientation of the
specimen under a binocular microscope. When the cement has hardened, the orientation
of the specimen is checked and a plasticine mould is built round it. The mould is then
filled with Ceemar cold setting resin, which, when hardened, allows the block (attached
to the slide) to be serial sectioned on a Croft parallel grinder (see Hendry el al. (1963)
for other methods). Although the theory of sectioning such small specimens at 0-02 mm.
intervals is sound it is found to be inaccurate in practice. Better results are obtained if
acetate peels are taken after a standard number of 30-50 revolutions (depending on the
area of the block face). The length of the specimen is divided by the number of peels
obtained, thus averaging out any error.

Residue obtained by the method described was found to constitute approximately
40% of the Oolite Marl from the Westington Locality. An analysis of the composition
and faunal content of the cleaned residue is shown in Table 1.

In all size ranges, the bulk of the residue is composed of brachiopod shell fragments
which may be identified as rhynchonellid and terebratuloid. Occasional specimens of
Globirhynchia subobsoleta (Dav.), Epithyris submaxillata (Morris), and Plectothyris
fimbria (Sow.) occur in the coarse residue and it is probable that the shell debris was
formed from the remains of these species. The bivalve shell fragments may be identified
as Ostrea and Lopha species. The echinoderm debris consists of unidentified echinoid
spines and plates, crinoid ossicles of Pentacrinus type, unidentified ophiuroid plates
and vertebrae. Three very small gastropods of Nerinea type were recovered from the
422-p sample. The polyzoa fragments have been identified by Dr. A. W. Medd as
Actinopora sp., Berenecea sp., MeJieeritites sp., and Spiropora sp. The ostracods are
Bairdia sp., Cytherella sp., and several unidentified cytheracean genera.

The analysis probably shows a bias towards brachiopod shell fragments as the true
nature of the shell type was frequently obscured by adherent matrix. It is possible there-
fore, that some fragments included in the brachiopod count may in fact have been
bivalve material. In addition, quite large echinoid spines appear in the 422-/x sample as
their diameter is the critical factor. In the 152-p. size range it was not possible to dis-
tinguish between brachiopod and bivalve shell material with any degree of accuracy and

Brachial valves

Pedicle valves

Complete specimens

Broken valves with significant detail

55

16

43

58

172

ANALYSIS OF RESIDUE

P. G. BAKER: BRACHIOPOD MOORELLINA GRANULOSA (MOORE) 391

they are therefore grouped. However, as the proportion of brachiopod shell material
is so large it is considered that the probable error is of no significance. Analysis of the
brachiopod content was very critical and it is unlikely that any specimens escaped
attention, even in the 152-jU, samples. Although the number of ostracods shows a marked
rise in this size range, they still represent approximately the same percentage of the sample.

The thecidellinid material shows the same ranges of variation of shell shape noted
by Elliott (1948) during his study of Bifolium and by Nekvasilova (1967) during her
study of Thecidiopsis. It appears to be correlated with the size of the area of attachment

0-5 10 1-5 2-0mm.

valve width

text-fig. 1. Histogram to show the size distribution of 126 brachial
valves of Moorellina granulosa (Moore) from Westington Hill Quarry.

Number of individuals plotted against width of the brachial valve.

and is almost certainly the result of environmental influence (Rudwick 1962). The
dangers of using a single character are evident but for the purpose of expressing size
distribution (text-fig. 1) width of the brachial valve is used, as this appears to be
one of the least variable growth characteristics. This feature is particularly useful as
over half the material consists of complete or broken brachial valves.

Scars of the area of attachment on the pedicle valve occasionally show portions
missing and occasionally, the pedicle valves are still cemented to shell fragments.
Nekvasilova (1967) has shown that the form of the area of attachment is consistent with
Thecidiopsis being attached, either to the valves of living oysters (liberated on the decay
of conchiolin), or settled directly to some firm rocky substratum. The author is in
agreement with her views on the ecology of Thecidiopsis and the evidence suggests that
Moorellina occupied a similar environment, that is, belonged to the epifauna of the
inner sub-littoral zone. This opinion is further supported by their association with
peri-reefal brachiopods and the nature of the organo-detrital remains with which they
are deposited (ref. Table 1).

A consideration of the evidence indicates that the collection represents a transported
death assemblage, the size-frequency distribution (text-fig. 1), showing moderate
positive skewness. This may be regarded as the product of a normal growth-mortality

392 PALAEONTOLOGY, VOLUME 12

rate if compared with the histograms plotted by Hallam (1967) in his work on molluscan
death assemblages.

The fragmentation of thecidellinids, other brachiopods, bivalves, and Polyzoa in-
dicates that the debris was formed in a turbulent environment. However, the lack of
abrasion of the fragments together with the persistence of occasional bridges and
brachial lobes in detached brachial valves of thecidellinids would suggest that the
organo-detrital remains were transported only a short distance. Reference to Table 1
will show that thecidellinids are most common in the 422-^ size range whereas ostracods
are most common in the 152-fi, size range. Brachiopod shell fragments represent a
high percentage of the material in all size ranges. These data and arguments advanced by
Hallam (1967) suggest that no significant sorting of the population has occurred. The
absence of appreciable numbers of small thecidellinids which would reflect the normally
expected high juvenile mortality rate may simply be the result of selective shell breakage
(Hallam 1967, p. 35). The presence of brachiopod shell fragments in excess of 2 mm.
diameter, the absence of large thecidellinids and particularly the absence of large theci-
dellinid shell fragments, is considered to indicate that thecidellinids with a brachial
valve width of more than 2 mm. are not represented in the population. Sedimentation
factors are not thought to have affected the upper size limit as ooliths larger than the
largest complete thecidellinid shells are common in the 422-p sample. Clearly, therefore,
the larger size-distributions of the sample may be regarded as adult shells. This is a
much smaller population than that studied by Nekvasilova (1967) but the ratio of
brachial valves to pedicle valves and complete shells within the sample is similar.

The material shows a number of progressive changes, such as, the development of the
cardinal process, the development of the border and the appearance and progressive
elaboration of the sub-peripheral rim. The brachial apparatus develops in the same
manner and, in the tuberculate stages, forms with from one to five pairs of brachial
tubercles are present. For reasons to be described later it is thought that some of these
latter are damaged adults. All the structures, with the exception of the adult brachial
lobes, show varying degrees of development in the material studied. This shows the
presence of an intimate relationship between the progressive development of the various
growth features with general size increase.

The collection of the material from one sample from a single bed, the size distribution
and the close parallel between the growth stages of Moorellina and the ontogenetic
development of Bifolium (Elliott 1948), Lacazella ( B .) lacazelliforme (Elliott) (Nek-
vasilova 1964), and Thecidiopsis (Nekvasilova 1967) leaves little room for doubt that
the material from Westington Hill quarry represents the ontogenetic stages of a single
species.

Comparison of the forms having a single pair of brachial tubercles with Thecidium
forbesi Moore 1855 indicates that there is no valid basis for the separation of T.forbesi
from Moorellina granulosa. It is proposed, therefore, that the adolescent growth stages
showing this degree of development of the brachial apparatus should be designated
forbesiform. It is considered that five ontogenetic stages may be recognized, each marked
by the appearance of a characteristic feature (text-fig. 3a-f). The dimensions recorded
are those of the specimens figured in Plate 73 and are intended to indicate only the general
size relationship, the stages represented being obviously subject to natural size varia-
tion.

P. G. BAKER: BRACHIOPOD MOO RELLINA GRANULOSA (MOORE)

393

Length mm.

Width mm.

Thickness mm.

A. Brephic valve

0-47

0-5

0-2

Pre-forbesiform stage

0-9

10

0-3

C. Early forbesiform stage

10

1-2

0-35

D. Late forbesiform stage

1-2

16

0-35

E. Post-forbesiform stage

1-4

1-7

0-4

F. Adult valve

1-1

13

0-4

MORPHOLOGY

The present paper is concerned only with the morphology of the interior of the brachial
valve (text-fig. 2a) but it is felt that some attempt should be made to reconcile the
interpretations of Elliott (1948) and Pajaud (1963) with the glossary laid out in the
Treatise on Brachiopoda but without adding to the existing confusion. Briefly the new
morphological terms and the justification for their introduction are as follows:

Border. The term is introduced to define the flat region of the valve between the sub-
peripheral rim and the periphery of the valve. This unites the limbe-marginal and bord
frontal of Pajaud and enables the distinction between anterior, lateral, and postero-
lateral regions to be made. It is felt that this is necessary as it is noted that the postero-
lateral border is the first to appear during the pre-forbesiform stage of development
(text-fig. 3b).

Brachial shelf. The term is introduced to define the raised flat areas, within the lopho-
phorian area, from which the brachial tubercles develop. The inner boundary of the
shelf is occasionally raised to form low ridges which may correspond, in part, to the
ridge extensions described by Elliott (1948) during the early ontogeny of Bifolium
faringdonense (Davidson).

Brachial tubercle. The dotted brachial ridge (Elliott) is not thought to be sufficiently
explicit. The rounded dots (Elliott 1948, p. 9) are by definition tubercles (Williams
1965, HI 54) and the term brachial tubercle is introduced as these structures are of
considerable importance in the development of the brachial apparatus.

Socket ridges. As defined in the glossary, inner and outer socket ridges are present (PI.
74, fig. 3).

The recommendation of the glossary (op. cit. H148) that the term marginal flange
should be replaced by sub-peripheral rim is adopted but it is noted that this has a
postero-lateral extension demarcating the outer boundary of the cardinal area. The
term brachial lobe introduced by Pajaud (19666) for the establishment of the genus
Rioultina is adopted, particularly as lobes are referred to in Moore’s original description
of MooreUina granulosa (Moore 1855). One must recognise however, that the brachial
lobes of M. granulosa are convoluted and not as in Rioultina , auriform.

Ontogenetic stages recognized. The brephic valve (text-fig. 3a; PI. 73, fig. 1) is the first
stage represented and is 0-5 mm. wide. The valve is almost circular, thick, and cap-like.
The cardinal margin is almost straight and approximately two-thirds of the width of
the valve. The cardinal process is the only recognizable thecidellinid feature, being low

394

PALAEONTOLOGY, VOLUME 12

and broad, occupying a little more than half the hinge-line and projecting slightly
beyond the cardinal margin (text-fig. 2b). There is no median septum but the postero-
lateral border is just discernible. The dental sockets are poorly developed but clearly
bounded anteriorly by the lateral divergence of the sides of the cardinal process, form-
ing inner socket ridges where they turn down to unite with the posterior margin of the
valve. No sub-peripheral rim is present at this stage. Three valves show this stage of
development.

Stage two is marked by the appearance of the median septum which is considered to
be a neanic feature (text-fig. 3b; PI. 73, fig. 2). The valve is still nearly circular, relatively
thick, markedly convex, and in the specimen figured, 1 mm. wide. The cardinal margin
is slightly curved and somewhat less than half the width of the valve. The cardinal
process is more strongly developed (text-fig. 2c) and turns down sharply into the floor
of the valve anteriorly. The postero-lateral border is now clearly defined, also the dental
sockets. The sub-peripheral rim is represented by a row of denticles which, posteriorly,
mark the inner boundary of the postero-lateral border, not, as in Elliott (1948), trending
inwards to form the sides of the cardinal process. Laterally the denticulate rim is almost
peripheral so that there is no noticeable border. It will be noticed that the bridge
extensions do not unite with the inner socket ridges and that they merge laterally with
the sub-peripheral rim. The median septum is fairly thick, developing from the anterior
margin and extending posteriorly, the posterior portion being free from the floor of the
valve. This stage is designated pre-forbesiform by virtue of the fact that the brachial
tubercles of T.forbesi Moore (1855) are not yet developed. Six valves show this stage of
development.

The two stages described correspond closely with the first two ontogenetic stages of
Bifolium faringdonense (Davidson) described by Elliott (1948) but the subsequent
stages showing the development of the brachial apparatus are quite different.

Stage three is marked by the appearance of the brachial tubercles characteristic of
T. forbesi and is accordingly designated the forbesiform stage, early and late features
being distinguishable (text-fig. 3c, d; PI. 73, figs. 3, 4).The valve is now slightly wider
than long. The cardinal margin is well-defined, slightly curved, and just less than half
the width of the valve. The cardinal process is by now a prominent feature, projecting
markedly beyond the cardinal margin. The dental sockets are deep and the postero-
lateral border inclined to the plane of the valve. The sub-peripheral rim is well developed
so that the lateral and anterior portions of the border are now visible. At this stage the

EXPLANATION OF PLATE 73

Figs. 1-6. Stereoscan photomicrographs of brachial valves of Moorellina granulosa (Moore) collected
from the Oolite Marl, Westington Hill Quarry near Chipping Campden. All the figures are interior
views of specimens coated with evaporated aluminium before photography. The bridge is broken
on specimens fig. 3-5. 1 . Brephic valve (37500), shell recrystallized. X 75. 2. Pre-forbesiform stage

of development (37503) showing the development of the median septum. Cardinal process slightly
damaged. X 50. 3. Early forbesiform stage (37505) showing the brachial shelf and appearance of

the first pair of brachial tubercles. X 50. 4. Late forbesiform stage (37507) showing four pairs of

brachial tubercles, well defined sub-peripheral rim and border. X 50. 5. Post-forbesiform stage

(37508) showing the expanded brachial tubercles, uniting posteriorly to form arches. Cardinal
process slightly damaged. X 50. 6. Adult valve (37509) showing the form of the right brachial

lobe. Cardinal process slightly damaged. X 50.

Palaeontology, Vol. 12

PLATE 73

BAKER, MooreUina granulosa

P. G. BAKER: BRACHIOPOD MOO RELLIN A GRANULOSA (MOORE) 395

bridge is usually complete (broken in the specimen figured during cleaning) but without
the marsupial notch (Elliott 1948). The posterior portion of the median septum is quite
clearly free from the floor of the valve and is usually slightly larger than that shown.
The floor of the valve is now characterized by two raised areas (brachial shelf), along

cardinal process
cardinal margin
postero - lateral sub-

peripheral rim '
lateral adductor L a Pi.
muscle sc

brachial shelf

brachial tubercle
median septum

border

outer socket

ridge

dental socket
inner socket

bridge rid9e
body cavity

brachial lobe

(reconstructed )

sub-peripheral

rim

text-fig. 2. a. Composite drawing from brachial valves 37509 and 37510 to show the morphological
features of the interior of the adult valve. Brachial lobe and median septum reconstructed from
specimen 37510. b-g. Drawings to show the form and modification of the cardinal process during
ontogeny, together with the development of the inner and outer socket ridges. b, brephic 37500.
c, preforbesiform 37503. d, e. early forbesiform 37505, 37506. f, late forbesiform 37507.

g, post forbesiform 37508.

the inner boundary of which the brachial tubercles develop, usually appearing in pairs.
Valves with one or two pairs of tubercles are said to constitute an early forbesiform
stage, whilst those with four to five pairs are said to constitute a late forbesiform stage.
Attention is drawn to the fact that the tubercles are regularly arranged and without
the random distribution figured by Pajaud (1966fi) in his work on material from the

396

PALAEONTOLOGY, VOLUME 12

Inferior Oolite of Dundry. During the early forbesiform stage the cardinal process
begins to lose its concavity posteriorly (text-fig. 2d, e) eventually forming a flattened
region which gives rise to the outer socket ridges during the late forbesiform stage
(text-fig. 2f; PI. 74, fig. 3). The bulk of the valves show this stage of development.

The beginning of stage four may be recognized by the fact that the distal ends of the
brachial tubercles expand and develop projections which ultimately unite with those
of adjacent tubercles to form delicate arches. This degree of organisation is taken to
represent a post-forbesiform stage (text-fig. 3e; PI. 74, fig. 1) The cardinal margin is, by
definition, a hinge line. The outer socket ridges, developed from the posterior of the
cardinal process are now clearly visible (text-fig. 2g; PI. 73, fig. 5). The median septum
extends back almost as far as the edge of the body cavity. The floor of the valve is
characterized by expanded brachial tubercles showing the development of arches
and inward pointing projections. The brachial shelf is now hardly visible, its decline
probably to be correlated with the higher development of the brachial apparatus. Five
valves show this stage of development.

In the adult state, stage five, the arches of the post-forbesiform stage are united,
thickened, and extended to form convoluted lobes (text-fig. 3f). Two specimens were
found at this stage of development, 37509 with the right brachial lobe intact (PI. 73,
fig. 6; PI. 74, fig. 2) and 37510 with the left brachial lobe intact. The brachial lobes of
both specimens could be seen to be joined to brachial tubercles anteriorly. An attempt
to clean specimen 37510 in order to provide conclusive evidence for the view that the
brachial lobes are attached to the valve solely by brachial tubercles, resulted in the loss
of the remaining brachial lobe. Specimen 37509 has therefore been subjected only to
partial cleaning, this in itself being sufficiently destructive to remove part of the bridge.
This specimen is remarkable in possessing a very young form cemented to its anterior
border in front of the median septum (PI. 74, fig. 4).

For reasons to be mentioned later, very little appears to be known about the adult
brachial valve of M. granulosa. The above specimens are almost certainly examples of
the type referred to by Davidson (1874, p. 110) in Moore’s collection from Dundry and
almost identical with Moore’s type specimens, M2850, Nos. 2, 5, and 6 held in the base-
ment of the City of Bath Reference Library. The Westington Hill specimens are smaller
than the examples of M. granulosa held at the British Museum. These have a brachial
valve width of 2-5-3-5 mm. whereas the largest specimens from Westington Hill have a

EXPLANATION OF PLATE 74

Figs. 1-8. Moorellina granulosa (Moore) 1 . Stereoscan photomicrograph (37508) showing the brachial
arches. Postero-lateral view, angle of incidence 48° to the plane of the valve. X 250. 2. Stereoscan

photomicrograph (37509) showing the brachial lobe free from the floor of the valve. X indicates the
point of attachment to one brachial tubercle. Antero-lateral view, angle of incidence 36° to the plane
of the valve. X 100. 3. Stereoscan photomicrograph (37507) normal to the plane of the valve

showing the cardinal process with inner and outer socket ridges. X 100. 4. Very young form

cemented to the anterior border of specimen 37509. X 250. 5. Photomicrograph, reflected light.

Vertical transverse section through specimen (37511) showing the posterior extensions of the brachial
lobes and median septum. X 46. 6. Retouched copy of fig. 5. 7, 8. Photomicrographs prepared

from acetate peels of vertical transverse sections of specimen (37511) at 0-68 and 0-7 mm. from the
umbo, showing the form of the brachial lobes and their attachment to the floor of the valve by
brachial tubercles. X 50.

Palaeontology , Vol. 12

PLATE 74

BAKER, Moore Hina granulosa

P. G. BAKER: BRACHIOPOD MOORELLINA GRANULOSA (MOORE)

397

text-fig. 3. Series of three-quarter profile drawings to show the development of the morphological
features of the interior of the brachial valve of Moorellina granulosa (Moore) during ontogeny.
a. brephic valve, b. pre-forbesiform stage, c. early forbesiform, d. late forbesiform, e. post-forbesiform

f. adult valve.

brachial valve width of only 1-7 mm. (Moore’s types 1-2-1 -8 mm.). In addition, the
posterior region of the sub-peripheral rim is different, the rim being inclined posteriorly
in the majority of the large Dundry specimens and forming quite pronounced angles
where it turns to unite with the bridge. In the Westington Hill and Moore’s type material
the rim remains vertical in this posterior region (PI. 73, figs. 3-6). It is felt, therefore,
that re-study of the larger forms must be undertaken in order to ascertain whether they

398

PALAEONTOLOGY, VOLUME 12

are properly assigned to M. granulosa. Although the stages show a general size increase,
size is found to be no criterion of stage of development. This morphological variation
is judged by Elliott (1948, p. 24) to be the natural result of intra-specific variation.

GENERAL OBSERVATIONS

As it appears almost impossible to clean brachial valves without destroying the brachial
lobes, if present, a technique for serial sectioning oriented complete shells at 0-02 mm.
intervals has been developed in order to discover the true nature of the brachial lobes
and the nature of their attachment to the brachial valve. Sections show that the brachial
lobes are extremely delicate (approximately 0-03 mm. thick) when first formed. They
are, in fact, developed from the brachial tubercles in the manner described and extend
posteriorly as crescentic horns, turned inwards, towards, but not uniting with, the
posterior termination of the median septum (PI. 74, fig. 5-8).

As, in the forms studied, the brachial lobes are only attached to the valve floor by
the slender brachial tubercles, one feels that this must surely be the explanation for
the general absence of brachial lobes in detached brachial valves of M. granulosa,
although the remains of the brachial tubercles are quite common. On separation of the
valves, such delicate structures could hardly be expected to survive in the accepted en-
vironment of the thecidellinids (Ager 1965, Nekvasilova 1967). It is probable that the
bridge also is usually broken in detached valves, a view supported by the fact that the
bridge is present in all sectioned shells with a brachial valve width of more than 0-8 mm.

Distinction between the post-forbesiform stage and the adult is not possible in
brachial valves, in which only the broken tubercles remain. As size is found to be no
criterion of stage of development, it might be better at present to include all forms with
expanded tubercles in the adult stage.

Study of the Oolite Marl material may resolve the difficulty observed by Pajaud
(19666) concerning the division of the sub-family Moorellininae Pajaud 1966 into the
genera Moorellina Elliott 1953 and Rioultina.

The criterion of distinction between these genera is said to be the absence of brachial
lobes ( Moorellina ) or the presence of well established auriform brachial lobes ( Rioultina ).
The genus Elliot tina Pajaud 1963 created on the form of the area is wisely reduced to
sub-generic rank. Pajaud maintains that Rioultina is evolved from Moorellina stock.
The Westington Hill specimens might logically be considered to occupy an intermediate
position in time ( Moorellina , Rhaetic to Bajocian, Rioultina, Pliensbachian to Oxfordian).
Careful comparison of the thecidellinid material of the British Museum and the In-
stitute of Geological Sciences with that collected from Westington Hill shows a range of
features in the Westington material which grade from moorellinid to rioultinid so that
the only real difference is that of size. Rudwick (1962, p. 334) notes the occurrence of
typically adult shells of Terebratella inconspicua (Sowerby) which are much below the
normal size and attributes this to phenotypic stunting. It is equally possible that the
material from Westington Hill represents a dwarfed population.

The ontogeny, however, shows the clear development of a ptycholophe (see Pajaud
1966a, p. 618) whereas Rioultina is said never to get beyond the schizolophe (Pajaud
19666). The problem therefore, appears to be one of definition; either one must accept
that some moorellinids do have brachial lobes or these forms must be referred to

P. G. BAKER: BRACHIOPOD MOORELLINA GRANULOSA (MOORE) 399

a new genus. One hesitates to create further new genera until more is known about the
relationship of the brachial with the pedicle valve. However, concerning the brachial
valves of moorellinids it appears that the form of the brachial apparatus has a higher
taxonomic value than the mere presence or absence of brachial lobes.

The presence of a ptycholophe in the genus poses the problem of its systematic
position at family level, as the ptycholophe is regarded as a thecideid character. Owing
to the difficulty of determining between post-forbesiform types and adults or even
whether the ptycholophe is universally an adult character in the population its taxonomic
significance can not yet be fully appreciated. However, the clear ability to develop a
ptycholophe is regarded as being important, particularly in view of the phyletic relation-
ships proposed by Rudwick (1968, p. 352) as it increases the probability that the simple
ptycholophous thecideaceans have evolved from moorellinid stock.

REFERENCES

ager, d. v. 1965. The adaption of Mesozoic brachiopods to different environments. Palaeogeography ,
Palaeoclimatol., Pcilcieoecol. 1, 143-72.

davidson, t. 1 874. Supplement to the British Jurassic and Triassic Brachiopoda, British Fossil Brachio-
poda, 4, 110, Sup. PI. 12. Palaeont. Soc. [Monogr.], London.
elliott, g. f. 1948. Palingenesis in Thecidea (Brachiopoda). Ann. Mag. nat. Hist. (12) 1, 1-30, pi. 1, 2.

1953. Classification of the Thecidean Brachiopods. Ibid. (12) 6, 693-701, pi. 18.

hallam, a. 1967. The interpretation of size-frequency distributions in molluscan death assemblages.
Palaeontology, 10, 25-42.

hendry, r. d., rowell, A. J., and J. w. Stanley. 1963. A rapid parallel grinding machine for serial
sectioning of fossils. Ibid. 6, 145-7, pi. 20.

moore, c. 1855. On new Brachiopoda from the Inferior Oolite of Dundry. Proc. Somerset Arch. Nat.
Hist. Soc. 5 (for 1854), 107-28, 3 pis.

nekvasilova, o. 1964. Thecideidae (Brachiopoda) der bohmischen Kreide. Sbor. geol. ved, Praha,
3, 119-162, 12 pis.

1967. Thecidiopsis ( Thecidiopsis ) bohemica imperfecta n. subsp. (Brachiopoda) from the Upper

Cretaceous of Bohemia. Ibid. 9, 115-36, 8 pis.

pajaud, d. 1963. Note sur les Thecideidae (Brachiopodes) jurassiques. Bull. Soc. geol. Fr. (7) 5,
995-1000, pi. 24b.

1966a. Note preliminaire a la classification des Thecidees (Brachiopodes). Ibid. (7) 8, 615-20.

19666. Problemes relatifs a la determination des especes chez les Moorellininae (Thecideidae,

Brachiopodes). Ibid. (7) 8, 630-7.

rudwick, m. j. s. 1962. Notes on the ecology of brachiopods in New Zealand. Trans, roy. Soc.
N.Z. Zoo/. 1, 327-35.

1968. The feeding mechanisms and affinities of the Triassic brachiopods Thecospira Zugmayer

and Bactrynium Emmrich. Palaeontology, 11, 329-60.
williams, A. 1965. Morphology in r. c. moore (ed).) Treatise on Invertebrate Palaeontology, Part H,
Brachiopoda. Geol. Soc. Am. and Univ. Kansas Press.

P. G. BAKER

Department of Biological Sciences
Derby and District College of Technology
Kedleston Road
Derby, de3 Igb

Typescript received 23 September 1968

A CONODONT ASSEMBLAGE FROM THE
CARBONIFEROUS OF THE AVON GORGE,

BRISTOL

by R. l. Austin and f. h. t. Rhodes

Abstract. A fused assemblage of four apatognathids and one Spathognathodus scilulus is described and illust-
rated. It was collected from relatively unfossiliferous carbonates from the D Zone of the Avon Gorge, Bristol.
Only two of the apatognathids are closely similar. The others are of different sizes and represent different species.
The assemblage is interpreted as the remains of a single conodont-bearing animal and this is supported by a
consideration of other assemblages, the rarity of conodonts in strata above and below those yielding the present
assemblage, and the similar occurrence and stratigraphic ranges of the components in this and other localities.

The recent growing interest in the value of conodonts in problems of stratigraphic
correlation is based largely upon the study of discrete specimens which have been isola-
ted from a carbonate matrix by digestion in dilute acid. In spite of the value of these
individual conodont ‘species’ there has long been a realization that different individual
forms of conodont may have been combined together within the body of a single animal.
This possibility was first suggested by Hinde(1879, p. 361) who described what he thought
to be a natural association of conodonts from the Devonian Genesee Shale of New
York. He argued that the intimate association of a large number of different types of
conodont on the bedding plane of the shale implied their original association.

Although this particular group is probably not a true natural assemblage (see Rhodes
1962, p. Wl\ for details), other workers have described associations of conodonts,
which are accepted by most workers as original ‘natural’ (i.e. biological) assemblages.
Most of these are from black fissile shales of the Carboniferous of North America and
Europe (Schmidt 1934, 1950; Scott 1934, 1942; Dubois 1943; Schmidt and Muller 1964;
Rhodes 1952, 1954, 1962). A full discussion of the arguments for regarding these as
natural assemblages is given by Rhodes (1962, p. WIT).

Other less well-documented Carboniferous assemblages are described by Cooper
(1945) and D. J. Jones (1956, p. 126).

Two other methods of recognising original associations of conodonts have been
developed within recent years. Rexroad and Nicoll (1964), Klapper (in Rexroad and
Nicoll 1964), and Barnes (1967) have described fused conodont assemblages from Or-
dovician and Silurian strata, in which individual conodonts are fused to others of similar
form. In some cases, e.g. Rexroad and Nicoll (1964), the fused elements are all of the
same size and belong to the same form species; in others, e.g. Barnes (1967), they are of
the same form genus, but represent individuals of different size and different form
species. Barnes, Rexroad, and Nicoll have discussed at length the basis for regarding
these fused specimens as natural associations.

Other workers have recently attempted to recognize original associations of conodonts
by analysing the statistical distribution of individual species in large samples of isolated
conodonts, e.g. Walliser (1964), Bergstrom and Sweet (1966), and Webers (1966).

[Palaeontology, Vol. 12, Part 3, 1969, pp. 400-405.]

AUSTIN AND RHODES: CONODONT ASSEMBLAGE FROM BRISTOL 401

These studies have involved strata of Ordovician and Silurian age in which, if the infer-
ences as to natural assemblages are correct, the arrangement of individual conodonts
is rather more simple than that found in Carboniferous assemblages.

During a recent study of lower Carboniferous conodonts (Rhodes, Austin, and Druce
1969) a group of fused conodonts has been recovered from a typical light grey, fine
grained calcarenite of the D Zone of the Avonian at the Avon Gorge, Bristol (ST
564734, sample D7, in the above study). 6 kg. of the rock was processed in acetic
acid and a heavy mineral separation in bromoform of the dried residue produced a
total of nine conodonts referable to the genera Apatognathus, Hindeodella , and Spathog-
nathodus. Five were originally fused together as an assemblage, but one specimen be-
came detached in preparing the specimen. The assemblage is deposited in the collection
of the Department of Geology, University of Southampton (Catalogue number 10412).

Description of the assemblage

The assemblage consists of three specimens of the genus Apatognathus Branson and
Mehl and a single specimen of Spathognathodus scitu/us (Hinde). The arrangement of
the specimens is illustrated in text-fig. 1. This may or may not be the original biological
arrangement. The biggest of the three apatognathids (unit 2) is orientated so that its
anterior and posterior limbs both lie in a horizontal plane. The posterior bar lies nearest
to the observer when seen in text-fig. 1. Its apical denticle region is fused to the anterior
distal end of unit 1, the most anterior of the three Apatognathus specimens. This specimen
is oriented at right angles to the plane of the posterior bar of the biggest specimen, and
it faces in a position directly opposed to it. The third specimen of Apatognathus (unit 3)
lies near the distal end of the posterior bar of unit 2, and is also arranged at right angles
to it, both in a horizontal and vertical plane. This specimen, however, points directly
posteriorly so that its denticles point in the direction of the distal end of the posterior
bar of unit 2. The small specimen of Spathognathodus parallels the apical denticle of
the Apatognathus (unit 2.).

The Apatognathus elements show individual differences. The anterior and posterior
apatognathid elements of the assemblage tend to resemble one another in overall size
and form. It is difficult to identify these in terms of existing species of Apatognathus
but they resemble Apatognathus chauliodus Varker and A. cuspidatus Varker. The biggest
Apatognathus unit possibly represents a distinct species.

The fusion of the four specimens with one another is very strong, and seems to be a
result of additional material having the same appearance as that from which the cono-
donts are made. There seems to be ‘fusing material’ in direct and continuous contact
with the opposed surfaces on the conodonts, but although we have looked at the assem-
blage under the highest magnification which we can obtain by use of optical microscopy,
we have been unable to see any detail of this material.

One remarkable feature of the present specimens is the fact that they are preserved
in such striking three-dimensional relief. They show little, if any, effect of compression.
The two most striking features of the assemblage are the opposed position of the three
elements at right angles to one another, in both a horizontal and a vertical plane, and
also that in each case the distal end of one of the bars is fused to the apical end of its
neighbour. It seems improbable that this is wholly fortuitous, but it is quite unlike the
parallel and common alignment which is characteristic of all other known assemblages.

402

PALAEONTOLOGY, VOLUME 12

denticles on oral
surface of
posterior bar

posterior bar

OUTER LATERAL VIEW
OF APICAL CUSP

POSTERIOR

BAR

UNIT 3

OUTER
LATERAL
SI DE

denticle on oral
surface of anterior
bar

anterior

bar

OUTER LATERAL SIOE

POSTERIOR BAR

apical

cusp

APICAL CUSP

text-fig. 1. Stereoscan photograph and drawing of assemblage of Apatognathus and Spathognathodus
scitulus. For details see text. Unit 1 is labelled in lower-case letters, unit 2 in upper-case italics, and unit
3 in upper-case roman. Magnification x 70 approx.

Systematic palaeontology

Specific identification of the hindeodellid fragment is impossible. One of the isolate
apatognathids is compared with A.petilus Varker, another is fragmentary and specifically
indeterminate, and the remaining two, which are similar, are well preserved, but juveniles.
They do not agree exactly with any descriptions or illustration of existing species,

AUSTIN AND RHODES: CONODONT ASSEMBLAGE FROM BRISTOL 403

though they show some resemblance to A. cuspidatus Varker. They differ most con-
spicuously from illustrations and descriptions of this species, however, in the relative
length of the apical denticle, and in the degree of lateral twisting of the anterior and
posterior bars. This is so extreme that the denticles curve upwards towards each other
and the centre of the arch, away from the plane of the anterior and posterior limbs.
The apical angle is extremely acute and in this it resembles A. cuspidatus as well as
A. chauliodus Varker.

One of the specimens (unit 1) fused in the assemblage, appears to represent the same
species as the two free specimens just described, but it is a somewhat larger individual
and it has a longer apical denticle than either of the other specimens.

Interpretation of the assemblage

We regard our specimen as a natural assemblage not only because of the enormous
improbability of any artificial association of this kind within samples which have been
subjected to such relatively violent disaggregation, but also because of other occurrences
of a similar kind of fusion in what are to us clearly natural conodont assemblages
(Rexroad and Nicoll 1964, Barnes 1967). The association must, in our view, clearly be
‘original’ in the sense that it represents a pre-depositional association. The rarity of
conodonts in this sample seems to make it wholly improbable that it could be regarded
as a fortuitous sedimentary inorganic association, which occurred after the death of the
conodont bearing ‘animal’.

It may then be asked what type of association this could represent if it is accepted as
an original biological association. It may represent some kind of pathologic condition,
in which elements normally freely associated together within a single animal, have
become abnormally fused together. This also seems to be the most acceptable interpre-
tation of the specimens described by Rexroad and Nicoll and by Barnes. Rexroad and
Nicoll interpreted a similar association as a possible case of tetanus, but this may be to
read more into the association of the conodonts than is justified. It seems unlikely how-
ever, in view of the very great rarity of other fused specimens, that this could have been
a ‘natural’ condition.

A striking feature of the present assemblage is the difference in size between the
elements that are associated together. It might be argued that the association of elements
of such very different sizes is against their original association in a natural assemblage,
but a fused assemblage from the Ordovician Coburg Formation of Ottawa ( Barnes 1 967),
contains an association of belodids, which show a comparable variation in individual
size. Barnes argues that this association may suggest either a process of continuous
replacement within a natural conodont assemblage or the original association of strikingly
similar conodonts of different sizes. Either interpretation could be applied to the present
group of apatognathids.

In contrast to this most of the elements in known Carboniferous conodont assemblages
are of almost identical size and seem, therefore, to have been directly paired. It could
be that Apatognathus, which is a genus with a very irregular occurrence, did not represent
a similar type of paired component in a natural assemblage and Cooper (1945) has
argued that at least some Carboniferous assemblages may have contained unpaired
components. Other workers (Schopf 1966, Webers 1966, Bergstrom and Sweet 1966)
have suggested that Ordovician assemblages were probably composed of different types

404

PALAEONTOLOGY, VOLUME 12

of individual components from those of Pennsylvanian age. The present assemblage can-
not therefore, be interpreted only by comparison with known Pennsylvanian assemblages
(Rhodes 1952).

It is difficult to compare the present conodont fauna of the assemblage with any
immediately adjacent to it, because from samples collected at 10 ft. intervals from 200 ft.
of strata immediately below and from 50 ft. immediately above the present sample no
conodonts have been recovered. It is noteworthy however that the genus Apatognathus,
which is not known to form more than 25% of faunas of broadly similar age in other
areas, constitutes 77% of the fauna of the present sample. Both the rarity of conodonts
in over 250 ft. of strata (138 kg. of which were digested in acid) and the relative rarity
of Apatognathus elsewhere support the possibility that the assemblage may be an original
biological association. The strong nature of the fusion of the elements, also seems to us
to favour such an interpretation.

As noted above, studies of some Silurian (e.g. Walliser 1964) and Ordovician
(Bergstrom and Sweet 1966) faunas have shown a constant numerical relationship and
identical stratigraphic range between certain isolated conodont ‘species’. These have
been interpreted as assemblages.

We have compared the relative abundance of individual components of our assem-
blage, when they occur as isolated conodont elements in strata of comparable age in
the North Crop of the South Wales coalfield, the Avon Gorge, Bristol, Yorkshire, and
Scotland. Comparison of the percentage frequency of all isolated apatognathids with
S. scitulus provides no consistent ratio between them, individual samples ranging from
1 : 2-50 to 1 : 0-20. Similar variable ratios are also found in other individual samples of
very large conodont faunas from which assemblages have been recognized (e.g. Schopf
1966, Webers 1966). Most authors attribute this to post mortem sorting (e.g. Schopf
1966, p. 16).

The stratigraphic ranges of S. scitulus and Apatognathus are similar in each area from
which they have been recovered. Also S. scitulus and Apatognathus , though both rela-
tively rare, are most frequently found in the same sample. This common association
is also present in the Visean rocks of North Wales (Aldridge, Austin, and Husri 1968).

We interpret the common association and similarity of stratigraphic range of S. scitulus
and Apatognathus, when found as isolated elements, as support for the suggestion that
they were originally associated together as a biological assemblage.

CONCLUSIONS

A fused assemblage of four Apatognathus with a S. scitulus suggests that these
elements were associated together in the same conodont-bearing animal. This inter-
pretation is supported by a general consideration of the occurrence and preservation
of the assemblage, comparison with other known assemblages, other common occur-
rences of the two components, and their generally similar stratigraphic ranges.

Variation in the size of the Apatognathus components of the assemblage suggests that
there may have been a form of continuous replacement of components within the assem-
blage or, more probably, that different sizes of the same element were present.

The relative positions of individual apatognathids in the assemblage may or may not
represent the original orientation within the conodont-bearing animal. It seems to us

AUSTIN AND RHODES: CONODONT ASSEMBLAGE FROM BRISTOL

405

improbable that they do, for all described conodont assemblages from the Ordovician,
Silurian, and Carboniferous display a broadly parallel alignment of associated elements.

The present assemblage throws little new light on the puzzling question of the affinity
and function of conodonts.

Acknowledgements. We are happy to acknowledge our gratitude for the use of the stereoscan micro-
scope at the Royal Aircraft Establishment, Farnborough, and the assistance of Miss V. M. Hale and
Mr. D. Clark. We are grateful to Miss Sonia J. Kostromin and Mrs. R. J. Aldridge for typing the
manuscript and to Mrs. A. Dunkley, Mrs. Beryl Fisher, and Mr. S. Osborn for their contributions to
the text-figure.

REFERENCES

aldridge, r. j., Austin, r. l., and husri, s. 1968. Visean conodonts from North Wales and Ireland.
Nature, Lend. 219, 255-8.

barnes, c. r. 1967. A questionable natural conodont assemblage from Middle Ordovician Limestone,
Ottawa, Canada. J. Paleont. 41, 1557-60.

Bergstrom, s. m. and sweet, w. c. 1966. Conodonts from the Lexington Limestone (Middle Ordo-
vician) of Kentucky and its lateral equivalents in Ohio and Indiana. Bull. Amer. Paleont. 40 (229),
271-441.

cooper, c. l. 1945. Microfauna of Pennsylvanian-Mississippian borderline formations (abstr.).
Bull. geol. Soc. Amer. 56, 1153.

dubois, e. p. 1943. Evidence on the nature of conodonts. J. Paleont. 17, 155-9.

hinde, g. j. 1 879. On conodonts from the Chazy and Cincinnati group of the Cambro-Silurian and from
the Hamilton and Genesee shale division of the Devonian in Canada and the United States. Q. Jl
geol. Soc. Lond. 35, 351-69.

jones, d. J. 1956. Introduction to Microfossils. Pp. 1-381. Harper and Bros., New York.

rexroad, c. b. and nicoll, s. 1964. A Silurian conodont with tetanus? /. Paleont. 38, 771-3.

Rhodes, f. h. t. 1952. A classification of Pennsylvanian conodont assemblages. Ibid. 26, 886-901.

1954. The zoological affinities of the conodonts. Biol. Rev. 29, 419-52.

1962. Recognition, interpretation, and taxonomic position of conodont assemblages, in r. c.

moore (ed.), Treatise on Invertebrate Paleontology, Part W, Miscellanea, pp. 70-83. Geol. Soc. Am.
and Univ. Kansas Press.

Austin, r. l. and druce, e. c. 1969. British Avonian (Carboniferous) conodont faunas, and their

value in local and intercontinental correlation. Bull. Br. Mus. Nat. Hist. (Geol.) Supplement No. 5,
pp. 1-313.

schmidt, h. 1934. Conodonten-Funde in urspriinglichem Zusammenhang. Paldont. Z. 16, 76-85.

1950. Nachtrage zur Deutung der Conodonten. Decheniana, 104, 11-19.

and muller, K. J. 1964. Weitere Funde von Conodonten-Gruppen aus dem oberen Karbon des

Sauerlandes. Paldont. Z. 38, 105-35.

schopf, t. j. m. 1966. Conodonts of the Trenton Group (Ordovician) in New York, Southern Ontario
and Quebec. Bull. N.Y. St. Mus. 405, 105 pp.

scott, h. w. 1934. The zoological relationships of the conodonts. J. Paleont. 8, 448-55.

1942. Conodont assemblages from the Heath Formation, Montana. Ibid. 16, 293-301.

walliser, o. w. 1964. Conodonten des Silurs. Hess. Landesamt fiir Bodenf. Abb. 41, 1-106.

webers, g. F. 1966. The Middle and Upper Ordovician conodont faunas of Minnesota. Minn. Geol.
Surv. Spec. Publ. SP-4, 1-123.

R. L. AUSTIN

Geology Department
University of Southampton
F. H. T. RHODES
Geology Department
University of Michigan
Ann Arbor

Revised typescript received 23 October 1968

THE TREMADOC TRILOBITE
PSEUDOKAINELLA IMPAR (SALTER)

by PETER H. WHITWORTH

Abstract. Salter’s species Oleiuis impar, 1866, from theTremadoc of North Wales is referred to Pseudokainella
and redescribed with the aid of new and better material. The record is the first for this genus in Europe. The
species is compared with other records of the genus, and correlations with North and South America are dis-
cussed.

During current research into the trilobite faunas of the Tremadoc rocks of North
Wales and Shropshire a number of specimens collected by the writer, and three others
from the Geological Survey collections, have provided evidence of a new record for
Europe.

A nearly complete, distorted specimen from Portmadoc (Geol. Soc. Coll. 6970) was
described and figured by Salter (1866) under the name of Olenus impar, and it is interest-
ing to note that he observed (1866, p. 303) a close resemblance of this species to
‘ Remopleurides' . His illustration with respect to the specimen Geol. Soc. Coll. 6970 is
reversed and evidently in part restored. Since then no further mention of the species
has been made, even in the monographs of Lake (1906-46) and Henningsmoen (1957).
The discovery of new and better material at Arenig substantiates my first suspicions
arising from the study of the Geological Survey material that this is a member of the
Remopleurididae, namely Pseudokainella Harrington, 1938.

Figured material is deposited at the Geological Survey Museum (GSM, Geol. Soc.
Coll.) or at Birmingham University together with topotype material (BU).

SYSTEMATIC DESCRIPTION
Family remopleurididae Hawle & Corda 1947

Remarks. The classification here adopted is that of the Treatise on Invertebrate Paleonto-
logy Part O (Moore 1959) except that the diagnosis of the Remopleurididae Hawle &
Corda 1847 should be adapted to include the narrower thoracic doublure of Pseudo-
kainella, where it extends only across the free pleural terminations. The term ‘intergenal
angle’ used in this paper refers to the prominent angulation of the posterior cephalic
margin, resulting in part from the rather forward position of the genal spines.

Subfamily richardsonellinae Raymond 1924
Genus Pseudokainella Harrington 1938
Type species. Pseudokainella keideli Harrington 1938.

Remarks. A recent diagnosis of the genus has been given by Harrington and Leanza
(1957). Kobayashi (1953) excludes this genus from the Richardsonellinae on the absence

[Palaeontology, Vol. 12, Part 3, 1969, pp. 406-413, pi. 75].

P. H. WHITWORTH: TRILOBITE PSEUDOKAINELLA IMPAR (SALTER) 407

of an intergenal angle and includes it in his (new) subfamily Kainellinae. Subsequent
work has however shown that this character can be present and even strongly developed,
as in P. keideli and P. impar. Consequently his diagnosis of the Kainellinae, based
partly on the presence of a nearly straight posterior cephalic margin (i.e. absence of
intergenal angle), requires amendment. His subgenus ParakaineUa (1953, p. 43) is
furthermore regarded as a subjective junior synonym for the reasons given by Harring-
ton and Leanza (1957, p. 133).

PseudokaineUa impar (Salter 1866)

Plate 75, figs. 1-8

1866 Olemis impar Salter, pp. 302-3, pi. 8, fig. 4.

1881 Olemis impar Salter, p. 496, pi. 8, fig. 4.

Holotype (by monotypy). Geol. Soc. Coll. 6970.

Other Material. GSM 70999, 71001; BU 398a and b, 399, 400.

Due to poor preservation of the holotype the following description is largely sup-
plemented by BU 398. The material comprises one large complete specimen, one smaller
disarticulated and nearly complete specimen, a number of small cranidia, and numerous
isolated free cheeks and pygidia from the Arenig area; one complete specimen and two
incomplete specimens (all distorted and one bearing a hypostome) from Portmadoc.

Diagnosis. A species of PseudokaineUa with nearly parallel-sided glabella, two pairs of
lateral glabellar furrows, prominent intergenal angles, and very long genal spines reach-
ing beyond the posterior end of the pygidium; width of frontal area (tr.) about twice
the posterior width (tr.) of the glabella; thorax with wide axis, and narrow doublure
widening backwards; pygidium with four axial rings, margin bearing four pairs of
evenly sized spines.

Description. Dorsal exoskeleton oval in outline. The cephalon is sub-semielliptical, slightly
more than twice as wide as long, gently convex transversally and less so sagittally. The
glabella is raised above the level of the librigenae, is gently convex, longer than wide,
tapers very slightly forwards, and is very gently curved in front with rounded antero-lateral
corners; it is expanded slightly opposite distal ends of posterior glabellar furrows and
at occipital ring; well-defined by axial furrows which are accentuated by distortion in
GSM 70999, 71001. Two pairs of lateral glabellar furrows are discontinuous across the
glabella and are more or less isolated from the axial furrows by slight swellings (dis-
tortion may give the impression that they are confluent across the glabella). Anterior
(2p) furrows short, faint, directed backwards and inwards in a slightly forward convex
curve. Posterior (Ip) furrows longer, very slightly sinuate especially in young cranidia,
directed backwards at a greater angle and stronger than 2p, being deepest at their mid-
length, and commence opposite mid-points of palpebral lobes. Occipital furrow wide
(sag. and exsag.), relatively shallow, bending forwards at the middle and more strongly
forward-outward abaxially, descending to meet the axial furrows just behind the posterior
ends of the palpebral lobes; occipital ring prominent, wider than the rest of the glabella,
turning forwards abaxially and well-defined laterally by an outward curve of the axial
furrows.

408

PALAEONTOLOGY, VOLUME 12

Fixigenae much reduced; palpebral areas very narrow (tr.), elongate-crescentic;
posterior areas extremely narrow, blade-like, extending out to a distance about equal
to two-thirds the width of the occipital ring; anterior areas expanded, triangular, some-
what inflated. Preglabellar area rather narrow, nearly flat, set below the level of the
glabella and lateral extensions of anterior fixigenal areas; it is bounded anteriorly by
the wider (sag. and exsag.), gently convex, forward-curved anterior border which nar-
rows abaxially, and by the anterior border furrow which carries a row of small pits.
Palpebral lobes moderately large, narrow, crescentic, raised well above level of cheeks
abaxially; situated slightly behind the mid-line (tr.) of the glabella and closer to it in
front than behind; they extend from opposite outer ends of 2p furrows to just in front
of occipital furrow. Palpebral furrows prominent but fairly shallow; visual surfaces of
eyes not preserved. Librigenae are gently convex with moderately wide lateral borders
which extend into strong spines without angular deviation from the general curvature
of the cephalic border; genal spines reach back beyond the posterior margin of the
pygidium in a gentle outwards curve; the lateral border furrows are wide and shallow.
The posterior cephalic border, at first directed slightly backwards, widens rapidly and
then turns sharply forwards through about 80° to form a prominent intergenal angle,
finally becoming narrower towards the genal angles. The intergenal angles encroach
strongly across the first and part of the second thoracic segments. The posterior border
furrow is flat and wide, commencing opposite the middle of the occipital ring; it more
or less follows the curve of the posterior border but bends through only 45°; its con-
fluence with the lateral border furrow is usually obtuse (but see below). Inner spine
angle (see Henningsmoen 1957, p. 13) acute, about 45°.

Sutural pattern kainelliform. Anterior sections of facial suture are short and strongly
divergent forwards with an angle of 145° between them; after turning abruptly around
the anterior border they become marginal and meet axially, continuing across the
doublure as a median suture. The anterior sections reach the axial furrow immediately
in front of the palpebral lobes. The posterior sections, which do not quite reach the
axial furrow, turn sharply backwards and outwards behind the palpebral lobes and
curve gently back to cut the posterior border of the cephalon just inside the intergenal
angle.

The hypostome, previously unrecorded, shows a typical Apatokephalid form, though
it is rather elongated by distortion. The general outline is ovate (probably originally
more or less oval) with length greater than breadth; the frontal margin is well rounded
and probably has a narrow, flat anterior border. A slight posterior increase in breadth

EXPLANATION OF PLATE 75

Figs. 1-8. Pseudokainella impar (Salter). 1, Flolotype, Geol. Soc. Coll. 6970, x 2£; Tremadoc (? Port-
madoc Flags), Pen-y-clogwyn, Portmadoc, Caernarvonshire. G.R. 56553830. Figured and partly
restored by Salter. 2, Flypostome, excavated and enlarged from fig. 3, GSM 71001, x4; Tremadoc
Slates, Portmadoc. 3, Cranidium, hypostome, and part of thorax, GSM 71001, x2J; Locality as
for fig. 2. 4, Attached librigena showing genal caeca and reticulate ornament, BU 399, X 2\ \ Upper
Tremadoc, Shumardia Beds, Ceunant-y-garreg-ddu, Arenig. G.R. 821 53603. 5, Small uncrushed
cranidium showing anterior border pits, BU 400, X4; Upper Tremadoc, Shumardia Beds, stream
section near Arnnodd Bwll, Arenig. G.R. 80753690. 6, Damaged cranidium and part of thorax
(note effect of deformation on glabellar furrows), GSM 70999, x21; locality as for fig. 2. 7, 8,
Internal mould (BU 398a) and latex cast of counterpart (BU 398b) respectively, both X 1£; Upper
Tremadoc, Shumardia Beds, locality as for fig. 5.

Palaeontology , Vol. 12

PLATE 75

WHITWORTH, Tremadoc trilobite Pseudokainella

P. H. WHITWORTH: TRILOBITE P SEU DO KAINELLA IMPAR (SALTER) 409

coincides with a widening of the lateral border. Anterior wings if present are not pre-
served. The middle body is strongly convex, reaching a maximum in front of centre;
it is oval and bounded laterally by a deep border furrow which continues around the
posterior margin. Maculae appear to be absent, and a posterior body is not defined
(except by a lowering of convexity which may be due to crushing). The lateral border
furrow becomes deep postero-laterally where the border is slightly expanded and upward-
turned. The posterior furrow is much narrower and shallower, defining a rounded
margin to the posterior lobe and a gently convex posterior border.

The cephalic doublure and dorsal surface of the cephalic border are ornamented
with some 9 or 10 well-spaced terrace lines, and the glabella, librigenae, and anterior
areas of the fixigenae have a close-set reticulate ornament.

The specimen from Ceunant-y-garreg-ddu (BU 399) shows two features not seen on
the other material. The first is the angle of intersection of the lateral border- and posterior
border-furrows which, normally rather obtuse, is here acute and directed strongly
backwards and outwards in a point. The second is the presence of a strong caecal ridge
on the librigena; this commences opposite the mid-point of the palpebral lobe and
crosses the librigena obliquely backwards and outwards at an angle of 40° with the
axial normal; at a position level with the posterior end of the palpebral lobe it turns
abruptly outwards and runs directly towards the genal angle, dying out before reaching
the same. These features could prove to be of varietal significance, but further material
is required before more definite conclusions can be reached.

Thorax of 12 segments. Axis wider than the pleurae for the first 8 segments, then
becomes increasingly narrower than the pleurae; it is moderately convex and raised
well above the pleural regions. The first axial segment is slightly wider (tr.) than the
occipital ring, and the second slightly wider again, thereafter each becoming successively
smaller. Axial rings are straight medially but bend forwards abaxially and form vaguely
defined marginal lobes; ring furrows are wide axially where they bend forwards, but
laterally they become narrower and deeper as they curve forwards to meet the axial
furrows. Articulating half-rings are well developed. Axial furrows are strong and slightly
outwards curved around the abaxial ends of each axial ring. 8th segment bears a broadly
based median spine which extends well beyond the posterior margin of the pygidium;
it appears to have a double base — the main ridge, developing from the anterior part
of the axial ring, is supported by a broader triangular base which originates from the
posterior part of the ring and extends back as far as the 10th segment before the two
parts merge into a single spine. Pleural regions are nearly flat and of constant width
(tr.); weak proximal fulcra can be made out in each of the first few anterior segments
only. Each pleura is crossed by a prominent oblique pleural furrow, commencing in
confluence with the axial furrow in a shallow axial socket at the anterior margin of the
pleura, and reaching the inner edge of the doublure in the posterior half of the segment.

The original convexity of the exoskeleton seems to have been rather low and the shell
material thin. As a result a thin layer of deposit lying between the dorsal exoskeleton
and the doublure remains attached to the external mould, and the pleural furrows have
been impressed through it on to the doublure. This layer is present on both thorax and
pygidium and is too thin and firmly attached to allow removal without damaging the
underlying surface. This is why the impression of the doublure is seen on the external
mould.

410

PALAEONTOLOGY, VOLUME 12

The pleural furrows can therefore be seen to continue into the pleural terminations
in each segment. Each furrow approaches the posterior pleural border and dies out
before reaching the acute, backward-pointed spinose termination. These pleural termina-
tions become progressively longer and more spinose posteriorly, an appearance which
is somewhat exaggerated by a corresponding increase in the width of the doublure with
its ornament of step-like terrace lines. Each axial ring bears a reticulate ornament of
Bertillon pattern as on the glabella; the wider posterior band of each pleura also has a
weak reticulate ornament.

text-fig. 1. Comparative diagrams of two species of Pseudokainella. (a) P. impar (Salter) from BU
398. ( b ) P. lata (Kobayashi) after Harrington and Leanza 1957. Both x 1-4.

Pygidium sub-elliptical and twice as wide as long. Axis is convex and raised above the
level of the pleural regions; it tapers gradually backwards and bears four segments plus
a rounded sub-triangular terminal piece, the latter being prolonged at a lower level into
a weak, flattened post-axial ridge which dies out shortly behind the anterior edge of the
pygidial doublure; the abaxial terminations of the first and second rings are feebly
lobate. Axial and ring furrows are straight and become weaker posteriorly. Pleural re-
gions, unlike those of the thorax, are somewhat inflated and the pleural terminations are
turned slightly upwards. Three well marked pairs of pleurae and a fourth less prominent
pair close to the terminal axial segment are seen. Interpleural furrows are weaker than the
oblique pleural furrows, but both curve backwards and are directed backwards at increas-
ing angles until the fourth pair lie nearly parallel to the axis. The first two pleural furrows

P. H. WHITWORTH: TRILOBITE PSEUDOKA1NELLA IMPAR (SALTER) 411

extend into the marginal spines, the third is considerably shorter, and the fourth does
not extend beyond the inner limit of the doublure. The pygidial margin is extended into
four pairs of lateral spines which correspond with the continuations of the pleurae;
they are short and triangular with wide bases, and each pair, except for the innermost
which are small and tooth-like, becomes only gradually smaller adaxially. Doublure
wide, reaching over half-way in towards the axis, and ornamented with terrace lines
which converge and become closely crowded behind the axis.

Measurements (in mm.) of BU 398
Width of cephalon (at occipital ring) 32-0

Length of cephalon c. 17 0

Width of cranidium (at eyes) 12-8

Width of glabella (at eyes) 9 0

Length of glabella (exc. occipital ring) 10-3

Width of occipital ring 9 3

Length of occipital ring (sag.) 2-8

Length of palpebral lobe 5 0

Length of thorax c. 25-0

Width of thorax (anterior) 26 0

Width of thoracic axis (anterior) 10 3

Width of pygidium (anterior) 19-3

Length of pygidium (excluding spines) 8-7

Width of pygidial axis (anterior) 5-5

Length of pygidial axis 6 5

The holotype is considerably less than half the overall size of BU 398 and is too
distorted to obtain reliable measurements from.

Localities. Tremadoc Slates, Portmadoc, and Pen-y-clogwyn (no other information known )\Shumardia
Beds, north bank of stream 150-60 paces upstream from east end of wall by forestry road and 200
yards NW. of Amnodd Bwll farm, G.R. 80753690; Shumardia Beds, south bank of Ceunant-y-garreg-
ddu gorge about 30 yards upstream from stone wall at bottom and 1 mile SE. of Amnodd Bwll,
G.R.821 53603.

Remarks. Pseudokainella impar seems to be closest to the Argentinian species P. lata
(Kobayashi 1935) in general outline and in the characters of thorax and pygidium. It
differs however in having a less constricted anterior lobe to the glabella, more strongly
divergent anterior sections to the facial suture, smaller palpebral lobes which do not
extend so far backwards, a weaker occipital furrow, wider posterior border furrows,
and a strong intergenal angle. The thoracic axis is also wider and the reticulate ornament
generally stronger. P. keideli Harrington 1938 is much smaller and shorter over-all.
Its cephalon has larger palpebral lobes, narrower anterior glabellar lobe, shorter
posterior sections to the facial suture, and more divergent genal spines. Its pygidium is
shorter and has a macropleural first segment. P. pustidosa Harrington & Leanza 1957 is
intermediate between keideli and lata and has a strong pustulose ornament.

Of the other recorded occurrences referred to this genus, P. armatus Hintze 1953
was believed by Ross (1957) to belong to an unnamed and ‘undescribed, more primitive
genus’, and was later redescribed by Lochman (1964) as Praepatokephahis armatus.
Sando’s (1958) two specimens from the Stonehenge Limestone of Pennsylvania, which
he tentatively compares with P. armatus, are, therefore, not to be included in the genus;
indeed, the glabellar shape and furrows are much more reminiscent of Apatokephalus

412

PALAEONTOLOGY, VOLUME 12

s.l. than of Pseudokainella. Kobayashi (1960) illustrates and describes a ?new species
(sic), but I prefer not to include this at present in Pseudokainella on the grounds of
insufficient and fragmentary material, especially as he himself expresses doubt as to its
correct assignment. P? macarenae (Harrington and Kay 1951) has a rather long pre-
glabellar field bearing radiating ridges suggestive of Kainella, but otherwise it closely
resembles P. lata; it has recently been placed in the new subfamily Artokephalinae
by Chugaeva (1964).

Until now Pseudokainella has been recorded only from Argentina, Korea, Columbia,
and recently North America (R. J. Ross Jr., personal communication), and the present
record lends credence to the idea that it may well be, like its close relative Kainella,
considerably more widespread than at present believed. Its presence strengthens the
relationship between the Kainella jCeratopyge faunas of South America and the Pharo-
stomina fauna of Balto-Scandia (Whittington 1966). The genus occurs largely in the
Lower Tremadoc of Argentina but only in the Upper Tremadoc of North Wales. This
seems to indicate a temporary influx into the Welsh area late in Tremadocian times after
a northward migration from South America. This migration could well have been along
the North American-Balto-Scandian-Russian zone, since it is now known to occur in
North America (see below). Such a migration route may perhaps be indicative of an
early development of the dispersal directions indicated by Whittington (1966, text-fig. 2)
for the Arenig-Llandeilo period.

R. J. Ross Jr. (personal communication) informs me that a species of Pseudokainella
is now known to occur in Clear Creek Canyon, Monitor Range, Nevada (Lowell 1965)
in association with Hypermecaspis and Parabolinella, an assemblage very reminiscent of
that in both Argentina and Britain (the British species Parabolinella rugosa Crosfield
& Skeat is thought to be a Hypermecaspis, see Harrington and Leanza 1957 and
Henningsmoen 1957). Precise correlation of the American assemblage is not yet settled,
although it ‘seems to correlate with the lowest Hystricurus-Symphysurina assemblage
further east’, i.e. Utah (Ross, personal communication). Since the underlying Saukia
Zone of North America, usually placed in the late Upper Cambrian, now seems for
certain to correlate with the lower part of the Mexican Tinu Formation (Lower Tre-
madocian) of Robison and Pantoja- Alor 1968, the Symphysurina Zone might be con-
sidered as Upper Tremadocian in age. As such it would correlate well with the British
Upper Tremadocian, and if Ross’s correlations are verified the mutual presence of
Pseudokainella would support this. That the Mexican and British Tremadoc are closely
equivalent can be readily seen from the presence in both areas of species of Geragnostus,
Asaphellus, Angelina, Bienvi/lia, Parabolinella, Leptoplastides, Peltocare, and Shumardia.
In addition to the presence of Hypermecaspis in Nevada, affinities between North
America and Britain are further strengthened by the mutual association of species of
Shumardia, Beltella , and Apatokephalus. These last all occur in the Goodwin Limestone,
Pogonip Formation of Nevada (Merriam 1963, Nolan and others, 1956) and some
occur in the middle part of the Garden City Formation, Utah (Ross 1951). Both may
therefore be considered as equivalent to some part of the Tremadocian (the Garden
City Formation in part only).

Acknowledgements. I wish to thank Dr. Adrian Rushton (Institute of Geological Sciences, London)
for the loan of material and for helpful discussion; Dr. Isles Strachan (Birmingham University) for
his continual help and encouragement during the preparation of this paper.

P. H. WHITWORTH: TRILOBITE PSEUDOKAINELLA IMPAR (SALTER) 413

REFERENCES

chugaeva, m. n. 1964. Trilobites of the Early and Middle Ordovician of the northeast of the U.S.S.R.,
and Analysis of trilobites in chugaeva, m. n., rozman, kh. s., and ivanova, v. a., Comparative
bio-stratigraphy of the Ordovician deposits of the northeast of the U.S.S.R. (In Russian.) Trudy geol.
Inst., Leningr. 106, 24-85, pis. 1-6 (not seen).

Harrington, h. J. 1938. Sobre las faunas del Ordoviciano inferior del norte Argentino. Revta Mus. La
Plata , N.S. 1 ( Paleontology ) 4, 209-89.

and kay, m. 1951. Cambrian and Ordovician faunas of Eastern Columbia. J. Paleont. 25, 655-68,

pis. 96, 97.

and leanza, a. f. 1957. Ordovician trilobites of Argentina. Univ. Kansas Spec. Publ. 1, 1-276,

137 figs.

henningsmoen, g. 1957. The trilobite family Olenidae. Skr. norske Vidensk.-Akad., Mat.-naturv.
Kl. 1, 1-303, pis. 1-31.

hintze, l. f. 1953. Lower Ordovician trilobites from western Utah and eastern Nevada. Bull. Utah
geol. miner. Surv. 48, 1-249, pis. 1-28.

kobayashi, t. 1953. On the Kainellidae. Jap. J. Geol. Geogr. 23 (3), 37-61, pis. 3, 4.

1960. The Cambro-Ordovician formations and faunas of South Korea; Part 6, palaeontology 5.

J. Fac. Sci. Tokyo Univ., sect. 2, 12 (2), 217-75, pis. 12-14.
lake, p. 1906-46. A Monograph of the British Cambrian trilobites. Palaeontogr. Soc. ( Monogr .),
parts 1-14, 1-350, pis. 1-47.

lochman, c. 1964. Basal Ordovician faunas from the Williston Basin, Montana. J. Paleont. 38, 453-75,
pis. 63-7.

lowell, j. d. 1965. Lower and Middle Ordovician stratigraphy in the Hot Creek and Monitor Ranges,
central Nevada. Geol. Soc. Am. Bull. 76, 259-66.

merriam, c. w. 1963. Palaeozoic rocks of Antelope Valley, Eureka and Nye Counties, Nevada. Prof.
Pap. U.S. geol. Surv. 423, 1-67.

moore, r. c. (ed.) 1959. Treatise on Invertebrate Paleontology, Part O, Arthropoda I . Geol. Soc. Am.
and Univ. Kansas Press.

nolan, t. b., merriam, c. w., and williams J. s. 1956. The Stratigraphic section in the vicinity of
Eureka, Nevada. Prof. Pap. U.S. geol. Surv. 276, 1-77.
robison, r. a. and pantoja-alor, 5. 1968. Tremadocian trilobites from the Nochixtlan region,
Oaxaca, Mexico. /. Paleont. 42, 767-800, pis. 97-104.
ross, r. j. 1951. Stratigraphy of the Garden City Formation in north-eastern Utah and its trilobite
faunas. Bulk Peabody Mus. nat. Hist. 6, 1-161, pis. 1-36.

1957. Ordovician fossils from wells in the Williston Basin, eastern Montana. Bull. U.S. geol.

Surv. 1021-M, 439-510, pis. 37-44.

salter, j. w. 1866. On the fossils of North Wales. Appendix to The Geology of North Wales by A. C.
Ramsay. Mem. geol. Surv. U.K., 3, 239-363.

1881. In Salter and Etheridge, On the fossils of North Wales. Appendix to The Geology of North

Wales, 2nd edn, by A. C. Ramsay. Ibid. 331-567.

sando, w. j. 1958. Lower Ordovician section near Chambersburg, Pennsylvania. Bull. geol. Soc. Am.
69, 837-54, pis. 1, 2.

Whittington, h. b. 1966. Phylogeny and distribution of Ordovician trilobites. J. Paleont. 40, 696-737.

P. H. WHITWORTH

Department of Geology
The University
Birmingham, 15

Final typescript received 9 January 1969

A NEW SPECIES OF AULACOTHECA
(PTERIDOSPERMALES) FROM THE MIDDLE
PENNSYLVANIAN OF IOWA

by DONALD A. EGGERT and RALPH W. KRYDER

Abstract. Aulacotheca iowensis sp. nov. is described from the Middle Pennsylvanian of Iowa; it is distinct
from previously described forms in synangial dimensions, number of sporangia per synangium, and size range
of pollen grains. The Iowa material demonstrates that pollen organs of the Aulacotheca type were borne in
large numbers on ultimate axes of either an entirely fertile frond or portion of frond lacking planated foliar
structures. Orientation of the synangia suggests that the fertile regions were three-dimensional and bushy
prior to fossilization.

The genus Aulacotheca was instituted by Halle (1933) for certain synangiate pollen
organs believed to have been produced by medullosan pteridosperms. The exact nature
of remains assigned to this genus was entirely problematical until the work of Halle,
and included leaves (see, for example, White 1900) and seeds (Kidston 1890-8, Crookall
1929). Older generic designations of remains now assigned to Aulacotheca include
Whittleseya, Rhabdocarpus, and Holcospermum. Four of the six previously described
species (A. elongata (Kidston) Halle 1933, A. hemingwayi Halle 1933, A. campbelli
(White) Halle 1933, A. idelbergeri Halle 1933, A. dixiana Hemingway 1941, and A.
hallei Hemingway 1941) occur in the Upper Carboniferous of Great Britain. Aulaco-
theca idelbergeri occurs in continental Europe in strata designated as Westphalian A.
In North America, Aulacotheca has been reported from the Pottsville Series of the
Appalachian region (White 1900), the Pocahontas and New River Beds of West Virginia
(Jongmans 1937), and the Michigan coal basin (Arnold 1949). Arnold (1949) has dis-
cussed the occurrence of the genus in North America and has compared this material,
which is referred for the most part to the species A. campbelli , with that from Europe.
Methods of study. Elucidation of the various features of the specimen was carried out
by excavation of the individual synangia to insure completeness. Isolated synangia were
treated with either Schulze solution or dilute sodium hypochlorite (Clorox) to remove
dark occluding materials. Subsequent to these treatments the synangia were carried
through a standard dehydration series and mounted between coverslips so that both
sides of the synangium could be examined and photographed at relatively high mag-
nifications. Attempts to separate the individual pollen grains in the adherent masses were
unsuccessful so that individual pollen grains were observed and measured within the

EXPLANATION OF PLATE 76

Figs. 1-5. Aulacotheca iowensis sp. nov. 1, Over-all view of one-half of specimen (part) showing
general organization; 1324a, Xl. 2, Over-all view of counterpart of specimen after additional
excavation had been carried out; 1324b, X2. 3, Two synangia apparently borne as a pair;
1324b, x5-5. 4, Single synangium showing the narrow stalk; 1324b, X 8. 5, Isolated synangium
after treatment with Schulze solution; Slide 2934, x22.

[Palaeontology, Yol. 12, Part 3, 1969, pp. 414-9, pis. 76-77.]

Palaeontology, Vol. 12

PLATE 76

EGGERT and KRYDER, Aulacotheca (Pteridospermales)

D. A. EGGERT AND R. W. KRYDER: AULACOTHECA CPTERIDOSPERMALES) 415

pollen masses. Measurements parallel and perpendicular to the long axis were carried
out on a sample of 250 grains to determine the dimensional range, percentage distribu-
tion, and average dimensions of the pollen.

SYSTEMATIC SECTION

Order pteridospermales
?Family medullosaceae
Genus aulacotheca Halle 1933

Aulacotheca iowensis sp. nov.

Diagnosis. Fertile frond material lacking planated foliar structures, primary laterals
apparently borne alternately and having numerous groups of stalked synangia along
their entire lengths. Individual synangia about 5 mm. in length, 1-5 mm. in maximum
width, and having either 3 or 4 pollen sacs. Pollen sacs with elongate pollen masses of
adherent bilaterally symmetrical Monoletes- type pollen grains 88 (135) 165/x long by
44 (81) 121 p. broad.

Holotype. Specimen 1324a and b (part and counterpart), and slide preparations 2926 through
2980, Paleobotanical Collections, Department of Biological Sciences, University of Illinois at
Chicago Circle.

Stratigraphic position. Clay pit approximately | mile south-west of Redfield, Iowa (Sec. 5T 78N,
R29W, Adel Quadrangle), Cherokee Group (Des Moines Division).

Age. Middle Pennsylvanian (Des Moines Series).

Description. Initial cleavage of the matrix combined with subsequent excavation for
more complete exposure yielded an extensive mass of synangia approximately 8 cm.
in length (PI. 76, figs. 1, 2). The arrangement of the synangia suggests that the specimen
consisted of a major axis bearing approximately 6 primary laterals. These laterals,
apparently borne alternately along the major axis, bear the individual synangia. Little
in the way of clearly definable remains of the major axis are preserved, but the presence
of such a structure may be inferred from the orientation of the synangium-bearing
laterals. Each bears a large number of synangia which are radially disposed in the matrix
surrounding the position of the lateral axis. This apparent 3-dimensional disposition
was presumably a feature of the plant in life and suggests that this portion of the
frond was non-planated and probably had a rather bushy appearance. Synangia are
stalked (PI. 76, fig. 4) and in some instances borne in pairs (PI. 76, fig. 3). Individual
synangia are spatulate in outline, broadest just back of the tip, and gradually tapered
toward the base into a delicate stalk. Average dimensions of the synangia are 5 mm.
long and T5 mm. in maximum width. Relatively few, somewhat distantly spaced dark
lines run longitudinally along the synangia (PI. 76 fig. 3) and apparently represent
intervening walls between individual sporangia. As is seen in PI. 76, fig. 3 and 4, these
dark lines designate furrows that separate somewhat curved ridges. A number of rep-
resentative synangia are shown in place in the matrix in PI. 77, fig. 1.

Entire synangia consist of either 3 or 4 elongate masses of pollen (PI. 76, fig. 5,
PI. 77, fig. 5). Each mass apparently represents the contents of a single sporangium,
little else remains. In some instances, a thin layer of dark material is present covering

e e

C 6685

416

PALAEONTOLOGY, VOLUME 12

portions of the synangium (PI. 76, fig. 5), which when cleared in Clorox consists of a
single layer of extremely thin-walled cells that are axially elongate and have horizontally
orientated end walls. Whether this tissue represents all or part of the sporangial walls,
or is some sort of tapetal structure, is not known. Individual pollen masses illustrate
that the sporangial cavities were relatively narrow in comparison to their lengths and
reflected the exterior shape of the synangium in being broader near the tip (PI. 77, fig. 3)
and gently tapering toward the base of the synangium. Disposition of the pollen within
the masses (PI. 77, figs. 3, 4) further illustrates that the sporangial cavities were probably
circular in cross section rather than radially elongate. Material embedded in paraffin
and sectioned did not provide any pertinent information concerning the central portion
of the synangium.

Pollen grains of A. iowensis are similar to those found in a large number of pollen
organs thought to have affinities with the Medullosaceae. Grains are bilaterally sym-
metrical, have a prominent suture on the proximal face (PI. 77 fig. 2), and have lengths
that range from 88 to 165 /j. (text-fig. 1) and breadths from 44 to 121 p. The percentage
size distribution is shown in the text-figure. Average dimensions are 135 ju. long by 81 p.
broad (based upon 250 grains). The walls of the pollen grains are relatively thick (ap-
proximately 8-9 p) and are essentially devoid of ornamentation although a few grains
have an extremely faint punctation. In the dispersed form similar pollen would be
assigned to the genus Monoletes Schopf, Wilson, and Bentall 1944.

Discussion. Information concerning published species is restricted to features of in-
dividual synangia although we can now suggest certain other features of fertile regions
of A. iowensis. Of these species, A. elongata and A. hemingwayi have the most completely
known synangia. In these forms the number of sporangia (8-9 in A. elongata, 9 in
A. hemingwayi) and range of pollen dimensions are fairly well established.

Aulacotheca is extremely rare in occurrence in North America. Initially described under
the name Whittleseya campbelli by White (1900) on the basis of material from several
horizons within the Pottsville of the Appalachian region, North American material
has subsequently been assigned to Aulacotheca campbelli (Arnold 1947, 1949) in most
instances; however, Jongmans assigned material from the Pocahontas and New River
Beds of West Virginia to A. elongata and A. hemingwayi (Jongmans 1937). Material
from Eastwood, Michigan described under the name A. campbelli (Arnold 1949) is fairly
well known and in this instance the number of sporangia per synangium is 6. Additional
material described by Arnold from the Sewell Formation of West Virginia as A.
campbelli, and the material originally studied by White has not been shown to have
spores present nor has the number of sporangia per synangium been determined,
although it appears that Arnold’s estimate of 6 is probably correct in comparison with
the better preserved material from Michigan. The number of sporangia is also not

EXPLANATION OF PLATE 77

Figs. 1-5. Aulacotheca iowensis sp. nov. 1, Portion of specimen showing numerous representative
synangia; 1324a, x5-5. 2, Portion of pollen mass showing detail of individual grains; Slide

2928, X300. 3, Individual pollen mass after treatment with dilute Clorox; Slide 2928, x43.

4, Portion of pollen mass showing disposition of grains that suggests the tubular form of the pollen
sac; Slide 2928, X 120. 5, Individual synangium having three pollen sacs partially separated;

Slide 2926, X 22.

Palaeontology, Vol. 12

PLATE 77

EGGERT and KRYDER, Aulacotheca (Pteridospermales)

D. A. EGGERT AND R. W. KRYDER: AULACOTHECA (PTE R IDO SPERM ALES) 417

known in A. dixiana and A. hallei. Hemingway (1941) suggested that A. Hallei had 6
‘loculi’, but apparently believed them to be synangia rather than individual sporangia.
A. iowensis with 3 to 4 sporangia per synangium has a lower sporangial number than
any other species for which this number is well established. Evidence for either the
presence or absence of a central hollow region is totally lacking, in so far as we can

40

35

30

P
E
R
C
E
N
T

15

25

20

I 1

Width of grain

Length of grain

44 55 66 77 88 99 110 121 132 143 154 165

MIC RONS

text-fig. 1. Aulacotheca iowensis sp. nov. Histogram illustrating range and percentage distribution
of pollen grain dimensions; based on a sample of 250 grains.

judge, for all of the species except A. elongata. In the latter Halle (1933) inferred a
centrally placed hollow on the basis of the disposition of the spore masses after com-
pression and on the lack of evidence of any relatively thick carbonized area in the centre
of the synangium. In some instances, a longitudinal splitting of the flattened synangia
into two halves has been used to suggest the presence of the central hollow area, and
Halle seems to have been impressed with this feature of the material since it allowed
a close comparison to be made with Whittleseya , in which a ring of fused sporangia
surrounded a large central hollow area and made up a cup-shaped pollen organ that
was open at the distal end. In cases where synangia have relatively large numbers of
sporangia it can be argued from their disposition, as Halle did, that the sporangia
originally formed a ring about a central sterile region, either hollow or of solid tissue.
However, in forms such as A. iowensis where 3 or 4 sporangia are present, compression
results in a disposition of the sporangia which does not permit determination of the
exact nature of the central part of the synangium. Thus, largely through limitations of
the material, most of the described species of Aulacotheca cannot be shown to exhibit

418

PALAEONTOLOGY, VOLUME 12

the full complement of features assigned to this genus by Halle in the original diagnosis
(Halle 1933, p. 40). In practice, however, most material assigned to the genus shows a
similar external form, evidence of slight longitudinal ribbing, and most often a toothed
or lobed distal end. In this respect, material assigned to Aulaeotheca is more or less
indistinguishable from that placed in the genus Boulaya Carpentier (Carpentier 1925)
the most complete study of which was published by Halle (1933). In Halle’s study, the
genus Boulaya is characterized by having relatively fine vertical striations but lacking
evidence of any external ribs. In other respects, the genera Boulaya and Aulaeotheca
are indistinguishable on the basis of external features. The internal structure of Boulaya
has never been satisfactorily determined, although Halle believed it to have a central
hollow region surrounded by a ring-shaped sporangial region which may or may not
have been subdivided into individual sporangia. We have assigned the present material
from Iowa to Aulaeotheca largely on the presence of shallow external ribbing and the
clear presence of several individual sporangia within each synangium.

In 1932 Dix suggested that synangia of the Aulaeotheca type may have been attached
to Neuropteris sehlehani, but, this appears to be an instance of close association in the
matrix. Thus, at present, nothing is known of the source plants which produced pollen
organs such as Aulaeotheca. Relationships with the Medullosaceae are deduced from
the type of pollen and general organization of the pollen organs, which compare
favourably with some better-known pollen organs such as Dolerotheca which are almost
certainly of medullosan origin. However, the general organization of Aulaeotheca is
not very different from that occurring in some pollen organs believed to have been
produced by members of the Lyginopteridaceae. For example, both Telangium and
Heterotheca Benson (1904, 1922) are synangiate and the central portion of Telangium
was hollow. Fusion of the sporangia apparently extended throughout their lengths in
both Aulaeotheca and Heterotheca while in Telangium the sporangia were free except
near their bases. The genus Telangium , is, in turn, questionably distinct from some forms
of Crossotheca and Kidston believed that the two genera were probably the sam?
( Kidston 1 906). Thus, although general accounts of these genera of pollen organs generally
assume that the forms are distinct types such is not the case and a great deal of continued
research is needed to properly determine the detailed structure of these presumed
pteridosperm pollen organs.

A few remarks should be made concerning features of the fertile frond or frond regions
of A. iowensis. Whether the synangia occurred on separate fertile fronds or merely
occupied portions of the same fronds having sterile foliar areas cannot be determined.
It is of interest to note that present evidence indicates that Aulaeotheca pollen organs
were borne on pinnate fronds with unequal branching of the frond axes rather than on
fronds with equal dichotomous branching. Also, the fertile regions of Aulaeotheca
lacked sterile foliar structures and were apparently 3-dimensional in the distal regions
where the pollen organs were borne all around the ultimate axes. In this respect,
Aulaeotheca is reminiscent of several Devonian genera such as Archaeopteris and
Tetraxylopteris (Carluccio, Hueber, and Banks 1966, Bonamo and Banks 1967),
and, in view of several studies by Long on Lower Carboniferous pteridosperm fronds
and structures such as cupules borne on fronds, illustrates that the frond of
Palaeozoic seed ferns was much more 3-dimensional and branch-like than has been sup-
posed.

D. A. EGGERT AND R. W. KRYDER: AULACOTHECA fPTERIDOSPERM ALES) 419

Acknowledgement. This investigation was supported by National Science Foundation grant GB 4126
to the senior author.

REFERENCES

Arnold, c. a. 1947. An Introduction to Paleobotany. New York.

1949. Fossil flora of the Michigan coal basin. Contrib. Mas. Paleont. Univ. Michigan 7, 131-269.

benson, m. 1904. Telangium scotti, a new species of Telangium ( Calymmatotheca ) showing structure.
Ann. Bot. 18, 161-76.

1922. Heterotheca grievii. The microsporangium of Heterangiwn grievii. Bot. Gaz. 74, 121-42.

bonamo, p. m. and banks, h. p. 1967. Tetraxy/opteris schmidtii: its fertile parts and its relationships
within the Aneurophytales. Amer. J. Bot. 54, 755-68.
carluccio, l. m., hueber, f. m. and banks, h. p. 1966. Archaeopteris macilenta, anatomy and mor-
phology of its frond. Amer. J. Bot. 53, 719-30.

carpentier, a. 1925. Note sur quelques empreintes de graines et micro-sporanges de pteridospermees
provenant du westphalien du Nord de la France. Rev. Gen. Bot. 37, 145-56.
crookall, r. 1929. Coal Measure Plants. London.

dix, e. 1932. On a sporocarp probably attached to a frond of Neuropteris schlehani Stur. Ann. Bot. 46,
1065-8.

halle, t. G. 1933. The structure of certain fossil spore-bearing organs believed to belong to pterido-
sperms. Kungl. Svensk. Vetenskap. Hand!., Ser. 3, 12, 1-103.
hemingway, w. 1941. On the coal-measure plant Aulacotheca. Ann. Bot. N.s. 5, 197-201.
jongmans, w. 1937. Comparisons of the floral succession in the Carboniferous of West Virginia with
Europe. Compt. rend. 2cme congr. strat. Carb., Fleerlen, 1, 393-415.
kidston, r. 1890-8. The Yorkshire Carboniferous flora. Trans. York. Nat. Union, Volume for 1888-98.

1906. On the microsporangia of the Pteridospermeae, with remarks on their relationship to

existing groups. Trans. Roy. Soc. London, 248, 413-45.
schopf, j. m., wilson, l. r. and bentall, r. 1944. An annotated synopsis of Paleozoic fossil spores and
the definition of generic groups. Illinois Geol. Surv. Rept. Invest. 91.
white, d. 1900. The stratigraphic succession of the fossil floras of the Pottsville formation in the southern
anthracite coal field of Pennsylvania. U.S. Geol. Surv. Ann. Rept. 20, 749-930.

DONALD A. EGGERT
Department of Biological Sciences
University of Illinois at Chicago Circle,
Chicago, Illinois

RALPH W. KRYDER

Department of Geology
University of Iowa
Iowa City, Iowa

Typescript received 30 September 1968

MIOSPORES FROM THE LOWER
CARBONIFEROUS BASEMENT BEDS IN THE
MEN AI STRAITS REGION OF
CAERNARVONSHIRE, NORTH WALES

by f. a. hibbert and W. S. LACEY

Abstract. A well-preserved miospore flora from the Basement Beds of the Lower Carboniferous in the Menai
Straits region of Caernarvonshire, North Wales is described. A total of 47 species is recorded from the deposits.
One new genus Umbonatisporites, and 7 new species are proposed. The assemblage contains spores characteristic
of both Tournaisian and Visean deposits, but is considered to be Visean in age.

The Lower Carboniferous succession throughout North Wales consists largely of a
series of limestones underlain by Basement Beds and resting unconformably on Lower
Palaeozoic rocks. Lower Carboniferous deposits outcrop on both sides of the Menai
Straits and in Caernarvonshire lie on Ordovician rocks. Greenly (1928) described
conglomeratic sandstones, shales and thin limestones which he placed at the base of
the Lower Dibunophyllum zone (Dx). There is a fragmentary fauna, mainly of brachio-
pods; the lowest horizon containing abundant faunal remains lies close to the base
of the overlying Brown Limestone. There is no clear indication of the precise age of the
Basement Beds and they have been variously assigned to the base of the Da or the top
of the S2 (Greenly 1928, Neaverson 1946, George 1958).

Three samples were taken from a lenticle of shale, approximately 40 yards long and
3 ft. in thickness, where the Basement Beds outcrop by the Britannia Tubular Bridge on
the Caernarvonshire side of the Menai Straits (Grid Ref. SH541708).

Plant remains from these beds were first described by Walton in Greenly (1928), the
list of species later being extended by Lacey (1952 a, b). The later work indicated the
presence of a rich assemblage of plant micro-fossils and seeds.

The three samples were collected from the shale in the following ascending order:
sample LC2 from the base of the shale outcropping on the foreshore; sample LC3 one
foot above LC2 and associated with the plant bed described by Walton and Lacey;
LC4 at the top of the shale band two feet above LC3. The three samples showed no
marked differences in miospore content and are accordingly treated as one assemblage,
characteristic of the Basement Beds.

Preparation of samples. The samples were immersed in 40% hydrofluoric acid at 40 °C
for up to four days, to remove the silicates. The residue was oxidized in fuming nitric
acid for up to two hours, then washed with progressively more dilute nitric acid and
transferred to a sinter-glass Buchner funnel. Here the residue was further washed with a
5% solution of potassium hydroxide and then, repeatedly, with distilled water using the
technique described by Neves and Dale (1963).

Permanent slides were made using ‘Cellosize with a thermosetting plastic as a mountant
(Jeffords and Jones 1959).

(Palaeontology, Vol. 12, Part 3, 1969, pp. 420-440, pi. 78-83.]

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION 421

The terminology used is that outlined by Couper and Grebe (1961) and expanded
by Smith and Butterworth (1967). The classification of the dispersed miospores follows
the scheme first proposed by Dettmann (1963) as revised and extended by Smith and
Butterworth (1967).

Only those species which are described for the first time, or are considered to be more
critical to the present study, are given systematic treatment. In addition to the illustra-
tions using the transmitted light microscope, a number of photographs are reproduced
using the scanning reflection electron microscope developed by Cambridge Scientific
Instruments following the technique described by Hibbert (1967).

The slides containing holotypes and other figured specimens have been deposited in
the School of Plant Biology, University College of North Wales. They are marked with
the preparation number and the co-ordinates are those of the Leitz Laborlux micro-
scope no. 582096 of the above Department. Single grain mounts bear the prefix MS.

SYSTEMATIC DESCRIPTIONS

Anteturma sporites H. Potonie 1893
Turma triletes (Reinsch) Dettmann 1963
Suprasubturma acavatitriletes Dettmann 1963
Subturma azonotriletes (Luber) Dettmann 1963
Infraturma laevigati (Bennie and Kidston) Potonie 1956
Genus punctatisporites (Ibrahim) Potonie and Kremp 1954

Type species. P. punctatus Ibrahim 1933.

Punctatisporites irrasus Hacquebard 1957
Plate 78, fig. 1

Description. Diameter 58-89 p, mean 74 p (45 specimens); amb circular to sub-circular.
Laesura distinct, straight, length one-half to three-quarters spore radius, occasionally
low lips are developed. Frequently the laesura are gaping with dark intertectal areas.

Remarks. Spores with dark intertectal areas were included in this species by Sullivan
(1964a). It is thought to be a miospore characteristic of Tournaisian assemblages
(Sullivan 1967).

Previous records. Horton Bluff (Tournaisian) Canada (Hacquebard 1957). Lower
Limestone Shales (Tournaisian) Forest of Dean Gloucestershire (Sullivan 1964a).
Cementstone group (Tournaisian) of Ayrshire (Sullivan 1968). Springer formation
(Mississippian/Pennsylvanian boundary) of Oklahoma (Felix and Burbridge 1967).

Infraturma apiculati (Bennie and Kidston) R. Potonie 1956
Subinfraturma granulati Dybova and Jachowicz 1957
Genus granulatisporites (Ibrahim) Potonie and Kremp 1954

Type species. G. granulatus Ibrahim 1933.

422

PALAEONTOLOGY, VOLUME 12

Granulatisporites visensis sp. nov.

Plate 78, fig. 4

Holotype. Slide LS9b, 57.2 104.9. Size 41 p.

Diagnosis. Diameter 26-51 p, mean 37 p (56 specimens); amb subtriangular with concave
interradial margins and rounded apices. Laesura simple, straight, length from three
quarters to equal the spore radius. Ornamentation consists of grana 1 -5-4-0 p wide at
the base and up to 2-0 p high; the grana may coalesce to form short, verrucate ridges.
Ornament well developed at the apices where it forms an indented margin; the inter-
radial margins mostly smooth. The grana are most strongly developed on the distal
surface and are frequently concentrated at the distal pole and along the triangular radii.
Exine punctate between the grana.

Remarks. The development of irregular ridges characterises this species; its development
is not strong enough to warrant different generic assignment.

Subinfraturma verrucati Dybova and Jachowicz 1957
Genus verrucosisporites (Ibrahim) Smith and Butterworth 1967

Type species. V. verrucosus Ibrahim 1932.

Verrucosisporites eximius Playford 1962
Plate 78, figs. 9, 10

Remarks. The present specimens show a larger size from 62 to 92 p, mean 82 p than those
described by Playford (mean 72 p); otherwise they are similar.

Previous records. Lower Carboniferous of Spitsbergen (Playford 1962, 1963#).

Subinfraturma nodati Dybova and Jachowicz 1957
Genus waltzispora Staplin 1960

Type species. W. lobophora (Waltz) Staplin 1960.

EXPLANATION OF PLATE 78

All figures X 500

Fig. 1. Punctatisporites irrasus Hacquebard 1957; Slide LC5b, 21-3 110 0.

Fig. 2. Waltzispora planiangulata Sullivan 1964; Slide LC2e, 41-4 101-7.

Fig. 3. Lophotriletes tribulosus Sullivan 1964; Slide LC9b, 33-3 95-4.

Fig. 4. Granulatisporites visensis sp. nov., Holotype; Slide LC9b, 57-2 104-9.

Figs. 5-6. Raistrickia nigra Love 1960; slide MS80. 5, proximal surface. 6, distal surface.

Figs. 7-8. Neora'strickia drybrookensis Sullivan 1964. 7, slide LC2b, 23-0 100-2. 8, slide MS14.
Figs. 9-10. Verrucosisporites eximius Playford 1962. 9, proximal surface; slide MS144. 10, proxi-

mal surface; slide MS182.

Fig. 11. Grumosisporites verrucosus (B. and W.) Smith and Butterworth 1967; LC2c, 32-9 98-5.

Figs. 12-13. Umbonatisporites variabi/is gen. et sp. nov. 12, distal surface; slide LC2c, 14-6 106 8
13, Holotype, proximal surface; slide LC2e, 44 0 99 0.

Palaeontology , Vol. 12

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HIBBERT and LACEY, Early Carboniferous miospores

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION

423

Waltzispora planiangulata Sullivan 1964
Plate 78, fig. 2

Description. Diameter 30-41 p, mean 35 p (50 specimens); amb triangular with bluntly
rounded apices, having angular junctions with the concave, interradial margins. Laesura
distinct, simple, straight, length from three-quarters 1o equal to spore radius. Exine
1-0-1 -5 /x thick, ornamented with grana and coni, 0-5 p high and 1-0-1 -5 /x in basal
diameter; ornament absent from the proximal contact area.

Remarks. The variation in ornament between the specimens was not as evident as
Sullivan describes. The angular junction between the apex and the concave side was
variable but never approaches the distinct angularity described by Staplin (1960) for
W. lobophora.

Previous records. Drybrook Sandstone (Visean) Forest of Dean Basin, Gloucestershire
(Sullivan 19646).

Genus lophotriletes (Naumova) Potonie and Kremp 1954
Type species. L. gibbosus (Ibrahim) Potonie and Kremp 1954.

Lophotriletes tribulosus Sullivan 1964
Plate 78, fig. 3

Remarks. The size range, diameter 28-39 p, mean 32 p (40 specimens), varies from that
originally given by Sullivan (30-45 p, mean 36-5 p). Otherwise the range of ornament
in the present specimens agrees with the original description.

Previous records. Drybrook Sandstone (Visean) Forest of Dean Basin, Gloucestershire
(Sullivan 19646).

Genus umbonatisporites gen. nov.

Type species. U. variabilis sp. nov.

Diagnosis. Radial, trilete miospores, amb circular to sub-circular. Faesura simple,
straight, one sixth of the spore radius; frequently indistinct. Ornament of variable
shape, predominantly narrow at the base, widening towards the apex and terminating
in a rounded head, which is topped by a short, sharply tapering spine. There may be
from one to three rounded ‘heads’ on the apex of the element (text-fig. 1). There are
tapering spines interspersed over the surface of the spore. Exine is frequently folded.

Umbonatisporites variabilis sp. nov.

Plate 78, figs. 12, 13; Plate 79, figs. 1-3
Holotype. Slide LC2e, 44-4 90 0. Size 120 p.

Diagnosis. Diameter 95-134 p, mean 106 p (34 specimens); amb sub-circular to circular.
Faesura one-sixth spore radius, simple, frequently indistinct. Exine 1-2 p thick, covered

424

PALAEONTOLOGY, VOLUME 12

with a distinctive ornament arranged in indiscriminate patterns. Ornament variable
in both size and shape; one element up to 4-5 p high and 1-0-1 -5 p in basal diameter,
widening towards the apex where it terminates in a rounded head which is topped with
a thin tapering spine. There may be from one to three rounded projections at the apex
of the element. These elements are interspersed with spines 0-5-1 -0 p at the base and from
2-0 to 4-0 i x long. The exine is commonly folded.

Remarks. The only other spore showing variable branching at the apex of the elements
making up the ornament is the megaspore Singhisporites (Potonie 1956), ‘die terminal
± kleine Verzweigungen aufweisen’. There is no indication of the short terminal spine
seen in Umbonatisporites nor of tapering spines interspersed with the ‘bacula’. The
ornament in Singhisporites is frequently adpressed on to the spore body as is typical of
Umbonatisporites (PI. 79, figs. 2, 3).

text-fig. 1. Profile view of sculpture of Umbonatisporites variabitis
gen. et. sp. nov.

Subinfraturma baculati Dybova and Jachowicz 1957
Genus raistrickia (Schopf, Wilson, and Bentall) Potonie and Kremp 1954

Type species. R. grovensis Schopf 1944.

Raistrickia nigra Love 1960
Plate 78, figs. 5, 6

Remarks. The size range of the present specimens, from 48 to 67 p, mean 56 p (33
specimens) is smaller than that given by Love, the bacula are also of a smaller dimension.
Love comments that his description is based on only a small number of specimens and
it is considered that the present material represents an extension of his original description.
The sizes do not differ markedly from those given by Sullivan and Marshall (1966).

Previous records. Lower Oil Shale group (Visean) of Scotland (Love 1960). Upper
Sedimentary Group (Visean) of Scotland (Sullivan and Marshall 1966).

Raistrickia cf. clavata (Hacquebard) Playford 1963
Plate 79, figs. 4, 5

Description. Diameter 34-128 p, mean 109 p (30 specimens); amb circular. Laesura
straight, length two-thirds to three-quarters the spore radius, with slight lip develop-
ment. Exine 3-0-6-0 p thick (excluding ornament) covered with a variable ornament

HIBBERT AND LACEY: MIOSPORES FROM THE MEN AI STRAITS REGION 425

of verrucae, mushroom-shaped processes and bacula; their basal diameter varies from
5-5 to 8-0 p and height from 2-0 to 9-0 p. The ornament is irregular and occurs on both
faces of the spore.

Remarks. The character and positioning of the ornament in R. clavata (Hacquebard)
Playford 1963) is very similar to the present specimens. The size, however, is that of
R. ponder osa Playford 1963, which has less verrucae and a more uniform ornament.

Genus neoraistrickia Potonie 1956
Type species. N. truncatus (Cookson) Potonie 1956.

Neoraistrickia drybrookensis Sullivan 1964
Plate 78, figs. 7, 8

Description. Diameter 31-53 ft, mean 45 p (35 specimens); amb triangular with rounded
apices and straight to slightly concave, or convex sides. Laesura often indistinct,
straight, length three-quarters of spore radius; slight lip development. The distal face
of the spore is ornamented with cones, bacula, and verrucae. The coni are often blunt,
up to 3-0 p in height and 4-0 p in basal diameter; the bacula are up to 9-0 p high and
5-0 p in basal diameter and the verrucae from 3-0 to 7-0 p high and up to 9-0 p in basal
diameter. Exine 2-0-2-5 p thick.

Remarks. The specimens agree closely with the description given by Sullivan; the size
range is extended. The large verrucae when occurring on the equator, in particular to-
wards the triangular apices, give the impression that the spore has a flange.

Previous records. Drybrook Sandstone (Visean) Forest of Dean Basin, Gloucestershire
(Sullivan 19646).

Infraturma murornati Potonie and Kremp 1954
Genus convolutispora Hoffmeister, Staplin, and Malloy 1955

Type species. C. ftorida Hoffmeister, Staplin, and Malloy 1955.

Convolutispora labiata Playford 1962
Plate 79, figs. 8, 9

Remarks. Diameter 47-89 p , mean 64 p (50 specimens). The size of the miospores from
the Basement Beds is considerably smaller than those described by Playford, diameter
82-114 ft, mean 99 p. Apart from size difference the present specimens have the same
characteristics as Playford originally described and they are therefore placed in C. labiata.

Previous records. Lower Carboniferous of Spitsbergen (Playford 1962).

Convolutispora vermiformis Hughes and Playford 1961
Plate 79, figs. 6, 7

1957 Convolutispora flexuosa forma minor Hacquebard, p. 312; pi. 2, fig. 10.

426

PALAEONTOLOGY, VOLUME 12

Remarks. A number of the present specimens have lower, more insignificant muri than
was originally described by Hughes and Playford. They form a continuous morphological
series to the more typical form and were all included under C. vermiformis.

Previous records. Lower Carboniferous of Spitsbergen (Hughes and Playford 1961,
Playford 1962). Horton Group (Tournaisian) of Canada (Hacquebard 1957, Playford
1963). Upper Devonian of Melville Island (McGregor 1960). Springer formation (Missis-
sippian/Pennsylvanian boundary) of Oklahoma (Felix and Burbridge 1967).

Genus dictyotriletes (Naumova) Smith and Butterworth 1967
Type species. D. bireticulatus (Ibrahim) Potonie and Kremp 1954.

Dictyotriletes tesselatus sp. nov.

Plate 80, figs. 1, 2, 4, 5, 7, 8
Holotype. Slide LC3a, 55-7 93-2. Size 95 p.

Diagnosis. Diameter 78-105 p, mean 91 p (50 specimens); amb circular to sub-circular.
Laesura distinct, straight, length from three-quarters to equal to the spore radius;
accompanied by prominent lips up to 6-0 p broad on each side of the mark, having a
number of blunt crests up to 5-0 p high. Ornamentation on both faces of the spore of
smooth muri, 2-5-4 0 p wide and up to 11-0 p high, frequently with a clavate profile
when seen equatorially. The muri are frequently expanded where they anastamose, and
may terminate abruptly on the proximal face. Lumina very irregular in shape, from
5-0 to 27-0 p in longest diameter, there may be clavate projections within them. Exine
2-5 to 4-0 p thick (excluding ornament).

Comparison. Retieulatisporites variolatus Playford 1962 is characterized by a higher
frequency of more clavate muri when seen in profile. The lumina are more regularly
arranged and are rounded to polygonal in shape; the exine is also thicker and the laesura
is not accompanied by lips. R. cancellatus Playford 1962 has lower muri which are not
clavate in section.

EXPLANATION OF PLATE 79
All figures X 500 unless otherwise stated

Figs. 1-3. Umbonatisporites variabilis gen. et sp. nov. 1, Details of ornament; slide LC2c, 14-6
106-8; X 1000. 2, 3, Stereoscan pictures showing detail of ornament. 2, negative S/28/32,

x 5650. 3, negative S/28/37, x 5650.

Figs. 4-5. Raistrickia cf. clavata (Hacquebard) Playford 1963. 4, Distal surface; slide MS174.

5, Proximal surface; slide MS 177.

Figs. 6-9. Convolutispora spp. 6-7, C. vermiformis Hughes and Playford 1961; Slide MS153. 6,

Proximal surface. 7, Distal surface. 8-9, C. labiata Playford 1962; Slide LC3a, 32-8 96-2. 8,

Distal surface. 9, Proximal surface.

Figs. 10-11. Dictyotriletes spp. 10, D. pactilis Sullivan and Marshall 1966; Slide LC5c, 16-5 106-7.
1 1, D. cancellatus Playford 1962. Proximal surface, slide LC3d, 37-8 1 10-9.

Palaeontology , Vol. 12

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HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION

427

Dictyotriletes cancellatus (Waltz) Potonie and Kremp 1955
Plate 79, fig. 1 1

1938 Azonotriletes cancellatus Waltz, in Luber and Waltz, p. 1 1 ; pi. 1, fig. 8 and pi. 5, fig. 73.
1955 Sphenophyllotriletes cancellatus (Waltz) Luber, pp. 41-2, pi. 4, figs. 78 a, b, 79.

1955 Dictyotriletes cancellatus (Waltz) Potonie and Kremp, p. 108.

1956 Dictyotriletes cancellatus (Waltz) Ishchenko, p. 43; pi. 7, figs. 88, 89.

1957 Dictyotriletes cancellatus (Waltz) Naumova; Kedo, p. 1166.

1957 Reticulatisporites varioreticulatus Hacquebard and Barss, p. 17, pi. 2, figs. 15, 16.

1962 Reticulatisporites cancellatus (Waltz) Playford, pp. 597-8; pi. 82, figs. 11-13 and pi. 83,
figs. 1, 2.

Remarks. The inclusion of this species within the genus Dictyotriletes follows the emen-
dation of Reticulatisporites by Neves (1964) and the subsequent emendation of Dictyo-
triletes by Smith and Butterworth (1967). In the comparison of their new genus
Corbulispora with Dictyotriletes Bharadwaj and Venkatachala (1962) separate the two
on the basis of the latter having ‘flat muri ... a simple trilete mark’ (p. 24). There is
no valid reason for emphasizing the simple trilete mark as an important difference
between the two and it would seem that the interpretation of flat muri is not objective.
It would seem that these characteristics are not of sufficient significance to separate the
two genera. A more detailed study of the type material is needed to resolve the problem.

Previous records. Lower Carboniferous of the U.S.S.R. (Waltz in Luber and Waltz
1938, Luber 1955, Ishchenko 1956, 1958 and Kedo 1957, 1958). Lower Carboniferous
of Canada (Hacquebard and Barss 1957) and of Spitsbergen (Playford 1962).

Dictyotriletes pactilis Sullivan and Marshall 1966
Plate 80, fig. 10

Description. Diameter 62-105 p, mean 85 p (50 specimens); amb circular to sub-
circular. Laesura not seen. Ornament of thin, tall muri 0-5 to 2-0 p wide and up to
18-0 p high, clearly visible as radial projections at the equator. Lumina irregular in
shape, from 5-0 to 33-0 p in longest diameter. Muri frequently folded. Exine 2-0 to
4-0 p thick.

Remarks. In measuring eleven specimens Sullivan and Marshall gave a size range of
52-63 p, mean 58 p. On the basis of a greater number of specimens this size range is
extended. Reticulatisporites sp. B recorded by Love (1960) would seem to be D. pactilis.
Love records a size of 74 p for his specimen.

Previous records. Lower Oil Shale group (Visean) of Scotland (Love 1960). Upper
Sedimentary Group (Visean) of Scotland (Sullivan and Marshall 1966). Goddard
formation (upper Mississippian) of Oklahoma (Felix and Burbridge 1967).

Dictyotriletes submarginatus Playford 1963
Plate 80, figs. 3, 6, 11, 12.

Description. Diameter 52-69 p, mean 60 p (25 specimens); amb sub-triangular. Laesura
distinct, sinuous or straight, extending to the equator, accompanied by elevated lips up to

428

PALAEONTOLOGY, VOLUME 12

3 p wide. Proximal surface laevigate, occasionally the muri run on to the proximal surface
in the equatorial region. Distal surface ornamented with low, narrow, sinuous muri,
which may both anastamose and terminate freely; the lumina formed are irregular in
shape. Equatorial outline irregular to deeply indented.

Remarks. The spores described here agree closely with the original description given by
Playford, with the exception that the ornament of the distal surface appears to be less
dense and the incisions at the equator are deeper than his figured specimens. It is not
clear if the equatorial structure is a true cingulum, or is a feature produced by the fusion
of muri.

Previous records. Horton Group (Tournaisian) of Canada (Playford 1963).

Subturma zonotriletes Waltz 1935
Infraturma cingulati (Potonie and Klaus) Dettmann 1963
Genus knoxisporites (Potonie and Kremp) Neves and Playford 1961

Type species. K. hageni Potonie and Kremp 1954.

Knoxisporites stephanophorus Love 1961
Plate 80, figs. 9, 10

Remarks. Diameter 46-84 p, mean 68 p (20 specimens). The distal thickenings and dis-
tinctive structure of the lips, thinning proximally, are characteristic of this species.

Previous records. Lower Oil Shale group (Visean) of Scotland (Love 1960). Upper
Sedimentary group (Visean) of Scotland (Sullivan and Marshall 1966). Springer forma-
tion (Mississippian/Pennsylvanian boundary) and Goddard formation (Upper Missis-
sippi) of Oklahoma (Lelix and Burbridge 1967).

Knoxisporites pristinus Sullivan 1968
Plate 81, figs. 5, 6, 9

Description. Diameter 53-89 p, mean 68 p (27 specimens); amb circular to sub-circular,
frequently irregular. Laesura distinct, length from three-quarters to almost equal to

EXPLANATION OF PLATE 80

All figures X 500 unless otherwise stated

Figs. 1, 2, 4, 5, 7, 8. Dictyotriletes tesselatus sp. nov. 1, Holotype, distal surface; slide LC3a, 55-7
93-2. 2, Holotype, proximal surface, 4, Proximal surface; slide LC3c, 31 0 99-8 5, Distal

surface; slide LC3c, 31-0 99 8 7, Stereoscan, proximal surface; negative S/26/40, X 600. 8,

Stereoscan, distal surface; negative S/26/29, X 630.

Figs. 3, 6, 11, 12. Dictyotriletes submarg inatus Playford 1963. 3, Proximal surface; slide MS122.

6, Distal surface; slide MS122. 11, Distal surface; slide MS81. 12, Proximal surface; slide

MS81.

Figs. 9, 10. Knoxisporites stephanophorus Love 1961 ; Slide MS 108. 9, Distal surface. 10, Proximal

surface.

Figs. 13, 14. Knoxisporites seniradiatus Neves 1961; Slide MS91. 13, Proximal surface. 14,

Distal surface.

Palaeontology, Vol. 12

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HIBBERT and LACEY, Early Carboniferous miospores

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION 429

the radius of the central body, occasionally lips are developed. Exine thickened on the
distal surface, the thickenings are irregular in shape and frequently only slightly
developed.

Remarks. The present specimens clearly fit into the description given by Sullivan. The
variability and often ill-defined nature of the thickenings make it likely that many spores
which are rather badly preserved will be placed in this species. Certainly some of the
present material approximates to K. hederatus (Ishchenko) Play ford 1963 and K. rot alas
Hoffmeister, Staplin, and Malloy 1956.

Previous records. Cementstone group (Tournaisian) of Ayrshire (Sullivan 1968).

Knoxisporites seniradiatus Neves 1961
Plate 80, figs. 13, 14

Remarks. Although very few specimens of this spore were seen, they were clearly refer-
able to this species; the laesura having wide, prominent lips, so distinguishing the
specimens from K. triradiatus Hoffmeister, Staplin, and Malloy 1955. Sullivan (1964a)
records K. cf. triradiatus from Tournaisian deposits; in these specimens the trilete has
narrow lips, narrower than those of K. seniradiatus. This may rather be a representation
of the morphological range of K. seniradiatus.

Previous records. Namurian of the southern Pennines (Neves 1961).

Genus cincturasporites Hacquebard and Barss 1957
Type species. C. altilis Hacquebard and Barss 1957.

Remarks. This genus includes specimens which have a cingulum and a distinct convolute
ridge, or boss distal ornament. It is likely that the genus Orbisporis Bharadwaj and
Venkatachala 1962 does possess an equatorial cingulum, although the authors do not
describe such a feature; this, together with its variable distal ornament, makes it difficult
to separate from Cincturasporites. Critical reassessment of the type material of the genus
Orbisporis is necessary to resolve the problem.

Cincturasporites intestinalis sp. nov.

Plate 81, figs. 11-13; Plate 82, figs. 1-3

Holotype. MS 132. Size 130 p.

Diagnosis. Over-all diameter 92-143 p, mean 104 p (70 specimens); amb circular to
sub-circular. Laesura distinct, straight, length two-thirds to equal to the central body
radius, often gaping and frequently accompanied by a development of the proximal
ornament. Cingulum from 10-0 to 19-0 p in width, showing a poleward overlap onto
the central body; the equatorial amb is irregular and has several thickened lobes.
Cingulum is concentrically thickened, having a peripheral band of thickening and a
further band adjacent to the body with a thinner area between. The distal and, to a
lesser extent, the proximal faces of the central body are ornamented with convolute.

430

PALAEONTOLOGY, VOLUME 12

vermiform ridges, only rarely anastamosing, from 5-0 to 30-0 p in length and 4-0 to 9-0 p
in width. The central body is most often displaced laterally.

Remarks. Orbisporis convolutus Butterworth and Spinner 1967 is similar but has a
thickened band on the proximal side of the equator and lacks proximal ornament.
Cincturasporites sp. Balme and Hassell 1962 seems to approach the structure of C.
intestinal is.

Suprasubturma laminatitriletes Smith and Butterworth 1967
Subturma zonolaminatitriletes Smith and Butterworth 1967
Infraturma cingulicavati Smith and Butterworth 1967
Genus murospora Somers 1952

Type species. M. kosankei Somers 1952.

Murospora intorta (Waltz) Playford 1962
Plate 81, fig. 8

1938 Zonotriletes intortus Waltz, in Luber and Waltz, p. 22; pi. 2, fig. 24.

1954 Simozonotriletes intortus (Waltz) Potonie and Kremp, p. 159.

1956 Simozonotriletes intortus (Waltz) Ishchenko, pp. 88-9; pi. 17, fig. 204.

Description. Diameter 50-69 p, mean 58 p (30 specimens) ; amb subtriangular with straight
to concave sides and rounded apices. Laesura simple, distinct, straight, length from two-
thirds to equal to the spore body radius. Cingulum laevigate, 6-12 p wide, may be
thicker and wider at the apices, overlaps the central body on the proximal side.

Remarks. The validity of the generic assignment of this species remains in doubt.
Staplin (1960) showed that Murospora Somers, Simozonotriletes (Naumova) Potonie
and Kremp, and Westphalensisporites Alpern could be included in a single genus having
patellate and capsellate forms, the equatorial feature being a tightly attached but
separate part of the spore and not a centrifugal extension of the spore body. He did
not amend the diagnosis of Somers. It is not known what is the true nature of the equa-
torial structure in the type material of Simozonotriletes and until this is understood the
present specimens are placed in the genus Murospora following the work of Staplin.

EXPLANATION OF PLATE 81

All figures X 500 unless otherwise stated

Figs. 1-4. Lophozonotriletes muricatus sp. nov. 1, Holotype, proximal surface; slide LC5b 46 7
101-4. 2, Holotype, distal surface. 3, Proximal surface; slide MS29. 4, Distal surface; slide

MS29.

Figs. 5, 6, 7, 9. Knoxisporites spp. 5, 6, 9. K. pristinus Sullivan 1968. 5, Distal surface; slide

MS26. 6, Proximal surface; slide MS26. 9, Distal surface; slide MS219. 7, K. literatus (Waltz)

Playford 1962; Slide MS145.

Figs. 8, 10. Murospora spp. 8, M. intorta (Waltz) Playford 1962, Proximal surface; slide LC5a,
43-6 97T. 10, M. aurita (Waltz) Playford 1962, Proximal surface; slide LC7b, 33-4 101-6.

Figs. 11-13. Cincturasporites intestinalis sp. nov. 11, Stereoscan, proximal surface; negative
S/26/41, 360. 12, Holotype, distal surface; slide MSI 32. 13, Holotype, proximal surface.

Palaeontology , Vol. 12

PLATE 81

HIBBERT and LACEY, Early Carboniferous miospores

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION 431

Previous records. Widely recorded from the Lower Carboniferous of the U.S.S.R.
(Luber and Waltz 1938, Ishchenko 1956) and Spitsbergen (Playford 1962). The Upper
Carboniferous of Britain (Sullivan 1958) and Upper Mississippian of Canada (Playford
and Barss 1963).

Murospora aurita (Waltz) Playford 1962
Plate 81, fig. 10

1938 Zonotriletes auritus (Waltz) in Luber and Waltz, p. 17, pi. 2, fig. 23.

1956 Simozonotriletes auritus (Waltz) Potonie and Kremp, p. 109.

1957 Cincturasporites auritus (Waltz) Hacquebard and Barss, p. 23, pi. 3, fig. 1.

1957 Cincturasporites irregularis Hacquebard and Barss, pp. 25-6; pi. 3, fig. 19.

1960 Murospora varia Staplin, p. 30, pi 6, figs. 16, 18.

1960 Murospora sp. cf. varia Staplin, p. 30, pi. 6, fig. 19.

1962 Murospora aurita (Waltz) Playford, pp. 609-10, pi. 87, figs. 1-6; text figs. 6a-q, s, 7.

Description. Diameter 49-73 p, mean 59 p (30 specimens); amb sub-triangular, margin
smooth to undulating. Laesura distinct, straight, reaching to the equator of the central
body, accompanied by lips 2-5-6-0 p broad and slightly elevated. Cingulum from 5-0 to
13-0 p wide, laevigate, showing variation in thickening and in equatorial outline,
thickenings commonly situated at the radial apices.

Remarks. The overlap of the cingulum onto the central body is not considered to be a
constant feature of M. aurita by Playford; he rejects the assignment to Cincturasporites.
Certainly the continuous morphological series of cingulum width and thickness which
he describes is present in the Basement Bed material, cingulum overlap occurring in-
discriminately throughout this series.

Previous records. Lower Carboniferous of the U.S.S.R. (Luber and Waltz 1938, 1941).
Upper Mississippian of Canada (Hacquebard and Barss 1957, Playford and Barss
1963). Lower Carboniferous of Spitzbergen (Hughes and Playford 1961, Playford 1962.)

Genus lophozonotriletes (Naumova) Potonie 1958
Type species. L. lebedianensis Naumova 1953.

Remarks. Potonie (1958) includes spores in the genus Lophozonotriletes which were
cingulate and had a prominent verrucate ornament. Playford (1963a) found an overlap
of the cingulum onto the central body in rather less than a half of the specimens of
Cincturasporites appendices Hacquebard and Barss which he examined. He discounted
this overlap and placed the specimens in Lophozonotriletes.

Lophozonotriletes muricatus sp. nov.

Plate 81, figs. 1-4

Holotype. Slide LC5b, 46 7 101-4. Size 59 p.

Diagnosis. Over-all diameter 48-69 p, mean 58 p (55 specimens); amb sub-triangular
with convex sides and rounded apices. Laesura distinct, simple, straight, length from
three-quarters to equal to the central body radius. Cingulum from 11-0 to 20-0 p in

C 6685 p f

432

PALAEONTOLOGY, VOLUME 12

width. Distal surface of both the cingulum and the central body bears an ornament of
verrucae which may coalesce to form ridges, from 3-4 to 7-5 p in basal diameter.

Remarks. L. appendices (Hacquebard and Barss) Playford 1963, has an irregular distal
ornament which is not elongate and is only rarely coalescent; it is also larger (1 10-70 p).

Genus vallatisporites Hacquebard 1957
Type species. V. vallatus Hacquebard 1957.

Vallatisporites vallatus Hacquebard 1957
Plate 82, figs. 6, 13.

Remarks. The present specimens agree with the descriptions given both by Hacquebard
1957 and by Staplin and Jansonius (1964). There is a variability in the size of the vacuoles
which are not considered by Staplin and Jansonius to be of secondary origin but rather
as a specific character.

Previous records. Horton group (Tournaisian) of eastern Canada (Hacquebard 1957,
Playford 1963). Banff formation (Tournaisian) of Alberta (Staplin and Jansonius 1964).
Cementstone group (Tournaisian) of Ayrshire (Sullivan 1968).

Vallatisporites ciliaris (Luber) Sullivan 1964
Plate 82, fig. 8

1938 Zonotri/etes ciliaris Luber, in Luber and Waltz, p. 25, pi. 6, fig. 82.

1964 Vallatisporites ciliaris (Luber) Sullivan, p. 370, pi. 59, figs. 14, 15.

Remarks. Over-all diameter 52-77 p, mean 62 p (50 specimens). The ornament described
by Sullivan as galeae and spines is variable in size and density. In the present material
there seems to be a continuous morphological series between this species and V. cf.
ciliaris Sullivan 1964, in which the ornament is more or less completely absent. In this
series there is considerable variation in the size and shape of the vacuoles.

Previous records. Drybrook Sandstone (Visean) Forest of Dean Gloucestershire (Sullivan
1964). Bewcastle Beds (Upper Tournaisian/Lower Visean) of north-west England (Butter-
worth and Spinner 1967).

EXPLANATION OF PLATE 82
All figures X 500 unless otherwise stated

Figs. 1-3. Cincturasporites intestinalis sp. nov. 1, Distal surface, slide MS69. 2, Proximal surface,

slide MS69. 3, Stereoscan picture, distal surface, negative S/28/25, x 625.

Figs. 4, 5, 7, 9-12. Vallatisporites microgalearis sp. nov. 4, Holotype, proximal surface, slide LC2a
23-5 102-2. 5, Holotype, distal surface, slide LC2a 23-5 102-2. 7, Distal surface, slide MS231.

9, Distal surface, slide LC2c 28-9 95-5. 10, Proximal surface, slide LC2c 28-9 95-5. 11, Stereoscan

picture, distal surface, negative S/28/27, X 600. 12, Stereoscan picture, proximal surface, negative

S/28/26, X 600.

Figs. 6, 8, 13. Vallatisporites spp. 6, V. vallatus Hacquebard 1957, Proximal surface; slide MS248.
8, V. cilliaris (Luber) Sullivan 1964, Proximal surface; slide MS233. 13, V. vallatus with Lycospora

uber, stereoscan, proximal surface; negative S/26/2, X 650.

Palaeontology, Vol. 12

PLATE 82

HIBBERT and LACEY, Early Carboniferous miospores

HIBBERT AND LACEY: MIOSPORES FROM THE MEN A I STRAITS REGION

433

Val/atisporites microgalearis sp. nov.

Plate 82, figs. 4, 5, 7, 9-12
Holotype. Slide LC2a, 23.5 102.2. Size 56 p.

Diagnosis. Over-all diameter 39-59 p, mean 49 p (56 specimens); amb sub-triangular
with convex sides. Laesura indistinct obscured by sinuous, elevated lips which are equal
to the over-all radius of the spore. The distal surface of the central body and the cingulum
is ornamented with verrucae and galeae, the bases of which are often fused, from 1-5
to 4-0 ju, in basal diameter and up to 3-0 p in height. The cingulum is internally vacuolate;
in addition there are a number of vacuoles opening into the proximal face of the spore.

Remarks. This spore is smaller than V. galearis Sullivan 1964, and the distal ornament
is found both on the central body and the cingulum.

Suprasubturma perinotrilites (Erdtman) Dettmann 1963
Genus perotrilites (Erdtman) ex Couper 1953

Type species. P. granulatus Couper 1953.

Perotrilites magnus Elughes and Playford 1961
Plate 83, fig. 7

Remarks. Diameter 101-54 p, mean 123 p (20 specimens); the perine is torn away from
several of the present specimens, otherwise the spore is as described by Hughes and
Playford.

Previous records. Lower Carboniferous of Spitsbergen (Hughes and Playford 1961,
Playford 1962). Horton Group (Tournaisian) of eastern Canada (Playford 1963.)

Perotrilites perinatus Hughes and Playford 1961

Plate 83, fig. 6

Remarks. Diameter 61-86 p, mean 65 p (22 specimens). The folding of the perine gives a
wrinkled appearance to the spore which can assume a reticulate pattern. Punctations
occasionally seen on the perine are likely to be due to corrosion.

Previous records. Lower Carboniferous of Spitsbergen (Hughes and Playford 1961,
Playford 1962). Upper Sedimentary Group (Visean) of Scotland (Sullivan and Marshall
1966). Springer formation (Mississippian/Pennsylvanian boundary) of Oklahoma (Felix
and Burbridge 1967).

Suprasubturma pseudosaccititriletes Richardson 1965
Infraturma monopseudosacciti Smith and Butterworth 1967
Genus grandispora Hoffmeister, Staplin and Malloy 1955

Type species. G. spinosa Hoffmeister, Staplin and Malloy 1955.

434

PALAEONTOLOGY, VOLUME 12

Grandispora reticulatus sp. nov.

Plate 83, figs. 1, 2, 4, 5, 8

Holotype. Slide MS 192. Size 82 p.

Diagnosis. Over-all diameter 75-130 p, mean 101 p, diameter of central body 52-96 /x,
mean 72 /x (36 specimens) ; amb circular to sub-circular. Laesura distinct, may be obscured
by lips, straight, length one-half to three-quarters over-all diameter. Distal and proximal
faces ornamented with simple spines from 2-0 to 5-0 p in basal diameter and 5-0-15-0 p
high; they may have expanded bases which anastamose to form a reticulate pattern
over the surface of the spore. The exo-exine is strongly punctate and the central body is
distinct and laevigate.

Remarks. It is not clear whether the reticulate nature of the exo-exine is caused by the
development of the bases of the spines or by its thickening. The reticulate pattern is
most clearly seen in those specimens where the ornament is well developed and rather
crowded. Grandispora sp. A of Sullivan and Marshall 1966 is likely to be G. reticulatus.
The size and disposition of the ornament distinguish G. reticulatus from other species
in the genus.

Genus hymenozonotriletes (Naumova 1937?, 1939) Potonie 1958
Type species. H. polyacanthus Naumova 1953.

Hymenozonotriletes? hastulus Sullivan 1968
Plate 83, fig. 3

Description. Diameter 51-72 p, mean 60 p (35 specimens); amb circular to rounded
triangular. Laesura indistinct, obscured by raised lips which are sinuous, length equal
to the radius of the central body or reaching onto the cingulum. The distal face of the
central body is ornamented with small cones grading into well-developed spines on the
cingulum which are 1 -5-4-0 p in basal diameter and from 2-5 to 7-0 p high, spines fre-
quently have a swollen base. Proximal face of the central body laevigate to finely punctate.
Spines occur on the proximal face of the cingulum.

Remarks. The genus cannot be placed with any certainty in the present system of classifi-
cation because details of the structure and exine stratification of the type species is not
fully known. The designation of these specimens as Hymenozonotriletes follows Sullivan
(1968).

Previous records. Cementstone group (Tournaisian) of Ayrshire (Sullivan 1968).

EXPLANATION OF PLATE 83
All figures X 500 unless otherwise stated

Figs. 1, 2, 4, 5, 10. Grandispora reticulatus sp. nov. 1, Holotype, distal surface; slide MS192.
2, Distal surface; slide MS 187. 4, Stereoscan, negative S/26/32, X1900. 5, Stereoscan, detail of

ornament; negative S/26/31, X630. 8, Proximal surface; slide MS76.

Figs. 3, 8. Hymenozonotriletes? hastulus Sullivan 1968. 3, Slide MS9. 8, Slide MS6.

Fig. 6. Tetrapterites visensis Sullivan and Hibbert 1964, Stereoscan, negative S/26/35, x 190.

Fig. 7, 9. Perotrilites spp. 7, P. magnus Hughes and Playford 1961; Slide MS147. 9, P. perinatus

Hughes and Playford 1961; Slide LC2e, 43-4 99-1.

Palaeontology, Vol. 12

PLATE 83

HIBBERT and LACEY, Early Carboniferous miospores

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION

435

THE MIOSPORE ASSEMBLAGE FROM THE BASEMENT BEDS

The assemblage from the Basement Beds contained a total of 47 species, as shown in
Table 1. It was dominated by Lycospora uber; species of Pimctatisporites and V al/atis-
porites were present in quantities greater than 10% of the total (500 spores were counted).
Cyclogranisporites lasius and species of Leiotriletes and Dictyotriletes had representation
between 1 and 8%, the remainder individually contributing less than 1% of the total.

The assemblage contains miospores which are restricted elsewhere to assemblages of
Tournaisian age: Dictyotriletes submar ginatus, Knoxisporites pristinus, Vallatisporites
vallatus, whilst others have been recorded mainly from deposits of Visean age : Leiotriletes
tumicius , Waltzispora planiangulata, Lophotriletes tribulosus, Raistrickia nigra , Neorai-
strickia drybrookensis, Convolutispora mellita, Dictyotriletes pactilis, Knoxisporites
stephanophorus, Gruniosisporites verrucosus, Remysporites magnificus, and Tetrapterites
visensis.

COMPARISON WITH OTHER LOWER CARBONIFEROUS
MIOSPORE ASSEMBLAGES

Comparisons between the miospore assemblage from the Basement Beds and others
of Lower Carboniferous age are set out in Table 1. This table is compiled from data
of the following authors: Butterworth and Williams 1958, Felix and Burbridge 1967,
Flacquebard 1957, Flacquebard and Barss 1957, Hoffmeister, Staplin and Malloy 1955,
Ishchenko 1958, Kedo 1958, Love 1960, Luber 1955, Luber and Waltz 1938, Playford
1962, 1963a, 19636, Playford and Barss 1963, Smith and Butterworth 1967, Staplin
1960, Sullivan 1964a, 1964 b, and Sullivan and Marshall 1966.

The correlation and zonation for the Carboniferous follows that set out by Francis
and Woodland (1964) in Table 1, p. 222. Correlations between faunal assemblages is that
of Prentice and Thomas (1965), and the S2/D! boundary is used to define upper from
lower Visean (Murray Mitchell pers. comm.).

The assemblage from the Caernarvonshire Basement Beds closely resembles that
described by Sullivan (1964) from the Drybrook Sandstone of the Forest of Dean,
which is Visean in age (S2). There are species which are only recorded from these two
deposits, namely Lophotriletes tribulosus Sullivan 1964, Waltzispora planiangulata
Sullivan 1964, Neoraistrickia drybrookensis Sullivan 1964, and Tetrapterites visensis
Sullivan and Hibbert 1964. Both of these deposits lack characteristic species which are
recorded from the Visean-Namurian deposits of the north of England and Scotland
(Sullivan and Marshall 1966).

The assemblages described by Knox 1948, Butterworth and Williams 1958, Love 1960,
Sullivan and Marshall 1966, Owens and Burgess 1965, and Butterworth and Spinner
1967 complete a range from middle Visean to Namurian A and have been grouped
together, on the basis of spore content, as the Grandispora suite by Sullivan (1965, 1967).
This suite has also been recognized from the mid continent of U.S.A., Spain, Poland,
Czechoslovakia, Romania, and Turkey (Sullivan 1967). None of the characteristic
species of this suite are recorded from the Basement Beds but other species commonly
found in the Grandispora suite do occur. Raistrickia nigra Love 1960 and Dictyotriletes
pactilis Sullivan and Marshall 1966 both indicative of upper Visean deposits in Scotland,

436

PALAEONTOLOGY, VOLUME 12

TABLE 1

Stratigraphic distribution of miospores found in the Menai Straits assemblage. Broken line indicates
uncertainty regarding stratigraphic dating or precise age limits; B & W = Butterworth and Williams.
Stratigraphic time scale as defined in Harland, W.B., et al, (eds.) 1967, The Fossil Record, Geological

Society of London.

PScu,

TOURNAISIAN

VISEAN

LOWER UPPER

NAMURIAN

BASHKIRIAN

LEIOTRILETES INERMIS (Waltz) Ishchenko 1952

L, SUBINTORTUS (Waltz) Ishchenko 1952

L. ORNATUS Ishchenko 1956

L TUMIDUS Butterworth & Williams 1958

PUNCTATISPORITES GLABER (Naumova) Playford 1962

P IRRASUS Hacquebard 1957

CALAMOSPORA MlCRORUGOSA(lbrahim)Schopf, Wilson & Bentall 1944

GRANULATISPORITES GRANULATUS Ibrahim 1933

G. MICROGRANIFER Ibrahim 1933

G. VISENSIS

sp, nov

CYCLOGRANISPORITES LASIUS (Waltz) Playford 1962

VERRUCOSISPORITES EXIMIUS Playford 1962

WALTZISPORA PLANIANGULATA Sullivan 1964

LOPHOTRILETES TRIBULOSUS Sullivan 1964

UMBONATISPORITES VARIABILIS gen. et sp. nov.

RAISTRICKIA NIGRA Love 1960

R. c.f. CLAVATA Playford 1963

NEORAISTRICKIA DRYBROOKENSIS Sullivan 1964

CONVOLUTISPORA TUBE RCULATA (Waltz) HoffmeisterStaplin&Mallcy1955

C. LABIATA Playford 1962

C. VERMIFORMIS Hughes & Playford 1961

C. MELLITA Hoffmeister, Staplin & Malloy 1955

DICTYOTRILETES CANCELLATUS (Waltz) Potonie & Kremp 1955

D. PACTILIS Sullivan 8 Marshall 1966

D, TESSELATUS sp. nov

D. SUBMARGINATUS Playford 1963

D. c.f. PELTATUS Playford 1962

KNOXISPORITES PRISTINUS Sullivan 1968

K. STEPHANOPHORUS Love 1960

K. SENIRADIATUS Neves 1961

K. LITERATUS (Waltz) Playford 1963

CINCTURASPORITES INTESTINALIS sp. nov.

MUROSPORA INTORTA (Waltz) Playford 1962

M, AURITA (Waltz) Playford 1962

GRUMOSISPORITES VERRUCOSUS (B. & W.) Smiths Butterworth 1967

LYCOSPORA UBER (Hoffmeister, Staplin & Malloy) Staplin 1960

LOPHOZONOTRILETES MURICATUS sp.nov

VALLATISPORITES VALLATUS Hacquebard 1957

V. CILLIARIS (Luber) Sullivan 1964

V. MICROGALEARIS sp nov

PEROTRILITES MAGNUS Hughes & Playford 1961

PERINATUS Hughes & Playford 1961

REMYSPORITES MAGNIFICUS (Horst) Butterworth 8 Williams 1958

ENDOSPORITES MICROMANIFESTUS Hacquebard 1957

GRANDISPORA RETICULATUS sp. nov.

HYMENOZONOTRILETES ? HASTULUS Sullivan 1968

TETRAPTERITES VISENSIS Sullivan & Hibbert 1964

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION 437

occur in the Basement Beds, as does Remysporites magnificus (Horst) Butterworth and
Williams 1958, restricted to Namurian A and younger deposits in Scotland.

Similarities also exist between the assemblage from the Basement Beds and the ‘Aurita’
assemblage described by Playford (1963a), from Spitsbergen. This latter assemblage
is characterized by zonate spores and is thought to be of Visean, possibly Namurian
age. Many of the species of this assemblage are found in North Wales, notably Ver-
rucosisporites eximius Playford 1962, Convolutispora labiata Playford 1962, C. vermi-
formis Hughes and Playford 1961, Murospora aurita (Waltz) Playford 1962, and M.
intorta (Waltz) Playford 1962, all of which have not been previously recorded in the
United Kingdom, also Dictyotriletes canceUatus (Waltz) Potonie and Kremp 1966,
Convolutispora tuberculata (Waltz) Hoffmeister, Staplin, and Malloy 1955, and Pero-
trilites magnus Hughes and Playford 1961 which have previously been recorded from
Tournaisian and Visean deposits of the British Isles. According to Sullivan (1967)
elements of the Spitsbergen miospore flora place it in his Monilospora suite.

The assemblages described from the Horton Group of Eastern Canada have similari-
ties with the flora of the Basement Beds, Dictyotriletes submarginatus Playford, Rais-
trickia cf. clavata, Perotrilites perinatus, and P. magnus being conspicuous in both
assemblages.

AGE OF THE BASEMENT BEDS

The miospore assemblage from the Basement Bed has affinities with other assemblages
of Visean age, in particular that of the Drybrook Sandstone (Sullivan 1964). It also
contains important species characteristic of the Tournaisian Vallatisporites suite (Sullivan
1968) and is similar to the Visean miospore floras from Scotland (Love 1960, Sullivan
and Marshall 1966), both of which are assigned to the Grandispora suite, and the "Aurita’
assemblage from Spitsbergen (Playford 1962, 1963a).

There is very little published information on lower to mid-Visean miospore floras with
which comparisons can be made. Also there is no opportunity to extend the strati-
graphic range of Lower Carboniferous fossils in the Menai Straits area because of the
limited nature of the deposits.

The appearance of miospores typical of Tournaisian assemblages is likely to be due to
reworking (Wilson 1962). The Basement Group of the Carboniferous Limestone is a
transgressive sequence and a shale lenticle in a sandstone-conglomerate is a situation
where such reworking would take place. Yet these spores are not less well preserved
than those more typical of the later deposits. The presence of the widely recorded
miospore Lycospora uber may be taken as evidence that the deposits are of Visean,
rather than Tournaisian, age. The presence of other upper Visean to Namurian mio-
spores would place a minimum age on the deposit without entirely excluding the possi-
bility that further work on early Visean deposits may extend the range of some of these
spores. Since, however, many of the typical late Visean types are absent it is believed that
the Basement Beds are of early upper Visean age.

This agrees well with the tentative Sa-Dj^ age assigned to the deposits on the basis
of fragmentary faunal evidence and the middle to upper Visean age suggested for the
overlying Brown Limestone elsewhere in North Wales on the basis of floral and faunal
macrofossil evidence (Lacey 1962).

438

PALAEONTOLOGY, VOLUME 12

Acknowledgements. The authors thank Professor P. W. Richards for the use of facilities in the Univer-
sity College of North Wales. Their thanks are also due to Dr. R. Neves, Dr. H. J. Sullivan, and
Dr. M. A. Butterworth for their advice and assistance at various times during the course of the work.
Cambridge Scientific Instruments provided valuable help with the scanning electron microscope.
This paper is part of a thesis prepared by F. A. H. while in receipt of a D.S.I.R. Studentship.

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New Phytologist, 66, 825-6, 1 pi.

hoffmeister, w. s., staplin, f. l. and malloy, R. E. 1955. Mississippian plant spores from the Hardins-
burgh Formation of Illinois and Kentucky. J. Paleont. 29, 372-99, 4 pi.
hughes, n. f. and playford, g. 1961. Palynological reconaissance of the Lower Carboniferous of
Spitsbergen. Micropaleontology, 7, 27-44, 4 pi.

Ishchenko, a. m. 1956. Spores and pollen of Lower Carboniferous deposits of the western extension
of the Donets Basin. Tr. Inst. geol. nauk, Akad. nauk Ukrainsk. S.S.R., ser. strat. paleont. 11,
1-185, 20 pi. (in Russian).

— — 1958. Sporo-pollen analysis of the Lower Carboniferous deposits of the Dnieper-Donets Basin.
Ibid. 17, 1-188, 13 pi. (in Russian).

jeffords, r. m. and jones, d. h. 1959. Preparation of slides for spores and other microfossils. J. Paleont.
33, 344-47.

kedo, G. i. 1957. On the stratigraphy and spore-pollen complexes of the lower horizons of the Carboni-
ferous in the B.S.S.R. Dokl. Akad. nauk S.S.S.R. 115, 1165-8 (in Russian).

• 1958. Characteristic spores and pollen of the lower horizons of the Carboniferous in the B.S.S.R.

Tr. Inst. geol. nauk, Akad. nauk B.S.S.R. 1, 44-56 (in Russian).
knox, e. m. 1948. The microspores in coals of the Limestone Group in Scotland. Trans. Inst. Min.
Engrs., Lond. 101, 98-112, 4 figs.

lacey, w. s. 1952a. Correlation of the Lower Brown Limestone of North Wales with part of the Lower
Carboniferous succession in Scotland and Northern England. Rep. 18th Int. Geol. Congr. Gt. Brit.
(1948), 10, 18-25.

HIBBERT AND LACEY: MIOSPORES FROM THE MENAI STRAITS REGION 439

lacey, w. s. 1952 b. Additions to the Lower Carboniferous flora of North Wales. C. R. 3rd Congr.
Strat. Geol. Carb. Heerlen (1951), 2, 375-7.

1962. Welsh Lower Carboniferous plants. I. The flora of the Lower Brown Limestone in the Vale

of Clwyd, North Wales. Palaeontographica, 111B, 126-60, 4 pi.
love, l. g. 1960. Assemblages of small spores from the Lower Oil Shale Group of Scotland. Proc.
Roy. Soc. Edinb. 67, 99-126, 2 pi.

luber, A. a. 1955. Atlas of the spore and pollen grains of the Palaeozoic deposits of Kazakhstan. Izd.
Akad. nauk Kazakh. S.S.R., Alma-Ata, 1-125 (in Russian).

and waltz, i. e. 1938. Classification and stratigraphical value of the spores of some Carboniferous

coal deposits in the U.S.S.R. Trans. Central Geol. Prosp. Inst. 105, 1-45, 10 pi. (in Russian).

■ 1941. Atlas of microspores and pollen grains of the U.S.S.R. Tr. All-Union Geol. Sci. Res.

Inst. ( V.S.E.G.E.I .) 139, 1-107, 16 pi. (in Russian).
mcgregor, d. c. 1960. Devonian spores from Melville Island, Canadian Arctic Archipelago. Palaeon-
tology, 3, 26-44, 3 pi.

nea verson, e. 1946. The Carboniferous Limestone Series of North Wales; conditions of deposition
and interpretation of its history. Proc. Liv. geol. Soc. 19, 113-44.
neves, r. 1961. Namurian plant spores from the southern Pennines. Palaeontology, 4, 247-79, 5 pi.

1964. The ‘Dispersed Spore’ genus Knoxisporites (Potonie and Kremp) Neves 1961. C.R. 5th

Congr. Strat. Geol. Carb. Paris, 1063-8, 1 pi.

and dale, b. 1963. A modified filtration system for palynological preparation. Nature, 198, 775.

owens, b. and burgess, i. c. 1965. The stratigraphy and palynology of the Upper Carboniferous outlier
of Stainmore, Westmorland. Bull. geol. Surv. Gt. Brit. 23, 17-44, 2 pi.
playford, g. 1962. Lower Carboniferous microfloras of Spitsbergen, Part 1. Palaeontology, 5, 550-
618, 10 pi.

1963a. Idem. Part 2, Ibid. 5, 619-78, 8 pi.

1963b. Miospores from the Mississippian Horton Group, Eastern Canada. Bull. geol. Surv. Can.

107, 47 pp„ 11 pi.

and barss, m. s. 1963. Upper Mississippian Microflora from Axel Heiberg Island, District of

Franklin. Geol. Surv. Pap. Can. 62-36, 5 pp.

potonie, r. 1956. Synopsis der Gattungen der Sporae dispersae. I. Teil: Sporites. Beih. Geol. Jahrb.
23, 1-103, 11 pi.

1958. Synopsis der Gattungen der Sporae dispersae. II. Teil: Sporites (Nachtrage), Saccites,

Aletes, Praecolpates, Polyplicates, Monocolpates. Ibid. 39, 1-189, 11 pi.

and kremp, g. 1955. Die Sporae dispersae des Ruhrkarbons, ihre Morphographie und Strati-

graphie mit Ausblicken auf Arten anderer Gebiete und Zeitabschnitte: Teil 1. Palaeontographica,
98B, 1-136, 16 pi.

1956. Idem. Teil 2. Ibid. 99B, 85-191, 22 pi.

prentice, j. e. and thomas, j. m. 1965. Prolecanitina from the Carboniferous rocks of North Devon.
Proc. Yorks, geol. Soc. 35, 34-46, 2 pi.

schopf, j. m., wilson, l. r. and bentall, r. 1944. An annotated synopsis of Paleozoic fossil spores
and the definition of generic groups. Rep. Inv. III. State geol. Surv. 91, 1-72, 3 pi.
smith, a. h. v. and butterworth, m. a. 1967. Miospores in the coal seams of the Carboniferous of
Great Britain. Spec. Paper Palaeont. 1, 324 pp., 27 pi.
somers, e. 1952. A preliminary study of the fossil spore content of the lower Jubilee seam of the Sydney
coalfield, Nova Scotia. Publ. Nova Scotia Found., Halifax, 1-30.
staplin, f. l. I960. Upper Mississippian plant spores from the Golata Formation, Alberta, Canada.
Palaeontographica, 107B, 1-40, 8 pi.

— and jansonius, j. 1964. Elucidation of some Paelaozoic Densospores. Ibid. 114B, 95-117, 4 pi.
Sullivan, h. j. 1958. The microspore genus Simozonotriletes. Palaeontology, 1, 125-38, 3 pi.

— — 1964a. Miospores from the Lower Limestone Shales (Tournaisian) of the Forest of Dean Basin,
Gloucestershire. C.R. 5th Congr. Strat. Geol. Carb. Paris (1963), 3, 1249-58, 2 pi.

19646. Miospores from the Drybrook Sandstone and associated measures in the Forest of Dean

Basin, Gloucestershire. Palaeontology, 7, 352-92, 5 pi.

1965. Palynological evidence concerning the regional differentiation of Upper Mississippian

floras. Pollen et Spores, 7, 539-63, 2 pi.

440

PALAEONTOLOGY, VOLUME 12

sullivan, h. j. 1967. Regional differences in Mississippian spore assemblages. Rev. Palaeobot. Palynol.
1, 185-92.

— 1968. A Tournaisian spore flora from the Cementstone Group of Ayrshire, Scotland. Palaeon-
tology, II, 116-31, 3 pi.

— and hibbert, a. f. 1964. Tetrapterites visensis, a new spore bearing structure from the Lower
Carboniferous. Ibid. 7, 64-71, 2 pi.

and marshall, a. E. 1966. Visean spores from Scotland. Micropaleontology, 12, 265-85, 4 pi.

wilson, l. r. 1964. Recycling, Stratigraphic Leakage, and Faulty Techniques in Palynology. Grana
Palyn. 5, 425-36.

winslow, m. r. 1959. Upper Mississippian and Pennsylvanian megaspores and other plant micro-
fossils from Illinois. Bull. III. State geol. Surv. 86, 135 pp., 16 pi.

F. ALAN HIBBERT

Department of Botany
University of Cambridge
Cambridge
WILLIAM S. LACEY

School of Plant Biology
University College of North Wales

Typescript received 4 October 1968 Bangor

MEGASPORE ASSEMBLAGES FROM VISEAN
DEPOSITS AT DUNBAR, EAST LOTHIAN,

SCOTLAND

by EDWIN SPINNER

Abstract. Megaspore assemblages are described from two coal seams in the lower Limestone Group. Two new
species are proposed; Setosisporites pseudoreticulatus and Zonalesporites fusinatus. The genera Lagenicula
(Bennie and Kidston) Potonie and Kremp 1954 and Setosisporites (Ibrahim) Potonie and Kremp 1954 are
emended. Two new combinations are proposed: Setosisporites indianensis (Chaloner), and Setosisporites
splendidus (Zerndt). The assemblages are compared with published records of megaspores from North America
and Europe.

Dunbar, East Lothian, is situated on the east coast of southern Scotland, some 25
miles east of Edinburgh. Exposed on the foreshore approximately 2 miles east of the
town centre near the Barns Ness lighthouse are a group of alternating limestones, shales
and sandstones, the prominent members of which are named as follows (from a detailed
succession given by Lumsden, in press) :

Upper Skateraw Limestone
shales etc.

Middle Skateraw Limestone
Skateraw Coal
shales, sandstones etc.

Lower Skateraw Limestone 25 metres
shales etc.

Upper Longcraig Limestone
shales etc.

Longcraig Coal

Middle Longcraig Limestone

Of two thin coal seams, approximately 10-22 cm. thick, the lower, named here Long-
craig Coal, can be seen at the base of a low ridge (high-water mark) formed by the over-
lying Upper Longcraig Limestone (NT 7 1 55 7730). The upper seam, named here Skateraw
Coal, occurs 5-6 m. above the Longcraig seam in the succession and crops out on a small
promontory 200 m. farther east towards the lighthouse. Both localities are clearly
indicated on the geological map permanently exhibited on a wall in the derelict lime-
stone kiln situated just above the shoreline as well as by marker pegs on the ground.

This succession is locally assigned to the Lower Limestone Group of the Carboniferous
Limestone and is probably upper l3! or basal P2 age (pers. comm. Dr. R. B. Wilson,
Institute of Geological Sciences, Edinburgh, based on the tentative correlation of the
Upper Longcraig Limestone with the Hurlet Limestone of the Glasgow area; see also
Currie 1954, p. 533). By its geographical position, this succession probably represents
part of a link between the depositional areas of the Midland Valley of Scotland, and the
Northumberland Trough of England in Upper Visean-Namurian time.

[Palaeontology, Vol. 12, Part 3, 1969, pp. 441-458, pis. 84-86.]

442

PALAEONTOLOGY, VOLUME 12

The succession was brought to the writer's attention by Dr. R. H. Wagner as a result
of a field excursion, led by A. Davies and D. C. Greig (Institute of Geological Sciences)
and subsequent discussion of the stratigraphic age of the strata.

Although the work began as a study of the presence, preservation, and variety of
megaspores in this deposit and of the possibilities of changes in megaspore assemblages
as an aid in the subdivision and correlation of Visean deposits in this area, it was found
during the work that some revision of the systematic descriptions was desirable, and
that some new taxa were necessary. A comparative account is also given of published
records of megaspore assemblages from deposits of Lower Carboniferous age in other
areas.

Channel samples were taken of both seams and megaspores were obtained by using
Schulze’s solution (2 days) followed by repeated washings with 5% Potassium hydroxide
solution. Specimens were prepared for both transmitted and reflected light examination.

SYSTEMATIC DESCRIPTIONS

No suprageneric classification is employed, and the arrangement is in alphabetical
order. The type material is deposited in the reference collections, Microplaeontology
Laboratory, Geology Department, University of Sheffield, England.

Genus lagenicula (Bennie and Kidston) Potonie and Kremp emend.

1 954 Lagenoisporites Potonie and Kremp, p. 1 52.

1962 Rostratispora Bharadwaj and Venkatachala, p. 25.

Type species. Lagenicula horrida Zerndt 1 934.

Emended diagnosis. Trilete megaspores with an apical prominence formed by the
progressive expansion of the laesurae along most of their length from the junction with
the curvaturae to the proximal pole, and the thickening of the exine of the greater part
of the contact areas.

Description. This expansion often, but not necessarily, results in a longer polar than
equatorial axis of the spore body. There is a tendency for lateral compression to be
most common, but polar oblique compressions are found. Spores compressed laterally
are bottle-shaped in outline, polar compressions circular to oval in outline. Originally
the spore body was more or less spherical in shape. Contact areas are usually distinct,
laevigate or ornamented with elements, similar in form to but smaller in size than those
which characterize the distal surface. Exine ornamentation is varied consisting of basic
elements e.g. verrucae, cones, spines, pilae, baculae, capillae ‘hair-like’ forms, or more
complex elements with features of two or more of the basic type, or the exine may be
laevigate. Often two layers of exine can be distinguished (transmitted light) the inner
layer when distinguishable being thin and often folded.

Comparison and remarks. Lagenicula is most similar to Setosisporites (Ibrahim). How-
ever, in Setosisporites the apical prominence is formed mainly by the expansion of the
laesurae, which generally occurs only in the area immediate to the proximal pole. Also,
the heightening of the laesurae is relatively abrupt in Setosisporites, not a gradual
increase towards the proximal pole as in Lagenicula.

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 443

Setispora Butterworth and Spinner 1967 may appear similar to Lagenicula, particularly
in ‘dry’ specimens. However, any indication of an apical prominence in Setispora is due
to sculptural elements on the laesurae.

Some specimens of Cystosporites Schopf 1938 may have an apical prominence similar
to Lagenicula (e.g. see Chaloner 1954), but the extremely long polar axis and mesh-like
structure of the sac-like spore body clearly distinguish the genus. Isolated abortive
specimens of Cytosporites may be confused with small laevigate specimens of Lagenicula,
e.g. L. nuda Nowak and Zerndt 1936.

Lagenoisporites Potonie and Kremp 1954 and Rostratispora Bharadwaj and Venka-
tachala 1962 are characterized by an apical prominence essentially similar to that of
Lagenicula. The former genus was separated by its authors from Lagenicula on the
presence of a more or less smooth or laevigate exine. Rostratispora was proposed on the
basis of a verrucose rather than "hair-like’ sculptural elements on the exine. Opinion
varies considerably on the use of differences in ornamentation to group otherwise similar
specimens at a generic level, but I doubt the value of further subdivision at this level
on the basis of differences in ornament. Many lageniculate forms are known with very
fine elements which are difficult to see on dry specimens. Others when examined by
transmitted light reveal elements which vary considerably in form and are difficult
to describe in terms of basic element types such as cones, verrucae, pila.

It is proposed that the close similarity in form of apical prominence and original
shape of body, suggests that the older name Lagenicula should be retained for the genus,
with Lagenoisporites and Rostratispora as junior synonyms; and that the ornament
differences are used at a specific level.

Botanical affinities. Lepidodendraceae; Potonie 1962.

Lagenicula subpi/osa (Ibrahim) forma major Dijkstra ex Chaloner 1954

Plate 84, figs. 1-4

1950 Triletes subpilosits forma major Dijkstra, p. 871 (nom. nud.).

1954 Triletes subpilosits forma major Chaloner, p. 27, pi. 1, figs. 4-8.

71957 Triletes subpilosus (Ibrahim) S.W. et B. forma major ; type 271/; Dijkstra, p. 14, pi. 9,
figs. 94-6, pi. 10, figs. 97-103.

71957 Triletes subpilosus (Wicher) S.W. et B. forma major Dijkstra; Dijkstra and Pierart,
pp. 12-13, pi. 11, figs. 126-7.

1959 Triletes subpilosus forma major (Dijkstra) ex Chaloner; Winslow, pp. 18-20, pi. 1,
figs. 1-9.

71962 Lagenicula subpilosa (Ibrahim) Potonie and Kremp; Ishchenko and Semenova, pars
p. 71, pi. 8, fig. 1 .

1967 Lagenicula subpilosa (Ibrahim) forma major Dijkstra ex Chaloner 1954; Butterworth
and Spinner, pp. 13-14, pi. 3, figs. 2-4.

71967 Lagenicula subpilosa (Ibrahim 1933) Potonie and Kremp 1955 pars', Karczewska, pp.
286-7, pi. 2, figs. 4-6.

Description. See Chaloner 1954, p. 27; Winslow 1959, pp. 18-20; Butterworth and Spinner
1967, pp. 13-14.

Remarks. The material assigned here to this forma agrees closely with an earlier de-
scription (Butterworth and Spinner 1967, pp. 18-20), but an additional feature noted

444

PALAEONTOLOGY, VOLUME 12

on several specimens is the presence of a thin, folded layer of exine within the ornamented
layer that is generally regarded as the spore wall.

Since there is some disagreement over the value of recognising this forma (Karczewska
1967, p. 286), and also in the variation in over-all size of spores, length of sculptural
elements etc., which appear to vary with stratigraphic horizon (Winslow 1959, pp. 1 8-20),
the following new results are recorded: Maximum diameter of the spore 960-1760 ft,
mean 1280 /x, based on 50 specimens mounted in a hydrous medium; One other specimen
measured 640 p in maximum diameter. The apical prominence ranged between 150 and
300 /x in height, 200-400 p in basal width (height measured on lateral compressions as
from beginning of projection from curved spore outline to apex). The spines on the
outer layer of exine, distal to the curvaturae, varied between 80 and 250 p in length,
16-40 ft in basal diameter, 4-10 ft in diameter at the apex or tip.

These figures are in approximate agreement with those previously recorded by Winslow
(1959, p. 18) from the Hardinsburg Formation, Kentucky, U.S.A. (low Chester Series,
Mississippi) and by Butterworth and Spinner (1967, p. 14) from the Bernician beds,
Cementstones, Cumberland, England (Visean S2). Dijkstra (1952, p. 103) in describing
this forma for the first time gave a size range of 500-1300 ft, mean 866 ft, based on 50
‘dry’ specimens.

The species Lagenicula subpilosa (Ibrahim) Potonie and Kremp 1955, which character-
izes strata of low Westphalian age, has a similar size range (320-1 100 ft, mean 652 ft,
Dijkstra 1946, p. 46 and 1952a, p. 103, ‘dry’ specimens; 550-1270 ft, Winslow 1959,
p. 17, wet specimens) to L. subpilosa forma major. This led Karczewska (1967, p. 286)
to reject the forma. However, as Winslow pointed out (p. 18) the distal spines on L.
subpilosa are generally less than 100 ft in length, and the maximum length of the spines
recorded by Karczewska is 115 ft. It is therefore suggested that on the basis of larger
mean diameter of spore, length of spines and different stratigraphic age, that the re-
tention on this forma is of value.

The relatively short dimensions of the spines, as given by Karczewska, makes the
inclusion of her record in the synonymy questionable. Also Dijkstra (1957, p. 14),
Dijkstra and Pierart (1957, p. 13) describe short elements 10 p long between the larger
distal spines. These were not reported by Winslow (1959) or by the author, either in.
1967 (Butterworth and Spinner) or during the present study.

Affinities. Lepidodendraceae; Potonie and Kremp 1954.

Stratigraphic distribution. Europe: Turkey, Namurian ABC (Dijkstra 1952); Ireland, Lower Carbonifer-
ous (Dijkstra 1957, Chaloner 1966); Scotland, Dinantian-Namurian (Chaloner 1954, ?Dijkstra 1957,
Sen 1964); England, Visean (Butterworth and Spinner 1967); ? Moscow Basin, Lower Carboniferous

EXPLANATION OF PLATE 84

All specimens by transmitted light.

Figs. 1-4. Lagenicula subpilosa (Ibrahim) forma major Dijkstra ex Chaloner 1954. 1, Proximal

surface, polar compression, X 100; slide D/10. 2, Lateral compression, X 50; slide D/ll. 3, Part

of specimen to illustrate cones on contact areas, x 100; slide D/12. 4. Part of specimen illustrating

spines projecting from distal surface, X 100; slide D/13.

Figs. 5-7. Setosisporites pseudoreticulatus sp. nov. 5. Proximal surface, polar compression, x 100;
slide D/7. 6, Holotype, oblique compression, x 100; slide D/8. 7, Lateral compression. X 100,

slide D/9.

Palaeontology , Vol. 12

PLATE 84

SPINNER, Scottish Visean megaspores

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 445

(Dijkstra and Pierart 1957); ? Poland, Visean (Karczewska 1967). U.S.A.: Illinois, Indiana, Kentucky,
Michigan, Mississippian (Chaloner 1954, Winslow 1959).

Genus setosisporites (Ibrahim) Potonie and Kremp 1954 emend.

Type species. Setosisporites hirsutus (Loose 1932) Ibrahim 1933.

Emended diagnosis. Trilete megaspores characterised by an apical prominence at the
proximal pole, formed mainly by the expansion of the laesurae at the pole, although
the polar part of the contact area may be involved. The greater parts of the contact
areas are not involved in the formation of the apical prominence whose size relative to
the spore body is such that the polar and equatorial axes are of similar dimensions.
There is no preferred type of compression; polar oblique compressions are circular to
oval in outline, lateral compressions bottle-shaped. The spore was originally more or
less spherical in shape. Distal to the apical prominence the laesurae are low and of
variable length. Contact areas are distinct, laevigate, or ornamented with elements
similar to those present on the distal surface, but smaller in size. Spore wall when
ornamented is covered with basic grana, cones, spines baculae, capillae or pilae type
elements of variable size and density.

Comparison. In their original diagnosis of this genus Potonie and Kremp (1954, p. 152)
referred to the exine ornamentation, except the contact areas, as being branched,
slightly pointed, short or also relatively long hairs. The emended diagnosis refers to
the original shape of the spore body, form of apical prominence and wide variation in
exine ornamentation. See also under emendation of Lagenicula above.

Botanical affinities. Porostrobus spp. (Chaloner 1958, Bharadwaj 1958); Bothrodend-
rostrobus watsoni (Chaloner 1967).

Setosisporites indianensis (Chaloner 1 954) comb. nov.

Plate 85, figs. 1-4

1954 Triletes indianensis Chaloner, p. 28, pi. 2 figs. 1-2.

1959 Triletes indianensis', Winslow, p. 26, pi. 6 figs. 1-3.

Size and shape. Trilete megaspores circular to oval in outline varying between 875 and
1615 p in max. diameter, mean 1195 p (based on 28 dry specimens). Polar, oblique and
lateral compressions are found. Lateral compressions are bottle-shaped due to the
projection of an apical prominence. However, the polar axis of the compressed spore
is not always larger than the equatorial axis. Originally, the spore body was more or
less spherical in shape.

Haptotypic features. An apical prominence is present at the proximal pole. Bluntly
pyramidal in shape, this structure is formed by the expansion of the contact areas
close to the proximal pole, and in part (?) by the laesurae. Basal diameter of apical prom-
inence varies 200-375 p, 50-150 p in height. The laesurae extend as low trilete ridges
from the apical prominence to the curvaturae, 30-60 p high and wide. On the apical
prominences of some specimens small ridges occur as (?) extensions of the laesurae.

446

PALAEONTOLOGY, VOLUME 12

Contact areas occupy approximately three-quarters of the proximal surface of the com-
pressed spore and ornamented with small grana approximately 6 p in diameter. The
position of the curvaturae are marked by low arcuate ridges which are only well developed
at the junction with the laesurae, 25-50 p wide, approximately 25 p high.

Exine structure and sculpture. The spore wall, distal to the contact faces is ornamented
with densely placed granulate to conate type elements 10-15 p up to 20 p in basal
diameter and height. These are generally more densely placed in the region of the curva-
turae. Internal to the granular layer, a thin membrane can be seen on broken specimens.
The outer granular layer of exine varies 30-50 p in thickness.

Comparisons and remarks. Chaloner 1954 and Winslow 1959 both assigned this species
to Triletes, sensu Schopf, Wilson, and Bentall 1944. An attempt to place this species
in one of the narrower genera proposed by Potonie and Kremp 1954, encounters
difficulties.

These authors proposed three genera of lageniculate megaspores i.e. Lagenicu/a
(Bennie and Kidston) emend., Lagenoisporites Potonie and Kremp and Setosisporites
(Ibrahim) emend. The primary feature used in distinguishing these genera being the
structure, form and size of the apical prominence relative to the spore body, and a
secondary feature being the type of (or lack of) ornament on the exine. In type of apical
prominence T. indianensis is more similar to Setosisporites e.g. S. praetextus (Zerndt)
Potonie and Kremp 1955, than Lagenicula e.g. L. horrida (Zerndt) Potonie and Kremp
1954. However, the exine ornamentation in Setosisporites is ‘hair-like’ e.g. S. hirsutus
(Loose) Ibrahim 1932. Comparing with Lagenoisporites T. indianensis is similar in the
small differences in length, between polar and equatorial axes, and in the small size of
sculptural elements on some specimens which give an approximately smooth appearance,
particularly on specimens not well preserved. The apical prominence in Lagenoisporites
is, however, formed by the expansion along most of the length of the laesurae and the
contact areas.

Rostratispora Bharadwaj and Venkatachala 1962 another lageniculate genus of
megaspore is similar to T. indianensis in the type of ornamentation (verrucose) but has an
apical prominence similar in form to Lagenicula and Lagenoisporites. Although lateral
compressions of Setispora Butterworth and Spinner 1967 appear lageniculate, the height
of laesurae is a result of capillate type ornamentation, and is not comparable with
T. indianensis. Triletes indianensis Chaloner is therefore recombined to the genus

EXPLANATION OF PLATE 85

All specimens by reflected light, X 40, unless stated otherwise.

Figs. 1-4. Setosisporites indianensis (Chaloner 1954) comb. nov. 1, Proximal surface, polar com-
pression, illustrating typical Setisosporites type apical prominence and large contact areas; slide
D/6. 2. Lateral-oblique compression; slide D/6. 3, 4, Part of specimen illustrating ornamenta-

tion at spore margin, X 120, by transmitted light; slide D/14.

Figs. 5-7, 10-12. Setosisporites splendidus (Zerndt 1937) comb. nov. 5, Proximal surface, polar
compression; slide D/5. 6, Distal surface, polar compression; slide D/5. 7, 10, Part of specimen

illustrating ornament at spore margin, xl20, by transmitted light; slide D/5. 11, 12, Part of

specimen illustrating types of ornament on distal surface, X 120, by transmitted light; slide D/15.

Fig. 8. Setosisporites sp. A, oblique compression; slide D/4.

Fig. 9. Setosisporites sp. B, lateral compression; slide D/3.

Palaeontology, Vol. 12

PLATE 85

SPINNER, Scottish Visean megaspores

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 447

Setosisporites as emended above in the characteristics of original shape of spore body,
form of apical prominence and type of ornamentation.

Stratigraphic distribution. North America, Beaver Bend Limestone, Indiana, Low Chester Series,
Mississippian (Chaloner 1954); Bethel Formation, Kentucky, Low Chester Series, Mississippian
(Winslow 1959).

Setosisporites pseudoreticulatus sp. nov.

Plate 84, figs. 5-7

71937 Type 13a, Triletes temiispinosus var. brevispinosa Zerndt pars, p. 6, pi. 3, figs. 1, 2, 5-7.
71957 Triletes hirsutus (Loose) var. brevispinosa Schopf, Wilson and Bentall forma I Dijkstra
pars, p. 13, pi. 7 figs. 67-71.

71959 Triletes g/obosus Arnold var. A Winslow, pi. 4, figs. 1-3.

Holotype. Plate 84, fig. 6.

Diagnosis. Trilete megaspores approximately circular to rounded triangular in outline
varying 420-800 p in maximum diameter. Apical prominence varies 60-120 p in height,
90-150 p in basal width, rounded at apex. Contact areas are ornamented with cones
4-6 p high. Distal to the contact areas, baculate to pilate elements, 10-30 p in length,
up to 6 p in basal diameter with small cone-like terminals, form an irregular reticulate
pattern of ornamentation. These elements are densely placed in the region of curvaturae
and may be partly fused to form a small flange-like structure 20-30 p in width.

Description: Size and shape. Trilete megaspores, approximately circular to rounded
triangular in outline, carrying between 420 and 800 p in maximum diameter, mean 620 p
(based on 50 specimens in a hydrous medium). Polar, oblique and lateral compressions
are all commonly found, there being no apparent preferred direction of compression.
On lateral compressions the curved outline of the spore body is interrupted by a small
‘neck-like’ protuberance from the proximal pole. Originally, the spore body was of
approximately spherical shape, the polar and equatorial axes being of similar dimensions.

Haptotypic features. Laesurae are straight to slightly sinuous in outline, approximately
two thirds the radius of the spore body in length. The line of commissure may be
ruptured, and is bounded by narrow (approximately 10 p wide) tecta raised 20 p high
near the junction with the curvaturae. The height of the laesurae increases gradually
20-40 p from the curvaturae towards the proximal pole, where there is a marked ex-
pansion forming an apical prominence. This structure varies 60-120 p in height (measured
in lateral compressions as projections from spore margin), 90-150 p in basal diameter
(measured perpendicular to height). The apex of the apical prominence is smoothly curved.

Contact areas are distinct occupying one-half to two-thirds of the proximal surface
of the compressed spore, and ornamented with small cones, approximately 4 p basal
width, up to 6 p high. On some specimens the contact areas have irregular areas of
thickened exine, which tend to be arranged radially from the proximal pole. The
curvaturae are represented by narrow thickenings on the exine.

Exine structure and sculpture. The spore wall excluding the contact areas, is covered
by what are basically baculate to pilate type elements. These vary 10-30 p in length,
up to 6 p in basal diameter, slightly swollen at the apex, where one or more small,
pointed, cone-like projections may occur. The elements may be curved or straight-sided

C 6685 G g

448

PALAEONTOLOGY, VOLUME 12

and are oftened joined at the bases. This feature, in addition to the elements being
adpressed to the spore exine, gives the appearance of an irregular interrupted reticulum.
On many specimens the elements are densely placed in the region of the curvaturae so
as to form a small flange-like structure 20-30 p wide.

Two layers of exine can easily be distinguished forming the spore wall. The outer
layer appears infrapunctate, approximately 20 p thick, measured in optical section,
yellow-brown in colour. The inner layer, distinguished under transmitted light, is
attached to the laesurae, thin and folded. In some specimens it is shrunken, but in others
it is in close proximity to the thick outer layer.

Under reflected light, the exine is yellow-brown in colour, the apical prominence is
not always clearly visible, and the sculptural elements appear more or less granular.
The pseudoreticulate nature of the ornament is not clearly distinguishable.

Comparisons and remarks. This type of spore illustrates some of the difficulties that arise,
when descriptions are given based on only one type of examination, i.e. reflected or
transmitted light.

Under reflected light examination S. pseudoreticulatus is similar in shape, size, and
ornamentation to S. hirsutus var. brevispinosa forma 1 (Zerndt) Potonie and Kremp 1955
as described by Zerndt (1937, p. 6) and Dijkstra (1957, p. 13). Zerndt noted that it was
difficult to distinguish a regular form of the sculptural elements present on his specimens,
but that he regarded the elements as small spines, up to 6 p in length, 4 p in basal width.
He also reported much smaller elements on the contact areas of his material. On most
specimens of S. pseudoreticulatus the elements are adpressed to the spore wall, and
appear under reflected light as small projections of the size given by Zerndt. However, both
Zerndt and Dijkstra emphasize the radial thickenings of the exine on the contact areas
as being the characteristic feature of S. hirsutus var. brevispinosa forma 1 . Although some
specimens of S. pseudoreticulatus were found with thickenings on the contact areas,
these were not common or as distinctive as suggested by Zerndt. It may be noted that
Dijkstra (1957) using reflected light found S. hirsutus var. brevispinosa forma 1 to be
common in the coals of the Limestone Coal Group of Scotland, slightly younger
(Ei) in age than the horizon studied here. Thus, there is evidence which suggests that
some of the material referred by earlier workers to S. hirsutus var. brevispinosa could
be assigned to S. pseudoreticulatus and this is indicated in the synonmy given above.
The major argument against such an assignment is the prominence of the radial thicken-
ings on the contact areas, and the size of ornament.

Under transmitted light many more of the details of morphology of S. pseudoreticulatus
are seen, and similarities can be seen with *S. globosus (Arnold) Potonie and Kremp 1955
in shape and general morphology. However, S. globosus has a smaller size range (390-
570 p Arnold 1950, p. 80; 385-640 p Winslow 1959, p. 42), and the sculptural elements
are larger (35-50 p length, up to 12 p diameter; Arnold p. 80), straight-sided with terminal
clefts. The sculptural elements are not joined to form a reticulate type pattern which
characterises S. pseudoreticulatus , but are discrete and widely spaced.

Winslow (1959, pp. 43-5) described three varieties of S. globosus. Of these varieties,
S. pseudoreticulatus is very similar to S. globosus var. A in length and height of laesurae,
size of apical prominence, type and size of sculptural elements forming an irregular
reticulate pattern (catenulate ornament of Winslow, p. 43). The size range of var. A is

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 449

smaller (330-630 p, mean 515 p; Winslow, p. 43) than S. pseudoreticulatus (420-800 p,
mean 620 p). Winslow (p. 43) also noted the similarities of var. A with S. hirsutus var.
brevispinosa when compared under reflected light examination, but did not consider the
two as being the same due to the absence of radial thickenings on the contact faces of
variety A.

The differences in ornamentation between S. globosus and S. pseudoreticulatus (in-
cluding Winslow’s var. A, see synonymy) are considered here to warrant separation at
specific level. No specimens with the type of ornamentation present on S. globosus
were found associated with S. pseudoreticulatus during the present study.

Winslow’s other two varieties of S. globosus can be distinguished from S. pseudoreticu-
latus by the larger size of the sculptural elements (15-57 p, generally more than 35 p,
var. B) and the type of element (tubercles 5-26 p length, 5-52 p basal diameter, var. C).

S. pseudoreticulatus also resembles S. reticulatus Karczewska 1967 in size, shape, and
ornament pattern. However, on S. reticulatus the outer layer of exine forms a uniform
fused reticulum on the distal surface. Small projections 3-7 p in length, are situated
between the lumina of the reticulum.

S. pseudotenuispinosus Pierart 1958 and S. pilatus Spinner 1965 are similar in shape
and size of the spore body to S. pseudoreticulatus , but these species have smaller,
discrete, sculptural elements which do not form a reticulate type pattern.

Triletes catenulatus Winslow 1962 can be distinguished from S. pseudoreticulatus
by the presence of highly developed laesurae (200 p or more high).

In my opinion many of the specimens previously assigned to S. hirsutus var. brevi-
spinosa forma 1 could be placed in S. pseudoreticulatus. However, rather than raise the
varietal name to species rank, a new name is proposed for the following reasons. Zerndt
(1937) recognised two forms (varieties 1,2; Zerndt 1937, p. 7) within var. brevispinosa.
These forms are markedly different in type and size of ornamentation (form 1 , appendices
up to 6 p long; form 2, spines 39-96 p wide at the base, decreasing to approximately
12 p near the tip). However, Zerndt emphasized the folding, thickening on the contact
areas as the common feature of both forms grouped within the named variety brevi-
spinosa. Later Schopf, Wilson, and Bentall (1944, p. 26) in dealing with the problem
of Zerndt’s varieties within a variety regarded variety 1 of Zerndt as being var. brevispinosa
and proposed a new varietal name secundus for variety 2 of Zerndt; whereas Dijkstra
(1946, 1956) interpreted varieties 1, 2 of Zerndt as forms within var. brevispinosa.
Due to the different interpretation of Zerndt’s work, the emphasis on folding and thick-
ening in his description of the variety, and the small size of the appendices on his form 1,
a new name pseudoreticulatus is proposed for the species.

Botanical affinities. Unknown.

Stratigraphic distribution. ?Limestone Coal Group, Scotland, Namurian E2 (Dijkstra 1957); Lower
Carboniferous, Ireland (Chaloner 1966); ? Hardinsburg Formation, Kentucky, Wattersburg Forma-
tion, Illinois, Chester series, Low Mississippian (Winslow 1959).

Setosisporites splendidus (Zerndt) comb. nov.

Plate 85, figs. 5-8, 11-12

1937 Type 28; Lagenicula splendida Zerndt, p. 13-14, pi. 18, figs. 1, 2; pi. 19, figs. 1-3, 5;
pi. 20, figs. 2, 3, 4.

450

PALAEONTOLOGY, VOLUME 12

1944 Triletes splendida (Zerndt) Schopf, Wilson, and Bentall, p. 25.

1946 Triletes splendidus (Zerndt) Dijkstra, p. 50, pi. 16, figs. 173-5.

1955 Lagenicula splendida Zerndt; Potonie and Kremp, p. 119.

1957 Triletes splendidus (Zerndt) S.W. et B.; Dijkstra, p. 14, pi. 8, figs. 85; pi. 9, figs. 86-8.

1959 Triletes splendidus (Zerndt) Schopf, Wilson, and Bentall; Winslow, p. 27, pi. 5, fig. 8.

71962 Lagenicula splendida Zerndt; Ishchenko and Semenova, pp. 72-3, pi. 8.

71962 Lagenicula verrucosa Ishchenko and Semenova, p. 74, pi. 9, fig. 3.

Description: Size and shape. Trilete megaspores more or less circular in equatorial
outline. Lateral compressions are most commonly found which have a marked ‘bottle-
shaped’ outline, the ‘neck’ being formed by an apical prominence at the proximal pole
of spore. The difference between the polar and equatorial axes is not large, and on some
lateral compressed specimens the polar axis is the smaller. Polar and oblique com-
pressions also occur. Maximum diameter of the spore body including apical prominence
on lateral compressions varies 935-1625 p, mean 1225 p based on 35 dry specimens.
Originally, the spore body was more or less spherical in shape.

Hapotypic features. The characteristic feature here is the bluntly pyramidal apical
prominence at the proximal pole, varying 200-300 p in basal diameter, 150-270 p
high on lateral compressions, and may appear slightly constricted at the base. This
structure is probably formed by an expansion of the exine of the immediate polar part
of the contact areas, and in part by the laesurae. On some specimens, small ridge-like?
continuations of the laesurae can be traced on to the prominence. In some specimens
the laesurae are ruptured, but the ruptures have not been observed extending on to the
prominence, whilst in other damaged specimens the prominence has been completely
removed. Distal to the apical prominence, the laesurae form low ridges 250-400 p long.

The contact areas occupy approximately three-quarters of the proximal half of the
compressed spore. These are delimited from the remainder of the spore body by the
smaller cone-like sculptural elements, generally less than 12 p in diameter, but occasion-
ally up to 30 p on some specimens. Arcuate ridges representing the curvaturae are not
well developed except at the junction with the laesurae i.e. curvaturae imperfectae.

Exine structure and sculpture. In the region of, and distal to the curvaturae the spore
wall is ornamented with cones and granular type elements. These are most densely
placed in the region of the curvaturae. Two groups of elements can be recognized based
on size. The most prominent group consists of relatively large cones 30-75 p in basal
diameter (generally larger than 50 p), 12-35 p high, and often mammillate in form.
Interspersed with these elements are smaller cone and granular type forms generally
20 p or less in basal diameter, 5-15 p high.

Broken specimens reveal a thin layer of exine, internal to the heavily ornamented layer
which is approximately 50 p thick.

Comparisons and remarks. The limits of this species are ill defined. Both Zerndt (1937,
pp. 13-14, pis. 18-20) and Dijkstra (1946, p. 50, pi. 16, 1957, p. 14, pi. 8) allow consider-
able range in spore diameter (700—1700 ^ ; Dijkstra 1957) and in size of sculptural elements
(16-97 p diameter, Zerndt 1937; 10-150 p diameter, Dijkstra 1957). At the same time
both authors recognised other species e.g. Lagenicula subtilinodulata Nowak and Zerndt
1936, which could be placed within the limits they described for L. splendida. Chaloner
(1954, p. 28) proposed Triletes indianensis for specimens having a distal ornamentation

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 451

of elements of up to, but generally less than, 20 g in length. These specimens could, on
the basis of the limits given by Zerndt and Di jkstra, be assigned to L. splendidci. Winslow
(1959, pp. 26-7) recognizes S. splendidus (Triletes splendida of Winslow) as being larger
in size of spore and ornamentation, than L. subtilinodulata and Trdeles indianensis.
Reference to the holotype is of no assistance for, as far as I am aware, neither Zerndt
nor any other worker has designated a holotype for this species.

I do not think that any particular value should be placed on relatively small differences
in the size of the spore body, but suggest that a useful character for distinguishing this
species is the two types of sculptural element. This character is seen on Zerndt’s speci-
mens illustrating the species, and plate 18, fig. 2 is designated here as the lectotype of the
species.

Triletes indianensis Chaloner 1954 differs from S. splendidus in the small size (generally
less than 20 g in basal diameter and height) of the elements forming the ornamentation
on the spore wall.

Lagenicula agnina Zerndt 1937 can be distinguished from S. splendidus by the type
of apical prominence (laesurae expanding along most of the length), small contact
areas, and close verrucose ornament on the distal surface.

Lagenicula verrucosa Ishchenko and Semenova 1962 can according to its authors
be distinguished from S. splendidus by its smaller size (1360x 1325 g) and ‘barbate’
sculpture. Judging from the drawings produced as illustrations of the two species,
L. verrucosa appears more similar to the specimens described here than does L. splen-
dida sensu Ishchenko and Semenova and this is indicated in the synonymy given above.

There is some reason to doubt the correctness of assigning this species to Lagenicula
as described by Potonie and Kremp (1955, p. 118). In their diagnosis these authors
maintain: ‘a long polar axis, an apical prominence larger than the part of the contact
areas not involved in its formation, and unornamented contact areas.’ These features
are not applicable to this species. The type and size of the apical prominence relative to
the other haptotypic features is more typical of Setosisporites e.g. S. praetextus (Zerndt)
Potonie and Kremp 1955. Consequently this species is assigned to Setosisporites as
emended above.

Stratigraphic distribution. Europe: Poland, Dinantian-Namurian A (Zerndt 1937); U.S.S.R. Donetz
Basin, ?Dinantian-Namurian, (Ishchenko and Semenova 1962); Ireland, Lower Carboniferous,
(Dijkstra 1957, Chaloner 1966). North America: Degonia Formation, Chester Series, Illinois,
Mississippian (Winslow 1959).

Setosisporites sp. A
Plate 85, fig. 9

Description. Trilete megaspores circular to oval in outline, approximately 1250 g
in maximum diameter (only two specimens found, 1310 and 1180 ft in maximum dia-
meter). Apical prominence at proximal pole, blunt pyramidal shaped, 300 g maximum
diameter. Laesurae relatively low, 25-60 g wide at junction with curvaturae. Contact
areas approximately 650 g in radius ornamented small cones 25 g diameter. Spore wall
distal to contact faces, ornamented with large cones 65-80 g basal diameter, 60-70 g
high, smaller cones approximately 40 g diameter in region of curvaturae.

Remarks. These megaspores closely resemble Setosisporites splendidus as described

452

PALAEONTOLOGY, VOLUME 12

here, but have more typical conate ornamentation in which the elements are higher,
approximately equal to basal diameter, and not mamillate in form as in S. splendidus.
The elements are also not so densely placed on the contact areas or distal surface.
Lagenicula agnina Zerndt 1937 has larger sculptural elements on the distal surface
(97-225 p width, 48-160 p high; Zerndt 1937, p. 14). L. subtilinodulata Nowak and
Zerndt 1936 is smaller in size of spore body and distal ornamentation, and the contact
areas are laevigate.

Setosisporites sp. B
Plate 85, fig. 10

Description. Trilete megaspores 912-1500 p maximum diameter, mean 1300 p based on
six dry specimens. Apical prominence is bluntly pyramidal, 150-225 p high, 250-375 p
basal diameter. Laesurae, distal to apical prominence, are approximately 50-60 p
high and wide, best developed near the junction with the curvaturae. Contact areas
occupy three-quarters proximal surface, smooth bounded by arcuate ridges 35-50 p
high, 50-80 p wide, most pronounced at junction with the laesurae. Distal surface
ornamented with irregularly scattered cones, 25-40 p high, and wide, 60-300 p apart.

Remarks. In type of apical prominence this species resembles S. splendidus and S.
indianensis.

Genus zonalesporites (Ibrahim) Potonie and Kremp emend. Spinner 1965
Type species. Zonalesporites brasserti (Stach and Zerndt) Potonie and Kremp 1956.

Zonalesporites fusinatus sp. nov.

Plate 86, figs. 1-4

71957 Triletes brasserti Stach and Zerndt, type 20; Dijkstra pars , p. 13, pi. 7, figs. 73-6.
Holotype. Plate 86, fig. 1.

Diagnosis. Trilete megaspores composed of spore body with a subequatorial corona.
Spore body varies 1300-2160 p in maximum diameter. Corona varies 415-700 p width,
composed of a number of layers of baculate type elements, 10-30 p width, with rounded
tips. Elements are more fused distally to form a rim with some dissections and ornamented
with small baculate processes. Margin of rim generally smooth, lacking ornamentation.

EXPLANATION OF PLATE 86

All specimens by transmitted light, x40, unless stated otherwise.

Figs. 1-4. Zonalesporites fusinatus sp. nov. 1, Holotype, proximal surface, polar compression, by
reflected light; slide D/1. 2, Distal surface, polar compression, by reflected light; slide D/2. 3, 4,

Parts of corona, X 120; slide D/17.

Figs. 5-8. Parts of corona of Zonalesporites spp. for comparison with Z. fusinatus. 5, 8, Z. brasserti
(Stach and Zerndt) Potonie and Kremp, X 120; slides WD/6, WD/8 (Westphalian B, South Wales
Coalfield, Great Britain). 6, Z. conacies Butterworth and Spinner, x 100; slide V4/21 (Dinantian,
S., age, Bernician Beds, Whitberry burn, Cumberland, England). 7, Z. cf. ramosus (Arnold) Spinner,
x 250 (early Pennsylvanian, Williamston Spore coal, Michigan, U.S.A.; specimen lent by W. G.
Chaloner, Botany Department, University College, London).

Palaeontology, Vol. 12

PLATE 86

SPINNER, Scottish Visean megaspores

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 453

Description: Size and shape. Trilete megaspores approximately circular to rounded,
triangular in outline consisting of a spore body with an equatorial corona. The spore
body varies between 1300 and 2160 p in maximum diameter, mean 1440 p (based on
36 ‘wet’ specimens). Entire specimens are not common, 10 such specimens measured
1550-2300 p in maximum diameter, mean 1950 p. Polar compressions are most common,
due to the longer equatorial axis of the entire spore. The spore body was originally
more or less spheroidal in shape.

Haptotypie features. Laesurae are straight or sinuous in outline, half to three-quarters
the spore body radius in length. Broadly based, the laesurae are raised 100-250 p at the
proximal pole. On most specimens the laesurae decrease slightly in height from the
proximal pole before attaining the greatest height at the position of the curvaturae,
150-290 p. Also on some denuded specimens the laesurae are seen projecting beyond the
spore body margin. An apical prominence is not present. Contact areas are laevigate
and occupy half to three-quarters of the proximal surface of the compressed spore body.
The positions of the curvaturae are marked by the bases of the elements forming the
corona.

Exine structure and sculpture. The corona is attached in a region slightly proximal to
the geometrical equator of the compressed spore body. The width of the corona varies
415-750 p, greatest width in line with the laesurae, hence the subtriangular outline of
the entire spore. The overlap of the corona onto the proximal surface of the spore body
approximately equals one third of the width of the corona. The corona is formed by
several layers of elements (five or more). These are approximately 30 p wide at the base,
densely spaced, and partly fixed to form a loose type of reticulum. The proximal elements
are most distinct 10-25 p width, baculate in form, with smoothly rounded tips. Distally
the layers are more completely fused and form an almost continuous rim to the corona.
Small baculate elements with rounded tips 10-30 p in width project from the fused part
of the corona, but only rarely from the rim (small dissections occur in the corona
approximately 100 p in diameter). On some small specimens the fused outer part of the
corona is not so clearly distinguishable and the rim has a crenulate appearance.

Distal to the corona the spore body is generally laevigate, but small verrucate type
elements 20-50 p in diameter may occur scattered on the distal surface. These tend to
become more elongate near the spore body margin and merge into the corona. The spore
body wall appears infrapunctate, approximately 40 p thick, as measured in optical
section. Within this layer a further thin folded layer of exine can be distinguished (trans-
mitted light) in most specimens.

Under reflected light the corona appears almost completely fused, with some striations,
lighter in colour than the spore body. Dissections can be distinguished.

Comparison and remarks. Under reflected light both Zonalesporites brasserti (Stach and
Zerndt) Potonie and Kremp 1956 and Z. fusinatus sp. nov. appear very similar, and the
differences in corona structure are not easily seen. Dijkstra (1952, 56, 57) recognizes wide
variation in size of spores and some variation in the corona of this species, but does not
consider these differences to warrant distinction at species level, but he distinguishes
several formae within the species. Potonie and Kremp (1956, p. 122) give no details of the
structure of the corona (cingulum of P. and K.) in their description. However, Winslow

454

PALAEONTOLOGY, VOLUME 12

(1959) using transmitted light, restricted Z. brasserti to those forms with a corona
(flange of Winslow) formed by ?layers of elements, fused laterally except at the point
of attachment to spore body and at the terminals. The outer layer forms ‘bar-like’ processes
(Winslow, p. 36) projecting at the margin. Only rarely are small dissections seen in the
corona.

I agree with this interpretation of Z. brasserti and separate the specimens described
here into a new species Z. fusinatus. The two species are similar in general appearance,
especially under reflected light. However, in Z. fusinatus the corona is less compact,
the elements are baculate processes with smoothly rounded tips, are only partly fused,
and the rim of the corona is generally smooth, lacking any projections. The laesurae
and contact areas are usually ornamented in Z. brasserti with verrucate type processes,
generally laevigate in Z. fusinatus.

Z. conacies Butterworth and Spinner 1967 can be distinguished from Z. fusinatus
by the elongate cones which characterize the corona (PI. 86, fig. 5). Z. radiatus (Zerndt)
Spinner 1965 lacks the fused margin of the corona, and the elements are spaced on the
spore body. Z. fusinatus differs from Z. rotatus (Bartlett) Spinner 1965 by the larger
spore body and the corona formed by several layers of elements. Z. ramosus (Arnold)
Spinner 1965 although having a corona formed by several layers of elements as in
Z. fusinatus, characteristically has relatively large projections (up to 52 p long, Winslow
1959, p. 33) from the rim of the corona (PI. 86, fig. 7).

The comparisons made above are based on transmitted light studies. Under reflected
light examination it is difficult to distinguish Z. brasserti from Z. conacies and Z.
fusinatus. However, the author considers the differences (see PI. 86, figs. 3-8) to be
distinctive and worthy of recognition at a specific level.

Affinities. ?Lycopsida (Potonie and Kremp 1954).

Stratigraphic distribution. ?Namurian, Scotland (Dijkstra 1957).

THE MEGASPORE ASSEMBLAGES

Both coal seams yielded well preserved megaspores of considerable variety. A count
of 500 specimens from each seam indicated the composition of the assemblage as follows :

Coal samples

Skateraw

Longcraig

Lagenicula subpilosa forma major

176

132

Setosisporites pseudoreticulatus

145

101

Setosisporites splendidus

53

30

Setosisporites indianensis

0

22

Setosisporites sp.

3

11

Cystosporites sp.

11

10

Zonalesporites fusinatus

34

194

Zonalesporites rotatus

78

0

At generic and specific level the two assemblages are closely similar in over-all
content and proportional representation. However, the differences may be of significance
in correlation with other areas in northern Britain, in that the Longcraig and Skateraw
coals are stratigraphically almost immediately below and above the upper Longcraig

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 455

limestone; these are the records of S. indianensis and Z. rotatus, and the dominance of
Z. fusinatus in the Longcraig coal. Accepting the suggested correlation of this limestone
with the Hurlet limestone of the Midland valley of Scotland by Wilson, the differences
between these assemblages may contribute further to the subdivision and correlation of
Lower Carboniferous deposits in the Midland valley of Scotland and northern England.

Although no plant megafossils have been recorded from the area, some indication
of the types of plant forming the forests at this time can be obtained from the megaspore
assemblages. Chaloner (1954, p. 31) and Winslow (1959, p. 94) commented on the im-
portance of the Lepidodendrid-Lepidocarpon elements in the forests during the Missis-
sippi (Chestererian) of central U.S.A. The same comment may be made here, based
on the occurrence of Lagenicula and Cystosporites. Probably a more important element
was the plant producing the Zonalesporites. This type of spore is known from Sporangio-
strobus Bode (Chaloner 1962). The plant bearing Sporangiostrobus is unknown, but
the large size and structure of the cones suggests a large arborescent form. Nemejc
(1946, p. 7) suggested that the most probable parent plants of Sporangiostrobus are
among the lycopods with long leaves, such as Ulodendron Lindley and Hutton. The
recorded species of Setosisporites are unknown from fructifications, although spores
assignable to the type species of this megaspore genus, S. hirsutus, are known from the
small ?herbaceous lycopod Porostrobus canonbiensis Chaloner (Chaloner 1962).

From the known broad association of Zonalesporites and Setosisporites with the
Cristatisporites/ Densosporites group of microspores (Chaloner 1962, pp. 82-3) it may
be suggested that the Zonalesporites and Setosisporites of the assemblages recorded
here represent elements of the open moor of Kremp (1952) or the densospore phase of
Smith (1957). Similarly, Lagenicula subpilosa forma major and Cystosporites sp. rep-
resent the forest moor or lycospore phase.

The absence of the megaspore genera Tuberculatisporites (Ibrahim) Spinner 1968,
Laevigatisporites (Ibrahim) Potonie and Kremp 1954, or similar forms, suggests an
absence of Sigillarians in the lycopod flora. A similar comment may be made regarding
the Selaginellales since there is an absence of any spores of Triangulatisporites Potonie
and Kremp 1954. In general, the megaspore assemblages indicate a flora in which
large arborescent lycopods with long leaves were important together with some more
diminutive forms.

COMPARISONS WITH RECORDS FROM OTHER AREAS

The relatively little published research on Lower Carboniferous has been largely
concerned with records of occurrence, discussion on the value of broad or narrow
species definitions, and the relationship of megaspores to fructifications. Consequently,
it is not yet possible to reach firm conclusions on the stratigraphic value of these mega-
spores.

In one of the earliest papers dealing with dispersed megaspores (Bennie and Kidston
1886), three forms were described and recorded from localities in the ‘Carboniferous
Limestone Series’ of Scotland: Triletes IV, Lagenicula I, and Lagenicula II of Bennie
and Kidston are very similar to S. pseudoreticulatus, L. subpilosa forma major and
Cystosporites sp ( Igiganteus ) recorded here. Much later, Dijkstra (1956) recorded the
same forms as well as Triletes rotatus, T. splendidus, T. brasserti (a form very similar

456

PALAEONTOLOGY, VOLUME 12

to Z. fusinatus ) from the Limestone Coal Group, which overlies the Lower Limestone
Group in Scotland. Dijkstra’s records were confirmed by Sen (1964) who studied the
Limestone Coal Group in Ayrshire, west Scotland. Also, Chaloner (1968, p. 79)
recorded megaspores closely similar to those reported by Dijkstra from part of the
Ballycastle coalfield. Northern Ireland. The present work extends the stratigraphic
record of these species into the Lower Limestone Group of the Carboniferous Limestone
Series and records for the first time the presence of S. indianensis outside North America.
In addition, Dijkstra and Sen reported Triletes praetextus Zerndt forma minor Dijkstra
1952, T. simplex (Zerndt) Schopf, Wilson, and Bentall 1944, T. mucronatus Nowak and
Zerndt 1936, T. horridus (Zerndt) Schopf, Wilson, and Bentall 1944, T. subsimplex
Dijkstra 1957, T. hirsutoides Dijkstra 1957 in the Limestone Coal Group. None of these
species have been recorded from either the Lower Limestone Group or older deposits
in Scotland.

Butterworth and Spinner 1967 described megaspores from coals and shales of the
Cementstones and Lewis Burn (Scremerston) Coal Group of Cumberland, north-west
England, which are C2 to S2 in age (coral-brachiopod zonation, Garwood 1931) and
equivalent to part of the Calciferous Sandstone of Scotland. The megaspores obtained
were not so abundant, nor were the assemblages as varied ( Butterworth and Spinner
1967, p. 21, table 1 ) as those obtained here. Of the seven species of megaspores recorded,
only one, L. subpilosa forma major has also been recorded here.

Summarizing, it appears that the megaspore assemblages from the Dunbar area
(upper Px to basal P2 in age) are different in content from those reported from the under-
lying Calciferous Sandstone formations (C2-S2 age) below and the Limestone Coal
Group above (Ei in age) in the Midland Valley of Scotland and northern England
respectively.

Studies of megaspores from Lower Carboniferous deposits in areas outside Great
Britain are also few in number. Chaloner 1954 recorded L. subpilosa forma major, and
S. indianensis from the Beaver Band limestone (low Chesterian), Indiana, U.S.A.
Two species, Setispora echinoides (Chaloner) Butterworth and Spinner 1967, S. palaeo-
cristata (Chaloner) Butterworth and Spinner 1967 were also first described by Chaloner
from this formation. Although these species were not found during the present study,
the only other record of this genus is of three similar species, S. pseudoreticulata Butter-
worth and Spinner 1967, Triletes pannosus Alvin 1966, T. subpalaeocristatus Alvin
1965, all reported from the underlying Calciferous Sandstone Series of this region.

Winslow (1959) published on the distribution of megaspores in the Mississippian
(Chester series; Winslow, p. 74) deposits of central U.S.A. In this work Winslow com-
mented (p. 77) that although megaspores were often abundant, there was little variety
within an assemblage. Most of the species recorded during the present study were also
reported by Winslow, a notable exception being the Zonalesporites. Winslow reported
(p. 80) this type of spore appearing for the first time in the Pennsylvanian (Caseyville
Group).

A markedly different impression of the variety of Lower Carboniferous megaspores
is given by workers in continental Europe. Dijkstra 1956 described twelve new species
from samples obtained from Egypt considered by Dijkstra to be older than the Lime-
stone Coal Group of Scotland, and later (1957) with Pierart proposed 29 new taxa
from the Moscow Basin.

E. SPINNER: MEGASPORE ASSEMBLAGES FROM DUNBAR, EAST LOTHIAN 457

From our present knowledge it appears that the megaspore assemblages described
here are more comparable with those of approximately equivalent age in North America
than in continental Europe and Asia. Further work may produce more evidence for
Sullivan’s (1965, 67) suggestion of regional differentiation of Mississippian floras, and
may answer the question as to whether the similarities and differences noted above
have stratigraphic value.

Acknowledgements. I am indebted to Dr. W. G. Chaloner for the critical reading of the systematic
description of spores and to Dr. R. H. Wagner for reading the manuscript.

REFERENCES

alvin, k. l. 1965. A new fertile lycopod from the Lower Carboniferous of Scotland. Palaeontology,
8, 281-93.

1966. Two cristate megaspores from the Lower Carboniferous of Scotland. Ibid. 9, 488-91.

Arnold, c. a. 1950. Megaspores from the Michigan Coal Basin. Contr. Mas. Palaeont. Univ. Mich.,
8, 59-111, 18 pi.

bennie, j. and kidston, r. 1886. On the occurrence of spores in the Carboniferous formation of Scotland.
Proc. R. Phys. Soc. Edinb. 9, 82-1 17, pi. 3-6.

bharadwaj, d. c. 1959. On Porostrobus zeilleri Nathorst and its spores with remarks on the systematic
position of P. bennholdi Bode and the phylogeny of Densosporites Berry. Palaeobotanist, 7, (1958),
67-76.

and venkatachala, b. s. 1962. Spore assemblage out of a lower Carboniferous shale from Spits-
bergen. Ibid. 10, 18-47, pi. 1-10.

bode, h. 1928. Uber eine merkwiirdige Pteridophyten — fruktifikation aus dem oberschlesischen Carbon.
Jb. Preuss. Geol. Landesanst. 49, 245-7.

butterworth, m. a. and spinner, e. 1967. Lower Carboniferous spores from North-west England.
Palaeontology, 10, 1-24, pi. 1-5.

chaloner, w. g. 1953. On the megaspores of four species of Lepidostrobus. Ann. Bot. London (N.S.),
17, 262-93.

1954. Mississippian megaspores from Michigan and adjacent states. Contr. Mus. Palaeont. Univ.

Mich. 12, 23-35, 2 pi.

1958. A Carboniferous Selagineltites with Densosporites microspores Palaeontology, 1, 245-53,

pi. 44.

1962. A Sporangiostrobus with Densosporites microspores. Palaeontology, 5, 73-85, pi. 10-11.

1966. in wilson and robbie Geology of the country around Ballycastle. Mem. Geol. Snrv. Great

Britain, 78-81.

1967. in boureau Traite de Paleobotanique, Paris, 2, 656-7.

currie, e. d. 1954. Scottish Carboniferous Goniatites. Trans. R. Soc. Edin. 62, 527-99, 4 pi.
dlikstra, s. j. 1946. Eine monographische Bearbeitung der Karbonischen Megasporen. Meded. Geol.
Stichting, ser. c— 1 1 1—1, 1, 1-101, 16 pi.

1950. Carboniferous Megaspores in Tertiary and Quaternary Deposits of S.E. England. Ann.

Mag. nat. Hist. 3, 865-77.

— 1952a. New Carboniferous megaspores from Turkey. Ann. Mag. nat. Hist. 12, 102-4, 2 pi.

1952b. Megaspores of the Turkish Carboniferous and their stratigraphical value. Rept. Int. Geol.

Congr. 18 th sess. Great Britain, 1948, 10, 11-17.

1952c. The stratigraphical value of megaspores. C. R. Congr. A r. Etud. Strat. Carb. Heerlen,

1951, 1, 163-8, pi. 5-7.

1956. Megaspores carboniferas de la Camocha (Gijon). Estud. geol. Inst. Mallada, 12, 245-55

(English summary 256-62) pi. 48-57.

— 1957. Lower Carboniferous Megaspores. Meded. geol. Stichting (n.s.), 10, 1956, 5-18.

— and pierart, p. 1957. Lower Carboniferous Megaspores from the Moscow Basin. Meded. geol.
Stichting, 11, 5-19.

458

PALAEONTOLOGY, VOLUME 12

ishchenko, a. m. and semenova, e. v. 1962. The Megaspores of the Carboniferous ages and their
stratigraphical importance. Academy of Sciences Ukrainian S.S.R. Proceedings of the Institute of
Geological Sciences Stratigraphy and Palaeontology series, 43, Kiev. 147 pp. (in Russian).
karczewska, j. 1967. Carboniferous Spores from the Chetm 1 Boring (Eastern Poland). Acta Palaeont-
ologica Polonica Warsaw, 12, 268-345, 12 pi.

kremp, g. 1952. Sporen-Vergesellschaftungen und Mikrofaunen-Horizonte im Ruhrkarbon. C.P.

Congr. Aw Etud. Strat. Carb. Heerlen (1951), 1, 347-57.
lumsden, g. I. (in press). Excursion A4, B4, The Midland Valley of Scotland. C.R. Congr. Aw Etud.
Strat. Carb. Sheffield, 1967.

nemejc, j. 1946. Further critical remarks on Sternberg's Lepidodendron dichotonmm and its relations
to the cone of Sporangiostrobus Bode. Bull. inst. Acad. Tscheque, 47, 35-45.
nowak, j. and zerndt, j. 1936. Zur Tektonik des ostlichsten Teils des Polnischen Steinkohlenbeckens.
Bull. Acad. Pol. Sci. Letts. A, 56-73.

pierart, p. 1958. Palynologie et Stratigraphie de la zone de Neeroeteren (Westphalian C superieur)
en Campine beige. Publ. Ass. Wtud. Paleont. Bruxelles, 30, 23-102, pi. 1-18.
potonie, r. 1962. Synopsis der Sporae in situ. Geol. Jb., 52, 204 pp.

and kremp, g. 1954. Die Gattungen der palaozoischen Sporae dispersae und ihre Stratigraphie.

Geol. Jb. 69, 111-94, pi. 4-20.

1955. Die Sporae dispersae des Ruhrkarbons, ihre Morphographie und Stratigraphie mit

ausblicken auf Arten anderer Gebiete und Zeitabschnitte, Teil 1. Palaeontographica , 98B, 1-136,
pi. 1-16.

sen, j. 1964. The megaspores of the Ayrshire coalfield and their stratigraphic value. Micropaleontology ,
10, 97-104.

schopf, j. m., wilson, l. r. and bentall, r. 1944. An annotated synopsis of Palaeozoic fossil spores and
the definition of Generic groups. Rep. Invest. III. geol. Surv. 91, 1-73, 3 pi.
smith, a. h. v. 1962. The Palaeoecology of Carboniferous Peats based on the Miospores and Petro-
graphy of Bituminous Coals. Proc. Yorks. Geol. Soc., 33, 423-74.
spinner, e. 1965. Westphalian D megaspores from the Forest of Dean Coalfield, England. Palaeont-
ology, 8, pp. 82-106, pi. 14-17.

1968. Contribution on the megaspore genus Tuberculatisporites (Ibrahim) Potonie and Kremp

1954. Pollen et Spores, 10, 395-410.

Sullivan, h. j. 1965. Palynological evidence concerning the regional differentiation of Upper Mississip-
pi floras. Ibid. 7, pp. 539-63.

1967. Regional differences in Mississippian spore assemblages. Rev. Palaeont. Palynol. 1, 185-92.

winslow, m. 1959. Upper Mississippian and Pennsylvanian megaspores and other plant microfossils
from Illinois. Bull. III. geol. Surv. 86, 7-102, 16 pi.

1962. Plant Spores and other Microfossils from Upper Devonian and Lower Mississippian rocks

of Ohio. Prof. Pap. U.S. geol. Surv. 364, 93 pp.

zerndt, j. 1937. Les Megaspores du Bassin Houiller Polonais, partie 2. Acad. Pol. Sci. Lett. Traw
Geol. 3, 1-78.

EDWIN SPINNER
Department of Geology
The University
Sheffield 1

Typescript received 15 January 1969

CRUSTACEAN BURROWS IN THE WEALD CLAY
(LOWER CRETACEOUS) OF SOUTH-EASTERN
ENGLAND AND THEIR ENVIRONMENTAL

SIGNIFICANCE

by W. J. KENNEDY and J. D. S. MACDOUGALL

Abstract. The trace fossil usually known as Ophiomorpha nodosa Lundgren occurs at several horizons and
localities in the Weald Clay (Lower Cretaceous) of Surrey and Sussex. By analogy with Recent occurrences
(as described by Weimer and Hoyt 1962), all of which are marine, these structures are ascribed to the burrowing
activities of decapod crustaceans (callianassids) in near-shore and littoral sands. Normally the traces are associ-
ated with a mixed assemblage of marine and brackish water fossils, and it is inferred that the Weald Clay
occurrences of Ophiomorpha by itself indicates a marine environment, close to shore, where such an assemblage
might occur.

Three variants of Ophiomorpha are recognizable in the Weald Clay sandstones: (1) a pellet-lined system;
(2) burrows with a meniscus fill; (3) unlined burrows with scratch marks. Each represents a different type of
burrowing activity by the callianassids.

The Weald Clay (Lower Cretaceous) of southern England consists of a thick (up to
360 m.) sequence of clays, sands, silts, and limestones of Barremian age (MacDougall
and Prentice 1964). It has long been realized that the Weald Clay shows marked evidence
of marine conditions, although indications of fresh, brackish/marine, and perhaps near-
terrestrial conditions have been recorded. Thus, as Kilenyi andAlIen (1968) noted, there
are two broad molluscan assemblages: a freshwater one, dominated by Viviparus , but
also yielding unionids, and a much less common assemblage, with Filosina (Casey 1955),
Cassiope, Melanopsis , Nemocardium , Ostrea, and other forms.

Other evidence is seen in the Isle of Wight, where the top of the Wealden Shales
are undoubtedly part marine, with marine molluscs and trace fossils. Echinoid debris
(again fully marine) is recorded from the Weald Clay by Casey (1961) and Allen and
Keith (1965). The latter suggest extensive variation in salinities during Weald Clay
times on the basis of carbon isotope ratios.

Detailed studies on the ostracods have been published; Anderson (1967) described
a complex variation in the ostracod fauna of the formation, which he took as evidence
of the presence of many marine phases. Kilenyi and Allen (1968) described a brackish/
marine microfauna from the lower part of the Weald Clay, but in reviewing Anderson’s
work (op. cit. p. 162), they concluded that only three undoubted marine bands occur in
the Weald Clay of Surrey and Sussex.

In this paper we describe the trace fossil Ophiomorpha nodosa , a marine indicator
not previously recorded from the Weald Clay. It occurs in sandstones at several horizons
and localities. Further detailed microfaunal work is needed to relate these sandstones to
the brackish/marine bands of the ostracod workers.

[Palaeontology, Vol. 12, Part 3, 1969, pp. 459-471, pis. 87, 88.]

460

PALAEONTOLOGY VOLUME 12

SYSTEMATIC DESCRIPTION

Ichnogenus ophiomorpha Lundgren 1891 (for synonymy see Hantzschel 1962)

Type species. Ophiomorpha nodosa Lundgren (1891, p. 114) from the ‘Wealden’ (Lower Cretaceous)
of southern Sweden, by monotypy.

Diagnosis. Medium-sized three-dimensional tunnel systems branching dichotomously
at acute angles, swollen at the point of branching. Tunnels internally smooth, sometimes
filled or lined with ovoid pellets, when the surface of the filling is mammillated.

Ridges on
surface of
burrow filling

A B C

text-fig. 1. Diagram of types of Ophiomorpha occurring in the Weald Clay sandstones. Sectioned
parts are stippled. X 1 approx, a, Type 1, agreeing with the typical form of O. nodosa, b. Type 2, with
meniscus fill (detailed structure of fill omitted), c, Type 3, comparable to Halymenites striatus.

Discussion. Hantzschel (1962) described Ophiomorpha as having ‘tubercle-like or wart-
like ornamentation of the outer wall but smooth inside’. This description is liable to
misinterpretation; Lundgren’s original figures and our material shows that the tunnel
is smooth, whilst the outer surface of the filling has a tubercle-like or wart-like orna-
ment (text-fig. 1a). This is usually clear in lithified sediments, but in unconsolidated
sands the distinction is usually lost.

Ophiomorpha nodosa Lundgren 1 89 1

71836 Ophiomorpha mantelli Nilsson; in Mantell, p. 25 ( nom . nud.)

1842 Spongites saxonicits Geinitz, p. 96 (pars), pi. 23, fig. 2 only. (non. fig. 1, = lectotype of
S. saxonicus).

1847 Cylindrites spongioides Goeppert; Goeppert, p. 359, pi. 35, figs. 1-2; pi. 36, figs. 2-3
(non Goeppert 1841, 1842 (see Hantzschel 1965)).

71852 Spongites saxonicus Geinitz; von Otto, p. 20, pi. 6, fig. 1.

71856 Cylindrites spongiodes Goeppert; Dunker, p. 183, pi. 25, fig. 5.

71858 Halymenites flexuosits Fischer-Ooster, p. 55, pi. 13, fig. 1. (Genolectotype of Ha/y-
menidium Schimpner 1879, vide Hantzschel 1965, 42).

1865 Cylindrites tuberosus Eichwald, p. 8, pi. 4, fig. 13; pi. 5, figs. 1 a, b.

W. J. KENNEDY AND J. D. S. MACDOUGALL: CRUSTACEAN BURROWS 461

1866

1873

71873

1875

1877
71878

1878

1878

1879

1889

1890

1891
1895
1900
1909

1916

1917
71917

1919

71919

1921

1928

1931

1937

1938
1948

1952

1953

1954
1956
1959
1961
1961

1961

1962
1962

1962

1963
1963

1963

1964

1964

1965
1965
1965
1965
1965
1965

Phymatoderma dienvalii Watelet, p. 24, pi. 4, fig. 1.

Halymenites major Lesquereux, pp. 373, 390.

Halymenites minor Fischer-Ooster; Lesquereux, p. 373.

Halymenites major Lesquereux; Stephenson, p. 406.

Broeckia bruxellensis Carter, pp. 382-92, pi. 18.

Halymenites striatus Lesquereux; Lesquereux, p. 37, pi. 1, fig. 6.

Halymenites major Lesquereux; Lesquereux, p. 38, pi. 1, figs. 7, 8.

Halymenites minor Fischer-Ooster; Lesquereux, p. 39, pi. 1, fig. 9.

Halymenites major Lesquereux; Stephenson, pp. 370, 371.

Halymenites major Lesquereux; Eldridge, p. 317.

Halymenites major Lesquereux; Stephenson, p. 532.

Ophiomorpha nodosa Lundgren, pp. 114-15, figs. 1, 2.

Astrophora baltica Deecke, p. 167, pi. 1.

Halymenites major Lesquereux; Knowlton, pp. 17, 18.

Cylindrites spongioides Goeppert emend. Richter; Richter, pp. 8-11 {parts), pi. 9,
fig. 7; pi. 8, fig. 6; pi. 12, fig. 3.

Halymenites major Lesquereux; Knowlton, pp. 86, 87.

Halymenites major Lesquereux; Lee and Knowlton, p. 242.

Halymenites striatus Lesquereux; Lee and Knowlton, p. 243.

Halymenites major Lesquereux; Knowlton, pp. 313-14 {cum. syn.).

Halymenites striatus Lesquereux; Knowlton, p. 314 {cum. syn.).

Halymenites major Lesquereux; Berry, pp. 55, etc.

Astrophora baltica Deecke; Voigt, p. 104.

Halymenites major Lesquereux; Mathias, p. 355, fig. 1.

Halymenites major Lesquereux; Carter, pp. 256-7, pi. 43, figs. 1, 2; pi. 34, figs. 1, 2.
Halymenites major Lesquereux; Stenzel, pp. 68-70, fig. 7.

Terebella sp. Beets, pp. 184-7, figs. 1-5.

Ophiomorpha nodosa Lundgren; Hantzschel, pp. 142-53, pi. 13, 14 {cum syn.).
Halymenites major Lesquereux; Eargle, pp. 143, etc.

Ophiomorpha nodosa Lundgren; Prescher, p. 59.

Ophiomorpha nodosa Lundgren; Seidel, pp. 489-93, figs. 1, 2.

Ophiomorpha Lundgren; Baatz, pp. 168-71.

Ophiomorpha nodosa Lundgren; Toots, pp. 165-70.

Halymenites major Lesquereux; Toots, pp. 165-70.

Halymenites major Lesquereux; Weimer, pp. 88, 89, 95.

Ophiomorpha nodosa Lundgren; Hantzschel, pp. W205-6, fig. 124 (4, 9).

Ophiomorpha Kilpper, pp. 55, 57.

Halymenites or Ophiomorpha', Weimer and Hoyt, p. 321.

Ophiomorpha Lundgren; Hillmer, pp. 137-41, fig. 1.

Callianassa burrows, Hoyt and Weimer, p. 530, fig. 3.

Ophiomorpha-, MacKenzie, pp. 141, 143, pi. 2a, b.

Ophiomorpha tuberosa (Eichwald); Vialov, pp. 163-7, figs. 1-4.

Ophiomorpha, Halymenites', Weimer and Hoyt, pp. 761-7, pi. 123, figs. 7, 8; pi. 124.
figs. 1-7.

Ophiomorpha nodosa Lundgren; Hantzschel, p. 63.

Halymenites major Lesquereux ; Hantszchel, p. 72.

Phymatoderma dienvalii Watelet; Hantzschel, p. 71.

Astrophora baltica Deecke; Hantzschel, p. 13.

Broeckia bruxellensis Carter; Hantzschel, p. 18.

Ophiomorpha nodosa Lundgren; Hoyt and Weimer, pp. 203-7, figs. 1, 2, 5-7.

Material. The present description is based largely on specimens collected from the Weald Clay of
Hambledon (Surrey) and Burgess Hill (Sussex). Much additional material has been examined in the
field, at these and several other localities and horizons.

462

PALAEONTOLOGY, VOLUME 12

Diagnosis. Ophiomorpha with tunnel wall ornamented by single elliptical pellets.
O. nodosa differs from O. borneensis Keij (1965, pp. 224-6, pi. 29, figs. 1-8, text-fig.
2 (1-4), text-fig. 3) by the predominance of vertical as opposed to horizontal elements in
the system and the ornament of single as opposed to bilobate pellets. Neither of these
criteria is particularly satisfactory.

Description. Ophiomorpha nodosa forms tunnel trails between 5 and 15 mm. diameter
with a number of distinct types of surface ornament (text-fig. 1); these variations indi-
cate the type of infillings of the tunnel; the tunnel itself is generally completely smooth.
The tunnel system is extensive and ramifying both vertically and horizontally. Horizontal
elements of the system predominate (though this may be an artefact of preservation)
and are concentrated along sediment interfaces; with the vertical elements they form a
three-dimensional network. The tunnels branch at acute angles at infrequent intervals
(usually about 25 cm.), branching is dichotomous and both branch tunnels are of the
same diameter as the original tunnel. The tunnel may be slightly swollen at the point of
branching. The tunnelling organisms were not phobatactic, and later tunnels cut earlier
tunnels indiscriminately.

There are three main variations in surface ornament, with transitional forms between;
all may occur in a single tunnel. The texture of the sediment within the tunnel is variable,
but always differs from that of the surrounding rock.

Type 1. (Text-fig. 1a; PI. 87, fig. 1 ; PI. 88, fig. 2). This agrees well with the typical form of
Ophiomorpha nodosa. The surface of the tunnel infilling is mammillate, and is built up
of small discoidal pellets, 2-3 mm. wide and 1 mm. thick in the largest burrows; the
size decreases slightly in smaller burrows.

Type 2. (Text-fig. 1b; PI. 87, figs. 1, 2; PI. 88, figs. 3, 4). The surface of the tunnel in-
filling is ornamented by annular ridges. Sections show that these ridges are related to a
well-defined fill of concavo-convex laminae (meniscus filling); these laminae may dis-
integrate to a honeycomb structure; the individual ‘cells’ of the honeycomb are clay
pellets. This type is found more commonly in vertical tunnels.

Type 3. (Text-fig. lc; PI. 87, fig. 1 ; PI. 88, fig. 1). Comparable to Halymenites striatus
Lesquereux. The surface of the tunnel fill is covered by a longitudinal, reticulate, ridge
system. Similar trails of ridges are present on clay/sand interfaces associated with the
sandstones in which Ophiomorpha occurs (PI. 88, figs. 1, 5).

Within the systems in the Weald Clay sandstones, there is little obvious distribution
of types as a whole, other than a suggestion that type 1 is more abundant in the upper
part of systems, type 2 in vertical elements, type 3 in the lowest, often horizontal parts
of burrow systems.

Discussion. We give an extensive synonymy for this trace-fossil; other references are
given by Hantzschel (1952, 1962), whom we follow in regarding Ophiomorpha nodosa as
the most satisfactory name for these burrows.

The association of scratched burrows (type 3) with O. nodosa has led us to include
other scratched burrows occurring associated with Ophiomorpha in the synonymy, i.e.
Spongites saxonicus von Otto {pars, non Geinitz) and Halymenites striatus Lesquereux,
but not scratched burrows not found associated with Ophiomorpha, e.g. Spongelio-
morpha de Saporta 1887 (p. 299, pi. 6, figs. 2, 3).

W. J. KENNEDY AND J. D. S. MACDOUGALL: CRUSTACEAN BURROWS 463

INTERPRETATION

There can be no doubt that Ophiomorpha is a crustacean burrow, a view already con-
vincingly stated by Hantzschel (1952). Weimer and Hoyt (1962, 1964) described the
burrows of the extant marine decapod Callianassa major Say from sand beaches of
Florida and North Carolina, convincingly demonstrating their presence in Pleistocene
deposits nearby, and the identity of these burrows with Ophiomorpha nodosa.

The material from the Weald Clay differs from that previously described, in the
dominance of preservation parallel to bedding planes, a result of the occurrence
of the burrows in, and on, thin, locally cemented sands in an economically exploited
formation, so that the blocks of sandstone accumulate in front of the working face.
Elsewhere, occurrences are in unconsolidated sands, and they are naturally seen in
vertical section only.

In the Weald Clay sandstones, the density of Ophiomorpha systems is low, and the
sediment is not burrowed intensively; depositional structures are never destroyed and
there is no intense bioturbation. Characteristically the tunnels are confined to sand
horizons, only passing through clays when these are thin, i.e. less than 15 cm., clearly
reflecting a preference by the crustaceans for particular depositional environments.

The animals populated only recently deposited unconsolidated sediments, so that
settling structures within the sand bodies, i.e. load casts and syndepositional faults
(see PI. 87, fig. 1) often affected the tunnels.

This is in strong contrast to the associated Equisetiles ( E . burkhartii Dunker), whose
underground stem, root, and tuber systems cut the burrows (and are thus later) and
are unaffected by the settling structures and compaction effects.

The three types of burrows represent different natural processes, which we inter-
pret as follows:

Type 1. This corresponds to the generally accepted form of Ophiomorpha nodosa.
Worked pellets of sediment, moulded and cemented by the callianassid, are pushed into
the sides of the burrow and scraped flat internally (see MacGinitie in Hantzschel
1952, p. 150) or smoothed off by the passage of the body of the inhabitant (as described
by MacGinitie 1930, p. 39, in burrows of the callianassid Upobegia pugeltensis Dana).
In the Weald Clay material the pellets can sometimes be seen to have a brown cement;
in Eocene examples from Upnor, Kent, microchemical tests indicate the presence of
phosphate. Weimer and Hoyt (1964, p. 763) record collophanite as the cementing agent
produced by Callianassa major to bind the sand pellets forming the lining of its burrow.

Type 2. The meniscus-filled Ophiomorpha tunnels can be interpreted in several ways.
According to MacGinitie (1934, p. 167), when individuals of Callianassa ealiforniensis
(Dana) break into adjoining burrow systems, offending tunnels are blocked off and
burrowed round; a similar explanation for our filled material seems quite likely.

Alternatively, this type of burrow may represent the filling-in of early parts of the
tunnel system with material from new excavations, rather than disposal by the ejection
of this material at the surface. Honeycomb structure, observed in some parts of this
type of filling, is a result of the deposition of loads of clay rather than sand alone. A
third interpretation is that these parts of the system were never open, but represent

C 6685 H ll

464

PALAEONTOLOGY, VOLUME 12

the activities of the callianassid as it passed through the sediment, leaving a meniscus
filling as the animal transferred sediment backwards.

These fills thus appear to be somewhat different from the ‘ Halymenites ’ noted and
figured by Brown (1939), which are packed with what are clearly callianassid faecal
pellets.

Type 3. The surface ornament of reticulate ridges on these burrow fillings represent
grooves or scratches on the inside of the burrow. These can be interpreted as scratch-
marks, produced by the appendages of the inhabitant when digging or passing through
the burrow system. Their preservation is the result of lack of subsequent lining or
packing of the burrow, so that the marks produced during the initial excavation are
preserved.

This type of burrow clearly resembles Spongeliomorpha [de Saporta 1 887, regarded here,
and by Reis (1922), Hantzschel (1962, 1965), and Kennedy (1967) as a burrow, although
de Laubenfels ( 1955) treated it as a sponge], the scratched tube of RhizocoraUium (Hantz-
schel 1962, p. W210, fig. 129 b), and the scratched fossil and Recent burrows figured by
Weigelt (1929, pi. 1, 2).

The boring ‘ Terebel/a ’ harefieldensis White (1923), widely distributed in the top of the
Upper Chalk immediately beneath, and filled by, the Lower Tertiary deposits of southern
England (Hester 1965) has a similar ornament (Kennedy 1967). In all cases these appear
to be crustacean burrows or borings.

In all types of Ophiomorpha observed, there are occasional swollen portions, often
close to, or at, the point of branching. These represent ‘turn arounds’ where the animals
were able to change direction in the burrow (see MacGinitie 1930, 1934, for a description
of similar features in Recent callianassid burrows).

The trails of ridges seen on the bottom surfaces of the sandstones containing Ophio-
morpha (PI. 88, figs. 1, 5) are interpreted as the surface tracks of crustaceans, produced
as scratches on a mud bottom. These were almost certainly produced by the Ophiomorpha
inhabitant. They are identical with the trails of Recent crustaceans figured by Weigelt
(1929, pi. 3, figs. 1, 2).

The small rod-like objects occurring abundantly on the bottom surfaces of some of
the sandstones containing Ophiomorpha are similar to the callianassid faecal pellets
described by Moore (1932), Pohl (1946), and Weimer and Hoyt (1964).

OCCURRENCE

Ophiomorpha ranges throughout the Weald Clay, but it is apparently confined to
the more sandy horizons: siltstones, sandstones, and interbedded sandstones and clays.
Table 1 gives the localities at which it has been found, and relates these occurrences to
the divisions of the Weald Clay erected by Topley (1875), and, as far as is possible to
those of Reeves (1953, 1958) and the ostracod zonation of Anderson (in Worssam 1963).

Hambledon , Surrey

Ophiomorpha occurs in thin, silty, largely siderite-cemented sandstones in Topley’s
(1875) no. 7 sand (the Fenhurst sandstone of Wooldridge (1950)), at the Nutbourne
Brickworks, Vann Lane, Hambledon, Surrey (SU 973375). The beds with Ophiomorpha

W. J. KENNEDY AND J. D. S. MACDOUGALL: CRUSTACEAN BURROWS 465

contain no calcareous fossils, but are notable for the occurrence of large numbers of
well-preserved plant remains, identified as Equiselites burkhartii Dunker. Three con-
taining lithologies can be recognized:

Lithology A. Thin sideritic silty sandstones in beds which vary in thickness from 40 to
120 mm., showing little lateral variation. In vertical sections these sandstones show
thinly laminated low-angle cross-bedding and ripple-drift bedding in the upper part.
Minor (20 mm.) textural rhythms occur within the cross-bedding. There is a general
lack of biogenic disturbance, except for burrows of Ophiomorpha (compare with the

Table 1

Ophiomorpha occurrences and Weald Clay stratigraphy

Ophiomorpha

occurrence

Local divisions

Ostracod zonation
(Anderson in Worssam 1963)

Topiey (1875)

Reeves (1953, 1958)

Hambledon

no. 7, sand

Group III

Cypridea vaidensis

no. 6, limestone, ‘Sussex
marble' (large Paludina)
no. 5, sand and sandstone
with calcareous grit
no. 4, limestone
(large Paludina )
no. 3, limestone
(small Paludina)
no. 2, sand and sandstone

newest red clay

Cypridea clavata

Capel, Burgess Hill

Group II

Cypridea tuberculata

oldest red clay

Cypridea dorsispinata

Southwater, Sedgewick
Park

no. 1, Horsham stone

Group I

observations of MacDougall and Prentice 1964), which pass through individual beds
of sandstone and are concentrated along major bedding planes, particularly the bases
of the sandstones. Ecpiisetites stems pass through Ophiomorpha and are filled by sediment
identical with the surrounding sandstone.

The bases of the sandstones are erosional, and show a variety of sole-markings, i.e.
broad, poorly defined load structures, prod-marks, groove-casts, and the ‘A’ and B’
trace-fossils of Prentice (1962). Dessication cracks in the underlying clay are preserved
as reticulate ridges on the base of some of these sandstones.

Lithology B. Massive sideritic sandstones, up to 240 mm. thick, with indistinct bedding.
The top of the sandstones is poorly defined, grading up into sandy clays. The main
mass of the sandstone is penetrated by vertical and horizontal Ecpiisetites stems, with
tubers arising from stems at the nodes. The base of these sandstones is covered by Ophio-
morpha tunnel systems. A syndepositional fault system cuts this surface and the Ophio-
morpha (PI. 87, fig. 1 ).

Lithology C. Micaceous sandstones, 200 mm. thick, breaking into 50-mm. slabs along
clay partings, which contain abundant drifted plant remains. The bases of individual
layers have load casts; other bottom structures also occur. The upper surfaces are

466

PALAEONTOLOGY, VOLUME 12

carious, and have a honeycomb texture due to intense penetration by Ophiomorpha,
with a lot of clay in the burrow filling, which has weathered out, leaving cavities.

Capel, Surrey

At Clock House Brickworks, If miles south of Capel Church (TQ 175385),
Ophiomorpha occurs in Topley’s (1875) no. 5 sandstone, in lithologies similar to those
at Hambledon. The section in the middle face has been described by Kirkaldy and Bull
(1948); it is now badly degraded, but has been confirmed by one of us (J. D. S. M.).
Bed 2 of Kirkaldy and Bull includes the sandstones with Ophiomorpha, and two litho-
logies can be recognized: micaceous sandstones, similar to the Hambledon lithologies,
in beds up to 200 mm. thick, sideritic, with abundant Ophiomorpha-, and thin siltstones,
(ripple-marked with an erosional base) in which basin cast structures (Prentice 1962,
pi. 3, fig. 2), varying from 50 to 250 mm. thickness are developed.

The base of these siltstones is covered by groove-casts and prod-marks, which are
occasionally interrupted by Ophiomorpha (PI. 87, fig. 2). The siltstones are penetrated
by obscure vertical tubes, similar in diameter to stems of Ecpdsetites burkhartii. Apart
from these, we have failed to detect the Equisetites recorded by Kirkaldy and Bull at
this level.

An interesting feature of this section is the occurrence of Filosina [ Neomiodon } in
some numbers, above bed 3 of Kirkaldy and Bull (1948).

In the lower pit at Capel, a further section includes clays with Paraglauconia strombi-
formis (Schlotheim), Filosina gregarea Casey, oysters, and an abundant ostracod fauna
including Schuleridia, arenaceous foraminifera, and cirripedes (Kilenyi and Allen 1968),
suggesting that these beds were laid down in marine or brackish waters.

Burgess Hill, Surrey

Ophiomorpha occurs at Keymer Brickworks, near Wivelsfield station, Burgess Hill,
Surrey (TQ 324193), in massive, micaceous sandstones up to 200 mm. thick, ripple-drift
bedded in the upper part. These sandstones appear to have been dredged up from the
lower part of the succession, where silty clays and shaly clays with ostracods, passing
up into red clays and sandstones, are seen; the succession is rather like that at Capel.
The beds with Ophiomorpha appear to be Topley’s (1875) no. 5 sandstones.

EXPLANATION OF PLATE 87

Figs. 1, 2. Ophiomorpha nodosa. 1, Bottom surface of a sandstone block from Hambledon, Surrey,
showing all three forms of Ophiomorpha (as indicated), load casts, syndepositional faults, and
Equisetites burkhartii (E); x 1. 2, Bottom surface of a basin cast from Clockhouse brickworks,

Capel, Surrey, showing a specimen of Type 2 Ophiomorpha, with prominent pelletal fill, cutting a
surface covered with aligned groove- and prod-marks; x 1.

EXPLANATION OF PLATE 88

Figs. 1-5. Ophiomorpha nodosa and associated trace fossils. 1, Type 3 Ophiomorpha running parallel
to bedding, with a trail of ridges to the left. 2, Type 1 Ophiomorpha, showing characteristic pelletal
surface; a tuber of Equisetites burkhartii is visible at the top right of the photograph. 3, Type 2
Ophiomorpha, showing ornament of burrow fill. 4, The same, vertical longitudinal section,
showing meniscus fill. 5, Trail of ridges from bottom surface of Ophiomorpha — bearing sandstone.
All photographs (except 4, a section) are the lower surfaces of sandstones from Hambledon,
Surrey; figs. 1-3, 5 are X 2; fig. 4 is x4.

Palaeontology, Vol. 12

PLATE 87

KENNEDY and MacDOUGALL, Crustacean burrows in Weald Clay

Palaeontology, Vol. 12

PLATE 88

KENNEDY and MacBOUGALL, Crustacean burrows in Weald Clay

W. J. KENNEDY AND J. D. S. MACDOUGALL: CRUSTACEAN BURROWS 467

Southwater, Surrey

The topmost bed of the Horsham Stone (Topley 1875, no. 1 sandstone) in Southwater
no. 2 Brickpit, Castle Wood, Southwater, is intensely penetrated by Ophiomorpha.
Fragments of crustacean carapaces and rootlets of Equisetites are occasionally found in
both sandstone and clay.

Sedgewick Park and Slinfold, Surrey

Ferguson (1926) recorded three types of sandstone in the Horsham Stone, very similar
to those recognized by us at Hambledon. At a quarry (now filled in) 800 m. west of
Southwater Church (TQ 152263) he recorded (p. 408) what may be Ophiomorpha :
‘peculiar cylindrical bodies, possibly animal tracks or burrows, are observed, and car-
bonaceous fragments are not uncommon’.

Ophiomorpha also occurs in Ferguson’s type A sandstone in the Upper Horsham Stone
at a natural exposure at Sedgewick Park, Surrey (TQ 179271).

OPHIOMORPHA AS AN ENVIRONMENTAL INDICATOR

Ophiomorpha structures in Recent sediments. Weimer and Hoyt (1962, 1964) have shown
beyond doubt that Ophiomorpha- like structures are being produced in present day
sediments by callianassids (marine decapod crustaceans), particularly Callianassa major
Say. They recorded this animal primarily from the low littoral zone, between mean
sea-level and low tide on beaches that face the open ocean (1964, fig. 2), in well-sorted
sand in strongly wave-agitated waters. They also noted some occurrences just above
mean sea-level, and in the shallow neritic zone, up to T5 m. below mean low water,
and suggested that the burrows extend to greater depths.

Fossil occurrences. Extending their observations to adjacent Pleistocene sands, Weimer
and Hoyt successfully demonstrated that the occurrence of Ophiomorpha in these indi-
cated their deposition in shallow neritic and/or littoral conditions. They suggested that
Tertiary and Cretaceous occurrences indicated a similar environment; burrows in
well-sorted massive-bedded sandstones indicated littoral or shallow neritic conditions;
burrows in poorly sorted, well-bedded silty sandstone suggested deeper neritic conditions.

In spite of this definitive view of the environment indicated by Ophiomorpha , i.e.
wholly marine, littoral/sub-littoral, there is a range of suggested environments in the
very extensive literature on this trace fossil (Baatz 1959, Hillmer 1963, Mathias 1931,
Ortmann 1925, Seidel 1956, etc.)

Ophiomorpha is recorded in sequences which contain lignites, marine fossils (including
ammonites), freshwater fossils (including unionids), quasimarine or brackish-water
forms, and mixed assemblages in the same bed. Other authors have recorded faunas
suggesting varying salinities in the same sequence, features which require explanation.

Considering the occurrence of a trace-fossil of this type, the environment indicated
by the burrow is not that of the sediment the burrows occur in, nor of the sediment
above the surface whence the burrows originate, nor any fauna these sediments may
contain. The environment indicated is that existing during part, or all, of the period
represented by the surface from which the burrows originate.

If this is borne in mind, it is readily understood how Ophiomorpha can occur associated

468

PALAEONTOLOGY, VOLUME 12

with marine, brackish, and freshwater fossils, and how the sediment they occur in
could become colonized by plants (for example, in the case of the Weald Clay, by horse-
tails). If the environment of the Recent producers of Ophiomorpha structures is con-
sidered, this is the very environment (the transition between marine/brackish-freshwater/
terrestrial conditions — a shoreline) where mixed assemblages could accumulate, and
where slight movements of the littoral zone would give the vertical faunal variation
found with so many occurrences of Ophiomorpha.

Ophiomorpha is a marine indicator of littoral and shallow neritic environments
(Weimer and Hoyt 1962, etc.). The present records are in keeping with the picture
emerging of the Weald Clay depositional environment. Thus the presence of Ophio-
morpha, together with the general form and lateral extent of the sandstones in which
it occurs, suggests that the latter were sand bodies, lying parallel to the contemporary
Weald Clay shoreline. These sands were colonized by the callianassids soon after
deposition, since early post-depositional (and perhaps even syndepositional) features such
as load casts and fault systems disrupt their burrows. At about the same time as coloniza-
tion occurred, the upper parts of the sand bodies were gently reworked, so that ripple-
drift bedding forms the upper part of many beds. Subsequently, either as a result of
local changes in geography, or the waning influence of marine conditions, the sand
bodies were colonized by semi-aquatic plants (horsetails, all apparently Equisetites
burkhartii). This colonization was later than the general compaction and settling of the
sediment, as the roots pass through burrows, load casts, and fault systems without
disturbance. This interpretation of the environment of the callianassids which inhabited
the Ophiomorpha burrow systems is in keeping with the general pattern of known
palaeosalinities as based on isotopic work (Allen and Keith 1965, plus additional
unpublished data), the occurrence of brackish water or marine molluscs, and ostracods
(Kilenyi and Allen 1968), and the well known occurrence of non-marine fossils and
rootlet beds.

Acknowledgements. We are grateful to Mr. B. C. Worssam for bringing the Weald Clay occurrences
of Ophiomorpha to our attention, and to Mr. J. N. Allen and Dr. T. I. Kilenyi for discussion of their
data on Weald Clay ostracods. Dr. J. M. Hancock gave considerable assistance in the field, Mr. R.
Cleevely helped in the location of some of the more obscure references. The assistance of the technical
staff of the Departments of Geology, Sir John Cass College (London), King’s College (London), and
the Department of Geology and Mineralogy, Oxford, is acknowledged. We are particularly indebted
to Dr. J. E. Prentice for much useful discussion and to Dr. W. S. McKerrow for criticizing early
versions of the manuscript of this paper.

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stenzel, h. b. 1938. The geology of Leon County, Texas. Univ. Texas Pub. 3818, 259. 1 pi.

W. J. KENNEDY AND J. D. S. MACDOUGALL: CRUSTACEAN BURROWS 471

Stephenson, j. j. 1875. Report on the geology of a portion of Colorado. Rept. U.S.geol. Surv. Terri-
tories, 3, 406.

1879. Note on the Fox Hills Group of Colorado. Am. J. Sci. (3)17, 369-73.

1890. In Discussion of newberry, j. g., The Laramie Group. Bull. geol. Soc. Am. 1, 529-32.

Sternberg, K. M. graf von. 1833. Versuch einer geognostisch-botanischen Darstellung cler Flora der
Vorwe/t. 5, 6, 80 pp. Leipzig and Prague.

toots, h. 1961. Beach indicators in the Mesa-Verde Formation. In Wyoming Geol. Ass. 16th. Ann.
Field Conference Guidebook, 165-70.

topley, w. 1875. The geology of the Weald. Mem. geol. Surv. U.K. 503 pp.

vialov, o. s. 1964. O Prirodya ees Palaeogyena priolya. Bjul. mosk. Obsh. Ispvtat. Prirody, Otdel.
geol. 39, 163-7.

voigt, e. 1928. Kocherbauten von Wiirmern in Sedimentargeschieben. Z. gescheibforschg. 4, 94-104.

watelet, a. 1866. Description des plantes fossiles du bassin de Paris. 264 pp. 60 pi. Paris.

weigelt, j. 1929. Fossile Grabschachte brachyurer Decapoden als Lokalgeschiebe in Pommern und
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neritic environments. Spec. Pap. geol. Soc. Am. 68, 321.

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white, e. i. 1923. Notes on a new species of terebelloid and other phenomena in the Great Pit at
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Wooldridge, s. w. 1950. Some features of the structure and geomorphology of the country around
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worssam, B. c. 1963. Geology of the country around Maidstone. Mem. Geol. Surv. U.K. 152 pp. 5 pi.

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W. J. KENNEDY

Department of Geology and Mineralogy
Parks Road
Oxford

J. D. S. MACDOUGALL
Department of Geology
Sir John Cass College
Jewry Street
London E.C. 3

Typescript received 27 November 1968

NEW SPIRIFERID BRACHIOPODS FROM THE
LOWER DEVONIAN OF NEW SOUTH WALES

by N. M. SAVAGE

Abstract. Five species and subspecies of spiriferid brachiopods are described from the early Siegenian Man-
dagery Park Formation, New South Wales: Cyrtina praecedens Kozlowski, Proreticularia beddiei sp. nov., and
the new subspecies Quadrithyris robusta molongensis, Ambocoelia praecox dorsiplicata , and Howellella nucida
australis. There is a close resemblance to species from eastern Europe and a genera! affinity to species from central
Asia.

Recent investigations in the vicinity of Manildra, New South Wales, have led to the
discovery of several interesting Lower Devonian brachiopod assemblages. Of these
the most important consists of delicately preserved silicified material of probable early
Siegenian age. Two new genera from this fauna have been described (Savage 1968 a , b )
and several complementary papers are in preparation. This paper is concerned with the
spiriferids.

The Lower Palaeozoic deposits of the Manildra district form part of the Cowra
Trough sediments of the Lachlan geosyncline (Packham 1960). Within this depositional
trough a thickness of at least 10 000 ft. of sediments accumulated during Silurian and
Early Devonian times. No detailed palaeontological work took place until Hill and
Jones (1940) and Hill (1942) established the Lower to Middle Devonian age of corals
from the Garra Limestones immediately to the east. More recently Strusz (1965 a , b ,
1966, 1967) has investigated the Garra Formation further and described many more of
the corals.

The stratigraphy of the Manildra district has been described in detail elsewhere
(Savage 1969) together with faunal lists and a discussion of possible correlations. It is
suggested that a new group, the Gregra Group, should consist of the Maradana Shale,
the Mandagery Park Formation, and the Garra Formation. The first two are new forma-
tions, of late Gedinnian and Siegenian age, respectively. The third is a relatively well-
known sequence of limestones and shales of probable late Siegenian to early Eifelian
age. All three formations are calcareous, shallow-water deposits with rich shelly
faunas.

The Lower Devonian spiriferids from the Mandagery Park Formation show a close
resemblance to forms from Bohemia and Podolia, and it is with these forms that the
present taxonomic discussion is largely concerned. There is also a general affinity
with other spiriferids from the whole Eurasian faunal province, which in Early Devonian
times extended from eastern Europe, across Asia, and into western North America.
These affinities are also evident in other groups of Manildra brachiopods.

Tn the systematic treatment below, specimen numbers used are those of the Palaeontology Collection,
Department of Geology and Geophysics, University of Sydney.

[Palaeontology, Vol. 12, Part 3, 1969, pp. 472-87, pis. 89-92.1

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

473

SYSTEMATIC PALAEONTOLOGY

Phylum BRACHIOPODA
Suborder spiriferoidea
Superfamily reticulariacea Waagen 1883
Family reticulariidae Waagen 1883
Genus quadrithyris Havlicek 1957

Type species. Spirifer robustus Barrande 1848, by original designation.

Quadrithyris robusta molongensis subsp. nov.

Plate 89

Diagnosis. A form close to Q. robusta (Barrande) but with a more prominent fold and
sulcus. The lateral slopes bear no trace of plications.

Material. The total of 61 silicified specimens consists of 16 complete or nearly complete conjoined
shells, 15 dorsal valves, and 30 ventral valves. Specimen SU 19590 is designated the holotype.

Description. Exterior. The shell is transversely elliptical in outline with the maximum
width between mid-length and the posterior margin. The cardinal margins are evenly
rounded and the anterior margin is straight or emarginate. In lateral profile the shell
is strongly biconvex with the thickness almost equal to the length and greatest near the
umbones.

The strongly convex ventral valve has a prominent umbo and an erect to slightly
incurved beak. An apsacline interarea is concave with an apical angle of 110-20°.
The delthyrium includes an angle of about 45°. Narrow, ribbon-like, deltidial plates
project normal to the interarea along the delthyrial margins (PI. 89, fig. 26).

The dorsal valve is less strongly convex than the ventral valve. It has a broad umbo
and a prominent, incurved beak. The interarea is low and orthocline to apsacline with
an open notothyrium which includes an angle of 125-35°.

The ventral valve bears a broad, rounded sulcus and the dorsal valve a corresponding
fold. Both commence at the umbones and extend to the strongly uniplicate anterior
commissure. The surface ornament consists of fine growth lamellae spaced at 5 or 6 per
mm.

Ventral interior. A deep delthyrial cavity is bounded by strongly receding, subvertical
dental lamellae which converge posteriorly before meeting the valve floor (PI. 89, fig. 33).
The teeth are small and stubby with a subtriangular cross-section. A prominent median
septum, extending half the valve length, is high in the middle with a concave edge
anteriorly and dorsally (PI. 89, fig. 30). The ventral muscle field is not sufficiently
impressed to be discernible in the material available.

Dorsal interior. Long, widely divergent sockets are supported on strong hinge plates
(PI. 89, figs. 28, 29). On their inner edges the sockets are bounded by strong inner
socket ridges, triangular in section and becoming higher and stronger anteriorly
(text-fig. 1). Narrow crural lamellae slope sharply down from the socket ridges but do
not reach the valve floor. Slender crural bases are attached along the inner edges of these
crural lamellae (PI. 89, fig. 28). A large, well-rounded, cardinal process is occasionally

474

PALAEONTOLOGY, VOLUME 12

preserved in this silicified material. In some specimens traces of several lobes are present
(text-fig. 1). A partly complete laterally directed spire with 4 volutions is present in
one specimen and it appears that a complete spire would have 5 or 6 volutions. The
muscle field is long and narrow and extends within the dorsal fold about one-third of
the distance to the anterior commissure (PI. 89, figs. 28, 31, 32).

text-fig. 1. Quadrithyris robusta molongensis subsp. nov.
Reconstruction of the dorsal cardinalia based on several
incomplete specimens. Approx. X 10.

Measurements. The

dimensions of 4 specimens are

given below in mm.

Length

Width

Thickness

SU

19588

Complete shell

5-8

70

4-7

SU

19589

Complete shell

80

101

6-9

SU

19590

Complete shell

8-9

11-5

7-2

SU

19597

Dorsal valve

9-6

13 6

—

Ontogeny. From the relatively small and broken collection, sufficient specimens of
different ontogenetic stages are present to allow an examination of the morphological
development during most of the growing period. The youngest form available has only
a very gentle fold and sulcus and a ventral umbo which does not project posteriorly

EXPLANATION OF PLATE 89

Figs. 1-35. Quadrithyris robusta molongensis subsp. nov. Mandagery Park Formation, Manildra.
1-25, Dorsal, ventral, posterior, anterior, and lateral views of five specimens of progressively in-
creasing size. 1-5, SU 19586, a very young stage with a low fold and sulcus (fig. 4) and a ventral
umbo which does not project posterior of the hinge-line (fig. 1). 6-10, SU 19587, a young stage

also with a low fold and small ventral umbo. 11-15, SU 19588, a larger form with a prominent
ventral umbo (fig. 15), a distinct ventral interarea and ribbon-like deltidial plates projecting normal
to the interarea (fig. 13). 16-20, SU 19589, a mature form with a pronounced fold and sulcus

(fig. 19), and incurved umbones (fig. 20). 21-5, SU 19590 (holotype), a relatively large form with a

ventral umbo projecting well past the hinge-line and a strongly uniplicate anterior commissure
(fig. 24). 26, Postero-lateral view of SU 19590 showing the ventral interarea and the thin ribbon-

like deltidial plates projecting normal to it. 27, Antero-ventral view of dorsal valve SU 19595.
28, 29, Antero-ventral and ventral views of dorsal valve SU 19597 showing the inner socket ridges
becoming higher and stronger anteriorly and the narrow inner hinge plates suspended from these
ridges and not meeting the valve floor (fig. 28). 30, Lateral view of large broken ventral valve

SU 19598 showing the long median septum with its concave anterior edge. 31, 32, Antero-ventral
and ventral views of dorsal valve SU 19591 showing the widely divergent sockets, the strong inner
socket ridges, and the adductor muscle scars. 33-5, Dorso-anterior, dorso-lateral, and dorsal
views of ventral valve SU 19592 showing the strongly receding dental lamellae (fig. 35), and the long
tooth ridges. (Figs. 1-26 x 3, figs. 27-35 x4.)

Palaeontology , Vol. 12

PLATE 89

SAVAGE, Quadrithyris

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

475

past the hinge line (PI. 89, figs. 1-5). As growth proceeds the fold and sulcus become more
pronounced and the ventral umbo extends well past the hinge line (PI. 89, figs. 21-5).
Both the dorsal and ventral beaks change from an erect position in the younger stages,
to an incurved position in the mature shell (PL 89, figs. 10, 15, 25).

Internally the ventral median septum becomes relatively longer and the dental lamellae
relatively shorter, whilst in the dorsal valve the hinge plates thicken and a low median
myophragm becomes visible in the muscle field.

Discussion. The relatively smooth lateral slopes and growth lamellae, in combination
with the high ventral median septum and prominent dental lamellae, place this form in
the genus Quadrithyris Havlicek. The closely related genus Quadrithyrina Havlicek
lacks dental lamellae.

Q. robusta molongensis is distinguished by receding dental lamellae, lateral slopes
with no trace of plications, and a high median septum extending half the valve length.
It closely resembles specimens of Q. robusta (Barrande) although the Bohemian type
species has a less-pronounced fold and sulcus. Other Bohemian species are Q. kotysensis
Havlicek and Q. falco (Barrande). Both have a sulcus bounded on each side by a low
plication and a fold bounded by weak furrows. Q. trisectus (Kayser), from the Rhenish
Schiefergebirge, differs from the Manildra form, and also from the type species, in
possessing a very long median septum flanked by almost equally long dental lamellae.

Q. robusta molongensis is one of the earliest species of Quadrithyris known. Siegenian
species have been recorded from Nevada by Johnson (1965) and from the Altai Moun-
tains by Kulkov (1963) but all other recorded species are from deposits of Emsian or
Eifelian age.

Genus proreticularia Havlicek 1957
Type species. Spirifer carens Barrande 1879, by original designation.

Proreticularia beddiei sp. nov.

Plate 90, figs. 22-38

Diagnosis. A wide Proreticularia with evenly rounded lateral margins and a strongly
incurved ventral umbo.

Material. A total of 120 silicified specimens consists of 29 complete or nearly complete conjoined
shells, 45 dorsal valves, and 46 ventral valves. Externally very fine growth lines and radial striations
are preserved but no spines have been observed. Specimen SU 16620 is designated the holotype.

Description. Exterior. In outline the shell is transversely oval with the greatest width at
about mid-length. The cardinal and lateral margins are evenly rounded and the anterior
margin is gently rounded to emarginate. The lateral profile is unequally biconvex
with the ventral valve the deeper. Maximum thickness is just posterior of mid-length.

The ventral valve is strongly convex with the greatest curvature in the high umbonal
region. An apsacline, strongly concave interarea has a curvature which increases apically
and a width slightly more than half the maximum shell width. The beak ridges are very
weak. An open delthyrium includes an angle of 40-50° and is bordered by prominent
deltidial plates which project normal to the interarea (PI. 90, fig. 24).

476

PALAEONTOLOGY, VOLUME 12

The dorsal valve is convex in lateral profile with a broad umbo terminating in a small
incurved beak. A concave interarea is apsacline and very short with an apical angle of
about 150°. The notothyrium is open and includes an angle of 125-35°.

The ventral valve has a faint sulcus and the dorsal valve a barely discernible fold.
Where they meet anteriorly the commissure is rectimarginate or weakly uniplicate
(PI. 90, fig. 25). A surface ornament of fine concentric growth-lines also shows traces
of very fine radial lines (PI. 90, fig. 22). No spines are visible, probably because of the
nature of the preservation.

Ventral interior. The ventral interior is without dental lamellae. Narrow tooth ridges
border the delthyrium and thicken markedly on the inner surfaces prior to projecting
as short strong teeth. In a single specimen the ventral muscle field is visible as a number
of linear radiating impressions (PI. 90, fig. 29).

Dorsal interior. Small hinge plates extend for about one-third of the maximum shell
width (PI. 90, fig. 28). The cardinal process, placed at the very apex of the notothyrium,
is poorly preserved in most of the material but in some specimens occurs as a wide
structure with several ill-defined lobes (text-fig. 2). Deep, narrow sockets are widely
divergent at about 140° and are supported on narrow hinge plates. The inner socket
ridges are low proximally but become wider and higher distally to terminate in pro-
jections which articulate with crural fossettes (PI. 90, figs. 27, 28). Thin, triangular
crural lamellae slope downwards from the inner edges of the socket ridges but do not
meet the valve floor. They extend between one-fifth and one-quarter the distance to the
anterior margin and are strongly convergent downwards and moderately divergent

EXPLANATION OF PLATE 90

Figs. 1-21. Ambocoelia praecox dorsiplicata subsp. nov. Mandagery Park Formation, Manildra.
1-5, Dorsal, ventral, posterior, anterior, and lateral views of SU 16614 showing the large ventral
interarea (fig. 3), and the small dorsal plication in the anterior commissure (fig. 4). 6-10, Dorsal,

ventral, posterior, anterior, and lateral views of SU 16609 showing the more inflated ventral umbo
(fig. 10), and the gently plicate anterior commissure (fig. 9). 11-15, Dorsal, ventral, posterior,

anterior, and lateral views of SU 16610 showing the slightly emarginate anterior margin (fig. 11)
and the high, gently concave interarea (fig. 15). 16-18, Three dorsal views of ventral valve SU

16612 showing the narrow tooth-ridges along the delthyrial margins and the absence of dental
lamellae and a median septum. 19, 20, Dorsal and ventral views of dorsal valve SU 16615 showing
the bifid cardinal process, the long widely divergent sockets, and the broad crural lamellae meeting
the valve floor posteriorly. 21, Ventral view of dorsal valve SU 22668 together with a crus broken
from it. (All figures x 9.)

Figs. 22-38. Proreticidaria beddiei sp. nov. Mandagery Park Formation, Manildra. 22-6, Dorsal,
ventral, posterior, anterior, and lateral views of SU 16620 (holotype) showing the distinct fold and
sulcus (fig. 25), the emarginate anterior margin (fig. 23), the narrow strip-like deltidial plates project-
ing normal to the interarea (fig. 24), and the large incurved ventral umbo (fig. 25). 27, Antero-

ventral view of dorsal valve SU 16622 showing the broad cardinal process and the large sub-triangular
crural lamellae. 28, Antero-ventral view of dorsal valve SU 1 6623 showing the hinge plates support-
ing the large crural lamellae. 29, Dorsal view of large ventral valve SU 16621 showing the narrow
tooth ridges, the absence of a median septum, and the large deeply striated muscle field. 30-4,
Dorsal, ventral lateral, anterior, and posterior views of SU 16617, a young form with an almost
perfectly elliptical outline and a rectimarginate anterior commissure (fig. 34). 35-8, Dorsal,

anterior, lateral, and posterior views of SU 16616, a very young form. (All figures x6.)

Palaeontology, Vol. 12

PLATE 90

SAVAGE, Ambocoelia and Proreticularia

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

477

anteriorly (PI. 90, figs. 27, 28). From the lower edges of the crural lamellae slender
crura curve upwards. Spires have not been observed. The dorsal muscle field extends
between one-third and one-half the distance to the anterior margin. It is divided into a
narrow, elongate pair of medial adductors confined to the faint internal impression of
the fold, and a pair of shorter, more rounded lateral adductors.

text-fig. 2. Proreticularia beddiei sp. nov. Reconstruction of
the dorsal cardinalia based on several incomplete specimens.
Approx. x20.

s'. The dimensions of 5 specimens are given

Length

below in mm.
Width

Thickness

SU 16616

Complete shell

1-5

20

1-3

SU 16617

Complete shell

1-9

2-6

1-7

SU 16620

Complete shell

4-5

60

3-6

SU 16621

Ventral valve

4-8

6-9

■ — -

SU 16622

Dorsal valve

4-2

6-8

—

Ontogeny. This species is not well represented in the collection and most of the specimens
are damaged to some degree. As with the other spiriferid genera described herein, the
most obvious ontogenetic changes are the accentuation of the fold and sulcus, the
increasing prominence of the ventral umbo posteriorly, and the incurving of the beaks
(PI. 90, figs. 22-6). In this species the dorsal fold is very gentle in even the mature forms,
and in the youngest stages it is scarcely visible at all (PI. 90, fig. 36). As growth proceeds
the ventral sulcus becomes more marked than the dorsal fold with the result that the
anterior margin is emarginate in the mature specimens.

Discussion. This Manildra form resembles the type species from the Ludlovian Kopanina
Limestone, Bohemia. However, specimens of Proreticularia carens, sent to the author
by Dr. Havlicek, are less wide and have a less incurved ventral umbo than P. beddiei
and this difference is also clear from the published illustrations (Havlicek 1959, pi. 25,
figs. 1, 2). P. Candida Havlicek, 1959, from the Koneprusy Limestone, is also narrower
than the Manildra species and it differs further in having shallow furrows bounding
the low dorsal fold. Proreticularia has not been recorded previously outside Bohemia
and Asiatic Russia.

478

PALAEONTOLOGY, VOLUME 12

Superfamily cyrtiacea Frederiks 1919 (1924)

Family ambocoeliidae George 1931
Genus ambocoelia Hall 1860

Type species. Orthis umbonata Conrad 1842, by original designation.

Members of the Ambocoeliidae show considerable variation, even from the same
locality, and precise distinctions between species, and even genera, are often difficult
for this reason. Vandercammen (1956) attempted to group the genera primarily on the
basis of dental lamellae, separating those with dental lamellae from those without,
whereas Havlicek (1959) laid particular emphasis on the presence or absence of a fold
or sulcus in the dorsal valve. Neither of the resulting groupings has proved satisfactory
and in the Brachiopod treatise (1965) Pitrat has not attempted to subdivide the Ambo-
coeliidae into subfamilies.

The Manildra species described below shows considerable variation in the degree
of plication (see PI. 90) and it is unlikely that minor departures of the anterior com-
missure from the rectimarginate condition are of generic significance. In the form of the
interarea, and the internal features of both the ventral and dorsal valves, the Manildra
form closely resembles the type species of Ambocoelia and it is to this genus that it is
referred herein.

Ambocoelia praecox dorsiplicata subsp. nov.

Plate 90, figs. 1-21

Diagnosis. A form of the species Ambocoelia praecox Kozlowski with a weak dorsal
fold and a uniplicate anterior commissure.

Material. Of a total of 55 silicified specimens, 30 are complete or nearly complete shells with conjoined
valves, 10 are dorsal valves, and 15 are ventral valves. The internal features, including spiralia and
crura, are often visible in this material but the finer external detail is not preserved. Specimen SU
16609 is designated the holotype.

Description. Exterior. The shell is small, subquadrate to semicircular in outline, and
widest near the hinge line. The cardinal margins are sharply rounded and the anterior
margin is often emarginate. In lateral profile the shell is very unequally biconvex.

The ventral valve is deep with a swollen, strongly arched umbo and a suberect beak
(PI. 90, figs. 5, 15). The lateral slopes are gently concave postero-laterally, and gently
convex antero-laterally. A high interarea is apsacline and concave, with the curvature
increasing apically; it has a width about two-thirds that of the shell and an apical angle
of about 90°. The large open delthyrium is triangular and includes an angle of 25-35°.

The dorsal valve is evenly convex with a broad umbo. A low, concave interarea is
anacline to orthocline and the notothyrium is open, often with the cardinal process
visible apically.

A low sulcus on the ventral valve extends from the umbo. The dorsal valve has a
low fold, bounded either side by a broad furrow. Anteriorly the fold and sulcus form
a gently uniplicate commissure (PI. 90, figs. 4, 9, 14). The shell surface is smooth, or
ornamented with fine concentric growth lines. Fine radial striations are present, and
often a further set of stronger growth lines is irregularly superimposed (PI. 90, fig. 19).
No spines have been observed.

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

479

Ventral interior. The ventral interior is without dental lamellae. Narrow tooth ridges,
which border the delthyrium, are continuous with the teeth. No median septum is
present. The muscle field has not been observed in the material available.

Dorsal interior. The hinge plates extend from one-half to two-thirds of the valve width.
A shallow notothyrial cavity is partly filled by a bifid cardinal process which has a long,
inclined shaft (PI. 90, fig. 20). The sockets are long and widely divergent (text-fig. 3).
They are bounded posteriorly by the valve margin and anteriorly by narrow inner socket

text-fig. 3. Ambocoelia praecox dorsiplicata subsp. nov.

Reconstruction of the dorsal cardinalia based on several
incomplete specimens. Approx. X 30.

ridges which are low proximally, but become higher distally. Broad crural lamellae
rest on the valve floor with their inner edges almost parallel (PI. 90, fig. 20), but occasion-
ally they only join the valve floor posteriorly and are suspended from the socket plates
for most of their length. The lamellae are strongly convergent downwards and moderately
divergent anteriorly. The crura are long and subparallel with short, hook-like, ventrally
curved processes projecting from them at about one-third the valve length (PI. 90, fig. 21).
The spines are directed laterally, each with 4 or 5 volutions. A long, narrow, muscle
field extends three-quarters of the valve length but the individual impressions are in-
distinct.

Measurements. The dimensions of 5 specimens are given below in mm.

Length

Width

Thickness

SU 16609

Complete shell

1-6

2-3

1-5

SU 16610

Complete shell

1-7

20

1-6

SU 16612

Ventral valve

1-7

2-2

—

SU 16614

Complete shell

19

2-6

1-9

SU 16615

Dorsal valve

2-6

3-5

—

Discussion. Considerable variation of form is present in this material and it is difficult
to find two specimens really alike. Examples of this variability are seen in Plate 90.
Ambocoelia praecox Kozlowski, from the Borszczow beds of Podolia, is very close to the
Manildra material. The only real difference is the rectimarginate anterior commissure
of the Podolia species compared with the dorsal fold in the form from New South
Wales.

Of the Bohemian species, A. operculifera Havlicek, from the early Emsian Reporyje
Limestones, differs from the Manildra form in being distinctly sulcate, though it is

480

PALAEONTOLOGY, VOLUME 12

worthy of note that one of the figured specimens (Havficek, 1956, fig. 17) shows very
weak plications laterally. The younger species, A. mesodevonica Havlicek, from the
Middle Devonian Acanthopyge Limestones, also shows weak plications anteriorly.
Both A. praecox from Podolia, and A. praecox dorsiplicata from Manildra, differ from
the North American type species in having a higher ventral interarea and a non-sulcate
dorsal valve.

Superfamily delthyridacea Phillips 1841
Family delthyrididae Phillips 1841
Subfamily acrospiriferinae Termier and Termier 1949
Genus howellella Kozlowski 1946

Type species. Terebratula crispus Hisinger 1826, by original designation.

Howellella nucula australis subsp. nov.

Plate 91

Diagnosis. A form of Howellella close to H. nucula (Barrande) but with a very small
median septum in the extreme apex of the umbonal cavity.

Material. A total of 716 silicified specimens consists of 426 complete or nearly complete shells with
conjoined valves, 157 dorsal valves, and 133 ventral valves. The internal features, including the spiralia
in some cases, are well preserved but the finer external ornament is not present. Specimen SU 16603
is designated the holotype.

Description. Exterior. The shell is small and transversely oval in outline with the
maximum width just posterior of mid-length. It has broadly rounded cardinal margins
and more gently rounded anterior and lateral margins. In lateral profile the shell is
strongly biconvex with the ventral valve deeper than the dorsal valve. The thickness is
often equal to the length with the maximum thickness at about mid-length and the
anterior slopes almost vertical.

EXPLANATION OF PLATE 91

Figs. 1-35. Howellella nucula australis subsp. nov. Mandagery Park Formation, Manildra. 1-5,
Dorsal, ventral, posterior, anterior, and lateral views of SU 16602. 6, Antero-ventral view of

dorsal valve SU 16608 showing the conspicuous cardinal process and the large triangular crural
lamellae. 7, Antero-lateral view of broken specimen SU 16605 showing part of the exposed spire.
8-12, Dorsal, ventral, posterior, anterior, and lateral views of SU 16601. 13-15, Dorsal, dorso-

lateral, and antero-dorsal views of ventral valve SU 16606 showing the dental lamellae and low
myophragm. 16-20, Dorsal, ventral, posterior, anterior, and lateral views of SU 19599, a young
stage showing a relatively gentle fold and sulcus (fig. 19), and dorsally directed beaks (fig. 20).
21-6, Dorsal, ventral, posterior, anterior, lateral, and postero-lateral views of mature specimen
SU 16603 (holotype) showing the prominent ventral umbo (fig. 25) and pronounced fold and sulcus
(fig. 24) which are typical of the larger specimens. 27, Ventral view of dorsal valve SU 16604 showing
the divided hinge-plate, the broad cardinal process, the widely divergent sockets, and the large
crural lamellae suspended from the socket ridges. 28-30, Dorso-lateral, antero-dorsal, and dorsal
views of ventral valve SU 16607 showing the gently advancing dental lamellae (fig. 28), the small
median septum at the extreme apex of the umbo (fig. 29), and the distinct myophragm (fig. 30).
31-5, Dorsal, ventral, posterior, anterior, and lateral views of SU 16600, a relatively young form.
(All figures x5-5.)

Palaeontology, Vol. 12

PLATE 91

SAVAGE, Howellella

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

481

The ventral valve has a concave, apsacline interarea with a width half the maximum
shell width and an apical angle of about 1 10°. This is bordered by low, poorly defined
beak ridges. An open, triangular delthyrium includes an angle of 30-40° and is bordered
by narrow deltidial plates which project normal to the interarea surface (PI. 91, fig. 10).
The ventral umbo is prominent and the beak incurved.

The dorsal valve has a small suberect beak and a low anacline interarea. An open
notothyrium includes an angle of about 140°.

The ventral sulcus and dorsal fold are both rounded in section and strongly defined.
They commence at the beaks and expand rapidly to occupy two-thirds of the shell width
at the anterior commissure. The lateral plications are broadly angular with straight
slopes and sharply rounded crests and troughs (PI. 91, fig. 11). Generally there are 3
plications each side of the sulcus and 2 each side of the fold. The finer ornament is
poorly preserved but numerous regularly spaced concentric growth-lines are visible.

text-fig. 4. Howellella nucula australis subsp. nov. Reconstruction of the dorsal
cardinalia based on several incomplete specimens. Approx, x 20.

Ventral interior. Well-developed slender dental lamellae extend one-quarter to one-
third of the valve length along the line of the plications bordering the median sulcus
(PI. 91, fig. 28). The lamellae diverge anteriorly and downwards, and have concave
anterior edges which drop sharply from the overhanging delthyrial margins to advance
along the valve floor (PI. 91, fig. 29). Small slender teeth have shallow crural fossettes
directed antero-medially (PI. 91, fig. 15). A very short median septum is present in the
posterior extremity of the umbonal cavity. The ventral muscle field, which is weakly
impressed in the material available, extends up to half the valve length and is divided
medially by a low myophragm (PI. 91, fig. 29).

Dorsal interior. Short hinge-plates extend laterally for two-fifths of the maximum shell
width and separated by a wide notothyrial cavity, (text-fig. 4). The cardinal process
is broad and strongly recurved posteriorly to protrude through the notothyrium (PI. 91,
fig. 6). Finer details of the cardinal process are seldom preserved but two specimens
have a bilobed process (text-fig. 4) and one has a trilobed process. Narrow sockets are
widely divergent at about 125°. They are variable in length and supported by the strongly
curved hinge-plates which arise from beneath the interarea. The inner socket ridges
are narrow with prominent projections distally which articulate with the crural fossettes
of the ventral valve (PI. 91, fig. 27). Large blade-like crural lamellae extend downwards
from the inner socket ridges but do not reach the valve floor. They meet the valve wall
posteriorly directly below the cardinal process (text-fig. 4). The lamellae are strongly

482

PALAEONTOLOGY, VOLUME 12

convergent downwards and divergent anteriorly, with concave anterior edges extending
one-third of the distance to the anterior margin (PI. 91, fig. 6). From the lower edges
of the crural lamellae arise crural bases from which long crura project anteriorly and then
curve sharply upwards to point postero-ventrally. Distally the crura are attached to the
laterally directed spires, each of which has 4 or 5 volutions. The faintly impressed dorsal
muscle-field is confined to the trace of the fold. It extends about one-third of the valve
length.

Measurements. The dimensions of 5 specimens are given

Length

below in mm.
Width

Thickness

SU 16601

Complete shell

2-5

3-3

2-7

SU 16602

Complete shell

3-4

4-6

30

SU 16603

Complete shell

3-7

4-7

3-5

SU 16604

Dorsal valve

5-7

9-3

—

SU 16606

Ventral valve

3-2

4-3

—

Ontogeny. From the

numerous specimens

in the collection a good ontogenetic series is

available. The main changes which occur with growth are the accentuation of the fold
and sulcus, the projection posteriorly of the ventral umbo, and the gradual incurving
of the ventral beak (PI. 91). As maturity approaches the thickness of the shell is increased
by the deposition of shell material along the lateral and posterior margins, and this is
responsible for the incurving of the beaks and the increasing prominence of the ventral
umbo. In the young internals examined the features differed little from the mature forms.

Discussion. Howellella nucula ( Barrande 1879), from the Ludlovian Kopanina Limestones
of Bohemia, is very close to the Manildra material. Externally the forms are identical,
with a similar fold and sulcus bordered by two or three gentle plications. Internally
the arrangement of the crural lamellae and dental lamellae is the same, and there is a
low median myophragm in both ventral valves (compare PI. 91, herein, with Havlicek
1959, figs. 41, 42). However, there is a median septum at the very apex of the umbonal
cavity in the Manildra form which is not present in the Bohemian species (Havlicek
1959, fig. 41). Though distinct, this median septum is very short and different from that
in Delthyris.

Other related Bohemian species include H. koneprusensis Havlicek, from the Kone-
prusy Limestones, which differs from the Manildra form in having an angular fold and
sulcus and more lateral plications. H. spuria (Barrande), from the Kopanina Beds, has a
broader fold and sulcus flanked by incipient lateral plications, and H. inchoans ( Barrande),
from the Lochkov Limestones, has numerous weak lateral plications.

H. laeviplicata Kozlowski, from the Borszczow beds of Podolia, is also close to the
Manildra species. Only a single ventral section is figured by Kozlowski (1929, fig. 60a),
and this does not show a myophragm. However, Kozlowski states in his description that
the internal features are essentially the same as in H. angustiplicata which he illustrates
with several serial sections (op. cit. , fig. 64). These show the crural lamellae suspended
from the hinge-plates for most of their length, as in the Manildra specimens. Material
with affinities both to the eastern European forms and those from New South Wales, is
found further east, in central Asia. The Siberian species H. khalfini Kulkov, 1963, from

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

483

the Lower Devonian Solovikha Limestone in the Altai Mountains, is less inflated than
the Australian form and does not possess advancing dental lamellae.

Of the species described from south-east Australia, H. scabra Philip, from the Cooper’s
Creek Formation at Tyers, Victoria, is less gibbous than the Manildra form. Both have
2 or 3 lateral plications each side of a strong median fold and sulcus, but whereas the
Victorian species has 2 very short dental lamellae, the lamellae of H. nucula australis
extend up to one-third of the valve length. Philip (1962, p. 222) states that the Tyers
material has extremely short crural bases but makes no mention of the crural lamellae,
possibly because the form of these structures is not apparent from slots visible in mould
material when species with ‘suspended’ crural lamellae are examined. H. textilis Talent,
from the Tabberabbera Formation, Victoria, is known only from deformed material
which is not easily compared with the silicified material from Manildra. Another Vic-
torian species, H. lirata Talent, differs from H. nucula australis in being wider and having
a lower fold and sulcus flanked by more numerous lateral plications.

Superfamily suessiacea Waagen 1883
Family cyrtinidae Frederiks 1912
Genus cyrtina Davidson 1858

Type species. Ca/ceo/a heteroclita Defrance, 1828, by the subsequent designation of Hall and Clarke,
1894.

Cyrtina praecedens Kozlowski 1 929
Plate 92

1929 Cyrtina praecedens Kozlowski; p. 207.

1954 Cyrtina praecedens Kozlowski; Nikiforova, p. 150.

Material. The collection consists of 472 silicified specimens of which 329 are complete, or almost com-
plete, with the valves united. Of the remainder 55 are dorsal valves and 88 are ventral valves. The
preservation is sufficiently delicate to show the external punctation.

Description. Exterior. The shell is small and hemipyramidal. In outline it is semicircular
with right-angular cardinal extremities and evenly rounded lateral and anterior margins.
The lateral profile is subtriangular with the maximum thickness at the posterior margin.

The ventral valve is hemipyramidal with a high umbonal region. The steep slopes
are slightly convex laterally and more strongly convex anteriorly. A prominent, suberect
beak projects just past the hinge line and the very high interarea is apsacline to catacline
with an apical angle of 85-95°. The interarea bears growth striations parallel to the
hinge line. A long, narrow delthyrium includes an angle of about 20° and is closed for
most of its height by a deltidium which is gently convex near the hinge-line but strongly
arched and hood-like about a round or oval apical foramen (PI. 92, fig. 28).

The dorsal valve is convex with a broad umbo and a suberect or nearly straight beak.
It has a very low, anacline interarea.

A dorsal fold and ventral sulcus are prominent. Both extend from the posterior margin
to the strongly parasulcate anterior commissure. The fold is elevated above the remainder
of the valve anteriorly and at the commissure it accommodates a semicircular tongue
from the ventral sulcus. Both valves have 3-5 pairs of strong, rounded plications which
arise along the cardinal margins (PI. 92, figs. 1, 2). The furrows bordering the fold, and

484

PALAEONTOLOGY, VOLUME 12

the plications bordering the sulcus, are particularly pronounced. Several strongly lamel-
lose growth-lines are usually present in mature specimens. The shell material is finely
punctate.

Ventral interior. Convergent dental lamellae project anteriorly from the delthyrial
margins but do not reach the valve floor. These lamellae meet to form a vertical spondy-
lium which extends anteriorly for about one-sixth of the valve length (PI. 92, figs. 35,
39). Above the spondylium the dendal lamellae become progressively reduced to form
stout, vertical pillars which border the delthyrial cavity and project as short rounded
teeth. Supporting the spondylium is a high median septum with a concave forward
edge. The septum protrudes into the spondylial cavity as a subvertical tichorhinum with
a hollow interior partly divided by a thin vertical plate, complete at the apex, but for
the most part consisting of two separate septa (PI. 92, figs. 35, 39). The closed apical
part of the tichorhinum is visible through the foramen (PI. 92, fig. 37). No ventral
muscle-scars have been observed in this material.

Dorsal interior. The dorsal interior has a broad cardinal process consisting of three
primary lobes and numerous secondary lobes (PI. 92, fig. 31). The sockets, which are
deep and triangular, are supported on strong, thickly developed hinge-plates (text-fig. 5).
Crural lamellae extend downwards from short, widely divergent inner socket ridges but
do not meet the valve floor; the lower parts form crural bases which diverge anteriorly
at 65-75°. Subparallel crura extend about one-quarter of the valve length and are joined
by a slender, V-shaped jugum with its pointed end directed anteriorly (PI. 92, fig. 27).
The spiralia are large and occupy most of the shell interior. Each is elongate and laterally

EXPLANATION OF PLATE 92

Figs. 1-44. Cyrtina praecedens Kozlowski. Mandagery Park Formation, Manildra. 1-25, Dorsal,
ventral, posterior, anterior, and lateral views of five specimens of progressively decreasing size.
1-5, Specimen SU 16628, a large mature form with successive growth thickenings at the valve margins
(fig. 5). The closed apical part of the tichorhinum and the horizontal striations on the ventral in-
terarea are also visible (fig. 3). 6-10, Specimen SU 16627, a mature form with the dorsal fold

distinctly elevated above the lateral plications (fig. 10). 11-15, Specimen SU 16626, a younger

stage with less pronounced plications. 16-20, Specimen SU 16625, a young form with only two
plications each side of the fold (fig. 16). 21-5, Specimen SU 16624, a very young form with gentle

plications and relatively rounded contours. 26, Antero-ventral view of dorsal valve SU 22669
showing the attachment of the crura to the crural lamellae. 27, Anterior view of broken specimen
SU 16634 showing the slender V-shaped jugum joining the subparallel crura. 28, Postero-lateral
view of specimen SU 16633 showing the strongly arched deltidium with a hood-like foramen.
29, 30, Ventral and antero-ventral views of dorsal valve SU 16636. 31, 32, Enlargements of figs.

29, 30 showing the trilobed cardinal process with its numerous secondary subdivisions (fig. 3 1 ), the deep
sockets partly overhung by the inner socket ridges (fig. 30), the widely divergent crural lamellae,
and the long narrow muscle-field divided by a low myophragm (fig. 30). 33, 34, Two dorsal

views of ventral valve SU 16629 showing the dental lamellae converging to form a spondylium
into which the high median septum protrudes as a hollow subvertical tichorhinum. The latter is
partly divided by a thin vertical plate which can be seen on both the posterior and anterior sides.
35-6, Dorsal and dorso-lateral views of ventral valve SU 16631. (The hollow nature of the median
septum within the spondylium is probably the result of incomplete silicification.) 37, Postero-dorsal
view of SU 16632 showing the partly exposed spires. 38, 39, Dorso-lateral and dorsal views of
ventral valve SU 16630. 40-44, Dorsal, ventral, posterior, anterior, and lateral views of SU 16635

showing unusually prominent growth lamellae. (Figs. 31, 32 x8, remaining figures all x4.)

Palaeontology, Vol. 12

PLATE 92

SAVAGE, Cyrtina

N. M. SAVAGE: SPIRIFERID BRACHIOPODS

485

directed with 5-6 volutions (PI. 92, fig. 37). The adductor muscle-field is long and narrow,
extending two-thirds of the distance to the anterior margin, and consisting of small
posterior adductors and larger, more elongate, anterior adductors. A low myophragm
divides the muscle field medially (PI. 92, fig. 32).

text-fig. 5. Cyrtina praecedens Kozlowksi. Reconstruction
of the dorsal cardinalia of the Manildra specimens based on
several incomplete specimens. Approx, x 35.

Measurements. The dimensions of 4 specimens are listed below in mm.

SU 16625

Complete shell

Length

2-7

Width

3-8

Thickness

2-2

SU 16626

Complete shell

3-2

4-5

2-8

SU 16627

Complete shell

4-0

5-7

40

SU 16628

Complete shell

4-5

6-5

5-2

Variation. There is some variation in the strength of the growth-lines and in a few
specimens these are particularly pronounced and produce a very uneven, lamellose
surface (PI. 92, figs. 40-4). In other specimens the growth-lines are more gentle, though
equally numerous (PI. 92, figs. 6-10). Differences in the form of the deltidium are com-
monly the result of breakage, but there is evidently some variation for a few specimens
have a strongly convex plate with a hooded foramen (PI. 92, figs. 28), while the majority
have a far less convex plate with a longer, non-hooded foramen (PI. 92, fig. 42).

Ontogeny. A good ontogenetic series is present for this species. There are few external
differences between stages apart from the increase in the number and height of the lateral
plications (PI. 92, fig. 1-25). In the gerontic stage successive growth lamellae are closely
spaced at the lateral and anterior margins and become more conspicuous (PI. 92, fig. 4),
than in younger specimens. Internally there is a thickening of the hinge-plates in older
specimens and a broadening of the cardinal process as the three primary lobes divide
and multiply.

Discussion. Although the Manildra specimens are smaller and have fewer plications
than those from Podolia described by Kozlowski, an attempt has been made to allow
for environmental stunting and to avoid the introduction of a new species name when
only size and related characteristics are distinctive.

The type species from Western Europe, Cyrtina heteroclita, is not well known and
its relationship to the Manildra material is difficult to assess. The form from Santa
Lucia identified as C. heteroclita by Oehlert (1901 ) is seen in his illustrations to have an
unbroken septum dividing the tichorhinum. If this is a constant characteristic it may

486

PALAEONTOLOGY, VOLUME 12

have taxonomic significance, for m specimens of C. praecedens from Podolia and from
Manildra the septum is not continuous across the tichorhinum. Another reported feature
of C. heteroclita which needs substantiation is the absence of an apical foramen (Koz-
lowski 1929, p. 210).

Of the other European species, C. teta Havlicek, from the late Emsian or early Eifelian
of Bohemia, resembles C. heteroclita externally although the interarea is considerably
longer. It is more angular than the Manildra form and has more prominent horizontal
striations across the interarea.

C. heteroclita gregale Talent, from Sandy’s Creek, Victoria, resembles the Manildra
form in shape and the number of plications but the growth lamellae are very weak or
absent in the material described. Another eastern Australian form, described by Dun
(1907) as the new species C. wellingtonensis, is also close to C. praecedens. It is larger than
the material from Manildra but possibly lived in a more favourable environment.

Acknowledgements. I wish to thank Dr. V. Havlicek and Mr. Galle of the Geological Survey of Czecho-
slovakia who generously made specimens available. I am also indebted to Professor H. B. Whittington
and Mr. A. G. Brighton for facilities at the Sedgwick Museum and access to specimens. Dr. N. P.
Kulkov of the Academy of Sciences, U.S.S.R., kindly sent specimens from the Altai Mountains.
Professor C. E. Marshall and Professor F. H. T. Rhodes provided facilities at University of Sydney,
and University College of Swansea, respectively. Professor A. J. Boucot and Dr. J. G. Johnson, of the
California Institute of Technology, kindly read the typescript and suggested improvements.

REFERENCES

barrande, j. 1879. Systeme Silurien du Centre de la Boheme. Pt. 1, Recherches paleontologiques
5, Brachiopodes , 226 pp., 153 pi. Prague, Paris.

conrad, t. a. 1842. Observations on the Silurian and Devonian systems of the U.S. with descriptions
of new organic remains. J. Acad. nat. Sci. Philad. 8, 228-80, pi. 12-17.
dun, w. s. 1907. Notes on Palaeozoic Brachiopoda and Pelecypoda from New South Wales. Rec.
geol. Surv. N.S. W. 8, (3), 269.

hall, j. and clarke, J. 1892-95. An Introduction to the Study of the Genera of Palaeozoic Brachio-
poda. Pt. I. Nat. Hist. N.Y. Palaeontology, 8, pt. 1, 1-367, pi. 1-20 (1892); pt. 2, 1-317 (1893),
319-94, pi. 21-84 (1895).

havlicek, v. 1956. Ramenonozci va pencil branickych a hlubocepskych z nejblizslho prazskeho okoli.

Sb. listred. Ust. geol. 22, 535-650, pi. 1-12 (English summary 651-65).

— — 1959. Spiriferidae v ceskem siluru a devonu (brachiopoda). Rozpr. ust fed. TJst. geol. 25, 219 pp.
28 pi. (English summary 221-75).

hill, d. 1942. Middle Palaeozoic rugose corals from the Wellington district, N.S.W. J. Proc. R. Soc.
N.S.W. 76, 182-9, pi. 5, 6.

and jones, o. a. 1940. The corals of the Garra Beds, Molong District, New South Wales. Ibid.

74, 175-208, pi. 2-8.

hisinger, w. 1826. Gotland geognostik beskrifting. K. Vet.-Akad. Hand!. Stockholm.

Johnson, J. G. 1965. Lower Devonian Stratigraphy and Correlation, Northern Simpson Park Range,
Nevada. Bull. Canadian Petrol, geol. 13, no. 3, 365-81.
kozlowski, r. 1929. Les brachiopodes Gotlandiens de la Podolie Polonaise. Palaeont. pol. 1, 254 pp.
12 pi.

kulkov, n. p. 1963. Brackhiopody Solovikhinskikh sloev Nizhnego Devona Gornogo Altaya. Akad.
Nauk SSSR , 131 pp, 9 pi.

Nikiforova, o. i. 1954. Stratigrafiya i brakhipody Siluriyskikh otlozheniy Podolii. Trudy Vsesojuz
geol. inst. Moskva. 218 pp.

oehlert, d. p. 1901. Fossiles Devoniens de Santa Lucia. Bull. Soc. geol. Fr. 1, 233-50.
packham, g. h. 1 960. Sedimentary history of part of the Tasman Geosyncline in South Eastern Australia.
21st. Int. geol. Congr. Copenhagen 1960, Regional paleography, 12, 74-83.

N. M. SAVAGE: SPIRIFERID BRACHIOPODS 487

philip, G. m. 1962. The palaeontology and stratigraphy of the Siluro-Devonian of the Tyers area,
Gippsland, Victoria. Proc. R. Soc. Viet. 75, 123-246, pi. 11-36.

pitrat, c. w. 1965. Suborder Spiriferidina, in moore, r. c. (ed).. Treatise on Invertebrate Paleontology,
Part H, Brachiopoda , H668-H727, Geol. Soc. Am. and Univ. Kansas Press.

savage, n. m. 1 968a. Planicardinia, a new septate dalmanellid brachiopod from the Lower Devonian
of New South Wales. Palaeontology, 11, 627-32, pi. 122.

19686. Australirhynchia, a new rhynchonellid brachiopod from the Lower Devonian of New South

Wales. Ibid. 11, 731-5, pi. 141.

1969. The Geology of the Manildra District, New South Wales. J. Proc. R. Soc. N.S. W. (in press).

strusz, d. l. 1965a. A note on the Stratigraphy of the Devonian Garra Beds of N.S.W. Ibid. 98,
85-90.

1965. Disphyllidae and Phacellophyllidae from the Devonian Garra Formation of New

South Wales. Palaeontology, 8, 518-71, 72-8.

1966. Spongophyllidae from the Devonian Garra Formation, New South Wales. Ibid. 9, 544-

98, pi. 85-96.

1967. Chlamydophyllum, Iowaphyllum, and Sinospongophvl/um (Rugosa) from the Devonian of

New South Wales. Ibid. 10, 426-35, pi. 67.

termier, h. and termier, g. 1949. Essai sur 1’evolution des Spiriferides. Notes Mem. Serv. Mines Carte
geol. Maroc. no. 74, 85-112.

vandercammen, a. 1956. Revision des Ambocoeliinae de la Belgique. Bull. Inst. r. Sci. nat. Belg. 32,
1-51, pi. 1, 2.

N. M. SAVAGE

Department of Geology
University of Natal
Durban, Natal

Typescript received 23 December 1968 South Africa

A CRETACEOUS ECHINOID WITH FALSE TEETH

by PORTER M. KIER

Abstract. The teeth described in 1911 in a specimen of Conulus subrotundus Mantell from theTuronian Middle
Chalk are from a Recent echinoid. No lantern was present in adults in Conulus or probably in other members of
the families Conulidae Lambert or Galeritidae Gray. The structures previously thought to be lantern support
structures (auricles) are considered to be related in function to the large buccal plates. Instead of being degenera-
ting structures as previously thought, they increase in size in later species.

Although echinoid workers generally have not become very aroused in their con-
troversies, two subjects have caused considerable heat: the question whether Both-
riocidaris was an echinoid (now resolved in the affirmative), and whether or not Conulus
had teeth. Conulus is one of the better-known and more ‘popular’ echinoid genera in
Europe because of its abundant occurrence in the Chalk. It was assumed that it was
toothed because it is an holectypoid, and most holectypoids have teeth. Forbes (1850,
p. 3) described and figured what he considered to be teeth and jaws in Conulus and
subsequent authors (see Hawkins, 1911, p. 70 for a complete history) accepted his
opinion until Duncan (1884, p. 11) in a paper considered to be dogmatic by those op-
posed to his views, disagreed with the previous workers and contended that the objects
they thought to be jaws were imaginary, or merely grooves made by a tool in the soft
matrix within the peristome. After this strong rebuttal the proponents for a lantern
retreated and most subsequent workers until 1911 accepted that it was lanternless.
However, Hawkins (1911, p. 71) found a specimen of Conulus subrotundus Mantell,
in the British Museum with four teeth protruding from its peristome. He was unable to
find any jaws. Since this time it has been assumed by all echinoid workers that Conulus
and the rest of the genera of the families Conulidae Lambert and Galeritidae Gray
had jaws and teeth when adults.

As part of a study of the lantern in echinoids, I was particularly anxious to see a
lantern in Conulus and dissected hundreds of specimens of C. albogalerus Leske. This
is a relatively easy task with an air abrasive machine because of the soft chalk matrix.
However, no fragments of a lantern or teeth were found in any of them. Inasmuch as
the peristome is very small in diameter, bits of the lantern would have been expected
to have been retained in the test. Furthermore, I dissected a specimen of C. albogalerus
in which all the buccal plates were still preserved in place but there were no fragments of
a lantern. These plates have been found on only a handful of the thousands of specimens
of this species that have been collected. Obviously the slightest movement of the speci-
men after death caused these plates to become separated, and if a lantern had been
present in this specimen it would be expected that parts of it would still be there.
Hawkins also dissected hundreds of C. albogalerus and never found any lantern frag-
ments. But the teeth in the specimen of C. subrotundus described by Hawkins were
positive evidence that could not be ignored. It could not be assumed that four teeth
were washed into a specimen all with their tips extending outward. But if it had teeth,
why were no pyramids preserved? Hawkins noted that it would not be possible for the

(Palaeontology, Vol. 12, Part 3, 1969, pp. 488-493, pis. 93, 94.]

P. M. KIER: A CRETACEOUS ECHINOID WITH FALSE TEETH 489

pyramids to slip out through the small peristome and have the teeth remain. He sug-
gested that perhaps the pyramids were noncalcified — a conclusion with which he was
not satisfied but could suggest no other alternative. Restudy of this specimen has re-
vealed the solution to this dilemma — the teeth do not belong to the specimen. Some
person excavated a cavity inside the peristome and inserted four Recent teeth mixed
with some cement into the cavity.

The following evidence indicates that the teeth did not belong to the echinoid:

1. The matrix around the teeth is much softer than the matrix in the rest of the test.
This difference was readily apparent when the air abrasive machine was used. The
machine had very little effect on the area away from the peristome but one blast of
abrasive in the area around the teeth removed considerable matrix. As is well known
to anyone who has worked with chalk fossils, the matrix in Middle Chalk specimens is
commonly quite hard as opposed to Upper Chalk specimens. The matrix in this specimen
is typical of the Middle Chalk except in the cavity where the teeth lie. Here it is not only
very soft and crumbly but also much coarser (PI. 93, fig. 5) in texture. Dr. Maurice
Black, an authority on chalk, examined this material and concluded that it was not
chalk. Only a few coccoliths were visible and he suggested that these had probably come
from the adjacent chalk matrix. He surmised that this material around the teeth was
probably some type of cement (perhaps dental). A. G. Brighton, curator of the Sedgwick
Museum, Cambridge, reports (personal communication, 1968) that there have been many
chalk ‘fossils’ faked by individuals anxious to sell specimens to museums, or interested
collectors. Commonly, the hoaxer mixes up a matrix of crushed chalk or lime, and some
type of cement and inserts into it a Recent specimen and then offers it for sale as a
‘perfectly preserved Cretaceous fossil with color markings’.

After hearing about this forgery in Conulus, Peter J. Moulds of Queen Mary College,
London, examined some specimens of Chalk echinoids which had been puzzling him
and discovered that at least two of them are forgeries. According to his letter to me
(1968): ‘one block of Chalk with several spines enclosed had an entirely different test
added later. This test had been sawed in half in order to fit to the block! I suppose the
main reason for these forgeries was to increase the interest and thus the value (financial
that is — many of the museum specimens have their original price on them).’

2. The quality of the preservation of the teeth indicates that they are from a Recent
echinoid and not fossil. All the teeth have a glistening, porcellanous sheen (PI. 94, fig. 1)
which T have never seen to this extent in a fossil tooth. Although a slight sheen may
rarely be preserved on an extremely well-preserved fossil tooth, it is never as pronounced
as on these teeth. Furthermore, the open meshwork of the microstructure of the tooth
is not permineralized as it would be in a fossil (pi. 93, fig. 4). The open interstices in
the tooth are normally filled with secondary calcite in a fossil tooth but in this specimen
they are not. Furthermore, the upper part of the teeth are soft and fibrous with the
asbestos-like structure found in a Recent tooth but never in a fossil.

The teeth are too small for a carbon-14 analysis, but Dr. Kenneth Towe pointed out
that Weber and Raup (1968, p. 42) have shown that skeletal magnesium is lost early in
diagenesis and that Recent echinoids therefore have a higher magnesium carbonate
content than fossil ones. Dr. Towe suggested that if these teeth were Recent, they should
contain a larger amount of MgCO:J than the rest of the fossil. He analyzed (using X-ray
diffraction) a portion of one of the teeth, part of the test of a Conulus subrotundus ,

490

PALAEONTOLOGY, VOLUME 12

and for comparison purposes a Recent tooth, and a tooth known for certain to have
come from a Chalk species, Phymosoma koenigi (Mantell). The fragment of the test of
C. subrotundus and the tooth from P. konigi contained no MgC03, whereas the tooth of
the Recent echinoid contained 6-8 mol per cent MgCC)3 and the tooth from the Conulus
contained 3-4 mol per cent. The fact that the Comdus tooth contained MgC03 and the
specimen from which it was supposed to have come contained none indicates that the
tooth is not from the Comdus.

3. The fact that all the teeth are broken and that all these broken ends are on the
ends of the tooth in the matrix must arouse considerable suspicion as to their authenticity.
It is very doubtful that any natural forces could break the inner ends of all four teeth
and still permit their outer ends to protrude unbroken from the echinoid test. Echinoid
teeth are quite strong and any force which broke all of them would surely disassociate
them enough so they would not all remain with their tips still protruding out the peri-
stome. Probably the hoaxer, in order to avoid drilling a much deeper hole into the test,
simply broke part of each tooth and inserted the broken ends into the hole.

4. The teeth themselves are unlike any found before in an irregular echinoid. Their
keels are far too narrow and sharp. Hawkins (1911, p. 72) stated that the teeth were very
like those found in Camerogalerus cylindricus (Lamarck), but the tooth of Cam. cylindricus
has a much broader keel, tapering from the edge of the keel to the sides of the tooth,
whereas the teeth in the Comdus have the sides of the keel parallel to each other. Al-
though, Durham and Melville (1957, text-fig. 1b) show a narrow sharp keel in Holectypus
r/e/?re.sms(Leske)Ihave made further preparations of the specimen they figured and have
found that the tooth has a broad keel very much like that in Pygaster as described by
Melville (1961). I know of no irregular echinoid tooth resembling the teeth attributed
to Comdus.

5. When Hawkins first saw the specimen he noted that someone had enlarged the
peristome by cutting. Presumably, he thought that this enlarging was done in an effort
to expose the teeth, but probably the hoaxer was unable to fit all the teeth in the small

EXPLANATION OF PLATE 93

Figs. 1-3. View of the interior region around the peristome in three species of Comdus showing the
thickened basicoronal plates which have been considered to be auricles. The pictures are arranged
stratigraphically with the earliest species, Comdus castanea (Brongniart) from the Cenomanian at
the base (fig. 3), the Turanian C. subrotundus Mantell in the middle (fig. 2) and the latest, the
Senonian C. albogalerus, at the top (fig. 1 ). Note that the structures formerly considered to be auricles
are more pronounced in C. albogalerus than in the older species contradicting the assumption that
these structures are degenerating lantern supports. 1, Comdus albogalerus Leske, Upper Chalk,
Gravesend, Sedgwick Museum B. 3623, Kent, x 8. 2, Comdus subrotundus Mantell, Middle Chalk,

Orbirhynchia cuvieri zone, Flitchin, Herts., Sedgwick Museum B. 408, X 10. 3, Conulus castanea

(Brongniart), Bed 13 Meyer, Beer Head, Devon, Sedgwick Museum B. 7577, X 13.

Fig. 4. Section through tooth considered to be Recent but found in specimen of Conulus subrotundus
figured in fig. 6. Note the microstructure which is normally visible on a Recent tooth but not on a
fossil, x37.

Figs. 5, 6. Comdus subrotundus Mantell. Specimen which H. L. Hawkins found in the British Museum
(Natural History) with four teeth protruding from the peristome. He cut the specimen in half and
excavated the area around the peristome but found no fragments of a lantern. Note the coarser
matrix around the teeth. Label for specimen, B.M. E 10743, only states Upper Chalk which is
presumably an error because this species is known only in the Middle Chalk. Fig. 5, X 6 5 ; fig. 6, x 2.

Palaeontology, Vol. 12

PLATE 93

KIER, Cretaceous echinoid with false teeth

P. M. KIER: A CRETACEOUS ECHINOID WITH FALSE TEETH 491

peristome and just widened it enough to accommodate them — which might also explain
why he inserted only four teeth.

6. The presence of teeth but absence of any of the numerous parts of the jaws is
nearly impossible to explain. The peristome of Conulus is so small in diameter that the
jaws could hardly have slipped out around the teeth without the teeth slipping out also.
Commonly, the teeth are the first to slip out of the test after the echinoid dies. They are
connected to the jaws by far less tissue (they must be able to move down the dental
slide as the echinoid grows) than the pyramids are to each other. I have found very few
teeth in comparison to the number of jaws during my excavations of fossil echinoids,
and I have never found teeth without there also being part of the jaws.

The only direct evidence of a lantern in Conulus was the presence of the teeth described
above. Now that they are shown to be fraudulent we must examine again the problem
as to whether Conulus, and for that matter any of the members of the Conulidae or
Galeritidae, had a lantern. Recent workers (Hawkins 1911, 1917, 1934, Mortensen
1948, p. 43, Wagner and Durham 1966, p. 455) have considered that the thickened
structures (PI. 93, figs. 1-3) in the interambulacra at the edge of the peristome were
auricles (lantern support structures). Although Hawkins’s illustration (1917, pi. 28,
fig. 1, reproduced in the Treatise, Wagner and Durham 1966, fig. 331, 4 c) does depict a
structure strongly resembling auricles, this figure is highly stylized and gives a misleading
impression of the structure. In this figure the auricle-like features are exaggerated.

Although the thickened basicoronal plates do resemble auricles or apophyses, they
differ from them in an important character. Auricles or apophyses consist of processes
which rise upward from the basicoronal ambulacral or interambulacral plates. These
tabs may be thick or thin but invariably they rise far above the general level of the
basicoronal plates. No such tabs are present in Conulus. Although minute knobs are
present on the edge of the thickened basicoronal plates in large specimens of some species
of Conulus, they are absent from most species and are far too small to be considered
as auricles.

Many workers including Hawkins (1911, p. 72) have considered that the ‘auricles’
in Conulus were degenerate structures and that their lack of strong development resulted
from the fact that the lantern and its supporting structures were gradually being lost
through time. If this were the case it would be expected that these ‘auricles’ would be
less pronounced in succeeding species, but just the opposite is the case. This thickening
of the basicoronal plates becomes more pronounced in later species. The earliest
Conulus in which I have been able to expose the interior is C. castanea (Brongniart)
from the Cenomanian. The basicoronal plates are slightly thickened (PI. 93, fig. 3)
and two slight depressions are present in each of these thickened interambulacra. In
the Turonian C. subrotundus Mantell the interambulacral plates are more thickened
(PI. 93, fig. 2) the paired depressions deeper and the angle of their faces greater. Finally
in the Senonian C. albogalerus Leske all these features (PI. 93, fig. 1) are even more
pronounced. Therefore, this thickening cannot be considered a degenerating character.

The position and character of these structures suggest that they are related to the
function of the ten plates around the peristome which are considered to be buccal plates.
These plates are interpreted as buccal plates rather than basicoronal plates because they
are not attached to the rest of the test by normal sutural tissue as is indicated by their
absence in most specimens (they have only been found in C. albogalerus but presumably

492

PALAEONTOLOGY, VOLUME 12

were present in the other species of Conulus ) and their loose connection on those few
specimens where they are found. Furthermore, the plates which precede them have the
arrangement and number characteristic of basicoronal plates, with a single plate in
each interambulacrum, and the ambulacral plates and their pores arranged according
to Loven’s law. Finally, these basicoronal plates have their edges curved inward with their
tubercles facing the peristomial opening.

The plates are very thick and large in C. a/bogalerus (PI. 94, figs. 3-5) and could have
their origin in the large buccal plates normally found in a regular echinoid or be derived
from the secondary buccal plates found in irregulars. The primary buccal plates of
regular echinoids have large buccal tubefeet whereas tubefeet are absent from the second-
ary buccal plates of irregular echinoids. The absence of pores in the Conulus buccal
plates indicates that they probably are derived from these secondary plates. Although
some authors (Mortensen 1948, p. 38) thought pores were present in these plates because
of the presence of pits on the interior of the plates. These pits (PI. 94, fig. 3) are shallow
and do not penetrate to the exterior.

Hawkins suggested that the ‘auricles’ may have been slots into which the buccal
plates were retracted. However, the exterior of the buccal plates has tubercles, pre-
sumably for spines or pedicellariae, which probably would have prohibited the sliding
of these plates back into the peristomial opening. Furthermore, the buccal plates curve
interiorly over the edge of the peristome almost in a joint which would make impossible
their sliding back over the edge of the peristome.

It is apparent, however, that there is some relation between the ‘auricles’ and the
buccal plates. The ten depressions in the thickened interambulacral plates are directly
behind the ten buccal plates. I agree with Hawkins that the pits on the interior of the
buccal plates are for the insertion of muscles and suspect that the deep depressions of
the ‘auricles’ were where these muscles were attached to the test. Perhaps the buccal
plates functioned as teeth, pushing food into the gut. The thickening of the basicoronal
plates would be necessary not only to provide a properly angled face for attachment
of these muscles, but also would strengthen these plates so that a large stress could be
exerted on them when the muscles contracted.

SUMMARY

There is no direct evidence that Conulus or any member of the Conulidae or Galeritidae
had a lantern when adult. Excavation of hundreds of specimens of Conulus has revealed
no fragments of a lantern. The teeth described by Hawkins are shown to be Recent

EXPLANATION OF PLATE 94

Figs. 1-2. Views of two of the teeth, believed to be Recent, from specimen of Conulus subrotundus
Mantell figured on Plate 93, figs. 5, 6. Note the glistening porcellanous sheen and the fibrous micro-
structure which are typical of a Recent tooth but never so well preserved on a fossil, X 16.

Figs. 3-5. Conulus albogalerus Leske. 3, Interior view of area around peristome showing the buccal
plates and the deep depressions in the thickened basicoronal interambulacral plates which may have
served for the attachment of muscles leading to buccal plates. The small pits on the buccal plates do
not pass through the plates and were probably for the insertion of these muscles, X 10. 4, View

of same specimen less enlarged, X 2. 5, Exterior view of same specimen showing buccal plates and

the small tubercles for the attachment of spines or pedicellariae, X 10. BM E 33079, Senonian,
Micraster coranguinum zone, Northfleet, England.

Palaeontology, Vol. 12

PLATE 94

KIER, Cretaceous echinoid with false teeth

P. M. K1ER: A CRETACEOUS ECHINOID WITH FALSE TEETH

493

and the fragment that he considered to be part of a pyramid, he later suggested was a
piece of a pelecypod. The structures considered to be auricles were probably related
functionally to the large massive buccal plates which occur immediately oral to them.
The obliquity of the peristomial opening in many species of Conulus and its relatives
is further evidence that no lantern was present in the adult. All echinoids having a
lantern have a symmetrically shaped peristome.

Acknowledgements. This study was carried out under a Guggenheim Fellowship at the Sedgwick
Museum, Cambridge. I thank Professor H. B. Whittington for making the facilities of the Museum
available to me, and A. G. Brighton and Maurice Black for their advice. Dr. R. P. S. Jefferies, Curator
of Fossil Echinoderms at the British Museum (Natural History), very kindly lent me specimens of
Conulus. J. Wyatt Durham and Richard E. Grant reviewed the manuscript and made several very
useful suggestions. Kenneth Towe not only pointed out the value of making a magnesium carbonate
analyses of the tooth, but also made the analyses.

REFERENCES

duncan, p. m. 1884. On Galerites albogalerus, Lamarck, syn. Echinoconus conicus, Breynius. Geol.
Mag. 21, 10-18.

Durham, j. w. and melville, r. v. 1957. A classification of echinoids. J. Paleont. 31, 242-72, text-figs.
1-9.

forbes, E. 1850. British fossils. Mem. Geol. Surv. 3, 3 pp., pi. 8.

hawkins, h. l. 1911. Teeth and buccal structures in Conulus. Geol. Mag. 48, 70-4, pi. 3.

1917. Morphological studies on the Echinoidea Holectypoida and their allies. 6, The buccal

armature of Conulus albogalerus, Leske. Geol. Mag. 54, 433-41, pi. 28.

1934. The lantern and girdle of some recent and fossil Echinoidea. Phil. Trans. Roy. Soc. Lond.

223B, 617-49, pi. 68-70, 26 text-figs.

melville, r. v. 1961. Dentition and relationships of the echinoid genus Pygaster J. L. R. Agassiz,
1836. Palaeontology, 4, 243-6, pi. 28-9.

mortensen, th. 1948. A Monograph of the Echinoidea. 4, 1. Holectypoida, Cassiduloida, 363 pp.,
14 pk, 326 text-figs., Copenhagen.

wagner, c. d. and Durham, j. w. 1966. Holectypoids, in r. c. moore (ed.), Treatise on Invertebrate
Paleontology, Part U, Ecliinodermata 3, (2) 440-50, figs. 329-34, Geol. Soc. Am. and Kansas Univ.
Press.

weber, j. n. and raup, d. m. 1968. Comparison of C13/C12 and 018/016 in the skeletal calcite of Recent
and fossil echinoids. J. Paleont. 42, 37-50, text-figs. 1-5.

porter m. kier
Department of Paleobiology
U.S. National Museum
Washington, D.C. 20560

Typescript received 6 December 1968

CERATOCYSTIS PERNERI JAEKEL-A MIDDLE
CAMBRIAN CHORDATE WITH ECH I NODERM

AFFINITIES

by R. P. S. JEFFERIES

Abstract. Ceratocystis perneri Jaekel 1901 is the oldest known member of the Order Cornuta Jaekel 1901,
and of the Subphylum Calcichordata Jefferies 1967; as such it is not an echinoderm, though having echinoderm
affinities, but is interpreted as the oldest known member of the Phylum Chordata.

Its anatomy has been studied in detail, and new observations have also been made on Cothurnocystis americana
Ubaghs, Cothurnocystis primaeva Thoral, Cothurnocystis elizae Bather, and Mitrocystites mitra Barrande. These
studies throw light on the origin of the chordate heart and pericardium, reproductive system and acustico-
lateralis system. They also suggest a basic similarity between the anatomy of Ceratocystis perneri and that of
a pterobranch hemichordate resting on its right side.

It is suggested that a population of pterobranch hemichordates, that took to resting on their right sides and
acquired calcite skeletons, gave rise both to the echinoderms and to the chordates.

Ceratocystis perneri Jaekel, from the Middle Cambrian of Bohemia, is interpreted as
the oldest and most primitive chordate known. It is also the oldest and most primitive
member of the subphylum Calcichordata Jefferies 1967, of the class Stylophora, of
the order Cornuta and of the family Ceratocystidae. Unlike any known descendants,
C. perneri had a hydropore, and this, with other features, connects it more closely than
later calcichordates with the echinoderms and makes it possible to suggest how echino-
derms and chordates are related to each other and to the phylum Hemichordata. Study
of C. perneri also throws light on the origin of the chordate heart, reproductive system,
acustico-lateralis system, and hypophyseal complex.

C. perneri was first described and figured by Barrande (1887, pi. 2, figs. 17-21)
as ‘plaquettes isolees’. Pompeckj (1896) described and figured C. perneri under the
names Trochocystites! (p. 503, pi. 13, figs. 9, 11) and Mitrocystites (p. 504, pi. 14, figs. 1,2).
Jaekel (1901) named the species and cursorily described and figured it. Bather (1913)
discovered the gill slits, whose nature was first recognized by Gislen (1930). Ubaghs
(1967, summarized in 1968) has given a well-illustrated account of the species and inter-
preted it in a way completely different from that here adopted (see also Ubaghs 1961,
1963, 1968, and Jefferies 1967, p. 205; 1968, pp. 263, 276, 289 ff. , 335). He has also
selected a lectotype and given precise references to previous work (Ubaghs 1967, p. 2),
except for that of Barrande and Pompeckj.

I have reconstructed C. perneri from natural moulds and rubber casts. As with my
previous work, several projections were drawn simultaneously on a drawing-board.
Reconstruction was difficult as the thecal plates have always been badly dislocated.
I have made complementary observations on two species which largely bridge the gap
between C. perneri and the previously studied Cothurnocystis elizae. These two species
are Cothurnocystis americana Ubaghs and Cothurnocystis primaeva Thoral (text-fig. 1 .)

(Palaeontology, Vol. 12, Part 3, 1969, pp. 494-535, pis. 95-98.]

R. P. S. JEFFERIES: CERATOCYSTIS PERNERl JAEKEL

495

Systematic position. Phylum Chordata; Subphylum Calcichordata Jefferies 1967; class
Stylophora Gill and Caster 1960; order Cornuta Jaekel 1901; family Ceratocystidae
Jaekel 1901 ; genus ceratocystis Jaekel 1901; species Ceratocystis perneri Jaekel 1901.

The family Ceratocystidae, as here understood, includes the family Cothurnocystidae
Bather 1913 and comprises Ceratocystis , Cothurnocystis, Phyl/ocystis, and Nevadaecystis
(whose only known species I prefer to regard as the earliest Cothurnocystis known).
It excludes the Scotiaecystidae Caster and Ubaghs 1968. This usage differs from Ubaghs
(1963, 1967, 1968) and Jefferies (1967, 1968).

IO>ars

Occurrence. All the specimens studied come from near Skryje, Bohemia. Ubaghs
(1967, p. 14) has given details of localities and horizon. The specimens occur in numbers
scattered over bedding planes in a greenish greywacke. They are remarkably complete,
and presumably died by burial. This agrees with the fact that the matrix, being a grey-
wacke, was presumably deposited from a turbidity current. Associated trilobites prove
marine, and suggest shallow-water conditions.

Material. The material of Ceratocystis perneri examined numbered about 100 specimens and is pre-
served in: Narodnl Muzeum, Prague (including lectotype 22123/7, 1924); Naturhistorisches Museum,
Vienna (specimens W1-W27); British Museum (Natural History), London (E16071-4); Geologisch-
Palaontologisches Museum der Humboldt Universitat, Berlin (Ca9, 26, 30-33); Geologisch-Palaeont-
ologisches Institut der Universitat, Greifswald, German Democratic Republic; High School of Mines,
Ostrava, Czechoslovakia; Ustredni Ustav Geologicky, Prague; Geologisch-Palaontologisches Institut
der Universitat, Freiberg-i-Br., Federal German Republic; Smithsonian Institution, Washington
(61503, 33324); Mineralogisk Museet, Copenhagen.

C 6685 K k

496

PALAEONTOLOGY, VOLUME 12

The two known specimens of Cothurnocystis primaeva Thoral were lent by the Institut de Geologie,
Universite de Lyon (1879/508), and the Institut de Geologie, Universite de Montpellier (holotype).
The only known specimen of Cothurnocystis (= Nevadaeeystis) americana Ubaghs (USNM 143237)
was lent by the Smithsonian Institution, Washington. Additional material of Cothurnocystis elizae
Bather and Scotiaecystis curvata was lent by the Hunterian Museum, Glasgow.

GENERAL SHAPE, THECAL PLATE NOTATION, AND
THECAL PLATE HOMOLOGIES

Like all calcichordates, Ceratocystis perneri consists of a theca and a stem (text-fig. 2).
The theca is boot-shaped. On the anterior face, right oral, left oral, and left appendages
can be recognized. These are equivalent to like-named appendages in Cothurnocystis
elizae and Coth. primaeva. However, the right oral appendage of Ceratocystis perneri
was fixed immoveably at the base, instead of being held by a hinge as in Coth. primaeva
and Coth. elizae. The posterior right and left angles of C. perneri correspond to what
Bather (1913, p. 399) called the ‘ball of the foot’ and the ‘heel’ of Coth. elizae.

The ventral surface of the theca of C. perneri is approximately flat but a ventral spike
(S3R) is always developed on plate M4RV, and sometimes there are one or two ventral
bosses (S1R, S2R) on M2R (PI. 95, fig. 1 ; PI. 96, fig. 7). Also the left appendage projects
somewhat downwards so that, as far as its form in C. perneri is concerned, it could
almost equally well be called a ventral spike. It is here named the left appendage because
it is homologous with the better-developed left appendages of later forms. The ventral
surface curves somewhat dorsally in two regions on each side of the stem. As Ubaghs
noticed (1967, p. 12), the stereom mesh of the ventral plates of C. perneri \s denser where
it would have touched the sea floor than elsewhere. It is particularly dense where it forms
the ventral spikes, and the ventral surfaces of the left appendage and of the right and
left oral appendages (text-fig. 17a; PI. 95, fig. 1). These particularly dense areas represent
portions of the ventral surface which would have been forced slightly into the sea floor
by the weight of the animal.

The dorsal surface of the theca (text-fig. 2a) is raised into three keels which meet
on the plate CM (PI. 95, fig. 2). The stereom forming these keels is denser than that
surrounding the keels so that each forms a sort of girder. These girders are most deve-
loped where the plates that they cross are thinnest, i.e. roughly in inverse proportion to
the distance from the meeting-point of the three keels.

A keel also exists dorsal to the left appendage, on plates M3LV and M3LD (text-fig. 2a;
PI. 95, fig. 2). In addition a rather sharp angle (peripheral keel) separates ventral from
lateral surfaces in most places.

The theca of C. perneri , unlike that of later cornutes, was almost entirely covered by
large plates. There was, however, a semicircular oral integument, (or in, in text-fig. 2c;
PI. 95, fig. 1) just ventral to and posterior to the mouth, and small, flexible flaps, con-
taining platelets, probably covered the gill slits (text-fig. 2a, d; fpl in PI. 98, fig. 1).
The large plates of the theca can be divided into: dorsal marginals (MD), ventral
marginals (Mv), marginals that are neither dorsal nor ventral (M), centro-dorsals
(C), right and left oral appendages (roap, loap) and infra-branchials (IB). The suffixes
used in the plate notation are: A = anterior; P = posterior; L = left; R = right;
D = dorsal; v = ventral; 4_5 = position in sequence starting just anterior to the stem.
The ascription of the same suffix number to dorsal and ventral marginals (e.g. M4RV,

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL 497

M4RD) is somewhat arbitrary. A plate is not necessarily homologous with a plate having
the same notation in some other species of cornute.

The relation of the present notation for C. perneri to that of Ubaghs (1967) is as

follows; MPV = M0; MPD = Am; M1RV = M4; M1RD = Ag; M2R = M2; M3R = I4;

^4rv = h43 M4RD = S4; roap = M4; M5R = I4; MAD = M6, S8.

M4Lv = M4; M1ld = Ad; M2L = M2; M3LD = S4; M4LV = M4; M4LD = S6;

loap = M5; M5L = I2.

CA = S5; CM = S2; CPL = S3; CPR = S4. IB4, IB2, IB3, not shown by Ubaghs.

I3 of Ubaghs is probably only a portion of M5R. M,; and S8 of Ubaghs are ventral
and dorsal aspects of the same plate (MAD). Ubaghs did not observe the oral integument
or the infra-branchial plates.

The homology of the thecal plates of C. perneri with those of the various species of
Cothurnocystis and Seotiaecystis can be established fairly certainly in most cases (text-
fig. 3). Ubaghs ( 1 967, p. 4) seems to have been over-cautious in this matter. One important
trend in the evolution of Ceratocystidae was the development of the bellows system of
pumping water through the gill slits. This had much influence on the thecal plating,
since it caused the thecal wall to become more and more flexible, and it must be borne in
mind in examining text-fig. 3. The bellows system was rudimentary in C. perneri ;
Coth. americana had a flexible roof to the theca and a rigid floor crossed by a thickened
strut. Coth. primaeva , Coth. elizae, and Seotiaecystis curvata had a flexible roof and a
flexible floor, crossed by a rigid strut.

The plate homologies call for the following additional comments:

1. The strut of Coth. tfme/7Ctf/7tf(strintext-fig.4;P1.98,fig.2), which has not previously
been described, corresponds in position to, and must have been homologous with, the
struts of later species. Comparison with Coth. americana allows the anterior and
posterior strut plates to be recognised in C. perneri (M1RV, M4LV).

2. The left appendage, right and left oral appendages and the ‘heel’ and ‘ball of the
foot’ provide good landmarks in all species, except that the right oral appendage is
absent in S. curvata.

3. Apart from Coth. americana , where the relevant anatomy cannot be worked out,
the plates just anterior to the stem (i.e. MPV, M1RV, M1RD, MPD, M1LD, M1LV) can be
shown to have the same connections with each other in all species of cornutes and mitra-
tes except that MPV is known only in C. perneri while Coth. elizae , S. curvata and the
mitrates have lost MPD. The relative sizes of these plates, however, vary greatly from
species to species.

4. M5LD of Coth. primaeva is homologised with M3LD of C. perneri, rather than with
M4LD, because (i) M4LD of C. perneri is obviously homologous with a plate C1A in the
dorsal integument of Coth. americana, and (ii) the shape of M4LV in the incomplete,
only known specimen of Coth. americana , suggests the existence of a plate M4LD (text-
fig. 3c, ? pos M4LD in text-fig. 4; PI. 98, fig. 2) in the complete animal that would be
intermediate in position between M3LD of C. perneri and M5LD of Coth. primaeva.

5. The plate M4RV of C. perneri is homologised with M3R of Coth. primaeva, Coth.
americana, and Coth. elizae, and with M4R of S. curvata, because a ventral spike is
present, except in Coth. elizae and some specimens of S. curvata.

6. The Montpellier specimen (holotype) of Coth. primaeva, on which text-fig. 3 a,b

498

PALAEONTOLOGY, VOLUME 12

largely based, lacks M3L, which exists in the only other known specimen of the species,
preserved at Lyon.

7. IB1; 2, 3 (text-fig. 2a, d; ib in PI. 98, fig. 1) of C. perneri correspond to some of the
posterior U-plates of Coth. amerieana, Coth. primaeva, and Coth. elizae.

The stem of C. perneri , like that of other calcichordates, was divided into anterior,
medial and posterior parts and ended abruptly. Ubaghs himself stressed that there is no
evidence for the pointed plate which he showed in his reconstruction at the posterior end
of the stem (1967, text-fig. 1, p. 13.)

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

499

C roaP

text-fig. 2. Ceratocystis perneri
Jaekel. Reconstruction of external
features, a, dorsal; b, right lateral;
c, ventral; d, posterior aspects. For
plate and spike notation, see text,
ag = accessory gaps; an = anus;
as = anterior stem; bs = branchial
slit ; dg = dorsal groove (median
eye); g = gonopore; h = hydro-
pore; loap = left oral appendage;
lpvp = lateral portion of a ventral
anterior stem plate; m = mouth;
ng = narrow groove (lateral line); or
in = oral integument; ps = posterior
stem; roap = right oral appendage;
stc = stylocone ; vo = ventral ossicle ;
vpr = ventral process of a ventral
anterior stem plate.

D bs ^pl Mpd bq M

IB3 /aq n9 / J m; an 9

3 M ^ KA M 'IRV

1 'ILD M|Lv 1 'PV

GAPS IN THE THECAL SKELETON

There are many places where the plates of C. perneri are penetrated or separated by
definite gaps, as distinct from ordinary sutures. Some of these gaps represent true
thecal openings which, in life, would carry organs or parts of organs through the thecal
wall. Such thecal openings would either connect external sense organs to the nervous
system, or connect internal cavities with the ambient sea-water. By contrast, other gaps
in the thecal skeleton merely represent places where the thecal wall existed but was not
calcified. These can be called uncalcified wall spaces. They would be filled in life with
muscle or connective tissue.

500

PALAEONTOLOGY, VOLUME 12

22

text-fig. 3. Plate homologies among Cornuta. a, b, Cothurnocystis primaeva Thoral, ventral and dorsal aspects; c, Cothurnocystis americana
Ubaghs, dorsal aspect; d, e, Ceratocystis perneri Jaekel, ventral and dorsal aspects; f, g, Scotiaecystis curvata (Bather), ventral and dorsal
aspects; h, i, Cothurnocystis elizae Bather, ventral and dorsal aspects. Homologous plates have the same stipple, lap = left appendage;

loap = left oral appendage; rap = right appendage; roap = right oral appendage.

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

501

Gill slits and the rudiments of the bellows system

A first complex of gaps in the thecal skeleton is disposed between the plates in the left
dorsal part of the theca (ag, bs in text-fig. 2a, d). This complex consists of two types of
gap. The first type is represented by seven large holes (bs). Three of these are between
CPL and IBj, IB2 and IB3, while four are between CPL and M2L. The second type of
gap, corresponding to what are here called accessory gaps (ag), is represented by holes
that are variable in form, number, size, and position. They are usually much smaller
than the seven holes bs. A first group of such accessory gaps is arranged anterior to the
gaps bs along the sutures M3LD/M3LV, CpL/M3LV and CPL/M2L. A more posterior group
of accessory gaps is arranged near the three most median of the gaps bs, mainly around
the infra-branchial plates, along the sutures IB3/M2L, IB3/M1LD, IB2/M1LD, 1B1/M1LD,
IBi/MpD, and MPD/M1LD.

The distinction between the gaps bs and the accessory gaps is emphasized by Plate 96,
fig. 2, showing the dorsal margin of M2L and M3LV, and by Plate 97, figs. 4, 5, showing
dorsal and ventral margins of infra-branchial plates.

The gaps bs must represent branchial slits for the following reasons:

1. They closely resemble in position and number the 7 (or perhaps 8) gill slits of
Coth. americana (bs in text-fig. 4; PI. 98, fig. 2); each of these, in turn, is almost identical
in structure to the previously studied gill slits of Coth. elizae (Jefferies 1967, p. 167;
1968, p. 253).

2. If the gaps bs are gill slits, the infra-branchial plates could well be homologous
with some of the posterior U-plates of Coth. americana (pu in text-fig. 4), Coth. elizae,
and Coth. primaeva.

3. Upper and lower margins of the gaps bs are not identical. Each upper margin
is sharply defined and truncated by a facet (faf in PI. 98, fig. 1) while the lower is rounded.
This suggests, by analogy with Coth. elizae, that the upper margin bore a soft flap,
attached to the facet, and this flap closed against the lower margin, so forming an outlet
valve. One specimen has platelets inside one of the gaps bs (fpl in PI. 98, fig. 1). In life
these could have been imbedded in a flap, much as in Coth. elizae.

If the gaps bs were gill slits, the accessory gaps represent uncalcified wall spaces.
Thus the posterior accessory gaps correspond closely in position to the gaps in Coth.
elizae and Coth. americana between the plates in the strip of dorsal integument posterior
to the gill slits. The anterior accessory gaps correspond, though less closely, to the gaps
between the dorsal integument plates anterior to the gill slits in Coth. elizae and Coth.
americana. Further, many of the integument plates of Coth. americana are star-shaped,
and contact their star-shaped neighbours by the points of the stars (text-fig. 4; PI. 98,
fig. 2). They could well have evolved from polygonal plates whose sutures were affected
by the enlargement of accessory gaps of the type seen in C. perneri.

As regards function, the accessory gaps of C. perneri probably contained muscles
which could slightly depress the roof of the theca, so forcing a sudden jet of water out
through the gill slits. This seems likely, since the gaps between the integument plates
of Coth. elizae to which the accessory gaps correspond, seem to have contained muscle
that pumped water out through the gill slits (Jefferies 1967, p. 168; 1968, p. 258) and
the same is true of Coth. americana. There was a shallow depression in the inner face of

502

PALAEONTOLOGY, VOLUME 12

the theca of C. perneri surrounding the posterior group of accessory gaps and correspond-
ing to a raised portion of the internal cast (iml in text-fig. 9b; see also text-fig. 10, and
PI. 95, fig. 9; PI. 97, figs. 5, 8). This depression could have contained a patch of muscle
that cooperated with the muscle of the accessory gaps in pumping.

internal surface of ventral plates

matrix

text-fig. 4. Cothurnocystis americana Ubaghs, dorsal aspect of only known specimen and holotype
(USNM 143237 from the Lower Tremadoc of Nevada), au = anterior u-plate; bs = branchial slit;
f flap; dpi = dorsal plates of stem base region (precise morphology not discernible); grg = gono-
rectal groove; in be = integument over buccal cavity; in ph = integument over pharynx; lap = left
appendage; ?pos M4LD = presumed position of plate M4LD; pu = posterior u-plate; str = strut.

Slight contractions of the right part of the thecal cavity, which would supplement
those of the left part of the theca, may also have occurred. This is suggested by the form
of the edges of some of the plates near the right margin. Thus the edge of M4RD where
it abuts against M4RV is cylindrical, rather than plane (rm in PI. 95, fig. 1 ; PI. 96, figs. 3,
6). Again the edge of M5R where it touches roap and of M3R where it touches M2R
are distinctly cylindrical. This suggests that a hinging movement was possible of the thecal
plates about a hinge-line, corresponding to the sutures M5R/roap, M4RV/M4RD, and M3R/
M2R, near the right-hand margin of the theca (hinge in text-fig. 3d, e). This presumed

R. P. S. JEFFERIES: CERATOCYSTIS PERNER1 JAEKEL 503

hinge-line runs along the middle of an elongate depression on the inner face of the
skeleton, corresponding to a raised portion of the internal cast (imr in text-fig. 9a, c;
PI. 95, fig. 2; PI. 96, figs. 1, 4). This depression presumably carried a sheet of muscle
like the depression internal to the posterior accessory gaps. It is likely that the action
of this sheet of muscle was mainly to raise the neighbouring plates of the thecal floor
by rotation about the presumed hinge line.

C. perneri therefore, could probably pump water through the pharynx by flexing the
thecal wall, just as later Ceratocystidae could, i.e. the bellows system of pumping water
through buccal cavity and pharynx had already begun to evolve. This method of
pumping, however, must have been much less efficient than in later forms, since the
volume of the thecal cavity when contracted could have been only a few per cent less
than the expanded volume. Indeed, in C. perneri flexing of the thecal walls could not have
been the only method of pumping water. Some other method must have been much more
important. This could have involved cilia on the inner surface of pharynx and buccal
cavity, or as seems less likely, some type of muscular pumping which has left no trace
in the skeleton.

The ability to flex the thecal wall in C. perneri was developed along sutures which,
with few exceptions, correspond to the dorsal margins of the frame in later forms (text-
fig. 3). This is obvious on the left side of the theca and also true on the right, for M4RV
of C. perneri, which is ventral to the right hinge-line, corresponds to the frame plate
M3R of Coth. aniericana, Coth. primaeva, and Coth. elizae, and M4R of S. curvata. This
situation is consistent with the fact that the thecal roof of Coth. americana was flexible,
but the floor was not. The muscles round the presumed right hinge line must have changed
their mode of action at an early stage from lifting the neighbouring floor, as in C. perneri,
to depressing the side wall, which soon became part of the roof.

The dorsal and peripheral keels of C. perneri would have stiffened the roof and floor
of the theca. They may owe their existence to the rudimentary bellows mechanism for,
without them, contraction of the bellows muscles would have forced water towards the
middle of the theca and made the floor, and especially the roof, belly out, instead of
expelling water through the gill slits. As the musculature of the roof gradually spread
inwards in the course of evolution, the stiffening provided by the dorsal keels became
both unnecessary and disadvantageous, and they were eventually lost. The dorsal
keels are still obvious in Coth. americana , however (text-fig. 4; PI. 98, fig. 2), and some
sign of them even occurs in Coth. elizae, though probably only as a functionless and
variable vestige (crestal plates, Jefferies 1968, p. 259). The peripheral keels of C. perneri,
on the other hand, became elaborated to form the thecal frame of Coth. americana and
later species and became better adapted to a stiffening function, the need for which in-
creased when the roof and floor of the theca became flexible.

The presence of 7 gill slits in C. perneri is of interest, since this is probably the primi-
tive number for cornutes and therefore for chordates in general. That 7 is the primitive
number for cornutes is supported by Table 1 which shows that all the early species of
cornutes with the unspecialized elizae type of slit (i.e. all except P. crassimarginata,
S. curvata, and Coth. elizae itself, which is a late form) have a similar number. The
lowest value among these species is perhaps 5, probably 7 ; the highest value is 9.

Certain other groups of chordates with a similar number of pairs of gill slits, may have
inherited this number from an unknown Upper Cambrian cornute with 7 slits on the

504

PALAEONTOLOGY, VOLUME 12

left side of the theca, by way of an Upper Cambrian mitrate with 7 gill slits on each side.
Examples of such groups are the Cyathaspididae, which include some of the earliest
known agnathous fishes and which show evidence of 6 or 7 pairs of gill slits (Denison
1964, p. 345), and gnathostomatous fishes which never have more than 8 pairs of gill
slits, including the spiracles (Bertin in Grasse 1958, p. 1306).

table 1. The number of gill slits in Cornuta

Name

Age

Type of Slit

Number

Ceratocystis perneri Jaekel

M. Cambrian

As here described

7

Cothurnocystis americana Ubaghs

L. Tremadoc

elizae

7 (or 8)

Cothurnocystis primaeva Thoral

U. Tremadoc

At least 5,

or

probably a

L. Arenig

,,

few more.

Cothurnocystis ubaghsi Chauvel

,,

,,

7 (or 8)

Phyllocystis btayaci Thoral

,,

,,

9

Phyllocystis crassimarg inala Thoral

,,

Specialized

13

Cothurnocystis elizae Bather

Ashgill

elizae

16

Scotiaecystis curvata (Bather)

Specialized

40

Thecal openings on the right side of the theca

There are 4 thecal openings on the right side of the theca of C. perneri (m, h, g and an
in text-fig. 2c, d; PI. 95, fig. 1 ; PI. 96, figs. 6, 7, 8, 10; PI. 98, fig. 5). Two of these open-
ings (g and an) are usually fused together to varying extents (cf. PI. 96, figs. 7, 8, 10).
The identification of these four holes follows from a comparison with rhombiferan

EXPLANATION OF PLATE 95

The lengths of the scales are in mm. E = British Museum (Nat. Hist.); GPMB = Geologisch-

Palaontologisches Museum der Humboldt Universitat, Berlin; NM = Narodni Muzeum, Prague;

USNM = U.S. National Museum, Washington; W = Natur-historisches Museum, Vienna.

Figs. 1-7, 9. Ceratocystis perneri. 1, Ventral aspect of W 13; latex impression to show oral integu-
ment or in; M2l is affected by a fault in the latex; an = anus; g gonopore; h = hydropore;
lap = left appendage; ng = narrow groove (lateral line); rm = rounded margin of M3RD i Sir, S3R —
first and third right ventral spikes. 2, Reconstruction of dorsal aspect (W 8) made by glueing latex
impressions of plates together. 3, Natural mould, dorsal aspect (NM 33723, same individual as
Plate 97, fig. 1); br = brain; gbc = groove between buccal cavity and pharynx; pbc = pit between
buccal cavity, pharynx and anterior coelom. 4, Reconstruction of ventral aspect (E 16074)
made in same way as fig. 2. 5, Latex impression of part of an individual (W 10), to show inside of

plate M4RD; cbcr ridge corresponding to cleft in natural mould, i.e. between buccal cavity and
anterior coelom; imr = limit of groove on internal cast housing the right internal muscle. 6,
Anterior aspect of natural mould representing the part of the brain in contact with MiRD (specimen
W 16); iskc = intraskeletal cones of right side; rf = right face of brain. 7, Anterior aspect rep-
resenting part of brain in contact with MPD (specimen W 16); magnification as fig. 6; adf = antero-
dorsal face; dll, dir = left and right dorsal lobes; dp dorsal process. 9, Natural mould of
inside of MPD, (specimen W 26); ag = accessory gaps; iml = limit of left internal muscle.

Figs. 8, 10. Cothurnocystis elizae. 8, Natural mould representing posterior coelom and adjacent
structures in contact with M1L+RV; cf. text-fig. 6; postero-ventral aspect, with specimen lying on its
back (specimen E 28644); gd gonoduct; grg = infilling of gonorectal groove; mini = left median
line nerve; pco = posterior coelom; pbr = right pyriform body; r = rectum. 10, Natural mould
of posterior surface of M1Ld of E 28658 representing terminal portion of rectum (r) and gonoduct
(gd) and part of antero-dorsal surface of posterior coelom (pco); cf. text-fig. 6.

Palaeontology, Vol. 12

PLATE 95

-T«f

rP M ■

cbcr

&bi 'MkC

Imlnl^pbr'

;[

JEFFERIES, Ceratocystis perneri, Cothurnocystis elizae

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

505

and diploporitan cystoids. Jaekel (1899) established the existence in those groups of
four holes in the theca, apart from pores connected with respiration (text-fig. 5). These
four holes were arranged roughly in a plane that included the oral end of the theca and
the base of the stem or, in stemless forms, the point of attachment. In sequence they were
the mouth, at the oral end of the theca, the hydropore, the gonopore, and the anus.
The holes m, h, g, and an of C. perneri are also distributed roughly in a plane which
includes the oral (anterior) end of the theca and the stem base, but which, in this case,
is parallel to the ventral surface and to the sea bottom. This comparison suggests that
m of C. perneri is the mouth, h the hydropore, g the gonopore, and an the anus.

text-fig. 5. Oral surface and thecal openings of the diploporitan
cystoid Glyptosphaerites leuchtenbergi (Volborth) redrawn after
Jaekel 1899. an = anus; fg = food groove; g gonopore;
h = madreporite (corresponding to hydropore of other cystoids
and of Ceratocystis)-, m = mouth.

The identification of the mouth (m) in C. perneri as a large opening at the anterior
end of the body, agrees with the situation in other calcichordates, and with chordates
in general. It agrees in particular with Coth. elizae (Jefferies 1967, p. 168; 1968, p. 255).
The account of the mouth region of C. perneri given here differs greatly, even in terms
of pure morphology, from that of Ubaghs (1967, p. 1 1 , under anus), who did not observe
the oral integument.

The identification of the holes g and an of C. perneri as gonopore and anus calls for
partial restudy of the anatomy of Coth. elizae. In this species I have previously described
an opening, which I called the ‘anus’, just left of the stem, where it would be in the
outwash from the most median gill slits. I described the ‘anus’ as connected by a ‘rectal
groove’ in the skeleton that ran under the posterior coelom to the anterior coelom in
the right-hand, posterior region of the theca. Re-examination, however, reveals that the
distal, vertical part of the ‘rectal groove’, between the ‘anus’ and the posterior coelom,
in fact consists of two grooves — a larger and a smaller (rg and gdg in text-fig. 6, cor-
responding to r and gd in PI. 95, figs. 8, 10). These two grooves must represent two
contiguous tubes, one wide and one narrow, opening into the ‘anus’. The narrower
tube, as judged by its groove, ran ventral and posterior to the wider tube across the
posterior coelom, but ran left of the wider tube where both climbed upwards after

506

PALAEONTOLOGY, VOLUME 12

leaving the posterior coelom, so that finally it lay dorsal and anterior to the wider tube
near the opening. The situation in Coth. primaeva (text-fig. 7; PI. 98, fig. 6) is essentially
the same as in Coth. elizae.

As already mentioned, the two tubes opening into the ‘anus’ of Coth. elizae are
connected by the ‘rectal’ groove to a point just right of the stem. This point corresponds

text-fig. 6. Cothurnocystis elizae Bather, internal anatomy of the theca (cf. Jefferies 1968, fig. 3).
a, Antero-dorsal aspect of posterior part of theca, with the dorsal integument removed and M1L+RD
lifted upwards, b. Posterior aspect of M1L+RD. c, Left lateral aspect of animal, lying on sea-floor,
to show lines of sight used in a and b. Only the portion of the theca posterior to X-X is shown in a.
1 , 2, 3, 4 = points where pharyngo-visceral line approaches the dorsal side of the theca ; bpc = portions
of skeleton in contact with posterior coelom; gdg = gonoductal groove; gmln = groove for median
line nerves; grg = gonorectal groove; htd = heart depression ; pbd = depressions for pyriform bodies;
pvl = pharyngo-visceral line; rg = rectal groove, sp = striations of pharynx; str = strut ;vint = ven-
tral integument.

to the position of gonopore and anus in C. perneri (cf. text-fig. 8b-f with 8a-e). This
suggests that the two tubes of Coth. elizae represent the rectum and the gonoduct, which
had migrated in evolution to just left of the stem, so as to be in the outwash from the
gill slits. Conversely, the position of these tubes in Coth. elizae, such that their products
could be washed away, confirms that they were outlet ducts, and supports the identi-
fication of gonopore and anus in C. perneri. It is likely that the narrower tube in Coth.
elizae was the gonoduct, and the wider one the rectum. Evidently the ‘rectal’ groove is
better called the gonorectal groove, and the ‘anus’ should be called the gonopore-anus.

The distal part of the rectum of a mitrate (text-fig. 8c-g) had the same basic course as
in Coth. elizae on the one hand (text-fig. 8b-f), or a tunicate tadpole on the other (Jefferies
1967, p. 181 ; 1968, pp. 287 ff., p. 317). In mitrates as in tunicate tadpoles it opened into
a left atrium. By comparison with Coth. elizae, the gonoduct of mitrates probably also

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

507

opened into the left atrium (text-fig. 8g) with the terminal portion of the gonoduct left
of the rectum.

In enterogonous tunicates (text-fig. 8h) the terminal portion of the gonoduct is right
of the rectum. Comparison of text-fig. 8g and h, however, shows that the mutual rela-
tions of rectum and gonoduct are nevertheless similar in mitrates and enterogonous
tunicates. In the adult tunicate the two tubes are merely turned through 150° compared
with the mitrate condition. The point x in the tadpole diagram (text-fig. 8d) indicates

text-fig. 7. Cothurnocystis primaeva Thoral. a, camera lucida drawing of the ventral
aspect of a portion of the natural mould of the Montpellier specimen (holotype) (cf. PI.

98, fig. 6). b. Ventral aspect of theca; stipple indicates portion of anatomy shown in a.
br = brain; fM1RD natural mould of the facet on plate M1RV which contacted plate
M1RD; gd gonoduct; grg = infilling of gonorectal groove; iskc = intraskeletal cones,
corresponding to those in Ceratocystis perneri; mini and rnlnr = left and right median-
line nerves; pbl and pbr = left and right pyriform bodies; r = rectum.

where the gonoduct will enter the left atrium later in ontogeny, and underlines the
comparison. There is no evidence whether cornutes and mitrates were hermaphrodite
like tunicates. What is here called the gonoduct may represent the male or the female
duct in different individuals or, as in tunicates, one inside the other.

In Coth. amerieana the position of the gonopore-anus cannot be established. A
gonorectal groove can be seen right of the stem (grg in text-fig. 4) as in other forms of
Cothurnocystis, but cannot be followed more distally. The migration of gonopore and
anus had therefore already begun, but may not yet have finished.

The leftward twist of gut and gonoducts was thus acquired by cornutes because gill
slits existed only on the left side. It was retained in mitrates and tunicates, where gill
slits existed on both sides, because it was no particular disadvantage.

The identification of the hydropore (h) in C. perneri, based on comparison with
cystoids, is confirmed by its absence in all cornutes later than C. perneri. Thus the cor-
nutes, in this respect as in others, evolved towards a more usual chordate condition.

508

PALAEONTOLOGY, VOLUME 12

□ ant. coelom [S3 post, coelom WIM pharynx
□atria and buccal cavity

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

509

The presence of a hydropore in C. perneri, as discussed below, has important implica-
tions for the internal anatomy and throws light on the connections between chordates,
echinoderms and hemichordates.

Thecal openings related to the nervous system

Near the anterior end of the stem are two openings, the narrow groove and the dorsal
groove (ng and dg in text-fig. 2a c, d), that are not represented in Coth. elizae (ng in
PI. 96, figs. 4, 10; dg in PI. 96 figs. 1, 9; PI. 98, fig. 1).

The narrow groove is excavated in the ventral surface of M1LD. Ubaghs (1967, pp.
8 ff.) described it as “?hydropore\ It is here regarded as the first rudiments of the acustico-
lateralis system and, since it is widely open at the surface, would have functioned as a
lateral line. It is reminiscent of the lateral line of mitrates (narrow groove (ng) in Jefferies
1968, pp. 283, 307, 314, 322; lateral line (11) in Jefferies 1967, pp. 178, etc.), and Ubaghs
homologized it with this feature. It is, however, on the left of the theca, whereas the
lateral line of mitrates is on the right. It will be further discussed under the nervous
system.

The dorsal groove is excavated in MPD, just anterior to the mid-line of the stem.
Ubaghs named it ‘encoche mediane’ and suggested (1967, p. 1 1) that it might be connected
with an internal mouth that he supposed to exist in this region. It is interpreted here as
having carried an upward extension of the optical part of the brain, which was presumably
light-sensitive. The dorsal groove certainly existed also in the cornutes Phyllocystis
b/ayaci and P. crassimarginata. It may perhaps have existed in Coth. primaeva , which
certainly had a plate MPD that could have carried it, but this plate is not well displayed
in either of the two known specimens. The dorsal groove is also further discussed under
the nervous system.

THE CHAMBERS OF THE THECA

Four thecal chambers can be recognized in C. perneri on the basis of superficial
internal anatomy, the positions of thecal openings, and the fundamental resemblance to
Coth. elizae. These four chambers are the buccal cavity, pharynx, anterior coelom,
and posterior coelom. Text-figs. 9a-c are reconstructions of the internal cast of C. perneri ,
combining details from many different specimens. Text-figs. 10a-c are an interpretation
of these reconstructions. The sculpture of the internal cast is complex and includes:

(1) Growth-lines, parallel to the edges of plates and not shown in the text-figures.

(2) Growth traces, extending inwards perpendicular to plate edges from notches (e.g.
gill slits gtbs in text-fig. 9b, c) in the plate edges (cf. PI. 97, fig. 8). (3) Indications of
muscles, associated with rudimentary flexibility of the theca, discussed above, and dis-
tributed in two areas, i.e. just ventral to the more median gill slits (iml in text-fig. 9b)

text-fig. 8. Homologies between various calcichordates and a living tunicate, a, e, Ceratocystis
perneri (cornute) ; b, f, Cothurnocystis elizae (cornute) ; c. g, Mitrocystella incipiens (mitrate) ; d, tadpole
of the tunicate Ciona; h, adult of Ciona (dorsal aspect ) a, b, c, and d compare the anatomy of the four
forms in dorsal aspect, e, f, and G show the relations of gonoduct, rectum, gill slits, and atria, at
atrium; ato = atrial opening; bs = branchial slit; g = gonoduct; h = hydropore; ht = heart;
lat = left atrium; Ip = left pharynx; m = mouth; ph = pharynx; r = rectum; rat = right atrium;
rp = right pharynx; st = stomach; x = the point where, in the developing tunicate, the gonoduct

later comes to enter the atrium.

510

PALAEONTOLOGY, VOLUME 12

and near the right border of the theca (imr in text-fig. 9a-c). (4) Intercameral features
and indications of internal organs other than thecal chambers.

The buccal cavity was situated much as in Coth. elizae, occupying the ‘ankle’ part
of the theca. Its posterior border is indicated on the internal cast: (1) By a groove
(gbc in text-fig. 9c; PI. 95, fig. 3; PI. 96, fig. 3). (2) By a pit (pbc in text-fig. 9c; PI. 95,
fig. 3 ; PI. 96, fig. 6) in the floor of the groove. (3) By a cleft (cbcr in text-fig. 9a, c; PI. 95,
fig. 5; PI. 96, fig. 1) which appears to have divided the right-hand strip of ‘bellows’
muscle (imr) into two parts. (4) By a cleft on the left (cbcl in text-fig. 9a; PI. 97, fig. 10).
On general grounds the clefts and the grooves would indicate where the skeleton had
filled the gap between two chambers, and the pit would indicate where it had filled the
gap between three chambers (in this case, buccal cavity, anterior coelom, and pharynx).

The anterior coelom must also have been situated as in Coth. elizae, i.e. mainly in
the posterior, right-hand part of the theca. It is from this region that gonopore, hydropore,
and anus emerged, indicating that the corresponding organs, and the chamber that
housed them, must have lain here. The boundary between posterior coelom and buccal
cavity is indicated : (1 ) By the part of the groove gbc to the right of the pit pbc. (2) By the
cleft cbcr. The junction of anterior coelom, pharynx, and buccal cavity is indicated by

EXPLANATION OF PLATE 96

The lengths of the scales are in mm.

Figs. 1-10. Ceratocystis perneri. 1, Latex impression in dorsal aspect of W 25 to show internal
features of M4EV, he. imr = limit of groove for right internal muscle ; cbcr = division between buccal
cavity and posterior coelom; dg = dorsal groove in plate MPD; imr = margin of groove for right
internal muscle on plate M4EV. 2, Latex impression in dorsal aspect (W19) to show distinction
between accessory gaps (ag) and branchial slits (bs.) 3, Latex impression in ventral aspect (Ostrava

specimen) to show features on internal surface of plate CA; gacpd = ridge corresponding to groove
on internal cast between pharynx and anterior coelom; gbc = ridge filling groove on internal cast
between buccal cavity and pharynx; rm = rounded margin of M4ED; stc = stylocone. 4, Latex
impression in ventral aspect of Ostrava specimen (different individual to fig. 3) to show internal
surface of M4ED and right oral appendage with limits of groove for right internal muscle (imr);
gan gonopore and anus; ng = narrow groove. 5, Latex impression of stem in dorsal aspect
(W 9); note how dorsal plates (dpi), though dislocated after death, have nonetheless remained articu-
lated to each other; aif = anterior imbrication facet of dorsal plate; bos = boss of ventral ossicle;
bs = branchial slit; DS = dorsal plate of anterior stem; lg = lateral groove; mg = median groove;
stc = stylocone; VS = ventral plate of stem. 6, Latex cast in ventral aspect of Ostrava specimen
(different individual to figs. 3 and 4) to show features of inside of CA with process (pbc) corresponding
to the pit on natural mould between buccal cavity, pharynx, and anterior coelom; gan = gonopore-
anus; h = hydropore; rm = rounded margin of M4ED. 7, Reconstruction of right posterior region of
theca in ventral aspect (W22), made by gluing latex impressions of plates together; an = anus;
g = gonopore; S1E, S2E = first and second right, ventral spikes. 8, Natural mould in ventral aspect
(GPMB Ca 33); an = anus;br = brain; g = gonopore; gacpv = groove on ventral surface between
anterior coelom and pharynx; gpcac = groove between posterior coelom and anterior coelom;
gppc = groove between pharynx and posterior coelom; pbl = left pyriform body. 9, Latex mould
in dorsal aspect (W15), to show anterior stem, with dorsal and ventral stem plates (DS, VS); dg =
dorsal groove ; gan = gonopore-anus ; gpcac = ridge corresponding to groove on internal cast between
posterior and anterior coelom; gppc = ridge corresponding to groove on internal cast between
pharynx and posterior coelom. 10, Latex mould in ventral aspect of posterior part of theca and
part of stem (NM 22123/7 1924 lectotype). Note narrow groove (ng) and the gonopore (g) and anus
(an) separated by a minute ridge, and lateral and ventral parts of a single ventral anterior stem plate
(lpvp, vpvp); h = hydropore; stc = stylocone.

Palaeontology, Vol. 12

PLATE 96

JEFFERIES, Ceratocystis perneri

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

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PALAEONTOLOGY, VOLUME 12

CD

text-fig. 10. The chambers of the theca of Ceratocystis perneri — an interpretation of the reconstructed internal cast (cf. text-fig. 9).

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

513

the pit pbc. The junction of anterior coelom and pharynx is indicated: (1) By the groove
gacpd (text-fig. 9c; PI. 96, fig. 3) which meets the groove gbc at the pit pbc. (2) By the
groove gacpv (text-fig. 9a; PI. 96, fig. 8) on the ventral surface, internal to M1RV. (3) By
the grooves gacpld and gacplv (text-fig. 9a, b; PI. 97, figs. 8, 9) on the left side of the inter-
nal cast, beneath the gill slits, which indicate an extension of the anterior coelom into
the left part of the theca, as in Coth. elizae. The boundary between anterior and posterior
coeloms is indicated on the right by the groove gpcac (text-fig. 9a; PI. 96, figs. 8, 9).
To the left it presumably ran left of the left pyriform body (pbl in text-fig. 9a, b; PI. 96,
fig. 8). A groove (gio in text-fig. 9a, b; PI. 96, fig. 1 ; PI. 98, fig. 5a) just beneath the gono-
pore and anus, and beneath and around the hydropore, may indicate the contact with
the skeleton of some visceral organ inside the anterior coelom.

The pharynx would have connected with both buccal cavity and gill slits, and so would
have been disposed as in Coth. elizae. The indications on the internal cast of its bound-
aries with buccal cavity and anterior coelom have been mentioned already. The
boundary between pharynx and posterior coelom is shown by the groove gppc (text-
fig. 9a; PI. 96, figs. 6, 9; PI. 97, fig. 2; PI. 98, fig. 3).

The posterior coelom probably occupied a small volume just in front of the stem.
The indications of its boundaries on the internal cast have already been mentioned.

ORGANS INSIDE THE ANTERIOR COELOM

The thecal openings from the anterior coelom are the anus, the gonopore, and the
hydropore. They indicate that the anterior coelom contained: (1) the rectum and the
rest of the alimentary canal behind the pharynx; (2) the gonad or gonads; and (3)
an axial complex, like that of echinoderms. The presence of an axial complex calls for
discussion.

Living echinoderms which compare with C. perneri in having an axial complex that
opens directly outwards by a hydropore or madreporite, comprise the echinoids, asteroids,
and ophiuroids. In all such forms the axial complex has basically the same structure
(Fedotov 1924, Smith 1940). This can be illustrated from an echinoid (text-fig. 1 1) where
the following parts exist: (1) An axial organ divided into the main portion (Millott and
Vevers 1968) and the head process. The main portion of the axial organ consists of a
close-packed mass of canals, some of which are haemal. (2) An axial sinus (lumen of
axial organ of Millott and Vevers 1968) which expands upwards into a madreporic am-
pulla beneath the madreporite. (3) The stone canal connecting the water vascular system
with the madreporic ampulla. (4) The dorsal sac which is a coelomic chamber that has
become invaginated on one side to contain the head process of the axial organ. (5) The
genital strand which runs from the axial organ to the gonads and is surrounded by (6) the
genital sinus.

The functions of the different parts of the axial complex are various. (1) The stone
canal, opening as it does by way of the madreporic ampulla and madreporite to the
outside, serves to keep the fluid of the water vascular system at the same hydrostatic
pressure as the ambient sea water (Fechter 1965). (2) The main portion of the axial
organ has three functions: (a) it is concerned in the degeneration of amoebocytes
(Millott 1966, Millott and Vevers 1968) and presumably expels their degeneration pro-
ducts by way of axial sinus, madreporic ampulla, and madreporite; ( b ) it is certainly
secretory (Millott and Vevers 1968) and possibly acts as an endocrine gland (Millott

514

PALAEONTOLOGY, VOLUME 12

1967, p. 63); (c) it helps to circulate fluid in the haemal system by means of contractile
vessels that it contains (Boolootian and Campbell 1964). (3) The head process and dorsal
sac are particularly concerned with pumping fluid through the haemal system. The head
process is in fact known to be pulsatile in all the living classes where it exists. Thus, in
echinoids, Boolootian and Campbell (1964) showed that it had the structure and action
of a two-chambered heart and pulsation was also observed by Narasimhamurti (1931)
and Prouho (1887, p. 331). In ophiuroids pulsation of the head process was recorded

by Narasimhamurti (1933, p. 79) and Gemmill
(1919), and in asteroids by Narasimhamurti
(1931) and Gemmill (1919) among others.

The head process and dorsal sac of echino-
derms are homologous with the heart and peri-
cardium of hemichordates, which they resemble
in structure, function and embryological origin
(Fedotov 1924, p. 298, Narasimhamurti 1931,
Gemmill 1914).

The head process and dorsal sac of C. perneri,
by comparison with living echinoderms, would
have lain near the aboral end of the axial
complex, i.e. in the anterior coelom near the
hydropore (ht in text-fig. 8a). In C. perneri the
superficial internal anatomy does not show
their exact location. In Coth. elizae and Coth.
primaeva, on the other hand, the position of the
dorsal sac is probably indicated by a bulge in
the internal cast (ht in PI. 98, fig. 4) ventral to a
culmination (4 in PI. 98, fig. 4) in the pharyngo-visceral line. This bulge corresponds to
a depression in the internal face of the skeleton (htd in text-fig. 6), and is similar in
position to the hydropore of C. perneri, since both lie near the median edge of plate
M2R (the ‘heel’ plate) which is homologous in the two species. The bulge is unlikely to
represent the position of an optic nerve (contrast Jefferies 1967, PI. 168; 1968, p. 251)
for it is too tumid and differs in shape from the space ventral to point 1, on the
left of the theca (PI. 98, fig. 4; text-fig. 6) which could well have carried such
a nerve.

Lying in such a position the dorsal sac and the presumably pulsatile head process of
C. perneri , Coth. elizae, and Coth. primaeva could well be homologous with the peri-
cardium and heart of other chordates. They would be situated (ht in text-fig. 8a, b) like
these features in other chordates, in the main visceral cavity, in the general region of
the gonads and post-pharyngeal gut. Berrill (1955, p. 112) was therefore mistaken in
asserting that, despite their morphological resemblance, the heart and pericardium of
chordates could not be homologous with those of hemichordates, because located in
different parts of the body. In the evolution of mitrates (text-fig. 8c) the pouching out
of the right pharyngeal chamber (rp) would separate the dorsal sac and head process
(henceforth called pericardium and heart) from the right wall of the theca and push
them towards the right side of the gut. Their position would thus become very similar
to the pericardium and heart of a tunicate larva (text-fig. 8d).

head process

of axial organ .rnadreporite

a^eV

dorsal sac

genital

sinus-

genital

strand

main portion
of axial organ

madreporic
ampulla

stone canal
axial sinus

text-fig. 1 1 . The axial complex of an
echinoid. Redrawn after Fedotov (1924,
fig. 80).

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL 515

In C. perneri, the stone canal, along with other parts of the water vascular system,
was probably degenerate or absent. The water vascular system would have been non-
functional if it did exist, for C. perneri shows no place that could have borne tube feet,
which are the effector organs of the system.

The axial sinus and main portion of the axial organ of C. perneri presumably stretched
forward from the hydropore, into which the axial sinus would have opened. It seems pos-
sible that the axial organ may correspond to the pituitary gland of vertebrates and the
homologous neural gland of tunicates. The function of the axial organ in expelling the
degeneration products of phagocytes (Millott 1966) is found also in the neural gland of
tunicates (Godeaux 1964, p. 59; Peres 1943; Millar 1953, p. 47). The possible endocrine
function of the axial gland (Millott 1967, p. 63) can be parallelled in the pituitary and
perhaps in the neural gland (Godeaux 1964, p. 57). The loss of the hydropore as Cerato-
cystis evolved into Cothurnocystis implies that a new way must have been found to
eliminate the degeneration products of amoebocytes. It may be that C. perneri , like
hemichordates, had a buccal diverticulum opening into the buccal cavity or the front
of the pharynx and in contact with the axial organ, just as the buccal diverticulum is in
contact with the equivalent glomerulus of hemichordates. Such a diverticulum could
have provided a new pathway for the expulsion of degeneration products. Komai
(1951) has already suggested that the buccal diverticulum of hemichordates (the so-
called ‘stomochord' or ‘notochord’) is homologous with the hypophysis.

THE STEM

The posterior stem (text-fig. 2a-c, 12; PI. 96, fig. 5; PI. 97, fig. 3) is fundamentally
similar to that of Coth. e/izae (Jefferies 1967, p. 171 ; 1968, p. 261) or S. curvata (Jefferies
1968, p. 275). Its skeleton consists dorsally of imbricating plates (dpi in text-fig. 12;
PI. 96, fig. 5), and ventrally of ossicles (vo in text-fig. 1 2). Unlike Coth. e/izae or S. curvata ,
however, there is more than one pair of dorsal plates for each ventral ossicle (PI. 97,
fig. 3; Ubaghs 1967, p. 13).

The ventral ossicles are wider than deep. Their dorsal surfaces are complicated;
they carry a median groove (mg in text-fig. 12; PI. 96, fig. 5; PI. 97, fig. 3) flanked by a
longitudinal lateral groove (Ig) on each side. Lateral to each longitudinal groove is a
series of low bosses (bos). In the more anterior part of the posterior stem these usually
number about three bosses on each ossicle on each side but in the more posterior part
about two bosses on each ossicle on each side. Sometimes, however, a boss straddles
an interossicular suture and, in addition, bosses on right and left sides of an ossicle
do not always exactly correspond.

Right and left series of dorsal plates meet at a lax suture in the mid-line and, together
with the ventral ossicles, they enclose a tunnel-like lumen. Each dorsal plate has a
rough external surface (exs), a well-defined anterior imbrication facet (aif), a less well-
defined posterior imbrication facet on the inner face, and an expanded base (bdp).
The anterior imbrication facet fits against the posterior imbrication facet of the plate
next in front. The expanded base articulates with the bosses on the ventral ossicles in
such a way that each base largely covers the anterior part of one boss and the posterior
part of the next boss in front. Also the base of each plate slightly overlaps the
base of the next plate in front. Successive plates were probably connected to each other
in life by connective tissue between adjacent imbrication facets. This is shown by

516

PALAEONTOLOGY, VOLUME 12

specimens in which the weak dorsal sutures have been parted after death, probably
by gas released by decay of the soft parts, while the plates of a series on each side are
nevertheless still articulated together as in life (PI. 96, fig. 5).

The soft parts of the posterior stem would have been much as in Coth. elizae or
S. eurvata. The median groove would have carried the equivalent of the crinoid cham-
bered organ (not in text-fig. 12), coated by the equivalent of the crinoid peduncular
nerve (dnc in text-fig. 12). Comparison with mitrates suggests that the peduncular nerve
(= dorsal nerve cord) was dorsal to the chambered organ (= notochord). Lateral blood
vessels (lv) probably went off from the notochord, as in Coth. elizae and S. eurvata ,
though in C. perneri they were not in contact with the skeleton. By comparison with
Coth. primaeva, which has two pairs of dorsal plates, and two pairs of grooves for
transverse vessels corresponding to each ossicle (personal observation), the transverse
vessels of C. perneri would have corresponded in number to the dorsal plates and the
bosses. The tunnel-like lumen between the dorsal plates and the ventral ossicles must
have contained muscle (mu), since the posterior stem with its imbricating dorsal plates
is adapted to flex upwards. Analogy with S. eurvata (Jefferies 1968, p. 276) suggests that
the muscle was in the form of muscle blocks, separated by the lateral vessels. Comparison
with mitrates suggests that lateral ganglia (lga) connected with the dorsal nerve cord
overlay the lateral vessels. There was probably a blood vessel down the middle of the
notochord (pv), as deduced for Coth. elizae and S. eurvata.

Ubaghs (1967, p. 12) has interpreted the posterior stem as carrying the water vascular
system of an arm in accordance with his earlier views (1961, 1963). However, the dorsal

EXPLANATION OF PLATE 97

The lengths of the scales are in mm.

Figs. 1-10. Ceratocystis perneri. 1, Natural mould in dorsal aspect to show brain and associated
structures (NM 33723/1951); same individual as Plate 95, fig. 3; dll and dir = left and right dorsal
lobes; lpr = left process; ng = infilling of narrow groove; nng = nerve to narrow groove; pbl
and pbr = left and right pyriform bodies; rf = right face of brain. 2, Natural mould in ventral
aspect to show brain and adjacent structures (W 24); gppe = groove between pharynx and posterior
coelom; lpr = left process; pbl, pbr = left and right pyriform bodies; vfl = left, ventral face of
brain; vs = ventral swelling. 3, Latex impression of stem in dorsal aspect (NM 221123/7 1924
lectotype); bos = boss; mg = median groove; stc = stylocone. 4, 5, Latex impression of infra-
branchial plates in internal aspect (GPMB Ca 32 and W 26) to show distinction between branchial
slits (bs) and accessory gaps (ag); iml dorsal margin of depression for left internal muscle.

6, Natural mould (W13) in posterior aspect of the interior of M1LD to show the nerve to the narrow
groove (nng) going round the left pyriform body (pbl) to the infilling of the narrow groove (ng).

7, Natural mould of the inside of Mpv (specimen W 4); sut = suture with M1RD; vfr = right ventral
face of brain; vs = ventral swelling. 8, Natural mould of the inside of M,L, posterior aspect (W 3);
ag = accessory gap; bs = branchial slit; gacpld = groove showing dorsal boundary between
anterior coelom and pharynx left of the stem; iml = limit of left internal muscle. 9, As fig. 8,
ventral aspect; gacplv = groove showing ventral boundary between anterior coelom and pharynx
left of the stem. 10, Natural mould of inside of M4LV (USNM 61503); cbcl = cleft between buccal
cavity and pharynx left of stem.

Fig. 11. Mitrocystites mitra. Natural mould of inside of posterior part of the theca in postero-dorsal
aspect (E 7517), to show (i) auditory nerve (aun) going round left pyriform body (pbl) to auditory
ganglion (aug) in left atrium (la), and (ii) lateral-line nerve (lln) going round right pyriform body
(pbr) to lateral line ganglion (llg); cf. Plate 98, fig. 7; hep = cast of hypocerebral processes; n5 =
nerve n5 of palmar complex; og = oblique groove; r = rectum.

Palaeontology, Vo/. 12

PLATE 97

JEFFERIES, Ceratoeystis perneri, Mitrocystites mitra

R. P. S. JEFFERIES: CERATOCYSTIS PERNER1 JAEKEL

517

plates do not resemble cover plates, for the base of each one is expanded and makes
contact with the bosses by a very broad area, and successive plates on the same side
seem to have been held together by connective tissue. The median dorsal suture was
lax, and for this reason often split open after death. The reason for this laxity was
mechanical, for when the stem flexed upwards the angle between opposite members of a
pair of plates would need to change as the pair was forced downwards on to the pair
next behind. The fact that left plates sometimes separated from right plates after death
resulted from this laxity of the median dorsal suture and does not show that they could
separate in life.

text-fig. 12. Block diagram of posterior stem of Ceratocystis perneri, left dorso-lateral
aspect, aif = anterior imbrication facet; bdp = base of dorsal plate; bos = boss on
ventral ossicle; dnc = dorsal nerve cord; dp = dorsal plate; exs = external surface
of dorsal plate; lg = lateral groove; Iga = lateral ganglion; Iv = lateral vessel;
mg = median groove; mu = muscle; not = notochord; pv = peduncular or noto-
chordal vessel; vo = ventral ossicle.

The skeleton of the medial stem consisted of a ventral stylocone (stc in text-fig.
2a-c; PI. 96, figs. 3, 5, 10; PI. 97, fig. 3) surmounted by about eight pairs of dorsal
plates. The stylocone has been accurately figured and described by Ubaghs (1967,
p. 13, fig. 6).

The skeleton of the anterior stem was more regular than Ubaghs has suggested (1967,
p. 12). It consisted of about 15 rings of plates and each ring, as in other calcichordates,
contained 4 plates— left and right dorsal, and left and right ventral (PI. 96, fig. 9).
Each ventral plate consisted of a lateral and a ventral portion (Ipvp and vpvp in PI. 96,
fig. 10) and the ventral portion extended across the ventral face to near the opposite
side (text-fig. 13). Successive rings of plates in general imbricated so that the front part
of each ring was inside the posterior part of the next ring in front. On the ventral face,
however, the ventral portions of ventral plates imbricated with one another, in such a
way that right and left ventral portions usually alternate (text-fig. 2c).

518

PALAEONTOLOGY, VOLUME 12

The soft parts of the anterior stem presumably included powerful muscles, almost
filling the large lumen. In addition there must have been, on functional as well as
comparative grounds, an anti-compressional structure extending through the lumen.
This would be the anterior part of the notochord or chambered organ whose posterior
continuation followed the median groove of the posterior stem. There would have been
a nerve, the dorsal nerve cord or peduncular nerve, from the brain to the posterior
stem. The functioning of the stem is dealt with below.

text-fig. 13. Transverse section through skeleton of anterior stem of
Ceratocystis perneri. dp = dorsal plate; Ipvp = lateral portion of
ventral plate; vpvp = ventral portion of ventral plate.

THE BRAIN AND CRANIAL NERVES

The brain of C. perneri (text-fig. 14a-e) can be reconstructed more fully than that of
later cornutes. It lay, as in all calcichordates, and like the aboral nerve centre of crinoids,
just anterior to the stem, where its position is indicated by facets on the internal cast.
These facets show that it was in contact with all those plates of the theca which touched
the anterior end of the stem, i.e. MPD, M1LD, M1LV, MPV, and M1RD. It did not touch
M1RV, which did not form part of the stem insertion in C. perneri. The cranial nerves of
C. perneri touched the skeleton in only a few places.

The morphology of the cerebral cast and of the associated structures may be described
as follows:

1. Right and left dorsal lobes (dir, dll in text-fig. 14b-e; PI. 95, fig. 7; PI. 97, fig. 1)
are situated on right and left sides of the dorsal part of the cerebral cast. They are mainly
in contact with MPD but they continue downwards in contact with M1LD and M1RD and
end in a number of points, the intraskeletal cones (iskcr and iskcl in text-fig. 14b-e;
PI. 95, fig. 6). These presumably indicate where nerves entered the dorsal lobes from
the skeleton.

2. The cerebral cast had an ant ero- dor sal face (adf) which was produced upwards
into a dorsal process (dp in text-fig. 14b-e; PI. 95, fig. 7). This latter was contained in
the dorsal groove leading to the dorsal face of the theca (see above dg in text-fig. 2a, d;
PI. 96, figs, i, 9; PI. 98, fig. 1).

3. The left face of the cerebral cast (If in text-fig. 14d) corresponds to a facet on

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI IAEKEL

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PALAEONTOLOGY, VOLUME 12

M1LD (PI- 97, fig. 1). Just anterior to it a powerful left process (lpr in text-fig. 14d, e;
PI. 97, fig. 1) went off leftwards.

4. Just left of the left process, and in contact with plates M1LD and M1LV, are indica-
tions of an approximately spherical body called the left pyriform body (pbl in text-fig.
14a, c-e; PI. 96, fig. 8; PI. 97, figs. 1, 2, 6; PI. 98, fig. 3), since it is homologous with the
like-named structure of mitrates and of other cornutes (Jefferies 1967, pp. 173, 177,
185 ff, 193; 1968, pp. 264, 276, 299 ff., 322). The right face (rf in text-fig. 14b, c; PI. 95,
fig. 6; PI. 97, fig. 1) of the cerebral cast has anterior to it the right pyriform body (pbr
in text-fig. 14a-c, e; PI. 97, figs. 1, 2; PI. 98, fig. 3). Both right face and right pyriform
body are in contact with M1RD. The right and left ventral faces (vfr and vfl in text-fig.
14a-d; PI. 97, figs. 2, 7; PI. 98, fig. 3) of the cerebral cast are in contact with the plates
MPV and M1LV. The anterior boundaries of these ventral faces are better defined towards
right and left sides of the cerebral cast than further ventrally where they pass into a
rather vague ventral swelling (vs in text-fig. 14a-d; PI. 97, figs. 2, 7; PI. 98, fig. 3).
Just left of the left pyriform body was situated the narrow groove (ng) in the external
surface of the theca in plate M1LD. This receives a projection to the narrow groove on
the internal cast (png in text-fig. 14a, e; PI. 97, fig. 6). This projection represents the
infilling of a canal in the skeleton.

EXPLANATION OF PLATE 98

The lengths of the scales are in mm.

Fig. 1. Ceratocystis perneri. Latex impression showing dorsal surface in dorsal aspect (W 8); ag
accessory gaps; bs = branchial slits; dg = dorsal groove; faf = flap attachment facet; fpl = prob-
able flap platelets; ib = infrabranchial plates.

Fig. 2. Cothurnocystis americana. Dorsal aspect of only known specimen and holotype (USNM
143237); bs = branchial slits; inbc = integument over buccal cavity; in p = integument over
pharynx; ? posM4LD = probable position of plate M4LD; str = strut.

Fig. 3. Ceratocystis perneri. Plaster cast of silicone rubber mould of Copenhagen specimen, simulating
natural mould; the ventral surface of the brain is better marked off anteriorly than in any other
specimen examined; gppc = groove between pharynx and posterior coelom; pbl and pbr = left
and right pyriform bodies; vfl = left ventral face of brain; vs = ventral swelling.

Fig. 4. Cothurnocystis elizae. Natural mould of inside of theca in postero-ventral aspect (E 28667),
with the specimen lying dorsal side downwards, cf. text-fig. 6; 1, 3, 4 = points where the pharyngo-
visceral line approaches the dorsal side of the theca; grg = gonorectal groove; ht = probable
position of heart; pbr = right pyriform body; pco = posterior coelom; sp = striations on the inside
of the pharynx; str = impression of strut.

Fig. 5. Ceratocystis perneri. (a) Plaster cast simulating natural internal mould of plate M2R, posterior
aspect; ( b ) Latex cast of plate M2R, posterior aspect. Magnification, of (a) and ( b ) equal. Both are
from same specimen. Narodni Muzeum, no number, h = hydropore; gio = groove round internal
organ.

Fig. 6. Cothurnocystis primaeva. Natural mould of inside of theca (Montpellier specimen) in ventral
aspect, to show region just anterior to stem; cf. text-fig. 7; br = brain; gd = gonoduct; grg = in-
filling of gonorectal groove; iskc = intraskeletal cones; mini = left median-line nerve; pbl, pbr =
left and right pyriform bodies; r = rectum.

Fig. 7. Mitrocystites mitra. Natural mould of inside of theca in postero-dorsal aspect (MCZ 566);
cf. text-fig. 15 and Plate 97, fig. 11; aug = auditory ganglion; aun = auditory nerve; la = left
atrium; llg = lateral line ganglion; lln = lateral line nerve; mp = median part of brain; n3 =
optic nerve; n4 + 5, n5 = other nerves of palmar complexes; og = oblique groove; pbl, pbr - left
and right pyriform bodies; pco = posterior coelom; pp = posterior part of brain; r = rectum;
ra = dorsal margin of right atrium.

Palaeontology, Vol. 12

PLATE 98

JEFFERIES, Ceratocystis perneri, Cothurnocystis spp., Mitrocystites mitra

R. P. S. JEFFERIES: CERATOCYSTIS PERNER1 JAEKEL 521

Interpretation of these features of the internal cast is possible by comparison with
other cornutes and, more particularly, with mitrates (text-figs. 15, 16; PI. 97, fig. 11;
PI. 98, fig. 7; Jefferies 1967, pp. 185 ff., 191 ff.; 1968, pp. 295 ff., 319 ft'.,). Taken to-
gether, the dorsal lobes (dl) and intraskeletal cones (iskc) are comparable with the
anterior part of the brain of mitrates, which likewise is dorsal in position and probably
also received nerves from the skeleton, sometimes by way of intraskeletal cones (e.g.
MitrocysteUa incipiens miloni , Jefferies 1967, fig. 10; 1968, figs. 19 a-c). The anterior part
of the brain in mitrates represented the telencephalon and was olfactory. Consequently
the intraskeletal nerves of C.perneri, which presumably entered the dorsal lobes by way
of the intraskeletal cones, would be the olfactory nerves, and the dorsal lobes would
represent the telencephalon. The antero-dorsal face (adf) of the brain in C. perneri
is ventral to the dorsal lobes, i.e. ventral to the telencephalon, and this part of the brain
in mitrates and all other chordates is optic in function. The dorsal process (dp) that
extended upwards on to the dorsal surface of the theca would therefore also be optic
in function. It would represent a sort of primitive median eye. This, however, was prob-
ably not homologous with the pineal or parapineal eyes of vertebrates, since it runs
upwards in front of the telencephalon, rather than behind it, and is absent from all
known mitrates. The right and left pyriform bodies (pbr and pbl) of C. perneri corre-
spond to the pyriform bodies of other calcichordates and these represent the trigemino-
profundus ganglia of vertebrates (Jefferies 1967, p. 190; 1968, p. 307). The ventral
swelling of the cerebral cast (vs) may possibly correspond to the places where ventrally
placed nerves left the brain. These would correspond to the median-line nerves of other
cornutes (1967, pp. 173, 177; 1968, pp. 264, 276). The left process (lpr) presumably
represents a large nerve to the theca (ner in text-fig. 14c, d) corresponding to either
posterior or medial part nerves, or both, of mitrates, i.e. to optic or medullary nerves,
or both. If a corresponding right nerve existed in C.perneri it did not touch the skeleton.

The narrow groove (ng) on the surface of M1LD recalls the narrow groove or lateral line
that exists in mitrates (lateral line (11) in Jefferies 1967, pp. 178, 190; narrow groove (ng) in
Jefferies 1968, pp. 283, 299 ft'., 314). If the projection to the narrow groove of C. perneri
(png) carried a nerve, as seems likely, then the nerve supply to the groove (nng in text-fig. 14
a, c, e) was like that to the groove of mitrates in coming round the outside of a pyriform
body. However, the narrow groove of C. perneri was left of the stem, and its nerve
supply came round the left pyriform body, whereas the narrow groove of mitrates was
right of the stem, and its nerve supply came round the right pyriform body.

This anomaly demands a partial restudy of the mitrates. In Mitrocystites mitra there
is a previously unrecorded ridge on the internal cast (aun in text-fig. 15; PI. 97, fig. 1 1 ;
PI. 98, fig. 7) which sweeps round the front and left side of the left pyriform body (pbl),
behind the rectum (r), and appears to enter the left atrium (la) to end there in a little
lump (aug). With respect to the pyriform body this ridge corresponds in its position on
the left of the theca to the ridge in M. mitra representing the course of the lateral line
nerve (lln) on the right. In addition, the lump (aug) is a left counterpart of the lump
representing the lateral-line ganglion (llg) on the right. It therefore seems that the two
sets of structures are right and left antimeres of each other, except that the left one ends
in the left atrium, while the right one ends at the surface.

The acustico-lateralis system of fishes includes two parts — the lateral line which lies
at or near the surface, and is directly sensitive to displacements of water (Dijkgraaf

522

PALAEONTOLOGY, VOLUME 12

1963, Bergeijk 1964), and the acoustic system, which is invaginated and sensitive to
pressure waves, gravity and angular acceleration. It seems that the acustico-lateralis
system of mitrates had two corresponding parts, i.e. external and functioning as lateral
line on the right, and invaginated and presumably functioning as ear on the left. The
ridge aun would therefore represent the auditory nerve and the lump aug would rep-
resent the auditory ganglion. The fact that the auditory nerve runs immediately in front
of the left pyriform body, behind the rectum, suggests that the lateral line nerve (lln)
probably ran immediately in front of the right pyriform body. Its proximal course was

text-fig. 15. a. Portion of internal mould of Mitrocystites mitra to show auditory nerve
and ganglion (MCZ 566, cf. PI. 98, fig. 7). b. Diagram of dorsal aspect of theca, to show
portion (stippled) included in a. aug = auditory ganglion; aun = auditory nerve; la = left
atrium; mp = medial part of brain; n2, n3, n4+5 = nerves of left palmar complex;
og = oblique groove; pco = posterior coelom; pp = posterior part of brain; r =
rectum. For meaning of anatomical terms see Jefferies 1967, p. 1 79fF. ; 1968, p. 315ff.

therefore probably different from what I previously supposed (1967, figs. 10, 14; 1968,
figs. 19, 27). Text-fig. 16 is a revised reconstruction of the cranial nerves of M. mitra
that takes these new conclusions into account.

The narrow groove (ng) of C. perneri and the nerve supplying it (nng) correspond
closely in position to the auditory ganglion and auditory nerve of Mitrocystites mitra.
The nerve nng must be homologous with the auditory nerve of M. mitra and the floor
of the narrow groove presumably housed the homologue of the auditory ganglion.
In C. perneri the narrow groove, being on the surface of the animal, would have func-
tioned as lateral line.

The early history of the acustico-lateralis system in the chordates would therefore
have been somewhat as follows. The system is first seen in C. perneri as a surface groove
just left of the stem that functioned as lateral line, being directly sensitive to displace-
ments of the sea water. In Cothurnocystis this lateral line must still have existed, but
probably lay just right of the gonopore-anus, without any separate opening in the

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

523

text-fig. 16. Reconstruction of brain and cranial nerves of Mitrocystites mitra in dorsal aspect
(modified from Jefferies 1968, fig. 27). ap = anterior part of brain; aug = auditory ganglion; aun
auditory nerve; be buccal cavity; e = eye; llg = lateral line ganglion; lln lateral line nerve;
nip = medial part of brain; mpn = medial part nerve; n0 = ventral nerves of uncertain connection
and homology; n,_5 = nerves of palmar complex including maxillary trigeminal (n2) and optic (n3)
nerves; n2AfP = branches of nerve n2; olo = olfactory openings; pal = palmar nerves; pbl and
pbr = left and right pyriform bodies (trigemino-profundus ganglia); pg peripheral groove; pp
posterior part of brain; ppn = posterior part nerves; r rectum.

524

PALAEONTOLOGY, VOLUME 12

skeleton. The nerve that supplied it would have swept round the front of the left pyriform
body and followed the posterior surface of the rectum to the lateral line. When the
left gill slits, anus, gonopore and lateral line became enclosed in the left atrium, which
happened either when the mitrates evolved, or somewhat before, the lateral line changed
its function. Displacement of the surrounding sea-water no longer affected it, but it
would be able, in a rudimentary way, to fulfil the functions of an ear. It would thus be
sensitive to pressure waves, to angular rotation that caused swirling of water in the
left atrium, and to gravity, if calcareous otoconiae existed in the cupolae of the neuro-
masts (Pumphrey 1950, p. 13). Among mitrates, the Mitrocystitidae again became sensi-
tive to displacements of the external sea-water by evolving a lateral line, with its nerve
supply, on the right side.

The appearance of organs on the right which had previously existed only on the left
was a feature of the evolution of mitrates from cornutes. It is seen in the pharynx, the
gill slits, and probably the atria, as well as the acustico-lateralis system. The earliest
known occurrence of a lateral line on the right side is found in Chinianocarpos thorali
Ubaghs (Upper Tremadoc or Lower Arenig), which is the oldest mitrocystitid known.
Such a feature is unknown in the Peltocystidae, which is the other family of mitrates
represented at the base of the Ordovician. The way in which the primitive mitrocystitid
acustico-lateralis system evolved into that of vertebrates cannot be worked out, since
it would have happened in soft-bodied animals. Presumably, however, the lateral-line
was replicated on the left side of the body and the acoustic system on the right. This
replication on opposite sides of the body would be similar, but opposite in sense, to
what had happened in the evolution of mitrates. It is interesting that the only skeletal
calcium carbonate which vertebrates possess is found in the acustico-lateralis system
(otoconiae and otocysts). This may represent an inheritance from calcichordates, in
which the acustico-lateralis system was better developed than I previously supposed.

The relation between brain and blood system in C. perneri is of interest. The brain
was in contact with the thecal skeleton all the way round the stem insertion. The blood
supply to the stem must therefore have gone through the middle of the brain, just as the
haemal strand goes through the aboral nerve centre of crinoids ( Reichensperger 1905).
Like the haemal strand, it probably continued backwards down the middle of the cham-
bered organ (= notochord) as a notochordal vessel (Jefferies 1967, pp. 172, 184;
1968, pp. 263, 293). Anterior to the brain the notochordal vessel would presumably be
connected to the axial organ, as the haemal strand is in crinoids (Reichensperger 1905).

POSTURE, HABITS, AND MODE OF FEEDING

The posture of C. perneri would have been very like that of Coth. elizae, i.e. the theca
rested ventral side downwards on the bottom with the stem stretched out almost horizon-
tal behind the theca. Certain portions of the ventral surface of the theca (stippled in
text-fig. 17a) were ventrally prominent and would have touched the sea floor more
frequently or sunk deeper into it than the less prominent parts. Those ventrally promin-
ent parts would have been the places that bore most of the weight of the theca and are
constructed of more massive stereom than the other parts of the ventral surface (PI. 95,
fig. 1). Indeed, the massiveness of the stereom is roughly proportional to the prominence
of the part which it forms. Massive stereom would be smoother than loose-textured

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

525

stereom, and would have less soft tissue within it that could be damaged by being scraped
over the sea floor.

In later members of the Ceratocystidae the ventrally projecting, weight-bearing parts
of the ventral surface became better developed (e.g. Coth. elizae, text-fig. 17b). In such
forms they had the function of raising the ventral integument clear of the sea floor.

L

text-fig. 17. Ventrally prominent areas (stippled) and observed positions of stem in a, Ceratocystis
perneri and b, Cothurnocystis elizae. or in = oral integument; stc = stylocone. R and L = especially
frequent positions of stem to right and left. The arrow a indicates probable easiest direction of slip in
Coth. elizae. In the histograms each dot indicates one specimen.

so that its pumping movements would not be hindered. This raising of the ventral
integument is particularly marked in Scotiaecystis curvata (Jefferies 1968, figs. 8 c-e)
but is also developed, to lesser degree, in Coth. elizae (figs. 1 d-g) and is incipient in
Coth. primaeva. The form of the ventral spikes and anterior appendages of these three
species suggests that the theca could have slipped backwards much more easily than
forwards (1968, pp. 265, 277) and this was probably also true of mitrates (1968, pp. 310,
323). The probable direction of easiest slipping in Coth. elizae is shown by the arrow a
in text fig. 17b.

The form of the ventrally prominent parts of the theca of C. perneri suggests that the
theca could already slip more easily backwards than forwards, though the tendency
was less pronounced than it became in later Ceratocystidae. Thus the anterior part of

526

PALAEONTOLOGY VOLUME 12

the spike S3R (text-fig. 2b; PI. 95, fig. i) would have stuck deeper into the sea-floor
than its posterior part, and the same is true of the left appendage (lap in PI. 95, fig. 1).

The stem of C. perneri is in all observed specimens stretched out on a bedding plane,
and could probably wave from side to side. Text-fig. 17a shows its position in 105
specimens. There are two especially frequent positions (L and R) which also existed
in Coth. elizae (text-fig. 17b). These perhaps correspond to the most usual end positions
of the stem in its side-to-side movements after burial and before death.

The massive texture of the stereom of the ossicles of the posterior stem, of the ventral
surface of the anterior stem, and of part of the ventral surface of the stylocone (as
stippled in text-fig. 17a, cf. PI. 96, fig. 3; Ubaghs 1967, pi. 2, fig. 8) resembles that of the
ventrally prominent parts of the theca. It is likely that the massive-textured parts of the
stem, like the similar parts of the theca, were habitually in contact with the sediment of
the sea bottom. Conversely, the loose-textured part of the ventral surface of the stylocone
would not usually have touched the sediment, or would have touched it more lightly.
The distribution of these variations in texture suggests that the stem stretched out almost
horizontally in life so that the anterior and posterior parts sank somewhat into the
sediment, while most of the stylocone was just clear of the sediment, or sank into it
less deeply. This agrees with the observed position of the stem in fossils, parallel to the
bedding. With such a posture, abrupt lateral flexion of the stem would have caused the
theca to slip over the sea bottom, mainly backwards, and would have provided a clumsy
but workable means of locomotion. Pulling an asymmetrical object is easier than pushing
it, being directionally stable. This was the probable reason why cornutes, as here be-
lieved, took to moving backwards.

C. perneri would have been a deposit feeder like C. elizae (Jefferies 1968, p. 258),
since its mouth was level with the bottom. It is likely that water was pumped through
the pharynx mainly by ciliary action, but that the animal could also ‘cough’ slightly,
using the rudimentary bellows mechanism of the theca.

EVOLUTION IN THE CER ATOCYSTID AE

Ceratocystis perneri, Cothurnocystis aniericana, Cothurnocystis primaeva, and Coth-
urnocystis elizae closely approximate to a single line of descent (text-fig. 1), and most of
the changes in this line of descent can be explained functionally.

C. perneri had a boot-shaped theca formed of calcite plates. This theca rested on the
sea bottom by the ventral side, which carried ventral spikes to hold the theca in the sea
floor and to carry most of the thecal weight. The animal moved by dragging the theca
backwards by the stem. The theca had a mouth at the anterior end, and hydropore,
gonopore, and anus existed right of the stem. Left of the stem were 7 gill slits. The theca
contained 4 chambers, i.e. buccal cavity, pharynx, anterior and posterior coelom. A
respiratory and feeding current was created, probably mainly by ciliae on the insides
of buccal cavity and pharynx. These two chambers, however, could also contract, slightly
but suddenly, by the action of muscles in the thecal wall. The dorsal thecal wall was
stiffened by keels which assured that this ‘coughing’ action caused contraction of the
thecal cavity, and expulsion of water, rather than merely changing the shape of the
thecal cavity without change in volume. Most of the food supply came from the organic
detritus lying on the sediment, i.e. the animal was a deposit feeder. Internal organs in the

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL 527

right-hand posterior corner of the theca (i.e. in the anterior coelom) included the post-
pharyngeal gut, the gonads, the heart and pericardium, probably an axial gland, and
perhaps parts of a degenerate water vascular system. The brain lay at the anterior
end of the stem and at the anterior end of notochord and dorsal nerve cord. Olfactory
and optic parts can be identified in the brain. The optic part gave rise to a sort of
median eye, which lay on the dorsal surface of the theca. Just anterior and lateral to the
brain were two ganglia (the pyriform bodies) homologous to the trigemino-profundus
ganglia of vertebrates. Just left of the stem was a groove on the surface that represented
the first beginnings of the acustico-lateralis system and was supplied by a nerve coming
round the left pyriform body.

Coth. americana was functionally a major advance on C. perneri but is unfortunately
not well known. The theca was still boot-shaped, but more like that of Coth. elizae
than that of C. perneri in shape. The theca had become much more flexible than it was
in C. perneri by the development of a dorsal integument. It seems likely that muscular
pumping had become the main way of expelling water through the gill slits. The most
important pumping muscles would be those in the roof of the pharynx, which would be
partly housed in large spaces between the plates. The roof of the buccal cavity was
flexible, but lacked comparable spaces for muscles. The gill slits of Coth. americana,
which still numbered 7, were very similar to those of Coth. elizae and were well adapted
as outlet valves. The hydropore had disappeared, and it is possible that the axial gland
had started to discharge through a duct in the anterior part of the pharynx, i.e. had
turned into the pituitary gland. Gonopore and anus had started to migrate leftwards,
so as to approach the exhalent current from the gill slits, though it is not possible to say
how far they had moved. The flexibility of the thecal roof meant that the floor of the
theca was liable to buckle when the muscles of the dorsal integument contracted. This
tendency was counteracted by thickening the floor along an antero-posterior line to
form a strut.

Coth. primaeva differed principally from Coth. americana in having a flexible floor
to the theca, crossed by a rigid strut. The pumping action of the theca therefore involved
movements of the floor, as well as the roof, and would have been more efficient. Gono-
pore and anus were just left of the stem in Coth. primaeva , as they were in Coth. elizae.
They would thus lie in the outwash from the gill slits. Heart, pericardium, gonads,
most of the post-pharyngeal gut, and the pituitary gland lay in the anterior coelom,
as in C. perneri. The rectum and the distal part of the gonoduct, however, ran across
the floor of the posterior coelom, from the anterior coelom to the gonopore-anus.
The brain of Coth. primaeva probably resembled that of C. perneri in most respects,
including the presence of a median eye.

Coth. elizae did not much differ from Coth. primaeva but is better known, because
more specimens are known. It lacked the median eye and had more gill slits (16 instead
of about 7). Spikes on the ventral surface would have lifted the thecal floor above the
sea bottom, so that movement of the ventral integument would not be hindered.

The foregoing history involves some oversimplification. Thus Coth. americana is only
known to occur at a horizon that also contains a cornute with a flexible thecal floor
( Phyllocystis sp. Ubaghs 1963). Coth. americana itself, therefore, must represent a late
survival of the stage with flexible roof and rigid floor, which other cornutes had already
surpassed. Again the outline of the theca of Coth. primaeva is not intermediate between

C 6685

m m

528

PALAEONTOLOGY, VOLUME 12

that of Coth. americana and Coth. elizae , but suggests affinities with Scotiaecystis curvata.
These discrepancies are probably not important.

The line connecting C. perneri with Coth. elizae was a conservative one. Other cornutes
diverged more from the C. perneri type. The line of descent leading to S. curvata pre-
sumably separated from the perneri-elizae line after a flexible thecal floor had been
acquired. It is represented by an undescribed form contemporary with Coth. primaeva
(Ubaghs, personal communication). S. curvata was a suspension feeder; its mouth opened
upwards and the ventral integument was lifted well above the sea floor by strong curva-
ture of the frame so as to improve its pumping efficiency. The gill slits are also highly
specialized.

The Phyllocystis line of descent, which includes bilaterally symmetrical forms, also
separated from the perneri-elizae line after the acquisition of a flexible thecal floor.
The increased symmetry of the theca was probably acquired by bending the ‘ankle’
part of the boot-shaped theca leftwards, so that the mouth also came to point leftwards.
Coth. ubaghsi Chauvel (1966, p. 98) in which the buccal cavity is strongly bent to the left,
may represent an intermediate stage in this process. The mitrates were probably derived
from Phyllocystis. Some primitive mitrates belonging to the Mitrocystitidae have a
leftward pointing mouth (e.g. Chinianocarpos thorali Ubaghs, Jefferies 1968, p. 314).
From primitive mitrates the other chordate subphyla are probably descended.

CALCICHORDATES, HEMICHORDATES, AND ECHINODERMS

The extraordinary asymmetry of Ceratocystis perneri, together with the position of
the anus right of the stem, suggest that it evolved from a bilaterally symmetrical animal
that lay down on its right-hand side. Moreover, it seems likely that the animal in question
was a pterobranch hemichordate. Thus text-fig. 18 compares the reconstructed anatomy
of C. perneri, based on echinoderm and tunicate analogies, with the anatomy of the
living pterobranch Cephalo discus. The disposition of organs is similar, if the ventral side
of C. perneri is taken to correspond to the right side of Cephalodiscus.

A digression on the anatomy of Cephalodiscus is now necessary, before pursuing this
comparison in detail. Cephalodiscus zooids (van der Horst 1 935) normally live in bundles
of horny tubes, but are capable of leaving the tubes to wander over the surface of the
bundle. Each zooid consists of three parts, which can be called protosome (head shield)
(prs in text-fig. 18b), mesosome (collar), and metasome (trunk), and the metasome is
extended to form a stem (st), at whose distal end buds are produced (not shown in
text-fig. 18b).

The protosome contains an unpaired coelom, the protocoel. It also contains, in a
dorsal position, the pericardial vesicle and heart, the buccal diverticulum (bd) (‘noto-
chord’, ‘stomochord’) and an excretory glomerulus. A right and a left protocoel pore
(hj), lateral to the heart, connect the protocoel with the outside.

The mesosome contains a right and left coelom, the mesocoels, separated by a mesent-
ery. It carries a right and a left array of tentacles and each tentacle contains an extension
of the mesocoel of the same side. Down the ventral side of each tentacle runs a food
groove to the region of the mouth. A ganglion (ga) is situated dorsally in the mesosome,
in the mid-line, between the right and the left sets of tentacles. Right and left mesocoels
open to the outside, each by a separate mesocoel pore (h2). The mouth (m) is slit-shaped,
and lies ventrally, between protosome and mesosome.

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

529

A

|notochord = extension of right metocoel into stem
posterior coelom = right metocoel

buccal cavity

janterior coelom = left metacoel (not shown) |° \ gonad

Jaxocoel = protocoel j pericardium

|mesocoel [ [stem muscle

i jph arynx ner ve

text-fig. 18. Homologies between Ceratocystis perneri (a) and the living hemichordate Cephalodiscus
(b): an = anus; axg axial gland; bd = buccal diverticulum; br = brain; bs = branchial slits;
dnc = dorsal nerve cord ; ga = mesosomal ganglion of Cephalodiscus; h = hydropore; hx = protocoel
pore; h2 = mesocoel pore; m mouth; prs = protosome; st = stem; stn = peduncular nerve.

The metasome contains a right and a left coelom, the metacoels, separated by a
mesentery. It also carries the whole of the gut behind the buccal cavity, and the gonads.
The gut is U-shaped and consists of buccal cavity, pharynx, oesophagus, stomach,
and intestine. Right and left walls of the pharynx are each penetrated by a single gill
slit (bs). There are 2 gonads, one in the left and one in the right metacoel, and each opens
by a single gonopore (g) to the outside.

530

PALAEONTOLOGY, VOLUME 12

The stem is muscular and highly innervated particularly on its ventral side (peduncular
nerve stn). The coelom of the stem is an extension of the right and left metacoels, which
in the stem of Cephalodiscus itself are not separated by mesenteries. In the related
Rhabdopleura, however, the mesentery separating right and left metacoels continues
to the end of the stem.

Animals of Cephalodiscus type have probably existed for a very long time. The fossil
remains of Eocephalodiscus, described by Kozlowski (1948, p. 195), from the Tremadoc
of Poland, differ very little from the skeletons of recent Cephalodiscus colonies. Also
Kozlowski has argued (1947, 1966) that graptolites were pterobranchs, and the first
graptolites are known from the Middle Cambrian (Obut 1964, p. 306).

The homologies of pterobranchs and echinoderms can be expressed in a table.

Pterobranch Echinoderm

protocoel

left protocoel pore

right protocoel pore

glomerulus

pericardium

heart

buccal diverticulum
left mesocoel
left mesocoel pore
right mesocoel
right mesocoel pore
mesosome ganglion
left metacoel
right metacoel
left gonopore
left gonad
right gonopore
right gonad
stem

right metacoel extension into stem
left metacoel extension into stem
innervation of stem

axocoel

hydropore (in part)
absent

main portion of axial gland
dorsal sac

head process of axial organ
absent
hydrocoel
hydropore (in part)
reduced or absent
absent
absent

left (oral) somatocoel
right (aboral) somatocoel
gonopore
gonad or gonads
absent
absent

stem

coelom of chambered organ
absent

peduncular nerve

These homologies are based largely on the discussion given by Fedotov (1924, p. 298)
except that his assertion that the pericardial sac is a right antimere of the protocoel is
disregarded. Also the stem homologies given here never seem to have been proposed
before. It is here assumed that the hydropore of echinoids represents the fusion of the
protocoel and metacoel pores of the left side of a pterobranch, instead of corresponding
to one rather than the other.

It is interesting that Grobben (1924) derived echinoderms from a pterobranch that
came to rest on its right side. His views on the changes involved are unnecessarily com-
plicated, however, by his attempt to derive the echinoderm stem from the region of the
hemichordate protosome. Other authors such as Bather (1900) have derived echino-
derms from an essentially hemichordate and bilateral ‘ dipleurula ’ ancestor that came to
rest on its right side. The conclusion that the bilateral ancestor of echinoderms lay down
on its right side is based on the absence in echinoderms of the right protocoel pore,
right mesocoel pore, and right gonopore and the reduction of the right mesocoel as

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL 531

compared with the left mesocoel. Also, if a pterobranch lay on its right side, the food
grooves of the right tentacles would touch the sea floor, and so would become useless
and disappear, while the food grooves of the left tentacles would face upwards. The
absence of the originally right mesocoel and tentacles, and the upward orientation of
the food grooves of the remaining, originally left, tentacles are observed features of
stemmed echinoderms.

Returning now to the comparison between Ceratocystis and Cephalodiscus , the simil-
arity can be expressed by saying that, starting from the stem of either, and proceeding
clockwise round text-fig. 18a and b one passes in succession: (1) the branchial slits
(bs), (2) the mouth (m), (3) the hydropore (h) or equivalent protocoel pore (fq) and
mesocoel pore (h2), (4) the gonopore (g), and (5) the anus (an).

There are many obvious differences between Cephalodiscus and Ceratocystis. Thus
Cephalodiscus : (1) Lacks a calcite skeleton; (2) has paired tentacles; (3) has paired proto-
coel and mesocoel pores, instead of a single hydropore; (4) has paired gonopores,
instead of one only; (5) has a relatively smaller buccal cavity; (6) has a small pharynx
with a single pair of gill slits, instead of a large pharynx with seven gill slits on the origin-
ally left (= dorsal) side; (7) has the anus, gonopores and equivalents of the hydropore
much further from the stem; (8) has an important ganglion near the tentacles which,
so far as known, is unrepresented in Ceratocystis ; (9) has the stem coelom continuous
with the coelom of the trunk (metacoel) instead of separated from it; (10) has no big
ganglion at the proximal end of the stem; (11) has the metacoels paired and equal, by
contrast with the probably equivalent, anterior and posterior coeloms of Ceratocystis,
one of which is nearer the stem than the other. Most of these differences would represent
either the loss by Ceratocystis of organs present in Cephalodiscus or else changes in
the relative sizes of parts.

C. perneri shares with stemmed echinoderms many of the features which separate
it from Cephalodiscus. The most important of such features are: (1) the calcite skeleton;
(2) the position of one perivisceral coelom (right somatocoel, posterior coelom) nearer
the stem than the other (left somatocoel, anterior coelom); (3) the separation of the
stem coelom (coelom of the chambered organ, cavity of the notochord) from the peri-
visceral coelom; (4) the presence of an important ganglion at the proximal end of the
stem (aboral nerve centre, brain); (5) the presence of a single hydropore (cf. cystoids),
instead of paired protocoel and mesocoel pores; (6) the singleness of the gonopore (cf.
cystoids).

These common features suggest that stemmed echinoderms and calcichordates have a
common ancestor more recent than the common ancestor of both with Cephalodiscus.
Also this more recent common ancestor would resemble a calcite-plated hemichordate
that habitually rested on its right side.

The resemblances and differences between Cephalodiscus, stemmed echinoderms and
Ceratocystis suggest the following evolutionary history. A population of Lower Cambrian
animals resembling Cephalodiscus vacated, as Cephalodiscus can, the tubes in which
they had normally dwelt. They took to moving over the sea floor, by the action of their
muscular stems, lying on the right side of their bodies. Because of this orientation
they lost the body openings of the right side, i.e. right gill slit, right protocoel pore,
right mesocoel pore, and right gonopore. Further, the right tentacles found them-
selves with the food grooves forced downwards into the sea-floor and so became useless

532

PALAEONTOLOGY, VOLUME 12

and disappeared. The left tentacles found themselves with food grooves upwards and
were retained. At about this time the left protocoel pore and left mesocoel pore fused
together to form a hydropore, the calcite skeleton appeared, the metacoels lost their
simple paired arrangement, so that the right metacoel came nearer the stem than the
left metacoel, and alone provided the stem coelom. This stem coelom then became sepa-
rate from the right metacoel in the adult, so that the stem could now be bent without
being shortened or lengthened, and a large ganglion evolved at its proximal end.

Evolution now proceeded in two directions. One population specialized in tentacle
feeding and lost the gill slits. It gave rise to the stemmed echinoderms. Some of these
lifted the mouth upwards, so that the left somatocoel came to overlie the right soma-
tocoel and the hydropore, gonopore and anus came to be arranged in a vertical plane,
as in cystoids. Another population specialized in pharyngeal feeding and lost the
tentacles. This group gave rise to the calcichordates, including Ceratocystis. The
expansion in the size of the buccal cavity and pharynx, and the increase in the number
of gill slits and their migration to the downstream end of the pharynx, were adaptations
for producing a more powerful pharyngeal current. The originally U-shaped gut, which
may have been an adaptation to the original tubicolous habit, had been lost at some stage
in the sequence described.

CALCICHORDATES AND OTHER CHORDATES

The present work does not much modify the picture previously given (Jefferies 1968,
p. 331) of the evolution of the extant chordate subphyla from the Calcichordata.
However, the beginnings of the acoustic part of the acustico-lateralis system seem
already to have existed in mitrates; also, the left atrium of mitrates received a gonoduct
as well as the rectum, and the heart and pericardium already existed in the anterior
coelom. It is possible, also, that the pituitary or neural gland may have existed in mitra-
tes, having been derived from the axial gland, and that the gill slits in mitrates may have
numbered about seven on each side.

CONCLUSIONS

1 . Ceratocystis perneri is here interpreted as the oldest chordate known, and the oldest
known member of the order Cornuta. It had affinities with echinoderms, but is best
regarded as a chordate; from it, or from very similar unknown forms, all later chordates
are probably descended.

2. The bellows system of pumping water through the pharynx, which is well developed
in later cornutes, had already begun to appear in C. perneri.

3. C. perneri had 7 gill slits, and this was probably the primitive number for cornutes,
and therefore for chordates in general.

4. C. perneri possessed mouth, hydropore, gonopore, and anus arranged in the same
order as in primitive stemmed echinoderms such as cystoids. Gonopore and anus also
existed in the later cornute Cothurnocystis, in which rectum and gonoduct had come to
be disposed fundamentally as in modern enterogonous tunicates.

5. The thecal cavity of C. perneri, like that of later cornutes, was divided into buccal
cavity, pharynx, anterior coelom, and posterior coelom.

6. The anterior coelom of C. perneri would have contained, near the hydropore,

R. P. S. JEFFERIES: CERATOCYSTIS PERNERI JAEKEL

533

a heart and a pericardium. These resembled in position, and were presumably homolog-
ous with, the heart and pericardium of other chordates. They were also homologous with
the heart and pericardium of living hemichordates and the head process of the axial
organ and dorsal sac of living echinoderms.

7. The main portion of the axial organ of echinoderms may be homologous with the
neural gland of tunicates and the pituitary gland of vertebrates.

8. The stem of C. perneri did not differ fundamentally from that of later cornutes.
The anterior stem, contrary to Ubaghs (1967), was regularly tetraserial.

9. The brain of C. perneri had paired olfactory dorsal lobes, homologous with the
telencephalon of vertebrates, and a dorsal median eye which was not homologous with
the median eyes of vertebrates. Paired pyriform bodies (trigeminal ganglia) existed as in
later calcichordates. A lateral line was already developed in C. perneri, and evidence
suggests that the acustico-lateralis system of some mitrate calcichordates was already
divided into acoustic and lateral-line subsystems. The blood supply to the stem of C.
perneri must have gone through the middle of the brain.

10. C. perneri lay habitually on its ventral side, and could drag itself backwards by
side-to-side wagging of the stem. It was a deposit feeder.

1 1. The reconstructed anatomy of C. perneri can be compared with the anatomy of a
modern pterobranch hemichordate lying on its right side. On this comparison the stem
of hemichordates would correspond to the stem of stemmed echinoderms and calci-
chordates and to the tail of other chordates. The extension of the right metacoel into
the hemichordate stem would be homologous with the chambered organ of stemmed
echinoderms and the notochord of chordates. It is suggested that a population of
Cambrian pterobranchs took to wandering over the sea floor with right side downwards
and acquired calcite skeletons. This population gave rise to the echinoderms, by elabora-
tion of the tentacles and loss of the gill slits, and to the chordates, by loss of the tentacles
and elaboration of the gill slits.

Acknowledgements. It is a pleasure to acknowledge the help of many Museum curators who have made
specimens available. In particular I wish to thank Drs. V. Zazvorka, R. Horny, and R. J. Prokop
(Prague), Dr. P. Kier (U.S.N.M., Washington), Professor B. Ruzicka (Ostrava), Dr. H. Kolmann
(Vienna), Drs. H. W. Rasmussen and N. Bonde (Copenhagen), Dr. H. Jaeger (Berlin), Dr. H. Nestler
(Greifswald), Professor M. Pfannenstiel (Freiberg i. Br.), M. J. Mattei and M. K. Revert (Montpellier),
and Professor L. David and M. B. Walter (Lyon). Professor G. Ubaghs (Liege) has, as always, done
all he could to assist me in work which ran exactly counter to his own ideas. Professor N. Millott
(Bedford College, London) helped to clarify ideas on axial complexes, and Dr. P. Toghill (British
Museum (Natural History)) helped with relevant graptolite references. Mr. P. Minton (Imperial
College, London) contributed many ideas on functional morphology. Dr. L. R. M. Cocks has helped
greatly by constructive criticism of the presentation.

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R. P. S. JEFFERIES
Department of Palaeontology
British Museum (Natural History)
Cromwell Road

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PALAEONTOLOGY

VOLUME 12 • PART 3

CONTENTS

Medusae from uppermost Precambrian or Cambrian sandstones, central
Australia. By mary wade 351

Specific frequency and environmental indicators in two horizons of the
Calcaire de Ferques (Upper Devonian), northern France. By peigi
WALLACE 366

On the structure and relationships of a new Pennsylvanian species of the
seed Pachytesta. By t. n. taylor and d. a. eggert 382

The ontogeny of the thecideacean brachiopod Moorellina granulosa
(Moore) from the Middle Jurassic of England. By P. g. baker 388

A conodont assemblage from the Carboniferous of the Avon Gorge,
Bristol. By r. l. Austin and f. h. t. Rhodes 400

The Tremadoc trilobite Pseudokainella impar (Salter). By p. h. whitworth 406
A new species of Aulacotheca (Pteridospermales) from the Middle
Pennsylvanian of Iowa. By d. a. eggert and r. w. kryder 414

Miospores from the Lower Carboniferous Basement Beds in the Menai
Straits region of Caernarvonshire, north Wales. By f. a. hibbert and
w. s. LACEY 420

Megaspore assemblages from Visean deposits at Dunbar, East Lothian,
Scotland. By e. spinner 441

Crustacean burrows in the Weald Clay (Lower Cretaceous) of south-
eastern England and their environmental significance. By w. J.
KENNEDY and J. D. S. MACDOUGALL 459

New spiriferid brachiopods from the Lower Devonian of New South
Wales. By n. m. savage 472

A Cretaceous echinoid with false teeth. By p. m. kier 488

Ceratocystis perneri Jaekel — a Middle Cambrian chordate with echino-
derm affinities. By r. p. s. jefferies 494

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COUNCIL 1969-70

President: Professor Alwyn Williams, The Queen’s University, Belfast
Vice-Presidents : Dr. W. S. McKerrow, Department of Geology, Oxford
Dr. C. Downie, The University, Sheffield

Treasurer: Dr. J. M. Hancock, Department of Geology, King’s College, London, W.C. 2
Membership Treasurer: Dr. A. J. Lloyd, Department of Geology, University College, Gower

Street, London, W.C. 1

Secretary: Dr. W. D. I. Rolfe, Hunterian Museum, The University, Glasgow, W. 2

Editors

Mr. N. F. Hughes, Sedgwick Museum, Cambridge
Dr. Gwyn Thomas, Department of Geology, Imperial College, London, S.W. 7
Dr. Isles Strachan, Department of Geology, The University, Birmingham, 15
Professor M. R. House, The University, Kingston upon Hull, Yorkshire
Dr. R. Goldring, Department of Geology, The University, Reading, Berks.

Other members of Council

Dr. F. M. Broadhurst, Manchester
Dr. L. R. M. Cocks, London
Dr. C. B. Cox, London
Mr. D. Curry, Northwood
Dr. A. Hallam, Oxford
Dr. Julia Hubbard, London
Dr. J. D. Hudson, Leicester

Dr. W. J. Kennedy, Oxford

Dr. J. D. Lawson, Glasgow

Dr. E. P. F. Rose, London

Dr. C. T. Scrutton, Newcastle

Dr. V. G. Walmsley, Swansea

Professor H. B. Whtttington, Cambridge

Overseas Representatives

Australia: Professor Dorothy Hill, Department of Geology, University of Queensland, Brisbane
Canada: Dr. D. J. McLaren, Institute of Sedimentary and Petroleum Geology, 3303-33rd Street
NW., Calgary, Alberta

India: Professor M. R. Sahni, 98 The Mall, Lucknow (U.P.), India

New Zealand: Dr. C. A. Fleming, Geological Survey, P.O. Box 368, Lower Hutt, New Zealand
West Indies and Central America: Mr. John B. Saunders, Geological Laboratory, Texaco Trinidad
Inc., Pointe-a-Pierre, Trinidad, West Indies

Western U.S. A.: Professor J. Wyatt Durham, Department of Palaeontology, University of California,
Berkeley 4, California

Eastern U.S. A. : Professor J. W. Wells, Department of Geology, Cornell University, Ithaca, New Y ork

© The Palaeontological Association, 1969

A NEW FAMILY OF CARBONIFEROUS
AMPHIBIANS

by ROBERT L. CARROLL

Abstract. The morphology of a new Carboniferous amphibian, Acherontiscus caledomae, combines cranial
characteristics typical of lepospondyls with a vertebral structure resembling that of embolomeres. The skull is
relatively small and the trunk region quite long. Limbs are apparently poorly developed. This form cannot be
placed in any of the recognized amphibian orders, but presumably represents an isolated lineage which originated
prior to the establishment of the definitive characteristics which differentiate all known lepospondyls and
labyrinthodonts. Acherontiscus is designated the type of a new family, Acherontiscidae. This genus is based on a
single specimen from the Royal Scottish Museum which had no horizon or locality data. The matrix contains
spores which indicate a horizon from the upper part of the Visean to about the middle of the Namurian.

Since the publication of Zittel’s Handbuch der Pacelontologie (1890), it has been
generally accepted that two major groups of Palaeozoic amphibians could be recog-
nized: labyrinthodonts and lepospondyls. The groups included by him in the Lepo-
spondyli — microsaurs, nectrideans, and aistopods — had been recognized by the Miall
commission somewhat earlier, but at that time (1875) they were only vaguely defined
and not really distinguished from labyrinthodonts. Zittel’s grouping was based pri-
marily on the structure of the vertebrae. Labyrinthodonts (or, to use Romer’s (1933)
term applying to the vertebrae, apsidospondyls) have distinct ‘arch’ centra: posteriorly
the pleurocentrum, frequently paired, and anteriorly the intercentrum. These clearly
correspond to the vertebral components in amniotes. Lepospondyls have typically been
described as having holospondylous or husk vertebrae — a single central ossification for
each segment. The gross similarity between the structure in lepospondyls and that in
adult salamanders has led to the assumption that embryological development followed
a similar pattern in both groups, with direct ossification from the perichordal sheath.
The presumed distinction in the pattern of embryological development in the two
groups of Palaeozoic amphibians makes it difficult, if not impossible, to homologize
their components, although it has been suggested that the lepospondyl centrum was
comparable with either the pleurocentrum (Parsons and Williams 1963) or the inter-
centrum (Thomson 1967, Carroll 1967) of labyrinthodonts.

In addition to the vertebral pattern, a series of cranial features also distinguish laby-
rinthodonts and lepospondyls. Labyrinthodonts (discussed at length by Romer in 1947,
1963, and 1964) typically have labyrinthine infolding of the enamel, large fangs on the
ectopterygoids, palatines, and frequently the vomers, and typically parasymphyseal
tusks. They generally possess an otic notch dorsal to the quadrate, and the stapes is
directed laterally or dorso-laterally. Lepospondyls (reviewed by Baird 1965), in contrast,
lack labyrinthine infolding of the enamel and distinct fangs on the palatal bones. They
all appear to lack an otic notch. The stapes is directed ventro-laterally toward the quad-
rate. Lepospondyls (as most clearly shown in microsaurs, Carroll and Baird 1968) have
a very well-developed articulation between the occipital condyle and atlas-axis complex
in which the atlas fits into a large strap-shaped recess formed by the exoccipitals and

[Palaeontology, Vol. 12, Part 4, 1969, pp. 537-48]

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basioccipital. Although the articulation is not as well defined in labyrinthodonts, the
occipital condyle is typically a knob-shaped structure which fits into a ring formed by
the atlas intercentrum, pleurocentrum, and paired arches. The latter pattern becomes
better defined in early reptiles.

In addition to these fairly clearly defined anatomical differences, lepospondyls can
also be characterized by their tendency to be small, aquatic forms, many of which have
feeble limbs. In contrast to labyrinthodonts, lepospondyls are not known to have a
distinct larval stage exhibiting external gills.

It has been possible to fit all adequately known Palaeozoic amphibians into one or
other of these major taxonomic categories. The distinctions appear complete even in
early Mississippian lepospondyls (notably the ai'stopods, Baird 1964) and the Upper
Devonian ichthyostegids (Jarvik 1952). Despite the obvious distinctions between laby-
rinthodonts and lepospondyls, however, it is considered that they have a common
ancestry, for, to quote Baird (1965, p. 293): ‘a duplicate origin of the tetrapod appen-
dicular skeleton is simply incredible.’

The reason for reviewing the established classification of Palaeozoic amphibians is the
discovery of a single specimen from the collections of the Royal Scottish Museum which
appears to combine the characteristics of both labyrinthodonts and lepospondyls. This
specimen, R.S.M. no. 1967/13/1, was discovered in a search for lepospondyls and early
reptiles made in 1964. The specimen appeared to resemble the pattern of typical micro-
saurs such as Microbrachis and Hyloplesion in having a small head and an elongate
body, with little or no evidence of limbs. The skull grossly resembled that of gym-
narthrids in having the orbits far forward and in possessing a small number of blunt
cheek teeth. As originally preserved, the post-cranial skeleton showed very few details.
The importance of the specimen was not recognized until a cast was made which
revealed a series of vertebrae preserved as impressions. Instead of a series of single,
elongate centra, as in microsaurs, the vertebrae were clearly formed on the pattern of
embolomeres, with two rather similar, spool-shaped centra per segment. The specimen
obviously belongs not only to a new genus, but also to an entirely new lineage, other-
wise unreported in the fossil record of the Carboniferous.

The purpose of this paper is to describe this particular specimen, and to discuss the
current concept of the classification of the lepospondyls in light of its anatomy.

Class AMPHIBIA

Subclass and Order Undesignated

ACHERONTISCIDAE llOV.

Diagnosis. Small stegocephalian amphibia with both pleurocentra and intercentra well-
developed cylinders. Skull with lateral line canals, orbits far forward, no otic notch,
teeth without labyrinthine infolding of enamel. Dermal pectoral girdle well developed.
Long trunk region.

acherontiscus gen. nov.

Type species. Acherontiscus caledoniae gen. et sp. nov.

Diagnosis. Same as for family. The generic name follows Cope’s practice of naming
serpentiform lepospondyls for tributaries of the Styx: Cocytinus, Phlegethontia, etc.

CARROLL: A NEW FAMILY OF CARBONIFEROUS AMPHIBIANS 539

Acherontiscus caledoniae gen. et sp. nov.

Holotype. 1967/13/1 in the Royal Scottish Museum, Edinburgh. Skull and associated postcranial
skeleton. This is the only known specimen.

Locality and horizon. The specimen bears no data as to horizon, locality or collector. The matrix is a
fine-grained coal shale which resembles that of amphibians from the Loanhead No. 2 Ironstone of the
Limestone Coal Group at Burghlee, Midlothian, which is lower Upper Carboniferous in British terms
and equivalent to the Continental Namurian A and the American Upper Mississippian. Ostracods
preserved with the specimen have been variously identified. According to Dr. Robinson, University
College, University of London, all are assignable to Carbonita ( Carbonia ) fabulina , which inhabits coal
shale facies from the Visean to the Coal Measures. Dr. Pollard, at Manchester, identifies most as
Carbonita humilis, and others dubiously as Carbonita infiata. He suggests that these specimens may
indicate Coal Measures, most likely Westphalian A or B.

Dr. A. H. V. Smith of the National Coal Board examined a piece of the matrix and reported that it:
‘contained an assemblage of spores rich in species including such forms as Cingulizonates cf capistratus,
Rotaspora knoxi and Tripartites trilinguis. By considering the stratigraphic ranges of all the species
recovered from the sample, it is possible to assign the miospore flora to a horizon within Assemblage
III of Smith and Butterworth 1967. In Scotland this Assemblage ranges from the upper part of the
Lower Limestone Group, through the Limestone Coal Group to upper part of the Upper Limestone
Group. In terms of the Heerlen classification, these lithological divisions range from Upper part of
Visean to about the middle of the Namurian. . . . The horizon is definitely not Coal Measures.’
Athough there is some question as to the exact age of this specimen, it is of sufficient anatomical
significance to warrant description.

Description. The skull is exposed primarily from the right side. It is flattened, with much
of the original bone surface badly damaged. In order to determine the position of the
sutures more accurately, an attempt was made to etch away the bone so as to expose the
impression of the ventral surface. The skull roof and lower jaws were crushed so closely
together that what matrix there may have been between them was lost in the etching of
the bone. This led to the etching of all three bone layers. It was hence felt better to save
the poorly exposed dorsal surface, rather than to attempt further preparation.

To judge from the apparently complete right lower jaw, almost the entire length of
the skull is preserved. The premaxillae are missing, however, and possibly small areas
of the nasal and lacrimal bones. These bones appear to extend almost to the external
naris, but the posterior margin of this opening is not clearly defined. As preserved, the
skull is 16-5 mm. in length. In contrast to typical labyrinthodonts, the orbit is very
small (approximately 2-4 mm. in diameter) and is located far anteriorly (its posterior
margin is 10-5 mm. from the rear margin of the skull). Where the original surface of the
bone is preserved, the posterior roofing bones are sculptured with shallow, irregular
pits, somewhat as in the microsaur Tuditanus. The more anterior bones, particularly the
frontal, are essentially smooth. Lateral line canals are evident on the supra-temporal,
post-orbital, prefrontal, and lacrimal. What is preserved of this system suggests an
arrangement typical of that of aquatic labyrinthodonts. As in the microsaur Micro-
brachis, the posterior portion of the skull is very wide.

The crushing and loss of much of the original surface makes it difficult to substantiate
the pattern of the dermal bones. Three bones may, nevertheless, be fairly clearly dis-
tinguished across the back of the skull. These may be designated as supra-temporal,
squamosal, and quadratojugal. One important point that can be safely established is
the large size of the supra-temporal, since much of the surface of this bone is preserved.
Such a large bone in this position follows the pattern of microsaurs and is distinct from
that of labyrinthodonts. Although poorly preserved, the posterior margin of the cheek

540

PALAEONTOLOGY, VOLUME 12

C

1 cm

‘==3*?

text-fig. 1. Acherontiscus caledoniae. A. Skeleton, skull and dermal shoulder girdle drawn as a mirror
image; postcranial skeleton drawn from rubber mould. Numbers indicate vertebral count, x 1
b. Skull and dermal shoulder girdle. X 3. c. Restoration of skull. X 3. d. Single ventral scale in

medial view, x 12.

CARROLL: A NEW FAMILY OF CARBONIFEROUS AMPHIBIANS 541

region appears to have been nearly vertical, without an otic notch. The anterior extent
of the supra-temporal, squamosal, and quadratojugal is difficult to ascertain, but they
may be reconstructed as meeting the jugal and post-orbital in the manner of Micro-
brachis (Steen 1938). The mid-dorsal region of the skull is not well preserved posteriorly;
there may have been small postparietals as in Microbrachis. The parietal appears to
extend a lappet laterally between the supra-temporal and post-frontal; the area of the
pineal foramen is missing. The left post-frontal, post-orbital, and jugal are disarticu-
lated and lie behind the remainder of the skull. The post-frontal is shown as an im-
pression of its ventral surface, the post-orbital as an impression of the lateral surface,
and the jugal is exposed medially. The latter shows very large areas of overlap with the
post-orbital, squamosal, and quadratojugal. The portion of these bones entering the
margin of the orbit seems to be somewhat different from those of the corresponding
bones preserved on the right side, but this is not surprising considering the extent of
overlap shown by the jugal. The frontal appears much longer than the nasal, but the
former bone was probably overlapped quite extensively by the parietal. Where the sur-
face is preserved, the maxilla appears smooth. The upper dentition is obscured by that
of the lower jaw. Neither the palate nor the braincase is exposed, nor can these structures
be readily prepared without danger to the skull roof.

The right jaw, which is displaced slightly posteriorly, reaches just beyond the posterior
margin of the cheek. The anterior end narrows, suggesting that only a very short
portion has been lost. Like the other bones, those of the lower jaw have lost most of their
surface. It remains only on the antero-ventral margin of the dentary, which shows no
sculpturing. No lateral line canal grooves are preserved on the lower jaw. Including the
impressions of the anterior teeth, 16 are present in the lower jaw, with room for at least
2 more. The posterior 7 teeth are bluntly-rounded cones, while those more anterior
are slim and sharply pointed. The posterior teeth were ‘sectioned’ by some earlier pre-
parator, showing that the pulp cavity is very large and that the enamel is definitely not
infolded. The tips of the posterior teeth are not well exposed in the right jaw, but those of
the left can be seen protruding through the skull roof in the area of the post-orbital.
They are laterally compressed and marked by vertical ridges, much as the teeth of
Cardiocephalus (Gregory, Peabody, and Price 1956, p. 18). They show little, if any, wear.
There is no evidence for more than a single row of marginal teeth. The posterior portion
of the lower jaw is very deeply worn, making it impossible to establish sutures between
the dentary, angular, and surangular. The medial surface of the left jaw is exposed
behind the skull. It is very poorly preserved. The articular surface appears to be at the
level of the posterior end of the jaw, with no retro-articular process.

Immediately behind the skull are a number of bones which presumably are remains of
the visceral arch apparatus. They resemble in general the bones described by Sollas
(1920, p. 513, fig. 39) in Lysorophus, but their disarticulation in Acherontiscus precludes
homologizing the individual elements.

What is visible of the structure of the skull resembles in general that of lepospondyls
such as the ‘typical’ microsaurs Microbrachis and Cardiocephalus. It shows none of the
characteristics expected in labyrinthodonts, such as infolding of the enamel, multiple
bones in the temporal region, or an otic notch.

Aside from the dermal shoulder girdle, the post-cranial skeleton of this animal was
prepared by etching away the bone with hydrochloric acid and casting the rssulting

542

PALAEONTOLOGY, VOLUME 12

impressions with silicone rubber. This results in the post-cranial skeleton being viewed
from the opposite side from the skull, and explains the necessarily composite nature of
text-fig. 1 . This method of preparation was necessary since the bone surface as exposed
was so damaged and weathered that very little structure could be determined.

The vertebral column is visible primarily in ventral view, with the neural arches only
occasionally exposed. The centra, in contrast, are readily seen for much of the length of

text-fig. 2. Acherontiscus cciledoniae. Postcranial skeleton drawn in three sections. X 3.

the column. It is immediately apparent from their configuration and from the arrange-
ment of the few neural arches present, that each vertebra consists of two centra, typi-
cally complete cylinders, which may be compared with the intercentra and pleurocentra
of embolomeres and other labyrinthodonts. There is absolutely no question of the
association of the skull and these vertebrae, which demonstrates a combination of
lepospondyl cranial features and apsidospondylous vertebral structure in a single form.

The first segment has only a single centrum. Posterior to this, some 32 central pairs
are exposed in sequence. More posteriorly, the column is intermittently visible, with

CARROLL: A NEW FAMILY OF CARBONIFEROUS AMPHIBIANS 543

much of it represented only by a depression in the matrix. A few vertebrae are more
clearly exposed at the extremity of the column. The entire trunk region and most, if not
all, of the tail were apparently in place, with the elements quite well articulated, when
the animal was buried. By extrapolation from the well-preserved sections, a total of
approximately 64 vertebrae were present. The posterior 3 or 4 show haemal arches and
so are definitely caudals. More caudal vertebrae may originally have been present, but
the series ends without reaching the margin of the block. Bones which may represent
elements from the pelvic girdle and rear limb are present in the area from the 26th
through the 31st vertebra. This suggests approximately 27 presacrals, although this is
only very tentatively established. At least as far back as the 36th vertebra the intercentra
are not modified to support haemal arches, although poorly preserved structures which
might be so identified are present in the area of approximately the 40th vertebra.

There is some regional variation in the centra. There was apparently only one element
in the 1st segment, a narrow crescentic pleurocentrum. It would be very interesting to
know the manner of articulation between the anterior cervicals and the brain-case, but
it is impossible to determine this on the basis of this skeleton as it is preserved. The 2nd
segment consists of a crescentic intercentrum, which bears facets for the articulation of
the capitulum of the first cervical ribs, and a rather short pleurocentrum. Apparently the
first 4 pleurocentra are crescentic rather than cylindrical, judging from the 4th which is
disarticulated. The 5th and those more posterior are complete cylinders, but retain a
large passage for the notochord. All (except the two most anterior) are considerably
longer than the intercentra. The 6th intercentrum is displaced and evidently crescentic.
The 25th, 26th, and 27th intercentra are visible end-on and are cylindrical, with most, if
not all, of the notochordal canal closed. With the centra oriented as they are, it is not
possible to judge where, between the 6th and 25th segments, they become cylindrical. In
addition to their short length, the intercentra are characterized by the presence of facets
for the articulation of the capitulum of the ribs. Dorsal to the area for rib articulation,
the surface of the intercentra is of unfinished bone, presumably marking the area for
articulation with the neural arch.

Both pleurocentra and intercentra are marked laterally and ventrally by a regular
pattern of deep pits, giving them a marked resemblance to those of embolomeres. The
posterior caudal vertebrae differ from those in the trunk region in being less deeply and
regularly pitted, and seem to have rather thin walls. The intercentra more closely ap-
proach the length of the pleurocentra in this region. The pleurocentra, and apparently
the intercentra as well, are complete cylinders as far posteriorly as the tail is preserved.
The haemal arches appear to articulate with rather than being solidly attached to the
intercentra.

Poorly preserved neural arches are present on the 26th, 30th, and 31st vertebra. The
most posterior is visible ventrally, showing both posterior zygapophyses. The two halves
of the arch are completely fused. The neural spines appear to be short and located far
posteriorly. The pedicle of the arch and the transverse process are never well exposed.
Where visible, the arches are disarticulated from the centra. It is probable from the
structure of the centra that, as in embolomeres, they were never solidly attached.

Fragments of ribs are visible throughout the column; none are well preserved, and the
articulating surfaces appear incompletely ossified. Since there are well-ossified facets for
the articulation of the capitulum on the intercentrum, it is probable that the ribs were

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PALAEONTOLOGY, VOLUME 12

double headed in the manner of embolomeres and most early reptiles and typical micro-
saurs. Their length can be nowhere adequately established, but there does not appear to
be any marked modification in length in the suggested area of the pelvic girdle, except
for the ‘sacral ribs’ themselves. Structures which appear to be quite long ribs extend nearly
to the end of the vertebral column.

The dermal shoulder girdle is represented by the interclavicle, in the form of a large
oval plate, and the clavicles. The left clavicle is complete, with a narrow, unsculptured
blade, and a short, thick stem. The interclavicle is marked ventrally by a very low
median ridge, and fine radiating grooves. It is complete posteriorly and definitely has no
stem. The anterior margin is covered by the clavicles, preventing determination of the
presence of a fimbrilated margin. Neither the cleithra nor the endochondral shoulder
girdle is visible.

No elements identifiable as belonging to the fore limb are visible. Considering the
size of the bones of the dermal shoulder girdle, it would be most surprising if this animal
lacked forelimbs. They may, however, have been small and/or poorly ossified, or lost
prior to burial.

Nothing can be very confidently identified as representing the pelvic girdle and rear
limb. There are a few elements in the area of the 26th through 31st vertebrae which are
definitely not normal ribs or vertebral elements. Two, apparently paired, blocks are
adjacent to the 28th vertebra. They might represent remnants of the pelvic girdle, but
they do not compare with any bones described from other Paleozoic tetrapods. They
are quite thick and well ossified, except for margins which appear to be surfaces of
articulation. They might conceivably be sacral ribs. A pair of bones reasonably identi-
fiable as limb elements is found adjacent to the 26th and 31st vertebra. Each is approxi-
mately the length of a single segment. They resemble in a vague way the tibia of other
Palaeozoic tetrapods, but, in the absence of other evidence, they could as well be the
femora of this animal. Some of the bones in this region, otherwise accepted as ribs, may
be part of the appendicular skeleton. It is unfortunate, in the light of the very interesting
evidence of the axial skeleton, that so little of the appendicular skeleton is preserved in
this animal.

Numerous fragmentary scales are associated with Acherontiscus. Most are poorly
preserved, represented by roughly oval patches of fine parallel rods, resembling in a
general way the dorsal scales of microsaurs. One scale, shown in text-fig. Id, is almost
complete and resembles closely the ventral scales of microsaurs, as viewed medially
(Carroll and Baird 1968, fig. 20). Although more closely resembling those of micro-
saurs, the scales of Acherontiscus also resemble in a general way those of such laby-
rinthodonts as Trimerorhachis (Colbert 1955).

The presence of lateral line canals indicates that Acherontiscus was primarily aquatic.
This habit would explain the small size and incomplete ossification of the elements of
the pelvic girdle and rear limb (assuming that they have been correctly identified).
Such poor ossification might also be attributed to immaturity. Judging from the solid
attachment of the bones of the skull and the degree of ossification of the vertebrae, this
animal appears to be essentially mature. In lepospondyls in general, however, there is
very little difference in the anatomy of individuals of different size within a given texon.

The fairly long trunk region, as well as the small size of the skull, suggests a snake-like
habitus. As has been suggested by Panchen (1966) and Parrington (1967, p. 277), the

CARROLL: A NEW FAMILY OF CARBONIFEROUS AMPHIBIANS 545

development of an embolomerous or lepospondylous vertebral structure are both
methods of lengthening the segments so as to assist a sinuous type of locomotion,
usually associated with an aquatic habit and anguilliform swimming.

Taxonomic position. The significance of this specimen is its combination of cranial
features accepted as typifying lepospondyls with a vertebral structure closely resembling
that of embolomerous labyrinthodonts. This combination makes classification of this
particular genus difficult, but contributes to our understanding of vertebral homologies
among other Palaeozoic amphibians.

The great range in vertebral structure among the labyrinthodonts indicates that this
feature alone is not a sufficient basis for classification within that group. Within both
the temnospondyls and anthracosaurs the pleurocentrum varies from being a major
structural element to being little more than an accessory. The relative importance of the
intercentrum likewise differs greatly within each group. Although such a range in
vertebral structure has not been recognized among the lepospondyls, the presence of
haemal arches in Pantylus and Lysorophus (Carroll 1968) indicates that multipartite
vertebral centra do occur within that group.

The pattern of the bones of the skull roof appears a much more valid basis for classi-
fying both labyrinthodonts and lepospondyls. Ichthyostegids, anthracosaurs, and tem-
nospondyls can all be defined on the basis of the relative position of the bones in the
temporal series (Romer 1947). No labyrinthodonts are known which have a pattern
which could be confused with that of any described lepospondyl. The patterns within
each of the lepospondyl groups seem similarly stereotyped. All typical microsaurs have a
particular pattern (Carroll and Baird 1968) as do the better-known lysorophids and
adelogyrinids (Carroll 1967).

The pattern of the skull roof of Acherontiscus resembles most closely that of the
microsaur Microbrachis. The absence of an otic notch and labyrinthine infolding of the
enamel support association with lepospondyls, as do the general body proportions. This
genus differs from all labyrinthodonts in these features. The presence of multipartite
centra indicates closer association with microsaurs and lysorophids than with aistopods
or nectrideans, but no more specific assignment of this genus among the lepospondyls is
possible. The dentition is similar to that of the gymnarthrid microsaurs, but this prob-
ably does not indicate any particularly close relationship. The fact that Acherontiscus is as
old or older than any of the known microsaurs makes it a possible ancestor to some or
all members of that group, but there is no very convincing evidence for this while the
well-developed embolomerous vertebral pattern suggests that it diverged from the
lineage leading toward microsaurs at a stage when neither had yet developed its defini-
tive vertebral pattern.

On the basis of our present knowledge of the one described genus, the family Acheron-
tiscidae should be recognized as representing an isolated lineage, of equivalent rank with
nectrideans, aistopods, lysorophids, adelogyrinids, and typical microsaurs.

DISCUSSION

Acceptance of Acherontiscus as a lepospondyl raises several problems in regard to our
concept of that group. This genus provides the first conclusive evidence of the presence
of multiple central elements in the trunk region. There seems no reason to argue against

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PALAEONTOLOGY, VOLUME 12

the obvious identification of these structures as intercentra and pleurocentra, or to
reject their general homology with their counterparts in labyrinthodonts. The presence
of haemal arches in Pantylus (Carroll 1968) and ‘ Hylonomus' fritschia (Credner 1885)
supports identification of the major central elements as pleurocentra in microsaurs as
well, although trunk intercentra have never been conclusively demonstrated in this
group. Haemal arches, but not trunk intercentra, are also present in the lysorophids
Lysorophus (Carroll 1968) and Molgophis. It seems plausible to assume that primitive
microsaurs and lysorophids derived their vertebral structure from an apsidospondylous
pattern. Unfortunately no members of either group are known prior to the Lower
Pennsylvanian, by which time all genera had lost all trace of trunk intercentra and most
typical microsaurs had eliminated the haemal arches as well. Acherontiscus, from the
earlier Carboniferous, appears to have retained a more primitive configuration.

If the central elements in typical microsaurs, lysorophids and acherontiscids can be
homologized with those of labyrinthodonts, can these forms be considered as lepo-
spondyls ? The term lepospondyl may be used in a descriptive manner, as does Romer in
1966: ‘. . . the centrum forms as a single, spool-shaped bony cylinder around the noto-
chord . . (p. 96), or to imply a particular mode of embryonic development \ . . in

which arch-centra preformed in cartilage do not occur; instead, the centrum forms
directly as a bony cylinder around the notochord’ (Romer 1945, pp. 157-8). In terms of
adult structure, there is no evidence for more than a single vertebral element in either
the trunk or tail of any nectridean or aistopod, although the vertebral anatomy is well
known. In both groups, outgrowths from the major central elements in the tail act as
haemal arches, precluding the presence of a separate element with this function. What-
ever their homology, the vertebrae in these forms are structurally lepospondylous.

At present we have no way of knowing whether the intercentra and pleurocentra of
Acherontiscus are preformed in cartilage, or ossify directly as bony sheaths around the
notochord. It is possible that the retention of multipartite central elements indicates
retention of the primitive developmental pattern, but there is no proof of this.

Unless one assumes that the developmental pattern in nectrideans, and presumably
ai'stopods, implies a separate origin for the vertebrae from that described for laby-
rinthodonts and other lepospondyls, the centra in all amphibian groups must be
generally homologous. This implies a suppression of the normal cartilaginous precursors
of the centra in nectrideans, if not in other forms which are structurally lepospondylous.
It seems probable that the small size of the vertebrae and their simple structure in the
adult made such abbreviated development possible.

If the vertebral structure and developmental pattern in nectrideans was initially
derived from the rhipidistian or labyrinthodont condition, then the adult configuration
in Acherontiscus , lysorophids, and microsaurs can be accepted as representing slightly
more primitive stages in the same general pattern. In all but Acherontiscus, there is a
strong tendency to develop a single central element. On this basis, all of these forms may
be retained within the Lepospondyli, whatever the pattern of embryonic development.

Despite the common tendency toward a holospondylous vertebral structure, small
size, and aquatic habit, there is little convincing evidence that all of the groups accepted
as lepospondyls share a single common ancestry separate from the labyrinthodonts
(Parrington 1967). Whatever the developmental pattern, the adult structure of the
vertebrae clearly separates ai'stopods and nectrideans from microsaurs, lysorophids, and

CARROLL: A NEW FAMILY OF CARBONIFEROUS AMPHIBIANS

547

acherontiscids. The morphology of each group is stereotyped and distinctive throughout
its known fossil record. They are united primarily by their common distinctiveness
from labyrinthodonts. It is quite conceivable that all of the known lineages evolved
separately from primitive labyrinthodonts or their ancestors among the rhipidistians.

While knowledge of Acherontiscus appears to confirm the homology of the major
central elements in microsaurs and lysorophids with the pleurocentra in labyrinthodonts
it contributes little to our understanding of the vertebrae in nectrideans or aistopods and
emphasizes the difficulties in classifying the lepospondyls in general.

Acknowledgements. I wish to thank Dr. Waterston and Dr. Miles of the Royal Scottish Museum for
permission to borrow, prepare and describe Acherontiscus. Dr. E. Robinson, University College
London and Dr. Pollard, Manchester University, were very helpful in identifying the ostracods
associated with this specimen. Particular thanks are due to Dr. A. H. V. Smith of the National Coal
Board for identification of the associated spores. Dr. Baird provided much useful information con-
cerning the ontogeny of labyrinthodonts and lepospondyls and many stimulating comments as to the
taxonomic position and anatomical significance of this genus.

REFERENCES

baird, d. 1964. The ai'stopod amphibians surveyed. Breviora , 206, 1-17.

1965. Paleozoic lepospondyl amphibians. Am. Zool. 5, 287-94.

carroll, r. l. 1967. An adelogyrinid lepospondyl amphibian from the Upper Carboniferous. Can. J.
Zool. 45, 1-16.

1968. The postcranial skeleton of the Permian microsaur Pantylus. Ibid. 46, 1175-92.

and baird, d. 1968. The Carboniferous amphibian Tuditanus [ Eosauravus ] and the Distinction

between Microsaurs and Reptiles. Am. Mas. Novit. 2337, 1-50.
colbert, e. h. 1955. Scales in the Permian amphibian Trimerorachis. Ibid. 1740, 1-17.
credner, h. 1885. Die Stegocephalen aus dem Rothliegenden des Plauenschen Grundes bei Dresden.
V Theil. Z. dtsch. geo/. Ges. 37, 694-736.

Gregory, j. t., peabody, f. e., and price, l. i. 1956. Revision of the Gymnarthridae, American Permian
microsaurs. Bull. Peabody Mas. 10, 1-77.

jarvik, e. 1952. On the fish-like tail in the ichthyostegid stegocephalians. With descriptions of a new
stegocephalian and a new crossopterygian from the Upper Devonian of East Greenland. Meddr
Gran land 114, 1-90, 21 pis.

miall, l. c. 1875. Report of the Committee, consisting of Professor Huxley, Professor Harkness,
F.R.S., Henry Woodward, F.R.S., James Thomson, John Brigg, and L. C. Miall, on the structure
and classification of the Labyrinthodonts. Drawn up by L. C. Miall, Secretary to the Committee.
Rep. Br. Ass. Advmt. Sci. 149-92, pis. iv-vii.

panchen, a. l. 1966. The axial skeleton of the labyrinthodont Eogyrinus attheyi. J. zool. Lond. 150,
199-222.

parrington, f. r. 1967. The vertebrae of early tetrapods. Colloques internationaux du Centre National
de la Recherche Scientifique, No. 163, Problemes actuels de paleontologie (evolution des vertebres),
269-79.

parsons, t. s. and williams, e. e. 1963. The relationships of the modern Amphibia: A re-examination.
Q. Rev. Biol. 38, 26-53.

romer, a. s. 1933. Vertebrate paleontology. 1st ed. Chicago.

1945. Ibid. 2nd ed. Chicago.

1947. Review of the Labyrinthodontia. Bull. Mus. comp. Zook Harv. 99, 3-352.

1963. The larger embolomerous amphibians of the American Carboniferous. Ibid. 128, 415-54,

pis. 1-2.

1964. The skeleton of the Lower Carboniferous labyrinthodont Pholidogaster pisciformis. Ibid.

131, 129-156, pi. 1.

1966. Vertebrate Paleontology. 3rd ed. Chicago.

548

PALAEONTOLOGY, VOLUME 12

smith, a. h. v. and butterworth, m. a. 1967. Miospores in the coal seams of the Carboniferous of
Great Britain. Spec. Paper Palaeont. 1.

sollas, w. j. 1920. On the structure of Lysorophus as exposed by serial sections. Phil. Trans. R. Soc.
Ser. B , 209, 481-527.

steen, m. 1938. On the fossil Amphibia from the Gas Coal of Nyrany and other deposits in Czecho-
slovakia. Proc. zool. Soc. Load. 108, 205-83.

Thomson, k. s. 1 967. Notes on the relationships of the rhipidistian fishes and the ancestry of the tetrapods.
J. Paleont. 41, 660-674.

zittel, K. A. 1 890. Handbuch der Palceontologie. Ill Band. Vertebrata ( Pisces , Amphibia , Reptilia, Aves).
Miinchen und Leipzig.

R. L. CARROLL
Redpath Museum
McGill University

Typescript received 11 February 1969 Montreal 110, P.Q.

A FA VREINA-THALASSINOIDES ASSOCIATION
FROM THE GREAT OOLITE OF OXFORDSHIRE

by W. J. KENNEDY, M. E. JAKOBSON, and R. T. JOHNSON

Abstract. The crustacean microcoprolite, Favreina decemlunulatus (Parejas) is present in several beds of the
Great Oolite Series at Kirtlington, Oxfordshire. It occurs in burrows of Thalassinoides type and was probably
produced by a brachyuran crustacean. It is associated with a molluscan and annelid infauna in bioturbated
sediments that have been interpreted as accumulating intertidally or just below low water mark (McKerrow
et al. 1969).

The ichnogenus Favreina Bronniman 1955 covers a variety of rod-like microfossils
perforated by many fine calcite-filled longitudinal canals. Objects of this type have been
described from many parts of the world, i.e. the Oligocene of Turkey (Altini, 1942),
Jurassic of Switzerland (Joukowsky and Favre 1913), French Mesozoic (Cuvillier and
Sacal 1956), Upper Jurassic and Lower Cretaceous of Cuba (Parejas 1948, Bronniman
1955), and the Triassic to Miocene of the Middle East (Elliot 1962, 1963).

Recognition of the true nature of these objects, as crustacean faeces, was first made by
Parejas (1935, 1948) from the work of Moore (19326) on recent forms.

We describe here the ichnogenus for the first time from Britain, on the basis of material
from the Great Oolite Series (Jurassic, Bathonian) White Limestone and Forest Marble
at Kirtlington Old Cement Works, Oxfordshire (National grid reference SP. 494199).
This quarry has been described elsewhere (McKerrow et al. 1969).

The only previous records of faecal pellets of this type in Britain are from the Chalk
(i.e. Type A faecal pellet of Wilcox 1953; see also Kennedy 19676, p. 137, and Bromley
1967, p. 172), although unpublished observations suggest they occur at other localities
in the Great Oolite and at other horizons in the Mesozoic.

SYSTEMATIC DESCRIPTION

Ichnogenus favreina Bronniman 1955
Ichnospecies Favreina decemlunulatus (Parejas) 1948

1948 Coprolithus decemlunulatus Parejas, p. 519, figs. 46- 8.

Emended diagnosis. Favreina with ten crescentic longitudinal canals. The pellet is orna-
mented externally by V-shaped transverse grooves.

Description of the Kirtlington specimens. The pellets are rod-shaped, up to 1 cm. long,
1 mm. in diameter, and circular in cross-section.

Externally (PI. 99, fig. 2), there is a distinctive ornament of V-shaped transverse grooves.
In section (PI. 99, fig. 1, text-fig. 1), there is a distinct separation of the pellet into two
zones, a central core and a surrounding envelope. The core consists of fine-grained
carbonate sediment, the envelope of fine-grained calcite. They are distinguishable as a
result of slight colour differences, the envelope being lighter than the core. The envelope
may also be detached from the core around part of the pellet (PI. 99, fig. 1).

[Palaeontology, Vol. 12, Part 4, 1969, pp. 549-54, pi. 991

550

PALAEONTOLOGY, VOLUME 12

There are ten crescentic calcite-filled canals in the core. They are arranged symmetri-
cally into an outer ring of eight about a central pair (PI. 99, fig. 1, text-fig. 1).

Discussion. Favreina decemlunulatus can be separated from all previously described
Favreina and Favreina- like pellets by virtue of the form, number and arrangement of
the canals (see Table 1 in Elliot 1962, also Elliot 1963, p. 299). It is also the only Favreina
in which an external ornament has been noted.

Interpretation and occurrence. Faecal pellets of recent invertebrates have been described
by a number of workers, i.e. Moore (I931u, b, 1932 a, b) and Edge (1934).

It is clear from these works that only one group of animals, brachyuran crustaceans,
could produce faecal pellets of Favreina type (Parejas 1948, Bronniman and Norton

text-fig. 1. Diagrammatic cross-section of Favreina decemlunulatus.

Actual diameter of pellet is 1 0 mm. approx.

1961, Elliot 1962, 1963). Here, material is passed down the stomach, and canals are pro-
duced by a system of fleshy processes which project inwards from the stomach wall.
These processes are at first flanges, but traced posteriorly, they become detached from
the stomach wall, projecting as fleshy cylinders.

As matter passes down the gut it is compacted around these cylinders, and faeces are
voided with minute canals inside. When fossilized, these canals become filled with
calcite, thus preserving traces of the passage of the soft parts of the animal concerned.

In the recent crustaceans, there is variation in the type of pellet produced, in particular
in the number and distribution of canals. This is a reflection of specific and generic
differences in stomach structure. In particular the Galatheidae produce a pellet which
has a well-differentiated ventral cap. We had thought to interpret the separation of the
Kirtlington pellets into core and envelope as a comparable phenomenon, presumably
the result of a comparable sorting mechanism in the Jurassic animal, but Dr. A. Kendall
(Reading University) has told us that petrographic studies on the White Limestone of
the Cotswolds show that such envelopes occur on non-skeletal grains. He suggests that

EXPLANATION OF PLATE 99

Fig. 1. Thin section of part of a Favreina packed burrow, X 6.

Fig. 2. Fractured surface of a Favreina packed burrow showing external ornament of pellets, X 6.

Palaeontology, Vol. 12

PLATE 99

KENNEDY, JAKOBSON and JOHNSON, Favreina

KENNEDY, JAKOBSON, AND JOHNSON: FAVREIN A-TH A LA SS IN O I D ES 551

these envelopes are a diagenetic replacement of organic sheaths. In the case of Favreina,
the sheath was probably a mucilaginous coat, present at the time of extrusion, but now
replaced by calcite.

The remarkable ornament present on the ‘solid’ Jurassic specimens of F. decemlunu-
latus (the original account is based on sections of Oligocene material) is the first record
of such structure in Favreina.

Pellets of recent brachyurans appear to be smooth; the ornament of our material may
have been produced in the gut of the animal, but it seems more likely that it is the result

50 cm

text-fig. 2. Sketch of Thalassinoides burrow systems on bottom surfaces of
limestones from the basal Forest Marble, Kirtlington, Oxfordshire.

of a rhythmic contraction of the anus during defecation. Miller (1910) records such
contractions in related Crustacea.

F. decemlunulatus ranges throughout the whole of the White Limestone at Kirtlington
although it is absent in some beds. It also occurs at the base of the Forest Marble (beds
3k, 4d, 6e, McKerrow et al. 1969).

Two modes of occurrence can be recognized, as loose pellets in sediment, and as
closely packed masses which are clearly filled burrows (PI. 99, figs. 1-2). In section, these
burrows are elliptical, compressed parallel to the bedding, and have a breadth of from
2 to 5 cm. On bottom surfaces the burrows weather out, and have a striking polygonal
branching form (text-figs. 2a, b).

The burrows lie horizontally, and branch at 5-15 cm. intervals, widening at the point
of branching. Several levels of burrowing can be recognized, but burrows rarely cut each
other; the sediment is not intensively disturbed.

These burrows can be referred to Thalassinoides. This ichnogenus was erected by
Ehrenberg (1944), for ramifying burrow systems with Y-shaped branching points and
local swellings, described by him from Miocene sands. These burrows were intimately
associated with remains of Callianassa.

552

PALAEONTOLOGY, VOLUME 12

Subsequently, these burrows have been recorded from many horizons and lithologies;
in the British Mesozoic they have been recorded by Hallam (1961), Farrow (1966),
Kennedy (1967 a , b ), and Bromley (1967). There is no doubt that these are crustacean
burrows. In the Chalk they have been described associated with ‘anomuran’ faecal
pellets (Kennedy 19676, Bromley 1967), and inferred to be the products of sediment
eating crustaceans.

This is also our interpretation of the Favreina-Thalassinoides association described
here, although the dense packing of the burrows is unusual.

The only comparable examples of this type of which we are aware are Ophiomorpha
(‘ Hcilymenites ’) burrows described by Brown (1939). These too are callianassid burrows
(Hantzschel 1952), and the pellets appear to be of Favreina type.

That these pellets belong to some organism other than the excavator of the burrows is
possible, but we see no evidence for this, i.e. traces of re-working within burrow fills.

The explanation of the filling is far from clear. The nature of the packing is such as to
preclude a passive filling by pellets washing into burrows by current action. It seems
that the animals stuffed their burrows with pellets, rather than moving them some dis-
tance to eject them at the surface.

CONCLUSIONS

Favreina and Thalassinoides occur associated in the Great Oolite Series at Kirtlington,
Oxfordshire. This is in keeping with previous suggestions that both trace fossils are
produced by anomuran crustaceans.

As with most other occurrences of these trace fossils (Kennedy 1967 b) actual crusta-
cean skeleta are rare in associated sediments. Three brachyurans are described from the
Great Oolite Limestone of the Midlands by Woods (1925-31). These are the palinurids
Mecochirus clypeatus (Carter) and Glyphea regleyana (Desmerest), and the astacurid
Eryma bedelta (Quenstedt). Phillips (1871) also records Glyphea rostrata Phillips from
Kirtlington.

Of these, M. clypeatus is of a suitable shape and size to have occupied our Thalassin-
oides burrows, and seems the most likely of the known crustacean fauna to have pro-
duced both burrows and faecal pellets. Some poorly calcified form, not preserved at all
could equally have been responsible.

The beds in which the trace-fossils occur consist of shelly, oolitic, and marly lime-
stones, often bioturbated, with a rich and variable fauna. Some beds, yielding brachio-
pods ( Epithyris ) and mytilid bivalves ( Modiolus ) are interpreted as subtidal channel
fills; other beds, with infaunal bivalves and annelids represent tidal flat deposits
(McKerrow et ah 1969).

The Favreina crustacean thus lived at, or close to low water mark; even if the animals
colonized tidal flat environments, their burrows probably reached waterlogged sedi-
ments.

The extension of the recorded range of Favreina decemlunulatus into the Jurassic is not
particularly remarkable. It does not automatically imply the altogether improbable
occurrence of the same crustacean during 120 million years, rather it should be com-
pared with the great time-range of other trace-fossils (Hantzschel 1962).

KENNEDY, JAKOBSON, AND JOHNSON: FA VREINA-THALASSINOIDES 553

Acknowledgements. We acknowledge most gratefully the encouragement of Dr. W. S. McKerrow, and
the assistance of Mr. G. F. Elliott and Dr. R. P. S. Jefferies towards the identification of our material.
Dr. A. C. Kendall has kindly allowed us to quote from his unpublished researches on the Great Oolite.
We thank Professor M. R. House for allowing us to publish his photographs of Favreina.

REFERENCES

altini, e. 1942. Etude geologique de la chaine cohere entre Bandirma-Gemlik. Istanb. Univ. Fen. Fak.
Mean., Ser. B, 8, 76-137, 6 pi.

bromley, r. g. 1967. Some observations on burrows of thalassinidean Crustacea in chalk hardgrounds.
Q. Jl. geol. Soc. Lond. 123, 157-82, pi. 7-11.

bronniman, p. 1955. Microfossils incertae sedis from the EJpper Jurassic and Lower Cretaceous of
Cuba. Micropaleontology , 1, 28-51, 2 pi.

and Norton, p. a. 1961. On the classification of fossil fecal pellets and description of new forms

from Cuba, Guatemala and Libya. Eclog. geol. Helv. 53, 832-42.
brown, r. w. 1939. Fossil plants from the Colegate member of the Fox Hills sandstone and adjacent
strata. U.S. Geol. Surv. Prof. Paper, Washington, 189-lc, 239-75, pi. 48-63.
cuvillier, j. and SACAL, V. 1956. Stratigraphic correlations microfacies in western Aquitaine. 2nd ed.,
1-33, pi. 1-100. Leiden.

ehrenberg, k. 1944. Erganzende Bemerkungen zu den seinerzeit aus dem Miozan von Burgschleinitz
beschreibenen Gangerkern und Bauten dekapoder Krebse. Paldont. Z. 23, 354-59.
edge, e. r. 1934. Faecal pellets of some marine invertebrates. Am. Midi. Nat. 15, 78-84.
elliott, g. f. 1962. More microproblematica from the Middle East. Micropaleontology, 8, 29-44,
pi. 1-6.

1963. Problematical microfossils from the Cretaceous and Palaeocene of the Middle East.

Palaeontology, 6, 293-300, pi. 46-7.

farrow, g. e. 1966. Bathymetric zonation of Jurassic trace-fossils from the coast of Yorkshire,
England. Palaeogeog. Palaeoclim. Palaeoecol. 2, 103-51, pi. 1-7.
hallam, a. 1961. A sedimentary and faunal study of the Blue Lias of Dorset and Glamorgan. Phil.
Trans. R. Soc. (B) 243, 1-44, pi. 1-2.

hantzschel, w. 1952. Die Lebensspur Ophiomorpha Lundgren im Miozan bei Hamburg, ihre welt-
weite Verbreitung und Synonymie. Mitt. Geol. St. Inst. Hamb. 21, 142-53, pi. 13-14.

1962. Trace-fossils and Problematica. In Treatise on invertebrate paleontology. Part W, 177-245,

figs. 109-49. New York.

joukowsky, e. and favre, j. 1913. Monographic geologique et paleontologique du Saleve (Haute-
Savoie, France). Mem. Soc. Phys. Hist, nat, Geneve, 37, 296-519.

Kennedy, w. j. 1967«. Field Meeting at Eastbourne, Sussex. Proc. Geol. Ass. 77, 365-70.

1967 b. Burrows and surface traces from the Lower Chalk of South-east England. Bull. Br. Mus.

(Nat. Hist.) (Geol.) 15, 3, 125-67, 9 pis.

mckerrow, w. s., Johnson, r. t. and jakobson, m. e. 1969. Palaeoecological studies in the Great
Oolite at Kirtlington, Oxfordshire. Palaeontology, 12, 56-83, 2 pi.
miller, f. r. 1910. On the rhythmic contractility of the anal musculature of the crayfish and lobster.

J. Physiol. 40, 431-44.

moore, h. b. 1 93 1 <7. The systematic value of a study of molluscan faeces. Proc. malac. Soc. Lond. 19,
281-90.

• 19316. The specific identification of faecal pellets. J. mar. biol. Ass. U.K. n.s. 17, 359-65.

— — 1932u. The faecal pellets of the Trochidae. Ibid. 18, 235-41.

19326. The faecal pellets of the Anomuran Crustacea. Proc. roy. Soc. Edinb. 52, 296-308.

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.

Phillips, J. 1871. The geology of Oxford.

C 6940

o o

554 PALAEONTOLOGY, VOLUME 12

wilcox, n. r. 1953. Some coprolites from phosphatic chalks in S.E. England. Ann. Mag. nat. Hist. (12)
6, 369-75, pi. 11.

woods, h. 1925-31. A monograph of the fossil Macrurous Crustacea of England. Palaeontogr. Soc.
(Monogr.).

W. J. KENNEDY

Department of Geology and Mineralogy
Parks Road
Oxford

M. E. JAKOBSON

Department of Biology
Royal Free Elospital School of Medicine
8 Hunter Street
London, W.C. 1

R. T. JOHNSON

Eleanor Roosevelt Institute for Cancer Research
University of Colorado Medical Centre
Denver

Typescript received 27 February 1969 Colorado, U.S.A.

A NEW SPECIES OF FOSSIL TURTLE FROM THE
UPPER SI WA LI KS OF PINJORE, INDIA

by b. s. tewari and G. l. badam

Abstract. A new species of freshwater fossil turtle, Geoclemys sivalensis n. sp. is described from the Pinjore
Stage (basal Pleistocene) of the Upper Siwaliks near Chandigarh, India.

Representatives of Testudines are known from the Lower as well as Upper Siwaliks
(Smith 1931). The Lower Siwaliks have yielded nine species of turtles of the genera
Chelonia, Trionyx, and Emys (Pascoe 1962). The Upper Siwaliks of the type region are
fairly rich in representatives of the Reptilia but these need further study.

This paper records the occurrence of the genus Geoclemys and describes a new species,
Geoclemys sivalensis n. sp., from about 1 km. south-east of Quranwalla (30° 46' 5";
76° 52' 50") in the type area of Pinjore. The geology of the area has recently been out-
lined by Sahni and Khan (1959).

The genus Geoclemys is a tropicopolitan freshwater land turtle and is represented in
the present-day Indogangetic system by Geoclemys hamoltoni (Gray 1831). The species
lives in well vegetated shallow, clear, freshwater of oxbow lakes and ponds. It is con-
sidered that G. sivalensis, which is a closely related form, might have inhabited a similar
environment. The Upper Siwalik sediments of the Pinjore Stage comprise alternations of
clays, gravels, and sands with common cross-stratification, indicative of a fluvial facies.

Acknowledgements. The authors are grateful to Dr. L. B. Halstead (Reading University) for his
criticism of the typescript, and to Miss Kamla Dutt (Panjab University) and Miss Lalita Kaw (Jammu
and Kashmir University) for help during the preparation of this paper.

SYSTEMATIC PALAEONTOLOGY

Order chelonia (testudinata)

Family emydidae Gray 1825
Genus geoclemys Gray 1855

Geoclemys sivalensi sp. nov.

Diagnosis. Carapace oblong and tricarinate, strongly arched transversely, lateral edges
straight, margin not flared, nuchal pentagonal, three neurals, and three pairs of pleurals.

Holotype: Carapace, A/665; Museum of Centre of Advanced Study in Geology, Panjab University,
Chandigarh, India.

Locality and horizon : 1 km. south-east of Quranwalla, which is 6 km. northeast of Chandigarh Lake.
Pinjore Stage (basal Pleistocene).

Description. The carapace is oblong and tricarinate being strongly arched breadthwise
and widest at about one-quarter the distance from the anterior end. The lateral edges of
the carapace are straight and taper slightly towards the posterior margin which is not

[Palaeontology, Vol. 12, Part 4, 1969, pp. 555-8.]

556

PALAEONTOLOGY, VOLUME 12

text-fig. 2. Geoclemys sivalensis, n. sp., dorsal view of carapace, P.U. Mus. A/665. X

TEWARI AND BADAM: PLEISTOCENE TURTLE

557

flared. Three interrupted keels or series of nodose prominences are present on the
carapace. The pentagonal nuchal abuts against the first neural. There are three rect-
angular neurals which tend to become hexagonal, the first having a convex anterior end,
with an almost straight posterior end and with the lateral edges somewhat concave
anteriorly. The second neural is the largest; a prominent median ridge with its highest
point near the posterior end runs along the length of the neural continuing from one to
the other. The pleurals appear to be in three pairs alternating with the neurals, the third

TABLE 1

Measurements in millimetres: (E) estimated, (R) right side, (L) left side

Carapace: length 230 (E), width 215, height 95 (E).

Nuchal: length 26, maximum width 20, anterior width 1 1.

Neurals:

length

width

1

71

52-5

2

67-7

48-5

3

51

45

Pleurals: (L) maximum

minimum

maximum

minimum

length

length

width

width

1 101

69

100

74

2 46

—

101

—

Pleurals: (R)

1 101

68

101

76

2 50

—

96-5

—

Peripherals: (L)

inner

length

outer

length

1

38

50

2

36

39

3

85

47

4

44 (E)

47 (E)

one is not preserved in the present specimen but its presence can be inferred from the
part of the third neural present in the specimen. The first pair of pleurals are roughly
rectangular with the antero-lateral edge arched and the surface marked with arched ribs
indicating growth stages. Near the proximal end a ridge extends from the posterior
margin towards the anterior side and gradually merges into the general level of the
pleural. The second pair of pleurals seems to be rectangular but is incomplete in the
present specimen, the ridges representing the growth-lines in this case are parallel to
the lateral margin. The ridge on the proximal side, extending from posterior margin, runs
anteriorly merging again into the general level of the pleural. It is also a continuation of
the ridge seen on the first pleural. The third pleural and suprapygal are not seen. Three
pairs of marginals are faintly discernible near the anterior margin of the carapace,
especially on the left side. The first marginal is the largest, having the shape of a roughly
elongated hexagon, and second and third are roughly rectangular. Sutures between
neurals and pleurals are prominent.

On the ventral side there are only parts of two fairly preserved mesoplastra; the

558

PALAEONTOLOGY, VOLUME 12

sutures of the mesoplastra are visible and form the longest part of the sagittal suture.
Fragmentary parts of the entoplastron and hypoplastron are seen. In fact the plastron
has been damaged to such an extent that many of the sutures have been obscured.

Remarks and comparison. The present form resembles in general appearance G. hamoltoni
(Gray), a living species of Indogangetic turtle, also described by Minton (1966) from
Sehwan (Pakistan). In G. sivalensis the shape and morphology of the pleurals and
neurals is similar to that of G. hamoltoni but differs in the number of pleurals and neurals
which are restricted to three instead of four in the latter. The dimensions of other parts of
the carapace differ. G. sivalensis resembles G. hamoltoni in the presence of three pairs of
central ridges on the lamina which also suggests that the forms represent young stages.

REFERENCES

minton, s. a. 1966. A contribution to the herpetology of West Pakistan. Bull. Amer. Mus. Nat. Hist.
134, 59-70.

pascoe, e. h. 1962. A manual of the geology of India and Burma. 3, Government of India Publication,
Calcutta.

smith, m. a. 1931. The fauna of British India including Ceylon and Burma. Reptila and Amphibia, 1,
49-116, London.

sahni, m. r. and khan, e. 1959. Stratigraphy, structure and correlation of the Upper Siwaliks east of
Chandigarh. Jl. Pal. Soc. India, 4, 61-74.

B. S. TEWARI and G. L. BADAM
Centre of Advanced Study in Geology
Panjab University

Final typescript received 24 April 1969 Chandigarh, India

LOWER DEVONIAN LAND PLANTS FROM
GRAPTOLITIC SHALE IN SOUTH-EASTERN

ALASKA

by MICHAEL CHURKIN, JR., G. DONALD EBERLEIN,
FRANCIS M. HUEBER, and SERGIUS H. MAMAY

Abstract. The discovery of vascular plants ( Drepanophycus sp. and Hostimella spp.) in graptolitic shale from
Noyes Island, south-eastern Alaska is the first record of such an association in North America and the oldest
confirmed occurrence of land plants in the western hemisphere. Monograptus aff. thomasi associated with the
plants on Noyes Island occurs in Australia with the famous Baragwanathia flora that has long been considered
Silurian, but now is regarded to be no older than early Devonian, about the same age as the earliest undoubted
vascular plants in Europe. Corals that occur in limestone interbedded with the Alaskan plant and graptolite-
bearing shale further indicate an early Devonian age.

The plant and graptolite shale of Noyes Island is part of a section composed predominantly of conglomerate,
sandstone, and coral limestone breccia, suggesting high-energy shallow marine sedimentation that was interrup-
ted by brief periods of accumulation of graptolitic mud that also preserved fragments of land plants that lived
on nearby uplifts.

For many years the earliest record of vascular land-plants was known with certainty
only from the Old Red Sandstone of early Devonian age in England. A few records from
older strata had been open to doubt either because the age of the beds was not proven
or because the plant remains were obscure. Then from Australia Lang and Cookson
(1935) described the remarkably well-preserved Baragwanathia flora from graptolite-
bearing shale in Victoria. The graptolites associated with these plants, often preserved
on the same slab of rock, were considered as definitely Silurian (and not younger than
lower Ludlow); thus the associated plants were considered the most ancient record of
vascular plants anywhere in the world.

Recently, however, re-examinations of the Australian graptolites (Jaeger 1966,
1967; Berry 1965), long regarded as conspecific with those from the British lower Lud-
low, indicate that they correlate with the considerably younger, Lower Devonian,
graptolite succession established by Jaeger (1959, 1962) in Europe. The Baragwanathia
flora accordingly is now considered post-Ludlow and approximately the same age as the
earliest undoubted vascular plants in Europe.

The presence of land plants in earliest Devonian or possibly latest Silurian grapto-
litic shale although very rare has been observed from widely separated regions: Bohemia
(Obrhel 1962); Germany (Zimmermann 1953, Roselt 1962); Russia (Obut 1957);
Australia (Lang and Cookson 1935).

The discovery of vascular plants in graptolitic shale from south-eastern Alaska
described in this paper is the first record of such an association from North America and
the oldest confirmed occurrence of land plants in this hemisphere. In addition, the close
association of corals with the plant and graptolite-bearing shale further defines the age of
the plants in terms of a marine shelly fauna.

Acknowledgements. This study is the result of finding plants in graptolite shale in 1965 by M. Churkin,
Jr., G. D. Eberlein, and A. T. Ovenshine while mapping parts of the Craig quadrangle in south-eastern

[Palaeontology, Vol. 12, Part 4, 1969, pp. 559-73, pis. 100-1.]

560

PALAEONTOLOGY, VOLUME 12

SOUTHEASTERN ALASKA

text-fig. 1 . Geologic map of north-eastern Noyes Island, south-eastern Alaska. Explanatory symbols

on opposite page.

CHURKIN ET AL.: LOWER DEVONIAN LAND PLANTS FROM ALASKA 561

Qag

Alluvial and glacial deposits

di

Diorite

UNCONFORMITY

=4.

S4.

O

<5

I

r

Karheen Formation

DSk, conglomerate and sandstone with interbedded chert
grit, limestone, and calcareous siltstone
DSks, black graptolitic shale and coral limestone
breccia

UNCONFORMITY

SOdg

Descon Formation

Graywacke and mafic submarine
volcanic rocks with inter-
bedded siliceous siltstone ,
conglomerate, and limestone

SOdl

Descon OFormat ion

Massive, fi ne-gra ined crystalline
limestone

cc -
uj Jr
h- ec
< <
3 Z
O'

P 3
uj O
or uj

o o

< _ «

g<^

cr c/7

o

,75

_^5°

Contact

Dashed where approximately
located; dotted where
concealed; queried where
doubtful

Strike and dip
of beds

Strike and dip
of cleavage

Fault, showing dip

Dashed where approximately
located; dotted where
concealed; queried where
doubtful

3~> 20

Minor anticline, showing plunge

Minor syncline, showing plunge

<r

FA

Folds axes, horizontal

Fossil locality

text-fig. 1 ( continued ). Explanation of symbols. Contours are in feet above and below sea level.

Alaska for the U.S. Geological Survey. We are indebted to Hermann Jaeger (Institut fur Palaontologie
und Museum der Humboldt Universitat, Berlin) for his identification of Monograptus aflf. thomasi ; to
A. M. Obut (Institute of Geology and Geophysics, Novosibirsk) for discussion of graptolite-plant
occurrences in the U.S.S.R.; to I. I. Chudinova, V. N. Dubatolov, A. B. Ivanovskii, K. A. Ermakova,
A. G. Kravtsov, F. E. Yanet, and other Soviet palaeontologists for their identifications and comments
on the Alaskan tabulate corals while Churkin visited the Soviet Union on the exchange programme
of the United States and Soviet Academies of Science. Claire Carter helped prepare the graptolite
descriptions. Publication authorized by the Director, U.S. Geological Survey.

562

PALAEONTOLOGY, VOLUME 12

STRATIGRAPHY

The beds containing the plant-graptolite assemblage described here crop out along the
north-east shore of Noyes Island, a small island in the Alexander Archipelago of south-
eastern Alaska (text-fig. 1). The predominantly greywacke and mafic volcanic rocks
with interbedded siltstone, shale, conglomerate, and limestone constitute one of the most
westerly exposures of the Palaeozoic Cordilleran geosyncline in North America.

The stratigraphic details in the Noyes Island area are complicated by rapid lateral
gradation from one lithology into another, especially among the volcanic rock types,
and by tight folding and high-angle faulting (text-fig. 1).

The oldest rocks in the immediate area, the Descon Formation (Eberlein and Churkin,
in press), consist mainly of greywacke and siltstone, with minor interbeds of basaltic
tuft' and limestone, bedded chert with graptolitic shale partings, and chert-rich pebble
conglomerate. The only fossils found in the formation are graptolites indicating a range
from upper Arenigian (Early Ordovician) at locality A, into upper Caradocian (Middle
Ordovician) at locality I, and finally into the lower Llandoverian zone of Akidograptus
acuminatus (earliest Silurian) at locality J (see text-fig. 1 for fossil locations, and text-
fig. 2 for stratigraphic positions of key graptolite faunules).

Overlying the Descon Formation with apparent unconformity is the Karheen For-
mation (Eberlein and Churkin, in press). Conglomerate, sandstone, siltstone, and shale
are the dominant lithologies of the Karheen also, but the formation generally is not as
indurated as the Descon, has conspicuous amounts of carbonate cement, and contains
far more limestone interbeds than the underlying Descon Formation. Interbedded with
fossiliferous limestone and calcareous siltstone is a thin black graptolitic shale unit
(text-fig. 1 ). At two localities (C and F), plants were found on the same shale bedding
planes with graptolites.

The best plant-graptolite locality is on a small point on the east shore of Steamboat
Bay (locality C3), where a wave-cut bench exposes about 75 ft. of nearly vertical black
shale that is underlain by limestone and limestone conglomerate and apparently over-
lain by the coral-limestone breccia that forms the seaward edge of the point. Widely
divergent orientations of cleavage and fractures within the shale imply that it is inter-
nally folded and faulted. In fact, what appears to be the basal part of the shale fossil
locality C2 surprisingly yields a Lower Silurian graptolite fauna far older than the plant
and graptolite-bearing upper part of the shale unit of early Devonian age. Thus, either a
similar black shale that contains much older graptolitic material has been faulted into the
base of the unit or the shale represents a condensed section about 75 ft. thick that ranges
in age from early Silurian into early Devonian.

The other plant-graptolite locality (text-figs. 1 and 3; locality F) is about 0-70 miles
east of locality C at the head of a rocky cove 0-85 miles south-east of Point Incarnation.
Plant remains are also present in shale at locality H (text-fig. 1).

GENERAL DESCRIPTION OF THE PLANTS AND FAUNAS

The Lower Devonian graptolites and plants are frequently encountered on the same
slabs of shale providing the shale splits parallel to the bedding. Where cleavage has
developed at angles to the bedding, no trace of these fossils is visible on fracture surfaces.

CHURKIN ET AL.: LOWER DEVONIAN LAND PLANTS FROM ALASKA 563

O

O

Top covered by water

■ o o

DESCRIPTION

Polymictic conglomerate, graywacke, and banded silt
stone; calcareous cement

0 \a%\:adb;o\\d?

Mostly covered in cove

Fossiliterous limestone blocks and boulders cemented by
C4 limestone with polymictic pebbles and cobbles; fossili
ferous limestone clasts. Forms small point of land

Black plant-graptolite shale; rare interbeds of calcareous
chert sandstone. Monograptus thomasi
Graptolitic cherty shale of lower Silurian age
Probable fault slice of Descon Formation
Limestone conglomerate
Orange weathering

Calcareous polymictic conglomerate and siltstone
v Poorly exposed in cove

Calcareous siltstone and sandstone; Forms most promi-
\nent point on east shore of Steamboat Bay
Polymictic pebble conglomerate
Massive limestone

Limestone conglomerate; fossiliterous limestone cobbles
in a sandy limestone cement

Argillaceous limestone and chert wacke
Argillaceous wacke

Polymictic conglomerate
Chert-rich graywacke

Very thin bedded, dark gray chert with graptolitic shale
partings

Pebbly graywacke
Covered

Massive, finely crystalline limestone,
nonfossiliferous

-50

-100 Feet

text-fig. 2. Detailed stratigraphy on eastern shore of Steamboat Bay, Noyes Island. For explanation
of symbols see text-fig. 3. For M. thomasi read M. aff. thomasi.

564

PALAEONTOLOGY, VOLUME 12

DESCRIPTION

Polymictic conglomerate and graywacke

Black graptolitic shale

Platy limestone and calcareous shale

Chert- pebble conglomerate
Polymictic conglomerate with blocks of
light -gray siltstone

Covered interval in sand spit

Polymictic conglomerate with blocks of
light-gray siltstone

Dark-gray calcareous plant-and graptolite-
bearing shale with minor interbeds of
silty to sandy limestone and calcareous
polymictic pebble conglomerate

Calcareous shale and siltstone
thinly interbedded with lime-
stone; chert pebbles and gran-
ules abundant in some layers;
minor plant-bearing black shale
(Loc. H). Abundant corals and
stromatoporoids in limestone
boulders and calcareous shale

Polymictic conglomerate
and graywacke

-0

-25

-50

— 75

L 100 Feet

EXPLANATION

Graptolites
Land plants

Stromatoporords

Corals

0

Brachiopods

text-fig. 3. Detailed stratigraphy on north-east coast of Noyes Island.

CHURKIN ET A L.\ LOWER DEVONIAN LAND PLANTS FROM ALASKA 565

Both the plants and graptolites are preserved as flattened silvery films with only a
trace of original relief present in a few specimens. Although nearly 100 individual plant
specimens were collected, most are fragmentary. Three types of plant structures are
preserved: stout axes possessing widely spaced lateral appendages; dichotomously
branched slender stems that have evenly paired branchlets; dichotomously branched
stems small, but robust.

More than 100 graptolites in association with plant fossils were collected at localities
C and F. Because of the fragmentary preservation, and in many cases intense defor-
mation of the graptolites, a first impression suggested several species of Monograptus
characterized by rapidly widening unbranched stipes having hooked thecae on one
side and a straight smooth margin on the other side. Fossils identified from these
collections from north-eastern Noyes Island, south-eastern Alaska, are listed below:

Locality C3. From a black shale section about 75 ft. thick in the Karheen Formation.
Exposed in a wave-cut bench just back from limestone breccia that forms the sea-
ward edge of a small point on the east side of Steamboat Bay. U.S. Geological Survey
field localities 65ACnll81, 66ACnll01B.

Plants: Drepanophycus sp. (rare), Hostimella sp. A (common), Hostimella sp. B
(rare).

Graptolite: Monograptus aflf. M. thomasi Jaeger (frequent).

Locality F. From about a 75-ft. -thick section of dark-grey slightly calcareous shale
containing some thin interbeds of laminated silty to sandy limestone and calcareous
polymictic pebble conglomerate, Karheen Formation. About 0-70 mile east of locality
C at the head of a rocky cove 0-85 mile south-east of Point Incarnation. U.S. Geo-
logical Survey field localities 65ACnl221 and 66ACnll21.

Plant: Hostimella sp. A (common).

Graptolite: Monograptus aflf. M. thomasi Jaeger (common).

Locality G. From 3-ft.-thick black shale containing thin beds of chert sandstone. Under-
lain by black platy limestone and calcareous shale with chert grit layers and overlain
by polymictic pebble conglomerate, all within the Karheen Formation. In small
bluff on north-east side of small cove across sandspit from locality F. U.S. Geological
Survey field locality 65ACnll92.

Graptolite: Monograptus aff. M. thomasi Jaeger (rare).

In addition to graptolites, the fossil plants in Alaska are associated with shelly faunas
consisting mainly of corals, stromatoporoids and brachiopods. (Fossil localities B, Cl5
C4, E, and H.) The best collections, mainly tabulate corals, were made from argillaceous
limestone at localities B, E, H, directly below the plant-graptolite shale, and from
a limestone boulder conglomerate and breccia, locality C4, directly above the plant-
graptolite shale (Table 1).

AGE AND CORRELATION

The graptolites associated with the plants from Noyes Island can be recognized by
their distinctively hooked thecae as belonging to the Monograptus hercynicus group.
Until recent years, monograptids were considered restricted in their age range to the
Silurian, but the most recent work on the Silurian-Devonian boundary using graptolites

566

PALAEONTOLOGY, VOLUME 12

(Jaeger 1962, Boucek et al. 1966) has shown that where monograptids of the
M. hercynicus type have been found in close association with the shelly fossils (upon
which the stages of the Silurian and Devonian are based), the shelly fossils have been
interpreted to be of post-Ludlow (post-Silurian) age. The graptolites associated with the
plants in localities C, and F compare most closely with M. thomasi Jaeger, one of the
youngest graptolites in Australia that is associated with the famous Baragwanathia
flora (Jaeger 1966, 1967).

TABLE 1

Coral faunas closely associated with plant and graptolite-bearing shale from Noyes Island, south-
eastern Alaska. Field locality numbers: Loc. B, 65ACnll72, 65ACnll62; Loc. C1; 66ACnll01D;
Loc. E, 65 ACn 1182, 65 ACn 1281; Loc. H, 65 ACn 1311; Loc. C4, 65 ACn 1181.

Species

Striatopora minuscula Tchud.
Syringopora sp.

Favosites sp.

Pachyfavo sites sp. transitional
to Favosites sp.

Thanmopora sp.

Disphyllum sp.

Tryplasma

Tryplasma altaica (Dybowski)
Pseudomicroplasma sp.
Thamnoporoid coral

Localities

B Cx E H C4

X

X X
X X

X

X

? ?

X

X X
X

X

According to Jaeger the typical form of Monograptus thomasi closely resembles M.
praehercynicus, but differs chiefly from praehercynicus in its thin, ‘stalk like’ habit of the
proximal part and in the tendency towards isolation of the first 3-5 thecae. M. thomasi
in Australia occurs with Baragwanathia in the most widespread development of plant-
graptolite beds, i.e. the Wilson’s Creek Shale and its equivalents. M. aequabilis (subsp.)
reported first to be stratigraphically below M. thomasi (Jaeger 1966) was later found to
be above M. thomasi (Jaeger 1967). M. aequabilis (subsp.) is also recognized by Jaeger
from the uppermost Pragian (Early Emsian) beds with M. atopus Boucek and M. sp. cf.

M. yukonensis Jackson and Lenz in Bohemia (Boucek 1967). In the Tanjil Formation
overlying the Wilson Creek Shale with M. thomasi occur the tentaculitids Nowakia cf.

N. acuaria and Styliolina sp. also indicating a Pragian age (Boucek 1967). This suggests
to Jaeger (1967) that the zone of M. thomasi in Australia associated with the Baragwan-
athia flora could also be early Pragian in age (Siegenian). The earliest adequately dated
occurrence of vascular land-plants in Australia and, by analogy, in south-eastern
Alaska is early Devonian, at about Siegenian time. This occurrence in Alaska consti-
tutes the earliest evidence of a land flora in the Western Hemisphere. It parallels other
occurrences, for example, in Britain (Croft and Lang 1942) and Belgium (Stockmans
1940) and Germany (Krausel and Weyland 1930) but is younger than the material
reported by Lang (1937) from Wales (Gedinnian) and by Obrhel (1962) from Czecho-
slovakia (uppermost Silurian).

The corals associated with the graptolite-plant shale are mostly long-ranging genera
of Silurian through Devonian and, in some cases, younger age. The presence of Tryplasma

CHURKIN ET AL.\ LOWER DEVONIAN LAND PLANTS FROM ALASKA 567

oltaica, a very widespread and easily recognizable large solitary coral restricted to the
Lower Devonian of Asiatic U.S.S.R., suggests an early Devonian age in agreement with
M. thomasi. In addition, the presence of Striatopora minuscula , a tabulate coral found
only in the lowest Devonian of the Kuzbas, U.S.S.R., strengthens the early Devonian
age assignment. A more detailed correlation of the Alaskan corals must await the results
of studies in progress of corals from other parts of Alaska and western North America
where the first Lower Devonian faunas have just been recognized (Churkin and Brabb
1968, Lenz 1968).

ENVIRONMENTAL SIGNIFICANCE OF THE PLANT-GR APTOLITE

ASSOCIATION

The plant-graptolite shale on Noyes Island is from the lower part of a widely distribu-
ted, mainly conglomerate, sandstone and siltstone unit (Karheen Formation) that in its
southern area of exposure on Noyes Island rests with a marked unconformity on a
greywacke sequence of varying age (Eberlein and Churkin, in press). Some of the more
richly fossiliferous limestone and limestone conglomerate of the Karheen Formation
probably represent reefs, shell bank deposits or their fragmented equivalents. The plant-
graptolite shale interbedded with these coarser-grained rocks reflects, as do numerous
thinner beds of siltstone, short periods of deposition under relatively lower energy
conditions. The presence of land plants in these, as in similar graptolitic shales, is
used as evidence of near-shore sedimentation (Miroshnikov 1956, Obut 1957). How-
ever, studies of modern ocean bottoms many miles from shore have shown land-plant
fragments of the size present in the much older graptolitic shales. For example, woody
fragments from sugar cane processing in the Hawaiian Islands literally cover the ocean
bottom at depths up to 12,000 ft. and up to 20 miles from shore (J. G. Moore — personal
communication 1968).

In Australia, the Baragwanathia flora with graptolites is in a thick succession of
siltstone, blocky mudstone, or shale that has numerous interbeds of conglomerate and
sandstone (Schleiger 1964). The coarser sediments have well-developed graded bedding
and are interpreted by Schleiger as products of turbidity currents induced by slumping of
coarse sediment into probably deep standing water accumulating the graptolite-plant
mud. The stratigraphic section at Noyes Island, Alaska, differs from the Australian
association of plants and graptolites by being mostly conglomerate and sandstone with
only occasional siltstone and shale interbeds. The Alaskan section, by its predominantly
coarse detritus including large blocks of coral head limestone (probably in part slump
blocks), limey cement, and limestone interbeds suggests rapid high-energy sedimentation
near the sediment source in shallow marine waters rich in carbonate constituents.
Therefore the source of the land-dwelling plants was probably from the nearby uplifts
within the geosyncline instead of from some distant continental area to the east.

The remarkable occurrence of the same or very similar species of Monograptus at
both ends of the Pacific in beds of about the same age supports the universally accepted
hypothesis of a drifting mode of life for graptolites, exclusive of dendroid types (Bulman
1957, 1964). It also indicates that the north and south Pacific areas were interconnected
perhaps in the form of an ancestral Pacific Ocean basin during early Devonian time.

568

PALAEONTOLOGY, VOLUME 12

DESCRIPTION OF THE GRAPTOLITES

Monograptus aff. M. thomasi Jaeger

Plate 100, figs. 1-4

1965 Monograptus sp. (of the M. hercvnicus type) forms A and B, Berry, pp. 9-13, figs. 1 a, b;
pi. 1-2.

1966 Monograptus thomasi Jaeger, pp. 403-11, text-figs. 1 a-c, o, pi. 41, figs. 3-5, pi. 42, figs.
2-7, pi. 43.

Material. About 70 compressed specimens, at least half of which are fragmentary. All are preserved as
silvery or whitish films on black slate, and show various degrees of deformation (stretching).

Description. According to Jaeger (1966, p. 404), the rhabdosome of M. thomasi is
‘medium-sized, straight, with stretched proximal portion and slightly reclined proximal
extremity. Thecae of uncinatus type, with hoods decreasing in size from proximal to
distal end; first few thecae with tendency towards isolation. Sicula with relatively long
(0-3 mm.), strongly incurved dorsal tongue. Dimensions for flattened, but tectonically
undeformed adult rhabdosomes: Length: 20-50 mm., commonly 20-30 mm. Width: 1
mm. at theca 1 across its hood, \ mm. just above the hood of theca 1 ; maximum width
attained at theca 10i2: T8-2-0 mm. (1-3—1 *6 mm. without hoods); number of thecae in
1 cm.; 9-10, practically constant throughout the rhabdosome’.

Discussion. Although the Alaskan specimens have been tectonically deformed, they
closely resemble the Australian species M. thomasi. Some of the less-deformed
specimens (PI. 100, figs. 2-4) exhibit the distinct stalk-like thinning in the proximal part
of the rhabdosome (0-6-0-7 mm. wide at theca 1) which is a characteristic feature of the
species. The initial thecae show the characteristic tendency towards isolation, most
pronounced in theca 1 .

The features which distinguish M. thomasi from species such as M. praehercynicus and
M. hercynicus which it otherwise resembles are (1) the thin, stalk-like habit of the proxi-
mal portion of the rhabdosome, (2) the tendency towards isolation of the first 3-5
thecae, and especially the strong projection of theca 1, (3) the constant number of thecae
in 1 cm. throughout the rhabdosome, and (4) the relatively long, ventrally bent dorsal

EXPLANATION OF PLATE 100

Graptolites and closely associated corals, Noyes Island, south-eastern Alaska.

Figs. 1-3. Thamnopora sp., X 2. 1, Transverse view, loc. B; USNM 162070. 2, Longitudinal view,
loc. B; USNM 162071. 3, Longitudinal and transverse views, loc. E; USNM 162072.

Fig. 4. Pseudomicroplasma sp., transverse and longitudinal views, X 2, loc. E; USNM 162073.

Fig. 5. Disphyllum sp., transverse view, x2, loc. C4; USNM 162074.

Fig. 6. Tryplasma altaica (Dybowski), transverse view, x2, loc. E; USNM 162075.

Figs. 7-11. Monograptus aff. M. thomasi Jaeger; Noyes Island, south-eastern Alaska. 7, 8, and 11,
Wide rhabdosomes with proximal stalk-like portion largely obliterated by tectonic compression along
length of rhabdosome, loc. C3; these deformed specimens cannot effectively be separated from M.
praehercynicus Jaeger. 7 and 8, x 3; USNM 162382. 11, X 5; USNM 162386. 9, 10, Specimens that
show stalk-like proximal part not appreciably tectonically modified, x3; lineation formed by
intersection of cleavage and bedding subparallel to length of rhabdosomes. 9, Loc. C3; USNM
162387. 10, Loc. F; USNM 162380.

Fig. 12. Type specimen of Monograptus thomasi Jaeger from 19 Mile Quarry, Yarra track, Australia;
x 3.

Palaeontology, Vol. 12

PLATE 100

CHURKIN et al., Lower Devonian corals and graptolites

CHURKIN ET AL .: LOWER DEVONIAN LAND PLANTS FROM ALASKA 569

tongue of the sicula (Jaeger 1966, p. 407). In the more deformed Alaskan specimens,
where the lineation or direction of stretching is nearly at right angles to the length of the
rhabdosome, i.e. PI. 100, fig. 1, the thin stalk-like shape of the proximal part is deformed
into a much broader form resembling M. praehercynicus. In these cases the thecal
count/unit length also increases (14 thecae/cm. in USNM 162382) as in the Australian
deformed specimens. Some of these specimens with wide proximal ends in Australia are
considered by Jaeger (1967) as new subspecies of M. thomasi and are found in beds below
typical M. thomasi (see also Addendum on p. 573).

DESCRIPTION OF THE PLANTS

Plant material is contained in small collections of black shale from fossil localities
C3, F, and G. Fragmentary axes are fairly abundant in these collections, but the collec-
tion from locality C3 contains the richest concentration of plant material, as well as
most of the larger specimens. The plants are preserved as thin, lustrous black films on the
otherwise dull black matrix, so that angles of oblique lighting are critical in observation
or photography. With one or two exceptions, the plant fragments show no relief what-
soever. No cuticular material or evidence of fructifications is preserved, and only one
specimen ( Drepanophycus sp.) shows evidence of emergences. Only one tiny fragment
of a xylem strand showing tracheidal wall ornamentation was found (locality C3), but
this is sufficient to establish without doubt that the material represents an assemblage of
vascular land plants.

With the exception of the single specimen of Drepanophycus, the plants are small,
naked axial fragments ranging from less than 1-0 mm. to 7-0 mm. in width and up to 5
cm. in length. Morphologically, most of the specimens are small parallel-sided fragments.
However, a few of the specimens show varying degrees of branching or other characters
that permit provisional recognition of the following entities in the assemblage.

Genus drepanophycus Goeppert 1852
Drepanophycus sp.

Plate 101, fig. 8

Material. One broken axial fragment forms the basis for the identification; both counterparts of the
upper half of the specimen were found.

Description. The specimen is a stout, unbranched axial fragment, measuring 6-5 cm. in
length and 1 cm. in width. A faint longitudinal ridge presents the suggestion of a com-
pressed, median vascular system, best seen in the lower half of the specimen. Also present
on the surface of the axis are four approximately circular, shallow depressions or
punctations that suggest points of attachment of spinose emergences. The best defined
of these is seen at about the middle of the upper half of the specimen as shown in Plate
101, fig. 8; it measures approximately 2-5 mm. in diameter.

The most definitive feature of this specimen lies in the presence of parts of several
stout leaf-like emergences. The two best examples of these are shown in the upper half
of Plate 101, fig. 8; the emergence on the right side of the specimen is about 8 mm. long
and 5 mm. broad at its base; its decurrent attitude is clearly shown in the photo-
graph. A somewhat smaller emergence is shown on the left side and somewhat lower
on the specimen. No evidence of vascularization is observable in the emergences.

pp

C 6940

570

PALAEONTOLOGY, VOLUME 12

Discussion. Although nothing is known of the branching pattern or reproductive parts
of the plant represented by this specimen, the very stout proportions of the lateral pro-
jections are distinctly more suggestive of the vascularized leaves of Drepanophycus than
of the slender spines of Psilophyton. The few characteristics of the leaves or emergences
that are preserved relate the specimen more closely to D. spinaeformis Goeppert than
to D. spinosus (Kreji) Krausel and Weyland, D. gaspianus (Dawson) Krausel and Wey-
land, or D. colophyl/us Grierson and Banks.

Genus hostimella Barrande 1882

Discussion. The generic designation Hostimella is applied to naked, dichotomously
divided, sterile, psilophyte-like plant axes, several excellent examples of which are
present in the Noyes Island collections; they are represented from both locality C and
locality F. Because of differences in degree of tapering in successive ramifications of the
axes, it is possible that two distinct taxa may be present.

Hostimella sp. A
Plate 101, figs. 1, 2, 3, 7, 9, 10, 1 1

Description. This form of Hostimella is characterized by repeated, equal dichotomies of
the slender axis, the successive divisions being not substantially more slender than their
parent axial segments. Two excellent examples are illustrated in Plate 101, figs. 1 and 10.

The largest specimen (PI. 101, fig. 10) is 6-5 cm. long and no more than 2-0 mm. wide.
Two successive dichotomies are indicated, although the left terminal segment of the
right initial segment apparently was broken away when the matrix was split, leaving
only 3 terminal segments. The angles formed by the dichotomies are about 30° in both
instances. Another similar, though less complete, specimen with one dichotomy is
shown in Plate 101, fig. 9; a yet smaller, Y-shaped fragment with a somewhat broader
angle of dichotomy is shown in fig. 3.

The most complete specimen on hand is shown in Plate 101, fig. 1. Parts of four
successive dichotomies are shown, which, if complete, would have resulted in 16 terminal
ramifications or branchlets. The dichotomies involve much broader angles than the
specimens shown in figs. 9 and 10; here they reach angles of 90° or more and display an

EXPLANATION OF PLATE 101

Plants from graptolitic shale, Noyes Island, south-eastern Alaska.

Figs. 1-3, 7, 9-11. Hostimella sp. A. 1, Most complete specimen showing four successive dichotomous
divisions, x2, loc. F; USNM 43045. 2-3, Small fragments showing different angles of divergence
of the dichotomies, X 2, loc. C3; USNM 43046 and 43047. 7, Smallest branched specimen showing

ultimate division of the branching system, x2, loc. C3; USNM 43048. 9-10, Long segments of
once and twice dichotomously branched axes; fig. 10 illustrates the largest specimen of the species,
x2, loc. F; USNM 43049 and 43050. 11, Scalariform and helical tracheids preserved by pyrite

in axis (comparable in diameter to the basal portion of axis in fig. 10), x 270, loc. C3; USNM
43051.

Figs. 4-6, Hostimella sp. B; axes stout and ultimate divisions seemingly foreshortened as compared
with H. sp. A (in figs. 1-2 especially), x2, loc. C3; USNM 43052, 43053, and 43054.

Fig. 8. Drepanophycus sp.; arrows in the illustration indicate base of emergence in face view and
emergence in side view, X 2, loc. C3; USNM 43044.

Palaeontology, Vol. 12

PLATE 101

CHURKIN et al., Lower Devonian land plants

CHURKIN ET AL.: LOWER DEVONIAN LAND PLANTS FROM ALASKA 571

essentially uniform, symmetrical pattern of division. Smaller specimens with similarly
wide angles of dichotomy are shown in figs. 2 and 7. Fig. 7 represents the smallest
branching specimen available; its base is only 1-0 mm. broad, and one of its ultimate
divisions, seen at the upper left, is a small process only 1-5 mm. long.

It should be noted that regardless of the angles of dichotomy involved, in all the
specimens included under this category of Hostimella, there is a generally slight, gradual
diminution in the width of the axis above each point of dichotomy.

A few tracheids have been found preserved as pseudomcrphs by iron pyrite in an
unbranched fragment of an axis comparable in size to the basal portion of H. sp. A
illustrated in Plate 101, fig. 10. The best-preserved tracheid is illustrated in fig. 11. It has
simple scalariform secondary thickenings. In the lower left of fig. 1 1 is a partially pre-
served tracheid with helical secondary thickenings. No information could be gained
with regard to the actual outline of the xylem strand nor of the sequence of its maturation
or development.

Hostimella sp. B

Plate 101 , figs. 4, 5, 6

Description. Three specimens of this form of Hostimella are present, all from locality C.
These each represent one more or less complete set of branch tips involving the three
ultimate orders of ramification, or the branchlets produced by the last two successive
dichotomies. The best specimen (PI. 101, fig. 5) is 13 mm. long and clearly shows two of
the ultimate axial divisions at the left; these are each slightly less than 1-5 mm. long,
while the penultimate division from which they were produced is about 3-5 mm. long.
The outstanding morphological feature common to these three specimens, and by
which they seem to be readily distinguishable from material of H. sp. A, is their stubby
appearance. This is a function of a relatively abrupt decrease in width of axial divisions
of successive orders, in contrast to a much more gradual size decrease in H. sp. A. As
shown in Plate 101, figs. 4-6, the basal widths of the specimens of H. sp. B are 2-5 to
3-0 mm.; this is substantially more robust than corresponding parts of the specimens of
H. sp. A. Furthermore, the penultimate divisions of H. sp. B are much more slender in
proportion to their parent axial segments than in H. sp. A. These differences are well
brought out by a comparison of fig. 6(B) with fig. 7 (A); in fig. 6 the penultimate division
is not more than one quarter the width of the parent division, whereas in fig. 7, this pro-
portion is more of the order of 1:2. In H. sp. B the penultimate divisions are also
relatively much shorter than in H. sp. A. As the aggregate effect of these seemingly dis-
proportionate dimensions of successive parts, the over-all appearance of Hostimella
sp. B is considerably more robust than that of H. sp. A.

SIGNIFICANCE OF THE ALASKAN PLANTS IN THE EVOLUTION OF

LAND PLANTS

It is recognized that the phenotypic differences between the two species of Hostimella
described in the foregoing paragraphs may eventually prove not to be of sufficient
magnitude to segregate the two on even a tentative taxonomic basis. It may, however, be
of more than coincidental significance that Hostimella sp. B is restricted to locality C,
while H. sp. A appears at both localities C and F. Although the localities are less than

572

PALAEONTOLOGY, VOLUME 12

a mile apart geographically, it is possible that two slightly different plant communities
are represented at the two localities and there may be some genetic basis for the morpho-
logical differences described here. These same differences are observed in plants being
studied from the Emsian of Gaspe Bay, Canada, and results so far show that truly
different plants are represented by such morphological characteristics.

A comparison of these few plants with those from other Lower Devonian floras
could have merit only in pointing up the broad morphological simplicity of earliest
Devonian land parts. The Alaskan plants are definitely the earliest record of land plants
for the Western Hemisphere, and yet we probably are not in possession of the earliest
land plants as such. The Devonian Period saw the rapid diversification of plants that
had become adapted to the rigours of growth on land, although those adaptations
probably occurred during Silurian time.

REFERENCES

berry, w. b. n. 1965. Description and age significance of M. hercynicus type Monograptids from
Eildon, Victoria. Proc. R. Soc. Victoria, 78, pt. 1, 1-14.
boucek, b. 1967. Significance of Dacryoconarid tentaculites and graptolites for the stratigraphy and
palaeogeography of the Devonian system. International Symposium on the Devonian System, Calgary,
2, 1275-81.

horny, R., and chlupac, I. 1966. Silurian versus Devonian — the present state of the Siluro-

Devonian boundary problematics and proposal of its solution. Acta Mas. not. Prag. 22B, 2, 49-66.
bulman, o. m. b. 1957. Graptolites, in Treatise on marine ecology and paleoecology. Mem. geol. Soc.
Amer. 67, 2, 987-91.

1964. Lower Palaeozoic plankton. Q. J. Geol. Soc. Lond. 120, 455-76.

churkin, michael, JR., and brabb, e. e. 1968. Devonian rocks of the Yukon-Porcupine Rivers area
and their tectonic relation to other Devonian sequences in Alaska. International Symposium on the
Devonian System, Calgary, 2, 227-58.

croft, w. n. and lang, w. h. 1942. The Lower Devonian flora of the Senni Beds of Monmouth-
shire and Breconshire. Phil. Trans. R. Soc. Lond. B231, 131-63.
eberlein, g. d., and churkin, michael, jr. (in press) Paleozoic stratigraphy in the northwest coastal
area of Prince of Wales Island, southeastern Alaska. Bull. U.S. geol. Surv. 1284.
jaeger, h. 1959. Graptolithen and Stratigraphie des jungsten Thiiringer Silurs. Abh. Deutsch. Akad.
Wiss. Berlin, Kl. Chem., Geol., Biol. 2, 197 pp.

1962. Das Silur (Gotlandium) in Thuringen und am Ostrand des Rheinischen Schiefergebirges

(Kellerwald, Marburg, Giessen). Symposiums — Band 2. hit. Arbeitstagung Silur/ De von-Grenze und
die Stratigraphie von Silur und Devon, Bonn-Bruxelles (1960), 108-35.

1966. Two late Monograptus species from Victoria, Australia, and their significance for dating the

Baragwanathia flora. Proc. R. Soc. Victoria, 79, pt. 2, 393-413.

1967. Preliminary stratigraphical results from graptolite studies in the Upper Silurian and Lower

Devonian of southeastern Australia. J. geol. Soc. Aust. 14, pt. 2, 281-86.
krausel, r. and weyland, h. 1930. Die Flora des deutschen Unterdevons. Abh. Preuss. Geol. Landesan-
stalt, n.f. 131, pp. 1-92, pi. 1-15.

lang, w. h. 1937. On the Plant-remains from the Downtonian of England and Wales. Phil. Trans.
R. Soc. Lond. 227B, 245-91.

and cookson, i. c. 1935. On a flora, including vascular land plants, associated with Mono-
graptus, in rocks of Silurian age, from Victoria, Australia. Ibid. 224B, 421-49.
lenz, a. c. 1968. Upper Silurian and Lower Devonian biostratigraphy. Royal Creek, Yukon Territory,
Canada. International Symposium on the Devonian System, Calgary, 2, 587-99.
miroshnikov, l. d. 1956. K voprosu o proiskhozhdenii graptolitovykh fatziy. Izv. Akad. Nauk SSSR,
ser. geol. 7, 25-32.

CHURKIN ET AL.: LOWER DEVONIAN LAND PLANTS FROM ALASKA 573

obrhel, j. 1962. Die Flora der Pridoli-Schichten (Budnany-Stufe) des mittelbomischen Silurs. Geologie,
Berlin, 11, 83-97, pi. 1-2.

1962. Die Silur- und Devon-Flora Bohmens. Symposiums — Band 2. Int. Arbeitstagung Silur/

Devon-Grenze und die Stratigraphie von Silur and Devon, Bonn-Bruxelles (1960), 180-5.
obut, A. m. 1957. Graptolitovye slantzy Silura i svyazannye s nimi nefteproyavleniya v Sredney Azii.
Geologiya i geokhimiya , 1, 7, 228-35.

roselt, g. 1962. Uber die altesten Landpflanzen und eine mogliche Landpflanze aus dem Ludlow
Sachsens. Geologie, Berlin, 11, 320-33.

schleiger, n. w. 1964. Primary scalar bedding features of the Siluro-Devonian sediments of the Sey-
mour district, Victoria. J. geol. Soc. Aust. 11, 1-31.
stockmans, f. 1940. Vegetaux Eodevoniens de la Belgique. Mem. Mus. R. Hist. Nat. Belg., 93 1-89.
zimmermann, w. 1953. Die Urlandpflanzen ( Psilophyten ) und ihre Bedeutung fur die Stammes-
geschichte. Aus der Heimat, 61 (7/8), 175-187.

MICHAEL CHURKIN, JR.

G. DONALD EBERLEIN

U.S. Geological Survey
Menlo Park
California 94025

FRANCIS M. HUEBER

U.S. National Museum

Washington, D.C. 20560

SERGIUS H. MAMAY

U.S. Geological Survey

Typescript received 1 November 1968 Washington, D.C. 20242

ADDENDUM

Further study of these and other monograptids from Alaska has shown that the Noyes Island
graptolites, although very similar to M. thomasi, are a distinct new species (H. Jaeger 1969, pers.
commun.).

MIOSPORES FROM THE PURBECK BEDS AND
MARINE UPPER JURASSIC OF SOUTHERN

ENGLAND

by GEOFFREY NORRIS

Abstract. Miospores from the marine Upper Jurassic and from the non-marine Purbeck Beds (Tithonian-
Berriasian) of southern England are described and illustrated. Three principal palynologic suites embracing a
total of 77 species (13 of which are new) are recognized, each delimiting broadly similar spore-pollen assem-
blages. Suite A characterizes the Upper Kimmeridgian (rotunda and pallasioides zones), the Portlandian, and
the basal Lower Purbeck Beds. Suite B characterizes the remaining Lower Purbeck and the lower part of the
Middle Purbeck. Suite C occupies the remaining Middle Purbeck and all of the Upper Purbeck. The Purbeck
Beds of central Sussex correlate palynologically with some of the Middle and the Upper Purbeck of Dorset and
with the Serpulite and Wealden 1 to 4 of north-central Germany and Holland. The lower limit of palynologic
Suite C approximately coincides with a possible Jurassic-Cretaceous boundary. The successive palynologic
suites represent a progressive diversification of the spore-pollen flora derived from principally gymnosperm-
pteridophyte vegetation. Although filicalean spores are the most diverse elements in the assemblages as a whole,
coniferalean pollen species are the most persistent as well as the numerically dominant elements. Bryophytic and
cycadalean elements remain unimportant throughout.

This paper describes miospore assemblages from the Purbeck Beds of southern Eng-
land and from the Upper Kimmeridgian and Portlandian marine sediments exposed on
the Dorset coast. Couper (1958) made an extensive study of British Jurassic and Lower
Cretaceous miospores but examined only a few samples from the Purbeck Beds of
Durlston Bay, Dorset and Mountfield, Sussex and three palynologic assemblages from
the Kimmeridgian of Scotland. He demonstrated that significant differences exist between
miospore assemblages from various Jurassic and Cretaceous stages and formations but
did not attempt detailed subdivision and correlation of the Upper Jurassic. Lantz
(19586), however, attempted a palynologic subdivision of the English Upper Jurassic in
a limited study. Hughes (1958) demonstrated a relatively detailed miospore zonation of
both marine and non-marine Lower Cretaceous sediments and correlated the Wealden
with the marine stages by this means. Recently, Hughes and Moody-Stuart (1969) have
noted a new method to correlate palynologically the Lower Cretaceous of southern
England.

The present study examines the stratigraphic value of miospores for zonation and
correlation of strata developed close to the Jurassic-Cretaceous boundary in southern
England and for age determination of the Purbeck Beds and equivalent strata in Dorset
and Sussex.

STRATIGRAPHY AND LOCATION OF SAMPLES

The stratigraphy of the sections is summarized in text-fig. 1. Location of the sections
examined are indicated in text-fig. 2. Precise stratigraphic positions of the samples
within these sections are given in the Appendix to Norris (1963). Reference should be
made to Arkell (1947) for a detailed account of the stratigraphy and structural setting
where also a full list of references to major works on the geology of the Dorset area may
be found.

[Palaeontology, Vol. 12, Part 4, 1969, pp. 574-620, pis. 102-113.]

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

575

SAMPLED

INTERVAL

DORSET

SAMPLED

INTERVAL

SUSSEX

cn

<

CL

<

LU

VARIEGATED
MARLS AND
SANDSTONES

ASHDOWN SAND

FAIRLIGHT CLAY

UPPER PURBECK

W//////A

MW

1IDDLE PURBECK

<_>

LlI

DO pj

nr UJ
Q_

W

LOWER PURBECK

UPPER" PURBECK

MIDDLE PURBECK

LOWER PURBECK

FREESTONE SERIES

<

Q

0C

O

Q.

WF*

CHERTY SERIES

wm.

<

Q

on

O

CL

PORTLAND

SANDSTONE

PORTLAND SAND

r

e>

o

KIMMERIDGE

CLAY

(UPPER PART)

text-fig. 1. Generalized succession of strata close to the Jurassic-Cretaceous boundary in Dorset and
Sussex. Correlations are not implied between the two areas.

Swindon

576

PALAEONTOLOGY, VOLUME 12

text-fig. 2. Locality map of Kimmeridgian, Portlandian, and Purbeck sections examined in southern England, plus other areas
mentioned in the text. Inset map shows localities on the Dorset coast within the area enclosed by a broken line on the main map.

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

577

Samples were selected on the basis of lithologies most suitable for palynomorph
preservation in each section. In general, the samples from the Kimmeridge Clay are
shales or silty shales and those from the Portland Sand are siltstones or fine-grained
sandstones. Clastic material is almost entirely lacking in the Portland Stone and all
samples from here are relatively pure limestones. Samples from the Purbeck Beds are
more varied but in general grey and brown shales, silty shales, and clays yielded the best
assemblages. Of the carbonate rocks in the Purbeck Beds, argillaceous limestones are
the most satisfactory for palynologic examination.

TABLE 1

Zonal classification of the Portlandian and upper part of the Kimmeridgian in Dorset, with thicknesses
of respective rock divisions exposed at West Weare Cliff (Isle of Portland) and Hounstout (Isle of
Purbeck). The strata comprising the upper two Portlandian zones constitute the Portland Stone and
those of the lower two zones the Portland Sand. Numbers of samples examined are indicated.

Stage Zone

Portlandian

Titanites titan

Kerberites

okusensis

Glaucolithites

gorei

Zaraiskites

albani

Kimmeridgian

Pavlovia

pallasioides

Pavlovia

rotunda

Marine Upper Jurassic

Isle of Portland

Rock Units

Isle of Purbeck

Barren

Freestone Series 25 ft.

Cherty Series 60 ft.
Basal Shell Bed 6 ft.

Not

sampled

Freestone Series
50 ft.

Cherty Series 65 ft.

' Portland Clay 14 ft.

West Weare Sandstone
40 ft.

22 samples <

Exogyra Bed 8 ft.

Upper Black Nore Beds

35 ft.

Black Nore Sandstone
6 ft.

Lower Black Nore Beds
25 ft.

9

l

samples <

Black Sandstones and
Parallel Bands 44 ft.
St. Alban’s Head
Marls 45 ft.

White Cementstone
2 ft.

Emmit Hill Marls
30 ft.

, Massive Bed 6 ft.

12 samples Kimmeridge Clay
100+ ft.

Not (Not exposed at West

sampled Weare Cliff)

' Hounstout Marl
50 ft.

Hounstout Clay 30 ft.
Rhynchonella Marls

10 samplesi

20 ft.

Lingula Shales 40 ft.
Rotunda Clays and
Nodules 98 ft.
Crushed Ammonoid
Shales 108 ft.

(a) Hounstout ( Isle of Purbeck). A complete succession from the Upper Kimmeridgian
to Portlandian, as shown in Table 1, is exposed in the face of Hounstout and surround-
ing cliff's at Chapman’s Pool, Emmit Hill, Pier Bottom, and St. Alban’s Head on the
Isle of Purbeck (Arkell 1935, 1947). Number of samples collected is shown in Table 1.

Arkell’s (1935, 1947) zonal classification of the Kimmeridgian and Portlandian is
followed, with modifications suggested by House (1958 a , b), and is also shown in
Table 1. The top two ammonite zones of the Kimmeridge clay were examined in this
study so that the typical shaley facies of the Kimmeridgian would be included.

578

PALAEONTOLOGY, VOLUME 12

( b ) West Weave Cliff (Isle of Portland). A continuous succession from Kimmeridge
Clay to Portland Stone is exposed in the cliff's on the west side of the Isle of Portland as
shown in Table 1 (Arkell 1933, 1935; House 1958u). The number of samples used is also
shown in the Table.

Pur beck Beds

The lower boundary of the Purbeck Beds in southern England is delimited by Port-
landian sediments which belong to different zones in different areas (Arkell 1953, Taitt
and Kent 1958, Falcon and Kent 1960). The Purbeck Beds are terminated by the in-
coming of the Wealden facies (Allen 1955). Arkell (1947, 1956) used ‘Purbeckian’ as
a stage of the Jurassic system, basing it on three ostracod zones recognizable in the
Purbeck Beds. However, the Purbeckian of north-west Europe is now considered as a
predominantly freshwater and continental facies developed at the top of the Jurassic
and the base of the Cretaceous (Maubeuge 1962, Oertli 1963), and to avoid confusion
the term will not be used.

It has been customary in British stratigraphy for more than a century to use the term
‘Purbeck Beds" for all those continental, freshwater, brackish, estuarine, lagoonal or
partly marine, usually fine-grained and calcareous sediments developed above the Port-
land Beds in southern England (Fisher 1856, Bristow 1857, Topley 1875, Woodward
1895, Arkell 1933, Howitt 1964). The desirability of using the same rock unit name for
similar sediments from widely separated localities is questionable. The term Purbeck
Beds is used in this study for all areas because it is still in widespread use. The erection of
other formational names requires more intensive work on the detailed stratigraphy at
each locality than was attempted here. The present work on the Purbeck palynologic
assemblages may help to resolve some problems of the stratigraphy and correlation of
the Purbeck Beds.

In a later section, reference will be made to the ‘Purbeckian’ of France and Switzer-
land developed in and around the Jura Mountains (Arkell 1956, Donze 1958). To avoid
confusion and to remove any connotation of time-concordance, these sediments will be
informally termed, for purposes of discussions, the ‘Swiss Purbecks’, ‘French Purbecks’,
or ‘Jura Purbecks’, depending on the context.

Some sections of the Purbeck Beds in southern England have been zoned using
ostracods. The history of the zonal classification of the Purbeck Beds has been given by
Anderson (1958, 1962), the later reference with particular regard to the Upper Purbeck.
Anderson (1940) proposed a more detailed subdivision of the Purbeck Beds than the
original tripartite division of earlier workers. This was subsequently revised by Sylvester-
Bradley (1949) and Anderson (1958). In consequence, the limits of the Lower, Middle,
and Upper divisions have undergone considerable fluctuations. Anderson (1958)
proposed 6 ostracod zones of which the lower 2 define the Lower Purbeck, the succeed-
ing 3 the Middle Purbeck, and the highest zone characterizes the Upper Purbeck
(Table 2). The limits of these three divisions now correspond exactly to those used by
Bristow (1857).

(a) Dorset Coast. The type section of the Purbeck Beds is exposed on the north limb
of the Purbeck anticline in Durlston Bay. There is a general attenuation of the Purbeck
Beds westwards from Durlston Bay (390 ft.), Worbarrow Bay (290 ft.). Bacon Hole
(250 ft.) to Lulworth Cove (179 ft.) (see Howitt 1964, fig. 7). Purbeck assemblages have

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

579

not been recovered westwards beyond Lulworth Cove. The detailed stratigraphy of the
Purbeck Beds has been described by Fisher (1856), Bristow (1857), and Damon (1884).
Bristow’s sections have usually been taken as a basis for later work, but his rock units
have been emended by Woodward (1895), Strahan (1898), and Arkell (1933). Some
workers still use Bristow’s rock unit nomenclature in preference to Arkell’s. The two
schemes are compared in Table 2, where also the ostracod zones of Anderson (1958)
are indicated.

TABLE 2

Ostracod zonal classification of the Purbeck Beds from Anderson (1958), with Arkell's and Bristow’s
alternative stratigraphic nomenclature of the Dorset Purbeck sections; the Mountfield Purbeck is
from Howitt (1964) including his suggested lithologic correlations with Dorset.

Zones

Upper Purbeck
Cypridea
setina

Middle Purbeck
Cypridea
propunctata

Cypridea

fasciculata

Cypridea

granulosa

Lower Purbeck
Fabenella
boloniensis

‘ Cypris ’
purbeckensis

Dorset

Arkel! 1953

Bristow 1857

Upper Cypris Clay and
Shales

Unio Beds
Upper Broken Shell
Limestone

Chief Beef Beds

Corbula Beds
Scallop Bed
Intermarine Beds
Cinder Beds
Cherty Freshwater
Beds

Marley Freshwater Beds

Soft Cockle Beds

Hard Cockle Beds
Cypris Freestone
Broken Beds
Soft Cap
Hard Cap

Viviparus Clays

Marble Beds and
Ostracod Shales
Unio Beds
Broken Shell
Limestone

Chief Beef Beds

Corbula Beds

Upper Building Stone
Cinder Bed
Lower Building Stone

Mammal Bed

Broken Beds
Caps and Dirt Beds

Mountfield, Sussex

Greys Limestone
Series

Shales with Beef and
Clay Ironstone

Arenaceous Beds
Cinder Beds
Plant and Bone Beds

Rounden Greys
Main Gypsiferous Beds

Marls with Gypsum and Blue Limestone Series
Insect Beds

Four Purbeck localities were examined along the Dorset Coast. The type section at
Durlston Bay was examined in most detail. One hundred and twenty samples were
collected from the 390-ft. section of Lower, Middle, and Upper Purbeck at this locality.
Of the 85 samples treated for palynomorphs, 69 yielded assemblages.

At Worbarrow Bay, 18 samples were collected, 15 were processed, and 12 yielded
palynomorphs in the 290-ft. section.

At Bacon Hole, 73 samples were collected from the 250-ft. Purbeck section. 47
samples were processed and 37 of these yielded palynomorphs.

At Lulworth Cove, 30 samples were collected from the 179-ft. section. Of these, 17
samples were processed for palynomorphs but only 5 yielded assemblages. The poor
yield at Lulworth may be related to the increased proportion of carbonates at this

580

PALAEONTOLOGY, VOLUME 12

locality compared with areas further east. Limestone proved to be generally barren of
palynomorphs. For the same reason, sections of the Purbeck Beds developed close to
the edge of the basin at Teffont in the Vale of Wardour (Andrews and Jukes-Browne
1894), at Swindon, Wiltshire (Sylvester- Bradley 1941), and at Hartwell, Buckingham-
shire (Ballance 1963) all proved to be barren of palynomorphs and are not considered
further here.

( b ) Mountfield, Sussex. Purbeck Beds are brought to the surface in Sussex along the
central line of the Wealden dome in the Brightling anticline (Edmunds 1954). The lower
part of the Purbeck Beds is known in subsurface sections in the mine of Gyproc Ltd., at
Mountfield and in various boreholes. The stratigraphy, compiled from surface and sub-
surface sections, has been summarized by Topley (1875), White (1928), and Allen
(19606). A more detailed account has been given by Howitt (1964). The general succes-
sion of the Purbeck Beds in the vicinity of the mine is shown in Table 2.

At Mountfield, 31 samples were collected between the lowest gypsum seam, at the
base, and the Greys Limestone Series at the top of the section, a total of almost 400 ft.
(Howitt 1964). These samples were from surface outcrops and from subsurface sections
in the mine and from Gypsum Mines borehole M 64 (grid reference TQ 716187).
Further details of this borehole are given in Appendix 1 of Norris (1963). Of the 23
samples processed from Mountfield, 21 yielded palynomorphs.

(c) Warlingham Borehole , Surrey. Preliminary accounts of the succession in this bore-
hole have been published in the Summary of Progress of the Geological Survey (1957,
p. 29; 1958, p. 48). The Purbeck-Wealden junction has been placed at 1,892 ft. on the
basis of ostracod faunas and the top of the Middle Purbeck at 1,935 ft. (Anderson
1962; Howitt 1964, p. 105). The top of the Portland Beds is recognized at 2,150 ft. but
below 2,040 ft. there appears to be a repetition of lithological and faunal types in the
Purbeck Beds, which is attributed to reversed faulting.

Twenty-three core samples between 1,900 and 2,027 ft. in the borehole were examined
for palynomorphs. Of these, 18 yielded assemblages.

Fairlight Clay

On the coast east of Hastings, the Fairlight anticline brings the Fairlight Clay to the
surface (Edmunds 1954). The Fairlight Clay is the lowest Wealden formation and is
generally thought to pass laterally north and westwards into, and to be overlain by,
the Ashdown Sand (Allen 1960n). The complete Fairlight Clay succession has been
described by Topley (1875) and by White (1928), who believed that the Purbeck Beds
underlie this section, close to the surface on the crest of the anticline. Allen (1955)
stated that the Upper Purbeck-Wealden junction as defined by ostracods is probably
situated low down in the Fairlight Clay near the base of the Wealden as traditionally
defined, i.e. the base of the Fairlight Clay. A series of calcareous clays and limestones
developed between 949 and 1,080 ft. in the Henfield borehole and regarded as equivalent
to the basal portion of the Fairlight Clay (on ostracod faunas) \ . . could equally well be
classed with the Purbeck . . .’ (Taitt and Kent 1958, p. 12). The lateral change of facies
at this horizon promotes the possibility that the lower part of the Fairlight Clay on the
Sussex coast may be a lateral equivalent of part of the Purbeck Beds of Dorset. Allen
(19606, p. 7) noted ‘The problem of how far the Purbeckian extends upwards (if at all)
into the Fairlight facies of the Wealden remains unsolved.’ A detailed palynologic

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

581

study of the entire Fairlight Clay sequence would be desirable to determine its probable
equivalence to the Purbeck but was not attempted in the present study. The preliminary
results of a study by Hughes and Moody-Stuart (1969) using a new correlation method,
however, suggest that the Fairlight Clay is in fact a lateral facies equivalent of the upper
Middle and Upper Purbeck.

EXTRACTION OF PALYNOMORPHS

Most samples examined in this study were grey or brown fine-grained clastic rocks
but a few limestones were represented. These samples were prepared by first crushing
about 5 gm. (20-50 gm. in the case of limestones) to pass through a 1-mm. mesh sieve.
Carbonates were removed with cold dilute hydrochloric acid. Silica and silicates were
dissolved in hot 50% hydrofluoric acid (40-60 minutes treatment was sufficient). By-
products from this reaction are finely crystalline or in part colloidal, gelatinous white,
pink, or brown precipitates usually occurring as a distinct layer or intimately mixed
with the organic residue. This by-product was reported by Norem (1953) to be composed
of either aluminium fluosilicates or a mixture of double fluorides of calcium, magnesium,
sodium, and potassium. It interferes with subsequent preparation procedures and
consequently must be removed after hydrofluoric acid treatment by repeated washing
and dissolution in hot 25% hydrochloric acid.

The organic residue remaining was treated with 20 kHz ultrasound for 20-30 seconds
using a generator with a 60-W power output and a magnetostrictive transducer coupled
to a steel probe with a 1 : 1 end area ratio. The probe tip has an acoustic power output of
2-74 W/cm2. Acoustic treatment was used to disperse the finely divided organic material
which characterized most residues. This finely divided material was removed from each
residue after ultrasonic treatment by repeated short centrifugation in water with a few
drops of non-ionic detergent added (Funkhouser and Evitt 1959). Ultrasonic treatment
was used before oxidation of the residues because spores and pollen are more fragile to
ultrasound after oxidation (McIntyre and Norris 1964).

Most samples were not highly carbonaceous and the residues did not require more
than a few minutes oxidation in concentrated nitric acid or Schulze’s solution. Humic
material remaining after oxidation was rare in most samples, but when present dilute
ammonium hydroxide removed it effectively. Safranine-O was used to stain some paly-
nomorph assemblages. Residues were mounted in glycerine jelly and stored in a mixture
of glycerine and water with phenol added to prevent microbial attack.

Rock samples used in this study are stored in the Sedgwick Museum, Cambridge.
Full locality details are listed for all samples in Appendix 1 of Norris (1963). Type
material currently in the palynology collection of the Department of Geology, Uni-
versity of Toronto, will be transferred to the Sedgwick Museum for permanent storage.

SYSTEMATIC PALYNOLOGY— SPECIES LIST

The species mentioned in the following list occur in the marine Upper Jurassic and
Purbeck Beds. The species are listed morphographically using Dettmann’s (1963)
modified scheme. Where applicable the original author’s name is followed by the plate

582

PALAEONTOLOGY, VOLUME 12

and figure number of the illustrations given in this paper. New species and new combina-
tions are treated thoroughly in the following section on Systematic Descriptions. The
stratigraphic significance of all species is discussed in a later part of the paper.

Turma triletes

Suprasubturma acavatitriletes
Subturma azonotriletes
Infraturma laevigati

Cyathidites australis Couper 1953; Plate 102, fig. 1.

Cyathidites minor Couper 1953; Plate 102, figs. 2, 3.

Deltoidospora rafaeli Burger 1966; Plate 102, fig. 11.

Deltoidospora psilostoma Rouse 1959; Plate 102, fig. 8.

Dictyophyllidites harrisii Couper 1958; Plate 102, figs. 9, 10.

Dictyophyllidites equiexinus (Couper) Dettmann 1963; Plate 102, figs. 4, 5.
Stereisporites antiquasporites (Wilson and Webster) Dettmann 1963; Plate 102, figs. 13,
14.

Concavisporites juriensis Balme 1957; Plate 102, figs. 6, 7.

Divisisporites sp. cf. D. euskirchenensis Thomson and Pflug 1952; Plate 102, fig. 17.

Infraturma apiculati

Acanthotriletes varispinosus Pocock 1962; Plate 102, fig. 12.

Osnumdacidites wellmanii Couper 1953; Plate 102, fig. 18.

Baculatisporites comaumensis (Cookson) Potonie 1956; Plate 102, figs. 15, 16.
Converrucosisporites variverrucatus (Couper) comb. nov. ; Plate 102, fig. 19.
Leptolepidites psarosus sp. nov.; Plate 103, figs. 2-5.

Leptolepidites epacrornatus sp. nov.; Plate 103, figs. 6-9, 11.

Rubinella major (Couper) comb, nov.; Plate 103, fig. 10.

Pilosisporites trichopapidosus (Thiergart) Delcourt and Sprumont 1955; Plate 103,
fig. 1.

Pilosisporites delicatulus sp. nov.; PI. 103, figs. 12-18; Plate 104, figs. 1, 2.

Infraturma murornati

Cicatricosisporites australiensis (Cookson) Potonie 1956.

Cicatricosisporites purbeckensis sp. nov.; Plate 104, figs. 5-11.

Cicatricosisporites angicanalis Do ring 1965; Plate 104, figs. 12-13; Plate 105, figs.

1,2.

Cicatricosisporites brevilaesuratus Couper 1958; Plate 105, fig. 3.

Reticulisporites semireticulatus (Burger) comb, nov.; Plate 105, figs. 4, 5.
Lycopodiacidites cerniiditcs (Ross) comb, nov.; Plate 105, figs. 6, 7.

Lycopodiumsporites austroclavatidites (Cookson) Potonie 1956; Plate 105, figs. 8, 9.
Klukisporites pseudoreticulatus Couper 1958; Plate 105, fig. 11.

Microreticulatisporites diatretus sp. nov.; Plate 105, figs. 12-15.

Foveosporites canalis Balme 1957; Plate 106, fig. 3.

Tripartina sp. ; Plate 105, fig. 10.

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

583

Subturma zonotriletes
Infraturma auriculati

Trilobosporites bernissartensis (Delcourt and Sprumont) Potonie 1956; Plate 106, figs.

1,2.

Trilobosporites apiverrucatus Couper 1958; Plate 107, figs. 9, 14.

Trilobosporites obsitus sp. nov. ; Plate 106, figs. 7, 8.

Trilobosporites domitus sp. nov.; Plats 106, figs. 9, 10, 12, 13.

Appendicisporites potomacensis Brenner 1963; Plate 107, figs. 1-4, 7, 10.

Plicatella abaca (Burger) comb, nov.; Plate 106, figs. 4-6, 11 ; Plate 107, figs. 5, 6.

Infraturma cingulati

Foraminisporis wonthaggiensis (Cookson and Dettmann) Dettmann 1963; Plate 107,
figs. 11, 13.

Contignisporites dorsostriatus (Bolchovitina) Dettmann 1963; Plate 107, fig. 12.
Duplexisporites problematicus (Couper) Playford and Dettmann 1965.

Infraturma tricrassati

Gleicheniidites senonicus Ross 1949; Plate 107, figs. 16, 17.

Sestrosporites pseudoalveolatus (Couper) Dettmann 1963; Plate 108, fig. 5.
Coronatispora valdensis (Couper) Dettmann 1963; Plate 108, figs. 1, 2.

Suprasubturma perinotriletes

Densoisporites perinatus Couper 1958; Plate 108, figs. 3, 4, 6.

Heliosporites sp. ; Plate 108, figs. 7, 8, 10, 11.

Turma hilates

Aequitriradites spinulosus (Delcourt and Sprumont) Cookson and Dettmann 1961;
Plate 108, fig. 9.

Couperisporites complexus (Couper) Pocock 1962; Plate 108, fig. 13.

Januasporites tumulosus sp. nov.; Plate 108, fig. 12; Plate 109, figs. 2-4, 7.

Turma monoletes
Suprasubturma acavatomonoletes
Subturma azonomonoletes
Infraturma sculptatomonoleti

Marattisporites scabratus Couper 1958; Plate 109, figs. 5, 6.

Anteturma pollenites
Turma saccites
Subturma monosaccites
Infraturma saccizonati

Cerebropollenites mesozoicus (Couper) Nilsson 1958; Plate 109, figs. 11, 12.

584 PALAEONTOLOGY, VOLUME 12

Subturma disaccites

Alisporites bilateralis Rouse 1959; Plate 109, figs. 14, 15.

Abietineaepollenites minimus Couper 1958; Plate 109, fig. 13.

Vitreisporites pallidus (Reissinger) Potonie 1960; Plate 109, figs. 8-10.

Podocarpidites sp. cf. P. eUipticus Cookson 1947; Plate 109, figs. 16, 17.

Parvisaccites radiatus Couper 1958; Plate 109, figs. 18, 19; Plate 9, fig. 1.

Subturma polysaccites

Callialasporites sp. cf. C. trilobatus (Balme) Sukh Dev 1961; Plate 110, fig. 8.
Callialasporites dampieri ( Balme) Sukh Dev 1961 emend.; Plate 110, figs. 2, 3.
Callialasporites obrutus sp. nov. ; Plate 1 10, figs. 6, 7.

Callialasporites sp.; Plate 110, figs. 4, 5.

Turma aletes
Infraturma psilonapiti

Inaperturopollenites dubius (Potonie and Venitz) Thomson and Pfiug 1953; Plate 110,
figs. 9, 10; Plate 111, fig. 19.

Inaperturopollenites sp.; Plate 110, figs. 11, 12.

Infraturma granulonapiti

Araucariacites australis Cookson 1947; Plate 110, fig. 17.

Spheripollenites subgranulatus Couper 1958; Plate 110, fig. 13.

Infraturma spinonapiti

Peltandripites tener sp. nov.; Plate 110, figs. 18, 19.

Infraturma reticulonapiti

Undulatasporites araneus sp. nov.; Plate 110, figs. 14-16; Plate 111, figs. 2-10.

Turma plicates
Subturma praecolpates

Eucommiidites troedssonii Erdtman 1948; Plate 111, figs. 13, 14, 16.

Eucommiidites minor Groot and Penny 1960; Plate 111, fig. 15.

Subturma monocolpates

Cvcadopites sp. cf. C. nitidus (Balme) comb, nov.; Plate 111, figs. 11, 12.

Cveadopites carpentieri (Delcourt and Sprumont) Singh 1964; Plate 111, fig. 18.
Monosulci tes sp. aff. M. minimus Cookson 1947; Plate 111, fig. 17.

Turma poroses
Subturma monoporines

Exesipollenites scabrosus sp. nov.; Plate 111, figs. 20-2.

Perinopollenites elatoides Couper 1958; Plate 112, figs. 6, 7.

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

585

MIOSPORES INCERTAE SEDIS

Classopollis torosus (Reissinger) Balme 1957; Plate 112, figs. 1-5.

ClassopoUis echinatus Burger 1965; Plate 1 12, figs. 8-13.

Classopollis hammenii Burger 1965; Plate 112, figs. 14-16; Plate 113, figs. 1-4.
Sehizosporis reticulatus Cookson and Dettmann 1959; Plate 113, figs. 5, 8.
Sehizosporis spriggi Cookson and Dettmann 1959; Plate 113, figs. 6, 13.
Sehizosporis parvus Cookson and Dettmann 1959; Plate 113, fig. 7.
Sigmopollis callosus sp. nov. ; Plate 113, figs. 9-12.

SYSTEMATIC DESCRIPTIONS
Turma triletes

Suprasubturma acavatitriletes
Subturma azonotriletes
Infraturma laevigati

Genus divisisporites Thomson and Pflug 1952
Divisisporites sp. cf. D. euskirchenensis Thomson and Pflug 1952

Plate 102, fig. 17

Description. Spores radiosymmetric, complexly trilete. Amb triangular convex to
irregularly sub-circular. Laesurae long, simple, straight but dichotomozing up to the
third order, reaching or almost reaching the equator. Both proximal and distal surfaces
scabrate to granular. Granules usually low (occasional up to 1 p high), closely spaced, up
to 2 p in diameter. Occasional verrucae developed at the equator up to 3 p high and 10 p
or more in width. Exine 3-5 p in total thickness; endexine distinct and 0-5-1 p thick.

Dimensions (3 specimens). Equatorial diameter: 53-69 p.

Distribution. Upper Purbeck, Dorset.

Remarks. These specimens differ from D. euskirchenensis by the possession of a thicker
exine, more complexly divided laesurae, and verrucae at the equator. Insufficient
specimens were available to erect a new species. Lower Cretaceous specimens originally
assigned to D. euskirchenensis by Cookson and Dettmann (1958) are now included in
Rouseisporites radiatus Dettmann (1963), which is distinct in structure from the present
specimens.

Infraturma apiculati

Genus converrucosisporites Potonie and Kremp 1954
Converrucosisporites variverrucatus (Couper) comb. nov.

Plate 102, fig. 19

1958 Concavisporites variverrucatus Couper, p. 142, pi. 22, figs. 4-5.

Remarks. This species is removed from Concavisporites because of the lack of curvaturae,
and recombined with Converrucosisporites on the basis of shape and ornament.

Q9

C 6940

586

PALAEONTOLOGY, VOLUME 12

Genus leptolepidites Couper 1953 emend. Norris 1968
Leptolepidites psarosus sp. nov.

Plate 103, figs. 2-5

Holotype. GN 109B/1, 40.4 124.5. Sample 59-1-6 (dark grey shale), Middle Purbeck, Chief Beef Beds,
Durlston Bay (from the top of Bed 75, Bristow 1857).

Diagnosis. Spores radiosymmetric, trilete, amb rounded triangular to circular. Laesurae
simple, reaching the equator, usually indistinct. Proximal face entirely granulate, or
only on the contact areas. Distal face with closely spaced verrucae. Exine thin in be-
tween the projections.

Description. Laesurae straight, reaching or almost reaching the equator, occasionally
quite distinct but usually difficult to see. Granules on proximal face rounded or poly-
gonal, 1-2 p in diameter, dense, occasionally joining up to give a sub-rugulate sculpture,
rugulae about 4 p long. When only contact faces are granular there is a distinct levigate
zone separating them from the verrucate distal ornament.

Distal verrucae 3-13 p in diameter, 1-5 p high, rounded sub-circular, rounded poly-
gonal, irregular or elongated. Solitary rugulae up to 17 p long and about 5 p wide may
be interspersed among the verrucae. Verrucae closely packed forming a distinct negative
reticulum with grooves about 0-5 yu. wide but varying slightly. The distal verrucae may
encroach to a variable extent on to the proximal face up to the contact areas.

Exine between projections less than 0-5-1 p in thickness. Exine on proximal face less
than 0-5 p in thickness.

Dimensions. Equatorial diameter: 20-44 p (holotype, 39 p).

Distribution. Middle and Upper Purbeck, Dorset, Sussex, and Surrey.

Remarks. Distinguished from Converrucosisporites proxigranulatus Brenner (1963) by
the more densely packed distal verrucae and by the presence of rugulae.

EXPLANATION OF PLATE 102
All figures X 750 unless otherwise stated.

Figs. 1-3. Cyathidites spp. 1, C. australis Couper, GN 161/1, 32.0 127.4. 2-3, C. minor Couper.

2, GN 317/2, 44.2, 128.5. 3, GN 138/2, 58.2 123.1.

Figs. 4, 5, 9, 10. Dictyophvllidites spp. 4-5, D. equiexinus (Couper) Dettmann. 4, GN 146/1, 24.7
128.3. 5, GN 161/2,’ 53.0 117.7. 9-10, D. harrisii Couper. 9, GN 316/1, 32.6 114.9. 10, GN 316/1,
58.0 124.8.

Figs. 6, 7. Concavisporites juriensis Balme. 6, GN 142/2, 43.1 112.5. 7, GN 142/1, 40.1 119.6.

Figs. 8, 11. Deltoidospora spp. 8, D. psilostoma Rouse. 11 13-1 3c, 38.3 116.9. 11 , D. rafaeli Burger,
GN 262/2, 32.2 107.5.

Fig. 12. Acanthotriletes varispinosus Pocock, GN 188/1, 44.0 114.5; X 1250.

Figs. 13, 14. Stereisporites antiquasporites (Wilson and Webster) Dettmann. 13, GN 476/1, 51.8 112.5;
X 1250. 14, GN 187/1, 29.5 108.6; X 1250.

Figs. 15, 16. Baculatisporites comaumensis (Cookson) Potonie. 15, 11 13-1 8c, 22.6 118.9. 16, GN 432/2,
46.9 104.3.

Fig. 17. Divisisporites sp. cf. D. euskirchenensis Thomson and Pflug, GN 345/1, 117.3 56.2.

Fig. 18. Osnmndacidites wellmanii Couper, GN 316/1, 22.2 106.6,

Fig. 19. Converrucosisporites variverrucatus (Couper) comb, nov., GN 338/3, 50.9 124.1.

Palaeontology, Vol. 12

PLATE 102

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

587

Leptolepidites epacroniatus sp. nov.

Plate 103, figs. 6-9, 1 1

Holotype. Slide GN 147/1, 27.8 119.1. Sample 60-5-22 (dark grey-brown shale). Upper Purbeck,
Marble Beds and Ostracod shales, Durlston Bay (6 ft. below the top of Bed 84, Bristow 1857).

Diagnosis. Spores radiosymmetric, trilete. Amb rounded triangular to circular. Laesurae
long but not reaching equator, simple or labiate. Proximate face levigate to subgranular.
Distal face verrucate with occasional echinate projections. Exine thin.

Description. Laesurae simple or with very narrow lips less than 0-75 p wide, straight or
sinuous. Ornament of proximal face very much reduced with respect to distal ornament,
but distal ornament encroaches on to proximal face at apices. Distal verrucae regularly
or irregularly rounded or elongate, 2-3 p in diameter, sometimes reaching 5 p long, up to
1 p high and spaced about 1 p apart. Exine 0-25-0-5 p.

Dimensions. Equatorial diameter: 12-22 p (holotype 20 p).

Distribution. Middle and Upper Purbeck, Dorset, Sussex, and Surrey.

Genus RubineUa (Maljavkina 1949) Potonie 1960
RubineUa major (Couper) comb. nov.

Plate 103, fig. 10

1958 Leptolepidites major Couper, p. 141, pi. 21, figs. 7-8.

Description. Spores radiosymmetric, trilete. Amb rounded triangular or occasionally
almost circular. Laesurae long, simple, not reaching the equator, rather indistinct. Both
proximal and distal surfaces ornamented with closely spaced or touching, almost
spherical, irregularly rounded or elongated verrucae. Verrucae 2-10 p in diameter,
1-5 p high. Exine thickness between verrucae 1-2-5 p.

Dimensions. Equatorial diameter: 39-80 p.

Remarks. It proved impossible to split off the larger specimens as a distinct species and
consequently all are included in one species with a larger size range than that indicated
by Couper in his original description. The species is transferred to RubineUa because of
the comprehensive verrucate sculpture. Leptolepidites is characterized by verrucae on
the distal face only (Norris 1968).

Genus pilosisporites Delcourt and Sprumont 1955
Pilosisporites deiicatulus sp. nov.

Plate 103, figs. 12-18; Plate 104, figs. 1, 2

Holotype. GN 428/1, 46.0 128.4 Sample 61-7-5 (bufiflignitic clay), Upper Purbeck 7 ft. above Paludina
Clays, Bacon Hole (middle of Bed 26 described in Norris 1963).

Diagnosis. Spores radiosymmetric, trilete. Amb triangular convex with broadly rounded
apices, occasionally becoming almost circular. Laesurae short, one-third to one-half of
the spore radius, straight, simple, frequently indistinct. Both proximal and distal
surfaces scabrate and covered with irregularly distributed echinulate processes. Pro-
cesses hair-like, very narrow, straight or curved, simply terminated or briefly bifurcate,

588 PALAEONTOLOGY, VOLUME 12

2-5 , u long, spaced 1-2 p apart at the equator but up to 5 p apart at the poles. Exine
0-5—1 /x thick.

At high magnifications (greater than 1,000 diameters) the exine is seen to be dis-
tinctly microreticulate. Lumina of reticulum less than 0-25 p in diameter and muri also
very narrow. Echinulate processes very narrow and consequently indistinct. Occasionally
the bases of the processes may widen up to 0-5 p and can be seen to be hollow.

Dimensions. Equatorial diameter: 28-40 /x (holotype 30 p).

Distribution. Upper Purbeck of Dorset.

Remarks. This species is easily overlooked on account of its delicate ornament, or it
may be confused with Stereisporites antiquasporites (Wilson and Webster) Dettmann.

Infraturma murornati

Genus cicatricosisporites Potonie and Gelletich 1933
Cicatricosisporites purbeckensis sp. nov.

Plate 104, figs. 5-11

Holotype. Slide GN 145/1, 38.8 121.9. Sample 60-5-16 (grey, slightly calcareous shale) Upper Purbeck,
Marble Beds and Ostracod Shales, Durlston Bay (Bed 84 of Bristow 1857).

Diagnosis. Spores radiosymmetric, trilete. Amb. triangular. Laesurae long, straight,
simple or labiate. Proximal face levigate. Distal face with 3 or 4 triangular sets of
widely spaced ribs running more or less parallel to equator. Ribs narrow and uneven in
width, height, and spacing.

Description. Laesurae simple or bordered by very narrow lips about 0-5 p wide, reaching
equator. Distal ribs straight or sinuous, occasionally bifurcating, 0-25-1 /x wide, 0-5-1 p
high, spaced 0-5-2 p apart (4 ribs and intervening lumina measure 9-12 p). Ribs project
at apices but not on sides of amb. Distal ribs encroach on to proximal face at apices.
Ribs are characteristically uneven in width with swollen nodes at irregular intervals
along their length. Exine about 1 p thick, with a very thin layer of endexine distinguishable.

Dimensions. Equatorial diameter 30-48 p (holotype 47 p).

Distribution. Lower, Middle, and Upper Purbeck of Dorset and Sussex.

EXPLANATION OF PLATE 103

All figures X 750 unless otherwise stated.

Figs. 1, 12-18. Pilosisporites spp. 1, P. trichopapiUosus (Thiergart) Delcourt and Sprumont, GN 338/2,
30.6 117.4. 12-18, P. delicatulus sp. nov. 12, proximal surface, GN 427/1, 56.6 124.9. 13-14, Holo-
type, median focus and distal surface respectively, GN 428/1, 46.0 128.4. 15-16, Holotype, median
focus, and distal surface respectively. X 1,250. 17-18, Proximal and distal surfaces respectively,
GN 338/2, 119.7 22.2; X 1,250.

Figs. 2-9, 11. Leptolepidites spp. 2-5, L. psarosus sp. nov. 2, Proximal surface, GN 428/1, 128.5 37.0.
3, Distal surface, GN 338/3, 36.2 1.262. 4-5, Holotype, proximal and distal surfaces respectively,
GN 109b/1, 40.4 124.5. 6-9, 11, L. epacrornatus sp. nov. 6, 8, Holotype, x 750 and x 1,250 respec-
tively, GN 147/1, 27.8 119.1. 7, GN 146/2, 52.6 125.3. 9, GN 146/2, 52.7 125.5. 11, GN 190/3, 41.6
108.9.

Fig. 10. Rubinella major (Couper) comb, nov., GN 255/1, 41.7 1 19.4.

Figs. 19, 20. Cicatricosisporites australiensis (Cookson) Potonie. 19, GN 142/1, 54.8 108.7. 20, GN 315/3,
45.5 122.8.

. Palaeontology , Vol. 12

PLATE 103

('•A »>
Cl' IvV

*>x

L«i

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

589

Remarks. Distinguished from other species of Cicatricosisporites by means of the narrow
ribs of uneven width and spacing.

Genus reticulisporites Pot. and Kremp in Weyl. and Krieger 1953
Reticulisporites semireticulatus (Burger 1966) comb. nov.

Plate 105, figs. 4, 5

1966 Lycopodiumsporites semireticulatus Burger, p. 247, pi. 14, fig. 4.

Remarks. The concave amb and low muri place this species in the genus Reticulisporites.

Genus lycopodiacidites (Couper 1953) Potonie 1956
Lycopodiacidites cerniidites (Ross 1949) comb. nov.

Plate 105, figs. 6, 7

1949 Lycopodium cerniidites Ross, p. 30, pi. 1, figs. 1-2.

1955 Lycopodiumsporites cerniidites (Ross) Delcourt and Sprumont, p. 32.

Remarks. The distal surface is rugulate, not reticulate, thus placing this species in Lyco-
podiacidites. There appears to be a morphological transition between those forms of L.
cerniidites (Ross) with more regularly arranged regulae, to forms of Coronatispora
valdensis (Couper) Dettmann with poorly developed circum-equatorial ridges. However
the latter species has a very much greater stratigraphic and geographic distribution in the
sediments examined and it is likely that each species was derived from different sources,
their morphological similarity being only apparent. Reticu/atisporites pudens Balme
from the Lower Cretaceous of Western Australia is similar but much smaller and carries
an imperfect reticulum on the distal face.

Genus microreticulatisporites (Knox 1950) Potonie and Kremp 1954
Microreticulatisporites diatretus sp. nov.

Plate 105, figs. 12-1 5

Holotype. Slide GN 148/1, 33.2 125.6. Sample 60-5-24 (grey shaley clay), Upper Purbeck Unio Beds,
Durlston Bay (Bed 82 of Bristow 1857).

Diagnosis. Spores radiosymmetric, trilete. Amb rounded triangular. Laesurae one-half
to one-third of the spore radius in length, usually with narrow lips. Both proximal and
distal surfaces ornamented with a perfect microreticulum with rather variable circular
to polygonal lumina never exceeding 2 p in diameter. Exine 1-2 p thick interradially,
thinning to 0-75-1 p at apices.

Description. Amb occasionally elongated along one median. The trilete scar is rather
variable in development, occasionally being simple or indistinct. Muri of the micro-
reticulum 0-5-1 p wide, 0-3— 1 -5 p in height, of slightly variable width usually widening
at the junctions. Lumina 0-5-2 p in diameter, rounded or rounded-polygonal, varying
in spacing from moderately widely spaced pits to closely spaced rounded-polygonal
lumina. The lumina of the proximal reticulum may be slightly radially attenuated, par-
ticularly near the ends of the laesurae.

590 PALAEONTOLOGY, VOLUME 12

Dimensions. Equatorial diameter: 30-40 p.

Distribution. Middle and Upper Purbeck, Dorset and Sussex.

Remarks. Distinguished from Foveotriletes subtriangularis Brenner 1963 by the micro-
reticulate ornament on both surfaces rather than foveolate ornament primarily de-
veloped on the distal surface.

Genus tripartina Maljavkina 1949 ex Potonie 1960
Tripartina sp.

Plate 105, fig. 10

Description. Spores radiosymmetric, trilete. Amb triangular, usually concave or straight-
sided, slightly undulating. Laesurae long, reaching the equator, with lips about 2 p
wide. Proximal surface unornamented. Distal surface with irregular radial grooves
0-5-2 p wide, spaced 1-2 p apart, occasionally anastomosing or coalescing with
adjacent grooves. Both proximal and distal face may be slightly undulose. Exine 1-2 p
thick.

Dimensions. Equatorial diameter 24-46 p.

Distribution. Middle and Upper Purbeck of Dorset, Sussex, and Surrey.

Remarks. This species is distinguished from T. sp. cf. T. variabilis Maljavkina described
by Dettmann (1963) from the Australian Cretaceous by the more broadly rounded apices
and less dense distal ornament.

Subturma zonotriletes
Infraturma auriculati

Genus trilobosporites Pant 1954 ex Potonie 1956
Trilobosporites obsitus sp. nov.

Plate 106, figs. 7, 8

Holotype. Slide GN 163/2, 59.2 127.5. Sample 60-5-1 (buff, silty, calcareous clay), Upper Purbeck,
Unio Beds, Durlston Bay (Bed 80 of Bristow 1857).

Diagnosis. Spores radiosymmetric, trilete. Amb triangular. Laesurae long, usually
simple, but rather variable. Both proximal and distal faces verrucate, but sculpture
sparse and more reduced on distal face. Apices each carry a very large and prominent
thickening which is circular in equatorial view and is restricted to the apical equatorial
region. Exine thick.

EXPLANATION OF PLATE 104

All figures X 750 unless otherwise stated.

Figs. 1, 2. Pilosisporites delicatulus sp. nov. 1, Median focus, GN 338/2, 122.3 23.0; X 1,250. 2, Proxi-
mal surface, GN 428/2, 31.4 108.3; x 1,250.

Figs. 3-13. Cicatricosisporites spp. 3, 4, C. australiensis (Cookson) Potonie, Distal and proximal
surfaces respectively; GN 146/2, 32.0 1 1 1.8. 5-1 1, C. purbeckensis sp. nov. 5, Proximal surface, GN
153/2, 29.3 119.7. 6, Holotype, median focus, GN 145/1, 38.8 121.9. 7, Median focus, GN 146/2,
20.9 113.2. 8, Median focus, GN 146/2, 44.7 108.2. 9, Equatorial view, GN 153/1, 21.9 109.0. 10,
Equatorial view, GN 142/2, 23.2 126.7. 11, Distal surface, GN 147/2, 32.5 125.6. 12-13, C. cingi-
canalis Doring, proximal and distal surfaces respectively, GN 163/2, 40.8 122.7.

Palaeontology, Vol. 12

PLATE 104

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

59J

Description. Amb with roughly straight or slightly concave sides which are undulating
because of verrucate ornament. Apices rounded and extended by the apical projections.
Laesurae may be narrowly labiate, two-thirds or more of radius in length, occasionally
reaching the equator. Proximal face with dense, rounded, or irregular verrucae, 6-12 p
in diameter, 2-3 p high, and spaced up to 3 p apart. Occasionally they may fuse into
large indistinct rugulae. Distal ornament variable, ranging from low, indistinct, sparse
irregular granules about 3 p in diameter to rounded, low or high sparse verrucae up to
11 /x in diameter. Apical thickenings usually restricted to the equator but occasionally
extending about 4 p towards the polar areas. Exine thickness 8-10 p over the apical
thickening which is 14-20 p wide and has a rounded aspect in equatorial views. Endexine
0*75—1 *5 p thick. Ektexine 1-5-3 p thick between the projections.

Dimensions. Equatorial diameter 56-69 p (holotype 63 p).

Distribution. Upper Purbeck, Dorset.

Trilobosporites domitus sp. nov.

Plate 106, figs. 9, 10, 12, 13

Holotype. Slide GN 345/1, 57.0 111.7. Sample 60-19-3 (grey calcareous marl), Upper Purbeck, Upper
Cypris Clays and Shales, Bacon Hole (6 in. above Bed 1 of Arkell 1933).

Diagnosis. Spores radiosymmetric, trilete. Amb triangular, concave. Laesurae long,
straight, labiate, commissures raised. Both proximal and distal faces entirely covered in
closely spaced, low, irregular granules. Exine at apices undulating and usually slightly
thickened in the equatorial region here, but these thickenings not extending polewards
as distinct valvae. Exine thicker on proximal than distal face.

Description. Laesurae almost reach the equator, raised 1 -5-2-5 p high at the centre and
provided with narrow (1 p) tapering lips along at least half their length. Scabrate to
sub-granular ornament on both proximal and distal surfaces. Granules rather irregular
in outline, very low, 0-25-1 p in diameter and spaced not more than 0-5 p apart. The
exine becomes rather undulating around the apices tending to assume a low, poorly
developed, verrucate ornament, and also is slightly thickened. Exine 2-4 p thick, in-
creasing at apices to between 4 and 7 p. Exine on proximal face distinctly thicker than on
distal face, about 6 p at the centre. Endexine 0-25-0-5 p thick. Occasionally the exine is
slightly thickened interradially at the middle of the concave sides. Both proximal and
distal faces convex, proximal face rather flatter than distal.

Dimensions. Equatorial diameter : 56-80 p (holotype 65 p).

Distribution. Upper Purbeck, Dorset.

Genus plicatella Maljavkina 1949
Plicatella abaca (Burger) comb. nov.

Plate 106, figs. 4-6, 11; Plate 107, figs. 5, 6
1966 Cicatricosisporites abacus Burger, p. 242, pi. 7, fig. 3.
Distribution. Middle and Upper Purbeck, Dorset, Sussex, and Surrey.

592

PALAEONTOLOGY, VOLUME 12

Remarks. This species is transferred to Plicatella on account of the apical thickenings at
the equator. The distinction of Plicatella , characterized by weak apical thickenings, from
Appendicisporites, with more prominent apical thickenings, is arbitrary but a convenient
procedure. Hughes and Moody-Stuart (1969) have questioned the validity of the genus
Plicatella on the basis of observations on associated spores of Cretaceous schizeaceous
ferns. They have noted that occasional apical thickenings occur in populations of
Cicatricosisporites-typc spores. These thickenings may be in part the result of compres-
sion in the equatorial plane. Dispersed spores, however, are difficult to relate to associ-
ated spore populations. Plicatella abaca has a distinctive morphology and distribution
and appears to be a discrete group. Consequently it is maintained as a distinct spore
species.

Suprasubturma perinotrilites
Genus heliosporites Schulz 1962
Heliosporites sp.

Plate 108, figs. 7, 8, 10, 11

Description. Spores radiosymmetric, trilete, zonate, consisting of a central body and an
outer more complex layer. Amb of outer layer, excluding the zone, convex triangular
with broadly rounded apices. Amb of inner body convex triangular with more sharply
rounded or pointed apices. Inner body excentrically placed in relation to outer layer.
Laesurae of inner body long, straight, simple, reaching the equator, occasionally in-
distinct. Laesurae of outer layer long, sinuous, labiate, reaching equator and frequently
the outer zone. Lips 1 p wide, slightly tapering, with fibrilar structure similar to that of
outer layer. Equatorial zone 4-7 p wide, roughly parallel to amb of outer layer, 2 p
thick and tapering very slightly towards its outer edge which is smooth except for
occasional projecting spines from the dorsal surface. Entire distal surface of outer
layer, including the distal surface of the zone, ornamented with irregularly distributed
spines, 3-5 p high, bases roughly circular and 3-6 p wide, spaced 4-7 p apart (tips
spaced 8 p apart), tapering rapidly at base but more gradually towards tips which are
pointed, truncate or occasionally bifurcate. Exine of inner layer levigate, 0-25 p or less in
thickness, apparently of simple, undifferentiated structure, frequently folded, particu-
larly near equator. Outer layer including zone, spines, and trilete mark with a fibrilar
structure. Fibrils about 0-25 p thick and anastomosing to form a ‘three-dimensional’

All figures X 750.

EXPLANATION OF PLATE 105

Figs. 1-3. Cicatricosporites spp. 1-2, C. angicanalis Doring. 1, Proximal surface, GN 140/2, 42.6
128.5. 2, Distal surface, GN 145/2, 48.0 128.6. 3, C. brevilaesuratus Couper, GN 265/1, 39.8 112.9.
Figs. 4, 5. Reticulisporites semireticulatus (Burger) comb. nov. 4, Median focus, GN 189/2, 46.0 122.5.
5, Distal surface, GN 195/1, 21.1 110.8.

Figs. 6, 7. Lvcopodiacidites cerniidites (Ross) comb. nov. 6, Proximal surface, GN 147/1, 29.8 128.2. 7,
Distal surface, GN 421/1, 111.8 60.8.

Figs. 8, 9. Lycopodiumsporites austroclavatidites (Cookson) Potonie. 8, Proximal surface, GN 147/2,
43.4 128.2. 9, Distal surface, GN 154/1, 29.2 127.8.

Fig. 10. Tripartina sp., distal surface, GN 182/3, 26.0 101.4.

Fig. 11. Klukisporites pseudoreticulatus Couper, GN 148/1, 45.3 119.5.

Figs. 12-15. Microreticulatisporites diatretus sp. nov. 12, Holotype, proximal surface, GN 148/1,
33.2 125.6. 13, Distal surface, GN 341/2, 26.8 121.6. 14, GN 145/1 121.3 39.2. 15, Proximal surface,,
GN 153/2, 34.7 111.8.

Palaeontology, Vol. 12

PLATE 105

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

593

reticulum with lumina 0-25-1 p in diameter. Outer layer very thin on proximal sur-
face where its fibrilar nature is difficult to determine. The outer layer is 1-2 p thick at
equator and on distal surface.

Dimensions. Equatorial diameter (including zone) : 30-42 p.

Distribution. Upper Purbeck, Dorset.

Remarks. Insufficient material was available to erect a new species. It was considered
appropriate to describe the specimens in detail, however, because they are extremely
distinctive and possibly of stratigraphic importance.

These specimens are tentatively referred to Heliosporites. Schultz interpreted the
type material of this genus as possessing a distal perispore. He did not distinguish a zone.
This latter feature, together with the restriction of the spinose ornament to the distal
surface, suggests affinities with the genus Styxisporites Cookson and Dettmann. This
genus, however, does not possess a central spore body.

Considerable difficulty was encountered in interpreting the structure of these speci-
mens owing, firstly, to the indistinct appearance of the central body; secondly, to the
thinness of the proximal surface of the outer layer; and thirdly, to the fibrilar nature of
this layer and the zone. All these features make elucidation of the equatorial structure,
in particular, very difficult. Consequently this interpretation must be considered tenta-
tive until further specimens are available. In view of this uncertainty these specimens
and the type material of Heliosporites may ultimately prove to have a similar structure.
If this is so, these specimens are distinguished from the Lower Jurassic Heliosporites
altmarkensis Schulz by their rather small size, thinner exine of the inner spore body,
and rather shorter distal spines which are not truncated.

Turma hilates

Genus januasporites (Pocock 1962) Singh 1964
Januasporites tumulosus sp. nov.

Plate 108, fig. 12; Plate 109, figs. 2-4, 7

Holotype. Slide GN 421/1, 55.5 123.3. Sample 61-6-2 (grey, clayey shale), Upper Purbeck, Paludina
Clays, Lulworth Cove, Dorset (from 2 ft. 6 in. below the highest Viviparus limestone).

Diagnosis. Spores radiosymmetric, probably trilete, with a distinct central body. Amb
rounded triangular to circular or oval. Inner body thicker and ornamented with a perfect
polygonal microreticulum and usually carrying a large circular aperture on one face.
Outer layer is very thin and loosely fitting around the antapical face, projecting beyond
the central body at the amb and ornamented with irregularly distributed granules
which are distinctly raised on hollow protuberances. Exine layers usually very thin and
of indeterminate thickness.

Description. Exine is composed of two distinct layers, an inner thicker and an outer,
thinner membraneous layer almost completely enclosing the inner body. The inner
layer bears an irregular but perfect polygonal micro-reticulum; muri 0-25-0-75 p wide;
lumina 1-3 p wide and of polygonal shape but occasionally irregularly rounded. Reticu-
lum always present at centre of antapertural face but usually becoming indistinct
towards edges, sometimes showing a trilete distribution by development of stronger,

594

PALAEONTOLOGY, VOLUME 12

elongated lumina along 3 rays. The outer layer is very thin and is covered with rounded
granules about 1-2 p high and varying from 0-5-2 p in diameter. Granules are distinctly
raised on protuberances of the outer layer giving a characteristic L-O-L pattern in
surface view. Granules spaced 1-2 p apart and irregularly distributed. Occasional
gemmate or papillate projections may be distributed amongst the granules. Outer layer
loosely lies over the antapertural face and projects 1-6 p from the amb. It occasionally
encroaches up to the edges of the aperture but usually becomes indistinct on the aper-
tural face.

Inner layer about 0-25 p thick but usually indeterminate; outer layer of exine very
thin with no distinguishable optical section visible under oil immersion, excluding the
thickened granules.

Dimensions. Maximum equatorial diameter: 47-62 p (holotype 47 p). Minimum equatorial diameter:
32-52 p (holotype 33 /x).

Distribution. Middle and Upper Purbeck, Dorset.

Remarks. The outer layer of Januasporites is not referred to in the description as either
a perine or a saccus since the stratification of the inner layer is not visible. This outer
layer is only loosely attached on the antapertural face but becomes closely attached to
the apertural face at or just beyond the amb.

Anteturma pollenites
Turma saccites
Subturma disaccites

Genus podocarpidites (Cookson 1947) ex Couper 1953
Podocarpidites sp. cf. P. ellipticus Cookson 1947

Plate 109, figs. 16, 17

Distribution. Kimmeridgian and Portlandian of Dorset. Lower, Middle, and Upper
Purbeck of Dorset, Sussex, and Surrey.

Subturma polysaccites

Genus callialasporites (Sukh Dev 1961) Potonie 1966
Callialasporites sp. cf. C. trilobatus (Balme 1957) Sukh Dev 1961

Plate 110, fig. 8

Description. Spores radiosymmetric, alete, with a distinct central body. Amb of central
body rounded triangular, surrounded by a distinctly trilobed equatorial saccus constricted

All figures X 750.

EXPLANATION OF PLATE 106

Figs. 1-2, 7-10, 12-13. Trilobosporites spp. 1-2, T. bernissartensis (Delcourt and Sprumont) Potonie-
1, Median focus, GN 345/1, 47.3 127.4. 2, Distal surface, GN 163/2, 51.0 122.9. 7-8, T. obsitus sp.
nov. 7, Holotype, median focus, GN 163/2, 59.2 127.5. 8, Median focus, GN 163/2, 24.2 114.7.
9-10, 12-13, T. domitus sp. nov. 9, Holotype, proximal surface, GN 345/1, 57.0 111.7. 10, Median
focus, GN 345/1, 57.4 120.6. 12, Equatorial view, GN 345/1, 51.6 1 16.5. 13, Median focus, GN 345/1,
38.4 127.8.

Fig. 3. Foveosporites canalis Balme, GN 168/1, 107.6 22.3.

Figs. 4-6, 11. Plicatella abaca (Burger) comb. nov. 4, Median focus, GN 145/2, 49.0 12.49. 5, Median
focus, GN 212/2, 45.9 126.1. 6, GN 261/1, 46.6 116.0. 11, Equatorial view, GN 212/2, 40.4 126.4.

Palaeontology, Vol. 12

PLATE 106

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

595

at the apices of the central body. Exine scabrate or indistinctly wrinkled. Saccus 8-10 p
wide. Exine of saccus and central body indistinct and thin.

Dimensions. Equatorial diameter 50-51 p.

Distribution. Middle and Upper Purbeck, Dorset and Sussex.

Remarks. These grains differ from C. trilobatus (Balme) Sukh Dev in the scabrate
rather than rugulate central body and in the overall smaller size.

Callialasporites dampieri (Balme 1957) Sukh Dev emend.

Plate 1 10, figs. 2, 3

1937 Nelumbium type Simpson, p. 673, fig. 2a.

1957 Zonalapollenites dampieri Balme, p. 32, pi. 8, figs. 88-90.

1958 Zonalapollenites cf. trilobatus Balme; Hughes and Couper, p. 1482, fig. 1 (e).

1958 Zonalapollenites dampieri Balme (partim); Lantz, p. 925, pi. 3, fig. 34.

1958 Zonalapollenites trilobatus Balme (partim); Lantz, p. 925, pi. 4, fig. 37.

1961 Callialasporites dampieri (Balme) Sukh Dev, p. 48.

1962 Pflugipollenites dampieri (Balme) Pocock, p. 72.

Restated diagnosis. Spores radiosymmetric, alete, with a distinct central body. Central
body amb circular to rounded triangular. One face of central body distinctly convex.
Equatorial saccus surrounds central body and imparts a circular outline to the entire
spore. Saccus attached by a narrow area at the equator of the central body, about one-
fifth to one-seventh of the total diameter in width but decreasing and becoming irregu-
larly constricted at the apices of the central body. The saccus is never constricted so
much as to completely separate into three distinct lobes. Saccus usually carries delicate
radial folds which may pass into rugulate folds on the attachment area, but is never
folded on the centre of the spore body. Saccus and spore body scabrate to subgranular.
Saccus wall about 1 p thick. Exine of central body 0-25-0-5 p thick.

Description. Attachment area of saccus 2-8 p wide, distinct or indistinct, sometimes with
a wrinkled appearance due to folding. Bladder usually 8 p wide between apices but
varying from 4-13 p. Constrictions at apices reduce bladder width to the range 1-8 p,
but these are not equal and all three are not necessarily developed on any one spore.

Dimensions. Equatorial diameter: 47-69 p.

Distribution. Kimmeridgian, Portlandian, Lower, Middle, and Upper Purbeck of Dorset, Sussex and
Surrey.

Remarks. C. dampieri (Balme) is similar to C. trilobatus (Balme). Balme (1957) noted
the similarity but in his description of C. dampieri did not mention any constrictions
of the saccus, which however, are clearly shown in his photographs (pi. 8, figs. 88-90).
In the present material the sacci are always constricted to a variable degree, frequently
deeply, in three places. The more deeply constricted examples are very similar to C.
trilobatus, particularly as Balme described the bladders of this species to occasionally
coalesce to form one trilobate bladder. The central body, however, is always scabrate
to subgranular rather than rugulate, the latter ornament being characteristic of C.
trilobatus.

596

PALAEONTOLOGY, VOLUME 12

C. dampieri is here emended to include forms in which the bladder is constricted to a
variable degree at the apices of the central body but never sufficiently deeply to clearly
delimit three bladders as are found in C. trilobatus. It is distinguished from C. trilobatus
by this character, by the scabrate to subgranular rather than rugulate central body, by
the narrow attachment areas of the bladders to the central body, and by the restriction
of the rugulate folds to this area.

Thus emended it embraces forms variously attributed to or compared with C.
dampieri (Balme) and C. trilobatus (Balme) by Lantz (19586) and Hughes and Couper
(1958). Some of these trilobed, but not trisaccate forms have a clear rugulate central
body (e.g. Lantz 1958, pi. 4, fig. 40) and on this character are best left in C. trilobatus. In
his original description of C. dampieri , Balme noted that ‘some specimens show vestigial
triradiate markings’ although these were not illustrated. Saccate grains with trilete scars
of various types and development have been illustrated by Hughes and Couper (1958)
and Lantz (19586) and attributed to Balme’s species of Zonalapollenites. It seems advis-
able to remove these from Callialasporites dampieri ( Balme) and C. trilobatus ( Balme) to
genera of the Triletisacciti since some of the trilete marks illustrated by these authors
appear to be quite well developed and not vestigial.

All figures x750. explanation of plate 107

Figs. 1-4, 7, 10. Appendicisporites potomacensis Brenner. 1, 2, Median focus and distal surface respec-
tively, GN 427/1, 48.0 117.6, 3, Proximal surface, GN 265/1, 122.3 44.1. 4, Median focus, GN
265/1, 50.2 111.0. 7, Median focus, GN 265/1, 48.2 123.4. 10, Median focus, GN 138/2, 50.0 102.6.
Figs. 5, 6. Plicatella abaca (Burger) comb, nov., distal and proximal surfaces respectively, GN 145/2,
45.5 108.4.

Fig. 8. Duplexisporites problematicus (Couper) Playford and Dettmann, GN 109B/1, 44.2 128.6.

Figs. 9, 14. Trilobosporites apiverrucatus Couper. 9, Proximal surface, GN 338/1, 34.9 124.5. 14,
Proximal surface, GN 341/1, 116.8 25.8.

Figs. 11, 13. Foraniinisporis wonthaggiensis (Cookson and Dettmann) Dettmann. 11, GN 428/1, 58.3

110.7. 13, GN 428/1, 44.6 114.0.

Fig. 12. Contignisporites dorsostriatas (Bolchovitina) Dettmann, GN 163/2, 32.4 119.1.

Fig. 15. Coronatispora valdensis (Couper) Dettmann; Damaged specimen showing structure of distal
surface, GN 154/1, 27.2 127.8.

Figs. 16, 17. Gleicheniidites senonicus Ross. 16, GN 146/2, 37.4 124.4. 17, GN 147/2, 28.2 116.9.

EXPLANATION OF PLATE 108
All figures X 750 unless otherwise stated.

Figs. 1, 2. Coronatispora valdensis (Couper) Dettmann. 1, Proximal surface, GN 187/2, 47.4 125.3.
2, Distal surface, GN 152/1, 42.5 112.3.

Figs. 3, 4, 6. Densoisporites perinatus Couper. 3, Proximal surface, GN 262/1, 41.5 123.3. 4, Median
focus, GN 338/1, 52.8 111.2. 6, Median focus, GN 338/3, 50.4 127.9.

Fig. 5. Sestrosporites pseudoalveolatus (Couper) Dettmann, GN 257/1, 44.1 116.4.

Figs. 7, 8, 10, 11. Heliosporites sp., Median foci. 7, GN 338/2, 37.1 110.7. 8, GN 427/2, 34.9 112.6.

10, GN 338/2,44.9 117.4. 11, GN 262/1, 53.3 114.2.

Fig. 9. Aequitriradites spimdosas (Cookson and Dettmann) Cookson and Dettmann, GN 428/2, 50.5

124.7.

Fig. 12. Januasporites tumulosus sp. nov., holotype, distal surface; GN 421/1, 55.5 123.3; X 1,250.

Fig. 13. Couperisporites complexus (Couper) Pocock; distal surface, GN 265/1, 59.0 115.3.

Palaeontology, Vol. 12

PLATE 107

NORRIS, Late Jurassic and Purbeck miospores

Palaeontology, Vol. 12

PLATE 108

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

597

Callialasporites obrutus sp. nov.

Plate 1 10, figs. 6, 7

Holotype. Slide GN 148/1, 48.4 109.6. Sample 60-5-24 (grey, shaley clay), Upper Purbeck, Unio Beds,
Durlston Bay (from Bed 82 of Bristow 1857).

Diagnosis. Spores radiosymmetric, alete, with a distinct central body. Amb of central
body circular to oval, surrounded by a saccus imparting an irregularly circular outline
to the entire spore. Central body outline not clear, saccus not distinctly attached to it.
Saccus very thin, standing out from central body one-tenth to one-twelfth of the total
diameter, scabrate to sub-granular, carrying coarse and also fine radial folds on equa-
torial bladder, these passing into irregular rugulate folds on central body. Central body
granular to rugulate, finer sculpture elements at poles. Exine of central body ( ?endo-
exine) thin, about 0-25 p or less in thickness. Bladders (?ektexine) also thin, about
0-25 p thick.

Description. Bladders very narrow, usually about 5 p wide but varying 4-9 p. Bladder
folded coarsely and approximately radially. Central body usually indistinct.

Dimensions. Equatorial diameter (including sacci): 38-69 p (holotype 61 p). Central body diameter:
29-44 p (holotype 44 p).

Distribution. Kimmeridgian, Dorset. Lower, Middle, and Upper Purbeck, Dorset, Sussex, and Surrey.

Remarks. Callialasporites obrutus sp. nov. is distinguished from Zonalapollenites
segmentatus Balme by the possession of a very thin-walled central body.

Callialasporites sp.

Plate 1 10, figs. 4, 5

Description. Spores radiosymmetric, trilete, with a narrow equatorially attached saccus.
Laesurae long, sinuous, labiate, 1-2 p wide reaching beyond amb of central body and
occasionally reaching amb of the saccus when they fan out into folds. Amb of both
saccus and central body circular to rounded triangular. Saccus scabrate but thrown into
rugulate, rather than irregular folds where attached to central body. Saccus projects
evenly beyond amb of central body a distance equal to one-sixth to one-tenth of the total
diameter (4-8 p). Saccus almost unfolded or carrying rather coarse, irregular radial
rugulate folds up to 2 p wide and 4 p long.

Dimensions. Equatorial diameter (including sacci): 39-64 p.

Distribution. Portlandian, Dorset. Middle and Upper Purbeck, Dorset and Surrey.

Remarks. Apart from the prominent trilete mark, this species is similar to Calliala-
sporites obrutus sp. nov. in possessing an equatorial, radially folded bladder which is
attached to a rugulate central body.

Turma aletes
Infraturma psilonapiti

Genus inaperturopollenites (Pflug ex Thomson and Pflug 1953) Potonie 1958

Inaperturopollenites sp.

Plate 1 10, figs. 11,12

598

PALAEONTOLOGY, VOLUME 12

Description. Spores small, spheroidal or occasionally of irregular shape owing to folding,
inaperturate. Exine relatively thick and rigid, unfolded or carrying short arcuate folds.
Exine 0-25-1 p in thickness, usually about 0-75 p.

Dimensions. Diameter: 9-15 p.

Distribution. Kimmeridgian and Portlandian of Dorset and Sussex. Lower, Middle, and Upper Purbeck
of Dorset, Sussex, and Surrey.

Infraturma spinonapiti
Genus peltandripites Wodehouse 1933
Peltcindripites tener sp. nov.

Plate 110, figs. 18, 19

Holotype. Slide GN 316/1, 42.0 123.7. Sample WM 2024/2 (grey, calcareous shale), Middle Purbeck,
Warlingham borehole, Surrey.

Diagnosis. Spores radiosymmetric, inaperturate, spherical, folded, entirely covered in

All figures X 750.

EXPLANATION OF PLATE 109

Fig. 1. Couperisporites complexus (Couper) Pocock; distal surface, GN 338/1, 43.4 1 1 1.0.

Figs. 2 -4, 7. Januasporites tumulosus sp. nov. 2, 3, Holotype, median focus and distal surface respec-
tively; GN 421/1, 55.5 123.3. 4, Median focus, showing circular aperture; GN 345/1, 41.51 28.5.
7, Median focus, GN 138/2, 40.4 117.2.

Figs. 5, 6. Marattisporites scabratus Couper. 5, Polar view, GN 163/2, 51.2 111.5. 6, Equatorial view,
GN 147/1, 27.8 119.0.

Figs. 8-10. Vitreisporites pallidus (Reissinger) Potonie. 8, GN 138/2, 28.0 122.0. 9, GN 154/2, 45.7
121.7. 10, GN 153/2, 30.7 1 13.5.

Figs. 11, 12. Cerebropollenites mesozoicus (Couper) Nilsson. 11, GN 152/1, 38.1 117.9. 12, GN 152/1,
48.0 111.1.

Fig. 13. Abietineaepollenites minimus Couper, GN 265/2, 46.4 111.3.

Figs. 14, 15. Alisporites bilateralis Rouse. 14, GN 143/2, 42.6 120.8. 15, GN 259/2, 28.6 127.2.

Figs. 16, 17. Podocarpidites sp. cf. P. ellipticus Cookson. 16, GN 431/1, 118.8 39.3. 17, GN 421/1,
47.5 123.1.

Figs. 18, 19. Parvisaccites radiatus Couper. 18, Oblique polar view, GN 146/2, 37.0 124.3. 19, Oblique
equatorial view, GN 186/2, 49.5 112.6.

EXPLANATION OF PLATE 110

All figures X 750 unless otherwise stated.

Fig. 1. Parvisaccites radiatus Couper, equatorial view, GN 138/2, 50.8 128.2.

Figs. 2-8. Callialasporites spp. 2-3, C. dampieri (Balme) Sukh Dev. 2, 111 3—1 3C, 44.2 119.4, 3. GN
421/1, 46.6 119.3. 4-5, C. sp. 4, GN 152/1, 29.5 123.2. 5, GN 152/2, 36.1 121.8. 6-7, C. obrutus sp.
nov. 6. Holotype, GN 148/1, 48.4 109.6. 7, GN 431/1, 23.9 118.2. 8, C. sp. cf. C. trilobatus (Balme)
Sukh Dev, GN 255/2, 49.7 120.2.

Figs. 9-12. Inaperturopo/lenites spp. 9-10, I. dubius (Potonie and Venitz) Thomson and Pfiug. 9, GN
196/1,56.8 127.7. 10, GN 345/1, 35.9 108.6. 11-12, /. sp. 1 1, GN 272/1, 44.9 1 17.0; X 1,250. 12, GN
421/1, 39.1 120.4; X 1,250.

Figs. 13. Spheripollenites subgramdatus Couper, GN 482/1, 34.6 109.3.

Figs. 14-16. Undidatasporites araneus sp. nov. 14, 16, High and median foci respectively, GN 421/2,
43.2 126.8. 15, High focus, GN 421/2, 47.1 110.0.

Fig. 17. Araucariacites australis Cookson, GN 265/2, 46.6 1 19.8.

Fig. 18, 19. Peltandripites tener sp. nov. 18, Holotype, GN 316/1, 42.0 123.7; x 1,250. 19, Tetrad, GN
281/1, 46.4 115.0.

Palaeontology, Vol. 12

PLATE 109

z.TA.

K^‘~- -i s a J ■ s

iV*'

NORRIS, Late Jurassic and Purbeck miospores

Palaeontology, Vol. 12

PLATE 110

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

599

short, closely spaced, irregularly distributed spines. Exine very thin, usually less than
0-25 p in thickness.

Description. Grains always carry many arcuate and crescentic folds. Spines 0-75-1 p
long, usually about 0-25 p wide but occasionally up to 1 p wide, spaced irregularly
0-25-1 p apart. In some corroded specimens some of the spines are reduced to low
granules. Rarely the grains occur in tetrads. Sometimes the grains are ruptured in an
arcuate fashion. Optical section of exine indistinct, occasionally reaching about 0-25 p
thick but never thicker.

Dimensions. Maximum diameter: 33-55 p (holotype 50 p).

Distribution. Kimmeridgian and Portlandian, Dorset. Lower and Upper Purbeck, Dorset.

Remarks. This species is easy to confuse with some acritarchs with short processes, ft is
only distinguished with difficulty but is clearly a pollen grain on account of its occur-
rence in tetrads.

Peltrandripites tener is distinguished from Araucariacites australis Cookson by its
smaller size, much thinner exine, and clearly echinulate rather than granular ornament.

Infraturma reticulonapiti
Genus undulatasporites Leschik 1955
Undulatasporites araneus sp. nov.

Plate 110, figs. 14-16; Plate 111, figs. 2-10

Holotype. Slide GN 345/1, 25.3 107.2. Sample 60-19-3, (grey calcareous marl), Upper Purbeck, Upper
Cypris Clays and Shales, Bacon Hole (6 in. above Bed 1 of Arkell 1933).

Diagnosis. Spores radiosymmetric, alete. Amb circular to oval. Exine very thin and
ornamented with irregular rugulae coalescing into a very imperfect, irregular micro-
reticulum consisting of narrow muri of constant width, and elongated irregular
lumina with a radially elongated arrangement towards the periphery. Exine 0-25-1 p
thick, possibly tectate.

Description. Muri and rugulae 0-25-1 p wide, of constant width along their length, 0-25 p
or less up to 0-5 p high. Luminae 0-5-1 p wide and up to 3 p long, tortuous and boun-
ded by anastomosing muri which show both angular and rounded bends in their courses.
Spores sometimes show concentric folds close to the periphery.

Dimensions. Maximum equatorial diameter: 21-30 p (holotype 26 p).

Distribution. Upper Purbeck, Dorset.

Remarks. Undulatasporites araneus sp. nov. is distinguished from Undulatasporites
anguineus Leschik by the overall smaller size, thinner exine, and shorter rugulae.

Turma plicates
Subturma monocolpates

Genus cycadopites Wodehouse 1933 ex Wilson and Webster 1946
Cycadopites sp. cf. C. nitidus (Balme 1957) comb. nov.

Plate 111, figs. 11, 12

600

PALAEONTOLOGY, VOLUME 12

1957 Entylissa nitidus Balme, p. 30, pi. 6, figs. 78-80.

1962 Ginkgocycadophytus nitudus (Balme) de Jersey, p. 12.

Distribution. Kimmeridgian, and Portlandian, Dorset. Lower, Middle, and Upper Purbeck, Dorset and
Surrey.

Remarks. These grains have a slightly greater over-all size range (18-39 p long; 13-27 p
broad) than those described by Balme but also differ in their tectate and scabrate to
infrapunctate exine (rather than ‘smoothly or faintly granulate’). The exine, however,
has a granular appearance in some corroded specimens. Balme (1957) believed that
C. nitidus was derived from plants of cycadalian or bennettitalian affinities. The strati-
fication of the exine of the present grains is not in conflict with this. Pollen grains of the
modern genera Bowenia, Cycas, and Macrozamia belonging to the Cycadaceae and are
illustrated by Erdtman (1957, figs. 10, 17, 46). All have tectate exines with rod-like
elements supporting the tectum and closely resemble C. sp. cf. C. nitidus. The exines of
some of the modern forms are distinctly crassitegellate whereas the tectum of C. sp. cf.
C. nitidus is frequently seen to be just thinner than the gap between it and the endexine
but merging into a crassitegellate exine.

Turma poroses
Subturma monoporines
Genus exesipollenites Balme 1957
Exesipollenites seabrosus sp. nov.

Plate 111, figs. 20-2

Holotvpe. Slide GN 316/1, 45.5 109.5. Sample WM 2024/2 (grey calcareous shale), Purbeck Beds,
Warlingham borehole.

Diagnosis. Spores radiosymmetric, monoporate. Amb rounded triangular to circular.
Both proximal and distal faces rather flattened. Distal face with a circular thickening
around the centre with an over-all diameter about half the equatorial diameter. At the
distal pole at the centre of the thickening is a rather thinner, distinctly depressed circular

EXPLANATION OF PLATE 1 1 1

All figures X 1,250 unless otherwise stated.

Fig. 1. Peltandripites tener sp. nov., GN 281/1, 54.5 121.1.

Figs. 2-10. Undulatasporites araneus sp. nov. 2-3, High and median foci respectively, GN 341/1, 27.8
128.0. 4-5, Holotype, high and median foci respectively, GN 345/1, 25.3 107.2. 6-7, GN 345/1,
30.5 112.4. 8, High focus, GN 345/1, 107.3 25.9. 9, Median focus, GN 421/1, 56.1 127.6. 10, GN
345/1, 24.3 108.9.

Figs. 11, 12. Cvcadopites sp. cf. C. nitidus (Balme) comb. nov. 11, GN 338/3 49.4 119.8; X750. 12,
GN 196/1 43.0 109.3; X 750.

Figs. 13-16. Eucommiidites spp. 13-14, 16, E. troedssonii Erdtman, x 750. 13, GN 163/2, 40.3 109.9.
14, GN 163/2, 48.7 125.9. 16, GN 148/1, 38.1 117.2. 15, E. minor Groot and Penny, GN 345/1,
25.9 107.4; X750.

Fig. 17. Monosulcites sp. aff. M. minimus Cookson, GN 344/1, 39.2 128.6; X 750.

Fig. 18. Cvcadopites carpentieri (Delcourt and Sprumont) Singh, GN 184/1, 41.9 108.8; x750.

Fig. 19. Inaperturopollenites dubius (Potonie and Venitz) Thomson and Pflug, GN 153/2, 23.2 122.1;
x 750.

Figs. 20-22. Exesipollenites seabrosus sp. nov. 20, GN 316/1, 109.3 45.3; X 750. 21-2, Holotype, distal
surface, X 750 and X 1,250 respectively; GN 316/1, 45.5 109.5.

Palaeontology , Vol. 12

PLATE 111

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

601

pore. Exine tectate, microreticulate, consisting of inwardly projecting elements. Occa-
sional scattered, irregular granules may occur on both faces. Exine just under 1 p in
thickness.

Description. Distal thickening distinct but not sharply delimited at its periphery, 11-
17 p in diameter (just greater or just less than half the equatorial diameter). Distal pore
at centre of distal thickening circular, depressed 1-5-3 p, 1-8 p in diameter (one-
twelfth to two-thirds the diameter of the distal thickening).

Microreticulate ornament developed over entire surface of grain, including the distal
pore and thickening, occasionally indistinct or scabrate. When well developed the muri
of the microreticulum are about 0-25 p wide and the lumina are about 0-25 p in diameter.
This tectate ornament may be accompanied by scattered groups of irregular granules up
to 1 p in diameter developed on the outer surface of the tectum. The tectate nature of the
exine is usually quite clear even though delicately marked. The inwardly projecting
elements of the ektexine are usually only seen as faint radial structures between the
tectum and endexine. The radial elements may become so faint as to simulate a simply
cavate exine.

Exine 0-25-1 p in total thickness. Endexine 0-25 p or less in thickness, occasionally not
clearly distinct from the ektexine.

Folding seldom occurs owing to the flattened nature of the spore but the amb is
occasionally irregularly crumpled.

Dimensions. Equatorial diameter: 23-36 p (holotype 36 p). Polar diameter: 4-6 p (holotype 6 p).

Distribution. Kimmeridgian and Portlandian of Dorset and Sussex. Lower, Middle, and Upper Purbeck
of Dorset, Sussex, and Surrey.

Remarks. Exesipollenites scabrosus is distinguished from ExesipoIIenites tumulus Balme
by the tectate nature of the thinner exine and by the microreticulate ornament. No
spheroidal forms with a triangular polar thickening which Balme ( 1 957) found associated
with E. tumulus were found associated with E. scabrosus. The Lower Jurassic assem-
blage in which E. tumulus occurred also contained abundant Classopo/lis (Balme 1957,
p. 41) as do the present samples in which E. scabrosus occurs.

MIOSPORES INCERTAE SEDIS

Genus sigmopollis Hedlund 1965
Sigmopollis ca/losus sp. nov.

Plate 1 13, figs. 9-12

Holotype. Slide GN 383/1, 48.7 108.3. Sample 60-19-22 (buff calcareous sandstone), Middle Purbeck
Corbula Beds, Bacon Hole (from the base of Bed 60 of Bristow 1857).

Diagnosis. Spores spheroidal. One face carries a distinct, markedly sinuous split in the
exine. Entire surface of grain ornamented with low bifurcating rugulae which do not
project at the amb. Exine about 1 p thick.

Description. Grains seldom folded and consequently more or less perfectly spherical
owing to the relatively thick and rigid exine.

Splitting is almost invariably developed and often takes an inverted S-shaped course

r r

C 6940

602

PALAEONTOLOGY, VOLUME 12

across the entire face. It does not appear to be associated with a thinning of the exine and
in optical section is seen to be a simple break perpendicular to the surface of the grain.

Rugulae 1-2 p long, up to 0-25 p wide, branching and occasionally partly forming an
imperfect microreticulum, usually not projecting at the amb but in a few specimens
giving the outline a slightly irregular appearance. Exine 0-75-1-25 p thick.

Dimensions. Diameter: 11-14 p (holotype 14 p).

Distribution. Middle and Upper Purbeck, Dorset, Sussex, and Surrey.

Remarks. SigmopolUs callosus is distinguished from Spheripollenites subgranulatus
Couper by the much thicker, more rigid exine, by the distinct rugulae which do not
project at the amb, and by the distinct sinuous rupture occurring on one face.

The split in the exine is considered to be significant (possibly connected with germina-
tion) rather than fortuitous owing to its constant and similar development in most
grains examined. Since it is not associated with any thinning of the exine it cannot be
considered a colpus. It is similar to a monolete mark but is not associated with a differ-
ential raising of the surface of the exine and appears to transgress the entire exine thick-
ness from the inner to the outer surface. Probably it is best considered an alete grain.

STRATIGRAPHIC PALYNOLOGY

All the samples examined from the Purbeck Beds and Upper Jurassic with few ex-
ceptions are dominated either by ClassopoUis torosus or Inaperturopollenites dubius. These
two species together usually constitute at least 70% and occasionally more than 90% of
the total spore-pollen flora. The relative abundance of these two species may have
palaeoecologic significance. They are of little use, however, for delimiting assemblages

EXPLANATION OF PLATE 112
All figures x 1,250 unless otherwise stated.

Figs. 1-5,8-16. ClassopoUis spp. 1-5, C. torosus (Reissinger) Balme. 1, Tetrad, GN 164/1, 43.9 128.5;
X 750. 2-3, Median and high foci respectively showing grain with endexine separated from ektexine;
GN 414/5, 49.5 125.2. 4, Equatorial view, GN 97C/1, 43.1 107.8; X 750. 5, polar view, GN 316/3,
37.6 128.0; x 750. 8-13, C. echinatus Burger. 8, Polar view, GN 316/1, 41.9 129.1. 9, Oblique polar
view, GN 316/1, 58.9 124.6. 10, Polar view, GN 316/1, 37.0 124.3. 11, Equatorial view, GN 203/1,

31.2 128.7. 12-13, Equatorial view, median and high foci respectively; GN 97C/1, 53.8 108.5. 14-16,
C. hammenii Burger. 14, Equatorial view, GN 316/1, 29.2 125.7. 15, Equatorial view, GN 162/2,
41.9 128.7. 16. Oblique equatorial view, GN 97C/2, 47.1 126.7.

Figs. 6, 7. Perinopollenites elatoides Couper. 6, GN 259/2, 50.8 1 18.1 ; X 750. 7, GN 190/3, 42.5 108.5;
X 750.

EXPLANATION OF PLATE 113
All figures X 1,250 unless otherwise stated.

Figs. 1-4. ClassopoUis hammenii Burger. 1, 2, Equatorial view, low and median foci respectively; GN
316/1, 32.0 128.5. 3, Polar view, GN 154/1, 52.7 120.3. 4, Oblique polar view, GN 316/1, 40.9 129.5.
Figs. 5-8, 13. Schizosporis spp. 5, 8, S. reticulatus Cookson and Dettmann. 5, GN 260/1, 43.3 116.3;
X 350. 8, GN 259/1, 53.7 127.9; X 350. 6, 13, S. spriggi Cookson and Dettmann. 6, GN 265/1,

46.2 1 16.5; x 350. 13, GN 163/1, 51.6 1 17.6; X 350. 7, S. parvus Cookson and Dettmann, GN 154/1,
47.8 125.6; X750.

Figs. 9-12. SigmopolUs callosus sp. nov. 9-11, Holotype, low, median and high foci respectively, GN
383/1, 48.7 108.3. 12, High focus, GN 383/1, 37.5 108.7.

Palaeontology, Vol. 12

PLATE 112

NORRIS, Late Jurassic and Purbeck miospores

Palaeontology, Vol. 12

PLATE 113

NORRIS, Late Jurassic and Purbeck miospores

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND 603

for stratigraphic purposes. On the other hand, the constituent species of the remaining
fraction of the miospore samples are very diverse and may be used to define spore-pollen
suites which appear to have stratigraphic significance. Suites have been recognized by
study of samples collected in sequences. They delimit broadly similar spore-pollen
samples, presumably derived from the same vegetational unit or units. They are de-
scribed on the basis of their persistent species (those present in about 50% or more of the
constituent samples in a suite) and spasmodic species (those occurring in less than half
of the samples on which the suite is recognized). The composition of the suites is shown
in text-fig. 3.

Marine Upper Jurassic Assemblages

The Upper Kimmeridgian and Portlandian of the Dorset coast and the Portland
Sand of Sussex have similar miospore assemblages which are included in one miospore
suite (Suite A). The basal Purbeck of Durlston Bay is also marine or brackish and is
characterized by the same Suite.

Suite A is restricted in composition, most assemblages consisting of 10-15 species
(listed in text-fig. 3). None of the species show restricted ranges within the intervals
examined. All species occur in the overlying Purbeck Beds. The extension of their
ranges into lower strata is not known in detail, owing to lack of adequate studies in the
lower horizons of the Kimmeridgian, but more than half the species comprising Suite A
have been reported from the Lower, Middle and Upper Jurassic of Britain by Couper
(1958), Lantz (19586), and Wall (1965).

The restricted nature of assemblages constituting Suite A is probably the result of the
combination of several factors. Both the Upper Kimmeridgian and Portlandian com-
prise marine sediments deposited in an offshore environment. Considerable sedimentary
sorting of the miospores by both wind and water is likely and may have resulted in only
a fraction of the total available spore-pollen population from the adjacent land areas
reaching the depositional site. This relative impoverishment may have been accentuated
by the extremely limited flora that occupied coastal sites at that time as evidenced by the
overlying Lower Purbeck assemblages.

At West Weare Cliff (Isle of Portland) Suite A occurs up to and including the Basal
Shell Bed. Samples above this bed from the Cherty Series and Freestone Series did not
yield palynologic assemblages. Suite A occurs in Portlandian samples from the West
Weare Sandstone, Upper Black Nore Beds, Black Nore Sandstone, and Lower Black
Nore Beds. Samples from the Exogyra Bed, the lower part of the West Weare Sandstone
and upper part of the Upper Black Nore Beds were not available. At Tar Rocks below
West Weare Cliff, Suite A characterizes the Kimmeridgian pallasioides zone to at least
135 ft. below the lowest row of nodules in the Black Nore Sandstone.

At Hounstout, Suite A characterizes similar horizons in the Lower Portlandian and
Upper Kimmeridgian. The highest occurrence of Suite A collected at this locality was
immediately below the Lower Parallel Band of the St. Albans Head Marls. Suite A
characterizes the Portlandian and Upper Kimmeridgian below this, comprising the
remainder of the St. Albans Head Marl, Emmit Hill Marl, Hounstout Marl, Hounstout
Clay, Rhynchonella Marl, Lingula Shale, Rotunda Clays and Nodules, and the highest
20 ft. of the Crushed Ammonoid Shales, below which samples were not collected.

At Durlston Bay, the only Purbeck assemblage obtained below the base of the Marls

6u4

PALAEONTOLOGY, VOLUME 12

text-fig. 3. Composition of microfloral suites showing persistent species (unbroken lines) and spasmodic
species (broken lines). N.B. Range in entry 16 is correct, and not that in entry 41.

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

605

with Gypsum and Insect Beds belongs to Suite A. This assemblage which was obtained
from a horizon 3 ft. below the top of the Broken Beds, contains dinoflagellate cysts
(Norris 1965 b) suggesting marine conditions of deposition, as suggested independently
by Brown (1964) using petrologic criteria.

Marine plankton. The spore-pollen assemblages of Suite A are accompanied by a large
proportion of dinoflagellate cysts and acritarchs; Norris (1965) has described a few of
these species. The relative numerical abundance of microplankton in the marine Upper
Jurassic (expressed as a ratio — number of microplanktonic forms/number of miospores)
varies from 0-2 to TO. The relative diversity of marine Upper Jurassic microplankton
species compared with miospore species (expressed as a ratio) varies between 0-6 and
3-0. In contrast the saline horizons in the Purbeck Beds containing microplankton have
relative numerical abundances of microplankton that rarely exceed 0-01 and relative
diversities of microplankton that never exceed 0-2 and frequently are less than 0T.

Purbeck Assemblages

The Purbeck miospore assemblages of Dorset, Sussex, and Surrey can be grouped into
two suites of distinctly different composition.

Suite B. This Suite (see text-fig. 3) is similar to Suite A in terms of most of the commoner
constituents; however, Classopollis echinatus and Inaperturopollenites sp. occur less
frequently, and Podocarpidites sp. cf. P. ellipticus, Araueariacites australis and Perino-
pollenites elatoides more frequently. Eucommiidites minor occurs in Suites A and C but
is unknown in Suite B. Suite B is distinguished from Suite A by the presence of the
following rarely occurring species:

Acanthotriletes varispinosus, Converrucosisporites variverrucatus, Leptolepidites psarosus , Micro-
reticelatisporites diatretes, Cicatricosisporites purbeckensis , PlicatelUi abaca, Couperisporites complexus,
Cycadopites carpentieri, Schizosporis spriggi , Parvisaccites radiates.

On the Dorset Coast, Suite B characterizes most of the Lower Purbeck and the lower
part of the Middle Purbeck. At Durlston Bay, Suite B occurs up to Bristow’s (1857)
Bed 53 in the Upper Building Stone, 17 ft. below the base of the Corbula Beds. At
Bacon Hole Suite B occurs up to the top of the Marly Freshwater Beds (Bristow’s Bed
42). Its lowest occurrence is not clearly defined at Bacon Hole where palynologic as-
semblages are rare in the Lower Purbeck. At Durlston Bay, however, Suite B definitely
occurs down to the base of the Marls with Gypsum and Insect Beds. Suite A occurs
almost to the top of the Broken Beds.

At Mountfield, Suite B characterizes the 4 gypsum seams at the bottom of the Purbeck
sequence, the highest sample containing Suite B assemblages occurring at the top of No.
1 seam.

In the Warlingham borehole, Suite B does not occur above 2,022 ft. but the section
was not examined below 2,027 ft. and so its lowest occurrence was not determined.
Suite C. This suite is similar to Suite B in terms of most of the persistent species (text-
fig. 3) but contains these additional persistent species (some of which, however, occur
rarely in other suites) :

Cyathidites minor , Klnkisporites pseudoreticidatus, Cicatricosisporites australiensis, Cicatricosi-
sporites purbeckensis, Plicatella abaca, Coronatispora valdensis, Parvisaccites radiates.

606

PALAEONTOLOGY, VOLUME 12

Suite C contains all the rare species that characterize Suite B but also contains the 32
additional spasmodic species listed in text-fig. 3.

Suite C occurs in the upper part of the Middle Purbeck and the Upper Purbeck of the
Dorset Coast. At Durlston Bay, this Suite ranges from a horizon 15 ft. below the top of
the Upper Building Stone (Bristow's Bed 55) to the highest sample examined, a bed of
limestone at the top of the Marble Beds and Ostracod Shales. The Viviparus Clays are
no longer exposed at this locality. At Bacon Hole the lowest occurrence of Suite C is
from near the base of the Corbula Beds (from the bottom of Bristow’s Bed 60). The
interval from the base of the Corbula Beds to the highest occurrence of Suite B close to
the lower middle Purbeck junction is barren of miospores. Consequently the exact
delimitation of Suites B and C at this locality cannot be determined accurately. Suite C
occurs up to the highest sample examined from Bacon Hole, a silty clay in Wealden
arenaceous facies about 8 ft. above the top of the Upper Purbeck Paludina Clays. The
latter is delimited from the basal Wealden by the incoming of sandstone.

At Lulworth Cove, Suite C occurs in the Paludina Clays but its lowest occurrence has
not been determined at this locality because of the lack of assemblages. Suite C is not
known at Worbarrow Bay, also because of lack of palynomorph assemblages in the
Upper Middle and Upper Purbeck.

At Mountfield, Suite C occurs from 13 ft. above the top of No. 1 gypsum seam
throughout the remaining ‘Middle’ and ‘Upper’ Purbeck to the highest sample examined
in the lower half of the Grey Limestone Series exposed in the River Line south-west of
the gypsum mine (Howitt 1964, p. 86).

In the Warlingham borehole. Suite C occupies the interval from 2,004 ft. to the highest
sample examined at 1 ,900 ft.

DISCUSSION OF ASSEMBLAGES AND SUITES

The three suites represent a progressive diversification of the plant microfossil assem-
blages (text-fig. 3). Species of Suite A are drawn from a total of 32 species. Most assem-
blages constituting Suite A are composed of 10-15 species. Suite B is represented by a
total of 42 species, although each assemblage in this suite usually consists of less than 10
species with occasionally up to 22 species present. Suite C is the most diverse, being
represented by a total of 73 species; most assemblages of Suite C contain 15-30 species
although this number can be (rarely) as low as 4 or as high as 42.

Each successive suite is characterized by the entrances of new forms. There are no
well-marked extinctions although local ranges may be restricted in some sections. Most
of the characteristic new forms of each suite are rare types. This leads to difficulties in
assigning some samples to a suite, but the use of sequential samples has allowed a more
meaningful evaluation of the absence of rare types in a sample.

This progressive diversification of the assemblages may be the result of evolution,
sedimentation, local phytogeography, or changing environments. The distribution and
ranges of characteristic species within each Suite is not constant from section to section,
which suggests that factors other than evolution and extinction are important in deter-
mining individual ranges. For example, Pi/osisporites delicatulus is characteristic of
Suite C in Dorset but is unknown in Sussex and Surrey. Thus local palaeoecologic
control may be severe and does not allow the use of individual miospores for minutely

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

607

detailed stratigraphic correlation, although the use of miospores for general correlative
purposes is possible as set out below.

There is no apparent relationship between the miospore Suites and the numerical
dominants in each section. Either Classopollis torosus or Inaperturopollenites dubius may
dominate the assemblages constituting Suite A. At Durlston Bay and in the Warlingham
borehole Suite B is characterized by relative abundances of Classopollis torosus greater
than 50%. At Bacon Hole and Mountfield, C. torosus dominates the lower assemblages
of Suite B but I. dubius becomes important in the higher assemblages of this Suite.
Suite C is characterized by an abundance of /. dubius at Durlston Bay and in the War-
lingham borehole. At Bacon Hole and Mountfield either I. dubius or C. torosus may
dominate the Suite C assemblages. Many assemblages in Suite B and C are characterized
by the freshwater chlorophycean alga, Botryococcus, presumably reflecting the domin-
antly non-marine conditions of deposition.

Botanical considerations. It is possible to assign many dispersed spore and pollen species
from the Jurassic and Cretaceous to natural plant orders and families (Potonie 1962,
Brenner 1963, Dettmann 1963, Pocock 1962, Singh 1964, Norris 1967). Species con-
stituting Suites A, B, and C are distributed amongst natural plant taxa at the ordinal or
higher levels as follows:

bryophyta: Slereisporites antiquasporites, Foraminisporis wonthaggiensis , Aequitriradites spinulosus,
Couperisporites complexus.

pteridophyta: Inaperturopollenites dubius (pars). Lycopodiales: Acanthotriletes varispinosus, Lyco-
podiacidites cerniidites, Lycopodiumsporites austroc/avatidites, Foveosporites canalis, Sestrosporites
pseudoalveolatus , Densoisporites perinatus. Filicales: Cyathidites minor, C. australis, Osmundacidites
wellmanii. Bacidatisporites comanumensis, Converrucosisporites variverrucatus , Leptolepidites psarosus,
Rubinella major, Pilosisporites trichopapillosus, Cicatricosisporites australiensis. C. purbeckensis, C.
angicanaiis, C. brevilaesuratus, Klukisporites pseudoreticulatus, Trilobosporites bernissartensis, T.
obsitus, T. domitus, Appendicisporites potomacensis, Plicatella abaca, Gleicheniidites senonicus, Maratti-
sporites scabratus. Pteridophyta incertae sedis: Deltoidospora rafaeli , D. psilostoma, Concavisporites
juriensis, Divisisporites sp. cf. D. euskirchenensis, Pilosisporites delicatidus, Reticidisporites semireticulatus,
Microreticidatisporites diatretus, Tripartina sp., Heliosporites sp., Contignisporites dorsostriatus,
Coronatispora valdensis, Januasporites tumulosus.

cycadophyta: Vitreisporites pallidas , Cycadopites sp. cf. C. nitidus, C. carpentieri.

coniferophyta. Coniferales: CerebropoUenites mesozoicus, Alisporites bilateralis, Abietineaepollenites
minimus, Podocarpidites sp. cf. P. ellipticus, Parvisaccites radiatus, Callialasporites sp. cf. C. trilobatus,
C. dampieri, C. obrutus, C. sp., Inaperturopollenites dubius (pars), Araucariacites australis, Spheri-
pollenites subgranulatus, Perinopollenites elatoides. Coniferophyta incertae sedis: Eucommiidites
troedssonii, E. minor, Exesipollenites scabrosus, Classopollis torosus, C. echinatus, C. hammenii.

spores and pollen incertae sedis: Inaperturopollenites sp., Peltandripites tener, Undulatasporites
araneus, Monosulcites sp. aff. M. minimus, Schizosporis reticulatis, S. spriggi, S. parvus, Sigmopollis
callosus.

From the above list it is evident that the assemblages spanning the Jurassic-Cretaceous
boundary in southern England as a whole represent principally pteridophyte-gymno-
sperm vegetation. Lycopsids are less important than filicalean elements but some of the
pteridophyte spores of uncertain affinities may belong to this group. Bryophytes and
cycads are both relatively unimportant groups.

Ascending the sequence from the Upper Kimmeridgian to the Upper Purbeck there is
a progressive diversification of the spore-pollen assemblages. Suite A consists principally

608

PALAEONTOLOGY, VOLUME 12

of coniferalean elements with a few filicalean, lycopsid, and other pteridophyte species.
In Suite B conifers remain important but pteridophytes begin to diversify. Bryophytes
appear in Suite B as rare elements. In Suite C conifers remain the most diverse persistent
elements but pteridophytes are the most diverse of the spasmodic species. Bryophytes
are commoner here than in Suite B but are still rare. Cycadophytes are rare elements in
all the suites. The numerical dominants in all the suites, Classopollis torosus and Inaper-
turopollenites dubius, are both coniferalean species but some of the inaperturate grains
could be derived from equisetalean plants (Batten 1968).

CORRELATION AND AGE OF THE PURBECK BEDS

Correlation

It is difficult to select ‘index’ forms or key species (Couper 1958) but the suites recog-
nized on the basis of several species are presumably time concordant.

Correlation of the Purbeck Beds and equivalent strata in southern England on the
basis of palynologic suites is shown in text-fig. 4. At Durlston Bay, Suite B is present
from the top of the Broken Beds (samples from the Caps and Dirt Beds below did not
yield palynomorphs) to the upper part of the Upper Building Stones. Suite C occupies
the remainder of the Upper Building Stone to the top of the Marble Beds and Ostracod
Shales. In Bacon Hole the boundary between Suites B and C occurs at a similar strati-
graphic position, i.e. at the approximate level of the Scallop Beds (of Bristow’s 1857
terminology). Due to the westward attenuation of the strata, however, the B/C boundary
is approximately 15 ft. above the Cinder Bed at Bacon Hole whereas the equivalent
interval at Durlston is almost double that thickness.

Samples from the ma jor part of the Purbeck Beds at Worbarrow Bay and at Lulworth
Cove were too sparse (perhaps due to the large amounts of limestone in the sections) to
locate the boundary between Suites B and C. At Mountfield, however, this boundary is
well marked in the subsurface section of the Purbeck Beds just above the No. 1 gypsum
seam approximately 50-60 ft. above the base of the section. This contrasts with the
position of the B/C boundary approximately 260 ft. above the base at Durlston Bay.
Thus the four gypsum seams and intercalated shales at the base of the Mountfield
Purbeck would appear to be either a condensed sequence equivalent to the Lower and
much of the Middle Purbeck of Dorset (some 260 ft.), or there is a hiatus between the
underlying Portlandian and the lowest Purbeck at Mountfield which seems more likely
for the following reasons.

Although the Purbeck of Dorset is apparently conformable with the underlying Upper
Portlandian, in other areas there is some evidence to suggest that the Upper Portlandian
is absent. In the Henfield borehole of Sussex, 2 ft. below the base of the Purbeck Beds,
faunas occur indicating the Lower Portlandian albani zone (Taitt and Kent 1958). In the
Portsdown borehole the Purbeck Beds rest on Portlandian equivalent to the top of the
Portland Sand (Lower Portlandian) in Dorset. At Mountfield the arenaceous sediments
immediately underlying the Purbeck Beds have yielded dinoflagellates characteristic of
the top part of the Lower Portlandian gorei zone in Dorset (Norris 1963). Falcon and
Kent (1960, p. 13) presented additional evidence from boreholes in southern England and
stated that there is ‘no evidence of the occurrence of Upper Portland anywhere in the
Wealden area, there being a non-sequence at the base of the Purbeck’. Taitt and Kent

LULWORTH BACON WORBARROW DURLSTON MOUNTFIELD WARL1NGHAM

COVE HOLE BOREHOLE

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

609

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610

PALAEONTOLOGY, VOLUME 12

(1958) also ascribed this break to a non-sequence due to a regional movement causing a
cessation of deposition since there are no signs of erosion and the break is of a uniform
nature. The occurrence of Lower Portlandian pebbles within the Lower Purbeck of
Dorset (Woodward 1895), the erosion of the Portland Beds prior to deposition of the
Purbeck Beds in the Aylesbury district (Woodward 1895), and the presence of derived
Kimmeridgian and Portlandian pelecypods from the basal Purbeck Beds of Swindon
similar to forms from the south of England and the Boulonnais (Arkell 1941) indicate
the possible existence of a pre-Purbeck unconformity.

The correlation of the Purbeck at Mountfield is particularly interesting whatever the
reason for the absence of marine Upper Portlandian at this locality. Details of Purbeck
ostracod faunas are not available. Howitt (1964) used lithologic criteria for correlation
of the Purbeck sequence of the Mountfield inlier. He assigned this relatively thick
sequence to the conventional tripartite subdivision: the ‘Lower’ Purbeck consists
largely of gypsum and limestone; the ‘Middle’ Purbeck consists of shale, some marine
limestone and arenaceous horizons; the ‘Upper’ Purbeck contains freshwater limestones,
shales, and calcareous sandstones. A different correlation is suggested on the basis of
miospores (text-fig. 4). Suite B characterizes the gypsiferous series at the base plus a few
feet of overlying shale as detailed in the previous section. Suite C occupies the remaining
Mountfield Purbeck to the middle of the Greys Limestone Series. This suggests that the
entire gypsiferous series at Mountfield is equivalent to the Middle Purbeck rather than
to the Lower Purbeck of Dorset as suggested by Howitt (1964). It is difficult to deter-
mine how much of the Durlston Suite B is represented at Mountfield. There appear to be
no breaks in sedimentation within Suite B strata which suggests in view of their thin
development that only the upper part of Suite B is represented, i.e. equivalent to part of
the lower Middle Purbeck. The correlation of the higher assemblages of Suite C is
difficult. Cicatricosisporites angicatialis , a distinctive species of limited stratigraphic
range, is restricted to the upper part of the Upper Purbeck in Dorset. At Mountfield,
C. angicanalis occurs in one sample of the Greys Limestone suggesting a correlation
with the upper part of the Upper Purbeck of Dorset. Thus if this correlation is correct,
sedimentation of the upper part of the Middle Purbeck (above the Cinder Bed) and the
Upper Purbeck proceeded at less than half the rate at Durlston Bay (183 ft.) compared
with the equivalent interval at Mountfield (396 ft.).

Age

At the type locality on the Dorset Coast the Purbeck Beds overlie with apparent
conformity the uppermost Portlandian. The Upper Portlandian has been correlated with
the Middle Tithonian of the continent by Arkell (1956) on the basis of the contained
ammonite faunas. Presumably the Upper Tithonian is represented by at least part of the
Purbeck sequence in Dorset. The top of the Purbeck Beds is delimited by the incoming
of coarse, clastic arenaceous and carbonaceous sediments of Wealden facies which have
not been precisely dated but in the past have been generally assumed to be basal Cre-
taceous (Allen 1955).

Ostracods have been used to correlate the Purbeck Beds of southern England (Ander-
son 1958) and similar European formations (Anderson 1962, Arkell 1956, Bartenstein
and Burri 1955). Age determinations of the Purbeck Beds, however, rest principally on
occurrences of ammonites (Arkell 1956). Arkell ’s arguments for a Late Tithonian age

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

611

for the Purbeck Beds have been reviewed and revised by Donze (1958) who suggested
that the ammonites indicate either a Late Tithonian or Berriasian age for the French
Purbeck.

The Jura Purbecks have been correlated with the Lower and Middle Purbeck by
F. W. Anderson using ostracods (Arkell 1956). Their Berriasian and possibly Late
Tithonian age is not inconsistent with other stratigraphic and palaeontologic evidence
from underlying and overlying deposits in this region. Arkell (1956) has noted that
undated limestones underlie the ammonite-bearing and other sequences of the French
Purbecks. Only the lower part of the thick limestones underlying the Jura Purbecks
contain ammonites and these indicate the Kimmeridgian gravesiana zone. The higher
part of this limestone sequence is customarily referred to as ‘Portlandian’ but diagnostic
fossils are lacking. The limestones overlying the French Purbecks are assigned to the
Berriasian with little or no palaeontologic evidence but are overlain by undoubted
Valanginian. The limestones and marls overlying the Jura Purbecks are considered to be
Berriasian on insecure evidence (Bartenstein and Burri 1955) and the one diagnostic
ammonite has been compared to a Valanginian form (Arkell 1956). The remaining
fauna is not diagnostic.

Similar correlations to the above for the Swiss and French Purbecks have been
suggested by other workers using fossils other than ammonites. Bartenstein and Burri
(1955) correlated the Swiss Purbecks with the Serpulite and Wealden 1 of NW. Germany
and with the higher Lower Purbeck and lower Middle Purbeck of England using ostra-
cods and charophytes. Donze (1958) correlated the French Purbecks with a regression
in the Berriasian of the ‘fosse vocontienne’ (indicated by microfaunal and algal
changes) and with the Lower Purbeck and part of the Middle Purbeck of Dorset (using
ostracods). Palynologic correlations differ slightly: these suggest a correlation of the
Serpulite and Wealden 1 of Germany and Holland with the upper Middle Purbeck and
part at least of the Upper Purbeck. Thus the Berriasian regressive facies in the West
Tethys may correlate with the Lower Purbeck and lower Middle Purbeck or with the
upper Middle Purbeck and Upper Purbeck. Bartenstein (1959, 1962) extended the
Berriasian correlation of the French Purbecks to the Jura Purbecks. He confirmed that
probably only the lowest part of the Swiss ‘Berriasian’ overlying the Jura Purbecks is to
be correlated with the top of the type Berriasian, most of the remainder being of
Valanginian age.

Thus on the evidence from ammonites and using ostracods to correlate to the west
Tethyan region, the Purbeck Beds in Dorset appear to span the Jurassic-Cretaceous
boundary occupying the interval Late Tithonian-Berriasian (text-fig. 5).

Assignment of the upper part of the Purbeck Beds to the Lower Cretaceous is also
suggested by palaeobotanical arguments. Hughes (1958) studied megaspores from the
English Wealden and compared them with those from Dutch and German sequences.
The lowest Wealden formation in the type area is the Fairlight Clay of the Sussex Coast.
The Fairlight Clay is believed to immediately overlie or be a partial facies equivalent of
some of the Purbeck (Allen 1960 a , b) but the base of this formation is not exposed and
so exact relationships cannot be confirmed. Overlying the Fairlight Clay is the Ashdown
Sand which Hughes (1958) correlated with the Lower Valanginian of the Netherlands
from the occurrence in both of Thomsonia megaspore floras. Hughes also noted that
Thomsonia floras are known from the German Wealden 4 and 5. The beds below the

612

PALAEONTOLOGY, VOLUME 12

Thomsonia horizon of the Lower Valanginian of the Netherlands contain Pyrobolospora
pyriformis (Dijkstra) Hughes which is known only in the Fairlight Clay in England.
These lower beds in the Netherlands are probably Berriasian or partly Lower Valangin-
ian, suggesting a similar age for the Fairlight Clay. The Upper Purbeck is a facies
equivalent of the Fairlight Clay, as suggested by Hughes and Moody-Stuart’s (1969)
correlation of the Lower Fairlight Clay with the upper Middle and Upper Purbeck using
a new palynologic method. This indicates that the upper Middle and Upper Purbeck
may also be Lower Cretaceous ( ?Berriasian) in age. The uppermost Purbeck Beds at
Bacon Hole contain megaspores at some horizons (see Appendix 1 in Norris 1963). More
intensive study of these megaspores may help to resolve the point.

Jurassic-Cretaceous boundary in southern England

As reviewed above, the Purbeck Beds on the Dorset Coast appear to occupy the
Upper Tithonian-Berriasian interval and hence straddle the Jurassic-Cretaceous
boundary which is conventionally placed between the Tithonian and Berriasian stages
(Arkell 1956). These stages are based on ammonite faunas occurring in marine lime-
stones of south-east France. Correlation with non-marine and transitional strata is
difficult, being based on sparse ammonite-bearing marine intercalations together with
the use of non-marine fossils to extend these age determinations into other areas of non-
marine rocks. In southern England (Dorset) this has led to a situation in which it is
possible to say that the lower part of the Lower Purbeck is probably Upper Tithonian
and that the higher part of the Upper Purbeck is Berriasian (perhaps Upper Berriasian).
But it is not possible to place the Jurassic-Cretaceous boundary precisely. Casey (in
Howitt 1964, p. 109) has discussed some of the problems involved in locating the base of
the Cretaceous in England. He suggested tentatively that the Cinder Bed of Dorset
might be taken to mark the Jurassic-Cretaceous boundary. Casey considered this to be a
convenient procedure because he thought the Cinder Bed to be the most extensive of
several saline horizons in the Purbeck of England. Furthermore Casey pointed out that
the flooding of the Wessex Basin that led to deposition of the Cinder Bed and supposed
correlative horizons was perhaps linked with the Osterwald phase of Saxonian folding in
north Germany that immediately preceded deposition of the Serpulite (a possible
correlative of the Cinder Bed), and whose effects might be traced in the Spilsby Sea
deposits further north. If the Cinder Bed is taken to mark approximately the base of the
Cretaceous, the base of miospore Suite C occurs just above the Jurassic-Cretaceous
boundary (text-fig. 4). At Mountfield, the Jurassic-Cretaceous boundary on palyno-
logic evidence would be placed at about the top of the Main Gypsiferous Beds. Hence the
gypsiferous series at Mountfield appear to correlate at least in part with the Cinder Bed
of Dorset (text-fig. 4). This is considerably different from the correlations suggested
by Howitt (1964) who equated the gypsiferous series at Mountfield with the gypsum
developed in the middle of the Lower Purbeck at Durlston, more than 100 ft. below the
Cinder Bed. Howitt correlated the Cinder Bed of Dorset with a 5-ft. marine limestone
containing Ostrea distorta, 182 ft. above the top of the Main Gypsiferous Beds. This
5-ft. limestone in fact would correlate with a horizon in the Chief Beef Beds at Durlston,
assuming the palynologic correlation is correct and sedimentation was twice as rapid at
Mountfield compared with Durlston (see the preceding sections). Dinoflagellate cysts
and acritarchs occur in samples from the Chief Beef Beds of Durlston Bay (Norris 1963)

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND 613

suggesting marine influence at these horizons and supporting the above correlation.
Dinoflagellate cysts and acritarchs also occur in small quantities in the Main Gypsi-
ferous Beds of Mountfield but comparisons with acid-insoluble microfossil assemblages
from the Cinder Bed of Dorset is not possible because this limestone bed has proved to
be barren of palynomorphs. If the microfloral correlation is correct it suggests that the
entire Purbeck sequence in Sussex is Early Cretaceous in age (text-fig. 5), accepting
Casey’s selection of the Cinder Bed in Dorset as a marker horizon for the base of the
Cretaceous. More detailed work is required in the Mountfield area to refine the corre-
lations suggested here.

COMPARISON WITH OTHER JURASSIC AND CRETACEOUS

ASSEMBLAGES

Britain

Couper (1958) noted that the spore-pollen assemblages from the Kimmeridgian of
Brora (Scotland) are impoverished in both number of species and specimens and in
general are similar to Oxfordian assemblages from Yorkshire and to Middle Callovian
assemblages from Brora. Although details on only three Kimmeridgian assemblages,
one Oxfordian assemblage, and one Callovian assemblage are available, the data are
sufficient to make a general comparison. All these Upper Jurassic assemblages (the
Callovian is classed variably as either Middle or Upper Jurassic by different workers)
are characterized by the dominance of Abietineaepollenites minimus and Abietineae-
pollenites microalatus Couper. This contrasts sharply with Suite A in the Upper Kimmer-
idgian and Portlandian of Dorset in which Classopollis torosus and Inaperturopol/enites
dubius are dominants or co-dominants. Couper’s Callovian, Oxfordian, and Kimmer-
idgian assemblages are clearly distinguished from the Dorset Upper Jurassic assemblages
by the presence of Sestrosporites irregulatus (Couper) Dettmann, Cingulatisporites
dubius Couper, and Pteruchipollenites microsaccus Couper. These three species are
unknown in the Upper Kimmeridgian, Portlandian, or Purbeck Beds of southern
England. Of the approximately twenty remaining species in Couper’s assemblages, a
little more than half are pteridophyte spores, just less than half are conifer grains and
the rest are cycads or pteridosperms. All of these occur in younger strata but the
following are absent in Suite A: Cyathidites australis Couper, Dictyophyllidites harrisii
Couper, Densoisporites perinatus Couper, Duplexisporites problematicus (Couper)
Playford and Dettmann; the distance of the parent vegetation from the site of marine
deposition in Dorset may have been important in determining what miospores entered
the sediment. With the onset of estuarine and non-marine conditions in southern
England in the Upper Tithonian and Lower Cretaceous these species presumably were
able to reach the depositional sites more easily. Species present in Suite A which are not
known in Couper’s Middle or Upper Jurassic assemblage include Exesipollenites
scabrosus sp. nov., Inaperturopol/enites sp., Classopollis echinatus, Classopollis hammenii,
Rubinella major , Retieulisporites semireticulatus, Klukisporites pseudoreticulatus, Cieatri-
cosisporites australiensis, Coronatispora valdensis, Podocarpidites sp. cf. P. ellipticus ,
Callialasporites dampieri, Callialasporites obrutus, Callialasporites sp., Peltandripites
tener, Eucommiidites minor , Cycadopites sp. cf. C. nitidus.

Couper (1958) and Hughes (1958) summarized the main differences between the

614

PALAEONTOLOGY, VOLUME 12

text-fig. 5. Correlation chart of strata close to the Jurassic-Cretaceous boundary in north-west Europe.

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND 615

Wealden microflora and the Purbeck assemblages known at that time. Many of their
conclusions are invalidated by the present study. Of the six species stated to characterize
the Wealden, five are now known to occur in the Purbeck or Upper Jurassic (particu-
larly in Suite C but some also in Suites A and B), viz. Tri/obosporites apiverrucatus ,
Parvisciccites radiatus, Appendicisporites potomacensis (= A. tricornitatus of Couper 1958
in part), Cicatricosisporites brevilaesuratus, and Coronatispora valdensis. Concavi-
sporites pane tat us was reported by Couper (1958) and by Hughes (1958) to range down
to the base of the exposed Fairlight Clay and this species is not known in the Purbeck
Beds. Apart from this species, however, the Fairlight Clay assemblages are closely
similar to those of Suite C, as far as it is possible to judge from Couper’s descriptions.
The numerical dominants in the Fairlight Clay, however, are quite different: about
70% of the grains are filicalean spores and the remainder are AbietineaepoUenites , con-
trasting with the abundance of Inaperturopollenites dubius in the Upper Purbeck. The
possibility that the Fairlight Clay at least in part is a lateral facies equivalent of the upper
part of the Purbeck Beds should be considered in view of recent palynological corre-
lations suggested by Hughes and Moody-Stuart (1969). Numerical differences could be
due to the clastic facies of the Fairlight Clay compared with the finer grained lithologies
in the Purbeck (limestone and shale).

Germany and Holland

Comprehensive accounts of the palynology of strata close to the Jurassic-Cretaceous
boundary have been given by Doring (1965) for north-central Germany and by Burger
(1966) for the eastern Netherlands. Several difficulties, however, are involved in making
comparisons with other areas. Strata close to the British Jurassic-Cretaceous boundary
show relatively restricted spore-pollen assemblages. This is due either to distance from
the site of the vegetation in the case of the marine sediments or to the apparently re-
stricted and perhaps specialized nature of the coastal vegetation in the case of the
estuarine and non-marine sediments. The assemblages described from the continent
appear to be more diverse which in part may be a taxonomic effect due to the less
conservative approach to miospore variability adopted by those continental workers.
But much of this diversity appears to be a real reflection of richer parent vegetation.
Detailed comparisons of individual ranges of miospores between Britain and the
Dutch and German localities is not appropriate for stratigraphic purposes in this case
because palaeophytogeography or palaeoecology appear to have been important in
controlling miospore distribution. A general comparison of the continental palyno-
logic sequences with the palynologic suites recognized in southern England may yield
more meaningful results.

Burger (1966) studied two cored sections of uppermost Jurassic and lowermost
Cretaceous strata located on the western edge of the Lower Saxon Basin in the Twente
area of Holland. He recognized 9 pollen zones spanning the higher Upper Malm (con-
ventionally placed in the Tithonian, e.g. Arkell 1956) and the predominantly non-
marine ‘Wealden’ up to the next fully marine horizon in the Middle Valanginian.
Burger noted that his 9 pollen zones based on appearances and disappearances of
miospore species may have only local significance. Of the 88 species occurring in the
Dutch sequence, only 29 species are common to the Upper Jurassic-Purbeck sequence in
England. All but 6 of these species range from Upper Malm to Middle Valanginian. Of

616

PALAEONTOLOGY, VOLUME 12

the 6 species of restricted range, all are present in the Upper Malm but do not occur
higher than the lower part of the Middle Valanginian. Thus the following 29 species
occur from Upper Malm to the base of Middle Valanginian sediments in Holland and
also occur in the Upper Kimmeridgian-Purbeck of England:

Cyathidites minor, C. australis, Deltoidospora rafaeli*, Concavisporites juriensis** , Dictyophyllidites
equiexinus, Osmundacites wellmanii, Rubinella major, Lycopodiumsporites austroclavatidites, Coronati-
spora valdensis, Reticulisporites semireticulatus, Plicatella abaca*, Trilobosporites bernissartensis**, T.
apiverrucatus**, Gleicheniidites senonicus, Callialasporites dampieri, C. trilobatus**, Abietineaepollen-
ites minimus, Parvisaccites radiatus*, Vilreisporites pallidus, Cerebropollenites mesozoicus, Eucom-
miidites troedssonii, E. minor, Monosulcites minimus**, Araucariacites australis, Splieripollenites sub-
granulatus* , Perinopollenites elatoides, Classopoltis torosus, C. echinatus, C. hammenii.

Twenty of the species in the above list occur in Suites A, B, or C in England. Species
marked * occur only in Suites B and C and species marked ** occur only in Suite C.
Two of the latter in England are comparable with but not identical to the Dutch forms
( Callialasporites trilobatus and Monosulcites minimus). Thus apart from the presence of
Trilobosporites apiverrucatus, T. bernissartensis, C. trilobatus, and Monosulcites minimus
the species from the Upper Malm that also occur in England suggest a correlation with
Suite B. If the presence of the few Suite C species is considered significant, a correlation
of this sequence with Suite C is indicated. But approximately forty other species also
occur in the Upper Malm of Holland that are not known in the Kimmeridgian-Purbeck
sequence of England. Of particular significance amongst these forms is Peromonolites
asplenioid.es Couper. This species was reported by Couper (1958) to occur in the higher
parts of the Wealden but not in the Fairlight Clay, Ashdown Sand, Wadhurst Clay or
Tunbridge Wells Sand (i.e. this species occurs in the Barremian-Aptian using Hughes
1958 correlation). In view of the local variations in entrances of particular species,
evidence from one species alone is not usually adequate for stratigraphic correlation.
However, the presence of this species down to the Upper Malm together with the evi-
dence from the microfloras reviewed above suggests that the Dutch Upper Malm
microflora is probably no older than Suite C of England and may be younger.

Doring (1965) gave details of the stratigraphic ranges of 109 spore species (mostly
trilete) in 7 boreholes penetrating strata close to the Jurassic-Cretaceous boundary from
an area about 50 miles east of Hamburg, Germany. His section extended from the Upper
Malm (Serpulite) to Wealden G which Doring equated with Wealden 5 and 6 of the
standard NW. German sequence i.e. immediately underlying the Middle Valanginian.
Only 12 of these species are known in Suites A, B, and C of England and of these, the
following range from Upper Malm to Wealden G:

Dictyophyllidites equiexinus, Osmundacidites wellmanii, Pilosisporites trichopapillosus, Klukisporites
pseudoreticulatus, Coronatispora valdensis, Trilobosporites bernissartensis, Cerebropollenites mesozoicus.

In England all these species occur in Suites A, B, and C except Trilobosporitites bernis-
sartensis and Pilosisporites trichopapillosus which only occur in Suite C, suggesting a
correlation of the Upper Malm (Serpulite) with Suite C. This correlation is reinforced by
the entrance in Wealden A (overlying the Serpulite) of Duplexisporites problematicus,
Aequitriradites radiatus, and Cicatricosisporites angicanalis. All these species appear in
England in Suite C. Trilobosporites apiverrucatus does not appear in the German se-
quence until Wealden G however, compared with Suite C in England. Thus on the basis

NORRIS: MIOSPORES FROM SOUTHERN ENGLAND

617

of trilete spores, the Serpulite and Wealden A (= Wealden 1) of Germany correlate
with Suite C of England. Correlations between the English, Dutch, and German sequences
are shown in text-fig. 5, incorporating palynologic correlations in the scheme based on
ostracods by Anderson (1955, 1962).

Acknowledgements. This work was started in the Sedgwick Museum, Cambridge and completed at the
New Zealand Geological Survey, Lower Hutt. I am indebted to Mr. N. F. Hughes for much helpful
discussion and for invaluable assistance at all stages of this study. Material from the Warlingham
Borehole was made available by the Institute of Geological Sciences, London. I am grateful to Dr. C. A.
Fleming and Mr. R. W. Willett of the New Zealand Geological Survey who facilitated continuation
of this work in New Zealand. I also acknowledge help received from Dr. M. E. Dettmann, Dr. W. F.
Harris, Mr. D. J. McIntyre, and Dr. G. Playford on palaeobotanical questions, and from Mrs. A. F.
Norris in technical assistance. Mr. P. B. O’Donovan made the photomicrographs and Mr. F. Jurgeneit
drafted some of the figures. Financial assistance for this work was received from the following
sources: Harkness Scholarship, Cambridge University; Frank Smart Studentship, Gonville and Caius
College, Cambridge; New Zealand Geological Survey; and National Research Council of Canada.

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GEOFFREY NORRIS
Department of Geology
University of Toronto

Typescript received 2 December 1968 Ontario, Canada

WEN LOCK I AN BRYOZOA FROM DUDLEY,
NIAGARA, AND GOTLAND AND THEIR
PALAEOGEOG RAPH IC IMPLICATIONS

by DAVID E. OWEN

Abstract. A comparison is made between the Wenlockian bryozoan faunas of Dudley and those of the
Niagaran region of New York State, and localities near Visby, in Gotland. Their occurrence fits a narrow
Atlantic, the Niagaran and Dudley faunas occupying a different climatic zone from that of Gotland. The following
new species are described from the British material : Meekopora dudleyensis, Stenopora primaeva , and Bato-
stomella maniformis. The new genus Asperopora is proposed for Silurian and lowermost Devonian species of
Leioclema with numerous, often cystose mesopores, and acanthopores which frequently inflect zooecial apertures.

In 1911, Bassler described 155 species and 6 varieties of bryozoa from numerous
Ordovician collections from the Baltic provinces, some made especially for the study.
He was clearly interested in possible links with American species and compared the
faunas from the two regions. He found 62 species and 1 variety (pp. 50-3) common to
both. He commented (p. 32) that, leaving out bryozoa from the Lyckholm and Borkholm
Limestones, which he believed to be of early Silurian age, the number of species com-
mon to Baltic and American regions was 52 out of 134. The above beds are now con-
sidered to be of late Ordovician age.

In 1965, I was able to examine Bassler’s material in the U.S. National Museum, but
would not be prepared to admit that any of the species was, in fact, common to the two
areas. Furthermore, he wrongly recorded the American Silurian forms Hallopora
elegantula (Hall) and Lichenalia concentrica Hall from Baltic Ordovician rocks. His
named sections in the Smithsonian Institution bear little resemblance to the American
types. With further study it is possible that some of the material which he believed to
be common to both areas will turn out to be conspecihc, though the Baltic bryozoan
fauna has a very different aspect from the American equivalents.

In my own work on Silurian bryozoa, 1 had been struck by the differences between the
British and American species. Further, material I had collected in Gotland in 1959
seemed quite different again. Until recently, it has been difficult to compare published
figures of species described from different countries. Many have been illustrated by
drawings and where photomicrographs have been used, sections have often been too
thick to show the detailed structures. In consequence, I collected material from the
Niagara region in 1965 for comparison with my own Wenlockian material, and Llan-
doverian and Wenlockian bryozoa from Gotland in 1959. In both Sweden and America
my time was short and my small collections were made largely from localized spoil
heaps. My British Wenlockian material is from the classic locality of Dudley in
Worcestershire, where I have had longer in the field and have collected my fossils in situ.

In order to make the closest comparisons, I have tried to compare faunas from strata
of similar lithology. This has not been difficult, for the shaly limestones of Dudley,
which lie between the two thick beds of reef limestone which have been extensively
quarried, are lithologically very like the Lower Rochester Shales of the Niagara region.

[Palaeontology, Vol. 12, Part 4, 1969, pp. 621 36, pis. 114-116.]

622

PALAEONTOLOGY, VOLUME 12

I was also able to collect from some thinly bedded, argillaceous limestone of the
Hogklint group near Visby, a rock of similar lithology to the other two.

In British Silurian and Carboniferous deposits, bryozoa occur most commonly in
fore-reef calcirudites rather than in more massive reef limestones. The latter frequently
contain fenestellids and certain incrusting forms, but their bryozoan fauna as a whole
is small in numbers and in variety. A similar distribution was noted by Cuffey (1967) in
Permian rocks. Stach (1936), working on the Tertiary Cheilostomata of New Zealand,
noted a number of zoarial shapes, some of which could stand strong wave action and
littoral conditions. These included certain fenestrate colonies and those incrusting
flexible algae. Articulated zoaria could also stand wave action though some lived in
waters of greater depth. The cylindrical forms, however, needed more shelter. Lagaaij
and Gautier (1965), working on modern bryozoan assemblages of the Rhone delta,
found them rarest in places of most rapid deposition, particularly in front of the main
distributory mouths, and the rate of sedimentation appeared to be a major factor con-
trolling distribution. Working with slight modifications of Stach’s zoarial shapes, they
showed that colonies incrusting a solid substratum and ramose forms occupied the
deeper, quiet waters, but that those incrusting flexible algae or of fenestrate habit could
stand stronger currents and wave action. Bifoliate forms tended to inhabit waters of
around 40-50 m. depth floored by sandy coralline beds.

Thus, the distribution of Palaeozoic bryozoa would not appear to have differed much
from that of Tertiary and Recent species and may have been influenced by the same factors.
A further possible cause may have been the apparent mutual exclusiveness of ramose
bryozoans and tabulate corals in some deposits. They may have competed for food.

Powell (1967), describing Recent bryozoans from north New Zealand, found the
richest fauna at around 53 fathoms. This would present quiet conditions which ap-
parently suit ramose forms. It would also be at the limit of, or just beyond the reach of
light, and consequently beyond the range of corals.

In order to compare the Wenlockian faunas from the three localities, I have calculated
the percentage distribution of the specimens collected. These figures could be improved
by more detailed and representative collecting, but nevertheless give a clear picture of
the relative frequency of the commoner species. Further, the percentages of American
species which 1 collected agree with the notes ‘common’ etc. used by Bassler (1906).
The percentages of British species collected are as follows:

COMPARATIVE FAUNA

Percentage

Fistulipora nummulina Nicholson and Foord
Eridotrypa cava Owen
Rhombopora mawi Owen
Hatlopora elegant ula (Hall)

Batostomella maniformis sp. nov.

Stenopora primaeva sp. nov.

Asperopora multipora (Bassler)

Asperopora aspera (Hall)

Trematopota sp.

Fenestrate Cryptostomata
Other species

24

20

14

5

4

4

4
3
3

5
14

OWEN: WENLOCKI AN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 623

Bifoliate Cryptostomata, though present, made up less than 1 per cent of the material
collected. They include Ptilodictya lanceolata (Goldfuss) and Pachydictya crassa (Hall).

In contrast, the American Lower Rochester Shale species from two localities were
represented as follows:

Percentage

Chilotrypa ostiolata (Hall) 57

Trematopora tuberculosa Hall 17

Hallopora elegantula (Hall) 16

Fenestrate Cryptostomata 5

Other species 5

The other species include specimens of Asperopom aspera (Hall), Asperopora multipora
(Bassler), and a few bifoliate Cryptostomata.

The Gotland Wenlockian fauna is quite different. The percentages are as follows:

Percentage

Helopora lindstromi Ulrich 26

Phaenopora lindstromi Ulrich 22

Fistulipora mutabilis Hennig 1 6

Hallopora spp. 15

Eridotrypa spp. 8

Ptilodictya lanceolata (Goldfuss) 6

Crepipora lunariata Hennig 3

Other species 4

Though species of Hallopora and Eridotrypa are common, their preservation is such
that 1 would not be prepared to identify them specifically. None of them, however,
bears any resemblance to species of Hallopora and Eridotrypa common in Britain and
America. The bifoliate Cryptostomata are beautifully preserved, and no doubt this
preservation led Hennig to devote the first of his three Gotland bryozoan papers to
them. The small Helopora lindstromi Ulrich, first described by Ulrich (1890, p. 343) from
Gotland and later by Hennig ( 1906, pp. 19-23), is also common and very well preserved.
The Trepostomata, on the other hand, are poorly preserved, commonly worn, and would
appear to have drifted some distance from their location in life. Perhaps the bifoliates
lived in a slightly different habitat from that occupied by ramose forms (as with living
bryozoa), and the latter were introduced as detrital material. If so, the Gotland per-
centages are not strictly comparable with those of Dudley and Niagara, but it is unlikely
that this factor would alone account for such marked differences.

The species common to more than one locality are as follows:

Species

Hallopora elegantula (Hall)
Asperopora aspera (Hall)
Asperopora multipora (Bassler)
Pachydictya crassa Hall

America
Very abundant
Common
Common
Fairly common

Ptilodictya lanceolata (Goldfuss) Missing

Britain Gotland

Very abundant
Abundant
Abundant
Occurs

Occurs

Missing

Missing

Missing

Missing

(occurs in Russia)
Common

Other species, notably Berenicea consimilis (Lonsdale) have been recorded in all three
localities (Lonsdale 1839, p. 675; Bassler 1906, pp. 16-17; Hennig 1906, p. 25). I have
not been able to examine specimens from each of the three localities, but the figures

624

PALAEONTOLOGY, VOLUME 12

suggest that the forms do not necessarily belong to one species. For instance, that shown
by Bassler (1906, pi. 5, fig. 5) has a very different appearance from the form illustrated
by Hennig (1906, pi. 3, fig. 8), while that of Lonsdale (1839, pi. 15, fig. 7) is too small to
illustrate the specific features.

It would seem that Ptilodictya lanceolata (Goldfuss) is the only bryozoan species
recognized with certainty both in Britain and in Gotland. This species has been restudied
by Phillips Ross (1960). The Gotland specimens examined were mostly Ludlovian,
though one was upper Llandoverian. Phillips Ross stated (p. 445): ‘Specimens of P.
lanceolata from the Wenlock limestone of Dudley, England, are more robust, longer
and thicker than specimens from the Ludlovian calcareous clay of Mulde, Gotland.’
These differences are relatively unimportant but serve to show that even this species is
not identical in the two localities. Fragments which I collected from the Wenlockian
beds of Gotland are not sufficiently complete to allow an accurate comparison with
British examples.

This brief faunal comparison suggests that the British bryozoan fauna had a few links
with the American but none with that of the Baltic area. Bryozoa today are worldwide
in distribution and known to inhabit the seas from tropical to polar regions. Further,
they are found in water of all depths, from littoral to oceanic. According to Flyman
(1959, pp. 417-30) Arctic species are circumpolar, and extend southwards some way
down the coasts. Boreal and Mediterranean faunas are a mixture of localized and widely
spread species. Arctic and some boreal species occur on both sides of the Atlantic but
most boreal forms are restricted to the east or the west. There is a temperature zoning,
though some species extend from Alaska to Central America and the Galapagos
Islands.

The Wenlockian British and American bryozoan faunas would appear to have
behaved in a similar way to their modern counterparts. Some genera were common to
both, but most species were localized, though a few appeared in both areas. A closer
look at the actual species is revealing. The commonest American form is Chilotrypa
ostiolata (Hall) and the commonest British one Fistulipora nummulina Nicholson and
Foord. These two fistuliporids are remarkably similar in many respects, the main differ-
ence being in form. The hollow cylinders of Chilotrypa ostiolata appear to have incrusted
a particular non-calcareous stem, probably algal. Fistulipora nummulina was also in-
crusting but is found covering a variety of objects. The genera Bcitostomella and Tretna-
topora are represented in both areas though by different species. The place occupied in
America by the very common Trematopora tuberculosa Hall appears to have been taken
in Britain by Eridotrypa cava Owen. Though species of Eridotrypa occur in America,
none has any resemblance to the British species.

A comparison of Atlantic and Baltic species yields markedly different results. The
exception to this is the common incrusting form Fistulipora mutabilis Hennig, which
compares with Fistulipora nummulina and probably occupied a similar niche. Otherwise
the dominant cryptostome element gives the bryozoan fauna of Gotland a distinctive
character.

PALAEOGEOGRAPHY

An examination of the British and American bryozoan faunas of Wenlockian age
shows similarities and contrasts not unlike those obtaining today. This suggests that

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 625

the areas were separated by an early Atlantic ocean, though that ocean need not have
been as wide as at present. Each side had its own local fauna but a few species, notably
Hallopora elegantula ( Hall), were common to both. Perhaps these species had a greater
climatic tolerance and were able to range along the northern shores of the ancient
Atlantic. Hallopora elegantula was not an incrusting species and is unlikely to have
floated across the waters in its adult state.

The difference in the faunas suggests that there was a considerable water barrier
between the two regions even at that time. Free-floating bryozoan larvae exist for a
limited period before becoming attached. It is not known whether the larvae of the
extinct orders were cyphonautes or were comparable to the cheilostome and cyclostome
brooded larvae of today. In the latter case, their unattached life would have been a few
hours only. As ovicells and brood chambers are only known from a few forms in the
extinct orders, their larvae may have been cyphonautes with an unattached life of as
much as two months in certain circumstances.

If the Silurian Atlantic was very narrow, such larvae may have drifted across it. In
like manner, the more fragile incrusting species might have been carried on drifting
algae. In these circumstances the same species of fistuliporid might well be looked for
on both sides. This has not happened. Today driftwood is known to cross the Atlantic.
Scheltema (1966) gave a table of drift bottle data which showed that the minimum time
taken by bottles drifting from the North American coast to the coasts of Europe or
Africa was 364 days. The shorter journey from North America to the Azores took no
less than 125 days. Such journeys were well within the survival limits of small immature
gastropods but too long for bryozoan larvae.

Had the width of the Silurian Atlantic been half the present distance from the
American coast to the Azores, it might still have formed a barrier to most species. If
much less, it would scarcely have been a barrier at all. These comments are necessarily
tentative, since the free-floating life of Palaeozoic bryozoa may have been much shorter,
and ocean currents and winds may not have carried across narrow waters in Silurian
times. Nevertheless, the existence of a narrower ocean which acted as a partial barrier
between the British and American Silurian basins is suggested by the bryozoan fauna.

An interesting study of the effect of a water barrier was made by Cheetham (1963),
describing the late Eocene zoogeography of the American eastern Gulf Coast. He postu-
lated a bank of considerable size, the Ocala Bank, separated from the continental shelf
by a strait, the Suwannee Strait. This was 50-70 miles across, with a variable depth of
about 500 ft., and the bottom waters were slightly colder than those on the surface. This
proved a barrier to a few species, though most occurred on both sides. It would seem
that the Silurian Atlantic must have been a much more formidable barrier than the
Eocene Suwannee Strait.

Spjeldnaes (1961) considered Ordovician climatic zones. He found evidence for a
number of faunal provinces in Western Europe, North Africa, and eastern America,
which could be considered to mark climatic zones if the poles and equator occupied
positions different from those of today. He showed that such zones would be consistent
with a pole somewhere in or west of Africa, and that such a position accorded with
palaeomagnetic observations. Most of Britain including the Welsh Borderland lay in the
Anglo-Scandic Province throughout Ordovician times, and this province extended into
the Oslo region but not into the Baltic. The Appalachian Province was intimately related

626

PALAEONTOLOGY, VOLUME 12

and was evidently in the same climatic zone. The climatic zones of Europe and America
were found to fit better if the continental blocks were considered to be 40° closer
together than now. Such a position would still allow for a narrow intervening Atlantic
Ocean. If the same situation obtained in Silurian times, and the Niagara and Welsh
Borderland regions were in the same climatic province but separated by a narrow
Atlantic, the similarities and differences of the bryozoan fauna would be explained.
The distinctive Baltic Silurian fauna might also be explained, if it lay in a different
climatic zone.

Bulman (1964) considered the distribution of lower Palaeozoic plankton; he plotted
the occurrences of an Ordovician and a Silurian graptolite, and found that they were
consistent with a pole west of Africa. In the case of the Ordovician graptolite, the occur-
rences showed a better fit with the continents in their present positions and the Atlantic
its present width. Though the occurrences of the Silurian graptolite would fit either the
present positions of the continents or a narrower Atlantic, it would be hard to picture
the Atlantic narrowed by Wenlockian times and wider again at the present.

Stormer (1967, pp. 209, 210) noted that the highly fossiliferous coral reefs of the Silurian
of Gotland may have formed the coastal facies of a syneclise, a basin of marine deposits
on the eastern European Platform. Such a basin might have had little or no physical
connection with the Caledonian geosyncline which included the Welsh Borderland
and possibly eastern American Silurian deposits. Such circumstances would explain the
similarities of the American and British bryozoan faunas and their differences from those
of Gotland.

In his work on Silurian cystoids, Paul (1967) found that the two families represented
in Britain, the Echinoencrinitidae and the Callocystitidae, originated from the Baltic
and from North America respectively. This would suggest separate provinces, but the
British intermingling would have called for connections at some period.

Thus, though no completely satisfactory answer is at present available, there must be
a reasonable and natural explanation for the links between the bryozoan faunas of
Britain and New York State, and the marked differences between those faunas and that
of the Baltic in Wenlockian and, I believe, in upper Ordovician times. It seems likely that
the distribution of climatic zones and presence of a narrower Atlantic Ocean offer the
best hypothesis.

SYSTEMATIC PALAEONTOLOGY

All material described is in the collection of the Manchester Museum, whose registration numbers
are quoted.

The following species have been identified in my collection from Dudley:

Amplexopora sp.

Asperopora aspera (Hall)

Asperopora densipora (Owen)

Asperopora multipora (Bassler)

Batostomella maniformis sp. nov.

Calamotrypa millichopensis Owen
Ceramopora sp.

Eridotrypa cava Owen
Eridotrypa sp.

Favositella interpuncta (Quenstedt)

Fistulipora crassa (Lonsdale)

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 627

Fistulipom nummulina Nicholson and Foord
Fistulipora sp.

Hallopora elegant ula ( Hall)

Meekopora dudleyensis sp. nov.

Monotrypa macropora Foord
Monotrypa paterella nom. nov.

Monotrypella benthallensis Owen
Monotrypella pulchella (Edwards and Haime)

Nicholsonella parva Owen
Pachydictya crassa (Hall)

Ptilodictya lanceolata (Goldfuss)

Rhombopora mawi Owen
Stenopora primaeva sp. nov.

Trematopora sp.

A description of new species, of species common to Dudley and Niagara, and those
requiring special comment is given below.

Order cystoporata Astro va 1 964
Family fistuliporidae Ulrich 1882
Genus fistulipora McCoy 1850
Fistulipora nummulina Nicholson and Foord
Plate 1 14, figs. 1 , 2

1885 Fistulipora nummulina Nicholson and Foord, pp. 506-7, text -fig. 4, pi. 15, figs. 2-2 b.
1885 Fistulipora dobunica Nicholson and Foord, p. 511, pi. 17, figs. 3-3 b.

Material. LL3214, 3215.

Description. Zoaria highly variable, disc-shaped, incrusting, or thin expansions, occa-
sionally rising into low domes or branches. When incrusting, cover shells, crinoid stems,
and other bryozoans. Low monticules, consisting of cystopores, occur at intervals of
c. 4 mm.

Zooecia, at first recumbent, turn up rapidly to reach surface at or near right angles.
Wall thickens in exozone on one side to form deep-seated lunaria. Cystopores, closed
at surface, separate zooecia, and occur in groups. In tangential section, zooecia have
prominent lunaria which sometimes give clover-leaf effect to apertures. Slightly larger
apertures surround monticules, lunaria directed towards each monticule. Apertures
average 0-2 mm. across; 8 occur in 2 mm. Lunaria average 0- 10-0-14 mm. across.

Remarks. 1 had considerable difficulty in deciding whether the many specimens which
I collected were F. nummulina or F. dobunica. The former is described as ‘discoid,
lenticular, concavo- or plano-convex, also forming thin, irregular crust-like expansions’,
and the latter ‘extremely thin, encrusting’. The former is dotted with monticules con-
sisting of mesopores, and the latter with maculae. In both, the lunaria are well developed,
the apertures in F. nummulina being larger, 4 in 1 mm., and those in F. dobunica smaller,
6 in 1 mm.

I have been unable to locate the types of either species. That of F. nummulina was
presented to either Nicholson or Foord by W. Madeley of Dudley and these authors
refer to the fact that other specimens occur in the Holcroft Collection, now in Birming-
ham University museum. The type of F. dobunica was also said to be in Madeley’s
Collection. Named specimens in the collections of the British Museum (Natural History)
appear to be indistinguishable from F. nummulina. Furthermore, my own collections

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PALAEONTOLOGY, VOLUME 12

show every gradation from one to the other and I regard F. dobunica as less mature
zoaria of F. nummulina. As F. nummulina was described first, F. dobunica appears to be
a junior synonym.

F. nummulina is the commonest species in the Dudley area, Chilotrypa ostioJata (Hall)
is by far the commonest in New York State, and F. mutabilis Hennig is the third most
common species in my Wenlockian Gotland collection. Careful examination shows these
three to have close similarities even though they are unrelated. Each is incrusting, though
C. ostioJata is always hollow. All have well-developed monticules, particularly in the
larger zoaria, and these are raised above the surface and consist of closed cystopores. All
have well-developed lunaria, though only those of F. nummulina are deep-seated. The
three species would seem to have occupied the same ecological niche.

According to Borg (1965) maculae and monticules originated as points of growth.
Whether this is correct in any or all instances, there is little doubt that raised monticules
in larger zoaria would have been protective. Whether they consisted of cystopores,
closed mesopores, or acanthopores, they would be a protection against abrasion,
whether from a free-swimming host, a series of waving fronds, or the tumbling of shells
by wave action.

Genus meekopora Ulrich 1889

Meekopora dudleyensis sp. nov.

Plate 1 14, figs. 3, 4

Material. LL3216, 3217 (paratypes).

Diagnosis. Meekopora with well-developed lunaria and maculae consisting of small
circular groups of cystopores.

Description. Zoaria bifoliate, variable in size, from strap-like forms 4-5 mm. across,
to leaf-shaped examples up to 15 mm. wide and 1-T5 mm. thick. Small circular maculae
occur, consisting of cystopores, surrounded by apertures whose lunaria lie towards each
macula. Zooecia arise from imperforate mesotheca which consists of two transparent
layers separated by dark laminar layer; zooecia separated by cystopores whose surface
layer is formed of dense, granular tissue. Apertures oval, averaging 0T2-0T6 mm. x
0-16-0-20 mm., with well-developed lunaria 0-08-0-10 mm. across.

Remarks. The strongly marked lunaria, the small apertures, the small circular maculae,
and the flat, strap-like or leaf-like zoaria serve to identify this species even when there
is no cross-section visible to show the mesotheca. It differs from Meekopora foliacea

EXPLANATION OF PLATE 114

Fig. 1. Fistulipora nummulina Nicholson and Foord from Dudley. Vertical section LL3214, X 50.

Fig. 2. Fistulipora nummulina Nicholson and Foord from Dudley. Tangential section showing clover-
leaf-shaped apertures and lunaria. LL3215, X 50.

Fig. 3. Meekopora dudleyensis sp. nov. from Dudley. Vertical section showing mesotheca. Paratype,
LL3216, x 50.

Fig. 4. Meekopora dudleyensis sp. nov. from Dudley. Tangential section showing small apertures with
lunaria facing macula. Paratype, LL3217, x 50.

Fig. 5. Batostomella maniformis sp. nov. from Dudley. Vertical section showing thick exozone con-
taining acanthopores. Holotype, LL3218a.

Fig. 6. Batostomella maniformis sp. nov. from Dudley. Tangential section showing small apertures
and acanthopores. Holotype, LL3218b, x50.

Palaeontology, Vol. 12

PLATE 114

OWEN, Wenlockian bryozoa

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 629

(Hall), which it most nearly resembles, in that the American species has larger apertures,
and narrow, elongate dividing maculae. Like most fistuliporids, the species is variable
but the characteristic shape of the maculae of M.foliacea seems to rule out the possibility
of the two species being conspecific.

Order trepostomata Ulrich 1882
Family batostomellidae Miller 1889
Genus batostomella Ulrich 1882
Batostoniella maniformis sp. nov.

Plate 1 14, figs. 5, 6

Material. LL3218 (holotype, specimen with five sections, LL3218a-e).

Diagnosis. Batostomella with a few large, deep-seated acanthopores in wide, laminate
exozone.

Description. Zoaria ramose, with many branches apparently rising from a disc, each
branch 1-1-5 mm. diameter, with small spines at surface marking position of acantho-
pores. Complete zoarium may be up to 30 mm. across with branches 25 mm. long.
Zooecia arise axially, thin-walled, and turn sharply into exozone to reach surface at or
near a right angle. Walls in exozone thick, laminate, with large, deep-seated, acantho-
pores with clear axes. No diaphragms. In tangential section, oval apertures widely
separated, ringed. Acanthopores with clear axes occur intermediately, 3 or 4 surrounding
each aperture. Mesopores absent. Apertures 0-1-0-12 mm. x 0-05-0-07 mm., 10 in 2 mm.
longitudinally, 15 in 2 mm. laterally or diagonally. Acanthopores 0-02 mm. in diameter.

Remarks. The holotype shows branches bifurcating at intervals of 5-8 mm. and the
whole zoarium has a hand-like appearance. In section, it bears no resemblance to any
described species. I have also collected fragments from the Lower Bringewood beds of
Millichope, near Ludlow. The form is slightly smaller than the American B. granulifera
(Hall), differing also in having relatively few acanthopores compared with the large
numbers in that species, a thicker exozone, and no mesopores. In other respects, it is
a closely allied species.

There is little difference between Silurian species of Batostomella and of Rhombopora
and its allies. The latter have more clearly marked vestibules, often partly closed by
hemisepta, but the genera must be closely related.

Family stenoporidae Waagen and Wentzel 1886
Genus stenopora Lonsdale 1 844
Stenopora primaeva sp. nov.

Plate 1 15, figs. 1, 2

Material. LL3219a-c, 3220 a, b (paratypes).

Diagnosis. Stenopora with large, clear centred acanthopores.

Description. Zoaria ramose, with small spines marking positions of large acanthopores.
Diameter 2-5-4 mm. Zooecia arise axially and curve gently to reach surface at c. 70°.
Moderately thin walled in endozone, walls thicken rapidly in exozone, which varies in

630

PALAEONTOLOGY, VOLUME 12

width from c. 0-5 mm. in thinner branches to 1 mm. in stouter specimens. Exozone walls,
commonly moniliform, have laminae which form wide, flat U distally. Within this region,
large acanthopores with clear axes occur. Diaphragms present, few in number except
in stoutest specimens. Examples frequently show an incipient development of exozone
marking slight pause in growth; this feature often seen in stenoporids. Mesopores
absent. Apertures oval, ringed, 0-15-0-3 mm. x 0-08-0- 15 mm., the longer axis approxi-
mately parallel with that of growth; 5 occur in 2 mm. longitudinally, 7-8 laterally.
Large acanthopores numerous, diameter 0-03-0-05 mm., sometimes inflect zooecial
wall, 5-8 ringing each aperture. Thickened walls with numerous acanthopores may form
maculae.

Remarks. Though Stenopora tasmaniensis Lonsdale is Permian, and the genus has not
been recorded in rocks older than the Carboniferous, this species is so similar in section
to the type material of S. tasmaniensis that I consider it congeneric. S. tasmaniensis is
considered to have no diaphragms. None was visible in the type section I examined, but
the solid specimen made me wonder if occasional diaphragms might occur. Diaphragms
are few in S. primaeva, occurring at the base of the exozone, especially in thicker
specimens.

S. primaeva differs from S. tasmaniensis in having fewer, much larger acanthopores
and generally a rather thinner exozone. A thick exozone may, however, merely reflect
greater longevity owing to congenial conditions.

Genus asperopora gen. nov.

Ulrich (1882) proposed the genus Leioclema for the Carboniferous species Callopora
punctata Hall, which he considered to be quite distinct from Callopora elegantula Hall.
The original description (p. 141) was as follows:

Zoarium ramose, branches slender, smooth and sometimes hollow. Cell apertures small, rounded,
and with two or three series of subangular interstitial cells surrounding them. Longitudinal sections
show the tubes in the axial portion of the branch to be thin-walled, and crossed by remote diaphragms.
In the peripheral region the interstitial tubes and spiniform tubuli are developed in great numbers.
The walls of all the tubes are much thickened, and diaphragms in the interstitial tubes are straight
and remote, while in the true tubes they appear to be wanting. In tangential sections the visceral cavity
of the proper zooecia is often indented by the encroachment of the rather large spiniform tubuli, of

EXPLANATION OF PLATE 115

Fig. 1. Stenopora primaeva sp. nov. from Dudley. Tangential section showing large, clear-centred
acanthopores. Paratype, LL3219a, x50.

Fig. 2. Stenopora primaeva sp. nov. from Dudley. Vertical section showing wide exozone and deep-
seated acanthopores. Paratype, LL3220a, x 50.

Fig. 3. Asperopora aspera (Hall) from Dudley. Vertical section showing deep-seated, clear -centred
acanthopores. LL3221, x 50.

Fig. 4. Asperopora aspera (Hall) from Dudley. Tangential section showing apertures surrounded by
mesopores, and acanthopores inflecting the apertures. LL3222, X 50.

Fig. 5. Asperopora aspera (Hall) from Niagara. Vertical section showing deep-seated, clear-centred
acanthopores. Diaphragms are fewer in this specimen. LL3223a, x 50.

Fig. 6. Asperopora aspera (Hall) from Niagara. Tangential section; the specimen is partly recrystallized
but the large acanthopores are seen to inflect the apertures. LL3223b, x 50.

Palaeontology , Vol. 12

PLATE 115

OWEN, Wenlockian bryozoa

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 631

which there are, in the type species, from four to seven round the orifice of each tube. The interstitial
tubes are small, and of irregular shape; two or three rows occupy each intertubular space.

He noted that the species was Carboniferous (p. 154). Ulrich (1890, p. 425) stated:

When this genus was proposed I was acquainted with the type species, Callopora punctata Hall,
only. Now by extending the limits, fifteen species are united under the name.

He suggested that more than one generic group might be included in the genus as he
then defined it. The 15 species occurred in rocks of Upper Ordovician to Pennsyl-
vanian age.

Since then many other species have been added. In 1965, Boardman drew my attention
to the fact that the Devonian species of Leioclema were markedly different from the
Carboniferous species. I examined the type material in the U.S. National Museum and
realized that, with Dyscritella Girty 1910, it appeared to be related to such genera as
Rhombopora Meek 1872 and other rhabdomesids which are normally placed in the order
Cryptostomata.

I then examined the sectioned types of many species of Leioclema described by Ulrich,
Bassler, Girty, and others which are housed in the U.S. National Museum collections.
Those from Silurian and Helderbergian deposits formed a moderately tight group which
probably also includes those from upper Ordovician rocks. These forms did not survive
the Helderbergian and differ markedly from Devonian and Carboniferous species.

Vinassa de Regni (1920) proposed a new type species for Leioclema , the Silurian L.
explanation Bassler, and suggested a new genus Leioclemina, based on L. ramulosum
Bassler, for species with few or no diaphragms. The first suggestion is invalid since
Ulrich’s type species exists and is well known. The diagnosis of Leioclemina precludes
this genus from embracing the many common typical Silurian species, and appears to
be quite unnecessary.

1 therefore propose the genus Asperopora based on Callopora aspera Hall, Leioclema
asperum (Hall) Bassler, for the Silurian species.

Diagnosis. Zoaria incrusting, massive or ramose; zooecial walls relatively thick; meso-
pores numerous, sometimes cystose; acanthopores large, with clear axes, commonly
deep-seated and inflecting the apertures, which may become petalloid; diaphragms few
to many, particularly in mesopores.

The species I have examined which belong to this genus are:

Leioclema asperum (Hall)

Leioclema conduction Bassler
Leioclema densiporum Owen
Leioclema explanation Bassler
Leioclema globulare Bassler
Leioclema halloporoides Owen
Leioclema inornatum Bassler
Leioclema ludlovensis Owen
Leioclema multiporum Bassler
Leioclema pulchellum Bassler
Leioclema ramosum Owen
Leioclema ramulosum Bassler
Leioclema subramosum Ulrich and Bassler
Leioclema variaporum Billings

Silurian: Niagaran and Wenloclcian

Devonian : Helderbergian
Silurian; Wenlockian
„ Niagaran
Devonian ; Helderbergian
Silurian

Wenlockian

Niagaran and Ludlovian

Ludlovian
Waldron Shale
Ludlovian

Niagaran and Wenlockian

632

PALAEONTOLOGY, VOLUME 12

Asperopora aspera (Hall)

Plate 115, figs. 3-6

1852 Callopora aspera Hall, p. 147, pi. 40.

1884 Fistulipora ludensis Nicholson, p. 120.

1890 Leioclema asperum (Hall); Ulrich, p. 425.

1900 Lioclema asperum (Hall); Nickles and Bassler, p. 302.

1906 Lioclema asperum (Hall); Bassler, p. 32, pi. 11, figs. 1-3; pi. 24, figs. 14-16.

1965 Leioclema asperum (Hall); Owen, p. 106, pi. 3, figs. 5, 6.

Material. LL3221, 3222, 3223, 3223 a, b.

Description. Zoaria incrusting to massive, often layered. Under certain circumstances
very large specimens may occur. No. 564 of the Holcroft Collection, Birmingham, com-
pletely covers upper and side surfaces of He/iolites sp., extending for 30 mm. over one
side of the base. Total size of specimen is 1 14 x 105 mm. diameter, 59 mm. high. Overlap
on to lower surface on one side suggests coral was already dead and had been dislodged
and tilted. Most other zooecia examined have been of much smaller size.

Slightly raised monticules containing zooecia of somewhat larger than normal size
are visible on careful examination. Spines marking large acanthopores are also seen on
unworn surfaces.

In longitudinal section, moderately thin-walled zooecia lie horizontally along epitheca
in short endozor.e and turn at right angles into broad exozone, where walls are thicker,
sometimes slightly beaded, and many mesopores separate zooecia. Large acanthopores
occur throughout exozone and protrude as spines. Zooecial diaphragms few, but those
in mesopores are about a tube width apart.

In tangential sections, circular or oval apertures are completely separated by meso-
pores and inflected by large, axially clear acanthopores, which are also associated with
mesopores. Structure of wall not apparent. Dimensions of zooecia, etc., as in Owen
(1965, p. 106).

Remarks. The species is relatively common in the Niagara region and in the Wenlock
Limestone of Dudley and Wenlock Edge. In both, the spiny acanthopores are a marked
feature of unworn specimens. The size of apertures is almost identical, as is the size and
disposition of both mesopores and acanthopores. Walls appear to be somewhat thicker
in the British specimens and diaphragms are more numerous but these features were
probably linked with environmental conditions, and I have no doubt that the species
is the same in both localities.

Needless to say such large zoaria have not escaped notice in Britain and have been
described under the name of Fistulipora ludensis Nicholson. There is an undoubted
similarity of appearance between the mesopores of this species and some fistuliporid
cystopores, though the general structure is typically trepostomatous. Further, the
numerous large acanthopores, noted and figured by Nicholson (1884, p. 120) are quite
distinctive.

Asperopora multipora (Bassler)

Plate 116, figs. 1, 2

1906 Lioclema multiporum Bassler, p. 34, pi. 11, figs. 4-6; pi. 23, fig. 18; pi. 27, fig. 20.

Material. LL3224.

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 633

Description. Zoaria incrusting brachiopods, crinoid stems, etc. Surface not unlike A.
aspera but petalloid apertures are smaller, mesopores more numerous and acanthopores
smaller and less protuberant as spines. No monticules or maculae visible. In vertical
section, zooecia are at first recumbent, and turn almost immediately into exozone to
run straight to surface. Diaphragms few. Mesopores very numerous, often cystoid.
Acanthopores axially clear, moderately deep-seated. In tangential section, deeply
inflected, petalloid zooecia separated by numerous oval and polygonal mesopores.
Acanthopores with clear axial zones exceedingly numerous. Structure of walls not
apparent. Diameter of apertures 0-15-0-20 mm. Diameter of mesopores 0-04-0-08 mm.
Diameter of acanthopores up to 0-05 mm. 7-9 apertures in 2 mm., each surrounded by
4-7 acanthopores. Zooecial wall at surface 0-02-0-03 mm. thick.

Remarks. This species is relatively rare in Britain but fairly abundant in the Niagara
region. The specimens I collected in America are partly recrystallized and do not illus-
trate the characters clearly. I have examined sections labelled by Bassler in the British
Museum (Natural History) and consider that the species is identical on both sides of
the Atlantic. Unfortunately, these sections are too thick to allow the features to show
well in a photomicrograph.

1851 Calopora elegantula Hall, p. 400.

1851 Chaetetes fletcheri Edwards and Haime, p. 271.

1852 Cal/opora elegantula Hall, p. 144, pi. 40, figs, \a-\m.

1854 Monticulipora fletcheri Edwards and Haime, p. 267, pi. 62, figs. 3-3 a.

1882 Callopora elegantula Hall; Ulrich, p. 250, pi. 11, figs. 6-6 b.

1884 Callopora nana Nicholson, p. 120, pi. 7, figs. 4-4 b.

1884 Callopora fletcheri (Edwards and Haime); Nicholson, p. 120, pi. 7, figs. 5-5 b.

1900 Callopora elegantula Hall; Nickles and Bassler, p. 188.

1906 Callopora elegantula Hall; Bassler, p. 41, pi. 17, figs. 11-15; pi. 26, fig. 12.

1911 Hal/opora elegantula (Hall); Bassler, pp. 325-6.

1938 Callopora fletcheri (Edwards and Haime); Stubblefield, p. 30.

1953 Hallopora elegantula (Hall); Bassler, G112, fig. 74, \a-\d.

1961 Calopora elegantula Hall; Phillips Ross, p. 55.

1965 Hallopora elegantula (Hall); Owen, pp. 109-10, pi. 4, figs. 3, 4.

Material. LL3225, 3226, 3227, 3228.

Description. As in Owen (1965, p. 109). Zoaria more variable than previously described,
commonly include small bosses up to 15 mm. diameter, 10-12 mm. height.

The following measurements show that differences between Dudley and Niagara
specimens are well within the limits of variation of individuals in each area:

Family halloporidae Bassler 1911
Genus hallopora Bassler 1911
Hallopora elegantula (Hall)

Plate 1 16, figs. 3-6

Percentage circular apertures

Dudley

56

Niagara

44

Average diameter
Greatest diameter
Smallest diameter

0-29 mm.
0-40 mm.
0-20 mm.

0-33 mm.
0-36 mm.
0-30 mm.

T t

C 6940

634

PALAEONTOLOGY, VOLUME 12

Average size of oval apertures
Greatest measured
Smallest measured

Number of apertures or major parts in 2 sq. mm.
Average number of diaphragms in 1 mm. exozone
Average number of diaphragms in 1 mm. meso-
pore

Thickness zooecial wall in exozone

Dudley

0-32x0-28 mm.
0-36 x 0-32 mm.
0-26 x 0-22 mm.
14-15
4

Niagara

0-30x0-28 mm.
0-36x0-34 mm.
0-22x0-20 mm.
15
3-5

15

0-02-0-03 mm.

15

0-02-0-04 mm.

Remarks. It has already been stated (Owen 1965, p. 110) that H. elegantula and H. nana
(Nicholson) are synonymous. Edwards and Haime (1851, p. 271) briefly described
Chaetetes fletcheri but left it unfigured. In 1854 (p. 267), they redescribed it as Monticuli-
pora fletcheri and illustrated it (pi. 62, figs. 3-3n). Stubblefield (1938) recognized
the actual specimen they had figured though it was shown the wrong way round on the
plate (a mirror image). He therefore designated the lectotype GSM5631. Though the
lectotype is unsectioned, I have examined it carefully and have no doubt that it is con-
specific with H. elegantula. Further, a series of sections labelled by Nicholson Callopora
fletcheri (Edwards and Haime) and now in his collection in the Royal Scottish Museum
are typical examples of H. elegantula. This poses a problem. Hall’s first description,
unfigured, was 1851 and his first figure 1852. Edwards and Haime’s first description
was 1851 and their first figure 1854. It would therefore appear that H. elegantula (Hall)
has priority as a properly described species.

Family trematoporidae Miller 1889
Genus trematopora Hall 1851
Trematopora sp.

Material. LL3229, 3230 a, b.

Description. Zoaria small, ramose, apparently without monticules. Zooecia thin walled
in endozone, turn sharply into short exozone to reach surface at right angles. Occasional
diaphragms occur, some incomplete, attached to proximal wall only and extending
three-quarters of the way across tube. Numerous short mesopores with thick diaphragms.
Acanthopores numerous. In tangential section, oval apertures separated by polygonal
mesopores; acanthopores occur both within area of mesopores and on aperture wall.
In the latter case, apertures may be inflected. Apertures average 0-18 mm. xO-12 mm.,
7-9 occurring in 2 mm.

EXPLANATION OF PLATE 116

Fig. 1. Asperopora multipora (Bassler) from Dudley. Vertical section showing cystose mesopores and
acanthopores. LL3224, x 50.

Fig. 2. Asperopora multipora (Bassler) from Dudley. Tangential section showing petalloid apertures
and numerous mesopores. LL3224, X 50.

Fig. 3. Hallopora elegantula (Hall) from Niagara. Vertical section showing wall laminae veeing
distally. LL3225, X 50.

Fig. 4. Hallopora elegantula (Hall) from Niagara. Tangential section showing ringed circular apertures
and polygonal mesopores. LL3226, X 50.

Fig. 5. Hallopora elegantula (Hall) from Dudley. Vertical section showing wall structure and dia-
phragms closely spaced in endozone and in mesopores. LL3227, x 50.

Fig. 6. Hallopora elegantula (Hall) from Dudley. Tangential section showing ringed circular apertures
and polygonal mesopores. LL3228, X 50.

Palaeontology, Vol. 12

PLATE 116

OWEN, Wenlockian bryozoa

OWEN: WENLOCKIAN BRYOZOA FROM DUDLEY, NIAGARA, GOTLAND 635

Remarks. Though this is a common form, and very similar to the three commoner
American Silurian species noted below, I have not found perforated diaphragms in the
mesopores. Furthermore, mesopores are very short and sometimes closed by thickened
diaphragms, but they seldom show the beaded appearance so typical in T. tuberculosa
Hall and allied species. The absence of tubercles also distinguishes it from that species
but it may yet prove to be closely related to such species as T. whitfieldi Ulrich or T. halli
Ulrich. The material is not sufficiently diagnostic to permit specific determination.

Monotrypa paterella nom. nov.

Dr. F. K. McKinney has informed me that the name Monotrypa patera Owen 1962
is preoccupied by Monotrypa ( Dianulites ) patera Vinassa de Regni 191 1 and that under
Article 57 (a) of the International Code of Zoological Nomenclature M. patera Owen
is a junior homonym. I therefore propose the name Monotrypa paterella for this species.

Acknowledgements. I am grateful to Dr. R. S. Boardman for making available collections in the U.S.
National Museum; the Director of the British Museum (Natural History) and Dr. C. Scrutton for
giving me access to their collections; the Director of the Institute of Geological Sciences and Mr. D.
White for allowing me to study the collections at the Geological Museum; the Director of the Royal
Scottish Museum and Dr. C. D. Waterston for letting me examine sections from the Nicholson
Collection; Professor F. W. Shotton and Dr. I. Strachan for permitting me to study the Holcroft
Collection at Birmingham University. I would like to thank Mr. W. J. Kilgour for taking me into the
field in the Niagara region and Dr. A. Hede for providing me with a marked map of the Visby region.
I also record my thanks to Dr. J. N. Carruthers and Professor Sir Alistair Hardy for advice on Atlantic
drift currents.

REFERENCES

astrova, g. g. 1964. A new order of Palaeozoic Bryozoa. Paleont. Zh. 2, Moscow, 22-31.
bassler, R. s. 1906. The bryozoan fauna of the Rochester Shale. Butt. U.S. geol. Surv. 292, 1-137.

1911. The early Paleozoic Bryozoa of the Baltic Provinces. Bull. U.S. natn. Mus. 77, 1-382.

1953. In moore, R. c. (ed.), Treatise on invertebrate paleontology, Part G, Bryozoa, G1-G253.

Geol. Soc. Am. and Univ. Kansas Press.

borg, f. 1965. A comparative and phyletic study on fossil and recent Bryozoa of the suborders Cyclo-
stomata and Trepostomata. Ark. Zook Ser. 2, 17 (1), 1-91.
bulman, o. m. b. 1964. Lower Palaeozoic plankton. Quart. J! geol. Soc. Lond. 120, 455-76.
cheetham, a. h. 1963. Late Eocene zoogeography of the eastern Gulf Coast region. Mem. geol. Soc.
Am. 91, 1-113.

cuffey, r. j. 1967. Bryozoan Tabulipora carbonaria in Wreford Megacyclothem (Lower Permian) of
Kansas. Univ. Kansas Paleont. Contrib. 1, 1-96.

edwards, h. m. and haime, j. 1851. Monographie des polypiers fossiles des terrains palaeozolques.
Arch. Mus. natn. Hist. not. Paris, 5, 1-502.

1854. Corals from the Silurian Formation. A monograph of the British fossil corals. 5,

Palaeontogr. Soc. ( Monogr .), London.

girty, g. H. 1910. New genera and species of Carboniferous fossils from the Fayetteville Shale of
Arkansas. Ann. N.Y. Acad. Sci. 20, 189-238.

hall, j. 1851. New genera of fossil corals from the report by James Hall, on the Palaeontology of
New York. Am. J. Sci. 11, 398-401.

1852. Natural history of New York. Palaeontology. 2, Albany.

hennig, a. 1905. Gotlands Silur-Bryozoer, 1. Ark. Zook 2 (10), 1-37.

1906. Ibid. 2. Ark. Zook 3 (10), 1-62.

1908. Ibid. 3. Ark. Zook 4 (21), 1-64.

hyman, l. h. 1959. The invertebrates. Smaller coelomic groups. Ectoprocta, 275-515. New York,
London, and Toronto.

636

PALAEONTOLOGY, VOLUME 12

lagaaij, r. and gautier, y. v. 1965. Bryozoan assemblages from marine sediments of the Rhone delta,
France. Micropaleontology, 11 (1), 39-58.
lonsdale, w. 1839. In murchison, r. i., The Silurian system, 577-712. London.
meek, f. b. 1872. Report on the paleontology of eastern Nebraska, with some remarks on the Carboni-
ferous rocks of that district. Final Rep. U.S. geol. Surv. Nebraska, 141-58.

Nicholson, H. A. 1881. On the structure and affinities of the genus Monticulipora and its sub-genera.
xvi, 240. Edinburgh and London.

1884. Contributions to micro-palaeontology. Notes on some species of Monticuliporoid corals

from the Upper Silurian rocks of Britain. Ann. Mag. nat. Hist. (5) 13, 1 17-27.

and foord, a. h. 1885. On the genus Fistulipora McCoy with descriptions of several species.

Ibid. (5) 16, 496-517.

nickles, j. m. and bassler, r. s. 1900. A synopsis of American fossil Bryozoa. Bull. U.S. geol. Surv.
173, 1-663.

owen, d. e. 1962. Ludlovian Bryozoa from the Ludlow district. Palaeontology, 5, 195-212.

1965. Silurian Polyzoa from Benthall Edge, Shropshire. Bull. Br. Mus. nat. Hist., Geol. 10 (4),

96-117.

Paul, c. r. c. 1967. The British Silurian Cystoids. Ibid. 13 (6), 297-355.

Phillips ross, j. R., 1960. Type species of Ptilodictva — Ptilodictya lanceolata (Goldfuss). J. Paleont.
34, 440-6.

1961. Ordovician, Silurian and Devonian Bryozoa of Australia. Bull. Bur. Miner. Resour. Geol.

Geophys. Aust. 50, 1-172.

powell, n. a. 1967. Polyzoa (Bryozoa) — Ascophora — from North New Zealand. ‘ Discovery' Rep. 34,
199-394.

scheltema, R. s. 1966. Evidence for trans- Atlantic transport of gastropod larvae belonging to the
genus Cymatium. Deep Sea Res. 13, 83-95.
spjeldnaes, n. 1961. Ordovician climatic zones. Norsk, geol. Tidsskr. 41, 45-77.
stach, l. w. 1936. Correlation of zoarial form with habitat. J. Geol. 44, 60-5.

stormer, l. 1967. Some aspects of the Caledonian geosyncline and foreland west of the Baltic Shield.
Quart. Jl geol. Soc. Lond. 123, 183-204.

Stubblefield, c. j. 1938. The types and figured specimens in Phillips’ and Salter’s Palaeontological
Appendix to John Phillips’ Memoir on ‘The Malvern Hills compared with the Palaeozoic District
of Abberley, etc.’ Mem. geol. Surv. Summ. Prog. 2, 27-51.
ulrich, e. o. 1882. American Palaeozoic Bryozoa. /. Cincinn. Soc. nat. Hist. 5, 121-75, 232-57.

1890. Palaeozoic Bryozoa. Geol. Surv. Illinois, 8, 283-728.

vinassa de regni, p. e. 1911. Trias-Tabulaten, Bryozoen und Hydrozoen aus dem Bakony. Residtate
Wiss. Erforschung des Balatonsees, 1 (1), No. 4.

— 1920. Sulla classificazione dei Trepostomidi. Atti Soc. Ital. Sci. nat. 59, 212-31.

DAVID E. OWEN

The Manchester Museum
The University
Manchester 13

Typescript received 27 January 1969

ORDOVICIAN STROMATOPOROIDS FROM
NEW SOUTH WALES

by B. D. WEBBY

Abstract. The stromatoporoid faunas from the Ordovician limestones in the central-west part of New South
Wales are described, and a local biostratigraphical scheme using stromatoporoids and corals for correlation of
the limestones is outlined.

The lower member of the Cliefden Caves Limestone and equivalent horizons has yielded only labechiids.
The six species include representatives of Pseudostylodictyon, Rosenella, Labechia, Cystistroma, and Strato-
dictyon gen. nov. (comprising .S’, ozakii sp. nov. and S. columnare sp. nov.). The upper part of the middle
member and the upper member of the Cliefden Caves Limestone, and its correlatives, have produced a fauna
comprising four labechiid species (including Pseudostylodictyon inequale sp. nov. and Cystostroma cliefdenense
sp. nov.) and four clathrodictyid species (including Ecclimadictyon nestori sp. nov.). In addition, the upper
member contains a distinctive new form, Cliefdenella etheridgei gen. et sp. nov., the sole representative of the
new Family Cliefdenellidae.

Approximately half the described species, including both labechiids and clathrodictyids, exhibit Asian and
east European affinities; the others appear to be endemic. None of the species is closely related to North
American forms. At generic level, Cystistroma, Stratodictyon, and Cliefdenella can only be doubtfully regarded
as endemic to Australia. Excluding the supposed Cambrian stromatoporoids, the occurrences of Clathrodictyon
and Ecclimadictyon are the earliest known, of possible Lower Eastonian (= middle Caradoc or ‘Trentonian’)
age.

Apart from the account of Cystistroma donnellii by Etheridge (1895), no Ordovician
stromatoporoids have been described previously from Australia. The type specimens
of C. doimellii were probably collected originally from an horizon at Fossil Hill, in the
lower member of the Cliefden Caves Limestone (Stevens 1952). The stromatoporoids
described are mainly from the Cliefden Caves Limestone, though they are supplemented
by material from other Ordovician limestones in central-western New South Wales
(text-fig. 1), particularly from the Regan’s Creek Limestone, the Reedy Creek Limestone,
the Bowan Park Limestone, the Cargo Creek Limestone, the Canomodine Limestone,
and an unnamed limestone near Gunningbland, 16 miles west of Parkes.

STROMATOPOROID DISTRIBUTION

Three stratigraphically distinct units in the lower member of the Cliefden Caves Lime-
stone contain labechiid stromatoporoids (text-fig. 2). The ‘lower coral’ unit has pro-
duced Stratodictyon ozakii gen. et sp. nov., Labechia regidctris Yabe and Sugiyama, and
Cystistroma donnellii ; the ‘ mixed fauna ’ unit contains Pseudostylodictyon aft', poshanense
Ozaki, S. columnare sp. nov., Rosenella woyuensis Ozaki, L. regularis , and C. donnellii ;
and the ‘upper big shell’ unit contains R. woyuensis. Cystostroma cliefdenense sp. nov.
is the next species to appear, towards the middle of the middle member. It occurs with
P. inequale sp. nov., L. variabilis Yabe and Sugiyama, and the first clathrodictyids,
Clathrodictyon aff. mammillatum (Schmidt), C. cf. microundulatum Nestor, and Ecclima-
dictyon nestori sp. nov. in the upper part of the middle member. In the ‘Island’ unit
of the upper member, the clathrodictyids, C. cf. microundulatum , E. nestori, and E.
amzassensis (Khalfina) occur with Cliefdenella etheridgei gen. et sp. nov.

[Palaeontology, Vol. 12, Part 4, 1969, pp. 637-62, pis. 117-129.]

638

PALAEONTOLOGY, VOLUME 12

text-fig. 1. Map showing location of main stromatoporoid-bearing Ordovician
limestones in New South Wales.

A similar distribution occurs in the Regan’s Creek Limestone (Stevens 1956) with the
labechiids, L. regularis and C. donnellii at the base, C. cliefdenense and Cryptophragmus ?
sp. through the middle, L. variabilis at the top of the middle, and the clathrodictyid
E. amzassensis in the upper part of the limestone.

L. regularis and C. donnellii have been collected from the lower part of the Reedy
Creek Limestone (Adrian 1956; Phillips Ross 1961) near Molong. Further north, near
Eurimbula, E. amzassensis and E. nestori occur in isolated limestone lenses.

E. amzassensis also comes from a thinly bedded unit in the lower part of the Cargo

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 639

Creek Limestone (Stevens 1950), and from a similar horizon in the Canomodine Lime-
stone (Stevens 1950).

In the Bowan Park Limestone (Stevens 1956), S. columnare and Cryptophragmus ? sp.
come from near the base of the formation, and Clathrodictyon aff. mammillatum and E.

7K 7F

CE

LU

Q_

_ Q.

i 4-

Al

ISLAND

I III

o o o

Q

Q

T

\ ‘ LITHIC’ “

'UPPER BIG SHELL'

'MIXED FAUNA'
'UPPER coral'
BRACHIOPOD'
‘LOWER CORAL'

Y-Yi'if

Al s l '

LOWER BIG SHELL

Clieldenella etheridgei, Clathrodictyon
cf. microundulatum. Ecclimadictyon
amzassensis, E. nestori

Cystostroma cliefdenense, Pseudostylodictyon
inequale, Labechia variabil is, Clathrodictyon
aft, mammillatum, C. cf. microundulatum,
Ecclimadictyon nestori

Cystostroma cliefdenense

r ioo

20-

Metres

Feet

0 -

L 0

Rosenel la woyuensis

Pseudostylodictyon aff. poshanense, Stratodictyon
columnare, Rosenel la woyuensis, Labechia
regularis, Cystistroma donnellii

Stratodictyon ozakii, Labechia regularis,
Cystistroma donnellii

text-fig. 2. Stratigraphical column showing subdivisions, stromatoporoid occurrences,
and distribution of ‘faunas’ in the Cliefden Caves Limestone. A fault in the main section
east of the Large Flat is depicted by the break in the column. It seems to have little
significance, at most involving the loss of only a few feet of beds.

amzassensis occur sporadically through it from just below the prominent thinly bedded
unit near the stratigraphical middle, to the top of the succession. In addition, E. nestori
comes from just above and below the thinly bedded unit, L. variabi/is from just below it,
and P. inequale from immediately above it. C. cf. microundulatum and P. inequale have
also been collected from the upper part of the formation.

Further west a belt of Ordovician limestone extends from Billabong (= Goobang)
Creek (Packham 1967) to Gunningbland, and thence seems to swing eastwards towards

640 PALAEONTOLOGY, VOLUME 12

Parkes. Cliefdenella etheridgei has been collected from a locality 1 mile north of
Gunningbland.

STROMATOPOROID AND CORAL BIOSTRATIGRAPHY

In her study of the Ordovician tabulate corals of New South Wales, Hill (1957)
observed certain differences in the faunal content of the respective limestones. The
differences between the faunas of the Cliefden Caves Limestone and the Bowan Park
and Regan's Creek Limestones may be attributed to random sampling of different parts
of the respective successions, and have no biostratigraphical significance. However, the
fauna containing Plasmoporella inflata Hill and Plasmopora cargoensis Hill, from the
Cargo Creek and Canomodine Limestones, is stratigraphically distinct from the faunas
described by her from the Cliefden Caves, Bowan Park, and Regan’s Creek Limestones.

It is now possible to recognize three biostratigraphically distinct stromatoporoid and
coral faunas, here referred to as Faunas I, II, and III. These have proved to be useful for
the correlation of the limestones in the region west of Orange (text-fig. 1). Detailed work
has shown the strikingly similar sequence of faunas through each local succession. Hill’s
fauna with Plasmoporella inflata and Plasmopora cargoensis belongs to the youngest.
Fauna III.

There are, however, some anomalies in using the faunas to correlate the limestones
near Gunningbland, some 55 miles to the west, and near Tamworth, in northern New
South Wales, and this emphasizes the essentially local nature of the biostratigraphical
scheme.

In terms of Packham’s (1960; 1967) palaeogeographical reconstructions, the lime-
stones west of Orange were formed on the Molong Geanticline. To the east lay the Hill
End Trough and to the west the Cowra Trough. Further west, the limestones at Gun-
ningbland accumulated on the Parkes Platform. The faunal scheme is entirely workable
in the region of the Molong Geanticline, but beyond is less certain.

The stromatoporoid and coral faunas are as follows (text-fig. 3):

Fauna I

Diagnosis. Characterized by abundant labechiids, like L. regularis and C. donnellii , a
varied tabulate element including several species of Tetradium, and one rugose coral
(a large species of Tryplasma).

Distribution. Typically developed in lower member of Cliefden Caves Limestone; also occurs in lower
part of Regan’s Creek and Reedy Creek Limestones. Slightly modified fauna is represented in lower
part of Bowan Park Limestone, consisting of several species of Tetraclium but lacking labechiids.

Fauna II

Diagnosis. Distinguished by first appearance of clathrodictyids, particularly E. amzas-
sensis and E. nestori, by occurrence of C. etheridgei , by abundant heliolitids, and by first
appearance of Palaeophyihim. Lichenariids and tetradiids are less common, and
apparently make their last appearance in this fauna.

Distribution. Characteristic of upper part of middle member and upper member of Cliefden Caves
Limestone, and upper part of Regan’s Creek Limestone. Also occurs through middle units of Bowan
Park Limestone, in limestone lenses correlated with Reedy Creek Limestone near Eurimbula, and in
lower part of Cargo Creek and Canomodine Limestones.

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 641
Fauna III

Diagnosis. Characterized by first appearance of halysitids, favositids, solitary strepte-
lasmatids, and favistellids, particularly Favistina. Plasmoporella inflata is also diagnostic.

Distribution. Middle and upper parts of Cargo Creek and Canomodine Limestones. Similar fauna,
though lacking Favistina , found in upper part of Bowan Park Limestone. Favistina collected from
limestone breccias of Malongulli Formation overlying Cliefden Caves Limestone.

text-fig. 3. Chart showing occurrence of Faunas I— III in the Ordovician limestones of New South
Wales, and their tentative correlation with the Victorian graptolite stages.

The Bowan Park Limestone has representatives of all the faunas in superposition.
Though it lacks the variety of fauna and lithology in the lower part, there can be little
doubt that it is at least in part equivalent to the lower member of the Cliefden Caves
Limestone. Fauna II is well represented in the middle thinly bedded unit and through
the more massive beds just above and below this unit. Fauna 1 1 1 occurs in the upper part
of the limestone, though Favistina is lacking. Its absence may be due to local environ-
mental differences rather than implying that the uppermost beds are not developed, for
the same species of Halysites occurs in the uppermost beds of both the Canomodine
and Bowan Park Limestones.

Fauna I is well represented in the lower member, and Fauna II in the upper part of
the middle member and upper member of the Cliefden Caves Limestone (text-fig. 2).
Conodonts studied by Dr. G. H. Packham (pers. comm.) from the lower member sug-
gest an upper Porterfield? or lower Wilderness age (text-fig. 4).

The overlying Malongulli Formation is interpreted as a deeper-water graptolitic shale
succession with lenses of limestone breccia containing Favistina , possibly derived by
slumping from neighbouring shallow-water carbonate banks. The species of Favistina
seems to be conspecific with the form found in the upper part of the Cargo Creek and
Canomodine Limestones. Apparently the slumps from fringing carbonate banks took
place during or immediately after the accumulation of Fauna III. The present distance
between the Cliefden Caves Limestone and the Cargo Creek and Canomodine Lime-
stones is less than 8 miles, but at least one great north-south trending thrust fault, the

642

PALAEONTOLOGY, VOLUME 12

Columbine Mountain fault (Stevens 1950) separates the two areas, and they may have
been much further apart prior to folding and thrusting.

Graptolites from the Malongulli Formation have been determined by Sherrard (1954)
as belonging to the Nemagraptus gracilis Zone. Moors (1966) restudied this fauna and
concluded that it exhibited an Eastonian aspect, possibly of the Dicranograptus hians
Zone. The underlying Cliefden Caves Limestone may therefore range from the Gis-
bornian to the Eastonian.

EUROPE

AUSTRALIA

NORTH AMERICA

cn

LU

Richmond

CL

=>

Maysville

Cincinnatian

Eden

Barneveld

'Trentonian'

LU

_l

a

Q

Wilderness

'Blackriveran'

i

Porterfield

Ashby

Chazyan

Marmor

Whiterock

Ashgil I

Caradoc

Llandei lo

Llanvirn

Bolindian

Eastonian

Gisbornian

Darriwilian

text-fig. 4. Tentative correlation table for the middle and upper parts of the Ordovician
of Europe, Australia, and North America. Based on Cooper (1956), Thomas (1960),
Whittington and Williams (1964), and Whittington (1966). The old subdivisions in the
North American succession are shown in quotation marks.

Packham (1967) recorded a varied fauna, including Tetradium and heliolitids, from
Billabong Creek. He tentatively correlated it with the upper part of the Wilderness
(= Upper Gisbornian, in terms of the Victorian graptolite succession). Near Gunning-
bland, Cliefdenella etheridgei, Streptelasma sp., Palaeophyl/um sp., and Plasmoporella
injiatci have been collected together. The species of Palaeophyllum appears to be con-
specific with the form occurring in the upper member of the Cliefden Caves Limestone.
The fauna thus includes diagnostic elements of both Faunas II and III. Possibly Strepte-
lasma and P. inflata appear earlier in the limestones of the Parkes Platform than the
limestones to the east, on the Molong Geanticline. The intervening Cowra Trough, or
some marked differences in the environments of the Parkes Platform and Molong
Geanticline, may have affected the faunal distribution.

Philip (1966) described an isolated occurrence of Ordovician limestone south-east of
Tamworth, northern New South Wales. Named the Trelawney Beds, they have pro-
duced an abundant fauna, including Favistina, Palaeophyllum, Streptelasma, and helio-

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 643

litids. The coral fauna, though apparently lacking halysitids and favositids, resembles
that found in the upper part of the Cargo Creek and Canomodine Limestones, and seems
to belong to Fauna III. Philip observed that the conodonts included forms previously
described from strata of Barneveld to Maysville age in North America. He suggested
correlation of the Trelawney Beds with the upper Caradoc, or with the Eastonian of
the Victorian type succession (text-fig. 4).

Thus, Faunas I and II are tentatively regarded as Lower? or Upper Gisbornian and
Upper Gisbornian? or Lower Eastonian, respectively, and Fauna III is Upper Eastonian
(text-fig. 3). There is no evidence at present to suggest that Fauna III extends into the
Bolindian, though the possibility cannot be entirely ruled out.

ZOOGEOGRAPHICAL RELATIONSHIPS

Little is known about the stromatoporoids from the Gordon Limestone of Tasmania,
and no systematic work has been published on the fauna. Banks (1962, pp. 164, 173-4)
reported Cryptophragmus and ‘aulacerid hydrozoans’ from the Chudleigh-Mole Creek
area, and the Queenstown area. The ‘aulacerids’ near Chudleigh occur with Catenipora,
Tryplasma, and other fossils in the upper beds of the limestone. Several other localities
are mentioned by Banks (1957, p. 50) as having stromatoporoids.

In New South Wales, the stromatoporoids are dominantly encrusting laminar and
hemispherical forms. The only records of a cylindrical form are Cryptophragmus ? sp.,
from the middle part of the Regan’s Creek Limestone and from the lower part of the
Bowan Park Limestone. The significance of the specific references to Tasmanian forms
with cylindrical coenostea remains in doubt. In the departmental collection at Sydney
there are laminar-hemispherical specimens of Rosenella and Labechia from the Gordon
Limestone. The virtual lack of post-Eastonian limestones in New South Wales may
explain the absence of Aulacera. In North America species of Aulacera are limited to
the Richmond (Galloway and St. Jean 1961, p. 25), which would correlate with Upper
Bolindian in the Victorian sequence (text-fig. 4).

The stromatoporoid faunas of New South Wales exhibit the closest relationships to
Asian and east European faunas described by Yabe and Sugiyama (1930), Ozaki (1938),
Yavorsky (1955), Khalfina (1960), and Nestor (1964). Six of the fifteen described species
bear resemblances to Asian, and two to east European forms. Four of these, Rosenella
woyuensis, Labechia regularis, L. variabilis, and Ecclimadictyon amzassensis are synony-
mous with Asian species, two others exhibit affinities to Pseudostylodictyon poshanense
(Asian) and Clathrodictyon mammillatum (east European) respectively, and one is com-
parable to C. microundulatum (east European). One further species, P. inequale , is closely
related to a Chinese species described by Ozaki (1938, p. 216) as Rosenella ? sp. nov.
Only the genera Stratodictyon and Cliefdenella appear to be endemic to Australia.

Cystistroma donnellii shows some similarities to Stromatocerium canadense Nicholson
and Murie from North America (Galloway and St. Jean 1961, p. 62), but none of the
other New South Wales forms is closely related to a North American species. The only
positively assigned genera common to the Ordovician of North America (Galloway and
St. Jean 1961) and New South Wales are Cystostroma, Rosenella , and Labechia.

The clathrodictyid genus Ecclimadictyon, which makes its appearance in Fauna 11
(Lower Eastonian = middle Caradoc), first occurs in south-west Siberia in horizons

644

PALAEONTOLOGY, VOLUME 12

correlated with the upper Caradoc, and in Estonia in the Porkuni stage (upper Ashgill).
C/athrodictyon, also found in Fauna II, first appears in Estonia in the Vormsi stage
(lower-middle Ashgill). Ordovician clathrodictyids have not been reported outside the
Australian-Siberian-Estonian region. Vlasov (1961) described a species of Clathro-
dictyon , C. formozavae, from the Cambrian of West Sayan, south-west Siberia; Nestor
(1966) thought it resembled the internal structure of Ecclimadictyon, but it remains too
incompletely known for its precise affinities to be settled. Other Cambrian species
formerly regarded as clathrodictyids are now assigned to the family Korovinellidae
Khalfina 1960.

The trilobites also exhibit strong Asian affinities in the Upper Ordovician (Caradoc).
Whittington’s (1966) ‘ Encrinurella ’ fauna is considered to be restricted mainly to
Australia and south-east Asia (particularly Burma, South China, and Korea), though
Dean (1967, p. 33) noted that it may reach into eastern Turkey in the late Caradoc and
overlap with a fauna of the ’trinucleid-homalonotid’ group, having a Bohemian aspect.
The ‘ Encrinurella ’ fauna is reported as occurring in the lower member of the Cliefden
Caves Limestone and in theMalongulli Formation of New South Wales (Stevens 1952),
and in the Gordon Limestone of Tasmania (Banks 1962).

Packham (1967) observed that most of the macrofossil genera in the Ordovician fauna
at Billabong Creek are found in the Ordovician of North America, but P/iomerina
is a notable exception, occurring in south-east and central Asia.

Relatively strong faunal linkages between New South Wales and North America are
suggested by some other Ordovician groups, notably polyzoans (Phillips Ross 1961),
conodonts (Philip 1966; Packham 1967), and edrioblastoids (Webby 1968).

SYSTEMATIC PALAEONTOLOGY

The registration numbers of specimens in the University of Sydney palaeontological collections
have the prefix SUP.

Order stromatoporoidea
Family labechiidae Nicholson 1885
Genus cystostroma Galloway and St. Jean 1957

Type species. C. vermontense Galloway and St. Jean 1957.

Cystostroma cliefdenense sp. nov.

Plate 117, figs. 1-5

Material. 8 specimens from upper part of middle member of Cliefden Caves Limestone, Licking Hole
Creek (SUP 28258-60), from eastern side of Large Flat (SUP 28261-4), and from middle part of
Regan's Creek Limestone (SUP 28162).

Holotype. SUP 28258; other specimens designated paratypes.

Description. Coenosteum mainly laminar, but sometimes laminar-hemispherical in
form ; up to 1 40 x 60 mm. diameter, 50 mm. high. Broad, updomed mamelons may occur,
about 15-20 mm. in diameter. Latilaminae developed in some specimens, from 1-5 to
5-5 mm. high.

Vertical section shows small cysts of variable size, chiefly with length slightly more
than twice height; cysts usually 0-25-1 -0 mm. long, 0-12-0-20 mm. high though

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 645

exceptionally in mamelons, may be 2 mm. long and 1 mm. high. Between 12 and 22 cysts
occur in 2 mm. vertically. Cyst wall mainly 1 5-20 p thick. Latilaminae separated by mud
and calcite infilling. No astrorhizae or villi visible. In tangential section, cysts exhibit
round to polygonal outline and variable size.

Remarks. C. c/iefdenense, although not particularly well preserved, is clearly distinguish-
able from other known species of Cystostroma. It resembles both C. minimum (Parks
1910) from the ‘Trentonian’ of Kentucky, and C. fritzae Galloway and St. Jean 1961
from the Richmond of Ontario, but differs in exhibiting, on average, smaller cysts.
C. minimum has astrorhizae and C. fritzae smaller mamelons.

Genus pseudostylodictyon Ozaki 1938
Type species. P. poshanense Ozaki 1938.

Discussion. Galloway (1957, p. 424) interpreted the structures on the cysts of Pseudo-
stylodictyon as crenulations rather than denticles. This may be true of some species, but
in the original description of the type species, Ozaki (1938, p. 209) referred to short
pillar-like structures, often thicker than cysts, and not extending beyond one inter-
laminar space, or more frequently ‘mere projections’ from the underlying cysts, sug-
gesting denticles.

Pseudostylodictyon is similar to Rosenella but differs in exhibiting more gently arched
to flat or sagging, low, elongate cysts, which sometimes approximate to laminae. It
may be distinguished from Cystostroma by having less arcuate cysts and denticles.
Aulacera Plummer 1843 has a cylindrical coenosteum with large axial cysts, smaller
arcuate cysts in medial and outer zones, and sporadically developed pillars in the outer
zone (Galloway and St. Jean 1961, p. 21).

Pseudostylodictyon aff. poshanense Ozaki 1938
Plate 1 17, fig. 6; Plate 1 18, figs. 1-3

Material. 5 specimens (SUP 26226; 26232-5) from ‘mixed fauna’ unit of lower member of Cliefden
Caves Limestone, west of shearing shed, Boonderoo.

Description. Coenosteum hemispherical to encrusting, up to 170 mm. diameter, 160 mm.
high. Latilaminae exhibited at irregular intervals through coenosteum. No astrorhizae.

Vertical section shows prominent mamelons with arching of latilaminae across them.
Preservation of finer elements in coenosteum is not complete, especially within mame-
lons. Cysts of variable size, updomed at mamelons and broadly sagging between; cysts
long, low, and gently convex away from mamelons. Denticles occur on upper surfaces
of cysts and rarely on outer surfaces of mamelons. Mamelons usually 1-2 mm. wide,
spaced 3-8 mm. apart. Number of cysts varies from 4 to 14 in 2 mm. vertically; occa-
sional large cysts 1-8-8 mm. long, 0-5-2 mm. high. Thickness of cyst wall varies from
10 to 40 p; mainly about 20 p. In tangential section, mamelons are round; denticles
show as fine specks.

Remarks. P. aff. poshanense ranges from a finer variety (PI. 118, figs. 1, 2) to coarser
varieties (PI. 117, fig. 6; PI. 118, fig. 3). It has affinities with P. poshanense from the
Ordovician of Shantung (Ozaki 1938, p. 208), but is distinguished by narrower and more

646

PALAEONTOLOGY, VOLUME 12

erect mamelons. P. aft', poshanense is not closely comparable with any of the North
American species of Pseudostylodictyon ? described by Galloway and St. Jean (1961).

Pseudostylodictyon inequale sp. nov.

Plate 119, figs. 1-3

Material. 3 specimens (SUP 28252-4) from upper part of middle member of Cliefden Caves Lime-
stone, Licking Hole Creek; 1 specimen (SUP 29141) from just above middle thinly bedded unit of
Bowan Park Limestone, Quondong; 1 specimen (SUP 29134) from upper part of limestone at Malachi’s
Hill.

Holotype. SUP 28252; other specimens designated paratypes.

Description. Coenosteum hemispherical to laminar, up to 90 mm. diameter, 85 mm.
high. In vertical section, cysts very variable in size, from large gently convex plates up
to 20 mm. long and 4 mm. high, to long, low cysts resembling laminae, spaced about
12-16 in 2 mm. vertically. Long, low cysts occur in groups arranged parallel to one
another, lying between scattered larger cysts. Cyst wall usually about 20-25 p thick, but
exceptionally 120^. Denticles prominent on upper surface of cysts in some areas;
especially conspicuous where they project into sediment infilling of a large overlying
cyst; seen to extend 0-2 mm. above upper surface of cyst. Mamelons not clearly differen-
tiated. In tangential section denticles appear as dots spaced about 0-1 mm. apart;
diameter 20-60 p.

Remarks. P. inequale bears close similarities to a stromatoporoid from the Middle?
Ordovician of Shantung described by Ozaki (1938, p. 216) as Rosenellal sp. nov.
Galloway (1957, p. 424) assigned Ozaki’s species to Pseudostylodictyon, evidently
because it exhibited the frequent grouping of parallel rows of long, low cysts simulating
laminae. The Shantung species is closely related to P. inequale but differs in having
mamelons.

EXPLANATION OF PLATE 117

Figs. 1-4. Cystostroma cliefdenense sp. nov., X 10, middle member, Cliefden Caves Limestone, Licking
Hole Creek. 1, SUP 28259, paratype, vertical section showing rhythmic alternation of latilaminae
and calcite with sediment-filled zones. 2-4, SUP 28258, holotype; 2, vertical section exhibiting
typical arrangement of cysts in coenosteum; 3, vertical section showing mamelon with large asso-
ciated cysts; 4, tangential section.

Fig. 5. Cystostroma cliefdenense sp. nov., SUP 28162, paratype, X 10, vertical section; middle part of
Regan’s Creek Limestone.

Fig. 6. Pseudostylodictyon aff. poshanense Ozaki, SUP 26235, x 5, vertical section exhibiting conical-
shaped mamelons and denticles on upper surface of the large cysts. From ‘mixed fauna’ unit, lower
member, Cliefden Caves Limestone, west of shearing shed, Boonderoo.

EXPLANATION OF PLATE 118

All figures from ‘mixed fauna' unit, lower member, Cliefden Caves Limestone, west of shearing shed,

Boonderoo.

Figs. 1-3. Pseudostylodictyon aff. poshanense Ozaki. 1, 2, SUP 26226, x 5; 1, vertical section showing
small and large cysts; 2, tangential section. 3, SUP 26234, x 5, vertical section.

Figs. 4-6. Stratodictyon columnare sp. nov., SUP 26229, holotype. 4, vertical section showing lati-
laminae and mamelons, X 5. 5, vertical section showing rows of fine, long, low cysts and poorly
developed short, denticle-like pillars, x 10. 6, tangential section exhibiting arched rows of cysts
around mamelons and dark specks which represent small pillars, x 10.

Palaeontology, Vol. 12

PLATE 117

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 118

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROMATOPOROIDS FROM NEW SOUTH WALES 647

Genus stratodictyon gen. nov.

Type species. S. ozakii sp. nov.

Diagnosis. Encrusting, laminar to hemispherical coenosteum, with latilaminae, rela-
tively long, low cysts in regular rows resembling laminae, and small, discontinuous
pillars. Mamelons may be present or absent. Scattered astrorhizae also occur.

Discussion. The short pillars extending vertically across up to 10 cysts and the presence
of scattered astro rhizal canals distinguish StratodictyonixomPseudostylodictyon.Aulacera
has small pillars in the outer zone of its cylindrical coenosteum, but the columnar form
is taken by Galloway (1957, p. 423) to be one of the fundamentally diagnostic features
of the genus. It is uncertain whether Aulacera exhibited astrorhizae or not. Galloway
(1957, p. 422) reported them to be rare or absent, but Galloway and St. Jean (1961,
p. 21) stated ‘astrorhizae absent’.

Plumatalinia Nestor 1960 also seems to be a related genus, but the columns are formed
of fine subreticulate tissue, and astrorhizae are absent (Nestor 1964). In the mamelon
columns of Stratodictyon columnare sp. nov., the rows of cysts are updomed and the
pillars diverge outwards.

Stratodictyon ozakii sp. nov.

Plate 1 19, figs. 4-5; Plate 120, figs. 1,2; Plate 124, fig. 1

Material. Based on 4 specimens (SUP 26247-8, 26252-3) from ‘lower coral’ unit of lower member of
Cliefden Caves Limestone, Licking Hole Creek.

Holotype. SUP 26252; other specimens designated paratypes.

Description. Encrusting to laminar-hemispherical coenosteum, 130x100 mm. across,
65 mm. high. Latilaminae prominent, even to gently undulating, usually 1-10 mm. high.
Encrusts Nyctopora, Labechia, Cystistroma, and a polyzoan resembling Prasopora.

In vertical section rows of long, low cysts approximating to laminae, averaging
16-19 per 2 mm., are visible. Alternating zones of well preserved cysts and poorly
calcified elements or calcite through coenosteum. In one specimen (SUP 26248, PI.
120, fig. 1) numerous closely spaced latilaminae occur, each successively thinning out
towards crest of an encrusted Labechia , and finally completely mantling it; alga present
between two of the latilaminae. Walls of cysts mainly c. 20 p thick. Pillars thicker than
cyst walls, flocculent in appearance, 40-50 p in diameter; seen to extend continuously
across up to 7 cysts. In tangential section pillars appear as fine circular specks, mainly
spaced 50-150 p apart. Vague scattered astrorhizal canals observed, mainly c. 0-2 mm.
wide.

Remarks. S. ozakii is closely allied to S. columnare , only differing in lacking mamelons.
Aulacera peichuangensis Ozaki (1938, p. 217) from the Ordovician of Shantung is
remarkably similar in the character of the latilaminae, the long, low cysts, and the
moderately persistent pillars of the outer zone, but not in the differentiated axial and
outer zones of the cylindrical or dendroid coenosteum.

648

PALAEONTOLOGY, VOLUME 12

Stratodictyon columnare sp. nov.

Plate 118, figs. 4-6; Plate 119, fig. 6; Plate 124, fig. 3

Material. 5 specimens (SUP 26227-31) from ‘mixed fauna’ unit of lower member of Cliefden Caves
Limestone, and 1 specimen (SUP 29142) from lower part of Bowan Park Limestone, east of Quondong.

Holotype. SUP 26229 ; other specimens designated paratypes.

Description. Encrusting to hemispherical coenosteum, up to 140x100 mm. across,
90 mm. high. Mamelons prominent. Latilaminae folded across mamelons, from 1 to
15 mm. high. Encrusts Coccoseris and Labechia. Algae encrust it, and occur between
latilaminae.

Vertical section exhibits rows of long, low cysts resembling laminae, averaging
15-18 per 2 mm. Walls of cysts mainly 15-20 p thick. Pillars thicker than cyst walls, c.
50 /x in diameter; usually only height of interlaminar space, but some more continuous,
extending across up to 10 cysts. Structures not well preserved in mamelons, but in one
part of holotype (see text-fig. 5) rows of cysts updome and pillars diverge outwards.
Mamelons c. 1-2 mm. in diameter, spaced c. 3-5 mm. apart. Astrorhizal canals scattered
in coenosteum. In tangential section, cysts seem to be arranged concentrically around
mamelons. Pillars show as fine specks.

Remarks. The close relationships between 5. columnare and S. ozakii are emphasized by
one specimen (PI. 124, fig. 3) which is assigned to S. columnare. Its coenosteum lacks
mamelons near the encrusted Coccoseris but has them away from it.

EXPLANATION OF PLATE 119

Figs. 1-3. Pseudostylodictyon inequale sp. nov. 1, 2, SUP 28252, X 10, holotype, upper part of
middle member, Cliefden Caves Limestone, Licking Hole Creek. 1, vertical section showing alter-
nations of rows of long, low cysts simulating laminae and large cysts, and prominent denticles.
2, tangential section shows denticles as dark dots. 3, SUP 29134, X 5, paratype, vertical section,
upper part of Bowan Park Limestone, Malachi’s Hill.

Figs. 4, 5. Stratodictyon ozakii gen. et sp. nov., SUP 26252, x 10, holotype, ‘lower coral’ unit, lower
member, Cliefden Caves Limestone, Licking Hole Creek (see also PI. 124, fig. 1). 4, vertical section
showing rows of long, low cysts with scattered short pillars. 5, tangential section exhibiting specks
which seem to represent pillars.

Fig. 6. Stratodictyon columnare sp. nov., SUP 26227, X 10, paratype, vertical section showing small
pillars crossing several rows of cysts. ‘Mixed fauna' unit, lower member, Cliefden Caves Limestone,
west of shearing shed, Boonderoo.

EXPLANATION OF PLATE 120

Figs. 1,2. Stratodictyon ozakii gen. et sp. nov., SUP 26248, paratype, ‘ lower coral ’ unit, lower member,
Cliefden Caves Limestone, Licking Hole Creek. 1, vertical section, x 5, showing latilaminae suc-
cessively thinning towards crest of encrusted specimen of Labechia regularis. Darker patch between
latilaminae represents a small algal growth. Rows of long, low cysts and small pillars well exhibited
in upper parts of coenosteum. 2, tangential section, X 10, exhibiting vague astrorhizal canals in
addition to pillars.

Figs. 3-6. Rosenella woyuensis Ozaki, ‘mixed fauna’ unit, lower member, Cliefden Caves Limestone,
east of Large Flat. 3, SUP 26216, x 5, vertical section showing rows of large cysts alternating with
sediment-filled spaces. 4, SUP 26217, x5, tangential section showing large mamelon surrounded
by cysts. 5, SUP 26216, X 10, vertical section showing denticles on upper surfaces of cysts. 6,
SUP 26216, x5, vertical section showing large cysts, including some infilled with sediment and
algal? structures.

Palaeontology, Vol. 12

PLATE 119

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 120

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 649

Genus rosenella Nicholson 1886
Type species. R. macrocystis Nicholson 1886.

Rosenella woyuensis Ozaki 1938
Plate 120, figs. 3-6

1938 Rosenella woyuensis Ozaki, p. 215, pi. 30, fig. 2; pi. 31, fig. 1 a-d.

Material. 4 specimens (SUP 26214—1 7) from ‘mixed fauna’ unit and 1 specimen (SUP 29140) from
‘upper big shell’ unit of lower member of Cliefden Caves Limestone, east of Large Flat.

Description. Laminar to hemispherical coenosteum composed of large convex cysts;
reaches 150 mm. across, 90 mm. high. Latilaminae with sediment-filled spaces at
irregular intervals between them. Broad mamelons developed.

Vertical section shows irregular-sized larger and smaller cysts with denticles arising
from upper surfaces. Wall of cysts varies considerably in thickness, usually from 25 to
1 50 /x, exceptionally up to 500 p. Cysts usually 1-3 mm. high, 4-10 mm. long, excep-
tionally up to 6 mm. high, 24 mm. long. Cysts spaced about 5 in 4 mm. vertically. Some
cysts contain calcite infilling, others filled with sediment which seems to include a variety
of algal? structures. In tangential section, broad mamelons prominent, with large
curved cysts surrounding them. No astrorhizae.

Remarks. The species closely resembles R. woyuensis from the Tsinan Limestone
(Middle? Ordovician) of Po-shan-hsien, province of Shantung, China (Ozaki 1938), and
is therefore assigned to it. R. amzassensis Khalfina from the Upper Ordovician of the
Siberian Platform (Khalfina 1960) is similar, but one of its type specimens exhibits three
pillars.

Genus labechia Milne-Edwards and Haime 1851
Type species. Monticularia conferta Lonsdale 1839.

Labechia regular is Yabe and Sugiyama 1930

Plate 120, fig. 1 ; Plate 121, figs. 3-6; Plate 124, figs. 1, 2

1930 Labechia regularis Yabe and Sugiyama, p. 56, pi. 18, figs. 5, 6; pi. 21, fig. 8.

1938 Labechia regularis Ozaki, p. 210, pi. 26, fig. 2 a-d.

1955 Labechia regularis Yavorsky, p. 59, pi. 24, figs. 4, 5.

Material. 13 specimens from ‘lower coral’ unit. Licking Hole Creek (SUP 26236, 26238-9, 26242,
26248, 26252, 26255), and from ‘mixed fauna’ unit, west of shearing shed, Boonderoo (SUP 26231,
26240, 26243-4) and east of Large Flat (SUP 26237, 26245). Also occurs in lower part of Regan's
Creek Limestone, south-east of Cargo, and in lower part of Reedy Creek Limestone between Molong
and Copper Hill.

Description. Coenosteum hemispherical to laminar-encrusting, up to 100 mm. diameter,
80 mm. high. Latilaminae mainly from 3 to 15 mm. high. No astrorhizae.

Vertical section shows moderately thick, persistent pillars, 0-2-0-3 mm. in diameter;
some exhibit lighter axial zones and some have zigzag edges, being broader just above
point where it intersects lamina or dissepiment. In specimen SUP 26236, the laminae
seem to persist as updomed extensions across pillars, and a pillar shows multiple
branching (PI. 121, fig. 6). At the upper boundary of a latilamina, pillars often appear to

C 6940

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650

PALAEONTOLOGY, VOLUME 12

extend up as rounded tubercles above level of adjacent lamina (PI. 121, fig. 3). Laminae
regularly spaced, flat to slightly concave, continuous between pillars over distances of 50
mm. or more in several examples; also smaller upcurved cysts (dissepiments) developed
between laminae in certain areas, from 0-2 to 1 mm. wide, 0-1 -0-3 mm. high. Wall
thickness of dissepiments same as for laminae, from 25 to 50 p thick. Laminae mainly
spaced from 9 to 12 in 2 mm. vertically. Pillars usually between 0-1 and 0-5 mm. apart.

In tangential section, some pillars exhibit light-coloured centres suggesting that they
were originally hollow, or originally infilled with different material from outer walls;
others appear to be undifferentiated. Pillars mainly circular, but in some sections irregular
to subangular in outline (PI. 121, fig. 4).

Remarks. The New South Wales material particularly closely resembles L. regularis, as
described by Yavorsky (1955) from Upper Ordovician localities in the basin of the Stony
Tunguska River and a tributary of the Kotuy River, Siberian Platform. The Russian
specimens appear to come from the Dolborsk stage, which is correlated with the upper
Caradoc by Ivanovsky (1965). The type material described by Yabe and Sugiyama (1930)
from Wu-hu-tzui, Fu-hsien, province of Liaotung, South Manchuria, probably from
the Toufangkou Limestone of Middle? Ordovician age, seems to have fewer updomed
cysts (dissepiments) but is otherwise similar.

L. ( Labeehiella ) mingshankouensis (Ozaki) from the Ordovician of South Manchuria
and Shantung also has regular laminae, but they are more widely spaced (4 to 5 in
2 mm.), and astrorhizae are exhibited (Ozaki 1938, p. 207).

Labechia variabi/is Yabe and Sugiyama 1930

Plate 121, figs. 1, 2

1930 Labechia variabilis Yabe and Sugiyama, p. 54, pi. 17, figs. 1-9.

1938 Labechia variabilis Ozaki, p. 211, pi. 28, fig. 1 a-d.

Material. 3 specimens (SUP 28163-4, 28248) from top of middle, massively bedded part of Regan’s
Creek Limestone, and 1 specimen (SUP 28251) from upper part of middle member of Cliefden
Caves Limestone, on banks of Belubula River, west of Large Flat. Also occurs just below middle,
thinly bedded unit of Bowan Park Limestone east of The Ranch.

Description. Coenosteum hemispherical, reaching 90x60 mm. across, 80 mm. high.
One specimen (SUP 28251) encrusts Tetradium cribriforme (Etheridge), and is associated
with heliolitids.

In vertical section, laminae gently undulating, concave upwards between closely
spaced pillars to flat and gently convex. Laminae spaced about 4-6 in 2 mm. vertically;

EXPLANATION OF PLATE 121

Figs. 1, 2. Labechia variabilis Yabe and Sugiyama, x 5. 1, SUP 28248, vertical section; top of middle
part of Regan's Creek Limestone. 2, SUP 28251, tangential section; upper part of middle member,
Cliefden Caves Limestone, on south bank of Belubula River, west of Large Flat.

Figs. 3-6. Labechia regularis Yabe and Sugiyama. 3, 4, SUP 26240, x5; ‘mixed fauna’ unit, lower
member, Cliefden Caves Limestone, west of shearing shed, Boonderoo; 3, vertical section showing
well-developed latilaminae; 4, tangential section exhibiting pillars with lighter centres, and irregular
to subangular outlines. 5, 6, ‘lower coral’ unit, lower member, Cliefden Caves Limestone, Licking
Flole Creek; 5, SUP 26242, X 5, tangential section; 6, SUP 26236, x 10, vertical section showing
branching pillar.

Palaeontology, Vol. 12

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WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 651

in few instances may be traced horizontally as continuous elements for more than
15 mm. Pillars appear to arise from more continuous laminae; persistent, extending to
height of at least 8 mm., and cone-shaped across interlaminar spaces; branching pillar
also observed (SUP 28251).

Tangential section shows great variability in diameter of pillars, from 0-1 to 0-5 mm.
owing to the cone-shaped form; pillars spaced from 0-4 to TO mm. apart, circular, oval
to subangular in outline; some appear to have hollow centres. Irregular tube-like struc-
tures about 0-3 mm. wide may represent astrorhizae, but they cannot be confirmed in
vertical section.

Remarks. The New South Wales specimens are closely comparable with L. variabilis
from the Asian Ordovician, especially with the material described by Ozaki (1938) from
South Manchuria. The only possible difference is the doubtful astrorhizae exhibited by
one New South Wales specimen. The original description of L. variabilis by Yabe and
Sugiyama (1930) is based on a number of specimens (syntypes) from various localities
in South Manchuria, North China, and Korea, and needs revision.

Genus cryptophragmus Raymond 1914
Type species. C. antiquatus Raymond 1914

Cryptophragmus ? sp.

Plate 122, figs. 1, 2

Material. 2 silicified specimens (SUP 28169, 28171) from middle part of Regan’s Creek Limestone.
Also occurs in lower part of Bowan Park Limestone, Paling Yard Creek.

Description. Coenosteum cylindrical, unbranched, more than 40 mm. in length, 13-
15 mm. in diameter; encrusted by colony of Propora. Axial column 7-10 mm. in diameter;
axial cysts unknown. Latilaminae present, though not prominent. No astrorhizae or
mamelons. Lateral cysts spaced 8-10 per 2 mm. radially; flat to concave between pillars;
seem to be slightly oblique to the outer wall rather than parallel to it. Pillars have long,
blade-like form; in silicified material, individual pillars can be traced as longitudinal
grooves on outer surface of coenosteum for up to 7 mm.; spaced about 0-5-1 mm. apart
around periphery of coenosteum.

Remarks. In the type species of Cryptophragmus , C. antiquatum Raymond, from the
‘Blackriveran’ of eastern North America, the pillars are described as ‘oval to prismatic
and tend to be round’ (Galloway and St. Jean 1961, p. 19). The New South Wales speci-
mens, in contrast, have blade-like pillars. They are therefore only tentatively assigned
to the genus.

Genus cystistroma Etheridge 1895
Type species. C. donnellii Etheridge 1895.

Discussion. The original type material of Labechial ( Cystistroma ) donnellii Etheridge is
being redescribed by Dr. J. Pickett (Geological and Mining Museum, Geological Survey
of New South Wales). Judging from the nature of preservation of the material it was
probably collected originally from the Tower coral’ unit in the Fossil Hill section. This
locality is near the Belubula River, in the Parish of Malongulli, as stated by Etheridge

652

PALAEONTOLOGY, VOLUME 12

(1895, p. 134). Pickett is raising the type species of Cystistroma to full generic rank, a
procedure which I support.

Etheridge (1895) originally discussed the relationships between Cystistroma and
Labechia , Rosenella and Beatricea Billings 1857 ( = Aulacera), but not Stromatocerium
Hall 1847. He observed (p. 139) that ‘ Cystistroma appears to be a Labechia assuming
certain Rosene/la-Yike features’.

Cystistroma is closely allied to Rosenella , but is distinguished by having large pillars.
It differs from Labechia and Stromatocerium in exhibiting denticles on the upper surface
of cysts and sometimes on the outer surface of pillars. Labechia has regular, cylindrical
pillars, whereas Stromatocerium has irregular to blade-like pillars. The variability of
pillars in Cystistroma typically extends to both Labechia and Stromatocerium types. Only
those with serrated outlines, owing to the intersection of denticles on outer surfaces of
pillars, are distinct.

Denticles have only been positively identified on the upper surface of cysts in one
species of Stromatocerium , namely. S', canadense Nicholson and Murie from the ‘ Black-
riveran’ and lower ‘Trentonian’ of North America (Galloway and St. Jean 1961, p. 62).
Nestor (1964) recorded ‘small monticles’ on the upper surface of cysts and ‘hollow’
pillars in Estonian examples of S. canadense from the Oandu and Pirgu stages (= middle
Caradoc and middle Ashgill, respectively), and S. sakuense Nestor from the Oandu
stage. Possibly these features result from incomplete preservation, and the ‘monticles’
are poorly preserved denticles. Perhaps these species should be assigned to Cystistroma.

Cystistroma donnellii Etheridge 1 895

Plate 122, figs. 3-8; Plate 123, figs. 1-5; Plate 124, fig. 2

1895 Labechia (?) {Cystistroma) donnellii Etheridge, p. 134, pi. 14, figs. 1-6; pi. 15, figs. 1,2; pi.

16, figs. 1-3.

Material. Based on specimens from following localities and horizons in lower member of Cliefden

EXPLANATION OF PLATE 122

Figs. 1, 2. Cryptophragmusl sp„ SUP 28169, x4, middle part of Regan's Creek Limestone. 1, cross-
section showing outer zone of cysts and pillars in silicified specimens. 2, oblique section showing
linear grooves representing impressions of blade-like pillars on outer surface of specimen.

Figs. 3-8. Cystistroma donnellii Etheridge, x 5. 3, 4, SUP 26263, ‘mixed fauna’ unit, lower member,
Cliefden Caves Limestone, west of shearing shed, Boonderoo. 3, vertical section with characters
resembling P. aff. poshanense. 4, tangential section showing pillars with serrated outline owing to
denticles on outer surfaces. 5, SUP 26258, vertical section showing denticles on outer surface of
pillars; ‘mixed fauna’ unit, lower member of Cliefden Caves Limestone, east of Large Flat. 6, SUP
26259, tangential section; ‘mixed fauna’ unit, lower member, Cliefden Caves Limestone, near
Cliefden Caves. 7, SUP 26250, assumed topotype, vertical section ; ‘lower coral ’ unit, lower member,
Cliefden Caves Limestone, Fossil Hill. Typical representative of C. donnellii var. A (see also PI. 123,
fig. 1). 8, SUP 26262, vertical section; lower part of Reedy Creek Limestone, just south of Molong.

EXPLANATION OF PLATE 123

Figs. 1-5. Cystistroma donnellii Etheridge, X 5, assume! topotypes, ‘lower coral’ unit, lower member,
Cliefden Caves Limestone, Fossil Hill. 1, SUP 26250, tangential section showing pillars with an
oval outline. 2, 3, SUP 26268, vertical and tangential sections showing irregular, angular pillars.
4, 5, SUP 26267, vertical and tangential sections exhibiting angular, blade-like pillars. Figs. 2-5
depict representatives of C. donnellii var. B.

Palaeontology , Vol. 12

PLATE 122

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 123

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 653

Caves Limestone: 'lower coral’ unit, Fossil Hill (assumed topotypes SUP 26250, 26267-8, 28246-7),
shearer’s quarters, Boonderoo (SUP 26260-1), Licking Hole Creek (SUP 26255), 'mixed fauna’ unit,
west of shearing shed, Boonderoo (SUP 26263), east of Large Flat (SUP 26257-8), east-south-east of
Cliefden Caves (SUP 26259). Also found in lower part of Reedy Creek Limestone, near Molong
(SUP 26262), and in lower part of Regan’s Creek Limestone, south-east of Cargo.

Description. Large hemispherical to encrusting coenosteum, up to 280 mm. diameter,
200 mm. high. Vertical section shows thick persistent pillars varying from 0-3 to 2 mm.
(but mainly c. 1 mm.) in diameter. Cysts updomed at contacts with pillars and sagging
between; where not intersected by pillars, tend to be convex to undulating plates of
variable size; usually spaced from 2 to 4 in 2 mm. vertically, but exceptionally up to
3 mm. apart. Wall thickness varies from 50 to 250 p. Upper surface of cysts and outer
surface of some pillars exhibit denticles.

In tangential section, pillars round to elongate, irregular; some round pillars have
serrated outline where denticles are intersected, and some irregular pillars have narrow
flanges. Pillars mainly composed of narrow, dense outer zone, and broad, granular axial
zone; former seems to be in continuity with adjacent cysts. Pillars spaced about 2-3 mm.
apart. No astrorhizae seen.

Remarks. The two morphological variants of C. donnellii, depicted in text-fig. 5, occur
together in the same beds of the ‘lower coral’ unit at Fossil Hill. C. donnellii var. A
exhibits large round to oval pillars (PI. 122, fig. 7; PI. 123, fig. 1), whereas C. donnellii
var. B shows large angular, blade-like pillars (PI. 123, figs. 2-5).

C. donnellii most closely resembles S. canadense, which is noted for exhibiting an
extreme of variability (Parks 1910, pp. 16-20). However, C. donnellii is on the whole a
much larger form.

Relationships between Stratodictyon, Pseudostylodictyon and Cystistroma. Strato-
dictyon ozakii, S. columnare, P. aff. poshanense, and C. donnellii from the lower member
of the Cliefden Caves Limestone exhibit a remarkably continuous range of morpho-
logical variation (text-fig. 5). S. ozakii resembles S. columnare except for the mamelons;
S. columnare is similar to P. aff. poshanense except that the small pillars are denticles
and the cysts are larger; and P. aff. poshanense is close to C. donnellii var. A, but the
mamelons are replaced by large pillars. As already noted, C. donnellii var. A has round
pillars and var. B irregular, blade-like pillars.

The small pillars of S. ozakii and S. columnare have much the same spacing as the
denticles in P. aff. poshanense and C. donnellii , and seem to have developed as extensions
of them. They are not seen to be continuous across more than 10 cysts. Galloway (1957,
p. 368) indicated that pillars were derived by upward growth of denticles.

Closely spaced mamelons in S. columnare and P. aff. poshanense , especially in the
latter (PL 117, fig. 6; PI. 118, fig. 3) are morphologically similar to the large, round
pillars of C. donnellii var. A (PI. 122, figs. 3, 4, 7, 8). Both structures have a similar
spacing, and have cysts updomed adjacent to them. The updomed ‘mamelon’-like pillar
structure of C. donnellii is well shown in the intergrowth with L. regularis and Prasopora
(PI. 124, fig. 2). The presence of denticles on the upper surface of cysts and the outer
surface of pillars in C. donnellii illustrates that the cyst tissue is in continuity with the
outer zone of the large pillars, and supports the view that the large pillars were derived
from mamelons. Although it may be a function of the relative size of cysts and

654

PALAEONTOLOGY, VOLUME 12

pillars, there seems to be little obvious updoming of cysts against small pillars (derived
from denticles), whereas the cysts adjacent to large pillars are appreciably updomed,
suggesting derivation from mamelons. The great size discrepancy between large and
small pillars in species occurring at similar horizons in the lower member of the Cliefden
Caves Limestone is thus explained by their different origins.

text-fig. 5. Relationships between Stratodictyon, Pseudostylodictyon and Cystistroma,
showing derivation of large pillars from mamelons, and small pillars from denticles.

Vertical and tangential sections approximately 7 times natural scale.

Regarding details of occurrences of these forms, P. aff. poshanense (5 specimens) and
S. co/umnare (5 specimens) are found together at one locality, in the ‘mixed fauna’ unit
west of the Boonderoo shearing shed. One specimen (PI. 124, fig. 3) of S. columnare
from this locality is very similar to S. ozakii. Another specimen (PI. 122, fig. 3), assigned
to C. donnellii, is closely related to P. aff. poshanense. Judging from the intermediate
forms (for example, PI. 118, fig. 1), specimens of P. aff. poshanense and S. columnare
could be interpreted as variants of one species.

The locality in the Tower coral’ unit at Licking Hole Creek yielded 4 specimens of
S. ozakii and 1 specimen of C. donnellii, but no S. columnare or P. aff. poshanense. At
Fossil Hill, also in the Tower coral’ unit, both variants of C. donnellii occur, but none
of the other species.

At each of these localities, with the possible exception of Licking Hole Creek, one
organism may have constructed the forms represented. This, however, leads to viewing

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 655

the whole range of variability from S. ozakii to C. donnellii as the product of one organ-
ism. Even if there were local differences in environmental conditions and preservation,
it is difficult to accept this proposal. Yet the possibility remains that we are here dealing
with form genera and species formed by one organism under differing local conditions,
rather than with true genera and species.

Family cliefdenellidae fam. nov.

Relationships. The family Cliefdenellidae is distinguished from the labechiids by having
coenostea with laminae which are downwardly indexed against pillars, and complex
astrorhizae. The presence or absence of astrorhizae has been considered by Galloway
(1957, p. 378) to have no more than specific importance, but it may be significant that
no other Ordovician stromatoporoid has such an elaborate canal and column system
as Cliefdenelia etheridgei gen. et sp. nov. The comparatively large size, the presence of
denticles on the upper surface of laminae (or cysts), and dissepiments are features that
the group shares with labechiids.

The cliefdenellids resemble representatives of the family Clathrodictyidae Kuhn 1939
in having downward inflexions of laminae at pillars, but differ in exhibiting persistent
solid to tube-like pillars, and denticles on the upper surface of laminae. The supposed
Cambrian stromatoporoid family Korovinellidae Khalfina 1960 is distinguished by
exhibiting slender, rod-like pillars, porous, perforated laminae, and by lacking denticles.
Nestor (1966) has maintained that the Cambrian forms are archaeocyathids con-
vergently similar to stromatoporoids, but Khalfina and Yavorsky (1967) have defended
the view that they are the oldest stromatoporoids.

Genus cliefdenella gen. nov.

Type species. C. etheridgei sp. nov.

Diagnosis. Coenosteum composed of laterally persistent laminae and continuous solid
to tube-like pillars. Laminae exhibit denticles on upper surfaces and downward in-
flexions against pillars. Domed tabulae occur in tube-like pillars. Interlaminar spaces
occupied by dissepiments and branching astrorhizal canals. Prominent astrorhizal
columns with updomed laminae and vertical ‘septal’ structures.

Discussion. Few genera show even broadly similar characteristics. The most closely
comparable labechiids, Cystistroma, Labechia, and Labechieiia Yabe and Sugiyama 1930
all differ in having updoming of cysts (or laminae) against pillars. In addition, Cysti-
stroma lacks astrorhizae, Labechia lacks denticles and astrorhizae, and Labechieiia lacks
denticles. The Ordovician species of Clathrodictyon do not typically exhibit dissepiments,
and they lack denticles and tube-like pillars. The Lower Cambrian genus Korovine/ia
Khalfina 1960 also lacks the tube-like pillars, and differs in having laminae with a reti-
culate tissue and lacking denticles.

Cliefdenella etheridgei sp. nov.

Plate 125, figs. 1-5; Plate 126, figs. 1, 2

Material. 8 specimens (SUP 24154-9, 26246, 26264) from ‘Island' unit in upper member of Cliefden
Caves Limestone, on Island, between Belubula River and Large Flat. 3 specimens (SUP 24160,

656 PALAEONTOLOGY, VOLUME 12

29138-9) from Ordovician limestones, south side of Parkes-Bogan Gate road, 1 mile north of Gun-
ningbland.

Holotype. SUP 24157; other specimens designated paratypes.

Description. Coenosteum hemispherical to laminar, up to 1 30 X 90 mm. diameter, 80 mm.
high. Regularly spaced laminae and pillars (appearing as tubes) show on broken surfaces.
Upper surface in one specimen (SUP 24155) undulatory, with updoming of laminae
around astrorhizal columns.

Laminae regularly spaced, continuous, with pronounced downward inflexions at
junctions with pillars; spaced on average 6 laminae in 5 mm. Walls of laminae from 50
to 100 p thick with sharp outer boundaries; composed of 3 layers, an inner and outer
clear zone, and central granulated zone. Smaller cyst-like dissepiments occur abun-
dantly between laminae, some obliquely inclined across laminae. Denticles present on
upper surfaces of laminae.

Pillars are well-developed cylindrical vertical elements, but variable in internal form;
some are hollow tube-like structures with updomed cysts (tabulae) crossing them, others
are infilled to form solid elements. In tangential section, circular in outline, mainly
0-5 mm. in diameter; outer wall of tube-like pillars 50-75 /x thick. Branching pillar
observed in one specimen (PI. 126, fig. 1).

Astrorhizal columns with associated updomed laminae and vertical spine-like ele-
ments are 2 mm. in diameter. Pattern of stellate branching astrorhizal canals, up to 0-5
mm. wide, ramifies through interlaminar spaces away from astrorhizal columns; astro-
rhizal canals also occasionally exhibit tabulae, spaced c. 1 mm. apart.

Remarks. "Labechia ? sp. (Gen. et sp. nov. ?)’ figured and described by Ozaki (1938,
p. 213) from Ordovician limestones of Shantung resembles C. etheridgei, but it is on the
whole smaller, having 3-4 laminae in 2 mm. and pillars 0- 1 2—0-2 1 mm. in diameter; the

EXPLANATION OF PLATE 124

Fig. 1. Association of Nyctopora sp., Labechia regularis Yabe and Sugiyama, and holotype of Strato-
dictyon ozakii gen. et sp. nov., SUP 26252, X 5. Both L. regularis and S. ozakii exhibit latilaminae.

Fig. 2. Association of Labechia regularis Yabe and Sugiyama, Cystistroma donnellii Etheridge, and
Prasopora sp., SUP 26255, X 5. Note mamelon-like pillars of C. donnellii.

Fig. 3. Intergrowth of Stratodictyon columnare sp. nov. and Coccoseris sp., SUP 26230, X 5. Note
latilaminae in S. columnare, and relationship to periods of growth in Coccoseris.

Figs. 1, 2 from ‘lower coral’ unit, lower member, Cliefden Caves Limestone, Licking Hole Creek;
Fig. 3 from ‘mixed fauna’ unit, lower member, Cliefden Caves Limestone, west of shearing shed,
Boonderoo.

EXPLANATION OF PLATE 125

Figs. 1-5. Cliefdenella etheridgei gen. et sp. nov., x 5. 1-4, from ‘Island’ unit, upper member, Cliefden
Caves Limestone, at the Island. 1, SUP 24157, holotype, vertical section showing prominent astro-
rhizal columns with vertical ‘septa '-like structures and updomed horizontal elements, tube-like
pillars, laminae with denticles and downward inflexions against pillars, dissepiments, and vague
tabulae in hollow pillars. 2, SUP 24156, paratype, oblique section showing tube-like pillars filled
by sediment. 3, 4, SUP 24154, paratype; 3, tangential section showing stellate, branching astro-
rhizal canals radiating from column; pillars are mainly solid elements, and denticles show as small
dark specks; 4, oblique section showing tabulae in astrorhizal canals, and both tube-like and solid
pillars. 5, SUP 24160, paratype, vertical section, from one mile north of Gunningbland, showing
slightly updomed tabulae in hollow pillars.

Palaeontology, Vol. 12

PLATE 124

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 125

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 657

laminae are less regular and the dissepiments are not clearly differentiated. Astrorhizae
are recorded, but nothing comparable with the complex network seen in C. etheridgei.

Family clathrodictyidae Kuhn 1939
Genus clathrodictyon Nicholson and Murie 1878

Type species. C. vesciculosum Nicholson and Murie 1878.

Clathrodictyon aff. mammillatum (Schmidt 1858)

Plate 126, figs. 3-5

Material. 1 specimen (SUP 28255) from upper part of middle member of Cliefden Caves Limestone,
Licking Hole Creek; 1 specimen (SUP 28250) from just below middle, thinly bedded unit of Bowan
Park Limestone, Quondong; 2 specimens (SUP 26218-19) from upper part of limestone, Malachi’s

Hill.

Description. Coenosteum laminar, up to 110x70 mm. diameter, 50 mm. high. Broad
mamelons 10-20 mm. apart. In one specimen (SUP 28250, PI. 126, fig. 3), coenosteum
intergrown with Propora and thick-walled tubes almost 1 mm. in diameter, interpreted
as caunopore tubes; also associated with Ecclimadictyon and algae.

In vertical section, laminae moderately continuous, gently undulating, spaced from
6 to 9 in 2 mm. vertically. Vesicular appearance where downward inflexions of laminae
meet pillars, and triangular space forms between each adjacent inflexion above pillar.
Pillars about 0-3-1 mm. apart; not seen to extend beyond one interlaminar space, and
occasionally do not reach underlying laminae. All tissue in coenosteum seems to be
compact. Wall thickness of laminae c. 50 p. Few vague astrorhizae appear to be present,
but not particularly restricted to mamelons. In tangential section, elements not well
preserved; faint speckling may represent pillars. Broad mamelons and downward
inflexions of laminae especially prominent.

Remarks. The New South Wales material closely resembles C. mammillatum (Schmidt
1858) from the late Ordovician of Estonia and the Ordovician or Silurian boulders on
the south coast of the Baltic (Nestor 1964, p. 42). The relatively poor preservation and
less prominent astrorhizae make it necessary to refer the material to C. aft', mammilla-
turn. The form of the pillars is the same as that depicted for C. striatellum (d’Orbigny)
by Stearn (1966, p. 90). According to him, the pillar spreads upwards to meet the
lamina, leaving a hollow space.

Clathrodictyon cf. microundulatum Nestor 1964
Plate 126, fig. 6; Plate 127, figs. 1-4

Material. 1 specimen (SUP 28257) from upper part of middle member of Cliefden Caves Limestone,
Licking Hole Creek; 6 specimens (SUP 26220-5) from ‘Island’ unit of upper member of limestone.
Island between Belubula River and Large Flat; I specimen (SUP 29133) from upper part of Bowan
Park Limestone, south-south-east of Malachi’s Hill.

Description. Coenosteum hemispherical to laminar, up to 120x90 mm. diameter,
120 mm. high. Broad mamelons spaced 10-20 mm. apart. Latilaminae range from 0-5-4
mm. in thickness, with sediment-filled spaces between some.

Vertical section shows undulating laminae of variable length and vesicular appearance,

658

PALAEONTOLOGY, VOLUME 12

spaced from 9 to 14 in 2 mm. vertically. Coenosteum frequently exhibits alternating
latilaminae and ill-defined or structureless, calcite-filled, or sediment-filled layers (pre-
sumably originally representing non-calcified tissue or pauses in growth). Alignment of
downward inflexions of laminae in some parts of coenosteum produces slightly zigzag
pillars which extend continuously through 4 or more interlaminar spaces; pillars spaced
from 0-2 to 0-6 mm. apart. Astrorhizae represented by a few scattered irregular tube-like
structures in coenosteum, about 0-5 mm. in diameter, including mamelons. In tangential
section, undulating laminae appear to be arranged concentrically around mamelons.
Only faint traces of pillars seen.

Remarks. Since C. microundulatum from the late Ordovician of Estonia (Nestor 1964,
p. 41) lacks mamelons and astrorhizae, the New South Wales material is only tentatively
assigned to it. The character of the undulating laminae is the same as that exhibited by
the Estonian species.

Specimens from the Island locality show a gradation from poorly calcified to more
completely preserved forms (PI. 127, fig. 1) resembling Ecclimadictyon amzassensis
(Khalfina 1960), a species which occurs in some abundance at this locality. Perhaps the
material should be assigned to Ecclimadictyon. However, Clathrodictyon and Ecclima-
dictyon are closely related (Nestor 1964, p. 107) and in practice it proves most difficult
to separate the more ‘primitive’ forms with undulating laminae.

EXPLANATION OF PLATE 126

Figs. 1, 2. Cliefdenella etheridgei gen. et sp. nov., x 5, ‘Island' unit, upper member, Cliefden Caves
Limestone, the Island. 1, SUP 24158, paratype, vertical section showing branching pillar, and
laminae with denticles. 2, SUP 24155, paratype, tangential section showing solid pillars, astro-
rhizal column with "septal’ structures, and laminae formed of compact tissue with denticles
represented by darker dots.

Figs. 3-5. Clathrodictyon aff. mammillatum (Schmidt), X 10. 3, 4, SUP 28250, from below middle,
thinly bedded unit, Bowan Park Limestone, Quondong; 3, vertical section showing association
with Propora and a caunopore tube; 4, tangential section. 5, SUP 26219, upper part of Bowan Park
Limestone, Malachi’s Hill; vertical section showing downward inflexions of laminae, with triangular
spaces developed between adjacent inflexions above pillars.

Fig. 6. Clathrodictyon cf. microundulatum Nestor, SUP 26221, x 10, vertical section, from ‘Island’
unit, upper member, Cliefden Caves Limestone, the Island; showing alternation of latilaminae
and poorly preserved to calcite-filled layers.

EXPLANATION OF PLATE 127

Figs. 1-4. Clathrodictyon cf. microundulatum Nestor, x 10. 1,2, from ‘Island’ unit, upper member,
Cliefden Caves Limestone, the Island. 1, SUP 26220, vertical section showing incipient develop-
ment of pillars like those in Ecclimadictyon amzassensis, and alternation of latilaminae and poorly
preserved layers; 2, SUP 26223, vertical section showing part of broad mamelon, and rhythmic
alternation of latilaminae and incompletely preserved layers. 3, 4, SUP 29133, upper part of Bowan
Park Limestone, SSE. of Malachi’s Hill ; 3, vertical section exhibiting undulating laminae of variable
size and prominent breaks in growth between successive latilaminae; 4, tangential section showing
undulating, cyst-like laminae concentrically arranged around mamelons.

Figs. 5-7. Ecclimadictyon amzassensis (Khalfina), X 10, ‘Island’ unit, upper member, Cliefden Caves
Limestone, the Island. 5, SUP 26206, vertical section showing zigzag-shaped laminae and slightly
zigzag-shaped pillars; note layer of irregular tissue at base of coenosteum (see also PI. 128, figs. 1,
3). 6, SUP 26211, vertical section showing astrorhizal canal. 7, SUP 26205, tangential section
showing dark dots which represent pillars.

Palaeontology, Vol. 12

PLATE 126

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 127

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROM ATOPOROIDS FROM NEW SOUTH WALES 659

Genus ecclimadictyon Nestor 1 964
Type species. Clathrodictyon fastigiatum Nicholson 1886.

Discussion. Nestor (1964) interpreted the genus Ecclimadictyon as closely related to
Clathrodictyon, but differing in having strongly crumpled (zigzag-shaped) laminae and
lacking pillars. He reported (1964, p. 107) a grained microstructure consisting of small
dark dots on a lighter background as seen in tangential section. Similarly, Stearn (1966,

р. 91), recorded dark dots in the type species, E. fastigiatum, and considered them to
be melanospheres formed from the originally compact tissue. Yet Nestor (1964, p. 69)
despite his interpretation, has assigned Clathrodictyon (?) kirgisicum amzassensis
Khalfma 1960 (a species which exhibits well developed pillars), to Ecclimadictyon and
has referred to ‘pillars’ developed as inflexions of laminae in another form, E. pandum
Nestor 1964. It therefore seems necessary to expand the conception of the genus
Ecclimadictyon to include species like E. amzassensis (Khalfma) exhibiting indubitable
pillars formed by downward inflexions of laminae.

Ecclimadictyon amzassensis (Khalfma 1960)

Plate 127, figs. 5-7; Plate 128, figs. 1-5

1960 Clathrodictyon (?) kirgisicum amzassensis Khalfma, p. 370, pi. O-l, figs. 1-3.

1964 Ecclimadictyon amzassensis Nestor, p. 69.

Material. 9 specimens (SUP 26205-13) from ‘Island’ unit of upper member of Cliefden Caves Lime-
stone, at Island between Belubula River and Large Flat; also occurs in upper part of Regan's Creek
Limestone, south-east of Cargo; from lower part of Cargo Creek Limestone, south of Cargo; from
lower part of Canomodine Limestone, north of The Glen. In addition reported from several localities
ranging stratigraphically from above to below the middle, thinly bedded unit of Bowan Park Lime-
stone, and from an Ordovician limestone lens near Eurimbula, north of Molong.

Description. Coenosteum hemispherical, up to 180 x 110 mm. diameter, 120 mm. high.
Some specimens show latilaminae, and rhythmic thickening and thinning of tissue.
Surface relatively smooth.

Vertical section exhibits network of zigzag-shaped laminae. Downward inflexions of
inclined lamina occur at different levels on either side of pillar giving zigzag appearance.
Pillars appear as long, zigzag-shaped vertical elements in well-orientated vertical sec-
tions; up to 12 pillars in 2 mm. laterally. Pillars and laminae form a compact (speckled?)
tissue, usually inaregular network, but in basal 0-5-2 mm. (PI. 127, fig. 5), or sometimes
at commencement of a latilamina, tissue is notably irregular. Zigzag-shaped laminae are
up to 10 mm. in length, spaced from 8 to 12 (usually 10) in 2 mm. vertically; rarely
updomed in small mamelons with astrorhizae; mamelons up to 2-5 mm. in diameter.
Latilaminae exhibited, from 4 to 33 mm. thick. Also rhythmic (possibly secondary)
thickening of tissue in some specimens (PI. 128, fig. 5), possibly due to slight fluctuations
in environmental conditions. Astrorhizal canals scattered through coenosteum, mainly

с. 0-5 mm. in diameter, and occur in mamelons and more commonly elsewhere.
Coenosteum of SUP 26209 encrusts the smaller E. nestori (PI. 128, fig. 1).

In tangential section, pillars usually spaced 0-2-0-3 mm. apart; appear as dots which
may be connected by lamellar elements to form bars, or a more complicated meshwork
of vermiculate to meandriform appearance. In a few places the meshes become com-
pletely closed.

660

PALAEONTOLOGY, VOLUME 12

Remarks. The New South Wales material is remarkably similar to E. amzassensis
described by Khalfina (1960, p. 370) from the Upper Ordovician of the Altai-Sayan
mountain region, south-west Siberia, and is therefore assigned to it. The holotype is from
the Amzass Formation of Gornaya Shoriya. Other figured specimens come from
localities in the Altai mountains. The Amzass Formation and its equivalents, the
Orlovsk Formation of Altai and Chumysh Formation of Salair, are correlated by
Ivanovsky (1965) with the upper Caradoc of Europe.

Other species may be compared with E. amzassensis ; E. kirghisicum (Riabinin 1931)
from the Upper Silurian of northern Kazakhstan bears similarities, but lacks astro-
rhizae (Yavorsky 1955), and E. pandum from the Llandoverian of Estonia is comparable
but has less well-pronounced pillars and cone-shaped astrorhizae (Nestor 1964).

Ecclimadictyon nestori sp. nov.

Plate 128, fig. 1 ; Plate 129, figs. 1-6

Material. 1 specimen (SUP 28256) from upper part of middle member of Cliefden Caves Limestone,
Licking Hole Creek; 7 specimens (SUP 26199-204, 26209) from 'Island' unit of upper member of
limestone, Island between Belubula River and Large Flat. Also occurs in horizons just above and
below middle, thinly bedded unit of Bowan Park Limestone, and in Ordovician limestone lenses near
Eurimbula, north of Molong.

Holotype. SUP 26203; other numbered specimens designated paratypes.

Description. Coenosteum laminar to encrusting, up to 180x 150 mm. diameter, 40 mm.
high. Latilaminae often well developed. Upper surface of one specimen (SUP 26201)
exhibits moderately large mamelons, 12-20 mm. apart.

Vertical section shows subreticulate structure with zigzag-shaped laminae and poorly
developed short pillars formed from downward inflexions of laminae; in most cases

EXPLANATION OF PLATE 128

All specimens come from 'Island' unit, upper member, Cliefden Caves Limestone, the Island.

Fig. 1. Ecclimadictyon amzassens is (Khalfina) and Ecclimadictyon nestori sp. nov., SUP 26209, X 10;
vertical section showing E. amzassensis encrusting paratype of the smaller E. nestori.

Figs. 2-4. Ecclimadictyon amzassensis (Khalfina), SUP 26206. 2, tangential section showing bar-like
connections formed from downflexed horizontal elements between pillars, x 10. 3, vertical section
exhibiting zigzag-shaped laminae, pillars, astrorhizae, and a prominent break in growth between
successive latilaminae, x 5. 4, tangential section showing bars and more complex meshworks of
intersected folded horizontal elements; note also astrorhizae; x5.

Fig. 5. Ecclimadictyon amzassensis (Khalfina), SUP 26207, x 5, vertical section showing prominent
rhythmic thickening and thinning of tissue.

EXPLANATION OF PLATE 129

Figs. 1-6. Ecclimadictyon nestori sp. nov., X 10, ‘Island’ unit, upper member, Cliefden Caves Lime-
stone, the Island. 1. SUP 26199, paratype, vertical section showing zigzag-shaped laminae and
poorly developed zigzag-shaped pillars. 2, SUP 26203, holotype, vertical section showing cauno-
pore tube between latilaminae, with offset extending vertically through coenosteum. 3, SUP 26200,
paratype, vertical section exhibiting zigzag laminae and bunchy astrorhizae. 4, SUP 26201, para-
type, tangential section showing dark specks representing pillars, some of which are connected in
meshwork of horizontal elements. 5, SUP 26204, paratype, vertical section showing break in growth
between successive latilaminae. 6, SUP 26203, holotype, tangential section showing laminae con-
centrically arranged around two caunopore tubes.

Palaeontology, Vol. 12

PLATE 128

WEBBY, Ordovician stromatoporoids from New South Wales

Palaeontology, Vol. 12

PLATE 129

WEBBY, Ordovician stromatoporoids from New South Wales

WEBBY: ORDOVICIAN STROMATOPOROIDS FROM NEW SOUTH WALES 661

laminae confined to interlaminar spaces, but exceptionally may extend as zigzag elements
through up to 4 interlaminar spaces. Laminae usually spaced 15-16 in 2 mm. vertically.
Bunchy astrorhizae associated with updomed laminae scattered through coenosteum;
not particularly associated in mamelons. Latilaminae frequently well developed, 4-12
mm. thick; may be in contact, or separated by sediment and calcite-filled cavities. At
commencement of each new latilamina, thin zone less than 0-4 mm. thick of less regular
tissue (PI. 129, fig. 5). Large thicker-walled cylindrical structures (probably representing
caunopore tubes formed by a commensal organism) about 1 mm. in diameter; tend to
be associated with mamelons; also horizontal tubes of same organism in sediment
between latilaminae. No dissepiments seen.

In tangential section, pillars appear as fine dots spaced 0- 1-0-2 mm. apart; usually
connected with obliquely cut laminae to form partly open meshwork having vermiculate
appearance. Large, relatively thick-walled caunopore? tubes clearly shown (PI. 129,
fig. 6), with laminae concentrically arranged around them.

Remarks. E. nestori bears the closest similarity to forms belonging to the E. micro-
vesiculosum group of Nestor (1964) from the Llandoverian of Estonia, and may be
regarded as a species of the E. microvesiculosum group, being distinguished from other
representatives of the group by having 15-16 laminae in 2 mm. vertically, bunchy astro-
rhizae, and lacking dissepiments. E. microvesiculosum (Riabinin) has dissepiments,
16-19 laminae in 2 mm., and lacks astrorhizae. E. microfastigiatum (Riabinin) has star-
like astrorhizae and 14-15 laminae in 2 mm. E. macrotuberculatum (Riabinin), the third
Estonian member of the group, has dissepiments and 12 laminae in 2 mm.

Acknowledgements. The author wishes to thank V. Semeniuk, R. A. McLean, and J. G. Byrnes for
providing specimens for study from areas at Bowan Park, Regan’s Creek, and north of Molong,
respectively. The work was supported by funds from the Australian Research Grants Committee.

REFERENCES

Adrian, j. 1956. The geology of the Molong District. Unpublished B.Sc. (Hons.) Thesis, University of
Sydney, 138 pp.

banks, m. r. 1957. The stratigraphy of Tasmanian limestones. Min Resour. Tasm. 10, 39-85.

1962. Ordovician system. The geology of Tasmania. J. geol. Soc. Aust. 9, 147-76.

cooper, g. a. 1956. Chazyan and related brachiopods. Smithsonian Misc. Colt. 127 (1), xvi+ 1-1024.
dean, w. t. 1967. The distribution of Ordovician shelly faunas in the Tethyan region. In Aspects of
Tethyan biogeography (ed. adams, c. g. and ager, d. v.). Syst. Assoc. Publ. 7, 1 1-44.
etheridge, r. jr. 1895. On the occurrence of a stromatoporoid, allied to Labechia and Rosenella, in
the Siluro-Devonian rocks of N. S. Wales. Rec. Geol. Surv. N.S.WA. 134-40.
galloway, j. j. 1957. Structure and classification of Stromatoporoidea. Bull. Am. Paleont. 37, no.
164, 341-480.

and st. jean, j. 1961 . Ordovician Stromatoporoidea of North America. Ibid., 43, no. 194, 1-1 1 1.

hill, d. 1957. Ordovician corals from New South Wales. J. Proc. R. Soc. N.S.W. 91, 97-107.
ivanovsky, A. b. 1965. Stratigraficheskiy i paleobiogeograficheskiy obzor rugoz ordovika i silura. Nauka,
Moscow, 150 pp. ( Akad . Nauk SSSR, Sib. Otd. Inst. Geol. Geofiz.)
khalfina, v. k. 1960. Stromatoporoidei. In khalfin, l. l. (ed.). Biostratigrafiya paleozoya Sayano-
Altayskoy Gornoy oblasti. Tom I, Nizhniy Paleozoy. Trudy sib. nauchno-issled. Inst. Geol. Geofiz.
miner. Syr., vyp. 19, pp. 82-4, 141-3, 357-8, 370-3.

andYAVORSKY, v. i. 1967. 0 drevneyshikh stromatoporoideyakh. Paleont. Zh. 1967, no. 3, pp. 133-6.

moors, h. t. 1966. Some of the stratigraphy and palaeontology of the Cliefden Caves District, near
Mandurama. Unpublished M.Sc. Thesis, University of Sydney, 115 pp.

662

PALAEONTOLOGY, VOLUME 12

nestor, h. e. 1964. Stromatoporoidei Ordovika i Llandoveri Estonii. Geoloogia-Inst. Uurim. 112 pp.

■ 1966. O drevneyshikh Stromatoporoideyakh. Paleont. Zh. 1966, no. 2, 3-12.

ozaki, k. e. 1938. On some stromatoporoids from the Ordovician limestone of Shantung and South
Manchuria. J. Shanghai Sci. Inst., Sect. 2, 2, 205-23.
packham, g. h. 1960. Sedimentary history of part of the Tasman Geosyncline in south-eastern
Australia. Repts. 21st lnt. Geo/. Cong. Norden, 12, 74-83.

1967. The occurrence of shelly Ordovician strata near Forbes, New South Wales. Anst. J. Sci.

30, 106-7.

parks, w. a. 1910. Ordovician stromatoporoids. Univ. Toronto Stud., geol. ser., no. 7, 1-52.
philip, g. m. 1966. The occurrence and palaeogeographic significance of Ordovician strata in Northern
New South Wales. Aust. J. Sci. 29, 112-13.

Phillips ross, j. 1961. Ordovician, Silurian, and Devonian Bryozoa of Australia. Bull. Bur. Miner.
Resour. Geol. Geophys. Aust. 50, 1-172.

sherrard, k. m. 1954. The assemblages of graptolites in New South Wales. J. Proc. R. Soc. N.S.W.
87, 73-101.

stearn, c. w. 1966. The microstructure of stromatoporoids. Palaeontology 9, 74-124.
stevens, n. c. 1950. The geology of the Canowindra district, N.S.W. Part 1. The stratigraphy and
structure of the Cargo-Toogong District. J. Proc. R. Soc. N.S.W. 82, 319-37.

1952. Ordovician stratigraphy at Cliefden Caves, near Mandurama, N.S.W. Proc. Linn. Soc.

N.S.W. 77, 114-20.

1956. Further notes on Ordovician formations of Central New South Wales. J. Proc. R. Soc.

N.S. W. 90, 44-50.

thomas, d. e. 1960. The zonal distribution of Australian graptolites. Ibid. 94, 1-58.
vlasov, a. n. 1961. Kembriyskiye stromatoporoidei. Paleont. Zh. 1961, no. 3, 22-32.
webby, b. d. 1968. Astrocystites distans sp. nov., an edrioblastoid from the Ordovician of eastern
Australia. Palaeontology 11, 513-25.

Whittington, h. b. 1966. Phylogeny and distribution of Ordovician trilobites. J. Paleont. 40, 696-737.

and williams, a. 1964. The Ordovician period. In The Phanerozoic time-scale. London (Geological

Society), 241-54.

yabe, h. and sugiyama, t. 1930. On some Ordovician stromatoporoids from South Manchuria, North
China and Chosen (Corea), with notes on two new European forms. Sci. Repts. Tohoku Imp. Univ.,
ser. 2 (Geol.), 14, no. 1, 47-62.

yavorsky, v. i. 1955. Stromatoporoidea Sovetskogo Soyuza, Pt. I. Trudy vses. nauchno-issled. geol.
Inst., N.s. 8, 1-173.

B. D. WEBBY

Department of Geology and Geophysics
University of Sydney
Sydney 2006, N.S.W.

Typescript received 27 February 1969 Australia

THE WENLOCK GRAPTOLITES OF THE LUDLOW
DISTRICT, SHROPSHIRE, AND THEIR
STRATIGRAPHICAL SIGNIFICANCE

by C. H. HOLLAND, R. B. RICKARDS, and P. T. WARREN

Abstract. Monograptus ludensis (Murchison 1839) sensu Wood 1900 (with its synonyms M. gotlandicus Perner
1899 and M. vulgaris Wood 1900) is fully described together with a new species Pristiograptus jaegeri , the two
having been frequently confused with M. vulgaris. A second new species described from the Wenlock rocks of
the Ludlow district, Shropshire, is Holoretiolites ( Balticograptus ) lawsoni, and another form Pristiograptus sp. 1
is left under open nomenclature. M. deubeli Jaeger 1959 is recorded for the first time in the British Isles. The
stratigraphical distribution of the graptolite fauna of the Wenlock and lowest Ludlow of the Ludlow district
is discussed and comparisons made particularly with North Wales. A poorly developed ‘ nassajdubius Inter-
regnum’ (Jaeger 1959) separates the C. lundgreni Zone, with its abundant M. flemingii (Salter), from the M.
ludensis Zone with its association of P. jaegeri sp. nov. and the index species. The problem of the correlation of
the Wenlock/Ludlow boundary is discussed and it is recommended that in the graptolite sequence the horizon
best correlated with this boundary is the base of the P. nilssoni Zone.

In their revision of the stratigraphy of the Silurian rocks of the Ludlow district, Holland,
Lawson, and Walmsley (1963) defined the Ludlow Series and its four component stages
by means of standard sections for the boundaries between these and adjacent strati-
graphical units. The higher Wenlock rocks cropping out in the south-western part of
the district were given detailed description in terms of lithology and fauna though,
naturally, no attempt was made at a definitive classification of the Wenlock Series. The
boundary between the Wenlock and Ludlow was, however, effectively designated by the
choice and accurate description of a standard section for the base of the Ludlow Series
and its lowest Eltonian Stage. At this standard locality, in an old quarry in Pitch Coppice
on the south side of the Ludlow-Wigmore road, nearly 5 m. of Wenlock Limestone are
followed by 1 or 2 m. of Lower Elton Beds. Conscious that they were stabilizing rather
than completely solving the problem of the Wenlock/Ludlow boundary, Holland et a/.
(1963, p. 141) referred to the remaining difficulty, viz. that the position of the base of
the Monograptus vulgaris Zone in relation to the Wenlock Limestone was not known.
‘The problem of the graptolite sequence in areas where the Wenlock Limestone is
developed might be solved by prolonged collecting throughout the Welsh Borderland
but the rarity of graptolites at this level in the shelf facies would make this a most
difficult task. In any event, it is desirable that at the standard locality the Wenlock
Limestone should be within the Wenlockian and the Ludlovian (Eltonian) should begin
above it.’

Subsequently, two of the present authors (C. H. H. and R. B. R.) decided to attempt
a revision of the Wenlock graptolites of the Ludlow district, the available background
of carefully documented localities permitting the possibility of rigorous collection from
all exposed Wenlock and basal Ludlow. The third author ( P. T. W.) was in the meantime
concerned with the precise revision of the graptolite sequence at the same level in the
geosynclinal area of North Wales and had encountered a nomenclatural question con-
cerning the species M. vulgaris. Thus, the present paper sets out not only to describe the

[Palaeontology, Vol. 12, Part 4, 1969, pp. 663-83, pi. 130.]

664

PALAEONTOLOGY, VOLUME 12

graptolite fauna of the Wenlock rocks of the Ludlow district but also to attempt to solve
the taxonomic and stratigraphical problems associated with M. vulgaris, the zone of
that name, and the Wenlock/Ludlow boundary. Two preliminary notes on these matters
have already been published (Warren, Rickards, and Holland 1966; Holland, Rickards,
and Warren 1967).

Localities within the Ludlow district are numbered as in Holland et al. (1963), where
grid references and other topographical details are given. Where it has proved necessary
to subdivide these localities the relevant information is provided herein. Our graptolite
collection from the Ludlow district has been deposited in the Geological Museum of
Trinity College, Dublin, and TCD numbers are given. Other material described herein
is in possession of the Sedgwick Museum, Cambridge [SM], Birmingham University
[BU], and the Institute of Geological Sciences [GSM GSC (Geological Society of
London Collection), GSM Zp (Boswell Collection)].

Acknowledgements. One of us (P. T. W.) thanks Dr. A. G. Brighton (Cambridge) and Dr. I. Strachan
(Birmingham) for the loan of specimens in their care and also Dr. H. Jaeger (Berlin) for helpful dis-
cussions on the ludensis Zone faunas of both North Wales and Germany. Both C. H. H. and P. T. W.
are grateful to Dr. L. Teller (Warsaw) for affording them the opportunity individually to see and
collect from the Pragowiec section in the Holy Cross Mountains. C. H. H. has had the benefit of a visit
to sections in the German Democratic Republic under Dr. Jaeger's expert and hospitable guidance.
C. H. H. and R. B. R. thank Dr. T. R. Lister, Christine Rickards, and Dr. J. H. Shergold, for assistance
in collecting from the Ludlow district. Dr. Warren's contribution is published by permission of the
Director, Institute of Geological Sciences.

THE ‘ VULGARIS ’ PROBLEM

In the systematic descriptions below it will be shown that Monograptus ludensis ( sensu
Wood 1900), M. got/andicus Perner 1899, and M. vulgaris Wood 1900 are conspecific.
We have already indicated our preference for usage of the name ludensis (Warren et ah
1966, Holland et al. 1967) and our views have been accepted by Martinsson (1967). In
addition we have received a number of personal communications variously advocating
the usage of ludensis, gotlandicus , or vulgaris. It would seem that several workers have
more or less concurrently concluded that these forms are conspecific.

Of these three the earliest published name is ludensis Murchison 1839, and subse-
quent to Wood’s (1900) description of this species nothing has been added to its dia-
gnosis. Although it has been but rarely recorded. Wood’s concept of ludensis has not been
changed or abused. Largely because of her definitive work ludensis cannot be considered
a confused species. Nevertheless, although she clearly interpreted the species by reference
to fig. 2 of Murchison's original illustrations (Murchison 1839, pi. 26), Wood did not
specify (as lectotype) a particular specimen from those shown crowded together on the
slab depicted there.

It is true that earlier M‘Coy (1851, p. 4) gave as ‘Ref.’ (i.e. synonymy) ‘Sil. Syst. t.
26. f. 1, & la. (Not. 2.)’, thus referring to those figures depicting a Monograptus priodon
(Bronn)-Iike form; but in describing fossils from the Geological Museum of the Uni-
versity of Cambridge he cannot be taken as ‘first reviser’ of ludensis in the sense of
the Rules. There is no question of M'Coy’s having selected a lectotype from the type
series as the Rules require; he was simply assigning material to one particular previ-
ously published figure and not to another.

Murchison himself in Si/uria( 1854, pi. 12, figs. 4 and 4a) chose to repeat only figs. 1 and
1 a of his original plate from the ‘Silurian System’ and referred to the form shown there

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 665

as ‘Graptolithuspriodon (Ludensis), Bronn’, thus leaving outside this implied synonymy
his original fig. 2. However, in Siluria (3rd ed., 1859) Murchison again included both
original figures under the name ‘Gr. priodon Bronn [Gr. ludensis, Sil. Syst.]’.

The principal objection to the use of gotlandicus Perner 1899 as the specific name for
the graptolite in question is that the original figure is of a distal rhabdosomal fragment.
However, modern Czechoslovakian and Polish workers have adequately redefined the
species, and again there has been no confusion in the literature. M. gotlandicus has been
widely recorded except in Britain (an omission explained once the conspecificity of
ludensis , vulgaris, and gotlandicus is appreciated), and the name is well entrenched in
the literature.

By contrast M. vulgaris Wood 1900 must be one of the most frequently misidentified
species in graptolite literature. The majority of records and illustrations of the species
are based upon incorrect identification. Our recent examination of the Das Gupta collec-
tion (Imperial College, London), for example, showed that some specimens labelled
‘ vulgaris ’ were referable to spinose graptolites of the M. leintwardinensis Lapworth type.
The confusion over the identification of vulgaris arises partly from the fact that Wood’s
(1900) original description involved two distinct species. One of these, a straight, slender
pristiograptid (P. jaegeri sp. nov. herein) has been commonly found and recorded as
vulgaris, whereas Elies and Wood (1911) clearly designate a specimen belonging to the
other form as ‘type specimen’. As implied above, the true ‘ vulgaris ’ has been widely
recorded outside Britain as gotlandicus , whilst in Britain some specimens in Elles's and
Wood’s collections were referred to ludensis and other material, notably in the Boswell
Collection, was assigned either correctly or to M. vulgaris cf. curtus Elies and Wood and
M. colonus. Few indications of vulgaris in the literature are, therefore, correct and none
clearly defines the morphology either of the pristiograptid or of M. vulgaris Wood (pars).

The true vulgaris we would retain in Monograptus since it has distinctly rounded lateral
aperture margin.s to th 1, and does not fall readily into either Pristiograptus Jaekel or
Co/onograptus Pribyl. M. ludensis as defined by Wood is, as we now know, identical
with this, and Wood’s failure to notice the conspecificity reflects the poor preservation
of specimens collected from the locality of the type specimen of M. ludensis. We are
fortunate in having in our possession isolated Polish specimens of M. gotlandicus
Perner. Comparison of these with our now extensive collections of ludensis and vulgaris
indicates that the three named species are conspecific and that the pristiograptid form
(common also in the Polish material) requires a new name.

The problem, therefore, is complex in that none of the three names is entirely satis-
factory. Our discussions with colleagues have indicated that to choose one name will
please but a fraction of the interested parties. At this point we shall state our views on the
matter, and then describe the species below under our preferred name. The immediate
aims are to establish the conspecificity of ludensis, gotlandicus, and vulgaris, and to apply
our preferred name to the relevant biostratigraphical unit. It is our view that the interests
of palaeontology and stratigraphy will best be served if the name vulgaris is not (by
special provision) adopted. It is the most junior name and yet is shrouded in the greatest
misunderstanding. Of the two remaining names we prefer properly to follow the Rules
still further in adopting the senior name ludensis, though we appreciate that it is less well
established in the literature than gotlandicus.

x x

C 6940

666

PALAEONTOLOGY, VOLUME 12
SYSTEMATIC DESCRIPTIONS

Family retiolitidae Lapworth 1873
Subfamily plectograptinae Boucek and Munch 1952
Genus holoretiolites Eisenack 1951
Subgenus balticograptus Boucek and Munch 1952

H. ( Balticograptus ) lawsoni sp. nov.

Text-figs 1 a-c

Diagnosis. Rhabdosome tiny, parallel-sided; thecae inconspicuous, apertures ventrally
facing and surmounted by loop-shaped processes which show, in the form of lists, traces
of the graptolite fusellar structure; thecal spacing 13 in 10 mm.; clathrial elements well-
developed, reticula imperfect and sparse; nema subcentral.

Holotype. The specimen illustrated as Fig. \a, number TCD 8473.

Material. Three well-preserved specimens in low relief, and other fragmentary specimens (TCD 8473-
5).

Horizon and locality. Wenlock Shale, Wenlock Series, ludensis Zone. Ludlow district, Burrington Farm
lane, section, locality 62 of Holland et al. (1963): see text-fig. 4. Associates are: Monograptus ludensis
and Pristiograptus jaegeri sp. nov.

Derivation of name. After Dr. J. D. Lawson, who collected the specimens during the 1964 Ludlow
Research Group meeting in the Ludlow district.

Description. The maximum observed rhabdosome length is 4-00 mm. and the dorso-
ventral width (low relief) 2-70 mm. Excluding apertural processes the rhabdosome
achieves a dorso-ventral width of 0-90-1 -00 mm. and is more or less parallel-sided. The
extreme proximal end is almost rectangular in outline and rapidly reaches the maximum
dorso-ventral width. The thecal spacing is 13 in 10 mm. Text-fig. 1 b depicts the apertural
processes at their maximum observed length of 0-80-0-90 mm.

Clathrial elements appear to form a very approximately hexagonal mesh on the lateral
walls. The free ventral thecal walls are roughly vertical or gently concave, turning out-
wards slightly in the region of the apertures, and are defined by quite robust paired
ventral lists. An imperfect and sparse reticulum is indicated by the presence and distri-
bution of much finer threads.

The thecal apertures are obscure, but are probably similar to those of H. (B.) baiticus
Eisenack, i.e. ventrally facing. Loop-shaped processes surmount the apertures and pro-
ject ventrally, or somewhat proximally, for a distance of 0-80-0-90 mm. The loop may
exhibit some transverse expansion away from the thecal aperture (text-figs. 1 a , b) and
is infilled with slender cross threads. The cross threads form a rough ‘zigzag suture’ and
could possibly be homologous with the growth bands of sclerotized graptolites. One
specimen (text-fig. lc) shows a slender nema in a subcentral position. Distally there is
an abbreviated appendix (text-figs. 1 a , b) and a general slight tapering, clearly indicating
that the rhabdosomes have reached their adult length.

Remarks. This species clearly belongs to the subgenus Balticograptus and is distinguished
from H. ( Holoretiolites ) by the presence of a poor reticulum, and a less tapering rhab-
dosome. H. (B.) lawsoni is perhaps closest to Eisenack’s species H. (B.) baiticus but
differs in having longer apertural processes, a less parallel-sided and more robust

c

text-fig. 1 a-d. Holoretiolites ( Balticograptus ) lawsoni sp. nov. a, TCD 8473, holotype, ludensis Zone,
Wenlock Series, locality 62 Burrington Farm; b, TCD 8475, same horizon and locality as holotype; c,
TCD 8474, same horizon and locality as holotype; d, Pristiograptus sp. 1, TCD 8654, part of specimen
on PI. 130, fig. 6, showing pattern in cortical tissue.

Figs. 1 c-cx 20, 1 dx 30; is, interthecal septum; stipple indicates ferruginous staining; oblique
ruling indicates nearside of rhabdosome in lc.

668

PALAEONTOLOGY, VOLUME 12

rhabdosome, and more closely spaced thecae. From Retiolites clathrospinosus Eisenack,
H. ( B .) lawsoni differs in having loop-shaped apertural processes as opposed to paired
bifurcating spines, a less well-developed reticulum, and thecae which approximate to a
‘climacograptid’ rather than an ‘orthograptid’ type. The general dimensions of the two
species are similar. Holoretiolites Eisenack is recorded for the first time in Britain.

Family monograptidae Lapworth 1 873
Genus pristiograptus Jaekel 1889
Pristiograptus jaegeri sp. nov.

Plate 130, fig. 1, text -figs 2k-r, 3/

1900 Monograptus vulgaris Wood, pp. 455-6 (pars), text-fig. 10« (non 106, pi. 25, fig. 2).

1911 Monograptus vulgaris Wood; Elies and Wood, pp. 378-9 (pars), text-fig. 248a (non
2486, pi. 37, figs. 10 a-e).

71935 Monograptus vulgaris Wood; Decker, pp. 443-4, figs. 32-4.

71935a Monograptus vulgaris Wood; Decker, p. 309.

1943 Pristiograptus vulgaris vulgaris (Wood); Pribyl, text-fig. 2l (non 2m), 7 pars, pp. 22-3.

71947 Monograptus vulgaris Wood; Ruedemann, p. 490, pi. 84, figs. 22-4.

71948 Pristiograptus cf. vulgaris vulgaris (Wood 1900); Pribyl, p. 78.

1963 Monograptus vulgaris Wood; Holland et a/., pp. 104, 136, 157.

non 1900 Monograptus vulgaris var. /3 Wood, p. 457, pi. 25, fig. 3.

non 1911 Monograptus vulgaris var. curtus Elies and Wood, p, 379, pi. 37, fig. 1 1.

non 1960 Monograptus vulgaris var. ashlandensis Berry, p. 1163, fig. 2h.

Diagnosis. Rhabdosome of moderate length, relatively slender, and ‘stiff’, particularly at
the proximal end; sicula 2 mm. long, reaching aperture of th 2; over-all thecal spacing
12-8 in 10 mm.; distal thecae inclined at 40°; maximum dorso-ventral width at 20 mm.,
T30-F50 mm (relief).

Holotype. The specimen figured herein as Plate 130, fig. 1, and text-fig. In, TCD 8661.

Material. Many specimens flattened and in relief, numerous isolated specimens.

Horizon, ludensis Zone in the Ludlow district and uppermost lundgreni and ludensis Zones in North
Wales; nilssoni Zone of Pragowiec, Bardo Syncline, Holy Cross Mountains, Poland.

Localities. See text-fig. 4 for Ludlow and Conway Valley (North Wales) areas; Holy Cross Mountains,
Poland (see Tomczyk 1962, fig. 9, p. 45).

Derivation of Name. After Dr. H. Jaeger (Humboldt University, Berlin).

Description. The rhabdosome reaches several centimetres in length but is usually of the
order of 20 mm., where the dorso-ventral width is approximately 1-30-1 -50 mm (in
relief ). Flattened and compressed specimens may have a distal dorso-ventral width of up

EXPLANATION OF PLATE 130

Figs. 1-8. 1, Pristiograptus jaegeri sp. nov., holotype, TCD 8661, ludensis Zone, Wenlock Limestone
locality 100a. 2, Monograptus ludensis (Murchison) sensu Wood 1900, TCD 8658, ludensis Zone,
Wenlock Shale, locality 61. 3, Monograptus deubeli Jaeger, TCD 8657, ludensis Zone, Wenlock
Limestone, locality 114d. 4, Gothograptus nassa (Holm), TCD 8653, ludensis Zone, Wenlock Shale,
Locality 40. 5, 6, Pristiograptus sp. 1, TCD 8660, 8654, lundgreni Zone, Wenlock Shale, locality 91.
7, Pristiograptus dubius (Suess), TCD 8663, ludensis Zone, Wenlock Shale, locality 40. 8, Monograptus
flemingii (Salter), TCD 8652, lundgreni Zone, Wenlock Shale, locality 79. Figs. 1, 2, 3, 6, and 8 x 10;
fig. 4 x7; figs. 5 and 7x5. None retouched. Localities within Ludlow district as in text.

Palaeontology, Vol. 12

PLATE 130

HOLLAND, RICKARDS and WARREN, Wenlock-Ludlow graptolites

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 669

to 1-90 mm. A typical feature of the species is the ‘stiff’ rhabdosome, particularly at the
proximal end (text-figs. 2, 3) which only occasionally shows very slight dubius-Uke curva-
ture. The sicula has a length of at least 2-00 mm., the apex reaching to the level of the
aperture of th 2. The over-all proximal thecal spacing is 12-10 in 10 mm. falling to
almost 8 in 10 mm. distally in the most extreme variants. Undistorted specimens in relief
usually have a thecal spacing of 1 1-10 in 10 mm. proximally and 10-9 in 10 mm. distally.
The thecae are of simple pristiograptid type throughout the length of the rhabdosome,
and are inclined to the axis at about 40°, in the distal regions. Rare specimens show a
slight rounding (incipient lappets) of the apertural region of th 1 (text-fig. 2k). Such a
degree of rounding of th 1 in P. jaegeri would be most difficult to detect in a specimen
not isolated from the matrix. Thecal overlap distally is approximately two-thirds. A small
number of specimens has been obtained from Ludlow in which a strange lateral thecal
spine is seen on th 2 at about the level of the thecal aperture of th 1. We consider that
these are probably abnormal and that the spine is not a character of specific importance.

Remarks. It is this species which has been most commonly recorded, and indifferently
figured and described, by numerous authors as M. vulgaris Wood. Some of Das Gupta’s
(1933) specimens, for example, although with a ‘stiff’ proximal end, are referable to
M. leintxvardinensis and not to ' M. vulgaris'. Pribyl (1948), however, selected as the
lectotype of M. vulgaris the specimen figured by Wood (1900) as pi. 25, fig. 2 and re-
figured by Elies and Wood (1911) as Monograptus vulgaris Wood (pi. 37, fig. 10#). It is
of interest that Elies and Wood (op.cit.) had already effectively designated this specimen
as ‘type’ (plate description of pi. 37). The species is distinct from the pristiograptid
described herein as P. jaegeri sp. nov. Monograptus vulgaris sensu Elies and Wood does
not merit retention in the genus Pristiograptus, an assignation adopted by some workers
(e.g. Pribyl 1943). M. vulgaris sensu stricto is considered below under the description
of Monograptus ludensis (Murchison).

Pristiograptus jaegeri sp. nov. occurs quite commonly associated with M. ludensis ,
although on evidence from North Wales its first appearance may be somewhat earlier
than that of the latter. Assemblages isolated from calcareous nodules from the Holy
Cross Mountains, Poland, almost invariably yield both species. The faint rounding of
th 1, rarely exhibited by P. jaegeri , perhaps suggests an evolutionary relationship
between this species and M . ludensis (Murchison).

Work in progress by one of us (P. T. W.) on the graptolites of North Wales suggests
that P. jaegeri may eventually be usefully subdivided into several subspecies, and that
some evolutionary connections may be unravelled. At the moment we would note that
more slender forms seem to occur at lower stratigraphical levels, in lundgreni and basal
ludensis Zone beds.

P. jaegeri may be distinguished from P. dubius (Suess) by the straight, ‘stiff’ proximal
end and the long sicula reaching to the level of th 2. As with all pristiograptids, identi-
fication is most difficult if few specimens are at hand. There is considerable variation
of such factors as dorso-ventral width, thecal spacing, etc., depending on whether the
rhabdosome is flattened or in relief and, further, on whether the apparent width has
been increased or decreased by compression in the strata. The North Wales specimens
tend, for example, to be less well preserved than the isolated Polish or the three dimen-
sional Ludlow specimens, and are usually flattened and often compressed. Consequently,

670 PALAEONTOLOGY, VOLUME 12

TEXT-FIG. 2

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 671

a greater dorso-ventral width is sometimes achieved (up to 1 -90 mm.) and the thecal
spacing shows a greater range of variation.

P. jaegeri differs from M. vulgaris ashlandensis Berry 1960 mainly in having a longer
and more conspicuous sicula. We consider that Berry’s form is at present best regarded
as a species in its own right distinct from M. ludensis, probably referable to the genus
Pristiograptus. P. ashlandensis is known only from flattened material and there may be
some doubt about its true thecal characters.

Pristiograptus sp. 1

Plate 130, figs. 5, 6, text-fig. Id

Material. Twenty-four specimens almost flat or in low relief.

Horizon and localities. Wenlock Shale, Wenlock Series, lundgreni Zone, localities 91 and 38 of Holland
et at. (1963). Associated are: Monograptus flemingii and Pristiograptus dubius.

Description. The rhabdosome reaches a length of at least 35 mm. and a distal dorso-
ventral width (low relief) of 2-40 mm. A width of 2-00 mm. is achieved quite close to the
proximal end, usually at about 7 mm. above the base of the sicula or approximately
th 9-th 10. The proximal thecal spacing is of the order of 13 in 10 mm., although some
badly squashed specimens reach 14 in 10 mm. Distally a value of 10 in 10 mm. is the
widest observed spacing.

The sicula has a length of 2-00 mm. the apex reaching the level of the second thecal
aperture. In the mesial and distal parts of the rhabdosome the thecal tubes are inclined
at 45-50° to the axis. Thecal overlap in the same regions is of the order of two-thirds.

text-fig. 2 a-j- Monograptus ludensis (Murchison 1839) sensu Wood 1900: a, TCD 8658, specimen in
relief, ludensis Zone, locality 61 Ludlow district (text-fig. 4); b, SM A60900, isolated specimen in full
relief, nodule from Pragowiec, Bardo Syncline, Holy Cross Mountains, Poland, probably nilssoni
Zone, presented to the Sedgwick Museum by Dr. A. Urbanek; c, d , SM A60904-5, isolated, mature
specimens in moderate relief from Pragowiec, nilssoni Zone, showing infilling of apertural region with
late tissue, nodule from Dr. A. Urbanek, specimens prepared by Dr. G. H. Packham 1960; e, SM
A60901, broad, immature, isolated, specimen in full relief, nodule from Pragowiec, probably nilssoni
Zone, angles of thecal apertures, not yet infilled with late tissue; /, GSM GSC 6584a, proximal end,
in full relief, of proposed lectotype, illustrated in text-fig. 3b. ludensis Zone; g, TCD 8744, external
mould of specimen in low relief, ludensis Zone, locality 41 Ludlow district (text-fig. 4); /;, TCD 8744a,
internal mould of th 1 and th 2 of previous specimen showing rounding of th 1 not apparent on external
mould; /, SM A60906, proximal end in full relief, nodule from Pragowiec, Holy Cross Mountains,
Poland, nilssoni Zon e;j, GSM Zp 5670, abnormal development of dorsal lip of sicula, specimen almost
flattened, ludensis Zone, Oerfa, Llanrwst, N. Wales (N.G.R. SH 8456 5950), Boswell's locality 482;
k-r, Pristiograptus jaegeri sp. nov. : k, BU 1470, specimen originally figured by Wood (1900) as Mono-
graptus vulgaris, text-fig. 10a and again by Elies and Wood (1911) as text-fig. 248a; /, GSM Zp 5720,
proximal end of specimen illustrated herein as text-fig. 3 f ludensis Zone, Cefn-y-Fran, Llanrwst, N.
Wales (SH 8280 6080) Boswell’s locality 536; m, TCD 8745 specimen with unusually curved sicula,
ludensis Zone, locality 100 Ludlow district (text-fig. 4); n, TCD 8661, holotype, a specimen in full
relief, ludensis Zone, locality 100a, Ludlow district (text-fig. 4); o, SM A60903, isolated specimen in full
relief, showing the rare slight rounding of th 1, nodule from Pragowiec, Bardo Syncline, Holy Cross
Mountains, Poland, probably nilssoni Zone; p, SM A60902, isolated specimen in low relief, nodule from
Pragowiec, probably nilssoni Zone; q, TCD 9218, ludensis Zone, locality 40, Ludlow district (text-fig.
4); r, TCD 8696 specimen in low relief with abnormal spine, ludensis Zone, locality 40, Ludlow district

(text-fig. 4). All figures x 10.

672

PALAEONTOLOGY, VOLUME 12

The length of the ventral thecal wall at th 10 is 2-50 mm. The thecal apertures approach
the horizontal, that is they are not at right angles to the long axis of the thecal tube, and
both proximal and distal thecae are strongly denticulate, almost spinose.

A single specimen (TCD 8654, text-fig. 1 d) shows a rather curious pattern of the
cortical tissue. This is composed of fine, arcuate ridges, which are mostly, but not
invariably, concave towards the ventral side of the rhabdosome. The ridges are laid down
over the dorsal wall of the rhabdosome and over the nema. In numerous instances earlier
ridge systems can be seen beneath later ones. The exposed portion of the sicula is
similarly affected, but not the interthecal septum of th 6-th 7.

Remarks. The cortical tissue on the specimen described above is clearly laid down in a
somewhat irregular manner, presumably by soft parts operating from the ventral side
of the rhabdosome. Dr. Berry has recently shown us electron microscope photographs
of this structure which he has found on Ordovician orthograptids.

Pristiograptus sp. 1 is a relatively robust form, and is difficult to equate with the many
previously described pristiograptids. In general appearance it resembles the form
figured by Pfibyl (1943) as P. cf. sardous sardous (Gortani), a flexi/is (linnarssoni) Zone
species. The Ludlow district species is, however, rather more slender and has more
closely spaced thecae, whilst the thecal angle is distinctly higher. P. kosoviensis (Boucek),
a Ludlow species, is perhaps closer to Pristiograptus sp. 1 in terms of general dimensions,
being only slightly broader. The long-ranging P. dubius stock seems repeatedly to pro-
duce narrow and broad species (and subspecies), and P. sp. 1 may reflect the laiter
tendency.

Genus monograptus Geinitz 1842
Monograptus deubeli Jaeger 1959

Plate 1 30, fig. 3

1959 Monograptus deubeli Jaeger, pp. 126-7, pi. 10, figs. 4-8.

Material. Six specimens, low to moderate relief.

Horizon and Locality, ludensis Zone, Wenlock Series; locality 1 14d, Ludlow District (see p. 676).

Description. The rhabdosome is almost straight or with gentle dorsal curvature (parti-
cularly at the proximal end), achieving a length of about 20 mm. and a distal dorso-
ventral width of L10-L25 mm. The thecal spacing varies from 12-13 in 10 mm.
proximally to 10 in 10 mm. distally. At the level of the aperture of th 1 the dorso-
ventral width is 0-90 mm.

The sicula has a length of approximately 2-00 mm. its apex reaching almost to the level
of the aperture of th 2. The dorsal wall of the sicula may have dorsal curvature whilst
the sicula as a whole expands rapidly towards its aperture. The sicula, therefore, appears
more broadly triangular than is usual in monograptids: the aperture has a diameter of
0-50 mm. (moderate relief).

Th 1 has a distinctly rounded aperture. Subsequent thecae are of simple pristiograptid
type, though a faint trace of rounding can be seen on th 2 and th 3. The rounded aper-
ture of th 1 is already formed when th 3 is half completed. Thecal overlap is about one-
half and the angle of inclination of the thecae some 30°.

Remarks. Although less well preserved our material agrees closely with that described

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 673

by Jaeger (1959) from Thuringia, particularly in the nature of the proximal end and the
shape of the sicula. M. deubeli has not previously been recorded in Britain. Dr. Jaeger
has recently obtained the species from Australia whilst one of us (R. B. R.) has identified
M. cf. deubeli in the Hassi Bedda-lA borehole of the Compagnie Frangaise des Petroles
d’Algerie, where it occurs in strata immediately overlying beds with P. bohemicus (Bar-
rande), M. incipiens Wood, and M. chimaera semispinosus Elies and Wood.

1839

71879

1890

1899

1900
71900

1900

1911

71911

1911

71935

71936
1936

71942

1943

71948

1956

1959

71965

non 1879
non 1879
non 1900
non 1900
non 1911
non 1911
non I960

Monograptus ludensis (Murchison 1839), sensu Wood 1900

Plate 1 30, fig. 2, text-figs. 2a- j, 3 a-e.

Graptolithus Ludensius Murchison, p. 694, pi. 26, fig. 2 (non fig. 1).

Graptolithus colonus Barrande; Quenstedt, pp. 198-201, pi. 150, fig. 43 X (non figs. 43/,

P, y, 2).

Monograptus sp. Holm, pi. 1, figs. 27-30.

Monograptus gotiandicus Perner, p. 12, pi. 14, fig. 22.

Monograptus vulgaris Wood, pp. 455-6 (pars), text-fig. 106, pi. 25, fig. 2.

Monograptus gotiandicus Perner; Wood, p. 460, pi. 25, fig. 7.

Monograptus colonus var. ludensis Murchison; Wood, p. 465, pi. 25, fig. 1 1.
Monograptus vulgaris Wood; Elies and Wood, pp. 378-9 (pars), text-fig. 2486, pi. 37,
figs. 10fl-c.

Monograptus cf. gotiandicus Perner; Elies and Wood, pp. 382-3, text-fig. 252, pi. 37,
fig. 8.

Monograptus colonus (?) var. ludensis (Murchison); Elies and Wood, pp. 394-5, text-fig.
262, pi. 38, figs. 9 a-c.

Monograptus vulgaris Wood; Boucek, pp. 9-10, figs. 4 i-j.

Monograptus cf. vulgaris Wood; Boucek, p. 6, fig. 1 k.

Monograptus gotiandicus Perner; Boucek, p. 6, figs. 1 a-c.

Pristiograptus gotiandicus (Perner); Munch, p. 251, pi. 3, figs. 1-3.

Pristiograptus gotiandicus (Perner 1889); Pribyl, p. 26, text-fig. 38, pi. 2, fig. 5.
Pristiograptus ( Pristiograptus ) gotiandicus (Perner); Pribyl, p. 70.

Pristiograptus ( Pristiograptus ) vulgaris vulgaris (Wood, 1900); Tomczyk, pp. 53-54,
fig. 15«, pi. 6, fig. 1, pi. 7, fig. 1.

Pristiograptus gotiandicus (Perner 1899); Urbanek, pp. 11-26, text-figs. 1-3, text-pl. 1,
pis. 1 and 2.

Pristiograptus gotiandicus (Perner), 1899; Obut, Sobolevskaya, and Bondarev, pp. 67-8,
pi. 11, figs. 4, 5.

Graptolithus ludensis Murchison; Quenstedt, pp. 192-3, pi. 150, figs. 29, 31, 32.
Graptolithus ludensis arcuatus; Quenstedt, p. 194, pi. 150, fig. 30 [= M. testis s.l . ].
Monograptus vulgaris var. /? Wood, p. 457, pi. 25, fig. 3.

Monograptus vulgaris Wood, text-fig. lOn.

Monograptus vulgaris Wood; Elies and Wood, text-fig. 248«.

Monograptus vulgaris var. curtus Elies and Wood, p. 379, pi. 37, fig. 11.

Monograptus vulgaris var. ashlandensis Berry, p. 1 1 63, fig. 2h.

Lectotype. The type slab contains at least 80 well-preserved specimens of the species. Murchison’s
original figure (pi. 26, fig. 2) appears to be diagrammatic, probably composite, and shows only a por-
tion or portions of the type slab. We have chosen GSM GSC 6584« as lectotype of the species. This
specimen is illustrated herein as text-figs. 2/and 36. The specimen figured by Murchison (1839) as pi.
26, fig. 1 is conspecific with Monograptus priodon (Bronn 1835).

Material and localities. Numerous specimens in low to moderate relief from the Ludlow District and
from North Wales (text-fig. 4). Many specimens isolated from the matrix, from nilssoni Zone nodules,
Bardo Syncline, Holy Cross Mountains, Poland.

674

PALAEONTOLOGY, VOLUME 12

Horizon. ludensis Zone in Ludlow District; ludensis and basal few metres of nilssoni Zone in North
Wales; elsewhere the species ranges into the nilssoni Zone.

Description. The rhabdosome is characteristically large and straight, often reaching a
length of 60-80 mm. and a dorso-ventral width (in relief) of over 2 mm. At 10 mm. from
the proximal end the dorso-ventral width in our material varies from 1-50 to 1-70 mm.

The proximal region shows rather variable but usually ventral curvature, whilst the
actual proximal end commonly has a most striking appearance (text-figs. 2 a, b ), often
with a ventrally curved sicula and a prominent apertural region. In these specimens the
ventral wall of th 1 is often concave, thus accentuating the rather claw-like appearance
of the proximal end. However, there is every gradation between this ‘typical’ proximal
end and the more robust type (text-figs. 2 e-g), in which th 1 may have a less concave
ventral wall and a higher angle of inclination and in which the sicula is less prominently
curved.

Th 1, and less often th 2, shows a distinct rounding of the apertural region (text-figs.
2a, e, h, i, j ) due to the development of incomplete half rings resulting in incipient lappets
(Urbanek 1959). The lappets are clearly visible on our British specimens. The sicula may
be slightly in excess of 2 mm. in length whilst its apex reaches the level of the aperture
of th 2. At th 2 the dorso-ventral width is from 0-80 to M0 mm. The thecal spacing
changes from 13 in 10 mm. proximally, to usually 10 in 10 mm. distally, but occasionally
to 8 in 10 mm.

In immature rhabdosomes (say 10-20 mm. long), thecae subsequent to th 1 or th 2
may have simple pristiograptid apertural regions, but adult specimens show a pro-
nounced infilling with late fusellar tissue of the angle between the original even aperture
and the subsequent free ventral wall (Urbanek 1959). This is clearly seen in mature
Ludlow specimens and is commonly observed in the North Wales examples (text-fig 3c).

Thecal overlap changes from rather more than one-half and up to two-thirds proxi-
mally, to three-quarters distally. However, when infilling of the apertural angle is
complete in mature specimens the rhabdosome appears almost parallel-sided and the
‘serrations’ of the ventral margin may be hardly visible and a free ventral wall non-
existent. In such material overlap is effectively complete.

Remarks. M. gerhardi Kiihne, 1955 is clearly very close to M. ludensis (Murchison),
differing only in having a higher proximal thecal count (14 in 10 mm.) and a dorso-
ventral width of 2-20 mm. at 10 mm. from the proximal end. Indeed, Jaeger (1959,
p. 64 n.) has suggested that gerhardi and vulgaris (large form, = ludensis ?) are con-
specific and reiterating this later (1964, p. 37) he added that he believed gotlandicus and
gerhardi also to be conspecific. It is possible that the variation noted above can be
ascribed to astogenetic changes (see above and Jaeger 1962, p. 37), but it may equally
represent geographical variation and short of a detailed study of gerhardi we prefer to
leave open the question of its precise relation to ludensis.

The appreciation of the conspecificity of M. ludensis (Murchison) sensu Wood, M.
gotlandicus Perner, and M. vulgaris Wood (pars) solves a number of problems concern-
ing the geographical records of this species. Thus M. gotlandicus was widely recorded
and common on the continent ( nilssoni Zone) and only doubtfully recognized in Britain.
The converse holds for M. vulgaris (pars). The latter has, however, been frequently and

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 675

a

c

e f

text-fig. 3 a-e. Monograptus ludensis (Murchison 1839) sensu Wood 1900: a, BU 1466 (a), specimen
figured by Elies and Wood (1911), pi. 37, fig. 10 b as M. vulgaris Wood on the same slab as the type
specimen, pi. 37, fig. 10a; b, GSM GSC 6584a, proposed lectotype, a specimen in full relief on the type
slab showing critical details of the proximal end, locality: Llanfair, Montgomeryshire; c, BU 1496,
specimen figured by Elies and Wood (191 1), pi. 38, fig. 9b as M. colonus (?) var. ludensis (Murchison);
d , BU 1466 ( b ), the type specimen of M. vulgaris Wood, figured by Wood (1900) as pi. 25, fig. 2 and
by Ellis and Wood (1911) as pi. 37, fig. 10a; e, GSM Zp 4003 b, somewhat compressed specimen,
ludensis Zone, locality Brynsylldy, Llanrwst, N. Wales (SH 8202 6164), Boswell’s locality 472; /,
Pristiograptus jaegeri sp. nov. GSM Zp 5720, ludensis Zone, locality given under explanation of text-

fig. 21. All figures x 2\.

widely misidentified, and is probably not as common in Britain as the literature would
suggest.

The form described by Elies and Wood as Monograptus vulgaris curtus nom. nov.
(for Wood’s M vulgaris var. /?) would seem to be a pristiograptid, the type lacking the

676

PALAEONTOLOGY, VOLUME 12

rounding of th 1 so typical of M. ludensis. We feel unable to comment at this stage on
the position of Pristiograptus curtus in relation to other pristiograptids.

STRATIGRAPHICAL DISTRIBUTION

The stratigraphical distribution of graptolites we have collected from the Wenlock and
lowest Ludlow of the Ludlow district is indicated in text-fig. 4. The locality numbers are
those of Holland et al. ( 1 963), except that a few subdivisions have been made as follows :

Locality 114 (Holland et al. 1963, p. 165) is a track section continuing some 500 m.
through Wenlock Shale. Our more precise Locality 1 1 4a refers to National Grid
Reference SO 45387228, whereas Locality 114b lies about 120 m. north-eastwards at
45487232, and Locality 1 14c is at 45667247, some 230 m. further north-eastwards. About
160 m. beyond this an exposure within the Wenlock Limestone at Grid Reference
45817253 is referred to as Locality 1 14d.

The north-eastern arm of the branching track with its exposures of Wenlock Shale
referred to as Locality 100 in Holland et al. (1963, p. 164) may be followed up slope
where some exposures of the lower part of the Wenlock Limestone are available. The
lowest of these, at the base of the Wenlock Limestone, is here referred to as Locality
100a (Grid Reference 44887244).

As will be appreciated from text-figure 4, some localities in the Wenlock Shale have
yielded no graptolites or but a sparse record, whereas others, such as Localities 79, 91,
and 40, are relatively rich. Holland et al. recorded no graptolites from the Wenlock
Limestone but the present investigation has revealed a sparse fauna in its lower part.
These same authors, (Holland et al. 1963, p. 108), record a graptolite fragment from the
Lower Elton Beds of Locality 69, but this and other exposures within these beds were
searched again with very little success. We have obtained but a further single specimen
which we have identified as Monograptus ? varians Wood.

Approximately the lower half of the Wenlock Shale in the Ludlow district falls within
the lundgreni Zone and is characterized by an assemblage of relatively abundant
Monograptus flemingii, associated with Pristiograptus dubius and another pristiograptid
here referred to as Pristiograptus sp. 1.

The ' nassa j dubius Interregnum" of Jaeger (1959) may be identified as a relatively thin
horizon in which M. flemingii and Pristiograptus sp. 1 have disappeared; Gothograptus
nassa is present; but Monograptus ludensis has not yet appeared. It is immediately suc-
ceeded by strata yielding M. ludensis; but between this first appearance of the species
and its further occurrences there is a considerable thickness of strata from which we
have recorded no graptolites at all (see text-fig. 4). The factors responsible for the
impoverishment of graptolite faunas at the level of Jaeger’s "Interregnum", and marked
there by the restricted fauna of G. nassa and P. dubius , may perhaps be reflected here in
the absence of graptolites through the basal part of the ludensis Zone, which we have
taken as beginning at the first appearance of the index species just over 100 m. below
the base of the Wenlock Limestone.

The characteristic assemblage of the ludensis Zone as seen in the highest part of the
Wenlock Shale is of M. ludensis itself together with Pristiograptus jaegeri. Both G. nassa
and M. dubius are also present at first.

Finally, it is important to note that the ludensis-jaegeri association is now known to

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GR APTOLITES 677

Wen lock Shale

50 m

M. flemin gii
Pristiograptus sp l
P dubius

G. nassa

M. lude n s i s
P jaegeri sp.nov.

H. lawsoni sp. nov
M. d e u b e 1 1

U>

o

X)

n>

i'll

l- ■

Wenlock

Limestone

LElton

Beds

D —
O o

o

>

30

m

>

M varians

<

m

xi

2

CO

lundgreni

/'

ludensis

'n il ssom

o p

■o

M f.flemingii
M f. aff ele g a n s
G. nassa
PI. aff. dubius
P dubius
P pseu d o du biu s
P jaegeri sp.nov.
M. luden si s

o

70

L. Mottled U Mottled

Mds. Mds.

L. Nantglvn FI aas Gp

disturbed bed
U N. F G

o ^

< >
B r~

® m

cn

WENLOCK

LUDLOW

text-fig. 4. Range chart of graptolites in the Ludlow, Malvern, and North Wales (Conway Valley)
districts. The section given here is that measured at Ty-mawr Farm (N.G.R. SH 820 682), Eglwysbach,
Conway Valley and the graptolite occurrences are local and hence can be directly related to this section.
However, the general stratigraphy and graptolite ranges are applicable to the whole of NW. Denbigh-
shire. In the North Wales graptolitic facies approximate peaks are indicated by fine dotted lines.
Dot/dash lines indicate approximate position of the nassa/dubius Interregnum. Alternating lined and
stippled beds are mudstones; ‘bricks’ are limestones; ‘broken bricks’ are calcareous mottled mud-
stones; and fine-lined beds are shales. P. nilssoni Zone species are omitted from the North Wales side
of the chart, whereas these only are shown for the Malverns. Abbreviations: Mds. = Mudstone;
Gp. and G. = Group; spec. = specimens. Details of graptolites (in the North Wales section) not
referred to in the text are as follows: M.flemingii aff. elegans Elies, Plectograptus (?) aff. dubius Boucek
and Munch, Pristiograptus pseudodubius (Boucek).

678

PALAEONTOLOGY, VOLUME 12

continue into the basal part of the Wenlock Limestone. Thus, we have factual
evidence that the upper part of the Wenlock Shale and part at least of the Wenlock
Limestone belong in the ludensis as distinct from the lundgreni Zone. The consequences
of this in terms of stratigraphical nomenclature are discussed in the next section of
this paper.

Also within the Wenlock Limestone, we may note the occurrence of Monogrciptus
deubeli at a somewhat higher level than in Thuringia from whence Jaeger first described
this species.

As mentioned above, it is unfortunate that the Lower Elton Beds of the Ludlow
district have still failed to provide conclusive evidence of their position in the graptolite
zonal sequence. We have, through the kindness of Mr. J. S. W. Penn, Miss J. Vinni-
combe, and Mr. D. G. A. Whitten, been allowed to examine a collection of
Silurian graptolites from the Malvern Hills assembled at Kingston-upon-Thames College
of Technology. Mr. Penn will undoubtedly be referring to the full results of our examina-
tion elsewhere, but we note here the presence of Monograptus various in the Lower Elton
Beds. The number of specimens is small and, in spite of the uniformity of shelf facies
Ludlow successions throughout the Welsh borderland, there remains the possibility
that the Lower Elton Beds of the Malverns are not of precisely the same age as those at
Ludlow. So, as M. various has never been recorded in pr e-uilssoui strata, here is slight
evidence that the Lower Elton Beds belong to the nilssoni Zone and we have added this
to our chart.

We have also been able to examine, and here report on, four graptolite specimens
obtained by Professor P. J. Lesperance (University of Montreal, Canada) from the
Lower Elton Beds of Millichope, Shropshire, which were forwarded to us by Dr. J. H.
Shergold (Canberra, Australia). They are: an indeterminate distal fragment; the part
and counterpart of a form probably referable to P. dubius ; and a specimen referred to
Monograptus sp. cf. various various Wood. The material is not well preserved and the
last mentioned specimen does not exhibit clearly the proximal thecal ‘hooks’ of various
although the dimensions of the rhabdosome accord in every way with that species. It
could possibly be assigned to M. aff. ludensis. Thus, whilst in no way providing of itself
an unequivocal age for the Lower Elton Beds, this material adds further slim evidence
to that from the Malverns.

Finally we would note the occurrence of M. uncinatus orbatus Wood, a nilssoni Zone
species, from the Lower Elton Beds of the May Hill Inlier (Birmingham University,
specimen no. My 82m).

Text-fig. 4 also includes a summary of uppermost Wenlock graptolite occurrences in
the Denbighshire Moors in North Wales. We provide this as an illustration of a typical
basin facies graptolitic sequence in contrast to that of the Ludlow district; and, more-
over, a basin sequence which has been subject to modern study by the Institute of
Geological Sciences. The Ludlow district is to be regarded as of shelf facies in terms of
higher Silurian stratigraphy, though certainly in a position marginal to the basin or
geosyncline. In the North Wales section, as in that at Ludlow, the ranges of Monograptus
ludensis and Pristiograptus jaegeri replace that of the original M. 'vulgaris'. Although
Gothograptus nassa is common within the basal M. ludensis Zone, its maximum occur-
rence is within a 'nassaldubius Interregnum’, of the same order of thickness as that at
Ludlow, which may be recognized immediately below the first occurrence of M.

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 679

ludensis. This interregnum is thus again in the same stratigraphical position as that
originally described by Jaeger (1959).

It is now clear that the sequence from the lundgreni Zone (with its characteristic occur-
rence of M. flemingii), through a ‘ nassajdubius Interregnum’, to the ludensis Zone
(characterized by both M. ludensis and Pristiograptus jaegeri ), and then into a more
varied nilssoni Zone assemblage, can be widely recognized in Europe. The comparison
with the very carefully described sequence from Thuringa is evident from Jaeger’s
diagram of graptolite ranges (1959, p. 39) and two of the present authors (C. H. H. and
P. T. W.) have had the opportunity of seeing the same sequence in the Polish sections of
the Holy Cross Mountains.

THE WENLOCK LUDLOW BOUNDARY

The question of the recognition of the boundary between the Wenlock and Ludlow
in the light of the stratigraphical facts referred to above must now be considered
against the historical background. A short review of the problem is given by Holland
(in press).

Murchison (1833) was originally confused between the two Silurian limestones at
Wenlock and Aymestrey respectively, and his Lower and Upper Ludlow Rock were at
first separated by the so-called Wenlock Limestone. This confusion was eliminated the
following year when the relevant part of his stratigraphical table (Murchison 1834)
showed the Lower Ludlow Rock following upon the Wenlock and Dudley Limestone
and succeeded by the Aymestry and Sedgley Limestone. Wood (1900), in her classical
account ‘The Lower Ludlow Formation and its Graptolite-Fauna’, referred to Mur-
chison’s line of demarcation, but noted (p. 421) that he himself had admitted that the
‘Lower Ludlow was simply an upward prolongation of the Wenlock Shale’. In some
districts he thought it impracticable to separate them. She referred also to Lapworth’s
(1880, p. 48) comment that the division was probably made ‘less from a palaeontological
than from an aesthetic point of view’. Wood’s own palaeontological studies had con-
firmed that ‘where there is a lithological transition between the Wenlock and Ludlow
Beds there is also a palaeontological transition’ (p. 421). Nevertheless, she was able to
draw a palaeontological line between the two ‘of considerable stratigraphical and prac-
tical value’ (p. 421). Working with Elies (1900), who had similarly described the grapto-
lite fauna of the Wenlock, Wood (1900, p. 421) compiled a short list of the comparative
graptolitic characteristics of the two divisions. Thus, the Wenlock was characterized by
the presence of Cyrtograptus and monograptids of the flemingii type, both of which were
absent in the Lower Ludlow. The Lower Ludlow, on the other hand, contained mono-
graptids of the colonus (Barrande) type as well as spinose forms such as M. chiniaera
( Barrande), none of which was present in the Wenlock.

More significantly for the future development of Wenlock/Ludlow biostratigraphy.
Wood divided the ‘Lower Ludlow Beds’ into five graptolite zones, of which the lowest
was that of M. vulgaris. This last followed upon the highest of the Wenlock zones defined
by Elies, that of Cyrtograptus lundgreni Tullberg. The vulgaris Zone was devoid of
graptolites in the type Ludlow district, though the fauna was to be found in both the
Builth and Long Mountain districts where Wood also established her detailed strati-
graphy.

The M. ‘ vulgaris ’ [= ludensis ] Zone has since been taken internationally as the base

680

PALAEONTOLOGY, VOLUME 12

of the Ludlow Series and numerous references from Central and Eastern Europe could
be quoted. The authoritative and comprehensive work of Jaeger (1959) on the graptolitic
Silurian of Thuringia has already been referred to above. In Britain, Boswell ( 1949), for
example, used it (though evidently with some difficulty) in his lengthy treatment of the
Silurian rocks of North Wales. In contrast, those working in the shelf facies of the higher
Silurian, where the Wenlock Limestone is developed, have followed Murchison’s revised
stratigraphy with the Ludlow beginning immediately above that limestone. A correla-
tion table and comprehensive reference list for the British Ludlow is given by Holland
et al. (1963).

Nevertheless, since Watts (1925) equated the M. ‘ vulgaris ’ [= ludensis] Zone of the
Long Mountain with the ‘Wenlock Limestone Stage’ of the Wenlock-Ludlow Area
(Watts 1925, p. 346) and went on to report that above the Wenlock zones ‘comes the
zone of M. vulgaris which Miss Wood (Dame Ethel Shakespear) has shown to be
equivalent of the Wenlock Limestone’ (p. 394), a sense of disquiet has tended to affect
those obliged to rely upon the assumed correlation of the barren beds above the
Wenlock Limestone with the ‘ vulgaris ' [ = ludensis ] Zone of the graptolitic geosynclinal
facies.

The Summary of Progress of the Geological Survey of Great Britain for 1926, published
in 1927, referring to the Much Wenlock district, suggested (pp. 42-3) that the ‘ vulgaris ’
[= ludensis ] Zone extended some 60 m. below the Wenlock Limestone whilst a P. nilssoni
zonal assemblage had been found about 18 m. above the limestone. An abundance of
Gothograptus nassa in the 60 m. of shales below the Wenlock Limestone ‘would, other
things being equal, be taken as clearly indicative of the zone of Monograptus vulgaris.
The correspondingly low position of the localities indicating the zone of Cyrtograptus
lungreni supports this view.’ Das Gupta (1933), however, found the range of Gotho-
graptus nassa at Wenlock Edge and in North Wales to be ‘closely comparable with its
occurrence abroad, as, for example, in Bohemia, where it ranges from the highest beds
of the zone of M. testis (= zone of C. lundgreni in Great Britain) to the lower beds of
the zone of M. nilssoni ’. Das Gupta’s comments were taken into account by the authors
of the Shrewsbury memoir (Pocock et al. 1938), where Robertson and Stubblefield
(p. 102) wrote as follows:

Subsequent ... to the completion of the survey of the Shrewsbury Sheet, Dr. Das Gupta, working
in the Long Mountain area, where there is no Wenlock Limestone, has found M. vulgaris in the
C. lundgreni Zone, and associated with the zone-fossil. He has also collected M. vulgaris from beds a
short distance above the Wenlock Limestone of Wenlock Edge, at a point some 9 miles south-west
of Much Wenlock. Manifestly, these discoveries are still not sufficient to establish whether the Wenlock
Limestone belongs to the C. lundgreni or the M. vulgaris Zone, or to both, and until that is done, it
should be clearly recognized that the position of the Ludlow-Wenlock boundary in the graptolite suc-
cession remains unsettled.

At this point we may simply refer back to the introduction to this paper, where
reference is made to the standard locality (Pitch Coppice) for the base of the Ludlow
Series. We accept this marker point in the shelf facies for the Wenlock/Ludlow boundary
(but see later). We are now in possession of new biostratigraphical facts as summarized
in text-fig. 4 above and our commentary upon it in the preceding section. We now know
that the M. ludensis Zone begins some 100 m. below the base of the Wenlock Limestone

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GR APTOLITES 681

in the Ludlow District and that some at least of that limestone is definitely to be included
therein.

Consequently, a difficult situation exists. The ‘ vulgaris ’ [= ludensis] Zone has been
widely regarded as the base of the Ludlow and yet we are convinced that the Wenlock
Limestone must remain in the Wenlock Series and not the Ludlow Series. Thus, we
have advocated that workers in the graptolitic facies should now take as the base
of the Ludlow Series the base of the nilssoni Zone (Holland, Rickards, and Warren
1967). Martinsson (1967) has already found this helpful in his ostracode correlations.

But though this course of action will restore stability and allow for reasonable accu-
racy of correlation from one facies to another, it does not dispose of the final problem
of the definition of the base of the Ludlow Series at a standard section in the Ludlow
district. As explained above, we still do not know that the base of the Lower Elton Beds
in that district coincides with the base of the nilssoni Zone. It is still possible that the
whole of the Lower Elton Beds are, as was originally assumed, of ‘ vulgaris ’ [= ludensis]
age or, again, the base of the nilssoni Zone may come within or even below these beds.
If the first of these three possibilities obtains there will be no problem as the base of
the Ludlow Series, as defined by Holland et al. (1963), will coincide with the biostrati-
graphical horizon at the base of the nilssoni Zone already advocated for use in correla-
tion. We have indicated above that there is some evidence from the Malvern Hills that
this is indeed the case.

If, on the other hand, fresh finds of graptolites or other aids to correlation do even-
tually demonstrate that the base of the nilssoni Zone comes at a higher level within the
Lower Elton Beds, we shall be left with a Wenlock/Ludlow boundary, as at present
defined, which comes within the ludensis Zone, though of course high in that zone. This
situation may not commend itself when the inevitable progress of international strati-
graphical procedure leads to a decision on this particular boundary, and it may prove
necessary to accept a slight readjustment of the base of the Eltonian to coincide (if
possible) with the base of the nilssoni Zone.

Our present study cannot contribute further to this particular problem and we can
only reiterate that we recommend the acceptance of the nilssoni Zone as the base of the
Ludlow Series in the graptolitic facies, and we are satisfied that any resulting correlations
from one facies to another will then in any event be close to the truth.

REFERENCES

berry, w. b. n. 1960. Early Ludlow Graptolites from the Ashland Area, Maine. J. Paleont. 34, 1158-63.
boswell, p. G. H. 1949. The Middle Silurian rocks of North Wales. London.

boucek, b. 1935. O silurske faune od Stlnavy (Zap. od Plumlova) na Drahanske vysocine. Casop.
vlast. spot, musej. Olomouc. 48, (3-4).

1936. Graptolitova fauna ceskeho spodniho ludlowu. Rozpr. II, tr. Ceske Akad. 46 (16).

- — -and munch, A. 1952. Central European Retiolites of the Upper Wenlock and Ludlow. Sb.
ustred. Ust. geol. 19, 104-51.

das gupta, t. 1933. The Zone of Monograptus vulgaris in the Welsh Borderland and North Wales.
Proc. Lpool. Geol. Soc. 16, 109-15.

decker, c. E. 1935. Graptolites from the Silurian of Oklahoma. J. Paleont. 9, 434-46.

1935a. Some tentative correlations on the basis of Graptolites of Oklahoma and Arkensas. Bull.

Amer. Ass. Petrol. Geol. 20, 301-11.

C 6940

Yy

682

PALAEONTOLOGY, VOLUME 12

eisenack, a. 1951. Retioliten aus dem Graptolithengestein. Paleontographica, 100A, 129-63.
elles, g. l. 1900. The Zonal Classification of the Wenlock Shales of the Welsh Borderland. Q. Jl.
geol. Soc. Loud. 56, 370-414.

and wood, e. m. r. 1901-18. Monograph of British Graptolites. Palaeontogr. Soc. [Monogr.]

Holland, c. h. (in press). Problems of classification and correlation in the Wenlockian, Ludlovian,
and post-Ludlovian pre-Gedinnian stratigraphy of the British Isles. Symposium volume of Third
International Symposium on the Silurian-Devonian Boundary, and the Stratigraphy of the Lower and
Middle Devonian, Leningrad, USSR, 1968.

lawson, j. d. and walmsley, v. g. 1963. The Silurian rocks of the Ludlow district, Shropshire.

Bull. Br. Mas. not. Hist. Geol. 8, 93-171.

— rickards, r. b. and warren, p. t. 1967. The position of the Wenlock/Ludlow boundary in the
Silurian graptolite sequence. Geol. Mag. 104, 395-7.

holm, g. 1890. Gotlands Graptoliter. Bihang Svenska Vet.-Akad. Handl. 16, Afd. 4, no. 7, 1-34.
jaeger, h. 1959. Graptolithen und Stratigraphie des jiingsten Thiiringer Silurs. Abh. deutsch. Akad.
1 Viss. Berl., Kl. Chem. Geol. Biol. 1959 (2), 1-197.

1964. Der gegenwartige Stand der stratigraphischen Erforschung des Thiiringer Silurs. Abh.

deutsch. Akad. Wiss. Berk, Kl. Bergb. Hiittenw. Montangeol. 1964 (2), 27-51.
kuhne, w. g. 1955. Unterludlow-Graptolithen aus Berliner Geschieben. Neues Jb. Geol. Paldont. Abh.
100 (3), 350-401.

lapworth, c. 1873. On an improved classification of the Rhabdophora. Geol. Mag. 10, 500-4,
555-60.

1880. On the geological distribution of the Rhabdophora. Ann. Mag. Nat. Hist. (5) 5, 45-62.

martinsson, a. 1967. The succession and correlation of ostracode faunas in the Silurian of Gotland.
Geol. For. Stockh. Fork. 89, 350-86.

m‘coy, f. 1851-5. Description of the British Palaeozoic fossils in the Geological Museum of the University
of Cambridge. 661 pp. London.

munch, a. 1942. Die Graptolithenfauna des unteren Ludlow von Ronneburg und Umgebung. Beitr.
Geol. Thiiring. 6, 241-66.

Murchison, r. i. 1833. On the sedimentary deposits which occupy the western parts of Shropshire and
Herefordshire, etc. Proc. geol. Soc. Load. 1, 474-7.

— 1834. On the Structure and Classification of the Transition Rocks of Shropshire, Herefordshire,
and part of Wales, etc. Proc. geol. Soc. Lond. 2, 13-18.

1839. The Silurian System, xxxii+768 pp. London.

1854. Siluria. xv+523 pp. London.

1859 Siluria. 3rd ed., London.

obut, a. m., sobolevskaya, r. f., and bondarev, v. i. 1965. [Graptolites of the Silurian of Taimyr.]
Acad. Sci. U.S.S.R., Siberian Section, Inst. Geol. Geogr. 1-119 (in Russian).
perner, j. 1894-9. Etudes sur les Graptolites de Boheme. Prague.

pocock, r. w., whitehead, t. h., wedd, c. b., and robertson, t. 1938. Shrewsbury District including
the Hanwood Coalfield. Mem. geol. Surv. Eng. and Wales. Sheet 152, New Series. 297 pp.
pribyl, A. 1943. Revision aller Vertreter der Gattung Pristiograptus aus der Gruppe P. dubius and P.
vulgaris aus dem bohmischen und auslandischen Silur. Rozpr. ceske Akad. 53, 1-49.

1948. Bibliographic index of Bohemian Silurian graptolites. Knihovna St at. geol. list. Csk.

Repub I iky, 22, 1-96.

quenstedt, f. a. 1879-1881. Petrefactenkuude Deutschlands 6, Korallen, 1-1093.
ruedemann, r. 1947. Graptolites of North America. Mem. geol. Soc. Am. 19, 1-652.
tomczyk, h. 1956. Wenlok i Ludlow w Synklinie Kieleckiej gor swietokrzyskich. Inst. Geol.Prace, 16,
1-129.

- 1962. Problem Stratygrafii Ordowiku i Syluru w Polsce w Swietle Ostatnich Badari. Inst. geol.
Prace, 35, 134 pp.

urbanek, a. 1959. Studies on Graptolites. I. Development and Structure of Pristiograptus gotlandicus
(Perner). Acta palaeont. pok 4, 1 1-26.

warren, p. t., rickards, R. b., and Holland, c. h. 1966. Pristiograptus liulensis (Murchison 1839) —
its synonomy and allied species — and the position of the Wenlock/Ludlow boundary in the Silurian
graptolite sequence. Geol. Mag. 103, 466-7.

HOLLAND, RICKARDS, AND WARREN: THE WENLOCK GRAPTOLITES 683

watts, w. w. 1925. The geology of South Shropshire. Proc. Geol. Assoc., Lond. 36, 321-63.
wood, e. m. r. 1900. The Lower Ludlow Formation and its Graptolite-Fauna. Q. Jl. geol. Soc. Lond.
56, 415-92.

C. H. HOLLAND

Trinity College
Dublin 2
Ireland

R. B. RICKARDS

Sedgwick Museum
Cambridge

P. T. WARREN

Institute of Geological Sciences
Ring Road Halton

Typescript received 24 February 1969 Leeds 15

PATTERNS OF TAXONOMIC AND ECOLOGICAL
STRUCTURE OF THE SHELF BENTHOS DURING

PHANEROZOIC TIME

by JAMES W. VALENTINE

Abstract. The taxonomic and ecological structure of the shelf biota are intimately related at the species-
population levels. Early Paleozoic faunas contained relatively few species representing relatively many higher
taxa, and ecosystems were relatively generalized. Medial and late Paleozoic faunas contained more species
representing fewer higher taxa, and ecosystems were relatively specialized. This suggests that, as higher taxa
became extinct, they were not replaced except at lower taxonomic levels; diversification was proceeding through
increasing specialization. After Permo-Triassic extinctions, rediversification was chiefly confined to low taxo-
nomic levels. Late Mesozoic and Cenozoic diversification at lower taxonomic levels has been remarkably great,
resulting not only from increasing specialization at the population level but from a marked increase in pro-
vinciality due to rising latitudinal temperature gradients on the shelves and to the fragmentation and isolation
of shelf environments by continental drift.

This paper examines the historical relationships between the ecological and taxonomic
structures of the marine biosphere, and attempts to account in a general way for the
patterns of their evolution. Each of these structures is hierarchic. The units composing
the levels of the ecological hierarchy include individuals, populations, communities,
and provinces, while the units composing the levels of the taxonomic hierarchy are such
categories as species, genera, and families.

There has been relatively little theoretical discussion of the evolution of these struc-
tures for marine invertebrates, yet the geological record of skeletonized taxa of the
shallow marine invertebrate benthos is longer and more complete than for any com-
parable group of organisms. This paper therefore deals with the rich and lengthy
record of shallow marine environments.

This restriction to a specific group of communities has some special advantages. The
diversity pattern for the world at large is obviously very much influenced by the deploy-
ment of organisms into new environments, such as the invasion of the terrestrial habitat
by vertebrates. By restricting the data to a limited group of communities it may be pos-
sible to investigate the patterns of diversity changes within ecosystems.

Much of the structural evolution which the taxonomic and ecological hierarchies have
undergone is a product of the diversification and extinction of species. There has been
much discussion of the patterns of taxonomic diversifications and extinctions through
geologic time, especially of higher taxa, and ecological relations are commonly invoked
to account for these patterns, particularly for extinctions. The processes of diversification
assumed herein are those of the synthetic theory of evolution based on Darwinian
selection and upon modern genetic concepts. Speciation and the origin of higher taxa
have been discussed from this viewpoint in a number of larger works (for example
Huxley 1942; Mayr 1963; Rensch 1947; and Simpson 1953). As diversity rises, there
must be a mechanism of accommodation of the new forms in ecological systems;
such mechanisms are discussed by Klopfer (1962), MacArthur and Wilson (1967), and

[Palaeontology, Vol. 12, Part 4, 1969, pp. 684 709.]

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 685

Miller (1967), among others. Possible causes of extinction, and hypotheses of the processes
of extinction that have operated to create the Permo-Triassic faunal change, have been
reviewed by Rhodes (1967).

THE ECOLOGICAL STRUCTURE OF THE BIOSPHERE

The ecological hierarchy is regarded as being composed of the levels that are depicted
in text-hg. 1. This paper is chiefly concerned with the functional aspects of the hierarchy,

GENETIC H

ERARCHY

ECOLOGICAL HIERARCHY

LEVEL

UNIT

COLLECTIVE

DESCRIPTIVE

UNITS

FUNCTIONAL

UNITS

*2

\

F3 —

-«*» Marine Shelf Biota

Shelf Realm of
the Biosphere

High

F1

\

\

^ —

r-E®» Province

Provincial System

Gene Pool

\

\

A1

r^®» Community

Community System

(Ecosystem)

Genotype

- \

\

Gene Pool —

Population

(Deme, Species)

Population System

(Niche)

Functional
Genetic Unit

\

Genotype —

Individual

Ontogenetic System

Low

*! Collection of gene pools

p

* Collection of gene pool collections

-*3 Col lection of col lected gene poo! collections

text-fig. 1. Some levels of organization in the ecological hierarchy employed in this paper

(after Valentine 19686).

that is, with the interacting systems of organisms and environments. From the highest
level down, each functional system is composed of subsystems representing the systems
of the next lower level. The lowest functional level in the figure, that of the individual,
is certainly capable of further subdivision into sorts of functions of ‘unit characters’,
each underpinned by a system of genes and its regulators. For the most part, however,
the present discussion concerns population and higher levels.

It is convenient to consider the ecological units in terms of the environment with
which they interact. Hutchinson (1957, 1967) has developed a formal conceptual model
that treats the environment as a multi-dimensional region (see also Simpson 1944, 1953).
Only an informal treatment, based on Hutchinson’s model, is required here. If each
separate environmental parameter is visualized as a single geometric dimension of this
region, then all possible environments are represented by the resulting multi-dimensional

686

PALAEONTOLOGY, VOLUME 12

space or hyperspace, which contains as many dimensions as there are possible environ-
mental parameters. The space extends along each dimension to the physical limits of
each parameter. It is assumed that this multi-dimensional environment model is stan-
dardized by having each axis allotted an arbitrary but permanent direction to form an

text-fig. 2. Highly diagrammatic representation of some aspects of environment-organism relations,
visualized as a multi-dimensional space, of which each dimension is some environmental factor,
physical or biotic. Each point within the lattice represents a unique combination of factors. Only three
of the many dimensions are depicted, a, the total possible range of all environmental factors repre-
sented as a multi-dimensional lattice, b, the portion of the environment that actually exists on earth,
the biospace ; it is available for occupation by organisms, c and d, the region of environmental space
that coincides with factors tolerated by an organism and that is bounded by its limits of tolerance — -
the ecospace of the organism. Only a portion of the ecospace is realized (c) ; the remainder is prospective
ecospace (d) that may be inhabited if the environment fluctuates so as to include more of that portion
of the lattice. The ecospace concept may be expanded to population, community, province or biosphere
levels, and to species, genus, family, and higher taxonomic levels.

environmental hyperspace lattice , hereinafter called simply a lattice for brevity. Only a
certain portion of the total possible lattice (the prospective lattice) actually represents
conditions of the environment. This ‘realized ecological hyperspace’ may be called
biospaee (text-fig. 2), a term employed in a similar but less generalized sense by Doty
(1957).

For any organism there is some more or less small volume (actually a hypervolume)

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 687

within the lattice corresponding to the range of environmental conditions under which
it may live. This functional hypervolume will be called the ecospace of that organism
(text-fig. 2). Each population also has its own ecospace, which is the hypervolume of
its niche within the lattice. Indeed, the ecological units at all levels have ecospaces. A
community ecospace is the multi-dimensional model of its ecosystem, and a provincial
ecospace is the model of the provincial system. Although the highest functional level
standing above that of the provincial system is the level of the biosphere, the system
of the shallow marine realm is being used here in its place as a matter of simplicity,
and this realm has its own ecospace. The total ecospace that an organism or other
ecological unit may utilize if it is physically available may be called the prospective
ecospace , while the portion of the ecospace that actually overlaps with realized biospace
may be called the realized ecospace. These terms are modelled on the discussion of Parr
(1926) and Simpson (1944, 1953).

Dimensions of the lattice which have special properties are those that represent the
real dimensions of space, in which discontinuities occur that permit the occupation of
similar functional regions in different geographic regions (Miller 1967) and of time, in
which the changing shapes and sizes of ecospaces and of biospace are perceived.

The structure of the ecological hierarchy may be illustrated by considering just one
level, for example the community level. Community ecospace is composed of the eco-
spaces of all the niches of the component populations, and includes some dimensions
that are not niche properties but are organizational properties of the ecosystem. The
size of the community ecospace, measured by the number of dimensions occupied and
the extent of occupation along each dimension, depends upon the sizes of the com-
ponent niche ecospaces and to a small extent upon the organizational properties. Into
a community ecospace of a given size, a relatively large number of small niches or a
relatively small number of large niches may be packed. All niches in a community
ecospace overlap to some degree, for all share a common tolerance for certain salinity
ranges, for example, and for certain oxygen concentrations, and for other parameters.
The more that niches overlap, other things being equal, the more populations that can
be packed into a community ecospace of a given size (Klopfer 1962; Miller 1967).

Consider, then, a community (A) composed of relatively few populations that have
very large niches that overlap only narrowly on the whole. The animals tend to be rather
generalized feeders, so that energy flows in relatively broad streams through the trophic
levels. This community, though of low diversity, may displace a large biospace in the
lattice, that is, may have a large ecospace. Consider another community (B) composed
of many populations of different species that tend to have very small niches which
overlap broadly on the average. The animals are highly specialized with relatively
narrow ranges of food sources, so that energy flows through the trophic levels in
relatively discrete paths along chains of organisms that tend to be rather isolated
owing to their high specialization. Energy flow is not like a stream but more like a
shower that breaks up into numerous jets. A community of this sort, though rich in
species, may displace no more biospace in the lattice than community (A) and may
displace considerably less.

These communities have vastly different structures in the lattice, and yet it seems
possible for one to evolve from the other. They may thus represent relatively early (A)
and advanced ( B ) stages in the evolution of a community ‘lineage’ that has inhabited

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PALAEONTOLOGY, VOLUME 12

a similar biotope through its history. In this event the ecospaces of the two communities
will approximately coincide, although the way in which each community biospace is
occupied by niches is different. The community structure has evolved.

Structural states of ecological systems at other levels may be described in an ana-
logous way. All the systems evolve by changes in the quality, relative proportions, and
diversity of their subsystems (Valentine 19686). Thus evolution of ecological systems
need not involve organic evolution, but may result merely from the readjustment of
existing populations in new patterns of association. However in the present discussion
the chief interest lies in changes that are based upon organic evolution, upon changes in
gene frequencies within populations that produce changes in niches, and upon the
accommodation of the changed niches in ecosystem structures. Enough is now known
of these processes to permit the construction of a provisional model of the diversification
of ecosystems. But before proceeding to the model, it is appropriate to examine the main
patterns of taxonomic structure during the Phanerozoic.

THE TAXONOMIC STRUCTURE OF THE BIOSPHERE

The taxonomic hierarchy is too well known to require any general remarks. For pur-
poses of this paper only a few levels need be considered: phylum, class, and order, which
will be called ‘higher’ taxonomic categories; and family, genus, and species, which will
be called ‘lower’ categories. It is possible to visualize the ecospace of any genus as com-
posed of the ecospaces displaced by all its component species, and the ecospace of a
family as composed of all the generic ecospaces, and so on. Thus defined, the ecospace
of a higher taxon displaces the actual regions of the lattice that have been occupied by
the members of that taxon. Thus the taxonomic hierarchy possesses a precise structure
at any time. This structure changes through time in well-defined patterns.

The main trends of evolution of the taxonomic structure may be characterized by con-
sidering the trends of diversity among higher and lower taxa through geologic time.
The fossil record of diversity, however, is certainly biased. An important source of bias
is the differential preservation of taxa. It seems possible to use the skeletonized taxa that
are best represented as a sample, from which to attempt to generalize to the entire biota.
The basic data from which generalizations will be attempted are the records of easily
fossilized shallow benthonic taxa of nine phyla: Protozoa, Porifera, Archaeocyatha,
Coelenterata, Ectoprocta, Brachiopoda, Mollusca, Arthropoda, and Echinodermata.
The ranges of these phyla and of their taxa are taken chiefly from the Treatise on
Invertebrate Paleontology (ed. Moore 1 953—67), the Fossil Record (ed. Harland et al.
1967), and the Russian Osnovy Paleontologii (Orlov 1958-64).

As the assignment of groups of organisms to taxonomic categories involves a large
element of subjectivity, it is fair to ask to what extent the trends in taxonomic diversity
are real. In the first place, if one constructs a hierarchical classification of fossils that
appear at different times, the average time of appearance of higher taxa will be earlier
than that of lower, simply because some of the lower taxa appeared later than others,
but none appeared earlier than the higher taxa to which they belong. The mode of first
appearance should shift progressively towards the present at lower and lower taxonomic
levels ( Simpson 1953, pp. 237-9). Similarly, the mode of highest diversity will tend to shift
towards the recent at progressively lower levels provided that the earlier taxa at each

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 689

level persist or are replaced. These considerations account for such shifts in the mode
of appearance and diversity in text-fig. 3.

Secondly, there is no doubt that the present data contain monographic artifacts (for

text-fig. 3. Stratigraphic variation in diversity of higher taxa of well-skeletonized
marine shelf invertebrates. Data chiefly from Harland et al. (1967) and Moore

(1953-7).

an instructive example see Williams 1957). However some of the main points to be dis-
cussed here concern relative diversities among the several taxonomic levels. Presumably,
monographic artifacts would tend to appear at all levels, and relative diversities would
be much less affected than absolute diversities. It is unlikely that there is a consistent

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PALAEONTOLOGY, VOLUME 12

monographic bias in the same direction among a majority of the taxa, and I therefore
believe that the major trends are real.

Another important consideration is the extent to which trends among skeletonized
taxa represent the biota as a whole. At present the non-skeletonized Invertebrata have
the same biogeographic and synecological patterns as skeletonized groups (Lipps, in
press), and there is no reason to expect that patterns of diversity of non-skeletonized
taxa would follow different trends than the skeletonized ones. Furthermore Lipps has
pointed out (pers. comm.) that the preserved groups are morphologically diverse and
unrelated, yet they often exhibit similar patterns. Therefore it is assumed that major
trends among skeletonized and non-skeletonized groups tend to be in phase.

Experience has shown clearly that the chances of preservation of an organism that
does not possess a well-mineralized skeleton are exceedingly small. Indeed, the lack
of a record of a taxon that does not have a relatively high probability of preservation
can hardly be taken as proof that the taxon was not living at the time. And the probabi-
lities of preservation cannot yet be specified even for taxa with highly mineralized
skeletons, under many of the stratigraphic situations common in the geologic record.
It is therefore difficult to assess the significance of negative records.

Most of the known phyla had appeared in the record by Cambrian time, although
even among our sample of nine, one (Ectoprocta) does not appear until the Lower
Ordovician. The phyla are well-differentiated and some contain relatively complex
organisms when they first appear, so that a fairly long period of evolution can be assumed
to have preceded their appearance in the record. However, it is possible to argue for many
phyla that their final organization into the ground-plans that are now considered as
characteristic may have only narrowly preceded their appearance in the record (Cloud
1949, 1968). It has been suggested that such a great evolutionary event may have been
permitted by an increase in atmospheric oxygen past a critical level (Berkner and
Marshall 1965). At any rate, it is likely that nearly all of the invertebrate phyla had
become established before the Cambrian, and the relative timing of their appearance in
the record may partly indicate the order in which they acquired hard parts or the chance
occurrences of unusual preservations. Nicol (1966) and others have suggested that the
acquisition of hard parts may have ensued as a result of widespread phyletic body-size
increases.

Many of the nine phyla in the sample contain taxa that are not members of the shelf
benthos or that have relatively low probabilities of preservation. Examples are the
planktonic Scyphozoa and the soft-bodied Keratosa. Such taxa are excluded from the
tallies. A few other taxa probably participated only partly in the benthonic ecosystems.
An important example is the Ammonoidea. The effects of such taxa on the diversity
curves are considered separately. Diversity graphs for phyla, classes and orders that seem
to be chiefly members of the benthos are presented in text-fig. 3; diversities are classed
by geological epochs, and therefore do not exactly represent the standing diversities at
any given time. The diversity levels depicted in text-fig. 3 represent a balance between
diversification and extinction, but the amount of taxonomic turnover that has occurred
in any epoch cannot be inferred from the diversity levels. Text-fig. 4 depicts the numbers
of appearances and disappearances (presumed to be extinctions) among the higher taxa
per epoch.

From text-figs. 3 and 4 the following history of higher taxonomic diversity can be

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 691

inferred. The higher the category the earlier it tends to reach its maximum diversity.
If the phyla were not all present throughout the Cambrian, at least they all appear by
early Ordovician time, but the highest diversity is recorded in the Middle and Upper
Cambrian. New classes continue to appear until the Lower Carboniferous, but the
highest class diversity is recorded in the Middle and Upper Ordovician. Orders have

text-fig. 4. Appearances and extinctions of phyla, classes and orders of
shelf benthos in the sample, classed by Epochs. Data as in text-fig. 3.

continued to appear until the end of the Cretaceous, but achieved their highest recorded
diversity in medial Ordovician time. Ordinal diversification, however, was great during
early Ordovician time, whereas the greatest class diversification was during the Cambrian
(text-fig. 4).

It also appears that the higher the taxonomic category the less it has been ravaged
by extinction, and the earlier extinction has stopped. Only one phylum (Archaeocyatha),
which comprises 11% of the sample, disappears, although the sample is so small that
this figure cannot be taken as very precise. However, 16 classes comprising 50%, and

692

PALAEONTOLOGY, VOLUME 12

75 orders comprising 64%, disappear. These extinct taxa are never fully replaced by
other higher taxa, at least not from among the taxa that we are considering, so that the
diversity of each higher taxonomic category has decreased, rather markedly in the cases
of classes and orders, since the early Paleozoic. For the taxa in the sample, the extinction
of phyla is complete by the end of Cambrian time, of classes by the end of Permian time,
and of orders by the end of the Cretaceous.

text-fig. 5. Stratigraphic variation in diversity of skeletonized families
of shelf benthos belonging to phyla included in text-fig. 3. Data chiefly
from Harland et al. (1967), Moore (1953-7), and Orlov (1958-64)

The lower taxa present a somewhat different pattern. Text-fig. 5 depicts the geological
record of family diversity in the sample, again excluding unsuitable taxa such as the
planktonic foraminiferal families, and text-fig. 6 depicts the record of appearances and
disappearances. About 100 families are recorded by the end of the Cambrian, and 300
occur in the Upper Ordovician; diversity remains near 300 until the later Paleozoic,
when it gradually falls off. A marked drop occurs in late Permian and early Triassic
times. The pattern has until this point been not too unlike that of the higher taxa, with
the mode of major diversity shifted towards the present. The Jurassic rise is even anti-
cipated on the ordinal level (text-fig. 3). Flowever it is in the great diversity rise of the
Cretaceous and Cenozoic that the pattern of the families departs in a fundamental way

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 693

text-fig. 6. Appearances and extinctions of the families of shelf benthos
in the sample, including Nautiloidea and Ammonoidea, classed by
Epochs. Data as in text-fig. 5.

from the pattern of the higher taxa, for the higher taxa reach and maintain rather steady
levels of diversity from Ordovician onwards for phyla. Lower Triassic onwards for
classes, and lower Jurassic onwards for orders. The Cretaceous and Cenozoic diversi-
fication of families marks a major change in the evolutionary trends of taxonomic
structure.

It can be seen from text-fig. 6 that the times of diversification of families tend to alter-
nate with times of extinction. This pattern is well shown in a figure by Newell (1967,

694

PALAEONTOLOGY, VOLUME 12

fig. 7) that is based upon a different taxonomic sample. The pattern certainly suggests
that times favourable for diversification and those favourable for extinction were distinct,
and that there is therefore a complementary relation between these processes. Newell
suggests in effect that there have been extinctions to provide unoccupied biospace
before there are major diversifications, and of course there must be many taxa resulting
from diversification before there can be major extinctions. However, the data in text-
figs. 5 and 6 do not entirely bear out this thesis. The very high extinction peaks in the
Ordovician, Devonian, and Cretaceous are not accompanied by massive reductions in
standing diversity. In fact, the early Ordovician and Cretaceous extinction highs are
nearly hidden in text-fig. 5 owing to the great contemporary diversifications. Late
Ordovician and Devonian extinction peaks reduce the diversity level somewhat, but only
by about 6 and 13% respectively, because diversification is fairly high at these times.
Only near the Permo-Triassic boundary, when diversification is exceedingly low (text-
fig. 5), does the diversity level suffer a major decline of about 50%. The unusually low
level of Permo-Triassic diversity is not unique because of the extinction peak alone, but
because of the lack of a corresponding peak of diversification.

It is interesting to examine the patterns of family diversification and extinction within
each of the higher taxa. Newell (1967) has presented graphs of family diversification
among a number of higher taxa; I have prepared similar charts for diversification as well
as for extinction of the families of higher taxa in the present sample which confirm the
patterns he has presented. In general, however, the high rates of family diversification
and of extinction within a higher taxon do not alternate in time but are highly correlated.
For example, the Brachiopoda diversify strongly during the Ordovician and Devonian,
but extinction peaks are found at these times also. Secondary levels of diversification
during the Silurian and Lower Carboniferous correlate with secondary peaks of extinc-
tion. Only in the Permian is there a lack of correlation; the great extinction is not accom-
panied (nor is it followed) by diversification at the family level, but is accompanied by
the lowest diversification rate known for brachiopods during the Phatierozoic.

Trilobites display a similar correlation. Cambrian diversification rises to a peak in
the late Cambrian and falls oft' progressively during Ordovician epochs; extinction levels
do precisely the same, except that they rise a bit in latest Ordovician. There is no follow-
ing rise in diversity, however, although there is a secondary peak of extinction in medial
and late Devonian time. Certainly there is no alternation of diversification and extinc-
tion. The same may be said of diversity patterns of the Porifera, the Echinoidea, and
several of the Paleozoic echinoderm groups. The Foraminiferida, Anthozoa, and Gastro-
poda have more complex patterns, but there is no suggestion of alternating extinction
and diversification except at the Permo-Triassic boundary. In the Ostracoda there is
some indication that extinction follows diversification and not the reverse, and similar
trends are found during parts of the record of other taxa. Even in the Ammonoidea and
Nautiloidea peaks of diversification and extinction tend to correlate and certainly do
not alternate.

The alternation of peaks of diversification and extinction of all families in the sample
(text-fig. 6), then, are chiefly due to the alternation of high rates of family extinction of
some higher taxa with high rates of diversification of different higher taxa (which is
usually accompanied by a rise in extinction among these different taxa also). Except at
the Permo-Triassic boundary, all this tends to be accomplished while diversity as a whole

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 695

remains surprisingly stable considering the magnitude of extinction and diversification
peaks. The major events that alter standing diversity on the family level are the diversi-
fication in Cambro-Ordovician times, the Permo-Triassic diversity low, and the
Cretaceous-Cenozoic diversification (text-fig. 5).

text-fig. 7. Stratigraphic variation in the diversity of benthonic shelf
families (dashed line) and genera (solid line) of Foraminiferida, excepting
the poorly skeletonized Allogromiina. Data from Loeblich and Tappan

(1964).

It is possible to demonstrate that with the genera as with the families, there is a striking
rise in numbers in the Cretaceous and Cenozoic. Text-fig. 7 depicts the diversity of
families and of genera of benthonic Foraminiferida through geological time. This is one
of the taxa that contributes strongly to the late Cretaceous and Cenozoic rise in family
diversity. Throughout the Paleozoic there are, on the average, about 3-4 genera per
family described. Across the Permo-Triassic boundary this ratio drops and then rises
again in the Jurassic and early Cretaceous. In the late Cretaceous the genus/family ratio
climbs to nearly 6, and in the early Cenozoic to over 8, where it stands at present after
a late Cenozoic decline. The size of families is somewhat a matter of opinion. There is no

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PALAEONTOLOGY, VOLUME 12

special reason, however, to believe that the disproportionate Cretaceous-Cenozoic rise
is a taxonomic artifact. The data are certainly subject to monographic and other biases,
but have all been reviewed by the same team of authorities (Loeblich and Tappan 1964).
It is interesting in this regard that the same trend can be inferred from the data charted
by Henbest (1952) based chiefly on the work of Cushman (1948). If the trend is an arti-
fact it is an enduring one. Incidentally, peaks of extinction of genera of foraminiferida
correlate rather than alternate with peaks of diversification, just as is common among
invertebrate families.

Another taxon that contributes heavily to the Cretaceous rise in family diversity is
the Gastropoda, among which the same pattern of disproportionate generic diversi-
fication is present. There is no satisfactory recent review of all marine gastropod genera,

ORDERS

GEOLOGIC RANGE

FAMILIES

Genera 8<
Subgenora

Genera - /

Subgenera/ Family

Archaeogastropoda1

U. Cambrian-Recent 2

32

7374

76.0

Mesogastropoda

U.Ord. (Caradocian)- Recent

75

7467

79.6

Opisthobranchia

L. Carb, (Visean)-Recent

9

240

26.6

Neogastropoda

L. Cretaceous (Albian)- Recent

20

7779

56.0

'including Bellerophontacea.

11 Possibly from L. Cambrian, depending upon ordinal assignment of early groups.

text-fig. 8. Families, genera, and subgenera of orders of shallow marine gastropods. The most
advanced orders contain higher numbers of genera and subgenera, on the average, than primitive orders.

Data from Taylor and Sohl (1962).

but Taylor and Sohl (1962) have published a census of gastropod genera and subgenera
combined by family and higher taxa. It is possible to show that the more recently an order
has appeared in the record, the more genera and subgenera per family it contains on
the average (text-fig. 8). The Neogastropoda, which appear in the Lower Cretaceous
(Albian or possibly earlier) and which diversify chiefly in the Upper Cretaceous and
later, have 56 genera and subgenera per family on the average. This is 3| times as many
as the average of the Archaeogastropoda. Furthermore, even the groups of archaeo-
gastropods with the largest living representatives, such as the trochaceans, tend to have
differentiated strongly at the generic level in the Upper Cretaceous and Cenozoic. For
the Trochacea, for example, there are 14 genera and subgenera recorded from the
Lower Cretaceous, 32 from the Upper Cretaceous, and 66 from the Upper Cenozoic
(data from Moore 1953-7). The other orders of Gastropoda are intermediate, both in
time of appearance and in genus-subgenus/family ratios, between the Archaeogastropoda
and the Neogastropoda. It follows from this situation that, for the shallow marine
shelled Gastropoda, the late Cretaceous-Cenozoic rise in family diversity (from 37
families in the Lower Cretaceous to 66 in the Upper Cretaceous and to 83 or 84 in the
Cenozoic) is disproportionately exaggerated on the generic level. This agrees with the
data for Foraminiferida. Two other groups that contribute especially strongly to
the rise in Upper Cretaceous and Cenozoic family diversity, the Ectoprocta and the

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 697

Echinoidea, appear to display similar trends (see Newell 1952 and the appropriate
Treatise volumes).

In contrast to the preceding groups, the phylum Brachiopoda displayed its greatest
familial diversity during the Paleozoic. It has not diversified during the Cretaceous-
Cenozoic but maintained an average of about 12 or 13 families and about 50-65 genera

text-fig. 9. Stratigraphic variation in brachiopod diversity by families (broken
line) and by genera (solid line). The genera/families ratio remains near 4.
Genera after Williams (1965), and family data from Williams et ah (1965).

during this time (text-fig. 9). It is interesting, therefore, that the brachiopods have about
the same genus/family ratio recorded for the Paleozoic as in the Mesozoic and Cenozoic
— about 4 times as many genera as families throughout the Phanerozoic. Even during
the greatest periods of diversification, the numbers of genera per family did not rise
disproportionately, although generic turnover was significantly greater than family
turnover. Judging from a comparison of the graphs of evolutionary rates among trilobite
genera (Newell 1952) with family diversity trends, disproportionate generic diversity is
not found among trilobites during their time of greatest family diversity either.

In summary, diversification on the family level during the Paleozoic and early Mesozoic

z z

C 6940

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PALAEONTOLOGY, VOLUME 12

seems to be accompanied chiefly by simple proportionate diversification on the generic
level. Diversification on the family level during the late Cretaceous and Cenozoic seems
to be accompanied by a disproportionately high diversification on the generic level.

It is impracticable to attempt a census on the species level for even a few higher taxa,
and there are strong reasons for doubting the significance of fossil species counts in any
event. It is necessary to approach species diversity at least in part from a theoretical
point of view. Species evolve at greater frequencies than genera or families or higher
taxa, and their standing diversities are therefore more volatile. The appearance of
isolated habitats can produce swarms of closely related species, and the development
of specialized communities may permit the development of a great number of species,
not necessarily closely related, but endemic to the community. Reef communities, for
example, appear to contribute large opportunities for both these types of speciation.
A great number of specializations are possible on reefs, and thus large numbers of
relatively specialized species from various phylogenetic backgrounds may appear. Reef
tracts are also characterized by patchy and discontinuous distributions of reefs, and the
isolation of outlying patches might often serve as a basis for speciation. Communities
such as those on reefs that appear, endure long enough for a highly specialized biota to de-
velop, and then disappear or become greatly reduced, can produce temporally localized
but significantly large fluctuations in standing species diversity. If species diversification is
great within such communities, generic diversity would also be enhanced. Since the
species endemic to such communities are normally specialized, the average niche size of
the shelf biota would be decreased while they flourish and increased when they wane.

At times the middle and upper Paleozoic record contains numerous reef associations
and at these times it is likely that species diversity reaches disproportionately high levels,
relative to families. It is expected that generic diversity might also rise disproportionately
at these times. Although early and middle Permian reefs are widespread and contain
probably the most specialized Brachiopoda recorded (Rudwick and Cowen 1968), a dis-
proportionate generic diversity peak does not appear at that time in the available data
(text-fig. 9). The description and evaluation of Permian reef biota is far from finished,
however.

Another important way in which specific (and generic) diversity may be dispropor-
tionately multiplied is through a rise in provinciality. For example, theoretical considera-
tions suggest that there are many more shelf species today than in the past, owing to
the high degree of shallow-water provinciality at present (Valentine 1967, 1968a). This
provinciality is both latitudinal, correlating with the great latitudinal temperature
gradients at present, and longitudinal, owing to the presence of efficient biogeographic
barriers of continents and ocean deeps. In the early Jurassic provinciality was not
strongly developed. A Middle and Upper Jurassic Boreal fauna that contains endemic
forms has been widely recognized (Neumayr 1883; Arkell 1956). Evidence has now been
advanced to suggest that the Boreal fauna of the Jurassic signifies a low-salinity facies
in a region of rather stable palaeogeography rather than a climatic province ( Hallam 1 969).
Whatever its environmental basis, the appearance of the widespread fauna marks an
increase in environmental heterogeneity on a sub-continental scale and a rise in species
diversity. The general trend towards increasing provinciality in late Cretaceous and
Cenozoic times must have greatly enhanced the numbers of species on the shelves
(Valentine 1967, 1968a). Many genera which contain several species in a given province

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 699

are now represented in different provinces by separate suites of species, so that the total
numbers of their species are immense. Such is the case with species of Nucula, Macoma,
and Mactra among the Bivalvia and Conus, Calliostoma , and Fissurella among the
Gastropoda, to choose a few of the many examples. This situation must have been much
less marked during times of low provinciality. For these reasons alone it is contended
that the number of species in the shelf environment has increased disproportionately
relative to the genera, especially during the Cretaceous and Cenozoic. Thus a species
diversity curve would have about the same pattern as a generic diversity curve, but the
peaks would be exaggerated (and the curve would be offset slightly towards the present).
There are still other reasons, discussed below, for believing this pattern to be correct.

The major diversity trends in time among the fossil taxa are assumed to reflect real
diversity trends among the ancient shelf biota, and they can be described in terms of the
structure of the taxonomic hierarchy (text-fig. 10). In the earliest Paleozoic each phylum
was represented by only a few classes, each class by relatively few orders, and so on
down the hierarchy. By the close of Ordovician time, however, the average phylum was
well differentiated into classes, and the average class into orders. Some phyla became
extinct, but were not replaced by other phyla. After the Middle Ordovician the diversity
of classes and of orders declined but the diversity of families rose, so that the structure
became relatively more diversified among the lower taxa (text-fig. 10). It is possible that
the average generic and specific diversity of the Upper Carboniferous indicated in text-
fig. 10 is too low, owing to a disproportionate diversification at these lower levels that
culminated during the early Permian. At about the Permo-Triassic boundary, both the
numbers of classes and of orders were reduced by just less than half relative to their
Middle Ordovician peaks, as were the families. The Permo-Triassic diversity low was
most marked at lower taxonomic levels (contrast the familial and ordinal diversity
decreases).

After the Permian the only gains in diversity registered among higher taxa are on the
ordinal level. The numerous lost classes are not replaced, and even the rise in ordinal
diversity is relatively small. The great climb in diversity at the familial level returns the
hierarchy to a pattern commensurate with the early Upper Paleozoic pattern by Middle
Jurassic time. Thereafter the number of families per higher taxon increases, especially
during the Upper Cretaceous, and the diversities of lower taxa follow suit (text-fig. 10).

The so-called nekto-benthonic cephalopod groups Nautiloidea and Ammonoidea
have not been included in the basic sample because of uncertainty as to the degree to
which they participated in benthonic ecosytems. Nevertheless, some of them were surely
regular members of a benthonic food chain. In text-fig. 5, the families of these cepha-
lopod taxa are added to the families of the sample. The pattern of diversity is not much
altered thereby; there was a slight increase in the steepness of the diversity rise from the
Triassic to the Lower Cretaceous and the appearance of a rough plateau during
the late Cretaceous and Cenozoic. The effect on the ordinal level is to emphasize
the Ordovician rise in diversity and the mid- and upper-Paleozoic decline. Therefore,
the exclusion of these Cephalopoda from the sample has a conservative effect insofar
as the major trends are concerned, and in no way contributes to special conclusions.

Another major group which might have merited some representation in the figures is
the fishes. At the family level their inclusion would raise the curve in text-fig. 5 to an even
higher Devonian peak, and somewhat steepen the Mesozoic trend of rediversification (as

700

PALAEONTOLOGY, VOLUME 12

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VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 701

would almost any additional taxon). At higher levels, they would raise ordinal diversity
in medial Paleozoic and class diversity in early Paleozoic times, emphasizing graph
patterns. Clearly, their exclusion does not affect the major diversity trends.

COADAPTATION OF ECOLOGICAL AND TAXONOMIC HIERARCHIES

The correlation of the structures of the two hierarchies under consideration is best
approached at the species level. The fluctuations in species diversity, which have changed
through a whole order of magnitude and more at times (Valentine 1967), naturally cause
wide fluctuations in the numbers of populations in the ecological hierarchy, which must
be accommodated in some manner.

In general there are three ways in which new species populations might be accom-
modated in ecological units (Klopfer 1962): (1) they may colonize parts of the environ-
mental lattice that were previously unoccupied, in which case their niches represent an
extension of ecospace in the ecological unit but average niche size is little affected; (2)
space for their niches may be created in the lattice by shrinking one or several of the
pre-existing niches, in which case the existing ecospace becomes more crowded by
partitioning and average niche size decreases; or (3) parts of their niches may overlap
with one or several pre-existing niches and thus crowd the lattice by overlap rather than by
partitioning. The last sort of accommodation, by niche overlap, occurs hand in hand with
one of the other sorts. The fitting of a new niche into the lattice may commonly involve
all these sorts of accommodation. Theoretical or practical aspects of niche partitioning,
which were touched on by Darwin (1859), have been considered in a modern perspective
by a number of workers (for example, Bray 1958; Brown and Wilson 1956; Klopfer and
MacArthur 1960, 1961; Klopfer 1962; McLaren 1963; MacArthur and Levins 1967;
MacArthurand Wilson 1967; Hutchinson 1967; and Miller 1967). Yet data on variations
in niche size and its relation to species diversity among marine shelf invertebrates is
scanty. Marine research includes the work of Kohn ( 1 959, 1 966) on the gastropod Conus
and Connell (1961) on some intertidal barnacles.

The most recent major rise inferred in species diversity from the late Cretaceous to
the present seems to have involved an extension of ecospace, an invasion of parts of the
lattice which were becoming newly realized, thus expanding the available environment.
This increase in environmental heterogeneity in the shallow marine realm was evidently
due partly to the cooling of shelf waters in high latitudes (Smith 1919; Durham 1950;
Valentine 1967, 1 968 a), which permitted the rise of new biogeographic provinces in separate
chains along north-south-trending coastlines. New provinces may also have been created
by the drifting apart of some continents, progressively isolating, from about medial
Cretaceous time, shelf regions that had previously been connected. This permitted an
increase in endemism.

Thus this expansion of ecospace is envisaged as due primarily to two factors: (1)
extension of the thermal factors and of numerous other parameters that are related to
temperature, creating one set of biogeographic barriers; and (2) the creation of another
set of barriers through the breaking up of formerly continuous or nearly continuous
epicontinental seaways and continental shelves through continental drift (which in
effect multiplies biospaces along dimensions of real space). The relative timing of these
events is not yet clear, although there is a suggestion that longitudinal provinciality was

702

PALAEONTOLOGY, VOLUME 12

strengthening in the late Cretaceous (Sohl 1961) while latitudinal provinciality was still
weak. The Cretaceous-Cenozoic expansion of ecospace was fundamentally on the level
of the biosphere and involved the rise of provinciality, which in turn permitted new com-
munities based upon endemic populations to appear in each province. The increase in
isolation among the populations living in separate provinces or separate communities
permitted the formation of many new species.

The increase in generic diversity follows from the multiplication of species and their
isolation in separate provinces or communities. Different but related species with similar
morphological adaptations would arise in similar habitats in different provinces, forming
associations such as Thorson's (1957) parallel bottom communities or becoming
‘geminate’ or twin species such as occur on opposite sides of the Isthmus of Panama
(Ekman 1953). The increase in generic diversity would be proportionately less than in
species diversity, owing to the multiplication of similar morphological types in distinct
provinces that would be grouped as genera by taxonomic practice. The same principle
seems to apply to the family level. A marked increase in the diversity and provinciality
of genera would lead to a more or less modest increase in family diversity. That the pre-
sent biogeographic pattern of diversity is similar at the familial, generic, and specific
levels has been well documented for the Bivalvia by Stehli, McAlester, and Helsley
(1967). Kurten (1967) suggested that mammalian diversity is high at the ordinal level
because of endemism arising through the isolating effects of continental drift.

At higher taxonomic levels on the marine shelves a different factor must be operating
to control diversity, since higher taxa do not much participate in the Mesozoic-Cenozoic
diversification. Perhaps this diversification has occurred too recently for evolution to
have proceeded to the level of higher taxa. The pattern of ordinal diversity suggests that
this may be the case. Yet class diversity has declined since the Ordovician. A better
explanation may be that the available biospace of the epicontinental seas and shelves
was nearly fully occupied since at least very early in the Phanerozoic, and most increases
in taxonomic diversity had to be accomplished by ecospace partitioning and overlap (see
Rhodes 1962, pp. 270-2; Nicol 1966). Thus only in times of unusual expansion of the
marine shelf biospace, when the realized parts of the environmental lattice increased
persistently, would significant diversity increases result simply from the invasion of new
biospace. Ecospace partitioning involves a decrease in the niche sizes of populations, at
least along dimensions where competition may occur (Miller 1967), and this is not a
process that lends itself to the appearance of organisms with wholly new ground-plans
or with major modifications thereof, such as are required for the development of higher
taxa. It is instead a process suited more to the modification in detail of pre-existing
morphological types, so as to accommodate to smaller ecospaces — in other words, a
process suited to the increase of specialization.

By Cambrian time or shortly thereafter the ground-plans that are the hallmarks of
the major invertebrate phyla had been established. Most of the species were by present
standards primitive and functionally generalized; modal niche size was no doubt far
larger than at present, though there certainly may have been some highly specialized
forms. Evidently, diversification in the Cambrian and Ordovician led to the presence
in late Ordovician time of a large number of higher taxa. This may have been partly due
to an expansion of biospace and should have included an increase in resources. As much
of the former marine shelf biospace may have been occupied by soft-bodied organisms,

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 703

the expansion of skeletonized forms may have involved the appropriation of some re-
sources that had formerly been utilized by soft-bodied groups. On the other hand, on
the assumption that soft-bodied taxa should have responded to the same opportunities
as skeletonized taxa, it seems even more likely that there was a concomitant diversi-
fication of soft-bodied lineages. These are debatable points, but it is clear that it cannot
be assumed that Cambro-Ordovician radiation was occurring in vacant biospace. Vast
regions of the lattice must have been occupied, and this may have served to channel the
evolutionary pathways of diversifying lineages.

Certainly there was unusual extinction among higher taxa during this time. Higher
taxa in the sample that appear in the Cambrian, but which are not known to have sur-
vived the Ordovician, include the phylum Archaeocyatha, the echinoderm classes
Homostelea and Helicoplacoidea, the inarticulate brachiopod orders Obolellida, Pateri-
nida, and Kutorginida, the monoplacophoran order Cambridoida, and the trilobite
orders Redlichiida and Corynexochida. Numbers of other taxa that are poorly known
also disappeared early, and may have represented higher taxonomic levels. They were
not included in the sample because of their questionable status. These include some
trilobitoids, some early echinoderm stocks, and early gastropod-like molluscan stocks.
Perhaps most of these are functionally generalized forms, the morphological architecture
of which proved unsuitable to the demands for specialization.

Biospace formerly occupied by populations of extinct lineages would soon be re-
colonized by populations of the lineages that remained, if such colonization did not
actually precede and contribute to the extinction. This easily leads to the diversification
of extant lower taxa, but not to the creation of taxa on a comparably high level. In the
early Phanerozoic the large average size of the former ecospaces of extinct taxa provided
ecological room, so to speak, which allowed the lineages that reoccupied vacated bio-
space a certain leeway for progressive morphological modification that could still lead
to the establishment of a higher taxon. Later, when vacated biospace was to be in smaller
parcels, opportunities for morphological modifications became limited, as during medial
Paleozoic time, when the diversity of classes declined markedly (text-fig. 3). Although
diversity of orders (text-fig. 3) and families (text-fig. 4) also declined during that time,
the decline was less marked, and order/class and family/order ratios were both rising.
This suggests that the extinction of a higher taxon was not usually accompanied by
replacement at the level of the higher taxon but at a lower level. It also suggests that
biospace was decreasing. In the Lower and Middle Permian the numbers of species and
genera may well have been disproportionately higher than is suggested by the number
of families, owing to the development of reef associations.

Towards the close of the Paleozoic the change in the taxonomic structure suggests
a great reduction in the heterogeneity of the shelf environment. Provinciality was
already low. It is not certain, therefore, whether a significant further reduction occurred
or indeed was even possible in late Permian time. However the numbers of communities
certainly decreased (for example, the late Paleozoic reefs disappear) and, although there
are not well-documented field studies, it appears from faunal lists that the numbers of
populations in the remaining communities also declined. Thus the ecological structure
shrank at all, or nearly all, levels, suggesting a general decline in biospace. There are
too few data on the ecological hierarchy of Permo-Triassic times, however, to support
speculation on the precise causes of the extinctions on this basis. Rhodes (1967) has

704

PALAEONTOLOGY, VOLUME 12

reviewed the major hypotheses of extinction and remarks that probably none of them
alone would cause the sorts of changes found at the Permo-Triassic boundary.

Rediversification evidently began by medial Triassic time, unless the increase in
family diversity then is an artifact. If it is real (and it includes the beginnings of Sclerac-
tinian radiation as well as the expansion of gastropod families), it suggests that generic
and specific diversification was proceeding at even higher rates. Presumably biospace
had expanded (or was expanding) once more and the newly realized parts of the environ-
mental lattice were being recolonized. However, compared with Cambro-Ordovician
lineages, Triassic lineages were rather specialized, with smaller modal niche sizes, so that
the average colonizing lineage must have occupied a relatively smaller part of the lattice.
The opportunities now presented for the formation of higher taxa could not be much
exploited by the relatively specialized populations. There may be exceptions; the Sclerac-
tinia may have taken advantage of biospace vacated by Paleozoic coelenterate lineages
to become skeletonized and reoccupy some of the same biospace. Finally, at some time
in the Mesozoic a diversification involving the marked rise in provinciality discussed
previously began to occur as well. It seems likely that the Jurassic and early Cretaceous
diversity increases, which are not inconsiderable and which take place at a high rate
(text-fig. 5), are partly owing to an increase of latitudinal provinciality due to cooling
poles and to an increase of longitudinal provinciality due to the separation of some
continental masses in Jurassic time. A thorough analysis of the biogeographic patterns
of Mesozoic diversification is badly needed.

The Cretaceous-Cenozoic boundary is marked by extinctions of some benthonic
marine groups (see Hancock 1967), but if there was any appreciable alteration in the
taxonomic diversity structure at the family and higher levels it was of so short a duration
that it does not appear in the present data. The structure of communities, provinces and
of the entire shelf realm must have undergone qualitative changes, but this is a more or
less continuous process on the broad scale we are considering. The Late Cretaceous rise
in diversity extends unabated across the Cretaceous-Cenozoic boundary. Surely there
was no large-scale reduction in biospace.

Two major modes of taxonomic diversification have been described, both proceeding
at progressively lower taxonomic levels through time. The first involves a biospace that
fluctuates about some size that does not vary much in time. Diversification at the popula-
tion level at first proceeds by colonization of untenanted biospace, but soon must be
accompanied by a progressive decrease in average niche size. Communities therefore
become increasingly packed with more and more specialized populations and begin to
fragment into portions, each of which has an energy flow that is partially independent of
the others. The isolation and independence of these portions will increase with further
specialization until they form ecosystems that are as independent as the original one
from which they fragmented. Slighter and slighter environmental discontinuities will
form community boundaries until the environmental mosaic of a given primitive com-
munity, relatively heterogeneous but occupied by primitive populations with large
niches, is broken up into a number of smaller environments, each more homogeneous
than the original and each occupied by more specialized populations. Provinces become
packed with more and more communities that have progressively smaller ecospaces. The
diversity of provinces is not so sensitive to this progressive specialization, although it may
eventually be affected if the trend continues long enough. Any widespread partitioning

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 705

of temperature or temperature-correlated parameters would lead to increased pro-
vinciality even in the absence of progressive changes in the latitudinal temperature
gradient. If partitioning were to continue, smaller and smaller changes in thermal regimes
would act to localize range end-points, and thus form provincial boundaries (Valentine
1966). In sum, in this mode the ecological structure evolves from lower levels towards
higher.

The other mode of diversification involves a biospace that expands to create new
environments or to add new dimensions (or at least to extend old dimensions) to old
environments. New environments may be created, for example, through climatic changes,
and old environments may be extended through the improvement of limiting factors,
that is through the amelioration of conditions which tended to inhibit diversification.

From what is known and can be inferred of the hierarchies of the Paleozoic, diversi-
fication (at least after an initial radiation of skeletonized taxa and probably before) was
proceeding chiefly in the first mode, from the bottom of the ecological hierarchy up-
wards, implying a relatively stable biospace. Rediversification following the Permo-
Triassic extinction was probably in the second mode, involving an amelioration of
factors that had inhibited diversification, and the Upper Cretaceous and Cenozoic
diversification seems to have also been in the second mode, but involved the creation
of new environments. Nevertheless there must have been continuing specialization
and thus the first mode was also active in the taxonomic and ecological evolution.
Processes that bring species that appeared during biospace expansion into sympatry
with older lineages, such as ‘species pumps’ of various kinds (Valentine 1967, 1968a),
may link these two modes into a single system of diversification. Finally, the more
lineages that exist the greater the opportunity for large-scale diversification under
appropriate circumstances in either mode. This factor is certainly at work in the dis-
proportionate multiplication of lower taxa during Cretaceous and Cenozoic times.

A number of authors have suggested that extensive changes in sea level may control
some of the diversification and extinction patterns (Newell 1952, 1956, 1963; Moore
1954; see Rhodes 1967 for other references). Widespread epicontinental seas, it is asserted,
provide more inhabitable area for shelf invertebrates and therefore more opportunity
for diversification, while regressions reduce the inhabitable area and thus the diversity.
There is some theoretical support for this position in the species-area work of Preston
(1962), Williams (1964), MacArthur and Wilson (1967), and others. Little work has
been done in marine environments; the areas of ancient shelf seas, especially during
regressive phases, are difficult to estimate (though see Ronov 1968); and the effects that
could be expected in a biosphere of vastly different ecological and taxonomic structure
and composition are largely uncertain. The problem is further complicated by facies
differences between epicontinental seas and shelves bordered by open oceans. Although
uncertainties in calculations must be great, preliminary estimates suggest that the species-
area effect would have been far too small to account for major diversifications and ex-
tinctions by itself. Moreover, we live at present in a time of great continental emergence
yet the shelves are richly diverse in lower taxa and in ecological units at all levels, pre-
cisely the opposite of the pattern of Permo-Triassic extinction. Indeed, a relative lower-
ing of sea level must commonly result in the emergence of land barriers which isolate
regions formerly connected and permit the rise of an endemic biota in each region. This
would have the effect of increasing the total number of species in these regions.

706

PALAEONTOLOGY, VOLUME 12

Nevertheless, the elimination or rise of species resulting from shelf-area fluctuations
would certainly contribute to diversity patterns, and further evaluation of this sub-
ject is clearly merited.

CONCLUSIONS: THE PROGRESSIVE CANALIZATION OF ECOSPACE

It is concluded that a major Phanerozoic trend among the invertebrate biota of the
world’s shelf and epicontinental seas has been towards more and more numerous units
at all levels of the ecological hierarchy. This has been achieved partly by the progressive
partitioning of ecospace into smaller functional regions, and partly by the invasion of
previously unoccupied biospace. At the same time, the expansion and contraction of
available environments has controlled strong but secondary trends of diversity. Present
marine biospace is in fact unusually extensive, and the world’s shelf seas are therefore
unusually heterogeneous and support a large number of ecological units today. The
relations of these trends to trends within the taxonomic structure of the benthonic
invertebrates are intimate.

Assuming for the moment that evolutionary trends among marine benthonic inverte-
brates will continue and that biospace does not change much (a dim prospect in view of
rising pollution), what might be predicted of the future structures of the ecological and
taxonomic hierarchies ? Speculation on this point may be of some value to underline the
sort of process that is postulated to have gone before. Clearly, the trend towards
specialization would further reduce the average niche sizes of species. It would be
increasingly difficult for evolving lineages to depart much from their modal functions
and morphologies, as biospace would become available only in increasingly smaller
compartments. The amount of change necessary to produce a new family would be
increasingly difficult to attain, and eventually no new families could appear. In fact,
some families would become extinct so that familial diversity would decrease, and
lineages from other families would fill any vacated biospace. After some time genera
could no longer appear, for biospace would be packed too tightly to permit morpho-
logical variation even at that level, and generic diversity would decline for a while as
some extinction, inevitably, occurred. Eventually, all the biospace would become filled
with evolving lineages with an incredibly small modal niche size, each lineage constrained
by the presence of all the others to evolve in only a narrow pathway directed by the trends
of evolution of the entire biota, and of changes in the entire biosphere. Ecological
units are now exceedingly small by today's standards, with virtually every few food-chains
forming a separate community and every moderate topographic irregularity forming
a provincial boundary. Canalization of ecospace is complete. The biosphere has become
a splitter’s paradise.

Although this extrapolation cannot be taken too seriously, it does point to some
important consequences of ecospace partitioning. First, average species of the
early Paleozoic, with their broad niches, may have had different patterns of morpho-
logical variation than the specialized species of today. Secondly, the occurrence in the
early Paleozoic of numbers of unusual ‘ aberrant ’ higher taxa that contain few lower taxa
is not necessarily due to a poor fossil record but is probably the natural consequence of
adaptive strategies that prevailed in primitive ecosystems of low diversity. Finally, extinc-
tion of taxa of high diversity is less likely than extinction of taxa of low diversity, other

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 707

things being equal (Simpson 1953), simply because so many more lineages must dis-
appear. Similarly, the markedly rising provinciality of the late Cretaceous and Cenozoic
will tend to make the extinction of the newly diverse taxa that have representation in
many provinces — a common situation even on the generic level — more difficult.

Acknowledgements. Gratitude for extensive discussion of the ideas presented herein is expressed to
Dr. A. Hallam (Oxford University) and Professor A. L. McAlester (Yale University). The manuscript
was carefully reviewed by Dr. W. S. McKerrow (Oxford University) and Professor R. Cowen, Pro-
fessor J. H. Lipps, and Robert Rowland (University of California, Davis). All this attention resulted
in much improvement. The manuscript was written during a Guggenheim Fellowship spent at the
Department of Geology and Mineralogy, Oxford University, and at the Department of Geology and
Geophysics, Yale University. The generosity of the Guggenheim Foundation and the hospitality ot
these departments is gratefully acknowledged.

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NicoL, d. 1966. Cope’s rule and Precambrian and Cambrian invertebrates. J. Paleont. 40, 1397-9.
orlov, y. a. 1958-64. Osnovy Paleontologii, Akad. nauk SSSR., Moscow (in Russian).
parr, a. e. 1926. Adaptiogenese und Phylogenese; zur Analyse der Anpassungserscheinungen und ihre
Entstehung. Abb. Theor. org. Entw. 1, 1-60.

preston, f. w. 1962. The canonical distribution of commonness and rarity. Ecology, 43, 185-215,
410-32.

rensch, b. 1947. Neuere Probleme der Abstammungslehre. Stuttgart.

Rhodes, f. h. t. 1962. The evolution of life. Baltimore, Md.

- 1967. Permo-Triassic extinction. In harland, w. b., et al. (eds.), The fossil record, 57-76. London
(Geological Society).

ronov, a. b. 1968. Probable changes in the composition of sea water during the course of geological
time. Sediment ology, 10, 25-43.

R ud wick, M. j. s. and cowen, r. 1968. The functional morphology of some aberrant strophomenide
brachiopods from the Permian of Sicily. Bol. Soc. Paleont. Ital. 6, 113-76.
simpson, g. g. 1944. Tempo and mode in evolution. New York.

— — 1953. The major features of evolution. New York.

smith, j. p. 1919. Climatic relations of the Tertiary and Quaternary faunas of the California region.
Proc. Calif. Acad. Sci. (4) 9, 123-73.

sohl, n. f. 1961. Archaeogastropods, Mesogastropods, and stratigraphy of the Ripley, Owl Creek and
Prairie Bluff Formations. Prof. pap. U.S. geol. Surv. 331 A, 151 pp.
stehli, F. g., mcalester, a. l., and helsley, c. e. 1967. Taxonomic diversity of Recent bivalves and some
implications for geology. Bull. geol. Soc. Am. 78, 455-66.
taylor, d. w. and sohl, n. f. 1962. An outline of gastropod classification. Malacologia, 1, 7-32.
thorson, g. 1957. Bottom communities (sublittoral or shallow shelf). In hedgpeth, j. w. (ed.), Treatise
on marine ecology and paleoecology. Mem. geol. Soc. Am. 67 (1), 461-534.
valentine, j. w. 1966. Numerical analysis of marine molluscan ranges on the extratropical north-
eastern Pacific shelf. Limnol. Oceanogr. 11, 198-211.

- 1967. Influence of climatic fluctuations on species diversity within the Tethyan Provincial System.
In adams, c. g. and ager, d. v. (eds.), Syst. Ass. Pub. 7, 153-66.

1968«. Climatic regulation of species diversification and extinction. Bull. geol. Soc. Am. 79,

273-76.

- 19686. The evolution of ecological units above the population level. J. Paleont. 42, 253-67.

1969. Niche diversity and niche size patterns in marine fossils. Ibid. 43, 905-15.

williams, a. 1957. Evolutionary rates in brachiopods. Geol. Mag. 94, 201-1 1.

VALENTINE: PATTERNS OF TAXONOMIC AND ECOLOGICAL STRUCTURE 709

williams, a. 1965. Stratigraphic distribution. In moore, r. c. (ed.), Treatise on invertebrate paleon-
tology, Part H. H237-50. Geol. Soc. Am. and Univ. Kansas Press.

— — et al. 1965. In moore, r. c. (ed.). Treatise on invertebrate paleontology. Part H. Geol. Soc. Am. and
Univ. Kansas Press.

williams, c. b. 1964. Patterns in the balance of nature and related problems in quantitative ecology.
New York.

j. w. valentine
Department of Geology
University of California
Davis, California, 95616

Typescript received 8 April 1969 U.S.A.

THE PALAEONTOLOGICAL ASSOCIATION

Annual Report of the Council for 1968-9

Membership. On 31 December 1968 there were 1330 members (716 Ordinary, 121 Student, and 493
Institutional), a net decrease of 23 members during the year, the first year of the higher subscriptions
for Ordinary and Student Members.

Finance. During 1968 the Association published Palaeontology in five parts at a cost increased to
£10674. The reprinting of Volume 3 was completed at a cost of about £1500, £426 of which has so
far been paid. Special Papers Nos. 2 and 3 were published at a total cost of £1594. Administration
costs remained a relatively small charge on the Association’s finances. In spite of expenditure increasing
to a record level, the Association's reserves were greatly strengthened during the year because of increased
income. The higher rates of membership subscriptions provided £940 more income, and sales increased
from £4378 to £5973, giving an excess of income over expenditure amounting to £2315. The excess has
been added to the Publications Reserve which now stands at £9540, sufficient to pay for one year’s
publication of Palaeontology. Sales of Special Papers have been proceeding at a reasonably steady
rate, and the cost of each has already been covered by sales and donations. The Association is indebted
to: the Shell Oil Co. for a further grant of £250 for Special Papers; Belfast University for a donation
of £150 towards the cost of No. 2; to Oberlin College for a donation of £475 towards the cost of
No. 3; and the the Royal Society for a loan of £1000 for the production of No. 2.

Publications. Five parts of Palaeontology were published during 1968 to form Volume 11 ; they con-
tained 61 papers and consisted of 824 pages and 154 plates. Special Papers in Palaeontology Nos. 2 and
3 (for 1968) were published; they contained a total of 137 pages and 32 plates.

Meetings. Six meetings took place during 1968-9. The Association is grateful to the Council of the
Geological Society of London, Dr. H. W. Ball (Keeper of Palaeontology at the British Museum —
Natural History), and Professor F. Hodson (University of Southampton) for generously granting
facilities for meetings, to Dr. A. Hallam for leading the field meeting, and to Mr. A. Rixon for giving
the demonstration.

a. The Eleventh Annual General Meeting was held in the rooms of the Geological Society of London,
Burlington House, London, W. 1, on Wednesday, 6 March 1968, at 5.00 p.m. The Annual Report
of the Council for 1967-8 was adopted and the Council for 1968-9 was elected. Dr. Maurice
Black of the University of Cambridge delivered the Eleventh Annual Address on ‘Taxonomic
problems in the study of coccoliths’.

b. A Field Demonstration Meeting was held on the Frodingham Ironstone on Saturday, 4 May 1968.
Dr. A. Hallam demonstrated features of palaeoecological interest.

c. A Joint Meeting with the Geological Society was held in the rooms of the Geological Society
on Wednesday, 19 June 1968. Professor Melvin Calvin gave a lecture on ‘Molecular Palaeonto-
logy’.

d. A Special Lecture by Professor Dr. E. Voigt ( University of Hamburg) on ‘Palaeohistological studies
on the soft parts of fossil animals from the Eocene lignites of the Geiseltal, near Halle (Saale)’
was delivered in the rooms of the Geological Society of London on Wednesday, 2 October 1968.

e. A Demonstration Meeting on ‘The preparation and preservation of fossils' was given by Mr. A.
Rixon in the Department of Palaeontology at the British Museum (Natural History) on Saturday,
16 November 1968, at 2.00 p.m.

f A Symposium on ‘Patterns of Evolution' was held in the Department of Geology, University of
Southampton, on Tuesday to Thursday, 17 to 19 December 1968. About 60 persons attended to
hear 1 1 papers and see a number of demonstrations. Professor Frank Hodson was local secretary.

THE PALEONTOLOGICAL ASSOCIATION

711

Council. The following were elected members of the Council of the Association for 1968-9 at the
Annual General Meeting on 6 March 1968: President'. Professor Alwyn Williams, F.R.S.; Vice-
Presidents : Dr. W. S. McKerrow, Professor C. H. Holland; Treasurer. Dr. C. Downie; Membership
Treasurer : Dr. A. J. Lloyd; Secretary: Dr. J. M. Hancock; Editors: Mr. N. F. Hughes, Dr. Gwyn
Thomas, Dr. I. Strachan, Professor M. R. House, Dr. R. Goldring; Other members: Dr. F. M. Broad-
hurst, Mr. M. A. Calver, Dr. C. B. Cox, Mr. D. Currey, Dr. Grace Dunlop, Dr. G. F. Elliott, Dr. A.
Hallam, Dr. Julia Hubbard, Dr. J. D. Hudson, Dr. R. P. S. Jefferies, Dr. J. D. Lawson, Dr. A. H.
Smout, Professor H. B. Whittington.

Shortly after the Annual General Meeting, Dr. W. D. I. Rolfe was co-opted as Assistant Secretary
under Rule 3. In order to keep the number of officers to 1 1, the maximum permitted by the rules of the
Association, Professor C. H. Holland graciously offered to resign as Vice-President.

Circulars. Three Circulars (Nos. 54-6), containing full details of the affairs of the Association were
distributed to Ordinary and Student Members during the year, and to Institutional Members on
request.

712

PALAEONTOLOGY, VOLUME 12

BALANCE SHEET AND ACCOUNTS FOR
THE YEAR ENDING 31 DECEMBER 1968

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Report of the Auditors to the Members of the Palaeontological Association. We have examined the above
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and fair view of the state of the Association’s affairs as at 31 December 1968 and of its income and expendi-
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JOSHUA WORTLY & CO.
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THE PALAEONTOLOGICAL ASSOCIATION

713

Income and Expenditure Account for the Year ending
31 December 1968

Expenditure

£

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d.

To provision for the cost of Publication of Palaentology, Vol. 1 1

, Part 1

2049

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To Depreciation .......

To Publications Fund, Excess of Income over Expenditure
To Special Papers .......

£ s. d.

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£13 870 1 7

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3688 2 3
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110 0 0
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46 15 9
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38 16 0
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545 8 9
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250 0 0
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1806 17 4
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2596 17 4
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£13870 1 7

3 A

C 6940

INDEX

Pages 1-171 are contained in Part 1 ; pages 173-350 in Part 2; pages 351-535 in Part 3; pages 537-709 in Part 4.

Figures in Bold Type indicate plate numbers.

A

Abathomphalus, 26; intermedia, 27, 2; mayaroensis,
26, 2.

Abietineaepollenites minimus, 109.

Acanthotriletes varispinosus, 102.

Acherontiscus, 538; caledoniae, 539.

Acinosporites saiopiensis, 225, 39.

lAcrocy there inornata , 144, 28.

Aequitriradites spinulosus, 108.

Alaska: landplants in graptolitic shale from, 559.

Alisporites bilateralis, 109.

Ambitisporites, 228; cf. avitus, 40; cf. dilutus, 229, 40.

Ambocoelia praecox dorsiplicata, 478, 90.

Amphibia: new family from Carboniferous, 537.

Amphicythere, 146; confundens, 146, 29; pennyi, 146,
29; sphaerulata, 147, 30.

Anomalinoides, 197; hyphalus, 197; velascoensis, 198.

Apatognathus, 401.

Apiculiretusispora, 219; cherata, 219, 37; microconus,
219, 37 ; spicula, 220, 38 ; synorea , 221, 38 ; sp. A,
221, 38; sp. B, 221, 38; sp. C, 222, 38.

Appendicisporites', jansonii, 108; potomacensis, 107.

Araucariacites australis, 110.

Arcellites medusus, 334, 62, 63, 67.

Archaeozonotriletes, 234; chulus, 234; c. chulus, 235,
43 ; c. inframurinatus, 237, 43 ; c. nanus, 238, 43 ; cf.
divellomedium, 238, 43; dubiits, 238, 42.

Archimedes', sp., 52, 53, 54; wortheni, 52.

Armstrong, J. The cross-bladed fabrics of the shells of
Terrakea solida (Etheridge and DunnJ and Strepto-
rhynchus pelicanensis Fletcher, 310.

Arthropoda. See Crustacea, Ostracoda, Trilobita.

Asperopora, 630; aspera, 632, 115 ; multipora, 632, 116.

Atlantic: benthonic foraminifera from Galicia Bank,
189; planktonic foraminifera from Galicia Bank, 19.

Aulacotheca iowensis, 415, 76, 77.

Austin, R.L., and Rhodes, F.H.T. A conodont assem-
blage from the Carboniferous of the Avon Gorge,
Bristol, 400.

Australia: late Precambrian/Cambrian medusae, 351 ;
Ordovician stromatoporoids, 637; spirifer id brach-
iopods, 472.

B

Babinka, 173; oelandensis , 174, 34; prima, 34.

Baculatisporites comaumensis, 102.

Badam, G. L. See Tewari, B. S.

Baker, P. G. The ontogeny of the thecideacean
brachiopod Moorellina granulosa (Moore) from the
Middle Jurassic of England, 388.

Bjtostomella maniformis, 628, 114.

Batten, D. J. Some British Wealden megaspores and
their facies distribution, 333.

Bivalvia: new Ordovician from Sweden, 173.

Bohemia: Cambrian chordate with echinoderm affini-
ties, 494.

Brachiopoda: interpretation of serial sections, 321;
ontogeny of Moorellina, 388; spiriferids from
Australia, 472; structure of shells, 310; trimerellids
from Canada, 161.

Brizalina incrassata, 194

Bryozoa: skeletal growth in fenestellids, 281; Wen-
lockian, 621.

C

Callialasporites, 594; dampieri, 595, 110 ; obrutus, 597,
110; sp., 597, 110; cf. trilobatus, 594, 110.

Cambrian: chordate with echinoderm affinities, 494.

Canada: Ordovician trimerellid Eodinobolus, 161.

Carboniferous: calamite cone from England, 253;
conodont fauna from Cornwall, 262; conodont
faunas from Devon, 276; fused conodont assem-
blage, 400; growth in fenestellids, 281 ; megaspores
from Scotland, 441; miospores from Wales, 420;
new amphibian, 537; pteridosperm from Iowa,
414; seed from Illinois, 382.

Carroll, R. L. A new family of Carboniferous amphi-
bians, 537.

Cephalopoda: scale modelling of shells, 48.

Ceratocystis perneri, 494; 95-8.

Cerebropollenites mesozoicus, 109.

Chamberlain, J. A. Jr. Technique for scale modelling
of cephalopod shells, 48.

Chelinospora, 242; cassicula, 242, 42; sp. A, 243, 41.

Chordate: Cambrian with echinoderm affinities, 494.
See also Vertebrata.

Churkin, M., Eberlein, G. D., Hueber, F. M., and
Mamay, S. H. Lower Devonian landplants from
graptolitic shale in south-eastern Alaska, 559.

Cicatricosisporites, 106, 588; abacus, 107; austra-
liensis, 103, 104 ; angicanalis, 104, 105 ; brevilaesu-
ratus, 105; crassistriatus, 108; globosus, 107;
grabowensis, 108; myrtellii, 108; lucifer, 108;
purbeckensis, 588, 104; recticicatricosus, 106; spp.,
13-22; ( Anemia ) crimensis, 108; (A.) sibirica, 107.

Cincturasporites intestinalis, 429, 81, 82.

Classopollis ; echinatus, 112; hammenii, 122, 113;
torosus, 112.

Clathrodictyon, 657; aff. mammillatum, 657, 126; cf.
microundulatum, 657, 126, 127.

Cliefdenella etheridgei, 655, 125, 126.

Coccoseris sp., 124.

Coelenterata: medusae from Australia, 351; rugose
corals from France, 178; stromatoporoids from
Australia, 637.

Coelopleurus melitensis, 42, 6.

INDEX

715

Concavisporites juriensis , 102.

Conodonts: fused assemblage from Avon Gorge, 400;
Lower Carboniferous from Cornwall, 262; Lower
Carboniferous from Devon, 276.

Contignisporites dorsostriatus, 107.

Conulus ; albogalerus, 93, 94; castanea , 93; subrot m-
dus , 93.

Converrucosisporites variverrucatus, 585, 102.

Convolutispora, 425; labiata , 425, 79; vermifonnis ,
425, 79.

Coronatispora valdensis , 108.

Cothurnocystis ; americana , 98 ; elizae, 95, 98 ; primaeva ,

98.

Couperisporites comp! exus, 108, 109.

Cretaceous; benthic foraminifera, 189; crustacean
burrow, 459; echinoid with false teeth, 488;
planktonic foraminifera, 19; stratigraphic correla-
tion by miospores, 84; Wealden megaspores, 333.

Crustacea: burrows from Weald Clay, 459; trace
fossils in Jurassic, 549. See also Ostracoda.

Cryptophragmus ? sp., 651, 122.

Cyathidites ; australis, 102; minor, 102.

Cycadopites, 599; carpentieri, 111 ; cf. nitidus, 599, 111.

Cymbosporites, 239; cf. catillus, 242, 41; dittonensis,
241, 41 ; echinatus, 239, 42; verrucosus, 241, 42.

Cyrtina praecedens, 483, 92.

Cystistroma, 651; donnellii, 652, 122-4.

Cystostroma cliefdenense , 644, 117.

Cytherella recta, 1 1 5, 23.

Cytherelloidea, 115; paraweberi, 116, 23; weberi, 115,

23.

Cytheropteron, 141; aquitanum, 141, 28; sp., 142, 28.

D

Deltoidospora; psilostoma, 102; rafaeli, 102.

Densoisporites perinatus, 108.

Devonian; palaeoecology of limestone, 366; rugose
corals from France, 178; serial sections of brachio-
pods, 321; spiriferids from Australia, 472; spore
assemblages, 201; land plants from Alaska, 559.

Dicrorygma (Orthorygma), 137; kimmeridgensis, 137,
31 ; sp. I, 137, 31.

Dictyophyllidites ; equiexinus, 102; harrisii , 102.

Dictyotriletes, 226, 426; cancellatus, 427, 79 ; pactilis,
421, 79 ; submar ginat us, 427, 80 ; tesselatus, 426, 80 ;
sp. A, 226, 39 ; sp. B, 226, 39.

Disphyllum sp., 100.

Divisisporites cf. euskirchenensis, 585, 102.

Doliognathus lata, 267 , 46.

Drepanophycus sp., 569, 101.

Duplexisporites problematicus, 107.

E

Eberlein, G. D. See Churkin, M.

Ecclimadictyon, 659; amzassensis, 659, 127, 128; nes-
tori, 660,128,129.

Echinodermata : affinities with Cambrian chordate,
494. See also Echinoidea.

Echinoidea: Cretaceous with false teeth, 488; from
Miocene of Malta, 42.

Ecology: Devonian limestones, 366; patterns in shelf
benthos, 684; studies in Great Oolite, 56.

Eggert, D. A., and Kryder, R. W. A new species of
Aulacotheca (Pteridospermales) from the Middle
Pennsylvanian of Iowa, 414. See also Taylor, T. N.

Emphanisporites, 222; epicautus, 223, 38 ; micrornatus,
222, 38 ; cf. neglectus, 224, 38 ; sp. A, 224, 39 ; sp. B,
225,39.

England: calamite cone, 253; conodont faunas, 262,
276; Cretaceous miospores, 84; crustacean burrows,
459; echinoid with false teeth, 488; fused conodont
assemblage, 400; Jurassic miospores, 574; ontogeny
of brachiopod, 388; ostracods from Dorset, 112;
palaeoecology in Gt. Oolite, 56; Siluro-Devonian
spores, 201 ; trace fossils in Gt. Oolite, 549; Wealden
megaspores, 333; Wenlock bryozoa, 621; Wenlock
graptolites, 663.

Eocytheropteron decor atum , 142, 28.

Eodinobolus, 163 ; canadensis, 167, 33 ; erect us, 169, 33 ;
magnificus, 164, 32.

Equisetites burkhartii, 88.

Eucommiidites ; minor. 111; troedssonii, 111.

Exesipollenites scabrosus, 600, 111.

Exophthalmocythere fuhrbergensis, 152, 28.

F

Falcodusl sp., 278, 51.

Favreina; decemhmulatus, 549, 99; sp., 12.

Fisher, M. J. Benthonic foraminifera from the
Maestrichtian Chalk of Galicia Bank, west of Spain,
189.

Fistulipora nummulina, Ell , 114.

Florinites, 72.

Foraminifera: benthonic from Cretaceous, 189; plank-
tonic from Cretaceous, 19.

Foraminisporis wontliaggiensis, 107.

Foveosporites canalis, 106.

France: Devonian rugose corals, 178; palaeoecology
of Devonian limestone, 366.

Friend, J. K. See Funnell, B. M.

Funnell, B. M., Friend, J. K., and Ramsay, A. T. S.
Upper Maestrichtian planktonic foraminifera from
Galicia Bank, west of Spain, 19.

G

Galicia Bank: Cretaceous benthonic foraminifera, 189;
planktonic foraminifera, 19.

Galliaecytheridea, 120; confudens, 125, 25; dissimilis,
120, 24; elongata, 123, 25 ; fragilis, 130, 27; malzi,
123, 25; cf. mandelstami , 126, 26; polita , 128, 26;
postrotunda, 129, 27; punctata, 121, 25; spinosa ,
128, 26 ; sp. 1, 1 3 1 , 25 ; sp. 2, 131,27;sp. 3, 131,27;
trapezoidalis, 124, 25; wolburgi, 121, 24.

Geoclemys sivalensis, 555.

Gleicheniidites senonicus, 107.

Globotruncana, 28; area, 28, 2, 3; cf. aspera, 35, 5;
conica, 29, 3 ; contusa, 29, 3 ; falsostuarti, 30, 3 ;
gansseri, 31, 4; havanensis, 32, 4; stuarti, 33, 4;
s. stuartiformis, 34, 4.

Gnathodus, 267, 279; delicatus, 267, 46, 51 ; punctatus,
267, 46, 51 ; aff. semiglaber, 279, 51 ; texanus, 268, 46.

Gothograptus nassa, 130.

Grandispora, 433 ; reticulatus, 434, 83.

Granulatisporites, 421; visensis, 422, 78.

716

INDEX

Graptolites: Devonian from Alaska, 559; Wenlock
from England, 663.

Grumosisporites verrucosus, 78.

H

Hallam, A. Faunal realms and facies in the Jurassic, 1.

Hallidaya brueri, 356, 68, 69.

Hallopora elegant ula, 633, 116.

Halymenites striatus, 460.

Heliosporites sp., 592, 108.

Hemitrypa hibernica , 55, 56.

Heterohelix, 20; globulosa, 20, 1 ; striata, 21, 1 ;
ultimatumida, 21, 1.

Hexagonaria; cf. longiseptata , 185, 36; namnetensis,
1 79, 35, 36 ; venetensis, 184, 35.

Hibbert, F. A., and Lacey, W. S. Miospores from the
Lower Carboniferous Basement Beds in the Menai
Straits region of Caernarvonshire, North Wales,
420.

Hindeodella segaformis, 47.

Holland, C. H., Rickards, R. B., and Warren, P. T.
The Wenlock graptolites of the Ludlow District,
Shropshire, and their stratigraphical significance,
663.

Holoretiolites ( Balticograptus ) lawsoni, 666.

Hornera frondiculata, 55, 56.

Hostimella , 570; sp. A, 570, 101 ; sp. B, 571, 101.

Howellella nucula australis, 480, 91.

Hueber, F. M. See Churkin, M.

Hughes, N. F., and Moody-Stuart, J. C. A method of
stratigraphic correlation using Early Cretaceous
miospores, 84.

Hymenozonotriletes ? hastulus, 434, 83.

I

Inaperturopollenites, 597; dubius, 110, 111; sp., 597,

1 10.

India: new Pleistocene turtle from, 555.

Iowa: new Carboniferous pteridosperm from, 414.

J

Jakobson, M. E. See Kennedy, W. J., and also Mc-
Kerrow, W. S.

Januasporites tumulosus, 593, 108, 109.

Jefferies, R. P. S. Ceratocystis perneri Jaekel — a
Middle Cambrian chordate with echinoderm affini-
ties, 494.

Johnson, R. T. See Kennedy, W. J., and also Me Ker-
row, W. S.

Jurassic: faunal realms and facies, 1 ; miospores from
Purbeck, 574; ontogeny of brachiopod, 388;
ostracods from Kimmeridge Clay, 112; palaeoeco-
logy in Gt. Oolite, 56 ; trace fossils in Gt. Oolite, 549.

K

Kennedy, W. J., Jakobson, M. E., and Johnson, R. T.
A Favreina-Thalassinoides association from the
Great Oolite of Oxfordshire, 549.

— and Macdougall, J. D. S. Crustacean burrows
in the Weald Clay (Lower Cretaceous) of south-
eastern England and their environmental significance,
459.

Kier, Porter M. A Cretaceous echinoid with false teeth,
488.

Kilenyi, T. I. The ostracoda of the Dorset Kimmeridge
Clay, 112.

Klukisporites pseudoreticulatus, 105.

Knoxisporites, 428; literatus, 81; pristinus, 428, 81;
semiradiatus, stephanophorus, 428, 80.

Kryder, R. W. See Eggert, D. A.

L

Labe chia, 649; regular is, 649, 121, 124; variabilis,
650, 121.

Lacey, W. S. See Hibbert, F. A.

Lagenicula, 442; subpilosa major, 443, 84.

Leptolepidites, 586; epacrornatus, 587, 103; psarous,
586, 103.

Lister, T. R. See Richardson, J. B.

Lophotriletes tribulosus, 423, 78.

Lophozonotriletes muricatus, 431, 81.

Lycopodiacidites cerniidites, 105.

Lycopodiumsporites austroclavatidites, 105.

Lycospora uber, 82.

Lyropora quincuncialis, 52, 54.

Lytoceras fimbriatum, 7.

M

Macdougall, J. D. S. See Kennedy, W. J.

Macrodentina ( Macrodentina) , 149; cicatricosa, 149,
30 ; macidata, 149, 31 ; sp. 1, 150, 30 ; ( Polydentina ),
150; parvapunctata, 151, 31 ; proclivis proclivis, 150,
31 ; p. striata, 1 50, 30.

Malta: new Cretaceous echoniod from, 42.

Mamay, S. H. See Churkin, M.

Mandelstamia ( Mandelstamia ), 133; angulata, 134,
29; rectilinea, 133, 29; triebeli, 133, 29; ( Xero -
mandelstamia) maculata, 135, 29; sp. 1, 136, 29.

Marattisporites scabratus, 109.

Martinottiella alabamensis, 193.

Matthews, S. C. A Lower Carboniferous conodont
fauna from east Cornwall, 262.

— Two conodont faunas from the Lower Carboni-
ferous of Chudieigh, south Devon, 276.

McKerrow, W. S., Johnson, R. T., and Jakobson, M. E.
Palaeoecological studies in the Great Oolite at
Kirtlington, Oxfordshire, 56.

Meekopora dudleyensis, 628, 114.

Microreticulatisporites diatretus, 105.

Minerisporites , 343 ; aliens, 344, 66, 67 ; marginatus,
343, 65, 66, 67.

Miocene: echinoid from Malta, 42.

Mitrocystites mitra, 97, 98.

Mollusca: new Ordovician bivalve, 173; scale models
of cephalopods, 48.

Monoceratina, 145; sp. 1, 145, 29; sp. 2, 145, 29.

Monograptus, 672; deubeli, 672, 130; flemingii, 130;
ludensis, 673, 130; thomasi, 100; vulgaris, 675.

Monosulcites sp. aff. minimus. III.

Monotrypa paterella, 635.

Moody-Stuart, J. C. See Hughes, N. F.

Moorellina granulosa, 388, 73, 74.

Murospora, 430; aurita, 431, 81 ; intort a, 430, 81.

INDEX

N

Neoeponides hillebrandti , 196.

Neoraistrickia drybrookensis , 425, 78.

New South Wales: Ordovician stromatoporoids from,
637; spiriferid brachiopods from, 472.

Nodophthalmocy there tripartita , 120, 24.

Norford, B.S. and Steele, H. Miriam. The Ordovician
trimerellid brachiopod Eodinobolus from south-
east Ontario, 161.

Norris, G. Miospores from the Purbeck Beds and
marine Upper Jurassic of southern England, 574.

Nuttallides galiciensis, 194.

Nuttallinella lusitanica, 195.

Nyctopora sp., 124.

O

Ontogeny: of thecideacean brachiopod, 388.

Ophiomorpha nodosa, 460, 87, 88.

Ordovician: bivalve from Sweden, 173; stromatopo-
roids from Australia, 637; Tremadocian trilobite,
406; trimerellid brachiopod from Canada, 161.

Orthonotacythere, 143; interrupta, 143, 28; pustulata,
143, 28; sp., 144, 28.

Osmundacidites wellmanii , 102.

Ostracoda: from Kimmeridge Clay, 1 12.

Ostrea ( Liostrea ) hebridica, 12.

Owen, D. E. Wenlockian bryozoa from Dudley,
Niagara and Gotland and their palaeogeographical
implications, 621 .

P

Pachytesta bsrryvi/lensis, 382, 71, 72.

Palmatolepis , 269; gonioclymeniae, 269, 47; gracilis
gracilis, 269, 47 ; perlobata schindewolfi, 269, 47 ;
nigosa trachytera, 270, 47 ; sp., 270, 47.

Paracalamostachys spadiciformis, 257, 44, 45.

Paracypris, 117; problematica, 177, 23; sp. C, 117,
23; sp, 1, 117, 23.

Parvisaccites radiatus, 109, 110.

Peltandripites tener, 593, 110, 111.

Perinopollenites elatoides, 1 1 2.

Permian: structure of brachiopod shells, 310.

Perotrilites, 226, 433; magnns, 433, 83; microbacu-
latus, 226 ; m. microbaculatus, 227 , 39 ; m. attenuatus,
227, 39; perinatus, 433, 83; sp. A, 228, 40.

Pilosisporites, 587; delicatulus, 587, 103, 104; tricho-
papillosus, 103.

Planoglobulina acervulinoides, 22, 1.

Plantae: Devonian landplants from Alaska, 559;
new British calamite cone, 253; new pteridosperm
from Iowa, 414; new seed from Illinois, 382. See
also Spores.

Pleistocene: new turtle from India, 555.

Plicatella abaca , 591, 106, 107.

Podocarpidites sp. cf. ellipticus, 594, 109.

Polygnathus ; communis, 48, 51 ; sp., 51.

Polypora; cestriensis, 53; sp., 52, 53.

Precambrian: medusae from Australia, 351.

Pristiograptus, 668; dubius, 130; jaeger i, 668, 130;
sp. 1,671, 130.

Procytheropteron sp. 1, 142, 28.

Proreticularia beddiei, 479, 90.

717

Protocythere, 139; neali, 140, 28; rodewaldensis, 139,
27 ; sigmoidea, 1 39, 27.

Pseudoguembelina , 24; costulata, 24, 1 ; excolata, 24, 2.

Pseudokainella, 406; impar , 407, 75.

Pseudomicroplasma sp., 100.

Pseudopolygnathus, 27 1 ; triangula triangula, 271, 48 ;
tr. pinnata , 271, 48, 51 ; sp., 272, 48.

Pseudostylodictyon, 645; inequale, 646, 119; aff.
poshanense, 645, 117, 118.

Pseudotextularia elegans, 23, 1.

Ptychomaletoechia cf. gonthieri, 61.

Punctatisporites irrasus, 421, 78.

Pyramidina szajnochae, 193.

? Pyrocytheridea sp., 133, 27.

Q

Quadrithyris rob list a molongensis, 473, 89.

R

Racemiguembelina fructicosa, 25, 2.

Raistrickia\ cf. clavata , 424, 79; nigra, 424, 78.

Ramsay, A. T. S. See Funnell, B. M.

Reptilia: new Pleistocene turtle from India, 555.

Reticidisporites semireticulatus , 105.

Retusotriletes, 214; dittonensis, 215, 37; dubius, 215,
38; cf. minor, 217, 37 ; sp. A, 218, 37 ; cf. triangu-
latus, 217, 37 ; warringtonii, 216, 37.

Richardson, J. B., and Lister, T. R. Upper Silurian
and Lower Devonian spore assemblages from the
Welsh Borderland and South Wales, 201.

Rickards, R. B. See Holland, C. H.

Rosenella woyuensis, 649, 120.

Rubinella major, 587, 103.

Rugoglobigerina, 36; pustulata, 36, 5; rotundata, 37,
5 ; scott i, 37, 5.

Rugosa: Hexagonaria from France, 178.

S

Savage, N. M. New spiriferid brachiopods from the
Lower Devonian of New South Wales, 472.

Scaliognathus anclioralis, 272, 49, 51.

Schizosporis; parvus, 113 ; reticulatus, 113 ; spriggi, 113.

Schuleridea, 118; triebeli, 1 18, 23 ; sp. 1 , 1 19, 23 ; sp. 2,
119, 24.

Scotland: Carboniferous amphibian from, 537;

Visean megaspore assemblages, 441.

Sestrosporites pseudoalveolatus, 108.

Setosisporites, 445 ; indianensis, 445, 85 ; pseudore-
ticulatus, 447, 84; splendidus, 449, 85; sp. A, 451,
85 ; sp. B, 452, 85.

Sigmopollis callosus, 601 , 1 13.

Silurian: spore assemblages, 201; Wenlock grapto-
lites, 663; Wenlockian bryozoa, 621.

SiphonodeUa, 273, 279; cooperi, 279, 51 ; isostichia,
279, 51 ; obsoleta , 273, 46 ; sexplicata, 51 ; sp., 51.

Skinnera brooksi, 361, 69.

Soot-Ryen, Helen. A new species of Babinka (Bivalvia)
from the Lower Ordovician of Oland, Sweden, 173.

Sorauf, J. E. Lower Devonian Hexagonaria ( Rugosa )
from the Armorican Massif of western France, 178.

Spathognathodus scitulus, 402.

Spheripollenites subgranulatus, 110.

718

INDEX

Spinner, E. Megaspore assemblages from Visean
deposits at Dunbar, East Lothian, Scotland, 441.

Spores: British Wealden megaspores, 333; Carboni-
ferous megaspores, 441 ; Carboniferous miospores,
420; early Cretaceous miospores, 84; late Jurassic
miospores, 574; Siluro-Devonian spore assem-
blages, 201.

Steele, H. Miriam. See Norford, B. S.

Stenopora primaeva, 629, 115.

Stereisporites antiquasporites, 102.

Stratodictyon, 647; columnare, 648, 118, 119, 124;
ozakii, 647, 119, 120, 124.

Streelispora, 230; granulata, 231, 41 ; newportensis, 230,

41.

Streptorhynchus pelicanensis, 311, 57, 58, 60.

Sweden: new Ordovician bivalve, 173; Wenlockian
bryozoa, 621 .

Synorisporites, 232; downtonensis, 232, 40; tripapil-
latus, 233, 40; verrucatus, 233, 40; sp. A, 234, 41.

T

Tavener-Smith, R. Skeletal structure and growth in the
Fenestellidae (Bryozoa), 281 .

Taylor, T. N., and Eggert, D. A. On the structure and
relationships of a new Pennsylvanian species of the
seed P achy test a , 382.

Tenakea solida, 3 1 2, 58, 59.

Tertiary: See Miocene.

Tetrapterites visensis, 83.

Tewari, B. S., and Badam, G. L. A new species of
fossil turtle from the Upper Siwaliks of Pinjore,
India, 555.

Thalaminoides, 551.

Thanmopora sp., 100.

Thomas, B. A. A new British Carboniferous calamite
cone, Paracalamostachys spadicifonnis, 253.

Thomsonia, 338; alata, 338, 63, 64, 67; fairlightensis,
340, 64, 67 ; pseudoteneUa, 341, 65, 67.

Trematopora sp., 634.

Trigonorhynchia pared, 61.

Trilobita: Tremadocian from Wales, 406.

Trilobosporites, 590; apiverrucatus, 107; bernissar-
tensis, 106; domitus , 591, 106; obsitus, 590.

Tripartina sp., 590, 105.

Tryplasma altaica, 100.

U

Umbonatisporites variabilis, 423, 78, 79.

Undu/atasporites araneus, 599, 110, 111.

U. S. A. : Devonian landplants from Alaska, 559; new
pteridosperm from Iowa, 414; Pennsylvanian seed
from Illinois, 382.

V

Valentine, J. W. Patterns of taxonomic and ecolo-
gical structure of the shelf benthos during Phane-
rozoic time, 684.

Vallatisporites, 432; ciliaris, 432, 82; microgalearis,
433, 82 ; vallatus, 432, 82.

Valvalabamina scrobiculata, 196.

Vermcosisporites eximius, 422, 78.

Vertebrata: new Carboniferous amphibian, 537; new
Pleistocene turtle, 555.

Vitreisporites pallidas, 109.

W

Wade, Mary. Medusae from uppermost Precambrian
or Cambrian sandstones, Central Australia, 351.

Wales: Carboniferous miospores, 420; Siluro-

Devonian spore assemblages, 201 ; Tremadoc
trilobite, 406.

Wallace, Peigi. Specific frequency and environmental
indicators in two horizons of the Calcaire de Ferques
(Upper Devonian), northern France, 336.

Waltzispora planiangulata, 423, 79.

Warren, P. T. See Holland, C. H.

Webby, B. D. Ordovician stromatoporoids from New
South Wales, 637.

Westbroek, P. The interpretation of growth and form
in serial sections through brachiopods, exemplified
by the trigonirhynchiid sepatalium, 321.

Whitworth, P. H. The Tremadoc trilobite Pseudo-
kainella impar (Salter), 406.

Z

Zammit-Maempel, G. A new species of Coelo-
pleurus (Echinoidea) from the Miocene of Malta, 42.

Zonalesporites , 452; brasserti, 86; coniacies, 86;
fusinatiis, 452, 86 ; cf. rawosus, 86.

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A new family of Carboniferous amphibians. By R. L. carroll 537

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Wenlockian Bryozoa from Dudley, Niagara, and Gotland and their palaeo-
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© The Palaeontological Association, 1971

A CUMULATIVE INDEX

TO

Palaeontology

VOLUMES 1-12

(1 957-1 969)

ISLES STRACHAN

PUBLISHED BY THE
PALAEONTOLOGICAL ASSOCIATION
LONDON
1971

© THE PALAEONTOLOGICAL ASSOCIATION I 9 7 I

PRINTED IN GREAT BRITAIN

CONTENTS

Page

INTRODUCTION v

DATES OF PUBLICATION OF PARTS OF VOLUMES vii

GENERAL INDEX 2

STRATIGRAPHICAL INDEX 15

AUTHOR LIST 25

INTRODUCTION

At the suggestion of the Council of the Palaeontological Association, I have attempted
to compile an index to the first twelve volumes of Palaeontology. It was originally
intended to have three ‘subject’ indexes (by age, locality, and fossil group or general
topic) in addition to the list of papers by author. Unfortunately it has proved impossible
to produce a satisfactory geographical index within a reasonable space and so only the
topical and stratigraphical indexes are here. Most of the papers in Palaeontology lend
themselves easily to classification under both heads and are therefore to be found in
both indexes. Others have either a general time span or deal with matters like techniques
and thus cannot be ‘dated’ even to an era. These will then only appear in the general
index. The basic sequence used in each index is noted before it.

This cumulative index is compiled on slightly different lines from the indexes to the
annual volumes (apart from not having the generic and specific entries for fossil taxa)
and it is not intended to supersede the annual indexes entirely. It is hoped that it will
provide a quicker access to the main topics which have been dealt with in Palaeontology
between 1957 and 1969. The final result includes improvements suggested by various
members of the Executive Committee and by my colleagues in Birmingham but many
arbitrary decisions on where to classify particular papers result from my own ideas
and prejudices.

ISLES STRACHAN

Geology Department,

University of Birmingham,

Birmingham B15 2TT
1 7 May 1971

DATES OF PUBLICATION

Vol.

1,

Part

1,

pp.

1-86

28

November 1957

2,

pp.

87-158

27

May 1958

3,

pp.

159-260

11

July 1958

4,

pp.

261-411

30

January 1959

Vol.

2,

Part

1,

pp.

1-160

17

October 1959

pp.

161-280

29

March 1960

Vol.

3,

Part

i,

pp.

1-128

30

May 1960

?

•^-9

pp.

129-244

19

August 1960

3,

pp.

245-396

18

December 1960

4,

pp.

397-624

9

March 1961

Vol.

4,

Part

1,

pp.

1-148

30

April 1961

a

^-9

pp.

149-312

27

July 1961

3,

pp.

313-476

28

October 1961

4,

pp.

477-662

30

January 1962

Vol.

5,

Part

1,

pp.

1-148

12

April 1962

a

•^9

pp.

149-354

20

July 1962

3,

pp.

355-618

2

November 1962

4,

pp.

619-826

5

February 1963

Vol.

6,

Part

1,

pp.

1-218

26

April 1963

2,

pp.

219-396

27

May 1963

3,

pp.

397-596

7

October 1963

4,

pp.

597-770

10

December 1963

Vol.

7,

Part

1,

pp.

1-172

14

April 1964

2,

pp.

173-350

21

July 1964

3,

pp.

351-524

22

September 1964

4,

pp.

525-707

14

January 1965

Vol.

8,

Part

1,

pp.

1-198

26

February 1965

2,

pp.

199-373

9

July 1965

3,

pp.

375-576

28

October 1965

4,

pp.

577-767

15

December 1965

Vol.

9,

Part

1,

pp.

1-181

21

March 1966

2,

pp.

183-354

18

July 1966

3,

pp.

355-522

25

October 1966

4,

pp.

523-705

23

December 1966

DATES OF PUBLICATION

viii

Vol. 10, Part 1, pp. 1-170

2, pp. 171-337

3, pp. 339-523

4, pp. 525-720

8 May 1967
29 June 1967
24 November 1967
22 December 1967

Vol. 11, Part 1, pp. 1-162

2, pp. 163-328

3, pp. 329-490

4, pp. 491-642

5, pp. 643-817

4 March 1968
28 March 1968
2 July 1968
28 November 1968
20 December 1968

Vol. 12, Part 1, pp. 1-172

2, pp. 173-350

3, pp. 351-535

4, pp. 537-718

25 April 1969
24 July 1969
11 September 1969
19 December 1969

NOTES ON THE GENERAL INDEX

This index includes both systematic categories and topical headings (such as fauna,
flora, functional morphology, technique, trace fossils) arranged in alphabetical order.
The primary systematic division is into Class, Phylum, or equivalent, depending on the
general practical recognition of fossils, i.e. molluscs are recognized as Bivalvia or
Gastropoda, etc., while brachiopods are initially recognized at the phylum level. Cor-
respondingly the further subdivision of these groups varies. There are sufficient entries
under Ammonoidea for a stratigraphical subdivision to be useful while recognition of
the superfamilies is not within the knowledge of many palaeontologists. Similarly
trilobites are subdivided by age even though this results in the separation of two papers
on Encrinurus. Brachiopods, on the other hand, are treated further systematically since
recognition of the main groups of articulate brachiopods is fairly common palaeon-
tological knowledge and the stratigraphical arrangement can be obtained from the
other index. This arrangement seems to be the best practical one and corresponds to
some extent at least with the interests of palaeontologists themselves.

The entries include reference to the volume and page of Palaeontology on which the
paper referred to begins as well as the index number (in italics) from the Author List.

GENERAL INDEX

AGNATHA

Cephalaspis lyelli Agassiz, 1, 99 (482).

Downtonian ostracoderm Corvaspis, 3, 217 (437).

New evidence on Jamoytius, 11, 21 (353).

Traquairaspis from Canada, 7, 210 (139).

ALGAE

Algal debris-facies in Middle East Cretaceous, 1, 254 (154).

Algal growths in English Rhaetic, 4, 324 (190).

Algomycetes and Tasmanacea from Brazilian Palaeozoic, 10, 640 (407).

Calcareous algae: Cretaceous Permo calculus, 4, 82 (755); from the Dalradian of Islay, 5, 268 (752);
new Tertiary, Subterraniphyllum, 1, 73 (153).

Coccoliths: from Atlantic seamounts, 7, 306 (40); taxonomic problems in, 11, 793 (47); Up. Cre-
taceous from Zululand, 11, 361 (333).

Codiaceae: some Cretaceous, 8, 199 (759); Namurian of N. England, 1, 147 (238); Visean of Scot-
land, 8, 192 (58).

Dasycladaceae : new Carboniferous genus of, 7, 181 (498); three new Tethyan, 11, 491 (160).
Girvanella: British Carboniferous, 6, 264 (497); Silurian from Welsh Borderland, 9, 48 (239); type
species of, 1, 22 (496).

Problematical microfossils from Middle East, 6, 293 (156).

Solenoporaceae, reproductive structures in Tertiary, 7, 695 (755).

AMMONOIDEA

Palaeozoic

Abnormal growths in Devonian goniatites, 3, 129 (205).

Alaskan goniatite from Ireland, 1, 158 (196).

Carboniferous goniatite zones in S.W. England, review, 3, 75 (60).

Cornish ammonoids and trilobites, 3, 153 (393).

Delepinoceras in North America, 7, 173 (777).

Devonian goniatites and correlations in Canada, 6, 491 (207).

Eumorphoceras, new from Ireland, 4, 54 (501).

Goniatite fauna from Germany, 11, 264 (165).

Goniatites striatus from Ireland, 1, 384 (797).

Namurian goniatites from Eire, 5, 355 (502).

New South Wales: clymeniid, 3, 237 (332); Famennian ammonoids, 11, 535 (235); Up. Devonian
Cheiloceras, 9, 458 (234).

Nuculoceras stellarum, 8, 226 (799).

Palaeoecology of Goniatite Bed, Castleton, 8, 186 (168).

Supposed earliest clymeniid, 3, 472 (206).

Visean goniatites from N. Staffs., 1,16 (39).

Mesozoic (general)

Yorkshire type ammonites, 5, 93 (208).

Jurassic

Epizoic oysters on Kimmeridgian ammonites, 11, 19 (706).

Juraphyllitidae in Britain, 7, 286 (209).

Kimmeridgian ammonites from drift of Lincolnshire, 6, 219 (75).

Liassic Neomicvoceras and Paracymbites, 9, 312 (141).

Propectinatites, 11, 16 (105).

Rasenia from Scotland, 5, 765 (509).

GENERAL INDEX

3

Toarcian Pseudolillia from Spain, 5, 86 (140).

Variation and ontogeny in Oxfordian ammonites, 9, 290 (314), 10, 60 (315).

Cretaceous

Ammonites from Bathurst Island, N. Australia, 6, 597 (500).

Barremian of North Bulgaria, 5, 527 (278).

Biometric study of Barremites subdijficilis, 6, 727 (345).

Gault Hengestites, 2, 200 (71).

Leymeriella in Britain, 1, 29 ( 70).

Scaphites, origin, limits and systematic position, 8, 397 (485).

Speeton Clay: from Lower D Beds, 5, 272 (301); new heteromorph from, 6, 575 (145); phyl-
loceratid from, 9, 455 (343).

AMPHIBIA

Growth stages of branchiosaurs, 6, 540 (472).

New Carboniferous family, 12, 537 (67).

ANGIOSPERMS

Probable angiosperm pollen from Britain, 11, 421 (254).

Wood: Eocene Anacardiaceae, 9, 360 (48); Eocene oak, 3, 86 (47).

ANTHOZOA, see Corals

ARTHROPODA

Eurypterida: median abdominal appendage of Slimonia, 3, 245 (469); new from Scottish Old Red
Sandstone, 5, 137 (470); ventral anatomy of Carboniferous Anthraconectes, 7, 474 (491).
External anatomy of Carboniferous ‘scorpions’, 1, 261 (488), 3, 276 (489).

Insecta: beetles from Antarctica, 1, 407 (505); Carboniferous nymph, 10, 307 (360); Liassic dragon-
fly, 1, 406 (504); Rhaetic and Liassic beetles, 4, 87 (172); Tertiary Homoptera of Stravropol,
10, 542 (36).

See also Crustacea, Ostracoda, Trilobita.

BELEMNOIDEA, see Cephalopoda.

BIBLIOGRAPHY: Catalogue of Begg Collection, Glasgow, 6, 378 (124).

BIVALVIA (= Lamellibranchia).

General

Babinka: affinities, life habits, 8, 231 (275); new species from Ordovician of Oland, 12, 173 (408).
Epizoic oysters on Kimmeridgian ammonites, 11, 19 (106).

Functional studies on Arctostrea, 11, 458 (69).

Intestine of Liassic Nuculana preserved, 2, 262 (115).

Ligaments preserved in Australian Permian bivalves, 11, 94 (376).

Microstructure and mineralogy of Jurassic mytilid, 11, 163 (213).

Mode of life of Jurassic ’’Posidonia' , 8, 156 (230).

Salinity-controlled assemblages from Middle Jurassic, 6, 318 (211).

Size-frequency and growth-ring analyses, 6, 731 (119).

Size-frequency distributions in death-assemblages, 10, 25 (187).

Devonian

Prothyris in Britain, 6, 1 36 (495).

Carboniferous

Caneyella, Posidonia and PosidonieUa, distinctions, 1, 405 (342).

Naiadites obesus from Fife, 4, 300 (38).

Namurian from Eire, 5, 355 (502).

‘Nebraskan’ fauna in Scotland, 4, 507 (494).

Non-marine: from East Fife, 3, 137 (37); from Pembrokeshire, 3, 104 (233); new, 5, 307 (147).
Wilkingia to replace AUorisma, 1 , 401 (493).

4

GENERAL INDEX

Permian

Dentition of Permophorus costatus, 7, 281 (269).

Eurydesma from Dwyka Beds, 4, 138 (134).

Merismopteria and origin of Pteriidae, 3, 387 (135).

New records from Eastern Australia, 4, 119 (137).

New Zealand Atomodesma, 6, 699 (467).

Palaeotaxodonts from New Zealand, 7, 630 (468).

Mesozoic: new genera and subgenera, 4, 592 (116).

Trias

From Oman Peninsula, 4, 1 (218).

Photonegative young in Lima lineata, 3, 362 (228).

Pteromya, 6, 582 (118).

Jurassic

Ecology and stratigraphical distribution in Great Estuarine Series, 6, 327 (212).
Malayomaorica from Indo-Pacific Upper Jurassic, 6, 148 (232).

Cretaceous

Characters and relationships of Pseudavicula, 3, 392 (136).

Tertiary

From Libya, 5, 1 (117).

Teredinid from Iraq, 6, 315 (157).

BLASTOIDEA, see Echinodermata.

BRACHIOPODA

General

Anchorage of articulates on soft substrata, 4, 475 (372).

Fabrics of shells of Terrakea and Streptorhynchus, 12, 310 (17).

Function of zig-zag deflexions in the commissure, 7, 135 (373).

Interpretation of serial sections, 12, 321 (480).

Life assemblages from the Lias, 4, 653 (184).

Origin of the loop in articulates, 4, 149 (486).

Predation and shell damage in Visean fauna, 9, 355 (52).

Sectioning of steinkerns, 7, 105 (415).

Shell structure of billingsellaceans, 11, 486 (487).

Faunas

Devonian: from Nevada, 9, 152 (240); from Somerset, 7, 1 (474); from Canada, 3, 208 (322).
Llandeilo from N. Wales, 4, 177 (272).

Triassic from Oman Peninsula, 4, 1 (218).

Inarticulata

Dinobolus from Silurian of British Columbia, 3, 242 (304).

Eodinobolus from Ordovician of Ontario, 12, 161 (305).

Valdiviathyris from Indian Ocean, 4, 542 (370).

Orthida

Cortezorthinae, new subfamily, 10, 142 (242).

Four British Ordovician species, 1, 292 (76).

Isorthis and Salopina in Ludlovian, 8, 454 (465).

Planicardinia from Devonian of New South Wales, 11, 627 (385).

Schizophoriids : Carboniferous from Europe, 11, 64 (337); Devonian from Europe, 9, 381 (336);

mantle canal patterns, 11, 389 (355).

Visbyella — a new genus from Silurian, 11, 306 (466).

Triplesiacea, morphology of superfamily, 5, 740 (499).

Pentamerida

Brachial structure, 7, 220 (12).

Unusual stricklandiids from Wales, 9, 346 (506).

GENERAL INDEX

5

Strophomenida

Delepineci from New South Wales, 7, 514 (66).

Feeding mechanism and affinities of Thecospira and Bactrynium, 11, 329 (375).

Feeding mechanism of Prorichthofenia, 3, 450 (371).

Horridonia from the Permian, 4, 42 (175).

Howesia, a new Permian genus, 6, 754 (268).

Leptaena from English Wenlock and Lr. Ludlow, 10, 590 (253).

Llandovery stropheodontids from Welsh Borders, 10, 245 (96).

Parachonetes from the Devonian, 9, 365 (241).

Pedicle sheath of young productaceans, 7, 703 (57).

Pennsylvanian Juresania nebrascensis, 7, 23 (162).

Scottish Visean productid, 9, 426 (394).

Thecideacea: morphology of Moorellina, 12, 388 (20).

Rhynchonellida

Austmlirhynchia , new Devonian genus, 11, 731 (386).

Carboniferous Pugnoides, 3, 477 (318).

Sensory spines in Acanthothiris , 8 , 604 (374).

The true Rhynchonella, 1, 1 (4).

Atrypida

Dayia navicula, 11, 612 (453).

Devonian atrypids from England, 8, 358 (108).

Eocoelia hemisphaerica and related species, 9, 523 (507).

Leptocoeliidae, revision of family, 6, 440 (11).

Spinatrypa and Spinatrypina, 10, 489 (109).

Spiriferida

Brachial skeleton of Attenuatella convexa, 11, 783 (16).

Delthyrial cover in Mucrospirifer, 11, 317 (114).

Eospiriferidae, 5, 682 (45).

Martiniopsis- like spiriferids from Queensland, 1, 333 (62).

New Devonian spiriferids from New South Wales, 12, 472 (387).

Shell development in Spirifer trigonalis, 4, 477 (146).

Type species of three Up. Palaeozoic punctate spiriferoids, 1, 351 (63).

Terebratulida

English Aptian Terebratulidae, 2, 94 (287).

Shell-growth in Recent terebratuloids, 10, 298 (384).

BRYOZOA (= Polyzoa)

Caradoc: trepostomes from Shropshire, 6, 1 (367); early Phaenopora, 5, 52 (365); Homotrypa and
A mplexopora ?, 8, 5 (369).

Chazyan leptotrypellid and atactotoechids, 5, 727 (366).

Cretaceous Pyripora and Rhammatopora, 3, 370 (447).

Cupuladria canariensis, 6, 172 (263).

Dionella, new Cretaceous genus, 8, 492 (284).

Fenestrate: new fenestrate bryozoan from Carboniferous, 8, 478 (438); micrometric formula in
classification, 9, 413 (439); Polypora , 6, 166 (291); skeletal structure and growth, 12, 281 (440);
type Fenestella, 4, 221 (289); Wenlockian from England, 5, 540 (290).

Permian from W. Australia, 6, 70 (368).

Silurian: from Canadian Arctic, 9, 517 (43); from Central Wales, 3, 69 (310); from Ludlow, 5,
195 (311); Wenlockian from Dudley, Niagara and Gotland, 12, 621 (312).

Types of seven Ordovician bifoliates, 3, 1 (330).

CEPHALOPODA

Belemnites: Dicoelites and Prodicoelites, 7, 606 (421); new belemnite from Indonesia, 7, 621 (422);

Op pel’s specimens of B. gerardi, 6, 690 (420).

Namurian fauna from Ireland, 5, 355 (502).

6

GENERAL INDEX

Nautiloid: Leurocycloceras from Welsh Silurian, 7, 525 (201).

Technique for scale modelling shells, 12, 48 (83).

Visean fauna from New South Wales, 7, 682 (50).

Yorkshire type ammonites and nautiloids, 5, 93 (208).

See also Ammonoidea.

CHITINOZOA

Ordovician chitinozoa from Shropshire, 10, 436 (236).

CHORDATA

Cambrian chordate with echinoderm affinities, 12, 494 (231). See also individual Vertebrate classes.
CIRRIPEDIA: See Crustacea.

COELENTERATA

Eocene sea-pen from New Zealand, 1, 266 (189).

Medusae from Precambrian/Cambrian, Australia, 12, 351 (459).

Medusoid (?) from Silurian of England, 11, 610 (424).

See also Corals, Stromatoporoidea.

CONODONTOPHORIDA
Apatognathus from Yoredales, 10, 124 (457).

Assemblage from Carboniferous, 12, 400 (18).

Cornwall: Carboniferous fauna, 12, 262 (281).

Devon: Carboniferous fauna, 12, 276 (282).

CORALS

General

Devonian corals from Somerset, 7, 1 (474).

Lower Palaeozoic corals from New South Wales, 4, 334 (425).

Periodicity in Devonian coral growth, 7, 552 (389).

Sequence of Ludlovian — Couvinian faunas in U.S.S.R., 10, 660 (195).

Rugose

Silurian: Microplasma from Monmouthshire, 9, 148 (481).

Devonian: Alaiophyllum from Canada, 6, 132 (323); correlation of Canadian M. Devonian, 7,
430 (324); Hexagoraria from France, 12, 178 (409); Marisastridae from Devon, 10, 266 (390);
Metriophyllum, 7, 108 (203); New South Wales corals, 8, 518 (426), 9, 544 (427), 10, 426 (428);
N. American Smithiphyllum, 8, 618 (325).

Carboniferous: corallum increase in Lithostrotion, 8, 204 (245); hysterio-ontogeny of Lonsdaleia
and Thysanophyllum, 10, 617 (246); Slimoniphyllum gen. nov., 4, 280 (247). Permian: corals
from northern Iraq., 1, 174 (215).

Tabulate

Cystihalysites from Wenlock and Dudley, 7, 452 (433).

Devonian from Devon, 11, 44 (204).

Dimorphism in Striatopora, 9, 488 (307).

Squamulate favositids of Victoria, 3, 186 (327).

CRINOIDEA, see Echinodermata.

CRUSTACEA

Cirripede: new species from Gault, 8, 629 (98); pedunculate from Silurian, 6, 161 (490); probable
burrows, 1, 397 (243).

Crabs in an Eocene nodule, 4, 85 (97).

Cuticle of Silurian ceratiocaridids, 5, 30 (359).

Cyclas from Mendip Hills, 10, 317 (177).

GENERAL INDEX

7

Phyllocarid: Caryocaris from N. America, 9, 371 (87); Palaeozoic from Czechoslovakia, 6, 97 (86).
Syncarid from Stephan ian of Warwickshire, 4, 546 (355).

Traces: burrows in Weald Clay, 12, 459 (255); Favreina-Thalassinoides association in Jurassic, 12,
549 (256).

See also Ostracoda.

ECHINODERMATA

General

Lower Palaeozoic faunas of the British Isles and Balto-Scandia, 2, 161 (344).

Blastoid: Devonian Belocrinus from France, 9, 244 (274); Pentablastus from Spain, 6, 471 (244).
Crinoid: British Ludlovian, 1, 106 (341); Devonian inadunates from Somerset, 4, 538 ( 473), 8,
11 (475); earliest known, 11, 406 (33); form and function of stem, 11, 275 (392); Jurassic from
New Zealand, 2, 150 (410); Ordovician from N. Wales, 8, 355 (32).

Cystoid: function of pore-structures, 11, 697 (321); Macrocystella , 11, 580 (320).

Ceratocystis, a chordate with echinoderm affinities, 12, 494 (231).

Edrioblastoid : Ordovician from Australia, 11, 513 (476).

Ophiuroid : Cretaceous from Australia, 6, 579 (400) ; traces from Namurian of Ireland, 7, 508 (265).
See also Echinoidea.

ECHINOIDEA

Cidarites moniliferus and the status of Eucidaris, 5, 785 (325).

Classification of cassiduloids, 6, 718 (329).

Cretaceous echinoid with false teeth, 12, 488 (257).

Dentition and relationships of Pygaster, 4, 243 (256).

Evidence for the age of Myriastiches gigas, 10, 171 (266).

New Albian echinoid from Kent, 3, 260 (73).

New species of Coelopleurus from Miocene, 12, 42 (503).

Pedicellariae of Silurian echinoids, 11, 576 (42).

Re-interpretation of Bothriocidaris , 10, 525 (319).

Salenia in the eastern Pacific, 7, 331 (510).

Spines and fascioles of Echinocorys, 6, 458 (418).

EURYPTER1DA, see Arthropoda.

FAUNAS, general
Precambrian

Fossils from Ediacara, South Australia, 9, 599 (174).

Palaeozoic

Described and figured specimens in the Begg Collection, 6, 378 (124).

Graptolite assemblages and zones of Birkhill shales, 11, 654 (448).

Specific frequency and environmental indicators in Upper Devonian of France, 12, 366 (464).
Palaeontology of Namurian rocks of Slieve Anierin, Eire, 5, 355 (502).

New South Wales : Lower Carboniferous fauna, 8, 54 (356); Carboniferous fossils from, 4, 428 (64).
Jurassic

Faunal realms and facies, 12, 1 (188).

Great Estuarine Series: ecology and stratigraphical distribution of fossils, 6, 327 (212); salinity-
control mollusc assemblages, 6, 318 (211).

Cretaceous

Stratigraphical palaeontology of the Lower Greensand, 3, 487 ( 74).

FLORA, general

Middle Triassic from Cachuta Formation, Argentina, 10, 564 (226).

FORAMINIFERA

Devonian: from Western Australia, 3, 397 (121), 31, 601 (99).

Carboniferous: fusulinids from Spitsbergen, 2, 210 (167); graphical aids in description of fusulines,
10, 322 (126).

8 GENERAL INDEX

Permian: from British Honduras, 5, 297 (563); from Peru, 5, 817 ( 364 ); from Spitsbergen, 2, 210
(767).

Mesozoic, general: adherent foraminifera, 1, 116 (25); 5, 149 (2).

Jurassic: arenaceous from type Kimeridgian, 1, 298 (267); from Ampthill Clay, 4, 520 (178);

Brotzenici and Voorthuysenia, 6, 653 (110); new information on Pfenderina, 4, 581 (405).
Cretaceous: alveolinid from Jamaica and Mexico, li, 526 (357); Bolivinoides from Britain, 9, 220
(29); Chalk Marl palaeoecology, 4, 599 (59); from Ireland, 9, 492 (30); from Isle of Wight, 4,
552 (28); Maestrichtian from Galicia Bank, 12, 19 (170), 189 (166); Marssonella and Pseudotex-
tulariella from England, 6, 41 (27); Polymorphinidae from England, 5, 712 (26).

Tertiary, general: classification and distribution of Globigerinaceae, 2, 1 (22); large forams from
Central America, 11, 283 (149); Globigeriiioides ruber, Miocene to Recent, 10, 647 (111).
Palaeocene: Heterostegina, 10, 314 (148); new nummulitid, 11, 435 (756).

Eocene: adherent from France, 5, 149 (2); Assam Discocyclina, 6, 658 (380); Assam Nummulites,
11, 669 (379); Cassigerinella from Florida, 11, 368 (772); Orbulinoides beckmanni, 11, 371 (773);
Operculinoides, 2, 1 56 (297).

Oligocene: stratigraphical distribution of Archaias, 1, 207 (404).

Miocene: distribution of Discospirina, 1, 364 (7).

Pliocene: Alliatina and Alliatinella, 1, 76 (68).

Holocene: forams in marsh cycles in Wales, 8, 27 (3).

FUNCTIONAL MORPHOLOGY
Cretaceous oyster Arctostrea, 11, 458 (69).

Dichoporite pore-structures in cystoids, 11, 697 (327).

Feeding mechanism of branchiopod Prorichthofenia, 3, 450 (377).

Feeding mechanism of Thecospira and Bactrynium 11, 329 (375).

Schizochroal eyes and vision in trilobites, 9, 1 (97), 464 (92).

Sensory spines in Acanthothiris, 8, 604 (374).

Stem of pseudoplanktonic crinoid, 11, 275 (392).

Zigzag deflexions in brachiopod commissures, 7, 135 (373).

GASTROPODA

Apical development in turritellid classification, 8, 666 (7).

Australasian Typhinae, 4, 362 (458).

Carboniferous from Queensland, 4, 59 (283).

Devonian Orecopia from Canada, 9, 142 (326).

Discohelix in the Tethyan Jurassic, 11, 554 (479).

Fossilized intestines from Lower Cretaceous, 2, 270 (72).

Nomenclatural corrections, Lower Greensand, 4, 312 (75).

Permian Platyteichum from Australia, 4, 131 (733).

Peruvispira from S. Africa, 4, 138 (134).

Sinus-bearing monoplacophoran, 11, 132 (361).

GRAPTOLITHINA

Assemblages and zones of Birkhill Shales, 11, 654 (448).

Development of a dicellograptid, 8, 41 (227).

Development of Lasiograptus harknessi, 8, 272 (352).

Diplograptids: from British Lower Silurian, 5, 498 (313); from British and Scandinavian Llan-
dovery, 11,1 (57).

Glyptograptus dentatus and allied species, 6, 665 (56).

Monograptus, new species from Yukon, 6, 751 (225).

New variety of Orthoretiolites, 2, 226 (399).

Sequence of graptolite faunas, 1, 159 (55).

Silurian: new from Howgill Fells, 8, 247 (357); from Illinois, 5, 59 (362).

Tremadocian from Norway, 6, 121 (413).

Wenlock from Ludlow district, 12, 663 (202).

GENERAL INDEX

9

GYMNOSPERMOPHYTA

Amyelon: in American coal balls, 7, 186 (722); revision of genus, 5, 213 (24).

Fossil cycads, 4, 313 (191).

Opposite-leaved conifer from Jurassic of Israel, 2, 236 (79).

Structure of Vertebraria indica, 11, 643 (317).

See also Pteridospermopsida.

HYSTRICOSPHERES, see Microplankton.

INSECTA, see Arthropoda.

LAMELLIBRANCHIA, see Bivalvia.

MAMMALIA

European Proviverrini, 8, 638 (456).

Growth gradients in monotremes and marsupials, 6, 615 (419).

Late Jurassic fossils in Cambridge, 6, 373 (95).

Miocene anthropoids from India, 7, 124 (340).

Odontoma in a northern mammoth, 7, 674 (223).

Wealden fossils, 6, 55 (94).

MEDUSAE, see Coelenterata.

MICROFOSSILS

Composition of Pyritosphaera, 6, 119 (271).

Micro-organisms and syngenetic pyrite, 5, 444 (270).

Problematical microfossils from the Middle East, 6, 293 (156).

U. Jurassic and L. Cretaceous microfossils from Hautes-Alpes, 8, 391 (454).

See also Foraminifera, microplankton, Ostracoda, Protozoa, spores.

MICROPLANKTON

General

Interpretation and status of hystricosphere genera, 6, 83 (143).

Silurian

Wenlock Shale acritarchs, 2, 56 (742); 6, 625 (144).

Devonian

Reef-controlled distribution in Alberta, 4, 392 (416).

Permian

Hystricospheres from Britain, 5, 770 (462).

Trias

Fine structure of some acritarchs, 9, 351 (285).

Jurassic

From Ampthill Clay of Yorkshire, 5, 478 (382).

From Australia and New Guinea, 2, 243 (100).

From Kellaways Rock and Oxford Clay of Yorkshire, 4, 90 (381).

New name for species of Gonyaulacysta, 7, 472 (383).

Cretaceous

From Cambridge Greensand, 7, 37 (104).

Tertiary

Tasmanites and Leiospheres from Louisiana, 8, 16 (163).

Quaternary

Fossil microplankton in Caribbean deep-sea cores, 10, 95 (463).

See also Chitinozoa.

MOLLUSCA

Sinus-bearing monoplacophoran and classification of primitive mollusca, 11, 132 (361).
See also Bivalvia, Gastropoda, Cephalopoda, Ammonoidea.

C 8439 B

10

GENERAL INDEX

NAUTILOIDEA, see Cephalopoda.

OSTRACODA

Ordovician

Siiicified fossils from South Wales, 6, 254 {414).

Silurian

From Stonehouse Formation, Nova Scotia, 3, 93 {107).

Carboniferous

Non-marine fauna from Durham and Northumberland, 9, 667 {338).

J urassic

Freshwater from Oxfordshire, 8, 749 {31).

From Dorset Kimeridge Clay, 12, 112 {260).

Mandelstamia from English Mesozoic, 3, 439 {299).

New genus, Oertliana, from NW. Europe, 8, 572 {258).

Cretaceous

From Bargate Beds in Surrey, 7, 317 {250).

From California, 7, 393 {198).

From Sutterby Marl of Lincolnshire, 8, 375 {251).

From Tealby Clay of Lincolnshire, 9, 208 (252).

Law of ostracod growth, 7, 85 (13).

Ontogeny of Theriosynoecum fittoni, 7, 72 (406).

Non-marine from Ghana, 11, 259 (262).

Speeton Clay: Neocythere, 6, 274 (248); Ortlionotacy there , 6, 430 (249).

Weald Clay marine brackish bands, 11, 141 (259).

Quaternary

Normanicy there and division of the Trachyleberididae, 2, 72 (298).

Normanicythere leioderma in North America, 4, 424 (300).

PALAEOECOLOGY

Actinocamax plenus Subzone in Anglo-Paris Basin, 4, 609 (229).

Brachiopod ecology and Lower Greensand palaeogeography, 5, 253 (288).

Colour markings in phacopids, 11, 498 (161).

Crustacean burrows in Weald Clay, 12, 459 (255).

Ecology and stratigraphical distribution of bivalves in Great Estuarine Series, 6, 327 (212).
Environmental causes of stunting, 8, 132 (186).

Foraminifera of the Chalk Marl, 4, 599 (59).

Frodingham Ironstone (Lower Jurassic), 6, 554 (185).

Goniatite Bed at Cowlow Nick, Derbyshire, 8, 186 (168).

Great Oolite at Kirtlington, Oxfordshire, 12, 56 (277).

Llandovery transgression of the Welsh Borderland, 11, 736 (508).

Marine benthos, substrate and palaeoecology, 9, 30 (120).

Mode of life of two Jurassic ‘ PosidonicC (Bivalvia), 8, 156 (230).

Patterns of taxonomic and ecological structure of the shelf benthos, 12, 684 (455).

Periodicity in Devonian coral growth, 7, 552 (389).

Photonegative young in Lima lineata, 3, 362 (228).

Population studies in the Visean of NW. Ireland, 9, 252 (210).

Predation and shell damage in a Visean brachiopod fauna, 9, 355 (52).

Size-frequency distributions in molluscan death-assemblages, 10, 25 (187).

Size-frequency and growth-ring analyses and their palaeoecological significance, 6, 731 (119).
Transition zone across an Upper Cretaceous boundary, New Jersey, 7, 266 (261).

PISCES

Antiarchs, additions to our knowledge of, 4, 210 (471).

Birgeria from Rhaetic, 9, 135 (388).

Dipnoan skull roof and palate, 8, 634 (65).

GENERAL INDEX

11

Haplolepid fauna from Pennsylvanian of Nova Scotia, 5, 22 (19).

Otoliths from English Jurassic, 11, 246 (423).

Palaeozoic fishes from South Africa, 5, 9 (173).

Triassic Saurichthys, 5, 344 (180).

POLLEN, see spores.

PROTOZOA, see Foraminifera, Radiolaria.

PTER1DOPHYTA

Devonian

Land plants from graptolite shale, Alaska, 12, 559 (88).

New plant from South Wales, 11, 683 (151).

Lycopsida

Fertile lycopod from Scottish Carboniferous, 8, 281 (8).

Revision of Eskdalia Kidston, 11, 439 (445).

Selaginellites with Densosporites microspores, 1, 245 (77).

Sporangiostrobus with Densosporites microspores, 5, 73 (81).

Sporophyll from Coal Measures of Somerset, 11, 445 (46).

Pteropsida

Biscalitheca from Pennsylvanian of Illinois, 11, 104 (331).

Structure and relationships of Radstockia, 10, 43 (441).

Sphenopsida

Calamitean plants from Scottish Lower Carboniferous, 6, 408 (84)

New British calamite cone, 12, 253 (446).

New calamitalean cone from Illinois, 8, 681 (193).

On the genus Pothocites Paterson, 8, 107 (85).

Probable dispersed spores of Equisetites, 11, 633 (34).

PTER1DOSPERMS

Calathospermum fimbriatum, a Scottish Carboniferous cupule, 3, 265 (23).

Mariopteris from NW. Spain, 10, 694 (460).

New species of Aulacotheca from Iowa, 12, 414 (152).

Peltaspermaceae, a Permian and Triassic family, 3, 333 (449).

Probable microsporangiate fructification from Illinois, 7, 60 (132).

Structure of leaves of Rhabdotaenia from India, 6, 301 (316).

Structure and relationships of a new species of Pachytesta, 12, 382 (442).

Three fructifications from Scottish Lower Carboniferous, 5, 225 (402).

Triassic plants of Argentina: trunk of Rhexoxylon, 11, 236 (49); the leaf Dicroidium and its relation-
ship to Rhexoxylon, 11, 500 (14).

RADIOLARIA

Radiolaria from the Namurian of Derbyshire, 9, 319 (200).

REPT1LIA

‘Dwarf' crocodiles of the Purbeck, 10, 629 (237).

Gastric contents of an ichthyosaur from Dorset, 11, 376 (339).

Histology of dinosaur bone, 5, 238 (123).

Jurassic dinosaur Scelidosaurus, 11, 40 (303).

New pleurodiran turtle from Eocene of Somalia, 9, 511 (461).

New turtle from Pleistocene of India, 12, 555 (444).

Pliosaurus brachyspondylus from Kimeridge Clay, 1, 283 (435).

Pliosaurus macromerus, scapula, 1, 193 (434).

Stretosaurus, a giant pliosaur, 2, 39 (436).

Vertebrate evidence of a southern transatlantic connection in Trias, 10, 554 (44).

GENERAL INDEX

12

SPORES and POLLEN
Megaspores

Assemblages from Visean deposits, Scotland, 12, 441 (412).

Devonian from Canada, 1, 321 (78).

Devonian from Wyboston borehole, Bedfordshire, 10, 1 89 (294), 706 (295).

Seed megaspore from Devonian of Canada, 7, 29 (82).

Two cristate megaspores from Scottish Carboniferous, 9, 488 (9).

Wealden megaspores and their facies distribution, 12, 333 (15).

Westphalian megaspores from Forest of Dean, 8, 82 (411).

Miospores (including microspores)

Silurian

Assemblages from Welsh Borders and South Wales, 12, 201 (349).

Devonian

Assemblages from Welsh Borders and South Wales, 12, 201 (349).

From Melville Island, Canadian Arctic, 3 , 26 (276).

From Vestspitsbergen, 8, 687 (5); 30, 280 (6).

Middle Old Red Sandstone: Cromarty, 3, 45 (346); Orcadian basin, 7, 559 (348).
Spores with bifurcate processes, 5, 171 (347).

Carboniferous

Assemblages from coals and sediments in Yorkshire, 7, 656 (279).

Distribution of assemblages in Western Pennsylvania, 9, 629 (181).

From Basement Beds, Menai Straits, N. Wales, 12, 420 (194).

From Drybrook Sandstone, Forest of Dean, 7, 351 (431).

From North-West England, 10, 1 (61).

From Spitsbergen, 5, 550, 619 (334).

From Springer Formation, Oklahoma, 10, 349 (164).

Microbiological attack on miospores, 6, 349 (293).

Namurian from southern Pennines, 4, 247 (302).

Sections of dispersed spores, 5, 247 (220), 679 (133).

Simozonotrilites, 1, 125 (429).

Spore-bearing structure, Tetrapterites visensis, 7, 64 (430).

Structure of spore wall in Cingulati, 3, 82 (401).

Toumaisian spores from Ayrshire, 11, 116 (432).

Permian

Assemblage from Iraq, 7, 240 (398).

British saccate and monosulcate miospores, 8, 322 (90).

New British spore, 4, 648 (80).

Trias

From Western Australia, 6, 12 (21).

Keuper from Worcestershire, 8, 294 (89).

Jurassic

From Purbeck Beds and marine Up. Jurassic, 12, 574 (306).

Further interpretation of Eucommiidites, 4, 292 (219).

Cretaceous

Hoegisporis from Australia, 3, 485 (101), 8, 39 (103).

Method of stratigraphic correlation using miospores, 12, 84 (222).

Reappraisal of Aequitriradites, 4, 425 (102).

Revision of some Belgian microspores, 6, 282 (131).

Schizaeaceous spores from macrofossils, 9, 274 (221).

Pollen

Angiosperm from British Lower Cretaceous, 11, 21 (254).

Aqudapollenites in British Isles, 11, 549 (280).

STRATIGRAPHY

Llandovery transgression of the Welsh Borderland, 11, 736 (508).

GENERAL INDEX

13

Method of stratigraphic correlation using miospores, 12, 84 (222).

Stratigraphic palaeontology of the Lower Greensand, 3 , 487 ( 74).

Time in stratigraphy, 8, 113 (292).

STROMATOPOROIDEA
Actostroma from Jurassic of Israel, 1, 87 (214).

Microstructure of stromatoporoids, 9, 74 (417).

New species of Komia and systematic position of genus, 6, 246 (492).

Ordovician from New South Wales, 12, 637 (477).

Revision of some Jurassic from Yugoslavia, 2, 28 (216).

Some Tethyan Jurassic from Middle East, 2, 180 (217).

TECHNIQUE

Glass fibre resin casts of fossils, 3, 124 (354).

Graphical aids for description and analysis of fusulines, 10, 322 (126).

Improved method of analysing distortion in fossils, 9, 125 (391).

Serial sections: interpretation of growth and form, 12, 321 (480)', of steinkerns, 7, 105 (475); rapid
grinding machine for, 6, 145 (192).

Technique for scale modelling of cephalopod shells, 12, 48 (83).

TRACE FOSSILS

Crustacean burrows in Weald Clay, 12, 459 (255).

Favreina-Thalassinoides association in English Jurassic, 12, 549 (256).

Kulindrichnus langi from Lias, 3, 64 (183).

Probable cirripede, phoronid and echiuroid burrows within an echinoid test, 1, 397 (243).

Starfish traces from Irish Namurian, 7, 508 (265).

TRILOBITA

Cambrian

Alimentary caeca of agnostids and other trilobites, 3, 410 (308).

Estaingia from South Australia, 7, 458 (335).

From Amanos Mountains, Turkey, 4, 71 (729).

From Pioche Shale, Nevada, 11, 183 (169).

From Purley Shales, Warwickshire, 6, 397 (403).

Irvingella nuneatonensis, 10, 339 (377).

Ontogeny of Peltura scarabaeoides , 1, 200 (483).

Revision of two Up. Cambrian trilobites, 11, 410 (378).

Tremadocian

New trilobites from Shropshire, 10, 47 (224).

Pseudokaiiiella impar , 12, 406 (484).

Ordovician

Ashgillian from Ireland, 1, 369 (478).

Duftonia from England and Wales, 2, 143 (727).

Encrinurus multisegmentatus and allied species, 1, 60 (450).

From Albany division, Girvan, 8, 577 (452).

From northern Yukon, 9, 39 (264).

Llandeilo from Berwyn Hills, N. Wales, 5 , 790 ( 273).

Oedicybele from Kildare Limestone, 8, 1 (443).

Silicified from South Wales, 6, 254 (414).

Tiresias M'Coy, 5, 340 (130).

Silurian

Acaste downingiae and related spp., 9, 183 (395).

A caste! la spinosa, revision of, 10, 175 (396).

Dalmanites myops (Konig), 2, 280 (128).

Denckmannites from New South Wales, 11, 691 (397).

14

GENERAL INDEX

Encrinums punctatus and related spp., 5, 460 (451).

From Upper Llandovery of Tortworth, 1, 139 (725).

Odontopleurids from Balto-scandia, 10, 214 (54).

Schizochroal eyes and vision, 9, 1 (91), 464 (92); 10, 603 (93).

Two new genera from Howgill Fells, 7, 541 (350).

Devonian

Colour markings in phacopids from New York, 11, 498 (161).

New odontopleurid from Bohemia, 9, 330 (55).

Carboniferous

Australosutura: new genus, 3, 227 (10); from Oklahoma, 9, 270 (309).
Tournaisian of Britain and Belgium, 1, 231 (176).

Permian

From Salt Range, West Pakistan, 9, 64 (179).

VERTEBRATA, see Agnatha, Amphibia, Mammalia, Pisces, Reptilia.

NOTES ON THE ST R AT IG R A PH I C A L INDEX

The entries are arranged in sequence by Periods, starting with the oldest. There are
some entries under Eras, usually where the subject is also broad and where there is no
restricted indication of ‘age’ in the title of the paper. The Tremadocian has been indexed
separately between the Cambrian and Ordovician and the Old Red Sandstone in-
cluded in the Devonian. Within each period, the initial subdivision is generally by class
or phylum, arranged alphabetically, and it is hoped that none of the resulting groupings
is too large for convenience. Papers covering specifically more than one period have
been entered under each period.

Entries include reference to the volume and page of Palaeontology on which the
paper referred to begins as well as the index number (in italics) from the Author List.

STRATIGRAPHICAL INDEX

PRECAMBRIAN

Late Precambrian fossils from Ediacara, S. Australia, 9 , 599 (174).

Medusae from uppermost Precambrian or Cambrian, central Australia, 12, 351 (459).

Calcareous algae from the Dalradian of Islay, 5, 268 (182).

PALAEOZOIC, general

Brazilian Palaeozoic algomycetes and Tasmanacea, 10, 640 (407).

Catalogue of the Begg Collection, Glasgow, 6, 378 (124).

Corals from New South Wales, 4, 334 (425).

Echinoderm faunas of the British Isles and Balto-scandia, 2, 161 (344).

Graphical aids for description in fusulines, 10, 322 (126).

Microstructure of stromatoporoids, 9, 74 (417).

Morphology and function of dichoporite pore-structures, 11, 697 (321).

Origin of the loop in articulate brachiopods, 4, 149 (486).

Sequence of graptolite faunas, 1, 159 (55).

CAMBRIAN

Ceratocystis — a chordate with echinoderm affinities, 12, 494 (231).

Shell structure of billingsellacean brachiopods, 11, 486 (487).

Trilobites: alimentary caeca of agnostide, 3, 410 (308); Estaingia, new genus from S. Australia, 7,
458 (335); from Pioche Shale, Nevada, 11, 183 (169); from Purley Shales, Warwickshire, 6, 397
(403); from Turkey, 4, 71 (129); Irvingella nuneatonensis , 10, 339 (377); ontogeny of Peltura, 1,
200 (483); revision of two Upper Cambrian, 11, 410 (378).

TREMADOCIAN

Graptolites from Norway, 6, 121 (413).

Macrocystella, the earliest glyptocystid cystoid, 11, 580 (320).

New trilobites from Shropshire, 10, 47 (224).

Pseudokainella impar (Trilobita), 12, 406 (484).

ORDOVICIAN

Alga: type species of Girvanella, 1, 22 (496).

Bivalvia: systematics, affinities and life habits of Babinka, 8, 231 (275); new species of Babinka
from Sweden, 12, 173 (408).

Bothriocidaridae, new from Girvan, 10, 525 (319).

Brachiopoda: Eodinobolus from Ontario, 12, 161 (305); four British dalmanelloids, 1, 292 (76);

Llandeilo from Wales, 4, 177 (272); morphology of Triplesiacea, 5, 740 (499).

Bryozoa: Caradocian from Shropshire, 5, 52 (365), 6, 1 (367), 8, 5 (369); Chazyan, 5, 727 (366);

types of seven bifoliate, 3, 1 (330).

Chitinozoa from Shropshire, 10, 436 (236).

Crinoidea: ‘ Dendrocrinus ’ cambriense, the earliest crinoid, 11, 406 (33); new from North Wales, 8,
355 (32).

Crustacea: morphology and range of Caryocaris from Alaska and Great Basin, 9, 371 (87).
Edrioblastoid, new from eastern Australia, 11, 513 (476).

Graptolites: development of a dicellograptid, 8, 41 (227); Glyptograptus dentatus and allied species,
6, 665 (56); Lasiograptus harknessi, 8, 272 (352); new variety of Orthoretoilites hand, 2, 226 (399).
Silicified fossils from South Wales, 6, 254 (414).

Stromatoporoids from New South Wales, 12, 637 (477).

STRATIGRAPHICAL INDEX

17

Trilobita: Ashgillian from Ireland, 1, 369 (478); Duftonia, 2, 143 (127); Encrinurus multisegmentatus
and allied species, 1, 60 (450); from Girvan, 8, 577 (452); from Yukon, 9, 39 (264); Llandeilo
from Wales, 5, 790 (273); Oedicybele from Ireland, 8, 1 (443); Tiresias, 5, 340 (130).

SILURIAN

Alga: Girvanella from Welsh Borderland, 9, 48 (239).

Brachiopoda: brachial structure in Pentameridae, 7, 220 (12); Cortezorthinae, new dalmanellid
group, 10, 142 (242); Dayia navicula, 11, 612 (453); Dinobolus from British Columbia, 3, 242
(304); Eocoelia hemisphaerica, 9, 523 (507); Eospiriferidae, 5, 682 (45); Isorthis and Salopina
from Welsh Borders, 8, 454 (465) ; Leptaena from England, 10, 590 (253); Leptocoeliidae, revision
of, 6, 440 (11); Llandovery stropheodontids from Welsh Borders, 10, 245 (96); unusual strick-
landiids from Wales, 9, 346 (506); Visbyella, new resserellid, 11, 306 (466).

Bryozoa: from Canadian Arctic, 9, 517 (43); from Central Wales, 3, 69 (310); Ludlovian of Ludlow,
5, 195 (311); Wenlockian fenestrate, 5, 540 (290); Wenlock palaeogeography, 12, 621 (312).

Cephalopoda: Leurocycloceras from Wales and Welsh Borders, 7, 525 (201).

Corals: dimorphism in Striatopora, 9, 448 (307); Cystihalysites from England, 7, 452 (433); Micro-
plasma lovenianum from Wales, 9, 148 (481); sequence of faunas in U.S.S.R., 10, 660 (195).

Crinoids from British Ludlovian, 1, 106 (341).

Crustacea: cuticle of ceratiocaridids, 5, 30 (359); ostracods from Nova Scotia, 3, 93 (107); phyllo-
carids from Czechoslovakia, 6, 97 (86); pedunculate cirripede from Esthonia, 6, 161 (490).

Echinoidea: age of Myriastiches gigas, 10, 171 (266); pedicellariae, 11, 576 (42).

Eurypterid: median appendage of Slimonia acuminata, 3, 245 (469).

Graptolites: assemblages and zones of Birkhill Shales, 11, 654 (448); diplograptids, 5, 498 (313),
11, 1 (57); new from N. England, 8, 247 (351); monograptids from Illinois, 5, 59 (362); Wenlock
from Ludlow district, 12, 663 (202).

Llandovery transgression of the Welsh Borderland, 11, 736 (508).

Medusoid (?) from England, SI, 610 (424).

Microplankton: age of primitive echinoid, 10, 171 (266); from Wenlock Shales, 2, 56 (142), 6, 625
(144).

Ostracoderms : Corvaspis kingi, 3, 217 (437); Jamoytius kerwoodi, 11, 21 (353).

Spores: assemblages from Welsh Borders, 12, 201 (349).

Trilobita: Acaste downingiae, revision 9, 183 (395); Acastella spinosa, revision, 10, 175 (396);
Dalmanites myops (Konig), 2, 280 (128); Denckmannites from New South Wales, 11, 691 (397);
Encrinurus punctatus and allied species, 5 , 460 (451); fine structure of eye of Phacops, 10, 603
(93); Llandovery from Tortworth, 1, 139 (125); odontopleurids from Balto-scandia, 10, 214 (54);
schizochroal eyes and vision, 9, 1 (91), 464 (92); two new phacopid genera, 7, 541 (350).

DOWNTONIAN, see Silurian.

OLD RED SANDSTONE, see Devonian

DEVONIAN

Bivalvia: Prothyris in Great Britain, 6, 136 (495).

Blastoid: Belocrinus from France, 9, 244 (274).

Brachiopoda: Atrypidae, unusual structures in, 8, 358 (108); Australirhynchia, new genus, 11, 731
(386); brachial structure in Pentameridae, 7, 220 (12); Cortezorthinae, new dalmanellid group,
10, 142 (242); delthyrial cover in Mucrospirifer, 11, 317 (114); Eospiriferidae, 5, 682 (45); from
Brendon Hills, Somerset, 7, 1 (474); from central Nevada, 9, 152 (240); from Western Canada,
3, 208 (322); interpretation of serial sections, 12, 321 (480); Leptocoeliidae, revision of, 6, 440
(11); new spiriferids from New South Wales, 12, 472 (387); Parachonetes, new genus, 9, 365 (241);
P/anicardinia, new dalmanellid, 11, 627 (385); schizophoriids from Europe, 9, 381 (336); Spina-
trypa and Spinatrypina, 10, 489 (109).

Cephalopoda: abnormal growths in goniatiites, 3, 129 (205); Acanthoclymenia, the earliest clymeniid,
3, 472 (206); ammonoids from Cornwall, 3, 153 (393); Cheiloceras from New South Wales, 9, 458
(234); clymeniid from New South Wales, 3, 237 (332); Famennian ammonoids from New South
Wales, 11, 535 (235); goniatites and stratigraphical correlations in Western Canada, 6, 491 (207).

Corals: Alaiophylliim from Canada, 6, 132 (323); correlation by tetracorals in Canada, 7, 430 (324);

18

STRATIGRAPHICAL INDEX

from Brendon Hills, Somerset, 7, 1 (474); from Garra Formation, New South Wales, 8, 518 (426),

9, 544 (427); from New South Wales, 10, 426 (428); Hexagonaria from France, 12, 178 (409);
Marisastridae from Devon, 10, 266 (390); Metriophyllum , 7, 108 (203); periodicity in coral
growth, 7, 552 (389); sequence of faunas in the U.S.S.R., 10, 660 (195); Smithiphyllum in North
America, 8, 618 (325); squamulate favositids of Victoria, 3, 186 (327); tabulates of North Devon,
11, 44 (204).

Crinoids: inadunate from Somerset, 4, 538 (473); Quantoxocrinus from Somerset, 8, 11 (475).
Crustacea: phyllocarids from Czechoslovakia, 6, 97 (86).

Eurypterid, new from Old Red Sandstone of Scotland, 5, 137 (470).

Foraminifera from Western Australia, 3, 397 (121); 11, 601 (99).

Gastropod: Orecopia in western Canada, 9, 142 (326).

Graptolites: Monograptus from Yukon, 6, 751 (225); with land plants in Alaska, 12, 559 (88).
Microplankton, reef-controlled distribution in Alberta, 4, 392 (416).

Mollusca, monoplacophora: a sinus-bearing monoplacophoran and the classification of primitive
molluscs, 11, 132 (361).

Palaeoecology : specific frequency and enviromental indications in the Calcaire de Ferques, 12, 366
(464).

Plants, pteridophytes : land plants from graptolite shale, Alaska, 12, 559 (88); new plant from South
Wales, 11, 683 (151).

Spores: assemblages from South Wales, 12, 201 (349); assemblages from Vestspitsbergen, 10, 280
(6); from Canadian Arctic, 3, 26 (276); from Vestspitsbergen, 8, 687 (5); megaspores from Cana-
dian Arctic, 1, 321 (78); megaspores from Wyboston bore, Beds., 10, 189 (294), 706 (295); middle

0. R.S. of Cromarty, 3, 45 (346); Middle O.R.S. of Orcadian basin, 7, 559 (348); seed megaspore
from Canada, 7, 29 (82); with bifurcate processes, from Scotland, 5, 171 (347).

Trilobita: colour markings in Phocops and Greenops, 11, 498 (161); fine structure of eye of Phacops,

10, 603 (93); from Cornwall, 3, 153 (393); new odontopleurid from Bohemia, 9, 330 (53).
Vertebrates: additions to knowledge of antiarchs, 4, 210 (471); Cephalaspis lyelli Ag., 1, 99 (482);

skull roof and palate of Dipnorhynchus, 8, 634 (65); Traquairaspis from Canada, 7, 210 (139).

CARBONIFEROUS

Algae: biostromes in Namurian of northern England, 1, 147 (238); British species of Girvanella, 6,
264 (497); Calcifolium from Visean of Scotland, 8, 192 (56’); Komia and its systematic position,
6, 246 (492); Nanopora, new dasycladacean, 7, 181 (498).

Arthropods: non-marine ostracod fauna from northern England, 9, 667 (338); Rochdalia, an insect
nymph, SO, 307 (360); external anatomy of some ‘scorpions’, 1, 261 (488), 3, 276 (489); ventral
anatomy of eurypterid, 7, 474 (491).

Bivalvia: distinctions between Caneyella, Posidonia, and Posidoniella, 1, 405 (342); evidence for
‘Nebraskan’ fauna in Scotland, 4, 507 (494); Naiadites obesus from Fife, 4, 300 (36); non-marine
assemblages from Pembrokeshire, 3, 104 (233); non-marine, from East Fife, 3, 137 (37); non-
marine, new, 5, 307 (147); Prothyris in Great Britain, 6, 136 (495); Wilkingia to replace Allorisma,

1, 401 (493).

Brachiopoda: Delepinea from New South Wales, 7, 514 (66); Juresania nebrascensis, taxonomy, 7,
23 (162); mantle canal patterns in Schizophoria, 11, 389 (355); new productid from Scotland, 9,
426 (394); pedicle sheath in young productaceans, 7, 703 (51); predation and shell damage in
a Visean fauna, 9, 355 (52); Pugnoides triplex, 3, 477 (318); schizophoriids from Europe, 11, 64
(337); shell development in Spirifer trigonalis , 4, 477 (146); type species of three punctate spiri-
feroids, 1, 351 (63).

Bryozoa: new fenestrate from Ireland, 8, 478 (438); Polypora M’Coy, 6, 166 (291); skeletal structure
and growth in Fenestellidae, 12, 281 (440); type specimens of Fenestella, 4, 221 (289).
Cephalopoda: Delepinoeeras in North America, 7, 173 (171); Entogonites cf. borealis from Ireland,
1, 158 (196); Eumorphoceras, new, 4, 54 (501); goniatite fauna from Visean/Namurian boundary,

11, 264 (165); Goniatites striatus and related forms, 1, 384 (197); goniatites from Staffordshire, 1,
16 (39); Nuculoceras stellarum , 8, 226 (199); palaeoecology of Goniatite Bed, Cowlow Nick, 8,
186 (168); palaeontology of Namurian rocks, Slieve Anierin, Eire, 5, 355 (502); review of goniatite
zones in Devon and Cornwall, 3, 75 (60); Visean fauna from New South Wales, 7, 682 (50).

STRATIGRAPHICAL INDEX 19

Conodonts: Apatognathus from Yoredale Series, 10, 124 (457); assemblage from Avon Gorge, 12,
400 (18); from East Cornwall, 12, 262 (281); from South Devon, 12, 276 (282).

Corals: increase in Lithostrotion, 8, 204 (245); hystero-ontogeny in Lonsdaleia and Thysanophyllum,
10, 617 (246); Slimoniphyllum , new genus from Britain, 4, 280 (247).

Crustacea: Cyclus martinemsis from Mendip Hills, 10, 317 (177); syncarid from Stephanian of
Warwickshire, 4, 546 (358).

Echinoderms: starfish traces from Namurian of Ireland, 7, 508 (265); structure and systematic
position of Pentablastus from Spain, 6, 471 (244).

Faunas: from Kuttung rocks of New South Wales, 4, 428 (64); from Trevallyn, New South Wales,
8, 54 (356); population studies in Visean of NW. Ireland, 9, 252 (210).

Foraminifera: fusulinidae from Spitsbergen, 2, 210 (167).

Gastropoda: faunas from Queensland, 4, 59 (283).

Gymnosperm: Amyelon in American coal balls, 7, 186 (122); revision of Amyelon, 5, 213 (24).

Lycopsida: compressed sporophyll from Somerset, 11, 445 (46); fertile, from Scottish Lower Car-
boniferous, 8, 281 (8); revision of Eskdalia, 11, 439 (445); Selaginellites with Densosporites, 1,
245 (77); Sporangiostrobus with Densosporites, 5, 73 (81).

Megaspores: cristate from Scottish Lower Carboniferous, 9, 488 (9); from Visean of Scotland, 12,
441 (412); Westphalian D, Forest of Dean, 8, 82 (411).

Micro-organisms: composition of Pyritosphaera barbaric t, 6, 119 (271); micro-organisms and
syngenetic pyrite, 5, 444 (270).

Miospores: assemblages from coals and associated sediments, Yorkshire, 7, 656 (279); distribution
of assemblages in Lr. Kittanning Coal Measures, Pa., 9, 629 (181); from Basement Beds, Menai
Straits, N. Wales, 12, 420 (194); from Drybrook Sandstone, Forest of Dean, 7, 351 (431); from
Lr. Carboniferous, NW. England, 10, 1 (61); from Lr. Carboniferous, Spitsbergen, 5, 550, 619
(334); microbiological attack on miospores, 6, 349 (293); Namurian, from southern Pennines,

4, 247 (302); sections of dispersed spores, 5, 247 (220), 679 (133); Simozonotriletes , 1, 125 (429);
Springer Formation, Oklahoma, 10, 349 (164); structure of wall of Cingulati, 3, 82 (401); Tetra-
pterites visensis, new spore-bearing structure, 7, 64 (430); Tournaisian flora from Ayrshire, 11,
116 (432).

Ostracoda: non-marine from English Coal Measures, 9, 667 (338).

Protozoa, radiolaria, from Namurian of Derbyshire, 9, 319 (200).

Pteridosperms : Aulacotheca from Iowa, 12, 414 (152); Calathospermum fimbriatum, cupule from
Scottish Lr. Carboniferous, 3, 265 (23); Mariopteris from Spanish Stephanian, 10, 694 (460);
probable pteridosperm microsporangiate fructification from Illinois, 7, 60 (132); structure and
relationships of Pachytesta, 12, 382 (442); three fructifications from Scottish Lr. Carboniferous,

5, 225 (402).

Pteropsida: Biscalitheca from Illinois, 11, 104 (331); structure and relationships of Radstockia, 10,
43 (441).

Sphenopsida : calamitean plants from Scottish Lr. Carboniferous, 6, 408 (84) ; new British calamite
cone, 12, 253 (446); new cone from Illinois, 8, 681 (193); on Potbocites Paterson, 8, 107 (85).

Trilobita: Australosutura, new genus from Australia and Argentina, 3, 227 (10); Australosutura
from U.S.A., 9, 270 (309); Tournaisian trilobites from Britain and Belgium, 1, 231 (176).

Vertebrates: haplolepid fish fauna from Nova Scotia, 5, 22 (19); Namaichthys and other fishes from
South Africa, 5, 9 (173); new family of amphibians, 12, 537 (67).

PERMIAN

Algae: new Tethyan Dasycladaceae, 11, 491 (160).

Bivalvia: Atomodesma from New Zealand, 6, 699 (467); Eurydesma from Dwyka Beds, S. Africa,
4, 138 (134); new records from eastern Australia, 4, 119 (137); Merismopteria and origin of
Pteriidae, 3, 387 (135); palaeotaxodonts from New Zealand, 7, 630 (468); Permophorus costatus,
dentition, 7, 281 (269); preserved ligaments from Australia, 11, 94 (376).

Brachipoda: Attenuateila, unusual brachial skeleton, 11, 783 (16); feeding mechanism of Prorich-
thofenia, 3, 450 (371); Horridonia, 4, 42 (175); Howseia, new genus from Durham, 6, 754 (268);
Martiniopsis-Uke spiriferids from Queensland, 1, 333 (62); shell structure in Terrakea and Strep-
torhynchus, 12, 310 (17); type species of three punctate spiriferoids, 1, 351 (63).

20

STRATIGRAPHICAL INDEX

Bryozoa, from Western Australia, 6, 70 (368).

Corals, from northern Iraq, 1, 174 (215).

Foraminifera: from British Honduras, 5, 297 (363); fusulinids from Peru, 5, 817 (364); fusulinids
from Spitsbergen, 2, 210 (167).

Gastropoda: Peruvispira from Dwyka Beds, S. Africa, 4, 138 (134); Platyteichum, from W. Australia,
4, 131 (138).

Microplankton: hystricospheres from Britain, 5, 770 (462).

Miospores: assemblage from Iraq, 7 , 240 (398); British saccate and monosulcate, 8, 322 (90);
new British spore, 4, 648 (80).

Plants, land: Peltaspermaceae, pteridosperm family, 3, 333 (449); structure of leaves of Rhabdo-
taenia from India, 6, 301 (316); structure of Vertebraria indica, 11, 643 (317).

Trilobita: from Salt Range, W. Pakistan, 9, 64 (179).

Vertebrates: growth stages in branchiosaurs, 6, 540 (472).

MESOZOIC, general

Adherent foraminifera, 1, 116 (25).

Calcareous adherent foraminifera from British Jurassic and Cretaceous, 5, 149 (2).

Fossil cycads, 4, 313 (797).

Reappraisal of microspore Aequitriradites, 4, 425 (102).

Yorkshire type ammonites and nautiloids, 5, 93 (208).

TRIASSIC

Algal growths in Rhaetic Cotham Marble, England, 4, 324 (190).

Arthropods: new Rhaetic and Liassic Beetles, 4, 87 (172).

Bivalvia: Rhaetic-Hettangian Pteromya, 6, 582 (118); from Oman Peninsula, Arabia, 4, 1 (218);
new genera and subgenera, 4, 592 (116); photonegative young in Lima , 3, 362 (228).

Brachiopoda: feeding mechanisms in Thecopira and Bactrynium, 11, 329 (375); from Oman Penin-
sula, Arabia, 4, 1 (218).

Microplankton: fine structure of some acritarchs, 9, 351 (285).

Plants, land: flora from Cacheuta Formation, Argentina, 10, 564 (226); leaf Dicroidium and relation
to Rhexoxylon, 11, 500 (14); pteridosperm family Peltaspermaceae, 3, 333 (449); trunk of Rhexo-
xylon, 11, 236 (49).

Spores, etc.: Keuper miospores from England, 8, 294 (89); plant microfossils from W. Australia,
6, 12 (27).

Vertebrates: Birgeria acuminata and absence of labyrinthodonts from Rhaetic, 9, 135 (388); his-
tology of dinosaur bone, 5, 238 (123); new evidence for a southern transatlantic connection, 10,
554 (44); Saurichthys krambergeri , 5, 344 (108).

JURASSIC

Bivalvia: epizoic oysters, 11, 19 (106); intestine of Nuculana from Lias, 2, 262 (115); Malayomaorica,
new genus from Indo-Pacific, 6, 148 (232); micro-structure of mytilid shell, 11, 163 (213); mode of
life of ‘ Posidonia ’, 8, 156 (230); new genera and subgenera, 4, 592 (116); Rhaetic-Hettangian
Pteromya, 6, 582 (118).

Brachiopoda: life assemblages from Marlstone Rock-bed, 4, 653 (184); ontogeny of Moorelline
granulosa, 12, 388 (20); sensory spines in Acanthothiris, 8, 604 (374); the true Rhynchonella, 1,

1 (4).

Cephalopoda: belemnite, new, from Indonesia, 7, 621 (422); Belemnites gerardi, 6, 690 (420);
Dicoelites and Prodicoelites, 7, 606 (421); epizoic oysters on ammonites, 11, 19 (106); Jura-
phyllitidae in Britain, 7, 286 (209); Kimmeridgian ammonites from Lincolnshire, 6, 219 (15);
Neomicroceras and Paracymbites, 9,312 (141) ; Propectinatites, new genus, 11,16 (105) ; Pseudolillia,
5, 86 (140); Rasenia from Scotland, 5, 765 (509); variation and ontogeny of some Oxfordian, 9,
290 (314), 10, 60 (375).

Crinoids: form and function of stem, 11, 275 (392); Phyllocrinus from New Zealand, 2, 150 (410).

Echinoids: Cidarites moniliferus and status of Eucidaris, 5, 785 (328); dentition and relationships of
Py gas ter, 4, 243 (286).

STRATIGRAPHICAL INDEX

21

Fauna: ecology and distribution of invertebrates of Great Estuarine Series, 6, 327 (212); faunal
realms and facies in the Jurassic, 12, 1 (188); recognition of salinity-controlled mollusc assem-
blages, 6, 318 (211).

Foraminifera : arenaceous from type Kimeridgian, 1, 298 (267); Brotzenia and Voorthuysenia, 6,
653 (110); from Ampthill Clay, Cambridgeshire, 4, 520 (178); new information on Pfenderina, 4,
581 (405).

Gastropoda: Discohelix as an index fossil, 11, 554 (479).

Gymnosperm: opposite-leaved conifer from Israel, 2, 236 (79).

Insecta: beetles from Antarctica, 1, 407 (505); Liassic dragonfly, !, 406 (504); new Rhaetic and
Liassic beetles, 4, 87 (172).

Microfossils: Up. Jurassic from Hautes-Alpes, 8, 391 (454).

Microplankton: from Ampthill Clay, south Yorkshire, 5, 478 (382); from Australia and New
Guinea, 2, 243 (100); from Kellaways Rock and Oxford Clay, Yorkshire, 4, 90 (381); new name
for species of Gonyaulacysta, 7, 472 (383).

Ostracoda: freshwater from Bathonian of Oxfordshire, 8, 749 (31); from Dorset Kimmeridge Clay,
12, 1 12 (260); Mandelstamia from England, 3, 439 (299); Oertliana, new genus from NW. Europe,
8, 572 (258).

Palaeoecology : in the Great Oolite, Kirtlington, Oxfordshire, 12, 56 (277); of the Frodingham Iron-
stone (Lr. Jurassic), 6, 554 (755); salinity-controlled mollusc assemblages, 6, 318 (211).

Spores and pollen: further interpretation of Eucommiidites , 4, 292 (219); miospores from Purbeck
and marine Up. Jurassic, S. England, 12, 574 (306).

Stromatoporoids: Actostroma, new genus from Israel, 1, 87 (274); revision of Actinostromina,
Astrostylopsis and Trupetostromaria, 2, 28 (216); Tethyan Stromatoporina, Dehornella and Astro-
porina, 2, 180 (217).

Trace fossils: Favreina-Thalassinoides association from Gt. Oolite, 12, 549 (256); Kulindrichnus
from Lias, 3, 64 (183).

Vertebrates: dinosaur Scelidosaurus, 11, 40 (303); ‘dwarf’ crocodiles of Purbeck, 10, 629 (237); fish
otoliths from English Bathonian, 11, 246 (423); gastric contents of ichthyosaur, 11, 376 (339);
giant Kimmeridgian pliosaur, 2, 39 (436); late Jurassic mammals, 6, 373 (95); Pliosaurus brachy-
spondylus, 1, 283 (435); scapula of Pliosaurus macromerus, 1, 193 (434).

CRETACEOUS

Algae: alga debris-facies in the Middle East, 1, 254 (154); coccoliths from Atlantic seamounts,

7, 306 (40); coccolithophorids from Zululand, 11, 361 (333); interrelationships of some Codiaceae,

8, 199 (159); sexual organization of Permocalcuhis, 4, 82 (155); Tethyan Dasycladaceae, 11, 491
(160).

Bivalvia: characters and relationships of Pseudavicula, 3, 392 (136); functional studies on Arcto-
strea, 11, 458 (69).

Brachiopoda: English Aptian terebratulids, 2, 94 (287).

Bryozoa: Dionella, new genus from Europe, 8, 492 (284); Pyripora and Rhammatopora, 3, 370 (447).

Cephalopoda: ammonites from Barremian of Bulgaria, 5, 527 (278); ammonites from Bathurst
Island, Australia, 6, 597 (500); ammonoids from Berriasian of the Speeton Clay, 5, 272 (301);
biometric study of Barremites, 6, 727 (345); Hengestites, new genus from Gault, 2, 200 (71);
Leymeriella in Britain, 1, 29 (70); new heteromorph from Yorkshire, 6, 575 (145); origin, limits,
and systematic position of Scaphites, 8, 397 (485); phylloceratid from the Speeton Clay of York-
shire, 9, 455 (343).

Cirripede: Arcoscalpellum comptum, new to the Gault, 8, 629 (98).

Echinoderms: echinoid with false teeth, 12, 488 (257); new echinoid from Albian of Kent, 3, 260
(73); new ophiuroid from Australia, 6, 579 (400); Salenia in the eastern Pacific, 7, 331 (510);
spines and fascioles of Echinocorys scutata, 6, 458 (418).

Foraminifera: Bolivinoides from the British Isles, 9, 220 (29); Chubbina, new alveolinid from Jamaica
and Mexico, 11, 526 (357); from Ballydeenlea Chalk, Ireland, 9, 492 (30); Maestrichtian from
Galicia Bank, 12, 19 (170), 189 (166); morphology and development of Marssonella and Pseudo-
textulariella, 6, 41 (27); palaeoecology of Chalk Marl, 4, 599 (59); planktonic, from Isle of Wight,
4, 552 (28); Polymorphinidae from England, 5, 712 (26).

22

STRATIGRAPHICAL INDEX

Gastropoda: fossilized intestines of, 2, 270 (72); nomenclatural corrections, 4, 312 (75).

Microfossils: from the Hautes-Alpes, 8, 391 (454) ; problematical, from Middle East, 6, 293 (156).

Microplankton, from the Cambridge Greensand, 7, 37 (104).

Ostracoda: first non-marine from Ghana, 11, 259 (262); from Aptian, Lincolnshire, 8, 375 (251);
from Bargate Beds, Surrey, 7, 317 (250); from Barremian, Lincolnshire, 9, 208 (252); from Cre-
taceous of California, 7, 393 (198); microfauna of marine brackish bands in Weald Clay, 11,
141 (259); Neocy there in Speeton Clay, 6, 274 (248); ontogeny of Theriosynoecum fittoni, 7, 72
(406); Orthonotacythere inversa from the Speeton Clay, 6, 430 (249).

Palaeoecology : Actinocamax plenus Subzone in Anglo-Paris Basin, 4, 609 (229); brachiopod ecology
and Lr. Greensand palaeogeography, 5, 253 (288); crustacean burrows in the Weald Clay, 12, 459
(255); transition zone across an Upper Cretaceous boundary in New Jersey, 7, 266 (261).

Spores and pollen: angiosperm pollen from British Barremian to Albian strata, 11, 421 (254);
Aquilapollenites in the British Isles, 11, 549 (280); dispersed spores of Equisetites, 11, 633 (34);
Hoegisporis, new Australian form genus, 3, 485 (101); new species of Hoegisporis, 8, 39 (103);
revision of some Belgian microspores, 6, 282 (131); schizaeaceous spores, from macrofossils, 9,
274 (221); stratigraphic correlation using miospores, 12, 84 (222); Wealden megaspores and facies
distribution, 12, 333 (25).

Stratigraphical palaeontology of the Lower Greensand, 3, 487 (74).

Trace fossils: probable cirripede, phoronid, and echiuroid burrows in an echinoid test, 1, 397 (243).

Vertebrates: Wealden mammalian fossils, 6, 55 (94).

TERTIARY, general

Australasian Typhinae (Gastropoda), 4, 362 (458).

Bivalvia from Libya, 5, 1 (117).

Classification of cassiduloid echinoids, 6, 718 (329).

Classification and distribution of the Globigerinaceae, 2, 1 (22).

Coccoliths from Atlantic seamounts, 7, 306 (40).

Cupuladria canariensis — portrait of a bryozoan, 6, 172 (263).

Development of Globigerinoides ruber, Miocene to Recent, 10, 647 (111).

Echinoid Salenia in the eastern Pacific, 7, 331 (510).

Growth gradients among fossil monotremes and marsupials, 6, 615 (419).

Homoptera of Stavropol and reconstruction of continental palaeobiocoenoses, 10, 542 (36).

Larger foraminifera from Central America, 11, 283 (149).

Neogene Tasmanites and leiospheres from Louisiana, 8, 16 (163).

Solenoporacean algae and their reproductive structures, 7, 695 (158).

PALAEOCENE

Heterostegina, 10, 314 (148).

Problematical microfossils from the Middle East, 6, 293 (156).

Sindulites, a new nummulitid, 11, 435 (150).

Teredinid (Mollusca) from Iraq, 6, 315 (157).

EOCENE

Coelenterate : sea-pen from New Zealand, 1, 226 (189).

Crustacea: crabs in London Clay nodule, 4, 85 (97).

Foraminifera: adherent from France, 5, 149 (2); Cassigerinella from Florida, 11, 368 (112); Disco-
cyclina from India, 6, 658 (380); morphology and phylogeny of Orbulinoides beckmanni, 11, 371
(113); note on Operculinoides, 2, 156 (297); Nummulites from India, 11, 669 (379).

Gastropoda: apical development in turritellid classification, 8, 666 (7).

Vertebrates: European Proviverrini (Mammalia), 8, 638 (456); new pleurodiran turtle from Somalia,
9, 511 (461).

Wood: Anacardiaceae from Britain, 9, 360 (48); oak wood from Britain, 3, 86 (47).

OLIGOCENE

Stratigraphical distribution of Archais (Foraminifera), 1, 207 (404).

Subterraniphylliim , new calcareous alga, 1, 73 (153).

STRATIGRAPHICAL INDEX

23

MIOCENE

Anthropoids from India, 7, 124 (340).

Coelopleurus (Echinoidea) from Malta, 12, 42 (503).

Geological distribution of Discospirina (Foraminifera), 1, 364 (1).

PLIOCENE

Foraminifera: distribution of Alliatina and Alliatinella, 1, 76 (68).

PLEISTOCENE

Microplankton in deep-sea cores from Caribbean, 10, 95 (463).

New turtle from Siwaliks of India, 12 , 555 (444).

Normanicythere and the ostracod family Trachyleberididae, 2, 72 (298).

Odontoma in a northern mammoth, 7, 674 (223).

RECENT

Brachiopoda: anchorage of articulates on soft substrata, 4, 475 (372); shell growth in terebratuloids,
10, 298 (384); Valdiviathyris, 4, 542 (370).

Foraminifera in Elolocene marsh cycles at Borth, Wales, 8, 27 (3).

Normanicythere leioderma in North America, 4, 424 (300).

NOTES ON THE AUTHOR LIST

This list provides an index to the papers by author but it is mainly useful as a
supplement to the other indexes for those who do not have a complete set of Palaeonto-
logy at hand. Each paper has been given a reference number which is quoted with the
entry or entries for the paper in the general and stratigraphic indexes.

In almost all cases, papers by the same author are listed in sequence of publication,
those with multiple authors being inserted in sequence under the first author. Cross-
references are provided for the second (and subsequent) authors.

AUTHOR LIST

1. adams, c. g. 1959. Geological distribution of Discospirina (Foraminifera) and occurrence of
D. italica in the Miocene of Cyprus. I, 364.

2. adams, c. g. 1962. Calcareous adherent foraminifera from the British Jurassic and Cretaceous and
the French Eocene. 5, 149.

3. adams, T. d., and haynes, J. 1965. Foraminifera in Holocene marsh cycles at Borth, Cardiganshire
(Wales). 8,27.

4. ager, d. v. 1957. The true Rhynchonella. 1,1.

5. allen, k. c. 1965. Lower and Middle Devonian spores of North and Central Vestspitsbergen.
8, 687.

6. allen, k. c. 1967. Spore assemblages and their stratigraphic application in the Lower and Middle
Devonian of North and Central Vestspitsbergen. 10, 280.

ALLEN, N. W., see KILENYI, T. I.

ALLISON, E. C., see ZULLO, V. A.

7. allison, r. c. 1965. Apical development in turritellid classification with a description of Cris-
tispira pugetensis gen. et. sp. nov. 8, 666.

ALVAREZ-RAMIS, C., See WAGNER, R. H.

8. alvin, k. l. 1965. A new fertile lycopod from the Lower Carboniferous of Scotland. 8, 281.

9. alvin, k. l. 1966. Two cristate megaspores from the Lower Carboniferous of Scotland. 9, 488.

10. amos, a. J., Campbell, k. s. w., and goldring, r. 1960. Australosutura gen. nov. (Trilobita) from
the Carboniferous of Australia and Argentina. 3, 227 .

11. amos, a., and boucot, a. j. 1963. A revision of the brachipod family Leptocoeliidae. 6, 440.

12. amsden, t. w. 1964. Brachial plate structure in the brachiopod family Pentameridae. 7, 220.

13. anderson, F. w. 1964. The law of ostracod growth, 7, 85. See also sohn, i. g.

ANDREWS, H. N., See PHILLIPS, T. L.

14. archangelsky, s. 1968. Studies on Triassic fossil plants from Argentina. IV. The leaf genus
j Oicroidium and its possible relation to Rhexoxylon stems. 11, 500.

15. arkell, w. j., and callomon, j. h. 1963. Lower Kimeridgian ammonites from the drift of Lincoln-
shire. 6, 219.

16. Armstrong, J. 1968. The unusual brachial skeleton of Attemiatella convexa sp. nov. ( Brachiopoda).
11, 783.

17. Armstrong, J. 1969. The cross-bladed fabrics of the shells of Terrakea solida (Etheridge and Dun)
and Streptorhynchus pelicanensis Fletcher. 12, 310.

18. Austin, r. l., and Rhodes, f. h. t. 1969. A conodont assemblage from the Carboniferous of the
Avon Gorge, Bristol. 12, 400.

badam, g. l., see tewari, b. s.

19. baird, d. a. 1962. A haplolepid fish fauna in the early Pennsylvanian of Nova Scotia. 5, 22.

20. baker, p. g. 1969. The ontogeny of the thecideacean brachiopod Moorellina granulosa (Moore)
from the Middle Jurassic of England. 12, 388.

21. balme, b. e. 1963. Plant microfossils from the Lower Triassic of Western Australia. 6, 12.

22. banner, f. t., and blow, w. h. 1959. The classification and stratigraphical distribution of the
Globigerinaceae, Part 1. 2, 1.

banner, f. t., see also eames, f. e.
barker, d., see kaye, p.

C. 8439

c

26 AUTHOR LIST

23. Barnard, p. d. w. 1960. Calathospermum fimbriatum sp. nov., a Lower Carboniferous Pterido-
sperm cupule from Scotland. 3, 265.

24. Barnard, p. d. w. 1962. Revision of the genus Amyelon Williamson. 5, 213.

25. barnard, t. 1958. Some Mesozoic adherent foraminifera. 1, 116.

26. barnard, t. 1963. Polymorphinidae from the Upper Cretaceous of England. 5, 712.

27. barnard, t. 1963. The morphology and development of species of Marssonella and Pseudo tex-
tulariella from the Chalk of England. 6, 41.

28. Barr, f. t. 1962. Upper Cretaceous planktonic foraminifera from the Isle of Wight, England.
4, 552.

29. barr, f. t. 1966. The foraminiferal genus Bolivinoides from the Upper Cretaceous of the British
Isles. 9, 220.

30. barr, f. t. 1966. Upper Cretaceous foraminifera from the Bally deenlea Chalk, County Kerry,
Ireland. 9, 492.

31. bate, r. h. 1965. Freshwater ostracods from the Bathonian of Oxfordshire. 8, 749.

32. bates, d. e. b. 1965. A new Ordovician crinoid from Dolgellau, North Wales. 8, 355.

33. bates, d. e. b. 1968. On ' Dendrocr inns' cambriensis Hicks, the earliest known crinoid. 11, 406.

34. batten, d. J. 1968. Probable dispersed spores of Cretaceous Equisetites. 11, 633.

35. batten, d. J. 1969. Some British Wealden megaspores and their facies distribution. 12, 333.

BATTEN, R. L., See ROLLINS, H. B.

36. bekker-migdisova, e. e. 1967. Tertiary Homoptera of Stavropol and a method of reconstruction
of continental palaeobiocoenoses. 10, 542.

37. bennison, g. m. 1960. Lower Carboniferous non-marine lamellibranchs from East Fife, Scotland.
3, 137.

38. bennison, g. m. 1961. Small Naiadites obesus from the Calciferous Sandstone Series (Lower
Carboniferous) of Fife. 4, 300.

39. bisat, w. s. 1957. Upper Visean Goniatites from the Manifold valley, North Staffordshire. 1, 16.

40. black, m. 1964. Cretaceous and Tertiary coccoliths from Atlantic seamounts. 7, 306.

41. black, m. 1968. Taxonomic problems in the study of coccoliths. 11, 793.

42. blake, d. b. 1968. Pedicellariae of two Silurian echinoids from western England. 11, 576.
blow, w. h., see banner, f. t. ; eames, f. e.

BOEKEL, N. M. VAN, See SOMMER, F. W.

boellistorff, j. d., see fagerstrom, j. a.

43. bolton, t. e. 1966. Some late Silurian Bryozoa from the Canadian Arctic Islands. 9, 517.

44. bonaparte, J. f. 1967. New vertebrate evidence for a southern transatlantic connection during the
Lower or Middle Triassic. 10, 554.

boucot, a. j., see amqs, a.; walmsley, v. g.

45. boucot, a. J. 1963. The Eospiriferidae. 5, 682.

46. boulter, m. c. 1968. A species of compressed lycopod sporophyll from the Upper Coal Measures
of Somerset. 11, 445.

BREIMER, A., See JOYSEY, K. A.

47. brett, d. w. 1960. Fossil oak wood from the British Eocene. 3, 86.

48. brett, d. w. 1966. Fossil wood of Anacardiaceae from the British Eocene. 9, 360.

49. brett, d. w. 1968. Studies on Triassic fossil plants from Argentina. III. The trunk of Rhexoxylon.
11, 236.

50. brown, d. A., Campbell, k. s. w., and Roberts, J. 1965. A Visean cephalopod fauna from New
South Wales. 7, 682.

51. brunton, h. 1965. The pedicle sheath of young productacean brachiopods. 7, 703.

52. brunton, h. 1966. Predation and shell damage in Visean brachiopod fauna. 9, 355.

AUTHOR LIST

27

53. bruton, d. l. 1966. A new odontopleurid trilobite genus from the Devonian of Bohemia. 9, 330.

54. bruton, d. l. 1967. Silurian odontopleurid trilobites from Sweden, Estonia, and Latvia. 10, 214.

55. bulman, o. m. b. 1958. The Sequence of Graptolite faunas. 1, 159.

56. bulman, o. m. b. 1963. On Glyptograptus dentatus (Brongniart) and some allied species. 6, 665.

57. bulman, o. m. b., and rickards, r. b. 1968. Some new diplograptids from the Llandovery of
Britain and Scandinavia. 11, 1.

bulman, o. m. b., see rickards, r. b.

BURBRIDGE, P. P ., See FELIX, C. J.

58. burgess, i. c. 1965. Calcifolium (Codiaceae) from the Upper Visean of Scotland. 8, 192.

59. burnaby, t. p. 1962. The palaeoecology of the foraminifera of the Chalk Marl. 4, 599.

60. butcher, n. e., and hodson, f. 1960. A review of the Carboniferous goniatite zones in Devon
and Cornwall. 3, 75.

61. butterworth, m. a., and spinner, e. 1967. Lower Carboniferous spores from North-west England.
10, 1.

callomon, j. h., see arkell, w. j.

62. Campbell, k. s. w. 1959. The Martiniopsis- like Spiriferids of the Queensland Permian. 1, 333.

63. Campbell, k. s. w. 1959. The type species of three Upper Palaeozoic punctate Spiriferoids. 1, 351.

64. Campbell, k. s. w. 1961. Carboniferous fossils from the Kuttung rocks of New South Wales. 4,
428.

65. Campbell, k. s. w. 1965. An almost complete skull roof and palate of the Dipnoan Dipnorhynchus
sussmilchi (Etheridge). 8, 634.

66. Campbell, k. s. w., and Roberts, J. 1964. Two species of Delepinea from New South Wales.
7, 514.

CAMPBELL, K. S. W., See AMOS, A. J.; BROWN, D. A.

67. carroll, r. l. 1969. A new family of Carboniferous amphibians. 12, 537.

68. carter, d. j. 1957. The distribution of the Foraminifer Alliatina excentrica (di Napoli Alliata)
and the new genus AlliatineUa. 1, 76.

69. carter, r. m. 1968. Functional studies on the Cretaceous oyster Arctostrea. II, 458.

70. casey, r. 1957. The Cretaceous Ammonite genus Leymeriella, with a systematic account of its
British occurrences. 1, 29.

71. casey, r. 1960. Hengestites, a new genus of Gault ammonites. 2, 200.

72. casey, r. 1960. A lower Cretaceous gastropod with fossilized intestines. 2, 270.

73. casey, r. 1960. A new echinoid from the Lower Cretaceous (Albian) of Kent. 3, 260.

74. casey, r. 1961. The stratigraphical palaeontology of the Lower Greensand. 3, 487.

75. casey, r. 1961. Stratigraphical palaeontology of the Lower Greensand: nomenclatural corrections.
4, 312.

76. cave, r., and dean, w. t. 1959. Four British Ordovician species of dalmanelloid brachiopod. 1,
292.

77. chaloner, w. g. 1958. A Carboniferous Selaginellites with Densosporites microspores. 1, 245.

78. chaloner, w. g. 1959. Devonian megaspores from Arctic Canada. 1, 321.

79. chaloner, w. g., and lorch, j. 1960. An opposite-leaved conifer from the Jurassic of Israel. 2, 236.

80. chaloner, w. g., and clarke, r. f. a. 1962. A new British Permian spore. 4, 648.

81. chaloner, w. g. 1962. A Sporangiostrobus with Densosporites microspores. 5, 73.

82. chaloner, w. g., and pettitt, j. m. 1964. A seed megaspore from the Devonian of Canada. 7, 29.
chaloner, w. g., see Mortimer, m. g.

83. chamberlain, j. a., jr. 1969. Technique for scale modelling of cephalopod shells. 52, 48.

84. chaphekar, M. 1963. Some calamitean plants from the Lower Carboniferous of Scotland. 6, 408.

85. chaphekar, m. 1965. On the genus Pothocites Paterson. 8, 107.

C 2

C. 8439

28 AUTHOR LIST

86. chlupac, ivo. 1963. Phyllocaricl crustaceans from the Silurian and Devonian of Czechoslovakia.
6, 97.

CHURKIN, M., see LENZ, A. C.

87. churkin, m., jr. 1966. Morphology and stratigraphic range of the phyllocarid crustacean
Caryocaris from Alaska and the Great Basin. 9, 371.

88. churkin, m., eberlein, G. d., hueber, F. M., and mamay, s. h. 1969. Lower Devonian land plants
from graptolitic shale in south-eastern Alaska. 12, 559.

CLARKE, R. F. A., See CHALONER, W. G.

89. clarke, r. f. a. 1965. Keuper miospores from Worcestershire, England. 8, 294.

90. clarke, r. f. a. 1965. British Permian saccate and monosulcate miospores. 8, 322.

CLARKE, W. J., See EAMES, F. E.

91. clarkson, e. n. k. 1966. Schizochroal eyes and vision of some Silurian acastid trilobites. 9, 1.

92. clarkson, e. n. k. 1966. Schizochroal eyes and vision in some phacopid trilobites. 9, 464.

93. clarkson, e. n. k. 1967. Fine structure of the eye in two species of Phacops (Trilobita). 10, 603.

94. clemens, w. A. 1963. Wealden mammalian fossils. 6, 55.

95. clemens, w. a. 1963. Late Jurassic mammalian fossils in the Sedgwick Museum, Cambridge.
6, 373.

96. cocks, l. r. M. 1967. Llandovery stropheodontids from the Welsh Borderland. 10, 245.
cocks, l. r. m., see also ziegler, a. m.

97. collins, J. 1961. Eocene crabs in a London Clay nodule. 4, 85.

98. collins, J. s. H. 1965. Arcoscalpelhtm comptum (Withers), a species of cirripede new to the Gault.
8, 629.

CONKIN, B. M., see conkin, j. e.

99. conkin, j. e., and conkin, b. m. 1968. A revision of some Devonian Foraminifera from Western
Australia. 11, 601.

COOGAN, A. H., See WILSON, E. C.

100. cookson, Isabel c., and eisenack, a. 1960. Upper Mesozoic microplankton from Australia and
New Guinea. 2, 243.

101. cookson, Isabel c. 1961. Hoegisporis, a new Australian Cretaceous form genus. 3, 485.

102. cookson, Isabel c., and dettmann, mary e. 1961. Reappraisal of the Mesozoic microspore genus
Aequitriradites. 4, 425.

103. cookson, i. c. 1965. On a new species of Hoegisporis Cookson. 8, 39.

104. cookson, i. c., and hughes, n. f. 1964. Microplankton from the Cambridge Greensand (mid-
Cretaceous). 7, 37.

105. cope, J. c. w. 1968. Propectinatites, a new Lower Kimmeridgian ammonite genus. 11, 16.

106. cope, J. c. w. 1968. Epizoic oysters on Kimmeridgian ammonites. 11, 19.

107. copeland, m. J. 1960. Ostracoda from the Upper Silurian Stonehouse Formation, Arisaig, Nova
Scotia, Canada. 3, 93.

108. copper, p. 1965. Unusual structures in Devonian Atrypidae from England. 8, 358.

109. copper, p. 1967. Spinatrypa and Spinatrypina (Devonian Brachipoda). 10, 489.

110. cordey, w. g. 1963. The genera Brotzenia and Voorthuysenia (Foraminifera) and Hofker’s
classification of the Epistomariidae. 6, 653.

111. cordey, w. G. 1967. The development of Globigerinoides ruber (d'Orbigny 1839) from the Miocene
to Recent. 10, 647.

112. cordey, w. g. 1968. A new Eocene Cassigerinella from Florida. 11, 368.

113. cordey, w. g. 1968. Morphology and phytogeny of Orbulinoides beckmannii (Saito 1962). 11, 371.

114. cowen, r. 1968. A new type of delthyrial cover in the Devonian brachipod Mucrospirifer. 11,
317.

AUTHOR LIST 29

115. cox, l. r. 1960. The preservation of moulds of the intestine in fossil Nuculana (Lamellibranchia)
from the Lias of England. 2, 262.

116. cox, l. r. 1962. New genera and subgenera of Mesozoic Bivalvia. 4, 592.

117. cox, l. r. 1962. Tertiary Bivalvia from Libya. 5, 1.

118. cox, L. r. 1963. The Rhaetic-Hettangian bivalve genus Pteromya Moore. 6, 582.

119. craig, G. y., and hallam, a. 1963. Size-frequency and growth-ring analyses of Mytilns echtlis and
Cardium edule , and their palaeoecological significance. 6, 731.

120. craig, G. y., and jones, n. s. 1966. Marine benthos, substrate and palaeoecology. 9, 30.

121. crespin, Irene. 1961. Upper Devonian Foraminifera from Western Australia. 3, 397.

122. cridland, A. a. 1964. Amyelon in American coal balls. 7, 186.

123. currey, J. d. 1962. The histology of the bone of a prosauropod dinosaur. 5, 238.

124. currie, ethel d., and george, t. Neville. 1963. Catalogue of described and figured specimens
in the Begg Collection in the Hunterian Museum of the University of Glasgow. 6, 378.

125. curtis, M. L. k. 1959. The Upper Llandovery Trilobites of the Tortworth Inlier, Gloucestershire.

1, 139.

126. cutbill, J. l., and forbes, c. l. 1967. Graphical aids for the description and analysis of variation
in fusuline foraminifera. 10, 322.

dean, w. t., see cave, r.

127. dean, w. t. 1959. Duftonia, a new Trilobite genus from the Ordovician of England and Wales.

2, 143.

128. dean, w. t. 1960. The Silurian Trilobite Dalmcmites myops (Konig). 2, 280.

129. dean, w. t., and krummenacher, r. 1961. Cambrian trilobites from the Amanos Mountains,
Turkey. 4, 71.

130. dean, w. t. 1962. The Ordovician trilobite genus Tiresias M’Coy, 1846. 5, 340.

131. delcourt, a. f., dettmann, m. e., and hughes, n. f. 1963. Revision of some Lower Cretaceous
microspores from Belgium. 6, 282.

delevoryas, t., see jain, r. e.

132. delevoryas, t. 1964. A probable pteridosperm microsporangiate fructification from the Penn-
sylvanian of Illinois. 7, 60.

DETTMANN, MARY E., See COOKSON, ISABEL C.; DELCOURT, A. F.

DETTMANN, MARY E., and PLAYFORD, G., SC’C also HUGHES, N. F.

133. dettmann, mary e., and playford, g. 1963. Sections of some spores from the Lower Car-
boniferous of Spitsbergen. 5, 679.

134. dickins, J. M. 1961. Eurydesma and Peruvispira from the Dwyka Beds of South Africa. 4, 138.

135. dickins, J. m. 1960. The Permian Leiopteriid Merismopteria and the origin of the Pteriidae.

3, 387.

136. dickins, J. m. 1960. Characters and relationships of the Mesozoic Pelecypod Pseudavicu/a. 3,
392.

137. dickins, j. m. 1961. Permian pelecypods newly recorded from Eastern Australia. 4, 119.

138. dickins, j. m. 1961. The gastropod Platyteichum in the Permian of Western Australia. 4, 131.

139. dineley, d. l. 1964. New specimens of Traquairaspis from Canada. 7, 210.

DONOVAN, D. T., See HOWARTH, M. K.

140. donovan, d. t. 1962. New information on the Toarcian ammonite genus Pseudolillia Maubeuge,
1949. 5, 86.

141. donovan, d. t. 1966. The Lower Liassic ammonites Neomicroceras gen. nov. and Paracymbites.

9, 312.

downie, c., see lister, t. r. ; wall, d.

142. downie, c. 1959. Hystrichospheres from the Silurian Wenlock Shale of England. 2, 56.

30 AUTHOR LIST

143. downie, c., and sarjeant, w. a. s. 1963. On the interpretation and status of some hystrichosphere
genera. 6, 83.

144. downie, c. 1963. ‘Hystrichospheres’ (acritarchs) and spores of the Wenlock Shales (Silurian)
of Wenlock, England. 6, 625.

145. doyle, J. c. 1963. A new heteromorph ammonite from the Lower Cretaceous of Yorkshire. 6,
575.

DROZDZEWSKI, G., See SEILACHER, A.

146. dunlop, grace m. 1962. Shell development in Spirifer trigonalis from the Carboniferous of
Scotland. 4, 477.

DURHAM, J. W., see ZULLO, V. A.

147. eagar, r. m. c. 1962. New Upper Carboniferous non-marine lamellibranchs. 5, 307.

EAMES, F. E., see SMOUT, A. H.

148. eames, f. e., and clarke, w. j. 1967. A Palaeocene Heterostegina. 10, 314.

149. eames, f. e., clarke, w. J., banner, F. t., smout, a. h., and blow, w. h. 1968. Some larger
foraminifera from the Tertiary of Central America. 11, 283.

150. eames, f. e. 1968. Sindulites, a new genus of the Nummulitidae (Foraminiferidae). 11, 435.
eberlein, g. d., see churkin, m.

151. edwards, d. 1968. A new plant from the Lower Old Red Sandstone of South Wales. 11, 683.

EGGERT, D. A., See HIBBERT, F. A.; TAYLOR, T. N.

152. eggert, d. a., and kryder, r. w. 1969. A new species of Aulacotheca (Pteridospermales) from
the Middle Pennsylvanian of Iowa. 12, 414.

eisenack, a., see cookson, i. c.

153. elliott, g. f. 1957. Subterraniphyllam, a new Tertiary calcareous alga. I, 73.

154. elliott, g. f. 1958. Alga debris-facies in the Cretaceous of the Middle East. 1, 254.

155. elliott, g. f. 1961. The sexual organization of Cretaceous Permocalculus (Calcareous Algae).
4, 82.

156. elliott, g. f. 1963. Problematical microfossils from the Cretaceous and Palaeocene of the
Middle East. 6, 293.

157. elliott, g. f. 1963. A Palaeocene teredinid (Mollusca) from Iraq. 6, 315.

158. elliott, g. f. 1965. Tertiary solenoporacean algae and the reproductive structures of the Soleno-
poraceae. 7, 695.

159. elliott, g. f. 1965. The interrelationships of some Cretaceous Codiaceae (Calcareous Algae).
8, 199.

160. elliott, g. f. 1968. Three new Tethyan Dasycladaceae (Calcareous algae). 11, 491.

161. esker, g. c., iii. 1968. Colour markings in Phacops and Greenops from the Devonian of New
York. 11,498.

162. fagerstrom, j. a., and boellstorff, J. d. 1964. Taxonomic criteria in the classification of the
Pennsylvanian productoid Juresania nebrascensis. 7, 23.

163. felix, c. J. 1965. Neogene Tasmanites and Leiospheres from Southern Louisiana, U.S.A. 8, 16.

164. felix, c. j., and burbridge, p. p. 1967. Palynology of the Springer Formation of southern
Oklahoma, U.S.A. 10, 349.

165. figge, k. 1968. A goniatite fauna from the Visean/Namurian boundary. 11, 264.

166. fisher, m. j. 1969. Benthonic Foraminifera from the Maestrichtian Chalk of Galicia Bank, west
of Spain. 12, 189.

167. forbes, c. l. 1960. Carboniferous and Permian Fusulinidae from Spitsbergen. 2, 210.

FORBES, C. L., see CUTBILL, J. L.

168. ford, t. d. 1965. The Palaeoecology of the Goniatite Bed at Cowlow Nick, Castleton, Derbyshire.
8, 186.

AUTHOR LIST

31

FRIEND, J. K., see FUNNELL, B. M.

169. fritz, w. h. 1968. Lower and early Middle Cambrian trilobites from the Pioche Shale, East-
central Nevada. U.S.A. 11, 183.

170. funnell, b. m., friend, J. k., and ramsay, a. t. s. 1969. Upper Maestrichtian planktonic Foramini-
fera from Galicia Bank, west of Spain. 12, 19.

171. furnish, w. m., quinn, J. H., and mccaleb, j. a. 1964. The Upper Mississippian ammonoid Dele-
pinoceras in North America. 7, 173.

172. gardiner, b. g. 1961. New Rhaetic and Liassic beetles. 4, 87.

173. gardiner, b. g. 1962. Namaichthys schroederi Gtirich and other Palaeozoic fishes from South
Africa. 5, 9.

GEORGE, T. NEVILLE, See CURRIE, ETHEL D.

174. glaessner, m. f., and wade, mary. 1966. The late Precambrian fossils from Ediacara, South
Australia. 9, 599.

175. gobbett, d. j. 1961. The Permian brachipod genus Horridonia Chao. 4, 42.

176. goldring, r. 1958. Lower Tournaisian trilobites in the Carboniferous Limestone facies of the
south-west province of Great Britain and of Belgium. 1, 231.

177. goldring, r. 1967. Cyclus martinensis sp. nov. (Crustacea) from the Upper Visean of the Mendip
Hills, England. 10, 317.

goldring, r., see amos, a. j.

GOOGAN, A. H., See WILSON, E. C.

178. Gordon, w. A. 1962. Some foraminifera from the Ampthill Clay, Upper Jurassic, of Cambridge-
shire. 4, 520.

179. grant, r. e. 1966. Late Permian trilobites from the Salt Range, West Pakistan. 9, 64.

180. Griffith, J. 1962. The Triassic fish Saurichthys krambergeri Schlosser. 5, 344.

181. habib, d. 1966. Distribution of spore and pollen assemblages in the Lower Kittanning Coal
Measures of western Pennsylvania. 9, 629.

182. hackman, b. d., and knill, j. l. 1962. Calcareous algae from the Dalradian of Islay. 5, 268.

183. hallam, a. 1960. Kulindrichnus langi, a new trace fossil from the Lias. 3, 64.

184. hallam, a. 1962. Brachiopod life assemblages from the Marlstone Rock-bed of Leicestershire.
4, 653.

185. hallam, a. 1963. Observations on the palaeoecology and ammonite sequence of the Frodingham
Ironstone (Lower Jurassic). 6, 554.

186. hallam, A. 1965. Environmental causes of stunting in living and fossil marine benthonic inverte-
brates. 8, 132.

187. hallam, A. 1967. The interpretation of size-frequency distributions in molluscan death-
assemblages. 10, 25.

188. hallam, a. 1969. Faunal realms and facies in the Jurassic. 12, 1.
hallam, a., see also craig., g. y.

189. Hamilton, d. 1958. An Eocene Sea-Pen from Dunedin, New Zealand. 1, 226.

190. Hamilton, d. 1961. Algal growths in the Rhaetic Cotham Marble of Southern England. 4, 324.
harper, c. w., see walmsley, v. g.

191. Harris, T. m. 1961. The fossil cycads. 4, 313.

HAUDE, R., see SEILACHER, A.

HAYNES, J., see ADAMS, T. D.

192. hendry, r. d., rowell, A. j., and Stanley, j. w. 1963. A rapid parallel grinding machine for
serial sectioning of fossils. 6, 145.

hibbert, f. a., see sullivan, h. j.

193. hibbert, f. a., and eggert, d. a. 1965. A new calamitalean cone from the Middle Pennsylvanian
of Southern Illinois. 8, 681.

32 AUTHOR LIST

194. hibbert, f. a., and lacey, w. s. 1969. Miospores from the Lower Carboniferous Basement Beds
in the Menai Straits region of Caernarvonshire, North Wales. 12, 420.

195. hill, d. 1967. The sequence and distribution of Ludlovian, Lower Devonian, and Couvinian
coral faunas in the Union of Soviet Socialist Republics. 10, 660.

HODSON, F., see BUTCHER, N. E.

196. hodson, F. 1958. Entogonites cf. borealis, an Alaskan goniatite from Ireland. 1, 158.

197. hodson, F., and moore, e. w. j. 1959. Goniatites striatus and related forms from the Visean of
Ireland. 1, 384.

198. holden, j. c. 1964. Upper Cretaceous ostracods from California. 7, 393.

199. holdsworth, B. k. 1965. The Namurian goniatite Nuculoceras stellarum (Bisat). 8, 226.

200. holdsworth, b. k. 1966. Radiolaria from the Namurian of Derbyshire. 9, 319.

201. Holland, c. h. 1965. On the nautiloid Leurocycloceras from the Ludlovian of Wales and the
Welsh Borderland. 7, 525.

202. Holland, c. h., rickards, r. b., and warren, p. t. 1969. The Wenlock graptolites of the Ludlow
district, Shropshire, and their stratigraphical significance. 12, 663.

203. holwill, f. J. w. 1964. The coral genus Metriophyllum Edwards and Haime. 7, 108.

204. holwill, f. j. w. 1968. Tabulate corals from the Ilfracombe Beds (Middle-Upper Devonian) of
North Devon. 11, 44.

205. house, m. r. 1960. Abnormal growths in some Devonian goniatites. 3, 129.

206. house, m. r. 1961. Acanthoclymenia, the supposed earliest Devonian clymeniid, is a Manti-
coceras.

207. house, m. R., and pedder, a. e. h. 1963. Devonian goniatites and stratigraphical correlations in
Western Canada. 6, 491.

208. howarth, M. k. 1962. The Yorkshire type ammonites and nautiloids of Young and Bird, Phillips,
and Martin Simpson. 5, 93.

209. howarth, m. k., and donovan, d. t. 1964. Ammonites of the Liassic family Juraphyllitidae in
Britain. 7, 286.

210. hubbard, J. A. e. b. 1966. Population studies in the Ballyshannon Limestone, Ballina Limestone,
and Rinn Point Beds (Visean) of NW. Ireland. 9, 252.

211. Hudson, j. d. 1963. The recognition of salinity-controlled mollusc assemblages in the Great
Estuarine Series (Middle Jurassic) of the Inner Hebrides. 6, 318.

212. Hudson, J. d. 1963. The ecology and stratigraphical distribution of the invertebrate fauna of the
Great Estuarine Series. 6, 327.

213. Hudson, J. d. 1968. The microstructure and mineralogy of the shell of a Jurassic mytilid (Bivalvia).
11, 163.

214. Hudson, r. g. s. 1958. Actostroma gen. nov., a Jurassic stromatoporoid from Maktesh Hathira,
Israel. 1, 87.

215. Hudson, R. G. s. 1958. Permian corals from northern Iraq. 1, 174.

216. Hudson, R. G. s. 1959. A revision of the Jurassic stromatoporoids Actinostromina, Astrostylopsis,
and Trupetostromaria Germovsek. 2, 28.

217. Hudson, r. G. s. 1966. The Tethyan Jurassic stromatoporoids Stromatoporina, Dehornella, and
Astroporina. 2, 180.

218. Hudson, r. g. s., and jefferies, r. p. s. 1961. Upper Triassic brachiopods and lamellibranchs from
the Oman Peninsula, Arabia. 4, 1.

hueber, f. m., see churkin, m.

hughes, n. f., see cookson, i. c.; delcourt, a. f.

219. hughes, n. f. 1961. Further interpretation of Eucommiidites Erdtman 1948. 4, 292.

220. hughes, n. f., dettmann, mary e., and playford, g. 1962. Sections of some Carboniferous dis-
persed spores. 5, 247.

AUTHOR LIST 33

221. hughes, n. f., and moody-stuart, j. 1966. Descriptions of schizaeaceous spores taken from early
Cretaceous macrofossils. 9, 274.

222. hughes, n. F., and moody-stuart, j. c. 1969. A method of stratigraphic correlation using early
Cretaceous miospores. 12, 84.

223. hunter, h. a., and langston, w. 1965. Odontoma in a northern mammoth, 7, 674.

224. hutchison, r., and ingham, j. k. 1967. New trilobites from the Tremadoc Series of Shropshire.
10, 47.

INGHAM, J. K., see HUTCHISON, R.

225. jackson, D. e., and lenz, a. c. 1963. A new species of Monograptus from the Road River Forma-
tion, Yukon. 6, 751.

226. jain, R. K., and delevoryas, t. 1967. A Middle Triassic flora from the Cacheuta Formation,
Minas de Petroleo, Argentina. 10, 564.

jakobson, m. e., see mckerrow, w. s., also Kennedy, w. j.

227. james, judith. 1965. The development of a dicellograptid from the Balclatchie Shales of Laggan
Burn. 8, 41.

228. Jefferies, r. p. s. 1960. Photonegative young in the Triassic Lamellibranch Lima Iineata (Schlo-
theim). 3, 362.

229. Jefferies, R. p. s. 1962. The palaeoecology of the Actinocamax plenus Subzone (lowest Turonian)
in the Anglo-Paris Basin. 4, 609.

230. jefferies, r. p. s., and minton, r. p. 1965. The mode of life of two Jurassic species of ‘ Posidonia '
(Bivalvia). 8, 156.

231. jefferies, r. p. s. 1969. Ceratocystis perneri Jaekel — a Middle Cambrian chordate with echino-
derm affinities. 12, 494.

jefferies, r. p. s., see also Hudson, r. g. s.

232. jeletzky, J. A. 1963. Malayomaorica gen. nov. (Family Aviculopectinidae) from the Indo-
Paciffc Upper Jurassic; with comments on related forms. 6, 148.

233. jenkins, t. b. h. 1960. Non-Marine Lamellibranch assemblages from the Coal Measures (Upper
Carboniferous) of Pembrokeshire, West Wales. 3, 104.

234. jenkins, t. b. h. 1966. The Upper Devonian index ammonoid Cheiloceras from New South Wales.
9, 458.

235. jenkins, t. b. h. 1968. Famennian ammonoids from New South Wales. 11, 535.

236. jenkins, w. a. m„ 1967. Ordovician Chitinozoa from Shropshire. 10, 436.

237. joffe, Joyce. 1967. The ‘dwarf’ crocodiles of the Purbeck Formation, Dorset: a reappraisal. 10,
629.

238. Johnson, G. a. l. 1958. Biostromes in the Namurian Great Limestone of northern England. 1,
147.

239. Johnson, h. m. 1966. Silurian Girvanella from the Welsh Borderland. 9, 48.

240. Johnson, j. g. 1966. Middle Devonian brachiopods from the Roberts Mountains, central Nevada.
9, 152.

241. Johnson, J. G. 1966. Paraclionetes, a new Lower and Middle Devonian brachiopod genus. 9, 365.

242. Johnson, J. G., and talent, j. a. 1967. Cortezorthinae, a new subfamily of Siluro-Devonian
dalmaneffid brachiopods. 10, 142.

JOHNSON, R. T., see MCKERROW, W. S.; also KENNEDY, W. J.

jones, n. s., see craig, g. y.

243. joysey, K. A. 1959. Probable Cirripede, Phoronid, and Echiuroid burrows within a Cretaceous
Echinoid test. 1, 397.

244. joysey, k. A., and breimer, a. 1963. The anatomical structure and systematic position of Penta-
blastus (Blastoidea) from the Carboniferous of Spain. 6, 471.

245. jull, r. k. 1965. Corallum increase in Lithostrotion. 8, 204.

34 AUTHOR LIST

246. jull, r. k. 1967. The hystero-ontogeny of Lonsdaleia McCoy and Thysanophyllum orientale
Thomson. 10, 617.

KAAR, R. F., see ZULLO, V. A.

247. kato, M., and mitchell, m. 1961. Slimoniphyllum, a new genus of Lower Carboniferous coral
from Britain. 4, 280.

248. kaye, p. 1963. The ostracod genus Neocythere in the Speeton Clay. 6, 274.

249. kaye, p. 1963. The ostracod species Orthonotacythere inversa (Cornuel) and its allies in the
Speeton Clay of Yorkshire. 6, 430.

250. kaye, p. 1964. Revision of the Ostracoda from the Bargate Beds in Surrey. 7, 317.

251. kaye, p., and barker, d. 1965. Ostracoda from the Sutterby Marl (U. Aptian) of South Lincoln-
shire. 8, 375.

252. kaye, p., and barker, d. 1966. Ostracoda from the Upper Tealby Clay (Lower Barremian) of
South Lincolnshire. 9, 208.

253. kelly, F. b. 1967. Silurian leptaenids (Brachiopoda). 10, 590.

254. kemp,e. m. 1968. Probable angiosperm pollen from the British Barremian to Albian strata. 11, 421.

255. Kennedy, w. j., and macdougall, j. d. s. 1969. Crustacean burrows in the Weald Clay (Lower
Cretaceous) of South-eastern England and their environmental significance. 12, 459.

256. Kennedy, w. J., jakobson, M. e., and Johnson, r. t. 1969. A Favreina-Thalassinoides association
from the Great Oolite of Oxfordshire. 12, 549.

257. kier, P. M. 1969. A Cretaceous echinoid with false teeth. 12, 488.

KILENYI, T. I., See NEALE, J. W.

258. kilenyi, t. i. 1965. Oertliana, a new ostracod genus from the Upper Jurassic of North-West
Europe. 8, 572.

259. kilenyi, t. i., and allen, n. w. 1968. Marine-brackish bands and their microfauna from the lower
part of the Weald Clay of Sussex and Surrey. 11, 141.

260. kilenyi, t. i. 1969. The Ostracoda of the Dorset Kimmeridge Clay. 12, 112.

261. krinsley, d., and schneck, m. 1964. The palaeoecology of a transition zone across an Upper Cre-
taceous boundary in New Jersey. 7, 266.

262. krommelbein, k. 1968. The first non-marine Lower Cretaceous ostracods from Ghana, West
Africa. 11, 259.

krummenacher, r., see dean, w. t.

KRYDER, R. W., See EGGERT, D. A.

263. lagaaij, R. 1963. Cupuladria canariensis (Busk) — portrait of a bryozoan. 6, 172.

LANGSTON, W., See FIUNTER, H. A.

LARGE, N. F., See SAVAGE, R. J. G.

LARWOOD, G. P., See THOMAS H. DIGHTON
LENZ, A. C., See JACKSON, D. E.

264. lenz, a. c., and churkin, m., jr. 1966. Upper Ordovician trilobites from northern Yukon. 9, 39.

265. lewarne, G. c. 1964. Starfish traces from the Namurian of County Clare, Ireland. 7, 508.

LISTER, T. R., See RICHARDSON, J. B.

266. lister, t. r., and downie, c. 1967. New evidence for the age of the primitive echinoid Myrias-
tiches gigas. 10, 171.

267. lloyd, A. J. 1959. Arenaceous Foraminifera from the type Kimeridgian (Upper Jurassic). 1, 298.

268. logan, a. 1963. The new brachiopod genus Howseia from the Permian Magnesian Limestone of
Durham. 6, 754.

269. logan, A. 1964. The dentition of the Durham Permian pelecypod Permophorus costatus (Brown).
7, 281.

LORCH, J., See CHALONER, W. G.

AUTHOR LIST 35

270. love, l. g. 1962. Further studies on micro-organisms and the presence of syngenetic pyrite. 5,
444.

271. love, l. g. 1963. The composition of Pyritosphaera barbaria Love 1957. 6, 119.

MACDOUGALL, J. D. S., See KENNEDY, W. J.

272. macgregor, a. r. 1961. Upper Llandeilo Brachiopods from the Berwyn Hills, North Wales. 4,
177.

273. macgregor, a. r. 1963. Upper Llandeilo Trilobites from the Berwyn Hills, North Wales. 5, 790.

274. macurda, d. b., jr. 1966. The Devonian blastoid Belocrinus from France. 9, 244.

275. mcalester, a. l. 1965. Systematics, affinities, and life habits of Babinka, a transitional Ordovician
lucinoid bivalve. 8, 231.

MCCALEB, J. A., See FURNISH, W. M.

276. mcgregor, d. c. 1960. Devonian spores from Melville Island, Canadian Arctic Archipelago. 3,
26.

277. mckerrow, w. s., Johnson, r. t., and jakobson, m. e. 1969. Palaeoecological studies in the Great
Oolite at Kirtlington, Oxfordshire. 12, 56.

mckerrow, w. s., see also ziegler, a. m.
mamay, s. h., see churkin, m.

278. manolov, J. r. 1952. New ammonites from the Barremian of North Bulgaria. 5, 527.

279. marshall, a. e., and smith, a. h. v. 1965. Assemblages of miospores from some Upper Car-
boniferous coals and their associated sediments in the Yorkshire coalfield. 7, 656.

280. martin, a. r. h. 1968. AquilapoUenites in the British Isles. 11, 549.

281. Matthews, s. c. 1969. A Lower Carboniferous conodont fauna from East Cornwall. 12, 262.

282. Matthews, s. c. 1969. Two conodont faunas from the Lower Carboniferous of Chudleigh, South
Devon. 12, 276.

283. maxwell, w. g. h. 1961. Lower Carboniferous gastropod faunas from Old Cannindah, Queens-
land. 4, 59.

MEADE, M. J., see RIXON, A. E.

284. medd, a. w. 1965. Dionella gen. nov. (superfamily Membraniporacea) from the Upper Cretaceous
of Europe. 8, 492.

285. medd, a. w. 1966. The fine structure of some Lower Triassic acritarchs. 9, 351.

286. Melville, r. v. 1961. Dentition and relationships of the echinoid genus Pygaster J. L. R. Agassiz,
1836. 4, 243.

287. middlemiss, f. a. 1959. English Aptian Terebratulidae. 2, 94.

288. middlemiss, f. a. 1962. Brachiopod ecology and Lower Greensand palaeogeography. 5, 253.

289. miller, t. g. 1961. Type specimens of the genus Penes tella from the Lower Carboniferous of
Great Britain. 4, 221.

290. miller, t. g. 1962. Some Wenlockian fenestrate Bryozoa. 5, 540.

291. miller, t. g. 1963. The bryozoan genus Polypora M’Coy. 6, 166.

292. miller, t. g. 1965. Time in stratigraphy. 8, 113.

MINTON, R. P., see JEFFERIES, R. P. S.

MITCHELL, M., See KATO, M.

MONROE, E. A., See SASS, D. B.

MOODY-STUART, J., See HUGHES, N. F.

MOORE, E. W. J., See HODSON, F.

293. moore, l. r. 1963. Microbiological colonization and attack on some Carboniferous miospores. 6,
349.

294. Mortimer, m. g., and chaloner, w. g. 1967. Devonian megaspores from the Wyboston borehole,
Bedfordshire, England. 10, 189.

36 AUTHOR LIST

295. Mortimer, m. g., and chaloner, w. g. 1967. Nomenclatural note: correction of name for a
Devonian megaspore. 10, 706.

296. NOTES FOR AUTHORS. 10, 707.

297. nagappa, y. 1959. Note on Operculinoides Hanzawa 1935. 2, 156.

298. neale, J. w. 1959. Normanicythere gen. nov. (Pleistocene and Recent) and the division of the
Ostracod family Trachyleberididae. 2, 72.

299. neale, J. w., and kilenyi, t. i. 1961. New species of Mandelstamia (Ostracoda) from the English
Mesozoic. 3, 439.

300. neale, j. w. 1961. Normanicythere leioderma (Norman) in North America. 4, 424.

301. neale, J. w. 1962. Ammonoidea from the Lower D Beds (Berriasian) of the Speeton Clay.

5, 272.

302. neves, r. 1961. Namurian plant spores from the southern Pennines, England. 4, 247.

303. newman, b. h. 1968. The Jurassic dinosaur Scelidosaurus harrisoni, Owen. 11, 40.

304. norford, b. s. 1960. A well-preserved Dinobolus from the Sandpile Group (Middle Silurian) of
northern British Columbia. 3, 242.

305. norford, b. s., and steele, h. miriam. 1969. The Ordovician trimerellid brachiopod Eodinobolus
from south-east Ontario. 12, 161.

306. norris, g. 1969. Miospores from the Purbeck Beds and marine Upper Jurassic of southern Eng-
land. 12, 574.

307. Oliver, w. a., jr. 1966. Description of dimorphism in Striatopora flexuosa Hall. 9, 448.

308. opik, a. a. 1961. Alimentary caeca of agnostids and other trilobites. 3, 410.

309. ormiston, a. r. 1966. Occurrence of Australosutura (Trilobita) in the Mississippian of Oklahoma,
U.S.A. 9, 270.

310. owen, d. E. 1960. Upper Silurian Bryozoa from Central Wales. 3, 69.

311. owen, d. E. 1962. Ludlovian Bryozoa from the Ludlow district. 5, 195.

312. owen, d. e. 1969. Wenlockian Bryozoa from Dudley, Niagara, and Gotland and their palaeo-
geographic implications. 12, 621.

313. packham, g. h. 1962. Some diplograptids from the British Lower Silurian. 5, 498.

314. palframan, d. f. b. 1966. Variation and ontogeny of some Oxfordian ammonities: Taramelliceras
richei (de Loriol) and Creniceras renggeri (Oppel), from Woodham, Buckinghamshire. 9, 290.

315. palframan, d. f. b. 1967. Variation and ontogeny of some Oxford Clay ammonites: Disticho-
ceras bicostatum (Stahl) and Horioceras baugieri (d’Orbigny), from England. 10, 60.

316. pant, d. d., and verma, b. k. 1963. On the Structure of leaves of Rhabdotaenia Pant from the
Raniganj coalfield, India. 6, 301.

317. pant, d. d., and singh, r. s. 1968. The structure of Vertebraria indica Royle. 11, 643.

318. parkinson, d. 1961. The Carboniferous rhynchonell id Pugnoides triplex (M’Coy). 3, 477.

319. paul, c. r. c. 1967. New Ordovician Bothriocidaridae from Girvan and a re-interpretation of
Bothriocidaris Eichwald. 10, 525.

320. paul, c. R. c. 1968. Macrocystella Callaway, the earliest glyptocystitid cystoid. 11, 580.

321. paul, c. R. c. 1968. Morphology and function of dichoporite pore-structures in cystoids. 11, 697.
pedder, a. e. h., see also house, m. r.

322. pedder, a. e. h. 1960. New species of brachiopods from the Upper Devonian of Hay River,
Western Canada. 3, 208.

323. pedder, a. e. h. 1963. Alaiophyllum mackenziense sp. nov., a Devonian tetracoral from Canada.

6, 132.

324. pedder, a. e. h. 1964. Correlation of the Canadian Middle Devonian Hume and Nahanni
Formations by tetracorals. 7, 430.

325. pedder, a. e. h. 1965. Some North American species of the Devonian tetracoral Smithiphyllum.
8, 618.

AUTHOR LIST

37

326. pedder, a. E. h. 1966. The Upper Devonian gastropod Orecopia in western Canada. 9, 142.
PETTIT, J. M., see CHALONER, W. G.

327. philip, g. m. 1960. The Middle Palaeozoic squamulate favositids of Victoria. 3, 186.

328. philip, g. m. 1963. The Jurassic echinoid Cidarites moniliferus Goldfuss and the status of Euci-
daris. 5, 785.

329. philip, g. m. 1963. Two Australian Tertiary neolampadids, and the classification of cassiduloid
echinoids. 6, 718.

330. Phillips, june r. p. 1960. Restudy of types of seven Ordovician bifoliate Bryozoa. 3, 1.

331. Phillips, t. l., and Andrews, h. n. 1968. Biscalitheca (Coenopteridales) from the Upper Penn-
sylvanian of Illinois. 11, 104.

332. pickett, j. w. 1960. A clymeniid from the Wockhimeria zone of New South Wales. 3, 237.

333. pienaar, r. n. 1968. Upper Cretaceous coccolithophorids from Zululand, South Africa. 11, 361.
playford, g., see also dettmann, mary e., and hughes, n. f.

334. playford, g. 1962-3. Lower Carboniferous microfloras of Spitsbergen. Part I. 5, 550. Part II.

5, 619.

335. pocock, k. J. 1964. Estaingia, a new trilobite genus from the Lower Cambrian of South Aus-
tralia. 7, 458.

336. pocock, y. p. 1966. Devonian schizophoriid brachiopods from western Europe. 9, 381.

337. pocock, y. p. 1968. Carboniferous schizophoriid brachiopods from western Europe. 11, 64.

338. pollard, j. e. 1966. A non-marine ostracod fauna from the Coal Measures of Durham and
Northumberland. 9, 667.

339. pollard, j. e. 1968. The gastric contents of an ichthyosaur from the Lower Lias of Lyme Regis,
Dorset. 31, 376.

340. prasad, k. n. 1964. Upper Miocene anthropoids from the Siwalik Beds of Haritalyanger, Hi-
machal Pradesh, India. 7, 124.

QUINN, J. H., see FURNISH, W. M.

RAMSAY, A. T. S., See FUNNELL, B. M.

341 . ramsbottom, w. h. c. 1958. British Upper Silurian crinoids from the Ludlovian. 1, 106.

342. ramsbottom, w. h. c. 1959. Distinctions between the Carboniferous Lamellibranch genera
Caneyella , Posidonia, and Posidoniella. 1, 405.

343. rawson, p. f. 1966. A phylloceratid ammonite from the Speeton Clay (Lower Cretaceous) of
Yorkshire. 9, 455.

344. regnell, g. 1960. The Lower Palaeozoic Echinoderm faunas of the British Isles and Balto-
Scandia. 2, 161.

345. reyment, r. a., and sandberg, p. 1963. Biometric study of Barremites sabdifficilis (Karakasch).

6, 727.

RHODES, F. H. T., See AUSTIN, R. L.

346. Richardson, J. b. 1960. Spores from the Middle Old Red Sandstone of Cromarty, Scotland. 3,
45.

347. richardson, j. b. 1962. Spores with bifurcate processes from the Middle Old Red Sandstone of
Scotland. 5, 171.

348. richardson, j. b. 1965. Middle Old Red Sandstone spore assemblages from the Orcadian basin,
north-east Scotland. 7, 559.

349. richardson, j. b., and lister, t. r. 1969. Upper Silurian and Lower Devonian spore assemblages
from the Welsh Borderland and South Wales. 12, 201.

350. rickards, r. b. 1965. Two new genera of Silurian phacopid trilobites. 7, 541.

351. rickards, r. b. 1965. New Silurian graptolites from the Howgill Fells (Northern England). 8, 247.

352. rickards, r. b., and bulman, o. m. b. 1965. The development of Lasiograptus harknessi (Nichol-
son 1867). 8, 272.

38

AUTHOR LIST

RICKARDS, R. B., See also BULMAN, O. M. B. ; HOLLAND, C. H.

353. ritchie, a. 1968. New evidence on Jamoytius kerwoodi White, an important ostracoderm from the
Silurian of Lanarkshire, Scotland. 11, 21.

354. rixon, a. e., and meade, m. j. 1960. Glass fibre resin casts of fossils. 3, 124.

ROBERTS, J., SCC Cllso BROWN, D. A.; CAMPBELL, K. S. W.

355. Roberts, j. 1968. Mantle canal patterns in Schizophoria (Brachiopoda) from the Lower Car-
boniferous of New South Wales. 11, 389.

356. Roberts, J. 1965. A Lower Carboniferous fauna from Trevallyn, New South Wales. 8, 54.

357. robinson, e. 1968. Chubbina , a new Cretaceous alveolinid genus from Jamaica and Mexico. 11,
526.

358. rolfe, w. d. i. 1962. A syncarid crustacean from the Keele Beds (Stephanian) of Warwickshire.

4, 546.

359. rolfe, w. d. i. 1962. The cuticle of some Middle Silurian ceratiocaridid Crustacea from Scotland.

5, 30.

360. rolfe, w. d. i. 1967. Rochdalia, a Carboniferous insect nymph. 10, 307.

361. rollins, h. b., and batten, r. l. 1968. A sinus-bearing monoplacophoran and its role in the classi-
fication of primitive molluscs. 11, 132.

362. ross, c. a. 1962. Silurian monograptids from Illinois. 5, 59.

363. ross, c. a. 1962. Permian foraminifera from British Honduras. 5, 297.

364. ross, c. a. 1963. Early Permian fusulinids from Macusani, Southern Peru. 5, 817.

365. ross, j. r. p. 1962. Early species of the bryozoan genus Phaenopora from the Caradoc Series,
Shropshire. 5, 52.

366. ross, J. r. p. 1963. Chazyan (Ordovician) leptotrypellid and atactotoechid Bryozoa. 5, 727 .

367. ross, j. r. p. 1963. Trepostome Bryozoa from the Caradoc Series, Shropshire. 6, 1.

368. ross, j. r. p. 1963. Lower Permian Bryozoa from Western Australia. 6, 70.

369. ross, .t. r. p. 1965. Homotrypa and Amplexoporal from the Caradoc Series, Shropshire. 8, 5.
ROWELL, A. J., see HENDRY, R. D.

370. rowell, a. J. 1962. The brachiopod genus Valdiviathyris Helmcke. 4, 542.

371. rudwick, m. J. s. 1961. The feeding mechanism of the Permian brachiopod Prorichthofenia . 3,
450.

372. rudwick, m. j. s. 1961. The anchorage of articulate brachiopods on soft substrata. 4, 475.

373. rudwick, m. j. s. 1964. The function of zigzag deflexions in the commissures of fossil brachio-
pods. 7, 135.

374. rudwick, M. J. s. 1965. Sensory spines in the Jurassic brachiopod Acanthothiris. 8, 604.

375. rudwick, m. j. s. 1968. The feeding mechanisms and affinities of the Triassic brachiopods Thecos-
pira Zugmayer and Bactrynium Emmrich. 11, 329.

376. runnegar, b. 1968. Preserved ligaments in Australian Permian bivalves. 11, 94.

377. rushton, a. w. a. 1967. The Upper Cambrian trilobite Irvingella nuneatonensis (Sharman). 10,
339.

378. rushton, a. w. a. 1968. Revision of two Upper Cambrian trilobites. 11, 410.

379. samanta, b. k. 1968. Nummulites (Foraminifera) from the Upper Eocene Kopili Formation of
Assam, India. 11, 669.

380. samanta, b. k. 1963. Two new species of Discocyclina (Foraminifera) from the Upper Eocene of
Assam, India. 6, 658.

SANDBERG, P., See REYMENT, R. A.

SARJEANT, W. A. S., See DOWNIE, C.

381. sarjeant, w. a. s. 1961. Microplankton from the Kellaways Rock and Oxford Clay of York-
shire. 4, 90.

AUTHOR LIST 39

382. sarjeant, w. a. s. 1962. Microplankton from the Ampthill Clay of Melton, South Yorkshire. 5,
478.

383. sarjeant, w. a. s. 1964. New name and diagnosis for an Upper Jurassic species of Gonyaulacysta
(Dinophyceae). 7, 472.

384. sass, d. b., and monroe, e. a. 1967. Shell-growth in recent terebratuloid brachiopoda. 10, 298.

385. savage, n. m. 1968. Planicardinia, a new septate dalmanellid brachiopod from the Lower Devonian
of New South Wales. 11, 627.

386. savage, n. m. 1968. Australirhynchia, a new Lower Devonian rhynchonelloid brachiopod from
New South Wales. 31, 731.

387. savage, n. m. 1969. New spiriferid brachiopods from the Lower Devonian of New South Wales.
12, 472.

savage, n. m., see also walmsley, v. g.

388. savage, r. j. g., and large, n. e. 1966. On Birgeria acuminata and the absence of labyrinthodonts
from the Rhaetic. 9, 135.

389. scrutton, c. t. 1965. Periodicity in Devonian coral growth. 7, 552.

390. scrutton, c. t. 1967. Marisastridae (Rugosa) from south-east Devonshire, England. 10, 266.

391. sdzuy, k. 1966. An improved method of analysing distortion in fossils. 9, 125.

392. seilacher, a., drozdzewski, g., and haude, r. 1968. Form and function of the stem in a pseudo-
planktonic crinoid ( Seirocrinus ). 11, 275.

393. selwood, e. b. 1960. Ammonoids and trilobites from the Upper Devonian and lowest Carboni-
ferous of the Launceston area of Cornwall. 3, 153.

394. sheils, k. a. g. 1966. A new productid brachiopod from the Upper Visean of Scotland. 9, 426.

395. shergold, j. h. 1966. A revision of Acaste downingiae (Murchison) and related trilobites. 9, 183.

396. shergold, j. h. 1967. A revision of Acastella spinosa (Salter 1864) with notes on related trilobites.
10, 175.

397. sherwin, l. 1968. Denckmannites (Trilobita) from the Silurian of New South Wales. 11, 691.

398. singh, h. p. 1964. A miospore assemblage from the Permian of Iraq. 7, 240.

SINGH, R. S., see PANT, D. D.

399. skevington, d. 1960. A new variety of Orthoretiolites hami Whittington. 2, 226.

400. skwarko, s. w. 1963. A new Upper Cretaceous Ophiuroid from Australia. 6, 579.

401. smith, a. h. v. 1960. Structure of the spore wall in certain Miospores belonging to the series
Cingulati Pot. and Klaus 1954. 3, 82.

smith, a. h. v., see marshall, a. e.

402. smith, d. l. 1962. Three fructifications from the Scottish Lower Carboniferous. 5, 225.

403. smith, j. d. d., and white, d. e. 1963. Cambrian Trilobites from the Purley Shales of Warwick-
shire. 6, 397.

404. smout, a. h., and eames, f. e. 1958. The genus Archaias (Foraminifera) and its stratigraphical
distribution. 1, 207.

405. smout, a. h., and sugden, w. 1962. New information on the foraminiferal genus Pfenderina. 4, 581 .
smout, a. h., see also eames, f. e.

406. sohn, i. g., and anderson, f. w. 1964. The ontogeny of Theriosynoecum fittoni (Mantell). 7, 72.

407. sommer, f. w., and van boekel, n. m. 1967. Brazilian Palaeozoic Algomycetes and Tasmanacea.
10, 640.

408. soot-ryen, h. 1969. A new species of Babinka (Bivalvia) from the Lower Ordovician of Oland,
Sweden. 12, 173.

409. sorauf, j. e. 1969. Lower Devonian Hexagonaria (Rugosa) from the Armorican Massif of Western
France. 12, 178.

410. speden, i. g. 1959. Phyllocrinus furcillatus sp. nov., a Cyrtocrinoid from the upper Jurassic of
Kawhia, New Zealand. 2, 150.

40

AUTHOR LIST

SPINNER, E., see BUTTERWORTH, M. A.

411. spinner, e. 1965. Westphalian D megaspores from the Forest of Dean Coalfield, England. 8, 82.

412. spinner, e. 1969. Megaspore assemblages from the Visean deposits at Dunbar, East Lothian,
Scotland. 12, 441.

413. spjeldN/ES, n. 1963. Some Upper Tremadocian graptolites from Norway. 6, 121.

414. spjeldN/ES, n. 1963. Some silicified Ordovician fossils from South Wales. 6, 254.

STANLEY, J. W., SCC HENDRY, R. D.

415. Stanley, J. w. 1964. Serial sectioning of steinkerns. 7, 105.

416. staplin, F. l. 1961. Reef-controlled distribution of Devonian microplankton in Alberta. 4,
392.

417. stearn, c. w. 1966. The microstructure of stromatoporoids. 9, 74.

STEELE, H. MIRIAM, See NORFORD, B. S.

418. stephenson, d. g. 1963. The spines and diffuse fascioles of the Cretaceous echinoid Echinocorys
scutata Leske. 6, 458.

419. stephenson, n. G. 1963. Growth gradients among fossil monotremes and marsupials. 6, 615.

420. stevens, G. r. 1963. The systematic status of Oppel’s specimens of Belemnites gerardi. 6, 690.

421. stevens, G. r. 1965. The belemnite genera Dicoelites Boehm and Prodicoelites Stolley. 7, 606.

422. stevens, g. r. 1965. A new belemnite from the Upper Jurassic of Indonesia. 7, 621.

423. stinton, f. c., and torrens, h. s. 1968. Fish otoliths from the Bathonian of southern England.
11, 246.

424. strachan, i. 1968. A medusoid (?) from the Silurian of England. II, 610.

425. strusz, d. l. 1961. Lower Palaeozoic corals from New South Wales. 4, 334.

426. strusz, d. l. 1965. Disphyllidae and Phacellophyllidae from the Devonian Garra Formation of
New South Wales. 8, 518.

427. strusz, d. l. 1966. Spongophyllidae from the Devonian Garra Formation, New South Wales.
9, 544.

428. strusz, d. l. 1967. Chalamydophyllum, Iowaphyllum, and Sinospongophyllum (Rugosa) from the
Devonian of New South Wales. 10, 426.

SUDGEN, W., see SMOUT, A. H.

429. sullivan, h. J. 1958. The microspore genus Simozonotriletes. 1, 125.

430. sullivan, h. j., and hibbert, a. f. 1964. Tetrapterites visensis — a new spore-bearing structure from
the Lower Carboniferous. 7, 64.

431. sullivan, H. J. 1964. Miospores from the Drybrook Sandstone and associated measures in the
Forest of Dean basin, Gloucestershire. 7, 351.

432. sullivan, h. j. 1968. A Tournaisian spore flora from the Cementstone Group of Ayrshire, Scot-
land. 15, 116.

433. sutton, i. d. 1964. The tabulate coral genus Cystihaly sites from Wenlock and Dudley. 7, 452.

TALENT, J. A., See JOHNSON, J. G.

434. tarlo, l. b. 1958. The scapula of Pliosattrus macromerus Phillips. 1, 193.

435. tarlo, L. b. 1959. Pliosaurus brachyspondylus (Owen) from the Kimeridge Clay. 1, 283.

436. tarlo, L. b. 1959. Stretosaums gen. nov., a giant Pliosaur from the Kimeridge Clay. 2, 39.

437. tarlo, L. b. 1960. The Downtonian Ostracoderm Corvaspis kingi Woodward, with notes on the
development of dermal plates in the Heterostraci. 3, 217.

438. tavener-smith, r. 1965. A new fenestrate bryozoan from the Lower Carboniferous of County
Fermanagh. 8, 478.

439. tavener-smith, r. 1966. The micrometric formula and the classification of fenestrate cryptom-
stomes. 9, 413.

440. tavener-smith, r. 1969. Skeletal structure and growth in the Fenestellidae (Bryozoa). 12, 281.

AUTHOR LIST 41

441. taylor, t. n. 1967. On the structure and phylogenetic relationships of the fern Radstockia Kid-
ston. 10, 43.

442. taylor, t. n., and eggert, d. a. 1969. On the structure and relationships of a new Pennsylvanian
species of the seed Pachytestci. 12, 382.

443. temple, J. t. 1965. The trilobite genus Oedicybele from the Kildare Limestone (Upper Ordovician)
of Eire. 8, 1.

444. tewari, b. s., and badam, g. l. 1969. A new species of fossil turtle from the Upper Siwaliks of
Pinjore, India. 12, 555.

445. thomas, b. a. 1968. A revision of the Carboniferous lycopod genus Eskdalia Kidston. 11, 439.

446. thomas, b. a. 1969. A new British Carboniferous calamite cone, Paracalamostachys spadiciformis.
12, 253.

447. thomas, h. dighton, and larwood, g. p. 1960. The Cretaceous species of Pyripora d’Orbigny
and Rhammatopora Lang. 3, 370.

448. toghill, p. 1968. The graptolite assemblages and zones of the Birkhill Shales (Lower Silurian)
at Dobb’s Linn. 11, 654.

TORRENS, H. S., See STINTON, F. C.

449. townrow, J. a. 1960. The Peltaspermaceae, a Pteridosperm family of Permian and Triassic age.
3, 333.

450. tripp, r.p. 1957. The trilobite Encriimrus multisegmentatus (Port]ock) and allied Middle and Upper
Ordovician species. 1, 60.

451. tripp, r.p. 1962. The Silurian trilobite Encrinuruspunctatus (Wahlenberg) and allied species. 5,460.

452. tripp, R. p. 1965. Trilobites from the Albany division (Ordovician) of the Girvan district, Ayr-
shire. 8, 577.

453. tucker, e. v. 1968. The atrypidine brachiopod Dayici navicula (J. de C. Sowerby). 11, 612.

454. turner, Judith. 1965. Upper Jurassic and Lower Cretaceous microfossils from the Hautes-Alpes.
8, 391.

455. valentine, j. w. 1969. Patterns of taxonomic and ecological structure of the shelf benthos during
Phanerozoic time. 12, 684.

456. van valen, l. 1965. Some European Proviverrini (Mammalia, Deltatheridia). 8, 638.

457. varker, w. J. 1967. Conodonts of the genus Apatognathus Branson and Mehl from the Yore-
dale Series of the North of England. 10, 124.

458. vella, p. 1961. Australasian Typhinae (Gastropoda) with notes on the subfamily. 4, 362.
verma, b. k., see pant, d. d.

WADE, M., see GLAESSNER, M. F.

459. wade, mary. 1969. Medusae from uppermost Precambrian or Cambrian sandstones, central
Australia. 12, 351.

460. wagner, r. h., and alvarez-ramis, c. 1967. Mciriopteris from the Stephanian of north-west
Spain. 10, 694.

waines, r. h., see wilson, e. c.

461. walker, c. a. 1966. Podocnemis somaliensis, a new pleurodiran turtle from the Middle Eocene of
Somalia. 9, 511.

462. wall, d., and downie, c. 1963. Permian hystrichospheres from Britain. 5, 770.

463. wall, d. 1967. Fossil microplankton in deep-sea cores from the Caribbean Sea. 10, 95.

464. Wallace, peigi. 1969. Specific frequency and environmental indicators in two horizons of the
Calcaire de Ferques (Upper Devonian), northern France. 12, 366.

465. walmsley, v. g. 1965. Isorthis and Salopina (Brachiopoda) in the Ludlovian of the Welsh Border-
land. 8, 454.

466. walmsley, v. g., boucot, a. j., harper, c. w., and savage, n. m. 1968. Visbyella — a new genus of
resserellid brachiopod. 11, 306.

42

AUTHOR LIST

WARREN, P. T., see HOLLAND, C. H.

467. Waterhouse, J. b. 1963. New Zealand species of the Permian bivalve Atomodesma Beyrich. 6,
699.

468. Waterhouse, J. b. 1965. Palaeotaxodont bivalves from the Permian of New Zealand. 7, 630.

469. waterston, c. d. 1960. The median abdominal appendage of the Silurian Eurypterid Slimonia
acuminata (Salter). 3, 245.

470. waterston, c. d. 1962. Pagea sturrocki gen. et sp. nov., a new eurypterid from the Old Red
Sandstone of Scotland. 5, 137.

471. watson, d. m. s. 1961. Some additions to our knowledge of antiarchs. 4, 210.

472. watson, d. m. s. 1963. On growth stages in branchiosaurs. 6, 540.

473. webby, b. d. 1962. A Middle Devonian inadunate crinoid from west Somerset, England. 4, 538.

474. webby, b. d. 1964. Devonian corals and brachiopods from the Brendon Hills, West Somerset.
7, 1.

475. webby, b. d. 1965. Quantoxocrinus, a new Devonian inadunate crinoid from West Somerset,
England. 8, 1 1.

476. webby, b. d. 1968. Astrocystites distans sp. nov., an edrioblastoid from the Ordovician of Eastern
Australia. SI, 513.

477. webby, b. d. 1969. Ordovician stromatoporoids from New South Wales. 12, 637.

478. weir, J. a. 1959. Ashgillian Trilobites from Co. Clare, Ireland. 1, 369.

479. wendt, j. 1968. Discohelix (Archaeogastropoda, Euomphalacea) as an index fossil in the Tethyan
Jurassic. 11, 554.

480. westbroek, p. 1969. The interpretation of growth and form in serial sections through brachipods,
exemplified by the trigonorhynchiid septalium. 12, 321.

WHITE, D. E., see SMITH, J. D. D.

481. white, d. e. 1966. The Silurian rugose coral Microplasma lovenianum Dybowski from Mon-
mouthshire. 9, 148.

482. white, e. i. 1958. On Cephalaspis lyelli Agassiz. 1, 99.

483. Whittington, h. b. 1958. Ontogeny of the trilobite Peltura scarabaeoides from Upper Cambrian,
Denmark. 1, 200.

484. whitworth, p. h. 1969. The Tremadoc trilobite Pseudokainella impar (Salter). 12, 406.

485. wiedmann, J. 1965. Origin, limits, and systematic position of Scaphites. 8, 397.

486. williams, a., and wright, a. d. 1961. The origin of the loop in articulate brachiopods. 4, 149.

487. williams, a. 1968. Shell structure of the billingsellacean brachiopods. 11, 486.

488. wills, l. J. 1959. The external anatomy of some Carboniferous 'scorpions’, Part 1. 1, 261.

489. wills, l. j. 1960. The external anatomy of some Carboniferous 'scorpions’, Part 2. 3, 276.

490. wills, l. j. 1963. Cyprilepas holmi Wills 1962, a pedunculate cirripede from the Upper Silurian

of Oesel, Esthonia. 6, 161.

491. wills, l. j. 1964. The ventral anatomy of the Upper Carboniferous eurypterid Anthraconectes
Meek and Worthen. 7, 474.

492. wilson, e. c., waines, r. h., and coogan, a. h. 1963. A new species of Komia Korde and the
systematic position of the genus. 6, 246.

493. wilson, r. b. 1959. Wilkingia gen. nov. to replace Allorisma for a genus of Upper Palaeozoic
lamellibranchs. 1, 401.

494. wilson, r. b. 1962. A review of the evidence for a ‘Nebraskan’ fauna in the Scottish Carboni-
ferous. 4, 507.

495. wilson, r. b. 1963. The lamellibranch genus Prothyris in the Upper Devonian and Carboniferous
of Great Britain. 6, 136.

496. wood, a. 1957. The type-species of the genus Girvanella (Calcareous Algae). 1, 22.

497. wood, A. 1963. The British Carboniferous species of Girvanella (Calcareous Algae). 6, 264.

AUTHOR LIST

43

498. wood, a. 1964. A new dasycladacean alga, Nanopora, from the Lower Carboniferous of England
and Kazakhston. 7, 181.

499. wright, a. d. 1963. The morphology of the brachiopod superfamily Triplesiacea. 5, 740. See also

WILLIAMS, A.

500. wright, c. w. 1963. Cretaceous ammonites from Bathurst Island, Northern Australia. 6, 597.

501. yates, Patricia J. 1961. New Namurian goniatites of the genus Eamorphoceras. 4, 54.

502. yates, Patricia J. 1962. The palaeontology of the Namurian rocks of Slieve Anierin, Co. Leitrim,
Eire. 5, 355.

503. zammit-maempel, g. 1969. A new species of Coelopleunts (Echinoidea) from the Miocene of
Malta. 12, 42.

504. zeuner, f. e. 1959. A new Liassic dragonfly from Gloucestershire. 1, 406.

505. zeuner, f. e. 1959. Jurassic beetles from Grahamland, Antarctica. 1, 407.

506. Ziegler, A. M. 1966. Unusual stricklandiid brachiopods from the Upper Llandovery Beds near
Presteigne, Radnorshire. 9, 346.

507. ziegler, A. M. 1966. The Silurian brachiopod Eocoelia hemisphaerica (J. de C. Sowerby) and
related species. 9, 523.

508. ziegler, A. M., cocks, l. r. m., and mckerrow, w. s. 1968. The Llandovery transgression of the
Welsh Borderland. 11, 736.

509. ziegler, b. 1963. Some Upper Jurassic ammonites of the genus Rasenia from Scotland. 5, 765.

510. zullo, v. A., kaar, F. f., Durham, J. w., and allison, E. c. 1964. The echinoid genus Salenia in the
eastern Pacific. 7, 331.

THE PALAEONTOLOGICAL ASSOCIATION

SPECIAL PAPERS IN PALAEONTOLOGY

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The following Special Papers are available. Members may obtain them at reduced
rates through the Membership Treasurer. Non-members may obtain them from
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Special Paper No. 1. (for 1967) : Miospores in the Coal Seams of the Carboniferous
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72 text-figs., 27 plates. Price £8 (U.S. $22.00), post free.

Special Paper No. 2 (for 1968): Evolution of the Shell Structure of Articulate
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Special Paper No. 10 (for 1971): Upper Cretaceous Ostracoda from the
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