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BULLETIN OF
THE BRITISH MUSEUM
(NATURAL HISTORY)
GEOLOGY
VOL. Ill
1956-1959
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1956-1959
DATES OF PUBLICATION OF THE PARTS
No. i. 1 8 April ....... 1956
No. 2. 12 October ...... 1957
No. 3. 13 December ...... 1957
No. 4. 31 January ...... 1958
No. 5. 14 February ...... 1958
No. 6. 20 February ...... 1958
No. 7. 20 February ...... 1958
No. 8. 26 August ...... 1958
No. 9. 12 September ..... 1958
No. 10. 6 February ...... 1959
PRINTED IN
GREAT BRITAIN
AT THE
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ADLARD AND SON, LTD.
CONTENTS
GEOLOGY VOLUME III
PAGE
No. i. The Structure, Evolution and Nomenclature of the Ostracod Hinge.
P. C. SYLVESTER-BRADLEY i
No. 2. Eocene Mollusca from Nigeria : A Revision. F. E. EAMES 23
No. 3. The Oligocene Flora of the Bovey Tracey Lake Basin, Devonshire.
M. E. J. CHANDLER 71
No. 4. The Structure of some Leaves and Fructifications of the Glossopteris
Flora of Tanganyika. D. D. PANT 125
No. 5. Lidgettonia, a new type of fertile Glossopteris. H. HAMSHAW THOMAS 177
No. 6. The Faunal Succession in the Caradoc Series of South Shropshire.
W. T. DEAN 191
No. 7. A new Labyrinthodont (Pamcyclotosaurus] from the Upper Trias of
New South Wales. D. M. S. WATSON 233
No. 8. An Early Pleistocene Mammalian Fauna from Bethlehem. D. A.
HOOIJER 265
No. 9. The Upper Permian Flora of England. H. M. M. STONELEY 293
No. 10. Blue-Green Algae from the Middle Devonian of Rhynie, Aberdeenshire.
W. N. CROFT AND E. A. GEORGE 339
Index to Volume 3 355
THE STRUCTURE, EVOLUTION
AND NOMENCLATURE OF THE
OSTRACOD HINGE
P. C. SYLVESTER-BRADLEY
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. i
LONDON: 1956
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following papers appeared in Volume I (i949~52) :
Pnce
No. i (1949). The Pterobranch Rhabdopleum in the English Eocene.
H. D. Thomas & A. G. Davis . • 75- ^d-
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu . 5*-
No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White.
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White . 7*. «•
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
Northern Iraq. L. F. Spath . . 1O5-
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum. T. H. Withers . . 5«-
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .
T. H. Withers ... 55-
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian
of Podolia. W. N. Croft . . IOS-
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsdale. .
No. 9 (1952). Australian Arthrodires. E. I. White *5S-
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard
The following papers appeared in Volume II (i953-I956) :
No. i (1953). The Deer of the Webourn Crag and Forest Bed of Norfolk.
A. Azzaroli ... • ' ' r '
No. 2 (1953). A Carboniferous Petrified Forest in Patagonia. Mary
Gordon Calder ....
No. 3 (1953). The Solution of the Piltdown Problem. By J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark • 3*. &•
No. 4 (1954). Some Upper Cretaceous & Eocene Fruits from Egypt.
Marjorie E. J. Chandler .
No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans . I5«-
No 6 (1955). Further contributions to the Solution of the Piltdown
Problem. J. S. WTeiner, W. E. Le Gros Clark, K. P. Oakley &
others ... • • • i1
No. 7 (1955). The Schizaeaceae of the South of England in early Tertiary
Times. Marjorie E. J. Chandler . I5$-
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson.
THE STRUCTURE, EVOLUTION AND
NOMENCLATURE OF THE OSTRACOD HINGE
BY
PETER C. SYLVESTER-BRADLEY
University of Sheffield
Pp. 1-21; Pis. 1-4 ; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. i
LONDON : 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. i of the Geological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued April, 1956 Price Twelve Shillings and Sixpence
THE STRUCTURE, EVOLUTION AND
NOMENCLATURE OF THE OSTRACOD HINGE
By P. C. SYLVESTER-BRADLEY
[University of Sheffield]
SYNOPSIS
A revision of Middle Jurassic ostracods from the Fullers' Earth Clay has revealed a series
of hinge-types which clearly demonstrate transitional stages between those commonly known
as merodont and those known as amphidont. Their study has justified the classification of
hinge-types according to the stage reached in a morphogenetic series. The primitive hinge is
tripartite in character, and more specialized hinge-types originate by the subdivision of the
median element of the basic three. The morphology of the ostracod hinge is discussed, a
rational classification is proposed, and in the light of this classification, the terminology of both
the hinge as a whole and that of its constituent parts is reviewed. The evolution of the post-
Palaeozoic Cytheracea is traced from Palaeozoic ancestors with a similar hinge but different
muscle-scar pattern, and a new superfamily, the Quasillitacea, is proposed for these Palaeozoic
forms. Two new genera are proposed, Acanthocythere (Middle Jurassic) and Dictyocythere
(Upper Jurassic). The latter is divided into two subgenera, Dictyocythere sensu stricto and
Rhysocy there nov. The distribution of the species of Dictyocythere in the Upper Jurassic of England
and northwest Germany suggests that the " Purbeck " Beds of the Aylesbury and Swindon
districts are earlier than Middle Purbeckian, and that the " Wealden 2 " of the German
sequence is probably equivalent to the upper part of the Middle Purbeck Beds of Dorset. The
development of the hinge in Dictyocythere is shown to be palingenetic.
i. TERMINOLOGY OF THE OSTRACOD HINGE
THE hinge structures of fossil ostracods play an important part in their taxonomic
determination, but unfortunately the terms used for their description are ambiguous.
Thus the terms " taxodont " and " heterodont " have been used to describe the
compound hinge of ostracoda, not always with quite the same meaning. As used by
Bold (1946), Kingma (1948), Grekoff (1952) and Malkin (I953),1 " taxodont "
refers to the presence of denticulate elements, and " heterodont " to the development
of high, pointed teeth and a hinge-bar. The terms (which have been borrowed from
those used to describe the lamellibranch hinge) are not altogether appropriate in their
new context, and seem likely to lead to ambiguity. The term " taxodont " as used
in lamellibranchs refers to the occurrence of an alternate tooth and socket arrange-
ment, essentially similar in both valves. An exactly similar structure is not known
to occur in the Ostracoda, although an alternate tooth and socket arrangement
(here termed " interdentate ") is not infrequently found in some or all of the elements
1 Malkin (1953) also defines the terms adont, dysodont, desmodont, crasidont and archidont. These
terms are not referred to further in this paper, as they do not describe hinge structures here dealt with.
GEOL. Ill, I. I
4 STRUCTURE, ETC. OF THE OSTRACOD HINGE
of the compound hinge. This arrangement might well be called " taxodont " and the
term has in fact been so used by Kesling (1951) ; this use of the term is not, however,
equivalent to that of the earlier definitions, and it would seem advisable to abandon
it to prevent further confusion.
Other authors have used different terms descriptive of the same structures ;
thus " merodont " = " taxodont ", and " amphidont " =" heterodont " (e.g. Triebel
1950 ; Pokorny, 1952). These terms are not open to the same objections, and are
adopted here and fully defined below. Additional terms are also introduced ; some
of these are new, some are after Triebel. Technical terms, in standard English, or
anglicized from Latin roots, are used to designate structures of the individual
elements ; terms derived from the Greek and terminating in " -dont " are used to
designate the hinge structure as a whole.
2. DEFINITIONS OF HINGE ELEMENTS
The hinge structure of the ostracod carapace may be simple or compound.
The simple hinge (" adont " of Grekoff, Bold and Kingma) may be thought of as
made up of a single element, which consists of a groove along the margin into which
fits the edge of the other valve (Text-fig, i, A). The hinge is not further differentiated.
(/anterior
median
posterior
—''anterior
"-xanterbmedkfri
posteromedian
posterior
A B
TEXT-FIG, i. — Diagrams of left valves of three ostracods, dorsal view, to illustrate division of
hinge into "elements". (Stippled areas represent sockets).
The compound hinge is divided into three or four elements in each valve. Primi-
tively there are three elements, anterior, median and posterior (Text-fig, i, B).
Usually these elements alternate so that if (as is usual in the right valve) the anterior
and posterior are ridges, the median element separating them will be a groove ; or
if (as is usual in the left valve) the terminal elements are grooves or sockets, the median
will be a ridge. Though genera are known which are exceptions to this (in Haplo-
cytheridea, for example, all three elements of the left valve are grooves, those of the
right being ridges), the distinction between the constituent elements remains clear,
the terminal elements being more coarsely dentate than the median.
It will be shown below that the more primitive compound hinge with three elements
STRUCTURE, ETC. OF THE OSTRACOD HINGE 5
evolves, by subdivision of the median element, into a more advanced type in which
there are four elements — anterior, anteromedian, posteromedian and posterior.
The division into two parts of the median element is at first not pronounced — the
anterior part being merely more coarsely dentate than the posterior (Text-fig. 2, c),
but later developed forms have a more clear-cut distinction, the anteromedian
becoming rather similar in proportions to the anterior element, the posteromedian
usually being much the longest element of the four (Text-fig, i, c) . Rarely the anterior
element also becomes differentiated, with denticles of two sizes (e.g. Amphicy there
and Dictyocythere ; see p. 14).
These hinge elements themselves display a great deal of variation, and the following
definitions are of terms used in their description :
A. Ridge and groove.
i. A hinge element may consist of a smooth ridge fitting into a corresponding
groove in the opposite valve, e.g. the median element of Camptocythere
(Text-fig. 2, A).
A. Lophodont
B. Merodont
C. Entomodont
D. Lobodont
E. Schizodont
F Amphidont
TEXT-FIG. 2. Diagrams illustrating the six main types of dentition developed in the ostracod
hinge. Each pair of valves in dorsal view. (Stippled areas represent sockets).
6 STRUCTURE, ETC. OF THE OSTRACOD HINGE
2. A dentate element consists of a ridge divided into projections (denticles)
which fit into a socketed groove termed loculate.
3. The median element may be furnished with finer denticles, when it is
termed denticulate, the groove locellate (Text-fig. 2, B).
4. A ridge may be divided into separate teeth between which lie alternating
sockets for the reception of projections from the complementary groove.
Such a ridge is termed interdentate, the groove interloculate.
5. The median element of the left valve (which is usually a ridge or a bar)
often lies under an elongated, sub-triangular, excavated area termed the
accommodation groove. This, it would seem, serves to receive the project-
ing dorsal edge of the right valve when the carapace is open.
B. Bar.
6. The median element may consist of a ridge which is raised up from the shell
margin behind it ; this is differentiated as a hinge-bar.
C. Boss and pit.
7. An element may be raised up as a hemispherical boss, which may show
division into two or more lobes, when it is referred to as bi-, tri-, or multi-
lobate ; a lobate boss fits into a loculate pit in the complementary valve
(Text-fig. 2, D).
D. Tooth and socket.
8. An element may consist of a single tooth, projecting from a platform which
itself projects more or less beyond the hinge margin, when it is referred to
as stirpate (e.g. the anterior element in Trachyleberis] (Text-fig. 2, F).
9. The tooth may be a simple, more or less conical projection ;
10. or it may have more or less parallel sides, when it is called pessular ;
11. or it may be split into two lobes, when it may be called bifid (Text-fig. 2, E) ;
12. or it may be crescentic, half-surrounding a socket ;
13. or it may be a slightly elongated and slightly curved projection, when it is
termed reniform (as in the posterior element of many amphidont hinges
(Text-fig. 2, F). A reniform tooth may be smooth (Hemicythere or Trachy-
leberis} or lobate (Bradley a] or dentate (some species of Cythereis}.
3. CLASSIFICATION OF HINGE TYPES
Using the above definitions of the elements of the compound hinge as a basis, it is
possible to classify hinges developed in the Cytheracea in the six main groups outlined
below. This classification does not take into consideration hinge-types developed in
other superfamilies of the Podocopa. For example, whereas the Quasillitacea (here
defined) have hinges very like the Cytheracea (though apparently confined to the more
primitive lophodont and merodont types), the Bairdiacea have a more complicated
compound hinge not always homologous with that in the Cytheracea — compare, for
example, Bairdoppilata (Coryell, Sample & Jennings, 1935) with Macrocypris
(Sylvester-Bradley, 1948^) and Triebelina (lophodont ; Triebel, 1948). The Cypridacea,
STRUCTURE, ETC. OF THE OSTRACOD HINGE 7
on the other hand, have mostly simple hinges with a single element, though a few
are lophodont (e.g. Cypridea ; Sylvester-Bradley, 1949).
i. Lophodont: e.g. Bythocythere, Camptocythere, in which the hinge elements are
divided into three in each valve, all consisting of ridges and grooves (Text-fig. 2, A).
The most usual arrangement is as follows :
Left valve Right valve
Anterior element . . Groove . Ridge.
Median element . . Ridge . Groove.
Posterior element . . Groove . Ridge.
2. Merodont: ( = " taxodont " of some authors) e.g. Cythere, in which anterior
and posterior elements are dentate or loculate, the median element either smooth or
denticulate, but not subdivided (Text-fig. 2, B). The most usual arrangement is as
follows :
Left valve Right valve
Anterior element . . Loculate groove . Dentate ridge.
Median element . . Denticulate ridge . Locellate groove.
Posterior element . . Loculate groove . Dentate ridge.
The elements in some merodont genera (e.g. the terminal elements of Pleurocythere)
are interdentate and interloculate rather than dentate and loculate. Many species of
Middle Jurassic age have a merodont hinge with smooth median element (e.g.
Schuleridea) , and this condition may well be the more primitive.
3. Entomodont: e.g. Lophocy 'there, Progonocythere and Xenocythere (Triebel,
1949, 1951), in which the median element becomes subdivided, the anterior part
remaining dentate, the posterior smooth or denticulate (Text-fig. 2, c). A common
arrangement is as follows :
Left valve Right valve
Anterior element . . Loculate groove . Dentate ridge.
Anteromedian element . Short dentate ridge • . Short, wide loculate groove.
Poster omedian element . Long denticulate ridge . Long, narrow locellate groove.
Posterior element . . Loculate groove . Dentate ridge.
The genus Macrodentina has a hinge transitional between entomodont and lobodont
(seep. 14).
4. Lobodont: e.g. Acanthocythere gen. nov., in which the anterior element and the
anteromedian elements are lobed bosses (see PL i, figs. 3, 4, 8, 9 ; Text-fig. 2, D).
Left valve Right valve
Anterior element . . Loculate pit . Lobate boss.
Anteromedian element . Lobate boss . Loculate pit.
Posteromedian element . Smooth or denticulate bar . Smooth or locellate groove.
Posterior element . . Loculate groove . Dentate ridge.
8 STRUCTURE, ETC. OF THE OSTRACOD HINGE
The genus Amphicythere has a dentition transitional between lobodont and amphi-
dont. The anterior element is lobate, but the antero-median element is almost entire
(see p. 14).
5. Schizodont: e.g. Schizocy there, Palmenella, Paijenborchella (Triebel, 1950), in
which teeth of the anterior and anteromedian elements are bifid (Text-fig. 2, E).
Left valve Right valve
Anterior element . . Biloculate socket . Bifid, stirpate tooth.
Anteromedian element . Bifid tooth . Biloculate socket.
Posteromedian element . Denticulate bar . Locellate groove.
Posterior element . . Loculate socket . Lobate, reniform tooth.
6. Amphidont: ( = " heterodont " of some authors ; Triebel uses the term
amphidont in a wider sense than that proposed here — to include dentitions of both
schizodont and lobodont types) e.g. Dictyocythere, Trachyleberis , in which the median
element is further differentiated into a single anterior tooth and a posterior bar or
ridge. A common arrangement (Text-fig. 2, F) is as follows :
Left valve Right valve
Anterior element . . Socket . Stirpate tooth.
Anteromedian element . Conical or pessular tooth . Socket.
Posteromedian element . Smooth or denticulate bar . Smooth or locellate groove.
Posterior element . . Loculate socket . Lobate, reniform tooth.
Young moults of amphidont species usually have merodont or entomodont hinges
(see p. 19). The ontogeny is therefore recapitulatory .
This classification is not comprehensive. The hinge structures listed form a useful
morphogenetic series, but complications are developed in some genera which are not
covered — for example, the differentiations of the median element found in some
genera of the Cytherideinae (e.g. Cytheridea, Cyprideis), and the development of
crescentic teeth in the terminal elements in Loxoconcha and other genera.
4. THE EVOLUTION OF THE HINGE AND ITS BEARING ON
THE CLASSIFICATION OF THE PODOCOPA
Well-defined compound hinge-structures are particularly characteristic of the
superfamily Cytheracea, which in post-Palaeozoic times provided a far greater number
of species of marine ostracods than all the remaining superfamilies taken together.
The series of hinge-types classified above is a morphogenetic series and not phylo-
genetic. Genera have passed through the various stages at different times during
their racial history. The most advanced stage reached by Middle Jurassic times is the
lobodont, but the amphidont hinge appears before the close of the Upper Jurassic
(e.g. Dictyocythere, see p. 14) . It is not unlikely that earlier genera with an amphidont
hinge remain to be discovered. In Recent and Tertiary ostracods, merodont and
amphidont hinge-structures predominate but entomodont hinges also occur (e.g.
Leptocythere) ; in Middle Jurassic times the Cytheracea have hinges which are pre-
STRUCTURE, ETC. OF THE OSTRACOD HINGE 9
dominantly either merodont (usually with a smooth median element) or entomodont ;
lobodont hinges are rare. Elsewhere (Sylvester-Bradley, 19486) attention has been
drawn to the series leading from the Middle Jurassic Oligocythereis (entomodont)
through the Cretaceous Cythereis to the Tertiary and Recent Trachyleberis (amphi-
dont). This series is thought to be broadly phylogenetic, and the three genera in
question are placed in the same subfamily, the Trachyleberidinae. Several other
undescribed genera from the Middle Jurassic have reached various stages in similar
series. It seems unlikely that any of these forms are directly related to the Trachyle-
beridinae ; they probably represent other lineages undergoing parallel evolution.
The acquisition of the amphidont hinge is considered to be a well-marked trend. If
this view is correct, the diagnosis of the subfamily Progonocytherinae (Sylvester-
Bradley, 19480 : 189) for Cytheracea with entomodont hinge, needs drastic revision,
and it is now suggested that Lophocythere, Progonocy there and Oligocythereis should not
be grouped together in one subfamily, although they all have entomodont hinges.
Before attempting to assess which of the hinge-types described can truly be taken
as primitive, it is necessary to refer to older faunas and seek possible Palaeozoic
ancestors of the Cytheracea.
The genus Monoceratina is the longest lived of the Cytheracea according to present
ideas. The type species is Carboniferous in age, and Recent species have been
recorded (Stephenson, 1946 ; Bold, 1946). However, the hinge structure is rather
obscure ; the shape differs much from the majority of the Cytheracea ; the muscle-
scar patterns differ sufficiently in different species to warrant query as to whether
there has not been a mistake in the identification of some of the post-Palaeozoic
material, and it is not in any case easy to postulate that Monoceratina exhibits an
ancestral type of hinge-structure.
Perhaps the Palaeozoic family that most invites comparison with the Cytheracea
is the Middle Devonian Ropolenellidae first described by Coryell & Malkin (1936),
especially the two genera Ropolenellus and Euglyphella. By the courtesy of Mr.
Raymond R. Hibbard, who presented me with several samples rich in ostracods,
which he collected from the Middle Devonian of New York State, I am able to describe
the more intimate details of the shell of some species of Euglyphella. Some of the
earlier authors who dealt with the Ropolenellidae nave oriented the ostracods so that
the higher end is posterior. The investigation of the hinge reveals that they are closely
analogous, if not homologous, with those found in the Cytheracea ; the higher end
is therefore here regarded as anterior in conformity with the known orientation of
the Cytheracea. Consequently the terms " left " and " right " used here have the
reverse meaning to those used in description by some previous authors.
The shape of these two genera of the Ropolenellidae, in marked contrast to many
other described species of Palaeozoic ostracods, is similar to many genera of the
Cytheracea (PI. 2, figs. 5-7). A broadly rounded anterior tapers to a narrow, rounded
or triangular posterior. Either or both ends may be spinose. The hinge-line is straight,
delimited by well-marked anterior and posterior cardinal angles and is shorter than
the length of the shell. The ventral border is straight or slightly concave.
The genus Euglyphella was proposed by Warthin (1934) with Strepula sigmoidalis
Jones as type species. Several other species have since been assigned to the genus.
GEOL. Ill, I. 2
io STRUCTURE, ETC. OF THE OSTRACOD HINGE
It possesses a striking ornament of carinae not unlike those developed in many post-
Palaeozoic Cytheracea (e.g. Lophocythere , from the Bathonian ; see Sylvester-
Bradley, 19480). The hinge in Euglyphella is divided into three elements as follows :
Left valve Right valve
Anterior .... Rather long, deeply over- . Projecting ridge, forming a
hung, loculate groove continuation of the selvage.
Median .... Bar, lying below a long, . Long groove.
narrow accommodation
groove
Posterior. . . . Short, curved loculate groove . Short, curved projecting ridge.
The hinge margin of this genus is in fact almost exactly similar to that of many
post-Palaeozoic Cytheracea. The loculate nature of the terminal elements in the
left valve cannot be made out in more than a few specimens, and I have not detected
any specimen showing a corresponding dentation on the terminal elements of the
right valve, but this may well be due to indifferent preservation.
The free margin is also similar to that seen in many Cytheracea. There is a wide
duplicature, no vestibule, but a pronounced selvage. The muscle-scar pattern,
however, is quite unlike anything known in the post-Palaeozoic Cytheracea ; a
central, circular muscle-scar pit bears a cluster of small oval scars in the form of
slight tubercles, which are in close juxtaposition.
Another genus referred by Coryell & Malkin (1936) to the Ropolenellidae is Bufma
(PI. 2, figs. 3, 4). Species of this genus differ somewhat in shape from Euglyphella
and Ropolenellus in that the dorsal margin of the left valve is markedly curved. The
characteristic ornament of the genus is unlike that of any post-Palaeozoic ostracod
known to the author, though Henningsmoen (1953) has pointed out that it is shared
by other Palaeozoic genera such as Ponderodictya (see below). The details of hinge,
duplicature and muscle-scar, however, are as described for Euglyphella, and though
Coryell & Malkin were clearly correct in concluding that the genera are related,
Henningsmoen is probably right in removing Bufina to the Quasillitinae.
The Ropolenellidae are apparently only known from Middle Devonian strata, in
which they are often fairly common. They seem to represent a successful though
short-lived line of evolution which in many respects anticipated the much later
Cytheracea.
Another family well represented in the Middle Devonian of North America is the
Quasillitidae. An examination of Quasillites (PI. 2, figs, i, 2) shows that it has a
hinge-structure very similar to that of the Ropolenellidae and the two families should
almost certainly be grouped together. Henningsmoen (1953) regarded them as
subfamilies of one family ; it is here suggested that they are related families in the
same superfamily, the Quasillitacea (see p. n). Near to Quasillites is the Carboni-
ferous genus Graphiadactylus.
It may also be mentioned that the genus Ponderodictya (described by Coryell &
Malkin from the Middle Devonian and placed by them in the Cytherellidae ; see figs,
in Triebel, 1954) bears a hinge comparable to that of the Ropolenellidae, differing
mainly in that the anterior element is longer in proportion (as long, in fact, as the
STRUCTURE, ETC. OF THE OSTRACOD HINGE n
median element). The ornament is not unlike that developed in Bufina. The genus
has been placed in the Healdiidae as a member of the Platycopa (Henningsmoen,
1953 ; Triebel, 1954), but the hinge makes this assignment a little doubtful. Pondero-
diclya may perhaps be a member of the Quasillitacea, and as such one of the Podocopa.
It is clear that in Devonian times the Healdiidae, Quasillitidae and Bairdiidae
possessed characters in common which suggest that all three families had been
derived from a common ancestor of not much greater antiquity than the Devonian.
Subsequently the three families diverged in morphology so that they have been
placed far apart in classification.
From the foregoing brief survey it will be seen that several families of Palaeozoic
ostracods possessed features in common with the later Cytheracea. It seems probable
that the first members of the Cytheracea were derived from one or more of the Quasil-
litacea towards the close of Palaeozoic time. Somewhat similar views have been
advanced by Kellett (1943). Thus the superfamilies Quasillitacea and Cytheracea
form part of the suborder Podocopa, and are distinguished by the muscle-scar
pattern, which in the Quasillitacea is aggregate (i.e. consisting of a group of individual
scars crowded in close juxtaposition ; PI. 2, figs. 3, 4, 6), and in the Cytheracea
discrete (i.e. consisting of a series of individual scars separated from each other).
The primitive hinge was therefore lophodont. Whether the elements were primi-
tively dentate or smooth cannot be certainly stated, for the apparently smooth
elements in much Palaeozoic material may well be due to a loss of finer structure on
recrystallization. The fact that loculate grooves have been detected in the terminal
elements of both Euglyphella and Quasillites shows that at least the terminal elements
were dentate in these genera by Devonian times. Some Palaeozoic specimens show a
well-developed accommodation groove, so that structure cannot be taken as an
advanced feature. It. seems to be the inevitable consequence of any considerable
difference in size between the two valves.
5. SYSTEMATIC DESCRIPTIONS
Suborder PODOCOPA
Superfamily QUASILLITACEA (Coryell & Malkin).
(First introduced here as a superfamily)
TYPE GENUS. Quasillites Coryell & Malkin. (Middle Devonian.)
DIAGNOSIS. Podocopa with a primitively tripartite hinge and an aggregate
muscle-scar pattern.
FAMILIES INCLUDED. Quasillitidae (including Graphiadactyllidae) and Ropolenel-
lidae. Other families no doubt remain to be described.
REMARKS. Distinguished from Cytheracea by muscle-scar pattern ; from
Cypridacea and Bairdiacea by hinge and muscle-scar pattern ; from Healdiidae
(Platycopa) by hinge.
DISTRIBUTION. So far known only from the Palaeozoic. Mesozoic Platycopa with
an aggregate muscle-scar pattern are, however, known (Ogmoconcha, Lower Lias)
GEOL. Ill, I. 2§
12 STRUCTURE, ETC. OF THE OSTRACOD HINGE
and it may be expected that the Podocopa underwent a parallel development ;
Mesozoic Quasillitacea may yet be discovered.
Superfamily CYTHERACEA
Family uncertain
Genus ACANTHOCYTHERE nov.
TYPE SPECIES. Cythere sphaerulata Jones & Sherborn, 1888 : 253, pi. i, fig. 6.
DERIVATION OF NAME. anavBa, a thorn or prickle + genus Cythere. Gender:
feminine.
DIAGNOSIS. Cytheracea with lobodont hinge, surface more or less spiny, carapace
plump, eye tubercles shiny, rather prominent.
OCCURRENCE. The two species of the genus here described are so far known only
from the Upper Fullers' Earth Clay (Bathonian) of the Bath district.
Acanthocy there sphaerulata (Jones & Sherborn)
(Plate i, figs. 1-4)
1888. Cythere sphaerulata Jones & Sherborn, p. 253, pi. i, fig. 6.
DIAGNOSIS. Carapace oblong, tumid, ornamented with closely-set blunt spines
arranged in a faint reticulate pattern.
HOLOTYPE. Geol. Dept. Brit. Mus. (N.H.) no. 1.1835. A complete carapace.
TYPE LOCALITY. " Yellow Fullers-earth Clay, Midford " (Jones & Sherborn,
1888 :• 254).
OTHER FIGURED SPECIMENS. Geol. Dept. Brit. Mus. (N.H.) nos. 1^.42433-34.
DESCRIPTION. Both valves with sub-parallel dorsal and ventral margins, no
taper, anterior and posterior ends evenly rounded. Sexual dimorphism rather pro-
nounced, the presumed males longer than females.
Dimensions (mm.) Proportions
L H W L H W
Carapace <J (1.1835 ; holotype) . . 0-51 0-31 0-32 . 1-65 : i : 1-03
Left valve $ (In. 42434) . . . 0-67 0-37 0-26 . 1-81:1:0-70
Right valve <J (In. 42433) . . . 0-69 0-38 0-22 . 1-82:1:0-58
The whole surface (with the exception of the eye tubercles) covered with very
fine spines set close together and forming a reticulate pattern which is clearer in
worn specimens than in those more perfectly preserved.
Normal pore canals large, sparse, about 20 to the valve in the female. Radial
pore canals simple, sparse, straight, about 8 to the anterior end. Duplicature fairly
wide at anterior end (PI. i, fig. 3), with flange-groove well developed in left valve.
Muscle-scar not seen. Hinge lobodont, in detail as follows :
STRUCTURE, ETC. OF THE OSTRACOD HINGE
Hinge element
Anterior
A nteromedian
Posteromedian
Posterior
Left valve
Right valve
Circular pit, presumably . 4-lobate boss
loculate
4-lobate boss
Smooth bar lying below ac-
commodation groove which
is rather wide in the female,
but narrow, and developed
only in posterior half in
male
Loculate groove
Oval pit, presumably loculate.
Narrow groove lying below
rather prominent dorsal mar-
gin which forms a ridge
complementary to the accom-
modation groove.
Prominent, slightly curved 5 to
7 dentate ridge, set at a slight
angle to the median element.
The state of preservation is not sufficiently perfect to make it quite certain that the
posteromedian element is not denticulate.
MATERIAL. Fourteen specimens (in addition to the holotype) from the same
horizon and locality as A . spiniscutulata.
Acanthocy there spiniscutulata n. sp.
(Plate i, figs. 5-9)
DERIVATION OF NAME. Latin spina, spine -f- scutulatus, diamond-shaped.
DIAGNOSIS. Carapace tapering, with prominent cardinal angles ; ornament of
subtriangular or diamond-shaped spinose ridges.
HOLOTYPE. Geol. Dept. Brit. Mus. (N.H.) no. 111.42435. A left valve.
TYPE LOCALITY. Upper Fullers'-Earth Clay (in top foot of economic " Fullers
Earth "), Fosse Way Fullers'-Earth Mine, near Bath. Nat. Grid, ref 31/727613.
Author's field ref. no. 47 FW g.
PARATYPES. Same locality and horizon. 1^42436-37.
DESCRIPTION. Carapace tapering, in side view, towards posterior. Ornament of
subtriangular or diamond-shaped rather coarsely spinose ridges. Sexual dimorphism
not observed.
Dimensions (mm.) Proportions
Carapace (In. 42437)
Left valve (In. 42435 ; holotype)
Right valve (In. 42436)
o-57
o-57
0-56
H
0-40
0-32
W
o-35
O-22
0-IQ
i-43
1-54
1-75
H W
i : 0-88
i : 0-59
i : 0-59
Details of hinge, duplicature and pore canals as in A. sphaendata.
MATERIAL. Nine specimens from the type-locality and horizon.
Genus DICTYOCYTHERE nov.
TYPE SPECIES. Cythere retimgata Jones, 1885 : 350, pi. 9, figs. 21-24.
DERIVATION OF NAME. SIKTVOV, a net -f genus Cythere. Gender : feminine.
DIAGNOSIS. More or less trapezoidal Cytheracea, usually reticulate, with amphi-
I4 STRUCTURE, ETC. OF THE OSTRACOD HINGE
dont hinge, the anterior element undifferentiated (subgenus Dictyocy there) or
differentiated into denticles of two sizes (subgenus Rhysocy there], the posterior
element dentate. No accommodation groove. Eye tubercles not developed.
OCCURRENCE. This genus includes several species from the Purbeck Beds of
Buckinghamshire and Swindon, the Middle Purbeck Beds of Dorset and the so-called
Wealden Beds of N.W. Germany. D. transiens is abundant in the Upper Portland
Beds of the Aylesbury district.
REMARKS. Species of this genus were placed by Martin (1940) in his genus
Macrodentina. However, the type-species of Macrodentina (M. lineata] is Upper
Oxfordian ; Triebel (1954) has shown that two homeomorphs were confused by
earlier authors under the name M. lineata, one of which (the true M, lineata] has an
advanced type of entomodont hinge, verging on the lobodont. The Kimmeridgian
genus Amphicy 'there, described by Triebel in the same paper, has an interesting hinge
transitional between lobodont and amphidont. In it the anterior element of the hinge
has denticles differentiated into two series. This rather rare variation is also found in
Macrodentina and in some species of Dictyocy there, which suggests that the three
genera may be related.1 The hinge of Dictyocythere resembles that of Amphicy there
but has moved nearer to the typical amphidont type, showing its primitive nature
only in those species possessing a differentiated anterior element (subgenus Rhyso-
cythere]. The details of the hinge of Amphicythere semisulcata, derived from Triebel's
(1954) description, are tabulated below for comparison with those of Dictyocythere.
Left valve Right valve
Anterior part of anterior element . Loculate groove . Low tridentate ridge.
Posterior part of anterior element Loculate pit . Trilobate boss.
Anteromedian element . . Faintly bilobate boss . Deep pit.
Posteromedian element . . Smooth ridge . Shallow groove.
Posterior element . . . Loculate groove . Slightly curved, y-dentate
ridge.
SUBGENERA. The fact that two hinge-types exist in the group of species here
regarded as forming Dictyocythere might be regarded as evidence that they should
be split up into two distinct genera. In other characters, however, D. rugulata seems
to be so close to D. retirugata that I prefer to separate the groups as subgenera only,
as follows :
(i) Subgenus Dictyocythere sensu stricto (with undifferentiated anterior element to
hinge] :
D. retirugata (Jones).
D. mediostricta n. sp.
D. decor ata (Anderson).
(ii) Subgenus Rhysocythere (with differentiated anterior element]:
D. rugulata (Jones).
D. transiens (Jones).
1 However, fairly prominent eye tubercles are developed in Amphicythere, but not in the other two
genera.
STRUCTURE ,ETC. OF THE OSTRACOD HINGE 15
STRATIGRAPHICAL VALUE. The distinction, made below, between species of the
genus found in the Purbeck Beds of Buckinghamshire and in the Swindon Series on
the one hand (D. (R.) rugulata, D. retirugata), and in the Middle Purbeck Beds of the
Dorset coast on the other (D. mediostricta) , lends further support to the supposition
that the Swindon Series (and the Aylesbury " Purbeck " Beds) were earlier than the
Middle Purbeck Beds of the Dorset coast (see Arkell & Sylvester-Bradley, 1941). It
appears that no species of Dictyocythere or of Cypridea is identical in the two regions1.
It is D. mediostricta that is found in the so-called Wealden of N.W. Germany, not
D. retirugata, as recorded by Martin. The abundance of D. transiens in undoubted
Portland Beds at Aylesbury (below horizons yielding Titanites) shows that the genus
was certainly in existence before the close of Portlandian time, and it still seems
possible that Blake (1885) may have been right when he suggested that the so-called
Purbeck Beds of Swindon and Aylesbury were deposited at the same time as the Port-
land Beds further south. D. (R.} rugulata and D. retirugata have been both recorded
from the Swindon Sands and Stone (Sylvester-Bradley, 1941 : 358), which has always
been regarded as Portlandian.
Subgenus DICTYOCYTHERE sensu stricto
DIAGNOSIS. Dictyocythere with undifferentiated anterior element to hinge.
Dictyocythere (Dictyocythere) retirugata (Jones)
(Plate 3, figs. 7-10 ; PI. 4, figs. 3, 4, n, 16, 17)
1885. Cythere retirugata Jones, p. 350, pi. 9, figs. 21-24 (including var. textilis).
[not Macrodentina retirugata (Jones) Martin, 1940.]
1941. Cythere retirugata Jones var. textilis Jones : Anderson, p. 374 (part), pi. 18, fig. 3, ? fig. 2.
DIAGNOSIS. Surface reticulate. Shape trapezoidal. Median constriction absent or
slight. Sexual dimorphism pronounced.
LECTOTYPE (here designated). Geol. Dept. Brit. Mus. (N.H.) no. In.486oi. Figured
Jones, 1885, pi. 9, fig. 23. (Jones' no. 253, n.) A right valve J2.
TYPE LOCALITY. " Last Portland bed ", Hartwell, Bucks.
DESCRIPTION. Surface evenly reticulate except immediately above muscle scar.
Shape trapezoidal, with pronounced cardinal angles. Sexual dimorphism pronounced,
the presumed males proportionately longer than the females.
1 Wolburg (1950) suggests that Ulwellia papulata Anderson from the Swindon Series may be synony-
mous with Cypridea sowerbyi Martin from N.W. Germany, but specimens of the latter species that Dr.
Wolburg kindly sent me show that this is not the case. On the other hand the specimen figured by
Anderson (1941, pi. 18, fig. 2) from the Middle Purbeck of Poxwell, Dorset, does appear to have been
correctly identified as Cythere retirugata.
2 Lectotype of " Cythere retirugata var. textilis Jones " (here designated) : Geol. Dept. Brit. Mus.
(N.H.) no. In. 48602, figured Jones, 1885, pi. 9, fig. 24. A right valve $. (Jones' no. 166, i). " Shaly
beds, Barnard's Pit, Hartwell ".
16 STRUCTURE, ETC. OF THE OSTRACOD HINGE
Dimensions (mm.) Proportions
L H W L H W
Left valve $ (In. 48609) . . . 0-92 0-47 0-25 . 1-97 : i : 0-53
Right valve $ (In. 48608) . . . 0-97 0-49 0-29 . 1-98:1:0-59
Left valve $ (In. 48610) . . . 0-80 0-51 0-28 . 1-58:1:0-55
Right valve $ (In. 48611) . . . 0-84 0-51 0-24 . 1-66:1:0-47
Normal pore canals large ; radial pore canals sparse, straight. Muscle-scar pattern
of usual Cytheracea type, with four scars in vertical superposition, and a single scar
in front. Hinge amphidont (see PI. 4. figs. 3, 4, n, 16, 17) details as follows :
Hinge element Left valve Right valve
Anterior . . Pit . Smooth protuberant boss with a
slight anterior swelling, perhaps
representing the relict of ant-
erior denticulation.
Anteromedian . . Smooth protuberant boss . Pit.
Posteromedian . Smooth, slightly projecting bar. . Smooth groove.
No accommodation groove
Posterior . . Loculate groove . Curved, dentate ridge.
REMARKS. Jones (1885) and Anderson (1941) regarded this species as composed
of a number of varieties based on differences in the strength and complexity of the
reticulate ornament. Two of these varieties are here raised to specific rank, since the
different ornament is correlated with slight differences in shape and with (in one case)
a difference in the hinge. Moreover there are no transitional forms, and though they
occur together in the same bed at Swindon, they have a different distribution at
other horizons and in other localities. I have been unable, however, to maintain a
distinction between the typical D. retirugata and the forms referred by Jones and
Anderson to var. textilis.
OCCURRENCE. This species is abundant in the Cythere Marl of the Swindon Series
at Swindon, and occurs also at higher and lower horizons. It is found at several
horizons and localities in the Purbeck Beds of Buckinghamshire.
Dictyocythere (Dictyocythere) mediostricta n.sp.
(Plate 3, figs. 2-6)
1940. Macrodentina retirugata (Jones) : Martin, p. 330, pi. 5, figs. 74-78.
DERIVATION OF NAME. Latin medius, middle — strictus, drawn tight.
DIAGNOSIS. Large, reticulate, trapezoidal Dictyocythere, with a slight constriction
in the mid-dorsal region.
HOLOTYPE. Geol. Dept. Brit. Mus. (N.H.) no. ^.48607. A male carapace (PI. 3,
figs. 3-6).
TYPE LOCALITY. Upper part of the Cypridea fasciculata subzone of the Middle
Purbeck Beds (shale 24 ft. above the Cinder Bed ; author's field ref. no. WT 33),
Worbarrow Tout, Dorset (Nat. Grid ref. 30/869796).
STRUCTURE, ETC. OF THE OSTRACOD HINGE 17
PARATYPE (same locality and horizon). In.486o6. A female carapace (PL 3, fig. 2).
DESCRIPTION. Strongly reticulate and trapezoidal, superficially much resembling
D. retirugata, but larger, and with a constriction about the middle of the back. Sexual
dimorphism marked, the females usually shorter than the specimen figured :
Dimensions (mm.) Proportions
L H W L H W
Carapace <$ (In. 48607 ; holotype) . . 1-12 0-63 0-56 . 1-79:1:0-89
Carapace $ (In.486o6) . . .1-02 0-67 0-52 . 1-54:1:0-78
Internal details not well displayed, owing to recrystallization of material. Hinge
distinctly amphidont, however, with anteromedian boss of left valve strongly
protuberant.
MATERIAL AND OCCURRENCE. This is a very abundant ostracod in the upper
part of the Middle Purbeck Beds of the Dorset coast, where it is associated with
species of Cypridea, but alternates in dominance with that genus in successive beds.
Hundreds of specimens have been isolated from several horizons at Worbarrow Tout,
and the species has also been found at other Dorset localities. Specimens from N.W.
Germany (sent by the kindness of Dr. J. Wolburg) from Lingen (Boring No. 29 at
1195 m.) are slightly smaller, but have an almost identical ornament to that of the
Dorset specimens. Wolburg (1950) classifies this horizon as " Wealden 2 ", and
correlates it with the British Upper Purbeck. The type-locality in Dorset is in the
upper part of the C. fasciculata subzone at the top of the Middle Purbeck.
Dictyocythere (Dictyocythere) decorata (Anderson)
(Plate 3, fig. i)
1941. Cythere retirugata Jones var. decorata Anderson, p. 374, pi. 18, fig. 4.
DIAGNOSIS. Reticulate Dictyocythere with a " second order " reticulation within
the main cells.
HOLOTYPE. Geol. Surv. Mus. no. 70339. Figured Anderson, 1941, pi. 18, fig. 4.
A right valve.
TYPE LOCALITY. Cythere Marl, Swindon Series, Swindon. Author's field ref.
no. TGA 7.
REMARKS. This is a rare species known only by three specimens from the
Swindon Series (Sylvester-Bradley, 1941). The hinge appears to be exactly as in
D. retirugata, but the shape is a little different, and the ornament more complex
(see PI. 3, fig. i) ; the " second-order " reticulation noticed by Anderson is not
confined, as suggested and figured by him, to the centre of the valve.
Dimensions (mm.) Proportions
L H W L H W
Left valve (In. 4861 8) .... 0-94 0-57 0-36 . 1-65:1:0-63
i8 STRUCTURE, ETC. OF THE OSTRACOD HINGE
Subgenus RHYSOCYTHERE nov.
TYPE-SPECIES. Cythere retirugata var. rugulata Jones, 1885.
DERIVATION OF NAME, pvaog, wrinkled + genus Cythere. Gender : feminine.
DIAGNOSIS. Dictyocythere with anterior element of hinge differentiated so that,
in the right valve, an anterior group of denticles lies in front of the usual boss.
Dictyocythere (Rhysocythere) rugulata (Jones)
(Plate 4, figs, i, 2, 5-10, 12-15)
1885. Cythere retirugata var. rugulata Jones, p. 350, pi. 9, figs. 17-20.
1941. Cythere retiviigata Jones var. rugulata Jones : Anderson, p. 373, pi. 18, fig. i.
DIAGNOSIS. Shape trapezoidal. Surface smooth over anterodorsal region,
strongly ridged along venter and anterior margin.
LECTOTYPE (here designated). Geol. Dept. Brit. Mus. (N.H.) no. In. 48600. A
carapace, $, not left valve, as stated by Jones. Figured Jones, 1885, pi. 9, fig. 17.
(Jones' No. 256, 2.)
TYPE LOCALITY. Hartwell, Buckinghamshire.
DESCRIPTION. Shape trapezoidal, dorsal and ventral margins being subparallel.
Moderate sexual dimorphism, males somewhat longer than females and much rarer.
Dimensions (mm.) Proportions
L H W L H W
Carapace $ (In. 48616) . . . 0-98 0-56 0-49 . 1-76
Left valve <$ (In. 4861 5) . . . 0-97 0-57 0-28 . 1-70
Left valve $ (In. 48604) . . . 0-84 0-55 0-29 . 1-54
Right valve $ (In. 4 8605) . . . 0-86 0-55 0-29 . 1-59
Left valve, juv. (In. 48614) . . . 0-68 0-45 0-24 . 1-50
Right valve, juv. (^.48613). . . 0-67 0-41 0-20 . 1-61
0-88
0-49
o-54
o-54
o-53
0-48
Valves almost smooth over the anterodorsal region of the carapace, but minutely
punctate over a small region just below and behind the centre of the carapace (PI. 4,
fig. 6). Venter and anterior border strongly ridged (PL 4, figs. 12-14). Normal pore
canals large and obvious in translucent specimens (PL 4, fig. 6) though they have
often been misinterpreted as pits or bosses. Radial pore canals few, straight, sparse.
Anterior duplicature fairly wide, with slight vestibule. Muscle-scar pattern (PL 4,
fig. 5) consisting of four scars in vertical superposition, with one oval scar in front
of them. Hinge of adult is amphidont, the anterior element clearly differentiated
into an anterior group of about four small denticles in front and to a well-marked
boss behind (PL 4, figs, i, 2 and 10) :
Adult Left valve Right valve
Anterior element . . Deep pit, with an anterior . Protuberant boss lying at the
loculate groove leading out posterior end of a dentate
from it ridge, the denticles decreasing
in size towards the anterior.
Anteromedian element . Rounded boss . Deep pit.
Posteromedian element . Smooth bar . Shallow groove.
Posterior element . . Loculate groove . Dentate ridge.
STRUCTURE, ETC. OF THE OSTRACOD HINGE 19
The hinge of juvenile moults (PI. 4, figs. 7-9) is particularly interesting, as
it is transitional between merodont and entomodont, with a subdivided, nearly
smooth, median element, but dentate terminal elements. The hinge of this species is
therefore palingenetic, and it seems that all investigations of the ontogeny of hinge
development so far published show that recapitulation takes place.
Juvenile Left valve Right valve
Anterior element . . Loculate groove . Dentate ridge.
Median element . . Smooth bar, the anterior end . Groove, the anterior end more
swollen and protruding fur- deeply excavated than post-
ther than posterior erior.
Posterior element . . Loculate groove . Dentate ridge.
Dictyocythere (Rhysocythere) transiens (Jones)
(Plate 3, figs. 11-13)
1885. Cythere transiens Jones, p. 349, pi. 9, figs. 13-16.
DIAGNOSIS. Small reticulate Dictyocythere, tapering strongly to posterior in side
view ; with Rhysocythere hinge
LECTOTYPE (here designated). Geol. Dept. Brit. Mus. (N.H.) no. ^.48603. A
left valve. Figured Jones, 1885, pi. 9, fig. 16. (Jones' No. 364, 2.)
TYPE LOCALITY. " Lower Purbeck, Swindon " (but see below).
DESCRIPTION. Fairly coarsely reticulate, tapering strongly to posterior in side view,
sexual dimorphism not observed.
Dimensions (mm.) Proportions
L H W L H W
Carapace (In. 48617) .... 0-60 0-37 0-31 . 1-61 : i : 0-82
Hinge appears to be exactly as in D. (R.) rugulata.
OCCURRENCE. Although the type locality is at Swindon, only a few specimens
have been found there by the author in the f " Swindon Series ", and these
may have been derived from the Portlandian " Swindon Sands and Stone " below.
The species is abundant at a certain horizon of the " Bugle Pit ", Hartwell, within the
Creamy Limestones of undoubted Portlandian age (author's field ref. no. BP 18 ;
this is from Bed 6 of Arkell, 1947 : 126). The species is characteristic, in fact, of the
Upper Portlandian, and only with some doubt can its range be said to include the
Lower Purbeck, despite Jones' contention. The horizon " Lower Purbeck" appears
to be an inference by Jones, and was not indicated by Blake, who collected the
specimens (see Jones, 1885 : 328).
ACKNOWLEDGMENTS
The author would like to express his thanks to Professor L. R. Moore, for critically
reading the manuscript, to Mr. Raymond R. Hibbard for presenting the material
from which the Devonian ostracods were isolated, and to Dr. J. Wolburg for material
20 STRUCTURE, ETC. OF THE OSTRACOD HINGE
from the Purbeck and Wealden Beds of N.W. Germany. The artist's fees (Pis. i
and 2) were met by a grant from the University of Sheffield Research Fund.
REFERENCES
ANDERSON, F. W. 1941. Ostracoda from the Portland and Purbeck Beds at Swindon. Proc.
Geol. Ass., London, 51 : 373-384, pis. 18, 19.
ARKELL W. J. 1947. The Geology of Oxford, vi + 267 pp. Oxford.
ARKELL, W. J. & SYLVESTER-BRADLEY, P. C. 1942. Notes on the Age of the Swindon Purbeck
Beds. Proc. Geol. Ass., London, 52 : 321-327.
BLAKE, J. F. 1885. In discussion on JONES, T. R., 1885.
BOLD, W. A. VAN DEN. 1946. Contribution to -the study of Ostracoda with special reference to
the Tertiary and Cretaceous Microfauna of the Caribbean region. 167 pp., 18 pis. Amster-
dam.
CORYELL, H. N. & MALKIN, D. S. 1936. Some Hamilton Ostracodes from Arkona, Ontario.
Amer. Mus. Novit., New York, 891': 1-20, pls./i, 2.
SAMPLE, C. H. & JENNINGS, P. H. 1935. Bairdoppilata, a new genus of Ostracoda,
with two new species. Amer. Mus. Navit., New. York, 777 : 1-5.
GREKOFF, N. 1952. Quelques donnees sur l-'utilisation des ostracodes dans les Etudes strati-
graphiques et paleographiques des series Mesozoique et Cenozoique. Principles de classi-
fication et de determination. Inst. Franc, du Petr., 47 pp., 5 pis. (Mimeograph).
HENNINGSMOEN, G. 1953. Classification of Paleozoic Straight-Hinged Ostracods. Norsk
geol. Tidskr., Kristiana, 31 : 185-288, pis. i, 2.
JONES, T. R. 1885. On the Ostracoda of the Purbeck Formation ; with Notes on the Wealden
Species. Quart. J. Geol. Soc. Lond. 41 : 311-353, pis. 8, 9.
JONES, T. R. & SHERBORN, C. D. 1888. On some Ostracoda from the Fuller's earth Oolite
and Bradford Clay. Proc. Bath Nat. Hist. Fid Cl., 6 : 249-278, pis. 1-5.
KELLET, B. 1943. Permian Ostracodes. /. Paleont., Chicago, 17 : 615-628.
KESLING, R. V. 1951. Terminology of Ostracod carapaces. Contr. Mus. Paleont. Univ.
Michigan, 9 : 93-171, pis. 1-18.
KINGMA, J. T. 1948. Contributions to the knowledge of the young-Cenozoic Ostracoda from the
Malayan region. 118 pp., n pis. Utrecht.
MALKIN, D. S. 1953. Biostratigraphic study of Miocene Ostracoda of New Jersey, Maryland,
and Virginia. /. Paleont., Chicago, 27 : 761-799, pis. 78-82.
MARTIN, G. P. R. 1940. Ostracoden des norddeutschen Purbeck und Wealden. Sencken-
bergiana, Frankfurt a.M., 22 : 275-361, pis. 1-13.
POKORNY, V. 1952. The Ostracods of the so-called Basal Horizon of the Subglobosa Beds at
Hodonin (Pliocene, Inner Alpine Basin, Czechoslovakia). Shorn, geol. Ust. csl., Praha, 19
(Pal.) : 229-396, pis. 7-12. (In Czech and Russian, with English summary.)
STEPHENSON, M. B. 1946. Weches Eocene Ostracoda from Smithville, Texas. /. Paleont.,
Chicago, 20 : 297-344, pis. 42-45.
SYLVESTER-BRADLEY, P. C. 1941. The Purbeck Beds of Swindon. Proc. Geol. A ss., London,
51 : 349-372.
19480. Bathonian Ostracods from the Boueti Bed of Langton Herring, Dorset. Geol.
Mag., London, 85 : 185-204, pis. 12-15.
19486. The Ostracode Genus Cythereis. J. Paleont., Chicago, 22 : 792-797, pi. 122.
1948^. The Shell of the Ostracod Genus Macrocypris. Ann. Mag. Nat. Hist., London
(12) 1 : 65-71, 3 figs.
1949. The Ostracod Genus Cypridea and the zones of the Upper and Middle Purbeckian.
Proc. Geol. Ass., London, 60 : 125-153, pis. 3, 4.
TRIEBEL, E. 1948. Zur Kenntnis der Ostracoden-Gattung Triebelina. Senckenbergiana,
Frankfurt a.M., 29 : 17-22, 5 figs.
1949- Mikropalaontologischen Kennzeichnung der Ostracoden-Gattungen Xenocythere
und Palmenella. Senckenbergiana, Frankfurt a.M., 30 : 185-192, pis. i, 2.
STRUCTURE, ETC. OF THE OSTRACOD HINGE
21
1950. Homoomorphe Ostracoden-Gattungen. Senckenbergiana, Frankfurt a.M., 31 :
313-330. Pis. i-4-
1951. Einige stratigraphisch wertvolle Ostracoden aus dem hoheren Dogger Deutsch-
lands. Abh. senckenb. naturf. Ges. Frankfurt a.M., 485 : 87-101, pis. 44-49.
1954- Malm-Ostracoden mit amphidontem Schloss. Senckenb. leth., Frankfurt a.M.,
35 : 3-16, pis. 1—4.
WARTHIN, A. S. 1934. Common Ostracoda of the Traverse Group. Contr. Mus. Paleont.
Univ. Michigan, 4 : 205-226, pi. i.
WOLBURG, J. 1950. Vergleichende stratigraphische Untersuchungen der bradisch-limnischen
Ablagerungen Europas an der Wende Jura-Kreide. Geol. Jb., Hannover-Celle, 64 : 159-
171.
GEOL. Hi, 1.
PLATE i
Acanthocythere sphaerulata (Jones & Sherborn)
>
FIGS, i, 4. Left valve male (external and internal lateral views). In. 42434 (0-67 mm.
long).
FIGS. 2, 3. Right valve male (external and internal lateral views). 1^42433 (0-69 mm.
long).
Acanthocythere spiniscutulata n. sp.
FIGS. 5, 8. Left valve (external and internal lateral views), holotype. 1^42435 (0-57
mm. long).
FIGS. 6, 9. Right valve (external and internal lateral views), paratype. ^.42436 (0-56
mm. long).
FIG. 7. Carapace (dorsal view), paratype. In. 42437 (0-57 mm. long).
All figures approx. x 90 (exact lengths given above), from pencil drawings by Miss
D. Robinson based on camera lucida sketches by the author.
All specimens from the economic Fullers' Earth (Upper Fullers' Earth Clay, Bathonian,
Middle Jurassic) from the Fosse Way mine, nr. Bath, collected by the author, now in the
British Museum (Nat. Hist.).
Ru)l. B. M.\N.H.) Geol. Ill, i.
1LATE I
ACANTHOCYTHERE GEN. NOV. (MIDDLE JURASSIC)
PLATE 2
Quasillites sp.
FIG. i. Right valve, external lateral view. The irregularly shaped blank space below the
centre is obscured by matrix. In. 48624 (1-16 mm. long).
FIG. 2. Lett valve, internal lateral view. In. 48623 (1-09 mm. long).
Bufina sp.
FIG. 3. Left valve, external lateral view. In. 48622 (0-89 mm. long).
FIG. 4. Right valve, external lateral view. In. 48621 (0-93 mm. long).
Euglyphella sp.
FIGS. 5, 6. Right valve, external lateral and internal lateral views. In. 48619 (0-99 mm.
long).
FIG. 7. Left valve, internal lateral view. In. 48620 (1-13 mm. long).
Figs, i, 2, approx. x 70 ; 3-7, x 65. Exact lengths given above. All are pencil
drawings by Miss D. Robinson based on camera lucida sketches by the author.
All specimens from the Pleurodictyum Bed (Ludlowville, Hamilton Group, Middle
Devonian) of Bay View, Erie County, New York, U.S.A., collected by Raymond R. Hib-
bard, now in the British Museum (Nat. Hist.).
Bull. B. M. (N.H.) Gcol. Ill, t.
PLATE 2
MIDDLE DEVONIAN OUASILLITACEA
PLATE 3
Dictyocythere (Dictyocythere) decorata (Anderson)
FIG. i. Left valve, external lateral view. In. 48618 (0-94 mm. long). Cythere Marl,
Swindon Series, Swindon (? Purbeckian) .
Dictyocythere (Dictyocythere) mediostricta n. sp.
FIG. 2. Carapace $, left lateral view. In. 48606 (1-02 mm. long).
FIGS. 3-6. Carapace $, right lateral, left lateral, dorsal and ventral views, holotype. In.
48607 (1-1-2 mm. long). Middle Purbeck Beds, Worbarrow Tout, Dorset.
Dictyocythere (Dictyocythere) retirugata (Jones)
FIG. 7. Carapace <$, ventral view. Specimen accidentally destroyed.
FIG. 8. Right valve $, external lateral view. In. 48611 (0-84 mm. long).
FIG. 9. Left valve $, external lateral view. In. 48610 (0-80 mm. long).
FIG. 10. Right valve $, external lateral view. In. 48608 (0-97 mm. long).
All from Cythere Marl, Swindon Series, Swindon (? Purbeckian).
Dictyocythere (Rhy so cy there) transiens (Jones)
FIGS. 11-13. Carapace (right), lateral, dorsal and ventral views. In. 48617 (0-60 mm.
long). Upper Portland Beds, Bugle Pit, Hartwell.
All figures from photographs by the author, x 60 approx. The specimens whitened
with magnesium oxide before being photographed.
Bull. 13. M. (N.H.} Geol. Ill, i.
PLATE 3
DICTYOCYTHERE GEN. NOV. (UPPER JURASSIC)
PLATE 4
Diet} ocvthere (Rhy so cy there] rugulata (Jones)
FIGS, i, 2. Lett and right valves $, dorsal views. In. 48604 (0-84 mm. long), In. 48605
(0-86 mm. long).
Dictyocythere (Dictyocy there) retirugata (Jones)
FIGS. 3, 4. Left and right valves $, dorsal views. In. 48610 (0-80 mm. long), In. 48611 (0-84
mm. long).
Dictyocythere (Rhy socy there) rugulata (Jones)
FIG. 5. Internal view of centre of right valve to show muscle-scar pattern. In. 4861 2
(x 200).
FIG. 6. Left valve $ photographed under water by reflected light to show normal pore canals
and minute punctation behind and below central region. In. 48604 (0-84 mm. long).
FIGS. 7, 8. Dorsal views of left and right valves of juvenile moults showing entomodont
hinge. ^.48614 (0-68 mm. long), 111.48613 (0-67 mm. long).
FIGS. 9, 10. Anterior hinge elements of juvenile and adult right valves in dorsal view. In.
48613, In. 48605 (x 200).
Dictyocvthere (Dictyocythere) retirugata (Jones)
FIG. ii. Anterior element of right valve in dorsal view. In. 48611 (X 200).
Dictyocythere (Rhy socy there) rugulata (Jones)
FIG. 12. External lateral view of left valve of <$. In. 48615 (0-97 mm. long).
FIGS. 13-15. (J carapace in right lateral, ventral and dorsal views. 111.48616 (0-98 mm.
long).
Dictyocythere (Dictyocythere) retirugata (Jones).
FIGS. 16, 17. Dorsal views of left and right valves $. In. 48609 (0-92 mm. long), 111.48608
(0-97 mm. long).
Figs, i, 2, 6-10, 12-15 from so-called Purbeck Beds of Bugle Pit, Hartwell. Figs. 3, 4,
5, n, 16, 17 from Cythere Marl, Swindon Series, Swindon (? Purbeckian).
All figures from photographs by the author. Figs. 1-4, 6-8, 12-17 X 55 approx.
Specimens in Figs. 12-14 whitened with magnesium oxide before being photographed.
null. B. M. (N.H.) Gcol. HI, I.
PLATE 4
PICTYOCYTHERE GEN, NOV. (UPPER JURASSIC)
EOCENE MOLLUSCA FROM
NIGERIA :
A REVISION
F. E. EAMES
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 2
LONDON : 1957
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following, papers appeared in Volume I (1949-52) :
Price
No. I (1949). The Pterobranch Rhabdopleura in the English Eocene. H. D.
Thomas & A. G. Davis . . . . . . . . 75. 6d.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P. Oakley
& M. F. Ashley Montagu ....... 55.
No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone of
the Welsh Borders. E. I. White.
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White 75. 6d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan, North-
ern Iraq. L. F. Spath ........ IDS.
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum. T. H. Withers ........ 55.
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae). T. H.
Withers .......... 55.
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian of
Podolia. W. N. Croft los.
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle East.
T. F. Grimsdale ......... 105.
No. 9 (1952). Australian Arthrodires. E. I. White .... 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.
The following papers appeared in Volume II (1953-56) :
No. I (1953). The Deer of the Weybourn Crag and Forest Bed of Norfolk.
A. Azzaroli . . . . . . . . . . £i 55.
No. 2 (1953). A Coniferous Petrified Forest in Patagonia. Mary G. Calder I2S.
No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner, K. P.
Oakley & W. E. Le Gros Clark . . . . 35 6d.
No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.
Marjorie E. J. Chandler ........ 165.
No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans . . 155.
No. 6 (1955). Further Contributions to the Solution of the Piltdown
Problem. J. S. Weiner, W. E. Le Gros Clark, K. P. Oakley &
others ........... £i
No. 7 (1955). The Schizaeaceae of the South of England in early Tertiary
Times. Marjorie E. J. Chandler . . . . . . 155.
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . . £i
EOCENE MOLLUSCA FROM NIGERIA
A REVISION
BY
FRANK E. EAMES
Pp. 23-70 ; Pis. 5-10
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 2
LONDON: 1957
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series, corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 2 of the Geological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued October, 1957 Price Twenty-five Shillings
EOCENE MOLLUSCA FROM NIGERIA:
A REVISION
By F. E. EAMES
CONTENTS
Page
I. INTRODUCTION AND GENERAL REVIEW ...... 29
II. SYSTEMATIC DESCRIPTIONS ........ 30
A. GASTROPODA .... . -3°
Family POTAMIDIDAE ........ 30
Genus Exechostoma Cossmann . . . . .30
Exechostoma cossmanni Newton ..... 30
Genus Terebralia Swainson ....... 30
Terebralia nigeriensis sp. nov. ..... 30
Terebralia amekiensis sp. nov. . . . . .31
Family TURRITELLIDAE . . . . . . . .31
Genus Turritella Lamarck . . . . . . .31
Turritella amekiensis sp. nov. . . . . .31
Subgenus Coeloconica nov. ....... 32
Turritella (Coeloconica) mauryana Newton . . .32
Family ARCHITECTONICIDAE ....... 32
Genus Architectonica (Bolten MS.) Roding .... 32
Subgenus Nipteraxis Cossmann ...... 32
Architectonica (Nipteraxis) bendeica sp. nov. ... 32
Subgenus Stellaxis Dall . . . . . . -33
Architectonica (Stellaxis) bicingulata (Newton) . . 33
Subgenus Solariaxis Dall ....... 33
Architectonica (Solariaxis) amekiensis sp. nov. . . 33
Family SCALIDAE ......... 34
Genus Acrilla H. Adams ....... 34
Acrilla nigeriensis sp. nov. ...... 34
Family CALYPTRAEIDAE ........ 35
Genus Calyptraea Lamarck ....... 35
Calyptraea newtoni sp. nov. ...... 35
Genus Turbocalyptraea nov. ....... 35
Turbocalyptraea scabrosa sp. nov. ..... 35
Genus Crepidula Lamarck ....... 36
Subgenus Concavimargo nov. ...... 36
Crepidula (Concavimargo) falconeri Newton ... 36
Family XENOPHORIDAE ........ 36
Genus Xenophora Fischer von Waldheim . . . 36
Xenophora nigeriensis (Newton) ..... 36
Family STROMBIDAE ........ 36
Genus Tibia (Bolten MS.) Roding 36
Tibia bidigitata (Newton) ...... 36
Genus Cyrtulotibia nov. . . . . . . .36
Cyrtulotibia unidigitata (Newton) . . . . -37
GEOL. Ill, 2.
26 EOCENE MOLLUSCA FROM NIGERIA
Genus Semiterebellum Cossmann . . . . . -37
Subgenus Africoterebellum nov. . . . . . -37
Semiterebellum (Africoterebellum) elongatum Newton . 38
Genus Amekichilus nov. ....... 38
Amekichilus suturocostatum (Newton) .... 39
Family AMPHIPERATIDAE
Genus Eovolva Schilder ........ 39
Eovolva nigeriensis (Newton) ..... 39
Genus Sphaerocypraea Schilder ...... 39
Sphaerocypraea sudanensis (Schilder) . . . -39
Family NATICIDAE ......... 39
Genus Neverita Risso ........ 39
Neverita amekiensis sp. nov. ..... 39
Genus Sinum (Bolten MS.) Roding ..... 40
Sinum africanum Newton ...... 40
Sinum nigeriense sp. nov. ...... 40
Family CYMATIIDAE ........ 40
Genus Varicohilda nov . . . ... . .40
Varicohilda turriculata (Newton) . . . . .41
Family MURICIDAE . . . . . . . . .41
Genus Hexaplex Perry . . . . . . . .41
Subgenus Paziella Jousseaume . . . . . .41
Hexaplex (Paziella) bendeica sp. nov. . . . .41
Genus Pterynotus Swainson . . . . . . .42
Pterynotus newtoni sp. nov. ...... 42
Family BUCCINIDAE ........ 43
Genus Bendeia nov. ........ 43
Bendeia africana (Newton) ...... 43
Genus Laccinum nov. ........ 43
Laccinum lugardi (Newton) ...... 44
Genus Janiopsis Rovereto ....... 44
Janiopsis nigeriensis Newton ..... 44
Family VOLEMIDAE ......... 44
Genus Pseudomazzalina nov. . . . ' . . . .44
Pseudomazzalina nigeriensis (Newton) . . . .45
Family FUSINIDAE ......... 45
Genus Clavilithes Swainson . . . . . . .45
Subgenus Africolithes nov. ...... 45
Clavilithes (Africolithes) africanus (Newton) ... 45
Genus Leucozonia Gray ........ 46
Leucozonia pseudominax sp. nov. . . . .46
Family VOLUTIDAE ......... 46
Genus Volutocorbis Dall ....... 46
Volutocorbis multispinosa (Newton) .... 46
Genus Bendeluta nov. ........ 46
Bendeluta conicoturrita (Newton) ..... 47
Family OLIVIDAE .......... 47
Genus Pseudoliva Swainson . . . . . . .47
Subgenus Buccinorbis Conrad . . . . . -47
Pseudoliva (Buccinorbis) kitsoni (Newton) . . '! . 47
Family STREPSIDURIDAE ........ 4^
Genus Strepsidura Swainson ....... 48
Subgenus Strepdiduropsis nov. ...... 48
Strepsidura (Strepsiduropsis) spirata Newton ... 48
EOCENE MOLLUSCA FROM NIGERIA 27
Page
Family CANCELLARIIDAE ........ 48
Genus Sveltia Jousseaume ....... 48
Subgenus Africosveltia nov. ...... 48
Sveltia (Africosveltia} multiplicis (Newton) ... 49
Genus Bonellitia Jousseaume ...... 49
Subgenus Admetula Cossmann ...... 49
Bonellitia (Admetula) amekiensis sp. nov. ... 49
Subgenus Africostoma nov. ...... 49
Bonellitia (Africostoma) decorata (Newton) ... 50
Family CONIDAE ......... 50
Genus Conus Linn6 . . . . . . . .50
Subgenus Leptoconus Swainson ...... 50
Conus (Leptoconus) amekiensis sp. nov. ... 50
Family TURRIDAE . . . . . . . .51
Genus Eopleurotoma Cossmann . . . . . .51
Eopleurotoma nigeriensis Newton . . . . .51
Genus Turricula Schumacher . . . . . .51
Subgenus Surcula H. & A. Adams . . . . .51
Turricula (Surcula) africana (Newton) . . . .51
Genus Surculites Conrad . . . . . . .51
Subgenus Clinura Bellardi . . . . . . .51
Surculites (Clinura) ingens (Mayer-Eymar) . . .51
Genus Mitrelloturris nov. . . . . . . .51
Mitrelloturris casteri (Chavan) . . . . .52
Genus Amekicythara nov. . . . . . . .52
Amekicythara douvillei (Newton) . . . . -53
B. LAMELLIBRANCHIA 53
Family NUCULIDAE ........ 53
Genus Nucula Lamarck . . . . . . .53
Nucula costaeimbricatis Newton . . . . .53
Family GLYCYMERIDAE . . . . . . . .53
Genus Africarca nov. ........ 53
Africarca nigeriensis (Newton) . . . . .53
Family NOETIIDAE ......... 54
Genus Protonoetia MacNeil . * . . . . .54
Protonoetia nigeriensis (Newton) ..... 54
Genus AY cop sis von Koenen . . . . . . .54
Arcopsis africana (Newton) ...... 54
Genus Rectangularca nov. ....... 54
Rectangular ca africana (Newton) . . . . -55
Family PLICATULIDAE . . . . . . . .55
Genus Plicatula Lamarck . . . . . . -55
Plicatula polymorpha Bellardi . . . . -55
Family OSTREIDAE ......... 55
Genus Ostrea Linn6 ........ 55
Ostrea amekiensis sp. nov. . . . . . -55
Ostrea pseudomarginidentata sp. nov. .... 56
Subgenus Crassostrea Sacco ...... 56
Ostrea (Crassostrea) lugardi (Newton) . . . .56
Family CARDITIDAE ........ 57
Genus Glans M. von Muhlfeldt . . . . . -57
Glans nigeriensis sp. nov. . . . . . -57
28 EOCENE MOLLUSCA FROM NIGERIA
Page
Subgenus Bendeglans nov. . . . . . . 57
Glans (Bendeglans} costaeirregularis (Newton) . . .58
Subgenus Amekiglans nov. ...... 58
Glans (Amekiglans} costaenodulosis (Newton) . . -59
Subgenus Divergidens nov. ...... 60
Glans (Divergidens} triparticostata (Newton) . . .60
Family LUCINIDAE ......... 60
Genus Phacoides Gray ........ 60
Phacoides eaglesomei Newton ..... 60
Genus Gibbolucina Cossmann . . . . . .61
Subgenus Eomiltha Cossmann . . . . . .61
Gibbolucina (Eomiltha} (?} subrhomboidalis (Newton) . 61
Genus Pompholigina Dall . . . . . . .61
Subgenus Eodivaricella Chavan . . . . . .61
Pompholigina (Eodivaricella) oppenheimi (Newton) . . 61
Family CARDIIDAE ......... 61
Genus Fragum (Bolten MS.) Rodin g . . . . .61
Subgenus Africofragum nov. ...... 61
Fragum (Africofragum} newtoni sp. nov. . . .61
Family VENERIDAE ......... 62
Genus Tivelina Cossmann . . . . . . .62
Tivelina newtoni sp. nov. ...... 62
Genus Pitar Roemer ........ 63
Pitar amekiensis sp. nov. ...... 63
Genus Chionella Cossmann . . . . . . -63
Subgenus Costacallista Palmer ...... 63
Chionella (Costacallista} elongatotrigona (Newton) . . 63
Subgenus Microcallista Stewart ...... 64
Chionella (Microcallista) kitsoni (Newton) . .64
Genus Sinodia Jukes-Browne ...... 64
Sinodia heward-belli Newton ..... 64
Genus Sinodiopsis nov. ....... 64
Sinodiopsis coxi sp. nov. ...... 65
Family MACTRIDAE ......... 65
Genus Spisula Gray ........ 65
Subgenus Crepispisula nov. ...... 65
Spisula (Crepispisula} amekiensis sp. nov. . . -65
Family TELLINIDAE ......... 66
Genus Macoma Leach ........ 66
Subgenus Bendemacoma nov. ...... 66
Macoma (Bendemacoma} nigeriensis (Newton) . . 66
Family MYIDAE ......... 67
Genus Raetomya Newton ....... 67
Raetomya schweinfurthi (Mayer-Eymar) . . -67
Family CORBULIDAE ........ 67
Genus Varicorbula Grant & Gale ...... 67
Varicorbula amekiensis sp. nov. . . . . .67
Family KITSONIIDAE nov. ....... 68
Genus Kitsonia nov. ........ 68
Kitsonia eocenica (Newton) ...... 68
III. ACKNOWLEDGMENTS ........ .68
IV. REFERENCES 68
EOCENE MOLLUSCA FROM NIGERIA 29
SYNOPSIS
The molluscan fauna described by R. B. Newton (1922) from the Eocene of Bende Ameki
(Nigeria) is revised. Fifteen new gastropod species and eight new lamellibranch species are
described ; ten new genera and seven new subgenera of gastropoda and four new genera and
six new subgenera of lamellibranchia are proposed, one of the lamellibranch genera belonging
to a new family. The Eocene age of the fauna is confirmed, but it is believed to be of Upper
Eocene (Bartonian), rather than Middle Eocene (Upper Lutetian), age.
I. INTRODUCTION AND GENERAL REVIEW
DURING the course of exploration in Nigeria by the joint Shell and British Petroleum
organization it has frequently been necessary to consult R. B. Newton's work on
Eocene Mollusca from Nigeria (1922). Certain problems concerning the age of the
beds arose, and these led the writer to investigate the fauna more fully. A detailed
study of the material described by Newton, which is in the collections of the British
Museum (Natural History), was undertaken, with the result that, while the con-
clusions agreed fairly closely with Newton's opinion as to the age, it became
apparent that many of the generic determinations required revision, and that not
only was only one European species present, but also that all forms compared by
Newton with European ones were specifically, and in some cases even generically,
distinct. A full description of many of the forms is given in Part II of this paper.
On the basis of the fauna he described, Newton concluded that the beds were of
Upper Lutetian age, although he recognized that the Bartonian facies was very
apparent. The result of the present revision has been to show that only two lamelli-
branchia— Plicatula polymorpha Bellardi and Raetomya schweinfurthi (Mayer-
Eymar) — occur outside Nigeria, in the Upper Mokattam Beds of Egypt ; the
Plicatula has been found also in the Priabonian of Italy, and the Raetomya in the
Eocene of Senegal and the Cameroons. Of the gastropod genera, Strepsidura (sensu
lato) ranges from Eocene to Oligocene, and Exechostoma and the subgenus Buccin-
orbis range from Maestrichtian to Eocene ; not one of the gastropod species has yet
been found elsewhere. The cephalopod genus Belosepia is restricted to the Eocene.
Although the bryozoan genus Cupuladria, hitherto believed to range only from
Miocene to Recent, has been found in the beds by palaeontologists of the Shell Oil
Company, the Eocene age indicated by the mollusca is also confirmed by the fish
remains recorded, which include the genus Cylindr acanthus. The coral Turbinolia,
which is also of fairly common occurrence in the beds, is known only from Eocene
and Oligocene deposits.
Although two of the lamellibranch species have been recorded from the Upper
Mokattam Beds of Egypt, these beds have also yielded Nummulites beaumonti and
Orbitolites complanatus ; these latter fossils suggest a rather earlier age than
Bartonian (to which stage the Upper Mokattam Beds have been assigned), and appear
to suggest that the Upper Mokattam is not entirely of Bartonian age.
In Angola, Dartevelle & Roger have recently clearly shown that the beds (Quim-
briz Beds) in which Platyodon klinghardti (mistakenly referred to Raetomya schwein-
furthi by Caster) and Macrocallista palmerae (to which the Nigerian form here named
Sinodiopsis coxi was erroneously referred) are really of Miocene age. Any similarity
30 EOCENE MOLLUSCA FROM NIGERIA
of the Angola fauna to that of the Nigerian Bende Ameki Beds is evidently purely
superficial, no species being in common. Dartevelle & Roger refer to Raetomya as
a mactrid, but Newton has clearly shown that it is a myid ; indeed, Dartevelle &
Roger had some doubts about referring the species klinghardti to the American
genus Platyodon, and it may well be that it also is a Raetomya.
As a result of the revision of the molluscan fauna from Bende Ameki, the writer
believes that the evidence and the general relationships of the fauna, while confirming
Newton's opinion as to the Eocene age, suggest that an Upper Eocene (Bartonian)
rather than a Middle Eocene (Upper Lutetian) age, is indicated.
II. SYSTEMATIC DESCRIPTIONS
GASTROPODA
Family POTAMIDIDAE
Genus EXECHOSTOMA Cossmann, 1889
Exechostoma cossmanni Newton
1922. Exechostoma cossmanni Newton, p. 45, pi. 5, figs. 8-9.
REMARKS. Specimen G.42I77 (Newton's fig. 9) is now selected as lectotype.
Genus TEREBRALIA Swainson, 1840
Terebralia nigeriensis sp. nov.
1922. Terebralia sp.A. Newton, p. 46, pi. 4, fig. 10.
MATERIAL. The holotype (G. 42 174).
DESCRIPTION. Apical whorls missing and aperture broken. Of medium size,
turriculate-conic, spire angle 19°. Whorls (of which four or five are preserved)
almost flat-sided, their height very slightly more than half their width ; sutures
deeply impressed, distinctly undulatory. Ornament consisting of moderately coarse
and fairly widely-spaced ribs crossed by strong spiral threads ; ribs somewhat
irregularly spaced, sometimes only slightly narrower than, but sometimes (especially
on the later whorls) only half the width of, their intervals, gently opisthocline
adapically, their forward-directed face slightly concave, more strongly so abapically,
where they tend to be slightly swollen. On the whorls preserved there are n strong
spiral threads with intervals of about the same width. There is no noticeable
development of varices. Abapical portion of last whorl poorly preserved, without
ribs, but with about nine incised spiral grooves separated by intervals of approxi-
mately half their width. Columella moderately concave and callous ; other apertural
characters not determinable.
DIMENSIONS. Height (incomplete) 49-7 mm. Diameter of last whorl 21-0 mm.
REMARKS. The above characters are sufficient for specific determination.
EOCENE MOLLUSCA FROM NIGERIA 31
Terebralia amekiensis sp. nov.
1922. Terebralia sp.B. Newton, p. 48, pi. 4, fig. n.
MATERIAL. The holotype (6.42175).
DESCRIPTION. Apical whorls missing and aperture broken, about five whorls
preserved. Of small-medium size, turriculate-conic, spire angle i7°-i8°. Whorls
flat-sided, their height about two-fifths of their width ; sutures linear and moder-
ately deep, gently undulatory. Ornament consisting of relatively fine and closely-
spaced ribs crossed by strong spiral threads ; ribs of about the same width as their
intervals, their forward-directed face sometimes gently concave, vertical or slightly
opisthocline abapically. On the whorls preserved there are eight to ten fairly strong
spiral threads with intervals of about the same width ; no noticeable development
of varices. Base broken, without ribs, but carrying strong spiral threads with
intervals of about the same width.
" DIMENSIONS. Height (incomplete) 38-4 mm. Diameter of last whorl 15-3 mm.
REMARKS. The above characters are sufficient to define the species.
Family TURRITELLIDAE
Genus TURRITELLA Lamarck, 1799
Turritella amekiensis sp. nov.
(PI. 5, figs, i, 2)
1922. Turritella cf. sulcifera Deshayes : Newton, p. 50, pi. 5, fig. 7.
MATERIAL. Numerous specimens, including the holotype (G. 42215, Newton's
fig. 7)-
DESCRIPTION. Of medium size, turriculate-conic, spire angle 15°, last whorl
about one-eighth of the total height. Protoconcti not preserved, evidently small.
Whorls distinctly convex, their greatest convexity slightly below the middle, upper
slope less convex than the lower. Sutures linear. Height of whorls about seven-
twelfths of their width. The earliest whorls seen carry four spiral threads, the
number increasing to about 18 on the last spire whorl, on which a few very fine
additional threads are present in some of the intervals. Base of last whorl with
threads of a similar type. Columella gently concave ; columellar lip narrow, especially
abapically. Aperture evidently rounded-subquadrate. No siphonal fascicle. Growth
lines concave forwards, distinctly prosocline adapically, very slightly prosocline
abapically, the maximum concavity at about three-fifths the height of the whorl.
DIMENSIONS. Height 27-4 mm., width 7-5 mm.
REMARKS. The Nigerian form is consistently smaller than T. sulcifera Deshayes
of the Paris Basin Lutetian ; its whorls do not tend to be concave above, and the
details of the spiral ornament are different.
32 EOCENE MOLLUSCA FROM NIGERIA
Subgenus COELOCONICA nov.
TYPE SPECIES. Turritella mauryana Newton.
SUBGENERIC CHARACTERS. Of moderate size to fairly large, distinctly coeloconoid.
Protoconch not preserved, evidently small. Early whorls subcylindrical to slightly
convex, with three sharp spiral threads the uppermost of which is fairly close to the
suture ; traces of a fourth thread at the lower suture ; all four threads with sharp,
closely-spaced, vertically diposed crenulations where crossed by growth lines ; rest
of surface with microscopic spirals. With growth the third thread from the adapical
suture gradually develops into a very prominent, sharp flange. Base of last whorl
with a second prominent, but feebler, keel and two obscure threads below, as well as
microscopic spirals. Aperture rounded-subquadrate, as in Turritella. Growth-lines
with a deep U-shaped sinus the apex of which is slightly above the second keel ;
markedly prosocline at the adapical suture, orthocline at the abapical suture, their
lower end immediately below the upper.
REMARKS. The high, gently coeloconoid spire, the details of ornament (including
development of the strong flange), and the form of the growth-lines differentiate this
from other described groups of Turritella.
Turritella (Coeloconica) mauryana Newton
1922. Turritella mauryana Newton, p. 48, pi. 5, figs. 4-6.
MATERIAL. Several specimens, including the lectotype (G. 42209, Newton's fig. 4)
here selected.
Family ARCHITECTONICIDAE
Genus ARCHITECTONICA (Bolten MS.) Roding, 1798
Subgenus NIPTERAXIS Cossmann, 1915
Architectonica (Nipteraxis) bendeica sp. nov.
(PI. 5, figs. 3«-c)
1922. Solariaxis cf. canaliculata (Lamarck): Newton, p. 54.
MATERIAL. The holotype (G.42373) and a few other specimens.
DESCRIPTION. Moderately small, low-conic to slightly cyrtoconoid, apical angle
I4O°-I45°. Protoconch smooth, anastrophic ; 3-4 flattened, very slightly convex
spire whorls with deep, sunken sutures. Ornament consisting of five crenulated
spiral threads, the uppermost one the most prominent and with the strongest
crenulations, the others subequal. Last whorl sharply rounded at the periphery
where there is a sixth and broader cordon, and with a fine subsidiary thread in each
interval ; on large specimens a second order of intercalaries may appear. Base
gently convex, with seven crenulated or beaded spiral threads, the outer three finer
and moderately widely spaced, the inner three coarser and with coarser beading,
the innermost one well within the umbilicus ; blunt, accentuated growth lines join
EOCENE MOLLUSCA FROM NIGERIA 33
the crenulations. Umbilicus deep, wide and completely visible, occupying about
two-fifths of the diameter of the base. Growth lines gently prosocline, slightly
sinuous, becoming orthocline at the upper suture. Aperture rounded kite-shaped,
peristome discontinuous ; columella thin, slightly excavated, with two small furrows
corresponding to the two innermost spiral threads on the base, the upper furrow
more distinct.
DIMENSIONS. Holotype : height 5-7 mm., width 12-0 mm.
A topotype (0.42374) has a width of 13-8 mm.
REMARKS. Compared with A. canaliculata (Lamarck), this form lacks the sharp
peripheral keel, and the details of the ornament are quite distinct.
Subgenus STELLAXIS Ball, 1892
Architectonica (Stellaxis) bicingulata (Newton)
(PL 5, ng. 4)
1922. Stellaxis bicingulata Newton, p. 52, pi. 5, figs. 10-11.
MATERIAL. Numerous specimens, including the lectotype (G. 42355, Newton's
fig. 10) here selected.
SUPPLEMENTARY DESCRIPTION. Of medium size, solarioid, spire slightly cyrto-
conoid ; spire angle decreasing during growth from about 125° to about a right
angle or slightly less ; last whorl forming slightly more than one-third of the
height. Protoconch smooth and loosely coiled, anastrophic. About five
spire whorls, which are gently convex, the main surface separated from a narrow
but distinct, smooth, abapical sutural cord by a spiral groove ; main surface
smooth in the earlier stages, later developing extremely vague, irregular spiral
threads. Sutures linear. Growth lines almost straight, prosocline. Marginal keel
on last whorl with a few very faint threads, occasionally appearing to be slightly
bifid. Base gently convex medially, flatter marginally where there is one smooth
spiral cord close to the peripheral keel. Umbilicus very deep, wide and completely
visible, occupying about a third of the diameter of the base, margined by blunt teeth
which are sometimes made to appear vaguely bifid by the accentuated growth-lines,
its inner wall vertical and with a spiral thread at about two-thirds of its height ;
a fine spiral groove limits the teeth externally in young forms, but is often obsolete or
absent in adults. Aperture kite-shaped, with a small indentation level with the row of
umbilical teeth, and a still fainter one by the umbilical thread above.
Subgenus SOLARIAXIS Dall, 1892
Architectonica (Solariaxis) amekiensis sp. nov.
(P. 5, figs. $a-c)
1922. Solariaxis cf. spectabilis (]. de C. Sowerby): Newton, p. 53, pi. 5, figs. 12-13.
MATERIAL. A few specimens, including the holotype (0.42361, Newton's fig. 12).
DESCRIPTION. Of medium size, moderately low, gently cyrtoconoid, spire angle
34 EOCENE MOLLUSCA FROM NIGERIA
decreasing during growth from about 130° to about 105° (excluding the peripheral
downturning of the last whorl). Protoconch not well preserved, anastrophic. About
five flattened spire whorls with deep sutures, slightly imbricate. Ornament consisting
of seven rather finely crenulated, not very coarse spiral threads, the fourth and
sixth (from the adapical suture) finer than the others, the second, third and fifth
coarser, the first even a little coarser than these. Last whorl bluntly angular at the
periphery, its upper four primary threads rather wide-spaced, with a faint double
intercalary, a fine single intercalary, and a stronger double intercalary in the upper,
median and lower intervals, respectively ; the five lower primary threads more
closely-spaced, the upper interval with a fine intercalary, the peripheral thread
double. Base flattened, only very slightly convex, with seven crenulated or beaded
spiral threads increasing in strength towards the umbilicus, the wall of the latter
with two finer, widely-spaced threads ; growth lines serrate but not thickened.
Umbilicus deep, wide and completely visible, occupying about a third of the diameter
of the base. Growth lines distinctly prosocline, fairly straight, but becoming orthocline
at the adapical suture. Aperture oval, kite-shaped ; peristome discontinuous. Columella
thin, gently concave, with four slight furrows corresponding to the two umbilical
threads and the two innermost spiral threads on the base.
DIMENSIONS. Holotype : height 13-4 mm., width 22-8 mm.
REMARKS. Compared with that of A. (Solariaxis) spectabilis (J. de C. Sowerby)
the spiral ornament is coarser and differently disposed.
Family SCALIDAE
Genus ACHILLA H. Adams, 1860
Acrilla nigeriensis sp. nov.
(PI. 5, ng. 6)
1922. Acrilla cf. affinis (Deshayes) : Newton, p. 50, pi. 3, figs. 10-12.
MATERIAL. The holotype (0.42285, Newton's fig. 10), and a few other specimens.
DESCRIPTION. Specimens incomplete, apical whorls missing and aperture broken.
The largest specimen (the holotype) has about eight whorls preserved, spire angle
11° ; they are rather loosely coiled, with deep, linear sutures, and are very strongly
convex, with a tendency to angularity at about three-quarters or more of their height.
Ornament consisting of fine, sharp, prominent axial ribs with subdued spiral threads
in the intervals. The ribs are considerably narrower than their interspaces, occasion-
ally slightly varicose, straight or gently concave forward, gently prosocline, and
more strongly bent forward at the adapical suture. There are some 30-35 vague,
irregular spirals on the last-preserved whorl of the holotype. A topotype (G. 42286)
shows a basal disk limited by a fairly sharp carina ; the ribs continue over the disk,
but are much flattened, and there are some 20 spirals similar to those on the spire
whorls. Aperture oval, a little higher than wide ; columellar lip slightly concave,
more callous abapically, where a small auricle is developed.
EOCENE MOLLUSCA FROM NIGERIA 35
REMARKS. This species was compared by Newton with A. affinis (Deshayes) of
the Paris Basin Eocene, but is distinctly more aciculate, the spire angle being smaller.
The whorls are distinctly more convex, and the sutures more deeply sunk, the shell
being more loosely coiled.
Family CALYPTRAEIDAE
Genus CALYPTRAEA Lamarck, 1799
Calyptraea newtoni sp. nov.
(PI. 5, figs. 7a, b)
1922. Calyptraea crepidularis Lamarck : Newton (pars), p. 59.
MATERIAL. The holotype (0.42438) and one topotype (0.42439).
DESCRIPTION. Of moderate size, calyptraeiform, irregularly conic. Protoconch
small, smooth, dextral, Nerita-like, obliquely set. Surface ornamented, at least in
the later stages, with oblique threads bearing papillae, the threads becoming more
nearly parallel to the apertural margin as they approach it. Aperture subcircular,
with margin lying in one plane, septum with very concave edge.
DIMENSIONS. Height 4-7 mm., width 12-8-13-5 mm.
REMARKS. C. crepidularis Lamarck, of the Paris Basin Eocene, with which
Newton identified this form, is much flatter, is subrectangular in outline, has a
markedly eccentric apical region, and does not possess oblique threads bearing
papillae.
Genus TURBOCALYPTRAEA nov.
TYPE SPECIES. T. scabrosa sp. nov.
GENERIC CHARACTERS. Of medium size to moderately small, calyptraeiform,
tending to turbinate, apex markedly eccentric. Whorls strongly convex, with several
spirally disposed rows of short, hollow, forward-directed spines together with
obliquely disposed threads. Aperture as in Calyptraea, subcircular to suboval. A
distinct, sunken, gently concave septum, the margin of which is noticeably concave
in its upper part and gently convex in its lower part, occupies nearly half the aper-
ture. A small, deep umbilicus is developed in the upper part of the septum.
REMARKS. The form, ornament, character of the septum, and presence of an
umbilicus together characterize this new genus.
Turbocalyptraea scabrosa sp. nov.
(PI. 5, figs. 8, ga, b)
1922. Calyptraea crepidularis Lamarck : Newton (pars), p. 59, pi. 4, figs. 18-19.
MATERIAL. A few specimens, including the holotype (0.42437, Newton's fig. 19).
DESCRIPTION. As above.
REMARKS. C. crepidularis Lamarck, from the Eocene of the Paris Basin, with which
Newton identified this species as well as the last, differs markedly in its almost flat
36 EOCENE MOLLUSCA FROM NIGERIA
form and subrectangular outline, its very small spire, its lack of coarse ornament,
and in the characters of the septum.
Genus CREPIDULA Lamarck, 1799
Subgenus CONCAVIMARGO nov.
TYPE SPECIES. Crepidula falconeri Newton.
SUBGENERIC CHARACTERS. Like Crepidula, but teleoconch whorls not in contact ;
surface smooth ; body cavity deep ; aperture oval ; septum rather deeply sunk
and with a cavity extending underneath the inner lip ; edge of septum distinctly
concave.
Crepidula (ConcavimargoJ falconeri Newton «
1922. Crepidula falconeri Newton, p. 58, pi. 2, figs. 13, 130.
MATERIAL. Several specimens, including the lectotype (0.42411, Newton's fig. 13)
here designated.
Family XENOPHORIDAE
Genus XENOPHORA Fischer von Waldheim, 1807
Xenophora nigeriensis (Newton)
(PI. 6, figs. la-c)
1922. Tugurium nigeriense Newton, p. 51, pi. 4, figs. 20-21.
MATERIAL. Numerous specimens, including the lectotype (0.42291, Newton's
fig. 21) here designated.
REMARKS. The presence of numerous agglutinated objects covering most of the
whorl surface indicates that the species is better regarded as a Xenophora ; an
umbilicus as small as the one it possesses may occur in this genus.
Family STROMBIDAE
Genus TIBIA (Bolten MS.) Roding, 1798
Tibia bidigitata (Newton)
1922. Rostellaria bidigitata Newton, p. 12, pi. 4, figs. 8-9.
MATERIAL. Several specimens, including the lectotype (0.41643, Newton's fig. 8)
here designated.
Genus CYRTULOTIBIA nov.
TYPE SPECIES. Rostellaria unidigitata Newton.
GENERIC CHARACTERS. Form somewhat like that of a Tibia with a very short,
inclined siphonal canal ; often developing a strong shoulder on the last whorl and
thus recalling Cyrtulus. Protoconch conic, of two or three smooth, moderately
EOCENE MOLLUSCA FROM NIGERIA 37
convex whorls. Spire like that of Tibia, conic, of five or six gently convex whorls ;
early stages with fine axial riblets, which are orthocline above and opisthocline below
(i.e., concave forwards), crossed by increasingly broad spiral threads ; coarse,
swollen varices developed occasionally. On later spire whorls the axial riblets be-
come obsolete and are represented by accentuated growth lines only. Last whorl
slightly to very strongly shouldered, the more strongly shouldered specimens with a
narrow callous band (an extension of the callus of the posterior sinus) extending back
some one and a half to two whorls along the sutural region. Last whorl smooth
except for spiral threads on the base, rather conic, base very slightly excavated, neck
oblique. Rostrum short, curved to the right. Aperture oval, with a narrow, slit-like
posterior sinus curving back on to the suture ; columellar lip callous, developing a
prominent, raised knob of callus limiting the inner side of the posterior sinus ; a
broad notch to the right of the rostrum is delimited on the right by a short spine.
Outer lip rather thick, not varicose, internally smooth, with no additional spines,
gently parasigmoidal, distinctly opisthocline as a whole.
Remarks. The general form, short inclined rostrum, single abapical labial spine,
the contour of outer lip, broad abapical notch, and extremely strong adapical
parietal callus (extending back along the suture for two or three whorls) readily
distinguish this genus from Tibia.
Cyrtulotibia unidigitata (Newton)
1922. Rostellaria unidigitata Newton, p. 14, pi. 4, figs. 3-7.
MATERIAL. Many specimens, including the lectotype (0.41688, Newton's fig. 3)
here selected.
Genus SEMITEREBELLUM Cossmann, 1889
Subgenus AFRICOTEREBELLUM nov.
TYPE SPECIES. Semiterebellum elongatum Newton.
SUBGENERIC CHARACTERS. Form much like that oT Terebellum, but more narrowly
fusiform. Protoconch consisting of some three to four smooth, moderately convex
whorls, less acute than the shell as a whole. About six spire whorls, at first gently
convex, becoming flatter with growth. Ornament in the earlier stages consisting of
a sharp, fine adapical thread finely crenulated by growth lines, with a narrow,
shallowly excavated band below it, the abapical half of the remaining portion of
the whorls with four to six fine, incised spiral lines ; ornament obsolete on later whorls.
Last whorl with numerous irregular spiral threads (finer and with broader intervals
abapically) on the base, which is only vaguely concave ; neck short, gently swollen.
Rostrum extremely short, its end barely projecting more than the outer lip abapically.
Aperture oval-subtriangular, narrower adapically, with a broad, shallow notch to
the right of the rostrum. Columellar callus thin, especially medially, with a low,
oblique ridge adapically, forming the upper edge of the short posterior sinus which
extends only slightly above and back along the suture. Outer lip opisthocline as a
38 EOCENE MOLLUSCA FROM NIGERIA
whole, forming a strongly projecting, rounded lobe to the right of the abapical
channel, thin, not varicose, internally smooth.
REMARKS. This subgenus differs from Semiterebellum (sensu stricto) in being more
lanceolate, in the less projecting rostrum, in the posterior sinus which does not ascend
partly up the spire, and in the more opisthocline outer lip which is more lobate below.
Semiterebellum (Africoterebellum) elongatum Newton
(PL 5, figs. ioa, b)
1922. Semiterebellum elongatum Newton, p. 17, pi. 2, figs. 14-15.
MATERIAL. Numerous specimens, including the lectotype (0.41762, Newton's
fig. 15) here selected.
Genus AMEKICHILUS nov.
TYPE SPECIES. — Semiterebellum suturocostatum Newton.
GENERIC CHARACTERS. Of small-medium to medium size, having the general form
of Ectinochilus. Protoconch helicoid-trochoid, consisting of about four smooth,
moderately convex whorls. Spire gently cyrtoconoid ; last whorl slightly more than
half the height of the shell. Whorls only slightly convex, sutures distinct ; five to
six spire whorls. An incised line separates a narrow juxtasutural band which is always
crenulated or beaded in the early stages ; later, the band is either smooth or beaded,
and may be vaguely bifid. Last whorl oval, base declivous ; neck short, not swollen.
Rostrum short, scarcely projecting, inclined gently to the right, with a broad, very
shallow notch to its right, the notch limited externally by a short, sharp spine.
Aperture oval, rather small, with a long, narrow, callous posterior sinus which
curves over and slightly down on to the suture of the last whorl, extending only a
short distance back. On those specimens which have a heavily beaded juxtasutural
thread, the upper part of the posterior sinus extends back right up the spire as a
gently convex callous band occupying the lower half of the whorls. Columella
gently concave, smooth, with thick callus developing an even thicker ridge along
the upper side of the posterior sinus. Outer lip thin, varicose, internally smooth, but
with a slight internal thickening, externally strongly varicose, especially at the
posterior sinus, nearly straight and orthocline, slightly convex adapically. Ornament
consisting of a few incised spiral lines with minute pits, widely spaced posteriorly,
often obsolete on the middle of the last whorl, changing to more closely spaced spiral
threads on the base.
REMARKS. This form is shorter and more oval than Semiterebellum, and has a less
projecting rostrum, a distinct juxtasutural band, and an abapical labial spine ; the
posterior sinus does not ascend above the suture of the last whorl. Its closest
relatives seem to be the Ectinochilus and Dientomochilus group of shells, but there
are no obvious varices and the posterior sinus curves over at the suture of the last
whorl. The posterior sinus in Africoterebellum is similar, but in that genus the form
of the shell, ornament and labial spine are quite different.
EOCENE MOLLUSCA FROM NIGERIA 39
Amekichilus suturocostatum (Newton)
1922. Semiterebellum suturocostatum Newton, p. 15, pi. 4, figs. 14-17.
MATERIAL. Many specimens, including the lectotype (0.41757, Newton's fig. 14)
here selected.
Family AMPHIPERATIDAE
Genus EOVOLVA Schilder, 1932
Eovolva nigeriensis (Newton)
(PI. 6, figs. 2, 3)
1922. Amphiperas nigeriensis Newton, p. 18, pi. 3, figs. 14-15.
1932. Eovolva nigeriensis (Newton) : Schilder, p. 212.
MATERIAL. Several specimens, including the lectotype (0.41786, Newton's fig. 14)
here selected.
REMARKS. This is the monotype of Eovolva.
Genus SPHAEROCYPRAEA Schilder, 1927
Sphaerocypraea sudanensis (Schilder)
(PI. 6, figs. 40, b)
1922. Cypraea cf. bowerbanki J. de C. Sowerby : Newton, p. 18, pi. 3, fig. 13.
1929. Sphaerocypraea bowerbankii (J. de C. Sowerby) : Schilder, p. 305.
1932. Eocypraea (Sphaerocypraea) bowerbanki (J. de C. Sowerby) var. sudanensis Schilder,
p. 218.
MATERIAL. A few specimens, including the lectotype (0.41780, Newton's fig. 13)
here selected.
Remarks. In spite of its general similarity to 5. bowerbanki, Schilder (1929) sus-
pected that the Nigerian form was probably specifically distinct. The details of the
fossula cannot be compared as they are not seen in available British specimens, but
the outer lip of the Nigerian form is somewhat broader and is distinctly more callous
and margined, and the shell was evidently a little more globose. It seems advisable
to regard the Bende Ameki specimens as constituting a distinct species for which
Schilder's name must be adopted, in spite of its unsuitability.
Family NATICIDAE
Genus NEVERITA Risso, 1826
Neverita amekiensis sp. nov.
(PI. 6, figs. 50, b)
1922. Neverita cf. calvimontana (Deshayes): Newton, p. 55, pi. 5, figs. 16-17.
MATERIAL. The holotype (G. 42383).
40 EOCENE MOLLUSCA FROM NIGERIA
DESCRIPTION. Of small-medium size, having the form of Polinices rather than
Neverita, a little less flattened than is usual in the latter genus. Protoconch conic,
very low, of about two smooth, gently convex whorls. Last whorl forming about
four-fifths of the height of the shell. Spire consisting of about two and a quarter
almost flat, smooth whorls with fine linear sutures. Last whorl very large, rather
flattened adapically, its flank moderately sharply rounded ; base declivous. Aper-
ture semilunar, not much produced to the right. Columella straight, its callus very
heavy, a solid, gently convex plug almost, but not quite, filling the umbilicus.
REMARKS. In N. calvimontana (Deshayes), from the Eocene of the Paris Basin,
the umbilical callus is less extensive and solid, the whorls are more distinctly convex,
and the sutures are not quite linear.
Genus SINUM (Bolten MS.) Roding, 1798
Sinum africanum Newton
(PI. 6, figs. 6a-c)
1922. Sinum africanum Newton, p. 57, pi. 4, figs. 12-13.
MATERIAL. Several specimens, including the lectotype (0.42406, Newton's fig. 12)
here selected.
Sinum nigeriense sp. nov.
(PI. 6, figs. 7«, b)
1922. Sinum cf. clathratum (Gmelin): Newton, p. 56, pi. 2, figs. 16-17.
MATERIAL. Several specimens, including the holotype (0.42390, Newton's fig. 16).
DESCRIPTION. Of small-medium size, not very thin-shelled, not very auriform,
outline more like that of Polinices. Protoconch conic, very low, of two smooth,
slightly convex whorls with a small nucleus. One and a quarter almost flat spire
whorls with 13-15 wavy, fine spiral threads the course of which is slightly deflected
at each growth line ; up to three microscopic spirals developed in the intervals on
the penultimate whorl. Last whorl very large, occupying most of the height of the
shell, adapical portion rather extensive and flattened, flanks rounded, base declivous ;
ornament as on the spire whorls, but primary threads more numerous (approximately
40 in number). Aperture rounded-subquadrate, somewhat produced abapically and
to the right. Umbilical callus narrow, but distinct, leaving a very small umbilical
opening. Outer lip markedly prosocline and gently convex.
REMARKS. S. clathratum is more auriform and compressed, has a lower spire and
a more ample aperture, and does not have the adapically flattened whorls which in
the new species produce a rather conic appearance.
Family CYMATIIDAE
Genus VARICOHILDA nov.
TYPE SPECIES. Hilda turriculata Newton.
GENERIC CHARACTERS. Of small-medium size, having the general form of Hilda.
EOCENE MOLLUSCA FROM NIGERIA 41
Protoconch not well preserved, evidently smooth and naticoid. Last whorl consti-
tuting about half the height of the shell. Spire consisting of four to five whorls which
are about half as high as wide and carry a quite sharp median angulation ; ornament
consisting of narrow, rather widely-spaced, straight, orthocline or slightly opistho-
cline axial ribs crossed by fairly prominent spiral threads which are a little broader
than their intervals ; no varices on spire whorls. Last whorl ornamented like the
spire whorls, with the addition of fine intercalary spiral threads near the shoulder
and a very strong varix 240° back from the outer lip ; base excavated abapically,
the ribs becoming fainter but the spiral ornament persisting ; neck short, swollen,
inclined to the left. Aperture narrow and parallel-sided, rather like that of Anachis,
a fairly broad, oblique adapical part limited by a spiral parietal fold ; a fairly short,
narrow siphonal canal, slightly inclined to the left, has a deep lateral notch at the
end. Siphonal fasciole swollen. Outer lip almost orthocline, with a very strong varix
slightly behind its sharp edge, internally thickened and with long lirae. Columellar
lip straight and vertical, with three strong, spiral columellar folds, forming angular
junctions with the parietal lip and with the siphonal canal ; callus well-developed,
spreading a little over the base, distinctly limited, becoming slightly detached by
the siphonal fasciole, with numerous knobs and transverse wrinkles between its
outer margin and the columellar and parietal folds.
REMARKS. Although generally resembling Hilda, this genus differs in having a
slightly shorter spire, angular whorls, a strong varix in addition to the labral varix
on the last whorl, a narrower and more parallel-sided aperture, the canal inclined to
the left, and three strong columellar folds in addition to wrinkles and knobs on the
inner lip.
Varicohilda turriculata (Newton)
(PI. 7, figs. la-c)
1922. Hilda turriculata Newton, p. 29, pi. 4, figs. 24-25.
MATERIAL. Many specimens, including the lectotype (0.41971, Newton's fig. 24)
here selected.
v
Family MURICIDAE
Genus HEXAPLEX Perry, 1811
Subgenus PAZIELLA Jousseaume, 1880
Hexaplex (Paziella) bendeica sp. nov.
1922. Poirieria cf. calcitrapa (Lamarck) : Newton, p. 31, pi. 3, figs. 22-23.
MATERIAL. The holotype (0.41983, Newton's fig. 22).
DESCRIPTION. Of small-medium size, fusiform, ribs aligned in seven irregular axial
series, each rib being, on the last three whorls, slightly behind that on the whorl
above. Protoconch not preserved. Last whorl forming about half the height of the
shell. Spire conic, consisting of five to six whorls which are distinctly angulated
medially, irregularly flattened and shelving above, subcylindrical or even slightly
GEOL. Ill, 2. c
42 EOCENE MOLLUSCA FROM NIGERIA
inturned below, with linear, wavy sutures. Adapical shelf with two or three very
vague spirals near the middle ; one spiral thread on the angulation, and one close
to the abapical suture, the latter on later whorls with a faint spiral just above it.
Ribs nodular and subspinose at the shoulder on early whorls, a little narrower than
their intervals, becoming considerably narrower than the intervals with growth and
also developing short upturned spines at the shoulder, the spines being channelled
on the forward-facing side. Last whorl inflated, base well excavated, neck moderately
long ; siphonal fascicle bulging, carrying widely-spaced scales. Three primary threads
and vague intercalaries on the flank of the last whorl. Aperture oval, with no distinct
adapical channel, with a moderately long, narrow, gently curved siphonal canal
which is gently inclined to the left. Columella gently excavated, smooth, twisted,
and with a vague fold at the beginning of the canal. Columellar lip callous, not
widely spread, becoming detached abapically, leaving a small false umbilicus be-
tween it and the siphonal fascicle. Outer lip thin, with a varix close behind it, gently
parasigmoidal and definitely prosocline adapically, with a deep lateral notch at the
spine on the shoulder, internally thickened, dentate below the notch. Growth lines
serrate on forward-facing side of varices.
REMARKS. H. (P.) calcitrapa has a much more inflated last whorl and a lower spire.
Poirieria has five, not seven, axial rows of varices.
Genus PTERYNOTUS Swainson, 1833
Pterynotus newtoni sp. nov.
1922. Pteropurpura cf. tricarinata (Lamarck) : Newton, p. 30, pi. 4, figs. 26-27.
MATERIAL. The holotype (G.4I982, Newton's figs. 26-27).
DESCRIPTION. Of medium size, fusiform, markedly triangular when viewed from
above on account of three rows of lamellar, non-spinose varices, each varix being
slightly behind the corresponding one on the preceding whorl. Protoconch (nucleus
missing) apparently rather tectiform, of about three smooth, gently convex whorls.
Last whorl forming slightly more than half the height of the shell. Spire conic,
consisting of nearly five convex whorls with linear, undulatory sutures. Early whorls
carry two ribs, later whorls only one, between the varices ; ribs nodular and crossed
by muricate spiral threads, three orders of which are present on the last whorl ;
serrate, rather widely-spaced growth threads are also present. Last whorl inflated ;
base well excavated, neck long and straight except for the protuberant, tubular
siphonal fasciole. Aperture oval, with an abapical channel and a long, narrow
siphonal canal which is slightly longer than the height of the aperture, inclined to
the left but vaguely curved towards the right, and then curved well back at the tip.
Columella gently concave, twisted at the start of the canal. Inner lip callous, a little
wider adapically, rather detached medially, well detached abapically, smooth
except for a spiral ridge limiting the adapical channel. Outer lip with a broad,
leaf-like, non-spinose varix, orthocline, its edge thin and dentate, internally thickened
and dentate. Growth lines serrate on forward-facing side of varices.
EOCENE MOLLUSCA FROM NIGERIA 43
REMARKS. Pterynotus (Pteropurpura) tricarinatus (Lamarck) has the varices up-
turned and spinose adapically.
Family BUCCINIDAE
Genus BENDEIA nov.
TYPE SPECIES. Liomesus africanus Newton.
GENERIC CHARACTERS. Of small-medium size, buccinoid, intermediate in general
form between Cominella and Strepsidura, oval-conic. Protoconch naticoid, of two
to two and a half smooth, moderately convex whorls. Last whorl forming slightly
less than three-quarters of the height of the shell. Spire consisting of about two and
a half gently convex whorls with conspicuous, deep sutures, slightly stepped ; two
incised spiral lines close to the adapical suture define two raised cords. Last whorl
inflated-oval, ornamented like the spire whorls, base well excavated and with 11-13
spiral threads which are narrower and stronger abapically ; neck moderately long
and swollen. Aperture rather narrowly oval, with an adapical channel, and with a
siphonal canal of moderate length which is inclined to the left and notched. Colu-
mellar callus not thick or extensive ; columella with a fairly strong fold limiting
the canal, and a series of wrinkles where the spiral threads of the base pass under
the columellar callus. Siphonal fasciole only moderately swollen, carrying five to
eight longitudinal threads, limited above by a strong, sharp, raised thread. Outer
lip thin, internally smooth, almost orthocline, slightly prosocline adapically.
REMARKS. Although superficially resembling Liomesus, this genus is more strep-
siduriform (i.e. it has a shorter spire, and is more produced abapically), has a
longer and more inflected canal, a less callous inner lip, and a distinct siphonal
fasciole limited above by a fine, raised carina ; moreover, spiral ornament is present
on the upper part of the whorls, the base of the last whorl is more excavated, the
aperture is narrower, and there is a distinct columellar fold in an abapical position.
The relationships evidently lie more with Cominella than with Liomesus or Strepsidura.
Bendeia africana (Newton)
(PI. 7, figs. 2a, b)
1922. Liomesus africanus Newton, p. 38, pi. 3, figs. 20-21.
MATERIAL. Several specimens, including the lectotype ^.42096, Newton's fig. 21)
here selected.
Genus LACCINUM nov.
TYPE SPECIES. Athleta lugardi Newton.
GENERIC CHARACTERS. Attaining a large size, thick-shelled, conic to buccinoid, with
a low spire. Protoconch (worn in available specimens) evidently not large and
bulbous as in many Volutidae. Shell completely smooth except for growth lines
which are somewhat accentuated on the siphonal fasciole, although extremely vague ;
fine spirals can sometimes be distinguished. Last whorl forming about ten-thirteenths
44 EOCENE MOLLUSCA FROM NIGERIA
(younger specimens) to seven-eighths (larger specimens) of the height of the shell.
Spire distinctly coeloconoid, composed of about five flattened, very gently convex
whorls with distinct, linear sutures, relatively narrower in later stages of growth.
Last whorl very large, its shoulder even more sharply rounded in large specimens
than in juveniles, flank subcylindrical, base slightly excavated, neck moderately
long and swollen by the siphonal fasciole. Aperture narrowly oval, with a broad,
flat channel situated adapically at the shoulder, and with a fairly short, poorly-
defined siphonal canal which is slightly inclined to the left and deeply notched.
Outer lip thin, smooth internally, orthocline as a whole, gently bisinuous. Columella
gently concave, vaguely bent at its junction with the canal ; inner lip with a thick,
moderately wide layer of callus which becomes slightly detached abapically at the
extensive siphonal fasciole, and which develops a large, protruding knob of callus
beside the adapical channel. No columellar folds.
REMARKS. The above characters indicate that this genus is not related to Athleta,
nor even a volutid. It is evidently a buccinid, and seems best placed near Lacinia,
from which it differs in its more cylindrical last whorl, the lack of an umbilicus,
and its less ample aperture and better defined siphonal canal.
Laccinum lugardi (Newton)
1922. Athleta lugardi Newton, p. 25, pi. 5, figs. 1-3.
MATERIAL. Several specimens, including the lectotype (0.41860, Newton's fig. i)
here selected.
Genus JANIOPSIS Rovereto, 1899
Janiopsis nigeriensis Newton
1922. Janiopsis nigeriensis Newton, p. 41, pi. 4, figs. 22-23.
MATERIAL. Several specimens, including the lectotype (0.42131, Newton's fig. 22)
here selected.
Family VOLEMIDAE
Genus PSEUDOMAZZALINA nov.
TYPE SPECIES. Bulbifusus nigeriensis Newton.
GENERIC CHARACTERS. Of large-medium size, rather thin-shelled, inflated-fusiform,
entirely smooth. Protoconch unknown. About six moderately convex spire whorls
which are broadest a little below the middle, height about two-fifths of the width.
Sutures linear. Last whorl forming one-half to three-fifths the height of the shell,
inflated, base evenly excavated, neck moderately long, straight, vertical. Aperture
oval, with a narrow adapical channel, vaguely constricted abapically where it is
extended into a moderately oblique and wide, notched siphonal canal the length of
which is about one-half the height of the aperture proper. No siphonal fasciole.
Columella gently excavated, rather vaguely bent at the beginning of the canal, with
a narrow, thin layer of callus, without columellar folds. Outer lip thin, strongly
EOCENE MOLLUSCA FROM NIGERIA 45
convex in its median and anterior parts, receding strongly adapically at an angle of
about 45° and becoming nearly orthocline close to the suture. Some 10-12 irregular,
elongate lirae are developed well inside the aperture in its upper half.
REMARKS. Compared with Mazzalina (of which Bulbifusus is a synonym) this
genus is less inflated and more fusiform, and has a higher spire, a narrower aperture
and canal, no columellar folds, no spiral ornament on the base, and an outer lip
which is deeply excavated adapically. Levifusus is distinctly ornamented and has a
longer canal. Sycostoma is less fusiform and has heavier columellar callus ; its
outer lip is less excavated adapically.
Pseudomazzalina nigeriensis (Newton)
1922. Bulbifusus nigeriensis Newton, p. 35, pi. 4, figs. 1-2.
MATERIAL. Several specimens, including the lectotype (0.42057, Newton's fig. 2)
here selected.
Family FUSINIDAE
Genus CLAVILITHES Swainson, 1840
Subgenus AFRICOLITHES nov.
TYPE SPECIES. Rhopalithes africanus Newton.
SUBGENERIC CHARACTERS. Of large-medium size, fusiform. Protoconch not pre-
served. Last whorl forming about half the height of the shell. Spire conic (apex rather
cyrtoconoid), of about eight gently convex whorls, with a slight spiral depression
just below the suture. First five or six whorls with moderately fine spiral threads
crossing strong nodular ribs which are not in alignment from whorl to whorl. In
the later stages the ribs become obsolete adapically, and on the last two whorls are
absent completely, the spiral threads also becoming feebler. Base of last whorl
excavate ; neck long, straight, vertical. No siphonal fascicle. Imperforate. Aper-
ture oval, with a small adapical channel and a long, straight siphonal canal only
slightly inclined to the left. Outer lip thin, broadly concave, internally smooth.
Columellar callus not widely spread, detached externally. Columella straight, joining
the canal without any twist. No columellar folds.
REMARKS. Clavilithes is considerably less fusiform. Rhopalites has rather shouldered
whorls, a more oblique canal, and a heavier adapical apertural callus, and it tends
to have a siphonal fascicle and small pseudumbilicus. Chiralithes, from the Upper
Eocene of Peru, is somewhat similar, but has a considerably broader aperture.
Perulithes, also from the Upper Eocene of Peru, has a higher spire, and its later
whorls are quite smooth.
Clavilithes (Africolithes) africanus (Newton)
1922. Rhopalithes africanus Newton, p. 32, pi. 2, figs. 9-10.
MATERIAL. Several specimens, including the lectotype (0.42004, Newton's fig. 10)
here selected.
46 EOCENE MOLLUSCA FROM NIGERIA
Genus LEUCOZONIA Gray, 1847
Leucozonia pseudominax sp. nov.
(PI. 6, fig. 8)
1922. Cornulina minax (Solander) : Newton, p. 34, pi. 3, figs. 6-7.
MATERIAL. The holotype (G. 42052, Newton's figs. 6-7).
DESCRIPTION. Of medium size, similar in form to Cornulina minax. Protoconch
not preserved. Last whorl forming a little more than half the height of the shell.
Probably about four spire whorls forming a conic spire ; whorls angulated at about
one-third of their height, cylindrical or with sides inclined slightly inwards abapically,
sloping and gently concave adapically. Ornament of spiral threads crossing vague,
broad ribs which form blunt, slightly upturned spines on the keel. Last whorl large,
base excavated, lower portion missing ; spines well developed and protruding both
on the main keel and on a subsidiary one developed abapically, with a narrow slit
on their forward-facing side and thus evidently hollow. Aperture rounded-oval, with
a small adapical channel limited below by a small spiral ridge ; evidently with a
fairly short siphonal canal distinctly inclined to the left. Columella concave, with at
least two strong columellar folds abapically. Outer lip thin, gently parasigmoidal,
orthocline as a whole, coarsely but vaguely fluted internally. The broken lower end
of the columella is solid.
REMARKS. Apart from the fact that the ornament on the lower part of the last
whorl differs from that of Cornulina minax in consisting of widely-spaced, sharp
spiral threads instead of numerous closely-spaced threads of several orders, the
presence of strong columellar folds indicates that the Nigerian form is not a
Cornulina. Since the genus Fascioplex has an umbilicus, a lower spire, and a last
whorl which is less inflated adapically, the Nigerian form seems best placed in the
genus Leucozonia.
Family VOLUTIDAE
Genus VOLUTOCORBIS Ball, 1890
Volutocorbis multispinosa (Newton)
1922. Volutospina multispinosa Newton, p. 28, pi. 3, figs. 3-5.
MATERIAL. Numerous specimens, including the lectotype (0.41949, Newton's
fig. 3) here selected.
REMARKS. This species is now removed from Volutospina, as the more scabrous
ornament, less spinose ribs, and more oval form indicate that it is a Volutocorbis.
Genus BENDELUTA nov.
TYPE SPECIES. Volutospina conicoturrita Newton.
GENERIC CHARACTERS. Of medium size, not very thick-shelled, in general form
somewhat similar to Volutospina and related genera. Protoconch conical, not large, of
EOCENE MOLLUSCA FROM NIGERIA 47
about three smooth, moderately convex whorls. Spire conic. Four flat-sided spire
whorls, a little wider than high, ornamented with rather weak and narrow, straight
axial ribs crossed by low, flat spiral ribbons ; whorls shouldered above, shoulder
bearing increasingly prominent but short, upturned, hollow spines, both shoulder and
horizontal sutural ledge free of spiral ornament ; sutures linear, undulatory. Last
whorl large, inflated, with a second row of short, laterally directed spines at the level
of the adapical end of the aperture, this level forming the widest part of the whorl ;
ribs dying out below and spiral ornament obsolete on flanks ; base moderately
sharply excavated at its junction with the rather broad neck at the middle of which
a vague spiral depression demarcates the slightly swollen siphonal fascicle ; base,
neck and siphonal fascicle with irregular spiral threads. Aperture elongate, rather
irregular in shape on account of the constriction of the shell above the neck and on
the flank, rather deeply notched below, and with a moderately short, oblique,
poorly-differentiated siphonal canal. Columella oblique, well set off above, slightly
convex, its upper half with three spiral folds of which the upper one is a little weaker
and is a little closer to the median fold than is the lower one. Parietal callus spread
adapically over half the ventral surface of the last whorl as a very thin glaze, almost
absent medially and abapically. Lip straight and orthocline as a whole, slightly
receding and sinuous abapically, strongly prosocline on the narrow shoulder, thin-
edged, internally smooth.
REMARKS. The more globose last whorl (the widest part of which bears the lower
row of spines), the thicker neck, and the shorter and more irregularly shaped aperture,
together with the other characters mentioned above, readily distinguish this from
Volutospina.
Bendeluta conicoturrita (Newton)
1922. Volutospina conicoturrita Newton, p. 27, pi. 3, figs. 1-2.
MATERIAL. Several specimens, including the lectotype (0.41901, Newton's fig. i)
here selected.
Family OLIVIDAE'
Genus PSEUDOLIVA Swainson, 1840
Subgenus BUCCINORBIS Conrad, 1865
Pseudoliva (Buccinorbis) kitsoni (Newton)
1922. Buccinorbis kitsoni Newton, p. 37, pi. 3, figs. 16-19.
MATERIAL. Many specimens, including the lectotype (0.42069, Newton's fig. 16)
here selected.
REMARKS. The umbilicus is mostly covered by the umbilical callus, but the
presence of such a distinct depression as is present in this species is more a feature
of Buccinorbis than of Pseudoliva.
48 EOCENE MOLLUSCA FROM NIGERIA
Family STREPSIDURIDAE
Genus STREPSIDURA Swainson, 1840
Subgenus STREPSIDUROPSIS nov.
SUBGENERIC CHARACTERS. Like Streps^d^lra, but with one strong spiral fold on
the columella at the beginning of the canal, and above it a convex pad on which
are grouped six more folds ; anterior part of shell more constricted, siphonal canal
a little longer and more twisted ; outer lip internally thickened and weakly crenulated.
REMARKS. Mazzalina has no carinate siphonal fascicle, and its canal is shorter
and less oblique.
Strepsidura (Strepsiduropsis) spirata Newton
(PL 7, ng. 3)
1922. Strepsidura spirata Newton, p. 33, pi. 3, figs. 24-25.
MATERIAL. Many specimens, including the lectotype (0.42008, Newton's fig. 25)
here selected.
Family CANCELLARIIDAE
Genus SVELTIA Jousseaume, 1887
Subgenus AFRICOSVELTIA nov.
TYPE SPECIES. Cancellaria multiplicis Newton.
SUBGENERIC CHARACTERS. Of small-medium size, having the form of Sveltia or
Sveltella, but with a rather shorter spire. Protoconch unknown. Spire conic ; three
to four spire whorls which are convex and tend to be slightly subangular just above
the middle ; sutures linear, undulatory. Ornament consisting of strong, widely-
spaced spiral threads crossing solid axial ribs which are narrower than their intervals ;
ribs fairly strong, becoming very solid on the last whorl, gently prosocline. Last
whorl oval, narrower abapically, base declivous. Aperture oval, narrower and
gently emarginate (not channelled) abapically. Columella straight, with three
columellar folds of which the uppermost is the strongest. Columellar callus moder-
ately widely spread adapically, narrower and detached abapically, leaving a narrow
but distinct pseudumbilicus which is limited externally by a broad, vague swelling
rather than by a siphonal fasciole. Lip straight to gently concave, slightly prosocline,
thin-edged, with a strong varix close behind it, lirate and slightly thickened
internally.
REMARKS. The aperture has no siphonal notch as in Cancellaria. Compared with
Sveltia s.str., in the new subgenus the spire is lower, there is a distinct pseudum-
bilicus, and there are three (not two) columellar folds. Sveltella has no noticeable
umbilicus, a higher spire, and only two columellar folds.
EOCENE MOLLUSCA FROM NIGERIA 49
Sveltia (Africosveltia) multiplicis (Newton)
(PL 7, figs. 4a-c)
1922. Cancellaria multiplicis Newton : p. 42, pi. 3, fig. 26.
MATERIAL. The holotype (0.42137).
Genus BONELLITIA Jousseaume, 1887
Subgenus ADMETULA Cossmann, 1889
Bonellitia (Admetula) amekiensis sp. nov.
(PL 7, figs. 5«-c)
1922. Bonellitia cf. evulsa (Solander) : Newton, p. 44, pi. 5, figs. 14-15.
MATERIAL. The holotype (0.42171, Newton's fig. 15) and several other specimens.
DESCRIPTION. Small, like Admete and Admetula in form. Protoconch unknown.
Last whorl slightly less than half the height of the shell. Spire conic, consisting of
about four strongly convex whorls with deep, linear, undulatory sutures. Ornament
consisting of solid axial ribs, practically straight and gently prosocline, equal to or
slightly narrower than their intervals, occasionally varicose, crossed by spiral threads.
Last whorl swollen-ovate, narrower abapically, base declivous and with the ribs
feebler. Aperture rounded-oval, not notched abapically, but with a broad, slightly
emarginate channel inclined to the left. Columella vertical, with three well-developed
columellar folds of which the lower two are the more closely spaced. Parietal callus
thin ; columellar callus narrow, its outer edge separated from the region of the
neck by a well-marked, linear, vertical depression, but not by an umbilicus. No
siphonal fascicle. Outer lip fairly straight, gently prosocline, blunt, limited behind
by a varix, internally thickened and lirate, the lirae continuing over the floor of the
aperture.
REMARKS. Although Admetula is placed in the synonymy of Bonellitia by Wenz,
the name seems worth retaining for those forms wj,th less muricate ornament. The
Nigerian form differs from B. (A.} evulsa in its more oval-conic outline and less
inflated form, its relatively finer and more numerous spiral threads, and its more
callous and more detached columellar lip.
Subgenus AFRICOSTOMA nov.
TYPE SPECIES. Trigonostoma decorata Newton.
SUBGENERIC CHARACTERS. Small, having the general form of a Bonellitia except
for the marked, canaliculate ramp behind the shoulder. Protoconch naticoid, of
two smooth, convex whorls. Last whorl forming slightly less than half the height
of the shell. Spire conic, of two and a half subcylindrical, gently convex whorls
which are slightly inturned below and have a sharp, upturned, undulatory carina
on the shoulder angle ; ramp horizontal as a whole, gently concave, smooth except
5o EOCENE MOLLUSCA FROM NIGERIA
for irregular continuations of the ribs. Ornament consisting of delicate, gently
prosocline axial ribs crossed by spiral threads, both being narrower than their
intervals ; the occasional very strong varices develop even stronger lobes on the
upturned carina than do the ribs. Last whorl large, slightly convex abapically,
ornamented like the spire whorls. No neck or siphonal fascicle. Aperture oval-
pyriform, wider adapically, slightly emarginate abapically (not notched), with a
slight projection at the shoulder carina. Columella vertical, with a double fold
abapically (limiting the very short, oblique siphonal canal), and with a well-separated
median fold. Parietal callus thin and not widely spread above, thicker and tending
to be slightly detached below. Lip straight, gently prosocline, thin-edged but with
a broad, strong varix just behind it, internally thickened and lirate.
REMARKS. The absence of an umbilicus together with other characters described
above indicate that this species is not a Trigonostoma ; it seems best regarded as
belonging to a new subgenus of Bonellitia.
Bonellitia ( Ajricostoma) decorata (Newton)
(PI. 7, figs. 6a-c)
1922, Trigonostoma decorata Newton, p. 43, pi. 5, figs. 18-19.
MATERIAL. Several specimens, including the lectotype (0.42138, Newton's fig. 18)
here selected.
Family CONIDAE
Genus CONUS Linne, 1758
Subgenus LEPTOCONUS Swainson, 1840
Conus (Leptoconus) amekiensis sp. nov.
(PI. 7, figs. 7«-c)
1922. Conospirus cf. parisiensis (Deshayes) : Newton, p. 24, pi. 3, figs. 8-9.
MATERIAL. Several specimens, including the holotype (0.41837, Newton's fig. 9).
DESCRIPTION. Of small-medium size, biconic. Protoconch rather tectiform, of three
smooth, slightly convex whorls. Last whorl forming at least five-eighths of the height
of the shell. Spire gently coeloconoid, of five to five and a half whorls with small
nodes on an angulation close to the abapical suture ; the narrow portion below the
angulation vertical or sloping slightly inwards and causing a slightly imbricate
appearance ; upper part flattened or vaguely concave, shelving, with four or five
spiral threads on its abapical two-thirds, the threads being slightly crenulated where
crossed by the deeply concave growth-lines. Last whorl inverted-conic, shoulder
with small, low nodes, base slightly excavated and neck slightly swollen ; coarse,
irregular spiral ribbons, which have narrow intervals and are fainter adapically,
are vaguely beaded where crossed by low, flat ribs extending from the small nodes
on the shoulder ; the ribs are of about the same width as their intervals. Aperture
EOCENE MOLLUSCA FROM NIGERIA 51
narrow and parallel-sided, vaguely constricted near the abapical end, emarginate
but not notched terminally.
REMARKS. Conus parisiensis, with which Newton compared this species, is more
produced abapically and has compound crenulations on the uppermost thread of the
spire whorls (not small nodes on the lower angulation) ; its spiral ornament is not
beaded and is restricted to the abapical half of the last whorl.
Family TURRIDAE
Genus EOPLEUROTOMA Cossmann, 1889
Eopleurotoma nigeriensis Newton
1922. Eopleurotoma nigeriensis Newton, p. 23, pi. 2, figs. 11-12.
MATERIAL. A few specimens, including the lectotype (0.41829, Newton's fig. 12)
here selected.
Genus TURRICULA Schumacher, 1817
Subgenus SURCULA H. & A. Adams, 1853
Turricula (Surcula) africana (Newton)
1922. Surcula africana Newton, p. 22, pi. 2, figs. 6-8.
MATERIAL. Several specimens, including the lectotype (0.41809, Newton's fig. 7)
here selected.
Genus SURCULITES Conrad, 1865
Subgenus CLINURA Bellardi, 1875
Surculites (Clinura) ingens (Mayer-Eymar)
1922. Surcula ingens (Mayer-Eymar) : Newton, p. 20, pi. 2, figs. 3-5 (cum syn.).
MATERIAL. Two specimens.
Genus MITRELLOTURRIS nov.
TYPE SPECIES. Asthenotoma (Endiatoma) casteri Chavan.
GENERIC CHARACTERS. Having the general form of Mitrella, elongate oval-conic
to rather turriculate-conic. Protoconch unknown. About six spire whorls (only
four preserved), which are flat-sided, their height a little more than half their width.
Sutures linear, slightly stepped. A narrow, flat, slightly raised juxtasutural band is
limited abapically by a vague, narrow, spiral depression ; whorls otherwise smooth.
Last whorl about two-fifths of the height of the shell, narrowly oval, base narrowly
and gently excavated, the neck of moderate length and swollen. Base and neck with
spiral threads becoming increasingly stronger and more closely spaced abapically.
52 EOCENE MOLLUSCA FROM NIGERIA
Aperture narrow and parallel-sided, with a short, distinct siphonal canal notched at
the end. Columellar callus not extensive, smooth. Small adapical channel present.
Outer lip (broken) evidently thin, not varicose, internally smooth, gently convex
abapically, orthocline or slightly prosocline at the suture, with a broad, shallowly
V-shaped sinus at a position corresponding to two-thirds of the height of the spire
whorls. The two last whorls with a very vague angulation of the surface at the
apex of this sinus.
REMARKS. Although obviously related to Asthenotoma and Endiatoma, this genus
differs in the complete lack of axial ornament, the reduced spiral ornament, the
smooth columella, and the absence of folds inside the outer lip.
Mitrelloturris casteri (Chavan)
(PI. 7, figs. 8a, b ; PL 8, fig. i)
1952. Asthenotoma (Endiatoma) casteri Chavan, p. 80, text-fig.
MATERIAL. The holotype (0.69600).
Genus AMEKICYTHARA nov.
TYPE SPECIES. Cominella douvillei Newton.
GENERIC CHARACTERS. Of small-medium size, form resembling that of the
Cythara group of the Turridae. Protoconch naticoid, consisting of two and a half
smooth, convex whorls. Last whorl forming four-sevenths of the height of the shell.
Spire conic, consisting of three to four gently convex whorls which are moderately
angulated at about two-thirds of their height ; sutures linear. Ornament delicately
cancellate, Ficus-like, with tiny crenulations at the intersections of the axial and
spiral elements. Last whorl oval, gently angulated adapically like the spire whorls
and similarly ornamented ; base declivous and gently excavated at the origin of
the moderately short neck ; axial ornament obsolete and spiral ornament irregular
abapically. Aperture narrowly oval, with a short siphonal canal which is gently
inclined to the left and flares a little at its deeply emarginate end. Columella gently
concave, with one prominent fold (well inside the aperture) at half the height of the
aperture proper. Columellar lip with thin callus which is moderately widely spread
adapically but narrow abapically. No siphonal fasciole. Imperforate. Outer lip
thin at edge, moderately convex medially, slightly concave at a level correspond-
ing with that of the columellar fold, receding adapically, and with a very shallow,
rounded sinus adjacent to the suture ; thickened internally and with prominent,
short lirae.
REMARKS. The above characters indicate that the species is a member of the
Turridae. Cominella, in which genus Newton placed the species, is a buccinid genus
with fundamentally different characters.
53
Amekicythara douvillei (Newton)
(PI. 8, figs. 2a-c)
1922. Cominella douvillei Newton, p. 39, pi. 2, figs. 18-19.
MATERIAL. Several specimens, including the lectotype (0.42100, Newton's fig. 19)
here selected.
LAMELLIBRANCHIA
Family NUCULIDAE
Genus NUCULA Lamarck, 1799
Nucula costaeimbricatis Newton
1922. Nucula costaeimbricatis Newton, p. 73, pi. 6, figs. 10-11.
MATERIAL. The holotype (£,.48096, Newton's figs. 10-11).
Family GLYCYMERIDAE
Genus AFRICARCA nov.
TYPE SPECIES. Glycimeris nigeriensis Newton.
GENERIC CHARACTERS. Of small-medium size, moderately thick-shelled, moder-
ately strongly inflated, oval-subtrapezoidal in outline. Beaks submedian, almost
orthogyrous. Hinge-line almost straight, forming only about two-thirds of the length
of the shell. Greatest height anterior to the median line, at about two-thirds of the
length, the shell being slightly produced antero-ventrally. Cardinal area inclined
mainly at an angle of about 45° to the surface of the hinge-plate. Hinge-plate fairly
solid, straight medially, gently arched downwards on the flanks. Teeth taxodont,
numerous, anterior ones slightly more numerous than the posterior, the eight most
anterior and six most posterior ones larger and oblique ; a minute gap between
anterior and posterior series. Muscle scars fairly large, not buttressed. No byssal
sinus. Numerous low, square-cut ribs. Surface without carina. Ventral margin
internally fluted.
REMARKS. Although the hinge is somewhat like that of Glycymeris, the sub-
trapezoid (rather Arcopsis-like) outline and other characters described above readily
distinguish this genus.
Africarca nigeriensis (Newton)
(PI. 8, figs. 3a, 6)
1922. Glycimeris nigeriensis Newton, p. 72, pi. 8, figs. 8-10.
MATERIAL. Several specimens, including the lectotype (1,48441, Newton's fig. 8)
here selected.
54 EOCENE MOLLUSCA FROM NIGERIA
SUPPLEMENTARY CHARACTERS. Beaks very slightly anterior to the middle line,
the extreme tips tending to be slightly prosogyrous. Hinge-line sloping vaguely
downwards and outwards away from the beak. Dorso-lateral angles obtusely
rounded, the anterior a little the more angular. Anterior end gently convex, joining
the ventral margin in a well-rounded curve. Ventral margin gently convex. Posterior
end obliquely truncated, slightly convex, joining the ventral margin in a rounded,
blunt angle. Cardinal area moderately narrowly triangular, placed symmetrically
beneath the beak, slightly concave where abutting against the slight projecting
ridges constituting the dorsal margin of the shell, carrying fairly numerous ridges,
slightly wider than their intervals medially and narrower than their intervals on the
flanks, at right angles to the hinge-line. Adult with 24 anterior and 20 posterior
teeth, distally converging ventrally ; median 30 teeth all small, especially medially.
Surface with nearly 70 rather low, square-cut, finely beaded ribs of about the same
width as their intervals, rather finer on the flanks. Surface evenly convex.
Family NOETIIDAE
Genus PROTONOETIA MacNeil, 1938
Protdnoetia nigeriensis (Newton)
1922. Anadara nigeriensis Newton, p. 70, pi. 8, figs. 4-7.
1938. Protonoetia nigeriensis (Newton) : MacNeil, p. 25, pi. 4, figs. 1-3.
MATERIAL. Several specimens, including the lectotype (L. 48544, Newton's fig. 4)
here selected.
REMARKS. This is the type species of Protonoetia.
Genus ARCOPSIS von Koenen, 1885
Arcopsis africana (Newton)
(PI. 8, fig. 4)
1922. Fossularca africana Newton, p. 68, pi. 8, figs. 14-17.
MATERIAL. Several specimens, including the lectotype (L. 481 13, Newton's fig. 14)
here selected.
REMARKS. Fossularca is a synonym of Arcopsis.
Genus RECTANGULARCA nov.
TYPE SPECIES. Striarca africana Newton.
GENERIC CHARACTERS. Of small-medium size, not very thick-shelled, rather
strongly inflated, subrectangular in outline. Beaks median, practically orthogyrous,
the extreme tips slightly opisthogyrous. Cardinal area narrowly triangular, the
portion anterior to the beak slightly the shorter ; with one fine chevron-shaped
groove very close to the two shorter sides, the remainder of the surface with
EOCENE MOLLUSCA FROM NIGERIA 55
numerous, fine, closely-spaced ridges perpendicular to the hinge-line ; when the
valves are in contact the two areas are in one plane, occasionally pouting slightly
along the line of junction. Hinge-line straight, forming about seven-ninths of the
length of the shell. Dorso-lateral angles obtuse, the anterior a little the more so.
Anterior end gently convex near the dorso-lateral angle, rather sharply rounded
below. Ventral margin almost straight, gently upturned near the ends. Posterior
end fairly straight, joining the ventral margin in a sharply rounded curve. No byssal
depression. Hinge-plate very narrow, straight, its base horizontal, slightly expanded at
the extreme ends ; a small, narrow gap, slightly depressed, beneath the beak, with 22-
23 taxodont teeth on each side, the teeth being short and vertical except for the
five or six flanking ones which are slightly longer and oblique (converging ventrally) .
Surface not carinate, with fine, very numerous radial riblets of three orders minutely
beaded by fine concentric threads. Muscle scars large, subequal, not buttressed.
Valve margins smooth.
REMARKS. The more elongate and subrectangular form, characters of the hinge,
lack of a buttress to the posterior muscle scar, and smooth valve margins distinguish
this genus from Striarca. Breviarca is much shorter and has a much more convex
ventral margin.
Rectangularca africana (Newton)
(PL 8, figs. 5«, b, 6)
1922. Striarca africana Newton, p. 69, pi. 8, figs. 11—13.
MATERIAL. Several specimens, including the lectotype (1^.48172, Newton's fig. n)
here selected.
DIMENSIONS. Height 9-6 mm., length 18-3 mm., thickness (two valves) 7-8 mm.
Family PLICATULIDAE
Genus PLICATULA Lamarck, 1801
Plicatula polymorpha Bellardi
1922. Plicatula polymorpha Bellardi : Newton, p 65, pi. 6, figs. 6-9 (cum syn.).
MATERIAL. Fourteen specimens.
Family OSTREIDAE
Genus OSTREA Linne, 1758
Ostrea amekiensis sp. nov.
(PL 8, fig. 7)
1922. Ostrea cf. ludensis Deshayes : Newton, p. 61, pi. 8, figs. 2-3.
MATERIAL. The holotype ^.48195, Newton's figs. 2-3).
DESCRIPTION. The single specimen is a right valve, rounded-subtriangular, a little
56 EOCENE MOLLUSCA FROM NIGERIA
narrower in the upper third near the ligament area which (although worn) is much
enrolled forward but otherwise typical. The valve is almost flat, very slightly concave
medially, vaguely out-turned laterally and at the rounded end. Outer surface, apart
from growth lines, with numerous, fine vermicular riblets like those of Placuna
(about six to the mm. near the rounded end). Muscle scar slightly posterior to the
middle. Valve margin smooth internally.
DIMENSIONS. Height 28-7 mm., length 26-3 mm.
REMARKS. Although known only by a right valve, this species is well character-
ized by its Placuna-like. ornament. As shown by Deshayes's illustration, the right
valve (called " left valve ") of 0. ludensis is larger and less transverse, and has a
larger ligament area and ornament of a different type (not Placuna-like).
Ostrea pseudomarginidentata sp. nov.
(PI. 8, fig. 8)
1922. Ostrea cf. marginidentata S. V. Wood : Newton, p. 60, pi. 6, figs. 2-5.
MATERIAL. Several specimens, including the holotype (1,48203).
DESCRIPTION. Of medium size, moderately thick-shelled, linguiform. Left valve
(incomplete) with a large, flattened attachment area occupying most of the surface,
leaving a narrow area, inturned ventrally almost at right angles, with numerous
rather small radial ribs of about the same width as their intervals (four occupying
5-2 mm. ventrally, where they are widest). Valve margin correspondingly frilled.
Right valve irregularly flat. Outer surface smooth except for growth lines and a
few irregularities. A narrow, inturned margin, flanking the ligament area, has rather
prominent, short ribs which are irregularly spaced but normally narrower than their
intervals ; further from the umbo these merge into relatively coarse, much wider-
spaced crenulations on the inner margin of the valve. Muscle scar fairly large,
posterior to the middle.
DIMENSIONS. Holotype (a left valve) : height (incomplete) 24-3 mm., length
19-6 mm. Topotype (Newton's illustrated right valve, 1.48202) : height, 36-5 mm.,
length 22-8 mm.
REMARKS. While showing some superficial similarity to 0. marginidentata S. V.
Wood, this species differs not only in being smaller, but in being usually higher, and
in having a larger attachment area, and plications which are finer on both valves.
Subgenus CRASSOSTREA Sacco, 1897
Ostrea (Crassostrea) lugardi (Newton)
1922. Crassostrea lugardi Newton, p. 62, pi. 6, fig. i ; pi. 7, fig. i ; pi. 8, fig. i.
MATERIAL. Several specimens, including the lectotype (£,.48217, Newton's pi. 6,
fig. i and pi. 7, fig. i) here selected.
EOCENE MOLLUSCA FROM NIGERIA 57
Family CARDITIDAE
Genus GLANS M. von Miihlfeldt, 1811
Glans nigeriensis sp. nov.
(PL 8, figs, ga, b)
1922. Cardita cf. planicosta J. Sowerby : Newton, p. 80.
MATERIAL. The holotype (1^.48318).
DESCRIPTION. The solitary right valve is small, subtrapezoidal in outline, and
rather strongly inflated. Umbo moderately prominent, distinctly prosogyrous,
situated at about one-third of the length from the anterior end. Lunule small, indis-
tinct, limited in its early stages by a fine, raised thread. Escutcheon well defined.
Antero-dorsal margin gently convex, more steeply descendent than the longer,
straight postero-dorsal margin. Anterior end rather sharply rounded, situated
rather low down. Ventral margin almost straight. Posterior end truncated slightly
obliquely, joining the ventral margin in a rounded angle which is only a little more
than a right angle, and the postero-dorsal margin in a rounded angle of about 155°.
Surface ornamented with 23 rather low, moderately broad, rather vaguely beaded
ribs which are slightly wider than their intervals over the main body of the shell but
more closely spaced at the extreme ends ; they have rather square-cut, shallow
intervals, and are flattened on top, although slightly depressed along the middle line.
Valve margins coarsely fluted. Right valve hinge : AI small, pointed, with a small,
shallow socket above it ; 3a broken, evidently thin and moderately oblique ; 3b
solid, rather narrowly triangular, its dorsal pointed portion projecting into the shell
cavity ; a small PHI may have been present, but cannot be observed as the margin
of the shell is rather worn posteriorly.
DIMENSIONS. Height 11-9 mm., length 14-7 mm.
REMARKS. From its general form and ornamentation, as well as from the charac-
ters of the hinge, it is evident that this specimen is not a juvenile of the large
Venericardia (Venericor) planicosta (Lamarck).
Subgenus BENDEGLANS nov.
TYPE SPECIES. Cardita costaeirregularis Newton.
SUBGENERIC CHARACTERS. Of small-medium size, only moderately thick-shelled,
subtriangular-cuneiform, moderately strongly inflated ; anterior and posterior parts
of surface with flat ribs separated by linear intervals, median portion with only
three very wide, rather high and flat-topped ribs separated by broad intervals, the
intervals showing as raised broad ridges on the inner surface of the shell ; right valve
with small AI, low 3a, solid 3b, lamellar 5b, and small PHI ; left valve with All,
fairly solid 2 and 4b, PII, and small PIV.
GEOL. Ill, 2. 6
58 EOCENE MOLLUSCA FROM NIGERIA
Glans (Bendeglans) costaeirregularis (Newton)
(PI. 9, figs, i, 2)
1922. Cardita costaeirregularis Newton, p. 81, pi. 9, figs. 30-31.
1944. Cyclocardia costaeirregularis (Newton) : Chavan, p. 35.
MATERIAL. Many specimens, including the lectotype (1.48324, Newton's fig. 30)
here selected.
SUPPLEMENTARY SPECIFIC DESCRIPTION. Beaks pointed, rather high, prosogyrous,
situated anterior to the middle line at about two-thirds of the length. Lunule small,
cordiform, concave. Escutcheon rather short, narrow. Anterior part of surface with
nine depressed, flatly rounded, unbeaded ribs separated by linear intervals ; posterior
part with n similar but rather narrower ribs ; median part with three very broad,
elevated, flat-topped, unbeaded ribs separated by broad, subrectangular intervals ;
the anterior ribs increase in size posteriorly, the ninth being almost as large as the
adjacent one on the median part and separated from it by a linear interval ; the
most anterior of the posterior group is a little larger than the remainder ; valve
margins fluted according to the ribbing, the three main depressions of the outer
surface showing as ridges on the inner surface. Posterior adductor impression of
normal shape, anterior one rather elongate. Right valve : AI small, low, close to
the end of 33 ; 3a obscure, thin, low, more or less fused to the valve margin, only
slightly inclined forwards ; 3b solid, elevated, not very broadly triangular, inclined
slightly backwards ; 5b long, thin and lamellar, almost straight, fused to the lower
edge of the nymph from which it is separated by a narrow groove ; PHI lamellar,
moderately long, fused to the continuation of the ridge limiting the escutcheon.
Left valve : All short, fused to the ridge limiting the inner margin of the lunule ;
2 solid, elevated, narrowly triangular, slightly inclined forwards ; 4b solid, elevated,
rather more narrowly triangular, oblique ; PI I small ; apparently a still smaller
tooth (PIV) above and behind PI I. Nymph narrow, relatively short. Escutcheon of
right valve with a groove outside it, radiating from beneath the beak and enlarging
with growth.
REMARKS. Chavan placed this species in Cyclocardia, but in C. borealis (Conrad),
as represented by specimens in the British Museum (Natural History), the shell is
suborbicular and has beaded ribs in youth, and vague, flatly V-shaped ribs in the
adult, with no median enlarged ribs ; there are no posterior lateral teeth, and the
cardinal teeth of the right valve differ in orientation. The hinge of Cardita (Cyclo-
cardia) granulata Say, as figured by Chavan, differs in a similar manner. The greatly
enlarged median ribs and the hinge characters warrant the placing of the Nigerian
species in a new subgenus.
Subgenus AMEKIGLANS nov.
TYPE SPECIES. Cardita costaenodulosis Newton.
SUBGENERIC CHARACTERS. Of small-medium to medium size, very thick-shelled,
oval-subtriangular to cuneiform, inflation moderate ; extreme posterior ribs flat
EOCENE MOLLUSCA FROM NIGERIA 59
and rather low, slightly wider than their intervals, the remainder very narrow and
high, with rounded tops, finely beaded, with deep, broader, smooth, U-shaped
intervals ; right valve with AI, low 3a, solid 3b, lamellar 5b, and distinct PHI ;
left valve with short, pointed All, solid triangular 2, narrower and longer 4b, and
distinct PI I.
Glans (AmekiglansJ costaenodulosis (Newton)
(PI. 9, figs. 3, 4)
1922. Cardita costaenodulosis Newton, p. 82, pi. 9, figs. 22-25.
1938. Cossmannella costaenodulosis (Newton) : Chavan, pp. 3-10, fig. i.
1944. Cossmannella costaenodulosis (Newton) : Chavan, p. 35.
MATERIAL. Many specimens, including the lectotype (1.48360, Newton's figs.
24-25) here selected.
SUPPLEMENTARY SPECIFIC DESCRIPTION. Beaks pointed, rather high, prosogyrous,
situated anterior to the middle line at about two-thirds of the length. Lunule small,
narrowly cordiform, depressed, bulging medially. Escutcheon narrow, rather short.
Extreme posterior end with five rather flattened, unbeaded ribs with slightly nar-
rower intervals, the remainder of the surface with 14 very narrow and high, finely
beaded ribs with rounded tops, separated by smooth, broadly U-shaped intervals
of nearly three times their width. Valve margins coarsely fluted. Right valve : AI
rather small, fairly close to and in line with the end of 3a ; 3a low, not very long,
sloping forwards at about 45°, partially fused to the inner margin of the lunule, from
which it is separated by a slight groove ; 3b solid, elevated, triangular, its anterior
edge vertical ; 5b rather long, very thin and lamellar, fused to the lower edge of
the nymph ; PHI distinct, moderately long, rather distant, situated fairly near the
valve margin. Left valve : AH small, pointed, situated at the forward end of the
inner edge of the lunule ; 2 solid, elevated, triangular, its posterior edge vertical ;
4b solid, more narrowly triangular, oblique at about 45° ; PII distinct. Posterior
adductor impression tending to be obliquely subrectangular ; anterior adductor
impression moderately elongate.
REMARKS. Chavan placed this species in Cossmannella. None of the specimens of
Cardita fayumensis Oppenheim (C. aegyptiaca Fraas non Monterosato) , the type
species of Cossmannella, in the British Museum (Natural History), shows the hinge.
It is not recorded as having any lateral teeth, but Chavan has intimated to me that
the hinges of the two species are identical as judged from specimens in his own
collections. The ribs of C. fayumensis are recorded as being " few, strong, sharp,
slightly tripartite ", but while specimens in the British Museum (Natural History)
do occasionally show tripartite ribs, this is due to wear. The ribs are considerably
more numerous and less high than in costaenodulosis, the intervals being much less
conspicuous, and the shell is not normally so pointed posteriorly, often being trun-
cated. In view of the doubts concerning the exact characters of the hinge of
Cossmannella, and since its form and ribbing apparently differ appreciably from those
of the Nigerian species, a new subgeneric name seems warranted for the latter.
Go EOCENE MOLLUSCA FROM NIGERIA
Subgenus DIVERGIDENS nov.
TYPE SPECIES. Cardita triparticostata Newton.
SUBGENERIC CHARACTERS. Of small to small-medium size, fairly thick-shelled,
rather strongly inflated, outline subtriangular, moderately high. A few simple, narrow
ribs posteriorly, the remainder of the surface with broad, low, square ribs with deep,
narrow, almost linear, square-cut intervals, the middle third of each rib bearing a
strong, rounded, beaded cord. Right valve with AI, obsolete and very oblique 3a,
solid and rather broadly triangular 3b, and distinct PHI ; left valve with All,
strongly divergent 2 and 4b (the latter the more oblique), PII and weak PIV.
Glans (Divergidens) triparticostata (Newton)
(PI. 9, figs. 50, b, 6, 7)
1922. Cardita triparticostata Newton, p. 83, pi. 9, figs. 26-29.
1938. " Venericardia " triparticostata (Newton) : Chavan, p. 7.
MATERIAL. Several specimens, including the lectotype (1^.48319, Newton's fig. 26)
here selected.
SUPPLEMENTARY SPECIFIC DESCRIPTION. Beaks pointed, prominent, moderately
prosogyrous, situated anterior to the middle line at about three-fifths of the length.
Lunule rather small, moderately broadly cordiform, smooth, not much sunk.
Escutcheon short. Extreme posterior end with seven low, gently rounded, unbeaded
ribs with rather narrower intervals, the remainder of the surface with 18 low, flat,
broad ribs with deep, narrow, almost linear, square-cut intervals, the middle third of
each rib carrying a strong, rounded, beaded cord. Valve margins coarsely fluted.
Right valve : AI small but distinct, slightly elongate ; 3a obscure, long, low, thin,
lamellar, very oblique and only gently descendent, fused to the inner margin of the
lunule ; 3b solid, elevated, rather broadly triangular, its anterior edge sloping gently
forwards, its posterior edge sloping more obliquely backwards ; PHI distinct. Left
valve : All distinct, slightly elongated ; 2 and 4b strongly divergent, solid, elevated,
narrowly triangular, their inner margins forming an angle of about 80°, 4b slightly
more oblique than 2 ; PII distinctly elongated ; PIV similar but weaker. Nymph
rather narrow, moderately long. Posterior adductor impression normal in shape ;
anterior adductor impression moderately elongate.
REMARKS. Chavan tentatively referred this species to Glyptoaxis, but the right
cardinal is not curved and elongate ; he has recently suggested to me that it may be
a Cardiocardita (group of Cardita beaumonti), but Cardiocardita has no anterior
laterals. In its form, ornament, and hinge characters the Nigerian species appears
to belong to a new subgenus.
Family LUCINIDAE
Genus PHACOIDES Gray, 1847
Phacoides eaglesomei Newton
1922. Phacoides eaglesomei Newton, p. 75, pi. 8, figs. 18-20.
MATERIAL. Numerous specimens, including the lectotype (£,.48237, Newton's
fig. 18) here selected.
EOCENE MOLLUSCA FROM NIGERIA 61
Genus GIBBOLUCINA Cossmann, 1904
Subgenus EOMILTHA Cossmann, 1910
Gibbolucina (Eorniltha) (?) subrhomboidalis (Newton)
1922. Phacoides subrhomboidalis Newton, p. 77, pi. 7, figs. 4-5.
MATERIAL. The holotype (1,48244).
REMARKS. The characters of the hinge are unknown since the valves of the single
specimen available cannot be separated. Chavan (in correspondence) has suggested
that the species may be an Eomiltha.
Genus POMPHOLIGINA Dall, 1901
Subgenus EODIVARICELLA Chavan, 1951
Pompholigina (Eodivaricella) oppenheimi (Newton)
1922. Divaricella oppenheimi Newton, p. 78, pi. 7, figs. 2-3.
1951. Pompholigina (Eodivaricella) oppenheimi (Newton) : Chavan, p. 23, fig. 27.
MATERIAL. A few specimens, including the lectotype (L.48O97, Newton's fig. 2)
here selected.
REMARKS. This is the type species of Eodivaricella.
Family CARDIIDAE
Genus FRAGUM (Bolten MS.) Roding, 1798
Subgenus AFRICOFRAGUM nov.
TYPE SPECIES. Cardium cf. obliquum Lamarck : Newton = Fragum (Africofragum)
newtoni sp. nov.
SUBGENERIC CHARACTERS. Small, outline Fragum-like, length and height nearly
equal, posteriorly subcarinate. Ornament of flattened ribs which have rather
narrower intervals, and are smooth except for the most anterior four or five which
bear small prickles ; ribs on posterior area more irregular and producing short spines
at the margin. Postero-dorsal margin, of left valve only, with a row of six short,
upturned and outward-bent spines. Hinge less arched than in Fragum, anterior
lateral teeth a little closer to cardinals than posterior lateral teeth. Nymph short.
Fragum (Africofragum} newtoni sp. nov.
(PI. 9, figs. Sa-c)
1922. Cardium cf. obliquum Lamarck : Newton, p. 74, pi. 7, figs. 6-9.
MATERIAL. Many specimens, including the holotype (L.484i6, Newton's fig. 6).
DESCRIPTION. Beaks moderately prominent, not large, prosogyrous, situated
slightly anterior to the median line. Antero-dorsal margin short, straight or slightly
62 EOCENE MOLLUSCA FROM NIGERIA
convex, joining the anterior end in an obtusely rounded angle. Anterior end well
rounded, receding a little ventrally. Ventral margin convex, its posterior part the
straighter, joining the posterior end in a blunt angle of a little more than 90°.
Posterior end obliquely truncated, gently convex. Postero-dorsal margin short,
nearly straight. About 34 ribs. Left valve : All obscure, below the forward end
of a well-developed AIV ; 2a massive, upturned, pointed ; 2b small ; PII and PIV
small. Right valve : AI larger than AIII ; 3a small ; 3b massive, upturned, pointed ;
PI well developed ; no PHI below the straight shell margin, above which are the
spines. Valve margins fluted.
REMARKS. Cardium obliquum Lamarck appears to be a Loxocardium, and is quite
different from the Nigerian form in that it is transversely oval in outline, not sub-
carinate, and has distinct transverse scales or beads on the ribs, but no spines.
Family VENERIDAE
Genus TIVELINA Cossmann, 1886
Tivelina newtoni sp. nov.
(PL 9, figs, ga, b, loa, b)
1922. Tivelina cf. sphenarium (Bayan) : Newton, p. 90, pi. 9, figs. 7-10.
MATERIAL. Many specimens, including the holotype ^.48504, Newton's fig. 7).
DESCRIPTION. Small to small-medium. Moderately thick-shelled, oval-subtri-
angular. Beaks small, prosogyrous, high, situated anterior to the middle line at
about two-thirds of the length. Inflation moderate ; greatest height anterior to the
middle line, at the position of the beaks. Lunule large, narrowly cordiform, limited
by a fine raised thread. Escutcheon narrow, rather short. Antero-dorsal margin
gently convex, steeply descendent. Anterior end well rounded, a little produced
antero-ventrally. Ventral margin convex, straighter (even vaguely emarginate)
posteriorly. Posterior end moderately produced, situated rather low down, sharply
rounded. Postero-dorsal margin long, slightly convex, rather steeply descendent.
Surface with numerous strong, rather irregular, fairly sharp, concentric threads.
Right valve : AI and AIII small, short, lamellar ; 3a rather short, simple, thin,
inclined forwards ; i vertical, narrowly triangular, simple ; 3b oblique, deeply
grooved. Left valve : All prominent ; 2a thin, simple, inclined forwards ; 2b
narrowly triangular, simple, inclined backwards ; 4b oblique, thin, lamellar, simple,
moderately long. Nymph moderately long, thin. Pallial sinus subtriangular, rather
short, its apex not reaching the middle line, its upper arm subhorizontal, its lower
arm steeply descendent. Valve margins smooth.
REMARKS. Comparison with specimens in the British Museum (Natural History)
shows that this is not Bayan's species ; the ornament is more serrate and less
Costacallista-like, the pallial sinus is larger, and the hinge-plate has no rectangularly
ending projection beneath the anterior lateral teeth, as in T. sphenarium.
EOCENE MOLLUSCA FROM NIGERIA 63
Genus PITAR Roemer, 1857
Pitar amekiensis sp. nov.
(PL 9, figs, na-c, 12 ; PL 10, fig. i)
1922. Cordiopsis incmssata (J. Sowerby) : Newton, p. 85 (pars).
MATERIAL. The holotype (1.48253) and two topotypes (L. 48245-6).
DESCRIPTION. Of medium size, moderately thick-shelled, rather strongly inflated,
equivalve, subtriangular in outline. Beaks rather prominent, prosogyrous, situated
anterior to the middle line at about one-fifth of the length. Escutcheon moderately
long, narrow. Lunule large, rather narrowly cordiform, limited by a vague incised
line. Antero-dorsal margin straight, steeply descendent. Anterior end rather sharply
rounded. Ventral margin convex, with a gentle median bulge, straighter anteriorly
and posteriorly. Posterior end vaguely truncated, joining the ventral margin in an
obtusely rounded angle. Postero-dorsal margin moderately long, gently convex.
Surface with irregular concentric threads, often with noticeably narrower intervals.
Right valve : AI and a smaller, shorter AIII ; 3a vertical, rather thin, its forward
face less perpendicular to the surface of the hinge-plate than its posterior face ; I
sloping slightly backwards, its posterior face the less nearly perpendicular to the
hinge-plate ; 3b oblique, moderately long, deeply grooved. Left valve : All pro-
minent, pointed ; 2a thin, lamellar, vertical ; 2b oblique at about 45°, narrowly
triangular, higher posteriorly ; 4b moderately long, simple, gently curved down-
wards near its end. Nymph of medium length (about twice as along as the posterior
cardinal tooth). Muscle impressions not very large. Pallial sinus rather acutely
triangular, its upper arm gently ascendent, its apex narrowly rounded and situated
slightly less than half-way across to the anterior adductor impression, its lower arm
very steeply descendent. Valve margins smooth.
DIMENSIONS. Holotype: height 27-0 mm., length 30-7 mm. Topotype (1,48245) :
height 30-9 mm., length 35-0 mm.
REMARKS. These three specimens, which were labelled " Cordiopsis incmssata "
by Newton, together with those here recorded as Sinodiopsis coxi sp. nov., are quite
different from that species ; they are more inflated anti more triangular, the beaks are
much higher and less anterior in position, the hinge-plate is placed less forward and
less inclined, the pallial sinus is a little shorter and more ascendent, the nymph is
shorter, the teeth differ in the details of their orientation, the ventral margin is more
bulging, and the concentric ornament is less serrate.
Genus CHIONELLA Cossmann, 1886
Subgenus COSTACALLISTA Palmer, 1927
Chionella (Costacallista) elongatotrigona (Newton)
1922. Callista elongatotrigona Newton, p. 88, pi. 9, figs. 1-5.
MATERIAL. Many specimens, including the lectotype (1,48065, Newton's fig. 3)
here selected.
64 EOCENE MOLLUSCA FROM NIGERIA
Subgenus MICROCALLISTA Stewart, 1930
Chionella (Microcallista) kitsoni (Newton)
(PI. 10, figs. 2a, b, 3)
1922. Callista kitsoni Newton, p. 89, pi. 9, figs. 6, 11-14.
MATERIAL. Several specimens, including the lectotype (L.48o69, Newton's fig. n)
here selected.
REMARKS. The left anterior cardinal tooth (2a) is simple, not grooved as in
" Callista " auct. ( = Costacallista Palmer, 1927).
Genus SINODIA Jukes-Browne, 1908
Sinodia heward-belli Newton
1922. Sinodia heward-belli Newton, p. 86, pi. 9, figs. 18-21.
MATERIAL. Several specimens, including the lectotype (L.48iO2, Newton's fig. 21)
here selected.
REMARKS. The left anterior cardinal tooth (2a) is not faintly grooved as in
Sinodia, but in all other characters the species agrees with that genus and seems best
retained in it.
Genus SINODIOPSIS nov.
TYPE SPECIES. Cordiopsis incrassata (J. Sowerby) : Newton = Sinodiopsis coxi
sp. nov.
GENERIC CHARACTERS. Of medium size, moderately well inflated, transversely oval-
subquadrate in outline, equivalve. Beaks rather small, prosogyrous, not prominent,
well recurved, situated anterior to the middle line at about one-quarter to one-fifth
of the length. Escutcheon fairly long, narrow. Lunule vague, large and moderately
narrowly cordiform, limited by a fine incised line. Surface ornamented with rather
irregular, strong, closely-spaced concentric threads. Hinge-plate moderately well
developed ; not as thick, as inclined, or as high as in Cordiopsis and Sinodia. Right
valve : AI rather solid and triangular ; AIII small ; 3a thin, lamellar, gently inclined
forwards ; i rather triangular, its forward edge vertical, high, lamellar, its surface
shelving downwards posteriorly ; 3b oblique and deeply bifid. Left valve : All
very prominent and pointed ; 2a thin, lamellar, vertical ; 2b not so thin as 2a but
lamellar at the apex, oblique at about 45° ; 4b simple, thin, long, gently arched.
Nymph long, narrow, gently arched. Muscle impressions rather large, situated rela-
tively more dorsally than in Cordiopsis and Sinodia. Pallial sinus bluntly and
moderately acutely triangular, its upper arm horizontal or slightly ascendent, its
apex sharply rounded, its lower arm descendent at about 45°, the apex situated at
about mid-length of the shell. Valve margins smooth.
EOCENE MOLLUSCA FROM NIGERIA 65
Sinodiopsis coxi sp. nov.
(PI. 10, figs. 4, 5)
1922. Cordiopsis incrassata (J. Sowerby) : Newton, p. 85 (pars), pi. 9, figs. 15-17 (non J.
Sowerby).
1938. Macrocallista palmerae Caster, p. 66 (pars) (non pi. i, figs. 9-10 ; pi. 8, fig. 7).
MATERIAL. The holotype (1^.48250) and several topotypes.
REMARKS. This form is quite distinct from Sinodia (Cordiopsis} orbicularis
(Goldfuss) (= Venus incrassata J. Sowerby non Brocchi) in its outline, hinge, and
pallial sinus ; the species was erroneously referred by Caster to the Angola Miocene
species Macrocallista palmerae, but it differs still more from Macrocallista. While
the hinge and pallial sinus are of the same type as in Sinodia, the form is quite
different, the hinge-plate is less massive and less inclined, and tooth 2a is not grooved.
Cordiopsis, which also has a much more massive and more inclined hinge and a
different outline, has a shorter pallial sinus. The species is named after Dr. L. R. Cox.
Family MACTRIDAE
Genus SPISULA Gray, 1837
Subgenus CREPISPISULA nov.
TYPE SPECIES. Mactra semisulcata Lamarck: Newton = Spisula (Crepispisula)
amekiensis sp. nov.
SUBGENERIC CHARACTERS. Of medium size, moderately thin-shelled, subtriangular,
rather Hecuba-like in outline, rather strongly inflated, subcarinate anteriorly and
posteriorly. No lunule or escutcheon. Ornament of coarse incrementals. Left
valve : All rather short, prominent ; 2a and 2b meeting in a right angle dorsally,
projecting ; resilium pit moderately narrowly triangular, its anterior side limited by
a high, thin lamella, its posterior side by a very slight ridge ; PII prominent, a
little longer than All ; posterior and anterior lateral teeth at about equal distances
from the cardinal teeth. Right valve : AI a little larger and stronger than All I ;
3a and 3b simple, divergent, 3a a little the more oblique ; resilium pit as in the left
valve ; PI and PHI better developed than the opposing anterior lateral teeth. A
narrow ligament slit extends from the dorsal side of the hinge-plate to the tip of the
beak. Pallial sinus narrow, forming a gently ascending tongue with its apex at mid-
length of the shell. Valve margins smooth.
Spisula (Crepispisula} amekiensis sp. nov.
(PI. 10, figs. 6a, b, 7)
1922. Mactra semisulcata Lamarck : Newton, p. 93, pi. 7, figs. 10-13.
MATERIAL. Several specimens, including the holotype ^.48224, Newton's figs.
10-11).
66 EOCENE MOLLUSCA FROM NIGERIA
SUPPLEMENTARY SPECIFIC DESCRIPTION. Beaks rather high, narrow, prosogyrous.
Antero-dorsal margin long, straight. Anterior end narrowly rounded, even bluntly
pointed. Ventral margin straightest posteriorly, bulging somewhat downwards
along its anterior half. Posterior end sharply and narrowly rounded to bluntly
pointed. Postero-dorsal margin arched, obtusely angulated in two places. Anterior
carination formed by a vague depression anterior to which the concentric ornament
becomes less closely spaced than on the middle of the shell. Posterior angulation
formed of two very obtuse carinae, the ornament behind the posterior one being
likewise relatively less closely spaced.
REMARKS. Although this form is somewhat similar in outline to the Recent genus
Scissodesma Gray, it has definite concentric ornament, its posterior carina is less
marked, the ligament slit from the dorsal side of the hinge-plate to the tip of the
beak is distinctly smaller, and the lateral teeth are not crenulated. Mactra semi-
sulcata Lamarck, a Paris Basin Eocene species with which Newton identified the
Nigerian form, possesses a similar ligament slit, but its outline differs in being less
triangular and less bulging antero-ventrally, it has a less definite anterior ridge,
and its ornament is weaker.
Family TELLINIDAE
Genus MACOMA Leach, 1819
Subgenus BENDEMACOMA nov.
TYPE SPECIES. Peronaea nigeriensis Newton.
SUBGENERIC CHARACTERS. Of large-medium size, rather thick-shelled, transversely
oval-subtriangular, length considerably exceeding height, inflation moderate. Beaks
small, moderately prominent, prosogyrous. Surface ornamented with accentuated
growth-lines, posteriorly with two very vague carinae. Escutcheon long, narrow.
Lunule narrow, shorter than escutcheon, limited by a fine incised line. Left valve :
2a vertical, distinctly grooved dorsally ; 2b moderately oblique, very thin and
lamellar, simple ; no lateral teeth. Right valve : 3a rather solid, directed moderately
forwards, grooved dorsally ; 3b a little longer than 3a, directed backwards moder-
ately obliquely, well grooved ; no lateral teeth. Nymph long and rather narrow.
Pallial sinus rather narrowly tongue-shaped, its upper part rising slightly for a short
distance from the posterior adductor impression, then gently descendent for most of
its length, narrowly rounded at its apex, its lower part coalescent with the pallial
line posteriorly for nearly half its length. Valve margins smooth.
Macoma (Bendemacoma) nigeriensis (Newton)
1922. Peronaea nigeriensis Newton, p. 91, pi. n, figs. 1-3.
MATERIAL. Several specimens, including the lectotype ^.48513 , Newton's fig. i)
here selected.
REMARKS. The characters of the hinge and the form of the pallial sinus indicate
that this species is not a Peronaea ; it appears to belong to a new subgenus of
EOCENE MOLLUSCA FROM NIGERIA 67
Macoma characterized by the shape of the shell, the grooved 3a, and the form of the
pallial sinus.
Family MYIDAE
Genus RAETOMYA Newton, 1919
Raetomya schweinfurthi (Mayer-Eymar)
1922. Raetomya schweinfurthi (Mayer-Eymar) : Newton, p. 96, pi. 10 (cum syn.).
1942. Raetomya schweinfurthi (Mayer-Eymar) : Rossi, p. 182, pi. n, fig. 3.
1952. Labiosa (Raeta) schweinfurthi (Mayer-Eymar) : Tessier, pp. 350-351, pi. 29, figs. 7-8.
1954. Labiosa (Raeta) schweinfurthi (Mayer-Eymar) : Dartevelle & Roger, pi. 5 fig. 4.
1955. Raeta schweinfurthi (Mayer-Eymar) : Dartevelle & Roger, pp. 164-167.
REMARKS. This is the type species of Raetomya. Tessier placed it as a subgenus
of the mactrid genus Labiosa Schumacher, 1817, which is evidently a synonym of
Anatina Lamarck, 1816. Raeta is also a mactrid genus, but schweinfurthi has been
well described by Newton and is undoubtedly a myid. Since it was recorded from
Egypt and Nigeria, it has also been recorded from the Eocene of Tripolitania and the
Cameroons and from the (reputed) Lower Lutetian or Ypresian of Senegal ; the
horizon stated in the latter record is rather low, and it might be advisable to re-
investigate the evidence on which the age was assigned. The Angola Miocene speci-
mens referred to this species by Caster actually belong to Platyodon klinghardti
( J. Bohm) ; Dartevelle & Roger express doubts concerning the generic assignation of
the latter species, and it may well be that it is a distinct Miocene species of Raetomya.
Family CORBULIDAE
Genus VARICORBULA Grant & Gale, 1931
Varicorbula amekiensis sp. nov.
(PI. 8, figs. 10, ii ; PI. 10, figs. 8a-c)
1922. Corbula rugosa Lamarck : Newton, p. 98, pi. 7, figs. 14-18.
MATERIAL. Many specimens, including the holotype ^.48264, Newton's fig. 14).
DESCRIPTION. Small, thick-shelled, inequivalve, right valve larger and more
strongly inflated than the left, beaks submedian (on the right valve a little anterior
to the middle line on account of the greater rostration), prosogyrous, right valve
umbo strongly enrolled. Outline subtriangular.
Right valve very strongly inflated, umbo prominent but not broad, posterior end
with a moderately short, slightly upturned rostrum, with two blunt carinae pos-
teriorly, limiting the upper and lower portions of the rostrum. Anterior end sharply
rounded. Ventral margin gently convex, straighter posteriorly. Ornament con-
sisting of strong, rounded concentric folds of about the same width as or slightly
wider than their intervals, regularly increasing in size with growth. Valve margin
internally smooth, with a fine incised line some distance from the edge for the recep-
68 EOCENE MOLLUSCA FROM NIGERIA
tion of the margin of the smaller left valve. Tooth i solid, triangular, strongly
upturned and pointed, with a deep, triangular chondrophore behind it ; PI short,
lamellar. Pallial sinus widely rounded, reaching about one-third of the distance
towards the anterior adductor impression. Inner half of rostrum with two short
grooves extending downwards and outwards, one near the top, the other half-way
down ; these are not associated with a left valve siphonal plate (such as is present
in Caestocorbuld), no trace of which has been found on any of the specimens.
Left valve fitting within the margin of the right, inner portion of right valve
rostrum remaining visible. Valve oval-subtriangular, inflated (a little flattened
posteriorly near the blunt carina), beak narrow, umbo with fine concentric folds,
main portion of surface with growth-lines only, apart from four very fine, widely-
spaced radial threads on its middle third. Tooth 2b oblique, simple, with a deep,
triangular chondrophore in front of it ; there is a trace of a very small, pointed All
at the anterior corner of the chondrophore. Pallial sinus as in the right valve. Valve
margin smooth.
REMARKS. The illustrations of Cossmann & Pissarro, and specimens from France
in the British Museum (Natural History), suggest that the name Corbula rugosa
Lamarck has been applied to two forms, one relatively finely ornamented, the other
(probably a Varicorbula) more strongly ornamented, higher, and more inflated. The
Nigerian form is not conspecific with either of these ; the rostrum and umbonal
region are narrower, the shell is more triangular, and there is no distinct initial stage
separated off by a constriction.
Family KITSONIIDAE nov.
TYPE GENUS. Kitsonia gen. nov.
FAMILY CHARACTERS. Lithophaga-like in form. No gape. Not nacreous externally.
Sinupalliate. Internally, that part of the surface behind the pallial sinus is highly
polished and nacreous. Ligament external. No hinge-plate, the two right valve
teeth and the one left valve tooth, all cardinals, project markedly beyond the plane
of commissure. Apparently a boring form.
REMARKS. This form belongs to a new family which is provisionally regarded
as being related to the Clavagellidae and is placed in the Clavagellacea.
Genus KITSONIA nov.
TYPE SPECIES. Coralliophaga eocenica Newton.
GENERIC CHARACTERS. Small, thin-shelled, equivalve, very elongate, ventral
margin slightly excavated, posterior end slightly curved downwards. Beaks small,
low, prosogyrous, situated slightly behind the anterior end. Surface smooth except
for growth lines. Anterior adductor impression moderately broad and large ;
posterior adductor impression situated medially just below the dorsal margin.
Pallial sinus apparently extending for two-fifths of the length of the shell. That
part of the surface anterior to the pallial sinus dull, white, with obscure, raised,
rather vermicular, radial markings (reminiscent of the Lucinidae), that part posterior
EOCENE MOLLUSCA FROM NIGERIA 69
to the sinus highly polished and nacreous. No lunule or escutcheon. Ligament
narrow, rather long, external. Right valve : two lamellar subumbonal teeth, sub-
horizontal and slightly ascending posteriorly, the anterior one relatively short,
highest distally ; posterior tooth very obliquely bifid, its posterior portion very
close to the anterior portion and largely behind it. Left valve : one long, lamellar,
simple, subumbonal tooth, highest distally, subhorizontal, and slightly ascending
posteriorly. No other teeth. Valve margins smooth internally.
REMARKS. As noted by Newton, this form is quite different from the edentulous
and gaping Gastrochaena and from the mytilid genus Lithophaga, which is integri-
palliate and likewise edentulous. However, the lack of a hinge-plate, the entirely
different dentition, and the lack of radial ornament and of wide-spaced concentric
frills distinguish it readily from Coralliophaga.
Kitsonia eocenica (Newton)
(Plate 9, figs. 13, 14)
1922. Coralliophaga eocenica Newton, p. 99, pi. n, figs. 4-5.
MATERIAL. Two specimens (a right valve and a left valve) ; lectotype ^.48192,
Newton's fig. 5) here selected.
III. ACKNOWLEDGMENTS
The writer is indebted to the authorities of the British Museum (Natural History)
for facilities to undertake the work ; to the Chairmen and Directors of the joint
Shell and British Petroleum organization for permission to publish the results ;
to Dr. L. R. Cox for advice at all stages of the investigation ; to palaeontologists
of the Shell organization for stimulating discussions concerning the fauna and its
age ; and to Mr. A. Chavan, who has collaborated in the study of the Lucinidae.
IV. REFERENCES
CASTER, K. E. 1938. Macroscopic fauna of the Quimbriz (Eocene) formation on the Luculo
River, Angola. Com. Serv. geol. Portugal, Lisboa, 20 : 53-85, pis. i-io.
CHAVAN, A. 1938. Un groupe africain des Carditides ; Cossm'annella Mayer-Eymar. Bull. Soc.
geol. Fr., Paris (5) 8 : 3-10.
1944- Sur la portee d'un remarquable 61argissement des cotes chez les V6n6ricardes. C. R.
Soc. geol. Fr., Paris, 1944, 4 : 34-36.
1951. Essai critique de classification des Divaricella. Bull. Inst. Sci. nat. Belg., Bruxelles,
27, 18 : 1-27.
• 1952. A new fossil gastropod from the Middle Eocene of Ameki, Nigeria. Proc. Malac.
Soc. Lond., 29 (2-3) : 80-82, fig. i.
DARTEVELLE, E. 1952. Echinides fossiles du Congo beige et de 1'Angola, I. Introduction, his-
torique et stratigraphique. Ann. Mus. Congo beige (8°) Sci. geol., Tervuren, 12 : 1-14,
23 figs.
DARTEVELLE, E. & ROGER, J. 1954. Contribution a la connaissance de la faune du Miocene de
1'Angola. Com. Serv. geol. Portugal, Lisboa, 35 : 227-312, pis. 1-5.
• !955- Platyodon klinghardti (J. Bohm, 1919). LameUibranche du Miocene
d'Afrique occidentale. Bull. Mus. Nat. Hist. nat. Paris (2) 27, 2 : 164-169.
70 EOCENE MOLLUSCA FROM NIGERIA
MACNEIL, F. S. 1938. Species and Genera of Tertiary Noetinae. U.S. Geol. Surv. Prof. Paper,
189A : 1-49, pis. 1-6.
NEWTON, R. B. 1922. Eocene Mollusca from Nigeria. Geol. Surv. Nigeria Bull., 3 : 1-114, P^s-
i-n.
Rossi, C. 1942. Molluschi paleogenici della Sertica. Pubbl. 1st. Geol. Univ. Milano (P) 33 :
109-193, pis. i-n.
SCHILDER, F. A. 1929. The Eocene Amphiperatidae and Cypraeidae of England. Proc. Malac.
Soc. Land., 18 : 298-311.
1932. Cypraeacea. Fossilium Catalogus, i Animalia. Pars 55. 276 pp. Berlin.
TESSIER, F. 1952. Contribution a la stratigraphie et a la paleontologie de la partie ouest du Senegal
(Crdtace et Tertiare). These, Univ. Marseille, IIIe Partie : Paleontologie.
WENZ, W. 1938-44. Gastropoda. In Schindewolf, O. H. Handbuch der Paldozoologie, 6, i.
Berlin.
WHITE, E. I. 1955. Notes on African tertiary sharks. Colonial Geology & Mineral Resources,
London, 5 : 319-325, pis. i, 2.
PLATE 5
FIG. i. Turritella amekiensis sp. nov. Holotype (0.42215). x i.
FIG. 2. Turritella amekiensis sp. nov. Paratype (0.42222). x 3.
FIGS. $a-c. Architectonica (Nipteraxis) bendeica sp. nov. Holotype (0.42373). X 3.
FIG. 4. Architectonica (Stellaxis) bicingulata (Newton). Syntype (0.42356). X i.
FIGS. 5<z-c. Architectonica (Solariaxis] amekiensis sp. nov. Holotype (0.42361). X i.
FIG. 6. Acrilla nigeriensis sp. nov. Holotype (0.42285). x 2.
FIGS, ja, b. Calyptraea newtoni sp. nov. Holotype (0.42438). x 2.
FIG. 8. Turbocalyptraea scabrosa gen. et sp. nov. Holotype (0.42437). x 2.
FIGS. ga, b. Turbocalyptraea scabrosa gen. et sp. nov. Paratype (0.42436). x 2.
FIGS. 100, b. Semiterebellum (Africoterebellum) elongatum Newton. Syntype (0.41761). x i.
Bull. B.M. (N.H.) Geol. 3, 2.
PLATE 5
5b
' *
5a
5c
PLATE 6
FIGS. la-c. Xenophora nigeriensis (Newton). Lectotype (0.42291). x 2.
FIG. 2. Eovolva nigeriensis (Newton). Lectotype (0.41786). x 2.
FIG. 3. Eovolva nigeriensis (Newton). Syntype (0.41787). x 2.
FIGS. 40, b. Sphaerocypraea sudanensis (Schilder). Syntype (0.45778). x i.
FIGS. 5«, b. Neverita amekiensis sp. nov. Holotype (G. 42383). x i.
FIGS. 6a-c. Sinum africanum Newton. Lectotype (0.42406). x 2.
FIGS, ja, b. Sinum nigeriense sp. nov. Holotype (G. 42390). x 2.
FIG. 8. Leitcozonia pseudominax sp. nov. Holotype (0.42052). x i.
Bull. B.M. (N.H.) Geol. 3, 2.
PLATE 6
7a
8
PLATE 7
FIGS. la-c. Varicohilda turriculata (Newton). Lectotype (0.41971). x 2.
FIGS. 2a, b. Bendeia africana (Newton). Lectotype (0.42096). x 2.
FIG. 3. Strepsidura (Strepsiduropsis) spirata Newton. Lectotype (0.42008).
X 2.
FIGS. 4«-c. Sveltia (Africosveltia) multiplicis (Newton). Holotype (0.42137).
X i.
FIGS. 5«-c. Bonellitia (Admetula) amekiensis sp. nov. Holotype (0.42171). x 2.
FIGS. 6a-c. Bonellitia (Africostoma) decorata (Newton). Lectotype (0.42138). x 2.
FIGS. ja-c. Conns (Leptoconus) amekiensis sp. nov. Holotype (0.41837). x 2.
FIGS. 8a, b. Mitrelloturris casteri (Chavan). Holotype (0.69600). x 2.
Bull. B.M. (N.H.\ Geol. 3, 2.
PLATE 7
la
•jH^*
I
PLATE 8
FIG. i. Mitrelloturris casteri (Chavan). Holotype (0.69600). x 2.
FIGS. 20,-c. Amekicythara douvillei (Newton). Lectotype (0.42100). x 2.
FIGS. 3<z, b. Africarca nigeriensis (Newton). Syntype (L. 48443). x 2. Right valve.
FIG. 4. Arcopsis africana (Newton). Syntype (L. 48135). x 2. Right valve.
FIGS. 5a, b. Rectangularca africana (Newton). Lectotype (L. 48172). x 2. 5« : left valve.
FIG. 6. Rectangularca africana (Newton). Syntype (L. 48173). x 2. Right valve.
FIG. 7. Ostrea amekiensis sp. nov. Holotype (L. 48195). x 2. Right valve.
FIG. 8. Ostrea pseudomarginidentata sp. nov. Holotype (L. 48203). x 2. Antero-ventEa.1 end
of left valve.
FIGS, ga, b. Glans nigeriensis sp. nov. Holotype (L. 4831 8). x 2. Right valve.
FIG. 10. Varicorbula amekiensis sp. nov. Syntype (L. 48265). x 2. Right valve.
FIG. ii. Varicorbula amekiensis sp. nov. Syntype (1^.48255). x 2. Left valve.
Bull. B.M. (N.H.) Geol. 3, z.
PLATE 8
2a
2b
y-
5a
5b
8
PLATE 9
FIG. i. Clans (Bendeglans) costaeirregularis (Newton). Lectotype (L. 48324). x 2. Left
valve.
FIG. 2. Clans (Bendeglans) costaeirregularis (Newton). Syntype (L. 48325). x 2. Right valve.
FIG. 3. Clans (Amekiglans) costaenodulosis (Newton). Syntype (1^.48364). x 2. Left
valve.
FIG. 4. Clans (Amekiglans) costaenodulosis (Newton). Syntype (L. 48365). x 2. Right
valve.
FIGS. 5<z, b. Clans (Divergidens) triparticostata (Newton). Syntyps (1^.48322). x 2. 50 :
left valve.
FIG. 6. Clans (Divergidens) triparticostata (Newton). Syntype (L. 48321). X 2. Right valve.
FIG. 7. Clans (Divergidens) triparticostata (Newton). Lectotype (1^.48319). x 2. Left valve.
FIGS. 8a-c Fragum (Africofragum) newtoni sp. nov. Holotype ^48416). X 2. 8a : right
valve. 8b : left valve. 8c : right valve.
FIGS. 9«, b. Tivelina newtoni sp. nov. Syntype (L. 48498). x 2. Left valve.
FIGS. loa, b. Tivelina newtoni sp. nov. Syntype (L. 48499). x 2. Right valve.
FIGS. na-c. Pitar amekiensis sp. nov. Paratype (L. 48245). x i. Left valve.
FIG. 12. Pitar amekiensis sp. nov. Holotype (L. 48253). x I. Right valve.
FIG. 13. Kitsonia eocenica (Newton). Lectotype (L. 48192). x 4. Right valve.
FIG. 14. Kitsonia eocencia (Newton). Syntype (L. 48193). x 5. Left valve.
Bull. B.M. (N.H.) Geol. 3, 2.
PLATE 9
PLATE 10
FIG. i. Pitar amekiensis sp. nov. Holotype (L. 48253). x i. Right valve.
FIGS. 2a, b. Chionella (Microcallista) kitsoni (Newton). Lectotype (L. 48069). x 2. Right
valve.
FIG. 3. Chionella (Microcallista) kitsoni (Newton). Syntype (L. 48071). x 2. Left valve.
FIG. 4. Sinodiopsis coxi gen. et sp. nov. Holotype (L. 48250). x i. Left valve.
FIG. 5. Sinodiopsis coxi gen. et sp. nov. Paratype (1^.48249). x i. Right valve.
FIGS. 6a, b. Spisula (Crepispisula) amekiensis sp. nov. Holotype (L. 48224). x 2. Left valve.
FIG. 7. Spisula (Crepispisula) amekiensis sp. nov. Paratype (L. 482 19). x 2. Right valve.
FIGS. 8a-c. Varicorbula amekiensis sp. nov. Holotype (1^.48264). X 2. 8a : right valve.
8b : left valve.
Bull. B.M. (N.H.) Geol. 3, 2.
PLATE 10
-
6a
6b
8a
8b
8c
THE OLIGOCENE FLORA OF THE
BOVEY TRACEY LAKE BASIN,
DEVONSHIRE
M. E. J. CHANDLER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 3
LONDON: 1957
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THE OLIGOCENE FLORA OF THE BOVEY
TRACEY LAKE BASIN, DEVONSHIRE
BY
MARJORIE E. J. CHANDLER
Pp. 71-123 ; Pis. 11-17 '' 3 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 3
LONDON: 1957
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THE OLIGOCENE FLORA OF THE BOVEY
TRACEY LAKE BASIN, DEVONSHIRE
By M. E. J. CHANDLER
SYNOPSIS
The Bovey Tracey lignite deposit suggests an accumulation of plant debris much of which
was swept from a steep warm valley into an isolated lake basin lying in Palaeozoic strata.
The lake was surrounded by marshland and tree-covered slopes which contributed to the fossil
flora. Previous work on the beds and their plant remains is shortly reviewed, and in a few
instances earlier determinations have been corrected. New material has been identified. The
rich abundance of Sequoia, Osmunda and Calamus is noted and there are a number of other
species represented by a few specimens only. Attention is called to the possibilities inherent in
the study of pollen from Bovey. Thirty-one families are listed from fruits and seeds and there
are at least thirty-eight species.
Many Tertiary species have a long range in time and cannot therefore be useful in determining
age. But the slow replacement of some species and genera by others with the passing of
successive Tertiary stages may afford a clue. The age which such evidence suggests for Bovey
is Middle Oligocene.
INTRODUCTION
A revision of the fossil flora of Bovey Tracey is long overdue. Hitherto, knowledge
of the deposit has been based largely on the work of Heer (1862) and Pengelley
(1863) and to a lesser extent on that of C. & E. M. Reid (1910). The first-named
workers carried out an exhaustive examination of the beds at a time when the coal
pit of Bovey Tracey was accessible owing to economic enterprise. Peculiarly good
exposures were then available. The sections were described by Pengelley and the
flora by Heer.
When the Bovey pit was no longer worked for fuel it became largely obscured
owing to flooding and to the growth of vegetation on the upper slopes. Yet
C. & E. M. Reid were able to procure material from sections near the surface of the
pit to which they added a few fruits collected in a newer pit at Heathfield owned
by Messrs. Candy & Co. They also re-examined what remained of Heer's material,
and published a revised account of the flora in 1910.
Shortly before the first world war the old Bovey pit was again worked, this time
by a German firm, but since 1914 it has remained untouched. E. M. Reid and
Chandler visited it again in 1932. Certain species were then available in rich
abundance (Sequoia, Osmunda, and Microdiptera an extinct genus of Lythraceae)
in some shaley beds just above the water line. There were also seeds in somewhat
GEOL. Ill, 3. 7
74 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
greater variety in sandy beds a few feet higher in the section. Both levels were at
the western end of the pit where at the very top a few seeds of Stratiotes
were obtained. The eastern end of the south face was also exposed but was so deeply
weathered that it yielded nothing. In a pit at Kingsteignton Sequoia was found but here
there was little time for prolonged search. In Candy's pit at Heathfield which was still
operated there were some good seeds and fruits in clays above a well-marked lignite
band. The clays known to the firm as " best black clay " were examined on the
spot and blocks kindly supplied later by Messrs. Candy were washed and sifted.
Samples supplied from the middle and lower beds at that time exposed were unpro-
ductive. Thus almost all the plants at present known come from the two large
pits at Bovey and Heathfield.
These Devonshire lignitic beds differ from all other plant-producing localities
in the South of England Tertiary Beds in that they occur in isolation in a deep
rock-basin formed of Palaeozoic strata. Other plant beds lately examined are inter-
bedded among marine strata and so can be dated within certain limits at least by
marine organisms. The Bovey basin is shown in the i-inch Geological Survey
Map (26th sheet), reproduced by Pengelley, as about 8 miles in length with
a maximum breadth of about 3^ miles. It is filled by gravel, sand, and pottery
clays with abundant seams of lignite. The nature of these deposits indicates their
derivation from decayed Dartmoor granites. C. & E. M. Reid (1910) state that
the tectonic rock-basin is " surrounded on every side by steep bluffs which
immediately to the northwest slope upwards into the heights of Dartmoor ".
Examination of borings by Reid suggested that throughout the great thickness
of the beds, exceeding 500 feet, the same flora occurs so that they appear to belong
to a single geological series. The beds near the surface at Bovey appeared to be
equivalent to those near the base of the boring at Heathfield which penetrated for
526 feet without reaching the bottom of the basin. Among the sediments are masses
of wood and other plant remains. Some may have been swept down from higher
ground, but the greater part are presumed by the Reids to have been derived from
the slopes of a forest-clad steep warm valley leading from Dartmoor and disgorging,
in the neighbourhood of the Bovey pit, into the old lake basin. Sequoia is easily
the dominant element in the vegetation of this valley. Ferns, such as Osmunda,
apparently flourished in the ravine or around the lake. Recent work shows the
presence of other marsh plants e.g. Caricoidea (Cyperaceae) , Myrica, Microdiptera
and Lysimachia. True aquatics are represented by Salvinia, Stratiotes, Potamogeton
and Brasenia. Climbing plants are represented by vines and Rubus. Trees and
shrubs such as Nyssa, the Lauraceae, Symplocos, Carpinus, Magnolia and Meliosma
probably overhung the water and dropped their fruits into it. The Heathfield pit
which lay nearer to the centre of the lake is less rich in Sequoia than the Bovey
pit, and shows a greater variety of types. Prolonged collecting by a local enthusiast
might prove very profitable here.
On the whole the flora is very limited but there is a great abundance of some
few forms, rarer species being represented by one or a few individuals only. This
may well be because dense Sequoia forests and Calamus jungle do not provide a
congenial habitat for a wide range of plants.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 75
PREVIOUS WORK ON THE FLORA
Although Heer described nearly sixty species, many of his determinations are
unsatisfactory as judged by modern standards. Some because his material was
poorly preserved or fragmentary as in the case of many of his leaves. This was
recognized by Heer himself as a cause of some uncertainty. In certain instances
his study of better material was too superficial, the possibilities inherent in intensive
detailed research being then not so well understood as they are at the present time.
Again to some extent comparable living forms were probably unknown. In spite
of these disadvantages, and to his lasting credit, a number of Heer's species have
stood the test of later critical study while it has now been possible to correct a few
erroneous determinations. A revision of Heer's leaves cannot be attempted in this
present work more especially as most of his specimens have decayed or seriously
deteriorated.
There are a few corrections also of the determinations made by C. & E. M. Reid
as will be apparent from the synonyms at the head of some descriptions in the
systematic part of this work. One or two plants named by them are omitted — the
evidence being inadequate — viz. Taxodium distichum (which may have been a
peculiarly preserved Sequoia cone-scale), Taxus baccata, Calla cf. palustris, Sagittaria
sp.?, Labiatae, Genus?, and Cornus.
In the following pages only earlier records about which there is additional evidence
and new additions to the flora based on fresh and reliable evidence are included.
A recent source of knowledge which calls for a separate study may be mentioned,
namely pollen. In 1932 Reid and Chandler obtained a few anthers from which
pollen grains were extracted, identified and photographed by Dr. J. B. Simpson.
These are included in this paper.
In 1950 the late Nils-Erik Ross examined a sample of lignitic clay. He wrote
from Uppsala (June, 1950) " The sample from Bovey contained an interesting
micro-flora. There are plenty of pollen-grains of Calamus indicating a dense Calamus
jungle at the time of the sedimentation of the clay. Other specimens of pollen and
spores I have provisionally determined as ? Picea, Pinus (several spp.), Podocarpus,
Engdhardtia (pollen-grains and star hairs from the cuticle), Myrica, Symplocos
(2 spp.), Ilex, Quercus (comparable with some evergreen spp., e.g. Q. ilex], cf. Tilia,
Ericaceae, Ulmaceae, Caprifoliaceae, PAnacardiaceae, Osmunda, Schizaeaceae, etc."
Although Ross's evidence has not been published, it is interesting to note in
passing that it confirms the determination of Calamus, Myrica and Symplocos from
other organs, gives support to Simpson's Tilia and confirms Heer's Quercus and
Ericaceae (the two last based on leaves).
The record of Engelhardtia is of interest because of its occurrence in the Oligocene
Bembridge Beds. Schizaeaceae range from the Palaeocene to the Bembridge but
it may be recalled that the Oligocene species of Anemia is different from that of the
Lower Bagshot and Lutetian (Bournemouth Freshwater Beds). It would be helpful
to know what genera and, if possible, what species Ross found of this family at Bovey.
Dr. J. W. Franks of the British Museum (Natural History) is presently continuing
the study of Bovey pollen and it is hoped that the important results of Ross's
investigations will be included in a forthcoming paper with additional evidence.
7& THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
THE MODE OF PRESERVATION OF THE FOSSILS
All specimens collected in recent years are carbonaceous entities. In the majority
of cases they are separable from the matrix of clay or sand only after prolonged
boiling with soda. All appear to have undergone intense compression so that they
are not only distorted and flattened but frequently the carpellary coats or testa have
been so carbonized that they break with a glassy fracture and the cell structure
may be more or less obliterated in many cases. So far as the fruits and seeds are
concerned the best specimens are not found in the lignite bands themselves but are
associated with the interstratified beds. It should, however, be noted that in
Ross's letter he asked for samples of pure lignite in which, he stated, the pollen
grains are much better preserved than in clays. It was clear when collecting in
the Bovey pit in 1932 that the richest variety of fruits and seeds occurred in the
coarser sandy beds. This is commonly the case in Tertiary deposits, for in sandy
beds the seeds and other remains are less distorted and more readily released and
therefore less liable to be damaged. Unfortunately sandy exposures are very
restricted in extent.
The laminated clay beds chiefly enclosed masses of matted Sequoia and Osmunda
at the horizons lately available, and very little else. Prolonged search of the washings
from such layers were rarely rewarded by any new discovery and were discontinued.
The leaves described by Heer were, in all probability, impressions with mummified
leaf-substance. Such at least is suggested by the mode of preservation of the
abundant Sequoia and Osmunda remains.
A revised list of Bovey plants is given below :
Family and its approximate
Genus and species
climatic range, after R. Good,
ct — tropical or chiefly
tropical,
tc = temperate or chiefly
temperate.
b — both tropical and
temperate.
Osmundaceae . b
Salviniaceae . . b
Taxodineae tc
Potamogetonaceae . b
Hydrocharitaceae . b
Cyperaceae . . b
Calameae (Palmae) . ct
Zingiberaceae . . ct
Myricaceae . . b
Betulaceae tc
Fagaceae . . tc
Ulmaceae . tc
Other fossil occurrences
in the South of England
Osmunda lignitum Giebel
Salvinia boveyana n. sp.
Sequoia couttsiae Heer
Potamogeton tenuicarpus C.
& E. M. Reid
Stratiotes websteri (Bgt.)
Caricoidea nitens (Heer)
Calamus daemonorops (Ung.)
Spirematospermum wetzleri
(Heer)
Myrica boveyana (Heer)
Corylus sp.
Carpinus boveyanus (Heer)
Fagus minima n. sp.
Zelkova boveyana n. sp.
Bournemouth Freshwater.
Genus, Lower Headon.
Palaeocene ? Lower Bagshot
(Studland) to Hamstead
Beds (Hamstead).
Hamstead Beds.
Hamstead Beds.
Bournemouth Freshwater ;
Cliff End, near Mudeford.
Lower Headon to Bern-
bridge.
Bournemouth Marine ; Cliff
End ; Lower Headon.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 77
Family and its approximate
climatic range, after R. Good.
ct = tropical or chiefly
tropical,
tc = temperate or chiefly
temperate.
b = both tropical and
temperate.
Moraceae . ct
Genus and species.
Other fossil occurrences,
in the South of England.
Nymphaeaceae .
Magnoliaceae
Lauraceae .
Capparidaceae .
Hamamelidaceae
Rosaceae .
Leguminoseae
Rutaceae .
Sabiaceae .
Vitaceae
Tiliaceae .
Lythraceae
Nyssaceae .
Myrtaceae .
Mastixioideae
Primulaceae
Symplocaceae
Solanaceae
b
b
ct
ct
tc
tc
b
ct
ct
ct
tc
b
b
ct
ct
tc
ct
Moroidea boveyana n. gen.
et sp.
Brasenia ovula (Bgt.)
Magnolia boveyana n. sp.
Various genera not deter-
mined
Capparidispermum bovey-
anum n. gen. et sp.
2 spp. or genera
Rubus micro spermus C. &
E. M. Reid
Genus ?
Rutaspermum exaratum
(Heer).
Meliosma reticulata (C. &
E: M. Reid)
Parthenocissus britannica
(Heer)
Parthenocissus boveyana n.
sp.
Vitis hookeri Heer
Vitis stipitata n. sp.
Tilia sp.
Microdiptera parva n. gen.
et sp.
Nyssa boveyana n. sp.
Myrtospermum boveyanum
n. gen. et sp.
Myrtospermum dubium n. sp.
Myrtospermum sp.
Mastixia boveyana n. sp.
Lysimachia boveyana n. sp.
Symplocos anglica n. sp.
Symplocos headonensis
Chandler
Solanispermum reniformis
n. gen. et sp.
Genus, Lower Headon.
Bournemouth Marine to
Hamstead.
Genus, London Clay and
Bournemouth Freshwater
Beds.
Palaeocene to Lower Hea-
don, Bembridge.
Family, Lower Bagshot to
Bartonian.
Family, Palaeocene to Lower
Headon.
Upper Headon.
Genus, London Clay to
Upper Headon.
Genus, London Clay ; Lower
Bagshot (Lake) ; Bourne-
mouth Freshwater Beds.
Genus, London Clay to Lower
Headon.
Genus, Palaeocene to Lower
Headon.
Bournemouth Freshwater.
Bournemouth Marine Bed
Lower and Upper Headon.
Genus, London Clay.
Family, Palaeocene to Upper
Headon.
Family, London
Lower Headon.
London Clay.
Clay to
Genus,
Genus, Oldhaven to Lower
Headon.
Species, Lower Bagshot to
Lower Headon.
Lower Bagshot to Bourne-
mouth Marine and Cliff
End Beds.
78 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
The grouping of the families into tropical or chiefly tropical, temperate or chiefly
temperate, and both tropical and extratropical is based on R. Good's classification
(1947). It is inevitably to some extent arbitrary. Thus it might be argued should
the Magnoliaceae, Lauraceae and Vitaceae be classified as chiefly tropical or wide-
ranging in that they have extensions into cooler regions ? or should the Rosaceae
and Hamamelidaceae be grouped as temperate or wide-ranging for they have limited
extensions into warmer areas? On the whole, however, the grouping appears to
give a representative picture and suggests a warm flora, for of thirty-one families,
eleven are chiefly tropical and only eight temperate. But for reasons shortly to
be stated these figures should not be unduly stressed. In this connexion it may be
noted that certain genera of temperate type e.g. Sequoia, Carpinus, are associated in
Eocene beds with a well-defined warm flora so that, in the past, they may have
had extensions into subtropical or tropical climates which have been lost at the
present day. Families such as Hamamelidaceae still retain limited extensions ;
hence their presence in a warm flora is not surprising. Further, the constant
association in the fossil record [in part awaiting publication] of the genus Micro-
diptera and the species Solanispermum reniformis with warm floras points to them
as warm representatives of the wide-ranging families Lythraceae and Solanaceae.
THE AGE OF THE BOVEY FLORA
Palaeobotanists have differed considerably in their views on the age of this isolated
undated deposit.
Heer, although on slender evidence, equated the Bovey lignites with the Middle
Oligocene Hamstead Beds of the Isle of Wight (18620). Gardner (1879 : I^)
regarded the flora as identical with the Eocene Bournemouth Freshwater Beds
with which, it is true, they have a number of genera in common e.g. Osmunda,
Calamus, Stratiotes, Magnolia, Rubus, Meliosma, Vitis, Mastixia and Symplocos.
C. & E. M. Reid (1910) compared the flora with that of the Wetterau. Reid
& Chandler (1926 : 25-28) used the percentages of European genera present in a
series of floras ranging from Eocene to Pliocene among which was Bovey with
38% (so far as it was then known). They reached the conclusion that the Bovey
flora was approximately of Middle Oligocene age and was considerably younger than
the Upper Eocene Hordle flora, and older than the Mio-Pliocene and Pliocene
floras of Pont-de-Gail and Reuver. The figure for Bovey would now be more like
44% still well below that for the Pliocene (47-58%). It cannot, however,
be stressed too emphatically that in dealing with small floras of relatively few
genera and species statistical methods may not be wholly satisfactory. A few new
finds or fresh determinations can alter the figures appreciably. It therefore seems
inadvisable to lay too much stress upon such methods in cases where the element
of chance may influence results to a large extent.
Another point to be borne in mind is this : The utmost caution is needed in
attempting to date one of the older Tertiary floras by its plant remains. Long
research on Eocene and Oligocene plants has underlined the fact that many species
have a far longer range in time than was once supposed. There are indications of
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 79
this in the Bovey table given on pp. 76, 77. Some examples are quoted here, using
species found at Bovey.
Sequoia couttsiae is a typical instance. The species is well known from twigs,
cones, seeds and cuticle. Its cuticular structure is distinctive. Its range in England
is certainly from the Lower Bagshot to Hamstead Beds and may be from the
Palaeocene.
Carpinus boveyanus is known in the Bournemouth Marine Beds, the Cliff End
Beds and the Lower Headon, possibly also in the Upper Headon.
Solanispermum reniformis ranges from the Cuisian, through the Bournemouth
Freshwater and Marine Beds to the Cliff End Beds.
A number of species not found yet at Bovey could also be used to illustrate the
long range of some older Tertiary types. For example the following awaiting record
or description which have been found in the Lower Bagshot of Lake or Arne, the
Lower Headon of Hordle, and sometimes in intermediate beds : Caricoidea obscura
(Cyperaceae), Protoaltingia (Hamamelidaceae) , Phellodendron, Natsiatum, Myrtoidea,
Myrtospermum variabile, Styrax, Eomastixia, Mastixicarpum.
It may be dangerous to rely on the presence or absence of particular species in
trying to assess age, for it is impossible to determine the part played by chance
preservation and discovery in the records of a flora.
Nevertheless time and further research may throw fresh light on the problem of
Tertiary plants as time-indicators. There is already some hint that although long
range of species is a common phenomenon, there is also a gradual replacement of
some species or genera by others during a long period.
Up to the present (and it is necessary to stress this qualification) Spirematospermum
wetzleri has not been found in this country below the Lower Headon (or possibly
the Barton Beds). It occurs at Hordle, and in the Middle Oligocene Bembridge
Beds. It also occurs at Bovey.
Bmsenia ovula (Bgt.) appears in the Bournemouth Marine Beds and Hengistbury
Beds and persists through the Barton, Lower and Upper Headon, Bembridge and
Hamstead Beds. Below the Bournemouth Marine Beds in the Freshwater
(Lutetian?) series and the Lower Bagshot, its place appears to be taken by another
characteristic and readily recognizable water-lily (awaiting description) which
disappears after the Lutetian. It is Brasenia ovula which occurs at Bovey.
Similarly, Microdiptera parva, an extinct genus of Lythraceae, occurs in the
Bournemouth Marine Beds and the Lower and Upper Headon. In the Bournemouth
Freshwater Beds there is a different species of Microdiptera. It is M. parva which
occurs at Bovey.
These records suggest that the Bovey flora is not older than the Bournemouth
Marine Series.
Again Potamogeton tenuicarpus is known in the Hamstead Beds (possibly also in
the Upper Headon). On the other hand a highly distinctive spiny species, P.
pygmaeus ranges, according to present knowledge, from the Bournemouth Marine
Beds, through the Lower Headon only up to the Bembridge Beds, after which P.
tenuicarpus is found. P. tenuicarpus is the species found at Bovey.
Stratiotes is a characteristic common Tertiary genus where water-plants are
8o THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
preserved. The successive species appear to have a very limited range in time
(Chandler, 1923). S. websteri (Bgt.), which occurs at Bovey, is found in the Hamstead
Beds, also in the Oligocene Cyrena-Marls of Offenbach on the Main. It does not,
so far as is known, range into the Miocene which is characterized by S. kaltennord-
heimensis Zenker. It is markedly different from the somewhat older Lower and
Upper Headon species S. headonensis Chandler. In deposits older than the Lower
Headon a much smaller species has now been found. It will be described as a variety
of 5. zinndorfi Kirch.
5. zinndorfi was thought by Kirchheimer to be Oligocene. The variety is of
Eocene age and disappears before the Lower Headon. The occurrence of the large
typical 5. websteri at Bovey therefore supports a Middle Oligocene age for the
deposit.
On the whole such evidence as there is indicates an Oligocene, and perhaps a
Middle Oligocene age for the Bovey flora.
One other genus perhaps deserves special mention, viz. Mastixia. It, or its
close allies, are among the most abundant Eocene plants ranging from the London
Clay through the Lower Bagshot, Bournemouth Freshwater and Marine, Cliff End and
Hengistbury Beds to the Bartonian and Lower Headon. So far they have not been
found in the Upper Headon or younger beds.
According to Kirchheimer (1936 : 125), who based his conclusions on evidence from
successive beds of the German Brown Coal, the genus disappeared in Europe after the
Oligocene. The sparse occurrence of a species, quite specifically distinct from any
of the British Eocene species, indicates, therefore, a pre-Miocene and post-Eocene
age for the Bovey lignites.
Further work on Bovey pollen may serve to throw more definite light on this
problem of age. At present there appears to be no other source from which
information is likely to come.
In conclusion it should be recorded that the writer had the advantage of collabora-
tion with E. M. Reid in the study of some at least of the material collected in 1932.
The Introduction to this paper was seen and approved by the late Mr. W. N. Edwards
shortly before his death and Mr. F. M. Wonnacott has given invaluable help in the
preparation of the manuscript for the press.
PTERIDOPHYTA
Order FILICALES
Family OSMUNDACEAE
Genus OSMUND A Linnaeus
Osmunda lignitum (Giebel)
(PI. u, figs. 1-6)
1862. Pecopteris (Hemitelia ?) lignitum Giebel : Heer, p. 1047, pi. 56, figs. 2-8.
1882. Osmunda lignitum (Giebel) : Gardner, p. 49 (reference to Bovey material).
DESCRIPTION. Pinnules : These have been fully described by Heer (1862).
Sporangia : Sub-globular, dehiscing longitudinally on one side from pole to
pole, the margins of the sutures formed by about three rows of narrow elongate
longitudinally aligned cells. Walls elsewhere formed of a layer of thiek-walled,
usually longitudinally elongate cells, often about 0-025 by °'°5 mm- m diameter.
Annulus a raised patch of thicker- walled more opaque cells at one pole adjacent to
the distal end of the split, occupying about half the length of the sporangium.
Diameter of sporangium about 0-45-0-55 mm.
Spores : Finely granular, sub-globular, tetrahedral, about 42 to 62 /* in diameter
(actual measurements, 50 x 45 fi and 62 X 42 fi).
REMARKS AND AFFINITIES. Almost as abundant in the Bovey coal pit as Sequoia
with which it is mixed in certain seams forming matted masses. Slabs of matrix
from such seams are very fissile on drying so that they quickly disintegrate. In
addition to a number of such blocks with barren fronds there are scanty remains
of fruiting organs in the washed residues. The disc-like annulus points to the family
Osmundaceae and to the genus Osmunda. Spores of the living 0. regale appear to
be somewhat thinner-walled with finer spines than those seen in the fossil. They
measured 45 /* in diameter. Heer found no sori in spite of repeated search
(1862 : 1047). Later his work was criticized by Gardner (1882 : 49) who identified
the Bovey barren pinnules with similar specimens from Bournemouth, and with
others from continental localities. Gardner pointed out that the absence of sori
on the pinnules should have directed Heer's attention to Osmunda in which the
fertile pinnules are segregated at the ends of the fronds.
The name 0. lignitum has been used frequently for indistinguishable barren
pinnules from widely scattered European localities. Probably, like Sequoia
couttsiae, the species was wide-ranging in space and time in the older Tertiary.
There seems no reason why the barren pinnules and isolated sporangia should not
belong to a single species.
Family SALVINIACEAE
Genus SALVINIA Linnaeus
Salvinia boveyana n. sp.
(PI. n, figs. 7-11)
1910. Spadix of aroid ? C. & E. M. Reid, p. 173, pi. 16, fig. 57.
DIAGNOSIS. Sporocarps sub-globular enclosing at least twenty to fifty globular
male sporangia. Microspores tetrahedral, the majority 0-025 mm- (25 /*) m diameter,
ranging from 0-02 to 0-03 mm. Vegetative parts unknown.
HOLOTYPE. A sporocarp. Brit. Mus. (N.H.), No. ¥.33834.
DESCRIPTION. Vegetative parts : Unknown.
Sporocarps: Sub-globular (incomplete). Walls thin, as shown by the manner
in which the sporangia distort the wall (PI. n, fig. 7), structure obscure, but small
equiaxial cells 0-006-0-009 mm- m diameter can be detected on the much corroded
surface. In many places abrasion has actually exposed the sporangia so that their
82 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
cell-structure and contents tend to obscure and confuse the cell-structure of the
sporocarp. As preserved the sporocarps enclose from about twenty to fifty sporangia,
but there must originally have been more for all are incomplete.
Sporangia : Globular, thin- walled, 0-15-0-2 mm. in diameter, walls apparently
one cell thick, formed of coarse polygonal cells. The cell-walls are obscure either
because they have decayed so that they are represented by impressions only, or
else because they are very thin. The sporangia are hollow in the middle, formed
of a frothy substance within which the microspores occur in fours, each spore with
characteristic triradiate markings. The average spore diameter is 0-025 mm-
(25 fi) but some are as much as 0-03 mm., others 0-028 mm., a few only are smaller
than 0-02 mm. and those are probably immature.
REMARKS AND AFFINITIES. Five sporocarps with numerous sporangia. The
microspores agree closely in size with an unnamed species described by Kirchheimer
(1931 : 102-113) from the Upper Miucene Brown Coal of Beuern in Vogelsberg
(spore diameter 25 /*), also with Salvinia hassiaca Kirchh. from Garbenteich (spore
diameter 26 /*), a species of similar age (19300 : 203). Among living species of which
spore measurements are available, the nearest to the fossil is S. auriculata Aubl.
(spore diameter 27 fi), a species from Central and South America and the West
Indies. In the living 5. natans (Linn.) the microspores measure only 18 /*, and in
S. ancillata Roxb. 22 /*, while in S. oblongifolia Mart they measure 23 /*. These
measurements are taken from Kirchheimer's detailed researches on the fruiting
organs of fossil Salvinia in German Brown Coal (1931) ; there are other papers by
this author on the fossil occurrence of the genus (1928 ; 1929 ; 1930 ; 1932 ; 1937).
His investigations are concerned chiefly with Miocene material, but an Oligocene
species was figured from the Niederpleis Brown Coal in 1937 (p. 897, text-fig. 4) ;
unfortunately it does not permit of satisfactory comparison ; the dimensions of the
microspores were not given.
No fruiting organs have been described previously from Britain, but vegetative
parts are known from the Lower Headon of Hordle (Chandler, 1925 : 10, pi. i,
figs. la-d ; text-fig, i) and also occur in numerous continental localities (Florin,
1919 : 243).
The living genus Salvinia is distributed throughout the north temperate zone in
the Old and New Worlds. It occurs also in the East Indies, Tropical Africa, the
Mascarene Islands, South and Central America and the West Indies.
GYMNOSPERMAE
Order CONIFERALES
Family TAXODINEAE
Genus SEQUOIA Endlicher
Sequoia couttsiae Heer
1862. Sequoia couttsiae Heer, p. 1051, pi. 59 ; pi. 60, figs. 1-46 ; pi. 61.
18620. Sequoia couttsiae Heer : Heer, p. 372, pi. 18, figs. 1-7.
1883. Sequoia couttsiae Heer : Gardner, p. 36, pi. 6, figs. 7, 10-17.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 83
1910. Sequoia couttsiae Heer : C. & E. M. Reid, p. 170, pi. 15, figs. 23-27.
1921. Sequoia couttsiae Heer : Chandler, p. 457.
1922. Sequoia couttsiae Heer : Chandler, p. 385.
1923. Sequoia couttsiae Heer : Bandulska, p. 257, pi. 21, figs. 31, 32.
The external morphology has been well described and figured by Heer (1862,
1862^) and by C. & E. M. Reid (1910). It may be added here that the seed-body
can be either straight or curved whereas Heer referred only to curved seeds. Twigs,
cones, detached cone-scales and seeds are common at Bovey, and occur less frequently
at Heathfield and Kingsteignton. The cuticle has been described and figured by
C. & E. M. Reid (1910) and Bandulska (1923). New details of cuticular structure
are here recorded.
The stomata are borne on both sides of the leaf.
Cuticle of upper surface : This has two well-marked stomatal bands one on each
side of the midrib showing thickly scattered stomata variously oriented, a few being
longitudinal or transverse but the majority oblique. They have a tendency to be
arranged in short longitudinal rows, but this is not a conspicuous feature. In the
broader parts of the leaves three or four, sometimes more, stomata may be arranged
abreast. Three bands of epidermal cells occur outside the stomatal bands, a narrow
band frequently about eight cells wide at each margin, and a broad median band over
the midrib which may be about twenty-five cells wide at the leaf base. The marginal
bands unite with one another and sometimes with the median band also at the apex
of the leaf. Occasionally the median band dies out a short distance below the
apex and then the two stomatal bands unite, or almost unite, just below the apex.
Towards the leaf-tip the epidermal cells nearest the margin tend to diverge in a
fan-like manner. Epidermal cells between the stomata very variable in form and
size, frequently more or less equiaxial, not infrequently transversely elongate and
aligned especially between stomata in the same longitudinal row. Stomata rarely
share any auxiliary cells but are often sufficiently close together for the auxiliaries
of adjacent stomata to be in contact without intervening epidermal cells. Stomatal
pores oval to sub-quadrangular except on the decurrent flanges of the leaf where
they may be sub-circular or broadly oval. Guard cells thinly cuticularized, the
slit between them conspicuous, occasionally they show fine 'granulation. The outer
pore ("aussere atemhohle" of Florin) is normally about 0-02 mm. long, occasionally
0-03 mm. The auxiliary (— subsidiary) cells are considerably thickened where they
unite with the guard cells, they vary from four to six and are arranged in a ring
around the guard cells, often end to end, but there is a tendency to vary in size and
shape so that the regularity of the ring is sometimes destroyed. Sometimes two
concentric cells ( or a cell which has divided into two by a longitudinal partition)
occur in parts of the ring. The epidermal cells outside the stomatal bands are
longitudinally elongate and aligned, in some leaves parallel-sided with rectangular
or oblique end walls, in others tending to be broader at the middle than at the ends
but such frequently have flat straight end walls. Cells brown and much cutinized
with a fine inconspicuous reticulate thickening at least in some cells. At the base
of the leaf the walls are always thin and colourless. Cells of the marginal epidermal
band often appreciably broader than those of the middle. Typical leaves measured :
84 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
(i) length, 0-7 mm.; breadth, i mm.; (2) length, 0-64 mm.; breadth, 0-48 mm.
Cuticle of lower surface : Also has two stomatal bands which are broader and less
sharply defined than those of the upper surface and do not extend so far towards the
tip of the leaf. The bands broaden considerably towards the base where sometimes
they almost merge, but at the extreme base they are reduced to one or two rows of
large oblique stomata adjacent to the leaf margin separated by a wide triangle of
thin-walled epidermal cells with large cavities. The stomata are more distant
and widely scattered than those of the upper surface with a tendency to occur in
short longitudinal lines often separated from neighbouring lines by several rows of
longitudinal ordinary epidermal cells ; they are transversely, obliquely, and longi-
tudinally oriented, but on the whole oblique orientation appears to predominate.
The " outer pore " may be oval but is frequently narrow-oval, usually about 0-02
mm. long or a little longer. The slit between the guard cells is clear. Auxiliary
cells vary from four to six and are arranged in a ring around the " outer pore ",
but there is a much more marked tendency for the cells in the ring to be uneven in
length, shape, and size than on the upper surface. There is also a marked tendency
for one or two of the cells to be prolonged into a row of ordinary epidermal cells
adjoining the stoma and from these they can scarcely be distinguished. Auxiliary
cells of adjacent stomata may be contiguous and are occasionally shared by adjacent
stomata, more often, epidermal cells intervene. There is sometimes a double
ring of auxiliary cells in part of the circumference. The auxiliary cells are not
more conspicuously cutinized than the ordinary epidermal cells except where they
abut on the guard cells ; here there is a thickening of the wall. The ordinary
epidermal cells in the stomatal bands are often irregularly arranged and are frequently
as broad as long, sometimes longitudinally elongate and aligned, but often between
two stomata in the same linear series transversely elongate and aligned. Marginal
and median bands of ordinary epidermal cells unite below the apex to form a broad
triangular apical area without stomata. The epidermal cells are frequently parallel-
sided, longitudinally elongate and aligned ; in the median band they often have
rectangular or oblique end walls ; in the marginal bands they are less frequently
rectangular, usually also narrower and longer than in the median. Also in the
median band they tend to be shorter at the apex of the leaf than below. In one
slide they appeared to be beset with small pits, about 0-002 mm. broad, irregular
in shape and distribution. They are occasionally very slightly sinuous, especially
their transverse walls. Such sinuosities are seen often on the decurrent leaf base
in the median band.
There is a greater resemblance to Sequoia gigantea in the arrangement of the stomata
on the upper-side of the leaf than to Sequoia sempervirens owing no doubt to the
greater similarity of form.
REMARKS. The species has lately been recognized at Studland (Lower Bagshot),
the Bournemouth Marine Beds of Southbourne, the Cliff End and Hengistbury Beds.
It is also represented in the Lower Headon at Hordle, and the Upper Headon at
Colwell Bay.
Detailed accounts and illustrations of these cuticles are included in a forthcoming
monograph on the Lower Bagshot flora where the species is fully discussed.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 85
ANGIOSPERMAE
Class MONOCOTYLEDONES
Family POTAMOGETONACEAE
Genus POTAMOGETON Linnaeus
Potamogeton tenuicarpus C. & E. M. Reid
(PI. n, figs. 12-14)
1910. Potamogeton tenuicarpus C. & E. M. Reid, p. 173, pi. 16, figs. 53, 54.
DESCRIPTION. Endocarp : Broadly obovate, originally somewhat inflated
(now much flattened), curved through almost a complete circle about a circular or
oboval central depression, the curved area forming the locule ; dorsal margin semi-
circular, ventral margin convex above and below, conspicuously indented between
the ends of the limbs i.e. between the convexities at a distance of about one-third
of the length from the base of the fruit ; sometimes remains of a small spine can be
seen just above the indentation. Style small, patent, terminal on the ventral
margin. Surface conspicuously ridged, one ridge outlining the central depression,
another the dorsal margin adjacent to the keel, a third down the middle of the keel ;
ridges sharp, forming small flanges. Keel broad reaching from the base almost to
the apex, with a groove on each side of the median ridge. Surface cells irregular
in shape with the long axes diverging from the central area but also showing an
alignment parallel with the curvature of the endocarp ; cells averaging in size about
0-012 by 0-02 mm. Length of endocarp, about 1-1-5 nun.; breadth, 0-75-1-2 mm.
Seed : (Formerly described as the embryo, C. & E. M. Reid, 1910.) Narrow,
elongate, curved in accordance with the curvature of the locule ; testa smooth,
shining, light brown, semi- translucent, the square cells measuring 0-017 nim. across
and aligned parallel with the direction of curvature.
Pollen : Preserved in detached anthers and determined by Dr. J. B. Simpson.
Smaller than that of P. natans, finely reticulate all over with a marked invagination
on one side of the grain exposed when the intine had been cleaned out (PL n, fig. 14).
Probably belongs to this species ; the only one present.
REMARKS AND AFFINITIES. This species is re-described above in greater detail.
The form and structure of endocarp, seed (and according to Dr. Simpson, pollen
also) indicate the presence of a species of Potamogeton. The living P. cristata
Regel & Maack is almost equally small but does not otherwise resemble the Bovey
fruit. The breadth of the dorsal keel in the Bovey fossil shows that in life the locule
was probably more or less triangular in cross-section, but the thin though coriaceous
carpel has collapsed and been variously distorted by pressure in fossilization. The
same species occurs in the Hamstead Beds of the Isle of Wight.
P. Pygmaeus Chandler from the Upper Eocene of Hordle and the Oligocene
Bembridge Beds, Isle of Wight, is distinguished by the normally smaller size, the
convexity of the ventral margin with median prominence, and especially by the
spines on the keel (bases only preserved except in impressions).
86 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Family HYDROCHARITACEAE
Genus STRATIOTES Linnaeus
Stratiotes websteri (Brongniart)
(PI. n, figs. 15-19)
1862. Carpolithes websteri (Brongniart) : Heer, p. 1075, pi. 70, fig. 6.
1910. Stratiotes websteri (Brongniart) : C. & E. M. Reid, p. 172.
1920. Stratiotes kaltennordheimensis (Zenker): E. M. Reid, p. 60, pi. 3, figs. 8, 9.
1923. Stratiotes websteri (Brongniart) : Chandler, p. 128, pi. 5, figs. 10, n ; pi. 6, figs. 2, 3.
DESCRIPTION. Seed : Oblong with rounded ends, hooked or slightly sigmoidal
in outline, laterally flattened. Keel narrow, beaked at the apex in sigmoidal seeds,
usually rounded externally, not continued round the base but merging gradually
into the collar. Collar usually large, rounded, testa woody ornamented over the
body with interrupted longitudinal ridges which run from neck to apex where they
curve towards the keel and converge to the raphe ; pitting fairly uniform on body,
collar and sides of keel, typical pits measuring about 0-05 mm.; along the dorsal
margin of the keel the pits are much finer. Keel broadening gradually towards the
apex as seen in longitudinal section of the seed (PI. n, figs. 18, 19).
Micropyle basal or sub-basal, very slightly oblique, hilum dorsal, associated with
the beak at the apex of the keel in the few specimens available. Raphe short,
transverse. Digitate cells of the interior of the keel straight, parallel to the length
of the keel.
Length of a large seed in Sedgwick Museum, Cambridge, 6-8 mm.; breadth,
2-75 mm. Length of a seed in the Geological Survey Collection 6-4 mm.; breadth,
3 mm. Length of a seed recently found at Bovey by the author, 5-25 mm.; maximum
transverse measurement of a seed flattened dorsi-ventrally, 3 mm.
REMARKS AND AFFINITIES. One seed and fragments of three others showing
respectively the raphe and the collar have been found lately at Bovey. Several
specimens from other collections have also been examined. The seeds have been
compared with specimens of Stratiotes websteri from the Hamstead Beds of the
Isle of Wight and are indistinguishable from that species.
Family CYPERACEAE
Section CARICOIDEAE
Genus CARICOIDEA nov.
A form-genus for fruits or endocarps belonging to the section Caricoideae of the
family Cyperaceae of which the nearer relationship is not known.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 87
Caricoidea nitens (Heer)
(PI. n, figs. 20-23 ; Text-fig, i)
1862. Carpolithes nitens Heer, p. 1078, pi. 70, figs. 15-23.
1910. Taxus ? nitens (Heer) C. & E. M. Reid, p. 172.
DIAGNOSIS. Originally globose with triangular calyx about 2-4 mm. in diameter ;
apex pointed, base with aperture closed by a plug about 0-8-0-9 mm. in diameter.
Epicarp shining, formed by longitudinally aligned cells with straight or sinuous
outlines. Endocarp 0-5-1 mm. thick. Diameter (crushed dorsi-ventrally) , 4-5 mm.
HOLOTYPE. A laterally compressed fruit ; also figured by Heer (1862, pi. 70,
fig. 20). Brit. Mus. (N.H.), No. ¥.33842.
DESCRIPTION. Fruit : Originally globose (now compressed sometimes laterally,
at others dorsi-ventrally), apex somewhat pointed, base somewhat truncate having
a triangular impression (as of a non-accrescent calyx), about 2-4 mm. in maximum
diameter, at the centre of which is a circular scar closed by a plug about 0-8-0-9
mm. in diameter. Surface of fruit and triangular impression (except over the plug)
shining, longitudinally striate (fine parallel striae varying in direction are the result
FIG. i . Caricoidea nitens (Heer) . Diagrammatic longitudinal section through specimen
dorsiventrally crushed, showing the cavity (c), thick wall (w) and the plug (p) filling the
basal canal, x 10.
»
of tensions in compression). This coat (epicarp) formed of irregular longitudinally
aligned cells with straight or sinuous outlines, 0-05 mm. or less in length, 0-016-0-025
mm. in breadth. This coat appears vitreous as seen in section and is intimately
fused with the subjacent wall. Endocarp, 0-5-1 mm. thick, formed of regular
parenchyma ; cells, 0-012 mm. in diameter, the layers lining the cavity being vitri-
fied. Locule small, often flattened by pressure, lined by irregular, slightly sinuous,
thin-walled cells, 0-25-0-5 mm. in diameter, and in one specimen by a series of parallel
transversely aligned cells (impression of testa or adherent testa). The locule
communicates with the exterior by a canal about 0-5-0-8 mm. broad and is closed
by the plug described above. Diameter of fruits, 4-5 mm.
REMARKS AND AFFINITIES. Seven fruits, two now broken. They are stated by
C. & E. M. Reid in their photographic records to be the originals of Heer's figures
(1862, pi. 70, figs. 15-23). It has been possible to identify the originals of figs.
GEOL. Ill, 3. 8
88 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
16, 18 and 20, but the other drawings are too inaccurate in detail to permit of further
identification of types.
In describing this species under the name Carpolithes nitens, Heer draws a com-
parison with seeds of Taxus. Later, C. & E. M. Reid (1910 : 172) refer the specimens
tentatively to this genus. The evidence does not support this ascription. The
characters indicate relationship with Cyperaceae, section Caricoideae (the shining
exocarp, basal triangular scar, basal canal and plug, and thick parenchymatous
wall). Similar surface cells occur in Cladium mariscus which differs in having a
much smaller fruit. No genus which corresponds closely with the fossil has been
found, while in most of the Caricoideae the exocarp is distinct from the wall of the
nut and not indistinguishable from it. The apparent fusion of the two in the fossil
is no doubt the result of the intense compression which has occurred.
Family PALMAE
Sub-section LEPIDOCARYINAE
Genus CALAMUS Linnaeus
Calamus daemonorops (Unger)
(PL 12, figs. 24-42)
1862. Palmacites daemonorops (Unger) Heer, p. 1056, pi. 55, figs. 7-15 ; pi. 60, figs. 50-53 ;
pi. 62.
1910. Palmacites daemonorops (Unger) : C. & E. M. Reid, p. 172, pi. 16, figs. 44-49.
DESCRIPTION. Spines: (PL 12, figs. 38-41). Common, either single or attached
to a piece of stem, especially at Bovey ; a few occur at Heathfield. They are black,
shining, thin, tapering to a fine point, varying in length from 4 to 50 mm., the longest
being only 3 mm. broad at the base. The spine has a shallow longitudinal furrow
on one surface ; frequently the spines are grouped in threes of which the middle
one is the longest. Often they are two, or solitary, while four, five, or even more
may occur in a group. Heer stated that they were distributed on a finely striate,
flat, minutely warty surface (Heer, 1862, pi. 55, figs, n, 12 ; pi. 62, fig. 7). In the
specimens recently examined this surface appears rough, formed of equiaxial cells
about 0-025 mm- m diameter.
Fruiting axes (PL 12, figs. 35, 36) : Abundant, some with scars only to which
bracts were formerly attached, others still carry striate overlapping bracts which
are usually broken at their free edges.
Flowers : Dioecious, surrounded by stiff coriaceous bracts and bracteoles as in
an amentaceous inflorescence. Male flowers (PL 12, figs. 32-34) with three pointed-
oval bracts having valvate aestivation. Usually the stamens have been shed but
one flower contained stamens with pollen. This was examined and photographed
by Dr. J. B. Simpson. In letters of 27 . iii . 38 and i . iv . 38 he provided the following
facts : " Pollen. Grains smaller than those of Tr achy carpus excelsis (the only
recent pollen available for examination) having a single furrow lined by a thin
membrane as in Trachy carpus , the membrane sometimes preserved in the fossils
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 89
but at other times destroyed. Also having the same type of irregular reticulations
in the exine " (see PI. 12, fig. 42). In his preliminary statement on Bovey pollen
grains the late Nils-Erik Ross reported abundant Calamus.
Female flowers or immature fruits : Numerous. One-loculed with three stigmas
(PI. 12, figs. 24-31). Better-developed specimens are ovoid or sub-globular (but
much compressed in fossilization), having six bracts, usually free with imbricate
convolute aestivation, arranged in two alternate whorls. Bracts longitudinally
striate through the occurrence of sub-parallel conspicuous nerves. Surface of fruit
formed by characteristic recurved scales of the Rotang palm type, frequently
almost obliterated owing to the highly compressed state of the material. Surface
of scales of oblong cells aligned in rows forming striae which diverge from the proximal
end of the scales and fan out to their thin slightly fringed edges. Inner wall of fruit
compact, formed of equiaxial cells o-oi2-o-oi6 mm. in diameter.
Seeds : Not seen in a fully developed condition, but C. & E. M. Reid (1910, pi. 16,
fig. 48) figured a seed partially enclosed by a fruit 6 or 7 mm. long. Heer, moreover,
stated that he had seen a drawing (natural size) by Pengelley, of a fruit which
measured 13 mm. in length and 10 mm. in breadth. The drawing is reproduced
by Heer (1862, pi. 60, fig. 50). Another fruit, recently examined by the author,
enclosed an immature seed (PI. 12, fig. 37) 1-75 mm. long and 1-5 mm. broad. The
seed was obovoid with two irregular longitudinal crumples on the dorsal face, and
with a sunk median area (chalaza ?) on the ventral face surrounded by a broad,
marginal, horse-shoe shaped, inflated area over which the cells radiate from the
centre. Cells from 0-025 to 0-03 mm. broad. No embryo-scar was seen.
REMARKS AND AFFINITIES. The evidence from all organs points to relationship
with the group Calameae of the sub-section Lepidocaryinae, family Palmae.
The fruits although so small and probably immature suggest Calamus as do the
pollen grains. Heer regarded the prickles as identical with organs from Laubach
in the Wetterau which linger (1860) described as Palaeospathe daemonorops on
account of their resemblance to the spines on the spathes of living Daemonorops.
Heer also identified them with similar organs from Salzhausen and Hessenbriicken
described by Ludwig (1860 : 86, pi. 20, figs. 2, 3) as Chamaerops feutonica. Palmacites,
however, is a form-genus containing different organs of palms which cannot yet be
ranged in well-defined genera. In view of the relatively full information now
available about the Bovey species it appears reasonable to place it in the palaeo-
tropical genus Calamus. Whether it should be referred to Unger's species
Daemonorops is open to question. But for the present it is retained in the species
after Heer.
Calamus daemonorops (Unger) ?
(PL 12, figs. 43, 44)
An obovoid seed, evidently immature, with two irregular longitudinal folds on
the dorsal side, and a narrow spathulate chalazal area on the ventral side, may
possibly be an isolated seed of Calamus. It appears to be related to Palmae as
evidenced by the general structure, possibly also by the presence close to the base
go THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
on the dorsal side of a small raised sub-circular scar, 0-3 by 0-2 mm. in diameter,
which may mark the embryo. The walls are 0-05 mm. thick, coarsely columnar
in section, the columns are about 0-016 mm. broad, but as the texture is very vitreous
they may not show their true structure. The surface is irregularly rugose, but its
cells are indistinct. Length of seed, 2-87 mm.; breadth, 1-6 mm.
Family ZINGIBERACEAE
Genus SPIREMATOSPERMUM Chandler
Spirematospermum wetzleri (Heer)
1862. Gardenia wetzleri Heer : Heer, p. 1069, pi. 69, figs. 1-6.
1925. Spirematospermum wetzleri (Heer) Chandler, p. 17, pi. i, figs. Sa-c ; text-fig. 5.
The characteristic spirally striate seeds were described and figured by Heer from
Bovey and referred by him to the genus Gardenia (Rubiaceae). More recent
researches by Chandler (1925) on similar material from the Eocene of Hordle demon-
strated that the true relationship lay with Zingiberaceae, a conclusion subsequently
corroborated by Kirchheimer (1936^ : 98 ; 1937^ : 50 ; 1 939*2 : 275). No further
specimens have been found in the Bovey Basin since Heer's discovery. The species
is also known from the Bembridge Beds (Reid & Chandler, 1926 : 84, pi. 5, figs.
6,7)-
DICOTYLEDONES
Family MYRICACEAE
Genus MY RICA Linnaeus
Myrica boveyana (Heer) pars
(PI. 12", figS. 45-48)
1862. Carpolithes boveyanus Heer, p. 1077, pi. 70, ? figs. 7-14 (in part).
Under the name Carpolithes boveyanus Heer may have included several species
or genera which in their highly compressed state are difficult to distinguish. All
are black, more or less flattened (sometimes laterally or marginally, sometimes
dorsi-ventrally). Heer's description and figures are inadequate although his figures
8 and 12 with " longitudinal furrow " suggest seeds of Zanthoxyleae showing the
typical hilar scar. The sections in figures 9 and 13, possibly also in figures 10 and
14 suggest either Myrica or Carpinus, probably the former.
In his description Heer points out that they are of " two different dimensions ;
some are 3 millims. long and 2\ millims. wide ; others are 4-4^ millims. long and
3 1 millims. wide".
An examination of hundreds of these small carpels suggests that while many
specimens cannot at present be sorted there is clear evidence in the material of two
genera viz. Myrica and Carpinus.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 91
Typical Carpinus shows the longitudinal fibre grooves of the persistent perianth.
The fibres terminate against a large basal scar where the fruit was formerly attached
to a wing-like bract. The surface of such specimens is usually smooth and shining.
Typical Myrica shows a more irregularly rugose dull surface with no clearly
defined large basal scar. Between these two types are others variously distorted,
which might belong to one or other. They cannot at present be separated with
certainty. Some, indeed, may be seeds, not fruits, and belong to other genera or
families not yet recognized.
The fruit of living Myrica is superior, the seed is erect and orthotropous, solitary.
There is a conspicuous basal placenta scar on the locule, and chalaza scar on the
seed. The form of the locule when the two valves have separated, as in germi-
nation, is similar to that of Carpinus, urceolate in outline, somewhat compressed at
right angles to the plane of dehiscence, but the wide sutures are very flat and more
close-textured in Myrica than in Carpinus. Unfortunately the fossil seeds, when
preserved at all, are much crumpled and do not well display the chalaza although
there are hints of its presence. The locule surface is usually much corroded so that
the placentation is not clear while evidence of a superior fruit is lacking. The
texture of the endocarp and the sutures nevertheless appear to point to Myrica
in many instances.
There may be more than one species of Myrica at Bovey. Some of the endocarps,
as Heer noted, appear so much smaller and more inflated than others. As there is
no clear line of demarcation under the conditions of preservation, all are grouped
for the present as Myrica boveyana (Heer). The relationship of these endocarps to
Myrica was recognized by Kirchheimer (1938 : 327, footnote).
DIAGNOSIS. Endocarp bisymmetric, sub-ovoid, usually somewhat compressed
and angled in the plane of symmetry. Locule not emarginate at the base. Basal
part of wall not thickened. Length about 3-5-4 mm.; breadth in plane of symmetry,
3-3-8 mm. Smaller specimens, length about 2-5 mm.; breadth in plane of symmetry,
2 mm.
NEOTYPE. Valve from a broad fruit. Brit. Mus. (N. H.), No. ¥.33868.
DESCRIPTION. Endocarp : One-loculed, somewhat rugose 'externally, rounded
to sub-ovoid but somewhat angled at the margins, bisymmetric, often but not
invariably compressed at right angles to the plane of symmetry. Dehiscing in
this plane into equal valves showing the sub-urceolate locule on the inner surface
which is not emarginate at the base. Locule narrowing into an apical stylar canal.
At the base the wall is pierced by a short straight vascular canal and is not here
thickened. No clear evidence of the attachment of the seed is yet available.
Sutures wide, flat, close-textured. Endocarp wall compact in structure, formed of
hard parenchyma, cells small, more or less radially aligned on the sutures. Locule
surface smooth (always much corroded), longitudinally striate due to finely toothed
cells which are oblong at the extremities of the locule to which they converge,
equiaxial over the middle. A second coat sometimes overlies the locule-lining
(= testa ?) and shows larger equiaxial cells about 0-05 mm. in diameter. Length
of typical fruit, 3-5-4 mm.; breadth in plane of symmetry, 3-3-8 mm. Small
specimens, length, 2-5 mm.; breadth in plane of symmetry, 2 mm.
92 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
REMARKS. It is impossible to say how many of the variously distorted fruits
should be referred to this species. A laterally crushed example is shown in PI. 12,
fig. 46 which may belong, but distorted specimens of this type are most difficult to
sort and determine.
The presence of Myrica is confirmed by Ross from pollen (p. 75). The Recent
genus has a wide range in the northern Hemisphere. It also occurs in South Africa
and the Andes. It is especially sub-tropical. It is also a widespread Tertiary
genus. The smaller Bovey specimens resemble endocarps found in the London
Clay of Nursling. The species appears to be distinct from the large much-inflated
Myrica supfiani Kirchheimer (1938 ; 1939) from the German Brown Coal (spherical
endocarps 2-3-5 mm- l°ng> locule cordate at the base, carpel wall basally thickened).
Family BETULACEAE
Genus CORYLUS (Tourn.)
Corylus sp.
(PI. 13, figs. 49-5i)
Part of a male inflorescence with anthers (PI. 13, figs. 49, 50) enclosing pollen
examined and determined by Dr. J. B. Simpson. The pollen, which was flattened,
contained no intine ; its condition made it clear that it was a true fossil, not Recent
material accidentally introduced. The grains, like those of Corylus, show the typical
thickening of the exine in the neighbourhood of the pores, and the zone of granules
around each pore, the granules being longer and not so close-set as in other regions
of the exine.
Various modern species of Corylus differ slightly from one another in the size
and prominence of the zone of granules, but Dr. Simpson has not determined the
nearest living species.
Genus CARPINUS Linnaeus
Carpinus boveyanus (Heer) pars
(PI. 13, figs. 52-67)
1862. Carpolithes boveyanus Heer, p. 1077, pi. 70, figs. 7-14 (in part).
DIAGNOSIS. Fruits sub-ovoid, 2-5-4 mm- l°ng. J '75-2*5 mm- broad. Sometimes
twinned.
NEOTYPE. A fruit showing fibres and scar of attachment. Brit. Mus. (H. N.),
No. V.3387O.
DESCRIPTION. Fruit : Attached to a bract rarely preserved and then only as a
fragment at the base. Very variable in shape and size, enclosed by the abraded
accrescent perianth rarely with remains of short superior perianth segments (PI. 13,
fig. 57). On the whole sub-ovoid, bisymmetric and slightly compressed. Scar of
attachment to bract basal, large, sub-circular (PI. 13, figs. 52-54). Surface in the
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 93
better-preserved specimens showing slender longitudinal vascular bundles which
arise from the margin of the basal scar ; the strands themselves are often abraded
but their position may be indicated by furrows (PI. 13, figs. 52, 54, 56). Surface
smooth, formed of small, oblong, longitudinally aligned cells about 0-008 mm.
in diameter.
A few specimens show two fruits grown together. If, as appears likely, they really
belong to Carpinus they may be due to the rare development of two fruits in a bract,
for two female flowers are present in the early stages in living Carpinus. I have
not been able to find any living twinned fruits to bear out this suggestion. Twinned
fossils are shown in PI. 13, figs. 59-65).
Endocarp : One-loculed, agreeing with the fruit in shape, style terminal forming
a mucro when well preserved, showing remains of two style bases. Dehiscence
along a marginal suture in the plane of symmetry. Cavity not well seen.
Seed : When preserved much shrivelled, its placentation obscure.
Length of best preserved fruits, 2-5-4 mm.; breadth, 1-75-2-5 mm. Scar breadth,
0-9-1 mm.
REMARKS AND AFFINITIES. A few fruits among a mass of small, black, crushed
fruits and seeds are attributable beyond doubt to Carpinus. Possibly some of these
were included in Heer's Carpolithes boveyanus which certainly included specimens
of Myrica. The presence of an undoubted accrescent superior perianth, and the
basal scar of attachment to a bract make the relationship to Carpinus clear. It
is unfortunate that there is no evidence so far of the pendulous anatropous seeds
which together with perianth and scar serve to distinguish this species from Myrica.
The same or a closely allied species of Carpinus occurs in the Bournemouth Marine
Beds, at Cliff End near Mudeford, and in the Lower Headon of Hordle.
Family FAGACEAE
Genus FAGUS Linnaeus
Fagus minima n. sp.
(PL 13, figs. 68-70)
DIAGNOSIS. Endocarp unusually small, 3-75-4-5 mm. long ; lateral faces, 2-5
mm., 2-2 mm. and i-i mm. broad respectively.
HOLOTYPE. An endocarp. Brit. Mus. (N. H.), No. ¥.33886.
DESCRIPTION. Endocarp : Trigonous with three unequal flat or concave sides.
Broadest outline semi-oval, style apiculate, base with scar of attachment, i mm. in
diameter, reaching the margin of the narrow face but not continued on to it as in the case
of the other two faces, thus showing it to be one of a pair of nuts which abutted along
the narrow face ; angles sharp but not flanged. Surface smooth, but cell walls
raised so as to form minute, irregular, sinuous, longitudinal crumples 0-008 mm.
apart, also having irregularly rounded dimples 0-012-0-019 mm. in diameter which
are so crowded towards the apex as to produce a rough surface ; they may represent
hair-bases. Length of endocarp, 3-75 mm.; breadth of broadest side, 2-5 mm.;
94 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
maximum breadth of narrowest side, i-i mm. (? originally about 1-75 mm. now
folded in) ; maximum breadth of medium side, 2-2 mm. Length of endocarp
found by C. & E. M. Reid (see below), 4-5 mm.
REMARKS AND AFFINITIES. One endocarp. Two others were figured by C. & E. M.
Reid from Bovey (1910, pi. 16, figs. 67, 68). The angled form, character of attach-
ment scar and surface, position and character of style, all indicate relationship with
Fagus. The small size distinguishes it from any living species seen. No other
fossil species so small has been recognized. The compressed form consequent on
the narrowness of one side, and the correspondingly narrow triangular scar which
terminates at the margin of the narrow side suggest that the endocarp was developed
in a laterally compressed cupule. Fagus leaves are of common occurrence in Cre-
taceous and Tertiary deposits. Leaves of Fagus and of Nothofagus (Bandulska,
1924) occur in the Bournemouth Freshwater Beds.
Family ULMACEAE
Section CELTIDOIDEAE
Genus ZELKOVA Spach.
Zelkova boveyana n. sp.
(PL 13, figs. 71-73)
DIAGNOSIS. Endocarp much inflated. Maximum diameter, 1-5 mm.; maximum
diameter in plane of symmetry, 0-9 mm.; maximum diameter at right angles to
plane of symmetry, 1-25 mm.
HOLOTYPE. An endocarp. Brit. Mus. (N. H.), No. ¥.33887.
DESCRIPTION. Endocarp : Approximately bisymmetric about a plane through
the attachment and style, marked by a conspicuous marginal ridge (PL 13, figs. 72
73) ; asymmetric in this plane of symmetry about a line between the attachment
and style so that the outline is gibbous on one margin near the attachment, and
on the other near the apex (PL 13, fig. 71). The endocarp is much inflated producing
a rounded-quadrilateral outline at right angles to the plane of symmetry (PL 13,
fig. 72). Attachment indicated by a small depression from which obscure ridges
diverge, a few also diverge from the marginal ridge. Surface rough with indefinite
depressions that give rise to an obscure network, wall formed of small equiaxial
cells 0-012 mm. in diameter.
Maximum diameter, 1-5 mm.; maximum diameter in plane of symmetry, 0-9 mm.;
maximum diameter at right angles to plane of symmetry, 1-25 mm.
REMARKS AND AFFINITIES. One endocarp. The form and structure so far as
it has been seen, relate the fruit to Zelkova, a genus represented by about six living
species in North Temperate regions.
All species seen are much larger than the fossil, but they vary much in size, so
that the relationship in size between the fossil and Z. keaki (for example) is comparable
with that between Z. keaki (3 mm. maximum diameter) and Z. sinica (7 mm.). Size
alone could not therefore exclude the fossil from this genus. Z. sinica resembles
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 95
the fossil closely in form, but most living species are more markedly curved, and some
are actually hooked in the stylar region. The apparent absence of superficial
fibres in the fossil may be due to its worn condition. There appear to be no other
grounds for excluding it from the genus Zelkova.
Family MORACEAE
Section MOROIDEAE
Genus MO RO IDE A nov.
DIAGNOSIS. Unidentified genera of the section Moroideae, family Moraceae.
Moroidea boveyana n. sp.
(PI. 13, fig. 74)
DIAGNOSIS. Fruit markedly asymmetric in plane of symmetry. Stylar projection
narrow. Length incomplete ; breadth in plane of symmetry, 1-5 mm.; thickness,
07 mm.
HOLOTYPE. A fruit, broken at the rounded base. Brit. Mus. (N. H.), No.
V.33888.
DESCRIPTION. Fruit : Sub-circular in outline, somewhat flattened laterally
but sub-cuneate in transverse section, the narrower edge crested along the whole
length preserved, the opposite edge and base of the fruit rounded. Style prominent,
terminal at the apex of the crested margin, closely adjacent to a sub-terminal
curved projection which marks the point of entry of the funicle to the sub-apical
placenta inside the rounded margin. Carpel wall 0-05-1 mm. thick, formed of a
few layers of small cells aligned radially so as to give a columnar appearance in section,
producing superficially a closely punctate surface with pits 6-oi2-o-oi6 mm. in
diameter.
Length of fruit, incomplete ; breadth, 1-5 mm.; thickness, 0-7 mm.
REMARKS. One fruit, broken at the rounded base. The interior and structure
of the seed have not been seen. A similar fruit was found at Hordle giving evidence
of both internal structure and of seed structure. In the Hordle specimen the
relation to Moraceae was clear, and the evidence indicated a connexion either with
the section Moroideae or with Artocarpoideae, probably with the former.
The chief distinctions between the Bovey and Hordle fossils lie in the style which
is a narrow projection in the Bovey fruit, and a broad flat one with a broad flat
stylar canal in the fruit from Hordle, and in the greater symmetry of the Hordle
fruit. Such differences are probably of specific value, but this conclusion can
only be established when a greater range of living and fossil material is available
for comparison. In the meantime the Bovey and Hordle fruits are treated as
specifically distinct.
96 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Family NYMPHAEACEAE
Genus BRASENIA Schreber
Brasenia ovula (Brongniart)
(PI. 13, fig. 75)
1862. Nymphaea doris Heer, p. 1072, pi. 70, figs. 32-37.
1925. Brasenia sp. (B. ovulum Brongn. ?) Chandler, p. 23, pi. 3, figs. ja-d.
1926. Brasenia ovula (Brongn.) : Reid & Chandler, p. 99, pi. 6, figs. 15-18.
DESCRIPTION. Seed : Obovoid, now much crumpled and distorted, having
an aperture at one end, measuring 0-45 mm. in diameter, from which the embryotega
has come away. Surface black, glistening, with longitudinal corrugations about
0-075 mm. in breadth at the middle of the seed. Surface cells, 0-05-0-075 mm.
in diameter, with interlocking digitations, the length of individual digitations often
more than one-third of the total diameter of a cell ; surface of cells finely punctate.
Length of seed, 2-25 mm.; breadth, 1-5 mm.
REMARKS. One seed from Heathfield. Heer recorded numerous seeds from
Bovey under the name Nymphaea doris. His seeds were 2-5-3-5 mm. long and 2-3
mm. broad, the diameter being slightly increased no doubt by the flattening of the
the seeds. The species ranges in the British Tertiary from Bournemouth Marine
to Hamstead Beds.
Family MAGNOLIACEAE
Genus MAGNOLIA Linnaeus
Magnolia boveyana n. sp.
(PI. 13, figs. 76-80)
1910. Magnolia attenuata Weber : C. & E. M. Reid, p. 165, pi. 15, figs, i, 2.
DIAGNOSIS. Seeds longer than broad. Length 6-7 mm., breadth 2-75-4-5 mm.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33890.
DESCRIPTION. Seed : Anatropous, ovate in outline, narrow, longer than broad,
much compressed (compression doubtless emphasized by f ossilization) , gently
convex, angled longitudinally on one face with a shallow longitudinal depression
on the other (the raphe side). Chalaza terminal at the broad end, marked by a
small plug or scar which is pierced at the centre. Surface marked by a fine " finger-
print " pattern due to polygonal cells, o-oi mm. in diameter, cells aligned in
rows, the rows being grouped in clusters. Testa (represented only by the inner
hard coat) 0-4 mm. thick at the middle of the seed, formed of equiaxial cells arranged
radially in a columnar manner, the columns about 0-016 mm. broad. Tegmen
thin, translucent, longitudinally striate, structure obscure.
Length of a seed, 6 mm.; breadth, 2-75 mm. Length of a second seed, 6 mm.;
breadth, 4 mm. Length of a seed found by C. & E. M. Reid (1910 : 165), 7 mm.;
breadth 4-5 mm.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 97
REMARKS AND AFFINITIES. Six seeds and several fragments. They resemble
Magnolia seeds of the American longer-than-broad type. The species is larger, more
ovate in outline, and less triangular in transverse section than M. angusta from the
London Clay (Reid & Chandler, 1933 : 177, pi. 5, figs. 6-8).
Comparable living species are M. grandiflora Linn., and M. glauca Linn., but
the former is a large, and the latter is a smaller species. C. & E. M. Reid (1910 : 165)
named these seeds M. attenuata Weber, identifying them with Magnolia seeds found
abundantly in the Rhine lignite ; but they were careful to indicate that as the
type of M. attenuata was a leaf, and as another Magnolia species also occurred in
the same deposit, the connexion of the Rhineland seeds with the leaves was not
conclusively established.
On this account Kirchheimer (1936^ : 85-86) recently instituted a new specific
name M . sinuata for a species from Salzhausen in which he included the relatively
narrow seeds of M. attenuata Weber and a broader-seeded species M. hoffmani
Ludwig on the grounds of histological identity. M. sinuata, although some of its
forms resemble the Heathfield specimens, appears to exhibit much greater variation
both of form and size. On the whole it is appreciably larger, the length of the seed,
6-10 mm.; breadth, 5-9 mm., whereas the maximum length of the Heathfield seeds
is 7 mm.; maximum breadth, 4-5 mm. Hence a distinct specific name, Magnolia
boveyana, has been given to the seeds from the Bovey basin.
Family LAURACEAE
Genus CINNAMOMUM Blume
Cinnamomum is represented at Bovey, according to Heer, by three species, two
based on leaves (C. rossmassleri and C. lanceolatum) , the third on leaves and flowers
(C. scheuchzeri). Probably some of the cupules and berries hereafter recorded may
belong to Cinnamomum. At present, however, they are referred to the family
Lauraceae only, the evidence being insufficient for definite determination.
C. rossmassleri Heer was represented by two leaf-fragments both with the apex
missing. Hence Heer himself regarded the determination as doubtful. The species
is therefore omitted in the list on p. 77 as the figures (Heer, 1862, pi. 67, figs. 17, 18)
are inconclusive and not distinguishable with certainty from those of C. scheuchzeri.
C. scheuchzeri and C. lanceolatum are better represented, and there can be no
reason to doubt the occurrence of the genus Cinnamomum in the Bovey Lake
deposits. No new leaves of either species have been found and the genus is not
therefore included in the plant list on p. 77.
VARIOUS GENERA AND SPECIES
(PI. 14, figs. 81-91)
In addition to the genus Cinnamomum, the Lauraceae are represented by leaves
referred to Laurus primigenia Unger and to Daphnogene ungeri (Heer, 1862 : 1064,
g8 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
pi. 65, figs, i, 2, 6). Neither of these determinations are wholly satisfactory. There
are a number of cupules and berries also. These were never seen by Heer or their
affinities were not recognized.
C. & E. M. Reid (1910, pi. 16, figs. 64-66) figured three small wrinkled cupules
clearly belonging to Lauraceae. Similar specimens (all small) are common both at
Bovey and Heathfield. They vary from about 1*75-3 mm. in diameter. Sometimes
they are cup-like with simple margins, sometimes they are notched or divided
above into sepals. In a few the small unripe berry still lies within the calyx. In
all the skin is much wrinkled, shining, formed of very small cells which may produce
a finely striate effect.
The flattened skins or epicarp of larger, detached, formerly ovoid berries are also
common. The epicarp is leathery, shining, often yellowish-brown and semi-
translucent, mainly formed of equiaxial cells which may vary considerably both in
form and size ; they are often about 0-025 rnm. in diameter.
In some specimens, overlying these cells at the apex is a thin layer of polygonal
cells about 0-05 mm. long and 0-025 rnm. broad. These diverge from the apical
scar, but quickly die out and become obscure so that they cannot be traced a short
distance below the apex. Remains of the mesocarp commonly adhere to the epicarp
and in some specimens enclose numerous ovoid or globular yellow oily (?) bodies
about 0-05 mm. in diameter. The largest berry seen is 6 mm. long (incomplete)
by 5-25 mm. broad (breadth increased by flattening).
It is possible, but unlikely in view of the immature condition, that detailed com-
parative study of cuticle and mesocarp structure in living and fossil material might
serve to distinguish some at least of the genera represented, but it would require
very long research and an abundance of living material for comparison. The
berries and cupules can therefore only be referred to the family Lauraceae without
suggestions as to the generic relationship.
Family CAPPARIDACEAE
Genus CAPPARIDISPERMUM nov.
DIAGNOSIS. A form-genus to embrace seeds of Capparidaceae of which the nearer
relationship is unknown.
Capparidispermum boveyanum n. sp.
(PI. 14, figs. 92-96)
DIAGNOSIS. Seeds transversely oboval about 2-3 mm. in minimum and 2-75-3-75
mm. in maximum diameter. Contiguous walls of the curved limbs form a condyle,
they appear to be fused for most of its length. Testa tubercled, the tubercles oriented
parallel with the margin of the seed. Surface cells equiaxial.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33904.
DESCRIPTION. Seed : Woody, transversely oboval in outline, almost flat,
approximately bisymmetric, splitting for germination in the plane of symmetry.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 99
Locule with curved unequal limbs separated by a narrow curved condyle, the
micropylar limb longer and narrower than the other (PI. 14, fig. 96). The sutures
in the plane of dehiscence are smooth finished surfaces both along the margins of
the seed and on the condyle. Marginal suture variable in breadth, 0-18 mm. broad
at the distal end of the seed ; 0-4 mm. broad near the proximal end on the concave
outer curve of the short limb. Hilar scar large, oval, marginal between the limbs,
sometimes sunk in an emargination (PI. 14, fig. 94). Micropyle small, terminal on
the longer limb. The walls which form the condyle appear to be completely fused
for the greater part of their length except at the hilar end where they enclose a shallow
V-shaped cavity (PL 14, fig. 96), 0-8 mm. long, 0-3 mm. wide at the margin.
The condyle is sometimes indicated externally by an elongate prominence which
is smoother than the rest of the surface (PL 14, fig. 92) . Testa tubercled externally,
the tubercles often elongate parallel with the margin of the seed ; breadth of bases
of tubercles, 0-05-1-5 mm.; height, 0-025-0-05 mm., surface cells equiaxial about
0-025 mm. in diameter giving the margins of the tubercles a clawed or digitate appear-
ance. Testa in section formed of equiaxial cells, 0-017 nun. in diameter. Lining
cells of seed-cavity, 0-017 mm. in diameter, equiaxial, in rows transverse to the length
of the cavity.
Dimensions of four seeds respectively : 2-75 by 2 mm., 3-75 by 3 mm., 3-5 by 2-25
mm., 3-5 by 2-25 mm. (the last represented by one valve only).
REMARKS. Four seeds and two fragments. The curvature of the locule, marginal
hilum between the limbs and the long narrow condyle indicate relationship with
Capparidaceae.
No living genus seen combines the characters of form, size and ornamentation,
hence the reference to a form-genus Capparidispermum. It is possible that the
specimen shown in PL 14, fig. 95, represents a second species, it is larger, flatter,
and the short limb has a more marked spiral incurve. In the absence of more
evidence it is regarded as a slightly abnormal seed of the same species.
Family HAMAMELIDACEAE
Genus ? sp.
(PL 14, figs. 97, 98)
DESCRIPTION. Seed : Sub-oboval in outline, rounded at the apex, slightly
excavated at the base, laterally compressed (compression exagerrated by fossiliza-
tion), with a large, bilobed, slightly concave hilar scar lying across the base having
one lobe on each flat face ; the lobes are unequal in length and breadth but occupy
about half the length of the seed. The arrangement of the other organs has not
been seen. Surface much abraded, rough as preserved, the polygonal cells being
about 0-03 mm. in diameter and somewhat sinuous. In a few places the testa has
cracked transversely on drying owing to the transverse alignment of one or more
of its layers. As seen in section near the apex it is 0-25 mm. thick but its cells cannot
ioo THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
here be distinguished although at one point there is a false appearance of curved
columns, 0-016 mm. broad, the results of the fracture of tissue which has become
vitrified in fossilization. Lining of seed-cavity formed of equiaxial cells o-o 12-0-016
mm. in diameter.
Length of seed, 5-25 mm.; breadth, 3-25 mm.; thickness, 2 mm.
REMARKS AND AFFINITIES. One seed. The form and the large lobed hilar scar
extending over the base and continued on each side, indicate a seed of
Hamamelidaceae. Although many seeds in this family show similar hilar scars,
no living genus seen has so large a scar as the fossil, while in many it is considerably
smaller. In certain living genera there are two distinct scars one on each side near
the base, but not united over it. Sinowilsonia (length of seed, 6-5 mm.; breadth,
4 mm.) has a similar scar occupying almost half the length of the seed. Fortunearia,
with a scar extending about one-third of the length, has a much larger seed as have
many species of Hamamelis. While the relationship of the fossil to the family
Hamamelidaceae is certain, the evidence is insufficient to determine the generic
position, but perhaps the closest living genus is Sinowilsonia.
It is interesting to note that Dr. J. B. Simpson (1936 : 99) records the occurrence
of pollen belonging to Bucklandia, Corylopsis, Fortunearia, Loropetalum, Dicoryphe
and Distylium in the Scottish Tertiary coals of Ardnamurchan and Mull.
Genus ? sp.
(PL 14, fig. 99)
DESCRIPTION. Fruit : Represented only by a fragment of septum and fibrous
axis adhering to the seed near its apex.
Seed : Originally ovoid (much compressed in fossilization) . Hilar scar sunk,
long and narrow, lateral but continued across the base where it terminates without
extending on to the opposite face. Testa black, shining, showing the cell-structure
fairly clearly ; around the scar the cells are elongate, approximately parallel with
its margin, but over most of the surface there are fine parallel striations diverging
obliquely from the middle which appear to be a secondary consequence of
compression.
Length of seed, 4-75 mm. ; maximum breadth, 2-5 mm. Length of scar on lateral
face, 2 mm.; breadth, 0-3 mm.
REMARKS AND AFFINITIES. One seed from Heathfield. The form, hilar scar,
and surface all indicate relationship with Hamamelidaceae. It has not been possible
to relate it to a genus. In Hamamelis the hilar scar is shorter and the seed larger.
In Corylopsis, which has a long, sunk scar, there is also a marked facetting on the
opposite side. F other gilla has a small, sunk scar and the walls of the surface cells
are much thicker.
The second species here described is quite distinct from the first in which the
hilar scar is large, broad and bilobed.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 101
Family ROSACEAE
Genus RUBUS Linnaeus
Rubus microspermus C. & E. M. Reid
(PL 14, figs. 100-109)
1910. Rubus microspermus C. & E. M. Reid, p. 169, pi. 15, figs. 13-17.
DESCRIPTION. Endocarp : Laterally compressed, semi-circular, semi-oval or
sub-ovate in outline, ventral margin straight or almost straight, dorsal margin
semi-circular or markedly convex, base rounded, margin rimmed all round. Surface
reticulate with conspicuous pits, angular in outline, separated by thin, sharp, clearly
defined ridges, occasionally the ridges are prolonged on to the margin. Carpel
wall formed superficially of small equiaxial cells o-oi mm. in diameter.
Length of endocarp, 1-2-5 mm- (commonly 2-25 mm.) ; breadth, 1-1-8 mm.
(commonly 1-25 mm.).
REMARKS AND AFFINITIES. Endocarps of this species are common at Bovey and
Heathfield. Their relationship to Rubus was discussed by C. & E. M. Reid (1910 :
169) but a larger range of material is here shown (PI. 14, figs. 100-109) to demonstrate
variation in size and form. Prickles of Rubus which were reasonably assumed to
belong to the same plant were also described and figured by C. & E. M. Reid (1910 :
169, pi. 15, figs. 16, 17). The endocarps have been compared with Rubus acutiformis
Chandler which occurs at Hordle, Cliff End, Sandbanks, Branksome Dene and Stud-
land, but the two species are clearly distinguishable although both are of unusually
small size. Rubus acutiformis is commonly more pointed and narrower at the
apex, and is therefore more ovate in outline as a rule, while the ventral margin is
not infrequently very slightly concave, the apex being curved somewhat towards
the ventral side. The well-marked marginal flange is most conspicuously developed
on the ventral margin.
>
Family LEGUMINOSAE
Genus ?
(PL 15, figs. 110-112)
DESCRIPTION. One perfect compressed seed, and fragments of three others must
be referred to Leguminosae. The original shape was probably lensiform (but the
seeds are now flat owing to compression), the marginal hilar scar (obscured by margi-
nal cracking) must have been small. The surface is formed of inconspicuous concave
equiaxial cells 0-02 mm. in diameter, the walls are 0-25 mm. thick as seen in one of
the incomplete specimens, 0-45 mm. thick around the hilar aperture, the cells being
arranged in a columnar manner, the radial columns about 0-012 mm. broad ; a
shallow pocket, presumably connected with the radicle, lies immediately beneath
the hilum, it is delimited on the surface of the cavity by elongate cells. Lining
of main seed-cavity of convex equiaxial cells 0-012 mm. in diameter. Diameter
of the perfect seed, 3-5-3-75 mm.; other specimens larger but incomplete.
102 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Family RUTACEAE
Section ZANTHOXYLEAE
Genus RUTASPERMUM nov.
DIAGNOSIS. A form-genus to include seeds of Rutaceae of which the nearer
relationship is unknown.
Rutaspermum exaratum (Heer)
1862. Carpolithes exaratus Heer, p. 1079, pi. 70, figs. 24-27.
The seed was described by Heer as 3-5 mm. long, 3-3 mm. broad ; the figures
showed it as sub-circular in outline, much inflated, with a long, narrowly-triangular,
sunk hilar scar (the " umbilical fissure " of Heer). The surface was ornamented
with nodular ridges aligned parallel with the rounded dorsal margin, much as in a
species to be described from Sandbanks, and ornamented also with fine polygonal
cells or pits (the " innumerable dots " of Heer) ; but the species is larger than
the Sandbanks seeds. The characters are clearly those of Zanthoxyleae, of the type
referred to the form-genus Rutaspermum. In the absence of actual specimens
and of more accurate figures than the diagrammatic illustrations given by Heer,
the species cannot be clearly defined. Available evidence, however, suggests that it
is of a very distinctive type.
Family SABIACEAE
Genus MELIOSMA Blume
Meliosma reticulata (C. & E. M. Reid)
(PI. 15, figs. 113-118)
1910. Calvarinus reticulatus C. & E. M. Reid, p. 169, pi. 15, figs. 18-20.
DIAGNOSIS. Endocarp about 5-5-25 mm. long, 3-75-4 mm. in maximum trans-
verse diameter. External surface with about eighteen to twenty raised ribs over
the proximal half which branch and anastomose to form a sharp network over the
distal half.
NEOTYPE. A perfect but laterally compressed endocarp. Brit. Mus. (N. H.),
No. ^33924.
DESCRIPTION. Endocarp : Woody, obovoid, slightly compressed laterally (com-
pression increased by fossilization), the transverse diameters being in the proportion
of 7 : 10 ; bisymmetric about a plane which passes through the attachment, funicular
canal and a marked marginal angle ; splitting in the plane of symmetry into two
valves. Funicular canal oblique, about I mm. long, placenta basi-lateral or sub-
basal. External surface ornamented with about eighteen to twenty raised ribs
diverging from the attachment over the lower half of the fruit and uniting, branching,
and anastomosing, to form a network over the upper half of the fruit. Surface
formed of polygonal cells o-oi mm. in diameter. Walls i-25-i'5 mm. thick. Surface
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 103
of locule rough, cell-structure very obscure, but the centipede-type of cells with
interlocking walls characteristic of Meliosma can be traced oriented parallel with
the lateral ribs on one small fragment ; the length of individual cells is obscure,
but near the base their width is about 0-037 nim.
Length of endocarp, 5-25 mm.; breadth, 4-25 mm. Length of a second endocarp,
5 mm.; breadth, 3-75 mm.
REMARKS AND AFFINITIES. Six endocarps or valves and a number of fragments.
The form, surface ornamentation, structure, and short oblique funicular canal
relate these fossils to Meliosma. So far as it has been possible to study the different
living species, the Bovey fossils are unique in the number of their longitudinal
ribs and the fineness and sharpness of the apical network, also in the narrow oboval
form. In living species the three diameters (two transverse and one longitudinal)
are usually approximately equal but occasionally the transverse diameters may be
slightly longer or shorter than the longitudinal. The considerable differences
in the diameters of the fossil which give rise to the elongate form and lateral com-
pression have not, however, been seen in the living forms.
C. & E. M. Reid (1910) described the species under the generic name Calvarinus,
and referred it to the family Boraginaceae. There were two specimens from Heath-
field and one from Bovey. Reid and Chandler also obtained endocarps from both
localities.
Family VITACEAE
There are several types of vine seeds at Bovey and Heathfield. Heer (1862 : 1070,
pi. 69, figs. 25-29) distinguished two species which he named Vitis britannica and
V. hookeri. The figures are poor and the descriptions inadequate but the outstanding
features are clear.
C. & E. M. Reid (1910 : 165, pi. 15, figs. 3-6) distinguished three species, V. hookeri
Heer, V. teutonica A. Br. and V. ludwigi A. Br. The greater number of specimens
are now referred to Parthenocissus britannica (Heer). One imperfect seed appears
to belong to V. hookeri Heer. The specific relationship is discussed in the following
pages where the species are described.
Two new species have been added, Parthenocissus boveyana from Bovey, and
Vitis stipitata from Heathfield.
Genus PARTHENOCISSUS Planchon
Parthenocissus britannica (Heer)
(PL 15, figs. 119-122)
1862. Vitis britannica Heer, p. 1071, pi. 69, figs. 25, 26.
1910. Vitis ludwigi A. Br. : C. & E. M. Reid, p. 166, pi. 15, fig. 6 (not fig. 4 as in text).
1910. Vitis teutonica A. Br. : C. & E. M. Reid, p. 166, pi. 15, figs. 4, 5 (not fig. 6 as in text).
DIAGNOSIS. Seed pointed-obovate in outline, smooth dorsally, slightly emarginate
at the apex, chalaza elongate-ovate, surface grooved between chalaza and base ;
sharply angled ventrally with infolds occupying more than half the length, deep,
GEOL. Ill, 3. 9
104 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
narrow, straight but diverging upwards. Length, 4-75 mm.; breadth, 2-25-2-8
mm.; thickness, 1-5-2 mm.
NEOTYPE. A typical seed. Brit. Mus. (N. H.), No. V.^g2y.
DESCRIPTION. Seed : Pointed-obovate in outline, slightly emarginate at the
apex, pointed at the base, with smooth contours, ventral and dorsal faces meeting
at an acute angle, ventral face sharply facetted so as to form a conspicuous
raphe ridge extending almost the whole length of the seed, the facets being flat
or concave and meeting approximately at a right angle ; at the apex the ridge gives
place to the shallow channel which produces the slight apical emargination. Ventral
infolds deep, narrow, straight, occupying more than half the length of the seed
arising near the base, diverging upwards so that the raphe ridge is broader above
than below. Dorsal face flat, slightly emarginate at the apex having a shallow
groove along which the raphe passes into the narrow elongate-ovate chalaza which
lies above the middle of the seed. From the chalaza a well-marked groove passes
to the base. Testa formed of two coats, the outer thin, its irregular elongate
polygonal cells about 0-02 mm. in shortest diameter producing transverse striations
which diverge from the chalaza and ventral infolds to the margin. The inner coat
is hard, averaging o-i mm. in thickness, formed of cells 0-016 mm. in diameter
arranged in radial columns. These cells give a finely pitted surface to the coat.
Length of seed, 4-75 mm.; breadth, 2-25-2-8 mm.; thickness, 1-5-2 mm. (somewhat
distorted).
REMARKS AND AFFINITIES. Ten seeds and several fragments from Bovey. The
smooth seed with long divergent infolds suggests relationship with the living Partheno-
cissus. The same species was apparently described by C. & E. M. Reid (1910)
under the name Vitis teutonica A. Br. They state that " Heer's type specimens
of V. brilannica appear to be nothing but badly compressed seeds of this vine
[V. teutonica}, though his figures are scarcely recognisable". Their description is
" ovate-acuminate, gradually narrowed into the beak, granulate all over, inner
face with long shallow pits, outer convex longitudinally sulcate with a narrow
pyriform chalaza, length 4 mm."
The identity of these specimens with Vitis teutonica A. Br. from the German
lignite cannot now be maintained. V. teutonica was originally based on leaves,
but as seeds were associated with them, Unger applied the name to the seeds also
(Braun, 1845 : 172 ; Unger, 1860 : 23, pi. 9, figs. 1-8). Unfortunately neither
Unger's figures or diagnosis really define the species. Later German workers have
also referred a variety of seeds from the Brown Coal to V. teutonica. Thus Krausel
(1920, pi. 25, figs, i, 2) illustrates seeds with short, wide, divergent lateral infolds,
a marked apical groove on the ventral side, a short elongate-obovate chalaza
and shallow furrow between the chalaza and base on the dorsal side (pi. 24,
figs. 20-23). His seeds appear more inflated than the Bovey specimens, and are
sometimes fluted. Kirchheimer (1934 : 35, pi. 9, figs. 3-6) describes and figures
a somewhat fluted seed under the name V. teutonica. It has a relatively small
oval chalaza and appears quite distinct from the Bovey seeds. Later Kirchheimer
(1938, pi. 4, figs. 12-15 i I939> pl- 2, fig. 3) figures other seeds under this name.
The first shows a small median oval chalaza and very wide sub-parallel ventral
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 105
infolds. The second shows a much larger chalaza which is elongate-oval in shape
occupying the upper half of the seed, while wide and divergent ventral infolds
and a markedly stipitate base arise out of the rounded, smooth, lower half of the
seed.
The Bovey fossil appears to be sufficiently distinctive in its shape and chalazal
character and in the narrow upwardly divergent infolds to merit a distinct specific
name and Parthenocissus britannica (Heer) is here retained for this purpose.
Another Bovey seed described and figured by C. & E. M. Reid (1910 : 166, pi. 16,
fig. 6) as V. ludwigi ? is also probably a differently distorted specimen of P. britannica,
although in the description it is said to have a more slender form with an oval, not
pyriform, chalaza. Such individual differences may, however, occur among living
seeds within a species.
Parthenocissus boveyana n. sp.
(PI. 15, figs. 123-125)
DIAGNOSIS. Seed obovate in outline, not emarginate at the apex, contours
smooth, chalaza median oval, angle of raphe ridge about 135°, ventral infolds
markedly divergent upwards. Length, 3-5 mm.; breadth, 2-75 mm.; thickness,
i mm.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33929.
DESCRIPTION. Seed : Obovate in outline, not emarginate at the apex, pointed
at the base, contours smooth, ventral face facetted, the facets being slightly concave
and meeting at an angle of about 135° so that the raphe-ridge is not very sharp
or conspicuous ; the raphe itself — a stout cord — is preserved above the middle
on the ventral face and is continued on to the dorsal face passing into
the external chalaza. Ventral infolds deep and narrow, extending from near the
base to a quarter of the length from the apex, they diverge above and are slightly
convex towards the raphe-ridge which is therefore triangular. Dorsal face almost
flat, very slightly convex, not emarginate at the apex, or if it is grooved, the groove
is obscured by the preservation of the raphe ; there is a shallow median groove
between the chalaza and the base. Chalaza oval situated rather above the middle
of the seed, gradually narrowing above into the raphe. Surface rather rough
especially at the base and apex and on the chalaza, cells diverging from the lateral
infolds and from the chalaza, many of them elongate in the direction of divergence,
others polygonal 0-012 mm. in diameter. Wall as seen in section columnar, the
columns about 0-012 mm. in diameter. Thickness of wall, o-i mm.
Length of seed, 3-5 mm.; breadth, 2-75 mm.; maximum thickness, i mm.
REMARKS AND AFFINITIES. One seed, and an imperfect specimen possibly
referable to this species. The seed is relatively broader than seeds of V. britannica,
and has a larger, broader chalaza, and more divergent lateral infolds. It is less
rounded and stipitate than Kirchheimer's figures of V. teutonica (1939, pi. 2, figs.
ja-e) and lacks the emarginate apex. Its chalaza is much larger than that in
Kirchheimer's figure (1938, pi. 4, fig. 14).
io6 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Genus VITIS Linnaeus
Vitis hookeri Heer
(PI. 15, figs. 126, 127)
1862. Vitis hookeri Heer, p. 1070, pi. 69, figs. 27-29.
1910. Vitis hookeri Heer : C. & E. M. Reid, p. 165, pi. 15, fig. 3.
DESCRIPTION. Seed : Broadly obovate in outline but scarcely emarginate
at the apex, pointed at the base, contours smooth and rounded. Ventral face
facetted so as to form a conspicuous raphe-ridge, ventral infolds broad widening
upwards, about half as long as the seed. Dorsal face rounded with slight flutings
diverging from the ovate chalaza which is situated above the middle of the seed.
Between chalaza and base is a deep median groove. Testa, 0-075 mm. thick,
formed of cells 0-0125 mm. in diameter which have a columnar radial arrangement ;
they give rise superficially to a finely and evenly but deeply punctate surface.
Length of seed, 3-75 mm.; estimated breadth (actually incomplete), 3-5 mm.
REMARKS AND AFFINITIES. One incomplete seed from Heathfield ; also a second
from Bovey which may possibly be referred to this species. This second specimen
is much distorted being compressed from base to apex, but its features can be seen
and measured, and agree with those described above. It is slightly stipitate.
Except that the better preserved specimen is not stipitate, it shows characters
which agree with those of Vitis hookeri Heer of which Heer found one seed at Bovey.
There is general agreement in size with Heer's specimen (length, 3-5 mm; breadth,
3 mm.), the contours are in general similar to those in Heer's inadequate figure;
the chalaza is of comparable size and, although not round, occupies a very similar
position on the dorsal surface when due allowance is made for its displacement
by distortion which has brought it nearer to the apex than it originally lay. In
the second much distorted specimen, the chalaza is round and but slightly above
the middle. Comparison of the ventral face cannot be satisfactorily made both
because of the distortion of the new material and because Heer's figure cannot really
represent the ventral aspect of any vine. While, therefore, identity with Heer's
species V. hookeri is not indisputably established, it seems probable that the new
specimens should be referred to that species. The presence or absence of stipitation
is not in itself of great importance, for Recent grape-seeds show that this character
varies considerably in a single species. C. &. E. M. Reid (1910 : 165, pi. 15, fig. 3)
described as V. hookeri Heer a seed which is probably correctly so named, although
its chalaza is somewhat smaller than that shown in Heer's type. But variation in
size of the chalaza also occurs in living species. The ventral side of this seed was
not shown.
Family TILIACEAE
Genus TILIA Linnaeus
Tilia sp.
(PI. 15, figs. 130-132)
Two groups of anthers were obtained from Heathfield and were sent to Dr. J. B.
Simpson for examination. In a letter (27.^.38) he reported that they yielded
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 107
typical pollen of Tilia, adding " I am not well acquainted with the pollen of many
of the tropical genera of Tiliaceae, and so cannot positively exclude the other genera
except Grewia and Luhea, but certainly they [the pollen grains] agree so perfectly
with Tilia, that I feel sure it must be this genus they represent ". A further report
after a greater variety of living material had been obtained and examined was
expected but has not come to hand. But in a letter dated i.iv.38 Dr. Simpson
adds in writing of the fossil pollen " The furrows are short and deep and have the
appearance of pits and occur in the middle of the sides not at the rounded corners
of the triangular grain. The exine ... is irregularly reticulate with a fine
mesh ".
Ross, working quite independently of Simpson recorded " cf. Tilia " based
on pollen.
Family LYTHRACEAE
Genus MICRODIPTERA nov.
DIAGNOSIS. Fruit many-seeded. Seeds anatropous, compressed at right angles
to the germination valve and in the plane of symmetry. Valve an oval operculum
as in Diclidocarya menzeli E. M. Reid in the lower part of the dorsal surface. Seed-
body flanked by two thin lateral wings with spongy internal tissue. Wings more
or less equally developed.
TYPE SPECIES. Microdiptera major n. sp. from the Eocene of Sandbanks (awaiting
full description). Brit. Mus. (N. H.), No. ¥.34249.
Microdiptera parva n. sp.
(PI. 15, figs. 133-149 '. Text-fig. 2)
DIAGNOSIS. Lateral wings very thin, markedly concave on the ventral side,
raphe straight and narrow, triangular form of seed rare. Maximum length of seed
so far recorded, 1-25 mm.; maximum breadth, 1-75 mm. (2 mm. in Cliff End
specimen) .
HOLOTYPE. A seed. Brit. Mus. (N. H.), No. ¥.33934.
DESCRIPTION. Seed : Anatropous, much compressed dorsi-ventrally ; oboval,
sub-circular, transversely oval, irregular in outline, or occasionally broadly triangular,
differentiated into a median elongate-oval body and thin lateral wings, convex on
the dorsal surface, concave on the ventral. Raphe prominent, linear, median
longitudinal on the ventral face flanked by the marked concavities of the wings,
hilum marginal. Germination by an oval operculum on the lower half of the dorsal
face, associated with the basal micropyle. Internal chalaza circular at the apex
of the seed (Text-fig. 2). Testa formed superficially of a shining, black coat giving
a netted surface with large cells or pits equiaxial at the apex, more irregular and
elongate below. Over the operculum the cells are very regular, equiaxial, about
0-05 mm. in diameter, arranged in about seven regular longitudinal rows. Some
specimens show in addition evidence of small cells, 0-01-0-012 mm. in diameter,
also aligned in longitudinal rows. Within the superficial coat and clearly seen on
io8 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
abraded seeds is a layer of fine cells 0-012 mm. in diameter ; they are arranged so as
to give rise to striations which curve around the operculum, across on to the wings,
and lie parallel with the margin of the wings near the circumference of the seed.
Near the margin, the wings are only 0-025 mm. thick, but they thicken towards the
seed-body close to which they measure 0-06 mm. in thickness. The external coats are
close-textured and no cells can be distinguished in section ; near the body the dorsal
coat is 0-025 mm- thick, the ventral 0-012 mm., the middle layer is 0-025 mm- thick
and is much less compact than the outer layers, but its cell-structure cannot be
clearly seen. The coat surrounding the seed-cavity is hard and compact, it is
0-05 mm. thick on the dorsal side, 0-037 mm. thick on the ventral side. The seed-
cavity is finely striate longitudinally. The tegmen is semi-translucent and is fused
with the testa at the large black chalazal scar.
FIG. 2. Microdiptera parva n. gen. et sp. Diagrammatic longitudinal
section through seed-cavity, x 45 approx.
The dimensions of a series of seeds are as follows :
Length Breadth
(mm.) (mm.)
(1) • 1-25 - i-75
(2) . i-o . 1-25
(3) • i-o . 1-5
(4) i-o i-75
(5) • i-25 • i-o
Typical dimensions of germination opercula are respectively 0-5 by 0-3 mm.,
0-5 by 0-4 mm., 0-6 by 0-4 mm., 0-5 by 0-4 mm., 0-55 by 0-35 mm., 0-4 by 0-3 mm.
REMARKS AND AFFINITIES. The characters are those of Lythraceae, more especially
the position of the organs, the form of the anatropous seed and the structure and
arrangement of the wings. The systematic position will be discussed in greater
detail when the type species from Sandbanks is described in a forthcoming catalogue.
At Bournemouth not only isolated seeds, but some still in the position of growth
in a fragment of fruit were found.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 109
Family NYSSACEAE
Genus NYSSA Gronov.
The genus Nyssa from Bovey is in urgent need of revision. Four supposed species,
truncate at one end, must now be referred to Symplocos. They are Nyssa europaea
Unger (Heer, 1862 : 1066, pi. 69, figs. 11-17 '> C. & E. M. Reid, 1910 : 67, pi. 15,
fig. 9), Nyssa laevigata Heer (1862 : 1066, pi. 69, fig. 18), Nyssa microsperma Heer
(1862 : 1067, pi. 69, fig. 24 ; C. & E. M. Reid, 1910 : 167) and Nyssa obovata Weber
(C. & E. M. Reid, 1910 : 168, pi. 15, fig. 10).
One specimen figured as Nyssa ornithobroma Unger by C. & E. M. Reid (1910 : 168,
pi. 15, fig. n) does not agree with Unger's species and may be a Mastixia.
Others are of uncertain relationship : N. striolata (Heer, 1862 : 1067, pi. 69,
figs. 20-23) • The figures show an ovoid ribbed fruit with finer, closer, more regular
ribbing than that of N. boveyana (now to be described). Heer's fig. 23 shows a mass
of fruits lacking both the finer ribbing seen in his figs. 20-22, and the coarser ribbing
of N. boveyana. The types in the Geological Survey Museum were so decayed in
1910 that the true nature of this endocarp could not be discovered, all finer features
being obliterated.
There remains in these earlier records Nyssa vertumni Unger (C. & E. M. Reid,
1910 : 168, pi. 15, fig. 12) which is undoubtedly Nyssa.
There is indeed abundant evidence of the genus which occurs both at Bovey
and Heathfield but the evidence is of one species only, now described as N. boveyana
n. sp.
Nyssa boveyana n. sp.
(PI. 16, figs. 150-157)
DIAGNOSIS. Endocarp sub-ovoid or less commonly sub-obovoid, tending to be
arched, the dorsal side being less convex than the ventral, ventricose in the upper
part of the ventral face. Germination valve triangular, relatively narrow in propor-
tion to its length, 2-5-4-5 mm.; breadth, 2-5-3-25 mm. Length of endocarp, 6-8-11-5
mm.; breadth, 3-1-5-5 mm.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33944.
DESCRIPTION. Endocarp : Sub-ovoid or less commonly sub-obovoid with a tendency
to be less convex on the dorsal than on the ventral side, frequently ventricose in the
upper part of the ventral surface, inflated and rather narrowed towards the base
(but now usually flattened and often distorted in f ossilization) . The surface shows
conspicuous, broad, rounded, longitudinal ribs and thin strands of fibres in the
intervening narrow furrows ; the ribs, six to twelve in number, extend from base to
apex on the ventral and lateral faces, are in general smoothly rounded but have a
tendency to become nodular especially over the ventricosity, they are symmetrically
no THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
placed, four or five being ventral and two lateral ; along the crests of the lateral pair
at the top of the endocarp are the sutures of the dorsal germination valve. Valve
sub-triangular, gaping at the apex and along the sides which are slightly incurved
towards the base, breaking irregularly along its base which is also that of the triangle.
On the dorsal surface there is a thin thread-like median rib, sometimes flanked
on each side near the base by a broad scarcely raised rib, these die out before reaching
the valve.
Length of endocarp, 6-8-11-5 mm.; breadth, 3-1-5-5 mm.
The dimensions of a series of endocarps, in millimetres, are as follows : 9-5 x 4-6,
10 x 4-6, 10 x 4, 9 x 5-1, 9 x 6, ii x 3-1, 9 x 4-2, n x 4-2,7-2 x 4-2, 9-8 x 4-5,
8 x 3-8, 9-8 x 5'5, 9x4, 10-5 x 4-8, 9 x 4-6, 7-4 x 4-9, 8x4, 7-2 x 4-1, 8 x 37.
9-5 x 5, n-5 x 5, 6-8 x 8-9, 8-4 x 4.
REMARKS AND AFFINITIES. One of the most abundant fossils in the Bovey
lignite basin occurring in profusion in the upper beds at Heathfield and in the lower
beds at Bovey (C. & E. M. Reid, 1910 : 167). The structure of the endocarp and the
characteristic short dorsal germination valve place them beyond doubt in Nyssa.
Although now almost invariably flattened they were obviously much inflated in
life. The many directions of distortion and compression show that originally they
were but slightly compressed dorsi-ventrally below the valve. The valve itself
was flat or slightly concave while there was considerable ventricosity in the
corresponding part of the ventral face. Some specimens have been folded upon
themselves from top to bottom, the folding always being towards the dorsal side.
A few have been flattened laterally ; these show the difference in curvature of the
two surfaces described. The position of crushing has affected the appearance
of the ridges which may be almost obliterated or folded longitudinally so as to appear
as sharp ridges.
The specific determination of fossil Nyssa presents great difficulties as was
recognized by Krausel (1920) and later, in a series of papers by Kirchheimer.
Initially Kirchheimer (1934) regarded the abundant Brown Coal species described
by Krausel (1920), Gothan & Sapper (1933) and himself as agreeing both in
morphology and histology with the Recent Nyssa sylvatica Marsh and indistinguish-
able from it. In later reviews of the subject (1938, 1939) he included all the Brown
Coal " species " (including Middle Oligocene to Pliocene forms) in a single " form-
species ", Nyssa disseminata (Ludwig), on the grounds that the characters did not
permit of true specific determination. The length of N. disseminata was given as
0-8-1-9 cm.; breadth as 0-5-1-2 cm. (1938 : 339) ; length, 1-2 cm.; breadth, 0-5-1-2
cm. (19390 : 270).
It has not been possible to make a study from actual Brown Coal material and
it is abundantly clear that at present species cannot be distinguished on histological
grounds. It may, however, be possible to separate some species at least by size
and shape of the endocarp plus shape of the germination valve. Thus the Pliocene
Reuverian endocarps recorded as N. sylvatica by C. & E. M. Reid (1915) are dis-
tinguishable on these grounds from the Bovey species although apparently agreeing
closely with the living N. sylvatica.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
N. sylvatica
Number of speci-
mens used
N. sylvatica
(Recent)
40
N. sylvatica
var. biflora
(Recent)
(Pliocene ;
(Reuver)
49
Length of endo- 6-9-8 mm. . 7-5-10-5 mm. . 5-5-11 mm.
carp
Breadth of endo- 4-5-6 mm. . 5-2-7 mm. . 3-5-6 mm.
carp
Length of valve . 3-3-5 mm. . 3-5-3-6 mm. . 2-25-3-25 mm.
(5 specimens (2 specimens (8 specimens)
only) only)
Breadth of valve . 4-5 mm. . 4-5-5 mm. . 3-4 mm.
N. disseminata
(Brown Coal) N. boveyana
Kirchheimer's . 42
measurements
(1938 ; I939«)
8-20 mm.
5-12 mm.
6-8-11-5 mm.
3-1-5-5 mm.
2-5-4-5 nun.
(13 specimens)
2-5-3-25 mm.
The valve measured in Kirchheimer's figure of a Brown Coal Nyssa (Kirchheimer,
1938, pi. 4, fig. 24) appears to be : length about 4-5 mm.; breadth, 3-5 or 4 mm.
It will be noted that the Bovey endocarps are relatively longer, and narrower
than those of N. sylvatica while the valves have a relatively narrow triangular
form. These characters can be seen in the published figures.
It is possible, basing the suggestion on experience of the wide range shown by
many other Eocene and Oligocene genera and species, that the Brown Coal species
of Nyssa, at least in the older beds, may indeed be a true single species. Whether
it should in that case be referred to N. disseminata Ludwig (1857) or to N. rugosa
Weber (1852) is not within the scope of this paper. It does, however, raise the
question of the relationship of the Bovey Nyssa to the Brown Coal forms, for it
might be reasonably supposed that the same wide-ranging species occurred in
Germany and Britain.
The figures already quoted, together with an examination of the . published
illustrations of N. disseminata (or N. rugosa) suggest that the Bovey Nyssa should
provisionally be regarded as distinct. The germination valve in the two species is
of similar narrow triangular character differing markedly from tljat of N. sylvatica
from any source. On the other hand the Bovey Nyssa is appreciably smaller on
the whole than the Brown Coal endocarps judging by Kirchheimer's measurements
quoted above. Moreover if his figures (1938, pi. 4, figs. 21-24) are typical they show
that N. disseminata tends to produce endocarps which are normally broadest above
the middle : those from Bovey show a majority which are broadest at the middle.
In view of these features a distinct name, Nyssa boveyana, has been given to the
British material. But it obviously more closely resembles N. disseminata than
N. sylvatica whether Recent or Pliocene.
Family MYRTACEAE
Section MYRTINAE
Genus MYRTOSPERMUM nov.
DIAGNOSIS. Seeds referable to the family Myrtaceae and probably to the section
Myrtinae, with curved or U-shaped cavities, marginal hilum at the end of a condyle
H2 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
between the limbs of the curved cavity. Chalaza close to the hilum, terminal
or sub-terminal on the inner side of one limb, micropyle adjacent to the hilum,
terminal on the other limb.
TYPE SPECIES. Myrtospermum variabile n. sp. Bournemouth Freshwater
Beds (awaiting description). Brit. Mus. (N. H.), No. ¥.34248.
Myrtospermum boveyanum n. sp.
(PL 16, figs. 160-168)
1910. " compressed winged seed " C. & E. M. Reid, p. 173, pi. 16, figs. 61, 62.
DIAGNOSIS. Seed of variable shape, much compressed, glossy, with thin testa.
External surface ornamented with very regular concentric rows of cells 0-05-0-1 mm.
in diameter (pits or convexities). Diameters of seeds, 1-25-2 mm.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33950.
DESCRIPTION. Seed : Bisymmetric, flat (the degree of compression emphasized
by fossilization, sometimes distorted and folded on itself) , sub-oval, transversely-oval,
sub-circular, or rounded-triangular, sometimes slightly emarginate at the hilum ;
limbs of the U-shaped cavity somewhat unequal in length and breadth, the micro-
pylar limb being the longer and narrower. Hilar-scar elongate-oval, marginal
between the limbs, micropyle small terminal on one limb, chalaza small sub-terminal
on the other. Surface shining, ornamented externally with polygonal or hexagonal
pits 0-05-0-1 mm. in diameter, but a few specimens give clear evidence that the pits
are actually highly convex areas which have now collapsed ; the pits are aligned
parallel with the margin of the seed except near the middle where they diverge
from the narrow median area between the limbs. In this median area individual
pits are usually very obscure but when visible they are longer and narrower in this
part of the testa than over the rest of the surface. In certain specimens the outlines
of the surface pits are confused possibly owing to the presence of fine parenchymatous
cells which form their walls as in other species of Myrtospermum. Testa only about
0-075 mm. thick in section. The outer part is formed by the coarsely pitted coat,
the inner part shows evidence of equiaxial or rectangular cells 0-012-0-016 mm.
in diameter ; its structure is often obscure owing to intense compression. Diameter
of seeds, 1-25-2 mm.
REMARKS AND AFFINITIES. Numerous seeds, all much crushed so that they are
reluctant to split in the plane of symmetry. Fortunately one imperfect seed had
split naturally and shows the U-shaped cavity and marginal germination. The
form of the cavity was confirmed in other specimens by treatment with nitric acid,
potassium chlorate and ammonia which rendered them semi-translucent. The
curved form and unequal limbs are also clearly indicated by the alignment of the
surface sculpture. The species occurs both at Bovey and Heathfield.
The relationship of the fossil to Myrtaceae is fully discussed in a forthcoming
catalogue on the Bournemouth flora.
The species here described as M. boveyanum differs from others from Bovey,
Heathfield, the Bournemouth Beds and elsewhere in its extreme degree of compression
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 113
which must be, in part at least, original. It differs also in its thin walls, glossy
surface and in the extreme regularity of its surface sculpture. It is also larger than
the common Tertiary species awaiting description in a forthcoming catalogue
as Myrtospermum variabile. Two specimens were figured and described by C. & E. M.
Reid (1910 : 173, pi. 16, figs. 61, 62) but not named.
Myrtospermum dubium n. sp.
(PI. 16, figs. 169-172)
DIAGNOSIS. Seed sub-circular or irregular in outline. Surface pits 0-025-0-05
mm. in diameter near the circumference, smaller near the hilum and between the
limbs. Diameter of seed, 1-25-2 mm.
HOLOTYPE. Brit. Mus. (N. H.), No. V.33957.
DESCRIPTION. Seed : Sub-circular or irregular in outline, somewhat inflated,
approximately bisymmetric but with a tendency to be more convex on one side than
on the other. Two specimens have a slight elevation over the condylar area between
the limbs of the U-shaped seed-cavity ; the exact form of the cavity is not exposed.
Marginal hilum large, oval, micropyle small. Surface pitted, pits polygonal or
hexagonal, about 0-025-0-05 mm. in diameter near the circumference, smaller and
more obscure towards the hilum and between the limbs, tending to be arranged in
rows parallel with the margin and to diverge from the area between the limbs.
Tegmen thin and translucent, cells not seen.
Diameter of four seeds respectively, 1-6 mm., 1-6 by 1-25 mm., 2 mm., 1-75 by
1-25 mm.
REMARKS. Five seeds all much compressed and carbonized. One was fractured
transversely whereupon it showed the two limbs of the cavity in transverse section;
owing to the mode of preservation, the structure of the wall is obscure. The
diameter is twice as great as that of typical seeds of M. variabile.
Myrtospermum sp.
(PI. 16, figs. 173, 174)
DESCRIPTION. Seed : Bisymmetric, broadly sub-oval in outline, slightly truncate
at the hilar end, inflated, but having a somewhat depressed median area ; U-shaped
cavity with a maximum diameter of 0-35 mm.; micropylar limb longer and narrower
than the chalazal limb, micropyle terminal at the end of the longer limb, chalaza
sub-terminal on the other. Surface deeply pitted, pits equiaxial and hexagonal
near the margin where they are 0-07 mm. in diameter, becoming irregular in form
and size away from the margin, narrow and elongate over the condylar area between
the limbs where they may measure o-i by 0-03 mm. Their walls appear not to be built
of small cells. They form thick ridges with a median groove along which there is
a marked tendency for splitting to occur. Testa thick, maximum thickness at the
end opposite to the hilum (0-12 mm.). The outer part of the testa as seen in section
is a single coat of large simple prismatic cells with their longest axes (0-09 mm.)
n4 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
at right angles to the surface. The collapsed outer ends of these cells form the surface
pits. Inner part of testa formed of uniform parenchyma, the cells being 0-025
mm. in diameter and radially arranged.
Diameter of seed, i-i by 0-9 mm.
REMARKS. One seed (¥.33960) , now broken at the hilar end and split marginally
(irregularly) so as to expose the internal structure.
It differs in its small size from M. boveyanum and M. dubium. From
M. boveyanum it also differs in its inflated form and less regular pitting. From
M, dubium it also differs in its surface sculpture.
The single layer of prismatic cells forming the outer coat of the testa is
characteristic. There is no indication, as in M. variabile from Cliff End, Sandbanks,
Woolwich and Reading Beds, that it was formed of fine parenchyma. The middle
lamella between the prismatic cells is clearly indicated by the narrow grooves along
the middle of the ridges between the pits.
Family CORNACEAE
Section MASTIXIODEAE
Genus MASTIXIA Blume
Mastixia boveyana n. sp.
(PI. 17, figs. 175-178)
1910. Mastixia n. sp. C. & E. M.Reid, p. 166, pi. 16, figs. 73, 74.
DIAGNOSIS. Endocarp with smoothly rounded, interrupted, longitudinal external
ribs ; median infold broad, rounded ; wall with an external coat of parenchyma.
Length about n mm. (estimated).
HOLOTYPE. A broken endocarp showing the infold. Brit Mus. (N. H.), No.
¥.33961.
DESCRIPTION. Endocarp : One-loculed, ribbed longitudinally, the ribs (about
nine at each end, the number in the middle doubtful) smoothly rounded, interrupted,
and with additional intercalated short ribs like elongate nodulations towards the
middle of the endocarp. A large longitudinal germination valve occupies nearly
half the breadth of the nut bearing on its inner surface a broad, rounded, median,
longitudinal infold (0-4 mm. across, 0-8 mm. deep in the one specimen in which it
could be measured) ; the length of the valve cannot be determined from the imperfect
material available. Endocarp wall hard and woody, variable in thickness from about
0-4 mm. as measured through the grooves, to 0-5 mm. through the ribs, the locule
being smooth, not ribbed in agreement with the external surface of the endocarp.
In section the walls show the following structure : An inner coat of horizontally
aligned sclerenchyma many layers thick giving a transversely striate locule-surface ;
the cells forming the outer layers of this coat gradually pass from a horizontal
to an oblique, and from an oblique to a radial alignment, the radially aligned portion
forming the main thickness of the wall. Along the margins of the valve, however,
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 115
the oblique alignment is never lost but passes across the walls to the periphery
constituting planes of weakness along which the valve separates ; outside the radial
cells are a few peripheral layers of parenchyma, superficially the cells which form
the layers are irregularly polygonal with an average diameter of about 0-025 mm.,
and no definite alignment. These cells give a smooth surface to the endocarp.
Length of endocarp unknown, estimated to be about n mm.; diameter of one
fragment (probably basal end), 2-7 mm. (possibly increased by distortion).
Seed : Not seen ; cells of the testa, 0-03-0-05 mm. in diameter, preserved as
impressions on the locule wall superposed on the transverse striations of the
locule-lining.
REMARKS AND AFFINITIES. Two fragments of Mastixia endocarp were figured
and briefly described by C. &. E. M. Reid (1910 : 166, pi. 16, figs. 73, 74) but no
attempt was made at specific diagnosis. Six more fragments have now been recog-
nized in their collection, two are the ends of nuts showing the complete circumference,
both being distorted obliquely. Another fragment is from the middle of a nut
with both ends and the valve missing ; a fourth, incomplete at the ends, appears
to have broken along the edge of the valve. It suggests a length of about n mm.
for the perfect specimen. All fragments are recognizable by their ribbing and micro-
scopic structure, they therefore appear to offer a sufficient basis for specific
determination.
Detailed histological studies of Mastixia and allied fossil genera from the Brown
Coal of Germany have been published by Kirchheimer in papers from 1934 to 1939.
Many details of the sclerenchyma and parenchyma of the endocarps are shown.
The genus Mastixia was first recognized as a fossil by C. & E. M. Reid (1910). It
has also been recorded more recently from the London Clay (Reid & Chandler,
1933:448, pi. 25, figs. 1-17).
Family PRIMULACEAE
Section LYSIMACHINAE
*
Genus LYSIMACHIA (Tourn.)
Lysimachia boveyana n. sp.
(PI. 17, figs. 179, 180)
DIAGNOSIS. Seeds as in Lysimachia, 0-8-0-85 mm. long, 0-62-0-7 mm. broad.
Surface rugosities forming small areoles with nodular boundaries. There are also
semi-translucent tubercles especially well seen around the margin.
HOLOTYPE. Brit. Mus. (N. H.), No. ¥.33964.
DESCRIPTION. Seed : Rounded triangular in outline, originally gently convex
on the dorsal face, facetted on the ventral face the two facets meeting to form
a longitudinal median angle extending from margin to margin. Hilum elongate
about the middle of this angle. Surface rugose, rugosities forming small areoles
with nodular boundaries about 0-032 mm. in diameter on the ventral side, and 0-032
mm. or larger in diameter on the dorsal side. On the ventral side they are aligned
n6 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
in rows directed from the median angle to the margin, on the dorsal side no such
definite arrangement is apparent. Around the margin semi-translucent tubercles,
0-025 mm- long and 0-022 mm. broad, are visible. Similar tubercles, or their remains
can also be seen over parts of the surface but they are less prominent than around
the margin.
Length of seed, 0-8-0-85 mm.; breadth, 0-62-0-7 mm-
REMARKS AND AFFINITIES. Two seeds. The form, character of testa, and
median hilum as described, are only to be found combined in the family Primulaceae.
Seeds of this shape with a simple hilar ridge extending from edge to edge have been
seen only in the Lysimachinae and closely comparable structure in the genus
Lysimachia.
The majority of species have larger seeds, but in L. japonica some seeds are
comparable in size (0-85 by 0-65 mm. and 0-9 by 0-65 mm. for example) although
average sized seeds are larger (1-05 by 0-9 mm and 0-9 by 0-75 mm.).
The genus Lysimachia is distributed through the temperate and sub-tropical
regions of the whole world.
Family SYMPLOCACEAE
Genus SYMPLOCOS Jacquin
Symplocos anglica n. sp.
(PL 17, figs. 181-186)
1862. Nyssa europaea Unger : Heer, p. 1066, pi. 69, figs. 11-17.
1910. Nyssa europaea Unger : C. & E. M. Reid, p. 167, pi. 15, fig. 9.
DIAGNOSIS. Endocarp ovoid to obovoid or oblong, three-loculed, apical depression
with gently sloping edges markedly truncating the apex of the endocarp. Base
rounded. Outer surface finely wrinkled longitudinally. Length, 3-5-6-7 mm.;
breadth, 2-2-4-4 mm.
HOLOTYPE. A somewhat compressed endocarp. Brit. Mus. (N. H.), No.
V-33965-
DESCRIPTION. Endocarp : Syncarpous, three-loculed, locules arranged around
a central canal, one or two occasionally being abortive, opening by wide apertures
into the large apical depression the edge of which slopes gently inward being neither
thickened nor rounded. At the rounded base is a small attachment scar from
which the funicle passes into the central canal. The endocarp may be either obovoid,
oblong with rounded base, or ovoid, markedly truncated at the top by the apical
depression, but the truncation may be obscured either by the persistent base of the
style or by oblique distortion when it appears ovoid. Wall possibly formed of
fused mesocarp or endocarp (the former preserved only in part if present), or the
mesocarp may be entirely absent ; thickness of wall averaging about 0-25 mm.;
its outer surface is finely wrinkled longitudinally, it usually shows about twelve to
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 117
sixteen irregular, sometimes interrupted and nodular rather obscure longitudinal
ribs ; surface cells mostly very small with obscure outlines but occasionally (when
the mesocarp is abraded ?) the wall is finely and evenly pitted.
Length of endocarp, 3-5-6-7 mm.; breadth, 2-2-4-4 mm- Average length, 4-8
mm.; average breadth, 3-3 mm.; average breadth if uncompressed, about 2-1
mm.; breadth of apical depression as compressed, 1-5-2 mm.
REMARKS AND AFFINITIES. About thirty-four specimens, all much crushed,
some symmetrically others obliquely. The species was described and figured as
Nyssa both by Heer and by C. & E. M. Reid. However it is clear that such
truncated specimens could not be Nyssa whereas they agree in character and size
with the species of Symplocos here described. Other fossil species based on fruits
are commonly much larger and differ in other ways. S. gregaria Unger (1866 : 31,
pi. n, figs, ig-h) is one of the more comparable species but is more variable in size
ranging from 4'i-i2-5 mm. in length, and from 2-1-6-2 mm. in breadth (Kirchheimer,
19360 : 95, pi. 9, figs. 50, b) gives the dimensions as 4-10 mm. length, 2-5-6 mm.
breadth. The fruits of 5. gregaria are represented as larger, more markedly truncate,
with finer, more regular and numerous grooves and with thicker walls. S. kirstei
Kirchheimer (1939 : 285, pi. 3, fig. 3) is also similar but larger (length, 5-5-11 mm.;
breadth, 3-5-6 mm.) and relatively longer and narrower.
A new name, Symplocos anglica, has therefore been given to the Bovey fruits.
Symplocos headonensis Chandler
(PL 17, figs. 187, 188)
1910. ? Nyssa obovata Weber : C. & E. M. Reid, p. 168, pi. 15, fig. 10.
1926. Symplocos headonensis Chandler, p. 40, pi. 7, fig. 3 ; text-fig. 24.
DESCRIPTION. Endocarp : Syncarpous, four-loculed, the locules arranged around
a central canal and opening above by wide apertures into a large apical depression ;
broadly ovoid, urceolate, or sub-globular, conspicuously truncated* by the apical
depression ; margin of the apical depression thickened, somewhat rounded ; attach-
ment indicated by a small sunk scar at the base, thickness of wall, 0-2 mm.; thickness
of septum, 0-05 mm. Surface without ribs, but uneven, surface cells angular,
unequal in size, with a tendency to be aligned in longitudinal rows near the base
thus producing obscure fine striations.
Length of an obscurely four-lobed and slightly urceolate specimen (possibly
immature), 6 mm.; breadth, 5-5 mm. Length of a second well-developed specimen,
7 mm.; breadth, 5-5 mm. Estimated diameter when uncompressed, 3-5 or 3-6 mm.
REMARKS AND AFFINITIES. Three specimens one of which is in the Geological
Survey Museum if the specimen figured by C. & E. M. Reid (1910, pi. 15, fig. 10)
really is of this character. The illustration indicates a larger, broader form than
S. anglica. This species is near in size, form, and general appearance to S. headonensis
Chandler from Hordle which see for the probable relationship of this species to
living forms. The species occurs both at Bovey and Heathfield.
u8 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Family SOLANACEAE
Genus SOLANISPERMUM nov.
DIAGNOSIS. Seeds of unknown generic relationship agreeing with Solanaceae
in form and structure.
Solanispermum reniformis n. sp.
(PI. 17, figs. 189-191)
1910. Carpolithus sp. 5 C. & E. M. Reid, p. 174, pi. 16, fig. 72.
A number of seeds of Solanaceae with distinctive rugose surface which characteristic-
ally weathers into fibres occur in the Lower Bagshot, Bournemouth Freshwater and
Marine Beds, and the Cliff End Beds near Mudeford as well as at Bovey. The
most perfect specimen was found at Branksome Dene and an extremely good seed
showing the hilar aperture was found at Sandbanks.
In order that the description should be as complete as possible it has been largely
based on material from the Bournemouth area.
DIAGNOSIS. Seed transversely oval or reniform in outline, occasionally hooked,
surface normally with coarse, interrupted, sinuous rugosities or tubercles which
produce a pitted effect in places. An outer coat, rarely preserved, shows " pits "
with sinuous outlines. The rugose coat shows fine striae at right angles to the
tubercles. Splitting along the striae on drying produces a fibrous effect. Inner
coat spongy formed of equiaxial cells. Maximum diameter of seeds about 3'5-4'8 mm.
HOLOTYPE. A perfect seed figured by C. & E. M. Reid (1910, pi. 16, fig. 72).
Geol. Surv. Colin. No. 1805.
DESCRIPTION. Seed : Bisymmetric, flattened or slightly inflated, transversely
oval or reniform in outline, occasionally hooked. Hilum usually marginal occupying
part of the concave margin in reniform seeds and one of the longer margins in oval
seeds, large and gaping, elongate-oval leading into a small cavity separated from the
main seed-cavity by a thin curved partition seen in longitudinal sections of the seed.
Probably the funicle lay close to this partition within the hilar cavity. A few seeds
are distorted (in growth) so that a gaping hilum is twisted on to one of the broad
surfaces. Dorsi- ventral flattening of such distorted seeds may have occurred.
Micropyle usually marginal, adjacent to the hilum. Its position is most apparent
in hooked seeds where it occupies the extremity of the hook. Surface occasionally
showing traces of an outer coat with coarse digitate cells, but in most specimens
this is worn away. As normally preserved ornamented with interrupted sinuous
rugosities which diverge from the hilum. They produce a pitted effect in places,
the pits being about o-05-o-i mm. in diameter. The rugose seed-coat is about
o-i mm. thick ; it appears striate, the striae crossing the rugosities more or less at
right angles and lying parallel with the margin near the circumference of the seed.
Splitting tends to occur along the striae in weathered or dried specimens producing
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 119
a fibrous effect. Close examination of the " fibres " shows them to be formed of
fine equiaxial cells 0-012 mm. in diameter. Several layers of such " fibres " occur
in this integument. Within it is a spongy coat, 0-4 mm. thick, formed of compact
soft parenchyma. The lining of the seed-cavity is striate, the striae diverging
from the neighbourhood of the hilum (actually from the closely associated chalaza) .
Maximum diameter of seeds, 4-8 mm.; commonly 3-5 mm.; diameter at right
angles to it, 2-25-3-6 mm.
REMARKS AND AFFINITIES. One seed figured by C. & E. M. Reid (1910) from
Heathfield and another (now broken) from the same pit collected by E. M. Reid
and Chandler in 1932. The curved outline, flattened seed, large gaping marginal
hilar cavity and even the mode of distortion which brings the gaping hilar cavity
on to one of the broad surfaces all suggest Solanaceae. Seeds of Capsicum and
allied genera show a general resemblance but the succession of coats cannot be
matched exactly in any genus examined. Striate fibrous coats, or striate fibrous
outgrowths of the testa are found in Lycopersicum and Cyphomandra.
Reference to a living genus cannot be made and even the reference to the family
is somewhat tentative until living seeds with a closely comparable succession of
coats have been found. Meanwhile these readily recognizable specimens are referred
to a new genus Solanispermum as it is in Solanaceae that the closest resemblance
has so far been traced.
INCERTAE SEDIS
Carpolithus sp.
(PI. 17, figs. 192-194)
Fruit : Inferior with remains of three small triangular patent perianth segments
at the apex. Elongate having three broad surfaces two of which are ventri-lateral
and one dorsal. Surfaces separated by longitudinal angles, the lateral angles sharp
and almost flanged, the ventral one fibrous. The broad perianth segments each
lie opposite one of the broad surfaces. In profile the fruit is long and narrow.
The ventral angle is straight. The dorsi-lateral angles are convex hence the two
ventri-lateral surfaces are more or less semi-oval (but unequal). The dorsal face
is narrowly oval. Ventral surfaces smooth formed of fine, close, elongate, obliquely
aligned fibres, 0-008 mm. broad. Dorsal surface with similar fibres transversely
aligned. Fibrous surfaces often concealed by a rough coat of equiaxial cells, 0-025
mm. in diameter.
Length, 2-25-3-5 mm.; breadth, 1-1-3 mm- Breadth of ventral faces in one
specimen, i and 0-3 mm. respectively, and in a second specimen, 0-5 and 0-52 mm.
respectively.
REMARKS. Thirteen endocarps from Heathfield. The relationship of these
small fruits has not been established. The difference in form of the dorsal and two
ventral faces or facets suggests that the fruits grew in close association with one
another, the ventral faces possibly in actual contact.
GEOL. Ill, 3. IQ
120 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
Carpolithus sp.
(PL 17, figs. 195-197)
A three-, four- or possibly five-partite capsule, the segments united still at the
base. Segments lanceolate, three only preserved, two being attached, the third
represented by a detached distal end. They diverge from one another and have
strongly incurved tips. The length of one segment, 4-2 mm.; breadth, 0-9 mm. (but
the full length is not shown owing to the strong incurving of the tip) . The second
(attached) segment 3-6 mm. with deeply incurved tip by 1-5 mm. The number of
segments in the perfect fruit can only be guessed by the angle between existing
segments.
Segments highly rugose and even nodular on both surfaces ; margins greatly
thickened and almost revolute. Hence the segments are concave externally, the
broader one having a slight median ridge which makes it biconcave. All show traces
of spines near the base, one each side of the median line in the shorter of the two
attached segments. Cells of outer surface equiaxial, about 0-016 mm. in diameter.
Carpolithus sp.
(PI. 17, figs. 198, 199)
Endocarp (?) : Syncarpous, sub-ovoid, three-lobed and three-carpelled, or two-
lobed and two-carpelled by suppression, the base being sunk between the lobes.
Dehiscing loculicidally from the apex (which is always broken irregularly), almost to
the base. No central axis seen. Wall close-textured, hard, 0-033 mm. thick
(cells indistinct owing to intensely carbonized condition in section) . Surface smooth,
of equiaxial cells 0-016 mm. in diameter usually evenly distributed but occasionally
aligned in obscure longitudinal rows. Septa very thin, columnar in section.
Locule lined by equiaxial cells, 0-012 mm. in diameter.
Length of longest specimen, 5-5 mm. (incomplete at apex) ; breadth, 3-25 mm.
Length of second specimen, 4 mm.; breadth, 2-1 mm.
Seed (lying within the locule near the broken apex) : Linear with a median ridge
on the concave side at one end, somewhat tufted at the other end, too decayed to
show cell-structure. Length, 2-7 mm.; breadth, 0-6 mm. at the broadest estimate,
but actually bent so as to appear only 0-5 mm. broad.
REMARKS. Two specimens from Bovey, one from Heathfield. Also three
very imperfect specimens from Bovey which may belong to this species. The
relationship is undiscovered.
Carpolithus sp.
(PL 17, figs. 200-202)
Several much collapsed and immature fruits are narrow-obovoid or urceolate,
three-lobed, three-loculed, the lobes being rather slender and much smaller than those
figured in PL 17, figs. 198, 199. Base sunk between the lobes. Apex in best
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 121
preserved specimens with three minute patent persistent perianth segments one
corresponding to each lobe. Dehiscence by loculicidal splitting from the apex
downwards showing apical canals (or canal) (style ?). Surface dull, finely rugose,
cells aligned in obscure longitudinal rows. Locule transversely striate, striae
0-0125-0-017 mm. apart due to small cells aligned in transverse rows. Thickness
of walls about 0-025 mm.; thickness of septum (where seen near the apex), 0-016
mm. Length and breadth of four specimens respectively : 4-6 mm. (broken at
one end) X 2-4 mm., 4-2 mm. x 2 mm., 4-2 X 2 mm., 4-6 mm. x 2 mm.
Seed : Occupying the whole length of the locule, flattened. Longitudinally
striate.
REMARKS. The form of this tiny fruit and the three minute patent perianth
segments recall the rather larger, flattened, immature fruits of Eomastixia bilocularis
found in the Bournemouth Beds and the considerably larger immature specimens of
another Eomastixia from Lake (awaiting description). There is nothing in the
limited evidence available to exclude such a relationship although it should be
noted that living Mastixia perianth segments are four- to five-partite. A species of
Mastixia occurs in the Bovey lignites but any suggestion as to the relationship
must be regarded as tentative only.
Carpolithus sp.
(PL 17, fig. 203)
An elongate ovoid seed with a few longitudinal angles or crumples. No definite
organs are visible but the alignment of cells at the pointed end and crumples at the
broad end suggest the presence of organs in these positions. Surface formed of
inflated longitudinally aligned cells about 0-025 mm. long and o-ooi mm. broad,
but over much of the surface the cells have been abraded so that no cell-structure
can be seen.
Length of seed, 2-6 mm.; breadth, 0-9 mm.
Bulbil ?
*
(PL 17, fig. 204 ; Text- fig. 3)
Two sub-spherical bodies, slightly flattened on one side with a large deep depression
at the middle of the flat surface, and a second similar depression on one side, may
FIG. 3. Bulbil ? Diagrammatic section showing overlapping
layers of thick bracts, x 4 approx.
GEOL. in, 3. to§
122 THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN
be bulbils or buds. They show superficially an obscure network of shallow furrows.
When fractured they reveal overlapping layers of thick bracts. Diameter about
6-6-7 mm. Relationship not known.
REFERENCES
BANDULSKA, H. 1923. A Preliminary Paper on the Cuticular Structure of certain Dicotyle-
donous and Coniferous Leaves from the Middle Eocene Flora of Bournemouth. /. Linn.
Soc. Bot., London, 46 : 241-268, pis. 20, 21.
1924. On the Cuticles of some Recent and Fossil Fagaceae. /. Linn. Soc. Bot., London,
46 : 427-441, pis. 39, 40.
BRAUN, AL. 1845. Die Tertiar Flora von Oeningen. N. Jb. Min. Geol. Palaont., Stuttgart,
1845 : 164-173.
CHANDLER, M. E. J. 1921. Note on the Occurrence of Sequoia in the Headon Beds of Hordwell,
Hants. Ann. Bot., London, 139 : 457.
1922. Sequoia couttsiae, Heer, at Hordle, Hants : A study of the characters which serve
to distinguish Sequoia from Athrotaxis. Ann. Bot., London, 143 : 385-390, 5 figs.
1923. The Geological History of the Genus Stratiotes : An Account of the Evolutionary
Changes which have occurred within the Genus during Tertiary and Quaternary Times.
Quart. J. Geol. Soc. Lond., 79 : 117-138, pis. 5, 6.
1925. The Upper Eocene Flora of Hordle, Hants, 1. 32 pp., 4 pis. Mon. Palaeont. Soc.,
London.
1926. The Upper Eocene Flora of Hordle, Hants, 2. vii + 20 pp., 4 pis. Mon. Palaeont.
Soc., London.
FLORIN, R. 1919. Eine Ubersicht der fossilen Salvinia — Arten mit besonderer Beriick-
sichtigung eines Fundes von Salvinia formosa Heer im Tertiar Japans. Bull. Geol. Inst.
Uppsala, 16 : 243-260, pi. n.
1938-45. Die Koniferen des Oberkarbons und des Unteren Perms, 1-8. Palaeonto-
graphica, Stuttgart, 85 (B) : 729 pp., 186 pis.
GARDNER, J. S. 1879-82. A Monograph of the British Eocene Flora, 1 : Filices. 86 pp.,
13 pis. Mon. Palaeont. Soc., London.
1883-86. A Monograph of the British Eocene Flora, 2 : Gymnospermae. 159 pp., 27 pis.
Mon. Palaeont. Soc., London.
GOTHAN, W. & SAPPER, J. 1933. Neues zur Tertiarflora der Niederlausitz. Inst. Palaobot.
Petrogr. Brennsteine, Berlin, 3 : 1-44, pis. 1-7.
HEER, O. 1862. On the Fossil Flora of Bovey Tracey. Philos. Trans., London, 152 : 1039-
1086, pis. 55-71.
i862a. On certain Fossil Plants from the Hempstead Beds of the Isle of Wight. Quart.
J. Geol. Soc. Land., 18 : 369-377, pi. 18.
KIRCHHEIMER, F. 1929. Die Gattung Salvinia in den Tertiarfloren der Wetterau und des
Vogelsberges. Ber. oberhess. Ges. Nat.-u. Heilk., Giessen (n.f.) 12 : 140-160.
192913. Die fossilen Vertreter der Gattung Salvinia Mich., I. Ein Beitrag zur kenntnis
der Mikrosporangien der Salvinia formosa Heer. Planta, Berlin, 9 : 388-406, 8 figs.
1930. Die fossilen Vertreter der Gattung Salvinia Mich., I. Die bisherigen Funde von
Sporangienresten und Sporen tertiaren Salvinien. Zbl. Min. Geol. Palaont., Stuttgart,
1930 : 339-349-
1930^. Die fossilen Vertreter der Gattung Salvinia Mich., II. Uber Sporangienrest
einer miozanen Salvinie. Planta, Berlin, 11 : 169-206, 19 figs.
1931. Die fossilen Vertreter der Gattung Salvinia Mich., III. Uber einen neuen Func
von Resten der Mikrosporangien einer miozanen Salvinie. Planta, Berlin, 13 : 102-1 13, 5 fi£
1932. Zur morphologie der Salvinia macrophylla Kirch, aus dem miozanen Ton vor
Lauterbach (Oberhessen) . Palaont. Z., Berlin, 14 : 309-314, 2 figs.
• 1934. Das Hauptbraunkohlenlager der Wetterau. 51 pp., 10 pis. Hanau.
THE OLIGOCENE FLORA OF THE BOVEY TRACEY LAKE BASIN 123
• 1935. Bau und botanische Zugehorigkeit von Pflanzenresten aus deutschen Braunkohlen.
Bot. Jb., Leipzig, 67 : 37-122, pis. 1-13.
1936. Zur Kenntnis der Friichte rezenter und fossiler Mastixioideen. Bot. Zbl., Dresden,
55 : 275-300, pis. 5-8.
19360. Beitrage zur Kenntnis der tertiarflora Friichte und Samen aus dem deutschen
Tertiar. Palaeontographica, Stuttgart, 82 (B) : 73-141, pis. 7-13.
1937- Palaobotanische Beitrage zur Kenntnis des Alters deutscher Braunkohlenschichten,
II. Die braunkohlefiihrenden Tone von Siegburg (Rheinland) und Karaenz (Oberlausitz).
Braunkohle, Halle, 50 : 893-931, 26 figs.
• 19370- Grundzeige einer Pflanzenkunde der deutschen Braunkohlen. 1. 53 pp., 117 figs.
Saale.
• 1938. Beitrage zur naheren Kenntnis der Mastixioideen-Flora des deutschen Mittel-
bis Oberoligozans. Bot. Zbl., Dresden, 58 (B) : 303-375, pis. 3-8.
— 1939. Tertiare Dikotyledonenreste und ihr systematischer Wert. Flora, Jena, 133 :
239-296, pis. 1-3.
i939a. Uber die botanische Zugehorigkeit weiterer Frucht- und Samenreste, besonders
aus den Braunkohlenschichten Sachsens. Planta, Berlin, 29 : 262-278, 3 figs.
KRAUSEL, R. 1920. Nachtrage zur Tertiarflora Schlesiens. Jb. preuss. geol. Landesanst.,
Berlin, 39 : 329-417, pis. 16-27.
LUDWIG, R. 1857. Fossile Pflanzen aus der jiingsten Wetterauer Braunkohle. Palaeonto-
graphica, Stuttgart, 5 : 81-110, pis. 16-23.
1860. Fossile Pflanzen aus der altesten Abtheilung der Rheinisch- Wetterauer Tertiar-
formation, Palaeontographica, Stuttgart, 8 : 39-154, pis. 6-60.
PENGELLEY, W. 1863. The Lignites and Clays of Bovey Tracey. Philos. Trans., London,
153 : 1019-1038.
REID, C. & E. M. 1910. The Lignite of Bovey Tracey. Philos. Trans., London, 201 (B) :
161-178, pis. 15, 16.
1915. The Pliocene Floras of the Dutch-Prussian Border. Meded. Rijksopsp. Delfst.,
Amsterdam, 6 : 178 pp., 20 pis.
REID, E. M. 1920. Recherches sur quelques graines Pliocenes du Pont-de-Gail (Cantal).
Bull. Soc. geol. Fr., Paris (4) 20 : 48-87, pis. 3, 4.
REID, E. M. & CHANDLER, M. E. J. 1926. The Bembridge Flora. Catalogue of Cainozoic
Plants in the Department of Geology, i. viii -f- 206 pp., 12 pis. Brit. Mus. (Nat. Hist.),
London.
1933. The Flora of the London Clay, viii + 561 pp., 33 pis. Brit. Mus. (Nat.
Hist.), London.
SIMPSON, J. B. 1936. Fossil Pollen in Scottish Tertiary Coals. Proc. Roy. Soc. Edinb.,
56 : 90-108, pis. 1-3.
UNGER, F. 1860. Sylloge Plantarum Fossilium, I. Denkschr. Akad. Wiss. Wien, 19 : 1-48,
pis. 1-2 1.
1866. Sylloge Plantarum Fossilium, III. Denkschr. Akad. Wiss. Wien, 25 : 1-76, pis.
1-24.
PLATE ii
Osmunda lignitum (Giebel)
FIG. i. Two sporangia which have burst and become interlocked. x 28. (¥.33833.)
FIG. 2. Diagram to explain Fig. i. Annulus at (a).
FIG. 3. The same pair of sporangia, opposite surface, x 28.
FIG. 4. Diagram to explain Fig. 3. Annulus at (a).
FIG. 5. A single burst sporangium showing the annulus at (a), x 60 approx. (V. 33833.)
FIG. 6. A group of spores embedded in remains of sporangium from the specimen in Figs,
i, 3. The spores show the fine granulations of the surface. x 400. (V. 33833^)
All the above are from Bovey.
Salvinia boveyana n. sp.
FIG. 7. A sporocarp showing the globular thin- walled sporangia projecting through the
wall, x 28. (V.33834.)
FIG. 8. Another less mature specimen showing a closely compacted mass of sporangia.
Also figured C. & E. M. Reid, 1910, pi. 16, fig. 57. x 12. Geol. Surv. Mus. Colin. No. 76682.
FIG. 9. A sporangium showing spores embedded in a froth-like mass, x 150. (¥.33835.)
FIG. 10. Another sporangium with stalk, x 150. ^.33835.)
FIG. ii. Part of another, x 400. (¥.33835.)
All the above are from Bovey.
Polamogeton tenuicarpus C. & E. M. Reid
FIG. 12. An endocarp, side, with keel on the right beginning to gape. The seed (s),
protrudes at the apex. x 15. (V. 33836) Bovey.
FIG. 13. Another endocarp showing the gap between the ends of the curved carpel. x 15.
(V.33837) Bovey.
FIG. 14. Pollen-grain probably belonging to this species. x 1000. J. B. Simpson Colin.
Heathfield.
Stratiotes websteri (Brongniart)
FIG. 15. A seed, ventral side, (c] collar. x 6-5. (¥.33838.)
FIG. 1 6. The same, dorsal, looking on to the keel. The specimen is somewhat crushed
dorsiventrally. x 6-5.
FIG. 17. Part of a seed which has begun to split and burst. It shows the smooth rounded
collar and the keel (k) arising out of it. x 6-5. (V. 33839.)
FIG. 1 8. Valve of a seed, broken at the micropylar end, inner surface showing the short
transverse raphe (r). x 6-5. (¥.33840.)
FIG. 19. Another broken valve as in Fig. 1 8. X 6-5. (¥.33841.)
All the above are from Bovey.
Caricoidea nitens (Heer)
FIG. 20. A laterally compressed fruit, truncate at the base. Also figured Heer (1862, pi. 70-
fig. 1 8). x 6 approx.
FIG. 21. Another fruit. The impression of the calyx is clearly seen in profile at the base.
Also figured Heer (1862, pi. 70, fig. 16). x 6 approx.
FIG. 22. Base of a dorsiventrally compressed fruit showing the basal calyx scar in the
centre of which is the plug closing the passage to the locule. x 6 approx.
FIG. 23. Holotype. Figured Heer (1862, pi. 70, fig. 20). A laterally compressed fruit from
which a tangential slice had been cut to display the small locule. x 6 approx. (¥.33842.)
All the above are from Bovey.
Figs. 8, 20-23: Photo C. Reid; Figs. 9-11 : Photo W. N. Croft; Fig. 14: Photo J. B.
Simpson.
Bull. B.M. (N.H.) Geol. 3, 3
PLATE 11
14 20 21 22
OSMUNDA, SALVINIA, POTAMOGETON, STRATIOTES, CARICOIDEA
PLATE 12
Calamus daemonorops (Unger)
FIGS. 24, 25. Two young fruits or female flowers showing three styles, x 15. (V. 33846 —
47-)
FIG. 26. Somewhat older fruit ; style bases are preserved but the three-fid style has
disappeared, x 15. (¥.33848.)
FIG. 27. A fruit showing clearly the inner and outer perianth segments. x 15. (V. 33849.)
FIG. 28. A better developed fruit also showing the two whorls of persistent perianth seg-
ments, x 15. (V. 33850.)
FIG. 29. Base of immature female fruit, the striate bracts in two whorls each of three bracts.
X 15- (V.3385I-)
FIG. 30. Small immature fruit showing overlapping reflexed scales, x 15. (V. 33852.)
FIG. 31. A larger, better developed but much compressed fruit showing the overlapping
reflexed scales, x 15. (¥.33853.)
FIG. 32. The three-partite bract-like perianth of a male flower, x 15. (¥.33854.)
FIG. 33, 34. Two more male flowers, x 15. (¥.33855-56.)
FIG. 35. Fragment of a fruiting axis, x 6-5. (¥.33857.)
FIG. 36. A smaller fragment of an axis. X 6-5. (V. 33858.)
FIG. 37. Immature seed extracted from a fruit, x 15-5 (¥.33859.)
FIGS. 38-41. Spines and spine bases, x 2-8. (¥.33860-63.)
FIG. 42. A pollen-grain (doubled on itself), x 1000. J. B. Simpson Colin.
FIG. 43. A crumpled immature seed possibly belonging to this species, x 15-5. (¥.33864.)
FIG. 44. The same, opposite side, x 15-5.
All the above (except Figs. 37 and 42 from Heathfield) are from Bovey.
Myrica boveyana (Heer)
FIG. 45. An endocarp showing somewhat rugose surface, x 6-5. (V. 33865.)
FIG. 46. A laterally flattened endocarp which may belong to this species. X 6-5. (¥.33866.)
FIG. 47. One valve of an endocarp, internal surface, showing coat of equiaxial cells :
(/) funicle, (st) stylar canal, x 15. (¥.33867.)
FIG. 48. Neotype. Another valve from a broader fruit. Interior. Lettering as above.
X 15- (V.33868.)
All the above are from Bovey.
Fig. 42 : Photo J. B. Simpson.
Bull. B.M. (N.H.) Geol. 3, 3
PLATE 12
CALAMUS, MYRICA
PLATE 13
Corylus sp.
FIG. 49. Fragment of an inflorescence of male flowers with anthers. X 15. (V. 33869.)
Heathfield.
FIG. 50. The same, opposite side. X 15.
FIG. 51. Pollen grain from the above, x 1000. J. B. Simpson Colin.
Carpinus boveyanus (Heer)
FIG. 52. Neotype. Fruit with accrescent calyx preserved showing longitudinal furrows
associated with fibres which arise from the margin of the basal scar (s) of attachment to the
wing-like bract, x 6-5. (¥.33870.)
FIG. 53. Another longer, narrower fruit ; (s) as above, x 6-5. (¥.33871.)
FIG. 54. Another fruit, x 6-5. (V. 33872.)
FIG. 55. Another, x 12. (¥.33873.) Heathfield.
FIG. 56. Another, x 15. (¥.33874.)
FIG. 57. A small fruit with remains of superior perianth at the apex of the accrescent
calyx ; one of the two styles is preserved, x 6-5. (¥.33875.)
FIG. 58. A specimen with styles preserved. Perianth much worn, x 6-5. (¥.33876.)
FIG. 59. A fruit, the lateral scar near the base indicates that it was one of a pair of fruits,
the other being but little developed, x 12. (¥.33877.) Heathfield.
FIG. 60. A twinned fruit, x 12. (¥.33878.) Heathfield.
FIG. 61. A small twinned fruit, x 12. (¥.33879.) Heathfield.
FIGS. 62-65. Four twinned fruits. In Fig. 62 part of the basal end of one of the pair is
broken away exposing the locule. The upper part is splitting in the plane of symmetry. Fruits
in Figs. 64, 65 very unequally developed, x 6-5. (¥. 33880-83.)
FIG. 66. Base of dorsiventrally compressed fruit. Carpinus ? x 6-5. (¥.33884.)
FIG. 67. One valve (interior) of another dorsiventrally compressed fruit probably Carpinus.
X 6-5. (V.33885.)
Fagus minima n. sp.
FIG. 68. Holotype. An endocarp : (s) basal scar of attachment, x 6-5. (¥.33886.)
FIGS. 69, 70. Two endocarps. x 6. The figures are reproduced from C. & E. M. Reid
(1910, pi. 16, figs. 67, 68). Geol. Surv. Mus. Colin. No. 76683.
Zelkova boveyana n. sp.
FIG. 71. Holotype. Endocarp, lateral aspect : (st) style, (a) attachment, x 15.
(V.33887.)
FIG. 72. The same, marginal view, x 15.
FIG. 73. The same, as in Fig. 72 but more tilted to show the attachment (a). X 15.
Moroidea boveyana n. sp.
FIG. 74. Holotype. Fruit, side. The rounded base is broken, (st) style; (/) projection
marking the point of entry of the funicle to the sub-apical placenta, x 15. (¥.33888.)
Brasenia ovula (Brongniart)
FIG. 75. A somewhat crumpled and distorted seed showing longitudinal corrugations of
the surface due to effects of contraction and alignment of the cells : (e) position of embryotega-
X 15- (¥.33889.) Heathfield.
Magnolia boveyana n. sp.
FIG. 76. Holotype. A seed, concave surface with raphe : (ch) chalaza. x 6-5. (¥.33890.)
Heathfield.
FIG. 77. The same, opposite convex surface. x 6-5.
FIG. 78. Another seed, raphe side. X 6-5. (¥.33891.) Heathfield.
FIG. 79. The same, opposite side, x 6-5.
FIG. 80. A seed longitudinally sectioned showing the canal (ca) through the chalaza-plug.
X 6-5. (¥.33892.) Heathfield.
Unless otherwise stated all the above are from Bovey.
Fig. 51 : Photo J . B. Simpson ; Figs. 55, 59-61, 70 : Photo C. Reid.
Bull. B.M. (N.H.) Geol. 3, 3
PLATE 13
76 ~ 77 ^78 ^ 79 80
CORYLUS, CARPINUS, FAGUS, ZELKOVA, MOROIDEA, BRASENIA, MAGNOLIA
PLATE 14
LAURACEAE
FIG. 81. An empty cupule with simple rim tilted to show the inner surface and scar of
attachment of the berry, x 6-5. (¥.33893.)
FIG. 82. Another, x 6-5. (¥.33894.)
FIG. 83. A wrinkled leathery type of cupule with entire rim. x 6-5. (¥.33895.)
FIG. 84. Leathery cupule with distinct sepals at the rim. x 6-5. (¥.33896.)
FIG. 85. Another of similar type but much more slender, x 6-5. (¥.33897.)
FIG. 86. Small cupule with distinct sepals enclosing berry, x 6-5. (¥.33898.)
FIG. 87. Cupule with entire rim, berry enclosed, x 6-5. (¥.33899).
FIG. 88. Small cupule with berry. x 6-5. (¥.33900.) Bovey.
FIG. 89. A detached berry, imperfect below. X 6-5. (¥.33901.)
FIG. 90. Incomplete berry having glandular secretions beneath the skin, x 6-5.
(V. 33902.) Bovey.
FIG. 91. Another, x 6-5. (¥.33903.) Bovey.
Unless otherwise stated all the above are from Heathfield.
Capparidispermum boveyanum n. sp.
FIG. 92. Holotype. A seed showing curved form and ornamentation : (h) hilum. x 15.
(V.33904.)
FIG. 93. The same, opposite side, x 15.
FIG. 94. Another seed showing the hilar aperture (h) very clearly, x 15. (¥.33905.)
FIG. 95. A larger but more compressed seed in which the cotyledonary limb is somewhat
more incurled. x 15. (¥.33906.)
FIG. 96. One valve of a seed, inner surface, showing the curved cavity. The form of the
partition between the limbs is highly characteristic of Capparidaceae. x 15. (V. 33907.)
All the above are from Bovey.
HAMAMELIDACEAE Genus ?
FIG. 97. A seed, side, showing a lobe of the large hilar scar at (/). x 6-5. (¥.33908.)
Bovey.
FIG. 98. The same seed at right angles to Fig. 97. The hilar scar (h) crosses the proximal
end of the seed and has a lobe on each of its broad faces, x 6-5.
FIG. 99. A seed of another species showing the long narrow hilar scar (h). At the opposite
end of the seed a small portion of the septum between two locules adheres and shows a strand
of fibres from the axis of the carpel, x 6-5. (¥.33909.) Heathfield.
Rubus microspermus C. & E. M. Reid
FIGS. 100-105. Six endocarps showing variations in form and size, x 15. (¥.33910-15.)
Bovey.
FIGS. 106-109. Four endocarps. x 15. (¥.33916-19.) Heathfield.
Bull. B.M. (N.H.) Geol. 3, 3
PLATE 14
106 ' 105 107 108 " 109
LAURACEAE, CAPPARIDISPERMUM, HAMAMELIDACEAE, RUBUS
PLATE 15
LEGUMINOSAE Genus ?
FIG. no. A crushed seed. X 6-5. (¥.33920.)
FIG. in. Another, x 6-5. (¥.33921.)
FIG. 112. Part of a third seed showing hilum and the testa in section. x 6-5. (¥.33922.)
The above are from Heathfield.
Meliosma reticulata (C. & E. M. Reid)
FIG. 113. Valve of an endocarp, external surface showing reticulations. X 6-5. (V. 33923.)
FIG. 114. The same, interior : (/) funicular canal. X 6-5.
FIG. 115. Neotype. A perfect but laterally compressed endocarp. X 6-5. (V. 33924.)
FIG. 116. The same, opposite surface, x 6-5.
FIG. 117. A dorsiventrally compressed endocarp, looking on to the attachment (centre of
figure), x 6-5. (¥.33925.)
FIG. 118. The lower part of one valve of an endocarp, interior showing the attachment
and short funicular canal (/). X 6-5. (¥.33926.)
All the above are from Bovey.
Parthenocissus britannica (Heer)
FIG. 119. Neotype. A typical seed, dorsal, showing long narrow chalaza. x 6-5.
(¥.33927.) Bovey.
FIG. 1 20. The same, ventral, showing lateral infolds diverging upwards, x 6-5.
FIG. 121. Another seed, dorsal. X 6-5. (¥.33928.) Bovey.
FIG. 122. The same, ventral, x 6-5.
Parthenocissus boveyana n. sp.
FIG. 123. Holotype. Seed, dorsal, x 6-5. (¥.33929.) Bovey.
FIG. 124. The same, ventral, x 6-5.
FIG. 125. Another seed with outer coat removed, dorsal (base broken) . X 6-5. (¥.33930.)
Bovey.
Vitis hookeri Heer
FIG. 126. Seed (imperfect above on the left), dorsal x 6-5. (¥.33931.) Heathfield.
FIG. 127. The same, ventral, x 6-5.
Vitis stipitata n. sp.
FIG. 128. Seed (imperfect on the left), dorsal, x 6-5. (¥.33932.) Heathfield.
FIG. 129. The same, ventral, x 6-5.
Till a sp.
FIG. 130. A group of anthers, x 15. ^.33933.)
FIG. 131. Another group of anthers, x 15.
FIG. 132. A pollen grain from anthers in Fig. 131. x 1000. J. B. Simpson Colin.
The above are from Heathfield.
Microdiptera parva n. gen. et sp.
FIG. 133. Seed, dorsal, with pitted oval germination valve and striate wing surface, x 15-5.
(V-33934-)
FIG. 134. The same, ventral, showing median longitudinal raphe ridge also a furrow on
each side of it flanked by a wing, x 15 -5.
FIG. 135. Seed with pointed wings, dorsal : (h) hilum, x 15-5 (¥.33935.)
Bull. B.M. (N.H.) GeoL 3, 3
PLATE 15
138 143 145
LEGUMINOSAE, MELIOSMA, PARTHENOCISSUS, V1TIS, TILIA. MCRODIPTERA
FIG. 136. The same, ventral, x 15-5.
FIG. 137. Seed, dorsal. The germination valve has come away and the cavity is full of
sand, x 15-5. (¥.33936.)
FIG. 138. The same, ventral, showing wings and raphe ridge very clearly, x 15-5.
FIG. 139. Seed, dorsal, pitted valve very clear, x 15-5. (V. 33937.)
FIG. 140. Same, ventral, x i5'5-
FIG. 141. Seed, dorsal. Xi5'5. (¥.33938.)
FIG. 142. Same, ventral. Cavities flanking raphe ridge show clearly as they are filled with
sand, x I5'5-
FIG. 143. Seed with asymmetrically developed wings and clearly denned valve beginning to
gape. x 15-5- (V-33939-)
FIG. 144. Seed, dorsal. Symmetric wings show striations very clearly, x I5'5- ^.33940.)
FIG. 145. Same, ventral, x 15*5.
FIG. 146. Small narrow seed, dorsal, x 15-5. (¥.33941.)
FIG. 147. Same, ventral, x 15-5.
FIG. 148. Small seed, dorsal, x 15-5. (V. 33942.)
FIG. 149. Same, ventral, x i5'5-
All the above are from Bovey.
Fig. 132 : Photo by J. B. Simpson.
PLATE 16
Nyssa boveyana n. sp.
FIG. 150. Holotype. Endocarp, dorsal, showing the valve (v). x 6-5. (V. 33944.)
FIG. 151. Same, ventral, x 6-5.
FIG. 152. Another endocarp, dorsal, valve beginning to open, x 6-5. ^.33945.)
FIG. 153. The same, ventral, x 6-5.
FIG. 154. An endocarp, dorsal, valve detached exposing part of locule. x 6-5. ^.33946.)
FIG. 155. The valve removed from the above. x 6-5.
FIG. 156. Apical end of an endocarp showing valve in position exceptionally clearly, x 6-5.
(V-33947-)
FIG. 157. A large laterally compressed endocarp. x 6-5. (V. 33948.)
FIG. 158. Detached valve from endocarp of Nyssa sylvatica var. europaea for comparison.
X 6-5. (V.33949.) Pliocene ; Reuver.
FIG. 159. Detached valve from N. sylvatica var. biflora for comparison, x 6-5. Recent.
Myrtospermum boveyanum n. sp.
FIG. 160. Holotype. Seed showing curved form and characteristic pitting, x 15.
(V.33950-)
FIG. 161. Typical seed, usual preservation, showing indications of curved form and condyle
between the limbs, x 15. (V. 3 3951.)
FIG. 162. Same, opposite side, x 15.
FIG. 163. Another seed with clear indications of curved cavity, x 15. (V. 33952.)
FIG. 164. Seed showing clear indications that the pits were originally inflated, x 15.
(V.33953-)
FIG. 165. Somewhat distorted seed, x 15. ^.33954.)
FIG. 1 66. Incomplete seed, internal surface of one valve showing condyle : (h) hilum
leading into raphe cavity in condyle. Structures are rarely seen in these highly compressed
seeds of which the valves are reluctant to separate, x 15. ^.33955.)
FIG. 167. Two closely adpressed seeds in position of growth, one ? abortive, x 15.
(V.33956-)
FIG. 168. The same pair of seeds, opposite side, x 15.
Myrtospermum dubium n. sp.
FIG. 169. Sub-circular seed, x 15. ^.33957.)
FIG. 170. Opposite side of same seed, x 15.
FIG. 171. Seed of irregular form, x 15. (V. 33958.)
FIG. 172. Another somewhat larger seed, x 15. ^.33959.)
Myrtospermum sp.
FIG. 173. Seed, imperfect at hilar end. X 15. (V. 33960.) Heathfield.
FIG. 174. One valve of the above, internal surface, showing curved cavity, broad triangular
canal in condyle and columnar testa cells as seen in section, x 15.
Unless otherwise stated all the above are from Bovey.
Bull. B.M. (N.H.) Geol. 3, 3
PLATE 16
169
170 ^HP~ 173
NYSSA, MYRTOSPERMUM
PLATE 17
Mastixia boveyana n. sp.
FIG. 175. Holotype. One end of an endocarp. x 6-5. (V. 33961.)
FIG. 176. Opposite aspect of same fragment showing locule in transverse section with its
valve and infold, x 6-5.
FIG. 177. One end of another endocarp, side. x 6-5. (¥.33962.)
FIG. 178. Fragment of endocarp, external surface, at about the middle showing ribbing.
X 6-5. (¥.33963.) The above are from Heathfield.
Lysimachia boveyana n. sp.
FIG. 179. Holotype. Seed, ventral, showing two facets and ridge on which the attachment
lies, x 28. (¥.33964.)
FIG. 1 80. The same, dorsal. The longitudinal shadow marks a crack in the testa. x 28.
Symplocos anglica n. sp.
FIG. 181. Holotype. An endocarp somewhat compressed and showing the apical depres-
sion, x 6-5. (¥.33965.)
FIG. 182. Endocarp, side, slightly distorted so as to show part of the apical depression.
X 6-5. (V.33966.)
FIG. 183. The same, more tilted, so as to show the three apical apertures. x 6-5.
FIG. 184. Another, side, showing truncated apex and wrinkled surface, x 6-5. (¥.33967.)
FIG. 185. Small endocarp, distorted so that two of the three apical apertures are shown.
X 6-5. (¥.33968.)
FIG. 186. A broader endocarp, probably belonging to this species. x 6-5. (¥. 33969.)
Symplocos headonensis Chandler
FIG. 187. Endocarp somewhat distorted so as to show the apex with four apertures. X 6-5.
(V.33970.)
FIG. 1 88. Another, side. The apical depression is seen in profile. X 6-5. (V. 3 3971.)
All the above are from Bovey.
Solanispermum reniformis n. gen et sp.
FIG. 189. Perfect seed, x 6. Geol. Surv. Mus. Colin. Na. 76684.
FIGS. 190-191. Two fragments of one seed which broke on removal from the matrix.
X 15-5- (V-33972-)
Carpolithus spp.
FIG. 192. A carpel, side, showing one of the ventral facets. ¥entral angle on the right.
Note three-fid perianth at apex, x 15. (V. 33973.) Heathfield.
FIG. 193. Another, dorsal side, dorsi ventral compression shows part of one of the ventral
facets on the right, x 15. (¥.33974.) Heathfield.
FIG. 194. The same, ventral side, showing two facets and ventral angle. Persistent superior
perianth can be seen, x 15.
FIG. 195. Two lobes of a dehisced capsule, inner surface, x 8. (¥.33975.)
FIG. 196. The same, outer surface, x 8.
FIG. 197. Outer surface of a third detached lobe from the same specimen, x 8.
FIG. 198. Endocarp, incomplete at one end, broken so as to show one of the septa, x 6-5.
(¥.33976.) Heathfield.
FIG. 199. Another, broken at one end. x 6-5. ^.33977.)
FIG. 200. A three-lobed slender fruit, x 6-5. (¥.33978.)
FIG. 201. Another somewhat broader specimen, x 6-5. (¥.33979.)
FIG. 202. Another fruit with three minute patent perianth segments, x 6-5. (¥.33980.)
FIG. 203. A seed, x 15-5. (¥.33981.)
Bulbil ?
FIG. 204. Bud or Bulbil. The large hollow may be the burrow of an insect, x 2-8.
(^.33982.) Unless otherwise stated all the above are from Bovey.
Fig. 189 : Photo by C. Reid.
Bull B.M. (N.H.) Geol. 3, 3
PLATE 17
198
199 200
201 202
MASTIXIA, LYSIMACHIA, SYMPLOCOS, SOLANISPERMUM, CARPOLITHUS, BULBIL?
THE STRUCTURE OF SOME LEAVES
AND FRUCTIFICATIONS OF THE
GLOSSOPTERIS FLORA OF
TANGANYIKA
D. D. PANT
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 4
LONDON: 1958
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following papers appeared in Volume I (1949-52) :
Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.
H. D. Thomas & A. G. Davis . . 75. 6d.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu . . 55.
No. 3 (1950) . The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White ..... . js.6d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
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No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede
Euscalpellum. T. H. Withers . . 55.
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .
T. H. Withers . . 5*-
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian
of Podolia. W. N. Croft ..... ios.
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsdale .... ios.
No. 9 (1952). Australian Arthrodires. E. I. White 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. WThittard 6s.
The following papers appeared in Volume II (i953~56) :
No. i (1953). The Deer of the Weybourn Crag and Forest Bed of
Norfolk A. Azzaroli . . . • • • ^ 5s-
No. 2 (1953). A Coniferous Petrified Forest in Patagonia. Mary G.
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No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark . . 3-s. 6d.
No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.
Marjorie E. J. Chandler
No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans .
No. 6 (1955). Further Contributions to the Solution of the Piltdown
Probem. J. S. WTeiner, W. E. Le Gros Clark, K. P. Oakley
& others ..... &
No. 7 (1955). The Schizaeaceae of the South of England in Early
Tertiary Times. Marjorie E. J. Chandler i.5s~-
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . £i
THE STRUCTURE OF SOME LEAVES AND
FRUCTIFICATIONS OF THE GLOSSOPTERIS
FLORA OF TANGANYIKA
BY
DIVYA DARSHAN PANT
(Botany Department, The University, Allahabad, India)
Pp. 125-175 ; Plates 18-21 ; 21 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 4
LONDON: 1958
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 4 of the Geological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued January 1958 Price Twenty-one Shillings
THE STRUCTURE OF SOME LEAVES AND
FRUCTIFICATIONS OF THE GLOSSOPTERIS
FLORA OF TANGANYIKA
By DIVYA DARSHAN PANT
CONTENTS
Page
I. INTRODUCTION .......... 127
II. DESCRIPTION OF THE MATERIAL ....... 128
Genus Glossopteris Brongniart . . . . . . .128
G. fibrosa n. sp. ........ 130
G. hispida n. sp. ........ 134
G. colpodes n. sp. . . . . . . . 141
Glossopteris sp. A . . . . . . . .149
Genus Rhabdotaenia nov. ........ 149
R. danaeoides (Royle) n. comb. . . . . .151
R. harkini n. sp. ........ 154
Scale leaves .......... 159
Sporangia .......... 160
Specimens from Mhukuru . . . . . . .161
Specimens from Australia . . . . . . .163
Genus Spermatites Miner . . . . . . . .165
S. crystallinus n. sp. ....... 165
S. tetrapterus n. sp. . . . . . . . .169
III. SUMMARY ........... 172
IV. ACKNOWLEDGMENTS . . . . . . . . .173
V. REFERENCES . . . . . . . . . .173
SYNOPSIS
The epidermal and cuticular structure of three new species of Glossopteris, a species of Rhab-
dotaenia gen. nov. (Taeniopteris in part), some scale leaves, microsporangia and two new species
of seeds (Spermatites Miner) from the Mhukuru Coalfield, Tanganyika are described. Taeniopteris
danaeoides Royle from India is referred to Rhabdotaenia on the basis of its epidermal structure.
Two discs bearing microsporangia, from Newcastle, New South Wales, are described and their
sporangia compared with those from Africa.
I. INTRODUCTION
THE study of the cuticular structure of fossil plant compressions has rapidly advanced
but comparatively little work has yet been published on the structure of plants
of the Glossopteris flora. The cuticular structure of Glossopteris itself, the most
widespread genus of the flora, is but briefly described for only two leaves, G. indica
Schimper (Zeiller, 1896 : 369, fig. 13) and G. angustifolia Bgt. (Sahni, 1923 : 277,
GEOL. Ill, 4. II
128 THE GLOSSOPTERIS FLORA OF TANGANYIKA
pi. 17, figs. 2, 3). These cuticles are, however, so dissimilar that they have always
pointed to the need for more extensive work on the subject and the present paper
is an attempt in this direction.
The material described here was obtained from six borehole cores from the
Mhukuru Coalfield in Tanganyika. The cores have been described by Harkin
(1953) and in the same paper Professor J. Walton (Appendix IV : 28) gave a pre-
liminary report on their flora, a typical Glossopteris assemblage. Professor Walton
later handed the cores to me for detailed study. The work was begun in his laboratory
in Glasgow and completed at Professor Harris's laboratory in Reading.
The cores are only about 5 cm. in diameter, hence the specimens are small, but
the matrix is fine and is entirely unoxidized and the plants are beautifully preserved
compressions. An unexplained feature of the preservation is that while some leaves
are complete, as is ordinarily the case, others show only one epidermis with more
or less of the mesophyll adherent to it ; the veins may be present or they may be
missing and merely represented by the epidermal cells. It looks as though the
leaf had split through the middle before preservation and the two halves were
preserved separately. As a result the single epidermis is thin enough to be trans-
parent and it shows its cell outlines very clearly, often far more clearly than can
be seen in the cuticle. It is easily removed from the rock by celloidin pulls. It
also shows the lignine thickenings of the guard cells which are destroyed when the
cuticle is prepared.
The smaller fossils (scale leaves, microsporangia and seeds) described in this
paper owe their interest to the excellent preservation of their cuticles. One of the
small Vertebraria axes which show their tracheids and ray tissue has already been
described (Pant, 1956).
The other fossils occurring in the cores consist of a single fragment of a leaf sheath
of Schizoneura, some equisetaceous stems, a few megaspores, abundant two-winged
pollen grains, a few three-winged, one-winged and unwinged examples and numerous
slender roots showing scalariform metaxylem and spiral and annular protoxylem
with or without a surrounding sheath of fibres. These are not described here.
Walton (1953 : 30) had, in addition, recorded the occurrence of two specimens
of Noeggerathiopsis in these cores (a third is a counterpart) but after examining the
specimens and preparing their cuticles I think that they are poorly preserved midrib
regions of Glossopteris leaves.
According to Harkin (1953 : 9) the plant-bearing shales at Mhukuru belong to
the " Upper Coal Measures " and to the " upper part of K3 bed " regarded as corres-
ponding to the upper part of the Ecca Series in South Africa (see Harkin, 1953 : 7,
Table 2). Walton (1953 : 28) also regards the age as Ecca. His determinations
were, however, based on a preliminary identification of the fossils which are here
regarded as new species and therefore do not indicate a precise age.
Genus GLOSSOPTERIS Brongniart
The epidermis of the various species described here is of a single general type and
it may well be that all these species belong to one true genus. Glossopteris indica
THE GLOSSOPTERIS FLORA OF TANGANYIKA
B
129
FIG. i. A-E, Glossopteris fibrosa n. sp. ; F, G. hispida n. sp. A, details of venation of a
typical leaf showing fibres in certain meshes. ¥.34444. x 10. B, leaf apex with short
meshes. ¥.34444. x 2. c, leaf apex with long meshes ; a few anastomoses may
have been missed. ¥.34446. x 2. D, leaf base showing narrow meshes near the
margin. The midrib shows lumps but no bundles. ¥.34447. x 2. E, two leaf frag-
ments, on the left, middle region, on the right, near base ; some anastomoses may
have been missed. Holotype, ¥.344400;. x 2. F, middle region of leaf showing
veins, midrib and part of the margin. Holotype, ¥.34450^. x 2.
130 THE GLOSSOPTERIS FLORA OF TANGANYIKA
of Zeiller (1896) however, appears from the figure to be different in cuticle and may
belong to another genus ; (I was unable to find Zeiller's original slide in Paris). The
fructifications described by Zeiller (1902) as Ottokaria, and by Plumstead (1952,
1956) as Scutum and Lanceolatus also suggest generic differences but at present
I consider it premature to make new genera out of Glossopteris.
Glossopteris fibrosa n. sp.
(PL 18, figs. 1-5 ; PL 19, fig. i ; Text-figs, i, A-E ; 2, 3)
DIAGNOSIS. Leaf long lanceolate, average length estimated at between 13 and
30 cm.; width in middle region 4-6 cm. Basal part tapering very gradually,
margins becoming almost parallel to midrib at base, two sides of lamina often of
unequal width. Apex more or less obtuse. Petiole not known. Midrib up to
0-5 cm. wide below, persisting to apex but becoming narrow, probably depressed
above and convex below. Midrib sometimes showing small lumps. Margins
normally entire or slightly undulate, rarely lobed, slightly curved downwards and
thickened. Veins arising from midrib at a very acute angle in all parts of the leaf
but soon bending outwards for about i cm. and then continuing at an angle of 6o°-8o°
to the midrib except in basal and apical parts where veins less arched making
angles of 40°-5o° only with midrib. Veins crossing lamina at a concentration of
about 20-30 per cm. near midrib and 32-44 per cm. near margin (measured at right
angles to majority of veins). Veins anastomosing in all parts of lamina but more
frequently near midrib. Average width of meshes 0-5 mm. (ranging from 0-4 to
0-9 mm.) near midrib and 0-3 mm. near margin. Meshes in middle of lamina of
varied length, average length 7 mm.; extreme base of leaf with lamina only one or
two short meshes wide. Veins bending forwards at margin. Veins normally
90-160 11 thick (some up to 250//,), prominent on underside.
Substance of lamina rather thin, meshes showing elongated fibres running parallel
with main veins, about 5-14 /£ wide, fibres occasionally crossing connecting veins
or moving to join main veins ; also showing palisade-like cells 15-30 JJL wide and
spongy mesophyll cells elongated transversely to the veins averaging 60 /* long
and 20 ju, wide.
Upper epidermis of lamina usually without stomata. Cells between veins
averaging 86 ju, long and 44 fi wide tending to form rows parallel with veins. Lateral
(anticlinal) walls about 3 /* thick, arched or nearly straight, never sinuous. Surface
wall either without any papilla but appearing finely mottled or occasionally with
several small papillae with or without an obscure larger median papilla. Sometimes
numerous small papillae tend to be in longitudinal rows or are replaced by
longitudinal striations. Cells over veins and often above fibres somewhat narrower
and longer. In basal part of leaf and near margin upper epidermal cells become
isodiametric but elongated along the margin itself. Cells over midrib with thicker
walls (about 6 /i thick), elongated or short, rectangular or polygonal, tending to
occur in longitudinal rows, stomata present but rare. Midrib cells often with a
moderately conspicuous papilla. Trichomes absent.
Upper cuticle of lamina rather thick (up to 3 /* thick). Cell outlines thin, straight,
THE GLOSSOPTERIS FLORA OF TANGANYIKA
V-\
131
:•': .:.'k-<s.' ,9 '.
•••••:f- .<:.J
•mm&m
mfcimmm
FIG. 2. Glossopteris fibrosa n. sp. A, leaf base showing slight asymmetry at the top,
venation mentioned on p. 130. "^34445. x i. B, upper cuticle of lamina showing cells
arranged in longitudinal rows parallel with the veins. V. 34448. X 125. c, cell of
upper epidermis over vein showing longitudinal striations with small papillae arranged
along them at some places. ¥.344490. x 800. D, epidermis (? upper) of midrib
showing stomata and thick-walled cells with median papillae. V.34444«. x 125.
E, a cell of upper epidermis showing papillae of various sizes. ¥.344490. x 800. F,
lower cuticle showing stoma with outlines of guard cells distinctly marked at the poles
but less distinct at the sides (see also PI. 18, fig. 4). ¥.34443. x 800.
132 THE GLOSSOPTERIS FLORA OF TANGANYIKA
not bordered ; surface of cells finely mottled, sometimes with obscure median
papilla or several small papillae ; otherwise as epidermis.
Lower epidermis of lamina showing isodiametric polygonal cells or cells somewhat
elongated in various directions between veins, but longitudinally elongated cells
over veins. Average width of cells about 44 /*. Lateral walls nearly straight or
curved, never sinuous, sometimes unevenly thickened. Surface wall finely mottled,
often showing a single ill-defined more or less prominent median papilla. Cells
over midrib as on upper side but stomata absent. Trichomes absent.
Stomata frequent in vein meshes, rarely over veins ; concentration typically
about 125 per sq. mm. Orientation inconstant, tending to be longitudinal near
veins and transverse between them.
Stomata haplocheilic, partly or completely amphicyclic but sometimes monocyclic.
Guard cells about 45 /t long and 25 fi wide ; sunken, usually aperture alone exposed
in a small pit about 30 fi deep. Subsidiary cells 4-8, forming a very irregular ring ;
often slightly smaller than ordinary epidermal cells, polar cells unspecialized ;
surface usually somewhat thicker than that of other epidermal cells ; usually
showing a rather prominent thickened papilla or with a thickened rim on the inner
side, papilla more or less hollow. Papillae pointing over the stomatal pit or pointing
outwards ; often a second faint papilla occurs in the middle of the cell. Encircling
cells unspecialized, sometimes tangentially elongated.
Lower cuticle rather thinner than upper (up to about 2 fi thick), anticlinal cell
walls thin without uneven thickenings. Cell surface finely mottled, usually with a
faint median papilla. Guard cells slightly thickened round aperture, outlines
clearly marked at poles, less clear at the sides. Lower cuticle otherwise like lower
epidermis.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.34440 (Text-fig. I, E).
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. G. fibrosa is by far the commonest of all Glossopteris remains in
the borehole cores. There are in all 146 fragments of different parts of its leaves
(including 16 apices and 15 bases). The diameter of the cores being about 5 cm.
no complete leaves can be seen.
In about half the specimens the midrib forms a fairly deep groove and the veins
are distinctly sunken. In the others the midrib is only very slightly raised and the
veins flat. I regard the first set as leaves preserved with the lower surface downwards,
the second set, preserved upside down (Walton, 1936). I conclude that the midrib
and veins were very prominent on the lower side but on the upper side the midrib
was slightly sunken and the veins flat. The margin also is slightly curved downwards.
The veins and midrib contain abundant scalariform tracheids from 14-26 ju, wide.
Tracheids with uniseriate or multiseriate bordered pits also occur in the midrib ;
some pits show crossed apertures (see Text-figs. 3, B, c).
In epidermal pulls, the guard cells of the more exposed stomata show lateral and
polar lignine lamellae of Gymnosperm type but these dissolve on maceration.
There is great variety in the surface of the upper epidermal cells of specimens
referred to G. fibrosa. A very common state is a finely mottled wall but other speci-
THE GLOSSOPTERIS FLORA OF TANGANYIKA
133
FIG. 3. Glossopteris fibrosa n. sp. A, lower epidermis showing a stomatal area between
two veins. ¥.34440^. x 125. B, tracheids of midrib showing bordered pits ; some
of them show crossed apertures. "^344400!. x 400. c, scalariform tracheids from a
vein. V. 34440^. x 800. D, stoma at a fold of the lower epidermis, seen from inner
side (see also PL 18, fig. 2). V. 34441. X 400. E, part of A, more magnified, showing
more and less exposed stomata. x 400. F, a stoma with four subsidiary cells and lateral
and polar lignine lamellae in guard cells. V. 34440^. x 400.
134 THE GLOSSOPTERIS FLORA OF TANGANYIKA
mens show a faint papilla and others several moderately developed small papillae,
a state which grades into mottling and also grades into longitudinal striations.
One leaf showed striations in cells over a vein and close to them cells with several
small papillae and one faint larger papilla. In some leaves the lower cuticle is
thin and looks much like that of G. hispida (details are however obscure), but differs
in the absence of hair bases.
The only additional characters to be noted are those seen in exceptional specimens
and which may be the result of peculiar preservation. The upper epidermis of a
leaf shows pale cell walls and dark cell contents, the cells being full of granules of
various sizes (PL 18, fig. 3).
It was noted that two- winged pollen grains, more commonly varying in size
from 40-60 ft across, occur abundantly on both sides of the leaves.
COMPARISON. G. fibrosa looks much like many of the figures given by various
authors (e.g. Brongniart, 1828 ; Dana, 1849 ; Feistmantel, 1878-79, 1879, 1880-81,
1882, 1886 ; Zeiller, 1896, 1902 ; Arber, 1905^ ; Walkom, 1922, 1928 ; Walton, 1929 ;
Plumstead, 1952, 1956) under the names G. browniana var. indica (G. indica), G.
angustifolia, G. browniana var. australasica (G. browniana) and G. communis. I have
seen the holotypes of the first two species and a syntype of G. browniana var.
australasica Brong. (No. 507) in the Natural History Museum in Paris. All are
clearly different because they lack fibres in vein meshes. None of these has satis-
factorily preserved cuticle but there is a little information about G. browniana
(see Table I). I have examined much other material attributed to these species in
the British Museum (Nat. Hist.), ficole de Mines, Paris and in Oxford University
Museum ; the only specimens which appear to have fibres are those described by
Bunbury (1861) as G. browniana var. indica (¥.19617) and by Walton (1929, pi.
c, fig. 19) as G. indica (¥.20778) both in the British Museum (Nat. Hist.). Two
slides showing fragments of Glossopteris leaves from Richmond Vale, Australia,
sent to the British Museum by Dr. A. B. Walkom, show similar fibres and venation.
A cuticle was prepared but cell outlines are very obscure. The only other Glossopteris
with similar fibres is G. hispida described in this paper (see p. 140 for comparison).
Glossopteris cuticles have been described from specimens identified as G. indica
by Zeiller (1896) and G. angustifolia by Sahni (1923). I assume that these determina-
tions are correct. G. fibrosa differs clearly in cuticle from both (see Table I). I
have examined a few other cuticles of Glossopteris leaves with this kind of venation
and all are different.
Glossopteris hispida n. sp.
(PL 18, figs. 6, 7 ; PL 19, fig. 3 ; PL 20, figs, i, 2, 7 ; Text-figs, i, F ; 4-6)
DIAGNOSIS. Leaf 4-3-7 cm wide, elongated, length unknown. Midrib up to
at least 0-5 cm. wide, showing numerous longitudinal strands anastomosing at
long intervals. Lateral veins departing from midrib at an acute angle (io°-20°)
but soon arching outwards and continuing at an angle of 6o°-8o° to midrib ; concen-
tration of veins 19-26 per cm. near midrib, 28-35 near margin (measured transversely
to the majority of veins) ; meshes shorter near midrib, longer towards margin but
THE GLOSSOPTERIS FLORA OF TANGANYIKA
135
TABLE I. — Comparison of Certain Glossopteris Leaves
with Narrow Vein Meshes and the Present New Species
Fibres
between
Material
Thickness
of cell
walls
Upper epidermis
or cuticle
Lower epidermis
or cuticle
i. With type G. browniana Absent
var. australasica Bgt.
Mus. Hist. Nat. Paris,
no. 507.
Loc. Hawkesbury River,
nr. Port Jackson,
N.S.W.
Thin
Lateral walls straight,
surface with several
small papillae
2. ? With type G. brown-
iana var. australasica
Bgt. Oxford Univ.
Mus., no. FW5.
Loc. " New Holland ".
(The type cannot be
traced)
3. Glossopteris sp. No.
¥.19164.
Loc. Burdwan, Bengal,
India
4. G. angustifolia (Sahni,
1923).
Loc. Raniganj, Bengal,
India
No
data
5. Glossopteris sp. Oxford Absent
Univ. Mus., no. FW8.
Loc. Newcastle, N.S.W.
6. Glossopteris sp. No. Present
V.34492.
Loc. Richmond Vale,
N.S.W.
Ditto
Lateral walls straight
to wavy, surface with
striations or several
small papillae (never
single)
Lateral walls wavy,
surface with " nu-
merous extremely
fine punctuations
closely arranged
along wavy parallel
lines "
Lateral walls straight,
surface with single
median papilla
Details and cell out-
lines very obscure in
cuticle
Lateral walls sinuous,
ordinary cells some-
times with median
papillae, subsidiary
cells with papillae
overhanging stoma-
tal pit. Trichome
bases absent.
Lateral walls sinuous,
ordinary cells some-
times with median
papillae, subsidiary
cells ?, trichome
bases absent.
Lateral walls slightly
wavy or almost
straight, ordinary
cells without median
papillae, subsidiary
cells with papillae
overhanging stoma-
tal pit. Trichome
bases absent.
Lateral walls almost
straight, ordinary
cells without papil-
lae, some subsidiary
cells with median
papillae. Trichomes?
(probably absent).
Lateral walls slightly
wavy, ordinary cells
with median papil-
lae, subsidiary cells
with papillae over-
hanging stomatalpit.
Trichomes absent.
Details and cell out-
lines very obscure in
cuticle.
136
THE GLOSSOPTERIS FLORA OF TANGANYIKA
Table I. — continued
Material
Fibres Thickness
between of cell
veins walls
7. Glossopteris sp. A. No. Absent . Thin
¥.34467.
Loc. Mhukuru Coalfield,
Tanganyika (2 speci-
mens)
8. G. fibrosa . . . Present
Loc. Mhukuru Coalfield,
Tanganyika (many
specimens)
9. G. hispida .
Loc. Mhukuru Coalfield,
Tanganyika (4 speci-
mens and some small
fragments)
10. G. indica (Zeiller, 1896).
Loc. Frances, Johannes-
burg, S. Africa
Upper epidermis
or cuticle
Lateral walls straight
or arched, never
wavy, bordered, sur-
face with a median
papilla or mottled
Lateral walls straight
or arched, never
wavy, not bordered,
surface mottled or
with single median
papilla or numerous
papillae or with
longitudinal stria-
tions
Lateral walls straight,
never wavy, surface
with numerous papil-
lae or mottled
No
data
. G. colpodes . . Absent
Loc. Mhukuru Coalfield,
Tanganyika (many
specimens)
Thick . No data
Thin . Lateral walls usually
wavy, straight near
midrib and margins
and over midrib
and veins, surface of
most cells with a
median papilla
Lower epidermis
or cuticle
Lateral walls almost
straight, ordinary
cells usually with
single median papil-
lae, subsidiary cells
with papillae over-
hanging stomatal pit.
Trichomes absent.
Ditto
Lateral walls straight
to sinuous, ordinary
cells without median
papillae, subsidiary
cells with papillae
overhanging stoma-
tal pit. Trichomes
and trichome bases
present.
Lateral walls straight,
ordinary cells and
subsidiary cells
without papillae.
Trichomes ?
Lateral walls marked-
ly wavy but straight
near midrib and mar-
gins and over midrib
and veins, surface
often with a median
papilla. Trichomes
absent.
often the ultimate mesh is again short. Meshes average about 0-8 mm. wide
(ranging from about o-6-i mm.) near midrib and about 0-3 mm. wide near margin,
average length of meshes about 4 mm. Veins and midrib prominent on the lower
side, midrib also slightly depressed on the upper. Veins up to 140 p thick. Margin
entire, slightly thickened and slightly curved downwards, occasionally lobed.
Substance of lamina thin, showing spongy mesophyll cells elongated transversely
THE GLOSSOPTERIS FLORA OF TANGANYIKA
137
v-v .... •
D M
FIG. 4. Glossopteris hispida n. sp. A, lower cuticle showing hair base. V. 344510.
X 400. B, lower epidermis showing three-celled hair base. V.3445oa. x 400.
c, D. fragments of lower cuticle showing stomata. V. 34452. x 400. E, middle
part of leaf showing venation and lobed margin. ¥.34451. x 2. F, upper cuticle
showing elongated cells in rows. ¥.344516. x 125. G, a cell from F further magnified
to show its numerous papillae (the small dots represent the substance of the cuticle).
¥.344516. x 800.
138
THE GLOSSOPTERIS FLORA OF TANGANYIKA
B
FIG. 5. Glossopteris hispida n. sp. A, hairs from matrix in contact with lower side of
leaf. B, lower epidermis of midrib showing trichome bases, c, lower epidermis of
stomatiferous area between two veins near midrib showing straight-walled cells,
stomata and hair bases. The rows of elongated cells in the middle are above a fibre.
D, lower epidermis of a mesh away from midrib showing sinuous-walled cells. The
elongated cells on the right are above a fibre (see Text-fig. 6 for details of stomata and
cells). All from Holotype (¥.344500). x 125.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 139
to the veins (about 39-52 fi long X 13-24 /* wide), and palisade mesophyll cells
(about 18-30 fji wide) also showing a few isolated fibres running parallel to veins
in vein meshes.
Upper cuticle of lamina about 2 fi thick, showing rectangular or polygonal cells.
Cells short or frequently elongated in the direction of veins and tending to be in
rows parallel to veins, lateral and end walls straight, thin, often obscure. Surface
wall with numerous small papillae which tend to be in longitudinal rows, sometimes
cell surface irregularly mottled. Cells typically about 30 /* x 57 /*.
Upper cuticle of midrib showing rectangular cells, short or longitudinally elongated,
tending to be in longitudinal rows. Surface mottled, papillae absent. Stomata
and trichomes absent on the upper side.
Lower epidermis of lamina showing frequent trichomes and trichome bases both
along veins and between them, stomata present in areas between veins. Cells in
meshes near midrib almost straight walled, polygonal, isodiametric or elongated
in various directions ; cells in meshes away from midrib irregularly shaped, often
more or less elongated in the direction of the veins, cells averaging about 46 ju, wide X
62 11 long ; lateral (anticlinal) walls slightly to markedly sinuous. Cells above
veins and sometimes above fibres in meshes, straight walled, narrow and elongated
in the direction of the veins. Lateral walls of lower epidermal cells about 3 fi
thick, sometimes unequally thickened. Surface of cells very finely mottled, usually
without any papillae.
Lower epidermis of midrib with longitudinally elongated or short, polygonal or
rectangular cells, lateral walls straight, about 6 /* thick, surface wall finely mottled,
trichomes and trichome bases present, stomata absent.
Trichome bases composed of a single oval or rounded cell, or two or three cells,
commonly overlapping a number of ordinary epidermal cells but occasionally
overlying a single cell. Trichomes simple, three- to six-celled, tapering, cells short
or long, apex of end cell acutely pointed. Trichomes pointing in different directions
but usually outwards and backwards on the midrib and backwards on the lamina.
Concentration of stomata about 70 per sq. mm., orientation irregular. Stomata
haplocheilic, monocyclic. Guard cells averaging 49 fi long x 21 /* wide, partly
overhung by subsidiary cells and their papillae or occasionally exposed. Subsidiary
cells 4-6, irregular or forming an irregular ring, like ordinary epidermal cells in shape
and size but usually with a prominent thick-walled hollow papilla pointing over the
stomatal pit or with a thickened rim towards stomatal aperture. Polar subsidiary
cells like lateral ones.
Lower cuticle delicate (about i /* thick), cell walls often obscure, thin and straight ;
surface wall smooth or granular, papillae absent in ordinary epidermal cells but
present in subsidiary cells. Lower cuticle otherwise like lower epidermis.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. V. 34450.
LOCALITY AND HORIZON. Mhukura Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. The material consists of four good specimens and a few small
fragments. In two of the leaves the lamina is torn (see Text-figs, i, F and 4, E).
The veins and the midrib show abundant scalariform tracheids like those of G.
I40
THE GLOSSOPTERIS FLORA OF TANGANYIKA
fibrosa. In the epidermal pulls the guard cells show lateral and polar lignine
lamellae. These dissolve on maceration. Some cells of the lower epidermis show
a dark inner area, slightly smaller than their lumen, which looks like their contracted
contents. Dark spots, resembling median papillae, are also seen in a few cells of
the lower epidermis. The spot is often placed near one end of the cell. No
corresponding structures were observed in the lower cuticle and these spots may be
extraneous particles.
FIG. 6. Glossopteris hispida n. sp. Part of Text-fig. 5, D further enlarged to show details
of stomata and two-celled hair base (see also PI. 20, fig. 2). x 400.
Two-winged pollen grains of varying sizes are frequently seen overlying the cuticle.
COMPARISON. G. hispida agrees with G. fibrosa in venation and in the fibres
between the veins. Though the vein meshes are usually of similar size, no specimens
of G. hispida are known with meshes as narrow as in some of G. fibrosa.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 141
The conspicous difference is in the hairs on the underside of G. hispida (none in
G. fibrosa) . These are seen clearly when the matrix in contact with the underside
of the leaf is moistened with oil ; their bases are also visible in the epidermis and the
cuticle. In G. fibrosa the cuticle is rather thicker and usually with clearly marked
cell outlines, the cuticle of G. hispida is thin and the cell outlines obscure. The cells
of the lower epidermis in G. hispida have sinuous walls (except over veins and midrib
and in meshes near midrib), in G. fibrosa they are always almost straight. In G. fibrosa
the stomata are partly amphicyclic, in G. hispida monocyclic. There may be other
differences between the two species but their constancy is unknown.
Glossopteris leaves with rather similar veins but whose fine details are unknown
have been described under such names as G. ampla, G. musaefolia, G. indica, G.
browniana, G. damudica, and others. I am unable to distinguish any difference
between them. A celloidin pull from one of the original specimens of G. ampla
Dana sent to me by Dr. S. H. Mamay did at least differ in possessing no fibres. I
also examined the types of G. indica Goppert (G. browniana, var. indica, Brongniart,
1828, pi. 62, fig. 2) consisting of two separate pieces (No. 506) in the Paris Museum
of Natural History ; G. musaefolia Bunbury (¥.19621 and also syntype material),
and a specimen attributed to G. damudica var. stenoneura (V. 19577) by Feistmantel
(1889, pi. 4, fig. 7) in the Palaeontological Department of the British Museum
(Natural History). They all differ from G. hispida in lacking impressions of fibres
in vein meshes (the original leaf substance is not preserved).
Glossopteris colpodes n. sp.
(PI. 19, fig. 2 ; Text-figs. 7-9)
DIAGNOSIS. Leaf elongated (length unknown), average width in middle region
about 6 cm. (extremes 4*5-7-5 cm.) basal part tapering with margins approaching
midrib at an angle of 10°; apex more obtuse, margins approaching midrib at an
angle of 30°. Midrib up to 0-4 cm. wide below, showing numerous longitudinal
strands and often small lumps in lower parts, fewer strands and lumps above. Leaf
margin entire, undulating or lobed, slightly thickened.
Veins arising from midrib at a small angle but soon bending outwards for about
i cm. and then continuing at an angle of 45°-65° to midrib (in extreme specimen
75°-8o°). Concentration of veins (measured at right angles to majority of veins)
11-15 Per cm- near midrib in lower and middle parts of leaf, increasing to about
20 per cm. near midrib in upper part. Near margins veins more crowded, about
30 per cm. (maximum about 36 per cm.) . Veins usually up to 140 /* thick, prominent.
Veins and midrib raised on the lower side of leaf and midrib also slightly depressed
on the upper. Meshes average 4 mm. long (extremes 2-8 mm.) and 0-9 mm. wide
(extremes 0-5-1-5 mm.) in lower and middle parts of leaf, longer and narrower
in upper part, averaging 7 mm. x 0-5 mm.; ultimate meshes near margin often
short and narrow. Veins usually bending slightly forwards at margin.
Upper epidermis of lamina without stomata or trichomes. Cells over veins
GEOL. Ill, 4. 12
I42
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 7. Glossopteris colpodes n. sp. A, lobed apex with narrow elongate meshes.
¥.34462. X 2. B, apical part of leaf. V.34463. X 2. c, details of venation at
margin of a leaf. V.34465. x 10. D, basal part of a leaf. ¥.34464. x 2. E,
middle portion of leaf . Holotype, ¥.34461. x 2.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 143
narrow, elongated, straight-walled. Cells between veins irregularly shaped or
polygonal, short or elongated in various directions, averaging 66 /* long and 40 /*
wide. Lateral (anticlinal) walls of cells about 3 ji thick, slightly to markedly sinuous,
sometimes appearing nearly straight especially near midrib and margins. Cells
at margin longitudinally elongated, nearly straight-walled. Surface of upper
epidermal cells always mottled, most cells showing a median papilla about 6 /* wide.
Cells over midrib in longitudinal rows, rectangular, short or elongated, lateral
walls straight, papilla sometimes seen, trichomes absent.
Upper cuticle moderately thick (about 3 fi), cell outlines between veins obscurely
marked, often appearing discontinuous but scarcely showing sinuous waves. Cell
walls clearer over midrib and margins, straight or nearly straight but always appear-
ing closely beaded. Papillae sometimes visible.
Lower epidermis of lamina showing narrow straight-walled cells above veins.
Cells between veins elongated in various directions or isodiametric, averaging
69 /* X 38 fi, often irregularly shaped, lateral walls more or less sinuous, often
unevenly thickened, but nearly straight near midrib and margins. Walls about
3 p thick. Surface of cells always mottled and often showing a median papilla.
Trichomes absent.
Stomata frequent between veins, often unevenly distributed, concentration
about 66 per sq. mm., orientation varied. Stomata haplocheilic, monocyclic,
occasionally incompletely amphicyclic. Guard cells about 65 fi X 25 /i, sunken,
frequently entirely covered by 4-8 subsidiary cells and their papillae, except some-
times from a region near aperture. Subsidiary cells irregular or forming an irregular
ring, polar cells resembling lateral ones, size of subsidiary cells almost as large as
that of other epidermal cells, but surface sometimes thicker ; each subsidiary cell
usually with a large hollow but thick-walled papilla pointing over the stomatal
pit (papillae often overlapping and completely concealing the stomatal aperture)
or papilla pointing upwards. Sometimes a second median papilla also present in
subsidiary cells. Encircling cells unspecialized but often tangentially elongated.
Lower cuticle of lamina thin (1-2 /* thick), cell walls between veins faintly marked,
usually interrupted and appearing nearly straight but never showing well-marked
waves, occasionally walls almost straight. Cell walls over veins more distinct,
straight, almost continuous but closely beaded. Papilla and surface mottling
often visible. Surface occasionally showing radiating striations extending over
several cells. Guard cell outlines clearly marked at the poles but faint on the
sides.
Lower cuticle of midrib with short or long polygonal cells arranged in longi-
tudinal rows. Lateral walls straight, surface mottled. Papillae often absent.
Stomata and trichomes absent.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.34461.
LOCALITY AND HORIZON. Mhukura Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. Sixty-seven fragments were attributed to G. colpodes. The
difference between the venation in the apical and the basal regions is considerable.
These were identified with one another because of their lack of fibres between veins
144
THE GLOSSOPTERIS FLORA OF TANGANKIYA
FIG. 8. Glossopteris colpodes n. sp. A, two protected stomata. ¥.344660. X 400.
B, lower epidermis of stomatiferous area between two veins. ¥.344666. X 125. c,
lower epidermis. ¥.344666. X 125. D, relatively exposed stoma from B. X 125.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 145
and also because of their agreement in epidermal or cuticular structure. Fourteen
specimens show epidermal cells of the upper or the lower sides and these have
sinuous walls, except above the veins and the midrib and in meshes near the midrib
and the margins. The other specimens were too thick to show the epidermal cells
but they all gave cuticles which showed the characteristic cells with very obscurely
marked walls which appear more or less straight but discontinuous (see Text-figs.
9, A ; 9, D).
Several epidermal specimens show structures not mentioned in the diagnosis.
When sufficiently exposed, the guard cells show lateral and polar lignine lamellae
of the Gymnosperm type (these are unrepresented in the cuticle) . Some pulls show
circles abut 30 ju, wide representing flattened palisade cells ; others show less distinct
spongy mesophyll cells often elongated transversely to the veins. At a few points
in the pulls the veins and the midrib show scalariform tracheids.
The only specimen showing the apex was unfortunately rubbed and damaged.
The lamina is lobed (Text-fig. 7, A) . There is nothing to show whether this is normal.
Two-winged pollen grains of various sizes ranging between 40-100 ju, across are
frequently found sticking to the epidermis or cuticle of either side.
COMPARISON. G. colpodes is distinguished from G. fibrosa and G. hispida by the
lack of fibres between the veins and in the lower parts by its wider meshes. The
epidermal cells in G. fibrosa are never sinuous walled (markedly sinuous in G. colpodes
except above the veins and the midrib and in the meshes near the midrib and the
margins). Glossopteris sp. A may have veins as wide as those in G. colpodes and also
lacks fibres in meshes but differs in the complete absence of sinuous-walled cells.
The lower epidermis of G. hispida resembles that of G. colpodes in having sinuous-
walled cells but differs in having trichomes (in G. colpodes trichomes are absent).
Among Glossopteris leaves of comparable venation are : Dana's Australian leaves
of G. reticulum and G. elongata (Dana 1849), ? Dictyopteris simplex Tate (G. Tatei
Feistmantel, 1889) from S. Africa (Tate, 1867), Feistmantel's Indian leaves of
G. retifera and G. conspicua (Feistmantel, 1880 ; 1881 ; 1886) and various other
specimens referred to them by Feistmantel and others, G. brancai Gothan (1914),
from Portuguese East Africa, and some specimens assigned to G. browniana by
Arber, (19050), Walton (1929) and others. Most of them differ from G. colpodes
in the size of meshes and their fine details are unknown (see table II) . However,
a specimen assigned to G. browniana by Arber (19050, : 56, pi. 3, fig. 2) and two
others figured by Walton (1929 : 70, pi. c, figs. 21, 22) as G. cf. browniana and
G. retifera respectively are very similar to G. colpodes in their venation but like
the rest their epidermal or cuticular structure is unknown.
I have also been able to examine some wide meshed undescribed Indian leaves at
the Oxford University Museum and an Australian specimen at the British Museum.
These yielded pulls showing epidermal cells and stomata very similar to those of
G. colpodes. The Indian leaves have veins at about the same concentration as in
G. colpodes (about 10-12 veins per cm. near midrib) and may belong to the same
species. The meshes of the Australian leaf are much wider (up to 2-5 mm. wide,
concentration of veins about 8 per cm. in lower part) and are therefore outside the
range observed in the present material of G. colpodes.
146
THE GLOSSOPTERIS FLORA OF TANGANYIKA
B
3|
V; ''^^^K^hSj^S^^
• •-' r^tjfa - ; VtfVAj':! '/• : : • •iiM&LMiftajL.V.- V '••".•"••."- .
FIG. 9. Glossopteris colpodes n. sp. A, upper cuticle as seen under phase contrast
microscope. V.34466c. X 400. B, upper epidermis showing sinuous-walled cells
between veins and straight-walled cells above a vein. ¥.34466^. x 125. c, epi-
dermis over midrib. V.34466d. x 125. D, lower cuticle showing a stoma as seen
under phase contrast microscope. ¥.344660. x 800. E, lower epidermis showing
a stoma with a ring of eight subsidiary cells and a few encircling cells. V. 34466(2.
X 400.
THE GLOSSOPTERIS FLORA OF TANGANYIKA
TABLE II. — Comparison of Certain Glossopteris Leaves with
Wide Vein Meshes
Angle of
veins to
midrib
Concentration of Character of
Material veins per cm. meshes
Walls of
epi-
after dermal
i cm. cells
i.G. colpodes
Loc. Mhukuru Coalfield,
Tanganyika, (many
specimens)
2. Glossopteris sp. A.
Loc. Mhukuru Coalfield,
Tanganyika (2 speci-
mens)
3. Glossopteris sp. Nos.
FY 14, 16-21, Oxford
University Museum.
Loc. Raniganj Coalfield,
Burdwan, India (15
specimens)
4. Glossopteris sp. No.
V.344QI
Loc. Newcastle, N ^.W.
(i specimen)
5. G. btowniana (Arber,
19050 : 56, pi. 3, fig. 2.
No. ¥.7207.
Loc. Port Stephens,
N.S.W.
6. G. cf. browniana (Wai- Near midrib up to
Near midrib 11-15
in middle and lower
parts ; up to 20 to-
wards apex ; near
margin up to 36
Near midrib 27-30 ;
near margins up to
36 (about 20 in
second specimen)
Near midrib 10-12
(veins ill-preserved
near margins)
Near midrib about
9 ; near margin
about 15 in lower
parts of leaf
Near midrib 14-16
up to 21 near mar-
gin (only middle
part known)
ton, 1929 : 70, pi. c,
fig. 21). No. ¥.20780.
Loc. Wankie Coalfield,
S. Rhodesia
15 ; up to 20 near
margin in lower
and middle parts.
Upper part of leaf
unknown
7. G. retifera. (Walton, Near midrib 11-16,
1929 : 70, pi. c, fig. about 20 near mar-
22). No. ¥.20781. gin (only basal part
Loc. Wankie Coalfield, of leaf known)
S. Rhodesia
8. G. retifera (Feistmantel, 9-14 all over lamina
1880 : 103, pi. 28A, (from figs.)
figs. 2, 7, 10 ; pi. 4iA.
1886, pi. 4A, fig. i).
Loc. Raniganj Coalfield,
Ramkola Coalfield,
Nakori River, nr. Gai
Nagar, Ganespur
River, India
Short and wide near . 45°-65° . Sinuous,
midrib and base,
elongated towards
apex and margin
Elongated but often . 4O°-5o° . Straight,
shorter near midrib
and margins
Shorter and wider . 6o°-7o° . Sinuous,
near midrib, elon-
gated elsewhere
Ditto . 8o°-85c
Ditto . 6o°-70° Unknown.
Short and wide near . 5o°-6o°
midrib in upper
part, elongated
elsewhere ; almost
uniformly short to-
wards base
Almost uniformly . 5o°-6o°
short and wide
Ditto
. 5o°-6oc
148
THE GLOSSOPTERIS FLORA OF TANGANYIKA
Table II. — continued
Material
9. G.retifera (Plumstead,
1952 : 300, pi. 49, fig.
5, pi. 51, figs. 1-6,
text-fig. 6.
Loc. Leeukuil, Vereeni-
ging, Transvaal, S.
Africa
10. ? Dictyopteris simplex
(Tate, 1867 : 141, pi. 6,
fig. 6 — G. tatei Feist-
mantel, 1889 : 44, pi.
4, fig. 8). No. V. 19579.
Loc. Bloemkop, S.
Africa
11. G. brancai (Gothan,
1914 : 13, pi. i, figs. 2,
3).
Loc. Tete, Portuguese
East Africa
Concentration of
veins per cm.
4 near midrib, 21
near margin in upper
part (from descrip-
tion) but 9-12 near
midrib, 21 near
margin, in Plum-
stead, 1956, pi. 12,
fig. 2
7-10 near midrib,
about 12 approx. 2
cm. from midrib.
Margin not pre-
served
4-6 near midrib, 10-
12 near margin in
upper and middle
part (from figs.)
14. G. reticulum (Dana,
1849 : 717, pi. 13, fig.
2)-
Loc. Newcastle, N.SW.
6 near midrib in
middle part, 12 near
midrib towards apex
and near margin.
Only upper part of
leaf known (from
description and fig.)
15. G. elongata (Dana, 7-11 below. Only
1849 : 718, pi. 13, fig. basal part of leaf
4). known (from des-
Loc. Newcastle, N.S.W. cription and fig.)
Character of
meshes
Short and wide near
midrib and base,
elongated in upper
part
Angle of
veins to
midrib
after
i cm.
Walls of
epi-
dermal
cells
"Almost .Unknown
90°"
Short and wide near . 7o°-8o°
midrib, elongated,
elsewhere.
Ditto
12. G. conspicua (Arber, 9-16 near midrib in .
igosa : 87, pi. 3, fig. various leaves in
3). No. ¥.2465. the slab
Loc. Mill River Drift,
Orange River Colony.
13. G. conspicua (Feist- 7-9 all over lamina .
mantel, 1881 : 104, pi. (from figs.)
28A. figs, i, 5, 6, 8,9).
Loc. Raniganj , Karan-
pura, and Auranga
Coalfields, India.
Ditto
Elongated
. 50°-6o°
. " About
65°"
Almost uniformly .
short and wide or
more
THE GLOSSOPTERIS FLORA OF TANGANYIKA 149
Glossopteris sp. A
(Text-fig. 10)
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
Only two specimens were recognized. The largest fragment (¥.34467) is shown
in Text-fig. 10, E, where all the details clearly visible are accurately represented.
The second specimen (¥.34493), a fragment with neither midrib nor margin has
slightly wider meshes, up to I mm. wide. The veins are sinuous but since the epi-
dermal cells show straight veins I have no doubt that the veins have been displaced
during preservation.
In both specimens the substance of the lamina is thin. In neither are there any
fibres in the vein meshes. The cuticle of both leaves was prepared but the figures
are all from the leaf shown in Text-fig. 10, E.
The upper cuticle of the lamina is about 3 fi thick. The cell outlines are clearly
marked as shown in Text-fig. 10, c. Lateral walls of cells are thin but have a border
on either side (Text-fig. 10, D). The cells tend to be in rows parallel to the veins.
The cells above the veins could not be seen in this specimen but in the second specimen
they are narrower and more elongated. Stomata are absent.
The lower cuticle is thinner (about 1-2 fi thick). The cell outlines are moderately
clear (Text-fig. 10, A). Lateral walls of cells are straight. The cell surface usually
shows a median papilla. The cells above the veins are narrow and elongated in
the direction of the veins, those between the veins are usually isodiametric or
elongated in various directions. Stomata are confined to areas between the veins.
Trichomes are absent.
The stomata are haplocheilic and possibly partly amphicyclic. The guard cells
are hidden by a ring of five or more subsidiary cells. Each subsidiary cell usually
has a prominent thickened papilla covering the stomatal aperture. The cells regarded
as encircling cells resemble ordinary epidermal cells but are often tangentially
elongated.
The cuticle in the second specimen is very similar but the papillae are very obscure.
Glossopteris sp. A resembles G. fibrosa in form, venation and cuticle. The only
difference is in the isolated fibres which are here absent but conspicuous in G. fibrosa.
It differs from G. hispida in lacking fibres, hairs and sinuous-walled cells. In the
complete absence of sinuous- walled cells it differs also from G. colpodes.
It has not been named as a species because too little is known about it.
Genus RHABDOTAENIA nov.
DIAGNOSIS. Leaf elongated, entire, with a strong midrib, lamina arising from the
sides of the midrib, lateral veins arising at a wide angle or at a smaller angle but
almost immediately bending outwards and crossing the lamina at a wide angle,
lateral veins occasionally forked, anastomoses between lateral veins extremely
rare. Stomata usually confined to areas between veins on lower epidermis, subsidiary
150
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 10. Glossopteris sp.A. A, lower cuticle showing elongated cells above veins and
stomatiferous areas between them. ¥.344670. X 125. B, stoma from A. X 400.
c, upper cuticle showing cells in rows parallel to veins. ¥.344670. X 125. D, part
of c more magnified to show bordered walls. X 400. E, middle part of leaf showing
venation. ¥.34467. X 2.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 151
cells haplocheilic, irregular, not forming a definite ring, guard cells exposed or covered
by subsidiary cell papillae not situated in a pit. Cuticle rather thin.
TYPE SPECIES. Rhabdotaenia danaeoides (Royle) n. comb.
DISCUSSION AND COMPARISON. Rhabdotaenia is made for certain fronds of
taeniopterid form of which the cuticular structure is known. Among such fronds
with haplocheilic stomata are Bjuvea Florin (1933 : 48) in which the subsidiary
cells are in a rather definite ring and form a pit above the guard cells, and Dorato-
phyllum Harris (1932 : 36) where they form a perfect ring arching over the guard
cells as a cone. The irregularity of the subsidiary cells distinguishes Rhabdotaenia.
The mode of origin of the lamina from the sides of the midrib may be important.
In R. danaeoides and in R. harkini there is no very sharp border to the midrib,
certainly no fold such as is seen in Taeniopteris vittata where the lamina arises rather
above the middle of the rachis and a good deal more rachis is seen from below than
from above. In Nilssonia it differs even more, arising from the top of the rachis.
The precise level of origin in Doratophyllum and Bjuvea has not been discussed.
In Glossopteris , from my own observations, the origin of the lamina is purely lateral ;
the midrib, however, being slightly depressed on the upperside and prominent below
may occasionally show folds at the sides but otherwise it grades imperceptibly into
the lamina. The epidermal structure of Rhabdotaenia also resembles that of Glossop-
teris and it may be that the two genera are naturally related.
Other genera which may be mentioned are the ill-defined Macrotaeniopteris
(type M, major) which proves to be a Bennettitalean leaf like several other species
and may be included in Nilssoniopteris , and Palaeovittaria (Feistmantel, 1876 :
368, pi. 19, figs. 3, 3«, 4, 4« ; Zeiller, 19020 : 81, pi. 16, fig. i) which differs in the
small angle of the veins from the midrib.
Rhabdotaenia danaeoides (Royle) n. comb.
(Text-fig, n)
1833. Glossopteris danaeoides Royle, p. 29, pi. 2, fig. 9.
1836. Aspidites danaeoides (Royle) Goeppert, p. 352.
1850. Pecopteris danaeoides (Royle) Unger, p. 170.
1901. Macrotaeniopteris danaeoides (Royle) Arber, p. 548 (pars).
19050. Taeniopteris danaeoides (Royle) Arber, p. 121, pi. 5, fig. I.
Similar leaves but with finer details unknown :
1850. Taeniopteris danaeoides (Royle) : McClelland, p. 56 pi. 15, fig. i.
1876. Taeniopteris danaeoides (Royle) : Feistmantel, p. 74.
18760. Macrotaeniopteris danaeoides (Royle) Feistmantel, p. 137.
18766. Macroteniopteris danaeoides (Royle) : Feistmantel, p. 305, pi. 19, figs, i, 2 ; pi. 21, fig. i.
1880. Macrotaeniopteris danaeoides (Royle) : Feistmantel, p. 88, pi. 200, figs, i, 2.
1886. Macrotaeniopteris danaeoides (Royle) : Feistmantel, p. 24, pi. 40, figs. 2, 3.
1893. Macrotaeniopteris danaeoides (Royle) : Oldham, pi. 2.
19050. Taeniopteris danaeoides (Royle) : Arber, p. 121 (pars).
EMENDED DIAGNOSIS. Leaf oval-oblong, widest in the middle region, typically
5-7 cm. wide, length possibly about 20 cm.; petiolate. Midrib up to 3 mm. wide
152
THE GLOSSOPTERIS FLORA OF TANGANYIKA
c iM
tftftt^X^S&P-fL
£'^•^•3 :::&••&•<'::'.
FIG. ii. Rhabdotaenia danaeoides (Royle). A, sinuous-walled cells of lower epidermis
between veins. V. 1959801. x 125. B, leaf showing venation. Neotype, V.4I9I.
X i. c, stoma showing exposed guard cells. ¥.195986. x 400. D, upper epidermis
showing straight-walled cells. V.i9598a. x 125. E, epidermis of midrib. ¥.41910.
X 125. F, stoma showing lateral lignine lamellae of guard cells. ¥.195980. x 400.
G, tracheid showing both bordered pits and transverse bars, from midrib. V. 19598^
X 800. H, xylem showing spiral, scalariform and pitted tracheids from midrib.
V.i9598c. x 800.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 153
below, narrower towards the apex, showing numerous longitudinal strands near leaf
base, fewer above. Margins entire or slightly undulate. Base more or less abruptly
contracted.
Lateral veins parallel, arising at right angles to the midrib or at a smaller angle
and almost immediately arching outwards and running at about 90° to midrib.
About one-third of lateral veins forked once, rest unbranched. Twice forked veins
very rare. Concentration of veins 12-16 per cm. near midrib, 15-20 per cm. near
margins.
Upper epidermis of lamina probably without stomata. Cells between veins
appearing polygonal with almost straight thick side-walls. Cells tending to be
arranged in rows parallel to the side veins. Cells above veins narrower. Upper
cuticle rather thick, like upper epidermis, surface showing longitudinal striations.
Upper and lower epidermis of midrib showing straight-walled rectangular cells
tending to be arranged in longitudinal rows. Surface sometimes showing longitudinal
striations, median papilla occasionally present, stomata absent.
Lower epidermis of lamina thinner than upper, stomatiferous. Cells between
veins irregularly shaped, typically about 40 fi wide X 95 /* long, lateral (anticlinal)
walls sinuous, about 3 fi thick, becoming gradually straight in the vicinity of
midrib, surface occasionally showing striations and a rather obscure median papilla.
Cells above veins straight-walled, rectangular, elongated in the direction of veins,
surface sometimes with fine longitudinal striations.
Stomata present in areas between veins. Subsidiary cells haplocheilic, numerous
(about seven), like ordinary cells, irregular, not forming a ring round guard cells,
guard cells exposed, typically 8 /* x 32 /*.
Lower cuticle thin, walls obscure.
NEOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.4191. Figured Royle (1833,
pi. 2, fig. 9) and Arber (19050, pi. 5, fig. i).
PARATYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.19598.
LOCALITY AND HORIZON. Burdwan Coalfield, India ; Permo-Carboniferous.
DESCRIPTION. The diagnosis is based on three leaves from Burdwan Coalfield,
India, namely Royle's type (¥.4191) and a block with two leaves (¥.19598). They
are very similar indeed but one of the leaves on ¥.19598 shows more of its petiole
which is 3 cm. long.
Without examining the originals it is impossible to say which of Feistmantel's
figured specimens belong to the same species ; all may well be the same. If so
the species includes quite small oval leaves (4 cm. X 2-5 cm.) as well as much wider
ones, up to n cm. wide and of unknown length.
Two cross connections occur between lateral veins in the neotype. Some guard
cells show lateral lamellae. The midrib shows abundant tracheids, sides 14-30 ji
wide. Some tracheids show spiral and annular thickenings but most are scalariform
(Text-fig, n, H). A few tracheids also show transverse bars between oval bordered
pits (Text-fig, n, G). There are in addition thick- walled elements showing usually
sparsely distributed, small, oval or rounded pinhole-like pits and cells with no pits
which may be tracheids or sclerenchyma.
Rhabdotaenia danaeoides is not the only species in the Lower Gondwana Coalfields
154 THE GLOSSOPTERIS FLORA OF TANGANYIKA
of India ; there is also Macrotaeniopteris feddeni figured by Feistmantel (1880 : 89,
pi. 2iA, fig. 3, pi. 22A, figs. 1-4 ; 1881 : 255, pi. 2, fig. i : 1882 : 31, pi. 21, fig. 5 ;
1886 : 24, pi. IA, fig. i). Dolianiti (1953 : 2, pis. i, 2) has recently reported this
species from Brazil.
Macrotaeniopteris feddeni is supposed to be distinguished chiefly by its more
crowded veins and thinner midrib. However, several specimens referred to M.
feddeni agree equally well with others referred to Macrotaeniopteris danaeoides e.g.
one (Feistmantel, 1882, pi. 21, fig. 5) has as few as n veins per cm. according to
the figure. I can see no difference in the midrib. No fine details are known.
Clearly M. feddeni is not, at present, adequately distinguished.
The Brazilian leaf, however, has 25-35 veins per cm. and is clearly distinct from
Rhabdotaenia danaeoides. The only other Lower Gondwana leaves are Taeniopteris
daintreei McCoy (1875 : 15, pi. 14, fig. i, 2), Angiopteridium spathulatum Etheridge
(1901 : 72), A. cf. mcclellandi Feistmantel (1880 : 92, pi. 2IA, figs. 4-7), Taeniopteris
criciumensis Dolianiti (1953, pi. 3) and some imperfectly known Taeniopteris leaves
which are all much narrower.
Rhabdotaenia danaeoides is compared with R. harkini below.
Rhabdotaenia harkini n. sp.
(PI. 20, fig. 5 ; Text-figs. 12-14)
DIAGNOSIS. Leaf about 6 cm. wide in widest part (presumably from middle
region of leaf), length unknown. Midrib at least about i mm. wide below, narrower
above. Margin entire, probably slightly undulate, very slightly converging towards
apex. Lateral veins parallel, arising almost at right angles or at a smaller angle
and almost immediately arching outwards and running at 8o°-9O° to midrib. Veins
forked once or twice or unbranched, usually dichotomising near midrib and often
elsewhere in the lamina. Cross connections between veins extremely rare. Some
veins bent slightly forwards near margin but no marginal vein present.
Concentration of veins 15-18 per cm. near midrib, 23-28 per cm. near margin.
Veins usually 145 fi thick but some veins up to 250 ju, thick at base. Substance of
lamina probably thin.
Upper epidermis of lamina without stomata. Cells between veins polygonal,
isodiametric or elongated in various directions typically 60 ju, x 45 ju, wide, lateral
(anticlinal) walls of cells straight, thick (up to 12 fi thick). Cells perhaps tending
to be arranged in rows parallel to veins. Surface of cells finely mottled, papillae
usually absent.
Upper cuticle about 2 fi thick, outlines of cells somewhat obscure, walls thin,
otherwise like upper epidermis.
Lower epidermis of lamina thinner than upper, stomatiferous. Cells between
veins irregularly shaped, isodiametric or elongated in various directions, typically
about 90 /* long x 60 /* wide, lateral (anticlinal) walls of cells sinuous but gradually
becoming straight near midrib. Cells near midrib and those in angles between
branches of veins sometimes unusually large and with almost straight walls. Cells
over veins narrow, elongated in the direction of veins, rectangular, lateral walls
THE GLOSSOPTERIS FLORA OF TANGANYIKA
155
straight. Lateral walls of lower epidermal cells often unevenly thickened, up to
3 /* thick. Cell surface usually showing a large thick-walled median papilla about
10-15 ju, in diameter. Papillae in some ordinary epidermal cells and subsidiary
cells of stomata (especially near imdrib) about 50 /* long, blunt and often with a
FIG. 12. Rhabdotaenia harkini n. sp. A, leaf showing venation. X 2. B, upper epi-
dermis showing straight- walled polygonal cells. X 125. c, a cell of the lower
epidermis from Text-fig. 13, B, showing the large median papilla and surface striations.
X 800. All from the Holotype (¥.34454).
rounded top. Cell surface sometimes showing a few ridges, but usually fine striations,
radiating from the base of the papilla. Striations often running parallel and con-
tinuous between adjacent cells and intervening walls, but usually in no definite
direction or arrangement in cells between veins, in cells above veins striations
usually parallel and running in the direction of the veins. Striations running longi-
tudinally along the surface of some laterally flattened papillae. Trichomes absent.
156
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 13. Rhabdotaenia harkini n. sp. A, lower epidermis of midrib (on left) and stomati-
ferous area between bases of two lateral veins showing sinuous-walled cells on the right.
X 125 B stomatiferous area between two veins. X 125. c, two closely placed
stomata, one (right) rather exposed, the other (left) protected by overlapping papillae.
X 400. D, cells below vein (lower side), x 125. All from the Holotype (¥.34454).
THE GLOSSOPTERIS FLORA OF TANGANKIYA 157
Stomata usually confined to areas between veins, rarely above veins, somewhat
unevenly distributed, concentration about 53 per sq. mm., orientation irregular.
Stomata haplocheilic, monocyclic or possibly partly amphicyclic. Guard cells
8-1 1 11 wide x 32-45 (i long, exposed or more or less covered by subsidiary-cell
papillae. Subsidiary cells 3-10, irregular, not forming a ring, polar subsidiary
cells like laterals, size and shape of subsidiary cells similar to ordinary epidermal
cells but often with a large hollow papilla on the side of the stomatal aperture, some-
times no well-marked papilla present but subsidiary cell margins thickened nearest
the guard cells. Papillae pointing upwards or over stomatal aperture. Surface of
subsidiary cells sometimes thicker than that of ordinary epidermal cells, showing
parallel striations radiating outwards from the side of the stoma.
Lower epidermis of midrib showing rows of straight-walled, longitudinally
elongated, polygonal cells, surface mottled, showing longitudinal striations, papillae
sometimes present but not prominent, stomata rarely present. Upper epidermis
of midrib unknown.
Lower cuticle of lamina thin (about i /i thick) walls of cells between veins obscure,
usually appearing broken but sometimes showing sinuous waves. Cell walls below
veins better marked, straight, thin. Cell surface mottled, occasionally showing
a median papilla and striations. Lower cuticle otherwise as lower epidermis.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.34454.
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. A single specimen was found (Text-fig. 12, A). The leaf substance
appears to have decayed considerably during preservation. It shows only the lower
epidermis and the substance of the veins. The upper epidermis is preserved only
at a few points (where it overlaps the lower and the intervening leaf substance).
In places the veins appear sinuous but this is clearly due to displacement during
preservation as the cells above veins run straight. There are two or possibly three
anastomoses between lateral veins. Occasionally veins and midrib show scalariform
tracheids with sides of varying width but usually about 14 /* wide and spiral tracheids
about 10 fi wide. Some exposed guard cells show lateral lamellae of Gymnosperm
type and thinner areas at the poles. A few two-winged pollen grains occur on the
epidermis.
The species is named after Mr. D. A. Harkin of the Tanganyika Geological Survey
who collected the material.
COMPARISON. Rhabdotaenia harkini differs from R. danaeoides in having a
comparatively higher concentration of veins especially near the margin (R. danaeoides
has 12-16 veins per cm. near the midrib, 15-20 near the margin ; R. harkini has
15-18 near the midrib, 24-28 near the margin). In R. harkini about half the veins
are forked and a number of these are forked twice ; in R. danaeoides only about
one-third of the veins are forked and twice-forked veins are very rare. The upper and
lower epidermis of the two species are, as far as they are known, similar ; the only
differences are : (i) the larger and more prominent papillae present in most cells
of the lower epidermis of R. harkini (in R. danaeoides the papillae are rather obscure
and only occasionally seen), and (2) in R. harkini, while some stomata are fully
GEOL. Ill, 4. 13
158
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 14. Rhabdotaenia harkini n. sp. A, stoma from stomatiferous area near midrib
protected by rather long papillae, x 400. B, part of Text-fig. 13, B showing details
of a moderately protected, possibly partly amphicyclic stoma. X 400. c, another
part of Text-fig. 13, B, showing details of two exposed stomata. X 400. D, a cell
under vein from Text-fig. 13, D, showing parallel longitudinal striations and median
papilla. X 800. All from the Holotype (¥.34454).
THE GLOSSOPTERIS FLORA OF TANGANYIKA 159
exposed exactly like those of R. danaeoides, there are others which are more or less
protected by epidermal papillae. No such protected stomata have yet been observed
in R. danaeoides.
These supposed differences are, however, based on the comparison of a single
specimen of R. harkini with three very incompletely preserved ones of R. danaeoides.
Thus the range of variation of neither species is known and, though they may
prove to be identical it seems advisable to distinguish them for the present.
Scale Leaves
(PI. 20, figs. 3, 4 ; Text-fig. 15)
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. Twelve scale leaves of varied form were examined and also some
fragments. Text-fig. 15, A shows a lanceolate scale overlying the cordate base of
another scale. They are 1-7-2 cm. long and 0-7-1-2 cm. wide, rather convex with
an ill-defined broad midrib and diverging veins which occasionally fork and
anastomose.
Text-fig. 15, B shows a small rounded scale with a broad base and Text-fig. 15,
c shows a group of somewhat larger rounded scales looking like a bud. The veins
are more distinct. There is no definite midrib but merely a number of longitudinal
strands. Again a few vein anastomoses were seen.
Both broad and lanceolate scales when isolated or transferred and seen in trans-
mitted light show a scarious margin. Some of them show fibres in vein meshes.
They usually have short simple hairs, about 300 p long, on their outer side and at
the margins, but some, e.g. the scale in Text-fig. 15, B, show none. The cuticles
of the different forms are rather similar. The cuticle of the outer (convex) side is
up to 2 /£ thick. Its cells are polygonal with straight or arched sides. The cells
above veins are narrower. Single-celled hair bases, occasionally with the cutinized
hair still attached, are unevenly distributed between the ordinary epidermal cells
(Text-fig. 15, D, E). Stomata are absent.
The cuticle of the inner (concave) side is more delicate with rather obscure cell
outlines. No definite stomata could be recognized and no hair bases were observed.
REMARKS. Similar scales have been described by Feistmantel (1880, 1882),
Zeiller (1896, 1902), Seward (1897, 1904), Arber (1905, 19050) Seward & Sahni
(1920), Walkom (1921, 1922, 1928, 1931), Walton (1929) and others. Zeiller (1896,
1902) figured some specimens clearly showing anastomosing veins as in Glossopteris.
Such scales have usually been attributed to Glossopteris, and sometimes to Noeg-
gerathiopsis , but with little evidence beyond association. In the cores from Mhukuru
they are found in association with leaves of Glossopteris fibrosa and G. colpodes
(as well as seeds, sporangia and roots). No specimens of Noeggerathiopsis occur in
the flora. The present specimens may be ordinary protective scales of the vegetative
bud of Glossopteris. Their form is fully consistent with this and their cuticles resemble
those of G. fibrosa. There is no reason at all to regard them as reproductive organs
i6o
THE GLOSSOPTERIS FLORA OF TANGANYIKA
and certain of these scales when transferred were proved to contain no sporangia,
seeds or other bodies.
Sporangia
Similar bodies :
1905. " Sporangium-like organs of Glossopteris browniana ", Arber, p. 324, pi. 30, figs. 1-3 ;
pi. 31, figs. 1-5.
19050. " Fructification of Glossopteris ", Arber, p. 39, text-figs. 12-15.
1907. " Sporangia ", Seward, p. 68, pi. 8, figs. 7, ja.
1919. " Sporangia ", Lundqvist, p. 12, pi. i, figs. 8, 9.
1928. " Microsporangia of Glossopteris (Arber, 1905) ", Walkom, p. 561.
B
D
(j
FIG. 15. Scale leaves. A, a lanceolate scale leaf and the cordate base of another.
V.34468. B, rounded scale leaf with a broad base (see also PI. 20 fig. 3). ¥.34453.
c, group of rounded scale leaves resembling a bud. ¥.34468. D, cuticle of the outer
(convex) side of a scale leaf with a cutinized hair and hair base. ¥.344680. E, cuticle
of the outer (convex) side of the same scale leaf showing two hair bases and epidermal
cells. F, two-celled hair from a scale leaf. V.34468&. G, H, single-celled hairs from
the same scale leaf, i, leaf epidermis with attached hairs. V.34468&. A-C, x 3 ;
D-I, x 125.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 161
1932. " Sporangia ", Du Toit, p. 384, pi. 40, figs. 9-12.
1955. " Male fructifications referable to Glossopteris ", Sen, p. 48, pis. 7, 8.
1956. " New type of fructification referable to Glossopteris ", Sen, p. 337, text-figs. 1-3.
Specimens from Mhukuru
(PI. 21, figs. 1-6 ; Text-figs. 16, 17, A-D)
Seventeen specimens, from Mhukuru Coalfield, Tanganyika, show a large number
of sporangia either isolated or occurring in groups. The best specimens were,
however, obtained by dissolving pieces of rock in HF. These show slender axes,
up to 200 fi thick, branching freely. Each ultimate branch ends in a single sporangium.
A longitudinal strand, at some points showing scalariform tracheids about 8 /* wide,
is present throughout the axes. Their surface cells are elongated but at a few points
in the thicker axes they are short.
Laterally compressed sporangia are oval with one side nearly straight, the other
strongly curved (Text-figs. 16, A, E, F, 17, c). They are up to 3 mm. X 1-25 mm. but
many are much shorter, some because they were compressed longitudinally (Text-
fig. 16, B) but others must have been smaller originally.
Most of the sporangia have dehisced along the whole length of their convex side
but the line of dehiscence is often oblique. Two flaps of the wall of such sporangia
are either compressed almost flat or contain rock matrix and stray spores. A few
sporangia are still closed and full of spores (PL 21, figs. 5, 6). These enclosed spores
are oval and two-winged, 40-55 fi long and 25-30 fi wide (Text-fig. 17, D). One
sporangium, however, which must be another species, contained winged spores
measuring about 70 JLL by 50 /£. In other respects this sporangium is similar to those
with smaller spores.
The wall of the sporangium consists of an outer layer of elongated fibrous cells
running lengthwise (parallel with the slit). The wall cells are 150-400 /* long
and 6-35 /* wide tending to be narrower towards the apex and base. The surface
of the wall cells is thick and uniformly dark ; a few show irregular paler areas
probably due to poor preservation. The sides of the cells are straight or slightly
sinuous (Text-figs. 16, G, H). The lines where these cells join are often represented
by narrow and occasionally by wide gaps (the cuticle, however, is without any corres-
ponding gaps). The outer surface of the sporangial wall cells projects and their
walls are sunken. The inner surface of the wall is almost smooth. No specialized
cells were recognized along the suture. At the apex of the sporangium the wall
cells radiate from a point which is sometimes slightly projecting and marks the
apical end of the slit. No specialized cells like an annulus occur anywhere.
At the base of the sporangium the narrowing fibrous cells of the wall converge
and enter the stalk which is up to i mm. long X o-i mm. thick. It is often attached
asymmetrically to the straight side of the sporangium (Text-figs. 16, A, E, F, 17, c).
The body of some sporangia is bent almost at right angles to the stalk (Text-fig.
16, E, F).
On maceration the sporangium yields two cuticular coats, an outer smooth coat
showing the outlines of the fibrous cells and an inner granular membrane showing
162
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 1 6. Sporangia from Mhukuru Coalfield, Tanganyika. A, branching stalks termi-
nating in sporangia ; some sporangia have broken off. ¥.34457. x 25. B, trans-
versely flattened sporangium. ¥.34469. x 50. c, forked axis bearing terminal
sporangia. ¥.34457. x 25. D, sporangium showing wall-cells radiating from a
point at the apex. ¥.34469. x 50. E, group of sporangia attached to a branching
axis (see also PI. 21, fig. 2). ¥.34457. x 25. F, sporangium, side view, bent almost
at right angles to the stalk. ¥.34470. x 25. G, sporangium wall showing dark
cell-interior from sporangium in PI. 21, fig. 3. ¥.34458. x 125. H, wall cells of a
sporangium. ¥.34471. x 125.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 163
straight or slightly sinuous- walled, isodiametric polygonal cell outlines typically
about 40 11 across. The cells of the outer membrane, particularly those near the
sporangial stalk, may show a median rounded mark (papilla) . Some of the dehisced
sporangia show spores in great variety which had evidently got in before preservation.
The sporangia occur in association with leaves of G. fibrosa and G. colpodes (as
well as scale leaves, Spermatites crystallinus, S. tetrapterus, Veriebraria, and some
thin roots).
Specimens from Australia
(PI. 21, fig. 7 ; Text-fig. 17, E, F)
A few sporangia from Newcastle, New South Wales, have been examined and the
following notes throw additional light on the African material.
1. Detached sporangia. Fourteen specimens from Newcastle (additional to those
described by Arber, 1905, 1905^) agree very closely with the African ones. None
of them is, however, as large as the largest African sporangia. They measure 1-2-5
mm. x 0-6-1-3 mm. A few of them show stalks about 100 /* thick. The wall
shows very similar cells to the African ones (projecting outwards but smooth inside)
and cuticles were obtained corresponding to the African ones. All have dehisced
but a few retain numerous spores, all of one type (Text-fig. 17, F).
The sporangia figured by Arber (1905, 1905^) are much like these specimens
(Text-fig. 17, E) except that those described by Arber have suffered complete
oxidation and no cuticle or other organic matter remains.
2. Attached sporangia. Two specimens are available. One (¥.24233) is a slightly
concave disc (PI. 21, fig. 7) thickly covered with sporangia compressed in various
planes. No stalks were seen and it is not known how the sporangia were attached.
The sporangia are empty. A little carbon of the wall remains but the wall cells are
replaced by delicate rods of a mineral. The margins of the disc show small epidermal
cells arranged in parallel rows. The rows are not concentric but meet the margin
at a small angle. The surface of the disc where it is not covered with sporangia is
wrinkled.
The other specimen, ¥.24244, is a fragment of what appears to be a similar disc.
Its sporangia are full of spores but they proved ill-preserved. They are represented
by rounded outlines about 14 /* wide and they are probably two-winged. This
specimen was transferred and the back of it proved to be smooth ; its surface
shows small epidermal cells arranged in parallel rows, running from what is regarded
as the base to the apex, i.e. longitudinally.
DISCUSSION. The Newcastle specimens were finally proved to be sporangia
by the discovery of a specimen (¥.24244) with sporangia still full of spores. They
look very much like the African ones, which also have been proved to be sporangia.
Arber (1905, 19050) who first noticed them believed them to be sporangia ; Seward
(1907) and Lundqvist (1919) agreed. Thomas, H. H. & Mrs. (1925) and Walkom
(1928) thought that they might be ramenta. The presence of spores adhering to
empty specimens as noted by du Toit (1932) is not enough to prove them to be
sporangia, but the occurrence of nearly ripe specimens full of spores, such as those
i64
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 17. Sporangia and spores. A, longitudinally dehisced sporangium. ¥.34457.
X 50. B, two-winged spores of diverse sizes adhering to the inner cuticle of a dehisced
sporangium (see also PL 21, fig. i). V. 34456. x 125. c, dehisced sporangium in side
view. The stalk is attached on the flat side. ¥.34457. X 50. D, two-winged spore
from an undehisced sporangium. ¥.34472. x 800. E, carbonized sporangium as
seen in a transfer preparation. ¥.34491^. x 50. F, typical two-winged spore from
a dehisced sporangium in a pull. The black spot is part of a wall cell. ¥.242450.
X 800. A-D from Mhukuru Coalfield, Tanganyika. E, F from Newcastle, New
South Wales.
in specimen ¥.24244 and the undehisced African specimens, make their identity
certain.
Sen (1955) described a few similar sacs but apparently with transverse dehiscence,
which suggests that they are different from the specimens described here. In 1956
he described others with longitudinal dehiscence which are even more similar.
Arber (1905 : 41, 19050 : 325) described and figured a specimen, ¥.7202, in which
he believed that these sporangia were borne on a scale (perhaps like one of the scales
here regarded as bud scales). There is little of this scale preserved, however.
Re-examination of the specimen merely shows that some of the sporangia lie beneath
the scale fragment while others are clear. There is no evidence at all of attachment.
No organ exactly like the disc has been described. Several incompletely known
fossils may, however, prove similar. These include Eretmonia natalensis du Toit
(1932, pi. 40, figs. 9-12), Conites Seward (1897) and Seward and Leslie (1908) and
the scale, " Glossopteris sp." (White, 1908, pi. 7, figs. 5, 50, 6), which shows small
rounded bodies on its surface. Ottokaria ovalis White (1908, pi. 7, figs. 7, 70) and
Ottokaria leslii Thomas (1921 : 285, text-figs, i, 2) may be similar.
Ottokaria bengalensis Zeiller (1902) and Seward & Sahni (1920), Lanceolatus and
Scutum Plumstead (1952, 1956) are assumed to be of a different nature. The rather
regular arrangement of rounded bodies in Scutum is apparently different from the
irregular arrangement of sporangia seen in the discs described here. I can see no
similarity between the much larger " male bracts " of that organ and the present
sporangia.
These Australian sporangia are of particular interest in relation to the African
ones because while they look so similar they are probably borne in rather different
ways. Not a single African specimen, in the present collection, shows a disc-like
structure and it cannot be assumed that the Australian and African sporangia
belong to closely related plants. Both of course occur in Glossopteris floras in asso-
ciation with various species of Glossopteris. It is possible that they belong to the
same plants as the associated leaves but so many varied organs have been assigned
to Glossopteris, on grounds of association, that I refrain from assigning these sporangia
to it.
Genus SPERMATITES Miner
This non-committal designation is used because the structure of this seed though
characteristic is not adequately understood. I feel sure that when it is so understood
it will be well worthy of generic rank.
Spermatites crystallinus n. sp.
(PI. 20, fig. 6 ; Text-figs. 18, 19)
DIAGNOSIS. Seed, flattened, orthotropous, 1-25-1-5 mm. long, i-i-i mm. wide
and 0-75 mm. thick, oval, micropylar end obtusely pointed, chalazal end rounded,
margins often bearing fragments of tissue (wing or possibly fruit substance?).
Surface of seed often showing longitudinal rows of polygonal cells about 30 /* wide.
Cuticles : nucellus thickly cutinized showing elongated cells 30-75 /* long and 20-40
166
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 18. Spermatites crystallinus n. sp. A-C, compressed seeds showing plates of
tissue at the margin. B, shows a narrower seed with a longitudinal ridge on the surface.
A, Holotype, "^34473. B, "^34474. c, V. 34475. D, surface cells of marginal plate
of tissue as seen before maceration of seed in A. E, surface cells of seed in A before
maceration. F, cuticle of plate of tissue at margin of seed in A (broken while mounting).
G, nucellus of seed in A with portions of outer membranes. H, cuticle of seed
immediately outside the nucellus (inner cuticle of integument). "^34476. i, j, cuticles
of seeds showing micropylar canals and chalaza. "^34477-78. K, cuticle of seed.
V.34479. L, cuticle in F more magnified ; a few two-winged pollen grains are sticking
to it. M, top part of G, more magnified, showing micropylar canal. The three dark
masses are unmacerated carbon. A-C, F-K x 25. D, E, L, M x 125.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 167
ju, wide, walls thick towards micropylar end, thinner elsewhere, straight or slightly
wavy. Micropylar canal 80-90 /£ long, chalaza about 300 /* wide. Megaspore
membrane absent but nucellus enclosing a dark-coloured tissue often showing
polygonal cells typically 15 fi long X 45 fi wide, walls up to 2 /* thick, usually
appearing hyaline. Outer cuticles delicate ; immediately outside nucellus a delicate
membrane (? inner lining of integument) showing narrow elongated cells which
become sinuous near micropyle, cells typically 70 fi long X 15 fi wide.
Stone of integument not very thick, composed of elongated fibres. Outer cuticle
(possibly surface of integument) showing short polygonal cells about 30 /i wide,
cell surface often showing a rhomboidal, hexagonal or octahedral mark like the
imprint of a crystal.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.34473.
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. Thirty-three of these seeds were isolated by dissolving pieces of
rock in HF. Most of them are rounded but a few are narrow and show a longitudinal
ridge on either surface (Text-fig. 18, B). On maceration their nucellus shows folds
corresponding to the surface ridges. I believe these are laterally compressed seeds,
they are about 0-75 mm. thick which is perhaps the original thickness of the seed.
Several seeds carry irregular plates of tissue at the margin and these may occur
at any point (Text-figs. 18, A-c). The irregular margins of these plates may be
original or may have been caused by breakage before preservation or during extrac-
tion. This tissue yields a delicate cuticle showing polygonal cells about 30 /* wide.
It might represent some sort of wing or may possibly be a relic of the tissue of a
fructification.
Maceration always yields the nucellus cuticle and usually fragments of other
cuticles around it. Their nature is, however, not as clear as that of the nucellus.
The inner cuticle of integument is best seen towards the micropyle. A few seeds
yielded internal casts of fibres in the form of rods covered with short outgrowths
which must have originally occupied pits. One seed yielded fragments of cuticle
showing elongated tracheid-like cells with scalariform marks (Text-fig. 19, i) probably
representing impressions of tracheids (cf. Harris, 1941 ; 88, text-fig. 5, F.). The
nature of the outer cuticle showing crystalline imprints is not clear and could not
be proved to be the same as the cells seen on the outside of the seed though it may
be so.
Seven out of twelve seeds showing suitable micropylar ends have pollen grains on
the surface of the nucellus around its micropylar end. In two seeds they also occur
farther away from the micropyle. They are underneath the delicate inner cuticle
of the integument and as far as can be seen most of them are of the same type.
Those seen clearly are two-winged, about 30 /i x 50 ju, in size (Text-fig. 19, G) but
many are so crushed that their shape is obscured ; their texture is, however, similar
to the clearer ones. Two exceptional round pollen grains about 60 fi in diameter
were also noticed (Text-fig. 19, E). The presence of these numerous pollen grains
suggests that they are not accidental but are the result of the normal pollination
processes. The grains look just like those of the microsporangia and they provide
168
THE GLOSSOPTERIS FLORA OF TANGANYIKA
H
FIG. ig. Spermatites crystallinus n. sp. A, nucellar cuticle showing elongated cells.
¥.34480. x 200. B, outer cuticle of seed showing polygonal cells and crystalline
imprints. ¥.344810. x 200. c. dark tissue inside nucellus showing hyaline cell
walls. V.3448a. x 200. D, portion of cuticle immediately outside nucellus in
Text-fig. 18, H, from near micropylar end of seed showing sinuous-walled elongated
cells and a pollen grain. V. 34476. x 200. E, unusual rounded pollen grain from the
nucellus of a seed. ¥.34475. x 800. F, part of nucellus of seed showing numerous
pollen grains at the micropylar end. A number of pollen grains are also seen sticking
to the plate of tissue outside the micropyle. ¥.34483. x 50. G, pollen grain from
the nucellus of a seed showing two wings and a body. ¥.34483. x 800. H, Chalazal
hole of seed shown in Text-fig. 18, i. x 125. i, membrane showing scalariform
imprints. ¥.34479. x 400.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 169
evidence that these seeds and microsporangia probably belong to the same plant
species. It is noteworthy that the micropylar canal is shrivelled and narrower than
the diameter of a pollen grain. It must have been wider when it admitted them.
Clearly the pollination is essentially gymnospermous. Similar pollen is found also
on the surface of the seed where, however, it is not significant.
COMPARISON. The most similar seed of the Glossopteris flora is Spermatites
indicus Srivastava (1954). It differs, however, in having surface cells twice as wide
as those in 5. crystallinus. Certain inner cuticles are present in S. indicus which
may correspond to those of S. crystallinus but their nature is very obscure. There
is nothing to suggest that S. indicus has any sort of marginal tissue.
Rather similar orthotropous seeds are known in the Mesozoic genus Caytonia
(Harris, 1940 : 724) where, however, no marginal flange exists, and in the isolated
seeds described by Harris (1932 : 14) under the name Amphorispertmtm.
Spermatites tetrapterus n. sp.
(Text-figs. 20, 21)
DIAGNOSIS. Seed, flattened, orthotropous, 3-4-5 mm. long and 2-2-5 mm- wide,
oval, micropylar end obtusely pointed, chalazal end rounded. Seed showing two
strong ridges on the surface, representing compressed wings, other specimens
without surface ridges. Surface showing rows of longitudinally elongated ceUs
about 150 11 long and 25 fi wide. Cuticles : Nucellus moderately thickly cutinized
showing elongated cells with straight or nearly straight walls, apical region or nucellar
cap sharply distinguished by thick-walled cells with straight or nearly straight
walls, up to 2 fi thick, lower part with thinner-walled cells, boundary region between
nucellar cap and lower part normally showing scattered round thickenings 6-12 /*
across. Micropylar canal very delicately cutinized, about 200 ju, long X 100 /* wide,
usually collapsed (chalaza not seen).
Megaspore membrane not present, nucellus often enclosing dark matter sometimes
showing transversely elongated cell outlines. This tissue extends only as far as the
base of the nucellar cap.
Inner cuticle of integument delicate showing obscurely marked longitudinally
elongated cells.
Stone of integument not very thick, containing elongated fibres.
Outer cuticle of integument about I /* thick, showing cells which are usually
elongated and narrow. Typical dimensions 150 /£ x 25 //. but sometimes short and
broad.
HOLOTYPE. Brit. Mus. (N.H.) Palaeont. Dept. no. ¥.34490.
LOCALITY AND HORIZON. Mhukuru Coalfield, Songea District, Tanganyika ;
Ecca Series (" Upper Coal Measures ").
DESCRIPTION. Twenty-seven seeds were seen, seventeen of which were obtained
by disssolving pieces of rock in HF. Some seeds are filled with rock matrix. If the
ridges in the seed represent wings, they may be four-winged. The nucellus of seeds
with surface ridges shows deep folds corresponding with the ridges, indicating that
the nucellus of the seed was itself angular. In some seeds the cells of the nucellar
1 7o
THE GLOSSOPTERIS FLORA OF TANGANYIKA
FIG. 20. Spermatites tetrapterus n. sp. A, upper part of macerated seed showing nucellus,
micropylar canal and outer membrane. V. 34484. X 25. B and c, two pollen grains
(from inside the nucellus) showing two wings and a striped body. ¥.34485.
X 800. D, unusual pollen grain from inside the nucellus of a seed. ¥.34486.
X 800. E, outer membrane of seed showing straight-walled elongated cells.
¥.34487. x 200. F, compressed seed showing a ridge on the upper surface. Its
interior is filled with rock matrix. Holotype, ¥.34490. x 25. G, surface cells of seed
in F. x 125. H, nucellar cuticle of a seed. The black spot at the top represents pollen
grains, a darker tissue is seen below the nucellar cap. ¥.34486. X 25.
THE GLOSSOPTERIS FLORA OF TANGANYIKA
171
cap form processes (Text-fig. 21, A-C) . Text-fig. 21, A shows the top view of a nucellar
cap in a seed which is probably obliquely compressed.
Seven out of ten seeds with suitable micropylar ends show numerous oval pollen
grains inside the top end of the nucellus. In one seed (Slide ¥.34494) a mass of pollen
grains is also seen inside the micropylar canal. Most of the measureable pollen
D
FIG 21. Spermatites tetrapterus n. sp. A, nucellar cap in top view showing radiating
cell rows. ¥.34488. x 50. B, nucellus and inner cuticle of integument showing
nucellar cap in side view. ¥.34489. x 25. c, upper part of B. There is a delicate
cuticle outside the nucellus (inner lining of integument) and the apex of the nucellus
forms processes. ¥.34489. x 125. D, part of B. The cells above belong to the
nucellar cap and the dark spots occur at its junction with lower part of the nucellus.
X 200.
172 THE GLOSSOPTERIS FLORA OF TANGANYIKA
grains are about 50 fi x 30 ju. They are two-winged and show a body with transverse
striations. One pollen grain which has a similar texture and shows no wings is
60 fi X 40 fi, another is a smaller two-winged one measuring 35 fi X 25 /*.
COMPARISON. Similar seeds have been described from Wankie, Rhodesia, by
Walton (1929 : n, pi. c, figs. 25, 26) but their fine details are unknown. These
seeds show wing-like expansions which Walton regarded as formed by the angular
inner tissue.
5. tetrapterus resembles S. crystallinus in its general organization but differs in
being larger and in having surface ridges, S. crystallinus shows ridges only when
laterally flattened and has a marginal tissue which is not present in 5. tetrapterus.
It has short surface cells while S. tetrapterus usually shows longitudinally elongated
cells on the surface. S. crystallinus has an outer membrane showing short cells
with crystalline imprints whereas the outer cuticle of 5. tetrapterus usually shows
elongated cells and no crystals.
III. SUMMARY
The paper describes details of the epidermal and cuticular structure of three
new species of Glossopteris (G. fibrosa, G. hispida, G. colpodes} and a possible fourth
(Glossopteris sp. A) from borehole cores in the Mhukuru Coalfield, Songea District
Tanganyika. All the Glossopteris leaves, described here, agree in their haplocheilic
stomata and in their guard cells having lignine lamellae of the Gymnosperm type.
They range from mono- to amphicyclic but all have subsidiary cell papillae covering
the stoma, palisade and spongy mesophyll cells and scalariform tracheids in veins.
Fibres in vein meshes are found in G. fibrosa and G. hispida. G. colpodes is a wider-
meshed form usually showing sinuous-walled epidermal cells on both the upper and
the lower sides of the leaf. G. hispida is unique in having multicellular hairs on the
lower surface of the leaf. The imperfectly known Glossopteris sp. A is rather like
G. fibrosa but has no fibres in vein meshes. The species are compared with others
of Glossopteris described by earlier authors.
A new genus Rhabdotaenia is made for the Indian leaf Taeniopteris danaeoides
(Royle) and its fine structure is described. A rather similar leaf, R. harkini, is
described from Mhukuru, Tanganyika. Rhabdotaenia shows haplocheilic stomata
and differs from other taeniopterid haplocheilic leaves, e.g. Doratophyllum and
Bjuvea (Florin, 1933) in having irregular subsidiary cells and rather exposed stomata
with or without protecting subsidiary cell papillae.
Some rounded or lanceolate scale-leaves showing a scarious margin, an ill-defined
midrib and anastomosing veins have been studied. They usually have short simple
hairs on their convex side and occasionally fibres in vein meshes. The epidermal
structure of the various forms is similar and resembles that of G. fibrosa of which
they may be bud scales.
Fine details of some well-preserved African and Australian microsporangia,
closely resembling Arber's " sporangium-like organs of Glossopteris browniana "
are described. Some undehisced specimens were found to be full of two-winged
spores. It was found that the African sporangia are borne terminally on branched
THE GLOSSOPTERIS FLORA OF TANGANYIKA 173
slender axes. Two discs bearing similar sporangia, from Newcastle, New South
Wales, are also described ; one of them has sporangia full of spores. The finding
of undehisced specimens, still full of spores, finally proves these bodies to be sporangia.
Two types of compressed seeds, Spermatites crystallinus and S. tetrapterus were
found in the cores. Their structure is described in detail. Both show gymno-
spermous pollination by two-winged spores. Both the sporangia and the seeds may
belong to the same plants as the Glossopteris leaves abundant in this collection.
Similar sporangia and seeds occur in Glossopteris bearing rocks at Ranigenj coalfield.
IV. ACKNOWLEDGMENTS
I take this opportunity to express my deep sense of gratitude to Professor T. M.
Harris for his keen interest, kind advice and very generous help throughout the
progress of this work. I am deeply grateful to Professor J. Walton for providing
me with the material and for his kind advice while I worked at his laboratory in
Glasgow. I am also highly indebted to the late Mr. W. N. Edwards, formerly
Keeper of Geology, and his successor Dr. E. I. White for permission to study the
extensive collections of the Glossopteris flora in the Department of Palaeontology,
British Museum (Natural History), and to Mr. F. M. Wonnacott for helping me in
many ways.
Special thanks are due to Mr. J. M. Edmonds, University Museum, Oxford,
Dr. E. Boureau, Laboratoire Anatomie Comparee Vegetale, Paris, and the Keeper
of the Museum, Scole des Mines, Paris, for permission to examine the specimens
of Glossopteris in their collections, Monsieur L. Grambast, Faculte des Sciences,
Paris University, for help in locating Zeiller's specimens in Paris and Dr. G. A. Cooper,
Curator of Invertebrate Palaeontology and Palaeobotany, and Dr. S. H. Mamay,
U. S. National Museum, Washington, for sending me photographs and a pull from
the types of Glossopteris ampla Dana.
I am grateful to Mr. L. C. Willis, Reading University, and to Mr. W. Anderson,
Glasgow University, for photographic assistance, and to Miss H. Wilkinson and
Mr. H. D. Scammell for facilitating my work at the British Museum (Natural History).
I owe my thanks to the Committee for Commonwealth Interchange, British
Council, for a travel grant and to the Government of Uttar Pradesh for financial
assistance enabling me to visit Britain. Lastly, I thank my wife, Radha Pant,
for much help and encouragement during difficult times.
V. REFERENCES
ARBER, E. A. N. 1901. Notes on. Royle's types of fossil plants from India. Geol. Mag.f
London, 8 : 546-549.
1905. On the Sporangium-like organs of Glossopteris Browniana, Brongn. Quart. J.
Geol. Soc. Lond., 61 : 324-338, pis. 30, 31.
• I9O5«. Catalogue of the Fossil Plants of the Glossopteris Flora in the Department of Geology t
British Museum (Natural History}. Ixxiv + 255 pp., 8 pis. Brit. Mus. (N.H.), London.
BRONGNIART, A. 1828. Prodrome d'une Histoire des Vegetaux fossiles. viii + 223 pp. Paris.
1828-38. Histoire des Vegetaux fossiles, ou recherches botaniques et geologiques sur les
vegetaux renfermes dans les diverse Couches du Globe. 2 vols. Paris.
BUNBURY, C. J. F. 1861. Notes on a collection of fossil plants from Nagpur, Central India.
Quart. J. Geol. Soc. Lond., 17 : 325-346, pis. 8-12.
GEOL. Ill, 4. 14
174 THE GLOSSOPTERIS FLORA OF TANGANYIKA
DANA, J. D. 1849. Wilkes' United States exploring expedition . . . 1838-1842, 10.
Geology, xii -j- 756 pp., atlas 21 pis. New York.
DOLIANITI, E. 1953. A Flora do Gondwana inferior em Santa Catarina, II. O Genero
Taeniopteris. Notas Prelim. Est., Min. Agric. Div. Geol. Min., Rio de Janeiro, 61.
DU TOIT, A. L. 1932. Some fossil plants from the Karroo System of South Africa. Ann.
S. Afr. Mus., Cape Town, 28 : 369-393, pis. 39, 40.
19320. Some fossil plants from the Gondwana Beds of Uganda. Ann. S. Afr. Mus.,
Cape Town, 28 : 395-406, pi. 41.
ETHERIDGE, R. Jun. 1901. Notes on fossil plants from the Saint Lucia Bay Coalfield, Ense-
leni River, Zululand. Rept. Geol. Surv. Natal &• Zululand, London, 1 : 69-76, pi. 13.
FEISTMANTEL, O. 1876. Notes on the age of some fossil floras in India. Rec. Geol. Surv.
India, Calcutta, 9, 3 : 63-79, pis. 3-5.
18760. On some fossil plants from the Damuda Series in the Raniganj Coalfield collected
by Mr. J. Wood-Mason. /. Asiat. Soc. Beng., Calcutta, 45 : 329-380, pis. 15-21.
1878-79. Palaeozoische und Mesozoische Flora des ostlichen Australiens. Palaeonto-
graphica, Stuttgart (Suppl.) 3, 2-4 : 53-195, pis. 1-30.
1879. The fossil flora of the Gondwana System, 3(1 and Suppl.). The flora of the Talchir-
Karharbari Beds. Palaeont. indica, Calcutta (12) : 1-64, pis. 1-31.
1880-81 . The fossil flora of the Gondwana System, 3, (2 and 3) . The flora of the Damuda
and Panchet Divisions. Palaeont. indica, Calcutta (12) : 1-149, pis. IA-47A.
1882. The fossil flora of the Gondwana System, 4 (i). The fossil flora of the South
Rewah Gondwana Basin. Palaeont. indica, Calcutta (12) : 1-52, pis. 1-21. x
— — 1886. The fossil flora of the Gondwana System, 4 (2). The fossil flora of some of the
Coalfields in Western Bengal. Palaeont. indica, Calcutta (12) : 1-66, pis. IA-I4A.
1889. Ubersichtliche dartstellung der Geologisch-Palaeontologischen Verhaltnisse Siid-
Afrikas, i. Die Karoo-formation und die dieselbe unterlagernden Schichten. Abh.
bohm. Ges. Wiss. Prag. (7) 3, 6 : 1-89, pis. 1-4.
FLORIN, R. 1933. Studien iiber die Cycadales des Mesozoikums. K. svenska VetenskAkad.
Handl., Stockholm (3) 12, 5 : 1-134, P^8- 1-16.
GOEPPERT, H. R. 1836. Die fossilen Farrnkrauter : Systema Filicum Fossilium. Nova Acta
Leop. Carol., Halle a.S., 17 (Suppl.) : 1-486, pis. 1-44.
GOTHAN, W. 1914. Die fossile Flora des Tete-Beckens am Sambesi. Branca-Festschrift,
Berlin, 1914 : 11-15, pi- *•
HARKIN, D. A. 1953. The geology of the Mhukuru Coalfield (Songea District). Tanganyika
Geol. Surv. Dept. Short Paper, 28 : 1-66.
HARRIS, T. M. 1932. The fossil flora of Scoresby Sound, East Greenland, 2. Description of
seed plants Incertae Sedis together with a discussion of certain Cycadophyte cuticles.
Medd. Gronland, Kobenhavn, 85, 3 : 1-112, pis. 1-9.
19320. The fossil flora of Scoresby Sound, East Greenland, 3. Caytoniales and Bennet-
titales. Medd. Gronland, Kobenhavn, 85, 5 : 1-133, pis- i-iQ-
1940. Caytonia. Ann. Bot., London (n.s.) 4 : 713-734, pi. 7.
1941- Cones of Extinct Cycadales from the Jurassic rocks of Yorkshire. Philos. Trans.,
London, 231 (B) : 75-98, pis. 5, 6.
LUNDQVIST, G. 1919. Fossile Pflanzen der Glossopteris Flora aus Brasilien. K. svenska
VetenskAkad. Handl., Stockholm, 60, 3 : 1-36, pis. i, 2.
MCCLELLAND, J. 1850. Report of the Geological Survey of India for the season 1848-49. 92
pp., 17 pis. Calcutta.
McCov, F. 1875. Prodromus of the Palaeontology of Victoria , or, Figures and Descriptions
Of Victorian Organic Remains. Dec. i and 2. Geol. Surv. Victoria, Melbourne.
MINER, E. L. 1935. Palaeobotanical examination of Cretaceous and Tertiary rocks. Amer.
Midi. Nat., Notre Dame, Ind., 16 : 585-625.
OLDHAM, R. D. 1893. A Manual of the Geology of India. 2nd. edit, xxiii + 543 pp.
PANT, D. D. 1956. On two compressed Palaeozoic axes : Stigmaria ficoides in Gymnostrobus
condition and Vertebraria indica. Ann. Bot., London (n.s.) 20 : 419—429, pi. 18.
THE GLOSSOPTERIS FLORA OF TANGANYIKA 175
PLUMSTEAD, E. P. Description of two new genera and six new species of fructifications borne
on Glossopteris leaves. Trans. Geol. Soc. S. Afr., Johannesburg, 55 : 281-328, pis. 43-52.
1956. Bisexual fructifications borne on Glossopteris leaves from S. Africa. Palaeonto-
graphica, Stuttgart, 100 (B) : 1-25, pis. 1-14.
ROYLE, J. F. 1833. Illustrations of the Botany and other branches of the natural history of the
Himalayan Mountains and of the Flora of Cashmere. 2 vols. Ixxxii + 472 pp., atlas
100 pis. London.
SAHNI, B. 1923. On the structure of the cuticle in Glossopteris angustifolia Brongn. Rec.
Geol. Sum. India, Calcutta, 54 : 277-280, pi. 17.
SEN, J. 1955. On some male fructifications referable to Glossopteris and the systematic
position of the genus. Proc. Nat. Inst. Sci. India, 21 (B) : 48-51, pis. 7, 8.
1956. A new type of fructification referable to Glossopteris. Nature, Lond., 177 : 337-338.
SEWARD, A. C. 1897. On the association of Sigillaria and Glossopteris in South Africa. Quart.
J. Geol. Soc. Lond., 53 : 315-338, pis. 21-24.
1904. Report on collections of Natal fossil plants from the Ecca Coal Series of Umhlali on
the north-east coast of Natal. Kept. Geol. Surv. Natal &• Zululand, London, 2 : 97-101,
pi. 4.
1907. On a collection of Permo-Carboniferous plants from the St. Lucia (Somkele)
Coalfield, Zululand, and from the Newcastle District, Natal. Trans. Geol. Soc. S. Afr.,
Johannesburg, 10 : 65-73, pis- 8, 9.
SEWARD, A. C. & LESLIE, T. N. 1908. Permo-Carboniferous plants from Vereeniging. Trans-
vaal.) Quart. J. Geol. Soc. Lond., 64 : 109-126, pis. 9, 10.
SEWARD, A. C. & SAHNI, B. 1920. Indian Gondwana plants : a revision. Palaeont. indica,
Calcutta (n.s.) 7, i : 1-54, pis. 1-7.
SRIVASTAVA, P. N. 1954. On some Lower Gondwana megaspores and seeds from Mangardaha
coal, West Bokaro, Bihar. Palaeobotanist, Lucknow, 3 : 113-115, pis. i, 2.
TATE, R. 1867. On some Secondary Fossils from South Africa. Quart. J. Geol. Soc. Lond.,
23 : 139-175. pis. 5-9-
THOMAS, H. H. 1921. An Ottokaria-like plant from South Africa. Quart. J. Geol. Soc. Lond.,
77 : 285-288, 2 figs.
& Mrs. 1925. On the ancestry of the Caytoniales. Rep. Brit. Ass. Adv. Sci., London
(93rd Meeting) : 364.
UNGER, F. 1850. Genera et Species Plantarum Fossilium. xl + 627 pp. Vindobonae.
WALKOM, A. B. 1921. On Nummulospermum gen. nov., the probable megasporangium of
Glossopteris. Quart. J. Geol. Soc., Lond., 77 : 289-295, pi. 21.
1922. Palaeozoic floras of Queensland, Pt. i. The flora of the Lower and Upper Bo wen
Series. Queensld Geol. Surv. Publn, 270 : 1-64, pis. 1-9.
1928. Notes on some additions to the Glossopteris flora in New South Wales. Proc. Linn.
Soc. N.S.W., Sydney, 53 : 555-564, pi. 36.
WALTON, J. 1929. The Fossil flora of the Karroo System in the Wankie District, Southern
Rhodesia. Bull. Geol. Surv. S. Rhodesia, Salisbury, 15 : 62-75, pis. A-C.
1936. On the factors which influence the form of fossil plants with descriptions of the
foliage of some species of the Palaeozoic Equisetalean genus Annularia Sterriberg. Philos.
Trans., London, 226 (B) : 219-237.
I953- Determination of fossil plants from Mhukuru Coalfield, Songea District, Tangan-
yika. Tanganyika Geol. Surv. Dept. Short Paper, 28 : 28-30.
WHITE, D. 1908. Fossil flora of the Coal Measures of Brazil. In White, I. C. Comissao
estudos das Minas de Carvao de Pedra do Brazil, Rio de Janerio : 337-617.
ZEILLER, R. 1896. F-tude sur quelques fossiles, en particulier Vertebraria et Glossopteris, des
environs de Johannesburg (Transvaal). Bull. Soc. geol. Fr., Paris (3) 24:349-378, pis.
15-18.
1902. Observations sur quelques Plantes fossiles des Lower Gondwanas. Palaeont.
indica, Calcutta (n.s.) 2 : 1-40, pis. 1-7.
19020. Flore fossile des Gites de Charbon du Tonkin. Etud. Gites min. Fr., Paris.
viii + 328 pp., pis. A-F ; atlas 56 pis.
GEOL. Ill, 4.
PLATE 18
Glossopteris fibrosa n. sp.
FIG. i. Epidermal pull showing a rather exposed stoma with lignine lamellae. ¥.344401!
x 400.
FIG. 2. Lower epidermis isolated with HF. The dark lignine lamellae of stomata are
obvious. There are two folds both exposing the inner side so that the stomatal pits appear
raised. ¥.34441. x 100.
FIG. 3. Upper epidermis showing pale cell walls and dark cell contents with granules of
various sizes. ¥.34442. x 450.
FIG. 4. Cuticle showing stoma. The polar and lateral lignine lamellae have dissolved by
maceration. ¥.34443. X400.
FIG. 5. Tracheids from the midrib. ¥.34440^. x 450.
Glossopteris hispida n. sp.
FIG. 6. Matrix in contact with the lower surface of leaf showing hairs when moistened with
oil. Holotype ¥.34450. x 10.
FIG. 7. Pull showing lower epidermis and a fibre between two veins. A few spongy meso-
phyll cells (dark rectangles) remain. ¥.344500. x no.
Bull. B.M. (N.H.) Geol. 3, 4
PLATE 18
GLOSSOPTERIS FIBROSA, G. HISPIDA
PLATE 19
Glossopteris fibrosa n. sp.
FIG. i. Epidermal pull showing stomatal aperture protected by thickened subsidiary cell
papillae. ¥.344490. x 800.
Glossopteris colpodes n. sp.
FIG. 2. Epidermal pull showing stoma protected by thickened subsidiary cell papillae.
¥.344660. x 800.
Glossopteris hispida n. sp.
FIG. 3. Epidermal pull showing stoma protected by subsidiary cell papillae. ¥.344500.
X 800.
Bull. B.M. (N.H.) Geol. 3, 4
PLATE 19
GLOSSOPTERIS FIBROSA, G. COLPODES, G. HISPIDA
PLATE 20
Glossopteris hispida n. sp.
FIG. i. Pull showing a one-celled hair base overlapping a number of epidermal cells.
¥.3445013. x 450.
FIG. 2. Pull showing a two-celled hair base overlapping epidermal cells. ¥.344500. x 450.
Scale Leaves
FIG. 3. Scale leaf isolated from rock with HF. ¥.34453 (i). x 6.
FIG. 4. Scale leaf isolated from rock with HF. ¥.34453 (2). x 6.
Rhabdotaenia harkini n. sp.
FIG. 5. Stoma protected by papillae of irregular subsidiary cells. ¥.344546. x 400.
Spermatites crystallinus n. sp.
FIG. 6. Outer cuticle showing crystals of various forms. ¥. 34455. X 400.
Glossopteris hispida n. sp.
FIG. 7. Upper cuticle showing numerous small surface papillae. ¥.344516. x 400.
Bull. B.M. (N.H.) Geol. 3, 4
PLATE 20
SPERMATITES, SCALE LEAVES, RHABDOTAENIA, GLOSSOPTERIS
PLATE 21
Sporangia
FIG. i. Dehisced sporangium showing outer cuticle with elongated cells (near stalk) and
inner cuticle with polygonal cells. Two- winged spores of varied size are seen inside the
sporangium. ¥.34456. x 40.
FIG. 2. Sporangia borne terminally on branches of an axis. V. 34457. X 8.
FIG. 3. Dehisced sporangium showing gaps between the wall cells. ¥.34458. x 40.
FIG. 4. Longitudinally flattened dehisced sporangium showing outer cuticle and a long
stalk. V.34459. x 40.
FIG. 5. Undehisced sporangium full of spores. ¥.34459. x 40.
FIG. 6. Two closed sporangia borne terminally on forks of a stalk. ¥.34460. x 40.
FIG. 7. A disc showing attached sporangia flattened in various planes. ¥.24233. x 6.
Figs. 1-6 from Mhukuru Coalfield, Tanganyika.
Fig. 7 from Newcastle, New South Wales.
Bull. B.M. (N.H.) Geol. 3, 4
PLATE 21
4
SPORANGIA
LIDGETTONIA,
A NEW 'YPE OF FERr LE
GLOSSOPTERIS
H. HAMSHAW THOMAS
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 5
LONDON : 1958
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following papers appeared in Volume I (1949-52) :
Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.
H. D. Thomas & A. G. Davis 75. 6d.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu ...... 55.
No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White 75. 6d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
Northern Iraq. L. F. Spath . . . . . . .105.
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum T. H. Withers 55.
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae).
T. H. Withers 55.
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian
of Podolia. W. N. Croft IDS.
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsda'le los.
No. 9 (1952). Australian Arthrodires. E. I. White .... 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard 6s.
The following papers appeared in Volume II (1953-56) :
No. i. (1953). The Deer of the Weybourn Crag and Forest Bed of
Norfolk. A. Azzaroli . . . . . . . £i 55.
No. 2 (1953). A Coniferous Petrified Forest in Patagonia. M. G. Calder 125.
No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark 35. 6d.
No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.
M. E. J. Chandler 165.
No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans . . 155.
No. 6 (1955). Further Contributions to the Solution of the Piltdown
Problem. J. S. Weiner, W. E. Le Gros Clark & K. P. Oakley et al. /i
No. 7 (1955). The Schizaeaceae of the South of England in Early Tertiary
Times. M. E. J. Chandler 155.
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . . £i
UDGETTQNIA,
A NEW TYPE OF FERTILE GLOSSOPTERIS
BY
H. HAMSHAW THOMAS, F.R.S.
Pp. 177-189 ; Pis. 22-23 / 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 5
LONDON : 1958
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they
become ready. Volumes will contain about three
or four hundred pages, and will not necessarily be
completed within one calendar year.
This paper is Vol. 3, No. 5 of the Geological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1958 Price Five Shillings
UDGETTQNIA,
A NEW TYPE OF FERTILE GLOSSOPTERIS
By H. HAMSHAW THOMAS
SYNOPSIS
Reproductive organs of two kinds have already been described attached to leaves of
Glossopteris. Cuticle studies suggest that this leaf is a form genus and that it was borne on
plants of several different types. The present paper describes leaves of another species which
are accompanied by fertile scale leaves bearing a number of delicate stalked cupules. The
cupules are empty but the matrix around them contains the remains of a very large number
of empty sporangia and of many small seeds. There was thus a third or perhaps a fourth
genus of plants with leaves of the Glossopteris type.
THE genus Glossopteris was instituted by Brongniart (1828) for leaves which were
entire, more or less lanceolate, with a midrib and fine secondary veins which showed
dichotomy and anastomosis. Two forms were mentioned as belonging to the new
group, G. browniana from Australia and India, and G. nilsoniana from Hor in
Sweden. Subsequently Brongniart added two further forms, G. angustifolia from
India and G. phillipsi from Yorkshire. Thus at the outset Glossopteris was a form
or artificial genus. The two European Mesozoic species were separated by Sternberg
(1838) and placed in the genus Sagenopteris ; they have proved to be leaflets of
members of the Caytoniales. The position of the remainder was uncertain, but the
study of their cuticular structure has shown that there are considerable differences"
in the epidermal cells of other species of Glossopteris. Zeiller (1896), Sahni (1923)
and Harris (1932) described the differing cuticles of three species, and recently
Srivastava (1956) has added considerably to our knowledge by descriptions of the
cuticles of fourteen additional species. Surange & Srivastata (1956) have suggested
that on this evidence six groups, possibly of generic rank, may be recognized. All
the epidermal structures suggest that the leaves known as Glossopteris and Ganga-
mopteris belonged to seed-bearing plants. Mrs. Plumstead (1952, 1956) has shown
that this view is true for five species, and although the preservation of her material
makes its interpretation very uncertain, it seems likely that the five species belong
to two genera which she has named Scutum and Lanceolatus.
The paper which follows shows that another species with Glossopteris leaves,
produced what must have been reproductive structures of a different type on small
leaves of the form called by some previous authors scale leaves. These are quite
distinct from Scutum and Lanceolatus, and merit the creation of a new genus.
Glossopteris was thus a leaf type belonging to several unrelated genera. It is still a
GEOL. Ill, 5. 15
i8o LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS
form genus and confusion is likely if it is used in the designation of a taxon of higher
rank as has been recently suggested (Plumstead, 1956^).
The material. The specimens here described were obtained from a narrow bed
of shale, 2-3 ft. thick, exposed in the sides and bed of a small stream on the estate
of Mr. S. Thomson at Lidgetton, about 35 kilometres north-west of Pietermaritz-
burg, Natal, South Africa. This plant bed was originally discovered by Mr. A. D. O.
Mogg, he kindly sent two large blocks to the author, who visited the locality in
1929* and made a collection of specimens, which have only recently been studied.
The plant remains occur in fine-grained shale which varies in colour and character
along the stream. In some places the rock is compact, dark in colour, and often
nearly black. The plants are well preserved as black carbonaceous compactions,
but are difficult to photograph because of the lack of contrast with the matrix (see
PI. 22, fig. i). At other places the matrix is coarser and light grey in colour, the
plant remains being less highly compressed. This again grades into fine-grained
material of a pale buff colour, in which the plant substance has entirely dis-
appeared, or is represented by ferruginous material lacking any trace of epidermal
structure (PL 22, fig. 3).
The macroscopic remains of plants in the accessible parts of this bed were of two
main types. A series of leaves of various sizes referable to the genus Glossopteris,
and including a number of shorter forms without a midrib, of the type called scale
leaves by previous authors. With these, towards the bottom of the bed, are parts
of equisetalean plants, stems, pith-casts, branches and leaves. These may belong
to the form called by Du Toit (1927 : 315) Neocalamites carreri Zeiller, but, from
the more abundant material present in this collection, it is clear that the plant
differed very considerably from the species described by Zeiller. In addition to
these larger remains there are very large numbers of detached sporangia scattered
in the matrix, and many isolated seeds of a uniform size and form.
Age of the specimens. The organic remains in the bed furnish no clear indications
of its age, and there is a heavy cover of soil and vegetation in the area. According
to the latest map published by the Geological Survey of South Africa the outcrop is
of Ecca age in the Karroo System. The boundary between the Ecca and the
Beaufort series is not far distant, and it is thus possible that the material is of Upper
Ecca age. But there is little lithological difference between the rocks of the Ecca
and those of the Beaufort series, as is stated by Du Toit, and it is also possible that
the forms described are of Beaufort age. In any event they are appreciably younger
than the fertile forms described by Mrs. Plumstead.
Genus LIDGETTONIA nov.
DIAGNOSIS. Sterile leaves of Glossopteris type, simple, lanceolate ; strong
tuberculate midrib almost to apex ; veins numerous, close, ascending ; forking and
anastomosing ; meshes narrow elongated, longer near midrib. Fertile leaves
distinct, short, spathulate-lanceolate ; midrib absent ; veins spreading from base,
* The author's thanks are due to the late Mr. J. A. Lidgett and to Mr. G. C. Lidgett who gave
valuable assistance in the re-discovery and working of the bed.
LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS 181
forking and anastomosing ; meshes considerably larger than in sterile leaves ; lower
(?) surface with two longitudinal rows of 4-6 small cupules on slender stalks, arising
from petiole or basal portion of leaf ; cupules open campanulate or disc-like, finely
striated, margins lobed.
Differences from comparable forms (Scutum, Lanceolatus) :
Several small cupules with slender stalks borne on small leaves differing from the
sterile leaves in size, shape and venation, instead of a single, large, bifid cupular
structure on a stout pedicel springing from the midrib or petiole of a leaf of normal
size and venation and containing massive fertile structures.
TYPE SPECIES. L. africana n. sp.
Lidgettonia africana n. sp.
(Plates 22, 23 ; Text-figs, i, 2)
DIAGNOSIS. As for genus.
HOLOTYPE. Fertile leaf with remains of cupules on either side of the petiole.
(Brit. Mus. N.H. No. V. 34633.)
DESCRIPTION. Sterile leaves : Most of the specimens are the remains of sterile
leaves (PI. 22, fig. i). From their simple, lanceolate form, their well-developed
midrib and their crowded anastomosing veins, they are clearly referable to the form
genus Glossopteris. They vary considerably in size, their venation is uniform, but
differs somewhat in appearance on the two sides of the leaf, and there is no reason to
suppose that more than one species is represented. No complete leaf has been found,
the longer pieces are about 15 cm. in length and 2-5-3 cm- broad in their widest part.
The lamina tapers very gradually towards the base, where the petiole was 4-5 mm.
broad. The length of the petiole is unknown, it is more than 2 cm. in some examples,
but in no case is its base visible. Towards the apex the lamina tapers to a blunt
tip. In the larger specimens the midrib is visible to within a short distance from
the apex. Below it merges without marked change into the petiole. As it becomes
broader it has a characteristic appearance due to the presence of 'small rounded
projections, 0-3-0-5 mm. in diameter, probably indicating the presence of groups
of hard cells in the original tissue. In addition, the well preserved specimens show
a series of slender longitudinal ridges, probably due to lines of thicker epidermal
or hypodermal cells.
The secondary veins are very numerous and crowded (Text-fig, i). They leave
the midrib at a very acute angle, and often run parallel to it at first for about 5 mm.,
then curving outwards they run steeply to the margin, diverging from the midrib
at an angle of 30°-40°. Between the midrib and the margin they show frequent
cross connections and they fork in an irregular way so that the number of veins
reaching the margin is rather more than double the number which leave the midrib.
Near the centre of the leaf the secondary veins are about 0-6 mm. apart, near the
margin they are only 0-3-0-4 mm. distant. The anastomoses of the veins are either
by small transverse veinlets, or by the fusion of veins which have come from dicho-
tomies at a lower level. The areolae are elongated and fusiform, becoming shorter
182 LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS
near the margin. In some hand specimens the anastomoses of the veins can only
be seen clearly in the lower part of the leaf and near the midrib. In the rather
rare cases where the lower surface of the leaf is seen the veins appear thicker and very
close together. The strength of the veins probably rendered possible the appearance
of the specimen represented in Text-fig, i ; here almost all of the tissue of the lamina
had disappeared leaving the midrib and the veins clearly seen in the matrix, but the
junction of the veins with the midrib was obscured in some places.
The surface of the upper side of well-preserved specimens shows, under the
binocular microscope, fine ridges above the secondary veins ; five such ridges are
visible following the course of the larger veins, some of them branching off when the
vein forks. Towards the margin of the leaf only one such ridge is seen. The outlines
of the epidermal cells are frequently visible, these cells were elongated above the
veins and rounded and isodiametric between the veins.
TEXT-FIG, i. — Part of a sterile leaf showing the venation. Drawn from a photograph of a
specimen in which only the midrib and the veins were preserved. Nat. size. ^.34639).
Although the preservation of some specimens appears very good it has proved
impossible, as yet, to obtain cuticle preparations, evidently the cuticle was very
thin and breaks into tiny fragments when stripped from the matrix.
Scale leaves. The leaves which have been described above are accompanied by
a number of smaller leaves, 2-6 cm. long and 1-1-5 cm- broad. They often show a
broad base and little or no trace of a midrib. Their veins are coarser, more distant
and show frequent anastomoses. Such structures have been described by previous
authors (see Arber 1905^) under the somewhat inappropriate name of scale-fronds.
They probably were organs intermediate between the normal photosynthetic leaves
and the fertile structures now to be described.
Fertile leaves. A large slab of grey shale, showing many compressed leaves,
and very similar to the specimen shown in PL 22, fig. i, contains the remains of
four shorter leaves close together (Text-fig. 2) . They appear to have been very thin
and present little contrast with the matrix. All are incomplete but three of them
are contracted at the base to form a petiole ; they show anastomosing veins but no
midrib. When examined in a strong beam of oblique light three of them were found
to bear small lateral appendages springing from their basal portions. These
structures are small campanulate bodies or peltate discs, 5-6 mm. in diameter,
LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS 183
borne on slender stalks about 5 mm. long and 0-6 mm. wide. They sprang from
the surface of the leaf and appear to have been produced in two rows, one on either
side of the longitudinal axis of the leaf. Fine striations, probably due to elongated
epidermal cells run from the stalks to the margins of the discs, which in the best
preserved specimens are seen to have a number of small lobes. One of the four
leaves in this group shows no appendages, but there are some small discs close to it
which may have sprung from the surface embedded in the matrix.
None of these structures contains any sporangia or seeds, but it seems highly
probably that they originally contained such reproductive organs. From a
comparison with the pteridosperms they may be termed cupules.
TEXT-FIG. 2. — Group of fertile leaves, showing stalked cupules. Nat. size.
A second specimen (PI. 23, fig. 4) shows a similar leaf preserved in a light grey
shale, the plant tissue being less compressed and altered. The apical part of the
lamina is not seen, at the top the lamina is 12 mm. broad, it tapers in a distance of
4 cm. to a petiole-like base, one and a half millimetres broad. In the expanded
portion fine veins run upwards and outwards from the basal region, there is no
midrib. As in the sterile leaves the epidermal cells above the veins were elongated
while those in the meshes were isodiametric and rounded. The petiolar portion
shows the remains of lateral appendages differing somewhat in appearance from
those described above, probably owing to the different preservation. Their stalks
are not clearly seen, they probably sprang from the lower side of the petiole. Parts
of four cupules can be detected on one side and parts of three on the other margin
of the petiole ; most of them are laterally compressed and are somewhat hemi-
spherical in outline. One example, 3 mm. in diameter, shows some of the marginal
184 LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS
lobes. In two places parts of seeds, like those to be described later, are seen in
contact with cupules ; they may possibly have been extruded when the original
structure became compressed in the matrix.
Another example shows a different type of preservation (PL 22, fig. 2). The plant
substance is converted to structureless red matter. The lamina has a different
shape, lacking the petiole-like base and the expanded upper portion ; it is 3 cm.
long, about 7 mm. broad at its widest part, and 5 mm. broad at the base. The
remains of six or possibly of seven cupules are seen on one margin, with indications
of two or three on the other margin. The stalks of several of them are seen on the
surface of the lamina but their exact place of origin cannot be determined. Some
of the cupules are conical in form, others appear as flattened discs. A well preserved
seed is seen lying in contact with one of the cupules, but its position may have been
accidental.
The form of the cupules is clearly shown in the specimen figured in PL 22, fig. 3.
Here the plant remains are reddish impressions or moulds in a light coloured matrix,
all the plant tissue has disappeared but the outlines of the original structures are
very clearly seen. The specimen and its counterpart show the impressions of seven
stalked cupular discs, which, from their positions, were probably attached to a
fertile leaf embedded in the matrix and not visible. The stalks of four of them are
more or less parallel and probably sprang from near one margin while three probably
came from near the other margin. The stalks are 07 mm. broad, they seem to
have expanded to form a somewhat thicker central portion of the cupule, about 3 mm.
in diameter, surrounded by a thinner zone, 2 mm. wide, with a lobed margin. The
surface of the outer zone shows a series of undulations associated with the lobes and
the whole is traversed by a series of fine radial striations. The cupules were
empty at the time of preservation, a seed is seen lying near to them. The same
hand-specimen contains also the remains of a scale leaf which is shown in the lower
part of PL 22, fig. 3. This is about 2 cm. long, narrow at the base but quickly
broadening to a width of i cm. and then rapidly contracting. No cupules are seen
attached to this leaf, but the remains of fragments of tissue and also surface irregu-
larities suggest that it may also have been fertile.
The specimens described show that associated with leaves of typical Glossopteris
form there were small (scale) leaves with a venation of the Gangamopteris type.
These structures often bore on their lower (?) surface rows of small and delicate
stalked cupular structures. It is very difficult to explain the presence of these
cupules except on the hypothesis that they formed the place of origin of the
reproductive structures of the plant. But there is at present no certain
direct evidence that this was true.
Sporangium-like organs. Arber (1905, 19050) described some characteristic
structures from several localities in New South Wales which he designated
sporangium-like structures. Almost identical structures are present in considerable
numbers in the Lidgetton shales associated with the plants just described. Isolated
examples occur abundantly, but they are also found in circular or elongated groups
about i cm. in diameter (PL 23, figs. 5, 6). The individual structures have a very
characteristic appearance, they are usually elliptical or ovoid in shape but sometimes
LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS 185
appear to have had one side flattened or concave, others are fusiform (PL 23, fig. 7).
They measure 1-1-5 mm. long and about 0-7 mm. broad. Well-preserved examples
show a series of longitudinal anastomosing ridges, these appear to have been specially
thickened cell walls which are more resistant than the rest of the wall substance to
maceration. Some of the specimens described by Arber show a " neck-like
prolongation " at one end, this was believed to be the part by which the sporangium
was attached to the organ on which it was borne. No evidence of a similar
prolongation has been found in the present specimens.
A number of these organs were removed from the rock and macerated in acid
oxidizing liquid. They dissolved away completely when transferred to dilute
ammonia leaving no trace of spores, but some of them contained a little of the matrix
material. Either the sporangia were completely empty when preserved or the
spores were not cutinized with material withstanding acid maceration. Several
of the fusiform specimens (like those shown in PI. 23, fig. 7) appear to have a
longitudinal slit probably representing the line of dehiscence. Though these
structures do not appear to contain spores the matrix contains large numbers of
winged spores of different sizes which are seen when pieces of the rock are dissolved
in hydrofluoric acid. Although there seems to be no proof that the structures in
question were sporangia or that they had any connection with the plants described
above, it seems difficult, in view of their large numbers, to regard them as uncon-
nected. They do not resemble the sporangia of Equisetalean plants, and in each
of the seven places where they have been found they are associated with Glossopteris.
But it is possible that they were derived from some plant not represented by
macroscopic remains in the present collection.
Isolated seeds. Mention has already been made of the presence of many isolated
seeds in close proximity to the sterile and fertile leaves. A considerable number
of these structures is present in the collection, and they are very uniform in size and
shape (PI. 22, figs. 2, 3 ; PI. 23, fig. 5). They are almost circular in outline, and
between 2 and 3 mm. in diameter, some of them are slightly flattened at their base.
They show two distinct portions, a central elliptical part evidently composed of
thick walled tissue, the sclerotesta, and a thinner peripheral part, or wing, which
does not seem to extend round the base of the seed. In the centre the sclerotesta
measures about 1-3 mm. across and the wing is about 0-8 mm. wide. Almost all
the examples have the same shape, which seems to show that the seeds were originally
flattened in form and that the rim round the sclerotesta had the form of a wing and
was not due to the squashing of a soft sarcotesta during preservation. Had the
original structures been spherical the appearance of the specimens would have varied
somewhat. In some specimens (PL 22, fig. 2) there is a slight indentation in the
wing at the apical end, in others the wing appears to extend beyond the apex of the
sclerotesta and shows a projecting point. No clearly defined micropylar canal can
be seen and there was no apical extension of the sclerotesta. No definite traces of
vascular tissue can be seen but in well preserved specimens there may be concentric
striations in the wing-like portion.
It would seem quite certain that these structures are the remains of seeds, and that
their size is such that they may have grown inside the cupular structures described
i86 LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS
above. But the specimens in the present collection do not establish this suggestion,
and much more material is needed.
COMPARISON WITH OTHER FORMS
A comparison of the sterile leaves from Lidgetton with those of the many species
that have been described is somewhat difficult and is probably untrustworthy in the
absence of cuticle preparations. In general form they resemble some of the
specimens that have been described as G. indica but they must belong to a different
type, probably to a distinct genus, because the fertile leaves are not at all similar
to those described by Mrs. Plumstead (1956) and found to bear reproductive organs
of the Scutum type.
They most closely resemble the specimens described and figured by Feistmantel
(1881) as G. communis. These had an acute apex, a gradually tapering lamina
at the base, closely crowded veins which run out to the margin at an acute angle,
especially near the tip of the leaf. The midrib was similar in being relatively broad
and distinct below but thinning out at the apex. Feistmantel noticed small rounded
projections on the midrib and the petiole, which he said had not been found on any
other species ; they are shown as rather smaller structures than those of the Lidgetton
leaves.
Another comparable form is the species named by Srivastava (1956) G. arberi,
but in this there is said to be not much difference in the size of the meshes at the mid-
rib and the margin, while in the present specimens the difference in size is very
noticeable.
It may be mentioned that the Lidgetton leaves are quite unlike the forms from
the Molteno beds of the Upper Umkomaas, Natal, which the author described under
the inapplicable name of G. longicaulis (Hamshaw Thomas, 1952) .x
The specimens which have been described above as fertile leaves have little in
common with the reproductive structures described by Mrs. Plumstead (1952, 1956)
from the Middle Ecca at Vereeniging. Although, owing to their mode of preservation
the morphological nature of these objects is still, in my view, uncertain, they were
large structures borne on a stout pedicel which arose from the midrib of a normal
foliage leaf. At the top of the pedicel was a cupule-like structure divided into two
halves, and containing a massive cone-like structure composed of a number of
closely packed bodies, probably containing seeds. The smallest specimen of Scutum
was more than three times the size of any of the cupules here described. The only
feature in common is that both were produced on foliar organs. In view of the
essential differences between the structures now discovered and those formerly
described, it seems desirable to make a new genus for their reception, in spite of
the fact that so many of the details of their structure are unknown. The name of
Lidgettonia is proposed for their designation.
It has already been mentioned that the sporangia seen at Lidgetton closely resemble
1 In transferring the plant which Du Toit (1927) had named Sagenopteris longicaulis to the form genus
Glossopteris, the fact was overlooked that Feistmantel had previously used the name longicaulis for a
different plant. Since the name was preoccupied it is proposed that the specimens from the Molteno
Series of the Upper Umkomaas described and figured by Hamshaw Thomas (1952) should be named
Glossopteris verticillata Thomas instead of Glossopteris longicaulis (Du Toit).
LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS 187
those described by Arber (1905) as sporangium-like organs. After the examination
of a large number of these structures in the present collection there seems to be no
reason for doubt that they are really the remains of sporangia. Our inability to
find in them the remains of cutinized spores may be due to the unsuitability of our
methods of treating the material or the spores may have been only lightly cutinized.
It may be noticed that the specimens described by Arber were also associated with
leaves of the normal type (G. browniana] and with " scale fronds ", they came from
two localities in New South Wales. He also referred to the discovery by Zeiller
(1896) of an example from the Transvaal among fronds and scale leaves of
Glossopteris.
Arber drew attention to the similarity between his sporangia and those of the
Recent cycad, Stangeria. He suggested the possible affinity of Glossopteris with
the Pteridosperms. Recent work gives strong support to this view. It may there-
fore be noticed that the sporangia may be further compared with those of the
pteridosperm Pteruchus, which occurs in the somewhat younger (Molteno) Beds of
Natal. Both are alike in size and shape, though those of Pteruchus have a broad
basal attachment. Both show longitudinal striations and a similar mode of
dehiscence.
Similar sporangia were described by Seward (1908) from Zululand, by Lundquist
(1919) from Brazil, and by Walkom (1928) from New South Wales, all associated
with Glossopteris leaves but without evidence as to their place of origin. But Du
Toit (1932) found in the Lower Beaufort beds of Natal examples of the same size
and form in close association with small spathulate structures which he named
Eretmonia natalensis. These were 15-35 rnm. long with a narrow stalk and a
spoon-shaped head ; anastomosing veins were faintly visible in the head. I have
recently examined these specimens by the kindness of Dr. Crompton, Director of
the South African Museum. This material also shows a graded series of leaves
referable to Glossopteris, the smallest being about 20 mm. long and 7 mm. broad
with a rounded apex and a narrow lamina. The sporangia are only seen in the
matrix near the Eretmonias, but there is no certain evidence of their attachment.
Poorly preserved remains of seeds, comparable in size to those fr6m Lidgetton,
also are present.
It seems very probable that Eretmonia was a fertile structure from another
plant with Glossopteris leaves, possibly G. cor data Feist. If this is so, Lidgettonia
and Eretmonia may well be related, though they could scarcely be regarded as
species of the same genus.
The isolated seeds described above are comparable in form to seeds found in other
places associated with Glossopteris and Gangamopteris , but they differ by being smaller
in size. Arber (1905(2) described winged seeds from India, Australia and South
Africa under the name Cardiocarpus, and Seward (1917) described somewhat similar
forms from a number of places under the name Samaropsis. Of the examples
described Samaropsis seixasi (White) from Brazil would seem to resemble the
present forms most closely. It seems to have had a sclerotesta 8-10 mm. long and
5 mm. broad, with a wing which completely surrounded it. Another small seed
from Tasmania was mentioned by Arber as having an oval sclerotesta, 5-5 mm.
i88 LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS
long with traces of a narrow wing. Walkom (1921) found winged seeds associated
with Glossopteris in a number of localities in Australia, which he called Nummulo-
spermum bowenense. This was a larger structure in which the sclerotesta had a
marked nucellar beak, and the sarcotesta, or wing extended all round the seed and
was broader in the micropylar region.
Although such winged seeds are often found associated with Glossopteris we have
no evidence of an original connection. Seeds of a somewhat similar form are known
to have been produced by plants of the Cordaites type, and Cordaitalean forms often
occurred in the Southern Hemisphere with Glossopteris. But it is far from certain
all the winged forms that have been found were derived from plants allied to
Cordaites. It seems likely that the seeds of structures like Scutum were set free and
dispersed when ripe, but we do not yet know their individual form.
CONCLUSION
The collection of Glossopteris leaves and the structures associated with them that
is here described, adds something to our knowledge of this puzzling form, but does
little to elucidate the morphological and taxonomic problems relating to this wide-
spread plant organ of Permian and Early Triassic times. It shows, however, that
in addition to the types which bore strobilus-like aggregates of reproductive structures
on stout pedicels springing from foliage leaves, there were other forms in which the
foliage leaves were accompanied by smaller fertile leaves. In these reproduction
was probably effected through the agency of structures produced in small and
delicate cupules borne on stalks on the fertile leaves. While there is no direct and
indisputable evidence as to the nature of the bodies borne in the cupules, there is
considerable likelihood that the cupules contained groups of small elongated sporangia
and that small seeds were borne in some of them ; both these structures occur in
considerable abundance in the matrix with the fertile leaves. Even if this suggestion
as to the probable nature of the actual reproductive organs is rejected as unproven,
it is clear that the plant remains at Lidgetton show certain fundamental differences
from the older types found in the Middle Ecca rocks at Vereeniging. These
differences support the view, based on the study of cuticles, that there were several
different groups of plants with leaves whose form and venation has led to their
inclusion in the form genus Glossopteris. This form of leaf may well have evolved
by parallel or even convergent development.
The problems raised by this work can only be solved by the discovery of much
well preserved material showing different stages in the growth of the structures
which have been described. When the vast extent of the beds containing
Glossopteris is remembered, the ultimate finding of such material does not seem
improbable. But a very careful search will be necessary as it has proved very easy
to overlook specimens like those described when collecting in the field.
REFERENCES
ARBER, E. A. N. 1905. On the sprangium-like organs of Glossopteris browniana. Quart. J.
Geol. Soc. Lond., 61 : 324-338, pis. 30, 31.
i 9050. Catalogue of the Fossil Plants of the Glossopteris Flora in the Department of Geology,
British Museum (Natural History}. Ixxiv -f- 255 pp., 8 pis. Brit. Mus. (Nat. Hist.),
London.
LIDGETTONIA, A NEW TYPE OF FERTILE GLOSSOPTERIS 189
BRONGNIART, A. 1828. Prodrome d'une Histoire des Vegetaux fossiles. viii + 223 pp. Paris.
1830. Histoire des Vegetaux fossiles. 1, 5 : 209-249.
Du TOIT, A. L. 1927. The Fossil Flora of the Upper Karroo Beds. Ann. S. Afr. Mus.,
Cape Town, 22 : 289-420, pis. 16-32.
— — 1932. Some Fossil Plants from the Karroo System of South Africa. Ann. S. Afr.
Mus., Cape Town, 28 : 370-393, pis. 39, 40.
FEISTMANTEL, O. 1881. Fossil Flora of the Gondwana System. Palaeont. indica, Calcutta
(2) 3, i (Suppl.) : 49-60.
HARRIS, T. M. 1932. The Fossil Flora of Scoresby Sound, East Greenland, 2. Medd. Gronland,
K0benhavn, 85, 3 : 1-112, pis. 1-9.
LUNDQUIST, G. 1919. Fossile Pflanzen der Glossopteris-Flora aus Brasilien. K. Svensk.,
Vet. Akad. Handl, Stockholm, 60, 3 : 1-36, pis. i, 2.
PLUMSTEAD, E. P. 1952. Description of Two New Genera and Six New Species of Fructifica-
tions Borne on Glossopteris Leaves. Trans. Geol. Soc. S. Afr., Johannesburg, 55 : 281-328,
pis. 43-52.
1956. Bisexual fructifications borne on Glossopteris leaves from South Africa. Palaeonto-
graphica, Stuttgart, 100, B : 1-25, pis. 1-14.
19560. On Ottokaria, the Fructification of Gangamopteris. Trans. Geol. Soc. S. Afr.,
Johannesburg, 59:211-236, pis. 33-39.
SAHNI, B. 1923. On the structure of the cuticle in Glossopteris angustifolia Brongn. Rec.
Geol. Surv. India, Calcutta, 54 : 277-280, pi. 17.
SEWARD, A. C. 1908. On a collection of Permo-Carboniferous plants from the St. Lucia
(Somkele) Coal-field, Zululand, and from the Newcastle district, Natal. Trans. Geol.
Soc. S. Afr., Johannesburg, 10 : 65-71, pis. 8, 9.
1917. Fossil Plants, 3. xviii + 656 pp. Cambridge.
SRIVASTAVA, P. N. 1956. Studies in the Glossopteris Flora of India, 4. Glossopteris, Gangam-
opteris and Palaeovittaria from the Raniganj Coalfield. Palaeobotanist, Lucknow, 5 : 1-45,
pis. 1-14.
STERNBERG, C. VON. 1838. Versuch einer geognostisch-botanischen Darstellung der Flora der
Vorwelt, 2 : 220 pp., 68 pis. Leipzig.
SURANGE, K. R. & SRIVASTAVA, P. N. 1956. Studies in the Glossopteris Flora of India, 5.
Generic status of Glossopteris, Gangamopteris and Palaeovittaria. Palaeobotanist, Lucknow,
5 : 46-49.
THOMAS, H. HAMSHAW. 1952. A Glossopteris with whorled leaves. Palaeobotanist, Lucknow,
1 : 435-438. pl- L
WALKOM, A. B. 1921. Nummulospermum bowenense gen. et. sp. nov. Quart. J. Geol. Soc.
Lond., 77 : 289-295, pl. 21.
1928. Notes on some additions to the Glossopteris Flora in New South Wales. Proc.
Linn. Soc. N.S.W ., 53 : 555-564, pl. 36.
ZEILLER, R. 1896. Etudes sur quelques plantes fossiles, en particulier Vertebraria et Glossop-
teris, des environs de Johannesburg (Transvaal). Bull. Soc. geol. Fr., Paris (3) 24 : 349-378,
pis. 15-18.
PLATE 22
Lidgettonia africana n.sp.
FIG. i. Large block of dark grey shale from Lidgetton, containing well preserved remains of
parts of sterile leaves. Groups of sporangia occur at the points marked by arrows, and some
isolated seeds are present, x 1/3. (¥.34637).
FIG. 2. Part of a fertile leaf in light buff shale. Remains of the stalked cupules are shown
on the right hand side, and a seed is seen in contact with one of them, x 3. (¥.34634.)
FIG. 3. Two series of impressions of stalked cupules are shown in the upper part of the
figure, which are believed to have sprung from one fertile leaf, which is not seen. An isolated
seed is in the centre of the figure, and a short " scale " leaf which may have been fertile is seen
below, x 3. (¥.34635.)
Bull. B.M. (N.H.) Geol. 3, 5
PLATE 22
LTDGETTONIA AFRICAN A
PLATE 23
Lidgettonia africana n.sp.
FIG. 4. Holotype. Fertile leaf with some preserved remains of tissues. The remains of
broken cupules are seen on either side of the petiole. x 2. (V. 34633.)
FIG. 5. Large group of sporangia. Remains of two seeds are shown at the bottom and on
the left hand side. x 6-5. (¥.34636.)
FIG. 6. Group of well preserved sporangia. x 10. (¥.34638.)
FIG. 7. Remains of two sporangia, which had probably dehisced before preservation. x 60.
(V.34638.)
Bull. B.M. (N.H.) Geol. 3, 5
PLATE 23
LIDGETTONIA AFRICANA
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THE FAUNAL SUCCESSION
IN THE CARADOC SERIES
OF SOUTH SHROPSHIRE
W. T. DEAN
BULLETIN OF
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THE FAUNAL SUCCESSION IN THE CARADOC
SERIES OF SOUTH SHROPSHIRE
BY
WILLIAM THORNTON DEAN
Pp. 191-231 ; Plates 24-26 ; 4 Text-figures
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Issued February, 1958 Price Fourteen Shillings
THE FAUNAL SUCCESSION IN THE CARADOC
1 SERIES OF SOUTH SHROPSHIRE
By WILLIAM THORNTON DEAN
SYNOPSIS
The history of the subdivision of the Caradoc Series in the type area is traced from the time
of Murchison to the present day. The faunally denned Stages proposed by Bancroft are
examined and redefined, and their relationship to established lithological units discussed. A
new interpretation of the correlation between shelly and graptolite faunas is suggested, with
some notes on the possible extension of the scheme to other areas.
I. INTRODUCTION AND ACKNOWLEDGMENTS
THE name " Caradoc " as applied to the succession of Ordovician rocks in south-east
Shropshire originated in 1839 when Murchison, in his great work The Silurian
System, gave the name " Caradoc Sandstone " to the strata cropping out along
the strip of country between the Wrekin in the north-east and Coston, near Clunbury,
in the south-west. Stratigraphically the beds lay between the igneous rocks forming
the Church Stretton Hills and the Wenlock Shales. The clearest section was said
by Murchison (1839 : 2I6) to be found in the valley of the River Onny near Horderley,
which may thus be taken as the " type " succession. Unfortunately the beds
assigned by Murchison to his " Caradoc Sandstone " included also horizons now
known to be Pre-Cambrian, Cambrian and Silurian in age, and the basal quartzites
of the Lower Cambrian were thought to be Caradoc Sandstone which had been altered
by the " igneous traps " of the Church Stretton range of hills.
It was not until 1854 that any detailed subdivision of the Caradoc Sandstone was
attempted, when Salter & Aveline published their classic results. Perhaps the most
important of these was the proving of the unconformity, displayed in the so-called
" Onny Section ", between what they termed the Trinucleus Shales (topmost
Caradoc) and the overlying Purple Shales (Upper Llandovery). They divided the
" Caradoc Sandstone " into five parts as follows, the youngest at the top of the table :
5. Trinucleus Shales
4. Flagstones of Cheney Longville, etc.
3. Sandstones of Horderley and Chatwall
2. Hoar Edge Grits
i. Shales of Harnage and Shineton
They failed to differentiate between the Shineton Shales (Tremadoc) and the
Harnage Shales (Caradoc), but the two were separated later by Callaway (1877 : 653)
who placed the Harnage Shales in their correct position above the Hoar Edge Grits,
GEOL. 3, 6. 16
194 THE CARADOC SERIES OF SOUTH SHROPSHIRE
and gave them the stratigraphical name by which they are now generally known.
In addition Callaway (p. 654) named Salter & Aveline's subdivision No. 3 the
Chatwall Sandstone.
In 1884 the Shropshire geologist J. D. La louche introduced the new names
Horderley Sandstone, Cheney Longville Flags and Onny Shales, corresponding to
Salter & Aveline's subdivisions 3, 4 and 5, and gave localities where the horizons
might be examined. Ten years later Lap worth & Watts (1894 : 320) erected the
name Acton Scott Beds for strata between the Cheney Longville Flags and the
Trinucleus Shales.
No further modifications or additions were made to the succession until 1916
when Lapworth proposed the name " Caradoc Series " for the Ordovician strata
concerned, and divided them into " Groups ", each being named after a locality
in south Shropshire. The Groups were subdivided further but Lapworth gave
neither type localities for his subdivisions nor any information regarding their
distinctive lithologies and faunas. Nevertheless the Geological Survey has attempted
to use Lapworth's subdivisions (Pocock et al., 1938 : 81-90), but these are in need
of more precise definition and will be examined in the following pages.
The most significant advance in our knowledge of Caradoc stratigraphy was made
in 1929 when B. B. Bancroft published the first of a number of papers on the sub-
division of the Series. All previous schemes of subdivision having been founded
on lithology, Bancroft pioneered the splitting of the Caradoc succession into units
defined by their contained faunas, in particular the brachiopods and trilobites.
He erected the three Stages, Soudleyan, Longvillian and Marshbrookian (19290 :
33-35), but later listed seven Stages, Girvanian, Harnagian, Soudleyan, Longvillian,
Marshbrookian, Actonian and Onnian in ascending order (19296 : table opposite
p. 76). Four years later (1933) appeared a series of tables showing the distribution
of the Stages Costonian to Onnian (the term Costonian not being denned, but
replacing Girvanian), and listing both lithological divisions and zonal fossils. Many
of the latter had not been described and were, therefore, nomina nuda. Additional
data relating to Bancroft's Stages were published posthumously (1945) ; a correlation
of the shelly and graptolitic faunas which was attempted is probably incorrect.
Much of Bancroft's earlier work did not receive the recognition it merited, due in
part to the fact that many of his zonal indices had not, at that time, been described.
In addition his results were marred by indiscriminate erection or suppression of
subdivisions without sufficient description, and sometimes by inadequate diagnoses
of critical fossils. Nevertheless, his pioneer work revolutionized research on the
detailed stratigraphy of the Upper Ordovician strata in much the same way as did
S. S. Buckman's on that of the Jurassic.
During recent work in south Shropshire Bancroft's major subdivisions, the Stages,
of the Caradoc Series have been found to be applicable, though some emendation
and redefinition are necessary in order to make the stratal classification more
effective. The Stage will serve as a basis for the following account of the subdivi-
sions, both large and small, of the Caradoc Series, and correlations with the graptolite
zones will be re-examined. Correlation Tables and Plates of critical fossils are
included.
THE CARADOC SERIES OF SOUTH SHROPSHIRE
SKETCH MAP OF
THE SOUTHERN
CARADOC AREA.
ORDOVICIAN ROCKS
UNSHADED
TEXT-FIG, i.
THE CARADOC SERIES OF SOUTH SHROPSHIRE
CARDINCTON HILL
\\\
CUTRTME*X+.
V . HOPE
M-BOWDLER
v
SOUDLEY + HOLLIES
CHELMICK
SKETCH MAP OF
THE NORTHERN
CARADOC AREA.
ORDOVICIAN ROCKS
UNSHADED
I MILE
TEXT-FIG. 2.
THE CARADOC SERIES OF SOUTH SHROPSHIRE 197
The field-work upon which these results are founded was carried out from the
University of Bristol during the tenure of a Post-Graduate Research Scholarship,
for the award of which I wish to express my thanks to the Shell Petroleum Co.
Ltd. Professor W. F. Whittard supervised the initial research, and I am also
grateful to him for much useful discussion, and for reading and criticizing this
manuscript.
II. THE STAGES OF THE CARADOC SERIES
(a) Costonian Stage
In the first instance, the strata constituting the Costonian were named the
Girvanian Stage by Bancroft (19296 : 67, table opposite p. 76), and divided into
five zones as follows : ^
5. Horderleyella plicata
4. Harknessella subquadrata
3. Harknessella subplicata
2. Reedolithus subradiatus
i. " Orthis " confinis
Of these, i and 2 were applied to the succession at Girvan, Ayrshire, 3 and 4
to the Cressage District, Shropshire, and 5 to the Horderley District, Shropshire.
Such a composite succession, founded on faunal provinces so distinct as those of
Girvan and south Shropshire, would appear to be unsafe ; it is not surprising that
when next Bancroft (1933) published the succession comprising the Stage, and at the
same time renamed the latter Costonian, he omitted the two lowest zones and retained
only those three from Shropshire.
The name Costonian implies in itself the use of Coston, near Clunbury, as the type-
locality, though Bancroft described it as being " typified by the grits of Horderley,
Hoar Edge and Coston in the East Shropshire area " (1945 : 182). At Coston the
succession is as follows :
U. Costonia ultima Beds nom. nov.
Coston Beds nom. nov. M. Harknessella Beds nom. nov.
L. Basal Conglomerates
The type-area for the three new stratigraphical terms in the above table is the
general vicinity of Coston Farm, one mile east-south-east of Clunbury.
No fossils have yet been found in the Basal Conglomerates. The succeeding
Harknessella Beds contain an abundant brachiopod fauna of Harknessella vespertilio
(J. de C. Sowerby), H. jonesi Bancroft, Dinorthis flabellulum (J. de C. Sowerby)
and Heterorthis patera (Davidson), occurring in lenticular shell-beds. The Costonia
ultima Beds represent Bancroft's Zone of Horderleyella plicata, and consist of thickly-
bedded sandstones throughout which the fossils are distributed more or less uniformly.
Horderleyella plicata Bancroft occurs throughout, though not commonly, but has
not yet been found in significant numbers outside the Coston-Horderley District,
and may prove to be no more than a local species. Costonia ultima suffers from
similar limitations to its distribution, but the genus is more widespread. The
ig8 THE CARADOC SERIES OF SOUTH SHROPSHIRE
topmost Costonian beds between the Onny Valley and Brokenstones have also
yielded Lichas (s.l.) sp. and Salopia salteri (Davidson) ; the latter is predominantly
a Harnagian species and a few specimens only are known from the Costonian.
Although Bancroft's Correlation Tables of 1929 and 1933 showed no subdivision
of the H. plicata Zone, in a later paper (1945 : 235, 244) he referred to two subzones
within it, those of Smeathenella strophomenoides Bancroft and Dinorthis robusta
Bancroft. The order of superposition of these two was not at that time stated,
but later Bancroft described the former as being the younger (1949 : 297). The
usefulness of such a subdivision is doubtful, at least outside the Coston District,
because the subzonal brachiopods have now been found associated at Brokenstones
Quarry, a few miles north of Coston. Costonia ultima is present in the Upper Coston
Beds of both Coston and Brokenstones, and constitutes probably the most useful
zonal index but only within those districts.
In the northern part of the Caradoc Area the detailed succession within the lower
beds of the Caradoc Series can be summarized as follows :
3. Rhynchonellid Grits
2. Sandy limestones with Harknessella subquadrata
i. Sandy shales and limestones with Harknessella subplicata
The lowest strata, constituting Bancroft's Harknessella subplicata Zone, contain
a fauna of few species which, in addition to the zonal brachiopod, includes Reacalymene
pusulosa Shirley and Rafmesquina, the interbedded shale-bands having yielded
Nemagraptus gracilis (Hall). Beds i to 3 were placed in the Costonian by Bancroft
(1933) who supposed them all to be older than the Horderleyella plicata Zone of Coston
and Horderley. This age-relationship is considered here to be incorrect ; Bancroft
gave no explanation but he probably relied on the supposed equivalence of the strata
with abundant Harknessella at Cressage and Coston, even though no species is common
to both districts. Such a correlation, founded solely on large, shallow-water brachio-
pods apparently confined to marginal marine deposits, appears to be unreliable.
Near Harnage the trilobite fauna of the H. subquadrata Zone presents a picture
quite different from that of the preceding H. subplicata Zone. Decoroproetus
[Proetidella Bancroft], Brongniartella, Eohomalonotus , Lichas (s.l.) and Costonia
sp. nov., of which the first two become more abundant in the basal Harnagian of the
Onny Valley, are associated with Reacalymene pusulosa. The topmost beds of the
so-called Hoar Edge Grits, that is those succeeding the H. subquadrata Zone and
termed the Rhynchonellid Grits, are imperfectly exposed in the Harnage area,
but at Stevenshill they contain, in addition to Salopia salteri and rhynchonellids,
the diagnostic cryptolithid Salterolithus, and must be included in the Harnagian
Stage, even though they immediately underlie the Harnage Shales (s. s.). There is
no evidence whatsoever for the assertion that a considerable break exists between
the Hoar Edge Grits and Harnage Shales of the Harnage area (Pocock et al., 1938 :
86), and even the H. subquadrata Zone may best be regarded as showing signs of
a transition towards the basal Harnagian, though the horizon must still be retained
in the Costonian on account of the presence of Costonia and the absence of
Salterolithus and Reuscholithus.
THE CARADOC SERIES OF SOUTH SHROPSHIRE
199
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DALMANELLA UNGUIS
DALMANELLA WATTS 1
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200 THE CARADOC SERIES OF SOUTH SHROPSHIRE
It can thus be claimed, with some degree of certainty, that the H. subquadrata
Zone of Cressage and Harnage is the equivalent of the Costonia ultima Beds (or
Horderleyella plicata Zone) of Cost on, at least in part. The H, subplicata Zone of
Harnage may reasonably be equated with the Harknessella Beds of Coston. The
strata containing abundant Harknessella would appear, then, to be diachronic,
becoming progressively younger from south to north, and it is probable that the
Basal Conglomerates of Coston may be the oldest members of the Caradoc Series
in the whole Caradoc Area, and unrepresented in the Harnage District.
The so-called Hoar Edge Grits of the Cwms, east of Church Stretton, represent
what is probably only a small portion of the Costonian succession, as trilobites
found therein include Decor oproetus together with Costonia sp. nov. identical with
that found in the H. subquadrata Zone of Harnage. The presence of S alter olithus
in the same area indicates a Harnagian age for, probably, the uppermost part of the
Hoar Edge Grits there.
(b) Harnagian Stage
The Harnagian was proposed by Bancroft (19296 : 67) and was said by him to
extend from the summit of the Girvanian to the base of the Soudleyan. It was
described merely as being characterized " by the caractaci group of Cryptolithus
s.L" (later S alter olithus}. In the same paper (table opposite p. 76) he divided the
Harnagian into four zones as given below, the two lower being described as occurring
at Cound Brook (near Cressage) and Trilobite Dingle, Welshpool, and the two upper
at Horderley and Trilobite Dingle.
Salterolithus caractaci
Salterolithus cf. intermedius
Reuscholithus reuschi
Salterolithus harnagensis
In a later work, however, Bancroft (1933) included only three zones in the Stage,
the zonal trilobite of the topmost of these being at that time undescribed and,
until 1949, unfigured :
Ulricholithus ulrichi
Salterolithus caractaci
Reuscholithus reuschi (misprinted as Reuschella [sic] reuschi)
It was in 1945 that Bancroft denned the type-locality of the Harnagian, stating
it to be in the " Harnage Shales of Cound Brook, Cressage ", but he also denned the
base as being marked by the appearance of Reuscholithus and Salterolithus.
Like those of the underlying Costonian, the strata assigned to the Harnagian Stage
in south Shropshire are marked by changes in lithology along the strike, with their
attendant variations in fauna. A further complication is introduced by Bancroft's
fixing of the type-succession as the highly-faulted area of "Harnage Shales" in Cound
Brook, near Evenwood, implying the inclusion of only the shale lithology at that
locality. Salterolithus is described by Bancroft (1945 : 182) as being a genus
THE CARADOC SERIES OF SOUTH SHROPSHIRE 201
especially characteristic of the Harnagian, and in the Evenwood District it first
appears in sandstones immediately and conformably overlying others of definite
Costonian age (see p. 198). It would seem advisable, therefore, to redefine the base
of the Harnagian and to describe it as being marked by the appearance of the trilo-
bite genera Reuscholithus and/or Salterolithus.
In the basal Harnage Shales of the Cound Brook section Reuscholithus reuschi
Bancroft is abundant and occurs at, apparently, the same horizon as Salterolithus
harnagensis Bancroft, both species inhabiting grey mudstones and, more rarely,
sandy shales. Farther south, however, near the Onny Valley, R. reuschi is relatively
uncommon, though attaining a larger size than at Cound Brook, and the basal horizon
of the Harnagian is marked by an abundance of Salterolithus smeathenensis Bancroft
and Phacopidina harnagensis Bancroft. S. harnagensis is not known from this area,
and it is not surprising that Bancroft found it advisable to discard the 5. harnagensis
Zone in favour of using the single index R. reuschi for this particular horizon. As
yet Phacopidina harnagensis, together with the uncommon accompanying form
Nieszkowskia stubblefieldi Bancroft, has not been recovered from the Harnagian of
Cound Brook, but Primaspis \Acidaspis} harnagensis (Bancroft) occurs there as well
as near the Onny, together with rare Decoroproetus [Proetidelld] fearnsidesi (Bancroft) .
It has already been shown (p. 200) that the topmost part of the Hoar Edge Grits
in The Cwms, east of Church Stretton, contains trilobites of Harnagian age, and on
the nearby Hazier Hill basal Harnagian deposits form the infillings of the well-known
neptunian dykes there. Details have been given of the fauna from these beds
(Strachan et al., 1948), but as long ago as 1929 Bancroft had stated that they were
of the same age as his Zone of Salterolithus harnagensis, that is, basal Harnagian
(19296 : 81). Apart from the large number of contained pebbles of Uriconian
material, the lithology of the Hazier Harnagian beds is closely similar to that of the
R. reuschi Zone in the Onny Valley, and the two may reasonably be equated. Certain
differences exist, probably as the result of a shallower-water environment at Hazier,
and the fauna there, while lacking most of the trilobites so abundant in the* Onny
Valley, includes the Baltic brachiopod genus Vellamo, a record new to Shropshire.
Bancroft's Zone of Salterolithus caractaci denotes an horizon stratigraphically
higher than that of Reuscholithus reuschi. Although Murchison's syntypes were
obtained from the Welshpool District, the zone was first established as a usable
stratigraphical horizon in the vicinity of Glenburrell Farm, near Horderley (Bancroft,
1949 : 294). A large collection was obtained by Bancroft from the excavations that
were dug here, and the material shows that there exists at this horizon in the
Horderley District a large cryptolithid fauna consisting of one species, which seems
best referred to Salterolithus caractaci (Murchison), though the latter has yet to be
described adequately. Murchison's species has not been discovered in association
with R. reuschi and its attendant fauna, and it seems advisable to retain the 5.
caractaci Zone as a separate unit. The state of preservation of the material from
Bancroft's excavations is closely similar to that of the R. reuschi Zone in the Horderley
District, and the S. caractaci Zone can be taken to represent the upper part of the
yellow, blocky mudstones of Harnagian age which crop out at Smeathen Wood,
just south of the Onny, and for which the name Smeathen Wood Beds nom. nov.
202 THE CARADOC SERIES OF SOUTH SHROPSHIRE
is here proposed. These comprise the strata in the Onny Valley belonging to the
Reuscholithus reuschi and Salterolithus caractaci Zones, and the type-locality is the
general area of Smeathen Farm, near Horderley.
The 5. caractaci Zone formed the highest subdivision of the Harnagian as
interpreted by Bancroft (19296, table opposite p. 76), but in a later paper (1933)
he introduced an overlying Zone of Ulricholithus ulrichi. Unfortunately the zonal
trilobite was not then figured or described, and the name remained a nomen nudum
until sixteen years later (Bancroft, 1949 : 295, pi. 9, fig. 14), although Bancroft had
cited the species from Shropshire, Welshpool and North Wales (1933). He described
Welshpool (1949 : 296) as being " the more important type locality " ; the text
suggests that he had intended to formulate the species on a number of syntypes from
Welshpool and from south Shropshire, but only an incomplete cephalon from Welsh-
pool was figured by him ; this is the holotype (British Museum (Nat. Hist.) In.
42371). In the Caradoc Area the species is known from only two localities, both
near Glenburrell Farm. Once again, a single crypt olithid species constitutes the
whole trilobite fauna, and the writer accepts provisionally the existence of an
Ulricholithus ulrichi Zone marking the horizon between the 5. caractaci Zone and the
base of the succeeding Soudleyan Stage, though the lateral distribution of the species
in the remainder of the Caradoc Area has yet to be demonstrated. In order to do
this, extensive excavations would be necessary as the strata, consisting of easily-
weathered shales and mudstones, are almost invariably poorly exposed. Near
Horderley they include the lower part of the Glenburrell Beds, comprising dark
green mudstones and shales which are a continuation of the mudstone phase of
sedimentation commenced in the Smeathen Wood Beds. The Ulricholithus ulrichi
Zone is retained in the Harnagian because it contains no specimens of Broeggerolithus,
the incoming of which genus indicates the base of the Soudleyan Stage.
Harnagian strata crop out elsewhere in the Caradoc Area, but the faunas are not
so completely known as in the districts already mentioned. At the southern end
of Ragleth Hill shales and limestones of Harnagian age rest unconformably on
Western Longmyndian sandstones, but their exact zonal position is difficult to deter-
mine owing to the scanty numbers and poor preservation of the fossils.
Grey sandy shales exposed in the stream-section west of Wallsbank, on the south-
eastern flank of Cardington Hill have yielded well-preserved specimens of Reuscholithus
reuschi, and other, more fragmentary, material from the same neighbourhood
suggests that at least one higher horizon may be present.
(c) Soudleyan Stage
The type-locality of the Soudleyan is not, as the name implies, at Soudley itself,
but was defined by Bancroft (1945 : 182) as being " in the Onny Valley beginning
in the Glenburrell Beds and terminating at the summit of the middle Horderley
Sandstone ". The reason for this apparent anomaly is, no doubt, the superior
series of exposures to be found throughout the Stage in the Onny Valley.
The name was first proposed by Bancroft (19290 : 33-34) to include " the highest
beds of the Harnage Shale and the lower half of the Horderley Sandstone in
Shropshire ". Faunally it was described as " corresponding to the upper part of
THE CARADOC SERIES OF SOUTH SHROPSHIRE 203
the series with the expansa group of Rafinesquina, and especially characterized by
a group of species of Cryptolithus in which the features of the gibbifrons group are
modified or not fully developed ".
Later in the same year (Bancroft, 19296, table opposite p. 76) the Soudleyan was
listed in a table and divided into four zones as follows :
Dinorthis sp.
Reuschella horderleyensis
Cryptolithus sp.
Cryptolithus broeggeri
In 1933 Bancroft gave a rather more detailed succession for the Stage, dividing
it into the following five zones :
Dinorthis multiplicata
Cliftonia persculpta
Broeggeria [sic] soudleyensis and Reuschella
Heterorthis retrorsistria, Broeggeria [sic]
Horderleyella and Broeggeria [sic]
The type-specimens of D. multiplicata Bancroft derived from near Glyn Ceiriog,
where the species is relatively abundant and occurs with Rafinesquina expansa.
It is not common in south Shropshire, and cannot be considered a satisfactory
zonal fossil there. Of the other forms, Cliftonia persculpta is a nomen nudum,
and Bancroft's genus Broeggeria has since been replaced by Broeggerolithus (Lamont,
1935 : 320). The two lowest zones were equated with part of what Bancroft termed
the Glenburrell Beds, and the three highest with part of the Horderley Sandstone.
In the Onny Valley to the east of Horderley the series of mudstones already
described under the Harnagian Stage continues upwards as far as the base of the
Horderley Sandstone. The beds are well exposed in the sections behind and .near
Glenburrell farmhouse, which may thus be taken as the type-locality of the Glen-
burrell Beds. These consist of dark green mudstones, similar to those with Ulricho-
lithus ulrichi at the top of the Harnagian. At Glenburrell they contain abundant
Broeggerolithus broeggeri (Bancroft). The beds crop out in the bed of the Onny
south-east of Glenburrell, and for a short distance to the south of the river. B.
broeggeri is fairly common, and a good zonal index for the horizon ; it is accompanied
by the characteristic brachiopod Soudleyella [Onniella] avelinei (Bancroft), with
Brongniartella and a new, though uncommon, species of S alter olithus. When
traced upwards in the Onny Valley the Glenburrell Beds become rather more
arenaceous, and it is believed that these levels constitute Bancroft's Zone of
" Heterorthis retrorsistria, Broeggeria " and, perhaps, correspond to the term
" Horderleyella Beds " used by him in his description of Horderleyella corrugata
but never defined (1945 : 238). The diagnostic species of cryptolithid is still B.
broeggeri, and Horderleyella corrugata ranges upwards just into the lowest sandstones
of the overlying Horderley Sandstone.
In turn the more arenaceous members of the Glenburrell Beds pass upwards
into the lowest, flaggy strata of the Horderley Sandstone group, known to many
204 THE CARADOC SERIES OF SOUTH SHROPSHIRE
geologists as the " Glyptocrinus Flags " on account of the abundance of the remains
of the crinoid Rhaphanocrinus [Glyptocrinus] basalis (McCoy). In the Onny Valley
these comprise greenish-brown, flaggy sandstones with lenticular shelly bands
in which brachiopods, especially Dinorthis, Sowerbyella, Rafinesquina expansa
and Reuschella horderleyensis Bancroft, predominate. Of the trilobites the most
important is Broeggerolithus soudleyensis (Bancroft). Early forms of this species
occur rarely in the B. broeggeri Zone (Bancroft, 1935 : 33), but B. soudleyensis
occurs in abundance over most of the Caradoc Area in the zone to which it gives its
name, and then disappears. The disappearance of this stock was only temporary,
as related forms reappeared later, though not in large numbers, in the Alternata
Limestone of the Upper Longvillian Substage (see p. 207).
As stated earlier, Cliftonia persculpta is a nomen nudum, and neither the C.
persculpta Zone nor the Dinorthis multiplicata Zone of Bancroft has been found to
be particularly valuable as a subdivision of the Soudleyan in south Shropshire. It
is preferred here to make one Broeggerolithus soudleyensis Zone for the whole of the
upper sandy portion of the Soudleyan in the Onny region, with Reuschella horder-
leyensis acting as a supplementary index (see table on p. 199). The B, soudleyensis
Zone is considered to be broadly equivalent to the Lower Horderley Sandstone, the
latter being used in a restricted sense, corresponding to the flaggy sandstones of the
Glyptocrinus Flags of the Onny Valley. Bancroft used the term Lower Horderley
Sandstone in a broader sense which is difficult to follow exactly in the field, including
as it does both the flaggy beds and some of the overlying massive sandstones.
The most southerly outcrops of the Soudleyan in the Caradoc Area are found near
Sibdon Carwood, west of Craven Arms, where quarries were opened in the
" Glyptocrinus Flags ". These have yielded Broeggerolithus soudleyensis, Soudleyella
and poorly-preserved graptolites. The Glenburrell Beds are not exposed in this
area, but may be seen in the valley to the north-east of Hopesay Hill.
North of the Onny Valley, strata of Soudleyan age crop out near Brokenstones,
but beyond this point they are cut out by faulting. They reappear near the southern
end of Ragleth Hill, near Little Stretton, but only the upper arenaceous beds have
been examined, the junction with the underlying Harnagian not being exposed.
Similarly, near Soudley the lowest beds are not seen, but at Soudley Quarry the
well-known purple and green sandstones are of Soudleyan age. These yielded the
type specimens of Broeggerolithus soudleyensis and, as they are disconformably
overlain by the Alternata Limestone (Upper Longvillian), the often used term
" Soudley Sandstone " must be restricted to the upper part of the Soudleyan.
Palaeontological evidence suggests that the diachronism known from the earlier
Caradoc strata is repeated in the Soudleyan, in the district north of the Cardington
Hills. In the Onny Valley the lower zone of the Soudleyan, that of B. broeggeri,
consists of dark green mudstones, followed by sandstones of the Glyptocrinus Flags
(B. soudleyensis Zone). Both near Chatwall and The Cwms typical Glyptocrinus
Flags are found to contain a form of Broeggerolithus close to B. broeggeri with
Rafinesquina expansa and Soudleyella cf. avelinei. This fauna suggests an horizon
somewhat earlier than that of the Zone of B. soudleyensis, a species not yet found in
these beds.
THE CARADOC SERIES OF SOUTH SHROPSHIRE 205
(d) Longvillian Stage
The name was first employed by Bancroft (19290 : 34), who also termed it "Kjaerina
Stage " and indicated that the lower and upper portions had distinct faunas ; his
description of the succession may be summarized as follows :
E. Middle Longville Flags
E2. Kjaerina geniculata Beds or Zone . . . . ... 120 ft.
Ei. Laminated micaceous flagstones . . . . . 165 ft.
D. Lower Longville Flags
Di. Heterorthis alternata Beds . . . . . . . . no ft.
(Kjaerina bipartita Zone)
C. Upper Horderley Sandstone
€7. Green sandstones. Wattsella sp. D Super-zone
C6. Green and buff sandstones. Wattsella sp. C Zone . . . 12 ft. -f-
€5. Grey sandstone. Kjaerina hedstroemi Zone ..... Few ft.
€4. Greenish and buff-grey sandstones. Kjaerina intermedia Zone . Few ft.
€3. Grey sandstone. Wattsella sp. A Super-zone . . . c. 25 ft.
€2. Green and purple sandstone . . . . . . 12 ft.
Ci. Buff sandstone. Wattsella sp. A Super- zone . . . . 18 ft.
Later in the same year Bancroft (i929&,table opposite p. 76) correlated some of these
horizons with strata in North Wales, and divided the Longvillian into six " Zones
and Super-zones " as follows :
Kjaerina geniculata Bancroft
Kjaerina bipartita (Salter)
Wattsella sp. D
Wattsella sp. C
Kjaerina hedstroemi Bancroft
Wattsella sp. A
Including as it did part of the Horderley Sandstone, together with the Alternata
Limestone and the Lower Cheney Longville Flags, the Longvillian constituted a
major and somewhat cumbersome subdivision of the Caradoc Series, and in a later
work (1933) Bancroft found it convenient to subdivide the Stage into Lower and
Upper Longvillian Substages. The dividing line was drawn at the base of the
Alternata Limestone, and in the present account the two parts will be discussed
separately.
Lower Longvillian Substage
When this subdivision was used by Bancroft for the first time, the previously
lettered species of Wattsella were given names, though nomina nuda, and the following
zones proposed :
Raymondella typa
Wattsella indica
206 THE CARADOC SERIES OF SOUTH SHROPSHIRE
Wattsella hpta
Wattsdla horderleyensis and Kjaerina
The brachiopod faunas of the Caradoc Series in south Shropshire and part of
North Wales were described by Bancroft in 1945. He did not recognize the fact
that certain of the species, such as Dalmanella horderleyensis, D. indica and Bancrof-
tina typa, had been described for the first time by Whittington (1938*2, 19386) who,
although the names had first been used in manuscript by Bancroft, thus became their
author.
In the present account it is recognized that a three-fold division of the Lower
Longvillian can be established on the basis of the brachiopods as follows :
Bancroftina typa (Whittington)
Dalmanella indica Whittington and D. lepta (Bancroft)
Dalmanella horderleyensis (Whittington)
The beds with Dalmanella horderleyensis constitute the best-known strata of the
Horderley Sandstone group, viz., the massive purple and green sandstones of Long
Lane quarries, north-west of Craven Arms, and of the Onny Valley, east of Glenburrell
Farm. The characteristic dalmanellid is abundant in lenticular limestone bands,
with less common Kjaerina jonesi Bancroft and gastropods such as Sinuites and
Lophospira cf. gyrogonia (McCoy). The overlying sandstones were divided into the
separate zones of Dalmanella lepta and D. indica by Bancroft, but the evidence for
the two distinct horizons is not altogether satisfactory and, for the present, it is
proposed to group them together as one. The Shropshire syntype of D. indica
came from Longville Plantation, whilst the type specimens of D. lepta were obtained
from near the south bank of the River Onny, a mile or so to the north along the strike
from Longville Plantation, and the division of the beds into two separate entities
cannot be accepted as satisfactorily established. The succeeding beds are charac-
terized by Bancroftina typa (Whittington), though Kjaerina also occurs in some
numbers. A conspicuous feature of the fauna of the Lower Longvillian is the
extraordinary abundance of Sowerbyella soudleyensis Jones, a form which occurs
throughout the Substage and constitutes a large proportion of the lenticular
limestones.
Tribolites are not uncommon in the Lower Longvillian, and include Brongniartella,
Eohomalonotus (rare), Phacopidina apiculata (Salter) and Reacalymene. The most
characteristic form is Broeggerolithus gloUceps (Bancroft) which occurs throughout
most of the Substage and is now taken as the zonal index. Bancroft recorded
Platylichas laxatus from the highest strata with Bancroftina typa, but intensive
searching has failed to substantiate this claim.
The sandstones with Dalmanella horderleyensis constitute the best-known building
stone in the Onny Valley district, but at higher faunal horizons the beds become
less suitable for building purposes, and are noticeably more flaggy, marking a transi-
tion to the overlying Upper Longvillian. Farther north, in the Soudley District,
the latter beds rest upon sandstones of Soudleyan age, the whole of the Lower
Longvillian being cut out by overstep.
THE CARADOC SERIES OF SOUTH SHROPSHIRE 207
Upper Longvillian Substage
In his first usage of the Substage Bancroft (1933) divided the beds into three
brachiopod zones :
Kjaerina geniculata Bancroft
Raymondella gigantea — manuscript name
Kjaerina bipartita (Salter)
Of these, Raymondella gigantea is still an undescribed species and, hence, a nomen
nudum. In practice it has been found most convenient to divide the Upper Long-
villian into only two parts. The lower is represented by the Alternata Limestone,
so-called from the abundance of Heterorthis alternata (J. de C. Sowerby), a species
which forms lenticular limestones at this horizon throughout the Caradoc Area.
The same brachiopod is known from higher horizons in south Shropshire and, for
zonal purposes, Bancroft's practice of employing Kjaerina bipartita is preferred.
Other brachiopods which appear to be restricted to this horizon are Bancroftina
robusta (Bancroft) and Marionites typa (Bancroft), though not in such abundance.
The Alternata Limestone is not a single calcareous band, but consists of imper-
sistent shelly lenses separated by dark green flaggy sandstones and siltstones.
Passing upwards, the limestone lenses die out, and there is a transition to the rather
monotonous series of green flaggy sandstones, known generally as the Lower Cheney
Longville Flags. Fossils occur mainly on bedding-planes throughout the group ;
Kjaerina typa Bancroft is abundant and is considered to be commoner than K.
geniculata, the index-species chosen by Bancroft. Other brachiopods found in
moderate abundance include Dolerorthis duftonensis (Reed) and a species of
Bancroftina (possibly Bancroft's Raymondella gigantea} , both of which occur through-
out the beds with Kjaerina typa.
Trilobites are frequent in parts of the Upper Longvillian but only one form can
be considered suitable as the zonal index. This is Broeggerolithus longiceps (Bancroft)
a species which is more abundant in the flaggy siltstones separating the shelly lenses
of the Alternata Limestone than in the lenses themselves. B. longiceps persists
through the higher beds of the Upper Longvillian, but in reduced numbers.
Phacopidina apiculata (Salter) is common in the Alternata Limestone, as is
Brongniartella bisulcata (Salter), the forma typica of which appears for the first
time at this horizon. Chasmops makes its first appearance in Shropshire in the
higher beds with Kjaerina typa, and individuals of both this genus and of B. bisulcata
often attain large sizes. An interesting feature of the trilobite fauna of the Upper
Longvillian, at least in the Soudley District, is the reappearance of a form of
Broeggerolithus, closely allied to B. soudleyensis (Bancroft), in which several pits
of the outermost or £2 row on the fringe are missing in front of the glabella. Nothing
like it is yet known between the Upper Soudleyan and the Upper Longvillian in
south Shropshire, nor from any strata later than the Upper Longvillian. In the
Llansantffraid-ym-mechain District of Montgomeryshire Whittington (i938c : 436,
451) records " Cryptolithus soudleyensis " from beds which are Lower and Upper
Longvillian in age.
GEOL. 3, 6. 1 8
208 THE CARADOC SERIES OF SOUTH SHROPSHIRE
The Upper Longvillian follows the Lower Longvillian with apparent conformity
in the Onny Valley, but farther north, at Soudley, it rests on the upper part of the
Soudleyan. At Chatwall the Upper Longvillian may include at its base both
conglomerates and sandstones which have been classified in the broad term of
" Chatwall Sandstone " ; this problem is discussed later (see p. 216).
(e) Marshbrookian Stage
The Marshbrookian Stage was first adopted by Bancroft, who also gave the
alternative name of Kjerulfma Stage, named after what he considered to be the
typical brachiopod genus (19290: : 34). The lower limit was taken arbitrarily at the
base of what he called the Wattsella wattsi Zone, owing to the fact that Kjerulfina
appeared just below the summit of the Longvillian Stage. In the same paper
(p. 39) he divided the Stage, which was described as being represented by the upper
130 ft. of the Cheney Longville Flags in east Shropshire, as follows :
F4. Transition Bed (Kjerulfina polycyma Zone)
F3. Wattsella unguis Beds (W. unguis Super-zone)
F2. Heterorthina praeculta Beds (H. praeculta Subzone)
Fi. Wattsella wattsi Beds (W. wattsi Zone)
F2 was stated to be a subzone of Fi, and, in another paper (19296, table opposite
p. 76), exactly the same subdivisions were again used. Bancroft claimed at first
that the fauna of the Marshbrookian was unknown elsewhere, but subsequently
(1933) he assigned strata in Westmorland, west Shropshire and North Wales to the
Stage and proposed a subdivision into the following three zones :
Onniella reuschi
Wattsella unguis
Wattsella wattsi
Not until 1945 did Bancroft define the type-section of the Marshbrookian as being
" through the highest beds of the Longville Flags as exhibited in the lane through
Marsh Wood, half a mile south of Marshbrook Station ".
Although Bancroft appeared to attach most importance to the brachiopods as
zone-fossils and used them in his first definition, in practice the fossil most charac-
teristic of the Marshbrookian as a whole is Broeggerolithus transiens (Bancroft).
This is not known from strata earlier than those lowest in the Marshbrookian,
and extends throughout the Stage in moderate numbers. Further, with reference
to the beds above the Marshbrookian, the absence of cryptolithids from the Actonian
in Shropshire is conspicuous and helps to fix the boundary between the two Stages.
The claim by Bancroft (1945 : 183) that Broeggerolithus disappears somewhat later
than the base of the Actonian has not been substantiated in Shropshire, and may
possibly refer to strata outside the county. Of Bancroft's zonal brachiopods,
Wattsella wattsi Bancroft and W. unguis (J. de C. Sowerby), both of which are now
referred to Dalmanella, appear to indicate successive horizons, but they have been
found to overlap slightly, at least in the Marshbrook District. Heterorthina praeculta
THE CARADOC SERIES OF SOUTH SHROPSHIRE 209
Bancroft is characteristic of the lower series of beds, but as it occurs in small numbers
throughout most of the strata containing D. wattsi its use as a subzonal fossil should
be discontinued. The third index-brachiopod, Onniella reuschi Bancroft, appears
to have some practical advantage, if not priority of publication, over Kjerulfina
polycyma Bancroft as an horizon indicator, and is retained for the topmost brachiopod
horizon of the Stage. Unless reasonably well-preserved, specimens of K. polycyma
may be difficult to distinguish from K. trigonalis Bancroft, a species which is found
only in the Dalmanella unguis Beds.
The type-section of the Marshbrookian, in Marsh Wood where the beds were
once quarried, is now almost overgrown and detailed collecting is difficult. The
topmost beds are not fully exposed, and probably the best continuous exposures
through the Stage are those in the Onny Valley, north-north-east of Cheney Long-
ville, where the lowest beds represent a transition from the underlying Upper Long-
villian, and comprise brownish-green flaggy siltstones with numerous lenticular
shelly bands. The latter are nearly always crowded with fossils, the most abundant
of which are Dalmanella wattsi and Tentaculites, with Broeggerolithus transiens,
Brongniartella bisulcata (Salter) and Chasmops. Individuals of the two last-named
often attain a large size.
Both lithologically and faunally the succeeding beds resemble those already
described, the only conspicuous difference being in the preponderance of Dalmanella
unguis which, although it overlaps slightly with D. wattsi, replaces that species as
the characteristic brachiopod. Rare and unusual trilobitic elements in the D. unguis
Beds of Marshbrook are Otarion and Encrinurus, the only horizon from which they
are known in the Caradoc of south Shropshire. The D. wattsi and D. unguis Beds
exhibit little change in lithology along the strike, but to the north-east of Soudley
the latter horizon includes some grey mudstones with abundant and conspicuous
ochreous-weathering fossils, such as D. unguis and B. transiens.
The upper beds of the Cheney Longville Flags consist typically of greenish-yellow
flaggy siltstones with lenticular fossil-bands. In the Onny. Valley they are followed
in the succession by beds of the Actonian Stage, at this, the type-locality, comprising
grey mudstones from which cryptolithids are absent, and in which small lamelli-
branchs, gastropods and cephalopods are dominant. The junction of the two Stages
is not usually exposed and there is some difficulty in mapping the precise line of
demarcation. In the Onny the highest visible Marshbrookian beds are flaggy
siltstones with Kjerulfina polycyma ; a gap in the exposures supervenes before
the Actonian mudstones are seen at Jack Slither. To the north, at Woolston,
what may be slightly higher beds with K. polycyma are exposed, and there is a
partial transition to a finer-grained mudstone lithology, some of the beds being
grey as well as the more usual yellowish-brown. Though the complete transition
is nowhere entirely exposed, the south-western portion of the track through Marsh
Wood, near Marshbrook, affords a good opportunity for further study, even though
partly overgrown. Flaggy siltstones, apparently the upper part of the D. unguis
Beds, are followed by a thin band of mudstone containing abundant worm burrows.
Next come mudstones resembling the Actonian beds of the Onny Valley, but contain-
ing a fauna which comprises both Marshbrookian and Actonian elements. It
2io THE CARADOC SERIES OF SOUTH SHROPSHIRE
includes small lamellibranchs, orthoceratids, Tropidodiscus acutus, Sinuites, Broeg-
gerolithus, Brongniartella bisulcata, Harknessella (s.l.), Hedstroemina fragilis,
Sowerbyella sericea and dalmanellids. Slightly higher in the succession flaggy
siltstones containing K. polycyma and Primaspis cf. caractaci are encountered.
The entry of new faunal elements seems to have been at least partly governed
by the lithology, and the line between the Marshbrookian and Actonian in south
Shropshire probably varied slightly from place to place, depending upon the condi-
tions at any one point. In this instance the transitional beds described are considered
to be best included in the Marshbrookian, primarily on account of their content
of cryptolithids, which are not found in the Actonian of the Onny Valley. This
description casts some doubt on the reliability of Kjemlfina polycyma as a zonal
index ; it does not usually occur in a mudstone environment, at least in Shropshire,
and appears to be restricted to the more arenaceous strata.
(f) Actonian Stage
The term Actonian was introduced first by Bancroft (19296, table opposite p. 76)
who, without giving definition or details, listed it in a stratigraphical table and
subdivided it into two zones, a lower one of Hedstroemina robusta, and an upper
one of Resserella (now Cryptothyris) paracyclica. The Stage was stated to be repre-
sented by part of the Acton Scott Beds of the Horderley District. Subsequently
Bancroft again listed the Stage in a stratigraphical table (1933), but on this occasion
used only the single brachiopod index of Onniella grandis, at that time an undescribed
species. Strata in south Shropshire, Westmorland and various parts of North
Wales were assigned to the Actonian.
It was not until twelve years later that Bancroft (1945 : 183) defined the base and
top of the Actonian, and designated the section exposed in the Onny Valley " east
of Burrells Coppice " as constituting the type-succession. His definition of the base
as being " marked by the disappearance of Kjerulfina and Tentaculites anglicus,
and the appearance of a large species of Onniella and Colpomya " is not found to be
sufficiently precise in practice, and does not take into account any variation of fauna
with environmental changes ; consequently, a detailed re-examination of the type-
succession is necessary.
The only place in Shropshire where a reasonably exposed succession through the
whole Actonian can be found is in the Onny Valley. Here it is possible, on the basis
of their faunas, to divide the beds roughly into three parts as follows, though some
difficulties arise when detailed correlation with other parts of the Caradoc Area is
attempted.
(i) The lowest Actonian comprises yellow and grey mudstones, with occasional
thin, nodular limestones, and shows a lithological transition from the topmost
beds of the Marshbrookian. Particularly noticeable is the sudden absence of crypto-
lithids ; instead, the mudstones of the Onny District were invaded by considerable
numbers of Platylichas laxatus (McCoy) and Chasmops, the latter usually considerably
larger than those from earlier strata. Kjerulfina has not yet been found in the
Actonian of the Onny, though Hedstroemina persists into the lowest beds, and the
THE CARADOC SERIES OF SOUTH SHROPSHIRE 2ii
most abundant brachiopod is Onniella depressa Bancroft. After a long absence
Heterorthis alternata (J. de C. Sowerby) returns to form a limestone band several
inches thick. The gastropods Holopea striatella (J. de C. Sowerby) and Sinuites
are often abundant, as are the corals Favosites and Coenites? , though there is no
suggestion of reef-conditions having existed.
(ii) The strata of the middle Actonian are marked by a decline in the number of
individuals of Platylichas and Chasmops, though these two genera are not altogether
uncommon. They are joined by another phacopid genus Calyptaulax, previously
unrecorded from Shropshire which, though not abundant, appears to be relatively
restricted in its vertical range. Primaspis [Acidaspis] caraclaci (Salter) is uncommon
in this mudstone environment. Of the brachiopods Onniella grandis is not very
abundant, but Cryptothyris paracyclica and Reuschella semiglobata Bancroft are
moderately common. All three species appear to be characteristic, but C. paracyclica
is here preferred to 0. grandis as index-brachiopod. Nicolella actoniae (J. de C.
Sowerby), abundant in some other districts of the Caradoc Area, is markedly
uncommon in the vicinity of the Onny.
(iii) The faunas of the upper Actonian in the Onny Valley appear to be less
prolific than those of the earlier beds, and diagnostic forms are less obvious, though
the strata are less well exposed. Platylichas and Chasmops are very much less
abundant, but they are accompanied by numerous Remopleurides sp. nov., a few
individuals of which first appeared late in the middle Actonian. Of the brachiopods
only Sowerbyella and Onniella are common, but Dolerorthis, Chonetoidea and Sampo
(recorded in Shropshire for the first time) occur. Onniella sinuata Bancroft is
known to be typical of an horizon fairly high in this upper third of the Actonian,
but its overall distribution is not yet fully known.
Northwards from the Onny Valley the beds are covered by Drift as far as the
neighbourhood of Acton Scott, where strata of Actonian age form the capping to
the high ground on which the village is situated. The detailed succession here is
much more difficult to follow, as well as being probably less complete, than in the
Onny Valley, and it has not been possible to examine fully the lowest beds and their
junction with the Marshbrookian. The valley running eastwards from Marshbrook
Village exposes strata which are probably rather higher in the Actonian succession.
They consist of yellowish-grey, sandy mudstones with a good fauna which includes
Platylichas, Primaspis caractaci, Remopleurides, Chonetoidea, Cryptothyris, Nicolella
actoniae and Onniella grandis, with common small gastropods and lamellibranchs.
A reasonable correlation is with the lower part of the middle Actonian of the Onny
Valley. The mudstones appear to pass upwards into more arenaceous beds, almost
quartzitic sandstones in part, which are more resistant to erosion than the underlying
mudstones, and which were once quarried extensively both at and near Acton Scott.
The most conspicuous elements of the fauna are Nicolella actoniae and Reuschella
bilobata (J. de C. Sowerby), associated with Cryptothyris paracyclica and Leptaena.
Trilobites also occur and include Platylichas, Chasmops, Primaspis caractaci and
Gravicalymene, with rare Illaenus and Atractopyge. Occasional bands of mudstone
yield locally abundant ostracods, such as Tetradella and Beyrichial '. The assemblage
suggests that the beds may be equated with part of the middle Actonian of the Onny
212 THE CARADOC SERIES OF SOUTH SHROPSHIRE
Valley. Higher horizons have not yet been confirmed, but this may be due to lack
of exposures. The assemblage of Platylichas, Illaenus, Atractopyge and Nicolella
is interesting in that it occurs at a much lower level in the Derf el Limestone (probably
basal Harnagian) of the Bala District (Whittington & Williams, 1955).
In the vicinity of Hatton, north-east of Acton Scott, Actonian beds are exposed
in the stream west of the village. The arenaceous strata of Acton Scott are not in
evidence, and the succession consists almost entirely of grey mudstones with bands
of impure limestone. The fauna is a prolific one, with Platylichas, Chasmops,
Reuschella bilobata and Onniella grandis in abundance, suggesting a lower to middle
Actonian age. The absence of higher beds is almost certainly due to the cover of
Drift or to the unconformable Upper Llandovery.
In the banks of Ticklerton Brook, between Ticklerton and Soudley, soft grey
mudstones occur below the basal Llandovery strata. Fossils are not abundant
but include Reuschella semiglobata, and the beds are probably middle Actonian
in age. The Marshbrookian/ Actonian junction is not exposed in this section.
It has been stated in recent years that strata belonging to Lap worth's Acton
Group do not occur to the north of the Cardington Hills (Pocock & Whitehead,
1948 : 51), but many years ago Salter & Aveline (1854 : 66) recorded them from the
then well-known fossiliferous locality of Gretton, near Cardington, cited again
by Cobbold (1900 : 55). Gretton Quarry, now unfortunately filled in, produced
an enormous fauna which clearly indicates a middle Actonian age. The large
number of forms found there include Platylichas, Chasmops, Illaenus, Gravicalymene,
Calyptaulax, Primaspis caractaci, Nicolella actoniae, Reuschella bilobata, Onniella
grandis, Sampo, Cryptothyris paracyclica, and abundant polyzoans and corals.
The beds are soft, yellow, flaggy sandstones with shelly lenses. The underlying
and overlying strata are insufficiently exposed for any detailed observations to be
made.
(g) Onnian Stage
This, the most restricted areally of all the Stages of the Caradoc Series in south
Shropshire, was introduced by Bancroft (19296, table opposite p. 76) ; it was divided
into three trilobite zones of Onnia cobboldi, 0. gracilis and 0. superba in ascending
order, and stated to occur only in the Onny Valley. Later it was again listed in a
tabular succession, but was correlated with strata in both south Shropshire and
Westmorland (Bancroft, 1933). In 1945 Bancroft named the Onny Valley as the
type-section and gave further details of the Stage (1945 : 183). The base of the
Onnian was described as being marked by the appearance of the cryptolithid genus
Onnia, and the summit by the disappearance of Onnia and the reappearance of
Tretaspis kjaeri, the latter being said to have a limited distribution in the Actonian.
Tretaspis is known from the Caradoc Series in Westmorland, but there is no evidence
that it has been found in south Shropshire.
Examination of the Onnian within the Caradoc Area is hampered by the uncon-
formable cover of Llandovery strata, as well as by extensive Drift deposits, and the
only accessible succession is in the Onny Valley south of Wistanstow. In practice
THE CARADOC SERIES OF SOUTH SHROPSHIRE 213
Bancroft's definition of the base has proved convenient ; the preceding Actonian
contains no cryptolithids, and their sudden reappearance can be easily followed in
the field. The three successive cryptolithid zones erected by Bancroft appear to be
well established in the Onny region, but unfortunately their lateral development
cannot be traced owing to paucity of exposures.
Lithologically the Onnia cobboldi Zone shows no obvious change from the Actonian,
but the trilobite fauna differs markedly, although occasional Actonian elements
remain. Plalylichas and Chasmops, genera so abundant in parts of the Actonian,
occur merely as occasional isolated individuals, and only the former genus has been
recorded from the succeeding Onnia gracilis Zone. Similarly, illaenids like those of
the Actonian are sometimes found. The occurrence of two specimens of Gravi-
calymene in the 0. gracilis Zone is interesting because it is apparently the same form
as is so abundant in some of the Actonian sandstones. The trilobite fauna of the
topmost zone of Onnia superba is particularly rich in individuals of the zonal crypto-
lithid, but other forms include Lonchodomas pennatus (La Touche), Raphiophorus
edgelli (Reed), Remopleurides burmeisteri Bancroft, Triarthrus, Pseudosphaerexochus
and Eobronteus? , the last two being extremely rare. The brachiopod fauna of the
two lower zones of the Onnian comprises mainly the genus Onniella. 0. inconstans
Bancroft in the Onnia cobboldi Zone is followed in the Onnia gracilis Zone by the
related Onniella broeggeri Bancroft, accompanied by abundant small Chonetoidea
and ostracods.
The strata of the Onnia superba Zone would appear to indicate quieter conditions
of deposition, and whole trilobites are of frequent occurrence. Coincident with
these conditions Onniella almost disappears, and the brachiopods usually found
are Chonetoidea (like that in the Actonian), and the very small " Rafinesquina "
holli (Davidson) which may be locally abundant.
The term Acton Scott Beds was used by Bancroft (19296, table opposite p. 76 ;
1933) to include the Actonian and the two lower trilobite zones of the Onnian,
leaving the Onnia superba Zone equivalent to the Onny Shales, which is apparently
the same usage as that of La Touche (1884). As stated earlier, however, the Acton
Scott Beds (s. s.) of the Acton Scott District include only part of the Actonian
Stage. Furthermore, the sharp colour change between the yellow-weathering
mudstones of the Onnia superba Zone at the well-known " Cliff Section ", with their
limonitic fossils, and the earlier Onnian strata is apparent rather than real, and all
the rocks appear as grey mudstones when seen in fresh section. Accordingly,
the strata belonging to the Onnian Stage in south Shropshire are here named Onnia
Beds nom. nov., with the Onny Valley between Cheney Longville and Wistanstow
as type-locality.
III. LAPWORTH'S SUBDIVISIONS OF THE CARADOC SERIES
Lapworth (1916) published a series of vertical sections covering the Lower
Palaeozoic rocks of Shropshire. He applied the name " Caradoc Series " to the
Ordovician rocks forming the elongated outcrop in the Caer Caradoc- Wrekin District
and subdivided them to give the following succession :
214
THE CARADOC SERIES OF SOUTH SHROPSHIRE
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THE CARADOC SERIES OF SOUTH SHROPSHIRE 215
~ f Upper Trinucleus Beds
Acton Group < A ~ , ,-, ,
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Longville Group . ^ Birrells Wood Flags
Chelmick Flags and Shales
Alternata Limestone
Upper Chatwall Sandstone
Chatwall Group . s T /-u n c j *.
I Lower Chatwall Sandstone
(Glyptocrinus Flags)
Harnage Group . Harnage Shales
Transition Bed
Upper Hoar Edge Sandstone
Hoar Edge Group . < TT -c-j T •
I Hoar Edge Limestone
Lower Sandstone and Conglomerate
Lapworth supervised extensive collecting by the Geological Survey from horizons
throughout the Series, but the results remained unpublished and the subdivisions
listed above were never defined or described. Some use has been made of the terms
(Watts, 1925 : 340 ; Pocock et al., 1938 : 81-90) and it is necessary to re-assess them
and ascertain their usefulness in the light of present-day knowledge.
Hoar Edge Group
No details having been given in addition to the table, it is difficult to fit the suc-
cession into the zonal sequence, but it has been pointed out (Pocock et al., 1938 : 86)
that the specimens of Nemagraptus gracilis identified from the Evenwood District
came from beds containing Harknessella subplicata near the top of the Hoar Edge
Group, which at that point is only about 100 ft. thick. These beds would, then,
coincide at least approximately with Lapworth's Lower Sandstone and Conglomerate,
the so-called Harknessella subquadrata Limestone above being equivalent to his
Hoar Edge Limestone. It is possible, though not certain, that the Upper Hoar
Edge Sandstone refers to the Rhynchonellid Grits, the latter, together with the
" Transition Bed ", being considered Harnagian in age. It is doubtful whether
Lapworth's detailed succession can be applied successfully to the southern part of
the Caradoc Area.
Harnage Group
As stated earlier, the strata referred to his " Transition Bed " by Lapworth
are almost certainly those topmost portions of the Hoar Edge Grits in the Evenwood
District which contain S alter olithus and are thus of Harnagian age.
The Harnage Shales of the type-area of Coundmoor Brook represent only the
Reuscholithus reuschi Zone of the Harnagian, the higher zones being cut out by
faulting. The Harnage Group, strictly interpreted, cannot therefore include more
than the lowest trilobite Zone of the Harnagian. It does not constitute a good
stratigraphical subdivision on faunal grounds and is probably best allowed to lapse.
GEOL. 3, 6. 19
216 THE CARADOC SERIES OF SOUTH SHROPSHIRE
Chatwall Group
The term " Glyptocrinus Flags " has already been referred to in the discussion
of the Soudleyan Stage, and is considered to be almost exactly equivalent to the
Broeggerolithus soudleyensis Zone in the Onny Valley, but farther north may include
beds of a slightly earlier date. The term appears to be in more general use than that
of " Lower Chatwall Sandstone ".
In order to summarize the stratigraphical position of those strata known as the
Upper Chatwall Sandstone, it is necessary to examine the succession in the type-
area around Chatwall Hall, one-and-a-half miles north-north-east of Cardington.
To the north-west of Chatwall Hall the Glyptocrinus Flags crop out and have been
quarried, for example, 70 yards north-north-west of the Hall, where they consist
of dark green flaggy sandstones with deeply-weathered shelly lenses containing
abundant Broeggerolithus cf. broeggeri. These beds appear to pass upwards into
more massive maroon and grey-green sandstones, such as those quarried just
south-east of Chatwall Farm. From this quarry a narrow cutting has been made,
leading south-eastwards to the adjacent crossroads, and at the eastern end of the
cutting slightly higher beds have been extensively quarried. These two series of
strata have generally been grouped together as the Chatwall Sandstone, and the
intermediate beds exposed in the cutting have been described by Robertson (in
Pocock et al., 1938 : 88). The higher sandstone beds, or Upper Chatwall Sandstone,
are followed in the succession by lenticular limestones of the Alternata Limestone
series (Upper Longvillian) , but there are certain features which make it difficult to
accept them without question as being of pre-Upper Longvillian age. For example,
the fauna includes Heterorthis alternata (J. de C. Sowerby) with abundant large
Sowerbyella sericea and Brongniartella bisulcata, all of which are particularly abundant
and characteristic in the lower part of the Upper Longvillian elsewhere in the Caradoc
Area. Furthermore, the lithology of the Upper Chatwall Sandstone, comprising
soft, yellow-weathering flaggy sandstones with shelly lenses, is quite different
from that of the Lower Chatwall Sandstone, and is separated from it by thick
bands of conglomerate which often contain abundant gastropods. The problem
is not yet satisfactorily resolved, but it is possible that the Upper Chatwall Sandstone
may eventually prove to be at least partly Upper Longvillian in age, forming what
is virtually an arenaceous development of the basal beds of the Alternata Limestone.
The exact horizon of the sandstones below the conglomerates is not yet known,
but the possibility of the further extension of the stratigraphical break below the
Upper Longvillian which is so apparent at Soudley cannot be excluded.
The Alternata Limestone was placed by Lap worth in his Chatwall Group, but
on faunal grounds there can be no doubt that its place is with the Lower Cheney
Longville Flags in the Upper Longvillian.
Longville Group
Lapworth's term " Chelmick Flags and Shales " presumably refers to the area
of Chelmick, about one mile south-west of Hope Bowdler. The derivation of
" Birrells Wood Flags " is less clear,- as no such place exists on the present-day
THE CARADOC SERIES OF SOUTH SHROPSHIRE 217
Ordnance maps, but it is here suggested that the name may refer to what is now
known as Burrells Coppice, by the south bank of the River Onny, rather less than
half-a-mile north of Cheney Longville. It is probably true that the beds near
Chelmick are fairly low in what generally are called the Lower Cheney Longville
Flags, and those exposed at the eastern end of Burrells Coppice are certainly high
in the same group. Owing to lack of good continuous exposures at both places it is
difficult to correlate the two subdivisions exactly, and the most practical solution
is to allow the two names to lapse, while retaining the term Lower Cheney Longville
Flags. The latter has the great merit of possessing a definite type-locality, in the
lane leading westwards from Cheney Longville, and the almost continuous section
available there does not suggest that further lithological subdivision would be of
any great value, though there are certain variations in the faunas.
The " Lower Trinudeus Shales " must be taken to represent the whole of what
is now regarded as the Marshbrookian Stage, but for a number of reasons the name
is highly unsuitable. The Marshbrookian, or Upper Cheney Longville Flags, consists
almost entirely of massive and flaggy siltstones and mudstones. Cryptolithid
trilobites occur throughout, but certainly not in numbers sufficient to give their
name to the beds. Furthermore, no type-locality was ever cited, and the name is
best rejected.
Acton Group
Though not defined as such, it is certain that Acton Scott must have been the
intended type-locality of the " Acton Calcareous Beds ", which are thus at least
approximately equivalent to the Acton Scott Beds (s. s.) discussed on p. 211. No
type-locality was chosen for the Upper Trinudeus Beds, though one can claim that,
by implication, they are equivalent to at least part of the Onnian Stage of the
Onny Valley.
If the Acton Calcareous Beds and the Upper Trinudeus Beds are, in fact, equivalent
to the Acton Scott Beds (s. s.} and part of the Onnian respectively, then, if they
are used in their restricted sense, they cannot be exactly equivalent to the Actonian
and Onnian Stages, as it has already been shown that the Acton Scott Beds (s. s.)
probably represent only a part of the Actonian Stage.
IV. FAUNAL LISTS
Each of the species in the following lists is accompanied by a number, or numbers,
followed by a letter, or letters. The former refers to the horizon as indicated in
Text-fig. 3 ; the latter refers to the district, or districts, within the Caradoc Area,
as listed below.
A.— Coston. F.— Little Stretton.
B.— Onny Valley. G.— Soudley-Ticklerton.
C. — Brokenstones, near Horderley. H. — The Cwms, near Church Stretton.
D. — Marshbrook. K. — Evenwood — Harnage Grange.
E. — Acton Scott.
218
THE CARADOC SERIES OF SOUTH SHROPSHIRE
Costonian Stage
Favosites fibrilla Smith .
Polyzoans (various)
Cliftonia cf . andersoni Reed
Dalmanella sp. (small)
Dinorthis flabellulum (J. de C. Sowerby)
D. aff. flabellulum (J. de C. Sowerby)
Dinorthis sp. A
Dinorthis sp. B
Dolerorthis sp.
Glyptorthis ? sp.
Harknessella jonesi Bancroft
H. subplicata Bancroft .
H. subquadrata Bancroft
H. vespertilio (J. de C. Sowerby)
Harknessella sp.
Heterorthis patera (Davidson)
Horderleyella plicata Bancroft .
Leptaena sp. .
Lingula cf . ovata McCoy
Orbiculoidea sp.
Rafinesquina cf . complanai
R. aff. expansa (J. de C. Sowerby)
Rafinesquina sp.
Rafinesquina sp. (? nov.)
Salopia salteri (Davidson)
Siphonotreta cf . micula McCoy
Smeathenella strophomenoides Bancroft
Sowerby ella sp.
Cyclonema cf. crebristria (McCoy)
Leseurilla balclatchiensis LongstafE
Liospira aequalis (Salter)
Rhaphistomina ? sp.
Hyolithes sp.
Ambonychia sp.
Ctenodonta cf. varicosa (Salter)
" Orthoceras " sp. indet. .
Asaphid indet.
Brongniartella aff. bisulcata (Salter)
Costonia ultima (Bancroft)
Costonia sp. nov. .
Eohomalonotus sp. .
Flexicalymene cf. acantha Bancroft
Flexicalymene sp. .
Metopolichas ? aff. verrucosa (Eichwald)
Reacalymene pusulosa Shirley
Primitia simplex (Jones)
P. strangulata (Salter)
Tetradella scripta Harper
2
A, C, K
2
A, B, C, H, K
I,
2 ; A, C, K
2
A, C
Sowerby) i
A
werby) ... 2
C, K
X
A
. .... 2
C
2
A, C
. . .2
A
. .... I
A
. . . I
K
2
K
y) . . i
A
2
C
I
A : 2 ; C
2
A, C
2
A, C
2
A
2
A
. de C. Sowerby) . . 2
A, K
rby) .... I
A
2
A, C
2
K
2
C
2
K
Bancroft ... 2
A,C
i,
2 ; A, C, K
:>y) . . i,
2 ; K
jtaff .... 2
K
2
K
I
A
. .... 2
A, K
2
K
I,
2 ; K
. .... 2
K
I
A
alter) .... 2
K
.2
A,C
2
H. K
. . . . .2.
K
croft . . . . 2
A, C
i
A: 2 ; K
Eichwald) ... 2
C, K
. . . . . i,
2; K
. . . .2
A, K
. . . .2
K
2
A, C, K ?
THE CARADOC SERIES OF SOUTH SHROPSHIRE
219
Harnagian Stage
Cystid plates ......
Crinoid ossicles ......
Lepidocoleus suecicus Moberg ....
Plumulites sp.
Turrilepas ? sp.
Polyzoans (various) .....
Chonetoidea ? sp. .
Cliftonia cf. andersoni Reed ....
Dalmanella sp. ......
Dinorthis sp. ......
Dolerorthis sp. ......
Harknessella cf. vespertilio (J. de C. Sowerby) .
Heterorthis aff. patera (Davidson)
Horderleyella sp.
Lingula sp. .......
Orbiculoidea sp. ......
Orthorhynchula sp. .....
Paterula cf . albida Reed .....
Rostricellula aff. triangularis Williams
Salopia salteri (Davidson) ....
Siphonotreta cf . scotica Davidson
Smeathenella harnagensis Bancroft .
S. cf. harnagensis Bancroft ....
Sowerby ella aff. sericea (J. de C. Sowerby)
Vellamo sp. .......
Carinopsis cf . gracilis «(Reed) ....
Cyrtolites sp. ......
Phragmolites sp. ......
Raphistoma sp. ......
Conularia (s.l.) sp. .....
Hyolithes sp. .......
Ctenodonta aff. coarctata (Phillips)
C. cf. varicosa Salter .....
Modiolopsis ? aff. postlineatus McCoy
" Orthoceras " sp. ......
Basilicus marstoni (Salter) ....
Brongniartella sp. .
Decoroproetus fearnsidesi (Bancroft)
D. cf . fearnsidesi (Bancroft) ....
Diacalymene ? praecox Bancroft
Flexicalymene acantha Bancroft
Flexicalymene sp. .
Nieszkowskia stubblefieldi Bancroft .
Parabasilicus powisi ? (Murchison) .
Reufcholithus reuschi Bancroft
Salterolithus harnagensis Bancroft
S. aff. harnagensis Bancroft ....
S. smeathenensis Bancroft ....
S. caractaci (Murchison) .....
Primitia nana Jones & Holl ....
P. simplex (Jones) .....
Tetradella scripta Harper ....
Ulrichia bicornis (Jones) ....
3 ; B, G, K
3, 4 ; B : 3 ; K
3 ; B, G, K
B, K
B
B, C, F, G, K
B
B, C
B
B, C, F, G
B, G
B, G
B, C
B
B, K
B
B, G
K
B
B, C, F, G
B
B, C, K ?
B
B, C, F, G, K
G
K
B, K?
B
K
B, K
B, K
B, K
B, G, K : 4 ; B
K
K
B
B, K
B, K
F
B, K
B, K
B
B
B
B, G, K
K
G,H, K
B
B, G ?
B, C, G, K
B, G ?, K
3 ; B, C, F, G, K : 4 ; B, G ?
3 ; B, C, G, K
220 THE CARADOC SERIES OF SOUTH SHROPSHIRE
Soudleyan Stage
Favosites fibrilla Smith . . . . . . 6 ; H : 7 ; B, G
Crinoid ossicles 6 ; H : 7 ; B, G, H
Rhaphanocrinus basalis (McCoy) . . . . 6 ; H : 7 ; B, G. H
Cliftonia cf . andersoni Reed . . . . . 6 ; B : 7 ; B, G, H
Craniops sp. . . . . . . . 6 ; B : 7 ; B, H
Dinorthis aff. flabellulum (J. de C. Sowerby) . . 7 ; B, G
Heterorthis retrorsistria (McCoy) . . . . 6 ; B : 7 ; B, G, H
Horderleyella corrugata Bancroft . ' . . . 7 ; B
Leptaena sp. . . . . . . . 6 ; B
Orbiculoidea sp. (small) . . . . . 6 ; B
Rafinesquina expansa (J. de C. Sowerby) . . . 7 ; B, G ?, H
Rafinesquina sp. . . . . . . 7 ; B
Reuschella horderleyensis Bancroft . . . . 7 ; B, G
Soudleyella avelinei (Bancroft) . . . . 6 ; B
Sowerbyella sp. . . . . . . . 6 ; B, H : 7 ; B, H
Carinopsis cf. gracilis (Reed) . . . . . 7 ; H
Cyclonema crebristria (McCoy) . . . . 7 ; B, H
Cyrtolites nodosus (Salter) . . . . . 7 ; B
Sinuites bilobatus (J. de C. Sowerby) . . . 7 ; B, H
S. soudleyensis Reed . . . . . . 7 ; H
Hyolithes sp. 7 ; H, K
Ctenodonta sp. . . . . . . . 6 ; B : 7 ; H, K
Orthonota sp. . . . . .. . 6;B:7;H
" Orthoceras " sp. . . . . . . . 6 ; B
Broeggerolithus broeggeri (Bancroft) . . . . 6 ; B, H ?
B. constrictus Bancroft . . . . . . 6 ; B
B. soudleyensis (Bancroft) . . . . . 7 ; B, G, H
Broeggerolithus sp. . . . . . . . 6 ; B
Brongniartella sp. . . . . . . . 6 ; B, H
Calymenid indet. . . . . . . . 6 ; B
Decoroproetus ? sp. . . . . . . 6 ; B, H : 7 ; B, H
Parabasilicus powisi (Murchison) . . . . 6 ; B : 7 ; B
Reacalymene cf . pusulosa Shirley . . . . 7 ; K
Salterolithus sp. . . . . . . 6 ; B
Primitia sp. . . . . . . . 6 ; B
Tetradella scripta Harper . . . . . 6 ; B
Climacograptus antiquus Hall var. . . . . 6 ; B
Orthograptus cf . apiculatus Elles & Wood . . » 7 ; B
Longvillian Stage
(i) Lower Longvillian Substage
Favosites fibrilla Smith . . . . . . 8 a, b, c ; B
Polyzoans (various) . . . . . . 8 a, b, c ; B
Bancroftina typa (Whittington) . . . . 8c;B
Cliftonia cf . spiriferoides (McCoy) . . . . 8 b ; B
Dalmanella horderleyensis (Whittington) . . . 8 a ; B
D. indica Whittington . . . . .. 8b;B
D. lepta (Bancroft) . . . ., ' . . 8 b ; B
Dinorthis sp. . . . . . . . 8 b ; B
Dolerorthis sp. . . . . . . 8 b ; B
Kjaerina hedstroemi Bancroft . . . . . 8 b ; B
THE CARADOC SERIES OF SOUTH SHROPSHIRE
221
K. horderleyensis Bancroft
K. intermedia Bancroft .
K. jonesi Bancroft
K. cf . jonesi Bancroft
cf. K. richteri Bancroft .
Leptaena sp. .
Lingula sp. .
Orbiculoidea sp.
Rafinesquina sp. .
Resserella canalis (J. de C. Sowerby)
Sowerbella soudleyensis Jones .
Clathrospira ? sp. .
Cyrtolites nodosus (Salter)
cf. Liospira aequalis (Salter)
Lophospira cf. gyrogonia (McCoy)
Murchisonia ? sp. .
Sinuites anceps Reed
S. bilobatus (J. de C. Sowerby)
5. soudleyensis Reed
Ambonychia ? sp. .
Ctenodonta sp.
Broeggerolithus globiceps (Bancroft) .
Broeggerolithus sp.
Brongniartella bisulcata (Salter) var.
Eohomalonotus sp. (? nov.)
Phacopidina apiculata (Salter)
P. aff. harnagensis Bancroft
Parabasilicus powisi ? (Murchison) .
Reacalymene sp. nov.
8a
8a
8 a
8 c
8 c;
B
B
B
B
B
B
B
8 a, b ; B
8 b, c ; B
8c; B
8c; B
8 a, b, c ;
8 a, b, c ;
8 a; B
8 a, b ; B
8a; B
8 a; B
8 a, b ; B
8 a; B
8 a; B
8 a, b ; B
8 a ; B
8 a; B
8 a, b, c ; B
8b; B
8 a, b ; B
8b; B
8 a, b, c ; B, D
8 a; B
8 a; B
8 a, b ; B, D
(ii) Upper Longvillian Substage
Favosites fibrilla Smith .
Streptelasma ? sp. indet. .
Lepidocoleus suecicus Moberg .
Bancroftina robusta (Bancroft)
B. aff. typa (Whittington)
Dolerorthis duftonensis (Reed) .
Harknessella (s.l.) sp.
Heterorthis alternata (J. de C. Sowerby)
Kjaerina bipartita (Salter)
K. geniculata Bancroft .
K. typa Bancroft ....
Kjaerina sp. .....
Lingula sp. .
Marionites typa (Bancroft)
Nicolella ? sp.
Orbiculoidea sp. ....
Philhedra drummuckensis Reed
Platystrophia sp. ....
Plectorthis ? cf. plicatella Reed
Rafinesquina sp. .
Schizocrania sp. ....
Sowerbyella sericea (J. de C. Sowerby)
9 a, b ; B, G
9b; B
9 b ; B, G
9 a; B
9b; B
9 a, b ; B, G
B
B. D, G
B, G
B
B
B
9b;
ga;
9 a ;
9b;
9b;
9b;
9 a, b ; B
9 a ; B, G
9b; B
9a
ga
9b
9b
9b
9 a
G
G
B
B
B
G
9 a, b ; B, D, G
THE CARADOC SERIES OF SOUTH SHROPSHIRE
Strophomena grandis (J. de C. Sowerby) . . . 9 a, b ; B, G
Trematis punctata (J. de C. Sowerby) . . . 9 a ; B, G
Cyrtolites nodosus (Salter) . . . . 9 a ; G
Lophospira cf . gyrogonia (McCoy) . . . . 9 a ; G
Sinuites soudleyensis Reed . . . . . 9 a ; G
Tentaculites cf. scalaris Schlotheim . . . . 9 a, b ; B, D, G
Ambonychia orbicularis (J. de C. Sowerby) . . 9 b ; B
Ctenodonta aff. varicosa Salter . . . . . 9 a ; G
Pterinea sp. . . . . . . . . 9 a ; G
Asaphid indet. . . . . . . 9 a, b ; B
Broeggerolithus longiceps (Bancroft) . . . . 9 a, b ; B, C, G
B. aff. soudleyensis (Bancroft) . . . 9 a ; G
Brongniartella bisulcata (Salter) . . . . 9 a, b ; B, D, G
Chasmops sp. . . . . . . . 9 b ; B
Flexicalymene aff. caractaci (Salter) . . . . 9 b ; B
Phacopidina apiculata (Salter) . . . . 9 a, b ; B, G
P. cf . apiculata (Salter) . . . . . . 9 a ; G
Primitia sp. . . . . . . . . g a.; G
Tetradella cf . scripta Harper . . . . . 9 a ; G
Marshbrookian Stage
Favosites fibrilla Smith . . . . . 10 a, b, c ; B, D
Polyzoans (various) . . . . . 10 a, b, c ; B, D
Craniops sp. . . . . . . . . 10 c ; B
Dalmanella multiplicata (Bancroft) . . . 10 a ; B, D
D. unguis (J. de C. Sowerby) . . . . . 10 b ; B, D
D. wattsi (Bancroft) . . . . . 10 a ; B, D
Dolerorthis sp. . . . . . . 10 b ; D
Hedstroemina fragilis Bancroft . . . . 10 c ; B, D
H. parva Bancroft . . . . . 10 b ; B
Heterorthina praeculta Bancroft . . . . 10 a, b ? ; B, D
Heterorthis alternata (J. de C. Sowerby) . . 10 c ; B
Kjaerina sp. . . . . . . . 10 c ; B
Kjerulfina polycyma Bancroft . . . . 10 c ; B, D
K. trigonalis Bancroft . . . . . 10 b ; B, D
K. cf. trigonalis Bancroft . . . . 10 a ; D
Lingula sp. . . . . . . . 10 a ; D
Lingula sp. (large) . . . . . . 10 a ; D
Nicolella cf. actoniae (J. de C. Sowerby) . . . 10 a, b, c ; D
Onniella reuschi Bancroft . . . . . 10 c ; B, D
Plectorthis ? virgata Reed . . . . . 10 a, b ; B
Reuschella aff. bilobata (J. de C. Sowerby) . . . 10 c ; B, D
Schizocrania crassa (Salter non Hall) . . . 10 a, b ; D
Strophomena grandis (J. de C. Sowerby) . . . 10 a, b, c ; B, D
Cyrtolites nodosus (Salter) . . . . . 10 b ; D
Sinuites sp. . . . . . . . . 10 c ; D
Tropidodiscus acutus (J. de C. Sowerby) . . 10 c ; D
Conularia (s.l.) sp. . . . . . . 10 b ; D
Hyolithes sp. . . . . . . . 10 b ; D
Tentaculites cf. scalaris Schlotheim . . . . 10 a, b, c ; B, D, G
Ambonychia obliqua (J. de C. Sowerby) . . . 10 b, c ; D
A. orbicularis (J. de C. Sowerby) . . . . 10 a, b, c ; D
Ctenodonta sp. . . . , . . 10 c ; D
THE CARADOC SERIES OF SOUTH SHROPSHIRE
223
Broeggerolithus transiens (Bancroft)
B. cf. transiens (Bancroft)
Brongniartella bisculata (Salter)
Chasmops sp. ...
Encrinurus sp.
Flexicalymene caractaci (Salter)
F. trigonoceps Bancroft .
Flexicalymene sp. .
Lichas (s.l.) sp. indet.
Otarion sp. ....
Phacopidina apiculata (Salter)
Primaspis caractaci (Salter)
Proetus (s.l.) sp. .
Tetradella cf . scripta Harper .
10 a, b, c ; B, D, G
10 c ; B, D
10 a, b, c ; B, D
10 a, b, c ; B, D
10 b; D
10 b, c ; B, D
10 a, b, c ; B, D
10 a, b ; D
10 b; D
10 b ; D, G
10 a, b ? ; B, D
10 c ; D, G ?
10 b; G
10 b, c ; D, G
Actonian Stage
Coenites ? sp. ......
Favosites fibrilla Smith .....
Lepidocoleus suecicus Moberg ....
Polyzoans (various) .....
Chonetoidea sp. ......
Cryptothyris paracyclica (Bancroft) .
Dolerorthis sp.
Heterorthis alternata (J. de C. Sowerby)
Kjaerina sp. . . . *
Leptaena sp. .......
Lingula cf. attenuata (J. de C. Sowerby) .
Lingula sp. .......
Nicolella actoniae (J. de C. Sowerby)
Onniella aspasia Bancroft ....
O. depress a Bancroft .....
O. grandis Bancroft .....
O. sinuata Bancroft .....
Orbiculoidea cf . perrugata (McCoy) .
Platystrophia sp. .
Rafinesquina sp. ......
Reuschella bilobata (J. de C. Sowerby)
R. semiglobata Bancroft .....
Sampo sp. .......
Sowerby ella aff. sericea (J. de C. Sowerby)
Strophomena grandis (J. de C. Sowerby)
Tr emails punctata (J. de C. Sowerby)
Triplesia sp. .......
Archinacella cf. oblongata (Portlock)
cf . Clathrospira trochiformis (Portlock)
Cyrtolites nodosus (Salter) ....
Ecculiomphalus sp. .....
Holopea striatella (J. de C. Sowerby)
Sinuites bilobatus (J. de C. Sowerby)
S. pseudocompressus Reed ....
Tropidodiscus acutus (J. de Sowerby)
Metaconularia cf . sowerbyi (de Verneuil) .
M. vesicularis (Slater) .....
GEOL. 3, 6.
II
B,
E,
J
II
B,
E,
J
II
B
II
B,
E,
G. J
II
B,
E,
J
II
B,
E,
J
II
B
II
B
II
B,
J
II
B,
E,
J
II
B
II
B
II
B,
E,
J
II
B,
E
II
B
II
B,
G,
J
II
B
II
B,
J
II
J
II
B,
E
II
E,
J
II
B,
E
?, G
II
B,
J
II
B,
E,
J
II
E
II
B
II
J
II
B
II
B
II
B
II
B
II
B,
J
II
B
II
B
II
B
II
J
II
B,
E,
J
20
224
THE CARADOC SERIES OF SOUTH SHROPSHIRE
Hyolithes sp.
" Orthoceras " sp. .
Ctenodonta varicosa Salter . .
Modiolopsis cf. modiolaris (Conrad) .
Orthonota cf. subcylindrica (McCoy) .
Orthonota sp. (large) . . .
cf. " Pectunculus " ambiguus Portlock
Pterinea sp. . . . . .*
Calyptaulax sp. . . . . .
Chasmops sp. . .
Flexicalymene cf . caractaci (Salter) .
F. laticeps Bancroft .
F. salteri Bancroft .
Gravicalymene sp. .
Illaenus sp. .
Lonchodomas pennatus (La Touche) .
Platylichas laxatus (McCoy) . . .
Primaspis caractaci (Salter) . . .
Raphiophorus edgelli (Reed)
Remopleurides sp. nov. . . . .
Beyrichia ? sp.
Primitia sp. ......
Tetradella cf . scripta Harper .
Diplograptus sp. .
Orthograptus of truncatus Lapworth group
ii ;
B
ii
B, G
ii
E, G
ii
B, E
ii
B
ii
E
ii
B
ii
B,E
ii
B, J
ii
B, E, J
ii
E
ii
B, E
ii
B?, E
ii
E, J
ii
B, E, J
ii
B, G
ii
B, E, G, J
ii
B, E, J
ii
B
ii
B, E
ii
E
ii
B, E
ii
E
ii
B
ii
B
Onnian Stage
Chonetoidea sp. ......
Onniella broeggeri Bancroft .
O. inconstans Bancroft .
Onniella sp. .......
Orbiculoidea cf . perrugata (McCoy) .
" Rafinesquina " holli (Davidson) . . ,
Sowerbyella sp.
Raphistomina ? sp.
Sinuites pseudocompressus Reed
Sinuites sp. .
Tropidodiscus acutus (J. de C. Sower by) .
Metaconularia cf. vesicularis (Slater)
Ceratotheca ? cf . subuncta Reed
Colpomya sp. .....
Ctenodonta sp. .....
Orthodesma sp. .
cf. " Pullastra " speciosa McCoy
Orthonota cf. subcylindrica (McCoy) .
" Cyrtoceras " sp. .
" Orthoceras " sp. . - .
Eobronteus ? sp. . »
Flexicalymene onniensis Shirley
F. aff. onniensis Shirley ....
Gravicalymene sp. .
Illaenus sp. . . .
Lonchodomas pennatus (La Touche) .
12, 13, 14; B
13: B
12; B
14 ; B
14 ; B
14; B
12 ; B
14; B
12, 13 ; B
14 ; B
12, 13 ; B
14 ; B
13: B
12; B
13; B
13: B
13; B
13; B
14 ; B
14 ; B
14 ; B
13, 14 ; B
12; B
13; B
12, 13, 14; B
12, 13, 14 ; B
THE CARADOC SERIES OF SOUTH SHROPSHIRE 225
Onnia cobboldi (Bancroft) . . . . . 12 ; B
O. gracilis (Bancroft) . . . . . . 13 ; B
O. superba (Bancroft) . . . . . . 14 ; B
Platylichas laxatus (McCoy) . . . . . 12 ; B
Pseudosphaerexochus sp. indet. . . . . 14 ; B
Raphiophorus edgelli (Reed) . . . . . 12, 13, 14 ; B
Remopleundes burmeisteri Bancroft . . . . 13, 14 ; B
Triarthrus sp. . . . . . . . 14 ; B
Climacograptus sp. . . . . . . 13 ; B
Orthograptus cf. apiculatus Elles & Wood . . 14 ; B
V. THE CARADOC/LLANDOVERY JUNCTION
The Caradoc and Llandovery strata of south Shropshire are separated by a
profound stratigraphical break, the importance of which was not, at first, fully
realized. Murchison claimed that the succession in the Onny Valley continued
unbroken from the " Caradoc Sandstone " to the Wenlock Shales, and it was left
to Salter & Aveline (1854 : 7°) to demonstrate the angular break below the Purple
Shales (called by them, Pentamems Beds) at the now classic " Cliff Section " in the
north bank of the River Onny, one mile south-west of Wistanstow ; this has been
described in detail by Whittard (1927 : 749). At different points along the outcrop
Llandovery beds rest on different horizons within the Caradoc Series, and in the
present account it is convenient to list these from the north to the south of the Caradoc
Area.
The succession employed by the Geological Survey for the Llandovery strata
(Robertson in Pocock et al., 1938 : 106) is as follows :
3. Hughley Shales
2. Pentamerus Beds
i. Kenley Grit
Beds i and 2 are only local names, and it may be an advantage in the present
discussion to follow Whittard (1927 : 738) in adopting stratigraphical terms which are
of more general application, such as :
c. Purple Shales
b. Pentamerus Beds
a. Arenaceous Beds
The most northerly point of the Caradoc /Llandovery junction is to be found
to the south-east of Harnage Grange, about seven-and-a-half miles south-east of
Shrewsbury. Here the Arenaceous Beds rest on Costonian strata, successively
higher horizons being transgressed as Church Preen is approached, to the south-
west of Kenley, where the Upper Longvillian is overlain. Near Plaish, and at Gretton
near Cardington, it is unlikely that the Caradoc outcrop includes any horizon higher
than the Actonian Stage.
The continuity of the Caradoc outcrop is broken by the Uriconian of Cardington
Hill, to the south-south-east of which the Pentamerus Beds rest on strata low in the
Caradoc sequence, though possibly no lower than Harnagian. The actual contact
is obscured by Drift. Farther south-westwards, about two-fifths of a mile east-
north-east of Hollies Farm, the highest Caradoc strata are flaggy Marshbrookian
226 THE CARADOC SERIES OF SOUTH SHROPSHIRE
siltstones with abundant Dalmanella unguis and Broeggerolithus transiens. In
Ticklerton Brook, and probably also at Hatton, one mile farther south-west, Penta-
merus Beds overlie mudstones of Actonian age.
As stated earlier, the highest Caradoc beds seen in the Acton Scott District are the
Acton Scott Beds (s. s.), probably equivalent to the middle part of the Actonian.
There is, however, some distance between their outcrop and the conjectured base of
the Pentamerus Beds ; the possibility that a small outcrop of Onnian exists
immediately south-east of Acton Scott cannot be ignored, and some support is
derived from Salter & Aveline's record of " Trinucleus Shales " (? Onnian) at Henley,
one mile to the south-south-west. These are not now exposed, and the solid geology
here is much obscured by extensive Drift deposits.
Tracing the lower limit of the Llandovery south-westwards, at the River Onny
the Purple Shales have overlapped the Pentamerus Beds to lie unconformably on
the Onnia superba Zone of the Onnian. South-west from the Onny the Purple
Shales transgress successively lower Caradoc horizons but, as has been shown by
Whittard (1927, pi. 57), they are in turn quickly overstepped by the Wenlock
Shales. Near Sibdon Carwood these rest on strata of Longvillian age.
Professor Whittard informs me that a boring made in search of water at Aston-on-
Clun some years ago proved the presence there of Wenlock Shales. These are not
exposed at the surface, but are presumed to overstep the Costonian near the village
and rest on Pre-Cambrian rocks (Western Longmyndian) . The inlier of Costonian
rocks at Coston is bounded on the west by the Church Stretton Fault, and on the
east by, presumably, unconformable Wenlock Shales, though these are covered by
alluvium of the Clun Valley.
VI. CORRELATION OF THE SHELLY AND GRAPTOLITIC FAUNAS
A great deal of attention has been paid to the problem of fitting the shelly succes-
sion of the Caradoc Series into the graptolite-zones, but no attempt has yet proved
entirely successful or satisfactory. The graptolite-zones were established in the
Lower Hartfell Shales of southern Scotland as follows :
Pleurograptus linearis
Dicranogmptus clingani
Climacograptus wilsoni
Climacograptus peltifer
Nemagraptus gracilis
More recently it has been shown by Jaanusson & Strachan (1953 : 695) that the
same succession cannot be used satisfactorily for both Scotland and Wales, and that
the C. peltifer and C. wilsoni Zones of the Southern Uplands are replaced by a single
Zone of Diplograptus multidens in the Welsh area. The succession of graptolite-
zones which must be applied to the Caradoc Area is then :
Pleurograptus linearis
Dicranograptus clingani
Diplograptus multidens
Nemagraptus gracilis
THE CARADOC SERIES OF SOUTH SHROPSHIRE
227
The presence of the N. gracilis Zone was recognized some years ago when Stubble-
field found the zonal graptolite in strata, loosely termed Hoar Edge Grits, which may
be referred to the Costonian (1930 : 87). In view of the small thickness of these
beds the presence of the whole of the N. gracilis Zone is improbable, and the
Costonian, even in the thickest development at Coston, may represent only part of
the Zone.
One of the fundamental differences in correlation between past works and the
present paper lies in the interpretation of the vertical extent of the D. multidens
Zone. In the Shrewsbury District Memoir (Pocock et al., 1938 : 82) the whole of
the Caradoc Series from the base of the Harnage Shales up to, and including, the
Cheney Longville Flags was assigned to the D. clingani Zone. This correlation made
necessary the acceptance of a large stratigraphical break between the Hoar Edge
Grits and the Harnage Shales, and the missing D. multidens Zone was thought to
be represented by Lap worth's so-called " Transition Bed ". In view of the evidence
put forward earlier in this paper against the existence of a large break at the base
of the Harnage Shales, it is essential to re-examine the foundations upon which the
Geological Survey's argument rests.
The graptolite-species recorded from the Harnage Shales of the Evenwood
District (Pocock et al., 1938 : 250) are listed below, together with their vertical
range according to Elles & Wood (1913 : 516-525) :
N. gracilis
Zone
D. multidens
Zone
Climacograptus cf. brevis Elles &
Wood
C. caudatus Lap worth
C. minimus Carruthers .
Dendrograptus sp. ....
Dictyonema cf . fluitans Bulman
Diplograptus multidens Elles & Wood
var. compactus Elles & Wood
Orthograptus calcaratus (Lapworth)
var. vulgatus Elles & Wood
O. truncatus Lapworth .
O. truncatus var. intermedius Elles &
Wood
O. truncatus var. pauperatus Elles &
Wood
D. clingani
Zone
xr
X
xe
X
P. linearis
Zone
It can thus be seen that, rather than proving conclusively the D. clingani Zone
age of the beds, the assemblage suggests at least the possibility of their belonging
to the D. multidens Zone. According to Stubblefield (in Pocock et al., 1938 : 87)
the three species which enabled both the Harnage and Chatwall Groups to be referred
to the D. clingani Zone were C. minimus, 0. truncatus and 0. truncatus var. intermedius.
Recorded occurrences of Climacograptus minimus suggest this species is more
characteristic of the D. clingani Zone but, on the other hand, Orthograptus truncatus
var. intermedius is common only in the D. multidens Zone. 0. truncatus itself has
228 THE CARADOC SERIES OF SOUTH SHROPSHIRE
often been regarded as characteristic of the D. clingani Zone, but nowadays it is
known to range from the D. multidens Zone upwards into the Ashgillian, with
consequent reduction in its value as a zonal index. A factor strongly in support of
the D. multidens Zone age of the Harnage Shales is that both 0. truncatus and 0.
truncatus intermedium are found with D. multidens and other characteristic forms
in the Caradoc mudstones and shales of the Pontesford outcrop, only a few miles
west of the Harnage Grange District (Pocock et al., 1938 : 91-92).
Of the other graptolites from the Harnage Shales, Climacograptus brevis is
confined to the AT", gracilis and D. multidens Zones, and the Dictyonema is close
to D. fluitans from the Aldress Shales of west Shropshire, now known to belong
to the D. multidens Zone (Whittard, 1955 : 5).
The available evidence for the age of the Harnage Shales thus tends towards
assigning them to the D. multidens, and not to the D. clingani, Zone. Confirmation
of this came when Bulman (1948 : 227) discovered D. multidens itself in the Harnage
Shales of Coundmoor Brook, the type-locality of the Harnagian Stage. The exact
place from which the species was collected is unknown but, as the upper beds of
the Harnagian are faulted-out at Coundmoor Brook, it may reasonably be assumed
that the specimen came from the Reuscholithus reuschi Zone. No graptolitic
evidence is yet available from the topmost beds of the Hoar Edge Grits, which have
yielded S alter olithus ; whether the junction between the N. gracilis and D. multidens
Zones should be placed at the extreme base of the Harnagian as now defined (see
p. n) or slightly higher is not clear but, as the thickness of strata concerned is small,
the possible error involved is correspondingly slight.
The base of the D. multidens Zone having been more or less satisfactorily
established, there remains the problem of defining the upper limit of the zone.
Apart from the Harnage Shales of the Harnage District, none of the horizons within
the Caradoc Series of the type-area has yielded an abundant graptolitic fauna.
A few specimens from the Glenburrell Beds of the Onny Valley have been identified
by Dr. Isles Strachan as Climacograptus antiquus Hall var., and are considered by
him to indicate an age earlier than the D. clingani Zone. The Caradoc shales of the
Pontesford District (Pocock el al., 1938 : 90) belong to the D. multidens Zone although
they contain, in addition to the zonal graptolite, such forms as Orthograptus truncatus
and 0. truncatus intermedius, which were considered to indicate the D. clingani
Zone when found in the Harnage Shales a few miles to the south-west. Re-
examination of the trilobite assemblage at Pontesford shows that it includes the
genera Brongniartella, S alter olithus and Broegget olithus, indicating a Lower
Soudleyan age, and the strata containing them can thus be correlated with the upper
part of the Glenburrell Beds of the Onny Valley, an horizon which they also resemble
lithologically.
Additional graptolitic material from the upper part of the Soudleyan in the
Onny Valley has been identified by Dr. Strachan as Orthograptus cf. apiculatus
Elles & Wood, and is probably indicative of a pre-clingani Zone age. The material
is, however, scanty owing to the arenaceous lithology.
The D. multidens Zone therefore should apparently include at least the lower half
of the Soudleyan and possibly the whole of that Stage. The massive sandstones
THE CARADOC SERIES OF SOUTH SHROPSHIRE 229
of the Lower Longvillian Substage have not yet yielded any graptolites, and their
inclusion in the D. multidens Zone is uncertain. For the present a provisional
line of demarcation between the D. multidens and D. clingani Zones is drawn between
the Soudleyan and Lower Longvillian, though a more convenient level might be at
the base of the Upper Longvillian, as this would coincide with a known faunal
break within the Caradoc Series in parts of the Caradoc Area. Further graptolitic
material will probably enable the margin of error to be reduced considerably, but
many difficulties are encountered when most of the strata involved are shallow-
water sandstones.
The remaining problem concerns the zonal position of the upper beds of the
Series, that is to say, the Actonian and Onnian Stages. The topmost Caradoc
strata of the Onny Valley, presumably the Onnia superba Zone of the Onnian, were
said by Wade (1911 : 445) to contain Orthograptus truncatus var. socialis Lapworth
in abundance, and he accordingly assigned them to the Ashgillian. Bancroft
(1933) equated the Actonian with the Pleurograptus linearis Zone ; no reasons for
this were given, but he may have been influenced by Wade's view that at least
part of the Onnian belonged to the Ashgillian. Whittard (1952 : 162) has drawn
attention to the conflicting statements made by Bancroft at various times regarding
the correlation of the shelly and graptolitic faunas ; these may be summarized as
follows. The upper Stages of the series Costonian to Onnian include the P. linearis
Zone and part or all of the succeeding Zone of Dicellograptus complanatus, the basal
zone of the Ashgillian in its usually accepted sense (Bancroft, 1945 : 181). The
Onnian was said to be succeeded in Westmorland by, first, the Pusgillian Stage,
and then the Ashgillian, the latter apparently being used in a restricted sense, though
few details were given. In the same paper Bancroft (p. 183) described the Actonian
as including " the earliest deposits with Tretaspis kjaeri, Phillipsinella and other
typical Upper Bala fossils ", but on a later page (p. 186) caused considerable con-
fusion by claiming that, at Girvan, " the Actonian and Onnian are represented in the
series of grey flags with f ossiliferous limestones underlying the Zone of Dicellograptus
complanatus (Pusgillian) ". It is thus difficult to see exactly what were Bancroft's
views, but his work shows a definite tendency to include in the Ashgillian the Pus-
gillian Stage and, perhaps also, the Onnian.
During extensive collecting from the upper strata of the Onny Valley Wade's
claim that they contain Orthograptus truncatus var. socialis in abundance has not
been substantiated ; indeed, well-preserved graptolites are exceedingly rare in
both the Actonian and Onnian. In view of this the shelly faunas, in particular
the trilobites, afford the most promising means of assessing the position of the highest
Caradoc beds. The assemblage in the Actonian includes Platylichas, Chasmops,
calymenids and raphiophorids, and closely resembles that of the 4b& fitage, or Upper
Chasmops Limestone, in southern Norway, an horizon known to belong to the
D. clingani Zone. As described earlier in this paper, the Onnian follows the
Actonian conformably and the faunas of the lower portion include some Actonian
elements, so that it would be difficult to claim a much later age for the Onnian.
Supporting data comes from the trilobite fauna, including Triarthrus and raphio-
phorids, which resembles one described by Thorslund (1940) from the D. clingani
230 THE CARADOC SERIES OF SOUTH SHROPSHIRE
Zone of southern Sweden. A few fairly well-preserved graptolites have been obtained
from the Actonian of the Onny, which Dr. Strachan has identified as Orthograptus
of the truncalus group, and Diplograptus (s. s.) sp. He considers that the last-named
cannot be later than the D. clingani Zone, and in view of the evidence of the shelly
faunas outlined above, the youngest strata of the Caradoc Series in the type-section
are concluded to be no later than the D. clingani Zone. There is no acceptable
evidence for the existence of the Pleurograptus linearis or Dicellograptus complanatus
Zones in the Ordovician of south Shropshire.
VII. REFERENCES
BANCROFT, B. B. 19290. Some new genera and species of Strophomenacea from the Upper
Ordovician of Shropshire. Mem. Manchr Lit. Phil. Soc. 73 : 33-65, pis. i, 2.
19296. Some new species of Cryptolithus (s.l.) from the Upper Ordovician. Mem.
Manchr Lit. Phil. Soc. 73 : 67-98, pis. i, 2.
J933- Correlation Tables of the Stages Costonian-Onnian in England and Wales. 4 pp.,
3 tables. Blakeney, Glos. [Privately printed.]
1935- The descent of Genera in Cryptolithinae. In Lamont, A. The Drummuck
Group, Girvan ; a stratigraphical revision, with descriptions of new fossils from the lower
part of the Group. Trans. Geol. Soc. Glasgow, 19 : 333-334.
1945- The Brachiopod zonal indices of the Stages Costonian to Onnian in Britain. /.
Paleont., Menasha, 19 : 181-252, pis. 22-38.
1949. Upper Ordovician Trilobites of zonal value in South-east Shropshire. Proc. Roy.
Soc., London (B), 136 : 291-315, pis. 9-11. [Edited by A. Lamont.]
BULMAN, O. M. B. 1948. Some Shropshire Ordovician Graptolites. Geol. Mag. Lond. 85 :
222-228, 3 figs.
CALLAWAY, C. 1877. On a new area of Upper Cambrian rocks in South Shropshire, with a
description of a new fauna. Quart. J. Geol. Soc. Lond. 33 : 652-671, pi. i.
COBBOLD, E. S. 1900. Geology. In Church Stretton, 1 : ii + 115 pp., 5 pis. Shrewsbury.
[Edited by C. W. Campbell-Hyslop.]
ELLES, G. L. & WOOD, E. M. R. 1913. A Monograph of British Graptolites, 10 : 487-526,
pis. 50-52. Palaeontogr. Soc. [Monogr.] London.
JAANUSSON, V. & STRACHAN, I. 1953. Correlation of the Scandinavian Middle Ordovician
with the graptolite succession. Geol. Foren. Stockh. Fork. 76 : 684-696.
LAMONT, A. 1935. The Drummuck Group, Girvan ; a stratigraphical revision, with descrip-
tions of new fossils from the lower part of the group. Trans. Geol. Soc. Glasgow, 19 :
288-330, pis. 7-9.
LAPWORTH, C. 1916. In " Palaeontological Work ". Summ. Progr. Geol. Surv. Gt Britian,
1915 : 36-38, 3 figs.
LAPWORTH, C. & WATTS, W. W. 1894. The Geology of South Shropshire. Proc. Geol. Ass.
Lond. 13 : 297-355, 23 figs.
LA TOUCHE, J. D. 1884. A Handbook of the Geology of Shropshire. 91 pp., 22 pis. London
& Shrewsbury.
MURCHISON, R. I. 1839. The Silurian System, xxxii + 768 pp., 36 pis. London.
POCOCK, R. W. & WHITEHEAD, T. H. 1948. British Regional Geology. The Welsh Borderland.
2nd edit, iv + 83 pp., n pis. Geol. Surv. Mus., London. .
POCOCK, R. W., WHITEHEAD, T. H., WEDD, C. B. & ROBERTSON, T. 1938. Shrewsbury
District, including the Hanwood Coalfield (One-inch Geological Sheet 152 New Series).
xii + 297 pp., 8 pis. Mem. Geol. Surv. Gt. Britain, London.
SALTER, J. W. & AVELINE, W. T. 1854. On the Caradoc Sandstone of Shropshire. Quart. J.
Geol. Soc. Lond. 10 : 62-73.
THE CARADOC SERIES OF SOUTH SHROPSHIRE 231
STRACHAN, I., TEMPLE, J. & WILLIAMS, A. 1948. The age of the neptunian dyke at Hazier
Hill. Geol. Mag. Lond. 85 : 276-278.
STUBBLEFIELD, C. J. 1930. " Palaeozoic rocks in Shropshire ". Summ. Progr. Geol. Surv.
Gt. Britain, 1929, i: 87-88.
THORSLUND, P. 1940. On the Chasmops Series of Jemtland and Sodermanland (Tvaren).
Sverig. Geol. Unders. Afh., Stockholm (C) 436 : 1-191, pis. 1-14.
WADE, A. 1911. The Llandovery and associated rocks of north-east Montgomeryshire.
Quart. J. Geol. Soc. Lond. 67 : 415-459, pis. 33-36.
WATTS, W. W. 1925. The Geology of South Shropshire. Proc. Geol. Ass. Lond. 36 : 321-363,
figs. 26-40.
WHITTARD, W. F. 1927. The Stratigraphy of the Valentian rocks of Shropshire. The main
outcrop. Quart. J . Geol. Soc. Lond. 83 : 737-758, pis. 56, 57.
1952. A Geology of South Shropshire. Proc. Geol. Ass. Lond. 63 : 143-197.
1955- The Ordovician Trilobites of the Shelve Inlier, West Shropshire, 1 : 1-40, pis. 1-4.
Palaeontogr. Soc. [Monogr.] London.
WHITTINGTON, H. B. 1938(2. The fauna of the Lluest Quarry, Llanfyllin (Wattsella horder-
leyensis Super-zone), and its correlation. Proc. Geol. Ass. Lond. 49 : 49-54, pi. 6.
19386. New Caradocian Brachiopods from the Berwyn Hills, North Wales. Ann. Mag.
Nat. Hist., London (n), 2 : 241-259, pis. 10, n.
I938c. The Geology of the district around Llandsantffraid ym Mechain, Montgomery-
shire. Quart. J. Geol. Soc. Lond. 94 : 423-457, pis. 38, 39.
WHITTINGTON, H. B. & WILLIAMS, A. 1955. The fauna of the Derfel Limestone of the Arenig
District, North Wales. Philos. Trans., London (B), 238 : 397-430, pis. 38-40.
GEOL. 3, 6. 21
EXPLANATION OF PLATES
All specimens in the British Museum (Natural History) except PI. 26, fig. 5, which is
in the Geological Survey and Museum (GSM.).
Photographs by J. V. Brown and C. Horton
PLATE 24
Harknessella vespertilio (J. de C. Sowerby)
Costonian Stage : CostonFarm, east of Clunbury.
FIG. i. Internal mould of dorsal valve. 66.24051. x ij.
Dinorthis flabellulum (J. de C. Sowerby)
Costonian Stage : Coston Farm, east of Clunbury.
FIG. 2. Internal mould of dorsal valve. 66.24049. x i.
Horderleyella plicata 6ancroft
Costonian Stage : disused quarry 500 ft. south of Coston Farm.
FIG. 3. Internal mould of ventral valve. 66.10343. x i£.
FIG. 4. Internal mould of dorsal valve. 66.10344. x i-|-.
Smeathenella harnagensis 6ancroft
Harnagian Stage : southern end of Smeathen Wood, near Horderley.
FIG. 5. Internal mould of ventral valve. 66.9149. x i.
FIG. 6. Internal mould of dorsal valve. 66.9281. x i.
Salopia salteri (Davidson)
Harnagian Stage : southern end of Smeathen Wood, near Horderley.
FIG. 7. Internal mould of dorsal valve. 66.24277. x i^.
FIG. 8. Internal mould of ventral valve. 66.24278. x i-|.
Soudleyella avelinei (6ancroft)
Soudleyan Stage : north-eastern corner of Smeathen Wood, near Horderley.
FIG. 9. Internal mould of dorsal valve. 66.10326. x 2.
Reuschella horderleyensis 6ancroft
Soudleyan Stage : near Glenburrell Farm, south-east of Horderley.
FIG. 10. Internal mould of dorsal valve. 66.9129. x i.
FIG. ii. Internal mould of ventral valve. 66.9134. x i.
Dalmanella horderleyensis (Whittington)
Lower Longvillian substage : Long Lane quarry, north-west of Craven Arms.
FIG. 12. Internal mould of ventral valve. 66.10316. x ij.
FIG. 13. Internal mould of dorsal valve. 66.10315. x i-J-.
Dalmanella indica Whittington
Lower Longvillian substage : Longville Plantation, near Cheney Longville.
FIG. 14. Internal mould of dorsal valve. 66.10325. x ij.
Dalmanella lepta (6ancroft)
Lower Longvillian substage : Rookery Wood, near Horderley.
FIG. 15. Internal mould of dorsal valve. 66.10318. x ij.
FIG. 16. Internal mould of ventral valve. 66.10319. x ij
Bancroftina typa (Whittington)
Lower Longvillian substage : New House quarry, south-east of Horderley.
FIG. 17. Internal mould of ventral valve. 66.10304. x i£.
FIG. 18. Internal mould of dorsal valve. 66.10309. x ij.
Kjaerina bipartita (Salter)
Upper Longvillian substage : Longville Lane, west of Cheney Longville.
FIG. 19. Internal mould of ventral valve. 66.10453. x i.
Kjaerina typa 6ancroft
Upper Longvillian substage : eastern end of 6urrells Coppice, south-east of Horderley
FIG. 20. Internal mould of ventral valve, 66.14383. x i,
Hull. li.M. (N.H.) Gcol. 3, 6
PLATE 24
ZONAL BRACHIOPODS OF THE CARADOC SERIES
PLATE 25
Dolerorthis duftonensis (Reed)
Upper Longvillian substage : eastern end of Burrells Coppice, south-east of Horderley.
FIG. i. Internal mould of dorsal valve. 66.24050. X ij.
Dalmanella wattsi (Bancroft)
Marshbrookian Stage : Marsh Wood quarry, south of Marshbrook.
FIG. 2. Internal moulds of ventral valves. 66.9127. x i.
FIG. 3. Internal mould of dorsal valve and external mould of ventral valve. 66.10341.
x ii.
Dalmanella unguis (J. de C. Sowerby)
Marshbrookian Stage : Marsh Wood quarry, south of Marshbrook.
FIG. 4. Gutta-percha cast from external mould of ventral and dorsal valves. 66.10337.
X I*.
Onniella reuschi 6ancroft
Marshbrookian Stage : Woolston, north of Wistanstow.
FIG. 5. Internal mould of dorsal valve. 66.10233. X Ii-
FIG. 6. External mould of dorsal valve. 66.10256. x i£.
Cryptothyris paracydica (6ancroft)
Actonian Stage : south bank of River Onny, north of Cheney Longville.
FIG. 7. Internal mould of dorsal valve. 66.24172. x 2.
Onniella inconstans 6ancroft
Onnian Stage : north bank of River Onny, south-west of Wistanstow.
FIG. 8. Dorsal aspect. 66.10258. x if.
FIG. 9. Ventral aspect. Same specimen as Fig. 8. x i£.
Onniella broeggeri 6ancroft
Onnian Stage : north bank of River Onny, south-west of Wistanstow.
FIG. 10. Dorsal aspect. 66.24052. x 2^.
FIG. ii. Ventral aspect. Same specimen as Fig. 10. x 2^.
" Rafinesquina " holli (Davidson)
Onnian Stage : " Cliff Section " in north bank of River Onny, south-west of Wistanstow.
FIG. 12. Various isolated valves. 6.13641. x 3.
Hull. 13. M. (N.H.) Geol. 3, 6
PLATE 25
8
10
12
11
ZONAL BRACHIOPODS OF THE CARADOC SERIES.
PLATE 26
Costonia ultima (Bancroft)
Costonian Stage : disused quarry 500 ft. south of Coston Farm.
FIG. i. Latex cast from external mould of incomplete cranidium. 111.48942. x z.
FIG. 2. Latex cast from external mould of incomplete cranidium. 111.48941. x 2.
Reuscholithus reuschi Bancroft
Harnagian Stage : southern end of Smeathen Wood, near Horderley.
FIG. 3. Internal mould. In. 42080. x i|.
Salterolithus smeathenensis Bancroft
Harnagian Stage : southern end of Smeathen Wood, near Horderley.
FIG. 4. Internal mould. ^.49560. x i|.
Salterolithus caractaci (Murchison)
Harnagian Stage : Welshpool, Montgomeryshire.
FIG. 5. Internal mould. GSM. 6829. x i.
Ulricholithus ulrichi (Bancroft)
Harnagian Stage : Middle House Dingle, near Welshpool, Montgomeryshire.
FIG. 6. Gutt-percha cast from external mould. In. 42371. x if.
Broeggerolithus broeggeri (Bancroft)
Soudleyan Stage : north-eastern corner of Smeathen Wood, near Horderley.
FIG. 7. Internal mould. ^.42077. x 2.
Broeggerolithus soudleyensis (Bancroft)
Soudleyan Stage : Soudley Pool quarry, south-east of Hope Bovvdler.
Fr?. 8. Internal mould. In. 493 14. x 2.
Broeggerolithus globiceps (Bancroft)
Lower Longvillian substage : Rookery Wood, near Horderley.
FIG. 9. Internal mould. In. 42076. x 2.
Broeggerolithus longiceps (Bancroft)
Upper Longvillian substage : south-west of Marshbrook.
FIG. 10. Internal mould. In. 42073. X ij-
Broeggerolithus transiens (Bancroft)
Marshbrookian Stage : Marsh Wood quarry, south of Marshbrook.
FIG. ii. Internal mould. In. 49026. x ij.
Platylichas laxatus (McCoy)
Actonian Stage : south bank of River Onny, north of Cheney Longville.
FIG. 12. Test intact. In 46445. x i.
Actonian Stage : Gretton quarry, east of Cardington.
FIG. 13. Latex cast from external mould. In. 49558. x i.
Onnia cobboldi (Bancroft)
Onnian Stage : south bank of River Onny, north-north-east of Cheney Longville.
FIG. 14. Testate specimen. In. 42071. x i£.
Onnia gracilis (Bancroft)
Onnian Stage : north bank of River Onny, north-east of Cheney Longville.
FIG. 15. Testate specimen. In. 42074. x 2.
Onnia superba (Bancroft)
Onnian Stage : " Cliff Section ", in north bank of River Onny, south-west of Wistanstow.
FIG. 16. Internal mould. In. 49031. x ij.
Bull. B.M. (N.H.) Gcol. 3, 6
PLATE 26
ZONAL TRILOBITES OF THE CARADOC SERIES.
FROM THE UPPER TRIAS OF
NEW SOUTH WALES
D. M. S. WATSON
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 7
LONDON : 1958
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following papers appeared in Volume i (1949-52) :
Price
No. i (1949). The Pterobranch Rhabdopleum in the English Eocene
H. D. Thomas & A. G. Davis . . . . ?5. 6rf.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu 55.
No- 3 (i95°)- The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White.
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White ^s. (>d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
Northern Iraq. L. F. Spath IO5.
No- 5 (I951)- Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum. T. H. Withers 5S
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae)
T. H. Withers .
55.
No- 7 (I952). A New Trochiliscus (Charophyta) from the Downtonian
of Podolia. W. N. Croft ..... IO5
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsdale Ios
No. 9 (1952). Australian Arthrodires. E. I. White . .,• . I5S.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.
The following papers appeared in Volume 2 (1953-56) :
No. i (1953)- The Deer of the Weybourn Crag and Forest Bed of
Norfolk. A. Azzaroli /i 55
No. 2 (1953)- A Coniferous Petrified Forest in Patagonia. M. G.
Calder I25
No- 3 (*953)- The Solution of the Piltdown Problem. J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark . . . . .'35. 6rf.
No. 4 (1954)- Some Upper Cretaceous and Eocene Fruits from Egypt
M. E. J. Chandler l6s
No- 5 (I954)- The Carboniferous Flora of Peru. W. J. Jongmans . 155.
No. 6 (i955)- Further Contributions to the Solution of the Piltdown
Problem. J. S. Weiner, W. E. Le Gros Clark & K. P. Oakley
gt &1- •-.... /T
No- 7 (1955)- The Schizaeaceae of the South of England in Early
Tertiary Times. M. E. J. Chandler .... 155.
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . /i
A NEW LABYRINTHODONT
(PARACYCLOTOSAURUS) FROM THE
UPPER TRIAS OF NEW SOUTH WALES
BY
DAVID MEREDITH SEARES WATSON, F.R.S.
Trustee of the British Museum
Pp. 233-263 ; Pis. 27-31 ; 16 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 7
LONDON: 1958
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, *s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 7 of the Geological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1958 Price Fifteen Shillings
A NEW LABYRINTHODONT
(PARACYCLOTOSAURUS) FROM THE
UPPER TRIAS OF NEW SOUTH WALES
By D. M. S. WATSON
CONTENTS
Page
I. INTRODUCTION .......... 235
II. SYSTEMATIC DESCRIPTION OF Paracyclotosaurus davidi gen. et sp. nov. 237
Braincase .......... 237
Palate ........... 243
Lower jaw ...."...... 244
Jaw musculature ......... 245
Vertebral column ......... 245
Ribs . . ... . ... . . . . 247
Shoulder girdle ......... 249
Pelvis ........... 251
Skin ........... 252
III. BODY SHAPE AND MODE OF LIFE ...... 253
IV. SYSTEMATIC POSITION ........ 254
V. DESCRIPTION OF Subcyclotosaurus brookvalensis gen. et sp. nov. . 258
VI. ASSOCIATED BRACHYOPID ........ 259
VII. ACKNOWLEDGMENTS ......... 260
VIII. REFERENCES .......... 261
IX. KEY TO ABBREVIATIONS IN TEXT-FIGURES ..... 263
I INTRODUCTION
THE Namurian rocks of New South Wales containing characteristic plants found
also in Scotland, are succeeded by a great series of beds with a total thickness of
14,000 feet. The upper part — some 3,500 feet of this — forms the Hawkesbury
Series, whose lowest element, the Narrabeen Beds, may perhaps be held to include
the fish fauna from Gosford, described by Woodward (1890) and revised by Wade
(1940). In the middle of the succeeding Hawkesbury Sandstone is the very rich
fish fauna of Brookvale, described by Wade (1935), and that of the Wianamatta
shales (which rest on the Hawkesbury Sandstone) of St. Peter's near Sydney,
described by Woodward (1908) and revised by Wade (1941). The St. Peter's fish
occur in shales, and the ironstone nodules in them, worked in a series of brick pits
near Sydney. They were collected by Mr. B. Dunstan of the Geological Survey
of New South Wales.
GEOL. 3, 7. 22
236 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
In one of these pits, about 1910, Mr. Dunstan found fragments of an enormous
ironstone nodule which by most careful search he was able to reconstruct completely.
The nodule contained a complete labyrinthodont skeleton which I saw in July 1914.
It was immediately obvious that the animal was closely related to Cyclotosaurus ,
then known only by specimens from the lower part of the Upper Trias of Germany.
This gave a well fixed correlation for at least one point in the Hawkesbury Series,
the only one, indeed, in the whole succession of rocks set out above. It was therefore
desirable that the position of " Cyclotosaurus " as a descendant of " Capitosaurus "
should be established, and the time range through which it lived should be determined.
This I undertook to do, and Sir Edgeworth David, after whom the species is named,
arranged for the British Museum (Natural History) to acquire the specimen.
MATERIAL. The nodule (B.M.N.H. R.6ooo), some 9 feet (2-75 metres) long,
is split roughly into dorsal and ventral halves, each composed of more than fifty
blocks, some of which weigh nearly a hundredweight. The matrix is an extremely
hard and brittle ironstone, quite impossible to work. As the bone was largely
rotten, and much of it already lost, Mr. F. O. Barlow of the Museum staff removed
all the remains of bone, even from the deepest fissures in the blocks. Many of the
blocks are very heavy and irregular in shape, and it was impossible to place them
together, run in a flexible casting medium, and so draw out complete casts of indi-
vidual bones. In practice the cavities in a block and its counterpart had to be cast
separately in glue. From the glue impression a waste mould was prepared from
which, in turn, a plaster positive was made. This was then trimmed until it fitted
accurately the cast of the same bone similarly prepared from the counterpart. The
two parts were then fitted together and cast in a jelly mould which yielded perfect
replicas that for all practical purposes are as good as the original bones. Details
of the braincase, however, can only be determined by supplementing them with
flexible plastic casts. Barlow's work covered many years, and I do not know of
any other man who could have done it ; it was a technical triumph.
The articulated skeleton is complete, but the skull and lower jaws are distorted
in a remarkable way. The mode of attachment of the head to the vertebral column
is very difficult to determine, and some bones were broken and misplaced, evidently
before burial. The fractures all lie in the same region, that of the hinder part of the
head, lower jaws, and interclavicle. The skull, seen from above, is asymmetrical,
the orbits evidently misplaced to the right. In direct dorsal view the lateral border
of the right orbit lies 7 cm. from the margin of the skull at the front of the squamosal,
the corresponding measurement on the left side being 11-5 cm. On the right side
the outer surface is smoothly rounded from the mid-line at the parieto-dermosupra-
occipital suture to the lower border of the quadratojugal. On the left the surface
is concave and broken by a crack which has a displacement of a centimetre. This
distortion extends forward to the anterior end, but scarcely affects the triangle
of bone between the pineal foramen and the occipital border. On the palate the
parasphenoid is broken at the ends of the sphenethmoid, the right pterygoid is
crossed obliquely by a crack running backward from the hinder end of the right
palatal vacuity, and the two occipital condyles are forced into contact. The left
lower jaw seems to be undistorted, but the right is crushed down just in front of
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 237
the opening through which the temporal muscles passed into it, so that its depth was
decreased, its width increased, and the jaw as a whole bent upward and inward. These
distortions agree with those of the palate. The interclavicle is crossed obliquely
by a crack, matching those on the palate and skull roof, which posteriorly, on the
left side, has a displacement of nearly one centimetre.
These breaks, and the displacements which result from them, lie almost entirely
in the region behind the eyes, and can only reasonably be interpreted as the results
of a single heavy blow, for it broke three horizontal sheets of bone — the skull roof,
palate and interclavicle — each almost two centimetres thick. All these bones were
surrounded and widely separated by soft tissues. Furthermore it appears from the
results that such a blow must have been delivered by a massive cylindrical body,
such as a tree trunk from the bank, falling across the animal ; the body afterwards
being washed into the bottom of the lake.
II. SYSTEMATIC DESCRIPTION OF PA RA C YCLOTOSA UR US DAVIDI
gen. et sp. nov.
Braincase. The braincase is almost completely underlain by the hinder part of
the parasphenoid, which except for a short border between the exoccipitals, has con-
tinuous sutural attachments to them and to the pterygoids, the details being obscure.
The ventral surface of the parasphenoid posteriorly bears a very low transverse
ridge, behind which its surface is depressed, a condition associated with the attach-
ment of recti capitis muscles. The greater part of its area is, however, shallowly
concave, continuous with that of the pterygoid attached to it, whose surface turns
down laterally. The whole directly supported the skin of the roof of the mouth.
At the hinder margin of the palatal vacuity, where the pterygo-parasphenoid suture
ends, the bone narrows, the narrowest point of the " processus cultriformis ", some
10 cm. in front, being only 19 mm. wide, with a cylindroid surface and a shallow
groove — presumably occupied in life by a cartilaginous septum — on its upper surface.
It then continues forward, widening and having a flat lower surface, until it is
overlapped by the vomers, and ends by rising up above the junction of these bones.
Each exoccipital bears a large hemispheroidal condyle covered by cartilage during
life to an extent shown by a projecting ridge, which marks the end of the dense
bony sheet that coated its outer surface. Viewed as a pair it is evident that the
condyles allowed dorso-ventral motion of the head, perhaps through a large arc,
and inhibited any horizontal movement. The lateral surface of the condyle passes
forward and outward to a suture with the pterygoid, forming a thick rounded flange,
the floor of the tympanic cavity. Above the inner part of the condyle a thin flange
passes horizontally inward above the space occupied by the persistently cartilaginous
basioccipital. Its upper surface forms the floor of the brain cavity and laterally
curves upwards and then slightly inwards to form the side walls of the brain cavity.
A foramen in the lateral grove leads into a canal passing through the bone, which
opens on the posterior surface of the exoccipital just above the condyle. It trans-
mitted a Xllth nerve. Damage obscures the exit of the vagus. The upper part of
the exoccipital is continuous with the descending process of the dermosupraoccipital
238
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
and of the tabular. These are massive bones separated by a triangular post-
temporal vacuity.
It is natural to suppose that, as in all earlier capitosaurs, there should be an
abruptly truncated projecting process, mesial to the junction of the dermosupra-
occipital and exoccipital, which supported a cartilaginous supraoccipital. On the
PMx
Int.Nw
Vo
FIG
, ExOc i .
PorSp Tab Q"
. i. Paracyclotosaurus davidi gen. et sp. nov. The palate, x 1/6. (For explanation
of lettering see p. 263.)
left side the surface of the two bones is perfectly shown and there is no trace of
the process, but the sculpturing seems to show that the surface abutted on to a
supraoccipital cartilage. Dorsally there remains a space on the under surface of the
dermosupraoccipitals, partially subdivided into three by low ridges, which was clearly
open and perhaps occupied by blood vessels above a supraoccipital. This unusual
arrangement seems to be well established here.
The position of the supraoccipital and paroccipital can only be inferred as there
is no trace of ossification in them. A small shapeless ossification, high up in the
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
239
prootic, is attached to the summit of the pre-tympanic flange of the left pterygoid.
On its inner side, at the hinder edge, is a narrow groove forming a quadrant of a
circle. Presumably it housed the anterior vertical semicircular canal.
Fortunately the stapes is in position on the right side, giving some indication of
the relations between the prootic, paroccipital and exoccipital. The cartilaginous
paroccipital must have been a small mass, meeting the prootic above the fenestra,
lying in contact above with the exoccipital, and separated from it below by the vagus.
Primitively the paroccipital reached the occipital process of the tabular. In the
present specimen that process passes downward and inward as a thick sheet of bone,
which meets and fuses with the corresponding process of the dermosupraoccipital
and the exoccipital, the sutures being invisible.
DSOc
FIG. 2. Paracydotosaurus davidi. The occiput, x 1/4.
The basisphenoid is entirely unossified. The impression of its lower surface on
the parasphenoid, however, is easily interpreted by comparison with Eryops, and with
a remarkable specimen of Rhinesuchus (B.n8, D. M. S. W. Coll.). The key feature
is the deep groove running transversely, housing a thickening of the lateral part of
the basisphenoid, which extends into a pit excavated mainly in the mesial border
of the pterygoid, and is roofed by the foot of the epipterygoid ; this is the basip-
terygoid process. The groove is supported behind by a deep flange which extends
outward, increasing in depth as it does so, to end on the border of the fenestra
ovalis, where the accessory process of the stapes is attached to it.
In front of the prootic the floor and side walls of the braincase are entirely unossified,
but the epipterygoids which lie immediately lateral to it (and are in part modelled
on the Gasserian ganglion) are well preserved. They are described later, with the
quadrate and pterygoid.
The anterior part of the brain was set within a massive bony sphenethmoid. This
is a single structure, lying in the grooved upper surface of the processus cultriformis
of the parasphenoid, and reaching the dermal roof of the skull between, and in front
of, the eyes. It has the following measurements : length about n cm., depth
(including the parasphenoid) posteriorly 6-5 cm., and anteriorly 7-5 cm., maximum
width at the upper edge 3-6 cm. posteriorly and 5-3 cm. anteriorly (all figures being
240
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
approximations). The brain cavity within, measured on a natural cast, is posteriorly
2-6 cm. deep and 2-0 cm. wide. This cavity was divided into two independent canals
(for olfactory lobes?) 7-0 cm. in front of the hinder end. There is no trace of any
further ossification of the braincase, and no evidence about the olfactory capsules.
The ossifications in the palatoquadrate cartilage are well shown, and as they are
attached to the braincase through the intermediary of the pterygoid, it is convenient
to begin by describing this bone. The pterygoid is attached by a long suture to the
lateral border of the parasphenoid and the front of the exoccipital. The attachment
is by interlocking surfaces about 2-5 cm. thick, and about 15 cm. long. From the
attachment the bone stretches outward, turning downward as it does so, the palate
between the subtemporal fossae being concave and cylindrical. The palatal ramus
PrOt.
ExOcCov
SpKEtK
FIG. 3. Paracyclotosaums davidi. Reconstruction of longitudinal section through the
braincase along the middle line, x 1/3. The " brain " in the sphenethmoid is a
drawing of a natural cast of the cavity.
turns forwards and outward to meet the narrow strip of palate lateral to the palatal
vacuity, the suture being unseen. The quadrate ramus, which is well shown (Text-
figs. 3, 4), is essentially a sheet of bone that rises abruptly from the flat lower part
attached by suture to the parasphenoid. It has a rounded edge admesially, which
rises immediately above the deep ridge on the parasphenoid limiting the basipterygoid
process behind, to extend upward and end some distance below the skull roof. The
prootic is attached to this edge. From this mesial border the paraotic flange of the
pterygoid rising to the skull roof, passes outward swinging round a conical surface
until it sweeps backward and outward to pass to the quadrate. There it ends, the
two bones meeting in a thick abutment, rather than in a deep overlap. On its
hinder surface (Text-fig. 2) the quadrate ramus of the pterygoid is divided into two
areas ; the mesial (the anterior wall of the tympanic cavity) is a hollow cone, trun-
cated below where the pterygoid and exoccipital meet, rising until it reaches the
anterior border of the hole in the skull roof which housed the tympanic membrane.
Laterally the pterygoid forms a triangular area that is attached to, but rises above,
the quadrate. The summit of this area is sculptured for some other attachment,
presumably that of a lateral wall of the tympanic chamber,
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
241
There are two ossifications in the palatoquadrate cartilage, which apparently
persisted between them. The posterior, the quadrate, is a massive bone whose very
wide lower surface is entirely occupied by the condyle for the articulation of the lower
jaw. Above this the bone rises as a solid sheet (22 mm. thick above the thickened
articular lower part) surrounded by the pterygoid, squamosal and quadratojugal,
which together hold it in place. The attachment to the pterygoid is almost edge to
FIG. 4. Paracyclotosanrus davidi. A, the braincase and the parts in relation to it seen
from above, with the dermal skull roof removed, and the right quadrate and quadrato-
jugal cut at a lower level. B, the left epipterygoid viewed directly from in front. The
drawing represents an actual glue cast from the original cavity, x 1/3.
edge, rather than the extensive overlap usual in labyrinthodonts. Ventrally,
just above and mesial to the condylar surface, the quadrate ends in a narrow surface
from which a cartilaginous strip 5 mm. thick extended forward, lying in a groove
in the pterygoid, until it met an abrupt face on the hinder and lower border of the
epipterygoid.
The epipterygoid is well ossified, and elaborate (Text-figs. 3, 4). Its base rests
on the upper surface of the pterygoid, where that bone passes forward and outward
behind the palatal vacuity, and was continued by cartilage back to the quadrate,
GEOL. 3, 7.
2 3
242
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
and perhaps a little forward and outward. On the inner surface the bone has a
projecting boss abruptly truncated where it was attached to the side of the cartila-
ginous basis cranii. Above this level the inner surface of the bone is crossed
horizontally by a deep groove, bounded above and below by shallow but projecting
flanges. In this the Gasserian ganglion lay, the ophthalmic branch of that nerve
passing outward and forward in front, and the maxillary and mandibular branches
PMx.
MX
Na
Lac
Orb.
FIG.
Tym M
5. Paracyclotosaurus davidi. The skull viewed from above, with its distortion
corrected, x 1/6.
outward and upward over a rounded border above the pterygoid. Above this
groove the bone extends upward until it ends, its admedian surface evidently
embedded in a thick cartilaginous side wall of the braincase (between the prootic
and the sphenethmoid) . On the outer surface the bone has a deep notch separating
the attachment to the pterygoid from the lower end of a special admedian process
extending downward and a little forward laterally to the basisphenoid cartilage, and
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
243
perhaps also to the parasphenoid. This notch must have been converted into a
foramen by the parasphenoid or pterygoid, and no doubt transmitted a blood
vessel.
The remainder of the skull consists of a series of membrane bones, attached to one
another by interdigitating sutures which are not often seen. The whole outer surface
is covered with bone except for orbits, nostrils, the pineal foramen, and the completely
enclosed openings in which lay the tympanic membranes. The dermal bone coating
the head is massive, about 2 cm. thick, and the individual bones of which it is com-
posed were so tightly attached to one another by interlocking sutures that they
withstood the blow from which the animal presumably died. In the cast actual
fe
No L,a
Nostril
PMx
SP1
FIG. 6. Paracyclotosaurus davidi. Skull and lower jaw seen from the left, x 1/6.
sutures are seldom visible, but in most places the distribution of the pitted and grooved
ornament determines their direction rather precisely. The general skull pattern
differs very little from that found in other capitosaurs ; the only striking peculiarity
is the closure of the otic notch by a fusion behind it of the outer end of the tabular
with a special shelf passing backward from the hinder border of the squamosal in
front. The tabular is an unusually large bone, forming a wide upper surface behind
the tympanic space. It is nearly as long from back to front as the dermosupra-
occipital.
Palate. The palate has the normal pattern of an advanced labyrinthodont.
The quadrate condyles, though ventral to the occipital condyles, lie essentially in
the same transverse plane, so that the quadrate ramus of the pterygoid is short
and unusually laterally directed. The sub-temporal opening through which the
jaw-closing muscles passed down to the lower jaw is large, but not unusually so ;
244 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
and the flange formed by the ectopterygoid and pterygoid, which in labyrinthodonts
is usually applied to the inner surface of the lower jaw, is vestigial. It is impossible
to determine the sutures between the pterygoid, ectopterygoid and palatine ; but
it is evident that the three bones form the outer border of the large palatal vacuity,
their respective parts being indeterminable. The pterygoid is toothless. The
palatine bears an alternative pair of large teeth, about 2-6 cm. high, immediately
behind the internal nostrils, set as always in a common recess. A similar tooth
seems to be shown on the vomer. Behind these " tusks " is a single series of teeth set
close to the maxillary suture of the palatine and ectopterygoid bones, numbering in
all 56 to 60. The series is essentially uniform, teeth being shed and replaced occasion-
ally. An individual tooth is attached to the bone on the bottom of a very shallow
depression. It is a triangular structure, flattened antero-posteriorly, attached to
the bone by a base rather more than twice as wide across the jaw as it is long. The
outer side is rounded and convex in profile, the inner a little concave, the point
(where it might be expected to meet the teeth of the lower jaw) is also rounded and
smooth. The tooth is as high as it is wide at its base, and separated from its neigh-
bours by about its own thickness, the height being uniformly 9 or 10 mm. only.
The elongated, narrow internal nostril, whose inner border is probably largely made
by the vomer, is bordered by a row of similar but much smaller teeth which ends
posteriorly at a short set of some six or seven very small conical teeth mesial of the
first palatine tusk. The maxilla bears a close-set series of teeth, which decrease
somewhat in size backwards. These teeth agree in structure with those of the palate,
from which they are separated only by a very narrow wedge-shaped continuous groove.
In all probability the series continued round the premaxilla.
The teeth of the lower jaws, carried entirely by the dentaries, are slightly enlarged
copies of those in the upper jaws. They bite within the maxillary row and laterally
to the palatal row.
The only large teeth — those on the palatines and vomers — are about an inch high,
little higher than the marginal teeth, and were evidently unsuitable for the capture
of any big animal, into whose body they could not penetrate far enough to kill.
Further consideration of the animal's diet is left until after the description of the
whole skeleton.
Lower jaw. The lower jaw is well preserved and has a normal labyrinthodont
structure. The articular is fused with the surrounding bones, and is articulated
with the quadrate, so that no complete description can be given. This articulation
is extremely wide, fitting the quadrate condyle and rising mesially as a powerful
process to articulate with the front face of the inner end of the quadrate condyle.
The bone has an analogous process rising behind the outer condyle, to
form the hinder border of a deep cylindroid notch, which assures accurate closure
of the mouth. Behind this a massive conical retroarticular process projects directly
backward for about 8 cm. behind the axis of the articulation. The surangular,
fused with the articular, projects forward, bounding the 19 cm. long suprameckelian
vacuity through which the masticatory muscles pass down into the hollow jaw. A
prearticular bone, attached to the inner side of the articular, stretches forward
parallel to the surangular forming the inner border of the muscle opening, which is
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 245
closed at the front by the third coronoid, a narrow bone wedged in between the dentary
and the prearticular. There is a series of apparently three coronoids attached to
the inner surface of the dentary, widely exposed above the splenial and postsplenial
bones. Their upper border may lie free above the shelf of the dentary to which they
are attached ; none bears any teeth, though they bound and held in place the strip
of gum from which dentary teeth arose.
Jaw musculature. The nature of the ornament on the outer surface of the lower
jaw makes it certain that the masticatory muscles can have had only an infinitesimal,
if any, attachment to the outer surface. The flat upper surface of the surangular,
which stretches forward from immediately in front of the articular surface, seems
unlikely to have given attachment to a muscle. Its character and width are similar
throughout its length, and a muscle attached immediately in front of the condyle
could have very little value. It probably represents the limit of the cheek, lateral
to the masticatory muscles. These would arise from the skull roof and pterygoid,
passing downward and inward, through the opening between the surangular and
prearticular, to the cavity of the jaw, and probably extending forward past the large
inner fenestra to an insertion on the inner surface. A muscle arising from the skull
roof, at the attachment of the pterygoid to it, and passing down to the upper surface
of the surangular at its mid-length, would.be only some 10 cm. long, and perhaps
placed about the same distance from the articulation ; a range of shortening by
a quarter of its length would allow the mouth to open only about 15 cm. at the front
of the jaw. An attachment to the point within the cavity of the jaw immediately
below would double the depth of the opening, and an extension forward to the front
of the internal mandibular vacuity might well double it again, the palate then being
nearly at right angles to the jaw. The relative position of the occipital and quadrate
condyles shows that if the lower jaw rested on the ground, as must often have been
the case, it would be moved forward as the mouth was opened by raising the skull.
Vertebral column. The vertebral column is articulated from head to tail, and in
general is undisplaced. In Mastodonsaurus (which is not remote from the Capito-
saurs) the ist vertebra is a continuous structure whose anterior face bears a pair of
articular facets for the occipital condyles, below and between which a canal for the
notochord opens. This passes obliquely through the bone, and opens behind, a
little below the neural canal. There is a small fused neural arch, and no rib facet.
No such structure can be found in this specimen. The blow which the body suffered
on the left side of the head disarticulated that structure, so that the occipital condyle
lies some 6 or 7 cm. to the right of the mid-line of the first two recognizable elements.
Of these the anterior, which is perhaps the intercentrum of the ist vertebra, differs
from all others throughout the column. The rest are crescentic — half rings surroun-
ding a large notochordal space — their upper ends narrowing and eventually becoming
rounded. The 2nd intercentrum is peculiar only in that it is divided into right and
left halves meeting below the notochord by extensive flat surfaces. The ist inter-
centrum is also paired, and the left element (which alone is known) lies nearly in
articulation with the corresponding bone of the 2nd, differing from this (like the
corresponding bone of Eryops) by the absence of an upstanding process lateral to the
notochord. Its anterior face does not show any definite cup for articulation with the
246 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
rounded condyle. A left half of a neural arch, though found behind and above the
left half of the 2nd intercentrum, probably belongs to the 3rd intercentrum, which
has none associated with it. It corresponds generally with those found further back
in the column, having similar cartilage-covered facets for articulation with pleuro-
centra, a short transverse process truncated by a rib facet, and a peculiarly small
anterior zygapophysis with a rounded articular surface. Thus we have to find
neural arches for vertebrae i and 2, and the right half of 3. Lying dorsal to and on
the right of the two anterior intercentra are six strange bones, forming a dorsal
series of two, a ventral of three, and still further down a fragment which may be a
rib head. These bones should be determinable as the missing neural arches, but I
am unable to identify them.
From this point backward the vertebral column is continuous, and the ribs
attached to it are identifiable. There is a systematic change in structure from front
to back. Such vertebrae as nos. 5, 6 and 13 (see Text-fig. 7) have a large semi-
cylindrical intercentrum whose upper border has a semicircular notch for the
notochord, on either side of which the bone was rounded and presumably capped
with cartilage. In side view the bone is somewhat wedge-shaped, its mid- ventral
border being widest, and the lateral surface is recessed between out-turned ridges.
Toward the summit, near the hinder edge, is a large shallow concavity, offset from
the outer surface, for the capitulum of the rib. The neural arch, apparently
composed of paired elements in many parts of the column, has a short neural spine
rising above the posterior zygapophyses and ending abruptly. Its anterior face
turns forward to form anterior zygapophyses, below which lie the two sides of the
neural canal, a relatively large cylindrical space. The lower surface of the neural
arch is cut out to form the roof of the neural canal, on each side of which it bears
a nearly rectangular facet for articulation with the cartilaginous pleurocentrum.
Laterally to this facet lies a transverse process of varied length whose outer end is
abruptly truncated by a rib facet.
Observation shows that mid-ventrally the intercentra met one another with a
minimum of ligament between them, so that a measured length of 20-5 cm. is occupied
by six intercentra — with individual lengths mid-ventrally of 3-4 cm., 2-9 cm. and
3-1 cm. in three directly measurable cases. The more anterior intercentra are
massive structures nearly as high as they are wide, with a cylindroid ventral surface.
No. 13 is perhaps the extreme of this condition ; its intercentrum is wide and high,
3-6 cm. in length, and remains massive almost up to the extreme dorsal points which
carry the outwardly and backwardly directed shallowly concave articulations of
the ribs. By no. 21 the intercentrum, still 3-5 cm. long, has become very shallow,
about 3-6 cm. in contrast with 5-5 cm. in no. 13. No. 27 has an intercentrum
3-9 cm. long, 7 cm. in maximum width (compared with 9-2 cm. in no. 13). The inter-
centrum has a flatly cylindrical lower surface and almost straight, vertical, lateral
surfaces, with large rib facets placed posteriorly and extending very nearly to its
summit. The transverse process is short, extending only 2 or 3 mm. on the ventral
surface beyond the intercentrum. No. 28, the sacral, does not show the intercentrum,
but its neural arch differs in the depth and massiveness of its transverse process.
In nos. 32 and 33 the neural arches are fused on the left side, all trace of zygapophyses
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
247
having vanished, but on the right the neural spines are not continuous, and the
anterior zygapophysis of no. 33 is well shown. The intercentra are very square cut,
with an essentially flat ventral surface, and a large facet for the rib head filling the
hinder part of the lateral surface. There is a very small, short, down-turned trans-
verse process, and the rib is articulated by two heads. The positions of the completely
cartilaginous pleurocentra are well shown.
NSP
Tran Proc.
I Cm.
FIG. 7. Paracyclotosaurus davidi. Vertebra No. 13. A, from in front. B, left side,
c, behind. D, from above. E, below, x 1/4.
These conditions continue down the tail, haemapophyses appearing at no. 36.
Here the intercentrum is little more than a quadrangular sheet of bone, a little
hollowed above, bearing a large downward and slightly backwardly directed chevron
below and with a rib facet on the posterior end of its lateral border. The neural
arch has a well-defined neural canal, lateral to which it extends out to a thickened
and rounded end, which is presumably the upper part of the rib facet. Pre- and
post-zygapophyses are well formed, with their articular faces at little more than
right angles to one another.
Ribs. Ribs are in position from the first vertebra until they cease at the 36th.
The first one is seen from below on the left side. It appears to be single
headed, and is 7-6 cm. long. Proxirnally it is a thin, nearly vertical, bony
sheet, but distally a peculiar ridge runs from its anterior surface making
the bone about 2 cm. in maximum thickness. It extends outward practically hori-
zontally. The upper end of the second rib is more than 3 cm. deep, and had a
cartilaginous head which articulated with the appropriate facet on the intercentrum
and the unknown neural arch. The bone is wide proximally, then becomes nearly
circular in section, but widens distally, foreshadowing the conditions in those which
follow. The ribs belonging to vertebrae 3, 4 and 5 are remarkable (cf. PL 29), for
they are attached to the flat outer ends of the transverse processes, bone to bone,
248 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
with little space remaining for cartilage between them. But the capitular part of
the wide upper end of the rib fails to meet the facet on the intercentrum, to which
it was attached in life, by about two centimetres, obviously a measure of the thickness
of the cartilaginous capitulum. The rib then stretches out laterally and backward
at about 45° to the mid-line. The ribs attached to vertebrae 3, 4, 5 and 6 are peculiar
because their distal parts are widened so that they overlap one another and provide
a large base of attachment to the muscles, the serrati anteriores, passing to the
scapula. This widening is abrupt ; the slender shaft of the rib, distal to a dorsal
ridge it bears, is roughly circular in section, and later rapidly widens to become at
least 8 cm. across, the lower border being inturned as a flange. Ribs, 3, 4 and 5 thus
combine to form a firm, powerful, flat surface some 8 x 20 cm. for the attachment
of the scapula. From the seventh or eighth vertebra backward the ribs are
FIG. 8. Paracyclotosaurus davidi. Occipital condyle and selected vertebrae numbered
below i, 2, etc. No. 28 is the sacrum ; note large rib facet. Nos. 32 and 33 are
abnormal in the continuity of their neural arches on this side, x 1/5.
essentially straight, simple, bony rods. Both sacral ribs are preserved, supple-
menting one another so that the complete structure is shown. The head, whose
confluent faces carried a single cartilage cap which articulated with the transverse
process and intercentrum of the sacral vertebra, is 6-2 cm. deep, with a maximum
thickness of 2 cm., so that its attachment to the vertebral column is powerful.
The bone rapidly narrows, and at 7 cm. from the attachment is only about 2 cm.
thick. It then widens, its anterior surface being hollowed out so that it ends in two
separated processes which must have met and been attached to the inner surface
of the ilium a little below its upper border. The area of such contact is very
small, the attachment being presumably reinforced by ligaments, as it presumably
was in Eryops and certainly was in embolomerous amphibia. It seems clear that
there is only a single sacral. The anterior caudal vertebrae still carry two-
headed ribs.
The vertebral column is thus largely cartilaginous, but with well-formed zygapo-
physeal articulations and rib attachments. It was evidently capable of lateral
movement though it is improbable that this would be through any extensive arc —
except in the tail which curves round abruptly at nearly a right angle. Flexure in
a dorso-ventral direction was probably very restricted,
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
249
Shoulder girdle. The primary shoulder girdle is most easily discussed after the
membrane bones which carry it have been described. The interclavicle is a sheet
of bone a little longer than wide ; the anterior end stretches forward as a narrow
process and the middle strip may have been visible on the lower surface between
the clavicles. The bone otherwise is rhomboidal with its three other corners rounded.
Its visceral surface is shallowly concave a little in front of the mid-point, and quite
posteriorly there is a small, shallow, but well-marked depression which is somewhat
asymmetrical, opening backward to the left of the mid-line and presumably support-
ing the heart. The ventral surface is very largely covered by the usual pitted and
FIG. 9. Paracyclotosaurus davidi. Interclavicle and clavicles in articulation, from below.
X 1/6.
ridged " ornament ", which extends to the borders of the bone in its posterior third,
but ends abruptly laterally where the ventral surface has a sudden step dorsally
for a centimetre or more, forming a recess into which the lower end of the clavicle
fitted.
The clavicles are almost completely preserved and undistorted. They fit the
recesses in the interclavicle sufficiently well to make the whole structure certain ;
it is possible that they met anteriorly below the interclavicle.
The dorsal process of the clavicle is formed by an upturning of the lateral border
of the bone, so that its outer surface lies at about 60° to the upper surface of the clavi-
cle, or some 55° to the ground. This dorsal process is firmly attached to the ventral
part of the bone by a base about 13 cm. long and (at its maximum) 3-5 cm. thick.
The process is about 14 cm. high and 12 cm. antero-posteriorly ; its upper border
slopes down a little in front, and the outer surface below it is excavated into a pocket,
3, 7. 2
250
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
about 4 cm. deep, across its whole width, presumably for a musculus cleido-mastoideus.
Below it the anterior part of the lateral surface is recessed on a definite oblique
line.
Clei.
C/eido-mast Mus.
\
a
FIG. 10. Paracyclotosaurus davidi. Left clavicle in articulation with the cleithrum and
scapula, x 1/6.
Ctei
Scop
Clav
i/
' Heart "
FIG. ii. Paracyclotosaurus davidi. The shoulder girdle in position with respect to the
vertebral column and ribs, viewed from behind, x i /6. (The intercentra and ribs belong
to the second vertebra, the neural arch, to the third.) The figure shows the inclination
of the dorsal processes of the clavicles towards one another as they pass forward and
upward, so that the structure may lie between the hinder ends of the lower jaws. The
modelling of the upper surface of the interclavicle to support the heart and other
structures is shown.
The left cleithrum remains firmly attached to the scapula. It is a massive rod (4 cm.
by 3 cm. in section at mid-height) which, beginning at a " blunt point " at the lower
end of the scapula, rises to overhang the upper end of that bone by some 4 cm.
The overhang slopes backward as a partial cap to a cartilage. The anterior face of
the lowest 9 cm. of the bone is shallowly concave, and articulated with the hinder
border of the dorsal process of the clavicle. Thus the position of the scapula is
known with certainty, and that of the glenoid cavity can be inferred with accuracy.
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
251
The shoulder girdle thus built up is placed so far forward that the anterior parts
of the clavicles and interclavicle lie directly below the braincase, the ascending
ramus of the clavicle lying well within, but close up to, the hinder border of the cheek.
The glenoid cavity for the humerus is about 18 cm. behind the quadrate.
The scapula is so small an ossification in the large cartilaginous scapulo-coracoid
that nothing of the glenoid cavity is preserved, though the great thickening of the
bone, below and behind, to more than 7 cm., shows its position. There is a large
supraglenoid foramen passing forward and downward above the glenoid cavity.
The position of the bones in the matrix, evidently still in articulation, gives a measure
of the thickness of the cartilaginous caps to long bones, and to the glenoid cavity of
the scapula ; it seems to be usually about one centimetre.
The humerus may be interpreted as a reduction of that of Eryops, differing in
being much less completely ossified, in having the rugged muscle attachments of that
animal reduced to very small proportions, and in presenting only the merest indica-
tion— by a small out-turning of the surface — of the huge hemispherical knob which
in Eryops gave attachment to the radius. The limb has become a paddle, not a leg
capable of supporting more than half the animal's weight in air. In fact all the
limb bones of this animal greatly resemble those of Trimerorhachis.
FIG. 12. Pamcyclotosaurus davidi. A, the right humerus from below. B, the left
femur from below, c, the left ilium, lateral aspect. All x 1/6.
The radius and ulna, about 8 cm. long, have widened extremities, once cartilage
covered ; and the sum of their widths distally is about 8 cm., the presumed width
of the entirely cartilaginous carpus, which is apparently about 5 cm. long and 8-5
cm. wide, judging from the widths of the heads of the five metacarpals. The middle
metacarpal supports a finger with three phalanges whose total length is just under
8 cm. The first digit has a short, very massive, metacarpal which is followed by two
phalanges, of which the second has a rounded and widened end, and a ligament
attachment well marked on its palmar surface. The metacarpals do not overlap
one another. The phalangeal formula is 2, 2, 3, 3, 2.
Pelvis. The pelvis is remarkable because it is much wider between the iliac
crests than it is between the acetabula. The ischium was evidently narrow, though
all its borders, except the lateral, were extended by cartilage. The pubis is largely
represented by an impression through a layer of skin and is a relatively large sheet
25-s A NEW LABYRINTHODONT FROM NEW SOUTH WALES
containing some ossification, the width across the pair much exceeding that across
the ischia (PI. 30). The wide expansion of the pubis, presenting the greatest possible
contrast to that of Eryops, illustrates the general flattening and widening of the body
found in the Triassic labyrinthodonts. It gives some indication that the tail was
sharply marked off from the body by its much smaller width.
The femur is longer and thicker in the shaft than that of Eryops with which it
was compared. The head bore a large cartilaginous cap ; the lower end, abruptly
truncated but continued in cartilage, seems to have carried two condylar knobs.
The massive shaft has a low crest running along its lower surface, ending proximally
at a ventral process, the homologue of the meeting point of the three great ridges
of the femur of Eryops. This bears a series of well-marked muscle (or ligament)
FIG. 13. Paracyclotosaurus davidi. Restoration of the pelvis, x 1/6. Cartilage is
restored in the dotted areas, and the meeting of the three bones in the acetabulum is
conventional. A, the ischia from in front ; B, the pelvis from above, c, the pelvis
from the left side.
insertions. The tibia, relatively massive, is incompletely known. No trace of the
fibula is preserved. The tarsus is unossified, but there remain a series of rather
scattered metatarsals and some phalanges. These merely show that the foot
greatly resembled the hand in both size and structure.
Skin. The animal's skin is largely preserved as an impression, over part of the
dorsal surface of the body, and of the ventral surface especially in the pelvic region.
Block 14, dorsal surface (PI. 31), lies from 15 to 30 cm. to the left of the mid-line
opposite vertebrae 23-25. It bears an impression of the skin in an area where
it has been rucked up in a manner which suggests that it was flexible, although it
contained closely set, bony scales. These are irregularly oval in plan, ranging
from 3-5 X 5 mm. to 9 X 6 mm. in size, devoid of definite ornament and set quite
irregularly, but not in contact with one another. Their thickness cannot be measured,
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 253
but appears to be less than i mm. They are calcareous, and presumably bony.
An area of 50 square cm. contains more than one hundred scales.
A plastic cast of the ventral surface below the pelvis shows a clearly recognizable
area of skin about 20 cm. square (cf. PI. 30). Posteriorly a strip of skin with a torn
anterior edge stretches across from the upper end of a femur on to a misplaced
ischium. It then spans a gap until it is supported by another flat bone (presumably
the other ischium), and then passes over another gap, where it is shown as an
unsupported sheet some 3 mm. thick that continues over the ventral surface of a
pubis, cracked by some accident with the cracks showing through the skin. It
continues on to the other (misplaced) pubis. It is puckered in a convincing manner
where the torn edge rests on the ischium. There is no suggestion that this ventral
skin contains any bony scales, but it has a delicate surface sculpture of small pits
which are clearly recognizable.
Such preservation of large areas and thicknesses of skin is remarkable, but it is
paralleled by an ichthyosaur skull (No. R.SOQ, D. M. S. W. Coll.) from the plani-
costatus zone between Lyme Regis and Charmouth. Here the skin is about 2 mm.
thick, and in part it retains its cellular structure (Whitear, 1956).
III. BODY SHAPE AND MODE OF LIFE
The mode of life of the labyrinthodont described above should be discoverable,
in part, from its unusually completely known structure. The small distortion
of the head can easily be allowed for, and the complete preservation of skull and jaws
makes its shape certain ; its length is 60 cm. The right hinder corner of the head
is essentially undistorted and shows that the skull at its point of greatest depth
is almost exactly as high as the lower jaw. Similarly the lower jaw symp-
hysis is the same height as the skull above it. Thus, at its point of greatest
height, the head was essentially of oval section, 22 cm. high and 44 cm. wide. The
nostril, orbit and tympanic membrane are all directed upward. Whether the eye
could be projected upward above the head to give a horizontal view, as in the frog,
is impossible to determine, but it may have been so.
The first rib is short, but the second, which is in undisturbed position, implies a
body width of at least 35 cm., 10 cm. behind the condyles. The shoulder girdle is
a rigid structure, the membrane bones lying in, or immediately below, the skin
having a maximum width of 44 cm. at a point a little behind the lower jaws. The
clavicles are upturned so that they incline inward, their minimum width of 23 cm.
lying between the lower jaws, the interclavicle projecting along the throat below
the skull for 15 cm. The maximum width of the body at the shoulder may well
have been 53 cm. The forelegs, in a position of rest, would give a total width between
the middle fingers of the forwardly directed hands of 87 cm. The distance of the
sacral vertebra behind the skull is 115 cm. The body thus has a minimum height
of 16 cm., and a width of more than 20 cm. The tail is about 50 cm. long, and
is probably flattened laterally. The total length is approximately 225 cm.
The weight could be estimated by making a number of assumptions, but has little
meaning in an aquatic animal. Roughly the creature is larger than a man in bulk
254 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
and presumably in weight, but it may well have had no weight when submerged,
depending on the size of the inflated lungs. On land it could no doubt walk to some
extent, making a track little less than a yard wide, with a stride of perhaps 20 cm.
As the individual prints of the well-preserved forefoot would be about 100 square
centimetres, the track would consist of two strips of forefoot prints, with similar
hindfoot tracks just within them, and with a wide streak of; body impressions.
In water it is probable that little but the ends of the fingers and toes would make
impressions, the body streak probably not being seen It seems probable that the
short tail could have had some use in swimming, though inadequate for rapid
movement.
The animal's food may be considered here. It has a feeble dentition ; the only
teeth which could be driven into the body of an animal seized in the mouth are those
on the vomers and palatines, and these stand very little above the maxillary teeth,
a close-set row lacking points of any kind. It is very likely that the creature caught
its food as the living Giant Salamander is said to do — by waiting until the prey came
near, then quickly opening its enormous mouth and swallowing it. The most
probable food would be small fishes, up to some 15 cm. long, which swam in shoals.
Fishes of such dimensions in the form of Promecosomina, a holostean, are those
most abundantly found with the labyrinthodont.
How the animal breathed is uncertain. Had it done so as Megalobatrachus
perhaps does, by moving the floor of the mouth by muscles attached to the hyoid
and branchial arches, some of these elements would probably have been ossified,
and none is. The ribs surround, at least, the upper part of the body, and articulate
with the vertebral column by a wide and essentially two-headed upper end, attached
obliquely to the column. They could presumably have been used in an essentially
mammalian manner for breathing, though the rigidity of the shoulder girdle must
have restricted rib movement there. But as the animal was probably as slow
moving as a Giant Salamander, and the rate of its oxygen consumption very low,
the buccal epithelium may have provided an efficient point of introduction of oxygen
into the body, for the skin of the ventral surface is thick, and the skin as a whole
was perhaps unlikely to be an important respiratory surface. It is difficult to imagine
any such individual moving far, and impossible to imagine it traversing any very
different environment.
IV. SYSTEMATIC POSITION
In 1914, when I first saw this animal as an impression, I referred it to the genus
Cyclotosaurus on the grounds that its skull was in general structure like that of any
capitosaur, and the otic notch closed by contact of the tabular and squamosal laterally.
At that time the only known cyclotosaur was still the type species " Mastodonsaurus
robustus ", given the new generic name Cyclotosaurus by E. Fraas in 1889 ; Smith
Woodward's little English form (C. stantonensis) was at that time referred to
Capitosaurus. Since then new discoveries have increased the number of species of
Cyclotosaurus to ten1, and added a new genus, Rhadalognathus Welles (1947), with a
1 Two other "species" have been referred to this genus without reason.
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 255
\
A
FIG. 14. Paracyclotosaurus davidi. Reconstruction of the skeleton in a walking pose.
Total length approximately 225 cm.
256 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
similarly enclosed tympanic membrane. It is therefore necessary to examine each
of these animals to distinguish which can rightly be referred to the genus.
The type species of Cyclotosaurus, C. robustus (Meyer & Plieninger) from the
Schliffsandstein of the Lower Keuper, was described by Quenstedt (1850) and later
by Fraas (1889). It is a large form with a triangular skull 53 cm. long in the mid-line
with nearly straight sides except for the rounding off of the snout. The width
posteriorly is about 43 cm. and the height at the occiput 5-1 cm. The skull is thus
extremely flattened, though there is no reason to doubt that its real shape is preserved.
Several skulls are known of similar proportions. There are large tusks on the vomers
and palatines. The lower jaw has a maximum depth of 12 cm. at the articulation.
Cyclotosaurus posthumus Fraas (1913), from the Stubensandstein of the Upper
Keuper, has a skull 53 cm. long and 42 cm. in maximum width, almost precisely
the same as that of C. robustus. The height in the mid-line at the occiput is 6-3
cm. again similar to C. robustus, but the face is narrower, its lateral borders are
concave, and the occiput lacks the deep concavity of the border found in C. robustus.
There are large tusks on the vomers and palatines, and the dentition in general
resembles that of C. robustus. It may well be placed in the same genus.
Cyclotosaurus mordax Fraas (1913), also from the Stubensandstein, was founded
on a skull broken off about 2 cm. behind the orbits. The skull is flat and differs
from C. robustus in having its interorbital width proportionally greater and its
suborbital width less than in that form. It is also much shorter in relation to its
width as judged from the palate. It differs from C. posthumus similarly, but may
well be placed in the same genus.
Cyclotosaurus papilio Wepfer (1923), from the top of the Muschelkalk, is based on
a fragmentary skull, clearly " Cyclotosaurus " by the complete enclosure of the
tympanic membrane. The structural details are incomprehensible. It is thus not
certainly determinable.
Cyclotosaurus ebrachensis Kuhn (1932), from the Middle Keuper of Oberfranken,
was described from a complete skull about half the size of C. robustus. The general
outline resembles that of C. robustus, but the orbits and tympanic membranes are
relatively more widely set, no doubt a scale effect. The greatest width is 23 cm.
and the height 3-2 cm., relatively more than in C. robustus and less than in C. mordax.
It is evidently a member of the genus Cyclotosaurus.
Cyclotosaurus hemprichi Kuhn (1942), from the Upper Keuper (PRhaetic) of
Halberstadt, is an admirably preserved but, in part, scattered skull, excellently
described. The skull is approximately 62-5 cm. long, 46 cm. wide, and the height
(dorsal surface to quadrate condyle) is 11-4 cm. The depth of the occiput distin-
guishes this skull from all those listed above, and cannot be explained by growth,
for it is little greater in size than the type species.
Cyclotosaurus stantonensis (A. S. Woodward) (1904), from Stanton, Staffordshire.
Skull length 20-5 cm., width 15-3 cm., maximum height 4-2 cm. The depth of the
occiput as a fraction of skull width agrees with C. hemprichi and differs from that of
the type species. The skull is remarkable because the sutural connection of the
exoccipital and pterygoid is short, whilst the quadrate ramus of the pterygoid has
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 257
a well-developed post-tympanic flange, known in " Capitosaurus "( = Parotosaurus)
but not in any other " Cyclotosaur us " (cf. Sushkin, 1927 : 273, 285).
" Cydotosaurus randalli " Welles (1947), from the Moenkopi formation of Arizona,
is founded on fragments including one showing a " closed otic notch " and part of
a basis cranii. This species differs greatly, in the relations of the exoccipital,
parasphenoid and pterygoid to one another, from any of the forms listed above and
is evidently not closely related to them.
Rhadalognathus boweni Welles (1947) is founded on a very slender labyrinthodont
lower jaw, impossible of comparison with those of capitosaurs. It is held by Welles,
on the basis of associated fragments, to have had a closed otic notch. As recon-
structed by Welles, the skull is unlike that of any of the cyclotosaurs mentioned
above.
" Cydotosaurus spitzbergensis " Wiman (1915) is an indeterminable fragment.
" Labyrinthodon pachygnathus " Owen (1842, pi. 46, figs. 6, 7) was recognized by
Welles as being an English Cydotosaurus.
Comparison of general proportions shows that the skull of the St. Peter's labyrin-
thodont differs from C. robustus (the type species of the genus), C. posthumus,
C. ebrachensis and, presumably, C. mordax, in being much deeper in proportion to
its length and width. In this matter it agrees closely with C. hemprichi and C.
stantonensis.
C. stantonensis differs very greatly in the persistence in it of the post-tympanic
flange of the pterygoid found in " Capitosaurus " ( = Parotosaurus) and in the propor-
tions of the tympanic cavity so far as it can be inferred. It retains more of the
structure and proportions of this region in Parotosaurus, and appears to be a " primi-
tive " form, not necessarily closely related to other cyclotosaurs and I therefore
establish for it a new genus, Procyclotosaurus.
C. hemprichi, however, very much resembles the Australian animal. It differs
in the shape of the tympanic membrane, which is " triangular " in contrast to
circular. But in the structure of the occiput (apart from the presence of a
well-marked shelf on the exoccipital for the supraoccipital cartilage) the two are
very alike ; the relation of the exoccipital to the pterygoid (Kuhn, 1942, pi. i,
fig. ib), the structures shown in his pi. i, fig. 3, and the structure of the quadrate-
pterygoid region (pi. 3, fig. ib) are extremely similar in the two animals.
The resemblances and differences set out above imply that the " Cydotosaur "
condition of total enclosure of the otic membrane by contact of the tabular and
squamosal lateral to it has arisen more than once, and hence cannot by itself charac-
terize a genus.
Thus Cydotosaurus, founded on C. robustus, may include C. posthumus Fraas,
C. mordax Fraas and C. ebrachensis Kuhn.
The species with deep skulls — the Australian skeleton which is the subject of this
paper, and C. hemprichi — differ noticeably from the true cyclotosaurs. That these
differences are significant is shown by the remarkable fact that their peculiarities
occur in three magnificent skulls from the Middle Trias of East Africa in Mr.
Parrington's collection (Field no. 48, Mkongoleko, Stockley's 6.9. " Upper Bone
Bed " (2 specimens) ; Field No. 135, Gingama, Stockley's B.26. " Upper Bone Bed ")
OEOL. 3, 7. 2;5
258 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
which might well be their ancestors, but not the ancestors of the typical flat-headed
Cyclotosaurus. There are accordingly two distinct lines.
Thus the St. Peter's labyrinthodont is made the type of a new genus and species,
Paracyclotosaurus davidi, and with it is placed C. hemprichi of the extreme
Upper Keuper of North Germany.
That the St. Peter's labyrinthodont and Paracyclotosaurus hemprichi belong to
the same genus and one manifestly distinct, by its ancestry, from true Cyclotosaurus
is of stratigraphical significance, for P. hemprichi comes from the " oberen Knollen
Mergel " at Halberstadt, at the extreme top of the Trias, or perhaps even in the
Rhaetic. Nothing in the structure of the St. Peter's animal is inconsistent with
such a late age, but nevertheless it might be somewhat older, that is, earlier in the
Upper Trias.
V. DESCRIPTION OF SUBCY CLOTOS AURU S BROOK V A LENSIS
gen. et sp. nov.
Parrington's East African animals, in so far as the dorsal surface of their skull is
concerned, resemble a small " Parotosaur " skull from the Brookvale clays of the
Hawkesbury Sandstone, shown in Text-fig. 15. This specimen is the mould of a
skull broken so that the right border is lost, and very slightly distorted by pressure
so that the orbits are no longer quite symmetrical. The proportions of the skull
resemble those of most others of Parotosaurus and Cyclotosaurus, the preorbital
length being 66% of the total mid-line length, the extremes amongst parotosaurs
being 61% in the small Cyclotosaurus ebrachensis, and 71% in Parotosaurus helgo-
landicus. The skull is characterized by the small tabular without any trace of a
" horn ", but with a round lappet that approaches the squamosal flange lateral to
the tympanic membrane, failing to meet it by about its own width. The occiput
between the otic notches is proportionately wide, a reflection of the small size of the
skull. The skull is otherwise of normal Parotosaurus structure, but has a small
internasal vacuity between the dorsal processes of the premaxillae. Lateral lines
are often shown as continuous grooves with well-defined borders. A deep groove
on the maxilla begins immediately behind and lateral to the nostril and passes
straight back to the lachrymal, on which bone it turns outward and forward and
ends abruptly. Another groove appears to begin on the maxilla, immediately
lateral to that described above. It passes back just above the insertion of the teeth
for the full length of the bone. The supraorbital groove begins abruptly on the
dorsal surface of the premaxilla, immediately passes on to the nasal, and extends
back on that bone close to its suture with the lachrymal, it then comes on to the
prefrontal, passing on to the frontal where that bone enters the orbital border.
Then as a well-defined groove it surrounds the hinder part of the orbit, turns vertically
on to the jugal, and then backward to cross the point where jugal, quadratojugal
and squamosal meet, continuing over the squamosal to pass back on to the body.
There is a canal, really a series of pits, crossing the supratemporal. For this skull,
whose characters are shown in the figure, I propose the new genus Subcyclotosaurus
and the trivial name brookvalensis.
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 259
FIG. 15. The skull of Subcyclotosaurus brookvalensis gen. et sp. nov. x 3/4. The missing
narrow strip on the right is restored from the left side, and the distortion corrected. The
short tabular processes approaching but not meeting the squamosal are characteristic.
[R. T. Wade Coll., Australian Museum, Sydney].
VI. ASSOCIATED BRACHYOPID
Some time after the discovery of Paracychtosaurus Mr. Dunstan found another
large piece of ironstone in the St. Peter's quarries which retained impressions of
characteristic labyrinthodont bones. A flexible cast from this mould shows part of the
dorsal surface of a skull, sometimes from both surfaces, and the upper surface of
an incompletely preserved left pterygoid. So far as the material goes it is excellently
preserved, but as the skull was partly disarticulated before burial, the bones displaced,
and only the hinder part— not including an orbit — available, it is difficult to determine
its systematic position with any assurance.
Text-fig. 16 represents the bones of the upper surface of the head as they lie, with
the upper surface of the pterygoid. The mesial part of the skull table was clearly
flat, but toward the outer end of the dermosupraoccipital it turns a little downward,
so extending to the end of the tabular. The hinder border is there carried by the
squamosal, continuing in the same direction until it rather suddenly turns vertically
260
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
and descends, presumably to meet a quadratojugal. The well-preserved left
dermosupraoccipital has a thickness of nearly 2 cm. on its hinder surface, which con-
tinues on to the tabular, where it is no longer seen. The tabular, shorter from back
to front than the dermosupraoccipital, is wide, ending in a point laterally, presumably
housed in a small groove in the hinder border of the squamosal, which continues
laterally to it, so that there can be no trace of an otic notch. The supratemporal,
which is cracked across and the two parts separated by about a centimetre, is
completely shown as a relatively large bone surrounded by the dermosupraoccipital,
tabular, squamosal, postorbital and presumably also by the missing postf rental.
The bone is crossed obliquely by an unusually wide and deep lateral line groove,
which ends abruptly before reaching the parietal border of the bone. The post-
orbital, lying lateral to the supratemporal and attached to it by visible suture,
FIG. 1 6. Fragment of associated Brachyopid skull, drawn from a plastic cast from a block-
of ironstone from the Wianamatta Shales of St. Peters, x 1/3.
bears a continuation of the wide lateral line groove, and has long sutures with the
supratemporal and squamosal, and with what is presumably the jugal. The squamosal
is firmly attached by suture to the jugal, postorbital and supratemporal, its lateral
border for the quadratojugal implying an unusual mode of attachment. No
recognizable piece of the orbital margin remains, but immediately in front and on
the left side of the parietal is a wide spread of scarcely ornamented bone, which
toward the middle line is attached to a small area of well-ornamented bone, probably
a frontal : these are obviously misplaced with respect to the rest of the skull.
There can be no doubt that this skull belonged to a Brachyopid, but it is scarcely
determinable generically, and is left unnamed.
VII. ACKNOWLEDGMENTS
The foregoing paper is a long delayed fulfilment of a promise made very many
years ago to Sir T. W. Edgeworth David. The development of the specimen, so
skilfully carried out by the late Mr. F. O. Barlow, required many years of patient
A NEW LABYRINTHODONT FROM NEW SOUTH WALES 261
work using the laborious methods then available, and the intervention of two wars
much increased the delay in finishing the task of preparation.
I can only express my gratitude to these two men, and to the original discoverer
Mr. B. Dunstan. I also owe thanks to Professor Medawar for the hospitality of the
Zoology Department of University College, London, and to the Royal Society for
enabling Miss J. Townend, who is responsible for all drawings which illustrate the
paper, to work with me, and also to Mr. W. Brackenbury for the photographs on
Pis. 29-31.
VIII. REFERENCES
BROILI, F. 1915. Uber Capitosaurus arenaceus Minister. Zbl. Min. Geol. Palaont., Stuttgart,
1915 : 569-575. 2 figs.
1927. Ein Stegocephalenrest aus dem Haupt-muschelkalk von Poppenlauer (Unter-
franken). Zbl. Min. Geol. Palaont., Stuttgart, 1927 (B) : 18-21, i fig.
BROILI F. & SCHROEDER, J. 1937. Beobachtungen an Wirbeltieren der Karrooformation,
27. liber einen Capitosauriden aus der Cynognathus-Zone. S. B. bayer. Akad. Wiss.,
Miinchen, 1937 : 97-117.
CASE, E. C. 1946. A census of determinable genera of the Stegocephalia. Proc. Amer.
phil. Soc., Philadelphia, 35 : 325-420.
EDINGER, T. 1937. Capitosaurus-Gaumen und Unterkiefer aus siiddeutschem Haupt-
Buntsandstein. Senckenbergiana, Frankfurt a.M., 19 : 181-192, 3 figs.
FRAAS, E. 1889. Die Labyrinthodonten der schwabischen Trias. Palaeontographica,
Stuttgart, 36 : 1-158, pis. 1-17.
1913. Neue Labyrinthodonten aus der schwabischen Trias. Palaeontographica, Stuttgart,
60 : 275-294, pis. 16-22.
HAUGHTON, S. H. 1925. Investigations in South African Fossil Reptiles and Amphibia,
13. Descriptive Catalogue of the Amphibia of the Karroo System. Ann. S. afr. Mus.,
Cape Town, 22 : 227-261, 19 figs.
JAEKEL, O. 1922. Neues uber Hemispondyla. Palaont. Z., Berlin, 5 : 1-25, pi. i.
KUHN, O. 1932. Labyrinthodonten und Parasuchier aus dem mittleren Keuper von Ebrach
in Oberfranken. N. Jb. Min. Geol. Palaont., Stuttgart, 69 (B) : 94-144, pis. 3-5.
1942. Uber Cyclotosaurus hemprichi Kuhn und einige weitere Tetrapodenreste aus dem
Keuper von Halberstadt. Beitr. Geol. Thuringen, Jena, 6 : 181-197, P^s- x~5-
MIALL, L. C. 1874. On the remains of Labyrinthodonta from the Keuper Sandstone of
Warwick, preserved in the Warwick Museum. Quart. J. Geol. Soc., London, 30 : 417-435,
pis. 26-28.
QUENSTEDT, F. A. 1850. Die Mastodonsaurier im Grunen Keupersandsteine Wiirttemberg's
sind Batrachier. 34 pp., 4 pis. Tubingen.
ROMER, A. S. 1947. Review of the Labyrinthodontia. Bull. Mus. Comp. Zool. Haw.,
Cambridge, Mass., 99 : 3-368, 48 figs.
ROMER, A. S. & WITTER, R. V. 1941. The Skin of the Rhachitomous Amphibian Eryops.
Amer. J. Sci., New Haven, 239 : 822-824.
SAWIN, H. J. 1945. Amphibians from the Dockum Triassic of Howard County, Texas.
Univ. Texas Publ. 4401 : 361-399, 12 figs.
SUSHKIN, P. P. 1927. On the modifications of the mandibular and hyoid arches and their
relations to the brain-case in the early Tetrapoda. Palaont. Z., Berlin, 8 : 263-321, 39 figs.
SWINTON, W. E. 1927. A new species of Capitosaitrus from the Trias of the Black Forest.
Ann. Mag. Nat. Hist., London, (9) 20 : 177-186, pi. 4.
WADE, R. T. 1935. The Triassic Fishes of Brookvale, New South Wales, xiv + 89 pp.,
10 pis. Brit. Mus. (N.H.), London.
— — 1940. The Triassic Fishes of Gosford, New South Wales. /. Roy. Soc. N.S.W., Sydney,
73 : 206-217, pl- JI-
262 A NEW LABYRINTHODONT FROM NEW SOUTH WALES
1941. Australian Triassic Fishes. /. Roy. Soc. N.S.W., Sydney. 74 : 377-396, pi. 17.
WATSON, D. M. S. 1919. The structure, evolution and origin of the Amphibia. The " Orders "
Rachitomi and Stereospondyli. Philos. Trans., London, 209 (B) : 1-73, pis. i, 2.
WELLES, S. P. 1947. Vertebrates from the Upper Moenkopi Formation of Northern Arizona.
Bull. Geol. Sci. Univ. Calif., 27 : 241-294, pis. 21, 22.
WEPFER, E. 1923. Cyclotosaurus papilio n.sp. aus der Grenzregion Muschelkalk-Lettenkohle
des nordlichen Baden, ein Beitrag zur Kenntnis des Stegocephalen H inter hauptes. Mitt.
bad. geol. Landesanst., Heidelberg, 9 : 367-390, pis. 6, 7.
WHITEAR, M. 1956. On the Colour of an Ichthyosaur. Ann. Mag. Nat. Hist., Londo'1
(12) 9 : 742-744, pi. 23.
WILLS, L. J. 1916. The structure of the lower jaw of Triassic labyrinthodonts. Proc.
Bgham Nat. Hist Soc., 14 : 1-16, pis. i, 2.
WOODWARD, A. S. 1890. The fossil fishes of the Hawkesbury Series at Gosford. Mem.
Geol. Surv. N.S.W., Sydney (Palaeontology), 4 : 1-55, pis. i-io.
1904. On two new labyrinthodont skulls of the genera Capitosaurus and Aphaneramma.
Proc. Zool. Soc. Lond., 1904 : 170-176, pis. n, 12.
1908. The fossil fishes of the Hawkesbury Series at St. Peter's. Mem. Geol. Surv. N.S.W.,
Sydney (Palaeontology), 10 : 1-29, pis. 1-4.
A NEW LABYRINTHODONT FROM NEW SOUTH WALES
VIII. KEY TO ABBREVIATIONS IN TEXT-FIGURES
Ang., angular.
Ant. Zyg., anterior zygapophysis.
Art., articular.
B. Pt., space formerly occupied by the
cartilaginous basipterygoid process.
Bd. Ves., foramen for a blood vessel.
Clav., clavicle.
Clei., cleithrum.
Cleido-mast. Mus., cleido-mnstoid muscle
insertion.
D. S. Oc., dennosupraoccipital.
Den., dentary.
E. Pt., epiterygoid.
EC. Pt., ectopterygoid.
Ept. Bas. Art., facet on epiterygoid which
articulates with the basis cranii.
Ex. Oc., exoccipital.
Ex. Oc. Cav., cavity in the exoccipital.
Ext. Nos., external nostril.
Fern., femur.
Fr., frontal.
Gass. Gang., groove in the epipterygoid for
the Gasserian ganglion.
7. Cen., intercentrum.
/. Clav., interclavicle.
//., ilium.
Int. Nos., internal nostril.
Isc., ischium.
Ju., jugal.
L. L., lateral line groove.
Lac., lachrymal.
MX., maxilla.
N. Sp., neural spine.
Na., nasal.
Noto., space for notochord.
Orb., orbit.
P. Cen., pleurocentrum.
P. MX., premaxilla.
P.O., postorbital.
Pal., palatine.
Par., parietal.
Par. Sp., parasphenoid.
Pin. For., pineal foramen.
Pr. Fr., prefrontal.
Pr. Ot., prootic.
Pt., pterygoid.
Pt. Fr., postfrontal.
Pu., pubis.
Qu., quadrate.
Qu. J ., quadratojugal.
S. Rib., sacral rib.
S. Tern., supratemporal.
Scap., scapula.
Sp. Ch., spinal chord.
Sph. Eth., sphenethmoid.
Spl., splenial.
Sq., squamosal.
St., stapes.
Sur. Ang., surangular.
Tab., tabular.
Tran. Proc., transverse process.
Tym. Cav., tympanic cavity.
Tym. Mem., tympanic membrane.
Vo., vomer.
//, notch in sphenethmoid probably for
optic nerve.
V1, ophthalmic branch of N. trigeminus.
V2 & 3, maxillary and mandibular branches
of N. trigeminus.
XII, foramen for nerve XII.
PLATE 27
Pavacyclotosauvus davidi
Photographs taken before 1914 by Mr. B. Dunstan showing how the many blocks of ironstone
in which the 9-foot skeleton is preserved were fitted together by him, so that one group, A,
has the impression of the upper surface of the whole animal, whilst B shows its lower surface.
The loose blocks to the right fit between the two main groups, showing vertebrae and ribs.
Bull. B.M. (N.H.) Ccol. 3, 7
PLATE 27
Paracyclotoscnirus davidi .
PLATE 28
Paracyclotosaurus davidi
Upper : Left lateral view of the mounted skeleton, approximately 9 feet (2-75 m.) long.
Lower : Three-quarter front view of the left side of the mounted skeleton to show the open
mouth and the character of the teeth. The animal could certainly have opened its mouth
even more widely.
Bull. B.M. (N.H.)Geol. 3, 7
PLATE 28
Paracyclotosaurus davidi.
PLATE 29
Paracyclotosaurus davidi
Photograph of a cast showing vertebrae 6-10 and the left ribs belonging to them, together with
the distal part of ribs 4 and 5 and the upper ends of the cleithrum and scapula.
The cast was made by Mr. F. O. Barlow to show how the skeleton looked before the individual
bones were disarticulated, x ^ approx.
PLATE 30
Paracyclotosaurus davidi
Plastic cast from a block showing (top, left and right) the ventral surface of the two pubes,
and (bottom, left and centre) the two ischia somewhat displaced.
Skin covers the whole structure except for a rent across the middle, where (centre) a surface
of ischium can be seen. A loose flap of skin crosses the left side of the tear. On the right
side the head of the left femur can be seen displaced posteriorly. x i approx.
Bull. EM. (N.H.) Geol. 3, 7
PLATE 30
Paracyclotosaurus davidi.
PLATE 31
Paracyclotosaurus davidi
Block No. 14 with skin of the dorsal surface with its included bony scales, from the region
25 cm. left of the mid-line a little in front of the sacrum. x \.
l3utt. 13.M. (N.H.) Gcot. 3, 7
PLATE 3:
Paracyclotosauras davidi.
AN EARLY PLEISTOCENE
MAMMALIAN FAUNA
FROM BETHLEHEM
D. A. HOOIJER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 8
LONDON: 1958
GEOLOGY
The following papers appeared in Volume I (1949-52) :
Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.
H. D. Thomas & A. G. Davis . . . 75. 6ti.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu 55.
No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White js. 6d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
Northern Iraq. L. F. Spath . . . . . . . ics.
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum T. H. Withers 55.
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae).
T. H. Withers . . 55.
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian
ofPodolia. W. N. Croft IDS.
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsdale 105.
No. 9 (1952). Australian Arthrodires. E. I. White .... 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. WT. F. Whittard 6s.
The following papers appeared in Volume II (1953-56) :
No. i. (1953). The Deer of the Weybourn Crag and Forest Bed of
Norfolk. A. Azzaroli . . . . . . . . £z $s.
No. 2 (1953). A Coniferous Petrified Forest in Patagonia. M. G. Calder 125.
No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark 35. 6d.
No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.
M. E. J. Chandler 16.9.
No. 5 (1954)- The Carboniferous Flora of Peru. W. J. Jongmans . . 155.
No. 6 (1955). Further Contributions to the Solution of the Piltdown
Problem. J. S. Weiner, W. E. Le Gros Clark & K. P. Oakley el al. /i
No. 7 (1955). The Schizaeaceae of the South of England in early Tertiary
Times. M. E. J. Chandler 15.-.-.
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . . /i
AN EARLY PLEISTOCENE MAMMALIAN
FAUNA FROM BETHLEHEM
BY
D. A. HOOIJER
(Rijksmuseum van Natuurlijke Historic, Leiden)
Pp. 265-292 ; Pis. 32-35
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 8
LONDON : 1958
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred Pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 8 of the Geological
series.
Trustees of the British Museum, 1958
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued August, 1958 Price Ten Shillings
AN EARLY PLEISTOCENE MAMMALIAN
FAUNA FROM BETHLEHEM
By D. A. HOOIJER
SYNOPSIS
The fossil mammalian fauna from Bethlehem contains characteristically Villafranchian forms
such as Archidiskodon cf. planifrons and Leptobos. It further comprises at least five species
known from Villafranchian sites in Eurasia. As at Villarroya (Spain) and at Tatrot (India),
which are likewise Villafranchian assemblages, there is Hipparion but no Equus. However,
the Bethlehem fauna differs from those of the Eurasian Villafranchian in the presence of Giraffa
cf. camelopardalis and in the absence of Cervidae ; this puts an African " stamp " upon the
Bethlehem fauna, as may have been expected a priori from its geographical position.
THE fossil mammalian remains described in the present paper were found at the
highest point of Bethlehem, Israel, 790 m. above sea-level. The first bones were
accidentally found by the owner of a garden digging for water ; in 1934 Miss D. M.
A. Bate drew attention to this discovery (Bate, 1934). In 1935 and 1936 under the
auspices of the Department of Antiquities, Palestine, the Wellcome Archaeological
Research Expedition to the Near East, with Miss Bate as palaeontologist and Miss
E. W. Gardner as geologist, undertook the excavation of the bone-bearing beds of
Bethlehem. Work was continued in 1937 through the support of the trustees of Sir
Henry Wellcome and Sir Robert Mond. Finally, in 1940 the excavation was completed
by Dr. M. Stekelis of the Hebrew University, Jerusalem. The geology and archae-
ology of the site was described by Miss Gardner, and Miss Bate contributed a discussion
of the fossil vertebrate fauna (Gardner & Bate, 1937). In this paper Miss Bate,
after a preliminary examination of the fossils, concluded that the Bethlehem fauna
" is not later than Early Pleistocene, using this term palaeontologically as indicating
the time of arrival of true Bos, Elephas, and Equus . . . Further, it is claimed for
this fauna that it will provide a faunistic link for this period between Asia and East
Africa . . . ". The following mammals were listed :
Felis sp. (size of Panthera leo) Hipparion sp.
Hippopotamus sp. Rhinoceros cf. etruscus
Bos sp. Stegodon sp.
Antelope. Elephas sp.
Giraffoid. Small carnivore.
In a subsequent note, dealing with the results of the 1940 excavation reported
upon by Dr. Stekelis, Miss Bate added " Equus sp. (? Hipparion) " (Bate, 1941).
GEOL. 3, 8 26
268 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
The fossil collections obtained at Bethlehem were sent to the British Museum
(Natural History) for study and description by Miss Bate, whose death in 1951
prevented completion of this phase of her Palestinian work, to which she had
devoted so much of her time and energy.
In the summer of 1956 I had the opportunity of examining the Bethlehem collec-
tion at the British Museum (Natural History), and Dr. E. I. White, Keeper of the
Department of Palaeontology, most kindly consented to send the material selected
for further examination on loan to me in Leiden. I am much indebted to Dr. White
for facilitating my study of this interesting fauna. To Dr. A. T. Hopwood I owe
valuable information and kind advice. Mr. A. J. Sutcliffe was very helpful in arrang-
ing the shipment of the collection. The photographs were taken by Mr. H. F. Roman
of the Rijksmuseum van Natuurlijke Historic at Leiden, with the exception of those
of PI. 32, which were made at the British Museum (Natural History). All this is
here gratefully acknowledged.
The results of my study are given in the present paper. The material is preserved
in the Department of Palaeontology of the British Museum (Natural History) ;
registration numbers are given in each case. Unless otherwise stated, the measure-
ments recorded in the present work are in mm.
CARNIVORA
CANIDAE
Nyctereutes megamastoides (Pomel)
(PI. 34, ng. 7)
In her second note on the Bethlehem fossils Miss Bate (Gardner & Bate, 1937)
mentioned a small carnivore, " but its remains are extremely fragile and have not
yet been extracted from the matrix ". When I received the collection the small
carnivore remains had been treated with preservative, and removed from the matrix.
It is evident that both the teeth and the bones were broken before removal from the
deposit ; a number of teeth and some vertebrae are partially embedded in plaster,
apparently because otherwise they could not be preserved.
Although the carnivore remains are in a bad state of preservation their specific
identity is certain : they belong to Nyctereutes megamastoides (Pomel), a species
characterized by its small teeth ; the large size of M1 relative to P4 ; the slightly
reduced M2 ; the high and laterally compressed lower premolars with a concave
posterior slope of the protoconid which bears an accessory cusp only in P4 ; the
elongated M2 ; and the oval-shaped M3. The mandibular ramus is quite slender,
and, above all, characterized by the peculiar development of the subangular process
or lobe ; this character cannot be observed in the Bethlehem material. There are
two portions of the left mandibular ramus (M 18521), one with P3_4, and the other
with M2_3, and also parts of the coronoid and the condyloid processes, apparently
of the same individual. The upper dentition is represented by a left canine (M 18516),
P3-4 sin. and M1"2 sin. (much worn down) (M 18512), and a P4 dext. (M 18521).
Post-cranial remains (M 18521) comprise several cervical vertebrae including a
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 269
partial epistropheus, proximal and distal portions of a right humerus, the proximal
ends of a left radius and ulna, the head of a right femur, a left astragalus and calcaneum,
and a great many fragments of the shafts of the long bones, of metapodials, foot
bones, and phalanges. In Table I the measurements of the Bethlehem teeth are
compared with those of Nyctereutes megamastoides from Perrier (Boule, 1889) and
from Villarroya (Villalta, 1952 : 42). There is no significant difference between the
Bethlehem teeth and those of the Villafranchian of Europe.
TABLE I. — Measurements of Nyctereutes megamastoides
Bethlehem Perrier Villarroya
P3, length 8-5 . 8
width ..... 3- i . 3
P4, length ..... 12 -7 . 13
width ..... 5 -6 . 6
M1, length . . . . . 10-2 . 10-5
width . . . . c. 12 . H'5
M2, length ..... 7-1 . 7
width ..... .9
P3, length .... 7-8 . 7-3 7-8
width . . . . . 3-0 . 2 • 5 . 2-3
P4, length 9-6 . 9 7-9*5
width 4-0 3 3-3-5
M2, length . . . . . 8-2 . 8-5 . 7-9'5
width ..... 5-6 . 5 . 5-6
M3, length 4-4 . 4-5
width . . . . . 3-8 . 3-5
Height of ramus between P2 and P4 . 12-5 . 13
Maximum width .... 6-5 . 6
Besides Perrier (Les Etouaires and Roccaneyra : Bout & Azzaroli, 1952) and Vil-
larroya, other Villafranchian localities that yield Nyctereutes megamastoides are the
Val d'Arno, Seneze, and Saint Vallier (Viret, 1954). The remains described as Canis
(Nyctereutes} sinensis Schlosser from the Villafranchian of Nihowan, China (Teilhard
de Chardin & Piveteau, 1930 : 88, pi. 17, figs. 1-3, pi. 18, figs. 2-3), and from the
Middle Pleistocene of Choukoutien (Pei, 1934 : 23, pi. 3, figs, i, 2, 4-6, 8, 10 ; pi.
4, figs. 1-3) appear to be at most subspecifically distinct from Nyctereutes mega-
mastoides. As related by Pei (1934 : 31), the Chinese Nyctereutes has been traced
back to the Middle Pliocene ; the Villafranchian remains are, on the average, a little
larger than those from the Middle Pleistocene, and the Upper Pleistocene remains
(Pei, 1940 : 15) as well as the protohistoric remains of Anyang (Teilhard de Chardin
& Young, 1936 : 7) are as small as, and have been referred to the Recent Nyctereutes
procyonoides (Gray), the " raccoon-dog " of Eastern Siberia, Japan, China, and
Northern Indo-China. In view of this fossil record it is probable that Nyctereutes
megamastoides arose in China, and should be considered a Villafranchian immigrant
to Europe. The remains in the Bethlehem collection indicate that Nyctereutes
megamastoides, in its east-west migration at the beginning of the Pleistocene, spread
also to the Eastern Mediterranean region. There are no records of the species from
Africa,
270 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
FELIDAE
Homotherium (?) sp.
(PL 34, figs, i, 2)
A large machairodontine is represented in the Bethlehem collection by a left
Mx (M 18511), incomplete in front as well as behind. The top of the paraconid is
missing. The powerful anterior root (below the paraconid) is for the most part
preserved ; the posterior root is broken off. Although the development of the
metaconid and of the talonid cannot be studied in the present incomplete specimen,
the wide concave area between the paraconid and the protoconid blades internally,
and the mode of wear of the crown are machairodontine. The greatest length of
the crown must have been over 30 mm.; the greatest width is 14-1 mm. These figures
suggest a form similar in size to Homotherium crenatidens Fabrini from Perrier
(length of MI} 32 mm.: Teilhard de Chardin & Piveteau, 1930 : 116, footnote), a
Villafranchian species also known from Seneze, the Val d'Arno, Chagny, Villarroya,
and Saint Vallier (Schaub, 1944 ; Viret, 1954, table opposite p. 184). Kretzoi
(1954 : 250), whose Epimachairodus is a synonym of Homotherium Fabrini (Simpson,
1945 : 121 ; Arambourg, 1947 : 438 ; Viret, 1954 : 78), records remains probably
referable to H. crenatidens from the Villafranchian site of Kislang in Hungary.
According to Schaub (1934), part of the material described as Machairodus
nihowanensis by Teilhard de Chardin & Piveteau (1930, pi. 22, fig. 2) does not seem
to be specifically distinct from H. crenatidens from the Villafranchian of Europe. This
Early Pleistocene form from Nihowan is succeeded in the Middle Pleistocene by
Homotherium ultimum (Teilhard de Chardin) from Choukoutien, the lower carnassial
of which measures 32 by 12 mm. (Teilhard de Chardin & Pei, 1941 : 40).
Homotherium (?) sp. from Bethlehem is too incompletely known to allow of a definite
identification. Similar large machairodontines are recorded from the Villafranchian
of East and South Africa (Arambourg, 1947 : 438 ; Ewer, 1955 : 612). Whether the
affinities of the Bethlehem form lie with the Eurasian or with the African species
cannot be decided until better specimens are available.
PROBOSCIDEA
ELEPHANTIDAE
Archidiskodon cf. planifrons (Falconer & Cautley)
(PI 32 ; PI. 34 ; figs. 3, 4 ; PI. 35, figs, i, 2)
A mandible without the ascending rami, collected by Dr. M. Stekelis in 1940
(M 18582, PI. 32, figs, i, 2), is one of the most diagnostic specimens found. Like
several of the others, this specimen is crushed, indicating the pressure to which the
bone-bearing beds were formerly subjected. The right horizontal ramus has been
pressed on from above and laterally, but the left ramus is only slightly distorted.
The last molars are in situ on both sides ; the M3 dext. is fractured between the 4th
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 271
and 5th plates from the front, and the two portions are displaced along the fracture,
but M3 sin. is undamaged. What remains of the ascending rami shows that their
anterior borders are on a level with the 6th plates of M3.
The symphysis is obliquely prolonged downward, forming a projection or " beak "
just as in the mandible of Archidiskodon planifrons from the Siwalik Hills figured
by Falconer & Cautley (1845, pi. 8, fig. 20). This is a very characteristic feature of
the species in question ; it is also present in the mandibles from Seneze and from
Chagny (Mayet & Roman, 1923 : 81, fig. 13). In the Bethlehem mandible as well as
in the Siwalik specimen, which I have studied in the British Museum (Natural
History) (regd. 36736), the beak is not complete ; in the French specimens the
length of the beak is about equal to the height of the ramus below M3. The portion
preserved in the Bethlehem mandible is about 7 cm. long ; it has a height of 7-8
cm., and is about 4 cm. wide, just as in the Siwalik specimen referred to above.
Because of the presence of this anterior downward projection of the symphysis the
lower border of the ramus is concavo-convex from before backward, the convexity
beginning approximately at the level of the anterior border of M3. Presumably
part of the M2 was still in place at the time of the animal's death, but is now lost.
The height of the ramus below the middle of M3 is 15 cm., equal to the greatest width,
measurements that correspond well with those of the mandible of A. planifrons
figured by Falconer & Cautley (1845, pi. 8, fig. 2) as well as with those of an unfigured
mandible of the same species in the British Museum (M 3090). There are two mental
foramina on each side, a large one at the middle of the height of the outer surface of
the ramus, on a level with the posterior border of the symphysis, and a smaller one
about 5 cm. below and anterior to the former. In the unfigured mandible of A.
planifrons from the Siwaliks the number and position of the foramina is the same ;
in the figured mandible the larger of the mental foramina is duplicated on the right
side but single on the left, and the distance between this and the smaller foramen
is 6-5 cm. On either side of the depressed anterior surface of the beak a ridge runs
upward to the alveolar edge ; the upper portion of this ridge is badly damaged
in the Bethlehem mandible, but its lower part forms an angle of about 120° with the
occlusal surface of the molar as in the two Siwalik specimens.
So far as the state of preservation of the Bethlehem mandible permits one to form
an opinion, there is nothing to distinguish it from the mandible of Archidiskodon
planifrons from the Upper Siwaliks.
The same is true of the molar, M3. In the Bethlehem mandible M3 is worn to and
including the 6th plate from the front, and there are ten plates in all, besides the
talonids. The greatest length of the crown, in a straight line from the middle of the
anterior border to the heel, is 290 mm.; the greatest width, at the 2nd plate from
the front, is 93 mm., cement included, and 84 mm. exclusive of cement. The height
of the foremost unworn plate, plate 7, is 94 mm.; the slightly worn 6th plate has the
same height and, therefore, must have been somewhat higher when unworn. In
height it cannot have exceeded/ 100 mm., however, for the dentine cores of the cone-
lets do not show yet, and the enamel thickness of the plates is about 6 mm. The
cement coat is heavy ; even the sides of the plates are completely covered.
The enamel figure of the anterior talonid is confluent with that of the first plate ;
only a labial enamel fold remains to indicate the boundary between the two. Plate i
GEOL. 3, 8. 26 §
272 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
has a postero-median enamel projection that makes contact with a similar projection
on the anterior surface of plate 2. Plates 2 and 3 are worn to single enamel figures,
with median anterior and posterior expansions blocking up the valleys in between,
which are open on either side, the antero-posterior width of the valleys being equal
to that of the plates. The enamel is 6 mm. thick, and is slightly crimped.
Passing backward along the crown, the enamel expansions of the plate figures
become less marked. In plate 3 the enamel border forms an almost entire loop, just
to the labial side of the median line of the crown, with a diameter of 13 mm. and
projecting 8 mm. posteriorly. The posterior expansion of plate 4 projects only 5 mm.
beyond the surface. The width of the enamel figure of plate 4 is only 70 mm., whereas
the greatest width of the same plate, including cement, is the same as that of plate 2,
93 mm. The enamel figure of plate 4 is constricted on either side of the median
expansion, forming a transversely elongated enamel figure on either side of the
central, antero-posteriorly extended figure. Of the lateral figures that on the lingual
side is longer. In the right M3 plate 4 is slightly less worn than that in the left, and
the three enamel figures are distinctly separate.
In plate 5 the grooves between the conelets are still visible. There appear to be
four conelets, the second from the labial side being more extended antero-posteriorly
than either of the others. It is this conelet that forms the central expansion of the
more worn plates in front. In plate 5 the two central conelets together measure
26 mm. transversely, the total width of the four enamel conelets being 63 mm.
The greatest (basal) width of this plate, cement included, is again 93 mm. Plate 6
has the four conelets just touched by wear, the two in the centre occupying a width
of 26 mm., as in plate 5. In plate 7 the conelets are still covered up with cement ;
evidently this plate had not erupted at the time of death.
From plate 7 backward the plates begin to diminish in width as well as in height ;
the basal widths cannot be measured as the base of the crown is not fully exposed,
but the heights decrease from 88 mm. in plate 8 to 80 mm. in plate 9, then to 70
mm. in plate 10, the terminal full plate. The talonid is barely 50 mm. high with a
width of about 35 mm., and is completely covered with cement. Because of the
outward curvature of the hinder end of the molar the talonid is rather obliquely
placed : it is in the same antero-posterior line as the labial ends of the foremost
plate-figures.
The laminar frequency of the molar, that is, the number of plates per 10 cm.
of antero-posterior length, varies somewhat with the place in which it is taken. The
laminar frequency is just under 4 in the middle of the occlusal surface, the distance
from the middle of the valley between plates i and 2 to the middle of the valley
between plates 5 and 6 (covering four plates and four cement intervals) being 106 mm.
When taken at the base lingually, which is exposed from the 4th plate backward in
the left M3, the laminar frequency is distinctly lower, not only because of the curva-
ture of the long axis of the crown with the convexity inward, but also because of the
rootward divergence of the plates, characteristic of lower molars. The laminar
frequency is only 3 at the base lingually ; at the labial alveolar margin it is 4.
The low plate formula (x 10 x), the low laminar frequency (3-4), the height of
the crown that hardly exceeds the greatest width (height-width index (plate height-
width indices cannot be given) just over 100) the presence of median looped expan-
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 273
sions of the enamel figures of the moderately worn plates, the great thickness of
the enamel (6 mm.), and the abundance of cement, are characters that leave no doubt
as to the specific identity of the Bethlehem elephant : it belongs to the most primitive
archidiskodont stage exemplified by Archidiskodon planifrons of the Upper Siwaliks
of India. In the characters of the mandible, notably in the downturned anterior
end of the symphysis, the present Bethlehem specimen agrees perfectly with those
from the Siwaliks and from certain Villafranchian sites in Europe.
A number of isolated molars and molar fragments in the Bethlehem collection
are described below.
A right upper second molar collected in 1940 (M 18523, PI. 34, figs. 3, 4) is only
slightly damaged. The molar carries nine plates, five of which are worn, as well as
the damaged anterior and posterior talons. The roots are preserved for the most
part. The crown as a whole is slightly curved : the lingual surface is concave from
before backward, and the labial surface is convex antero-posteriorly. The base of
the crown is concave antero-posteriorly towards the roots. The total length of the
crown is 262 mm.
The enamel figures of the anterior worn plates show the characteristic median
expansions (just to the lingual side of the median line), the posterior loop of the 2nd
and the anterior loop of the 3rd plates almost make a contact across the valley, which
is filled with cement. The enamel thickness is 5-6 mm. Plate 5 has four conelets,
the second from the lingual side being the largest. The remaining plates are fully
embedded in cement, as well as the incomplete and very small terminal plate, the
talon. Measurements are given in Table II.
TABLE II. — Measurements of Mz dext. of A. cf. planifrons (M 18523)
No. of plate
Width
Height
As shown by the height-width indices of the unworn plates, the plates are either
lower or slightly higher than wide. The laminar frequency of the present specimen
varies from 3 (in the middle of the worn surface) to 3! (at the base lingually). There
is one powerful anterior root supporting the first two plates, a very large single
root supporting the posterior five plates, and three intermediate roots, two labial
and one lingual. The apical portions of these roots are broken off ; the portions
preserved are as long as the height of the (unworn) crown.
In the Upper Siwalik A. planifrons the number of plates in M2 varies from eight
to nine. The length of the crown of the Bethlehem M2 (262 mm.) is greater than that
of any M2 of A. planifrons from the Siwalik Hills as recorded by Falconer and by
Osborn (see Osborn, 1942 : 949, 954 : 191-221 mm.), but this is not a matter of great
moment, for an unquestionable M2 of A. planifrons from the Punjab (Hooijer, 1955 :
99-101) is 261 mm. long against 178-204 mm. in Falconer's and Osborn's series of
r
i
2
3
4
5
6
7
8
9
Talon
—
IO2
103
IOO
IOO
IOO
97
83
55
. c. 30
—
—
—
—
84
81
74
c. 58
—
—
—
—
—
84
84
89
c. 105
—
274 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
Siwalik M2, which tends to show that upper second molars of the size of the Bethlehem
specimen occur in the Upper Siwalik A. planifrons also.
A left upper second molar (M 18524) is unworn (PI. 35, figs, i, 2). There are eight
plates, plus the talons. Much of the enamel along the base of the crown is gone :
the lingual edges of plates i, 2, 7, and 8, and the external conelets and edges of plates
3 to 7, inclusive, are missing. However, it is mostly only the enamel coat that is
lost, and the enamel thickness, wherever exposed in the present specimen, is 6 mm.
Therefore, it is possible to give the basal widths of most of the plates by taking the
actual width, and adding to it 6 mm. for the missing enamel layer. Cement is heavily
developed ; it covers up the conelets and the edges of all but the hindmost three
plates. The conelets of plates 4 and 5 are destroyed, and the full height of these
plates cannot be measured.
TABLE III. — Measurements of M2 sin. of A. cf. planifrons (M 18524)
No. of plate
12345678 Talon
Width 93 100 105 — 105 103 95 71 . 50
Height . 69 74 70 70 65 54 . 33
Height-width index . 74 74 67 68 68 76 . 66
The number of conelets in the anterior plates is difficult to see because of the
cement cover. There is a slight dislocation in the median line of plates 1-3, the labial
portions being placed slightly more forward than those on the lingual side. Plate
6 bears five conelets, the two on the labial side are broken off. In plate 7 the
number of conelets is likewise five, but in plate 8 there are four conelets only. The
talon consists of three cones, and a very small accessory cone on the lingual side.
The length of this M2 sin. is 242 mm., less than that of the M2 dext. recorded
above. The number of plates is also less (eight instead of nine), but the laminar
frequency is the same in both specimens, viz., 3 to 3^. The characteristic median
expansions do not show because all the plates are unworn. It will be seen from Table
III that the full height of the plates is only two-thirds to three-fourths the basal
width (exclusive of cement), which indicates that the M2 sin. is lower-crowned than
the M2 dext.
The anterior root is divided in the middle, and is preserved for a few centimetres
only ; the main root is broken off almost entirely, and the junction between the two
is on a level with that between the 3rd and 4th plates of the crown.
Another specimen with unworn plates is the hinder end of an M3 dext. (M 18527).
It is broken off anteriorly through the 7th plate from behind, and is much corroded.
The enamel is lost along the lingual edges of the plates, exposing the dentine cores.
A large portion of the labial surface is covered with plaster. The enamel of the
conelets of the plates is broken. Of the roots nothing is preserved. However, allowing
for an enamel thickness of 5 mm. it is possible to estimate the widths at base of all
the plates, and the height measurements of all but the last two plates and the talon,
which are so much damaged that their height cannot be measured. The laminar
frequency is 4.
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 275
TABLE IV. — Measurements of Mz dext. of A. cf. planifrons (M 18527)
No. of plate from behind
VII VI V IV III II I Talon
Width . . . 107 95 88 84 75 70 55 . c. 25
Height ... 88 80 75 69 65 —
Height- width index 82 84 85 82 87 —
Table IV shows that the unworn plates are less high than wide ; this is also true
of the unworn M2 sin. from Bethlehem.
A portion of a molar consisting of seven fragments (M 18561) contains parts of
two unworn, and two worn plates. The width and height of one of the unworn plates
(the enamel thickness is 6 mm.) can be determined as 95 mm., and 85 mm., respective-
ly, giving a height-width index of 89, the maximum figure for this index found in
the molars described above. The present specimen is broken through the median
line of the crown, and clearly shows the V-shaped valleys between the plates, open
down to the bottom as is characteristic of archidiskodonts. The laminar frequency
of the present fragment is 3^. Its serial position cannot be determined with certainty,
but because of the relatively great basal width and low laminar frequency it presum-
ably formed part of a penultimate or a last molar.
A fragment of a molar, probably of the same specimen as the last (M 18525),
comprises part of an unworn plate 84 mm. high but of unknown basal width. There
are at least five conelets to this plate, the grooves between which remain distinct
rootward to over one-half the height of the crown.
Two last lower molars, one right and one left, evidently of the same individual
(M 18528 and M 18529) are very much worn down. Their anterior portions are missing ;
the left M3 still comprises seven plates, the right, five only . The anterior plates have
their dentine surfaces coalesced as the valleys between them are partially or entirely
worn out. The laminar frequency of the occlusal surface is 3|. The enamel is very
thick : 5-6 mm. The plate figures of the last four plates are irregularly expanded
in the centre ; their lingual portions are placed more forward than the labial, and the
enamel bands make contact in the median line of the crown. The greatest width of
the crown (at the 5th plate from behind) is c. 100 mm., exclusive of cement which is
well developed all round the crown. At the 2nd plate from behind the width is 85
mm., and at the talonid, 40 mm. There are accessory enamel cusps labially of the
talonid and of the last plate as well as at the lingual entrances to the valleys. Height
measurements cannot be given as even the talonids are worn. The main posterior
root supports at least six plates and is of great length, the depth of the root below the
talonid is 15 cm. at least.
There remain a number of molars, most of them very much worn down, that are
either too incomplete or too much damaged for anything of value to be deduced
from them. It is evident, however, that they belong to the same primitive archi-
diskodont elephant as the better-preserved specimens above described ; their
enamel is very thick, the laminar frequency is low, the valleys are V-shaped, and the
roots are very long. Two much worn molars, evidently of the lower jaw as their
276 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
occlusal surfaces are concave from before backward (M 18557, M 18560) display
these characters very clearly. The laminar frequency is approximately 4 ; the long
roots are distinctly recurved backward. M 18557 seems to have had seven plates
only, and presumably represents Mx sin. The roots indicate that the length of the
crown probably did not exceed 165 mm. Width measurements cannot be given.
The median expansions of the enamel figures show well in a posterior fragment of
another lower molar (M 18554) tne root of which is 13 cm. long as preserved. A much
worn crown fragment without the roots (M 18538) holds four plates in 10 cm. of
length ; the same laminar frequency obtains in a fragment of an upper molar
(M 18526) that is interesting because it has had small stones pressed down on the
plates with such force that these have been partially broken, and the crown surfaces
deflected from their normal position. The following specimens are too fragmentary
for their position to be determined : nos. M 18530, M 18531, M 18539, M 18540,
M 18541, M 18543, M 18544, M 18551, M 18559.
As mentioned above the most complete and first described specimen of the Bethle-
hem elephant, viz., the mandible obtained during the 1940 season (M 18582) is
indistinguishable from Archidiskodon planifrons from the Upper Siwaliks of India.
However, in the isolated and (partially or entirely) unworn molars we observe a
character in which it appears to differ from the Siwalik A. planifrons.
All the relevant data on the molars of the Upper Siwalik A . planifrons described
and figured by Falconer and by Osborn are contained in two tables by Osborn
(1942 : 949, 954). Upper Siwalik specimens of A. planifrons collected by Eug.
Dubois in the Punjab, and now in the Leiden Museum, including a fine skull closely
resembling the British Museum specimen (Falconer & Cautley, 1845, pis. 9, 10),
have been described by the author (Hooijer, 1955 : 96-100). The material of A.
planifrons obtained by the Yale North India Expedition of 1932 (Hooijer, 1955 :
100-102 ; 1956) includes the geologically oldest specimen yet obtained, viz., a portion
of M3 collected near the base of the Tatrot zone first recorded by Lewis (1937 : 198),
basal Upper Siwaliks and basal Pleistocene. This specimen (Hooijer, 1956) cannot
be distinguished from other M3 of A . planifrons (the bulk of the material is presum-
ably from the Pin j or zone overlying the Tatrot zone), and the plates are higher than
wide (height-width indices 110-124). The M3 in situ in the British Museum skull
of A. planifrons has a height-width index of 115. An M3 from the Punjab in the
Leiden Museum (Hooijer, 1955 : 102, 103) has five unworn plates very nearly as
high as wide, the height-width indices varying from 97 to 105.
As can be seen from the tables by Osborn (1942 : 949, 954) the height of the molars
of A . planifrons as given by Falconer and by Osborn is either greater or much less
than the width of the same specimens. Fortunately, most of the specimens have been
figured, and from the figures it is evident that where the height exceeds the width
the crown has unworn plates, e.g., M3 : Falconer & Cautley, 1845, pi. 10 (width
89 mm., height 102 mm., index 115) ; M1 : Falconer & Cautley, 1845, pi. 6, fig. 5
(width 69 mm., height 77 mm., index 112) ; M3 : Falconer & Cautley, 1845, pi. 12,
fig. 13 (width 91 mm., height 114 mm., index 125) ; pi. 12, fig. 12 (width 89 mm.,
height 102 mm., index 115), Osborn, 1942, fig. 842 (width 101 mm., height 124 mm.,
index 123) ; fig. 835 (width 109 mm., height 114 mm., index 105). On the other
hand, in those molars where the height is less than the width, the height has been
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 277
taken at a worn plate, e.g., M3 : Falconer & Cautley, 1846, pi. 14, fig. 8 (width 89
mm., height 63 mm.), Osborn, 1942, fig. 834 (width 100 mm., height c. 88 mm.) ;
M1 : Osborn, 1942, fig. 833 (width 94 mm., height c. 66 mm.) ; fig. 832 (width 80 mm.,
height 65 mm.) ; fig. 831 (width 90 mm., height 67 mm.) ; M3 : Falconer & Cautley,
1845, pi. n, fig. 5 (width 105 mm., height 89 mm.), Falconer & Cautley, 1846, pi.
i8A, fig. i (width 104 mm., height 81 mm.) ; pi. 14, fig. 9 (width 89 mm., height
77 mm.), Osborn, 1942, fig. 855 (width 80 mm., height 53 mm.) ; fig. 839 (width
78 mm., height 45 mm.) ; M1 : Osborn, 1942, fig. 838 (width 80 mm., height 45 mm.).
The examples given above will suffice to show that the unworn molar crowns
of A. planifrons from the Upper Siwaliks are either about as high as wide, or higher
than wide. In those specimens where the height is less than the width (see above)
it is clear from the figures that the height recorded is not the height of the unworn
plate, and height-width indices based on these figures are meaningless. They are
only indicative of the degree of wear shown by the molars in question, and should
be excluded from comparison.
Among the Archidiskodon molars from Bethlehem, only the third molars in the
mandible M 18582 have a height-width index of just over 100 ; in the remaining
specimens the plates are neither fully nor even nearly as high as wide, the highest
height-width index found being 89, the lowest (talons excluded) being 67.
It is therefore evident that the Bethlehem Archidiskodon does not belong to the
species A. planifrons as known from the Upper Siwaliks (Tatrot and Pin j or zones)
of India. The mandible is within the variation limits of the Siwalik specimens,
but the unworn plates of the upper molars (two M2 and one M3) show that the Beth-
lehem Archidiskodon is more primitive than A. planifrons in the height of the unworn
crown being less than the basal width.
Certain primitive archidiskodont molars from the Vaal river gravels in the Trans-
vaal, South Africa, have been described by Osborn (1934, 1942 : 983-988) as A.
subplanifrons , and A. proplanifrons, respectively. Although the Vaal river specimens
are claimed by Osborn to be much more primitive than the most primitive molars
thus far discovered in the Siwaliks (^4. planifrons} the point is somewhat difficult
to make as none of the South African molars is unworn. The holotype of A . subplani-
frons, an M3 dext. (Osborn, 1934, fig. i ; 1942 : 987, fig. 874) is worn to and including
the hind talonid, and is broken off in front through the 5th plate from behind. The
laminar frequency is 4, the enamel thickness, 4 mm. The valleys are V-shaped in
longitudinal section, and the enamel figures of the 3rd and 4th plates from behind
have median anterior and posterior expansions. In all these characters the specimen
resembles the Siwalik A . planifrons, and as neither the plate formula nor the exact
height of the crown can be determined it would seem unjustified to create a new
species for the inclusion of this specimen. The estimated height of the plates is given
by Osborn as 53-63 mm.
The holotype of A. proplanifrons Osborn (1934 : 10, fig. 2 ; 1942 : 986, fig. 873)
is an M3 dext. with all the plates and the hind talon worn ; it is broken off in front
of the 5th plate from behind. The laminar frequency is 3, the thickness of the
enamel is 5 mm. Again, there is no way of telling the full number of plates or the
height of the unworn crown ; Osborn gives the height of the last plate as 55 mm.;
the basal width of the same plate (measured from the figure) is 66 mm. These
278 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
measurements are intermediate between those of the last plates in the two second
upper molars from Bethlehem recorded above (height c. 58 mm., and 54 mm., respec-
tively, and width 55 mm. and 71 mm. respectively), and the resulting height-width
index is also intermediate : 83 in A. proplanifrons, and c. 105, and 76, in the two
Bethlehem specimens. However, in the Bethlehem molars the height increases to
84 mm., and 74 mm., respectively, in the more anteriorly placed plates, and Osborn's
reconstruction of the type of A . proplanifrons, in which all the plates are shown of
the same height as the last, is extremely improbable. The last plate is always lower
than the others. Osborn further compares the M3 of A. proplanifrons with an M3
of A. planifrons (Amer. Mus. 19965 ; Osborn, 1934, fig. 4; 1942, figs. 840, 876)
all the plates of which are worn but boldly reconstructed in longitudinal section
to a height of about 55 mm. This reconstruction, again, is very improbable, for in the
unworn M3 of A. planifrons the crown height is seen to rise to 124 mm. (Amer.
Mus. 19951 ; Osborn, 1942, figs. 842, 845, and 855). Osborn (1934 : 9; 1942 : 986)
says that the worn M3 of A. planifrons is " the most primitive stage found in the
large series of the Archidiskodon planifrons molars collected by Barnum Brown in
the Upper Pliocene Pin j or horizon of the Siwaliks, India ", but it has a laminar
frequency of 4^, whereas the partially unworn M3, thrice figured by Osborn, has a
laminar frequency of 3^-4$, and another (Amer. Mus. 19952 ; Osborn, 1942,
fig. 835) is only 3^-4, which brings A. planifrons within the range of laminar fre-
quencies of the South African and Bethlehem types (3-4).
As stated by Dietrich (1942:79) and by Cooke (1947:455) A. subplanifrons
and A. proplanifrons are conspecific. In my opinion there is no proof at present
that they can be specifically distinguished from the Upper Siwalik A. planifrons
proper, a point that cannot be settled until unworn, entire molars have been found in
the same deposits from which the types of A. subplanifrons and A. proplanifrons
were obtained. In revising the South African " species " of Archidiskodon, Cooke
(1947 : 507) placed A. subplanifrons (including A . proplanifrons} in group I : " Archi-
diskodonts (with broad crowns and thick enamel). A : Low Crowned (less than
75 mm.) ", thereby accepting Osborn's low estimates of the crown heights. It seems
to me that such a group for the South African types is invalid prior to the discovery
of molars with unworn full plates.
Archidiskodon exoptatus Dietrich (1942 : 72) from the Lower Pleistocene of the
Southern Serengeti in Tanganyika, East Africa, is very close to the Upper Siwalik
A . planifrons ; the unworn full plates of the molars are higher than wide. The molars
from Kanam and Laetolil, East Africa, described by Maclnnes (1942 : 86) as Archi-
diskodon planifrons nyanzae, are all worn ; their laminar frequency is about 3^.
None of the molars described by Dietrich (1942) as A. exoptatus has a laminar
frequency of less than 4.
The North African Elephas africanavus Arambourg (1952), found in the Villa-
franchian (Lower Pleistocene) of Ichkeul, Tunisia, has molar crowns that are either
very slightly lower, or higher than wide (height-width index of type M3, 97 ; of
M1, no), which is within the limits of the Upper Siwalik A. planifrons (97-125,
see above). The laminar frequency is 3^ to 4|, the enamel thickness, 4-5 mm. These
specimens also appear to be indistinguishable from those of A. planifrons.
Archidiskodon planifrons has also been recorded from Villafranchian sites in Europe
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 279
(Osborn, 1942 : 961-969 ; Hooijer, 1953) ; the most primitive specimens have laminar
frequencies of 3|— 4, and their crowns are not higher than wide. It should, however,
be realized that these specimens might equally well represent primitive variants of
Archidiskodon meridionalis (Nesti) ; the molars of these two species grade imper-
ceptibly into one another, without a break to indicate a convenient boundary line
between low-crowned A. planifrons and higher-crowned A. meridionalis.
At the end of this brief survey of primitive Eurasian and African archidiskodonts,
let us reconsider the Archidiskodon from Bethlehem. It is apparently identical
with A. planifrons in mandibular characters, but has upper molars that are lower
than wide in the unworn state. There are no counterparts of such molars among the
known material of the Upper Siwalik A. planifrons ; even the geologically oldest
specimen, viz., the M3 from the base of the Tatrot zone (Hooijer, 1956), has unworn
plates higher than wide. The East and North African archidiskodonts mentioned
above agree with A. planifrons in relative crown height, and so do the most primitive
archidiskodonts from the Villafranchian of Europe. The crucial question is whether
the South African A. " subplanifrons " really has lower molar crowns than the
Upper Siwalik species ; this is still unknown. The Bethlehem elephant appears to
represent a somewhat less advanced evolutionary stage than A. planifrons proper,
and should perhaps be sub-specifically distinguished. As long as the " subplanifrons "
problem has not been solved it would seem best to place the Bethlehem elephant
on record as Archidiskodon cf. planifrons.
A large portion of an elephant's tusk, length almost 170 cm. (M 18584), diameter
13 cm. proximally, is so much distorted that the amount of curvature is uncertain.
The tusk, however, appears to have been nearly straight as far as preserved. There
are also a number of vertebrae and limb bones of the Bethlehem elephant, but
except for an atlas they are in a very bad state of preservation. The measurements
of the atlas are given in Table V.
TABLE V. — Measurements, in centimetres, of Atlas of Archidiskodon cf.
planifrons M 18579)
Total height 26
Facies articularis cranialis, vertical . . 15
transverse . c. 9
Antero-posterior diam. of corpus . . . 9-5
Height of foramen vertebrale . . . 13 -5
Least width of idem . . . . . c. 6
Foramen transversarium, diameter . . 3
Foramen for first cranial nerve, diameter . 1-5
The height of the atlas of the A. planifrons skeleton from Chagny-Bellecroix is
27 cm. (Mayet & Roman, 1923 : 82).
The following approximate measurements have been taken from a poorly preserved
radius : length c. 80 cm., greatest proximal width c. n cm., and greatest distal width
c. 16 cm.
The measurements of a left femur (M 18583) are, length from head to media
280 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
condyle c. 112 cm., greatest proximal diameter over head and great trochanter
c. 33 cm., least width of shaft c. 14 cm, greatest distal width c. 24 cm.
In the skeleton of A. planifrons from Chagny-Bellecroix (Mayet & Roman, 1923
84) the femur measures 133 cm. from head to medial condyle.
In a portion of the pelvis from Bethlehem (M 18585) the diameter of the aceta-
bulum is 21 cm.
A right and a left patella, evidently of the same individual (M 18535 and M 18588),
agree in the following dimensions : Height 17-5 cm., width, 13 cm., greatest antero-
posterior diameter, 9 cm.
The reference of the tusk and post-cranial material to A . cf . planifrons is provisional
but it is probable that these specimens belong to the same species as the mandible
and the molars described above because there is no evidence of the presence of more
than one species of elephant in the Bethlehem fauna.
PERISSODACTYLA
EQUIDAE
Hipparion sp.
(PL 35, figs. 3-6)
An isolated right lower last molar (M 18518, PL 35, figs. 3-4) gives evidence of the
presence of Hipparion in the Bethlehem fauna. The tooth is in the germ stage and
lacks the cement coating ; the height of the crown unfortunately cannot be deter-
mined. However, several slightly worn M3 of Hipparion gracile (Kaup) from the
Pontian of Pikermi in the British Museum (Natural History) agree with the Bethle-
hem M3 in every visible detail of their structure. The sharp vertical groove between
metaconid and metastylid is a character common to Hipparion and the zebras,
but Equus stenonis, the smallest zebrine horse of the Villafranchian, has an M3
larger than the Bethlehem tooth (see Major, 1877-80, pi. 7, figs. 21, 30, 31). The
Bethlehem M3 has a length of 25 mm. and a greatest width of 9-5 mm., exclusive
of cement. A Pikermi specimen measures 25-5 mm. antero-posteriorly near the top,
and 26-5 mm. near the base of the crown (also exclusive of cement), and the greatest
transverse diameter of the crown is 11-5 mm., some cement included.
In Hipparion anielopinum (Falconer & Cautley) M3 measures 28 by 12-5 mm.;
in H. theobaldi (Lydekker) M3 measures 30 by 13 mm. (Colbert, 1935 : 148). These
are Middle Siwalik species, but an isolated M3 dext. of Hipparion from the Tatrot
zone, basal Upper Siwaliks, exhibited in the British Museum (Natural History)
is very similar : length 29 mm., and width, 11-5 mm., inclusive of cement. Hipparion
also occurs in the Pin j or zone of the Upper Siwaliks (Pilgrim, 1938 : 447, 449 ; 1944 :
32) where it is associated with Equus ; the latter genus is absent from the underlying
but likewise Villafranchian Tatrot zone (Hooijer & Colbert, 1951).
Hipparion is known to occur in several European Villafranchian faunas, such as
those of Roccaneyra near Perrier, France (Stehlin, 1904, 1929 ; Bout & Azzaroli,
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 281
1952 : 39), of Villarroya, Spain (Villalta, 1952 : 107), and of Kislang, Hungary
(Kretzoi, 1954 : 251). Of these faunas only that of Villarroya lacks Equus, but, as
remarked by Viret (1954 : 182) there is no reason to consider Villarroya any older
than the Villafranchian faunas with Equus. The last lower molar of the Roccaneyra
and Kislang hipparions is unknown ; three specimens of the Villarroya Hipparion
crusafonti Villalta measure 26-2 to 29-9 mm. in length, and 10-1 to 11-5 mm. in width,
cement included (Villalta, 1952 : 119).
Although the crown height of the Bethlehem M3 cannot be measured exactly it
seems improbable that it belongs to the very hypsodont hipparion (Stylohippariori)
which occurs in the Villafranchian of North and East Africa (Arambourg, 1947,
1949 ; Dietrich, 1942) in association with Equus.
There is a second specimen in the Bethlehem collection referable to Hipparion,
viz., the distal portion of a small metapodial (M 18576, PI. 35, figs. 5,6). This specimen
seems to represent the lateral metacarpal or metatarsal, either II sin. or IV dext.
The total length of the fragment is only 4 cm. The shaft is much compressed trans-
versely : the surface facing the large median metapodial is flat, the abaxial surface
convex. The anterior edge is more marked than the posterior, which is rounded.
The distal extremity, somewhat curved backward on the line of the shaft, likewise
is much flattened transversely, and consists of a single condyle with a fossa on either
side. The fossa on the abaxial surface is marked, and surmounted by a weak tubercle.
The distal antero-posterior diameter is 17 mm., the distal condyle width only 8 mm.
The shaft is 7 mm. transversely, and diminishes in antero-posterior diameters
from 15 mm. just above the distal condyle to u mm. at a point only 2 cm. higher
up, where the bone is broken off.
The present metapodial fragment closely resembles the fourth metatarsal of Hip-
parion sp. from Roccaneyra figured by Stehlin (1929, fig. 2B), a cast of which has
been figured as ? Hipparion crusafonti by Villalta (1952 : pi. 23). Whether the speci-
men from Bethlehem belongs to the fore or to the hind foot, and whether it is the
outer or inner lateral metapodial cannot be made out. Determination of its exact
position must await the discovery of more complete material, but this fragment
provides definite proof of the presence of a three-toed horse in the Bethlehem fauna.
RHINOCEROTIDAE
Dicerorhinus etruscus (Falconer)
(PI- 33, ng. 5)
In the Bethlehem collection there is an almost entire skull (M 18542, PI. 33, fig. 5)
which unfortunately is crushed to a considerable extent. All the parts are firmly
cemented with plaster, evidently in the positions in which they were found. A frag-
ment of the palate with P3~4 sin. lies upside down in the right lateral nasal notch,
a partially exposed tooth (? P2 sin.) is seen in the right temporal fossa, and a portion
of distinct bone, possibly of the mandible, projects upward through the left temporal
fossa. The ventral surface of the skull is concealed by a thick mass of cement. The
left zygomatic arch is missing, and the right is broken.
282 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
The occipital portion of the skull is rather well preserved, but somewhat distorted ;
the right maxillary and the zygomatic process of the jugal are displaced out ward and
upward, and are almost in the plane of the dorsal surface of the skull. The right
upper tooth series is more or less in place but comprises only P3-M2, the anterior
premolar (P2) and the third molar being lost.
The dorsal surface of the skull shows numerous cracks and fractures, and the shape
of the dorsal profile, therefore, cannot be relied upon. The nasals (incomplete on the
left side) show a slight rugosity for a horn, and there is no trace of a nasal septum
on their ventral surface ; the skull, therefore, is probably that of a female (cf .
Thenius, 1955) . The rugosity for the second, frontal horn is likewise slight, and hardly
raised ; behind it the skull surface is slightly concave antero-posteriorly, and
gently rises into the nuchal crest, which overhangs the occiput. The infra-orbital
foramen is just behind the nasal notch, which ends posteriorly on a level with the
anterior border of P4. The anterior border of the orbit is above the anterior portion
of M2. The post-tympanic process has united with the post-glenoid process below
the sub-aural channel.
In all these structural characters the skull closely resembles those of Dicerorhinus
etruscus figured by Falconer (1868, pi. 26), Schroeder (1903, pi. i, figs, i, ia), and
Schaub (1944, fig. i), except for the absence of the nasal septum, which apparently
is confined to males (Thenius, 1955 : 11-14). The greatest length of the Bethlehem
skull, from nuchal crest to tip of nasals, is about 68 cm. the greatest length of the
Florence Museum skull figured by Falconer (1868) as well as by Schroeder (1903,
pi. i, figs. 2, 20) is slightly less, 64 cm. The length from the anterior border of the
right orbit to tip of nasals is about 34 cm. in the Bethlehem skull, against about
32 cm. in the Florence Museum skull. The height of the nuchal crest, right side,
from lower surface of condyle, is 15-5 cm. against 16-5 cm. in the Florence Museum
skull. The greatest width of the f rentals (right half only) is 11-5 cm. in both skulls
(Falconer, 1868 : 358). These figures tend to show that the Bethlehem skull agrees
well with that of D. etruscus in size ; further measurements cannot be given because
of the crushing of the specimen.
The dentition of the present specimen is badly preserved ; all the internal borders
of the crowns are incomplete, and the external surfaces broken and distorted. It is,
therefore, impossible to measure the crowns exactly. However, there is little or
no crown cement, the anterior cingulum is prominent and slightly inclined upward,
there is a horizontal lingual cingulum (P3 sin.), and a high and wide entrance to
the medisinus ; all characters pointing to D. etruscus (Falconer, 1868 : 354-368 ;
Dawkins, 1868 ; Wiist, 1901 : 273 ; Schroeder, 1903 ; Bernsen, 1927 ; Vialli, 1956).
In all the premolars and molars present the crochet is rather large, and united to
the crista so as to cut off a medifossette ; this is an individual peculiarity that is
apparently rare in D. etruscus, although in many of the specimens figured by the
above cited authors there are traces of cristae beside the large crochets (Falconer
1868, pi. 29 ; Dawkins, 1868, pi. 8 ; Wiist, 1901, pi. 4, figs. 2, 4, 7, 8 ; Schroeder,
1903, pi. 4 ; Bernsen, 1927, pis. i, 2 ; Vialli, 1956, pi. i, fig. 5). Small cristae are
also seen in the complete upper dentition of D. etruscus figured by Tuccimei (1891,
pi. n).
In the collection there is also a fragment of the maxillary with the right P3-M2
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 283
(M 18563, M 18570). The two premolars have broken off at their bases, but the
anterior portion of the ectoloph of P4 dext. is preserved (M 18577). Of M1 a small
posterior portion, with the laterally compressed post-sinus, only remains. M2 is
the best preserved tooth, lacking, however, the hinder portion. The anterior cingulum
is only slightly inclined upward toward the external side, the crown is low, and does
not show any cement, there is an inner cingulum forming a knob at the wide entrance
to the medisinus, and the crochet is large. There is also a very weak crista, which
would, however, never unite with the crochet. The antero-transverse diameter of
the crown base of M2 is 59 mm.
Nos. M 18563, M 18572, and M 18577 comprise a number of fragments, some of
which may well have formed part of the dentition just mentioned.
An isolated M1 or M2 sin. (M 18562), partially restored with plaster, has all the
characters distinctive of D. etruscus enumerated above. There is no crista, but a
small antecrochet. The lingual cingulum is stronger than that in M 18563.
A left (?) M1 (M 18564), found in 1935, is very incomplete but shows the character-
istic inner cingulum.
A broken and distorted portion of the left ramus of the mandible with part of M3
in situ (M 18558) is of the 1940 season. Portions of lower molars are further contained
in specimens M 18574 and M 18514.
Dicerorhinus etruscus (Falconer), to which the Bethlehem rhinoceros may be
confidently referred, is characteristically a Villafranchian type widely spread in
Europe (see, e.g., Viret, 1954, table opposite p. 184).
ARTIODACTYLA
SUIDAE
Sus cf. strozzii Meneghini
(PI. 33, ng. i ; PI. 34, fig- 8)
In the Bethlehem collection there are two specimens that belong to a suid very
close to or identical with Sus strozzii Meneghini, a species thus far known from the
Villafranchian of Italy, France (Seneze), and the Netherlands (Tegelen) only. The
first of these (M 18515) is a fragment of the right mandibular ramus with the second
premolar (P2) unerupted. The crown measures 12 '8 mm. antero-posteriorly. In a
mandible from Tegelen described by Schreuder (1945 : 188) P2 measures 13*0 by
7-0 mm. The Bethlehem specimen also displays part of the alveolus of the canine ;
the internal upper angle of the alveolus is approximately a right angle, which shows
it to have lodged a canine of the verrucosus type (Azzaroli, 1954 : 44, fig. 2). The
width of the upper surface of the alveolus is over 10 mm. as far as preserved, and
this surface is perfectly straight and the canine thus agrees with those of male
individuals of Sus strozzii (Azzaroli, 1954 : 63, fig. 7). The second specimen (M 18520)
is a left upper incisor, probably I2 sin. , rather worn. The length and width of the crown
are 14 mm. and 6-5 mm, respectively. A left second upper incisor is in situ in a
skull of Sus strozzii from the Val d'Arno figured by Azzaroli (1954 : pi. 10, fig. 40),
284 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
but its measurements have not been given. The upper incisors of the Tegelen Sus
strozzii are unknown as yet, and in the sub-adult Seneze specimen (Schaub, 1944 :
276, fig. 3 ; Azzaroli, 1954 : pi. 15, fig. 46) they are missing.
The existence of Sus cf . strozzii, at any rate of a large suid with a verrucosus-like
lower canine, at Bethlehem is interesting in view of the fact that Sus strozzii of the
Val d'Arno, Seneze, and Tegelen is generally accepted to be an invading form from
Asia. There is no fossil species of Sus known from Asia that meets all the conditions
ancestral to Sus strozzii, however, and both Schaub (1944 : 277) and Azzaroli (1954)
derive Sus strozzii from Sus minor Deperet of the Pliocene of Roussillon.
GlRAFFIDAE
Giraffa cf. camelopardalis (L.)
(PI. 34, ngs. 5, 6)
A small fragment of a molar, the distal portion of a metapodial and a crushed
distal condyle of another metapodial give evidence of the presence of a giraffe in
the Bethlehem fauna.
The molar fragment (M 18517, PI. 34, figs. 5, 6) is part of the external surface of
a left upper molar, probably M3. It comprises the portion just behind the prominent
median rib of the antero-external cusp (paracone) backward to and including the
weak median rib of the postero-external cusp (metacone). The preserved portion
of the paracone slopes inward, and abuts against the metacone with a small but
distinct, hook-shaped, outward turn. Between the paracone and the anterior style
of the metacone there is an outer valley, 2 mm. wide and about 4 mm. deep, that
extends rootward to 7 mm. from the crown base. The anterior style of the metacone
is very prominent, and marked off behind by a sharp fold that likewise extends
rootward to 7 mm. from the border of the enamel. It is worn down to a height of
13 mm. from the crown base ; the highest points of the paracone in front, and of
the metacone behind it are 16 mm. high from the crown base, as worn. The median
external rib of the metacone is weakly developed, and is more inclined inward than
the paracone. The enamel is rugose. Internally, the molar fragment is broken
off vertically just along the high and narrow pulp cavities of the external cusps.
I have compared the present fossil fragment with a number of dentitions of the
Recent giraffe, and found the closest resemblance with the external portion of the
left M3. Doubtless there is much individual variation in the development of the
anterior style of the metacone, which is very strongly marked in some, and only
slight in others. The closest approach to the condition seen in the fossil is in two
specimens (Leiden Museum, 7085 and 4216), in which the anterior metacone style
is distinctly marked off behind by a groove, and as prominent laterally as that
in the fossil, although not quite so thick (3-4 instead of 5-6 mm.). The valley just
anterior to it is a little wider in the Recent specimens than in the fossil, and the basal
cingulum in the Recent specimens is a trifle less high (5-6 mm. instead of 7 mm.).
However, had more Recent dentitions been available it is quite probable that the
condition observed in the fossil would be seen to fall within the limits of individual
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 285
variation in the Recent giraffe. It is important to note that the fossil specimen does
not appear to be larger than the Recent ; the distance between the apices of the
external surfaces of para- and metacone is about 14 mm. in the fossil as well as in the
Recent molars.
The next specimen referable to a giraffe is the distal portion of a metatarsal
(M 18508, found in 1934. It has largely been restored with plaster. The dorsal
surface has a shallow vascular groove, as in the metatarsals of the Recent giraffe, but
the foramen perforating the shaft distally does not show. The volar surface is
restored with plaster for a height of 3-4 cm. above the condyles, but more proximally
shows a weak median ridge, emerging about 6 cm. above the condyles, and flattening
out at about n cm. above the condyles, at which level the bone is broken off. At
this point the volar surface is still slightly convex transversely. In Recent meta-
tarsals of Giraffa camelopardalis there is exactly the same condition, the posterior
median longitudinal groove that extends along most of the length of the shaft fades
away some 12-15 cm. above the distal condyles, to be replaced distally by a weak
median ridge. The greatest distal width of the fossil metatarsal is about 85 mm.
(85-90 mm. in three Recent metatarsals), and the greatest dorso- volar diameter of
the condyles in the fossil specimen is 54 mm. (54-55 mm. in the Recent). The distal
condyle width is greater in the fossil than in the Recent metatarsals (about 84 mm.
as opposed to 78-80 mm.), but the condyles are slightly less closely approximated
in the fossil, which is apparently due to a slight fault in the reconstruction. The
condyles are shaped exactly as those in the Recent specimens used for comparison.
The last specimen (M 18507) is an isolated distal condyle of a metapodial, broken,
and with the abaxial part displaced upward relative to the axial portion. Exact
measurements cannot be given, but the specimen seems to be slightly larger than the
corresponding part of the metatarsal, and, therefore, may have formed part of a
metacarpal instead.
The remains described above differ slightly, if at all, from the Recent giraffe,
and may be identified as Giraffa ci. camelopardalis (L.). This species has never been
recorded in the fossil state from Europe, but remains apparently indistinguishable
from the living species of Giraffa have been recorded from Villafranchian deposits
in East and South Africa (Dietrich, 1942 : 112 ; Arambourg, 1947 : 375 ; Cooke
& Wells, 1947). The presence of this form sets a decidedly African stamp upon the
Bethlehem fauna.
BOVIDAE
Leptobos sp. nov.?
(PL 33, fig. 4)
A right horn core (M 18522) broken off approximately at the base, and without
the tip, must be referred to Leptobos. The core is slightly compressed vertically
at the base, and is curved backward and outward in a gentle curve ; the distal
half is curved upward. In the basal portion there are distinct longitudinal grooves
along the posterior upper and the anterior lower surfaces ; these grooves indicate
a slight anti-clockwise torsion. They flatten out toward the middle of the length
286 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
except for one along the lower surface anteriorly that continues to about 10 cm. from
the broken distal end. Another, along the lower surface posteriorly, originates only
in the middle of the length, and flattens out some 15 cm. from the broken end. Apart
from these grooves the distal portion is smooth.
The length of the horn core from base to broken end is 65 cm. along the outer curve ;
it describes about one-fourth of a circle. The basal diameters are about 94 mm.
horizontally, and 86 mm. vertically ; these diameters have diminished to 76 mm.,
both ways, at the middle of the length. At the broken distal end the core still measures
48 mm. in diameter, and remains perfectly round in cross section.
The present specimen, although incomplete, resembles the horn core of Leptobos
stenometopon (Riitimeyer, 1867, pi. i, figs. 3, 4 ; 1878, pi. 7, fig. 3 ; Merla, 1949,
pi. 8, fig. i a, c) except in size : the horn core of the latter species is only two-thirds
as long as the incomplete Bethlehem specimen (Table VI). The Bethlehem horn
core differs from the other European Villafranchian species of Leptobos, viz., L.
etruscus (Falconer) and L. vallisarni Merla, in having the convexity backward and
inward instead of outward (Merla, 1949 : 71, fig. 2). The horn cores of the two last-
mentioned species, moreover, are shorter than the Bethlehem specimen. In Leptobos
falconeri Riitimeyer of the Pinjor zone of the Upper Siwaliks the complete horn core
is unknown ; the American Museum specimen restored by Pilgrim (1937 : 816,
817), although it has the convexity backward and inward just as in the Bethlehem
specimen, is less curved than either the Bethlehem core or that of L. stenometopon.
TABLE VI. — Measurements of Horn Core of Leptobos species
Bethlehem stenometopon etruscus vallisarni falconeri
Length along outer curve . 650+ 420 500 330 c. 590
Basal diameters . . . 94 x 86 . 72 x 58 . 79 X 73 . 88 x 77 . 85 x 77
Idem, middle of length . . 76 x 76 . 56 x 46 . 50 x 50 . 75 x 62
Idem, 7 cm. from tip . . (48) . 35 x 32 . 32 x 34 . 45 x 40 .
The Bethlehem core undoubtedly represents a species of Leptobos, but does not
seem to be identical with any of the European species1 (Merla, 1949) or with the Upper
Siwalik L. falconeri (Table VI). There are further specimens of Leptobos in the Beth-
lehem collection, viz., a horn core fragment about 71 by 66 mm. in diameters
(M 18556), a broken distal condyle of a metapodial (M 18552), a proximal sesamoid
(M 18567), and, finally, two fragments of what appear to be upper premolars (M 18519)
but these specimens are of no use for comparative purposes. To indicate the possi-
bility that the Bethlehem Leptobos eventually may prove to be a new species it is
recorded provisionally as Leptobos sp. nov.?
Gazellospira torticornis (Aymard)
(PI. 33, figs. 2, 3)
The collection contains four mandibular rami, six isolated molars and molar
fragments, as well as the distal epiphysis of a right radius and a second phalanx
that belong to an antelope. The upper dentition is badly represented : two very
1 Leptobos iithe only bovine in the Villafranchian of Europe (Pilgrim, 1938 : 451, 466 ; 1944 : 29).
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 287
incomplete upper molars (M 18548, and one fragment M 18519) do not show any
diagnostic characters. The lower dentition, however, is very characteristic.
A left horizontal ramus of the mandible (M 18536) has the full permanent dentition
P2-M3 (PL 33, fig. 3) . The ramus is fractured below and in front of the premolars, and
the fractures are filled with matrix, leaving a diastema between P4 and Mj, while P2
is displaced backward and outward. The teeth are undamaged except for P2, which
lacks part of the anterior border of the crown. P2 is simply built ; its external surface
is about as wide as high, and shows a single cusp and a very weak posterior fold.
The internal surface is not exposed. P3 has an inner cusp, two anterior and two
posterior wings. The anterior valley is open internally, but the inner cusp is united
to the posterior wing, the posterior outer fold is stronger, and the external surface is
higher than wide. P4 has a closed internal wall ; there is a small but marked anterior
internal fold, while the posterior lobe is marked off by a deep external fold. The
molars are characterized by weak internal ribs, strong anterior folds both internally
and on the external side, and the apparent absence of basal pillars. The crowns
are hypsodont and rather narrow ; M.^_2 possess a postero-internal fold, M3 has a
laterally compressed third lobe.
A right ramus of the mandible (M 18534) lacks the two anterior premolars, and the
third lobe of M3 ; the ramus is broken and also distorted at Mx. In the closed internal
wall and marked external fold P4 resembles that of the preceding ramus closely.
The molars likewise agree with those of the foregoing specimen in the presence of
strong anterior folds, internally as well as externally, and in their weakly ribbed
internal surfaces. Basal pillars are not exposed either.
M 18533 is a younger specimen than the foregoing rami ; the milk molars DM2_4
are still in place, and M3 just appears above the alveolar margin. DM2 and DM3
resemble their successors in the permanent dentition in crown structure but are
narrower and more elongated ; DM4 is three-lobed ; the cingulum forms a knob
at the base of the groove separating the second and third lobe externally. As in the
molars, there is a postero-internal fold. The molars show the anterior folds very
distinctly ; the internal ribs are more marked apically than those in the more worn
molars of M 18536 and M 18534, but become less distinct toward the base. The
antero-posterior diameter of the crown diminishes rootward so that the length of
an unworn or slightly worn molar is greater than that of a much worn specimen.
The hypsodont character of the molars is shown by a fragment of the left mandibular
ramus (M 18532) with M3 unerupted ; the height of the slightly worn M2 in this
specimen is not less than 30 mm. by a transverse diameter of the crown of only
8 -5 mm.; the antero-posterior diameter of the crown is 19-5 mm. apically.
An isolated right Mx (M 18546), moderately worn, and a right M3 (M 18545),
the second and third lobes of which are incomplete, possess the characteristic anterior
folds and weak ribs of the preceding specimens. In these specimens the base is exposed,
and there are cingular knobs at the external grooves between the lobes basally.
The transverse diameters of the crowns slightly increase rootward, while the antero-
posterior diameters decrease from the apex toward the base of the crown. Finally,
a much worn M3 sin. (M 18537) shows that the third lobe is shorter antero-posteriorly
at base than the second lobe, and much compressed laterally. There is no basal
knob in the groove between the second and third lobes, but there is a regular basal
288 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
pillar between the first and second lobes externally. The folds at the anterior end
of the crown are very marked on both sides. In Mj the postero-internal fold is also
marked.
TABLE VII. — Measurements of Gazellospira torticornis
DM2, ant. post.
transv.
DM3, ant. post.
transv.
DM4, ant. post.
transv.
Length DM2_4
P2, ant. post.
transv.
P3, ant. post.
transv.
P4, ant. post.
transv.
Length P2_4
M1( ant. post.
transv.
M2, ant. post.
transv.
M3, ant. post.
transv.
Length M1-3
Height of ramus below Mx
Idem, below P,
Mi8533
7'5
3'9
ir-i
5'5
17-8
8-1
37'5
Mi 8545
9'3
20-0
9-0
24
13-0
7'7
32
16-5
9'9
19-0
9-6
8-7
27
23
c.g
10-8
7-0
13-0
7-8
15-9
Pilgrim &
Schaub,
1939
8-2
TO-5
19
36
9-5-10
14
c.i6
37
9-1
18-7
9-5
22-8
7'9
58
26
17-9
9-0
19-5
8-5
10-2
6 1 -5-72
24
21
Although nothing of the skull or of the horn cores is preserved, the lower dentition
of the Bethlehem antelope leaves no doubt as to its specific position : it clearly
belongs to Gazellospira torticornis (Aymard) as described by Pilgrim & Schaub
(1939). There is such a close resemblance between the Bethlehem specimens and
those from the Villafranchian of Roccaneyra, La Loubiere de Pardines, and Seneze
described and figured by Pilgrim & Schaub (1939), not only in structural details
but also in size (Table VII), that their conspecificity is rendered certain. The ascending
ramus of the Bethlehem mandibles is not preserved, but the anterior end is more
complete than in the specimens figured by Pilgrim & Schaub ; the distance between
P2 and the mental foramen is 30 mm. both in M 18534 and in M 18533, and the height
of the ramus half way between, 15 mm.
With the aid of Pilgrim & Schaub's monograph it has also been possible to identify
a few limb bones in the Bethlehem collection : the distal epiphysis of a right radius
(M 18510) agrees perfectly with that of Gazellospira from Roccaneyra (Pilgrim &
Schaub, 1939, pi. 3, fig. 20), and differs from that of deer in the same points mentioned
(p. 15) : the lunar and scaphoid facets are wider, the former almost reaches the ulna,
and extends more distally, the latter is less convex antero-posteriorly, etc. The
greatest distal width cannot be measured in the Bethlehem specimen, but that of the
distal articular surface is 35 mm. as in a Seneze specimen (p. 19). There is also a
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 289
second phalanx (M 18575), damaged proximally, the length of which is 30 mm. and
the proximal width, 15 mm., as in a Gazellospira phalanx II from Seneze (Pilgrim
& Schaub, 1939 : 20, pi. 3, fig. 6. lower figs.).
Gazellospira lorticornis is a characteristically Villafranchian species known from
France, Italy, and Spain (Villarroya : Schaub, 1944 : 280). Its occurrence at Beth-
lehem is of interest as according to Pilgrim & Schaub (1939 : 29) the species may be
considered to be an immigrant from Asia.
AGE AND COMPOSITION OF THE BETHLEHEM FAUNA
The faunal list of the mammals from Bethlehem resulting from the present study is
as follows :
Nyctereutes megamastoides (Pomel).
Homotherium (?) sp.
Archidiskodon cf. planifrons (Falconer & Cautley).
Hipparion sp.
Dicerorhinus etruscus (Falconer).
Sus cf. strozzii Meneghini.
Giraffa cf. camelopardalis (L.).
Leptobos sp. nov.?
Gazellospira torticornis (Aymard).
This faunal assemblage as a whole leaves no doubt as to the age of the bone-bearing
beds of Bethlehem. It is a characteristic Villafranchian fauna, and there is general
agreement nowadays among palaeomammalogists that the Villafranchian should
be assigned to the basal Pleistocene. Thus, Miss Bate's preliminary statement
of the age of the Bethlehem fauna is borne out by the present study.
When the list given above is compared with Miss Bate's provisional list of 1937
given in the introduction to the present paper certain discrepancies will be observed.
I have found no evidence of the presence of Hippopotamus in the Bethlehem fauna,
and none of Stegodon either. It is suggested that Miss Bate's record of the hippopo-
tamus is based on a deceptive fragment of an Archidiskodon molar, in which the median
looped expansions of the plates, at a certain stage of wear of the crown, may resemble
the trefoil pattern of the cusps in a Hippopotamus molar. There is such a misleading
fragment in the Bethlehem collection (M 18543). Likewise, it would seem evident
that Miss Bate's record of Stegodon in reality is based on an Archidiskodon molar.
The Bethlehem Archidiskodon cf. planifrons is an exceptionally low-crowned form,
and such forms may be easily mistaken for stegodonts. On the other hand, I have
been able to add one faunal element, viz., Sus cf. strozzii. This form is poorly repre-
sented in the collection, and the two small fragments must have been overlooked in
the provisional study of the Bethlehem fauna.
We know nothing as yet of the so-called microfauna, the small mammals such as
insectivores, bats and rodents. The condition in which the small carnivore (Nyctereutes
megamastoides) was found, however, is an indication that the small animals would
have been unable to withstand the conditions of the bone-bearing deposit.
On comparing the Bethlehem fauna as we now know it with well-known European
ago AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
Villafranchian faunas such as those of Perrier, Seneze, and Saint- Vallier (Schaub,
1944 ; Viret, 1954) we notice two important differences : the Bethlehem fauna
lacks Cervidae, and contains Giraffa. In the absence of cervids and the presence of
the giraffe the Bethlehem fauna agrees with those from the Villafranchian of East
and South Africa (Arambourg, 1947). The Bethlehem fauna, however, differs
from the East and South African faunas of the same age in the presence of Nyctereutes,
Dicerorhinus, and Gazellospira, which are characteristic of the Villafranchian of
Europe.
Therefore, the Bethlehem fauna is predominantly European, or rather Eurasiatic
in character, with an African stamp upon it, as, in fact, might have been expected
a priori from its geographic position.
As such, the Bethlehem fauna provides a faunistic link for the Villafranchian
between the continents to the north and that to the south. It is one of those rare
Villafranchian faunas such as that of the Tatrot zone of the Upper Siwaliks and that
of Villarroya in Spain in which Hipparion lingers on and from which Equus is absent.
Its Archidiskodon would appear to be more primitive even than that of the Tatrot
(Hooijer, 1956), but the faunal assemblage of Bethlehem is as characteristically
Villafranchian as any, and adds materially to our knowledge of the distribution of
the sites over the Old World that give evidence of the great mammalian migration
that occurred, we believe, at the beginning of the Pleistocene.
The so-called industry of the Bethlehem bone-bearing beds (see Gardner & Bate,
1937) has been re-examined by Dr. J. Desmond Clark who concluded that all the
flaking has the characteristics of a natural origin. His report will be published later.
REFERENCES
ARAMBOURG, C. 1947. Contribution a 1'etude geologique et paleontologique du bassin du lac
Rodolphe et de la basse vallee de 1'Omo. Deuxieme partie, Paleontologie. Mission
scient. Omo 1932-1933, 1, Geol.-Anthrop. : 232-562, 40 pis., 91 text-figs.
1949. Les gisements de vertebres villafranchiens de 1'Afrique du Nord. Bull. Soc. geol.
Fr., Paris (5) 19 : 195-203, 2 figs.
1952. Note preliminaire sur quelques Elephants fossiles de Berberie. Bull. Mus. Hist.
nat. Paris (2) 24 : 407-418, pi i.
AZZAROLI, A. 1954. Filogenesi e biologia di Sus strozzii e di Sus minor. Palaeontogr. ital.,
Pisa (N.S.) 18 : 41-76, pis. 9-20.
BATE, D. M. A. 1934. Discovery of a fossil elephant in Palestine. Nature, Lond., 134 : 219.
1941- The bone-bearing beds of Bethlehem. Nature, Lond., 147 : 783.
BERNSEN, J. J. A. 1927. The geology of the Teglian clay and its fossil remains of Rhinoceros.
iii -+- 108 pp., 12 pis. 's Hertogenbosch.
BOULE, M. 1889. Le Canis megamastoides du Pliocene moyen de Perrier (Puy-de-D6me).
Bull. Soc. geol. Fr., Paris (3) 17 : 321-330, pi. 7.
BOUT, P. & AZZAROLI, A. 1952. Stratigraphie et faune du creux de Peyrolles pres Perrier
(Puy-de-D6me). Ann. Paleont., Paris, 38 : 37-56, n figs.
COLBERT, E. H. 1935. Siwalik mammals in the American Museum of Natural History. Trans.
Amer. Phil. Soc., Philadelphia (N.S.) 26 : 1-401, 198 figs.
COOKE, H. B. S. 1947. Variations in the molars of the living African elephant and a critical
revision of the fossil Proboscidea of Southern Africa, Part I. Amer. J. Sci., New Haven,
245 : 434-457, 7 figs. ; Part II : 492-517, 7 figs.
COOKE, H. B. S. & WELLS, L. H. 1947. Fossil mammals from the Makapan Valley, Potgieters-
rust, III. Giraffidae. S. Afr. J. Sci., Johannesburg, 43 : 232-235, i fig.
AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM 291
DAWKINS, W. BOYD. 1868. On the dentition of Rhinoceros etruscus Falc. Quart. J. Geol.
Soc. Land., 24 : 207-218, pis. 7, 8.
DIETRICH, W. O. 1942. Altestquartare Saugetiere aus der siidlichen Serengeti, Deutsch-
Ostafrika. Palaeontographica, Stuttgart, 94, A : 43-133. Pls- 3~23-
EWER, R. F. 1955. The fossil carnivores of the Transvaal Caves : Machairodontinae. Proc.
Zool. Soc. Land., 125 : 587-615, pis. 1-3.
FALCONER, H. 1868. Palaeontological Memoirs and Notes. 2 vols. London.
FALCONER, H. & CAUTLEY, P. T. 1845-49. Fauna Antiqua Sivalensis. Pis. 1-12 (1845) ;
pis. 13-24 (1846) ; pis. 25-80 (1847) ; pis. 81-92 (1849). London.
GARDNER, E. W. & BATE, D. M. A. 1937. The bone-bearing beds of Bethlehem : their fauna
and industry. Nature, Lond., 140 : 431-433.
HOOIJER, D. A. 1953. On dredged specimens of Anancus, Archidiskodon, and Equus from
the Schelde estuary, Netherlands. Leid. geol. Meded., Leiden, 17 : 185-201, pis. r, 2.
1955- Fossil Proboscidea from the Malay Archipelago and the Punjab. Zool. Verh.,
Leiden, 28 : 1-146, pis. 1-17.
1956. Archidiskodon planifrons (Falconer & Cautley) from the Tatrot zone of the Upper
Siwaliks. Leid. geol. Meded., Leiden, 20 : 110-119, pi. i.
HOOIJER, D. A. & COLBERT, E. H. 1951. A note on the Plio-Pleistocene boundary in the
Siwalik series of India and in Java. Amer. J. Sci., New Haven, 249 : 533-538.
KRETZOI, M. 1954. Bericht uber die Calabrische (Villafranchische) Fauna von Kislang, Kom.
Fej&r. Jber. ung. geol. Anst., Budapest, 1953, i : 239-264.
LEWIS, G. E. 1937. A new Siwalik correlation. Amer. J. Sci., New Haven (5) 33 : 191-204.
MACINNES, D. G. 1942. Miocene and post-Miocene Proboscidea from East Africa. Trans.
Zool. Soc. Lond., 25 : 33-106, pis. 1-8.
MAJOR, C. J. F. 1877-80. Beitrage zur Geschichte der fossilen Pferde insbesondere Italiens.
Abh. schweiz. paldont. Ges., Zurich, 4 : 1-16 (1877). 7 : 17-153 (1880), pis. 1-7.
MA YET, L. & ROMAN, F. 1923. Les Elephants Pliocenes, Part I. Ann. Univ. Lyon (N.S.)
42 : 1-88, 15 figs.
MERLA, G. 1949. I Leptobos Rutim. italiani. Palaeontogr. ital., Pisa (N.S.) 16 : 41-155, pis. 5-11.
OSBORN, H. F. 1934. Primitive Archidiskodon and Palaeoloxodon of South Africa. Amer.
Mus. Novit., New York, 741 : 1-15, 5 figs.
1942. Proboscidea, II. Stegodontoidea, Elephantoidea. xxvii -\- 805-1675 pp., 18 pis.
Amer. Mus. Nat. Hist., New York.
PEI, W. C. 1934. On the Carnivora from Locality i of Choukoutien. Palaeont. sinica,
Peking (C) 8, i : 1-216, pis. 1-24.
1940. The Upper Cave fauna of Choukoutien. Palaeont. sinica, Peking (N.S., C) 10 :
1-100, pis. 1-8.
PILGRIM, G. E. 1937. Siwalik Antelopes and Oxen in the American Museum of Natural
History. Bull. Amer. Mus. Nat. Hist., New York, 72 : 729-874, 81 figs.
1938. Are the Equidae reliable for the correlation of the Siwaliks with the Coenozoic
stages of North America ? Rec. Geol. Surv. India, Calcutta, 73 : 437-472, 479-482.
1944- The lower limit of the Pleistocene in Europe and Asia. Geol. Mag., London,
81 : 28-38.
PILGRIM, G. E. & SCHAUB, S. 1939. Die schraubenhornige Antilope des europaischen Ober-
pliocaens und ihre systematische Stellung. Abh. schweiz. palaont. Ges., Zurich, 62 : 1-30,
pis. 1-3.
RUTIMEYER, L. 1866-67. Versuch einer naturlichen Geschichte des Rindes in seinen
Beziehungen zu den Wiederkauern im Allgemeinen. N. Denkschr. schweiz. naturf. Ges.,
Zurich, 22. Pt. i : 1-102, pis. i, 2 (1866). Pt. 2 : 1-175, pis. 1-4 (1867).
1877-78. Die Rinder der Tertiar-Epoche, nebst Vorstudien zu einer naturlichen
Geschichte der Antilopen. Abh. schweiz. paldont. Ges., Zurich, 4 : 1-72 (1877), 5 : 73-208,
pis. 1-7 (1878).
SCHAUB, S. 1934. Observations critiques sur quelques Machairodontides. Eel. geol. Helv.,
Lausanne, 27 : 399-406.
1944. Die oberpliocaene Saugetierfauna von Seneze (Haute-Loire) und ihre verbreitungs-
geschichtliche Stellung. Eel. geol. Helv., Lausanne, 36 : 270-289, 8 figs.
292 AN EARLY PLEISTOCENE MAMMALIAN FAUNA FROM BETHLEHEM
SCHREUDER, A. 1945- The Tegelen fauna, with a description of new remains of its rare
components (Leptobos, Archidiskodon meridionalis Macaca, Sus strozzii). Arch, neerl.
Zool., Leiden, 7 : 153-204, pi. 3.
SCHROEDER, H. 1903. Die Wirbelthier-Fauna des Mosbacher Sandes, I. Gattung Rhino-
ceros. Abh. preuss. geol Landesanst., Berlin (N.S.) 18 : 1-143, P^s- 1-14-
SIMPSON, G. G. 1945. The principles of classification and a classification of mammals. Bull.
Amer. Mus. Nat. Hist., New York, 85 : 1-350.
STEHLIN, H. G. 1904. Une faune a Hipparion a Perrier. Bull. Soc. geol. Fr., Paris (4) 4 :
432-444, pis. ii, 12.
1929. Bemerkungen zu der Frage nach der unmittelbaren Ascendenz des Genus Equus.
Eel. geol. Helv., 22 : 186-201, 4 figs.
TEILHARD DE CHARDIN, P. & PEI, W. C. 1941. The fossil mammals of Locality 13 of Chou-
koutien. Palaeont. sinica, Peking (N.S., C) 11 : 1-106, pis. 1-6.
TEILHARD DE CHARDIN, P. & PIVETEAU, J. 1930. Les mammiferes fossiles de Nihowan
(Chine). Ann. Paleont., Paris, 19 : 1-134, pis- J-23-
TEILHARD DE CHARDIN, P. & YOUNG, C. C. 1936. On the mammalian remains from the
archaeological site of Anyang. Palaeont. sinica, Peking (C) 12, i : 1-78, pis. 1-8.
THENIUS, E. 1955. Die Verknocherung der Nasenscheidewand bei Rhinocerotiden und ihr
systematischer Wert. Abh. schweiz. paldont. Ges., Zurich, 71, 4 : 1-17, pi. i.
TUCCIMEI, G. 1891. Alcuni mammiferi fossili delle provincie umbra e romana. Mem. Acad.
Nuovi Lincei, Roma, 7 : 89-152, pis. 5-11.
VIALLI, V. 1956. Sul Rinoceronte e 1'Elefante dei livelli superiori della serie lacustre di
Leffe (Bergamo). Mem. Soc. ital. Sci. nat., Milano, 12 : 1-70, pis. 1-6.
VILLALTA, J. F. DE. 1952. Contribucion al conocimiento de la fauna de mamiferos fosiles
del Plioceno de Villarroya (Logrono). Bol. Inst. geol. Esp., Madrid, 64 : 1-203, pis. 1-27.
VIRET, J. 1954. Le loess a banes durcis de Saint- Vallier (Drome) et sa faune de mammiferes
villafranchiens. Nouv. Arch. Mus. Hist. nat. Lyon, 4 : 1-200, pis. 1-33.
WtJST, E. 1901. Untersuchungen iiber das Pliozan und das alteste Pleistozan Thuringens.
Abh. naturf. Ges. Halle, 23 : 21-368, pis. 1-9.
PLATE 32
FIG. i. Mandible, right view
FIG. 2. Same, top view, x 2/11.
Archidiskodon cf. planifrons (Falconer & Cautley)
right view, x 2/11. M 18582. Bethlehem.
f rioixr V O /T T
Bull. B.M. (N.H.) Geol. 3, 8
PLATE 32
PLATE 33
Sus cf . strozzii Meneghini
FIG. i. Fragment of right ramus of the mandible with P2 and alveolus of C, outer view.
X ij. M 18515. Bethlehem.
Gazellospira torticornis (Aymard)
FIG. 2. Left ramus of the mandible with P2-M3, inner view, x f. M 18536. Bethlehem.
FIG. 3. Same, outer view, x f .
Leptobos sp. nov. ?
FIG. 4. Right horn core, anterior view, x 2/9. M 18522. Bethlehem.
Dicerorhinus etruscus (Falconer)
FIG. 5. Skull (crushed), right view. X \. M 18542. Bethlehem.
Bull. B.M. (N.H.) Geol. 3, 8
PLATE 33
*
w,' «
i* •
PLATE 34
Homotherium (?) sp.
FIG. i. Left lower carnassial (Mx), outer view, x i£. M 18511. Bethlehem.
FIG. 2. Same, crown view. X ij.
Archidiskodon cf. plamfrons (Falconer & Cautley)
FIG. 3. M2 dext., crown view. x 5/11. M 18523. Bethlehem.
FIG. 4. Same, inner view, x 5/11.
Giraffa cf. camelopardalis (L.)
FIG. 5. External fragment of left upper M, crown view. x i^. M 18517. Bethlehem.
FIG. 6. Same, outer view, x ij.
Nyctereutes megamastoides (Pomel)
FIG. 7. Fragment of left ramus of the mandible with P3_4, outer view. X ij. M 18521.
Bethlehem.
Sus cf. strozzii Meneghini
FIG. 8. Left upper I, inner view, x i£. M. 18520. Bethlehem.
Bull. B.M. (N.H.) Geol. 3, 8
PLATE 34
PLATE 35
Archidiskodon cf. planifrons (Falconer & Cautley)
FIG. i. M2 sin., crown view, x 5/11. M 18524. Bethlehem.
FIG. 2. Same, inner view, x 5/11.
Hipparion sp.
FIG. 3. M3 dext., crown view. x i£. M 18518. Bethlehem.
FIG. 4. Same, external view. X ij.
FIG. 5. Distal portion of metapodial II sin. or IV dext., abaxial view. x i\. M 18576.
Bethlehem.
FIG. 6. Same, front view, x ij.
Bull. B.M. (N.H.) Geol. 3, 8
PLATE 35
THE UPPER PERMIAN FLORA
OF ENGLAND
H. M. M. STONELEY
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 9
LONDON: 1958
BULLETIN OF
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THE UPPER PERMIAN FLORA OF ENGLAND
BY
HILDA M. M. STONELEY
— to
Pp. 293-337 ; Pis. 36-40 ; 16 Text-figures
BULLETIN OF
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GEOLOGY Vol. 3 No. 9
LONDON: 1958
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THE UPPER PERMIAN FLORA OF ENGLAND
By HILDA M. M. STONELEY
SYNOPSIS
The paper brings together existing information relating to the scanty flora of the Upper
Permian beds of northern England. Of the 24 fossil plant species described, the majority are
also known from the German Zechstein, but there are new species of the algal genera Piaea
and Calathella, a new cycadophyte provisionally included in Pseudoctenis, and a representative
of Hiltonia, recently described as a new genus elsewhere. Other plant remains not referable to
Zechstein species closely recall Cordaites aequalis (Goppert) and Psygmophyllum cuneifolium
(Kutorga), both well-known Russian Permian species ; these records, however, need confirma-
tion from more adequate material. The cuticle structure of several species is described. Leaf
fragments with preserved cuticles of distinctive types show that further species are represented
in the English Upper Permian, but the material is too imperfect for description. Certain non-
vegetable structures previously regarded as fossil plant remains are also discussed.
I. INTRODUCTION
IN England beds of established Upper Permian (Thuringian) age have two main
outcrops, one east of the Pennines, extending from the south of Northumberland
to near Nottingham, and the other west of the Pennines. There are also small
outcrops still further west. These beds lie unconformably on Upper Carboniferous
rocks which are not the latest known in this country. Rocks of intermediate age
succeed the Coal Measures conformably in Warwickshire, Staffordshire and some
adjacent areas. Some must unquestionably be included in the Upper Carboniferous,
but the youngest, a series of red marls, sandstones and breccias known as the Enville
or Corley Group, have been assigned by some writers to the Lower Permian (Autunian) .
They have yielded a scanty flora of which it is hoped to give an account elsewhere.
No plant-bearing beds thought to be of Middle Permian (Saxonian) age occur in this
country.
The Upper Permian rocks contain a flora which has hitherto been little investigated
and is the subject of the present paper. Identifiable specimens are, however, very
limited in number and are largely from localities no longer available for collecting.
An attempt has, therefore, been made to locate and examine all such specimens
found in the past. Material was collected personally, mainly at Hilton and Kimberley,
where plant remains still abound but are mostly ill-preserved. The writer thanks the
authorities of the following institutions for access to and the loan of specimens :
GEOL. 3, 9. 27
296 THE UPPER PERMIAN FLORA OF ENGLAND
British Museum (Natural History)1 ; Geological Survey2, Royal Scottish, Hancock
(Newcastle-upon-Tyne), Yorkshire, Manchester, Carlisle, Dorman Memorial (Middles-
brough), Sunderland, Warwick, Wollaton Hall (Nottingham), and Birmingham
City Museums ; Geological Departments of Birmingham, Nottingham, and Leeds
Universities, King's College, Newcastle-upon-Tyne, and University College, Galway
(where W. King's collection is deposited). Dr. C. T. Trechmann kindly lent specimens,
since presented to the British Museum (Natural History), from his own collection,
and Professor R. Krausel sent for comparison type specimens of German Upper
Permian algae belonging to the Senckenberg Museum. The writer also wishes to
acknowledge most helpful advice given her by Professor R. Florin and Dr. Britta
Lundblad during a visit to Stockholm, and by Professor T. M. Harris. The investiga-
tion, undertaken at the suggestion of the late Mr. W. N. Edwards, was carried out
with the aid of a grant from the Department of Scientific and Industrial Research
in the Sedgwick Museum, Cambridge, to the authorities of which the writer tenders
her thanks.
2. HISTORY OF PREVIOUS WORK ON THE ENGLISH
UPPER PERMIAN FLORA
The first record of the occurrence of plant remains in English Upper Permian
beds was by Sedgwick (1829 : 76, 77, 120), who mentioned the discovery of impres-
sions of " ferns " in the Marl Slate of Middridge, S. of Bishop Auckland, during the
construction of the railway. This locality long continued to yield fossil plants, includ-
ing the best found in the Permian of this country. Lindley & Hutton (1837 : I23»
pi. 195) described only one Permian species, Voltzia phillipsii [= Ullmania frumen-
taria (Schloth.)], from Whitley, Northumberland. King (1850) included descriptions
and figures of the plant species Caulerpa ? selaginoides (Brongniart) [== Ullmannia
frumentaria], Neuropteris huttoniana, andPolysiphonia? sternbergiana, from Durham,
and of Chondrus binneyi (an inorganic structure) from red Permian marls near Man-
chester. This work also included a brief description of that important organism
(possibly an alga) of the Upper Magnesian Limestone, Filograna ? permiana, tenta-
tively referred to a worm genus.
For many years Kirkby and Howse were the most active students of Durham
Permian fossils. Among Kirkby's papers was an important one (1867) on the Marl
Slate and Lower Magnesian Limestone with records of several fossil plants, but a
more important list of Permian plants was included in Howse's (1890) catalogue of
the fossils in the Museum at Newcastle (now the Hancock Museum) . From the Marl
Slate were recorded, besides Ullmannia selaginoides, two other well-known German
Upper Permian species, U. bronni Goppert and Zonarites digitatus (Brongniart),
a specifically unidentified Calamites, and two supposedly new species given the
nomina nuda Taeniopteris duffiana and Ctenis permiana. From the Upper Magnesian
Limestone were recorded U. selaginoides and a supposedly new species of Calamites
which had been first mentioned by Kirkby (1864) and was assigned the nomen
nudum C. wakei. The algal species Chondrites virgatus Munster was recorded from
several horizons, but Howse seems to have confused under this name both King's
1 Abbreviated to 'B.M.N.H.' in the systematic descriptions.
2 The abbreviation ' G.S.M. ' is used for the Geological Survey Museum in the systematic
descriptions.
THE UPPER PERMIAN FLORA OF ENGLAND 297
Polysiphonia ? sternbergiana and his Filograna ? permiana. Since 1890 little attention
has been paid to the Permian plants of Durham, with the exception of Filograna ?
permiana, which has lately attracted notice as an index fossil of the Upper Magnesian
Limestone (Trechmann, 1925, 1942, 1952 ; Dunham, 1948).
The few records of plant remains from the Permian outcrop between Co. Durham
and Nottingham are mentioned on p. 299. Wilson (1876) .first recorded their
presence in beds exposed in the railway cuttings between Kimberley and Cinderhill,
and stated (1881 : 122) that Carruthers had pronounced them to " disclose a flora
known on the Continent ". Neither in these nor in later references (Gibson & others,
1908 : 104 ; Lamplugh & Gibson, 1910 : 67 ; Smith, 1913 : 215) to the presence of
fossil plants in the Kimberley Marl Slates is a single specific identification cited.
Borings in Nottinghamshire and Yorkshire passing through Permian rocks east
of the outcrop have brought to light a number of plant remains, hitherto almost
uninvestigated. An exception is a well-preserved cone-scale of Pseudovoltzia liebeana
(Geinitz), the structure of which was described in detail by Walton (1929).
It was not until 1862 that Harkness (p. 215) recorded the discovery of the now
well-known plant beds at Hilton Beck, near Appleby, Westmorland. As identified
by the palaeobotanist Heer, they included " ferns referable to Neuropteris and
Sphenopteris ", among them a species close to 5. erosa (Morris), of the Russian
Permian ; remains " allied to Weis sites (Goppert) ", as figured by Geinitz from the
German Zechstein ; a form " having the aspect of Caulerpites selaginoides (Sternb.) ";
and " detached leaves, resembling Cupressites Ullmanni Brongn." Two years later
Murchison & Harkness (1864 : 154) published a more satisfactory list of Hilton
plants, this time determined by Etheridge. The species mentioned are Sphenopteris
naumanni Gutbier, 5. dichotoma Althaus, Alethopteris goepperti Naum., Ullmannia
selaginoides, U. bronnii, Odontopteris ? sp., Sphenopteris ? sp., and Cardiocarpon
triangulare Gein. To these records Goodchild (1889) added Walchia piniformis
(Schloth.) and Noeggerathia cuneifolia Geinitz, but the first may be dismissed as
incorrect as the species in question belongs to the Lower Permian, while the second
may also be ignored as Geinitz himself had already withdrawn this species as based
on generically indeterminate remains. Brockbank (1892) described a visit to Hilton
and published three plates of unnamed and mostly indeterminate plant remains.
Later authors who have mentioned the Hilton plants have merely quoted Etheridge's
identifications.
Ill-preserved plant remains found by Roeder in the Upper Permian Manchester
Marls were submitted to Geinitz for identification (Geinitz, 1889, 1890 ; Roeder,
1890, 1890^, 18906). The species identified were Voltzia liebeana Geinitz, ? Ullmannia
selaginoides (Brongn.), and Filograna permiana King, together with two non- vegetable
structures (Guilielmites permianus and Spongillopsis dyadica] which Geinitz had
previously described from the German Permian and regarded as plant remains. In
the present paper doubt is cast on the reliability of the plant determinations on
account of the poorness of the material. Records of plant remains from borings in
Permian rocks in SW. Cumberland are referred to on p. 303.
It is uncertain if the Permian rocks found in Devon, Shropshire, Ayrshire, northern
Scotland, Ulster, and certain other areas include beds of Thuringian age, and so
far they have yielded no plant remains,
298 THE UPPER PERMIAN FLORA OF ENGLAND
3. NOTES ON THE PLANT LOCALITIES,
WITH SPECIES FOUND AT EACH
(a) Eastern Outcrop, Localities Enumerated from North to South
Culler coats Bay (Trechmann, 1931 : 247 ; Hickling & others, 1948 : 6). The Marl
Slate, with fish and plant remains, formerly cropped out on the foreshore 50 yards
from the tip of the promontory, but it has been almost entirely removed by collectors.
It has yielded :
Paracalamites kutorgai (Geinitz)
Annular ia ? sp.
Ullmannia frumentaria (Schlotheim)
Unidentifiable fossil wood, including large branches
Whitley. At this locality, near Cullercoats, was formerly a quarry exposing Yellow
Sands overlain by Marl Slate and Lower Magnesian Limestone. It yielded the type
specimen of Voltzia phillipsi L. & H. (= Ullmannia frumentaria).
Westoe, South Shields. Here was a quarry in Lower Magnesian Limestone mentioned
by Kirkby (18670 : 188). Bedding planes were covered with Algites sternbergianus
(King).
Fulwell Hill Quarries (Kirkby, 1864 : 345 ; Woolacott, 1912, section facing p. 260).
On the north side of Sunderland and famous for concretionary structures. Here
the Upper Magnesian Limestone yielded plant remains (now lost) consisting, according
to Kirkby, of a stem of Calamites, Ullmannia selaginoides (= U. frumentaria),
and a large, reed-like plant. These were a unique occurrence at this horizon.
Fulwell Water Works. A specimen of Paracalamites kutorgai (Geinitz) came from
this locality. There is no published record of the beds that were exposed there.
Claxheugh (Kirkby, 1867 : 197 ; Woolacott, 1898 : 14 ; 1903 : 211 ; 1912, figs.
3, 4, facing p. 256). A river bluff 2 miles west of Sunderland, now in a shipbuilding
yard and inaccessible. There was formerly an interesting section exposing Permian
beds ranging from Yellow Sands to Middle Limestone. Specimens of Ullmannia
frumentaria are from the Marl Slate.
Thrislington Gap (King, 1850 : xii). A railway cutting about i mile N. of Ferryhill
Station, exposing Coal Measure Sandstone, Marl Slate and Lower Magnesian Lime-
stone. The type locality of Mixoneura huttoniana (King).
Ferryhill (Calvert, 1884 : 73)- Railway cuttings near the one mentioned last. From
them come :
Ullmannia frumentaria (Schlotheim)
Pseudovoltzia liebeana (Geinitz)
Cornforth. Presumably from the old quarries near the village King recorded
" Caulerpa selaginoides " ( — Ullmannia frumentaria).
Raisby Hill Quarry (Hickling, 1931). Two miles E. of Cornforth and just north of
the railway. This large quarry, i mile long, affords the best section of the lower beds
of the Magnesian Limestone now to be seen in Durham. The Marl Slate, thickly
developed at the western end, yielded the large branch of Ullmannia bronni Goppert
represented in PI. 38, figs, la, b.
THE UPPER PERMIAN FLORA OF ENGLAND 299
Brusselton. A farm S. of Bishop Auckland with quarries near-by. Cited by King
as a locality for " Caulerpa selaginoides " and " Neuropteris " huttoniana.
East Thickley Quarry (Trechmann, 1921). Adjoining Shildon railway station and
exposing Coal Measure Sandstone, Marl Slate, and Lower Limestone. Recorded as
the type locality of Algites sternbergianus (King), which probably came from the
Lower Limestone. From the Marl Slate of this quarry come :
Ullmannia frumentaria (Schlotheim)
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Middridge (Sedgwick, 1829 : 76 ; Hancock & Howse, 1870 : 356 ; Trechmann,
1921 : 538). Quarries adjoining the railway E. of Shildon Station and S. of Middridge
village formerly showed Coal Measure Sandstone succeeded by Marl Slate and Lower
Limestone, but only unfossiliferous Lower Limestone is now visible. The Marl Slate
at this locality was the source of the best Permian plants yet found in England.
Species identified by the writer are :
Algites virgatus (Munster)
Taeniopteris eckhardti Kurtze
? Psygmophyllum cuneifolium (Kutorga)
Sphenobaiera digitata (Brongniart)
Pseudoctenis middridgensis sp. nov.
Ullmannia bronni Goppert
Ullmannia frumentaria (Schlotheim)
? Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Glass Houghton, NW. of Pontefract, Yorks. Wilson (1881 : 122) recorded plant
remains from beds underlying Lower Magnesian Limestone in sandpits at this locality.
No specimens have been traced.
Glapwell, Derbyshire. Five miles NW. of Mansfield, Notts. In the Geological Survey
Museum are obscure leaf impressions resembling Ullmannia bronni Goppert and other
remains said to have come from Permian beds here.
Fackley Hill, near Skegby, Notts. (Aveline, 1861 : 5 ; Lamplugh, in Gibson &
Wedd, 1913 : 80). The locality of leaf impressions here identified as cf. Cordaites
aequalis (Goppert), a unique record for this country. The specimens occur as impres-
sions in yellowish limestone.
Fulwood Top, near Sutton-in-Ashjield, Notts. About 2 miles due S. of the last
locality. Aveline (1861 : 6) recorded fossil wood from sandstones representing Lower
Magnesian Limestone here.
Kimberley, near Nottingham (Wilson, 1876 ; 1881, pi. 7 ; Smith, 1913, pi. 380 ;
Carr, 1913, pi. 146). Two parallel railway cuttings extending eastwards from Kimber-
ley Station for about a mile expose Upper Coal Measure Sandstone overlain by the
lower beds of the Magnesian Limestone series, with a breccia at the base. The plant-
bearing Marl Slates (about 30 ft. thick) have been exposed in the more northerly
cutting by a recent landslip, and the writer was able to collect from and make a
300 THE UPPER PERMIAN FLORA OF ENGLAND
detailed measurement of the section. The following plant remains have been identi-
fied from Kimberley :
Callipteris martinsi (Kurtze)
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Strobilites bronni Solms-Laubach
Samaropsis triangularis (Geinitz)
Cinderhill. A colliery 2 miles on the Nottingham side of Kimberley. A temporary
exposure here some years ago enabled Dr. H. S. Holden to collect fossil plants,
now in the British Museum (Natural History), from the Marl Slate. They include
isolated leaves of Ullmannia bronni Goppert and U '. frumentaria (Schlotheim).
(b) Borings and Colliery Shafts in S. Yorkshire, Lincolnshire and Nottinghamshire
Utteskelf Nurseries Boring (Edwards, Mitchell & Whitehead, 1950 : 81). About
8 miles NW. of Selby, Yorks. The Upper Magnesian Limestone yielded Tubulites
permianus (King) and structures resembling Chondrus binneyi King.
Wressell Boring (Edwards, 1951 : 263). About 7 miles E. of Selby, Yorks. At
depth 1,982 ft. the Lower Magnesian Limestone yielded a fragment of a shoot of
Pseudovoltzia liebeana (Geinitz).
Bentley Colliery No. 2 Shaft (Edwards, 1951 : 127). Plant-bearing specimens from
the Lower Marl, in the Wollaton Hall Museum, contain Ullmannia bronni Goppert,
an unidentified fructification, and fossil wood.
Sutton Boring (Edwards, 1951 : 247). Two miles NNE. of East Retford, Notts.
The Lower Marl, at depth 1,085 ft., is rich in plant remains, including the following :
Callipteris martinsi (Kurtze)
Ullmannia sp.
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
West Drayton Boring. Four miles S. of East Retford. Lower Limestone at depth
1,073 ft. yielded an indeterminate conifer fragment recorded as Pinnularia capillacea
L. & H. (Edwards, 1951 : 101).
Haughton Hall Boring. Five miles S. of East Retford. Lower Marl at depth in ft.
yielded well-preserved leaves of Hiltonia rivuli Stoneley, recorded as Cordaites sp.
by Edwards (1951 : 99).
Doddington Boring (Edwards, 1951 : 162). About 6 miles W. of Lincoln. Lower
Limestone or Lower Marl (unseparated in record of boring) yielded the following :
Callipteris martinsi (Kurtze), depth 2,320 ft.
Ullmannia bronni Goppert, depths 2,300, 2,303, 2,310 ft.
Ullmannia frumentaria (Schlotheim) ?, depth 2,327 ft.
Hiltonia rivuli Stoneley, depths 2,310, 2,330 ft.
Markham Moor Boring. About 6 miles S. of East Retford. Plant remains occurred
through a range of 248 ft. of Permian beds, as follows :
301
Piaea anglica sp. nov., depth 1,140 ft.
Piaea sp., depth 1,105 ft-
Calathella dictyonemoides sp. nov., depth 1,056 ft.
Ullmannia frumentaria (Schlotheim) , depth 1,055 ft.
Ullmannia sp., depths 1,046 ft., 1,047 ft-
Pseudovoltzia liebeana (Geinitz), depth 1,105 ft-
Hillonia rivuli Stoneley, depth 1,294 ft.
Thoresby Colliery Shaft. About 7 miles NE. of Mansfield, Notts. Ullmannia
frumentaria (Schlotheim), depth unrecorded.
Wellow Boring (Edwards, 1951 : 259). About 14 miles S. of East Retford. Ullman-
nia bronni Goppert from Lower Marl or Lower Limestone at an unrecorded depth.
Farnsfidd Boring (Edwards, 1951 : 167). About 9 miles W. of Newark. Cuticle
preparations made by Professor R. Florin from bulk-macerated material from Lower
Limestone yielded, besides several new cuticle types, the following :
Ullmannia sp.
Hiltonia rivuli Stoneley
Calverton Colliery Shaft. About 5 miles NNE. of Nottingham. Material labelled
" Marl Slates 134-152 yards " has yielded the following :
Callipteris martinsi (Kurtze)
Ullmannia bronni Goppert
Lowdham (or Cliff Mill] Boring (Edwards, 1951 : 201). About 7 miles ENE. of
Nottingham. Unidentifiable plant remains were found in Middle Marl at depth
688 ft. and in Lower Marl at depth 782 ft.
Bulcote Boring (Edwards, 1951 : 140). Six miles ENE. of Nottingham. Lower
Marl at depth 711 ft., i ft. above the Basal Breccia, yielded the interesting fossil,
Conites sp. (p. 329).
(c) Eden Valley (Westmorland and SE. Cumberland)
Hilton Beck. The chief exposure of Hilton Plant Beds is in a low cliff near the
southern bank of Hilton Beck at a spot called Ash Bank, about \ mile W. of Hilton
village and close to the road to Appleby. The strata are also exposed along the stream
course, and in the northern bank, just above the Penrith Sandstone, was found a
bed full of well-preserved leaves, study of which gave the clue to the existence of the
genus Hiltonia. The plant beds consist of sandstones, which frequently are current-
bedded and have ripple-marked surfaces, and of shales, often finely laminated and
highly micaceous. No fossils other than plant remains were found during several
days' collecting. The list from this locality is :
Callipteris martinsi (Kurtze)
Sphenopteris bipinnata (Minister)
Schutzia ? sp.
Sphenobaiera digitata (Brongniart)
Ullmannia bronni Goppert
302 THE UPPER PERMIAN FLORA OF ENGLAND
Ullmannia frumentaria (Schlotheim)
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Strobilites bronni Solms-Laubach
Strobilites ludwigi (Weigelt)
Murchison & Harkness (1864 : 154) recorded the seed Cardiocarpon (now Samar op-
sis} triangularis Geinitz from Hilton, but the specimen recorded (Geological Survey
Museum) is not that species and may not be a seed. The writer has collected small
cordate seeds at Hilton.
Belah Brook (Eccles, 1871 : 34). Plant beds occur in the section exposed along this
stream between Belah Bridge, Brough Sowerby, and Robridding. They consist of
grey micaceous shale rich in plant remains which, however, are even more ill-preserved
than those at Hilton. The following species are represented in material collected at
Belah Brook in August 1957 :
Callipteris martinsi (Kurtze)
Ullmannia bronni (Goppert)
Ullmannia frumentaria (Schlotheim)
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Strobilites bronni Solms-Laubach
Birkshead Gypsum Mine, Marlon Moor. Three and a half miles N. of Appleby.
Plant remains are associated with the gypsum and may be seen in the roof of the mine.
The following species were collected in August 1957 :
Callipteris martinsi (Kurtze)
Pseudovoltzia liebeana (Geinitz)
Hiltonia rivuli Stoneley
Opencast Gypsum Mine (Thos. M'Ghie & Sons), i mile E. of Kirkby Thore. Situated
4^ miles NNW. of Appleby. About 5 ft. of plant beds occur just above " gypsum
bed B " of Hollingworth, and have yielded Hiltonia rivuli Stoneley.
Acorn Bank Gypsum Pit (Russell, in Dakyns, Tiddeman & Goodchild, 1897 :
81). Situated NNE. of Temple Sowerby. Penrith Sandstone is overlain by 20 ft.
of massive gypsum, and this by shale with ill-preserved plants. From the gypsum
itself came the remarkable specimen of Callipteris martinsi (Kurtze) illustrated in
Text-fig. 5^ (p. 313).
Little Salkeld. Five and three-quarter miles NE. of Penrith. A few specimens
from Hilton Plant Beds at this locality are in the Carlisle Museum. They include
leaves of the following :
Ullmannia bronni Goppert
Ullmannia frumentaria (Schlotheim)
Hiltonia rivuli Stoneley
(d) Manchester District
Fallowfield (Roeder, 1892) . A railway cutting 3| miles S. of the centre of Manchester.
THE UPPER PERMIAN FLORA OF ENGLAND 303
From the red marl at this locality Geinitz (1889, 1890) identified Voltzia liebeana
Geinitz and Guilielmites permianus Geinitz, and, from limestone bands, Filograna
permiana King. Specimens in the Manchester Museum labelled as the first species
are now indeterminate, Guilielmites is an inorganic structure, and the identification
of the last species is uncertain. The best specimen seen from here is a red marl im-
pression of a small shoot with broad, linguiform leaves recalling those of Hiltonia
rivuli Stoneley.
(e) South-West Cumberland and Furness
Old Layriggs Borehole, Kirksanton (Smith, 1919 : 17). Obscure plant remains were
found in sandy shale between depths 227 ft. and 243 ft.
Haverigg Haws Borehole, near Kirksanton (Dunham & Rose, 1949 : 20). Dark
grey shales at depth 1,675 ft. overlying Lower Brockram yielded plant remains
recorded as Ullmannia bronni, but actually Callipteris martinsi (Kurtze).
Davy Street Boring, Barrow (Dunham & Rose, 1949 : 21). Plant remains, not
traced, occurred in grey shale between depths 2,786 ft. and 2,790 ft.
4. STRATIGRAPHICAL AND GEOGRAPHICAL DISTRIBUTION
OF THE PLANTS DESCRIBED
The following are lists of the fossil plant species from (a) the Marl Slate or approxi-
mately equivalent beds near the base of the Magnesian Limestone Series in eastern
England (either along the outcrop or in borings) from the south of Northumberland
to near Nottingham ; (6) higher Permian beds in the same area ; (c) the Hilton
Plant Beds. Plant remains from the Manchester Marls and from Furness and SW.
Cumberland are too scanty to be considered here.
Marl Slate or Approximately Equivalent Beds in Eastern Area
Algites virgatus (Munster)
Paracalamites kutorgai (Geinitz)
Annularia ? sp.
Callipteris martinsi (Kurtze)
Mixoneura sp. (huttoniana (King))
Taeniopteris eckardti Kurtze
? Psygmophyllum cuneifolium (Kutorga)
Conites sp.
Sphenobaiera digitata (Brongniart)
Cordaites aequalis (Goppert)
Pseudoctenis middridgensis sp. nov.
Ullmannia bronni Goppert
Ullmannia frumentaria (Schlotheim)
Hiltonia rivuli Stoneley
Pseudovoltzia liebeana (Geinitz)
Strobilites bronni Solms-Laubach
Samaropsis triangularis (Geinitz)
304 THE UPPER PERMIAN FLORA OF ENGLAND
Higher Beds in Eastern Area
Piaea anglica sp. nov. Horizon not yet certain.
Calathella dictyonemoides sp. nov. Ditto.
Algites sternbergianus (King). Lower Limestone.
Tubulites permianus (King). Upper Limestone.
Paracalamites kutorgai (Geinitz). Upper Limestone.
Ullmannia frumentaria (Schlotheim) . Upper Limestone.
Hilton Plant Beds
Callipteris martinsi (Kurtze)
Sphenopteris bipinnata (Miinster)
Schiitzia ? sp.
Sphenobaiera digitata (Brongniart)
Ullmannia bronni Goppert
Ullmannia frumentaria (Schlotheim)
Hiltonia rivuli Stoneley
Pseudovoltzia liebeana (Geinitz)
Strobilites bronni Solms-Laubach
Strobilites ludwigi (Weigelt)
5. FLORAL EVIDENCE ON CORRELATION OF
ENGLISH UPPER PERMIAN ROCKS
The marine Upper Permian beds of England were probably laid down in the same
sea as the German Zechstein. The folio wing are the subdivisions of the latter formation
in Thuringia :
Oberer Zechstein
Obere Letten
Plattendolomit
Untere Letten
Mittlerer Zechstein
Hauptdolomit
Unterer Zechstein
Zechsteinkalk
Kupferschiefer
Zechsteinkonglomerat
The palaeontological investigation of these deposits has been much more thorough
than in the case of the English Permian beds, but the number of fossil plant species
known to occur in them is not great. The majority have been found only in the
Kupferschiefer, but a few species range upwards. Fossil plants are not infrequent
in the Upper Zechstein, although apparently they are not common in the intermediate
beds. The following is a list of species which have been described from the Kupfer-
schiefer, omitting synonyms and Weigelt's two species of the unsatisfactory genus
Archaeopodocarpus, which appear to have been founded mainly on specimens of
Ullmannia frumentaria.
THE UPPER PERMIAN FLORA OF ENGLAND 305
List of Fossil Plants of the German Kupferschiefer
Algites virgatus (Minister)
Paracalamites kutorgai (Geinitz)
Sphenopteris geinitzi Goppert
Sphenopteris bipinnata (Miinster)
Sphenopteris kukukiana Gothan & Nagalhard
Sphenopteris gibbelsi Gothan & Nagalhard
Sphenopteris gothani Weigelt
Sphenopteris densifolia Weigelt
Sphenopteris ropkei Weigelt
Sphenopteris gillitzeri Weigelt
Odontopteris goepperti Weiss
Callipteris martinsi (Kurtze)
Taeniopteris eckardti Kurtze
Sphenobaiera digitata (Brongniart)
Cordaites pangerti Weigelt
Ullmannia bronni Goppert
Ullmannia frumentaria (Schlotheim)
Ullmannia orobiformis (Schlotheim) (doubtful species)
Pseudovoltzia liebeana (Geinitz)
Strobilites bronni Solms-Laubach
Strobilites ludwigi (Weigelt)
Strobilites major Weigelt
Strobilites elongatus Weigelt
Strobilites minor Weigelt
Strobilites dentatus Weigeit
Samaropsis triangularis (Geinitz)
Cyclocarpon eiselianum Geinitz
Carpolithes rotherianus Geinitz
The following is a list of fossil plants recorded from the Upper Zechstein. The
chief localities are in Hesse (including the deposits at Frankenberg from which
Ullmannia bronni was first described) and Saxony (Schuster, 1933).
List of Fossil Plants of the Upper Zechstein
Sphaerococcites dyadiccus Sterzel
Piaea punctata Florin. Obere Letten.
Piaea gigantea Florin. Obere Letten.
Calathella krduseli Florin. Obere Letten.
Algites virgatus (Miinster)
Tubulites permianus (King). Plattendolomit.
Sphenopteris sterzeli Schuster. Plattendolomit.
Sphenopteris sp. I, Schuster. Obere Letten.
Callipteris martinsi (Kurtze)
306 THE UPPER PERMIAN FLORA OF ENGLAND
Taeniopteris eckardti Kurtze. Untere Letten.
Ullmannia bronni Goppert. Untere Letten and Plattendolomit.
Ullmannia frumentaria (Schlotheim). Untere Letten, Plattendolomit and
Obere Letten.
Pseudovollzia liebeana (Geinitz). Plattendolomit and Obere Letten.
Strobilites bronni Solms-Laubach
Cyclocarpus spongioides Geinitz
Rhabdocarpus klockeanus Geinitz. Plattendolomit.
The possibility must not be overlooked that deposition of the basal member of
the Upper Permian series in the eastern counties of England may have been diachro-
nous, as Sherlock and others have suggested. The view is here taken, however, that
any time that may have elapsed between the incoming of marine conditions at
different localities was not great in comparison with the duration of Upper Permian
time, so that species found at all the localities may be considered together. The great
majority of the plants in the first of the foregoing lists have, moreover, been found
in the Marl Slate at the same Durham locality, Middridge.
It has always been considered that the Marl Slate should be correlated with the
German Kupferschiefer. Their stratigraphical positions are similar, separated as
they are from the base of the Upper Permian succession only by a basal breccia or
by sands, and they are similarly rich in fossil fishes and plant remains. The flora
of the Marl Slate supports this conclusion, for several species are common to this
formation and to the Kupferschiefer. The stratigraphical importance of some of these
species is, however, limited, for the ranges of Callipteris martinsi, Taeniopteris
eckardti, Ullmannia bronni, U. frumentaria and Pseudovoltzia liebeana extend to
the Upper Zechstein, while Paracalamites kutorgai, although not recorded from
the Upper Zechstein in Germany, occurs in England in the Upper Magnesian Lime-
stone, if (as is probable) an untraced " Calamites " recorded by Kirkby belonged
to this species. Sphenobaiera digitata and the seed Samaropsis triangularis, both
common to the Kupferschiefer and the Marl Slate, have not, however, been found
in the Upper Zechstein.
Of great interest, however, is the occurrence in the Marl Slate of certain species
not known from the Kupferschiefer or any other horizon in Germany. Two of these,
Hiltonia rivuli (abundant in England) and Pseudoctenis middridgensis, are new,
while Mixoneura huttoniana, known only by isolated leaves, was described long ago
by King (1850). An important discovery is that of specimens possibly belonging
to Psygmophyllum cuneifolium, previously known only from the Permian of the
Urals. A leaf which seems to belong to the well-known species Cordaites aequalis,
from Central Asia, Pechoraland, and Siberia, and barely distinguishable from the
widespread Gondwana species Noeggerathiopsis hishpi, comes from near the base of
the Upper Permian at a locality in Nottinghamshire which deserves further investi-
gation.
The re-appearance of Ullmannia in the Upper Magnesian Limestone of Durham is
in agreement with our knowledge of its range in the German Zechstein. The most
important fossil plants found at horizons above the Marl Slate in the English beds
are, however, the algae. Tubulites permianus (hitherto called Filograna permiana
THE UPPER PERMIAN FLORA OF ENGLAND 307
in England) is an index fossil of the Upper Magnesian Limestone throughout Durham,
Yorkshire and Nottinghamshire, and occurs in similar abundance in Germany in the
Plattendolomit, confirming the correlation of the English and German beds. In
Germany the algal genera Piaea and Calathella have been found only in the Obere
Letten of the Upper Zechstein. The beds in the Nottinghamshire borehole in which
they occur, although not assigned to their exact stratigraphical positions, are well
above the base of the Upper Permian.
Earlier workers considered that the Hilton Plant Beds should be correlated with the
Marl Slate and with the Kupferschiefer, but Trotter & Hollingworth (1932 : 124)
put forward the view that they are the equivalent of the Middle Permian Marl of
Durham. This conclusion was based on a comparison of the lithological succession
in the two areas, the fossil plants being considered useless for correlation. While
several plant species range throughout the Zechstein, three found at Hilton (Spheno-
pteris Hpinnata, Sphenobaiera digitata and Strobilites ludwigi) have been recorded
only from the Kupferschiefer in Germany, and so support the earlier view as to the
age of the Hilton beds. Hiltonia rivuli, moreover, is common to the Hilton beds and
the Marl Slate and is not known from later beds east of the Pennines.
6. SYSTEMATIC DESCRIPTIONS
THALLOPHYTA
ALGAE
Genus PIAEA Florin, 1929
Piaea anglica sp. nov.
(PL 36, fig. 3)
DIAGNOSIS. A Piaea with axes unbranched in their upper parts ; axes about i
mm. wide, bearing whorls of filaments at intervals of about 0-25 mm.; filaments
about 10 in a whorl, up to 2 mm. long, and mostly arising at an angle of about
60°.
OCCURRENCE AND MATERIAL. Magnesian Limestone Series, Markham Moor
borehole, Notts, (p. 300) ; types (G.S.M. no. 76607) from depth 1,140 ft.; a single
specimen from depth 1,105 ft.
DESCRIPTION. There are about 30 axes lying mostly parallel on a bedding plane
in a core ; those shown in the figure are typical. The length of the longest fragment
is 5 cm. None is branched ; the lower parts, however, are not seen, being broken
or outside the core. Two axes show in places transverse rows of dots thought to
have been bases of the lateral filaments ; the number (about 4) visible in each row
suggests that there were about 10 in a complete whorl. The substance is carbonaceous
and there is no evidence of original calcification.
COMPARISON. Professor R. Krausel kindly sent me the original specimens of
Piaea punctata Florin (1929) from the Zechstein. Though very similar to the English
specimens, they have narrower axes, shorter filaments and more conspicuous filament
scars, and they branch freely.
308 THE UPPER PERMIAN FLORA OF ENGLAND
Genus CALATHELLA Florin, 1929
Calathella dictyonemoides sp. nov.
(PL 36, fig. 4 ; Text-figs, i, 2)
DIAGNOSIS. A Calathella with longitudinal strands, 0-2-0-3 mm. wide, connected
by transverse strands of the same width to form meshes mostly rectangular and about
0-5 mm. long and 0-2-0-3 mm. wide.
OCCURRENCE AND MATERIAL. Magnesian Limestone Series, Markham Moor bore-
hole, Notts, (p. 300) ; core with several specimens (G.S.M. no. 76608) from depth
1,056 ft.
DESCRIPTION. The figures show all that can be seen of two typical specimens.
With them is associated a second type of structure (Text-fig. 2) which may be a dif-
ferent part of the same plant. It consists of a group of axes diverging fan- wise from
FIG. i. Calathella dictyonemoides sp. nov. Magnesian Limestone Series ; depth 1,056 ft.,
Markham Moor borehole. Geological Survey, no. 76608. x 5.
a point. These axes vary in width, the majority between o-i and 0-2 mm., and
some are narrowest near the base. They are often curved and occasionally fork,
but are not joined by cross-connections.
COMPARISON. The original specimens of C. krduseli Florin from the Zechstein bear
at their apices what seem to be basket-shaped bodies composed of anastomosing
raKpH ,i €tt&8£*>*\
%^ r^test^
HS%
FIG. 2. Calathella dictyonemoides sp. nov. Rectangular mesh (at top) and associated
structures. Magnesian Limestone Series ; depth 1,056 ft., Markham Moor borehole.
Geological Survey, no. 76608. x 2-5.
strands. These bodies somewhat resemble the English specimens but have narrower
strands. At the apex of one specimen of C. krduseli is a bunch of narrow, irregularly
spreading filaments like those arising from the lower parts of axes in that species. The
THE UPPER PERMIAN FLORA OF ENGLAND 309
brush-like structures in the material now described may similarly have been terminal
on some axes.
The new specific name recalls the graptolite Dictyonema for which the present
specimens were originally mistaken.
Genus ALGITES Seward, 1894
Algites virgatus (Minister)
(Text-fig. 3)
1842 Chondrites virgatus Minister, p. 102, pi. 15, fig. 18.
1862 Chondrites virgatus Mtinst.: Geinitz, p. 132, pi. 24, fig. 5.
ENGLISH OCCURRENCE AND MATERIAL. Marl Slate, Middridge, Durham ; one
specimen (Hancock Museum).
DESCRIPTION. The figure shows all the features that can be seen. The distal ends
of some axes seem to bear plume-like structures, but no details are visible.
COMPARISON. The holotype from the Kupferschiefer is in the Palaeontological
FIG. 3. Algites virgatus (Miinster). Marl Slate, Middridge. Hancock Museum. X 2.
Department of the British Museum (no. 46674). It much resembles the English
specimen, and the axes, some of which seem to branch, bear some sort of rounded
body at the apex ; these bodies are, however, smaller than the " plumes " of the
English specimen.
It is clear from the recent paper by Simpson (1957) that the generic name Chondrites
GEOL. 3, 9.
28
310 THE UPPER PERMIAN FLORA OF ENGLAND
Steinberg should not be used for true algal structures. It is here replaced by Algites
Seward.
Algites sternbergianus (King)
(PL 36, figs, i, 2, 5)
1850 Polysiphonia (?) Sternbergiana King, p. 3, pi. i, fig. 2.
1 867*3 Chondrites virgatus Munster : Kirkby, pp. 192, 197.
1890 Chondrites virgatus Munster : Howse, pp. 244, 248.
OCCURRENCE AND MATERIAL. Lower Magnesian Limestone of Westoe, South
Shields ; numerous specimens in the Hancock, Geological Survey, and Manchester
Museums. Also recorded (as C. virgatus) from Claxheugh. The holotype (not traced)
was stated by King to be from the Marl Slate of Thickley, but was probably from the
Lower Magnesian Limestone.
DESCRIPTION. The specimens are axes up to i mm. wide and 8 cm. long, straight
or curved but not branched. Some are naked but others are clothed with irregular
fine filaments up to i cm. long, and some bear a brush of these filaments at their
distal end. No scars of filaments can be seen on the naked axes.
REMARKS. These specimens are too imperfectly characterized to be placed in a
more definite genus such as Calathella. It is to be noted that Geinitz (1858 : 6 ;
1862 : 132) thought this species inseparable from A. virgatus, and in this he was
followed by Kirkby, Howse, and others. To me they seem different.
Genus TUBULITES Bein, 1932
Tubulites permianus (King)
(PI. 36, figs. 6-9)
1850 Filograna (?) Permiana King, p. 56.
1 86 1 Filograna Permiana King : Geinitz, p. 41.
1932 " Stabchenkalk ", Prager, p. 358, pi. 9, figs. 1-3.
1932 Tubulites articulatus Bein, p. 798, pi. 27, figs. 3, 4.
1934 Tubulites articulatus Bein : Naumann, p. 194, pi. i$a, fig. i.
1942 Filograna (?) permiana King : Trechmann, p. 317.
1948 Filograna permiana King : Dunham, p. 220, pi. 10, fig. 3.
1952 Filograna permiana King : Trechmann, p. 307.
OCCURRENCE AND MATERIAL. Upper Magnesian Limestone throughout Durham
and Yorkshire. Localities of specimens studied are Blackball Rocks, 5 miles N. of
Hartlepool (material from here includes King's types, University College, Galway,
nos. 130, 131) ; shore at Easington, Durham (B.M.N.H., C. T. Trechmann Colin.) ;
Low Park Farm, Yorks. ; east of Hepworth Wood, Yorks. ; near Toulson Lodge,
Yorks.; and near Cross Newton, Doncaster (last four, G.S.M.). Borehole material
is from depth 4,240 ft., Eskdale, near Whitby (G.S.M.) ; also recorded from depth
638 ft., Sutton borehole, Notts. (Edwards, 1951 : 102). Very doubtful specimens
(Manchester Mus. nos. L.7o6, L. 12336) from the Manchester Marls of Strangeways
and Fallowfield, near Manchester, have also been examined ; they were the basis of
records by Geinitz (1890 : 548) and Roeder (1892 : 15).
THE UPPER PERMIAN FLORA OF ENGLAND 311
DESCRIPTION. The broken calcareous tubes from Easington shown in PL 36, fig. 7,
are typical of this abundant organism ; they are 0-5 mm. wide and up to 7 mm. long.
No carbon is present in them and there is no fine sculpture of any kind. Specimens
were examined in thin section, but the original structure was not clear on account
of metasomatic changes. The calcareous wall of the tube is usually 30-40 ju, thick
and in one case an inner tube 100 fi thick was observed. In no instance are the tubes
arranged like strings of beads. Specimens from the Eskdale borehole are a little
wider (0-75 mm.). The length of the longest single tube seen from any locality was
about 2 cm.
Some specimens from Blackball Rocks (PL 36, fig. 6) are only 0-3 mm. wide and may
occur in bundles, sometimes as many as 24 side by side ; one bundle of 5 can be
traced for 2 cm. There is a suggestion that certain of these groups may arise by
the branching of one tube. It is not certain that all the specimens mentioned are
specifically identical.
DISCUSSION. King gave no figure of this organism, which he referred with doubt
to a genus of living worms, Filograna, and his description was too brief to distinguish
it. His type specimens (kindly lent to me by Professor J. Mitchell) are too eroded to
be worth figuring here, but show that the species has been correctly identified by
modern authors. Kirkby (1861 : 309), however, who gave a description of this
organism, seems not to have known of King's name for it, while Howse (1890),
in a list, doubtfully placed Filograna permiana as a synonym of Chondrites virgatus.
Trechmann (1925), who revived the name F. permiana, called attention to the impor-
tance of the organism as a characteristic fossil of the Upper Magnesian Limestone,
and considered it an alga because he occasionally saw one tube branching out of
another. None of the specimens I have seen, however, shows this feature unmistak-
ably.
In 1932 Prager gave the name " Stabchenkalk " to an Upper Zechstein limestone
full of tubes i mm. wide and up to 8 mm. long. In the same year Bein gave reasons
for considering them algal and assigned to them the new name Tubulites articulatus
in ignorance of the earlier name for English specimens. Bein (1932) and Naumann
(1934) noted the frequent presence of an inner tube inside the outer one.
PTERIDOPHYTA
EQUISETALES
Genus PARACALAMITES Zalessky (1927) 1932
This generic name was first published in Zalessky's Atlas (1927) without a diagnosis.
The species there figured were Paracalamites decor atus (Eichwald), P. kutorgae
(Geinitz), and P. striatus (Schmalhausen), all of which had been originally described
under Catamites . The name was subsequently published (Zalessky, 1932 : 112) in
a paper on Upper Palaeozoic plants from Siberia, and the new species Paracalamites
sibiricus was described at the same time. Even in this paper no formal generic
diagnosis was given, but a few remarks were made on the reasons for founding a
new genus, and seem just sufficient to validate it with P. sibiricus as the type species.
It was explained that the casts of the pith-cavities of Calamites-like stems and
312 THE UPPER PERMIAN FLORA OF ENGLAND
rhizomes found in the Permian have the ridges of successive internodal segments in
alignment instead of alternating in position and interdigitating at the nodes, as in
true Catamites. In this respect they resemble the Lower Carboniferous genus Aster o-
calamites. Knowledge of their foliage, Zalessky thought, might eventually establish
their identity with either Phyllotheca, Schizoneura, or Lobatannularia, but for the
time being it seemed advisable to refer them to a new genus, Paracalamites. The
foliage of one of these Permian species has, however, been described by Weigelt
(1928 : 470, pi. 10, fig. i), who founded a new species Asterocalamites mansfeldicus
on specimens from the German Kupferschiefer. The characters of Paracalamites,
in so far as they are known, may now be summarized as follows.
Stems and the casts of their pith-cavities with numerous narrow longitudinal
ridges which are in alignment along successive internodal segments. Leaves lanceolate,
not joined, arranged as star-shaped verticils on slender foliage shoots which arise in
pairs and opposed at each node, and are directed upwards at an angle of about 45° ;
number of leaves in each verticil small.
The genus resembles Asterocalamites in the alignment of the internodal ridges,
but differs in the form of its leaves, which are not filiform and repeatedly forked.
Paracalamites kutorgai (Geinitz) Zalessky
(PL 37, fig- i)
1880 Catamites Kutorgai Geinitz, p. 14, pi. 6, fig. i.
1887 Catamites Kutorgae Geinitz : Schmalhausen, p. 33, pi. i, figs. 7-17.
1927 Paracalamites Kutorgae (Geinitz) Zalessky, p. 36, pi. i, fig. I ; pi. 44, figs. 3, 7.
1928 Asterocalamites mansfeldicus Weigelt, p. 470, pi. 10, figs. 1-4, 7-14.
1930 Asterophyllites (Neocalamites) mansfeldicus (Weigelt) Weigelt, p. 649.
OCCURRENCE AND MATERIAL. Marl Slate of Fulwell Water Works, near Sunderland
(one specimen) and of Cullercoats (two specimens), Hancock Museum.
DESCRIPTION. All three specimens are flattened stem fragments in shale. One
specimen is 25 cm. long and 17 cm. wide and shows three nodes with internodes
45 mm. long. One node is shown enlarged in PL 37, fig. i. The ribs on one internode
are in alignment with those on the next. Another specimen is 25 mm. wide and shows
internodes about 50 mm., 25 mm., 35 mm., and 25 mm. in length.
DISCUSSION. Stems such as these agree with those of Lower Carboniferous Astero-
calamites, but they match specimens from the German Kupferschiefer identifiable as
Paracalamites kutorgai. It may be noted that Kirkby (1864) recorded " Calamites
arenaceus ?" from the Upper Magnesian Limestone of Fulwell Hill, and that the
specimen was catalogued by Howse (1890) under the name Calamites wakei. The
specimen, which has not been traced, may have belonged to P. kutorgai.
Genus ANNULARIA Sternberg, 1821
Annularia ? sp.
(Text-fig. 4)
OCCURRENCE AND MATERIAL. Marl Slate of Cullercoats, two specimens (Hancock
Museum).
THE UPPER PERMIAN FLORA OF ENGLAND
313
Description. Each specimen shows a whorl of what are taken to be n very small
leaves, not more than 3 mm. long, round a circular space representing the node. No
FIG. 4. Annularia ? sp. Marl Slate, Cullercoats. Hancock Museum, x 2.
midrib is, however, visible in these " leaves ". They occur in the same bed as Para-
calamites kutorgai but differ in number and shape from the leaves of that species as
described by Weigelt (1928).
PTERIDOSPERMAE
Genus CALLIPTERIS Brongniart, 1849
Callipteris martinsi (Kurtze) Zeiller
(PI. 37, figs. 2, 5 ; Text-figs. 5, 6)
Alethopterin (sic) Martinsii Germar MS.: Kurtze, p. 34, pi. 3, fig. 2.
Callipteris Martinsi (Germar) : Gothan, pp. 1-4, figs. 1,2. (See this for full synonymy.)
Callipteris Martinsi (Germar) : Weigelt, p. 457, pi. 2, figs. 14, 15, 26 ; pi. 7 ; pi. 35, figs.
7, 14, 21, 22, 26.
OCCURRENCE AND MATERIAL. Marl Slate of Kimberley, Watnall, and Cinderhill,
near Nottingham ; several specimens (B.M.N.H.). Lower Marl of Magnesian Lime-
stone Series, Doddington borehole (p. 300), depth 2,320 ft.; fragment (G.S.M.). Hilton
1839
1907
1928
FIG. 5. Callipteris martinsi (Kurtze) Zeiller. (a), Marl Slate, Kimberley. British Museum
(Natural History), no. ¥.59630 ; (b), Marl Slate, Kimberley. British Museum (Natural
History), no. ¥.59636 ; (c), Hilton Plant Beds, Hilton. British Museum (Natural
History), no. ¥.5975 ; (d), Main gypsum horizon, Acorn Bank quarry, near Temple
Sowerby. Private collection, All x i.
3i4 THE UPPER PERMIAN FLORA OF ENGLAND
Plant Beds, Hilton and Belah Brook ; several small shoots (B.M.N.H. and G.S.M.).
Gypsum of Acorn Bank quarry, near Temple Sowerby (Text-fig. 5^, private coll.)
and of Birkshead Mine (B.M.N.H.). Kirksanton Beds, Haverigg Haws borehole
(p. 303); one fragment (G.S.M. no. TW 3846). Fragments were also identified in
bulk-macerated cores from the borings at Sutton (p. 300), depth 1,085 ft-. Lowdham
(p. 301), depth 688 ft., and Farnsfield (p. 301) ; also from the Calverton Colliery shaft
(p. 301).
DESCRIPTION. The best English specimens are from Kimberley ; the largest is
shown in Text-fig. 5«. The rachis has transverse ridges which may represent the
pinnules known to be borne directly on it in German specimens, and the pinna
FIG. 6. Callipteris martinsi (Kurtze) Zeiller. Cuticle, (a), Marl Slate, Cinderhill. British
Museum (Natural History), no. ¥.26971. x no ; (b), Marl Slate, Kimberley. British
Museum (Natural History), no. ¥.351 14. x 150 ; (c), Hilton Plant Beds, Hilton. British
Museum (Natural History), no. ¥.35115. x 270.
rachises have similar but smaller ridges. Another Kimberley specimen (PI. 37, fig. 5)
has scars of unknown origin at some pinna bases ; a previous worker who labelled
this specimen regarded these as seed scars, but they may be the remains of pinnules ;
seeds of C. martinsi are unknown. Text-fig. 5& represents a specimen from the same
locality in which the pinnules are lobed and relatively large. A few specimens from
THE UPPER PERMIAN FLORA OF ENGLAND 315
this locality show a midrib and lateral veins. Small fragments with deeply lobed
pinnules (Text-fig. 5c) are common at Hilton Beck and were recorded by Murchison
& Harkness (1864) as Alethopteris goepperti (Munster), a synonym of C. martinsi.
The specimen shown in Text-fig. $d is remarkable for its preservation in gypsum.
CUTICLE. Preparations were made from several specimens from Kimberley,
Watnall, Cinderhill, and Hilton Beck. Text-fig. 6 and PI. 37, fig. 2, show the cuticular
structure. The ordinary epidermal cells of both cuticles are polygonal and do not
show a median papilla, but the subsidiary cells often have a prominent papilla over
the stomatal pit. The subsidiary cells number either five or six, the latter number
appearing to be more usual on deeply lobed pinnules. The stomata are rare on one
face of the leaf and frequent on the other.
REMARKS. Although there are slight differences in their form, I consider that the
numerous English fragments all belong to one species, as they have the same general
type of cuticular structure. All of these forms can be matched (macroscopically)
with German specimens included in C. martinsi, and they are, therefore, identified
with that species. It is, however, surprising that there is a striking difference between
their cuticle and that of an Upper Zechstein specimen figured by Gothan & Nagalhard
(1922) and of another from the same locality, Frankenberg, figured by Florin (1931),
which I examined in Stockholm ; for in the German specimens the epidermal cells
have a clearly marked dark spot, representing a thickening of the cuticle. It may be
that the presence of a papilla on epidermal cells is inconstant in C. martinsi, but it
is also possible that some forms from the Zechstein once thought distinct species
but now included in C. martinsi are, after all, distinct. Hence, the English specimens
cannot be satisfactorily identified until the cuticles of many more German specimens
have been studied.
Genus SPHENOPTERIS Brongniart, 1822
Sphenopteris bipinnata (Munster) Geinitz
(PI. 37, fig. 6 ; Text-fig. 7)
1842 Caulerpites bipinnatus Miinster, p. 102, pi. 14, fig. 3.
1846 Sphenopteris dichotoma Althaus, p. 30, pi. 4, fig. i.
1928 Sphenopteris dichotoma Althaus : Weigelt, p. 448, pi. 8, figs. 3, 4.
FIG. 7. Sphenopteris bipinnata (Munster) Geinitz. Hilton Plant Beds, Hilton, British
Museum (Natural History), no. ¥.35117. x 1-5.
316 THE UPPER PERMIAN FLORA OF ENGLAND
OCCURRENCE AND MATERIAL. Hilton Plant Beds, Hilton Beck ; several fragments
(B.M.N.H. and G.S.M.).
DESCRIPTION. In some of the fragments the leaves show their veins excellently,
but in others the form alone is visible as the veins are not preserved in the coaly
substance. Some small fragments resemble S. patens (Althaus), which Geinitz
(1848) and Schenk (1864) but not some later authors have considered synonymous
with S. bipinnata. No cuticle is preserved in the specimens examined.
Genus MIXONEURA Weiss, 1869
Mixoneura sp.
(PL 37, ng. 4)
1850 Neuropteris Huttoniana King, p. 6, pi. i, fig. 4.
OCCURRENCE AND MATERIAL. Marl Slate of Thrislington Gap, Durham. The type is
preserved in University College, Galway. It is the only specimen seen, although
King also recorded the species from Middridge, Brusselton, and East Thickley.
REMARKS. The type specimen shows four pinnules on a single small piece of rock.
This may have been broken and re-assembled and it is impossible to say how close
they were originally. One pinnule is shown in PL 37, fig. 4 ; others seem to have been
slightly broader or slightly longer. No further details are visible.
These fragments much resemble Mixoneura neuropteroides (Goppert), from the
Lower Permian of Germany ; it is interesting that no Mixoneura is known to occur
in the German Upper Permian.
Genus TAENIOPTERIS Brongniart, 1828
Taeniopteris eckardti Kurtze
(PL 37, figs. 3«, b)
1839 Taeniopteris Eckardti Germar MS., Kurtze, p. 34, pi. 3, fig. i.
1851 Taeniopteris Eckhardi Germar : Ettingshausen, p. 99, pi. 13, figs. 2, 3.
1890 Taeniopteris duffiana Howse, p. 248 (nom. nud.).
1928 Taeniopteris Eckardti Germar : Weigelt, p. 461, pi. 2, fig. 21 ; pi. 9, figs. 7-18.
1930 Taeniopteris eckardti Germar : Weigelt, p. 647, figs. 13-16.
1933 Taeniopteris eckardti Germar : Schuster, p. 85, fig. 5 ; pi. 7, fig. 8.
OCCURRENCE AND MATERIAL. Marl Slate of Middridge, Durham ; one specimen
(Hancock Museum).
DESCRIPTION. The figures show the only English specimen, catalogued by Howse as
T. duffiana. Its veins, clearly seen, are all parallel, forming an angle of 60° with the
midrib, and number 8-9 to the centimetre. Its cuticle is not preserved. It agrees
with T. eckardti from the Upper Permian of Germany.
Genus PSYGMOPHYLLUM Schimper, 1870
? Psygmophyllum cuneifolium (Kutorga) Schimper
(PL 40, figs. 14, 15 ; Text-figs. 8, 9)
1838 Sphenopteris cuneifolia Kutorga, p. 32, pi. 7, fig. 3.
1927 Psygmophyllum cuneifolium (Kutorga) : Zalessky, pi. 8, figs. 3, 4 ; pi. 9, figs. 2, 3 ; pi.
12, fig. i ; pi. 14, fig. 5 ; pi. 16, fig. 6 ; pi. 32, fig. 6 ; pi. 33, fig. 8 ; pi. 37, fig. 6.
1927 Sphenopteris incerta (Fischer) : Zalessky, p. 40, pi. n, fig. 2.
1937 Psygmophyllum cuneifolium (Kutorga) : Zalessky, p. 64, text-fig. 27.
THE UPPER PERMIAN FLORA OF ENGLAND
317
OCCURRENCE AND MATERIAL. Marl Slate of Middridge, two specimens (B.M.N.H.).
DESCRIPTION. Satisfactory photographic illustration of the specimens is not possible
FIG. 8. ? Psygmophyllum cuneifolium (Kutorga) Schimper. Some remaining irregular
patches of leaf substance are not indicated. Marl Slate, Middridge. British Museum
(Natural History), nos. V. 35 131, ¥.35132. X i.
as they are dirt-stained and the leaf substance, once complete, has peeled off in places.
The form of the leaves, as shown in Text-figs. 8a, b, can, however, be made out. One
318 THE UPPER PERMIAN FLORA OF ENGLAND
clearly shows forking of the leaf below and perhaps the beginning of a repeated forking
at the apex. The other specimen indicates probable forking below (where broken
away). Where the cuticle has disappeared the veins are visible as parallel longi-
tudinal strands on the impressions of both rachis and pinnae.
CUTICLE. The preparations were fragile and broke into small pieces. It was impos-
sible to distinguish the two sides, but different kinds of epidermis were recognized
along the veins and between them. Along strips assumed to be the veins the cells
mostly form elongated polygons. Between the veins they are either isodiametric
polygons, or polygons elongated in various directions often with one or more walls
curved. At intervals on both types of epidermis are round scars most of which
probably represent unicellular trichome bases although some are stomata. Six or
more subsidiary cells form an imperfect ring round a circular pore in which the guard
cell aperture is occasionally visible.
DISCUSSION. These specimens resemble the one from the same locality (but appar-
ently not from the same bed) described below (p. 321) as Pseudoctenis middridgensis.
Unfortunately comparison of the cuticular structure is impossible as only very poor
preparations are obtainable from that specimen. Although it is not impossible that
future discoveries may prove their specific identity, the following points suggest
FIG. 9. ? Psygmophyllum cuneifolium (Kutorga) Schimper. Cuticle. Marl Slate,
Middridge, British Museum (Natural History) ; (a), no. V. 35 133 ; (b), no. V. 35 134(3 ; (c),
no. V.35I340. All x 150.
that the two forms are distinct. In P. middridgensis the rachis is not known to fork ;
the pinnae are mostly narrower at the base and taper to a pointed apex ; the veins
are more delicate ; and the cuticle appears to have been thinner. Most of the pinnae
in the present specimens are incomplete owing to fracture of the rock, and their
apparent truncation elsewhere may in some cases be due to breakage before fossiliza-
tion. There is, however, no indication that a single one tapered to a point, and in
one case the impression of what appears to have been the undamaged apex can just
be seen to be bluntly rounded.
These specimens much resemble a figure of Psygmophyllum cuneifoliiim published
THE UPPER PERMIAN FLORA OF ENGLAND 319
by Zalessky (1937) and are therefore referred with a query to that species. Previously
described specimens are from the Permian of Russia, where the species is well repre-
sented although the cuticle has not been described. It is questionable whether
Psygmophyllum is the best generic name for the species (cf. Arber, 1912), and it
might be transferred to Zalessky's genus Bardia.
Genus SCHUTZIA Geinitz, 1863
Schutzia ? sp.
(PL 37, figs. 7-12)
OCCURRENCE AND MATERIAL. Hilton Plant Beds of Hilton ; several specimens,
B.M.N.H. and G.S.M.
REMARKS. The specimens illustrated look like the cup-like sporangial aggregates
of Schutzia anomala figured from the Lower Permian by Geinitz (1863), Goppert
(1864), Schuster (1911) and Gothan (1937). It is not, however, possible to prove
that they are of this nature. Compression has distorted many of them in various
ways. Gothan thought that S. anomala belonged to the species Sphenopteris germanica
Weiss, and the present specimens could belong to S. bipinnata, with which some were
found associated.
Genus SPHENOBAIERA Florin, 1936
Sphenobaiera digitata (Brongniart) Florin
(PI. 38, fig. 6 ; Text-figs. 10, n)
1828 F-ucoides digitatus Brongniart, p. 69, pi. 9, fig. i.
1862 Zonarites digitatus (Brongn.) Geinitz, p. 336, pi. 26, figs. 1-3.
1876 Baiera digitata (Brongn.) Heer, p. 7, pi. 21, figs. 1,2.
1880 Schizopteris digitata (Brongn.) Geinitz, p. 16, pi. 6, figs. 13, 14.
1928 Baiera digitata (Brongn.) : Weigelt, p. 476, pi. n, figs. 1-19 ; pi. 12, figs. 1-16.
1932 Baiera digitata (Brongn.) : Weigelt, p. 148, pi. 4, figs. 1-3.
1936 Sphenobaiera digitata (Brongn.) Florin, p. 108.
OCCURRENCE AND MATERIAL. Marl Slate of Middridge, Durham ; 3 specimens
(Hancock Museum). Hilton Plant Beds, Hilton Beck ; several specimens (B.M.
N.H. and author's coll.).
DESCRIPTION. Relatively good leaves from Middridge are shown in Text-fig. ioa
and PI. 38, fig. 6 ; unfortunately their cuticles are not preserved. Some recognizable
although less perfect leaves (Text-fig. 106) were found at Hilton and some of these
retain their cuticles. The cuticle proved delicate ; Text-fig, n shows parts of prepared
fragments. The cuticle has the rather elongated cells and ring of subsidiary cells
seen in many Ginkgoales. The stomatal pit is partly overhung by blunt papillae but
the ordinary epidermal cells have none.
A curious structure found at Hilton (PI. 38, fig. 4), looking like a capsule on a stalk,
rather resembles the groups of microsporangia of the Rhaetic species Baiera miln-
steriana Heer figured by Schenk (1884 : 261, fig. i8o&), but it is larger. The nature of
the specimen was not determined, and although it was found in the same bed as
leaves of Sphenobaiera digitata there is no proof that it belongs to that species.
320
THE UPPER PERMIAN FLORA OF ENGLAND
FIG. 10. Sphenobaiera digitata (Brongniart) Florin, (a), Marl Slate, Middridge. Hancock
Museum ; (b), Hilton Plant Beds, Hilton. British Museum (Natural History), no.
V.35ii8. Both x i.
FIG. ii. Sphenobaiera digitata (Brongniart) Florin. Cuticle. Hilton Plant Beds, Hilton.
(a), British Museum (Natural History), no. V. 35 135 ; (b), British Museum (Natural
History), no. ¥.35136. Both X 200.
THE UPPER PERMIAN FLORA OF ENGLAND 321
Genus CORDAITES Unger, 1850
cf. Cordaites aequalis (Goppert) Zalessky
(PL 38, fig. 3)
1845 Noeggerathia aequalis Goppert, p. 385, pi. 27, fig. 7.
1879 Rhiptozamites Goepperti Schmalhausen, pp. 32, 49, 81, pi. 4, figs. 2-4 ; pi. 7, figs. 23-27 ;
pi. 15, figs. i-n.
1912 Cordaites aequalis (Goppert) Zalessky, pi. i, figs, i, 3 ; pi. 2 ; pi. 3, figs, i, la, 3, 30, 4 ;
pis. 4, 5 ; pi. 6, figs. 1-3 ; pi. 7, fig. i.
1914 Cordaites aequalis (Goppert): Zalessky, p. 71, pi. 2, figs. 8, 8a, 10, ioa, 11-14,
6 bis, 6a.
OCCURRENCE AND MATERIAL. Fackley Hill (p. 299), probably from basement beds
of Magnesian Limestone Series ; a leaf impression in cream-coloured limestone
(G.S.M. no. 76680).
DESCRIPTION. The specimen (PI. 38, fig. 3) is the impression of the upper part of a
leaf with the base missing. It shows some obscure longitudinal ridges (visible in
the figure) and also fine longitudinal cellular striations. There are impressions of
similar but smaller bodies on the same piece of rock. No organic substance remains.
The specimen resembles leaves from the Russian Permian figured by Zalessky,
although with such scanty material the identification is not very convincing. If it
can be confirmed by the discovery of more material it will be interesting as C. aequalis
is unknown in the German Zechstein.
Genus PSEUDOCTENIS Seward, 1911
Pseudoctenis middridgensis sp. nov.
(PI. 38, fig. 5 ; Text-fig. 12)
DIAGNOSIS. Leaf as a whole broadly lanceolate, typically about 18 cm. x 6 cm.;
midrib straight, bearing pinnae laterally at an angle of 45° ; pinnae typically 40
mm. x 4-5 mm., widely spaced, with lower margin strongly decurrent on rachis,
contracted just above the base, and with upper part tapering from the middle to
an acute point ; base of pinna with two veins, each forking once or twice near the
pinna base to give about 6, which continue without branching or anastomoses to the
apex ; cuticle of upper side of rachis with straight-walled elongated cells, that of
other parts of leaf unknown.
OCCURRENCE AND MATERIAL. Marl Slate of Middridge, Durham ; holotype only
(Hancock Museum).
REMARKS. This specimen, catalogued by Howse (1890) under the nomen nudum
Ctenis permiana, is probably the best preserved fossil plant ever found at Middridge.
It is interpreted as consisting of two leaves on the same bedding plane, one lying
across the other. Detached and disarranged pinnae surround the lower end of the
rachis of the larger leaf. Parts of some pinnae are represented only by their impressions
on which the venation is faintly visible, and the remaining parts by a thin carbonaceous
film. The veins have no cross-connections. The cuticle is ill-preserved. A few frag-
322 THE UPPER PERMIAN FLORA OF ENGLAND
FIG. 12. Pseudoctenis middridgensis sp. nov. Marl Slate, Middridge. Hancock Museum.
X i.
THE UPPER PERMIAN FLORA OF ENGLAND 323
ments of cuticle from the upper side of the lamina were prepared, but the only ones
showing any structure were from the rachis. These have elongated cells 20 fi wide
in bands alternating with bands of less elongate cells very variable in size and shape.
No stomata could be recognized. The ordinary epidermal cells have simple anticlinal
walls which do not undulate, showing that the species does not belong to the
Bennettitales.
This specimen resembles various Mesozoic leaves placed in Pseudoctenis, but in
the absence of adequate knowledge of its cuticle its reference to that genus must be
considered provisional, and it is also not definitely separable from Pterophyllum,
though in typical species of that genus the pinnae arise rather above the lateral
margins of the rachis. No comparable leaf is known from the Zechstein, but several
from the Upper Carboniferous and Lower Permian have been referred to Pterophyllum,
They differ in their more crowded, parallel-sided pinnae, which are almost at right
angles to the rachis. The Permian leaf Ctenis renaulti Zalessky (1928 : 153) has
broader and more crowded pinnae. Plagiozamites Zeiller (1894), from the Carboniferous
and Permian, differs considerably in its crowded, broadly lanceolate pinnae.
CONIFERALES
Genus ULLMANNIA Goppert, 1850
Ullmannia bronni Goppert
(PI. 38, figs. ia,b, 2 ; PI. 39, figs. 3, 4)
1828 Cupressus Ullmanni Bronn, p. 526, pi. 4, figs. 1-7, 9 (?), 10 (for figs. 8, n see Strobilites
bronni) .
1850 Ullmannia Bronnii Goppert, p. 185, pi. 20, figs. 1-19, 236 (for figs. 20-22, 230, 24-26 see
Strobilites bronni).
1862 Ullmannia Bronni Goppert : Geinitz, p. 154, pi. 30, fig. 2 ; pi. 31, figs. 22-27 (non figs.
ii, 21, 28-30).
1876 Ullmannia Bronnii Goppert : Heer, p. 8, pi. 21, figs. 3-5.
1880 Ullmannia Bronni Goppert : Geinitz, p. 23, pi. 4, figs. 8-n, 14.
1884 Ullmannia Bronnii Goppert : Solms-Laubach, pp. 13-28, pi. 2, figs. 11-15.
1922 Ullmannia Bronni Goppert : Gothan & Nagalhard, p. 443, pi. 5, figs, i, 2 ; pi. 6, fig. i.
1928 Ullmannia Bronni Goppert : Weigelt, p. 564, pi. 35, figs. 1-5, 9.
1954 Ullmannia Bronnii Goppert : Florin, p. 2, pi. 2.
OCCURRENCE AND MATERIAL. Marl Slate of Raisby Hill quarries (p. 298), one fine
specimen (King's College, Newcastle-on-Tyne) , and of Middridge, one specimen
(G.S.M.). Isolated leaves (some as cuticle preparations) from Marl Slate of Cinderhill
(p. 300) (B.M.N.H.) and of Calverton Colliery shaft (p. 301), depth 402-456 ft., (B.M.
N.H.) ; from Lower Marl of Bentley Colliery (p. 300) (Wollaton Hall Mus.) and of
Sutton borehole (p. 300), depth, 1,085 ft. (B.M.N.H.) ; from Lower Limestone or
Lower Marl of Doddington borehole (p. 300), depths 2,300, 2,303 and 2,310 ft. (G.S.M.),
and of Wellow borehole (p. 301) (B.M.N.H.) ; and from Hilton Plant Beds of Hilton
(B.M.N.H.), Belah Brook, and Little Salkeld (Carlisle Mus.).
DESCRIPTION. The specimen from Raisby Hill (PL 38, figs. la, b) which is nearly
35 cm. long, consists of a main stem about 20 mm. wide and 20 cm. long which
324 THE UPPER PERMIAN FLORA OF ENGLAND
divides into four secondary branches about 13 mm. wide, two of which ultimately
bifurcate. Two of the secondary branches cross as they lie flattened on the bedding
plane. The main stem and branches are densely covered with imbricating leaves,
the impressions of which are clearly preserved. The leaves, have the broadly lance-
olate form and bluntly angular extremity characteristic of Ullmannia bronni.
Their median keel and longitudinal striations are visible in places. The leaves on
the main stem are about 4-5 mm. wide and twice as long and there are about five to
the width of the stem. The size of the leaves decreases towards the distal end of
each branch, but their proportions remain the same. No cuticle is preserved.
There is a poorly preserved specimen of U. bronni from Middridge, but most speci-
mens from that locality so labelled in museums belong to U. frumentaria or to
Hiltonia rivuli. Many of the isolated leaves from the borings and colliery shafts
have well-preserved cuticles. The preparations show characteristic dark bands with
stomata in single files, and agree with the cuticle of U. bronni as figured by Gothan
& Nagalhard (1922) and Florin (1944, 1954) from German and Belgian specimens,
and as prepared by myself from German material.
Goppert was not justified in changing the trivial name of this species when found-
ing the genus Ullmannia, but his name bronni is here retained in accordance with
long-established usage.
Ullmannia frumentaria (Schlotheim) Goppert
(PI. 39, figs, i, 2 ; Text-figs. 13, 14)
1820 Carpolithes frumentarius Schlotheim, p. 419, pi. 27, fig. i.
1828 Fucoides selaginoides Brongniart, p. 72, pi. 9, fig. 3.
1836 Voltzia Phillipsii Lindley & Hutton, p. 123, pi. 195.
1850 Caulerpa ? selaginoides (Brongn.) King, p. 3, pi. i, figs. 3, 30.
1922 Ullmannia frumentaria (Schloth.) : Gothan & Nagalhard, p. 445, pi. 5, fig. 3 ; pi. 6, figs.
3, 4. (See this for further literature.)
1928 A rchaeopodocarpus germanicus Weigelt, pp. 485-553, pi. 2, figs. 24, 25 ; pi. 3, figs, i, 4-6,
8 ; pi. 4, figs, i, 2, 7, 11-13 ; pi. 6, figs. 3-12 ; ? pi. 13, figs. 7, n, 13, 15, 17 ; pis. 23-38
(most figs.) ; pi. 31, figs. 1-3/6, 7, n.
1928 Strobilifer frumentarius (Schloth.) Weigelt, p. 553, pi. 30, figs, i, 2, 11-17, 28-31, 34;
pi. 35, figs. 10 a, b.
1944 Ullmannia frumentaria (Schloth.) : Florin, p. 447, pi. 169/170, figs. 10-12 ; p. 449, text-
figs. 44«, b (microspores) .
1944 Ullmannia Bronnii Goppert : Florin, p. 484, text-fig. 52 ; pi. 179/180, figs. 17-19 ;
pi. 181/182, figs, i, 2.
OCCURRENCE AND MATERIAL. Marl Slate of Cullercoats, Middridge, Ferryhill,
Thickley, Claxheugh, and Cinderhill ; numerous specimens in the chief collections
studied. Hilton Plant Beds, Hilton Beck and Belah Brook, fragments of shoots and
isolated leaves (B.M.N.H. and G.S.M.). Isolated leaves, some now as cuticle prepara-
tions, from Lower Marl of Sutton borehole (p. 300), depth 1,085 ft. (B.M.N.H.) ;
from Lower Limestone or Marl of Doddington borehole (p. 300), depth 2,327 ft.
(G.S.M.), and of Thoresby Colliery shaft (p. 301) (B.M.N.H.) ; and from Hilton Plant
Beds of Little Salkeld (Carlisle Mus.).
DESCRIPTION. Text-fig. 130 shows a shoot apex with densely overlapping leaves ;
PI. 39, fig. 2, the lower part of a shoot with more widely spaced leaves ; and Text-
THE UPPER PERMIAN FLORA OF ENGLAND
325
FIG. 13. Ullmannia frumentaria (Schlotheim) Goppert. (a), Marl Slate, Middridge.
British Museum (Natural History), no. "^35130 ; (b), Hilton Plant Beds, Hilton. Geo-
logical Survey, no. 19075 ; (c), Marl Slate, Thickley. British Museum (Natural History),
no. V.3239O ; (d), Marl Slate, Co. Durham. Dorman Memorial Museum, Middlesbrough.
All x i.
GEOL. 3, 9. 29
326
THE UPPER PERMIAN FLORA OF ENGLAND
fig. 13^ a shoot with rather short spreading leaves. It is noticeable that in some
of the compressed shoots the leaves look less regular than in others. There are
considerable differences in the size of the leaves on different shoots. Text-fig. 13^
and PL 39, fig. i show fertile shoots similar to those from Germany described by Florin
(1944). Unfortunately no details of the structure of the female cones can be made
out. These fertile shoots and the majority of sterile ones from Durham are unsuitably
preserved to give cuticle preparations. Rather poor preparations were obtained from
a specimen from Cullercoats and good ones from isolated leaves from Thoresby
Colliery shaft and from Hilton Beck. The details are like those of U, bronni ; the
stomata are in single files along strips darker than those between the files. The struc-
FIG. 14. Ullmannia frumentaria (Schlotheim) Goppert. Cuticle, (a), Marl Slate, Culler-
coats. British Museum (Natural History), no. V.35I37. x 150 ; (6), Magnesian Lime-
stone Series, Thoresby Colliery shaft. British Museum (Natural History), no. ¥.35138.
X 200.
ture agrees with that of the German specimens described by Gothan & Nagalhard
(1922) and by Krausel (1923).
The English specimens had previously been identified in museums as U. frumentaria,
U. selaginoides, and U. bronni. The second was considered a synonym of U. fru-
mentaria by Gothan & Nagalhard (1922), whose views are here accepted. The
difference between U. bronni and U. frumentaria lies in the shape of the leaves. In
U. bronni these are broadly lanceolate and mostly obtusely pointed, although some
have rounded tips ; their length does not exceed 2\ times their width. In U. fru-
mentaria they are relatively and often absolutely longer and the apex is acute. There
has, however, been some disagreement about the boundary between the two species.
THE UPPER PERMIAN FLORA OF ENGLAND 327
By the criterion stated, most English specimens labelled as U. bronni belong to U.
frumentaria.
The holotype of Voltzia phillipsii Lindley & Hutton, an early synonym of U.
frumentaria, has not been traced ; it came from the Marl Slate of Whitley, Durham.
Genus PSEUDOVOLTZIA Florin, 1927
Pseudovoltzia liebeana (Geinitz) Florin
(PL 40, figs, i, 4, 6 ; Text-figs. 15, 16)
1862 Cyclopteris Liebeana Geinitz, p. 140, pi. 26, figs. 4-6.
1880 Voltzia liebeana (Geinitz) Geinitz, p. 26, pi. 5, figs. 1-8, 10-25 (non ngs- 9. 26)-
1928 Voltzia Liebeana (Geinitz) : Weigelt, p. 565, pi. 5, fig. 4; pi. 16, figs. 4, 8, 9 ; pi. 31, figs.
4, 5, 8-10 ; pi. 32, figs. 1-29 ; pi. 33, figs. 1-19 ; pi. 34, figs. i-n.
1929 Voltzia Liebeana (Geinitz) : Walton, p. i, pi. i ; text-fig, i.
1929 Pseudovoltzia liebeana (Geinitz) Florin, p. 257, pi. 4, fig. 10.
1944 Pseudovoltzia Liebeana (Geinitz) : Florin, p. 413, text-fig. 26a ; p. 479, text-figs. 50, 51 ;
pi. 179/180, figs. 1-15.
OCCURRENCE AND MATERIAL. Marl Slate of Thickley (B.M.N.H. and King's
College, Newcastle-on-Tyne), of Ferryhill (G.S.M.), of Middridge ? (Hancock Museum),
and of Kimberley (B.M.N.H.). Lower Marl, Sutton borehole (p. 300), depth 1,085 ft-
(G.S.M.) and Markham Moor borehole (p. 300), depth 1,105 ft- (G.S.M.). Lower
Magnesian Limestone, Wressell borehole (p. 300), depth 1,982-1,984 ft. (G.S.M.).
FIG. 15. Pseudovoltzia liebeana (Geimtz) Florin. Cone scales, (a), Marl Slate, Kimberley.
British Museum (Natural History), no. ¥.35128 ; (b, c), Hilton Plant Beds, Hilton.
Geological Survey, nos. 19073, 19074 ; (d), Hilton Plant Beds, Hilton. British Museum
(Natural History), no. ¥.35119. All x 1-6.
Hilton Plant Beds, Hilton (B.M.N.H. and G.S.M.) , Belah Brook, and Birkshead Mine.
DESCRIPTION. The English material consists of shoots, isolated leaves, and isolated
cone scales. Typical shoots are shown in PI. 40, figs. 1,4; fig. 4 represents the largest
seen. The leaves in all the specimens are elongated, parallel-sided, and rounded at
the apex. Fig. i shows the short basal leaves produced when the bud first began
growth ; as Florin pointed out, this is a character of the species (cf. Geinitz, 1880,
pl- 5, ng. i).
The cuticle has perished in the Durham specimens, but moderately good preparations
were made from leaves from the Markham Moor borehole (Text-fig. 16). They look
just like the figure given by Florin (1944 : 413) and some preparations I made from
328
THE UPPER PERMIAN FLORA OF ENGLAND
German shoots. They are very similar to the cuticles of the two species of Ullmannia
— so similar, indeed, that isolated fragments not showing the form of the whole leaf
cannot be distinguished. There is, however, a possibility (which needs confirmation
from further material) that in Pseudovoltzia the subsidiary and ordinary cells along the
stomatal bands are less heavily cutinized ; that the ordinary epidermal cells are often
FIG. 1 6. Pseudovoltzia liebeana (Geinitz) Florin. Cuticle. Magnesian Limestone Series ;
depth 1,293 ft., Markham Moor borehole. Geological Survey, no. PF623. x 200.
shorter ; that the subsidiary cells have a more regularly pentagonal shape ; and that
the stomata are more often grouped in pairs or in threes. At present, however, the only
reliable distinction between leaves of Pseudovoltzia and Ullmannia lies in their very
different shape.
An excellent isolated cone scale from the Sutton borehole has been figured by
Walton (1929) and Florin (1944, pi. 179/180, figs. 14, 15). Text-fig. 15 shows isolated
cone-scales from Hilton and Kimberley preserved in carbonized form ; the lobes
are strikingly varied.
The specimens from the Manchester Marls preserved in the Manchester Museum
as the basis of records of this species by Geinitz (1890 : 549) and Roeder (1892 : 16)
are indeterminate.
Genus HILT ONI A Stoneley, 1956
Hiltonia rivuli Stoneley
1956 Hiltonia rivuli Stoneley, p. 714, text-figs. 1-4.
OCCURRENCE. Marl Slate, Middridge, Thickley, and Kimberley. Lower Marl,
Haughton Hall borehole (p. 300), depth i,n6ft. ; and of Sutton borehole (p. 300), depth
1,085 ft. Lower Marl or Lower Limestone, Doddington borehole (p. 300), depths
2,310 and 2,330 ft. ; of Wellow borehole (p. 301) ; and of Farnsfield borehole (p. 301).
Hilton Plant Beds, Hilton, Belah Brook, Birkshead Mine, and opencast mine near
Kirkby Thore.
THE UPPER PERMIAN FLORA OF ENGLAND 329
Genus CONITES Steinberg, 1823
Conites sp.
(PI. 40, fig. 5)
OCCURRENCE AND MATERIAL. Bulcote borehole (p. 301), depth 711 ft. (3 ft. 6 in.
above base of Upper Permian beds) ; one specimen (G.S.M. no. 76681).
DESCRIPTION. The only specimen is shown in the figure. The lateral appendages
seem irregularly spaced, but this is perhaps a result of compression of those originally
above or below the bedding plane on which the specimen lies. Both main axis and
appendages are longitudinally striated. There is nothing to show whether the
expanded distal ends of the appendages bore pollen sacs or seeds. The specimen
consists of crumbly carbonaceous matter and a fragment yielded no cuticle. The matrix
contained no leaf or other plant remains throwing light on its identity.
REMARKS. Nothing like this specimen has been previously described from the
Permian. There are, however, a good many obscure cones from the Mesozoic which
look something like it, and it is referred to Conites at the suggestion of Professor
T. M. Harris.
Genus STROBILITES Lindley & Hutton, 1833
Strobilites bronni Solms-Laubach
(PI. 40, figs. 7-11).
1850 Ullmannia Bronnii Goppert, p. 185 (in part), pi. 20, figs. 20 (?), 21, 22, 230,, 24-26.
1884 Strobilites Bronnii Solms-Laubach, p. 34, pi. 2, figs. 2, 3, 4 (?), 5-9, 16 (?) 17 (?) 18 (?)
I9(?).
1928 Strobilites Bronni Solms-Laubach : Weigelt, p. 474, pi. 10, fig. 5.
1930 Strobilites bronni Solms-Laubach : Weigelt, p. 644, pi. i, fig. 2.
1931 " Rosetten von Ullmannia bronni ": Weigelt, p. 106, 108, text-figs. 36, c.
1944 Strobilites Bronni Solms-Laubach : Florin, p. 447, pi. 169/170, figs. 13-22.
OCCURRENCE AND MATERIAL. Marl Slate, Kimberley ; several specimens (B.M.
N.H.). Hilton Plant Beds, Hilton ; several specimens (B.M.N.H., G.S.M.) ; also
of Belah Brook.
DESCRIPTION. The specimens occur carbonized or as mere impressions. They
show no fine details. The figures show the range of form. All can be matched among
published figures of German specimens.
REMARKS. There has been much doubt about the nature of these curious little
fossils, specimens of which from the Upper Zechstein were described as " Stern-
graupen " by Waldin (1778) and Ullmann (1802). Heer (1876) and Solms-Laubach
(1884) doubted Goppert's assumption that they belonged to Ullmannia, Weigelt
at first (1928) doubted but later (1931) accepted this view. Florin (1944) interpreted
them as isolated segments of Ullmannia male cones. They vary so much in form that
they may not all be of the same nature
330 THE UPPER PERMIAN FLORA OF ENGLAND
Strobilites ludwigi (Weigelt)
1928 " Weibliche Blute von Archaeopodocarpus ": Weigelt, p. 495, pi. 13, fig. 19.
1930 Araucarites ludwigi Weigelt, p. 660, text-fig. 39 ; ? also text-fig. 41 (p. 663).
1931 Araucarites ludwigi Weigelt : Weigelt, p. 107, 115, text-figs. 2, 12.
OCCURRENCE AND MATERIAL. Hilton Plant Beds, Hilton ; one specimen (B.M.
N.H., C. T. Trechmann Colin.).
DESCRIPTION. The specimen shows three rounded or bell-shaped groups of elongated
leaf-like organs and closely resembles the fossil figured by Weigelt. The specimen is
carbonized and no cuticle could be obtained from it ; no finer details are apparent.
REMARKS. Weigelt first named his species Araucarites, but this generic name has
been used by some authors for cones or cone scales which agree in structure with
Araucaria (e.g. in having one seed on the upper surface of each scale), and nothing
of the sort has been proved in this species. Weigelt abandoned his original view that
it was the female inflorescence of " Archaeopodocarpus " (Ullmannia). The non-
committal name Strobilites is here adopted for it.
Genus SAMAROPSIS Goppert, 1864
Samaropsis triangularis (Geinitz) Seward
(PI. 40, figs. 2, 3)
1862 Cardiocarpon triangulare Geinitz, p. 145, pi. 31, figs, n*, 12-15.
1880 Ullmannia frumentaria (Schloth.), " Fruchtschuppen mit Abdruck des Samens, fruher
Cardiocarpon triangulare ": Geinitz, p. 22, pi. 3, figs. 11-15.
1884 Cardiocarpon triangulare Geinitz : Solms-Laubach, p. 34, pi. 2, fig. 20.
1917 Samaropsis triangularis (Geinitz) Seward, p. 338.
1928 Cyclocarpon triangulare (Geinitz), " Fruchtschuppen, vielleicht aus Zapfen von Strobilifer
frumentarius ": Weigelt, p. 560, pi. 30, figs. 23-26.
OCCURRENCE AND MATERIAL. Marl Slate of Kimberley and Watnall ; several
specimens (B.M.N.H.).
REMARKS. The English specimens, which seem locally abundant, show the central
part of the seed as a solid body surrounded by a flat membrane. The surface of the
central part is slightly wrinkled. Geinitz (1880) and later Weigelt (1928) thought
that this might be the seed of Ullmannia frumentaria, but Florin has found a different
kind of seed in the seed-scale complex of that species.
UNIDENTIFIED CUTICLE TYPES
Bulk-macerated material from several boreholes and from the Hilton Plant Beds
yielded a number of well-characterized cuticle types distinct from any of those
prepared from identifiable plant remains. In some cases it was possible to ascertain
the form of the complete leaf, but in others the material consisted only of small
fragments. A set of descriptions and illustrations is available for consultation in
the British Museum (Natural History), where the cuticle slides are also deposited.
The identification of these cuticle types must await future research. More complete
remains of the plant species to which the majority belong probably have yet to be
THE UPPER PERMIAN FLORA OF ENGLAND 331
discovered and described, although a few of the new types may belong to described
species the cuticular structure of which has not yet been investigated owing to lack
of suitably preserved identifiable material.
NON-VEGETABLE STRUCTURES WHICH HAVE BEEN
REGARDED AS PLANT REMAINS
Guilielmites permianus Geinitz
(PI. 40, fig. 12)
1858 Guilielmites permianus Geinitz, p. 19, pi. 2, figs. 6-9.
1889 Guilielmites permianus Geinitz : Geinitz, p. 56 (English specimens).
1890 Guilielmites permianus Geinitz : Geinitz, p. 550 (English specimens).
18906 Guilielmitis (sic) permianus Geinitz : Roeder, p. 16 (English specimens).
OCCURRENCE AND MATERIAL. Upper Permian Marl of Fallowfield, near Manchester ;
specimens recorded by Geinitz and by Roeder are now in the Manchester Museum.
REMARKS. First described from the German Rotliegende and originally regarded
as a fructification of a palm related to the living Guilidma, Guilielmites is a peculiar
structure now known from many geological formations. Authors who have discussed
it include Weiss (1868 : 94 ; 1872 : 211), Carruthers (1871 : 446), Roemer (1880 :
246), Sterzel (1881 : 242), Hoffmann & Ryba (1899 : 102), Potonie (1921 : 16),
Pruvost (1930 : 260) and Wood (1935). The best figures are those of Hoffmann &
Ryba (1899, pi. 19, figs. 19, 20). Wood attributes such structures to the collapse
under pressure of some body, such as a shell or plant fragment, which has been
embedded in the sediment. As a result, a series of polished, striated surfaces came
into existence owing to local slipping of the rock, which is always a compact, fine-
grained shale.
Spongillopsis dyadica Geinitz
1889 Spongillopsis dyadica Geinitz, p. 56 (non S. dyatica Geinitz, 1862).
1890 Spongillopsis dyadica Geinitz : Geinitz, p. 550 (non S. dyatica Geinitz, 1862).
OCCURRENCE AND MATERIAL. Upper Permian Marl of Stockport, near Manchester ;
the specimen recorded by Geinitz, now in the Manchester Museum.
REMARKS. It is doubtful if the short, irregularly curved, occasionally branching
wrinkles which stand out on the surface of this specimen of red marl are of the same
nature as Geinitz's original 5. dyatica, itself problematical. The structures are
undoubtedly of inorganic origin and the name can be expunged from the list of
English Permian plants.
Palaeophycus insignis Geinitz
1 86 1 " Cast of a laterally compressed tube ..." Kirkby, p. 309.
1862 Palaeophycus insignis Geinitz, p. 131, pi. 24, fig. 4.
1890 Palaeophycus insignis Geinitz : Howse, pp. 237, 244.
1930 Palaeophycus insignis Geinitz : Weigelt, p. 645, text-fig. la, b.
OCCURRENCE AND MATERIAL. Kirkby (1861) described specimens which must have
332 THE UPPER PERMIAN FLORA OF ENGLAND
belonged to P. insignis from the Lower Magnesian Limestone of Hampole Stubbs,
near Doncaster. Howse (1890) recorded P. insignis from the Lower Magnesian
Limestone of Thickley and the Middle Magnesian Limestone of Tunstall Hill. The
only specimen seen by the present writer is from Thickley and is in the Hancock
Museum.
REMARKS. The specimen from Thickley closely resembles that described by Geinitz,
which came from the Dolomitic Zechstein near Gera and was a fragment, 40 mm.
long and 7 mm. wide, of a smooth, solid, rod-like body elliptical in cross-section.
He listed the species as an alga, but Weigelt (1930), who figured specimens from the
Kupferschiefer of Ilmenau, thought that they were the borings of arthropods. There
have been various theories as to the origin of other " species ", from different forma-
tions, described under Hall's genus Palaeophycus (Richter, 1927 : 198, 200 ; Schinde-
wolf, 1928 : 39 ; Wilckens, 1947 : 47 ; Korn, 1932 : 17). No modern authorities
consider them to be algae.
Chondrus binneyi King
(PI. 40, fig. 13)
1850 Chondrus ? Binneyi King, p. 2, pi. i, fig. i.
OCCURRENCE AND MATERIAL. Upper Permian Marl, just above Collyhurst Sand-
stone, of a river section at Newton, near Manchester (Binney, 1839 : 55 > I&55 '• 226).
The holotype, formerly in the Binney Coll., has not been traced, but a topotype
belonging to the Geological Survey has been examined. The specimen (G.S.M. no.
Be 4081) from the Ulleskelf Nurseries boring recorded as " cf. Chondrus binneyi
King " (Edwards et al., 1950 : 42) has also been examined.
DESCRIPTION. On the bedding planes of the red marl, in the topotype, are numerous
irregularly scattered circular structures, each about 2 mm. in diameter and having
the form of a raised ring with a central depression, or (as seen on the reverse surface)
of a central boss surrounded by a moat-like depression. Between these structures
each bedding plane is covered with sinuous fibrous markings, the general direction
of which varies considerably. The structures are composed of the same material
as the rest of the marl and carbonaceous matter is absent. They were regarded by
King (1850) as " seed vessels " sessile on a " broad frond ", but there is little doubt
they are of inorganic origin. They seem comparable to the " pit and mound "
structures described by Kindle (1916), Shrock (1948 : 132) and other authors.
The Ulleskelf specimen consists of hard grey limestone with comparable structures,
which, however, are more variable in size and often larger than in the topotype.
REFERENCES
ALTHAUS, J. 1846. Ueber einige neue Pflanzen aus dem Kupferschiefer von Riechelsdorf.
Palaeontographica, Stuttgart, 1 : 30-33, pis. i, 4.
ARBER, E. A. N. 1912. On Psygmophyllum majus sp. nov. from the Lower Carboniferous
rocks of Newfoundland, together with a revision of the genus and remarks on its affinities.
Trans. Linn. Soc. Land. (Bot.), (2), 7 : 391-408, pis. 42-44.
AVELINE, W. T. 1861. The geology of parts of Nottinghamshire and Derbyshire. Mem.
geol. Surv. U.K., Explan. J-Sheet 82 SE. 15 pp.
THE UPPER PERMIAN FLORA OF ENGLAND 333
BEIN, G. 1932. Die Stellung des Richelsdorfer Gebirges zum Thiiringer Walde und Rheinischen
Schiefergebirge. Z. dtsch. geol. Ges., Berlin, 84 : 786-829, pis. 25-27.
BINNEY, E. W. 1839. Sketch of the geology of Manchester and its vicinity. Manchr. geol.
Min. Soc. Trans., 1 : 35—66.
1855. On the Permian beds of the North-West of England. Mem. Manchr. lit. phil. Soc.
(2) 12 : 209-269.
BROCKBANK, W. 1892. On the Permians of the NW. of England. Discovery of two plant
beds in the St. Bees Sandstone at Hilton, Westmorland. Mem. Manchr. lit. phil. Soc.
(4) 5 : 66-76, pis. 2-4.
BRONGNIART, A. 1828. Prodrome d'une histoire des vegetaux fossiles. viii + 223 pp. Paris.
BRONN, H. G. 1828. Untersuchung der versteinerten Kornahren und anderer Pflanzentheile,
zu Cupressus Ullmanni gehorig, aus den Frankenberger Erz-Flozzen. Z. Miner., Heidelberg
(N.F.) 1828 (2) : 509-531, pi. 4.
CALVERT, R. 1884. Notes on the geology and natural history of the County of Durham. 184 pp.
Bishop Auckland.
CARR, J. W. 1914. Permian. In J. W. Carr & H. H. Swinnerton. Report of an excursion to
the Nottingham district. Proc. Geol. Ass., Land., 25 : 88-89.
CARRUTHERS, W. 1871. On some supposed vegetable fossils. Quart. J. geol. Soc. Land., 27 :
443-448, pi. 19.
DAKYNS, J. R., TIDDEMAN, R. H. & GOODCHILD, J. G. 1897. The geology of the country
between Appleby, Ullswater and Haweswater. Mem. geol. Surv. U.K., Explan. J-Sheet
102 SW. no pp.
DUNHAM, K. C. 1948. A contribution to the petrology of the Permian evaporite deposits of
North-Eastern England. Proc. Yorks. geol. Soc., 27 : 217-227, pi. 10.
DUNHAM, K. C. & ROSE, W. C. C. 1949. Permo-Triassic geology of South Cumberland and
Furness. Proc. Geol. Ass., Lond., 60 : 11-40.
ECCLES, J. 1871. On some sections of Permian strata near Kirkby Stephen. Manchr. geol.
Min. Soc. Trans., 10 : 30-37.
EDWARDS, W. 1951. The concealed coalfield of Yorkshire and Nottinghamshire. 3rd edit.
Mem. geol. Surv. U.K. x + 285 pp., 5 pis.
EDWARDS, W., MITCHELL, G. H. & WHITEHEAD, T. H. 1950. Geology of the district north
and east of Leeds. Mem. geol. Surv. U.K., Explan. Sheet 70. 93 pp.
ETTINGSHAUSEN, C. VON. 1851. Beitrage zur Flora der Vorwelt. Naturw. Abh., Wien (Haidinger),
4, i : 65-100, pis. 8-13.
FLORIN, R. 1929. tiber einige Algen und Koniferen aus dem mittleren und oberen Zechstein.
Senckenbergiana, Frankfurt a.M., 11 : 241-266, pis. 1-5.
1931. Untersuchungen zur Stammesgeschichte der Coniferales und Cordaitales. Erster
Teil : Morphologie und Epidermisstruktur der Assimilationsorgane bei den rezenten
Koniferen. K. svenska VetenskAkad. HandL, Stockholm (3) 10, i : 588 pp., 58 pis.
1936. Die fossilen Ginkgophyten von Franz-Joseph-Land, nebst Erorterungen iiber
vermeintliche Cordaitales mesozoischen Alters. Palaeontographica, Stuttgart, 81, B : 71-173,
pis. 11-42.
1938-45. Die Koniferen des Oberkarbons und des unteren Perms. Palaeontographica,
Stuttgart, 85, B. 729 pp., 186 pis.
1954- Note on Ullmannia from the Upper Permian formation of north-eastern Belgium.
Ass. Etude Paleont. Stratigr. houilleres, Publ., 18. 6 pp., 2 pis.
GEINITZ, H. B. 1848. Die Versteinerungen des deutschen Zechsteingebirges. 26 pp., 8 pis.
Dresden & Leipzig.
1861-62. Dyas, oder die Zechsteinformation und das Rothliegende. xviii -f- 130 pp., pis.
1-23 (1861) ; viii + 131-342 pp., pis. 24-42 (1862). Leipzig.
1863. Uber zwei neue dyadischen Pflanzen. Neues Jb. Min. Geol. Paldont., Stuttgart,
1863 : 525-530, pis. 6, 7.
1880. Nachtraege zur Dyas, I. Mitt, min.-geol. praehist. Mus. Dresden, 3. 43 pp., 7 pis.
1889. Ueber die roten und bunten Mergel der oberen Dyas bei Manchester. S.B. Isis
Dresden, 1889 : 48-57.
334 THE UPPER PERMIAN FLORA OF ENGLAND
GEINITZ, H. B. 1890. On the red and variegated marls of the Upper Dyas, near Manchester.
Manchr. geol. Min. Soc. Trans., 20 : 537-554.
GIBSON, W. & OTHERS. 1908. The geology of the southern part of the Derbyshire and Notting-
hamshire coalfield. Mem. geol. Surv. U.K., Explan. Sheet 125. 199 pp.
GIBSON, W. & WEDD. C. B. 1913. The geology of the northern part of the Derbyshire coalfield
and bordering tracts. Mem. geol. Surv. U.K. viii + 186 pp., 6 pis.
GOODCHILD, J. G. 1889. Excursion to North-West Cumberland and Edenside. Appleby,
Helton Beck and Roman Fell. Proc. Geol. Ass., Lond., 11 : xciv-xcvi.
i88ga. An outline of the geological history of the Eden Valley or Edenside. Proc. Geol.
Ass., Lond., 11 : 258-284.
GOPPERT, H. R. 1845. (Descriptions of fossil plants.) In P. de Tchihatcheff, Voyage scienti-
fique dans I' Altai oriental : 379-392, pis. 25-35. Paris.
1850. Monographic der fossilen Coniferen. 286 + 73 PP-, 5§ pis. Leiden.
1864-65. Die fossile Flora der permischen Formation. Palaeontographica, Stuttgart, 12.
316 pp., 64 pis.
GOTHAN, W. 1907. Callipteris Martinsi. In Potonie", H. Abbildungen und Beschreibungen
fossiler Pflanzen-Reste (K. preuss. geol. Landesanst.), 5, 96, 4 pp., 2 figs. Berlin.
1937- Zwei interessante Funde von Rotliegendpflanzen in Thuringen. Jb. preuss. geol.
Landesanst., Berlin, 57 : 507-513, pis. 27-29.
GOTHAN, W. & NAGALHARD, K. 1922. Kupferschieferpflanzen aus dem niederrheinischen
Zechstein. Jb. preuss. geol. Landesanst., Berlin, 42 : 440-460, pis. 5-7.
HANCOCK, A. & HOWSE, R. 1870. On a new Labyrinthodont Amphibian from the Magnesian
Limestone of Middridge, Durham. Quart. J. geol. Soc. Lond., 26 : 556-564, pi. 38.
HARKNESS, R. 1862. On the Sandstones and their associated Deposits in the Vale of the
Eden, the Cumberland Plain, and the South-east of Dumfriesshire. Quart. J. geol. Soc.
Lond., 18 : 205-218.
HEER, O. 1876. Ueber permischen Pflanzen von Fiinfkirchen in Ungarn. Mitt. Jb. ung.
geol. Anst., Budapest, 5 : 1-18, pis. 21-24.
HICKLING, G. 1931. The Summer Field Meeting, 1931, in Northumberland and Durham.
Proc. geol. Ass., Lond. 42 : 378-385.
HICKLING, G., ROBERTSON, T. & OTHERS. 1948. North-East Coast. Internal, geol. Congr.,
1 8 Sess., Excursion Guide C 9. 26 pp. London.
HOFFMANN, A. & RYBA, F. 1899. Leitpflanzen der palaeozoischen Steinkohlenablagerungen in
Mittel-Europa. 108 pp., atlas, 20 pis. Prag.
HOWSE, R. 1890. Catalogue of the local fossils in the Museum of the Natural History Society.
Nat. Hist. Trans. Newcastle, 10 : 227-288.
KINDLE, E. M. 1916. Small pit and mound structures developed during sedimentation. Geol.
Mag., London, 6 : 542-547, pi. 13.
KING, W. 1850. A Monograph of the Permian fossils of England, xxxviii + 258 pp., 28 pis.
[Mow. Pal. Soc., London].
KIRKBY, J. W. 1 86 1. On the Permian rocks of south Yorkshire ; and on their palaeontological
relations. Quart. J. geol. Soc. Lond., 17 : 287-325, pi. 7.
1862. On the remains of fish and plants from the Upper Limestone of the Permian series
of Durham. Ann. Mag. nat. Hist., London, 3 : 267-269.
1864. On some remains of fish and plants from the " Upper Limestone " of the Permian
series of Durham. Quart. J. geol. Soc. Lond., 20 : 345-358, pi. 18.
i864a. On some fossils from the Lower Magnesian Limestone of Sunderland. Trans.
Tyneside Nat. Field Club, 6 : 212-220.
1867. On some remains of fish and plants from the " Upper Limestone " of the Permian
series of Durham. Nat. Hist. Trans. Newcastle, 1 : 64-83, pi. 9.
18670. On the Fossils of the Marl-slate and Lower Magnesian Limestone (Durham,
No. n). Nat. Hist. Trans. Newcastle, 1 : 184-200.
KORN, H. 1932. Entgasungsspuren in Sedimenten. Chem. d. Erde, Jena, 7 : 1-23.
KRAUSEL, R. 1923. Palaobotanische Notizen. VII. Uber Papillenbildung an den Spaltoff-
nungen einiger fossiler Gymnospermen. Senckenbergiana, Frankfurt a.M., 5 : 81-96, pis.
3-4-
THE UPPER PERMIAN FLORA OF ENGLAND 335
KURTZE, G. A. 1839. Commentatio de petrefactis quae in schisto bituminose Mansfeldensi
reperiuntiur. 38 pp., 3 pis. Halae.
KUTORGA, S. 1838. Beitrag zur Kenntniss der organischen Ueberreste des Kupfersandsteins am
westlichen Abhange des Urals. 38 pp., 7 pis. St. Petersburg.
LAMPLUGH, G. W. & GIBSON, W. 1910. The geology of the country around Nottingham. Mem.
geol. Surv. U.K. vi + 72 pp.
LINDLEY, J. & HUTTON, W. 1835-37. The Fossil Flora of Great Britain, 3. 205 pp., pis. 157-230.
London.
MUNSTER, G. 1842. Ueber die Fucoiden des Kupferschiefers. Beitr. Petrefacten-Kunde,
Bayreuth, 5 : 100-102, pis. 14, 15.
MURCHISON, R. I. & HARKNESS, R. 1864. On the Permian rocks of the north-west of England,
and their extension into Scotland. Quart. J. geol. Soc. Lond., 20 : 144-165.
NAUMANN, E. 1934. Zur Stratigraphie und Lagerung des Konigseer Zechsteins. Jb. preuss.
geol. Landesanst., Berlin, 54 : 194-203, pis. i$a, b.
POTONIE, H. 1921. Lehrbuch der Paldobotanik. Edit. 2. 537 pp., 326 figs. Berlin.
PRAGER, H. 1932. Die Perm-Trias-Grenze nordlich des Kellerwaldes. Z. dtsch. geol. Ges.,
Berlin, 84 : 317-360, pi. 9.
PRUVOST, P. 1930. La faune continentale du terrain houiller de la Belgique. Mem. Mus.
Hist. nat. Belg., Bruxelles, 44 : 105-282, pis. 1-14.
REMY, W. 1953. Untersuchungen iiber einige Fruktifikationen von Farnen und Pterido-
spermen aus dem mitteleuropaischen Karbon und Perm. Abh. dtsch. Akad. Wiss. Berl.,
1952, 2 : 38 pp., 7 pis.
RICHTER, R. 1927. Die fossilen Fahrten und Bauten der Wurmer. Paldont. Z., Berlin, 9:
193-235. Pis. 1-4-
ROEDER, C. 1 890. Some newly discovered species in the Upper Permian deposits of Manchester.
Manchr geol. Min. Soc. Trans., 20 : 535-537.
iSgoa. Notes on the Upper Permians, etc., at Fallowfield, lately laid open. Manchr
geol. Min. Soc. Trans., 20 : 615-627.
18906. Description of fossils found in the Permian Upper Series, at Fallowfield. Manchr
geol. Min. Soc. Trans., 21 : 13-16.
1892. Notes on the Permians and superficial beds at Fallowfield. Manchr geol. Min. Soc.
Trans., 21 : 104-113.
ROEMER, F. 1880. Systematische Aufzahlung und Beschreibung der bezeichnenden palao-
zoischen Fossilien. I. Pflanzen. Lethaea geognostica, I, Lethaea palaeozoica, 1 : 115-258.
Stuttgart.
SCHENK, A. 1864. Beitrage zur Flora der Vorwelt. Palaeontographica, Stuttgart, 11 : 296-308,
pis. 46-49.
1884-90. Completion of W. P. Schimper, Palaeophytologie. Handb. Palaeont. (edit. K.
A. Zittel), 2 : 233-958. Miinchen & Leipzig.
SCHINDEWOLF, O. H. 1928. Studien aus dem Marburger Buntsandstein, III- VII. Sencken-
bergiana, Frankfurt a.M., 10 : 16-54.
SCHLOTHEIM, E. F. VON. 1820. Die Petrefactenkunde. Ixii + 437 pp., atlas 29 pis. Gotha.
SCHMALHAUSEN, J. 1879. Beitrage zur Jura-Flora Russlands. Mem. Acad. Sci. St-Petersb.
(Sci. math., phys., nat.), 27, 4 : 96 pp., 16 pis.
— 1887. Die Pflanzenreste der artinskischen und permischen Ablagerungen im Osten des
europaischen Russlands. Mem. Com. geol., St-Petersb., 2, 4 : 1-42, pis. 1-7.
SCHUSTER, A. 1933. Oberrotliegendes und Zechstein in Sachsen. Abh. sdchs. geol. Landesamts,
Leipzig, 13 : 1-85, pis. 1-8.
SCHUSTER, J. 1911. Uber die Fruktifikation von Schuetzia anomala. S.B. Akad. Wiss. Wien,
120, i : 1125-1134, pis. i, 2.
SEDGWICK, A. 1829. On the geological relations and the internal structure of the Magnesian
Limestone, and the lower portions of the New Red Sandstone series in their range through
Nottinghamshire, Derbyshire, Yorkshire, and Durham, to the southern extremity of North-
umberland. Trans, geol. Soc. Lond., 2, 3 : 37-124.
336 THE UPPER PERMIAN FLORA OF ENGLAND
SEWARD, A. C. 1912. Mesozoic plants from Afghanistan and Afghan-Turkistan. Palaeont.
indica, Calcutta (N.S.) 4, 4 : 1-57, pis. 1-7.
1917- Fossil Plants, 3. xviii -f- 656 pp. 253 figs., London.
SHROCK, R. R. 1948. Sequence in layered rocks. 507 pp. New York.
SIMPSON, S. 1957. On the trace-fossil Chondrites. Quart. J. geol. Soc. Lond., 112 : 475-499,
pis. 21-24.
SMITH, B. 1913. The geology of the Nottingham district. Proc. geol. Ass., Lond., 24 : 205-240,
pis. 38-41.
1919. Iron ores : haematites of West Cumberland, Lancashire and the Lake District.
Mem. geol. Surv. U.K., Special Repts. min. Resources, 8. 182 pp.
SOLMS-LAUBACH, H. 1884. Die Coniferenformen des deutschen Kupferschiefers und Zech-
steins. Palaont. Abh., Berlin, 2 : 81-116, pis. 12-14.
STERZEL, J. T. 1881. Palaontologischer Charakter der oberen Steinkohlenformation und
des Rotliegenden im Erzgebirgischen Becken. Ber. naturw. Ges. Chemnitz, 7 : 155-270.
STONELEY, H. M. M. 1956. Hiltonia, a new plant genus from the Upper Permian of England.
Ann. Mag. nat. Hist., London (12) 9 : 713-720, 5 figs.
TRECHMANN, C. T. 1921. Some remarkably preserved brachiopods from the Lower Magnesian
Limestone of Durham. Geol. Mag., London, 58 : 538-543, pi. 12.
1925. The Permian formation in Durham. Proc. geol. Ass. Lond., 36 : 135-145.
1931. Contributions to the geology of Northumberland and Durham. V. The Permian.
Proc. geol. Ass., Lond., 42 : 246-252.
1942. Borings in the Permian and Coal Measures around Hartlepool. Proc. Yorks. geol.
Soc., 24 : 313-327.
1952. (Contribution to discussion of paper by L. R. Raymond on exploration for potash
in NE. Yorkshire.) Quart. J. geol. Soc. Lond., 108 : 307.
TROTTER, F. M. & HOLLINGWORTH, S. E. 1932. The geology of the Brampton district.
Mem. geol. Surv. U.K., Explan. Sheet 18. 223 pp.
ULLMANN, J. C. 1802. Mineralogische Beobachtungen, 1. 160 pp., 4 pis. Marburg.
WALDIN, J. G. 1778. Die Frankenberger-Versteinerungen, nebst ihren Ursprunge. 32 pp., i pi.
Marburg.
WALTON, J. 1929. On the structure of a Palaeozoic Cone-Scale and the evidence it furnishes
of the primitive nature of the Double Cone-Scale in the Conifers. Mem. Manchr lit. phil.
Soc., 73 : 1-6, pi. i.
WEIGELT, J. 1928. Die Pflanzenreste des mitteldeutschen Kupferschiefers und ihre Ein-
schaltung ins Sediment. Fortschr. Geol., Berlin, 6 : 395-592, 35 pis.
1930. Neue Pflanzenfunde aus dem Mansfelder Kupferschiefer. Leopoldina, Halle,
6 : 643-668, 2 pis.
1931. Neue Pflanzenfunde aus dem Mansfelder Kupferschiefer. Z. Naturw., Leipzig,
89 : 104-125, 19 figs.
1932. Uber die Fruktifikationsverhaltnisse von Kupferschiefer-Coniferen und andere
neue Pflanzenfunde. Palaont. Z., Berlin, 14 : 137-149, pis. 3, 4.
WEISS, C. E. 1868. Begrundung von funf geognostischen Abtheilungen in den Steinkohlen-
fiihrenden Schichten des Saar-Rheingebirges. Verh. naturh. Ver. preuss. Rheinl., Bonn,
25 : 63-134.
1869-72. Fossile Flora der jiingsten Steinkohlenformation und des Rothliegenden im Saar-
Rhein-Gebiete . 250 pp., 20 pis. Bonn.
WILCKENS, O. 1947. Palaontologische und geologische Ergebnisse der Reise von Kohl-Larsen
(1928-29) nach Siid-Georgien. Abh. senckenb. naturf. Ges., Frankfurt a.M., 474 : 1-66,
pis. 1-9.
WILSON, E. 1876. On the Permians of the North-East of England (at their southern margin)
and their relations to the under- and overlying formations. Quart. J. geol. Soc. Lond.,
32 : 533-537.
1 88 1. The Permian formation in the north-east of England, with special reference to
the physical conditions under which these rocks were formed. Mid. Nat., Birmingham,
4 : 97-101, pi. 6 ; 121-124, pi. 7 ; 187-191 ; 201-208.
THE UPPER PERMIAN FLORA OF ENGLAND 337
WOOD, A. 1935. The origin of the structure known as Guilielmites. Geol. Mag., London,
72 : 241-245, pi. 10.
WOOLACOTT, D. 1898. An explanation of the Claxheugh section (Co. Durham). Quart. J.
geol. Soc. Land., 54 : 14.
1903. An explanation of the Claxheugh section (Co. Durham). Nat. Hist. Trans. Newcastle,
14 : 211-221.
1912. The stratigraphy and tectonics of the Permian of Durham (northern area). Proc.
Univ. Durham phil. Soc., 4 : 241-313.
ZALESSKY, M. D. 1912. Sur le Cordaites aequalis Goppert sp. de Siberie et sur son identite
avec la Noeggerathiopsis hislopi Bunbury sp. de la flore du Gondwana. Mem. Com. geol.,
St.-Petersb. (N.S.) 86 : 1-43, pis. 1-7.
1914. Flore gondwanienne du bassin de la Petchora. Bull. Soc. oural. Sci. not., Ekaterinburg,
33 : 55-85, pis. 1-4.
1927. Flore permienne des limites ouraliennes de 1'Angaride. Mem. Com. geol., Leningr.
(N.S.) 176 : 1-52, pis. 1-46.
1928. Sur 1'extension du continent de 1'Angaride et premieres donnees sur la flore de ses
limites oussouriennes. Ann. Soc. geol. Nora, Lille, 53 : 118-138.
1932. Observations sur les vegetaux nouveaux paleozoiques de Siberie. Ann. Soc. geol.
Nord, Lille, 57 : 111-134.
1937- Sur la distinction de 1'etage Bardien dans le Permien de 1'Oural et sur sa flore
fossile. Probl. Palaeont., Moscow, 2-3 : 31-101.
ZEILLER, R. 1894. Notes sur la flore des couches permiennes de Trienbach (Alsace). Bull.
Soc. geol. Fr., Paris (3) 22 : 163-182, pis. 8, 9.
PLATE 36
FIG. i. Algites sternbergianus (King). Fine filaments are visible on the curved axis. Lower
Magnesian Limestone, Westoe, South Shields. Geological Survey, no. 90165. x i.
FIG. 2. Same species, formation, and locality. Group of axes. Geological Survey, no. 90164.
X i.
FIG. 3. Piaea anglica sp. nov. Group of axes. Magnesian Limestone series, depth 1,140 ft.,
Markham Moor borehole, near East Retford. Geological Survey, no. 76607. x 2 approx.
FIG. 4. Calathella dictyonemoides sp. nov. Magnesian Limestone series, depth 1,056 ft.,
Markham Moor borehole, near East Retford. Geological Survey, no. 76608. x i.
FIG. 5. Algites sternbergianus (King). Brush of filaments. Same formation and locality as
Figs, i, 2. Geological Survey, no. 49956. X 2-2.
FIG. 6. ? Tubulites permianus (King) . Bunched tubes. Upper Magnesian Limestone, Black-
hall Rocks, 5 miles N. of Hartlepool. C. T. Trechmann Colin., British Museum (Natural History),
no. V.35IH. x 3.
FIG. 7. Tubulites permianus (King). Upper Magnesian Limestone (concretionary beds),
Easington. C. T. Trechmann Colin., British Museum (Natural History), no. "^35113. x 2-4.
FIG. 8. Same species. Transverse sections of tubes. Upper Magnesian Limestone, depth
4,240 ft., Eskdale borehole no. 3, Sleights, near Whitby. Geological Survey, no. PF625. x 5.
FIG. 9. Same species. Longitudinal sections of tubes. Same borehole and depth as the
preceding. Geological Survey, no. PF624. x 5.
Bull. B.M. (N.H.) Geol. 3, 9
PLATE 36
PLATE 37
FIG. i Paracalamites kutorgai (Geinitz) Zalessky. Enlarged view of part of flattened stem
fragment, with a node near left of figure. Marl Slate, Fulwell Water Works, near Sunderland.
Hancock Museum, x 3, approx.
FIG. 2. Callipteris martinsi (Kurtze) Zeiller. Cuticle, showing a stoma with a prominent
papilla. Marl Slate, Kimberley. British Museum (Natural History), no. ¥.5963^. x 200.
FIG. 3. Taeniopteris eckardti Kurtze. Marl Slate, Middridge. Hancock Museum, (a) x i ;
(b) part of specimen, x 2, showing details of veiris.
FIG. 4. Mixoneura sp. Pinnule from the holotype of Neuroptsris huttoniana King. Marl
Slate, Thrislington Gap, near Ferryhill. University College, Galway. x 2-3.
FIG. 5. Callipteris martinsi (Kurtze) Zeiller. The pinnules show a midrib and veins, also,
at their base, scars of unknown origin. Marl Slate, Kimberley. British Museum (Natural
History), no. ¥.5960. x 1-5.
FIG. 6. Sphenopteris bipinnata (Munster) Geinitz. Hilton Plant Bads, Hilton. British
Museum (Natural History), no. V.35ii6. x 1-4.
FIGS. 7, 8. Schutzia ? sp. Crushed specimens possibly belonging to the same species as
Figs. 9-12, but with the sporophylls (?) relatively few and distinct. Hilton Plant Beds, Hilton.
British Museum (Natural History), nos. V.35I.2O-2I. X i.
FIG. 9. Schutzia ? sp. Two associated cup-shaped sporangial aggregates, preserved almost
uncompressed but with their structure obscure. Hilton Plant Beds, Hilton. C. T . Trechmann
Colin., British Museum (Natural History), no. V.2O554. x i.
FIG. 10. Schutzia ? sp. Laterally crushed specimen, with the opened-out sporophylls
clearly seen. Hilton Plant Beds, Hilton. British Museum (Natural History), no. ¥.35122. x 3.
FIG. ii. Schutzia ? sp. Specimen crushed so as to present much the same aspect as Fig. 10.
Hilton Plant Beds, Hilton. Royal Scottish Museum, no. 1957. 15.1. X 3.
FIG. 12. Schutzia ? sp. Crushed specimen ; the under side with a hollow for stalk
attachment is seen, but the outlines of individual sporophylls are mostly indistinct. Hilton
Plant Beds, Hilton. Geological Survey, no. 19068. x i.
Bull. B.M. (N.H.) Geol. 3, 9.
PLATE 37
PLATE 38
FIGS. la. b. Ullmannia bronni Goppert. Impression of a forking branch. Marl Slate,
Raisby Hill Quarries, near Coxhoe. King's College, Newcastle-upon-Tyne. (a) Complete speci-
men, x 0-38, (b) part of specimen, showing densely imbricating leaves, x 0-77.
FIG. 2. Same species. Marl Slate, Middridge. Geological Survey, no. 19067. x i.
FIG. 3. cf. Cordaites aequalis (Goppert) Zalessky. Leaf impression. Basement beds of
Magnesian Limestone series, Fackley Hill, near Skegby. Geological Survey, no. 76680. x i.
FIG. 4. Capsule of uncertain affinities, from bed containing Sphenobaiera digitata (Brongniart) .
Hilton Plant Beds, Hilton. British Museum (Natural History), no. V. 35 123. x i.
FIG. 5. Pseudoctenis middridgensis sp. nov. Portion of rachis and bases of several pinnae,
showing veins. Marl Slate, Middridge. Hancock Museum, x 1-8. (see also Text-fig. 12).
FIG. 6. Sphenobaiera digitata (Brongniart) Florin. Marl Slate, Middridge. Hancock Museum.
X i
Bull. B.M. (N.H.) Geol. 3, 9
PLATE 38
PLATE 39
FIG. i. Ullmannia frumentaria (Schlotheim) Goppert. Female cone. Marl Slate, Middridge.
British Museum (Natural History), no. V. 35 129. x i.
FIG. 2. Same species. Marl Slate, Thickley. C. T. Trechmann Colin., British Museum'
(Natural History), no. ¥.20553. x I-
FIG. 3. Ullmannia bronni Goppert. Cuticle. Near base of Magnesian Limestone series,
Wellow borehole. British Museum (Natural History), no. V. 35139. X 150.
FIG. 4. Same species. Cuticle with two stomata. Near base of Magnesian Limestone series,
Calverton Colliery shaft. British Museum (Natural History), no. ¥.35140. x 270.
FIG. 5. Stvobilites ludwigi (Weigelt) ; also two leaves of Hiltonia, near right and left margins.
Hilton Plant Beds, Hilton. C. T. Trechmann Colin., British Museum (Natural History), no.
¥.20555. x i.
Bull. B.M. (N.H.) Geol. 3, g
PLATE 39
PLATE 40
FIG. i. Pseudovoltzia liebeana (Geinitz) Florin. Marl Slate, Thickley. King's College,
Newcastle-upon-Tyne, no. 2525. x i.
FIGS. 2, 3. Samaropsis triangularis (Geinitz) Seward. Marl Slate, Kimberley. British
Museum (Natural History), nos. ¥.6210, ¥.6206. x i.
FIG. 4. Pseudovoltzia liebeana (Geinitz) Florin. Marl Slate, Thickley. British Museum
(Natural History), no. 46651. x i.
FIG. 5. Conites sp. Near base of Upper Permian Beds, Bulcote borehole, near Nottingham.
Geological Survey, no. 76681. x 2.
FIG. 6. Pseudovoltzia liebeana (Geinitz) Florin. Impression of cone scale. Marl Slate,
Kimberley. British Museum (Natural History), no. ¥.6228. x i.
FIG. 7. Strobilites bronni Solms-Laubach. Carbonized specimen. Hilton Plant Beds,
Hilton. British Museum (Natural History), no. ¥.35124. x 1-5.
FIG. 8. Same species. Impression in sandstone. Hilton Plant Beds, Hilton. British Museum
(Natural History), no. ¥.35125. x i.
FIG. 9. Same species. Marl Slate, Kimberley. British Museum (Natural History), no.
¥.35126. x i.
FIG. 10. Same species. Hilton Plant Beds, Hilton. Geological Survey, no. 19070. x i.
FIG. ii. Same species. Marl Slate, Kimberley. British Museum (Natural History), no.
¥.35127. x i.
FIG. 12. Guilielmites permianus Geinitz. A non-vegetable structure. Manchester Marls,
Fallowfield. Manchester Museum, no. 1^.707. x 3-5.
FIG. 13. Chondrus binneyi King. A non-vegetable structure. Manchester Marls, Newton,
Manchester. Geological Survey, no. 19048. x 1-7.
FIGS. 14,15. ? Psygmophyllum cuneifolium (Kutorga) Schimper. Cuticle. Marl Slate, Middridge.
British Museum (Natural History), nos. ¥.35133-34. x 150.
Bull. EM. (N.H.) Geol. 3, g
PLATE 40
BLUE-GREEN ALGAE FROM THE
MIDDLE DEVONIAN OF
RHYME, ABERDEENSHIRE
W. N. CROFT and E. A. GEORGE
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 10
LONDON : 1959
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY
The following papers appeared in Volume I (1949-52) :
Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.
H. D. Thomas & A. G. Davis 75. 6d.
No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.
Oakley & M. F. Ashley Montagu ...... 55.
No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E.I. White
Pteraspis leathensis White, a Dittonian Zone-Fossil. E. I.
White 75. 6d.
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,
Northern Iraq. L. F. Spath . . . . . 105.
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-
pellum. T. H. Withers 55.
No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .
T. H. Withers 55.
No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian
of Podolia. W. N. Croft los.
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle
East. T. F. Grimsdale los.
No. 9(1952). Australian Arthrodires. E. I. White .... 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 65.
The following papers appeared in Volume II (1953-56) :
No. i (1953). The Deer of the Weybourn Crag and Forest Bed of
Norfolk. A. Azzaroli . . . . . . . . £i $s.
No. 2 (1953). A Coniferous Petrified Forest in Patagonia. M. G. Calder 125.
No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner,
K. P. Oakley & W. E. Le Gros Clark 35. 6d.
No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.
M. E. J. Chandler 165.
No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans . 155.
No. 6 (1955). Further Contributions to the Solution of the Piltdown
Problem. J. S. Weiner, W. E. Le Gros Clark & K.P. Oakley et al. £i
No. 7 (1955) . The Schizaeaceae of the South of England in Early Tertiary
Times. M. E. J. Chandler . 1:55.
No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . £i
BLUE-GREEN ALGAE FROM THE MIDDLE
DEVONIAN OF RHYNIE, ABERDEENSHIRE
BY
WILLIAM N. CROFT*
British Museum (Natural History)
and
ERIC ALAN GEORGE
Botany School, Cambridge
PP- 339-353 / Plates 41-44
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 3 No. 10
LONDON : 1959
* Deceased.
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 10 of the Geological series.
Trustees of the British Museum, 1959
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1959 Price Eight Shillings
BLUE-GREEN ALGAE FROM THE MIDDLE
DEVONIAN OF RHYNIE, ABERDEENSHIRE
By the late W. N. CROFT AND E. A. GEORGE
SYNOPSIS
Three new blue-green algae, Langiella scourfieldi nov. gen. et sp. (Stigonemataceae), Kidstoni-
ella fritschi nov. gen. et sp. (Stigonemataceae) and Rhyniella vermiformis nov. gen. et sp.
(incertae sedis) are described from the Middle Devonian Rhynie Chert Bed in Scotland. The
material is well preserved and details of the sheath, cell wall, cell contents and heterocysts are
clearly seen. It supports the view that the Myxophyceae are an ancient stock and indicates
that the Stigonemataceae which are generally regarded as being among the most advanced
members of the class existed as early as the Devonian age.
I. INTRODUCTION
SINCE the classical memoirs of Kidston & Lang (1917-21) on the vascular cryptogams
and thallopttytes of the Rhynie Chert Bed, no further work has been published
on the plants of this deposit.
Archaeothrix (Kidston & Lang, 1921) is the only fossil so far described from this
chert which could reasonably be placed within the blue-green algae. The material
on which the present account is based confirms the presence of Myxophyceae in
the deposit and adds very materially to the knowledge of the fossil history of this
important group of plants.
The specimens, placed in three new genera, occur in a single small chip of the chert.
This was found labelled " ? Blue-green alga " in D. J. Scourfield's collection which was
bequeathed to the British Museum (Natural History) in 1950.
The rarity of remains of blue-green algae in the deposit is in contrast with the
abundance of other plants, especially fungi. It is the more striking when it is remem-
bered that these algae are generally abundant at the present day in warm springs
in which siliceous material is being deposited. It suggests that the habitat was little
favourable to their growth, but is in no way contradictory to the well founded view
that the peat bog was overwhelmed by flooding (Kidston & Lang, 1921 : 892).
The method adopted for studying the plants in the chip was that used by Scour-
field (1926 : 154) for the crustacean remains ; direct examination in oil with an
oil-immersion objective. Examination and illustration of the plants are rendered
difficult by the thickness of the chip and the amount of debris contained in it. It was
often necessary to readjust the setting of the substage mirror even for examination
of different parts of the same plant. Moreover the cell walls often contrast little with
the matrix and the outlines and visibility of the cells and of the sheaths vary consider-
ably with the setting of the mirror and the width of the illuminating cone.
GEOL. 3, IO. 30
342 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
II. DESCRIPTION OF THE MATERIAL
The chip is about I mm. thick and nearly 5 mm. across. The topography of the
two sides is shown at a magnification of X 14 in PI. 41, figs. 1,2. In the upper right-
hand corner of fig. i numerous trichomes of Langiella, including the type specimen,
are present. They tend to be similarly orientated (PI. 43, fig. 17). Numerous trichomes
of the same species are seen in the same corner of the chip from the other side (PL 41,
fig. 2), but they are not aligned. Near the middle of the upper edge of the chip
(PL 41, fig. i) is the only specimen of Rhyniella. Associated with it are a few filaments
of Langiella. Running obliquely through the lower right-hand portion of the chip
in PL 41, fig. 2 is part of what is presumed to be a vascular axis about 1*4 mm. in
diameter. Along the edge of this axis and partly detached from it is a narrow zone
of trichomes of Kidstoniella. As suggested below (p. 348) the plant probably grew
epiphytically upon this axis. A few trichomes showing branching, near the middle
.of the face of the chip (PL 41, fig. i) may belong to the same species. Many fragments
of plant tissue are present together with abundant fungal hyphae. These latter are
occasionally branched and rarely septate. Spores have not been observed.
The occurrence of these three species in such close proximity is not unlike present-
day communities of Myxophyceae. Mats and cushions consisting of several species
or genera almost inextricably interwoven are a characteristic covering of damp
surfaces to-day.
That so much has been learnt of the algal flora of the Rhynie Chert from one small
chip encourages the hope that further fortunate discoveries may add still more to
our knowledge.
III. SYSTEMATIC DESCRIPTIONS
MYXOPHYCEAE
Order STIGONEMATALES
Family STIGONEMATACEAE
Genus LANGIELLA nov.
DIAGNOSIS. Mature thallus heterotrichous. Prostrate axis multiseriate, cells
large and moderately thick-walled with both transverse and longitudinal divisions,
also rounded cells in small aggregations, branches few, heterocysts absent, sheaths
thin. Erect filaments essentially uniseriate, unbranched, trichomes more or less
strongly torulose, cells generally smaller and except in the proximal region of some
filaments, thinner-walled than those of the prostrate system ; sheaths generally
thin, but thicker in middle and upper parts of some filaments ; in the upper parts of
some filaments trichomes much narrower with more elongate cells or no clear evidence
of cell walls ; ends of some filaments multiseriate with cells formed by oblique
divisions. Heterocysts, at or near the base of most erect filaments, absent from the
basal system, walls refractive, contents depleted, shape subspherical to cylindrical.
Reproduction by fragmentation of erect filaments at heterocysts, probably by single
akinetes formed in basal system, possibly by catenate akinetes formed in lower part
of erect filaments, possibly by hormogonia released from the tips of erect filaments.
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 343
Diameter of prostrate axes, 22 fi to 40 fi ; diameter of prostrate axes cells, 12 /*
to 20 [i.
Length of erect filaments about 100 /£ to 300 fi.
Maximum width of trichome (lower part), 16 /^. Minimum width of narrowest
trichome, 8 fi.
Length of heterocysts, 6 fi to 10 /* ; width 6 JJL to 8 //.
TYPE SPECIES. Langiella scourfieldi n. sp.
Langiella scourfieldi n. sp.
(Plate 41, figs, i, 3-8, 10 ; PI. 42 ; PL 43, figs. 17, 19-24)
DIAGNOSIS. As for genus.
HOLOTYPE. Specimen No. ¥.32409, Palaeont. Dept. Brit. Mus. (Nat. Hist.)
OCCURRENCE. Middle Devonian (Old Red Sandstone) ; Muir of Rhynie, Aberdeen-
shire, Scotland.
General morphology. The holotype (PI. 41, fig. 10 and PI. 42, fig. 12) is markedly
heterotrichous and the same habit is indicated by some of the obscurely seen specimens
alongside it (to the left in PI. 41, fig. i). A multiseriate basal system, curved round into
the shape of an asymmetrical horse-shoe, gives rise on its outer edge to a number of
uniseriate branches. Most of these, including two or three apparently detached
filaments are subparallel, similarly orientated and lie in almost the same plane. Some
at least of the uniseriate branches were probably fertile. Possibly the basal system
was originally more or less straight and creeping with erect branches arising on the
side opposite the substrate. Most of the specimens in the chip are, however, clearly
unattached and there is no direct evidence that they ever were attached to a sub-
strate. A description of a few associated forms follows that of the holotype (p. 346).
Basal system. The compacted cells of the basal system are generally broader and
thicker-walled than those composing the erect filaments. They are not arranged in
regular transverse rows as is usually the case in present-day species of Stigonema.
The left-hand portion of the basal system (PI. 41, fig. 10) attains a thickness of about
40^. A row of broad thick-walled cells formed by transverse and longitudinal
divisions, up to 18 /* across, is clearly seen on its surface. Beneath these, at a deeper
focus, a few small cells with polygonal outlines suggesting mutual pressure are visible.
The dense right-hand portion shows a row of three large thick-walled cells 20 /£ in
diameter. The middle, curved portion of the basal system consists of rounded cells
one of which with a thick wall and a diameter of about 18 fi is suggestive of an akinete
(PI. 42, fig. 12). Some of the cells of the basal region may be " chroococcoid ", but
this is uncertain ; heterocysts seem to be quite absent from the basal system.
Branches Nos. 12 and 14 (PI. 41, fig. 10) are dark like the basal system and are
probably branches of it. Unlike the erect branches they are crooked and lack hetero-
cysts. The status of branch No. i is uncertain. The cells are rather short in branch
No. 12 and a few of them are wedge-shaped. The distal portions of branches Nos.
i and 14 are narrow and composed of elongate cells similar to those of the hair-like
terminations of a few of the erect branches.
344 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
Erect system. — The erect filaments of Langiella are quite unbranched and essentially
uniseriate. Though generally curved or slightly sinuous they give the appearance of
being somewhat rigid. The longest filament measures about 300 //.
The cells of the filaments present some diversity of size and shape. Many of the cells
are slightly wider than long, others nearly twice as broad as long, while others are
depressed globose, cask-shaped or truncate ovoid. In some filaments there are
cells once to twice as long as broad, wedge-shaped cells also occur. The small cells
which are clearly heterocysts also show a diversity of form ; they are described in
detail below (p. 345). The trichomes are generally markedly torulose and may
be nearly moniliform. The two short branches, Nos. 4 and 9, are of nearly uniform
width throughout and the end cells are subspherical. The larger branches, however,
show a more or less clear differentiation into a proximal portion resembling these
two branches and a distal portion which usually has an elongate cell at the base.
The distal portions are of two main types. In one type it is hair-like and slightly
club-shaped with comparatively long and narrow cells giving the whole branch a
somewhat tapering form, e.g. Nos. 2 and 5. The other type is composed of cells which
are shorter and thinner-walled than those of the proximal portion and the apex
may include a few rather large cells or cells formed by oblique divisions, e.g. Nos.
3 and 7. There is a tendency for the end cells of all the filaments to be somewhat
longer than the penultimate cells and they are always rounded, never pointed or
attenuate. The sheath surrounding the proximal part is thin, usually fairly dark
and well defined and often slightly torulose. That around the distal cells, however,
is less distinct but thicker and at the thickest part the width of the filament may be
nearly twice that of the trichome (PI. 42, figs. 13, 15).
Basal portion of erect filaments. The catenate basal cells of some of the erect
filaments are large and torulose with well-defined walls and a dark narrow sheath.
A good example of this is seen in branch No. 8 (PI. 42, fig. 12) where the basal hetero-
cyst is succeeded by a row of seven cells which contrast rather sharply with the less
well-defined, less torulose and generally smaller cells of the distal portion. These
proximal cells have a diameter of about 10 /a and those of other filaments from 9 fi
to 12 /*. They recall the lower cells of the uniseriate filaments of certain Rivulariaceae,
lying next to the heterocyst which as in the fossil is not enclosed in the sheath. In
filament No. n the four cells lying between the two heterocysts are unusually large
and well defined, and the dark sheath closes round the end cells. The largest has a
diameter of 13 /i. It is possible that these are resting cells (akinetes) as developed
in e.g. Nodularia. They may, however, merely be thick-walled older cells like those
met in the filaments of Stigonema and Hapalosiphon. If they are resting cells it
seems possible that the basal cells of some of the other filaments may be incipient
akinetes.
Distal portions. The modifications shown by the distal portions of the filaments
have already been briefly noted. The hair-like form is well shown in filament No. n
(PI. 42, fig. 15) in which the end four cells are considerably smaller, though unchanged
in shape, giving a sudden reduction by about one-third in the width of the trichome.
The basal cell of the four is subcylindrical and about twice as long as broad, whereas
the remaining cells are subspherical. The hair-like terminal portions of branches
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 345
Nos. 2, 5 and 10 also have elongate basal cells and increase slightly in width towards
the apex where two or three rounded cells can be faintly seen. The cells in the middle
part of the trichome are long and attenuated. The sheath around these latter
trichomes is very faintly seen and is narrow, whilst that of filament No. n is more
distinct and also wider.
Branch No. 3 is an example where the trichome does not end in a hair-like portion.
The swollen apex which follows after a series of rather small cells with indistinct
walls, consists of perhaps six or eight cells formed partly by oblique divisions. Their
walls are not all distinct and it is impossible to say whether any of them are ruptured.
The sheath around the terminal portion of the trichome which is separated from the
proximal portion at an intercalary heterocyst is ill-defined but apparently of moderate
width. The last few cells of branches Nos. 6 and 7 include some wedge-shaped
sublateral cells probably indicating an early stage in the development of a multi-
cellular termination like that of branch No. 3. However there is a rather similar
wedge-shaped cell at the proximal end of branch No. 10 which can hardly be inter-
preted thus. At the end of branch No. 13 a short length of trichome, about 35 fi
long with five rather ill-defined partitions, appears to be detached from the rest of the
filament and to be emerging from the end of the sheath (PL 42, fig. 14). Its maximum
width (13 fi) is somewhat greater than that of most of the trichome. Proximal to it
is a shorter piece of trichome, apparently consisting of two cells, which is similar
to that near the end of filament No. 8. It is possible that these pieces of trichome,
especially the first, had a propagative function like that of hormogonia.
The sheaths of filaments Nos. 8 and 13 are rather wide and increase gradually
and markedly in width from the lowest cell of the terminal portion of the filament as
is well shown in PI. 42, fig. 13. A similar contrast between the sheaths of the basal
and of the terminal portion is shown by branch No. u. In each of these three
filaments the sheaths taper both proximally and distally and the widest part, where
the filament is nearly twice the width of the trichome, occurs nearer to the apex
than to the base. The hair-like termination of filament No. n is enclosed by the
sheath. In filament No. 8 (PL 42, figs. 13, 16) and more clearly in the isolated piece
of trichome shown in PL 43, fig. 23 there are indications of dark, more or less trans-
verse lines or bars in the sheath, the only indications of lamination to be observed.
Heterocysts. All the erect filaments, with the possible exception of No. 10, have a
heterocyst at or near the point of origin of the filament from the prostrate system.
In two branches (Nos. 3 and n) there is also a second, intercalary, heterocyst
near the middle of the branch. The basal heterocyst is usually more or less symmetri-
cally placed in relation to adjoining cells. In filament No. 4, however, the heterocyst
is somewhat wedge-shaped and in a rather lateral position. The heterocyst may be
ovoid to cylindrical, spherical or subquadrate, the long axis is usually parallel to
that of the trichome. Examples of cylindrical trichomes are seen in filament No. 2
and in the isolated piece of filament shown in PL 43, fig. 24. These heterocysts,
which are always distinctly smaller than the adjoining cells, range in diameter from
5 fi to 8 fi and in length from 5 fi to 10 (i. In several instances the heterocyst is broader
and flatter at the proximal pole. The heterocysts are characterized chiefly by the
hyaline nature of the wall, giving the appearance of a thin bright line. As may be
346 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
seen in PI. 43, fig. 22, the inner boundary of the wall is defined by a dark line, perhaps
formed by deposits on the wall. The outer boundary of the wall is more faintly
demarcated. In no case does the heterocyst appear to be enclosed in the sheath.
The absence of a sheath would no doubt facilitate fragmentation of the trichome at
the heterocyst. That fragmentation did occur is clearly shown by the isolated
filament in PI. 43, fig. 23 and by filament No. 2. This last example suggests that
fragmentation normally took place distal to the heterocyst and this is supported by
the observation that the distal end of the heterocyst is more often pointed than the
other. Branch No. 6 which has no basal heterocyst was quite possibly detached in
this way. The isolated filament in PI. 43, fig. 24 shows that the heterocyst may
permit considerable displacement of the filament without fragmentation occurring.
Cell contents. In some cells of a few branches of the erect system there are indica-
tions of cell contents. These consist of slightly darker central areas with a defined,
rounded outline which often follows the outline of the cell wall. These areas appear
minutely granular. The diameter is usually less than half that of the containing cell.
In several examples the area lies slightly to the left of the centre. In a cell near the
middle of filament No. 6 the faintly granular contents are but slightly contracted
away from the cell wall and there are also signs of partial contraction in the cell
next but one below it. The prominent black dendritic contents in branch No. 4
(PI. 43, fig. 19) largely disregard the boundaries between the cells. It is probable
that these, like the irregular black markings in a few other cells of the plant (PL 42,
figs. 12, 15, 16) are merely inorganic, probably pyritic, deposits like those scattered
through the matrix.
The heterocysts are scarcely paler than most of the other cells of the filaments. A
few of them have irregular black deposits on the inner side of the wall. The example
illustrated in PL 43, fig. 19 is the only heterocyst to show clear indications of a central
dark area. This also may be inorganic.
It should be noted that in Archaeothrix Kidston & Lang (1921 : 875) from the
same geological deposit the cell contents are also preserved.
Other specimens of Langiella. Closely associated with the holotype and clearly
belonging to the same species are numerous filaments, few of which can be seen
clearly. Most of them are uniseriate. A few are straight uniseriate filaments with
heterocysts like those of the erect system of the holotype (PL 43, figs. 23, 24). The
majority are more or less crooked (PL 41, figs. 3, 6, 7 ; PL 43, figs. 20, 21) and
resemble branches Nos. 12 and 14 of the holotype which are regarded as vegetative
branches of the prostrate system. They include small intercalary cells which,
because of their size, are possibly incipient heterocysts. But they lack the distinctive
characters of those of the erect system, especially a hyaline wall and elongation
parallel to the axis of the filament. Also they are more or less clearly enclosed in
the sheath, which is usually rather well developed. A few of the cells show cell
contents as a dark central area, as in the holotype. These additional specimens
do not include any cells which can definitely be recognized as akinetes or hormogonia
and do not give any further evidence as to the methods of propagation or of the growth
of the plant. They do, however, provide additional evidence of the modes of cell
division and of branching.
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 347
The smallest group of cells seen, consisting of a row of three cells of graded size
with a definite sheath, is shown in PI. 41, fig. 5 ; whether they were formed by germina-
tion of a spore or by fragmentation is uncertain. PI. 41, fig. 8 shows a short row of
rather wide cells up to 16 /* in diameter. The small basal cell, which does not appear
to be enclosed in the sheath is probably a heterocyst. Of particular interest is the
fact, clearly demonstrated by focussing at different planes, that the distal cell has
undergone a longitudinal division, and one of the daughter cells is further divided
in a plane at right angles to the previous division. The piece of filament in PI. 41,
fig. 4 shows at its widest part a pair of collateral cells, presumably formed by division
of an apical cell rather than by longitudinal division of an intercalary cell. The bent
piece of filament in PI. 41, fig. 7 shows a small cell at the bend formed by oblique
division. The specimen in PI. 41, fig. 6 and PI. 43, fig. 21 appears to consist of two
short filaments crossing one another. The large dark-walled cell near the centre may
possibly be an akinete which has germinated. Lastly in PI. 41, fig. 3 the upper filament
appears to be branching out of the more horizontal filament. The downwardly
directed filament appears to originate at the junction of the filaments. The upper
filament shows what is perhaps an incipient heterocyst where the filament bends.
DISCUSSION. As is general in the filamentous blue-green algae, the cell division in
Langiella is predominantly transverse. In the Stigonematales, where true branching
occurs, the branches arise from the longitudinal division either of an apical cell or
of an intercalary cell. Branches produced in this latter way push through the sheath
of the filament and increase in length by the usual transverse divisions. In the holotype
and the other fossil material examined this type of longitudinal division of an inter-
calary cell is nowhere conclusively shown. Presumably such must have occurred
both to increase the thickness of the prostrate axis and to produce the numerous
erect filaments arising from it. It is considered unlikely though not impossible that
all these arose from apical cells.
Elongation of the erect filaments was probably in part at least from intercalary
divisions in the distal portion of the filaments. In branches Nos. 3, 7 and 8 a series of
short cells suggests recent cell divisions, such cells are not found in the proximal,
presumably older, parts of the filaments. The subterminal lengths without cross-walls
in filaments Nos. 2, 5 and 10 recall the filaments of Recent species of Fischerella
(Geitler, 1932, text-figs. 296-299) where hormogonia are in the process of leaving the
sheath of the parent filament. However it seems that this is not the case here, as the
non-septate portion appears not to be an empty space but more like an attenuated
cell. The possibility of necrotic lengths of trichome must also be considered,
especially as Kidston & Lang (1921 : 835) report necrotic and wound areas in Rhynia.
An interesting feature clearly shown by the holotype and other specimens is the
prevalence of wedge-shaped cells, often small and lying at a bend of the filament ;
they were presumably formed by the oblique division of an apical or perhaps an
intercalary cell. This is especially well shown by the ends of branches Nos. 6 and 7
in the holotype where several cells are cut off by oblique walls.
348 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
Genus KIDSTONIELLA nov.
DIAGNOSIS. Thallus heterotrichous, epiphytic. Basal system consisting of a mass
of cells, some of which are polygonal, forming thin parenchymatous tissue, others
larger, rounded and thicker-walled. Erect branches uniseriate, straight, more or
less strongly torulose, once or twice unequally branched by longitudinal division of
the apical cell, or unbranched. Intercalary transverse divisions rare or absent.
Cells variable in shape and size, mostly somewhat wider than long, end cells sub-
spherical. Sheath seldom well preserved. Heterocysts small, nearly spherical,
intercalary in erect filaments. Cell contents conspicuous in many cells of the thallus.
Method of reproduction uncertain.
Cells of pseudoparenchymatous tissue of basal system 10 fi to 14 /i broad ; large
rounded cells of basal system 16 fi broad ; erect filaments up to 160 ju, long and up
to 24 jn broad ; length of mature cells 5 //. to 14 /i ; heterocysts 5 /* to 6 /* long and
6 fi to 7 ju, broad.
TYPE SPECIES. Kidstoniella fritschi n. sp.
Kidstoniella fritschi n. sp.
(PI. 41, ngs. 2, 9 ; PI. 44)
DIAGNOSIS. As for genus.
HOLOTYPE. Specimen No. ¥.32409, Palaeont. Dept. Brit. Mus. (Nat. Hist.).
OCCURRENCE. Middle Devonian (Old Red Sandstone) ; Muir of Rhynie, Aber-
deenshire, Scotland.
GENERAL MORPHOLOGY. As noted above (p. 342), this species appears to have
grown epiphytically. PI. 44, fig. 29 shows a dark band of multiseriate filaments
forming a basal layer lying parallel to, but partly detached from the edge of an
oblique section through a nearly opaque plant axis. A number of uniseriate filaments,
a few of which are branched, arise from this basal layer and stand away from it at
a wide angle. It seems likely that the axis, perhaps growing more or less erect,
formed the substrate for the heterotrichous thallus of the alga.
Basal system. Much of the basal system shown in PI. 41, fig. 9 is composed of a
dense mass of rather ill-defined cells. Some of these are rather large, spherical or
ovate, with thick walls and clear indications of dark rounded contents. These cells
attain a diameter of 15 fi. In part of the basal system a number of cells which are
smaller than those forming the erect filaments have become polygonal through
mutual pressure and form a thin pseudoparenchymatous tissue which in part at
least seems to be monostromatic. Another part of the basal system is formed of
imperfectly seen rounded cells up to 16 jLt in diameter. There is little or no evidence
of any mucilaginous sheath around the basal system.
Erect system. The more or less erect branches are essentially uniseriate and often
strongly torulose. The cells vary considerably in shape and size. They are mostly
somewhat wider than long, and may be twice as wide as long, ranging in shape
from discoid or wedge-shaped to depressed — or truncate-globose. A few are slightly
longer than wide, cask-shaped or ovoid. The end cells are rounded and when mature,
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 349
subspherical. The filaments range in width from about 8 /* to 14 /i. An unusually
large cell in filament No. i is 15 ju, in diameter.
A feature of special interest is the unequal dichotomous branching shown by two
of the erect filaments. In filament No. i the two divisions are but slightly divergent.
The shorter and narrower division consisting of shorter but more numerous cells,
is less developed than the other. In filament No. 2 the very short right-hand division
is indistinctly preserved but appears to be multiseriate. The terminal cell of the left-
hand division is itself divided by a faint but definite longitudinal wall (PI. 44, fig. 27).
From the fortunate preservation of the filament at this stage of growth it may be
inferred that the other dichotomous divisions have also resulted from the longitudinal
division of an apical cell. There is no clear indication in the thallus of the longitudinal
or transverse division of an intercalary cell and it might therefore be concluded that
growth was wholly apical. However, in filaments Nos. i and 4 there are rather short
cells which might be products of recent intercalary divisions. The large end cell of
filament No. 2 seems to be in process of dividing longitudinally, for the contents
consist of two rounded bodies in place of the usual one. In the left-hand filament in
PI. 44, fig. 25 the distal end of the filament is divided into a number of imperfectly
seen cells somewhat reminiscent of the multicellular tips seen in Langiella. The large
end cell of the next filament seems to be in the process of dividing longitudinally
for two rounded bodies are seen inside it. Springing from the right-hand side of
this filament is a short club-shaped body whose contents are divided into three or four
very small portions not obviously separated by septa. This appears to be an incipient
branch and may be compared with the branch of very short cells in filament No. i.
The pair of cells just below the branch in filament No. 2 may have resulted from a
transverse intercalary division.
The erect filaments have few and discontinuous pieces of a surrounding sheath.
The clearest examples are round the distal parts of filaments Nos. 2 and 3 and around
filament No. 4.
Hderocysts. The small nearly spherical cells indicated in filaments Nos. 3 and 4
are possibly heterocysts. Their diameters are about 6 /*. Like those in Langiella
they are small and have a hyaline wall delimited by dark lines of which the inner
seems to be the darker. A few of the normal sized cells, for instance those on each side
of the heterocyst in filament No. 4, also have hyaline walls, though in these the outer
limiting line seems to be the darker. Most of the cells having a thin dark wall in
optical section, show a similar shining wall at a slightly higher focus. In shape and
appearance these two cells resemble the small intercalary cells thought to be hetero-
cysts in some of the filaments of Langiella.
In PI. 44, fig. 27 several of the cells, notably in filament No. 3, show a clear wall
bounded internally by a fine dark line. The small intercalary cell in this filament
recalls by its size the cells which are regarded doubtfully as heterocysts in filaments
of Langiella. The wall, however, is not hyaline, cell contents are present and the
sheath appears to enclose it. This may well be an example of a small cell which has
not yet developed into a heterocyst.
Cell contents. In most of the cells in the uniseriate filaments, as well as in the basal
region, there are more or less clear indications of contents (PI. 44, figs. 27-29).
350 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
As in Langiella, these consist of darker, central areas with a rounded outline. As
a rule the larger bodies are found in the larger cells. The shape of the central area
corresponds generally with that of the cell, being wedge-shaped in wedge-shaped
cells and transversely elongate in discoidal cells. The central areas are particularly
well denned in the distal regions of filaments Nos. I and 2. In the main division of
branch No. i the central areas of four cells are more or less sharply delimited and
surrounded by a narrow clear zone or halo which is paler than the outer zone. In
certain cells in filament No. 2 the clear zone appears to be delimited from the outer,
slightly dark, contents of the cell by a fine line. This suggests that the zones outside
the central dark area may represent mucilaginous or other contents of the cell. With
the highest resolution which could be used the dark central areas appear to be finely
granular. The cell immediately below the dichotomy in filament No. i appears to
be empty and to be ruptured on its left side.
MYXOPHYCEAE
INCERTAE SEDIS
Genus RHYNIELLA nov.
DIAGNOSIS. Thallus filamentous, essentially uniseriate, vermiform, non-hetero-
trichous. Trichomes mainly sinuous within a mucilage sheath, often two or three
sharing a common sheath, branching doubtful, heterocysts doubtful, some with
multicellular apices. Cells usually broader than long, wedge-shaped at the sharper
bends in the trichome. Terminal cells rounded or rarely bluntly pointed. Method
of reproduction unknown.
Trichome up to 140 fi long ; cells 4 ju, to 10 fi long and 10 fi to 14 fi broad.
TYPE SPECIES. Rhyniella vermiformis n. sp.
Rhyniella vermiformis n. sp.
(PI. 41, ngs. i, ii ; PI. 43, fig. 18)
DIAGNOSIS. As for genus.
HOLOTYPE. Specimen No. ¥.32409, Palaeont. Dept. Brit. Mus. (Nat. Hist.).
OCCURRENCE. Middle Devonian (Old Red Sandstone) ; Muir of Rhynie, Aber-
deenshire, Scotland.
GENERAL MORPHOLOGY. The species represented in PI. 41, fig. n is found near the
middle of the upper edge of the chip in PL 41, fig. i and does not appear to be present
elsewhere in the chip. It consists of numerous subparallel trichomes many of which are
distinctly vermiform. The drawing is considerably condensed, but the thickness of the
bundle of trichomes is much less than the width. The cells are typically broader than
long ; wedge-shaped cells are found at the sharper bends in the trichome. Several
trichomes are more or less isolated and enclosed in their own sheaths. Some of the
trichomes, however, are closely associated for part of their length and it is practically
certain that they share a common sheath. No clear case of branching has been
observed. Nor are heterocysts certainly present, though one cell may be of this type.
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 351
There is no defined basal system ; the multicellular, and presumably distal, termina-
tions of many of the filaments are, however, with one exception similarly orientated
and the direction of growth was probably to the right in PI. 41, fig. n.
Trichomes. Some of the more distinct trichomes attain a length of 140 ju,, and
vary in width from 10 fi to 14 fi. A few of the indistinctly seen trichomes are nearly
straight, but the majority are sinuous, and a few are bent at right angles. All but the
terminal cells are shorter than wide and many are twice as wide as long. At the
bends in the trichome the cells are more or less markedly wedge-shaped, in some
instances not reaching across the full width of the trichome. There are generally
distinct constrictions at the septa. The end cells vary considerably in shape, they
may be segmental, hemispherical, subspherical or spherical. If spherical two or
more may form a group. A few of the hemispherical cells are bluntly pointed. At
A (PL 41, fig. n) a trichome is terminated by an unequal pair of rounded cells :
and at B the penultimate cell is divided longitudinally. At c an unusually large,
oval-shaped end cell (length 20 ju, ; breadth 14 fi) appears to be dividing. The various
groups of terminal cells may have had a propagative function.
To the left of the centre of PL 41, fig. n there is a small cell almost surrounded
by a curved piece of trichome. This cell is subquadrangular and measures 5 /*by
6 /I. It is distinctive not only because of its size and shape but because the wall
is hyaline and marked on its inner boundary by a heavy dark line. These features
characterize those cells of Langiella which are confidently regarded as heterocysts ;
this cell must therefore be regarded as quite possibly a heterocyst. Its position in
the thallus is, however, unusual and its true nature is uncertain.
Some of the trichomes are divided into lengths composed of from 4 to 12 cells. The
longest piece of trichome without such a segmentation has 18 cells. The end cells
of each segment are usually hemispherical or subspherical. There may be a small,
nearly spherical cell connecting two segments as in the central trichome in the lower
part of PL 41, fig. ii. These small cells do not have a hyaline wall.
In a few places, the trichomes are closely parallel for part of their length indicating
that they share a common sheath. The mode of branching, if this does occur, is not
clearly demonstrated at any point.
Cell contents. As indicated in PL 41, fig. n a few of the cells show rounded contents.
Though indistinct, they do not appear to differ significantly from those in the associated
species.
Sheaths. The trichomes are enclosed in more or less clearly defined sheaths which
vary in thickness from a quarter to a half of the trichome width. In one or two
places where the edge of the sheath is clearly defined it is irregularly sinuous. The
sheaths also show indications of longitudinal folds. Two of the more isolated trichomes
are completely enclosed in their own individual sheaths. The majority of the trichomes
appear to share a common sheath, at least over part of their length.
DISCUSSION AND COMPARISON. Rhyniella differs from the other two genera in
the apparent absence of heterotrichy, of heterocysts and in its shorter and less
rounded cells. It calls to mind forms like Tolypothrix elenkinii and Scytonema
(Diplocolon) crustaceum (Geitler, 1932 : 738, 784). Both these Recent species are
heterocystous and non-heterotrichous with many contorted trichomes in a common
352 BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE
sheath. However the possibility must not be overlooked that this specimen grew
from a basal system which is not now in evidence. From its general appearance and
cell morphology it could not be part of either Langiella or Kidstoniella which,
however, it resembles in the possession of multicellular apices. Nothing quite like
these structures occurs in present-day members of the class.
IV. CONCLUSIONS AND GENERAL DISCUSSION
It is now evident that in Middle Devonian times the blue-green algae had reached
an evolutionary grade which is generally regarded as advanced among Recent forms,
namely a heterotrichous thallus with a pseudoparenchymatous basal system. The
enigma of the heterocyst, so well presented by Fritsch (1951) is no nearer solution,
but it is clear that the evolutionary origin of the heterocyst must be sought in still
older fossils.
The multicellular tips to the trichomes seen in Langiella and Rhyniella and rarely
in Kidstoniella are of considerable interest in that they have no close parallel in
present-day forms. Rather similar structures in Stigonema dedroideum were described
by Fr6my (1930) as conidia, but there the cells are rounded and separated within
mucilage. It is of course possible that the fossil structures are an early stage, either
phylogenetically or ontogenetically, in the development of such conidia.
There is a certain similarity in general appearance between Langiella and Kidstoniella
but it is considered most unlikely that they belong to the one species or even the same
genus. Langiella has unbranched erect filaments with distinct heterocysts and
occasional multicellular tips, while Kidstoniella shows true branching of the erect
filaments which have few multicellular tips and doubtful heterocysts. How constant
these characters were in the living population one cannot say but on the present
evidence and in accordance with taxonomic practice, both in phycology and in
palaeobotany they must be considered separate genera. There can be little doubt
that these two genera belong in the Stigonematales (Geitler, 1925; Fritsch, 1942)
and there are no characters on which one could satisfactorily found a new family
apart from the Stigonemataceae.
The position of Rhyniella is less certain. On the present probably incomplete
evidence it is non-heterotrichous and unbranched, suggesting a position in the
Nostocales (Geitler, 1925 ; Fritsch, 1942) and perhaps within Oscillatoriaceae
but the presence of multicellular tips as in Langiella suggests Stigonematales and
possibly a close relationship with Langiella. This difficulty in assigning a taxonomic
position to a blue-green alga is by no means limited to fossil forms ; there is wide
lack of agreement over the classification of present-day forms (see Papenfuss, 1955).
V. REFERENCES
FR^MY, P. 1930. Les Myxophycees de 1'Afrique equatoriale fran9aise. Arch. Bot. Mem.,
Caen, 3 : 1-508, 362 figs.
FRITSCH, F. E. 1942. The interrelations and classification of the Myxophyceae (Cyano-
phyceae). New Phytol., Cambridge, 41 : 134-148, 5 figs.
1945- The Structure and Reproduction of the Algae, 2. xii + 939 pp., 336 figs. Cambridge.
1951- The Heterocyst : A Botanical Enigma. Pvoc. Linn. Soc. Lend., 162, 2 : 194-211,
pl-5-
BLUE-GREEN ALGAE FROM THE MIDDLE DEVONIAN OF RHYNIE 353
GEITLER, L. 1925. Synoptische Darstellung der Cyanophyceen in morphologischer und
systematischer Hinsicht. Beih. hot. Zbl., Cassel (2) 41, 3 : 163-294, pis. 17-20.
1932. Cyanophyceae. In Rabenhorst, L. Kryptogammenflora, 14: 1196 pp., 780 figs.
Leipzig.
KIDSTON, R. & LANG, W. H. 1921. On Old Red Sandstone Plants showing structure, from
the Rhynie Chert Bed, Aberdeenshire, V. The Thallophyta occurring in the Peat-Bed,
the Succession of the Plants throughout a Vertical Section of the Bed ; and the Conditions
of Accumulation and Preservation of the Deposit. Trans. Roy. Soc. Edinb., 52 : 855-902,
pis. i-io.
PAPENFUSS, G. F. 1955. A Century of Progress in the Natural Sciences, 1853-1953 :
pp. 115-224. San Francisco.
SCOURFIELD, D. J. 1926. On a New Type of Crustacean from the Old Red Sandstone (Rhynie
Chert Bed, Aberdeenshire) — Lepidocaris rhyniensis gen. et sp. nov. Philos. Trans.,
London (B) 214 : 153-187, pis. 21-23.
PLATE 41
Langiella scourfieldi, Kidstoniella fritschi and Rhyniella vermiformis
FIG. r. Drawing of the chip showing position of the fossils, in particular Langiella scourfieldi
(top right-hand corner) and Rhyniella vermiformis (centre), x 15 (V.324OQ).
FIG. 2. The same, reverse side, to show position of Kidstoniella fritschi. x 15.
FIGS. 3-8. Drawings of isolated filaments of Langiella scourfieldi associated with the
holotype. x 280.
FIG. 9. Drawing of the holotype of Kidstoniella fritschi constructed from photographs and
direct observation, x 280.
FIG. 10. Drawing of the holotype of Langiella scourfieldi constructed chiefly from photo-
graphs at different focal levels, but partly from direct observation, x 280.
FIG. ii. Drawing of the holotype of Rhyniella vermiformis from photographs and direct
observation, x 280.
Bull. B.M. (N.H.} Geol. 3, 10
PLATE 41
PLATE 42
Langiella scourfieldi sp. nov.
FIG. 12. A supposed akinete near the base of filament No. 4 (latter not in focus) and pyritic
granules in filament No. 7 ; also filament No. 10. x 230 approx.
FIG. 13. Filaments Nos. 4 and 8. x 230 approx.
FIG. 14. Terminal portion of filament No. 13. x 560.
Fig. 15. Filaments Nos. 9 and n. x 230 approx.
FIG. 16. Filaments Nos. 4 and 8 and cells of the basal system, x 375 approx.
Bull. B.M. (N.H.) Geol. 3, 10
PLATE 42
PLATE 43
Langiella scourfieldi and Rhyniella vermiformis
FIG. 17. Holotype of Langiella scourfieldi showing habit. x 44.
FIG. 1 8. Holotype of Rhyniella vermiformis. x 375 approx.
FIG. 19. Langiella scourfieldi showing branches Nos. 4 and 8. x 560.
FIGS. 20, 21. Isolated filaments of Langiella scourfieldi. Fig. 20 x 105, Fig. 21 x 185.
FIG. 22. Langiella scourfieldi. Two heterocysts in filament No. n and one in filament
No. 9. x 560.
FIG. 23. Langiella scourfieldi. Isolated piece of erect filament showing break at the hetero-
cyst and laminated sheath, x 560.
FIG. 24. Langiella scourfieldi. Isolated fragment of filament with a cylindrical heterocyst
showing displacement at the heterocyst. x 560.
Bull. B.M. (N.H.) Geol. 3, 10
PLATE 43
24
PLATE 44
Kidstoniella fritschi sp. nov.
FIG. 25. Drawing of paratype. The distal end of the left-hand filament is divided into
a number of obscurely seen cells, x 280.
FIG. 26. Holotype. Right branch of filament No. i and a divided apical cell of filament
No. 2. Note the granular contents. x 435 approx.
FIG. 27. Photograph showing general features of holotype including the basal system and
granular cell contents. x 435 approx.
FIG. 28. The left branch of filament No. i and the apical cell of the right branch. x 435
approx.
FIG. 29. Photograph showing habit. x 44.
Bull. B.M. (N.H.) Geol. 3, 10
PLATE 44
INDEX TO VOLUME III
New taxonomic names and the page numbers of the principal references are printed in Clarendon type.
An asterisk (*) indicates a figure.
Acanthocythere i, 7, 12
sphaerulata 12, 13 ; PI. i, figs. 1-4
spinisculata 13 ; PI. i, figs. 5-9
Acrilla 25, 34
affinis 34
nigeriensis 25, 34 ; PL 5, fig. 6
Admete 49
Admetula 27, 49
Africa 127-175, 179-189
Africarca 27, 53
nigeriensis 27, 53 ; PI. 8, fig. ^a, b
Africofragum 28, 61
Africolithes 26, 45
Africostoma 27, 49
Africosveltia 27, 48
Africoterebellum 26, 37, 38
Alethopteris goepperti 297, 315
martinsi 313
Algae 307, 341
Algites 309, 310
sternbergianus 298, 299, 304, 310 ; PI. 36, figs.
i, 2, 5
virgatus 299, 303, 305, 309*. 310
Ambonychia obliqua 222
orbicularis 222
sp. 218, 221
Amekichilus 26, 38
suturocostatum 26, 39
Amekicythara 27, 52
douvillei 27, 53 ; PI. 8, fig. za-c
Amekiglans 28, 58
American Museum 286
Amphicythere 5, 8, 14
semisulcata 14
Amphiperas nigeriensis 39
Amphiperatidae 26
Amphorispermum 169
Anacardiaceae 75
Anachis 41
Anadara nigeriensis 54
Anatina 67
Anemia 75
Angiopteridium mcclellandi 154
spathulatum 154
Annularia 298, 303, 312, 313*
Araucaria 330
Araucarites 330
ludwigi 330
Archaeopodocarpus 304, 330
germanicus 324
Archaeothrix 341, 346
Archidiskodon 277-279, 289, 290
exoptatus 278
meridionalis 279
planifrons 267, 270-28o, 289 ; PI. 32 ; PI. 34,
figs. 3, 4 ; PL 35, figs, i, 2
nyanzae 278
proplanifrons 277, 278
subplanifrons 277-279
Archinacella oblongata 223
Architectonica 25, 32
affinis 35
canaliculata 33
(Nipteraxis) bendeica 25, 32 ; PL 5, fig. 3a-c
(Solariaxis) amekiensis 25, 33 ; PL 5, fig. 5«-c
spectabilis 34
(Stellaxis) bicingulata 25, 33 ; PL 5, fig. 4
Architectonicidae 25, 32
Arcopsis 27, 53, 54
africana 27, 54 ; PL 8, fig. 4
Artocarpoideae 95
Asaphid 222
Aspidites danaeoides 151
Asterocalamites 312
mansfeldicus 312
Asterophyllites mansfeldicus 312
Asthenotoma 52
(Endiotoma) casteri 51, 52
Athleta 44
lugardi 44
Atractopyge 211, 212
Australia 127, 145, 163, 165, 172, 179, 187, 188,
235
Baiera digitata 319
munsteriana 319
Bairdiacea 6, u
Bairdiidae n
Bairdoppilata 6
Bancroftina 207, 221
robusta 207, 221
ty^>a 199, 206, 220, 221 ; PL 24, figs. 17, 18
Bardia 319
Basilicus marstoni 219
BATE, D.M.A. 268, 289
Belosepia 29
356
INDEX
Bendeglans 28, 57
Bendeia 26, 43
africana 26, 43 ; PI. 7, fig. 2a, b
Bendeluta 26, 46
conicoturrita 26, 47
Bendemacoma 28, 66
Bethlehem Fauna 267, 289
Betulaceae 76, 92
Beyrichia 211, 224
Bjuvea 151, 172
Bonellitia 27, 49, 50
evulsa 49
(Admetula) amekiensis 27, 49 ; PI. 7, fig. 5#-c
evulsa 49
(Africostoma) decor ata 27, 50 ; PI. 7, fig. 6a— c
Boraginaceae 103
Bos 267
Bovey Tracey Flora 71-123
Bovidae 285
Brachyopid skull 259, 260*
Bradleya 6
Brasenia 74, 96
ovula 77, 79, 96 ; PI. 13, fig. 75
Breviarca 55
Broeggeria 203
soudleyensis 203
Broeggerolithus 202—204, 207, 208, 210, 228
broeggeri 199, 203, 204, 216, 220 ; PI. 26, fig. 7
constrictus 220
globiceps 199, 206, 221 ; PI. 26, fig. 9
longiceps 199, 207, 222 ; PI. 26, fig. 10
soudleyensis 199, 204, 207, 216, 220, 222 ;
PI. 26, fig. 8
transiens 199, 208, 209, 223, 226 ; PI. 26,
fig. ii
Sp. 22O, 221
Brongniartella 198, 203, 206, 228
bisulcata 207, 209, 210, 216, 218, 221-223
sp. 219, 220
Buccinidae 26, 43
Buccinorbis 26, 29, 47
kitsoni 47
Bucklandia 100
Bufina 10, n ; PI. 2, figs. 3, 4
Bulbifusus 45
nigeriensis 44, 45
Bulbil 121,* 122 ; PI. 17, fig. 204
Bythocythere 7
Caestocorbula 68
Calameae (Palmae) 76, 89
Calamites 296, 298, 306, 311, 312
arenaceus 312
kutorgae 312
wakei 296, 312
Calamus 73-75, 78, 88, 89
daemonorops 76, 88, 89 ; PI. 12, figs. 24-44
Calathella 295, 307, 308, 310
dictyonemoides 301, 304, 308*; PL 36, fig. 4
Calathella — contd.
krduseli 305, 308
Calla palustris 75
Callipteris 313
martinsi 300-306, 313*, 314*, 315 ; PI. 37,
figs. 2, 5
Callista 64
elongatotrigona 63
kitsoni 64
Calvarinus 103
reticulatus 102
Calyptaulax 211, 212, 224
Calyptraea 25, 35
crepidularis 35
newtoni 25, 35 ; PI. 5, fig. 7a, 6
Calyptraeidae 25, 35
Camptocy there 5*, 7
Cancellaria 48
multiplicis 48, 49
Cancellariidae 27, 48
Canidae 268
Canis (Nyctereutes) sinensis 269
Capitosaurus 236, 254, 257
Capparidaceae 77, 98, 99
Capparidispermum 98, 99
boveyanum 77, 98 ; PI. 14, figs. 92-96
Caprifoliaceae 75
Capsicum 119
Caradoc Series 193-230 (S. Shropshire)
Cardiidae 28, 61
Cardiocardita 60
Cardiocarpon triangular e 297, 302, 330
Cardiocarpus 187
Cardita aegyptiaca 59
beaumonti 60
costaeirregularis 57, 58
costaenodulosis 58, 59
fayumensis 59
planicosta 57
triparticostata 60
(Cyclocardia) granulata 58
Carditidae 27, 57
Cardium obliquum 61, 62
Caricoidea 74, 86
nitens 76, 87*; PL n, figs. 20-23
obscura 79
Caricoideae 86, 88
Carinopsis gracilis 219, 220
Carpinus 74, 78, 90—93
boveyanus 76, 79, 92 ; PL 13, figs. 52-67
Carpolithus boveyanus go, 92, 93
exaratus 102
frumentarius 324
nitens 87, 88
rotherianus 305
websteri 86
spp. 118-121 ; PL 17, figs. 192-203
Caulerpa selaginoides 296, 298, 299, 324
Caulerpites bipinnatus 315
selaginoides 297
INDEX
357
Caytonia 169
Celtidoideae 94
Ceratotheca subuncta 224
Cervidae 267
Chamaerops teutonica 89
CHANDLER, M. E. J. 71-123
Chasmops 207, 209-213, 222-224, 229
Chionella 28, 63
(Costacallista) elongatotrigona 28, 63
(Microcallista) kitsoni 28, 64 ; PI. 10, figs.
•za, b, 3
Chiralithes 45
Chondrites 309
virgatus 296, 309-311
Chondrus binneyi 296, 300, 332 ; PI. 40, fig. 13
Chonetoidea 211, 213, 219, 223, 224
Cinnamomum 97
lanceolatum 97
rossmassleri 97
scheuchzeri 97
Cladium mariscus 88
Clathrospira trochiformis 223
sp. 221
Clavagellacea 68
Clavagellidae 68
Clavilithes 26, 45
(Africolithes) africanus 26, 45
Cliftonia andersoni 218-220
persculpta 203, 204.
spiriferoides 220
Climacograptus antiquus 220, 228
brevis 227, 228
caudatus 227
minimus 227
peltifer 226
wilsoni 226
sp. 225
Clinura 27, 51
Coeloconica 25, 32
Coenites 211, 223
Colpomya 210, 224
Cominella 43, 52
douvillei 52, 53
Concavimargo 25, 36
Conidae 27, 50
Conites 165, 329
sp. 301, 303, 329; PI. 40, fig. 5
Conospirus parisiensis 50
Conularia sp. 219, 222
Conus 27, 50
parisiensis 51
(Leptoconus) amekiensis 27, 50 ; PI. 7, fig.
Coralliophaga 69
eocenica 68, 69
Corbula rugosa 67, 68
Corbulidae 28, 67
Cordaites 188, 300, 321
aequalis 295, 299, 303, 306, 321 ; PI. 38, fig. 3
Cordaites — contd.
pangerti 305
Cordiopsis 64, 65
incrassata 63-65
Cornaceae 114
Cornulina 46
WMwa.* 46
Cornus 75
Corylopsis 100
Corylus sp. 76, 92 ; PI. 13, figs. 49-51
Cosmannella 59
costaenodulosis 59
Costacallista 28, 62-64
Costonia 198
ultima 197-200, 218 ; PI. 26, figs, i, 2
sp. 200, 218
Craniops sp. 220, 222
Crassostrea 27, 56
lugardi 56
Crepidula 25, 36
(Concavimargo) falconeri 36
Crepispisula 28, 65
CROFT, W. N. 339-353
Cryptolithus 200, 203
broeggeri 203
soudleyensis 207
Cryptothyris 211
paracyclica 199, 211, 212, 223 ; PI. 25, fig. 7
Ctenis permiana 296, 321
renaulti 323
Ctenodonta coarctata 219
varicosa 218, 219, 222, 224
sp. 220-222, 224
Cupressites ullmanni 297
Cupressus ullmanni 323
Cupuladria 29
Cyclocardia 58
borealis 58
costaeirregularis 58
Cyclocarpon eiselianum 305
triangulare 330
Cyclocarpus spongioides 306
Cyclonema crebristria 218, 220
Cyclopteris liebeana 327
Cyclotosaurus 236, 254, 256—258
ebrachensis 256-258
hemprichi 256-258
mordax 256, 257
papilio 256
posthumus 256, 257
randalli 257
robustus 256, 257
spitzbergensis 257
stantonensis 254, 256, 257
Cylindr acanthus 29
Cymatiidae 26, 40
Cyperaceae 74, 76, 79, 86, 88
Cyphomandra 119
Cypraea bowerbanki 39
Cypridacea 6, n
358 INDEX
Cypridea 7, 15, 17
fasciculata 16, 17
sowerbyi 15
Cyprideis 8
Cyrtoceras 224
Cyrtolites nodosus 220-223
sp. 219
Cyrtulotibia 25, 36
unidigitata 25, 37
Cyrtulus 36
Cythara 52
Cytharacea i, 6, 8-13, 16
Cythere 7, 12, 13, 18
retirugata 13, 15
var. decorata 17
rugulata 18
textilis 15
sphaerulata 12
transiens 19
Cythereis 6, 9
Cytherellidae 10
Cytheridea 8
Cytherideinae 8
Daemonorops 89
Dalmanella 208
horderleyensis 199, 206, 220 ; PI. 24, figs. 12, 13
indica 199, 206, 220 ; PI. 24, fig. 14
te^a 199, 206, 220 ; PI. 24, figs. 15, 16
multiplicata 222
unguis 199, 209, 222, 226 ; PI. 25, fig. 4
wattsi 199, 209, 222 ; PI. 25, figs. 2, 3
sp. 218, 219
Daphnogene ungeri 97
DEAN, W. T. 191-231
Decoroproetus 200, 220
fearnsidesi 219
(Proetidella) 198
fearnsidesi 201
Dendrograptus sp. 227
Diacalymene praecox 219
Dicellograptus complanatus 229, 230
Dicerorhinus 290
etruscus 281-283, 289 ; PI. 33, fig. 5
Diclidocarya menzeli 107
Dicoryphe 100
Dicranograptus clingani 214, 226-230
Dictyocythere i, 5, 8, 13-15
decorata 14
mediostricta 14, 15
retirugata 14-17
var. textilis 16
rugulata 14
transiens 14, 15
(Dictyocythere) decorata 17 ; PI. 3, fig. i
mediostricta 16 ; PI. 3, figs. 2-6
retirugata 15 ; PI. 3, figs. 7-10 ; PI. 4, figs.
3, 4, ii, 16, 17
(Rhysocy there) regulata 15, 18, 19 ; PI. 4,
Dictyocythere — contd.
figs, i, 2, 5-10, 12-15
transiens 19 ; PI. 3, figs. 11-13
Dictyonema 228, 309
fluitans 227, 228
Dictyopteris simplex 145, 148
Dientomochilus 38
Dinorthis 204
flabellulum 197, 199, 218, 220 ; PI. 24, fig. 2
multiplicata 203, 204
robust a 198
sp. 203, 218-220
Diplograptus multidens 214, 226-229
multidens var. compactus 227
sp. 224, 230
Distylium 100
Divaricella oppenheimi 61
Divergidens 28, 60
Dolerorthis 211
duftonensis 199, 207, 221 ; PI. 25, fig. i.
Sp. 2I8-22O, 222, 223
Doratophyllum 151, 172
EAMES, F. E. 23-70
Ecculiomphalus sp. 223
Ectinochilus 38
Elephantidae 270
Elephas 267
africanavus 278
Encrinurus 209
sp. 223
Endiatoma 52
Engelhardtia 75
Eobronteus sp. 224
Eocypraea (Sphaerocypraea) bowerbanki var.
sudanensis 39
Eodivaricella 28, 61
Eohomalonotus 198, 206
sp. 218, 221
Eomastixia 79, 121
bilocularis 121
Eomiltha 28, 61
Eopleurotoma 27, 51
nigeriensis 27, 51
Eovolva 26, 39
nigeriensis 26, 39 ; PI. 6, figs. 2, 3
Epimachairodus 270
Equidae 280
Equus 267, 280, 281, 290
stenonis 280
Eretmonia 187
natalensis 165, 187
Ericaceae 75
Eryops 239, 245, 248, 251, 252
Euglyphella 9-11 ; PI. 2, figs. 5-7
Exechostoma 25, 29, 30
cossmanni 25, 30
Fagaceae 76, 93
INDEX
359
Fagus 93, 94
minima 76, 93 ; PI. 13, figs. 68-70
Fascioplex 46
Favosites 211
fibrilla 218, 220-223
Felidae 270
Felis sp. 267
Ficus 52
Filograna 311
permiana 296, 297, 303, 306, 310, 311
Fischerella 347
Flexicalymene acantha 218, 219
caractaci 222-224
laticeps 224
onniensis 224
salteri 224
trigonoceps 223
sp. 218, 219, 223
FLORIN, R. 301, 327
Fortunearia 100
Fossularca 54
africana 54
F other gilla 100
Fragum 28, 6 1
(Africofragum) newtoni 28, 61 ; PI. 9, fig.
8a-c
Fucoides digit atus 319
selaginoides 324
Fusinidae 26, 45
Gangamopteris 179, 184, 187
Gardenia go
wetzleri go
Gastrochaena 69
Gazellospira 288-290
torticornis 286, 288, 289 ; PI. 33, figs. 2, 3
GEORGE, E. A. 339-353
Gibbolucina 28, 6 1
(Eomiltha) subrhomboidalis 28, 61
Giraffa 285, 290
camelopardalis 267, 284, 285, 289 ; PI. 34,
figs. 5, 6
Giraffidae 284
Glans 27, 57
nigeriensis 27, 57 ; PI. 8, fig. ga, b
(Amekiglans) costaenodulosis 28, 59; PI. 9,
%s. 3, 4
(Bendeglans) costaeirregularis 28, 58 ; PI. 9,
figs, i, 2
(Divergidens) triparticostata 28, 60 ; PL 9,
figs. 5a, b, 6, 7
Glossopteris 127, 128, 130, 132, 134-136, 141,
145, 147, 149-151, 159-161 (scale leaves,
sporangia), 165, 169, 172, 173, 179-181,
184-188
ampla 141, 173
angustifolia 127, 134, 135, 179
arberi 186
brancai 145, 148
Glossopteris — contd.
browniana 134, 141, 145, 147, 160, 172, 179,
187
var. australasica 134, 135
indica 134, 141
colpodes 127, 136, 141, 142*, 143, 144*. 145,
146*, 147, 149, 159, 163, 172 ; PL 19, fig. 2
communis 134, 186
conspicua 145, 148
cor data 187
damudica 141
var. stenoneura 141
danaeoides 151
elongata 145, 148
fibrosa 127, 129*, 130, 131*, 132, 133*, 134,
136, 140, 141, 145, 149, 159, 163, 172 ; PL
18, figs. 1-5, PL 19, fig. i
hispida 127, 129*, 134, 136, 137*, 138*. 14°*.
141, 145, 149, 172 ; PL 18, figs. 6, 7, PL 19,
fig. 3, PL 20, figs. i. 2, 7
indica 127, 128, 134, 136, 141, 186
longicaulis 186
musaefolia 141
nilsoniana 179
phillipsi 179
reticulum 145, 148
retifera 145, 147, 148
tatei 145
verticillata 186
sp. A 149, 150*
Glycymeridae 27, 53
Glycymeris 53
nigeriensis 53
Glyptoaxis 60
Glyptocrinus 204, 215, 216
Glyptorthis 218
Graphiadactyllidae n
Graphiadactylus 10
Gravicalymene 211-213, 224
Grewia 107
Guilielma 331
Guilielmites 331
permianus 297, 303, 331 ; PL 40, fig. 12
Hamamelidaceae 77-79, 99, 100 ; PL 14, figs.
97-99
Hamamelis 100
Hapalosiphon 344
Haplocytheridea 4
Harknessella 197-200, 210, 218, 221
jonesi 197, 218
subplicata 197, 198, 200, 214, 215, 218
subquadrata 197, 198, 200, 214, 215, 218
vespertilio 197, 199, 218, 219 ; PL 24, fig. i.
Healdiidae n
Hedstroemina 210
fragilis 210, 222
parva 222
robust a 210
Hemicythere 6
INDEX
Heterorthina praeculta 208, 222
Heterorthis alternata 205, 207, 211, 216, 221-223
patera 197, 218, 219
retrorsistria 203, 220
Hexaplex 26, 41
(Paziella) bendeica 26, 41
calcitrapa 42
Hilda 40, 41
turriculata 40, 41
Hiltonia 295, 301, 328
n'zm/i 299-304, 306, 307, 324, 328
Hipparion 267, 281, 290
antelopinum 280
crusafonti 281
gracile 280
theobaldi 280
sp. 280, 289 ; PI. 35, figs. 3-6
Hippopotamus 267, 289
Holopea striatella 211, 223
H omotherium 270
crenatidens 270
ultimum 2 jo
sp. 270, 289 ; PI. 34, figs, i, 2
HOOIJER, D. A. 265-292
Harder ley ella 203, 219
corrugata 203, 220
plicata 197-200, 218 ; PI. 24, figs. 3, 4
Huttoniana 303
Hydrocharitaceae 76, 86
Hyolithes 218-220, 222, 224
77e# 75
Illaenus 211, 212, 224
Janiopsis 26, 44
nigeriensis 26, 44
Kidstoniella 342, 348, 352
fritschi 341, 348 ; PI. 41, figs. 2, 9, PI- 44
Kitsonia 28, 68
eocenica 28, 68 ; PI. 9, figs. 13, 14
Kitsoniidae 28, 68
Kjaerina 205, 206
bipartita 199, 205, 207, 221 ; PI. 24, fig. 19
geniculata 205, 207, 221
hedstroemi 205, 220
horderleyensis 221
intermedia 205, 221
jonesi 206, 221
richteri 221
/y/>a 199, 207, 221 ; PI. 24, fig. 20
sp. 221-223
Kjerulfina 208, 210
polycyma 208-210, 222
209, 222
Labiatae 75
Labiosa 67
(Raeta) schweinfurthi 67
Labyrinthodon pachygnathus 257
Labyrinthodont 235
Laccinum 26, 43
lugardi 26, 44
Lacinia 44
Lanceolatus 130, 165, 179, 181
Langiella 342, 344, 346, 347, 349-352
scourfieldi 341, 343 ; PL 41, figs, i, 3-8, 10,
PI. 42, PL 43, figs. 17, 19-24
Lauraceae 74, 77, 78, 97, 98 ; PL 14, figs. 81-91
Laurus primigenia 97
Leguminoseae 77, 101 ; PL 15, figs. 110-112
Leiden Museum 276, 284
Lepidocaryinae 88, 89
Lepidocoleus suecicus 219, 221, 223
Leptaena 211, 218, 220, 221, 223
Leptobos 267, 285, 286
etruscus 286
falconeri 286
stenometopon 286
vallisarni 286
sp. 285, 289 ; PL 33, fig. 4
Leptoconus 27, 50
Leptocythere 8
Leseurilla balclatchiensis 218
Leucozonia 26, 46
pseudominax 26, 46 ; PL 6, fig. 8
Levifusus 45
Lichas 198, 223
Lidgettonia 179, 180, 186, 187
africana 181, 182*, 183* ; PL 22, figs. 1-3, PL
23, figs. 4-7
Lingula 219, 221-223
attenuata 223
ovata 218
Liomesus 43
africanus 43
Liospira aequalis 218, 221
Lithophaga 68, 69
Lobatannularia 312
Lonchodomas pennatus 213, 224
Lophocythere 7, 9, 10
Lophospira gyrogonia 206, 221, 222
Loropetalum 100
Loxocardium 62
Loxoconcha 8
Lucinidae 28, 60, 68, 69
Luhea 107
Lycopersicum 119
Lysimachia 74, 115, 116
boveyana 77, 115 ; PL 17, figs. 179, 180
japonica 116
Lysimachinae 115, 116
Lythraceae 73, 77-79, 107, 108
Machairodus nihowanensis 270
Macoma 28, 66, 67
(Bendemacoma) nigeriensis 28, 66
Macrocallista 65
palmerae 29, 65
INDEX
361
Macrocypris 6
Macrodentina 7, 14
lineata 14
retirugata 15, 16
Macrotaeniopteris 151
danaeoides 151, 154
feddeni 154
major 151
Mactra semisulcata 65, 66
Mactridae 28, 65
Magnolia 74, 78, 96, 97
angusta 97
attenuata 96, 97
boveyana 77, 96, 97 ; PI. 13, figs. 76-80
glauca 97
grandi flora 97
hoffmani 97
sinuata 97
Magnoliaceae 77, 78, 96
Marionites typa 207, 221
Mastixia 78, 80, 109, 114, 115, 121
boveyana 77, 114; PI. 17, figs. 175-178
Mastixicarpum 77
Mastixioideae 77, 114
M astodonsaurus 245
robustus 254
Mazzalina 45, 48
Megalobatrachus 254
Meliosma 74, 78, 102, 103
reticulata 77, 102 ; PI. 15, figs. 113-118
Metaconularia sowerbyi 223
vesicularis 223, 224
Metopolichas verrucosa 218
Microcallista 28, 64
Microdiptera 73, 74, 78, 79, 107
major 107
parva 77, 79, 107, 108*; PI. 15, figs. 133-148
Mitrella 51
Mitrelloturris 27, 51
casern 27, 52 ; PI. 7, fig. 8a, b, PI. 8, fig. i
Mixoneura 316
huttoniana 298, 303, 306
neuropteroides 316
sp. 303, 316 ; PI. 37, fig. 4
Modiolopsis modiolaris 224
postlineatus 219
Monoceratina 9
Moraceae 77, 95
Moroidea 95
boveyana 77, 95 ; PI. 13, fig. 74
Moroideae 95
Murchisonia 221
Muricidae 26, 41
Myidae 28, 67
Myrica 74, 75, 9°-93
boveyana 76, 90, 91 ; PI. 12, figs. 45-48
suppani 92
Myricaceae 76, 90
Myrtaceae 77, in, 112
Myrtinae in
Myrtoidea 79
Myrtospermura 77, 111-113
boveyanum 77, 112, 114 ; PL 16, figs. 160-168
dubium 77, 113, 114 ; PI. 16, figs. 169-172
variabile 79, 112-114
sp. 113 ; PI. 16, figs. 173, 174
Myxophyceae 341, 342, 350
Naticidae 26, 39
Natsiatum 79
Nemagraptus gracilis 198, 214, 215, 226-228
Neocalamites carreri 180
mansfeldicus 312
Nerita 35
Neuropteris 297
huttoniana 296, 299, 316
Neverita 26, 39, 40
amekiensis 26, 39 ; PI. 6, fig. 50, b
calvimontana 39, 40
New South Wales 163, 173, 233-263
Nicolella 212, 221
actoniae 211, 212, 222, 223
Nieszkowskia stubblefieldi 201, 219
Nigeria 23-70
Nilssonia 151
Nilssoniopteris 151
Nipteraxis 25, 32
Nodularia 344
Noeggerathia aequalis 321
cuneifolia 297
Noeggerathiopsis 128, 159
hislopi 306
Noetiidae 27, 54
Nostocales 352
Nothofagus 94
Nucula 27, 53
costaeimbricatis 27, 53
Nuculidae 27, 53
Nummulites beaumonti 29
Nummulospermum bowenense 188
Nyctereutes 269, 290
megamastoides 268, 269, 289 ; PI. 34, fig. 7
procyonoides 269
Nymphaea doris 96
Nymphaeaceae 77, 96
Nyssa 74, 109-1 u, 117
boveyana 77, 109, in ; PI. 16, figs. 150-159
disseminata no, in
europea 109, 116
laevigata 109
microsperma 109
obovata 109, 117
ornithobroma 109
rugosa in
striolata 109
sylvatica no, in
var. biflora in ; PI. 16, fig. 159
europaea PI. 16, fig. 158
vertumni 109
Nyssaceae 77, 109
362
INDEX
Odontopteris 297
goepperti 305
Ogmoconcha n
Oligocythereis 9
Olividae 26, 47
Onnia 212-214
cobboldi 199, 212, 213, 225 ; PI. 26, fig. 14
gracilis 199, 213, 225 ; PI. 26, fig. 15
superba 199, 212, 213, 225, 226, 229 ; PI. 26,
fig. 16
Onniella 210, 211, 213, 224
aspasia 223
broeggeri 199, 213, 224 ; PI. 25, figs. 10, n
depressa 211, 223
grandis 210-212, 223
inconstans 199, 213, 224 ; PI. 25, figs. 8, 9
reuschi 199, 208, 209, 222 ; PI. 25, figs. 5, 6
sinuata 211, 223
Orbiculoidea perrugata 223, 224
sp. 218-221
Orbitolites complanatus 29
Orthis confinis 197
Orthoceras 218-220, 224
Orthodesma 224
Orthograptus 230
apiculatus 220, 225, 228
calcaratus var. vulgatus 227
truncatus 224, 227, 228
var. intermedius 227, 228
pauperatus 227
socialis 229
Orthonota 220, 224
subcylindrica 224
Orthorhynchula 219
Oscillatoriaceae 352
Osmunda 73-76, 78, 80, 81
lignitum 76, 80, 81 ; PL n, figs. 1-6
regale 81
Osmundaceae 76, 80, 81
Ostrea 27, 55
amekiensis 27, 55 ; PL 8, fig. 7
ludensis 55, 56
marginidentata 56
pseudomarginidentata 27, 56,; PL 8, fig. 8
(Crassos/fea) lugardi 27, 56
Ostreidae 27, 55
Otarion 209, 223
Ottokaria 130
bengalensis 165
/es/n 165
ovalis 165
Paijenborchella 8
Palaeophycus 332
insignis 331, 332
Palaeospathe daemonorops 89
Palaeovittaria 151
Palmacites 89
daemonorops 88
Palmae 88, 89
Palmenella 8
PANT, D. D. 125-175
Panthera leo 267
Parabasilicus powisi 219-221
Paracalamites 311, 312
decor atus 311
kutorgai 298, 303-3 06, 311, 812,313; PI. 37, fig. i
sibiricus 311
striatus 311
Paracyclotosaurus 235, 237
davidi 235, 237, 238*, 239*, 240* 241*, 242*,
243*, 247*, 248*. 249*, 250*, 251*, 252*,
255*. 258 ; Pis. 27-31
hemprichi 258
Parotosaurus 257-259
helgolandicus 258
Parthenocissus 103, 104
boveyana 77, 103, 105 ; PI. 15, figs. 123-125
britannica 77, 103, 105 ; PI. 15, figs. 119-122
Pater ula albida 219
Paziella 26, 41
Pecopteris danaeoides 151
(Hemitelia] lignitum 80
Pectunculus ambiguus 224
Pentamerus 225, 226
Peronaea 66
nigeriensis 66
Perulithes 45
Phacoides 28, 60
eaglesomei 28, 60
subrhomboidalis 61
Phacopidina apiculata 206, 207, 221—223
harnagensis 201, 221
Phellodendron 79
Philhedra drummuckensis 221
Phillip "sinella 229
Phragmolites 219
Phyllotheca 312
Piaea 295, 301, 307
anglica 301, 304, 307 ; PI. 36, fig. 3
gigantea 305
punctata 305, 307
Picea 75
Pinnularia capillacea 300
Pinus 75
Pi/ay 28, 63
amekiensis 28, 63; PI. 9, figs. n«-c, 12, PI.
10, fig. i
Placuna 56
Plagiozamites 323
Platylichas 211-213, 229
laxatus 199, 206, 210, 224, 225 ; PI. 26, figs.
12, 13
Platyodon 30
klinghardti 29, 67
Platystrophia 221, 223
Plectorthis plicatella 221
virgata 222
Pleurocythere 7
Pleurograptus linearis 226, 227, 229, 230
INDEX
363
Plicatula 27, 29, 55
polymorpha 27, 29, 55
Plicatulidae 27, 55
Plumulites 219
Podocarpus 75
Poirieria 42
calcitrapa 41
Polinices 40
Polysiphonia sternbergiana 296, 297, 310
Pompholigina 28, 6 1
(Eodivaricella) oppenheimi 28, 61
Ponder odicty a 10, n
Potamididae 25, 30
Potamogeton 74, 85
cristata 85
natans 85
pygmaeus 79, 85
tenuicarpus 76, 79, 85 ; PI. n, figs. 12-14
Potamogetonaceae 76, 85
Primaspis caractaci 210-212, 223, 224
(Acidaspis) caractaci 211
harnagensis 201
Primitia nana 219
simplex 218, 219
strangulata 218
Sp. 22O, 222, 224
Primulaceae 77, 115, 116
Proboscidea 270
Procyclotosaurus 257
Proetus 223
Progonocythere 7, 9
Progonocytherinae 9
Promecosomina 254
Protoaltingia 79
Protonoetia 27, 54
nigeriensis 27, 54
Pseudoctenis 295, 321, 323
middridgensis 299, 303, 306, 318, 321, 322* ;
PL 38, fig- 5
Pseudoliva 26, 47
(Buccinorbis) kitsoni 26, 47
Pseudomazzalina 26, 44
nigeriensis 26, 45
Pseudosphaerexochus 213, 225
Pseudovoltzia 327, 328
liebeana 297-306, 327*, 328* ; PI. 40, figs.
i, 4, 6
Psygmophyllum 316, 319
cuneifolium 295, 299, 303, 306, 316, 317*,
318* ; PI. 40, figs. 14, 15
Pteridospermae 313
Pterinea 222, 224
Pterophyllum 323
Pteropurpura tricarinata 42
Pteruchus 187
Pterynotus 26, 42
newtoni 26, 42
(Pteropurpura) tricarinatus 43
Pullastra speciosa 224
Quasillitacea i, 6, 10-12
Quasillites 10, 11 ; PI. 2, figs, i, 2
Quasillitidae 10, n
Quasillitinae 10
Quercus 75
i/e# 75
67
schweinfurthi 67
Raetomya 28-30, 67
schweinfurthi 28, 29, 67
Rafinesquina 198, 203, 218, 220, 221, 223
complanata 218
expansa 203, 204, 218, 220
AO//Z 199, 213, 224 ; PI. 25, fig. 12
Raphiophorus edgelli 213, 224, 225
Raphistoma 219
Raphistomina 218, 224
Raymondella gigantea 207
£y^>a 205
Reacalymene 206, 221
pusulosa 198, 218, 220
Rectangularca 27, 54
africana 27, 55 ; PI. 8, figs. 50, fc, 6
Reedolithus subradiatus 197
Remopleurides 211, 224
burmeisteri 213
Resserella canalis 221
(Cryptothyris) paracyclica 210
Reuschella 203
bilobata 211, 212, 222, 223
horderleyensis 199, 203, 204, 220 ; PL 24,
figs. 10, ii
reuschi 200
semiglobata 211, 212, 223
Reuscholithus 198, 200, 201
reuschi 199-202, 215, 219, 228 ; PI. 26, fig. 3
Rhabdocarpus klockeanus 306
Rhabdotaenia 127, 149, 151, 172
danaeoides 127, 151, 152*, 153, 154, 157, 159
harkini 127, 151, 154, 155*, i56*-i58*, 159,
172 ; PI. 20, fig. 5
Rhadalognathus 254
boweni 257
Rhaphanocrinus basalis 220
(Glyptocrinus) basalis 204
Rhinesuchus 239
Rhinoceros etruscus 267
Rhinocerotidae 281
Rhiptozamites goepperti 321
Rhopalites 45
africanus 45
Rhynia 347
Rhyniella 342, 350-352
vermiformis 341, 350; PI. 41, figs, i, n, PI.
43. fig- 18
Rhysocythere i, 14, 18, 19
Rivulariaceae 344
Ropolenellidae 9—11
Ropolenellus 9, 10
INDEX
Rosaceae 77, 78, 101
Rostellaria bidigitata 36
unidigitata 36, 37
Rostricellula triangularis 219
Rubiaceae 90
Rubus 74, 78, 101
acutiformis 101
microspermus 77, 101 ; PI. 14, figs. 100-109
Rutaceae 77, 102
Rutaspermum 102
exaratum 77, 102
Sabiaceae 77, 102
Sagenopteris 179
longicaulis 186
Sagittaria 75
Salopia salteri 198, 199, 218, 219 ; PI. 24, figs 7, 8
Salterolithus 198, 200, 201, 203, 215, 220, 228
caractaci 199-202, 219 ; PI. 26, fig. 5
harnagensis 200, 201, 219
intermedius 200
smeathenensis 199, 201, 219 ; PI. 26, fig. 4
Salvinia 74, 81, 82
ancillata 82
auriculata 82
boveyana 76, 81 ; PI. n, figs. 7-11
hassiaca 82
natans 82
oblongifolia 82
Salviniaceae 76, 81
Samaropsis 187, 302, 330
seixasi 187
triangularis 300, 303, 305, 306, 330 ; PI. 40,
figs. 2, 3
Sampo 211, 212, 223
Scalidae 25, 34
Schizaeaceae 75
Schizocrania crassa 222
Schizocythere 8
Schizoneura 128, 312
Schizopteris digitata 319
Schuleridea 7
Schutzia 319
anomala 319
sp. 301, 304, 319 ; PI. 37, figs. 7-12
Scissodesma 66
Scutum 130, 165, 179, 181, 186, 188
Scytonema (Diplocolon) crustaceum 351
Semiterebellum 26, 37, 38
elongatum 37, 38
suturocostatum 38, 39
(Africoterebellum) elongatum 26, 38 ; PL 5,
fig. ioa, fe
Sequoia 73-76, 78, 81, 82
couttsiae 76, 79, 81, 82, 83
gigantea 84
sempervirens 84
Sinodia 28, 64, 65
heward-belli 28, 64
(Cordiopsis) orbicularis 65
Sinodiopsis 28, 64
coxi 28, 29, 63-66 ; PI. 10, figs. 4, 5
Sinowilsonia 100
Sinuites 206, 210, 211, 222, 224
anceps 221
bilobatus 220, 221, 223
pseudocompressus 223, 224
soudleyensis 220-222
Sinum 26, 40
africanum 26, 40 ; PI. 6, fig. 6o-c
clathratum 40
nigeriense 26, 40 ; PI. 6, fig. 7^, fe
Siphonotreta scotica 219
micula 218
Smeathenella harnagensis 199, 219 ; PI. 24,
figs. 5, 6
strophomenoides 198, 218
Solanaceae 77, 78, 118, 119
Solanispermum 118, 119
reniformis 77-79, 118 ; PI. 17, figs. 189-191
Solariaxis 25, 33
canaliculata 32
spectabilis 33
Soudleyella 204
avelinei 199, 204, 220 ; PI. 24, fig. 9
(Onniella) avelinei 203
South Africa 180
Sowerbyella 218, 220, 224
sericea 210, 216, 219, 221, 223
soudleyensis 206, 221
Spermatites 127, 165
crystallinus 127, 163, 165, 166*, 168*, 169,
172, 173 ; PI. 20, fig. 6
indicus 169
tetrapterus 127, 163, 169, 170*, 171*, 172, 173
Sphaerococcites dyadicus 305
Sphaerocypraea 26, 39
bowerbanki 39
sudanensis 26, 39 ; PI. 6, fig. 40, 6
Sphenobaiera 319
digitata 299, 301, 303-307, 319, 320*; PI. 38,
figs. 4, 6
Sphenopteris 297, 305, 315
bipinnata 301, 304, 305, 307, 315*, 316, 319 ;
PI. 37, fig. 6
cuneifolia 316
densifolia 305
dichotoma 297, 315
erosa 297
geinitzi 305
germanica 319
gibbelsi 305
gillitzeri 305
gothani 305
incerta 316
kukukiana 305
naumanni 297
patens 316
ropkei 305
sterzeli 305
INDEX
365
Spirematospermum go
wetzleri 76, 79, 90
Spisula 28, 65
(Crepispisula) amekiensis 28, 65 ; PI. 10, figs.
6a, b, 7
Spongillopsis dyadica 297, 331
Stangeria 187
Stegodon 267, 289
Stellaxis 25, 33
bicingulata 33
Stigonema 343, 344
dedroideum 352
Stigonemataceae 341, 342, 352
Stigonematales 342
STONELEY, H. M. M. 293-337
Stratiotes 74, 78, 79, 86
headonensis 80
kaltennordheimensis 80, 86
websteri 76, 80, 86 ; PI. u, figs. 15-19
zinndorfi 80
Strepsidura 26, 29, 43, 48
spirata 48
(Strepsiduropsis) spirata 26, 48
Strepsiduridae 26, 48
Strepsiduropsis 26, 48
Streptelasma 221
Strepula sigmoidalis 9
Striarca 55
africana 54, 55
Strobilifer frumentarius 324, 330
Strobilites 329, 330
bronni 300, 302-306, 323, 329 ; PI. 40, figs.
7-1 1
dentatus 305
elongatus 305
ludwigi 302, 304, 305, 307, 330 ; PI. 39, fig. 5
major 305
minor 305
Strombidae 25, 36
Strophomena grandis 222, 223
Stylohipparion 281
Subcyclotosaurus 258
brookvalensis 235, 258, 259*
Suidae 283
Surcula 27, 51
africana 51
ingens 51
Surculites 27, 51
(CKwwra) ingens 27, 51
SMS 284
minor 284
sfrozn 283, 284, 289 ; PL 33, fig. i ; PL 34,
fig. 8
Sveltella 48
Sveltia 27, 48
(Africosveltia) multiplicis 27, 49 ; PL 7, fig.
Sycostoma 45
SYLVESTER- BRADLEY, P. C. 1-21
Symplocaceae 77, 116
Symplocos 74, 75, 78, 109, 116, 117
anglica 77, 116, 117 ; PL 17, figs. 181-186
gregaria 117
headonensis 77, 117 ; PL 17, figs. 187, 188
117
Taeniopteris 127, 154, 316
criciumensis 154
daintreei 154
danaeoides 127, 151, 172
duffiana 296, 316
eckardti 299, 303, 305, 306, 316 ; PL 37, fig.
3«, 6
vittata 151
Tanganyika 125-175
Taxodineae 76, 82
Taxodium distichum 75
Taxus 88
baccata 75
nitens 87
Tellinidae 28, 66
Tentaculites 209
anglicus 210
scalaris 222
Terebellum 37
Terebralia 25, 30
amekiensis 25, 31
nigeriensis 25, 30
Tetradella 211
scripta 218-220, 222-224
Tibia 25, 36, 37
bidigitata 25, 36
T*7ia 75, 77, 106, 107 ; PL 15, figs. 130-132
Tiliaceae 106, 107
Titanites 15
Tivelina 28, 62
newtoni 28, 62 ; PL 9, figs. 90, 6, ioa, b
sphenarium 62
Tolypothrix elenkinii 351
Trachycarpus 88
excelsis 88
Trachyleberidinae 9
Trachyleberis 6*, 8, 9
Trematis punctata 222, 223
Tretaspis 212
kjaeri 212, 229
Triarthrus 213, 225, 229
Triebelina 6
Trigonostoma 50
decorata 49, 50
Trimerorhachis 251
Trinucleus 193, 194, 215, 217, 226
Triplesia 223
Tropidodiscus acutus 210, 222-224
Tubulites 310
articulatus 310, 311
permianus 300, 304-306, 310 ; PL 36, figs. 6-9
Tugurium nigeriense 36
Turbinolia 29
366 INDEX
Turbocalyptraea 25, 35
scabrosa 25, 35 ; PI. 5, figs. 8, ga, b
Turricula 27, 51
(Surcula) africana 27, 51
Turridae 27, 51, 52
Turrilepas 219
Turritella 25, 31, 32
amekiensis 25, 31 ; PI. 5, figs, i, 2
mauryana 32
sulcifera 31
(Coeloconica) mauryana 25, 32
Turritellidae 25, 31
Ullmannia 300, 301, 306, 323, 324, 328-330
bronni 296-306, 323, 324, 326, 327, 329 ;
PI. 38, figs. la, b, 2, PI. 39, figs. 3, 4
frumentaria 296, 298-306, 324, 325*, 326*.
327. 330 ; Pi- 39, figs, i, 2
orobiformis 305
selaginoides 297, 326
Ulmaceae 75, 76, 94
Ulrichia bicornis 219
Ulricholithus ulrichi 199, 200, 202, 203 ; PI. 26,
fig. 6
Ulwellia papulata 15
Varicohilda 26, 40
turriculata 25, 41 ; PI. 7, fig. la-c
Varicorbula 28, 67, 68
amekiensis 28, 67; PI. 8, figs. 10, n, PI. 10,
fig. 8a-c
Vellamo 201, 219
Venericardia triparticostata 60
(Venericor) planicosta 57
Venertdae 28, 62
Venus incrassata 65
Vertebraria 128, 163
Vitaceae 77, 78, 103
Vitis 78, 106
brilannica 103-105
hookeri 77, 103, 106 ; PI. 15, figs. 126, 127
ludwigi 103, 105
stipitata 77, 103 ; PI. 15, figs. 128, 129
teutonica 103-105
Volemidae 26, 44
Voltzia liebeana 297, 303, 327
phillipsi 286, 298, 324, 327
Volutidae 26, 43, 46
Volutocorbis multispinosa 26, 46
Volutospina 46, 47
conicoturrita 46, 47
multispinosa 46
Walchia piniformis 297
WATSON, D. M. S. 233-263
Wattsella 205
horderleyensis 206
indica 205
lepta 206
unguis 208
wattsi 208
Weissites 297
Xenocythere 7
Xenophora 25, 36
nigeriensis 25, 36 ; PI. 6, fig. la-c
Xenophoridae 25, 36
Zanthoxyleae 90, 102
Zelkova 94, 95
boveyana 76, 94 ; PL 13,
keaki 94
sinica 94
Zingiberaceae 76, 90
Zonarites digitatus 296, 319
71-73
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